Annals of the Missouri Botanical Garden Volume XLIX — L! 1962 E G c ` Published quarterly by the Board of Trustees of the Missouri Botanical Garden and Washington University Press, St. Louis, Mo. Annals of the Missouri Botanical Garden A quarterly journal containing scientific contributions from the Missouri Botanical Garden and the Henry Shaw School of Botany of Washington University in affiliation with the Missouri Botanical Garden. Information THE ANNALS OF THE Missourt BOTANICAL GARDEN appears four times during the calendar year: eg May, September, and November. Four numbers constitute a volum Beginning with Volume 45, 1958: Subscription Price $15.00 per volume Single Numbers ... 4.00 each Contents of previous issues of the ANNALS or THE Missourt BOTANICAL GaRDEN are listed in the Agricultural Index, published by the H. W. Wilson Company. Beginning ae 1, 1962 the Stechert-Hafner Service Agency, Inc., 31 East 10th St., New York 3, N. Y., will become sole agent for Annals of the Missouri S aim: ginning with volume 50 (1963) and = IE for back issues. Out-of-print red. nder the arrangement with the Stechert-Hafner Service Agency, the Agency will handle also all diee of any portion of the Annals eet known as the “Flora of Panam All matters regarding exchange m continue to be handled by the Missouri Botanical Garden as in the pas D TABLE OF CONTENTS Studies in the New World Moraceae: Trophis, Clarisia, Acanthinophyl- William C. Burger lum Pollination and Variation in the Subtribe Catasetinae (Orchidaceae) Calaway H. Dodson Comparative Morphology of the Osmundaceae ................ Walter Hewitson The Floral Morphology of Secamone and the Evolution of the Pollinating Apparatus in Asclepiadaceae Fuad M. Safwat Index of Orchid Names— 1961 Robert L. Dressler Flora of Panama. Part IV, Fascicle 5 (Nymphaeaceae to Monimiaceae) Robert E. Woodson, Jr., Robert W. Schery and Collaborators General Index to Volume XLIX 1-34 35-56 51-93 95-129 131-136 137-255 256-259 MISSOURI BOTANICAL GARDEN STAFF Director Frits W. WENT Epcar ANDERS Curator of eege Plants Henry N. ANDREWS, Paleobotanist HucH C. CUTLER, Executive Director CARROLL W. DODGE, Mycologist Caraway H. Dopson Taxonomist and Curator of Living Plants Rosert L. DRESSLER, Taxonomist and Editor of the ANNALS James A. DUKE, Assistant Curator of the Herbarium N D. DWYER Research Associate RAYMOND FREEBORG, Research Associate Norton H. NICKERSON, Morphologist TRIFON VON SCHRENK, Assistant Curator Museum of Economic Plants Owen J. SEXTON, Research Ecologist GEORGE B. VAN SCHAACK, Curator më Grasses and Librar Rosert E. Woopson, JR. Curator of Herbarium BOARD OF TRUSTEES sident Henry HITCHCOCK Vice-President LEICESTER B. FAUST Second Vice-President Henry B. PFLAGER Howarp F. BAER DANIEL K. CATLIN SAM L. C. Davis Dons FrencH (Honorary) Joun S. LEHMANN Rosert W. OTTO Warren McK. SHAPLEIGH ROBERT BROOKINGS SMITH EX-OFFICIO MEMBERS GEORGE L. CADIG Bishop of m dl of Missour DANIEL President, Board òl Education Lou of St. RAYM STRATFORD LEE MORTON, President of the ee of Science of St. L Tuomas H Er Chancellor of E University Mayor of the City of P "Louis C. RaurET Lincorn, Secretary Volume XLIX T a ` ` Number 1 and 2 Annals of the Missouri Botanical Garden A quarterly journal containing scientific contributions from the Missouri Botanical Garden and the Henry Shaw School of Botany of Washington University in affiliation with the Missouri Botanical Garden. ae Tue ANNALS or THE ppears four times during the calendar year: ee gen ve hee de eva: Four numbers constitute a volume, Beginning with Volume 4, 1958: Subscription Price .... $1200 per volume Single Numbers .... 3.00 each Contents of previous issues of the ANNAIS or THE Misourt BOTANICAL Gear ae lid in de Agua Inder, published by the H. W. Win pany. Agency will ginning with volume 50 (1963) and all requests for back issues Out-of-print numbers will be reprinted as may be required. > Under the. srrazigemens the Sechert Hales Service the Agency will handle ab all purchase o ary porton ofthe Annals msec known as the “Flora of Panam : An ters ming echange will continue 1o, be handled the Missouri Botanical E a 7 SC Annals of the Missouri Botanical Garden Vol. XLIX February and May, 1962 No. 1 and 2 STUDIES IN NEW WORLD MORACEAE: TROPHIS, CLARISIA, ACANTHINOPHYLLUM* WILLIAM C. BURGER This study includes revisions of three New World genera of the family Moraceae which have often been mistaken for one another. Two have been considered synonymous. Trophis nophyllu revised by Ge in 1936, who treated them as a single genus, In herbaria, critical speci- mens of one genus have frequently been filed with those of one of eh other genera, and new species have ee been assigned to the wrong genus. Studying the peed .. un permitted a consideration of their ffi iius and made den Do misplaced collec- tions. The revisions are based upon the study of herbarium specimens from seventeen impor- tant herbaria of the world. These collections include most of the type specimens of the species, — and varieties treate includes discussions f he important morphological characteristics of the Ee relationship of the genera are also briefly discussed. The genus Sorocea is included n the discussi Four species are recognized in the genus Trophis. Clarisia and Acanthinophyllum are treated as separate genera, each with two species. e subspecies of T. racemosa and C. biflora and the species of Acanthinophyllum are new Wee Wax C. Bur- GER, Imperial Ethiopian College, Box 138, Dire Dawa, Ethiopi INTRODUCTION This study originally included a revision of the genus Sorocea. Only after material had been received from a number of herbaria and the study was well under way did I come to know that Professor J. Lanjouw had been studying Sorocea for many years. In July 1961 I had the good fortune of meeting Professor Lanjouw and his assistant Mr. Wessels Boer while visiting Utrecht. There we discussed Sorocea and found that our delimitation of species was very similar. We hope to publish a revision jointly in the near future. Sorocea is discussed in the following pages and included in a key to the genera. The genera considered in this study are distinguished from most other dioecious New World Moraceae by their racemose, spicate, or occasionally capitate inflores- cences. The pistillate perianth, with rare exceptions, is fused and tubular, while the alternate leaves are distichous and pinnately veined on unarmed branches. Trophis has never been monographed, but Clarisia and Acanthinophyllum were revised by Lanjouw in 1936. The vegetative resemblance and similar flowers and inflorescences of these genera have resulted in numerous misidentifications. Fre- rt of an investigation carried out in the graduate oe of the Hen ibis of. Botany of Washington University and submitted as a thesis in partial fulfillment of the requirements for the degree of Doctor of Philosophy. (1) [Vor. 49 2 ANNALS OF THE MISSOURI BOTANICAL GARDEN quently critical specimens of one genus have been filed with those of one of the other genera, and new species have occasionally been assigned to the wrong genus. Thus, Trophis macrostachya, and nearly all the specimens which were referred to it, belong with Sorocea. Lanjouw concluded that Clarisia and Acanthinophyllum are congeneric and it has been suggested more recently (Woodson, 1960, p. 114) that Trophis and Sorocea might be congeneric. Studying the genera together has permitted a consideration of their relationships and made available misplaced collections which would not have been possible had only one genus been studied. INFLORESCENCES The staminate inflorescences are racemose or spicate, or condensed into small capitula in several species of Sorocea. These inflorescences are most commonly paired in the axils of leaves. In Trophis there is a reduction to solitary inflores- cences in T. chiapensis. The spikes of Clarisia and Acanthinophyllum are especially interesting in that the stamens arise directly from the rachis and the tetramerous organization of tepals and stamens has been lost. The most elaborate type of inflo- rescence found among the genera studied is that of Clarisia racemosa. These “racemes” of staminate spikes are interpreted as leafless flowering shoots and are discussed in more detail under the special remarks on that genus. The pistillate inflorescences parallel the staminate in Trophis and Sorocea. The pistillate flowers of Clarisia may be the result of a similar reduction. The peltate bracts at the base of the receptacle and the paired, axillary flowers of C. biflora may be interpreted as evidence for such a reduction. As in the staminate spikes the development of leafless flowering shoots has apparently resulted in the elongate pistillate “racemes” of C. racemosa. The pistillate capitula of Acanthino- phyllum resemble those found in some species of Sorocea but are distinguished by the uncinate trichomes of the peduncle and the paucity of bracts subtending the flowers. The nature and origin of the spikes and racemes, characteristic of the genera under consideration, remain hypothetical. Bernbeck (1932), in an important morphological study, concluded that the inflorescences of Urticaceae and Moraceae are basically cymose. The spikes and racemes that I have seen are noteworthy in usually lacking flowers along one or sometimes two sides of the inflorescence and thus simulating helicoid cymes. This is similar to the inflorescences of Myriocarpa cordifolia and Urtica membranacea studied by Bernbeck and interpreted by him to be derived from a cymose type. While Bernbeck did not study Moraceae with inflorescences of the type found in the genera under discussion, it seems safe to assume that they, too, arose from a cymose type. STAMINATE FLOWERS The staminate flowers fall into two categories: those that are definitely organ- ized and tetramerous and those where the organization has been lost and the flowers are reduced to single stamens. The tetramerous flowers are readily distin- guishable in Trophis and Sorocea. The stamens are inflexed before anthesis and the perianth is valvate in aestivation in Trophis, while the stamens are straight and 1962] BURGER—STUDIES IN MORACEAE 3 the perianth is decussate-imbricate in Sorocea. Dehiscence of the anthers is introrse in Trophis, but is usually extrorse in Sorocea; a pistillode is always present in Trophis, but is rare in Sorocea. These distinctive characteristics clearly serve to separate the two genera despite the similarity of the pistillate flowers. The reduced flowers of Clarisia and Acanthinophyllum, lacking a definite perianth, are more difficult to distinguish. The stamens of both genera are straight before anthesis, but the anthers of Clarisia are subbasifixed while those of Acan- thinophyllum are usually dorsifixed. The smaller size of the anthers (ca. 0.3 mm.) and the retrorse uncinate trichomes of the peduncle further distinguish Acanthi- nophyllum. I believe that the reduction of the staminate flowers in these two genera does not reflect a true relationship but rather a parallel evolution. Without intermediate stages or morphogenetic studies it is not possible to indicate how this reduction took place. Woodson (1960, p. 125) has described an interesting series among several species in Brosimum where there is a gradual reduction in stamen number and a striking change in the mode of dehiscence. However there may be another way in which the staminate flowers became reduced. This would require that the flowers first became broadly sessile with the stamens and perianth parts arising separately from the rachis as in some species of Sorocea. Following this, reduction in the perianth parts together with a loss of symmetry might give rise to spikes such as are found in Clarisia and Acanthinophyllum. PIsTILLATE FLOWERS The pistillate flowers of the four genera under discussion are very similar. This is not unexpected since the basic plan of the gynoecium is quite uniform in the Urticales. The fusion of perianth parts of the pistillate flower is frequent in Moraceae and is not in itself indicative of close relationship. I have placed some emphasis on the length and form of the style branches which would separate Trophis and Clarisia from Sorocea and Acanthinophyllum. The short style branches and free perianth tube of Acanthinophyllum ally it to Sorocea rather than to Clarisia, with which it had been made congeneric by Lanjouw. The fruit of the genera are quite similar, but the embryo of Sorocea may be separated on the basis of its fused cotyledons and prominent hypocotyl 2-3 mm. long. The cotyledons of the other genera are free and the hypocotyl is minute. ANATOMICAL CHARACTERS The trichomes, while often not consistent among all the species of a genus, may nevertheless be useful in suggesting relationships with other genera. Sorocea, for example, usually possesses minute (0.05 mm.) multicellular gland-tipped trichomes. Though species of Sorocea in Central America may lack these trichomes, I believe it is significant that they are not found in any of the other genera studied. Acanthinophyllum can be immediately distinguished from the other genera under consideration by its small (0.1 mm.) retrorse uncinate trichomes. The leaf surfaces of Trophis can be distinguished from other genera by the minute (0.02 mm.) broadly aculeiform trichomes bordered by about a dozen small epidermal cells. These trichomes are numerous in the usually scabrous leaves of T. racemosa but [Vor. 49 4 ANNALS OF THE MISSOURI BOTANICAL GARDEN sparse in the other species. Clarisia lacks distinguishing trichomes that might serve to aid in its placement. The epidermal cells of Acanthinophyllum have a deeply sinuate outline unlike those of the other genera. These and the smaller trichomes can be seen on dried herbarium specimens under low power of the compound microscope by reflected light; clearing is unnecessary. RELATIONSHIPS OF THE GENERA The subfamilies Moroideae and Artocarpoideae have been defined on the basis of whether the stamens are inflexed or straight in bud. As a result, Trophis has been placed in the Moroideae and Clarisia, Acanthinophyllum, and Sorocea in the Artocarpoideae. These subfamilies also have been divided on the basis of leaf vernation. The Moroideae have conduplicate leaf vernation while the Artocar- poideae have involute leaves. Clarisia has conduplicate leaf vernation but straight stamen aestivation, a contradiction of the subfamilial criteria. If the staminate flowers have become disorganized there is no reason to assume the orientation of the stamens could not change radically. I believe that the criterion of staminal position prior to anthesis cannot be critically applied to such reduced forms as Clarisia and that other characters must be sought in determining relationships. Inflorescence type has been used in classification at the tribal level. In Sorocea and Clarisia, however, the inflorescences are quite variable and anatomical criteria may prove more important in determining generic relationships. In the absence of a modern study of the family it is not possible to define the relationships of the four genera under discussion. The short remarks that follow indicate what I believe to be their aftınities. Trophis, with the stamens inflexed in bud, is clearly a member of the Moro- ideae related to Paratrophis of the western Pacific. The valvate aestivation of the staminate flowers and consistently tubular perianth in the pistillate flowers separate Trophis from Paratrophis despite the fact that P. philippinensis has a tubular pistil- late perianth very similar to that of Trophis. Clarisia, with long slender style branches and subbasifixed anthers, resembles Trophis more closely than the other genera I have studied. This would imply that the stamens were originally inflexed in bud and agrees with the fact that the leaves are conduplicate in bud; a characteristic of the subfamily Moroideae. I would therefore transfer Clarisia from the subfamily Artocarpoideae to the Moroideae and place it near Trophis among New World genera. Acanthinophyllum was considered congeneric with Clarisia by Lanjouw but I believe that actually they are not closely related. The short-styled pistillate flowers with free perianth tube resemble those of Sorocea. The retrorse uncinate trichomes and sinuate epidermal cells are also found in Trymatococcus of the Moroideae and Helianthostylis of the Artocarpoideae among American genera. On the basis of the characters of the pistillate flowers I would retain Acanthinophyllum in the Arto- carpoideae and place it near Helianthostylis. Sorocea is clearly referable to the Artocarpoideae where it was placed in the fundamental study of Trécul (1847) and with which later investigators have con- 1962] BURGER— STUDIES IN MORACEAE 5 curred. Inasmuch as the New World Artocarpoideae are in need of a thorough general study there is no point to suggesting a change in the placement of Sorocea by Engler in “Das natürlichen Pflanzenfamilien” which was based on the work of Bentham and Hooker (1880). ILLUSTRATIONS The illustrations are my own. The foliage is nearly half natural size while the staminate and pistillate flowers are about 8x. The inflorescences with young fruit are reproduced at a magnification of about 4x. MATERIALS This study is based on specimens from the following herbaria. The abbrevia- tions are taken from Lanjouw & Stafleu’s ‘Index Herbariorum,’ Part 1 (Regnum Vegetabile vol. 15, 4th edition, 1959). A—Arnold Arboretum, Harvard University, Cambridge B—Botanisches Museum, Berlin-Dahlem BM—British Museum, London C—Botanical Museum and Herbarium, Copenhagen F—Chicago Natural History Museum, Chicago GH—Gray Herbarium of Harvard University, Cambridge LIL—Instituto Miguel Lillo, Tucumán MEXU—Herbario Nacional del Instituto de Biología, Mexico MG—Museu Paraense Emilio Goeldi, Belem MICH—University Herbarium, University of Michigan, Ann Arbor MO—Missouri Botanical Garden, Saint Louis NY—New York Botanical Garden, New York P—Muséum National d’Histoire Naturelle, Paris PB—Jardim Botanico, Rio de Janeiro S—Botanical Department, Naturhistoriska Riksmuseum, Stockholm US—U. S. National Museum, Smithsonian Institution, Washington Y—Yale School of Forestry, New Haven I am indebted to the curators of these herbaria for the loan of specimens which included many valuable types. I also wish to thank Dr. George Van Schaack for assistance in the use of the library of the Missouri Botanical Garden. I am espe- cially indebted to Professor Robert E. Woodson, Jr., for suggesting this study and for his advice and guidance throughout the course of its completion. KEY TO THE GENERA A. Cotyledons distinct in the seed; staminate flowers tetramerous with valvate aestivation or not definitely organized; lacking multicellular gland-tipped EC B thers 0.5 mm. long or longer; pistillate flowers with long slender style branches; peduncles lackin i retri uncinate trichomes; leaves conduplicate in bud, midvein impressed above ....—— C C. Staminate flowers tetramerous, the tepals valvate and the stamens inflexed in aestivation; pistillate flowers ile or if pedicellate lacking bracts at the base of the rec ROPHIS eptacle Staminate flowers not definitely organized; pistillate flowers pedicel- ptacle C late with peltate bracts at the base of the receptacle ........................ II Crarısıa a [Vor. 49 6 ANNALS OF THE MISSOURI BOTANICAL GARDEN BB. Anthers 0.2—.03 mm. long the staminate flowers not definitely or- ganized; pistillate flowers with short broad style branches; paduncies densely invested "n minute (0.1 mm.) KR uncinate trichom II A aves involute in , the midvein prominulous ............ CANTHINOPHYLLUM AA. Cotyledons fused in the Enos style branches usually short and broad; stam nate flowers tetramerous, the tepals decussate-imbricate the stamens straight in aestivation; leaves involute in bud, multicellular gland-tipped trichomes usually present on the leaves, midvein impressed above ........... IV Sorocea TAXONOMY Ld TROPHIS P Br. Tropuis P. Br. Hist. Jamaica 357. 1756, nom. conserv. [T.: T. Americana L. = T. racemosa L.)-Urb. Bucephalon L. Sp. Pl. 1190. D aw rejic. [T.: B. racemosum L.] Skutchia Pax & Hoffm. ex Morton, in Journ. Wash. Acad. Sci. 27: 306. 1937. [T.: S. caudata Pax & Hoffm 1 Dioecious laticiferous unarmed shrubs and trees. Leaves alternate, distichous, bistipulate, pinnately veined, the petioles sulcate above. Stipules lateral, caducous or occasionally persistent, their scars encircling less than half the stem. Inflores- cences axillary on the leafy stems, paired or solitary, racemose or spicate, with triangular or suborbicular bracts along the rachis. Flowers unisexual, regular and basically tetramerous, usually lacking along one side of the inflorescence. Staminate flowers: sessile or pedicellate; perianth 4-parted or 4-lobed, the tepals equal and valvate; stamens 4, antetepalous, strongly inflexed in the bud but erect at anthesis, filaments slender and glabrous, the anthers subcentral with 2 somewhat divergent thecae, dehiscing longitudinally, introrse; pistillode always present, cylindrical to rectangular, puberulent. Pistillate flowers: sessile or pedicellate, the perianth tubu- lar, irregularly or minutely 4-lobed at the apex; pistil 1, the ovary superior to inferior by adnation of the perianth tube, 2-carpellate, unilocular, the ovule solitary, anatropous and pendulous from below the apex of the locule, the style deeply bifid, the style branches long and slender with the inner stigmatic surface densely and minutely papillate. Fruit drupaceous, the perianth tube accrescent and succulent, the stone globose, the embryo large and straight with thick equal cotyledons and minute hypocotyl, without endosperm. The genus was first recorded by Plumier in 1703 (Nova Plant. Amer. Gen. p. 52) as Bucephalon and this was adopted by Linnaeus in his Genera Plantarum (1737) and Species Plantarum (1753). However, preferring a more adequate de- scription, Linnaeus later (Systema Naturae ed. 10, p. 1289. 1759) chose to use the name Trophis described by Patrick Browne in 1756 (p. 357). Usage has concurred with Linnaeus and has led to the conservation of Trophis. The name is derived from the Greek word tpogr (nourishment) referring to its frequent use as forage. Trophis has not received monographic study since Bureau's treatment of the Moraceae in De Candolle’s ‘Prodromus’ (1873) and although the species are rather closely knit, their identification is usually not difficult. Determining a phylogenetic sequence for the four species, however, is a somewhat conjectural matter. I have begun the sequence with T. racemosa, in some respects the most specialized species. T. racemosa rejoys a much wider distribution, has adapted to a greater range of habitats, and possesses greater morphological variation than is found in any other 1962] BURGER—STUDIES IN MORACEAE T species. These considerations lead me to suspect that it is the oldest element of the genus. The lack of fusion of the staminate perianth parts corroborates this view, but the usually inferior ovary and condensed inflorescences are definitely advanced. I believe that Trophis is closely related to Paratrophis of the southwest Pacific in which the pistillate perianth is usually 4-parted and imbricate. In Para- trophis the pistillate flowers are broadly sessile, quite similar to T. racemosa and the major argument for the antiquity of the latter. If this view is correct, the long pedicels of pistillate flowers and fruit in the other species of Trophis are secondarily evolved. The fact that the fruit of T. racemosa is never pedicellate while the fruit of the other species are occasionally sessile may indicate that the latter, less stable, condition is derived. The geographical center of the genus is in southern Mexico where all four species are found. KEY TO THE SPECIES oe flowers quite sessile and not constricted at the rachis; fruit sessile, a nsely velutinous; staminate flowers sessile and crowded, the perianth parts Geematte free; leaves frequently scabrous on either or both surfaces. Mexico to Peru, Venezuela and the West Indies l. T. RACEMOSA aa. Pistillate flowers pedicellate or rarely subsessile but constricted at the rachis; fruit usually pedicellate, sparsely puberulent to glabrescent; staminate flowers sessile or pedicellate, usually not crowded, the perianth parts connate toward the base; leaves never scabrous. Leaves glabrous or very ry spar rsely piene beneath; branchlets slender; eine eg a n inerous-pube o sessile; Se poc nce to 10 cm. long; leaves lips a GC eg oblong or obovate, usually over 3 e wide. 2. T. MEXICANA n fees pedicellate; pistillate inflorescences to Be cm. long; e e N kann linear-lanceolate to narrow elliptic, less than S wide. rn Mexico to Guatemala 3 'T. CHIAPENSIS bb. Leaves eege short- er beneath; ‘peng pa thick; pistillate flowers densely ferrruginous-puberulent. Sout E 4. T. CUSPIDATA l. Tropuis RACEMOSA (L.) Urb. Symb. Ant. 4:195. 5: Bucephalon racemosum L. Sp. Pl. 1190. 1753. [T.: Linn. Herb. London (1156.1) (9 4) Trop hi aneii i L. Syst. Nat. ed. 10. 1289. 1759. Shrubs or trees to 18 m. tall, the trunk to 50 cm. in diameter and often divided, with rough brown bark and milky latex; leafy branchlets usually stout, 2.0-3.8 mm in diameter, hirtellous becoming glabrescent, conspicuously lenticellate on older parts. Leaf blades obovate to oblong or elliptic, 5-23 cm. long, 2.4-10 cm. broad, acuminate to subcaudate-acuminate, obtuse to acute and often slightly oblique at the base, subcoriaceous to membranaceous, often scabrous, glabrous, above an glabrous or minutely strigose beneath, the midvein plane or impressed above, prom- inent below with 4-8 pairs of ascending secondary veins arching upward near the entire or serrulate, occasionally deeply sinuate margin; petiole 4-16 mm. long, 0.7-2.0 mm. in diameter, minutely puberulent and glabrescent. Stipules caducous or persistent, lanceolate, 2.0-3.8 mm. long, their scars small and inconspicuous. Staminate inflorescences: spicate, paired or solitary in the axils of leaves or rarely of undeveloped or fallen leaves, 1.5-5.0(-7.5) cm. long, the peduncles 1.3-11.0 mm. long, minutely velutinous, the rachis with numerous triangular or suborbicular [Vor. 49 8 ANNALS OF THE MISSOURI BOTANICAL GARDEN peltate or basally attached bracts 0.5-1.1 mm. broad; flowers 15 to many and densely crowded, sessile or rarely subsessile, about 4 mm. long and 4 mm. wide, the peri- anth parts 1.6-2.2 mm. long, 1.0-1.5 mm. wide, acute or obtuse at the apex, free or basally connate, minutely puberulent, the trichomes about 0.1 mm. long; stam- inal filaments 2.0-2.6 mm. long, the anthers 0.8-1.2 mm. long; pistillode 0.3-0.6 mm. long. Pistillate inflorescence; spicate, paired or solitary in the axils of leaves or rarely fallen or undeveloped leaves, 1.8-4.0(-6.0) cm. long, the peduncle 1.2-10 (-14) mm. long, minutely velutinous, the rachis with numerous triangular or suborbicular peltate or basally attached bracts 0.5-1.2 mm. broad; flowers 4-15, broadly sessile, ovoid or conic, the perianth tube indistinctly or minutely 4-lobed at the apex, 2.0-4.5 mm. long, 1.4-2.5 mm. in diameter, densely velutinous, the tri- chomes about 0.1 mm. long, ovary inferior or rarely half-inferior, the style branches slender, 2.0-5.5 mm. long. Fruit globose or ovoid, occasionally with a narrow collar at the apex, about 1 cm. in diameter, smooth or ridged, minutely velutinous, becoming bright red and sometimes blackish. This species is quite common throughout a range that extends from southern Sonora, Mexico, to eastern Peru and the West Indies. It is found on hillsides and river valleys in Mexico and moist forests in Central and South America. Within this wide area it has begun to differentiate into three morphologically distinct populations which I am recognizing as subspecies. Subspecies ramon and racemosa with their relatively thick leaves have apparently become adapted to more xeric habitats than ssp. meridionalis. The elongate inflorescences and occasionally half- inferior ovaries of ssp. meridionalis add to my impression of its more primitive nature. Subspecies ramon and meridionalis intergrade quite imperceptibly in west- ern Panama and northern Colombia. I have used the Panama-Colombia boundary to separate the two subspecies realizing, of course, that no real discontinuity exists. The membranaceous, serrulate and long-acuminate leaves of most Colombian speci- mens make their assignment to ssp. meridionalis quite natural even though some may have the congested pistillate inflorescences characteristic of ssp. ramon. T. racemosa is closely related to T. mexicana with which it is occasionally con- fused. The densely velutinous fruit and scabrous leaves are diagnostic. KEY TO THE SUBSPECIE a ru sis genii to 45 cm. long at anthesis, dit flowers distant, usually . apart; ine smooth on both surfaces, tapering to the ae b. eaves wéi ire, firmly chartaceous to subcoriaceous, acuminate; rachis of the a inflorescence I ër 5 mm. in diameter. West Indies....lc. ssp. RACEMOSA bb. mbra to chartaceous, na uminate; dw of Me "pictis iSo ce be 0.6-1.2 mm. in diameter. Colombi t zuela la. ssp. MERIDIONALIS aa. Pistillate inflorescences to 2.5 cm. long at anthesis, the flowers crowded, rarely e than rt; leaves aia or scabrescent on either or both sur- faces, GH chartaceous to subcoriaceous, usually abruptly short-acuminate. Mexico to Panam Ib. ssp. RAMON la. TROPHIS RACEMOSA ssp. meridionalis (Bur.) W. Burger, comb. & stat. nov. Trophis americana y meridionalis Bur. in DC Prodr. 17: 253. 1873. [T.: Bonpland 1512 P) (8)] Sorocea colombiana Standl. in Trop. Woods, no. 19: 39. [T.: Curran 186 (F) (9)] Trophis racemosa var. meridionalis (Bur.) Macbr. in POR Mus. Pub. Bot. 13°: 308. 1937. 1962] BURGER—STUDIES IN MORACEAE 9 ma ris i EN ni i i . ssp. meridionalis: from Killip 35580 (1 [upper right] & Figure 1. Trophis racemosa. ssp sa Herd peer A 2). ssp. ramon: from Burger 148 (3). ssp. racemosa: [Vor. 49 10 ANNALS OF THE MISSOURI BOTANICAL GARDEN Leaf blades elliptic to elliptic-oblong, (5.0-) 7.5-19.0 cm. long 2.5-6.5 cm. broad, gradually tapering to a long and narrowly acuminate apex, acute to obtuse at the base, membranaceous to chartaceous, smooth on both surfaces, glabrous, the margin serrulate. Staminate inflorescences: 2.5-6.0 cm. long, the flowers numerous and densely crowded or rarely somewhat distant. Pistillate inflorescences: 1.5-4.5 cm. long, the rachis 0.6-1.2 mm. in diameter, the pistillate flowers usually more than 1 mm. distant along the rachis. Flowering from late October through February and May to June. Moist forests and river valleys from sea level to altitudes of 1000 meters. ernacular names: Uspai-manchinga (Peru-Ll. Williams) ; Marfil (Venezuela- Pittier). OLOMBIA: Borívan: San Martín de Loba and icu Curran 102 18) wee Ze [(9) E US], 205 [(9) S, US], 372 (C9) US]. caou : Solano Esquinas, Little & Little 9670 [( 8) US]. cauca: ec Wie: en en jon A]. cHocó: Bahia de solano; Fée: 310 [(8) US] bank of of Rio San Juan, near Andagoya, Killip 35392 [(2) F, US]. vALLE pet cauca: La Trojita, Rio Calima, Sieger 16348 [(&) F], 16810 [(9) pe ae Killip 35580 [( 2) BM, F, S, US]; co Tr S. of Carraipia, Haught 4224 I8 ) F. US]. purumayo: La Concepció ión, eins 10840 [(8) F, A MART. H. Smith 1752 [F (2), MO (2), P (4), S (8), (8)], 2732 [(4) BM, E MO, : NDER: vicinity of Puerto Berrio, Haught 1864 [(& S]. WITHOUT PRECISE LOCALITY: ad Fluvium Magdalenam, Bonplan 1512 Ae e. Gi (Dir "mutis 2992 [(9 A: Patanemo Valley, Pittier 8659 [(4) NY, US]; Parque Nacional, LE Williams 10719 TG EE e R: NAPO na. Asplund 9387 [(9 ) NY, P]; Tiputini-Lagarto Cocha, Premi. d Wibom oe See S], 2403 [(2) S]; Chimbo River valley, Rimbach 187 [(9) E N ERU: Ta Aramango, Woytkowski 5645 [(?) MO]. muanuco: Tingo Maria, Stork & Horton 9506 [(&) F]. LORETO: Yurimaguas, lower Río Huallaga, Killip & Smith 85 {19 Fl d 276 F]; Florida, Rio Putumayo, Klug 20709 [(&) BM, F, MO, S]; above Pongo de Manseriche, Mexia 6282 [(4) BM, F, LIL, MICH, MO, S], 6303 [(9) BM, F, LIL, MI ]; Santiago-Mund am se de Manseriche, Sonne 4083 S]; lower Rio Huallaga, d Williams e [(9) F], san martin: Juanjui, Alto Río Huallaga, Klug 3794 [( 4) BM, F, MO, S]; near Tarapoto, Spruce 4521 [(9) 8. "EMI. Ule 6838 [( 2 ) ], LI. Williams oU I?) FT, 6573 [(9) F]. Ib. TE RACEMOSA ssp. ramon (Schlecht. & Cham.) W. Burger, comb. & stat. Tr Bos Steg Schlecht. & Cham. Linnaea 6: 357. 1831. [T.: Scheide & Deppe s.n. near Misantla (B no. 6750) (9 Trophis americana B. Ramon Bur. in DC Prodr. 17: oe 1873. aha gunia urophylla Donn. Sm. in Bot. Gaz. 40: 11, 1905. Et: här 54 (US) (9)]. 3: 263. 1936 Leaf blades abonati to eigu oian ovate or elliptic, 5.0-23 cm. long, 2.5-10 cm. wide, short-acuminate to subcaudate-acuminate or rarely tapering to a long- acuminate apex, obtuse or occasionally acute at the base, subcoriaceous to thickly chartaceous, scabrous or scabrescent, glabrous above and glabrous or minutely strigose beneath, the margin entire or occasionally serrulate, the new shoots often with deeply sinuate leaves. Staminate inflorescence: 1.2-4.0(-5.8) cm. long, the staminate flowers densely crowded. Pistillate inflorescences: 0.6-1.8(2.7) cm. long at anthesis, the flowers densely clustered or closely approximate, the rachis 0.8-2.0 mm. in diameter. The scabrescent, pergamentaceous leaves and congested inflorescences are appar- 1962] BURGER—STUDIES IN MORACEAE 11 ently associated with an adaptation to more xeric conditions than those tolerated by other members of the genus. This is the only subspecies of T. racemosa sympatric with other species of the genus. Dry hillsides, ravines, river valleys and forests; flowering intermittently throughout the year, from sea level to 2500 m. altitude. Vernacular names: Ramön (Mexico to Panama); Capomillo (Mexico-Ortega) ; Ushi (Mexico-Mexia); Cha cox (Mexico-Flores); White ramon, Ramön colorado, Yaxox (British Honduras-Lundell); Ojushte (El Salvador-Standley); Ujushte (El Salvador-Calderön); Hoja tinta (Honduras-von Hagen); Breadnut, Ojoche macho (Panama-Cooper & Slater). Mexico: cHIAPAS: Escuintla: Matuda 2598 [(9) F, mS n 4022 [( A MICH, MO], Esperanza, Matuda 16794 [(9) F], La Grada, Matuda 16799 [(4) F]; S. Terena Acapa- : OLIMA canyon slope, Mexia 1142 [(&) F, BM, MICH, MO]. mexico: ben under Acatitlán, Hinton 3165 [(&) US], 3186 i) US], vede Hinton 3643 [CS a , MO, US], Pun- garancho, Hinton 6750 [(9) F, MO, US]. : Coalcom : San edro, Hinton 15901 [(9) BM, LIL, MICH, US], Aquila, Hin "16291 DES? US]. NAYARIT: vicinity of Acaponeta, Tepic, Rose, Standley & Russell 14428 [(9) US]. oaxaca: Trapiche de la Concepción, Liebmann 14267 [( 9 ) C, F, P]; S. Jago Amatlán, Liebmann 14269 [(9 ) CFI Between S. Miguel del Puerto & Guatulce, Liebmann 14271 [(9 ) C]; Chinantla, Liehmann 14270 [(&) C]. san Luts porost: edge of El Salto River, El Naranjo, Burger 148 [(9 ) MO], 150 [(2) MO]; Serra: E 667 [(9) F]; 7 mi. W. of N. Morelos, Tam., Kenoyer & Crum 3770 [(8 ICH]. stnatoa: Cerro Colorado vicinity of Culiacán, Brandegee S. N. [(9 4) US]; oe Malquesön, Cerro Colorado, Gentry 5147 [(9 ) MICH, ]; Capadero, Sierra Tacuichamona, Gentry 5582 [(9) ebrada del gore Balboa, San. Ignacio, Montes & Salazar 855 [(9) US]; Rosario, " Cacalotán, Ortega 7100 [(8) F, "poe Marastan, Ortega 7493 [( d US]. sonora: Quiricoba, Dist. Alamos, en 736M [(9) MICH]. tamauipas: Tampico de Tamaulipas, Berlandier R2 F) Fi; Rancho Las Yucas, Sa. de eoe ange Dosis 2372 [(9 ) MO]; Julillo to Río Sabinas . of Mante, Duke 3636 [( 9) MO]; near Rancho de Barberena, LeSueur 570 [(9 ) F, US]; vicinity of Tampico, ear? 337 dei "e VERA CRUZ: Bonpland 4483 [(9) P]: Fortin, Kerber 395 [(2) BM, C, US]; Mirador, Liebmann s.n. [(?) C]; Mirador and Zacuapa Liebmann 14268 [C (9), C (2), F (?)]; Zacuapan, Liebmann 15111 [( 2 S Cordoba, Matuda 0685 AG MICH, MO, US]; Hda. Mirador, Nelson 66 [(?) U of Juan Ramirez, 56 km. S. of Tampico, Palmer 454 [(?) ee Zacuapan and vic a Pave 2316 [(9) F, MO, US], 4386 [(2) BM, MO, US]. 5049 [(9) MO, US 5988 [(4) BM, F, MO, US], 10837 [( 4) MICH, US], 12072 [(4) F, MO, us) 14372 [(4 ) F]; near Rancho Remudadero, Purpus 11070 [( 4) F], 11044 [(&) US], 1638 AA F]; Misantla, Schiede & ioc: sn I($) , MO]. xucaráw: Progreso, Flores, s.n. [(?) F]. WITHOUT PRECISE órdova, "Bourgeau 7069 [(9) P]; acuit 1468 [( 4) F]; Sesse, Mociño, Castillo P "Maldonado 4647 [(4) F]. Bnrris ONDURAs: BELIZE: Bright Lookout Bank, Sibun River, Gentle 1447 [(?) F, MICH, MO, US, S]. er cayo: Mountain Pine Rides. faciet 11867 [(?) MICH]; Monkey Falls, Lundell 6970 [(&) C, F, MICH, S]. ORAN ALK: Honey Camp, Lundell LP19 US], Meyer 157 [(?) F]. sraNN K ^19 nil Stann railway, Gentle 2704 [(2) F, M ; Silk Grass Creek Reserve, Gentle 3097 [(9 ) crie Punta Gorda, Hu Do [(4) BM, E, M , S]; Big Rock, Toledo, Schipp Jide [(9) 2 O, S]; Tomash River, Stevenson 153 [(2) F]; Stevenson & Smar 140 ite d fis aye ALTA VERAPAZ: Saquijá, 43 km. NE. of SE Standley 70185 [(&) F]; region “of Chelae, NE. of ER die eet? [(?) F]; Near Pancajché, Standley 70793 F]; mountains between Tactic and Coban, vo n Tuerckheim 11 = St 12169 [(9) C, US]; Cubilquitz, von erer 3581 [(4) F, US]; alon r, Chama, [5 ovd 514 [(4) MO, US]. cuigurwura: Volcán Quezaltepeque See "3151 Er 4) [Vor. 49 12 ANNALS OF THE MISSOURI BOTANICAL GARDEN PROGRESO: near Finca Piamonto, Steyermark 43664 UO) F]. sscuiNTLA: along Río Missions Standley 89114 [(?) F]. HUEHUETENANGO: between Finca San Rafael and Finca Providencia, Steyermark 49549 [(9) F]; along Río Trapichillo, below La Libertad, Steyermark 51030 [(&) F]; Paso del Boquerön, along Rio Trapichillo, Steyermark 51185 [(?) F]. IZABAL: near Quiriguá, Standley 72483 [(&) im PETÉN: Uaxactün, Bartlett 12354 [(?) F, MICH, US]; Santa Cruz, Bartlett 12375 [(9 ) F, MICH]; Santa Teresa , Subin River, Lundell 2917 [(9 ) BM, E MICH, S]. rETALHULEU: vicinity of Retalhuleu, Standley 88452 [(?) F], 88549 [(?) F], 88729 [(&) F]. sawrA rosa: vicinity of Chiquimulilla, Standley 79291 [(9) El: along Avellana Road, S. of Guazacapan, Standley 79513 [(9 ) F]; 79524 [(?) F]. sucurrEPÉQUuEz: vicinity of Tiquisate, Steyermark, 47670 [(?) F], 47774 [(?) F, US]. zacapa: Loma EI Picacho, Steyermark 42744 [( 2) F]. EL SALVADOR: CHALATENANGO: Calderón 2178 le n SAN SALVADOR: San Salvad Calderón 845 [(&) US], 1065 [(4) MO, US], 1448 [(9) MO, US], Standley 22458 [(?) US] san vicENTE: vicinity of San Vicente, Standley 21404 [(9 Gel SONSONATE: Izalco, Calderön 1698 [(?) US]; vicinity of Sonsonate, Standley 22340 (oi S, ES. WITHOUT PRECISE LOCALITY: GEN 112. (9 dai NDURAS: ATLANTIDA: Triunfo, near Tela, Standley 53841 [(9) F, Fu Lacus Valley, Standley 54312 [(?) F, US]; vicinity of Tela, Standley 54727 [(?) F ARA: area de pinoroble de Rio Ulúa, Molina 3810 [(2) BM, F, 1 osi EGUCI- : Mont. de la Flor, Hagen & Hagen 1130 [(?) F], 1177 12) F]. vono: Concepción, Edwards P- 644 [(&) F, US]; near Progres, Standley 54997 [(?) F, US]. wm OUT PRECISE LOCALI Highland Creek, Puerto Sierra, Wilson 54 [(2) US], 250 LES) Ad e UA: GRANADA: near Lakes, Baker 21 [(4) MICH, MO], d ES US]. MANAGUA: between EI Crucero and Finca Santa ulia, Standley 8357 [(?) F]. ZELAYA: vicinity of El neo Long 215 [(&) F], Standley 19797 [(9) F]; Río Grande, Molina 2497 [(2) RU CosTA he PUNTARENAS: Jicaral, Orozco 406 [(9 ) F]. gen TORO: Changuinola Valley, region of Almirante, Cooper 349 [( 2 ) CG REUS YU, © SCENE n. [(2) F, US, YU], 4 [(2) US, YU], 4a [(9 ) US, YU], 326 [(4) F, YU]. Dune 16 [(9) F] 40 [(9) F]; Seibert 1581 [( MO, US], 1583 [(?) MO, : ui Lagoon, Water Valley, [(9) MO, US]; Old ? v Bank Island, Chiriqui re von Wedel 2075 [(?) MO, SU]; Rio Cricamola, between Viii St. Louis and Konkintoé, Woodson, Allen & Seibert dé [(2) F, MO]; wirHout Liry; von Wede | "196 ( MO]. cawar zone: Barro Colorado Island, Bailey & Bailey 669 [(9) El; Shattuck 827 [(?) F}, 1164 [(9) FI; Standley 41080 [(2) US]; around Culebra, Pittier 2255 [(9) US], 3627 [(9) US]; around Gamboa, Pittier 6637 [( 2) US], 6652 [(2) US]; near Gatun, Standley 27287 [Q) US]; vicinity of Fort Sher- ) Standley 30235 [(?) US], 30296 [(?) US]. varıen: between Payo and Pa as Letras, Stern et al. 21 OJ]. panamá: Taboga, Hayes 658 [(8) BM]; eee Sta., Trinidad River, Pitter 6636 [(&) C, P, US]. WITHOUT PRECISE LOCALITY: Western Panama, Stork 16 [(9 lc. TROPHIS RACEMOSA ssp. RACEMOSA Leaf blades elliptic to oblong or ovate, (6.0-) 8.0-18.5 cm. long, 3.0-8.5 cm. wide, acuminate, the acumen usually short and gradually tapering from a broad base, obtuse or occasionally acute at the base, chartaceous to pergamentaceous, smooth on both surfaces, glabrous, the margin entire or occasionally with several deep sinuses. Staminate inflorescences: 3.7-7.5 cm. long, the staminate flowers numerous and densely crowded. Pistillate inflorescences: 0.9-3.8 cm. long, the rachis 0-1.8 mm. in diameter, the flowers usually distant along the rachis. The absence of this subspecies in the Lesser Antilles and its resemblance to ssp. ramon suggest that the West Indian plants originally migrated from Central Amer- ica. Ssp. racemosa is quite uniform in its pattern of variation and I have been unable to detect any insular differentiation. 1962] BURGER—STUDIES IN MORACEAE 13 Hillsides and forests from sea level to 1000 meters altitude; flowering from September through March. Vernacular names: Ramön (Cuba, Haiti, and Jamaica); Ramoon (Jamaica- Harris); Ramön de bestia (Cuba-Ekman). UBA: CAMAGUEY: La Gloria to Columbia, Shafer 617 [(9) F]. Hasana: near Villa Real, Ekman 532 [(?) S]. ras virras: Soledad, Cienfuegos, Jack 5002 [(9 ) P, S], 5842 H . ORIENTE: Bayate, Ekman 2981 [(4) S]. 4540 [(?) S], 4594 [(9) S], 7665 [(4) S], 9005 [(4) F, S, (?) S]; Baracoa, Ekman 4200 [(?) S]; Sierra de Nipe, at Río Piloto, Ekman 6687 [(9 ) S]; La Ermita, Hioram 4931 [( 4 icini Shafer 3272 [(9) F]. Without precise locality, Wright 589 [BM (8), S (9 6], 589 & 592 [MO (9), S (9 8)] 592 [(9) F, MO, S], sn. [(9) MO]. Par er Rio: Morillo, woods bordering the Manglares, Ekman 17430 [(?) S]; Las Martinas to the coast, Shafer 11078 [(9) F, MO]. WITHOUT PRECISE LOCALITY: Sagra 490 [(9 ) P]. Harr: ouest: Massif du Nord, Marmelade, Ekman 8285 [(9 ) S]; Massif de la Selle, Papette, Ekman 9387 [( 4) S]. AMAICA: CORNWALL: Trelawny: vicinity of Troy, Perkins 1005 [(9) B]; vicinity of Burnt Hill, Cockpit Country, Proctor 20588 [(9) BM]. Westmoreland: Kentucky Hill, Harris 10211 [(&) BM, F]. mippresex: Clarendon: Tom's Cave Wood, Harris 10858 [(9 . SURREY: Portland, John Crow Mts., SW. of Ecclesdown, Howard, Proctor & Stearn 14782 [(9) BM]. WITHOUT PRECISE LOCALITY: Cedarhurst, Harris 6075 [(9 ) BM, C]; Hall’s Delight, Harris 5550 [(9 ) BM, F]. Puerto Rico: ArEcıBo: Utuado, Buysman s.n. (1877) [(2) MICH], Sintenis 6309 [(9) BM, C, F, MO, S]. san JUAN: Candelaria, near Bayamón, Britton, Hess & Marble 2853 [(9 ) F, MO]; prope Naranjito, Stahl 862 [(9 ) S]. REPUBLICA DOMINICANA: BARAHONA: Fuertes 1086 [(4) BM, F. S]. SAN PEDRO DE macoris: Rose, Fitch & Russell 3719 [(9 ) F]; Consuelo, Taylor 146 [(2) F], 229 [(4) F]. WITHOUT PRECISE LOCALITY: Poiteau, s.n. (ca. 1882) [(9) P]; prope Puerto Plata, Eggers 1769 [(8) S]. West INDIES: WITHOUT PRECISE LOCALITY: Swartz s.n. [(4 ) S]. 2. TROPHIS MEXICANA (Liebm.) Bur. in DC. Prodr. 17:253. 1873. Sorocea mexicana Liebm. in Kon. Danske Vidensk. Selsk. Skr. 5. 2: 335. 1851. [T.: Lieb- mann 14260 (2) (C)]. Bucephalon mexicanum (Liebm.) O. Ktze. Rev. Gen. 2: 624. 1891. Trophis chorizantha Standl. in Field Mus. Pub. Bot. 4: 302. 1929. [T.: Standley 56771 (F) Skutchia caudata Pax & Hoffm. in Journ. Wash. Acad. Sei. 27: 307. 1937. [T.: Skutch 2383 Trophis St desi in Lloydia 2: 81. 1939. [T.: Matuda 2091 (MICH) (9)]. Shrubs or trees to 20 m. tall; leafy branchlets slender, 1.5-2.6(-3.2) mm. in diameter, glabrescent or infrequently puberulent, the lenticels small and inconspic- uous. Leaf blades elliptic to oblong or occasionally obovate, 8-20 cm. long, 2.5-6.5 cm. broad, acuminate, acute to obtuse at the base, firmly membranaceous to char- taceous, smooth on both surfaces, glabrous above and below or rarely sparsely puberulent below, the midvein plane above, prominent below with 4-9 pairs of ascending secondary veins arching upward near the bluntly serrulate or entire margin; petiole 3-15 mm. long, 0.7-1.5 mm. in diameter, glabrescent. Stipules caducous or occasionally persistent, lanceolate, 2.0-3.6 mm. long, minutely puberu- lent, their scars small and inconspicuous. Staminate inflorescences: spicate, solitary or less often paired in the axils of leaves or fallen leaves 4-11 cm. long and about 1 cm. in diameter at anthesis, the peduncle 3-12 mm. long, minutely puberulent, the rachis with scattered triangular or suborbicular, basally attached or rarely peltate bracts 0.4-0.8 mm. broad; flowers white or greenish, 13 to many and some- [Vor. 49 14 ANNALS OF THE MISSOURI BOTANICAL GARDEN what distant though appearing crowded at anthesis, sessile or rarely subsessile, about 4 mm. long and 4 mm. wide, the perianth parts 1.5-2.6 mm. long, 0.5-1.3 mm. wide, acute at the apex, connate (0.5-1.3 mm.) toward the base, minutely puber- ulent, the trichomes less than 0.1 mm. long; staminal filaments 2.0-3.2 mm long, the anthers 0.6-1.2 mm. long; pistillode prominent about 0.6 mm. long. Pistillate inflorescences: racemose, solitary or occasionally paired in the axils of leaves or fallen leaves, (2-)4-10 cm. long, the peduncles 4-25 mm. long, densely puberulent, the rachis with scattered triangular or suborbicular, basally attached or rarely pel- tate bracts 0.5-1.1 mm. broad; flowers (4-)6-22 and distant, the pedicels (0-)0.5-2.3 mm. long and elongating in fruit, the perianth tube minutely 4-lobed at the apex, ovoid, 1.8-5.0 mm. long, 1.2-2.8 mm. in diameter, sparsely puberulent, the ovary half-inferior, with style branches 2.8-4.5 mm. long and minutely puberulent. Fruit globose, about 5-7 mm. in diameter, strongly ridged or ridged and tuberculate, sparsely puberulent to glabrescent, becoming red, the fruiting pedicels 2.0-6.0(-13) mm. long or the fruit occasionally subsessile. Plants of Chiapas with subsessile fruit and broad glabrescent leaves have been segregated as T. matudai. These specimens resemble T. racemosa and are the basis for my impression that T. mexicana represents a first divergence from T. racemosa. If this is true, T. mexicana probably represents the ancestral stock from which closely related T. chiapensis and T. cuspidata were derived. Central Mexico to Costa Rica, in moist forests and along water courses; flower- ing from October to March, from sea level to 2000 m. elevation. Vernacular names: Sweet breadnut, Masico dulce (British Honduras-Gentle) ; Ramön blanca a See EES Raspa-lengua (El Salvador-Standley). Mexico: pare do, Escuintla, Matuda de^ AO MICH, der 0576 iv MICH, NY, US], 2 MO D F, MICH, NY, US], 2648 [(9) A, MICH, NY], 397 [(8) F MICH, Vos NY], 3979 [(9) A, MICH, NY], ee [(4) A, MICH, MO, So 16157 [(&) E MICH]. Jarisco: gea Quadrilla to San Sebastian, Mexia 1701 [(&) BM, 2 MI OAXACA: Cafetal Nueva Esperanza utla, Conzatti, Reko & Makrin 3064 [(9) US]; Cafetal Concordia, Morton & Malin 2478 (9) A, ES USE Cafetal Calvario, Cerro Espino, Reko 3519 [US (4) not (9)]. vera cruz: Haci enda de Jovo. Liebmann 14260 [(9 ) F, photo C]. British HoNDURAS: EL cayo: Va un Lundell 6422 [(9) F, MICHI. STANN ee Creek valley, Gentle 3211 [(&) A, MICH, MO, NY]. roLevo: Temash River, Schipp 1334 [(9 ) A, BM, F, GH, MICH, MO, NY, S]. WITHOUT DEFINITE E LOCALITY: Pueblo Viejo, Schipp 1261 [(2) A, "BM, F, GH, MICH, Ma ie S]. UATEMALA: ALTA VERAPAZ: betwee and San Pedro Carchá, Standley 89897 [(?) F]; Pansamalá, von Turckheim 1231 UTE E MO, P, US]. cHiMALTENANGO: slopes of Volcán Fuego, Steyermark 52069 [(?) F]. ESCUINTLA: Aquilar 1645 [(&) F]; Río Guacalate, Standley 60197 [(9 ) A, F, MICH, NY]; below Las Lajas, Standley e [(?) F], 64815 [(?) F]; Rio Burriön, ‘Standley 89574 [(?) F]. Gapat: Río Juyamá, SE. of Cheyenne, . A 39157 [(9) F]; Río Dulce, near Livingston, Steyermark 30451 ICS) E, NTI. tween Finca Yalpemech and Chinajá, Steyermark 45445 [(9) F, NY, S]. QuE- an Colomba, Skutch 1986 [(&) A, BM, F, US]; between Finca Pirineos and Patzulín, Standley 86697 [(?) F]; slopes of Volcán Santa María, Steyermark 33499 [(9 ) F]. SAN MARCOS: slopes ud Volcán Tajumulco, Steyermark 37426 [(&) F], 37576 [(4) F]. SANTA ROSA: Cerro Redondo, Heyde & get 6239 [(8) GH, MO, US]. sucHITEPEQUEZ: Finca Moca, Skutch py K 9) A, BM, F, NY, US]. E Lus R: AHUACHÁPAN: Finca Coline. Standley 20183 [(9 ) GH, NY, US]. erer ar Tela, Standley 56771 [(2) E US]. COMAYAGUA: above El Moses Vouk: Dawson, Y Youse 6171 [(9) F, MICH, MO, NY, S, US]. MORAZÁN 1962] BURGER—STUDIES IN MORACEAE 15 Figure 2. Trophis mexicana: from Skutch 2386 (1), Skutch 1986 (2), Schipp 1261 (3), and Steyermark 45945 (4). [Vor. 49 16 ANNALS OF THE MISSOURI BOTANICAL GARDEN n and near Mt. Uyuca: duet 2445 [(9 ) MICH], s 909 [(9 ) F, GH, MO, US], 1108a [(9) F, MO, US], 1300 [(2) F. GH, MO], 1443 [(9) F, MO], Pfeifer 1423 [(9 ) MOI, Standley 12431 [(2) F], 20680 [(2) F], L. O. Williams 15799 [(9) F, LIL, MO, PA n F, ae MO, US], Williams & Molina 5 [(9) F, GH] 15601 [( 3 TEGUCIGALPA: San Juancito, Edwards P-43 [(?) oe . voro: 10 mi. south of Progreso, Record & Saken H.40 [(&) GH, NY, US]. NICARAGUA: JINOTEGA: Finca Aventina, Standley 9965 [(?) F]. eer Bach: ALAJUELA: Zarcero, Alfaro Ruiz, A. T H1558 Ié) F, MO, NY]. UANA 1 Arenal, Standley & Valerio 45271 [(4) US]. san José: vicinity of El General. Skutch 2383 [(2) A, MO, NY, S], 2386 [(4) A, Zub MO, US, S]. 3. TROPHIS CHIAPENsIS Brandeg. in Univ. Calif. Pub. Bot. 6: 178. 1915. [T.: Purpus 7091 (UC) (9)] Trophis nubium Standl. in Field Mus. Pub. Bot. 22: 17. 1940. [T.: Skutch 925 (F) (2)] Shrubs or trees to 15 m. tall; leafy branchlets slender 0.8-1.8 (-2.6) mm. in diameter, minutely puberulent and glabrescent, the lenticels small and inconspicu- ous. Leaf blades narrowly oblong-lanceolate to linear-lanceolate or narrowly ellip- tic, 5-16 cm. long, 1.2-3.0(-3.8) cm. broad, gradually tapering to the long-acumi- nate apex, the acumen 1.0-2.5 mm. wide and 5-23 mm. long, acute at the base and often slightly oblique, membranaceous to chartaceous, smooth on both surfaces, glabrous above and below, midvein plane or slightly impressed above, prominent below, with 5-11 pairs of ascending secondary veins arching upward near the serrulate margin; petiole 3.8-12 mm. long, 0.5-1.3 mm. in diameter, sparsely and minutely puberulent, glabrescent. Stipules persistent or caducous, narrowly deltoid to lanceolate, 1.2-3.3 mm. long, minutely puberulent or glabrescent, their scars small and inconspicuous. Staminate inflorescences: racemose, solitary in the axils of leaves or fallen leaves, 1.8-4.2 cm. long, the peduncle 2.4-9.0 mm. long, the rachis with triangular or suborbicular, basally attached bracts 0.4-0.8 mm. broad; flowers white to lavender, 10-35 and usually distant along the rachis, about 5 mm. long and 4 mm. wide, pedicels 0.4-2.8 mm. long, the perianth parts 2.0-2.8 mm. long, the lobes 0.8-1.4 mm. wide, connate by about 25-50% (0.5-1.3 mm.) of their length, minutely puberulent, the trichomes about 0.1 mm. long; staminal filaments 2.0-3.2 mm. long, the anthers 0.8-1.2 mm. long; pistillode prominent, 0.6-0.8 mm. long. Pistillate inflorescences: racemose, solitary in the axils of leaves or fallen leaves, 9-32 mm. long, elongating in fruit, the peduncles 2-7-11.0 mm. long, minutely puberulent, the rachis with numerous triangular or suborbicular, basally attached bracts; flowers 3-14 and distant along the rachis, pedicels (0.3-)0.8-3.0 mm. long, elongating in fruit, the perianth tube 4-lobed at the apex, narrowly ovoid, 1.2-2.6 mm. long, 0.8-1.5 mm. in diameter, sparsely appressed-puberulent, the vesture greenish becoming cinereous; ovary superior to half-inferior, the style branches 2.0-4.2 mm. long. Fruit globose, about 6-8 mm. in diameter, conspicuously tuberculate, glabrate, the fruiting pedicels 3-14 mm. long. The consistent correlation of narrow leaves and short racemose inflorescences distinguishes this species which ranges from central Mexico to Guatemala. Central Mexico to Guatemala, in moist forests at altitudes from 800 to 2600 meters; flowering from May to June and October to December. 1962] BURGER—STUDIES IN MORACEAE 17 Figure 3. Trophis chiapensis: from Matuda 3933 (1: 12x), Skutch 1511 (2), and Matuda 4241 (3). [Vor. 49 18 ANNALS OF THE MISSOURI BOTANICAL GARDEN Vernacular name: Cereza de montafia (Guatemala-Steyermark). Mexico: cHıapas: Mt. Ovando, Matuda 3933 [(9 ) A, MICH, MO, US]; near Escuintla, Matuda 4241 Léi A, F, MO, US], 15989 [( 4) F, MICH]; Cerro del Boquerón, Pur pus 7091 [C9 )- F, O, US]. GUERRERO: Montes de Oca, Hinton 10298 [(&) GH, US]. vera CRUZ: Orizaba, Bote eds [C8 ) ~~ GH, P]; Palo Dulce, Mell 685 [( 3) F, NY, US]; Jalapa, Schiede & D s.n. [(?) BM, (8) F]. wrrHouT ee, LOCALITY: Sierra San Pedro Nolasco, T. UNE 941 te &) BM]; Mohr 465 [( 2) U UATEMALA: QUEZALTENANGO: Volcán Zunil, Skutch 925 UPS F, MICH, NY, US]; r Santa María de Jesüs, Standley 84572 [(9 ) F, S]; between Finca Pirineos and Patzulín, Standley 86808 [(?) F], "86994 LO) F], 87012 [(9) F]; Volcán Santa Pa Steyermark 33636 [(?) F]. san marcos: Volcán Atitlán, Skutch 1511 [(9 ) A, BM, F, NY, US]: near Rodeo, Standley 68956 [(?) F]. sororá: Volcan Atitlán, Steyermark 47833 E) Bi, 47349 [(&) F], 47392 [(?) F, US]. sucurrepéguez: Volcán Santa Clara, Steyermark 4663 [(2) F], 46658 [(?) F], 46687 [( 4) F, S], 46730 [(9 ) F]. 4. TROPHIS CUSPIDATA Lundell, in Amer. Midl. Nat. 19:427. 1938. [T.: Matuda 1051 (MICH) (8)] Shrubs or trees to 12 m. tall, the trunk to 30 cm. in diameter; leafy branchlets usually stout, 1.8-4.0 mm. in diameter, puberulent and glabrescent, prominently lenticellate. Leaf blades oblong to elliptic-oblong or elliptic, 10-25 cm. long, m. broad, narrowly caudate-acuminate, the acumen 1.2-3.0 mm. wide and 1.4-2.8 cm. long, acute to obtuse at the base, firmly membranaceous to chartaceous, smooth and glabrous above, short pilose beneath, the midvein slightly impressed above, prominent below with 7-13 pairs of ascending secondary veins arching upward near the entire margin; petiole 10-24 mm. long, 1.0-2.2 mm. in diameter, densely puberulent. Stipules persistent or caducous, lanceolate, 2.5-3.8 mm. long, their scars often conspicuous. Staminate inflorescences: solitary in the leaf axils, 2-4 cm. long (immature), the perianth parts densely hirtellous. Pistillate inflores- cences: racemose, solitary in the axils of leaves or fallen leaves, 2.8-8.0(-12)cm. long, elongating in fruit, the peduncles 7-22 mm. long, densely puberulent, the trichomes about 0.2 mm. long, the rachis with scattered triangular or suborbicular, basally attached bracts 0.5-0.9 mm. broad; flowers 12-30 and distant, pedicels 0.8-3.0 mm. long, elongating in fruit, the perianth tube minutely 4-lobed at the apex, ellipsoid to obovoid, 2.0-4.5 mm. long, 1.5-2.6 mm. in diameter, densely appressed- puberulent, ferruginous or becoming so; ovary half-inferior to inferior, the style branches 2.0-3.7 mm. long, densely and minutely puberulent. Mature fruit not seen. Characterized by a dense tomentum on floral and vegetative parts and a nearly inferior ovary, this species is closely related to T. mexicana. Southern Mexico and probably extending into Guatemala, at altiudes from 2000 to pec meters; flowering in December. EXICO: CHIAPAS: Mt. Ovando, Matuda 1051 [( 4) MICH, MO]; Mt. Tacaná, Matuda 2370 ms A, F. MICH, NY]; Pinabeto, Motozintla, Matuda 15478 [(9 ) F]. ExcLUDED SPECIES Trophis aurantiaca Herzog, in Fedde, Rep. Nov. Spec. 7: 51. 1909.— Olmedia sp. Trophis Hilariana Cazar. Nov. Strip. Bras. Dec. 80. 1845.=Sorocea guilleminiana Gaud. Trophis macrostachya Donn. Sm. in Bot. Gaz. 40: 10. 1905=Sorocea pubivena Hemsl. All the Old World species proposed for Trophis are excluded following Bureau in DC. Prodromus 17:254. 1873. 1051 1962] BURGER—STUDIES IN MORACEAE 19 aue 7 y e ^ i j SN DAE ? SNO SR M SR, SRI, Pe OA ls a Ae, d d 1 Figure 4. Trophis cuspidata: from Matuda 2370 (1), Matuda 15478 (2), and Matuda (3). [Vor. 49 20 ANNALS OF THE MISSOURI BOTANICAL GARDEN I. CLARISIA R. & P. Crarisia R. & P. Fl. Per. 128. t 28. 1794, nom. conserv., Clarisia Abat. [T.: C. race- mosa R. & P.] Geier deeg Liebm. in Kon. Danske Vidensk. Selsk. Skr. 5. 2: 316. 1851. [T.: S. mexicana iebm. Soaresia, Fr. Allem. in Rev. Brazil. 1: 210: 1857, non Soaresia Sch. Bip. [T.: S. nitida Fr. Allem.] Dioecious, laticiferous, unarmed shrubs and trees. Leaves alternate, distichous, bistipulate, pinnately veined, the petioles sulcate above. Stipules lateral, caducous or rarely persistent, their scars encircling less than half the stem. Inflorescences axillary or cauliflorous on the older branchlets; staminate inflorescences spicate with bracts and stamens usually lacking along a narrow line along one or two sides, paired in the axils of leaves or fallen leaves or borne in alternate pairs along a leafless racemiform shoot. Staminate flowers reduced to single stamens arising directly from the rachis, interspersed among numerous bracts, the stamens in well or- dered rows or paired but usually irregularly placed, with filaments short and slender, anthers subbasifixed, dehiscing longitudinally. Pistillate flowers pedicellate, with 3-7 suborbicular peltate bracts at the base of the receptacle; perianth tubular, adnate to the inferior ovary; pistil 2-carpellate and 1-locular, the ovule solitary, anatropous and pendulous, the style deeply and narrowly bifid with the stigmatic surface mi- nutely papillate. Fruit drupaceous, the perianth tube accrescent and succulent, the stone ellipsoid, without endosperm, the embryo large with thick equal cotyledons and minute hypocotyl. Clarisia was revised by Lanjouw in 1936 and many recent collections have been identified according to his revision. The accumulation of new collections with the frequent misplacement of specimens of Trophis and Sorocea made necessary an examination of material filed under Clarisia. In studying this material I have come to conclusions differing from those of Lanjouw though fully cognizant of the con- tribution his study has made. Lanjouw clarified the nomenclatural problems asso- ciated with the name Clarisia, which led to the conservation of the genus of Ruiz and Pavón. The major differences between my conclusions and those of Lanjouw are, first, that I have separated the species with capitate pistillate inflorescences as a distinct genus (Acanthinophyllum), and secondly, that I interpret C. biflora as a polymorphic species of wide distribution. The genus defined in this manner is composed of but two species. Clarisia presents an interesting sequence in the development of a racemiform inflorescence. The basic condition is probably best illustrated by the pistillate flowers of C. biflora. These are usually paired in the axils of leaves but in some specimens they arise in alternating pairs along a shoot that terminates in one to several leaves. In occasional specimens of C. biflora the flowering shoots with alternating pairs of flowers are completely leafless [Allen 3741, Weberbauer 7057 (GH)]. The cauliflorous *racemes" of C. racemosa with alternating pairs of pistillate flowers or staminate spikes may thus be interpreted as leafless flowering shoots. The genus extends from central Mexico to Bolivia and eastern Brazil. The trees often reach 30 meters in height and perhaps for this reason are infrequently col- ected. 1962] BURGER—STUDIES IN MORACEAE 21 KEY TO THE SPECIES a. Inflorescences axillary on the leafy branchlets; pistillate flowers in axillary pairs o afless racemiform shoots; leaves wi -12 pairs of ascending econdary veins. Mexico to Bolivia and the upper Amazon basin ............ 1. C. BIFLORA aa. Inflorescences cauliflorous on the older branchlets; pistillate flowers on long leafless racemiform shoots; leaves with 10-18 pairs of subhorizontal secondary veins. Ecuador to Bolivia and eastern Brazil 2. C. RACEMOSA CLARISIA BIFLORA R. & P. Syst. Veg. Fl. Peruv. et Chil. 255. 1798. [T.: Ruiz & Pavön s.n. in Herb Madrid] Shrubs or trees to 35 m. tall, the trunk to 1.2 m. in diameter with smooth brown bark, yellowish wood, and milky latex; leafy branchlets 2.0-4.5 mm. in diameter, sparsely appressed-puberulent, the lenticels small and inconspicuous. Leaf blades narrowly oblong, elliptic-oblong or broadly elliptic, 8-25 cm. long, 2.5-9.0 cm. broad, acuminate, acute to obtuse at the base, firmly membranaceous to stiffly chartaceous, glabrous above, glabrous or sparsely puberulent beneath, the midvein slightly impressed above, prominent below with (4-)6-12 pairs of ascending sec- ondary veins arching upward near the entire margin; petiole (4-)6-22 mm. long, 1.0-2.2 mm. in diameter, minutely and sparsely puberulent, glabrescent. Stipules caducous or rarely persisting, narrowly to broadly cuneate, 3.5-7.2 mm. long, their scars inconspicuous. Staminate inflorescences: spicate, paired in the axils of leaves or fallen leaves or in alternate pairs on leafless racemiform shoots, the primary peduncle 5-35 mm. long, minutely puberulent and bracteate only below the attach- ment of the secondary peduncles (peduncles of the spikes); spikes 2-10 cm. long, secondary peduncles 3-6 mm. long, sparsely puberulent, the rachis with numerous spatulate to broadly cuneate, basifixed or occasionally peltate bracts; stamens inter- spersed among the bracts, the filaments straight or sometimes bent below the anther, about 1.0 mm. long, anthers 0.4-0.8 mm. long, emarginate in Mexico, slightly apiculate in Central America, or the connective forming a small apical disc in South America. Pistillate flowers in pairs in the axils of leaves or fallen leaves or occasionally in alternate pairs along a leafless racemiform shoot, peduncles to 2 mm. long, the pedicels 0.5-6.0 mm. long, densely and minutely puberulent, with 3-7 peltate bracts 0.6-1.4 mm. broad at the base of the receptacle; perianth tube 2.2-5.0 mm. long, 1.1-5.0 mm. in diameter, narrowly ovoid to globose, glabrous except at the minutely 2-4 or irregularly lobed apex, style branches 2-6 mm. long. Fruit ovoid to ellipsoid, to 25 mm. long, green or yellowship, smooth and glabrous is species was interpreted as a group of four by Lanjouw in his revision of 1936. I believe that the additional material now available requires a broader cir- cumscription. The leaves are quite variable and isolated collections have given the impression that this variation is correlated with geography. However, plants with narrow elliptic leaves have now been collected in Costa Rica, Colombia, Peru, Bolivia, and Brazil while broadly elliptic leaves were collected by Ruiz and Pavön in Peru and recently in Panama. The variation in inflorescences likewise is not worthy of taxonomic recognition. The same collection, for example, may have solitary or paired staminate spikes in the axils of leaves [Skutch 2004 (US)] or leafless racemiform shoots of alternating pairs of spikes [Skutch 2004 (F)]. — H [Vor. 49 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN The anthers of the only staminate collection from Central American (Skutch 2004) possess a slight apical enation that is lacking in specimens from Mexico. I suspect that further collections from Central America will exhibit a transition to the definite disc-like structure of South American specimens. This character of the anthers together with differences in leaf form justifies, I believe, the recognition of two subspecies. The geographical demarcation of these subspecies, however, can only be provisional due to the lack of staminate collections between Guatemala and Colombia. KEY TO THE SUBSPECIES a. Anthers emarginate or with the connective only slightly expanded at the apex of the thecae; pistillate flowers globose at anthesis; pedicels 0.5-3.0 mm. long; leaves narrowly oblong to elliptic-oblong. Mexico and Guatemala ......la. ssp. MEXICANA aa. Anthers with a definite disc at their apex; pistillate flowers narrowly ovoid to globose at anthesis, pedicels 1-6 mm. long; et ot elliptic to obovate o elliptic oblong. Costa Rica to Brazil and Boliv Ai BIFLORA la. CLARISIA BIFLORA ssp. mexicana (Liebm.) W. Burger, comb. & stat. nov. Sahagunia mexicana Liebm. in Kon. Danske Vidensk. Selsk. 5°: 316. [T.: Liebmann 14259 (C) (8)] Clarisia mexicana (Liebm.) Lanj. in Rec. Trav. Bot. Néerl. 33: 270. 1936. Leaves narrowly oblong to elliptic-oblong or occasionally narrowly obovate, 8-22 cm. long, 2.5-4.5(-7.0) cm. broad, acuminate, acute to obtuse at the base, the secondary veins similar to the lower leaf surface in pigmentation. Staminate spikes to 4.5 cm. long, the anthers eglandular or with a small projection at the apex. Pis- tillate pedicels 0.5-3.0 mm. long, the flowers globose at anthesis. Fruit not seen. F gë from December through March. XICO: CHIAPAS: Escuintla, Matuda 4023 [(8) MICH, rae 17439 [(9) F]. vera CRU! kr ears: de Tlatetla, E. of Mirador, Liebmann 14259 [( 8 ) Fl; poe Lieb- mann 14273 [( 2) F]; aei re Purpus 5989 [(9 ) US], 6492 aor GUATEMALA: QUEZALTENANGO: Colomba, Skutch 2003 [(9) BM, Ay Tr 2004 [( 2) BM, F, US]. lb. CLARISIA BIFLORA ssp. BIFLORA Sahagunia colombiana Rusby, Descr. New Sp. S. Am. Pl. 10. 1920. [T.: H. H. Smith 424 (NY : Clarisia colombiana (Rusby) Lanj. in Rec. Trav. Bot. Néerl. 33: 266. 1936. Clarisia mattogrossensis Lanj. loc. cit. 268. 1936. [T.: Gë M ip (8)]. Clarisia panamensis Woods. in Ann. Missouri Bot. Gard. 4 1960. [T.: Allen 3741 (MO) (9 Leaves Re elliptic to obovate or elliptic-oblong, 7-24 cm. long, 3.2-8.5 cm. broad, often abruptly acuminate, obtuse or occasionally acute at the base, the secon- dary veins often darkly pigmented. Staminate spikes to 8.5 cm. long, the anthers with a conspicuous gland-like disc at their apex. Pistillate pedicels 1-6 mm. long, the flowers narrowly ovoid to globose at anthesis. Fruit ellipsoid to narrowly oblate, becoming yellowish, with thickened style branches. The differences in stamen size used by Lanjouw to distinguish C. colombiana from C. mattogrossensis appear to be due to individual variation and stage of development. Flowering from December to February and June through September at alti- tudes to 2100 meters. 1962] BURGER—STUDIES IN MORACEAE 93 Figure 5. Clarisia biflora. ssp. mexicana: from Liebmann H. H. Smith 424 (2), Klug 3698 (3), and Weberbauer 7057 (4) 14259 (1). ssp. biflora from [Vor. 49 24 ANNALS OF THE MISSOURI BOTANICAL GARDEN Vernacular names: Lechero colorado (Colombia-Garcia-Barriga) ; Leche caspi, Cuchara caspi (Peru-Ll. Williams). Costa Rica: PUNTARENAS: Palmar Norte de Osa, Allen 5955 [(9 ) F, US]. PANAMÁ: COCLÉ; N. eR El Valle de Antón, Allen 3741 [(9 ) BM, F, G, MO]. COLOMBIA: CUNDIN. CA: between San Francisco and Subachoque, Garcia.Barriga Pici [(4) US]. santa marta: Don Amo Road, H. H. Smith 424 [(8) F, NY, USI- LLE DEL CAUCA: Cordillera Occidental, Rio Sanguinini, Cuatrecasas 15487 [(9) F], 15493 [(4) Fl. PERU: LIBERTAD: Pataz, below Ongón, Weberbauer, 7057 [(2) F, GH]. Loreto: Caballo- cha, Amazon River, LI William 2086 [(9) F]. san MARTÍN: Zepelacio, near Moyobama, B H MO, NY, S]. Lamas, near Tarapoto, LI. ies 6471 . SANTA ROSA: Urubamba Valley, Cook & Gilbert 1719 [(8) F, US]. wırHouT PRECISE LOCALITY: Ruiz & Pavón 24 26 [(9) F], 24/28 [(9) F]. from Río Ucayli, cult. in Hort. aisi m Brazil I" Borr San Bartolomé, Krukoff 10410 [(9) F, LIL, MO, S], Larecja, Tuiri. Kru Eo "Gei A, F. LIL, MO, S], 10873 [( 3) F, LIL, MO]. L: ACRE: aulo, Lewin 2076/w17 [(?) LIL]. matro Grosso: Tombador, on Rio See d eps 131 [(8) NY]. Pará: in Hort. Belém 2. CLARISIA RACEMOSA R. & P. Syst. Veg. F. Peruv. et Chil. 255. 1798 [T.: Ruiz & Pavón s.n., Chinchao et Pozuzo, in Herb Madrid. Soaresia nitida Fr. Allem. in Rev. Brazil 1: 210, cum tab. 1857; Arch. Palestr. Scient. Rio : 142. ic Olmedia erythrorhiza Huber, nom. ex Ducke in Arch. Jard. Bot. Rio Jan. 3: 40. 1922, pro syn. Clarisia nitida (Fr. Allem.) Macbr. in Field Mus. Pub. Bot. 11: 15. 1931. Trees to 40 m. tall, the trunks slender with rough brown bark, yellowish wood, and white latex; leafy branchlets 1.5-3.7 mm. in diameter, minutely puberulent, the lenticels small and inconspicuous. Leaf blades narrowly to broadly oblong or ovate, (4-)6-17 cm. long, (2.0-)3.5-7.0 cm. broad, acuminate, the base often rounded, subtruncate to obtuse or occasionally acute, thickly chartaceous to subcori- aceous, often lustrous above, glabrous, the midvein slightly impressed above, promi- nent below with 10-18 pairs of subhorizontal secondary veins, the margin entire or slightly undulate; petiole 4-16 mm. long, 0.7-1.8 mm. in diameter, sparsely puberu- lent and glabrescent. Stipules caducous or rarely persistent, broadly cuneate, 1.2-2.0 mm. long, their scars small and inconspicuous. Staminate inflorescences: cauliflo- rous, solitary or several arising together from the older branchlets, usually of alter- nate pairs of spikes along a leafless racemiform shoot, 6-15 cm. long, the priraary peduncle 5-43 mm. long, minutely puberulent or glabrescent, the primary rachis bracteate only below the attachment of the secondary peduncles (peduncles of the spikes); spikes 2-9 cm. long, secondary peduncles 2-10 mm. long, densely and minutely puberulent, the rachis of the spike with numerous spatulate to broadly cuneate, basifixed or occasionally peltate bracts; stamens interspersed among the bracts, the filaments about 1.0 mm. long, anthers about 0.5 mm. long, emarginate. Pistillate inflorescences: racemiform, cauliflorous on the leafless branchlets or rarely in the axils of leaves, solitary to several arising together, 6-14 cm. long (to 25 cm. in fruit), the 10-24 flowers usually borne in alternate pairs along a leafless race- miform shoot (inflorescence) ; peduncle 12-35 mm. long, the rachis glabrescent an bracteate only below the attachments of the pedicels; pedicels 2.4-4.5 mm. long (to 20 cm. in fruit), densely and minutely puberulent, occasionally with a bract 1962] BURGER—STUDIES IN MORACEAE 25 i isi /X/1935) Figure 6. Clarisia racemosa: from Ducke (30/V1/1942) 948 (1), Ducke (26 59 (2), Krukoff 5380 (3), Krukoff 10744 (4), and Krukoff 8312 (5). [Vor. 49 26 ANNALS OF THE MISSOURI BOTANICAL GARDEN or two along their length, with 3-5 suborbicular peltate bracts 0.7-1.5 mm. broad at the base of the receptacle; perianth tube narrowly to broadly ovoid, glabrous but appressed-puberulent at the irregularly lobed apex, 3.5-8.0 mm. long; style branches 5-10 mm. long. Fruit ellipsoid or somewhat asymmetric, to 35 mm. long and 14 mm. in diameter, orange becoming black, smooth and glabrous. The inflorescences show little of the variation found in C. biflora and appar- ently have become stabilized. An interesting exception is the British Museum specimen of Krukoff 1450 with two solitary, long-pedicelled pistillate flowers in the axils of leaves. Other specimens of the same collection possess the normal cauli- florous “racemes.” The disjunct plants of Rio de Janeiro exhibit no morphological distinctions that might be correlated with their isolation. Ecuador to Bolivia and eastern Brazil; in moist forests and the matta de terra firme and várzeas of the Amazon basin. Flowering from July to January. Vernacular names: Moral babo, Moral comida de mono, sota, zota (Ecuador- Little); Capinuri (Peru-Ll. Williams); Tulpay (Peru-Burgos); Guariuba (Peru and Brazil-Ll. Williams, Ducke, Krukoff, et al.). ECUADOR: ESMERALDAS: n" Quininde, Little 6219 [(?) F]. ERU: HUÁNU 42 Hu qtd ue Burgos 59 [(?) F]. JUNÍN: La dore eeh 1912 Tee 4) F]; Río Negro, Woytkowski 5865 [(9) MO]. LORETO: Ucayli, Tessmann 5433 [(4) S]; La Victoria, L E William ics A FL aa F) Alto Rio Itaya, LI. Wiliams 3380 [(?) F]. OUT PRECISE 24/24 AS? ) zm Bor : S. Yungas, Río Bopi, Krukoff 10533 [(4) F, LIL, MO, S], 10679 [(?) m Lares, Tuiri, Krukoff 10744 [(?) F, LIL, MO, S]. ACRE: mouth of e Macauhan, Krukoff 5380 [(?) A, BM, F, MO S]: Rio here, Ule “9317 [(&) MG]. AZONAS: Manaus, Ducke (26/X/1935) 59 [(&) A, F, MO, S]; basin of Rio Solimóes, a Oel 132 [(9) LIL]; São Gabriel, Rio Ak go Froes 410 to LIL], 852 [(?) LIL]; mouth of Rio Embira, Krukoff 5173 [(9) BM, F. MO, S]; Humayata, near Tres Casas, Krukoff 6327 [(9) BM, LIL, MO, S]; near icra Krukoff 1450 [(9) A Si; P yr Tapajos, "Baa Vista, Capucho 568 [(9) F]; Faro, Ducke (2/1/1920) er no ) RB, S]; Gurupá, Ducke CIA/VITI/1918) [(?) MG]; Rio Rapajoz, Sáo Luiz, Ducke 5601919] 13052 [(9 ) P, RB, S], (14/XII/1919) 13052 [(4) S]; Rio Trambetas, eech (1 re 16993 vx MG], EE bac d n) RB, SS]; Rio Xingu, Ducke (14/X/1916) 1 EnS : ens rane re 20489 = ` FAMULI 2 A Cosmo o Velho, e 8: 8289 no P]; Rio de Janeiro, Kuhlmann (VIII/1920) ines [RB (9 3), 8 (8)], We E [(9) S]. EXCLUDED SPECIES Clarisia ilicifolia (Spreng.)—Lanj. & Rossb. in Rec. GH Bot. Néerl. 33:717. 1936. =Acanthinophyllum ilicifolia (Spreng.) W. Bur Clarisia mollis Standl. in Ann. Missouri Bot. Gard. 30: e 194. — Sorocea pubi- vena Hemsl. Clarisia spruceana Lanj. in Rec. Trav. Bot. Néerl. 33:272. 1936. = Acanthino- phyllum spruceana (Lanj.) W. Burger. Clarisia strepitans (Fr. Allem.) Lanj. in Rec. Trav. Bot. Néerl. 33:272. 1936. = Acanthinophyllum ilicifolia (Spreng.) W. Burger. Clarisia urophylla (Donn. Sm.) Lanj. loc. cit. 270. 1936. =Trophis racemosa (L.) Urb, 1962] BURGER—STUDIES IN MORACEAE 21 III. ACANTHINOPHYLLUM Fr. Allem ACANTHINOPHYLLUM Fr. Allem. in Rev. Brazil. 1:368, 1858; Arch. Palestr. Scient. Rio Jan. 1:215. 1858. [T.: A. strepitans Fr. Allem., =A. ilicifolia (Spreng.) W. Burger] Dioecious, laticiferous, unarmed shrubs and small trees. Leaves alternate, distichous, bistipulate, pinnately veined, the petioles sulcate above. Stipules lateral, caducous or persistent, their scars encircling less than half the stem. Inflorescences axillary or cauliflorous on the leafless branchlets, solitary or several clustered on leafless brachyblasts or rarely the staminate alternate along a short leafless racemi- form shoot, the staminate spicate with the bracts and stamens lacking along a narrow strip along one side, the pistillate capitate; peduncles densely invested with minute retrorse uncinate trichomes. Staminate flowers reduced to single stamens arising directly from the rachis and interspersed among numerous variously shaped bracts, occasionally in groups of 2-4; filaments short and slender, the anthers small and usually dorsifixed, dehiscing longitudinally. Pistillate flowers sessile on a short peduncled capitulum; perianth tubular, free or partially adnate to the superior or subinferior ovary; pistil 2-carpellate, l-locular, the ovule solitary, pendulous, and anatropous, the style shortly bifid with the inner stigmatic surface glabrous. Fruit drupaceous, the perianth tube accrescent and succulent, the stone globose to ellip- soid, without endosperm, the embryo large, straight, and erect with thick, usually equal cotyledons, the hypocotyl minute. The genus ranges from British Guiana to eastern Brazil and the Amazon basin of Venezuela, Brazil, and Bolivia. Acanthinophyllum is characterized by its staminate flowers reduced to single stamens and its pistillate flowers borne on a short peduncled capitulum. Liebmann described Sahagunia from staminate plants he had collected in Mexico. When Bentham and Hooker (1880) described the pistillate flowers of the genus Sahagunia they unfortunately drew their details from specimens of Acanthinophyllum, which is restricted to South America. The staminate spikes of Acanthinophyllum resemble those of Sahagunia but the dorsifixed anthers and pistillate capitula readily distin- guish the two genera. The retrorse uncinate trichomes present on leaves, twigs, and especially peduncles of Acanthinophyllum are not found in Clarisia (an earlier name for Sahagunia). In addition to the trichomes the midvein of the leaves is prominulous above and the epidermal cells have a deeply sinuate outline. These vegetative characters are shared, to my knowledge, only by Trymatococcus and Helianthostylis among South American Moraceae. The genus is represented by two species. Key TO THE SPECIES a. Staminate spikes condensed, to 2.0 cm. long; pistillate inflorenscences of 8-15 closely contiguous flowers; che spinulose-serrate to entire. British Guiana to iA Brazil and Bolivia ILICIFOLIA aa. Vm spikes lax, to 6.5 cm. long; pistillate flowers not seen; leaves entire. nezuela: Amazonas 2. A. SPRUCEANA T ea ilicifolia (Spreng.) W. Burger, comb. n Excoecaria ilicifolia Spreng. Neue Entdeck. 2: 117. 1821. [T.: Wind Collector, "in Brasilia,” in Herb. Berlin, (4)] [Vor. 49 28 ANNALS OF THE MISSOURI BOTANICAL GARDEN Acanthinophyllum strepitans Fr. Allem. in en m 1: 36, cum tab. 1858; and in Arch. da Palestr. Scient. Rio Jan. 1: 215. 1858. Pseudosorocea Poeppigii Baill. in Adansonia M 297. 1875. Sahagunia strepitans (Fr. Allem.) Benth. & Hook. Gen. Pl. 3: 377. 1889. Sahagunia racemifera Huber, in Bol. Mus. Para. 5: 334. 1908. [T.: Ducke (27/V11/1902) Sorocea stenophylla Standl. in Trop. Woods no. 43: 18. 1935. [T.: Capucho 447 (F) (9)] Clarisia strepitans (Fr. Allem.) Lanj. in Rec. Trav. Bot. Néerl. 33: 274. 1936. Clarisia strepitans var. guianensis Lanj. loc. cit. 274. 1936. [T.: em 120 (U) [(9)] Clarisia strepitans var. paraensis Lanj. loc. cit, 275. 1936. [T.: Huber 6965 (U) (9)] Clarisia strepitans var. micranthera Lanj. loc. cit. 275. 1936. [T.: Glaziou 12172 (K) (&)] Clarisia strepitans var. cuyunensis Lanj. loc. cit. 275. 1936. [T.: Forest Dept. 1029 (U) EA Clarisia ilicifolia (Spreng.) Lanj. & Bosch loc. cit. 717. 1936. Clarisia ilicifolia var. guianensis (Lanj.) Lanj. & Rossb. loc. cit. 718. 1936. Clarisia ilicifolia var. paraensis (Lanj.) Lanj. & Rossb. loc. cit. 1936. Clarisia ilicifolia var. micranthera (Lanj.) Lanj. & Rossb. loc. cit. 718. 1936. Clarisia ilicifolia var. cuyunensis (Lanj.) Lanj. & Rossb. loc. cit. 718. 1936. Shrubs or small trees to 8(-15) m. tall with white or yellowish latex; leafy branchlets sparsely invested with small retrorse uncinate trichomes, glabrescent, the lenticels small and inconspicuous. Leaf blades elliptic to elliptic-oblong, obovate or occasionally linear-lanceolate, 3-25 cm. long, 1.7-7.0(-9.0) cm. broad, acuminate to abruptly caudate-acuminate, the apex acute, acute to obtuse at the base, char- taceous to pergamentaceous, glabrous, the midvein prominulous above, prominent below with 6-14 pairs of ascending secondary veins arching upward near the spinu- lose-serrate to entire or occasionally deeply sinuate margin; petiole 5-17 mm. long, 0.7-2.5 mm. in diameter, glabrescent or sparsely invested with retrorse uncinate trichomes. Stipules caducous or persistent narrowly lanceolate, 1.0-1.8 mm. long their scars small and inconspicuous. Staminate inflorescences: spicate, axillary or cauliflorous on the older branchlets, paired, solitary, or clustered on leafless brachy- blasts or rarely on leafless racemiform shoots, 4-20 mm. long; peduncles 2-5 mm. long, densely invested with small retrorse uncinate trichomes, the rachis with numerous cuneate to suborbicular, basally attached or occasionally peltate bracts; stamens interspersed among the bracts, the filaments about 1 mm. long, the anthers 0.2-0.3 mm. long, occasionally paired or rarely in threes. Pistillate inflorescences: capitula of 8-15 closely contiguous flowers, solitary, paired or clustered on leafless brachyblasts in axils of leaves or cauliflorous, 10-18 mm. long at anthesis; peduncles -9 mm. long, densely invested with retrorse uncinate trichomes about 0.1 mm. long; flowers occasionally subtended by basally attached bracts, elliptic to obovoid or variously shaped by contact with adjacent flowers, sessile, the perianth tube 1.2-3.0 mm. long, 1.0-3.0 mm. in diameter, minutely hispidulous, the ovary superior to subinferior, tapering to the slender style, style branches broad, about 1-2 mm. long, glabrous. Fruit globose or ellipsoid, to 25 mm. long and 15 mm. in diameter, smooth and glabrous, orange at maturity. The leaves of this species are extremely variable, ranging from broadly elliptic to linear-lanceolate and from spinulose-serrate to entire. This variability is found 1962] BURGER—STUDIES IN MORACEAE Figure 7. Acanthinophyllum ilicifolia: from Ducke (2/IX 1923) 18326 (1 & 2: stamen X20), Krukoff 1669 (3), Krukoff 5991 (4), and Capucho 447 (5 & 6). [Vor. 49 30 ANNALS OF THE MISSOURI BOTANICAL GARDEN throughout the range of the species except for the very narrow-leaved specimens which have only been collected in Para. Standley’s Sorocea stenophylla (Plate 7, fig. 5) is based on a collection with linear-lanceolate leaves and may be worthy of varietal status. Collections from Amazonas have more ellipsoid fruit and fewer spinulose leaves than specimens from Para, but these characters are too inconstant to establish subspecies and I am not describing categories below that rank. Lan- jouw described four varieties of which var. guianensis with large broadly elliptic leaves appears to be most distinct; however I have not seen var. cuyunensis. Flowering from June to December in moist forest and the mata de terra firme of the Amazon basin. Vernacular names: Bainha de espada (Rio de Janeiro-Allemäo) ; Folha de serra (Minas Mrd an oe Janita (Para-Ducke). FRE ANA: Karoumany, Sagot 530 [BM (?), P (&), (?)]; Without precise —_ Poiteau s.n. 1844 [(&) B]. o Serra, Kuhlmann 571 [(ļ) oe crE: Rio Abunan, Kuhlmann Gäns 675 [(?) RB]. Amazonas: Calam Ma deles | River, ae 1309 Ke ) BM, MO, NY, P, S]; Borba, oe? Bella Vista, Krukoff 5991 [(?) S]; Manicoré, near S ée Fé, Krukoff 6049 [(2) BM, LIL, T e B Humayatá, near Livramento, koff 9 , BM HF Madeira River, Kuhlman x ran 257 Han RB, Sh C Viii) 2d CT) RB]; Mane Pires 11 " NY); Santarém, Ginzburger e 4 [C*3-EL APORÉ: i [C a M ara 3 M, F, M Ouro Preto, Kuhlmann ( Se "487 [( 9) R B]. MINAS GERAIS: Campo de Cementes, Magalhäes 843 [(9 ) US]; Barbado, Mexia 5158 [(&) A, BM, F, GH, MICH, MO, NY]; Lagóa Santa, E ae in. [0 81. vs Belterra, Black 473715 (2) NY], 47-1728 [@) E Tapajos, Bóa Vista, Capucho 447 [(9) F]; Obidos, Ducke (29/V1I/1902) 2885 [(&) M Ġ, RB]. (23/XII/1907) 9190 [(9) MG, RB], (28/XII/1913) 15270 [(9) BM, MG, RB], (13/1%/1926) 19473 [( 4 ) RB, S]; Alemquer, Ducke bec 4899 [( 4 ) BM, MG, RB]; Rio Cuminá, Ducke (13/X/1913) 14978 [(8) BM, MG, RB]; Monte Alegre, Ducke (24/IV/1916) "16059 [(&) MG]; Braganca, Ducke dg 18323 [(2) RB]; Rio Tapajoz, (22/V1II/1923) 18324 [(4) RB, S]; Serra de Santarém, Ducke (2/IX/1923) 18325 [(8) RB], 18326 [(&) RB, S]; Belém, Bosque Municipal, Ducke (10/VII/ 1945) 1726 [(&) A, F, MG]; Manaus, Francisco- INPA 2025 [(8) MG]; Moema, Herb. 9572 [(?) MG, RB]; Belém do Pará, Santa tie Huber 6894 Gi MG, RB]; ious : à : rBo 3) M 96 RB]; e y opidi e sab Be u 12172 [(8) B, C, E P]; S. José, Peckholt s.n. Nov 46 Te) B]. Glaziou 1470 [Q) C]; Castelnovo, Riede l s.n. Sept. 1822 [(?) US]; Salle e s.n. 1. 1815. 1817 C; (4) BM]. 2. ACANTHINOPHYLLUM spruceana (Lanj.) W. Burger, comb. nov. Clarisia Spruceana Lanj. in Rec. Trav. Bot. Néerl. 33: 272. 1936. [T.: Spruce 3279 (K) (8)] Small trees with slender glabrescent branchlets, the lenticels small and incon- spicuous. Leaf blades oblong to elliptic-oblong, 8-17(-21) cm. long, 3.2-6.7 cm. broad, acuminate, acute to slightly rounded at the base, chartaceous to pergamen- taceous, glabrous, the midvein prominulous above, prominent below with 7-13 pairs of ascending secondary veins arching upward near the entire margin to form an arcuate marginal vein; petiole 8-12 mm. long, 1.2-1.7 mm. in diameter, glabrescent 1962] BURGER—STUDIES IN MORACEAE 31 and rugose. Stipules caducous, their scars small and inconspicuous. Staminate inflorescences: spicate, 2-4 clustered leafless brachyblasts, 2.0-6.5 cm. long; penducle 4-10 mm. long, densely invested with retrorse uncinate trichomes, the rachis with numerous cuneate to suborbicular, basally attached or subpeltate bracts; stamens interspersed among the bracts or in groups of 2-4, the filaments about - 1 mm. long, somewhat flattened, anthers 0.2-0.3 mm. long. Pistillate flowers not seen, A. spruceana is quite distinct though represented solely by Spruce’s original collection. The isotype in the British Museum does not possess the rounded leaf base of the original description. VENEZUELA: AMAZONAS: ad flumina Casiquieri, Vasiva et Pacimoni, Spruce 3279 [(4) BM]. LITERATURE CITED Bentham, G., & J. D. Hooker. 1880. Urticaceae, in Genera Planetarum 3: 341-395. Bernbeck, F. 1932. Mee enam Morphologie der Urticaceen- und Moraceen-infloreszenzen. ot. Abh. ed. Goebel 19: 1-100. Browne, P. 1756. Civil = History of Jamaica. London, 503 p Bureau, E. 1873. Moraceae, in A. P. de Candolle, Prodromus ee Naturalis Regni Vegetabilis 17: 211-279. ege J. 1936. Studies in Moraceae II. The genus Clarisia Ruiz et Pavon and its onyms with a discussion of the generic name. Rec. Trav. Bot. Néerl. 33: 254-276. Trécul, A. 1847. Mémoire sur la famille des Artocarpées. Ann. ui Nat. Bot. III 8: 38-157. Woodson, R. E., Jr. 1960. Moraceae, in Woodson & Schery, Flora of Panama. Ann. Missouri Bot. Gard. 47: 114-178. ENUMERATION OF THE GENERA AND SPECIES I. TRropuis IL Crarisia l. racemosa Sos Urb. E — . & P. la. ssp. meridionalis (Bur.) p. mexicana (Liebm.) W. urger lb. ssp. ramon (Schlecht. & lb. ssp. biflora Cham.) W. Burger 2. racemosa R. & P lc. ssp. racemo 2. mexicana (Liebm.) Bur. III. ACANTHINOPHYLLUM 3. chiapensis Brandeg. l. ilicifolia (Spreng.) W. Burger 4. cuspidata Lundell 2. spruceana (Lanj.) W. Burger INpEx TO COMMON NAMES Bainha 2 espada III 1 Lechero colorado II 1b Breadnu II 1b Marfil I la C nit H2 H2 Capomillo I Ib Moral comida de mono ................—. II 2 Cereza de montaña 13 j I lb cox I lb Ojushte I lb Cuchara caspi II 1b mön Ilb Folha de serra IH 1 Ramön I2 Guariuba II 2 Ramón colorado I lb Hoja tinta I Ib de bestia I lb Janita II 1 oon I lb Leche caspi II 1b Raspa-lengua I2 [Vor. 49 32 ANNALS OF THE MISSOURI BOTANICAL GARDEN A H2 saani EEN Ib SC Breadnut lix du. -manchiäfl „nen I la Tillo prieto ....... IV 7 ite BR peer eee Seda Sate Peete iei ben I lb Lets 12 Yaxox A CAT Ujushte ....... I lb Dll i align ac ER II 2 INDEX TO EXSICCATAE Italicized numerals refer to collectors numbers, s.n. (sine numero) to unnum- bered collections; parenthetical numerals refer to the numerals of taxa conserved in this revision: the Roman to genera, and the Arabic to species; lower case letters refer to subspecies. Allen, P. H. 3741, See (II 1b). I 2). I la). Bailey, L. H. & E. Z. Bailey 669 (I lb). Baker, C. F. 21, 784 (I 1b Bartlett, H. H. 11867, 12354, 12375 (I 1b) Berlandier, J. L. 1 Black, G. A. 47-1715, 47-1728 (III 1). (I 1a); 4485 (I 1b). Brandegee, T. S. s.n. (I 1 b). "eitis N. „ W. E. Hess & D. W. Marble 285 c Burger, W. C. 148, 150 (I 1b). u 9 (II 2). . (I Ic). Calderón, * 845, 1065, 1448, 1698, 2178 ( Capachb, P. 447 (III 1); 568 (II 2). Carlson, M. 2445 ( Collenette, C. S. eg (II lb). Conzatti, C., B. P. Reko & E. Makrinius Cook, O. & G. B. Gilbert 1719 (II 1b) Cooper, G. P. 349 Cooper, G. G. M. Slater 4, 4a, 326 (I Grates J. 10840, e 16348, 16810 la); 15487, 15493 (II se n, H. M. 102, r 205. 372 (I 1a) Dressler, R. 2372 (I 1 ( GE 18324, (2/IX/1923) 18- 325, (2/IX/1923) 1 8326, (13/IX/ 1926) I uere 17204, (25/X/1919) 13- 54, (4/XII/1919) 13052, wiren Hei (2/1/1920) 13051, 59, (30/V1/1942) 948 (11 2 Ducke, A. & J. G G. Kuhlmann 6153, 19697 (III 1). Duke, J. 3636 (I 1b). | C. 16, 40 ( 1b). dwards, J. B. P-43 (I 2 P-644 (I lb). [rd ds, M. T. 667 (I 1b) ggers, H. 1769 (I 1c). Ekman, E. L. 532, 2981, 4200, 4540, 4594, 6687, 7665, 8285, 9005, 9387, 17430 (I c Emrick, G. M. 133 (I 1b). Fagerlind, F. G. Wibom 2376, 2403 (I la). Fernández, A. 310 vem E Flores, R. S. s.n. (I 1b). Froes, R. 132, 10, 852 (II 2); 1869 (III 1). Fuertes, M. 1086 (I 1c Garcia-Barriga, H. 11030 (II 1b). Gentle, P. H. 1447, 2704, 3097 (1 1b); 3211 (I 2.) Gentry, H. S. 736m, 5147, 5582 (I 1b) Clan A. 4919, 5989, 16356, 1470, 12172 (III 1); 7851, 11565, 18493, (IV 13); 8289, 20489 (II 2). oe T Aig (I 1b). on & W. von Hagen 1130, 1177 Herb. Amaz. 9572 (III SCH Heyde, Nee ee 12). Hinton, G. B. 3165, 3643, 6750, 11540, 15901, vg ad a 15) 10298. a > Howard, R., or & W. T. S 14782 Huber, J. 6894, : 6965 = 1). Jack, J. G. 5002, 5842 (I Ic). Johnson, H. 514 (I TA Jurgensen, C. 941 (I Kenoyer, L. A. A Crum 3770 (I 1b). 0 Killip, E. P. & A. C Smith 27685 TO la) ue G. 2079, aus A m: 3698 (II 1b). A. 5991, 6040, 6849 Hi 15 7 de 5380, 6327, 6628, 8312, 8666, 10533, 10744, 10679 (II 2); 10410, 10728, GE (II 1b). Kuhlmann, J. G. 284, 487, 578, 675, 1906, 12292 (III m "13055 1:2): 1962] BURGER—STUDIES IN MORACEAE 33 LeSueur, H. 570 (I 1b). Lewin, ët ME ar by. Liebmann, F. M. s.n. 14267, 14268, 14269, 14270, Tar, i5111 (I 1b); 14259; 14273 I Little, E ^ 6219 (II 2). Little, E. L. & R. R. Little 9670 (I la). Long, L. E. 215 (I1 Lundell, C. L. LP19, 2917, 6970 (I 1b); 6422 (I 2). Magalhaes, G. M. 843 (III Matuda, E. 0433, 0576, OG 2648, 3978, 3979, 4014, 16157 (I 2); 0685, 2598, 4022, et 16794, 16799 (I 1b); 1051, 2370, 478 (I a 3933, 4241, 15989 (I 3); 9 (II la). a exia, Y. 1142, 8844 (I 1b); 1701 (12); 5158 (III 1); 6303, 6262 (I la). e 157 (I 1b). 5 13). Molina R, A. 909, 1108a, 1300, 1443 (I 2); 2497, 3810 (I 1b). Montes, M. N. & A. E. Salazar 855 (I 1b). Morton, C. V. & E. Makrinius 2478 (I 2). Moses, 1, 69 (HIE 1), ). Pittier, H. 2255, 3627, 6636, 6637, 6652 (I 1b); 8659 (I la). Poiteau, A. s.n. (I a p (III 1). Purpus C. A. 231 16 oe 5049, 5988, 108 37, 11044, (I1 b). Riedel, L. s.n. (III 1); 19, 303 (IV 14). sai A. 187-11 ts , W. R. Fitch & P. G. Russell is (1 lc). ose, J. N., P. C. Standley & P. G. Russell 14428 (I Ib). AT dr Pavón 24/24 (II 2); 24/26, 24/28 (II 1b). Sagot, P. A. 530 (III ds de la Sagra, R. 490 (I Ic). NI C. J. W. & F. Deppe sn. (I 1b); E ar W. A. 1010, 1074, (I 1b); 1261, 1334 (12). Seibert, a s 1581, 1583 (I 1b). Sello, F. (III 1). Sessé, M. r M. Mociño 4647 (I 1b). Signiera, R. 3657, 4 (III 1). Shafer, J. A. 617, 3272, ipy (I 1c). Shattuck, O. 827, 1 He (I 1b). Sintenis, P. kutch, A. F. 925, 1511 a3: Geck 2094, 2383, 2386 (I 2); 2003, 2004 (II Smith, Austin 141, A391, rey "P2313, 2720, 2851, wd I 5); HI558 (I 2). Smith, H. H. 424 (II 1b); 1752, 2732 (I la). Spruce, R. s.n., xi: (III 2); 4521 (I la). Stahl, A. 862 (11 c). Standley, P. C. 8376, 19797, FT 22458, 27287, 30235, 30296, 30970, 4 1080, 53841, 54312, 54727, 54997, 70185, 70403, 70793, 72483, 79291, 79513, 70524, 88452, 88549, 88729, 89114, (I 1b); 9965, 13431, 20183, 20680, 56771, 60197, 64766, 64815, 86697, 89574, 89897 (I 2); 68956, 84572, 86808, (I C. & A Valerio 45271 (I 2). 153 Stev Stevenson, N. S. & H. P. Smart 140 (I Ib). Steyermark, J. A. 31511, 42744, 43664, 47670. 47774, 49549, 51030, 51185 (I 1b); 33499, 37496, 37576, 39157, 39451, 4 52069 (I 2); 33636, 46630, 46658, 46687, 46730, 47333, 47349, 47592 (13). Stork, H. E. 16 (I Ib). Stork, H. E. & O. B. dde 9506 (I la). Taylor, N. 146, 229 (I 1 essmann, G. 4083 o la) 5433 (II 2). 1 (1 2); 112167, 112169, TS Ule, E. 6838 (I lo; "vA (II 2). Warming, E. s.n. (III 1). Weberbauer, A. 182 (II 2; 7057 A Ib). Wedel, H. von. 196, 1599, 2075 (I 1b). Williams, Llewelyn 2086, 6471 (II 1b); 2 3380 (II 2); 4658, 5243, 6542, 6573, 1071 1o (I la). Wi lliams, L. O. 15799, 16850 (I 2). Williams, L. O. & A. Molina, 14815, 15601 Wilson, P. 54, 250 (I 1b). Woodson, R. E., P. H. Allen & R. J. Seibert 1924 (I 1b). Woodson, R. E. & R. W. Schery, 866 (I Ib). Woytkowski, F. 5645 (I la); hr (II 2). an s.n., 589, 592 (I Ic). ncker, TG, R. F. Dawson & H. R. um 6171 (I 2). [Vor. 49 34 ANNALS OF THE MISSOURI BOTANICAL GARDEN INDEX TO TAXA Roman type indicates accepted, preexisting names; italics indicate synonyms; bold face indicates new names, new combinations, and newly described taxa. Acanthinophyllum 27 Pseudosorocea Poeppigii 28 ilicifolia 21,29 Sahagunia 20 spruceans 30 colombiana 22 strepitans 27, 28 mexicana 20, 22 Bucephalon 6 racemifera 28 Mexicanum 13 strepitans 28 racemosum 6, 7 urophylla 10 Clarisia 20 Skutchia 6 biflora 21 caudata 6,13 ssp. biflora 22,23 = Soaresia 20 ssp. mexicana 22, 23 nitida 20, 24 colombiana 22 Sorocea ES ilicifolia 26, 28 colombiana 8 var. cuyunensis 28 mexicana 13 var. guianensis 28 stenophylla 28 var. micranthera 28 Trophis 6 var, paraensis 28 americana 6, 7 mattogrossensis 22 meridionalis 8 mexicana 22 Ramon 10 mollis 26 aurantiaca 18 nitida 24 chiapensis 16, 17 panamensis 22 chorizantha 13 racemosa 24,25 cuspidata 18, 19 spruceana 30 Hilariana 18 strepitans 26, 28 macrostachya 18 var, cuyunensis 28 Matudai 13 var, guianensis 28 mexicana 13.19 var. micranthera 28 nubium 16 var. paraensis 28 racemosa 7 urophylla 10, 26 ssp. meridionalis 8, 9 Excoecaria ilicifolia 27 ssp. racemosa 9, 12 Helianthostylis 4 ssp. ramon 9, 10 Myriocarpa cordifolia 2 var. meridianalis 8 Olmedia erythrorhiza 24 Ramon 10 Paratrophis 4,7 Trymatococcus 4 philippinensis 4 Urtica membranacea 2 1962] DODSON—-VARIATION IN CASTASETINAE 35 POLLINATION AND VARIATION IN THE SUBTRIBE CATASETINAE eins Ce H. DODSO The natural history ui d subtribe ant ae is briefly discussed. The present taxo- f the gen och rm the triggered by the relative amounts of sunlight received by the plants. Those plants which i ers female flowers. The pollen throwing apparatus and the pollination mechanisms of the sufficiently arid the: species to come together and pollination Ko mo nt has roduced several highly variable hybrid swarms. Caraway H. Dopson, Missouri Botantical Garden, 2315 Tower Grove Ave., St. Louis 10, Missouri. INTRODUCTION This paper has been written to discuss in a general manner, the pollination, pollination mechanisms, taxonomy, relationships among the members, sexuality and population variation of the Catasetinae. It is hoped that in this manner a basic reference will be formed for forthcoming papers dealing with specific phe- nomena in the group. The present status of taxonomy in the group is deplorably bad and classic herbarium techniques of study are of little or no help in critical groups, due to the general paucity of specimens and the destruction of important characters in their preparation. Field studies of population variation will surely be important in clarifying the taxonomy of the group. The subtribe Catasetinae has been of interest to both botanists and horticul- turists since the members of this group became known. The complicated mechan- isms enabling forcible discharge of the pollen apparatus found in most species have been described and commented upon by various authors. Such highly evolved mechanisms capture the interest of anyone from the orchid enthusiast to the evolutionist and considerable literature is available on the mechanics of these systems. However, very few accurate observations have been made upon the natural pollination of these plants in the field. Authoritative accounts have been published of pollination in Cycnoches by Allen (1952) and in Catasetum by Darwin (1872), based on observations by Criiger. Unfortunately, Criiger’s account was based on very superficial observation and both papers contain some errors and omit con- siderable pertinent information. This group is unique among the Orchidaceae for having members which produce separate staminate and pistillate flowers. They are also unique in having the capacity of forcibly flinging the pollinarium at an insect pollinator. The pol- linarium is thereby fixed in such a position on the insect that it may easily com- plete the pollination phase of the life cycle by delivering the pollen to the stigma of another flower. * Much of the work done for this paper was supported by a grant from the National Science Foundation (G-19957). [Vor. 49 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN TAXONOMY OF THE GROUP As presently constituted the subtribe Catasetinae consists of three genera, Catasetum Rich. with approximately 70 nominal species, Mormodes Lindl. with about 40 nominal species and Cycnoches Lindl. with seven species. The group is limited to the New World tropics where its members occasionally form a conspicu- ous element in the orchid flora. Its closest affinities probably lie with Cyrtopodium Br. and Galeandra Lindl. of the subtribe Cyrtopodiinae. While the linking members between these genera and the Catasetinae are now extinct, the gap is not great. Catasetum has been divided into two subgenera; Clowesia and Catasetum. The subgenus Clowesia, as treated by Mansfeld (1932) and Hoehne (1942), con- sisted of eight species all of which produce bisexual flowers and have no antennae on the column to trigger the expulsion of the pollinaria. Two quite dissimilar groups have been placed together in Clowesia and it is my opinion that they should eventually be separated. Castasetum roseum Rchb. f., C. warczewitzii Lindl. & Paxt., C. glaucoglossum Rchb. f. C. thylaciochilum Lem. and C. russellianum Hook., have the lip relatively thin in substance, not adnate to the column and usually fimbriate. In C. glaucoglossum and C. russellianum the anther is only slightly sensitive and the viscidium springs down only slightly when released. These species have many characters in common with certain species of Galeandra. The other group consists of C. dilectum Rchb. f., C. eburneum Rolfe and C. suave Ames an Schweinfurth. Unfortunately, there appear to be only two species in this group, but there are three names; two of which are of uncertain application. One species, corresponding to C. suave of Ames and Schweinfurth (the type of this species, at the Smithsonian Institution, has been examined) occurs from Costa Rica to Ecuador. The other species is only to be found in Costa Rica and Panama. Both species are somewhat variable and the type descriptions are not satisfactory to differentiate between the two kinds of plants. These species have the lip entire, fleshy and adnate to the thick, swollen column. I feel that this group is adequately distinct from the other group of species in Clowesia and should be recognized as a valid genus. The subgenus Catasetum as treated by Mansfeld and Hoehne consists of nearly 60 species, all producing separate male and female flowers. 'The subgenera Clowesia and Catasetum are not far separated in terms of evolution and the development of unisexual flowers from the bisexual group is easily traced. The column of most species of the subgenus Catasetum has long projections or antennae extending from each side of the anther to the center or cavity of the lip. These antennae, when touched, release the tightly stretched stipe of the pollinarium so that the viscidium is flung out and onto the thorax of the insect. Mansfeld divided the subgenus Catasetum into two groups; Pseudocatasetum and Meizocatasetum, the former having male flowers which are devoid of antennae and the latter having antennae. He then went further to divide Meizocatasetum into two groups; Isoceras and Anisoceras, depending on whether the antennae were parallel or one was bent back under the other. I have found this division to be unworkable on many occasions 1962] DODSON—VARIATION IN CASTASETINAE 37 in which some flowers had the antennae parallel, while others on the same in- florescence had one bent. In other instances one inflorescence would have flowers with antennae bent and another inflorescence of the same plant would have flowers with parallel antennae. The flower morphology is not particularly distinct between the two subgenera and the more advanced members of Clowesia closely approach the species which lack antennae in the subgenus Catasetum. Even in the more advanced members of the subgenus Catasetum hermaphroditic flowers are occa- sionally produced which are quite reminiscent of some members of Clowesia. Cycnoches has been revised recently by Allen (1952). Allen’s treatment of this genus is unusual in modern taxonomic work on the tropical Orchidaceae in that it is based on personal knowledge of the plants as living populations in the field. Cycnoches is composed of two distinct subgenera. In the subgenus Cycnoches the male and female flowers are essentially the same in form with the only signifi- cant difference being that the column is shorter and thicker with the stigma functional in the female flowers while the column is long and slender with only the anther present in the male flowers. Allen lists four species in this subgenus. The subgenus Heteranthae produces male and female flowers which are strikingly different. The female flowers are morphologically similar to the female flowers of the subgenus Cycnoches but are smaller. The male flowers are much smaller than the female flowers, are produced in considerable quantity on long pendant racemes, have a flexible claw on the lip and usually have a toothed fleshy callus. Allen lists three species in this subgenus. Mormodes has never been revised. This is probably due to the formidable variation encountered within the populations. In several cases, characters which were previously used to separate species have been found to vary widely on one inflorescence. Certainly many of the 40 nominal species will be found not to be valid when a careful study is made of the genus. PHYLOGENETIC RELATIONSHIPS WITHIN THE CATASETINAE The ancestral types linking the subtribe Catasetinae with its closest allies in the Cyrtopodiinae are now extinct but sufficent correlation between the two sub- tribes remains to indicate relationship. A phyletic series can be observed from Catasetum glaucoglossum and C. thylaciochilum as the simplest types through C. russellianum to C. roseum and C. warczewitzii in the subgenus Clowesia. The sensitive anther which is character- istic of the Catasetinae is not highly developed in some of the members of this group. Catasetum discolor of the subgenus Catasetum is morphologically quite similar to C. warczewitzii except that separate male and female flowers are pro- duced. More advanced species are then encountered with unisexual flowers and are furnished with complicated antennae for triggering pollinia expulsion. In the latter group the species with rather similar hooded male and female flowers are probably less advanced than the species with open male flowers (such as C. barbatum Lindl.) which have highly developed fringes and papillae on the lip of the male flower. The basic separation between the types of flowers produced by the major groups [Vor. 49 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN of Catasetum is morphological but is founded upon different pollination mecha- nisms. In the portion of the subgenus Clowesia which produce less fleshy flowers the column is of necessity placed relatively near the lip in order for the pollinator to brush the anther with its upper thorax and receive the pollinarium. Catasetum roseum, C. thylaciochilum, C. warczewitzii, C. russellianum and C. glaucoglossum all appear to be of this type. Catasetum dilectum, C. eburneum and C. suave appear to be based on another system in which the bee approaches the flower, lands on the front of the open flower with its head toward the apex of the lip and its abdomen over the column, touches the triggering mechanism and receives the pollinia on the ventral side of the thorax either in front of or among the legs. In the more advanced species of Catasetum the bee lands upon the lip and while approaching the source of the fragrance, located in a depression or within a saccate hood, inadvertantly touches one of the antennae, releasing the pollinarium which is flung for some distance to strike the pollinator. The pollinator is usually positioned in such a manner that the viscidium is attached to the upper thorax. Cycnoches may have developed from the fleshy flowered group of species in the subgenus Clowesia of Catasetum. Though the general features of the labellum are not entirely similar in the two groups, the apparatus for expulsion of the pollinarium is very similar. Both types of flowers are designed for placement of the pollinarium on the ventral surface of the pollinator. In Catasetum suave, for example, the column is short and fleshy with the anther positioned on the apex of the column. The simple extension of the anther on a long, slender column would result in the same type of apparatus as found in Cycnoches. The triggering mech- anisms for expulsion of the pollinia in the two groups are based upon leverage and tension released by lifting the apex of the anther. These two groups are the only members of the Catasetinae which use this system for propulsion of the pollinia (fig. 1). The fleshy members of Clowesia are normally quite uncommon in nature, perhaps an indication that they are not particularly successful. They are only found in the tropical cloud forest regions of Central and South America. These cloud forests seem to be particularly hospitable for relict and marginal epiphytic species. Cycnoches has apparently left the bisexual flowered condition and normally only exhibits unisexual flowers. 'The occasional production of hermaphroditic flowers indicates that the capacity for production of bisexual flowers is not com- pletely lost. The genus Mormodes is apparently in the process of developing unisexual flowers at present. There have been reports of the production of unisexual flowers in certain species of Mormodes and I have encountered the same condition in a species in Ecuador. This genus is perhaps not as highly successful as Catasetum or Cycnoches. Within their range they are usually not encountered in such great quantity as the other two genera and are often quite rare. The plants are not as tolerant of adverse ecological conditions as are the related genera. 1962] DODSON—VARIATION IN CASTASETINAE 39 SEXUALITY The production of unisexual flowers in these orchids has been the subject of considerable comment and discussion. It is, of course, unique in a family nearly typified by bisexual flowers. This development of polygamo-dicecious flowers appears to be in a stage of transition in certain species within the Catasetinae. The species within the subgenus Clowesia of Catasetum apparently produce bisexual flowers only, while the members of the subgenus Catasetum ordinarily produce unisexual flowers. Species of the subgenus Catasetum do occasionally produce bisexual flowers, and both male and female flowers are sometimes found on the same inflorescence. Within the genus Cycnoches no species occur which produce bisexual flowers ex- clusively, but bisexual flowers are found on these plants on rare occasions. Recent evidence (Allen 1959) indicates that two types of flowers are found in Mormodes lineatum Batem.; staminate flowers, which apparently function only as male flowers, and perfect flowers which produce functional pollinia as well as a functional stigma. The two types of flowers are different morphologically but a transition can be found between them. I have observed the same situation in M. buccinator Lindl., from central Ecuador. Certain plants were noted which produced the same large flowered type with little contortion of the column and a very large stigmatic sur- face which Allen noted in M. lineatum. On other occasions the same plants pro- duced smaller flowers with strongly contorted columns and reduced stigmas. mum Hh WB nisi 7) igure 1. Action of the pollinaria une apparatus in the various .. ege the Catasetinae. The arrow indicates int of contact which releases . Catasetum Geer Hook. (Slightly exaggerated) c. Catasetum discolor Lindl. d. Catasetum platyglossum Schltr. e. Catasetum aff. s A. & S. f. Cycnoches ventricosum Batem. g. Mormodes buccinator Lin L [Vor. 49 40 ANNALS OF THE MISSOURI BOTANICAL GARDEN Figure 2 shows two views of a flower of M. colossus, the first taken with the anther in place and the second taken 24 hours after removal of the anther. The column has straightened and reorganized itself so as to function as a female flower. This phenomenon has not yet been observed in other species of Mormodes. Figure 2. Twisting of the colum in TE colossus Rchb. d a. Flower with the anther in paw b. The same flower 24 hours after removal of the pollinarium. The factors which make it possible for a plant of this group to produce male flowers during one season, female flowers in the following season and perhaps a mixed inflorescence in the succeeding season have never been clearly explained. There has been considerable speculation about temperature relations triggering one type or the other. Also an intrinsic ability on the part of the plant to decide whether it should produce one sex or the other has been suggested. Studies made by the author over a 14 month period in coastal Ecuador involving Catasetum macroglossum Rchb. f., C. platyglossum Schltr., Cycnoches lehmannii Rchb. f., and Mormodes buccinator Lindl., indicate that the production of male versus female flowers depends primarily upon ecological factors. When a plant growing in full sunlight has adequate moisture during the growing season and has its roots em- bedded in a suitable substrate (rotting wood is apparently most favorable) it will be robust and will generally produce female flowers. If on the other hand, either sunlight or humidity is lacking, or the plant is growing on the bark of relatively smooth-barked tree, it will normally not be as robust and will produce male flowers. The substrate does not appear to be as important as the other two factors for plants with female inflorescences have occasionally been observed on living trees, though in all such cases the trees had deeply fissured bark. The factor of sunlight appears to be particularly important. Experiments conducted at the Instutito Botánico of the University of Guayaquil indicated that the sex of the flower could be changed during one season. Thirty robust plants of Catasetum macroglossum with female flowers which had been in full sunlight under natural conditions were brought in and 15 were placed in dense shade and 15 in full sun. These plants had produced their first inflorescences before the full development of the pseudobulb and still retained their leaves. New inflorescences were produced and all plants in shade flowered male. Thirteen plants placed in full sun continued to produce female inflorescences. Two plants which were not quite as robust as others produced inflorescences with both male and female flowers present. Large plants which had produced male inflorescences in 1962] DODSON—VARIATION IN CASTASETINAE 41 the early part of the season produced female inflorescences when placed in full sun for the remainder of the season. It seems clear from these experiments that determination of sex depends largely upon growing conditions of the individual plant. Robust plants growing in full sun usually produce female inflorescences. The chemical differences in the plants which are producing one sex as opposed to plants producing the other have not been investigated. There is the possibility that the production of female flowers may be due to carbohydrate balance or, on the other hand, it may also be due to a hormone balance which is affected by sunlight. A similar situation has been reported in Cannabis sativa in which the production of male or female flowers was controlled by presence or absence of hormones. As a general rule very few plants in a population are growing under conditions conducive to the production of female flowers. In the Catasetum macroglossum populations growing in the tropical deciduous forests of coastal Ecuador, approxi- mately 21 male inflorescences for each female inflorescence were counted. In popu- lations growing in the regions of greater humidity which are now planted to cacao and coffee, the trees provide considerable shade and the count was 46 male inflo- rescences to one female. In populations of Cycnoches lehmannii growing in old cacao plantations around Quevedo the count was 27 male inflorescences to one female. No accurate results were obtained in the population of Mormodes buc- cinator due to the shortage of specimens, however it was noted that the only two plants which produced the female type inflorescences were growing in full sun in ant nests and had extremely large and robust pseudobulbs. Seven other plants were found growing in partial shade which did not have large pseudobulbs and all produced male inflorescences. The discrepancy in numbers between the male and female inflorescences pro- duced is apparently important in the ecology of the plants. As a rule the male flowers in all species are short lived, lasting at most four or five days. Their seg- ments are usually thin in substance and wilt within hours after the pollinaria have been discharged. Within two or three days after opening their fragrance becomes strong and they wilt shortly thereafter. If the pollinator does not discharge the pollinarium within a day or two after the fragrance becomes strong they are often discharged automatically or by the slightest movement of the wind. The female flowers, on the contrary, are very long lasting, occasionally persisting up to six weeks. They are usually large and of very heavy, fleshy substance and do not emit their fragrance until seven or eight days after they have opened. It would seem logical that more male flowers would be needed since insect visitation is often rather haphazard and it would be highly important to the fertilization of the female flowers to have a large quantity of short-lived, fragrant male flowers in the vicinity. With a succession of numerous short-lived male flowers available to the pollinators the possibilities of the pollinator having pollinaria attached when he visits the female flower would be greater. POLLINATION Pollination within the subtribe Catasetinae is very poorly documented and the accounts which have been published are inaccurate in many points. We have [Vor. 49 49 ANNALS OF THE MISSOURI BOTANICAL GARDEN observed pollination in Catasetum macroglossum, C. platyglossum, C. suave, Cycnoches lehmannii, C. egertonianum and Mormodes buccinator (Dodson & Frymire 1961b). Allen (1952) has reported the pollination of Cycnoches ventri- cosum, Catasetum oerstedii and Mormodes histrio. Hoehne (1933) reported on Catasetum cernuum. Ostlund (unpublished) very accurately recorded the pollina- tion of Mormodes histrio, M. uncia and M. lineatum. Unfortunately, these species represent a very small portion of the actual number of species in the subtribe but they do represent both subgenera in Catasetum and Cycnoches, and four species of Mormodes. A discussion of pollination in each of the species which we observed should establish the basic patterns of pollination involved. It would appear that all species in the Catasetinae are pollinated by bees of the family Apidae, subfamily Euglossini. Two genera important in pollination of the Catasetinae are Eulaema and Euglossa. We have not observed any other visitors to the flowers. For a more extensive discussion of these bees and their habits see Dodson (1962). Pollination of Catasetum suave (this name is tentatively used until the iden- tity of earlier names can be definitely fixed) was observed in June of 1960 at a site 12 kilometers east of Balzapamba on the road to Guaranda in the Province of Bolívar. This locality is on the western slope of the Andes at an elevation of 1400 meters. The plants are terrestrial and grow on the faces of road cuts. Pollination is effected by males of Eulaema cingulata (L.). The bees approach the flowers, which are carried on racemes of 10-20 flowers each, and land upon the open sur- face of a flower attempting to reach the source of the strong clove oil-like fragrance which emanates from the cavity of the lip (fig. 3). The flowers are nonresupinate, therefore the lip is uppermost in the flower. In attempting to thrust its head into the cavity of the lip, the bee moves the small hump on the anther cap and the stipe is released, flinging the sticky viscidium upwards where it strikes the bee either on the trochanters of his legs or in front of the first pair of legs. The stipe of the pollinarium, immediately after attachment to the bee, is curled and does not dry and straighten out for several minutes. This tends to insure that the bee will have flown on to another plant and will effect cross-pollination. The viscidium covers the stigmatic surface of the flower so that the pollinarium must be removed before pollination can occur. When a bee with pollinarium attached comes to a flower and repeats the attempt to enter, the pollinia are guided into the stigmatic cavity, trapped in the narrow extremities of the cavity, and pulled from the bee upon his exit from the flower. Lankester (1960) reports the pollination of the other species from this group, which he (possibly quite correctly) called C. dilectum. The flowers of this species are essentially similar to C. suave. The primary differences between the two species lie in the sub-capitate inflorescence, strongly reflexed tepals, smaller flowers and enlarged callus of C. dilectum. According to Lankester C. dilectum is pol- linated by a species of Euglossa. His description indicates that the activity of the bees is similar to what I observed in the pollination of C. suave. 1962] DODSON—VARIATION IN CASTASETINAE 43 * Figure 3. Eulaema cingulata (Fab.) visiting the flowers of Catasetum aff. suave A. & S. Catasetum russellianum is representative of the closed flowered types of the subgenus Clowesia and experiments with living male bees of Eulaema cingulata at the Missouri Botanical Garden indicate that this species of Catasetum with its only slightly sensitive anther uses a different mechanical system for attaching the pollinarium. The bee, in attempting to reach the saccate portion of the lip, near the base of the column, touches the stipe of the pollinarium and the viscidium is released and swings down into such a position that in withdrawing from the flower the bee will push against the viscidium with its metathorax. Upon subsequent entrance and exit from another flower—allowing time for the anther cap to dry and fall—the pollinia are left behind in the narrow stigmatic cavity. The subgenus Catasetum has a rather different system of pollination primarily because there are two sexes of flower involved. Two species from this section were observed and have essentially the same system although the flower form is different. Two or three days after opening, the mature hood shaped male flowers of C. macro- glossum begin emitting a strong musky odor. Male bees of four species of Eulaema were observed visiting the flowers. These were E. cingulata (L), E. tropica (L), E. bomboides Friese and E. speciosa Moc. The bees grasp the labellum, which in this species of Catasetum is uppermost in the flower, and enter upside down (Fig. 4). After entry they do not extend their tongues, but scratch the inner surface of the labellum with their fore legs using the same action as a dog digging a hole. At the same time their antennae brush over the scratched surface. Chemoreceptive pads on the tarsi of the bees (which are present in all male bees of the Euglossini) apparently allow the bee to taste the liquid which is emitted from the scratched surface of the labellum. While scratching at the inner surface of the saccate [Vor. 49 44 ANNALS OF THE MISSOURI BOTANICAL GARDEN pollinia being guided into the stigmatic pocket of the column where they are removed by leverage as the bee struggles free, 1962] DODSON—-VARIATION IN CASTASETINAE 45 labellum the bee contacts one of the antennae which are extensions of the sides of the rostellum and hold the anther cap and stipe of the pollinarium under tension. The tips of the antennae are located just over the deepest point of the saccate labellum where the strongest odor is emitted. The movement of the antennae triggers the release of the pollinarium. The pollinarium is propelled upward and outward, the sticky surface of the viscidium foremost, and is stuck to the thorax of the bee. The adhesive substance of the viscidium sets rapidly, and the stipe hinges back at the point of attachment to the viscidium lying flat along the dorsal mid-line of the abdomen of the bee. The bee may then proceed in its wandering to a mature, aromatic, female flower. The female flower takes somewhat longer to mature than does the male and begins to give off a strong odor on the third or fourth day after opening. The non- resupinate female flowers last up to a month if not pollinated and continue to emit their strong fragrance during the day; however, the fragrance disappears within hours after pollination, and the stigma swells shut. Bees are attracted to and enter the female flowers much as they do in the male flowers. When a bee with a pol- linarium attached to its thorax enters a flower, the stipe, hinging at the viscidium, swings down in line with the stigmatic cleft. As the bee backs out to leave the flower, one or both of the pollen masses are guided down on the back side of the lip and are caught in the stigmatic cleft. They are then wrenched free—by leverage—from the weak bonds with the stipe and are left to fertilize the flower. Pollination in C. platyglossum is essentially identical, the only major difference being that the male flowers are resupinate with the open lip lowermost in the flower (Fig. 5). The bee simply lands on the open labellum, walks to the basal cavity and touches the antennae, setting off the same process as in C. macroglossum. The female flowers are so nearly identical to C. macroglossum that it is not possible to separate the two species from the female flower alone. Hoehne (1933) has reported on the pollination of Catasetum cernuum (Lindl) Rchb. f., in Brazil, by Euglossa violacea. Catasetum cernuum is similar in many respects to C. platyglossum, having male flowers which are open rather than hooded as in C. macroglossum and its allies. The flowers of C. cernuum are considerably smaller than those of C. platyglossum and are highly colored with orange and red igure 5. a. Male Eulaema tropica (L.) visiting the male flowers of Catasetum aff. platyglossum Schltr. b. Male Eulaema cingulata (Fab.) visiting the female flowers of Catasetum platyglossum Schltr. [Vor. 49 46 ANNALS OF THE MISSOURI BOTANICAL GARDEN spots. This species is representative of a large number of species from South America which have open male flowers and more or less fimbriate lips. Hoehne’s description indicates that the pollination process is essentially the same in this species as in C. platyglossum with the exception that C. cernuum utilizes bees of the genus Euglossa as pollinators. Probably most of the species in the group allied to C. cernuum are pollinated by Euglossas. The pollination process in Cycnoches lehmannii appears very complex but in reality is quite simple. The system depends on perfect placement of the bee in relation to the lip and sexual apparatus of the flower. In order to place the bee properly the flower has developed a lip which is nonresupinate, forcing the bee to land in an inverted position, with the source of the odor located at the apex of the callus of the lip. The female flowers appear to have an odor that is a little stronger which will be caught by the hooks of the apex of the column) 1962] DODSON—VARIATION IN CASTASETINAE 47 and more penetrating than the male flowers. The male bee of Eulaema cingulata approaches the male flower and lands on the inverted lip, gradually swinging around into an inverted position with the 2nd and 3rd pair of legs grasping the edges of the lip. (Fig 6). The first pair of legs are brushed back and forth on the apex of the callus. The odor appears to emanate from between the apex of the callus and the lip. The extended callus of the male flower forces the bee—in attempting to get closer to the source of the odor—to swing his abdomen down and release his lower pair of legs. This is an extremely awkward position for the bee and the lower portion of the adbomen of the bee brushes the trigger mechanism of the column. The brushing of the tip of the anther cap releases the viscidium which is held under tension in what would normally be the stigmatic region in a bisexual flower. The stipe whips the viscidium around striking the bee on the last sternum of the abdomen where it sticks by means of its rapid drying cement. The anther cap remains over the pollinia and the stipe is curled around the tip of the bee’s abdomen. The stipe of the pollinarium, through differential drying of the two surfaces, gradually straightens out and after about 40 minutes has straightened to a stiff pendant position. The anther cap remains in place covering the pollinia for about two to three hours before drying sufficiently to fall off. The pollinarium is then ready for the bee to pollinate the female flower. The female flower is constructed differently. The position of the column and stigma is quite different from the male flower in relation to the callus, being much shorter and thicker. The callus is not as extended as in the male flower and the bee has little difficulty in hanging and scratching on the callus tip. When the bee is ready to fly he must fall for a short distance to be able to revert to a flying posi- tion. In falling, the bee, through his predetermined position on the callus is oriented in such a manner that in passing the column the pollinia are caught by the finger- like processes of that structure. One of the pollinia is caught and is stripped from the stipe. Within a few hours the stigma swells, closing around the pollinia, the odor ceases, and the flower parts wither; the pod ripens about a year after pollination. Pollination of Cycnoches egertonianum Batem. was observed in the Amazon drainage area of the southeast Ecuador. Cycnoches egertonianum is a member of the heteromorphic group of Cycnoches in which the male flowers—unlike the group to which C. lehmannii belongs—are strikingly different from the female flowers. The male flowers of this species are small and are carried on long pendant racemes. The lip is somewhat different from that of C. lehmannii in that the claw is thin and flexible and the blade is divided into numerous fleshy terete divisions. Male bees of Euglossa viridissima were observed pollinating this species. (Fig. 7). The bees would land on the blade of the lip which would then swing down with the weight of the bee. This action caused the tip of the bee’s abdomen to touch and lift the anther cap on the apex of the column, discharging the pol- linarium which became affixed to the bee much as in Cycnoches lehmannii. The female flowers are nearly identical to those of C. lehmannii except that they are somewhat smaller and are darker green in color. The pollination of the female flower is essentially the same as discussed in C. lehmannii above. [Vor. 49 48 ANNALS OF THE MISSOURI BOTANICAL GARDEN Pollination of Mormodes has been reported for four species (Allen 1952 and Ostlund unpublished). We observed bees with pollinia attached to their thorax visiting the female flowers of M. aff. buccinator in Ecuador. The bees, Euglossa * viridissima, landed on the outstretched lip and proceeded to scratch the surface ol the lip with their front pair of feet. (Fig. 8). In moving around on the lip the pollinaria on their backs came in contact with the very broad stigmatic surface and the pollinia became detached. Though the anther of the pistillate flower in Mormodes can eject the pollinarium, it is held up and far away from the lip of the flower in a position which would be very difficult for a bee to encounter and dis- charge. In the male flowers the column is twisted in such a way that the apex of the column lies in contact with the lip. Usually the twisting places the back of the column in contact with the lip and the anther, located on the ventral surface, is facing outward. A small terete finger or tubercle formed from the tip of the column is in contact with the tip of the anther cap. This tubercle lies on the surface of the lip and the slightest movement moves the anther cap allowing the viscidium to be released. The viscidium, carried by the stipe, springs around and strikes any object which has moved the tubercle. The cement on the viscidium quickly dries leaving the pollinarium attached to the insect. After about 30 minutes have passed the stipe e T. Male Euglossa viridissima Fr. pollinating Cycnoches egertonianum Batem. a. Male bee hanging from the flexible lip of the male flower (note the pollinarium extending from the apex of the abdomen of the bee). b. Male bee scratching at the callus of the lip of the female flower (note the pollinarium which has been Pier te in the stigma). 1962] DODSON—VARIATION IN CASTASETINAE 49 straightens and the pollinia are held erect from the thorax of the bee in such a position that when the bee walks under the stigma of a flower, contact is made. that the stipe of the pollinarium has straightened placing the pollinia in position to be deposited on the stigma in the background. In no instance of pollination observed by me, in any of the above mentioned species of the Catasetinae, did the bees gnaw on the lips of the flowers as reported by Darwin (1877), Criiger (1865), Porsch (1955), Allen (1952 & 1959) and as so eloquently described by Ames in Catasetum (1945). The bees were never noted by us to even touch the surface of the labellum with their mandibles and they did not extend their tongues. It is possible that chewing or gnawing do actually occur during pollination of these orchid species in Panama or Trinidad but this hardly seems reasonable since the same species of bees are involved as in Ecuador. Ostlund (unpublished) reports the same phenomen of scratching in the pollination of Mormodes lineatum in Mexico. He also pointed out that the bees did not gnaw the flowers. We have noted numerous small bees and wasps which do gnaw the flowers of these and other orchids but which are too tiny to actually effect pollination. None of the Catasetinae provide nectar or has nectaries. The bees, apparently in all cases, scratch the surface of the lip at the point of origin of the odor and then touch the liquid released, with the sensory pads on their feet. This liquid appears to have an intoxicating effect on the bees because they cease to be wary and then can be approached without being frightened. The bees often spend considerable time at one plant and will enter the same flower repeatedly. On each occasion, they enter, scratch the surface of the lip for about 60 seconds, leave the flower and hover for about 40-60 seconds, apparently checking for danger, and then reenter the flower. After the bees have been visiting a particular flower for some time, it can even be removed and the bee will continue to search out that flower from the odor. Perhaps the most significant factor brought out by these observations is the fact that in both Catasetum and Cycnoches, species groups have become both mor- phologically and ecologically distinct by their adaptation to different kinds of pollinators. Of even greater interest is the fact that the two kinds of pollinators are [Vor. 49 50 ANNALS OF THE MISSOURI BOTANICAL GARDEN closely related, have similar habits and similar morphology. Their principal dif- ferences are in size and response to different fragrances produced by the flowers for their attraction. Indications are that the bees have been on the scene for a much greater period of time than the orchids. The orchids have been able to utilize the two types of bees as a crutch, so to speak, in speciation. The opportunity of using different kinds of pollinators, which would not overlap and thereby would not nullify the adaptation to distinct mechanical systems, may be as important in speciation to a group of this nature as are the development of incompatibility and genetic barriers to crossing in many other groups of flowering plants. Speciation in these orchids could occur by adaptation to one pollinator and later to another and then back again. Such speciation would, of necessity, be on a large scale with drastic changes in morphology of the flowers expected. Elimination of inter- mediates by early extinction would leave large discontinuity between groups and would present to the systematist clear-cut groups even at the generic level. The development of such strikingly different morphological features as found in the Catasetinae and the Stanhopeinae (Dodson and Frymire 196la) may be attribut- able to just this sort of speciation. VARIATION IN POPULATIONS Population variation in the subtribe Catasetinae has long been a problem in species classification. The problems in this group are not so great on the generic level. For the most part the genera are amply distinct, but the delineation of species has been unusually difficult. Taxonomists working with preserved material have been plagued by this problem and the all too common solution has been to describe each troublesome specimen as a new species, based upon the obvious floral differences from any other known species. Unfortunately, though this has been the easiest method for handling the problem, it has not reflected species dif- ferences. Until recently no studies have been attempted on extensive living popu- lations in their natural habitats. The problem of extreme population variability is not peculiar to the Catase- tinae. It is also quite evident in certain species of nearly all genera of the Stan- hopeinae, the Maxillariinae and in many genera of the Oncidiinae and Epiden- drinae. The only usable taxonomic treatments of large variable groups in the orchids have been done recently by Paul Allen in Cycnoches (1952) and Chysis (1955). His treatments of these two troublesome genera reflect his deep knowledge of the extent of population variation gained by personal knowledge of the taxa involved. His reduction of a series of 50 named species of Cycnoches to 7 with 4 subspecies demonstrates, for the first time in the group, not only the excessive number of named variants but also phylogenetic trends within the group. As an example, he reduced 14 named species to one, Cycnoches egertonianum and re- tained three subspecies which represent geographical and population trends. Similar work is needed in the genera Catasetum and Mormodes. Many names have been applied to individual variants from variable populations of Catasetum, particularly in South America where population studies have been nonexistent and few preserved specimens are available in herbaria. In many cases names have been applied to plants without any knowledge of their origin. 1962] DODSON—VARIATION IN CASTASETINAE 51 1217077 Le NS AMBATO “OAM BA x X. LIVAS d x Ve Qo CATASETUM MACROGLOSSUM 8 C. PLATYGLOSSUM HYBRIDS S T | Figure 9. Map of western Ecuador showing the distribution of Catasetum macro- iba Rchb. f., C. platyglossum Schltr. and their hybrids. In coastal Ecuador, Ben of two highly confused species of Catasetum are found in quantity. These are C. macroglossum and C. platyglossum. During the early months of "1988, 1060, and 1962 I had the opportunity to observe and study hundreds of flowering plants of these species and some rather startling ob- servations were made. The two species were apparently once separated by a range of mountains, the Cerros de Colonche, which run north and south through the coastal plain paralleling the Andes. These mountains effectively split the coastal plain into two valleys with the eastern valley about 50 miles wide and flanked by [Vor. 49 52 ANNALS OF THE MISSOURI BOTANICAL GARDEN the Andes on the eastern side. The western valley is about 30 miles wide and is flanked by the Pacific Ocean. A large rather dense and variable population of Catasetum macroglossum occupies the eastern valley and C. platyglossum occurs in a sparse population in the western valley (see Fig. 9). P9 NS * Y e e 10. Catasetum macroglossum Rchb. f. showing two inflorescences, the female to = left and the male to the right. 1962] DODSON—VARIATION IN CASTASETINAE 53 | IM N A ke SESCH s M [S S S V SN BOX D. ^ In ER ST Figure 11. Catasetum platyglossum Schltr. showing the male inflorescence. [Vor. 49 54 ANNALS OF THE MISSOURI BOTANICAL GARDEN Until recently [probably less than 200 years as established by comparison of the grazing areas of livestock during the colonial period (1600-1700) and the map in Wolfs Geografía y Geología del Ecuador (1892)] the Cerros de Colonche with its rainforest-clad slopes provided an effective barrier to gene exchange. A drying trend throughout the coast, caused by an apparent shift in the Humboldt current, has reduced the width of the rainforest, particularly in the passes between the higher areas. In these passes the two species have come together and the result has been the production of a series of hybrid swarms of bewildering variation. Figure 10 shows a plant with male and female inflorescences of what could be considered as typical C. macroglossum. Figure ll represents a plant with a male inflorescence of typical C. platyglossum. 'The female flowers of both are so nearly identical that no statistical difference could be noted between the two. Figure 12 shows a series of specimens taken from a representative group at Las Pefias, a pueblo some 30 kilometers south of Jipi-japa in the province of Manabí on the western slopes of the Cerros de Colonche. All specimens were collected within a radius of 50 meters. Several of these individuals could easily be given specific names and in fact a plant very similar to number 4 of figure 12 which I collected from this same locality was donated to the Los Angeles County Arboretum where it was seen by Fowlie (1962) who suggested very strongly that it be named as a new species since it did not fit any of the descriptions contained in Mansfeld's monograph of the genus. The problem became doubly significant when it was discovered that the same bees visit and pollinate all the various types of flowers indescriminately, regardless of their color or form. Bees of Eulaema tropica were observed passing from the flowers of the plant illustrated as number 4 of figure 12 to number 8 of the same figure. One bee visited a flower of number 4 and received pollinia on its thorax and then visited number 8. Another bee visited the same flower of number 4 and did not receive pollinia from that flower since it had already been triggered. He then flew to number 8 and did receive pollinia from a flower on that inflorescence. Both bees could then presumably fly to female flowers of either type and effect pollination. This behavior on the part of the bees is apparently the reason for the variation noted where populations of the two species come in contact. The im- portance however, is that variation is apparently permitted by a lack of selection for a particular type of flower since visits by bees were frequent for all types of flowers and no particular preferences were noted. These species have evolved morphologically and physiologically to withstand a wide range of ecological habitats, from the xerophytic conditions of the coastal thorn forests to the wet tropical cloud-forests on the slopes of the Andes. Though having diverged in the past to form separate populations with very characteristic floral structures, the genus has maintained integrity of interfertility within all of its taxonomically definable species. Regardless of extremes in other floral structures, that part of the male flower critical in ensuring cross-fertilization, the column and its incumbent ejectable pollinarium, show remarkably little variation. Any con- sistant deviation from the condition that allows the successful ejection of the 1962] DODSON—VARIATION IN CASTASETINAE 55 Figure 12. A series of flowers, taken from a population at Las Piñas, Zoe sn: Strating the extremes from a hybrid swarm between Catasetum macroglossum Rchb. f. an C. platyglossum Schltr. [Vor. 49 56 ANNALS OF THE MISSOURI BOTANICAL GARDEN pollinarium and its precise deposition on the thorax of the bee would tend to elimi- nate that plant as a contributor to the gene pool of the population. A situation which is apparently quite comparable to these populations and their hybrids in Ecuador occurs in Venezuela between C. macrocarpum A. Rich ex Kunth and C. pileatum Rchb. f. Catasetum macrocarpum is very similar to C. macro- glossum and C. pileatum is quite close to C. platyglossum. A vast series of inter- mediates between these two concepts have been encountered and many were named by early taxonomists before the situation was understood. Plate 502 in Lindenia illustrates a series of these presumed hybrids which is quite similar to a comparable series which could be selected from the C. macroglossum-C. platyglossum series in the Cerros de Colonche. Although Mormodes as a genus is amply distinct, the species boundaries or con- cepts within it are nearly nonexistent. Several recent papers, i.e., Teuscher (1954), Allen (1959) and Correll (1941) have pointed out the confusion involved but have done little to clarify the situation. There are three distinct kinds of Mormodes; those with strongly 3-lobed lips, those with obcordate lips and those with cordate lips. The factor which makes classification almost impossible, however, is that nearly every conceivable intermediate between these three basic types can be encountered. Not a great amount of information is as yet available but there may be definite, distinct populations of certain of these intermediate types. In other cases, there may be extremely variable populations demonstrating all types of variations on the three themes. This group promises to present formidable taxo- nomic problems, and there is the possibility that a simple clearcut account will not be possible. LITERATURE CITED Allen, P. H 1952. The Swan orchids, a revision of the genus Cycnoches. Orchid Jour. 1: 173-184, 225-230, 273-276, 349-403. —. 1955. A revision of the genus Chysis. Am. Orchid. Soc. Bull. 24: 665-666. —. 1959. Mormodes lineatum: A species in transition. Am. Orchid Soc. Bull. 28: 411-414. Ames, O. 1945. The strange case of Catasetum barbatum. Am. Orchid Soc. Bull. 13: 289- C “n 1865. A few notes on the fecundation of orchids and their morphology. Jour. n. Soc. Bot. 8: 129-1 Darwin "C. H. 1877. On the fertilisation of orchids by insects. New York. 2nd ed. Dodson, C. H., & rymire. a. Preliminary studies in the genus Stanhopea. fod Missouri Bot. Ga rd. 48: 137-172. 196lb. Natural pollination of orchids. Missouri Bot. Gard. Bull 49: 133-152. Dodson, C. H. 1962. The importance of pollination in the evolution of the orchids of Tropical America. Am. Orchid Soc. Bull. 31: 525-534. Hoehne, F. C. 1933. Contr. para o Conh. do genero Catasetum L. C. Rich. e especialmente o hermafroditismo e Seet das sus flores. Bol. da Agricultura 133-196. 1942, Mormodes, no. 102; Catasetum, no. 103; Cycnoches, no. 104. Flora Brasilica 12(6): 94-114. Lankester, C. H. 1960. A reminiscence and its cause. Orchid Rev. 68: 354. per 1932. Die Gattung Catasetum L. C. Rich. Repert. Sp. Nov. 30: 257-275, 31: Porsch, O. 1955. Zur Biologie der Catasetum-bliite. Österr. Bot. Zeitschr. 102: 117- dare H. 1954. Two forms of Mormodes lineatum Batem. ex Lindl. Zeg um 3: 178-180. COMPARATIVE MORPHOLOGY OF THE OSMUNDACEAE WALTER HEWITSON ABSTRACT Thirteen species of the Osmundaceae were compared in the Tru respects: gross morphology of the fronds, sporangial structure, hae structure, leaf base anatomy and rhizome morphology. The purpose of comparing rhizome structure was to try to find ea zomes. Further study on larger rhizomes of these two Se may indica S ihe alee pathway by which the stele in the dc has undergone reduction in cage nd n umber of INTRODUCTION The Osmundaceae is a relatively small family of ferns that consists of three genera, Osmunda, Todea and Leptopteris; it contains about 21 species, depending upon how many of the described species are considered to be varieties of Osmunda regalis L. (Copeland, 1947; Christensen, 1906-34). Indications are that the family has been more abundant and more widely distributed in the geologic past; it can be traced back to the Permian by a series of well-preserved fossil stems. Because of the fine fossil record, the small number of extant species and the isolated phylogenetic position of the family, the Osmundaceae has always been of interest to students of fern evolution. In particular, two questions have attracted a great deal of attention. The most difficult one concerns the origin of the family; for over 100 million years it has displayed a remarkable morphological constancy in those structures where living and fossil members can be compared. Because of this stability, a large phyletic gap exists between the most ancient forms and any supposed ancestors. The second question concerns evolution within the family—in particular, evolution of the stele. Answers to this question have come from comparisons of fossil species with living species. An apparent failing in this method is that fossil stems have generally been compared with the three American species, O. regalis, O. cinnamomea and O. claytoniana, because of the inaccessibility of other species distributed in parts of the world remote from leading universities. It seems tenuous to base evolutionary theories for the family merely on the accidental juxtaposition of investigators and available material. Part of an investigation carried out in the graduate laboratory of the Henry Shaw School of Botany of Washington University and submitted as a thesis in partial fulfillment of the requirements for the degree of Doctor of Philosophy. d. [Vor. 49 58 ANNALS OF THE MISSOURI BOTANICAL GARDEN Andrews (1961) asserts: “it may be worth noting that Engler and Diels recognized 17 living species in their Syllabus der Pflanzenfamilien and since they have not all been studied critically, significant information may be forthcoming from studies of living plants as well as the fossils.” The main purpose of the present work, then, was to investigate as many species as possible in an attempt to relieve the uncertainty which will always exist so long as the morphology of some species in the family remains unknown. MATERIALS AND METHODS Source of the Material—One of the most difficult aspects of this problem was assembling material; the family has an almost cosmopolitan but discontinuous distribution. Thirteen species are included in the present study. The collectors and countries from which material came are summarized below. Osmunda regalis L. H. N. An ba Jr. New Hampshire, India J. W. Clibur ississippi N. H. Ca Massachusetts O. japonica Thunb. S. Hattori Japan . Kodama Japan S. Miki & S. Kokawa Japan O. lancea Thunb. S. Miki & S. Kokawa Japan O. cinnamomea L. H. N. Andrews Jr. New pe T. H. Chung L W., ge Gen ER N. H. Massachusetts K. eebe "Thailand O. claytoniana L. H. N. Andrews Jr. New Hampshire O. javanica Blume T. H. Chung Korea N. Seneviratne Ceylon K. Suvatabandhu Thailand O. vachellii Hook. Herbarium of the a. Botanical Garden with permission of the Curator, R. E. Woodson Jr. O. bromeliaefolia (Pr.) Copel. S. Hattori Japan ari Japan 1962] HEWITSON—-MORPHOLOGY OF OSMUNDACEAE 59 O. banksiaefolia (Pr.) e S. Miki & S. Kokawa Todea barbara (L.) pec Japan R. H. Michie New Zealand N. Potts New Zealand H. B. Ryeroft South Africa S. L. Thrower Australia Leptopteris superba MR Presl T. S. Baylis New Zealand A eben Zealand . Curtis; Botanic Gardens, Glasgow, Scotland L. hymenophylloides (A. Rich.) Presl G. T. S. Bayli New Zealand E. W. Curtis Scotland N. Potts New Zealand L. fraseri (Hook. et Eo Presl H. Anderson Australia 5 L Thrower Australia Treatment of the Material—All species in the Osmundaceae are terrestrial with erect, non-paleate stems. The stems are always deceptively bulky, being covered with a large mass of wiry roots and persistent leaf e structure of such “stems” therefore demands special treatment. To obtain smooth stem sections one must trim away the roots and leaf bases. The rhizomes were then cut up into pieces from one to two centimeters in length in such a way that the proper sequence could be recognized and these pieces were softened in a 1: 1 solution of 48% hydrofluoric acid and 95% ethyl alcohol for a period of two weeks. At the end of this period the material was washed in running tap water for at least 24 hours. The material was then dehydrated in an ethyl alcohol series and embedded in celloidin ac- cording to Method 2 of Johansen (1940). Cross sections were made at 20 microns and longitudinal sections were cut at 10 microns. The trimmed-off leaf bases were treated in a similar manner. All sections were stained in 1% aqueous safra- nin and 0.5% fast green in 95% ethyl alcohol. For studies of the leaf epidermis, pinnae were cleared in 5% NaOH and then transferred to 5% chloral hydrate to complete clearing. After washing, the pinnae were stained according to the ferric chloride-tannic acid method of Nevling (1951). Sporangia from herbarium sheets were rehydrated and mounted directly in glycerine jelly. OBSERVATIONS Osmunda can be distinguished from Todea and Leptopteris by the facts that in Osmunda the fertile pinnae are contracted with little or no vegetative lamina present and the pinnae are articulated at the point of attachment to the rachis. Diels (1898) recognized three subgenera in the genus Osmunda, subgenus Osmunda, subgenus Osmundastrum and subgenus Plenasium. Copeland (1947) pointed out that they are quite natural and could be treated as separate genera, but since the genus Osmunda is equally natural and not overly cumbersome, 60 ANNALS OF THE MISSOURI i : > o N IT MA NG Um SAN: = $ NA VAS Ne 7 : 4 YA T7 T TTLLLD MÀ VARESE 4 Mm NA Sr H MAr ze Ihn Ke LASS 3 EI ANA 8 SS NA ZN dë c N M 4 d Ge WZ AL, A We Bessen Yee 3 ad 4 44 1 AL MAS -— M M 1 > APY se: 3 E Hr P d NA DYES VAN bee. PY RE N Rau. rA Na ise EE Nes A Vie > SZ LEIP», eS WA Se». E N ike AM 7 Y "ugs ZEN A N24 7». “tes D SÉ, Se M A E EZ së D B + bi EE YA bb. Old stems not conspicuously, lenticellate; leaves longer than broad, cori- aceous; sepals Petals so greatly involute as to almost Br eg stamens fixed on the claw; leaves ly exceeding 10 c ; dru rarely less than 16 mm. in length, albuminous ............ NOMOSPERMUM Petals mob eg not dp maces Ae the s — erg rarely less 0 cm in length, Re BUE ana ls connate, the stamens 4, the filaments pistillate flower with 1 sepal and 1 5. CissAMPELOS e, E y RE Staminate flower with 4 sepals, the peta coherent forming a staminal column; petal £5 £5 Ted [Vor. 49 158 ANNALS OF THE MISSOURI BOTANICAL GARDEN l. CHONDRODENDRON Ruiz & Pav. CHONDRODENDRON Ruiz & Pav. Prodr. Fl. Peruv. 132. 1794 (as Chondodendron) ; Miers, in Ann. Nat. Hist. ser. 3. 19: 187. 1867. Botryopsis Miers, loc. cit. ser. 2. 7:43. 1851. Detandra Miers, loc. cit. ser. 3. 13:124. 1864. Twining shrubs. Leaves alternate, petiolate. Staminate inflorescence axillary, fasciculate, paniculiform. Staminate flowers 3-merous, actinomorphic: Sepals 6-22, the exterior minutely bracteoliform, the interior 6 conspicuously larger, usually reflexed or sometimes cucullate at the apex in anthesis; petals 6, minute, ante- sepalous, appressed to the androecium; stamens 6, antepetalous, greatly surpassing the petals, inflexed, the filaments free or slightly coherent at the base, the longitu- dinally dehiscing anthers 2-celled, the cells separated by and sometimes immersed within a fleshy connective which is often produced beyond the thecae. Pistillate inflorescence axillary, solitary or fasciculate, racemose. Pistillate flowers: Sepals and petals similar to the staminate; staminodia minute or none; carpels 6, erect, gibbose-ellipsoid, the style almost wanting, the stigma lingulate and reflex Drupes 6 or a few aborted, gibbose-ellipsoid; endocarp chartaceous-coriaceous, divided by a septum which extends from the base beyond the center; seed deeply hippocrepiform, exalbuminous. About 10 species. Panama, South America. Throughout the literature involving the Menispermaceae Ruiz and Pavön’s original spelling Chondodendron has been adopted by some authors, including Diels, while others have utilized the form Chondrodendron. As pointed out by Sandwith (in Kew Bull. 58. 1955), Ruiz and Pavön stated that they called their new genus Chondodendron because of the “abundance of burrs” on the trun and branches of the “tree,” referring to the verrucose-lenticellate cortex of the woody stems. The name therefore clearly must have been derived from the Greek xóv6poc, “burr” and d£vöpov, “tree” and the name should be spelled Chondroden- dron. Sandwith proposes that the original spelling Chondodendron be treated as an orthographic error, and that the correct spelling Chondrodendron be adopted in the future for this genus. On the basis of the evidence, the spelling Chondroden- dron is adopted for use in this paper. : 1. CHONDRODENDRON TOMENTOSUM Ruiz & Pav. Syst. Veg. 261. 1798. Epibaterium ? tomentosum Pers. Syn. Pl. Š 561. 1807. 1009. Chondodendron scabrum Miers, in Ann. Nat Hist. ser. 2. 7:44. 1851. Botryopsis spruceana Eichl. in Flora 47:394. 1864. pee vues cretosum Miers, in Ann. Nat. Hist. ser. 3. 19:192. 1867, nom. nud.; Contrib. Goce hypoleucum Standl. in Contrib. Arn. Arb. 5:65, pl, 10, 1933. Shrubby twiners; stems striate with the bark of older stems verrucose-lenti- cellate, minutely puberulent to tomentose, rarely glabrous. Leaves petiolate, ovate, deltoid or suborbicular, entire to crenate, the apex obtuse or acute, mucronate or sometimes emarginate, the base cordate, subcordate or truncate, 10-18 cm. long, (470) 1962] FLORA OF PANAMA (Menispermaceae) 159 Figure 148. Chondrodendron tomentosum 9-18 cm. wide, at length subcoriaceous, 5-, 7- or 9-pli-nerved, glabrous above, white and woolly beneath; petioles distally and proximally swollen, 4-14 cm. long, puberulent to tomentose. Staminate inflorescence fasciculate in the leaf axils, paniculiform, 6 cm. or more in length, the rachis tomentose; bracts and bracteoles ovate, about 0.5 mm. long. Staminate flowers (in Panama): Sepals 9, 14, or 16, irregularly imbricate, the 3, 8, or 10 exterior small, broadly ovate, rounded or obtuse at the apex, 0.5-1.4 mm. long, 0.5-1.2 mm. wide, exteriorly tomentose or with the margins ciliolate, the 6 interior conspicuously larger, oblong-ovate, 2.5-3.0 mm. long, 1.5-1.7 mm. wide, glabrous or with the margins ciliolate; petals 6, minute, broadly ovate or lanceolate, acute or acuminate at the apex, 0.5-0.8 mm. long, 0.3-0.5 mm. wide, glabrous; stamens 6, 1.5 mm. long, inflexed, the connective produced beyond the thecae, the filaments free or coherent at the base. Pistillate flower or fruit not seen. (471) [Vor. 49 160 ANNALS OF THE MISSOURI BOTANICAL GARDEN Peru, Bolivia, Colombia and Panama. arro Colorado Island, shore east of Gross Trail, Woodworth & Vestal 438, Gatun “= ee 120 meters or less, Standley 40849, 41007, 41155. Three distinct types of Ge flowers are recognized by Krukoff and Moldenke (in Brittonia 3: 17. 1938) in Chondrodendron tomentosum. Type I is described as having 15 sepals So the outer 9 being 0.7-1.6 mm. long and 0.7-1.3 mm. wide, acute at the apex, fleshy, and slightly connate at the base. The 6 inner sepals are 3-4 mm. long, 1.7-2.5 mm. wide, campanulate-tubular at the base and glabrate except for the more or less fringed-ciliolate margins. The 6 petals are well- developed, broadly elliptic or obovate, about 0.5 mm. long and 0.25 mm. wide and rounded at the apex. The 6 stamens are about 2 mm. long. This is the Botryopsis spruceana type of staminate flower. Type II has (15-) 19-22 sepals about 0.7 mm. long and 0.5 mm. wide which are glabrous throughout. The 6 petals are very minute, about 0.2 mm. long and wide, broadly elliptic and obtuse or rounded at the apex. The 6 stamens are about 1.2 mm. long with the anthers about as long as the filaments. This type represents the true Chondrodendron tomentosum type of staminate flower. Type III is described above and is based upon the Chondro- dendron hypoleucum of Standley. Representatives of the three groups cannot be distinguished vegetatively. 2. ODONTOCARYA Miers. OpoNTOcARYA Miers, in Ann. Nat. Hist. ser. 2. 7: 38. 1851. Chondodendron acc. to Benth. & Hook. f. Gen. 1:34. 1862, not Ruiz & Pav. Twining or scrambling shrubs. Leaves alternate, petiolate. Staminate in- florescence axillary, solitary, racemiform, rarely paniculiform. Staminate flowers 3-merous, actinomorphic: Sepals 6-18, the exterior bracteoliform, the interior conspicuously larger, imbricate; petals 6, carnose, antesepalous; stamens 6, ante- petalous, the filaments more or less coherent, dehiscence longitudinal. Pistillate inflorescence axillary, solitary, racemose. Pistillate flowers: Sepals and petals similar to the staminate; staminodia 6 or wanting; carpels 1-3, gibbose-oblong, the stigma sessile, 3-lobed. Fruit an ovoid-ellipsoidal drupe. About 10 species. Lesser Antilles, Central America and South America. a. Leaves lustrous; staminate inflorescences borne on p congested secondary branches; sepals 18; petals 0.6-0.7 mm. long, pubesc 1. O. NITIDA D £0 Leaves dull; staminate inflorescences arising Case from main stem; sepals 6; petals 0.7 mm. or more in length, glabrous, . Leaves with glandular areas between the base of the midrib and lowest diverging nerves, the base usually cordate; staminate inflorescence racemi- form; petals 1.5-2.6 mm. long ee ae .2. O. PAUPERA bb. Leaves not glandular, pu base usually truncate; staminate inflorescence Se E petals 0.7-1.2 mm. long 3. O. TRUNCATA (472) 1962] FLORA OF PANAMA (Menispermaceae) 161 1. ODONTOCARYA NITIDA Riley, in Kew Bull. 119. 1927. Scrambling shrubs; stems striate, puberulent. Leaves petiolate, the lamina broadly ovate, entire, the apex obtusely caudate-acuminate, the base cordate, 5-11 cm. long, 3-6 cm. wide, subcoriaceous, glabrous above, dark and lustrous, palmately 5- to 7-nerved, puberulent below; petioles 2-4 cm. long, pubescent. Staminate in- florescence a racemiform thyrse, axillary, borne on a short congested leafy secon- dary branch, 3-6 cm. long, the rachis pubescent; bracts linear, about 1 mm. long. densely pubescent; bracteoles linear, 0.5 mm. long, densely pubescent. Staminate flowers: Sepals 18, ovate to elliptic, 0.8-2.1 mm. long, 0.5-1.2 mm. wide, carnose, the exterior densely pubescent; petals 6, obovate, the apex truncate or rounded, m. long, 0.3-0.5 mm. wide, carnose, the exterior densely pubescent; stamens 6, 0.7-0.8 mm. long, glabrous, the filaments coherent at the base. Pistillate flower and fruit not seen. Endemic to Panama. CANAL ZONE: Cocoli, Riley 136. Riley recognizes no calyx or corolla and refers to the perianth as being com- prised of tepals. The six innermost perianth segments, however, are conspicu- ously small and interrupt a transitional increase in size from the outermost whorl of segments inward. An anatomical distinction exists in that the six innermost segments are provided with a single median nerve characteristic of petals while the outer segments are provided with a single median plus a pair of lateral nerves characteristic of sepals. Bearing in mind the anatomical evidence present and the fact that the family is characterized by the presence of comparatively small petals, I consider the perianth to be dichlamydeous. 2. ODONTOCARYA PAUPERA (Griseb.) Diels, in Engl. Pflanzenr. 4.94: 172. 1910. Chondodendron hederaefolium Miers, in Ann. Nat. Hist. ser. 2. 7:38. 1851, nom. nud. Cocculus pauper Griseb. in Gotting. Abh. 7:162. 1857. Chondodendron tomentosum Benth. in Journ. Linn. Soc. 5: suppl. 2. 47. 1861. Odontocarya hederaefolia Miers, in Contrib. Bot. 3. 64. 1871. Odontocarya scabra Miers, loc. cit. 65. 1871 Climbing or scrambling shrubs; older stems thick and fleshy, conspicuously lenticellate, the younger stems striate, glabrous or essentially so. Leaves petiolate, the lamina broadly ovate to subhastate, entire, the apex acute or obtuse, mucronate, the base cordate or rarely obscurely truncate, 4.5-9.5 cm. long, 3.0-6.0 cm. wide, membranous, pubescent or puberulent above with conspicuous glandular areas between the base of the midrib and lowest diverging nerves, pubescent below; petiole 2.0-8.0 em. long, pubescent or puberulent. Staminate inflorescence axil- lary, racemiform, about 7 cm. long, the rachis puberulent; bracts and bracteoles about 1.0-1.5 mm. long. Staminate flowers: Sepals 6, the 3 exterior ovate or elliptic, sparsely toothed, 1.2-2.0 mm. long, 0.6-1.0 mm. wide, membranous, essentially glabrous, the 3 interior conspicuously larger, obovate, the apex obtuse or rounded, 1.8-3.5 mm. long, 1.5-2.2 mm. wide, glabrous; petals 6, elliptic, carnose, 1.5-2.6 mm. (473) [Vor. 49 162 ANNALS OF THE MISSOURI BOTANICAL GARDEN Figure 149. Odontocarya paupera (474) 1962] FLORA OF PANAMA (Menispermaceae ) 163 long, 1.0-1.8 mm. wide, glabrous; stamens 6, 1.0-1.8 mm. long, the filaments more or less coherent. Pistillate inflorescence an axillary raceme, about 9 cm. long, the rachis puberulent; bracts and bracteoles about 1.0 mm. long. Pistillate flowers: Sepals and petals similar to the staminate; carpels 1-3, gibbose, free, sessile, the style not evident, the stigma 3-lobed. Drupe ovoid, about 1 cm. long, 0.5 cm. wide; endocarp semi-ellipsoid, verrucose, ribbed. Panama, Lesser Antilles and northern South America. : Balboa, moist thicket, Standley 27130; in lati Garden, oe. 41189; vicinity o Wise G. White 121; locality sree Johansen 70. PANAMÁ: Isla Taboga, ca. 0-186 m., Woodson, Allen & Seibert 1486; s Sa eneen Zeg thicket, Ski 40778, Paul 599; San José Island, Perlas Archipelago, Goen 581, 546, Johnston 7, 786. There has been considerable taxonomic confusion and misdetermination be- tween this species and Odontocarya nitida. The two species are readily dis- tinguished, however, as Odontocarya paupera possesses distinctive glandular areas on the leaf, an essentially glabrous flower with six sepals, and staminate in- florescences which are borne upon the main stem of the vine. Odontocarya nitida has a leaf devoid of glandular areas, a pubescent flower with 18 sepals, and staminate inflorescences which are borne on short congested secondary branches. 3. ODONTOCARYA TRUNCATA Standl. in Journ. Arn. Arb. 11: 121. 1930. Twining shrubs; stems striate, conspicuously verrucose-lenticellate, glabrous. Leaves petiolate, the lamina ovate, the apex acuminate, the base truncate or ob- scurely cordate, 7-12 cm. long, 5-9 cm. wide, at length subcoriaceous, glabrous, somewhat bullate above; petioles 3-4 cm. long, glabrous. Staminate inflorescence axillary, paniculiform, 8-24 cm. long, the rachis glabrous; bracts 1.0-1.5 mm. long, mm. wide; bracteoles scarious, ovate to lanceolate, 0.5-1.0 mm. long. Staminate flowers: Sepals 6, glabrous, the 3 exterior obscurely coherent at the base, ovate to lanceolate, membranous, 0.5-0.8 mm. long, 0.3-0.6 mm. wide, the 3 interior con- spicuously larger, ovate to elliptic, the apex obtuse or rounded, the base truncate or rounded, 1.3-1.7 mm. long, 1.0-1.4 mm. wide; petals 6, cuneate, carnose, 0.7-1.2 mm. long, 0.5-0.8 mm. wide, glabrous; stamens 6, about 1 mm. long, the filaments coherent at the base. Pistillate flower not seen. Drupe ovoid, yellow about 1.5 cm. long, 1.0 cm. wide. Endemic to Panama. CANAL : Barro Colorado Island, Bangham 541, Salvoza 960; Las Cascadas Planta- tion, near Sumali, Standley 29519; Margarita Swamp, south of France Field, Maxon & Valentine 7046. 3. ANOMOSPERMUM Miers. ANOMOSPERMUM Miers, in Ann. Nat. Hist. ser. 2. 7: 39. 1851. Woody climbers. Leaves alternate, petiolate. Staminate inflorescence axillary, solitary or fasciculate, racemiform or paniculiform. Staminate flowers 3-merous, actinomorphic: Sepals 6, the exterior bracteoliform, the interior conspicuously larger, imbricate; petals 6, so greatly involute as to almost conceal the stamens (475) [Vor. 49 164 ANNALS OF THE MISSOURI BOTANICAL GARDEN by forming a “pseudodisc”; stamens 6, free, dehiscence longitudinal. Pistillate in- florescence axillary, racemose or sessile and umbelliform. Pistillate flowers: Sepals and petals similar to the staminate; staminodia 6; carpels 3, free; stigma sub- sessile or sessile, lingulate. Drupes 1-3, excentrically stipitate, erect or curved; albumen copious, ruminate. About ll species. South America and Panama. The only representative of this genus thus far collected in Panama (COCLE, El Valle de Antón, Allen 3538) is perhaps a new species but the absence of flower- ing material prevents an adequate analysis at this time. "This specimen is closely related to Anomospermum reticulatum (Mart.) Eichl. of the Amazon basin in that the extremely dense venation reticulum of the lamina is as conspicuous be- neath as above, but differs in the glabrous petioles and the absence of any pubes- cence on the lower leaf surface. 4. HYPERBAENA Miers. HyPERBAENA Miers, in Ann. Nat. Hist. ser. 2. 7: 44. 1851. Pachygone Eichl. in Denkschrift. Bot. Gesellsch. Regensb. 5:1. 1864. Woody twiners or sometimes erect shrubs or small trees. Leaves alternate, petiolate. Staminate inflorescence axillary, fasciculate, spiciform or racemiform. Staminate flowers 3-merous, actinomorphic: Sepals 6, the exterior bracteoliform, the interior conspicuously larger, imbricate; petals 6, antesepalous; stamens 6, ante- petalous, free or the filaments connate at the base, dehiscence longitudinal. Pistil- late inflorescence axillary, solitary, racemose. Pistillate flowers: Sepals and petals similar to the staminate; staminodia 6; carpels 3, free, gibbose; style reflexed. Drupes sessile, curved; exalbuminous. About 40 species. Mexico, Central America, South America and the Antilles. Woody twiners; es palmately 5-nerved; inner sepals 2.5-4.0 mm. long, dency ee nt; petals pubescent; stamens cucullate, pubescent ........ 1. H. PANAMENSIS aa. Small trees; leaves ER erved; inner sepals 1.1-1.4 mm. long, glabrous or essentially so; petals glabrous; stamens erect, glabrous ..............————— 2. H ALLENII 1. HYPERBAENA PANAMENSIS Standl. in Journ. Wash. Acad. Sci. 15:474. 1925. Hyperbaena hondurensis Standl. in Field Mus. Pub. Bot. 4:305. 1929. vines; stem striate with the bark of older stems conspicuously lenticellate, puberulent or glabrous. Leaves petiolate, elliptic, ovate or oblong, entire, the apex abruptly acute, acuminate or obtuse, mucronate, the base obtuse, rounded or truncate, 6-24 cm. long, 3-14 cm. wide, at length coriaceous, palmately 5-nerved, frequently with a pinnate aspect, the basal nerves frequently slender and inconspicuous, glabrous above, sparsely and minutely puberulent or glabrous be- low; petioles swollen distally, 1-9 cm. long, glabrous to puberulent. Staminate inflorescence fasciculate or solitary in the leaf axils, racemiform, up to 20 cm. in length, the rachis pubescent to tomentose; bracts and bracteoles ovate to linear, about 1 mm. long, densely pubescent. Staminate flowers: Sepals 6, the 3 outermost small, ovate, 0.7-1.2 mm. long, 0.5-0.8 mm. wide, exteriorly pubescent, the 3 (476) 1962] FLORA OF PANAMA (Menispermaceae) 165 innermost conspicuously larger, broadly ovate, 2.5-4.0 mm. long, 2.2-2.5 mm. wide, exteriorly pubescent; petals 6, minute, spatulate to oblong, about 0.5 mm. long, exteriorly pubescent; stamens 6, cucullate, about 1.3 mm. long, densely pubescent, the anthers immersed within the connective, the filaments free or rarely coherent at the base. Pistillate inflorescence solitary or fasciculate in the leaf axils, racemose, about 8 cm. long, the rachis tomentose; bracts and bracteoles ovate to linear, about 0.5-1.0 mm. long, densely pubescent. Pistillate flowers: Sepals and petals similar to the staminate; staminodia 6, about 2.5 mm. long, pubescent at the base; carpels 3, gibbose, sessile, the style not evident, the stigma radiate-incised. Drupe obovoid, 1.5-2.5 cm. long, puberulent or glabrous. Panama, Costa Rica, Guatemala, Honduras and British Honduras. CANAL ZONE: Barro Colorado Island, Bangham 484, Bailey 101, Standley 40978, 40800, 40787, 41077, Wetmore & Woodworth 76; Gamboa, Standley 28417. Considerable deliberation involved the placing of Hyperbaena hondurensis into synonomy. In 1925 Standley described Hyperbaena panamensis (Standley 28417) from sterile material. The distinguishing characteristic was the venation. A palmately 5-nerved condition provided a pair of slender and inconspicuous basal nerves, a prominent midrib, and a prominent pair of inner nerves extending al- most to the apex of the leaf so as to impart a subparallel effect. In 1929 Standley described Hyperbaena hondurensis (Standley 55220), once again from sterile material. A palmately 5-nerved condition existed here also but the inner pair of nerves extended approximately half the length of the leaf and several lateral nerves originated from the midrib. This afforded a pinnate aspect to the leaf. Specimens from Panama and Costa Rica illustrate the Hyperbaena panamensis type of venation while those of Honduras, Guatemala and British Honduras the H. hondurensis type. Unfortunately, no specimens from Nicaragua are available. The Hyperbaena panamensis type of leaf is provided with somewhat more pubes- cence on the lower surface and has a somewhat shorter petiole than the Hyper- baena hondurensis type. Vegetatively, it appears that two species exist. However, floral examination involving the two types indicates the opposite view as no floral distinction can be formulated between the two. Furthermore, the pubescent petals, cucullate stamens, and connivent carpels present very distinctive floral characteristics thereby making their appearance in separate species quite unlikely. Along this line of thought is the marked contrast between the floral characteristics noted above and allied species of the genus, for example, Hyperbaena allenii. In summary it can be stated that although considerable variation exists vegetatively, the floral similarity between Hyperbaena panamensis and H. hon- durensis unite them as one species—H. panamensis by precedence. 2. HYPERBAENA ALLENI Standl. in Ann, Missouri Bot. Gard. 27 :310. 1940. Small trees; young stems striate, densely pubescent. Leaves petiolate, lanceo- late-oblong, entire, the apex acute or obtuse, the base obtuse or rounded, 14-23 cm. (477) 166 ANNALS OF THE MISSOURI BOTANICAL GARDEN Figure 150. Hyperbaena allenii (478) [Vor. 49 1962] FLORA OF PANAMA (Menispermaceae) 167 long, 4.5-6.0 cm. wide, subcoriaceous, pinnately nerved, glabrous above, puberulent below; petioles swollen distally, 1.5-3.0 cm. long, densely pubescent. Staminate inflorescence axillary, fasciculate, spiciform, 0.5-3.0 cm. long, the rachis densely pubescent; bracts and bracteoles about 0.5 mm. long, densely pubescent. Staminate flowers; Sepals 6, the 3 outermost small, obovate, obscurely toothed, 0.7-0. long, 0.5-0.6 mm. wide, exteriorly pubescent, the 3 innermost conspicuously larger, obovate, obscurely toothed, 1.1-1.4 mm. long, 0.7-0.9 mm. wide, glabrous or with the margins ciliolate; petals 6, obovate, 0.5-0.8 mm. long, 0.3-0.5 mm. wide, glabrous; stamens 6, free, about 1 mm. long, glabrous. Pistillate flower not seen. Fruit ovoid, about 2 cm. long, glabrous. Endemic to Panama. coc a rim of El Valle ie Antón, 600-1000 m., Allen 1656; north of El Valle de Antón, trail 4 Las Minas, alt. a 000 m., Allen 2465; Loma del Tigre, region north of El Valle de Ann alt. 1000 m., Allen 3807. 5. CISSAMPELOS L. CissaMPELos L. Sp. Pl. 1031. 1753. Dissopetalum Miers, in Ann. Nat. Hist. ser. 3. 17:267. 1866. Twining shrubs, rarely erect. Leaves alternate, petiolate, more or less peltate or basifixed. Staminate inflorescence axillary, generally fasciculate, normally originat- ing in a leafy axil as multiflowered dichasia or the dichasia originating on axillary secondary branches in the axils of reduced leaves or bracts. Staminate flowers 2-merous, actinomorphic: Sepals 4, usually exteriorly pubescent; petals connately cupulate; stamens 4, the anthers sessile on a column formed by the connate fila- ments, dehiscence transverse. Pistillate flowers fasciculate in the axils of reduced leaves or bracts upon secondary, frequently fasciculate, axillary branches, zygomor- phic: Sepal 1, obovate, exteriorly pubescent; petal 1, antesepalous; carpel 1, free, gibbose, the style short, the stigma 3-lobed. Drupes subglobose; endocarp osseous, verrucose, ribbed. A troublesome genus of about 25 species. North and South America, Africa, Asia and Australia. a. nit conspicuously peltate, the base usually truncate or rounded, indument present sericeous with long whitish hairs; bracts of secon d axillary branche large and foliaceous; carpels essentially glabrous; gie mm. ong C. TROPAEOLIFOLIA aa. Leaves generally obscurely mw. or basifixed, the base cordate, rarely trun- cate or rounded, in us or tomentose with short reddish-yellow, yellowish-brown itish geg bracts of secondary axillary branches 3r t Le E c © g = (b o = d $ c Bos S, i a SR Do et 5 zu "n ve a monly arising from the axils of bracts upon secondary, gener s of EES and pistillate "erf with m. long. 2. C. FASCICULATA Leaves usually more or less pu — a. am below; staminate inflor- escence commonly arising from normal leaves; sepals of staminate and pistillate flowers duni opi i red spotting; drupes 4-5 mm. long. EE 3. C. PAREIRA c c [Vor. 49 168 ANNALS OF THE MISSOURI BOTANICAL GARDEN The frequent appearance of Cissampelos grandifolia Triana & Planch. in Costa Rican and Colombian collections suggests its probable presence in Panama. Although several specimens have been misdetermined as Cissampelos grandifolia, I do not find the species to have been collected in Panama. The species is dis- tinguished by its much-branched and spreading paniculiform inflorescence and the large, conspicuously peltate, broadly ovate to suborbicular, long-petiolate leaves. 1. CissAMPELOS TROPAEOLIFOLIA DC. Syst. 1:532. 1818. Cissampelos membranacea Triana & Planch. in Ann. Sc. Nat. ser. 4. 17:43. 1862. Subherbaceous twiners; stems striate, pubescent or puberulent, rarely glabrous. Leaves petiolate, conspicuously peltate, ovate to suborbicular, entire to crenate, the apex obtuse, rounded or acuminate, mucronate, the base truncate or rounded, rarely cordate, 5-11 cm. long, 5-10 cm. wide, membranous, palmately nerved, sericeous with long whitish hairs or glabrous above, paler and occasionally glaucous below, sericeous or puberulent; petioles 4-9 cm. long, tomentose to glabrous. Staminate inflorescence multiflowered fasciculate dichasia in the leaf axils or upon short secondary axillary branches within the axils of reduced leaves or bracts, the two forms frequently occurring together; bracts of secondary branches large and foliaceous, basifixed, sessile or obscurely petiolate, broadly cordate to subreniform, mucronate, 5-15 mm. long, 4-25 mm. wide, membranous, puberulent to densely pubescent; bracteoles linear, about 0.5 mm. long, pubescent. Staminate flowers: Sepals 4, ovate, oblong or obovate, 1.0-1.4 mm. long, 0.8-1.0 mm. wide, glabrous or exteriorly puberulent; corolla campanulate, 0.5-1.0 mm. in diameter, glabrous; anthers 4, glabrous. Pistillate inflorescence composed of individual flowers fascicu- late in the axils of bracts upon secondary axillary branches. Bracts and bracteoles similar to the staminate. Pistillate flowers: Sepal 1, ovate to obovate, 1.0-1.2 mm long, 0.8-1.0 mm. wide, glabrous or puberulent; petal 1, suborbicular, 0.5-0.7 mm. long, 0.5-1.0 mm. wide; carpel 1, gibbose, sessile, essentially glabrous, the stigma 3-lobed. Drupe red, obovoid, compressed, 5-7 mm. long, 4-5 mm. wide, pubes- cent, rarely glabrous; endocarp ribbed, verrucose. Southern Mexico, Central America and northern South America. BOCAS DEL TORO: Chiriqui Lagoon, Von Wedel 1853, 1119, 1376, 1372, 1035; Isla Colon Von Wedel 46, 2947; Old Bank Island, Von Wedel 2083, 1888, 1991, 1985; Water Valley Von Wedel 673, 1819, 1490; Chiriqui Lagoon region, Punta Rovalo and Rovalo River, Seibert 1563; Columbus Island, Von Wedel 23; no sa er but bus bled. Chiriqui s ace LON: betw t y Remedios, ca. 15-50 m., Woodson, Allen & Seibert 1187; between R. Tinta and R. Tabasará, along main highway, Woodson; Allen & Seibert 414; vicinity of San Félix, alt. 0 to 120 m., Pittier 5458. DARIEN: Cana and vicinity, 2000-6500 ft. el., Williams 778; Paca below Cana, Williams 767. (480) 1962] FLORA OF PANAMA (Menispermaceae) 169 rh IL S fh Loi C. m x £d Di H PA ds 2 tru e chat Wil f Ge A ek f nu H Las Figure 151. Cissampelos tropaeolifolia (481) [Vor. 49 170 ANNALS OF THE MISSOURI BOTANICAL GARDEN 2. CissAMPELOS FASCICULATA Benth. in Lond. Journ. Bot. 2:361. 1843. Cissampelos scutigera Triana & Planch. in Ann. Sc. Nat. ser. 4. 17:42. 1862. Cissampelos myriocarpa Triana & Planch., loc. cit. 1862 Cissampelos floribunda Miers, in Ann. Nat. Hist. ser. 3. 17:135. 1866. Cissampelos coriacea Standl. in Field Mus. Pub. hot 18:437. 1937. Shrubby twiners; stems striate, puberulent to tomentose. Leaves petiolate, conspicuously or obscurely peltate or basifixed, broadly ovate to suborbicular, entire, the apex obtuse, rounded or acute, mucronate, the base cordate or truncate, rarely rounded, 4-14 cm. long, 5-15 cm. wide, membranous to subcoriaceous, palmately 5- to 9-nerved, puberulent and dark above, paler below and sericeous or tomentose with reddish-yellow, yellowish-brown or whitish hairs; petioles twisted and swollen proximally, 4-11 cm. long, pubescent or more commonly tomentose. Staminate inflorescence multiflowered fasciculate dichasia originating upon secondary axillary branches within the axils of reduced leaves or bracts, rarely axillary from normal leaves; peduncle of cymes at length 1.5 cm. long, densely pubescent; bracts of secondary branches basifixed, sessile or petiolate, ovate to subreniform, mucronate, generally conspicuously involute, pubescent; bracteoles about 1 mm. long, pubes- cent. Staminate flowers: Sepals 4, obovate, 0.8-1.5 mm. long, 0.5-0.8 mm. wide, exteriorly pubescent, the interior with conspicuous red spotting; corolla cupulate, 0.6-1.0 mm. in diameter, sparsely pubescent or glabrous; anthers 4, glabrous. Pistillate inflorescence composed of individual flowers fasciculate in the axils of bracts upon fasciculate secondary, frequently branched, axillary branches; bracts and bracteoles similar to the staminate. Pistillate flowers: Sepal 1, obovate, 1.0-2.0 mm. long, exteriorly pubescent, the interior with conspicuous red spotting; petal 1, suborbicular, 0.5-0.8 mm. long, 0.7- 1.0 mm. wide; carpel 1, gibbose, pubescent. Drupe obovoid, compressed, 2-4 mm. long and wide, pubescent; endocarp ribbed, verrucose. Nicaragua, Costa Rica, Panama and northern South America. Qui: vicinity of Bajo Mona and Quebrada Chiquero, alt. 1500 m., Woodson & Soha | 545. vicinity of Bajo Chorro, alt. 1900 m., Woodson & Schery 682 After extensive examination of available material no significant distinction could be ascertained between Cissampelos coriacea Standl. and C. fasciculata, there- fore the former is placed in synonomy. Vegetatively this species tends to be quite variable. Specimens from Costa Rica habitually bear large obscurely peltate or basifixed leaves which are broadly cor- date. Panamanian and Colombian material, on the other hand, has a tendency toward smaller suborbicular leaves which are conspicuously peltate. 3. CissAMPELOS PAREIRA L. Spec. Pl. 1031. 1753. Cissampelos pareira « L., loc. cit. 1753. Cissampelos caapeba L. loc. cit. 1032. 175 1962] FLORA OF PANAMA (Menispermaceae) 171 Cissampelos discolor DC. loc. cit. 1818. Cissampelos tomentosa DC, loc. cit. 1818. Cissampelos dei Willd. ex DC. loc. cit. 536. 1818. Cissampelos argentea H.B.K. Nov. Gen. : Sp. 5:67. 1821. Clan ua yaquilens’s HR K. loc. cit. 1 Cissampelos orinoce H.B.K. loc. cit. 68. et monoica ce Hil. Fl. Bras, Mer. SCH 1825. Cissampelos australis St. Hil. loc. cit. 56. 182 Cissampelos glaucescens Triana & Planch, in Ann. Sc. Nat. ser. 4. 17:41. 1862. i . 18 Cissampelos longipes Miers, in Ann. Nat. Hist. ser. 3. 17:134, 1866, nom. nud.; Contrib. Bot. 3:139. 1871. Cissampelos testudinaria Miers, loc. cit. 1866, nom. nud.; loc. cit. 143. 1871. Cissampelos limbata Miers, loc. cit. 1866, nom. nud.; lo T 7 i i nom.n issampelos pareira var. 6. monoica Eichl. in Mart EE 1894. Cissampelos pareira f. reinformis and f. emarginato- ki Chod. & Hassl. in Bull. Herb. ser. 2. 3:420. 1 Shrubby twiners; stems striate, glabrous to tomentose. Leaves petiolate, ob- scurely peltate or basifixed, broadly ovate, entire, the apex obtuse, rarely acuminate, frequently somewhat emarginate, mucronate, the base truncate or cordate, 2-12 cm. long and wide, membranous or papery, palmately 3- to 7-nerved, in youth more or less sericeous-tomentose on both sides, at length above more or less pubescent, rarely glabrous, below frequently paler, tomentose or pubescent, rarely entirely glabrous. Staminate inflorescence multiflowered fasciculate dichasia in the leaf axils, sometimes upon secondary axillary branches within the axils of reduced leaves or bracts; bracts of secondary branches basifixed, sessile or obscurely petiolate, ovate to subreniform, mucronate, about 1 cm. long and wide to minute, mem- branous, pubescent to tomentose; bracteoles linear, about 0.5 mm. long, pubescent. Staminate flowers: Sepals 4, ovate to obovate, exteriorly pubescent, 1.0-2.0 mm. long, 0.5-1.2 mm. wide; corolla campanulate, 0.8-1.8 mm. in diameter; anthers 4, glabrous. Pistillate inflorescence composed of individual flowers fasciculate upon secondary axillary branches within the axils of reduced leaves or bracts; bracts basifixed, sessile or obscurely petiolate, broadly ovate to reniform, mucronate, about 1.5 mm. long, 2.0 mm. wide to minute, membranous, puberulent to tomentose. Pistillate flowers: Sepal 1, obovate, 1.0-2.0 mm. long, 0.5-0.8 mm. wide, exteriorly pubescent; petal 1, broadly obovate to reniform, 0.5-0.8 mm. long, 0.7- 1.5 mm. wide, glabrous or puberulent; carpel 1, gibbose, densely pubescent. Drupe broadly obo- vate or suborbicular, compressed, 4-5 mm. long, 3-4 mm. wide, pubescent or puberulent; endocarp verrucose, ribbed. (483) [Vor. 49 172 ANNALS OF THE MISSOURI BOTANICAL GARDEN Southern Mexico, Central America, South America, Antilles, eastern Africa, India, Malaya, Indonesia, Philippines, northern Australia and Madagascar. AS DEL TORO: Chiriqui Lagoon, Von Wedel 1309; Shepherd Island, Von Wedel 2691. BOC CANAL ZONE: Barro Colorado Island, Aviles 921, 57, 888, Shattuck 476, Bailey & Bailey 575, Wetmore & Abbe 177, 178; Bismark above Penonomé, Williams 536; Balboa, Standles 5 e m. Pittier 3796; Obispo, Standley 31773; Rio Goen near Culebra, alt. 50 to 100 m., Pittier 2092; Fort Kobbe road, Woodson, Allen & Seibert 1425, 1426; thickets north of Summit, Woodson, Allen & Seibert 766; vicinity of Miraflores, P. White 120, G. White 122; western slope of Ancón Hill, Woodson, Allen & Seibert 1324; around Culebra, alt 50 to 150 m., Pittier 2218; Aes Experiment Station, 3 miles east of Panama City, Maxon, Harvey & Valen- tine ; iRIQUÍ: Boquete, Davidson 611; Cerro Vaca, eastern Chiriquí, in forest or in thickets, E "900 t o 1136 m.. Pittier 5323. ‘cock: El Valle de Anton, alt. 600 m., Allen 3705; i i m DE y missing, Macbride 2699. panamá: Bella Mase Bro. Heriberto 2 = ds of Bella Me s : Dias Sedi Paul 21, 40; Sabanas near Chepo, 30 m., Hur & Allen 9 Diels divides Cissampelos pareira into nine varieties of which only two, var. a. typica and var. y. haenkeana, occur in Panama. The former is characterized by having comparatively small, ovate, more or less peltate leaves which are pale be- low and more or less pubescent or tomentose above and below. This is the most common variety and is pantropic in distribution. The latter variety has similar characteristics but differs in having a larger leaf, 4-9 cm. long and wide, which is obscurely peltate and rarely ovate. To avoid possible taxonomic confusion the remaining varieties will not be discussed as they have little or no bearing on the Panamanian flora. The synonomy for Cissampelos pareira has been restricted to American varieties for similar reasons. (484) 1962] FLORA OF PANAMA (Magnoliaceae) 173 MAGNOLIACEAE By J. E. DANDY Trees or shrubs, glabrous or with an indumentum of simple hairs; wood heteroxylous; branches annulate at the nodes with the scars of fallen stipules. Leaves alternate, stipulate, petiolate, simple; stipules large, free from the petiole or adnate to it, at first enclosing the young growths, later deciduous and leaving an annular scar round the node; lamina penninerved, entire or rarely 2- or more-lobed. Flowers terminal or axillary, usually solitary, pedunculate, actinomorphic, herma- phrodite or rarely unisexual, entomophilous, usually large and fragrant; peduncle bearing 1 or more deciduous spathaceous bracts which at first enclose the flower- bud and after falling leave annular scars. Perianth 2- or more-cyclic, 3-6-merous; tepals 6 or more, free, imbricate, usually subequal and fleshy but the outer whorl sometimes reduced in size or texture so as to simulate a calyx. Androecium of numerous free stamens spirally arranged; filaments short or more or less elongated; anthers linear, 2-thecous, introrse to extrorse, dehiscing by longitudinal slits; connec- tive often produced above the anther-thecae into an appendage. Gynoecium super- ior, sessile or stipitate; carpels numerous to few (rarely 1), spirally arranged, free or sometimes concrescent; ovules 2 or more, biseriate on the ventral suture. Fruit apocarpous or sometimes syncarpous; fruiting carpels dehiscing longitudinally or more rarely circumscissile or indehiscent, sometimes samaroid. Seeds 1 or more in each fruiting carpel, large, suspended (when carpel dehiscent) by a silky thread; testa arilloid or sometimes adherent to the endocarp; endosperm copious, oily; embryo minute. Species about 220, mostly in southeast Asia, the others in tropical America and southeastern North America. The two tropical American genera, Magnolia and Talauma, belong to the tribe Magnolieae and are both more strongly represented in Asia than in America. Some species of the Asiatic genus Michelia L., notably M. champaca L. and M. figo (Lour.) Spreng., are widely planted in the tropics; this genus is distinguished from Magnolia and Talauma by its axillary flowers with a stipitate gynoecium. a. Carpels free, longitudinally dehiscent in fruit, the valves persistent; stipules free from the petiole, the latter therefore unscarred 1. MAGNOLIA aa. Carpels concrescent at least towards the base, woody and circumscissile in fruit, the upper portions falling away, the lower portions persistent and bearing the suspended seeds; stipules adnate to the petiole, leaving a scar on its upper surface Eeer 2. TALAUMA 1. MAGNOLIA L. Macnozia L. Sp. Pl. 1: 535. 1753. Evergreen or deciduous trees or shrubs. Stipules free from or adnate to the petiole. Flowers terminal, hermaphrodite, at first enclosed in 1 or more spatha- ceous bracts. Gynoecium sessile or shortly stipitate; carpels numerous or few, free; ovules 2, rarely 3-4 in the lower carpels. Fruiting carpels dehiscing longitudinally, the valves persistent. (485) [Vor. 49 174 ANNALS OF THE MISSOURI BOTANICAL GARDEN Species about 80, the majority in southeast Asia, the others in tropical America and southeastern North America. The Panama species belongs to Sect. Theorhodon Spach, an American section which extends from the southeastern United States to the Greater Antilles and through Mexico and Central America to northern South America. This section differs from the other American sections in its free stipules. It includes M. grandiflora L., native of the southeastern United States, which is widely planted in the tropics. Many other species of Magnolia are cultivated in temperate gardens. 1. MAGNOLIA soRORUM Seibert, in Ann. Missouri Bot. Gard. 25: 828. 1938. Tree up to 30 m. tall; indumentum rufous or tawny; branchlets densely villous-pubescent when young. Leaf-lamina elliptic or ovate-elliptic to elliptic- or ovate-oblong, cuneate to obtuse or rounded at the base, obtuse to subacute or subacuminate at the apex, up to about 20 cm. long and 10 cm. broad, coriaceous, at first pubescent above on the midrib towards the base, densely pubescent beneath; lateral nerves about 10-14 on each side of the midrib; petiole up to about 3 cm. long, at first densely villous-pubescent, sometimes glabrescent; stipules villous- pubescent outside. Flowers very fragrant; bud ellipsoid; bracts 1-4, the uppermost or single one inserted immediately below the flower, densely villous-pubescent outside; peduncle stout, about 1.5-4 cm. long, densely villous-pubescent. Tepals 9, 3-merous, white; outer 3 obovate-oblong, about 4-7 cm. long, sparingly pubescent outside at the very base; inner 6 up to about 8 cm. long, glabrous. Stamens about 13-15 mm. long. Gynoecium ellipsoid; carpels about 20-45, densely villous- pubescent. Fruit ellipsoid to ellipsoid-oblong, about 4.5-6 cm. long; fruiting carpels shortly beaked. Endemic to the mountains of western Panama, at 1100-2500 m. altitude. Hess reports the vernacular name vaco in Chiriquí. CHIRIQUÍ: valley of the upper Rio Chiriquí ae alt. 1300-1900 m., White & White 21, 239; vicinity of Casita Alta, Volcán de Chiriqui, alt. P red > Woodson, Allen & Seibert 916; vicinity of Casita Alta, north fores ted | ace GË Copete, me spur of Volcán de Chiriqui, alt. 2400-2550 m., Allen 4874; vicinity of Finca Lérida, a 1650 m. n 4729; Chiriquí, Hess 261, 263. cocı&: El Valle de Antón, crest of Cerro ps alt. 1100 m., Allen 3752. The handsome rufous or tawny indumentum readily distinguishes this species from the related M. poasana (Pittier) Dandy, which occurs to the west in Costa Rica. 2. TALAUMA Juss. TALAUMA Juss. Gen. Pl. 281. 1789. Svenhedinia Urb. in Fedde, Rep. Sp. Nov. 24:3. 1927. Evergreen trees or shrubs. Stipules adnate to the petiole. Flowers terminal, hermaphrodite, at first enclosed in 1 or more (usually several) spathaceous bracts. Gynoecium sessile; carpels numerous or few, concrescent at least towards the base; ovules 2. Fruiting carpels woody, circumscissile, the upper portions falling away (486) 1962] FLORA OF PANAMA (Magnoliaceae) 175 Figure 152. Magnolia sororum (487) [Vor. 49 176 ANNALS OF THE MISSOURI BOTANICAL GARDEN either separately or in irregular masses, the lower portions persistent and bearing the suspended seeds. Species about 45, mostly in tropical and subtropical southeast Asia, the others in tropical America in the Lesser Antilles, Cuba, southern Mexico and through Central America to Brazil. The genus is essentially tropical and lacks the strong North Temperate element present in Magnolia. a. Leaves rounded to obtuse at the apex, rather laxly reticulate .................... 1. T. GLORIENSIS aa. Leaves bluntly acuminate to subacute or apiculate at the apex, closely reticulate 2. T. SAMBUENSIS l. TALAUMA GLORIENSIs Pittier, in Contrib. U.S. Nat. Herb. 13:94. 1910 Magnolia allenii Standl. in Field Mus. Publ. Bot. 22: 331. 1940. Tree up to 30 m. tall; branchlets glabrous. Leaf lamina ovate to ovate-elliptic or elliptic-oblong (occasionally obovate), cuneate to obtuse at the base, rounded to obtuse at the apex and sometimes slightly emarginate, up to about 31 cm. long and 16 cm. broad, coriaceous, glabrous; lateral nerves about 10-14 on each side of the midrib; petiole up about 5 cm. long, glabrous; stipules glabrous. Flower bud ovoid; bracts and peduncle glabrous. Tepals 9, 3-merous, greenish-white or creamy-white, glabrous; outer 3 oblong to oblong-elliptic, about 4-4.5 cm. long. Stamens about 13-15 mm. long. Gynoecium ovoid; carpels about 25-36, puberulous. Fruit ovoid; fruiting carpels up to about 4 cm. long, often becoming bifid at the apex. Mountains of Costa Rica and western Panama, at about 1000 m. altitude. According to Pittier this species serves as a shade tree in the coffee plantations at La Gloria in the Cartago Province of Costa Rica. cocLE: hills north of El Valle de Antón, alt. 1000 m., Allen 2200, 3574. The Panama plant was described by Standley as a new species of Magnolia, M. allenii; his description was inaccurate in its reference to free stipules, connate bracts and glabrous carpels. 2. TALAUMA SAMBUENSIS Pittier, in Contrib. U.S. Nat. Herb. 20: 105. 1918. Tree up to 40 m. tall; branchlets glabrous. Leaf lamina obovate- to elliptic- or ovate-oblong, cuneate to obtuse at the base, bluntly acuminate to subacute or apic- ulate at the apex, up to about 27 cm. long and 12.5 cm. broad, coriaceous or thinly coriaceous, glabrous; lateral nerves about 10-14 on each side of the midrib; petiole sometimes up to about 6 cm. long, glabrous; stipules glabrous. Flower-bud ovoid; bracts and peduncle glabrous. Tepals white. Fruit (according to Pittier) subglobose, about 8 cm. long and 7.5 cm. bro Endemic to southeastern Panama, at low altitudes up to 240 m. : Rio Cuasi, Chepigana District, alt. 240 m., Terry & Terry 1420; Boca de Paua- Ses on KI Sambi, alt. 20 m., Pittier 5681. The available material of this species is too scanty for its characters to be fully described and understood. Its leaves are somewhat differently shaped at the apex and more closely reticulate than in T. gloriensis, which occurs further to the west and at a higher altitude. (488) 1962] FLORA OF PANAMA (Winteraceae) 177 WINTERACEAE By LE DANDY Trees or shrubs with homoxylous wood. Leaves alternate or sometimes sub- opposite or verticillate, exstipulate, simple; lamina penninerved, entire. Flowers terminal or axillary, solitary or in fascicles or simple or compound umbels, actino- morphic, hermaphrodite or unisexual. Perianth heterochlamydeous; calyx either closed round the bud and at length splitting into 2-3 lobes, or open and not en- closing the bud; petals 2-numerous (sometimes reduced to | or 0), free, imbricate. Stamens numerous or few, free, in 2 or more series; filaments more or less elongated; anthers small, 2-thecous, extrorse, dehiscing by longitudinal slits, the thecae parallel or convergent. Gynoecium superior, sessile or stipitate; carpels numerous or few (sometimes 1), in 1 or more series, free or sometimes concrescent; ovules several or numerous. Fruit apocarpous of 1 or more baccate carpels, or sometimes a syncarpous berry. Seeds with copious endosperm and a minute embryo. An austral family of about 90 species, of which 4 are distributed in Central and South America, the remainder in the region of eastern Malaysia, the Solomon Islands, New Caledonia, Australia and New Zealand. The American species belong to the genus Drimys which is much more strongly represented in Malaysia and Australia. 1. DRIMYS J. R. & G. Forst. Drimys J. R. & G. Forst. Charact. Gen. Pl. 83. 1776. Wintera Murr. Syst. Veg. ed. 14, 507. 1784. Trees or shrubs, sometimes epiphytic. Leaves alternate. Flowers hermaphrodite or unisexual, solitary, fasciculate or umbellate. Calyx at first completely closed round the bud, at length splitting into 2-3 often caducous lobes; petals 2 or more, sometimes reduced to 1 or 0. Carpels 1 or more, in 1 series, free; ovules 2-seriate on the ventral suture. Fruit apocarpous of | or more baccate carpels. Species about 40, 4 in Central and South America, the others distributed in eastern Australia and in Malaysia from the Philippines and Borneo to New Guinea. The biggest concentration of species is in New Guinea. l. Drimys cranapensis L. f. Suppl. Pl. 269. 1781. Wintera granadensis (L. f.) Murr. Syst. Veg. ed. 14, 507. 1784. Shrub or small tree up to about 13 m. tall, glabrous. Leaf-lamina elliptic- oblong or narrowly elliptic-oblong to oblanceolate, cuneate or attenuate at the base, obtuse to rounded at the apex, up to about 17.5 cm. long and 5.5 cm. broad, coriaceous or subcoriaceous, glaucous or pale beneath; petiole up to about 2.5 cm. long. Flowers umbellate or fasciculate or sometimes solitary, hermaphrodite. Calyx 4.5-12 mm. long; petals 8-25, oblong to elliptic- or ovate-oblong, 5-25 mm. long. Stamens about 25-65, in 2-5 series. Carpels 3-24; ovules 7-12. (489) [Vor. 49 178 ANNALS OF THE MISSOURI BOTANICAL GARDEN Southern Mexico to western Venezuela and Peru. The species is divided by A. C. Smith into five geographical varieties, the Panama population being regarded as endemic and named D. granadensis var. chiriquiensis A. C. Smith, in Journ. Arnold Arb. 24: 25. 1943; this variety occurs in rain-forest at about 1800-2250 m. altitude. CHIRIQUÍ: Bajo Chorro, Boquete District, alt. 1800 m., Davidson 328. Figure 153. Drimys granadensis (490) 1962] FLORA OF PANAMA (Annonaceae ) 179 ANNONACEAE By R. E. FRIES Trees and shrubs with alternate and exstipulate leaves, always quite entire at the margins. Flowers generally trimerous. Sepals 3, valvate or imbricate. Petals generally 6, in two series, valvate or imbricate, free or rarely connate at the base, mostly conspicuously longer than the sepals; the inner petals often smaller, some- times rudimentary or even absent. Stamens generally numerous; filaments short; anther cells adnate, the connective broad, usually expanded in a more or less truncate or hoodlike disc above the anther; the anther cells sometimes transversely septate by thin, horizontal membranes (anthers locellate). Carpels numerous, rarely few or solitary, generally free; ovules 1 or more. Fruiting carpels free (mono- carps) or united in a dry or fleshy mass. Seeds with or without an aril, with copious ruminate endosperm; embryo minute. This large family, which is of pantropic distribution, is represented in America by 36 genera. Two of them (Annona and Xylopia) occur also in Africa, two also in tropical Asia (Artabotrys and Xylopia). Only 14 genera and 41 indigenous species are recorded at present from Panama but there is no doubt that further investigations will show that more are to be found in the country. Some species of Annona (in particular muricata L., squamosa L., reticulata L., and Cherimolia Mill.) are grown in the tropics for their edible fruits, and one or two of them are probably also cultivated in Panama. Only in respect to the last named, however, was any material from Panama available at the time when this was written. In the following, only the species known to grow wild in the Republic of Panama will be considered. The Annonaceae of the Panama flora have been dealt with, inter alia, by Standley in his “Flora of the Panama Canal Zone” (in Contr. U.S. Nat. Herb. 27: 1-415. 1928) and “The Flora of Barro Colorado Island, Panama” (in Contr. Arnold Arboretum 5: 1-178. 1933). The first of these papers is very rich in valuable in- formation on the native names of the species, for which readers are referred, as they are not quoted below. a. Anthers not transversely septate. b. Petals not caudate-cuspidate. c. Flowers axillary. d. Petals imbricate. e. Connective of the stamens not expanded above the anther. 2. NDRA. ee. Connective expanded into a truncate disk above the anther. Ovule attached close to the upper end of the eo. and Leiden: us, sometimes located lower a but no basal ; pedicel nearly always with a small bract above the artic Wee a ee E CR EMASTOSPERMA. Ovule basal, erect; pedicel without a bract er the articulation ...---...... ee 3. GuATTERIA. be joa] m [Vor. 49 180 ANNALS OF THE MISSOURI BOTANICAL GARDEN dd. Petals valvate. s not keeled on the inner surface; a ate usually few to several, rarely solitary and, i attac near the lower or upper end of the ovary; geese irregularly bursting 4. UNONOPSIS ee. All petals, or only the inner, keeled on the inner surface above the middle; ovules 2, erect, basal; monocarps open- ing on the inside by a longitudinal split .................... 6. ANAXAGOREA. cc, Flowers not axillary, more or less opposite the leaves. d. Petals imbricate e. Pubescence of simple hairs; monocarps stipitate, free....9. MALMEA. ee. Pubescence o Lite haírs or stellate scales; monocarps 1 sesle DER: Concrete ae nee 2. DUGUETIA. dd. e at Pass Kë outer, ien Monocarps free, stipitat 7. Desmopsıs. ee. Mice sessile, gege into a dry or fleshy mass. Outer petals without wings or spurs 13, ANNONA. ff. Outer petals produced into wings or spurs ............ 14. RoLLINIA. bb. Petals long alien ovules lateral 8. STENANONA. aa. Anthers transversely septate. b. Pedicels with bracts 5. XYLOPIA. bb. Pedicels ion d without bracts. c. Petals not saccate 10. PORCELIA. cc. Inner imi spi saccate, with involute margins ................ ll. CyMBOPETALUM. 1. CREMASTOSPERMA R. E. Fries CREMASTOSPERMA R. E Fries, in Acta Hort. Berg. 10:46. 1930; 11: 323. 1931. Shrubs or trees with 1, rarely 2, axillary flowers, or sometimes cauliflorous. Flowers medium-sized, generally glabrous; pedicels articulate above the base, nearly always with one small bract above the articulation. Sepals free, imbricate in bud, with thin, ciliolate margins. Petals much longer than the calyx, rigid, imbricate, with thin edges. Stamens numerous, the connective expanded above the anther into a disk. Carpels several; ovule 1, attached close to the upper end of the ovary and pendulous, or sometimes located lower down, even near the base and then erect. Monocarps free, stipitate, 1-seeded. Seventeen species are known, inhabiting the most tropical parts of South America, from Panama and Colombia to Peru, Bolivia, Amazonian Brazil and French Guiana. Only one species is known from Panama. CREMASTOSPERMA ANOMALUM R. E. Fries, Svensk. Vet.-Akad. Handl. ser. 3, 241°: 4, pl. 1 c-d. 1948 Tree up to 10 m.; young branchlets glabrous, reddish. Leaves on petioles 5-8 mm. long, quite glabrous and lustrous on both sides, elliptic or oblong and broadest at the middle, rotundate at the base, abruptly cuspidate at the apex, (10-) 15-28 cm. long and 5-10 cm. broad. Flowers solitary, white but blackening when dried; pedicels quite glabrous, 1-2 cm. long, articulate, without any bract above the artic- ulation. Sepals glabrous, connate into a disk 8-10 mm. broad with ciliolate mar- gin. Petals orbicular, 1.5 cm. diam., very shortly sericeous-tomentellous. Stamens (492) 1962] FLORA OF PANAMA (Annonaceae) EAM m " Figure 154. Cremastosperma anomalum adi] | WI NUOVO V DA D Hr YY 181 few, 2.0-2.5 mm. long. Carpels very numerous; ovule 1, attached near the base of the ovary, erect. Monocarps ellipsoid, 15-18 mm. long and 8 mm. across, black. Panama and Colombia. Allen 348 (493) DARIEN: vicinity of El Real, ca. 15 m. Allen 969; near the mouth of Rio Yape, ca. 20 m., - [Vor. 49 182 ANNALS OF THE MISSOURI BOTANICAL GARDEN 2. OXANDRA A. Rich. OxanDrA A. Rich. in Sagra, Cub. 10: 45. 1845; R. E. Fries, in Acta Hort. Berg. 10: 153. 1931 Trees or shrubs. Flowers small, solitary in the axils or in few-flowered axillary inflorescences; pedicels articulate a little above the base, bearing a few small, distichous bracts below and 1 above the articulation. Sepals 3, connate at the base, imbricate in bud, persistent. Petals 6, thin, black when dried, all rather alike, orbicular, ovate or oblong, imbricate. Receptacle hemispheric. Stamens 6-20, lanceolate-oblong, the connective elongated above the anther into a triangular to lanceolate appendage. Carpels rather few, (1-)4-13; ovaries cylindric-ovoid; stigma sessile, capitate or shortly clavate; ovule 1, basal, erect. Monocarps free, nearly sessile or shortly stipitate, l-seeded. Seeds without an aril. A rather large genus with 23 species in tropical America, distributed from the West Indies and Panama to southern Brazil; only 2 species are recorded from Panama. a. Leaves lanceolate, acute at the base ]. O. PANAMENSIS. aa. Leaves elliptic or oblong-elliptic, rounded at the base ......................... 2. O. LONIGIPETALA. 1. OXANDRA PANAMENSIS R. E. Fries, in Acta Hort. Berg. 10: 168, fig. 5, c, 1931. Tree up to 6 meters high; young branchlets covered with crisp hairs. Leaves on petioles 3-5 mm. long, finally glabrous on both sides, but densely verruculose beneath and thinly appressed-hairy on the midrib beneath, lanceolate, broadest at the middle and equally narrowed and acute at both ends, 8-12 cm. long and 2.0-3.5 cm. broad. (Flowers unknown). Fruits on very short rigid pedicels (2-3 mm.); monocarps glabrous, black when dried, ellipsoid, 12-15 mm. long and 9-10 mm. across; stipes 1-1.5 mm. long. DARIEN: Marraganti and vicinity, 10-200 ft., Williams 1017; Boca de Cupe, Williams 669. 2. OXANDRA LONGIPETALA R. E. Fries, in Acta Hort. Berg. 10: 170, pl. 5. 1931. A small tree; young branchlets hirsute-tomentose. Petioles very short (about l mm. long). Leaves glabrous on the upper side, the midrib tomentellous when young, glabrous beneath and sparsely appressed-hairy on the midrib, elliptic or oblong-elliptic, rotundate and even subcordate at the base, rather long-acuminate at the obtuse apex, 7-10 cm. long and 2.5-3.5 cm. broad. Flowers solitary; pedicels 9-6 mm. long, appressed-hairy; flower buds oblong. Sepals rotundate-ovate. Petals linear-oblong, rotundate at the apex, up to 16 mm. long. Carpels about 11. (Fruit unknown). ZONE: near Salamanca Hydrographic Station in the gorge of the R. Pequení, 70- 80 m m., hee Steyermark & Allen 16968; Lion Hill Station, Hayes, 385. (494) 1962] FLORA OF PANAMA (Annonaceae) 183 A N A E W Figure 155. Oxandra longipetala (495) [Vor. 49 184 ANNALS OF THE MISSOURI BOTANICAL GARDEN 3. GUATTERIA Ruiz & Pav. GUATTERIA Ruiz & Pav. Fl. Peruv. & Chil. Prodr. 85, tab. 17. 1794; R. E. Fries, in Acta Hort. Berg. 12: 291. 193 Shrubs or trees, rarely climbing; the pubescence of simple hairs. Flowers axillary, solitary or in few-flowered inflorescences; pedicels articulate, with a few small bracts (rarely foliaceous) below the articulation, no bracts above this. Sepals valvate. Petals imbricate, usually hairy, especially so outside at the base. Stamens numerous; connective expanded above the anther into a truncate disc; anther cells not locellate. Carpels numerous; ovule 1, basal, erect. Monocarps ovoid or ellipsoid, or shortly cylindrical-ellipsoid, l-seeded, stipitate or rarely almost sessile. Seeds without an aril The genus is the largest in the family Annonaceae, the species numbering about 240. It is distributed from southern Mexico and the West Indies to Bolivia and southern Brazil, centering in the Amazonian region and Guiana. a. Leaves not verrucose. b. Pubescence appressed. Leaves -5 cm. Se LC LUCENS cc. Leaves nn than 6 cm d. gi vine! and a little emarginate at the base ........ 2. G. AMPLIFOLIA dd. pee acute at the base. Petals oblong, all of about the same length .................. 3. G. INUNCTA ee. Petals broadly ovate to ic die Meis inner shorter Petioles 1.0-1.5 cm. long, the upper half narrowly winged; 2 nerves nado S ae upper s = of e leave 4. G. SLATERI IE Páloles 3. 7 cm. long; lateral nerves € on the upper side ........ 5. G. PANAMENSIS bb. Pubescence spreading. c. Leaves oblanceolate to obovate; flower pedicels 8-10 mm. long. ....6. G. ALLENII ce, Leaves linear-oblong; flower pedicels about 20 mm. long ....7. G. CHIRIQUIENSIS p EI Leaves very verrucose on both sides b. Branchlets, pedicels and der oe of the leaves with spreading pubes- m. lon cence; leaf-petioles 1-2 m 8. G. AERUGINOSA bb. Branchlets, pedicels and under a of the leaves with ee pubes- cence; leaf-petioles 6-8 mm. lon 9, G. DUMETORUM l. GUATTERIA LUCENS Standl. in Trop. Woods 42: 22. 1935; R. E. Fries, in Acta Hort. Berg. 12: 481, pl. 34. 1939. A small tree, the young branchlets sparsely sericeous, soon glabrous and blackening. Leaves with petioles 6-8 mm. long, lustrous on both sides, finally glabrous above except on the more or less hirsute midrib, sparsely appressed-hairy beneath, oblong-lanceolate, acute and decurrent at the base, tapering at the apex into an acumen 1-2 cm. long, 10-16 cm. long and 3.5-4.5 cm. broad. Flowers 1 or 2 in the axils of persistent or newly fallen leaves; pedicels thin, rigid, black, sparsely sericeous, glabrescent, 10-13 mm. long, articulate 3-4 mm. above the base. Sepals rounded-ovate, about 2.5 mm. long. Petals green but blackening when dried, puberulous, sericeous outside at the base, oblong-spathulate, rounded at the apex, (496) 1962] FLORA OF PANAMA (Annonaceae) 185 Figure 156. Guatteria amplifolia the outer up to 12 mm. long and 4 mm. broad, the inner a little larger. Stamens 1 mm. long, connective disc very shortly setulose. (Fruits unknown). SAN BLAS: Permé, Cooper 280. 2. GUATTERIA AMPLIFOLIA Tr. & Pl. in Ann. Sc. Nat., ser. 4. 17: 35. 1862; R. E. Fries, in Acta Hort. Berg. 12: 376, fig. 12, a. 1939. Branchlets glabrous. Leaf petioles 4-5 mm. long, rugose, glabrous; blades papyraceous, glabrous above, sparsely provided beneath with very short appressed hairs, elliptic or oblong-elliptic or ovate, rotundate and a little emarginate at the very base, more or less abruptly cuspidate, 20-40 cm. long and 7-12 cm. broad; lateral nerves 18-25 on each side, more or less impressed on the upper side. Flowers solitary or sometimes 2; pedicels sericeous, 1-1.5 cm. long. Sepals triangular-ovate, reflexed, sericeous outside, tomentellous inside, 6-8 mm. long. Petals ferruginous- tomentellous on both sides, oblong-lanceolate, finally oblong-obovate, rotundate at the apex, 15-18 mm. long, 8-12 mm. broad. Stamens 1.8 mm. long. Monocarps ellipsoidic-pyriform, rugose, apiculate, 8-9 mm. long and 5 mm. across; stipes 10-15 mm. long, rigid. (497) [Vor. 49 186 ANNALS OF THE MISSOURI BOTANICAL GARDEN CANAL ZONE: imc Fendler 3; Lion Hill Station, Hayes 346; shores of Gatun Lake, Bangham 464; Barro Colorado Island: Barbour Point, Shattuck 406, and Chapman Trail, Starry 170, and Ce katie Trail, Starry 325. 3. GUATTERIA INUNCTA R. E. Fries, in Acta Hort. Berg. 12: 378. 1939. Guatteria Ouregou Griseb. Fl. Brit. W. Ind. 7.1860, pro parte quoad specimen a St. 'Thomas citatum, non Dunal. A shrub or tree up to 8-10 m. high, young branchlets at first shortly ferrugi- nous-sericeous. Leaves on petioles 5-8 mm. long, chartaceous, pale green and soon quite glabrous above, under side olivaceous and densely appressed-hirsute at first, finally nearly glabrous, elliptic to oblong-elliptic, broadest at the middle, acute and a little decurrent at the base, rather gradually tapering to the blunt apex, 15-28 cm. long and 6-12 cm. broad. Flowers 1-2; pedicels very thin but rigid, 1.5-2.0 cm. long. Sepals rotundate-ovate, ferruginous-sericeous outside, glabrous inside, 4-5 mm. long. Petals oblong and rotundate-truncate at the apex, ferruginous-sericeous outside, 10-15 mm. long, 5-8 mm. broad. Stamens 1.3-1.5 mm. long. Monocarps ellipsoidic, very shortly apiculate at the apex, acute at the base, about 7 mm. long and 4 mm. across; stipes thin, about 7 mm. long Costa Rica and Panama. TORO: vicinity of Chiriquí Lagoon, Water Valley, Von Wedel 1460, 1716; Old Bank bland, Isla Colon, alt. 0-120 m., Von Wedel 1932. GUATTERIA INUNCTA, var. CAUDATA R. E. Fries, in Ann. Missouri Bot. Gard. 42: 152. Differs from the type in the leaves, relatively narrower, narrowly cuneate at the base, with a narrowly caudiform apical acumen 2.0-3.5 cm. long, and in the more slender pedicels of the flowers. BOCAS DEL TORO: vicinity of Chiriqui Lagoon, Old Bank Island, Von Wedel 2108. 4. GUATTERIA SLATERI Standl. in Field Mus. Publ. Bot. 4: 206. 1929; R. E. Fries, in Acta Hort. Berg. 12: 517. 1939 Tree up to 10 m. high, the young branchlets ferruginous-sericeous. Leaf petioles 1.0-1.5 cm. long, the upper part narrowly winged; blades blacken when drying, glabrous on upper side from the first, with sparse short appressed hairs beneath (longer on the midrib) and more or less glabrescent, elliptic to obovate, acute and long-decurrent at the base, rather abruptly contracted into an obtuse cusp 1-2 cm. long, 15-20 cm. long and 5-8 cm. broad. Flowers solitary, pedicels 1.5-2.0 cm. long, rigid. Sepals rotundate-ovate, about 4 mm. long and broad, soon recurvate, deciduous, sericeous outside, glabrous inside. Petals broadly ovate, rotundate at the apex, 8-12 mm. long and broad, golden yellow-sericeous outside, tomentellous inside, the inner a little smaller. Stamens 1.2-2.0 mm. lon HIRIQUI: Progreso, Cooper 2 Slater, 177; Bajo Chorro, Boquete District, in rain forest, E 6000 Gs Davidson 233; Boquete Region, Cerro Horqueta, cloud forest, alt. 6500 m., n Hagen & Von Hagen 2135 BOCAS DEL TORO: Robalo Trail, northern slopes of Cerro Se ak 6000-7000 ft., Allen 4802. (498) 1962] FLORA OF PANAMA (Annonaceae) 187 5. GUATTERIA PANAMENSIS R. E. Fries, in Arkiv Bot. andra ser. 1: 335. 1950. Guatteria costaricensis subsp. panamensis R. E. Fries, in Acta Hort. Berg. 12: 515. 9 A tree 10-15 m. high; branchlets ferruginous-sericeous. Leaves on petioles 3-7 mm. long, membranaceous, more or less blackening when dried, smooth above and hirsute at first at the midrib but soon quite glabrous, with appressed rigid hairs beneath, finally presumably glabrous, oblong or oblong-obovate, broadest at the middle or a little above the middle, acute at the base, abruptly contracted into a broad obtuse cusp 1 cm. long, 20-28 cm. long and 6-9 cm. broad. Flowers solitary or 2 in the axils of persistent or fallen leaves; pedicels sericeous, about 3 cm. long. Petals finally horizontally expanded, ovate and rotundate at the apex, the outer densely sericeous outside, tomentellous on the upper half inside, about 1 cm. long, the inner tomentellous outside and glabrous inside, a little smaller than the outer. Monocarps very numerous, ellipsoid to obovoid, shortly acute, 9-10 mm. long and 5.0-5.5 mm. across, abruptly tapering into a thin stipe 5-7 mm. long. Known only from Panama. BOCAS DEL TORO: region of Almirante, Daytonia Farm, Cooper 382; Old Bank Island vicinity of Chiriquí Lagoon, Von Wedel 1965. 6. GUATTERIA ALLENI R. E. Fries, in Arkiv Bot. andra ser. 1: 336. 1950. A tree about 12 m. high; young branchlets very densely covered with rigid ferruginous erect-patent hairs. Leaves on petioles 2-3 mm. long, membranaceous, glabrous above except on the more or less tomentose midrib, pale green beneath and rather thinly clothed with decumbent or patent hairs, more densely along the thicker nerves, oblanceolate-obovate, tapering into a cuneate acute base, abruptly contracted at the apex into an acumen 0.5-1.0 cm. long, 10- 18 cm. long and 3-7 cm. broad. Flowers solitary in the axils; pedicels thick, 8-10 mm. long, articulate about 2 mm. above the base. Sepals rotundate, about 4 mm. long and broad. Petals ob- long, tapering towards the blunt apex, densely golden-sericeous on the outside, especially near the base, 10-15 mm. long and 5-6 mm. broad. Stamens with con- nectives 1.5 mm. long, disk velutinous. (Fruit unknown). COCLE: north rim of El Valle, Allen 1900. 7. GUATTERIA CHIRIQUIENSIS R. E. Fries, in Svensk. Vet-Akad. Handl. ser. 3. 241°: 11. 1948 Young branchlets shortly ferruginous-hirsute. Leaf petioles 1-3 mm. long; blades glabrous above except on the densely tomentose midrib, underside rather sparsely clothed with patent ferruginous persistent hairs, linear-oblong, acute and shortly decurrent at the base, rather abruptly contracted into an obtuse cusp about 1.5 cm. long. Flowers solitary (or more?), their pedicels about 2 cm. long, rigid, articulate about 5 mm. above the base, the lower part sericeous, the upper provided with more patent ferruginous hairs. Sepals broadly ovate, acute, ferruginous- sericeous outside, glabrous inside, about 5 mm. long. Petals thick, horizontally expanded, elliptic-oblong and rotundate at the apex, densely golden-sericeous (499) [Vor. 49 188 ANNALS OF THE MISSOURI BOTANICAL GARDEN outside, subglaucous inside, pulverulent upwards on both sides. Stamens 1.7-2.0 mm. long. Monocarps black, narrowly ellipsoidic-clavate, 9-10 mm. long and 4-5 mm. broad on stipes 6-7 mm. long. cHIRIQUÍ: vicinity of San Félix, alt. 0-120 m., Pittier 5132, 5748. 8. GUATTERIA AERUGINOSA Standl. in Field Mus. Publ. Bot. 4: 206. 1929. Tree about 15 m. high; young branchlets densely clothed with short fer- ruginous hairs. Leaves on very short petioles (1-2 mm. long), 14-18 cm. long and 4-5.5 cm. broad, oblong to oblanceolate and broadest above the middle, shortly narrowed to the base, long-cuspidate at the apex, verruculose-punctate on both sides, ferruginous-pilose on the midrib above and finally glabrous, densely rubi- ginous-pilose beneath, later thinly ferruginous-hirsute, the hairs rather long-patent and soft. Flowers solitary; pedicels thick, rigid, densely pilose, 1-3 cm. long, ar- ticulate 3-4 mm. above the base, Petals thick, finally horizontally expanded, seri- ceous-tomentellous on the outside, tomentellous on the inside except at their glabrous base, ovate to oblong-elliptic, the outer petals up to 13 mm. long and 6-8 mm. broad, the inner a little shorter. Stamens 1.5-1.7 mm. long. Monocarps black, narrowly ellipsoid, acute at both ends, 10 mm. long and 4 mm. across. : Von Wedel 155, 490; Isla Colón vicinity of Chiriquí Lagoon, Von Wedel 285 2856: na near Almirante, Cooper 526, 526a. GUATTERIA DUMETORUM R. E. Fries, in Svensk. Vet.-Akad. Handl. ser. 3, 241°: 12. 1948 Young branchlets appressed-hairy, glabrescent. Leaf petioles 6-8 mm. long; blades densely verrucose on both sides, hirsute at first above but soon quite glabrous, underside covered at first with long more or less ferruginous appressed hairs, finally nearly glabrescent but still with some hairs especially on the midrib, oblanceolate, gradually tapering and decurrent at the base, rather abruptly contracted at the apex into an obtuse acumen 1-1.5 cm. long, about 15 cm. long and 4-5 cm. broad. Flowers 1-2 in the axils of leaves; pedicels thin, sparsely sericeous, about 2 cm. long, articulate about 0.5 cm. above the base. Sepals finally recurvate, ovate, acute, densely sericeous outside, glabrous inside, 3-4 mm. long. Petals oblong to rhombic- ovate, narrowed to the obtuse apex, 10 mm. long and 4-5 mm. broad, flavescent- tomentellous outside; outer petals pubescent at the base, inner petals with longer appressed golden-yellow hairs nearly up to the apex. Stamens 1 mm. long, the connective disk plane, glabrous. (Fruit not known). COLÓN: along Rio Paté, in forests and thickets, alt. 10-100 m., Pittier 3915. 4. UNONOPSIS R. E. Fr. Unonopsis R. E. Fries, in Svensk. Vet.-Akad. Handl. 345: 26. 1900. Shrubs or small to medium-sized trees. Inflorescences axillary or sometimes produced from older branches or from the trunk; pedicels articulate above the basal bract and provided with a second bract above the articulation. Flowers small, the (500) 1962] FLORA OF PANAMA (Annonaceae) 189 buds spherical. Sepals minute, valvate. Petals rigid, valvate, ovate or rotund, con- cave, the inner thick in particular; pollen sacs not locellate; connective expanded above the anther into a truncate disk. Carpels several to numerous; ovules (up to 6-8), lateral, or else solitary and attached near the lower or upper end of the ovary. Monocarps free, stipitate. A large genus of about 30 species, inhabiting a vast area from British Honduras and the West Indies to south Brazil; only 2 species recorded from Panama. a. Leaves 2 to obovate-elliptic, 20-30 cm. long; pedicels thin, 2-4 cm. rend ovules solitary . PITTIERI. aa. Leaves oblong-oblanceolate, 15-20 cm. long; pedicels short and E m m. long—or more?); ovules 3-4, Jateral . PANAMENSIS l. Unonopsis PITTIERI Saff. in Journ. Washington Acad. Sci. 15: 102. 1925. Trees of moderate size, 10 m. tall, with pyramidal crown; young branches minutely sericeous, becoming glabrous and blackish, densely lenticellate, the stouter with grayish bark. Leaf petioles at first puberulent, becoming glabrous and blackish, 3-5 mm. long, flat or channelled above; blades membranaceous-papyraceous, al- most concolorous, at first sericeous on both sides but soon glabrescent, beneath with some more or less persistent hairs along the midrib or completely glabrous, the whole surface finely verruculose-punctate, elliptic or obovate-elliptic, narrowed toward the base and broadly acute or obtusish, cuspidate at the tip, 20-30 cm. long and 5-9 cm. broad, the veins on both sides prominent, 12-15, arcuate, ir- regularly gathered toward the margin. Inflorescences on leafless branches, about l cm. thick; pedicels thin, minutely sericeous, 2-4 mm. long, articulate about mid- way and bearing a minute bracteole. Sepals 1.5 mm. long, sericeous without, tri- angular and strongly connate. Petals valvate, broadly ovate, acute, the outer rather thick, glaucous-sericeous without, about 9 mm. long and 7 mm. broad, the inner thicker, 5-6 mm. long and 4-5 mm. broad, the midrib sericeous without but other- wise glabrous. Stamens 1.5 mm. long. Pistils 14-18, the ovaries cylindric-prismic, glabrous save the hairy angles, 1.5-1.7 mm. long, the stigmas conic-ovoid, ; long, the ovules solitary, attached below midway, horizontal. Monocarps brown, globose, 11-12, glabrous, with a stipe about 1 cm. ONE: d of Rio Chinilla, above Nuevo Limón, Max alon Sieg Rio Fató, in forests or thickets, alt. 10-100 m., Pittier 3871 als reported by Baraily from "Gei Colore Island. This species occurs also in Costa Rica, from whence the very closely related U. schippii is described. 2. Unonopsis panamensis R. E. Fries, in Ann. Missouri Bot. Gard. 42: 151. 1955. Young branches with short ferruginous erect or appressed hairs. Leaf petioles terete or flattened above, tomentellous, 4-5 mm. long; blade chartaceous, above glabrous save for the minutely hairy glabrescent midrib, beneath hirsute-glabres- cent along the midrib, oblong-oblanceolate, broadest above the middle, shortly acute at the base, abruptly contracted at the tip with a cusp 1.0-1.5 cm. long, 15-20 cm. long and 4-5 cm. broad, the midrib elevated on both sides, the stronger veins (501) [Vor. 49 ANNALS OF THE MISSOURI BOTANICAL GARDEN 190 Figure 157. Unonopsis panamensis (502) 1962] FLORA OF PANAMA (Annonaceae) 191 about 15 on both sides, arcuately ascending and converging about 1-2 mm. from the margin, slender and prominent above. Inflorescences springing from the axils of fallen leaves; pedicels thick, ferruginous-sericeous, 7 mm. long or longer, articulated slightly above the base. Sepals free, rotund-ovate, acutish, about 2 mm. long. Petals stiff, blackening in desiccation, the outer rotund, about 7 mm. in diameter and ferruginous-sericeous without, the inner smaller, ferruginous- tomentose along the midrib. Stamens 1.8-2.0 mm. long, truncate at the tip. Ovaries 1.5 mm. long, prismatic and shortly strigose, tipped with a globose-ovoid stigma; ovules 3-4, lateral. (Fruit unknown.) Known only from Panama. PANAMA: Cerro Campana, trail from Campana to Chica, alt. 600-800 m., Allen 2647. 5. XYLOPIA E XvLoPi L. Syst. ed. 10. 1759; R. E. Fries, in Acta Hort. Berg. 10: 85. 1930. Shrubs or trees. Flowers usually small, sessile or shortly pedicellate, in axillary inflorescences or sometimes produced from old branches or from the trunk. Sepals small, valvate in bud, nearly free or connate into a cupular calyx. Petals thick, valvate, linear-oblong or sometimes deltoid. Stamens numerous, the anther cells locellate; connective expanded above the anther. Ovaries hidden in a deep cavity in the center of the torus; ovules lateral. Monocarps free, cylindric or sometimes short, clavate, containing a row of rather few seeds. About 50 species in America, distributed from Central America and the West Indies to southern Brazil and Paraguay; also in the tropics of the Old World. Only 4 species are known at present from Panama. a. Sepals nearly free or connate up to the middle; carpels few (4-8). ers numerous in axillary inflorescences; flower buds elongate, cylindric- trigonous; petals linear-oblong ..............— l. X. FRUTESCENS. bb. Flowers solitary; flower buds ovoid; petals flat, the outer ovate.....2. X. BOCATORENA. aa. Calyx cup-shaped with almost wholly connate sepals; carpels numerous (25-45). axill inflorescences; flower buds narrowly broa 3, X. AROMATICA Flowers numerous SE CRANTHA. 1. XyLopra FRUTESCENS Aubl. Pl. Guy. Fr. 1:602, t. 292. 1775, excl. syn. Linn. et Brownei. Xylopia setosa Poir. in Lam. Encycl. Méthod. 18: 812. 1808. Xylopia muricata Vell. Fl. Flumin. 9: t. 127. 1827; non Linn. A medium-sized tree or shrub; young branchlets densely covered with about 2 mm. long, more or less ferruginous, erect-patent hairs. Leaves rigid, glabrous on upper side, densely covered underneath with long appressed hairs, lanceolate, acute at the base, rather long-acuminate, 4-6 cm. long and 0.8-1.5 cm. broad; petioles 2-4 mm. long. Inflorescences 1-5-flowered, axillary. Sepals ovate, about 2 mm. long, (503) [Vor. 49 192 ANNALS OF THE MISSOURI BOTANICAL GARDEN Figure 158. Xylopia frutescens shortly connate at the base. Outer petals 8-11 mm. long and 2.5 mm. broad, oblong- obtuse, sericeous; inner petals a little shorter and narrower. Carpels 6 (or more ?); style with stigma 4 mm. long, glabrous, curvate and thickened above the base. Monocarps 10-13 mm. long and 9-10 mm. broad, rhomboid-globose, obtuse, con- tracted at the base into a stipe 1-2 mm. long. Seeds 2 (or 1), ovoid, black, 6-7 mm. ong. Distributed from Central America (Guatemala and British Honduras) to southern Brazil (Rio de Janeiro and Minas Geraés). CANAL ZONE: low hill north of Balboa, Gillespie P 33; Barro Colorado Island, Shattuck H 250. PANAMÁ: Sabana, Pittier 2539 Panamá, Haenke 167, Seeman s.n.; vicinity of rraijan, Allen 1765; Isthmus of Panama, in woods, Hayes 670; Punta Paitilla, Piper 5410; vicinity of Juan Franco Race Track, Standley 27760. 2. XYLOPIA BOCATORENA Schery, in Ann. Missouri Bot. Garden 30:86. 1943. Tree, its young branchlets very shortly and densely tomentellous. Leaves rigid, on petioles 2-3 mm. long, elliptic-lanceolate, acute at the base, long acuminate at the apex, glabrous above, thinly provided beneath with short appressed hairs, 8-11 (504) 1962] FLORA OF PANAMA (Annonaceae ) 193 cm. long, and 2.3-3.5 cm. broad. Flowers solitary; pedicels sericeous, often recurved, 5-6 mm. long; flower buds ovoid, about 1 cm. long. Sepals connate up to the middle. Petals flat; the outer ovate, sericeous outside, about 12 mm. long and 6 mm. broad; the inner rhombic-lanceolate, 11 mm. long and 4-5 mm. broad. Stamens 3 mm. long, connective appendage ellipsoid. Carpels about 8. Monocarps 1-4 (or more), obovoid to clavate, ca. 2 cm. long. Seeds 2, black. BOCAS DEL TORO: Isla Colön, vicinity of Chiriqui Lagoon. Von Wedel 2965. 3. XYLOPIA AROMATICA (Lam.) Eichl. in Mart. Fl. Bras. 131: 43. 1841; R. E. Fries, in Acta Hort. Berg. 10: 106. 1930 (where 23 synonyms are quoted). A tree; young branchlets tomentose. Leaves large, 8-15 cm. long and 2.0-4.5 cm. broad, rigid, lanceolate or elliptic-lanceolate, gradually narrowing at the apex, rounded or shortly acute at the base, glabrous on the upper side from the first except on the midrib, covered underneath with short patent or appressed hairs or even glabrous, glaucous; petioles 3-6 mm. long. Flowers numerous in axillary inflo- rescences; flower buds long and narrow, trigonous, densely sericeous. Calyx cup- shaped, with almost wholly connate sepals. Outer petals linear, up to 30 mm. long and 2-3 mm. broad; inner petals a little shorter, about 1 mm. broad. Stamens 1.0-1.5 mm. long. Carpels very numerous (about 25). Monocarps cylindric, toru- lose, 2.0-2.5 cm. long and 6-7 mm. thick. Seeds about 6, ovoid, black, 5-6 mm. long. This is the most widely distributed species of all American Xylopias, covering a large area from Central America and the West Indies down to southern Brazil and Paraguay. PANAMA: sabanas near Chepo, 30 m., Hunter & Allen 49; Rio Tataré, Woodson & Schery 1018; Isla Taboga, ca. 0-186 m, Woodson, Allen & Seibert 1447; Bohio, Pittier 3417; savanas north of Panama city, Brother Paul 443; Island San José in sinu Panamensi, Andersson s.n. 4. XYLOPIA MACRANTHA Tr. & Pl. in Ann. Sc. Nat., sér. 4. 17: 38. 1862. R. E. Fries, in Acta Hort. Berg. 10: 112, fig. 8. 1939. Young branchlets sericeous-villous. Leaves 3-4 mm. long, petiolate, coriaceous, glabrous and lustrous above, covered beneath with long appressed hairs, finally glabrescent, oblong, rounded at the base, gradually narrowed into an obtuse apex, 9-12 em. long and 3-4 cm. broad. Flowers solitary in the axils of fallen leaves; pedicels thick, 5-6 mm. long. Sepals nearly wholly connate, forming a cup 8-10 mm. high and 10-11 mm. broad, ferruginous-sericeous on the outside. Petals thick, rigid, ligneous, 20-23 mm. long, the outer plane, oblong, sericeous on the outside, grey-tomentellous on the inner side, 8 mm. broad; inner petals quadrangular-pris- matic, very acute, about 3 mm. in diameter, broadened at the base. Stamens 1.5-2 mm. long. Carpels numerous (about 45). Fruit unknown. At first described from Colombia, but later found in many localities in Panama and Venezuela. (505) [Vor. 49 194 ANNALS OF THE MISSOURI BOTANICAL GARDEN AL ZONE: along Capo Quebrado, Pittier 6822; Barro Colorado Island in Gatün Lake, e 120 m. or less, Standley 41085; hills north of Frijoles, Standley 27483; hills west of e Canal, near Gatün, Standley 27211; Barbour Point, Bangham 490; end of Van Tyne Trail Barro Colorado Island, Zetek 3903. corów: vicinity of Camp Pina, Allen 3676. 6. ANAXAGOREA St. Hil. ANAXAGOREA St. Hil. in Bull. Soc. Philom. Paris 91. 1825; R. E. Fries, in Acta Hort. Berg. 12:6. 1934 Small trees or shrubs. Flowers small, axillary, sometimes produced on the older, leafless branches; pedicels bracteolate. Sepals thin, valvate in bud, free or sometimes connate at first and irregularly splitting, often deciduous. Petals valvate, all rather alike, but the inner mostly a little smaller and thicker, all, or only the inner, keeled on the inner surface above the middle. Stamens several, the innermost often sterile; connective of the fertile stamens not forming a disk above the pollen sacs, elongated into a ligule, sometimes very short. Carpels several; ovules 2, erect, produced from the basal part of ovary cavity. Monocarps free, dry, stipitate, club- shaped, opening along the inner sides. Seeds 2, very smooth and lustrous, appressed against each other and therefore plane on one side side and convex on the other. A rather large genus, containing 20 American species occupying a vast region from Central America and Colombia to southern Brazil, centering along the Amazon. a. Leaves u. (8-16 cm. nr acute and long-decurrent at the ar? the secondary nerves 7-8 per s PANAMENSIS. aa See Ge Mee cm. Mery rounded at the base; the ud gu about 15 per si 2. A. ALLENI: L ANAXAGOREA PANAMENSIS Standl. in Journ. Washington Acad. Sc. 15: 101. 1925; R. E. Fries, in Acta Hort. Berg. 12: 24. 1934. A shrub up to 2 m. high; the young branchlets minutely ferruginous-puberulous, soon glabrescent. Leaves on petioles 4-7 mm. long, membranaceous, paler and puberulous beneath, finally glabrous on both sides, lanceolate-oblong, acute and de- current at the base, rather gradually tapering at the apex, 8-16 cm. long and 3.0-4.5 cm. broad; secondary nerves 7-10 per side. Flowers solitary; pedicels 2.0-2.5 cm. long, finally glabrous, provided near the calyx with an orbicular, amplectant bract, L mm. long. Sepals membranaceous, ovate, acuminate, ferruginous-puberulous, 8 mm. long. Outer petals linear-oblong, obtuse, 2 cm. long; inner petals triangular- ovate, acuminate, ca. 1 cm. long. Monocarps glabrous but rugose, apiculate at the apex, 2.2-2.5 cm. long, together with the stipe. ANAMA: in wet forest along the Rio Tapia, near sea-level, Standley 26168.—According to Standley also collected on the hills north of Frijoles, Canal Zone: Standley 27589. 2. ANAXAGOREA ALLENII R. E. Fries, in Ann. Missouri Bot. Gard. 42: 151. 1955. Tree about 6 m. tall; young branches very shortly rufous-tomentellous. Petiole of leaves 10-12 mm. long, thick; blade chartaceous in desiccation, yellowish green, very glabrous and smooth above, glabrous and paler beneath, oblong-lanceolate, rounded at the base, gradually narrowed toward the tip, 30-35 cm. long and 8-10 cm. broad, (506) 1962] FLORA OF PANAMA (Annonaceae) 195 Figure 159. Anaxagorea allenii the costa flattened above, strongly prominent beneath, terete, roughish, lateral veins about 15 on each side, broadly ascending, both the veins and the veinlets somewhat elevated beneath. Pedicels of flowers short, 3-4 mm. long, thick, rufous-tomentellous like the sepals and the petals, provided with a vaginiform bract near the calyx. Sepals ovate-orbicular, rounded at the tip, not recurved, 7-10 mm. long. Petals thick, the outer oblong, rounded and not narrowed toward the tip, but carinate toward the tip within, up to 12 mm. long and 5 mm. broad, the inner lanceolate, very acute, pilose to the middle and carinate above the middle within. Stamens linear, 3 mm. long, the appendage of the connective slender, about 0.7 mm. long, truncate-rounded. Fruit unknown. CANAL ZONE: Quebrada Löpez, alt. 30 m., Allen 2143. (507) [Vor. 49 196 ANNALS OF THE MISSOURI BOTANICAL GARDEN Another species of the genus Anaxagorea certainly also occurs in Panamä, as the material available to me from the herbarium of the Missouri Botanical Garden includes a third, collected in the vicinity of the Chiriqui Lagoon (Von Wedel 2210). This lacks flowers, however, and it is therefore impossible to determine with any certainty its position in relation to hitherto known species of Anaxagorea. 7. DESMOPSIS Saff. Desmopsis Saff. in Bull. Torrey Bot. Club 43: 184. 1916; R. E. Fries, in Acta Hort. Berg. 10: 16. 1930 Shrubs or small to medium-sized trees; the pubescence of simple hairs. Flowers medium-sized in few-flowered inflorescences (rhipidia) opposite the leaves or rarely produced from the trunk; pedicels usually long with 2 bracts. Sepals valvate. Petals valvate in bud or a little imbricate at the apex, all rather alike, linear-oblong to lanceolate. Stamens numerous; pollen sacs not locellate; connective expanded above the anther into a truncate disk. Carpels several to numerous with 1-8 lateral ovules. Monocarps stipitate or rarely subsessile, spherical to shortly ovoid-cylindric. About 13 species, distributed from Mexico down to Panama, one (neglecta) endemic on Cuba a. kon orescence potancie i near the apex a foliaceous bract 1-2 cm. long; ower pedicels m. lon b. Petals => at SE tips; monocarps 10-12 cm. across ...............- 1 D. PANAMENSIS bb. Petals not incurved at the tips; monocarps 6-8 mm. across. c. Monocarps with thin stipes 5-8 mm. long. d Leaves acutely acuminate at the apex 2. D. MICROCARPA dd. Leaves obtuse or rotundate at the apex 3. D. BIBRACTEATA. cc. Monocarps with thick stipes 1-3 mm. long 4. D. MAXONII aa. All bracts very small; flower pedicels about 1 cm. long 5. D. BREVIPES. l. Desmopsis PANAMENSIS (Rob.) Saff. in Bull. Torrey Bot. Club 43: 185, pl. 7. 1916; R. E. Fries, in Acta Hort. Berg. 10: 18, fig. 3, b. 1930 Unona panamensis Rob. in Amer. Journ. Sci. 50: 175. 1895. Tree 5-8 m. high, young branchlets ferruginous-tomentose. Leaf petioles tomen- tose, 3-4 mm. long; blades glabrous on the upper side except on the midrib when young, subglabrous underneath with the midrib and thicker nerves tomentellous, elliptic or elliptic-lanceolate, cuneate and acute at the base, obtusely acuminate at the apex, 10-20 cm. long and 4-8 cm. broad. Inflorescences opposite the leaves, l- to 2-flowered; their peduncle 3-10 mm. long, provided near the apex with a cordate-orbicular, acute, foliaceous bract 1.5-2.0 cm. long; flower pedicel about 7 cm. long, tomentellous, generally curved, bearing below the middle and at the base on the secondary flower small bracts 1-3 mm. long. Sepals triangular, 2-3 mm. long. Petals linear-oblong from a broad base, revolute at the margins and incurved at the apex, 15-25 mm. long. Ovaries 7-15, ovules 2-8. Monocarps shortly cylindric, rounded at both ends, 14-18 mm. long and 10-11 mm. across; stipes 8-10 mm. long. (508) 1962] FLORA OF PANAMA (Annonaceae) 197 Figure 160. Desmopsis microcarpa CANAL ZONE: woods near Gattn Station on the old Panama Railway, Hayes s.n.; hills around the Agua Clara Reservoir, near Gatun, alt. 20-30 m., Pittier 2660. Reported also from Barro Colorado Island (Standley, 1933). 2. Desmopsis MICROCARPA R. E. Fries, in Acta Hort. Berg. 10: 23. 1930. Desmopsis glabrata Schery, in Ann. Missouri Bot. Gard. 28: 428. 1941. Small tree 2-3 m. high, with the young branchlets thinly appressed-hirsute, very soon glabrescent. Leaves on petioles 3-4 mm. long, lanceolate-elliptic to broadly elliptic, rounded to cuneate at the base, rotundate and abruptly acuminate and more or less gradually tapering at the apex, glabrous above except on the more or less hirsute midrib, glabrous beneath, 10-20 cm. long and 4-10 cm. broad. Inflo- rescences up to 5-flowered; their peduncles about 1 cm. long and bearing near the apex a rotundate-cordate or foliaceous bract 1-2 cm. long; flower pedicels 5-8 cm. vided near the base and below the middle with 2 bracteoles long, very slender, pro 0.5-1.0 mm. long. Sepals ovate, 3-4 mm. long. Petals narrowly lanceolate, 4-6 mm. (509) [Vor. 49 198 ANNALS OF THE MISSOURI BOTANICAL GARDEN broad at the base, gradually tapering to the blunt apex, sericeous outside, glabrous and black inside. Monocarps 7, spherical or shortly cylindrical, 7-8 mm. across; stipes thin, 6-8 mm. long. Costa Rica and Panama. DEL TORO: vicinity of Chiriqui Lagoon, Water Valley, Von Wedel 1265, 1459, 1565, Joer 2464; exact locality lacking, Von Wedel 279. 3. DzsMoPsis BIBRACTEATA (Bob) Saff. in Bull. Torrey Bot. Club. 43: 190, pl. 9, Unona bibracteata Rob. in Amer. Journ. Sci. 50: 175, 1895. Shrub or small tree. Leaves on petioles 2-3 mm. long, lustrous on both sides, glabrous above, hirsute beneath at first but soon quite glabrous, oblong-lanceolate or elliptic or rhombic, cuneate at the base, shortly angustate into an obtuse or rotundate apex, 5-14 cm. long, 2.5-5.5 cm. broad. Inflorescences on peduncles 2-5 mm. long, these bearing at the apex a rotundate-cordate, foliaceous bract 0.5-1.5 cm. long; flower solitary; pedicel slender, thinly appressed-hairy, 1.5-2.5 cm. long, provided below the middle with a small foliaceous bract 1-3 mm. long. Petals oblong 10(-18) mm. long and 4-6 mm. broad, sericeous outside. Ovaries 14-20, densely sericeous; ovules 3. Monocarps on stipes about 5 mm, long, subglobose or shortly cylindric, truncate-rotundate or even umbilicate at both ends, constricted between the seeds, 5-10 mm. long and 6-8 mm. broad. Nicaragua, Costa Rica and Panama. CHIRIQUÍ: Pittier 3367, 5747 (according to Safford; the specimens not seen). 4. DesMopsis Maxont Saff. in Bull. Torr. Bot. Club 43: 188, pl 8. 1916; R. E. Fries, in Acta Hort. Berg. 10: 24, fig. 3, d-e. 1930 Tree 5-7 m. high with appressed-hairy branchlets. Leaves on petioles 5 mm. long, glabrous above, the underside pale fulvous-hirsute at first, fmally quite gla- brous, broadly elliptic or elliptic-oblong or lanceolate, acute and decurrent at base, acuminate, 10-14 cm. long and 4-8 cm. broad. Inflorescence peduncle 7-14 mm. long, bearing near the apex a suborbicular or broadly ovate, amplexicaul bract, 7-15 mm. broad, flower solitary, pale yellow; its pedicel 3.5-6.0 cm. long, curved, slender and thickened upwards, soon quite glabrous, below the middle bearing a small bracteole. Sepals broadly triangular, 4 mm. long. Petals all uniform, linear- oblong, obtuse, 13-18 mm. long and 4-5 mm. broad, clothed outside with gray ap- pressed minute hairs. Pistils 10-15, ovules 2-5. Monocarps globose or obovoid, about 8 mm. across, subsessile or shortly stipitate, the stipes 1-3 mm. long and 1.5-2.0 mm. thick. : in forest near El Boquete, alt. 1000-1300 m., Pittier 3154; in the humid forest ie HAS sss Caldera River, above El Boquete, alt. 1450-1650 m., Maxon 5564. 5. Desmopsis BREVIPES R. E. Fries in Acta Hort. Berg. 13: 107. 1941. A shrub or tree up to 10 m. tall, the young branchlets ferruginous-strigose, soon glabrescent. Leaf petioles 2-4 mm. long, appressed ferruginous-hirsute on both sides (510) 1962] FLORA OF PANAMA (Annonaceae) 199 when young, soon glabrescent, rounded or shortly acute at the base, rather abruptly cuspidate at the apex, 7-15 cm. long, 3-7 cm. broad. Inflorescences with 1-2 (or more ?) flowers opposite the leaves on young branchlets or rarely produced from the trunk; bracts small; pedicels rather thick, ferruginous-sericeous at first, more or less glabrescent, 8-10 mm. long, provided below the middle with a small bract (0.5-1.0 mm. long). Sepals rounded-deltoid, sericeous outside. Petals all uniform, linear-elongate from a dilated base, gradually narrowed into a more or less incurved apex 20-25 mm. long. Ovaries densely sericeous; ovules 3. Monocarps about 12, tomentellous at first, finally glabrescent, black when dried, very shortly stipitate, ellipsoid to more or less spherical, about 1.5 cm. across. BOCAS DEL TORO: vicinity of Nievecita, Woodson, Allen & Seibert 1864. 8. STENANONA Standl. STENANONA Standl. in Field Mus. Publ. Bot. 8: 205. 1929; R. E. Fries, in Acta Hort. Berg. 10: 151. 1931; 13: 103. 1941. Small trees. Flowers medium-sized, on short pedicels. Sepals 3, valvate, caudate. Petals 6, biseriate, all similar, long caudate-cuspidate, the outer imbricate and connate with the inner at the base. Stamens numerous; filaments short; anthers oblong, extrorse; pollen sacs not locellate; connective elongated into a deltoid ligule above the anthers. Carpels numerous; ovules 2, lateral. A small genus of 2 species, 1 of them occurring in Costa Rica, l in Panama.—The genus is very little known so far, and more complete material is desirable for a determination of its systematic position. l. STENANONA PANAMENSIS Standl. in Field Mus. Publ. Bot. 8: 205. 1929. Tree about 6 m. high; branchlets densely covered with fulvous patent hairs 1.5- 2.0 mm. long. Leaves on petioles 2-4 mm. long, membranaceous, sparsely hairy on the upper side, glabrescent, fulvous-hirsute beneath, oblong or obovate-oblong, abruptly acuminate, rotundate or subcordate at the base, 8-18 cm. long and 2.5-6.5 cm. broad. Pedicels hirsute, up to 5 mm. long, bracteate; bract superior, lanceolate, long-acuminate, about 1 cm. long. Sepals ovate, abruptly long-acuminate, 15 mm. long, 5 mm. broad at the base, hirsute on the outside. Petals about 7 cm. long, fleshy, reddish, pilose, 4-5 mm. broad at the base, 0.6 mm. at the apex. BOCAS DEL TORO: Daytonia Farm, region of Almirante. Cooper 427. 9. MALMEA R. E. Fries MarMza R. E. Fries, in Arkiv Bot. 5*: 3. 1905; in Acta Hort. Berg. 10: 37. 1939. Trees and shrubs. Flowers medium-sized or large, in few- to several-flowered inflorescences opposite the leaves, or rarely pseudo-axillary; pedicels articulated immediately above a nearly basal bract and bearing another bract above the artic- ulation. Sepals small, with thin margins, imbricate in bud. Petals much longer than the sepals, more or less blackening when dried, orbicular-elliptic, imbricate in bud, their margins thin and more or less crispate. Stamens very numerous, the (511) [Vor. 49 200 ANNALS OF THE MISSOURI BOTANICAL GARDEN Figure 161. Stenanona panamensis (512) 1962] FLORA OF PANAMA (Annonaceae) 201 connective expanded above the anther into a truncate disk; pollen sacs not locellate. Carpels numerous, containing 1 basal, erect ovule. Monocarps free, stipitate, l-seeded. Twelve species are known of this genus, 8 inhabiting Ecuador, the Ama- zonian region and Guiana; 1 is native to Bahia, and 3 are found in Central America (Mexico to Panama). Only 1 species is recorded from Panama. l. MALMEA HYPOGLAUCA (Standl.) R. E. Fries, in Acta Hort. Berg. 10: 321. 1931. Guatteria hypoglauca Standl. in Field Mus. Publ. Bot. 4: 207. 1929. A tree up to 10 m. high; young branchlets glabrous. Leaves on petioles 5-6 mm. long, glabrous and glaucous beneath, elliptic or oblong-elliptic, rounded at the base and shortly decurrent, shortly acuminate at the obtuse apex, 12-14 cm. long and 5-6 cm. broad. Inflorescences terminal (or opposite the leaves ?); pedicels about 5 mm. long, thinly ferruginous-hirsute. Sepals 2-3 mm. long. Petals thick and glabrous, blackening when dried, elliptic-obovate and rotundate at the apex, 15-18 mm. long and 11-12 mm. broad. Stamens 2 mm. long. Fruit unknown, SAN BLAS: Permé, Cooper 661. This species is also found in Colombia (Department of Antioquia). 10. PORCELIA R. & P. Porcetia R. & P., Fl. Peruv. & Chil. Prodr. 84, tab. 16. 1794; R. E. Fries, in Acta Hort. Berg. 10: 28, fig. 4. 1930. Trees. Leaves ovate to lanceolate-oblong, nearly glabrous, with translucent glands. Flowers medium-sized, bisexual or polygamous, terminal or opposite the leaves, often produced on axillary, very reduced branchlets (pseudo-axillary flowers) ; pedicels articulate at the base, without bracts. Sepals small. Petals im- bricate in bud and finally widely spreading, all of about the same size, oblong- ovate or obovate. Stamens numerous; pollen sacs locellate; connective expanded above the anther into a truncate disk. Ovules numerous, lateral. Monocarps (sometimes very) large, free, stipitate, ellipsoid-cylindric, with large, reniform seeds. A small genus of 5 species. Two of them (macrocarpa and goyazensis) are indigenous in eastern Brazil, 1 (nitidifolia) in Peru and Ecuador, 1 (stein- bachii) in eastern Bolivia and adjacent parts of the Amazonas basin and | (magnifructa) in Panama and Venezuela. l. PoRcELIA MAGNIFRUCTA (Schery) R. E. Fries, in Arkiv Bot. andra ser. 1:343. 1950. Cymbopetalum magnifructum Schery, in Ann. Missouri Bot. Gard. 30: 87. 1943. A tree about 12 m. high; young shoots and pedicels densely tomentellous, ochraceous, finally glabrous. Leaves on petioles 2-8 mm. long, ovate to oblong, rotundate to shortly acute at the base, longer acuminate at the apex, grey-ochraceous (513) [Vor. 49 202 ANNALS OF THE MISSOURI BOTANICAL GARDEN Figure 162. Porcelia magnifructa tomentellous at first on both sides, especially beneath, soon glabrescent with translucent glands, up to 15 cm. long and 4-5 cm. broad. Pedicels 2-3 cm. long, without bracts. Sepals ovate, 3-5 mm. long, grey-tomentellous. Petals ovate, the apex rotundate and more or less incurved when dried, up to 17 mm. long, tomentellous on both sides. Stamens 2.5-3.0 mm. long, pollen sacs locellate. Carpels 10-15. Monocarps large, 6-10 cm. long and about 6 cm. across, ovoid; peri- carp thick, ligneous. Seeds 2.5-3.0 cm. long. PANAMA: vicinity of Bejuco, alt. ca. 50 m., Allen 2455.—Apparently also in Venezuela. 11. CYMBOPETALUM Benth. CyMBOPETALUM Benth. in Journ. Linn. Soc. 5: 69. 1861; R. E. Fries, in Acta Hort. Berg. 10: 180. 1931. Trees or shrubs with large, rather thin leaves. Flowers solitary, usually pro- duced from the internode above the axil, sometimes terminal or rarely pseudo- axillary; pedicels often long, articulate at the base, without bracts. Perianth seg- ments valvate in bud; sepals small; outer petals flat and thin, the inner longer, thick and fleshy with involute margins, cymbiform. Stamens numerous; pollen sacs (514) 1962] FLORA OF PANAMA (Annonaceae ) 203 locellate; connective expanded above the anther into a truncate disk. Fruit of several free, oblong-cylindric monocarps, containing several (4-14) lateral seeds. A small genus of 19 species, distributed from Mexico to Brazil (Rio de Janeiro). Only 3 species are found in Panama. a. Pedicels 5 cm. long or EE? much longer; flowers large; inner petals 2.5-3.5 cm long; stamens 4.0-4.5 mm. lon Leaves obovate- GE long-cuneate at the base. .................. l. C. BRASILIENSE. bb. Leaves narrowly oblong-lanceolate, rounded-acute at the base. 2. C. LANUGIPETALUM. Pedicels about : cm. long; flowers small; inner petals 1.5 cm. long; stamens 2.0-2.5 mm. lon 3. C. COSTARICENSE, m e l. CYMBOPETALUM BRASILIENSE (Vell.) Benth. in Journ. Linn. Soc. 5:69. 1861; R. E. Fries, in Svensk. Vet.-Akad. Handl. ser. 3. 241°: 29. 1900. U varia ry Vell. Fl. Flumin. a 1825; 5: tab. 122. 1827; Eichl. in Mart. Fl. Bras: 13°: 39, tab, 15, fig. 2 Unona n Splitg. in Nederl. i, Arch. 1: 224. 1848. Uvaria viridiflora Walp. Ann. Bot. Syst. 2: 19. 1851. Eschweilera simplex Miers, in Trans. Linn. Soc. 30: 264. 1875. Trigynaeia anastomosans Rusby, Descr. New Species of S. Amer. Plants 19. 1920. Uva brasiliensis O. Ktze. Rev. Gen. 7. 1891. A small tree with shortly puberulous branchlets, soon glabrescent. Leaves on petioles 3-4 mm. long, membranaceous, glabrous and densely pellucid-punctate, obovate, oblanceolate, or lanceolate-elliptic, cuneate-angustate toward the base, 15-30 cm. long and 6-10 cm. broad. Pedicels 5-9 cm. long, glabrous. Sepals con- nate at the base, rounded-reniform, obtusely acuminate, 4-6 mm. long and 6-9 mm. broad. Petals tomentellous on both sides, the outer ovate or rotundate, flat, 2.0-2.5 cm. long and 2-3 cm. broad, the inner thick, rounded-elliptic, obtuse, zd -3.5 cm. long and 1.8-2.3 cm. broad, the margins involute. Stamens 4.0-4. long. Monocarps numerous, oblong-cylindric, and more or less curved, torulose, rounded at both ends, up to 3.5 cm. long and L5 cm. broad. Seeds up to 6, but generally fewer. This species is distributed in eastern South America from Trinidad and Venezuela down to Rio de Janeiro; in western South America from Panama to south Colombia. DARIEN: Río Cuasi, alt. 800 ft., Terry & Terry 1417. 2. CYMBOPETALUM LANUGIPETALUM Woods. & Schery, in Ann. Missouri Bot. Gard. 28: 427. 1941. Tree; young branchlets very shortly appressed- -hirsute. Leaves on very short petioles (1-2 mm. long), rigid, glabrous on both sides, oblong-lanceolate, rounded or very shortly acute at the base, gradually tapering toward a long narrow cusp, 15-30 cm. long and 5-7 cm. broad, the nerves impressed on the upper side, very prominent beneath. Flowers solitary; pedicels glabrous, pendulous, about 20 cm. (515) [Vor. 49 204 ANNALS OF THE MISSOURI BOTANICAL GARDEN Figure 163. Cymbopetalum lanugipetalum long. Sepals small, about 0.3 cm. long. Outer petals flat and thin, subtriangular, l cm. long and 1.0-1.6 cm. broad, the inner fleshy and tomentellous, tri- angular-ovate, cymbiform, about 3 cm. long and 1.5 cm. broad. Stamens 4-5 mm. long, the pollen sacs locellate, the connective-appendage globose, setulose. cocL£: north rim of El Valle de Antón near Cerro Turega, alt. 650-700 m., Woodson & Schery 191a. (516) 1962] FLORA OF PANAMA (Annonaceae) 205 3. CYMBOPETALUM COSTARICENSE (Donn. Sm.) R. E. Fries, in Acta Hort. Berg. 10: 186, fig. 6 a-c d. 1931. Asimina costaricensis Donn. Sm. in Bot. Gaz. 23: 2. 1897. Tree or shrub; young branchlets sparsely clothed with very short appressed hairs, soon glabrescent and densely provided with lenticels. Leaves on petioles -6 mm. long, membranaceous, finally glabrous, verruculose along the thicker nerves, oblong, elliptic-oblong or obovate, cuneate at the base, rather abruptly cuspidate, 10-22 cm. long, 4.0-6.5 cm. broad. Pedicels about 3 cm. long. Sepals rounded-ovate, shortly apiculate, about 4 mm. long. Petals grey-tomentellous, the outer flat, broadly ovate, acute, up to 12-13 mm. long, the inner obovate-rotundate, about 17 mm. long and 10 mm. broad, narrowed at the base into a petiole 5 mm. long, the margins of the upper part striate and involute. Stamens 2.5 mm. long, connective-disk plane and tomentellous. Monocarps 8-10, shortly cylindric, not torulose, 3-5 cm. long and about 1.7 cm. thick. Seeds 12-14. Costa Rica and Panama. BOCAS DEL TORO: Daytonia Farm, Cooper 624; vicinity of Chiriquí Lagoon, Water Valley, Von Wedel 693, 1768. 12. DUGUETIA St. Hil. Ducurria St. Hil. Fl. Bras. Mer. 1: 35, t. 7. 1825; R. E. Fries, in Acta Hort. Berg. 12: 28. 1934 Trees or shrubs with stellate hairs or stellate scales. Flowers medium-sized or large in few- or multi-flowered inflorescences; these more or less strictly opposite the leaves, or sometimes produced from older branches or from the trunk. Sepals valvate. Petals free, generally imbricate in bud. Stamens numerous; connective generally expanded above the anther into a truncate disk. Carpels numerous; ovules solitary, basal, erect. Monocarps sessile, closely crowded, ligneous or fleshy, easily separable or more or less coherent. A large genus (about 70 species), distributed from Panama and the West Indies to southern Brazil and Paraguay; especially rich in the Amazonian region and Guiana. a. Stellate scales abundant; monocarps very rugose above the middle ......1. D. PANAMENSIS. aa. No scales, only stellate hairs; monocarps smooth ..... nn 2. D. vALLICOLA. l. DUGUETIA PANAMENSIS Standl. in Field Mus. Publ. Bot. 4: 207. 1929; R. E. Fries, in Acta Hort. Berg. 12: 53. 1934. A tree about 10 m. high, the young branchlets densely covered with golden- yellow stellate scales. Leaves on 3-4 mm. long petioles, 12-22 cm. long and 3.5-5.0 cm. broad, chartaceous, glabrous on upper side from the first, densely scaly under- neath when young, finally nearly glabrescent, oblanceolate, cuneate and very acute at the base, contracted at the apex into an obtuse acumen 1.5-3.0 cm. long. In- florescences 1- to 3-flowered, opposite the leaves, shortly pedunculate. Flowers on pedicels 2-4 mm. long, small, fragrant. Sepals about 10 mm. long, ovate, yellow- (517) [Vor. 49 206 ANNALS OF THE MISSOURI BOTANICAL GARDEN scaly outside. Petals oblong, obtuse, rather densely silvery-scaly outside, up to 13 mm. long and 5 mm. broad. Stamens 1 mm. long, the connective-disk glabrous. Ovaries glabrous. Fruit spherical, 3.5-4.5 cm. across; monocarps numerous, free, clavate, 16-18 mm. long and 7-8 mm. thick, very rugose above the middle, rotun- date and very shortly apiculate at the apex. BOCAS DEL TORO: region of Almirante, Daytonia Farm, Jan. 1928, Cooper 418. The appearance of the fruit of this species is unique in its genus. Towards their upper parts, the monocarps are densely covered by protuberant, conical, and acuminate excrescences, about a millimetre long, making their appearance so foreign to the genus that the specimen might easily be taken for a monstrosity (galls ?). Nevertheless, the species is well differentiated from the other Duguetias also by other characters. 2. Ducueria varııcor.a Macbr. in Contr. Gray Herb. n. ser. 56: 51. 1918; R. E. Fries, in Acta Hort. Berg. 12: 85. 1934. A shrub or tree, up to 20-25 m. high. Leaves on petioles 4-7 mm. long, provided at first with small stellate grayish hairs on the upper side and finally glabrescent, densely stellate-hirsute at first beneath, finally nearly glabrous with scattered hairs, oblong-lanceolate or narrowly elliptic, rather gradually tapering at the apex, narrowed into the rotundate-obtuse or shortly acute base, 15-30 cm. long and 4.5- 8.0 cm. broad. Flowers opposite the leaves or below the nodes; pedicels gray- tomentellous, 12-15 mm. long (in fruits up to 25 mm.). Sepals reflexed, ovate, gray-tomentellous on both sides, 8-12 mm. long. Petals ovate, obtuse, longitu- dinally striate at the base, densely covered on both sides with very short, gray. stellate hairs, up to 2.5 cm. long and 1.5 cm. broad. Stamens 1.3-1.5 mm. long. Fruit 4.0-4.5 cm. diam.; monocarps free, glabrous but more or less pruinose, obovoid, pentagonal, shortly apiculate, up to 18 mm. long and 7-8 mm. across. DARIÉN: forest around Pinogana, Pittier 6565. Described from Colombia, Magdalena Valley; probably also occurring in Venezuela. 13. ANNONA L. ANNONA L. Sp. Pl. 536. 1753 [Anona]. Shrubs or trees with simple or stellate hairs. Flowers solitary or in few- flowered inflorescences, terminal or opposite the leaves, or extra-axillary from the internodes. Sepals 3, small, valvate. Petals 6, free or connate at the base, biseriate, or the inner rudimentary or absent, the outer valvate, the inner imbricate or valvate. Stamens numerous, the anther cells not locellate, the connective generally terminat- ing in a swollen head or hoodlike process above the pollen sacs. Carpels numerous or rarely few; ovule 1, basal, erect. Fruit fleshy, formed by the concrescence of the carpels and the torus, usually areolate on the surface, the areoles, indicating the united carpels, being often gibbous or acuminate. (518) 1962] FLORA OF PANAMA (Annonaceae) Figure 164. Duguetia panamensis (519) 207 [Vor. 49 208 ANNALS OF THE MISSOURI BOTANICAL GARDEN A large genus of about 110 species in all the tropics of America from Florida, Mexico and the West Indies to southern Brazil and Paraguay. A few species also in tropical Africa. Five indigenous species are recorded from Panama. a. Petals 6. b. € free Branch ints and leaves glabrous; sepals 3-5 mm. long; petals SCH mm. long, the inner valvate A. GLABRA. cc. Branchlets and leaves hirsute; sepals l-2 cm. long; outer pt up to 5 cm. long, the inner imbric 2. A PURPUREA. bb. Petals connate We ‘bile 9. A. HAYESII aa. Petals 3, the inner missing or very rudimentary b. Leaves thickly tomentose beneath, 10-20 cm. long 3. A. SPRAGUEI. bb. Leaves glabrous, 6-8 cm. long 4. A. ACUMINATA. l. ANNONA GLABRA L; Sp. Pl. 537. 1753. Anona palustris L. Sp. Pl. 757. 1162. Anona laurifolia Dunal, Monogr. Anonac. 65. 181T. Anona. peruviana Humb. & Bonpl. ex Dunal, 1. c. 67. 1817. Anona uliginosa H. B. K. Nov. Gen. & Sp. Pl. 5: 65. 1821. Anona australis St. Hil. Fl. Bras. Mer. 1: 33. 1825. Anona pisonis St. Hil. & Tul. in Ann. Sci. Nat. sér. 2. 17: 131. 1842, non Mart. A tree up to 10 metres high; branchlets very soon glabrous. Leaves on petioles 1.0-1.5(-2.5) cm. long, rather thin, pale green, ovate-elliptic or oblong-elliptic, rotundate or acute and decurrent at the base, shortly acute or rarely obtuse at the apex, 7-14 cm. long and 3-8 cm. broad; network of veins dense and nicely prominent on both sides. Flowers solitary, issuing from about the middle of the internodes below the leaves; pedicels 1.5-2.0 cm. long, glabrous. Sepals rotundate, apiculate, 3-5 mm. long. Petals glabrous on outside, the outer ovate, 2.5-3.0(-4.0) cm. long, the inner a little smaller. Stamens 3-4 mm. long, connective disk broad, papillose but not setose. Fruit globose-ovoid, rounded at the top, 7-12 cm. long, smooth, areoles hardly obvious. This species is distributed from Mexico and the West Indies southwards to Ecuador on the west side and to southern Brazil on the east, also on the west coast of tropical Africa; always growing near the sea in littoral forests, in mangrove swamps and on riverbanks. CANAL ZONE: Chagres, Fendler 1; forest at An Indio de Gatün, near sea-level, Pittier 2809; in swampy places near Panama, S. Hayes s.n. socas DEL TORO: Old Bank Island, Von "Wedel 2082. Also reported from Barro Colorado Island (Standley, 1933). 2. ANNONA PURPUREA Moc. & Sesse, ex Dunal, Monogr. Anonac. 64, tab. 2. 1817. Anona manirote H. B. K. Nov. Gen. & Sp. Pl. 5: 59. 1821. Anona involucrata Baill. in Adansonia 8: 265. 1867-68. Anona prestoei Hemsl. in Hook. Ic. Pl. 4: 6, t. 2519-20. 1897. (520) 1962] FLORA OF PANAMA (Annonaceae) 209 Tree with ferruginous-tomentose branchlets. Leaves deciduous, on 3-5 mm. long petioles, membranaceous, on the upper side shortly and thinly, on the nerves more densely hirsute, finally more or less glabrate, underneath longer ferruginous- hirsute along the nerves, obovate or elliptic-obovate, rotundate at the base, shortly acuminate at the apex, 12-30 cm. long and 6-14 cm. broad. Flowers solitary, sub- sessile; flower bud enclosed at first by an involucre composed of 2 sessile acuminate bracts. Calyx lobes triangular-ovate, acuminate, 1-2 cm. long. Outer petals valvate, thick, rigid, ovate-lanceolate, gradually tapering into a long, obtuse apex, up to 5 cm. long and 2 cm. broad, ferruginous-sericeous on the outside, inner petals imbricate, thinner, elliptic-oblong, rotundate at the apex, 2.5 em. long. Stamens 5-6 mm. long. Fruit globose, up to 20 cm. in diameter, bearing numerous pyramidal protuberances and clothed with brown felt-like tomentum. Seeds 28-30 cm. long. This species is recorded from Mexico, British Honduras, Costa Rica, Panama, Venezuela, Ecuador and Trinidad. CANAL ZONE: Ancón Hill, alt. 200 ft. Allen 4521; Hospital grounds, Ancon, Pittier 3955; Barro Colorado Island, Bangham 610; Matachín, on Panamá Railway, Hayes s.n. CHIRIQUI: forest of San Félix, Pittier, 5749; Chiriqui, Cooper & Slater P 314. The species *is common on the Pacific slope" (Standley, 1928, p. 180). 3. ANNONA SPRAGUEI Saff. in Contr. U. S. Nat. Herb. 16: 270, fig. 43, pl. 92-93 1913. Anona uncinata Sprague, in Bull. Herb. Boiss. ser. 2. 5: 701. 1905, non Lam. A tree up to 16 m. high; young branchlets ferruginous-tomentose, soon glabres- cent. Leaves on petioles 8-15 mm. long, membranaceous, pellucid-punctulate, sparsely pubescent above, at first very densely and softly sericeous-pubescent beneat with appressed grayish olivaceous hairs, rufous-tomentose along the midrib and lateral nerves, lanceolate to oblanceolate or narrowly elliptic, shortly acute or some- times subrotundate at the base, acuminate at the apex, 15-35 cm. long and 5-11 cm. broad. Flowers solitary; pedicels about 1 cm. long, in fruit up to 2 cm. long (or longer ?); flower buds globose. Sepals long-caudate from an ovate base, 8-10 mm. long. Petals thick, rotundate-ovate, 18-25 mm. long, tomentellous on the outside. Stamens 3.0-3.5 mm. long. Fruit globose or globose-ovoid, about 5 cm. in diameter; the areoles produced into long-attenuate protuberances. Seeds oblong, 7-9 mm. long. Known only from Panama. CANAL ZONE: Gamboa, Pittier 3409; R. Cocoli, opposite lighthouse, P. White 99; Frijoles, Standley 27570. panamá: San Jose Island, Perlas Archipelago, Johnston 734; arbacoas, Hayes 127; Tapia River, Juan Diaz region, Maxo arvey . DARIEN: Marraganti and vicinity, Rio Tuyra, 10-200 ft. elev., Williams; forest around Yaviza, Pittier 6540. Also reported from Barro Colorado Island (Standley 1933). 4. ANNONA ACUMINATA Staff. in Contr. U. S. Nat. Herb. 16: 274. pl. 97. 1913. Anona echinata Hemsl. Biol. Centr.-Amer. Bot. 1: 19. 1879, non Dunal. Tree, 5-7 m. high; young branchlets clothed with very minute appressed hairs, soon glabrate. Leaves on petioles 3-5 mm. long, membranaceous, finally glabrous, pellucid-punctulate, lanceolate or oblong-elliptical, acute and decurrent at the base (521) [Vor. 49 210 ANNALS OF THE MISSOURI BOTANICAL GARDEN \ 7 if L d t Figure 165. Annona acuminata 1 and gradually acuminate at the apex, 5-9 cm. long and 1.5-2.5 cm. broad. Flowers solitary, more or less opposite the leaves; pedicels 12-16 mm. long, with 2 linear- lanceolate bracts 2-4 mm. long; flower buds subglobose, about 15 mm. in diameter. Sepals high-connate, appressed-pilose on the outside. Petals 3, ovate-rotundate, connate at the base, 12-15 mm. long and broad. Stamens 2.5 mm. long; connective disk muriculate with short stiff points. Fruit globose, 2.0-2.5 cm. in diameter, with conical spines 1 mm. long. Seeds yellow, 7-8 mm. long. (529) 1962] FLORA OF PANAMA (Annonaceae) 211 Recorded only from Panama. NE: vicinity of Salamanca Hydrographic Station, Rio Pequeni, alt. ca. 80 m., wm: len & Seibert 1564; drowned forest of Quebrada Tranquilla and its Stiech odge & Allen 17331; Bohio station, Panamá railroad, Hayes 142. PANAMÁ: Rio Tapia Standley 26212, 28036. Reported from Barro Colorado Island (Standley 1933). 5. ANNONA HAYEsII Saff. ex Standley, in Journ. Washington Acad. Sci. 15: 102. 1925; R. E. Fries, in Acta Hort. Berg. 10: 273, tab. 21. 1931. A high shrub or small tree; branchlets densely sericeous-tomentose, ferruginous. Leaves on petioles 3-7 mm. long, thin-membranaceous, underneath pale and at first appressed-hirsute, soon glabrous except on the nerves, obovate or elliptic, abruptly acuminate and rather rotundate at the base, 8-12(-25) cm. long and 4-9 cm. broad. Inflorescences opposite the leaves or produced from the middle of the internodes below the leaves, l- or few-flowered; pedicels 12-15 mm. long, densely ferruginous-hirsute; flower buds long-caudate from a globose base. Sepals ovate, acuminate. Outer petals up to 30 mm, long, connate up to 5 mm. above the base, the base ovate, abruptly narrowed into a linear obtuse acumen nearly 20 mm. long, rufous sericeous-tomentellous on the outside; inner petals minute, 5-7 mm. long, connate with the outer at the base. Stamens 1.2-15 mm. long, the con- nective disk narrower than the anther. Fruit subglobose, smooth, 5 cm. long or more. Found only in Panama. one: Ancón Hill, alt. 200 ft., A ar a Bahr 141. DARIÉN: forest at Yen viza, Pieter 6592; La Palma, alt. 50 m., er 6598. A: thickets and forests near Arraijan, alt. ca. 15 m., Woodson, Allen & dos I372; "Matías Hernández, Pittier 6749. 5 Also reported from Beo Colorado Island (Standley, 1933). 14. ROLLINIA St. Hil. Rorınıa St. Hil. Fl. Bras. Merid. 1: 28, t. 5. 1825; R. E. Fries, in Acta Hort. Berg. 12: 112. 1934. Trees and shrubs. Flowers usually clustered in inflorescences opposite the leaves, or extra-axillary from the internodes. Sepals small, valvate. Petals valvate, connate at the base, the outer produced on their outside into a wing or spur. Stamens numerous, their anther cells not locellate; connective expanded above the anther into a truncate disk. Carpels several to numerous, united into a fleshy syncarpium; ovule solitary, basal, erect. A large genus of about 62 species, distrubuted from Central America and the West Indies to southern Brazil, Paraguay and northern Argentina. Only 3 species are recorded from Panama. it pals shin, fas er. inde; pal man are PLR — aa. Sepals more or less concave, the midrib thickened and keeled on the weet Sepals about 3 mm. long; petal wings recurved ne E long; petal wings spreading but not bb. meet small, 1.0-1.5 mm, long; pe EE i LI PITTIERI Gees EE EEN EG [Vor. 49 212 ANNALS OF THE MISSOURI BOTANICAL GARDEN Rollinia chocoensis R. E. Fr., which, according to E. P. Killip, occurs “along the Pacific coast of Colombia from the Darién region of Panama into northern Ecuador” (cf. R. E. Fries in Sv. Vet.-Akad. Handl., ser. 3, 241%: 18. 1948), might possibly be added to the above. As, however, no specimen from Panama has been available and as it is hard to distinguish it from closely related species, it is perhaps better for the time being not to include it in the flora of that country. 1. RoLLINIA PERMENSIS Standl. in Field Mus. Publ. Bot 4: 208. 1929. A small tree about 6 m. high; young branchlets, petioles and flower-pedicels covered with ferruginous appressed to patent hairs. Leaves membranaceous, 20-30 cm. long and 8-13 cm. broad, obovate or elliptic-obovate, cuneate or shortly ro- tundate-acute at the base, 2-3 cm. long, cuspidate at the apex; upper side thinly hairy when young, soon glabrescent, with persistent hairs on the impressed mid- rib and thicker nerves, densely hairy underneath, finally nearly glabrescent, the midrib with short appressed glossy hairs. Inflorescences more or less opposite the leaves on peduncles about 3 mm. long, bearing up to 8 flowers; pedicels up to 2.5 cm. long. Sepals rotundate-ovate, ferruginous-sericeous, 3-4 mm. long. Corolla 2.0-2.5 cm. across, tomentellous, gray at the base, more densely ferruginous upward; wings divergent and curved upwards a little, oblong, about 10 mm. long and 5 mm. broad. CHIRIQUÍ: Cooper & Slater 211. san BLAs: Permé, Cooper 645. 2. ROLLINIA PITTIERI Saff. in Journ. Washington Acad. Sci. 6: 376. 1916. Tree with the young branchlets very shortly sericeous. Leaves on petioles 7-12 mm. long, membranaceous, 8-20 cm. long and 4.5-8.5 cm. broad, elliptic or obovate- elliptic, shortly rotundate-acute at the base, abruptly contracted at the apex into a cusp about 1 cm. long, nearly glabrescent above, glaucous beneath, the nerves cin- namomeous and rather densely covered with short appressed white hairs. In- florescences sessile, opposite the leaves or issuing a little below the nodes; flowers up to 6, on pedicels 3.5(-5.0) cm. long. Sepals triangular, 2-3 mm. long, the mid- rib thickened and keeled on the outside. Corolla 2.5-3.0 cm. across; wings very densely and shortly silvery-tomentellous, obviously recurved, 1.0-1.5 cm. long and 0.7-1.0 cm. broad. (Fruits unknown). SAN BLAS: plain of Sperdi, near Puerto Obaldia, near sea level, Pittier 4358. 3. ROLLINIA MICROSEPALA Standl. in Field. Mus. Publ. Bot. 4: 208. 1929. Young branchlets fulvous-sericeous, the hairs short-appressed. Leaves on peti- oles 6-10 mm. long, 10-18 cm. long and 3.5-5.0 cm. broad, oblong or lanceolate- oblong, gradually narrowed at the apex, acute or rotundate at the base, glabrous above and shortly sericeous beneath. Inflorescences about 3-flowered, opposite the leaves or produced a little below the nodes; pedicels densely sericeous, 15-18 mm. long. Sepals minute (1.3-1.8 mm. long), curvate, concave, sericeous. Corolla gray, tomentellous; petal wings cuneate or narrowly obovate, broadly rotundate at the (524) 1962] FLORA OF PANAMA (Annonaceae ) 213 apex, spreading but not recurved, 15-18 mm. long and 8-10 mm. broad. Stamens 0.6 mm. long. Fruit globose, rather smooth and very shortly tomentellous, 15-17 mm. diam. OCAS DEL TORO: Changuinola Valley, Cooper & Slater 9, 102. This species is also recorded from Costa Rica: San Carlos, Kotschny. Figure 166. Rollinia permensis (525) [Vor. 49 214 ANNALS OF THE MISSOURI BOTANICAL GARDEN MYRISTICACEAE By J. A. DUKE Aromatic trees or shrubs, not infrequently with buttressed bases, whorled branches, stellate pubescence, and reddish sap. Leaves alternate, exstipulate, penninerved, entire, often coriaceous, occasionally with pellucid punctation or stellate pubescence. Flowers unisexual, usually dioecious, actinomorphic, basically trimer- ous, solitary or more usually fascicled in racemes or panicles, or in dichotomous cymes. Perianth uniseriate, of typically 3 partially fused carnose tepals. Stamens 2-30, the filaments united into a column, the 2-locular extrorse anthers free or co- herent to the column. Pistillodes absent. Ovary superior, l-carpellate, the stigma subsessile; with a single basal anatropous ovule. Fruit fleshy, the thick pericarp splitting longitudinally into 2 valves, the seed often with a reddish laciniate aril, the endosperm ruminate. A tropical group of about 15 genera centered in America, Africa and Asia, this family is represented in the Americas by 6 genera, 1 of them including the nutmeg of commerce Myristica fragrans, locally introduced. Little can be added to the excellent monograph of the American representatives by A. C. Smith (in Brittonia 2: 393. 1937). Four of the 6 American genera occur in Panama. a. E and young twigs glabrous; secondary veins 4-12 pairs, the tertiary veins etimes cons p ap allel, almost i es tte to the midrib; anthers 4- 30, at least as long as umn b. Tertiary veins not Fee? perpendicular to the midrib; staminate flowers few in dichotomously branching cymes, the perianth 3-7 mm. long; anthers 12-30; pistillate pedicels 6-20 mm. long; arils laciniate ............ 1. MYRISTICA Er SE Tertiary veins strikingly perpendicular to the midrib; staminate flowers several, fasciculate in subspicate racemes, the perianth 1.5-3.0 mm. long; anthers 4-8; piedini pedicels 2-4 mm. long; arils subentire ............ 2. COMPSONEURA HI D Leaves and young twigs pubescent, occasionally glabrate; secondary veins 5-35 re the tertiary veins not nrg es perpendicular to the midrib; anthers gue longer or shorter than the e em "e the young branchlets Sr Ee with only 1 or 2 conspicuous s; secondary veins 5-25 pairs; staminate er A subcorymbosely to ger disposed, the Deas usually free to their bases, pus connate dorsally DIALYANTHERA e e Hairs of the young ae oe petioles with several gare branches; secondary veins 10-35 pairs; staminate fasicles racemosely eat disposed, the anthers docsolly connate or distally Grae never free to their base 4. VIROLA 1. MYRISTICA Boehm. Myristica [L.]Boehm. in Ludw. Def. 3: 513. 1760, nom. conserv. Comacum Adans. Fam. 2: 345. 1763, nom. rejic. Aromatic, glabrous, dioecious trees, the bases occasionally buttressed. Leaves alternate, subdistichous, glabrous, subcoriaceous, petiolate, exstipulate, the tertiary veins obscure. Flowers rather large for the family, the bracteate pedicels glabrous. (526) ^ 1962] FLORA OF PANAMA (Myristicaceae) 215 Staminate flowers 1-20 in dicho- tomously branching cymes; tepals partially connate, usually 3, the anthers 12-30, usually longer than the column. Pistillate flow- ers commonly solitary, the tepals partly connate, the Learpellate ovary with a subsessile 2-lobed stigma. Fruits fleshy, the thick pericarp dehiscing longitudinally into 2 valves; aril usually brightly colored and laciniate; seed ellip- soid to globose. Of this austral Asian genus of around 75 species, only the commonly cultivated nutmeg, a native of the Moluccas, is found in Panama. Figure 167. Myristica fragrans 1. MYRISTICA FRAGRANS Houtt. Handleid. Hist. Nat. Linn. 2: 333. 1774. Myristica officinalis L.f. Suppl. 262. 1781. Myristica moschata Thunb. in Vet. Akad. Handl. Stockh. 49. 1782. Myristica aromatica Lam. Act. Acad. Sci. Paris 1788: 155. 1791. Aromatic, dioecious, glabrate trees to 20 m. high, the older bark rather smooth and olivaceous, mottled with white, the younger branches often mottled with red. Leaves subcoriaceous, glabrous, lanceolate to ovate or obovate, apically acute to acuminate, basally acute, 6-12 cm. long, 3-6 cm. broad, with usually 6-11 pairs of secondary veins, the tertiary nerves obscure, not conspicuously perpendicular to the midrib, the petioles 5-15 mm. long, ca. 1 mm. broad. Staminate flowers | to few in dichotomous cymes; pedicels glabrous, 5-15 mm. long; bracts ca. 1 mm. long; perianth tardily 3-parted, 3-7 mm. long; anthers 12-30, 2-3 mm. long, dorsally co- herent to the column, the infra-antheral portion of the column 1-2 mm. long. Pistillate flowers solitary or rarely paired in the axils; pedicels glabrous, 5-15 mm. long; bracts ca. 1 mm. long; perianth accrescent, tardily 3-parted, the segments del- toid, the subsessile stigma obscurely 2-lobed. Fruit fleshy, ovoid to pyriform, the pericarp splitting longitudinally into 2 valves, 3-6 cm. long, 2.5-45 cm. broad; seed 1.5-4.5 cm. long, 1.0-2.5 cm. broad, the laciniate aril reddish. Native to the Moluccas, widely cultivated in tropical America. BOCAS DEL TORO: vicinity of Chiriquí Lagoon, Von Wedel 1232. (527) [Vor. 49 216 ANNALS OF THE MISSOURI BOTANICAL GARDEN Many spices are extracted from this utilitarian tree, now largely cultivated in southeastern Asia and Grenada. Mace is derived from the aril, and nutmeg from the pulverized seed; a non-drying oil, the so-called nutmeg butter, is also ex- pressed from the seeds. Rumor has it that imperial bureaucrats of little botanic bent once ordered a speedup in the culture of nutmeg trees and a cutback in pro- duction for mace trees. 2. COMPSONEURA Warb. Figure 168. Compsoneura sprucei (528) CoMPSONEURA Warb. in Ber. Deutsch. Bot. Ges. 13:83, 1895, hyponym; in Nov. Act. Acad. Leop.-Carol. 68: 125. 1897. Dioecious glabrous shrubs or trees, the sap often reddish. Leaves alternate, glabrous, char- taceous to coriaceous, petiolate, exstipulate, the tertiary veins par- allel, conspicuously perpendicu- lar to the midrib. Flower fascicles in spikes, racemes or panicles, the axes glabrous, the bracts absent or inconspicuous. Staminate flowers 3-25 per fascicle, the 3 (-5) tepals partially united; an- thers 4-10, as long as or longer than the column. Pistillate flow- ers 1-8 per fascicle, the tepals partly connate, the 1-carpellate ovary with a subsessile 2-lobed stigma. Fruits fleshy, the thin pericarp dehiscing longitudinally into 2 valves; aril usually brightly colored and subentire; seed ellip- soid. In this genus of 8 species, that which occurs in Panama essen- tially covers the entire range of the genus, from southern Mexico through Central America to Am- azonian Peru and Brazil. 1962] FLORA OF PANAMA (Myristicaceae) 217 1, CoMPSONEURA SPRUCEI (A. DC.) Warb. in Nov. Act. Acad. Leop.-Carol. 68: 143. 1897 Myristica sprucei A. DC. in DC. Prodr. 14: 199. 1856. Myristica mexicana Hemsl. Biol. Centr.-Am. Bot. 3:67. 1882. Compsoneura costaricensis Warb. in Fedde, Repert. Sp. Nov. 1:71. 1905. Glabrate shrubs or trees to 13 m., the sap reddish. Leaves chartaceous or sub- coriaceous, glabrous, oblong to obovate, apically acute to acuminate, basally atten- uate to rounded, 9-30 cm. long, 3.5-10.0 cm. broad; secondary veins (4-)6-10 (-12) on either side; tertiary veins parallel, conspicuously perpendicular to the midrib; petioles 7-30 mm. long, 1-3 mm. broad. Staminate fascicles 3- to 15-flowered, in subspicate racemes (in Panama), the racemes 1-5 per axil, 2-8 cm. long; pedi- cels 0.5-2.0 mm. long, perianth tardily 3(-4) parted, 1.5-3.0 mm. long, anthers 4-8, 1.0-1.5 mm. long, free or somewhat connate basally, the infra-antheral portion of the column less than 0.5 mm. long. Pistillate fascicles 2-8 cm. long, l- to 8- flowered; pedicels 2-4 mm. long; perianth accrescent, tardily 3(-4) parted; ovary subglobose, with a subsessile 2-lobed stigma. Fruit somewhat fleshy, ellipsoid, the perianth splitting longitudinally into 2 valves, 20-37 mm. long, 13-21 mm. broad, the reddish aril entire except at the tip. Southern Mexico to Amazonian Peru and Brazil, at low elevations. CAS DEL TORO: Water Valley, vicinity of Chiriqui Lagoon, Von Wedel 598, 946, Ki ? vicinity of Chiriquí Lagoon, Von Wedel 1004, 2382, 2462; Daytonia > arm, region o Almirante, Cooper 420; Changuinola Valley, Cooper & Slater 84. coró de dE eg la Gloria, near Fató (Nombre de Dios), 10-104 m., Pittier 4099, 3846. a BLAS: Permé, Cooper 241. Dr. A. C. Smith observes (in Brittonia 2: 411. 1937) that the Central American material varies from the Amazonian material in having the fascicles subsessile on the rhachis, but he concludes that the variation is of no taxonomic consequence. 3. DIALYANTHERA Warb. DIALYANTHERA Warb. in Ber. Deutsch. Bot. Ges. 13:83. 1895, hyponym; in Nov. Act. Acad. Leop.-Carol. 68: 126. 1897. Dioecious trees often with reddish sap, the younger branchlets pubescent. Leaves alternate, glabrous above, strigillose below with 2- to few-branched hairs, charta- ceous or subcoriaceous, petiolate, exstipulate, the tertiary veins obscure. Staminate ae in subspicate racemes or subcorymbose, the axes pubescent with few- anched hairs; bracts and bracteoles inconspicuous or absent; flowers 2-40 per Zen the perianth deeply but tardily 3(-4)-parted; anthers 2-6, usually free to the base, shorter or longer than the column. Pistillate flowers 2-5 per fascicle, the tepals partially connate; ovary l-carpellate with a subsessile obscurely 2-cleft stigma. Fruits 1-4 per inflorescence, the ligneous pericarp dehiscing longitudinally into 2 valves; seed ellipsoid to globose, the aril laciniate. A genus of six species, ranging from Costa Rica through Central America to Amazonian Peru and adjacent Brazil. (529) [Vor. 49 218 ANNALS OF THE MISSOURI BOTANICAL GARDEN p Petioles winged to the base, 6-14 mm. broad; secondary veins 20-35 on either side; staminate pedicels 5-10 mm. long L D. LATIALATA. Petioles narrowly if at all winged, 1-6 mm. broad; secondary veins 5-18 on either side; staminate pedicels 1-6 mm. long Leaves 10-28 cm. long, 4-16 cm. broad, the secondary, veins 8-18 on either side; staminate inflorescences subspicate, 3-15 mm. long, the perianth 3.0-4.5 mm. long; androecium 2.4-3.0 mm. long, the anthers de dorsally at the base 2. D. OTOBA. Leaves 5-15 cm. long, 2.5-6.0 cm. broad, the on veins 5-10 on séier? side; staminate inflorescences subcoryml ose, 1.0-2.5 cm. long, the perianth ong; androecium 1.7-2.1 mm. long, org SE eiert Be pee near r the middle 3. D. ACUMINATA. D P = > o l. DIALYANTHERA LATIALATA Pittier, in Contr. U. S. Nat. Herb. 20: 454, 1922. Dioecious trees to 20 m., the younger branchlets strigillose, glabrescent. Leaves rugulose, minutely strigillose below, elliptic or obovate-elliptic, apically cuspidate or short-acuminate, basally attenuate, 20-40 cm. long, 7-13 cm. broad; secondary veins 20-25 on either side, the tertiary veins obscure; petioles winged, 15-40 mm. long, 6-14 mm. broad. Staminate fascicles in subspicate racemes, the racemes 1-3 per axil, 10-16 cm. long; fascicles 10- to 40-flowered, the pedicels 5-10 mm. long; perianth tardily 3-parted, 2.5-3.5 mm. long; bracts deciduous; anthers 2-3, ca. 0.25 mm. long, free to their bases, the infra-antheral portion of the androecium ca. 1.5 mm. long. Fruits 2-few per inflorescence, the rugulose ligneous pericarp ultimately splitting longitudinally into 2 valves, compressed-subglobose, short-stipitate, ca. 25 mm. long, 18 mm. broad; seed compressed-subglobose, the aril subentire. Panama to Colombia, at very low elevations. CANAL ZONE: forest between Peluca Hydrographic Station and Quebrada Peluca, d» R. Boquerón, ca. 70 m., Steyermark & Allen 17227. coLön: along R. Paté, 10-100 m Pittier 4193. 2. DIALYANTHERA OTOBA (Humb. & Bonpl.) Warb. in Ber. Deutsch. Bot. Ges. 13. 89. 1895 Myristica otoba Humb. & Bonpl. in Willd. Sp. Pl. 4: 869. 1805. Dioecious trees to 30 m., the younger branchlets rugulose, strigillose, glabres- cent. Leaves verruculose, strigillose but glabrescent below, broadly elliptic to obovate, 10-28 cm. long, 4-16 cm. broad; secondary veins 8-18 on either side, the tertiary veins very obscure; petioles canaliculate, slightly winged, 15-40 mm. long, 1-6 mm. broad. Staminate fascicles in subspicate racemes, 1-3 racemes per axil, 3-16 cm. long; fascicles 2-12, 8- to 15-flowered; pedicels 1-6 mm. long, the fugaceous orbicular bracts 3-4 mm. long; perianth tardily 3-parted, 3.0-4.5 mm. long; anthers 0.5-0.8 mm. long, free to their bases or slightly connate dorsally, the infra-antheral portion of the androecium 1.5-2.5 mm. long. Pistillate flowers 2-5 per fascicle, the perianth 3-lobed nearly to the base, to 5 mm. long; ovary l-carpellate, with an obscurely 2-lobed stigma on a short curving style. Fruits 2-4 per inflorescence, the verrucose ligneous pericarp dehiscing longitudinally into 2 valves, subglobose to ellipsoid, 21-33 mm. long, 16-23 mm. broad; seed ellipsoid to subglobose, the aril laciniate. (530) 1962] FLORA OF PANAMA (Myristicaceae) Figure 169, Dialyanthera latialata (531) 219 [Vor. 49 220 ANNALS OF THE MISSOURI BOTANICAL GARDEN Costa Rica to Colombia, usually below 1000 m. BOCAS DEL TORO: vicinity of Guabito, Stern & Chambers 112; loc. indet., Von Wedel 440; region of ee Cooper & Slater 7. cHırıqui: Progreso, Cooper & Slater 257 & 451. Called saba, bogamani verde, roble, miguelario, “wine wood” and “white cedar” in Panama, fruta dorado and sebo in Costa Rica, and otoba in Colombia, this rather large tree has somewhat foul-smelling fruits reputedly used as is the nutmeg. Locally, the wood is utilized in carpentry. 3. DIALYANTHERA ACUMINATA Standl. in Field Mus. Publ. Bot. 4: 209. 1929. Dioecious trees to 20 m., the younger branchlets strigose, glabrescent. Leaves verrucose, strigillose and glabrescent below, ovate to elliptic, apically acuminate, basally acute or attenuate, 5-15 cm. long, 2.5-6.0 cm. broad; secondary veins 5-10 on either side, the tertiary veins very obscure; petioles canaliculate, 8-20 mm. long, 1-2 mm. broad. Staminate fascicles subcorymbosely disposed, 1-3 “corymbs” per axil, 1.0-2.5 cm. long; fascicles 1-3, 3- to 10-flowered, the pedicels 1-4 mm. long; perianth 1.5-3.0 mm. long, tardily 3- to 4-parted, the bracts to 1 mm. long; anthers 0.25-0.5 mm. long, attached dorsally near the middle, the infra-antheral portion of the androecium ca. 1.5 mm. long. Pistillate and fruiting inflorescences not seen. Known only from Panama where it is called saba. ono: Daytonia Farm, region of Almirante, Cooper 395. cock: vicinity of El Valle « Pe hates ca. 600 m., Allen 2004; region north oF El Valle de Antón, 1000 m Allen 3645. 4. VIROLA Aubl. VıroLA Aubl. Pl. Guian. Fr. 2: 904. 1775. Sebophora Neck. Elem. 2: 188. 1790. Dioecious shrubs or trees, the younger portions usually pubescent with stellate hairs, the sap often reddish or brownish. Leaves alternate, glabrous above, stellate- pubescent and often glabrescent below, submembranaceous to coriaceous, petiolate, exstipulate, the tertiary veins obscure. Flowers in fascicles, racemes or panicles, the axes usually stellate-pubescent; bracts membranaceous, deciduous; bracteoles absent. Staminate flowers in fascicles of 3-15, the perianth deeply or shallowly 3- to 4- parted; anthers 2-6, dorsally connate, at least basally, longer or shorter than the column. Pistillate flowers solitary or in fascicles of 2-7, the tepals partially connate. Fruits 1-40 per inflorescence, the ligneous pericarp dehiscing longitudinally into 2 valves; seed globose to ellipsoid, the aril laciniate. The largest American genus, this has 38 species concentrated largely in the Amazon basin. Five of the six Central American species occur in Panama, the sixth being thus far reported only from British Honduras. a. Hairs of lower leaf surfaces stalked, persistent. b ondary veins of leaves desidia n either side, averaging less than 1 per cm. wont midrib; stamin ie Déi much branched; mature fruits 10-30 2 inflorescenc gon ped carp 0.5-2.0 mm. thick 1. V. SEBIFERA. bb. Segen veins of leave 18.35 6 either side, averaging more that 1 per cm. along the midrib; shit te inflor cences l- or 2-branched; mature fruits L5 per inflorescence, the pericarp 2-3 mm. thick zy KOSCHNYI, (532) 1962] FLORA OF PANAMA (Myristicaceae) 221 aa. Hairs of the lower leaf surfaces sessile, usually evanescent. ages veins of leaves 25-30 on either side, ae 1.5-2.5 per cm. along the midrib; bracts of the staminate inflorescences 3-5 mm. long; fruits Sa -Stipitate, 3-8 per inflorescence, on weine 6-9 mm. long 3. V. NOBILIS. Q cc. Secondary veins of leaves 9-21 on either side, averaging 0.8-1.3 per cm. along the midrib; bracts of the staminate inflorescences 2-3 mm. long; fruits sessile, Pedicels of the staminate flowers 0.5-1.5 mm. long, subtended by a conspicuous discoid expansion of the ultimate peduncle; fruits 1-5 per inflorescence, more than 2 cm. long, the pedicels 5-10 lon 4. V. GUATEMALENSIS. p Pedicels of the staminate flowers 2-3 cm. pr the subtending peduncle not conspicuously flared in the form of a disk; fruits 5-40 per inflorescence, less than 2 cm. long, the le 2-5 mm. lon 5, a m V. ELONGATA. 1. ViROLA SEBIFERA Aubl. Pl. Guian. Fr. 2: 904. 1775. Myristica sebifera Sw. Prodr. 96. 1788. Myristica virola Raeusch. Nom. ed. 3. 292. : Myristica sebifera var. cordifolia A. DC. in DC. ie 14: 195. 1856. Myristica sebifera var. curvinervia A. DC. loc. cit. 1856. Myristica mocoa A. DC. loc. cit. 1856. Myristica panamensis Hemsl. Biol. Centr.-Am. Bot. 3: 67. 1882. Virola sebifera var. curvinervia Warb. in Nov. Act. Acad. Leop.-Carol. 68: 174. 1897. Virola venezuelensis Warb. loc. cit. 182. 1897. Virola mocoa (A. DC.) Warb. loc. cit., 183. 1897. Virola boliviensis Warb. loc. cit. 184. 1897. Virola panamensis (Hemsl.) Warb. loc. cit. 185. 1897. Virola peruviana var. tomentosa Warb. loc. cit. 189. 1897. Virola mycetis Pulle, in Rec. Trav. Bot. Néerl. 4: 125. 1907. in part. Virola warburgii Pittier, in Contr. U. S. Nat. Herb. 18: 143. 1916. Dioecious, often buttressed trees to 40 m., the younger branchlets persistently tomentose or glabrescent. Leaf blades glabrous above, with persistent, ochraceous stalked-stellate hairs below, coriaceous, oblong to elliptic-ovate or obovate, acute to acuminate, cordate, truncate or acute, 10-47 cm. long, 4-15 cm. broad; secondary veins 10-28 per side, averaging less than 1 per cm. along the midrib, the tertiary veins rather prominent below; petioles canaliculate, 8-25 mm. long, 2-5 mm. broad. Staminate flowers in much-branched panicles; pedicels 0-3 mm. long; bracts inconspicuous or absent; perianth tardily 3(-5)-lobed, 1.3-3.0 mm. long; anthers 3(-5), 0.7-1.5 mm. long, usually connate to the apex, the infra-antheral portion of the androecium 0.2-1.0 mm. long. Pistillate flowers solitary or clustered in racemes 3-7 cm. long, 2-7 cm. broad; pedicels 1-4 mm. long; tepals partially connate, with subpinnate ochraceous pubescence; ovary l-carpellate, with a sessile, obscurely 2-lobed stigma. Fruits 10-30 per inflorescence, the velutinous ligneous pericarp ultimately dehiscing longitudinally into 2 valves, subglobose to ellipsoid, 10-21 mm. long, 7-17 mm. broad; seed ellipsoid to subglobose, the aril laciniate. Nicaragua to southern Brazil, Bolivia and Peru, usually below 1500 m. (533) [Vor. 49 229 ANNALS OF THE MISSOURI BOTANICAL GARDEN BOCAS DEL TORO: region of Almirante, Cooper 648. CANAL zONE: Barro Colorado Island, Barbour Point to next point south, Bangham 493, Barro Colorado ee Dwyer S Barro Colorado viid. Ebinger 300, Kenoyer 361, 493; Barro Colorado Island, Gatún Lake, Standley 31287, 31440, 41091; road along R. Pina-R. Media divide, Johnston 1601, í „Stern mbe al pe 2; vicinity. of Lá Palma, 0-50 M. Pittier 6615. PANAMÁ: Cerro Azul, Dwyer 1383; Ke along R. Chagres Geen Alhajuela, Pittier 3505; Juan Diaz, Standley 30615; R. Tecumen, Standley 29373; R. Tapía, Standley 28249. Colloquial names reported in Panama are malagueta de montana, mancha, bogamani and fruta dorado. Stern & Chambers 194 and Kenoyer 361 approach V. guatemalensis in that the subsessile stellate hairs are rather fugaceous, but A. C. Smith (in Brittonia 2:468. 1937) reveals that Central American specimens of V. sebifera deviate from Amazonian specimens in having larger fruits and more pronounced tendencies towards glabrescence and acute rather than cordate leaf bases. Warburg (in Nov. Act. Acad. Leop.-Carol. 68: 171. 1897) explains that many of the economic uses attributed to this species may be due to confusion with other members of the family. 2. VIROLA KOSCHNYI Warb. in Fedde, Repert. Sp. Nov. 1: 71. 1905. Virola merendonis Pittier, in Contr. U. S. Nat. Herb. 20: 453. 1992. Dioecious trees to 40 m., the younger branchlets densely tomentose to hirsute with ochraceous, usually evanescent hairs. Leaves glabrescent above except for the midrib, with persistent, ochraceous, stalked-stellate hairs below, coriaceous, oblong to narrowly ovate or obovate, apically acuminate or attenuate, basally cordate to acute, 13-38 cm. long, 4-13 cm. broad; secondary veins 18-35 on either side, averag- ing more than 1 per cm. along the midrib, the tertiary veins rather prominent below; petioles canaliculate, 7-16 mm. long, 1.5-4.0 mm. broad. Staminate flowers in few- branched panicles, the pedicels 1-5 mm. long; bracts ca. 5 mm. long, deciduous; perianth tardily 3-parted, 1.5-3.0 mm. long; anthers 3, 0.6-0.9 mm. long, usually connate to the apex, the infra-antheral portion of the androecium 0.5-1.2 mm. long. Pistillate flowers in clusters of 3-7 in racemes to 5 cm. long; pedicels 0.5-3 mm. long; tepals connate, with simple or few-branched hairs, the l-carpellate ovary with a subsessile obscurely 2-cleft stigma. Fruits 1-5 per inflorescence, on pedicels 5-8 mm. long, the glabrescent ligneous pericarp ultimately dehiscing longitudinally into 2 valves, subglobose to ellipsoid, 18-30 mm. long, 16-21 mm. broad; seed ellipsoid to subglobose, the aril laciniate. Guatemala to Panama. {: Progreso, Cooper & Slater 175; Almirante region, Cooper & Slater 24. DARIEN: foothills c of Ch Sambü basin, southern Darién, 30-500 m., Pittier The assignment of Pittier 5623 to this species is somewhat FONS The cuneate leaf bases, with the secondary veins somewhat distantly spaced, suggest a tendency to intergrade with Central American variants of Virola sebifera. The specimen cer- tainly departs from typical Virola koschnyi and probably merits varietal status, but the thick pericarp of the fruits tends to corroborate its determination as Virola koschnyi. (534) 1962] FLORA OF PANAMA (Myristicaceae) 223 , X ap EP MU LY Sz e JL EN, ei d d Figure 170. Virola sebifera (535) [Vor. 49 224 ANNALS OF THE MISSOURI BOTANICAL GARDEN 3. VIROLA NOBILIS A. C. Smith, in Brittonia 2: 490. 1937. Dioecious trees to 65 m., the younger branchlets sparsely puberulent with canescent hairs, glabrescent. Leaves glabrous above, with fugaceous canescent sessile- stellate hairs below, coriaceous, narrowly oblong, apically short-acuminate, basally acute to attenuate, 9-16 cm. long, 2.3-4 cm. broad, the secondary veins 25-30 per side, the tertiary veins rather obscure; petioles canaliculate, puberulent, 5-10 mm. ong, 1-2 mm. broad. Staminate fascicles in panicles up to 4 cm. long, the bracts deciduous, 3-5 mm. long; anthers 3, ca. 0.8 mm. long, connate to apex, the infra- antheral portion of the androecium 0.1-0.3 mm. long. Pistillate flowers in clusters of 3-several in racemes up to 5 cm. long, the connate tepals densely puberulent with much-branched hairs; ovary l-carpellate, with a subsessile obscurely 2-cleft stigma. Mature fruits 3-8 per inflorescence; pedicels 6-9 mm. long; pericarp densely puberulent, ultimately dehiscing longitudinally into 2 valves, short-stipitate, ellip- soid, 19-32 mm. long, 14-22 mm. broad; aril laciniate, the seed ellipsoid. Known only from the Canal Zone and adjacent Panama. NE: back of clearing at laboratory, Barro .. Island, Wetmore, Abbe & ck 1 155: hills north of Frijoles, Standley 27507; Barro Colorado Island Aviles 29; clearing, Barro Colorado iud Shatt uck 694; Barro Calera Island, Carpenter 43. PANAMA: Cerro Azul, Dwyer 1530. 4. VIROLA GUATEMALENSIs (Hemsl.) Warb. in Nov. Act. Acad. Leop.-Carol. 68: 220. 1897. Myristica guatemalensis Hemsl. Biol. Centr.-Am. Bot. 3: 66. 1882. Virola laevigata Standl. in Field Mus. Publ. Bot. 4: 209. 1929. Dioecious trees to 30 m., the younger branchlets puberulent with evanescent ferruginous or cinereous hairs. Leaves glabrous above, sparsely puberulent with fugaceous sessile-stellate hairs below, coriaceous, oblong to narrowly obovate, apically acuminate or cuspidate, basally attenuate to obtuse, 13-25 cm. long, 4-8 cm. broad; secondary veins 14-21 per side, the tertiary veins rather obscure; petioles canaliculate, 5-14 mm. long, 1-3 mm. broad. Staminate fascicles in many-flowered panicles 5-12 cm. long and nearly as broad; pedicels 0.5-1.5 mm. long, subtended by a conspicuous discoid expansion of the peduncle; bracts deciduous, 2-3 mm. long; perianth tardily 3-parted, ca. 2 mm. long; anthers 3, 0.6-0.8 mm. long, connate to apex, the infra-antheral portion of the androecium 0.4-0.7 mm. long. Pistillate flowers in few-flowered racemes 5-9 cm. long; tepals densely tomentose with fer- ruginous stellate or branched, fugaceous hairs; ovary l-carpellate with a subsessile obscurely 2-cleft stigma. Fruits 1-5 per inflorescence; pedicels 5-10 mm. long, the glabrescent, ligneous pericarp ultimately dehiscing longitudinally into 2 valves, ovoid-ellipsoid, 27-36 mm. long, 21-27 mm. broad; aril deeply laciniate, the seed ellipsoid. Guatemala to western Panama. CHIRIQUÍ: Progreso, Cooper & Slater 308. Standley and Steyermark (in Field Mus. Publ. Bot. 244: 298. 1946) report several uses for this species in Guatemala: the seeds are employed in flavoring (536) 1962] FLORA OF PANAMA (Monimiaceae) 225 beverages, oil expressed from the seeds is utilized in candle-making, and branches with the characteristic whorl of branchlets are used as egg beaters. A. C. Smith (in Brittonia 2: 499. 1937) lists cacao volador, quieche, sangre fruta dorado and boga- mani as common names for this Central American species. 5. VIROLA ELONGATA (Benth.) Warb. in Ber. Deutsch. Bot. Ges. 13:89. 1895, hyponym; in Nov. Act. Acad. Leop.-Carol. 68: 178. 1897. Myristica elongata Benth. in Hooker’s neg Bot. & Kew Mise, 5: 5. 1853. Myristica punctata Benth. loc. cit. 6. Myristica theiodora Benth, loc. cit. To Myristica cuspidata var. rufula A. DC. in DC. Prodr. 14: 696. Virola elongata var. punctata (Benth.) Warb. in Nov. Act ae FR -Carol. 68: 179. 1897. Virola elongata var. longicuspis Warb. loc. cit. Virola elongata var. subcordata Warb. loc. cit. 180. 1897. Virola rufula Warb. loc. cit. 181. 1897. Virola theiodora (Benth.) Warb. loc. cit. 187. 1897. Dioecious shrubs or small trees to 25 m., the younger branchlets puberulent with ochraceous evanescent hairs. Leaves glabrous above, sparsely puberulent below with fugaceous sessile stellate hairs, subcoriaceous, apically acute to acumi- nate, basally obtuse to acute, 7-32 cm. long, 1.5-11.0 cm. broad; secondary veins 9-20 per side, the tertiary veins obscure; petioles canaliculate, 4-16 mm. long, 1.0-2.5 mm. broad. Staminate fascicles in broad many-flowered panicles 4-18 cm. long, 3-15 cm. broad; pedicels 2-3 mm. long, the subtending peduncle not conspicuously expanded; bracts deciduous, 2-3 mm. long; perianth tardily 3(-4) parted, 1.5-3.0 mm long; anthers 3(-6), 0.6-1.6 mm. long, connate to apex or distally divergent, the infra-antheral portion of the androecium 0.2-0.6 mm. long. Pistillate flowers few in racemes 4-7 cm. long; tepals densely tomentose with ochraceous, stellate fugaceous hairs; ovary l-carpellate with a subsessile, obscurely 2-cleft stigma. Fruits 5-40 per inflorescence; pedicels 2-5 mm. long, the glabrescent ligneous pericarp ultimately dehiscing longitudinally into 2 valves, ellipsoid or subglobose, 11-16 mm. long, 8-12 mm. broad; aril deeply laciniate, the seed ellipsoid. Panama to Amazonian Brazil and Peru. CANAL ZONE: area west of Limón Bay, Gatün Locks and Gatün Lake, Johnston 1796. The fruits of the Panama specimen are apparently malformed but the indu- ment and venation of the leaves compare favorably with more southern representa- tives of the species, formerly not known from Central America. MONIMIACEAE Bx J. A. DUKE Aromatic, often lemon-scented shrubs or small trees. Leaves mostly opposite, rarely ternate or subalternate, exstipulate, penninerved, entire or irregularly dentate, membranaceous to coriaceous, glabrous to densely pubescent with simple or stellate hairs. Flowers unisexual (in America), perigynous, mostly dioecious, actinomorphic or asymmetrical, in axillary dichotomous or trichotomous cymes or rarely solitary, (537) [Vor. 49 226 ANNALS OF THE MISSOURI BOTANICAL GARDEN the pedicellate hypanthia urceolate to campanulate. Staminate flowers with few to several tepals, these usually uniseriate, often reduced to a subentire annulus en- circling the hypanthium; stamens 2-many, the outer often broader and occasionally sterile, the anthers oblong and valvate, or hippocrepiform and longitudinally de- hiscent (in North America), the filaments occasionally with a pair of lateral appendages; pistillodes usually absent. Carpellate flowers usually fewer and larger than the staminate; tepals 4-several, occasionally caducous and leaving a repand discoid hypanthium exposing the several separate carpels, more frequently persistent at the summit of the fleshy hypanthium tube (valve) in which the carpels are permanently imbedded. Fruiting carpels several, l-seeded, free or imbedded in the hypanthium, the seeds mostly erect in the drupaceous or coriaceous carpels. Confined largely to the tropics of both hemispheres, this family of some 30 genera is represented in North America by only 2. In the Americas the family is of only casual economic importance, with the Brazilian Mollinedia schottiana (Spreng.) Perk. and the Chilean Peumus boldus Mol. finding a limited demand in the timber trade. Some aromatic species are used in making teas supposed to have medicinal virtues. Considerable phylogenetic significance has been attributed to the combination of magnoliaceous and lauraceous characters in the family, which, according to the anatomical findings of Garratt (in Trop. Woods 39: 18. 1934), is more nearly lauraceous. Specific determinations in the American representatives are very difficult and a revision of existing specific concepts will ultimately be necessary. If one adopts the narrow concept followed by Perkins (in Engl. Pflanzenreich 4. Fam. 101. 1901), he will discover still many undescribed species in the Americas. Too often Perkins’ key characters are based on indumental and numerical variations. More reliable characters should be incorporated into a much-needed reorganization of this puzzling family. a. Anther ers. valvately oS tepals of the pistillate flowers 4-several, usually persistent, the fru s enclosed i in the hypanthium; leaves gla aie or with a, or stellate hai . SIPARUNA. aa apices hippocrepiform, Geste dehiscent; tepals of the pistillate "^u n deciduous, de fruits not enclosed in the hypanthium; leaves glabrous or with simple hai 2. MOLLINEDIA. 1. SIPARUNA Aubl. SiPARUNA Aubl. Hist. Pl. Guian. Fr. 2: 864. 1775. Citrosma Ruiz & Pavón, Fl. Per. & Chil. Prodr. 134. 1794. Leonia Mutis, ex Kunth, Synops. Pl. 1:462. 1822. Conuleum A. Rich. in Act. Soc. Hist. Nat. Paris 1: 391. 1823. Citriosma 'Tul. Monogr. 311. 1855. Angelina Pohl, ex Tul. loc. cit. 363. 1855. Dioecious or monoecious aromatic shrubs or small trees. Leaves usually oppo- site, glabrous to densely hairy, the hairs simple or stellate, the blades membrana- (538) 1962] FLORA OF PANAMA (Monimiaceae) 227 ceous to pergameneous, entire to irregularly dentate. Flowers pedicellate to sub- sessile, usually in axillary dichotomous or trichotomous cymes, the hypanthium often surmounted by an annular fusion of the tepal bases. Staminate flowers 3-many, with 4-8 subequal exappendiculate tepals united below to form an annulus interior to which is often a flat or hemispherical, apically perforate velum; stamens l-many, scattered or in circular disposition, the filaments often dilated or condupli- cate, the anthers oblong, valvate. Carpellate flowers similar to the staminate but usually fewer and larger; annulus and velum often present; carpels 4-many, the styles elongate, usually exserted, often basally connate. Fruits drupaceous or coria- ceous, permanently enclosed in the accrescent hypanthium; seeds tuberculate, ulti- mately exposed by the irregular bursting of the aggregate fruit. A puzzling economically trivial genus of about 100 “species,” at least 8 of them found in Panama. Several others are to be found in northern Central America and Mexico, but far more occur in South America. a. Lower leaf surfaces, young twigs and outer surfaces of the tepals hirsute or densely tomentose with a matted indument; plants dioecious; stamens 4-31. b. Stamens 8-30; leaves tomentose with a velvety indument, mostly 8-16 mm. broad LS PAUCIFLORA. bb. Stamens 4-6; leaves hirsute or, if tomentose, less than 8 cm. broad: c. Leaves tomentose, equilateral, entire or minutely denticulate. 2. S. GRISEO-FLAVESCENS. TONDUZIANA. ce. oe hirsute, often falcate, denticulate, some of the teeth at least . long zen r leaf surfaces, young twigs and outer surfaces of the tepals glabrous or A Kee appressed stellate hairs; plants dioecious or monoecious; stamen 2-14: £5 KI d. Leaves pergameneous, entire; plants monoecious; stamens (8- "ies velum lacking, the tepals erect, usually obscure in fruit .................. 4. S. GUIANENSIS. Leaves membranaceous, if pe neous denticulate; plants dioecious; stamens 2-8; velum present, pus beh erect or reflexed, usually obvious in fruit: e. Stamens t 8; leaves denticulate or, if entire, mostly more than 8c dd. jen Live denticulate; outer circle of stamens separate or con- nivent: g. Outer circle of stamens connivent; leaves equilateral, 10-20 cm. lo 5. S. TETRACEROIDES. 2g. haer circle m stamens not connivent; leaves falcate, ostly 5-10 cm. lon 6. 5. If. mie Po outer u af i stamens not connivent....7. S. NICARAGUENSIS. S. DIANDRA. ee, Stamens 2; leaves entire, less than 8 cm. long .. l. Sıparuma PAUCIFLORA (Beurl.) A. DC. in DC. Prodr. 16*: 696. 1868. Citriosma pauciflora Beurl. in Vet. Akad. Handl. Stockh. 1854: 144. 1856. Siparuna cauliflora Hemsl. Biol. Centr.-Am. Bot. 3: 69. 1882. Dioecious aromatic shrubs or small trees to 10 m., often branching from the base, the ultimate branchlets terete to quadrangular, gray-tomentose, the hairs to (539) [Vor. 49 228 ANNALS OF THE MISSOURI BOTANICAL GARDEN 0.5 mm. long. Leaves opposite, membranaceous to subpergameneous, narrowly el- liptic to obovate, apically acute to acuminate, basally cuneate to rounded, coarsely serrate to almost entire, densely gray stellate-tomentose below, densely to sparsely so above, the hairs to 0.8 mm. long; blades (10-)15-40 cm. long, (6-)8-16 cm. broad, with 10-16 pairs of lateral veins, cin a petioles 1-6 cm. long. Staminate inflorescence of 5- to 15-flowered subsessile axillary cymes, often cauli- florous, gray to yellow stellate-pubescent throughout; peduncles, 1-6 mm. long; pedi- cels 2-10 mm. long; tepals connate, forming an undulate subentire annulus; velum conical, 4-5 mm. in diameter, the apical pore 1.0-1.5 mm. in diameter; stamens (8-) 12-30, the outermost filaments dilated, ovate to orbicular, ca. 1 mm. long, the valvate anthers minute, introrse, usually slightly exserted. Carpellate inflorescences similar to the staminate but the cymes usually sessile, the 3-15 pedicels 2-10 mm. long; tepals connate forming an undulate annulus, the velum ultimately con- cave, the many separate styles scarcely exserted through the orifice. Aggregate fruits ecallose, globose, capped by the persistent annulus, yellowish-green, drying black, 10-15 mm. in diameter, bursting irregularly exposing the pinkish interior and the several tuberculate seeds. Peru to Colombia and Costa Rica, mostly at low elevations; called limoncillo in Costa Rica. DEL TORO: vicinity of Chiriqui Lagoon, Von Wedel 162, 1968, 2085, 2165; Almi- rante Cooper & Slater 69; Daytonia Farm, region of Almirante, Cooper 422; Farm Six, Changuinola valley, Dunlap 489. CANAL ZONE: Moji inga Swamp near mouth of R. Chagres, below 1 m., Allen 908; Barro Colorado Island, Ebinger 62, Kenoyer 469, Standley 31297, 31301, 31428, 40854, Wilson 32; Chagres, Isthmus of Panama, Fendler 196; Gatun, E s. n.; area west of Bay, Gatün Locks and Gatün Lake, Johnston 1711; 1- from Gorgona above the iue 40-150 m axon 4742; vicinity of Frijoles, Piper 5283; near old Fort Lorenzo, mout . Chagres, Piper 5913; around Las Cruces, 20-100 m Pittier 2620; along Cano Quebrado, Pittier 6660. cocrí: north rim of El Valle de Anton, 600-1000 m., Allen 1636. corów: forests around Porto Bello, 5-100 m., Pittier 2430; between France Field, Canal Zone and Catival, Province of Colén, Standley 30300. varıkn: banks co a oF 2), Chepigana, 2000 ft, Terry & Terry 1474. PANAMÁ: Cerro Azul, Dwyer 1523; . Tecumen, Standley 26752, 29348; R. Tapía, unde 28265; Juan Tia. Standley 30588. 2. SIPARUNA GRISEO-FLAVESCENS Perk. in Engl. Bot. Jahrb. 38: 693. 1901. Dioecious shrubs 2-3 m. high, the ultimate branches terete and densely yel- lowish- or grayish-tomentose, the hairs ca. 0.5 mm. long. Leaves opposite, mem- branaceous, densely yellow stellate-tomentose below, scantily so above, the hairs ca. 0.5 mm. long; blades oblong to obovate, apically acuminate, basally rounded to subcordate, entire or minutely denticulate, 10-17 cm. long, 4-8 cm. broad, with 9-11 pairs of secondary veins, the canaliculate petioles 1-4 cm. long, densely yellow-tomentose. Staminate inflorescences of 2- to 8-flowered axillary corymbi- form cymes, yellow-tomentose except for the velum; peduncles ca. 5 mm. long, the pedicels ca. 1 mm. long; tepals 4, minute, rounded, glabrous within; velum more or less flat, 3-5 mm. in diameter, the aperture ca. 1 mm. in diameter; stamens 6, exserted, the outermost much broader than the innermost. Carpellate in- (540) 1962] FLORA OF PANAMA (Monimiaceae) 229 florescences of usually 2 2- to 4-flowered cymes in each axil, densely yellow- tomentose except for the velum, the peduncles 2-4 mm. long, the pedicels 2-3 mm. long; tepals connate forming an entire or undulate repand annulus; velum convex, ca. 3 mm. in diameter, the aperture ca. 1 mm. in diameter, the 12-18 styles clearly exserted and tending to be connate at the aperture. Fruits not seen (“red” fide Standley, in Field Mus. Publ. Bot. 182: 449. 1937.) Costa Rica to Panama. CHIRIQUI: between Hato del Jobo and Cerro Vaca, eastern Chiriqui, 700-1000 m., Pittier 5404. The specimen cited, determined by Standley as S. griseo-flavescens, is carpel- late, but agrees well with Perkins' description of the species, based on a staminate plant (Pittier 1879 from Costa Rica). Siparuna patelliformis Perk. would seem to be very closely allied indeed. 3. SIPARUNA TONDUZIANA Perk. in Engl. Bot. Jahrb. 31: 746. 1902. Dioecious aromatic shrubs or small trees to 8 m., the ultimate branches terete, reddish-brown, hirsute with yellow to orange, mostly simple hairs to 3 mm. long. Leaves opposite, membranaceous, densely hirsute, at least below, with mostly simple hairs to 3 mm. long; blades elliptic to oblong-obovate, mostly inequilateral, apically long-acuminate, basally cuneate to cordate, coarsely and irregularly denticulate, 8-28 mm. long, 4-10 mm. broad, with 10-15 pairs of secondary veins, the terete to canaliculate petioles 0.5-3 cm. long. Staminate inflorescences usually 2 in each axil, corymbiform, with 3-7 flowers, or subspiciform with 3-15 flowers, the branching often unequal, yellow-pubescent throughout except for the velum; peduncles 4-15 mm. long, the pedicels 0-6 mm. long; tepals mostly 5-6, minute, ultimately forming a repand undulate annulus, the velum convex, 3-4 mm. in diameter, the aperture 1-1.5 mm. in diameter; stamens mostly 6, separate, the outer 4 broader and flatter, usually exserted, ca. 1 mm. long, the valvate anthers minute, introrse. Carpellate inflorescences like the staminate but the flowers usually fewer and slightly larger; styles 8-20, connivent near the apical aperture. Aggregate fruits callose, subreniform, capped by the minute lobate annulus, pinkish-red, ca. 10 mm. long and 15 mm. broad, bursting irregularly and exposing the pinkish interior and the tuberculate seeds. Panama to Guatemala, where it is called colloquially salvia and cerbatanero. Toro: Cricamola Valley, region of Almirante, Cooper 194; Buena Vista Camp on 2 Trail, region of Almirante, alt. 1250 ft., Cooper 596; without locality, Von Wedel 1 4. SIPARUNA GUIANENSIS Aubl. Hist. Pl. Guian. Fr. 2: 865. 1775. Citrosma discolor Poepp. & Endl. Nov. Gen. & Sp. 2: 48. 1817. Citriosma guianensis (Aubl.) Tul. Monogr. 361. 1855. Cistriosma guianensis B nuda Tul. loc. cit. 362. 1855. Citriosma guianensis y divergentifolia Tul. loc. cit. 1855. (941) [Vor. 49 230 ANNALS OF THE MISSOURI BOTANICAL GARDEN Siparuna panamensis A. DC. in Journ. Bot. 3: 219. 1865. Siparuna discolor A. DC. in DC. Prodr. 16°: 656. 1868. Siparuna guianensis B glabrescens A. DC. loc. cit. 654. 1868. Siparuna guianensis y longifolia A. DC. loc. cit. 1868. Siparuna guianensis 8 divergentifolia A. DC. loc. cit. 1868. Siparuna foetida Barb. Rodr. in Vellosia 21: 68. 1891. Siparuna guianensis var. nitens O. Ktze. Rev. Gen. 3?: 276. 1898. Monoecious aromatic shrubs or trees to 20 m., the ultimate branches terete or slightly flattened, reddish to greenish-brown, tomentose or glabrescent. Leaves opposite, pergameneous, mostly appressed-strigillose and glabrescent above, stellate- pubescent and glabrescent below; blades oblong to elliptic, apically acute to long- acuminate, basally cuneate to truncate, entire, usually symmetrical, 5-25 cm. long, 3-10 cm. broad, with 7-11 lateral veins on each side, the petioles 5-10(-15) mm. long. Staminate inflorescences hermaphroditic or unisexual, of 2-4 3- to 21-flowered cymes in each axil, yellow stellate-pubescent throughout, often with one con- spicuous dichotomous branch to the peduncle, the unbranched portion to 15 mm. long, the pedicels to 3 mm. long; tepals 4-6, deltoid to somewhat hemispherical, the velum lacking; stamens (8-)10-14. Carpellate inflorescences similar to the male, the flowers with several long-exserted, basally connate styles, the velum, if present, obscured by the tepals, conical. Aggregate fruits callose, contorted, ob- scuring the tepals, pinkish, 8-14 mm. long, 8-14 mm. broad, bursting irregularly and exposing the yellowish interior with its few tuberculate grayish seeds. Peru and Brazil to Colombia and Costa Rica. ZONE: Barro Colorado Island, Aviles 16, ponies 457, Shattuck 793, 1074; Chagres, Re of Panama, Fendler 188; area w. of Limón Bay, Gattin Locks and Gatün Lake, road alon ng R. Pin a-R. Media divide, Johnston 1808; near Ft. Randolph, Maxon & n. en DARIÉN: er of La Palma, 0-50 m., Pittier 6699. HERRERA: Ocu, Ebinger 1091. PANAMÁ: La Cam Cerro Campana, Ebinger 336; R. Tapia, Standley 28093, 28296. vERAGUAs: San Re Dwyer 1280. Standley (in Contr. U. S. Nat. Herb. 27: 182. 1928) reports that in Panama, where it is called hierba de pasmo, the plant is supposed to be used as a remedy for colic and as a vermifuge on fowl. This is the only confirmedly monoecious species of the Monimiaceae in Central America, but, when the monoecious con- dition is not obvious, it is difficult to distinguish from S. nicaraguensis. 5. SIPARUNA TETRACEROIDES Perk. in Engl. Bot. Jahrb. 28: 689. 1901. Dioecious shrubs or small trees, the branchlets terete, reddish, with gray hairs to 0.4 mm. long. Leaves opposite, the blades membranaceous, narrowly elliptic to obovate, apically with a mucronate or emarginate acumen, basally cuneate, marginally undulate-denticulate, 8-20 cm. long, 5-8 cm. broad, glabrescent above, scantily stellate-pilose below, with 8-12 pairs of arcuately ascending veins, the petiole gray-pilose, 2-5 cm. long. Staminate inflorescences of few-flowered cymes mostly paired in the axils of the leaves, with gray stellate hairs, the peduncles (542) 1962] FLORA OF PANAMA (Monimiaceae) 231 DNA, A 77 \ A T N oe M : NN \\ \ Ai Ai 7 D: Y Z 74 ZA Nm NA : N JA A" N Ww ip S A N Ä WY S Figure 171. Siparuna tetraceroides 2-4 mm. long, the pedicels 1-3 mm. long; flowers rotate, ca. 2 mm. in diameter, the receptacle obconic, the tepals obsolescent, ultimately connate and repand, the velum hemispheric, the orifice ca. 1 mm. in diameter, stamens 5-8, the 4 exterior in a whorl and often connate or connivent, 1.5-2.2 mm. long, the anthers minute, introrse, exserted. Carpellate inflorescences similar to the staminate, perhaps fewer-flowered, the tepals obscure, deltoid, ultimately forming an undulate annulus, the styles more or less connivent at the aperture. Fruits callose, reniform, capped by the persistent tepals. Costa Rica to western Panama. v cumiQuí: between El Hato (Volcán) and Costa Rica, Ebinger 813; Puerto Remedios, 0-30 m., Pittier 3386; in forest, Cerro de la Plata, near San Félix, eastern Chiriquí, 12-150 m., Pittier 5156. The cited Panama specimens depart rather radically from Costa Rican ma- terial and might be deserving of some nomenclatural status. Panama material has more delicate, less coarsely toothed leaves, and the veins are not so con- spicuously yellow as those specimens from Costa Rica which I believe to be properly called S. tetraceroides (Pittier 16144, cited by Perkins, and Skutch 2220, 3813, 3985). The staminate Panama plant (Pittier 5156), illustrated in Fig. 171, has the outer whorl of stamens connate, and in this respect represents an approach to the Colombian S. steleandra Perk. (543) [Vor. 49 232 ANNALS OF THE MISSOURI BOTANICAL GARDEN 6. SIPARUNA RIPARIA (Tul.) A. DC. in DC. Prodr. 16?: 647. 1868. Citriosma riparia Tul. in Ann. Sci. Nat. 4%: 36. 1855. Siparuna riparia var. macrophylla Perk. in Engl. Bot. Jahrb. 28: 690. 1901. Siparuna riparia var. grandiflora Perk. in Engl. Pflanzenr. 4. Fam. 101: 99. 1901. Siparuna riparia var. calantha Perk. in Notizbl. Bot. Gart. Berl. 10: 164. 1927. Dioecious shrubs or small trees to 6 m., the ultimate branches terete or quadrangular, reddish-brown or gray, tomentose with yellowish or grayish stellate hairs to 1 mm. long. Leaves opposite, membranaceous, pilose, especially below, with stellate hairs to 1 mm. long; blades elliptic to obovate, often inequilateral, apically acute to caudate-acuminate, basally cuneate to truncate, serrate, 5-12 mm. long, 3.5-6.0 cm. broad, with 8-10 lateral veins on either side, the subterete to canaliculate petioles 1.0-3.5 cm. long. Staminate inflorescences of 2-3 3- to 15- flowered cymes in each axil, grayish to yellowish stellate-pubescent except for the velum; peduncles mostly 4-10 mm. long before branching, the pedicels 3-12 mm. long; tepals 4-7, at first triangular, ultimately rounded and somewhat reflexed; stamens mostly 5-6, the outer 4 disposed in a circle but not connate; velum ca. 2 mm. in diameter, the orifice ca. 1 mm. in diameter. Carpellate inflorescences similar to the staminate, the styles 6-12, connate near the orifice, long-exserted. Aggregate fruits callose, reniform, reddish, ca. 10 mm. broad, 10-15 mm. broad. Mexico to Colombia. cHIRIQUÍ: rain forest, Bajo Chorro, 6000 ft., Davidson 242; vicinity of Bajo Mona and Quebrada Chiquero, 1500 m., Woodson & Schery 600. The identity of the cited specimens is not without question. Although Perkins has said that the stamens are 10-12 in S. riparia, Bourgeau 1748, identified by Per- kins as S. riparia, has 5-6 stamens, and I believe the latter numbers to be more prevalent. T. SIPARUNA NICARAGUENSIS Hemsl. Biol. Centr.-Am. Bot. 3: 69. 1882. Dioecious aromatic shrubs or small trees to 8 m., the ultimate branches terete, ferruginous stellate-pubescent, the hairs evanescent. Leaves opposite, membrana- ceous, glabrous or subglabrous above, scantily stellate-pubescent to subglabrous elow; blades elliptic to obovate, apically more or less acuminate, marginally entire to undulate or minutely denticulate, 6-22 cm. long, 3.5-12.0 cm. broad, with 5-10 lateral veins on either side, the canaliculate petioles 1-9 cm. long. Staminate inflorescences with 1-4 1- to many-flowered cymes in each axil, grayish- to yellow- ish-puberulent except for the inner portion of the tepals and the velum; peduncles 1-15 mm. long, the pedicels 1-5 mm. long; tepals (3-)4-7, basally connate, the lobes free, rounded or deltoid; velum 2-4 mm. broad, the pore ca. 1 mm. in diameter; stamens 5-8, the outer 4 or 5 separate, occasionally disposed in a circle embracing the inner 1-3 which are often conduplicate, ca. 1 mm. long; anthers valvate, minute, introrse. Carpellate inflorescences similar to the staminate but with fewer flowers, only 1 in a given cyme usually developing into fruit, the numerous styles exserted ca. 0.5 mm. Aggregate fruits smooth or slightly callose, capped by the (544) 1962] FLORA OF PANAMA (Monimiaceae) \ n m r4 y 2 N. d fer wid 1 e x 8 AN Laer NW / x : A ` " tad) Een y e Ze EM pom em N VAN i NN ^ d a d ` A N hm, M C GC PIR N W ch i , 7h Ws. WM P Z AZ “ry Ta lta, v Z o d 4 TA 7A KA WW ` A YG NOM ‘ 1 , G GR ZZ g KH Torre 7 / RL 2 pan Figure 172. Siparuna nicaraguensis persistent erect tepals, orange to red, 10-20 mm. in diameter, bursting irregularly to expose the several tuberculate seeds. Panama to Mexico. TORO: region of ee zer 616. cocL&: north rim of EI Valle, Allen 1841; north r rim is El Valle de a on & Scher 159. can con Hill, D \ SCH a de la Gloria, near Faté (Nombre ge Dios), 10-104 m., Pittier 4077. DARIÉN: along the Sam bon River, southern Darién, above tide limit, Pittier 5553; vicinity S Cana, alt. 1750 ft., Stern, Chambers et al 468, 6. Paca, below Cana, Williams 718. panamá: La Campana, Cerro Campana, Ebinger 333. The aromatic leaves of this species, known in various places in Central America as cerbatana, chuché, hormiguillo, kex, limoncillo, salvia, “wild coffee and yaya de mono, are locally brewed to form a tea supposedly remedial for respiratory ailments. Material from Darién, as illustrated in Fig. 172, departs rather disconcertingly from most Central American specimens referred to this species. Both Standley and I have suffered the temptation to treat such Darién material as a new taxon differing from typical S. nicaraguensis in having the shorter male tepals subdeltoid and erect at anthesis and uncommonly large leaves and petioles. Such variation is more common to the south but may be found also in specimens from Mexico and northern Central America. It is for that reason that I am reluctant to recognize the variants formally. (945) 233 [Vor. 49 234 ANNALS OF THE MISSOURI BOTANICAL GARDEN Figure 173. Siparuna diandra 8. SrparuNA diandra J. Duke sp. nov. Frutices dioeci 2-4 m. alti ramulis teretibus ferrugineis griseo-pilosiusculis pilis glandulosis ad 0.3 mm. longis. Folia plerumque opposita lamina membranacea obovata vel subelliptica apice caudate acuminata basi attenuata cuneatave utrinque parce stellato-pilosiuscula integra nervis lateralibus plerumque 6-8 arcuate ascen- dentibus in utroque latere petiolo griseo-pilosiusculo 5-10 mm. longo. Inflorescentiae masculae pauciflorigerae corymbosae in axillis foliorum partibus exterioribus parce grisiter stellateque pilosiusculis pilis ad 0.3 mm. longis pedunculo plerumque 1.0-1.5 mm. longo pedicellis 1.0-3.5 mm. longis. Flores rotati receptaculo obconico ca. 1.0-1.5 mm. in diam. tepalis plerumque 4 triangularibus erectis nigrescentibus ca. 0.5 mm. longis intus glabris; velo tholiforme ore parvissimo staminibus 2 carinatis cucullatisque carnosis glandulosis ad 1.5 mm. longis vix exsertis. Flores feminei nunc ignoti. N: Cana-Cuasi Trail (Camp 2), Chepigana, 2000 ft., Mar. 11, 1940, Terry & Terry 1496. SE MO). (546) 1962] FLORA OF PANAMA (Monimiaceae) 235 An interesting addition to the flora of Panama, this peculiar species seems to have no close allies in Central America. Among South American species, it would seem to be most closely related to the Peruvian S. gilgiana Perk., which differs in having much larger leaves and petioles and the tepals connate into a repand flange, and the Brazilian S. brasiliensis Perk., which also has larger leaves, petioles and peduncles. 2. MOLLINEDIA Ruiz & Pavén MorriNEDIA Ruiz & Pavón, Fl. Peruv. & Chil. Prodr. 83, t. 15. 1794. Tetratome Poepp. & Endl. Nov. Gen. & Sp. 2: 46, t. 163. 1838. Paracelsia Mart. ex Tul. in Mart. Fl. Bras. 4!: 314. 1857. Dioecious shrubs or small trees. Leaves opposite, glabrous or slightly pilose below, membranaceous to coriaceous, petiolate, exstipulate, entire or irregularly dentate, with few secondary veins confluent at some distance from the margin. Flowers pedicellate, solitary, cymose or corymbiform in the axils, the hypanthium campanulate or urceolate, glabrous or pubescent, the hairs simple. Staminate flowers 3-many per inflorescence, with 4 tepals, the 2 inner frequently smaller and provided with a terminal laciniate appendage; stamens 8-many, included on the inner surface of the hypanthium, the filaments minute, the anthers (in Central America) hippocrepiform, longitudinally dehiscent. Carpellate flowers mostly 1-3 per in- florescence, the 4 petals soon deciduous from the urceolate hypanthium; carpels 6-many, glabrous to pilose, ellipsoid, the styles filiform. Drupes fleshy or coriaceous, sessile or stipitate on the reflexed undulate hypanthium. With the exception of 1 dubious Australian species, all of the nearly 75 species of this genus are found in tropical and subtropical America. Economically unimportant, the genus is taxonomically perplexing and little can be done to correct this until a monographic study is undertaken. Four of about 15 Central American species occur in Panama. a. Leaves ovate to obovate, basally rounded to cuneate, weakly toothed; fruits drying black or brown, scarcely stipitate: Drupes smooth, glabrous, drying black; leaves glabrous, EEN branch- lets glabrous M. DARIENENSIS. bb. Drupes corrugated, often — drying brown; leaves rentas rarely glabrescent, obtuse to shortly acuminate; branchlets pubescent 2. M. COSTARICENSIS. aa. Leaves lanceolate to thomb-elliptic, basally m. often strongly toothed; fruits drying brown, occasionally long-stipita c. read 10-13 mm. long, sessile or obscurely stipitate; leaves membrana- ceous, usually puberulent; plants dioec 3. M. PINCHOTIANA. ec. Drupes 15-20 mm. long, on stipes l- $ mm. long; leaves pergameneous, glabrous; plant possibly monoecious 4. M. STIPITATA. l. MOLLINEDIA DARIENENSIS Standl. in Field Mus. Publ. Bot. 22: 332. 1940. Dioecious shrubs 1.0-2.5 m. high, the branches glabrous and rather dark in drying. Leaves chartaceous, glabrescent, lance-ovate, apically acuminate, basally cuneate to sub-rounded, 9-16 cm. long, 3-6 cm. broad, with 5-9 irregular teeth on (947) [Vor. 49 236 ANNALS OF THE MISSOURI BOTANICAL GARDEN either side, the secondary veins 4-5 on either side, arcuately anastomosing 2-5 mm. from the margin; petioles ca. 10 mm. long, drying dark brown, glabrous or glabrate. Staminate flowers not known. Carpellate flowers solitary or geminate in the axils of the leaves, the pedicels glabrous or glabrate, ca. 10 mm. long, the peduncles 1-5 mm. long; bracts and bracteoles not apparent; flowers at anthesis not known. Fruits fleshy ellipsoid, rounded at both ends, smooth, drying black, 13-15 mm. long, ca. 10 mm. broad; receptacles reflexed, ca. 10 mm. broad; seeds ca. 12 mm. long, 9 mm. in diameter, the testa mottled. Panama, known only from the type collection. DARIEN: Cana-Cuasi Trail (Camp 2), Chepigana District, 3000 ft., Terry & Terry 1456. 2. MOLLINEDIA COSTARICENSIS Donn. Sm. in Bot. Gaz. 33: 257. 1902. Dioecious shrubs or small trees to 7 m. high, the branchlets pubescent with grayish hairs. Leaves pergameneous, appressed-strigillose, especially below, rarely glabrescent, narrowly to broadly ovate or obovate, apically obtuse to shortly acumi- nate, basally acute to rounded, 8-18 cm. long, 4-9 cm. broad, subentire or with up to 25 minute irregular teeth on either side, the secondary veins 5-11 on either side, arcuately anastomosing 3-8 mm. from the margin; petioles 5-16 mm. long, drying rufous, puberulent. Staminate inflorescences 1-6 in 3- to 5-flowered cymes in each axil, the bracteate peduncles and pedicels with yellowish puberulence; peduncles 1-20 mm. long, the pedicels 2-5 mm. long. Flowers campanulate, 6-8 mm. in diameter at anthesis; tepals 4, exteriorly with a yellowish puberulence, the inner 2 with an apical laciniate appendage; stamens 16-34, ca. 1 mm. long, the anthers hippocrepiform. Carpellate flowers mostly solitary or paired in the axils, the bracteate peduncles 5-20 mm. long, the bracteolate pedicels to as much as 6 cm. long; flowers turbinate, 2-4 mm. in diameter, the tepals ca. 2 mm. long, caducous, the numerous carpels ellipsoid, with a grayish or yellowish pubescence. Fruiting receptacles discoid, repand, densely strigillose on both surfaces, 10-15 cm. broad; fruits ellipsoid, numerous, brown, corrugated, usually puberulent, 8-13 mm. long, -8 mm. broad, the seeds slightly smaller. Panama and Costa Rica. CHIRIQUI: rain forest, Bajo Chorro, Boquete Distr., 6000 ft., Davidson 370. M. costaricensis is not constantly separated from M. pinchotiana, but a sum- mation of the following characters seems to define the former rather well: leaves pergameneous, with the lateral veins prominent below, the base rounded, the apex rounded, acute or briefly and abruptly acuminate, the indument yellowish or grayish, the staminate flowers 6-8 mm. in diameter at anthesis, the stamens 16-34. 3. MOLLINEDIA PINCHOTIANA Perk. in Engl. Bot. Jahrb. 31: 743. 1902. Dioecious shrubs or small trees to 6 m. high, the branchlets often densely puberulent with evanescent brownish hairs. Leaves membranaceous, glabrous or lightly puberulent, narrowly elliptic to obovate, apically acuminate, basally cuneate, with 5-12 irregular arcuate teeth on either side, 5-16 cm. long, 2-6 cm. broad, with (548) 1962] FLORA OF PANAMA (Monimiaceae) 237 4-5 lateral veins on either side, these conspicuous to inconspicuous below, arcuately anastomosing 2-10 mm. from the margin; petioles 5-18 mm. long, light brown, puberulent or glabrate. Staminate inflorescences 1-4 in the axil of a leaf, usually of 3-flowered cymes, the bracteate peduncles and pedicels with a grayish or reddish- brown puberulence; peduncles 5-20 mm. long, the pedicels 3-30 mm. long. Flowers campanulate, 4-6 mm. in diameter at anthesis; tepals 4, with grayish or reddish- brown puberulence, the inner 2 with a laciniate apical appendage; stamens 23-36 (44 in some Costa Rican deviates with a pinkish puberulence on the leaves), ca. 1 mm. long, the anthers hippocrepiform. Carpellate flowers mostly solitary in the axils, the bracteate peduncles 3-20 mm. long, the bracteolate pedicels 3-30 mm. long; flowers turbinate, 2-3 mm. in diameter, the acute tepals ca. 2 mm. long, promptly deciduous, the numerous carpels ellipsoid, with a grayish puberulence. Fruiting receptacles discoid, reflexed, puberulent on both surfaces, ca. 8-12 mm. broad; fruits ellipsoid, numerous, brown, carnose, 10-12 mm. long, 6-7 mm. broad, the seeds slightly smaller, brown, mottled with purple. CHIRIQUI: humid forest between Alto de las Palmas and top of Cerro de la Horqueta, 2100-2268 m., Pittier 3246. Apparently Standley at one time considered this specimen to represent a new species. The leaves are rather large and basally rounded and the fruits are immature, but the plant appears to be M. pinchotiana, a variable species which grades into several “microspecies” in Costa Rica. At one extreme is a “microspecies,” appar- ently unnamed at present, from the region of Zarcero. This is characterized by narrow rhomb-elliptic leaves with a pinkish indument and the stamens numbering about 40. At the other extreme is M. minutiflora Standl. & Williams, with multi- florous male inflorescences, the flowers 1.5-2.5 mm. in diameter at anthesis, the stamens 10-16, the leaves glabrous, elliptic and with few or no teeth. Closely allied to M. mexicana Perk. (scarcely distinct from M. nigrescens Tul.) is a group of specimens from the vicinity of El General, Costa Rica. This group, to which Standley also applied an herbarium name, is characterized by lanceolate leaves with 2-6 teeth on either side, multiflorous staminate inflorescences, the flowers ca. 3 mm. in diameter, the stamens 16-24 in number. Whether all these should be treated as separate species or merely as components of one polymorphic species is a serious question for the monographer. 4. MoLLINEDIA stipitata J. Duke, sp. nov. Frutices dioeci vel interdum fortasse monoeci ad 3 m. alti ramis griseis subteretibus ramulis glabrescentibus griseis vel rufis. Folia opposita lamina glabra anguste lateve elliptica apice acuta vel acuminata basi cuneata vel acuta in utroque margine denticulata dentibus aristatis 1-5 irregulariter remotis 6-11 cm. longa 15-45 mm. lata utrinque glabra nervis prominentibus lateralibus plerumque 5 in utroque latere prope marginem confluentibus petiolo glabro canaliculato 5-10 mm. longo. Inflorescentiae masculae plerumque 3- vel 5-florigerae binae in axillis foliorum pedunculis bracteatis 10-14 mm. longis pedicellis ca. 5 mm. longis. Flores rotati ca. (549) [Vor. 49 238 ANNALS OF THE MISSOURI BOTANICAL GARDEN Figure 174. Mollinedia stipitata 2 mm. in diam. tepalis 4 glabris 2 interioribus laciniose appendiculatis staminibus ca. 26 1.0-1.5 mm. longis antheris hippocrepiformibus. Inflorescentiae femineae plerumque uniflorae solitariae vel binae in axillis foliorum pedunculis bracteatis ca. 8 mm. longis pedicellis bracteolatis ca. 10 mm. longis. Flores ca. 4 mm. in diam. glabri tepalis 4 ca. 2 mm. longis mox deciduis carpellis ca. 20 separatis stylis longis. Receptacula fructifera disciformia repanda 7-15 mm. lata pedunculo cum pedicello 10-45 mm. longo drupis stipitatis fuscis striatis ellipsoideis mucronatis 16-20 mm. longis ca. 10 mm. latis basem attenuatis. cHIRIQUÍ: between the R. Ladrillo and Los Siguas, southern slope of Cerro de la Horqueta, 1200-1700 m., Pittier 3280 (soLorype, NY); cloud forest, Cerro Horqueta, 6500 ft., Von Hagen & Von Hagen 2064; humid forest between Alto de las Palmas and top of Cerro de la Horqueta, 2100-2268 m., Pittier 3262. Among North American species this seems most closely related to a specimen from Chiapas (Matuda 442) determined by Standley as M. viridiflora 'Tul. Stand- ley's determination is doubtful, however, because a photograph of the type of M. viridiflora alarmingly resembles what is currently passing as M. guatemalensis Perk., a species with large pergameneous leaves, the staminate flowers 5-8 mm. in diameter at anthesis, with 30-48 stamens and with large short-stipitate drupes. Although Perkins has stated that the stamens in M. viridiflora are *usque 24," this seems to (550) 1962] FLORA OF PANAMA (Monimiaceae) 239 be the only character separating it from M. guatemalensis. The new species is readily distinguished by its small narrow leaves, the small male flowers, and the long-stipitate, apically mucronate, basally attenuate drupes. The Von Hagen specimen poses an interesting problem as it consists of both staminate and carpellate branches. Their data say “tree, 8 ft tall.” By the singular usage of tree, we would forced to conclude that the species is sometimes monoecious, but until the physical connection of male and female flowers is verified, this is pure speculation. There are no reports of monoecism in Mollinedia, but the condition is not uncommon in Siparuna. (551) INDEX TO PART IV (PIPERACEAE - MONIMIACEAE) bers represent italicized page m at bottom of pages. Synonyms in italics; Num new species or combinations in bold-face Abanico, 354 Abrojo, Acanthinophyllum, 163 Acetosella acetosella, 30 Achyranthes, 364; altissima, 361; arge > Ed muris 366, var. indica, 366, bosa, 4: halimifolia, corym 384 hilos 369; indica, 366; laguroides, 386; m , 376; lehmannii, 387; linkiana, 363; ana, 387; mucronata, 383; ob- eben. "366: anamensi. 7; philoxeroi- des, 385; ren. 382; prostrata, 67; radicans, 383; ens, cula, 366 stenophylla, 386: vermicularis, E wil- iamsii, 3 Acrolobus, 270 Ac rocarpidium Guildingianum, 106; hispidu- lum, 111; reueg 96; pulicare, 106; ene 106; scandens, 106; Sellowi- oe 1 um, Afer m, 433 enogramma oppositifolia, 381 Adenonema, 439 Adicea, Adorno de nifio, 377 Agathophytum, 344 Agonandra, 273; brasiliensis, 274 acatia, Alfaroa, 130; costaricensis, 132; manningii, 132 Alicastru Allaganthera forskalii, 381 chlam v we a O inata var. ferruginea, 134; ferruginea, 134: goa eat var, ferruginea, 1 Alsinanthe, 433 Alsinanthus, 433 Alsine, 432 Alsineite 433, 439 Alsinopsis, 433 ternanthera, 380; a 383; am 383; asterotricha, 384; ttzickianna, 383. SC 384; denticulata, on ficoidea, 383; ficoidea, 382, var. flavogrisea, 384, var. linearis, 375; mexicana, 387; nodif ora, 381; panamensis, 387; paniculata, 376; paronchyoides, 282; Mb ed 383; philox- eroides, olyg 382 poly- gonoides, 383, var. Pod ens, 383; re pe ns, GEN sessilis, Sieg stenophylla, 386; tenuis- ; triandra, 381; versicolor, 383; villiflora, 383, "williamsii 385 iini unas i 348 Am EN us, 354; ee Sa eh 356; EEN Sege 356; ch "358, v var. KEE 2207 cruentus, 357; diacanthus, 360 dubius, 358, var. flexuosus, 358, var. leptostachys, 358 stach paniculatus, 357; ege ndriacus, 358; in- comptus, 358; laetus, 358; paniculatus, 357, var. cruentus, 357, var. purpurascens, 357; parisiensis, 357; purpureus, 354; retroflexus 358; var. leptostachys, 358, var. xantho- sins 358; viridis, 356 Amar FR 389; globosus, 391, var. albi- florus, 391 Par 211 Ambrina, 344: ambrosioides, 347; andicola, 347; anthelmintica, 347; chilensis, 347; rieng 347 Ambu Ammodeni, 432 Amor 4, 2:35 don 152; hottlei, 152 : Anaxagorea, 506; allenii, 506; panamensis, 506 Ancistrocarpus, 415 Ancistrostigma, 424 Anderssoniopiper, 75; panamense, 75 Andriapetalum, ph see S 242; suaveolens, 242 ag na Bn acuminata, 521; cherimolia, 491: glabra, 520; hayesii, 523; gent ; purpurea, i IT 491; 21; squamosa, 491 SE 91 (552) 1962] FLORA OF PANAMA—INDEX 941 Anomospermum, 475; reticulatum, 476 Anomu, 410 Anona, 220, 518; australis, 520; echinata, 521; involucrata, 520; laurifolia, 520; manirote, 520; palustris, 520; peruviana 520; pisonis, 520; prestoei, 520; uliginosa, 20; uncinata, Anonocarpus, 176; amazonicus, 176 343 Antidaphne, 248; amazonensis, 248; fendleri, 248; paraensis, 248; viscoidea, Antigona, Antigonon, m d 317, 318; corda- (és 8; an tensis, 434, var. quitensis, n ite 434; 'polycaulos, 435; 435, cinerascens, 435, var. mearnsii, 435; MPa Aristolochia, 292; videi 298; appendi- 94 - 294: ferruginea, 300; foetens, 300; gemini- flora, 304; Le sa, 297; gigas, 300; grandi- flora, 300, hookeri, 300; haughtiana, 300; infit 997; loriflora, 298; macroura, SE dies teg a, 296; mathewsii, ; ; 3 d cordata, 304, var. geminiflora, over. axima, ST oam 296; Ser folia, 294; odoratissima, 296, var. grandi- flora, 296; panamensis, 295; pusilla 294; pyrinea, 294; reticulata, subclausa, 294; sylvicola, 303: eer Parte 295; tenera, 294; trifida, 295; triloba, 295; trilobata, 295: v veraguensis, 30 Aristolochiaceae, 291 Artanthe adunca, 31; aequalis, 47; alaris, 15; au, rev ta, ; aurita calocoma, 61; catalpaefolia, 17; caudata, 15; celtidifolia, 31; crassispicata vi en rens, 63; geniculata, 64; gran 68; Hartwegiana, 69; heterophylla, 9: Ce. 32; imperialis, 23; Lessertiana, 64; Lusch- nathiana, 64; macrophylla, 64; marginata, I . Bs re? Gaich 9; riparia, 68; s à 92; iana, 2 ptuplinervia, 46; trinervia, 65; tuberculata, 63; sa 64; xestophylla, 64; xylopodioides, 64 Asimina costaricensis, 51 N Atrag Atri Sta " ambrosioides, 347; anthelmintica, 347. muralis, 344 Aylmeria, 446 B Bajan, 354 Balanophoraceae, 285 Balanostreblus, 161 Ballarion, 439 Barba ei viejo, 459 Batidac Batis maritima, 08 Batocarpus, 174; costaricensis, 174, 176; orin- a, 461 ejuco Soe 321; Bejuco de crispillo, 459 Bellisima, 318 Geess 466 Berberis, 466; dealbata, 468; nigricans, 468 Bessera, Bigelovia, 433 Bledo, 356, 358; Bledo macho, 361 Bliton, 354 Blitum ambrosioides, 347 Blochmannia, 335 Bloodberry. Blulkfaren. 377; brevifolium, 378; repens, no 227; angustifolia, 236; aspera, 228; cuspidata, 229; cylindrica, 228; fallax var. Lage a, 228; Zeene 234; obliqua, 234; pavonii, 229; ramiflora var. cuspidata, 29; rugosa, eg ulmitolia, 228 Boerhaavia, ; adscen ; atomaria, coccinea, 395, L ere Deg 396; ixodes, 396; laxa, 396; paniculata, 396, var subacuta, bn? polymorpha, sedis nor 396; virgat Ga viscosa, 396, v icu- lata, 396, ena, diffusa, 396; elongata, 394; erecta, 394; hirsuta, 396; ramulosa, 396; squamata, 396; thornberi, 394 Déeg 534, 537; Bogamani verde, 532 Boheravia, 394 Boria, E Botón, 392 Botoncillo, 391 (553) 242 Botrydium, 344 Soe, 470; spruceanan, 470 Botrys, 344; ambrosioides, 347; anthelmintica, Brachystemma, 433 des antia, 389 andesia mexicana, 387 een 1 64 e panamense, ramonense, 167; sapii- folium, 167; spurium, 185; terrabanum, 173; utile, 17 Broussonetia brasiliensia, 158; fps 158; tinctoria, 158; xanthoxylon, 158 Bucephalon, 159; racemosum, 159 Bucholzia ficoidea, 383; philoxeroides, > EE 382, var. diffusa, 383, a, 383, var. radicans, 383 ere William C.: Sorocea, 161 Buttercup, 457 C Cabalyxtez ; 383 Cabello de ángel, 459 Cabomba, 454; australis, 455; palaeformis, 455; piauhye ensis, 454, f. albida, 454; pu- bescens, 454 Cacao volador, 537 urs e 178 Ge elaria, 364; indica, 366; sicula, 366 Cadena p^ amor, 31 Cadillo, Mere ly 319; leptobotrys, 320; tamnifo- lium, "m Calalü, 356, 358 Gare 286; mexicana, 286 Calete, ‚360 Cam End eria ege 329 TI mM ula, 338; Canilla = pava, 312 Car id vermicularis, 3 3 Cardiolochia, 2 Ca Caryophyllaceae, 432 alea Castalia, p rudgeana , 45 —_— 179; SE 180; fallax, 180; pana- Castilloa, 179, Caucho, 180 Cecropia, 211; arachnoidea, 213; asperrima, 213; bicolor, 215; commutata, 215; eh I5; eximia, 214; humboldtiana, 213; ampla, 451; 1 Weër, ec? iana, 178 blanda, 452; [Vor. 49 ANNALS OF THE MISSOURI BOTANICAL GARDEN signis, 215; jongipes, 214; maxonii, 215; mexicana, . macrostachya, 215; Ee 215; panamensis 215; peltata, ; schiedeana, 215; vogeleri, 215 Celedonia, 412, , 353; argentea, 354; buttonii, 354; coccin ea, 354; cristata „ 354; margaritacea, 354; marilandica. 354; pallida, 354; pani- culata, 361; pleiogyna, 352; pyramidalis, 354; tomentosa, 363 Celtis, 150; aculeata, 151, var. laevigata, 152, r. serrata, canescens, 148; hiki noides, 151; riparia, 148; schiedeana, 148; zizyphoides, 152 Centrostachys, 364; aspera, 366; indica, 366 Centunculus, Ce phalotomandra, 397; fragrans, 397; pana- mensis, 397 E 437; consanguineum, 437; glome- ratum, 437; nemorum, 439; viscosum, 437, var. consanguineum, 43 Ceratocephala, Ceratococca, Ceratophyllaceae, 456 Ceratophyllum, 456; demersum, 456; natum, p floridanian, 456; llerenae, ez 5 actez, Chaetoptelea, xA meia 145 Chamissoa, 361; ssima, 361; subsp. pou 363, var. ron 363, var. dioe paniculata, 363, var. glabrata, 363, grandispicata, 363, f. sem ispicata, 363, v lax iflor ra, E var. rubella, Gg celosioides, ie 264. E closes, Tr ar. uch cumbens, 364 v r. pubescens, 364 Chancanil, 377 Cheiropsis, 457 E ; chi- 347; ME. 344; ec er var. leptophylloides, 345; murale, 344, var. albescens, 344, var. carthagenense, 344, var. var. leptophylloides, 345, ssp. prater- 1962] FLORA OF PANAMA—INDEX icola var. leptophylloides, pt que forme, ; retusum, anta- dette 347; spathulatum, 347; affe loin 347; vagans Chetrophis, 433 Chile de perro, 367 Chilillo, 312, 314, 315; Chilillo de clucho, 314; Chili llo de perro, 312 Chloranthaceae, 121 Chlorophora, 156; ieu 156; tinctoria, 158 Chondodendron, 470, 472; cretosum, 470; hederaefolium, 473; hypoleucum, 470; sca- r tomentosum, Chondrodendr, 470; hypoleucum, 470; tomentosu Chuché, 55 Cissampelos, 479; acuminata, 483; andro- rgentea, 483; auriculata, vp eor 483; e deracea, 483; Ae trop 482; hirsutis- ba ; kohautiana, 483; limbata, 483; carpa, 482; monoica, 483; myriocarpa, 482; orinocensis, 483; a : f. emargi- nato- mucronata, enormi 2 var. var. er 484, v shot, 483, var. typica, 484; neg Dee 482; subreniformis s, 483; tamoides, 483; testudinaria, 483; tomentosa, 483; tropaeo- lifolia, 480 Citriosma, 538; guianensis, 541; diver- gentifolia, 541, var. nuda, 541; ‘wisest: ; riparia, Citrosma, 538; discolor, 541 Clarisia, 163; colombian na, 163; mexicana, ee , 429; patens, 429; re- = d d i c glabra, 459; goudotian grossa, 458; Mees set a, 458; E 459; integra, 459; medusae, 458; polyc phala, 458; p epuitlolia. 458; rho Fasc 458; sericea, , 458 thalictroides, 458 Clethropsi Cockscombs, Coccoloba, 332; acuminata, 332, var. glabra, 332, var. deese Me 32: allenii, 332; bracteolosa, ` acasana, 333, f. ra, i ibaea, 326; changui- nolana, 332; coronata, 326; darienensis, 30; Goudot tiana, 327; lasseri, 324; sis, 328; riparia, Ser: strobulifera, 332; raaa, 324; uvifera, 329; venosa, 1 omues 326; williamsii, 332 Coccoli Co wg KEN 470; pauper, 473 apa, Comida de canario, 446 Compona, 446 Compsoneura, 528; costaricensis, 529; sprucei, 52 Confederate vine, 31 8 arenis 318, 319; Confite rojo, 318 obaea, 319 Con Cane. 217 Conuleum, ams 461 ral vine, 18 assa, 288 poa, brevipes, 211; chagresiana, 209; magnifo olia, 209; nymphaeifolia, 209; panamensis, 209: rekoi, 1 Cow tree, 168, 171 Coyontura, 377; Coyontura de pollo, 377 Cremastosperma, 492; anomalum, 492 Cresta de gallo, 354 Cruzeta crassifolia, 378 Cua Cuartillero , 440 un de monte, 440 Cuq Cyathola, 367; achyranthoides, 367, var. densiflora, 369, var. glabrescens, 369; geniculata, 367; prostrata, 367, var. achy- ranthoides, Cymbopetalum, a. brasiliense, 515; costari- cense, 517; ugipetalum, 515; magni- fructum, Se Cyrtorhyncha, 460 SN io E.: Magnoliaceae, 485; Wintera- Fence ion, 292 (555) 244 De Wolf, Gordon P., Jr.: , 186 EIER 261; en "262: costari- s, 262 nn ee ee Ser E 0; micro- carpa, 509; neglecta, 508; panamensis, 508 Detandra Dialyanthera, 529; acuminata, 532; latialata, 530; .otoba, 530 Dorstenia, 217; contrajerva, 217, ssp. tenui- loba, 217, var. houstoni, 217, var. tenui- loba, 217; a, 217° Drimys, 489; granadensis, 489, var. chiriqui- ensis, Drymaria, 442; es 445; barrancae, 444; cordata, 445, diandra, 445, var. pacifica, 445, var. Ee 444, var. puberula 445; cubensis, 442; diandra, 445; hirsuta > procumbens, 445; stylosa, 444; tepi- na illosa, 443 3 3 Duguetia, 5/7; panamensis, 517; vallicola, Duke, J. A.: Amaranthaceae, 348; Berberida- M Nymphaeaceae, 449; rer 305; era 456 Duschekia, 133 Eggersia, 399 Einomeia, 292 Enckea lata, 15; orthostachys, 19; planta- inea, 19; reticulata, 15; Sieberi, 19; olia, 15 anus, 135; humboldtii, 141 Pack pete simplex, 315 [Vor. 49 ANNALS OF THE MISSOURI BOTANICAL GARDEN Esmarchia, 437 Euthalia, 433 Euxolus, 354; caudatus, 356, var. gracilis, 356, var. maximus, 356 Facchinia, 433 460 e "186, subg. Pharmacosycea, 187, subg. Urostigma, 186, 188; adhatodaefolia, 192; amazonica, 198; americana, 194; anguina, 202; angustifolia, 198; anthelminthica, 189, I e . guinea, 196; sii, 200; costari St 203; coybana, 190; crassa, 192; crassinerv 00; crassiuscu 92; davidsoniae, gandii, 199; duquei, 202; eggersii iud "sedie 195; erythrosticta, 196; eugeni haughtii, 201; CEA eue 199; hernan- phoides, 202; obtusifolia, 203; ochroleuca, 196; cerstedianum, 194; om- ee 194; padifolia, 196; palmicida, ; panamensis, 201; paraensis, 201; ; proctor-cooperi, 203; 191; Re 190; putumayonis, 201; (556) 1962] FLORA OF PANAMA—INDEX pyrifolia, 198; radula, 189; radulina, 192; 1 rubricosta, 190; rubri- nervis, 198; sapida, 196; segoviae, 192; sintensii, 194; sono $ cans, 189; su lei cipe 97; trigonata, 200 erckheimii, ; turbinata, 196, 199; uberrima, 201; ulei 0; velutina, 193, 200; vermifuga, 192; 99; vicencionis, $ 189; werckleana, 192; wilsoni, 194; yuca- tanensis, 200 a ead 193 F Fle a, 230; aestuans, 23] Flor de chayutal, 313: Flor de mano, 354; Flor de san miguel, 3/8 nn. 34 . E.: Annonaceae, 491 Pius de pan, 156; Fruta dorado, 532, 534 Fustic, 158 G Gaiadendron, 246; poasense, 247 Gun sere ta 164; utile, 170 128 oe, 422 Galliaria, 355; hybrida, 358, patula, 357; Garlic weed, 410 Glinus, 422: radiatus, 422 Glomeraria, 354 a mphrena, 389; aggregata, 378; crassifolia, 378: decumbens, 389, f. albi iflora, 389, var. carinata, r. genuina, 390, v grandifolia, 390, f. roseiflora, 389; - persa, 390; ficoidea, 383; globosa, 391, var albiflo 391, var. aure’flora, 391 r. carnea, 391; ixiamensis, 390; nitida, 391; paniculata, 371, n RER: 370; perennis subsp. pseudodecumben m simplex, 390; sessilis, 381; Vou MER 378 onufas, Gossypianthus decipiens, 370; hookerianus, 370 Gouffeia, 433 Greniera, 433 uaco Guniabara, x uvifera, 329 vr 03; costaricana, 404; standley- ien uae 205, 211 245 Guatteria, 496; aeruginosa, 500; allenii, ang amplifolia, 497; chiriquiensis, 499; costar e panamensis, 499; dëse: 198 Gui Gyna Cole 432 Haenkea, 283 Hagaea agea, Halerpestis, pl Halianthus, 4 Halimus, va ‘portulacastrum 426 Hashish plan Hebanthe, 369; acts: 370; hookeriana, 370; paniculata 371; virgata, 371 Hecaton Heber. 72; peltata, 72; scutata, 73; sidae- folia, 74; ie "73: 1 mbellata, 74 Hedyosmum, 121; brenesii, 121: calloso- serratum, 122; nutans, 122; scaberrimum, oe ay. at? concinna, 278; Sieg . Euheisteria, 277; fatoensis, 278; aot, E longipes, 280; gege Ha Ae Helicostylis tl, Hl montana, 167 Helosis, 286; a a, 286; mexicana, 286 Asian E ri «299 Heymassoli, 28; spinosa, 281 Hierba de gato, 377; Hierba de pasmo, 542; Hierba Co nn Higo, 186 Ben 186 Higu Er pe 292 Ho on cordatum, 445; diandrum, 445 -o 432 Hormig Hormi igui illo, 545 Howard, Richard A.: Coccoloba, 3 ur ardia, 293; anguicida, 298; benthamii 297; costaricensis, 300; foetens, 300; niflora, -p meh en grandior ; hoffmanni, 295; maxima, ww Tita, 295; pe vg 295; veraguensis, Hueso, 330 Husecudifts, 360 Huisquelite, 360, 361 Hule, 180 Hyala, 446 (557) 246 Hylebia, 439 Hypatia 476; allenii, 477; hondurensis, 476; panamensis, 47 I — ind Pe pne rues 383; ork mense, 4; ei KE per le. 381. 7 ees d Inmortal, 392 Inophloem, 182; armata, 182 Treneis, 37 Iresine, 373; acuminata, 376; aggregata, 378; a N sia, 376; celosioides, 376 i usa, , var. 76; racemosa, 375; vermicu- aris, 378; verticillata, 376 92 Itoasia, p crassa, 288 Ixtez, 361 oe 413 156 Julandacea, 130 Tuglans, 132 Karkinetron, 319 Kex, Kidd, Harold J. (collab.): Nyctaginaceae, 393 Killip, Ellsworth P.: Urticaceae, 219 Kix-xtez, Kokera, 361: acuminata, 364; celosioides, 364; paniculata, 363 Kosaria, 217 En apfıa, 46 Krascheninikovi, 439 Kumlienia, 461 L Labtzetz, 361 Lacistema, 124; aggregatum, 124; elongatum, 124; myrie oides , 124, var. stipitatum, 124; oblongum, 124. pedicellatum, 1 126 Lacistemace re a 446; corymbosa, 447 gsm e 422 [Vor. 49 ANNALS OF THE MISSOURI BOTANICAL GARDEN Langsdorffia, 289; hypogaea, 289; janeirensis, 289; moritziana, 289; rubiginosa, 289 Langsdorfia, rbrea, 439 Lathraeophila, 286 elle 286 Lechosa, 159 Lechuga de agua, 315 Leinkeria, 239 Lemia, 430 Lengua de caballo, 308; Lengua de vaca, 308 Leonia, Lepianthes, 72; peltatum, 72; umbellatum, 74 x Leucococeus, 234; occidentalis, 235 Leucocoma, 462 Litanum, 42 Diech vermiculata, 378 Llanto, 404 Llovizna, blanca, 445 Lophox = eege e, 2 45 ng ee 2315 conduplicatus, eet alenae, 257; margin 66; polystachyus, ioter a 232; spicatus, 261; theobro- e, 257 a 147; enantiophylla, 148; trema- toides, 1 Lozania, 126; montana, 126; pedicellata, 126 Macano, 158 Mace, 528 Maske affinis, 158; chlorocarpa, 158; plu- miera, 158; polyneura, 158; poms: sed ame agin, 158; tinctoria, 158; velutina, 158; xanthoxylon, 158 Maroketi $, 135 Macropiper sien 15 Magnolia, : sect. Theorhodon, 486; nn 488; grandiflora, 486; poasana, 486; Dante e, 485 Mahonia eeben, 466 Mala sombra, 404 Malachium, 439 Malagueta de montana, 534 Malmea, 511; rend 513 Mancha, Manning, Wayne, E.: Juglandaceae, 130 Mano de leön, 354 (558) 1962] FLORA OF PANAMA—INDEX Manwood, 276 Maandi. 330 Maquira, 183 Margarocarpus, 234; obliquus, 234 Marijuana, 219 Mastate blanco, 180 Mata palo, 194, 196; Matapalo, 186 Meclatis, 457 Melanocarpum, 351; sprucei, 351 Mengea, 355; pegs 356 KEE 433 0; utata, 152; iguanea, 191, laevigata, 151: Gët 151; zizy- a ees Michelia, ds champaca, 485; figo, 485 Micropetalon, 439 Micropetalum lanuginosum, 434 Micropiper iilo dont. "98. pellucidum, 79 Minuartia 432. ensis, 547; gu 949; minutiflora, 9; nigr $ pinchotiana, 548; schottiana, 538; stipi- emed 549; viridiflora, 550 Mollugo, 422; arenaria, 423; axillaris, 423; LEE 423; usa, 423; gracillima, 3; hoffmannseggiana 3; juncea, 423; schrankii, 423; spergulaefolia, 423; tri- phylla, 423; Se HEN. 442 aculeata, . 152; anfractuosa, omisia, 152; in ee 152; oe 152; laevigata, 151; zizyphoides, 1 Monandrodendron, Monimiaceae, 537 Mora, 3 Morus plumiera, 158; tataiba, 158; tinctoria, 158; xanthoxylon, 158 Mosquito, 377 247 Mozote, 367 ae SCH Mozotlex iere Sec 319; Benthamii, x lept a, 320, var. add. 320; stuebelii, 320: volcanica, 320 Mulato, 338 Muller, C. H.: Fagaceae, 135 38 Myrica, "128; mexicana, 128; xalapensis, 128 Myricaceae, 128 Myriocarpa, 237; longipes var. yzabalensis, 38; yzabalensis, 238 Myristica, 526; aromatica, 527; cuspidata var. rufula elongata, 537; fragrans, 527; guatemalensis, 536; mexicana, 529; mocoa, ia, curvi- "ei sprucei, 529; theiodora, 537; ine p 533 Myristicaceae, 526 Nag 128 Naraveli ia, 457 andin 6 Neer. 454 Neea, 399; amplifolia, Bea anisophylla, 400; delicatula, 399; laetevirens, 400; orosiana, ai pittieri, 402; Kat oide es, 400; pyc- ntha, 402; urophylla, 402; xanthina, 400 Nau nia, Nematospermum, 124 Nervillo, 446 Nettles, 219 Neumayera, 433 Nevling, Lorin I., Jr.: Aizoaceae, 422; Balan- ophoraceae, ; Chloranthaceae, 121; Corylaceae, 133; Lacistemaceae, 124; Myri- caceae, lacaceae, 275; iliaceae, 73; Portulacaceae, 427; Proteaceae, 239; Ulmaceae, 145 Nigua, 361 Niphus, 292 Ne negro, 276 Nutmeg, 526 Nyctaginaceae, 393 Nymphaea, 450; ampla, 451; blanda, 452; Ageing 452; jamesoniana, 452; rudg a, 451 es 449 (559) [Vor. 49 248 ANNALS OF THE MISSOURI BOTANICAL GARDEN O Oaks, 135 Obeah bus h, 410 ln 472; ere 473; PEN 473; era, 473; , 473; tom tosum, EC truncata, s Odontostemma, 433 Ogcodeia, 15 Ojoche, 167; Ojoche macho, 159 275 rthospermum, 344 Orthosporum, 344; ambrosioides, 347; suf- fruticosum, 347 Ortigas, 219 Oryctanthus, 258; cordifolius, 259; occiden- talis, 260; spicatus, 261 Otoba, 532 Ottonia, 71; glaucescens, 6; Thiemeana, 71 Oxandra, 494; longipetala, 494; panamensis, 494 Oxybasis, 344 Oxygraphis, 460 PONE 476 Paico, 348 Palitaria, 446 Pallavia, 406 Palo > vaca, 168; Palo mulato, 338; Palo san Sone. 242; Be ites 242; mucronata, 242; suaveolens, 242 Papaturra rg 327 Papaturro blanco, 335 Papularia, 424 Paracelsia Parietaria microphylla, 221 Paronychia achyrantha, 383; ficoidea, 383; frutescens, 384; sessilis, 381 Pe eperomia, 76; acuminata, & acuminati- 3 folia, x adscendens, 77; agapatensis, 89; albescens, 97; allagotacta, “00: Alleni, 85; eri mplexicaulis var. mag- noliaefolia, 110; angulata, 85, var. orbicans, ; angustata v. mar 110, f. fertilior, 110, f. lutea, reducta, 108; apoda, 88; MM. 101. : (560) Baileyae, 101; Bakerii, 107; barbinodis, 91; Meg din 77; bayatana, 108; bifrons, ; bocasensis, 85; brachyphylla, 97; brevi- a, 79, var. major, 79; Carlo- , va a, , var. eylindri- bacca, 104, var. longependula, 104, var marragantina, 105; eeh en var. galana, 97; io , galioides, 89, var. aprica, 89, var. longi- olia, 89, var. minutifolia, 89, var. nigro- punctulata, 89, var. umbrosa, 89; gatu- tidifolia, 108; herbaceum 106; olia, var. ciliifera, 114; Heydei, var. brevi- peduncula, 79; hispidorhachis, 112; hispi- 3 perhispi ELT var Sellowiana, 111; hygrophiloides, 94; hyme- deg "d ; : soni, 89; u e 93; novae-helvetiae, 100; mularifolia, 96; iania, 78; oblongibacc, rg ; 105, var. ar. s bvillo osa, 106; o folia, 701, var. fatoana, 101, var. minor, 101; obtusifolia, 108, var. emarginata, 109, var. longibracteata, 109; palm raed lata, 117; parietariaefolia, 101; ek? 100; pellu om 79; pelluci ida, 79, var. minor, 79, pygmaea, 79; T ta, Gei percuneata, 99: perglandulos , H6r p hispidula, 111; pcitamibusetél. 117; il. nervia, 111; piperorum, 104; portobellensis, 1962] FLORA OF PANAMA—INDEX 103; producta, 107; pruinosa, 72; dependens, 79; pulicaris, 106; p bere 7 quadrangularis, 85; Minis d E 2. quate nata, 84; reflexa var. americana, 82; re- lexa f. americana, "3 , var. enervulosa, 82; reniformis, 106; re 106; reptans, EN rivi-vetusi, 100; CS 6; rotun folia, f. ovata, 96, var. subelliptica, 06, var. pre “96; Ruiziana 78: saginans, 85; sarcocarpa, 86; sarcodes, 77; scandens, 106, var. fertilior, SS Lee eem na, 86; bertii, 92; septuplinervia, 115: serpens, 106; Kiemen 74; similis, 98; simulatio, 101; specio. 72; suaveolens, 89; subcorym- bosa, 89: sublaxiflora, 97; subquadrifolia, 83; sulrotanda, 110; substriata, 119; tene- braegaudens, 115; tenera, 111; GC 96; trinervis, 97; umbellata, 74; ri- gaudens, 8]; urocarpoides, 106; asian 91; variegata, 115; viridisp 86, var. perejil, 86; Wedelii, 118; Williamsii, 118; W oodsonii peltata, 178; tessmannii, 178; trophophylla, 179; en Persicaria acumin 313; hodroriprrpidn, 312; Ero unctata var. ecilia 310, var: robustior, 310, var. en na, € robustior, 310 Petatillo, 446 SE 408; alliacea, 410, var. grandifolia, 4 4 var. octandra, 410; foetida, 410; hex- 410; ochroleuca, 410; octandra, 410; ih 410 Pettera, 433 Peumus boldus, 538 Pfaffia, 369; pea 371; grandiflora, 370, var. hooker riana, 370; hookeriana, 370; paniculata, 37 Pfeiffer, Howard W.: Aristolochiaceae, 291 Phanerocalyx Phase ca, 186; angustifolia, Ger glau- cescens, 189; gran ndaeva, tly anensis, 189; hernandezii, 189; mexic a 189: peru- viana, 196; i CA P 189; gers 189; vermifuga, 192 aceum arenariu , 423; 423; ho ianum, 403: side? 423 Phehertnihas Phenax, 236; xu UNS 236; hirtus, 236; mexicanus, 236; rugosu Philoxerus, 377; aggregatus, 378; crassifolius, 378; vermicularis, 378 ee hes 262; allenii, 267; cooperi, 270; rynarthron, 267, var. seibertii, 267; Se 266; davidsoniae, 267; emargi- 249 natum, 270, var. minor, 270; Er pga 268; herrerense, oe latifol ium, 266; m branaceum r, 270: Lis i d 270; aon ze E pergranulatum, 263; piperoides, 266; rigidum rubrum Sa brevispica, el ia, 271; seibertii, 267; sonanum, : EE 264; tonduzii, 268; trinervium, 271; rapi 268; venezue- lense, 271; woodsonii, 266 Phrygilanthus, Zei pos Martiella, 252; Imeri, 252; panamensis, 252 hrygilanthus subg. N 46 Phthirusa, 257; adun agdalenae, 257; 276; en 257; xA 258; pyrifolia, 258; theobromae, 257 Physocarpum, 46 PREIS BERN, Du acuminata, 413, KA bogo- t , 414; malabarica, 413; ; novahispania, 413; poly- gma, e Si pur, a icum. 413; rivinoides, 414; rugosa, 412: sessili- flora, 413; triquetra, "rd Douai s ung 408 Picas, 219 Pie de e mA 377 ier 416; acuminata, 418; glabra, 418; Kem 418; tomentosa, 418 xs de gato, 367 ;-219: acumin nata, 226; auriculata, 224; hiriquin a, 223; chrysosplenioides, 226; ciliaris, 221; cornmanae, 222; costaricensis, 223. donnell-smithiana, 223; gra acilipes, alina, 225; im- m SG IN 223; var. a u^ tria naeana Pimecaria, 128, 281 Pinosia, 44 Piper, 3; acuminatum, 77; MN 66, var. trichopus ; aduncum, 30, var. brachya threm, 32, var. laevijolium, 31: aequale, 47, var. — — , var. substenocarpum, 47; 23; aggre gatum, 124; alare, 15: e" erf mum, 5; i, 42; Alstoni, 23; alveatum, pain 40; amphioxys, 47, var. ifolium, 48; amphoricar gustum leanum, 61; auritum, 22, ‘var. ie (561) 250 folium, 22, var. Seemannianum, 22; barbi- nerve, 39; Beatin 32: lo asense, 38; bogotense, granulatum, 65; crassinervium var. t t m, 8; crassispic , 49; cricamo- le 47; Cuatrecasasi, 32; culebranum, 36; cumarglenà se LIE bricola 32, var emarginatum, 109; erectamentum, 32; eru- eer 35; erubescentispicum 3 exiguicaule, 8; exserrens var. to m, agopyricarpum, 6; falcifolium, 65; 3 49; gal ioidos, 89; 69; run 21; Howe 69; erai 22. ar. ampli 22. var. cocleanum, : hi bostes 106; hirsutum, die 2; hireliipeti olum, 58; var. Harvey 58, var. tapianum, 58; hi en n hiepidürn, he var. ellipticifoli ium, 34, var. gambo- um, sé var. trachydermum, 33; ier 108; humorigaudens, 18; ibagu- ense, bi illudens, 32; imperiale, 23; in- fralut uid .nsignifolium, 69; insuli- colum Get intersitum, 31, var. porcecitense, 31; Johnstoni, E Killi ipi, 3, var. calder- (562) [Vor. 49 ANNALS OF THE MISSOURI BOTANICAL GARDEN anum, 32; konkintoense, 32, ladrillense, racteum, 64; lati- bracteum, 1 um, 15; cum, 6, var. latifolium, 6; leptocladum, 52; lincoln ense, 6; lin , 55; liratinerve, 54; lu gnifolium, 66; luci- gaudens, 42, var. i Afleniks 42; Luschnathi- anum, 64; macrophyllum, 63; macropunc- tatum, 10; macrourum, 63; maculosum, 115; magnantherum, l1; magnoliaefolium, bracteum, 39; palmasanum, 44; p 55; papyraceum, 8; en ES paso- anchoense, 45; paulownifolium, 5; pedi- cellare, 61; pellucidum, a peltatum e 74; peracuminatum ; N 51, var. subgla ee 51; perfugii, 58; perlasense, 67; Permari, 6; persubu- latum, 34; peruligerum, 7; e esar sanum, 3 peta pay ytolaccae- ee mE 68; var. ollium, ai var. pachypu um, 15; reticulatum, 19; riparium, 68; rivi-vetusti, 32; rotundifolium, 96; sala- mancanum, 19; salinasanum var. parvi folium, 39; buanum, 48; sancti-felicis, 32; san-joseanum, 16, var. aristolochi foli 18, kobense, 16, var. lorenzanum, 65; saururoidei, 19; im. serpens, 106; Ree 74; Siebert, 19; 1962] lans, 9; singulare, 9; smi see FLORA OF PANAMA—INDEX 251 en 308; sect. ae os are 334; Pena 309; s Tinaria, 309; simu 15; speciosum, 72; sperdinum, 50; s nervium, 68; Stevensi, 62; Storkii. 42: subcaudatum, 59, var. maternale, 59; subdilatatum, 39; subnudibracteum, 66; um, y trınerv: SEN, cie ibt n dime el Ca taba- gamboanum, 39; viridicaule, 36; vita- bundum, 34; Wagneri, 43, var. minuti- spicum, 43; Wedelii, 56, var. retrohirsu- tum, 57 hiteae, 70; Williamsii, = xylopioides, 64; yapeanum, 38; zedio iense, 47 Piperaceae, 1 Piratinera, 164; guianense, 169; panamensis, 169; oe 173 Pircunia, Pisonia, 406; aculeata, ed dein 405; loranthoides, 406; monotaxadenia, 406; Sieberi, 406; villosa, 406. Pistolochia, 292 abeat crassifolius, 383 Pleuropetalum, 351; calospermum, 351; taricense, 351; pleiogynum : T 1; stan leyi, Sons tucurriquense, 351 Plinthine, 433 Poleo, 445 olia, 446 Policarpaea, 446 Polycarpa, 446; brevifolia, 447; corymbosa, oyara 446; ee 447; brasili- var. ramosissima, 447; brevi- 447; corymbosa, 447, var. me 447, var. typica, 447; densiflora, 447; folia, 447: indica, 447; spadicea, 447. Polycarpoea, 446 Polycarpus, 446 Polyeonaceae, 305 SES 310, var. erg 310, var. brach var. riparium f ar. ee TUR var. capense, 318, var. nd. atum, 312; convolvulus, 315; cuspida- 313; erectum, 313; flexuosum, a for eh 313; rim eee 319; temalense, 313; iis m, 309; hydro pipe. 310, f. leu je ranthum, 312, A strigosum 12, var ás rifoliu 312, var. bushianum, 312, var. deer um, 312, 88 9 = SR Le nerianum, 316; mexicanum, 314; mite, 312; pennsylvanicum, 314; persicarioides, A portoricense, 3/2; punctatum, k r. aquatile, 310, f. qua tum . 310, var. con- tius, 310; segetum, 315; setigerum, 313; tamnifolium, 320; uvifera, 328; virgatum, Polyscalis, 367 Pomatotheca, 424 Porcelia, 513; goyazensis, 513; macrocarpa, GIER ‘magnifructa, 513; nitidifolia, 513; steinbachii, Portulaca, 430; marginata, 430; oleracea, 430, var. macrantha, 430, var. micrantha, 430, var. parvifolia, 430; penioulata, P3 parvi- ora, 430; patens, 428; reflex Sieg ion 424; monogynum, 424 Georg Potho morphe, 12: ies 74; almirantensis, auci Do a aie ^ age folia, 74; speciosa, 73; smi "73. var. grandis, 73; umbellata, 74 Poulsenia, 182; pana 182; armata, 182 Pourouma, 205; a , 206; en 208; johnston ii, 206; oraria, 208; palma 208; radula, 206: scobina, 207 252 Pouzolzia, 234; guatemalana, 234; obliqua, 234; occidentalis, 235 Pseudolmedia, 184; havanensis, 185; spuria, SC seudosorocea, 161 Pattacenthus, 249; allenii, 250; americanus, alyculatus, 249; ismarii, 2 lateriflorus, 250; scheryi, 249; Br 252 Psophiza, 293 Pteriphis, 293 Pupalia densiflora, 369; prostrata, 367 um vidiu um, a Parisa 426 Quaternella, 437 aad 358, 360 Quer 135; subg. Erythrobalanus, 136; AE se Pet pine 139; N 143; boquetensis, 142; ern. 142; chiri- qui 2; copeyensis, 140; co u 139; davidsoniae, 136; Be nial 142; humboldtii, 141; insignis, 136; oocarpa, 1 panamandinaea, 139; seemannii, 142; Weem 136 uiec tes, Quieche, 537 Quintoniles, 360 R Rabo de chanco, 367 Racoma, 4 Raeder, Katherine: Phytolaccaceae, 408 Ram Rarer d 456 Ranunculus, 460; amarillo, 462; E 461: intermedius, 461; pilosus, 462; r 461, var. ee ere 461, var. bidon EN 277 Rem Ste iguanaeus, 151; micranthus, 148 Rhaptostylum, 276 Rhodalsine, 433 Rhodes, D. G.: oe - Rhopala. 239; boissieriana, mpli icata, > 3 a Nds c 240; media, 240; o valis, 240; pii ge var. integrifolia, 240. veragu- 240 ae aert 461 Ribeirea, 283 [Vor. 49 ANNALS OF THE MISSOURI BOTANICAL GARDEN Rivina, d acuminata, 418; americana, 419; e bens, a 419; tetrandra, 418; tinctoria, 418: viridi- flora, 418; viridis, 418 Rivinia, 4 Rizzini, Carlos Toledo: Loranthaceae, 245 Robbairea, 446 , 524; microsepala, Roemeria, 354 Rellinia. 523; GE 524; permen S. 524; ee 524 Ropala, 239 Rosea, 373; elatior, 375 Rottboelia, 281 Ro ubieva, 344 anthelmintica Roupala, 239; d eege? 240; darienensis, 240; discolor, 240; dissimilis 240; montana, 240, var. dentata, 240, var. montana, 242; pana- mensis, 240; pyr yrifo lia, 7240; repanda, 340 Ruibarbillo, 308 Rumex, 306; acetosella, 308; costaricensis, crispus, 308 Rupala, 239 Ruprechtia, 336; martii, 340 Ruprechtia, 462 m, 383 ae ia, 163; urophylla, 159 Salica er 127 Salix chinensis, 127 1, 545 Sangrinaria, 39] Sanguinaria, 383, 39 Sangre fruta E 537 Sarcoca, 410 Sarcogonum, 319; tamnifolium, 320 eg 74; incurva, 75; naranjoana, 75 rratia, San a, 150 Schilleria . er 47; oe 22; ca folia, 17; caudata, - m andi SH "68: marginata, 15; riparia, ia, 68. (564) 1962] FLORA OF PANAMA—INDEX Schizotechium, 439 Schoepfia, 283: americana, 284; arborescens, 284; schreberi, 284; vacciniiflora, 284 Schoepfiopsis, 283 =~ sat 415; debilis, 416 us, 354 eg Sue? 329 Sebo, 532 Sebophora, 532 un 39 Secretania, 275; loranthacea, 276 ie. 133 Senftenbergia, 289; moritziana, 289 Serturnera, 369 S, Sesuvium, 426; ortegae, 426; pedunculatum, 426; portu ulacas trum, 426; revolutifolium, 426; sessile, 426; sessiliflorum, 426 Siebera, 433 Sieboldia, 457 Siempreviva, 391]; Siempreviva de monte, Siparuna, - brasiliensis, 547; — 539; diandra, 546; discolor, 542; foeti 542; ois ger griseo seo-flavesce ens, guianensis, 541, var. divergentijolia, 542, var. glabrescens, 542. var. longifol olia, rab var. nitens, 542; nicaraguensis mensis, 54); pa Sieg 541: pauciflora, alan gii m WM T var. ee 163 Solanoide. 416; laevis, 418; pubescens, 418; undulata, 418 Someraue uera, 433 Solenostigma, 150 Sorocea, 161; affinis, 161; pubivena, 162 148; riparia, 148; schiedeana, 149 Squibbia, 426 Stachyarpagophora, 364, aspersa, 366 Stachyphyllum, 248 Steffensia, adunca, 31; pois 64; hirsuta 32; Luschnathiana, 64; nitida, 64; erste = tuberculata, 63; oe 64; xylop es, ` Steiremis dor oie 383; repens, 382 Stellaria, 438; baldwini, 439; ciliata var. 253 439; subsp. glochidosperma, 439, montana, 439, f. reichenbachii, 439, japonica, 439; ovata, 440; pu SC 439 E Stenanona, 511; panamensis, 511 Strophium, 433 Struthanthus, 254; aduncus, 257; marginatus, 255; orbicularis, 255; polyrhizus, 257; achyus, 254; rotundatus, 2 usp Svenhedini Sychinium, 217 Synzyganthera, 124 Taba de giiegiiecho, 3 Tabaco, 338; Tabaco é monte, 338 Tabacóti; 38 Tabudo, 377 Tafalla, 121 Talauma, 486; gloriensis, 488; sambuensis, Talinium, 428 429; roseum, 429; See 428; spathulatum, rana, dec tangarana blanca, 340 is Zeg wd ata, ee manillensis, 383 Teloxys, 343 den 547 Thalictrum, 462; panamense, 464; viridulum, 464 Thonningia janeirensis, 289; mexicana, 289 Tithona, 4 Tithonia, 416; humilis, 418, var. canescens f. albiflora, 418, var. glabra, 418 Tmuul 39 2 Torrubia, 403; costaricana, 404; panamensis, = chichilea, 149; 148; canescens, 149; 149; " 49; melinona, 149; micrantha, r dogm 49, var. strigillosa, Le? mollis, 149, riparia, 149; ia, SE 149; schiedeana, 149. gear iie 254 Trencilla, 446 Tri rianthema, 424; flexuosa, 424; mdi Ss 424; port rtulacastrum, 424; procum ben een 419; octandrum, 419; poly- ect , 421; rivinoides, 419 Trig eia anastomosans, 515 Tripa i Fa pollo, 440 Triplaris, 335; am » 339; arnottiana, i ; bo colom- Mac et éi 3356. avonii, 339; poeppigiana, 340; 340; Ce, 339; pyramidalis, 330; s seto 340; surinamensis, 339; vahliana, williamsii, 340 Pb sii 373 Tropexa Trophis, 150. americana, 159, var. me nalis, . ramon, 159; chorizantha, 159; macrostachya, 162; racemosa, 159 Troxirum quadrifolium, 83 Tryphane, 433 Tsetz. U Ubero, 335 Ule, 178; Ule-ule, 180 Ulmaceae, 145 Ulm probes’ u. 145 Una a Unona a 910; panamensis, 508; viridiflora, 5 Unonopsis, Sa panamensis, 501; pittieri, 501; schip Urera, 231: ‘alceifolia, 233; baccifera, 232 SE SZ , $ Ka > MP a I o Le +s S + = "3 ® m, folium, 194; geminum, 196; hartwegii, 194; involutum, 203; i stedianum, 194; paraensis, 201; sapidum, 1 i 196; sulcipes, 196; docs aestuans, xe baccifera, .232; caraca- 232; cilia sicut 233; ew ia, 225; parietaria, 2t osa, 232 KE 219 Uva brasiliensis, 515 [Vor. 49 ANNALS OF THE MISSOURI BOTANICAL GARDEN Uvaria, 515; brasiliensis, 515; viridiflora, 515 Uvero, 324 V aco, 486 Valeriana "e 394 sn eg 335; melaenodendron, 336 Verdolaga Verhuellia pes 106 Vi T Villamilla, 419; octandra, 419; polyandra, 421 ze. 454 457 Virola, 1532; boliviensis, 533; elongata, 537, 536; panamensis, 533; peruviana S tomentosa, 533; rufula, 537. me 533, var. curvinervia, 533; ee $37; venezuelensis, 533; warburgii, 5. Viscum cordifolium, 259; latifolium, 266; mucronatum, 270; undulatum, 268 Viticella, 457 Vulvaria, 344; trachiosperma, 345 ee, 545 Wild Tomato, 418 Wilhelmsia, 433 Wine wood, 532 Wintera, 489; granadensis, 489 Winteraceae, Wormseed, 348 X 389; decumbens, 389; globosa, 391 SE 433 Xerandra, 373; celosioides, 376; elatior, 375 Xerea paniculata, 37 Xerosiphon, 389 etz, 358 gien and decre dd americana, 281; bor 282; elliptica, 281; flumi- ifolia, mo flora, 281; oblonga, 282; spinosa, 281 ylopia, 503; aromatica, 505; bocatorena, 504; frutesce ns, 503; macrantha; 505; muri- cata, 503; setosa, 503 Xtez, 361 (566) 1962] FLORA OF PANAMA-—INDEX Zacxiu, 377 Yaya de mono, 545 eia, 424 Yuncker, T. G.: Piperaceae, 1 Zaleya, 424 Zallia, 424 Zacpaiché, 367 Zactezxiu, 377 Zizyphus commutata, 151; iguanea, 151 Zorillo blanco, 367 (567) 255 256 [Vor. 49 ANNALS OF THE MISSOURI BOTANICAL GARDEN GENERAL INDEX TO VOLUME XLIx New scientific names of plants and the final members of new combinations are printed bold face type; syno nyms italics; and all other matter in ordinary ty A Acanthinophyllum: key to species, 27 Acanthinophyllum, 1, 27; ilicifolia, 26, 27, 29; spruceana, 26, 30; strepitans, Anaxagorea, 194; allenii, 194, 195; pana- mensis, | Angelin Anomospe 163 anama, 179 SECH 209, 210; glabra, 208, hayesii, 211; purpurea, 208; spraguei, 209 Anona australis, 208; echinata, ustris, 208; peruviana, 208: 208; prestoei, 208; uliginosa, 208; uncinata, 209 Aphanostemma, 148 Arcteranthis, 14 Asclepiadaceae: bibliography, 118; discussion, 109; evolution of the pollinating ie 95; floral morphology, 1/1, 126, 129; polli. nium and translator development, 106; summ fs Asimina inier 205 Aspidophyllum, 149 Atragene, 1 5 Batrachium, 148 Beckwithia, 149 Berberidaceae of Panama, 154 Berberis, 154; nigricans, 155. 156 Botryopsis 158; spruceana, 158 Bucephalon, 6; mexicanum, 13; racemosum, 6, Burger, William C.: Studies in New World oraceae: Trophis, Clarisia, Acanthino- phyllum, 1 Cabomba, 142; a 142, 143, forma ao 142; pubescens, 142 ale Castalia, 138; ampla, na, | 139; blanda, 140; ci 41; sexuality, 39; taxonomy of the group, 36; variation in populations, 50 Clematis, "M5: am age m having reference to figures and plates, in Catasetum macroglossum, = a Ge platy- glossum, 45, 53, 55; aff. s Ceratocephala, 148 Ceratophyllaceae of Panama, 144 M demersum, 144; echinatum, 44; floridanum, 144; llerenae, 144 ropsis, l ondodendron, 160; hederaefolium, 161; hypoleucum, 158; scabrum, 158; tomen- 161 m, Chondrodendron, 158; cretosum, 158; tomen- tosum, 158, 159 Cissampelos, 167; acuminata, 171; andro m argentea, 171; eege 171; benthamiana, 171; caapeba, ; canescens, 171; naria, 171; tomentosa, 171; tropaeolifolia, Citrios sma, 226; guianensis, 229, y divergen- tifolia, 229, En uda, 229; pauciflora, 227; riparia, 232 Citrosma, 226; discolor, Ca Clarisia: key to species, Clarisia de ir biflora, bas 23, subsp. biflora, p. mexicana, 22. 23; colom- biana, 2; ilisifolia, 26, 28, var. cuyunensis, va mollis, “ob: nitida, 24; p mensis, Ee racemos a, 24, 25; sprussans, 26, 30; strepitans, 26, 28, var. cuyunensis, 28, var. homes ensis, 28, var. micranthera, araensis, ird urophylla, 10, 26 a, 147; b 28, var. ana, var. var. B brasiliana, 147; discolor, 147; flori- bunda, 146; glabra, 147; goudotiana, 146; 1962] INDEX grahami, Nine Sea re haenkeana, 146; havanens 147; gra, 147; medusae, 146; vinee ira "46. populifolia, 146; dt 146; sericea, 146; thalictroides, 146 Cocculus chondodendron, 158 Comacum, 21 Cometae Morphology of the Osmun- dac Compsoneura, 216; costaricensis, 217; spru- ei 217 Coptid remastosperma, 180; anomalum, 180, 181 yenoches egertonianum, 48; lehmannii, 46 Cymbopetalum, 202; brasiliense, 2 - taricense, 205; lanugipetalum, 203, 204; magnifructum, 201 Cyrtorhyncha, 148 D IE: erg of Panama, 173; Winteraceae of Panama, 177 Desmopsis, 196; en 198; a slabrab, 197; maxonii, 198; carpa, 197; panamensis, 196 Das n. 158 Dialyanthera, 217; "Tec 220; latialata, 8, 219; otoba, Dissopetalum, 16 Dodson, Calaw way H.: Pollination and Vari- ation in the subtribe Catasetinae (Orchi- da Sieten aceae Dressler, Robert L.: Index of Orchid Names Drimys, pe granadensis, IT, Duguetia, 205; panamensis, ais Gi valli- cola, Duke, James A.: Berberidaceae of Panama, d Panama, 214; E Epibaterium? tomentosum, 158 Eschweilera Koch ex, 2 Euglossa viridiss , 49 Hilana dacs. E 44, 46; tropica, 45 Excoecaria ilicifolia, 27 Ficaria, 148 Flammula, 149 Floral Morpholo: of Secam and the Evolution of the Pollinating ‘ha in age: aceae, Fries, R. E.: Annonaceae of Panama, 179 257 G Guatteria, 184; re € allen nii, ae amplifo lia, 185; ice isi 188; hypoglauca, 20h inuncta, "186, var. caudata, 186; lucens, 184; Ouregou, 186; panamensis, 187; slateri, 186 Halerpestes, 149 Hecatonia, 148 Hewitson, Walter: Comparative Morphology of the Osmundaceae, 57 Hyperbae 164; allenii , 165, 166; hon- durensis, 164; panamensis, 164 I Index of Orchid Names—1961, 131 Krapfia, 148 Kumlienia, 149 E Leonia, 226 Leptopteris fraseri, 62; hymenophylloides, 62; superba, Leucocoma, 150 M Magnolia, 173; allenii, 176; sororum, 174, 175 Magnoliaceae of Panama, 173 Malmes, 199; h ypoglauca, 201 Meclatis, 145 Kee of Panama, 157 Mollinedia, 235; costaricensis, 236; darienen- sis, 235; pinchotiana, 236: stipitata, 237, Monimiaceae of Panama, 225 Moraceae, Studies in nr World, | Moraceae: Kuer Cannes 3; enumer- ation of the genera and species, 31; mo to com na 32; index to taxa, 34; inflorescen to genera, 5; literature cited, 31; peee d flowers, 3; zeien onships of the genera, 4; pee ve rs, 2; taxonomy, rmodes p Misses 49: colossus, 40 Mag Gen 145 Myristicaceae o of Panama, 214 Myristica, 214; aon. 215; cuspidata var. elongata, 225; fragrans, ne 258 Naravelia, 145 Nectris, 1 Nymphaea, 138; ampla, 139; blanda, 140, 141; rudgeana Nymphaeaceae Ve Panama, 137 ee 160; "ree 161; pim aupera, 161, 162; scabra, 161; Olmedia erythrorhiza, 24; species, 18 Orchid Names, Index, 131 Osmunda banksiaefolia, 63; bromeliaefolia, 63; cinnamomea, 62; claytoniana, 62; japonica, 60; javanica, 63; leno: 60: regalis, 60; Kee el Osmu e: bases, 69, 71; literature cited, 90; materials and methods, 58; observations, 59; rhi- zomes, 70; roots, 81; sporangia, 64, summary and conclusions, 88, venation, Mene 182; longipetala, 182, 183; pana- 182 Beste 148 Pachygone, 164 Paracelsia, 235 Physocarpum, 150 Pollination and Variation in the subtribe Pseudosorocea Poeppigii, R Ranunculaceae of Panama, 144 8; uniortiio 150; clintonii, Ranunculus, 149; intermedius, 149; pilosus, 150, 151; repens, 149, o glabratus, 149, var. pleni- Rhodes, D. G.: Menispermaceae of Pan- T Rhopalopodium , 149 Rollin Org microsepala, 212; permensis, 212, "A3: pittieri, 212 Ruprechtia, 150 S Safwat, Fuad M.: The dede Morphology of Secamone and the Evolution of the Pollinating Apc um in Asclepiadaceae, Sahagunia, 20; colombiana, 22; cana, ; pa diab 28; strepitans, e uro- phylla, [Vor. 49 ANNALS OF THE MISSOURI BOTANICAL GARDEN Sebophora, 220 Secamone: development of anther, 105 floral deeg, 95, 121; frutescens, floral morphology, 122; floral morphology and ana ; OB, floral bieten, 101: materials and methods Secamone alpinii, floral deeg 125; frutescens, floral morphology, 122 Sieboldia, 145 Siparuna, 26; cauliflora, 227; diandra, 234; disc o: isl d'Ee 230; griseo-flavescens, s 229, Ò divergen e St p eabnesens, E y longifolia, 230, v nitens, 230; nicaraguensis, 232, § Weed mensis, 230; ege ora, 227; riparia, 232, var. calantha, 232, var. grandiflora, 232, var. macrophylla, 232; tetraceroides 230, 231; ana, Skutchia, 6; DECH 6, 13 Soaresia, 20; niti Sorocea eji daana 8; guilleminiana, 18; m 13; pubivena, 18, 26; steno- hylla.. 28 Stenanona, 199; panamensis, 199, 200 5 nn umnera, venhedinia, 174 E Talauma, 174; gloriensis, 176; sambuensis, 176 Tetratome, 235 Thalictrum, 150; panamense, 152; viridulum, 153 Todea barbara, 62 Trigynaeia anastomosans 203 Trophis, 1, 6; key to species, T; americana, B ny hax. Ramon, 10; aurantiaca, 18; chiapensis, 16, 17; chori- zantha, 13; cuspidata, 18, 19; Hilariana, $ D XI- u 9, subsp. racemosa, 9, 12, subsp. ‚9, 10; Ramon, 10 ramon U Unona EE 198; panamensis, 196; i. 188; panamensis, 189, 190; pit- diee eri, i. 189 Uva brasiliensis, 203 Uvaria brasilensis, 203; viridiflora, 203 boobs 142 Viorna, 145 Virola, ech a 221; elongata, 225, var. pun 225, var. subcordata ; ee 224; Ween 222; laevi- 1962] INDEX 259 gata, 224; merendonis, 222; mocoa, 221; W mycetis, 221; nobilis, 224; beet ag Wintera, 177; granadensis, 177 i i i 17 Winteraceae of Panama, 177 225; sebifera, 221, var. urvinervia, 221; theiodora, 225; Ce 22]: warburgii, X 22 Xylopia, 191; aromatica, 193; bocatorena, Viticella, 145 192; frutescens, 191, 192; macrantha, 193; muricata, 191; setosa, 19] MISSOURI BOTANICAL GARDEN STAFF Director Farrs W. WENT EDGAR ANDERSON, Jonn D. Dwrek, Curator of Useful Plants Research Associate Henry N. ANDREWS, RAYMOND FREEBORG, Paleobotanist Research Associate Huch C. CUTLER, Norton H. Nickerson, ive Director orphologist Carrott W. Dopcs, TRIFON VON SCHRENK, Mycologist Assistant Curator Mien of Caraway H. DODSON, Economic Plants Taxonomist GG Curator of Owen J. SEXTON, Living Plan : Rosznr L. Dod EE ad Editor GEORGE B. Van SCHAACK, Curator of Grasses and James A. D eee AMES U Assistant ie RosrRr E. Woonson, Jr. of the Herbarium Curator of Herbarium BOARD OF TRUSTEES ` President Henry HrirCHCOCK Vice-President Lascesren B. Faust Second Vice-President Henry B. PrFLAGER eer F.BAER . - Jonn S. LEHMANN Danter K. CATLIN Roser W. Orto Samt. C. Davis Warren Mi Dn se r FRENCH aa: Rozert Brooxınos SMITH EX-OFFICIO MEMBERS L. Capican, Srrarrorp LEE Mor pore of the Diocese of — ~ President of the Academy of ee, > Science of St. Louis Dann Scan, 0. . Tromas H. Enor, "3 = BEI E ` Chancellor of — A (0 00 Ems RAYMOND onp R. Tucker, Messe of the a of S Louis `