ANNALS OF THE MISSOURI BOTANICAL GARDEN Annals of the Missouri Botanical Garden Volume XXXVI 1949 With 41 plates, and 81 figures Published то ас Galesburg, Illinois, Бу the Board of Trustees of issouri Botanical Garden, St. Louis, Mo. Entered аз second-class matter at the post-office at Galesburg, Illinois, under the Act of March 5, 1879. 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 arden. Information The ANNALS OF THE MISSOURI BOTANICAL GARDEN appears four times during the calendar year: February, May, September, and November. Four numbers constitute a volume. Subscription Price -$10.00 per volume Single Numbers оо 250 Qc Contents of previous issues of the ANNALS OF THE MISSOURI BOTANICAL GARDEN are listed in the Agricultural Index, published by the H. W. Wilson Company. 5ТАЕ OF THE MISSOURI BOTANICAL GARDEN Director Скоксе T. Moore HERMANN VON SCHRENK, RopERT ХУ. 5СНЕВУ, Pathologist Research Associate JessE M. GREENMAN, Gustav А. L. MEHLQUIST, Curator ы of the Herbarium Research Horticulturist CARROLL x: РОрсЕ, Кола M. Tryon, Mycologi Assistant Curator of the Herbarium EpGAR ANDERsON, Скоксе B. VAN SCHA Geneticist Honorary san yes кој е. RoBERT E. ХУоорвом, Jr. JULIAN А. STEYERMA Curator of the шеніне Honorary Research үе Henry М. ANDREWS, NELL C. Hor Paleobotanist Librarian oe Edit tor of Publications GERALD ULRICI, Business Manager BOARD OF TRUST OF THE MISSOURI noram ur GARDEN President RICHARD J. Lock woop Vice-President DANIEL К. CATLIN Second Vice-President EuGENE PETTUS Jonn S. LEHMANN DupLEY FRENCH GEorGE T. Moore Henry Ниснсоск A. WESSEL SHAPLEIGH THAN А. H. SHEPLEY L. Ray CARTER EX-OFFICIO MEMBERS ARTHUR H. Com WILLIAM P. GRU me of Washingto on University ЈОЅЕРН M. Акт, Mayor of the City of St. Louis ERT O. WINTERE President of ы жұла EE оғ Science of St. Louis ILLIAM SCARLETT, Bishop of the Diocese of Missouri R HERB President of the Board of Education of St. Louis Секлір Urmici, Secretary utr 7 TABLE ОҒ CONTENTS Flora of Panama. Part III, Fascicle 4 (Orchidaceae, third part) Robert E. Woodson, Jr., Robert W. Schery and collaborators Flora of Panama. Part III, Fascicle 5 (Orchidaceae, fourth part) Robert E. Woodson, Jr., Robert W. Schery and collaborators Index to Part III of Flora of Panama puce Evidence excluding Mutations, ТЕЛЕ id РО, ploidy as Possible Causes of Non-Mendelian Segre- gations in Saccharomyces alaji D. Mundkur New Species of еи iron Prod. Frederick j Hermann A Fist Record for the Genus Qualea (Vochysiaceae) from North America (Panama) .. Robert W. Schery Some Pteridosperm Stems and Fructifications with Particular Reference to the Medullosae.. _ КоБеге W. Baxter ЊЕ ЕЕ ш Hill Peoples “ Aan C. R. Stonor and Edgar Anderson On Some Us 7 Maize in the Sierra of Ancash 0 Charles M. Rick and Edgar Anderson ын of the Genus Selaginella in North America North of M Alice Е. Tryon The Cytology oe ы Ке ЕК, Henry А. McQuade Stegnosperma: А New Species and a Generic Com- David J. Rogers N кешел». А New Genus of Fossil Seeds Prev- iously Assigned to Lebidocarbon Henry N. Andrews, Jr. A Rhen of the Genus H get _ Ко Ко Тау New Apocynaceae о# South Am avid de Azumbuja ind тл: E. Woodson, Jr. General Index to Volume XXXVI PAGE 1-132 133-245 247—257 259-280 281-284 285-286 287-352 355-404 405-412 413—431 433—472 475—477 479—504 507—541 543—548 551-554 ЕГОКА ОЕ РАМАМА BY ROBERT E. WOODSON, Jr. AND ROBERT W. SCHERY AND COLLABORATORS PART III JUNCACEAE-ORCHIDACEAE From ANNALS or THE Missouni BOTANICAL GARDEN Vols. XXXII-XXXVI — 1945—1949 СОМТЕМТ5 Juncaceae Liliaceae Smilacaceae (Morton) Haemadoraceae Amaryllidaceae Bomarea (Killip) Velloziaceae Dioscoreaceae (Morton) Iridaceae Burmanniaceae (Jonker) Musaceae Zingiberaceae Cannaceae Marantaceae Orchidaceae (Williams) Orchidaceae (Allen) Index SHELVE . ў STACKS Voluine XXXVI : V Mab 1 Annals of the FEBRUARY, 1949 Flora of Panama. Part III. Fascicle 4 (Orchidaceae, third part) . . . . Robert E. Woodson, Jr., Robert W. Schery and Collaborators 1-132 BLISHED QUARTERLY AT GALESBURG, ILLINOIS, BY THE BORED OF усе > daa we iin petii BOTANICAL GARDEN, MISSOU Entered as second-class matter at the ее ас Galesburg, Illinois, under the Act of March 3, 1879. 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 arden. Information The ANNALS OF THE MISSOURI BOTANICAL GARDEN appears four times sinis the calendar year: February, May, September, and November. Four umbers constitute a volume, Subscription Price ---- ———.$10.00 per volume Single Namban a ZA Е аЙ Contents of previous issues of the ANNALS. OF THE Mists BOTANICAL GARDEN åre listed in the Agricultural Index, published by the H. W. Wilson ЕГОКА ОЕ РАМАМА Part ІП. Fascicle 4 ORCHIDACEAE! By PAUL H. ALLEN 39. CALANTHE R. Br. CALANTHE R. Br. in Bot. Reg. 7: sub Z. 573. 1821; Lindl. Gen. & Spec. Orch. РІ. 249. 1833; Benth. & Hook. Gen. Pl. 3:520. 1883. Ghiesbreghtia A. Rich. & Gal. Ann. Sci. Nat. ІП, 3:28. 1845. Erect terrestrial, or rarely epiphytic herbs, with leafy or sometimes pseudo- bulbous stems. Leaves few, broadly plicate, with prominent veins. In some of the Old World species the sheathing bases of the leaves envelop the angulate pseudobulbs, the leaf blades becoming deciduous at the end of the season’s growth, while in other species the leaves persist for more than one year, with the pseudo- bulbs much reduced in size or entirely absent. In the single species known to occur in Panama, the two broadly plicate leaves are contracted below into a sheath- ing petiole arising directly from the rhizome and without any enclosed pseudobulb. Inflorescences erect, many-flowered racemes, equaling or exceeding the leaves in length, the basal portions enveloped by the sheathing leaf bases. Flowers small, on slender pedicels, subtended by linear bracts. Sepals free, subequal, spreading. Petals similar to the sepals or smaller. Lip 3-lobed, prolonged at the base into a spur, the claw of the lip connate to the column. Column short, erect, broadly winged at the apex, the base without a foot. Anther subterminal, operculate, in- cumbent, 2-celled; pollinia 8, waxy, elongate-pyriform, in groups of 4 in each cell of the anther A large genus of the Old World tropics, widely distributed in Africa, Asia and Oceania, with a single species in Central America and the West Indies. 1 is врне E to acknowledge the cooperation of Professor Oakes Ames of the Botanical Mus = University in affording the writer the ыты Эш: use of the Ames Orchid Parapet and y. The есе баат candy by Mrs. Blanche Ames and Ьу Mr. Gordon W. Dillon, v A out the tex ould iot have been salable without Profesor Ames’ permission. It is also desired to men tion е е cooperation of О, lign in p prompt determinations during the long E. of field w 57 іп Ж == ата, апа, recently, the very generous help of Mr. Charles Schweinfurth in На solving of many. of yem езеді И іп the course Ei the preparation of the manuscript. Issued March 21, 1949. (1) (337) [Vor. 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN ан cad md КЕС = Mey ча 7 NM. oc а БАр. Б нана eii. SEEN ен w. я. т а нан ~~ Ы --2 ДЕК основ 2 ji coge T 252 ——-—-- ES / ^P P E ШЕЕ A: қар ме” НИ NEC LE Sos Сааи ж ы ee ы ыз Sa ы Sl ВР, >>... ж mom anf Calantbe mexicana Fig. 148. (338) 1949] FLORA OF PANAMA (Orchidaceae) 3 1. CALANTHE MEXICANA Rchb. f. іп Linnaea 18:406. 1844; Xenia Orch. 1:205, t. 70, figs. 1—3. 1856. Ghiesbreghtia calantboides A. Rich. & Gal. in Ann. Sci. Nat. ІП, 3:28. 1845. Ghiesbreghtia mexicana A. Rich. & Gal. ex Rchb. f. Xenia Orch. 1:205. 1856, in synon. Erect terrestrial herbs, 25—70 cm. tall. Leaves 2, broadly oblong-elliptic, acute, rather thinly plicate, 30—70 cm. long and 3—14 cm. broad, contracted at the base into a sheathing petiole, arising directly from the rhizome, the basal por- tions enclosed in 2—3 thin leafless bracts. Inflorescence an erect raceme equaling or exceeding the leaves in length, produced from between the sheathing leaf bases. Flowers few to many, small, white, pedicellate, subtended by linear acuminate bracts 8-20 mm. long and 1-2 mm. broad. Sepals white, free, subequal, spreading, ovate, acute, 7-10 mm. long and 3-6 mm. broad. Petals white, free, spreading, obovate-lanceolate, obtuse, 5—6 mm. long and 2-3 mm. wide. Lip 3-lobed, 8—10 mm. long, prolonged at the base into a spur; lateral lobes erect, minutely puberu- lent, the upper margins parallel with the column; frontal lobe spreading, yellow, minutely puberulent, subrotund, shortly apiculate, 3—4 mm. long. Ovary puberu- lent. Capsule ellipsoidal, 2.5—3 cm. long and 1—2 cm. wide.! Mexico, Guatemala, Costa Rica, Panama, and the West Indies. cHiRIQUÍ: Upper Río Chiriquí Маш. about 5000-6000 ft., Seibert 171; vic. Casita Alta, Volcán de Chiriquí, 1500—200 . Woodson, Allen & Sei ibert t 825; Finca Lérida to Peña Blanca, 1750—2000 m. RUNE 5 Scbery 303; ridges south of Finca Lérida to Loma Sardina, 6000-7000 ft., Allen 4758; rain forest, Volcán de Chiriquí, 8000 ft., Davidson 934; Bajo Chorro, 7500 ft., Davidson 203; Cerro Punta, 2000—2500 m., Allen 8 Fairchild 3517. А frequent and attractive terrestrial species іп «һе bamboo-oak zone of the upper slopes of Chiriqui Volcano. 40. BLETIA Ruiz & Pavon ВЕГЕТА Ruiz & Pavon, Fl. Peruv. & Chil. Prodr. 119, 7. 26. 1794; Benth. & Hook. Gen. Pl. 3:513. 1883. Gyas Salisb. in Trans. Hort. Soc. Lond. 1:299. 1812. Bletiana Raf. in Amer. Monthly Mag. 47:268. 1818, nomen. T biebautia Colla, Hort. Ripul. 139. 1824. Jimensia Raf. Fl. Tellur. 4:38. 1836. Regnellia Barb. Rodr. Gen. et Spec. Orch. Nov. 1:81. 1877. Bletilla Rchb. f. in Fl. des Serie 8:246. 1853 Erect terrestrial herbs, with a few йі. lanceolate, plicate, ultimately decidu- ous leaves which are contracted at the base into a sheathing petiole arising from the apex of a subglobose or sometimes more or less flattened, corm-like pseudobulb considerably resembling that of a Gladiolus. Inflorescences slender, erect, leafless scapes, equaling or exceeding the leaves in length, the basal portion either pro- 1Since all measurements have been taken from dried material, it is to be expected that fresh flowers will exceed these figures by at least a third. (339) [Vor. 36 4 ANNALS OF THE MISSOURI BOTANICAL GARDEN duced from the sides of the corm or enveloped in the sheathing leaf bases, the upper portion racemose or sometimes paniculate. Flowers of moderate size or small. Sepals free, subequal, spreading. Petals subequal to the sepals but usually broader. Lip entire or 3-lobed; the lateral lobes broad, usually rounded, erect, in natural position parallel to the column, or sometimes with lobulate, spreading apices; mid-lobe of the lip spreading, with or without an isthmus, the apex often crisped and reflexed, retuse or 2-lobed; the disk with 5—7 more or less fleshy, longi- tudinal crests or keels. Column elongate, semi-terete, arcuate, the apex winged and the base auriculate. Anther operculate, incumbent, 2-celled; pollinia 8, waxy, flattened, obovate or roughly triangular in outline. A seemingly natural but perplexing genus in need of revision, now embracing about 50 species distributed from Florida and the West Indies to Mexico and Central and South America as far as Argentina, with three or four species described from the Old World tropics. It seems probable that careful comparison of available material would reduce this list to about a dozen valid species, some of which evidently are widely distributed. a. Inflorescence enveloped by the sheathing leaf bases. Flowers about 3 cm. long, produced during the rainy season (summer). Hig hand species . B. REFLEXA aa. Inflorescence not enveloped by the sheathing leaf bases. Flowers T . long, produced during the dry season (winter). Lowland чи 1. B. PURPUREA 1. BLETIA PURPUREA (Lam.) DC. in Mem. Soc. Phys. Hist. Nat. Genëve 9:97, 100. 1841; Huit. Not. Р]. Rar. 23. 1841. Limodorum purpureum Lam. Encycl. Meth. Bot. 3:515. 1791. Li n n Bletia pallida Loddises, Bot. Cab. 7: f. 620 Erect terrestrial herbs with 1—4 lanceolate, acuminate, ultimately deciduous leaves 30—70 cm. long and 0.6—4.8 cm. wide, which are contracted below into а sheathing petiole arising from a subglobose or flattened corm-like pseudobulb 2.5—3 cm. in diameter. Inflorescence a slender, erect raceme or panicle, the degree of development depending on the strength of the individual plant, the scape pro- duced from the side of the corm and equaling or exceeding the leaves in length, sometimes flowering after the leaves have fallen. Flowers few to many, rosy purple. Sepals free, subequal, spreading, 1.6-1.8 cm. long and .6—7 ст. wide, the dorsal sepal ovate-lanceolate, acute, the laterals somewhat oblique at the base, ovate, acute. Petals subequal to the sepals, 1.5-1.7 cm. long and .6—7 cm. broad, obovate, acute. Lip 3-lobed, about equaling the petals in length, the lateral lobes erect and paralleling the column in natural position, the frontal lobe extended, spreading, more or less crisped and reflexed, the apex bilobed, the inner lip with 7 erect, yellow, parallel, longitudinal, fleshy keels, only 5 of which extend to the frontal lobe. Column slender, arcuate, the apex winged, the base auriculate. (340) 1949] FLORA OF PANAMA (Orchidaceae) Fig. 149. Bletia purpurea (341) [Vor. 36 6 ANNALS OF THE MISSOURI BOTANICAL GARDEN Mexico, Guatemala, British Honduras, Honduras, Costa Rica, Panama, Colombia, Venezuela, and the West Indies. РАМАМА: foothills east of Panama City, sea level, Powell 45; vic. San Carlos, on walls of EN canyons, 0—10 m., Allen 1149; along Río Tecümen, north of Chepo road, about 30 m., Hunter 9 Allen 214; Perlas Archipelago, San José Island, sea cliffs at East Bay, prse 567; Ісасо pop. top of large rocks near coast, Johnston 022. СОСТЕ: El Valle, banks of Río Antón, 600 m., Allen 2766; Río Mata Ahogado, region southeast of El Valle, 200 m., Allen 3820; vic. El Valle, White 9 White 60. A very common and attractive species of the lowlands of the dry Pacific slope, where it is found from sea level to about 3000 ft. elevation, being frequently seen on the nearly vertical walls of the river canyons and in rocky places. The rela- tively small but brightly colored flowers are produced during the dry season (winter) after the corms have matured and usually after the leaves have fallen. 2. BLETIA REFLEXA Lindl. in Bot. Reg. 21: Ё. 1760. 1835.1 Limodorum Lankesteri Ames & Schweinf. in Sched. Orch. 10:78. 1930 Bletia Lankesteri Ames, Hub. & Schweinf. in Bot. Mus. Leaf. Harv. aly. 3:41. 1934, Erect terrestrial herbs with 2-3, usually linear-lanceolate, acuminate, plicate, strongly veined leaves which are ultimately deciduous. Leaves 25-60 cm. long and 0.8-2.5 cm. wide, the basal portions contracted into a sheathing petiole en- veloped іп 2-3 short, tubular, leafless bracts which are produced from the’ apex of a short, subconic, corm-like pseudobulb averaging about 1.5 cm. in diameter. Inflorescences erect, the basal portions enclosed in the sheathing leaf bases, the terminal portion an unbranched raceme equaling or exceeding the leaves in length. Flowers relatively large and conspicuous, in our specimens described as being purplish rose, produced on slender pedicels subtended by ovate, acuminate bracts. Sepals free, spreading, subequal, lanceolate, acute, 3—3.2 cm. long and .7-.9 cm. wide, the dorsal usually somewhat narrower. Petals subequal to the dorsal sepal oblanceolate, acute, 3-3.5 cm. long and about .7 cm. wide. Lip 3-lobed, about 3 cm. long and 2-2.2 cm. wide when spread out, the lateral lobes rounded, erect in natural position, the mid-lobe entire, obtuse, spreading, about 8 mm. long and 8 mm. wide. Inner lip (disk) with 5 fleshy longitudinal crests. Column slender, semi-terete, 2.3—3 cm. long, somewhat dilated ас the apex. Mexico, Guatemala, Costa Rica, and Panama. CHIRIQUÍ: Llanos del Volcan, 1120-1200 m., Seibert 328. 116 seems possible that this species may be that Не део онаа as Bletia нсі Llave & Lex. (in Nov. Veg. Descr. Fasc. 2:17. 1825), but the ripti . Hes is obscure, and seem- ingly the type no longer exists. he present material asas аа marked difference "from the "Botanical Register’ plate of Bletia reflexa, particularly in the shorter central даја е wever, since Bletia Lankesteri is about intermediate in this character between our spec d the typical Bletia reflexa, it seems likely that both represent local forms of a widely. Жее, somewhat polymorphic specie (342) 1949] FLORA OF PANAMA (Orchidaceae) 7 41. CHYSIS Lindl. Cuysis Lindl. in Bot. Reg. 23: t. 1037. 1837; Benth. & Hook. Gen. Pl. 3:514. 1883. T borvaldsenia Liebm. in Bot. Not. 103. 1844. Epiphytic herbs with fleshy, fusiform or clavate, often more or less pendent, pseudobulbous stems. Leaves broadly plicate, the sheathing bases enveloping the pseudobulbs, the leaf blades becoming deciduous at the end of the current season's growth. Inflorescences usually solitary, produced from the axils of one of the lower bracts of the flush of leafy new growth, the scape terminating above in a short, few-flowered raceme. Flowers relatively large and conspicuous. Sepals subequal, free, spreading, the dorsal sepal erect, the broader lateral sepals oblique and adnate to the foot of the column. Petals free, subequal to the sepals, but usually narrower. Lip fleshy, 3-lobed, the lateral lobes rounded or falcate, erect in natural position, or converging over the column, the mid-lobe erect or reflexed, often more or less 2-lobed or emarginate; disk with 3 to 5 longitudinal fleshy crests. Column short, erect, arcuate, broadly winged or roughly triangular in cross-section, produced at the base into a foot. Anther subrotund, operculate, in- cumbent; pollinia 8, waxy, oblong-ovoid, 4 in each cell of the anther. Examination of available material in the Ames Herbarium and the Herbarium of the Missouri Botanical Garden would indicate that the genus is probably limited to two somewhat variable species differing fundamentally in the number of fleshy crests of the disk and in the frontal margin of the mid-lobe of the lip. The known geographic range is from Mexico to Peru and Venezuela. A small-flowered, richly colored variety of one of these species is known in Panama from a single collection of flowering material made by Mr. C. W. Powell. Although a few subsequent, sterile specimens have been seen in the field by the writer, it is to be considered as one of the local rarities. 1. Снүзї$ aurea Lindl. var. МАСОТАТА Hook. in Bot. Mag. 7. 4570. 1851. Epiphytic herbs, with fleshy cylindric, fusiform or clavate, pendent pseudobulbs 15—20 cm. long and 1—2 cm. wide. Leaves plicate, lanceolate, acute or acuminate, 6—40 cm. long and 1.2-3.5 cm. wide, the imbricating bases persistent, enclosing the pseudobulbs, the blades ultimately deciduous. Inflorescences usually solitary, short racemes produced from the axils of one of the bracts of the leafy new growth. Flowers 3-7, 2.5-3 cm. long in dried specimens but probably somewhat larger when fresh. Dorsal sepal free, erect, elliptic-oblong, ob- тизе, 2-2.2 cm. long and 1.2-1.5 cm. wide, lateral sepals obliquely cuneate, 2.2—2.5 cm. long and 1.6— “= 1.8 ст. broad, соппасе ас the Базе Fig. 150. Chysis aurea var. maculata and adnate to the column foot, form- (343) [Vor. 36 8 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN ing a rounded mentum. Petals obovate-lanceolate, acute, 2—2.2 cm. long and 0.9—1.1 cm. broad. Lip 3-lobed, the lateral lobes subfalcate, obtuse, erect, the apices somewhat convergent, the mid-lobe rounded in general outline, more or less cymbiform, the apex 2-lobed or emarginate; disk with 5 fleshy, longitudinal crests. Column short, stout, broadly winged, 0.9—1.0 cm. long and 0.9—1.0 cm. broad at the base, which is produced into a foot. Costa Rica, Panama, and Colombia. PANAMÁ: near Arraiján, west of the Canal, about sea level, Powell 309. Chysis aurea var. maculata differs from the type in the smaller, more richly colored flowers. The sepals and petals are described as tan, often shaded purple. Lip yellow, the mid-lobe purple, with paler markings. 42. BULBOPHYLLUM Thou. BULBOPHYLLUM du Petit-Thouars, Hist. Pl. Orch. Iles Aust. d’Afr., Tabl. des Езрес. ІП, 4. 03-110. 1822; Benth. & Hook. Gen. Pl. 3:501. 1883. Didactyle Lindl. in Fol. Orch. 1:57. 1852 Bolbophyllaria Rchb. f. in Bot. Zeit. 10: 934. 1852. Epiphytic herbs, with short, subconic or angulate, 1- to 2-leaved pseudobulbs often widely spaced along a creeping rhizome. Inflorescences erect or arching spikes or racemes produced from the base of the pseudobulbs. Flowers numerous, inconspicuous, subtended by a bract, free on short pedicels or sessile in shallow pits on the fleshy peduncle. Sepals free, or the laterals connate and obliquely dilated at the base, adnate to the foot of the column. Petals smaller than the sepals, with entire or ciliate margins. Lip simple, fleshy, or with a thickened basal callus, con- tracted at the base and hinged to the column foot. Column short, erect, produced at the base into a foot. Anther terminal, operculate, incumbent, depressed-hemis- pherical or obtusely conical, usually 2 celled; pollinia normally 4, waxy, in pairs in the cells of the anther. There is an extensive list of additional generic synonyms which are not cited because they apply to Old World members of the genus. Since nearly the entire genus is of Old World tropical distribution, where it attains a polymorphic devel- opment similar to that of Epidendrum in the Americas, it seems futile to attempt a generic description here which will cover the maximum possible limits of floral and vegetative structure. The few species found in the American tropics bear little resemblance to their Old World relatives, and might, for all practical pur- poses, be separated; but in this case, as in many others, it seems best to follow established usage. The description as given here is intended only to cover the two Panama species, which are identical in superficial appearance, but differ markedly in the size and shape of the petals, and to a lesser extent in the size and form of the lip. (344) 1949] FLORA OF PANAMA (Orchidaceae) 9 a. Petals acuminate to aristate, exceeding the length of the column; nearly equaling the sepa 1. B. ARISTATUM ю ы Petals о; хаду ырайы: the length of the column, less than -— the length of the . B. PACHYRRHACHIS 1. BULBOPHYLLUM ARISTATUM (Rchb. f.) Hemsl. Biol. Centr.-Amer. Bot. 3:213. 1883. Bolbophyllaria aristata Rchb. f. Beitr. Orch. Centr.-Amer. 60. 1861. Epiphytic herbs 10—35 cm. tall, with short, subpyramidal, angulate pseudo- bulbs 2.5—5 cm. long and 1-2 cm. wide, usually spaced at intervals along the creeping rhizome. Leaves 2, linear-lanceolate, acute, coriaceous, 6—25 cm. long and 1.5—3 cm. wide. Inflorescences erect racemes 10—40 cm. tall, from the base of the pseudobulbs. Flowers in- conspicuous, nearly sessile, subtended by a bract which nearly equals the flower in length. Dorsal вера! free, concave, ovate, acuminate, papillose on the outer surface, 5—6 mm. long and 1.5—2 i 2. mm. wide, lateral sepals ovate, acuminate, papillose on the outer ‚ 151, surfaces, 4—5 mm. long and 1.5-2 mm. broad, connate at the base оз. : petals: 1, of B, and adnate to the foot of the column. Petals lanceolate, acumi- aristatum, 2, nate to aristate, 4—7 mm. long and 1.5-3 тт. wide. Lip very d pacbyrrbachis fleshy, entire, obtuse, linguiform, 2 mm. long and 1-1.5 mm. wide, hinged to the column foot. Column very short, 1.5-2 mm. long, with two apical, lateral, denticulate processes, the base produced into a foot. Mexico, Guatemala, Honduras, Costa Rica, and Panama. PANAMÁ: foothills east of am sea level, Powell s.n.; San Juan Range, sea level, Powell 3402. СОСТЕ: south'rim of El Valle de Antón, 650 m., Allen 2915; El Valle lower valley and marshes send Rio Anton, about 500 m., Hunter Allen 375. енто: Гана (и 4500 ft., Davidson 1270. 2. BULBOPHYLLUM PACHYRRHACHIS (Rchb. f.) Griseb. Fl. Brit. W. Indies, p 613. 1864 (as Bolbophyllum и: сы Bolbophyllaria bacbyrrbachis Rchb. f. іп Walp. Ann. 6:241. 1861. Bulbobbyllum vinosum Schltr. Beih. Bot. См. 362:411. 1918. Ерірһугіс herbs 10-45 ст. tall, with short, зиБсопјса!, strongly 4-angulate pseudobulbs relatively widely spaced along а creeping rhizome. Leaves 2, linear- lanceolate, acute or acuminate, 7—20 cm. long and 0.8—2.4 cm. wide. Inflorescence erect or arching from the base of the pseudobulbs, 10—45 cm. tall, the apical flowering portion of the peduncle often fleshy. Flowers small and inconspicuous, sessile in shallow pits on the peduncle, subtended by broadly triangular bracts. Dorsal sepal free, concave, ovate-acute, 4—5 mm. long and arching over the column, the laterals ovate-acuminate, 4 mm. long and 1.5 mm. broad, connate at the base and adnate to the foot of the column. Petals scarcely equaling the column and less than М; the length of the sepals, oblong-elliptic, obtuse. Lip entire, very (345) [Vor. 36 10 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 152. Bulbophyllum pachyrrhachis (346) 1949] FLORA OF PANAMA (Orchidaceae) 11 fleshy, more or less 3-angled in cross-section, linguiform, the apex obtuse, con- tracted at the base, and articulated to the column foot. Mexico, Guatemala, Honduras, Costa Rica, Panama, and the West Indies. CANAL ZONE: near Frijoles, sea level, Powell 364; along Las Cruces P between Ft. Clayton and Corozal, Standley 29102; foothills east of city, sea level, Powell s.n. PANAMÁ dro Mac forest of Quebrada Anc hi. Madden Lake region, 70 m., ieri 9 Allen 17104. ҮЕКАСОАЅ: Río де Jesús, sea level, Allen 4243. 43. EULOPHIA R. Br. EuLopPHiA R. Br. in Bot. Reg. 8:/. 686. 1823; Benth. & Hook. Gen. Pl. 3:535. Cyrtopera Lindl. in Wallich, Numer. List, n. 7362-64. 1832; Gen. & Sp. Orch. Pl. 189. 1833. Erect, terrestrial herbs with lanceolate, plicate, ultimately deciduous leaves, the basal portions contracted into a sheathing petiole arising from the apex of the short, subconical, tuber-like pseudobulb. Inflorescence an erect, leafless raceme, pro- duced from the base of the tuber-like pseudobulb, equaling or exceeding the leaves. Flowers few to many. Sepals free, subequal, the laterals adnate to the foot of the column. Petals subequal to the sepals, but narrower. Lip 3- lobed, the rounded, lateral lobes erect and converging over the column, the mid- lobe erect or recurved, sometimes 2-lobed, base of the lip produced into a saccate mentum continuous with the foot of the column. Column short, produced at the base into а foot. Anther terminal, operculate, incumbent. Pollinia 2-4, waxy. A large genus of mainly Asiatic and African plants represented in Panama by a single species. There seems no point in citing here the extensive list of generic synonyms applicable only to the Old World plants. 1. EULOPHIA ALTA (L.) Fawc. & Rendle, Fl. Jam. 1:112, 2. 22, figs. 4-8. 1910. Limodorum altum L. Syst. ed. 12, 2:594. 1767. Dendrobium longifolium HBK. Nov. Gen. et Sp. 1:360. 1815. yrtopodium Woodfordii Sims, in Bot. Mag. Ё. 1814. 1816. Cyrtopera Wood fordii we Gen. & Sp. Orch. Pl. 189. 1833. Cyrtopera longifolia Rchb. f. in Walp. Ann. 6:668. 1861 Eulophia У oodfordii Rolfe, in E Trop. Afr. 7:68. Eulophia longifolia (HBK.) Schltr. Die ‘Orchideen, р. 347. 1914. Erect terrestrial herbs with thickened, tuberous rhizomes usually made up of roughly triangular, subconic, corm-like annual sections. Leaves several, erect, lanceolate, acuminate, plicate, strongly veined, ultimately deciduous, 60—120 cm. long and 2—10 cm. wide, contracted below into a cylindric leafy petiole, the broad bases enveloping the corm-like pseudobulb. Inflorescences erect, lateral, leafless scapes, produced from the base of the corm, terminating in a raceme 70—100 cm. tall. Flowers many, of moderate size. Sepals free, erect, subequal, lanceolate to oblanceolate, acute, 2—2.5 cm. long and 0.7—0.8 cm. wide, the laterals adnate at (347) [Vor. 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN ) Jh f 1378 ў Fii Fig. 153. Eulophia alta (348) 1949] FLORA OF PANAMA (Orchidaceae) 13 the base to the column foot. Petals erect, oblong, obtuse, subequal to the sepals, 1.5—1.7 cm. long and 0.7-0.8 cm. broad. Lip 3-lobed, lateral lobes rounded, erect, the apices spreading, mid-lobe entire, usually reflexed, with numerous minutely denticulate longitudinal nerves; disk with 2 erect, alate processes, the base sub- saccate, forming a rounded mentum with the foot of the column. Column winged, arcuate, 0.8—1.0 cm. long, produced at the base into a foot. Florida, Mexico, British Honduras, Guatemala, Panama, the West Indies, Colombia, Venezuela, Peru, and Brazil. PANAMÁ: Mata Redonda, sea level, Powell 3442; hills and valleys east of city, sea level, Powell 10; vic. La Chorrera, sea level, Allen 2080. сосі.Е: vic. El Valle de Antón, 600 m., Allen 1 Rather кеша terrestrial plants found іп areas of wet savanna and along roadways, sometimes persisting in vacant lots in Panama City. The leaves are reminiscent of those of Bletia or Peristeria. The sepals and petals are green or greenish tan, the lip variously shaded with rose or purple. 43-A. CYRTOPODIUM R. Br.! CvnTOPODIUM R. Br. in Ait. Hort. Kew., ed. 2, 5:216. 1813; Benth. & Hook. Gen. Pl. 3:541. 1883. Tylochilus Nees in Verh. Bot. Gart. Berlin 8:194, Ё. 3. 1832. Epiphytic or semi-terrestrial herbs with stout, cylindric or fusiform pseudo- bulbs. Leaves broadly plicate, strongly veined, more or less distichous, the imbri- cating bases enclosing the pseudobulb, the blades ultimately deciduous. Inflorescences erect, leafless scapes from the base of the pseudobulbs, racemose or paniculate, equaling or exceeding the leaves. Sepals subequal, free, spreading, the laterals often broader and more or less adnate to the foot of the column. Petals subequal to the sepals but usually broader. Lip 3-lobed, the laterals rounded, erect, parallel with the column or the apices spreading, the mid-lobe entire, retuse or crisped, with a fleshy basal crest, the base adnate to the column foot. Column short, subterete, arcuate, produced at the base into a foot. Anther operculate, imperfectly 2-celled; pollinia 2—4, waxy. A small genus of American orchids represented in Panama by a single species. 1. CYRTOPODIUM PUNCTATUM (L.) Lindl. Сеп. & Spec. Orch. Pl. 188. 1833. Epidendrum punctatum L. Syst. Nat. 2:1246. 0. Cymbidium trinerve Meyer, Prim. Fl. ч 258, 1818. Epidendrum gigas Velloso, Fl. Flum. Ic. 9: Ё. 1827. Cyrtopodium Willmorei Knowles & Westcott, FL Cab. 1: 4. 4. 1837. Oncidium palmophilum Mart. es ç ex Lindl. Sert. Orch. sub £. 12. 1838. Cyrtopodium speciosiss imum Hort. ex Du Buysson, L' Orchid. 299. 1878. С í Cyrtopodium Saintlegerianum Rchb. f. in Flora 68:301. 1885 Cyrtopodium punctatum var. Saintlegerianum Hort. in Stein's Orchideenbuch, p. 181. 1892. 1The genus Cyrtopodium was unrecorded from Panama when the generic key was written. (349) ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN ЖАУ, Fig. 154. Cyrtopodium punctatum (350) [Vor. 36 1949] FLORA OF PANAMA (Orchidaceae) 15 Robust, epiphytic or semi-terrestrial herbs with stout, fusiform pseudobulbs 12—40 cm. long and 2—6 cm. wide, the old pseudobulbs naked or enclosed in the imbricating, persistent leaf bases which are armed at the apex with stout, sharp spines. Leaves linear-lanceolate, acute or acuminate, plicate, distichous, strongly veined, 30-120 cm. long and 2-7.5 cm. wide. Inflorescence a lateral leafless panicle from the base of the pseudobulb, 60-120 cm. tall, the bracts oblong- lanceolate, with undulate margins, resembling the sepals in color and markings. Flowers many. Sepals free, subequal, spreading, oblong-lanceolate, acute or apiculate with strongly undulate margins, greenish yellow spotted irregularly with reddish brown. Dorsal sepal 0.8-2.5 cm. long and 0.8-1.1 cm. wide, lateral sepals 1.5-2 cm. long and 0.7-1.0 cm. wide. Petals free, spreading, obovate, obtuse or apiculate, yellow irregularly spotted red-brown, 1.5—2.0 cm. long and 0.9-1.2 cm. wide. Lip 3-lobed, the lateral lobes erect, more or less convergent over the column, rounded, usually exceeding the mid-lobe in size, reddish-brown shading to yellow at the base, mid-lobe strongly undulant, crisped, yellow, marked purple or reddish brown, crest tuberculate, verrucose, base of lip an oblong limb articulated with the column foot. Column short, arcuate, with a dorsal crest, dilated or semi-auriculate above, narrowed below, produced at the base into a foot. Florida, Mexico, Guatemala, Costa Rica, the West Indies, Colombia, Venezuela, the Guianas, Brazil, Peru, Paraguay, and Argentina. PANAMA: San José Island, Perlas Archipelago, sea level, Walker s.n. (under Allen 3542). Mr. Walker describes the plants as forming colonies on rocks. Although widely distributed in the Americas this is the only known station for the species in Panama. 44. WARREA Lindl. Warrea Lindl. in Bot. Reg. n. s. 6: Misc. 14. 1843; Benth. & Hook. Gen. РІ. 3:545. 1883 Erect, terrestrial, highland herbs with lanceolate, acute or acuminate, plicate, strongly veined leaves contracted at the base into a short, stout, sheathing petiole, either with or without an enclosed, short, cylindric, tapering pseudobulb. In- florescence an erect, leafless raceme produced from the axil of the lowest basal, petiolar bract. Flowers relatively large and conspicuous. Sepals subequal, con- cave, the bases of the laterals oblique and adnate to the column foot. Petals sub- equal to the sepals but usually broader. Lip entire, the lateral margins erect; apex spreading, emarginate, 2-lobed, obscurely apiculate or entire; disk lamellate; base of the lip adnate to the column foot. Column elongate, semiterete, with a short basal foot. Anther terminal, operculate, incumbent; pollinia 2 to 4, broadly semiglobose, waxy. (351) [Vor. 36 16 ANNALS OF THE MISSOURI BOTANICAL GARDEN A small, poorly understood genus of American terrestrial orchids at present including about six described species and several varieties ranging from Costa Rica to Colombia, Venezuela, Brazil, and Peru. А single species is known to occur in Panama in the highlands of Chiriqui Province. 1. WARREA COSTARICENSIS Schltr. іп Fedde Rep. Sp. Nov. 16:446. 1920. Erect terrestrial herbs to 70 cm. tall. Leaves 25—60 cm. long and 4—7.5 cm. broad, contracted at the base into a short, stout, sheathing petiole. Inflorescences erect racemes, equaling or slightly exceeding the leaves, produced from the sheath- ing base of the leafy petiole. Flowers relatively large, reddish bronze, the lip lighter with reddish bronze markings. Dorsal вера! oblong-ovate, obtuse, concave, 3.2-3.5 cm. long and 1.5 cm. broad, lateral sepals subequal to the dorsal sepal, the bases oblique, forming a short rounded mentum with the foot of the column. Petals obovate, obtuse, oblique, 2.8-3 cm. long and 1.3-1.5 cm. wide. Lip entire, 2.5-3 cm. long, and 2.5-3 cm. wide, suborbicular when spread out; basal callus narrow, about 1.5 cm. long. Column slender, arcuate, 2.5 cm. long, produced at the base into a foot. Costa Rica and Panama. CHIRIQUÍ: Potrerillos, 3000 ft., Kieswetter s.n.; in heavy forest near Potrerillos, Dunn s. n The present specimens differ from typical material of Warrea costaricensis in the entire, not emarginate apex of the lip, and in other minor details. Schlechter's sketch, however, does not seem to agree entirly with his description, which is in some respects nearer to our plant. Since our specimens compare in general quite well with the Costa Rican material it is thought best to consider them as slight geographical variants of an identical species, at least pending more adequate col- lections upon which to base comparisons. 45. GOVENIA Lindl. GOovENIA Lindl. in Lodd. Bot. Cab. 18: А. 1700. 1831; Benth. & Hook. Gen. РІ. 3:542. 1883. Eucnemis Lindl. Gen. & Sp. Orch. Pl. 161. 1833. Erect, terrestrial herbs, with 1 or 2 broadly plicate leaves, the basal portions forming a stout sheathing petiole and enclosing a short, stout, subconical tapering pseudobulb. Inflorescences erect, subcapitate or racemose, equaling or exceeding the leaves, the base enveloped by one of the leafless petiolar bracts. Flowers few to many, of small to medium size, borne on short or elongate pedicels, subtended by short or elongate bracts. Sepals subequal, connivent at the base, the dorsal sepal concave, erect, the laterals more or less falcate or decurved, adnate to the foot of the column, forming a mentum. Petals subequal to the lateral sepals. Lip entire, somewhat concave at the base, spreading, articulated to the foot of the column. Column arcuate, auriculate near the apex, or broadly winged throughout its (352) 1949] FLORA OF PANAMA (Orchidaceae) 17 length; base produced into a short foot. Anther terminal, operculate, incumbent, 1-celled; pollinia 4, waxy. A few American terrestrial orchids of remarkably uniform vegetative and floral structure. It seems evident that relatively trivial differences in size, color, length of the raceme, and geographic locality have been considered as adequate for the segregation of most of the so-called species. Only two entities showing any consistent structural differences are known to occur in Panama, where they seem to be confined to the Chiriquí highlands. The sterile plants greatly resemble those of Calantbe mexicana. Mte lc 1. С. CILILABIA a. Lip lanceolate, lateral margins minutely cil 4 АСЕ T A i, 2. С. LILIACEA аа. Lip broadly ovate, lateral margins пос cili 1. GOVENIA CILILABIA Ames & Schweinf. in Sched. Orch. 10:80. 1930. Erect terrestrial herbs 30-36 cm. tall. Leaves 2, broadly plicate, elliptic, shortly acuminate, 30—36 cm. long and 5.5—9 cm. broad, contracted at the base into an elongate sheathing petiole enclosed by 2—3 obtuse or very shortly acute, tubular, imbricating bracts. Inflorescence erect, the sheathing petiole enclosing a slender raceme scarcely equaling the leaves. Flowers small for the genus, mem- branaceous. Dorsal sepal elliptic-oblong, acute, concave, 10—13 mm. long and 4.2—5 mm. broad, the lateral sepals obliquely elliptic-lanceolate, acute, 9.5-10.5 mm. long and 3.5—3.8 mm. wide. Petals oblique, subfalcate, acute, 10-11 mm. long and 5—5.2 mm. wide. Lip entire, oblong-lanceolate, with minutely ciliate margins; base articulated to the foot of the column. Column arcuate, fleshy, broadly winged, dilated above, 6—7 mm. long, produced at the base into a foot. Costa Rica and Panama. CHIRIQUÍ: vic. Casita Alta, Volcán de Chiriquí, 1500—2000 m., Woodson, Allen & Seibert 947. The flowers of the single specimen bear the description: "yellow, purple striped; lip violet rose, with a white tip." 2. СОУЕМА LILIACEA Lindl. in Bot. Reg. 21: £. 1705. 1836; 24: Z. 13. 1838. Govenia utriculata (Sw.) Lindl. in Bot. Reg. 25: Misc. 47. 1839. j 788. Limodorum utriculatum Sw. Prodr. 119. 1 Cymbidium utriculatum Sw. in Nova Acta Soc. Sci. Upsal. 6:75. 1799. Govenia Powellii Schltr. in Fedde Rep. Sp. Nov. 17:51. 1922. Erect terrestrial herbs 45—65 cm. tall. Leaves 2, plicate, elliptic, shortly acute or lanceolate-acuminate, contracted at the base into a sheathing petiole, the bases enclosing a short, stout, fleshy, cylindric, tapering pseudobulb. Inflorescences erect, leafless scapes, racemose or subcapitate, less than, equal to, or exceeding the leaves, the bases enveloped by the tubular imbricating petiolar bracts. Flowers few to many. Dorsal sepal lanceolate, shortly acuminate, 10—16 mm. long and 4—5 mm. broad, lateral sepals oblique, lanceolate, subfalcate, acute, 8-11 mm. long and 4—5 mm. wide. Petals oblanceolate, acute, 10—11 mm. long and 5—6 mm. wide. Lip entire, broadly ovate, acute, 6—9 mm. long and 4.5-7.5 mm. broad, strongly (353) [Vor. 36 18 ANNALS OF THE MISSOURI BOTANICAL GARDEN nerved, the nerves often branching; base articulated to the column foot. Column short, arcuate, 7-8 mm. long, broadly winged and often subauriculate at the apex, produced at the base into a foot. Mexico, Guatemala, Salvador, Honduras, Costa Rica, Panama, the West Indies, Venezuela, Brazil, Argentina, and Bolivia. cHIRIQUÍ: in damp, shady places, 4000 ft., Powell 205; valley of the upper Río Gariché, 1050-1100 m., Seibert 320; Jaramillo, Boquete, 5000 ft., Davidson 798; Casita Alta to Cerro Copete, 2300-3000 m., Woodson 9 Scbery 345; Volcán de Chiriqui, 8000 ft., Davidson 934, 946. The last three specimens cited seem to show consistent differences from the rest in the longer racemes, the abundant linear bracts, and in the somewhat smaller flowers. In spite of this, however, no significant structural differences could be detected in the flowers. Under the circumstances it is considered best to include them here, at least until the genus as a whole is critically studied. 46. MORMODES Lindl. MonMonpzs Lindl. Introd. Nat. Syst. Bot. ed. 2, 446. 1836; Benth. & Hook. Gen. Pl. 3:552. 1883. Cyclosia Kl. in Allgem. Gartenzeit. 6:305. 1838. Epiphytic herbs, with cylindric or fusiform pseudobulbs usually tapering uni- formly upward. Leaves plicate, strongly veined, distichous, the unarmed imbricat- ing bases enclosing the pseudobulbs; leaf blades deciduous. Inflorescences arching racemes produced from the base or lower sides of the pseudobulbs. Flowers few or many, often richly colored. Sepals free, subequal, spreading or strongly reflexed. Petals subequal to the sepals or sometimes broader. Lip entire or 3-lobed, often fleshy, the lateral lobes or margins usually strongly reflexed, base often with a claw which is adnate to the base of the column. Column usually curved and twisted to one side so that the dorsal surface of the apex rests against the lip, the anther and stigmatic surface being thus exposed; rostellum without antennae, base of the col- umn without a foot. Anther terminal, operculate, incumbent, 1- or imperfectly 2- celled, extremely sensitive; pollinia 2 or 4, waxy, often ejected during the process of pressing the flowers and found adhering to the lip, or to one of the other floral parts. An interesting genus of American epiphytic orchids, ranging from Mexico to Peru and Brazil, vegetatively reminiscent of Catasetum, but readily distinguishable even when not in flower by the absence of spines on the upper margins of the im- bricating leaf bases enveloping the old pseudobulbs. Many species have been described, often founded on single specimens flowering in European greenhouses. Although many entities enter into the present confused picture, it seems probable that most represent color forms of either Mormodes igneum or M. atropurpureum. In spite of this abundance of named color forms, valid diagnostic characters seem to exist, based mostly on the lobes in the lip, whether present or absent, and their (354) 1949] А FLORA OF PANAMA (Orchidaceae) 19 relative size, shape or indument. Although even these characters are subject to а certain amount of individual interpretation, four fairly well-marked entities can be separated in Panama. Due to the extreme tortion of all of the floral parts, it is generally necessary to spread the lip out fully to obtain any adequate idea of its shape. a. Lip distinctly 3-lobed, about as long as broad; 552 йр conspicuously Һеуо ке the laterals, the apex sometimes recurved............ . M. ATROPURPUREUM aa. Lip entire, or of two lateral lobules with a shortly со арех. b. Blade of lip cllipti c-ovate to rhombic-ovate, acute or acuminate, not abruptly ко about twice аз long as broad 2. M. coLossus bb. Blade of lip obovate, oval or suborbieular, obtuse, or truncate, arrap K. about as long as bro 4. M. IGNEUM Cc г. el 3. M. HookzERI 1. Мокморвв ATROPURPUREUM Lindl. Introd. Nat. Syst. Bot., ed. 2, 446. 1836, non Hook. Epiphytic herbs with very stout pseudobulbs. Inflorescences erect racemes, produced from near the base of the pseudobulbs. Sepals subequal, free, spreading, anceolate, acute or acuminate, the dorsal sepal 2.4-2.6 cm. long and .6—7 ст. wide, laterals lanceolate, acute or acumi- nate, 2.6-2.8 cm. long and .7—8 cm. wide. Petals elliptic-lanceolate, acute, 2.5 cm. long and .9-1.1 cm. wide. Lip distinctly 3-lobed, 1.8-2 cm. long and 1.8-2 cm wide when spread out, the lateral lobes rounded and strongly reflexed, the entire mid-lobe projecting beyond the laterals, subcuneate, abruptly apiculate. Column twisted to one side. Flowers described as "red-brown, with small spots. Lip rose, spotted with red-brown." Costa Rica and Panama. Fig. 155. Mormodes atropurpureum—lip HIRIQUÍ: Caldera ке 3500 ft., Powell s. n.; Concepción, 800 ft., Powell s. n.; vic. Қ 1000 ft., Dun Seemingly rare. Most of the specimens previously whe Е as Mormodes atropurpureum Lindl. are dark red color forms of М. igneu 2. Мокмореѕ coLossus Rchb. f. in Bot. Zeit. 10:636. 1852. Mormodes macranthum Lindl. & Paxt. in ge Flow. яз 3:98. 1852-53. Mormodes Wendlandii Rchb. f. in Walp. Ann. 6:581. 18 Mormodes Powellii Schltr. in Fedde Rep. Sp. Nor. 17: 5. E Epiphytic herbs. Pseudobulbs often long, cylindric, tapering. Leaves plicate, deciduous, the persistent imbricating bases unarmed. Inflorescences long arching racemes produced from near the base of the pseudobulbs. Sepals subequal, spreading, linear-lanceolate, acuminate, 3.5—5 cm. long and .6—8 cm. wide. Petals subequal to (355) [Vor. 36 20 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN the sepals, lanceolate, acuminate, 3.2—4.5 cm. long and .7—1.0 cm. wide. Lip elliptic- ovate to rhombic-ovate, acute or acumi- : УЛ Ж ет ае = `\ Gongora maculata Fig. 172. (399) [Vor. 36 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN elongate, pendulous racemes from the base of the pseudobulbs. Flowers many, grotesque, strongly fragrant, relatively large, on elongate, slender, nearly straight pedicels. Sepals membranaceous, pale yellow, spotted or banded reddish brown (іп our specimens), dorsal sepal free, erect, the base inserted about midway on the column, lanceolate, acuminate, the margins often strongly recurved, 1.5—1.8 cm. long and .4—.6 cm. wide, the base not connivent with the bases of the lateral sepals, lateral sepals strongly reflexed, inserted on the column foot, obliquely ovate, acuminate when spread out, the lateral margins strongly recurved, 20-25 mm. long and 9-14 mm. wide. Petals colored similarly to the sepals, resembling stelidia, the bases inserted on the column and connivent with the base of the dorsal sepal, linear-lanceolate, acuminate, the attenuate apices somewhat recurved, 6—8 mm. long and 1-2 mm. broad. Lip fleshy, 1.5—2 cm. long, pale yellow, marked reddish brown (in our specimens), complexly 2-parted, the short, ligular, basal claw ap- parently articulated with the column foot; the hypochile saccate, of 2 complex, erect, lateral lobes, the upper closely appressed margins of which combine to form a dorsal keel, the hypochile when seen from above with 2 slender, lateral, ligular, somewhat recurved projections from the base, the apex in profile abruptly truncate and deeply concave, with two lateral erect elongate antennae prolonged into short acuminate teeth; the epichile laterally compressed, about as long as the hypochile, subsaccate and gibbose above the basal constriction, the apex in profile cuneate, with an attenuate recurved tip. Column erect, somewhat arcuate, 15-20 mm. long, terete below, somewhat dilated and subclavate above the point of insertion of the dorsal sepal and petals. Mexico, British Honduras to Brazil and Ecuador. CANAL ZONE: Mojinga Swamp, near the mouth of the Río Chagres, sea level, Allen 870; Ancon, in hospital grounds, sea level, Pittier 6627, 6630. PANAMA: Matías Hernán- dez, east of Panama City, sea level, Pittier 6630; hills east of Panama City, sea level, Powell 63, 70, 76, 1 76, COLÓN: forests along the Río Води uerón, above the Peluc a Hydro- каре Station, 90 m., Hunter 9 Allen 640 (in part). сос:®: El Valle de re 500— 70 , Hunter & Allen 352; El Valle Chiquito, 600 m., Руни s, n.; region north of El Valle de Anton, 800 m., Allen 2043; mountains beyond La Pintada, 400-600 m., Hunter & Allen 647. Bocas DEL токо: Water Valley, vic. Chiriqui Lagoon, von Wedel 1472. This common and widely distributed species is found in damp shaded locations at low to moderate elevations throughout our range. It seems possible that this may be the species described by Ruiz & Pavon as Gongora quinquenervis, but their description is obscure and entirely inadequate, and a photograph of the type shows it to have been based on fruiting material only; hence it has been considered best to adopt the next available name. As would be expected in a species having such a tremendous geographic range, the flowers in other areas are variable in color and to a lesser degree in size. In Panama, the flowering season is from November to May, individual plants usually producing two or more successive inflorescences. (400) 1949] FLORA OF PANAMA (Orchidaceae) 65 4 GONGORA MACULATA Lindl. var. LATIBASIs C. Schweinf. & Allen, in Bot. Mus. Leafl. Harv. Univ. 13:144. 1948. Vegetative material not seen. Pseudobulbs described as round and deeply grooved, about 5 cm. wide, the apex with 2 grayish green leaves 30 cm. long and 10 cm. wide. Sepals membranaceous, dark blood-red, the dorsal sepal erect, in- serted on the column, elliptic-lanceolate, acute, 2-2.2 cm. long and 1.2—1.4 cm. broad, the lateral sepals inserted on the column foot, strongly reflexed, obliquely ovate, shortly acuminate, the lateral margins strongly recurved, about 3 cm. long and 2 cm. broad when spread out. Petals inserted on the column, their bases con- nivent with the base of the dorsal sepal, light green spotted red, linear-lanceolate, acuminate, subfalcate, the attenuate apices incurved, 12—14 mm. long and about 2 mm. broad. Lip very fleshy, complexly 2-parted, dark blood-red; the hypochile when seen from above about 2 cm. long and 1.3 cm. broad at the abruptly truncate base, the lateral lobes broadly auriculate; the apex of the hypochile, seen in profile, with 2 erect, slender antennae prolonged below as short, acute teeth; the epichile typical of the species, except for the basal projection, which in this case is pro- longed into an almost conical spur. Panama. COLÓN: about 2 miles west of Gatun Dam, 600 ft., Butcher s. n. A very striking and well-marked variety, differing from the type in the larger, more richly colored flowers, the broader, auriculate base of the hypochile, and the prolonged subconic projection at the base of the epichile. 5. GONGORA MACULATA Lindl. var. TRICOLOR Lindl. in Bot. Reg. n. s. 10: £. 60. Gongora fulva Lindl. loc. cit. 2: t. 51. 1839. Gongora bufonia Lindl. loc. cit. 4: t. 2. 1841. Plants identical with those of the type. Sepals rich yellow to orange, usually spotted or blotched dark red or dark purple. Petals greenish yellow, usually spotted or marked dark red. Lip very fleshy, complexly 2-parted, rich yellow or orange, usually marked dark red, or dark purple. The hypochile when seen from above with a relatively broad base, with two short, fleshy, rounded horns on each side, otherwise identical with the type. Panama and Peru, and probably adjacent territory. CANAL ZONE: Mojinga Swamp, near the mouth of the Río Chagres, sea level, Allen QII. PANAMÁ: p aps east of Panama City, sea level, Powell 32, 60, 71, 93. COLON: forests along the Río Boquerón, above the Peluca Hydrographic Station, about 90 m., Hunter & Allen А (in part), 650. сосі.Е: El Valle de Antón, 1800 ft., Fairchild 5.7. Distinguished from the суре Бу the somewhat larger, more richly colored flowers, the somewhat broader base of the hypochile, and the rounded, fleshy, basal orns. (401) š [Vor. 36 66 ANNALS OF THE MISSOURI BOTANICAL GARDEN 57. CORYANTHES Hook. ConvaANTHES Hook. in Bot. Mag. t. 3102. 1831; Benth. & Hook. Gen. Pl. 3:549. 1883; Schltr. іп Orchis 10:67-82. 1916. Meliclis Raf. Fl. Tellur. 2:99. 1836. Panstrepis Raf. loc. cit. 4:41. 1836. Corynanthes Schlecht. in Bot. Zeit. 6:65. 1848. Corythantes Lem. in d’Orbigny, Dict. Hist. Nat. 4:259. 1849. Erect epiphytic herbs. Pseudobulbs slender, subcylindric, tapering, con- spicuously ridged, the apex with 2 or rarely 3 lanceolate or elliptic-lanceolate, plicate, strongly veined, subcoriaceous, acute or acuminate leaves. Inflorescences elongate, slender, pendent racemes from the base of the pseudobulbs. Flowers large, fragrant, often brightly colored, usually 2—3, rarely as many as 7, of very complex structure, on slender arching pedicels. The genus has been divided by Schlechter into two sections EuconvANTHES and LaMELLUNGUIS. In both sec- tions the sepals broad, membranaceous, free, spreading or strongly reflexed, the margins undulate or recurved, the laterals obliquely subfalcate and much larger than the dorsal sepal; the petals inconspicuous, linear-lanceolate, often subfalcately recurved at the apex, with undulate margins. Lip very fleshy, usually complexly 4-parted, the narrow basal claw continuous with the base of the column; the hypochile spreading and concave, or galeate, glabrous or pubescent; the mesochile more or less elongate and canaliculate. In EucoryANTHEs transverse lamellae or excrescences entirely lacking; in LAMELLUNGUIs, these conspicuously present. In both sections epichile inflated, very large and galeate or cup-shaped, in nature usually containing a considerable quantity of clear liquid. Column elongate, terete below, without a foot, the base with 2 short, fleshy, often subfalcate horns or glands from which the liquid is excreted, the apex inflexed-clavate and usually shortly 2-alate on either side of the clinandrium. Anther terminal, operculate, incumbent, 2-celled; pollinia 2, waxy. About 20 species of American epiphytic plants ranging from Guatemala to Peru and Brazil. They usually occur as a conspicuous element in the unique arboreal myrmecophyllous gardens, in the nests of ants of the genera Camponotus and Azteca, the association often including a purple- or orange-flowered, erect, tufted Epidendrum апа several apparently specialized succulent-leaved non- orchidaceous plants, among the most frequent being Peperomias and members of the Gesneriaceae. The flowers are probably the most complex and fascinating of the entire Orchidaceae, every detail of the floral structure having been profoundly modified to attract insects and to assure cross-fertilization To accomplish insect attraction, the flowers, although lasting only 3—4 days, are usually brightly colored, large, and strongly fragrant, secreting some substance on the inner margins of the epichile which is extremely attractive to hymenop- terous insects. The sepals, although large, soon wither, the direction of the insect's (402) 1949] FLORA OF PANAMA (Orchidaceae) 67 path being left almost entirely to the intricate contrivances of the marvelous labellum. This organ is unique in the Orchidaceae, the apical lobe or epichile re- sembling an inverted helmet or waxy cup, on one side prolonged below the apex of the column into a short, usually 3-cornute spout-like channel. The column, in its turn, is sharply reflexed at the apex, which rests precisely over the channel, exposing the anther and stigmatic surface to any insect seeking a way out from the interior of the lip. It would seem that the broad expanse of the mouth of the epichile would pro- vide ample space for an insect to take flight, with only an occasional individual choosing the lateral exit. If the flowers were of longer duration, this might have been left to pure chance, but since time is also an important factor, this possibility of escape by flight has been ingeniously circumvented by the plant in the following manner. Above the cup, on the base of the column, 2 fleshy glands have been developed. When the flowers are fully open and numbers of bees have been at- tracted to gnaw at the inner margins of the cup, these glands begin secreting a clear liquid, drop by drop, filling the bottom of the cup to the level of the apical channel. Any bee that loses its footing in the competing swarm on the upper margins is precipitated into this liquid, and once its wings have been wet, it has no other choice than to force its way out through the narrow channel below the stigma and anther. Thus the first bee to make the circuit receives the pollinia firmly cemented to its back, to be inserted on the stigma of this or another flower, on the next turn through. А single species of this most remarkable of all orchid genera has thus far been found in Panama. 1. CORYANTHES MACULATA Hook. in Bot. Mag. t. 3102. 1831. Coryantbes maculata Lindl. in Bot. Reg. 21: f. 1703. 1835. Coryanthes Albertinae Karst. Ausw. Neuer Gew. Venez. 5:1. I. 1848. Coryantbes splendens Barb. Rodr. Gen. & Sp. Orch. Nov. 1:103. 1877. Coryantbes Hunteriana Schltr. in Fedde Rep. Sp. Nov. Beih. 17:63. 1922. Erect epiphytic herbs with elongate, subcylindric, tapering, strongly ridged pseudobulbs 6.5-15 cm. long and 2.5—4 cm. wide, bearing at the apex 2 firm, lanceolate, plicate, strongly veined, acute or acuminate leaves 30—60 cm. long and 4-10 cm. wide. Inflorescences slender, elongate, pendent racemes 30—50 ст. long, from the base of the pseudobulbs. Flowers usually 2—3, large and attractive, of variable color, the lip complexly 4-parted. Sepals membranaceous, free, clear yellow with a few purple spots, pale purple, or reddish-brown, dorsal sepal sub- orbicular-ovate, broadly acute and minutely apiculate, the apical half strongly reflexed, the lateral margins revolute, 2—2.5 cm. long and 3—3.5 cm. broad when spread out, lateral sepals very strongly reflexed, from a broadly lobulate base, obliquely subfalcate, or the pair when spread out obliquely dolabriform, 5—6.5 cm. long and 2.5-4 cm. broad. Petals membranaceous, usually colored similarly to the sepals, ligular-lanceolate, the apices obliquely subacute, the margins undulate, (403) [Vor. 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN 68 WDY УУУУ) УХ “ NS SH) SD > Dy) DM, у) тата! ULA Nn BENZ == oF слог 7 SE РВ T p My Ex >> y) ) yy ))5» DD mm NS № 22222 ЕР 2% Coryantbes maculata (404) Fig. 173. > Ns See оо b : 22 рч че мар SAP MG RENA я e a a A E Р 1949] FLORA OF PANAMA (Orchidaceae) 69 3—3.5 cm. long and .6-1.0 cm. wide. Lip very fleshy, complexly 4-parted, clear waxy yellow, pale purplish spotted with blood red, or the hypochile yellow, the mesochile and epichile reddish brown on the outer surfaces, the inner epichile white spotted red-brown. Claw of the lip semiterete, continuous with the base of the column; hypochile broadly galeate, glabrous, the apex obtuse or subaoute, in profile 2-2.5 cm. long and 2—2.5 cm. broad; mesochile canaliculate, glabrous, the basal half covered by the hypochile, in profile 2.5—3 cm. long and .8-1.5 ст. wide; epichile deeply cucullate or galeate, broadly obtuse, seen in profile 3—3.5 cm. long and 2.5—3.5 cm. broad, in fresh material resembling а waxy сир or bucket, pro- longed below the apex of the column into a 3-cornute spout or channel. Column subterete, without a foot, near the base with 2 broad, lateral, subfalcate or ob- liquely auriculate glands, 8—10 mm. long and 5-10 mm. wide, the apex strongly reflexed and shortly 2-alate on either side of the clinandrium. Panama, Venezuela, the Guianas, Brazil, and probably other adjacent territory. CANAL ZONE: Río Indio, near Gatün, sea level, Bue s. n., Allen 2054. РАМАМА: hills east of Panama City, sea level, Powell 19, 156, In Panama the plants are frequently found in the tops of slender trees, in ants' nests, often associated with Epidendrum imatophyllum. They are well protected by the belligerent ants and are painful subjects to collect, and still more painful to transport. Moreover, they seldom thrive in cultivation, possibly from lack of some essential element contributed by the ants in their natural association. It seems possible that this may be only a well-marked variety of Coryantbes speciosa. 58. XYLOBIUM Lindl. XvLoBIUM Lindl. in Bot. Reg. 11:sub Z. 897. 1825; Benth. & Hook. Gen. РІ. 3:547. 1883. Onkeripus Raf. Fl. ber 4:42. 1836. Pentulops Raf. loc. cit. 1836. Epiphytic herbs with short or elongate, ovoid or semiterete pseudobulbs, the bases enveloped in 2—4 papery, brown, imbricating bracts, the apices with 1—2 lanceolate, plicate, subcoriaceous, strongly veined, acute or acuminate leaves. In- florescences short or elongate, erect or arching racemes from the base of the pseudobulbs. Flowers few to many, on short pedicels, reminiscent of those of a small-flowered Maxillaria. Sepals subequal, membranaceous, spreading, the dorsal sepal free, the laterals broader, adnate to the foot of the column, forming a short mentum, the dorsal surface, particularly near the apex, usually with an erect central keel. Petals membranaceous, subequal to the dorsal sepal. Lip entire or 3-lobed, the lateral lobes or margins erect, the base articulated with the foot of the column; disk smooth, callused or with parallel prominent nerves or lamellae. Column short, erect, somewhat arcuate, semiterete, the apex subclavate or nar- rowly winged, the base produced into a foot. Anther terminal, operculate, incum- bent, 1-celled or imperfectly 2-celled; pollinia 2 or 4, waxy. (405) [Vor. 36 70 ANNALS OF THE MISSOURI BOTANICAL GARDEN About 20 species of American epiphytes, found from southern Mexico to Peru and Brazil, and in the West Indies. The flowers considerably resemble those of some species of Maxillaria. However, the genus is amply separated by the racemose flowering habit and the plicate rather than coriaceous leaves. Three species are known from Panama. a. Lip conspicuously 3-lobed. . Lip densely рарШозе. Pseudobulbs elongate, semi-terete, diphyllous.... 1. X. ELONGATUM bb. Lip not papillose. Pseudobulbs ovoid, stout, di- or triphyllous............ 2. X. FOVEATUM gie entire or obscurely 3-lobed. oe short, и. ophyllous ог very rarely diphyllou . X. POWELLII 1. XvLoBIUM ELONGATUM (Lindl.) Hemsl. Biol. Centr.-Amer. Bot. 3:252. 1885. Maxillaria elongata Lindl. & Paxt. in Paxton's Flow. Gard. 3:69, fig. 264. 1852—53. Epiphytic herbs with erect, elongate, semiterete pseudobulbs 10—27 cm. long and .4-1.2 cm. wide, the bases enveloped in 3—4 brown, papery, imbricating bracts, the apices with 2 lanceolate or elliptic-lanceolate, plicate, strongly veined, acute or acuminate leaves 15-36 cm. long and 2.5-9 cm. wide. Inflorescences erect, slender racemes 10—15 cm. long, produced from the base of the pseudobulbs, the lower portion below the flowers enveloped in several broad, papery, closely imbricating bracts. Flowers few to many, comparatively large, on short pedicels. Sepals membranaceous, white or pale yellow, spreading, subequal, the dorsal sepal free, lanceolate, acute or acuminate, 15-25 mm. long and 4-7 mm. broad, lateral sepals adnate at the base to the column foot, forming a short mentum, somewhat obliquely lanceolate, acuminate, with a strong central keel on the dorsal surface, 15-30 mm. long and 4—8 mm. wide. Petals membranaceous, white or pale yellow, spreading, subequal to the dorsal sepal, lanceolate, acuminate, 15—20 mm. long and 4—7 mm. wide. Lip 3-lobed, articulated at the base with the foot of the column, 14-18 mm. long and 4—8 mm. broad, white or pale yellow, with purple or maroon stripes, lateral lobes conspicuous, erect, about equaling the column, apical lobe densely papillose, the lateral margins strongly inflexed; disk with 5 thickened cen- tral nerves or lamellae. Column short, stout, terete, produced at the base into a long broad foot. Anther terminal, operculate, incumbent, 1-celled; pollinia 4, waxy. Costa Rica and Panama. cocLÉ: hills north of El Valle de Antón, 1000 m., Allen 2705, 2745, 3739. CHIRIQUÍ: near Boquete, 3800—4000 ft., Powell 167; Bajo Mono-Robalo trail, western slopes of Cerro Horqueta, 5000—7000 ft., Allen 4787 A frequent plant of the highland forests above about 2500 ft., readily dis- tinguished from the other local species by the extremely elongate, slender, di- phyllous pseudobulbs, and the densely papillose lip. Usually to be found in damp situations in heavy shade, often on the lower trunks of large trees. The flowering season is during the fall and winter months, from September to March. (406) 1949] FLORA OF PANAMA (Orchidaceae) 71 2. XYLOBIUM FOVEATUM (Lindl.) Nicholson, Dict. Gard. 4:225. 1887. Maxillaria foveata Lindl. in Bot. Reg. n. s. 2: Misc. 2. 1839. Maxillaria concava Lindl. loc. cit. 7: Misc. 4. 1844. Maxillaria stachybiorum Rchb. f. in Bot. Zeit. 10:735. 1852. Xylobium concavum Hemsley, Biol. Centr.-Amer. Bot. 3:252. 1883. Xylobium stachybiorum Hemsley, loc. ск. 1883. Xylobium Filomenoi Schltr. in Fedde Rep. Sp. Nov. Beih. 9:100. 1921. Epiphytic herbs. Pseudobulbs stout, ovoid or subconic, tapering, smooth or somewhat ridged, 4—9 cm. long and 1.5—4 cm. wide, the base enveloped in 2—3 brown, fibrous, imbricating bracts, the apex with 2-3, plicate, lanceolate, strongly veined, acute or acuminate, subcoriaceous leaves 24-40 cm. long and 2.5—7 cm. wide, the blades usually deciduous after the second year, the apex of the old pseudobulbs unarmed. Inflorescences erect racemes 12-30 cm. tall, from the base of the pseudobulbs, the lower rachis below the flowers enveloped in several broad, papery bracts. Flowers usually many, fragrant, on short pedicels. Sepals mem- branaceous, subequal, spreading, creamy white, the dorsal sepal free, lanceolate- acuminate, 10-14 mm. long and 3—4 mm. wide, laterals adnate to the foot of the column, forming a mentum, subfalcately lanceolate-acuminate, 12—14 mm. long and 3-4 mm. wide, the dorsal surface with a prominent central keel. Petals subequal to the dorsal sepal, membranaceous, creamy white, lanceolate-acuminate, 8—10 mm. long and 2-3 mm. wide. Lip creamy white with reddish veining on the disk, conspicuously 3-lobed, 10-12 mm. long, articulated at the base with the foot of the column, the lateral lobes rounded, erect, about equaling or somewhat exceeding the column, apical lobe obtuse, spreading; disk with 5 thickened nerves or keels. Column very short, semiterete, the apex obtuse; the base produced into a long broad foot. Anther 1-сеПед; pollinia 4. Mexico to Peru and British Guiana; Jamaica. CANAL ZONE: Barro Colorado Island, Aviles 32. PANAMA: Ceiba Tierra, sea level, Powell 3060; Ana Lago, sea level, Powell 3064; hills east of Panama City, sea level, Powell 27; Chiva-Chiva, sea level, Powell 3053; vic. Pacora, sea level, Powell 3026, Allen 821; forest along telephone cable, vic. Rio Indio Hydrographic Station, Upper Madden region, Steyermark & Allen 17466; San José Island, Perlas Archipelago, Johnston 1017, 1117. A very frequent species found throughout the lowlands of our area, flowering during the fall and winter months from November to March. 3. Хүговюм PowELLI Schltr. in Fedde, Rep. Sp. Nov. Beih. 17:66. 1922. Xylobium sublobatum Schltr. loc. cit. 19:51. 1923. Erect epiphytic herbs with slender subterete (rarely cylindric-tapering) pseudobulbs 4-7 cm. long and .7-1.8 cm. wide, enveloped at the base in 2—3 fibrous, imbricating bracts, the apex with 1 (rarely 2) lanceolate, plicate, strongly veined, long-petioled, acute or shortly acuminate leaves 20-60 cm. long and 2—5.5 cm. wide. Inflorescences erect racemes, 9—15 cm. tall, the bases below the flower- (407) [Vor. 36 72 ANNALS OF THE MISSOURI BOTANICAL GARDEN ing portion enveloped in several papery, imbricating bracts. Flowers few to many, on short pedicels, yellow or tan, sometimes tinged with light green. Sepals mem- branaceous, subequal, spreading, the dorsal sepal elliptic-lanceolate, acute, 10—12 mm. long and 4—6 mm. wide, laterals adnate to the foot of the column, forming a mentum, from an oblique base, lanceolate, acute, 12-14 mm. long and 4—6 mm. wide, the dorsal surface with a prominent central keel. Petals subequal to the dorsal sepal, linear-lanceolate, acute, 10-12 mm. long and 3—4 mm. broad. Lip entire or obscurely 3-lobed, articulated at the base with the column foot, 10-12 mm. long; the narrow lateral lobes, if present, or the margins erect; the apical lobe somewhat concave, subacute; disk with 3 prominent erect nerves or keels. Column very short, subterete, produced at the base into a foot. Anther 1-celled; pollinia 4. Costa Rica and Panama. Peña Blanca, 1750-2000 m., Woodson & Schery 325; Quebrada Velo, 1800 m., Woodson 9 Schery 281; vic. Bajo Mono and Quebrada Chiquero, 1500 m., Woodson ë Scbery 534; Bajo Chorro, 6000 ft., Davidson 241. A very frequent species of the Chiriqui highlands, flowering in July and August. Although the type specimen of Xylobium Powellii was described as having two leaves, nearly all plants seen had a single leaf. Туре material has not been available for comparison, but our species seems to agree very well with an earlier monophyllous species from Guatemala, Xylobium Tuerckheimii Kranzl., and it seems likely that subsequent investigation may prove them to be identical. 59. BIFRENARIA Lindl. BIFRENARIA Lindl. Gen. & Spec. Orch. Pl. 152. 1833; Benth. & Hook. Gen. Pl. 3:546. 1883. Adipe Raf. Fl. Tellur. 2:101. 1836. Stenocoryne Lindl. in Bot. Reg. n. s. 6: Misc. 53. 1843. Lindleyella Schltr. Die Orchideen, p. 414. 1914. Epiphytic herbs. Pseudobulbs short, fleshy, ovoid, subconic or subpyramidal, often abruptly 4-angulate or laterally compressed, the bases enveloped in 2—3 fibrous, imbricating bracts, the apex with 1-2 lanceolate or elliptic-lanceolate, plicate, strongly veined, acute or acuminate leaves. There are two quite distinctive groups of species, one of which has been segregated by Schlechter as a separate genus under the name Lindleyella. Schweinfurth (Bot. Mus. Leafl. Harv. Univ. 11:246. 1944) has pointed out that the floral characters proposed by Schlechter for such separation are equally applicable to species of the other group, so that it is impossible to accept Lindleyella as a valid generic concept. However, since the groups are sufficiently distinctive to be separated on gross characters alone, the name proposed by Schlechter is retained as a section, simply for purposes of con- venience. On this basis, those species superficially resembling Bifrenaria atropur- фиғеа, the generic type with 1—3 large flowers on short erect racemes, are considered (408) 1949] FLORA OF PANAMA (Orchidaceae) 73 as representing the section EUBIFRENARIA, while those resembling Bifrenaria picta and B. aurantiaca, with many small flowers on elongate arching racemes, аге con- sidered as representing the section LINDLEYELLA. In both sections of the genus, the sepals are free, subequal, spreading, the laterals adnate to the column foot, forming a short or elongate spur-like mentum. Petals subequal to the dorsal sepal or smaller. Lip 3-lobed, articulated at the base with the foot of the column, lateral lobes erect, the mid-lobe spreading, entire or bifid; disk fleshy, lamellate, denticulate or hirsute. Column erect, thickened, semiterete, wingless, produced at the base into an elongate foot. Anther terminal, operculate, incumbent, strongly convex, sometimes cristate on the dorsal surface, 1-celled or imperfectly 2-celled; pollinia 2 or 4, waxy. A variable genus of mostly Brazilian epiphytes, a few species ranging to Peru, the Guianas, Trinidad, Venezuela, and Colombia, with a single species of the sec- tion LINDLEYELLA known from eastern Panama. 1. ВЕКЕМАВЈА PICTA (Schltr.) C. Schweinf. in Bot. Mus. Leafl. Harv. Univ. 11:246. 1944. Lindleyella picta Schltr. іп Fedde Rep. Sp. Nov. Beih. 27:173. 1924. Epiphytic herbs. Pseudobulbs ovoid, laterally compressed, somewhat ridged, 2.5-5 cm. long and 2-2.5 cm. wide, enveloped at the base in 2-3 fibrous imbri- cating bracts, the apex with a single, long-petiolate, elliptic-lanceolate, plicate, strongly veined, shortly acuminate leaf 38—46 cm. long and 5-6.8 cm. wide. Inflorescences elongate, slender, arching racemes produced from the base of the pseudobulbs. Flowers many, relatively small, on short pedicels, typical of the section LINDLEYELLA. Sepals subequal, spreading, membranaceous, deep red, or red marked with yellow particularly at the base, dorsal sepal free, elliptic- lanceolate, acute, 10-14 mm. long and 3-5 mm. wide, lateral sepals somewhat oblique, elliptic-lanceolate, acute, adnate to the foot of the column, 10—14 mm. long and 4-8 mm. wide. Petals subequal to the dorsal sepal, yellow, spotted red, elliptic-lanceolate, acute, 10-12 mm. long and 4-5 mm. wide. Lip 3-lobed, 10—12 mm. long, red marked with yellow, the base shortly clawed, lateral lobes erect, subfalcate, obliquely obtuse, longer than broad, the anterior margin undulate and crisped; disk between the lateral lobes with a convex, rather obscurely verrucose callus; mid-lobe subreniform or subquadrate, broadly spreading, contracted at the base into an isthmus with a short, suberect, fleshy, obtuse, linguiform callus, the apex obscurely bilobulate, shallowly emarginate, the lateral lobules undulate and crisped. Column semiterete, somewhat arcuate, 6-8 mm. long, produced at the base into a foot. Anther imperfectly 2-celled; pollinia 2, subglobose. Panama and Colombia. DARIÉN: Chepigana District, Cana-Cuasi trail, 800 ft., Terry 6 Terry 1616. The only record for the genus in North America. (409) [Vor. 36 74 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN 60. LYCASTE Lindl. LvcasrE Lindl. in Bot. Reg. Misc. n.s. 6: Misc. 14. 1843; Benth. & Hook. Gen. Pl. 3:547. 1883. Перріа Raf. Fl. Tellur. 2:51. 1836. Erect, epiphytic or sometimes pseudo-terrestrial herbs. Pseudobulbs short, fleshy, ovoid or ellipsoid, tapering, often laterally compressed, smooth or pluri- sulcate, the bases enveloped in several closely imbricating, papery or coarsely fibrous bracts, the upper 1 or 2 of which are usually foliaceous; the apex with 1 to several, usually broad, plicate leaves which are ultimately deciduous, the apex of the old pseudobulbs with or without 2 sharp marginal spines; roots fibrous, often pubescent or conspicuously lanuginose. Inflorescences 1 to many erect, stout or filiform, single-flowered scapes from the base of the pseudobulbs, enveloped in several closely imbricating or distant, tubular or spathaceous, papery bracts. Flowers small to large, usually more or less nodding. Sepals subequal, membra- naceous, spreading, the laterals somewhat broader than the dorsal sepal, sometimes somewhat reflexed, adnate at the base to the foot of the column, forming a short mentum. Petals subequal to the dorsal sepal or shorter. Lip conspicuously or obscurely 3-lobed, the base continuous with, or articulated with, the foot of the column, lateral lobes or margins erect, mid-lobe spreading or reflexed; disk with a thickened callus. Column slender, semiterete, rather arcuate, produced at the base into a foot. Anther terminal, operculate, incumbent, 1-celled; pollinia 4, waxy. A rather perplexing genus of perhaps 25 species of American epiphytes, ranging from southern Mexico to Peru, Brazil, and the West Indies. Although many of the species have been put at one time or another into the allied genus Maxillaria, they are readily separable by the plicate, rather than conduplicate leaves. Five species are known to occur in Panama. a. Apex of the old pseudobulbs armed with 2 conspicuous sharp spines. .. коне after the leaves have fallen, ог еге with the о flush о growth. Pseudobulbs ас flowering time without mature 2 b. Lip рени out about as long ав broad; apical lobe without а basal constric or . L. BREVISPATHA . 2. L. CAMPBELLII bb. Lip when spread out оер twice as long аз broad; apical lobe with a basal constrictio . L. TRICOLOR aa. РА of the old peudobuli P^ суд with 2 жен н те Flowers produced at the f the current season's growth, r the pseudobulbs have MM e before pon leaves bu fallen b. Lip conspicuously 3-lobe ore еф — as long as kasi when spread out; apical lobe ovate or subquadrate - 3. L. MACROPHYLLA bb. Lip obscurely 3-lobed, less than twice as јан as broad when spread out; apical lobe broadly obtuse, separated from the laterals by pheste сс. St S а et ОНЫ . L. POWELLII (410) 1949] FLORA OF PANAMA (Orchidaceae) 75 1. LYCASTE BREVISPATHA КІ. ex Lindl. in Paxton’s Flow. Gard. 3:44. 1852-53. Lycaste candida Lindl. ex Rchb. f. in Walp. Ann. 6:604. 1863. Epiphytic (rarely pseudo-terrestrial) herbs. Pseudobulbs fleshy, usually ellip- tic-ovoid, rarely suborbicular, laterally compressed, 4.5-6.5 cm. long and 2-3.5 cm. wide, enveloped at the base in several coarsely fibrous bracts, the apex with 2—4 elliptic-lanceolate, plicate, acute or shortly acuminate leaves which are ulti- mately deciduous, 15—20 cm. long and 3—5 cm. wide, the apex of the old pseudo- bulbs armed with 2 conspicuous sharp spines; roots usually conspicuously lanuginose. Inflorescences 1 to many slender, erect scapes 5—7 cm. tall, usually produced concurrently with the flush of new growth from the base of the old leafless pseudobulb, the filiform rachis provided with several distant tubular sheath- ing bracts, the apical bract usually about half the length of the ovary. Flowers more or less pseudo-campanulate, solitary, somewhat nodding. Sepals subequal, membranaceous, usually pale green to olive-green, rarely rose, the dorsal sepal free, erect, elliptic-lanceolate, acute, the apex strongly recurved, 2.5-2.8 cm. long and 10—15 mm. wide, laterals adnate to the foot of the column, forming a short mentum, from the oblique base elliptic-lanceolate, acute, the apices strongly re- curved, 2.5-3 cm. long and 1.0-1.5 cm. wide. Petals subequal to the sepals, white marked rose, or rarely pure white, elliptic-oblanceolate, acute, the apices recurved, 2.4-2.6 cm. long and 1.0-1.5 cm. wide. Lip 3-lobed, articulated at the base with the foot of the column, white with rose markings, or rarely white, nearly as broad as long when spread out, 2.2-2.5 cm. long and 1.8-2.2 cm. wide, lateral lobes erect, rather obscurely emarginate at the subacute apices, mid-lobe broadly obtuse, the apex somewhat reflexed; disk with an elongate, acute, 2-costate lingui- form callus. Column semiterete, rather arcuate, 8—9 mm. long, the under-surface conspicuously pubescent, the base produced into a foot. Costa Rica and Panama. RIQUÍ: without definite locality, shady places, 4000-4500 ft., Powell 140; vic. Finca ern upper forested margins of the Quebrada Velo, 5000 k Allen 4749. A fairly frequent species of the Chiriquí highlands, usually being found in shaded places between 4000 and 5000 ft. elevation. The flowers are variable in color, usually being produced from February to about April. 2. LYCASTE CAMPBELLI C. Schweinf. in Johnston, Sargentia 8:103, fig. 1949. ined. Epiphytic herbs with ovoid-ellipsoid, somewhat laterally compressed pseudo- bulbs 3.5-5 cm. long and 1.5-2.5 cm. wide, enveloped at the base in several coarsely fibrous, imbricating sheaths; apex of the pseudobulbs (during the growing season) with 1—3 plicate, elliptic-lanceolate leaves which are ultimately deciduous, the old pseudobulbs armed at the apex with 2 sharp spines; roots fibrous, lanuginose. Inflorescences 1 to several erect, filiform scapes less than 8 cm. tall, produced from the base of the current pseudobulb after the leaves have fallen. Flowers pseudo- campanulate, more or less nodding, the smallest of the genus in Panama. Sepals (411) [Vor. 36 76 ANNALS OF THE MISSOURI BOTANICAL GARDEN subequal, membranaceous, green, the dorsal sepal ovate-elliptic, acute, about 1.8 cm. long and 1 cm. wide when spread out, laterals adnate to the foot of the column, forming a short conical mentum, from the oblique base oblong-ovate, acute, 1.8 cm. long and about 1 cm. wide. Petals elliptic-ovate, obtuse, abruptly apiculate, yellowish green, about 1.5 cm. long and 1 cm. wide. Lip 3-lobed, contracted at the base and articulated with the foot of the column, yellow, about 1.6 cm. long and .9—1.0 cm. wide when spread out, lateral lobes erect and somewhat incurved, the apices rather obliquely acute, mid-lobe ovate, subacute, with a recurved apicule; disk between the lateral lobes with a concave oblong-obtuse callus. Column short, stout, semiterete, somewhat arcuate, about 6-8 mm. long; obscurely puberulent on the under-surface, produced at the base into a foot. Panama. PANAMÁ: San José Island, Perlas "aia ИН high on trees in stream-side forest in area 11-B, Johnston 1371, Сеты Пт Known only from the type collection. Although vegetatively reminiscent of Lycaste candida or L. tricolor, it may readily be distinguished by the much smaller, differently colored flowers. "Тһе season of flowering is given Ьу Dr. Johnston as February. 3. ГУСАЗТЕ MACROPHYLLA Lindl. in Bot. Reg. n. s. 6: Misc. 14. 1843. Maxillaria macrophylla Poepp. & Endl. Nov. Gen. & 1:37, Ё. 04. 1836. Lycaste plana Lindl. in Bot. Reg. п.5. 5: Misc. 85. 184 Lycaste Dowiana Endres & Rchb. f. in Gard. Chron, n. 5. 22: 194. 1874. Lycaste Filomenoi Schltr. in Fedde, Rep. Sp. Хоу. Beih. 9:100. 1921. Epiphytic herbs with stout, ovoid, somewhat laterally compressed, often rather angulate or plurisulcate pseudobulbs 4—10 cm. long and 3—6 cm. wide, the bases enveloped in 3—4 closely imbricating, papery bracts, the upper 2 of which are foliaceous, the largest blade about 40 cm. long and 10—12 cm. broad; the apex of the pseudobulbs with 2—3 broad, plicate, oblanceolate, acute or acuminate leaves 40—75 cm. long and 10—12 cm. broad, which are ultimately deciduous; the apices of the old pseudobulbs not armed with sharp spines. Inflorescences few to many erect, single-flowered scapes 8—18 cm. tall, from the base of the mature pseudo- bulbs, the rachis enveloped in several broad, papery, spathaceous bracts, the upper- most of which is usually broadly cucullate, acute, about equaling or somewhat exceeding the ovary. Flowers large and conspicuous, fragrant, more or less nod- ding. Sepals subequal, broadly spreading, olive-green, sometimes shaded on the margins with reddish-brown, the dorsal sepal free, erect, lanceolate to elliptic- lanceolate, acute, 4.5-6 cm. long and 1.5-2.5 cm. wide, laterals adnate to the foot of the column, forming a short mentum, from the oblique base lanceolate to ellip- tic-lanceolate, acute, 4.5—6 cm. long and 2-3 cm. wide. Petals subequal to the sepals, not spreading, more or less parallel with the column, white, often spotted rose-pink, linear-lanceolate to elliptic-oblanceolate, acute, the apices recurved, 4—5 cm. long and 1.8—2.2 cm. broad. Lip 3-lobed, 3—4.5 cm. long and 1.4-1.6 cm. wide when spread out, usually not equaling the petals, narrowed at the base and (412) 1949] FLORA OF PANAMA (Orchidaceae) 77 articulated with the foot of the column, white, usually with the margins of the apex spotted or blotched rose-red, lateral lobes erect, the apices rather irregularly oblique, mid-lobe spreading, ovate to subquadrate, acute or obtuse, the margins slightly ciliate and incurved; disk with a fleshy, lanceolate, acute, concave callus. Column slender, semiterete, somewhat arcuate, white, 18-23 mm. long. Costa Rica, Panama, Colombia, Venezuela, Peru, and Bolivia. CocLÉ: region north of El Valle de Antón, 3000 ft., ај 2665, 5067, 5130; western slope and summit of кори Valle Chiquito, 700-800 m., Seibert 644. VERAGUAS: for slopes of Cerro Тисе, of Santa Fé, 2500—3000 ft., Allen 3 Fairchild 4400. adt without definite EC 000 ft., Powell 246. A frequent, rather variable species of the wet highland forests of Coclé, Veraguas, and Chiriquí provinces. The Coclé specimens are rather consistently smaller than those from Veraguas and Chiriquí, particularly in the vegetative parts, but otherwise appear to be very nearly identical. The Coclé form is usually considered as representing Lycaste Dowiana, but the supposed differences (shorter floral bract and narrower lip) have been found on careful examination to be inconstant characters, scarcely any two specimens being exactly alike, and, in any event, hardly differences warranting specific segregation. From the evidence now available, the Coclé plants appear to be at most a local form of the widespread and variable Lycaste macrophylla. The flowering season is from March to about July. 4. Lycaste PowrrLn Schltr. in Fedde Rep. Sp. Nov. Beih. 17:65. 1922. Epiphytic, or often pseudo-terrestrial herbs with elliptic-ovoid, laterally com- pressed, smooth, or somewhat ridged pseudobulbs 3—6 cm. long and 2-3 cm. wide, the bases enveloped in 3—4 papery, closely imbricating bracts the upper 2 of which are usually foliaceous; the apices of the pseudobulbs with 2—3 plicate, lanceolate or elliptic-lanceolate, acute or acuminate, ultimately deciduous leaves 20—35 cm. long and 4—7 cm. broad; the apices of the old pseudobulbs unarmed. Inflorescences 1 to about 4, slender, erect, 1-flowered scapes 5—14 cm. tall, produced from the bases of the mature pseudobulbs before the leaves have fallen, the rachis provided with several distant, papery, tubular, acuminate bracts the uppermost of which is spathaceous and conspicuously exceeds the ovary. Flowers relatively large, very fragrant. Sepals widely spreading, pale translucent green, heavily blotched with chestnut-brown, or wine-red with yellow margins, dorsal sepal free, linear-lanceo- late, acute, 3—4 cm. long and .8—1.4 cm. wide, the apex lightly recurved, laterals adnate to the foot of the column, forming a short mentum, from the oblique base, linear-lanceolate, acute, 3—4.2 cm. long and .6—1.4 cm. wide. Petals subequal to the sepals, cream-yellow to nearly white, spotted rose-pink or wine-red, elliptic- lanceolate, obtuse or subacute, 2.5—3.5 cm. long and 1.2-1.8 cm. wide, more or less parallel with the column, the apices strongly reflexed. Lip elliptic-obovate, obtuse or subacute when spread out, 2.6-2.8 cm. long and 1.4-1.7 cm. wide, ob- scurely 3-lobed, the lateral lobes or margins erect, the mid-lobe short, spreading, obtuse or subacute, separated from the laterals by plicate folds; disk with a short, (413) [Vor. 36 78 ANNALS OF THE MISSOURI BOTANICAL GARDEN fleshy, ligular, obtuse, concave callus. Column semiterete, 1.8—2 cm. long, slight- ly arcuate, minutely papillose. Panama Қ аса omy east of Panama City, Powell 15; Cerro Campana, 2500 ft., Fairchild COCLE: . El Valle de Antón, floor and forested ravines in dry hills to RN south, 600-800 m., Allen 761, 2666; floor of El Valle de Antón, 600 m., Fairchild 5 A frequent and attractive species of the wooded ravines of the intermediate highlands of Cerro Campana and El Valle de Antón, where they are often found growing on rocks or on low, gnarled trees of the genus Coccoloba. The flowering season is usually from July to September. 5% 5 У SZ э» E rra. "ists АЛ Fig. 174. Lycaste tricolor (414) 1949] FLORA OF PANAMA (Orchidaceae) 79 5. LvcasrE TRICOLOR (КІ.) Rchb. f. in Walp. Ann. 6:603. 1863. Maxillaria tricolor Kl. in Allgem. Gartenzeit. 20:186. 1852. Lycaste Bradeorum Schltr. in Fedde Rep. Sp. Nov. Beih. 19:138. 1923. Epiphytic herbs with elliptic-ovoid, tapering, laterally compressed, smooth or ridged pseudobulbs 6—9 cm. tall and 2.5—3 cm. wide, the bases enveloped in several coarsely fibrous bracts, the apex (during the growing season) with 3—4 elliptic- lanceolate, plicate, acute or acuminate, deciduous leaves; the apex of the old pseudobulbs armed with 2 short, sharp spines; roots fibrous, lanuginose. Іп- florescences 1 to several erect, single-flowered scapes 9—12 cm. tall, from the base of the pseudobulbs, produced after the leaves have fallen or concurrently with the flush of new growth. Flowers pseudo-campanulate, relatively large. Sepals spread- ing, subequal, the apices strongly recurved, pale green, often spotted with pink, the dorsal sepal free, elliptic-lanceolate, acute, often minutely apiculate, 2.8—3.2 cm. long and 1.2-1.5 cm. wide, lateral sepals adnate at the base to the foot of the column, forming a short mentum, from the oblique base lanceolate, acute, often minutely apiculate, 2.8—3.2 cm. long and 1.0-1.4 cm. wide. Petals subequal to the sepals, spreading, the apices strongly recurved, white, often spotted pink, elliptic-oblanceolate, acute, 2.4-2.6 cm. long and 1.4-1.6 cm. wide. Lip con- spicuously 3-lobed, about twice as long as broad when spread out, narrowed at the base and articulated with the foot of the column, white, usually spotted pink, 2.8-3 cm. long and 1.4-1.6 cm. wide when spread out, lateral lobes erect, the anterior margins obscurely emarginate at the obliquely acute apices, mid-lobe sub- quadrate or ovate, obtuse, with a basal constriction, the apex strongly reflexed; disk with an elongate, acute, concave callus. Column semiterete, somewhat arcuate, 12—14 mm. long, produced at the base into a foot. doe Costa Rica, and Panama, and probably adjacent territory. Río a Ahogado, 1200 ft., Fairchild 5.п.; Río Mata Ahogado, volcanic badlands 442 Ре El Valle de Antón, 1500 ft., Allen 4526. УЕКАСОА8: Río Santa María, vic. Santa Fé, 1000 ft., Allen 4567. A frequent and attractive species often found on low-spreading trees over- hanging streams in the volcanic badlands of Coclé and Veraguas provinces, usually at about 1000-1500 ft. elevation. It seems possible that subsequent investigation may prove this to be simply a well-marked variety of Lycaste brevispatba. The flowering season is in May and June. 60-A. KOELLENSTEINIA Rchb. f. KoELLENSTEINIA Rchb. f. in Bonplandia 2:17. 1854; Walp. Ann. 6:551. 1863; Schltr. in Orchis 12:24. 1918. Terrestrial or epiphytic herbs, with or without minute subterete or 4-angulate pseudobulbs. Pseudobulbs when present completely enveloped by the imbricating bases of the foliaceous bracts, the apex with 1 or 2 long-petiolate, slender, plicate (415) [Vor. 36 80 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN leaves; in plants without pseudobulbs, the several slender, plicate leaves resembling those of a Bletia, more or less contracted at the base into a short, sheathing petiole, the basal portions of which are enveloped in several imbricating bracts. In- florescences lateral, erect, slender racemes, usually more or less equaling the leaves, produced from the base of the current season's growth. Flowers few to many, of moderate size, on short pedicels. Sepals subequal, free, more or less spreading, the laterals usually somewhat broader. Petals subequal to the dorsal sepal or a little less. Lip 3-lobed, contracted at the base into a short claw which is adnate to, or apparently articulated with, the foot of the column, the lateral lobes erect or some- what spreading, the mid-lobe spreading, rounded or subquadrate, or subreniform, entire, or broadly 2-lobed; disk between the lateral lobes with a thickened callus. olumn short, semi-terete, somewhat arcuate, the apical margins projecting and forming a short hood over the clinandrium, the base produced into a short, broad foot. Anther terminal, operculate, incumbent, 1-celled; pollinia 2, waxy. A South American and West Indian genus previously unrecorded from Panama. 1. KoELLENsTEINIA KELLNERIANA Rchb. f. in Bonplandia 2:17. 1854. Warrea graveolens Hort. ex Rchb. f. Xenia Orch. 1:65. 1854. Erect terrestrial herbs, with small, subterete or tetragonous, tapering pseudo- bulbs 2—3 cm. long and 4—6 mm. wide. Leaves 1 or 2, linear-lanceolate, plicate, acute or acuminate, the bases contracted into a long petiole, 45—70 cm. long and 10-15 mm. wide. Inflorescences erect, 40—60 cm. tall, produced from the base of the current flush of new growth, the peduncle provided with several distant, papery, tubular, acute or acuminate bracts. Flowers of moderate size, few to many, reminiscent of those of a small Cyrtopodium or Матеа. Sepals rather fleshy, free, subequal, green, the dorsal sepal incurved, rather concave, elliptic- lanceolate, acute, 10—12 mm. long and 4—5 mm. wide, laterals somewhat spreading, rather obliquely elliptic-lanceolate, acute, 10—12 mm. long and 5-6 mm. wide. Petals subequal to the dorsal sepal, submembranaceous, pale green, elliptic- lanceolate, acute, 9-10 mm. long and 3.5-4 mm. wide. Lip rather fleshy, 3-lobed, white transversely barred lavender or purple, 6—8 mm. long and 8—10 mm. wide when spread out, the base abruptly contracted into a short claw which apparently is articulated with the foot of the column, lateral lobes obliquely rhombic-tri- angular, obtuse, erect, the mid-lobe broadly spreading, somewhat concave, more or less transversely oblong-reniform, the apex sometimes shallowly emarginate; disk with a low, fleshy, tuberculate callus. Column short, stout, semi-terete, rather arcuate, 3-4 mm. long, the margins of the apex slightly projecting and forming a short hood over the clinandrium. Panama, Colombia, Venezuela, British Guiana, and Brazil. PANAMA: grassy ridges, vic. Cerro Jefe, hills east of Panama City, 2500 ft., Rolland . COCLÉ: Lo Jones s.n. (under Allen 3701) OCLE: ma del Tigre, hills north of El Valle de Anton, 3000 ft., Allen 3562. (416) 1949] FLORA OF PANAMA (Orchidaceae) 81 61. ZYGOPETALUM Hook. ZYGOPETALUM Hook. in Bot. Mag. /. 2748. 1827; Benth. & Hook. Gen. РІ. 3:542. 1883. Zygopetalon Rchb. Consp. 69. 182 i Galeottia A. Rich. in Ann. Sci. Nat. Bot. III, 3:25. 1845. Epiphytic or terrestrial herbs. Pseudobulbs ovoid or subcylindric, tapering, the bases enveloped in several short or elongate, non-foliaceous or foliaceous bracts, the apex with 1 or 2 lanceolate or elliptic-lanceolate, plicate leaves. Inflorescences short or elongate, erect, racemose, produced from the base of the current growth; scape bracts small or large. Flowers few to many, small, or large and conspicuous. Sepals subequal, spreading, free or slightly connate at the base, the laterals adnate to the foot of the column, forming a short, broad mentum. Petals subequal to the sepals. Lip conspicuously or obscurely 3-lobed, the base affixed to, or articulated with, the column foot, the lateral lobes spreading or erect, the mid-lobe broadly spreading, or the apex strongly recurved; the disk with a prominent, fleshy, often lunate, tuberculate or cristate callus. Column rather short, stout, semi-terete, somewhat arcuate, the apex with or without conspicuous lateral wings, the base produced into a short foot. Anther terminal, operculate, incumbent, 2-celled; pollinia 4, waxy. A polymorphic genus of American epiphytic and terrestrial herbs, ranging fror southern Mexico to Peru and Brazil. a. Plants қ алаты Flowers large and conspicuous; lip 3-lobed, the margins 1. Z. GRANDIFLORUM aa. Plants ers Flowers small; lip subpandurate, the margins entire... 2. Z. PARVIFLORUM — . ZYGOPETALUM GRANDIFLORUM (A. Rich.) Benth. 8: Hook. ех Hemsl. Biol. Centr.-Amer. Bot. 3:251. 1883. Galeottia grandiflora A. Rich. in Ann. Sci. Nat. Bot. НЕ $ 25. 1845. Batemannia grandiflora Rchb. f. in Bonplandia 4:323 Epiphytic herbs, with ovoid, somewhat furrowed Кы 3.5—6 ст. long and 2—3 cm. wide, enveloped at the base in 3—4 papery, imbricating bracts; the apex of the pseudobulbs with 2 elliptic-lanceolate, plicate, acute leaves 25—50 cm long and 4—6.5 cm. wide. Inflorescences 15—18 cm. tall, produced from the base of the current flush of new growth. Flowers 1 to about 5, large and conspicuous, the pedicels subtended by broad, spathaceous, papery bracts. Sepals subequal, spreading, the apices recurved, green striped reddish brown, dorsal sepal free, lanceolate-acuminate, 3.0-4.5 cm. long and 1.0—1.5 cm. wide, laterals connate at the base and adnate to the foot of the column, forming a gibbose mentum, rather obliquely lanceolate, acuminate, 3.0-4.5 cm. long and 1.2-1.8 cm. wide. Petals subequal to the sepals, obliquely falcate, acuminate, spreading, the broad bases adnate to the sides of the column foot, the apices recurved, green striped reddish- brown, 3.0-4.0 cm. long and 1.2-1.8 cm. wide. Lip conspicuously 3-lobed, ab- (417) [Vor. 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN 7” Ly, >, 7 = £ 7 D z Еј z 2 Е тт тт P Fig. 175. Zygopetalum grandiflorum (418) 1949] FLORA OF PANAMA (Orchidaceae) 83 ruptly contracted at the base, with a short claw which is articulated with the column foot, white with dull red or purple longitudinal markings, lateral lobes erect, rather obliquely acute, with fimbriate margins, mid-lobe obovate, acuminate, somewhat concave, the attenuate apex strongly recurved, the lateral margins fimbriate; disk between the lateral lobes with a broad, erect, lunate, prominently ridged and furrowed, multi-denticulate callus. Column broad, stout, arcuate, 1.5-2 cm. long, the apex with 2 short, denticulate processes on either side of which are 2 subfalcate, obliquely obtuse wings with ciliate apical margins; the base of the column produced into a short, broad foot. Mexico, Guatemala, Costa Rica, and Panama. CHIRIQUÍ: vic. Cerro Punta, headwaters of the Río Chiriqui Viejo, 2000 m., Allen 3604 2. ZYGOPETALUM PARVIFLORUM L. Wms. apud Woodson & Schery, in Ann. Mo. Bot. Gard. 28:424, 7. 25. 1941. Erect, robust, terrestrial herbs, with slender, subcylindric, tapering pseudo- bulbs 6—9 cm. tall and about 1.5 cm. wide; leaves and blades of the foliaceous bracts linear-lanceolate to elliptic-lanceolate, plicate, acute or acuminate, the bracts contracted at the base into a short sheathing petiole, the imbricating bases completely enveloping the small pseudobulb. Leaves 30-70 cm. long and 2.5-7 cm. wide. Inflorescences 60—80 cm. tall, produced from the basal portion of the current flush of new growth. Flowers several or many, among the smallest in the genus, described as "purple, the lip violet; column white." Sepals subequal, rather fleshy, spreading, oblong-ovate, acute, 7-10 mm. long and 3.5-5 mm. wide. Petals subquadrate, obtuse, 7-8 mm. long and 4—4.5 mm. wide. Lip obscurely 3-lobed, subquadrate, panduriform, 7-8 mm. long and 5-6 mm. wide, abruptly contracted at the base, with a short claw which is articulated with the column foot, lateral margins rounded and spreading, mid-lobe rectangular, abruptly truncate, spread- ing; the disk with a broad, transverse, subreniform, fleshy callus. Column short, semi-terete, somewhat dilated at the apex, 4-5 mm. long. Anther and pollinia typical of the genus. Panama cHIRIQUÍ: vic. Bajo Chorro, headwaters of the Río Caldera, 1900 m., Woodson t$ Schery 605; vic. Bajo Chorro, in rain forest, 6000 ft., Davids son 345. 62. CHONDRORHYNCHA Lindl. CHONDRORHYNCHA Lindl. in Orch. Linden. 12. 1846; Rchb. f. in Walp. Ann. 6:663. 1863; Benth. & Hook. Gen. Pl. 3:548. 1883. Kefersteinia Rchb. f. in Bot. Zeit. 10:633. 1852. 2 js i 58. Warszewiczella Benth. & Hook. Gen. E y е, 1883. (419) [Vor. 36 84 ANNALS OF THE MISSOURI BOTANICAL GARDEN Tufted epiphytic herbs without pseudobulbs. Leaves distichously arranged in the form of an open fan, erect or arching, plicate, lanceolate, acute or acuminate, contracted below into conduplicate petioles. Inflorescences slender, erect, arching or semi-pendulous, 1-flowered, scapose, less than half the length of the leaves, pro- duced from the axils of the lower leaves or bracts. Flowers small to large and conspicuous. Sepals subequal, free, spreading, membranaceous, the dorsal sepal erect, the laterals often retrorse, obliquely inserted on the short column foot. Petals usually spreading, subequal to the dorsal sepal or broader. Lip usually cucullate, obovate, suborbicular or subquadrate, sometimes obscurely or con- spicuously 3-lobed, the lateral lobes or margins erect, or the lip explanate and divided into a narrow basal and a broad apical part, contracted at the base and adnate to, or articulated with, the very short column foot, sometimes forming a very short mentum; disk with a broad or narrow, more or less fleshy callus, the apex free and usually denticulate, or rarely with the basal callus conspicuously pedicellate, the apex scutellate. Column semi-terete, slender or broadly clavate above, sometimes narrowly winged, the ventral surface with or without a keel, rarely with a broad, 2-alate plate below the stigma, the base of the column pro- duced into a very short foot. Anther terminal, operculate, incumbent, 1-celled or imperfectly 2-celled; pollinia 4, waxy. a. Lip UN subpandurate, never concave or cucullate, divided into narrower basal and broader apical p aa. Lip concave or 1. entire ог prp suborbicular or obovate when spread out. b. Lip entire when те out. эн also alternate ББ) 1. C. AROMATICA c. Flowers small; 1.2 cm. long or les . Basal callus 2. ы at the apex 2-scutellate...... 3. C. COSTARICENSIS dd. Basal callus sessile, the free apex bifid 5. C. LACTEA cc. Flowers relatively large and conspicuous; lip 2.5 cm. long or more. d. Lateral эи s of the basal callus erect, thickened, and con- fluent w 4, erect incurving lateral margins of the lip. Low- and species dd. Lateral margins of the basal callus not erect, not thickened, тегімен us the incurving lateral margins of the lip. Hi ў 6. С. МАКСІМАТА land s e. Paul “callus linear, the free apex bifid 2. C. CALOGLOSSA ee. et callus radiate, re pd bow projecting apex with elongate teeth of unequal 1 4. C. DISCOLOR bb. жа — ог ан 3- lobed 2. spread out, 2.5 cm. 6; Later мч of the basal callus erect, thickened, and confluent erect incurving lateral margins of the lip. Lowland species 6. C. MARGINATA cc. Lateral puc of и basal callus not erect, not thickened, nor onfluen ith the erect incurving lateral margins of the lip. Highland hn d. Basal callus linear, with a free bifid apex 2. С. CALOGLOSSA dd. Basal callus radiate, digitate, the free projecting apex with 5—7 elongate teeth of unequal lengt 4. С. DISCOLOR (420) 1949] FLORA OF PANAMA (Orchidaceae) 85 1. CHONDRORHYNCHA aromatica (Rchb. f.) Р. H. Allen, comb. nov. Zygopetalum aromaticum Rchb. f. in Bot. Zeit. 10:668. 1852. Warscewiczella aromatica Rchb. f. in Walp. Ann. 6:654. 1863. Zygopetalum Wendlandi Rchb. f. Beitr. Orch. Centr.-Amer. 74. 1866. Bollea Wendlandiana Hort. ex Gard. & For. 1:315. Warscewiczella Wendlandi (Rchb. f.) Schltr. in Beih. Bot. Centralbl. 362:494. 1918. Erect tufted epiphytic herbs without pseudobulbs. Leaves linear-ligular to elliptic-lanceolate, plicate, acute or shortly acuminate, 15—30 cm. long and 1.5-2.5 cm. wide. Inflorescences erect or arching 1-flowered scapes 7-9 cm. long, from the axils of the non-foliaceous basal bracts. Flowers relatively large and con- spicuous. Sepals subequal, free, spreading, pale green or yellowish green, the dorsal sepal elliptic-lanceolate, acuminate, 2.5—3 cm. long and .6—.8 cm. wide, the laterals lanceolate, acuminate, 2.8—3.2 cm. long and .9-1.0 ст. wide. Petals sub- equal to the dorsal sepal, spreading, pale or yellowish green, lanceolate, acuminate, 2.5-3 cm. long and .5-.6 cm. wide. Lip explanate, subpandurate, the base ob- scurely lobed, abruptly contracted into a short claw, lavender or violet, usually with white margins, 2.2-2.5 cm. long and 1.5-1.7 cm. wide, divided into a more or less rectangular basal half and a broader undulate and reflexed apex; disk with a lunate to radiate or rhombic, spreading, plurisulcate, violet-blue callus. Column short, erect, white, semi-terete below, the apex clavate and narrowly winged. Anther terminal, operculate, incumbent, 1-celled. Costa Rica and Panama. CHIRIQUÍ: without definite locality, 4000-5000 ft., Powell 248. The flowers of our specimens seem to be somewhat smaller than typical Costa Rican material, but otherwise are identical. 2. CHONDRORHYNCHA caloglossa (Schltr.) P. H. Allen, comb. nov. Warscewiczella caloglossa Schltr. іп Fedde Rep. Sp. Nov. 12:216. 1913. Chondrorhyncha estrellensis Ames, Sched. Orch. 4:54. 1923. Erect tufted epiphytic herbs without pseudobulbs, 23-28 cm. tall, the plants typical of the genus. Inflorescences erect or arching, 1-flowered scapes 10—15 cm. long, from the axils of the lower foliaceous bracts. Flowers relatively large and conspicuous. Sepals subequal, membranaceous, free, white or pale yellow, the dorsal sepal erect, lanceolate, acute or acuminate, 2.8—3.2 cm. long and .6—.7 cm. wide, laterals obliquely inserted, deflexed, concave, lanceolate, acute or acuminate, 3-3.5 cm. long and .6—.7 cm. wide. Petals spreading, white or pale yellow, elliptic-oblanceolate, acute, 3—3.5 cm. long and 1.2-1.4 cm. wide. Lip concave, entire or obscurely or conspicuously 3-lobed when spread out, white or pale yellow, reticulated purple, 2.8-3.5 cm. long and 2—2.5 ст. wide; disk smooth, with a linear-ligular yellow callus, the free projecting apex bifid. Column semi- terete, 1.2-1.4 cm. long. Anther and pollinia typical of the genus. Costa Rica and Panama. (421) [Vor. 36 86 ANNALS OF THE MISSOURI BOTANICAL GARDEN IRIQUÍ: humid forest around Los Siguas Camp, southern slope of Cerro Horqueta, 1700 : m., Pittier 3176; Cuesta de las Palmas, Cerro Horqueta, 1700-2100 m., Pittier 3214, Maxon 5510; vic. Bajo Chorro, rain forest, 6000 ft., Davidson 93; trail бекий Cerro Punta to the headwaters of the Río Caldera, 2250-2500 m., Allen 1462. 5-; А somewhat variable species, the flowers reminiscent of those of Chond- rorbyncba discolor, but readily separable by the reticulated color pattern and narrow, bifid callus of the lip. 3. CHONDRORHYNCHA costaricensis (Schltr.) P. H. Allen, comb nov. Kefersteinia costaricensis Schltr. in Beih. Bot. Centralbl. 367:413. 1918. Dwarf tufted epiphytic herbs without pseudobulbs, 12—18 cm. tall. Leaves plicate, lanceolate, acuminate, 10—18 cm. long and 1.5-2 cm. wide. Inflorescences of slender, arching or semi-pendulous scapes 2.5—4 cm. long, from the base of the leaves. Flowers small, solitary. Sepals subequal, spreading, white or cream, the dorsal sepal elliptic-lanceolate, acute or subacute, 8-10 mm. long and 4—5 mm. wide, lateral sepals rather obliquely elliptic-lanceolate, acute or apiculate, 10—12 mm. long and 5-6 mm. wide. Petals subequal to the dorsal sepal, white or cream, often spotted maroon, obliquely oblong, acute, 7-9 mm. long and 4—5 mm. wide. Lip entire, suborbicular, contracted at the base and adnate to the foot of the column, 8—10 mm. long and 7-8 mm. wide, white or cream, spotted maroon, the apex entire or with an obtuse apicule; disk with an erect pedicellate callus, the apex 2-scutellate. Column erect, 5-7 mm. long, the apex broadly clavate, the ventral surface with a longitudinal keel which is expanded into a broad biauricu- late, abruptly apiculate plate below the narrow transverse stigma. Anther terminal, operculate, incumbent, 1-celled. Costa Rica and Panama. OLÓN: Río Llano Sucio, vic. Puerto Pilón, about 65 m., H. P. Butcher s. n. (under Allen. 2457). Known in Panama from a single fragmentary specimen preserved in liquid. Our material shows some slight differences from the Costa Rican type, notably in the subrhombic rather than scutellate divisions of the apex of the basal callus, and the apparently fleshier winged plate below the stigma, which has a more prominent central spur or apicule. Although it has been decided to consider our specimen as representing Chondrorhyncha costaricensis, it would seem well to note that this, and apparently one or two other species from outside our range, exhibit a radical departure from the typical concept of either Chondrorhyncha or Kefersteinia iu the conspicuously pedicellate basal callus of the lip and the broad plate on the under-side of the column below the stigma. It seems possible that these may actually represent an unrecognized entity of generic rank. Unfortunately, these characters are not always easily seen in dried specimens. However, the necessity for adequate material to show points of difference in no way alters the basic fact that they may exist. (422) 1949] FLORA OF PANAMA (Orchidaceae) 87 Due to our present very limited and somewhat fragmentary material, it is thought best to follow established usage for the purposes of this treatment. 4. CHOoNDRORHYNCHA discolor (Lindl.) P. H. Allen, comb. nov. Warrea discolor Lindl. in Jour. Hort. Soc. Lond. 4:265. Warscewiczella discolor (Lindl.) Rchb. f. in Bot. Zeit. 10: 636. 1852. Zygopetalum discolor (Lindl.) Rchb. f. in Walp. Ann. 6:655. 1861. Erect, tufted, epiphytic or pseudo-terrestrial herbs without pseudobulbs, 20—35 cm. tall, the plants typical of the genus. Inflorescences of slender, erect or arching, 1-flowered scapes 7-15 cm. long, produced from the axils of the lower non- foliaceous bracts. Flowers relatively large and conspicuous, sometimes nodding. Sepals membranaceous, subequal, free, spreading, white, often with yellowish apices, dorsal sepal erect, elliptic-lanceolate, acute, the apex recurved, 2.8-3 cm. long and 1.0—1.5 cm. wide, laterals deflexed, concave, lanceolate, acute, 3—3.2 cm. long and .8—1.0 cm. wide. Petals spreading, elliptic, obtuse to subacute, white, often suffused with purple toward the apex, 2.5-3 cm. long and 1.2-1.5 cm. wide. Lip cucullate, obscurely to conspicuously 3-lobed when spread out, 2.5—3.2 cm. long and 2.2-3 cm. wide, deep violet-purple, sometimes with narrow whitish mar- gins, lateral lobes or margins erect and incurving over the column, the apex sub- orbicular and slightly retuse; disk with a radiate, digitate, plurisulcate callus, the free projecting apex with elongate teeth of unequal length. Column short, semi- terete, narrowly clavate, 10—12 mm. long, white, sometimes blotched with purple. Anther and pollinia typical of the genus. Cuba, Costa Rica, and Panama. cHIRIQUÍ: without definite locality, 4500-5500 ft., Powell 155; upper Río Chiriqui Viejo, vic. Monte Lirio, С. White 42; llanos and slopes of Chiriqui Volcano and along the Río Chiriqui Viejo, 1200 m., Allen 014. A very frequent species of the Chiriquí highlands, where the plants are often found in association with those of Odontoglossum Schlieperianum, as epiphytes on low trees or sometimes as pseudo-terrestrials on boulders and steep mossy banks. 5. CHONDRORHYNCHA LACTEA (Rchb. f.) L. Wms. in Caldasia 5:16. 1942. Zygopetalum lacteum Rchb. f. in Gard. Chron. 1290. 1872 Kefersteinia lactea Rchb. f. apud B. D. Jackson in Index Kew. 2:4. 1895. Epiphytic herbs without pseudobulbs, the plants usually consisting of tufted clusters of several distichous crowns of relatively broad plicate leaves, which are oblong to elliptic-lanceolate, acute, 6-12 cm. long and 1.2—2 cm. wide, the basal portions somewhat contracted into a stout petiole. Inflorescences very short, filiform, more or less arching, or erect scapes about 1.5 cm. long, produced from the lower leaf axils. Flowers small, solitary. Sepals subequal, free, spreading, elliptic-lanceolate, obtuse to subacute, 8-10 mm. long and 3—4 mm. wide. Petais subequal to the sepals, elliptic-lanceolate, acute, 7—9 mm. long and 3—4 mm. wide. Lip undivided, suborbicular, 6-10 mm. long and 6—10 mm. wide when spread out, (423) [Vor. 36 88 ANNALS OF THE MISSOURI BOTANICAL GARDEN abruptly contracted at the base into a short claw which is apparently articulated with the base of the column, the apex entire, shallowly emarginate or with an obtuse apicule; disk with a low, fleshy, sessile, bifid callus. Column semi-terete, 5-6 mm. long, the ventral surface below the stigma with a longitudinal keel. Anther and pollinia typical of the genus. Costa Rica and Panama. CHIRIQUÍ: without definite locality, Wallis s. n. The species was described by Reichenbach from plants presumably collected in Chiriquí Province, the actual locality being unknown. Since there have been several subsequent collections from the Pacific slope in adjacent Costa Rica, there is every reason to believe that this rather obscure record may be good. 6. CHONDRORHYNCHA marginata (Rchb. f.) P. H. Allen, comb. nov. Warczewiczella marginata Rchb. f. in Bot. Zeit. 10:636. 1852. 1 52. Huntleya marginata Hort. ex Rchb. f. loc. cit. 18 Tufted epiphytic herbs without pseudobulbs. Leaves 5-7, linear, elliptic- oblanceolate, acute, 12—30 cm. long and 2-4.5 cm. wide, firm, arching, contracted below into conduplicate petioles the lowest pair being little more than foliaceous bracts. Inflorescences of slender, arching or semi-pendulous, 1-flowered scapes, 5.5-10 cm. long, from the axils of the lowest non-foliaceous bracts. Flowers the largest of the genus in Panama, with a very agreeable and characteristic fragrance during the morning hours, reminiscent of cinnamon or cloves. Sepals subequal, free, membranaceous, elliptic-lanceolate, acute, white, the dorsal sepal erect, some- what concave, the apex recurved, 2,2-3 cm. long and .9-1.2 cm. wide, the laterals usually more or less conduplicate, obliquely reflexed, 2.8—3.2 cm. long and .8-1.0 cm. wide. Petals membranaceous, spreading, white, usually with a lavender median line, elliptic-lanceolate, acute, 2.5—3 cm. long and 1.0-1.2 cm. wide, the apices recurved. Lip white, with a conspicuous pattern of radiating, often divari- cate, lavender or pinkish lavender lines, cucullate, subquadrate to obscurely 3- lobed when spread out, 3.5—4.5 cm. long and 3—3.5 cm. wide, contracted at the base and adnate to the short column foot, the lateral lobes or margins erect and incurved over the column, the frontal margin broadly spreading, often suffused with lavender or pinkish lavender, entire, or emarginate at the apex; the disk between the erect lateral lobes or margins with a broadly spreading, fleshy, pluri- sulcate callus, white striped lavender, the lateral margins of which are thickened and confluent with the erect lateral lobes of the lip and the apex more or less broadly truncate, free, the many short projecting teeth all of about equal length. Column semi-terete below, dilated and broadly clavate above, pure white, 12—18 mm. long. Anther terminal, operculate, incumbent, 1-celled. (424) 1949] FLORA OF PANAMA (Orchidaceae) Fig. 176. Chondrorhyncha marginata (425) [Vor. 36 90 ANNALS OF THE MISSOURI BOTANICAL GARDEN Panama and Colombia. ZONE: Cacao sg near Summit, Lipscomb s. п.; Pieces Powell 346 3: йе ‘Colorado Island, J. Zetek s.n.; vic. Frijoles, Powell 3460; Gatún Lake и, Powell II, 47; Cacao | i vic. Summit, Floren idi 9 James Barrett s western arm of the Quebrada Salamanca, 70 m., Dodge, Steyermark 8 Allen 16080. cocLÉ: damp shaded valleys south of El Valle de Antón, 600 m., Allen 2 2643. A fairly frequent and attractive species, found in damp shaded locations usually at low elevations. 63. PESCATOREA Rchb. f. РЕЗСАТОВЕА Rchb. f. in Bot. Zeit. 10:667. 1852; Benth. & Hook. Gen. Pl. 3:543. Erect, tufted, epiphytic herbs without pseudobulbs. Leaves plicate, contracted below into conduplicate imbricating petioles, which are distichously arranged in the form of an open fan. Inflorescences of short, slender, arching, 1-flowered scapes produced from the axils of the non-foliaceous basal bracts. Flowers relatively large and conspicuous. Sepals more or less fleshy, subequal, concave, the dorsal sepal erect, free, the laterals connate at the base and obliquely inserted on the column foot. Petals subequal to the sepals. Lip rather fleshy, 3-lobed, abruptly contracted at the base into a conspicuous ligular claw which is continuous with the foot of the column, the limb forming an obtuse angle with the column foot, the base apparently with a deep concavity below the column, surrounded by an erect semicircular plurisulcate callus; apical lobe of the lip more or less convex or ventricose, the lateral margins ‘often recurved. Column stout, semi-terete, pro- duced at the base into a short foot. Anther terminal, operculate, incumbent, 1- celled; pollinia 4, waxy. A small genus of tufted pseudobulbless epiphytes, ranging from Costa Rica to Colombia. The genus is sometimes considered as being referable to Warscewiczella, and has been so treated in the main generic key. In the exceedingly perplexing association of closely allied generic concepts embracing Bollea, Chondrorhyncha, Kefersteinia, Huntleya, Pescatorea, Warscze- wiczella, and Zygopetalum, it becomes largely a matter of individual opinion which are to be rejected and which retained. Probably something very nearly approach- ing a logical treatment of the situation was proposed by Reichenbach filius (in Walp. Ann. 6:650-662. 1863) in his reduction of nearly all of these to sections of Zygopetalum. This solution has never been universally accepted, however, the more recent tendency being to recognize some as valid genera and to reduce others to synonymy. Неге, as in most other cases of a similar nature, по hard-and-fast rule can be applied. Most genera, when examined critically, are found to be more or less arbitrary segregations, and should be recognized as being conveniences and а means to an end. Dr. Louis O. Williams has suggested that Kefersteinia is referable to Chond- rorbyncba, a view in which I, for the most part, concur since the generic type, (426) 1949] FLORA OF PANAMA (Orchidaceae) 91 K. graminea (Lindl) Rchb. f., and most of the subsequently described species are structurally nearly identical with Chondrorhyncha; yet there аге other Kefer- steinias, notably K. costaricensis Schltr., which differ radically from this generic concept. The type species of Warszewiczella, W. discolor (Lindl.) Rchb. f., also is almost identical with the earlier Chondrorhyncha, and apparently should be re- duced; but not including Pescatorea, which seems to have ample characters upon which to base segregation. For the purposes of this treatment, the following alternate key is proposed for the separation of the genera in our area: KEY TO THE GENERA a. Plants with pseudobulbs. Inflorescences racemose, several- to many- flowered 1. ZYGOPETALUM aa. Plants without pseudobulbs. Inflorescences of short 1-flowered scapes. b. Lip truly clawed at the base, explanate or convex, never cucullate. c. Apex of the column semiterete, never with broad projecting lat- eral wings. Basal callus plurisulcate, never fimbriate 3, PESCATOREA cc. Apex of the column with broad projecting lateral wings, forming a conspicuous hood over the clinandrium. Basal callus fimbriate.... 64. HUNTLEYA bb. Lip obscurely clawed at the base, usually convex or cucullate, rarely te 6 explana 2. CHONDRORHYNCHA 1. РЕЗСАТОВЕА CERINA (Lindl.) Rchb. f. in Bot. Zeit. 10:667. 1852. Huntleya cerina Lindl. & Paxt. in Paxton's Flow. Gard. 3:62. 1852—53. Zygopetalum cerinum (Lindl.) Rchb. f. in Walp. Ann. 6:651. 1863. Tufted epiphytic herbs without pseudobulbs. Leaves erect or arching, plicate, subcoriaceous, linear-lanceolate to elliptic-lanceolate, acute or acuminate, 15—60 cm. long and 3—5 cm. wide. Inflorescences of short, 1-flowered, erect or arching scapes 3.5-10 cm. long, from the axils of the basal non-foliaceous bracts. Flowers large and conspicuous. Sepals subequal, rather fleshy, concave, spreading, the dorsal sepal free, white, linear-elliptic to obovate, obtuse, 2.5-3.2 cm. long and 1.6-1.8 cm. wide, laterals somewhat connate at the base, obliquely inserted on the column foot, white with a long greenish-yellow blotch near the base, elliptic- lanceolate to oblanceolate, obtuse, 2.5—3.5 cm. long and 1.8-2 cm. wide. Petals spreading, subequal to the dorsal sepal, white, inserted on the base of the column, spatulate to oblanceolate, obtuse, 2.5-3 cm. long and 1.5-1.8 cm. wide. Lip rather fleshy, 3-lobed, 2—3 cm. long and about 2.5 cm. wide, rich yellow, the basal callus marked reddish brown, abruptly contracted at the base into a conspicuous ligular claw which is continuous with the column foot, the limb forming an ob- tuse geniculate angle with the claw, the subfalcate lateral lobes resting against the base of the column, forming a deep concavity which is surrounded by a con- spicuous erect, semicircular, plurisulcate callus, the frontal lobe convex to slightly ventricose, the apical margins often reflexed. Column short, stout, white, semi- terete, 1.3—1.5 cm. long, produced at the base into a short foot. Anther terminal, lavender. Costa Rica and Panama. PANAMA: Cerro Campana, in cloud forest near summit, 2500-3000 ft., Allen 4447, (427) [Vor. 36 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN A — ШІК Š AG Ў Vi Ш ІШ “ӨЛШ. MIT = ІҢ; ‘ i | ВН ЖЗ” , M, i ж” Fig. 177. Реѕсаіотеа сетіпа 5160. cocLÉ: vic. La Mesa, region north of El Valle de Antón, 1000 m., Allen 2362. VERAGUAS: forested slopes of Cerro Tuté, region west of Santa Fé, 2500—3000 ft., Allen 4564, 5171. CHIRIQUÍ: without definite locality, 4500 ft., Purdom s. n. An attractive species of the wet highland forests of the Pacific slope, usually being found in shaded situations at about 3000 ft. elevation. (428) 1949] FLORA OF PANAMA (Orchidaceae) 95 64. HUNTLEYA Batem. ex Lindl. HuNTLEYA Batem. ex Lindl. in Bot. Reg. 23: post 7. 1001. 1837; Benth. & Hook. Gen. Pl. 3:543. 1883. Erect, tufted, epiphytic herbs without pseudobulbs. Leaves plicate, subcoria- ceous, linear-lanceolate, acute, the bases somewhat conduplicate and imbricating, distichously arranged in the form of a fan. Inflorescences of slender, elongate, erect, 1-flowered scapes produced from the axils of the central leaves. Flowers large and conspicuous. Sepals rather fleshy, subequal, spreading, the dorsal sepal free, the laterals connate at the base and obliquely inserted on the column foot. Petals rather fleshy, spreading, subequal to the sepals, inserted on the foot of the column. Lip fleshy, abruptly contracted at the base, with a conspicuous claw which forms a geniculate angle with the column foot; the basal callus with an erect, semicircular, fimbriate margin, the apical lobe articulated with the base of the callus plate. Column short, somewhat arcuate, the apex dilated, with a dorsal keel, the lateral margins broadly winged and confluent at the apex, forming a con- spicuous hood over the clinandrium. Anther terminal, operculate, incumbent, 1- celled or imperfectly 2-celled; pollinia 4, waxy. About three or four species of American epiphytes, ranging from Costa Rica to Brazil. They often are listed as Zygopetalums. One species is known from Panama. 1. HuNTLEYA MELEAGRIS Lindl. in Bot. Reg. 23: post Ё. 1001. 1837. Batemannia Burtii Endres & Rchb. f. in Gard. Chron. 1099. 1872. Zygopetalum meleagris Benth. in Jour. Linn. Soc. 18:321. 1880. Zygopetalum Burtii Benth. & Hook. ex Hemsl. Biol. Centr.-Amer. Bot. 3:251. 1883. Huntleya Burtii (Endres & Rchb. f.) Rolfe, in Orch. Rev. 24:236. Erect, tufted, epiphytic herbs without pseudobulbs. Leaves Eo subcoria- ceous, linear-lanceolate to elliptic-lanceolate, acute, 15—30 cm. long and 3—4.5 cm. wide. Inflorescences of erect 1-flowered scapes 10—15 cm. tall produced from the axils of the centralleaves. Flowers large and conspicuous. Sepals subequal, fleshy, spreading, elliptic-lanceolate, acute to acuminate, the margins undulate, the apical 25 a waxy reddish brown, usually with some yellow spots, the basal !4 white or pale yellow, the dorsal sepal free, 4-6 cm. long and 2-2.5 cm. wide, the laterals connate at the base and obliquely inserted on the column foot, 4.5—6 cm. long an 1.5-2.5 cm. wide. Petals fleshy, spreading, subequal and colored similarly to the sepals, with purple blotches or streaks at the base near the insertion on the column foot, broadly elliptic-lanceolate, acuminate, with undulate margins, 3.5—5.6 cm. long and 2—3 cm. wide. Lip fleshy, the base nearly white or yellowish, the frontal half of the apical lobe usually а rich waxy reddish brown or brownish purple, 2.5-3.5 cm. long and 2-3 cm. wide, the base of the lip abruptly contracted, with a conspicuous claw which forms a geniculate angle with the foot of the column, the basal callus with an erect semicircular fimbriate crest, the posterior margins of which are incumbent on the foot of the column, the apical lobe obscurely 3- (429) ња “.. Жж tf Fig. 178. Huntleya meleagris (430) [ Vor. 36 1949] FLORA OF PANAMA (Orchidaceae) 95 lobulate, obovate, acute to acuminate, contracted at the base and articulated with the apex of the callus plate, the lateral lobules rounded, spreading, the apical lobule broadly triangular, the acuminate apex recurved. Column stout, erect, 1.5-2 cm. long, semiterete below, dilated above, with a conspicuous dorsal keel and broad lateral wings which are confluent at the apex, forming a hood over the clinandrium. Anther terminal, operculate, incumbent, imperfectly 2-celled. Costa Rica, Panama, Colombia, and Brazil. OCLÉ: western slope and summit of Cerro Valle Chiquito, 700-800 m., Seibert 643; trail to Las Minas, region north of El Valle de Antón, 1000 m., Allen 2804. A rare species of the wet highland forests, growing in deep shade at elevations above 2500 ft. The plants are often associated with a Brassia of nearly identical vegetative appearance, and are exceedingly difficult to distinguish when not in flower. The conduplicate bases of the leaves of the Brassia are often slightly broad- er below the suture line, while those of the Huntleya are of uniform width. The Central American plants have hitherto been called Huntleya Burtii, but other than some slight difference in size and color our material seems identical with specimens collected in Brazil. 65. MAXILLARIA Ruiz & Pavon MAXILLARIA Ruiz & Pavon, Fl. Peruv. & Chil. Prodr. 116, 7. 25. 1794; Lindl. in Bot. Reg. n.s. 6: Misc. 10. 1843; Benth. & Hook. Gen. Pl. 3:555. 1883. Ornithidium Salisb. in Trans. Hort. Soc. ук M 293. 1812. 44. Psittacoglossum La Llave & Lex. Nov. Veg. T 2:29. 1825. Pentulops Raf. loc. cit. 4:42. 1836. Epiphytic herbs with very short to elongate, often branching, erect, arching or pendulous stems, with or without clustered or distant, 1- to 3-leaved pseudobulbs. Leaves persistent, conduplicate in vernation, usually coriaceous or fleshy, rarely thin, usually strap-shaped, without prominent veins. Inflorescences of 1 to many, reduced to elongate, 1-flowered scapes from the bases of the pseudobulbs, from the axils of the leaves, or the flush of new growth. Flowers small to large and con- spicuous. Sepals subequal, free, or the laterals somewhat connate at the base, ad- nate to the foot or base of the column, often forming a short mentum. Petals subequal to the sepals or somewhat smaller. Lip concave, 3-lobed or entire, sessile or contracted at the base into a short claw, articulated with or adnate to the foot or base of the column, lateral lobes or margins erect, the mid-lobe membranaceous or thickened, spreading or reflexed; disk rarely without a fleshy callus. Column erect, semi-terete, somewhat arcuate, not winged, the base with a short foot, or footless. Anther terminal, operculate, incumbent, 1-сеПед or imperfectly 2-celled; pollinia 4, waxy. (431) [Vor. 36 96 ANNALS OF THE MISSOURI BOTANICAL GARDEN About 250 species of tropical American epiphytes, ranging from Mexico to Peru, Brazil, and the West Indies. As would be expected from a large group of plants having a great geographic range, they vary considerably in size and vege- tative habit. Тһе species can roughly be separated into two main divisions. In the first division the pseudobulbs are conspicuously present, either as sessile clusters or distributed along the rhizome; while in the second division the pseudobulbs are inconspicuous or entirely absent, the plants usually either erect canes with 2-ranked foliage, or with sessile clusters of leaves in the form of a fan. However, there are many species in which conspicuous pseudobulbs are at first produced at the base, the subsequent growth becoming elongate, often branching or scandent, lacking pseudobulbs, or with pseudobulbs small and hidden by the imbricating leaf bases. Plants of some of these modified types are often indistinguishable from those of Camaridium and Ornitbidium, the first of which in particular is a highly technical and arbitrary generic concept. In accordance with recently accepted usage, both of these are here considered to be Maxillarias. Ас the present time, some 41 species are known from Panama. a. spen stems usually conspicuously thickened into pseudobulbs. (See also sectio: b. Plants cae T Rhizome very short; 5” егесі, usually ы (rarely ar. apex mono phy llou . Bra enveloping the base of a Е conspicuously foliaceo d. Bases ncs e foliaceous bracts completely covering the poorly dev е. 4. ilb dd. Bases of the foliaceous bracts covering only the basal half of the well- décide: pseudobulb. e. Plants sma ee bracts 1-2. Sepals 5 mm. wide or M ate ww 14. M. CRASSIFOLIA less. Mentu 13. M. CONFUSA ee. Plants large. асс esti 3 or more. Sepals 8 mm. wide or mo Mentum short M. MALEOLENS cc. көм enveloping. the bases of s pseudobulbs not or apparently t folia d. E е p he lip № or more of the total length of the lip. На the lateral lobes of the Пр acute, fiie аа) M. RUFESCENS ee, Arce E the lateral lobes of the lip not acute, not con- usly pro jec cting dd. Mid. rios of the lip less than / the total length of the lip. e. 2 и меди cylindric, apparently continuous with the leaf petiole 16. M. CUCULLATA 5. M. ARACHNITIFLORA ee. nd ovoid to elliptic- ovoid, кане, but never pied broader than -— leaf petioles f. sey бар up to 25 ст. ta Mid. about ie de total. length of the lip. Apex of je» hos MV te 23 gg. Mid-lobe about i i» otal а of the lip. Apex of the речни broadly pene h. short, acute. Sepals acute, 15 mm. or less M. LONGIPETIOLATA ng 9. M. BREVIPES hh. Mentum жн. acuminate. Sepals acuminate, 17 13. M. CONFUSA mo н. Tene large, more doe 25 >= tall unies di өөө mw Flowers relatively large. Sepals 4 cm. 3 TE Sepa 15 маа use to suba с about 1 101 mm. пра на ЕРИДА 24. M. LUTEO h. minate, i | -АТВА ера!5 аБ mm. wide . M. ANGUSTISEGMENTA gg. Flowers intr d 20 3.5 cm. or less long. (432) 1949] FLORA OF PANAMA (Orchidaceae) h. Mid-lobe about № d i length of the lip. Apex of the pseudobulb s 23 hh. Mid-lobe about 14 m А ажа of the lip. Apex of the pse кедді; broadly trun i Sepals 2.5 cm. long or more 34. Sepals 2 cm. long or less 32. bb. Plants зара а Rhizomes or canes elongate, sometimes branch- ing; pseudobulbs approximate or distant, usually inserted on the rhizome at an oblique angle, sometimes confined to the base of the plant, erect, solitary or pn ere c. Pseudobulbs 22 Mods the rhizome, approximate, the ternodes not equaling the me of the pseudobulbs, often more or less prin ing ч ji ^ with the base of each pseudobulb covered by the apical portion CU preceeding it d. Flowers produced ien the base of the current mature pseudo- bulb. e. Bracts enveloping the bases of the pseudobulbs foliaceous. f. Apical lobe of the lip about 74 of the total length.............. 38 ff. Apical lobe of the Пр about 1 of the total length.............. 28 ee. pin enveloping a bases of the pseudobulbs not or ap- ently not foliace P білі» d usually completely hidden by the papery imbricating bracts. Leaves very fleshy, ы conduplicate, rarely lanceolate-elliptic. Lip concave, entire, ; 1 linear-spatulat 3 ff. Pseudobulbs нари әде not hidden by the bases of the imbricating brac Leaves coriaceous, ligular, or subulate- conduplicate. Lip. entire or 3-lobed. g. Scapes Г ru in dense subsessile fascicles. Flowers enclosed 2 conspicuous glumaceous bracts. скат broadly ovate, about 6 mm. long. Lip 3-lobed. Colum footless к” gg. Scapes conspicuously pedicellate, solitary or in loose fas- cicles. Sepals acute or obtuse. Lip entire, or obscurely 3-lobed. Column produced into a рање foot. h. Apex of the pseudobulbs 2- to 3-leaved. Scapes к, loosely fasciculate. Sepals acuminate hh. e = the pseudobulbs monophyllous. Scapes _ E solita i v or more long. Pseudobulbs strongly i 28 > rt 1j nii es 25 о ancipitous. "Sepals acute to acu ii. "niti 15 cm. long or less. Pseudobulbs puer not obtu ongly ancipitous. Sepals broadly obtuse................ 0. dd. iic Neues from the bract axils of the flush of new е. кыы 25 ст. ог more long. £. Bracts of the flush of new growth acuminate. Mid-lobe оѓ ned ff. Bracts of the flush of new growth broadly obtuse. Mid- 1 flush of ne lobe of the lip not conspicuously thickened 0. ee. Leaves 18 cm. long or less. f. Lip conspicuously 3-lobed, the mid-lobe more than !4 the total length of the lip ff. Lip €T M -lobed, the mid-lobe about 1% the total length o 40. cc. Pseudobulbs pe to | very distant on the rhizome, or confined the base of the plant. d. Pseudobulbs distant to very distant on the rhizom е. ңы rtm produced from the axils of A. current flush wth. et ° d Pseudobulbs diphyllous. (433) 8. 97 M. LONGIPETIOLATA M. RINGENS M. PowELLIr . M. VAGANS . M. OREOCHARIS 7. M. UNCATA . M. NEGLECTA . M. FRIEDRICHSTHALII M. OREOCHARIS M. VARIABILIS M. ALBA M. CAMARIDII M. VAGANS M. VARIABILIS [Vor. 36 98 ANNALS OF THE MISSOURI BOTANICAL GARDEN g. Leaf petioles very short, igi apices of the leaves obtuse. Bracts of the flush of new growth broadly obtuse............ 10. M. CAMARIDII gg. "d petioles elongate, the apices of the leaves acute. cts of the flush of new growth acuminate.................. 15. М. CTENOSTACHYA ff. Dei monophyllou owers minute, the ak less than 8 mm. long. ч Pseudobulbs suborbicular 26. M. MINOR hh. Pseudobulbs linear M. WERCKLEI gg. Flowers of moderate size, the sepals more than 8 mm. ong. h. Pseudobulbs linear. Leaves less than 3 cm. long.......... 41. M. WERCKLEI ud hh. —— = he ptic-ovate to elliptic-oblong. Leaves m. long. i. Lip соло 3-lobed when spread out. j. Mid-lobe а bout 1⁄4 the total length of the lip........ 1. M. ALBA 36. M. UMBRATILIS ounde acute, the mid-lobe obtuse or subacute. Sepals cu 38. M. VAGANS ii. че entire ог obscurely 3-lobed, subpandurate when ead out 8. M. DIUTURNA ее. = ng gene produced from the base of the current ature pseudobulb. * Flowers small, дне in dense subsessile fascicles. Col- umn without 27. M. NEGLECTA ff. Flowers of ae rate x usually 1—3, each flower solitary in the bract axil, never scii in dis subsessile fas- cicle. Column with a . foot. g. Leaves em е ней less than 1 ст. wide. Sepals about 28 gg. 2 ойу pns more than 3 cm. wide. Sepals ut long = OREOCHARIS 31. M. PLANICOLA dd. Билди ақы to the base of the plant . Flowers large and conspicuous. Sepals 3.5 cm. long or more. 1 ng ip obscurely 3-lobed, about 1.5 cm. long 22. M. INAUDITA ip conspicuously 3-lobed, about 1 cm. long 8. M. BRADEORUM ee. Flowers small. Sepals 1.5 cm. long or less. s NE ovate. Lip 4. sigmoid when seen 21. M. FULGENS £f. “N acute to о пос broadly ovate. Lip пос geniculate sigmoid i g. Lip ү эбе se ры. oni lobes auriculate, erect........ 6. M. BIOLLEYI 30. M. PITTIERI ња їр епиге аа. Secondary stems not, ог rarely, ог obscurely 7. into pseudo- bulbs, or the pseudobulbs confined to the base of the pla b. Plants c caespito с; amd c conduplicate, imbricating, distichously arranged in the for a broad d. pp fleshy. Flowers with short pedicels, subsessile in the дир он 2 axils. . Leaves many, equitant, gladiate, acuminate. Plants pendent... 39. M. VALENZUELANA ee. рдім few, ligular, acute ог obtuse, the conduplicate bases completely RE a small pseudobulb. Plants erect. f. Plants more than 15 cm. tall. Leaves more than 15 mm. wide 14. M. CRASSIFOLIA ff. Plants less гт 10 ст. tall. Leaves less than M. BRACHYBULBON dd. que subcoriaceous. Flowers with long pedicels, han т: f rum basal conduplicate bracts 11. M. CHARTACIFOLIA the axils o cc. Leaf dans т below into narrow sheathing petioles, ог ар- arently (434) 1949] FLORA OF PANAMA (Orchidaceae) d. Leaves fleshy, obtuse, with short, ру iet d imbricating petioles. 52277 giis in the leaf a dd. Leaves coria nate. ird usually long- pedicellate, Mir ме уш іп се Тр xils. e. Leaves broad, 2.5 cm. wide or more. Flowers large, the sepals 4 cm. long or more ee. din narrow, .8 cm. wide or less. Flowers small, the sepals m. long or les a ы ts dwarf, usually less than 6 cm. tall ff. Plants 52. ерің rd cm. or s long bb. Plants caulescent, the gate ca mes repent, erect or eae imd sometimes vun ibus s at e pe Leaves 2-ranked on the C Case or as undivided, or apparently 50. 4. ст Ын fri cm. м. ог diu ee. s eav rie or Le. apices retuse dd. 5 more than 25 cm. tall. e. Leaves D a distichous, fasciculate cluster at the ape of the the imbricating bases sometimes enveloping a small pse ER ‘bulb ee. "sedg Danny distributed along the upper stems, the bas t fasciculat £. срби ves 9 ст. long ог a the apex retuse. Flowers small, the sepals 1.5 Se hes (ba 5. g. Sepals about cm Lip conspicuously 3-lobed, the basal lobe awaq erect, the apical lobe large, 2. ob gg. Sepals а m. long. Lip linear, pice И rather е F е 4, е d Vp small; СОСЕ. ff. Leaves 15 cm. or e lon pex br ы, СХ со acuminate. оен: а Ге Зои the sepals long or more. g. Lip about 9 mm. long, conspicuously 3- ыны the lateral lobes acute, broadly triangular to subfalcate.................... gg. Lip На ut 15 mm. long, rather pee lob bed. h. acuminate, 1.5 г less wide, the bases Guide clasping the суши ‹ canes hh. к, broadly obtuse, 3.5 ст. ог more wide, the nt bases conspicuously flattene cc. Canes or е mes xp edm d. Rhizome repent, Е freely from the bases of нң отне 59 14. M. CRASSIFOLIA 5. ^ 1 M. ARACHNITIFLORA . M. BRASHYBULBON M. ANGUSTISSIMA M. UNCATA . M. WERCKLEI 21. M. FULGENS 6. M. BIOLLEYI . M. ALLENII 8. M. BRADEORUM 19. 22, M. EXALTATA M. INAUDITA leaf clusters, 2. аге өске: ог terminal оп the stems............ 33. M. REPENS dd. Canes erect or penden reely rooting from стерот leaf kic the p «уйнау distributed оп the Кс; s dwarf, 15 ст. ог t: Ves coriaceous, poo broad obtu 41. М. WERCKLEI ff. edis fleshy, бара late condor АНИ ог pem lanceolate, acute, never Eb btus 37. M. UNCATA ee. Plants 3 rm all. Canes ni на Leaves very short, 6 cm. or less long (usual- ly m ff. ru ашлау stout. Leaves 10 ст. lon Flow conspicuously мено на d. over 15 mm. tinct foo dig ong g olumn with a distinct foot... | gg. ры іп dens subsessile fiiit: sepals 10 mm. or m long. Colu нао а foot. Plants Bia with a ad uisu Мс; the base. ip gen бги. дь? іп profile. hh. РВ not “g culate, not sigmoid in profile. 1. Арех of ds lip 2-lobed 1. Apex of the lip subarite tiie. EEA 17. M. DENDROBIOIDES . M. PARVILABIA M. CONDUPLICATA . M. Prr [Vor. 36 100 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1. MaxiLLARIA ALBA (Hook.) Lindl. Gen. & Spec. Orch. Pl. 143. 1832. Dendrobium album Hook. Exot. Fl. 1. N^. ршина Brougbtonia alba Spreng. Syst. Veg. 735. 1826. Epiphytic herbs with elongate, often branching, more or less pendulous, rhizo- matous stems enveloped in closely imbricating, persistent, brown, papery bracts. Pseudobulbs approximate, inserted at an acute angle on the stems and more or less imbricating, linear-elliptic, strongly ancipitous, 4—5 cm. long and 2—2.5 cm. wide, the rather oblique truncate apex with a solitary leaf. Leaves ligular, coriaceous, persistent, acuminate, 25-40 cm. long and 1.5-2 cm. wide. Inflorescences short 1-flowered scapes 3-5 cm. long from the bract axils of the flush of new growth. Flowers few to about 8, relatively small, apparently produced more or less simul- taneously rather than in successive flowerings as in some of the allied species. Sepals subequal, free, spreading, creamy white, ligular, acuminate, 2-2.5 cm. long and .4—5 cm. wide, the laterals adnate to the column foot, forming a short sub- acute mentum. Petals subequal to the sepals, creamy white, lanceolate, acute to acuminate, 1.8—2 cm. long and .4—5 cm. wide. Lip concave, slightly arcuate, obscurely 3-lobed, yellow; more or less elliptic-oblanceolate, acute, when spread out, 12-15 mm. long and about 5 mm. wide, contracted at the base and articulated with the foot of the column, the lateral lobes erect, the mid-lobe about № the total length of the lip, ovate, acute, somewhat thickened; disk with an elongate, ligular, fleshy callus. Column semi-terete, lightly arcuate, 7-9 mm. long, somewhat dilated and obscurely auriculate at the apex, the base produced into a distinct foot. Anther terminal, operculate, incumbent, obscurely 2-celled. Guatemala, Honduras, Costa Rica, Panama, Cuba, Jamaica, Trinidad, British Guiana, Surinam, Brazil, and probably other adjacent territories. PANAMÁ: hills near Panama City, sea level, Powell 127; vic. Juan Diaz, sea level, Cope s. n. (under Allen 3822). снікюоі: vic. Boquete, 3800 ft., Davidson 826; in low woods along small streams on rocky plains я 5 miles south of Boquete, 3000 ft. ” Allen 4706 2. MAXILLARIA ALLENII 1. Wms. in Ann. Mo. Bot. Gard. 27:282 Ё. 35. 1940. Erect epiphytic herbs without pseudobulbs, the complanate-cylindric canes 25—40 cm. tall, the lower half to two-thirds leafless and enveloped in the coarse, closely imbricating, persistent leaf bases. Leaves 2-ranked, equidistantly dis- tributed along the upper half to third of the stem, coriaceous, ligular, emarginate, 3.5-6.5 cm. long and 1.0-2.0 ст. wide. Inflorescences slender 1-flowered scapes produced in loose fascicles from the leaf axils. Flowers small. Sepals subequal, free, not spreading, lanceolate, acute to acuminate, 7-10 mm. long and 2-2.5 mm. wide, the laterals adnate to the foot of the column, forming a very short subacute mentum. Petals subequal to the dorsal sepal, lanceolate to oblanceolate, acute, pale yellow, 6—8 mm. long and about 2.5 mm. wide. Lip slightly concave, reddish tan to reddish orange, obscurely 3-lobed, articulated at the base with the foot of the column, 6—7 mm. long and about 3 mm. wide, lateral lobes erect, the acute apices somewhat spreading, the mid-lobe about half the total length of the lip, (436) 1949] FLORA OF PANAMA (Orchidaceae) 101 the acute apex somewhat thickened and with erect margins; disk with а low, fleshy, linguiform callus. Column semi-terete, somewhat arcuate, about 4 mm. long, produced at the base into a foot. Anther terminal, operculate, incumbent, 1-celled. Panama. PANAMÁ: iig summit of ber Campana, 800-1000 m., Allen & Fairchild va COCLÉ: wet mossy forest on Pajita, hills north of El Valle de Antón, 1200 m Cer Allen & Fairchild. Sod; region PNA of El Valle, 1000 m., Allen 1650, 2002, 3021. 3. MAXILLARIA ANGUSTISEGMENTA Ames & Schweinf. іп Sched. Orch. 10:86. 1930. Erect epiphytic herbs with stout ovate to rectangular-elliptic, somewhat сот- pressed pseudobulbs 4—6 cm. tall and 1.5—3.5 cm. wide, the bases enveloped in several imbricating papery bracts which become fibrous with age, the truncate apex of the pseudobulb with a single leaf. Leaves coriaceous, persistent, the blades oblong, acute, contracted below into an elongate, conduplicate petiole, 16—50 cm. long and 2-5 cm. wide. Inflorescences usually several, erect, 1-flowered scapes 12-15 cm. tall, enveloped in several approximate, tubular, acuminate, papery bracts. Flowers large and conspicuous. Sepals subequal, free, widely spreading, yellow, linear-lanceolate, long-acuminate, 4.5—5.5 cm. long and .5-.6 cm. wide, the laterals adnate at the base to the foot of the column, forming a short mentum. Petals subequal to the sepals, white, linear-lanceolate, acuminate, the apices in- curving, 4—4.5 cm. long and .35—5 cm. wide. Lip 3-lobed, 12-15 mm. long, contracted at the base and articulated with the foot of the column, the lateral lobes elongate, erect, the anterior margins rounded, the mid-lobe about М the total length of the lip, suborbicular to subquadrate, obtuse; the disk with a fleshy ligular callus. Column semi-terete, somewhat arcuate, 8-10 mm. long, produced at the base into a foot. Costa Rica and Panama. cHIRIQUÍ: without definite locality, Terry 1610; Bajo Mono, mouth of the poni Chiquero and along the Rio Caldera, 1500-2000 m., Woodson, Allen & Seibert 1010 Our specimens vary among themselves, none agreeing entirely with the Costa Rican type, yet they seem to form an association more readily referable to that concept than to any other species, differing from Maxillaria ringens in the larger flowers and more robust habit, and from Maxillaria luteo-alba in the much more attenuate floral segments. It seems likely that this, and many others of this asso- ciation, will eventually be reduced to synonymy. 4. MAXILLARIA ANGUSTISSIMA Ames, Hub. & Schweinf. in Bot. Mus. Leafl. Harv. Univ. 3:41. 1934. Maxillaria acutifolia Schltr. in Fedde Rep. Sp. Nov. Beih. 19:229. 1923, non Lindl. Caespitose, usually pendulous, epiphytic herbs without pseudobulbs, the short rhizome with one to several, sometimes divaricate, clusters of long-attenuate (437) [Vor. 36 102 ANNALS OF THE MISSOURI BOTANICAL GARDEN foliage. Leaves coriaceous, very narrow (in our specimen), 45—75 cm. long and .2—.3 ст. wide, the bases enveloped in several papery, imbricating bracts, forming a short complanate petiole. Inflorescences apparently solitary, short, 1-flowered scapes from the lower leaf axils. Sepals subequal, free, apparently not spreading (in our specimen), lanceolate, acuminate, about 2.5 cm. long and 5-6 mm. wide, the very oblique bases of the laterals adnate to the long column foot, forming a conspicuous, elongate, acuminate mentum. Petals apparently subequal to the sepals. Lip obscurely 3-lobed, concave, the slender base articulated with the column foot, the lateral lobes erect, the apical lobe spreading, about !4 the total length of the lip; disk with a low, fleshy, linear callus about equaling the lateral lobes in length. Column short, semi-terete, somewhat arcuate, produced at the base into a very long narrow foot. Costa Rica and Panama. COLÓN: summit of Cerro Santa Rita, 1200 ft., Allen 9 Fairchild 5108. АП the plants seen of this curious species have had the leaves consistently longer and narrower than in the typical Costa Rican material. The single flower of our specimen is fragmentary, and hence cannot be determined with certainty, but the plants and flowers in general aspect apparently compare better with this than with any other known species. It may be that adequate collections will prove it to be distinct. 5. MAXILLARIA ARACHNITIFLORA Ames & Schweinf. in Sched. Orch. 10:87. 1930. Erect, caespitose, epiphytic herbs, up to about 38 cm. tall. Pseudobulbs com- planate-cylindric, monophyllous, 4—5.5 cm. long and .5-1.0 cm. wide, the bases enveloped in several imbricating, sometimes foliaceous bracts. Leaves coriaceous, persistent, lanceolate, acute, 15—35 cm. long and 2.5-4.5 cm. wide, contracted below into an elongate, conduplicate petiole which is apparently continuous with the apex of the narrow pseudobulb. Inflorescences few to many erect 1-flowered scapes, 7—12 cm. tall, produced from the bases of the pseudobulbs, enveloped throughout in tubular, acuminate, papery bracts. Flowers relatively large and conspicuous. Sepals free, subequal, spreading, greenish yellow, linear-lanceolate, attenuate, 5—7.5 cm. long and .5-.6 cm. wide, the oblique bases of the laterals adnate to the column foot, forming a conspicuous acute mentum. Petals subequal to the sepals, white, erect with incurving apices, linear-lanceolate, long-acuminate, 5—6 cm. long and .4-.5 ст. wide. Lip concave, slightly arcuate, nearly entire but obscurely 3-lobed near the apex, clear yellow, oblong-ovate when spread out, 1.5—2 cm. long and .9—1.1 cm. wide, the base articulated with the foot of the column, the lateral margins erect, the apices bluntly acute, the obtuse mid-lobe somewhat thickened, with a fleshy callus on the under-side of the apex; disk with a fairly prominent, fleshy, slightly concave, linguiform callus about half the length of the lateral margins. Column short, stout, semi-terete, 6—7 mm. long, produced at the base into a very conspicuous foot which is about twice the length of the upper column. (438) 1949] FLORA OF PANAMA (Orchidaceae) 103 Costa Rica and Panama. cocLÉ: vic. El Valle de Antón, 600-1000 m., Allen 1250; trail to Las Minas, region north of El Valle, 1000 m., Allen 2875. 6. MAXILLARIA BIOLLEYI (Schltr.) L. Wms. in Ann. Mo. Bot. Gard. 28:425. 1941. Ornitbidium Biolleyi Schltr. in Fedde Rep. Sp. Nov. 9:29. 1910. Camaridium Biolleyi Schltr. in Beih. Bot. Сеш ШЫ, 362:498. 1918. Егесе ерірһугіс herbs, 30-50 ст. tall, the stout complanate-cylindric сапев enveloped below іп the broad persistent leaf bases, the base of the cane often with one or more oblong-elliptic, compressed pseudobulbs, 2.5—3 cm. long and 1.5-2 cm. wide. Leaves coriaceous, spreading, 2-ranked and equidistantly distributed on the upper stem, the blades ligular, shallowly and unequally 2-lobed to acute, 7-27 cm. long and 1.5-3.5 cm. wide, the conduplicate bases closely imbricating. In- florescences slender 1-flowered scapes, 6—9 cm. long, produced in dense fascicles from the leaf axils. Flowers small. Sepals subequal, free, spreading, white, lanceolate, acuminate, 10—12 mm. long and 3.5-4 mm. wide, the laterals somewhat obliquely inserted. Petals subequal to the sepals, rather obliquely lanceolate, acuminate, white, 9-10 mm. long and 2—3 mm. wide. Lip 3-lobed, shortly clawed at the base, red, 5—6 mm. long, the lateral lobes auriculate and erect, the mid-lobe spreading, oblong-elliptic, obtuse, about 24 the length of the lip; disk between the lateral lobes with a broadly obtuse, Ды: 520 fleshy callus. Column short, stout, semi-terete, 2—2.5 mm. long, produced at the base into a very short foot. Costa Rica and Panama. CHIRIQUÍ: Bajo Chorro, in rain forest, 6000 ft., Davidson 125. 7. MAXILLARIA BRACHYBULBON Schltr. іп Fedde Rep. Sp. Nov. Beih. 19:55. 1923. Dwarf, caespitose, epiphytic herbs, 3.5—8 cm. tall, usually without pseudobulbs. Leaves coriaceous, ligular to oblanceolate, obtuse, 2-4.5 cm. long and .3—8 cm wide, contracted below into elongate, conduplicate petioles, the bases of which are imbricating and more or less 2-ranked. Inflorescences short, usually solitary, 1- flowered scapes produced from the base of the leaves. Flowers small, but rela- tively large in relation to the size of the plants. Sepals subequal, free, yellow, lanceolate, acute, 12—15 mm. long and 3—4 mm. wide, the laterals obliquely in- serted on the column foot, forming a short subacute mentum. Petals subequal to the dorsal sepal, lanceolate, acuminate, yellow, 10-12 mm. long and 2-2.5 mm. wide. Lip 3-lobed near the apex, red-brown, contracted below and articulated with the column foot, the lateral margins erect, the apices acute, mid-lobe fleshy, ligular, acute, minutely papillose, about 15 the total length of the lip. Column short, semi-terete, somewhat arcuate, about 5 mm. long, the base produced into a short foot Honduras, Costa Rica, and Panama. PANAMA: Cerro Campana, 1000 m., Allen 4026. (439) [Vor. 36 104 ANNALS OF THE MISSOURI BOTANICAL GARDEN 8. MaxILLARIA BRADEORUM (Schltr.) L. Wms. in Ann. Mo. Bot. Gard. 28:425. 1941. Camaridium grandiflorum Ames, i in Proc. Biol. Soc. Wash. 34:149. 1921. Camaridium Bradeorum Schltr. in Fedde Rep. Sp. Nov. Beih. 19:141. 1923. Maxillaria ampliflora C. Schweinf. in Bot. Mus. Leafl. Harv. Univ. 8:188. 1940. Erect or pendulous, epiphytic herbs, 45—75 cm. tall, with elongate, cylindric canes, the lower portion usually with inconspicuous, distant, complanate-elliptic pseudobulbs, the internodes naked or enveloped in the persistent, imbricating leaf bases, the upper stem without pseudobulbs and with 2-ranked foliage. Leaves coriaceous, ligular-lanceolate, acute, 12-27 cm. long and 2-3.5 cm. wide; the con- duplicate petioles imbricating and 2-ranked on the stem. Inflorescences erect 1- flowered scapes 6.5—8 cm. tall, enveloped in several tubular, papery, acuminate bracts. Flowers relatively large and conspicuous. Sepals free, spreading, subequal, the apices recurved, white with a central rose-red blotch, to wine-red with yellowish margins, lanceolate, acuminate to long-acuminate, 3-5 cm. long and 1.0-1.2 cm. wide, the dorsal sepal with a keel, the laterals inserted on the very short column foot barely forming an obscure mentum. Petals subequal to the sepals and similarly colored, lanceolate, acuminate to long-acuminate, 2.5—4 cm. long and 7-8 mm. wide. Lip very short, 3-lobed, rich orange, 8—10 mm. long and 12-14 mm. wide when spread out, abruptly contracted at the base into a short claw which is articulated with the foot of the column, the subfalcate, acute, lateral lobes erect, the obtuse, concave mid-lobe about 25 the total length of the lip, the lateral margins lightly incurving; the disk "id a truncate, sulcate or obscurely 3-lobed fleshy callus which about equals the lateral lobes in length. Column short, semi-terete, 6—8 mm. long, the base with a very short foot. Costa Rica and Panama. COCLÉ: summit of кер Pajita, hills north of El Valle de Antón, 1200 т., Allen Fairchild 3942. CHIRIQUÍ: humid forest of the cordillera east of the Rio Caldera, ei m., Killip 3565; vic. Bajo fidem 6000 ft., Davidson 118. The species is somewhat variable in regard to the presence or absence of pseudo- bulbs and in the size and color of the flowers. 9. MAXILLARIA BREVIPES Schltr. in Fedde Rep. Sp. Nov. Beih. 19:302. 1923. Dwarf, erect, epiphytic herbs up to about 8 cm. tall. Pseudobulbs clustered, ovoid-complanate, 8—10 mm. tall and 6—8 mm. wide, the bases enveloped in sev- eral imbricating papery bracts, the apices monophyllous. Leaves coriaceous, elliptic-lanceolate, acute, 4-6.5 cm. long and .8-1.2 cm. wide, contracted below into slender, elongate, conduplicate petioles. Inflorescences usually solitary 1- flowered scapes, 2—2.5 cm. tall, produced from the base of the pseudobulbs. Flowers relatively small. Sepals subequal, free, apparently not spreading, white, ovate-lanceolate, acute to subacute, 10-13 mm. long and 3—4 mm. wide, the ob- lique bases of the laterals adnate to the column foot, forming a short, acute (440) 1949] FLORA OF PANAMA (Orchidaceae) 105 Fig. 179. Maxillaria Camaridii (441) [Vor. 36 106 ANNALS OF THE MISSOURI BOTANICAL GARDEN mentum. Petals subequal to the dorsal вера], somewhat obliquely lanceolate, acute, white, 10-11 mm. long and 2.5-3.5 mm. wide. Lip concave, elliptic-obovate, subacute when spread out, reddish brown, obscurely 3-lobed near the apex, 8-10 mm. long and 4—5 mm. wide, the cuneate base articulated with the foot of the column, the lateral margins erect, the apices rounded, the mid-lobe creamy yellow, obtuse, about 14 the total length of the lip; disk with a linear, obtuse, yellow callus about equaling the lateral margins in length. Column short, stout, semi- terete, produced at the base into a long narrow foot. Costa Rica and Panama. COCLÉ: north rim of El Valle de Antón, near Cerro Turega, 650—700 m., Woodson 9 Schery 201a; mountains beyond La Pintada, 400—600 m., Hunter & Allen 550; mossy forest on crest of Cerro Pajita, hills north of El Valle de Antón, 1200 m., Allen 9 Allen 4172. Possibly a dwarf form of Maxillaria Reichenheimiana. 10. MaxiLLARIA Слмакшп Rchb. f. in Hamb. Gartenzeit. 19:547. 1863. Camaridium ochroleucum Lindl. in Bot. Reg. 10: t. 844. 1824, non Maxillaria ochroleuca Lodd. ex Lindl. Cymbidium ochroleucum Lindl., Сеп. & Spec. Orch. РІ. 168. . Ornithidium album Hook. in Bot. Mag. t. 3306. 1834, поп Maxillaria alba (Hook.) Lindl. Camaridium affine Schltr. in Fedde Rep. Sp. Nov. Beih. 17:72. 1922. Epiphytic herbs with elongate, complanate-cylindric, pendulous stems, the lower portions with more or less approximate, elliptic, ancipitous, diphyllous pseudobulbs 3-7 cm. long and 1.5-3 cm. wide inserted at an acute angle, those toward the apex produced at more distant intervals; the internodes and the bases of the pseudobulbs enveloped in the persistent, chartaceous, brown, imbricating bases of the obtuse, foliaceous bracts. Leaves subcoriaceous, linear-lanceolate, obtuse, retuse, or unequally 2-lobed at the apex, 15—30 cm. long and 12-18 mm. wide, the conduplicate bases contracted into very short petioles. Inflorescences short 1-flowered scapes 4—5 cm. long, usually produced in successive pairs from the bract axils of the flush of new growth, each plant thus flowering 3 or 4 times during a given season. Flowers very fragrant, relatively large and conspicuous. Sepals subequal, free, widely spreading, somewhat concave, pure white, elliptic- oblanceolate, acute, 2.5-3.5 cm. long and 1.0—1.4 cm. wide, the laterals adnate at the base to the short column foot, forming an inconspicuous rounded mentum. Petals subequal to the sepals, widely spreading, somewhat concave, pure white, elliptic-oblanceolate, acute, 2.2-3 cm. long and 9-10 mm. wide. Lip white on the outer surface, rich yellow within, with reddish brown or reddish purple transverse lines, conspicuously 3-lobed, 10-12 mm. long and 10-12 mm. wide when spread out, contracted at the base and articulated with the foot of the column, the lateral lobes erect, rounded, the anterior margins obtuse to acute, the mid-lobe acute to suborbicular, more or less canaliculate to spreading, 25 to V5 the total length of the lip; disk with a linear-lanceolate, concave, yellow callus, % to 34 the length of the lateral lobes, the obscurely tridenticulate apex of which is fleshy and glabrous, (442) 1949] FLORA ОЕ PANAMA (Orchidaceae) 107 the basal 34 densely and conspicuously рарШозе. Column semi-terete, somewhat arcuate, 6-8 mm. long, pure white, with a reddish brown or deep purple blotch at the base. Guatemala, Costa Rica, Panama, Trinidad, British Guiana, Surinam, and prob- ably other adjacent territory. CANAL ZONE: Río Pedro Miguel, near Paraiso, Standley 29988. PANAMA: hills east of рч City, sea level, Powell 7; Río Тесішеп, Standley 20413; Juan Diaz, sea level, s.".; Cerro Campana, vic. Campana, 2000 ft, D. Allen 5087; San José Island, Perlas Archipelago, Harlow 58, Johnston 206, 1403. cocLÉ: hills south of El Valle de Antón, 800 m., Allen 2667. A common and attractive species, widely distributed throughout the lowlands of our area. The flowers are produced usually in pairs in several successive flower- ings lasting but a single day in each instance. The fragrance is reminiscent of that of Narcissus. 11. MAXILLARIA CHARTACIFOLIA Ames & Schweinf. in Sched. Orch. 10:92. 1930. Erect, epiphytic herbs without pseudobulbs, the foliage distichously arranged in the form of a fan. Leaves subcoriaceous, linear-lanceolate, acuminate, 8—30 cm. long and 1.0-2.2 cm. wide, the broad, conduplicate, chartaceous bases closely imbricating and persistent. Inflorescences usually about 1-3, slender, 1-flowered scapes 6—9 cm. tall, enveloped in several papery, tubular, acuminate bracts. Flowers relatively small, dull purplish red or reddish brown. Sepals subequal, free, spreading, elliptic-lanceolate, acute, 14-18 mm. long and 5.5-7.3 mm. wide, the laterals somewhat obliquely inserted on the short column foot, forming an incon- spicuous, rounded mentum. Petals subequal to the dorsal sepal, elliptic-oblanceo- late, acute, 10-13 mm. long and 3.5—5 mm. wide. Lip entire or very obscurely 3-lobed, oblong-ovate, obtuse or subacute, 10-14 mm. long and 2.5—3.5 mm. wide, contracted at the base and articulated with the short column foot, the lateral margins somewhat involute, the apical lobe ovate to oblong-ovate, obtuse to subacute, spreading. Column semi-terete, somewhat arcuate, 6—7.5 mm. long, produced at the base into a short foot. Costa Rica and Panama. cocLÉ: vic. El Valle de Antón, 600-1000 m., Allen 1256, 2074, 4450. A curious species, unlike most other Maxillarias, the plants somewhat reminis- cent of a Chondrorhyncha. The last two collections cited have shorter and nar- rower leaves and somewhat smaller flowers than those of the type, but otherwise appear to be more nearly referable to this species than to any other. 12. MAXILLARIA CONDUPLICATA (А. & 5.) L. Wms. in Ann. Mo. Bot. Gard. 29:348. 1942 Ornithidium conduplicatum Ames & Schweinf. in Sched. Orch. 8:66, #. 5. 1925. Stout, erect, epiphytic herbs with cylindric, ascending stems apparently often branching and enveloped in 2-ranked foliage; the base of the cane with one or (443) [Vor. 36 108 ANNALS OF THE MISSOURI BOTANICAL GARDEN more stout, ovoid, monophyllous pseudobulbs, in our specimen 5.5 cm. tall and 5 cm. wide. Mature leaves absent in our specimen, but apparently large, numerous and ligular, the upper stem covered by the broad, coarse, com- planate, 2-ranked, imbricating, per- sistent bases. Inflorescences short 1-flowered scapes about 2.5 cm. long, produced in dense fascicles from the leaf axils. Flowers small. Sepals sub- equal, free, apparently not spreading, elliptic-lanceolate, acute, about 6 mm. long and 2-2.5 mm. wide, the laterals somewhat oblique and concave at the base. Petals subequal to the dorsal sepal, elliptic-obovate, acute, about 5 mm. long and 2 mm. wide. Lip cana- liculate, with a deep median constric- tion, more or less subquadrate and 4-lobed when spread out, about 4 mm. long and 2.6 mm. wide, the narrowed base adnate to the base of the column, the lateral lobes rounded, erect in natural position, the mid-lobe suborbicular, con- spicuously emarginate at the apex, the lateral concave, rounded lobules erect, the median constriction with a transverse, fleshy callus. Column very short, stout, about 1.75 mm. long. Panama. CHIRIQUÍ: Раю Alto Hill, 4000-5000 ft., Powell 341. Fig. 180. Maxillaria conduplicata Known only from the type collection. 13. MaXILLARIA CONFUSA Ames & Schweinf. in Sched. Orch. 8:57. 1925. Small, erect, epiphytic herbs up to about 25 cm. tall, with clustered, elliptic- oblong, compressed, monophyllous pseudobulbs 12-18 mm. tall and 8-12 mm. wide, the bases enveloped in several imbricating bracts, the upper pair of which is usually foliaceous. Leaves coriaceous, grayish green, ligular to elliptic- lanceolate, acute, 7-16 cm. long and 2-3.5 cm. wide, contracted below into elongate, slender, conduplicate petioles. Inflorescences usually 6-8 slender, 1- flowered scapes 3—4 cm. long, produced from the base of the pseudobulbs. Flowers of moderate size, but large in relation to the size of the plants. Sepals subequal, free, spreading, white, linear-lanceolate, abruptly acute, 17-20 mm. long and 4—5 mm. wide, the oblique bases of the laterals inserted on the elongate column foot, forming a conspicuous acuminate mentum. Petals subequal to the dorsal sepal, white, from an oblique base lanceolate-acuminate, 15—17 mm. long and 4—5 mm. wide. Lip canaliculate, reddish brown, obscurely 3-lobed near the apex, oblong- (444) 1949] FLORA OF PANAMA (Orchidaceae) 109 obovate, about 12 mm. long and 6 mm. wide when spread out, contracted at the base and articulated with the foot of the column, the lateral margins erect, the apices rounded, mid-lobe ovate, acute, about "4 the total length of the lip, con- spicuously thickened and with a tubercle on the under-surface of the apex; disk with a fleshy linguiform callus, about 24 the length of the lateral margins. Column short, stout, semi-terete, somewhat arcuate, produced at the base into a long narrow foot. Costa Rica and Panama. acuas: forested slopes of Cerro Tuté, region west of Santa Fé, 2500 ft., Allen 9 Fairchild 4405. It seems quite possible that this species, as well as Maxillaria brevipes and M. arachnitiflora, may prove to be but well-marked varieties of Maxillaria Reichen- beimiana. 14. MAXILLARIA CRASSIFOLIA (Lindl.) Rchb. f. in Bonplandia 2:16. 1854. Epidendrum sessile Swartz, Prodr. Veg. Ind. Occ. 122. 1788, non Maxillaria sessilis Lindl. Heterotaxis crassifolia Lindl. in Bot. ќа 12: Е. 1028. 6 Dicrypta Baueri Lindl. Gen. & Spec. Orch. Pl. 44. 1830 Dicrypta crassifolia Lindl. ex Loud. Hort. Brit. Suppl. 3: 536. 1839. Maxillaria sessilis Fawcett & Rendle, Fl. Jam. 1:120. 1910, non Lindley. Maxillaria gatunensis Schltr. in Fedde Rep. Sp. Nov. Beih. 17:68. 1922. Erect, caespitose, epiphytic herbs averaging about 30 cm. tall. Leaves fleshy, linear-lanceolate, obtuse to acute, 6—35 cm. long and 1-3.8 cm. wide, the con- duplicate bases imbricating and forming a short, more or less complanate petiole, sometimes enveloping an inconspicuous, poorly developed, oblong, monophyllous pseudobulb, all but the leaf of the pseudobulb actually being foliaceous bracts. Inflorescences usually short, solitary, 1-flowered scapes, nearly sessile in the upper bract axils. Flowers of moderate size. Sepals subequal, free, spreading, usually yellow or pale yellow, lanceolate, acute, 16—20 mm. long and 5—6 mm. wide, the dorsal sepal somewhat concave, the laterals adnate to the column foot, forming a short, rounded mentum. Petals usually yellow, subequal to the dorsal sepal, linear- lanceolate to oblanceolate, acute, 14-17 mm. long and 3—4 mm. wide. Lip nearly entire or obscurely 3-lobed, yellow with red spots to dark red, concave, 12—14 mm. long and 5—7 mm. wide when spread out, contracted at the base and articu- lated with the column foot, lateral margins erect, the mid-lobe about / to V2 the total length of the lip, somewhat thickened and minutely papillose; the disk with a fleshy, thickened keel. Column elongate, semi-terete, somewhat arcuate, 7—9 mm. long, produced at the base into a short foot. Mexico, Guatemala, British Honduras, Honduras, Costa Rica, Panama, Colombia, Venezuela, Brazil, Florida, Cuba, Jamaica, Hispaniola and probably other adjacent areas. п Lake, sea level, Powell 207. PANAMA: San José Island, Perlas Жыр. Due PR 1275-А. CHIRIQUÍ: without definite locality. 4000 ft., Powell (445) [ Vor. 36 110 ANNALS OF THE MISSOURI BOTANICAL GARDEN 118. BOCAS DEL TORO: without definite locality, von Wedel 477. DARIÉN: vic. Pinogana, Rio Tuira, 20 m., Allen 917. A very common, weedy species widely distributed throughout the lowlands of our area. 15. MAXILLARIA CTENOSTACHYA Rchb. f. in Gard. Chron. 39. 1870. Maxillaria ctenostachys Rchb. f. ex чо іп Beih. Bot. се i :495. 1918. Camaridium arachnites Schltr. in Fedde Rep. Sp. Nov. Beih. 922. Camaridium stenostachys Schltr. loc. cit. 19: 238. 1923, Elongate, slender, erect or pendulous, epiphytic herbs. Pseudobulbs elliptic, ovoid, complanate, rugose, tapering, diphyllous, those at the base of the plant clustered, 3.5—6.5 cm, tall and 2.5—4 cm. wide, those on the upper branching stem becoming very distant and inserted at an acute angle, 2.5—5 cm. tall and 1.5-2 cm. thick, the bases provided with several elongate, foliaceous bracts; the long internodes closely enveloped in long, acuminate, persistent, papery bracts. Leaves coriaceous, linear-lanceolate, acute, 15—35 cm. long and 1—2.5 cm. wide, contracted below into short conduplicate petioles. Inflorescences usually many short, slender, 1-flowered scapes 1.5-2 cm. long, from the bract axils of the dis- tichous flush of new growth, which at flowering time much resembles the stem of a Lockbartia in vegetative appearance. Flowers relatively large, most or all being produced. simultaneously, the flowering habit reminiscent of that of МахШана alba. Sepals subequal, free, more or less spreading, white or pale yellow, lanceolate, long-acuminate, 2.5—4 cm. long and .4-.5 cm. wide; the laterals adnate to the column foot, forming a short, rounded mentum. Petals subequal to the sepals, white or pale yellow, rather obliquely elliptic-lanceolate, long-acuminate, 2—3 cm. long and .3—5 cm. wide. Lip conspicuously 3-lobed, white or pale yellow, the inner surface with brown or tan markings, 8-12 mm. long and 6-8 mm. wide when spread out, contracted at the base and articulated with the foot of the column, lateral lobes erect in natural position, the anterior margins acute, mid- lobe ovate, acute, about 1⁄2 the total length of the lip, the apex reflexed and some- what thickened; disk with 5 erect, parallel keels, about 34 the length of the lateral lobes, the central 3 of which are rounded at the apex and conspicuously thickened. Column semi-terete, somewhat arcuate, 4—6 mm. long, produced at the base into a short foot. Costa Rica and Panama. cHiRIQUÍ: Río Caldera, 4500 ft., Powell 210. 16. MAXILLARIA CUCULLATA Lindl. in Bot. Reg. 26: 7. 12. 1840. Maxillaria meleagris Lindl. loc. cit. 30: Misc. 3. 1844. Maxillaria Lindeniana Rich. & Gal. in Ann. Sci. Nat. Bot. III, 3:24. 1845. Maxillaria obscura Linden & yes f. in Rchb. Beitr. Orch. Centr.-Amer. 31. 1866. Maxillaria puncto-striata Rchb. f. in Linnaea 41:28. 1877 (446) 1949] FLORA ОЕ PANAMA (Orchidaceae) 111 Erect, epiphytic herbs with clustered, oblong-ovoid to elliptic-ovoid, com- pressed, rugose, monophyllous pseudobulbs 2.5—4 cm. tall and 1.5—2.5 cm. wide, the bases enveloped in several imbricating, papery bracts. Leaves coriaceous, acute, or retuse and obscurely 2-lobed at the apex, 15—30 cm. long and 1.5—2 cm. wide, contracted below into a conduplicate petiole. Inflorescences usually 3-4 short to elongate, 1-flowered scapes from the base of the pseudobulbs, enveloped in num- erous, often spreading, conspicuous, tubular, papery, acuminate bracts. Flowers of moderate size, dark reddish or purplish brown with a maroon lip, or with yellow sepals and petals which may be marked with varying amounts of the darker color. Sepals free, subequal, spreading, lanceolate to elliptic-lanceolate, acute, 1.8—2.2 cm. long and .4-.6 cm. wide, the laterals adnate to the foot of the column, forming a short, subacute mentum. Petals lanceolate, acute, 1.5—1.8 cm. long and .4—6 cm. wide. Lip conspicuously 3-lobed, 1.4-1.6 cm. long and .6—.8 cm. wide when spread out, contracted at the base and articulated with the foot of the column, the lateral lobes short, conspicuously erect in natural position, the mid-lobe elliptic-lanceolate to elliptic-ovate, acute to subacute, about ?4 the total length of the lip, the apex thickened, with a fleshy keel on the under-surface; disk with a broad, obtuse, fleshy callus about equaling the lateral lobes in length. Column short, stout, somewhat arcuate, rather dilated at the apex, produced at the base into a foot. Mexico, Guatemala, Nicaragua, Costa Rica, Panama, and probably adjacent territories. CHIRIQUÍ: Bajo Chorro, in rain forest, 6000 ft., Davidson 115; in woods on Llano del Volcán, 1500 m., Allen 3504 17. MAXILLARIA DENDROBIOIDES (Schltr.) L. Wms. in Ann. Mo. Bot. Gard. 27:283. 1940. Camaridium dendrobioides Schltr. in Beih. = ТРЭ 362:415. 1918. Camaridium Jimenezii Schltr. loc. ск. 416. Camaridium simile Schltr. in Fedde Rep. Sp. Nov. Beih. 19:239. 1923. Slender, erect or pendulous, epiphytic herbs without pseudobulbs, 30—45 cm. tall, the stems usually branching, the lower portions enveloped in the closely imbricating, papery leaf bases; the foliage of the apex 2-ranked. Leaves spreading, coriaceous, ligular, the apices usually retuse or rather unequally 2-lobed, very variable in size, 1—6 cm. long and .25-1.0 cm. wide (in our specimens 1.5—2 ст. long and about .3 cm. wide), the conduplicate bases closely imbricating and per- sistent, enveloping the stems. Inflorescences short, slender, 1-flowered scapes 10-15 mm. long, produced from the leaf axils. Flowers small. Sepals free, sub- equal, apparently not spreading, lanceolate, acute or obtuse, 9—11 mm. long an 2—2.5 mm. wide, the laterals somewhat obliquely inserted on the column foot, forming a short, rounded mentum. Petals subequal to the dorsal sepal, rather obliquely elliptic-oblanceolate, obtuse or shortly acute, 7-8.5 mm. long and 2-2.5 mm. wide. Lip obscurely 3-lobed, about 7.5 mm. long and 3.5 mm. wide when (447) [Vor. 36 112 ANNALS OF THE MISSOURI BOTANICAL GARDEN spread out, contracted at the base and articulated with the column foot, the lateral lobules or margins rounded, erect in natural position, the mid-lobe ovate to ligular, obtuse, about !⁄ the total length of the lip; disk with 3 parallel, thickened nerves, about / the length of the lateral margins. Column semi-terete, about 3.5 mm. long, produced at the base into a short foot. Costa Rica and Panama. CHIRIQUÍ: vic. Bajo Chorro, in heavy rain forest, 6000 ft., Davidson 240. 18. MAXILLARIA DIUTURNA Ames & Schweinf. in Sched. Orch. 8:58. 1925. Erect or pendent, epiphytic herbs with elliptic-oblong to elliptic-ovate, strongly ancipitous, monophyllous pseudobulbs 2-3 cm. long and .1—2 cm. wide, rather distant and inserted at an acute angle on the stems; the internodes and bases of the pseudobulbs enveloped in closely imbricating, persistent bracts the upper pair of which is foliaceous. Leaves persistent, coriaceous, ligular to elliptic-lanceo- late, acute ог minutely and unequally 2-lobed, 6—15 cm. long and 1.2-3 cm. wide, contracted below into short, conduplicate petioles. Inflorescences short, slender, 1-flowered scapes, apparently produced singly in succession from the bract axils of the flush of new growth. Flowers pseudo-campanulate, of moderate size. Sepals subequal, free, not spreading, bright yellow or sometimes reddish yellow on the outer surfaces, 12-14 mm. long and 5-6 mm. wide, ovate-lanceolate, acute, con- cave, with a low central keel, the laterals adnate to the column foot forming a prominent, rounded mentum. Petals elliptic-ovate, acute, bright yellow, 10—12 mm. long and 6—7 mm. wide. Lip entire, bright yellow with scarlet markings, oblong-pandurate, 10—12 mm. long and 5-6 mm. wide when spread out, somewhat contracted at the base and articulated with the column foot, lateral margins some- what erect in natural position, the apex truncate, obscurely emarginate and 2- lobed, the lobules somewhat erect in natural position; the disk with a linear to oblong, obtuse, fleshy callus. Column semi-terete, somewhat arcuate, 5-6 mm. long, produced at the base into a foot. Costa Rica and Panama. рен cloud forest оп the summit of Cerro Campana a, 3000 ft, Allen 5183. co Cativo-Porto Bello trail, sea level, Powell 376. сосіЕ: vic. La Mesa, region wat of El Valle de Antón, 1000 m., Allen 2306, 2035. 19. MAXILLARIA EXALTATA (Krinzl.) C. Schweinf. in Bot. Mus. Leafl. Нагу. Univ. 11:272. 1945. Camaridium exaltatum Kránzl. in Engler’s Bot. Jahrb. 37:386. 1906. Tall, erect, epiphytic herbs, the cylindric, sometimes branching stems 60—75 cm. tall, apparently entirely without pseudobulbs, the lower portions enveloped in the closely imbricating, persistent, chartaceous leaf bases which ultimately weather to loose fibers, the upper stem with 2-ranked foliage. Leaves coriaceous, linear-lanceolate, acuminate, 9—25 cm. long and .8—1.5 cm. wide, the broad, con- duplicate bases imbricating and equidistantly distributed on the stem. Inflores- (448) 1949] FLORA OF PANAMA (Orchidaceae) 113 cences usually several, slender, 1-flowered scapes 5—6 cm. long, produced from the upper leaf axils. Flowers of moderate size. Sepals free, subequal, spreading, pale pinkish tan (in our specimens), the dorsal sepal linear-lanceolate, acute, with a central keel, 2-2.5 cm. long and .35-5 cm. wide, the laterals ligular, acute, 2.3-2.8 cm. long and .5—7 cm. wide, the bases rather oblique and adnate to the column foot, forming a prominent acute mentum. Petals linear-lanceolate, acute, pale pinkish tan (in our specimens), 2-2.2 ст. long and .3—4 cm. wide. Lip 3-lobed near the apex, reddish-brown, 13-15 mm. long and 6—8 mm. wide when spread out, contracted at the base and articulated with the column foot, lateral lobes erect in natural position, the anterior margins subacute and somewhat projecting, the mid-lobe ovate, abruptly acute, about / the total length of the lip, conspicu- ously thickened and minutely papillose, with a prominent tubercle on the under- side of the apex; the disk with a ligular, rather concave, obtuse callus, about 34 the length of the lateral lobes, the apex rather conspicuously thickened. Column short, stout, semi-terete, produced at the base into a long foot. Panama and Peru. COCLÉ: mossy forest on the crest of Cerro Pajita, hills north of El Valle de Antón, 1200 m., Allen 9 Fairchild 3945. This collection represents a tremendous extension of range for this rather poorly known Peruvian species. Our specimens match the excellent photograph of the type and the type description in the Ames Herbarium almost exactly, even to the size of the floral parts, which is remarkable in view of the wide separation of the areas of collection. It is of course to be expected that the species also exists in Colombia and Ecuador, and will eventually be collected there. 20. MAXILLARIA FRIEDRICHSTHALII Rchb. f. in Bot. Zeit. 10:858. 1852. Maxillaria aciantha Rchb. f. loc. cit. 1852 Lycaste aciantha Rchb. f. in Bonplandia 3:216. 1855 Maxillaria turialbae Schltr. in Beih. Bot. Centralbl. 362:414. 1918. Maxillaria rbodosticta Kranzl. in Fedde Rep. Sp. Nov. 24:223. 1928. Erect or semi-pendent, epiphytic herbs with complanate, elliptic-oblong, rugose pseudobulbs 1.5—5 cm. tall and 7-15 mm. wide, approximate and inserted at an acute angle on the usually arching rhizome; the short internodes and bases of the pseudobulbs enveloped in the persistent, chartaceous bases of the closely imbricat- ing bracts, the upper pair of which is usually foliaceous. Leaves coriaceous, 2-3 (very rarely 1 or 4) from the apex of the pseudobulbs, ligular, the apex obscurely and unequally 2-lobed, 3.5-18 cm. long and .7-1.5 cm. wide, contracted below into very short, conduplicate petioles. Inflorescences 1 to several, often loosely fasciculate, 1-flowered scapes 1.8-4 cm. long, produced from the base of the mature pseudobulb, the peduncles closely enveloped in numerous broad, acute, imbricating, chartaceous bracts. Flowers small to relatively large, the size ap- parently correlated with the size of the plant. Sepals subequal, free, not spread- (449) [Vor. 36 114 ANNALS OF THE MISSOURI BOTANICAL GARDEN ing, linear-lanceolate, acute to acuminate, 1.5-3 cm. long and .3-.6 cm. wide, greenish yellow to greenish lavender, the laterals adnate to the column foot, form- Fig. 181. Maxillaria Friedricbstbalii ing a short, acute mentum. Petals lanceolate to linear-lanceolate, acute to acuminate, greenish yellow, 1.2-2.5 cm. long and .2—3 cm. wide. Lip entire, linear-oblanceolate, obtuse to subacute, canaliculate, 1.2-2.5 cm. long and .3-.6 cm. wide, usually pale yellowish green, the long narrow base articulated with the foot of the column, the lateral margins somewhat erect, the apex conspicuously thickened, usually dark maroon to nearly black; the disk with a rather obscure, yellow central keel, which extends from the base about 25 the total length of the lip. Column elongate, semi-terete, somewhat arcuate, 8—20 mm. long, produced at the base into a short foot. (450) 1949] FLORA OF PANAMA (Orchidaceae) 115 Mexico, British Honduras, Guatemala, Honduras, Nicaragua, Costa Rica, Danama, and probably adjacent South America. CANAL ZONE: Barro Colorado Island, Shattuck 543. PANAMA: vic. Paja, sea level, Powell 3210; Cerro Campana, vic. Campana, 800 m., Allen 4025. СОСТЕ: vic. El Valle de Antón, 600-1100 m., Allen 1254, 1257, 3992; Bismarck, 2000-3000 ft., Williams 444. cHIRIQUÍ: without definite locality, 4000-5000 ft., Powell 3516; Bajo Mono, 4500 ft., Davidson 524 A very common species, widely distributed throughout the lowlands and inter- mediate highlands of our area. 21. MAXILLARIA FULGENS (Rchb. f.) L. Wms. in Ann. Mo. Bot. Gard. 28:425. 1941. Ornitbidium fulgens Rchb. f. Beitr. Orch. Centr.-Amer. 76. 1866. Erect, epiphytic herbs 40—75 cm. tall, with elongate, often branching, cylin- dric, woody canes usually arising from a basal cluster of a few stout, ovoid, fleshy pseudobulbs 3.5—4 cm. tall and 2-3 cm. wide; the long internodes or basal por- tions of the stems closely enveloped in the persistent, imbricating leaf bases, or becoming naked after these have weathered away; the apex of the cane with 2-ranked foliage. Leaves coriaceous, linear-lanceolate, acute, 12-38 cm. long and 1.5—3 cm. wide, forming a compact, distichous fascicle at the apex of the stems, portions of which are often still in evidence at the earlier nodes along the more elongate canes, the conduplicate leaf bases sometimes enveloping a small, poorly developed pseudobulb. Inflorescences usually about 15—30 slender, 1-flowered scapes, 1.5—3 cm. tall, in dense subsessile fascicles from the lower leaf axils. Sepals rather fleshy, subequal, free, not spreading, broadly ovate, acute, concave, reddish orange, 7-8 mm. long and 3-3.5 mm. wide, the laterals adnate to the somewhat produced base of the column, forming a short, rounded mentum. Petals lanceolate, acute, 5-6 mm. long and about 2.5 mm. wide, suffused reddish orange shading to yellow at the base. Lip entire, bright yellow or orange, 4-4.5 mm. long and 1.5-2 mm. wide, geniculate and more or less sigmoid in profile, the orbicular basal half deeply concave or subsaccate, adnate to the base of the column, the strongly re- flexed apical half elliptic-oblong, obtuse, rather fleshy, with thin, erect, often somewhat undulant margins, the apex becoming rather shallowly emarginate, with a short, acute, fleshy, carinate projection on the under-surface. Column semi- terete, somewhat arcuate, bright yellow, 2.5—3 mm. long, the base without a foot. Costa Rica and Panama. СТЕ: region north of El Valle de Antón, vic. La Mesa and La Loma del Tigre, about Р = Allen 2256, 2388, 3770. снікюші: Caramilla, 5000 ft., Powell 283. A frequent, attractive species of the wet highland forests of Coclé and Chiriquí provinces. 22. MAXILLARIA INAUDITA Rchb. f. Beitr. Orch. Centr.-Amer. 76. 1866. Robust, erect or pendulous, epiphytic herbs 45—60 cm. tall, the elongate (451) [VoL. 36 116 ANNALS OF THE MISSOURI BOTANICAL GARDEN cylindric canes provided with distichous, spreading foliage at the apex, the basal portions enveloped in the broad, complanate, persistent, imbricating leaf bases; the plants seemingly without pseudobulbs but sometimes with 1 or more elliptic- oblong, strongly ancipitous, monophyllous pseudobulbs up to 8 cm. tall and 4.5 cm. wide at the base of the leafy cane. Leaves coriaceous, ligular to elliptic- lanceolate, broadly obtuse, 15—25 cm. long and 3.5—4.5 cm. wide, contracted at the base into narrow conduplicate petioles, the broader rugose bases persistent and equidistantly distributed on the stems. Inflorescences short 1-flowered scapes 4—5 cm. long, produced from the axils of the leaves. Flowers relatively large and conspicuous. Sepals subequal, free, spreading, lanceolate, acuminate, white, 2.5—4 cm. long and 6-8 mm. wide, the laterals adnate to the column foot, forming an inconspicuous, rounded mentum. Petals lanceolate, acuminate, white, 2.2-3 cm. long and .4—5 cm. wide. Lip rather obscurely 3-lobed, ovate, obtuse to subacute when spread out, 1.5 cm. long and 1 cm. wide, contracted at the base into a short claw which is articulated with the foot of the column, the lateral lobes white, erect in natural position, the mid-lobe ovate, obtuse to subacute, yellow, about 15 the total length of the lip, separated from the lateral lobes by plicate folds; the disk with a concave, ligular, obtuse callus which is about 3⁄4 the length of the lateral lobes. Column semi-terete, somewhat arcuate, 6-8 mm. long, produced at the base into a short foot. Costa Rica and Panama. VERAGUAS: forested slopes of Cerro Tuté, region west of Santa Fé, 3000 ft., Allen t$ Fairchild 4338. 23. MAXILLARIA LONGIPETIOLATA Ames & Schweinf. in Sched. Orch. 8:61. 1925. Erect, epiphytic herbs about 23 cm. tall, with elliptic-ovoid, compressed, some- what finely wrinkled monophyllous pseudobulbs 2.5 cm. tall and 1.5 cm. wide, which apparently are approximate on an abbreviated rhizome, the bases enveloped in several papery bracts which become fibrous with age. Leaves subcoriaceous, linear-lanceolate, acuminate, 16 cm. long and 2 cm. wide, contracted below into an elongate, slender, conduplicate petiole. Inflorescences apparently short, soli- tary, l-flowered scapes from the base of the pseudobulbs. Flowers of moderate size. Sepals subequal, free, spreading, oblong-lanceolate, acute, Indian red, 16—18 mm. long and 5.5-5.8 mm. wide. Petals subequal to the dorsal вера], Indian red, linear-lanceolate, rather obliquely acute, 1.5—2.5 cm. long and about 3 mm. wide. Lip conspicuously 3-lobed, dark maroon, about 1.4 cm. long and .8 cm. wide when spread out, contracted at the base and articulated with the foot of the column, the lateral lobes erect in natural position, the acute apices spreading, the mid-lobe ligular, about V2 the total length of the lip, the rounded apex lightly retuse; the disk with a linear-lanceolate callus about 24 the length of the lateral lobes, the obtuse apex rather fleshy. Column semi-terete, somewhat arcuate, about 7 mm. long, produced at the base into a short foot. Anther yellow. (452) 1949] FLORA OF PANAMA (Orchidaceae) 117 Г ZW Fig. 182. Maxillaria luteo-alba (453) [Vor. 36 118 ANNALS OF THE MISSOURI BOTANICAL GARDEN Panama. PANAMÁ: hills east of Corozal, sea level, Powell 307. Known only from the type collection. 24. MAXILLARIA LUTEO-ALBA Lindl. Orch. Linden. 20. 1846. Maxillaria luteo-grandiflora Hort. in Flor. Mag. 10: Z. 550. 1871. Erect, epiphytic herbs with stout, oblong-ovoid to elliptic-ovoid, somewhat compressed, monophyllous pseudobulbs 3—5 cm. tall and 2—3 cm. wide, the bases enveloped in several long, papery, imbricating bracts which become fibrous with age. Leaves coriaceous, linear-lanceolate to elliptic-lanceolate, acute, 25—50 cm. long and 2.5-4 cm. wide, contracted below into short or elongate, conduplicate petioles. Inflorescences usually about 3 erect, 1-flowered scapes 9-14 cm. long, produced from the bases of the pseudobulbs, enveloped in several papery, acute, tubular bracts. Flowers conspicuous, the largest of the genus in Panama. Sepals subequal, free, widely spreading, ligular, broadly obtuse, white on the outer sur- faces and pale yellow within, 4-5 cm. long and 1.0—1.2 cm. wide, the laterals adnate to the column foot, forming a prominent, acute mentum. Petals ligular, shortly subacute, the apices incurving in natural position, white without and pale yellow within, 3.5—4 cm. long and .5—7 cm. wide. Lip conspicuously 3-lobed, deep yellow, margined white, about 2.5 cm. long and 1.8 cm. wide when spread out, contracted at the base and articulated with the foot of the column, the lateral lobes erect in natural position, the rounded apices somewhat projecting, mid-lobe ovate, obtuse, about 14 the total length of the lip; the disk with a linguiform, laterally bicarinate, acute callus, about 24 the length of the lateral lobes. Column semi-terete, somewhat arcuate, about equaling the length of the lateral lobes of the lip. Costa Rica, Panama, Colombia, and Ecuador. СТЕ: hills north of El Valle de Antón, crest of Cerro Pajita and La Loma del Tigre, 3000-3200 ft., Allen 2870, 3817, 4192 A common, large-flowered, attractive species of the high rainy hills north of El Valle, and to be expected in other similar situations throughout our range. 25. MAXILLARIA MALEOLENS Schltr. in Fedde Rep. Sp. Nov. Beih. 19:233. 1923. Erect, epiphytic herbs 30-45 cm. tall, with approximate or solitary, strongly ancipitous, monophyllous, oblong-elliptic pseudobulbs 4.5-8 cm. tall and 2-3.5 cm. wide, the bases enveloped іп 3-4 complanate, imbricating, foliaceous bracts. Leaves coriaceous, ligular, broadly obtuse, 15—45 cm. long and 2.5-5 cm. wide, contracted below into conduplicate petioles, the complanate-conduplicate bases distichously arranged in the form of an open fan. Inflorescences usually solitary, 1-flowered scapes 5—8 cm. long, produced from the upper bract axils. Flowers of moderate size, fragrant. Sepals subequal, free, spreading, yellow, elliptic- lanceolate, acute, 2-2.5 cm. long and .8-1.0 wide, the laterals rather obliquely (454) 1949] FLORA OF PANAMA (Orchidaceae) 119 inserted on the column foot, forming a short, rounded mentum. Petals yellow, ligular, the apices obliquely acute, 18—20 mm. long and 4—5 mm. wide. Lip rather obscurely 3-lobed, reddish brown to rich purple throughout, or sometimes with the lateral lobes yellow, 2-2.3 cm. long and 1-1.2 cm. wide when spread out, contracted at the base and articulated with the foot of the column, the lateral lobes or margins rounded, erect in natural position, the mid-lobe ovate, obtuse, rather fleshy, about 14 the total length of the lip, reflexed at the apex; the disk with a narrow, thickened, central nerve, about equaling the lateral lobes in length. Column semi-terete, somewhat arcuate, 12-15 mm. long, produced at the base into a foot. Honduras, Costa Rica, and Panama. NAL ZONE: Gatün Lake, sea level, Powell 276. сосі.Е: region north of El Valle de Penis 3000 ft., Fairchild 5. п. (under Allen 2932). 26. MAXILLARIA MINOR (Schltr.) L. Wms. in Amer. Orch. Soc. Bull. 10:273. 1942. Camaridium minus Schltr. in Beih. Bot. Centralbl. 36?:417. 1918. Erect, epiphytic herbs with slender, branching stems provided with small, very distant, suborbicular, monophyllous, compressed pseudobulbs 6—15 mm. long and 6—15 mm. wide, the bases covered by 2—3 conspicuous, foliaceous bracts, the long internodes very closely enveloped in complanate, imbricating, acuminate, papery sheaths. Leaves subcoriaceous, lanceolate to elliptic-lanceolate, acute, 3.5—15 cm. long and 1—2 cm. wide, contracted below into slender, conduplicate petioles. In- florescences several very short, 1-flowered scapes 5-10 mm. long, apparently produced simultaneously from the bract axils of the flush of new growth. Flowers very small, often enveloped in 2 glumaceous bracts. Sepals subequal, free, not spreading, concave, oblong-lanceolate, acute to subacute, about 4 mm. long and 2 mm. wide, the laterals adnate to the base of the column, forming a rounded, con- spicuous mentum. Petals subequal to the dorsal sepal, ligular, obtuse, about 3 mm. long and 1.5 mm. wide. Lip fleshy, 3-lobed, about 3 mm. long, articulated with the base of the column, the lateral lobes rounded, erect, on either side of a basal concavity which has a short, ligular, fleshy callus, the mid-lobe ovate, subacute, nearly ! the total length of the lip, the basal constriction below the lateral lobes with a transverse, semicircular thickening. .Column short, stout, the lateral mar- gins thickened toward the obscurely produced base. Costa Rica and Panama OCLÉ: mossy forest on summit of Cerro Pajita, hills north of El Valle de Antón, 1100 m., Allen 3001. 27. MAXILLARIA NEGLECTA (Schltr.) L. Wms. іп Ann. Mo. Вог. Gard. 29:348. 1942. Ornit bidium э Rchb. f. Beitr. Orch. Centr.-Amer. 75. 1866, поп Maxillaria anceps Ornitbidium neglectum Schltr. in Fedde Rep. Sp. Nov. Beih. 19:242. 1923. (455) [Vor. 36 са ) N M Fig. 183. Maxillaria neglecta (456) 1949] FLORA OF PANAMA (Orchidaceae) 121 Erect or pendulous, epiphytic herbs, of very variable vegetative habit, usually with elongate, often branching rhizomes. Pseudobulbs monophyllous, ligular to suborbicular, strongly ancipitous to very thick and fleshy, 1.5—4.5 cm. long and .5—1.5 cm. wide, approximate to very distant, inserted at an acute angle on the stems, the long cylindric internodes and the bases enveloped in closely imbricating, papery bracts. Leaves subcoriaceous, ligular, acute, 6—18 cm. long and 1.2—2.5 cm. wide, the bases contracted into very short conduplicate petioles. Inflorescences very short, 1—flowered scapes, produced in dense subsessile fascicles from the bases of the mature pseudobulbs. Flowers small, usually enveloped in 2 glumaceous bracts. Sepals free, apparently not spreading, concave, yellow or white, about 6 mm. long, the dorsal sepal elliptic-ovate, acute, the laterals broadly rhombic with oblique, acute apices, the oblique bases forming an acute or rounded, conspicuous mentum. Petals subequal to the dorsal sepal, oblong-ligular, subacute, about 4.5 mm. long, similarly colored with the sepals. Lip yellow, about 6 mm. long and 3 mm. wide when spread out, geniculate in profile, the long narrow base continuous with the base of the column, the limb dilated and conspicuously 3-lobed near the apex, the obliquely triangular lateral lobes erect, joined below the apices by a narrow transverse callus, the mid-lobe rather fleshy, triangular to broadly ovate, acute to obtuse, inserted at an abruptly deflexed angle below the apices of the lateral lobes. Column short, about 1.7 mm. long, the base continuous with the claw of the lip. Honduras, Nicaragua, Costa Rica, and Panama. CANAL ZONE: hills north id Bar Td 27669. PANAMÁ: hills east of Panama City, sea level, Powell 324. : Río uení, upper Madden Lake region, 200 ft., Fairchild s. n.; Cerro Santa Rita 1 їс Allen & Fairchild 5101. cocrÉ: vic. El Valle de Antón, 800—1000 m., Allen 780. cHimiQuí: upper Río Chiriqui Viejo, 1300— E m., White ед White 37; D vds Lérida, ie i slopes of Chiriquí Volcano, 5000 ft., К. Morris s. . Bajo o and Quebrada Chiquero, 1500 m., Woodson & Schery 563; Bajo M Robalo = western slopes of Cerro Horqueta, 5000-7000 ft., Allen 4783. A common species of very variable vegetative habit, widely distributed throughout our area. 28. MAXILLARIA OREOCHARIs Schltr. in Fedde Rep. Sp. Nov. Beih. 17:69. 1922. Erect or pendulous, epiphytic herbs. Pseudobulbs strongly ancipitous, oblong- elliptic, monophyllous, 3-4 cm. long and 1.0-1.5 cm. wide, approximate and often more or less imbricating, obliquely inserted on the rhizome, the bases provided with several imbricating bracts, the upper 1—3 of which are conspicuously long- foliaceous, the short internodes enveloped in closely imbricating, more or less complanate, persistent, acute, papery bracts. Leaves subcoriaceous, linear, acute or acuminate, 25—40 cm. long and .6—1.0 cm. wide, contracted below into very short, conduplicate petioles. Inflorescences usually solitary, slender, 1-flowered scapes 3—4 cm. long, produced from the base of the mature pseudobulbs. Sepals subequal, free, more or less spreading, lanceolate, acute, white with red markings, (457) [Vor. 36 122 ANNALS OF THE MISSOURI BOTANICAL GARDEN about 1.5 cm. long and .3-.4 cm. wide, the laterals adnate to the column foot, forming an inconspicuous, subacute mentum. Petals subequal to the dorsal sepal, lanceolate, acute, white, about 12 mm. long and 2.5-3 mm. wide. Lip oblong- ligular, acute, 3-lobed near the apex, about 12 mm. long and 5 mm. wide when spread out, the base articulated with the foot of the column, the lateral lobes erect in natural position, dark red, the mid-lobe ovate, subacute, somewhat thickened, about № the total length of the lip, yellow, the upper surface and margins minutely papillose-puberulent; the disk with a linear, obtuse, fleshy callus, about 3⁄4 the length of the lateral lobes. Column slender, semi-terete, somewhat arcuate, the apex dilated, about 7 mm. long, white, the base produced into a foot. Costa Rica and Panama. cHIRIQUÍ: without definite locality, 4000—5000 ft., Powell 256. 29. MAXILLARIA PARVILABIA Ames & Schweinf. in Sched. Orch. 8:62. 1925. Erect, epiphytic herbs without pseudobulbs; the woody, often branching, cylindric canes 1-1.25 m. tall, the lower portions closely enveloped in the persistent leaf bases, the apices with 2-ranked foliage. Leaves coriaceous, linear-lanceolate, obtuse or retuse and obliquely 2-lobed at the apex, 4-15 cm. long and .6—1.2 cm. wide, the tubular, clasping bases closely imbricating and equidistantly distributed on the stems. Inflorescences few to many slender, 1-flowered scapes about 4 cm. long, produced singly or in loose fascicles from the leaf axils. Flowers pale yellow, of moderate size. Sepals subequal, free, spreading, lanceolate, acute, 1.6-2 cm. long and .4-.5 cm. wide, the laterals adnate to the foot of the column, forming a short, subacute mentum. Petals subequal to the dorsal sepal, elliptic-lanceolate, acute, 1.3-1.5 cm. long and .4-.5 cm. wide. Lip small, obscurely 3-lobed, elliptic- obovate, 5—6 mm. long and 3—3.5 mm. wide when spread out, contracted at the base and articulated with the foot of the column, the lateral margins somewhat erect in natural position, distinct only at the small acute apices, mid-lobe fleshy, ovate, acute, about 25 the total length of the lip, the apex lightly recurved; the disk with a small fleshy linguiform callus, about И, the length of the lateral lobes. Column short, stout, semi-terete, about 2.5 mm. long, produced at the base into a short foot. Costa Rica and Panama. cHIRIQUÍ: vic. Bajo Chorro, in rain forest, headwaters of the Rio Caldera, 6000 ft., Davidson 126. 30. MAXILLARIA Рітттекі (Ames) L. Wms. in Ann. Mo. Bot. Gard. 29:349. 1942. Ornithidium Pittieri Ames, in Sched. Orch. 2:35. 1923. Erect or pendulous, epiphytic herbs with woody, branching, cylindric canes up to about 1.5 m. tall, the lower portions naked or enveloped in the persistent, tubu- lar leaf bases, the apex with sparse clusters of 2-ranked foliage, portions of which may persist at distant intervals along the stems. Leaves coriaceous, ligular, acute, 7-15 cm. long and 1—2 cm. wide, the conduplicate bases forming a sheathing (458) 1949] 123 FLORA OF PANAMA (Orchidaceae) \ NY Maxillaria planicola Fig. 184. (459) [Vor. 36 124 ANNALS OF THE MISSOURI BOTANICAL GARDEN petiole. Inflorescences short 1-flowered scapes, produced in dense fascicles from the axils of the leaves. Flowers small, pink. Sepals free, subequal, not spreading, concave, ovate, acute, with a central keel, 8-9 mm. long and about 4 mm. wide, the laterals adnate to the somewhat produced base of the column, forming a short, rounded mentum. Petals subequal to the dorsal sepal, concave, oblong-ovate, acute, 7 mm. long and about 3 mm. wide. Lip fleshy, entire, about 4 mm. long and 2.5 mm. wide, the base deeply concave and adnate to the base of the column, the apex lanceolate, obtuse to subacute, conspicuously thickened. Column short, stout, about 2.5 mm. long, the base with conspicuous lateral thickenings. Costa Rica and Panama. CHIRIQUÍ: Bajo Chorro, in rain forest, headwaters of the Rio Caldera, 6000 ft., Davidson 117 31. MAXILLARIA PLANICOLA С. Schweinf. in Bot. Mus. Leafl. Harv. Univ. 8:188. 1940, Camaridium oe Schltr. in Fedde, Rep. Sp. Nov. Beih. 17:74. 1922, non Maxillaria latifolia Epiphytic herbs with elliptic-ovate to elliptic-oblong, compressed, monophyl- lous pseudobulbs 3.5—6 cm. long and 1.5—4 cm. wide, the internodes of the mature plants 6—8 cm. long and 8-12 mm. thick, enveloped in closely imbricating, tubular, acute, papery bracts, the upper 3—6 of which are complanate-spathaceous, the conduplicate bases covering the lower half of the pseudobulb. Leaves coriaceous, ligular to elliptic-lanceolate, acute, 15—35 cm. long and 3-6 cm. wide, contracted below into narrow, conduplicate petioles. Inflorescences usually 2-4 slender 1- flowered scapes, 4—5 cm. long, produced from the axils of the spathaceous, sheath- ing bracts after the current pseudobulb has nearly matured. Flowers relatively large and conspicuous. Sepals subequal, free, not spreading, concave, the dorsal sepal lanceolate, acute, 3—3.5 cm. long and .8-1.0 cm. wide, the laterals elliptic- lanceolate, acute, 3—3.5 cm. long and 1.0—1.2 cm. wide, adnate to the foot of the column, forming an inconspicuous, rounded mentum. Petals white, lanceolate, acute, 2.8—3.2 cm. long and .6—7 cm. wide. Lip conspicuously 3-lobed, white, 18-20 mm. long and 18-20 mm. wide when spread out, contracted at the base and articulated with the foot of the column, the lateral lobes erect and incurving over the column in natural position, the acute apices projecting, the mid-lobe triangular, acute, about 15 the total length of the lip; the disk with a short, fleshy, ligular, obtuse callus about 3⁄4 the length of the lateral lobes, the basal half yellow, densely and conspicuously pilose. Column semi-terete, somewhat arcuate, 10-12 mm long, produced at the base into a short foot. Panama. ANAL ZONE: vic. San Juan de Pequeni, upper Madden Lake SERA Powell 3438; vic. Vigia and San Juan де Pequení, 66 m., Dodge, 5... 6 Allen 16580. PANAMA: hills east of Panama City, Powell 8; Cares Campana, vic. Campana, on poe rock face, 2500 ft., Allen 5078. corów: vic. Porto Bello, sea lev]: Powell 3458. ( 460 ) 1949, FLORA OF PANAMA (Orchidaceae) 125 32. MAXILLARIA PowELun Schltr. in Fedde Rep. Sp. Nov. Beih. 17:70. 1922. Егесе, epiphytic herbs with approximate, fleshy, ovoid to elliptic-ovoid, com- pressed, monophyllous pseudobulbs 2—4 cm. long and 1.5-2 cm. wide, the bases enveloped in several papery, imbricating bracts which often weather away to loose fibers. Leaves coriaceous, ligular, obtuse to subacute, 15—40 cm. long and 2-3 cm. wide, contracted below into slender, conduplicate petioles. Inflorescences usually 2—4 slender, 1-flowered scapes 4—6 cm. long, produced from the bases of the pseudobulbs. Flowers of moderate size. Sepals subequal, free, spreading, yellow or tan, the dorsal вера! concave, 1.5-1.8 cm. long and .4-.6 cm. wide, ligular to elliptic-lanceolate, obtuse, with a minute apicule, the laterals ligular to elliptic-lanceolate, obtuse, 1.8-2 cm. long and .4-.8 cm. wide, the oblique bases adnate to the foot of the column forming a short, subacute mentum. Petals sub- equal to the dorsal sepal, yellow or tan, rather obliquely ligular, acute, 1.5-1.7 cm. long and .4—.5 cm. wide. Lip conspicuously 3-lobed near the apex, 1.4-1.6 cm. long and .7-.8 cm. wide when spread out, contracted at the base and articulated with the foot of the column, the lateral lobes yellow or tan, erect in natural posi- tion, the acute to obtuse apices projecting, the mid-lobe ovate, acute, fleshy, rugulose, minutely pilose, usually reddish brown, about 15 the total length of the lip, the under-surface of the apex with a triangular keel; the disk with a low, ligular callus about 3⁄4 the length of the lateral lobes. Column semi-terete, some- what arcuate, 7-8 mm. long, produced at the base into a foot. Anther terminal, operculate, incumbent, 1-celled. Panama. CANAL ZONE: vic. Frijoles, Gatün Lake, sea level, Powell 28, PANAMA: Cerro Cam- pana, 2400 ft., Allen 9 Fairchild 3073, 4235, 4237. cocLÉ: vic. El Valle de Antón, 600 m., Allen 2073; mossy forest on the crest of Cerro Pajita, hills north of El Valle, 1200 m., Allen 4254. A common small-flowered species, rather doubtfully distinct from Maxillaria ringens. 33. MAXILLARIA REPENS L. Wms. in Amer. Orch. Soc. Bull. 10:273, #. 0. 1942. Repent, epiphytic herbs with elongate, slender, often branching stems en- veloped in tubular, papery sheaths which often weather away, the caespitose 2- ranked foliaceous growths usually freely rooting from the base, distant or terminal on the stems. Leaves 3-5, coriaceous, 3-10 cm. long and 1-2 cm. wide, elliptic- lanceolate, acute, the conduplicate bases closely imbricating, forming a complanate, sheathing petiole. Inflorescences usually several slender, 1-flowered scapes 1.5-2 cm. long, produced from the axils of the leaves. Flowers small. Sepals subequal, green with reddish brown margins, lanceolate, acute, 7-10 mm. long and 2.5-3.5 mm. wide, the dorsal sepal concave and incurving over the column, the laterals adnate to the foot of the column, widely spreading or reflexed. Petals subequal to and parallel with the dorsal sepal, green with reddish brown margins, linear- lanceolate, acute, 6.5—7.5 mm. long and 2-3 mm. wide. Lip conspicuously 3- (461) [Vor. 36 126 ANNALS OF THE MISSOURI BOTANICAL GARDEN ч MA XII L.AR.IA аны Ты d "eoens adio, nas == AN w S X DS Шаты Fig. 185. Maxillaria repens (462) 1949] FLORA ОЕ PANAMA (Orchidaceae) 127 lobed, 6—8 mm. long and 3.5-4.5 тт. wide, the basal half canaliculate and adnate to the foot of the column, the lateral lobes erect and parallel with the column, the area between the truncate apices with a short, obtuse, concave, fleshy callus; the mid-lobe broadly spreading, subquadrate, strongly reflexed, the truncate apex deeply emarginate, the center orange-yellow with pale yellow, undulate margins, the under-surface with a short, fleshy, apiculate keel. Column stout, semi-terete, about 5 mm. long, produced at the base into a short, stout foot. Panama. СОСТЕ: region north of El Valle de Antón, trail to Las Minas, 1000 m., Allen 2868; mossy forest on the crest of Cerro Pajita, hills north of El Valle, 1200 m., Allen 9 Fair- child 3046, Allen 8 Allen 3792. 34. MAXILLARIA RINGENS Rchb. f. in Walp. Ann. 6:525. 1863. Maxillaria Tuerckbeimii Schltr. in Fedde Rep. Sp. Nov. 10:295. 1912. Maxillaria Rousseauae Schltr. in Beih. Bot. Centralbl. 36?:413. 1918. Maxillaria pubilabia Schltr. in Fedde Rep. Sp. Nov. Beih. 17:71. 1922. Maxillaria lactea Schltr. loc. cit. 233. 1923. Erect, epiphytic herbs with fleshy, approximate, elliptic-ovoid, compressed, monophyllous pseudobulbs 2.5-5.5 cm. long and 2-3 cm. wide, the bases en- veloped in several papery, imbricating bracts which become more or less fibrous with age. Leaves coriaceous, ligular, obtuse to acute, 15-40 cm. long and 2-4.5 cm. wide, contracted below into short or elongate, conduplicate petioles. Іп- florescences 1 to about 6 slender, 1-flowered scapes 5—12 cm. long, produced singly or in loose fascicles from the bases of the pseudobulbs. Flowers relatively large. Sepals free, subequal, spreading, yellow, white, or tan, sometimes shaded pink or lavender, ligular, obtuse to lanceolate, acuminate, 2.5-4 cm. long and .3-.6 cm. wide, the laterals adnate to the column foot, forming a short, rounded or subacute mentum. Petals subequal to the dorsal sepal, white or pale yellow, lanceolate, acute to acuminate, 2.2-3.5 cm. long and .25-.4 cm. wide. Lip 3-lobed near the apex, 10-15 mm. long and 6-8 mm. wide when spread out, white marked with lavender to yellow marked maroon, contracted at the base and articulated with the column foot, the lateral lobes erect, the obtuse to acute apices projecting, the mid- lobe rhombic-ovate to obovate, obtuse to acute, fleshy and more or less rugose or minutely papillose on the upper surface, about 15 the total length of the lip, the under-surface of the apex transversely thickened or with an acute central keel; the disk with a ligular callus about % the length of the lateral lobes. Column semi-terete, somewhat arcuate, about equaling the lateral lobes of the lip, the base produced into a short foot. Mexico, British Honduras, Guatemala, Honduras, Nicaragua, Costa Rica, Panama to Peru. cocLÉ: region north of El Valle de Antón, 800-1000 m., Allen 2264, 2871, 2873; (463) [Vor. 36 128 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN mountains beyond La Pintada, 400—600 m., Hunter & Allen 560. CHIRIQUÍ: without definite locality, 4000 ft., Powell 3443, 3466.. A common, very variable species, widely distributed throughout the inter- mediate highlands of central and northern South America. 35. MAXILLARIA RUFESCENS Lindl. in Bot. Reg. 21: sub /. 1802. 1836. Maxillaria Abelei Schltr. in Fedde Rep. Sp. Nov. Beih. 9:101. 1921. Erect, epiphytic herbs with approximate, ovoid-elliptic to linear, compressed, fleshy, monophyllous pseudobulbs 2.5-4.5 cm. long and 6—22 mm. wide, the bases enveloped in several papery, imbricating bracts which become fibrous with age. Leaves coriaceous, ligular, acute or rarely acuminate, 15-32 cm. long and 1.2-3.5 cm. wide, contracted below into short or elongate, conduplicate petioles. Іп- florescences slender, usually solitary, 1-flowered scapes 3.5—4 cm. long, produced from the bases of the pseudobulbs. Flowers of moderate size. Sepals subequal, free, spreading, creamy white or yellow, elliptic-oblong, obtuse to subacute, 1.4-2 cm. long and .5—7 cm. wide, the laterals adnate to the foot of the column, form- ing a short, rounded mentum. Petals subequal to the dorsal sepal, white or yellow, elliptic-lanceolate, obtuse to shortly acute, 12-15 mm. long and 5—7 mm. wide. Lip conspicuously 3-lobed, orange to yellow with reddish brown or purple mark- ings, 10-12 mm. long and 5-6 mm. wide when spread out, the base contracted and articulated with the column foot, the lateral lobes erect in natural position, the acute apices conspicuously projecting, the mid-lobe broadly oblong, about 34 the total length of the lip, the truncate apex usually emarginate; the disk with a fleshy, ligular callus equaling or exceeding the lateral lobes in length. Column stout, somewhat arcuate, semi-terete, 10-12 mm. long, the base produced into a short foot. Guatemala, Honduras, Nicaragua, Costa Rica, Panama, British Guiana, Peru, Brazil, Cuba, Hispaniola, Jamaica, Trinidad, and probably other adjacent areas. COCLÉ: mountains beyond La Pintada, 400—600 m., Hunter & Allen 563; vic. El Valle de Antén, 600-1000 m., Allen 1255. 36. MaxILLARIA UMBRATILIs L. Wms. in Ann. Mo. Bot. Gard. 28:425. 1941. Camaridium nutantiflorum Schltr. in Beih. Bot. Centralbl. 362:417. 1918, поп Maxillaria nutantiflora Schltr. Erect or pendulous, epiphytic herbs with elongate, sometimes branching stems. Pseudobulbs elliptic-ovoid, ancipitous, monophyllous, 2.5—4 cm. long and 1.5-2 cm. wide, very distantly and obliquely inserted on the canes, the lower half covered by the conduplicate bases of the 3—5 long-foliaceous bracts; the long internodes naked or enveloped in the persistent bases of the imbricating bracts. Leaves of the apex of the pseudobulb and those of the basal long-foliaceous bracts sub- coriaceous, ligular, acute or obtuse, 8—30 cm. long and 1.2-2.2 cm. wide. In- florescences slender, 1-flowered scapes 3—5 cm. long, produced from the bract axils of the elongate, complanate flush of new growth. Flowers of moderate size. Sepa!s (464) 1949] FLORA OF PANAMA (Orchidaceae) 129 free, subequal, spreading, yellowish green, lanceolate, acute, 2—2.5 cm. long and 4—6 mm. wide, the laterals adnate to the foot of the column, forming a short, rounded mentum. Petals subequal to the dorsal sepal and similarly colored, ob- liquely lanceolate, acute to acuminate, 15—18 mm. long and 3.5—5 mm. wide. Lip conspicuously 3-lobed, red or marked with red, 10—12 mm. long and 10—12 mm. wide when spread out, the base contracted and articulated with the foot of the column, the lateral lobes subfalcate, acute, erect in natural position and somewhat incurving over the column, the apices projecting, the mid-lobe ovate, acute, about И; the total length of the lip; disk with a short, broad, truncate, fleshy callus about 14 the length of the lateral lobes. Column very short, stout, about 5 mm. long, produced at the base into a short foot. Costa Rica and Panama. HIRIQUÍ: vic. Bajo Chorro, headwaters of the Río Caldera, in rain forest, 6000 ft., тон 308. 37. MAXILLARIA UNCATA Lindl. in Bot. Reg. 23: sub /. 1080. 1837. Maxillaria Macleei Batem. ex Lindl. in Bot. Reg. 26: Misc. 70. 1840. Maxillaria stenostele Schltr. in Beih. Bot. Centralbl. 367:414. 1918. Slender, erect or pendulous, epiphytic herbs with short to elongate, often branching stems 4—40 cm. long. Pseudobulbs minute, linear-complanate to terete, monophyllous, 4-10 mm. long and 2-3.5 mm. wide, approximate to rather remote, obliquely inserted on the stems, pseudobulbs and internodes enveloped in broad, brown, imbricating, scarious sheaths. Leaves fleshy, subulate-conduplicate, semi- terete or rarely lanceolate-elliptic, acute to acuminate, 2-6 cm. long and .2-1.0 cm. wide. Inflorescences short, usually solitary, 1-flowered scapes 10-20 mm. long, produced from the base of the current mature growth. Flowers small, trans- lucent with red or purple stripes. Sepals free, not spreading, the dorsal sepal ob- long, obtuse to acute, concave, 7-9 mm. long and about 3 mm. wide, the laterals lanceolate, acute, 10-12 mm. long and 3-4 mm. wide, the very long, oblique bases adnate to the foot of the column, forming an elongate, conspicuous, obtuse to subacute mentum. Petals ligular to oblanceolate, acute, about equaling the dorsal sepal in length. Lip linear-oblanceolate, obtuse to subacute, entire, somewhat ex- ceeding the lateral sepals, 12-14 mm. long and 3—3.5 mm. wide at the apex, the long narrow base slightly concave and articulated with the foot of the column, the subrotund to subacute apex often with undulate margins; the disk with a linear, obtuse callus about № the total length of the lip. Column elongate, semi- terete, somewhat arcuate, the apex with short, lateral, subfalcate, deflexed wings; the base produced into an elongate foot. British Honduras, Guatemala, Honduras, Costa Rica, Panama, Peru, British Guiana, Brazil, and probably adjacent areas. CANAL ZONE: Río Pequení, upper Madden Lake region, 66-70 m., Dodge, Steyer- mark 8 Allen 16752. PANAMÁ: hills east of Panama City, sea level to about 200 ft., Powell 120. corów: Quebrada López, lower forested slopes of Cerro Santa Rita, about 30 m., Allen 2141. состЕ: hills north of El Valle de Antón, 600-1000 m., Allen 1682, 3715. BOCAS DEL TORO: without definite locality, von Wedel 480. (465) [Vor. 36 130 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN 38. MAXILLARIA VAGANS Ames & Schweinf. in Sched. Orch. 8:65. 1925. Erect or pendulous, epiphytic herbs with stout, simple or branching stems 15—45 cm. long, the lower portions usually conspicuously enveloped in numerous fibrous roots. Pseudobulbs fleshy, monophyllous, elliptic-oblong, compressed and rugose, 1.5-3 cm. long and 1.0-1.5 cm. wide, approximate or distant, inserted at an acute angle on the stems, the current pseudobulb partly covered by the condup- licate bases of several long-foliaceous bracts, the internodes and old pseudobulbs entirely invested by the persistent bract bases. Leaves of the apex of the pseudo- bulbs and the blades of the foliaceous bracts coriaceous, ligular, obtuse, or the apex retuse and unequally bilobed, 4.5-18 cm. long and 1-2.5 cm. wide. In- florescences slender, 1-flowered scapes 3-4 cm. long, usually produced from the base of the current mature pseudobulb but sometimes from the axils of the flush of new growth. Flowers of moderate size. Sepals free, spreading, red or pink with yellow or green margins, the dorsal sepal linear-lanceolate to elliptic-lanceolate, acute, arching over the column, 1.7-2.3 cm. long and .5—7 cm. wide, the dorsal surface with a keel which terminates in an apicule, the lateral sepals spreading, elliptic-lanceolate, acute, 1.8—2.2 cm. long and .5—7 cm. wide, adnate at the base to the column foot, forming a short, rounded mentum. Petals ligular to oblanceo- late, acute, similarly colored with the sepals, 1.5-2 cm. long and .5—.6 cm. wide. Lip conspicuously 3-lobed, red margined white, or yellow with the apical lobe spotted rose-pink, 12-16 mm. long and 6-8 mm. wide, contracted at the base and articulated with the foot of the column, the lateral lobes rounded, erect in natural position, the mid-lobe fleshy, obovate, acute, canaliculate, with ascending margins, more than half the total length of the lip, the lower surface with a strongly developed keel; disk with a short, fleshy, emarginate callus. Column semi-terete, somewhat arcuate, 6—8 mm. long, produced at the base into a short foot. Anther terminal, operculate, incumbent, 1-celled Costa Rica and Panama. CLE: mountains beyond La Pintada, 400—600 m., Hunter & Allen 598; region north of El Valle de Antón, trail to Las Minas, 1000 m., Alle en 2803; mossy forest on crest о Cerro Pajita, hills north of El Valle, 1000 m., Allen 4177. iE Palo Alto, 4500— 5000 ft., Powell 348. A fairly frequent highland species closely allied to Maxillaria cucullata. 39. MAXILLARIA VALENZUELANA (A. Rich.) Nash, in Bull. Torr. Bot. Club. 34:121. 1907. Dicrypta iridifolia Batem. in Loud. Hort. Brit. Su ppl. 2:630. 1839, nomen nud E valenzuelana A. Rich. in Sagra, Hist. Isla Cubs 11 [Fl. Cub. Po 234. 18 Ий iridifolia (Batem.) Rchb. f. їп Bonplandia 2:16. 1854. Dicrypta irisphyta Barb. Rodr. Gen. & Spec. Orch. 1:126. 1877. Pendulous epiphytic herbs without pseudobulbs. Leaves distichously arranged in the form of a broad fan, rather fleshy, gladiate, lanceolate, acuminate, 9—30 cm. long and 1.0—2.5 cm. wide, the broad conduplicate bases closely imbricating. In- (466) 1949] * FLORA OF PANAMA (Orchidaceae) 131 florescences short, 1-flowered scapes, solitary or in dense fascicles in the axils of the leaves. Flowers of moderate size. Sepals free, subequal, spreading, yellow, elliptic- lanceolate, acute, 10—12 mm. long and 4—6 mm. wide, the laterals somewhat ob- lique, the bases adnate to the column foot, forming ап inconspicuous, rounded mentum. Petals lanceolate, acute, yellow, 8-10 mm. long and 2.5—3.5 mm. wide. Lip entire or obscurely 3-lobed, yellow with purple or maroon markings, elliptic- ovate, acute, 9-11 mm. long and 4—5 mm. wide when spread out, the base con- tracted and articulated with the column foot, the lateral margins erect in natural position, the mid-lobe thickened at the apex and on the margins, about 77 the total length of the lip; disk with a ligular callus running the full length of the lip, some- what thickened near the apices of lateral margins. Column short, stout, somewhat arcuate, produced at the base into a short foot. Cuba, Honduras, Costa Rica, Panama, Colombia, Venezuela, Brazil, and prob- ably adjacent areas. CHIRIQUÍ: vic. Boquete, 3800 ft., Davidson 636. A frequent, pendulous, fan-leaved epiphyte of the Chiriquí highlands. 40. MAXILLARIA VARIABILIS Batem. ех Lindl. in Bot. Reg. 23: sub 7. 1086. 1837. Maxillaria Hencbmanni Hook. in Bot. Mag. /. 364. 2 ; 8. п. Maxillaria revoluta Kl. in Allgem. Сагон 20: 186. Maxillaria cbiriquensis Schltr. іп Fedde Rep. Sp. Nov. ва, a 68. 1922. Maxillaria panamensis Schltr. loc. cit. 70. 1922. Maxillaria costaricensis Schltr. loc. cit. 19:232. 1923. Erect or pendulous, epiphytic herbs with short or elongate, usually simple stems of very variable vegetative appearance, 5—30 cm. long. Pseudobulbs linear to elliptic-oblong, subcylindric to strongly ancipitous, usually rugose, 1.5—6 cm. long and .3-2.0 cm. wide, approximate and inserted at an acute angle on the stems, the truncate apices monophyllous, the internodes and bases enveloped in closely im- bricating, papery bracts. Leaves coriaceous, ligular, obtuse or 2-lobed at the apex, 3-15 cm. long and .3-1.6 cm. wide, contracted below into very short, condupli- cate petioles. Inflorescences slender, 1-flowered, often solitary scapes produced from the bract axils of the flush of new growth, or sometimes from the base of the current mature pseudobulb. Flowers usually of moderate size or small, white or yellow marked with dark red, or with a red lip, or entirely dark red. Sepals sub- equal, free, more or less spreading, linear-lanceolate to elliptic-lanceolate, obtuse, with a minute apicule or sometimes acute, 6—12 mm. long and 2-4 mm. wide, the laterals adnate to the foot of the column, forming a short, subacute mentum. Petals lanceolate to oblanceolate, acute, 5-10 mm. long and 1.5—3.5 mm. wide. Lip entire or obscurely 3-lobed near the apex, 6-12 mm. long and 3-6 mm. wide when spread out, the base articulated with the column foot, the lateral margins rounded and erect in natural position, the mid-lobe subacute to truncate and (467) [Vor. 36 132 ANNALS OF THE MISSOURI BOTANICAL GARDEN shallowly emarginate, about !4 the total length of the lip; the disk with a ligular, obtuse callus nearly equalling the lateral lobes. Anther terminal, operculate, in- cumbent, 1-celled. Mexico, British Honduras, Guatemala, Salvador, Honduras, Nicaragua, Costa Rica, Panama, British Guiana, and probably adjacent territories. PANAMÁ: hills near Panama City, sea level, Powell 124. состЕ: vic. El Valle de Antón, 600-1000 m., Allen 1252, 3716, 3813. снікюоі: without definite locality, 4000 ft., Powell 125; vic. Boquete, 4000-4500 ft., Powell 418; Cerro de Lino, vic. Boquete, 1000-1300 m., Pittier 3017; vic. Boquete, 3800 ft., Davidson 645; vic. Monte Lirio, upper Río Chiriqui Viejo, б. White 38; vic. New Switzerland, Rio Chiriquí Viejo, 1800—2000 m., Allen 1339. 41. MAXILLARIA WERCKLEI (Schltr.) L. Wms. in Ann. Mo. Bot. Gard. 27:284. 1940. А Ornithidium Wercklei Schltr. іп Fedde Rep. Sp. Nov. Beih. 19:60. 1923. Maxillaria aurantiaca Schltr. loc. cit. 27:87. 1924. Maxillaria Lankesteri Ames, in Sched. Orch. 7:11. 1924. Erect or pendulous, epiphytic herbs with slender, branching stems 3—20 cm. tall. Pseudobulbs linear, compressed, rugose in dried specimens, 4—25 mm. long and 1.5-5 mm. wide, distantly inserted on the stems, the truncate apices mono- phyllous, the bases with 2-3 conspicuous, long-foliaceous bracts, the long inter- nodes enveloped in persistent, closely imbricating, complanate sheaths. Leaves of the apex of the pseudobulbs and the blades of the foliaceous bracts coriaceous, ligular to elliptic-lanceolate, broadly obtuse, or retuse at the apex, 1—3 cm. long and .2—1.2 cm. wide, contracted below into very short, subsessile, conduplicate petioles. Inflorescences slender, 1-flowered scapes 4-10 mm. long, produced from the bract axils of the flush of new growth. Flowers very variable, small or of moderate size, usually tan striped red or brown to dark reddish purple. Sepals sub- equal, free, spreading, lanceolate to elliptic-lanceolate, acute, 5-12 mm. long and 2.5-5 mm. wide, the apices sometimes carinate, the laterals adnate to the foot of the column, forming a short, rounded mentum. Petals subequal to the dorsal sepal, oblong-lanceolate, obtuse to subacute, 4-11 mm. long and 2-4 mm. wide. Lip 3-lobed near the base, 4-10 mm. long and 3—6 mm. wide, contracted at the base and articulated with the column foot, the short, lateral lobes rounded or subacute, obliquely erect in natural position, separated near the apices by a transverse, fleshy callus, mid-lobe elliptic-ovate to oblong, convex, the margins often deflexed or revolute, the apex obtuse to subacute, 34 to 3⁄4 the total length of the lip. Column short, stout, semi-terete, somewhat arcuate, produced at the base into a distinct oot. Anther terminal, operculate, incumbent, 1-celled. Costa Rica, Panama, and Colombia. СОСТЕ: hills north of El Valle de Antón, 1000 m., Allen 1253. (To be concluded in Part Ш, Fasc. 5) (468) OF THE MISSOURI BOTANICAL GARDEN Director СЕОБСЕ T. Moore HERMANN VON SCHRENK, Pathologist JEssE М. GREENMAN, e Emeritus o£ the Herbarium Саввотт. W. Dopcz, Mycologist EDGAR ANDERSON, —- Geneticist Ковевт E. Woopson, Ja. gee of the Herbarium ‘Henry М. ANDREWS, Paleobotanist Rosert W. SCHERY, Gustav А, L. MEHLQUIST, | — воша M. TRYON Assistant Curator of the Herbarium жс Ku шымы. 27 о а a ono Кои Ç ae FLORA ОҒ РАМАМА Part ІП. Fascicle 4 ORCHIDACEAE Ву PAUL H. ALLEN 66. TRIGONIDIUM Lindl. TRIGONIDIUM Lindl. in Bot. Reg. 23: 4. 1023. 1837; Benth. & Hook. Gen. Р], 3:567. 1883. Caespitose, erect, repent, ог scandent epiphytic herbs with short ovoid or elliptic-ovoid, compressed, usually ridged and sulcate pseudobulbs, the apices with 1-5 linear to elliptic-lanceolate, coriaceous or subcoriaceous leaves. Inflorescences simple, slender, short or elongate, 1-flowered scapes produced from the bases of the pseudobulbs. Flowers relatively large. Sepals subequal, the bases connivent, forming a tube, the apical portions spreading or reflexed. Petals much smaller than the sepals. Lip distinctly 3-lobed, very small, shorter than the petals, the lateral lobes erect and paralleling the column, the mid-lobe spreading, often thickened or tuberculate, the apex usually recurved; disk with a fleshy, usually ligular callus. Column short, rather stout, semiterete, wingless, the base without a foot. Anther terminal, operculate, incumbent, 1-celled; pollinia 4, waxy. A small genus of tropical American epiphytes, ranging from Mexico to Brazil. Two species are known from Panama. a. eee сы ба: аке the арех with 2 leaves. Petals пос apicu- Lowland sp l. T. EGERTONIANUM aa. БЕЗІ ub кра separated оп а cylindric rhizome, the арех with 3—5 leaves. Petals E yan ы apiculate. ‘Highland species................ 2. T. LANKESTERI 1. TRIGONIDIUM EGERTONIANUM Batem. ex Lindl. in Bot. Reg. n. s. 1: Misc. 73. 1838. Trigonidium Seemanni Rchb. f. in Seem. Bot. Voy. Herald, p. 214. 1854. Erect epiphytic herbs with approximate, ovoid to elliptic-ovoid, ridged and sulcate, diphyllous pseudobulbs 3.5-7 cm. long and 1.5-4 cm. wide, the bases en- veloped in 3—5 brown, imbricating bracts which become fibrous with age. Leaves subcoriaceous, linear-lanceolate, acute, 20-60 cm. long and .8-3 cm. wide, con- tracted below into slender, conduplicate petioles. Inflorescences 1 to several Issued June 18, 1949. (133) (469) ANNALS ОҒ THE Fig. 186. MISSOURI BOTANICAL GARDEN Trigonidium Egertonianum (470) [Vor. 36 1949] FLORA OF PANAMA (Orchidaceae) 135 slender, erect, 1-flowered scapes, 20—40 cm. long, produced from the bases of the pseudobulbs, enveloped throughout in tubular, papery, imbricating bracts. Flowers relatively large. Sepals free, the bases connivent, forming a tube, usually greenish yellow to pinkish tan, with brown or purple stripes; the dorsal sepal elliptic- oblanceolate, acute, spreading, 2.5—4.5 cm. long and 1—2 cm. wide, the laterals obliquely oblong to elliptic-lanceolate, acute, the apices strongly recurved, 27—42 mm. long and 8-14 mm. wide. Petals lanceolate to oblanceolate, acute, greenish yellow to pinkish tan, striped brown or purple, 12—22 mm. long and 4—6 mm. wide, with a brown to purple gland-like thickening near the apex. Lip 3-lobed, about № the length of the petals, yellowish tan striped brown or red, 5-6 mm. long and 3—4 mm. wide when spread out, contracted at the base and articulated with the base of the column, the lateral lobes erect and paralleling the column, the mid-lobe ovate, acute, pale yellow with a darker center, rather fleshy, № to 25 the total length of the lip, the apex recurved and verrucose on both the upper and lower surfaces, the disk with a fleshy ligular callus about equaling the lateral lobes in length. Column semiterete, purple, 4—5 mm. long, the base footless. Mexico, British Honduras, Guatemala, Salvador, Honduras, Nicaragua, Costa Rica, Panama, and Colombia. CANAL ZONE: vicinity Frijoles, sea level, Powell 3023; drowned forest of the upper Rio Pequení, Madden Lake region, 70 m., ‘Alles 17281; forest along banks above Rio Indio Hydrographic Station, Madden Lake region, 70-75 m., Steyermark 17426; vicinity Gamboa, 20-100 m., Pittier 2613; Barro Colorado Island, Gatun Lake, Zetek s.n., Wood- worth & Vestal 503. PANAMA: along road to Pacora, about 50 m., Allen 820; hills east of Panama City, Powell 44. согом: Cerro Santa Rita, 1200 ft., Allen 9 Fairchild 5194; Rio Gatuncillo, vicinity Santa Rosa, 100 ft., Allen 9 Allen 4159. veERAGUAS: Bahia Honda, Taylor 1515; Elmore Н-48; vicinity ‘Soné, about 100 ft., Allen & Allen 4240. BOCAS DEL TORO: vicinity Chiriqui Lagoon, von Wedel 2006. A very common lowland species widely distributed in Central America. It seems probable that it may represent a northern variant of the generic type, Trigonidium obtusum Lindl. of British Guiana and Brazil, differing only in the narrower leaves and the absence of an apparently distinct verrucose callus on the under-surface of the apex of the lip. 2. TRIGONIDIUM LANKESTERI Ames, in Sched. Orch. 5:32. 1923. Repent or scandent, epiphytic herbs with ovoid, compressed, ridged and sulcate, tapering pseudobulbs 4—8 cm. tall and 2—3 cm. wide at the base, very distantly spaced on the elongate, cylindric rhizome, the internodes 15—40 cm. long, en- veloped in numerous tubular, papery bracts which weather away, the basal portion of the pseudobulbs freely rooting and covered by several rigid, imbricating sheaths which soon weather into loose fibers, the apex of the pseudobulbs with 3—5 leaves. Leaves subcoriaceous, oblong-lanceolate, acute, 15—27 cm. long and 3—4 cm. wide, contracted below into slender, conduplicate petioles which are more or less dis- tichously arranged on the apex of the pseudobulbs. Inflorescences short, usually solitary, 1-flowered scapes 8—16 cm. long, produced from the bases of the pseudo- bulbs. Sepals subequal, free, the basal half, together with the petals, connivent (471) [Vor. 36 136 ANNALS OF THE MISSOURI BOTANICAL GARDEN and forming a tube, the petals visible between the margins of the dorsal and lateral sepals, the apical half of the sepals strongly reflexed, light greenish tan to cinna- mon, veined brown or purplish, elliptic-oblanceolate, acute, 4.5-5 cm. long and 1.6-2 cm. wide, the dorsal sepal a little narrower than the laterals. Petals broadly oblanceolate, the apices abruptly spreading and conspicuously apiculate or aristate, light greenish tan, veined brown, with purple spots, 20-24 mm. long and 8-10 mm. wide, with an irregular subventricose swelling on the inside below the apicule. Lip 3-lobed, white spotted brown, about 15 mm. long and 5 mm. wide, oblanceo- late when spread out, the base contracted and articulated with the base of the column, the lateral lobes erect in natural position, the triangular acute apices pro- jecting, the mid-lobe ovate, obtuse, fleshy, strongly recurved, about 1⁄2 the total length of the lip, the upper-surface minutely glandular, the under-surface strongly verrucose, the disk with a ligular, fleshy callus about equaling the lateral lobes in length. Column semiterete, somewhat arcuate, about 1 cm. long, the base without a foot. Costa Rica and Panama. СОСТЕ: hills north of El Valle de Antón, 800—1000 m., Allen 2290. An exteremely distinctive highland species, immediately separable from the common lowland Trigonidium Egertonianum by the elongate rhizome, the 3- to 5-leaved pseudobulbs, the larger flowers, and the apiculate petals. 67. CRYPTOCENTRUM Benth. CRYPTOCENTRUM Benth. in Jour. Linn. Soc. 18:325. 1881, nom. nud.; Benth. & Hook. Gen. Pl. 3:557. 1883. Pittierella Schltr. in Fedde Rep. Sp. Nov. 3:80. 1906. Small erect, epiphytic herbs with 1 to several short stems, usually without pseudobulbs, the cylindric basal portions of the stems enveloped in the rigid, per- sistent leaf bases. Leaves approximate, 2-ranked, linear to ligular, coriaceous, usually confined to the upper portions of the stems. Inflorescences few to num- erous, slender, erect or arching, 1-flowered scapes produced from the bases of the plants, usually less than (but sometimes exceeding) the leaves, the peduncles en- veloped in tubular, acute or acuminate, papery sheaths, the uppermost of which is much the longest, completely covering the ovary, only the perianth segments of the flowers being exserted. Flowers relatively small. Sepals narrow, the free portions widely spreading, the dorsal sepal usually a little shorter than the laterals, the bases of which are connate and produced into a long, narrow spur which is appressed to the ovary and nearly equaling it in length. Petals subequal to the dorsal sepal or smaller. Lip continuous with the base of the column, produced into an elongate tubular projection contained within, and nearly equaling the sepaline spur, the free limb entire or more or less divided into a basal and an ap- ical half, the basal half concave to subsaccate, the erect lateral margins enclosing (472) 1949] FLORA OF PANAMA (Orchidaceae) 137 the column, the apical half usually lanceolate or elliptic-lanceolate and spreading. Column short, very stout, wingless but with the apex usually with a pair of lateral auricles covering the clinandrium, the base without a foot. Anther terminal, operculate, incumbent, 1-celled; pollinia 2—4, waxy. A small genus of tropical American highland epiphytes, ranging from Costa Rica to Peru. Until more adequate collections are available for study, it would be profitless to speculate on the possible number of valid entities. Although the described species vary considerably in size, the flowers of most seem to be remark- ably constant in structure, so that many of the names may eventually be reduced to synonymy or some perhaps retained as local varieties of a few somewhat poly- morphic species. Four entities are separable in Panama, largely on a basis of size and vegetative habit. a. Plants less than 3 cm. tall. Inflorescences conspicuously exceeding the eaves 4. C. STANDLEYI aa. Plants more = 6 cm. tall. Inflorescences less than, ог barely equal- b bans € 6-12 cm. tall. Leaves 3.5 mm. wide or less. Spur 1.2-2.0 1. C. GRACILIPES bb. Pina more than 12 cm. tall. Leaves 3.5 mm. wide or more. Spur ME 3 cm. long. ves narrowly linear, acuminat 2. Са “gee pars am Co: ees broadly ligular, п emarginate 3. C. LATIF — . CRYPTOCENTRUM GRACILIPES Schltr. іп Fedde Кер. Sp. Nov. Beih. 19:246. 12925. Erect, usually caespitose, epiphytic herbs without pseudobulbs, the plants 6—12 cm. tall. Leaves coriaceous, spreading, approximate, linear-ligular, the apices obtuse or unequally emarginate, 5-10 cm. long and 0.2-0.5 cm. wide, the sheathing bases contracted into short cylindric petioles. Inflorescences numerous slender, erect or arching, 1-flowered scapes usually nearly equaling the leaves in length, enveloped throughout in several papery, tubular, imbricating sheaths. Flowers small, tan or brownish olive. Sepals oblong, obtuse, widely spreading, the free portions 10-13 mm. long and 2.5—3 mm. wide, the laterals oblique and rather narrower than the dorsal sepal, the bases connate and produced into an elongate spur 1.2-2 cm. long, parallel with the ovary and about equaling it in length. Petals obliquely elliptic-lanceolate, acute, 8-10 mm. long and 2—2.5 mm. wide. Lip entire, the free limb lanceolate, obtuse to subacute, 6—8 mm. long and 2.5—3 mm. wide, the base continuous with the base of the column and produced into a long narrow tube which is contained within the elongate, sepaline spur. Column short, stout, about 2 mm. long, the base footless. Costa Rica and Panama. СОСТЕ: hills north of El Valle de Antón, 1000 m., Allen 1011. N CRYPTOCENTRUM INAEQUISEPALUM C. Schweinf. in Bot. Mus. Leafl. Harv. Univ. 12:186. 1946. Erect, epiphytic herbs without pseudobulbs, the plants 15—25 cm. tall, the short, cylindric stems enveloped below in the persistent leaf bases, the apical por- (473) (Vor. 36 138 ANNALS OF THE MISSOURI BOTANICAL GARDEN tions with 2-ranked foliage. Leaves approximate, linear, acuminate, coriaceous, 12-23 cm. long and 0.35—0.6 ст. wide. Inflorescences numerous slender, 1-flowered scapes produced from the base of the stem, the peduncles entirely invested by sev- eral papery, tubular sheaths. Flowers relatively large for the genus, described as being greenish yellow and translucent. Sepals widely spreading above, lanceolate, acute, the dorsal sepal about 15 mm. long and 3.7 mm. wide, the lateral sepals oblique, the free parts about 19 mm. long and 3 mm. wide, the bases connate and produced into an elongate spur which is closely appressed to and about equaling the ovary in length. Petals obliquely lanceolate, acuminate, about 13 mm. long and 3 mm. wide. Lip continuous with the base of the column, the free limb ovate-lanceolate, acute to acuminate when spread out, about 10 mm. long and 4.2 mm. wide, the lower half deeply concave to subsaccate in natural position, the lateral margins enclosing the column, the base produced into a long slender tube contained within the sepaline spur. Column very short, stout, about 2 mm. long, the clinandrium shielded by 2 broad lateral auricles, the base without a foot. Panama and Peru. То be expected in Colombia and Ecuador. CHIRIQUÍ: vicinity Bajo Chorro, headwaters of the Río Caldera, in rain forest, 6000 ft., Davidson 160. Our specimens have thicker leaves, but in other respects seem to agree with the Peruvian type. 3. CRYPTOCENTRUM LATIFOLIUM Schltr. in Fedde Rep. Sp. Nov. Beih. 19:247. 1922. Erect, epiphytic herbs without pseudobulbs, the plants 18-25 cm. tall, the short cylindric stems often solitary and apparently monopodial, enveloped below in the rigid, brown, persistent, imbricating leaf bases. Leaves approximate, spread- ing, broadly ligular, coriaceous, 16-23 cm. long and 1.5—2 cm. wide, the apices rounded and unequally emarginate. Inflorescences a few erect or arching, 1- flowered scapes 8—16 cm. long, produced from the bases of the stems, the peduncles completely invested by several acute, tubular, papery sheaths. Flowers relatively large, rather fleshy, greenish olive or greenish tan. Sepals widely spreading above, the free portions narrowly ligular-lanceolate, acute, 3.5-5 cm. long and 0.5-0.6 ст. wide, the margins often strongly revolute, the lateral sepals oblique, the bases connate and produced into an elongate spur which is closely appressed to the ovary and about equaling it in length. Petals obliquely lanceolate, acute, 3—4 cm. long. Lip continuous with the base of the column, the free portion lanceolate, subacute, about 1.5 cm. long and 0.5 cm. wide, the base produced into an elongate tube which is contained within the sepaline spur, as in the genus. Column short, stout, about 4 mm. long. Costa Rica and Panama. СОСІ.Е: region north of El Valle de Antón, vicinity La Mesa, 1000 m., Dunn s. n. (under Allen 2762). (474) 1949] FLORA OF PANAMA (Orchidaceae) 139 A curious highland species, vegetatively reminiscent of a small Aerides or Vanda. Our specimen has somewhat smaller measurements than those given for the Costa Rican type, but otherwise seems nearer to it than to any other. Sterile plants of evidently the same species are fairly frequent on the eastern slopes of Chiriquí Volcano between 5000 and 7000 ft 4. CRYPTOCENTRUM STANDLEYI Ames, in Sched. Orch. 9:55. 1925. Dwarf, epiphytic herbs without pseudobulbs, the tufted plants less than 5 cm. tall. Stems cylindric, rather stout, the lower half covered by the persistent, im- bricating leaf bases, the upper half with approximate, more or less 2-ranked foliage. Leaves narrowly linear to oblanceolate, obtuse to subacute, often con- duplicate, coriaceous, 5-15 mm. long and 1-2.5 mm. wide. Inflorescences usually 1—2 erect, filiform, 1-flowered scapes 3—4 cm. long, produced from the lower axils of the leaves and conspicuously exceeding the foliage in height. Flowers small, dull red or tan. Sepals spreading, the free portions elliptic-lanceolate, acute, 6—8 mm. long and 1.5-2 mm. wide, the laterals connate at the base and produced into an elongate, slender spur which is closely appressed to the ovary and about equal- ing it in length. Petals lanceolate, acute, 5-7 mm. long and 1.5-2 mm. wide. Lip continuous with the base of the column, the free portion strongly concave, lanceolate, acute, 6-8 mm. long and about 2 mm. wide, the base prolonged into a slender tube contained within the sepaline spur. Column 2.2-2.5 mm. long, the apex with two lateral auriculate projections. Costa Rica and Panama. СОСТЕ: hills north of El Valle de Antón, 600-1000 m., Allen 1685. 68. TRICHOCENTRUM Poepp. & Endl. TricHOCENTRUM Poepp. & Endl. Nov. Gen. & Sp. 2:11, £. 115. 18371; Benth. € Hook. Gen. Р]. 3:559. 1883. Acoidium Lindl. in Bot. Reg. 23: /. 1051. 1837, nom. nud. Small, epiphytic herbs with very short, monophyllous stems which are thick- ened into short, inconspicuous pseudobulbs. Leaves elliptic-lanceolate to ligular, coriaceous or fleshy. Inflorescences usually simple, rarely branching, 1- or rarely 2-flowered scapes, usually very short, but sometimes nearly equaling the leaves. Flowers of moderate size to relatively large. Sepals subequal, free, spreading. Petals subequal to the sepals. Lip entire or obscurely 3-lobed, produced at the base into a spur. Column stout or slender, erect, the apex clavate or with prominent lateral wings or auricles, the base without a foot. Anther terminal, operculate, incumbent, 1-celled, often pubescent or papillose; pollinia 2, waxy. Perhaps fifteen species of tropical American epiphytes, ranging from Mexico to Peru and Brazil. One species is known from Panama. l'The date of p appearing on the title page of this volume is 1838, but evidently wn of it must have been publish in some way available for use earlier since E edle ey (in Bot. Reg. 23: 7. 1951, April L 1837) p the generic name Trichocentrum and the place of publication. (475) [Vor. 36 140 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1. TRICHOCENTRUM PANAMENSE Rolfe, in Kew Bull. 341. 1913. Dwarf, epiphytic herbs with minute, monophyllous pseudobulbs, the plants 3-12 cm. tall. Leaves elliptic-lanceolate to ligular, acute, fleshy, 2.5-10 cm. long and 0.8-2.5 cm. wide. Inflorescences slender, horizontal or pendulous, usually un- branched scapes 0.8—4 cm. long, which apparently elongate by progressive stages, producing a succession of moderate-sized solitary flowers. Sepals subequal, free, spreading, pale green or greenish yellow, lanceolate, acute, 10—12 mm. long and 3-4 mm. wide. Petals pale green or greenish-yellow, oblong-lanceolate, acute, 10—12 mm. long and 3—4 mm. wide. Lip elliptic-ovate, obtuse to acute, explanate or concave, white with a reddish-brown or reddish-purple blotch at the base, the free portion 10—12 mm. long and about 6 mm. wide, the base adnate to the base of the column and produced into a broad, flattened, rather obscurely 4-lobed spur. Column stout, about 5 mm. long, the apex with a pair of broad, fleshy, spreading, subfalcate, obtuse wings on either side of the clinandrium, the base without a foot. Anther white, papillose. Panama. CANAL ZONE: trail from Fort Sherman to the mouth of the Rio Chagres, sea level, Powell 371; upper | Rio Chagres, sea level, Powell 410. PANAMA: Cerro Campana, vicinity Campana, 600-800 m., Dorothy Allen 3989, 4455. lower valley and marshes along the Río pei 500—600 m., Hunter & Allen йу pem Allen 3953, 5077. This species is very variable in regard to the size and shape of the leaves, and the length of the inflorescences. 69. IONOPSIS НВК. Ionorsis НВК. Nov. Gen. & Sp. 1:348, Ё. 83. 1815; Benth. & Hook. Gen. РІ. 31302. 1859. Inopsis Steud. Nom. ed. 1, 432. 1821. lantba Hook. Exot. Fl. 2: £. 113. 1825. Cybelion Spreng. Syst. 3:679. 1826. Epiphytic herbs with short, subcylindric stems invested in 2-ranked foliage, the sheathing leaf bases with or without minute pseudobulbs. Leaves approximate, narrow, coriaceous, the bases imbricating. Inflorescences 1-3 slender, short to elongate, erect or arching, simple or paniculate scapes subterminal or lateral from the axils of the leaves. Flowers of moderate size to small. Sepals subequal, erect or spreading, the dorsal sepal free, the laterals connate at the base and produced into a short sac below the lip. Petals subequal to the dorsal sepal or broader. Lip clawed and adnate to the base of the column, the apical portion broadly expanded, entire or emarginate, conspicuously exceeding the sepals. Column short, stout, erect, the lateral margins not winged, the base without a foot. Anther terminal, operculate, incumbent, 1-celled or imperfectly 2-celled; pollinia 2, waxy. A small genus of American epiphytes, ranging from Florida and Mexico through Central America and the West Indies to South America, as far as Bolivia and Paraguay. One species is known from Panama. (476) 1949] 141 FLORA OF PANAMA (Orchidaceae) 4 A 3 x - _ b js ; » ж $ "m C Z » ` P T 23 dpt Q < №379 e d WS 3⁄4 4 үу 3 s. 2 5 A As ах š lonopsis utricularioides Fig. 187. (477) [Vor. 36 142 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1. IONOPSIS UTRICULARIOIDES (Sw.) Lindl. Coll. Bot. 7. 30-А. 1821. Epidendrum utricularioides Sw. Prodr. 122. 1788. Dendrobium utricularioides Sw. in Nov. Act. Soc. Upsal. 6:83. 1799, lantba рој n Exot. Fl. i раху 4. lonopsis tene Lindl. n Bot. Re eg. 22: f. 1004. 1836. жеде о Lindl. loc. cit. ^ t. 1004. 1836. lonopsis paniculata Lindl. loc. cit. 1836. Ionopsis zonalis Lindl. & Paxton, in Paxton’s Flow. Gard. 2:13. 1851-53, in textu. Epiphytic herbs 5-12 cm. tall. Leaves approximate, linear-lanceolate to ob- long-lanceolate, acute, the blades 3.5—12 cm. long and 0.5-1.5 cm. wide, the mid- vein forming a keel on the under-surface, the sheathing bases sometimes enveloping a minute pseudobulb. Inflorescences 1-3 slender, erect or arching, simple or paniculate scapes 15—60 cm. long, produced from the lateral or subterminal leaf axils, provided with a few distant, inconspicuous sheaths. Flowers few to many, small to relatively large and conspicuous, lavender, rose-purple, or white, super- ficially resembling those of some species of Utricularia. Sepals subequal, not spreading, oblong-lanceolate to elliptic-lanceolate, acute, 3-6 mm. long and 1—2 mm. wide, the dorsal sepal free, concave, the laterals connate at the base and pro- duced into a short, broad, obtuse sac. Petals subequal to the dorsal sepal, elliptic- ovate, acute, or obtuse and shortly apiculate, 2.5-5 mm. long and 1.5-3 mm. wide. Lip about twice as long as the sepals, 5-12 mm. long and 5—12 mm. wide, the apex broadly spreading, deeply emarginate and 2-lobed, contracted at the base into a fleshy, biauriculate claw which is adnate to the base of the column, the apex of the claw with 2 suborbicular, flattened calli. Column short, rather stout, with- out wings, about 2 mm. long, the base without a foot. Mexico to Brazil and Paraguay; Florida; West Indies. CANAL ZONE: vicinity Gamboa, 100 ft., Dorothy Allen 4572. PANAMÁ: hills east of Panama City, sea level, Powell 181, А common epiphyte widely distributed throughout the lowlands of the Ameri- can tropics and sub-tropics. The species is very variable in regard to the size of flowers and width of leaves. 70. RODRIGUEZIA Ruiz & Pavon RopniGUEZIA Ruiz & Pavon, Fl. Peruv. & Chil. Prodr. 115, £. 25. 1794; Benth. & Hook. Gen. Pl. 3:559. 1883. Burlingtonia Lindl. in Bot. Reg. 23: f. 1027. 1837 Physanthera Bert. ex Steudel, Nom. ed. 2, 2:330. 1841, nom. nud. Epiphytic herbs with short, ovoid or elliptic-oblong, compressed, 1- to 2-leaved pseudobulbs distantly inserted on an elongate rhizome, or approximate, usually almost completely hidden by the conduplicate, imbricating bases of 2 to several conspicuous, foliaceous bracts. Leaves of the apex of the pseudobulbs and the blades of the foliaceous bracts coriaceous, ligular to elliptic-lanceolate, obtuse to (478) 1949] | FLORA OF PANAMA (Orchidaceae) 143 acute, ог unequally emarginate. Inflorescences 1 to several simple, erect or deflexed racemes, produced from the axils of the foliaceous bracts. Flowers few to many, of moderate size to relatively large and conspicuous. Sepals usually dissimilar, the dorsal sepal free, concave and petaloid, the laterals usually connate for their entire length, forming a single conduplicate segment below the lip, often strongly genic- ulate or gibbose when seen in profile, or rarely connate for only about half their length, with the acute apices spreading. Petals subequal to the dorsal sepal. Lip usually exceeding the sepals, the base clawed, continuous with the base of the column and produced into a short spur or gibbose projection which is appressed to the ovary, the claw parallel with the column, the free limb spreading, obovate or obcordate, the disk often cristate. Column erect, slender, subcylindric, the apex dilated or subclavate, often with variously shaped arms or auricles, the base with- out a foot. Anther terminal, operculate, incumbent, 1-celled; pollinia 2, waxy. А small genus of attractive tropical American epiphytes, ranging from Costa Rica to Peru and Brazil. Two species are known from Panama. a. Inflorescences erect, equaling or exceeding the leaves. Sepals about 1 cm. long 2. В. SECUNDA aa. Inflorescences deflexed, not equaling the leaves. Sepals about 2.5 cm. long 1. R. COMPACTA 1. RopRIGUEZIA COMPACTA Schltr. іп Fedde, Rep. Sp. Nov. Beih. 19:144. 1923. Epiphytic herbs with oblong-elliptic, compressed, monophyllous pseudobulbs 1.5-2.5 cm. long and 1.0-1.5 cm. wide, nearly hidden by the conduplicate im- bricating bases of several conspicuous, foliaceous bracts. Leaves from the apex of the pseudobulbs and the blades of the foliaceous bracts oblong-ligular to ligular, coriaceous, the apices obtuse or obscurely emarginate, 6—14 cm. long and 1.5-2.5 cm. wide. Inflorescences 1-2 lax, slender, deflexed, few-flowered racemes 3-4.5 cm. long. Flowers of moderate size, conspicuously larger than those of Rodriguezia secunda, pale yellow or greenish yellow with a yellow lip. Sepals dissimilar, the dorsal sepal narrowly oblong, obtuse, concave, about 2.5 cm. long, the laterals connate for their entire length, forming a single conduplicate segment below the lip, about 2.6 cm. long, the apex shortly bifid, the base geniculate in profile. Petals obliquely oblong, obtuse, about 2.5 cm. long. Lip adnate to the foot of the column, about 2.5 cm. long and 1 cm. wide, the free limb obovate-spatulate, the apex deeply emarginate and 2-lobed, the mid-nerve thickened and projecting be- tween the apical lobes as a short apicule, the base produced into a short, curved, subulate spur. Column subcylindric, the apex dilated and subclavate, with pro- jecting arms, the basal half minutely puberulent, the base without a foot. Costa Rica and Panama. ВОСА5 DEL TORO: Río Cricamola, between Finca St. Louis and Konkintóe, about 10—50 m., Woodson 1888. An apparently attractive species, readily separable from the common Rodri- guezia secunda by the short, deflexed inflorescences and the relatively large, greenish white or pale yellow flowers. (479) [Vor. 36 144 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN 2. RODRIGUEZIA SECUNDA НВК. Nov. Сеп. & Sp. 1:367, £. 02. 1816. Pleurotballis coccinea Hook. Exot. Fl. 2: Ё. 120. 1815. Rodriguezia lanceolata Lodd. Bot. Cab. 7: t. 676. 1822, non R. lanceolata R. ез secunda НВК. var. banamensis Schltr. іп Fedde, Кер. Sp. Хо». E 17:75. Erect, epiphytic herbs with oblong-elliptic, compressed, monophyllous pseudo- bulbs 2-3.5 cm. long and 1.0-1.6 cm. wide, almost completely covered by the conduplicate, imbricating bases of the 4—7 conspicuous, foliaceous bracts. Leaves of the apex of the pseudobulbs and the blades of the foliaceous bracts coriaceous to rather fleshy, linear-ligular to elliptic-oblong, obtuse, subacute or unequally emar- ginate at the apex, 7-24 cm. long and 1-3.5 cm. wide. Inflorescences 1-6 erect or arching, unbranched, unilateral racemes 15—38 cm. tall. Flowers many, rather small, pink to rose-red. Sepals dissimilar, not spreading, the dorsal sepal concave, ovate, obtuse with a minute apicule, or subacute, 9—12 mm. long and 5-6 mm. wide, the lateral sepals connate for their entire length, forming a single, con- duplicate segment below the lip, 10—15 mm. long, the base conspicuously gibbose or geniculate when seen in profile. Petals obovate, shortly acute or apiculate, 9-12 mm. long and 6—7 mm. wide. Lip entire, obscurely clawed, the biauriculate claw adnate to the base of the column and produced into a very short spur, the free limb oblong-obovate, with undulate margins, 12—15 mm. long and 5—7 mm. wide, the apex deeply emarginate and 2-lobed, the disk with a prominent fleshy bicarinate callus. Column subcylindric, the apex rather dilated and subclavate, with 2 short projecting teeth on the under-side. Panama, Colombia, Venezuela, British Guiana, Surinam, and Trinidad. L ZONE: vicinity Frijoles, sea level, Powell 3184; along Rio и pede between Gamboa and Alahuela, 30-60 m., Allen 962; Gatün Lake, sea level, Powell s. Á: hills east of rcg 2 Powell s. n.; vicinity Paja, near Empire, sea ед Pouell 3104; vicinity Juan Mina, Río рү ы 100 ft, Allen & Allen 4133. corów: Cativo-Porto Bello trail, sea level, Powel 71. TRIZEUXIS Lindl. TrizEuxis Lindl. Collect. Bot. Z. 2. 1823; Benth. & Hook. Gen. Pl. 3:566. 1883. Trixeuxis Lindl. Orch. Sel. 15. 1826. Small epiphytic herbs with short, distichous, foliaceous stems, the imbricating leaf bases distichously arranged in the form of an open fan and usually enveloping a small, monophyllous pseudobulb. Leaves falcate to gladiate, acute. Inflores- cences paniculate scapes produced from the bases of the pseudobulbs and con- spicuously exceeding the leaves. Flowers subglobose, minute, in dense racemes terminating the branches of the panicle. Sepals dissimilar, of about equal length, the dorsal sepal arching, deeply concave, the laterals connate for nearly their entire length, forming a single bifid segment below the lip. Petals elliptic-ovate, obtuse or subacute, about as long as the sepals. Lip entire, concave, articulated with the (480) 1949] " , FLORA OF PANAMA (Orchidaceae) 145 base of the column, the lateral margins erect and parallel with the column, the apex rather fleshy and recurved. Column subcylindric, erect, the base without a foot. Anther terminal, operculate, incumbent, 1-celled; pollinia 2, waxy. Two or three species of small American epiphytes ranging from Costa Rica to Trinidad, Brazil, Bolivia, and Peru. One species is known from Panama. 1. TRIZEUXIS FALCATA Lindl. Collect. Bot. £. 2. 1823. Trizeuxis andina Schltr. in Fedde, Rep. Sp. Nov. Beih. 10:52. 1922. Small to minute, erect, epiphytic herbs 2.5-10 cm. tall, with short, distichous, foliaceous stems, the imbricating leaf bases usually enveloping a small, subquadrate, compressed, monophyllous pseudobulb. Leaves falcate or gladiate, acute or acum- inate, the complanate blades 1.5—8 cm. long and 0.3—1.0 cm. wide, the imbricating bases distichously arranged in the form of an open fan. Inflorescences slender, erect, paniculate scapes 3—20 mm. long, produced from the bases of the pseudo- bulbs, conspicuously exceeding the leaves. Flowers subglobose, minute, in dense, subcapitate or elongate racemes terminating the branches of the panicle. Sepals dissimilar, of about equal length, green or pale yellow, the dorsal sepal ovate, acute, arching and deeply concave, about 2 mm. long, the laterals connate for nearly their entire length, forming a single obovate, bifid segment about 2 mm. long below the lip. Petals elliptic-ovate, obtuse or subacute, about as long as the sepals. Lip entire, lanceolate, acute, about 2.5 mm. long, the base concave, the lateral margins erect and closely appressed to the sides of the column, the apex rather fleshy and recurved, ovate, acute, yellow or orange, about № the total length of the lip. Column short, stout, subcylindric, about 1 mm. long, the apex somewhat dilated, the base without a foot. Anther very large in.proportion to the size of the flower, about 1.2 mm. long. Costa Rica, Panama, Venezuela, Trinidad, Brazil, Bolivia, and Peru. uas: vicinity Santiago, 700 ft., Powell 3526. снікюсі: vicinity Concepción, 800 y ^" Powell 286. 72. TRICHOPILIA Lindl. TRICHOPILIA Lindl. Nat. Syst. Bot. ед. 2, 446. 1836; Benth. & Hook. Gen. РІ. 3:559. 1983. Pilumna Lindl. in Bot. Reg. n. s. 7: TH. 73. 1844. Helcia Lindl. loc. cit. % Misc. 17. 184 Leucobyle Kissed in Ind. Sem. Hort. ү: Вего]. Арр. 1. 1854. Tricbopbila Pritz. Icon. Bot. Ind. 1115. 1855, sphalm. Erect epiphytic herbs with approximate monophyllous pseudobulbs. Leaves subcoriaceous to coriaceous or sometimes fleshy, usually elliptic-lanceolate to ligular, rarely narrowly linear or semiterete. Inflorescences slender, short or elongate, erect, arching or pendulous, 1- to 7-flowered scapes produced from the bases of the pseudobulbs. Flowers usually large and attractive. Sepals narrow, (481) [Vor. 36 146 ANNALS OF THE MISSOURI BOTANICAL GARDEN widely spreading, often twisted, usually subequal and free, the laterals sometimes connate at the base, rarely as far as the middle. Petals subequal to the dorsal sepal. Lip entire or 3-lobed, clawed at the base and adnate to the base of the column, the limb spreading or the lateral lobes convolute and forming a tube, the mid-lobe spreading, the disk smooth or lamellate. Column erect, semiterete, the margins of the apex projecting, entire or lobed, usually dentate or fimbriate, the base of the column without a foot. Anther terminal within the clinandrium, operculate, incumbent, 1-celled; pollinia 2, waxy. A small genus of tropical American epiphytes ranging from southern Mexico to Peru, Brazil, and the West Indies. It is another of the frequently recurring and perplexing instances in which the specific concepts within the genus are by no means of equal value. In this case there are several groups of closely related entities, each group centering on an archetypal species, and more closely related to it, and each to the other, than to other similar groups within the genus. For example, Trichopilia subulata is obviously closely allied to, if not actually identical with Т. mutica; and both markedly distinct from the archetype T. suavis, and its associate Т. leucoxantha, ог from the archetype Т. tortilis, with its closely allied species T. marginata, T. maculata, T. turialbae, T. Galeottiana, etc. Due to the lack, in most instances, of types or adequate series upon which to base comparative studies, I hesitate at this time to undertake any such wholesale overhauling of names as would seem to be required on the basis of probable relationships. Тһе present treatment, for the most part, follows established usage, but it is to be expected that many of these minor concepts may ultimately be reduced to varietal status or synonymy. Ас the present time six separable specific concepts are гер- resented in Panama. a. — ЭЕ Leaves fleshy, narrowly linear to semi- lar terete, 10 or less. Lip not tubu 5. T.SUBULATA aa. Pseudobulbs лк ағайы linear то suborbi cular. 2" coriaceous to subcoriaceous, ligular to elliptic-lanceolate, 15 wide or more. Lip tubular. b. Құ dobulbs a p ЖК more than 4 times as long as broad. dh e nate to about the middle. . T. TURIALBAE ral s nate at the base........ T. MARGINATA bb. Рита к 5 со suborbicular, less than 3 times as long s broad. c. Leaves 2—3.5 wide. Basal imbricating vine of the pseudo- к“ conspicuously maculate. Scapes 1-flower the inner lip conspicuous keel not equaling the inde in length...... T. MACULATA сс жк ауе vi cm. wide. Basal imb era racts of the pseudobulbs not maculate. Scapes several-flowered, the inner lip with a prom- inent . "nd e oer ог еа the column іп length. d. Sep 5—3 попри ..1. T. LEUCOXANTHA аа. бер жу 2 cm. loni Bertie etalk |. 4. Т. SUAVIS — TRICHOPILIA LEUCOXANTHA L. Wms. іп Am. Orch. Soc. Bull. 10:137. 1941. Erect, epiphytic herbs 20—25 cm. tall, with approximate, elliptic-oblong to suborbicular, compressed, monophyllous pseudobulbs 2.5—4.5 cm. long and 2—3.5 cm. wide, the bases enveloped in several thin, papery, imbricating bracts which (482) 1949] FLORA OF PANAMA (Orchidaceae) 147 “ГРЛСНОРПЯА : `, leucacantha: LWA Fig. 188. Trichopilia leucoxantba (483) [Vor. 36 148 ANNALS OF THE MISSOURI BOTANICAL GARDEN soon weather away. Leaves elliptic-lanceolate, acute, subcoriaceous, 15—20 cm. long and 4—6 cm. wide, contracted at the base into a short, conduplicate petiole. Inflorescences arching or pendulous, 1- to 4-flowered scapes 5—7 cm. long, pro- duced from the bases of the pseudobulbs. Flowers of moderate size, white with a pale yellow blotch at the base of the lip. Sepals free, spreading, the dorsal sepal oblanceolate, acute, 2.5-3.5 cm. long and 0.7-0.8 cm. wide, the laterals lanceolate, acute, 2.5-3 cm. long and 0.6-0.8 cm. wide. Petals subequal to the dorsal sepal, elliptic-oblong, obtuse with a minute apicule, 2.5-3.2 cm. long and 0.9-1.0 cm. wide. Lip 3-lobed, broadly obovate when spread out, 3—3.5 cm. long and 3-3.5 cm. wide, the narrow base adnate to the base of the column, the lateral lobes rounded, incurving over the column, the anterior margins more or less undulant, the mid-lobe spreading or reflexed, deeply emarginate, the lateral lobules strongly undulate, the disk with a prominent central keel about equaling the column in length. Column slender, terete, 15-18 mm. long, the margins of the apex pro- jecting and forming а fimbiiate hood over the anther, the base of the column without a foot. Panama. LE: western slope and summit of Cerro Valle Chiquito, 700-800 m., Seibert 515; Ри El Valle де Antón, 600-1000 т., Allen 2401, 3564; доог of El Valle, 600 m., aircbi Thus far known only from the region of El Valle de Antón, in Coclé Province. The plants considerably resemble those of Trichopilia suavis, and may possibly rep- resent only a small-flowered variety of that species. 2. TRICHOPILIA MACULATA Rchb. f. in Bonplandia 3:215. 1855. Ттісбор а Powellii Schltr. іп Fedde, Rep. Sp. Nov. Всіһ, 17:77. 1922. Dwarf, epiphytic herbs; pseudobulbs elliptic-oblong, strongly ancipitous, 1- leaved, 2—5 cm. long and 1.5—2.5 cm. wide, approximate and inserted at an acute angle on the short rhizome, the bases enveloped in several densely spotted, imbri- cating, papery bracts. Leaves elliptic-lanceolate, acute, subcoriaceous, 5-12 cm long and 2-3.5 cm. wide. Inflorescences usually solitary, slender, arching or semi- pendulous 1-flowered scapes 4—6 cm. long, produced from the bases of the pseudo- bulbs. Flowers of moderate size. Sepals subequal, free, widely spreading, lanceolate, acuminate, often somewhat twisted, pale yellow or greenish yellow, 3-4 cm. long and 0.35—0.5 cm. wide. Petals subequal to the sepals, lanceolate, acuminate, pale or greenish yellow, about 3.5 cm. long and 0.5—0.6 cm. wide, often with undulant margins. Lip obscurely 3-lobed, obovate when spread out, about 3.5 cm. long and 1.6-1.8 cm. wide, contracted at the base and adnate to the base of the column, white, aging pale yellow, the inner lip yellow with numerous fine orange-red lines, the rounded lateral margins convolute, forming a tube, the mid- lobe emarginate, with spreading or reflexed lobules, the disk with a short, incon- spicuous, central keel not equaling the column in length. Column slender, 1949] FLORA OF PANAMA (Orchidaceae) 149 Fig. 189. Trichopilia maculata semiterete, the margins of the apex distinctly 3-parted and minutely denticulate, forming a hood over the anther. Panama. CANAL ZONE: vicinity Salamanca Hydrographic Station, upper Madden Lake region, 70-80 m., Dodge, Steyermark & Allen s. n.; Balboa, sea level, Powell 3422; drowned forest along Río Chagres, Madden Lake region, 66 m., Steyermark & Allen 16770; Barro Colo- rado Island, Gatün Lake, Shattuck 555; San Jose Island, Perlas Archipelago, Johnston 146, 1276. РАМАМА: hills near Panama City, Powell 3006; Orange River valley, 75 m., Killip 3247; vicinity La Chorrera, 20 m., Allen 2842; Río La Maestra, 0-25 m., Allen 51. Small, but attractive plants fairly frequent in the lowlands of the central and eastern Isthmian area. The species apparently is confined to Panama. Guatemalan specimens identified as this species seem more readily referable to the Mexican T. tortilis. (485) [Vor. 36 150 ANNALS OF THE MISSOURI BOTANICAL GARDEN 3. TRICHOPILIA MARGINATA Henfr. in Gard. Mag. Bot. 3:185. 1851. Tricbopilia coccinea Warscz. in Paxton's Flow. Gard. 2:79. 1851-52. Tricbopilia crispa Lindl. in Gard. Chron. 342. 1857. Trichopilia lepida Hort. ex Flor. Mag. n. s. t. 08. 1874. Erect, epiphytic herbs with approximate, linear, broadly truncate, strongly ancipitous, l-leaved pseudobulbs 6-14 cm. long and 1-2.5 cm. wide, the bases enveloped in several imbricating, papery, usually conspicuously maculate sheaths. Leaves elliptic-lanceolate to lanceolate, acute to acuminate, coriaceous, 12-30 cm. long and 3—5.5 cm. wide, contracted at the base into very short, conduplicate petioles. Inflorescences short, arching or pendulous, 2- to 3-flowered scapes pro- duced from the bases of the pseudobulbs. Flowers large and conspicuous, very variable in color, most frequently with the sepals and petals reddish with lighter margins, the lip usually white on the outer surface, rarely red, the inner tube a deep rose-red, the reflexed margins of the mid-lobe often margined white. Sepals narrow, subequal, free, widely spreading, lanceolate to oblanceolate, acute, usually not twisted but sometimes with undulant margins, 4.5-6 cm. long and 0.6—1.0 cm. wide. Petals subequal to the sepals, spreading, oblanceolate, acute, 4-5.5 cm. long and 0.8—1.2 cm. wide, often with undulant margins. Lip tubular at the base, the apex obovate when spread out, 5—8 cm. long and 4—5 cm. wide, the nar- row base adnate to the base of the column, the lateral lobes rounded and convolute forming a tube, the mid-lobe deeply emarginate, the lateral lobules spreading or recurved, with undulant margins, the disk without a prominent keel. Column terete, the margins of the apex projecting, forming an obscurely 3-lobed, fimbri- ate hood over the anther. Guatemala, Costa Rica, Panama, and Colombia. үчин =s Caldera River flats, 3800 ft., Powell 3333; without definite locality, 4000 ft., Powell 154, 3201; forests around El Boquete, 1000-1300 m., Pittier 2060; in A eavy forest along road below El Hato, s. w. slopes of Chiriquí Voleano, 3800 ft., Allen s. A large-flowered, attractive species first collected by von Warscewicz in 1849 on the slopes of Chiriqui volcano and introduced into cultivation under the name of Trichopilia coccinea. The plants are fairly common in heavy forest ас 3500— 5500 ft. elevation, growing usually in dense shade on the lower trunks of the trees. They are strikingly similar to the earlier Т. tortilis of Mexico and Guatemala, and it seems very probable that our plants represent at most a somewhat more robust variety of that species. There are several well-marked color forms, of which the following are recorded from Panama. TRICHOPILIA MARGINATA var. ALBA Rchb. f. Flowers pure white, the inner lip lemon-yellow. TRICHOPILIA MARGINATA var. LEPIDA Veitch, Man. Orchid. Pl. 2:183. 1893. Flowers somewhat larger than in the type; the sepals and petals blotched rose- (486) 1949] FLORA OF PANAMA (Orchidaceae) 151 pink, as is also the frontal lobe of the lip; the sepals and petals with very broad white margins. Fig. 190. Trichopilia suavis (487) ` [Vor. 36 152 ANNALS OF THE MISSOURI BOTANICAL GARDEN TRICHOPILIA MARGINATA Var. OLIVACEA Rchb. f. Beitr. Orch. Centr.-Amer. 13. 1866. Sepals and petals olive-green. 4. TRicHOPILIA suavis Lindl. & Paxton in Paxton’s Flow. Gard. 1:44. 1850-51. Tricbopilia Kienastiana Rchb. f. in Gard. Chron. n.s. 20:166. 1883. Erect, epiphytic herbs with approximate, rather fleshy, oblong-ovoid, elliptic or suborbicular, laterally compressed, monophyllous pseudobulbs 3.5—8 cm. long and 2.5-6 cm. wide, the bases enveloped in several thin, papery, imbricating bracts which soon weather away. Leaves elliptic-lanceolate, acute, subcoriaceous, 10—40 cm. long and 3.5-8 cm. wide, contracted below into short or elongate, condupli- cate petioles. Inflorescences short, arching or pendulous, 2- to 5-flowered scapes produced from the bases of the pseudobulbs. Flowers large and attractive, fragrant, the sepals and petals white or creamy white sometimes spotted with pale rose-pink or red, the lip white or creamy white usually heavily spotted rose-pink, the inner tube most frequently with yellow or orange markings, very rarely blotched red. Sepals subequal, free, widely spreading, usually with undulate mar- gins, lanceolate, acute, with a central thickened nerve or keel on the dorsal surface, 3—5.5 cm. long and 0.6-1.0 cm. wide, the lateral sepals rarely somewhat connate at the very base. Petals subequal to the sepals, lanceolate, shortly acute, with undulate margins, 2.8-5 cm. long and 0.8-1.2 cm. wide. Lip tubular, 3-lobed, obovate when spread out, 4.5-6.5 cm. long and 3-5 cm. wide, the narrow base ad- nate to the base of the column, the rounded lateral lobes convolute forming a tube, the anterior margins usually crisped and undulate, the mid-lobe retuse or emar- ginate, the lobules undulate, crisped and reflexed, the disk with a prominent, erect, central keel exceeding the column in length. Column elongate, terete, the mar- gins of the apex projecting, forming a conspicuous, fimbriate, obscurely 4-lobed hood over the anther. Costa Rica, Panama, and Colombia. CHIRIQUÍ: without definite locality, 4000-5500 ft., Powell 135, 3334, 3342, ae = on southwest slope of Chiriqui Volcano and along Rio Chiriqui Viejo, This is one of the most attractive species of the genus. It was first discovered by von Warscewicz in 1848, very probably in Costa Rica; but by far the finest forms have been found on the slopes of Chiriqui volcano in Panama, where they are fairly common in low mossy woods at 3800—5500 ft. elevation. The fragrant flowers are variable in color, a pure white form being rather frequent. They are usually produced in March and April. 5. TRICHOPILIA SUBULATA (Sw.) Rchb. f. in Flora 48:278. 1865. Epidendrum subulatum Sw. Prodr. 123. 1788. Cymbidium subulatum Sw. іп Nov. Act. Soc. Upsal. 6:73. 1799. Leucohyle Warscewiczii K]. in Ind. Sem. Hort. Bot. Berol. App. 1. 1854. (488) 1949] FLORA OF PANAMA (Orchidaceae) 153 Fig. 191. Trichopilia subulata Tricbopilia bymenantba жаш: f. in Бона 2:90. 1854. Trichopilia jamaicensis Fawc. & Rendle, in Jour. Bot. 48:107. 1910. Leucobyle subulata (Sw.) Artis Die РАСТ = 469. 1914. Small, caespitose, epiphytic herbs 10-25 cm. tall, with short, inconspicuous, approximate, subcylindric, monophyllous pseudobulbs 1-2.5 cm. long and 0.2-0.5 cm. wide, the bases enveloped in several brown, papery, imbricating sheaths. Leaves fleshy, linear-lanceolate to semiterete, acuminate, 9—22 cm. long and 0.3-1.0 cm. wide, contracted at the base and apparently continuous with the apices of the slender pseudobulbs. Inflorescences pendulous, unbranched racemes 2.5-6 cm. long, produced from the bases of the foliaceous stems. Flowers small for the genus, usually 3-8, on slender pedicels, each subtended by a broad, acute to acuminate, spathaceous, papery bract. Sepals subequal, free, spreading, pure white to pale yellow, linear-lanceolate, acuminate, 20-23 mm. long and 3-4 mm. wide. [Vor. 36 154 ANNALS OF THE MISSOURI BOTANICAL GARDEN Petals subequal to the sepals, spreading, linear-lanceolate, acuminate, pure white to pale yellow, 18—20 mm. long and 2.5—3 mm. wide. Lip entire to obscurely 3- lobed, concave, obovate to elliptic-oblong, abruptly acute, or obtuse and apiculate, the margins minutely denticulate to conspicuously lacerate, white, usually irreg- ularly spotted rose-purple, 15—20 mm. long and 12-18 mm. wide, the base adnate to the base of the column, the lateral margins erect at the very base and somewhat incurving over the column, the limb spreading, the disk with a short, fleshy, bifid or bicarinate callus. Column semiterete, 7-9 mm. long, the margins of the apex projecting, forming an entire, denticulate to fimbriate hood over the anther, the sides of the clinandrium below the hood with a pair of short to elongate, denticu- late processes. Panama, Colombia, and Jamaica. CANAL ZONE: Barro BÉ Island, Wheeler trail, Shattuck 548; Upper Chagres sina sea level, Powell 241; Cerro Campana, vicinity ое 3000 ft., Allen 5142. LÉ: hills north of El Valle de буни 1000 m., Faircbild s Small but attractive epiphytes, vegetatively reminiscent of a slender form of Brassavola nodosa. The flowers in the specimens examined are quite variable. In the El Valle and Cerro Campana collections the margins of the lip are conspicuous- ly lacerate while in the lowland specimens they are only minutely denticulate; also, the lateral subulate processes on either side of the clinandrium have been found to be quite variable, scarcely any two specimens being exactly alike. Тһе simpler of these forms very closely approximate the earlier South American Tricbopilia mutica, and it seems quite probable that a more complete series of collections from the intermediate area would prove the two concepts to be one polymorphic species. 6. TRICHOPILIA TURIALBAE Rchb. f. in Hamb. Gartenzeit. 19:11. 1863, non Batem. Erect, epiphytic herbs with approximate, linear, strongly ancipitous pseudo- bulbs 5.5-10 cm. tall and 1.2-1.8 cm. wide, the truncate apices with a single leaf, the bases of the pseudobulbs enveloped in several papery, imbricating bracts. Leaves elliptic-lanceolate, acute, subcoriaceous, 15—20 cm. long and 3.5-5 cm. wide. Inflorescences short, arching or pendulous, 1- to 3-flowered scapes produced from the bases of the pseudobulbs. Flowers of moderate size. Sepals spreading, pure white, 2.5-3 cm. long and 0.4-0.5 cm. wide, the dorsal вера] free, lanceolate, acute, the lateral sepals connate to about the middle, acuminate. Petals subequal to the dorsal sepal, pure white, lanceolate, acuminate, 2.5-3 cm. long and 0.6-0.8 cm. wide. Lip tubular, white with pale orange lines in the throat, obscurely 3- lobed, obovate when spread out, 3.5-4 cm. long and 2.5-3 ст. wide, the narrowed base adnate to the base of the column, the lateral margins convolute and envelop- ing the column, the mid-lobe retuse at the apex, the lateral lobules reflexed and undulate, the disk with an elongate, central keel exceeding the length of the (490) 1949] FLORA OF PANAMA (Orchidaceae) 155 column. Column elongate, semiterete, the margins of the apex projecting and forming a distinctly 3-parted, minutely denticulate hood over the anther. Costa Rica and Panama. VERAGUAS: mountains of western Azuero, about 600 m., Е. R. Dunn s. n. Trichopilia turialbae Rchb. f. is very close to the earlier T. tortilis, differing in the smaller size and in color of the flowers, and in the conspicuously connate lateral sepals. However, several Guatemalan collections of T. tortilis have been ex- amined in which the flowers closely approximate ours in size. In all the material seen of T. tortilis, the lateral sepals were found to be connate at the base in some degree, although none to the middle as in our specimens. It seems rather likely, however, that intermediate forms will eventually be found and that T. twrialbae will be re- duced to varietal status under the archetypal species. 73. MESOSPINIDIUM Rchb. f. MzsosPINIDIUM Rchb. f. in Bot. Zeit. 10:929. 1852; Walp. Ann. 6:856. 1864. Epiphytic herbs with slender, complanate, monophyllous pseudobulbs enveloped at the base in several conspicuous, foliaceous bracts. Leaves at the apex of the pseudobulbs and the blades of the foliaceous bracts ligular to elliptic-lanceolate, subcoriaceous to pergameneous, the conduplicate, imbricating bases o the foli- aceous bracts forming a 2-ranked, more or less flabelliform cluster. Inflorescences slender, erect or arching panicles or racemes produced from the base of the cur- rent pseudobulb. Flowers small, on short, slender pedicels subtended by incon- spicuous bracts. Sepals rather fleshy, not widely spreading, the dorsal sepal free, the laterals connate for more than half their length, forming a single bifid seg- ment supporting the lip. Petals oblong, acute to cuneate and aristate, the broad bases inserted on the sides of the column. Lip entire, the base shallowly subsaccate and continuous with the base of the column, the limb oblong, obovate or subpandurate, the margins usually revolute, the apex acute to obtuse and emarginate, the disk with 2 fleshy, converging keels. Column short, very stout, apparently without wings or appendages, the base without a foot; pollinia 2, waxy. Two or three species of tropical American epiphytes, apparently closely allied to Oncidium and Odontoglossum. One species is known from Panama. 1. MzsospPiNIDIUM WanscEWwiCZIU Rchb. f. in Bot. Zeit. 10:929. 1852. Solenidium Endresii Kranzl. in Engl. Pflanzenr. IV, Fam. 50 (Heft 80):317. 1922. Erect, epiphytic herbs with distichous clusters of foliaceous bracts, the con- duplicate bases somewhat enveloping a slender, strongly ancipitous, nearly linear, monophyllous pseudobulb, the plants 12-22 cm. tall. Leaves of the apex of the pseudobulbs and the blades of the foliaceous bracts ligular to elliptic-lanceolate, acute to acuminate, pergameneous, 5-17 cm. long and 1-2.8 cm. wide, contracted below into slender, conduplicate petioles, the persistent bases of the bracts often somewhat broader than the petioles, in dried specimens superficially resembling (491) [Vor. 36 156 ANNALS OF THE MISSOURI BOTANICAL GARDEN the pseudobulbs. Inflorescences 1—2 slender, erect, racemose or paniculate scapes equaling or exceeding the leaves, 12—27 cm. tall, produced from the bases of the pseudobulbs. Flowers small. Sepals rather fleshy, not widely spreading, greenish yellow to tan, often spotted reddish brown, the dorsal sepal ovate, deeply concave, the apex conspicuously aristate, 6-8 mm. long and 2.5-3 mm. wide, the lateral sepals connate for more than half their length, forming a single bifid segment supporting the lip, 6—8 mm. long and 4—5 mm. wide. Petals rather fleshy, oblong- lanceolate, the apices aristate, the broad bases inserted on the sides of the column, the margins scarcely imbricating with those of the sepals, 4-5 mm. long and 2.5-3 mm. wide. Lip fleshy, entire, subpandurate when spread out, 5—7 mm. long and 2.5-3 mm. wide, white to pale yellow, usually with minute red spots, the oblong, subsaccate base continuous with the base of the column, the apex broader, with revolute margins, the disk with 2 converging fleshy keels. Column short, very stout, apparently without wings or appendages, about 4 mm. long and 4 mm. wide, the base without a foot. Costa Rica and Panama. PANAMA: Cerro Campana, vicinity Campana, in cloud forest near summit, 800-1000 m., Allen 3066. состЕ: vicinity El Valle de Antón, 650 m., Fairchild s. n.; hills north of El Valle de Antón, 1000 m., Allen 2761, 2785. 74. ODONTOGLOSSUM НВК. ODONTOGLOssuM НВК. Nov. Сеп. & Sp. 1:350, 7. 95. 1816; Benth. & Hook. Gen. Pl. 3:561. 1883. Cuitlauzina La Llave & Lex. Nov. Veg. Descr. 2:32. 1825. Cuitlanzina Lindl. Orch. Sel. 15. 1826. Cuitlauzinia Rchb. Nom. 54. 1841 Lichterveldia Lem. Illustr. Hortic. 2: f. 50. 1855. Osmoglossum Schltr. in Orchis 10:162. 1916, as subgenus. Epiphytic herbs with approximate or rarely distant, usually ovoid or elliptic- oblong, compressed, 1- to 3-leaved pseudobulbs, the bases enveloped in a few distichous, papery or foliaceous bracts. Leaves coriaceous or fleshy. Inflorescences produced from the bases of the pseudobulbs, sometimes short and 1-flowered, more frequently elongate, erect or arching, very rarely pendulous, few- to many-flowered racemes or panicles. Flowers usually large and conspicuous, but sometimes small. Sepals subequal, usually spreading, free or rarely with the lateral sepals united. Petals usually subequal to the dorsal sepal but sometimes broader. Lip 3-lobed or entire, the base continuous with the base of the column, the limb erect or parallel with the column, sometimes shortly adnate to it, the lateral lobes (if present) spreading or erect, the mid-lobe usually deflexed, less frequently spreading or concave, the apex acute, obtuse or emarginate, the disk at the base of the lip variously cristate, denticulate, lamellate, or rarely smooth. Column usually longer and more slender than in Oncidium, often clavate, the apex without appendages or sometimes with the margins produced into lobes, auricles, or teeth. Anther terminal, operculate, incumbent, 1-celled or imperfectly 2-celled; pollinia 2, Waxy. (492) 1949] FLORA OF PANAMA (Orchidaceae) 157 A large, polymorphic group of highland epiphytes having centers of rich specific and varietal development in the mountains of Mexico and Colombia, with some species ranging as far south as Brazil and Bolivia. As has been pointed out by Dr. Louis Williams in a footnote to the generic key (Ann. Mo. Bot. Gard. 33:7 [Fl Pan. 3?:113]. 1946) the concepts of Odontoglossum, Miltonia, Meso- spinidium, Aspasia, Brassia, Leochilus, and Osmoglossum are technically insepar- able from that of the earlier Oncidium, and might much more logically be treated as subgeneric sections of that genus. However, the species in each category are generally sufficiently distinctive to be readily recognizable by the average amateur, only the border-line species presenting any great difficulty. Practical ends seem to be better served by the maintainance of these entities as distinct, particularly for a treatment of this type, with the single exception of Osmoglossum, which, it appears, was never properly described or published, having been listed by Schlechter merely as a subgenus. Five species have been known from Panama, and a sixth, Odontoglossum convallarioides, which is very common in the highlands of adjacent Costa Rica, may possibly be represented in our series of specimens by a single fragmentary collection. a. Apex of the poni cn monophyllou b. Inflorescences racem Plant i к the pseudobulbs distantly in- serted on an Dat dus O. CHIRIQUENSE bb. cina 1-flowered. Hee dwarf, the pseudobulbs approxi- : 5. O. OERSTEDII aa. ee E the Зоран е ха ог triphyllous. b. Inflorescenc € 1. O. CARINIFERUM bb. Inflorescence ra Flow mall, ар 15 тт. long ог less. Leaves narrowly linear. ui is но; “sepals connate = more than half their length................ 4. O. EGERTONI Lateral sepals not connate 3. O. CONVALLARIOIDES cc. Flowers large, sepals 30 mm. long or more. Leaves elliptic-lanceo- . O. SCHLIEPERIANUM 1. ODONTOGLOSSUM CARINIFERUM Rchb. f. in Bot. Zeit. 10:638. 1852. Oncidium cariniferum (Rchb. f.) Beer, Prakt. Stud. Orch. 283. 1854 Odontoglossum hastilabium Lindl. var. fuscatum Hook. in Bot. Mag. t. 4019. 1856. Large, erect, epiphytic herbs with ovoid to elliptic-oblong, compressed, usually furrowed pseudobulbs 6-12 cm. long and 2.5-8 cm. wide, the apices with 2-3 leaves, the bases partially enveloped in several more or less distichous, imbricating bracts, the upper 2-3 of which are conspicuously foliaceous. Leaves coriaceous, linear-ligular, acute, 30—45 cm. long and 2.5-7 cm. wide. Inflorescences large, stout, erect or arching, many-flowered panicles, conspicuously exceeding the leaves, up to about 1 m. in length. Flowers relatively large, on long, slender pedi- cels, each subtended by an inconspicuous bract. Sepals subequal, usually free, spreading, brown, usually with yellow tips and margins, lanceolate, acuminate, somewhat concave, with a strongly developed dorsal keel, 2.5-3 cm. long and 0.4-0.6 cm. wide. Petals subequal to the sepals and similarly colored, lanceolate, acuminate, sometimes subfalcately incurving, 2-2.5 cm. long and 0.6-0.8 cm. (493) [VoL. 36 158 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN wide. Lip inconspicuously 3-lobed, 2—2.2 cm. long and 2—2.2 cm. wide, the much narrower basal half elliptic-oblong, adnate to the base of the column, the narrow lateral margins erect or spreading, the mid-lobe broadly biauriculate, white aging pale yellow, the apex very shortly acute or shallowly emarginate, with a short central apicule, the disk at the base of the lip often stained red, with 2 lateral subfalcate wings, the center with several projecting, fleshy teeth. Column slender, erect, about 10 mm. long, the ventral surface of the lower half conspicuously thickened, with 2 obscure lateral keels, terminating in 2 short, acute, fleshy wings below the stigma. Costa Rica, Panama, Venezuela, and probably adjacent areas. IQUÍ: without definite locality, 4000-5000 ft., Powell 1 142, Kieswetter s. n.; vicinity Cerro Punta, headwaters of the Río Chiriquí Viejo, 7000 ft., Allen 1 1517. A robust species much resembling ап Oncidium іп vegetative habit, found іп our area in Chiriquí Province, at 4000—7000 ft. elevation, where the plants seem to be confined to the tops of the tallest forest trees. 2. ODONTOGLOSSUM CHIRIQUENSE Rchb. f. in Bot. Zeit. 10:692. 1852. но нија Гони (Rchb. f.) Beer, Prakt. Stud. Orch. 283. 1854 Odontoglossum coronarium Lindl. Fol. Orch. Odontog. (21) oa 60, 1852. св coronarium (Lindl.) Веег, Зана Stud. Orch. 285. Robust, epiphytic herbs with stout, elongate, repent or semi-scandent rhizomes, the oblong-ovoid, compressed, monophyllous, usually dull purple pseudobulbs 7—11 cm. long and 4—6 cm. broad, distantly and obliquely inserted on the stems, the bases freely rooting, provided with 2—3 lateral, conspicuously foliaceous bracts, the long internodes completely enveloped in numerous papery, imbricating sheaths. Leaves and the blades of the foliaceous bracts elliptic-oblong, obtuse or retuse, coriaceous, 15-30 cm. long and 6—9 cm. wide, contracted at the base into short, conduplicate petioles. Inflorescences stout, erect, many-flowered racemes 30-45 cm. long. Flowers large and conspicuous, bright yellow richly blotched with reddish brown. Sepals free, subequal, widely spreading, elliptic- oblong to obovate, obtuse, with strongly undulate margins, 2—3 cm. long and 1.6—2 cm. wide. Petals subequal to the sepals and similarly colored, elliptic-oblong to obovate, obtuse to subacute, with strongly undulate margins, 2—3 cm. long and 1.5-1.8 ст. wide. Lip 3-lobed, subpandurate, 1.8—2.5 cm. long, the basal half conspicuously narrower than the apex and shortly adnate to the base of the column, the small lateral lobes erect, rather obliquely auriculate, obtuse, with undulate margins, the mid-lobe broadly spreading or reflexed, obovate, the apex obtuse or emarginate, the disk tuberculate at the base. Column short, stout, somewhat arcuate, 8-10 mm. long, the margins of the apex membranaceous and spreading. Costa Rica, Panama, Colombia, Peru, and probably adjacent territories. VERAGUAS: “Wild in Veragua; on the Ени of Miei зы at the height of 9000 ft., on decayed tree trunks. Flowers in October. Warcze (494) 1949] FLORA OF PANAMA (Orchidaceae) 159 Тһе type collection of this large-flowered and handsome species was presumably made in Panama, but it has not been found in recent years in our area, although it is fairly well known from adjacent Costa Rica and Colombia. It is often listed in horticultural publications as a variety of Odontoglossum coronarium. The reason for such treatment is a bit obscure, since Lindley, on the page following the type description of the latter species (Folia Orchidacea. Odontog. (22) No. 62. 1852), cites the place and date of the prior publication of O. chiriquense. 3. ODONTOGLOSSUM CONVALLARIOIDES (Schltr.) Ames & Correll, in Bot. Mus. Leafl. Harv. Univ. 11:19. 1943. Osmoglossum convallarioides Schltr. in Fedde, Кер. Sp. Nov. Beih. 19:148. 1923. Slender, erect, epiphytic herbs 30—45 cm. tall, the plants nearly identical with those of Odontoglossum Egertoni. Flowers fragrant, white, sometimes tinged with pink or lavender, the disk of the lip usually yellow. Sepals free, subequal, some- what spreading, elliptic-oblong, acute or subacute, concave, 7-10 mm. long and 3-6 mm. wide. Petals broader than the sepals, obovate to suborbicular, obtuse or minutely apiculate, 6-9 mm. long and 5-7 mm. wide. Lip obscurely 3-lobed, obovate, the apex subacute to obtuse or emarginate, 8-10 mm. long and 6—8 mm. wide, the lateral lobules spreading or semi-erect, the mid-lobe usually concave, the disk with two short keels, produced at the front into 2 erect, incurving, parallel, approximate, denticulate processes, the central area below the base of the column with a low, narrow, cuneate callus. Column short, very stout, about 3 mm. long and 3 mm. wide, the apex with obscure, subentire, membranaceous lobules sur- rounding the clinandrium, the base without a foot. Mexico, Guatemala, Honduras, Costa Rica, and probably Panama. A widespread Central American highland species, very closely allied to Odonto- glossum pulchellum, but differing in the much smaller flowers, the concave rather than arcuate-deflexed lip, and in the less conspicuous lobules surrounding the clinandrium. Although there are no authentic records to substantiate its occur- rence in Panama, one fragmentary collection (Woodson, Allen & Seibert 875) looks rather suspiciously like it, and in any event its frequent occurrence in the adjacent Costa Rican highlands would strengthen the probability that it will ultimately be found in Chiriqui. 4. ODONTOGLOssUM EGERTONI Lindl. in Bot. Reg. n. s. 8: Misc. 50. 1845. Osmoglossum Egertoni (Lindl.) Schltr. in Orchis 10:166. 1916, in зупоп. Osmoglossum acuminatum Schltr. in Fedde, Rep. Sp. Nov. Вей. 17:79. 1922. Osmoglossum anceps Schltr. loc. cit. 19:147. 1923. Erect, epiphytic herbs with approximate, narrowly elliptic-oblong or ovoid, tapering, compressed, often ridged, 2- to 3-leaved pseudobulbs 3.5—10 cm. long and 1.5-2.2 cm. wide, the bases partially enveloped in 2—3 distichous, conspicu- ously foliaceous bracts. Leaves and the blades of the basal bracts usually rather rigidly erect, narrowly linear-ligular, acute or acuminate, coriaceous, 25—45 cm. (495) [VoL. 36 160 ANNALS OF THE MISSOURI BOTANICAL GARDEN long and 0.8—1.5 cm. wide. Inflorescences erect, few-flowered racemes, usually not equaling the leaves, produced from the bases of the pseudobulbs, 20-30 cm. tall, the rachis conspicuously flattened, provided with a few, distant, complanate, conduplicate, acuminate bracts. Flowers small for the genus, white, the disk of the lip yellow with reddish spots. Sepals not widely spreading, lanceolate to elliptic-lanceolate, acute to shortly acuminate, concave, the dorsal sepal free, 1.2—1.5 cm. long and 0.4-0.6 cm. wide, the laterals united to beyond the middle, forming a single bifid segment below the lip, 1.3-1.5 cm. long and 0.6—0.8 cm. wide. Petals rather oblique, subequal to the dorsal sepal, ovate-elliptic, acute or shortly acuminate, 1-1.3 cm. long and 0.5-0.8 cm. wide. Lip entire, elliptic- oblong, acute or shortly acuminate, 10—12 mm. long and 6-7 mm. wide, somewhat concave or spreading, the apex sometimes reflexed, the disk with two low keels converging at the front and produced into 2 erect, parallel, fleshy teeth, the cen- tral area below the base of the column with a low, narrow, cuneate callus. Column very short, stout, 2-3 mm. long and 2—3 mm. wide, the apex with short, mem- branaceous, 3-parted, fimbriate to denticulate projections surrounding the clinandrium. Mexico, Guatemala, Honduras, Costa Rica, and Panama. CHIRIQUÍ: without definite locality, 6000 ft., Powell 255. Our specimens have somewhat narrower sepals and petals, and а somewhat more acuminate lip than in the type, but otherwise do not differ in the presence or absence of any essential character. It seems possible that all of the species of this alliance will ultimately be reduced to sub-specific rank or synonymy under the two distinctive archetypes, O. pulchellum and O. Egertoni. 5. ODONTOGLOssUM OznsrEDI Rchb. f. in Bonplandia 3:214. 1855. Dwarf, erect, epiphytic herbs up to 22 cm. tall, with approximate, ovoid, usually somewhat compressed, monophyllous pseudobulbs 1—3 cm. long and 0.8—1.8 cm. wide, the bases enveloped in several thin, papery, imbricating, non-foliaceous bracts which soon weather away. Leaves linear-ligular to elliptic-lanceolate, acute, coriaceous, 5-18 cm. long and 0.8—3 cm. wide, contracted below into elongate, slender, conduplicate petioles. Inflorescences erect, slender, 1-flowered scapes equaling or exceeding the leaves, 5-15 cm. long, produced from the bases of the pseudobulbs. Flowers of moderate size, solitary, white, fragrant, the base of the lip with a golden-yellow blotch, the callus densely spotted orange. Sepals free, subequal, spreading, elliptic-oblong, obtuse to subacute, sometimes with a short apicule, 12-20 mm. long and 6—10 mm. wide. Petals subequal to the sepals or sometimes broader, oblong-obovate to elliptic-oblong, obtuse, 12-20 mm. long and 8—12 mm. wide. Lip obovate, shortly clawed at the base, the narrow claw adnate to the base of the column, the apex of the lip broadly spreading, deeply emarginate and 2-lobed, 1.5-2.5 cm. long and 1.5-2.5 ст. wide, the disk with a short, erect, subquadrate, bicarinate, fleshy callus, the center concave, the posterior margin (4%) 1949] FLORA OF PANAMA (Orcbidaceae) 161 Fig. 192. Odontoglossum Schlieperianum (497) [Vor. 36 162 ANNALS OF THE MISSOURI BOTANICAL GARDEN directly below the column minutely ciliate. Column short, stout, 5-7 mm. long, without wings or appendages, the base without a foot. Costa Rica and Panama. cHIRIQUÍ: Potrero Muleto to the summit of Chiriquí Меш 3500-4000 т., Wood- son & Schery 455; Loma Larga to the summit, 2500-3380 m., Woodson, Allen & Seibert 1030; Casita Alta to Cerro Copete, 2300-3300 m., nee hy 3 Schery 337; without defi- nite locality, 10,000 ft., Davidson 1001; summit of Cerro Copete, on dead branches in Ericaceous barrens, 9000 ft., Allen 4005. А small but attractive species rather frequent on the highest slopes of Chiriquí Volcano at 7,000—10,000 ft. elevation. Descriptions in horticultural publications usually indicate that the scapes are 2- to 5-flowered, yet none of the fairly ex- tensive series of specimens in the Ames Herbarium show this to be the case, all having solitary flowers, which has also been the writer's observation in the field in the Chiriqui highlands and in Costa Rica. It seems possible that the plant illustrated in the “Botanical Magazine’ (7. 6820) under the name of Odontoglossum Oerstedii may be a white-flowered form of O. Krameri. 6. ODONTOGLOSSUM ScHLIEPERIANUM Rchb. f. in Gard. Chron. 1082. 1865. Odontoglossum Insleayi Bark. var. macrantbum Lindl. Fol. Orch. prid (4). 1852. Odontoglossum Warscewiczii Bridges ex ^ Orchideenb. 398 Odontoglossum Powellii Schltr. in Fedde, Rep. Sp. Nov. Beih. 17: гу Odontoglossum grande Lindl. var. pallidum Hort. ex. Sanders, Orchid ius 313. 1927. Erect, epiphytic or sometimes pseudoterrestrial herbs with approximate, fleshy, elliptic-oblong or ovoid, compressed, centrally ridged, grayish green, diphyllous pseudobulbs 4—12 cm. long and 2.5—5.5 cm. wide, the bases enveloped in several thin, papery, imbricating bracts which soon weather away. Leaves lanceolate to elliptic-lanceolate, acute, subcoriaceous, 10—30 cm. long and 3—7 cm. wide, con- tracted below into short or elongate, conduplicate petioles. Inflorescences stout, erect, 2- to 8-flowered racemes equaling or exceeding the leaves, 18—35 cm. long, produced from the bases of the pseudobulbs. Flowers large, yellow or greenish yellow, conspicuously barred and blotched with reddish brown. Sepals subequal, free, spreading, lanceolate, acuminate, 3-4.5 cm. long and 1—1.5 cm. wide. Petals broader than the sepals, oblong-oblanceolate, obtuse to subacute, 3—3.5 cm. long and 1.5—2 cm. wide. Lip obscurely 3-lobed, subpandurate when spread out, 2-3 cm. long and 1.0-1.5 cm. wide, the base contracted into a narrow claw which is adnate to the base of the column, the lateral lobes small, rounded or suborbicular, erect in natural position, the mid-lobe spatulate to obovate, obtuse or retuse, about 3⁄4 the total length of the lip, the disk with a short, central keel, from the mid- section and apex of which are produced 2 pairs of short, spreading, fleshy auricles. Column short, about 10 mm. long, terete below, dilated above, with a slender, acuminate, incurving tooth on each side of the stigma. Costa Rica and Panama. (498) 1949] FLORA OF PANAMA (Orchidaceae) 163 HIRIQUÍ: vicinity Casita Alta, Finca Lérida, eastern slopes of Eu. Volcano, 1500-2000 m., Woodson, Allen & Seibert 700; vicinity Boquete, 3800—5500 ft., Pring s. n., Davidson 797; without definite locality, “In gulch, in damp shady places,” 3800 f Powell 178; valley of the upper Río Chiriquí Viejo, 1300—1900 m., Seibert 227; vicinity Paso Ancho, Bambito Woods, along Rio Chiriqui Viejo, 4500—5000 ft., Allen 54. A common species of the Chiriqui highlands, closely allied to Odontoglossum grande of Mexico and Guatemala. 75. ASPASIA Lindley Aspasia Lindl. Сеп. & Spec. Orch. Pl. 139. 1833; Benth. & Hook. Gen. Pl. 3:560. 1883. Trophianthus Scheidw. in Otto & Dietr. Allg. Gartenzeit. 12:218. 1844. Epiphytic herbs with short, erect, approximate, usually complanate, cylindric stems conspicuously thickened above into compressed, 1- to 2-leaved pseudobulbs, the lower stems and bases of the pseudobulbs invested by the conduplicate, dis- tichously imbricating bases of several conspicuous, foliaceous bracts. Leaves and bract blades coriaceous or subcoriaceous. Inflorescences 1—2 short, erect, few- flowered racemes or rarely 1-flowered scapes, not equaling the leaves, produced from the axils of the uppermost foliaceous bracts. Flowers relatively large to small. Sepals of about equal length, spreading, the dorsal sepal subequal to the petals, the bases of the 3 upper segments connivent and adnate to the column usually somewhat above the insertion of the free lateral sepals. Lip shortly and narrowly clawed at the base, the claw adnate to the lower half of the column, the free limb spreading or with the apical half somewhat porrect, obscurely 3-lobed, subpandurate, subquadrate, suborbicular or obovate, sometimes entire, the lateral lobes (if present) rather broad, subdistinct or confluent with the usually larger mid-lobe. Column erect, semiterete, sometimes somewhat arcuate, the lower half connate with the claw of the lip, the ventral surface below the stigma broadly or narrowly sulcate, the margins sometimes with 2 short, denticulate projections, the apex truncate or sometimes continued into a short membranaceous hood over the anther, the column otherwise wingless, the base without a foot. Anther terminal, operculate, incumbent, 2-celled; pollinia 2, waxy. About ten species of tropical American epiphytes, ranging from Guatemala to Brazil. Two of these and one variety are thus far known from Panama. a. nd monophyllous. Plants 25 cm. tall or less. Column less than . long . A. PUSILLA aa. һамы diphyllous. Plants more than 25 cm. tall. Column 1.5 m. lon b. ateral “sepals 152 long, with broad transverse markin Column with an тибу petens: pir. the stigma. Apical half of ect . А. EPIDENDROIDES - Є" the li . Lateral enu 2.5 cm. long or more, with narrow longitudinal wea, Column with a narrow linear groove below the stigma. Apical half of the lip not porrect 2. A. EPIDENDROIDES Var. PRINCIPISSA (499) [Vor. 36 164 ANNALS OF THE MISSOURI BOTANICAL GARDEN bs. НИ © ы SSS ы сщ SS AN Ж nt oe Fig. 193. Aspasia epidendroides (500) 1949] FLORA OF PANAMA (Orchidaceae) 165 1. ASPASIA EPIDENDROIDES Lindl. in Hook. Jour. Bot. 1:6. 1834. Asbasia fragrans К}. Ind. Sem. Hort. Bot. Berol. 12. 1852. Odontoglossum Aspasia Rchb. f. in Walp. Ann. 6:851. 1861. Erect, epiphytic herbs with linear to oblong-elliptic, laterally compressed, diphyllous, stipitate pseudobulbs 5—14 cm. long and 2—6 cm. wide, the more robust plants having one side of the pseudobulb flattened, while the other surface is conspicuously convex, the complanate-cylindric lower portions and bases of the pseudobulbs invested with several closely imbricating bracts, the uppermost 2—3 of which are distichously arranged and conspicuously foliaceous. Leaves lanceolate to ligular, acute, subcoriaceous, 10—28 cm. long and 1.5-5 cm. wide. Inflorescences 1—2 simple, erect, few-flowered racemes 9—25 cm. long. Flowers of moderate size, the sepals greenish with broad transverse bands of brown or brownish lavender, the petals pale lavender to greenish brown, the lip white, with conspicuous purple or lavender markings in the center, the disk yellow, the column and anther tinged with lavender. Sepals of about equal length, elliptic-obovate, shortly acute, the apex concave with a short fleshy apiculate keel or thickening on the dorsal surface, 1.3-2 cm. long and 0.7-1.0 ст. wide, the base adnate to the base of the column slightly above the insertion of the petals, the lateral sepals strongly reflexed, nar- rowly and somewhat obliquely oblanceolate, acute, 1.5-2.2 cm. long and 0.6-0.8 cm. wide, the dorsal surfaces thickened into short apiculate keels at the apices. Petals subequal to the dorsal sepal, elliptic-obovate, concave, 1.5-1.8 cm. long and 0.6—0.9 cm. wide, the apices thickened into short, acute keels on the dorsal sur- faces. Lip clawed at the base, the claw adnate to the lower half of the column, the free limb abruptly deflexed, obscurely 3-lobed, subquadrate, 1.2-1.6 cm. long and 1.4-1.8 cm. wide, the basal and Apical halves of about equal width, the lateral margins at the base spreading, the mid-lobe emarginate and 2-lobed, with a thick- ened central keel, usually more or less porrect, the disk with 2 narrow parallel keels about equaling the lateral lobes of the lip in length. Column semiterete, slightly arcuate, about 1.5 cm. long, with a narrowly elliptic concavity below the stigma. Guatemala, Honduras, Nicaragua, Costa Rica, and Panama. ANAMA: along road to Pacora, about 50 m., Allen 822. cocrÉ: mountains beyond La PAREN 400—600 m., Hunter 9 Allen 633; Penonomé and vicinity, 50—1000 ft., R. S. Williams 445. VERAGUAS: vicinity Santiago, 600—800 ft., Powell 3542; vicinity kis de Jesús, sea level, Allen & Allen 4273. cutrigui: without definite locality, Powell 3017. 2. ASPASIA EPIDENDROIDES var. PRINCIPISSA (Rchb. f.) P. H. Allen, stat. nov. Aspasia principissa Rchb. f. in Bot. Zeit. 10:367. 1852. Aspasia Rousseauae Schltr. in Gartenfl. 71:76. 1922. Epiphytic herbs often nearly identical to Aspasia epidendroides, but usually somewhat less robust and with larger flowers. Sepals subequal, spreading, lanceo- late, acute to acuminate, pale green, longitudinally striped brown, the apices thickened and somewhat concave, usually 2.5-3 cm. long and 0.5-0.7 cm. wide, (501) [Vor. 36 166 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 194. Aspasia epidendroides var. principissa the dorsal sepal connivent at the base with the bases of the petals and adnate to the column somewhat above the insertion of the lateral sepals. Petals a little broader than the sepals, spreading, lanceolate, acute, 2.2-2.8 cm. long and 0.6-0.9 cm. wide. Lip narrowly clawed at the base, the claw adnate to the lower half of the column, the free limb abruptly deflexed, explanate, ob- scurely 3-lobed, subpandurate, 2-3 cm. long and 1.8-2.6 cm. wide, the basal half usually conspicuously broader than the apical half, the lateral lobes rounded and spreading, the mid-lobe emarginate, the lateral lobules somewhat undulate but never porrect, the disk with numerous radiating thickened nerves, the central 2 of (502) 1949] FLORA OF PANAMA (Orchidaceae) 167 which are parallel and exceeding the others, about equaling the lateral lobes of the lip in length. Column semiterete, lightly arcuate, 1.8-2 cm. long, creamy white aging pale yellow, adnate to the claw of the lip for about half of its length, the ventral surface below the stigma with a narrow groove, the base of the column without a foot. Costa Rica and Panama. AL ZONE: wet forest bordering Саша годи 30 m., Killip 3447; Culebra, 50-100 m., Pittier 3396; Cruces, sea level, Powell 3108. PANAMA: foothills near Panama City, sea level, Powell 39; Juan Díaz, sea a Powell per Chiva-Chiva, sea level, Powell 3029; Río La Maestra, Pacific coast between the Río Bayano and the Gulf of San Miguel, 0—25 m., Allen Қай San Jose ERE: Perlas Archipelago, Johnston 136, 702, 1015, 1288, Erlanson 199. LÓN: Que a Лаа, upper Madden Lake region, 70 m., Dod ge, graphic Wap 90 m., Hunter 9 Allen 653; Río Viejo, vicinity Puerto Pilón, sea level, Allen 4252. DARIÉN: forests around Yaviza, Pittier 6581; vicinity Marraganti, 10—200 ft., R. S. Williams 060. An attractive, large-flowered variety of Aspasia epidendroides, usually con- fined to areas of higher rainfall, being known from the vicinity of Puerto Limón, on the Atlantic coast of Costa Rica, and ranging eastward along the Atlantic seaboard, probably as far as Colombia. In the Canal Zone area, it follows the Río Chagres drainage to the headwaters of Madden Lake, crossing to the Pacific slope in the forested hills east of Panama City and continuing into Darién Province and the Perlas Islands, replacing the typical form from the Canal area eastward. 3. AsPASIA PUSILLA C. Schweinf. in Bot. Mus. Leafl. Harv. Univ. 10:21, /. 1. 1941. Small, erect, epiphytic herbs with approximate, linear or ellipsoid-complanate, monophyllous pseudobulbs 2.5—4 cm. long and 1.0—1.5 cm. wide, the complanate- cylindric, stipitate bases invested by several distichously imbricating, conduplicate sheaths the upper 2-3 of which are conspicuously foliaceous and enveloping the lower portions of the pseudobulbs. Leaves linear-ligular, obliquely acute, sub- coriaceous, up to 17 cm. long and 1.5 cm. wide, contracted below into short con- duplicate petioles. Inflorescences slender, erect, few-flowered racemes up to about 8 cm. long. Flowers small for the genus, on long slender pedicels, the sepals and petals yellow to greenish yellow with a dark brown basal blotch, the lip white or pale yellow, the disk orange marked with mauve or maroon stripes. Sepals sub- equal, free, spreading, the dorsal sepal elliptic-lanceolate to oblanceolate, acute, about 13 mm. long and 4.5 mm. wide, the laterals lanceolate, acute, about 13.5 mm. long and 4 mm. wide. Petals subequal to the dorsal sepal, elliptic-oblanceolate, obliquely acute, about 12 mm. long and 4 mm. wide. Lip clawed at the base, the claw adnate to the lower half of the column, the free limb suborbicular to obovate, entire, about 11 mm. long and 12 mm. wide, the apex slightly retuse, broadly spreading, the base somewhat concave, the disk with a short pubescent, divaricate callus. Column very short, stout, about 5.5 mm. long, the apex with a denticulate (503) [Vor. 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN 168 Aspasia pusilla Fig. 195. (504) 1949] FLORA OF PANAMA (Orchidaceae) 169 hood over the anther, the lateral margins below the stigma with 2 short, porrect, obliquely triangular teeth. Costa Rica and Panama. DARIEN: Cana-Cuasi trail, hills near Chepigana, 2000 ft., Terry 9 Terry 1502. A very distinctive dwarf species, known in our area only from the type col- lection. 76. BRASSIA R. Brown Brassia R. Br. in Ait. Hort. Kew. 2:215. 1813; Benth. & Hook. Gen. Pl. 3:564. 1883 Erect, epiphytic herbs with short stems usually thickened into conspicuous pseudobulbs that are infrequently rudimentary and inconspicuous or rarely entirely absent, the 1-3 leaves at the apex enveloped in several foliaceous or papery bracts, the conduplicate leaf bases distichously imbricating and forming a broad or narrow an. Inflorescences 1—2 erect or arching, few- to many-flowered racemes produced from the bases of the pseudobulbs or concurrently with the flush of new growth, or, if the plants are pseudobulbless, from the axils of the leaves. Flowers usually large and conspicuous, subtended by small and inconspicuous, or elongate spath- aceous bracts. Sepals free, spreading, narrowly acuminate or caudate, sometimes of about equal length but more frequently with the lateral sepals conspicuously longer. Petals subequal to the dorsal sepal or smaller. Lip entire or obscurely 3-lobed, spreading, shorter than the sepals, the base sessile and adnate to the base of the column, the disk usually bilamellate. Column short, erect, without wings or appendages, the clinandrium scarcely prominent, usually truncate. Anther terminal, operculate, incumbent, 1-celled or imperfectly 2-celled; pollinia 2, waxy. A small genus of American epiphytes, ranging from southern Florida and the West Indies to Mexico, Central America, and northern South America to Brazil, Bolivia, and Peru. They are technically inseparable from Oncidium, yet the majority of the species can readily be distinguished by the conspicuously elongate lateral sepals. Five species are known from Panama. a. Plants without pseudobulbs, or apparently so b. Leaves many, forming a broad ins px КР without pseudobulbs. 3 m. Е ог то 1 Lateral sepals 3.5 С В. ALLENII ББ. Leaves few, for nar ichous petiole, usually іе сел а small rudimentary ыс “Laer sepals 2 cm. long or less....3. В. CHLOROPS 22: Plants withou ееси к ; b. Lateral sepals 2 cm. long o 3. B. cHLOROPS bb. Lateral sepals 3 cm. long or more. c. Pseudobulbs monophyllous at the ap 5. B. LONGISSIMA cc. Pseudobulbs 2.” ог rarely t кз: е ас the а d. Apical half of the lip ups broader than de bead half. Vai d into broad plates at the front. Highland ecies . B. GIREOUDIANA dd. e half of the lip not conspicuously broader than the basal alf. Calli at the front with 2 separate and distinct slender teeth. Lowland species 2. B. CAUDATA (505) [Vor. 36 170 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1. Brassia Аттеми L. О. Wms, ex C. Schweinf. in Bot. Mus. Leafl. Harv. Univ. 13:145, £. 12. 1948. Erect, epiphytic herbs entirely without pseudobulbs, the plants 30—38 cm. tall, nearly identical in vegetative appearance with Huntleya meleagris. Leaves 8-14, distichously arranged in the form of a broad fan, linear-ligular to lanceolate, usually shortly and subfalcately acuminate, subcoriaceous, 15—35 cm. long and 2—4 cm. wide, the conduplicate bases 2-ranked and closely imbricating, usually slightly broader at the suture line than the leaf petioles. Inflorescences erect or arching, 5- (о 8-flowered, congested racemes V5 to 25 the length of the leaves. Flowers of moderate size, the lip usually uppermost in natural position, very fragrant, subtended by broad, acute or acuminate, papery, spathaceous bracts. Sepals subequal, free, spreading, reddish tan to olive-ocher, shading to cinnamon- buff at the base, linear-lanceolate, long-acuminate, 3.5—4 cm. long and 0.4-0.5 cm. wide. Petals subequal to the sepals and similarly colored, with a small tri- angular barium-yellow spot at the base, obliquely lanceolate, long-acuminate, the attenuate apices subfalcately incurving, 3—4 cm. long and 0.5—0.6 cm. wide. Lip entire, subquadrate to suborbicular, spreading, slightly convex, 15—20 mm. long and 13-16 mm. wide, barium-yellow, with a narrow, semicircular band of Mikado brown spots surrounding the disk, the base sessile and adnate to the base of the column, the apex abruptly contracted into an elongate, acuminate apicule, the disk white, with 2 short, stout, erect, parallel, strontian yellow keels, terminating at the front in 2 short, obtuse, subconic teeth. Column very short, stout, terete, about 5 mm. long, green or white, with 2 lateral Mikado brown blotches, without wings or appendages, the base without a foot. Panama. diens cloud forest on summit of Cerro uses 3000 ft., Allen & Fairchild 5150. ntains beyond La Pintada, 400-600 m., Hunter & Allen 502; hills north of El Valle %- Antón, 800-1000 m., Fairchild s. n., ү s. n., Allen 374, 2830, 2922, 3717. A very distinctive species entirely without pseudobulbs, apparently most nearly allied to Brassia glumacea Lindl. of Venezuela. The plants are nearly identi- cal in superficial appearance with those of Huntleya meleagris, sometimes differing in the shortly and subfalcately acuminate apices of the leaves, and in the condup- licate leaf bases which are often rather noticeably dilated at the suture line and not uninterruptedly confluent with the leaf petioles, as in the latter. All specimens seen have originated in the cool wet forests north of El Valle de Antón, the cloud ferest cap of Cerro Campana, or in the mountains west of Santa Fé, in Veraguas; the only presently accessible areas in Panama representative of the extensive and poorly known intermediate highlands of the Atlantic slope. 2. Brassta CAUDATA (L.) Lindl. in Bot. Reg. 10: /. $32. 1824. Epidendrum caudatum L. Sp. Pl. ed 2, 1349. 1763. 1805. Malaxis caudata Willd. Sp. Pl. 4:93. Oncidium caudatum Rchb. f. in Walp. Ann. 6:766. 1863. (506) 1949] 171 FLORA OF PANAMA (Orchidaceae) mec mcm E а i // [| A Fig. 196. Brassia Allenii (507) | [Vor. 36 172 ANNALS OF THE MISSOURI BOTANICAL GARDEN Brassia Lewisii Rolfe, in Orch. Rev 99, 189 Brassia MM (Rchb. +.) Schlen. var. minor Ж Schltr. in Fedde Rep. Sp. Nov. Beih. 17:8 242. \ WI SW >. pt || n" | / Fig. 197. Brassia caudata (508) 1949] FLORA OF PANAMA (Orchidaceae) 173 Erect, epiphytic herbs with approximate, linear to oblong-elliptic, ancipitous pseudobulbs 6—15 cm. long and 2—4 cm. wide, the truncate apices usually with 2 or rarely with 3 leaves, the bases of the pseudobulbs enveloped in 4—6 papery, closely imbricating bracts, the uppermost sometimes foliaceous. Leaves ligular to elliptic-oblong, obtuse to shortly acute, coriaceous, 16—27 ст. long and 2.5—6 cm. wide, contracted below into short conduplicate petioles. Inflorescences 1 or 2 arching, unbranched, usually 6- to 12-flowered racemes 15—30 ст. long, produced from the lateral bases of the pseudobulbs, the rachis often dull red, often rather complanate. Flowers large, with conspicuous caudate lateral sepals. Sepals free, spreading, greenish yellow to yellow, usually spotted reddish brown, particularly near the base, the dorsal sepal erect, linear-lanceolate, long-attenuate, 4—9 cm. long and 0.4-0.7 cm. wide, the apical half usually incurving, the lateral sepals conspicuously longer, caudate, of quite variable length, 12-30 cm. long and 0.5—0.6 cm. wide at the base, the outer surfaces with a strongly developed central keel. Petals free, greenish yellow or yellow, usually heavily spotted reddish brown near the base, the long attenuate apices falcately incurving, 2-3 cm. long and 0.3—0.4 cm. wide. Lip entire, oblong-lanceolate, long-acuminate, somewhat con- vex, often with deflexed lateral margins, yellow with reddish brown markings at the base, 3—5 cm. long and 1.0-1.2 cm. wide, the base sessile and adnate to the base of the column, the disk with 2 erect parallel lamellae in front of which are 2 short, distinct, acute teeth. Column stout, 4—5 mm. long, without wings or ap- pendages, the base without a foot. Mexico to Brazil and Bolivia; Florida; Greater Antilles. CANAL ZONE: Miraflores, Powell 3231; Frijoles, Powell 3117, 3256, Killip 3448; Pedro Miguel, Powell 3258, 3530, 3566; Ancón, Pittier 6631; Barro Colorado Island, Gatún Lake, Shattuck 202. PANAMA: hills east of Panama City, Powell 3275, Allen 4560. DARIÉN: vicinity La Palma, 0—50 m., Pittier 6619. CHIRIQUÍ: exact locality lacking, sea level, Powell 87. i А very common and widely distributed species of the lowlands of the American tropics and subtropics. The flowers are somewhat variable in size, those with the longest sepals sometimes being confused with Brassia longissima, a very different species with monophyllous pseudobulbs, of the cool intermediate highland forests of the Atlantic slope. Brassia longissima var. minor is simply а small-flowered form of B. caudata. 3. BRASSIA CHLOROPS Endres & Rchb. f. in Gard. Chron. 542. 1873. Brassia parviflora A. & S. in Sched. Orch. 8:74. 1925. Erect, caespitose, epiphytic herbs usually with slender, complanate-elliptic, rudimentary, monophyllous pseudobulbs often completely enveloped by the con- duplicate leaf bases. Leaves lanceolate, shortly acuminate, subcoriaceous, 8—30 cm. long and 1.4—3.0 cm. wide, the conduplicate bases distichously imbricating, often forming a short, narrow, complanate petiole, the leaves toward the base be- coming progressively shorter, the lowest reduced to non-foliaceous bracts. Іп- (509) [Vor. 36 174 ANNALS OF THE MISSOURI BOTANICAL GARDEN florescences erect, 3- to 7-flowered racemes 15—25 cm. long, produced from the axils of the upper leaves. Flowers very small for the genus. Sepals rather fleshy, free, subequal, spreading, linear-lanceolate, acuminate, 1.4-2 cm. long and 0.2—0.3 cm. wide. Petals subequal to the dorsal sepal, obliquely lanceolate, acuminate, 1.1-1.5 cm. long and 0.15-0.25 cm. wide. Lip entire, rather fleshy, elliptic-lanceo- late to linear-lanceolate, acute, 1-1.1 cm. long and about 0.45 cm. wide, the apex usually dorsally carinate and recurved, the base adnate to the base of the column, the disk with 2 fleshy parallel pubescent keels, about V2 the total length of the lip, the apices terminating in 2 more or less prominent teeth. Column short, stout, about 5 mm. long, the base without a foot. Costa Rica and Panama. CHIRIQUÍ: vicinity Casita Alta, Finca Lérida, eastern slopes of Chiriquí Volcano, 1500-2000 m., Woodson, Allen & Seibert 874. An aberrant highland species, having the aspect of a small-flowered Odonto- glossum. It differs from most species of Brassia in the short sepals which are of about equal length, green with darker spots or brown with yellow markings. Our single specimen differs in color, and somewhat in structural detail from the type, but seems more readily referable to this species than to any other. 4. Brassia GIREOUDIANA Rchb. f. & Warscz. in Otto & Dietr. Allg. Gartenzeit. 22:273. 1854. Oncidium Gireoudianum Rchb. f. in Walp. Ann. 6:768. 1863. Erect, epiphytic herbs with approximate, fleshy, elliptic-ovoid, rather com- pressed, usually longitudinally ridged, diphyllous pseudobulbs 7-11 cm. long and 2.5—4 cm. wide, the bases enveloped in several papery, imbricating bracts the uppermost 1—2 of which are sometimes foliaceous. Leaves ligular to lanceolate, acute, coriaceous, 22—35 cm. long and 2-5 cm. wide, the bases contracted into short conduplicate petioles. Inflorescences usually solitary, elongate, arching, many-flowered racemes equaling or exceeding the leaves, usually produced con- currently with the flush of new growth from the axils of the expanding leaves or sometimes from the bases of the newly completed pseudobulbs. Flowers large and conspicuous, with elongate filiform sepals and petals. Sepals free, widely spread- ing, long-attenuate, greenish yellow with a few brown blotches near the base, the dorsal sepal narrowly linear-lanceolate, 9—10 cm. long and 0.3—0.4 cm. wide near the base, the lateral sepals somewhat longer, narrowly linear-lanceolate, 10—15 cm. long and 0.4-0.5 cm. wide near the base. Petals subequal to the dorsal sepal, spreading, the apical half greenish yellow, the basal half brown, narrowly linear- lanceolate, long-attenuate, 5-6 mm. long and 3-4 mm. wide near the base. Lip entire, rather rhombic in outline, 3—4.5 cm. long and 1.5-2.5 cm. wide, pale yellow with sparse brown spots and blotches, the basal portion subquadrate to oblong, adnate to the base of the column, the apical portion abruptly dilated and cordate, shortly acuminate or apiculate, the disk with 2 fleshy, minutely puberulent (510) 1949] FLORA OF PANAMA (Orchidaceae) 175 keels, the basal portions erect and parallel, terminating at the apex in 2 larger subquadrate spreading lobules. Column very short, stout, 3.5-5 mm. long, the base without a foot. Costa Rica and Panama. cHIRIQUÍ: “On banks of rivers", 3500-4500 ft., Powell 143; Llano del Volcán, and along Rio Chiriqui Viejo, 1200 m., Allen 015. 5. BRASSIA LONGISSIMA (Rchb. f.) Nash in Bailey, Stand. Cyclop. Hort. 1:541. March, 1914. Brassia Lawrenciana Lindl. var. longissima Rchb. f. in Gard. Chron. 1513. 1868. Brassia longissima (Rchb. f.) Schltr. Die Orchideen, 496. October 1914, as to basinym, but only in part as to plant described. Erect, epiphytic herbs with approximate, very strongly flattened, oblong to elliptic-oblong, monophyllous pseudobulbs, with very thin, acute margins, 4—16 cm. long and 3—5 cm. wide, the bases enveloped in several conduplicate, imbri- cating, papery bracts, the uppermost one of which is usually shortly foliaceous. Leaves oblong-lanceolate, acute, coriaceous, 25-55 cm. long and 4—7 cm. wide, the bases contracted into very short, conduplicate petioles. Inflorescences 1 or rarely 2, arching, few-to many-flowered racemes 25—45 cm. long, produced from the bases of the pseudobulbs. Flowers large and conspicuous, with elongate, fili- form, lateral sepals. Sepals yellow to greenish yellow, spotted or blotched reddish brown near the base, the dorsal зера! narrowly linear-lanceolate, long-attenuate, 8—10 mm. long and 3—4 mm. wide at the base, the lateral sepals conspicuously longer, narrowly linear-lanceolate, caudate, 10-18 cm. long and 0.3—0.4 cm. wide. Petals shorter than the dorsal sepal, yellow to greenish yellow, reddish brown near the base, narrowly linear-lanceolate, the attenuate apices subfalcately incurving, 4—5 cm. long and 0.3—0.4 cm. wide at the base. Lip entire, pale yellow to greenish white, with sparse reddish brown spots near the base, lanceolate, acuminate, 3—5 cm. long and 1.0—1.5 cm. wide, the base somewhat concave with erect margins, adnate to the base of the column, the disk with an erect, fleshy, minutely puberu- lent, bilamellate, obliquely truncate callus. Column short, semiterete, about 8 mm. long, the base without a foot. Costa Rica, Panama, Peru, and probably adjacent territories. СТЕ: region north of El Valle de Antón, 1000 m., Fairchild s.n.; Loma del Tigre, hills Ed of El Valle, 3000 ft., Allen 4562. This very distinctive species has thus far been found only twice in Panama, both times in the wet highland forests north of El Valle de Antón, in Coclé Province, at about 3000 ft. elevation, although the species is to be expected gen- erally in similar areas on the Atlantic slope. Since there seems to be some confusion in regard to the application of the name Brassia longissima, it might be well to give something of its history. Both (511) [Vor. 36 176 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 198. Brassia longissima Nash (in Bailey Stand. Cyclop. Hort. 1:541. March, 1914) and Schlech- ter (Die Orchideen, 496. October, 1914) decided that the entity hitherto known as Brassia Lawrenciana Lindl. var. longissima Rchb. f. actually represented a dis- tinct species, citing Reichenbach's variety as the name-bringing synonym. The (512) 1949] FLORA OF PANAMA (Orchidaceae) 177 accompanying description given by Nash, whose publication antedated that of Schlechter by some six months, essentially followed Reichenbach's original, while that of Schlechter was apparently based on two distinct elements: the Reichenbach variety of B. Lawrenciana, and a very different plant that we now know was a large-flowered form of B. caudata (Powell 87). Although under the rules, the name of B. longissima (Rchb. f.) Schltr. must be regarded as synonymous with the name-bringing synonym, or the first published derivative name therefrom, there can be no doubt that the plant which Schlechter had in mind was a large- flowered form of B. caudata. This is further borne out by his subsequent descrip- tion of another, smaller-flowered form of B. caudata under the name of B. longis- sima var. minor (in Fedde Rep. Sp. Nov. Beih. 17:80. 1922). 77. MILTONIA Lindl. MILTONIA Lindl. in Bot. Reg. 23: sub Z. 1976. 1837; Benth. & Hook. Gen. РІ. 3:563. 188, ? Gynizodon Raf. Fl. Tellur. 4:40. Macrochilus Knowles & Westcott, Fl. (3n 1:93, K 45. 1837. Epiphytic herbs, with short, usually rather inconspicuous, compressed pseudo- bulbs bearing 1—2 leaves at the apex, the base enveloped in few to many distichous, imbricating, conspicuously foliaceous bracts. Leaves and bract blades elliptic- lanceolate to narrowly linear, coriaceous to subcoriaceous, contracted at the base into short or elongate conduplicate petioles. Inflorescences axillary from the base of the pseudobulb, often short, erect or arching, 1- to few-flowered scapes or sometimes elongate, many-flowered racemes. Flowers relatively large and con- spicuous, on long slender pedicels subtended by elongate, spathaceous, or sometimes minute and inconspicuous bracts, the perianth segments and the lip usually being all on one plane so that the flowers are typically flat. Sepals subequal, spreading, free, or the laterals very shortly connate at the base. Petals subequal to the sepals or a little broader. Lip entire, broadly spreading, the apex often bifid, the base sessile or very shortly and broadly clawed and affixed to the base of the column usually at a right angle, the disk inconspicuously or sometimes prominently lamel- late. Column short, the apex or anterior portion variously 2-auriculate or 2-alate, the clinandrium short, truncate, or the apex 2-lobed, or sometimes membranaceous and dilated, 2- to 3-lobed, the base without a foot. Anther terminal, operculate, incumbent, 1- or imperfectly 2-celled; pollinia 2, waxy About 20 species of attractive, mostly highland ка. closely allied to Oncidium and Odontoglossum, having centers of development in the mountains of Colombia and southern Brazil, with a single species known from as far north as Costa Rica. 1. MirroNiA Enprestt Nichols. in Dict. Gard. 2:368. 1888. Odontoglossum Warscewiczii Rchb. f. in Bot. Zeit. 10:692. 1852. Miltonia superba Schltr. in Fedde Rep. Sp. Nov. 3:249. 1907. (513) [VoL. 36 178 ANNALS OF THE MISSOURI BOTANICAL GARDEN Erect, caespitose, epiphytic herbs with distichous, foliaceous stems 16—38 cm. tall, the conduplicate, imbricating leaf bases enveloping a small complanate, ellip- tic-oblong, monophyllous pseudobulb 3—5 cm. tall and 1.2—1.5 cm. wide. Leaves narrowly linear to lanceolate, obliquely acute to acuminate, coriaceous, the blades 12—30 cm. long and 1-2.5 cm. wide, contracted below into short or elongate conduplicate petioles. Inflorescences 1 or 2 erect or arching, 2- to 5-flowered racemes 15—30 cm. long, produced from the lateral bases of the pseudobulbs. Flowers large and conspicuous, on elongate pedicels, white with a rose-purple blotch at the base of the perianth segments and two blotches at the base of the lip, or sometimes entirely white with a golden yellow disk. Sepals broadly ovate, the laterals acute, the dorsal sepal usually obtuse with a minute apicule, 2.5—3 cm. long and 1-1.5 cm. wide. Petals a little broader than the sepals, spreading, elliptic- obovate, obtuse or minutely apiculate, about 2.5 cm. long and 1.2-1.6 cm. wide. Lip entire, broadly panduriform, 3—3.5 cm. long and 2.5-4 cm. wide, the apex broadly emarginate and 2-lobed, sometimes with a minute central mucro, the base of the lip with a very short, broad claw which is affixed at right angles to the base of the column, the disk with 3 short, fleshy, obtuse, puberulent keels. Column stout, 4—5 mm. long, the apex dilated, nearly wingless. Costa Rica and Panama. HIRIQUÍ: trail from Paso Ancho to Monte Lirio, upper valley of the Río Chiriquí Viejo, 1500-2000 m., Allen 1508. An attractive highland species, often forming clumps in the tops of the tallest forest trees. Our specimens have pure white flowers, with a yellow disk, and narrower leaves than in the Costa Rican material, but otherwise seem to be identi- cal. Mr. Walter Cope, of Pedro Miguel, reports that Miltonia Roezlii Nichols., a lowland species of the Colombian Choco, has been found in the upper Madden Lake region. 78. ONCIDIUM Swartz ONcIDIUM Sw. in Vet. Akad. Nya Handl. Stockh. 21:239. 1800; Lindl. Folia Orch. Oncidium, (1). 1855; Benth. & Hook. Gen. Pl. 3:562. 1883. ? Phadrosanthus Neck. Elem. 3:153. 1790. Cyrtochilum НВК. Nov. Gen. & Sp. 1:349. 1815. Cyrtochilos Spreng. Syst. Veg. 3:729. 1826. Coppensia Dum. in Mem. Acad. Brux. 9:10. 1835, in nota. Palumbina Rchb. f. in ale Ann. 6:699. 1861. Papiliopsis E. Morr. in Belg. Hortic. 24:261. 1874. Baptistonia Barb. Rodr. Gen. & Sp. Orch. Nov. 1:95. 1877. (514) 1949] FLORA OF PANAMA (Orchidaceae) 179 Erect or pendulous, epiphytic or infrequently terrestrial herbs with short, foliaceous stems, most frequently thickened into conspicuous, more or less ancipi- tous, 1- to 2 (rarely more)-leaved pseudobulbs, the bases enveloped in several papery or foliaceous bracts, or less frequently with the pseudobulbs very short, rudimentary, cylindric, subconic, or complanate-cylindric, enveloped in several papery sheaths, or the plants rarely with many distichously equitant leaves and entirely without pseudobulbs. Leaves subcoriaceous, coriaceous or fleshy, equitant, flat, terete or triangular. Inflorescences usually 1 or 2 short or elongate, often flexuose, erect, arching or laxly pendulous, branching panicles, simple racemes, or 1-flowered scapes produced from the lateral bases of the pseudobulbs, or in the equitant-leaved species from the axils of the leaves. Flowers often large and con- spicuous, usually yellow or brown. Sepals usually subequal, spreading or reflexed, free, or the laterals somewhat connate rarely nearly to the apex, the dorsal sepal rarely much longer and narrower than the laterals. Petals subequal to the dorsal sepal or sometimes larger. Lip usually 3-lobed, often pandurate, rarely nearly entire, the base shortly clawed or sessile and adnate to the base of the column, usually forming a right-angle with it, the lateral lobes porrect, spreading or re- flexed, sometimes obsolete, the central portion of the lip usually with an isthmus, the mid-lobe spreading, usually very broad and transversely dilated, rarely narrow, often emarginate or bifid, the disk usually conspicuously cristate or tuberculate. Column usually short, stout, the lateral margins near the stigma with or without auriculate or petaloid projections, the clinandrium very short and truncate or ovate and obliquely erect, entire or with the apex shortly bidentate, the base foot- less or sometimes apparently with a short foot, rarely produced into a prominent, erect, horn-like process. Anther terminal, operculate, incumbent, strongly convex, semiglobose or cucullate, 1-celled, imperfectly or rarely 2-celled; pollinia 2, usually deeply sulcate, waxy. A very large, polymorphic genus of often attractive, usually epiphytic Ameri- can orchids ranging from Florida and Mexico to the West Indies and South America as far as Peru, Bolivia, and Brazil. Twenty-six species are thus far known from Panama. a. Plants е pseudobulbs, ог the pseudobulbs rudimentary and іп- conspic b. Уа ро fleshy, from the x of a very short subconic or subcylindric stem. Basal shea Mus braces ae у. c. Leaves broadly lanceolate to ellip 6. O. CARTHAGINENSE cc. Leaves terete. Lip more than twice as lon the lateral sepals. Basal callus occupying less than half of Tus total груда. of the va us e. Flowers very small, the lip 1 cm. lon ess, t with 2 separate sate е ра: dd in өч тнт on either side of t ntral b 9. O. EBRACHIATUM ee. Flowers of j size, the li cm. long or more, at the base without кеш ог tubercles оп d cd side of the promi- nent central b 24. O. STIPITATUM dd. Lip less nih twice as long as the lateral icm Ет callus very pr ominent, occupying x; all of the central isthmus...... 25. О. TERES bb. Leaves many, coriaceous or subcoriaceous, the E d. bases (515) [Vor. 36 180 ANNALS OF THE MISSOURI BOTANICAL GARDEN distichously imbricating. @ зе warf, less than tall. d. ves not equitant, а pedir ea bases enveloping a small ро Peduncles filiform, not conspicuously flattened at the 8. O. CRISTA-GALLI e. d. Leaves equitant, the y entirely. without ро Ре- duncles conspicuously M ned, particularly near the арех....... 22. O. PUsILLUM ee. pe robust, more than 3 ta d. of the lip with dd ine nt, wal: developed lateral lobes...... 17. O. ОСНМАТОСНЦ ИМ ‚ос аа. i of the lip нај moe , well-developed lateral lobes.. 19. О. PANDURIFORME aa. Plants with conspicuous pseud . Lip conspicuously exceeding pn і sepals in length. c Ей е = bulbs аек about as broad as long. d. — very distantly inserted on an elongate, slender, rhizome 11. O. GLOBULIFERUM dd. Pseudobulbs approximate on a short rhiz e. Bracts enveloping the base of ај pseudobulb foliaceous. f. Apex of the pseudobulb diphyl m — л O. NEBULOSUM g. Apical leaf of the pm much reduced, conspicu- tha la usly less than the bract blades in length O. CRISTA-GALLI Bg. Apical m of the ая about equal to the bract bla T n length. h. with a х a isthm 16. O. oBRYZATUM hh. Lip with а sharp media consrition, but without a distinct Peleus hiq isth 7. O. CHEIROPHORUM . Bracts рање илы the base of ЊЕ pseudobulb not foliaceous.... 1. O. AMPLIATUM сс. Pseudobulbs ү оа реи со oblong-ovoid, usually тоге than as lon oad. 4 "Pscudobulbs К PMN at the apex. Sepals about 5 mm. 0. O. PARVIFLORUM dd. Preudobuis 2- to 3-leaved at the . Inflorescences with both normal an d abortive sterile flowers....12. О, HETERANTHUM ee. Inflorescences with only normal ike flow f. Sepals obovate-spatulate, broadly obtuse or truncate............16. О. OBRYZATUM ff. Mir lanceolate to elliptic- . acute to subacute. g. Isthmus broad, at least half the width of the mid-lobe... 5. O. CABAGRAE gg. Imus narrow, usually less than 1⁄4 the width of the b -lobe 13. O. IsTHMI bb. Lip TON ін to the lateral sepals or shorter. c. Lateral sepals connate nearly to the apex 26 cc. Lateral sepals free, or only very s hortly connate at the sere sepal and petals elongate, linear-spatulate, very disimilar = дерін ы MD sepals. Flowers solitary, at the apex of a O. WARSCEWICZII 4. O. KRAMERIANUM dd. ES d РК more ог less similar іп shape. ‚ Flor — -— и ап “ hern uous. Ў Flow diameter, the mid-lobe of the cies Ў то yon dunt ал! іп width ff. Flowers less than 3 cm. in diameter, the mid-lobe of the lip conspic nN — O. PowELLII conspicuously exceeding the dorsal sepal in width.......... 4. O. BRACTEATUM ee esa bracts small or slender and inconspicuous. f. Leaves be id va ligular, acute or acuminate. Pseudo- bulbs sually 2- to 3-leaved at the g. Lower portions y" the foliaceous nde not articulated, withou suture gg. Lower dign of the foliaceous bracts articulated, with a distinct suture. O. ENSATUM . Lip with a distinct isthmus, the callus with 2 crenu- late or denticulate, lateral plates at the base................ 3. O. BAUERI hh. Lip without a distinct PUN the callus surmounted by 4 divergent, crenulate keels 18. O. PANAMENSE ff. Leaves ihe dly lanceolate, L1 use to subacute. Pseudo- bulbs usually monophyllous at the apex (516) 1949] FLORA OF PANAMA (Orchidaceae) 181 g. Lip distinctly shorter than the lateral sepals. h. өт БУ С Ме: Пр about equaling the dorsal sepal in of the callus with a broad transverse 21. О. POWELLII hh. t lobe of the lip conspicuously exceeding the dorsal sepal in width. Apex of the callus with 3 fleshy teeth.. 23. O. sTENOTIS r what tdi prominent longitudinal ridges, the condupli icate ова 2. O. ANSIFERUM with rominent longitudinal ridges, the со dpud bos of t b pa hh. Pseudobulbs not exceptionally thin ind di ttened, P of the Rn bracts 5 fibro . O. BAUERI 1. ONCIDIUM AMPLIATUM Lindl. Gen. & Sp. Orch. РІ. 202. 1833. Oncidium Bernoullianum Kránzl. in Engler, Pflazenr. IV, Fam. 50 (Heft 80):231. 1922. Epiphytic herbs with ovoid to suborbicular, strongly compressed, or sometimes angular, pseudobulbs which are often longitudinally ridged and transversely wrinkled, 3-12 cm. tall and 3-9 cm. wide, usually flecked with red or brown, rarely entirely raisin-purple, the apex with 1—3 leaves, the base enveloped in sev- eral papery, imbricating bracts. Leaves elliptic-oblanceolate to ligular, obtuse to subacute, coriaceous, 8—40 cm. long and 3-12 cm. wide, contracted below into short conduplicate petioles. Inflorescences 1 or 2 erect or arching, few- to many- flowered racemes or panicles conspicuously exceeding the leaves in length, produced laterally from the bases of the pseudobulbs. Flowers variable in size but averaging about 2.5 cm. in diameter, bright yellow, nearly white on the reverse surfaces, the sepals spotted with reddish brown, the callus white spotted red. Sepals free, sub- equal, spreading, oblong-spatulate, obtuse, the dilated apices deeply concave, 6—10 mm. long and 3.5-5 mm. wide. Petals much broader than the sepals, clawed at the base, with flat suborbicular blades, 7-11 mm. long and 5-9 mm. wide. Lip very broadly spreading, 3-lobed, 1.5-2 cm. long and 2—3 cm. wide, the lateral lobes relatively small, subauriculate, obtuse, the central portion of the lip with a short constriction, the mid-lobe abruptly dilated, deeply emarginate and 2-lobed, transversely oblong or reniform, the base of the lip contracted into a short claw and adnate to the base of the column, the disk with an erect, fleshy callus, sur- mounted by a transverse biauriculate plate and terminating at the apex in a promi- nent tridenticulate process. Column very short, 3—4 mm. long, the apex with 3 denticulate wings, the base without a foot. Guatemala to Peru, Venezuela, and Trinidad. NE: Chagres, sea ге Fendler 331; near Culebra, 50-100 т., Pittier 3307; rro Colorado Island, Gatún Lake, Woodworth 9 Vestal С 706, Sbattuck 203; Frijoles, Powell 3211; ы Hill, 300 ft., Powell 3200. PANAMÁ near Bejuco, 100 ft., С. Fairchild s. n. cocrÉ: near La pintadas; 150 m., Allen 3542. vERAGUAS: Bahia Honda, Taylor 1512 An attractive lowland species with bright yellow flowers, widely distributed in the American tropics. In Panama they were formerly very frequent in the Chagres River valley, and in general along streams where they were often found high on the trunks and branches of the common Espavé (Anacardium excelsum). The (517) [Vor. 36 182 ANNALS OF THE MISSOURI BOTANICAL GARDEN Š Š 5; fae ‚Г у 5 ҒА ; A 47 “yy | %, À Fig. 199. Oncidium ambliatum (518) 1949] FLORA OF PANAMA (Orchidaceae) 183 larger-flowered forms are often offered for sale under the name of Oncidium ampliatum var. majus. 2. ONCIDIUM ANSIFERUM Rchb. f. in Bot. Zeit. 10:696. 1852. idium ensatum Hort. ex Rchb. f. in Xenia Orch. 1:232. 1858, in synon., non Lindl. Oncidium hieroglyphicum Hort. pro parte ex Rchb. f. x cit. 1858, in synon. Oncidium Lankesteri Ames, in Sched. Orch. 4:53. 192 ET naranjense Schltr. in Fedde Rep. Sp. Nov. ER 19:259. 1923. Epiphytic herbs with very strongly compressed, exceptionally thin, ovate- elliptic or oblong-elliptic, usually smooth pseudobulbs 6—10 cm. tall and 3—6 cm. wide, the apices with 1 or infrequently 2 leaves, the bases with several distichously imbricating bracts the uppermost 2-3 of which are conspicuously foliaceous. Leaves lanceolate to elliptic-lanceolate, subacute to obtuse, pergameneous, 15—33 cm. long and 2.5—5.5 cm. wide. Inflorescences 1 or 2 erect or arching panicles, usually much exceeding the leaves, up to about 1 m. in length, produced from the lateral bases of the pseudobulbs. Flowers of moderate size, averaging about 3 cm. in diameter, the sepals and petals brown, sometimes with yellow margins and apices, the mid-lobe and lateral lobes of the lip bright yellow, the central isthmus brown with a bright yellow crest. Sepals subequal, free, spreading, elliptic-lanceolate, obtuse to acute, with strongly undulate margins, 1.2-1.7 cm. long and 0.4-0.6 cm. wide. Petals somewhat broader than the sepals, spreading, elliptic-oblong, obtuse to subacute, with undulate margins, 1.4-1.6 cm. long and 0.5—0.65 cm. wide. Lip pandurate, 3-lobed, 1.4—1.6 cm. long and 1-1.5 cm. broad at the trans- versely reniform, emarginate apex, the mid-section of the lip abruptly contracted in front into a broad isthmus, the basal portions more or less confluent with the short, suborbicular, lateral lobes, the disk with a 5-lobed puberulous crest, termi- nating in a central porrect tooth. Column short, 4—6 mm. long, the apex with dolabriform wings. Costa Rica and Panama. CHIRIQUÍ: Río Chiriquí Viejo, vicinity Paso Ancho, 5000 ft., Н. Dunn s. n.; Llano del D western slopes of Chiriqui Volcano and along Río Chiriqui Viejo, 1200 m., Allen An attractive highland species, readily distinguished from other Panama On- cidiums by the broad, smooth, exceptionally thin pseudobulbs. 3. ONCIDIUM Bauert Lindl. in Bauer & Lindl. Ill. Orch. Gen. Ё. 7. 1830-38. Oncidium altissimum var. B Lindl. Gen. & Spec. Orch. AA 200. 1833. : eg. n 5%. Oncidium bolycladium Rchb. f. ex Lindl. Fol. Orch. Oncidium, (47). 1855. В 1892. Erect, robust, epiphytic herbs with approximate, oblong-ovoid, strongly ridged pseudobulbs up to about 18 cm. tall and 4 cm. wide, the apices with 1 or 2 leaves, the lower portions enveloped in the coarsely fibrous, conduplicate, imbricating (519) [Vor. 36 184 ANNALS OF THE MISSOURI BOTANICAL GARDEN bases of several conspicuously foliaceous bracts. Leaves and bract blades linear- ligular, acute, subcoriaceous, up to about 75 cm. long and 2.5-4 cm. wide. In- florescences 1 or 2 stout, erect, arching or pendulous, many-flowered panicles up to about 3 m. in length, produced from the lateral bases of the pseudobulbs. Flowers of moderate size, 2.5—3 cm. in diameter. Sepals free, subequal, spreading or reflexed, shortly clawed at the base, yellow marked with brown, the dorsal sepal elliptic-lanceolate, acute, 1-1.5 cm. long and 0.4-0.5 cm. wide, the lateral sepals obliquely linear-lanceolate, acute or shortly acuminate, the dorsal surface with a distinct central keel, 1.2-1.7 ст. long and 0.35-0.45 cm. wide. Petals subequal to the dorsal sepal and similarly colored, elliptic-lanceolate, acute, with undulate margins, 1—1.4 cm. long and about 0.5 cm. wide. Lip pandurate, 3-lobed, yellow with a reddish brown central blotch, 1.2-1.5 cm. long and 0.9—1.2 cm. wide, the lateral lobes small, suborbicular to subquadrate, obtuse, the central portion of the lip contracted into a distinct isthmus, the mid-lobe abruptly dilated, emarginate and bilobed, transversely reniform in outline, the disk with an erect, fleshy callus, at the base with 2 lateral crenulate or denticulate wings or plates, the apex with 3 short, fleshy teeth. Column about 5 mm. long, with the lateral wings often bifid, the lower lobules obtuse and spreading, the apices acute or acuminate, often more or less converging. Mexico to Brazil and Peru; Virgin Islands and Martinique. N : without definite locality, Pacific coast lowlands, Fairchild s.m. CHIRIQUÍ: without definite locality, von Warscewicz s. n. (fide Reichenbach). The species of this type form a closely allied, very perplexing association in- cluding Oncidium altissimum Sw., О. Baueri Lindl., О. sphacelatum Lindl., О. polycladium Rchb. f., O. stenotis Rchb. f., О. isthmi Schltr., and O. panamense Schltr. Oncidium stenotis and О. isthmi can be separated easily, and although О. sphacelatum and О. panamense аге very nearly alike, they differ from the other species in the broad basal half of the lip, without a distinct isthmus. Тһе most troublesome series involves the remaining species, which have a broad but distinct isthmus to the lip, a transversely reniform mid-lobe, and variously developed multi- denticulate basal calli. These are extremely difficult to distinguish from one another, at least on the basis of herbarium material, since most of the described differences consist more nearly in the degree of development of essentially identical parts, rather than in any basic structural deviation. In particular, there seem to be no consistent characters upon which to base the further segregation of О. polycladium from O. Baueri, the former concept being here reduced to synonymy, thus ex- tending the range of the widespread South American and West Indian О. Baueri as far north as Mexico. In our material, the latter species is separable from O. panamense by the distinct isthmus of the lip; from О. isthmi by the broader isthmus and narrower, shorter mid-lobe; and from О. stenotis by the smaller flowers, in which the lip is about equal to the lateral sepals in length. It seems quite probable that further study may reduce some of these, and other closely allied concepts, to varietal status or synonymy. (520) 1949] FLORA OF PANAMA (Orchidaceae) 185 4. ONCIDIUM BRACTEATUM Warsc. & Rchb. f. in Bot. Zeit. 10:695. 1852. Erect, epiphytic herbs with slender brown, compressed pseudobulbs, more or less linear in outline but usually somewhat broader at the base and tapering grad- ually to a narrower apex, 6—12 cm. tall and 1.5—3 cm. wide at the base, terminating above in 1 or 2 leaves, the base enveloped in several distichously imbricating bracts, the uppermost 2 or 3 of which are conspicuously foliaceous. Leaves ligular, obtuse, coriaceous, 15-45 cm. long and 2-3.2 cm. wide, contracted below into elongate, slender, conduplicate petioles. Inflorescences 1 or 2 erect or arching panicles much exceeding the leaves, up to about 1 m. in length, the lateral branches short, usually 3-flowered, provided with several conspicuous, elongate, acute, pale brown, spath- aceous bracts. Flowers of moderate size, the sepals and petals greenish yellow or yellow, heavily blotched and spotted dark brown or maroon, the apical and lateral lobes of the lip bright yellow, the central isthmus brown or maroon. Sepals sub- equal, free, spreading, with undulate margins, the dorsal sepal oblong-lanceolate, acute, 10-15 mm. long and 3.5—5 mm. wide, the lateral sepals linear-lanceolate, obliquely acute, with a distinct central keel on the dorsal surface, 12-17 mm. long and 2.5—4.5 mm. wide. Petals subequal to the dorsal sepal, elliptic-lanceolate, acute, with undulate margins, 10—15 mm. long and 4—6 mm. wide. Lip panduri- form, with a broad central isthmus, 12-15 mm. long and 10—12 mm. wide at the 2-lobed emarginate apex, the base with 2 small, suborbicular, lateral auricles, the disk with an erect, more or less triangular, multidenticulate callus. Column 5-6 mm. long, with narrow lateral wings, the base without a foot. Costa Rica and Panama. Although absent from recent collections in our area, the species was originally described from plants presumably found in Panama by von Warscewicz, the data being given as “Chiriqui Cordilleren 6—9000 ft. auf Bäumen.” They are readily distinguished by the narrow brown pseudobulbs and the very conspicuous, pale brown, spathaceous floral bracts. 5. ONciDIUM САВАСВАЕ Schltr. in Fedde Rep. Sp. Nov. 9:292. 1911. Oncidium Rechingerianum Kránzl. in Engler, Pflanzenr. IV, Fam. 50 (Heft 80):202, fig. 18, C:a-c. 1922 Erect, epiphytic herbs with slender, elliptic-ovoid to nearly linear, ancipitous pseudobulbs usually 6—11 cm. tall and 2-3 cm. wide, densely spotted dark brown or black, often becoming longitudinally ridged with age, the apex with 2 or 3 leaves, the lower portions enveloped in the conduplicate bases of several distichous- ly imbricating bracts, the uppermost 2 or 3 of which are conspicuously foliaceous. Leaves ligular, acute, subcoriaceous, 15-25 cm. long and 1.5-3 cm. wide, con- tracted below into slender, conduplicate petioles. Inflorescences usually solitary, erect or arching, many-flowered panicles much exceeding the leaves, up to about 80 cm. in length, produced from the lateral bases of the pseudobulbs. Flowers of moderate size, the sepals and petals heavily blotched rich chestnut-brown, usually (521) [Vor. 36 186 ANNALS OF THE MISSOURI BOTANICAL GARDEN with yellow margins and apices, the mid-lobe and lateral lobules of the lip bright yellow with the central isthmus a rich reddish brown, the callus white spotted with brown. Sepals subequal, free, usually reflexed, with recurved apices, the dorsal sepal elliptic-obovate, acute, usually 8-12 mm. long and 4—6 mm. wide, the laterals linear-lanceolate to rather obliquely ligular, subacute to acute, with a distinct dorsal keel, usually 8—12 mm. long and 3—4 mm. wide. Petals somewhat broader than the sepals, slightly reflexed, with rather undulate margins, elliptic-oblanceo- late, obtuse or acute, usually 10—12 mm. long and 5—6 mm. wide. Lip pandurate, usually 12—16 mm. long and 10-14 mm. wide at the 2-lobed, emarginate apex, the central portion abruptly contracted into a broad isthmus, the base with 2 small, lateral, obtuse, subquadrate auricles, the disk with an erect, fleshy, truncate keel, the upper margins with 2 pairs of fleshy teeth, the basal pair larger than the apical pair. Column short, erect, about 5 mm. long, with broad, lateral, spreading, rather bilobed or minutely dentate wings, the base of the column without a foot. Costa Rica and Panama. ОСТЕ: region north of El Valle de Antón, 1000 m., Allen 2361, 3681, 2/70; moun- tains beyond La Pintada, 400—600 m., Hunter & Allen 580. снікюоі: without definite locality, 4000—5000 ft., Powell 161, 177; vicinity Palo Alto, 4000 ft., Powell. p Llano del Volcán, 1000 m., Kieswetter s. n. An attractive species of the Coclé and Chiriqui highlands, usually found growing in the tops of tall trees. 6. ONCIDIUM CARTHAGINENSE (Jacq.) Sw. in К. Vet. Akad. Stockh. Nya Handl. 21:240. 1800. Epidendrum cartbaginense Jacq. Select. Stirp. Amer. 228, 7. 133, fig. 4. 1763. Epidendrum undulatum Sims, in Bot. Ma 7. 1804. Oncidium undulatum Salisb. in Trans. Roy. Hort. Soc. 1:295. 1812, non Lindl. Oncidium panduriferum Kunth, in HBK. ae tuer К Spec. 1:346, 7. 82. 1815. Oncidium Oerstedii Rchb. f. in Bonplandia 2: Oncidium сеи" (Таса.) Sw. var. eae (Kaw: f.) Lindl. Fol. Orch. Oncid. (40). 18 Oncidium cartbaginense (Jacq.) Sw. var. Swartzii Lindl. loc. cit. 1855. Oncidium obsoletum Rich. et Gal. ex Lindl. loc. cit. 41. 1855. Oncidium ERU Kránzl. in Engler, Pflanzenr. IV, Fam, 50 (Heft 80):112. 1922. Oncidium Oerstedii Rchb. f. var. crispiflorum Schltr. in Fedde, Rep. Sp. Nov. Beih. 17:85 Robust, epiphytic herbs with very short, subquadrate, compressed or sub- cylindric, nearly obsolete pseudobulbs 1-2 cm. long and 1-1.5 cm. broad, the truncate apex with a single persistent, broadly lanceolate to elliptic-oblong, acute, usually fleshy leaf, which usually is more or less spotted reddish brown, 15—60 cm. long and 3.5-8 cm. wide, the lower surface with a strongly developed central keel, the pseudobulbs and the bases of the new leaves enveloped in 3—4 papery, closely imbricating bracts. Inflorescences elongate, erect, usually solitary, arching or sometimes pendulous, many flowered panicles up to 1.5 m. in length, produced from the bases of the pseudobulbs. Flowers variable in size and color, averaging (522) 1949] FLORA OF PANAMA (Orchidaceae) 187 N IN i % 1 МР: WER N y ` ў \ ыл e 9. We N N , 55 : N N SA ENa ) N № у; © i ap (е; c ШІ А 5 Ñ | | «ЧА y ; “ЖА NY | AN Fig. 200. Oncidium carthaginense (523) [Vor. 36 188 ANNALS OF THE MISSOURI BOTANICAL GARDEN about 2 cm. in diameter, usually more or less heavily blotched and spotted with purplish rose on a white base. Sepals of about equal length, spreading, or with the laterals somewhat reflexed, distinctly clawed at the base, the blades with strongly undulate margins, the dorsal sepal free, the claw narrowly linear, the blade abruptly dilated and suborbicular, somewhat concave, 8-11 mm. long and 5-7 mm. wide, the lateral sepals very shortly connate at the base, the blades obovate- spatulate, obtuse to acute, 7-12 mm. long and 4—6 mm. wide. Petals distinctly clawed at the base, spreading, with undulate, often crisped margins, the blades oblong to elliptic-oblong, obtuse, 8-12 mm. long and 5-7 mm. wide. Lip pan- durate in outline, 3-lobed, abruptly contracted at the base and adnate to the base of the column, 8-12 mm. long and 6-10 mm. wide, the mid-lobe transversely sub- reniform, the anterior margin entire or sometimes shallowly emarginate, the cen- tral portion of the lip with a median constriction often prolonged into a short, broad isthmus confluent at the base with the oblong, obtuse, or triangular, rather auriculate lateral lobes, which are often somewhat reflexed, the disk with a promi- nent, more or less 4-parted, fleshy, erect, tuberculate crest. Column 2-3 mm. long, with large, spreading, unequally 2-lobed, lateral wings. Florida and the West Indies; Mexico to Venezuela and Brazil. CANAL ZONE: en Fort Kobbe, on margins of mangrove йр a sea level, Allen 2754. PANAMÁ: margin of the Pacific Ocean, northwest of Panama City, on slope of hill rising from the sea, Powell I. CHIRIQUÍ: near city of David, sea level, Powell 270. A rather variable species closely allied to Oncidium guttatum (L.) Rchb. f., widely distributed in the lowlands of the American tropics. Most of the available specimens from Panama would correspond fairly well to the concept of О. cartba- ginense var. Oerstedii, yet the supposed differences (shorter lip, longer isthmus, and brighter coloring) are found in an extensive series of specimens from the entire geographic range to merge imperceptibly with the type, leaving no clear-cut grounds for separation. 7. ONCIDIUM CHEIROPHORUM Rchb. f. in Bot. Zeit. 10:695. 1852. Oncidium ii Mero Kránzl. in Engler, Pflanzenr. IV, Fam. 50 (Heft 80):197, Z. 17, fig. F; 922. Dwarf, caespitose, epiphytic herbs averaging 12—15 cm. in height, with ovoid to suborbicular, compressed, monophyllous or rarely diphyllous pseudobulbs 1.5—3 cm. long and 1.4—2.5 cm. wide, at first smooth, often becoming finely wrinkled with age, the bases enveloped in several imbricating bracts the uppermost 2 or 3 of which are conspicuously foliaceous. Leaves linear-ligular, obtuse to subacute, pergameneous, 5-15 cm. long and 0.8-1.4 cm. wide, contracted below into short, conduplicate petioles. Inflorescences 1 or 2 slender, erect or arching, densely paniculate scapes usually exceeding the leaves, up to about 30 cm. in length, pro- duced from the lateral bases of the pseudobulbs. Flowers small, averaging 1—1.5 cm. in diameter, bright yellow and fragrant. Sepals of about equal length, the dorsal sepal erect, shortly clawed at the base, the orbicular blade deeply concave, (524) 1949] FLORA OF PANAMA (Orchidaceae) 189 often with а minute арісше, about 5 mm. long and 4 mm. wide, the lateral sepals elliptic-obovate, obtuse, often reflexed, apparently very shortly connate at the base, 5-6 mm. long and 3—3.5 mm. wide. Petals shortly clawed, spreading, obo- vate-spatulate, obtuse, about 4—5 mm. long and 4 mm. wide. Lip conspicuously 3-lobed, averaging about 8 mm. long and 12 mm. wide, the sessile base adnate to the base of the column, the broadly spreading, auriculate, lateral lobes subquadrate to suborbicular, usually with reflexed margins, the central portion of the lip with an abrupt, narrow constriction, the mid-lobe transversely subreniform or 2-lobed, the lobules often porrect in natural position, the disk with a prominent, fleshy keel, the upper margins with 2 conspicuous subquadrate wings, between which at the apex there is a short membranaceous hood or concavity. Column very short, with large obovate, usually porrect, lateral wings, the base produced in front into a prominent, erect, fleshy horn. Anther with a long beak, otherwise typical of the genus. Costa Rica, Panama, and Colombia. cocrÉ: El Valle de Antón, 500-1000 m., D. Allen 3987, Hunter & Allen 350, Allen 236, 1681, 2014, 2030, 4211. cHirigui: without definite locality, 5000 ft., Powell 54; Volcancitos, 5500 ft., Davidson 1261; Piedra de Lino, 1600 m., Killip 3571; valley of the upper Rio Chiriqui Viejo, near Monte Lirio, 1300-1900 m., Seibert 224; trail from Paso Ancho to Monte Lirio, Rio Chiriqui Viejo, 1500-2000 m., Allen 1499. A small-flowered, dwarf, but attractive species very common in the highlands of Coclé and Chiriqui provinces in our area, where the plants often form dense clumps in the tops of tall trees or on the ends of spreading branches, often fully exposed to the sun. 8. ONCIDIUM CRISTA-GALLI Rchb. f. in Bot. Zeit. 10:697. 1852. Oncidium iridifolium Lindl. in Bot. Reg. 22: f. 1011. 1836, non НВК. Oncidium decipiens Lindl. Folia Orch. Oncidium, (22). 1855. Dwarf, caespitose, epiphytic herbs less than 10 cm. tall, with small, ovoid, compressed pseudobulbs 10—20 mm. tall and 8-18 mm. wide, the apices with a single very short, conduplicate, ensiform, abortive leaf sometimes reduced to a very abbreviated foliaceous apicule, 2—15 mm. long and 1—2 mm. wide, the bases of the pseudobulbs almost completely enveloped by the distichously imbricating, conduplicate bases of 4 to 6 conspicuous, foliaceous bracts, the blades ligular, acute, pergameneous, 2—8 cm. long and 0.5—1.0 cm. wide, usually explanate but some- times conduplicate and ensiform. Inflorescences 1 to 4 erect or arching, filiform, 1-flowered scapes about equaling the leaves in length, produced from the axils of the foliaceous bracts, the peduncles provided with 3 or 4 distant, perfoliate, acumi- nate, spathaceous bracts 6—10 mm. long and about 3 mm. broad. Flowers very large in relation to the size of the plant, 2—3 cm. long and 1.8-2.2 cm. wide. Sepals free, spreading, the dorsal sepal erect, elliptic-ovate, slightly concave with a dorsal keel, terminating in a short apicule, the blade greenish yellow, 5—6 mm. (525) [Vor. 36 190 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 201. Oncidium crista-galli long and 3—4 mm. wide, the lateral sepals very inconspicuous, appressed to the back of the lip, rather obliquely linear-ligular, acuminate, greenish yellow, 5—6 mm. long and 2-2.5 mm. wide. Petals broadly spreading, oblong, rather obliquely obtuse and apiculate, bright yellow with transverse bands of reddish brown, the margins often undulate, 8-10 mm. long and 4—5 mm. wide. Lip complexly 3- lobed, 1.5-2.2 cm. long and 1.7-2.2 cm. wide, bright yellow, the margins undulate, the base shortly contracted and adnate to the base of the column, the lateral lobes spreading, obovate-spatulate, the dilated suborbicular blades 7-8 mm. long and 6-8 mm. wide, the mid-lobe much broader at the base than at the apex, divided into 4 lobules, the lateral basal pair rounded and spreading, the apical pair project- ing, obliquely cuneate, separated by a deep central sinus, the disk convex, white with reddish brown blotches, with a prominent, spreading, fleshy, suborbicular plate and ruffled and crisped margins, subtended on each side by semicircular, crisped extensions, the apex terminating in a short, complex, crisped, fleshy hood. Column very short, about 3 mm. long, with conspicuous, lateral, dolabriform wings. Mexico to Colombia and Peru. COCLÉ: mountains beyond La Pintada, 400—600 m., Hunter 9 Allen 588; El Valle de Antón, 600-1000 m., D. Allen 5158, Allen & Allen 1251, 1675, 2883, 3800. A remarkable, attractive, dwarf species of the wet highland forests of Coclé Province, with relatively large, bright yellow flowers, the plants superficially reminiscent of those of Oncidium pusillum, but readily distinguished by the presence of small suborbicular pseudobulbs and the filiform, rather than com- planate, scapes. (526) 1949] FLORA ОЕ PANAMA (Orchidaceae) 191 9. ONCIDIUM EBRACHIATUM А. & S. in Sched. Orch. 8:75, fig. 6. 1925. Pendulous, epiphytic herbs with fleshy, terete, longitudinally sulcate, acuminate leaves up to about 60 cm. in length, the plants identical in appearance with those of Oncidium stipitatum. Inflorescences usually solitary, arching, many-flowered panicles, produced from the bases of the plants. Flowers very small for the genus, about 1 cm. in length, the sepals and petals spotted with reddish brown, the lip yellow on both surfaces. Sepals free, spreading, the dorsal вера! suborbicular, strongly concave, incurving over the column, with an obscure, terminal apicule, the blades 3—3.5 mm. long and about 2.5 mm. wide, the lateral sepals obovate- spatulate, obtuse, minutely apiculate, the blades somewhat concave and incurving, about 4 mm. long and 2-2.2 mm. wide. Petals widely spreading, rather obliquely elliptic-oblong, obtuse, about 4 mm. long and 2-2.1 mm. wide. Lip pandurate, 3-lobed, about 8 mm. long and 6 mm. wide, the sessile base adnate to the base of the column, the lateral lobes subfalcate, obtuse, spreading, or with the apices somewhat antrorse in natural position, the lower margins confluent with the nar- row central isthmus, the mid-lobe dilated and emarginate, flabellate-reniform in outline, often porrect in natural position, the minutely papillose disk with a broad, fleshy, flat, lunate, porrect plate below the column, terminating below the apex in a central, subtrilobed tubercle, on each side of which are lightly converging keels, terminating in low tuberculate swellings. Column very short and stout, about 1 mm. long, without prominent lateral wings. Panama. АМА: without definite locality, M. D. Hunter s. n.; coastal swamps east of Panama ity, da ina een the Jagua Hunt Club and Congor Hill, sea level, Hunter & Allen 471; San jos Island, Perlas Archipelago, Johnston 1340. DARIEN: Cana and vicinity, 2000-6500 ‚ К. 5. Williams 975; Rio Tuira, near Pinogana, 100 ft., Allen 4300. A small-flowered, terete-leaved species very closely allied to O. nudum Lindl. of South America; apparently confined to the lowlands of the eastern half of Panama. 10. ONCIDIUM ENSATUM Lindl. in Bot. Reg. n. 5. 5: Misc. 17. 1842. Oncidium cerebriferum Rchb. f. in Bot. Zeit. 10:696. 1852. Oncidium confusum Rchb. f. in Xenia Orch. 1:234. 1858. Erect, terrestrial herbs with fleshy, ovoid, somewhat compressed, usually smooth pseudobulbs 5.5—8 cm. tall and 3.5—5 cm. wide, the truncate apices with 1 to 3 leaves, the bases enveloped in 4—6 imbricating bracts the uppermost 2 or 3 of which are conspicuously foliaceous, the blades persistent, not articulated and not separating by a suture near the base. Leaves and bract blades linear-lanceolate, acuminate, subcoriaceous, 45—90 cm. long and 2—3 cm. wide, rather rigidly erect, with a strong central keel, the margins somewhat conduplicate, particularly for the lower half. Inflorescences 1 or 2 erect or arching, many-flowered panicles equaling or exceeding the leaves, produced from the lateral bases of the pseudo- (527) [Vor. 36 192 ANNALS OF THE MISSOURI BOTANICAL GARDEN bulbs. Flowers of moderate size, in fresh material averaging about 3 cm. in diameter when spread out. Sepals subequal, free, spreading or reflexed, greenish or brownish olive, usually with undulant margins, the dorsal sepal oblanceolate, acute, 10—12 mm. long and 4—5 mm. wide, the lateral sepals lanceolate, acuminate, 13-15 mm. long and 3.5-4 mm. wide, with a pronounced dorsal keel. Petals subequal to the dorsal sepal and similarly colored, spreading or reflexed, with undulant mar- gins, obliquely oblanceolate, acute, 12-14 mm. long and 4-5 mm. wide. Lip panduriform, 3-lobed, bright yellow, about 1.5 cm. long and 1.5 cm. wide, ab- ruptly contracted at the base and adnate to the base of the column, the small lat- eral lobes oblong, obtuse or obscurely auriculate, the anterior margins confluent with the margins of the broad central isthmus, the mid-lobe dilated, 2-lobed, trans- versely subreniform, the disk olive-green, with a prominent, fleshy, white, 7- toothed callus. Column short, with prominent, lateral wings. British Honduras to Panama. ANAMÁ: foothills east of Panama City, sea level, Powell 267; vicinity Juan Díaz, Powell 3484; in low thick scrub, along Río Tecámen, east of Panama City, about 30 m., Hunter 9 Allen 266, Allen 5150. A fairly common terrestrial species of the Pacific slope, often found growing in low scrub in association with Peristeria elata, or sometimes in open grassland. In herbarium specimens, the species considerably resembles Oncidium panamense, but may be distinguished readily by the narrower basal callus, the terrestrial habit, and in the complete lack of articulations near the bases of the foliaceous bracts. 11. ONcIDIUM GLOBULIFERUM HBK. Nov. Gen. & Sp. 1:347. 1815. Oncidium scansor Rchb. f. in Linnaea 22:844. 1849. Oncidium convolvulaceum Lindl. in Paxt. Flow. Gard. 1:102, sub /. 21. 1850-51. Oncidium globuliferum HBK. var. кеу рин Rchb. f. in Сағд, Chron. 1678. 1871. Oncidium Wercklei Schltr. in Fedde Rep. Sp. Nov. Beih. 19:68: 1923. Epiphytic herbs with solitary or clustered, elliptic-oblong to suborbicular, compressed, monophyllous pseudobulbs 1.2-2.5 cm. long and 1-2 cm. wide, very distantly distributed along the slender, flexuose, scandent rhizomes, the bases of the pseudobulbs partially enveloped in the conduplicate bases of several distichously imbricating bracts the uppermost pair of which being conspicuously foliaceous. Leaves elliptic-oblong to elliptic-lanceolate, obtuse to subacute, coriaceous, 2.5—6 ст. long and 1—2 cm. wide, contracted below into very short, conduplicate petioles. Inflorescences 1 or 2 short, erect, 1-flowered scapes about equaling the leaves in length, produced from the lateral bases of the pseudobulbs. Flowers very large in relation to the size of the plant, 2.5-3 cm. long and 2-2.5 cm. wide. Sepals free, spreading, yellow barred with reddish brown, the dorsal sepal elliptic- oblong, shortly acute, 8—12 mm. long and 4—5 mm. wide, the lateral sepals rather obliquely lanceolate, acute, 10—12 mm. long and 3—4 mm. wide. Petals somewhat broader than the sepals, yellow with reddish brown bars, elliptic-oblong, obtuse to subacute, 10-12 mm. long and 5-6 mm. wide. Lip 3-lobed, broadly spreading, (528) 1949] FLORA OF PANAMA (Orchidaceae) 193 Бір. 202. Oncidium globuliferum (529) [Vor. 36 194 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1.7-2 cm. long and 2—2.5 cm. wide, abruptly contracted at the base and adnate to the base of the column, the small subquadrate, auriculate, obtuse, lateral lobes spreading, the anterior margins confluent with the margins of the short, broad, central isthmus, the mid-lobe spreading, emarginate, 2-lobed, broadly and trans- versely reniform, bright yellow, about 25 the total length of the lip, the disk with a more or less triangular, 7-toothed, fleshy callus. Column short, about 5 mm. long, with a pair of spreading lateral wings. Costa Rica to Venezuela and Peru. жұла region north of El Valle де en 1000 m., Allen 5201. cHiRIQUÍ: vicinity Bajo Mono and Quebrada Chiquero, 1500 m., Woodson & Schery 572; without definite чаи 5000 ft., Kieswetter s.n.; 5. Cerro Punta, headwaters of the Río Chiriqui Viejo, 2000 m., Allen 1567. Unique among the Oncidiums of Panama in having elongate, slender, flexuose rhizomes, the plants often forming large tangled colonies in the tops of tall trees in the wet highland forests. They are seldom seen except as fragments which have fallen from above, usually being rendered conspicuous by the yellowish rhizomes which superficially resemble the stems of a dodder. The solitary flowers are bright yellow, very large in relation to the size of the plant, often exceeding the pseudobulbs in diameter. 12. ONCIDIUM HETERANTHUM Poepp. & Endl. Nov. Gen. & Sp. 1:34, 2. бо. 1836. Oncidium bryolopbotum Rchb. £. in Gard. Chron. 738. 1871. Oncidium ionops Cogn. & Rolfe in Jour. des Orchid. 3:346. 1892. Oncidium megalous Schltr. in Fedde Rep. Sp. Nov. 9:30. 1910. Erect, caespitose, epiphytic herbs with linear or s ovoid, rather com- pressed pseudobulbs up to about 5 cm. tall and 2 cm. wide, the truncate apices with 1 or 2 leaves, the bases enveloped in several imbricating bracts the uppermost 2 of which are conspicuously foliaceous. Leaves narrowly linear to elliptic-lanceo- late, acute, subcoriaceous, 4—18 cm. long and 0.8—3.5 cm. wide, contracted below into very short conduplicate petioles. Inflorescences 1 or 2 slender, erect or arching panicles usually much exceeding the leaves, up to about 75 cm. long. Flowers dimorphic, each lateral branch of the inflorescences terminating in a single normal flower averaging about 2 cm. long and 1.2 cm. wide, subtended by many very dissimilar, small, stelliform, abortive, sterile flowers. Perianth segments of the sterile flowers broadly spreading, narrowly linear to spatulate, yellowish, 3—4 mm. long, the lip generally entirely aborted. Normal flowers with the sepals free, elliptic-lanceolate to ligular, shortly acute to obtuse, the bases shortly clawed, the blades often with undulate margins, yellow blotched with reddish brown, 6—10 mm. long and 2-3 mm. wide. Petals elliptic-oblong to obliquely obovate-spatu- late, obtuse to subacute, clawed at the base, yellow blotched with reddish brown, 7-10 mm. long and 2.5—3.5 mm. wide. Lip broadly panduriform, 10—12 mm. long and 10-15 mm. wide, subequally 4-lobed, the lateral lobes broadly triangular, (530) 1949] FLORA OF PANAMA (Orchidaceae) 195 obtuse, or rounded, the anterior margins converging and forming a short isthmus or narrow, median constriction above the abruptly dilated, spreading, transversely reniform, 2-parted, bright yellow mid-lobe, the disk with an erect, fleshy, tubercu- late callus terminating at the apex in 3 short teeth. Column short, 3—4 mm. long, with 2 very prominent lateral wings. Costa Rica, Panama, Colombia, Peru, Bolivia, and probably adjacent territories. CHIRIQUÍ: without definite locality, “On large trees, in dense shade”, 5000 ft., Kies- wetter s. n., Powell 48; central valley of the Río Chiriquí Viejo, vicinity of New Switzer- land, 1800—2000 m., Allen 1 A curious highland species distinguished by the abundant yellowish, stelliform, abortive florets, subtending the normal flowers on the branches of the paniculate inflorescences. 13. ONCIDIUM ISTHMI Schltr. in Fedde Rep. Sp. Nov. Beih. 17:84. 1922. Robust, erect, epiphytic herbs with approximate, narrowly ovoid to linear- oblong, compressed, longitudinally ridged, usually diphyllous pseudobulbs 8.5—12 cm. long and 3—4 cm. wide, the lower portions enveloped in the conduplicate, dis- tichously imbricating bases of 6—7 bracts, the uppermost 4—5 of which are conspic- uously foliaceous. Leaves and bract blades linear-lanceolate, acute, subcoriaceous, 15—45 cm. long and 1.5-3.2 cm. wide, the foliaceous bracts articulated near the base, with a distinct suture, the blades ultimately deciduous. Inflorescences 1 or 2 erect or arching, many-flowered panicles up to about 1 m. in length, produced from the lateral bases of the pseudobulbs. Flowers of moderate size, 2.5-3 cm. long and 1.5+2.2 cm. wide. Sepals free, spreading, clawed at the base, yellow barred with brown, the dorsal sepal elliptic-oblong, acute, with undulate margins, 9—12 mm. long and 4—5 mm. wide, the lateral sepals obliquely oblong, subacute, with un- dulate margins, 10-13 mm. long and 4—5 mm. wide. Petals rather obliquely ob- long, subacute, with undulate margins, yellow barred with brown, 10—12 mm. long and 4—5 mm. wide. Lip panduriform, 3-lobed, bright yellow, the small auriculate, lateral lobes subquadrate to suborbicular, the frontal margins abruptly contracted into a narrow central isthmus, the mid-lobe broadly dilated and 2- lobed, transversely reniform, occupying about 24 of the total length of the lip, the disk brown, with an erect, compact, 7-toothed basal callus. Column short, about 4 mm. long, with prominent lateral wings. Costa Rica and Panama. CANAL ZONE: San Juan, upper Chagres River, sea level, Powell 307. PANAMA: with- out definite locality, sea level, Fairchild s.m.; vicinity Pacora, sea T Allen 3442. DARIEN: Río Sambu, 200 m., Pittier 5569. Robust lowland epiphytes, vegetatively nearly identical with O. panamense and O. Baueri but readily distinguished by the larger, conspicuously expanded mid- lobe and the much narrower isthmus of the lip. (531) [Vor. 36 196 ANNALS OF THE MISSOURI BOTANICAL GARDEN 14. ONciDIUM KRAMERIANUM Rchb. f. in Otto & Dietr. Allgem. Gartenzeit. 2319. 1855. Oncidium Papilio var. Kramerianum Lindl. Folia Orch. Oncidium, к. 1855. Oncidium nodosum Ed. Morren іп Belg. Hortic. 24:258. 1874, in a Oncidium papilioniforme Regel, in Act. Hort, Petrop. 6:292. Oncidium Kramerianum var. resplendens Rchb. f. in Gard. Chron. 3: 360. 1888. Erect, caespitose, epiphytic herbs with suborbicular to subquadrate, truncate, compressed, more or less rugose, monophyllous pseudobulbs 2.5—4 cm. in diameter, the bases enveloped in several papery, imbricating bracts which soon weather away. Leaves elliptic-oblong, acute, persistent, rigidly coriaceous, 15-25 cm. long and 3.5—6.5 cm. wide, contracted below into very short, conduplicate petioles, the lower surfaces of the leaves often more or less suffused with purple and with a strong central keel, the upper surfaces deep green, sometimes mottled darker green or purple. Inflorescences 1 or 2 erect or arching, prominently jointed scapes up to about 75 cm. in length, produced from the lateral bases of the pseudobulbs, the nodes conspicuously swollen, each provided with an elongate, acute, tubular, papery sheath clasping the lower halves of the cylindric internodes, the apex of the scape with a subconic, acute, terminal "bud" made up of several very closely imbricating sheaths, representing a much condensed raceme, the 5-8 flowers pro- duced singly in succession from the axils of the sheaths, each flower lasting about a week. After the initially produced raceme has been exhausted, the scapes fre- quently elongate from one of the upper nodes, and again produce a series of flowers so that any given scape may remain in flower for as much as a year. Flowers large and richly colored. Sepals free, the dorsal sepal and the petals similar, erect, nar- rowly linear-spatulate, acute, rich reddish brown, the upper portions with un- dulate margins, 5.5-8 cm. long and 0.5—0.7 cm. wide, the lateral sepals very dissimilar, narrowly clawed at the base, the blades abruptly dilated above the claw, obliquely elliptic-ovate, obtuse to subacute, subfalcately deflexed with strongly undulate margins, yellow heavily blotched with rich reddish brown, 3—5 cm. long and 1—2.5 cm. wide. Lip about equaling the lateral sepals in length, subpandurate, 3-lobed, 3-4.5 cm. long and 3-4.5 cm. wide, richly blotched reddish brown, the lateral lobes rounded, the basal third of the lip suborbicular to transversely oblong in outline, undulate-margined, the central portion with a deep median constriction, which may sometimes be prolonged into a short, broad isthmus, the mid-lobe very large, spreading, transversely subquadrate to suborbicular, with a conspicuous bright yellow central blotch, the margins very strongly undulate, the disk at the base of the lip with a prominent, fleshy, minutely papillose, 5-parted, tuberculate crest. Column 6-8 mm. long, with 2 prominent lateral wings above which are 2 elongate, linear processes, thickened at the apices into small blackish glands. osta Rica to Colombia and Ecuador. A very handsome species, often cultivated for its large, richly colored flowers. Although no authentic specimens have been collected in Panama, it has been several times reported from our area, notably from the vicinity of Chiriquí Lagoon and (532) 1949] FLORA OF PANAMA (Orchidaceae) 197 from the vicinity of Puerto Armuelles. Since it is fairly frequent in closely ad- jacent territory in both Costa Rica and Colombia, there is every reason to believe that it will eventually be collected somewhere in the intermediate area. 15. Омсшгум NEBULOSUM Lindl. in Bot. Reg. n. s. 4: Misc. 175. 1841. Oncidium к Morren, іп Ann. Soc. Roy. d'Agr. ес Вог. Gand 4:55, 7. 170. 1848. Oncidium caesium Rchb. f. in Gartenfl. 75. 185 Oncidium Ко кти Rchb. f. in Walp. ree 6:802. 1861. Erect, epiphytic herbs with short, fleshy, approximate, ovoid to suborbicular, somewhat compressed, diphyllous pseudobulbs up to about 4 cm. long and 3 cm. wide, the bases usually with 2 foliaceous bracts. Leaves and bract blades narrowly lanceolate, obtuse to subacute, subcoriaceous, 9-20 cm. long and 2—2.5 cm. wide, contracted below into short, conduplicate petioles. Inflorescences 1 or 2 slender, erect or arching, few-flowered racemes up to about 36 cm. long, produced from the lateral bases of the pseudobulbs. Flowers relatively large, up to about 5 cm. long and 3 cm. wide. Sepals free, subequal, usually reflexed, pale yellow to greenish yellow, shaded or spotted with red or purple, lanceolate, acute, the lateral sepals somewhat oblique, 1.2-1.5 ст. long and 3-5.5 mm. wide. Petals spreading, sub- equal to the sepals or a little broader, ligular to elliptic-lanceolate, obtuse to sub- acute, yellow or greenish yellow, shaded or spotted with red or purple, 1.2-1.5 cm. long and 0.4-0.7 cm. wide. Lip pandurate, 3-lobed, 2-2.5 cm. long and 2-2.5 cm. wide, bright yellow, the lateral lobes small, obliquely triangular or subquad- rate, obtuse, the anterior margins contracted into a narrow isthmus, the mid-lobe abruptly and broadly dilated and 2-lobed, transversely reniform, the disk with a short, fleshy, 5-toothed callus. Column about 5 mm. long, with prominent lateral wings. Mexico, Guatemala, and Panama. Absent from recent collections, but cited by Lindley (Fol. Orch. Oncidium (48). 1855) as "Wild in Veragua, Chiriqui, at 4—5000 ft.—-Warczewicz.” Prob- ably a rather doubtful record, since it is not known from adjacent Costa Rica, or from any of the other Central American countries south of Guatemala. 16. ONCIDIUM OBRYZATUM Rchb. f. in Bonplandia 2:108. 1854. Oncidium obryzatoides Kranzl. іп Engler, Pflanzen. IV, Fam. 50 {нен 80):240. 1922. Oncidium fulgens Schltr. іп Fedde Rep. Sp. Nov. Beih. 17:83. 192 Oncidium Brenesii Schltr. loc. cit. 257. n Oncidium graciliforme C. Schweinf. in Boe Mos Leafl. Harv. Univ. 5:96. 1938. Erect, epiphytic herbs with approximate, suborbicular, ovoid, or elliptic-oblong, compressed, monophyllous pseudobulbs 2.5-9 cm. tall and 0.8—3 cm. wide, usually conspicuously ridged and wrinkled, often spotted or suffused with dark brown or black, the lower portions enveloped in the conduplicate, distichously imbricating bases of several bracts, the uppermost 2—4 of which are conspicuously foliaceous, the plants very variable in size, 1—4.5 dm. tall. Leaves and bract blades elliptic- oblong, ligular or sometimes narrowly linear, subacute, coriaceous, 10—35 cm. long (533) [Vor. 36 198 ANNALS OF THE MISSOURI BOTANICAL GARDEN and 0.8—3.5 cm. wide, contracted below into slender conduplicate petioles. In- florescences usually solitary, erect or arching, many-flowered panicles apparently always exceeding the leaves, in robust plants up to 1.5 m. in length, although often much less, in our specimens averaging about 4.5—6 dm. long. Flowers very vari- able in size, 1.5—3.5 cm. long and 1—2.5 cm. wide. Sepals subequal, free, spread- ing, obovate-spatulate, obtuse or truncate, golden yellow with chestnut-brown blotches at the base, 6-14 mm. long and 2.5-5 mm. wide. Petals usually broader than the sepals, obovate-spatulate, obtuse or truncate, golden yellow with basal chestnut-brown blotches, 6—12 mm. long and 3-6 mm. wide. Lip pandurate, 3- lobed, 10—20 mm. long and 8-20 mm. wide, golden yellow with a U-shaped reddish brown blotch surrounding the yellow basal callus, the lateral lobes subquadrate, obliquely triangular or rounded, the basal 25 of the lip roughly triangular to sub- orbicular, the central portion of the lip abruptly contracted into a short, narrow isthmus, the mid-lobe dilated, spreading, 2-lobed with a deep central sinus, often transversely reniform in outline, the disk with an erect, fleshy basal crest, some- times 5 (or more) -lobulate with 3 distinct apical teeth, or obscurely lobulate with only 2 distinct apical lobes. Column 3-4 mm. long, with prominent, usually porrect, lateral wings, the apices often more or less converging over the anther. Costa Rica, Panama, Colombia, Ecuador, and Peru. COCLÉ: region north of El Valle de Antón, 800-1000 m., Allen 225, 2023, 2038, 3424. CHIRIQUÍ: without 2. инде 4000-5000 ft., Р Powell 3150, 3150, 3160, 3227; vicinity Boquete, Svibla s A common, very variable species of the tree tops of areas of wet highland forest, distinguished from most other Panama Oncidiums by the obovate-spatulate, usually truncate sepals and petals. 17. ONcIDIUM OCHMATOCHILUM Rchb. f. in Bot. Zeit. 10:698. 1852. Oncidium cardiochilum Lindl. Fol. Orch. Oncidium, (27). 1855. Erect, usually robust, epiphytic herbs up to about 75 cm. tall, the many, con- duplicate, distichously imbricating bases of the foliage contracted into a com- planate petiole, often completely enveloping the usually inconspicuous, compressed, narrowly ovoid, diphyllous pseudobulb which may be up to about 10 cm. long and 2.5 cm. wide. Leaves narrowly lanceolate, acute to acuminate, coriaceous with prominent veins, sometimes conduplicate for their entire length, more fre- quently explanate, 25—70 cm. long and 1.5-6 cm. wide. Inflorescences 1 or 2 stout, erect or arching, many-flowered panicles up to about 2 m. in length but sometimes much shorter, produced from the axils of the uppermost basal leaves, the terminal portion of the inflorescence and the lateral branches distinctly fracti- flex. Flowers of moderate size, averaging about 3 cm. long and 2 cm. wide when spread out, with a strong lilac-like fragrance when fresh. Sepals free, pale green marked with brown, the dorsal sepal lanceolate, acuminate, often strongly reflexed, 1.5-2 cm. long and 0.3—0.4 cm. wide, the lateral sepals obliquely linear-lanceolate, acuminate, the dorsal surface with a prominent central keel, 1.5—2 cm. long and (534) 1949] FLORA OF PANAMA (Orchidaceae) 199 0.3—0.4 cm. wide. Petals broader than the sepals, elliptic-lanceolate, acute, brown or greenish brown, sometimes mottled white or with paler apices, 1.3-1.7 cm. long and 0.4—0.55 cm. wide. Lip panduriform, 3-lobed, 1-1.3 cm. long and 1-1.3 cm. wide, white spotted with brown or reddish purple at the base, the lateral lobes sub- quadrate, obtuse, projecting, the base of the lip subequal to, or sometimes exceeding the mid-lobe in width, the central portion contracted into a short distinct isthmus, the mid-lobe abruptly dilated, entire or apiculate, transversely semiorbicular or subcordate, the disk with a prominent, fleshy, multi-tuberculate callus terminating at the apex in a short, erect, obtuse, laterally compressed keel above and below which are 2 pairs of short fleshy teeth. Column 5—6 mm. long, apparently without lateral wings, the base without a foot. Guatemala, Costa Rica, Panama, Colombia, Peru, and probably adjacent terri- tories. cocLÉ: El Valle de Antón, 3600 ft., A. Bouché 0. снитоиТ: without definite local- ity, 4500 ft., Powell 160. The plant collected by Mr. Bouché was reported to have been a massive speci- men weighing in the neighborhood of fifty pounds, with nearly a thousand flowers evidently borne on several inflorescences, the fragrance having been perceptible for some considerable distance. 18. ONCIDIUM PANAMENSE Schltr. in Fedde Rep. Sp. Nov. Beih. 17:85. 1922. Erect, robust epiphytic herbs with approximate, oblong-ovoid, compressed, strongly ridged, diphyllous pseudobulbs 6—16 cm. long and 2—6 cm. wide, en- veloped below in the conduplicate, distichously imbricating bases of 3—4 elongate, foliaceous bracts. Leaves and bract blades linear-lanceolate, acute, subcoriaceous, 12—75 cm. long and 1.5—4 cm. wide, contracted below into elongate, conduplicate petioles. Inflorescences 1 ог 2 erect, arching, pendulous or sometimes scandent, many-flowered panicles 1-3.5 m. in length, produced from the lateral bases of the pseudobulbs. Flowers of moderate size averaging about 2.5 cm. in diameter. Sepals free, subequal, spreading, shortly clawed at the base, yellow heavily barred and blotched olive-brown, elliptic-obovate, acute, with somewhat undulate margins, 10-13 mm. long and 4—5 mm. wide. Petals subequal to the sepals and similarly colored, spreading, often with recurved apices, elliptic-obovate to elliptic-lanceo- late, acute, with somewhat undulate margins, 10—12 mm. long and 4—5 mm. wide. Lip obscurely 3-lobed, broadly pandurate, yellow with a large, transverse, reddish brown to yellowish brown blotch below the crest, the lip 10—15 mm. long and 10—15 mm. wide, the lateral lobes rounded or broadly and obtusely triangular, the central lip with a sharp constriction but without a distinct isthmus, the mid-lobe emarginate, obscurely 2-lobed, transversely oblong or subreniform, the basal and apical halves of the lip of about equal width, the disk with an erect, white, fleshy crest surmounted by 4 divergent, usually crenulate keels, terminating at the apex in 3 distinct fleshy teeth. Column short, 3—5 mm. long, with 2 prominent, spread- ing, usually serrate, lateral wings. Panama. (535) [ Vor. 36 200 ANNALS OF THE MISSOURI BOTANICAL GARDEN NAL ZONE: San Juan, upper Chagres River, sea level, Powell 416, 3201. PANAMA: hills east p DN City, sea level, Powell 3228; without definite locality, 50 m., Kies- wetter s. n.; San José Island, Perlas Archipelago, Erlanson 209; vicinity Pacora, 25 m., G. Fairchild 5. -— vicinity Chepo, 20 m., Allen 2360 А very common, robust, lowland species closely allied to the widely distributed Central American O. sphacelatum, but apparently differing in the broader basal callus and the larger column wings. Among the vegetatively similar or identical local species, it is separable from О. іі ті and О. Baueri by the uniformly broader basal half of the lip and the absence of a true isthmus, and from О. ensatum by the epiphytic habit and the presence of distinct articulations near the bases of the foliaceous bracts. 19. ONcIDIUM PANDURIFORME А. & 5. in Sched. Orch. 8:77. 1925. Erect, caespitose, epiphytic herbs 35—55 cm. tall, with distichous foliage, the conduplicate, imbricating bases enveloping a small, linear to narrowly ovoid, com- pressed, 1- or 2-leaved pseudobulb 5—7 cm. long and 0.8-2 cm. wide, the plants reminiscent of those of Oncidium ochmatochilum. Leaves linear-lanceolate, acute, subcoriaceous, rather coarsely veined, the blades up to about 40 cm. long and 1.5-3 cm. wide. Inflorescences usually solitary, stout, erect or arching, many- flowered panicles much exceeding the leaves, up to about 1 m. in length, produced from the axils of the foliaceous bracts, the terminal portion and the lateral branches usually distinctly fractiflex. Flowers rather small, averaging about 2 cm. in diameter. Sepals subequal, free, spreading or strongly reflexed, yellowish brown with yellow apices, the dorsal sepal lanceolate, acute, 9—12 mm. long and 2.5-3 mm. wide, the lateral sepals linear-lanceolate, acute, 10—12 mm. long and 2-2.5 mm. wide, the reverse surfaces with prominent central keels. Petals usually broader than the sepals, similarly colored, elliptic-lanceolate, acute, spreading or reflexed, 9—10 mm. long and 4—5 mm. wide. Lip pandurate, obscurely 3-lobed, white, 9-12 mm. long and 8-10 mm. wide, the lateral lobes small and rounded ог obtusely triangular, the anterior margins converging to form a short, broad isthmus, the mid-lobe abruptly dilated, spreading, transversely semiorbicular, the under-surface at the apex with a short, strongly developed keel terminating in an apicule, the disk with a low, fleshy, obscurely tricarinate callus at the base. Column about 5 mm. long, without lateral wings but with 2 pronounced, parallel, marginal thickenings on the under-surface below the stigma. Costa Rica and Panama. CHIRIQUI: vicinity Bajo Chorro, headwaters of the Rio Caldera, 1900 m., Woodson & Schery 704. Apparently closely allied to Oncidium ochmatochilum, but differing in the obscure, lateral lobes of the lip, and the much less complexly developed basal callus. 20. ONCIDIUM PARVIFLORUM L. Wms. in Amer. Orchid Soc. Bull. 11:33. 1942. Small, erect, epiphytic herbs, with approximate, elliptic-oblong, compressed, longitudinally ridged, monophyllous pseudobulbs 5—7 cm. long and 2—3 cm. wide, (536) 1949] FLORA OF PANAMA (Orchidaceae) 201 densely spotted or suffused with brown, enveloped below in the conduplicate, distichously imbricating bases of several bracts, the uppermost 2 of which are con- spicuously foliaceous. Leaves and bract blades ligular, obtuse, subcoriaceous, X и y E ONCIDIUM' ME m a | P Ж pou 7 (Ы; Ex Рап) огит 797 di JJ DANN Fig. 203. Oncidium parviflorum (537) [Vor. 36 202 ANNALS OF THE MISSOURI BOTANICAL GARDEN 11-13 cm. long and 1.2-1.6 cm. wide, contracted below into narrow, conduplicate petioles. Inflorescences solitary, erect or arching, many-flowered panicles up to about 75 cm. long, produced from the lateral bases of the pseudobulbs, the lateral branches very short, 1- to 3-flowered. Flowers small, 1.2-1.5 cm. in diameter. Sepals free, spreading, shortly clawed at the base, brown with yellow tips, the dorsal sepal elliptic-oblong, obtuse, shortly apiculate, 5—6 mm. long and about 3 mm. wide, the lateral sepals rather obliquely lanceolate, acute, with prominent dorsal keels, about 6 mm. long and 2.5 mm. wide. Petals clawed at the base, some- what broader than the sepals, spreading, brown with yellow tips, oblong-ovate, obtuse, about 6 mm. long and 3.5 mm. wide. Lip pandurate, 3-lobed, 8—9 mm. long and 5—6 mm. wide, the frontal and lateral lobes yellow, the isthmus brown, the lateral lobes small, auriculate, with subacute deflexed apices, the central portion of the lip abruptly contracted into а broad isthmus, the mid-lobe emarginate and 2-lobed, subreniform, slightly exceeding the lateral lobes in width, the disk with a prominent, pubescent, 4-lobulate callus at the base, at first pure white, becoming yellow with age. Column about 3 mm. long, with 2 prominent lateral wings, which are rounded below and produced above into 2 erect, parallel, acuminate processes on either side of the anther. Panama. COCLÉ: region north of El Valle de Antón, about 1000 m., Allen 2937. A. small-flowered species, found in the tops of tall trees in the wet highland forests north of El Valle de Antón, in Coclé Province. Known only from the type collection. 21. Омсшгум Powe Ltn Schltr. in Fedde Rep. Sp. Nov. Већ. 17:86. 1922. Erect, epiphytic herbs, with approximate, oblong-ovoid, compressed, longitud- inally ridged, monophyllous pseudobulbs 6—13 cm. long and 2.5-5 cm. wide, partially enveloped below in the conduplicate, distichously imbricating bases of 3—4 foliaceous bracts. Leaves and bract blades oblong-lanceolate to ligular, usually obtuse or slightly retuse at the apex, or sometimes subacute, subcoriaceous, up to about 45 cm. in length but averaging about 20-25 cm. long and 2.5-5 cm. wide. Inflorescences usually solitary, arching, few- to many-flowered, flexuose panicles much exceeding the leaves, sometimes up to nearly 1.5 m. long but averaging about 7.5—9 dm., produced from the axils of the uppermost foliaceous bracts, each of the short lateral branches of the inflorescence with 1—3 flowers subtended by a few conspicuous, white, papery, spathaceous sheaths. Flowers variable in size but always large and attractive, 3-6 cm. in diameter. Sepals subequal, spreading, shortly clawed at the base, the blades with strongly undulate margins, rich chocolate-brown barred and margined yellow, the dorsal sepal free, erect, broadly elliptic-oblong to obovate, obtuse to subacute, 1.5-2.5 cm. long and about 1 ст. wide, the lateral sepals free or sometimes with the basal claws shortly connate, rather obliquely oblong, obtuse to subacute, 1.7—3 cm. long and 0.8—1.0 cm. wide. (538) 1949] FLORA OF PANAMA (Orchidaceae) 203 Fig. 204. Oncidium Powellii (539) [VoL. 36 204 ANNALS OF THE MISSOURI BOTANICAL GARDEN Petals spreading, with strongly undulate margins, subequal to the sepals or some- what broader, the bases sessile, the blades rather obliquely oblong, obtuse to sub- acute, rich chocolate-brown with yellow margins and markings, 1.5-2.5 cm. long and 0.6-1.0 cm. wide. Lip pandurate, 3-lobed, shortly clawed at the base, 1.5-2 cm. long and 0.8—1.2 cm. wide, always distinctly shorter than the lateral sepals, the lateral lobes of the lip small, yellow, auriculate to subquadrate, truncate to subacute, the central portion of the lip contracted into a short, distinct, reddish brown isthmus, the mid-lobe bright yellow, quite variable in shape, usually shallowly emarginate, with a minute central apicule, the 2-lobed blade transversely reniform but sometimes only an obovate, obtuse, entire dilation of the isthmus, the disk with a prominent, yellow, multidenticulate callus, terminating at the apex in a broad, transverse, porrect, more or less verrucose or crenulate plate. Column 5-6 mm. long, without conspicuous lateral wings. Panama. CANAL ZONE: Gatun Lake, sea level, Powell 58; without definite locality, Pring s. п.; hills near Frijoles, sea level, Powell 3480. СОСТЕ: vicinity of El Valle de Antón, 800 m Allen 5. Ап apparently rare species, with very handsome chocolate-brown flowers, seem- ingly confined to the lowland forests of the Atlantic slope. It is very closely allied to the Colombian O. anthocrene, and may ultimately prove to be identical. 22. ONcIDIUM PUsILLUM (L.) Rchb. f. in Walp. Ann. 6:714. 1863. Epidendrum pusillum L. Sp. Pl. ed. 2, 1352. 176 Cymbidium pusillum Sw. in Nov. Act. Soc. Upeal 6:74. 1799. Oncidium iridifolium HBK. Nov. Gen. & Sp. 1:344. 1815. Dwarf, epiphytic herbs up to about 8 cm. tall, entirely without pseudobulbs, Leaves distichously spreading, forming a broad fan, equitant, linear, often falcate or ensiform, the apices obliquely acute, 2-6 cm. long and 0.4-1.0 cm. wide. In- florescences 1 to about 6 erect or arching scapes, about equaling the leaves in length, the apices with several acute, strongly compressed, imbricating sheaths, forming a short, complanate raceme from the axils of which the flowers are pro- duced in succession. Flowers large in relation to the size of the plant, averaging 2-2.5 ст. long and 1.8-2 cm. wide. Sepals free, spreading, bright yellow, the dorsal sepal erect, elliptic-lanceolate to subquadrate, obtuse, up to about 5 mm. long and 3-4 mm. wide, the dorsal surface with a pronounced keel, prolonged a: the apex into an erect, acute spur, the lateral sepals linear-lanceolate, acuminate, very closely appressed to the under-surface of the lip, 4-5 mm. long and 1-1.5 mm. wide, the reverse surfaces with a strongly developed keel. Petals spreading, oblong, obtuse, with undulate margins, bright yellow irregularly barred with reddish brown, up to about 8 mm. long and 4 mm. wide. Lip broadly pandurate, 3-lobed, up to 1.8 cm. long and 2 cm. wide, bright yellow, the lateral lobes sub- orbicular to obovate-spatulate, obtuse, spreading, the central portion of the lip contracted into a short, broad isthmus, blotched with reddish brown, the mid-lobe (540) 1949] FLORA OF PANAMA (Orchidaceae) 205 Fig. 205. Oncidium pusillum abruptly dilated, with a deep central sinus, the lateral lobules broadly spreading, with undulate margins, the disk white, spotted reddish orange, with a spreading, subquadrate plate, terminating in a second broad, spreading, semi-orbicular plate with a short central keel and undulate margins. Column short, about 3 mm. long, with prominent, denticulate, lateral wings. Mexico to Brazil and Bolivia; Trinidad. CANAL ZONE: Las Cascadas Plantation, near Summit, Standley 25761, Powell 3018; San Juan, upper Chagres River, sea level, Powell 3346; drowned forest along the Rio Chagres, upper Madden Lake area, Steyerm ark 9% Allen 16771; vicinity Salamanca Hydro- graphic Station, 80 m., Woodson, Allen © Seibert 1559; vicinity Alajuela, 90-100 m., Madden Dam area, Dodge, Steyermark & Allen 16510. PANAMA: Rio La о a tween the Río Bayano and the Gulf of San Miguel, 0-25 m., Allen 17. ВОСА5 DEL T Water Valley, vicinity Chiriquí Lagoon, von Wedel 1383, 1474, 1482, 2153; Western Rice vicinity Chiriqui Lagoon, von Wedel 2777; Almirante Bay, von Wedel 14. Attractive plants, with relatively large, bright yellow flowers, often found as colonies on old orange trees or scattered through the slender, twiggy ends of dead branches, often overhanging water. Widely distributed and common in the Amer- ican tropics. 23. ONcIDIUM sTENOTIS Rchb. f. іп Linnaea 41:67. 1877. Erect, epiphytic herbs with approximate, linear-oblong, compressed, longi- tudinally ridged, monophyllous pseudobulbs 7-14 cm. long and 2.5-4 cm. wide, the lower portions enveloped in the conduplicate, distichously imbricating bases of several bracts the uppermost 3 of which are conspicuously foliaceous. Leaves and bract blades linear-oblong, obtuse to subacute, coriaceous, 15—60 cm. long and 2.5—5 cm. wide, contracted below into very short, conduplicate petioles. In- florescences 1 or 2 erect or arching, few- to many-flowered panicles up to about (541) [Vor. 36 206 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1.5 m. in length, the lateral branches short, usually less than 15 cm. long, the 1 to 5 flowers subtended by thin, white, acuminate, papery bracts. Flowers of moderate size, 3—4 cm. in diameter. бера] free, subequal, spreading, with recurved apices and undulate margins, yellow heavily blotched with brown, the dorsal sepal oblong-lanceolate, shortly acute to apiculate, 1.3-1.8 cm. long and 0.6-0.8 cm. wide, the lateral sepals linear-lanceolate to lanceolate, acute, obliquely deflexed, the reverse surfaces with a strong central keel, 1.5-2 cm. long and 0.5-0.7 cm. wide. Petals subequal to the sepals and similarly colored, spreading, with recurved apices and undulate margins, lanceolate, acute, 1.4—1.6 cm. long and 0.4-0.6 cm. wide. Lip pandurate, 3-lobed, distinctly shorter than the lateral sepals, 1.2-2 cm. long and 1.2-1.8 cm. wide, bright yellow with a brown isthmus, the lateral lobes shortly clawed at the base, spreading, obovate-spatulate, with subquadrate to sub- orbicular blades, the central portion of the lip contracted into a narrow isthmus, the mid-lobe abruptly dilated, emarginate and 2-lobed, often with a short central apicule, transversely subreniform in outline, about equaling the extended lateral lobes in width, the disk with an erect, fleshy callus, the sides with 2 or 3 short teeth, the apex abruptly tridenticulate. Column up to about 7 mm. long, with 2 rather obscure lateral lobes, the under-surface below the stigma conspicuously thickened into 2 parallel, fleshy lobules. Costa Rica and Panama. CHIRIQUÍ: without definite locality, Pring s. n., Pring © Hunter s. п. BOCAS DEL TORO: Water Valley, vicinity Chiriqui Lagoon, von Wedel 1654, 1829, 2156. Allied to О. Baueri, but apparently always differing in the larger flowers, іп which the lip is distinctly shorter than the lateral sepals, and with a conspicuously narrower isthmus. The smaller plants are nearly identical in vegetative appearance with those of O. Powellii, and have sometimes been mistaken for that species. 24. ONCIDIUM sTIPITATUM Lindl. in Bot. Sulphur, 172. 1843. Oncidium lacerum Lindl. in Bot. Reg. n. s. 7: Misc. 30. Oncidium stipitatum Lindl. var. blaiyomyx Rchb. E in D ` Chron. 1:788. 1878. Caespitose, epiphytic herbs with very short, subcylindric to subconic, mono- phyllous pseudobulbs 5-10 mm. long and 5-10 mm. wide, the broadly trun- cate apices with elongate, fleshy, terete, acuminate, longitudinally sulcate leaves, the pseudobulbs and leaf bases of the current season’s growth enveloped in 4-6 closely imbricating, papery bracts which soon weather away. Leaves at first horizontal or ascending, becoming pendulous and wrinkled with age, often more or less spotted with reddish brown, 24-70 cm. long and 0.6—1.0 cm. in diameter. Inflorescences solitary, horizontal or ascending, many-flowered panicles usually about equaling the leaves in length, produced from the bases of the pseudobulbs. Flowers very variable in size, up to about 2.5 cm. long and 1.8 cm. wide, always brightly colored and attractive. Sepals subequal, free, spreading, shortly clawed at the base, yellow richly marked with reddish brown, the dorsal sepal obovate- spatulate, obtuse with a minute apicule, the blade slightly concave, 4.5-6 mm (542) 1949] FLORA OF PANAMA (Orchidaceae) Fig. 206. Oncidium stibitatum (543) 207 [Vor. 36 208 ANNALS OF THE MISSOURI BOTANICAL GARDEN long and 2.5-4 mm. wide, the lateral sepals obliquely obovate, shortly acute, 5-7 mm. long and 3-4 mm. wide. Petals spreading, elliptic-oblong, obtuse, with un- dulate margins, colored and marked similarly to the sepals, 5-8 mm. long and 2—3 mm. wide. Lip pandurate, 3-lobed, 1-2 cm. long and 1-1.8 cm. wide, bright yellow on both the frontal and reverse surfaces, the basal half of the isthmus reddish brown, the lateral lobes linear-oblong, obliquely obtuse, spreading or ascending, the central portion of the lip contracted into a narrow isthmus, the mid-lobe abruptly dilated and 2-lobed, transversely semiorbicular to oblong, often with undulate margins, the disk at the base with a suborbicular transverse plate terminating at the apex in a prominent, erect, fleshy, semiorbicular, somewhat laterally compressed tubercle occupying only the basal half of the isthmus. Column 2-2.5 mm. long, with 2 narrow, spreading, lateral arms. Panama. CANAL ZONE: vicinity of Empire and Culebra Cut, 80 m., Hunter 9 Allen 784; Mira- flores, sea level, Powell 3207; Las Cruces, Powell 3222; Las Cascadas, Powell 3124; Barro Colorado Island, Gatán Lake, Woodworth & Vestal 520; San Juan, upper Chagres River, 0-200 ft., Powell 3137. COLON: forests along the Rio Boquerón, above the Peluca Hydro- graphic Station, Upper Madden Lake region, 90 m., Hunter 8 Allen 654. PANAMA: hills A very common, attractive lowland species apparently confined to the central Isthmian area, particularly on the Pacific slope, where the plants are very abundant in areas of deciduous forest, with spontaneous seedlings often spreading to culti- vated trees and shrubs of the Canal Zone town sites. 25. ONCIDIUM TERES А. & S. in Sched. Orch. 8:78, 7. 7. 1925. Epiphytic herbs with slender, fleshy, terete, longitudinally sulcate leaves. In- florescences paniculate, many-flowered, up to about 4.5 dm. long, produced from the bases of the rudimentary, subcylindric pseudobulbs. Flowers small, up to about 1.5 cm. long and 1 cm. wide. Sepals free, subequal, spreading, shortly clawed at the base, obovate-spatulate, the blades suborbicular, obtuse, concave, minutely apiculate, yellow very heavily spotted reddish brown, about 6 mm. long and 3.5—4.5 mm. wide. Petals spreading, marked and colored similarly to the sepals, oblong-obovate, obtuse, with undulate margins, 5.2-6 mm. long and 3—4 mm. wide. Lip panduriform, 3-lobed, 7-8 mm. long and 6.5-8 mm. wide, the frontal surface bright yellow, the reverse surface and the disk heavily spotted reddish brown, the lateral lobes small, ligular or oblong-spatulate, obtuse to obliquely acute, minutely puberulent, antrorse in natural position, the central portion of the lip contracted into a distinct isthmus, the mid-lobe abruptly dilated, obscurely 2-lobed, transversely reniform, sometimes porrect in natural position, the disk with (544) 1949] FLORA OF PANAMA (Orchidaceae) 209 Бір. 207. Oncidium teres a prominent, complex, fleshy, tuberculate callus, occupying nearly the entire isthmus. Column about 3 mm. long, with 2 lateral, acute, incurving wings. Panama. VERAGUAS: vicinity San Francisco de Veraguas, 1000 ft., Powell 383. cnumuquí: vicinity David, 100 ft., Allen 4242, 4453. A small-flowered lowland species thus far known only from Chiriquí and Veraguas provinces in western Panama. The plants are vegetatively identical with those of Oncidium stipitatum, but the flowers differ in the shorter isthmus of the lip, the much more prominent and complex basal callus, the reddish brown spotting on the reverse surface of the lip, and in other characters. 26. Омсштом Warscewiczu Rchb. f. in Bot. Zeit. 10:693. 1852. Oncidium bifrons Lindl. in Gard. Chron. 84. 1857. Erect, caespitose, epiphytic herbs with approximate, elliptic-ovoid, compressed, usually diphyllous pseudobulbs 4—8 cm. tall and 2—3 cm. wide, the bases enveloped in several conduplicate, imbricating bracts, the uppermost bract being conspicu- ously foliaceous. Leaves ligular, obtuse to subacute, subcoriaceous, 15—30 cm. long and 1.5-3.5 cm. wide, contracted below into slender, conduplicate petioles. Inflorescence a solitary, erect or arching, 4- to 12-flowered, unilateral raceme ex- ceeding the leaves, up to about 5 dm. long, produced from the axil of the upper- most foliaceous bract. Flowers of moderate size, about 3 cm. in diameter, uniformly yellow throughout, the elongate pedicels subtended by conspicuous, papery, spathaceous bracts. Sepals dissimilar, the dorsal sepal free, oblong, obtuse, with slightly undulate margins, 10-12 mm. long and 5—6 mm. wide, the lateral sepals connate nearly to the apex, forming a single obovate, bifid segment below the lip, the lower surface with 2 distinct keels, 1.4—1.6 cm. long and 0.7—0.9 cm. wide. Petals obliquely oblong, obtuse, with slightly undulate margins, 1.2-1.4 cm. long and 0.7-0.8 cm. wide. Lip subpandurate, obscurely 3-lobed, 1.5-1.8 cm. long and 1.5—1.8 cm. wide at the apex, the lateral lobes reduced to inconspicuous auricles, the basal half of the lip nearly oblong, the apical half abruptly dilated and 2-lobed, reniform, about twice as wide as the basal half, the disk with a nar- (545) [Vor. 36 210 ANNALS OF THE MISSOURI BOTANICAL GARDEN row, fleshy crest, the base under the column semiterete, terminating at the apex in 5 short, divergent teeth. Column 8—10 mm. long, the apex dilated, with narrow lateral wings. Costa Rica and Panama. Originally described from plants collected by von Warscewicz, presumably on the slopes of Chiriquí Volcano, at 8000—10,000 ft. elevation. Although the species is not represented in recent collections from our area, its frequent occurrence in the adjacent Costa Rican highlands would lend weight to the supposition that this one citation may be authentic. DOUBTFUL SPECIES ONcIDIUM ADVENA Rchb. f. in Hamb. Gartenzeit. 16:422. 1860. Originally described from flowering material only, from plants presumably imported from Venezuela. An Endres collection is cited by Krünzlin, from the vicinity of the old Obispo railway station, in the Canal Zone. The figure given of a single flower looks like Oncidium ensatum. Absent from all recent collections. ONCIDIUM ONUSTUM Lindl. Gen. & Sp. Orch. Pl. 203. 1833. A poorly known South American species, sometimes doubtfully listed as occur- ring in Panama. Absent from all recent collections. ONCIDIUM PELIOGRAMA Linden & Rchb. f. in Gard. Chron. 1451. 1871. Described from flowers only. Оп the basis of the sketch of the type in the Ames Herbarium, the species which we list as Oncidium stenotis might possibly be referable to this concept, but the drawing is too poor for positive identification. In the absence of authentic material for comparison, or even any description of vegetative parts, it seems best to exclude it as a species of doubtful identity. 79. LEOCHILUS Knowles & Westcott Leocnitus Knowles & Westc. Fl. Cab. 2:143. 1838; Benth. & Hook. Gen. РІ. 3:564. 1883; Kränzl. in Engler, Pflanzenr. IV, Fam. 50 (Heft 80):291. 1922. Cryptosanus Scheidw. in Otto & Dietr. Allg. Gartenzeit. 11:101. 1843. Rhynchostele Rchb. f. in Bot. Zeit. 10:770. 1852. Cryptosaccus Scheidw. ex Rchb. f. Xenia Orch. 1:15. 1854. Leiochilus Benth. in Jour. Linn. Soc. 18:328. 1881, non Hook. uewa Regel, in Gartenfl. 40:89, /. 7341. 1891. Rhynchostelis Jackson, in Index Kew. 2:718. 1895, sphalm. Small, caespitose, epiphytic herbs with short, compressed, monophyllous or rarely diphyllous pseudobulbs, the lower portions enveloped in several membra- naceous or foliaceous sheaths. Leaves elliptic-lanceolate to ligular, coriaceous, contracted below into a petiole. Inflorescences 1 or 2 erect or arching racemes or panicles produced from the lateral bases of the pseudobulbs. Flowers small. Sepals subequal, spreading, free, or more or less connate. Petals subequal to the sepals or sometimes broader. Lip entire or 3-lobed, adnate to the base of the (546) 1949] FLORA OF PANAMA (Orchidaceae) 211 column, spreading, usually longer than the lateral sepals, the disk fleshy ог callous. Column short, erect, not laterally winged but usually biauriculate below the stigma, the clinandrium truncate, the rostellum elongate, the base of the column without a foot. Anther terminal, operculate, incumbent, 1-celled, produced in front into a hooded appendage; pollinia 2, waxy. A small genus of tropical American epiphytes closely allied to Oncidium, ranging from Mexico and the West Indies to northern Argentina. Two species are known from Panama. a. Lateral cba ne for more than half their length aa. Lateral sepals f 1. LEOCHILUS LABIATUS (Sw.) O. Ktze. Rev. Gen. Pl. 2:656. 1891 (as Leio- chilus) . Epidendrum labiatum Sw. Prodr. Veg. E. Occ. 124. 1788. Liparis labiata Spr. Syst. Veg. 3:741. Rodriguezia cocblearis Lindl. in Ann. 5 Mis, Nat. Hist. 5:116. 1840. Leochilus cochlearis Lindl. in Bot. Reg. : Misc. 23. 1842. Oncidium labiatum Rchb. f. in Walp. iud rs 1863 L. LABIATUS L. scRIPTUS Ne Dwarf, epiphytic herbs 4—9 cm. tall, with elliptic oblong, compressed, mono- phyllous pseudobulbs 10-15 mm. long and 5-8 mm. wide, the bases with 2—3 foliaceous bracts, the leaves and pseudobulbs often reddish brown. Leaves and bract blades elliptic-lanceolate to ligular, obtuse to subacute, coriaceous, 2—6 cm. long and 0.6—1.5 cm. wide. Inflorescences 1 or 2 slender, erect, few-flowered racemes or shortly branching panicles much exceeding the leaves in length, 4—25 cm. long. Flowers small, averaging about 1 cm. in diameter. Sepals of about equal length, yellow striped or spotted reddish brown, the dorsal sepal free, elliptic- lanceolate, shortly acuminate, deeply concave, 3—5 mm. long and 1.5-2 mm. wide, the lateral sepals connate to above the middle, forming a single bifid segment below the lip, 4—6 mm. long and 2.5-3 mm. wide, with 2 distinct keels on the lower surface. Petals subequal to the dorsal sepal and similarly colored, elliptic- oblong, subacute, with distinct central keels on the reverse surfaces, 3—5 mm. long and 2-3 mm. wide. Lip entire, slightly concave, yellow with a red or reddish brown spot at the base, oblanceolate, obtuse or sometimes slightly retuse, the blade with an obscure median constriction, 5—8 mm. long and 2.5-4 mm. wide, the disk with an erect, bicarinate, fleshy callus. Column 1—1.5 mm. long, semiterete, with- out lateral wings but with 2 short, porrect teeth below the stigma. Antilles, Panama, and Trinidad. cocLÉ: El Valle de Antón, 800 m., Allen 2295. A dwarf species, fairly frequent on calabash trees in cultivated plots, the plants being rendered conspicuous by the reddish brown leaves and pseudobulbs. Our specimens are nearly identical with typical West Indian material, but differ con- siderably from specimens of Leochilus gracilis Schltr. from Guatemala, Honduras, and Costa Rica, which would seem to be distinct. Іп L. labiatus the plants are (547) [Vor. 36 472 ANNALS OF THE MISSOURI BOTANICAL GARDEN from 4—9 cm. in height, with elongate scapes up to 25 cm. long, the flowers averaging slightly larger, with the lateral sepals connate to above the middle; while in L. gracilis the plants seem to be 2-4.5 cm. tall, the scapes only up to about 6 cm. long, with the lateral sepals entirely free. 2. LEOCHILUS sCRIPTUS (Scheidw.) Rchb. f. Xenia Orch. 1:15, 7. б. 1854. Cryptosanus scriptus Scheidw. in Otto & Dietr. E “qoqqa, 11:101. 1843. Oncidium scribtum Rchb. f. in Walp. Ann. 6:772 Leochilus major Schltr. in Fedde Rep. Sp. Nov. 15: iM. тој: инт РошеШ Ші Schltr. loc. cit. 17:81. Leiocbilus retusus Schltr. loc. cit. 19:256. 1923. Erect, caespitose, epiphytic herbs with approximate, oblong-ovoid to elliptic- oblong, compressed pseudobulbs 2-5 cm. long and 1—2.5 cm. wide, the truncate apices usually monophyllous, the lower portions of the pseudobulbs enveloped in the conduplicate bases of the several distichously imbricating bracts the uppermost 2 or 3 of which are conspicuously foliaceous. Leaves ligular, obtuse or shortly and unequally 2-lobed at the apex, coriaceous, 5.5-14 cm. long and 1-2.8 cm. wide, contracted below into very short, conduplicate petioles. Inflorescences 1 or 2 erect or arching, few- to about 12-flowered racemes or weakly branching pani- cles, equaling or somewhat exceeding the leaves, up to about 25 cm. in length. Flowers relatively large for the genus, averaging about 1.5—2 cm. in diameter, gteenish white to greenish yellow with purple or reddish brown markings. Sepals free, subequal, spreading, the dorsal sepal elliptic-lanceolate, acute, somewhat con- cave, the reverse surface with a distinct keel, 9—12 mm. long and 3.5-5 mm. wide, the lateral sepals spreading or somewhat reflexed, ligular to oblong-lanceolate, obtuse to subacute, 8—12 mm. long and 2.5-5 mm. wide, the reverse surfaces with a distinct central keel. Petals subequal to the dorsal sepal, lanceolate, acute, 9—12 mm. long and 2.5-3.5 mm. wide. Lip entire, spreading or slightly convex, the blade obovate-oblong, truncate, up to about 12 mm. long and 7 mm. wide, the narrower base conspicuously thickened and adnate to the base of the column, the broader apex shallowly emarginate, the reverse surface with a distinct central keel, the disk with a short, subquadrate, basal boss terminating at the apex in 2 distinct, parallel, erect, fleshy, puberulent calli. Column up to about 3 mm. long, semi- terete, the under-surface below the stigma with 2 distinct, ligular, porrect arms. Guatemala, Honduras, Costa Rica, and Panama. 1 4” Las Cascadas, sea level, Powell 3140, Standley 25755, 20700; vicinity Gat tcher s. n. PANAMA: Juan Díaz Range, sea level, Powe 120; Arraijan, fed С е Río La Maestra, 0-25 m., Allen 50; vicinity Chepo, 30 m., Allen 2355. 80. SIGMATOSTALIX Rchb. f. SIGMATOSTALIX Rchb. f. in Bot. Zeit. 10:769. 1852; Benth. & Hook. Gen. Pl. 3:565. 1883; Schltr. in Fedde Rep. Sp. Nov. 15:142. 1918. Ornithophora Barb. Rodr. Gen. & Spec. Orch. Nov. 2:225. 1882. (548) 1949] FLORA OF PANAMA (Orchidaceae) 213 Small, caespitose, epiphytic herbs with approximate, elliptic-oblong to ovoid, compressed, monophyllous or rarely diphyllous pseudobulbs, the lower portions partially enveloped in several distichously imbricating bracts the uppermost 2 to 5 of which are usually foliaceous. Leaves linear to elliptic-lanceolate, subcoriaceous to coriaceous. Inflorescences 1 to about 5 erect or arching, few- to many-flowered racemes or panicles about equaling or sometimes much exceeding the leaves in length, produced from the axils of the foliaceous bracts. Flowers small to minute. Sepals subequal, free, or with the laterals shortly connate at the base. Petals sub- equal to the sepals. Lip usually with an elongate claw, sometimes subsessile, the blade entire or 3-lobed, subquadrate, suborbicular or trulliform. Column slender, elongate, often somewhat arcuate, the apex more or less dilated opposite the stigma. Anther terminal, operculate, incumbent, 1-celled; pollinia 2, waxy. A small genus of tropical American epiphytes, ranging from Mexico to Brazil. Four species are known to occur in Panama. a. x^ арта without an elongate claw ас the bas b. Basal c of the lip subquadrate, with a ЊЕ ыңаа idc i . S. RACEMIFERA о к = m % о ч ы e] 2 on XE pri nt avity bb. Basal s of the lip transversely subreniform, bifid, with a E tubercle in the center of each > % шені рде 3. 5. НҮМЕМАМТНА аа. Lip with а disti tinct, elongate claw e bas b. Blade of the lip алы Ды уы фе auriculate lobules ас the base 1. S. ABORTIVA bb. Blade of the i ovate-sagittate to cordate, with 2 distinct auriculate lobules at the base 2 S. GUATEMALENSIS 1. SiGMATOSTALIX ABORTIVA L. Wms. in Ann. Mo. Bot. Gard. 27:284, #. 34, figs. T warf, caespitose, epiphytic herbs, with oblong-elliptic, compressed, mono- phyllous pseudobulbs 12—20 mm. long and 3—11 mm. wide, enveloped below in the conduplicate, distichously imbricating bases of 2 foliaceous bracts. Leaves and bract blades narrowly linear-ligular to elliptic-lanceolate, acute, subcoriaceous, 2—4 cm. long and 0.5-1.0 cm. wide, contracted below into very short, conduplicate petioles. Inflorescences usually solitary, slender, arching, few-flowered racemes up to about 10 cm. long. Flowers minute, averaging about 5 mm. in length. Sepals subequal, free, spreading or reflexed, yellow, lanceolate, acute, about 3.2 mm. long and 0.8—1 mm. wide. Petals somewhat broader than the sepals, yellow, elliptic- lanceolate to oblanceolate, subacute and obscurely apiculate, about 3.2 mm. long and 1.3 mm. wide. Lip with an elongate claw at the base, obovate-spatulate, white, about 5 mm. long and 3.5 mm. wide, the abruptly dilated blade truncate at the apex, subquadrate to suborbicular in outline, the disk with a fleshy, cucullate callus covering the entire upper surface of the claw, the obtuse apex abruptly erect. Column slender, about 3 mm. long, dilated at the apex. Panama. COLÓN: Quebrada Lopez, lower slopes of Cerro Santa Rita, 30 m., Allen 2121. (549) [Vor. 36 214 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 208. Sigmatostalix guatemalensis (550) 1949] FLORA OF PANAMA (Orchidaceae) 215 2. SIGMATOSTALIX GUATEMALENSIs Schltr. in Fedde Rep. Sp. Nov. 10:253. 1911. Sigmatostalix costaricensis Rolfe in Kew Bull. 78. 1916; Bot. Mag. /. 8825. 1919. Sigmatostalix poikilostalix Kranzl. in Engler, Pflanzenr. IV, Fam. 50 (Heft 80):310, fig. 27, D:a-e. Caespitose, epiphytic herbs with approximate, elliptic-oblong, compressed, monophyllous pseudobulbs 1.5-3.5 cm. long and 0.6-1.5 cm. wide, partially en- veloped below in the conduplicate bases of several distichously imbricating bracts the uppermost 2 or 3 of which are conspicuously foliaceous. Leaves and bract blades ligular to elliptic-lanceolate, acute, subcoriaceous, 3.5-12 cm. long and 0.8—1.5 cm. wide, contracted below into short, conduplicate petioles. Inflorescences 1 or 2 slender, erect, few- to many-flowered racemes, usually much exceeding the leaves in length, 11—32 cm. long, produced from the axils of the uppermost foliaceous bracts. Flowers relatively large for the genus, averaging about 12-15 mm. in diameter when spread out. Sepals free, subequal, usually strongly reflexed, pale green to yellow, usually marked with brown, ligular-lanceolate, acute, 6—9 mm. long and 1.5-2.5 mm. wide. Petals subequal to the sepals and similarly colored, usually strongly reflexed, rather obliquely lanceolate, acute, 6—9 mm. long and 2-3 mm. wide. Lip with a distinct, narrow, fleshy claw at the base, 2-3 mm. long, produced in front into a short, subacute spur, the blade of the lip abruptly dilated, rather obscurely 3-lobed, spreading or slightly convex, broadly trulliform, ovate-sagittate to cordate, usually reddish brown with a dark yellow apex, rarely entirely yellow, 5-6 mm. long and 4—5 mm. broad, the lateral lobules small, auriculate or with subfalcately acute, incurving projections from the pos- terior margin, the apex of the blade obtuse to acute, with a short, distinct, central keel on the under-side. Column elongate, slender, somewhat arcuate, up to about 6 mm. long, the lower portion terete, the apex shortly dilated on either side of the stigma, the base without a foot. Mexico, Guatemala, Costa Rica, and Panama. CHIRIQUÍ: without definite locality, 5000 ft., Powell 230. 3. SiGMATOSTALIX HYMENANTHA Schltr. in Вей. Bot. Centralbl. 367:419. 1918. Small, caespitose, epiphytic herbs with approximate, ovoid to elliptic-oblong, compressed, monophyllous pseudobulbs 1.2-3.5 cm. long and 0.8-2 cm. wide, the lower portions partially enveloped in the conduplicate, distichously imbricating bases of several bracts, the uppermost 2 or 3 of which are conspicuously foliaceous. Leaves and bract blades linear-ligular, acute, subcoriaceous, 3.5-14 cm. long апа 0.4—0.8 cm. wide, contracted below into narrow, conduplicate petioles. Inflores- cences 1 or 2 slender, erect, shortly branching panicles, usually about equaling the leaves in length, produced from the axils of the uppermost foliaceous bracts, the lateral branches of the panicles usually very short and densely bracteate. Flowers minute. Sepals and petals oblong, acute, about 2 mm. long and 0.5 mm. wide. Lip obscurely 3-lobed, about 1.5 mm. long and 2 mm. wide, subsessile or obscurely (551) [Vor. 36 216 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN clawed at the base, the blade broadly ovate, obtuse, spreading or slightly convex, the disk with a fleshy, 2-lobed, transversely subreniform callus, with an obscure, rounded tubercle іп the center of each of the lateral lobules. Column stout, terete, erect, slightly dilated at the apex, about 1.5 mm. long, the base without a foot. Costa Rica and Panama. DARIEN: vicinity Chepigana, Cana-Cuasi trail, 2000 ft., Terry 9 Terry 1437. SIGMATOSTALIX RACEMIFERA L. Wms. in Ann. Mo. Bot. Gard. 27:285, ¢. 36. 1940. Small, caespitose, epiphytic herbs 7—15 cm. tall, with elliptic-oblong, com- pressed, monophyllous pseudobulbs 2-3.5 cm. long and 1-2 cm. wide, the lower portions enveloped in the conduplicate, distichously imbricating bases of several bracts, the uppermost 3 to 5 of which are conspicuously foliaceous. Leaves and bract blades linear-ligular to elliptic-lanceolate, acute, subcoriaceous, 3-13 cm. long and 0.5-1.8 cm. wide, contracted below into short, conduplicate petioles. Inflorescences 1 to 5, slender, erect or arching, few- to many-flowered, shortly branching panicles about equaling or somewhat exceeding the leaves in length, produced from the axils of the foliaceous bracts. Flowers small. Sepals subequal, spreading or reflexed, greenish white, the dorsal sepal free, lanceolate, acute, about 3 mm. long and 0.75 mm. wide, the lateral sepals shortly connate at the base, elliptic-lanceolate, acute to shortly acuminate, about 2.5 mm. long and 0.8 mm. wide. Petals somewhat broader than the sepals, greenish-white spotted maroon, elliptic-lanceolate, acute, about 3 mm. long and 1.25 mm. wide. Lip subsessile or obscurely clawed at the base, the blade subquadrate, truncate, with a distinct median constriction, orange, about 3 mm. long and 3.5 mm. wide, the disk with a fleshy, subquadrate callus, with a deep longitudinal excision on each side of the central linear concavity. Column elongate, slender, about 5 mm. long, terete below, dilated above, with 2 acute auricles on each side of the stigma. > Panama. cocrÉ: hills north of El Valle de Antón, 800-1000 m., Allen 1232, 2266, 2744, 2878, 3745. 81. LOCKHARTIA Hook. Госкнакттл Hook. in Bot. Mag. t. 2715. 1827; Benth. & Hook. Gen. Pl. 3:570. 1883; Kränzl. in Engler, Pflanzenr. IV, Fam. 50 (Heft 83):6. 1923. Fernandezia R. & P., sensu Lindl. Сеп. & Sp. Orch. РІ. 207. 1833, in part. Caespitose, epiphytic herbs without pseudobulbs, the linear, undivided, pseudo- monopodial, foliaceous stems erect or pendulous, entirely enveloped in the short, equitant, distichously imbricating, coriaceous leaves. Inflorescences short 1- or 2- flowered scapes or sometimes abbreviated few-flowered panicles, produced from the upper leaf axils. Flowers of moderate size to small, usually yellow, on elongate, filiform pedicels, often subtended by conspicuous membranaceous bracts. Sepals (552) 1949] FLORA OF PANAMA (Orchidaceae) 217 subequal, free, spreading or with the laterals reflexed. Petals subequal to the sepals or broader. Lip usually complexly 3-lobed, rarely entire, usually conspicuously exceeding the sepals in length, the lateral lobes (if present) often linear, rather long, divaricate or antrorsely incurving, the mid-lobe variously 2- to 4-lobulate, the disk with a papillose, denticulate or tuberculate callus. Column very short, with two broad wings or auricles, the base without a foot. Anther terminal, oper- culate, incumbent, imperfectly 2-celled; pollinia 2, waxy. A rather perplexing genus of perhaps 30 species of American epiphytes, ranging from Mexico to Peru and Brazil. Six species, and one variety, are known to occur in Panama. a. Inflorescences relatively large, spreading, divaricate panicles ‚ L АСОТА Е аа. Inflorescences very short, compact, subsessile Жар ог 1- to 2- d scapes b. Lip entire or subentire, without distinct linear lobes at the base. Inflorescences 1-flowered. c. Leaves acute. Apex of the lip deeply emarginate 7. І. PrrTIERI cc. Leaves broadly obtuse. Apex of the lip entire 5. L. OBTUSATA bb. Lip with distinct linear е at the base. Inflorescences 1-flowered or shortly racemose or pan c. Flowers very small, the p mer be than 4 mm. long. Leaves bt the obliquely truncate or obtuse at t . L. MICRANTHA cc. Flowers сеф base the lip 7 mm. go or more. Leaves acute or subacut d. Mid- Бе of the lip ‘subpandurate, Pa spi oi broader at the apex than e base, with a dist isthmus or constriction between ey Acl portion and the pete: la iid lobules.......... 6. L. OERSTEDII dd. Mid-lobe of the lip = г subquadrate or conspicuously broader apex. e. Mid-lobe E us composed of 4 distinct or obscure, subequal lobules L. AMOENA ee. Mid-lobe quadrate-triangular, more or less distinctly 3- parted OENA Var. TRIANGULABIA 1. LockHarmTia ACUTA (Lindl.) Rchb. f. in Bot. Zeit. 10:767. 1852. ernandezia acuta Lindl. in Bot. Reg. 21: Ё. 1806. 1836. уин ballida Rchb. f. іп Bonplandia 2:14. 1854. Caespitose, epiphytic herbs with flattened, pendulous, foliaceous stems, 15—50 cm. long and 1—2 cm. wide. Leaves, as normally seen in profile, rather obliquely triangular, acute or sharply apiculate at the apex, coriaceous, 12—30 mm. long and 4—10 mm. wide. Inflorescences usually 1 to 3 relatively large, spreading, divari- cate, few- to many-flowered panicles up to about 8 cm. long, produced from the subterminal leaf axils. Flowers small, white with a yellow lip, less than 10 mm. іп diameter when spread out, on slender filiform pedicels subtended by small, sub- cordate, papery bracts. Sepals free, subequal, spreading, ovate, obtuse, concave, -4 mm. long and about 2 mm. wide. Petals elliptic-oblong, obtuse, 4-5 mm. long and about 2 mm. wide. Lip more or less rectangular in outline, about 6 mm. long and 4 mm. wide, the basal half subquadrate, slightly concave, with narrow, erect, lateral margins, the sessile base adnate to the base of the column, the mid- lobe 4-parted, usually somewhat reflexed, the lateral lobules acute, spreading or (553) [Vor. 36 218 ANNALS OF THE MISSOURI BOTANICAL GARDEN | 2 ` 55 e Vcore WAG (m V^ b ve Pas . ру ® Fig. 209. Lockhartia acuta (554) 1949, FLORA OF PANAMA (Orchidaceae) 219 divaricately extrorse, the projecting central lobule oblong to subquadrate, the apex deeply emarginate, the disk with a bifid, puberulent callus, the concave base under the column with a minute, semiglobose, densely papillose, shortly stipitate process. Column very short, 1-1.5 mm. long, with broadly spreading triangular or auricv- late wings on each side of the stigma, the base without a foot. Panama, Colombia, Venezuela, and Trinidad. CANAL ZONE: Pedro Miguel, sea level, Powell 3051; pres Chagres River, Madden ues region, 70-75 m., Steyermark 17515. PANAMA: hills near Panama City, sea level, owell pem oh of Juan Díaz, near Panama City, 20—50 m., n Pittier Esi Chiva- jeg ons sea id. Powell 3014, 3019; San José Island, Perlas АтсЬ аро, Johnston 1217. сото Río Gatuncillo, vicinity Santa Rosa, 25 m., Allen n 4253. A common lowland species with very small flowers, readily recognizable by the relatively long, pendulous, foliaceous stems and the conspicuous, broadly paniculate inflorescences. 2. LocKHARTIA AMOENA Endres & Rchb. f. in Gard. Chron. 666. 1872. Lockhartia costaricensis Schltr. in Fedde Rep. Sp. Nov 190 Lockhartia-grandibracteata Kránzl. in Englert кей. гу, Pa N (Heft 83):15. | 1925. Caespitose, epiphytic herbs with erect or pendulous, flattened, foliaceous stems 12-40 cm. long and 1.4-2.5 cm. wide. Leaves narrowly triangular in profile, subacute, coriaceous, 1.5—3.5 cm. long and 0.4-1.0 cm. wide. Inflorescences short, compact, few-flowered panicles, up to about 3 cm. long. Flowers relatively large for the genus, on long filiform pedicels subtended by conspicuous, ovate- cordate, membranaceous bracts. Sepals free, subequal, slightly concave, yellow, the dorsal sepal erect or somewhat reflexed, ovate, obtuse, with a minute, terminal apicule, 5-6 mm. long and about 4 mm. wide, the lateral sepals strongly reflexed in natural position, elliptic-lanceolate, subacute, 5-6 mm. long and about 4 mm. wide. Petals somewhat longer than the sepals, yellow, elliptic-oblong, obtuse, in- curving, the lateral margins conduplicately reflexed in natural position, 5-7 mm. long and about 4 mm. wide when spread out. Lip complexly 3-lobed, yellow, with reddish brown markings at the base, about 10 mm. long and 8 mm. wide, the lateral lobes linear-ligular, acute, antrorsely incurving in natural position, the mid-lobe subquadrate, distinctly or obscurely 4-lobulate, with undulate margins, the truncate apex with a deep central sinus, the basal lobules equaling or somewhat exceeding the 2-parted mid-lobule in width when spread out, very strongly re- flexed in natural position, the disk with a linear, truncate, papillose callus. Column very short, with broadly spreading, lateral wings. Costa Rica and Panama. COCLE: p pum of El Valle de Antón, 1000 m., Allez 2313; mountains beyond La m 400— жи 9 Allen 586. cutrigui: Palo Alto Hill, 4000-5000 ft., Рош 362; рија а Сй Alta, Finca Lérida, eastern slopes of Chiriquí Volcano, 1500- 2000 m., Woodson, Allen P. Seibert 066. (555) [Vor. 36 220 ANNALS OF THE MISSOURI BOTANICAL GARDEN A variable and rather perplexing highland species with relatively large, attrac- tive flowers. It is probably allied to Г. Oerstedii, but differs in the mid-lobe of the lip, which is more or less subquadrate and 4-lobulate, the basal half equaling or exceeding the apical half in width. 3. LOCKHARTIA AMOENA Endres & Rchb. f. var. TRIANGULABIA С. Schweinf. & Р. H. Allen in Bot. Mus. Leafl. Harv. Univ. 13:150. 1948. Lockhartia triangulabia А. & S. in Sched. Orch. 8:80. 1925. Vegetative parts and inflorescences apparently as in the type. Sepals subequal, free, probably more or less reflexed in natural position, about 5.4 mm. long an 3 mm. wide, the dorsal sepal ovate, subacute, the lateral sepals rather obliquely elliptic-oblong, obtuse, all three sepals dorsally mucronate at the apex. Petals oblong, obtuse, about 5.4 mm. long and 3 mm. wide. Lip broadly quadrate- triangular in outline, complexly 5-lobulate, about 7.2 mm. long and 10 mm. wide at the base, the lateral lobes rather obliquely ligular, obtuse, the mid-lobe more or less 3-parted, the basal lobules triangular, the basal half of the mid-lobe about twice as wide as the subquadrate, emarginate apical half, the disk with a linear- oblong, minutely papillose callus, the base under the column thickened into a semiorbicular minutely papillose boss. Panama. CHIRIQUÍ: without definite locality, 4000-5000 ft., Powell 362-А. Lochbartia amoena, а represented Бу the quite numerous specimens available for study, exhibits a distinct tendency to vary from a condition in which the basal and apical halves of the 4-parted mid-lobe are of subequal width, to one in which the bifid, apical segment is considerably narrower. Our present single specimen apparently represents an extreme example of this tendency and seems sufficiently distinctive to warrant varietal rank. 4. LOCKHARTIA MICRANTHA Rchb. f. in Bot. Zeit. 10:768. 1852. Lockhartia chiriquiensis Schltr. in Fedde Rep. Sp. Nov. 12:215. 1913. Lockhartia Lankesteri Ames, in Sched. Orch. 5:36. 1923. Caespitose, epiphytic herbs with erect or pendulous, flattened, foliaceous stems typical of the genus, 5—40 cm. long and 1—2 cm. wide. Leaves equitant, coriaceous, in profile narrowly triangular with obliquely truncate or obtuse apices, 0.8—2 cm long and 0.35—1.0 cm. wide. Inflorescences simple 1- to 3-flowered scapes or abbreviated, few-flowered panicles up to about 2 cm. long, produced from the central or subterminal leaf axils. Flowers small, yellow, on slender, filiform pedicels subtended by ovate-cordate, membranaceous bracts. Sepals subequal, free, spreading, concave, ovate to elliptic-oblong, acute and apiculate, 2.5-4 mm. long and 2—3 mm. wide. Petals broadly ovate to elliptic-ovate, obtuse, 2.5—4 mm. long and 1.5-2 mm. wide. Lip complexly 3-lobed, 2.5-5 mm. long and 2.5-5 mm. wide, the lateral lobes linear-ligular, obtuse, spreading or reflexed, the mid-lobe (556) 1949] FLORA OF PANAMA (Orchidaceae) 221 Жа LOCKHARTIA ANN obtusata «C. Wms Fig. 210. Lockhartia obtusata (557) [Vor. 36 222 ANNALS OF THE MISSOURI BOTANICAL GARDEN rhombic-obovate to obovate, the lateral margins obscurely lobulate, the apex deeply retuse, the disk with a low, slightly concave, more or less bifid callus. Column very short, with spreading lateral wings, the base without a foot. Nicaragua to Surinam (fide Kranzlin). AL ZONE: Balboa, on mango tree, Standley 25404; Río Pedro Miguel, near East NL Standley 20084; Lu Cruces, Powell 3189; Miraflores, Powell 3203; Pedro рн Powell 3040; drowned forests of the Quebrada Tranquilla, Madden Lake region, Dodge 9 Allen 17325; Río Indio, Madden Lake region, Steyermark & Allen 17437, 17440. PANAMA: io La Maestra, coastal area east of the Río Bayano, 0-25 m., Allen бі, сос: hills north of El Valle de Antón, 1000-1200 m., Allen 2318, 2903, 3420, 3939, 4259. VERA- GUAS: vicinity Santa Fé, 1000 ft., Allen 4429; vicinity Bahía Honda, Taylor 1510. IRIQUÍ: vicinity San Felix, 0-120 m., РИНег 5286; without definite locality, 4000 ft., Powell 3476. A common small-flowered species somewhat reminiscent of Lockhartia acuta, but distinguished by the usually shorter foliaceous stems, much shorter inflores- cences, yellow flowers, and differently shaped lip. 5. LockHaRTIA OBTUSATA L. Wms. in Amer. Orch. Soc. Bull. 9:209, 7. 8. 1941. Caespitose, epiphytic herbs with broad, strongly flattened, foliaceous stems up to about 35 cm. tall and 2.5-3.5 cm. wide. Leaves equitant, distichously imbri- cating, coriaceous, obliquely oblong as seen in profile, obtuse, 2-4 cm. long and 0.5-1.0 ст. wide. Inflorescences short, condensed racemes from the upper leaf axils. Flowers relatively large and attractive, bright yellow with a conspicuous orange callus at the base, apparently produced singly in succession from the term- inal cluster of ovate, acute, papery bracts. Sepals subequal, free, spreading, ovate, obtuse to shortly acute, minutely apiculate, about 9 mm. long and 6 mm. wide. Petals elliptic-oblong, obtuse, 9-10 mm. long and 5-6 mm. wide. Lip entire, suborbicular, slightly convex, 9-10 mm. long and 9—10 mm. wide, the anterior margin sometimes slightly retuse, the disk with an elliptic-oblong to obovate, obtuse callus, about 5 mm. long and 3-4 mm. wide, the apex conspicuously thick- ened and papillose, with a short, erect, acute, central spur. Column about 2 mm. long, with conspicuous lateral wings, the base without a foot. Panama. cocrÉ: hills north of El Valle de Antón, 1000 m., Allen 2160, 3550. A strikingly handsome highland species apparently allied to Г. Pittieri, but distinguished by the broad, obtuse leaves, the larger flowers, and the unique, entire, suborbicular lip. 6. LockHartia Овквтерп Rchb. f. Xenia Orch. 1:100, 105, /. 40. 1855. Lockhartia verrucosa Rchb. f. in Hamb. Gartenzeit. 15:53. 1859. Lockhartia е Rchb. f. loc. cit. 16: EN 1860 Fernandezia robusta Batem. in Bot. Mag. t. 5502. Lockhartia М. (Batem.) Schltr. in Fedde Rep. Sp. ‘Nov. 3:82. 1906. (558) 1949] FLORA OF PANAMA (Orchidaceae) 223 Caespitose, epiphytic herbs with strongly flattened, foliaceous stems 7—40 cm. long and 0.8—1.5 cm. wide. Leaves narrowly triangular as seen in profile, acute, 1-3 cm. long and 0.4-0.8 cm. wide. Inflorescences very short, 1- to 3-flowered racemes produced from the axils of the upper leaves. Flowers very variable in size, but often relatively large and attractive, the pedicels subtended by papery, ovate- cordate bracts. Sepals usually strongly reflexed, sometimes slightly concave, yellow, the laterals sometimes spotted with red, elliptic-oblong to ovate, obtuse to shortly acute, 4-8 mm. long and 3-5 mm. wide. Petals yellow, sometimes spotted red, oblong-ovate to broadly and obliquely ligular, obtuse or slightly retuse at the apex, antrorsely incurving in natural position, the lateral margins usually more or less undulate and strongly reflexed, 4-8 mm. long and 3—6 mm. wide. Lip complex- ly 3-lobed, 7-12 mm. long and 6—14 mm. wide, yellow with red or reddish brown markings at the base, the lateral lobes linear-ligular, obtuse or obliquely acute, spreading or antrorsely incurving in natural position, the mid-lobe more or less pandurate, 4-lobulate, the basal half distinctly narrower than the apical half, the small basal lobules rounded or obliquely triangular, strongly reflexed in natural position, sometimes confluent with the bases of the linear lateral lobes, the center of the mid-lobe with a distinct constriction which is often prolonged into a short isthmus, the apical lobule usually abruptly dilated, 2-parted with a deep central sinus, the disk with a narrowly flabellate or sometimes linear, strongly verrucose callus, extending to the median constriction. Column with spreading, minutely denticulate or crenulate lateral wings. Mexico, Guatemala, Honduras, Costa Rica, and Panama. снікюоі: Llano del Volcán, southwestern slopes of Chiriquí Volcano, 1500 m., Allen 8 Fairchild 3502; vicinity Cerro Punta, headwaters of the Rio Chiriqui Viejo, 2000 m., Allen & Fairchild 3403; valley of the upper Rio Chiriqui Viejo, vicinity Monte Lirio 1300-1900 m., Seibert 132, 133, 186; without definite locality, 4000-4500 ft., Powell 3344, 3352; vicinity Bajo Mono, headwaters of the Rio Caldera, 4500 ft., Davidson 584; vicinity Finca Lérida, eastern slopes of Chiriqui Volcano, 1750 m., Woodson & Schery 236; forested slopes along Quebrada Velo, 5000 ft., Allen 4743; bamboo-oak forest, south of Finca Lérida, eastern slopes of Chiriqui Volcano, 6000-7000 ft., Allen 4764; forested hills east of Boquete, 4500-6500 ft., Allen 4666. A common, attractive highland species apparently closely allied to the South American L. lunifera (Lindl.) Rchb. f. In Panama, the species is readily separable from its nearest ally, L. amoena, by the subpandurate mid-lobe of the lip, which is always broadest at the apex. 7. LOCKHARTIA PITTIERI Schltr. in Fedde Rep. Sp. Nov. 12:216. 1913. Lockhartia variabilis A. & S. in Sched. Orch. 8:81. 1925. Caespitose, epiphytic herbs with distichous, foliaceous stems up to about 20 cm. tall and 2.5-4.5 cm. wide. Leaves linear-lanceolate to narrowly triangular as seen in profile, obliquely acute to acuminate, 2—3.5 cm. long and 0.4—0.9 cm. wide. Inflorescences very short, apparently solitary, 1-flowered scapes produced from the axils of the upper leaves. Flowers of moderate size for the genus, yellow (559) [Vor. 36 224 ANNALS OF THE MISSOURI BOTANICAL GARDEN with an orange callus at the base. Sepals elliptic-lanceolate, acute, with apiculate tips, 3.9-4.9 mm. long and about 2 mm. wide. Petals somewhat broader than the sepals, elliptic-oblong to suborbicular, obtuse to acute, 4-5 mm. long and 2.4-3.2 mm. wide. Lip oblong-quadrate, 7-8 mm. long and 5-6.4 mm. wide, slightly con- vex, the apex deeply emarginate, the lateral margins sometimes obscurely lobular, the disk with an elliptic-oblong to ovate-oblong callus, the margins elevated, the apex conspicuously thickened and minutely papillose, with an erect, subacute, central spur, the inner elliptic depression with a low, rounded boss. British Honduras, Costa Rica, and Panama. L ZONE: vicinity Bohio, Pittier 3401; Fort Sherman and mouth of the Chagres River, Powell 372; wooded hills near Frijoles, sea level, Powell 355; Cativo-Porto Bello trail, Powell 361. DARIÉN: vicinity Marraganti, 10-200 ft., R. S. Williams 1008. DOUBTFUL SPECIES LOCKHARTIA MIRABILIS Rchb. f. Xenia Orch, 1:100, 106, 2. 40, figs. 11-12. 1855. Apparently described from flowering material only, without vegetative parts. The figures in the ‘Xenia Orchidaceae’ might be poor drawings of what we know as Lockhartia Oerstedii, seen in natural position, with the lateral lobules of the mid-lobe reflexed. Since both of these concepts seem very close to that of the South American L. lunifera Rchb. f., and since L. mirabilis is absent from all recent collections in Panama, it would seem best to exclude it, pending the rather remote possibility of direct examination of type material of both this and L. lunifera. LocKHARTIA ELEGANS Hook. in Bot. Mag. £. 2715. 1827. A South American species, apparently having the northern limit of its range in Trinidad. Listed by Schlechter from Panama, but absent from all collections. LocKHARTIA OBTUSIFOLIA Regel in Ann. Sci. Nat. VI, 2:378. 1856. A Colombian species, doubtfully listed by Kranzlin from Panama. Absent from all collections. 82. ORNITHOCEPHALUS Hook. ORNITHOCEPHALUs Hook. Exot. Fl. 2:/. 127. 1825; Benth. & Hook. Gen. Pl. 3:568. 1883. Small, epiphytic herbs without pseudobulbs, the equitant, fleshy or coriaceous leaves distichously imbricating and forming a broad or narrow fan, the blades articulated below, those at the base of the fan caducous, the persistent sheathing bases sometimes thickened and resembling pseudobulbs. Inflorescences 1 to several slender, erect or pendulous racemes produced from the leaf axils. Sepals subequal, free, spreading, often concave. Petals subequal to the sepals or larger and flabellate. Lip entire or 3-lobed, the subsessile base continuous with the base of the column, the lateral lobes (if present) membranaceous or sometimes thickened, the mid- lobe short or more frequently elongate, often inflexed, the disk with a fleshy, bifid, bialate or bicornute callus. Column short, rather stout, without wings or ap- (560) 1949] FLORA OF PANAMA (Orchidaceae) 225 pendages, the rostellar process very long and attenuate, the base of the column without a foot. Anther terminal, operculate, incumbent, imperfectly 2-celled, produced above the cells into an attenuate appendage resting on the rostellar process, sometimes exceeding it in length, the column and anther resembling the head and beak of a bird (genus unknown!) ; pollinia 4, waxy. A small genus of dwarf, pseudobulbless, epiphytic herbs, ranging from Mexico to Brazil. Four species are known from Panama. a. s of the inflorescence densely glandular-hispid. b. Lip linear-navicular, incurving, the fleshy basal plate with 2 eiut a horns ‚ Q. BIC b. Lip subquadrate- cochleate, without 2 divergent horns at the base......2. О. ses RN aa. Rachis of the in еа родени glabrous ог minutely puberulent : Flowers T ly large. Lip panduriform, broadest at “Whe bn with a ct fae " parted callus . O. PowELLII bb. Flowers peri Lip linear-navicular, incurving, the lateral uc thickened and erect 3. O. INFLEXUS 1. ORNITHOCEPHALUS BICORNIS Lindl. in Bot. Voy. Sulphur, 172. 1843. Ornit bocepbalus ж muu ` in PEE p. Sp. Nov. 3:251. 1906. Zygostates costaricensis Nash, in Bull. Tor 5. "Club 34:122, Z. 8. 1907. Ornit bocepbalus L АЙЕ un d in Ped Biol. Soc. Wash. 34:152. 1921. Ornitbocepbalus diceras Schltr. in Fedde Rep. Sp. Nov. Beih. 17:87. 1922. Small, epiphytic herbs 3.5—12 cm. tall, the equitant leaves distichously ar- ranged in the form of a broad fan. Leaf blades articulated to the conduplicate, imbricating, persistent bases, coriaceous, lanceolate to gladiate as seen in profile, obliquely acute, 2-9 cm. long and 0.4-1.2 ст. wide. Inflorescences 1-10 slender, many-flowered, axillary racemes about equaling the leaves in length, the rachis densely hispid with spreading glandular hairs. Flowers small, greenish or yellowish, on short hispid pedicels subtended by ovate, acute, membranaceous, ciliate bracts. Sepals suborbicular, concave, about 2 mm. in diameter, the reverse surfaces some- what hispid, with a central elevated keel, terminating in a short mucro. Petals subequal to the sepals, very shortly clawed at the base, the blades orbicular, con- cave, about 2 mm. in diameter, the reverse surfaces more or less hispid, with a ciliate central keel, terminating іп a short mucro. Lip entire, linear-navic- ular, acute, strongly incurving, about 4-5 mm. long when spread out, the base with a fleshy, papillose, subquadrate to suborbicular, cushion-like callus, produced at the sides into 2 short, divergent, horn-like appendages each about 1 mm. in length. Guatemala, Honduras, Costa Rica, and Panama. CANAL ZONE: drowned forest of the Quebrada Ancha, upper Madden Lake region, of t ío Pequení, Madden Lake area, 70-80 m., Dodge, Steyermark & PANAMA: hills ма of Panama Сігу, sea level, Powell 174; edge of forest along Panama- Pacora road, near Rio Tecümen, Killip 3216 without definite locality, Sinclair s.n. VERAGUAS: with- out Шы. locality, Hinds (561) [Vor. 36 226 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 211. Ornithocephalus bicornis А common, small-flowered lowland species distinguished from O. cochleari- formis by the elongate, narrowly navicular, incurving lip and from O. inflexus by the densely hispid rachis and the divergent horns of the basal callus. 2. ORNITHOCEPHALUS COCHLEARIFORMIS C. Schweinf. in Bot. Mus. Leafl. Harv. Univ. 4:124. 1937. Epiphytic herbs 3.5—12 cm. tall. Leaves as seen in profile narrowly elliptic- oblong to broadly ensiform, acute or obliquely acute, 3-7.5 cm. long and 0.7-1.5 cm. wide, the blades articulated to the oblong, persistent, conduplicate bases. Inflorescences 1—8 slender, arching or pendulous racemes up to about 9 cm. long, the rachis densely glandular-hirsute. Flowers of moderate size for the genus, on slender hirsute pedicels subtended by broadly ovate to suborbicular bracts with glandular-ciliate keels and margins. Sepals suborbicular to obovate, concave, (562) 1949] FLORA OF PANAMA (Orcbidaceae ) 127 white, about 3 тт. іп diameter, the reverse surfaces and margins densely gland- ulaz-hirsute, the lateral sepals slightly oblique, with lacerate keels on the outer surfaces. Petals subequal to sepals, white, broadly obovate, concave, the outer surfaces ciliate, with a conspicuously lacerate central keel. Lip deep green, entire, broadly calceiform, deeply concave, about 4.2 mm. long and 3.2 mm. wide, the truncate or subcordate base adnate to the base of the column, the shortly acute apex strongly inflexed, the disk transversely thickened, minutely papillose, the Jateral margins decurrent, with 2 short, lateral, acute swellings, finally truncately converging below at the base of the inner concavity of the lip. Panama. сосгЕ: lower valley and marshes along the Rio Antón, 600 m., Hunter & Allen 383, Allen 3020, 4233; rim of El Valle de Antón, 2500 ft., Purdom s.n.; hills north of El Valle de Antón, 1000 m., Allen 2202. A common highland species of the region of El Valle de Antón, in Coclé Province. It is distinguished by the densely glandular-hirsute outer surfaces of the sepals and petals, and by the broadly calceiform lip. 3. ORNITHOCEPHALUS INFLEXUS Lindl. in Ann. Nat. Hist. I, 4:384. 1840. Ornithocephalus mexicanus A. Rich. & Gal. in Ann. Sci. iy i Ш, 3:24. 1845. Ornitbocebbalus elepbas Rchb. f. in Walp. Ann. 6:493. Ornithocephalus Pottsiae n Wats. in T. Brigh. Guatem. M E: 429. 1887. Ornitbocepbalus Tonduzii Schltr. in Beih. Bot. Centralbl. Үр. :420. 1918. Dwarf, ерірһутіс herbs, the equitant, imbricating leaves forming а fan as in the genus. Leaves linear-lanceolate to ensiform as seen in profile, acute to acuminate, coriaceous, 4.5—8 cm. long and 0.25-0.5 cm. wide, the blades articulated to the persistent, conduplicate bases. Inflorescences 1 to several slender racemes up to about 8 cm. long, produced from the axils of the leaves, the rachis glabrous or minutely puberulent. Flowers small, on slender pedicels subtended by oblong-lanceolate, acute, minutely ciliate bracts. Sepals suborbicular, concave, about 2 mm. in diameter, the reverse surfaces essentially glabrous, with distinct central keels, terminating in a short mucro. Petals broadly flabellate-cuneate to dolabriform, about 2 mm. long and 3 mm. wide. Lip entire, linear-ligular, acute to acuminate, navicular, usually inflexed, about 5 mm. long and 2 mm. wide, the base thickened into a short bialate or biauriculate callus, the fleshy rounded or subquadrate wings usually erect or with the upper margins converging. Mexico, British Honduras, Guatemala, Honduras, Costa Rica, and Panama. CHIRIQUÍ: without definite locality, 4000 ft., Powell 431. > ORNITHOCEPHALUS PowELLN Schltr. іп Fedde Rep. Sp. Хоу. Вей. 17:88. 1922: Epiphytic herbs 5-8 cm. tall, the equitant leaves distichously arranged in the form of a broad fan, as in the genus. Leaves subfalcately ligular as seen in profile, obliquely acute, often shortly apiculate, coriaceous, 3.5-6.5 cm. long and 0.6-1.2 (563) [VoL. 36 228 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN cm. wide, the blades articulated with the very short, conduplicate, persistent bases. Inflorescences 1 or 2 slender racemes up to about 9 cm. long, produced from the leaf axils, the rachis laterally compressed, narrowly winged, subglabrous to puberu- lent. Flowers rather large for the genus, deep green with a white basal callus, the short, angulate pedicels subtended by elliptic-ovate, acute bracts which have a strongly developed keel and minutely ciliate margins, the apices of the bracts shortly mucronate. Sepals spreading or reflexed, ovate to oblong-ovate, concave, about 4 mm. long and 2.5-3 mm. wide, the outer surfaces with a distinct central keel, terminating in a short mucro, the margins of the sepals ciliate or serrulate. Petals much broader than the sepals, obovate-cuneate to obovate-flabellate, about 6 mm. long and 8 mm. wide, the upper rounded margins minutely ciliate to serrulate. Lip panduriform, obscurely 3-lobed, about 1 cm. long and 0.5 cm. wide at the base, the broadly ovate to subcordate basal half adnate to the base of the column, the anterior margins converging in an elongate isthmus, the subflabel- late apex rather abruptly dilated and truncate, with serrulate margins, the apical half of the lip usually porrect in natural position, the disk with a prominent fleshy, porrect, divergently 2-lobulate callus. Panama. PANAMÁ: foothills east of Panama City, sea level, Powell 231; San Juan de Pequení [now under water in the Madden Lake area], Powell 3279. The largest-flowered species of the genus in our area. The pair of low, di- vergent, linear calli described by Schlechter is an optical illusion produced in translucent liquid material by the shadows of the margins of the lateral sepals which rest against the back of the lip. 82-A. OAKES-AMESIA C. Schweinf. & P. H. Allen OAKES-AMESIA C. Schweinf. & P. H. Allen in Bot. Mus. Leafl. Harv. Univ. 13:133, і. IO. 1948. Dwarf, epiphytic herbs without pseudobulbs, the equitant, coriaceous leaves distichously imbricating and forming a broadly radiate cluster. Leaf blades ensi- form as seen in profile, obliquely acute, articulated to the conduplicate bases. Inflorescences erect or arching racemes produced from the subterminal leaf axils. Flowers small, the short pedicels subtended by elliptic-ovate, acuminate bracts. Sepals free, subequal, spreading, or the laterals reflexed, elliptic-oblong, subacute, the reverse surfaces with a distinct central keel. Petals much larger than the sepals, spreading, broadly obovate-flabellate. Lip somewhat fleshy, sharply 3-lobed, exceeding the lateral sepals in length, the base sessile and continuous with the base of the column, the lateral lobes very prominently developed, erect in natural posi- tion, the mid-lobe more or less subquadrate when seen from above, somewhat dilated and obscurely biauriculate at the abruptly truncate apex, the small lateral auricles obliquely acute, the under-surface of the mid-lobe produced into a very prominent, obliquely triangular, obtuse to subacute, laterally compressed keel (564) 1949] FLORA OF PANAMA (Orchidaceae) 229 which is apparently hollow with a narrow transverse opening at the apex of the lip, the disk with a low, linear-ligular, truncate, densely papillose callus. Column very slender and subterete below, somewhat dilated above, wingless and footless, a little below the lightly arcuate apex with a very large, distinctly 3-lobulate rostellar process. Anther suborbicular, terminal, operculate, incumbent, imper- fectly 2-celled; pollinia 4, apparently cartilaginous. A genus of dwarf highland epiphytes thus far known only from Panama. The plants strongly resemble those of the allied genus Ornithocephalus, but the flowers are amply separated by the very large, 3-lobulate rostellar process, the relatively very short terminal appendage of the anther, the short, abruptly truncate, sharply 3-lobed lip, the prominent, apparently hollow keel of the lip, the linear-ligular callus, and other characters. 1. OAKES-AMESIA CRYPTANTHA C. Schweinf. & P. H. Allen in Bot. Mus. Leafl. Harv. Univ. 13:134, ¢. IO. 1948. Dwarf, epiphytic herbs, the leaves forming a broadly radiate cluster 4 cm. tall and 7.5 cm. wide. Leaf blades ensiform as seen in profile, obliquely acute, 1.8—3.9 cm. long and 0.3—0.5 cm. wide, articulated to the conduplicate bases, the plants identical in vegetative appearance with those of Ornithocephalus inflexus. In- florescences 1 to 3 erect or arching racemes about 5.5 cm. long, produced from the subterminal leaf axils, the rachis slightly fractiflex and rather complanate, narrowly triangular in cross-section. Flowers small, the sepals and petals white, the lip dark green marked with white, the short pedicels subtended by long, elliptic- ovate, acuminate bracts which have strongly developed Кее!5 ‘оп the lower sur- faces, the bracts conspicuously exceeding the floral pedicels in length. Sepals free, subequal, elliptic-oblong, subacute, slightly concave, unevenly serrulate, irregularly glandular-verrucose on both surfaces, the reverse surfaces with a distinct central keel, terminating in an elongate spinous process, the dorsal sepal spreading or slightly reflexed, about 2.7 mm. long and 1.25 mm. wide, the lateral sepals strongly reflexed, about 2.9 mm. long and 1.5 mm. wide. Petals much larger than the sepals, spreading, broadly obovate-flabellate, the abruptly truncate blades with shallow transverse concavities below the irregularly serrulate upper margins, the blades about 3 mm. long and 3 mm. wide at the apex, both the frontal and reverse surfaces irregularly glandular-verrucose. Lip rather fleshy, sharply 3-lobed, the sessile base continuous with the base of the column, the very prominent lateral lobes rather obliquely oblong, truncate, erect in natural position and somewhat antrorse, about 1.5 mm. long and 1 mm. wide, minutely ciliate, the mid-lobe sub- quadrate when seen from above, somewhat dilated and obscurely biauriculate at the abruptly truncate apex, the small lateral auricles obliquely acute, the under- surface of the mid-lobe produced into a very prominent, obliquely triangular, obtuse to subacute, laterally compressed keel which is apparently hollow with a narrow, transverse opening at the apex of the lip, the entire lip sparsely and irregularly glandular-verrucose, about 4 mm. long and 2.5 mm. wide at the apex, (565) [Vor. 36 230 ANNALS OF THE MISSOURI BOTANICAL GARDEN the disk with a low, linear-ligular, abruptly truncate, densely papillose callus. Column very slender below, wingless, footless, below the slightly arcuate apex extends a very large, 3-lobulate rostellar process which considerably exceeds the rest of the column in length and bulk, 3 mm. long and 1 mm. wide, the lateral lobules semiorbicular, abruptly deflexed in natural position, the porrect, slightly undulate apical lobule more or less terete, all 3 lobules of the rostellar process rather densely glandular-verrucose. Column above the rostellar process short, slightly arcuate, the anther suborbicular, terminal, operculate, incumbent, imperfectly 2-celled, with a slender, ligular, acuminate appendage, the rostellar process and anther rather resembling the head and folded legs of a praying mantis when seen in profile. Panama. COCLÉ: Cerro Райса, hills north of El Valle de Antón, 1000-1200 m., Allen & Allen 4106. 83. NOTYLIA Lindl. NorYLIA Lindl. in Bot. Reg. 11:sub 2. 930. 1825; Rchb. f. Xenia Orch. 1:49. 1854; Benth. & Hook. Gen. Pl. 3:586. 1883. Tridachne Liebm. ex Lindl. in Paxt. Flow. Gard. 3:45. 1852-53, nom nud. Macroclinium Barb. Rodr. Gen. & Spec. Orch. Nov. 2:236. 1882. Small, epiphytic herbs with or without pseudobulbs. Leaves usually flat, rarely equitant, coriaceous or fleshy. Inflorescences arching or pendulous, few- to many- flowered racemes, or rarely panicles, produced from the bases of the pseudobulbs ог axils of the leaves. Flowers usually small, rarely of moderate size, оп slender pedicels, the subtending bracts usually narrow. Sepals subequal, usually narrow, erect or spreading, free or with the laterals more or less connate. Petals subequal to the sepals or smaller. Lip entire or obscurely lobed, usually clawed at the base, rarely sessile, continuous with the base of the column, the lamina triangular or hastate, acuminate, the disk smooth or with a carinate callus. Column erect, terete or sulcate, glabrous, papillose or velutinous, without wings or appendages, the apex usually slightly recurved, the rostellum more or less elongate, acuminate, erect, the base of the column without a foot. Anther erect, oblong, imperfectly 2-celled, above the cells more or less long-appendaged and applied to the rostellum; pollinia 2, waxy. Small tropical American herbs ranging from Mexico to Brazil They have been divided by Cogniaux into two well-marked subgenera, in the first of which, EUNOTYLIA, the plants have small but distinct, monophyllous pseudobulbs, with flat leaves; while in the second, MACROCLINIUM, the leaves are equitant and dis- tichously imbricating, the conduplicate bases sometimes enveloping a small com- planate pseudobulb. Five species, representing both subgenera, are known from Panama. (566) 1949] FLORA OF PANAMA (Orchidaceae) 231 a. Leaves equitant, ensiform. Lip cordate-hastate, distinctly 3-lobed. (Subgenus 2. N. Сокреѕп аа. е I ligular to oblong- lanceolate. Lip trulliform. (Subgenus NOT сао distinctly t papillose 5. N. PENTACHNE bb. Ier glabrous. . Lip about 5 mm. wide at the base 3. N. LATILABIA cc. Lip less than 3 mm. wide at the base. d. Dorsal "m серен лс Зай С obtuse, about 5 mm. wide, distinctly bro; than 4. N. PANAMENSIS d. Dorsal id linear- qst 2 v de 2 mm. wide or less, dis- tinctly narrower than the lip [ N. BARKERI 1. Мотуша BARKERI Lindl. in Bot. Reg. n. s. 1: Misc. 90. 1838. Notylia multiflora Hook. in Lond. Jour. Bot. 3:315, 7. 10. 1844, non Lindl. Notylia Huegelii Fenzl. in Denkschr. Akad. Wien, Math. Nat. Kl. 2:255. 1850. Notylia ag кот Lindl. & Paxt., in Paxton’s Flow. Gard. 3:45. 1852-53; Bot. Mag. t. 8306. Tridachne virens ш ех Lindl. & Paxt. voe cit. 1852-53. -53. Notylia tamaulipensis Rchb. f. in Hamb. Gartenzeit. 16:281. Notylia guatemalensis S. Wats. in Proc. Amer. Acad. 22:477. 1887, non Schltr. Notylia angustilancea Schltr. in Orchis 8:135, 7. 4, figs. 8-13. 1914. Notylia guatemalensis Schltr. in Fedde Rep. Sp. Nov. 15:208. 1918, non S. Wats. Notylia Bernoullii Schltr. in Beih. На o 362:502. 1918, nomen. Notylia Pittieri Schltr. loc. cit. 418. Notylia turialbae Schltr. in Fedde Rep. Se. Nov. Beih. 19:145. 1923. Notylia Brenesii Schltr. loc. cit. 249. Small, epiphytic herbs with me compressed, monophyllous pseudobulbs up to about 2.5 cm. long and 1.0 cm. wide, the bases enveloped in several imbricating bracts, the uppermost 1 or 2 of which are conspicuously foliaceous. Leaves and bract blades ligular, obtuse to subacute, coriaceous, up to about 18 cm. long and 0.8-4 cm. wide. Inflorescences 1 or 2 arching or pendulous, densely flowered racemes up to about 30 cm. long but often considerably less, produced from the bases of the pseudobulbs. Flowers small, white, sometimes spotted yellow, оп filiform pedicels, subtended by narrow, acuminate bracts. Sepals subequal, more or less spreading, the dorsal sepal free, linear-lanceolate, subacute, concave, some- what inflexed, 3-6 mm. long and 1-2 mm. wide, the lateral sepals usually more or less connate, rarely free, forming a single bifid segment under the lip, 3—5 mm. long and 1-2 mm. wide, rather obliquely subacute. Petals subfalcately linear- lanceolate, acute, 2.5—5 mm. long and 1—1.5 mm. wide. Lip clawed at the base and continuous with the base of the column, the lamina narrowly triangular, acuminate, 3—5 mm. long and 1—2 mm. wide at the base, the disk usually with a distinct carinate callus. Column slender, erect, terete, 2-3 mm. long, the base without a foot. Mexico to Panama. AL ZONE: г Balboa, sea level, Powell 405. VERAGUAS: Santa Fé, 2500 ft., Powell 40r. 22 without definite locality, 4000 ft., Powell 427. (567) [Vor. 36 432 ANNALS OF THE MISSOURI BOTANICAL GARDEN The plants and flowers are quite variable in size and in the degree to which the lateral sepals are connate, but the floral structure otherwise is rather remarkably uniform, particularly іп view of the great geographic range. It is difficult to believe that any great weight should be attached to the degree of connation of the lateral sepals since they are found to be very easily separable, both in specimens in liquid and in flowers boiled out for examination, so that one specific concept can be transformed into another simply by a little carelessness with the dissecting instruments! 2. NorvLiA Corpvesu L. Wms. in Ann. Mo. Bot. Gard. 26:286, t. 21, figs. 3-4. 1939, Dwarf, epiphytic herbs with equitant, distichously imbricating leaves, the conduplicate bases enveloping a small, oblong-complanate, monophyllous pseudo- bulb 1—1.5 cm. long and about 0.5 cm. wide, the plants reminiscent of those of an Ornitbocepbalus. Leaves lanceolate to linear-lanceolate as seen in profile, ensi- form, rather obliquely acuminate, the blades articulated to the conduplicate bases, 4-6 cm. long and 0.3-0.5 cm. wide. Inflorescences slender, essentially simple, subumbellate racemes, 4—6 cm. long, with 1 or 2 slender, simple, secondary in- florescences usually being produced from the nodes of the primary scape. Flowers of moderate size for the genus, on spreading, filiform pedicels subtended by lanceo- late, acute or acuminate, papery bracts. Sepals free, subequal, the dorsal sepal linear-lanceolate, acuminate, about 10 mm. long and 1.5 mm. wide, the lateral sepals rather obliquely linear-lanceolate, acuminate, about 12-13 mm. long and 0.6—1 mm. wide. Petals obliquely lanceolate, acuminate, about 10 mm. long and 1.2 mm. wide. Lip with an elongate basal claw, about 4 mm. long, the middle with a biauriculate thickening, the anterior margin of which is papillose-pubescent, the blade of the lip abruptly dilated, cordate-hastate, acuminate, about 4 mm. long and 2 mm. wide, the retrorse lateral lobes distinctly serrulate. Column slender, terete, about 3 mm. long, typical of the genus. Panama and Costa Rica. BOCAS DEL TORO: Mosquito Hill, Woodson, Allen 9 Seibert 1932. Apparently closely allied to Notylia Wullschlaegeliana Focke. Known only from the type collection and a single subsequent collection from the Dulce Golfo region in Costa Rica. 3. NOTYLIA LATILABIA А. & S. in Sched. Orch. 8:71. 1925. Epiphytic herbs unusually large for the genus, up to about 25 cm. tall, with short, oblong-complanate, striate-rugose, monophyllous pseudobulbs up to about 1.5 cm. long and 0.8 cm. wide, completely concealed by the large ovate-lanceo- late, papery bracts which become fibrous with age. Leaves oblong-lanceolate to elliptic-lanceolate, subacute, coriaceous, up to about 20 cm. long and 4—5 cm. wide, contracted below into short, conduplicate petioles. Infloresences 1 or 2 arching or pendulous, densely flowered racemes 13—21 cm. long, produced from (568) 1949] FLORA OF PANAMA (Orchidaceae) 233 the base of the current pseudobulb. Flowers relatively large for the genus, on slender pedicels subtended by narrowly triangular, acuminate bracts. Sepals con- cave, orange, the dorsal sepal free, fornicate, oblong-lanceolate, acute when flat- tened out, about 9—10 mm. long and 2.3-3 mm. wide, the lateral sepals connate to nearly the apex, forming a single bifid segment below the lip, about 8-9 mm. long and 4 mm. wide when spread out, the apices strongly reflexed. Petals fal- cately linear-ligular, acute, entirely white, antrorsely incurving in natural position, the apices deflexed, 7.6-8.4 mm. long and about 1.6 mm. wide. Lip shortly clawed at the base and obliquely inserted on the base of the column, the blade broadly trulliform, 6.3—7.1 mm. long (including the claw) and 4-5 mm. wide, the abruptly acuminate apex strongly recurved in natural position, the disk with а low carinate callus. Column slender, terete, glabrous, about 4.9 mm. long, typical of the genus. Panama. CANAL ZONE: Frijoles, sea level, Powell 406. Reminiscent of Notylia Barkeri, but differing in the larger vegetative habit and much broader lip. 4. NorTYLIA PANAMENSIS Ames, Orchidaceae 7:112. 1922. Epiphytic herbs rather large for the genus, up to about 18 cm. tall, with ob- long, compressed, monophyllous pseudobulbs about 2 cm. long and 1 cm. wide, partially enveloped in the papery imbricating bracts which apparently soon weather away. Leaves oblong-lanceolate, obtuse and unequally bilobulate, coria- ceous, up to 15 cm. long and 3—3.5 cm. wide, contracted below into very short, conduplicate petioles. Inflorescences pendulous, densely flowered racemes up to about 20 cm. long, produced from the base of the current pseudobulb. Flowers relatively large for the genus, white, on slender pedicels subtended by scarious, narrowly triangular, acuminate bracts. Sepals spreading, the dorsal sepal broadly elliptic-oblanceolate, obtuse, concave, about 7 mm. long and 5 mm. wide, the lateral sepals connate to the apex, forming a single linear-lanceolate segment below the lip, about 8 mm. long and about 2 mm. wide. Petals elliptic-lanceolate, acute, about 7 mm. long and up to 2.5 mm. wide. Lip clawed at the base, obliquely in- serted on the base of the column, the blade sagittate, about 6 mm. long (including the claw) and 3 mm. wide across the base, the disk with a short, carinate callus. Column slender, terete, glabrous, about 3 mm. long. Panama. DARIEN: Marraganti and vicinity, 10-200 ft., К. 5. Williams 977. A very distinctive species by reason of the broadly elliptic-obovate, obtuse dorsal sepal. Known only from the type collection. 5. NOTYLIA PENTACHNE Rchb. f. in Bonplandia 2:90. 1854. Notylia gracilispica Schltr. in Fedde Rep. Sp. Nov. Beih. 17:75. 1922. (569) [Vor. 36 254 ANNALS OF THE MISSOURI BOTANICAL GARDEN Epiphytic herbs of variable size, up to about 25 cm. tall, with oblong, com- pressed, monophyllous pseudobulbs up to about 2.5 cm. long and 1.2 cm. wide, usually partially enveloped in the conduplicate bases of the foliaceous bracts. Leaves and bract blades oblong-lanceolate to ligular, obtuse to subacute, coriaceous, up to about 20 cm. long and 2—5 cm. wide, contracted below into short condupli- cate petioles. Inflorescences 1 or 2, usually elongate, slender, pendulous, many- flowered racemes up to about 35 cm. long, produced from the base of the current pseudobulb. Flowers of moderate size for the genus, on filiform pedicels sub- tended by inconspicuous linear-lanceolate, acuminate bracts. Sepals pale green, concave, the dorsal sepal lanceolate, acuminate, 8—10 mm. long and about 2-2.5 mm. wide, the apex slightly recurved, the lateral sepals connate for more than half their length, forming a single bifid segment below the lip, about 10 mm. long and 2.5 mm. wide, the subfalcate, acuminate apices divergently spreading and retrorse in natural position. Petals about 8 mm. long and 1.5 mm. wide, rather obliquely lanceolate, shortly acuminate, white with 2 minute orange spots in the center. Lip with an elongate, slender basal claw, the blade abruptly dilated and trulliform, acuminate, white, about 6 mm. long (with the claw) and up to about 2 mm. wide at the base, the disk with a short, carinate callus. Column slender, terete, distinctly papillose, about 5 mm. long. Panama. CANAL ZONE: along Rio Chagres between Gamboa and Alajuela, 30-60 m., Allen 961; vicinity Gamboa, Allen 3026, Pittier 2610; Las Cruces, Powell 3130, Frijoles, Powell 3318; hills east of Panama City, Powell 185, 3230. veERAGUAS: Santa Fé, about 2500 ft., Powell 400. A common lowland species, closely allied to the widespread N. Barkeri, but differing in the papillose column. 84. MACRADENIA К. Br. MACRADENIA R. Br. in Bot. Reg. 8: 7. 012. 1822; Benth. & Hook. Gen. Pl. 3:586. 1883. Rhynchadenia A. Rich. in Sagra, Hist. Fis., Pol. y Nat. Cuba 12 (Fl. Cub. Fanerog. 2) :248, t. 85. 1853. Serrastylis Rolfe, in Gard. Chron. 16:726. 1894. Small, epiphytic, pseudobulbose herbs, the plants reminiscent of a Notylia in habit. Leaves flat, ligular to lanceolate, fleshy or coriaceous. Inflorescences nodding or pendulous racemes, produced from the base of the current pseudo- bulb. Flowers of moderate size or sometimes rather small, on slender pedicels subtended by small bracts. Sepals subequal, free, more or less spreading. Petals similar to the sepals or a little smaller. Lip continuous with the base of the column, erect, deeply 3-lobed, the lateral lobes rather broad, erect, embracing the column, the mid-lobe spreading. Column erect, subterete, wingless, footless, the apex sulcate, the rostellum slender, erect or slightly inclined, the clinandrium broadly cupulate, with fimbriate or dentate margins. Anther erect at the base of (570) 1949] FLORA OF PANAMA (Orchidaceae) 255 the clinandrium, oblong, imperfectly 2-celled, produced into a long appendage embracing the rostellum; pollinia 2, waxy. A small genus of tropical American epiphytes, ranging from Florida and the West Indies to Mexico, Peru, and Brazil. One species is known from Panama. 1. MacRADENIA BRAsAVOLAE Rchb. f. in Bot. Zeit. 10:734. 1852. Epiphytic herbs with narrowly linear to oblong, compressed, monophyllous pseudobulbs 2.5—4.5 cm. long and 0.4—1.0 cm. wide, enveloped below in a few imbricating, papery bracts which soon weather away. Leaves ligular, acute, coriaceous, 6.5—16 cm. long and 1.2-2.2 cm. wide, contracted below into short, slender, conduplicate petioles. Inflorescences usually solitary, arching or pendu- lous, many-flowered racemes up to about 25 cm. long, but often less. Flowers of moderate size, on slender pedicels, subtended by inconspicuous, linear-lanceolate, acuminate bracts. Sepals subequal, free, spreading, reddish brown with trans- lucent green margins, the dorsal sepal lanceolate, acuminate, concave, about 18 mm. long and 5 mm. wide, the laterals linear-lanceolate, acuminate, about 20 mm. long and 4 mm. wide, the reverse surfaces with a low central keel. Petals nar- rower than the sepals, reddish brown with translucent green margins, linear- lanceolate, acuminate, about 16 mm. long and 3.5 mm. wide. Lip conspicuously 3-lobed, about 12-15 mm. long and 7 mm. wide when spread out, contracted at the base and continuous with the base of the column, the lateral lobes semi- orbicular, white, erect, and embracing the column in natural position, the ob- liquely subacute anterior margins reflexed, the mid-lobe reddish brown, narrowly linear, acuminate, about 9 mm. long and 1 mm. wide, the disk with a low, fleshy, linear-oblong, obscurely striate callus. Column 6-7 mm. long, subterete, nar- rowed below, gradually dilated above, the margins of the clinandrium conspicu- ously lacerate, the rostellum narrowly linear-acuminate. Guatemala, Nicaragua, Panama, Colombia, and probably adjacent territories. hills east of Panama City, vicinity Juan Diaz, Powell 3401, 3501; without dinde pes Fairchild s. п.; vicinity Pacora, Allen 2357. 85. TELIPOGON HBK. Тыпгосоч НВК. Nov. Сеп. & Spec. 1:335, 7. 75. 1815; Benth. & Hook. Gen. Pl. 3:587. 1883; Kranzl. іп Ann. Nat. Hist. Hofmus. Wien 33:9. 1919. Telopogon Mutis, ex Spreng. Anleit. 21:291. 1817. Thelypogon Spreng. Syst. Veg. 3:742. 1826. Epiphytic herbs without pseudobulbs, the foliaceous stems short or elongate. Leaves usually narrow, distichous, congested or relatively distant, the blades articulated to the conduplicate bases, often deciduous, pergameneous, coriaceous or fleshy. Inflorescences slender, erect, few-flowered, pseudoterminal racemes pro- duced from the uppermost leaf axils and usually exceeding the leaves in length. Flowers usually large in relation to the size of the plant, rarely small, on slender (571) | [Vor. 36 236 ANNALS OF THE MISSOURI BOTANICAL GARDEN pedicels subtended by small, inconspicuous bracts. Sepals subequal, free, broadly spreading, narrow, usually almost completely hidden by the petals and lip. Petals very broad, spreading, much exceeding the sepals in width, distinctly veined. Lip sessile at the base, broadly spreading, usually undivided, similar to the petals or a little broader, rarely smaller, prominently radiate or reticulate-veined. Column very short, stout, wingless, footless, densely setosé or hispid on all sides or some- times only at the apex, the rostellum terminal, erect, usually prominent. Anther erect, applied to the rostellum, distinctly 2-celled; pollinia 4, waxy. Perhaps 60 species of small, tropical highland epiphytes, ranging from Costa Rica to Brazil and Peru. Two species are known from Panama. a. Lip more than 2 cm. wide, radiate-veined, without secondary transverse nerves 2. T. RADIATUS aa. Lip less than 2 cm. wide, reticulate-veined, with distinct secondary transverse nerves 1. T. DENDRITICUS 1. TELIPOGON DENDRITICUS Rchb. f. Otia Bot. Hamb. 1:6. 1878. Dwarf, epiphytic herbs with short foliaceous stems 3 cm. tall (in our speci- men). Leaves linear-lanceolate, acute, subcoriaceous, the blades 1.5-2 cm. long and 0.25—0.3 cm. wide, contracted below into a short, sheathing petiole. Inflores- cences 1- to 2-flowered scapes up to about 6 cm. long, produced from the sub- terminal leaf axils. Flowers in our specimen about 2.5 cm. in diameter, pale greenish yellow, veined brown. Sepals narrowly triangular, acuminate, apparently 3-nerved, about 1 cm. long and 0.3 cm. wide. Petals incomplete in our specimen, apparently broadly rhombic, 9-nerved, probably about 1.2 cm. long and 1.2 cm. wide. Lip similar to the petals, transversely rhombic-ovate, obtuse, minutely apiculate, about 1.5 cm. long and 1.9 cm. wide, distinctly reticulate-veined, ap- parently with 19 primary veins and many secondary transverse nervules, the disk minutely puberulent. Column densely setose, the slender rostellum prominently projecting, as in the genus. Panama and Colombia. CHIRIQUÍ: vicinity Bajo Chorro, headwaters of the Río Caldera, 1900 m., Woodson t$ Schery 694. The vegetative portion of our single specimen is very small, but the flower, although lacking the apices of the petals, seems to compare very well with the type description. The species apparently is allied to T'elipogon parvulus of Costa Rica, but differs in the larger flowers and the distinct reticulate venation of the lip. 2. TELIPOGON RADIATUS Rchb. f. іп Linnaea 41:70. 1877. Dwarf, caespitose, epiphytic herbs, the distichous, congested, foliaceous stems 3.5-8 cm. tall Leaves elliptic-lanceolate to ligular, acute, subcoriaceous, the blades 2-5.5 cm. long and 0.5—1 cm. wide, contracted below into short, con- duplicate, imbricating petioles. Inflorescences 1 ог 2 slender, erect, few-flowered racemes up to about 9 cm. long, produced from the subterminal leaf axils. Flow- ers averaging about 3.5 cm. in diameter, golden yellow striped with rich brown, (572) 1949] FLORA OF PANAMA (Orchidaceae) 237 the elongate, slender pedicels subtended Бу very short, inconspicuous, narrowly triangular bracts. Sepals narrowly triangular, acuminate, 3-nerved, 1.2-1.5 ст. long and 0.35-0.4 ст. wide, the central nerves thickened and carinate on the outer surfaces. Petals broadly rhombic-ovate, shortly and distinctly acute, much exceeding the sepals in width, 9- to 11-nerved, 2—2.5 cm. long and 1.8—2.1 cm. wide. Lip subequal to the petals, the sessile base adnate to the base of the column, the blade transversely rhombic-ovate, obtuse, minutely apiculate, 19- to 21-nerved, the nerves radiate without secondary transverse nervules, about 2 cm. long and 2.2—2.7 cm. wide, the disk with a short, rounded, densely pilose callus. Column about 5 mm. long, densely setose, the slender rostellum prominently projecting. Panama, Colombia, and Peru. CHIRIQUÍ: vicinity Casita Alta, Finca Lérida, eastern slopes of Chiriqui Volcano, 1500-2000 m., Woodson, Allen & Seibert 961. Our specimens differ from the combined concepts of Reichenbach f. and Kránzlin in the slightly smaller, apiculate lip, but otherwise appear to be identical. Evidently closely allied to Telipogon ampliflorus C. Schweinf. of Costa Rica, but differing in the radiate, rather than reticulate, venation of the lip and in the much тоге. densely setose column. 86. DICHAEA Lindl. DicHAEA Lindl Сеп. & Spec. Orch. РІ. 208. 1833; Benth. & Hook. Gen. РІ. 3:556. 1883; Kränzl. in Engler, Pflanzenr. IV, Fam. 50 (Heft 83):33 1923. Fernandezia R. & P. Fl. Peru & Chil. Prodr. 123. 1794 (in part). Epithecia Knowl. & Westc. Fl. Cab. 2:167, f. 87. 1838. Epithecium Benth. & Hook. Gen. Pl. 3:529, 1239. 1883, sphalm. Dicbaeopsis Pfitz. in Engler & Prantl, Pflanzenfam. 29:207. 1888. Epiphytic herbs without pseudobulbs, the erect or pendulous, monopodial, foliaceous stems enveloped in the conduplicate, distichously imbricating leaf bases. Leaf blades short to elongate, pergameneous to coriaceous, articulated and ultimate- ly deciduous, or connate with the sheathing bases and persistent. Inflorescences 1 to several short, 1-flowered scapes produced from the axils of the leaves. Flowers relatively small. Sepals subequal, free, spreading, the laterals rather obliquely inserted, the bases sometimes forming a mentum with the column foot. Petals subequal to the sepals or narrower. Lip affixed to the base of the column, usually clawed, rarely sessile, the blade usually 3-lobed, often more or less anchoraeform when spread out, infrequently entire, the lateral lobes (if present) triangular to linear, short or elongate, usually retrorse in natural position, sometimes reduced to acute angular projections at the base of the blade, the disk usually without a callus. Column short, erect, wingless or rarely narrowly winged, the margins of the clinandrium often denticulate, the under-surface of the column sometimes with a glabrous or pubescent infra-stigmatic ligule, the base of the column pro- duced into a short foot. Capsule ovoid or oblong, muricate, setose or smooth. (573) [Vor. 36 238 ANNALS OF THE MISSOURI BOTANICAL GARDEN A difficult genus of tropical American epiphytes, ranging from Mexico and the West Indies to Brazil and Peru. Five species are known to occur in Panama. a. Leaf blades not articulated, not deciduous. b. Leaf margins densely Саке. 1. D. СПЛОГАТА 3. D. MURICATA bb. Leaf margins not ciliate aa. Leaf blades articulated to the sheathing bases, deciduous. b. Leaves broadly oblong-ligular, obtuse, minutely apiculate, 8 mm. SR ON MN as а na adea р 2. D. Моккип bb. Leaves narrowly linear-lanceolate, acute or acuminate, 6 mm. wide or less c. Leaves 4 cm. long or less, usually of alternating groups of unequal о ОЛЕНИ Ре ЧАНАК ОНИН ОЛАН eta cea 4. D. PANAMENSIS cc. Leaves more than 6 cm. long, all of about equal length................ 5. D. Power Ln к ГИСНАЕА СПЛОТАТА Rolfe, in Kew Bull. 83. 1917. Small, epiphytic herbs, the distichous, often branching, densely foliaceous stems up to about 15 cm. tall and 8-10 mm. wide. Leaf blades elliptic-oblong, acute, distinctly keeled below, the margins densely ciliate, 2.5-3 mm. long and 2.5-3 mm. wide at the base, the blades not articulated to the sheathing bases, not deciduous. Inflorescences slender, 1-flowered scapes 4—6 mm. long, produced from the upper leaf axils, the apex with 2 short, spathaceous bracts enveloping the ovary. Flowers small, the sepals and petals pale buff with red-purple spots and bars, the lip white marked with purple. Sepals free, the dorsal sepal oblong, acute, 5.5—7 mm. long and about 2 mm. wide, the lateral sepals ovate, acute to shortly acuminate, rather concave at the base, 6—8 mm. long and 2-3 mm. wide, the outer surfaces of the sepals sparsely verrucose. Petals rather obliquely elliptic-lanceolate, acute, about 7 mm. long and 3 mm. wide. Lip shortly clawed at the base, the claw more or less geniculate in profile and continuous with the base of the column, the blade anchoraeform when spread out, 3-lobed, the narrowly linear, subfalcate, lateral lobes retrorsely incurving, the mid-lobe triangular, broadly acute, the lateral margins more or less erect in natural position, the entire lip (including the claw) about 4.5—6.4 mm. long and 6-8 mm. wide. Column rather stout, wingless, 3—5 mm. long, the lower surface with a short infra-stigmatic ligule. Capsule densely muricate. Costa Rica and Panama. CHIRIQUÍ: without definite locality, 4000 ft., Powell 435. This concept is very closely allied to that of Dichaea bystricina Rchb. f. and may ultimately prove to be conspecific. 2. DicHAEA Мовк15п Fawcett & Rendle, in Jour. Bot. 48:107. 1910. Epithecia Morrisii Schltr. in Orchis 9:26. 1915. Dicbaea Bradeorum Schltr. in Fedde Rep. Sp. Nov. Beih. 19:154. 1923. Epiphytic herbs, the erect or pendulous stems up to about 40 cm. tall, en- veloped in the very broad, conduplicate bases of the distichously imbricating leaves, the plants superficially resembling some of the pseudobulbless Maxillarias. Leaf blades articulated to the sheathing bases and ultimately deciduous below, (574) 1949] FLORA OF PANAMA (Orchidaceae) 239 elliptic-oblong to broadly ligular, obtuse and minutely apiculate, subcoriaceous, 3—7 cm. long and 0.8—1.5 cm. wide. Inflorescences 1-flowered scapes, about 8 mm. long, produced from the upper leaf axils, the peduncle provided at the apex with 2 large spathaceous bracts which completely envelop and conspicuously exceed the ovary. Flowers large for the genus, the sepals and petals pale green striped déep lavender, the lip deep lavender. Sepals lanceolate, acuminate, slightly concave, the margins minutely ciliate, about 11-15 mm. long and 5-7 mm. wide, the lateral sepals rather oblique and a little broader than the dorsal sepal, with a rather indistinct central keel on the outer surface. Petals subequal to the dorsal берді, lanceolate, acuminate, slightly concave, about 10-12 mm. long and 3.5-4 mm. wide, the margins minutely ciliate. Lip fleshy, the linear-oblong, slightly arcuate claw narrowed at the base and affixed to the very short column foot, the blade dilated and 3—lobed, more or less anchoraeform, the lateral lobes short to elongate, ligular to subular, sometimes falcately triangular-linear, usually more or less retrorse in natural position, the mid-lobe rather narrowly triangular, usually reflexed, acute or shortly acuminate, minutely papillose on both sufaces, the entire lip (including the claw) about 9—10 mm. long and 5-6 mm. wide. Column very stout, the lateral margins narrowly winged, 4—6 mm. long, the clinandrium very broad, with serrulate margins, the under-surface of the column with a subquad- rate, puberulent, slightly emarginate, infra-stigmatic ligule, the base of the column produced into a very short foot. Capsule densely setose. Jamaica, Santo Domingo, Costa Rica, and Panama. cocLÉ: hills north of El Valle de Antón, 1000 m., Allen 2874. снікюсі: vicinity Bajo Chorro, 6000 ft., Davidson 221 Apparently a very variable species. The collection cited above from El Valle de Antón is rather atypical in the narrower leaves and the much reduced lateral lobes of the lip. 3. Піснавл MURICATA (Sw.) Lindl. Gen. & Sp. Orch. Pl. 209. 1833. Cymbidium muricatum Sw. in No P Act. cig Mcr 6:71. 1799. Dichaea latifolia Lindl. Сеп. & Sp. Orch. РІ. Dichaea Moritzii Rchb. f. i а Kr uidk. en 4: 328. 1858. Dichaea muricata Lindl. В latifolia Lindl. ex Griseb. Fl. Brit. W. Ind. 624. 1864. Dicbaea muricata Lindl. var. Moritzii Cogn. in Mart. Fl. Bras. 35:488. 1906. Dicbaea ovatipetala Schltr. in Fedde Rep du Nov. Beih. 19:266. 1923. Dichaea verrucosa Ames & Жы іп l Orch. 8:83, 1925. Epiphytic herbs with elongate, pendulous, monopodial, foliaceous stems, up to about 60 cm. long, completely enveloped in the persistent, distichous leaves. Leaf blades elliptic-oblong to ligular, obtuse to acute, with entire margins, the under- surface distinctly keeled, terminating at the apex in a minute to relatively elongate apicule, the blades often rather obliquely spreading, not articulated to the sheathing bases, not deciduous, 10—20 mm. long and 4—7 mm. wide. Peduncles slender, 1-flowered, 1—2 cm. long, produced from the upper leaf axils. Flowers of moderate size for the genus. Sepals elliptic-lanceolate, (575) [Vor. 36 240 ANNALS OF THE MISSOURI BOTANICAL GARDEN shortly acuminate, about 7.5 mm. long, 2.5 mm. wide, the outer surfaces usually more or less verrucose. Petals subequal to the sepals, elliptic-lanceo- late, acute to shortly acuminate, about 7 mm. long and 2 mm. wide. Lip anchoraeform in outline when spread out, the broad claw contracted at the base and affixed to the base of the column, the blade abruptly dilated and distinctly 3-lobed, the short, acute or acuminate lateral lobes more or less falcately retrorse in natural position, the mid-lobe shortly triangular, acute or apiculate, the entire lip, including the claw, about 6 mm. long and 3.2 mm. wide. Column stout, about 2.5-3 mm. long, the under-surface with a small infra-stigmatic ligule, the broad clinandrium hooded, as in the genus. Capsule densely muricate. Mexico to Brazil; Cuba to Lesser Antilles. COLÓN: summit of Cerro Santa Rita, 1200-1500 ft., Allen &$ Allen 5112. сос: к slope апа summit of Cerro Valle — 700-800 m., ong 512; vicinity El alle de Antón, 600-1000 m., Allen 1240. снікшюіі: vicinity Casita Alta, Finca Lérida, eastern slopes of Chiriqui Volcano, 1500—2000 m., Woodson, Allen 4 Seibert 834; vicinity Cerro Punta, headwaters of the Rio Chiriqui "Viejo, 2000 m., Allen 1 Most of the specimens cited above are sterile, and the determinations must be regarded as provisional Іп particular, the collections from Chiriquí Province appear quite atypical in the narrower leaves and may possibly represent Dichaea trichocarpa (Sw.) Lindl. 4. DICHAEA PANAMENSIS Lindl. Gen. & i Orch. Pl. 209. 1833. Epithecia panamensis Schltr. in Orchis 9:25 Dichaeopsis panamensis (Lindl.) Schltr. in Beih. [n Centralbl. 36?:519. 1918. Epiphytic herbs with slender, monopodial, often rather flexuose foliaceous stems, 4 to about 18 cm. tall. Leaves 2-ranked on the stems, narrowly linear- lanceolate, acute and apiculate or acuminate, subcoriaceous, often glaucous, usually in alternating groups of unequal length, the blades articulated to the sheathing bases and ultimately deciduous below, 1—4 cm. long and 0.2-0.45 cm. wide. Peduncles filiform, 1-flowered, produced from the axils of the leaves. Flowers small, translucent, white spotted with pink or purple. Sepals free, the dorsal sepal oblong-lanceolate, acute, slightly concave, about 5 mm. long and 2 mm. wide, the lateral sepals obliquely oblong-lanceolate, acute, slightly concave, about 6 mm. long and about 2.5 mm. wide at the base. Petals shorter and broader than the sepals, rather obliquely elliptic-oblong, shortly acute, about 4.5 mm. long and 3.5 mm. wide. Lip sagittate to obovate-spatulate in outline when spread out, the ligular claw slightly arcuate in profile, abruptly contracted at the base and con- tinuous with the short column foot, the blade abruptly dilated and 3-lobed, the short, acute lateral lobes retrorse in natural position, the mid-lobe broadly tri- angular, shortly acute with a short central keel on the lower surface at the apex, the entire lip, including the claw, about 7 mm. long and 6 mm. wide. Column wingless, stout, about 3 mm. long, the apex truncate, the under-surface with a short, infra-stigmatic ligule, the base produced into a very short foot. Anther and pollinia typical of the genus. (576) 1949] P e а ROS | Š va VM, = e. | \ te дра ғы 4 > FLORA OF PANAMA (Orchidaceae) == — an 2 Fig. 212. Dichaea panamensis (577) 241 [Vor. 36 242 ANNALS OF THE MISSOURI BOTANICAL GARDEN Mexico, British Honduras, Guatemala, Honduras, Costa Rica, and Panama. CANAL ZONE: hills between Rio Grande and Puerto Vidal, on the road to Arraiján, 50-150 m., Pittier 2716; Barro Colorado Island, Gatün Lake, 100 m., Wetmore & Abbe 249; Miraflores, Powell 3133; Pedro Miguel, Powell 3033; Rio Indio, Madden Lake area, 70-80 m., Dodge & Allen 17312; Chagres, Fendler 333. PANAMA: without definite locality, Cuming 1202; low thick scrub along the Rio Tecúmen, north of the Chepo road, about 30 m., Hunter & Allen 221; Chiva-Chiva, Powell 3021, 3006; Rio Tapia, Standley 30671; hills east of Panama City, Powell 175; vicinity Paja, Powell 3130, 3240; Chorrera, Powell 3078; Panama Bay, Perlas Archipelago, San José Island, Johnston 400, 1013, 1216, 210. сосі.Е: Bismarck, above Penonomé, 2000-3000 ft., R. S. Williams 441; vicinity El Valle de Antón, 600-1000 m., Fairchild s. п., Allen 1240, A very common lowland species, widely distributed in Central America, 5. Піснавл PoweLLN Schltr. in Fedde Rep. Sp. Nov. Вей. 17:90. 1922. Dichaea Brenesii Schltr. loc. cit. 19:264. 1923. Erect, epiphytic herbs with unbranching, monopodial, foliaceous stems 15—45 cm. tall, enveloped in the conduplicate bases of the many, distichously imbricating leaves. Leaf blades spreading, linear-ligular, acute to acuminate, coriaceous, 6—13.5 cm. long and 0.3—0.6 cm. wide, the blades articulated to the persistent. sheathing bases and ultimately deciduous below. Inflorescences short, slender, 1-flowered scapes about 1.5-2 cm. long, produced from the leaf axils, terminating at the apex in a short, broadly cucullate bract which envelops and exceeds the ovary. Flowers relatively large for the genus, the sepals and petals pale green to white, the lip purple or pale green heavily marked with purple. Sepals free, glabrous, ovate- lanceolate to oblong-lanceolate, acute, 8-10 mm. long and 4 mm. wide. Petals elliptic-ovate, obtuse and apiculate to shortly acute, slightly concave, about 10 mm. long and 4 mm. wide. Lip with a broad ligular claw at the base, the blade dilated and 3-lobed, broadly trulliform when spread out, deeply concave in natural position, the short, obtuse to acute lateral lobes somewhat retrorse, the mid-lobe semi-orbicular when spread out, more or less conduplicate and shortly acute in natural position, the entire lip, including the claw, about 8 mm. long and 8 mm. wide. Column broad, stout, about 4—6 mm. long, the under-surface with a short, minutely papillose, infra-stigmatic ligule. Capsule glabrous. Honduras, Costa Rica, and Panama. PANAMÁ: foothills east of Panama City, Powell 23. COLÓN: Cerro Santa Rita, 1200 ft., Allen & Fairchild 5186, 5103. 87. CAMPYLOCENTRUM Benth. CAMPYLOCENTRUM Benth. in Jour. Linn. Soc. 18:337. 1881; Benth. & Hook. Gen. Pl. 3:585. 1883 (as Cam pylocentron). Todaroa A. Rich. in Ann. Sci. Nat. II, 3:28. 1845. (578) 1949] FLORA OF PANAMA (Orchidaceae) 243 Pseudobulbless, epiphytic herbs with either distichous, monopodial, leafy stems, or stemless and leafless clusters of thickened roots. Leaves oblong or ligular, coria- ceous, sometimes absent. Inflorescences short, usually densely flowered racemes, produced from the axils of the leaves or from the centers of the root clusters. Flowers minute, often more or less distichously arranged on the scape. Sepals sub- equal, free, connivent, or with the apices spreading. Petals subequal to the sepals or sometimes smaller. Lip sessile at the base of the column and produced into an elongate, often recurved spur, the blade entire or 3-lobed, subequal to the sepals in length, often with the lateral margins more or less convolute at the base. Column very short, wingless, the base without a foot. Anther terminal, operculate, іп- cumbent, 2-celled; pollinia 2, waxy. About 30 species of tropical American epiphytes, ranging from Florida, Mexico, and the West Indies to Brazil. Three species are known from Panama, but one is represented only by sterile material. a. Plants entirely leafless, consisting of spreading clusters of thickened roots 3 - 5р. aa. Plants with monopodial, foliaceous stems. . Leaves less than 1.5 cm. long. Highland species 1. C. BRENESII bb. Leaves 4—9 cm. long. Lowland species 2. C. MICRANTHUM 1. CAMPYLOCENTRUM ВВЕМЕЗИ Schltr. in Fedde Rep. Sp. Nov. Вей. 19:268. 1923. Small, epiphytic herbs with erect, distichous, monopodial, foliaceous stems 2-12 cm. tall. Leaves spreading, the blades oblong to elliptic-oblong, rather obliquely acute, coriaceous, about 7-14 mm. long and 3.5-5 mm. wide, articulated to the sheathing bases and ultimately deciduous below. Inflorescences short, lateral, 5—10-flowered racemes produced from the leaf axils. Flowers minute, distichously arranged on the scape. Sepals narrowly lanceolate-ligular, acute, about 1.75 mm. long, the laterals usually rather oblique. Petals subequal to the sepals, rather obliquely linear-lanceolate, acute. Lip 3-lobed, about 1.75 mm. long and 1 mm. wide, the lateral lobes short, obtuse, the mid-lobe narrowly triangular, the base produced into a somewhat laterally compressed, obtuse spur about 1 mm. long. Column very short, typical of the genus. Guatemala, Costa Rica, and Panama. CHIRIQUÍ: vicinity Bajo Mono and Quebrada Chiquero, 1500 m., Woodson & Scbery 2. CAMPYLOCENTRUM MICRANTHUM (Lindl.) Rolfe in Orch. Rev. 11:245. 1903. Angraecum micranthum Lindl. in Bot. Reg. 21: t. 1772. 1836. Angraecum brevifolium Lindl. loc. cit. n. s. 3: sub 2. 68. 1840. Angraecum Lansbergii Reichb. f. in Nederl. Kruidk. Arch. 4:316. 1859. eranthus micranthus Reichb. f. in Walp. Ann. 6:901. 1864 Pise ни Lansbergii Reichb. f. loc. cit. 1864. Campylocentrum panamense Ames, Orchidaceae 7:88. 1922 Campylocentrum peniculus Schltr. in Fedde Rep. Sp. Nov. Бе. 17:91. 1922. (579) [Vor. 36 244 ANNALS OF THE MISSOURI BOTANICAL GARDEN MAMAS yii U^ МИ VAS АЧУ LE W МА S К) % 2 I BAM 220 и D іс ‘oor Auh N ЖК V N № W үу 5 "S ey >, к; 2 (Қ) аң АУ Үү nn» ғ “Ж P А d. 2 A 4: @ = Ve ‘A $6 M SÀ J $ ж - | ay" m е” dI. Fig. 213. Campylocentrum micranthum (580) 1949] FLORA OF PANAMA (Orchidaceae) 245 Epiphytic herbs, with more or less flexuose, monopodial, foliaceous stems, up to about 35 cm. long, enveloped in the conduplicate bases of the distichous leaves. Roots alternating with the leaves and equidistantly distributed along the stem. Leaves broadly ligular to elliptic-oblong, obtuse to unequally bilobed at the apex, the blades about 4—9 cm. long and 1.2—2 cm. wide, articulated to the persistent, sheathing bases and ultimately deciduous below. Inflorescences short, lateral, densely flowered racemes 3-4 cm. long, produced from the sides of the stems just below the point of emergence of the roots. Flowers very small, white or yellowish white, distichously arranged on the scape. Sepals of about equal length, linear- ligular, shortly acuminate, slightly concave, the margins of the lower portions connivent, with the apices spreading, 4-4.5 mm. long and about 0.75 mm. wide; lateral sepals usually rather oblique. Petals narrowly ligular, rather concave, acute, subequal to the dorsal sepal in length, about 4 mm. long and 0.75-1 mm. wide. Lip 3-lobed, about equaling the lateral sepals in length, the sessile base affixed to the base of the column and produced into an elongate, obtuse, incurving spur, about 4 mm. long, the short, lateral lobes of the blade acute or semiovate, erect in natural position, about 1.5 mm. wide when spread out, the mid-lobe narrowly triangular or linear-ligular, rather acute, about 2.5 mm. long and 0.75 mm. wide. Cuba, Santo Domingo, Jamaica, Trinidad, Mexico to British Guiana, Brazil and Peru. CANAL ZONE: Las Cascadas, Powell 184, Standley 25748, 20600; drowned forest near Vigia and San Juan de Pequeni, Madden Lake area, 66 m., Dodge, Steyermark & Allen 6582; in woods, near Gattin, Sutton Hayes 988; vicinity Gamboa, 100 m., Allen 3444, Pittier 2612. BOCAS DEL ToRO: Water Valley, vicinity Chiriqui Lagoon, von Wedel 1386, 1451, 1483, 1655, 1707. 3. CAMPYLOCENTRUM sp. There is a specimen in the Ames Herbarium of a leafless species of Campylo- centrum from Panama, the single inflorescence being very immature and without flowers. It seems likely that it represents Campylocentrum fasciola (Lindl.) Cogn. which is widely distributed from Mexico and the West Indies to Brazil. PANAMA: hills east of Panama City, Powell 320. (581) INDEX ТО PART III (JUNCACEAE-ORCHIDACEAE) Numbers represent italicized page numbers A , Abacá, Achroanthes, 234; mianthemifolia, 235; montana, 23 Acineta, 387; oe: 382; densa, 382; fulva, 382; Colmani, 382; Humboldtii, oe Po etn 382; superba, 382; Warscewiczii, 382 Acostaea, 100; costaricensis, 101 жш а, 305; armeniaca, 306; Adi a ‚ 408 Aeranthus ашды, 570; micrantbus, 570 i cornuta, 3e panamana, 14; picta, 14 Alis, ЕТ Н., 337. 460 Allium Albinia, 57; aromatica, 59; exalt tata, 62; en lalis, 50; Renealmia, 62; spicata, 21; spiralis, бо; tubulata, 62 Alstroemeria edulis, 17; birsuta, 16 Amaryllidaceae, I2 Amaryllis cane 22; longifolia var. longi- ora, ; nervosa, 22; rosea, 22; tubi- spat tha а, Ames, Bue 337 Ames, Oakes, 3 Am аура 247 Androsolen, Angraecum (P TP 470; Lansbergii, ^M 2 š. © x a ЕЗ ©" = Anonymos capitatus, 43 Anthericum, 2; apodastanthum, 3; за тшге ioides, 3; maerophylium, 3 3; panamense, 3 Apteria, 46; aphylla, 47; гене 47; setacea, 4 Arbol de Viájero, 48 Arethusa picta, 156; racemosa, 150 Aspasia, pon epidendroides, 501, var. prin cipissa, 501; fragrans, 501; асы. 501; pusilla, 503; Rousseauae, 50I at bottom of pages. Synonyms in italics. Pu ieiunii ie 5 Atamosco Auliza, Y ЖН 257 Aurota, Banana, 45 Banter española, 75 hsea Це 514 Barkeria, 247 22222 Burtii, 429; grandiflora, 417 Bauxia, 35 Beef- steak Heliconia, 51 Beloglottis, 152; costaricensis, 154 Benitzia, d suaveolens, 44 B iana, Bifrenaria, ; p a, 409 ibai, 48; aur iden 56; barqueta, 51; Bibai, 51; E bs Cham pneiana, 51; chocon 55; densa, 50; elongata, %1; „кы, 50; Ici buts, 52; ba. 54; marginata, 54; platystachys, 54; pun- icea, 50; purpurea, 51; reticulata, 50; — 54; rutila, 51; straminea, 56; tortuosa, 53 а 48; acuminata, 55; Bourgeauana, idea, 56; conferta, 50; лым. 56; Матае, 50; bendu 26; vellerigera, 54; villosa, 53; Wagn Blebbentberll 2 Bletia, 230; capitata, 120; Lankesteri, 342; pallida, 340; purpurea, 340; reflexa, 342; F i 3 O 339 Boop, 344; aristata, 345; pachyr- rac Bollea Wend шеше 421 Bomarea, 14; Allenii, 16; Caldasiana var. concolor, 16; rune 15; chontalensis, 16; edulis, 17; edulis var. chontalensis, 16; var. деб НА Bonatea pauciflora, 12 Bothriochilus, 330; macrostachyus, 330 Bracbystele, 152; aguacatensis, 153; guy- anensis, 153 (247) (583) 248 Brassavola, 316; acaulis, 318; lineata, 318; n о chlorops, 509; Gireoudiana, 510; renciana var. longissima, 511; Lewisii 508; аш, 511; longissime, SIT, var minor, 508; parviflora, 500 енге др albe, 436 Bulbophyllum, 344; к= 345; расһуг- ; т, 34 ` Burmannia, 43; capitata, 43 Burmanniaceae, 42 C АЊА 13; blanca, 13 а, б3 Calanthe, 337; mexicana, 330 Calathea, 82; albicans, 04; Allenii, 80; Allouia, 86, var. violacea, 89; altissima, 04; Casupito, Ӛз; chrysoleuca, 03; cylind- rica, 87; dasycarba, 06; discolor, 83; бе esa. 92 srondifols 87; hirsuta, 85; indecora, 89; insignis, 84; желерін 0; кона, е leucocephala, 08; leuco- a, 08; ‚ 83; macrosepala, 87; зани a 85; жиги; 2а: microcephala 04; panamensis, 02; par ‚ $5; picta, 01; quadratispica, 84; villosa 85; Wars- cewiczii, 02 Camaridium, 431; affine, 442; enechnites, 446; Biolleyi, 430; Bradeorum ctenostachys, 446; dendvobioMe, 447; ane altatum, 448; grandiflorum, 440; Jime- пеші, 447; latifolium. 460; minus, 455; nutantiflorum, 464; ochroleucum, 442; simile Campylocentron, 578 Campylocentrum, 578; Brenesii, 570; fasci- ola, 581; micranthum, 579; panamense, 579; peniculus, 570: sp, 581 Caña de Mico, 64 Cañagria, 7 Canna, 74; angustifolia, 76; aurantiaca, 75; coccinea, 75; commutata, 75; densiflora, 75; edulis, 78; elegans, 76; flaccida, 76; flava, 76; floribunda, 75; glauca, 76; indica, 77; indica var. patens, 77; liliiflora, 78; liturata, 70; lutea, 75; maculata, 75; patens, 77; Schlechtendaliana, 76; stricta, 76; sylvestris, 77; Warscewiczii, 7 Cannaceae, 74 orus, 74; indicus, 77; ovatus, 77 Catacbaetum, 350 Catasetum, 358; bicolor, 360; Brenesii, 363; [ Vou. 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN eburneum, 362; gongoroides, 360; Оет- stedii, 363; rostratum, 303; scurra, 367; serratum, 305; suave, 305; viridiflavum, 5; Warczewitzii, 367 Cattleya, 312; Skinneri var. autumnalis, 313 entury Plant, 14 Ceratochilus, 389; 5 301 Chondrorhync 9, 427; aromatica, 421; ostaricensis, 422; dis- color, 423; ИЫН дын 421; lactea, 423; Libscombiae, 424; marginata, 424 Choretis, 24 Chysis, 343; aurea var. maculata, 343 graminea, 36; bumilis, 36; 8 Cleistes, 123; rosea, 123 Clowesia, 350 Coelia macrostachya, 330, var. genuina, 331, var. integrilabia, 331 Coeliopsis, 376; Бус бонда, 377 Coppensia, 51 Coryanthes 402; Albertinae, 403; Hunter- ‚ 403; н 403; Роше li. 409) нат A. Corymbis, ira Alava Corymborchis, 161; iei IÓI Corynantbes, 4 Corytbentes, 402 63; ен 67; Bakeri, 67; cernuus, 585 conicus, 71; cylindricus, 71; formosus, 69; Friedr beni 65; giganteus, 07; hirsutus, od lae 5, 67; laxus, 67; а, 67; lima 5 Wedelions, 4 Malor- ие var. amazonicus, 67; maximus, 67; nutans, 71; Pisonis, юн MM 56; ruber, : var. B. pubescens, 64; spiralis, 69; spiralis, 69; splendens, 67; uniflorus, 73; villosis- simus, 64; Weberbaueri, 67 Craniches, 148; muscosa, 146; 146; stachyodes, 147 с 10 Crinum, 10; Commelini, 20; darienensis, 21; erubescens, 20; Kunthianum, 20; longi- florum, 20 Келу аин, 472; gracilipes, 473; іп- aequisepalum, 473; latifolium, 474; Stand- leyi, 475 Cryptophoranthus, ig acaulis, 208; dotus, 178; Powe Crybloseccus, 5 Cryptosanus, 546; scriptus, 548 Ctenanthe, 05; dasycarpa, 06 2 oligantba, lepi- Cuitlauzina, 402 (584) 1949] INDEX Cuitlauzinia, 492 Curculigo, 10; scorzoneraefolia, 10 Curcuma, 57; americana, 86 Cybelion Cyclobogon, 152; РтазорРу ит, 154 Cyclosia, 354 Cycnoches, 368; Amparoanum, 373; aureum, 360; barbatum, 380; chlorochilon, 370; densiflorum, 373; Dianae, 372; Egertoni- anum, 373; glanduliferum, 373; guttu- latum, 373; pachydactylon, 373; pauci- florum, 373; peruvianum, 373; Powellii, 72; Rossianum, 373; stelliferum, 3⁄3; stenodactylen, 373; Tonduzii, 374; ventri- cosum, 376; ventricosum v gertoni- anum, 373; Warszewiczii, 370 Cymbidium muricetum ‚ 575; О UG CAI 442; busillum, 540; subulatum, 488; t nerve, cie смит, 353 Cymbocarpa, Cypella a E 5; gracilis, 35 Cypripedium са um, po var. согор уаг. puit 116; Hincksi- anum, II7; Humboldt, c add II7; Warszewiczianum, rf 16 Cyrtochilos, 514 Cyrtochilum, 514 Cyrtopera, 347; longifolia, 347; Woodfordii, 347 ы odium, 340; punctatum, 340, var intlegerianum, 349; Saintlegerianum, iy ‘ip speciosissimum, 340; tigrinum, 349; Willmorei, 349; Woodfordii, 347 D Dendrobium album, 436; longifolium, 347; ruscifolium, 206; utricularioides, 478 Deppia, 410 Diacrium, 310; bilamellatum, 312; bilamel- latum var. Reichenbachianum, 312 Dichaea, ; Bradeorum, 574; Brenesii, 576; ciliolata, 574; latifolia, ТА Moritxia, 575; Morrisii, 574; muricata, 575; muri- cata var. latifolia, 575, var. Moritzii, 575; ovatipetala, 575; panamensis, 576; Powellii, 578; similis, 575; trichocarpa, 576; ver- исоза, 57 Dicbaeopsit, 573; нур 576 Dicrypta, 431; i, 445; сте зијан 445; iridifolia, ps ptc Didactyle, 344 Dillon, Gordon W., 337 Dimerandra, 247; sfenopetala, 270 Dimerocostus, 72; strobilaceus, 73; uniflorus, 73 Dinema, 247; baleaceum, 256 249 ов 26; alata, 26; anconensis, 32; Bill- ergiana, 32; borealis, 31; bulbifera, 26; сей var. glabra, 30; cayennensis, 26; ida, 30; macrostachya, 32; pana- mensis, 30; bermollis, 32; pilosiuscula var. panamensis, — polygono „рија 22: rac mosa, 31; sa 20, var. iir en- sis, 29; sede: 28 Standleyi, КЕ tri- fida, 31; pov hylla, Dioscoreacea Diot bonaea ЗР 245; obbositifolia, 245 Distemon, 74 Dracaena, 2 Е Echeandia, 4; prolixa, 5; venusta, 4 Echeandiaea, 4 Echthronema, 36 Eckardia, 38 Elleanthus, 728; aurantiacus, 130; capitatus, 120; hymenophorus, 120; laxus, 132; lini- folius, 132; muscicola, 133; Tonduzii, 130; trilobatus, 130 Encyclia, 247; atropurpurea, 255, var. leu- cantha, 255, var.rhodoglossa, 2553 Brene esit, 262; ‘campylostalix, 260; gravida, 262; Powellii, 261; ramonensis, 261; tessalata, 250; Tonluziana, 262 Endocodon Endogona, 2 Epidanthus, 335; paranthicus, 33Ó Epidendrum, 247; aberrans, 276; abbrevi- atum, 254; Adolphi, 284; Ж ы. 262; Allenii, 270; aloifolium, 264; amandum, Sut anceps, M фоне 255, r. laciniatum, 255; a m, 256; auro- бањи 4 255; о. 260; Бе- nignum, 200; biflorum, 301; bilamellatum, columna, 281; cuspidatum, 257; Cycno- (585) 250 stalix 268; жет 250; dentiferum, 270; dentilobum m difforme, 287, v rmum, 83, . simulacrum, эз; eburneum, 283; ШЫ Ын. 284; Еп- dresii, 284; equitans, 285; equitantifolium, I 286; ibaguense, 286; itatoshylluns, 287; imbricatum, 301, var. angustifolium, 301; incomptum, 287; ion- ophlebium, 261; isomerum, 280; Isthmi, 203; labiatum, 547; latilabrum, 281; lep- rosum, 276; lividum, 310; lockhartioider, 280; lorifolium, 267; leucocardium, 283; macrochilum, 255, var. roseum, 255; mag- nibracteatum, 278; majale, 283; micro- dendron, 280; modestiflorum, 301; Moore- anum, 262; moyobambae, 200; musci- ferum, 273; myodes, 208; nocturnum, 201, var. panamense, 201; nodosum, 316; obesum, 202; Oerstedii, 259; nanidioides 2 M ; рогрруторруват, pa Powellii, 208; prismatocarpum, 265; probi- florum, 200; prorepens, 254; prostratum, 277; udepidendrum, 0; pudicum, 209; punctatum, 349; pusillum, 540; Pygmaeum, 2051 quinquelobum, 207; radicans, 28 аг. chiriquense, 286; R adlkoferianum, pom бт; ramonianum, 205; ramosum, 301, var. angustifolium, 307, var. imbricatum, 301; reflexum, 203; repens 5 ы. Yo Uo с 1 a ^ mn е eH = 3 =p S ° © c с ед B c et ы 2 2 © ; su patens, 290; subulatum, 488; рање cens, 278; tenuiflorum, 276; teres, 308; [Vor. 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN teretifolium, 308, var. Powellianum, 308; 257; volutum, Epitbecia, 573; Morrisii, 574; panamensis, 576 б HR 573 Eriopsis, 3⁄7; rutidobulbon, 378 Eribbilem ШС, 160; Killipii, 160 Ethanium, Eucharis, 23; Bouchei, 24; candida, 23 Eucnemis, 35 Eulophia, 3475 alta, 347; longifolia, Woodfordii, 3 2. 74; flaccida, 76 Eutbonaea, 245; imbricata, 245; oppositi- 347; Evelyna, ; 128; aurantiaca, I 30; capitata, 129; bymenobbora; 1 120 Fabricia, 18 Қы, 552, 573; acuta, 553; robusta, 8 Fourcroea, 13 Fourcroya, 13; gigantea, 13; tuberosa, 13 Franquevillea, 18 Fregia, 133; amebili is, 136 Funium, 1 жене 13; cabuya, 13, var. integra, 13 Furcroea, 13 G Galatbea, 34; coerulea, 35; speciosa, 35 Galeandra Batemanii, 332; Baueri, 332; 3 Galzottís, pies grandiflora, 417 Gethyra occidentali 5, 50 Ghiesbreghtia, 337; реа 330; mexi- Gongora, 305; arm "os a, а уаг. Ы- а, 308; bufonia, 01; жеді Р, 308; leuco- cbila, 308; Рот аға, 308, var. latibasis, 401, var. tricolor, 401; negrita, 308; (586) 1949] INDEX odoratissima, 398; Pouellii, 308; quadri- rnis, 308; r ie 398; ae dei 398; superflua ; truncata var. a 398; unicolor, 308; 308 Goodyera guyanensis, Govenia, 22. ВЫ = ZUM liliacea, 353; i, 353; utriculata, 353 Gymnadenia, 117 Сутповірһоп, 44; panamensis, 45; suaveo- lens, 44 Gynizodon, 513 Gyrostachys, 152; aguacatensis, 153; costa- ricensis, 154; picta, 156; ше. 54 н Habenaria, ti alata, 118; avicula, 110; bi- corni y 120; petalodes, 119; p crantba, 110; repens, 122; setifera, 120; atbacea, 120; Warszewiczii, 110 Н 21 Haemodoraceae, 11 ыл 57; coronarium, 57 Не1сїа, Fic conu: DER К, 51 Heliconia, 46; рона n уж E surentisca, 50; aurco-stria " ps Biabij, 51; 4 I; Bourgaeana, 51; buccinata, 51; cannoidea, 56; caribaea, Jis ; barqueta, EJ Biha Рој hirsuta, 50, var. 50; D 51; Lankesteri, 52; latispatha, 52; longa, 54; marantifolia, 56; margin- ata, js ariae, 50; meridensis, 52; nutans, 53; pe ndula, 54; platystachys, 54; 54; rutila, 51; Seemannii, a 56; Бе, 55: tortuosa, 53; v vellerigera, 54; villosa, 53; Wagneriana, оа 48; amboinensis, 51 Hellenia, 6 Helmia recemosa, 31 Hemp, Manila, 48 Heterotaxis, 431; crassifolia, 445 E 318; Acostaei, 320; be np indeniana, 320; mic js а, мау 320; Powellii, Нехіѕеа, bidentata, 2 ptum 245; Neues: 245 251 мне 12 Hoja blanca, 54 по 247; bseudobygmaeum, 265 Humboltia, 162 Huntleya, ` 427, 429; Вит, 420; cerina, 427; marginata, 424; meleagris, 429 Н ydastylus, 36 H ymenocallis, 24; acutifolia, 25; americana, 25; disticha, 25; Dryandri, 25; littoralis, 5 EA 25 Hym eei 2 oxis, 18; сег, 18; decumbens, 19; decumbens var. mexicana, 18; da 18; humilis, 19; seorzoneraejolis, 1 І lantba, 476; ballidiflora, 478 ian Shot, 75 Indian Inobsis, 476 Ionopsis, 476; pallidiflora, 478; paniculata, , 478; utricularioides, 478; e, 34 Bschacsiphon, 07; leucophaeus, 08; Morlaei, ; Pittieri, IOI; pruinosus, 00 hoch 3293 cbiriquengis, 320; linifolium, ; major, 329; prolifera, 321 Ixia КОТ ЭП асиапда, 6 eae, 2 ] wF x giganteum, I K Katubala, 74 Kefersteinia, 410; costaricensis, 422; lactea, 423 Kegelia, 370; Нон еапа, 380 je 379; Houtteana, 380; Kupperi, O DM I03 Killip I4 Koellens bull H 415; Kellneriana, 416 Kranzlinella, 101; blatyracbis, 230 bal Laelia, 374; Lueddemanii, 315; peduncularis, 14; rubescens, 314 Languas, 57 (587) 252 Leiochilus, 546; retusus, 548 Leo pim 546; cochlearis, 547; labiatus, 547; lii, 548; scriptus, 548 м со 3 + a ° ect с 8 е. -- ° а % м со 2? = с 3 ы < > 4: һы Leucobyle, 481; subulata, 480; Warsceuiczii, 488 EN 492 icinia Liliaceae 2 Limnorchis, 117 Limodorum altum, 347; Lankesteri, 342; purpureum, 340; utriculatum, 353; vere- cundum Lindleyella, 708: ; picta, 400 Lindsayella, 133; amabilis, 137 Liparis, 243; elata, 243; eustachys, 242; fratrum, 242; labiata, 547; tipuloides, 242; Wendlandii, 242 piles 10 Loic арш, ockhartia, pe acuta, 553; amoena, 555, var. triangulabia, 556; iii dp 556; costaricensis, 555; elegans, 560; grandi- bracteat а, 555; lemellosa 558; АНА ida, 553; Pittieri, 550; robusta, 558; triangulabia, $56; variabilis, 550; 8 Luzula, I; gigantea, I; gigantea var. vul- canica, 1; latifolia, 1; baniculata, I Lycaste, 410; aciantba, 449; Bradeorum, 415; brevispatha, 411; Campbellii, 411; candida, Кина ; Dowiana, 412; Filomenoi, 412; сову, 412; рапа, 412; Powelli. 2 tricolor, 415 M Macradenia, 570; Brasavolae, 571 Macrochilus, 513 трупса, 235; Lan macro- stachya, n Bicis P 235; mon- [Vor. 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN tana, 236; Parthonii, 240, var. denticulata, 241; Pittieri, 236; simillima, 238; Soulei, 236; tipuloides, 242; uncinata, 238; Wendlandii, 242; Wercklei, 241; Wood- sonii, 23 Manila Hemp, 48 Maranta, 101; Allouia, 86; arundinacea, ТОТ; e IO4; lutea, micans, 04; protracta, IOI; pruinosa, тоо; silvatica, 101; Warscewiezii, 02 Marantace Ме, И lutea, 104 Marica, 34, 35; caerulea, 35; gracilis, 35; тістен. 29; baludosa, 36; tinctoria, 40 Masdevallia, 770; Allenii, 185; aperta, 232; attenuata, 180; chontalensis, 182; collina, n diantha, 182; pp Resi 182; Lauch- а, 180; Livingstoneana ‚ 182; banamen- ss Sud 2; platyrachis, 230; simula, 185; iu 4 23 майн, 431; Abelei, 464; aciantba, 440; acutifolia, 437; alba, 436; Allenii, 436; Ambaroana, 463; атр Нота, 440; angusti- folia, 467; angustisegmenta, 437; angus- cullata, 446; dendrobioides, 447; diuturna, 448; elongata, 406; exaltata, 448; foveata, 407; Friedrichsthalii, 440; fulgens, ns gatunensis, enchmanni, 467; i audita, 451; о, 466; lactea, 463; 56; Pittieri, 458; planicola, 460; Powellii, 461; pubilabia, 463; buncto-striata, 446; repens, 461; revoluta, 467; rhodosticta, 440; ringens, 463; Roussesuae е, 463; rte. 464; sessilis, 445; stachybiorum, 407; stenostele, 465; tricolor, 415; Turckbeimii, turialbae, 449; umbratilis, 4 405; vagans valenzuelana, 466; variabilis, 467; Wercklei, 468 Medora (588) 19491 INDEX Meliclis, 402 Menadena, 431 Mesocbloa Mui 401; Warscewiczii, 40I Microstylis, 234; blephariglottis, 241; bra НВА 240; Втеп 41; ae 235; ЕНУ 235; ex- 238; fastigiata, 240; hastilabia, 236; SRM 240; ichthiorrhyncha, 235; majantbemifolia, 235; montana, 236; Parthonii, 241, var. denticulata, 241; Pittieri, 236; simillima, 238; tipuloides, 242; Wercklei, 241 Miltonia, 513; Endresii, 513; Roezli, 514; superba, 513 Monacantbus, 359 Monacbantbus, 358 Monostiche, 82 оотеа, 386; irrorata, 386 Moraea, 34; acorifolia, 41; cbimboracensis, 41; gladioloides, 41 Mormodes, 354; atro-purpurea, 356; purpureum, 355; barbatum, 356; colons 355; Hookeri, 356; igneum, 357; macran- thum, 355; Powellii, 355; W endlandii, 355 Morton, C. V., б, 26 Musa, 48; Bibai, SI; Cavendishii, 48; Ensete, 48; paradisiaca, 48; sapientum, 48; textilis, Musaseae, 48 Myanthus, 358 me 05: dasycarpa, 06; соси 104; Hoffmannii, 06; panamensis, М Мате, 26 Nanodes, 24; discolor, 304 Мага о, Nore 4 chrysantha, 382 Bin КИ 4 5,7 хун нт 143 Neolexis, pi 34; qe AU gracilis, 35 Neomoorea, 380; irrora Neottia ae 156; Ps eos 150; orcb- ioides, 158; picta, 156; speciosa, 157 Nidema Boothii, 256, var. triandrum, 264; tonis, 2 ея 18; pratensis, 18 Могуһа, 566; АН ШПНЕ, 567; B 5 arkeri, 253 tamaulipensis, 567; tridachne, 567; tri- sepala, 567; turialbae, 567 О Oakes-Amesia, 504; cryptantha, 565 Obsitila, 2 Odontoglossum, 402; Asbasia, 501; carini- ferum, 403; chiriquense, 494; convallar- 4; Egertoni, labium, 403; Insleayi хат. тастап 496; Oerstedii, 496; Powellii, 408; Schlie- perianum, 408; Warscewiczii, 408, 513 o llewiradenia, 276; centropetala, Olgasis, 514 Olsynium, 36 Oncidium, 514; advena, 519, var. В, Н atum, 517, var. majus, 510; 510; Baueri, ү к Bernoullienum, 546; altissimum, caesium, 533; С ferum, 493; “nha ue] 522; cartbag- ; ar. Swartzii, 525; Dielsianum, 524; ensatum, 527; ensatum, pes fulgens, 2333; Gireoudianum, офи | 535; р duriforme, 536; Papilio var. Kramerianum, pa arviflorum, 1 T Же; sii A 5333 Warscewiczii, 545; Wercklei, 528 Onkeripus, 405, 431 (589) 254 Opbiomeris banamensis, 46 Orchidaceae, 37, то, "ua 337 biza m, 442; anceps, 455; Biolleyi, 439; usi to sey 443; ful- gens, 451; neglectum, 455; Pittieri, 458; Wercklei, 468 Otsithocephalus, 560; bicornis, 561; coch- leariformis, 562; diceras, 561; e 563; inflexus, 563; lanuginosus, 561; m icanus, 563; Pottsiae, jj Powellii, ње НА nduzii, 563; xiphochilus, 561 Ornitbopbora, 548 Orthrosanthus, 41; pesi Зеница 41 Osmoglossum, 492; ттаѓит, 405; а серз, 405; Pisce bei 495; E 495 Palma mer dei т» 12 Рајт І Palmorchis 143; РочеШі, 144; trilobulata, 144 Palumbin Deut НЫ 10 Pancratium, 24; americanum, 25; littorale, 2 Paneguia, 36 ipe eo Sa Paphiopedilum, и caudatum, 116; Hincks- 117; longifolium, 117 Pephiopedium longifolium, 1 117 "id ес 514; n e 532 но 4, 152; со - a, 160; ; Funckiana, 158; temalensis, Es Pringle ei, 158 Pentalobs, 405, 431 Peperidium, 58 Peristeria, 384; elata, 384; Humboldtii, 382 Peristera, 384 426, 427; сегіпа, 427 Phadrosanthus, 51 Pbaedrosant bus айды. 257 Phalanganthus, коо 114; caudatum, 1106; longi- folium, 116 аты 114; Warszewiczianum, 116 Pbrynium waqa sQ 04; cylindricum, 87; floribun 89; microcephalum, 94; 9 villosum, ү, violaceum, во; Warscewiczii, Phollodes е 94 Pbysant bera, Physinga, 247 [ Vor. 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN Physosiphon, 176; minutiflorus, 176 Physurus, 160 Pilumna, 481 ipio. 472 Planera, 63 Plantain, 78 Plantanillo, 46, 75 Plátano, 48 Platantbera, 11 Platyglottis, 327; coriacea, 327 Plat ystele, ТОТ lectronema, 21 Pleiostachya, 07; Morlaei, 101; Pittieri, 101; Pruinosa, I Pleurothallis, 101; соз 210; Alex- nderae, 208; Allenii, Ji ak 104; calyptrostele, 225; canae, 220; cardiochila, 107; cerea, 206; j quensis, 218; cobraeformis, 203; coccinea, Rowleei, 221; ruscifolia, 206; Maio. 229; tim alas, 201; sororoa, 231; specta- bilis, 231; — 223, var. sed HN 223; ubserrata, _ 233; EE у$.205 (590) 1949] INDEX 208; Tuerckheimii, 228; uncinata, 208; valenzuelana ‚ 466; Yiliticsulis, 218; vere- cunda, 214; vittariifolia, 212; vittata, agneri, 220; Williamsii, 104; xanthophthalma, 233; Pogadelphia, 36 Pogonema 2I 5 bulissthes 1 E 387; barbata, 380; gratiosa, 380 ygonastrum Polystachya, 331; cerea, 331; costaricensis, 331; Masayensis, 332; minor, 331; pana- mensis, 331; Powellii, 331 Ponera albida, 322; ametbystina, 322; Bebrii, 322; bilineata, Fisk mescopsis, 3 Ponthieva, 148; Brenesii, 152; X жыш 150; Ephippium, 150; glandulosa, 150; guatemalensis, 150; maculata, 152; ob- longifolia, 150; racemosa, 150; rostrata, Prescottia, 146; oligantha, 146; panamensis, 140; stachyodes, Pseudepidendrum spectabile, 200 Pseuderiopsis, 377 Psittacoglossum, 431 Psychopsis, 514 P eria, 44; suaveolens, 44 Pyrolirion, 21 Pyxa, 03 R Ravenala madagascarensis, 48 Regnellia, 339 iP nee 58; aromatica, 50; ИЕН б ua, 58; chiriquina, 62; conc 60; costaricensis, бт; densiflora, hee ex- 62; mexicana, "бо; occidentalis, 50; duzii, 233; xantbopbtbalma, 233 Rhynchadenia, 570 Rhynchostele, 546 Rhynchostelis, 546 Rodriguezia, 478; cochlearis, 547; compacta, 479; lanceolata, 480; secunda, 480; зе- cunda var. banamensis, 400 Rodwaya, 45 Rolfea, 143; Powellii, 144 S Sagu, 91, 103 Sansevieria, 2 Sarantbe, 95 255 Sarcoglottis, 152; Hunteriana, 156; picta, 156; Powellii, 156; Purpusiorum, I Ep inni orcbioides, 158; plantagineum, 157 Scadi IQ Scaphosepalum, 186; elasmotopus, 186; longi- repens ; panamense, 102 имиња. 318; Acostaei, 320; albida, 22; amethystina, 322; Веһги, 322; БИ- ineata, 325; brachiata, 322; cuneata, 320; dolichophylla, 327; gracilis, 321; labia, 321; Lindeniana a, 320; ongicaulis, 323; prolifera, 320; tenella, 325; unguic- ulata, 321; Wercklei, 3 Schomburghia, 314; Lueddemanii, 315 Schweinfurth, Charles, 337 Selenipedium, 114; chica, 114; longifolium, Lb Serapias speciosa, I57; flava, 161 Serrastylis Sievekinsia, 378; suavis, 378 Sigillaria, 5 Sismatostalix, 548; censis, 551; guatemalensis, 551; an. 551; poikilostalix, 551; о. costa- abortiva, 540; 552 Siphotoma —— | 36; alatum, 36; arizonicum, iricanum, 40; convolutum, 36; fridifolium, 38; Mandoni, 39; mic дир „ 37; micranthum, 375 Moritzianum, pes таш, 40; tinctorium, 40; fingens, 40 Smilacaceae, Ó — 5; laxiflora, 5; nervulosa, 5; panic- ata, 5; thyrsoidea, 5; Gigas, solaz, 6; pelocerdis, IO; chiriquensis, 9; ton Solenipedium, 114 Souza, Spathiger, 247; rigidus, 303; strobiliferus, 306 (391) 256 Spathirachis, 37 Spider lily, 25 Spiranthes, 152; acaulis, 150; aguacatensis, osa, 157; subpandurata, 154; Woodsonii, Spirostalis, 104 Sprekelia, 12 Stanhopea, 380; amoena, 304; aurea, 304; bucepbalus, 301; costaricensis, 390; grav- eolens, 390; inodora, 300; oculata, 301, var. constricta, 300; pulla, 302; venusta, 304; Wardii, 304; Warscewicziana, 300 Stanbopeastrum, 380 Stelis, 261; aemula, 174; Allenii, 164; atro- rubens, 166; barbata, 165; bryophila, 165; cascajalensis, 160; chiriquensis, 165; cin- erea, I65; collina, 175; costaricensis, 165; crescentiicola, 160; cuspidilabia, 169; de- spectans, 105; Endresii, 171; eximia, 160; 169; leucopogon, qua is, 164; Літ, 5; montana, 106; nutanti- Skutchii, 167; Storkii, 172; vestita, 174; Williamsii, 175 Stellarioides, 2 Stemoptera, 46 Stenocoryne, Stenoptera, 145; costaricensis, I45 Stenorrbyncbus, 152; jaliscana, 158; navar- rensis, 157; orchioides. 1 "s deu 2 57 Stromanthe, 103; lutea, 104 Styrandra, 5 T Taetsia, 2 Tangbekolli, 19 Telipogon, pid dendriticus, 572; radiatus, 572 Telopogon, 571 T'etragamestus gracilis, 320 Thalia, 704; altissima, 105; angustifolia, 105; erecta, 104; geniculata, 104 ТРеіуродоп, 571 T hiebautia, 330 Thismia, 45; panamensis, 46 T borvaldsenia, 343 Tigridia, 34 [Vor. 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN Todaroa, 578 Tolumnia, 514 Tomodon, 24 Tovaria, 5; laxiflora, 5; nervulosa, 5; thyrsoi- dea, Trachinema Trichocentrum, 475; panamense, 470 Trichophila, 481 Trichopilia, coccinea, 480; crispa, 480; ymenantha, 480; jamaicensis, 480; K r. alba, 486, var. lepida, 486, var. olivacea, 488; Powellii, 484; suavis, 488; subulata, 488; turialbae, 490 Tridachne, 566; virens, 567 Trigonidium, 460; Egertonianum, 460; Lankesteri, 471; obtusum, 471; криење Tripbora, 123; cubensis, 125; mexicana, 125; agneri, Tripterella, 43; capitata, 43 Tritonia, 34 Trixeuxis, 480 Trizeuxis, 480; andina, 481; falcata, 491 Trophianthus, 499 63 T'siana, erose, 12 T ylochilus, 340 ү Vagnera, 5 Vanilla. ` 126; fragrans, ^d Pittieri, 128; planifolia, 126; pompona, 128 Vellosia, 2 Vellozia, 25; panamensis, 26 Velloziaceae, Vellozoa, 25 Vogelia, 43; capitata, 43 ү Waluewa » 546 Warczewiczella, 410 Warrea, 351; costaricensis, 352; discolor, 423; graveolens, 410; marginata, 424; drata, 424 Warscewiczella, 410; aromatica, 421; calo- p 5а, с: discolor, 423; Wendlandii, ; velat Wanszewiczells 410; marginata, 424 ‚ 48 Williams, КА О., 107, 247, 337 (592) 1949] INDEX Xylobium, 405; сопсауит, 407; elongatum, ; Filomenoi, 407; foveatum, 407; Powellii, 407; stacbybiorum, 407; sub- lobatum, 407 Xyphostylis, 74 Yam, 26 2357 7, Zelmira Zephyranthes 2I; citrina, 21; Eggersiana, 2; Lindleyana, 22; Mesoc blo oa, 22; ner- 0044, 22; rosea, 22; tubispatha, 2 Zingiber, 57 Zingiberaceae, 57 Zygopetalon, 417 Zygopetalum, 417, 427; aromaticum, 421; Burtii, 420; cerinum, 427; discolor, 423; grandiflorum, 417; lacteum, 423; mele- agris, 429; parviflorum, 410; јаре ч 4; Wendlandii, 421 Zygostates costaricensis, 561 (ТТЫ Director Скоков T. Moore HERMANN MM SCHRENE,. Pathologis Jesse M. GREENMAN с Emeritus of the Herbarium тт. W. Doper, таи об EDGAR ANDERSON, Geneticist T E. Woopson, Ја. СУ вите of the Не rbariüm Henry М. ANDREWS, Paleobotanist Number 3 Annals of the ` 2 _ Missouri Botanical | D. Garden’ Xtanical Garden dental in , Scientific Contributions from the | 2 ак Аппа} of the Missouri Botanical Garden Vol. 36 SEPTEMBER, 1949 No. 3 EVIDENCE EXCLUDING MUTATIONS, POLYSOMY, AND POLYPLOIDY AS POSSIBLE CAUSES OF NON-MENDELIAN SEGREGATIONS IN SACCHAROMYCES! BALAJI D. MUNDKUR? Tetrad analysis, the direct genetical characterization of the segregation products of meiosis, has frequently revealed the occurrence of ratios differing from the 1:1 gametic segregations implicit in the postulates of classical Mendelian genetics. Although many of these exceptions to regular segregation have been reported in the fungi, which offer advantages for genetical analysis denied to the geneticist studying higher organisms, the bases for inferring such divergent segregations were not beyond criticism. Previous workers using the fungi as genetic material in- vestigated inadequately marked stocks (generally only one marker was used) and the deduction of truly non-Mendelian ratios was therefore rendered ambiguous by the possible occurrences of chromosomal aberrations. The few markers used were not sufficient to establish hybridity of the stocks analyzed, and, usually being morphological, introduced obvious difficulties in diagnosing the segregants with accuracy. The occurrence of mutations in the segregants during vegetative growth prior to diagnosis was not excluded as a possible cause obscuring an initially Mendelian segregation. The experiments reported in this paper were designed with the above consid- erations in view and present data eliminating the questionable features involved in evaluating non-Mendelian inheritance. In the tetrad analyses of the five hybrids Nus investigation carried out in the Henry Shaw School of Бұрау of Washington University, St. s, Mo., and Southern Illinois Чача. Carbondale, and submitted as a thesis іп partial ful- Sl БЕ Yan requirements for the degree of Doctor of Philo ка іп the Henry Shaw School of Bota This work was supported by ке from Anheuser-Busch, Іпс., and the U. S. Public Health COMMO 2 Department of Microbiology, Southern Illinois University, Carbondale, Illinois. (259) [Vor. 36 260 ANNALS OF THE MISSOURI BOTANICAL GARDEN discussed on the following pages, use has been made of a sufficient number of marker genes to enable inclusion of only legitimate matings in the data. Being biochemical, these markers in addition permit precise diagnosis of the segregants. The analyses, furthermore, have eliminated the possibilities of polysomy and polyploidy as mechanisms causing irregular segregations and have provided evi- dence that the latter may originate in the heterozygote as a result of gene con- versions. TETRAD ANALYSIS OF YEAST HYBRIDS Lindegren and Lindegren (1946) analyzed extensive pedigrees of Saccharo- myces hybrids for the inheritance of abilities to ferment galactose and melibiose. They discovered that, in some pedigrees, the capacity of haplophase clones to fer- ment each of these sugars was controlled by single genes G or ME, respectively. Haplophase clones bearing the recessive alleles (g and те) were incapable of fer- menting galactose and melibiose; that is, they were incapable of producing detect- able gas under the conditions of the Durham tube fermentation test. Іп asci heterozygous for G/g and ME/me, these alleles segregated in a Mendelian manner yielding two fermenter and two non-fermenter haplophase clones from each ascus. In other words, hybrid asci in these pedigrees were of the constitution G G g g (or ME ME me me). In other pedigrees, tetrad analysis of hybrids heterozygous for abilities to fer- ment these sugars revealed occasional high frequencies of non-Mendelian segrega- tions. In such exceptional tetrads, three or four of the single ascospore cultures exhibited the fermenter phenotype instead of the expected ratio of 2 fermenter : 2 non-fermenter clones; that is, some asci in these pedigrees segregated G G G g and G G G G (or ME ME ME me and ME ME ME ME) phenotypes. Non-Mendelian irregularities in segregations of this nature were ascribed (Linde- gren and Lindegren, 1946) to gene-initiated cytogenes. А non-uniform distribu- tion of the cytogenes between homologous alleles was supposed to have occurred preceding spore formation, resulting in the transformation of recessive to ap- parently dominant alleles. Further analyses of irregular segregations were made in a recent book (Linde- gren, 1949) in which the Cytogene Theory was modified by assuming that cyto- genes (called gene-products to prevent confusing them with plasmagenes) were solely gene-produced and were incapable of self-duplication. The multiple-factor hypothesis was shown to be inadequate for the explanation of the non-Mendelian segregations encountered. The Plasmagene Theory elaborated by Spiegelman (1946) is an alternative view of the mechanism of non-Mendelian segregations in Saccharomyces. Accord- ing to this theory, plasmagenes are autonomous (self-perpetuating) cytoplasmic entities capable of producing enzymes, and an unequal distribution of these entities in the cytoplasm at the time of spore formation is held to account for irregular ratios of fermenter: non-fermenter phenotypes. Arguments against the Plasma- 1949] MUNDKUR—NON-MENDELIAN SEGREGATIONS IN YEAST 261 gene Theory have been considered in detail by Monod (1947) and Lindegren (1949). Irregular segregations in heterozygotes with well-marked chromosomes have excluded the view that cytoplasmic components are involved in irregularly segre- gating tetrads. On the basis of these data, Lindegren (19492) has proposed a gene- to-gene transfer of gene components in the heterozygote, and has embodied this concept in the Conversion Theory. This concept envisages the gene as a loosely organized, labile complex capable of sharing part of its essential components with its allele, thus endowing the allele with new or increased capacities. In parting with some of its essential constituents a gene, according to this view, is susceptible to partial degradation, and the deprivation, if excessive, may result in a weakening or even loss of the original capacities of the gene donating the constituents. The occurrence of tetrads with more than the expected number of fermenter segre- gants is explicable on the basis that recessives have been transformed to dominants in the heterozygote. Experiments have adequately shown that the transformed recessive allele is capable of perpetuating the new capacity (acquired from the dominant gene) through successive generations. The acquisition of the capacities of the dominant gene by the recessive allele and the ability of this new character to segregate after crossing is, indeed, equivalent to mutation. The term "conversion" has been used, however, to denote a mutation occurring in and dependent on the heterozygous condition. Mundkur and Lindegren (1949) demonstrated that the phenomenon of long- term adaptation to galactose could be explained as the result of the mutation of non-fermenter clones (g) to the fermenter allele (С). The capacity for mutation was shown to be characteristic of certain lines in which it occurred with unusually high frequency. This raised the question whether the non-Mendelian segregations of galactose and melibiose phenotypes encountered in previous pedigrees were indeed instances of conversion in the heterozygote during meiosis, i.e., true conversions, or merely mutations occurring after the spore had germinated. However, the fact that the extra-fermenters in non-Mendelian tetrads ferment as rapidly as bona fide fermenter segregants makes this phenomenon different from that encountered in "slow"-fermenter pedigrees. Іп the latter case, the variable number of days elapsing before a "slow" clone ferments is indicative of chance mutation; however, an irregular tetrad is diagnosed as such only if more than the expected number of segregants ferment at the same time, usually within 18 hours; or if more than the expected number of segregants fail to ferment over an extended period of time. Tetrad analysis involves the transfer of single-spore, glucose-grown micro- colonies to glucose agar slants. When the four haplophase clones have attained mature growth on the slants they are tested for their mating-type specificity and their abilities to ferment various sugars and synthesize vitamins, amino acids, and nucleic acid components. This initial growth on glucose may not involve selection [Vor. 36 262 ANNALS OF THE MISSOURI BOTANICAL GARDEN of phenotypes, and mutants of non-fermenter haplophase segregants (g or те) to fermenter phenotypes (G or ME) produced in genetically unstable lines should have equal opportunities for growth. When a hybrid ascus heterozygous for G/g and ME/me is dissected, one expects two ascospores to be non-fermenters. However, if mutation occurred on glucose after these two spores germinated, each segregant culture would comprise mixtures of original g and mutant G clones. When this mixed population is used as inoculum in testing for galactose (or melibiose) fer- mentation the mutant cells would be selected and would achieve fermentation. A highly mutable (g to С; те to ME) haplophase segregant is thus apt to be diagnosed as an extra-fermenter although the ascospore giving rise to the mutant clones was genotypically g or те when isolated from the ascus. Similarly, loss mutations (С to g; МЕ to те) are apt to yield fewer than the expected number (two) of fermenter phenotypes when mature glucose-grown clones are tested for fermentation. Situations such as these could obscure Mendelian segregations and lead to the incorrect conclusion that gene transformation had occurred. Originally G G g g spore tetrads (where either allele is highly mutable) might be finally diagnosed as СССС, G G G g, G £ £ g, ot g: gg g. A helpful criterion for determining whether the non-Mendelian segregations are cases of gene conversions or of mutations during growth on glucose would be, of course, the abilities of the four segregants to ferment at the earliest stage of growth, viz., the spore stage. In the absence of a test sufficiently sensitive to detect fermentation by a single spore, other means must be employed. Тһе experi- ments reported herein were designed to test the fermentative abilities of tetrads derived from "converting" pedigrees under conditions in which their multiplication was restricted and therefore before their chances of mutation had obscured the results. Previous pedigrees in which irregular segregations of galactose and melibiose markers were analyzed (Lindegren and Lindegren, 1946, table 2) were derived from crosses in which culture СТА was used as one of the parents. It is a single- spore culture derived from S. carlsbergensis (culture 126, Dr. Mrak) and its use as a parent causes marked disturbances in segregations іп the progeny. It is an а mating type capable of fermenting sucrose, galactose, melibiose, alpha methyl glucoside and maltose; СТА is ап adenine-independent white culture and is thiamin-, inositol-, and pantothenate-independent but is pyridoxine-dependent. The asci descended from СІА analyzed Бу Lindegren and Lindegren were characterized for only mating type specificity and galactose and melibiose fermentations. Моге- over, these asci were produced by inducing mass copulations of haploid parents and the limited number of marker genes made their truly hybrid nature a debatable question. In the present investigation, a sufficient number of marker genes was used to establish hybridity beyond any question. The use of the adenine-dependent, pink 1949] MUNDKUR—NON-MENDELIAN SEGREGATIONS IN YEAST 263 haplophase cultures as mates for СТА and its close descendants afforded an added advantage. Тһе pink cultures were descendants of a white clone treated with nitrogen mustard gas by Dr. E. L. Tatum and Mr. S. E. Reaume. EVIDENCE THAT CONVERSIONS OF FERMENTER GENOTYPES ORIGINATE IN THE HETEROZYGOTE Materials and methods: Five different crosses involving СТА and three of its direct descendants (607, 608, and 609) were made:—C1A 3349; 609 X 3190; 607 X 3168; CIA Х 5859; and 608 X M317. Тһе spores were isolated on natural medium with glucose as a carbon source. Only 34 four- and three-spored viable asci from a total of 87 dissected have been included in the present analyses. After all the four (or three) spores in a tetrad had germinated to produce microcolonies ranging in population from 100,000 to 500,000 cells per colony, each entire microcolony was transplanted and streaked over a small area on a glucose agar slant. This procedure insures a thorough mixing of mutant fermenter sectors that might have arisen during growth of the microcolony. А small num- ber of cells was then immediately scraped off the slant surface and inoculated into molten 1.75 per cent nutrient agar at 40° C. containing 2 per cent galactose as the carbon source. This molten agar was poured over a layer of solidified non- nutrient 3 per cent agar in a sterile Petri plate. А third layer of non-nutrient agar was poured over the galactose agar after the latter had solidified. This method prevents the fermenter colonies in close proximity from being confluent in growth and also minimizes contaminations. The same procedure was employed in making melibiose pour-plates. Fig. 1 describes the method diagramatically. The residual haplophase yeast on the slant was allowed to incubate 24 hours and was then streaked over the entire slant surface to obtain luxuriant growth. This tube was retained as the stock culture. Mature inoculum from the glucose slant was subsequently tested for mating type specificity and fermentative and vitamin- synthesizing abilities. Simultaneous tests for galactose and melibiose fermentation by colonies developing on galactose and melibiose pour-plates were conducted. The data are presented in tables I-V. The capitals G, ME, MG, S, and MA indicate phenotypes fermenting galactose, melibiose, alpha methyl glucoside, sucrose, and maltose, respectively; the corresponding small letters denote the recessive, non- fermenter alleles. The number of days elapsing before a "slow" clone ferments is indicated by subscript numerals. "White cultures are indicated by W and pink ones by P. Adenine-independence is denoted by AD while dependence is shown by a4. The alleles TH /#b, PN / pn, IN /in, PY /py indicate abilities (or inabilities) to synthesize thiamin, pantothenate, inositol and pyridoxine, respectively. EXPERIMENTAL RESULTS The widespread disturbances in segregations of fermenter phenotypes (shown in tables I-V) are noteworthy, since the stocks to which СТА, 607, 608 and 609 were mated had been carefully selected for regular segregations for the characters. | Vor. 36 264 ANNALS OF THE MISSOURI BOTANICAL GARDEN Calle from smear immediat ely transferred inte golectose © © pour- plate, incubated and tested 9) Ге quactese fermentation Haplophase micrecolonies of a single tetrad on Cells fram smear immediately J transferred inlo melibiese MEME c pour» plate, incubated and tesled for melibiose Fermentation glutose» ager droplet $ Inoculum from glucose slant, after luxuriant growth, i 15 tested divectly fov mating type, abi l: dy То ferment myers and synthesize Vitam "m$ Diagram showing the procedure adopted to minimize mutations in single spore SN and bn fermentative capacities during two growth periods of a nes fe em segregan In spite of the extensive non-Mendelian ratios, the truly hybrid nature of the asci analyzed is evident from the regular segregations of at least two marker genes in each ascus. Further support derives from the fact that cultures of C1A, 607, 608, and 609, and their mates are unable to copulate illegitimately and produce asco- spores. The possibility that a high rate of mutation of the non-fermenter recessive genes to dominance or of the fermenter gene to recessiveness would explain the irregularities : segregation was excluded by the fact that the haploid clones to which СТА, 607, 608, and 609 were mated had been derived from pedigrees in which the Ps of the genotypes was pronounced. Further confirmation of the stability of the recessive parents was obtained by the procedure for detecting fermentation mutants described previously (Mundkur and Lindegren, 1949). Мо reversions to melibiose fermentations were detected in fifteen melibiose pour-plates 1949] MUNDKUR—NON-MENDELIAN SEGREGATIONS IN YEAST 265 made with heavy inocula of glucose-grown, melibiose non-fermenter clones; the rates of reversion of galactose recessives were very low. SEGREGATIONS OF GALACTOSE AND MELIBIOSE FERMENTER PHENOTYPES It has been pointed out above that the ambiguity in deciding whether the extra- fermenters in G G G g and G G G G (or ME ME ME me and ME ME ME ME) tetrads originate from gene transformations or from mutations can be resolved by minimizing the chances of mutations in a haplophase segregant. ‘The galactose and melibiose pour-plates made with inoculum obtained directly from the microcolonies provided data on this point. Since the mutation rates observed are usually of the order of 1 X 1077 and the number of cells in a microcolony at the time of pouring plates does not exceed 500,000, the probability of recovering fermenter mutants may be assumed to be rather remote at this stage of growth. This assumption is strengthened, moreover, by the demonstration (cf. above) of the stability of the genotypes in the recessive parents. The colonies in pour-plates of both galactose and melibiose agar are principally of three sizes on the fourth or fifth day after plating. These have been arbitrarily designated as large, medium, and minute colonies. The large colonies range in diameter from approximately 0.8 to 6 mm., while the medium-sized colonies are approximately 0.2 to 0.5 mm. in diameter. The minute colonies are visible as specks in the agar. When a haploid clone is plated out, it may develop into uniformly sized colonies falling in any one of these categories, or a combination of two of these size classes; or it may show no growth at all. In tables I-V when one class alone is indicated for a segregant, it is implied that the colonies in that particular pour- plate are of uniform size. Plate 1 shows melibiose pour-plates of tetrad М532- M535, exemplifying these size classes. The medium- and large-sized colonies are obviously fermenters and consistently corroborate the phenotype of the segregant obtained from the corresponding glucose slant. However, the minute colonies may or may not be fermenters. For instance, on plates where both minute and large colonies occur, single minute colonies inoculated with a micropipette into sugar broth achieve fermentation and confirm the characteristic of the clone from the corresponding slant. Іп other cases, however, minute colonies occurring uniformly in a plate were generally non-fermenters. Evidence that the differences in colony sizes in a fermenter clone result not from differences in fermentative ability but really from differing growth rates was obtained from the following experiment: А suspension of a single minute fermenter colony (clone M533) from a galactose plate was plated out in galactose agar and was found to yield only minute colonies. А galactose pour-plate of a suspension of a single large fermenter colony of the same clone produced both large and minute colonies. Therefore, the minute colonies are slower-growing clones descended from the large-colonied clones. This explains the occurrence of [Vor. 36 266 ANNALS OF THE MISSOURI BOTANICAL GARDEN TABLE I TETRAD ANALYSIS OF CONVERTER BY STANDARD HYBRID ASCI, C14 BEING THE CONVERTER (СТА а С ME MG MA AD(W) TH PN IN py X 3349 ад me mg ma ad (P) TH pn in py) Fermentation by inocula pour-plates Size of colonies rom Gal. plates Ме]. plates in pour-plates Tetrad phenotypes Inocula derived from al. Mel. to to glucose slants plates plates gal. broth те]. broth M528 а G МЕ MG MA AD (W) pn in med lge + +- M529 а С ME MG MA AD (W) pn IN теа тіп d EE М530 а С ME MG MA AD (W) PN IN| med min + Ен М531 а G МЕ МС MA АР (W) PN in| тей ще + + M532 a G МЕ МС MA ad (W) pn i lge 4- + M533 ас ME MG ma ad (W) pn in| тіп. & ве med + + М534 а МЕ MG MA AD (W) PN je ња 4 M535 аз me MG МА AD (W) PN IN min min -- M536 a g ME mg та ad (Р) pn i in me — -- M537a С МЕ mg MA AD (W) pm in min med + de M538 a g me mg MA ad (P) PN I in i — -- M539 a С те МС та AD (W) PN IN тіп тіп + — M540 ? G ME MG MA AD (W) pn IN med min + + M541 2 G МЕ МСМ AD (W) pn IN med min + + M542 ? С ME MG MA AD (W) pn i min + + M543 ? G те» MG MA ad (W) PN іп |med.& min min + + М544 aG me MG МА ad (W) PN IN lge no growth + — M545 a G ME MG MA AD (W) pn IN med med. + + M546 a G me MG MA ad (W) pn med. no growth -- — M547 а С МЕ MG MA АР (W) PN in| med.& lge. med. + -+ M548 a G ME MG ma ad (P) PN in med med + + M549 a G me МС МА ad (P) PN IN| lge min + а M550 a g ME mg та AD (W) pn IN тіп тей = + M551 a g me mg; MA AD (W) pn in min min — — M552 a G ME mg МА AD (W) PN IN| med. &lge. | med.&lge. | + + М553 а С МЕ MG MA АР (W) pn IN| min.& lge. теа. -- E M554 а теа MG MA ad (W) рп in min. no growth +: -- M555 a С me, MG ma AD (W) PN IN min. no growth + — M556 а ME тео та аа (P) pn IN тіп тей i ie Mss7ag ME то MA ad (D) PN min min T " М558 a С me mg ma AD (W) pm IN med min + Tie M560 a С ME mg MA AD (W) PN IN med she + М561 а g те МС ma ad (W) pn іп min no growth — — M562 a G ME MG MA ad (W) PN in ]gë + е M563 a G МЕ МС МА AD (W) т IN| lge be Es ia М564 а С ME MG MA AD (W) PN IN med in. & lge + + М565а G me MG MA AD (W) PN in med no growth + -- М566 а С МЕ МС MA AD (W) PN in | min. & med + + M567 a G МЕ MG МА AD (W PN IN med Ige + + M568 а G МЕ MG МА ad (D) PN IN med med + + M569 a С МЕ MG mwa D(W) pn in med med + + 1949] MUNDKUR—NON-MENDELIAN SEGREGATIONS IN YEAST 267 fermenter colonies of two size classes in the same plate. However, the fact that when only minute colonies occur on a plate, they are generally non-fermenters indicates that the genotype is intrinsic in the spore from which the clone descended. The uniformity in size of pour-plate colonies in a large number of tetrads is, in itself, strong evidence that mutations with regard to fermentative ability did not occur with sufficiently high frequency to be detectable in the microcolony. In determining the fermentative abilities of the minute colonies, large numbers of them were inoculated into the same broth tube to establish their phenotype with greater certainty. This is especially important when the corresponding glucose slant clone is a non-fermenter. The use of a large number of minute colonies insures that corroboration of the non-fermenter phenotype from glucose was not coincidental. Оп the other hand, the extra-fermenters in a tetrad would have been diagnosed as such if only a single mutation (g to G; me to ME) occurred on a slant; or, if there were a single fermenter colony amidst the more numerous non- fermenter colonies and this exceptional one were inoculated into sugar broth to- gether with its sister, non-fermenter colonies. To test for variation among the individual colonies on a plate in the tetrads with extra-fermenters, the following experiment was performed: Two representa- tive tetrads having extra-fermenters were selected (M532-M535; M544-M547) and the clones known to be galactose- and melibiose-fermenters were plated out in glucose nutrient agar directly from the original glucose slant. Ten individual colonies picked at random from each plate were inoculated each into a separate galactose (or melibiose) Durham fermentation tube. All tubes contained gas within 24 hours. This experiment indicates that individual colony variation as regards galactose and melibiose fermentations did not occur in the pour-plate. It also confirms the fact that after an originally recessive, non-fermenter clone has acquired fermentative ability (and thus become the extra-fermenter in the tetrad) this new capacity is not affected by an intervening period of growth on glucose. Tetrads M488—M491 and M500—M503 (table П) exemplify a situation one would expect as a corollary to the cases of the extra-fermenter tetrads discussed above. "Тһе haplophase parents СТА and 607 involved in the crosses analyzed in tables I, III and V are members of the "converting pedigree" and bear the domi- nant С and ME alleles. The occurrence of extra-fermenters in these crosses has been explained above as being due to a transfer of gene material from the G to the $ (or ME to the me) allele. However, 609 (a parent of M488-M503, table II), which is also a member of the "converting pedigree," bore the recessive me allele and was mated to a ME clone (3190) derived from a regularly segregating (ME/me) pedigree. Tetrads M488—M491 and M500—M503 obtained from this cross include more than the two expected me clones, indicating that the recessive allele in the converter, 1.е., in 609, can degrade the ME allele of its mate which descended from a pedigree in which the МЕ/те alleles segregate regularly. At the same time an excess of G phenotypes are produced in the same asci. [Vor. 36 268 ANNALS OF THE MISSOURI BOTANICAL GARDEN Тһе occurrence of three non-fermenter segregants in each tetrad is explained on the following basis: During meiosis the me gene from the converter parent acquires some essential gene component from the homologous dominant allele of its mate (3190). This acquisition is not sufficient to elevate the me allele to func- tional activity, but the degradation of the ME gene is sufficiently severe to negate its normal function. The single fermenter segregant in each tetrad was presum- ably not affected or, at least, was apparently not degraded below the critical threshold. TABLE II TETRAD ANALYSIS OF CONVERTER BY STANDARD HYBRID ASCI, 609 BEING THE CONVERTER (609 а G те MA AD(W) TH IN py pn X 3190 a g МЕ ma ad (P) th in PY PN) Size of colonies Fermentation by inocula Tetrad phenotypes in pour-plates from pour-plates Gal plates | Mel. plates Inocula derived from Gal. ; t о glucose slants plates plates gal. broth | mel. broth М488 а С те MA AD (W) TH in med. no growth + -- М489 а С те МА AD (М) TH ІМ med. no growth + — М490 а С me МА AD (W) ТН ІМ med. no growth -} -- М491:а С МЕ та AD (W) № in med. med. + + М492 а С МЕ MA AD (W) ТН in| med.& іре. med. + + М493 а G me MA ad (W) ТН ІМ med.& lge. min. + -— M494 a G me ma ad (W) tb in| med.& lge. тіп. -- -- М496 а С МЕ МА AD (W) TH I med. lge. -+ + M497 a g me MA ad (W) thb in| nogrowth min. — EE M498 а g me MA ad (W) № in| nogrowth min. — -- М500 а G те MA ad (P) ТН ІМ min.& med. min. + — М501 а С те MA ad (W) № in med. no growth -- -- 502 а МЕ МА аа (Р) ТН іп med. med. + -- M503 а g me MA ad (W) tb in min. min. -- -- | TABLE ІП TETRAD ANALYSIS OF iris BY STANDARD HYBRID ASCI, 607 BEING E CONVERTER (607 a G ME ma пе X 3168 а g me MA ad(P) ) Size of к уын Fermentation by inocula Tetrad phenotypes in pour-plat rom pour-plates Gal. plates | Mel. plates Inocula derived from Gal. Mel. о о glucose slants plates plates gal. broth | mel. broth M508 ? С ME та d (W) med. med. -- + M509 ? g ME та ай (W) no growth med. — + М510 ? G ME MA AD (W) lge. med. FS re M511 ? С ME МА AD (W) lge. med. d + M512 ? G ME ma AD (W) lge. lge. + + 513 ? С ME та ad (ХУ) lge. lge. E + M514 ? g ME MA AD (W) min. med. -- + 1949] TETRAD ANALYSIS ОЕ CONVERTER BY T MUNDKUR—NON-MENDELIAN SEGREGATIONS IN YEAST 269 TABLE IV CONVERTEI (608 a g me mg S MA AD(W) TH IN PN ру X M317 a G ME MG S та ad(P) th in pn py) STANDARD HYBRID ASCI, 608 BEING 7 R Tetrad phenotypes Size of colonies in pour-plates Fermentation b from pour- у; inocula ates plate Gal. plates | Mel. plates to Inocula derived from Ga Mel. о glucose slants plates plates gal. broth | mel. broth M591 a G me МС SMA АР (W) TH IN pm ру med in. + = M592 а С me MG SMA ad (W) № т pn py med. no growth + — M593 а G те mg SMA ай (W) th т рп py med. no growth E = M594 ag me mg SMA AD (W) TH IN pn БУ nogrowth min. — — M595 а G me mg; SMA AD (W) TH IN pn ру min.& med. min. + — M596 a G me MG SMA AD (W) TH IN PN p| med. ín. ЕЗ EE Ms97 ас me mg S МА ad (W) № in рп py min. no growth — а M598 a G me mg SMA ad (W) № in pn фу| тіп. & те. | no growth -- — M599 а gs me MG S MA ad (W) № т Py ру med min. +. — M600 a g me МС SMA АР (W) TH IN іп ру med in. +. — M601 aG me mg S МА ad (W) th in PN py med no growth + — M602 aG те mg SMA ad (W) № т jm py lge. no growth + To M603 a gs me mg Sma AD (W) TH IN PN py med in. + = M604 a G те mgs SMA ad (W) № т pn py Ige. no growth + — M605 ag me MG 5 ma аа (W) № т pn by min no growth — = M606 а gs me тє SMA АР (W) TH IN PN py med min. + E M607 a С me МС SMA AD (W) TH in фт ру med. min. + = M608 a С me mg SMA AD (W) TH IN pm ру med + = M609 ag me mg 5 МА ad (W) № т pn фу med no growt +s -- M610 ag me МС SMA ad (W) № т pn py тіп по growth -- — М611 aG me mg SMA ad (W) № in pn py 1ге. по growth + — M612 a gs те mgs Sma ай (W) № т pn py med. no growth — - M613 а G me mg, SMA AD (W) TH IN т ру lge. min. + — Mél4aG me МС SMA АР (W) TH IN pn py med min. + — M615 аС me МС SMA AD (W) TH IN PN фу lgd sni "m = M616 aG те mg SMA ad (Ж) № т pn py med min + — M617 а т me MG Sma ad (W) № т pn фу med min +, — M618 aG me mg SMA ad (W) № т pm ру lge. min + = M619 a ж me MG SMA ad (W) № in pn py med min -- — М620 аС me MG SMA AD (W) TH IN pm ру 1де. тіп. БЕ — M621 а gs me mg SMA ad (W) № т pn py med no growth + — M622 ag те mg Sma АР (W) TH IN РМ ру тіп по growt — — M623 ag те МС SMA AD (W) TH IN pn ру min in. -- -- M624 aG me mg SMA ad (W) № т pn py lge. no growth -- — M625 а G me mg SMA AD (W) № IN рп py lge. no growth + — M627 аз т С $ то ad (W) tb in pn БУ nogrowth | no growth — — M628 a G me mg SMA ad (W) № in pn ру lge. min. — M629 ag me mg SMA АР (W) th in pn py min. min. — — M630 ag me MG SMA AD (W) TH IN рп ру nogrowth | по growth — — M631 а G me МС SMA ad (W) № т фп фу Ige. no growth + -- M632 aG me mg SMA ай (W) tb т pn ру lge. min. -- -- M633 а" те mg S МА AD (Ж) TH IN PN ру тей min. +: -- 270 [V sz, 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN TABLE V TETRAD ANALYSIS OF А eee BY T мыш HYBRID ASCI, C14 BEING CONVE (СТА a С ME S MG MA AD TH На € by X TP a g ME s MG MA AD TH IN PN PY) Tetrad phenotypes | oo by inocula | from ur-plates Inocula derived from lucose slants | Size of colonies in gal. pour-plates Gal. plates to gal. broth M464 a G ME s MA MG TH ру large + M465 a G ME $ MA MG TH py large + М466 а С МЕ $ МА МС ТН РУ medium — М467 а С МЕ 5 МА МС ТН РУ medium + M468 a G ME s MA MG TH ру medium + M469 a G ME S MA MG TH PY medium + M470 a g ME S MA MG TH ру minute — M471a g ME S MA MG TH PY minute — М472 а С ME s MA MG TH PY large + M473 a G ME S MA MG TH PY large T M474 a G MES MA MG TH py large + М475 а С МЕ 5 МА МС ру large + M476 a G ME S MA MG TH PY medium & large + M477 a G ME S MA MG TH PY medium + M478 a С ME S MA MG TH ру medium + М479 а С МЕ 5 МА МО ТН ру medium + M480 a G ME S MA MG TH PY large + M481 a g ME S MA MG TH PY no growth -- М482 a G МЕ 5 MA MG TH P large UE М484 a С ME s МА MG TH ру edium + М485 а С МЕ 5 МС ТН РУ medium + М486 a G МЕ 5 МА MG TH ру medium + Even more striking instances of gene degradations are evident in clones M591 to M633. In contrast to the mating 609 X 3190, which involved the melibiose gene as the only recessive converter among the sugar markers used, mating 608 X M317 (table IV) involved four heterozygous sugar markers with the converter parent carrying the recessives g, те, and mg. The total absence of progeny capable of producing gas from melibiose in the Durham tubes in all the tetrads derived from this cross is notable, and is perhaps indicative of a marked instability of the dominant gene controlling the fermentation of melibiose. It is also interesting that, with one exception, each tetrad comprises two clones yielding minute colonies in melibiose pour-plates and two showing no growth. Three tetrads from the same cross also include a deficiency of G clones, while three other tetrads include an excess of G segregants. The absence of any 4:0 G:g tetrads is significant. In addition, 10 "slow"-fermenters of galactose were obtained. The converter 608 carried the dominant MA allele, and tetrads obtained by mating this clone to M317 (which carried the recessive, ma) generally comprise ап excess of maltose fermenters (usually 4:0). Similar cases of 4:0 segregations when 1949] MUNDKUR—NON-MENDELIAN SEGREGATIONS IN YEAST 271 the converter parent carried dominant alleles for fermentative ability can be seen in other crosses. The significant point is that the phenomenon is non-specific. "That is, the presence of the recessive gene in a converter parent need not always result in tetrads with an excess of recessive progeny, for it is conceivable that the dominant allele may not necessarily be completely degraded. Тһе recovery of 4:0, fermenter: non-fermenter progeny from crosses of dominant converters by recessive, stand- ard clones suggests that the dominant gene in the converting parent is not de- graded so severely as in cases involving crosses of dominant, standard clones by recessive converters. Converter haploids may thus differ from standard hap- loids in possessing a more pronounced lability of the genotype, although the detec- tion of truly non-Mendelian segregations in crosses involving two standard clones is by no means rare. The data presented above eliminate the possibility that non-Mendelian tetrads arise from mutations during the growth of originally Mendelian tetrads оп glucose. The exclusion of mutations as a possible cause of irregular ratios leaves open poly- somy and polyploidy as possible mechanisms. Rather, the data suggest that ir- regular segregations of the abilities to ferment galactose and melibiose may originate in the heterozygous condition owing to gene conversions. NON-MENDELIAN SEGREGATIONS AT OTHER LOCI The view that non-Mendelian ratios arise from gene conversions in the hetero- zygote rather than from chance mutations has been confirmed above only with respect to galactose and melibiose fermentations. Мо attempt was made to verify irregular ratios of other sugar markers (sucrose, alpha methyl glucoside, and maltose) using pour-plates of tetrads as checks. The widespread irregular segre- gations of these markers suggests, however, that conversion of alleles was operative here also. A few "slow"-fermenter clones are listed in tables I and ГУ. "Slow" fermenta- tion of galactose was analyzed in a previous report (Mundkur and Lindegren, 1949) where it was shown to result from selection in galactose broth of G mutants descended from originally g segregants. In the present analyses, no attempt was made to determine whether the "slow" fermenters are similar to those previously analyzed or result from a delayed phenotypic expression of gene conversions because of the obvious difficulties involved in making such a distinction. Segregations of vitamin-synthesizing abilities were also irregular, exceptional tetrads occurring with higher frequency than are encountered in standard pedi- grees. Table VI summarizes the frequency of irregular tetrads. [Vor. 36 472 ANNALS OF THE MISSOURI BOTANICAL GARDEN TABLE VI NON-MENDELIAN INHERITANCE OF VITAMIN-SYNTHESIZING ABILITIES IN TETRADS SHOWN IN TABLES I TO V. ONLY 4-SPORED AND OBVIOUSLY NON-MENDELIAN 3-SPORED ASCI ARE INCLUDED. Vitamin Number of heterozygous asci Frequency of non- marked for the vitamin Mendelian asci Thiamin 11 4 Inositol 20 7 Pantothenate 19 11 Pyridoxine 5 2 In addition to the disturbances in segregations of sugar markers, the irregulari- ties in inheritance of adenine-synthesizing ability are very pronounced. It has been shown (Lindegren and Lindegren, 1947) that pink haploids result from the in- ability to synthesize adenine and that white clones are generally adenine-inde- pendent. Occasional tetrads with excess of white progeny out of a pink X white mating have been interpreted as being due to depletions of pinks. The pink color was restored to activity following outcrosses. The irregular ratios for the AD(W) /ad (P) alleles in the present analyses are presumably conditioned by the same mechanism. Several tetrads shown in the tables comprised four white progeny out of a pink X white cross but in some, two clones are ad (W) and two are AD(W). This fact further confirms the hybridity of the asci analyzed. Disturbances in mating reactions are also noteworthy. Іп testing for mating type specificity, a heavy inoculum of each of the four segregants from a tetrad is held in glucose broth together with a standard a clone known to copulate vigor- ously with other haploids. А positive mating reaction identifies its mate as ап “а” mating type; in the absence of copulations, the mate is diagnosed as a clone of like mating type, ie., as an a mating type. Copulations are generally effected within a few hours. In the present analyses, mating type specificity segregated 2а/2а. Occasional disturbances in mating reactions were, however, encountered. For instance, tetrad М528-М531 (W W W W) was mated to the standard a clone but no copulations were observed until the fifth day; tetrads М536-М539, and M548—M551 (2P:2 W) had the normal mating strength and copulated vigorously within twelve hours. Other tetrads (table I), none of whose haplophases copulated with the standard a clones even on the fifth day, were subsequently mated with a standard a clone, and two of the haplophases from each of these tetrads copulated vigorously. Still others, M540—M543 (table I) and M508—M514 (table ПТ) mated with neither the а nor the a standard clones. Similar instances of W W W W tetrads failing altogether to copulate are cited in Lindegren (1949) and Mundkur and Lindegren (1949). 1949] MUNDKUR—NON-MENDELIAN SEGREGATIONS IN YEAST 273 The fact that mating strength is normal in 2 P:2 W asci but is impaired or absent in some W W W W asci (obtained from P XX W crosses) suggests that the phenomenon is in some way associated with disturbances at the AD/ad loci. Further investigation of mating type reactions was not undertaken. DISCUSSION Winkler's (1930, 1932) proposal of the Konversions- Theorie was an attempt to explain deviations from the expectations of tetrad analysis which could not be satisfactorily explained by the theory of crossing-over. Winkler’s 1932 paper is the most ambitious attempt at refuting arguments levelled against his theory by Stern (1930, 1931, 1932), who insisted that the deviations cited by Winkler did not invalidate the theory of crossing-over. Winkler's Konversions- Theorie postulates a physiological transition of the gene from one physiological condition to another during the maturation divisions. Ac- cording to him, “monogenic” conversions involve the transformation of the re- cessive gene alone into a dominant one; or a transformation of the dominant alone into the recessive condition. When, however, the recessive gene acquired dominance while, simultaneously, the dominant allele transformed into recessiveness, the con- version was "digenic." The experiments reported in the present paper have justi- fied Winkler's insistence on the necessity of performing tetrad analyses as a means of determining whether irregular segregations arise from conversions or exclusively from cross-overs or other chromosomal aberrations. However, owing to the pre- cision with which the crossing-over theory can predict map distances, his view that conversions can explain all cases interpreted as cross-overs seems far fetched. When adequate tetrad analyses are available it is possible to distinguish those cases in- volving conversions from those involving crossing-over. Monogenic conversions can be detected unambiguously when segregations in а monohybrid tetrad show unequal frequencies of the dominant and recessive genes. It is simpler to assume that what Winkler termed digenic conversions are actually cross-overs since the results of this phenomenon lead so clearly to the prediction to map distances. Zickler (1934) made exhaustive genetical investigations on the heterothallic ascomycete Bombardia lunata. "Тесгай analyses of pedigrees heterozygous for rubiginosa/viridis revealed high frequencies of non-Mendelian ratios among several thousands of asci. Such exceptional asci contained spore tetrads exhibiting 6:2 and 8:0 rubiginosa/viridis segregations. These markers involved spore color and, though morphological, the characters could be distinguished from each other with great precision, in contrast to many morphological markers whose diagnosis is often ambiguous. То explain deviations in segregations from the Mendelian expecta- tions in B. lunata, Zickler invoked the Konversions- Theorie and held that crossing- over cannot explain the monogenic conversions encountered. Walker (1935) analyzed numerous crosses in Neurospora sitophila, and observed significant divergences from Mendelian expectations. Не described perithecia in which the asci exhibited either a 4:4, 0:8, or 2:6 segregation, exclusively; other [Vor. 36 274 ANNALS OF THE MISSOURI BOTANICAL GARDEN perithecia—the so-called “Mischperithecien”—i which all these three ascus types occurred were also found. The high frequencies of irregular segregations he dis- covered are evident from his data for the mixed perithecia: 4:4 segregations — 30 asci (19.9 per cent) 2:6 segregations — 92 asci (60 per cent) 0:8 segregations — 29 asci (19.2 per cent) Perithecia showing only one ascus type: 4:4 segregations — 0 asci 0:8 segregations — 89 asci However, in spite of these discrepancies, Wülker could not decide in favor of any one of the three explanations for divergent asci he suggested as possible: the labile nature of the gene, conversions, and the influences of environmental factors. This, indeed, is not surprising for he investigated only a single pair of markers L/l (the formation of Luftmyzel or its absence) in the non-Mendelian pedigrees. In addition to the non-Mendelian segregations discovered in Bombardia and Neurospora, numerous instances of exceptions to Mendelian ratios are known in the fungi. Such irregularities occur in Coprinus fimetarius (Brunswik, 1926); Ustilago levis (Dickinson, 1928); Aleurodiscus (Kniep, 1928); Phycomyces blakesleeanus (Burgeff, 1928); Ustilago zeae (Hanna, 1929; Christensen, 1931); Sphacelotheca (Rodenhiser, 1932); Sphaerocarpus (Allen, 1930); Hypomyces Ipomoeae (Dimock, 1939). However, in none of these fungi are chromosome maps available; and these are required to distinguish actual non-Mendelian segre- gations from aberrant ratios arising from segregations of polymeric hybrids or from segregations of hybrids carrying duplications. The unique advantages of tetrad analysis are obvious, and it is not surprising that irregular segregations are encountered more frequently in the fungi, where tetrad analysis is feasible, than in higher organisms, where it is not. When higher organisms are used as genetic material, the conclusive detection of irregular segre- gations is either impossible or the chances of detecting true conversions are, at best, greatly reduced. This limitation derives from the fact that since tetrad analysis is not possible in these organisms regular gametic segregations have been assumed to occur. During animal oogenesis, for instance, a regular 2:2 segregation of genes among the nucleus in the egg cell and the three polar bodies is only inferred provided an approximately 1:1 ratio is obtained when а heterozygous female is mated to a recessive male. Similar inferences are also made with regard to microsporogenesis and megasporogenesis in higher plants. These inferences re- garding the ratios of segregations among the gametes can be based only on the re- combinations of the markers in the Еј generation since the products of meiosis are not directly accessible for characterization. Apparent instances of irregular segre- gations can be explained on the basis of ploidy or crossing-over if they can be verified by cytological methods. However, significant deviations from 1:1 segre- gations among the progeny of a hybrid higher plant or animal are assumed to be 1949] MUNDKUR—NON-MENDELIAN SEGREGATIONS IN YEAST 275 due to preferential survival of sperm or eggs carrying the dominant allele, or to some chromosomal aberration. Tetrad analyses of all the nuclei originating from the same mother cell have, in yeast, yielded vital information inconsistent with the assumptions and inferences involved in the genetical analyses of higher plants and animals. А knowledge of linkage groups in Saccharomyces has been of especial usefulness. The necessity for tetrad analysis in evaluating the status of non-Mendelian in- heritance lies in the accessibility to all the haploid products of meiosis for a direct characterization. Unlike higher organisms where the Еј generation is analyzed, tetrad analysis in the fungi is not obscured by dominance effects or by the am- biguities concerning survival of the products of reduction or of competitions be- tween them resulting in selective fertilization. Moreover, when enough markers are employed, recourse to tetrad analysis makes genetical characterizations of large numbers of progeny a relatively unimportant matter. When biochemical criteria are used, as in the present investigation, tetrad analysis acquires an added signifi- cance. Segregations of abilities or inabilities to ferment various sugars, synthesize vitamins and other cell components are gene-controlled and can therefore be diag- nosed without ambiguity. Where the recessive markers used are especially "good" (examples: g, me, ma, mg, ad, tb, in, pn, ру) the segregations are exceptionally clear-cut. TABLE VII EXPECTATIONS OF virgi als AND RECESSIVE PHENOTYPES WHEN DOUBLE DOMINANT OR DOUD ECESSIVE DISOMICS (PP; рр) ARE CROSSED WITH ORMAL, EI CLONES OF OPPOSITE PHENOTYPE Ratio of phenotypes Mating Type of ascus (dominants : recessives) Frequency Pp genotype — P phenotype a) PP PP p фр 2:2 1 b) PP P рр p 3:1 c) PP P p pP 3:1 PP X p d) P PP рр p 3:1 4 o? РР p jP 1 f) P P pP БР 4:0 7 a) РЕ Pp b b 8:2 1 b) Pp P pp b 8:2 1 c) Pp P b pp 2:2 1 хе d)P Pb bb b 2:2 1 e) Pb b bb 2:2 1 f) Р E bb bb E:2 1 [Vor. 36 276 ANNALS OF THE MISSOURI BOTANICAL GARDEN Among the objections raised against a gene-component transfer mechanism as an explanation for non-Mendelian inheritance, polysomy and polyploidy (Leder- berg, 1948) are noteworthy. Such mechanisms, if they were operative in pro- ducing irregular ratios in Saccharomyces, would yield the phenotypes shown in tables VII and VIII. If we designate a dominant phenotype by P, and its alterna- tive, recessive manifestation by р, tetrad analysis of the crosses PP X р (where PP is the disomic! double dominant parent) and pp X P (where pp is the disomic parent) should yield the tetrad types designated in table VII. On the other hand, if a Pp disomic haploid originating from a heterozygous (P/p) diploid were crossed to a standard haploid clone of opposite phenotype the following tetrad types should result: TABLE VIII EXPECTATIONS OF DOMINANT AND RECESSIVE PHENOTYPES WHEN А Рр DISOMIC IS CROSSED WITH A NORMAL HAPLOID CLONE OF CONTRASTING PHENOTYPE Ratio of phenotypes Mating Туре of ascus (dominants : recessives) Frequency РР genotype = P phenotype (a) БР pP p р 2:2 1 (b) PP p pP p 212 1 (c) P p p pP 2:2 1 ка wy wa 2:2 1 (e) p pp БР 2:2 1 2:2 1 (р P bP bP А comparison of the frequencies of ascus types expected оп the basis of polysomy shown in tables VII and VIII with the frequencies actually encountered in the five crosses described (tables I-V) shows obvious discrepancies. The pos- sibility of a рр disomic effecting the irregularities is precluded by the occurrence of frequent deviations in segregation ratios in heterozygous crosses where the con- verter parent carried the recessive phenotype, although the only ratio expected (if a pp disomic parent were involved) is 2:2. The most striking divergences are seen in table IV where a 0:4 melibiose fermenter:non-fermenter ratio is the only one encountered; at no locus is there the required consistent 2:2 segregation in crosses where the converter carried the recessive phenotype. The possibility of a Рр disomic is also negated for the same reason since a 2:2 segregation would invariably be expected if such a disomic were involved in a cross with a normal haploid of contrasting phenotype. On the other hand, there is a significant deviation from the ratios expected if a PP parent were involved. The term "trisomic" describes diploid organisms having an extra chromos some. Since this term was ебе: particularly for normally diploid organisms, the term “disomic” may be used to describe a baploid individual bearing an extra chromosome. 1949] MUNDKUR—NON-MENDELIAN SEGREGATIONS IN YEAST 277 TABLE IX DISTRIBUTION OF TETRADS PRODUCED IN RECESSIVE CONVERTER x STANDARD DOMINANT CROSSES nly Re asci оК for p pati pes " recessive in E. converter parents (tables II and IV) are include he ce dicted tetrad classes makes statistical analysis PADI and excludes ce Војник јот NE in the haploid 2. is involved.) Ratio of Observed frequencies p fermenter:non-fermenter е оеш е Gal. Mel. a Methyl gluc. 2:4 2 0 6 100 0:4 0 9 0 0 13 3 2 3 0 3:1 4 0 0 0 TABLE X SUMMARIZED DISTRIBUTION OF TETRADS PRODUCED vj DOMINANT CONVERTER x RECESSIVE STANDARD CROSSE (Only those hybrids heterozygous for sugar markers (tables I to V) are included. Significant deviations from expected ratios for galactose and melibiose markers exclude polysomy in the ominant converter parents.) Ratio of Observed frequencies fermenter:non-fermenter рассол Сево сб Һепогу Gal. Mel. Malt. |а Methyl gluc. 2:2 4 4 3 1 8.33 3:1 4 5 8 2 22523 4:0 Z: 2 8 4 58.23 1:3 0 0 0 1 0 = 9.8 15:5 51 0 <.001 <.30 P value Polyploidy is only an extension of polysomy in that every linkage group is represented more than once in an ascospore, while in disomy only one linkage group is represented twice in an ascospore. Some markers in the present analyses belong in different linkage groups (for example, mating type specificity and abili- ties to synthesize pantothenate and ferment galactose are determined by genes located in separate chromosomes). However, the fact that in the same ascus some markers segregate in a precisely Mendelian manner while others segregate irregularly is а priori evidence for the exclusion of polyploidy as the mechanism yielding ir- regular ratios. Lindegren (1949а) has shown that the concept of modifying genes is inadequate to explain irregular segregations in yeasts. In the present analyses, polymeric genes affecting melibiose fermentative ability are excluded by the oc- currence of non-fermenter progeny among the offspring of 608, a non-fermenter converter. [Vor. 36 278 ANNALS OF THE MISSOURI BOTANICAL GARDEN 'ТҺе transformed (converted) recessive haplophase clones maintain the trans- formed type through successive sub-culture generations, within the limits imposed by а normal mutation rate; since back mutations of transformed recessives (e. g., me to ME) do not occur in excess of expectation among the segregants recovered from a hybrid, the transformation of ME to me is not of the type already described as depletion mutation. Тһе high frequencies of non-Mendelian segregations discovered in the present work derive from the use of parent stocks especially conducive to evoking irregular ratios; exceptional ratios that can be interpreted only on a basis of gene conversions are, however, by no means rare even in some so-called regularly segregating pedi- grees. The complete loss of ability to ferment melibiose among the offspring of a culture capable of vigorous fermentation of the sugar—a phenomenon not re- corded previously—is difficult to explain on the basis of the Plasmagene Theory, and questions a long-held genetical tenet: the integrity of the gene as a Mendelian unit uncontaminable in the heterozygote. SUMMARY Tetrad analyses of five different crosses of Saccbaromyces haploids with the use of biochemical markers revealed high frequencies of irregular segregation ratios. These irregularities derived from the use of certain haplophase parent stocks which, when crossed to standard haplophase clones, produced marked disturbances in seg- regations in the progeny. Clones effecting such disturbances have been designated as "converters." These findings are contrary to the regular 1:1 gametic segregations expected by conventional Mendelian genetics. In heterozygous crosses, when converter parents carrying dominant, `#ег- menter phenotypes were mated to clones obtained from standard pedigrees, the progeny comprised an excess of fermenter segregants. Converter parents carry- ing the recessive, non-fermenter phenotypes, when crossed with standard clones of contrasting phenotype, yielded tetrads frequently comprising an excess of non- fermenter segregants. Irregularities in segregation of vitamin-synthesizing abilities, and disturbances in mating type specificity were also encountered. These experiments have yielded data that are not amenable to adequate interpre- tation on the bases of the established concepts of Mendelian genetics. Conversion of alleles in the heterozygote effected by gene-to-gene transfer of gene components has therefore been invoked as an explanation for the anomalies discovered. ACKNOWLEDGMENTS I am deeply indebted to Dr. Carl C. Lindegren and Mrs. Gertrude Lindegren for generous facilities and helpful suggestions during the course of this work. I am also thankful to the Trustees of the R. D. Sethna Foundation, Bombay, India, for sponsoring my studies under Dr. Lindegren. 1949] | MUNDKUR—NON-MENDELIAN SEGREGATIONS IN YEAST 279 LITERATURE CITED Allen, C. F. (1930). Gametophytic inheritance in Sphaerocarpus. IV. Further studies on tuftedness and polyclady. Genetics 15:150-188. Врвни H. (1926). Die Reduktionsteilung bei den Basidiomyzeten. Zeitschr. f. Bot 18:481-498. Burgeff, H. (1928). Variabilitat, ' за und Mutation bei Phycomyces Blekesleeenus. Zeitschr. f. ind rbgsl. 49:2 . ind. Christensen, J. (1931). Studies on ae сакано оғ Ustilago zeae. сече pu Zeitschr. 4:129—188. Dickinson, S. (1928). Experiments on the physiology an ee tics of the smut fungi. Calera chara I. Their permanence and segregation. Pro . Roy . Soc. Lond. B 103:547-5 Dimock, A. W. (1939). Studies on ascospore variants of Ера se Ipomoeae. Mycologia 3l: 709— 27 7275 Hanna, W. F. (1929). Were in = physiology and cytology of Ustilago zeae and Sorosporium reilianum. Phytopath. 19:4 Kniep, H. (1928). Die Sexu a ч p ren Pflanzen. Jen Lederberg, Ј. (1948). Problems in microbial genetics. eed it су 2:14 Lindegren, C. C. (1949). Тһе Yeast Cell. Its Genetics and Cyt 22. егі Тағ Publ. Inc. St. Loui ‚ (1949a). Chromosome maps of Saccharomyces. Hereditas Suppl. Vol., pp. 338—355 ------ and 222 С. (1946). Тһе cytogene theory. Cold Spring Harbor Symp. Quant. Biol. т: 115-129 3 ae ` (1947). Depletion mutation in Saccharomyces. Proc. Nat. Acad. Sci. (U.S.) 3:314-318. icd J. p Rd The phenomenon of enzymatic adaptation. Growth Symp. 11:223— Mund B. D., and Lindegren, C. C. ( 9). An analysis of e AN dp did i Thag- term on to galactose by ИЕ зн (А тег. Jour. Bot., in е Н.А: ы co EE and hybridization in а са sorgbi and S. cruenta. Jour. Agr. Res. 45:28 96. s S. (1946). Moke and ud; алас a controlling enzymatic constitution. Cold ring Harbor Symp. Quant. Biol. 11:256-27 T C. (1930). Konversionstheorie und ш ipae Biol. Zentralbl. 50:6 (19315: сов ТУИ Untersuchungen als Beweise für die ы Theorie des Faktorenaustauschs. 1:547-587. -----,( Белі P VIEN TN Ibid. 52:367—379. Winkler, H. (1930). e Konversion der Gene. 186 pp. Jena. "(1932 onversions-Theorie und Austausch-Theorie. Biol. Zentralbl. 52:16 ЕЕЕ ES ersio eu е LEE 935). Untersuchungen über Tetradenaufspaltung bei Ne 2544 нео hes et Е Zeitschr. f. ind. Abst. - u. Vererbgsl. 59:210—248. Zickler, 934). Genetische ААА ae an einem heterothallischen Askomyzeten (Bom- ba s lunata nov. spec.). Planta 22:57 7 [ Vor. 36 280 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION Or РЕАТЕ PLATE I Melibiose pour-plates of tetrad M532-M535. Three plates (left to right) have colonies each of which is individually capable of fermenting melibiose. Inocula derived from glucose slants corresponding to each clone can also ferment melibiose. Pour-plate of M535 has minute, non-fermenter colonies, and the clone derived from the corresponding glucose slant cannot ferment melibiose. Тһе medium-sized colonies of M533 are not the result of crowding. ANN. Mo. Вот. Garb., Vor. C € SIN <) SI ( t + SESW 36, 949 PLATI 1 NEW SPECIES ОЕ LONCHOCARPUS FROM PANAMA! FREDERICK J. HERMANN U. S. Department of Agriculture Three Panamanian collections of Lonchocarpus which Dr. Robert Schery was unable to place among the species known from Central America were recently sent to the writer for study. All of them proved to be novelties for that region, as Dr. Schery had concluded: two of them undescribed species, of which the first is represented also in Costa Rica, and the third a tree of British Guiana, as follows: LowcHocanPus oli$anthus, sp. nov.—Frutex vel arbor gracilis ca. 10 m. alta; folia 5—7-foliolata; foliola oblonga vel elliptica multipunctata petiolulis sub- quadrangularibus; paniculae axillares 3—4 subspiciformes laxifloraeque foliis multo breviores; pedunculi secundarii minimi uniflori (raro biflori); pedicelli 0.5-0.7 mm. longi; flores 8—9 mm. longi, purpurei; calyx cupuliformis, margine integra vel aliquantum undulata; vexillum late oblongum vel oblongo-ovatum, externe dense argenteo-sericeum, margine valde inflexa; alae oblongae; carinae petala plus minusve falcata marginibus inferioribus solum per intervallum brevissimum apici propinquum connatis; stamen vexillare in fenestra et prope apicem columnae fila- mentarum liberum; ovarium 5—6-ovulatum; legumen ignotum. Shrub or slender tree about 10 m. high, 17 cm. in diameter at base, with "dark brown, almost glabrous, shallowly striate bark" (Smith), and terete, glabrous, shallowly and irregularly sulcate branchlets dotted with small but conspicuous lenticels; stipules scale-like, oblong, 1 x 0.7 mm., dark brown, closely appressed; leaves 5—7-foliolate, 18—30 cm. long, the petiole subterete, canaliculate and often sulcate, glabrous or very sparingly strigose, 3.5—6.5 (averaging 4) cm. long, the rachis deeply canaliculate, sparsely strigose to glabrate; petiolules 4.5—6 mm. long, subquadrangular, minutely strigose to glabrous, dark brown to black, more or less canaliculate above; leaflets thin-chartaceous, oblong to elliptic, the blade 6—15 cm. long, 2.5-7 cm. wide, dark green (sometimes paler beneath), faintly but heavily mottled above with purplish brown, copiously strigose when young, at maturity glabrous or minutely and sparingly strigose beneath, multipunctate (each cell with several to many semitranslucent puncta), caudate at the apex, rounded or tapering at the base, the margin minutely crenulate, the lateral veins 9—10 pairs; panicles 3-4, borne singly іп the upper leaf-axils, short (6—10 cm. long), much surpassed by the leaves, slender, subspiciform, loosely flowered; primary peduncles subterete to subquadrangular, glabrous or glabrate, slender (1 mm. or less wide at the base), floriferous to within 1.5—4 cm. of the base; secondary peduncles rudimentary, about 0.5 mm. long by 0.7-1 mm. wide, 1-, rarely 2-, flowered (the second flower commonly aborted); bracts and bractlets similar, squamiform, ovate, 0.7-1 mm. Issued September 30, 1949. (281) [Vor. 36 282 ANNALS OF THE MISSOURI BOTANICAL GARDEN long, densely strigose, the bracts caducous, the bractlets attached near the base of the calyx; pedicels 0.5—0.7 mm. long; flowers 8-9 mm. long, "purple" (von Wedel); calyx cupuliform, 2.5-3 x 3 mm., reddish brown, minutely tawny- strigose, the margin entire or slightly undulate, the teeth, except the broadly deltoid carinal tooth (0.1-0.4 mm. long), obsolete; standard broadly oblong to oblong-obovate, 9 x 7 mm., reflexed, copiously silvery-sericeous without, glabrous and lineate within, "center and margin pale green-yellow enclosing an area of violet" (Smith), deeply emarginate and somewhat cucullate at the apex, the margin strongly inflexed, irregularly truncate at the base, the auricles very short (0.5 mm.), the claw 1.5 mm. long, a short (0.2 mm.) membranaceous crest between each auricle and the claw; wings 9 mm. long (the claw 2.5 mm.), 2.5 mm. wide, oblong, glabrous except for a sericeous median band (0.6 mm. wide), little widened at the blunt apex, the vexillar half of the blade slightly prolonged at the base to form a shallow auricle (0.7 mm.), adnate to the keel near the base; keel petals 8.5 mm. long (the claw 2.5 mm.), averaging 2.5 mm. wide, somewhat falcate, much broadened toward the blunt apex, their lower margins united only at a point below the apex, glabrous except for the lower margin which is in- creasingly sericeous toward the apex; stamens monadelphous, glabrous, the tube fenestrate at the base with the margins of the opening thickened, the vexillar stamen free at the opening (1 mm.) and also for the terminal 3 mm.; anthers versatile but attached near the base, 2-celled, narrowly elliptic, 0.8 тт. long; ovary compressed, linear, sessile or substipitate, densely white-strigose; ovules 5—6; style strigose at base, glabrous above; stigma capitellate; pod unknown. PANAMA: PROVINCIA DE BOCAS DEL TORO: Gray Creek, vicinity of Chiriquí Lagoon, Sept. 8, 1941, H. von Wedel 2634 (МО түре, US). OSTA RICA: PROV. ALAJUELA: open ibade in rain forest alt. 850 m., Villa Quesada, San Carlos Canton, Feb. 21 1939, Austin Smitb H 1613 (F, MO). The small, sericeous flowers of this species, together with the only slightly coalescent keel petals, almost diadelphous androecium, narrowly oblong wings, and inflexed standard, place it in the subdivision of МЕСКОЗСАРНА designated by Pittier as PUBIFLORI. Its nearest ally is L. parviflorus Benth., from which it differs in having leaflets which are twice to three times as large, relatively long (6—10, rather than 2.5 cm.), loosely flowered panicles, and larger flowers (8—9, rather than 6, mm. long). The larger leaflets of L. oligantbus set it off likewise from the related L. atropurpureus Benth., in which the pedicels are also occasionally, though not predominantly, uniflorous, and from this it is further distinguished by having petiolules twice as long, rudimentary secondary peduncles (0.5, rather than 2—3, mm. long), narrower and densely sericeous standard with claw 1.5 instead of 0.8 mm. long, and densely pubescent ovary. LoNcHocanPus calcaratus, sp. nov.—Arbor 9 m. alta, ramulis crassis verru- cosis; stipulae squamiformes, dense strigosae, caducae; folia 9—11-foliolata; foliola subcoriacea, elliptico-lanceolata, epunctata, subtus strigosa; paniculae laterales, 25 1949] HERMANN—NEW SPECIES OF LONCHOCARPUS 283 ст. longae, ахі ргітагіо florigeno simplici, valido, recto; axis secundarius 10-18 mm. longus, gracilis, 5-9 florus; pedicelli gracillimi, 5-6 mm. longi, bracteolis subulatis caducis strigosis prope basim calycis praediti; flores 16-17 mm. longi; calyx late cupuliformis vel cyathiformis, margine subintegra vel aliquantum undulata; vexillum orbiculare prope apicem emarginatum externe sparsissime sericeum; alae cymbiformes margine superiore basi leviter lobata; carinae petala oblongo-falcata ad basim lateraliter calcarata marginibus inferioribus vix connatis; stamen vexillare solum in fenestra columnae filamentarum liberum; ovarium 6-7- ovulatum; legumen ignotum. Tree 9 m. high, with thick, subterete, often warty branches copiously marked with large, coarse lenticels; stipules squamiform, 2 mm. long, membranaceous, densely strigose, caducous; leaves 9- to 11-foliolate, 15-23 cm. long, the subterete petiole 2—5.5 cm. long, shallowly canaliculate, glabrous or glabrescent, the rachis sparsely strigose to glabrate; petiolules 3.5—5 mm. long, verrucose, conspicuously hirtellous, brown or occasionally olive-green, usually deeply but narrowly canalicu- late above; leaflets subcoriaceous, elliptic-lanceolate, the blade 3.5—11 cm. long, 2—4 cm. wide, epunctate, glabrous or glabrescent above, strigose beneath, the apex obtuse, the base cuneate to abruptly acute, about 8—10 of the lateral veins promi- nent, not impressed, the margin entire, indurated; inflorescence lateral, paniculate, 25 cm. long; primary peduncle and rachis stout (3 mm. in diameter near the base), straight, unbranched, angular, very sparingly strigose, floriferous to within 3 cm. of the base; secondary peduncles 10—18 mm. long, slender, strigose, 5- to 9- flowered; bracts squamiform, 1 mm. long, densely strigose, promptly deciduous; pedicels very slender, 5—6 mm. long, strigose, the caducous copiously strigose bractlets subulate, 1 mm. long, attached near the base of the calyx; flowers 16—17 mm. long, "pink" (Allen); calyx broadly cupulate to cyathiform, firmly char- taceous, 3.5—4 x 8 тт., densely sericeous-strigose, the teeth prominent and broadly deltoid in the bud but the margin in anthesis subentire to shallowly undulate except for the apiculate (0.5 mm.) two lower teeth; standard orbicular, 15 x 15 mm., very sparingly sericeous without toward the emarginate apex, otherwise glabrous, the blade truncate to shallowly cordate at the base, the lobes almost obsolete, the cuneate claw covered by two inflexed, fleshy, partly adherent marginal flaps, their free edges meeting in the center; wings 16 mm. long (the claw 4.5 mm.), 7 mm. wide, cymbiform, very sparsely sericeous without toward the apex, adnate to the keel near the base, the vexillar margin abruptly rounded above the claw to form a broad, shallow lobe; keel petals 15 mm. long (the claw 5 mm.), 4 mm. wide, oblong-falcate, their lower margins united for a distance of 2.5 mm., 3.5 mm. below the obtuse to subacute apex, finely sericeous along the lower margin toward the apex, each petal bearing on its outer face an elongate (3 mm. long, 1.5 mm. wide), hollow spur or pocket midway between the margins, beginning 1 mm. for- ward from the claw; stamens monadelphous, the tube laterally compressed, fenestrate at the base, the vexillar stamen free only at the opening (2 mm.); anthers versatile but attached very near the base, 2-celled, ovate-oblong, 0.8 mm. [Vor. 36 284 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN long; ovary linear, laterally compressed, densely white-strigose; ovules 6—7; style essentially glabrous; stigma capitellate; pod unknown. PANAMA: PROV. DE COCLE: infrequent, El Valle, floor to 1800 ft., April 8, 1947, Paul H. Allen 4472 (МО Tyre). The prolonged, several-flowered secondary peduncles of this species in con- Junction with epunctate leaflets allocate it in Bentham's section PANICULATI— a group otherwise unknown from Middle America but comprising a single species (L. praecox Benth.) in Minas Geraes, Brazil, and four species in tropical Africa. Although no material of any species of the section is available for study, it is evi- dent from Bentham's descriptions (even though the 3-line characterization of L. praecox leaves much to be desired) that it is more closely related to the Brazilian species than it is to the African members of the section. The illustration of Lonchocar pus praecox in Martius’ ‘Flora Brasiliensis’ (Vol. 15, pt. 1, /. 105. 1862) depicts a plant with much shorter panicles than those of L. calcaratus, much stouter and more prolonged secondary peduncles (these almost equalling the primary in diameter and length), smaller flowers with the calyx sparsely hirtellous and more prominently toothed, and elliptic-oblong, not at all lanceolate, leaflets. The re- semblance of the present species is actually much closer to the plate in Martius (t. тоб) designated as Lonchocarpus glabrescens Benth. Тһе figure of the in- florescence in this plate presents a stout and elongated primary floral axis with short and very slender secondary peduncles as in L. calcaratus, but such a con- spicuously paniculate inflorescence is not in agreement with Bentham’s diagnosis ("floribus fasciculatis”) nor with modern collections from the valley of the Amazon which seem to be correctly referred to this species in the light of the original description. In these specimens the primary floral axis is decidedly woody with the characteristic rudimentary secondary peduncles of Bentham’s section FascicULATI to which he referred the plant. According to both Bentham’s and modern accounts, Lonchocar pus glabrescens is, moreover, a liana with flowers having a subrostrate, strongly arcuate keel, usually 10 ovules, and more prominent calyx teeth. The pronounced hollow spur in the carinal petals of Lonchocarpus calcaratus appears to be a feature almost unique in the genus, occurring otherwise, so far as the writer knows, only in L. lineatus Pittier of Gautemala. In the latter species the spur is much more shallow, and the two plants have few other characteristics in common. LONCHOCARPUS DENSIFLORUS Benth. A member of Bentham’s section FASCICULATI, a group heretofore not known to be represented in Middle America. The possession of stipellate leaflets by L. densiflorus is an anomalous feature readily setting it off from other species. PANAMA: PROVINCIA DE BOCAS DEL ТОКО: Laguna de Chiriqui and its neighborhood, Nov.—Dec., 1885, John Hart 99 (US) (distributed as L. sericeus) ; Almirante, Sept. 12, 1920, W. W. Rowlee 8 H. E. Stork 1002 (US) (distributed as Andira sp.). CANAL ZONE: vicinity of Mindi, 1 13, 1947, Рам! H. Allen 5110 (МО, МА). А FIRST RECORD FOR THE GENUS QUALEA (VOCHYSIACEAE) FROM NORTH AMERICA (PANAMA)! ROBERT W. SCHERY ОЏАТЕА cymulosa Schery, n. sp.—Arbor 30 m. alta, ramis junioribus teretibus puberulentis, vetustioribus glabris cortice rimosa et longitudinaliter lenticellata, lenticellis numerosissimis fulvis elevatis; foliis simplicibus plerumque oppositis aut suboppositis aliquando alternatis, 10—15 cm. longis 4—6 cm. latis, integris ellipticis, ad apicem abrupte acuminatis, ad basim angustatis brevi-cordatis coriaceis glabris reticulatis inter nervos laterales; nervis lateralibus ca. 5—7 jugis arcuatis marginaliter confluentibus facere nervum costae parallelum 2—5 mm. ab margine, subtus nervis prominentibus; petiolis ca. 7 mm. longis, supra sulcatis, puberulentis fuscis; stipulis glandulis ovatis crateriformibus callosis ad petiolorum insertionem, duabus glandulis rotundatis inferius ad basin petioli decurrentis; inflorescentia 15 cm. longa 8 cm. lataque, terminali subterminalique, paniculis cymularum trichotomarum cinereo- puberulentis parvi-bracteatis vel subglandularibus; floribus magnis roseis, calcye cinereo-pubescenti coriaceo, 2 lobis lateralibus orbicularibus ca. 4—5 mm. longis, brunneis leviter pubescentibus, 2 lobis intermediis obovatis ca. 7 mm. longis, 1 loba interiora ca. 1 cm. longa, extus dense cinereo-pubescenti intus glabra, calcare bursiculato ca. 6 mm. longo ad basin inserto; petalo 1 obcordato glabro, ca. 2 cm. longo et 2.5 cm. lato apice profunde bilobato; stamine 1 crasso glabro, ca. 9 mm. longo, anthera biloculari; ovario rotundato dense cinereo-hirsuto 3-loculari, loculis biovulatis; fructibus ignotis. Tall forest tree, the bark of the young (leaf-bearing) twig puberulent and scarcely lenticellate, the bark of the older twig glabrous and very lenticellate. Leaves simple, usually opposite or subopposite but sometimes alternate, glabrous, elliptic, rather abruptly acuminate, narrowed toward the base and briefly cordate at the extreme base, the lateral veins usually 5—7 pairs and confluent a few mm. from the margin into an undulate marginal vein subparallel to the prominent costa; petioles dark, puberulent, sulcate above; stipules apparently modified (at insertion of petiole) into crateriform "glands" with the elevated callous margins light in color, and with 2 smaller globose prominences a few mm. lower down to the side of the subdecurrent petiole. Inflorescence terminal but its lower subdivisions fre- quently axillary from upper leaves, thrice-compound, paniculate as a whole but the lateral branchings mostly trichotomous and determinate (cymose), bearing at its branches "glands" similar to the stipular ones, the ultimate and penultimate pedicels with small bracts. Flowers large, showy, rose-pink; calyx coriaceous, cinereous- pubescent, 5-parted, the outer (lateral) 2 lobes smaller, flat and lightly pubescent, the next inner 2 more or less coiled into a cylindric form, the innermost markedly curled into a cone-like cylinder about 1 cm. long, bearing at the base a broad but lIssued September 30, 1949, (285) [Vor. 36 286 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN flattened thick spur about 6 mm. long and 4 mm. broad, this rounded basally; petal solitary, obcordate, about 2 cm. long and 2.5 cm. broad, short-clawed at the base, deeply cleft at the apex, glabrous; stamen solitary, about 9 mm. long, glabrous, thickish, inserted to the side of the petal, bearing terminally a large, fleshy, bi- locular anther less than 2 mm. long; ovary densely cinereous-hirsute, completely 3-carpellate, each carpel 2-ovulate; style glabrous, about equalling the stamen; stigma prominent, capitate. PANAMA: DARIEN: headwaters of the Río Chico, June 1947, P. H. Allen 4645 (Mo. Bot. Gard. ТУРЕ). This new species constitutes the first record of the genus north of South America. It is thus another example of a typically Amazonian plant having а northern range extension into the dense forest of eastern (southern) Panama. The species apparently falls into Warming’s section СОТАТАЕ (Fl. Bras.) of the genus, and as far as can be told from the literature is distinct from all of the multitude of Amazonian species. There is nothing even similar to it in the relatively meager collections of Qualea in the herbarium of the Missouri Botanical Garden, but it may resemble certain Brazilian species such as О. rupicola Ducke, to judge from the original description only. SOME PTERIDOSPERM STEMS AND FRUCTIFICATIONS WITH PARTICULAR REFERENCE TO THE MEDULLOSAE! ROBERT W. BAXTER* INTRODUCTION Theophrastus, one of the first to attempt the classification of plants, recognized as his major groups, trees, shrubs, and herbs. Although this division was long ago realized to be an artificial one it nevertheless provided then, as it does today, con- venient categories for different types of plant habit. Since the study of paleobotany involves not only the search for the ancestral types of present-day plants but also attempts to visualize and illustrate the gross appearance of past floras we may still find it convenient to use these major habit groupings in classifying fossil plants. We now know that the Carboniferous forests were made up of the tree-like Pitys, Cordaites, Lepidodendron, Sigillaria and Calamites, which attained diameters of several feet and reached 100 feet or more in height. Growing among these trees were numerous plants of creeping, climbing, and shrubby habit, characterized in general by small stems with little or no secondary growth. It is also generally true that this last group (ferns and seed-ferns) had developed large leaves or fronds (megaphyllous) while the larger tree-lihe plants were generally small-leaved (microphyllous). The term "microphyllous" is used in this paper in a broad sense to include not only the groups lacking leaf gaps but also those living and fossil gymno- sperms (Coniferophyta, Arnold, 1948) in which the seeds are stem-borne (Sahni's (1920) Stachysperms) and the leaves are small, simple, linear or fan-shaped growths, borne in dense spirals or whorls on the trunk and branches. In plant evolution the early development of a single large trunk (tree-like) seems to have some correlation with microphyllous habit, as the development of numerous small branches of equal size (shrubby) may be correlated with megaphyllous form. Accordingly, it seems possible that a classification based on external form and size may, considered in relation to the origin of plant groups, not be entirely without some phylogenetic meaning. Ав might be supposed, the larger tree-like fossils n investigation eiat out с Ew graduate laboratory of the Henry Shaw School of Botany of Washington University and s ted as a thesis іп partial qose of the requirements for the degree of Doctor of "Philo phy in ps Henry Shaw Scho e УВ 2 Assistant Professor of Bot атте ње Капвав, Lawre ?Wlhile it is not our экеа incen han to suggest di possibility of the foregoing n. ments, there is ample evidence t ihe Coniferophyta. masi. md унган апі p been trees and microphyllous since ps origin daiteae in U vonian and Carboniferous times (Arnold, 1948). n jr краја i vos dan dia _trees "(Calamites, Lepidodendron, etc.) are, on the other hand, represented today by only a few EM је, m era in the so-called fern allies. Thus it would seem that the only possible ancestral ч ga- phyllous gymnosperms, and possibly the angiosperms, must be sought in the shr rudis а ог hi the P ç this h group through stem ramifications the Zelome units would be provided for the development of the large frond and broad leaf, which when fertile may have evolved into sporophylls or carpels (Wilson, 1942). Issued September 30, 1949. (287) [VoL. 36 288 ANNALS OF THE MISSOURI BOTANICAL GARDEN have received a proportionate amount of attention and are today relatively well known. On the other hand, while numerous genera and species of the smaller plants have been discovered and described, it is becoming increasingly apparent that the field is open for the discovery of many more. This is due in part to improved techniques and increasing amounts of research material but probably even more to the undoubted greater diversity of the smaller plants. Certainly in many forest areas today, the genera and species of tree-like habit are but a small fraction of the total flora as compared to those of associated shrubs and herbs. In the present paper we intend to deal with some of these newly discovered small plants (or plant organs) which appear to be of pteridosperm affinity. As is well known to all paleobotanists, the fragmentary nature of the material preserved in coal balls makes it necessary to describe the isolated plant organs (stems, roots, leaves, fructifications, etc.), with the hope that subsequent work will show their proper assemblage. Thus the seed-ferns as a group were originally described as the Cycadofilicales on the basis of stem anatomy, which seemed to combine both fern and cycad characters along with a constant association with fern-like foliage. It was not until 1905 that seeds were actually found attached to some of these fronds and the present conception of the groups was created. Even now the possession of seeds is definitely known for only the genus Lyginopteris (Calymmatotbeca Hoeningbausi) and can only be inferred for the numerous other stems which have been carried over into the new classification on the basis of their similar anatomy. Briefly, the characters on which this relationship is based are: small stems, pith or mixed protostele with exarch or mesarch primary wood, sec- ondary wood of characteristic angular (in cross-section) tracheids, reticulate- bordered pitting, usually numerous leaf traces, and an outer cortex containing vertical or horizontal sclerotic bands. Additional similarities and differences are shown in the accompanying table which lists most of the better-known pterido- sperm stem genera. Our new genus (page 289) is validly included through possession of most of the above-listed characters although it lacked leaves and may or may not have had seeds. Its reconstructed habit (plate 5) does, however, show a close flattened branching pattern which offers implications as to the possible megaphyllous de- velopment of large fern-like fronds. Such ramification in a single plane, along with the lack of leaves, is suggestive of the Psilophytales, and it is our opinion that Micros permopteris may constitute a direct link between that primitive group and the larger seed-ferns. The constant small size (5 mm. or less) of its stems indicates that it would at best have been an inconspicuous element of the forest group and must have been either prostrate or climbing. The three new species of Medullosa described are of course generically identifi- able by their polystelic structure as well as by the character of the wood and pitting and, where the latter tissue is preserved, by the fibrous cortex. Here again the evidence of size and form seems to indicate that the habit was either creeping or climbing rather than tree-like. 1949] BAXTER—FOSSIL PTERIDOSPERMS 289 Тһе two new species of Dolerotbeca ате offered primarily as а supplement to Schopf's (1948) comprehensive work on the genus, although we believe our ma- terial does present some basis for additional phylogenetic interpretation as well as possible additional support for the more definite connection of the fructification to the Medullosae. During the latter course of this study several additional small stems of ap- parently primitive pteridosperm character have been discovered which we hope to describe in a future paper. One in particular is of interest since, as іп Micro- spermopteris, it appears to be leafless. Thus at least a part of the undergrowth of Carboniferous times seems to have occupied a position comparable to that of the living fern allies in that both present a retention of characters primitive for their time so that the excellent fossil record of the coal beds may yet provide the missing links from earlier eras just as those primitive living genera (Lycopodium, Equisetum and Psilotum) have done in part for the present-day flora. Microspermopteris aphyllum, gen. et spec. nov. The following description is based on a total of eight stems occurring in three different coal balls. Four stems were followed through one coal ball for approxi- mately 6 cm. and three stems extended through another ball for approximately 4 cm. The eighth stem was isolated ia the third coal ball and was cut as a longi- tudinal section 1 cm. in length. = All material is from the What Cheer Clay Products Co. coal mine, one-half mile west of What Cheer, Iowa. This horizon lies in the Des Moines series of the Pennsylvanian and is accordingly pe Carboniferous age. The stems followed a relatively straight to slightly sinuous course. Mixed among them were numerous eU of their smaller, but otherwise identical branches. “he stems are of a constant small size even when maturity is indicated by considerable secondary growth. “The total diameter including cortex and epi- dermal tissues never exceeds 5 mm., while the single stele averages 2 to 2/ mm. The external form of the stems is quite variable, particularly in the smaller specimens where the transverse configuration is almost “amoeboid” in outline (fig. 3). The more mature specim ns with considerable secondary growth appear oval to circular in cross-section (figs. 1 and 7). The primary tissues consist of a protostele in which the large metaxylem tracheids are divided into groups of 12—14 cells by a radiating network of paren- chyma, the individual strands of which are usually not more than one cell in thick- ness and in longitudinal section are seen to be vertically elongated cells, five to six times as long as wide. The inconspicuous protoxylem groups are situated on the periphery of the primary tissue and appear to be exarch though their exact position is vague due to the very small size of the adjacent innermost secondary wood. The metaxylem tracheids increase slightly in size towards the center, averaging 150 p in diameter as against 50 м for the secondary wood. АП walls of the metaxylem are reticulately bordered pitted with the opposing orifices at a slight angle to one another (fig. 12). 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"шо Z :шәз$ : put o[o3s0301d ive[nSut Tt: Ч: П ГР Suruuni spueq ешАцә ио]Кшоиә]$ put чегиолә(Т aie] 19310 :onssm 913022]5$ uo 313314 рәләрлод ° P 21215 ајдито : -11] ЈО s19U102 шо; ДЫ а -пәтей {2123503024 : 55јреј xo3105 1ouu[| -одејпоцат ‘pədopəaəp әзешІ8І20 s02t13 је рәхтш лејпдиеиј, IPA poom Атерпо>әс : vSO{IN Pa JO 25043 qSrq чао 2-1 * (92 + 5122 420391295 эшэ ZI :шәз$ 03 лерии$ so[onoq rso]|rpayy оз 1epuurs " ) mys skey "(вот yum ешАцәпәлеа (sə]ə3srrəuu) sno19gruoq1t^) 12407 *sə|ə3srrəur шол} | spueq 91701979S |е2122ӘА I 'Ч M put 52 шәүАхезәш 5риғ115 Áitrpisqns nt 2375 ‘5113495405 SIJI J, yam x23102 12310 +=) рәдорләр Ај -чеәл poos Arepuosag одаеј цата Ə|93so1oid рәхїш е *Ч2техЯ ПЕшв Ад popunouins әүәзѕ |еиә2 2818 I иолцојЦ əjonəd pue $99е21 fray X33100 91njonJjs Avi рие poom Áitpuooog poom Ајеипад ләјәштір шәз$ pue sə]ə3s jo 1oquinu әзешіхолай4у еләцә8 шә1$ U193]-poos Jo элозоелеЧо əaneredwop [Vor. 36 292 ANNALS OF THE MISSOURI BOTANICAL GARDEN The secondary wood ranges from 3-4 rows in thickness in the young small stems to around 16 in the larger more mature specimens. Тһе first-formed cells are quite small, being hardly distinguishable in size from the protoxylem. Their diameter increases gradually towards the outside but never approaches that of the metaxylem. Longitudinal sections of the secondary tissue show the radial walls pitted in the same way as the metaxylem, with a dense reticulum of bordered pits, while the tangential walls have fewer scattered pits, with more lineal openings, arranged in irregular rows up and down the central area. Wood rays аге incon- spicuous and relatively sparse. They are small, uniseriate and from 1 to 4 cells in height (fig. 9). А rather frequently observed character of the secondary wood of the mature stems is its unequal development (fig. 7) which is possibly due in some cases to differing cambial activity and in others represents the unfilled gaps left by departing branch steles. The tissues immediately surrounding the wood are seldom well preserved and are represented in all of our specimens as merely a thin dark brown ring of crushed cells. There is no evidence of periderm formation. The cortex is divided into an inner and outer zone of approximately equal thick- nes. The inner part is composed of poorly preserved isodiametric thin-walled cells containing numerous, scattered dark secretory cells. The outer zone is thicker- walled and usually better preserved, consisting of a compact tissue of brick-like cells (hexagonal to circular in transverse section), about twice as long as wide, the elongation being parallel to the axis of the stem. This merges into an outermost tissue which is 2—2 cells thick and differentiated by being made up of cells four to six times as long as broad. Ап unusual character of this tissue is the frequent occurrence of multicellular emergences which are commonly broad at the base, tapering to a sharp point. They are sometimes forked and horn-like (figs. 5 and 55) and may occur singly or in clusters (fig. 8). In length they average .6 mm. There is no evidence that they were glandular like the emergences of Lyginopteris, nor do they appear to have been arranged in parallel rows as has been described for the latter genus (Scott, 1923). Оп the contrary, they are produced more or less indiscriminately with the exception that they are nearly always present at the point of departure of a branch. A feature of the outer cortical zone which is probably the most unique in rela- tion to the comparable pteridosperm genera is the presence of a vertical series of horizontally aligned sclerotic plates instead of the usual vertical strands. These consist of thick-walled stone cells, 2—4 cells deep and 4—6 cells in diameter (fig. 10). In tangential section they are occasionally seen to anastomose with adjoining plates at the same level and with plates above and below, thus appearing as a loosely arranged horizontal network (fig. 11). One of the most difficult problems in interpretation has been the question of whether Microspermopteris bore leaves. As stated earlier, we have followed four of the stems for approximately 6 cm., three for 4 cm., and another for about 1 cm. 1949] BAXTER—FOSSIL PTERIDOSPERMS 293 This produced a close check on nearly 37 cm. of stem, all, of course, not of the same plant. However in this rather respectable amount of material we encountered only five instances of vascular tissue being given off from the main stele. In every case the details are identical and are illustrated for both a young and mature speci- men in figs. 1, 2, 3, 5 and 55. The branch is first evident as a departure from the stele of a complete segment of primary and secondary wood, with the crushed phloem and cambium tissues on its outer, abaxial surface. At this point it consists of 8 to 12 metaxylem cells with possibly 1 to 2 strands of protoxylem, bordered on the outer side by almost the complete amount of secondary wood present at that particular point on the main stele. This results іп a conspicuous gap being produced іп the mature stems and a relatively smaller one in the young stems where the proportion of secondary growth is much les. The angle of departure is acute (around 30^), the trace following this steep course through the cortex for approximately 2 to 3 mm. During this space the secondary wood and enclosing parenchyma tissues completely surround the primary core and produce a concentric bundle identical to the main stele except in size. The bundle then turns sharply outwards and proceeds at right angles to the stem axis for a distance approximately equal to the normal diameter of the stem before again resuming an upward course. Ав shown in fig. 5, the branch, where it projects horizontally out from the stem, almost equals the stem in size and exhibits all of its characteristic sclerotic plates and cortical zones. Ас no time does the bundle show any tendency to subdivide further, while a transverse section of one of the branches at a point around 4 mm. from the main stem shows it initiating a further small branch of its own in the same plane as the primary one. The branching was apparently distichous at relatively long intervals, so that the habit was probably fairly close to the partial reconstruction shown in pl. 5. Adventitious roots, which are primarily limited to the axis of the branches, are similar to those of Lyginopteris. They are commonly hexarch or heptarch with conspicuous protoxylem groups at the points of the projecting arms (fig. 4). The phloem lies between these arms and, as іп Kaloxylon, there is a small amount of conjunctive parenchyma in the metaxylem. The stele is enclosed by a single poorly preserved dark ring which probably represents the endodermis. The cortex is homogeneous and identical to the inner cortical zone of the stem. The peripheral zone is undifferentiated, so that in transverse section the loosely packed cortical tissue with scattered secretory cells appears to extend clear to the edge of the ir- regularly lobed outer margin. This lack of a differentiated epidermal zone is one of the distinguishing features between these roots and those of Lyginopteris.! Secondary growth is rather rare but when present is again similar to that of Kaloxylon, cambial activity initiating in the depressions between the protoxylem lWhile the information in the literature is often vague on this point, similar roots have also been described for Heterangium, although they are usually listed as diarch or triarch (Scott, 1923). [Vor. 36 294 ANNALS OF THE MISSOURI BOTANICAL GARDEN points and producing wedges of typical secondary wood alternating with wedge- shaped rays opposite the protoxylem. The pitting is identical to that of the cor- responding tissues in the stem. Ав might be expected, the roots are of constant small size, with the most mature specimens showing 6—7 layers of secondary growth not exceeding 2 mm. in total diameter. The point of their insertion on the stem offers another similarity to Kaloxylon as roots are most frequently found im- mediately above the branch attachment (see Scott, 1923, p. 49) where their hori- zontally departing bundle seems to function, in part, in filling up the branch gap. As it has been possible to follow these roots from their insertion on the stem (fig. 6) out to typical transverse sections as in fig. 56, their identity is beyond doubt. Discussion: At first glance, in transverse view Microspermopteris presents a close resem- blance to Heterangium. ‘The size is comparable to H. minimum (Scott, 1917), and the primary wood with its thin network of parenchyma separating large groups of tracheids is more closely approximated by H. Kukuki (Hirmer, 1933) although the total amount of parenchyma is much less іп Microspermopteris. In the anatomy and insertion of the roots and the presence of multicellular emergences on the stem and branches we have structures comparable to Lyginopteris. However, the almost solid protostele contrasts with that of Lyginopteris, while the absence of leaf traces and of a vertical fibrous hypoderma, along with the exarch position of the proto- xylem, removes all possibility of assigning the specimens to Heferangium. While the horizon of our petrifactions is late Upper Carboniferous and H. Grievii Scott is of Lower Carboniferous age, there seems little doubt but that we are dealing with a more primitive plant which possibly represents the ancestral stock of Heterangium. In fact, if it were possible to increase slightly the amount of parenchyma in the primary wood, fuse (through webbing) the closely grouped parallel branches into a multi-bundle rachis (Lignier's (1908) theory of mega- phyllous development), and add a hypodermal zone of vertically elongated fiber bands we would have a stem quite similar to Heterangium. The sclerotic plates lying in the extreme outer cortical zone of Microspermopteris (fig. 10) are in radial view somewhat similar to the plates found in the middle cortex of all of the Heterangium species (with the exception of H. minimum which, according to Scott, lacks fibrous and sclerotic tissue altogether). The most obviously primitive and unusual feature, the leafless condition, can be assumed to be demonstrated since our material permitted examination of approxi- mately 37 cm. of different stems. The only similar character in plants of possible pteridosperm affinities is іп Eospermopteris (Апеиторруют) of the middle 1 Devonian." We must assume, then, partly on the lack of any intervening evidence, that we are dealing with a link leading the pteridosperm complex clear back to the 1 Although the fructifications originally described as seeds were later shown to be sporangia the anatomy of the stem still provides some basis for including the genus in the pteridosperms. 1949] BAXTER— FOSSIL PTERIDOSPERMS 295 Psilophytales. Indeed, Dawson’s restoration of Psilopbyton princeps (known only from compressions) is disturbingly similar to our own restoration of Microsperm- opteris, particularly in the flattened branching pattern and the presence of spine-like emergences on the stem. Since what we know of Psilophyton stelar structure in- dicates that it possessed only spiral and scalariform pitting there obviously cannot be any direct comparison in internal anatomy with that of the secondary wood and highly developed bordered pitting of Microspermopteris. However, the gross appearance is close enough to intimate that a compression form of M. aphyllum might be identified as а Psilopbyton. The middle Devonian genus Schizopodium offers a closer parallel in pitting and beginning of secondary growth. However, we consider it here just an example of the early, contemporaneous origin of these characters in contrast to the more conventionally primitive scalariform pits and solitary primary tissues, since the present evidence indicates its relationship to the Cladoxylaceae rather than the Pteridospermae. In Andrews’ (1940) account of the stelar anatomy of the pteridosperms he makes the i cu observation: on be stated that i view that the pteridosperms represent an и piu the ferns and cycads is no э od tenable. Rather we must loo common кш like ancesto mde term uis ne sporangia, a solid protostele жар primitive secondary wood for the origin of the ferns iid pteridosperms . While we are more inclined to feel that the ferns arose ER. in many ways our present genus fulfils the theoretical requirements outlined. The possibility that M. apbyllum was parasitic and its leafless condition а result of degeneracy rather than primitiveness has not been overlooked. The fact that the habit was probably epiphytic (climbing) or prostrate in a swamp humus tends to lend some weight to such a view as does the apparent absence of stomata and potential photosynthetic tissues. However, since no organs resembling haustoria are present and the roots resemble so closely the aerial ones of Lyginop- teris we cannot consider a parasitic condition too seriously. Ас the same time the combination of so many characters common to both Lyginopteris and Heterangium, along with the more primitive stelar structure and lack of differentiation into rachis and lamina, seems to point rather definitely to an ancestral form from which both genera may have developed independently. This conflicts with Scott's (1923) theory of Lyginopteris arising from a developmental series through the subgenus Lyginangium, and it may be necessary to reconsider some of Kubart’s transitional Heterangium species as being actually primitive Lyginopteris species leading back to Micros permo pteris. Since no seeds or any fertile parts whatever have been found, it is necessary to include Microspermopteris among the numerous other organ genera assigned to the seed-ferns on purely anatomical characters. It may be open to question whether in its leafless condition it could have borne seeds. If they were present they were [Vor. 36 296 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN more than likely terminal on the smallest branch divisions. However, there is no reason for not considering the possibility that the stem anatomy constituting the organ genera of pteridosperms did not precede the development of seeds (i.e. Eospermopteris (Aneurophyton). Certainly in many of the Devonian and Lower Carboniferous Calamopityaceae the presence of seeds is still to be proven, although the degree of differentiation into stem and leaf-bearing organs had obviously reached а more advanced stage than that shown by Microspermopteris. It can be hypothesized that certain small psilophytalian plants struggling for survival in an environment of increasing lushness and density might well have developed a vine-like habit, involving additional conductive stelar tissue as well as supporting sclerotic and fibrous zones, to be followed chronologically by differen- tiation into leaves (as a more efficient light-catching mechanism) and still later by seeds as a product of the changing and possibly drying environment. Viewing evolutionary change as a response to environmental change, it is difficult to see seed production ever replacing reproduction by spores if there had not been a constant impetus to survival under conditions unfavorable (lack of proper moisture) to spore germination. It also seems likely that the struggle for living space and light preceded the above conditions. From the original psilophytalian stock larger and anatomically more complex forms would arise independently, leading by gradual ecological adaptations to the recognized groups of Carboniferous plants. Certainly, the contemperaneous asso- ciation, even as early as the Upper Devonian period, of widely differing stelar structures for the ferns and pteridosperms would seem to indicate that they had a separate individual origin. Some of the earliest known forms assigned to the ferns such as Aracbnoxylon and Reimannia of the Upper-Middle Devonian show definite affinities to the Carboniferous Zygopteroideae, while Eospermopteris of about the same age shows many seed-fern characters im spite of apparently producing only spores. Thus, although the picture is complicated by the existence of numerous genera from compressions of which we have no knowledge of the internal anatomy (i.e. Archeopteris, etc.), the safest and most consistent distinction between the two groups would seem to be stelar structure. Using this as a criterion, it would be possible to have a phylogenetically correct classification including, on the one side, pteridosperms, having only terminal sporangia and, on the other side, ferns ex- hibiting heterospory. There certainly can be no reasonable objection to recognizing that the seed-ferns must have passed through a sporangiate stage or that ferns may not have achieved heterospory as a terminal development rather than as a stage in seed production. The designation of the group as Pteridospermae becomes increasingly unfor- tunate if we recognize their independent origin from the Psilophytales and the fact that the most primitive genera undoubtedly lacked seeds. The same situation has 1949] BAXTER—FOSSIL PTERIDOSPERMS 297 been dealt with іп the Lepidodendrales (i.e. Lepidocarbon vs. Lepidostrobus) with much less confusion. Here the general plant structure and anatomy have been accepted as the basis for the order, while seed production has been recognized as an advanced stage of heterospory within certain species and genera. For some reason it has never seemed necessary to create the Lycopsidospermae to include Lepidocarpon although considered on the same basis as the pteridosperms such should be the case. The following seem to the author to be the important points to bear in mind in relation to Pteridospermae in question. (1). They are psilophytalian in origin rather than arising from the ferns. (2). They are possibly of dual origin, with Eospermopteris leading to the Calamopityaceae complex and Microspermopteris leading to the Lyginopteris and Medullosa complex. (3). Leaves when present are large and fern-like; seeds when present are basically terminal. (4). Тһе stem and petiole anatomy is constant and characteristic enough to delimit clearly the group irrespective of known possession of seeds. (5). Included on the basis of the above anatomical criteria are genera such as Microspermopteris, which may or may not have had seeds, and Eospermopteris, in which the possession of seeds seems doubtful. (6). While genera known only from compressions such as Archeopteris may be links in the pteridosperm chain, they are just as likely, in the absence of anatom- ical evidence, to be true ferns. Heterospory, as shown by the modern genera Salvinia and Marsilia, as well as the fossil lycopods, does not necessarily always imply later seed development. (7). Seed development in the pteridosperms may in some isolated cases have preceded the differentiation of fern-like leaves with the seeds being borne terminally on naked branches. In other words, we are possibly dealing with early seed-plants which through parallel evolution developed fern-like leaves rather than, as orig- inally thought, with ferns which developed seeds. "Тһе distinction may seem trivial, but while the original premises are close the phylogenetic conclusions (depending on which starting point is selected) are quite distinct. The terminal position of the seed in Lyginopteris oldbamia and Aneimites fertilis are cases in point since these species are among the oldest forms in which seeds are known.’ (We can only hope that the discovery of fertile material of Microspermopteris will illuminate this point.) Diagnosis: Stems of constant small size, the stele never exceeding 2.5 mm. or the entire stem 5mm. Меѓахувет of large cells (150 р diameter), diminishing towards the lwhile these seeds are "terminal" on leafy fronds rather than on naked stems it seems not unlikely that their apical position involves a carry-over from a developmental stage prior to the acquisition of leaves. [Vor. 36 298 ANNALS OF THE MISSOURI BOTANICAL GARDEN periphery. Protoxylem exarch, of numerous indistinct groups. Xylem parenchyma arranged іп a thin network enclosing groups of 12 to 14 metaxylem cells. 5ес- ondary wood of small angular (in cross-section) tracheids from 3 to 16 layers in thickness, increasing in size towards the outer edge. Reticulate-bordered pitting with crossed orifices in the metaxylem and on radial walls of the secondary wood. Scattered bordered pits with lineal openings on tangential walls. Protoxylem spiral or annular. Rays rare or lacking; when present, small, uniseriate, 1—4 cells high. Cortex differentiated into inner and outer zones. The inner zone composed of isodiametric thin-walled cells, poorly preserved, with scattered secretory cells but lacking sclerotic tissue. Outer zone of thicker-walled, longer cells containing vertical series of horizontal sclerotic plates. Epidermis of similar but more elon- gated cells producing numerous multicellular pointed emergences. Leaf traces lacking. Branching distichous and at right angles to the stem. Roofs adventitious, hexarch to heptarch, commonly inserted above point of branch departure. Horizon: Des Moines Series of Pennsylvanian. Type: fig. 1. Slide 1602 in the paleobotanical collection of the Henry Shaw School of Botany of Washington University. SOME NEW AMERICAN MEDULLOSAS In the interval since Thiessen (1920a, 1920b) discovered and identified a pteridosperm stem as M. anglica (later to be assigned varietal distinction by Schopf, 1939) three new species and one variety of Medullosa from American coal balls have been brought to light. They are: M. Noei (Steidtmann, 1937), M. distelica (Schopf, 1939), М. Thompsonii (Andrews, 1945), and М. anglica var. ioensis (Andrews & Kernen, 1946). With the exception of M. Noe; all of these new species and varieties have formed a fairly closely integrated group assignable to the subgenus Anglorota along with the European species M. anglica, M. centrofilis, and M. pusilla. М. Noei, showing affinities with M. Leuckarti, is held to be transitional between the Carboniferous form cycle and the Permian species. Our knowledge of this very interesting group has been expanding rather rapidly, and with the hope of throwing still further light on the subject three new species of Medullosa are herewith described and additional new data are presented on M. Noei. Medullosa primaeva, sp. nov.: The following description is based on a single coal ball specimen from the Urbandale Mine located 1.2 miles west of Des Moines, Iowa, on U. S. Highway No. 6. This is the same location from which Andrews (1945) described M. Thompsonii. The horizon is the Des Moines Series of the Pennsylvanian. The material was collected and cut by Mr. Frederick O. Thompson of Des Moines, who generously donated it to the Henry Shaw School of Botany of Washington Uni- versity. It consists of a cut block 5.5 x 5.5 cm. and 1.3 cm. thick, with the stem 1949] BAXTER—FOSSIL PTERIDOSPERMS 299 exposed оп опе side through а fracturing away of the adjoining material. The tem's course is through the thickness of the block at a slight angle from the verti- cal, the total available length being 1.4 cm. Figures 14 and 15 show the аррсаг- ance of the top and bottom surfaces of the specimen prior to any grinding of the fractured edge. Approximately one half of the cortex and external tissues have been lost on the side of the fracture, though the multiple stele appears to be nearly intact. As shown in fig. 14, the upper! surface of M. primaeva exhibits 5 rather large and uniform-sized steles, 2 of them partially fused. There are also 2 small steles and a portion of a third on the fractured margin. The lower surface shows the same 8 steles, though somewhat altered in orientation and size, with 3 of the steles partially fused. It is not implied, however, that 8 was the constant number of steles as even in the short length of stem available stelar fusions and divisions were num- erous and the diversity of vascular growth even included the production of steles running horizontally through the stem for short distances (fig. 13). Still, even with these variations the stele number seems to have been constantly 6 or more. The stem measures 2.0 cm. in diameter, the individual steles ranging from around 5 mm. for the average large ones to .5 mm. for the very smallest. The 5 largest steles are closely comparable in size to those of M. T'bompsonii even though the stem of the latter is considerably larger, averaging 3.5 cm. in diameter. The steles are essentially radially symmetrical with a very slight tendency towards endocentricity. It may be argued that some of the smaller vascular strands (fig. 14, 53 and 58) do not constitute true steles. However, even the smallest possesses 8 or more layers of secondary wood and can be seen to be independent in its course through- out the entire length of the specimen (fig. 13). Neither can they be designated as accessory strands from which leaf traces might be produced (as in Su£cliffia and M. anglica) since the two largest of the 8 steles are, in our specimens, the sole observed source of trace departure. The primary wood is small in amount and is characterized by the almost com- plete absence of parenchyma; what little there is being restricted to a thin network running through or enclosing the primary cylinder, from which the high uniseriate rays extend out through the secondary wood. The primary wood forms a cylin- drical to irregularly shaped core around which it is often difficult to determine the exact inner limits of secondary growth. The protoxylem is not distinguishable (figs. 14 and 19). The secondary wood is variable as to cell size and radial growth, the first- formed layers being often composed of large tracheids (similar in size to the metaxylem) which merge some distance out into rows of much smaller cells. On The interpretation as to lower and upper surfaces of the stem is made оп the basis of the outward passage of leaf traces adli in a series of longitudinal peels. [Vor. 36 300 ANNALS OF THE MISSOURI BOTANICAL GARDEN the other hand, the same stele may, on the opposite side, show the first-formed secondary wood to consist of very small (in diameter) tracheids and the successive outermost cells to be large (fig. 19). Each stele is enclosed by a dark tissue of crushed cells, probably of cambium and phloem, in which are occasional small resin canals. The entire stelar assembly is surrounded by a very thin, parenchymatous "periderm" which is never over 2 to 3 cells thick and is often so poorly preserved as to be indistinguishable from the cortex. The rays are up to 1 cm. or more in height and apparently constantly uniseriate. In this regard they probably come closest to Andrews’ (1940) classification H B in which he originally placed all of the Anglorota group as well as M. Noei and M. Leuckarti. The cortex is composed of cells rounded in transverse view but elongated vertically two to three times their width. Traversing upward and outward through the cortex are numerous small collateral leaf traces consisting solely of 3—4 proto- xylem tracheids with spiral thickening (fig. 27) and an abaxial phloem group. There is definitely no secondary wood associated with the trace. Mucilage canals with their "resin rodlets" are scattered sparingly through the cortex and are un- usually small, having a constant diameter of less than 60 и. They are commonly closely associated with the leaf traces and, in contrast to the majority of the other American species, seem to have seldom invaded the hypodermal, sclerotic strand zone. The sclerotic strands form a single layer and in transverse view appear circular to slightly radially elongated. They are separated by 2-3 layers of cortical-like cells from а conspicuously darkened epidermis. This latter tissue consists of brick- shaped cells with a slight tangental elongation and rather thick walls which oc- casionally show evidences of a cuticle. Discussion: Our specimen shows a rather close similarity to Medullosa T bom bsonii in the secretory canals—their small size, their comparative scarcity in the stem as a whole, and their almost complete absence in the sclerotic fibrous zone. However, although Andrews (1945) points out that the number of steles in the Medullosae is generally an unreliable taxonomic character (stelar divisions and fusions often resulting in variations of 2—3 or more steles), the possession by Medullosa primaeva of 7 to 8 steles, which tend to pursue horizontal courses through the stem and almost com- pletely lack conjunctive primary parenchyma, clearly makes it of specific im- portance and distinct from M. Thom psonii. Medullosa centrofilis and M. pusilla are the only other species which seem similar enough to deserve consideration. The former comes closest to M. primaeva in number of steles (considering the "star ring" as a fourth stele) but lacks the hori- zontal stelar meanderings and exhibits many more and larger secretory canals. М. 1949] BAXTER—FOSSIL PTERIDOSPERMS 301 pusilla is comparable іп over-all stem size but again is separated by the stele num- ber, the difference between 3 and 7-8 being held as more than specific variation. The slight tendency toward endocentricity of the secondary wood and paucity of conjunctive parenchyma in the primary area, along with its generally netted arrangement, make the individual steles more closely comparable to those of Microspermopteris than to Heterangium. According to Bower's (1930) "size and form" principles one would expect relatively little parenchyma in steles of such small size, but comparison with the equally small stele of M. Thompsonii seems to show greater differences than can be accounted for on the basis of Bower's hypoth- esis. Thus although the M. primaeva stelar number is much larger than in any of the other species of equivalent age the general stele structure appears to be more primitive. This is in contrast to what might be considered advanced development as evidenced by the absence of secondary wood in the leaf traces and leads one to believe that the presence of such secondary wood іп M. anglica is not necessarily indicative of a primitiveness either for the character or the species as a whole. On the contrary, it seems most logical at present to consider M. primaeva with its numerous small, radially symmetrical steles as having developed from a monostelic stem through stelar proliferation and accordingly representing a primitive stage of the genus. The specific name primaeva has therefore been assigned. Diagnosis: Stem small, not exceeding 3 cm. Steles numerous, 6-8, frequently fusing and dividing, ranging from .5 to 5 mm. in diameter, often pursuing a sinuous hori- zontal course through the stem. Primary wood parenchyma small in amount or lacking. Resin canals small and few, almost entirely absent from fibrous hypo- dermal zone. Rays uniseriate; secondary wood of variable cell size; irregular radial growth, only slightly endocentric. Horizon: Des Moines Series, Pennsylvanian. Type: Fig. 14. Slide 1615 in the paleobotanical collections of the Henry Shaw School of Botany of Washington University. Medullosa ејопваќа, sp. nov. The following description is also based on coal-ball material collected by Mr. F. О. Thompson and his associates of Des Moines, Iowa. The source was a large open pit mine of the Atlas Coal Co. located in Wilson County, Iowa. Тһе horizon is Pennsylvanian, Des Moines Series. The greater part of material from this locality was too heavily pyritized to permit good plant preservation, but the stem fragments which were identifiable indicate a flora consisting primarily of Lepidodendron and Calamites. Pteridosperms were relatively rare, the present description being based on a single stem specimen 7 cm. long and 6 x 1.5 cm. in diameter. There evidently had been some crushing, which would account partially, but not altogether, for the asymmetry (fig. 16). [Vor. 36 302 ANNALS OF THE MISSOURI BOTANICAL GARDEN The outstanding feature of the three steles is their extreme endocentricity in which character they resemble Schopf's (1939) M. distelica. The steles maintain their individuality, i.e., no stelar fusion, and relative position throughout the 7 cm. of stem length with the endocentric secondary development also showing a con- stant approximate 4 to 1 ratio in thickness as compared to the exocentric secondary tissues. The three steles are of equal size, averaging 10 X 2 mm., the plane of elongation following that of the stem as a whole and constituting a factor in the evidence that the general asymmetry was a natural character of the living plant. Two of the steles occupy a position directly opposite one another and near the center of the stem while the third is isolated off to one side (fig. 17). The primary xylem consists of a long narrow area following the elongation of the stele. It is seldom over 2 cells in width and runs up to 8 mm. in length. Тһе metaxylem tracheids are large (.2 mm.), and usually triangular in cross-section. There is relatively little conjunctive parenchyma, while the protoxylem is indiscernible. The secondary wood is rather unusual for a Medullosa in that the majority of the cells are square in transverse view, while the others are of the more character- istic alternately arranged pentagonal shape. Also, in contrast to the somewhat similar M. distelica, the radial rows of secondary tracheids are not conspicuously divided by rays nor are the outermost cells differentiated. In addition, the rays are uniseriate, not particularly numerous, and almost impossible to detect in transverse view. Surrounding each stele is a thin dark layer of crushed cells containing numerous small resin canals. The periderm is thin (3—4 cells), poorly preserved, and ap- parently interrupted at the narrow ends of the steles where the leaf traces are given off. The cortex is rather poorly preserved due to pyritization, with numerous rather large slime canals (.2 mm. in diameter) scattered throughout it and also conspicuously associated with the fiber strands in the sclerotic-fiber hypodermal zone. The fiber strands are arranged in 2-3 alternating rows, the individual strands being somewhat radially elongated. А rather striking character of the species is the large “compound” resin canals running horizontally through the stem. They measure 1.3 mm. in diameter and are approximately six times the size of the vertical canals. Figure 18 illustrates several of these canals at the same magni- fication as the vertical canals in fig. 17. They are lined with numerous small triangular and diamond-shaped secretory cells which appear to contain the same black resin substance as the canal itself. The leaf traces are given off from the narrow ends of the flattened steles and are initially quite large, consisting of 10—12 or more primary tracheids which divide farther out in the leaf bases into numerous smaller bundles. Three leaf bases are shown in fig. 16, two detaching themselves from the narrow transverse extremities of the stem and approximately parallel to it, while the third is being given off from the stem's broad side and at right angles to it. Thus it would ap- pear that in M. elongata we are dealing with a creeping stem of possibly dorsi- 1949] BAXTER—FOSSIL PTERIDOSPERMS 305 ventral symmetry. As stated earlier, the dimensions of 6 X 1.5 cm. for the stem can be only slightly attributable to crushing while the equivalent asymmetry of the steles and general lack of any crushed tissue indicate the form to be more or less characteristic of the living plant. In addition, the 7 cm. of stem length of our specimen exhibits rather short internodes, with several leaf bases given off in the same bilateral plane from the narrow opposite stem sides. The position of the stem on the edge of the containing coal ball prevented observation of more than the one dorsal leaf base shown in fig. 16. However, it seems plausible that these erect leaves were produced between internodes as short as the lateral ones. As implied previously, M. elongata is most closely comparable to M. distelica on the basis of the extreme endocentric secondary growth. However, in the possession of three steles, which at no time show any tendency towards fusion, the difference in primary wood, fewer and uniseriate rays, and square secondary tracheids, as well as the large compound resin canals and apparent dorsi-ventral habit, M. elongata is clearly distinct and of specific importance. | The species name elongata is assigned in recognition of the asymmetric trans- verse elongation of the steles (10 XX 2 mm.), as well as the equivalent transverse elongation of the stem as a whole. Diagnosis: Stem asymmetric to bilaterally symmetric, approximately 6 Х 1.5 cm. in diam- eter, only slightly crushed in the smaller dimensions. Steles 3, extremely endo- centric, 2 opposite one another in stem center, the third isolated to one side, retain- ing their relative positions and not fusing or dividing in over 7 cm. of stem length. Vertical resin canals numerous, averaging .2 cm. in diameter; fewer horizontal, large compound resin canals averaging 1.3 cm. in diameter. Habit creeping or climbing with leaves being given off from sides and dorsal surface of stem. Locality: Atlas strip mine, Wilson County, Iowa. Horizon: Pennsylvanian, Des Moines Series. Type Specimen: Fig. 16. Slide 1617, in the paleobotanical collections of the Henry Shaw School of Botany of Washington University. Medullosa endocentrica, sp. nov.: This third new medullosan stem comes from coal-ball material found in a stream bed outcropping near Berryville, Illinois. "Тһе location is near to that from which Steidtmann (1937) described M. Noei, and the horizon is the same, being determined as the upper part of the McLeansboro group of the Pennsylvanian of Illinois. The coal ball containing the specimen was 15 cm. long by 8 cm. wide. The stem followed a straight course through almost the complete longer axis, being itself 12 cm. long and 1.2 X .7 cm. in diameter. These dimensions are of the stelar system only, the cortex and other external tissues having been lost. How- ever, as can be seen by referring to fig. 20, the unique appearance of the steles seems [Vor. 36 304 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN to justify recognizing the specimen as a new species and possible culmination of endocentric development, in spite of the lack of information as to the stem's external anatomy. The stem consists of 3 steles, 2 of which are 5 X 5 mm. in diameter, the third, 2 X 1.5 mm. in diameter, situated on the side equidistant between them. Sec- ondary growth is almost entirely endocentric, only one of the larger steles showing a very small amount of exocentric growth. These characters are constant for the entire 12 cm. of stem length as are also the relative stelar positions. Thus there can be little doubt that this extreme centripetal growth was a distinct feature of the living plant, and the evidence also suggests that the stelar system had achieved a degree of uniformity not found in any of the other Anglorota species. The primary area of each stele is oval in transverse view, and measures 1.5 x1 mm. in the 2 larger steles. It consists of scattered groups of large metaxylem cells intermixed with considerable conjunctive parenchyma. The protoxylem appears to consist of two small exarch groups of tracheids with annular thickening while the metaxylem shows the characteristic dense reticulate bordered pitting. The secondary growth is produced in a fan-like pattern, radiating towards the stem center for an average of 15 cells in the large steles and 6 cells in the small stele. The transverse dimension of the cells increases gradually from the innermost to the outer rows but on the average is quite large, around .2 mm. The rays are of great height, more than 1.7 cm., and in the wood are uniseriate or at the most 2 cells wide. They commonly separate radial rows of 2—3 tracheids (fig. 22). The pitting of the secondary wood is bordered and densely reticulate on the radial and oblique-tangential walls, with relatively few scattered pits on the directly tangential surfaces. The length of the individual tracheids is also quite extreme, an almost perfect tangential section showing them to be over 1.7 cm. Adjoining the outermost secondary wood (and continuous with it in general pattern and ray position) is a rather disorganized tissue which we were at first inclined to interpret as undifferentiated secondary wood. It consists of large thin-walled cells, similar in size and shape to the secondary tracheids but completely unpitted. On entering this zone the thin rays of the wood become greatly expanded, averaging from 4 to 8 or more cells in width, and appearing to be completely mature in their develop- ment (fig. 21). This point, along with the total absence of any rudimentary pitting, in either the tangential or radial walls of the vertical elements, in our opinion, makes it very unlikely that the cells are immature secondary tracheids. On the other hand, one or two of our tangential peels through this zone shows vestiges of what appear to be disarranged transverse walls with extremely small pores similar to the sieve plates of living plants (fig. 23). Scott, in his description of M. anglica, has pointed out that the rays become much wider in the phloem zone, while excep- tionally large sieve tubes are common to plants of vine-like habit. Therefore, on the basis of the above evidence (which may be not entirely conclusive), along with the position of the tissue, we are inclined to consider it tentatively as phloem- like in nature. 1949] BAXTER—FOSSIL PTERIDOSPERMS 305 Lying between this phloem region of the three steles is а thin parenchymatous zone of vertically elongated cells in which are scattered numerous resin canals con- taining their opaque resin rodlets. The entire stelar assembly is completely sur- rounded by a thin layer of dark, crushed cells of possible periderm nature. The specific name endocentrica is assigned on the basis of the complete and constant centripetal secondary growth. Diagnosis: Steles small, not exceeding 5 XX 5 mm., 2 opposite one another and of equal size, the third smaller, situated on the side between them. Secondary growth completely centripetal. Primary xylem exarch. Metaxylem with considerable conjunctive parenchyma. Rays 1—2 cells wide, expanding in the conspicuous phloem zone to 4—8 cells in width, exceeding 1.5 cm. in height. Horizon: Upper part of the McLeansboro group, Pennsylvanian of Illinois. Type: Fig. 20. Slide 1619 in the paleobotanical collections of the Henry Shaw School of Botany of Washington University. Discussion: As stated previously, Medullosa endocentrica appears to represent the ultimate in centripetal secondary development, and accordingly its closest affinities lie in the direction of M. distelica and M. elongata. It differs from the former primarily in possessing a third distinct small stele, constant for the whole stem and in the more perfect symmetry of the opposing "twin" steles. It differs from M. elongata in stele shape (M. elongata averaging 10 XX 2 mm.), as well as in the character of the primary wood and amount of conjunctive рагепсһута. M. endocentrica differs from all the other Medullosas in its constant degree of endocentricity and the possession of a unique phloem zone of exceptionally large sieve tubes. It is possibly significant that here, as in M. elongata, we appear to be dealing with a stem of more bilateral (dorsi-ventral) than radial symmetry. From the form of stele development in M. endocentrica (fig. 20) leaf traces could only have been given off in the three planes opposite the abaxial sides of the primary xylem groups. Thus while the orientation of the steles is somewhat different in the two species it is likely that the phyllotaxy was similar, the leaves being produced laterally and vertically (dorsally) with the side lacking a leaf trace source being considered ventral! The predominance of 3 stele forms in the Medullosae is perhaps indicative of this trend, which, however, reaches its inescapable climax only in the extremely endocentric species. M. endocentrica has apparently almost reached the limit of its potential stem enlargement through secondary growth but it is still much too small to permit interpretation as an arborescent plant. Therefore it could not possibly uld be pointed out here that since the traces show no ке of "girdling," the direction sho of dur initial departure is a valid indication of the stem's phyllotax [Vor. 36 306 ANNALS OF THE MISSOURI BOTANICAL GARDEN have attained a habit comparable to that suggested in the reconstruction of M. Thompsonii (Andrews, 1945). Also if it produced relatively large Alethopteris- type fronds, as appears likely from the numerous associated leaves, it must have been a prostrate or climbing vine since the small stem would be incapable of sup- porting any such great weight. Thus the line of endocentric development from M. distelica and M. elongata to M. endocentrica appears to be one leading to constantly diminishing size accompanied by a change from radial symmetry of the stem with complete spiral phyllotaxy to a bilateral, dorsiventral symmetry with lateral and dorsalleaves. Extreme asymmetry can accordingly be seen to be an advanced rather than primitive character, which as it reached its climax resulted in a line of "dead- end" development in which the stem size was small and fixed. In this viewpoint we agree with Schopf (1939, page 204) where he states: It seems more tenable to consider the distelar condition in the new га Medullosa as reduced from a more polystelar type = than as directly derived from an ancestry reminiscent of Sutcliffia. It is also кра" пе that the extreme endocentric ақа ыы found in distelica is not a primitive featur We have included in the above quotation Dr. Schopf's statement on a distelar condition probably resulting from the fusion of a polystelar type in that it also agrees with our conception of what seems to be another and more important line of development in the Medullosas. Ав pointed out in the discussion of M. primaeva, a polystelar condition with numerous small radially symmetrical steles would seem to have arisen most logically by stelar proliferation of a monostelic stem with a similar small symmetrical stele. Then through subsequent tendencies to stelar fusion the line would lead through the small tetra- and tri-stelar forms to the large asymmetric stelar forms such as M. anglica and M. Noei. The fact that the smallest species (M. primaeva and M. pusilla) have the most radially symmetrical steles while the larger species (M. Thompsonii, M. elongata, M. distelica and M. anglica) all have greatly elongated (transversely) asymmetric steles is evidence, in our opinion, that the latter have been derived from polystelic fusion. (А good example of this point is illustrated in Andrews’ (1945) diagrams of stelar fusion in M. Thompsonii where it can be seen that the fusion produces а larger, more asymmetric and more endocentric stele). Where the fusion standard- ized itself at 3 steles of approximately equal size, or with one slightly smaller, a balance appears to have been reached which was only upset by the origin of the erratic cambial growths of M. Noei. Regarding the phyllotaxy of the predominately 3-steled forms it may also be pointed out that while the leaf arrangement has been mainly interpreted as spiral, Scott, in working with a specimen of M. anglica one foot in length, showed that the leaves were still produced on only three sides of the stem ( text-fig. 1). While Scott himself did not enlarge on this evidence, it seems to imply (when correlated with forms such as M. elongata and M. endocentrica) that the significance of the normal number of 3 steles in the Carboniferous Medullosas lies in the essentially 1949] BAXTER—FOSSIL PTERIDOSPERMS 307 dorsi-ventral habit of the plant as a whole. As may be noted in text-fig. 1 of M. anglica (which is typical for most of the 3-stele species), the steles are arranged in the shape of a flattened triangle, the obtuse apex being always towards the flattened side of the stem from which the dorsal leaf bases are produced, while the steles at the opposite acute angles are oriented towards the position of the lateral leaves. o АЦ о ) гусли бы, == ч. о 2522 y н stone жеуі F e е с Ja < Text-fig. 1. Medullosa Hi ee о.с., outer cortex; m leaf trace; 57, stele; pd, y ur es m sclerotic strands separating petiole from stem; r, t; p 5 stran and BC represe т che Pod д nt departure with the phyllotaxy being 2/5. Note fes vits arrangemen eaves оте side tbe stem completely devoid of leaves; so tbat tbe ent s babit may be BO to have been PSU ventral (after Scott). This bilateral symmetry is borne out also in the over-all stem shape of nearly all of the 3- to 4-steled group. М. centrofilis had a stem diameter of 5 X 1.5 cm.; M. pusilla, 2.2 >< 1.3 cm. (Scott points out that the larger diameter here had lost considerable tissue and was probably still greater in life); M. Thompsonii, 7 X 2.5 cm.; M. anglica, 10.5 XX 3.7 ст.) M. elongata, 6 X 1.5 ст. M. endocentrica also undoubtedly had a flattened stem but the lack of preservation of the outer tissues in our specimen precludes such measurements. АП of the above figures include the attached leaf bases, but since it has been shown that these were always decurrent for long distances on the stem and more or less overlapped, the measurements give a truer habit proportion than those of the stem alone would. The asymmetry can be seen to be quite constant in a proportion of 3:1, which fact alone makes it ex- tremely doubtful that it is altogether due to crushing. De Fraine states for M. centrofilis that “the shape of the stem with its covering of leaf bases was thus distinctly flattened.” It would seem that the dorsi-ventral habit, while most obvious іп species such as M. elongata and M. endocentrica was also the rule in the majority of the other species. M. Thompsonii is a possible exception since rather careful study of the type specimen seems to indicate that in this particular stem the orientation of the stelar assembly tends to vary, so that the planes of leaf trace departure are dis- [Vor. 36 308 ANNALS OF THE MISSOURI BOTANICAL GARDEN tributed more generally in the four opposing radii and thus supply petioles in a true radially symmetrical spiral phyllotaxy. However, such variations are not too unusual. Selaginella, for example, in the two present-day species, S. rupestris and S. apoda, shows a variation from erect plants with radial symmetry to creeping plants with bilateral symmetry. This view, accordingly, makes it necessary to visualize the Medullosae as primarily an assemblage of creeping or climbing plants rather than as comparable to present-day tree ferns. ‘The prostrate habit has previously been postulated by several earlier workers on the basis of the small stem of the Anglorota plants, while the great stem length recorded by Weber and Sterzel (1896) for some of the Permian specimens is almost conclusive proof of the vine-like habit of the much younger species. Weber and Sterzel state: сараланды wie М 43, das bei 92 ст. Linge nur еїпеп Durchmesser von durch- Жы: 8,8:4.4 cm. besitzt, kónnte man wohl an die von Góppert & Stenzel vermutete E i óppert & Stenze Schlingpflanzen denken kónnte. Е re Pin des Wachstums liegt ми о das “e нож háufige Vorkommen von Bruchstücken dieser Art begründet. In addition to the evidence offered by the dorsi-ventral flattened stem, the phyllotaxy, and the great stem length in relation to its width, there is also consider- able anatomical proof that the Medullosae were primarily prostrate or climbing vines. Solereder (1908) has shown that unequal xylem growth and flattened stems are characteristic of many living lianas, while Westermaier and Ambronn (1881) have pointed out that tracheids and sieve tubes of more than average diameter are typical of living climbing plants. As noted by Andrews (1940), the tracheids of M. Noei and M. anglica may reach a diameter of 250 м (larger than most vessels) while M. endocentrica has extremely broad sieve tubes. Although the acquisition of concentric rings of exocentric secondary wood permitted an increase in size of the Permian Medullosas, along with a possible return to a radially symmetrical spiral phyllotaxy, the vine-like habit of their Carbo- niferous predecessors appears to have been largely retained. The absence of a vine-like habit in the living or fossil cycads need not present any insurmountable difficulties in retaining the theories of their origin from the Medullosas, since, with the increased girth afforded by centrifugal secondary growth, the developmental trend may be assumed to have been away from climbing towards arborescent forms. This change may also have been influenced and accompanied by a disappearance from the scene of the potentially supporting Psaromius, Cala- mites and Lepidodendron through a gradual shift to a xerophytic environment. To summarize and restate our evidence: 1. A large number of small symmetrical steles with almost solid primary wood represents the primitive condition. Of the different specimens which have so far 1949] BAXTER— FOSSIL PTERIDOSPERMS M. stellata M.Levcharći Celi flr Su Lf £i zz M. Mo е; M. an /1са 8 №. distelica _ > /1. elongata Ис IM. endocentríca У. Thompsonit М. centrofilis 27. primaeva PsiLorHYTALES ( Mrcrospermopteris 2) Text-fig. 2. Proposed Medullosa phyletic chart. Explanation in text. been described, M. primaeva seems to most nearly represent this stage. In this light it seems probable that the origin was not from Heterangium but rather from a more ancient form possibly akin to Microspermopteris. Arnold (1940) has de- scribed a small polystelic stem from the Middle Devonian of New York to which he has given the name Xenocladia medullosina, It consists of 9-10 small (not much over 1 mm.) steles with a small solid protostele. He states that: “Xeno- cladia may eventually prove to be a representative of some intermediate stage be- tween the Psilophytales and certain of the polystelic Pteridosperms.” While we 309 [Vor. 36 310 ANNALS OF THE MISSOURI BOTANICAL GARDEN аге not inclined to attach too much weight to any direct connection between Xeno- cladia and a stem such as M. primaeva, it does show the possible origin of a poly- stelic structure contemporaneous with Rhynia and Hornea and, along with Micro- spermo pteris, indicates a psilophytalian origin for the pteridosperms. 2. Through subsequent tendencies to fusion the stele number eventually Бе- came standardized at 3, with occasional variations of 2 or 4 through further fusion or division. 3. Accompanying the above transition there was an increase in the size of the plants and consequently of the leaves, which (since in a closely grouped poly- stele the leaf traces of necessity are produced from the abaxial sides) tended to inhibit exocentric (centrifugal) secondary growth and thus resulted in an endo- centric (centripetal) asymmetry of the steles. 4. As this endocentricity became pronounced the production of leaf traces also became more localized on the abaxial stelar sides, resulting in a phyllotaxy of lateral and dorsal leaves (text-fig. 1, M. anglica). M. elongata and M. endo- centrica represent a climax of this trend which in the latter undoubtedly formed an end development. 5. Where the endocentricity did not reach such extremes, the plants tended to increase in size through the production of accessory vascular strands (M. anglica), while eventually tangential secondary growth (to compensate for the limitations of endocentric growth), along with the origin of prolific erratic cambial zones (M. Noe?), led to the larger and more complex Permian species. Text-fig. 2 shows a provisional and tentative phyletic chart based on the above points. Al- though the horizon of all of the American species is considerably higher than that of the European ones, the evidence of the stems themselves still seems to justify the reverse grouping. For the reasons given above M. primaeva constitutes the base with the main line of development leading through M. Noei and M. Leuckarti to the Permian forms, while the three extremely endocentric species constitute a dead-end side chain of development. Medullosa anglica is placed near the top because of its relatively large size and erratic "periderm" resembling that of М. Noei. The large leaf traces with sec- ondary growth may be viewed as advanced and of the nature of specialized ac- cessory vascular strands to supply the increased needs of the enlarged petioles. Also Sutcliffia, differing from M. anglica primarily only in its monostelic structure, may be regarded as resulting from a fusion of the three steles. Medullosa Noei Steidtmann: Steidtmann (1937, 1944) established М. Noei as the first clearly recognizable American species of the genus and gave us an excellent description of the salient features and peculiarities of the stem as well as the associated leaves, roots, and seeds. However, the holotype on which he based most of his description had only 1949] BAXTER—FOSSIL PTERIDOSPERMS 311 one stele with portions of two others, while our present specimen consists of a nearly entire stem with three large complete steles. It was collected from the same outcrop near Berryville, Illinois, that produced M. endocentrica and is exceptionally well preserved in its cellular details. The coal ball, of which the stem constituted almost the entire bulk, was 28.6 cm. long and approximately 16 XX 7 cm. wide in the middle portion, tapering to blunt points at either end. The essential features offering possibilities of an enlarged interpretation and understanding of the species are: 1. The presence of a distinctive phloem zone characterized Бу broadened ex- tensions of the medullary rays. 2. “Periderm rings" which show a transitional development of secondary wood, a stelar origin, and a remarkable similarity to the Permian Medullosae "star- rings." 3. Adventious roots, which run for long distances through the cortex parallel to the steles, range from triarch to pentarch, and show a well-preserved aerenchy- matous cortex. Before discussing these points in detail a brief description of the stem as a whole will be given. As shown in text-fig. 3, there are three large steles, tan- gentially elongated, which occupy the major portion of the area as seen in trans- verse section. Throughout the 28.6 cm. of stem length they remain essentially independent although steles а and c occasionally partially fuse and then re-separate. Their orientation with relation to one another is similar to that of the steles in M. elongata, two of the steles being side by side while the third occupies a position off to one side. They average 6 Х 1.5 cm. in diameter and show no particular endo- centricity, the secondary growth being the greatest in a tangential plane rather than towards the stem center. This is a greater elongation than was usual in Steidtmann's specimen, although he records that the stele did at times reach a measurement of 6 X 1.8 cm., which is certainly close to the above. For the rest, the primary and secondary wood are essentially equivalent to Steidtmann's speci- men and exhibit primary tracheal bundles, prolific and erratic "periderm" growths, and tetrarch roots originating in and running through the primary area. The cortex is crowded with “регійегт” rings and tangentially elongated bands, as well as with the numerous large adventitious roots with their many small branches. The hypodermal sclerotic zone is present as the outermost limiting tissue, the leaf bases being absent. While there are apparent general differences between the present specimen and Steidtmann's diagnosis of М. Мое? (mainly in the lack of any pronounced endocentric growth, and absence of a definite broad periderm zone), it is felt that the variations fall within those allowable to a species, partic- ularly in the Medullosas. The following descriptions of the previously outlined characters are therefore intended solely as an addition to the M. Noei diagnosis and not as of new specific or varietal importance. [Vor. 36 312 ANNALS OF THE MISSOURI BOTANICAL GARDEN OLA BAA m оз CL ” d, ж. Y ew н 6, A A P £M BAXTER—FOSSIL PTERIDOSPERMS [Vor. 36, 1949] 326 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 4 Micros permopteris aphyllum Fig. 8. Radial section of stem: x^, sec ondary xylem; un inner cortex; OC, outer cortex; “ер, sclerotic plate; ет, emergence. From slide 1609, Fig. 9. Tangential view of secondary wood showing small uniseriate rays and pitting in tangential walls. From slide 1610, X 135 10. Radial section of stem showing sclerotic plates in outer cortex just within epidermis. From slide 1611 53. Tangential un through outer cortical zone showing anastamosing hori- zontal plates. From slide 1612, X 25. Fig. 12. Reticulate bordered pitting in wall of a metaxylem tracheid. From slide 1613, X 450. PLATE 4 36, 1949 Mo. Bor. Слкр., Vor. ANN. „Мэ e © wis f bsc и “ EI = 4 е : : E > š р: 1 427 4 “ - Ae mex LE 3 мовы” - = : 4 . ЖР. заа 4# РЯ, 40-7 E = iy WM - xil ute s ЗСК < at BAXTER—FOSSIL PTERIDOSPERMS [Vor. 36, 1949] 328 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 5 A partial reconstruction of Microspermopteris abbyllum, Х 4. Explanation in text. PLATE 5 ANN. Mo. Bor. GaAnp., Vor. 36, 1949 IT a 5% ота бр олым» Жа ap м. ха? КЖ же ы. вон ord BAXTER—FOSSIL PTERIDOSPERMS [Vor. 36, 1949] 330 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 6 Medullosa primaeva 13. Transverse-longitudinal view of type specimen showing branching and fusing of the ле Рея а, small stele іп а horizontal course through the stem; Sr, 52, $3, and S4 show portions of steles indicated in next figure. From slide 1614, X 4.5. А ansverse section of top surface of type онн Sr, S2, 53, S4, 55, 50, 57, and % Же ate the eight steles. From slide 1615, 5. Transverse section of bottom surface of type specimen. Steles have partially m dn slide 1616 PLATE 6 Mo. Вот. Сакр., Vor. 36, 1949 ANN. ERMS BAXTER—FOSSIL PTERIDOSP [Vor. 36, 19491 332 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 7 Medullosa elongata Transverse view of сотор еке суре specimen: а апа Ё, lateral leaf bases; с, dorsal leaf bais: From slide 1617, X 2 Fig. 17. Enlargement of central portion of stem shown in fig. 16 to show the details of the РА steles. Note narrow line of primary wood in lower left-hand stele. From slide 1617, X 6.5. Fig. 18. Compound resin canals which run horizontally through the зет. From slide 1618, X 6.5. Medullosa primaeva Enlargement of steles 57 and S2 from fig. 14. Note almost solid protosteles. From "lide 1615, X 16. 7 PLATE GARD., VoL. 36, 1949 Bor. ANN. Mo. m „е є, CNET gm. 4 BAXTER—FOSSIL PTERIDOSPERMS 334 [Vor. 36, 1949] ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 8 Medullosa endocentrica g. Transverse section of stelar assembly of type specimen with enclosing "peri- derm": a, third small stele; b, phloem zone; Xr, primary xylem. From slide 16 10 Fig. 21. Transverse- tangential section showing very bro ad rays in “phloem” area. Above point indicated by (а) is transverse, below tangential. From slide 1620, X 40 Fig. Tangential-transverse section through secondary xylem showing narrow (1—2 cells wide). rays; а indicates line between transverse and tangential views. From slide 1621, x 40. 23. Portion of a transverse “sieve plate" from one of the large “sieve tubes” of the phloem zone. From slide 1620, Х 65 8 PLATE GARD., Vor. 36, 1949 Mo. Bor. ANN. Ben Б N sé E бы ња „^^, fe: 2 NX Exi E 5522 У v , f gy 7 eo [ £4 eR CM равој ad n >> се М б" 5 НАВЕ. 4 жж ма, E т 2 к. de v9 40. ұсақ ж s. = во“! Sie aces: 2724 : > ГЕ; ava Ж бы К . a ы PTERIDOSPERMS BAXTER—FOSSIL [Vor. 36, 1949] 336 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 9 Medullosa Noei Fig. 2 single “cambial ring” showing development of secondary wood. е сеп- tral area is енын of primary tracheids and parenchyma. From slide 1622, . A single well-preserved 2. ар нк root. Note enclosing periderm апа aerenchymatous соггех. From slide 162 Fig. 26. Marginal portion of a stele showing fringe of expanded phloem rays. From slide 1624, X 12 Medullosa primaeva Fig. 27. vs ria section through cortex showing spiral thickening in a leaf trace From slide 1625 PLATE 9 1949 36, ANN. Мо. Bor. Сакр., VOL. BAXTER—FOSSIL PTERIDOSPERMS [Vor. 36, 1949] 338 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION Or PLATE PLATE 10 “Star Rings” 28. А “cambial” or “periderm” ring 2. M. Noei in which radial rows of secondary xylem have formed. From slide 1622, X 5.5. Fig. 29. A “star ring” from М. stellata у. Cotta a. typica, at the same magnification as the above (after Weber and Sterzel), X 5.5. Fig. 30. An “aggregate leaf trace" from M. Thompsonii (see fig. 59b). From slide 1626, X 13.7. Fig. 31. А single small stele of M. primaeva. From slide 1615, X 20.7. Fig. 32. “Star Ring" of M. centrofilis (after De Fraine), X 20.7. ANN. Мо. Вот. GARD., Vor. 36, 1949 PLATE 10 BAXTER—FOSSIL PTERIDOSPER MS 340 [ Vor. ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 11 Diagrams of the stelar assemblies with а periderm (where present) of all of the Carboniferous species of Medullosa with t pee of M. Leuc is 2.8, and all of t s. 34, 36, 42, and 43 are taken fro (1945, 1946). The primary area of the stele is ы. as solid e diagrams other than figs WS i ‘ black, radiating lines, and the periderm as enclosing double or dotted вы depending оп whether it was actually observed or was partially assun karti. The the secondary wood as Fig. 33. M. centrofilis De Fraine; Fig. 34. M. DE Baxter; Fig. 35. M. 2” они Andrews; Fig. 36. M. endocentrica Baxter: Fig. 3 anglica Scott; Fig. 38. distelica Schopf; Fig. 39. M. pusilla Scott; Fig. 40. M. jt var. Thiesseni Schopf. (Continued on pl. 12) 36, 1949] ANN. Мо. Вот. САК BAXTER—FOSSIL PTERIDOSPERMS [VoL. 36, 1949] 342 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 12 241. M. anglica var. ioensis Andrews; Fig. 42. . elongata Baxter; Fig. 4 M. Noei Steidtmann, showing only about half of two of the с. three steles. (М. ч 15 50 much larger than апу of the other species represented here that опе сотр! tele at X 2.8 magnification would more than НИ ап entire plate.) The white ring within the black oe ain 4. the upper of the two stele halves) represents а “cambial ring” as shown n figs. ANN. Mo. Bor. Garb., Vor. 36, 1949 PLATE 12 BAXTER—FOSSIL PTERIDOSPERMS [Vor. 36, 1949] 344 ANNALS OF THE MISSOURI BOTANICAL GARDEN ExPLANATION оғ PLATE PLATE 13 Dolerotheca formosa . 44. Transverse-longitudinal view of a portion of a complete campanulum. Below the Be a-b is the longitudinal cut; above, the transverse. In comparison with the рај figure посе the fi lysigenous tubes and absence of a strong sclerotic framework. Fro slide 1627, X 9 PLATE 13 36, 1949 VOL. ANN. Мо. Вот. GARD., BAXTER—FOSSIL PTERIDOSPERMS [Vor. 36, 1949] 346 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 14 Dolerotheca sclerotica 5. 45. Transverse-longitudinal view of entire fructification. Below the line a-b is the longitudinal cut showing the curving of the marginal sporangia; above 1 is the transverse cut showing ты bifurcation of the thick sclerotic bands, between which can be seen the par- enchymatous-secretory tissue forming the sporangial walls. The upper он margin shows cu наса hypodermal zone. For more detailed view see figs. 48 and 49 from slide 1628, X 9, И ЧАЧУООТ ЧАЈ 4 TISSOJA—CLLXVG ANN. Mo. Bor. Garb., Vor. 36, 1949 PLATE 14 ГУог. 36, 1949] 348 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 15 Fig. 46. Dolerotheca Schopfii. Transverse view of a double row of tubular sporangia showing four empty sporangia separated by t n the left and right by the thick Kun bands. From slide 1629, X 22. X campanulum wall showing pubes- See fig. 57 for enclosed o tube between sporangia; sp, sporangium. Fig. 47. Dolerotheca Schopfii. Transverse view c cent epidermis and secretory- 2. ee mal zone; a, glandular hairs. detail of hairs. From slide 1629, X 4 Transverse view of a double row of sporangia sep- Fig. 48. Dolerotbeca sclerotica. Note thin-walled arated from portions of two other rows by the thick sclerotic bands. lysigenous tube alternating radially with the sporangia and bifurcation of the sclerotic nds; sp, sporangia; [, lysigenous tubes. From slide 1628, X 32. Transverse view of campanulum 1. тй ері- x 4 Fig. 49. Dolerotheca sclerotica. From slide 1628, dermis and sclerotic hypodermal zone; sp, sporangia. ‘ig. 50. Dolerotheca formosa Schopf. Transverse view of a double row of sporangia with the alternating smaller lysigenous tubes. а and b indicate the much less conspicuous sclerotic bands than in figs. 46 and 48; sp, sporangia; /, lysigenous tubes. From slide 1627, X 32. ANN. Мо. Bor. Одкр., VOL. 36, 1949 PLATE 15 +: ы ( tol т } = i 1154 M4 * 4 (ТД та * ` BAXTER—FOSSIL PTERIDOSPER MS [Vor. 36, 1949] 350 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 16 Fig. 5 Dolerotheca Schopfii. Transverse view of marginal portion of campanulum: a, broad slime canal and sclerotic zone forming the wall of the fructification partly broken away from inner tissue. See figs. 46 and 47 for detail. From slide 1629 Fig. 52. ee 21. енетін possible attach- ment to Myleoxylon sp. at point indicate ©; P longitudinal view of hypodermal zone of Myleoxylon sp. ойк іп fig. 54. From е1 x 8. Fig. 53. Dolerotheca Schopfi. Longitudinal view of marginal portion of campanulum showing the ascending origin of the outermost sporangia. From slide 1629, Х 8. Fig. 54. Myleoxylon sp. Transverse view of petiole shown in fig. 52b. From slide 1630, 16 PLATE 36, 1949 ANN. Mo. Bor. GARD., Vor. P а Á e “Ey UE а = а-а; же x еф RA, + дө, goes Iu BAXTER—FOSSIL PTERIDOSPERMS [ Vor. 36, 1949] 352 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 17 Fig. 55. Microspermopteris aphyllum. Cross-section of small stem or branch showing departure of branch bundle. This represents a stage shortly above that shown in fig. 3. From slide 1632, X 26. 56. n rospermopteris aphyllum. Cross-section of an adventitious root. From "ms 1635, хз Fig. 57. Dolerotbeca Schopfii. Drawing of epidermal hairs. From slide 1629, X 350. 5 Dolerotheca sclerotica. Drawing of one of the rare glandular epidermal T "Momus slide 1631 59. Medullosa Meu Diagram of central portion of the stem: а, stelar NERO b, line of "aggregate leaf traces" at point just 8 the departure of а petiole as indicated by absence of hypodermal fiber strands shown at c. See fig. 30 for detail of за a single “aggregate trace.” From slide 1626, ANN. Мо. Bor. Слңр., Vor. 36, 1949 PLATE 17 << Y CH fi 9506 95522579 20% ° í ee L ro рање: ad i ; (572 [r Y 2: CHRON ) - Р N KA ред Sd АСА y > = % НЕА ЈАНО 9/2. «529. 9 2, D O9 50256 quo 007007: 97. 29 ГА ii (y A е0. - i ж 76196 9 Doe BL ORR aD ВАН ое A Ф q ..0 94 290986726220 Do фа? (YO e Y "а; p ж 2-6 e ¿ Wades wes ns saat 52466,4 Qu ы BO y : жы лозе by «Жо 9 9 «9 ы POSH COSY , 54 We % 00674000709; ал огои SEE Хр 2 «>»? T sie 44, 5 BAXTER—FOSSIL PTERIDOSPERMS MAIZE АМОМС THE HILL PEOPLES ОҒ А55АМ C. R. STONOR Амр EDGAR ANDERSON INTRODUCTION Maize is widely grown in the Orient and is used there for a variety of purposes. Authorities once quite commonly believed that it originated there, but the demon- stration that it was almost universal in the New World in pre-Columbian times made an American origin seem most likely. From a meticulous investigation of the historical evidence Laufer (1907) concluded that maize did not reach the Orient until post-Columbian times, and Merrill (1941, 1946) produced convincing evi- dence that maize and other New World crops had been carried to the Philippines at an early date by the Spaniards and had been widely spread on the continent of Asia. The subject seemed closed and the lack of any evidence for pre-Columbian maize in the Orient became one of the most powerful arguments against any ef- fective trans-Pacific communication in pre-Columbian times. Several years ago the two authors of this paper came independently to the con- clusion that the subject needed to be reopened and examined on its own merits. The senior author, working in the mountains of Assam, found distinctive varieties of maize widely cultivated by the primitive Nagas. The ethnological and linguistic evidence suggested that these varieties had been in that area a very long time and most probably must have arrived there in pre-Columbian times. Тһе junior author, having made a beginning at distinguishing between the various races of Zea Mays (1942, 1943, 1946), found that the history of maize in the Orient was apparently complex. There was abundant evidence that Merrill was right and that varieties quite similar to those grown in the Caribbean had been brought to the Philippines and Guam by the Spaniards and have since that time been widely spread and ex- tensively grown in the Orient. However, the popcorns, green corns (ie. those used as a fresh vegetable) and brewing corns did not fit into this picture at all. Almost without exception they are grown by primitive peoples. Their distribution is notoriously spotty and is mostly confined to various ethnological back corners. The Oriental popcorns, furthermore, are not at all like the popcorns of Central America. They are on the whole similar to varieties of maize grown in Peru and Chile in early prehistoric times. Ав soon as one was able to distinguish effectively between different races of maize, Laufer's conclusions were no longer valid. His evidence can now only be interpreted as showing that maize in the Orient has had a long and complicated history. Ас an early date the popcorns, green corns, waxy corns, etc., spread widely in the Orient. Аса much later date different varieties of field maize were introduced by the Spaniards, and over wide areas are the only type being grown today. Through the good offices of Dr. W. B. Turrill and Mr. C. E. Hubbard of the Royal Botanic Gardens at Kew, the two authors were put in touch with each other (355) [Vor. 36 356 ANNALS OF THE MISSOURI BOTANICAL GARDEN and since that time have worked together as closely as the distances involved and disturbed world conditions would permit. The collections of native kinds made by Stonor have been grown, pressed, measured, and photographed in Assam. Samples of the same varieties were grown by Anderson at the California Institute of Tech- nology (through the courtesy of Dr. E. G. Anderson) and were pressed, measured and photographed. Pachytene smears were made of each culture with the assistance of Dr. A. E. Longley and Dr. W. L. Brown. In the following paper the evidence relating to these Assamese varieties, evi- dence ethnological, linguistic, distributional, morphological, and cytological, is set out as objectively as possible. Part I was written by Stonor, Part II by Anderson. To the authors the conclusion seems inescapable, that there are at least two races of maize in Asia and that one of these must have crossed the Pacific in pre-Columbian time. The direction (or directions) in which it travelled, however, is still uncer- tain. This new evidence, in other words, tells us little or nothing about the origin of maize. It does, however, enlarge the possibilities which must be considered in any serious investigation of that fascinating problem. PanT I C. R. STONOR Before detailing the uses of maize among the hill tribes of Assam, it is necessary to give a very short general account of the tribes themselves and particularly from the angle of their probable origins and directions of migration. Taking first the tribes of the Assam-Burma border: 1. МАСА TRIBEs:— The Naga Hills are inhabited by at least ten tribes, who, although they possess a number of features of culture indicative of common ele- ments in their origin, show many sharp differences in culture, language, traditions, temperament, and physical characters. The Мара tribes аге, in fact, more correctly described as the tribes inhabiting the Naga Hills. It is well established that they have elements in their culture complexes indicating wide diversity of origin. There are well-established links with Indonesia, Burma (including the Burma-China border), the Pacific and India. The dominant element today is widely different from any of the main cultures of India, and far more bound up with tribal peoples of Indonesia and southeastern Asia (including Burma). It is certain that there has been continuity of settlement in the Naga Hills for many centuries or millenia. Stone celts are commonly found in the hills, although no tribe today has any tra- dition of ever having used them. "There has probably been a good deal of movement to and from the hills from the Brahamaputra Valley. 2. LUSHAI-CHIN-KUKI TRIBES:—These tribes have limited affinities with the Nagas; and their immediate origin seems to be from hill regions of Burma, where they are closely related to the Karens. There are also undoubted kinships with Indonesia. They are pronouncedly Mongolian and have little or nothing in common with any part of India. 1949, STONOR % ANDERSON—MAIZE OF А55АМ Зах 90° 92° в 94° 96° \ B [3 RIBES MISHMIS oR 2 п -28° c < T < тах! SJ j zz S (n = BH UTA N = 4 зс MONBAS AE AKAS це BRA VALLEY = е са HMAPUTRA ; << ко ak NA QW st a . қ RENGMA NAGA наб кој SN омой mi ; ANGAMI NAGA a «e PLAINS оғ ASSAM —— 2+ ERAST PAKISTAN SHOWING THE LOCATION OF 9 tHe Ник Tries Sca ONE NCH = 52 mes © ка NX RSSRM <. *'$1*")H10043 9NO09ULLIHD 3. MANIPUR STATE:—Manipur State is inhabited by а hill population of mixed Kuki and Naga elements, and the big population of the great valley of Manipur is partly derived from these (and particularly the Nagas), and partly from an ancient Mongolian element of uncertain origin. Taking the tribes of the ranges running east and west and bordering the Brahamaputra Valley on the South in Assam: 1. MIKIR TRIBES:—The Mikir Hills are inhabited by mixed tribes of uncertain origin. There are probably links with both Kukis and Nagas and with the peoples on all sides of them. [Vor. 36 358 ANNALS OF THE MISSOURI BOTANICAL GARDEN 2. KHASI TRIBES:— These аге classed with the Mon-Khmer group, and their main origin is generally regarded as being the region of Indo-China. It is quite possible that there is an ancient strain in the population akin to the aboriginal stocks of peninsular India and originating from further West. Тһе Khasis are widely different in many respects from the other tribes of Assam. 3. GARO TRIBES:—The Garo tribes have definite affinities with elements among the Nagas and with tribal peoples of the Plains of Assam. There is also some indi- cation of links with Bhutan, and there may well be kinship with parts of India to the West. Turning to the tribes of the Assam Himalayas lying north of the Brahamaputra: 4. МІЅНМІ, ABOR, APA TANI, DAFLA AKA, МОМВА and LAMAI:—The tribes of the Assam Himalayas, divided extremely roughly into Mishmi, Abor, Dafla, Apa Tani, Aka, Monba and Lamai as the main groups, have been very little studied. There has undoubtedly been much migration from the east and I am of the opinion that a big element in the population is akin to the Kachin peoples of northern Burma. There are marked cultural similarities with the Naga tribes, mainly among the Abors. There has been infiltration down river valleys from Tibet and very possibly in the reverse direction into the hills from the Brahamaputra Valley. The Monbas, a small tribe living near the Bhutan border, may have originated from that state. Even these meagre notes show the hill tribes of Assam to be as mixed a popula- tion as сап well be imagined, who are linked with literally every part of Asia. Prehistoric (?) stone celts are found throughout the hills, indicating very ancient population. "These celts are recorded in the literature from all hill areas south of the Brahamaputra, and I have celts in my possession from the Abor and Dafla Hills of the Outer Himalayas. | Except for the Angami Nagas, the Ара Tanis and the Monbas, and to some extent the Khasis, the basis of subsistence economy is still dry cultivation, carried out by cutting and burning the forest and raising crops for one or more seasons in the area so cleared. The main subsistence crop in the majority of cases is rice, but millets, Job's Tears, sorghum, maize, are all important and are grown in varying amounts by nearly all tribes. In a few instances, one or other of these cereals is as important as rice, as, for example, among the Abor tribes, who grow rice and Job's Tears in almost equal amounts, or the Chin tribes of Burma and the Monba tribe of the Bhutan Border, who grow more maize than they do rice. It seems probable that the complex of Job's Tears-millets-maize preceded rice as the main food supply of many tribes. In general, the tribal peoples of the Assam mountain tracts live in the so-called Neolithic stage of culture. The community is a self-contained опе, growing its own food, weaving its own cloth, regulating its own affairs by tribal law. There is no writing, and the religion is animistic. Тһе level of culture is very similar to the tribes of Borneo, the mountain tribes of the Philippines, and some at least of 1949] STONOR & ANDERSON—MAIZE OF ASSAM 359 the South American Indians. In the Naga, Lushai, Khasi, Mikir and Garo hills there has been missionary activity for some decades, and the tribes have been ad- ministered for the same period, so that far-reaching changes have set in. This is not, however, of consequence for my present purpose, the more so since administra- tion has done remarkably little to change or to develop the economic life. In the Himalayan tracts north of the Brahamaputra, administration has only been started since the end of the War. Missions are banned from entry, and large tracts are still unexplored. DETAILS OF VARIETIES, CULTIVATION, USE, ЕТС., OF MAIZE AMONG THE HILL TRIBES OF ASSAM The notes which follow are all based on information obtained at first hand in the field, either by direct observation or questioning members of the tribes con- cerned, or more usually by a combination of both. Where I have had to rely on the work of others, or have not visited an area, I have drawn attention to the fact. 1. ANGAMI NAGAS:— ( Varieties grown: Nos. 5—18, particularly the larger flint types. Ecology and technique: The Angami Tribe lives in hills from 2500 to over 6000 feet in altitude. Тһе great bulk of the tribe lives above the 4000 feet level and in a sub-temperate climate. Maize is universally grown, and the larger flint types predominate. The Angamis have a remarkable system of irrigated rice-culti- vation, and only a small proportion of their crops is grown on dry fields. Маше is grown mixed either with millet and Job's-tears or else as a pure crop, usually in small plots in the immediate vicinity of the village. Sowing is by dibbling. Uses: Their irrigated rice-cultivation leaves the tribe well provided for. Maize, along with millets and Job's-tears, is important for human consumption mainly among poorer people who are short of rice fields. It is, together with subsidiary crops (notably millets), a catch-crop to "fill up corners" before the rice is har- vested. It is eaten fresh, either boiled or slightly roasted, and is largely consumed by children. The main use among this tribe, in years of good harvests at least, is for pig feed, the grain being parboiled and roughly mashed. It is only so used when actually in season, and is not stored for feeding the pigs. Small quantities of the popcorn types are stored for popping, but this is not important. It is used oc- casionally as an ingredient in beer, along with other cereals. Storage: The ears are hung up on the rafters of the dwelling house for next season's seed. Folklore, traditions, etc.: The Angamis I have talked to simply state that they have grown maize from time immemorial. The tribal name of corn is T'süke. 2. LHOTA NAGAS:— | Varieties grown: "The varieties grown in the village of Yimbang at elevations of less than 2000 feet in the sub-tropical lower ranges of the area are: 1. Tchetum-sopfu (Serial L. 1), meaning "Small late,” regarded as an in- digenous variety. [Vor. 36 360 ANNALS OF THE MISSOURI BOTANICAL GARDEN e Moro (Serial L. 2), meaning "Quick or early eating," regarded as indigenous. Aorr-chemyang (Serial No. 6 of first consignment), meaning "Ao Nagas blood." This refers to traditional origin from the neighbouring Nagas, the term blood having reference both to the small red grains, and to the former enmity between Lhotas and Ao’s. Konoma-tsunghundhro (Serial No. 8 of first consignment), meaning "Konoma Maize," the name Konoma being that of a large Angami Naga village to the south of the Lhota country, and indicating that it probably derived from there. А very similar type is known as Wokotsu, because it was first got by Yimbang from a Lhota village of that name. Epuk (Serial No. 12 of the first consignment), meaning “Bursting, is only grown for popcorn. Kor-chak, meaning “Horse-Food.” This variety, a coarse flint maize grown in many parts of India, was obtained within living memory from Nepalese immigrants into this region. The name is derived from the custom of the “Ы ” > as it мл - Nepalese of keeping ponies. The main varieties grown in the village of Yekhum, in the higher areas of the Lhota country and almost within sight of Yimbang, where conditions are between sub-tropical and sub-temperate and the altitude about 4,000 feet, are only two: (1) Korcbak and (2) Konoma-tsungbundbro. Both are said to be of fairly recent introduction, and supplanted other types. Other types may be grown in small amounts. Ecology and technique: А great part of the Lhota Naga country is sub-tropical, and there are few villages situated over 4,000 feet. The types grown are naturally those most suited to this climate. Maize is grown as a mixed crop among the rice and subsidiary cereals. It is commonly sown in rows along the edges of field paths ог inter-field boundaries. The seed is dibbled, and the sowing season is March- April, while the harvest is from June to early August. Uses: Maize is not a very important crop among the Lhota Nagas and, as among the Angamis, it is grown wholly as a catch-crop, utilized before the rice harvest is in. But it is grown entirely for human consumption, and except for the popcorn Epuk variety, it is entirely consumed as it ripens. A little is used now and again for beer-making. The reasons for growing the different types is gone into in more detail below. Storage: The ears for next season's seed are simply hung up on the rafters of the dwelling house. Traditions, folklore, etc.: The general word for maize is Tsungbundbro, which and I have been told of a М means "something obtained from the Angami Nagas,’ vague tradition that the Lhotas first got maize from their Angami neighbours. On the other hand, the variety Tchetum-sopfu, listed above, is regarded by the Lhotas as their own maize and does not seem to be grown by the Angamis. 1949] STONOR & ANDERSON—MAIZE OF ASSAM 361 The Lhotas have been in their present country for a matter of centuries. They state that they have had maize from time immemorial. 3. АО NAGAS:— Varieties grown: Some or all of Serial Nos. 5—18. Ecology and technique: Closely parallels that of the Lhota tribe. Uses: As among the Lhotas, it is entirely for human consumption. Storage: As for the Lhotas. Traditions, folklore, etc.: The Ao Nagas have a tradition that they have always grown maize among their field crops. The usual name for maize is Mentia, for which I can get no translation. A small group of villages which have a different language know it as Achang-Tangba, which means “bearing rice already husked.” A curious local name has been recorded by J. P. Mills (“Тһе Ao Nagas,” 1926. p. 125, footnote) as used in the Ao village of Changtongia: Moya zungkbum, mean- ing “бета Naga Lentils.” 4. SEMA NAGAS:— Varieties grown: Some or all of Nos. 5—18. Ecology, technique, etc.: Very much as for the Lhotas and Ao’s, except that maize is a much more important crop and is planted thicker on the ground. Sowm during March-April, and harvested in June—August. Uses: Is of considerable importance for human consumption, both fresh on the ear and (to a lesser extent) stored and pounded to mix with rice. It is also used to make several types of strong beer, again to a greater extent than among other Naga tribes. My information from the Sema area is scanty. However, the im- portance of maize is very likely linked with the shortage of land for cultivation and the large population of poor people, so that the staple food of rice is apt to be finished well before the next harvest. Catch-crops are consequently of much importance, and rice insufficient for beer-making. Traditions, folklore, etc.: The Semas regard maize as an ancient crop. The name for it is Kolakithi. This is usually translated as "Foreigner's Job’s-tears,” and some Europeans have assumed without evidence “Foreigners” to mean the British and that maize was introduced into the Naga Hills by the British (!). Apart from this, I have been given another meaning for the word by a Sema Naga: "Grain that is eaten by plucking singly.” Another word is Amehukethi, meaning “a grain ripe and eaten before the rice." 5. CHANG NAGAS:— Varieties grown: Some of Nos. 5-18. Ecology, technique, etc.: As for other Naga tribes. Most of the Chang Naga country is high and cold with sub-temperate climate, and the larger coarser flint types predominate. A small popcorn (Serial No. 12) is grown, and the African dent maize is also cultivated. [Vor. 36 362 ANNALS OF THE MISSOURI BOTANICAL GARDEN Uses: Except in a few low-elevation villages, the Chang Nagas even today depend largely for their subsistence on millets, Job's-tears, maize and taro, rather than rice. Maize is thus an important crop. It is, however, largely a catch-crop, eaten fresh, but appreciable quantities are stored and are mixed with other cereals for food during the cold weather. It is used preferably in making beer, and to a small extent for popcorn. Some, particularly in seasons of good harvest, is fed to the pigs. Storage: In round baskets, stripped from the cob, in separate granaries. Traditions, folklore, etc.: The Chang name for Maize is Hangi, for which I can get no translation. It is certain from their traditions and recent history that the Chang Nagas have been pushing west from the Burma side for some generations, and they themselves claim that at least a large element in their composition came from further east. Their economy is as noted, still based very much on Job's- Tears, millets, and maize. This is partly due to their living largely in high alti- tudes and possessing no suitable type of rice for the climate; but I have several times been informed by Changs that it is only since they have had contact with the more westerly Naga Tribes that they have taken to rice at all; and rice is widely recog- nized as being a more recent crop than the other cereals, maize not excluded. Among this tribe each major crop has its own minor tutelary deity, maize among the rest. The African dent maize is known as Bilati Hangi, meaning "English Maize." 6. YIMCHUNGRR, KONYAK and KALYO-KENGYU NAGAS:— I have very little information from any of these tribes. Maize is grown in varying amounts by them all. It is said to be an important crop among the Kalyo- Kengyu, but not very much grown among the Konyak Nagas, who are among the most ancient stock in these hills. I have visited the extreme northeast area of the Konyak country, where a poor type of one of the larger-grained flint maizes is grown in small quantities. I was told that the people were not keen on it as it attracted bears to the fields. The Konyak name for maize is Tongi. 7. KACHA NAGAS and NZEMI NAGAS:— These two closely allied groups inhabit the southern Naga Hills. I have been there once and can confirm that maize is grown in the fields along with other cereals, and my original stock of small-grained types came from this area. Тһе area varies a great deal in elevation (from 1,000 to 7,000 feet) and most or all of Nos. 5—18 are grown. I have a rather vague report from a Government subordinate concerning a branch of the tribe living in the hotter areas, about 3,000 feet, that three main types are recognized: (1) Imbaume mei Mitak, early; (2) Lingtak, second to ripen; (3) Lingtak tiingne, late. The first two are said to be the most widely grown. 1949] STONOR & ANDERSON—MAIZE ОҒ ASSAM 363 The Kacha Nagas have а tradition that the tribe first emerged from a cave іп their area, and brought their old established crops with them, maize being among them. I have no details of its use except that most or all the crop is used as it ripens and is not normally stored. 8. SANGTAM NAGAS:— There are two geographically separated branches of this small tribe. I have visited the northern branch. Varieties grown: Y have obtained the following list and notes from a Govern- ment subordinate, himself a member of the tribe. Chemese, sown in March and harvested June to July. Mesease, meaning "sweet maize." Is regarded as good for making beer and is grown mostly in rocky fields. Nuracbese, a small popcorn maize. Hengchimerem, used both for human consumption and for pig feed. Yengcbengese, used mainly for beer and for pig feed. Abocbese, not regarded as very good, but grown as it is ripe earlier than the rest. Unfortunately, my small collection from the northern Sangtam area got rather mixed and labels lost. I can, however, note that of the above No. 2 15 a small red- grained type similar to but larger than Serial 6. Хо. 2 (Mesease) is a very small white-grained popcorn identical with Serial 12. Мо. 4 (Hengchimerem) is a fair- sized maize with large, coarse grains, and a proportion at least are red. No. 6 ( Abocbese) is a stout rather coarse type with large hard yellowish-white grains. Ecology, technique: The northern Sangtams live in steep country, and grow their maize mixed with millet in their ordinary fields. Use: The above list shows the main use of maize and I have little to add. Most is said to be eaten when soft, but a proportion is stored for the winter, when its main use, except in seasons of rice shortage, is for beer. It is a fairly important catch-crop, probably more so than among the Ao and Lhota Nagas. Much of the Sangtam area (as among the Chang Nagas) is high and cold and unsuited for rice No ~ nu + У growing. Storage: In baskets, after shelling, in the granary. Folklore, traditions, etc.: The Sangtams simply informed me that they have always had maize among their crops, as far back as their traditions reach. 9. RENGMA NAGAS:— I have never visited this little tribe, which is split into two geographically sep- arated parts, of which the eastern branch is very primitive. Writing of the eastern Rengmas, J. P. Mills ("The Rengma Nagas," 1937, p. 86) states: bbs g R maize is a far more ques = than it is in the Western Rengma oth eaten boiled and used for bre Ordinarily it is sown scattered among is "Millet es the Sorghum) with which it is k; ira, but some men grow whole бе Ids of it. [Vor. 36 364 ANNALS OF THE MISSOURI BOTANICAL GARDEN Of the western branch of the tribe, the same writer says (l.c., p. 77): “Маше —though an important crop among the Eastern Rengmas, is little grown by the Western Rengmas. At most a few plants are grown among the rice and the heads roasted and given to children to eat. Occasionally a brew is made from it now- adays, but the resulting beer is not popular." The words for maize are (both from Mills): Western Rengma—Samphuruchi, and in Eastern Rengma—Akbhuzi. Mr. Mills has recorded from the Eastern Rengmas (p. 88): Early in August there is a series of five © Jue ' days. The first day is called Vei желе food. 2 апа 15 held со prevent waste of he next four are called Tsate, = "wam x p ps first fruits, that marks the beginning o Fs: maize and millet harvest. fields and bring back leaves E D x nly the women eat, or pre also bring in Maize-heads t e are not eaten that day, but ња с ‘ull the next чу but опе following, when i eats roasted Maize. Throughout this “genna” all wo ceases and people entertain woe friends at dizi partie 10. THE LUSHAIS:— Varieties grown: 11-18 as sent. A list of the types grown, as sent me Бу a reliable government subordinate is: 1. Ralte Vaimin (Serial L1), regarded as indigenous. 2. Pawi Vaimin (Serial L5), meaning "Chin Maize." 3. Sap Vaimin, meaning "European Maize." This variety is a dented maize recently introduced by the British. 4. Bawngpu Vaimin (Serial L7), meaning “Herdsmen’s Maize.” This is a large coarse flint maize introduced by Nepalese immigrants (the "Herds- men"). 5. Chingzo, meaning not given. 6. Vai Vaimim (Serial L4), meaning not given. 7. Lenliam (Serial L6), meaning not given. Nos. 5, 6, and 7 are regarded as similar, and have small, dark-grained ears noted for the stickiness of the grain while soft. 8. Puakzo (Serial No. L8), meaning "Burst all," as it is grown for use as pop- corn. Ecology апа technique: I have not visited the Lushai Hills, but have first-hand information that, as among the Nagas, maize is grown among the other crops in the dry jhum fields, and occasionally as a pure crop. In the latter case I am in- formed that it is usually followed by a legume crop. The seed is dibbled, and the sowing season is said to begin in April, with the harvest from July to August. Uses: The bulk of the crop is eaten, boiled or roasted, as it ripens, and is thus a catch-crop as among the Naga tribes. It is also used to some extent as pig food, and a little fed to poultry. Limited amounts are stored (presumably by poorer people, and in lean years), and the grain is roughly pounded and mixed with rice. The small Puakzo is probably grown entirely for popcorn. I have no information of maize being used for beer, and I think it to be so used very seldom. Some of the 1949] STONOR & ANDERSON—MAIZE OF А55АМ 365 Lushais іп the high-altitude eastern areas with а cold climate use maize as а main crop; this is due to lack of a high-level rice, and will be described under the Chin tribe. Storage: I have no information. Traditions, folklore, etc.: Lt. Col. J. Shakespear ("The Lushai Kuki Clans,” 1912, p. 87) states that there is a festival known as Mim-Kut, named after the Maize, as it takes place when the crop ripens. It is of but little importance and seems likely to die out. Cakes of Job’s-tears are eaten, and the next day is brilb. N. E. Parry (“А Monograph on Lushai Customs & Ceremonies,” 1928, p. 91) refers to this feast as being in honor of persons who have died during the past year. He mentions that the “Fresh vegetables, maize, bread, necklaces, & cloths are placed on the memorials of the dead.” Shakspeare remarks of the chiefly clan of Fanai, who originated from the Chin country to the East (l.c., p. 139): “А dead Fanai is buried in the usual Lushai way, but no rice is placed in the grave. An offering of Maize is however suspended above it. It may be noted that in the Zahao country rice is not cultivated, the staple crop being maize.” It is possibly an exaggeration that maize is the only main crop in this area: but I have had it confirmed by members of the Lushai tribe that it is more important than rice. 11. THE LAKHERS:— I have not visited this tribe, which lives next to the South Lushais. Their sys- tem of cultivation is the same as among the Lushais, and it is reasonably certain that maize is grown in the same way, and is of the same importance. The Lakher word for Maize is Chhamei. Rice is the staple crop. A minor use of the dry grain is mentioned by Mr. N. E. Parry (“Тһе Lakhers,” 1932, р. 199), who states that it is used for counting. The same authority describes a very interesting dance performed in one village in connection with the maize harvest (1.с., р. 434): Іп Chapi village, to mone the gathering in of the Maize harves st, a dance called Pazutawla is performed. The de hands and form a ring; the girls stand in front o them; one girl е between two men, and puts ап arm around the Hoddi of the men on each side of her. The j. par and round, singing to dba cco ompanim ent of gongs and drums. ТҺе dance is peculiar to C It is апа, to dance it except in celebration of the maize harvest, and were it Portion к any other time those taking part would suffer from carbuncles. 12. THE CHINS:— In the Chin Hills, which are politically within the borders of Burma, maize is a very major crop. All my information is from “The Economics of the Central Chin Tribes," by H. N. C. Stevenson (1943). Stevenson lists maize as among the staple crops (p. 35) and states that it is grown along with millets and beans. The system of cultivation is as a mixed crop in the jbum fields. The ears are stored, hung from the rafters of the dwelling- house. He implies that it is of equal importance with rice. It seems to be used [Vor. 36 366 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN both fresh and after being stored; it is used, mixed with rice and millet, for beer- making. Stevenson gives interesting notes on uses of stored maize. Thus under “Travel Rations,” which he describes as being the most common form of cooked food in pre-annexation days, owing to constant danger of raids, he includes . . .: Vainiim Аап.—" Аз the name implies, this is roasted maize, and it is prepared in exactly the same way as Vai kan. Occasionally, when it is to be eaten the same day, bananas are pounded into the grain as a change of flavour." He describes the Vai kan just referred to as follows: Тһе name means roasted millet, еч klawr again being the variety used. The grain is soaked in water for a minute or о damp it ae and is ig eyar E if t Lite it for fermenting, after dore it is spread o о dry in e final ges of preparation the swollen grain is roasted in a s Рд vol it rase КА рата и Тһе whole 15 then pounde чу p alt or Hw made into convenie which are taken on m unting parties, e ai kan is ы, as Ка 2 all foods for ku to the ribs” but it is very dry, е M is salted variety, and for this reason is not used during hot жыды іп areas where water is scarce...... The peibung is the vaina grain-roasted pot, shaped rather like a bee yee with a large hole in one side. Its existence proves the antiquity of the uU ptc of cooking. The same authority lists under the heading "Daily food in the home": Var: This is the commonest food of the present day, and it is generally made with maize, T millet = reserved for beer- -preparation or the travel ration he maize is pounded t to break the grains and then sieved and boiled in water. It is eaten with ind sauce t es is going, after the liquid res been drained off. When consumed as a bro ith the water in which it is cooked it is called 77 sawp, and if pumpkin and other {йык аге рија asa res it becomes bub ber The harvest is apparently later than in most of the Assam Hills, presumably for climatic reasons. Stevenson states (1.с., p. 41): sooner is the millet crop du gathered than the maize ripens in August and Sep- нане Та = rvesting maize also the Chin use no knife but tear the cobs off by hand, re- moving the outer cover and turnin gom inner ones back to act as ties when the cobs are sod чи d under the rafters of the house. With regard to the use of maize for beer-making, Stevenson records (l.c., p. 113): the Central Chin Hills there are three recognized types of beer: zu ba, made from вое quality 1 — -— ти pi, made of hu ^ d millet mixed with its chaff; and vainiim made from Maize ...... vainiim zu] is made of pe Maize, which ril IM we ^ for Apre often is treated like the mille: used for а. Li 4, it will n keep for hls ву is therefore Prepared shortly — use. "This zu being кај к 15 most often used іп «һе home оп day-to-day occa Maize seems to be too important for human consumption to be given directly to domestic animals. Stevenson notes, however (p. 114): “Incidentally, fermenta- tion of grain for beer does not waste the solid residue: this by-product is added to the pig-food and gives strength and substance to the unappetizing bulk of banana stalk which 15 the other main ingredient.” Ав а corollary to this, 1 may add that this use of the fermented grain of millet, rice, maize, etc., for feeding pigs is а 1949] STONOR & ANDERSON—MAIZE OF ASSAM 367 universal practice among all the Nagas and other tribes. The Chin word for Maize is Vainiim. 13. THE KUKIS:— The Kukis live in the Southern Naga Hills, and in Manipur State. I have once visited a few Kuki villages in the former area, among whom maize was grown and used in precisely the same manner as among their Naga neighbors: as a catch-crop for consumption while fresh, for popcorn, and to a small extent for beer-making. The same types are grown. The name for Maize among the Thadou branch of the tribe is Kolbu, which is said to mean "Burma Food": an interesting point since the Thadou Kukis are generally regarded as having close kinship with Northern Burma. 14. THE MIKIRS:— The Mikir tribes have been very little studied. Their system of shifting culti- vation is very primitive. Maize is among their subsidiary crops. I have been told by a member of the tribe that the large-grained types are most grown, but the small types are also used. The Mikir word for maize is Thengthe. I have not been into the Mikir Hills. 15. THE KHASIS:— Varieties grown: 'The Khasis have for generations been in close contact with the British, Nepalese and Indians, and Asiatic, African and American varieties have been introduced and are by now inextricably mixed with older types. Most of the tribe cultivates at over 4,000 feet, and it is doubtful if the small-grained types were ever used to any extent. Ecology and technique: Аз a mixed crop, together with rice, millet and Job's- tears. Nowadays the Khasis have a large-scale potato-growing industry, and maize is often grown in the potato plots, interspersed between the rows of potatoes. It is also grown as a pure crop. Uses: The original uses were for food, mainly as a catch-crop, the ears being eaten boiled or roasted. Some is put aside for popcorn, and in parts of the area it is prepared by placing the grain, with some sand, in the cooking-pot, and heating it over a fire. Maize is become a fairly important cash crop for sale in the bazaar at Shillong. It is used to some extent as pig food. Sforage: Very little is stored, except for seed. Тһе ripe ears are hung up in the house. Traditions, folklore, etc.: The Khasis state that maize is among their ancient crops. The word for it is Riew Hadem, meaning "Grain of the Hadem People." he Hadems are a small branch of the Kuki tribes who live in close contact with the Khasis in the extreme southeast of their area. 16. THE GAROS:— Varieties grown: G1-G5. Ecology and technique: The seed is dibbled in lines among the rice and other crops in the jbum fields. It is sown about March in most parts and harvested in [Vor. 36 368 ANNALS OF THE MISSOURI BOTANICAL GARDEN June. In some of the warmer areas it is often planted earlier, and I have seen plants with the ear already formed in April. Uses: Although the staple crop of the Garos is rice, they are hard-pressed for land and food is often short. Maize is an important catch-crop, and is almost entirely used as it ripens. The fresh ears are either boiled or roasted. Very little is stored, and this stored grain is pounded and eaten boiled, either alone or with rice. The smallest variety grown is used only for popcorn, and is prepared by heating it in a cooking-pot with a little sand. Maize is too precious for human consumption ever to be used as pig food; and is said not to be used for beer, which is made entirely from rice. Storage: When stored for winter use, it is stripped from the ear and put in round baskets. Stock for seed is hung up in the dwelling-house. Traditions, folklore, еѓс.: Тһе Garos say they have always grown maize, and got it with their main crops "out of the ground." % o All the foregoing tribes live south of the Brahamaputra, which has always been an effective, but by no means absolute, barrier to contacts of culture. The notes which follow describe the cultivation and uses of maize among the tribes of the outer Himalayas, within the political boundaries of Assam. 17. THE DAFLAS:— Varieties grown: Serial Nos. 1—4. Ecology and technique: Maize is grown as a mixed crop along with rice and millets, under the same system as among the Naga tribes. It is the earliest crop sown, and is often dibbled in before the rice, the usual month being March, although I have seen fields sown in February. Uses: Rice is the staple food among this large tribe, but their technique of shifting cultivation is slovenly and haphazard, and probably for this reason, sub- sidiary cereal crops of millet and maize are of considerable importance to supple- ment the rice supply. Maize is both grown more than among the Naga tribes, and much more, perhaps most of the crop, is stored for winter use. I have not been to the Dafla country while the maize was ripening, and cannot say to what extent it is eaten fresh and functions as a catch-crop. However, in normal seasons, it is a subsidiary crop to help out the rice in the winter rather than a catch-crop for im- mediate use. In the summer following a poor harvest, the people would, of course, be forced to use their maize as soon as it was developed enough to eat. The main use of the stored maize is for food, since the grain is more easily broken up and mixed with rice. For this reason, the smaller-grained of the main types is preferred since the grain is more easily broken up by the pestle-and-mortar grain- pounders. Sometimes it is crudely milled between flat stones. А proportion of the maize is mixed with millet (Eastern Daflas) or with rice (Western Daflas) for beer-making. Іп good seasons it is used for pig food. Even the dried ears are 1949] STONOR % ANDERSON—MAIZE OF А55АМ 369 often roasted and eaten off the cob. I have several times seen them given (о chil- dren as "snacks" between meals. All types are used for popcorn. For some years past, Daflas living fairly near the Plains of Assam have carried on a small trade in maize with tea-garden coolies. The Daflas grow only the larger-grained types, and have far fewer types, partially accounted for by the different status of the crop since the large-grained types are clearly more economical for storage than are the small-grained. Climatic factors may, however, enter into it, and it is also perfectly possible that the Daflas have never had the smaller types as grown on the Assam- Burma border. Storage: Maize is stored in the granary along with other grains. Among the eastern (and more primitive) section of the tribe it is kept on the cob, and the ears are heaped up in a roughly-made rectangular bin in one corner. The western Daflas strip the grain from the cob and store it in large round baskets, usually with the cobs wedged over the top as a protection against rats. Traditions, folklore, etc.: Y have several times questioned Daflas оп this. They simply state that they have always grown maize. Тһе Dafla word for maize is Topotbe. The smaller type grown (Serial 4) is known as Nyamatup. 18. THE ABOR TRIBES:— I have only visited the Padam and Minyong Abors. The other main tribe are the Galong Abors, whose country is adjacent to the Daflas and who are probably very closely related to them. Тһе following note deals only with the Minyong and Padam Tribes. Varieties grown: Nos. 1—4 (of first consignment). Ecology and tecbnique: 'The Abors grow their maize as do all other shifting cultivators as a mixed crop among their rice and millet. They are, however, unique in that they practice double cropping. The first crop is sown during the month of February and is harvested in June. As soon as it is ripe, a second sowing is made in the same field, and is harvested in October. Uses: The first crop of maize is used purely as а catch-crop, and is eaten while the ears are soft, either boiled or roasted. "The second crop is left standing until the ears are thoroughly ripe and the grain hardened off. It is then stored and used during the winter either as а food or for making beer. The proportion that goes for beer-making or for food is said to vary with the economic position of the household and the general abundance of the crops in any one year. In general, maize is an important crop, although rice, Job’s-tears, and millet are the staple foods. It is used to some extent for pig feed. Popcorn is also made, but not to any extent. Methods of Storage: The grain from the second sowing is stored, stripped from the ear, in round baskets in the granary. Folklore, traditions, etc.: The Abor tribes simply state that they have always had maize among their crops. Their folklore credits them with having obtained [Vor. 36 370 ANNALS OF THE MISSOURI BOTANICAL GARDEN all the main crops “Оп the horns of the tame Bison," and I have been told specifi- cally that maize is included. The name among the Minyong and Padam sections of the tribe is Sepa. Тһе main type used (a large golden or red flint maize) is called Pade-Pasing Sepa. 19. AKA TRIBE:— I have not visited this small tribe, but I have met and talked with several mem- bers of it. They all told me that their most important crop is maize, which is grown in their fields along with dry rice. According to my informants, the Aka tribe originated in the Assam Valley, whence they were driven up into the hills many generations ago. At that time their main (or only) crop was rice, and they learned to use maize from the Monba tribe immediately to their west. They in- formed me that they grow the same varieties as the Monba tribe. Тһе average altitude of cultivation in the Aka Hills is of the order of 5,000 feet. 20. MONBA TRIBE:— Varieties grown: 1. Phentang. Serial Nos. 1 & 2 of first consignment. 2. Num Phentang. Serial No. MI. 3. Khana Phentang. The first two are indigenous: the Khana is a South African dentate maize intro- duced a few years back by the government. Khana means tooth. Ecology and technique: The Monbas differ from all other tribes of the Assam Hills in that they are a civilized people whose whole culture and social organization is of Tibet. They have а well-developed system of farming on permanent fields. Their maize is grown as a pure crop in rotation with other cereals and in the permanent fields. It is sown in May and June, and is harvested from late September to early November. The lateness of the sowing as compared with other areas is due to the need of fitting it in with their system of rotation. Sowing is entirely by broadcasting, and the ears are pulled off by hand. The variety Phentang is the most grown, and is in large fields. The variety Num Phentang is grown in small amounts, and is relegated to odd corners, edges of fields and to the few areas of shifting cultivation. The introduced dentate maize, Khana Phentang, is almost entirely relegated to the areas of shifting cultivation. The reason for the different technique will be clarified below. Uses: Тһе Monbas, with their advanced economy, grow a variety of grain crops, all of some importance. These include maize, barley, rice, buckwheat, millets, Chenopodium, cockscomb. However, with the exception of a few villages at 9,000 feet and above, the most important crop is maize, which is said to be more important than the others put together, and is the staple food. The main uses of the grain are: 1. For the ordinary food, the variety Phentang is ground to a coarse flour Бу 1949] STONOR & ANDERSON—MAIZE OF ASSAM 371 means of the ordinary Tibetan water mill. The flour is usually boiled. This flour is known as Phentang buku. 2. А small proportion of the same variety is eaten when the ears are still soft. 3. The Phentang is slightly roasted and then flattened and partially broken up by light pounding. In this form it is chewed as a relish, especially while liquor is being drunk, and plates of it are always offered to visitors. The local name is kakung. A great deal is consumed in this way, and the kakung is also an article of trade in Tibet. 4. The Phentang is used to a fair extent in brewing of beer and distillation of a weak spirit. This is said to be the use of the introduced variety Khana Phentang. 5. For popcorn, the variety Num Phentang only is used, and it is grown for that purpose only. It is popped by heating in an earthenware vessel with a little sand. As far as I know, virtually all the maize crop is used for human consumption only, except that the "leavings" from manufacture of liquor go to the pigs. Ву ar the most important type is thus the Pbentang, а strong-growing, large-grained golden-yellow or occasionally red, flint type. Methods of Storage: This aspect is also interesting. When the maize is ripe, the ears are left for some weeks on the plant to dry. They are then gathered, and the enclosing leaves removed. The ears are then stacked on the fields in solid rectangular panels up to 20 feet long, by 10 feet high, and perhaps 2 feet thick, the structure being raised off the ground by a slight framework of sticks. The panels are often L-shaped. А field, when the crop is stacked, is a beautiful sight dotted with these solid patches of golden yellow, the more so since the few scarlet ears are suspended over the top for color effect!. The crop is left thus for a month or less to dry off thoroughly, after which the ears are carried home to the village, and the grain removed either by hand or threshing with a short stick. It is stored in large round baskets in the upper room of the two-storied house. The seed for next season is left on the ear and hung up on the rafters. Traditions, folklore, etc.: The Monbas simply state that they have always grown maize, and that as far as their traditions go back it has been the staple food crop. On the other hand, they have a well-established tradition that they first got their rice from the Plains of Assam. Ву religion they are Buddhists (every other tribe. in the Assam Hills is Animist), and it is both interesting and significant that the first ears of maize to be gathered are placed in the village temple as an offering, while they deny any such practice in the case of their barley which (together with their Buddhism) they must have obtained from Tibet. It is also a common sight to see a few ears of maize hung up as offerings inside the temple, or placed on small wayside shrines. I have asked members of the tribe if they have any special rites, dances or festivals for their maize, and in all instances this was denied. I wou “It is possible that the original purpose of suspending the red ears was to ward off evil spirits from the fields. [Vor. 36 372 ANNALS OF THE MISSOURI BOTANICAL GARDEN not, however, like to state categorically that my informants were accurate. In dealings with tribal peoples knowledge of religious custom can only be got by long and close acquaintance or direct observation. 21. THE APA TANI TRIBE:— Varieties grown: Serial Nos. 1-4 of first consignment (as for Райа Tribe). Ecology and technique: 'The Apa Tanis, although a primitive people without writing, etc., have a very highly developed system of irrigated rice cultivation for their staple food supply, and practice shifting cultivation extremely little, if at all. The maize is grown in small quantities in little garden plots in and around the village. The tribe inhabits a small area in the Dafla Hills and the varieties grown are the same. Uses: Maize is not an important crop. А proportion of it is eaten soft on the ear. Some is stored, and is prepared by first roasting it in a clay pot and then pounding it to a coarse flour. It is said to be eaten mainly by old people with decayed teeth, but this needs checking. Popcorn is prepared in considerable amounts, and is made by putting the grain into the glowing embers of the fire, and picking it out with bamboo tongs as it bursts. I am not sure if the other method of heating the grain in an earthenware pot is used. Methods of Storage: The small quantity stored is kept with the rice in separate granaries. Traditions, folklore, etc.: Y have no information. 22. THE MISHMI TRIBE:— I have once visited a small section of this very primitive tribe, and have only the scantiest information. Their shifting cultivation is rough and ready. Maize is included among the crops and Mishmis have told me that it is of moderate im- portance. І have seen Mishmi women making popcorn by putting the grain in the edge of the house fire, and picking it out as it bursts. MAIZE IN HILL AREAS OUTSIDE THE BOUNDARY OF ASSAM BHUTAN:— I have talked to many traders and others from Bhutan and am told by them that maize is grown everywhere, and is a fairly important crop. А large yellow type is said to be the most widely grown, and is presumably the same as the Phentang of the Monba tribe on the Assam-Bhutan border. SIKKIM :— As for Bhutan. Writing a hundred years ago, Sir Joseph Hooker (“Himalayan Journals," Vol. II, p. 78, footnote) states that he was given popcorn in North Sikkim. He describes it as, "Called pop-corn in America, and prepared by roasting the maize in an iron vessel, when it splits and turns partly inside out, exposing a snow-white spongy mass of farina. It looks very handsome, and would make а beautiful dish for dessert." Hooker also records (Volume I, p. 157) seeing, in May, 1848, the maize just sprouting in North Sikkim. Не goes on to record the 1949] STONOR & ANDERSON—MAIZE OF ASSAM 373 curious statement that “This plant is occasionally hermaphrodite in Sikkim, the flowers forming a large drooping panicle and ripening small grains: it is, however, a rare occurrence, and the specimens are highly valued by the people.” Un- fortunately, he does not tell us if he actually saw such plants, or if he was recording from hearsay. Maize is among the crops of the small Lepcha tribe of Sikkim. In his book “Himalayan Village” (published in 1938) Mr. G. Gorer refers several times to maize cultivation. He notes (p. 95) that wheat and maize are used for food when rice is short, but states that “they are not liked.” Не notes that the maize is sown in March (p. 94). This is confirmed by Major J. Morris in his book “Living with the Lepchas” (1938, Chapter 9) where he several times refers to maize as being among the crops raised by this tribe. NEPAL:— Of this region, I have the scantiest information. Nepalese immigrants into Assam have told me that they grow considerable quantities of maize in their own country. Maize is an invariable crop of these immigrants, even when they settle in the hot plains of the Brahamaputra Valley. They grow large yellow or white forms, the same or very similar to the main type of the tribes of the Assam Himalayas (Abors, Daflas, Monbas). BURMA:— Unfortunately, I have been able to get no information concerning the Shans, Wa’s, Karens, or any of the hill peoples in Burma except the Chins as recorded above. Of the large Kachin tribe, who live in the extreme north of the country and whose hills are contiguous to the Mishmi Hills of Assam, Mr. J. L. Leyden has recorded (introduction to a pamphlet “The Kachins of the Hukawng Valley” by Kawlu Ma Mawng, 1944, p. ix): | The Kachins аге rice eaters, and their agriculture is mainly concerned with the produc- tion of ұла staple food. Kachin agriculture is chiefly concerned in the extremely wasteful and inefficient shifting cultivation ...... he 275 апа towing i crudely рах out the Kachins invariably find themselves «i a poor crop of rice and are compelled t work subsidiary crops of maize, millet and yams to көм starvation. While this somewhat bald statement as to why maize and millet are grown can hardly be accepted at its face value, especially vis a vis the tribes of Assam, it at least indicates that maize is an established subsidiary crop, probably in very much the same way as among the Naga tribes. MANIPUR STATE:— The Naga and Kuki tribes of Manipur State are of the same stock as those of the southern Naga Hills (see above under Cachar Nagas and Kukis), and it is reasonably certain that they grow the same maize and in the same way as do their neighbors. The Manipuris proper inhabit the highly irrigated plain of the Manipur Valley. I have no information as to whether they grow maize, but consider it un- likely that it is of any importance as their fields give them a considerable surplus of rice. [Vor. 36 374 ANNALS OF THE MISSOURI BOTANICAL GARDEN HISTORY IN THE ASSAM HILLS As we have seen, maize is grown by all the multifarious peoples of the region. This wide distribution does not in itself mean that it is of necessity an ancient crop. Once introduced and given favorable conditions for its spread, any crop can spread with rapidity, even in remote areas during a very few generations. This has taken place in the Assam Hills, a good instance being the potato. Another such crop is manioc, which is now grown in at least three widely separated ranges of hills. But both these crops are universally recognized as being introduced after the coming of the British. In the Garo Hills I was told by illiterate villagers that manioc has been grown by them for about twenty-five years; and in the northern Мара Hills it is said to have been with them for two or three generations. But in the case of maize I was simply told by every tribe that they have always had it among their crops, and any suggestion that it came from outside was ridiculed. An exception to this, as recorded above, is the Aka tribe, a small tribe of the Assam Himalayas, who believe their maize to have been obtained from neighboring tribes many generations back. I have constantly inquired in tribal villages, and particu- larly among the old men of the community, as to when and how they first. got their main crops. The reply was always the same in regard to millets, taro, maize, Job's-tears, and yams. They have always been grown, and no one can say when ог how they were first obtained. In some cases this is true with rice, now the staple crop among the majority, but there are indications that rice is a more recent food than the rest, although it is undoubtedly a very ancient crop and has been grown in Asia for some thousands of years. Thus, among the Chang, Yimchungrr, and some of the Konyak Nagas, the people even now depend on roots and cereals other than rice for their food supply and state quite openly that they are older crops, maize not excluded. The same is true of the Monbas, who have a definite tradition that maize has "always" been their main crop, while rice was obtained many gene- rations ago from the Plains of Assam. Furthermore, in the case of rice there are established legends to account for its origin. They have been recorded for the Naga Tribes by Professor Hutton and Mr. Mills (in the series of monographs on the Naga Tribes), and for the Lakhers by Mr. Parry (“Тһе Lakhers," 1928), while I myself have been told folk tales by the Daflas, Minyong, Abors, and Khasis. But there is no legend known to account for the origin of the other cereals, millet, maize, and Job's-tears, the inference being that rice is more recent while the others are lost in the mists of antiquity. The existence of a distinct name for maize is everywhere indicative of a respect- able age, the more so when we note that in several cases where a variety has been introduced by the British or by Nepali immigrants, the fact is well-known. Among a few tribes, notably the Khasis and the Lhota Nagas, the tribal name is indicative of origin from neighboring peoples. This is not, however, quite conclusive, as the generalized name could be based on a variety got from the tribe in question and which supplanted older and more indigenous types. 1949] STONOR & ANDERSON—MAIZE OF ASSAM 375 Му notes on the religious aspect are particularly scanty. However, the dance of the Lakhers, the use of maize in funeral rites among the Lushais in deliberate preference to rice, its importance as a votive offering among the Monbas, the part it plays among the agricultural ritual of the Rengma Nagas, and the existence of a special tutelary deity among the Chang Nagas, all point to its being a well- established crop, the more so since primitive peoples with animistic religion are invariably shy of incorporating new crops into their agricultural ritual. The wide distribution, the positive statements of the peoples themselves, the position relative to the crop complex, the existence of distinctive tribal names, and the place in religion, all point to a long history, going back for centuries at least, for maize among the hill tribes of this region. Ав we shall consider immediately, this is fully supported by the general economics of the crop and the variety of uses to which it is put. GENERAL ECONOMICS This is a major study in itself, and the factors involved are many. Among them may be listed: (1) climate; (2) the varieties available; (3) techniques of cultivation; (4) other crops grown; (5) uses for which the crop is grown. Taking first the climatic factors, the climate of the Assam Hill areas varies from sub-tropical to sub-temperate, and maize is grown at all elevations from a few hundred feet to 6,000 feet. In general, it is of more importance to the tribes living at high altitudes, a state of affairs not unconnected with absence of rice varieties suitable for cold elevations, as among the Chang Nagas and surrounding tribes, and possibly among the eastern branch of the Lushais and the Chins. Varieties grown in any one area or by any one tribe are clearly dependent, with- in the limits of the climate, on the culture contacts, the purpose for which maize is needed, and so on. There is a far greater variety of types grown along the Assam-Burma border (Nagas, Lushais, and Chins) than in the outer Himalayas (Abors, Daflas, Monbas), although maize is on the whole more important in the Himalayan region. Since all tribes of the Himalayas store part of their maize, in- spection of granaries after the crops had been harvested have enabled me to survey with reasonable accuracy the varieties grown, and the peoples of this region seem to cultivate only the larger coarser types, the small several-eared forms being entirely absent and apparently confined to the Assam-Burma border. These very distinctive small varieties are moreover grown at low altitudes, and I do not think them to be cultivated at elevations higher than 4,000 feet, so that they are all associated with the sub-tropical rather than the sub-temperate zones. Although the climatic conditions of the inhabited parts of the Himalayas are, on the whole, colder than in the other hill areas with which we are concerned, there are thickly populated areas in the low sub-tropical foothills of the Abor country which are very similar to the outer Naga Hills; and the inference is that the tribes of the Himalayan region have never had these small maizes. This seems to link up [Vor. 36 376 ANNALS OF THE MISSOURI BOTANICAL GARDEN with the larger number of tribal groups in the Burma region and the greater fre- quency of migration to and from the area, as well as local movements within these particular hills, each group contributing its quota to the complex of crops over the passage of time and passing them on by local diffusion. Leaving aside any possi- bility of a variety having originated at some remote period in the area where it is now grown, each tribe seems to have accumulated its types of maize by culture contact. For reasons given I do not consider that there has been any great production of varieties by hybridization. In the data recorded above some support is given to this origin of varieties within a tribe. Thus the Lushais regard a proportion of their maize varieties as their own, one as obtained from their Chin neighbors, and two as of recent introduction from the Nepalese and the British. Тһе Lhota Nagas of the hotter foothills have three varieties they regard as indigenous, or at least of very ancient introduction, one obtained from the Angami Nagas, one from the Ao Nagas, and one from the Nepalese. It is stressed that the word "indigenous" im- plies only that a variety has been grown for a period for its ultimate origin to have been forgotten in the traditional memory of the tribe, and in no case necessarily means that it originated with them. Technique of cultivation:—As indicated already, in the majority of cases the maize is grown in lines, small patches, or single plants among the other cereal crops. Only among the Monbas with their advanced farming, the Angami Nagas and the Apa Tanis with their system of irrigated rice growing, and perhaps among the Lushais, the Chins, and Akas, is it normally a pure crop. Ас first sight, this rather haphazard technique seems conducive to hybridization, particularly where five or six types are grown by a single village. But examination of granaries shows but a small proportion of parti-colored ears. "The reasons are not far to seek. In the first place, there is а general prejudice which I have often heard expressed against saving seed from any plants which do not look pure bred; secondly, as I have demonstrated by experiment, different types flower at long enough intervals apart to act as a fairly effective check on cross-pollination; thirdly, all types are not extensive, but in every area I have visited, a walk through the fields shows different types grown in different stretches of land and certain types are definitely looked on as more suited to distinct soils and elevations . In clarification of this, it is important to remember that a village community cultivating on hill slopes will almost invariably have in use at any one time fields varying up to several hundred feet in height above sea-level, with corresponding diversity of exposure to winds, etc. Іс therefore stands to reason that where different varieties of any crop are grown they will be dispersed according to individual suitability to local variations in climate and soil conditions. Ав a general rule I have seen the larger-grained types of maize relegated to the higher-altitude fields, while the small popcorns and several-eared forms are sown on the lower, warmer slopes. Finally, the tribes are perfectly well aware that sowing different varieties mixed together leads to the 1949] STONOR & ANDERSON—MAIZE OF ASSAM 2/7 hybridization they are anxious to avoid, and for obvious reasons deliberate segre- gation is practiced. Uses to which maize is but: Within the limits of the climate and the varieties available, the purpose for which maize is grown is naturally the determining in- fluence of every aspect of a crop, maize as much as any other. As the whole back- ground of the economics, is the major fact that in the Assam Hills rice is the cereal crop preferred above all others, and wherever it can be grown and suitable types are available it is now the staple cereal. In qualification of this, we have already referred to maize as being the main grain crop among the Monba tribe of the Assam-Bhutan border. Members of the tribe have, however, told me that they prefer rice when they can get it, but since their arable land is limited and there is very little fit for irrigation, it is more economical to grow maize as a main crop. In a somewhat similar way, among the Chang Nagas and neighboring smaller Naga tribes, millets, maize, and Job's Tears are the most important cereal crops largely because the people have no rice varieties suitable for the cold altitude of their lands. Within the past two or three years the Chang tribe has made requests to the government to help obtain seeds of rice suitable for this cold climate. I do not know if this applies among the eastern Lushais and the Chins, but the in- ference is that it does, since the Lushais dwelling in warmer parts of the hills are all rice cultivators. The detailed uses of maize as outlined above are: 1. A catch-crop, eaten while the grain is soft. (АП tribes.) 2. Stored for food in the winter, either as a reserve secondary to rice, or more occasionally as a main crop. For beer-making. For popcorn. For pig food. As an article of trade outside the village. ON VA RS The first named is the most universal use of the crop, and applies to all tribes. For eating fresh, as a catch-crop, the smaller, several-eared types (other than the popcorn) are often preferred by those who grow them. This is apparently due to their soft, sugary grains which are appreciated both in themselves and as an alternative to the harder less sweet types. I think this to be the main reason for survival of these small types, since the grain is not nearly so economical for storing as winter-food as are the larger, "coarser" forms, and I know of no tribe which stores the smaller varieties for any purpose other than popcorn and perhaps for beer (vide infra) to a very limited extent. А point of some importance in con- nection with the use of maize as a catch-crop and for immediate consumption is the fact that different types grown in the area in question differ very appreciably in the time taken to mature, the small several-eared forms being several weeks slower than the larger types. It is of course well known to the peoples who grow them, and who naturally therefore space out the crop on this basis, to cover as long a period as is necessary. [Vor. 36 378 ANNALS OF THE MISSOURI BOTANICAL GARDEN The dried, ripened grain stored for the winter is, as we have seen, of primary importance among the Monbas, Changs and neighboring tribes, the Chins, and Eastern Lushais. It is of secondary importance among the Daflas and the Abor tribes, and the Sema Nagas, and is of fairly minor importance among the rest. It is, however, essential to maintain a balanced perspective and to remember that while statements of this kind are accurate in general terms, the whole balance of crops and their use is naturally fluid, especially among a primitive tribe with a comparatively unstable economy, and any general picture is only true for the actual time a survey is made. Thus, local conquests of tribe by tribe were taking place in the Naga Hills and elsewhere before British Administration was intro- duced, conquests which were accompanied by varying degrees of change in the pattern of culture, the crops not excluded. Nor has change ceased since more settled administration was started. In most hill areas of Assam, the government has constantly been trying to persuade tribal peoples to change over from shifting technique for mixed crops to permanent irrigated cultivation for rice. Similarly, in a more restricted sense still, a season of poor rice harvest in the autumn means that the supplies for the winter and following spring will soon be exhausted, and under these conditions a people such as the Daflas who normally consume a proportion of their maize as it ripens and store a proportion for beer and a reserve of food during the winter, will naturally use a greater amount for food and less for liquor, and vice-versa, after a good harvest, when there will probably be even a little to spare for the pigs. То narrow this down even further, from the tribe or the village to the individual household among peoples for whom rice is the most important of many crops, the subsidiary crops of maize, millets, and roots are naturally most important for the poorer people of the community whose rice supply runs out before the end of the season and the start of the next year's main harvest, and who live rather from "hand to mouth" depending to a greater or lesser extent on the "catch-crops" for their main food supply. The use of maize for beer is governed by the same factors as for food. Among most tribes rice is preferred for beer-making. Possible exceptions to this are the Sema Nagas, who seem to use maize from choice, and the Eastern Daflas, who grow considerable quantities of millet solely for beer. Те follows as a natural sequence that the worse the food crops in any one season, the less will be spared for beer and the more will be needed for human consumption. Very often beer is made from a blend of a variety of ingredients, the technique varying even from village to village within the same tribe, and maize will normally be included in the list of grains used for the brew. Thus, among the Chang Nagas I have been told that beer is made by mixing maize, Job's-tears, millets, and Chenopodium in fairly definite proportions. I have not been able to discover that any maize variety is used or grown specifically for beer, but this is a point calling for more detailed investigation. 1949] STONOR & ANDERSON—MAIZE ОЕ А55АМ 379 Рорсогп is used everywhere, and special varieties аге normally grown for this purpose only. The popcorn is made either by roasting the grain in a pot, often mixed with a little sand, or by simply placing the grains at the edge of the fire and picking out the popped grains with bamboo tongs as they burst. Тһе above notes deal with maize among the tribal peoples of the Assam Hills, and the plains areas of the Brahamaputra Valley and the southeast of Assam and Pakistan have not been considered since maize is not grown there by the indigenous population. It is, however, possible to grow it in the plains, and it is a common crop among Nepali immigrants. It is therefore perfectly feasible that it was grown in past epochs by the aboriginal population of the Assam Plains who might well have abandoned it with the development of a highly organized system of irrigated cultivation for rice. PART II EDGAR ANDERSON The data reported in this paper are basically simple, being essentially a morpho- logical survey of the varieties of maize grown by the Naga. However, the accurate assembling of these critical data required the cooperation of a number of individuals and institutions. The varieties collected Бу Stonor were numbered by him, and selections from most of them were grown in his experimental plot in Shillong, Assam. Herbarium specimens of the tassels and photographs of several of the more outstanding varieties were then forwarded to the Missouri Botanical Garden. Samples of the original ears collected by Stonor were sent to the Royal Botanic Garden at Kew where they were photographed. The ears were then shelled and the seeds, identified by their original numbers, were sent to the United States, where they were fumigated and forwarded to the Missouri Botanical Garden. The empty cobs were imported separately and were sterilized by heat before being re- leased. These extraordinary precautions were necessary because two of the worst diseases of maize are found in southeastern Asia. At the Missouri Botanical Garden samples of the seeds were germinated and the seedling characters were studied. Several representative cobs were turned over to Dr. L. W. Lenz for histological examination and were included in his recent (1948) survey. Early the following spring the seeds were forwarded to Dr. E. G. Anderson of the California Institute of Technology where they were planted in the maize-breeding plot at Arcadia, California, one of the most favorable sites in this country for tropical maize. With the cooperation of Dr. A. E. Longley of the U. S. Department of Agriculture and Dr. William L. Brown of the Pioneer Hi-Bred Corn Company, material for cyto- logical examination was obtained from many of the cultures and chromosome knob numbers and knob positions were determined from pachytene smears. During the growing season I worked at Arcadia for ten weeks where all the cultures, aside from a few late-maturing varieties, were scored for plant color, representative plants [Vor. 36 380 ANNALS OF THE MISSOURI BOTANICAL GARDEN were photographed to scale, herbarium specimens were made of tassels, internode diagrams were made of mature plants, and the details of tassel and ear morphology were recorded. Eventually all the data and materials were assembled at the Mis- souri Botanical Garden—the herbarium specimens and photographs from Shillong, the photographs and shelled cobs from Kew, the photographs, herbarium specimens, and notes from California, the knob counts made by Dr. Brown and myself, and the histological information from Dr. Lenz. We are also indebted to Dr. Herschel Roman and Mr. Earl Patterson for internode measurements and specimens of varieties which matured after I left California. The following collections were grown; the information concerning each variety was supplied by Mr. Stonor: SERIAL LOCALITY TRIBE REMARKS No 1 Е. Райа E. рай Sown Feb.-March; ripe June. Much grown at 3000 ft. Outer Himalaya — Local name Topothe. Number of ears 2-4. 5 N. Cachar Hills | Kuki Grown at 2000—3000 ft. Number of ears 4—5. 6 N. Cachar Hills | Kuki As for 5, with which it is grown. 7 бас Hill | Zani Mowe Grown ас 1500-4000 ft. 9 precise details. Said to give 2-3 ears. Sown about Ма 8 N Chu Hills | Zonk Naga I 2000—3000 ft. Sarwa to be slow-ripening. Gives Sown about M 9 Naga Hills Angami Naga | Grown at 3000 ft. Details as for 8. 10 Naga Hills Angami Naga | Grown at 3000—4000 ft. No other details. 11 Naga Hills Angami Naga | Grown at 3000—4500 ft. No other details. . Grown at 3000—4000 ft. Sown Feb. to March; ripe 12 Naga Hills Ao Naga June-Sept. Said to bear 7-8 ears. Local name Lozar Grown at н 000 ft. Sown late March; гіре Јипе- 13 Мара Hills Ао Мара July. Has I Said to be very sweet. Local name Mabok Mem : Grown at 3000 ft. Sown — to 9 ripe June to 14 Naga Hills Sema Nega fuu Hei eu Lol уым . Grown at 2000-3000 ft. Sown кегин и June- 15 Naga Hills Sema Nega July. Said to have 6 ears. Local name Ате : Grown at 2000—3000 ft. Sown pon ges ves ios Iur 1% Naga Hills Sema Nega Said to be sweet. Has 2—3 ears. Local name Mes | I at 3000 ft. Sown oer шы, ripe June-July. d Naga Hills Sema Nega s 2—3 ears. Local name Kolam 18 Naga Hills Sex Mii Grown at 2000-3000 ft. бозуп Feb.-May; ripe May to Sept. Has 6—7 ears iiio d. Local name Anila. 1949] STONOR & ANDERSON—MAIZE OF А55АМ 381 Іп addition, seven varieties collected іп the Lushai Hills were grown. Since they were in general very similar to the above and included no peculiarities not rep- resented in the Naga collection, no detailed account of them has been prepared. General Appearance: — As compared with most collections of native varieties from South and Central America, one of the most outstanding characteristics of these Assamese varieties was the uniformity of several of the varieties and their differences from each other. This bears out Mr. Stonor's remarks as to the skill and determination of the Naga in keeping their varieties pure. Several of the cultures, though grown directly from seeds collected among the Naga, were as uniform as a good inbred line. It is probably significant that those varieties which were most unlike anything previously known in our studies of exotic maize were the most uniform, while the one variety most closely resembling the maize of Latin America was among the most variable (see below under "Caribbean"). The conspicuous differences between certain of the varieties make it difficult to generalize about them as a group; nevertheless there were certain definite trends which characterized the entire collection. Almost without exception these trends were most strongly marked in those kinds which morphologicaly were the most extreme such as "Late Upright" and "Late Side- wise." F Internode diagram of one plant of Stonor 18 ("Late ey" mc grow at Arcadia, Cal. Circles represent tassels, elliptical figures ears. The diagram is as if the stalk were cut at the nodes and the dissected edap кој were laid side by side in succession, the lowest at the left, с ppermost at the right, and then a line (the were over nodes on the P ant, none Ë: which was over 8 cm. long; that there 4 ears; and only 4 short internodes betw the upper ear and the tassel. Small lines at the left of the diagram абер Ав ас which id were well-developed prop- roots. Figure 1 shows a drawing to scale of a typical seedling. The short, narrow tap- root, the numerous adventitious roots from the mesocotyl, and the broad mesocotyl and coleoptile are characteristic. One of the most unexpected characteristics of these varieties was their green color. Not only were they mostly without the bright plant, tassel, and silk colors of so many Latin American varieties, but some of them had no visible anthocyanin pigment in any part of the plant. The group as a whole had a strong tendency to green silks, green anthers, green leaves, and green culms. There was also a strong tendency for the leaves to have a more evenly green appearance, like certain varie- [Vor. 36 382 ANNALS OF THE MISSOURI BOTANICAL GARDEN ties of Sorghum. In nearly all the maize of the New World there are minute dif- ferences in the intensity of the green above and between the veins, giving the leaves macroscopically a kind of longitudinal grain. The Assamese varieties tended to be evenly green throughout like a green plastic dish. The collections among the Naga were outstanding in their lack of vigor, and this was quite as true in Shillong, Assam, as in Arcadia, California. Some were mid-season, others late or very late, but all of them developed slowly. From the time the tassel made its first appearance until it was completely out of the leaves and shedding pollen, as much as four weeks might elapse. One of the most out- standing characteristics of the maize of the New World is its vigor. Nearly all maize varieties grow and develop rapidly; the controlled heterosis of hybrid maize is merely an extreme standardized example of a tendency nearly universal in the New World. This vigor was absent in the Assamese varieties. Whether midseason or late in their maturity they poked along from week to week. The internodes were short; in the most extreme variety none of them was over 10 cm. in most plants. Nor was this the result of their having been transplanted from Assam to California, since the samples grown by Stonor in Shillong exhibited these same characteristics. The internode patterns were highly peculiar. In addition to short internodes, the varieties from the Naga tended to have many ears (as many as four or five in some varieties), and the internodes above the ears were so short and crowded that the silks of the uppermost ear were sometimes tangled in among the lower branches of the tassel (see plate 22). Not only were the upper leaves crowded together but as Collins noted (1909) in his description of waxy maize from the Orient, they all tended to be gathered at one side of the culm and to hang over the developing tassel like a kind of spathe. While this character was more extreme in some varieties than in others, there was a marked tendency in that direction throughout the collection. The arched and drooping spathe-like upper leaves were accented by the drooping and semi-included tassels. Tassel branches tended to be more slender than those of New World varieties, and in the more extreme varieties they hung down verti- cally until after the pollen was shed or even later. Though the tassel branches were long, the glumes were small. Because of the short upper internodes the tassels were never exserted from the upper leaves as pollen started to shed and many of them were not exserted even when fully mature. For the ear the prevailing tendencies of the collection were to small cobs, promi- nent glumes, small, isodiametric kernels, and complete absence of row-pairing. The colors of their kernels were mostly a pale straw-yellow or a dull reddish-blue. In their even green color, included tassels, slender culms, slender tassel branches, isodiametric kernels, straw-colored or dull blue kernels, these Assamese varieties resembled sorghum more closely than do New World varieties of maize, and one 1949] STONOR & ANDERSON—MAIZE OF А55АМ 382 variety even had a bluish-green bloom оп its leaves rather similar to that which is so characteristic of certain varieties of sorghum. Among the collections from the Naga a number of different sub-types could be distinguished. It seems unwise to dignify them with permanent names until we know more about the kinds of Oriental maize than we do at present. For the pur- poses of this discussion they may be provisionally designated as follows: I—Carib- bean; II—Early Slender; III—Late Upright; IV—Early Upright; V—Late Side- wise; VI—Drooping Waxy. Some of their characteristics are shown in tabular form in table 1. I—Caribbean: 'lypes of maize fairly similar to those grown around the Carib- bean basin are widely distributed in the Orient and were apparently introduced by the Spanish. They seem to be the prevailing type in the Philippines and in Guam and they or mixtures with them make up the bulk of the maize grown in Asia, particularly at lower elevations and along the coasts. In previous years I have grown collections from India, Guam, the Philippines, Sumatra, the valley of the Irrawady in Burma, and from China. Опе such variety was collected by Stonor (No. 17), though he commented as follows: “Evidently not quite a pure strain." The ear- to-row test of this collection showed more plant-to-plant variation than did Stonor's other collections among the Naga. In it the distinctive characters of the rest of the collection are less strongly developed. It grew more quickly, had more pronounced plant color and colored silks, it had coarse tassels, large ears with white kernels conspicuously capped with soft starch, and was the earliest of anything in the collection. It showed Assamese tendencies іп its somewhat included tassel. It probably represents a fairly recent mixture between Assamese maize and one of those Caribbean types which were spread so widely around the world by the Spanish and the Portugese. II—Early Slender: In its slender stem, long slender leaves, and most pronounced spathe at flowering time, this variety was very similar to VI. It differed in being the earliest of the distinctive varieties and in the morphology of the mature tassel. Although the tassel branches were completely pendent when they first appeared and even after they began to shed pollen they eventually stiffened to produce a tassel more like broom-corn than any other known variety of maize. Тһе long, slender, wiry branches became stiffly distended when mature, and the small glumes (5—6 mm. long) were quite closely appressed. Аз in many of the varieties with short upper internodes, the auricle of the uppermost leaf was developed into a conspicu- ous tuft of long white hairs. Though to casual observation this variety seemed to be completely green, careful examination at the base of the plant showed a faint flush of color. Ш-— Та Upright: The leaves of this type were dark green, were held crisply erect until after the tassel appeared, and were twisted, usually one complete revolu- tion and sometimes more. The plants had many short internodes and grew and [Vor. 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN 384 z Á1OA] иоѕеәѕ-рту{ 3001225 300125 +] Z 0< 222495 pews 91 | 2801 | 91 Áxt A, Surdoo1(T—IA ғ | моќ зт $ әзе] Á12A 390136 =- ғ 61 212445 yews FI | 6801 81 + | aop ЈО z әзе] ÁNA 300135 + + 81 212445 pews 21 €80I 21 әзтмәріс әзет—Д £ моүәќ 31 9—r әзер ÁNA Buong — £ £c 212445 yews 91 2801 II + рәм Арея штрәуү 390135 + £ 81 212445 pews Аләд 01 4401 9 2484 K[1*4— ATI ajdind-pay aI -- 71 81 әләц4в4п< adie] #1 <801 #1 enm mad | zs aT iure 0| Z | 61 222495 Wes | 91 |1801 | от 2119 ца r әзер Á19A 8 zz 212445 yeas 02 | 0801 | 6 эч3їл&г\ү ə1e1—TIII с usi|danq әзе] Á19A 300135 зше 0| mj zz 212495 Пеш oz | 6401 | 8 - 2119 ца Арея 340135 300125 0 4 oz 212495 pews +I 9/01 ç I моПәА 31 Арея штрәу 390136 — 9 02 ərenbç шпірәу{ УТ | $801 çI зәрчә[$ Á[1t4—II I WYA Арея 3481$ зи + 11 92 зе э81е] #1 8801 41 utoqque5—T[ 22 $E | 28 f F TEE AERHE: ЕЯ Бк | 22) го |22 ЧЕ ЕЕ = E 28. = Sá | sm) 22 $32 83 54| 55 |75 а 2, 33 E E 2.5 Нә) 89. e Bo я к 3 adÁ[ "a со = 9 t РА = Z 8 РА 8 B s K ° 58 ШЕ à ao = SHILAIYVA VOVN AO SOLLSDISIOVUIVHO I TIgV.L 1949] STONOR & ANDERSON—MAIZE OF ASSAM 385 developed slowly. The internode pattern of a typical plant is shown in fig. 1. The kernels are small, with a dull, purplish blue aleurone and are used as a popcorn (see Stonor’s notes). Similar popcorns (as well as the actual popped kernels) have been received from correspondents in Siam. Stonor’s different collections made among various Naga tribes were almost identical in growth type aside from one or two obviously out-crossed individuals. In their small kernels, upright twisted leaves, large number of ears, and pedicel and cob anatomy, this type resembles certain ancient popcorns of South America. Ears and popped kernels have been obtained from various graves and trash-heaps in coastal Peru and Chile (Anderson, 1947), and through the kindness of Dr. Paul Weatherwax a living example of such a pop- corn was grown from one of Parodi’s collections in Argentina. Since Ica times (the Ica preceded the more widely known Inca) these kinds of small-kernelled popcorns have been rare. At earlier times they were apparently the prevailing type along the coast of Peru and Chile and have been recovered from graves in the Argentine. A somewhat similar popcorn, of unknown origin, is sometimes grown in the United States in spite of its long season and small ears, because of its high quality. Though it has been extensively confused with the early-seasoned “Тот Thumb,” its proper name is apparently “Ladyfinger,” by which name it was first called in the United States at least a century ago (Emmons, 1849). The tassels of Late Upright plants exhibited little or no condensation and from slight to pronounced multiplication (Cutler, 1946). The central spikes were pre- vailingly in whorls of three. Most of the plants grew four good ears with partly developed nubbins at lower nodes. Collection Nos. 5, 8, and 10 showed no base color (in other words they were like ‘aa’ plants) and No. 14 which showed base color had larger and redder kernels and may well have been derived from a cross between a typical Late Upright and some other sort of maize. One cob of No. 8 was examined by Lenz (1948) and is figured by him, plate 38, fig. 5. It has the longest, slenderest pedicel ever reported for any variety of maize. IV—Early Upright: This one collection (No. 6) was in every way like the Late Uprights except that it was earlier, had red seeds, and pronounced base color. V—Late Sidewise: Both in Assam and in the United States this variety in the vegetative stage looks unlike anything previously reported for Zea Mays. The leaves and especially the culms are bluish-green with a distinct bloom. The tillers are practically horizontal during the first month or so of their development. On both tillers and main culm the internodes are very short, and even in the mature plants few ever exceed 10 cm. in length. Most agronomists, seeing the plants be- fore they had tasseled out, would have wondered if they were some kind of sorghum. Base color varied from very faint to strong, and the leaves, even early in development, stood out at right angles to the culm. The ears were long and slender with small yellowish kernels. The large number of internodes (30 or more [Vor. 36 386 ANNALS OF THE MISSOURI BOTANICAL GARDEN per plant) was evidently not a response to a longer day length, since similar results were obtained in Shillong and since there were prop-roots at only one or two of the lower nodes. (In tropical maize grown in the United States there may be prop- roots on 10 or more nodes as one result of the change in length of day.) VI—Drooping Waxy: Up to flowering time this variety was much like II ex- cept for the later season and consequent higher number of leaves. It had the same slender culm, slender green leaves, and, if possible, an even more highly developed spathe from within which the slender tassel branches drooped straight downward. Even at the end of the season, both in California and Assam, the tassel, though arching horizontally, was still quite pendent and much more extreme than any tendency of this sort in South American maize. All the plants of this variety had waxy pollen, and an analysis of the kernels at the Northern Regional Laboratory at Peoria, Ill., bore out this diagnosis. Stonor reported the young ears as being out- standingly sweet, and, like much of the waxy maize in the Orient, this is probably a specialized type used for green corn. Waxy maize is cultivated in the vicinity of Chungking, China, as a table corn, and Kuleshov's monograph (1928) illustrates the same pendent tassel in waxy varieties from various parts of Asia. To summarize: 'These remote Asiatic aborigines cultivate a number of excep- tionally well-differentiated varieties of maize. The following unusual characters typify one or more of these varieties: Uniformly green leaves, culms, silks, and anthers. Slender, pendent tassel branches. Straw-yellow endosperm, dull bluish-red aleurone. Small, isodiametrical kernels. Many short internodes, lack of vegetative vigor. Upright, twisted tassel branches, short silks. Tassel included in upper leaves at flowering time; leaves immediately below it falsely monostichous, forming a sort of spathe; tassel not completely ex- serted even when mature. Waxy pollen and endosperm. As we shall demonstrate below, this complex of characteristics is widely dis- tributed in the back corners of Asia. It would be of primary significance to know where it is most closely approached in the New World. Certainly nothing like it is known from Mexico, Guatemala, or other parts of Central America. The only United States variety showing any of these characteristics is "Ladyfinger" popcorn, a variety of unknown origin which has been in this country for at least a century (Emmons, 1849). In South America this complex of characters is rare, and most of the collections and published descriptions demonstrate radically different types of maize through- out that continent. However, a popcorn collected at Chiu Chiu, a remote oasis in 1949] STONOR & ANDERSON—MAIZE OF ASSAM 387 the Atacama desert of northern Chile, Бу Carl and Jonathan Sauer, has а number of features in common with these Oriental varieties ( Anderson, 1943). Further- more, as mentioned above, an indigenous popcorn collected in Argentina by Parodi, shows many of these same characters. А photograph by Jonathan Sauer of still another variety from Chiu Chiu, Chile, shows the multiple ears, the short silks, and the short upper internodes of the oriental varieties. In a collection of native corns from the Bolivian edge of the eastern lowlands, kindly turned over to me by Dr. H. C. Cutler, there were several plants which had the green plant color, pendent tassels, multiple ears, and spathe-like upper leaves of the Oriental varie- ties. Dr. F. C. Brieger informs me that he noted some of these same characteristics in other collections from these valleys. Insofar as one can judge from characters of the mature ear, a fairly similar set of varieties was once common on the west coast of South America. АП the collections of early prehistoric maize from that area (and the museum material is so rich that it has not yet been possible to measure and record all of them) shows a small-cobbed variety with isodiametrical kernels, much of it apparently a popcorn, since the prehistorically popped kernels are known from a number of sites. This type, uniform in the lower archaeological levels, became gradually more variable and was supplanted by larger-kernelled types sim- ilar to modern Andean maize, when the Icas extended their influence down to the coast in times preceding the Inca domination. In the Orient, on the other hand, similar varieties are widely, though very spottily, distributed. They are associated with the most primitive cultures in south- eastern Asia, principally with the Tibeto-Burmans and allied peoples. In 1909 G. N. Collins published an exhaustive account of a variety of maize with a waxy endosperm which had been collected in China. In 1920 the same author, in a short communication, reported that waxy maize and other curious varieties had been ob- tained from the hill tribes of Upper Burma. Collins’ description of waxy maize would apply equally well to most of the Naga varieties: While in only about 25 per cent of the plants were the upper leaf blades 2 PINE № of them showed a tendency in w: сиы. This one-ranke vo ance is brought abou d a twisting of the leaf chee ths, actual insertion of th ves мени орровіге аз іп all Ки es. Іп addition to the un сз phil of the leaves the tea of d upper nodes w erect ni и е or dropping as іп other varieties e internodes on the upper part of the plant were also much shortened, so that the а was not carried up, as in ae: varieties ie и 8). His description (loc. cit., p. 13) of the aleurone color is also applicable to many of the Naga collections: The color of the Bp laye r was distinct Мес anything that has been observed іп other varieti es. г varied greatly іп intensity. Іп rare cases it approached the bluish black of our common ack” ы but for es most p varied from a dull ruby to maroon. The color was bes confined to the top of the seed, fading out toward the base Under Vavilov the Russians made a comprehensive survey of Oriental maize which was reported upon by Kuleshov in 1928. It is extensively illustrated and accompanied by a summary in English (pp. 371—374). It demonstrates that [Vor. 36 388 ANNALS OF THE MISSOURI BOTANICAL GARDEN varieties similar to those described from the Naga country are widespread in cen- tral Asia from Persia and Turkestan to Tibet and Siberia. Figure 6, a variety from Persia, shows the characteristic short silks described by Collins and fig. 11 (3—6) shows tassels covered by a spathe of leaves in Persian and Turkestan varieties. Ex- tremely pendent tassels from a number of points are described and illustrated. On pages 373-374 Kuleshov summarizes his morphological evidence. Asiatic maize shows a series of characters which are ре unknown ог very rare іп тегісап! maize. These characters manifest themselves in connection with a definite area, sometimes a very large one. Be the waxy endosperm is Sinn: from 5 to 45? North pha the short plant habit is peculiar to the vast expanse of Central Asia; panicles covered by t upper leaves and silks "c in c» axil of the leaf are met with in specimens from eastern Asia to Transcaucasia, etc The question of the introduction of maize into the Orient was gone into ex- haustively by B. Laufer (1909). Не came unequivocally to the conclusion that it was not introduced into China from the coast like other American crops but spread overland via Tibet. He was an accomplished linguist and bibliographer but with- out botanical training and had had to take the word of the botanists of his day that maize in the Orient was morphologically not different from maize in the New World. With this as а premise he could have come only to the conclusion he finally reached: that maize somehow got to Indian ports at an early post-Columbian date and spread overland via various primitive peoples to China. Не quotes no botanical authorities for the morphological equivalance of Asiatic and New World maize, merely stating (footnote p. 224) his premise "If maize were indigenous to Asia, we should expect to find there either a wild form, from which the cultivated species are derived, or the Asiatic species to be differentiated from that of America, neither of which is found." Had the botanists of his day studied Oriental maize in even a cursory fashion they would have found that Asiatic maize is indeed ‘“‘dif- ferentiated from that of America," most particularly in that very area between China and India which Laufer decided must have been the route by which the crop eventually reached China. His monograph is a mine of information as to the frequency of maize among these various primitive peoples and the role it plays in their economy: In the remarkable culture of the small mountainous tribe of the Lepcha in Sikkim, a people closely related in language to the Tibetans, maize plays a veri role, and it is о surprising to note йөк а rich terminology they have developed with regard to its economy. There are four words for maize, two fo or the flowers of maize; ој o less. than gene eans mes acc е to its grow a speci tal Ж for a young head when pes for d when ri oc. cit., p. 242] . . 50- дена aboriginal tribes of western and southern ina, favorite and principal food and is more highly appreciated by these tribes i Tey ЊЕ үйө, ос, cit., p. 244]. ‘It should be emphasized that Kuleshov (193 0) produced a ане оп the фини ый maize made Бу Vavilov and his collaborators іп Central and South America and was probably familiar with the morphology of New World maize, as a whole, as any einn Раб 1949] STONOR & ANDERSON—MAIZE ОЕ ASSAM 389 Laufer quotes (loc. cit., p. 245) В. C. Henry who says that in China maize "is now very extensively cultivated by the Chinese, but especially by the aboriginal peoples, among whom it seems to be almost as great a favorite as among the Ameri- can Indians. It forms a main portion of the sustenance of both the aborigines in the north-western corner of Kwantung and of those in Hainan." Through the courtesy of Dr. Ian Khambanonda it was possible to import a sample of a popcorn from Siam. It has violet-blue kernels similar to those of the Assamese varieties received from Stonor. Dr. T. W. Whitaker kindly grew a few plants of it for me at the U. S. Dept. Agr. Vegetable Breeding Laboratory at La Jolla, California. In its long season, upright leaves, slender tassel branches, short internodes, and large number of ears it closely resembled the Assamese popcorns. Popped kernels were also forwarded from Siam but Dr. Khambanonda was unable to supply very much detailed information because maize in Siam is grown prin- cipally by the aboriginal hill-tribes, seldom by the Siamese. To summarize: А number of distinctive maize varieties are grown in the re- motest parts of southeastern Asia, particularly by the aborigines of various hill tribes. Without exception it is more common among these primitive folk than among their civilized neighbors. In China it is more common and more appreciated among the aboriginal tribes of western and southern China than by the Chinese (Laufer, loc. cit.). Dr. Khambanonda testifies to its almost exclusive cultivation in Siam by the aborigines. Interviews with Ko Ko Lay, now an exchange scholar from Burma, have produced similar testimony for Burma. Dr. Pierre Larroque, the former maize breeder for Indo-China, tells me that among the primitive Meo, ethnologically related to the Tibetans, it is very commonly grown and that on the Yunnan border it seems almost to run wild. Mr. Stonor, the author of the first section of this paper, having been transferred to New Guinea, informs me that although it is either unknown or very recent along the coast, it is found among the primitive peoples of the interior. Its importance among the Lepcha of Sikkim and the aboriginal Li of the Island of Hainan has already been alluded to. That maize could in post-Columbian times have spread to each of these various hinterlands without entering into the economies of the more civilized people who would have handed it on almost passes belief. Най it spread only to the Мара, опе would have wondered what special circumstances caused its adoption by a people so remote that one ethnographer describes them as having lived for thousands of years “іп these hills, as on some happy island, almost untouched by the waves of civiliza- tion which from time to time have surged through the plains of Assam and the valleys of Upper Burma. Ancient cultures which were once spread over great parts of southeastern Asia and which in most countries had finally to give way be- fore the higher Indian and Chinese civilizations have been preserved here in a com- paratively untouched form, and allow us to observe with our own eyes, early types [Vor. 36 390 ANNALS OF THE MISSOURI BOTANICAL GARDEN of human culture” (von Fürer-Haimendorf, 1939). Furthermore, when we ex- amine the maize of these people it is not the dominant world crop of Central and North America. It is relatively unproductive and with less vigor than other known types of maize. То believe that in post-Columbian times maize could have репе- trated not only to the Naga but to the hill tribes of Upper Burma, and of Siam, to the Lolo in central Asia, to the aborigines of Hainan, to the hill peoples of Sik- kim, and to the interior of New Guinea, in each case passing over the more civilized peoples along the coast is beyond credulity. То have these conservative people somehow learning to use maize as a popcorn and as a green corn and as a cereal for brewing, to have them growing types of maize which are similar to each other yet rare or unknown in the New World puts the burden of proof on any one who would ascribe all this development to separate post-Columbian acquisitions. It seems more likely that there have been at least two major movements of maize in Asia. Тһе latter in early post-Columbian times brought what is essentially a Caribbean type of maize to the Philippines and to many countries actively colonized by the Europeans. Back in the hills, however, are much more primitive types, unaggressive, not particularly productive, grown by conservative people. If one asks why they did not spread more the answer is that they did spread in Asia from Persia to Sumatra and New Guinea, which is virtually as far as the Asiatic Sorgbum (which was their companion crop) has been carried. The general evidence for the main kinds of maize in South America and in southeast Asia is crudely and diagramatically summarized іп fig. 2. Тһе letters A, B, and C represent three of the major races which make up Zea Mays. Letters within parentheses indicate that a particular type is present but is relatively un- common. Each of these major races has a core of morphological characters in common in spite of great variation from variety to variety within the race (Ander- son and Cutler, 1942). A represents the small-kernelled types we have just been describing and mixtures with them. In South America they are found today either as rare native popcorns or as one of various primitive tendencies in the highly heterozygous maize of the eastern river basins (Cutler, 1946). Іп Asia they are practically confined to hill areas and in the most isolated are the only type. In western South America, where we have stratigraphic archaeological evidence, we know that гасе A was for centuries the only type of maize. В, in fig. 2, represents the distinctive large-seeded frequently large-cobbed types of the Andean region, which are practically exclusive at higher altitudes, which have dominated (at least in mixtures) the west coast ever since prehistoric times (late Ica) and which are one of the elements in the maize of the Amazon basin. Such types are apparently unknown in Asia, a most significant fact. C stands for the widely adaptable types of the Caribbean basin which spread so rapidly and extensively around the world in post-Columbian times. 1949] STONOR & ANDERSON— MAIZE OF ASSAM 391 This diagram is factual and though some of these facts are matters of judg- ment; with other types of maize it has already proved possible to make objective records of such racial differences (Anderson, 1944; Brown and Anderson, 1947; and Anderson, 1947), thus taking such questions out of the realm of acrimonious quibbling. The interpretation of these facts is quite another matter. The facts are in themselves fantastic; any satisfying hypothesis must border on the miracu- lous. One fact seems to be clear. Race C, which we know to be archaeologically post-Columbian in South America, was widely spread in post-Columbian times. If we therefore remove C from consideration, the problem to be solved is how could race А get to a number of isolated hill areas in Asia without anywhere leaving a very definite record along the coast of Asia? ТЕ, for the sake of argument, we grant that it might somehow have spread there from the Upper Amazon in post- Columbian time, who brought it and how? Was there somewhere in the New World a reservoir of various A varieties which has since disappeared? Could all the A varieties in Asia have differentiated themselves in all these backward areas into a set of unique but similar varieties under the stimulus of a new environment? These are possibilities but they certainly seem fantastic. If we admit (with a growing minority of archaeologists) the possibility of trans-Pacific contact in very early pre-Columbian times, then race A might have crossed the Pacific at an early date when maize was stil an unaggressive little popcorn, to be carried across the Pacific again when the dominant world crop which we now know had been developed in the New World. Аз to which way maize made its first crossing, whether from Asia to the New World, or vice versa, the facts of fig. 2 do not even suggest an hypothesis. From the generally accepted facts as to the relationships of maize a good case could be made out for either Asia (oe Late Ica ТА: Е . Distribution of the major races of maize in South America and in Asia. Very активниот А, B, and С oe three of ho main ње of paite, each Mi ons of various nud nd innumerable Шайды Though these race pa ре” ach has own ore related tendencies: А, sca eological чај More ears, large gr а. seeds, Mir id drooping and in y x^ cach goes plant color; p Кои Andean—large Ке m cag cig шісі um, large cobs, strong plant color; Race C, t coid—long ears, stiff tassel, bon etters in parent rid indicate areas k лина а Aude ike race is i. only in 15 P ne races, In the Orient race C is represented in "Caribbean Flints" a mixture he C and other races. Asia is _diagrammed to the left of the figure, South America to the right, with the Pacific in bet For western South America there is extensive archaeologica evi iden s. some of it а” which allows us to establish a sequence of maize types fen At А [Vor. 36 392 ANNALS OF THE MISSOURI BOTANICAL GARDEN or the New World as a primary center. There can be no doubt that the New World was certainly the secondary center, or rather a whole set of secondary centers. The physical possibilities of transfer by the early Polynesians are much more likely than most botanists are aware. These early people were skilled navigators who could deliberately set out for Hawaii from Samoa and arrive at Hawaii. Fur- thermore, they were skilled cultivators who carried vegetatively propagated varieties of various crops to many of the islands of the Pacific (Buck, 1938). Getting the improved taro and bread-fruit to Hawaii (which they are known to have done) would have called for more skill as navigators and cultivators than to have taken maize from New Guinea to Peru. Nor would maize have been exceptional among cultivated plants had it been taken across the Pacific. It is known that the gourd crossed the Pacific in very early pre-Columbian times and it is admitted by most authorities that the cultivated sweet-potato originated in the New World but spread in pre-Columbian times to Polynesia and even as far as New Zealand. The sword bean (Canavalia), widely cultivated throughout the Pacific and always considered to be of Old World origin, is now known from prehistoric sites along the coasts of both South America and Mexico. The hypothesis here suggested as to the origin and developments of maize is paralleled almost exactly by the conclusions concerning cotton reached after long cooperative research by the British cotton experts (Hutchinson, Silow & Stephens, 1947). In the case of cotton, polyploidy operated to keep certain mixtures of old and New World germ-plasm from losing their identity in the mixture, thus making the story quicker to untangle and more difficult to controvert. For many minor domesticated plants, the rather complete neglect of cultivated plants and weeds by taxonomists leaves any general discussion of the problem at the stage of mere guesswork. Neither the cotton story nor that of maize could be put together until a beginning had been made at the fiendishly difficult problem of classifying and cataloguing their various cultivated varieties. Both Amaranths and Chenopodiums are cultivated by the hill peoples of Asia and of the New World as cereal crops, pot-herbs, ornamentals, magic plants, and food colors. Not until these groups have been meticulously collected and monographed (the cultivated strains, the weeds, and the genuinely wild entities) can we be in a position to dis- cuss the evidence. Similar careful studies are needed of the various strains of Bixa Orellana, of Job's tears (Coix), of Pachyrhizus, all of which are widely distributed in both the Old World and the New. There is little really critical taxonomic evidence on origin and diffusion for any of these groups at present. It is disap- pointing to find, half-way through the twentieth century, that our botanical evi- dence is not yet at a stage where such fundamental questions can be authoritatively discussed. For the plants most directly associated with man, the cultivated plants 1949] STONOR & ANDERSON—MAIZE ОЕ ASSAM 393 and weeds, aside from a few collections by such pioneers as Ames, L. H. Bailey, and Merrill, we do not have the specimens let alone the critical studies. The average botanical collector is so intrigued by cloud forests and river jungles that he does not even think about the more difficult problems posed by the vegetation of dump- heaps, clearings, and cultivated fields. For maize itself, two facts suggest how complicated a story may be involved: (1). The relationship of Sorghum, Assamese maize, and prehistoric North Ameri- can maize. Whatever the explanation, it is clear that the maize of Assam is more like Sorghum in a number of different ways than is any other modern maize as yet examined in detail. Whatever the explanation, it is also clear that the earliest pre- historic maize cob described from Bat Cave, New Mexico, by Mangelsdorf and Smith (1949) is even more Sorgbum-like in the details of its inflorescence, while cobs from the upper layers of the same cave are as radically un-Sorghum-like as it is possible for maize inflorescences to Ье. It may be that maize and Sorgbum have had a parallel evolution in the Orient under the stimulus of a similar set of environ- ments. It may be that they are related in some way; it is known that they have the same chromosome number. Whatever the explanation, the whole story must be a complicated one. (2). Much of the maize of Central and North America has knobs on its chromosomes, a character which Mangelsdorf and Reeves ascribed (1939) to introgression from Tripsacum, a New World grass. Much of the maize of South America is knobless. The maize of Assam has only a few knobs but they are frequently quite large and they tend to occur at positions and in combinations which are either rare or unknown in New World maize. Here again a complicated history of exchange and evolution is suggested. Only one thing is certain. We must have extensive and critical collections of Oriental maize if we are to understand Zea Mays and utilize it most effectively. We need this information for practical and theoretical purposes. Kuleshov (loc. cit.) describes and illustrates dwarf, drought-resistant types from Persia. Larroque tells (personal communication) of a small-grained sort raised in Indo-China for chicken feed which germinates effectively in the soaking wet soil of rice paddies. More important and far-reaching is the need for such collections if we are to under- stand the history of maize. Along with Drosopbila and Neuros pora, the science of Genetics is built on work with Zea Mays. The brilliant work of Stadler and of McClintock, for instance, might have quite different implications for the funda- mental nature of evolution or for the protein chemistry of the germ-plasm, depend- ing upon the actual history of the germ-plasm which they have been using, the germ-plasm of Zea Mays. Certainly we cannot even discuss the probable history of Zea Mays in an intelligent fashion until we have at least an approximate notion of what varieties of maize are being grown by the Lolo in central Asia, by the hill tribes of Burma, Siam, and Indo-China, by the aboriginal remnant in Hainan and Formosa, and in the isolated interior of New Guinea. š [Vor. 36 394 ANNALS OF THE MISSOURI BOTANICAL GARDEN GENERAL SUMMARY | Part 1:— 1. The various hill tribes of Assam and neighboring regions are enumerated and their relationships to each other and to outside peoples are briefly described. Іп general, these tribal peoples grow their own food, weave their own clothing, and regulate their own affairs by tribal law. "They have no writing and their religion is animistic. 2. The varieties and uses of maize are enumerated tribe by tribe. It is used for human food, particularly when immature (i.e., as "green corn"), for beer-making, as popcorn, and for pig-feeding. 118 importance varies from tribe to tribe but, along with the cereal forms of Job’s-tears (Coix) and millet (Sorghum), it is traditionally one of the ancient foods of the region. 3. It is usually grown in lines or patches or as single plants among other cereal crops. In spite of this practice, the varieties are kept remarkably pure, even when five or six distinct types are grown by a single village. 4. The wide distribution in the area, the traditions and positive statements of the peoples themselves, the existence of tribal names, and the place of maize in their religions all point to a long history in this region, going back for centuries at least and most probably antedating rice culture there. Part П:— 1. The assembling of all the evidence on the morphology of these Assamese varieties was a complicated affair, because their growing season is long and because the importation of maize from southeastern Asia is necessarily restricted!. They were collected directly from the Naga and grown in duplicate in Assam and California. A set of the original ears was photographed on the ear in London, and plants from ear-to-row tests were photographed in California and in Assam. Pachytene smears were made of many of the cultures, and a few of the original cobs were studied in celloidin section by Lenz. 2. Тһеге are considerable differences between the Assamese varieties (most of which are remarkably uniform within the variety). As a whole they tend to have: slight or no plant color, many short internodes, semi-included tassels, spathe- like upper leaves, pendent tassel branches, several small ears, long slender leaves, short silks, dull aleurone colors, small subspherical kernels. They are outstanding, both in Assam and in this country, for their lack of vigor. 3. This complex of characters is unknown in Mexico and Central America. In South America it is approached only in mixtures from the eastern rivers and in certain primitive popcorns. Іп prehistoric times, varieties with similar cobs and kernels were the only type of maize along the west coast of South America for a very long period. ! Due to the presence of two of the world's worst maize diseases in this area, in itself a significant fact. 1949] STONOR & ANDERSON—MAIZE OF ASSAM 395 4. Тһе distribution of these and similar varieties іп the Orient 15 reviewed. They are widely though spottily distributed from Persia and Turkestan to Tibet and the Island of Hainan, nearly always among primitive and conservative peoples. 5. It is concluded that there have been at least two major movements of maize in Asia. The later one, in early post-Columbian times, brought what is essentially a Caribbean type of maize to the Philippines and to many of the countries actively colonized by the Europeans. Back in the hills, however, are more primitive types whose progenitors must have crossed the Pacific in pre-Columbian times, though in which direction (or directions) the evidence does not indicate. 6. The possibility of trans-Pacific transfer of a primitive and unaggressive race of maize by early Polynesians is discussed. It is shown to parallel the conclusions reached by the most recent workers on cotton. For the study of such problems, the imperative need of more critical taxonomic evidence on cultivated plants and weeds is discussed and illustrated by examples. 7. It is concluded that maize has had а long and complicated history. As a dominant crop it certainly developed in the New World. Ав a primitive, relatively unproductive crop, utilized for brewing, for popping, and for green corn, it is almost universal among the primitive peoples of central and southeastern Asia. Presumably it must either have originated іп Asia or have been taken there in pre- Columbian times. Before we can discuss the history and origin of maize intelli- gently we shall need an approximate survey of the kinds of maize being grown by these peoples. BIBLIOGRAPHY Anderson, Edgar (1943). A variety of maize from the Rio Loa. Ann. Mo. Bot. Gard. 30:469- 474. -------, (1944). Maíz reventador. Ibid. 31:301-31 ‚ (1946). Maize in Mexico. Ibid. 33:147-247. , (1947). Corn before кы 24 рр. Des Моіпев, la. ‚ and William L. Brown (19 . The northern "nias corns. Ann. Mo. Bot. Gard. 34:1-22. Р" Н. C. Cutler (1942). Races of Zea Mays. 1. Their recognition and classification. I 29:69-86. Buck, Peter Н. (1938). Vikings of the Sunrise. New Уог Bc new type of Indian corn from M U. S. Dept. Agr. Bur. Plant Ind. 5 E Lil Z P E => -------, (1920). 4. maize from Upper Burma. Science, N. S. 52:4 ШР Hugh C. (1946). Races of maize in South America. Bot. Mus. ү Нагу. Univ. 12:257— ева , Ebenezer (1849). Agriculture of New York 2:263-265. Nat. Hist. М. Y. Pt. V, Vol. 2 Von Fürer- xu МА (ор: (1939). Тһе Naked Мадаз. 243 рр. London. Hutchinson, J. В., R. А. Silow, and S. G. Stephens (1947). The evolution of Gossypium. 160 pp. Oxfo 5 Kuleshov, N. М. (1928). Some peculiarities in the maize of Asia. Bull. Appl. Bot. and РІ. Breed. 19:325-374 ------ (19 25). = geographical distribution of the varietal diversity of maize in the world. Ibid. 20:506-50 7 СЕЛЕ The maize of Mexico, Guatemala, Cuba, Panama, and Colombia. Ibid. Suppl. Laufer, B. (1907). The introduction of maize into eastern Asia. Cong. Int. des Americanistes. pp. 22 2 ) 3—253. Quebec 396 [Vor. 36, 1949] ANNALS OF THE MISSOURI BOTANICAL GARDEN ж = W. (1948 ). ннн histology of the female inflorescence of Zea Mays L. Ann. Мо. Gard. 35:5 Manga Paul С, and R G. Reeves . Sta. Bull. j ge . Earl ed Jr. (1949). New archaeological evidence on evolution in maize. Вог. E n TAM Univ. 13:213-247. T E (1941 шш species. Proc. (1939). The origin of Indian corn and its relatives. Texas Man's influence on the — of Polynesia with special reference to Sixth Pacific Cong. 4:629—639. EXPLANATION OF PLATE PLATE 18 Fig. 1. Monba tribe. Threshing maize, Dirang village. (November, 1946). Fig. 2. Monba tribe. Maize stacked for ripening, Pakung village. (November, 1948). 18 PLATE Bor. GARD., Vor. 36, 1949 ANN. Mo. зе at инста ы, А55АМ ОЕ STONOR & ANDERSON—MAIZE [Vor. 36, 1949] 398 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION Or РЕАТЕ PLATE 19 Fig. 1. Eastern Dafla tripe. Carrying in the maize from the fields. (November, 1946). Lhota Naga tribe. Corner of a "jhum" field, with lines of maize growing among the rice. (May, 1947). PLATE 19 GARD., Vor. 36, 1949 ANN. Мо. Вот. ERSON—MAIZE ОЕ А55АМ ОК & АМО STON [Vor. 36, 1949] 400 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 20 Three views of Stonor No. 13 (Early Slender). Scale indicated by the lines in the background which were originally spaced at 50 cms. Below: habit of plant when tassel gan to shed pollen. Note the "spathe" of upper leaves which is directed towards the the same, viewed from the side. Upper right: close-up of tassel after pollen shedding was complete. Note ТОВА of upper leaf to base of tassel. Photo- graphs courtesy of California Institute of Techno ANN. Мо. Вот. Сакр., Vor. 36, 1949 PLATE 20 STONOR % ANDERSON—MAIZE ОЕ А55АМ [Vor. 36, 1949] 402 ANNALS OF THE MISSOURI BOTANICAL GARDEN ExPLANATION Or PLATE PLATE 2 wer left: habit of Stonor No. 16 (Drooping Waxy). Scale indicated by the lines in the background, originally spaced at 50 cms. Note ч of upper leaves due го short internodes. Tassel had been shedding for some tin was nearly mature when the 22 was taken. Photograph courtesy of California Institute of Technology. Right: ar of 5 г No. 18 (Late Sidewise). Approximately natural size. Above: portion of car No. риса natural size. Photographs of еагѕ, courtesy of Royal Botanic Gardens, dede England. ANN. Mo. Вот. Ganp., Vor. 36, 1949 PLATE 21 STONOR & ANDERSON—MAIZE OF ASSAM [Vor. 36, 1949] 404 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 22 Ears and plates of Late Upright and Early Upright. Upper left (with stone back- ground), photograph of Stonor No. 8, taken in Shillong, Assam. Note erect and ribbed leaves, still slightly twisted, and that ears scarcely extend from the axils of the leaves. Center above: Stonor No. 5, photographed at a slightly later stage just as the silks were beginning to wither and after all the pollen had been shed. Note that tassel is not com- pletely exserted from upper leaves and that mature ears scarcely exceed their subtending sheaths. Center below, tassel of same plant with its uppermost leaf. per right: Stonor No. 6, habit just as tassel was beginning to shed pollen. Note tassel practically hidden by the upper leaves, which are twisted, upright, and ribbed. Scale in this and previous two figures indicated by lines which were originally spaced at 50 cms. Photographs courtesy of the California Institute of Technology. Lower right: ear of Stonor No. 8; lower left: ear of Stonor No. 6, both a little less than natural size. with th These two ears are almost identical е popcorns found in early prehistoric graves in coastal Peru (Paracas, Cañete, etc.). Photographs of ears courtesy of Royal Botanic Gardens, Kew, England. 1949 36, GARD., Vor. Мо. Вот. ANN. STONOR & ANDERSON—MAIZE OF ASSAM "HERMANN Moa Scil, Patholog Jesse М. GREENM Саке Paes pr ihe Herbarium Савко W. Dopct, Mycologist * EDGAR дынан, Genetic "P RT E. Жоор R. "arator of the Ныша ы N. es Paleohotanist oo — Number 4 Annals of the issouri Botanical Garden Uses of Maize in the Sierra of Ancash 24. rles M. Rick Annals of the Missouri Botanical Garden Vol. 36 NOVEMBER, 1949 No. 4 ON SOME USES OF MAIZE IN THE SIERRA OF ANCASH CHARLES М. RICK! anv EDGAR ANDERSON In its long association with man, maize has had a complicated career. It has been used by various peoples and in various ways. One might compare its whole history to a complex fabric, its warp the multitudinous varieties of this versatile crop, its woof the myriad uses to which Zea Mays has been put by the peoples who have grown it. In interpreting and understanding this history and in developing the broadest use of this world asset, one can never predict which of these various strands will be most useful in unravelling some particular problem. Опе finds ethnological curiosities leading to modern technological progress; for example, waxy maize, developed by Asiatic aborigines (Collins, 1909), became the clever solution to wartime shortages of industrial carbohydrates (Sprague and Jenkins, 1948). In reporting these rather unusual uses of maize in the South American highlands we would not venture to predict whether their greatest significance might be to an historian, a biochemist, an archaeologist, a plant breeder, or to some imaginative industrialist. Ancash is a small department including part of the Coast and Sierra of Peru north of Lima. It is largely drained by the Río Santa, one of the largest rivers of the western slopes of the Peruvian Andes. "Throughout most of its length the Santa is flanked on the west by the Cordillera Negra and on the east by the Cordillera Blanca, the latter completely dominating the scenery of Ancash with its chain of very high peaks and their glaciers and perpetual snowfields. Excepting its upper and lower extremities, the large trough between these two ranges is known as the Callejon de Huailas (sensu latu). The Santa varies in elevation in this region from 3,370 m. at Recuay to 2,150 m. at Villa Sucre (Weberbauer, 1945). Sugar cane and other tropical and subtropical crops are cultivated under irrigation in the lower reaches, maize and small grains in the upper part, the latter lCollege of Agriculture, University of California, Davis; Fellow, John Simon Guggenheim Me- morial Foundation, 1948-49. (405) [Vor. 36 406 ANNALS OF THE MISSOURI BOTANICAL GARDEN crops being irrigated in the few areas in which the ground is reasonably flat and in which water is accessible. On the higher slopes, up to 4,300 m., pastures, potatoes, barley, fava beans, and other crops are grown without irrigation. Maize and other crops dependent upon rainfall are grown in the rainy season from October to May. The Callejon de Huailas is one of the most populated valleys of the Peruvian Sierra. The great majority of the people are pure-blooded Indians or descendants of Indians. They live in the larger centers, in small outlying villages, or in solitary dwellings in the fields. Typical of the Peruvian Indians, they live according to the same primitive customs that their forbears have followed for centuries, almost without modification by contact with modern civilization. Ав a closely knit group, however, they differ in varying degrees from the serranos of other sections: they dress in their own characteristic costumes; their quechua language is greatly modified; and they deviate in certain uses of food plants. Much of the commerce and agriculture of the region is managed by a small group of pure whites, or nearly pure whites, whose life is influenced to a surprising extent by the Indians with whom they live. This influence is evident in their language and foods. Part of the information for this publication was gathered by the senior writer from informants and from observations made during a stay of two weeks in January, 1949, in the Callejon de Huailas. We are deeply indebted to Señora Elola Haro de Guzmán, our chief informant, who offered generous hospitality and answered innumerable questions with utmost patience and understanding. She has always lived in the vicinity of Huaráz, capitol of the department of Ancash. From the time of her husband's early death, some 20 years ago, she has personally man- aged the affairs of the family farm or chacra, іп which, like many Peruvian chacras, the Indian laborers live and work in what might be called a benevolent sharecropper basis. Through this experience and her many other contacts with people in the area, Sefiora de Guzmán has become thoroughly versed in the life of the Ancash Indians. Maize, the most important food plant in the Sierra of Ancash, is used in a great variety of recipes. The most unusual use of maize and one which apparently is not known outside Ancash is tocos de maíz or fermented maize. Tocos—The most popular maize variety for the preparation of tocos is cusqueno blanco, a variety typically having eight-rowed ears of enormous grains with soft floury endosperm. If cusqueno blanco is lacking, other white-grained varieties are used. The maize is used only in the mature state. The fermentation or rotting process is conducted in the following fashion. The whole ears, husked, but with grains still attached, are placed in any quantity in a sack of linen (wool or cotton cannot be used presumably because they would dis- integrate in the process). The sack is tied shut securely and is submerged in water in a hole that has previously been dug in an irrigation ditch where the water is flowing freely. The hole must be deep enough so that the sack and its contents are covered by at least two inches of water. Stones are placed on top in order to 1949] RICK & ANDERSON—USES OF MAIZE 407 prevent the sack from rising above water. Aside from the importance of keeping the sack submerged, the depth of submergence does not seem to matter. A very important part of the process is to place the stones and to arrange the surrounding ground and grass so that the cache cannot be readily detected. Tocos are very popular and therefore valuable items of commerce, and their loss by theft is by no means uncommon. The maize can also be fermented in large pools of standing water, but it is considered that a product of better quality is produced in running water. Whether the water is standing or running it is essential that the maize be completely sub- merged in water in a hole in the ground. Whatever changes occur, the fermentation must be anaerobic or semianaerobic. The length of time required for the process depends on the age of the maize and probably also on the temperature of the water. When recently harvested ears are used the fermentation is complete in two months, but if the ears are older and the grains harder it may be necessary to wait as long as three months. The stage of fermentation is determined by touch, the process being completed when the grains are soft, at which stage they are bloated and have a somewhat water-soaked appearance. When the desired stage of fermentation has been reached, the pericarp is re- moved from the grains, which still remain on the ear, by rubbing with the fingers. It is not possible to do this before the fermentation is completed. The ears are then washed in cold running water. They are washed well, but gently, so that none of the starch is lost. At this stage the /осо$ are ready for cooking and can be stored in a moist con- dition for no more than seven or eight days. If it is desirable to keep them for а longer period, they are dehydrated in the sun. The grains are shelled from the ears and are spread thinly in a place where they will receive the maximum amount of sunlight. Depending upon the light intensity, from four days to two weeks are required to complete the dehydration. If the grains are well dried they can be stored for one or two years under the generally cool household conditions of the Sierra. Fresh focos are considered to make a better product than dried ones. They are marketed in both forms. The usual cooking process consists of stewing them in an olla, or low earthen- ware pot with a mouth nearly as large as the largest diameter. The /ocos are placed in the olla and enough boiling water added to cover them. Either refined cane sugar or cancbaca, a crude brown sugar considered to be superior to refined sugar for this purpose, is added, the amount depending upon taste. Sugar is required to offset the natural acidity of the product. Sometimes herbs are added for flavoring, but the uses and purposes of these are not well understood and are said to be secrets of the Indians. The mixture is cooked for 20 or 30 minutes. The odor of this dish, which is not the least bit dissipated by cooking, is reminiscent of vases of flowers in which the water has not been changed often enough. It is just barely possible for the uninitiated to stay in the same room when [Vor. 36 408 ANNALS OF THE MISSOURI BOTANICAL GARDEN tocos are served. They are much more agreeable to taste than to smell, according to the experience of the senior writer. They are eaten in great quantities without ill effect and are very popular both among Indians and whites of the region. They are probably no more offensive to us than our sauerkraut or highly scented cheeses would be to the Ancash Indian. It is conceivable also that certain valuable vita- mins, possibly of the B complex, might be elaborated in the rotting process. Tocos are generally eaten for lunch and dinner, but there is no great regularity, they being consumed also at other hours of the day. They are served warm like a stew, but are also very popular chilled after cooking. "They are considered effective as a remedy against colds. There is no distinction as to age of the person eating focos or occasion on which they are served. In the recollection of Señora de Guz- mán they have retained a constant popularity in her time. In the Sierra of Ancash possibly 20 per cent of the maize is consumed in the form of /ocos. More extensive use is probably limited because they are more diffi- cult to prepare than other maize foods. Yungay (2,535 m.) and Carhuas (some- what higher elevation) in the Callejon de Huailas are the principal centers of prep- aration of /ocos. From these centers they are transported to other markets in Ancash. Because they are in brisk demand, they sell rapidly whenever offered. "Тһе reasons for the development of this use in these places is not well understood. Climatic conditions can hardly be responsible for the restriction to Yungay and Carhuas because focos of good quality can be prepared at Huaráz, which lies at a considerably higher elevation (3,080 m.) We inquired extensively in both the Sierra and Coast of Peru, but found no evidence of the use, and very little evidence of the knowledge of tocos outside the Sierra of Ancash. Informants would nearly always indicate familiarity with tocos upon inquiry and upon mention that it was a fermented maize product, but further questioning would generally reveal that they were thinking of chicha or some other product and that they had never actually heard of /ocos. It is impos- sible to state the antiquity of this use of maize. Since it is used universally by the Indians of Ancash and since it is known only by a name in the quechua language, it seems likely that its use antedates the colonial period. We are indebted to Dr. Carl O. Sauer, of the Department of Geography, Uni- versity of California, for calling our attention to the following quotation from the works of de Champlain, which leaves no doubt that a similar product was used by the Huron Indians: They have another way of eating Indian c pid to vp “7 they take it in the ear and put it in water under the mud, leaving it tw three mo in that state until they udge that it is putrid; then they take it out a it баш» meat ог fish and th t They also roast it, and it is better that way pe I but I assure you that nothing smells so bad i when it comes ter all covered wit ud; yet the women ut of the y and children take it and suck it like n cane, 4. being nothing they like better, as they plainly show.—[Biggar, 1929, 3:129—130.] 1949] RICK % ANDERSON—USES OF MAIZE 409 A similar product, ѓосоѕ de papa, is prepared from a potato variety called anco having rather dry white flesh. Methods of fermenting and even of drying the fermented tubers are almost identical to those used for maize. According to Señora de Guzmán /ocos have a medicinal value in addition to their putative value in curing colds. They are used in the following manner to cure filmy eye of the burro or horse. Dehydrated tocos de papa are finely ground and passed through a fine screen. The fine powder thereby obtained is blown into the infected eye through a small tube of paper. Cancha—In the vicinity of Huaraz this name applies to parched maize, but in other parts of Peru it appears to be a general term pertaining to both parched and popcorn. Even in its restricted use to parched corn, it is by far the most popular form of prepared maize in the Callejon de Huailas. Perhaps 50 per cent of all the corn in the Sierra of Ancash is used as cancha. Almost any form of maize can be used for cancha, but tercio pelo or maiz dulce, a variety having starchy, hard, rounded grains of reddish brown color with a yellow tip, is preferred. Another variety, pacchus, apparently a true sweet corn, is also considered satisfactory for this use. For the preparation of cancha (and probably also popcorn) the Indians of Ancash use a baked clay vessel called the tiesto, which is mound-shaped and slightly rounded on the bottom. It has a small opening on the side and may or may not have a handle. They are generally 20-30 cm. in diameter. Valcarcel (1934) described and illustrated a similar vessel that was unearthed in the ruins of Sacsahuaman (department of Cuzco). This specimen or one very similar to it was seen by the senior writer оп a visit to the Instituto Arqueológico Fig. 1. Olla canchera, a prehistoric vessel used to parch maize. Reproduced from Valcárcel (1934). [V^L. 36 410 ANNALS OF THE MISSOURI BOTANICAL GARDEN del Cuzco and is copied by photostat from the cited paper as the accompanying fig. 1. Valcárcel describes it as follows (p. 228): 08--ОПа canchera tripode de barro cocido. Con asa. Sin pulir y sin ornamentar. Leva s seis рн cerca del asa y tiene la base ennegrecida наа as m por el fuégo. mifina. Por lo diminuto de su tamafio parece juguete, réplica de otros mayores. Alto ре ст. . Diam. de la boca 3.7 cm., Diam. de la base 6.2 cm. (Vessel of fired и with three legs, used for preparing cancha. With handle. Without polish or ornamentation. It has six holes near the handle and its base is blackened, probably by fire. Semifine Меен Бог the smallness of its size іс appears со Бе а toy, а ге г of other larger ones. Height 6,2 cm., Diameter of mouth 3.7 cm., Diameter of base 6.2 cm.) Another example described as a brazier or small stove was illustrated by Bing- ham (1930). This artefact, dug from the ruins of Machu Picchu, closely resembles the preceding one in shape, but is larger, being 17 cm. high. In his more recent book, ‘Lost City of the Incas’ (1948), one is illustrated opposite page 42, where it is described (erroneously, we suspect) as “А Brazier for Annealing Bronze Articles.” The tiestos of Ancash resemble these precolumbian ollas in general shape except that they have smaller mouths, lack the legs, and may or may not have the handle. The hooded form of these vessels prevents the escape of grains that jump during the parching. The senior writer has also seen cancha prepared in Huaraz in open earthenware vessels having the general shape of our frying pans. To prepare cancha only mature grains are used. These are placed without other ingredients in a tiesto that is heated as hot as possible over a wood or charcoal fire. The grains are soon toasted and slightly expanded by the heat. After they have reached this stage the grains are removed and cleaned on paper or cloth. They may be flavored with melted lard and salt. A supply of cancha is usually not kept for more than one day. Two batches—one in the morning and another in the afternoon—are usually prepared per day. Cancha is immensely popular in Ancash and throughout the Peruvian Andes. The parched grains may be eaten at any time of the day and at all times of the year. Everyone, children included, eat it. It is very convenient for the worker to carry a pouch of it with him at all times and to crunch away on the grains when- ever hungry. When cancha is lacking in a household, it is bought in the market or from a neighbor. It is veritably “el pan de los indios mejor dicho.” Та Ancash cancha is not ground to prepare a flour. The only type of arina or flour that is prepared from maize is ground from untreated grains or from chochaca. Chicha—This mildly alcoholic beer-like drink is by far the most important bev- erage of the Sierra. We are aware of several methods of preparation, but we did not investigate in detail the processes employed in Ancash since they are similar to the well-known methods of the Peruvian Sierra. Methods of preparing chicha in vari- ous sections of Bolivia are described in detail by Cutler and Cárdenas (1947). Chicha de jora, considered the chicha of best quality, is prepared from malted maize. Grains of several different yellow or red varieties are sprouted and then 1949] RICK & ANDERSON—USES ОҒ MAIZE 411 dried. The grains are stored or marketed in the dried sprouted condition. These are ground and mixed with water in large earthenware jars for fermenting. Chicha morada, another well-known product, is prepared from the maize variety known as morada having dark purplish grains and cobs. The dried grains are ground and the consequent flour is stewed with the cobs in water. The mixture is filtered and allowed to ferment. Other chichas are prepared from barley, wheat bread, fava beans, and other crops in Ancash. Mote—This is a form of hominy prepared from mature grains of preferably a white variety of maize. The grains are treated in a boiling lye solution (lejía) prepared from water and ashes, for one-half hour. The pericarp is then removed and the grains are boiled further until they burst. Mote is eaten іп this form or is ground for the preparation of masamoras ( puddings) or tamales. Mote, like tocos, can be dehydrated for storage. Chocbaca—Mature grains are shelled from the ear and cooked in a great quan- tity of water until they split slightly. The grains are then spread in the sun to dry and are stored for whatever time they might be needed. The drying process is facilitated by first exposing the grains to frost on a cold night before spreading them in the sun. Chochaca is used mostly for soups, for which purpose it is ground. One might point up this picture of maize in these remote highlands by con- trasting it with maize in the United States where it is primarily our medium for producing the maximum amount of beef and bacon per man hour per acre, and secondarily a most delicious vegetable. Or contrast either of these with uses in Mexico where it is indeed the immediate staff of life for nearly every citizen and ordinarily is eaten directly at every meal of every day in the year, as fortillas, tamales, and a variety of lesser-known foods such as atole, pinole, posole, etc. It is worth noting that one of these Peruvian recipes begins by parching the kernel, another by prolonged soaking. The parching (or popping) of the kernel (usually followed by grinding it into a fine meal) is apparently one of the oldest and most widespread uses of maize. It is almost universal in the New World and is widespread in the mountains of central and southeastern Asia. Though many of the products are delicious, they have tended to disappear under the impact of modern sophistication, ultimately to reappear as the ultra-modern, ultra-sophisti- cated, standardized, mass-produced, trade-marked cocktail wafers such as “Fritos.” As to the prolonged soaking of the kernel, this idea seems to have its roots deep in western South America. Of a collection of recipes obtained from an old family in Antiochia, Colombia, through the kindness of Srta. Julia Guzmán Naranjo, nearly half began by soaking the kernels from overnight to four or five days or longer. It is noteworthy that recipes involving long soaking are apparently un- known to Mexican cooks or considered too trivial for polite discussion. [Vor. 36 412 ANNALS OF THE MISSOURI BOTANICAL GARDEN SUMMARY 1. The common uses of maize are described for the Sierra of Ancash, a some- what isolated region of Peru north of Lima, with a conservative, predominantly Indian population. The two commonest maize foods in the Sierra of Ancash are focos de maíz and cancha. 'The former is made from kernels fermented under water for two months or longer. The latter consists of whole kernels parched in a special dome- shaped vessel called a tiesto or olla cancbera. Prehistoric examples of these utensils are known from the southern highlands of Peru. 3. These foods are discussed briefly in relation to the history and geographic distribution of uses of maize. LITERATURE CITED Biggar, H. P. (1929). The works of Samuel de Cham edi 6 аа ее бос. Тогопсо. Bingham, H. n Machu Picchu, a sag of D Inc Haven. — — ———, (1948). Lost City of the Incas. 263 pp. New nh Collins, с. > Ж 9). А new type of Indian x {ын Pies Џ. 5. Dept. Agr. Bur. Plant Ind. Bull. Cutler, H. С. T boves M. (1947). Chicha, a native South American beer. Bot. Mus. Leafl. Harvard len 13:33-6 Sprague, С. F., and Jenkins 2 D н The development of waxy corn for industrial use. Iowa State Coll. post Sci. 22.2 Weberbauer, A. (1945). El Ша: i etal de los andes peruanos. Min. de Agr. (Lima). Valcárcel, L. E. (1934). Los trabajos arquelógicos del Cuzco. II. Sajsawaman rediscubierto. Rev. Mus. Nac. (Lima) 3:211-234 5РОКЕ5 ОЕ THE GENUS SELAGINELLA ІМ NORTH AMERICA NORTH OF MEXICO ALICE F. TRYON Іп 1901 L. M. Underwood remarked that the old Selaginella rupestris was опе of several examples in which segregation of species had "successively expanded to the bounds that would cause the botanists of twenty years ago to suffer acute paralysis." Among American workers Underwood had already started this segre- gation three years earlier. The critical studies of Van Eseltine (1918), Maxon (1918, 1920, 1921), and most recently of Weatherby (1944, 1946) have added further new species. Additional collections and critical study will no doubt bring to light still other new species. Dr. Maxon has noted the need for an examination of the species complexes S. densa and S. Wallacei. The spores, particularly the characters of the megaspores, have appeared to be an integral part of recent descriptions of species. Mr. Weatherby has expressed the opinion that when used cautiously the pattern of spore sculpture is useful in species definition. The present study was undertaken to present a survey and an illustrated account of the spores of our native Selaginellas. The megasporangia are not very generously supplied with spores, each usually containing but four megaspores. Each spore has a hemispherical base and three plane triangular faces. The base ог outer face is the free surface in the tetrad and the three plane triangular surfaces—collectively, the commissural face—are the sides in contact in the tetrad. The commissural face is marked by three prominent commissural ridges which are united at the apex and radiate out at nearly equal angles to the vicinity of the equator. The ends of the ridges are sometimes con- nected by an equatorial ring. The spore surface may be smooth, granular, rugose, rugose-reticulate, or tuberculate wholly or in part. The enclosed areas in a reticulate pattern are called areolae. 'The photographs convey the type of sculpture more accurately than a descriptive statement, and, unless remarks on variations accompany the data for each species, the illustrations can be regarded as representative of the material examined. It rarely happens that one or two of the megaspores within a sporangium may develop at the expense of the others, which are then much dwarfed. The mega- sporangium of S. rupestris usually bears two spores although occasionally three or а single spore may develop. The spores of this species characteristically lack com- missural ridges but some spores bear a single straight or circular ridge. Occasionally sporangia contain one or two peculiar "dumb-bell"-shaped spores in which there has been an incomplete division (Lyon, 1905). One megasporangium of S. pilifera was observed to have eight megaspores. The size of the megaspores range from 0.15 mm. in diameter in S. armata to 0.53 mm. in S. selaginoides. The measurements given in the text are of the (413) [Vor. 36 414 ANNALS OF THE MISSOURI BOTANICAL GARDEN greatest diameter. The mode is used to express the central position of the measure- ments. It is printed in italic between the extremes. The color of the spores was determined at 45 diameters, using natural light and a black background. The color range is expressed according to Ridgway’s ‘Color Standards.’ The popular term of color approximately equivalent to that of Ridgway’s is in parentheses. There is marked variation in color in some species, immature spores often being of a different shade than the mature spores. The megasporangium was removed from the strobilus and soaked in 50 per cent alcohol. After two or three minutes it was sufficiently pliable to dissect out the spores. Some care had to be used to prevent them from escaping for they were easily blown or jarred from the dissecting field after the alcohol dried. The spores were mounted dry, on a glass or mica slide, within a ring of cement built up to support the cover glass, and this was sealed with cement. The megaspores were photographed through a compound microscope using a 48-mm. microteliplat lens as the objective and а 15 X hyperplane lens as an ocular. Additional magnification was achieved through suspending a 10 ocular above the microscope. The camera, a Bausch & Lomb Model K, with 8-inch bellows, was placed over this. An image in focus at the third lens was also in focus on the ground glass of the camera. Four Leitz microscope lamps with green filters were placed around the microscope at the level of the stage. These supplied a very satisfactory source of illumination, for the intensity and angle of the light could be adjusted for each lamp. The film used was Eastman Ortho-X, and the exposure time varied from 20 to 90 seconds. Each microsporangium bears several hundred microspores which are borne in tetrads and have the same shape as the megaspores and are sculptured in essentially the same manner. The commissural face usually is marked with three commissural ridges although some have a single ridge. The spores of some species have a promi- nent wing at the equator. The surface of the microspores also bears distinctive markings such as spines, tubercules, or rugae, but it is difficult to find and to rec- ognize mature material. Photographs of the microspores are included when they are representative and show good detail for the species. Twelve spores were measured for each species. The microspores of S. rupestris are similar to the mega- spores in that they frequently lack the commissural ridges and an occasional dumb- bell type is found. The microspores are roughly about one-eighth as large as the megaspores, rang- ing from 23 p in diameter in several species to 64 и in S. fortipila. There is no definite correlation between the size of mega- and microspores of the same species. For example, S. armata and S. selaginoides have microspores of a similar size while the mode of the megaspores of the former is 0.19 mm. and that of the latter is 0.53 mm. А magnification of at least 400 diameters is necessary to study the surface detail of the microspores, and 70 or 80 diameters is desirable for examina- tion of surface detail of the megaspores. 1949] TRYON-—SELAGINELLA SPORES 415 The microspores are generally more deeply colored than the megaspores, tending toward deep orange or red rather than the lighter yellows, although in some species, as S. selaginoides, both mega- and microspores are of the same color. Slides were prepared by breaking open a microsporangium on a glass slide, adding a drop of lactic acid, and sealing the mount with ringing cement. А lactic acid preparation was more satisfactory than a dry mount, for sculpture not other- wise evident could be distinguished. H. W. Morris, photographer for the University of Minnesota Hospital, photographed the microspores. А homal 3-ocular and а 8-mm. objective were used with а 46-cm. bellows extension of the camera. Illumi- nation was from a carbon arc lamp with a Kaisering green filter. Polychrome plates were used. The collections studied, for the most part, were identified by such authorities on the genus as Maxon, Van Eseltine, Weatherby, and Wherry. Specimens were examined from the collections of the American Fern Society, Chicago Natural History Museum, Gray Herbarium, University of Minnesota, Missouri Botanical Garden, United States National Museum, and the University of Wisconsin. I wish to express my appreciation to the curators of the herbaria of these institutions who have so kindly lent material for study. This study has been mainly done at the Missouri Botanical Garden where facili- ties were generously granted by the Director. Тһе photography was done at the University of Minnesota and the costs incurred supported by the Department of Botany. The problem was initiated at the University of Wisconsin, where a study of Selaginella spores was presented as a master's dissertation. I am especially grate- ful to my husband for the preparation of the photographic unit for the megaspores and for the suggestions and aid given in the study. SYNOPSIS OF SELAGINELLA IN NORTH AMERICA NORTH or Mrexico The following synopsis has been primarily adapted from the current literature, and references are given to the papers upon which it is based. Since there may be some question regarding the disposition of certain species, now generally reduced to synonymy, it was thought desirable to include photographs and discussion of the spores of these. Characters presented in the synopsis allow it to be used to a cer- tain extent as a key. However, the characters are general ones relating to whole groups and are not necessarily without exception. Each species bears the same number in the text and plates as it does in the synopsis. SUBGENUS EUSELAGINELLA! Vegetative leaves uniform; sporophylls uniform. GROUP or S. sELAGINOIDES. Strobilus cylindric 1. S. SELAGINOIDES L.) Link Group or S. RUPESTRIS. Strobilus tetragonous. a. Stems typically erect, rooting only at the base. . Setae tortuous. SUBGROUP OF S. TORTIPILA 2. S. токтірп.А А. Br. 2. S. Sberwoodii Underw. (see Wherry, 1936) l' The subgenera, groups and series are taken from Walton and Alston (1938). 416 b. бесае straight. SUB GROUP OF s. BIGELOvII. Megaspores rugose оп the outer face... SUBGROUP OF S. ARENICOLA. Megaspores smooth or very nearly so о п the outer face a. Stems prostrate, rooting throughout. e sein мнр Minen НЕ Stems elongate, slender, branches remote. а Weatherby, p SUBGROUP OF 5. RUPESTRIS. Stems short, stout, branches congested.. e w e w Оо © N © © 10. — N - > 14. . S. WALLA . S. siBIRICA (Milde Hi [Vor. 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN S. RUPINCOLA Underw. S. Con v1 Weatherby . S. Всегоун Underw. . S. NEOMEXICANA Maxon S. RipprLLI Van Eselt. Ве (see Clausen, . E ACANTHONOTA Underw. . S. floridana Maxon (see Clausen, 1946) S. ОКЕСАМА D. C. Eat m T Ra OMNES Hieron. . S. MUTICA D. C. E aton . 5. CINERASCENS А. А. Eaton S. VATSONI Underw. S. STANDLEYI Maxon . S. LEUCOBRYOIDES M axon СЕТ Hieron, ieron. S. ASPRELLA Maxon Rydb. . S. DENS T RUPESTRIS (L.) pring . S. SCOPULORUM 22 Maxon 25. S. ХУкіснті Hieron, c. Stems strongly dorsiventral. SUBGROUP OF S. HANSENI. Vegetative leaves uniform or nearly so.. 24. + SHELDONI Мах 25. S. HANSENI Мили. SUBGROUP S. ани Vegetative leaves dimorphous. (see Weatherby, 1943 26. 5. ARIZONICA Maxon 27. S. EREMOPHILA Maxon SUBGENUS STACHYGYNANDRUM Vegetative leaves dimorphous, sporophylls uniform. SERIES DECUMBENTES. Widely creeping, prostrate, stems rooting through- out 28. 29. 30. S. Dovcrasn (Hook. G S. APODA (S. ludoviciana hi Br.) 1949] TRYON—SELAGINELLA SPORES 417 SERIES CiRcINATAE. Tufted xerophytes, ascendent, rooted only at the base.. 31. S. LEPIDOPHYLL (Hook. & Gr 25 Sprin . $. PILIFERA А.В G 2 hh orton, 1939, and Weatherby, 1943b) DESCRIPTION SUBGENUS EUSELAGINELLA GROUP OF S. sELAGINOIDES.— 1. S.sELAGINOIDES (L.) Link. Pih 1. Megaspores Barium Yellow (yellow, green-tinged); 0.48—0.52—0.53 mm. in diameter, 18 spores measured. Microspores Barium Yellow (yellow, green-tinged) ; 26—34 y in diameter. GROUP or S. RUPESTRIS.— SUBGROUP OF S. TORTIPILA.— The two members of this group cannot be distinguished on the basis of spores. This would agree with Wherry's (1936) conclusion that S. Sherwoodii is merely an ecological form of S. tortipila. The megaspores of S. Sherwoodii in the collec- tion examined are strongly tuberculate-rugose. Those of S. fortipila are of a similar pattern of sculpture and some are less prominently marked. 2. S. токтірп.А А. Br. В Е 0, Megaspores Straw Yellow to Lemon Chrome (lemon-yellow) ; 0.25—0.34—0.41 mm. in diameter, 26 spores measured. Some spores are more prominently rugose on the outer face than in fig. 2b. Microspores Pinard Yellow (pale yellow); 41- 64 и in diameter. 2. S. Sherwoodii Underw. Pl. 23, fig. 2 Megaspores Apricot Yellow (pale orange); 0.32—0.40 mm. in diameter, 9 spores measured. Some spores are less prominently tuberculate-rugose on the outer face than in fig. 20. Microspores Apricot-Yellow (pale orange); 38-63 y in diameter. SUBGROUP OF S. BIGELOVII.— Megaspores of S. rupincola show a marked variation in prominence of sculptur- ing. Fig. 3b represents an extreme phase with a well-marked equatorial ring. For the most part, the megaspores examined are sculptured to a less degree as illustrated in fig. 3a, but with a more strongly pronounced equatorial ring. Megaspores of S. Bigelovii examined are finely rugose and usually bear an equatorial ring. Some megaspores are marked with subechinate, lace-like prominences which can be seen along the margins of the spores illustrated. Commissural face of megaspores with moderately coarse rugae; outer face rugose to rugose-tuberculate, ridges low and crowded, areolae narrow if present . S. RUPINCOLA Commissural face of megaspores with fine rugae; outer face with 1 uh scure rugae, areolae small, scarcely distinct from the low, rounded geo 5. S. BicEL ОУП [Vor. 36 418 ANNALS OF THE MISSOURI BOTANICAL GARDEN 3. S. RUPINCOLA Underw. Pl. 24, fig. 3. Megaspores Apricot Yellow (pale orange); 0.19-0.20-0.34 mm. in diameter, 29 spores measured. Microspores Deep Chrome (bright orange); 38-64 p in diameter. 4. S. Совут Weath. Am. Fern Jour. 36:51—53. 1946. Spores of this species were not available for photography but two megaspores from the туре have been examined. They are rather similar to those of S. rupin- cola except that they lack an equatorial ring. They are about 0.4 mm. in diameter and Apricot Yellow (pale orange). The rugae are prominent on the outer face and there are distinct areoles between the rugae. 5. S. Bicer ovi Underw. РІ. 24, fig. 5. Megaspores Apricot Yellow (pale orange); 0.25-0.34-0.40 mm. in diameter, 49 spores measured. Microspores Apricot Yellow (pale orange); 38-49 p in diameter. 6. S. NEOMEXICANA Maxon. Megaspores unknown. SUBGROUP OF S. ARENICOLA.— The members comprising this group are difficult to distinguish on the basis of the spores. Clausen (1946) has recognized three species and remarked that of these S. arenicola and S. acantbonota are not wholly separable. The commissural face is more strongly marked than the outer face in all of the species. However, the sculpturing is rather variable and there appears to be no marked character which distinguishes the species except the degree of prominence of the sculpturing. The megaspores of S. Riddellii are more strongly marked on the outer face than those of the other species. In the collections studied the megaspores of S. acantbo- nota are as prominently rugose-reticulate оп the commissural face as those of S. Riddellii, and phases of sculpture оп the megaspores of S. acanthonota and S. arenicola are indistinguishable. Megaspores Sa ie on outer face 7. S. RIDDELL Megaspores smooth on outer face. Scarcely sek on the commissural face Rugose to rugose-tuberculate on commisural face 8. S. ARENICOLA 8. S. funiformis 9. 5.А 9. 5. анде апа 7. S. Ктрркглп Van Eseltine. РІ, 24, fig. 7. Megaspores Lemon Chrome (lemon yellow); 0.21—0.32 9 0.38-0.46 mm. in diameter, 16 spores measured. "The outer face is usually more prominently rugose than іп fig. 7b. Microspores Apricot Yellow (pale orange); 34-53 p in diameter. 8. S. ARENICOLA Underw. Pl. 24, fig. 8. Megaspores White to Apricot Yellow (pale orange); 0.25-0.20-0.42 mm. in diameter, 35 spores measured. The outer face is smooth or minutely punctate. Microspores Apricot Yellow to Deep Chrome (pale to bright orange); 34-47 p in diameter. 1949] TRYON-—SELAGINELLA SPORES 419 $. S. funiformis Van Eseltine. Pl. 25, fig. Š. Megaspores Straw Yellow to Pinard Yellow (pale yellow); 0.21-0.34 mm. in diameter, 6 spores measured. The spores examined are similar in pattern of mark- ing to those of S. arenicola but are of a deeper yellow. Місғоѕротеѕ Deep Chrome (bright orange) ; 30-41 шіп diameter. 9. 5. ACANTHONOTA Underw. РІ 25, fig, 9. Megaspores White to Lemon Chrome (lemon yellow); 0.29—0.36—0.42 mm. in diameter, 28 spores measured. The pattern of sculpture is similar but the degree of prominence varies between that represented in fig. 9а and 9b. Microspores Deep Chrome (bright orange); 34-45 и in diameter. 0. S. floridana Maxon. Pl. 25, fig. 9. Megaspores White to Lemon Chrome (lemon yellow); 0.19—0.32—0.40 mm. in diameter, 36 spores measured. Microspores Deep Chrome (bright orange); 32- 53 шіп diameter. SUBGROUP OF $. OREGANA.— The group of S. oregana as treated Бу Weatherby (1944) includes several Mexican species and is characterized on the basis of habit. The spores of the species treated here are all distinct and do not appear to indicate relationships between the species. a. Commissural eri of E more prominently M than outer face, commissural ridges connected by equatorial ring o b. Commissural гэ Ыы ur. rugose, оа Д thin and delica С. Gute face slightly reticulate, meshes Bor loue areolae scarcely more distinct than those of the commissural f 10. S. OREGANA c. Outer face strongly reticulate, m alid forming e shallow areolae more distinct than those on ms commissural fac 11. S. UNDERWOODII b. Commissural face coarsely and remotely rugose, сэн бои ridges road and coarse; outer m lightly rugose to smooth 12. S. MUTICA кле face of megaspores lightly rugose, less prominently sculp- tured than outer face, commissural ridges connected by a prominent, thin и ring 13. S. CINERASCENS 10. S. onEGANA D. C. Faton. РІ; 25, hs. 10. Megaspores Pinard Yellow (pale yellow); 0.27—0.32—0.36 mm. in diameter, 25 spores measured. Some spores bear cross ridges connecting the commissural ridges Microspores Pinard Yellow (pale yellow); 41-56 м in diameter. S. (мрекчоорп Hieron. Pl. 26, fig. 11. Megaspores Apricot Yellow (pale orange); 0.27—0.36—0.42 mm. in diameter, 38 spores measured. Microspores Deep Chrome (bright orange); 30-45 м in diameter. 12. S. MuUTICA D. C. Eaton. Pl. 26, fig. 12. Megaspores Apricot Yellow (pale orange); 0.21-0.20-0.42 mm. in diameter, 31 spores measured. Some spores bear an equatorial ring; some are more strongly rugose-reticulate than fig. 12a. Microspores Apricot Yellow (pale orange); 30-53 шіп diameter. 13. S. CINERASCENS А. A. Eaton. PL 26 Бр. 15. Megaspores Pinard Yellow (pale yellow) ; 0.32—0.37—0.38 mm. in diameter, 16 [Vor. 36 420 ANNALS OF THE MISSOURI BOTANICAL GARDEN spores measured. Microspores Apricot Yellow (pale orange); 38-53 шіп diameter. SUBGROUP OF S. RUPESTRIS.— The subgroup of S. rupestris includes the largest number and the most perplex- ing species of the subgenus. Several of the species are undoubtedly aggregate groups which need to be extensively collected and critically examined. Maxon (1920, 1921) has investigated some of these and has remarked on relationships between them, but for the most part a natural series has not been worked out. There is a remarkable similarity between the pattern of sculpture of the mega- spores of some of the species. The megaspores of S. asprella and one phase of the megaspores of S. densa cannot be distinguished. However, there is much variation in the spore sculpture between collections of 5. densa. Some collections of S. Watsoni have megaspores scarcely rugose and very similar in pattern to those of S. Standleyi; others have prominently marked spores rather like those of S. scopu- lorum. The megaspores of S. Wallacei are variously sculptured but frequently characterized by an equatorial ring. a. Megaspores 4 per sporangium; 3 commissural ridges. hrs face obscurely rugose- reticulate to nearly ink ula . Commissural face finely rugose-ret 14. S. WATSONI 15. S. STANDLEYI c. Commissural face with coarse, low ruga 16. S. LEUCOBRYOIDES b. Both Xm strongly rugose to лена ы; . Commissural face with coarse, remote мын outer face strongly reticulate, surface of the areolae broad and smooth . S. ASPRELLA 20. S. DENSA d. Commissural face with и. compact rugae; outer face with areolae на ог над арраге е. Commissural face ngly ru igose to rugose- -reticulate; nearly all rugae on outer poi joined in a net forming areolae of moderate size. f. Equatorial ring us dard present 17. S. WALLACEI f. Without equatorial ri 14. S. WATSONI 18. S. srBIRICA 2. S. SCOPULORUM e. ге faces with prominent flexuose rugae which have free ends, eolae narrow if present... S. WRiGHTII . Mega EN ўр per sporangium; commissural ridges lacking or а single ridge [ present 21. S. RUPESTRIS 14. S. Watrsont Underw. Pl. 26, fig. 14. Megaspores Apricot Yellow (pale orange); 0.27—0.40—0.50 mm. in diameter, 75 spores measured. The commissural ridges are usually straight and the commis- sural face somewhat less prominently sculptured than fig. 14a. Microspores Deep Chrome (bright orange); 30-53 и in diameter. 15. 5. STANDLEYI Maxon. РІ, 27, fig. 15. Megaspores Apricot Yellow (pale orange); 0.34—0.40—0.48 mm. in diameter, 24 spores measured. Some spores are more prominently rugose than fig. 15. Micro- spores Apricot Yellow (pale orange); 26-41 и in diameter. 16. S. LEUCOBRYOIDES Maxon Pl. 27, fig. 16. Megaspores Apricot Yellow (pale orange); 0.32-0.47 mm. in diameter, 8 spores measured. Microspores Flame Scarlet (red orange); 38-56 p in diameter. 1949] TRYON—SELAGINELLA SPORES 421 17. S. WALLACEI Hieron. Рі. 27, fig. 17. Megaspores Apricot Yellow (pale orange); 0.27-0.34-0.38 mm. in diameter, 51 spores measured. Some spores are less prominently sculptured than fig. 17, and some are without an equatorial ring. Microspores Deep Chrome (bright orange); 38—49 џ in diameter. 18. S. stprrica (Milde) Hieron. Pl 27, hp. 18: Megaspores Pinard Yellow to Apricot Yellow (pale yellow to pale orange) ; 0.38-0.40-0.42 mm. in diameter, 5 spores measured. The commissural ridges аге usually twice as long as those in fig. 18a. Microspores Deep Chrome (bright orange); 38-49 y in diameter. 19. 5. ASPRELLA Maxon. FEL ve he Megaspores Straw Yellow to Apricot Yellow (pale orange); 0.29—0.34—0.38 mm. in diameter, 6 spores measured. Some spores have an equatorial ring. Micro- spores Pinard Yellow to Deep Chrome (bright yellow to bright orange); 34-53 м in diameter. 20. S. DENsA Rydb. ШІ 295, 89.20. Megaspores Apricot Yellow (pale orange); 0.36—0.40—0.50 mm. in diameter, 48 spores measured. The spores photographed are from type material and illustrate the pattern of sculpture in several of the collections studied. Місғоѕротеѕ Deep Chrome (bright orange); 34—49 џ in diameter. 21. S. RUPESTRIS (L.) Spring. Pl. 28, fig. 21. Megaspores Deep Chrome (bright orange); 0.32—0.46—0.53 mm. in diameter, 53 spores measured. The sculpture of the spores examined is generally of the same pattern, varying in the prominence of reticulation. Microspores Apricot Yellow (pale orange); spores monolete, 49—75 и in greatest dimension. 22. $. scopULORUM Maxon. Pl. 28, fig. 22. Megaspores Deep Chrome (bright orange); 0.32-0.42-0.48 mm. in diameter, 35 spores measured. Microspores Apricot Yellow (pale orange); 34-56 џ in diameter. 23. S. WnicHrTII Hieron. PL 29.52.25. Megaspores Apricot Yellow to Deep Chrome (pale to bright orange); 0.18- 0.25-0.55 mm. іп diameter, 44 spores measured. There is considerable variation in size of spores of this species, since one large spore may develop at the expense of three small spores within a megasporangium. The pattern of sculpture of the spores is generally similar. Містоѕротеѕ Deep Chrome (bright orange); 34-56 џи in diameter. SUBGROUP OF S. HANSENI.— The two species of this subgroup are allied on the basis of strongly dorsiventral stems and uniform leaves. The spores are distinctive for each species. Commissural face of megaspores without equatorial ring; strongly reticulate on outer face 24. S. SHELDONI Commissural face of megaspores with prominent equatorial ring; lightly rugose on outer face 5. S. HANSENI [Vor. 36 422 ANNALS OF THE MISSOURI BOTANICAL GARDEN 24. S. SHELDONI Maxon. Pl. 29, fig. 24. Megaspores Deep Chrome (bright orange) ; 0.34—0.36-0.44 mm. in diameter, 28 spores measured. Microspores Deep Chrome (bright orange); 34-45 м in diameter. 25. S. Нлмвемі Hieron. Pl. 29, fig. 25. Megaspores Lemon Chrome (lemon yellow) ; 0.29--0.36—0.42 mm. in diameter, 40 spores measured. Microspores Apricot Yellow (pale orange); 38-49 џ in diameter. SUBGROUP OF S. PARISHII.— Two members of this group are Mexican and are not treated here. Тһе mega- spores of S. eremophila are for the most part more prominently sculptured than the megaspores of S. arizonica. In both species a single, large megaspore and three small spores may be found within a sporangium. Commissural face of megaspore finely rugose- TUNE: vee ie marked ith equatorial ring; outer face coarsely reticulate to w М Commissural face of теразроге еу sharp, projecting гирае, prominent equatorial ring; outer face deeply rugose-reticulate 26. S. ARIZONICA 27. S. EREMOPHILA 26. S. ARIZONICA Maxon. Р]. 29, fig. 26. Megaspores Apricot Yellow (pale orange); 0.25—0.32—0.46 mm. in diameter, 36 spores measured. Some spores are more prominently marked than fig. 26b; some have an obscure equatorial ring. Microspores Deep Chrome (bright orange); 23- 38 p in diameter. 27. S. EREMOPHILA Maxon. Pl. 25, fig. 27. Megaspores Pinard Yellow (pale yellow); 0.23-0.34-0.50 mm. in diameter, 25 spores measured. Microspores Deep Chrome (bright orange); 38—53 и in diameter. SUBGENUS STACHYGYNANDRUM The species included in this study belonging to this subgenus are all char- acterized by strikingly distinct spores. Тһе microspores are as strongly marked as the megaspores. The microspores of S. apoda and S. Douglasii have a pebbled ap- pearance, those of S. armata are beset with papillae and those of S. pilifera have globules of a waxy substance adhering to them. SERIES DECUMBENTES.— Megaspores less than 0.22 mm. in diameter; deep yellow to orange; ob- e to nearly smooth scurely rugos 29. S. ARMATA Megaspores more than 0.36 mm. in diameter; straw-yellow or lighter with i r «i 1 28. S. DoucLasit Megaspores marked with жөніндегі thin, free rugae 30. S. APODA 28. S. Doucrasn (Hook. & Grev.) Spring. Pl. 30, fig. 28. Megaspores Straw Yellow; 0.36—0.35—0.40 mm. in diameter, 7 spores measured. Microspores Deep Chrome (bright orange) ; 23—41 џ in diameter. 29. S. ARMATA Baker. РІ, 30, fig. 29. Megaspores Deep Chrome (bright orange) ; 0.15-0.10-0.21 mm. in diameter, 15 spores measured. Microspores Flame Scarlet (red orange); 26-34 м in diameter. 1949] TRYON—SELAGINELLA SPORES 423 30. S. АРОрА (L.) Fern. Pl. 30, fig. 30. Megas pores Barium Yellow (yellow, green-tinged) ; 0.34—0.36 mm. in diameter, 7 spores measured. Microspores Flame Scarlet (red orange); 23-34 м in diameter. SERIES CIRCINATAE.— Megaspores spinulose Megaspores granular 31. S. LEPIDOPHYLLA (Hook. & Grev.) Spring. Pl. 30, fig. 31. Megaspores Deep Chrome (bright orange) ; 0.23-0.20-0.42 mm. in diameter, 23 spores measured. Microspores adhere in tetrads which are Flame Scarlet (red 22 S. LEPIDOPHYLLA . 5, PILIFERA orange). 32. S. PILIFERA А. Вг. Ба, 20-10 За. Megaspores Straw Yellow to Pinard Yellow (pale orange); 0.25-0.29 mm. in diameter, 11 spores measured. Some spores have an equatorial ring. — Microspores Flame Scarlet (red orange); 26-38 p in diameter. BIBLIOGRAPHY Baker, J. G. 154525 Ep of the 22 allies. London roun, M. (1938). Index t rth American ferns. Orleans, Сізш sen, R. Т. (1946). Selgin armies Euselaginella, in хен cas United States. Am. our. 36:65- Fern J Hieronymus, G. (1900). Белеги species novae vel non satis cognitae. I. Hedwigia 39:290- Lyon, F. `M. 901). A study of the болға and gametophytes of Selaginella apus and Selaginella puis. "Bo. Gaz. 32:124-141 -------, (1905). The spore coats d Sda, Ibid. 40:285—29 Maxon, W. R. (1918). A new Selaginella from Oklahoma ES "Tos. Proc. Biol. Soc. Wash. 31:171-172. ‚ (1920). New Selaginellas from the western United States. Smithson. Misc. Coll. 725:1-10. , (1921). Notes оп American e£. Ug Am. dex F 11:34-39. Е ‚ (4923). Notes оп American fer pee Ibid. Я Morton, C. V. eo Our sot која resurrection plant ^ d. 9:1 Reeve, R. M. (1935). De Du of the genus Selaginella in peat эү ud north eastern United St hodora 37:3 45. Ridgway, Robert (1912). cé теи and color nomenclature. Washington, D. = Selling, О. H. (1946). Studies in Hawaiian pollen statistics. Pt. I. Bishop Mus. eg nee bl. Underwood, L. | (1898). pol de rupestri and its allies. Bull. 27 Bot. Club 25: ipfa Gs: ------, (19 А changed conception of species. Fern Bull. 9:49-53. Van Eseltine, с Р. (1918). The allies ЊЕ Selaginella rupestris in x: southeastern United States. ontr. U. 5. Nat. Herb. 20:159-172. Ee J; 54 АНС. x. mr Lycopodiinae in Verdoorn, Manual of Pteridology, pp. 3—506. Weatherby, г (1943а). Тһе group of жа Parisbii. Am. Fern Jour. 33:113- — 3b). E dice in Johnston, I. Plants of northern Mexico, I. rd Arn. 2 1 о: Тһе group of Selaginella oregana in Ero 271 Ibid. 25:407-419. -------, (1946). А new Selaginella from western Texas. n Jour. 36:51—53 Wherry, E. Т. (1936). Observations on Selaginella ED [on 45 А. Бог. Club 1:65—69. [Vor. 36, 1949] 424 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE In this and the following plates, which are reproductions of photographs, o magnifi- cation for the megaspores is 85 diameters and for the microspores 450 diame PLATE 23 Fig S. SELAGINOIDES: Butters 9 Holway, 4. 1905 (Univ. Міпп.). la. Сот- Қ Тыны, ы of megaspore. 1b. Outer face of тер Fig. 2. S. rortipma: Donnell-Smith, South cat 1881 (Chicago Nat. Hist. Mus.). 2a. Commissural face of megaspore. 2b. Outer face of a moderately marked ге. 2 Sherwoodii: Donnell-Smitb, North Carolina, 1882 (Mo. Bot. Gard.) 24. Commissural face of megaspore. 2b. Outer face of a strongly marked megaspore. 2c. Microspore. ANN. Мо. Вот. Garp., Vor. 36, 1949 PLATE 23 S. Sherwoodi TRYON —SELAGINELLA SPORES Ann. Mo. Вот, Garb., Vor. 36, 1949 PLATE 24 S. ARENICOLA TRYON —SELAGINELLA SPORES [Vor. 36, 1949] TRYON—SELAGINELLA SPORES 425 EXPLANATION OF PLATE PLATE 24 S. RUPINCOLA: 3a. Commissural face of megaspore, Goodding, Arizona, 1921 т Nat. Hist. Mus.). 3b. Commissural face of prominently sculptured megaspore, Wooton, New vae 1907 (Univ. Minn.). 3c. Outer face of megaspore, same specimen as from the Fig. 5. S. BIGELOVII: Commissural es of megaspore Мис ие prominent equa- torial фы Козе 34405, California 1932 (Chicago Nat. Hist. Б. megaspore, Grant & Wheeler, California 1904 (Univ. Minn.). same specimen as 5а. uter face of Es bilcoppore from the Fig. S. RippELLI: Cory 41000, Texas, 1943 йа; НегЬ.). 7а. Commissural face of megaspore. 7b. Outer face of a lightly marked megaspore. 7c. Microspore 8. S. ARENICOLA: Deam 63023, Florida, 1946 (Mo. Bot. Gard.). 8a. Commis- sural hos of megaspore. 8b. Outer face of megaspore. 8c. Microspore. [Vor. 36, 1949] 426 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 25 S. funiformis: O'Neill 7580, bg sie 1933 ғ Мас. Hist. Mus.). ба. Суан face of megaspore. Sb. Outer face of megaspor S. ACANTHONOTA: 9a. Commissural face of a lightly marked megaspore, ры 5 McVaugh 3101, Georgia, 1938 (Мо. Bot. Gard. 9b. Commissural face of a prom nently marked megaspore, Harper 1057, Georgia, 1903 (Mo. Bot. Gard.). 9c. Outer ho of megaspore, from same specimen as . S. floridana: Nasb 1449, Florida, 1894, ичи. хера Nat. Hist. Миз.). да. ыы face of megaspore. 00. Outer face of meg Fig. 10. S. orEGANA: Piper, Washington, 1893 (Univ. Minn.). 10а, Commissural face of megaspore. 10b. Outer face of megaspore. 10с. Microspore. ANN. Мо. Bor. Garb., Vor. 36, 1949 PLATE 25 S. OREGANA TRYON —SELAGINELLA SPORES ANN. Mo. Вот. Garb., Vor. 36, 1949 PLATE 26 S. WATSONI TRYON —SELAGINELLA SPORES [Vor. 36, 1949] TRYON-—SELAGINELLA SPORES 427 EXPLANATION OF PLATE PLATE 26 S. UNDERWOODII: 11а. Commissural face of megaspore (var. dolichotricha Weath.), pt: alfe 001, New Mexico ds Bot. Gard.). 11b. Outer face of megaspore, Clark 8968, Arizona, 1940 (Mo. Bot. Gard.). Fig. 12. S. мотіса: Drouet, Richards 9 Rubinstein 4007, Colorado, 1941 (Mo. Bot. Gard.). 12a. Commissural face of a lightly marked spore without an equatorial ring. 12b. Outer face of megaspore. 12c. Microspore. S. CINERASCENS: Abrams 3399, California, 1903 сн Мас. Hist. Mus.). 13а. Сой face of megaspore. 13b. Outer face of megaspor Fig. 14. S. Watsoni: Maguire, Hobson & Maguire 14720, Utah, 1936 (U. S. Nat. Herb.). 14a. Commissural face of a prominently marked megaspor е with unusual Но а commissural ridges. 14b. Outer face of теразроге. 14c. Outer йн of megaspore with large areolae. 14d. Microspore. [Vor. 36, 1949] 428 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 27 Fig. 15. S. STANDLEYI: Wherry, Colorado (U. S. Nat. Herb. 1730847). 15а. Com- missural face of a lightly marked megaspore. 15b. Outer face of a lightly marked mega- spore. Fig. 16. S. P uai de Munz & Harwood 3769, pices на Isotype жай садо Nat. Hist. Mus.). ‚ Commissural face of a lightly e aspore. microspore attached near а of one commissural der 16 6b. Cute M pi Во Fig. 17. S. WALLACEI: Constance 9 Rollins 008, Washington, 1935 (Univ. Minn.). 17a. Commissural face of a prominently marked megaspore with prominent equatorial ring. 17b. Outer face of a prominently marked megaspor Fig. 1 S. 5ГВЕЕТСА: Porsild 150, Alaska, 1926 (Gray Herb.). 18a. Commissural i of а megaspore with unusually short commissural ridges. 18b. Outer 2. of megaspor ANN. Mo. Вот. Garp., Vor. 36, 1949 PLATE 27 S. STANDLEYI S. WALLACEI S. SIBIRICA TRYON —SELAGINELLA SPORES ANN. Mo. Bor. Garp., Vor. 36, 1949 PLATE 28 . ASPRELLA S. RUPESTRIS TRYON —SELAGINELLA SPORES [Vor. 36, 1949] TRYON—SELAGINELLA SPORES 429 EXPLANATION OF PLATE PLATE 28 Fig. 19. S. AsPRELLA: Johaston 1807, California, 1917, Paratype (U. S. Nat. Herb.). 19a. C Une а face of megaspore. 19b. Outer face of megaspore. 19c. Microspore. g. 20. S. ремзА: Rydberg & Bessey 3517, Montana, 1897, Isotype ic Nat. Hist. Mus.). 20a. Commissural face of megaspore. 20b. Outer face of prominently marked m de Fig. 21. S. RUPESTRIS: 21a. Commissural face of а megaspore with a single, straight commissural ridge: B. C. Taylor 1573, Wisconsin, 1892 (Univ. Minn.). 21b. Outer face of megaspore: eles 5106, SPRE” 1910 (Mo. Bot. Gard.). 21c. Dumb-bell mega- spore, from same specimen as 21 214. Microspore, Tryon 9 Tryon 5005, Wisconsin, 1947 (Mo. Bot. Gard.). Бір. 22. 5. SCOPULORUM: . Commissural face of megaspore, Sbaw 002, British Columbia, ipee dU es (Univ. m n.). 22b. Outer face of megaspore, Umbach 856, Montana, Isoparatype (Chicago Nat. Hist. Mus.). 430 [Vor. 36, 1949] ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 29 S. \Мвюенти: Reverchon, Texas, 1883 (U. S. Nat. Herb.). 3. 233. Commis- sural face of megaspore. 23b. Outer fa ce of megaspore. SHELDONI: Cory 30143, Texas, 1938 (Mo. Bot. Gard.). Fi 5. 24а. Commis- m > Ж of megaspore. 24b. Outer face of megaspore. | 24c. Microspore. Fig. 25. S. HANSENI: Heller 11802, California, 1915 хез Nat. Hist. Mus.) 25a. Commissural face of megaspore. 25b. Outer face of megaspor Fig. 26. S. ARIZONICA: Goodding, Arizona, 1921 (Chicago Nat. Hist. Mus.). 26a. Commissural face of megaspore. 26b. Outer face of a lightly marked megaspore. S. EREMOPHILA: Cook, California, 1921 (U. S. Nat. Herb. g. ). 27a. Com- missural fact of а megaspore with obscure equatorial ring. 27b. Outer face of megaspore. ANN. Мо. Вот. Ganp., Vor. 36, 1949 PLATE 29 S. HANSENI Š. ARIZONICA Š. EREMOPHILA TRYON —SELAGINELLA SPORES Ann. Mo. Вот, Сакр., Vor. 36, 1949 PLATE 30 S. Douctasit o S. ARMATA . PILIFERA со BE S. APODA S. LEPIDOPHYLLA TRYON —SELAGINELLA SPORES [Vor. 36, 1949] TRYON—SELAGINELLA SPORES 431 EXPLANATION OF PLATE PLATE 30 . Doucrasm: Peck, Oregon, 1914 (Мо. Bot. Gard.). 28a. Commissural re. Fig. 28b. Outer face of megaspore. 28c. Microspore face of Moin Fig. 29. 5. ARMATA: Small 9 Carter, Florida, 1903 (Univ. Minn.). 29a. Commissural face of megaspore. 29b. Outer face of megaspore. 29c. Microspore. 30a. Commissural face of -— Ó Rutb, Pennsylvania, 1890 Fig. 30. S. APODA: еп as 30a. 30с. Micro- (Univ. Minn.). 30b. Outer face of megaspore, from same specim spore, E. L. Nielson 1564, м Иб: 1932 (Univ. Minn.). 31. S. LEPIDoPHYLLA: E. J. Palmer 11366, Texas, 1917 (Мо. Bos Gard.). 31a. Е И face of megaspore. 31b. Outer face of megaspore. Fig. 32. S. РИЛЕЕВА: С. H. Mueller 8257, Texas (Mo. Bot. Gard.). 32a. Commis- sural ae of megaspore without equatorial ring. 32b. Outer face of megaspore. 32с. Microspore. THE CYTOLOGY OF PAPHIOPEDILUM MAUDIAE HORT.! HENRY A. McQUADE STATEMENT OF THE PROBLEM The hybrid Paphiopedilum Maudiae Hort. is the offspring of Paphiopedilum callosum var. Sanderae and P. Lawrenceanum var. Нуеапит. It has been used in relatively few crosses despite its quality, and its offspring have rarely, if ever, been able to exceed it. It was felt that an orchid of such superiority which produced little and often no seed should be examined cytologically to ascertain, if possible, the causes of its sterility. Accordingly, cytological examination was made of the parents and the hybrid and some observations were made on somatic tissues of cer- tain related species. I. PAPHIOPEDILUM Pfitz. Paphiopedilum was originally incorporated іп the genus Cypripedium described by Linnaeus. From Cypripedium were later extracted the three other genera of the Tribe CYPRIPEDILINAE as they are generally accepted today; the residue of the large genus continues under Linnaeus' original designation. The orchid-grower continues to refer to various species of Paphiopedilum as “Cypripedium,” and the confusion oftimes extends to the remaining genera. Linnaeus arrived at the name Cypripedium in an effort to latinize what he thought were the proper Greek words for the "Slipper of Venus." These are Kuzpis (Latin Veneris) and тобюу (calceus). The last, however, does not signify the Latin calceus, or shoe, but rather is the word for a small foot. Ascherson? changed the generic name to Cypripedilum (pedilum representing the latinization of the Greek equivalent of calceolus), and this was adopted by Pfitzer (1886) and published as such. Buser (1894) has concluded that both names represent the latinization of "very mediocre Greek," but that by virtue of priority Cyfri- pedium would be correct. Since Linnaeus had used Cypripedium consistently in both 'Species Plantarum' and 'Genera Plantarum' Pfitzer would have no right under present-day rules to suggest a chang Selenipedium was changed to а A Бу Pfitzer іп 1886 on the same basis that the change was made in Cypripedium. The genus was originally de- scribed as Selenepedium by Reichenbach filius (1859) who used this spelling con- sistently. Оп the basis of priority as determined by the present-day rules of nomenclature, Selenipedium is the proper expression. The use of Phragmopedilum rather than Pbragmopedium suggested by Rolfe (1896) is mandatory today since the genus was published originally as Pbragmo- — by Pfitzer. In the same manner Paphiopedilum (rather than Paphiopedium An investigation carried out at the Missouri Botanical Garden and submitted as a thesis in partial ын of the DE ki the degree of Doctor of Philosophy in the Henry Shaw School of Botany of Washington Univer ? Ascherson, P. Flora d. даја Depdedbücg: p. 700, in noía. 1864. [Vor. 36 434 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN of Kerchove, 1894, or Rolfe, 1896) takes precedence. The regularity that Rolfe sought for the endings of the generic names within the group, and the change sought by Pfitzer on the basis of derivation are tempered by priority. The fol- lowing keys to the genera, with necessary modification to generic names, are taken from Rolfe and Pfitzer in that order and are offered for comparison. ROLFE’S KEY TO ORCHIDEAE (ORCHIDACEAE OF PFITZER) Suborder DIANDRAE (PLEONANDRAE of Pfitz ribe CypRIPIDIEAE (CYPRIPEDILINAE r elled with axile placentas; sepals valva Leaves plicate; perianth persistent; seeds ые ЕН Selenipedium Rchb. f. [3 species in South America and Brazil] Leaves conduplicate; perianth deciduous; seeds fusiform... ии (Pfitz.) Rolfe [11 tropical American species] Ovary 1- = with parietal si = fusiform. Leav “. а th persistent; sepals valvate Cypripedium 1. [ab widely йер воде іп Europe, ннн Asia, e то rth America Leaves ae до тады perianth deciduous; sepals їтїһгїсаїе...................... hiopedilum Pfitz. [46 species in tropical Asia] PFITZER’S KEY TO ORCHIDACEAE PLEONANDRAE Tribe CYPRIPEDILINAE Vernation of the leaves convolute, perigonium marcescent, persisting in the a. Ovary trilocular, seed almost ee with я outer coat.. анат Rchb. f. gile vary unilocular, seed elongate with fragile coat... Cypripedium L. B. Vernation = leaves rs ind Бе rayuan ous. . Ovary trilocular, vernat of sepals valvate, margin of slipper- shaped labellum 54; involute ог Нана Pfitz.) Rolfe b. Ovary unilocular, vernation of sepals venne margin of slipper- aped abellum simple and straight, lightly incurved or recurved edilum Pfitz. Paphiopedilum callosum (Rchb. f.) Pfitz. and its var. eun Hort.— Paphiopedilum callosum was described as Cypripedium callosum by Н. G. Reich- enbach filius (1896). It was brought from Siam by Alexandre Regnier, of Paris, іп 1885, and very little time elapsed between its importation and the possession of a plant by the Sander's Company of St. Albans in England. It was early recognized as a graceful and vigorous form and was soon in culture wherever there were orchid enthusiasts. (See pl. 31.) The albino form, variety Sanderae, appeared in culture їп 1893 or 1894. Whether it flowered for the first time in England or France is a matter of debate; an anonymous article in the ‘Journal of Horticulture’ of 1912 says that it flowered for the first time in the autumn of 1893 in the collection of R. H. Measures, Esquire, at The Woodlands, Streatham. Be that as it may, in the Seventh Annual Show of the Royal Horticultural Society at the Inner Temple Gardens of May 23—25, 1894, it won a First Class Certificate when exhibited by Messrs. Sander and Sons of St. Albans. Both the Sander's plant and that of Measures appear to have come from the same source. For several years the variety Sanderae remained something of a rarity but came at length to be well represented in collections everywhere; its similarity to the 1949] McQuapE—Pabbiobedilum Maudiae нокт. 435 albino form of Lawrenceanum was commented upon but it was considered by most to be more graceful. Paphiopedilum Lawrenceanum (Rchb. f.) Pfitz. and its var. Hyeanum (Rchb. f.) —Papbiopedilum Lawrenceanum (see pl. 31) was described as Cypri- pedium Lawrenceanum by H. G. Reichenbach filius in 1878. The plant was found on the bank of the Lawas River by Mr. F. W. Burbidge while on a collecting trip in North Borneo in the service of the Veitch Company. Іп December, 1878, it was brought to flower in the Veitch greenhouses. Dr. Reichenbach named it for Sir Trevor Lawrence, Baronet, M. P., a contemporary orchid enthusiast whose col- lection is described as "being of exceptional richness and beauty" (Reichenbach, 1878). The albino form appeared spontaneously in the Linden cultures of ordinary Lawrenceanum which had been imported by Messrs. Lowe & Co. of Clapton. Mr. Jules Hye of Ghent, a fancier, was most eager to secure such a rarity for his own ollection, and apparently Mr. Linden, after having sold it to him, decided to name the plant for his customer. It was shown in the April 27, 1886, meeting of the Royal Horticultural Society by two exhibitors, the Compagnie Continetal, and R. B. White, Esquire, of Earlsfield, Surrey. The new form was at first called Cypripedium H yeanum, by various authors, but it was Reichenbach who first regarded it as a variety.! His description ap- peared іп 'Lindenia, of 1885, changing the nomenclature from Cypripedium Hyeanum (L.) Lind. & Rod. to Cypripedium Lawrenceanum Rchb. f. var. Н yeanum Rchb. f. x Paphiopedilum Maudiae Hort.—The hybrid is a cross between Р. callosum var. Sanderae and P. Lawrenceanum хат. Hyeanum, with the latter as the seed parent (Wilson, 1923). It flowered for the first time in the early fall of 1900 in the houses of Messrs. Charlesworth & Co. of Heaton, Bradford, England, and on September 27 of that year took a First Class Certificate at the Manchester Orchid Society exhibition. A similar award was given when it was shown by Mr. G. W. Law-Schofield at the Royal Horticultural Society on July 30, 1901. Since that time it has been highly prized as a show plant, decorative plant, and commercial orchid. (See pl. 31). Just who is to be credited with making the cross is a matter of conjecture. It has been reported that it was made in the Charlesworth establishment and again as having been made by two amateur growers, Major Mason and Mr. Charles Winn. In any event, the pot containing the seeds came into the possession of the Charles- por (Gard. Chron. 18:748) states: “Мг. Jules Hye Leysen, of Gand, Coupure 8, was so very kind as to send me the only flower we Europeans have had the орбіту of seeing . I immediately “hough it might be an albino of Sir Trevor мор Cypripediu d ат рег- suaded we must regard it vi Cypripedium — — variety Hyeanum, p name hae g been given, otherwise we might better call it “Mons. Hye Leysen’s individu ait. Тһе history is š simply, that it was found at she old arene S E at Linden aa a mass of typical С. Lawrenceanum.’ [Vor. 36 436 ANNALS OF THE MISSOURI BOTANICAL GARDEN worth Company which is recorded as having produced the first flower. Тһе plant was named for the daughter of Mr. Baguley, the Charlesworth foreman. Тһе cross, of course, was inevitable since the parent forms were collector's items valued at more than 100 guineas a plant, a price which indicated the value then placed on such desirable forms. There is a peculiarity in the nomenclature of the hybrid Maudiae which should be mentioned. If the specific, rather than the varietal, forms of the parent species had been crossed, the name Mazdiae would have denoted a colored form. The albino form would then have to be designated by some such additional term as albescens. Ав it happened, X Paphiopedilum Maudiae denotes the albino form, and the colored hybrid which appeared six years later had to be designated P. Maudiae var. coloratum. From the albino form, several selections have been made. Among them are P. Maudiae var. magnificum, P. Maudiae WESTONBIRT variety, P. Maudiae DELL variety, and P. Maudiae BANK House variety. The interspecific hybrid Paphiopedilum Maudiae exhibits hybrid vigor in both growth and flowering, for it is sturdier than either of its parents and flowers more frequently. There are certain striking characters in the hybrid which are readily seen in one parent or the other. The following table compares some of these char- acters in parents and offspring: P. Lawrenceanum P. callosum x Maudiae Leaves Tessellated Marbled Tessellated (a) Horizontal Pendent or drooping Intermediate (if otherwise it is closer to callosum) Petals (b) Tip acute Acuminate Acute (с) Warted on upper and | Warted on upper edge only | Warted on upper edge only lower edges | (а) Warted ас base Not warted ас base Not warted at base Sepals (b) Somewhat ciliate Less ciliate As in callosum In order to clarify the positions of the species within the genus and the appear- ances of the entities involved, the following outline is added. The system is that of Pfitzer which is at present accepted but the only varieties mentioned are those bearing on this problem. All but the descriptions of Maudiae are from Sander’s Orchid Guide of 1927. Genus Paphiopedilum Subgenus: ОтоРЕриим. (There аге 3 subgenera. The others are BRACHYPETALUM and ANATOPEDILUM Section: PHACOPETALUM. (There are 11 sections of subgenus OTOPEDILUM) Species: 1. P. Curtisii 6. P. callosum . P. ciliolare P. callosum var. Sanderae 3. P. superbiens 7. P. Lawrenceanum 4. P. Argus P. Lawrenceanum var. Нусапит 5. P. barbatum 1949] McQuape—Paphiopedilum Maudiae новт. 437 P. callosum: Foliage marbled. Scape 1 to 2 feet, 1- or 2-flowered. Flowers large, variable, dorsal sepal white, shaded or green at base, striped and often flushed with dark crimson, petals pale green with rose-purple apices, warted and ciliated at upper margins, lip brown-purple. Winter to summer. Var. Sanderae: Dorsal sepal white with apple-green stripes and radiating dorsal veins, petals pale green, whitish on the upper edges, lip pale green. Winter to summer. P. Lawrenceanum: Foliage brightly tessellated. Ѕсареѕ 1-2 feet high. Flowers large, bold, dorsal sepal white with purple-red stripes, greenish at base, petals hori- zontal, greenish with purple shading, chiefly at the tips and margins, black-warted on both edges. Lip dull purple tinted with brown, variable. Summer. Var. Hyeanum: Dorsal sepal pure white with green stripes, petals yellowish green, lip greenish. Summer. X P. Maudiae (Rolfe, 1900): An albino. Dorsal вера! very broad and rounded, white closely veined with bright green. Petals somewhat falcate, light green with darker veins and a few darker warts on the upper margins. Lip and scape light green. Flowering time various. II. Stratus оғ Paphiopedilum Maudiae as a PARENT PLANT The hybrid P. Maudiae combines the winter-flowering habit of one parent with the summer-flowering habit of the other and is one of a small group of albinos within its genus. It is, furthermore, an exceptionally beautiful plant. These features would seem to make it a highly desirable parent yet it has not been used to great extent perhaps because of its sterility. When Maudiae has been used in what would appear to be advantageous crosses there has been little seed produced and the extended flowering period characteristic of Maudiae has, as a rule, not been obtained in the offspring. Albino-flowered forms are rare in Paphiopedilum, the great majority of species as yet not having produced any. Although there is a considerable number of ad- vanced! albino hybrids today, they may be traced back to one or more of these basic forms (Black, 1933; Cooper, 1946). 1. P. bellatulum var. album and P. niveum (subgenus BRACHYPETALUM ). These аге the only species in which true whites are formed. Even so P. niveum shows a faint peppering of purple when viewed closely. 2. P. Lawrenceanum (var. Hyeanum), P. callosum (var. Sanderae), and P. Curtissii (all in sect. PHAcoPETALUM) have albino forms in which the white is striped and shaded with green. These belong to the “mottled group" characterized by marbled or tessellated foliage and petals ciliated and warted. 3. Р. Charlesworthii var. album (sect. NEUROPETALUM) which may no longer be in cultivation. lTn this study the expression primary hybrid is construed to mean species or variety hybrid. An oen nord is one in which one or both parents is of hybrid origin. - A. Sta [Vor. 36 438 ANNALS OF THE MISSOURI BOTANICAL GARDEN 4. P. insigne vars. Sanderae and Sanderianum (sect. NEUROPETALUM) 15 a soft yellow of variable depth according to the form. It is not striped, and white appears in the dorsal sepal. It is the chief member of the "insigne group." Some success has been achieved by combining Maudiae with other albino varie- ties (Wilson, 1923): Ногреми (Maudiae X callosum var. Sanderae).—Exhibited in 1909. ALMA GavaERT (Maudiae X Lawrenceanum var. Hyeanum).—Flowered in 1911. WARDEN (Maudiae X Holdenii).—Flowered 1920. EMERALD (Maudiae X Curtissii var. Sanderae) .—Exhibited in 1920. ENCHANTRESs (ALMA Слулект X Curtissii var. Sanderae).—Exhibited in 1921. Козетти (Maudiae >< insigne var. Sanderianum).—Exhibited in 1908. Before entering into a discussion of the crosses in which P. Maudiae and its parents have been involved, a further breakdown of the genus is necessary. The key given is taken from Pfitzer and may be used in conjunction with Charts I, II and III and their appendices: KEY TO THE SUBGENERA OF PAPHIOPEDILUM A. Labellum a Piper, exauriculate (without ears or auricles), bag-shaped, having a very short claw at the for the narrow inwardly rolled үз, petals very broadly elliptical or almost more ess purple RE ed бен е elliptical, tessellated above, or bifl В. 22 а slipper, exauriculate, inclining downward, having а claw of alm to that of the slipper on the front of the simple non- ттш margin; petals ea strap-shaped, green on both sides, scape several-flower Subgenus 2. бајт ын Um Pfitz. C. Labellum a slipper, auriculate, bag-shaped, having a claw almost as long as the pw at the fore of the шари, non-involute margin; petals longa di varying, scap several or single c OTOPEDILUM ба. KEY ТО THE SECTIONS OF THE SUBGENUS ANATOPEDILUM A. t striate between simple curving nerves; petals deflexed (bent outwardly), elongate with ciliate margins; бк зн» суйп ia ker the lower ascending part = зый hes the upper үнүн part glabrous “н with two lobed арех Sec TOPEDILUM Pfitz. B. Sepals striate between simple curving nerves; petals deflexed, hin nd margins adorned with hairy warts; ны directed upward апа fo iic arched ен Ler- below the concave margin Sect. ConvYoPEDILUM Pfitz. C. -— with extra striate curving nerves betw enuous reticulate nerves; шүлен curved, elongate, margin almost naked; анте like that °. the preceding section. PRENIPEDILUM Pfitz KEY TO THE SECTIONS OF THE SUBGENUS OTOPEDILUM minode inversely heart- shaped, forked, enlarged on the back by a basal boss, acute, pilose. rame ves а Же uniform in tint, almost erect, green. Scapes several-flowered, flowering multan a wi narrow, hanging, - twisted оп the lower margin, стандын “о hairy warts, yalin minutely spatulate at the apex. Leaves very narrow with margin ts MYSTROPETALUM Pfitz. Petals narrow from linear bae си» the apex, вино expanding into а blade extremely divergent, nearly twisted, margin without warts. Leaves very scolis yellow-margined S PARDALOPETALUM Hallier ! Bracbypetalum is not further divided to sections but is divided directly into species. аа! — - 1949] McQuape—Paphiopedilum Maudiae нокт. 439 В. уш г forked, almost with an undivided boss at the back. Leaves broadly ped, rved, uniform in tint, green or glaucous. Scape several-flowered, bracts = алаң flow won successively Sect. COCHLOPETALUM Hallier . Staminode nearly ania square, heart-shaped or almost heart-shaped. Sepals with tenuous reticulate ner дари simple curved nerves. Leaves green, uniform іп tint, scapes аа бе уегу Meran floral. a minode nearly * i enlarged on the back by 3 low, slightly prominent ian ; petals spatulate Sect. STICTOPETALUM 2. b. Staminode almost reverse heart- shaped, blunt, enlarged on the conv plan back by one large central boss; petals more or less inris into a blade © to “map; the apex Sect. NEUROP M Hallier c. Staminode heart- shaped, furrowed at the back, slightly LE pal реа Sect. Тн ALUM ome d. Staminode almost round, split on the posterior with gi ек o чоң tals oblong with wavy margins Sect АТОРЕТАТ, da " Hallier . Staminode half-moon-shaped, the 14 part equally three- рей ES with Min curving nerves uniting reticularly at the apex; petals strongly sigmoid, ben wards at the apex, erect. Leaves green, бена іп tint, зсаре unifloral Sect. CERATOPETALUM oe е half-moon- ping with double horse-shoe-shaped E" and the е arly rhombic boss. Sepals net-veined or simply veined, p clearly expan idi ing into a blade NOME e apex. Leaves more or less t e gis (checkered), scape unifloral ect. SPATHOPETALUM Pfitz. . Staminode moon-shaped or semi-rounded, without а boss; sepals simply iut petals not expanding or hardly expanding =ч ; за toward the ex ma aves clearly tessellate, scape unifloral, rarely etal margins naked or haired with pu сне | p^ UNE т ect. BLEPHAROPETALUM Pfitz. b. Petal margins spotted or por with prominent warts provided with small brush-like hair tuf Sect. PHACOPETALUM Pfitz. Maudiae has been crossed to members of nine of the eleven sections of the sub- genus OTOPEDILUM or to individuals whose antecedents lie within these sections. (It has not been crossed to individuals belonging to sections PARDALOPETALUM or MYSTROPETALUM.) The section contributing the most germ plasm to individuals crossed with Maudiae appears to be NEUROPETALUM; PHACOPETALUM, the section from which Maudiae stems, and CvMATOPETALUM contribute somewhat less. Individuals emanating from the NEUROPETALUM group were used freely, probably in efforts to combine Maudiae's everblooming character with some of the desirable "insigne" characters. Only one cross has been recorded with a BRACHYPETALUM species (niveum) and one with ANATOPEDILUM (Rothschildianum in sect. СомА- TOPEDILUM). Of the 35 crosses in which Maudiae has been used (Sander's List, 1946), 11 have been with species, 3 with primary hybrids, and 21 with advanced hybrids. (See Chart I). Paphiopedilum callosum has been crossed to members of nine out of the eleven sections of the subgenus ОТОРЕрпим, or individuals having their antecedents therein. (It has not been crossed to individuals with a background in Mvsrno- PETALUM ог PARDALOPETALUM). PHACOPETALUM, to which callosum and Law- renceanum belong, is the section contributing the greatest number of individuals to these crosses with NEUROPETALUM and others contributing lesser numbers. It has been crossed to all of the species within the subgenus BRAcHYPETALUM and with sections CORYOPEDILUM and PRENIPEDILUM of the subgenus ANATOPEDILUM. [Vor. 36 440 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN Of its 55 crosses, 29 have been with species, 13 with primary hybrids, and 13 with advanced hybrids. (See Chart II and Appendix). Paphiopedilum Lawrenceanum has been crossed to representatives of all of the eleven sections of OroPEDILUM. Again, PHACOPETALUM and NEUROPETALUM contribute the most germ plasm to the individuals used in these crosses, with other sections contributing lesser amounts. It has been crossed with all the Вкасну- PETALUM species and with individuals belonging to two of the sections of the sub- genus ANATOPEDILUM (Gonatopedilum and Coryopedilum). Of the 62 crosses in which Lawrenceanum has been involved, 30 have been with species, 17 with primary hybrids, 15 with advanced hybrids. (See Chart Ш and Appendix). The only reasonably complete list of orchid hybrids available is Sander's 'Com- plete List of Orchid Hybrids’ (1946). This lists Maudiae as having been used in 35 crosses, Lawrenceanum in 62, and callosum іп 55. Of the 35 offspring of Maudiae only 7 have been used to produce 41 named offspring of their own. Twenty Lawrenceanum offspring have been used in 123 crosses, and 15 callosum offspring in 74, with Maudiae being the most productive offspring in either case. It must be remembered, however, that Sander's List does not necessarily present a true picture of sterility or fertility. It simply lists the crosses giving rise to named and reg- istered varieties and gives no account whatever of the crosses made wherein the grower considered the result not worth saving. It is thought, however, that most of the early crosses were named and registered since even the poor ones were regarded as having botanical interest. That both Lawrenceanum and callosum were crossed to more species than Maudiae is to be expected since they are older and have been known to horticul- turists for a much longer time. It is reasonable to assume from the record that both of these species cross freely with other members of the genus. Maudiae ap- pears to be slightly restricted, in comparison to its parents, in the number of sections со which it has been crossed. Іп part, this restriction тау simply be a reflection of the horticulturist's discrimination; crosses made with callosum апа Lawrenceanum that had proved inferior were probably not repeated with Maudiae. Also it is to Бе noted that many of the individuals to which Mawdiae has been crossed have been advanced hybrids, and this has probably contributed to a lack of seed set. Ас any rate, hybridizers have not been able to obtain sufficient seed when using Maudiae as a parent to combine its undoubted qualities with those of other parents; large numbers of crosses must be made to obtain a small amount of seed. Ш. AwNatysis or Roor-Tips An attempt was made to analyze the chromosome complements of P. callosum, P. Lawrenceanum (the albino forms were not available), and several forms of Ж Maudiae. For this purpose root-tip smears were used. It was felt that in addition to observation with the microscope and scrutiny of camera-lucida drawings the con- struction of ideograms would be fruitful. The value of ideograms in this case will be discussed in a following paragraph. 1949] McQuape—Paphiopedilum Maudiae нокт. 441 Root-tips were killed and fixed, after quartering, in a solution containing six parts of absolute alcohol to three parts of chloroform to one part of glacial acetic acid. It was necessary to leave the tips in the killing and fixing fluid for at least fifteen minutes although they were often, for reasons of necessity, allowed to re- main at least an hour. Tips that could not be squashed and stained immediately after removal from the killing and fixing fluid were kept in an aqueous solution of 45 per cent glacial acetic acid. The first smears were made from tips of P. callosum. Aceto-orcein as well as aceto-lacmoid was used as a stain. After completing observations on P. callosum aceto-lacmoid only was used in studying P. Lawrenceanum and P. Maudiae. The squash procedure used was that outlined by Mehlquist (1947). It was found later in the investigation that the Feulgen technique outlined by Meyer (1943) gave excellent results generally but that for studies involving the centromere region aceto-lacmoid was the best agent when used on root tips that had not been in the Carnoy’s fluid for more than fifteen minutes. lin | ІШ Тех 1. P. callosum. Comparative Text-fig. 2. P. Ti i Compara- ied x chromosomes tive length of chromos The construction of ideograms was based upon camera-lucida drawings of mid- metaphase cells so flattened that all parts of a chromosome were in one plane. Ten such plates were obtained for P. callosum but only four could be found for Law- renceanum and five for XX Maudiae. It was the original plan to study ten of each, but, although many root-tips were examined, only this small number was found fit for use. Indeed, it was often difficult to obtain tips showing any divisions at all. On the basis of such small numbers the ideograms are presented here only as indi- cators and not as final pictures of chromosome complements. Haploid ideograms of P. callosum and P. Lawrenceanum based upon excellently flattened anaphase cells clearly indicating centromere position are presented. These were not used in the preparation of the final ideograms showing length relationships. After preparation of camera-lucida drawings, the chromosomes were paired ас- cording to length, position of centromere, and placement of secondary constric- tions. The basic ideogram thus consisted of the diploid number. For the fina ideogram the lengths of the two members of a pair were averaged and the averages of all corresponding pairs were in turn averaged. The final ideogram therefore [Vor. 36 442 ANNALS OF THE MISSOURI BOTANICAL GARDEN shows the haploid number. Тһе haploid ideograms of P. callosum and P. Law- renceanum are fundamentally alike. Both show a noticeably long chromosome number 1. Тһе difference in length between chromosomes 1 and 2 is less in P. Lawrenceanum than in the other. Aside from chromosome 1 all chromosomes show a very close intergradation in size. (See text-figs. 1 and 2.) Chromosome counts in various "horticultural Cypripediums," including P. callosum, P. Lawrenceanum, and the interspecific hybrid Maudiae, have been made by Mehlquist (1947a), Duncan (1945, 1947), and others. Some of these counts were verified during this investigation: P. callosum has a diploid number of 32; P. Lawrenceanum, 36; X Maudiae, 34 (see pl. 32); P. barbatum, 38; P. Curtissii, 36; and P. superbiens, 38. Paphiopedilum callosum, with а 2n number of 32, was analyzed first and ideo- grams were prepared. All but a few of the sixteen pairs of chromosomes have no identifying morphological features and exhibit a subtle intergradation in lengths. The following chromosomes are best recognized at sight: l. The longest pair has median centromeres; the secondary constriction is about midway on one arm. 2. А pair intermediate in length often appears to be composed of three pieces of chromatin to each individual. An end piece is a satellite. 3. One pair, longer than average, each with one ball type end, appears to have subterminal centromeres. к Шалы 7 7 8 9 m" и 42 4 n «T 5 „ 48 аа ------- - 3. Diagrammatic representation of the chromosomes of P. ee (L) an P. callosum (c), based on anaphases in root- tips which clearly disclosed the position of the centro- meres. Chromosomes arr данын according to length. Тһе numbers аге merely for pni and do not signify that chromosomes voii di м same numbers аге alike. The lines соппестіп c and L — suggest p mech homologue ү! ond secondary constriction is оп а chromosome of intermediate length with subterminal centromere ріш опе species and оп a chromosome with interstitial centromere іп the other. The соп- striction is not shown because vius positions of these ance are uncertain 1949] McQuave—Paphiopedilum Maudiae нокт. 445 4. One, or possibly two, pairs shorter than the average, with one ball type end, appears to have subterminal centromeres. The remaining chromosomes are so alike morphologically and vary so little in length that they cannot be identified as individuals upon observation. However, it can be shown that within the callosum complement there are six terminal or subterminal pairs and ten pairs in which the centromeres are interstitial. (See text-fig. 3.) The chromosomes of P. Lawrenceanum, by virtue of their similarity, remain as cryptic as those of P. callosum. Неге the longest pair is readily identifed by having a median, or close to median, centromere and is seen to possess a secondary constriction. Even the smallest chromosomes of Lawrenceamum cannot be dis- tinguished readily because of their great similarity in length to the next smallest pair. Thus, aside from the longest pair, the remaining seventeen pairs verge closely оп one another in size. The difference іп the chromosome numbers of P. callosum and P. Lawrenceanum appears to be in an increase of small chromosomes in the complement of P. Lawrenceanum. It can be shown that in P. Lawrenceanum there are ten pairs of terminal or subterminal chromosomes and eight pairs in which the centromeres are interstitial (See text-fig. 3.) The chromosomes of the interspecific hybrid Maudiae presented the same morphological anonymity as those of the parental forms. Again the long pair with median centromere and clearly defined secondary constriction was readily noted. There are two pairs longer than the average with centromeres between the median and subterminal points, and two pairs of average length with median centromeres but these are not often readily discernible. The satellited pair of intermediate length seen іп callosum was seen in Maudiae and presented an interesting situation. In regard to this chromosome, Maudiae var. magnificum is visibly an inversion heterozygote. A cell was found in which it was apparent that half of one of the chromosomes of this pair is inverted. Тһе in- verted half bears the centromere, and since the centromere is subterminal in one, it becomes interstitial in the other. The 2n chromosome configuration of Maudiae should consist of 10 chromo- somes with terminal or subterminal centromeres and 8 with interstitial centromeres from the Lawrenceamum parent, while of the 16 chromosomes contributed by callosum 6 should be terminal or subterminal and 10 would be interstitial. Attempts to construct an ideogram showing the diploid number were not rewarding. It may be assumed from these attempts, however, that the close intergradation of chromo- some lengths which characterize the parents is characteristic of the offspring. The value of the ideograms in the analysis of these chromosome complements would appear to be at best only moderate. Ideograms can be used only as approxi- mations for various reasons: 1. The lengths of members of a chromosome pair are not always equal; the longest chromosomes in the basic ideograms, for example, rarely agree exactly in 5 y ag x [Vor. 36 444 ANNALS OF THE MISSOURI BOTANICAL GARDEN length. This may be a result of a squashing which leads to distortion not only in length but in other dimensions. 2. Lengths of chromosomes vary from metaphase to metaphase. 3. The length ratio of one pair to another may vary from one metaphase to another because of pressure applied in squashing. The order of 16, 17 or 18 pairs in an ideogram must undoubtedly differ to some degree when there are many of approximately the same lengths lacking dis- tinct morphological features and have been, in addition, subjected to pressure. 5. Centromeres may vary in size or become so small as to be apparently lacking. 6. Satellites vary in size as do also the portions of a multi-piece chromosome. 7. А chromosome may break under pressure not only at the centromere but at other regions. Thus a chromosome could appear to have a median centromere when it is actually of the subterminal type, or a median type, when broken in two, appear to be two subterminal chromosomes. Despite these reasons for not placing too much faith in the ideogram as an actual picture of the chromosome complement there was sufficient resemblance among the basic ideograms to predict with considerable accuracy the appearance of the final ideogram. Ideograms of the haploid numbers of various Paphiopedilum species published to date by Duncan and MacLeod (1948a, 1948b) include representatives of sub- genera BRACHYPETALUM and ANATOPEDILUM and of the sections of OTOPEDILUM except SPATHOPETALUM, BLEPHAROPETALUM and PHACOPETALUM. These, for the most part, indicate a chromosome number 1 recognizable by its length and possessing a median or near median centromere. Chromosome number 2 may ap- proximate or nearly approximate it in length but there is usually a difference. From chromosome number 2 on there is a close intergradation in length down to the smallest chromosome. In this they agree with ideograms published earlier by Francini (1931, 1932, 1934). Among the intermediate chromosomes there is one with a secondary constriction. These same generalities are true for the chromo- somes of Р. callosum, Р. Lawrenceanum, and Р. Maudiae, except that these three appear to have an additional secondary constriction on chromosome number 1. Only one species, P. exul, ideogrammed by Duncan and MacLeod, shows a sec- ondary constriction on number 1 and none on any other chromosome. (Two species ideogrammed by Duncan and MacLeod exhibit a secondary constriction on number 2 and on no other chromosomes. These are P. Druryi and P. Spicerianum.) The following list presents the 2n chromosome numbers to date of species within the genus: 1949] McQuapE—Pabbiobedilum Maudiae нокт. 445 CHROMOSOME NUMBERS OF SPECIES OF P UM Chromosome Authority* Date umbers Subgenus BRAcHYPETALUM НаШег P. bellatulum (Reichb. f.) Pfitz. si Le БИ P. concolor (Batem.) Pfitz. 26 D&ML '48 P. niveum (Reichb. f.) Pfitz. 26 GM bid 26 D 47 va ° 26 D 47 P. Delanatii Guill. 26 GM 47 Subgenus ANATOPEDILUM Pfitz. Sect. GONATOPEDILUM Pfitz. Ет: ; 26,28 D 747 P. Rothschildianum (Reichb. f.) Pfitz. 26 D&ML 49 Sect. CoRYOPEDILUM Раст. t 28 D '47 P. praestans (Reichb. f.) Pfitz. 28 D&ML 49 P. philippinense (Reichb. f.) Pfitz. 26 D&ML 749 Sect. PRENIPEDILUM Pfitz. P. Stonei (Hook. f.) Pfitz. 26 D '47 Mary REGINAE Hort. 26 D&ML '49 Subgenus OroPEDILUM Pftz. Sect. MysTROPETALUM Pfitz. P. Parishit (Reichb. f.) Pfitz. > To) MS Sect. PARDALOPETALUM НаШег 23 Ç 26 D 47 P. Lowii (Lindl.) Pfitz. 26 D&ML 49 : А 26 р 47 P. Haynaldianum (Reichb. f.) Pfitz. 26 D&ML 49 Sect. CocHLOPETALUM Hallier P. Chamberlainianum (O’Brien) Pfitz. 26 D 747 P. glaucophyllum J. J. Smith 36 D 47 Sect. STICTOPETALUM Hallier P. hirsutissimum (Lindl.) Pfitz. 26 D&ML "49 Sect. NEUROPETALUM НаШег 26 F '34 28 F 241 P. villosum (Lindl.) Pfitz. 26 GM '47 26 D '47 26 D&ML 748 -T ; 26 GM 47 P. Boxallii (Reichb. f.) Pfitz. 26 D&ML '48 * 'The abbreviations used for authorities are: Raymond A. MacLeod; F, Eleanora Francini; GM, Gustav A. L. Mehlquist. D, Robert E. Duncan; D&ML, Robert E. Duncan and [Vor. 36 446 ANNALS OF THE MISSOURI BOTANICAL GARDEN Chromosome m | ора ана Authority Date 32 F 731 P. insigne (Wall.) Pfitz. 26 GM ^47 26 D 47 26 D&ML 48 P. exul. (O'Brien) Pfitz. 26 D 747 26 D&ML 49 P. Charlesworthii (Rolfe) Pfitz. | ч ТА А P. Gratrixianum Rolfe | i а. ња Sect. THIOPETALUM НаШег | . Druryi (Bedd.) Pfitz. 26 a р 47 ыы. re 26 D&ML '49 Sect. CvMATOPETALUM Hallier 30 F 731 P. Spicerianum Pfitz. 28 D 47 28,30 D&ML 749 Sect. CERATOPETALUM НаШег 26 D '47 P. Fairricanum (Lindl.) Pfitz. 26 GM 47 26 D&ML 749 Sect. SPATHOPETALUM Pfitz. P. venustum (Wall.) Pfitz. 1 5 es Sect. BLEPHAROPETALUM Pfitz. P. tonsum (Reichb. f.) Pfitz. 34 D | 47 P. Mastersianum (Reichb. f.) Pfitz. 32 D 47 P. javanicum (Reinw.) Pfitz. 36 р 47 P. Пауапит (Reichb. f.) Pfitz. 34 D 47 P. Мани Summerhayes 40-45 D '45 Sect. PHACOPETALUM Pfitz. е А 36 D 47 P. Curtisii (Reichb. f.) Pfitz. 16 GM 47 Р. superbiens (Reichb. f.) Pfitz. 38 D 47 32 F 731 : 38 F 34 P. barbatum (Lindl.) Pfitz. 8 D ya 38 GM '47 ; 32 D '47 P. callosum (Reichb. f.) Pfitz. 32 GM 47 А 36 р 747 P. Lawrenceanum (Reichb. f.) Pfitz. 36 GM 47 1949] McQuapr—Paphiopedilum Maudiae Hort. 447 IV. Meiosis Aceto-lacmoid and aceto-orcein gave metaphase I smear preparations which were clear enough to determine the nature of pairing but the outlines of the chromosomes were often not sharply defined. A modification of the Feulgen technique, although somewhat better, gave much the same result. These methods also proved unsatisfactory for a study of the prophase chromosomes; all of these methods seemed reasonably good for material in anaphase I or later stages. Тһе difficulty was not remedied even when an iron aceto-carmine mordant was added to the Carnoy's Fluid used for killing and fixing. The situation was further comp- licated by restrictions on the number of buds available; forty were used in the course of the study of pollen mother cells. It must be remembered that none of the plants involved are frequent bloomers, the parental forms blooming once yearly and Maudiae at most twice. It was found after considerable experimentation that the best preparations were obtained by making permanent slides. Anthers were dropped in Carnoy's Fluid for 10—15 seconds and then put in CRAF for 24 to 48 hours. They were then washed and run through the usual tertiary butyl alcohol series and, after infiltration, were embedded. Sections were cut at 10 м and then stained in crystal violet and saf- ranin (Stockwell, 1934). "This method gave results in pachytene and early diplo- tene which could not be obtained with the smear methods mentioned above, and the metaphase chromosomes were sharply outlined. А peculiarity in the staining effect was noted with the use of crystal violet and safranin. When the slides were left in the stain for a long time both meiotic and mitotic chromosomes were stained dark purple. When exposed to the stain for about 15 minutes the meiotic chromo- somes were red while the mitotic nuclei or chromosomes were blue. Ас an ex- posure of about 12 minutes both meiotic and mitotic chromosomes were red. No satisfactory study of chiasmata was made because of the extremely re- duced size of the bivalents at mid-metaphase and because the procedure adopted does not spread the chromosomes sufficiently as the smear method so often does. Diakinesis, a stage favorable for the study of chiasmata in many other plants, could not be prepared suitably by any of the methods cited above. P. Maudiae Hort. var. magnificum furnished most of the hybrid material studied although WESTONBIRT and DELL varieties were examined; the last do not appear to differ from magnificum. PAPHIOPEDILUM CALLOSUM (2n — 32) Prophase I.—Pairing appears to be normal when viewed in the pachytene stage. Permanent preparations of diakinesis in callosum resemble those of Maudiae prepared by aceto-lacmoid smears or the Feulgen technique. Within the various techniques employed in this study it may be said that diakinesis in the two species and the hybrid is not a favorable stage for study. The chromosomes appear to be in a fluid or somewhat fluid state and are therefore poorly defined by the stain; large patches of chromatin are occasionally seen where the chromosomes appear to [Vor. 36 448 ANNALS OF THE MISSOURI BOTANICAL GARDEN have stuck together. This perhaps represents а time when the matrix is being elaborated most freely and is therefore quite thin in consistency. Some chiasmata can be seen but no accurate over-all picture can be given. Occasionally they appear as relatively colorless strands between the ill-defined chromosomes and again they appear to be surrounded with enough matrix to render them stainable. Among the chromosomes of intermediate length, two ring configurations have been noticed: one with no free ends and another in which one of the chiasmata is apparently not terminalized so that a short portion of one end of each chromosome is free. The same pair may be involved in both cases. Among the shortest chromosomes (at least those with subterminal centromeres) bivalents are formed with only one chiasma. Тһе longest pair appears to form three or four chiasmata. The nucleolus is plainly visible at diakinesis. Metaphase I.—Metaphase chromosomes have not been stained as clearly as desired by smear methods but were clear enough to determine pairing; chiasmata could not be properly interpreted. Twenty-five metaphase cells examined showed pairing to be in 16 bivalents. (See pl. 32). Of 114 metaphase plates seen in side view, 107 appeared to have all their chromosomes in the plate in the normal fashion. One cell appeared to have one lagging chromosome, 2 were doubtful as to fragments or isolated chromosomes, and 5 appeared to have isolated chromosomes or fragments in each. It is entirely pos- sible that these apparent abnormalities existed as a result of smearing. Anaphase I.—Fifty-six anaphase cells were examined and no anaphase bridges were observed. One cell appeared to have a fragment between the two poles. Anaphase appears to be normal. Telophase I.—No cell wall is formed at the close of division I. An interphase nucleus is formed. Тео phase П.—Еіүе hundred tetrads were examined. No micro-grains or small extra cells were found within the thick wall. Pollen. grains.—Pollen grains were mounted in aceto-lacmoid for staining but were not pressed. The grains are elliptical to spherical in shape. Of 500 grains examined only 8 were found to be empty; no shrunken or distorted grains were found. The grains are slightly smaller than those of P. Maudiae. (See section on pollen grains, X Paphiopedilum Maudiae). P. UM LAWRENCEANUM (2n — 36) Propbase 1.— Мо leptotene or zygotene nuclei have been studied but pachytene and early diplotene nuclei have been observed. The crystal violet-safranin tech- nique was used. Pairing as observed in pachytene appears to be regular throughout and no abnormalities were noticed. Тһе early diplotene stages examined show the initial repulsion between the paired chromosomes. Metapbase I.—Only 2 metaphase I cells suitable for study were found. These indicated pairing to be in 18 bivalents. 1949] McQuaApE—Pabbiobedilum Maudiae Hort. 449 Anapbase I— Observations on a large number of these cells indicate no apparent abnormalities. Bridges are not formed nor do there appear to be any lagging chromosomes or fragments in evidence. The chromosomes do not appear to be split in readiness for division 2 at this stage. Telophase I.—An interphase nucleus is formed at this stage but no cell wall is formed. Eighteen chromosomes were counted at one pole in an early telophase. Metaphase I].—Examination of metaphase II cells indicated that division had been regular. The regularity with which 18 chromosomes are disposed to each pole serves as additional evidence that 18 bivalents are formed at metaphase. (See ple 32); Anaphase and Telophase II—No abnormalities are evident and the tetrads formed resemble those of P. callosum. A tetrad was found in which 18 chromo- somes could be counted at one pole. Pollen grains.—Pollen grains are elliptical to spherical and about the same size as the callosum grains. Of the 500 grains examined, only 2 were found to be empty. (See section on pollen grains, X Paphiopedilum Maudiae.) ХР UM MAUDIAE Hort. (2n = 34) Prophase I—In the pachytene of P. Maudiae the nucleolus appears as а well- defined almost colorless body in intimate contact with chromosomal material. Although in sectioning the knife may frequently remove the nucleolus from cells, it is visible nonetheless in many others. Its attachment to a chromosome is notice- able. It is impossible to tell which chromosome is involved since the pachytene nucleus represents a tangle of long slim threads whose ends, except in chromosomes cut through by the knife, are often not visible. As with callosum and Lawrence- anum two nucleoli are occasionally visible in the mitotic cells forming the jacket around the pollen mother cells but only one nucleolus has been seen in the PMC's. The nucleoli seen in the PMC's are larger than those seen in mitotic cells. There have been observed in the pachytene and pachytene-early diplotene chromosome configurations which suggest inversion and deletion but the chromo- somes are so attenuate and the suspected loops or buckles so small that it has been impossible to ascertain this. The presence of inversion loops in pairing is to be expected in view of the presence of anaphase bridges. As in Lawrenceanum the beadlike effect of the chromomeres is evident at this stage. Metaphase I.—The staining with smear methods was suitable enough to examine pairing. Of the 27 metaphase cells of P. Maudiae examined, 26 showed 17 bivalent chromosomes. (See pl. 34). One cell showed 16 bivalents and 2 univalents. It appears that two of the Lawrenceanum chromosomes are pairing. Anaphase 1.—А total of 483 anaphase I cells were examined and several ab- normalities were found: [Vor. 34 450 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN a. Cells Mor 1 bridge but no fragments 17 b. Cells with 2 bridges but no fragments 1 c. Cells with 1 bridge and 1 fragment 13 d lls with 1 bridge and 2 fragments 1 e. Cells with 2 bridges and 1 fragment 2 f. Cells with 1 fragment (or small isolated chromosome) 15 TOTAL 49 Of the cells examined 10.1 per cent showed abnormalities. A further analysis of anaphase I was made possible through the observation of 4 cells in which the chromosomes could be counted at one or both poles. While this number is far too small to be regarded as significant the information is in- cluded to amplify the notes above. 1. Two cells showed clearly 17 chromosomes at one pole but a count could not be made at the other pole. There were no lagging chromosomes in evidence. The chromosomes were split in readiness for division 2 but this partial separation may have been caused by smearing. 2. One cell showed 17 chromosomes at one pole and 16 at the other. One chromosome lagged but appeared to be slightly closer to the pole with 17. Again the chromosomes were split. 3. One cell had 17 chromosomes at each pole; these were split for division 2. Tetrads.—Examination of 527 tetrads was made. The arrangement is pri- marily that of 2 cells in each of two planes although 4 in one plane is not rare. The presence of а micro-grain or micro-cell, a small, extra cell within the tetrad wall, was occasionally noted. Hui ciii = „йок: 1. Tw їп of two planes 417 2. Four b in one plane 72 Apparently abnormal groups: 1. Two cells in two planes plus one micro-grain 28 2. Four cells in one plane plus one micr 4 3. Two cells in one plane (diad) plus one micro-g 1 4. Three cells in one plane (triad) plus one micro-grain 5 TOTAL 527 Of tetrads examined 7.2 per cent showed irregularities. Pollen.—In general the pollen grains range from elliptical to spherical. Of 500 grains examined 54 (10.8 per cent) were empty and some of them were distorted. (See pl. 33). Four of the empty grains were about one-third the normal size. The full regular cells are slightly larger than those of P. callosum and P. Lawrenceanum. Measurements of 100 Maudiae pollen grains and 100 each of callosum and Lawrenceanum indicated that those of Maudiae average .0580 mm. along the long axis, while those of callosum and Lawrenceanum are .0530 and .0526 respectively. That the slight difference in size between the grains of the offspring and the parents is not merely based on variation within the samples was confirmed with a standard deviation test. 1949] McQuapE—Pabbiobedilum Манае нокт. 451 In both species and in the hybrid, leptotene апа zygotene probably occur when the bud is quite small. Just how soon pachytene begins is not known but appar- ently pachytene and pachytene-early diplotene go on for considerable time, some- times even a matter of days, depending upon environmental conditions. The ге- maining stages, particularly diakinesis and metaphase I, progress rapidly. The lengths of some buds, divested of enfolding bracts, are listed together with the meiotic stages exhibited: P. callosum 13.0 mm Pachytene 15.0 mm Pachytene Pachytene to diakinesis 16.5 mm Early metap Metaph 17.0 mm Primarily diads. А few metaphase and anaphase I cells 18.0 mm Tetrads P. Lawrenceanum 16.5 mm Pachytene, early diplotene ( A few metaphase and anaphase I cells but chiefly division 2 18.0 mm Division 2 Tetrads 18.5 mm Division 2 19.0 mm Pollen grains x P. Maudiae 13.0 mm Pachytene 15.5 mm Pachytene 16.5 mm Diakinesis to early metaphase I ( Pachytene 17.0 mm 1 Pachytene to diakinesis | Pachytene to mid-metaphase I 18.0 mm 1. Metaphase I 2. Telephase I through division 2 18.5 mm Tetrads V. Discussion It will be noticed in the anaphase I data of the hybrid that of the 49 cells listed as showing disorders, group c (13 cells) is characterized by one bridge and one fragment. ‘This is the typical result of pairing between two chromosomes when one of them has an inversion not involving the centromere and a crossover occurs within the inverted region. = Group а (17 cells) exhibits the bridge but no fragment. ‘Two possibilities exist here: either the fragment was not visible or it did not exist. Since none of the fragments observed either with or without bridges were large it is possible that in these cases it had been displaced by smearing or that it had been concealed by [Vor. 36 452 ANNALS OF THE MISSOURI BOTANICAL GARDEN the chromosomes at one or the other of the poles. There being evidence that in- versions exist, this possibility must be given the more credence. The other pos- sibility is that in these cells no fragment existed as a result of an inversion bridge. is would necessitate another hypothesis for the origin of a bridge without a fragment, and there is no evidence here to support such a hypothesis. The single cell in group Ё with two bridges but no fragments should be considered in the same light except that in this group two chromosome pairs were probably involved. Group e (2 cells) is characterized by two bridges and one fragment. The most logical assumption here is that one of the fragments was rendered invisible as in the case above. Тһе single instance of one bridge and two fragments (4) may well indicate the breakage of one bridge before the other. In these cases, as in Ё, two chromosome pairs would be involved. Group f is composed of 15 cells which showed one fragment or small isolated chromosome. It is thought that these were true fragments since they were quite small. In such a case the bridge would have been already broken. However, the possibility that these might have been lagging chromosomes cannot be overlooked since many of the typical mid-metaphase I bivalents of the hybrid are small. Ап interesting case is presented by the pair of chromosomes of intermediate length with prominent secondary constriction. It is this pair which exhibits the inversion mentioned in the section on root-tip analysis. (See pl. 32). Half of the chromosome is involved in the inversion for which the hybrid examined is hetero- zygous, and the inversion, when viewed under the microscope, appears to be terminal. The possibility must not be overlooked that there is chromatin material here which cannot be seen and would render the inversion interstitial. It has ong been felt that inversions generally did not involve chromosome ends (Darlington, 1937; Kossikov & Muller, 1935; Muller, 1938). However, there is considerable evidence to date that terminal inversions do occur (Kauffmann, 1937; Sutton, 1940; Carson, 1944; Carson & Stalker, 1947), and the possibility that the in- version mentioned here is truly terminal must not be brushed aside. This inversion, since it involves the centromere, would not bring about the formation of a bridge. A single crossover within the inversion would result in a duplication-deficiency chromatid as well as one complete chromatid going to each pole. Of the complete chromatids, one would show the inversion (Sturtevant & Beadle, 1939). It will be noticed that although 10.1 per cent of the anaphase I cells exhibited disorders, only 7.2 per cent of the hybrid tetrads were visibly abnormal. This probably can be taken to mean that some of the tetrads, although appearing normal, аге not во. Іс cannot be said that there is any absolute correlation between the 10.8 per cent visibly non-viable pollen and the similar percentage of visible anaphase I abnormalities even though it is suggestive. No doubt there are other factors to be considered. The 10.8 per cent of non-staining pollen grains is not a true estimate of the hybrid's sterility in view of growers’ experiences in attempting to use Maudiae as a parent plant. There are, no doubt, many pollen grains which appear quite normal but whose viability is terminated at later stages. Male gametes 1949] McQuapE—Pabbiobedilum Maudiae нокт. 453 may become non-viable in the pollen tube; the pollen tube may grow too slowly; the egg may be non-viable; the zygote may be non-viable; or the seedling perish early in its existence. That the 10.8 per cent non-viable pollen is not a complete picture of the sterility of the hybrid is supported by what is known of pairing in Maudiae. It will be remembered that pairing in Maudiae was typically indicated by 17 bivalents at metaphase I. This suggests pairing between two Lawrenceanum chromosomes, and if two chromosomes are like enough to pair consistently this would in turn suggest polyploidy. It is entirely reasonable to assume that such pairing is the result of polyploidy. A glance at the list of chromosome numbers at the close of section III reveals that throughout subgenera BRACHYPETALUM and ANATOPED- пом the n number is 13. In the subgenus OTOPEDILUM, the group to which all of the individuals dealt with in this study belong, higher 7 numbers approach the triploid level. It is suggested that this increase in chromosome number is the result of hybridization between groups that cannot be named at this time for lack of evidence. The mere statement that pairing іп Mazdiae is in 17 bivalents and that the 2 remaining Lawrenceanum chromosomes pair is probably ап over-simplification. P. callosum is characterized, as we have seen, by a chromosome set of 6 pairs of terminal or subterminal chromosomes and 10 interstitial pairs while P. Lawrence- anum possesses 10 terminal or subterminal pairs and 8 interstitial pairs. The dif- ficulties of pairing a Lawrenceanum and a callosum chromosome for each of 16 bivalents (in addition to the bivalent already formed by each of two Lawrenceanum chromosomes), in view of the differences in morphology, are apparent. It is highly probable that some callosum chromosomes are pairing amongst themselves as well as with Lawrenceanum chromosomes, and this possibility would exist equally weli for chromosomes of Lawrenceanum. Under such conditions gametes could be pro- duced which would lack necessary genic elements. From the foregoing facts it seems logical to conclude that the causes of sterility in the hybrid are: 1. Visible disorders in anaphase I due to inversions. 2. Invisible disorders at anaphase I due to an inversion which includes the centromere and therefore results in duplication-deficiencies. 3. Pairing of some callosum chromosomes and some of the Lawrenceanum chromosomes among themselves (because of their polyploid background) takes place so that some of the gametes are without necessary chromatin material. VI. SUMMARY 1. Acytological study was made of the X Paphiopedilum Maudiae Hort., the result of a cross between the albino forms of P. callosum and P. Lawrenceanum. Because of its high sterility, P. Maudiae as a parent plant has rarely, if ever, pro- duced any offspring of a quality equal to, or exceeding, its own. 2. А short discussion of the genus and the histories of the parental forms and the offspring are set forth. [Vor. 36 454 ANNALS OF THE MISSOURI BOTANICAL GARDEN 3. Some observations are made on the status of X Paphiopedilum Maudiae Hort. as a parent plant. 4. А cytological analysis of root-tips of the parental forms and the hybrid gave the following results: a. Chromosome numbers of P. callosum, P. Lawrenceanum, and P. Maudiae are confirmed as being 32, 36, and 34, respectively. Two other species counts were also confirmed (P. barbatum, 38; P. Curtissii, 36; P. superbiens, 38). b. P. callosum has 6 pairs of chromosomes with terminal or subterminal centromeres and 10 pairs with interstitial centromeres. с. Р. Lawrenceanum has 8 pairs with interstitial centromeres and 10 with terminal or subterminal centromeres. Maudiae is heterozygous for an inversion which seems to Бе terminal. 5. Study of meiosis in the parental forms and the hybrid gave the following results: a. P. callosum and P. Lawrenceanum undergo normal meioses with bi- valent pairing. b. Pairing in P. Maudiae is in 17 bivalents. Some disorders are visible at anaphase I. c. Only 10.8 per cent of the Maudiaze pollen grains are visibly non- viable. This does not give a true picture of the sterility of the plant. 6. Some conclusions are offered as to the several causes of sterility of the hybrid: a. Inversions that give rise to anaphase I disorders. b. An inversion, visibly terminal upon microscopic examination, which does not give rise to visible anaphase I disorders because it includes the centromere. с. Аз a result of polyploidy in the genus some of the callosum chromo- somes pair with themselves as do some of the Lawrenceanum chromosomes in meiosis of hybrid pollen mother cells. Some of the gametes are therefore deprived of necessary chromatin material. LITERATURE CITED Anon. (1912). Cypripedium callosum Sanderae. Jour. Hort. 65:565. Black, J. М. (1933). Albinism in Cypripediums. Orchid Rev. 41:69-72, Buser, R. (1894). Cypripedium ou Cypripedilum? Bull. Herb. Boiss. 2:642-644. Carson, H. L. (1944). An analysis of natural chromosome variability in Sciara impatiens Johannsen. Jour. 311 чч orph. 75:11—59. ------, and Stalker, D. (1947). 7 arrangements іп natural populations of Drosophila robusta Sturtevant. ыа. 1:113—13 Cooper, E. (1946). White P Leap rd on hid Rev. 54:106- ас P. D. 2 Recent Advances in ЦЕ Ки: 2па i ” Philadelphia. Duncan, R (1945). ње жасын of variable — n of chromosomes within the root tip ба Paphiopedilum Wardi Am. Jour. Bot. vi —50 , (1947). The P ни Slipper. Orchid Dig. ти ------, апа чи од, . (19483). M ns of the ka i Am. Orchid Soc. Bull. 17: :170-17 1949] McQuave—Paphiopedilum Maudiae Hort. 455 Л : ОВА Chromosomes of the insigne complex of ladyslippers. Ibid. 4 —— — 49а). ar wero of the continental species of е ich solid green leaves. Ii 18:84—8 — -----, 49Б). The j^ m of some of the Polyantha. Ibid. 159—163. disci ЕЗ (1951): m embriologiche e cariologiche sul genere “Cypripedium”. Nuovo Giorn. Bot. Ital. N. S. 38:154-212 ы (19 32) m e сакен nella Fo di “X Paphiopedilum Тееапит”. Ibid. 39:251- 25 1934). Ibridazione interspecifica пе! genere “Paphiopedilum.” Ibid. 41:189-237. 632. Hooker, ТА 5 (1879). Cypripedium Laurenceanum. Curtis’s Bot. Mag. tab. Kauffmann, B. P. (1936). А terminal inversion in Drosophila ananassae. Proc. Nat. Acad. Sci. 91—594. konkor, R. з апа Muller, H. }. Seles Invalidation of the genetic evidence for branched chromonenas. Jour. Here 305—3 Mehlquist, G. "t `L (1947а). Polyploidy in с genus кнн Pfitz. (Cybribedium Hort.), and its practical implications. Mo. B ard. Bull. 35:2 25 ). Some smear а ија bu counting Kec НА in SE Meyer, J. R. (1943). Colchicine- ока leaf smears. Stain Technol. 18:53 Muller, H. 1. (1938). Тһе remaking of chromosomes. Collecting Net 13, No. Pfitzer, E. (1903). Orchidaceae Радне E o r's Das Pflanzenreich, IV, Pam. 50 (Heft 12). Reichenbach, т С. (1854). Xenia Orchidac ------, (1878). ое D IDE Gard. Chron. II, 18:748. ў «ее 2. 9, pl. — E (1886). OE ia callosum: n. sp. Gard. сн П, 26:3 Rolfe, R. А. (1896). Тһе Cypripedium group. Orchid то 327—334; prm 367. РЕА. (1900). crm x Maudiae. dou 8:3 Sander, G ES В. К, апаш ІК (1927). уаде Orchid G uide. v. ed. кө ЦЕН Sander, 5 К. WA Sander's Lia plete List of Orchid Ed p E^ mk P. A. A d: 4). А stain px dica plant material. Science N. S. "80: 121-122. Sturtev ü „ and Beadle, G. W. (1939). An Introduction to дот s Philadelphia and Ibid. 229-233. > de E (1940). Terminal deficiencies in the XX chromosome of Drosophila Melanogaster. Genetics 4—540. [ Wilson, Gurney], (1933). Cypripedium Maudiae. Orchid Rev. 31:299—300. [Vor. 36 456 ANNALS OF THE MISSOURI BOTANICAL GARDEN CHART 1 A LIST OF THE CROSSES IN WHICH 2 МАУО/АҒ НАЈ BEEN USED ALA PARENT. THE NUMBERS BELOW EACH VARIETAL МАМЕ INDICATE HOW MANY TIMES THE VARIETY WAS BEEN VIED Af A PARENT. TQ THE RIGHT OF THE ANTECEDENT SPECIES ARE THE SECTIONS OF OTOPEDILUM OR THE /UBBEWERA TO WHICH THEY BELOWG. MAUVOIAE (SECTION PWACOPETALUM OF SUBGENUS 270260407) J^ &/ СА "Атм MAUD a ACTAE UJ өс? LEEANUM ии 1м tro. TALUM огешат CYMATOPETALUPM PHALO. MAULIAE < LOJU^P VAR. JANDERAE А! рор /7 VAR. MYFANUM _ PAACOPETALUPM 2.Х ALMA GAVAERT : ALMAUO. ALMA GAVAERT “ e CAWRENCEANUM VAR. HYEANUPT _ PHACOPETALUM а EURO, ЕЛА ACTAEUJ < ANTINIUS IX ARMISTICE + ARMAMENT. ARMISTICE Z <, goi СРАМИ" жас сти Ами г 2 ERU ME FURÜPETALUP LOSUN WE WEURDPETALUPM 4х AUREUM = BENITA. AUREUM — Ај ae 58 2 SPICER/A NUI "AL e AVWVIIPDPFYA / si, BARBATUM PHALOPETALUM SX BINGLEVENSE + COLOURFUL. BINGLEVENSE < 2/ f АРКАИМ са СОРОМ NEURUPETALUM é x CALLOIO-BARBA TUM = METEORITE. CALLOIO- BARBATUM = PARERE PARAT BARBATUM .. TALUM zx сои “ HOLDENI me UP) WACOPETALUPM ах И ms . IRIS IE СНА ВЕС ДРАМА МОУ (Ја) 9х жые = COLORADO Кы OLARE (Ја) PNACOPETALUN er о СИКТИ (SANDER ии PETALUM EMERALDS др СКО is VAR JANDERAE — PHACOPETALUM LAW, — VAR. HYEANUN рін COPETALUM /Q. X CLAIRE DE LUNE » PIT YLEWE CLAIRE DE LUNE OIUH VAR. SANDERAE PHACOPETALUM 2 2 — електі а NUM VAR HYEANUM — PHACOPETALUM i далда “ала OAVAERTN се дут VAR HYEANUIT РНАСОРЕТАСИН / = авесил (OED/PPE) ( ініні jm LOSUITNEUROPETALU/T У“ 24 2/СЕА/АЛМШ/7Т очни (CUM LL СУМАТОРЕГАСИТ ‘LT OP. —— ПИЯ STILT ÜPETALUP Wf. DE CURTE c — geri ИХ CRUSADER = PALESTINE. CRUSADER м0. 440.07 VAR BOXALLHNEUROPETALUM 8 o ИШОЈИТ VAR BOXALL -- TALUM HERA (EURYADES) © , cea yy < "MIENE — MEURDPETALUM LUCIFER SPICERIANUIT _— CYMATOPETALUM P DICE AAJ NIOBE - DICA АР ына /2.х 7 We CURT MH. ENERALO veru (Ја) бурн 7273 рендано ACTAEUS (NIIGNE 741407 ££A НИТ x DIRA г ATAUDORA. DORA 4 JPLCERO/A NUM CY/TATOÜPETALUPN + 2 mo <= СУМАТОРЕТА И — WEUPDPETALUPP “4 vs TRE ЛИВ АЕ А; NITENS (SANDERI) < фо ИШАО NEUPDPETALUM CHARL ESI NORT HI NEU ROPETALU SF BWELEVENWSE аншы Аны ы NOT лі ІІІ MEAD АХ EARL OF CHESTER = SUNSET, EARL OF CHESTER wae NEUROPETALU IT 2 LIRO ROBERTS жаға А МИТ BARBATU/AA_. #WACOPETA ГУ ском м == FUP S. NEUROPETALUI LENANTHE ST? 1949] McQuapzr—Paphiopedilum Маите новт. 457 &427//// (JANOERAE ) PHALOPETALL//F ж /7 А « VERT. EMERALD а CAL ^7 VAR. Ni ONAE <“ VAR. SANDERAE ____ PHACOPETALU 4 А ALITA GAVAEAT. fi : 15. X ENCHANTRESS = ESMERALDA. ENCHANTREIT i амн ашр й WR. MVEAMUAT o PMALDPETALUM 2 > LAWRENCEANU/T VAR КУРААР PWACOPETALU/T [477 дА (JANDERAE ) PHACQOPETALUPMPM т X FAIRRIEANU/ = FA ii - МАШО. FAIRRIEANUTST С. CERATOPETALU/F 85 VILLOSUM VAR. E WEUROPETA ГТ м pco ttl КЕ 7 LORD KULME tr сул DAETALUTY POMER LHLORIS < LEEANUM ^ e INE. a £TALUM м МАУ. НАЛУ 22 1 "ФЕ в — САРА в ONADES MUN б ель гит) сетти CYMATOPETALUP (ULUSTRIOUS ) NONI.DE CURTE <=“ GHE NAE ae FULMINITER С OSUI VA BDYALLIL. MEURDPETA LU M REWA ear S py < MIONE EUR OPETALUP PICERI, /8.x FULHINITER s PADDINGTON (CHAROWAR ) "e LER. moe es AUREL E x DILLON HITENI а | КА SUH. NEUROPETALUFPT UALLIEA AUREUM) КАКИМ, СУМАТОРЕТА ИМ AUREUM pes ВМ MEURDPETALUPM VLLOJUAP —— NEURDPETALUP INSIONE NEUROPETALU/T sat АЙАНА семан СУРА алые ALCIBIADES ефора ОМГОЕ CORTE АА poe: VAR. BOXALLIL. europera LUM ШЕМИ ГЕД JLLUJUM VAR, BOXALL //—— NEUR OPE: wera L INSIGNE. NEUROPET: ons LORO WOLMER адр EC jii ins СУМАТОРЕТАЕ 0Р7 м 7 сыа won МЕНЕ. NEURO PETALUM aR < оо. LCYMATOPETALLIT IRS. HILARY JENKINS: ом< <; N х GÜULTENIANUF + ONYX. GOULTEN/ANUTT = 2410707 к ыы 5 2 CURTHM PHALOPETALUM CALLOSUM VAR. SANDE, PHACOPETALUM 20.Х HOLDENI = WARDEN. HOLDENI i Я Ў pru <“ pec VAR. JANDERAE . PHACOPETALUM LAWRENCEANU/T VAR. HYEANUPM. . PHACOPETALUM 2.x INSIGNE + ROSIE TTI. INSIGNE (ER) MEURÜPETALUPM 22. X 4A Ж ХОЛ A ма GAVAERT. LAWRENCEANUM (ОДА) PHACOPETALYH 23. x VERUM HANBUR Ds LEEANUM КЕ. = 2.2004 ny р: J4PICERIANUPM CYMATUPETALUPM INIIGN. pow с. fo — — ЖЕИФОРЕГА ЦИ ғамал «CC ИБМ... MEUPPETALUM CHRISTOPHER SPILERIANU 4. WIIGNE ; ; 24. X IIRI ELEY МОИТЕ // 247722 p cent а NUM — СУМАГОРЕГАИТ #3 о ¿wan —<— 2 зо AEURDPETALIM ALCIBIADES ии САУ... CYITATOPETALU M 47. NEUROPET; СОТТОООФЕ (TE 0/07 М UFULODDEFYA! EIPS prever SPICERIANUTI. CYAATOPETALU ^f JERNINGAHA /Т/АЕ WLLOSU/7_____. NEWROPETALUST WNNIAN UM 4 емен < деи фул. THIOPETALUYM 24. X NIVEUM = PURITY. NIVEUM (Ја) ЈГУФСЕМЕЈ BRACHYPETALLTS #2 . э [Vor. 36 458 ANNALS OF THE MISSOURI BOTANICAL GARDEN ж? pov GNE NEUROPETALUI . (8/GAWT EUM) ТРЕКАМИ CYHATOPETALUPM ALCIBIADES sa том DE CURTE < Боов АЙИН 26.X NIRVANA = TAUDANA.N/RVANA VALQIUM VAR BOXALL NEUROODETA LUP - a meo JP/CERIANUM CYPATODETALUM INSIGNE NEUROPETALUIT (VIRG/NALE) я —UO з, rs АА VILL OSU M ——_ WEURDPETALUPM INIIGNE “¿ee Aa С ANTINOUS САСА uon py р ARMISTICE Z нош ? LEEANUN < огке/дуу РАСИ 27.Х РИСК и FOLLY. PUCK INIIGNE | э 2 NITENS —— V/LLOJU^H INSIGNE NEUROPEVALUST LEEANUIT (GRATRIXIAE ) қ sed as parent ве їп о: 1. РНАСОРЕТА ОМ X ALMA САУАЕКТ (Chart І #2)ї 11$|NEREID 0+ 2. PHACOPETALUM X APPLETONIANUM (App. Chart. П #1) 4 |SIAMENSE 1 NEUROPETALUM 3. PHACOPETALUM x Argus 40 |CALLOSO-ARGUS 0 4. PHACOPETALUM X ASHBURTONIAE (App. Chart П #2) 12 [ZENOBIA 1 NEUROPETALUM 5. NEUROPETALUM X AUREUM (Chart I #11) 58 |ALTRICHAMENSE 0 CYMATOPETALUM 6. PHACOPETALUM X barbatum 49 |CALLOSO-BARBATUM 10 7. Subg. BaacHvPETALUM X bellatulum 57 |Моттоми 1 8. NEUROPETALUM x Boxallii (var. of sp. villosum) 6 42 |J. BARTELS 0 9. PHACOPETALU CALLO-ROTHSCHILDIANUM 1 |FRANCONIA 0 Subg. Азы (App. Chart П #3) Sect. CORYOPEDILUM 10. PHACOPETALUM X CALOPHYLLUM (App. Chart II, #4) 13 |PArras 2 SPATHOPETALUM 11. COCHLOPETALUM X Chamberlainianum 38 |ALCIPPE 0 12. NEUROPETALUM x Charleswortbii 66 |RosITa 0 13. PHACOPETALUM x ciliolare 27 |Zeus 0 14. Subg. BRACHYPETALUM Ж concolor 27 |CONCO-CALLOSUS 0 15. PHACOPETALUM xX Curtisii 44 |GOULTENIANUM 6 16. Subg. BRACHYPETALUM X Delenatii 14 |MME. MARTINET 1 17. THIOPETALUM x Druryi 33 |А. R. SMITH 0 18. NEUROPETALUM X exul 20 |Ок. Conway 0 19. CERATOPETALUM X Fairrieanum 85 |JUNO 1 20. PHACOPETALUM X gigas (App. Chart II, #5) 8 |E. J. SEYMOUR 0 NEUROPETALUM 21. Subg. BRACHYPETALUM X Godefroyae 31 |FELIXx FAURE 0 22. PHACOPETALUM X GouLTENIANUM (Chart II, 2:15) 6 |MALHERBE 0 23. PHACOPETALUM X GowrRiANUM (App. to Chart П, #6) 19 [HORTENSE 0 24. PHACOPETALUM X HARRISIANUM (App. to Chart II, #1) 72 |LEDouxIAE 1 NEUROPETALUM 25. STICTOPETALUM x bhirsutissimum 33 |DONCASTERIANUM 0 26. NEUROPETALUM х HITCHINSIAE (App. to Chart II, 3:7) 17 |SoONIA 0 27. PHACOPETALUM >x HorpENm (Chart II, #37) 6 |GLORIOSUM 0 28. ЗРАТНОРЕТАТОМ >x Hookerae 26 |FoRTUNA 0 29. NEUROPETALUM X insigne 148 |LEONIAE 2 30. BLEPHAROPETALUM X javanicum 7 |ТАУА 0 *Species names аге “o ida їп italics; hybrids, ipie: and a pim in caps. TKey to explanations of varietal backgrounds are given in parent iThe figures following nex in шка the пит сы ег of times T pom has been used in crosses. §Treated as P. villosum va xallii in Sander’s ‘Complete List of Orchid Hybrids’; treated as species Р. Boxallii by G. A. L. Macte (1947) and R. E. Duncan (1947). 460 [Vor. 36 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN Section of OTOPEDILUM 1 hybrid Resulti (or other subgenera) species ór hy ú ези ing ; used ав parent y used in cross 31. NEUROPETALUM x J. M. Brack (App. Chart II, #8) 46 JAMES 0 CYMATOPETALUM Subg. BRACHYPETALUM 32. CYMATOPETALUM X LATHAMIANUM (App. Chart II, #9) 39 |CALLIOPE 0 NEUROPETALUM 33. PHACOPETALUM X Lawrenceanum 62 |MAUDIAE 35 34. NEUROPETALUM X LrrANUM (Chart I, #1) 132 |ANGELIAE 0 CYMATOPETALUM 35, ? X MADAME COFFINET 2 Марлм MAXINE Oprorx 0 36. BLEPHAROPETALUM X Mastersianum 24 |Рүтно 0 37. PHACOPETALUM X Maupiar (Chart I, #2) 35 |Ногреми 6 38, NEUROPETALUM X NITENS (Chart I, #3) 66 |WENDIGO 0 39. Subg. BRACHYPETALUM X niveum 48 |WINIFRED HOLLINGTON 1 40. PHACOPETALUM X OENANTHUM (App. Chart Ш, #2) 28 |OrcaA BOGSHAWE 0 NEUROPETALUM 41. Pied ANATOPEDILUM X pbhillippinense 22 |MILLMANII 0 Sect. COR YOPEDILUM 42. Subg. ANATOPEDILUM X Rothschildianum 53 |сліло-Котнв5- Sect. CoORYOPEDILUM CHILDIANUM 1 43. Subg. ANATOPEDILUM X Sanderianum 13 PRINcEss MAY 0 Sect. CORYOPEDILUM 44. BLEPHAROPETALUM X SEMENTA (App. Chart II, #10) 8 |AURELIANENSE 0 15. CYMATOPETALUM X Spicerianum 62 |MLLE. GABRIELLE OENS 46. Subg. ANATOPEDILUM X Stonei 27 |FORDIANUM 0 Sect. PRENIPEDILUM 47. PHACOPETALUM X superbiens 39 |MOUSSETIANUM 0 48. PHACOPETALUM X SUPERCILIARE (App. Chart II, #11) 18 |MOREAUANUM 0 49. PHACOPETALUM TAUTZIANUM (App. Chart II, #12) 2 [NANDI 0 Subg. BRACHYPETALUM 50. BLEPHAROPETALUM X tonsum 33 [FELICITY 0 51. oo X TRIUMPHANS (App. Chart II, #13) 1 RAJAH 0 EUROPETALUM 52. SPATHOPETALUM X venustum 29 |OnPHEUS 3 53. NEUROPETALUM x villosum 60 INDRA 0 54. PHACOPETALUM WILLIAM MATTH BLEPHAROPETALUM (App. Chart П, Thie 1 ERNEsT READ 2 55. PHACOPETALUM WINIFRED Hor GTON Subg. BRACHYPETALUM (App. Chart Т. 15) 1 |WINSUM 0 1949] McQuapE—Pabbiobedilum Maudiae нокт. 461 APPENDIX TO CHART 77 PHALOPETALUMT 4 APRLETONIANUIT (APPLETONIAE 7) =. 6 Ф408470/7 WACOPETALUM MA RRHTANUM <<, М RORETAL UM AABGATULL PHECOPEVALGST 2 AIMBURTON/AE ека УУРУУ. NEUROPETAL LALLOSUST. COPETA un 9. CALLOSO -ROTHICHILDIAN UM отим pian са Саон SECTION | CORYOPSQ/LUTT BARBATUST AM COPETALY 4 CALOPHYLUUMN емуру OAFKOPETA ИМ ТЕУ ЕРТІСТЕ = е In COPETALU TTE eer аА V/LLOJUPM WEUPDPETALUPMP | LAVARENCEA НИТ PHALOPETALUST P277 22777 VACOPETALUM ^ 4 GOWERIANU T UA нра руни CHARLES WORTHH ег, 7 АИРСИИМИ/ АЕ —— € /А2/6 NE NEUROPETALU/T NEUROPETALUIT LEER WU CYHATOPETALUM INIIGNE MK ALC/BIADES я ре FR SERINO INIIENE NEURDDODETALUPM AO) 22. MOIS атуына т R CUPERBU т) RD ода (IONS. ФЕ CURTE ња шей Л MERA VALOJUT VAR. ВОИ а. им. 54ACK (NEW MALL HEY) INSIGNE BEE CKIIAN// < 4/7 BER fcn Lo VAR. BOXALLIL NEUROPETALU IT MEUWROPETALU/T OPETALL/F —— УРАЙ, SUBGENUS BRACHYPETA 4 4” S ИМ. /7О/7 VN JO/CER/AN UT САТМА МТА МОМ m — f. LATHAMIA бор ин JAVAW/CUPM 0. SENENTA —=——— ПА ВС ему ARBAT ж SUPERCHIARE —————— REMARE SYPE. ore “Ç РАД /2. TAUTZIANU ————— 2 PHALCOPETALU/T NIVEL ur J0868€WUJX BRACHYPETALL/T Me NITENS (SALLIERI) == Са 4. < ра СУЛАТОСОЕТА:И/7 NEUROPETALUYIT MA TRIVIPHANS 77 MALRDIC IAM QENANTHUST (SUPERBU IT) “52 P NE WA COPETA Tg 2 EUR OPETALUIT BARBEATY, PAACOPETALL ұй иисои. NEUYROPETAL UIT EUROPETA A MULIAMT PIATTHE YS — ш 4VOENCEANWUM 4... ОМАСОРЕГАИМ МАЈГЕЮЈГА МИ ST Жж WINIFRED HOLLINGTON = Е = NIVELI SUBGENUS BRACHYPETALU/T [Vor. 36 462 ANNALS OF THE MISSOURI BOTANICAL GARDEN CHART III LIST OF CROSSES IN WHICH P. LAWRENCEANUM HAS BEEN USED AS А PARENT* Section of OTOPEDILUM (or other subgenera) Species or hybrid Resulting used in cross progeny used as parent 1. PHACOPETALUM X ALMA САУАЕКТ (Chart I, #2) 11 ELEANOR ROoZILLA 0 2. PHACOPETALUM xX Argus 40 По 15 3. PHACOPETALUM X barbatum 49 |ALMUM j 4. Subg. BRAcHvPETALUM X bellatutum 57 |LAunrE-Brr 2 5. NEUROPETALUM X villosum var. Boxallii 42 |THAYERIANUM 0 6. PHACOPETALUM X CALLOSO-BARBATUM (Chart I, #6) 10 |MvrH 0 7. PHACOPETALUM X callosum 55 |MAUDIAE 35 8. NEUROPETALUM X CAPPAMAGNA (App. to Chart III, #1) 46 [MONTROSE 0 CYMATOPETALUM CERATOPETALUM 9. COCHLOPETALUM X Chamberlainianum 38 [HIERO 0 10. NEUROPETALUM X Charleswortbii 66 |DECIPIENS 0 11. PHACOPETALUM X ciliolare 27 |8мітні 4 12. SPATHOPETALUM X Сікоратва (App. to Chart Ш, #2) 1 |RESPLENDENS 0 HACOPETALUM NEUROPETALUM 13. NEUROPETALUM X CoruMBUs (App. to Chart П, #3) 11 |Sanpow 0 CYMATOPETALUM 14. Subg. BRAcHvPETALUM Ж concolor 27 |Cowco-LAURE 0 15. PHACOPETALUM X Curtisii 44 |GOWERIANUM 19 16. BLEPHAROPETALUM X Dayanum 26 |LITTLEANUM 0 17. THIOPETALUM X Druryi 35 |CYBELE 0 18. NEUROPETALUM X exul 20 |Тотла 0 19. CERATOPETALUM X Fairrieanum 85 |STREATHAMENSE 0 20. Subg. BRACHYPETALUM X Godefroyae 31 [Don CARLOS 0 21. PHACOPETALUM X GowERIANUM (App. Chart II, #6) 19 |LAURE-GOWER 0 22. NEUROPETALUM X GnacraE (App. Chart III, #5) 2 |GRIGNA 0 Subg. BRACHYPETALUM 23. PHACOPETALUM X HaRRISIANUM (App. to Chart Ш, #2) 72 |сїсл$ 8 NEUROPETALUM 24. STICTOPETALUM X bhirsutissimum 33 |Muras 0 25. PHACOPETALUM X Нотреми (Chart I, #20) 6 |PAULIAE 0 26. SPATHOPETALUM x Hookerae 26 |ENFIELDENSE 2 27. NEUROPETALUM X insigne 148 |UMLAUFTIANUM 0 28. PHACOPETALUM Х lo (App. to Chart Ш, #6) 15 |VANNINII 0 29. NEUROPETALUM X Kine Актнук (App. Chart ІП, #7) 9 Евг, KING 0 PHACOPETALUM 30. CYMATOPETALUM X | LATHAMIANUM (App. Chart II, #9) 39 |PvNAERTII 0 NEUROPETALUM 31. NEUROPETALUM X LrEEANUM (App. Chart Ш, #3) 132 |MAGNEI 0 CYMATOPETALUM 32. NEUROPETALUM X LowcwoopENsE (App. Chart ІП, #9) 18 VENIZELOS 0 CYMATOPETALUM *See footnotes Chart II for explanation. 1949] McQuaApE—Pabbiobedilum Maudiae Hort. 463 Section of OTOPED DILUM (or other subgenera) sed in cross Species or hybrid used as parent Resulting 33. PARDALOPETALUM ? . BLEPHAROPETALUM . Subg. BRACHYPETALUM ALUM 5РАТНОРЕТА . PHACOPETALUM . SPATHOPETALUM NEUROPETALUM . Subg. ANATOPEDILUM Sect. CORYOPEDILUM PHACOPETALUM . CERATOPETALUM CYMATOPETALUM . NEUROPETALUM . Subg. BRACHYPETALUM . PHACOPETALUM S A ubg. ANATOPEDILUM Sect. CORYOPEDILUM . PHACOPETALUM NEUROPETALUM . MYSTROPEDILUM . Subg. ANATOPEDILUM Sect. CORYOPEDILUM . PHACOPETALUM . PHACOPETALUM SPATHOPETALUM NEUROPETALUM . Subg. BRACHYPETALUM Subg. ANATOPEDILUM Sect. GONATOPEDILUM Subg. ANATOPEDILUM Sect. Con YoPEDILUM . PHACOPETALUM Subg. ANATOPEDILUM Sect. CORYOPEDILUM BLEPHAROPETALUM PHACOPETALUM . CYMATOPETALUM . Subg. ANATOPEDILUM Sect. CORYOPEDILUM . PHACOPETALUM . PHACOPETALUM . PHACOPETALUM BLEPHAROPETALUM . BLEPHAROPETALUM . SPATHOPETALUM . PHACOPETALUM NEUROPETALUM . NEUROPETALUM x x x x X X x x x x x Lowii 20 L'vsER 6 Mastersianum 24 MARSHALLIANUM (App. Chart III, 210) 1 MauniAE (Chart I, #2) 35 MEASURESIANUM (App. Chart III, #11) 4 MonGANIAE (App. Chart III, #12) 1 „а Мове (App. Chart Ш, #13) 28 NITENS (Chart I, #3) 66 niveum 48 NuMa (App. Chart III, #14) 2 OENANTHUM (App. Chart III, #2) 28 Parishii 5 philippinense 22 Purocroprs (App. Chart ІП, #15) 1 PoLLETTIANUM (App. Chart III, #16) 7 РзуснЕ (App. Chart III, #17) 15 Rothschildianum 53 Sanderianum 13 SELLIGERUM (App. Chart ІП, #18) 21 SEMENTA (App. Chart II, #10) 8 Spicerianum 62 Stonei 27 superbiens 39 SUPERCILIARE (App. Chart III, #15) 18 SwANIANUM (App. Chart III, #19) 13 tonsum 33 venustum 29 VERNIXIUM (App. Chart III, #20) 5 villosum 60 IMACFARLANIANU (1f "тутр ET x ALAIN GERBAULT 6 WILLIAM MATTHEWS 1 HENRY GRAVES 0 ALMA GAVAERT 13 HEBE 0 VENETIA 0 WELLESLEYI 2 JOHNSONIANUM 0 ANTIGONE 2 STANDENSE 0 Втуоц 0 ELIZABETHAE 0 CHARLES STEINMETZ 0 NIGRUM 0 FABIA 0 CONOPUS 0 | WIERTZIANUM 0 ULTOR 0 LADY LLANGATTOCK 0 CRASSIFOLIUM 0 RADIOSUM 2 NUMA 2 EURYALE 5 AUGUSTUM 2 ROGERSII 0 МАРАМ ВАКВЕУ 0 AUROREUM 2 JULIEN COFFIGNIEZ 0 LURIDUM ___ 0 123 [Vor. 36 464 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN APPENDIX FO CHART ПТ V/LLOITUTI NEUROPETALU 14 CARDINAL HERCIER = А LATHAMUANUM HYBRIO <— АМ” CYMATOPETALU ¿Z FAIRRIEANUIT CERATOPETA 4 CAPPAMA BNA NIOBE << у, "c гек уна ПА ГОРЕГАСИТ NUBIA OLUM NEURO PETALUPT СУМАГОРЕЛАХ U ^7 TWOMPIQNI ur SPICERIA —— CALYOLO ee hs. mos ане WEURDPETA LUP HOOKERA ғ BARBATUST ALOPETALY ғ 2. CLEOPATRA < US 777227754 WEUROPETALUP ФЕМАМТ HUP "m ыру INSIGNE NEURIPETAL U PT ACTAEUS < INIIGNE NEUROPETALUY CHRISTOPHER Sor SPICERIAN UM _ __ CYHATOPETALUM INSIGNE EUROPETALU I 4 COLUn BU UAM ESL он ру CYMATOPETALUM INIIGNE NEUR OPE, 7 NITENS ——— урина NEUROPETALU MH MITENS- 44 АМУ / WEURQPETALU P FEN р EPCOT уттук СИМАТОРЕТАШИУ сог нен еа ныи PHACOPETA 4 GOVERIANUST — 24 s NUI ОНА СОФЕТА LUI ФохА "ry (VALOSUM VAR. BOXALLI) EUR OPETALU PI — l. F. SRAACEA& € € U^ Е SUBGENUS ANINE TALUS 4.70 ——— l PETAL U Ë zu са Per NUM mas COPETALU AT z ORTH NEUROPETALUTT OINGLEVYENTSE k ечи м ———— BGARBAT UST PHALOPETALU TT 4 ж 7 KING ARTHUR = НАМА ПА меге x CANAL О, мк өңді; АРҒА АЕ MEVROPETAL rt жол” DE CURTE "UMS жыка а но а найы ы опе VAR. BOXALL on ыйы MEYROPETALU IT CERAM д СУКА у. 8. LATHAMIANU YY ———— /2/ V/Z2 2707 ЕЕ MN INE SEMINA ES tst WEURQPETALUPT ORTHI/ AFURODETALUP CHIAPLEJM. —— Ci РАСА ENS Я LONGWOOO Е азай 77/6 NE — WEUROPETALUP^T £££AWUP SPICER/A УТ I сута йы еу Ад CONCOL vip SUBGENUS BRACHY, TALS 10. МАМА ЕАМ —— Ашы ы VENUZTU JPATWÜPETALUM VENUITUM JPATHOPETALU MW AIEASURESIANUM —— с ————————— ч VALEAMUS әх» DETALUM STONES “ғ Фенг ANATOPEO/LU M 12, SIQRGANIAE SECT. LIRVOPE DIL UM SUPER BIEN S PHA PD£FAL UH CERATOPETALU M FAIRRIEANU II 2. NIOBE ———— 3E тесе АМО” — I mee $$ TOD ETAL UN Айла АСА U S UUO PWACOPETA LU ПОЕ ——— C /JUBSENUL ANATOPEOILUM “cr PHACOPETALU/ /. ^7 — V ена. NE ——— ?9WACODETAIM 4UPERCIITARE ——— ұстынын, A — — a масон л ^f PHLOGIODES PROBABLE — š: Ë i ^48ATU^ CAL OPHYLLUM 18. POLLETTIANUM venus Tut ЈЛАвГУОВЕА ИТ BATU PWACOPETALUM OEFMANTMH UM ucc EL ore ALUN INIIGNE WEURDDETA UP их PLYCHE чес O44 AT ULUI SUBGENUS BOACHAY, 4744 ^T NIVEDUP SUBSEENUS BRACHYPETAL ИМ BAREATUM 48. SELL) ————— 8. J£44/0€ еи PERIPDINENÁE SECTS 24, ^. Е me iibi дый, Un a PHACODETALUI rere OL PNLRODETA Шу 22. VERNIXIGH диам PETALUMg е” via < бит ———— NEUROPETAL [Vor. 36, 1949] 466 ANNALS OF THE MISSOURI BOTANICAL GARDE EXPLANATION OF PLATE PLATE 31 Flowers about №; habits about 14. Figs. 1 and 2. Paphiopedilum callosum. Figs. 3 and 4. Paphiopedilum Maudiae var. magnificum. Figs. 5 and 6. Paphiopedilum Lawrenceanum. WOATIdAdOLHdVd—AGVAOOW AVIGQAVW “LYOH ANN. Mo. Bor. GARD., Vor. 36, 1949 PLATI [Vor. 36, 1949] 468 ANNALS OF THE MISSOURI BOTANICAL GARDEN A. MTM a т — МЈ — — EXPLANATION Or PLATE PLATE 32 Anaphase in root-tip of Paphiopedilum callosum, X 1350. Aceto-lacmoid. Anaphase in root-tip of P. Lawrenceanum, Х 1350. Feulgen. , Metaphase of root-tip of P. Мамас, X 1350. Feulgen. The two homologues showing the inversion are marked. Camera-lucida drawing of the chromosome pair showing an apparently terminal in version. These chromosomes are of intermediate length and have a prominent secondary constriction. The centromere is subterminal in one and median in the other (centromere marked with a line). Magnification X about 1800. P. callosum, Metaphase 1, 16 bivalents, X 1350. Crystal violet and safranin. Р. ТЩ Metaphase ПІ, 18 chromosomes, X 1350. Crystal violet and safranin. ANN. Мо. Вот. Сакр., Vor. 36, 1949 PLATE 32 McQUADE—PAPHIOPEDILUM MAUDIAE HORT. | VoL. 470 ANNALS OF THE MISSOURI BOTANICAL GARDEN 25( EXPLANATION OF PLATI PLATE 33 Pollen of Paphiopedilum callosum, of P. Lawrenceanum, and of P. Манас, ); 36, 1949 | about ANN. Mo. Вот. GARD., Vor. 36, 1949 PLATE 33 P. Maudiae McQUADE—PAPHIOPEDILUM MAUDIAE HORT. [Vor. 36, 1949] 472 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 34 Paphiopedilum Maudiae Hort. А. Pachytene-early diplotene, X about 1350. В. Metaphase I, 17 bivalents, X about 650. C. Anaphase I, bridge and fragment, X about 650. D. Metaphase II, Х about 750. E. Tetrads (note micro-grain), X about 700. ANN. Мо. Bor. Garp., Vor. 36, 1949 PLATE 34 McQUADE—PAPHIOPEDILUM MAUDIAE HORT. STEGNOSPERMA: А NEW SPECIES AND A GENERIC COMMENTARY DAVID J. ROGERS The genus Stegnosperma (Phytolaccaceae) has been considered monotypic since Walter's treatment for Engler's "Pflanzenreich'. However, an examination of specimens in the major North American herbaria shows the inclusive species 5. balimifolium Benth. of Walter to be rather heterogeneous. Actually, three species exist, two of which have been described and published, the third noted by S. Wat- son on an herbarium label but never published. А description of the third species is provided here and is named for Dr. Watson. STEGNOSPERMA Watsonii D. J. Rogers, n.sp. Frutices aut scandentes aut crassi patulique 1—5 m. alti, 1-5 m. diam., cortice griseo vel rufo-brunneo. Folia anguste spathulata vel elliptica emarginata vel rotunda vel acuta 1.0-3.5 cm. longa 0.5—2.5 cm. lata, petiolo 0.1—0.3 cm. longo. Inflorescentia cymulis axillaribus aut terminalibus 1—8-floris; calycis lobis ellipticis vel ovatis 0.3—0.7 cm. longis 0.2-0.4 cm. latis; petalis ovatis rotundatis basi abrupte constrictis; fructu capsula 5- loculata plerumque in 5 valvis dehiscente; seminibus plerumque 5 aliquando 4 ovoideis vel ellipsoideis circa 0.3 cm. longis 0.2-0.3 cm. latis, cicatrice funiculari laterali, raphe in jugum dorsalem, testa levi fulgenti rufo-brunneo. Sprawling vine or coarse spreading shrub, 1—5 m. tall, 1-5 m. diameter spread; bark gray to reddish brown. Leaves narrowly spathulate to elliptic, emarginate to rounded to acute, 1.0-3.5 cm. long, 0.5—2.5 cm. wide, petiole 0.1-0.3 cm. long. Inflorescence of axillary or terminal 1- to 8-flowered cymules; calyx lobes elliptic to ovate, 0.3—0.7 cm. long, 0.2-0.4 cm. wide; petals ovate, rounded, abruptly con- stricted at base; fruit a 5-celled capsule, usually dehiscing by 5 valves; seeds usually 5, occasionally 4, ovoid to ellipsoid, about 0.3 cm. long, 0.2-0.3 cm. wide, funicular scar lateral, raphe on a dorsal ridge, testa smooth, shiny, reddish brown. XICO: BAJA CALIFORNIA: Wiggins 7681. SINALOA: Jones s.m. SONORA: Abra Prin Coville 1646; Dawson 1058; Drouet, Richards 8 Alvarado 3443; Ferris Srat: Gentry 2195, 2975; Goldman 399; Keck 4067; LeRoy s.n.; Lumboltz 9; McGee s. n.; William кеші 1226 (н Е іп wee Missouri Botanical Garden, isotypes in Herb. N. У. Bot. Gard. and U. S. Nat. Herb.) ; ngle s. n.; Rose I211, I211a; Rose, Standley 8 Russell 12390, 12566, 13138, P Ede ye 5092; Wiggins 6247. This species seems to be most closely related to $. halimifolium Benth., from which it may be distinguished by its scattered, few-flowered cymules, its ovate, abruptly constricted petals, and by its lateral funicular scar. Stegnosperma Watsonii grows on hillsides along rivers, thickets in palm groves, thorny foothills, from sea level to 300 meters. It flowers from about the first of February through March, and fruits from the last of February through April. That there are actually three species of Stegnosperma is most easily demonstrated by the following key: Walter in Engl. Pflanzenr. IV, 83:124. 1909. Issued November 30, 1949. (475) [ Vor. 36 476 ANNALS OF THE MISSOURI BOTANICAL GARDEN A. т” а terminal, many-flowered racemiform thyrse; petals rather gradually narrowed to the base B. Sepals lincar to elliptic, petals ve to spathulate; capsule dehiscing by rarely 5 valves, «prd мш 1 to 3 seeds; е with a lateral to sub- bal E scar, рони п а flattened do рерна ВВ. -— ovate; hn ovate to elliptic; PIRE dehiscing by 5 valves, ic with 5 eeds; seeds with a basal funicular scar, the raphe on a dorsal ridge S. balimifolium AA. tae an axillary, 1- to 8-flowered cymule; petals abruptly constricted at the base S. Watsonii cubense STEGNOSPERMA CUBENSE A. Rich. in Sagra, Hist. Nat. Cuba 10:309; 12:/ab. 44°. [Tricbilia] scandens, foliis simplicibus, ovatis alternis, екс. A. Robinson ex Lunan, Fl. Jam. 2:519. 1814. Trichilia scandens Lunan ex B. D. Jackson, in Index Kewensis 2:1105. 1895. Based on the preceding. Stegnosperma scandens (Lunan ex B. D. Jackson) Standley, in Field Mus. Publ. Bot. 23:6. 1943. Stegnosperma halimifolium of authors, not Benth. XICO: TRES MARIAS ISLANDS: Fisher s. n.; Howell 10409; Maltby 45; Mason 1702; РЕА 4185; Solis 4, 22, 45. REVILLA GIGEDOS ISLANDS: ason 1846. siNALOA: T. 5. Brandegee s. п.; Eyerdam & Beetle 8652; Lamb 465; ipi 45, Te 1005; Ortega 4480, 5150, 5649, 6453, 7232, 7488; E. Palmer 1503; Rose s. n., 1535; Rose, Standley & Russell 13721. NAYARIT: Ferris 5300; Nelson 4340. COLIMA: Goldsmith 00; Jones 13; E. Palmer 1280. MICHOACAN: Ranges 2627; Leavenworth & Hoogstraal 1304. MEXICO: Hinton 3704. GUERRERO: on 5431, 5719, 5062. oaxaca: Matuda 0664; Nelson 2507; Orcutt 3307. CHIAPAS: p 2608; Morley 710. VERA CRUZ: Purpus 8050, 10080, 13066. GUATEMALA: ESCUINTLA: Salas 376. SAN JOSE: Worth, Morrison & Horton 8632. RETALHULEU: Standley $7707. SAN MARCOS: Steyermark 37762, 37773, 37881. SUCHITEPEQUEZ: Steyermark 47825. ?АСАРА: Standley 74066; Steyermark 42084. EL SALVADOR: LA LIBERTAD: Standley 23210. NICARAGUA: CHINANDEGA: Baker 2065. MANAGUA: Chaves 262; Garnier 1071; 5 ABANA: Ekman 13403. PINAR DEL RIO: Baker & Van Hermann 4247; Ekman 13039, 10749; Leon 9 Коса 7132, 8810; Shafer 2. 11148; Wilson 11400, 11404. WITHOUT LOCALITY: Wright 20. рој vicinity of Spanish Town; Britton 3062. Healthshire Hills, Harris & Britton 10522. Dominican REPUBLIC: Beata Island, Fairchild 2605, 2606, s.n.; Ostenfeld 319. Massif des Cahos, Ekman Н0005. BARAHONA: Ekman HÓQÓI. SIERRA DE OCOA: Н13360. WITHOUT LocaLiTY: Bertero Puerto Rico: Asomante, Horne 9 Britton 9628. This species, although placed in synonymy under S. halimifolium by Walter?, is sufficiently distinct to be maintained. The description and plate provided by Richard demonstrate its characters accurately. Further characters which support my interpretation are found in the seed. These characters are used in the key. An interesting nomenclatorial problem concerning this species arose when Standley? made an apparently valid transfer of a "species" of Trichilia ascribed to Lunan by B. D. Jackson in ‘Index Kewensis'*. Lunan’s "publication" of A. Robin- * Walter, | Standley, in тыа Mus. Publ. Bot. 23:6. 1943. D. Jackson in Yandex Kewensis 2:1105. 1895. 1949] КОСЕК5--5ТЕСМО5РЕКМА 477 son's manuscript description of a new species of Trichilia was as а polynomial. Jackson's interpretation of the name as a binomial possibly could be explained by a method used in early printing in which the first word of a page was placed at the bottom of the preceding page on a line by itself. In this case, the first word to appear in the first sentence at the top of page 320, Lunan's Fl. Jam. Vol. 2, and accordingly at the bottom of the preceding page was “scandens.” I do not think that Lunan intended a binomial since he did not mention "scandens" as a species in his Classical Index of this work although other properly published binomials are listed, nor did he use the same form in his discussion of the plant in question as he consistently used throughout the text for species designation. STEGNOSPERMA HALIMIFOLIUM Benth. Bot. Voy. Sulph. 17:}/. 12. 1844 (as halimi- $i BAJA CALIFORNIA: T. S. Brandegee s. n.; Carter d Carter, nie : m pr 2II5, 2497; Collins, , Kearney © Kempton 186; Epling & Robison s. n.; Ferri 8617; Gentry 4032, 7603, 7864; Hammerly 102; Harvey 609; Johnston 3166, 552 5488, 3512, 3503, 3825; Jones 24481, 2/465; Nelson & Goldman 7147, 7249, 7323, 7395, 7502; Palmer 31, 258, 400, 870; Purpus s. n., 5; Rose 16289, 16415, 16616, 16690; 16924; Еф Shreve 6073; Wiggins 5415, 5051, ` боо, 7703; Xantus 9b. sonora: MacDougal 6 Shreve 40, 47; Pringle s. n. The generic ending of Bentham's specific epithet has been altered to comply with the International Rules of Botanical Nomenclature, Section 14, Art. Z2 (2). As I have interpreted this species, it occupies a rather narrow range in Baja California and occasionally in the adjoining state of Sonora, Mexico. I have been able to examine specimens from several of the major herbaria of the United States, but have not seen the type specimens nor any other material from Europe. The herbaria where specimens have been obtained for study аге as follows: Gray Herbarium of Harvard University, Chicago Natural History Museum, Missouri Botanical Garden, New York Botanical Garden, University of California at Berkeley, and the United States National Herbarium. I wish to acknowledge my indebtedness to the curators of these institutions. NUCELLANGIUM, А NEW GENUS ОҒ FOSSIL SEEDS PREVIOUSLY ASSIGNED TO LEPIDOCARPON HENRY N. ANDREWS, JR. Among the more abundant fossils found in the Iowa coal balls are the highly unique "seeds" which have been named Lepidocarpon glabrum. These were de- scribed by W. C. Darrah in 1941, and in a more recent publication (1949) the same author has continued the discussion with descriptions of included structures which are claimed to be gametophytes and embryos. It is the purpose of this paper to add somewhat to the information given in the published accounts, to point out what appear to the present writer as erroneous statements of fact, and to correct the corresponding conclusions. The fossil is not referable to any known genus, and a new generic name, Nucellangium, is proposed herewith for its reception. Origin of tbe specimens.— The specimens on which the present descriptions are based were collected by Mr. Frederick O. Thompson from the Urbandale coal mine located on the western outskirts of Des Moines, Iowa, the exact location having been given in the results of a previous study from this laboratory (Andrews and Kernen, 1946). It should be noted that these specimens and the ones described by Darrah come from the same locality, and through the cooperation of Dr. Elso Barghoorn I have also been able to study a series of similar preparations from the Botanical: Museum of Harvard University. There is, therefore, no possibility of confusion in the identity of Darrah's specimens and the ones on which this account is based. The material is from beds of Middle Pennsylvanian (Des Moines) age; un- fortunately the precise stratigraphical equivalence of the Urbandale coal is not known but presumably the material is a little younger than floras known from Illinois No. 6 coal or from above the upper Freeport coal of Pennsylvania. General introduction to tbe nature of tbe fossils.— We are involved in this discussion with two sets of fossil plants, the first being ovoid bodies presenting certain anatomical characters which lend some justification to their being considered as seeds, and the second less regularly shaped bodies with highly distinctive convolutions extending into their interior which are alleged to represent gametophyte and sporeling. Certain competent botanists who have examined the fossils іп my collection have expressed doubt that the two phases or forms belong to the same species. Mr. Darrah has based his case on the supposition that they represent different growth stages of the same organ. I agree with him to that extent yet it must be remem- bered that it is not beyond the realm of possibility that we are wrong in this belief. It seems most convenient to refer to these two forms as proliferated and normal depending on whether they do or do not contain the supposed sporelings. In view Issued November 30, 1949. (479) [Vor. 36 480 ANNALS OF THE MISSOURI BOTANICAL GARDEN of the incomplete nature of the previously published accounts it will be necessary to present rather detailed descriptions of the two phases. I have little doubt that many morphologists will take issue with the usage of terms as they are applied to these fossils. It is becoming clear, however, in groups such as the psilophytes and early coenopterid ferns, that the fossils are not going to make a special effort to comply with our preconceived terminology. It is hoped that the following pages contain descriptions that may be readily comprehended, but I believe that these fossils present structures which do not correspond precisely with known morphological entities. Insofar as the evidence allows it seems clear that the fossils are sporangia that may or may not have been integumented. It is not known how they were borne on the parent plant and Darrah's restoration of the "strobilus" (1949, fig. 39) is based, so far as I am able to judge, on the supposed general lycopod affinities of the fossil rather than on conclusive evidence. There is a trace of conservatism in the caption to that figure which reads, "Sporophylls not sufficiently known to warrant reconstruction." The fact is that nothing whatsoever is known of the supposed sporophylls. The general organization of the fossil, with its vascularization and complicated wall structure, seems to allow a closer comparison with a cordaite seed than with a lycopod sporangium. We shall return to such speculations on a later page. The fossil will be referred to in the following pages as a sporangium, as a nucellus, or as ап (unintegumented) seed. Тһе last term is used advisedly and as a matter of convenience, although it seems probable that at least the specimens containing a "seed megaspore" did function as such. It seems most expedient to present first а detailed description of the "normal" fossils and then consider the morphology of the principal structures involved. T he “normal” seeds.— These are very abundant in the Urbandale coal balls as well as in those from other localities which probably represent the same or a close horizon. In many of the coal-ball specimens examined a half dozen or more are exposed in a single saw cut and, due to the distinctive structure and preservation of the epidermal layer, they are often partially exposed on the broken surfaces of the petrifactions. It is occasionally possible to isolate the seeds intact from the surrounding matrix. While the following description is supplemented by observations on dozens of specimens it is based primarily on a series of transverse sections prepared through a single specimen. Although there is some variation in the size of the specimens it is not great. They are broadly ovate (figs. 1, 2), averaging 12 mm. long, and in the median region the large and small diameters are 9.5 by 6 mm. Many specimens, particu- larly the more poorly preserved ones, are crushed and distorted, yet there can be no doubt that the shape and dimensions as given here represent the life form of the seeds. 1949] ANDREWS—NUCELLANGIUM 481 At one end of these slightly elongated structures, which we will refer to as the proximal end, there is a tiny circular "hilum" scar (fig. 5) representing the point of attachment. Ас the other end, which will be referred to as the distal end, the seed tapers to a blunt point. Fairly conspicuous ridges lead to this point from the median region of the seed, following the narrow lateral faces. ‘The specimens shown in figs. 1 and 2 present the broad side of the seed and the ridges here form the outline of the photo of the upper half of the seed. The hilum scar may be seen at the proximal end in fig. 1 and the blunt point at the opposite extremity. A series of peel preparations has been made by first carefully smoothing a flat surface at the hilum end. Seventy successive peels were then made to within less than a half mm. of the distal end. Particular саге was taken to obtain a nearly perfect serial series at the hilum end in order to trace accurately whatever vascular system might be present. When it became evident after working through about one quarter of the length of the specimen that sudden changes in anatomy were no longer taking place the sections were taken further apart in the median region. It is perhaps apparent that this is a case in which the peel technique is quite indispensable, for it would be only through the greatest good fortune and the use of numerous well-preserved specimens that somewhat comparable results could be obtained by reliance only on ground sections. It is probable that if the specimen had been properly imbedded even better preparations could have been made. How- ever, they were generally removed with little difficulty by using a sharp razor under the low power of a dissecting microscope. Occasionally the epidermis was par- tially destroyed but since this remains constant in structure from one end to the other there was по loss. Text-fig. 1 indicates the approximate position from which the respective peels were taken. To present an effective description of Nucellangium this series of preparations will be followed from proximal to distal end. It may be an aid in following the discussion to note at the outset that three characters set this fossil apart from previously described species of Lepidocarpon. "These are: a well-developed vascular system with two strands running nearly the entire length of the seed; a thick complex wall including an inner sclerotic layer; and a mode of attachment unlike that of the radially elongated sporangia of other species of Lepidocarpon. This combination of characters, and particularly the vascular system, clearly prevents the inclusion of the fossil in that genus. In the first peel prepared, which does not quite reach the inner limit of the epidermis, the central vascular strand may be distinguished. It is circular in transverse section and is composed of a considerable number of conducting ele- ments (fig. 8). It is apparently purely tracheidal, no parenchyma cells having been observed. The conducting elements of this basal strand, as well as those of the lateral traces, are distinctive in that they are thin-walled, follow a slightly sinuous course, and the bands composing the secondary thickenings are fine and delicate, a condition, judging from the generally good preservation through the specimen, that is natural and not the result of decay. It is not possible to determine [Vor. 36 482 ANNALS OF THE MISSOURI BOTANICAL GARDEN whether the secondary thickenings were of a typical annular or scalariform nature; if the latter, it seems evident that the border of the thickenings was not strongly developed. 12 11 10 Ра 9 ZL "A ZI ШЕ 6 _ s 4 | A NN 2__\ 1 № 0 онај xt-fig. 1. Diagrammatic longitudinal view Tex Diagrammatic media presented as an aid in following the description longitudinal 5 section Bae the major of the series of transverse peels described on axis of the seed showing the entrance of e accompanying Pages. Figures at left are the vascular strand at the base and the millimeters; figures at right represent peel cour o branch traces through numbers. the eas y a wall. At peel No. 4 a disturbance of the thin-walled parenchyma surrounding the strand suggests a departing trace and when peel No. 6 is reached the tracheids of a branch trace may be observed, not actually leaving the strand but more than half way to the periphery of the seed. Ас first it was thought that the point of de- parture had been missed but in the next peel the trace was noted both departing (fig. 6) and in the outer region, as noted above. It is evident, as shown in fig. 10, as well as in text-fig. 2, that the trace dips down slightly after leaving the central strand to follow its course up through the seed. In peel No. 7 the departure of а second trace appears on the opposite side of the central strand. No other branch traces were observed although a careful search was made, since in his original description Darrah (1941) notes, with reference to the vascular system: "At the proximal end of this seed-like sporangium there is а vascular trace which forks twice, but the four branches quickly exhaust themselves. The bifurcations are at right angles to each other, and by serial sections it has been observed that the two forkings take place one above the other." (p. 97). The presence of these vascular strands, as I have described them, is of the greatest importance since they are typically absent from lycopod sporangia. It is understandable that they might be readily overlooked in longitudinal sections but 1949] ANDREWS—NUCELLANGIUM 483 I cannot feel that there is adequate excuse for failing to observe them in the serial sections Darrah indicates were prepared (1941, p. 85). The pair of traces continues to within less than one half mm. of the distal end of the seed and may be clearly observed in most of the peels throughout the length of the seed; and in the specimen described here there is no suggestion of a second pair of traces. А possible explanation of the apparent departure of such will be offered below. It is clear from all of these transverse peels, with the exception of the basal two or three, that the seeds are bilaterally symmetrical in their anatomy as well as in their gross external form. Taking as an example a nearly median section (fig. 16) the ovate form of the fossil is evident and the two traces may be seen at either end of the great diameter, the traces in this peripheral region occupying a position in extensions of the inner sclerotic layer. Figure 7 shows the trace rather well at a point where it and the surrounding tissues are quite well preserved. For the purpose of considering the extra-vascular structure of the seed a nearly median point will be taken where a typical sequence of the tissues is displayed. Selecting peel No. 34 (figs. 12, 16) the following may be clearly defined: The thick-walled, palisade-like epidermis (fig. 9) forms the outer cell layer of the seed over its entire surface with the exception of the hilum scar at the base. These cells are arranged with their long axis approximately parallel to а radius of the seed. They are uniform in size and shape, being about 125 м long and, when observed in surface view, about 25 џи т diameter. These cells also present an interesting preservation problem. They seem immune to the action of hydrochloric acid, unsuccessful attempts having been made to etch the outer face in order to obtain surface peels. Apparently they are little, if at all, mineralized. It is not surprising that with such an external tissue, so seemingly resistant to an infiltrating mineral solution, the more delicate internal tissues are poorly preserved in most specimens. Within this epidermal layer is a broad zone of nearly isodiametric, rather thin-walled cells (the outer parenchyma, o. p. of figs. 12, 16). Ав may be noted in the photos this tissue comprises a major portion of the sporangium wall as a whole. There is a tendency for approximately the outer third of this tissue to have somewhat thicker cell walls than the inner region although there is no sharp distinction into two zones. It is highly significant to the discussion of the morphology of the seed to note that this is clearly ¿z con- tinuous tissue connection with the conspicuous columnar epidermis. Forming a third layer is a very prominent, dark and semi-sclerotic tissue (inner sclerotic layer, i. s. of figs. 12, 16). Тһе term “sclerotic” is perhaps misleading although the cells are somewhat thicker walled than those of the outer parenchyma. It will be noted (fig. 12) that the cells of this tissue increase appreciably in diameter towards the inner periphery, and they are longitudinally elon- [Vor. 36 484 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN gated, being at least twenty times as long as they are broad. Where the preservation is good, and nearly perfect longitudinal sections are obtained, the cell walls appear to be strongly pitted. The end walls are transverse or only slightly oblique. The pits (fig. 4) are generally more or less oval-shaped and apparently simple, but whether an actual membrane separated one cell from the next in life cannot be determined. The pitting in some cells is more complex and may even approach reticulate banding. In certain of the more roximal sections in the series of peels taken through the specimen some of the cells of this tissue resemble the tracheidal cells of the traces. It is my suggestion that Darrah may have mistaken these for the second pair of traces mentioned in his account. The abundant pitting of these cells and their great length as compared with the other non-vascular tissues suggest that their primary function was the conduction of fluids. In studying the seed from base to apex the pres- ence of this tissue is first noted at the level of peel No. 7. From this point to approximately peel No. 11 it develops in abundance in two separate groups, sheathing the departing traces. The two groups soon expand in two C-shaped masses until they unite as a continuous band at the level of peel No. 29. Тһе radial width of this band thereafter gradually increases as is shown in text-fig. 3. Within the sclerotic layer there is a fourth tissue consisting of very thin- walled cells which in most specimens has been lost through decay. In a few instances, however, it is possible to observe that this tissue did consist of rather thin-walled parenchymatous cells. Within this fourth tissue layer it is possible, in most sections of the series, to follow a distinct, light yellow band around the periphery of the central cavity. The presence of this structure has been checked in num- erous other specimens, and there seems to be no doubt that it is correctly identified as the megaspore membrane. Darrah has succeeded in isolating it very nicely by maceration, a fine illustration being given in his 1949 paper (fig. 11). No tissue has been observed within this megaspore membrane. The shape of the internal cavity will, however, be described briefly. In following the series of transverse sections from proximal to distal end, at peel No. 10 a small cavity ap- peared in the position occupied in previous peels by the trace and its accompanying tissue. In peel No. 15 a similar cavity made its appearance on the opposite side and in peel No. 17 the two cavities merged. Тһе fact that one cavity appeared before the other is due to a very slight obliquity in the internal structure of the seed, possibly a slight aberrancy of the particular specimen. It thus appeared from a study of the serial sections that the internal cavity would be heart-shaped if viewed in median longitudinal section. It has been pos- sible to confirm this supposition from such a nearly median section in the collec- tions of the Harvard Botanical Museum. The basal portion of the specimen is 1949] ANDREWS—NUCELLANGIUM 485 shown in figure 10. This is a trifle oblique to the median plane so that the actual entrance of the trace into the seed is not shown. "Тһе trace does appear, however, as a conspicuous mass of tracheids (fig. 10, t), flaring upwards and terminating the upper part of the cushion or "archesporial pad." A portion of one of the two traces is shown curving down and upward to the left in the outer parenchymatous tissue. . А series of diagrams prepared from representative points in the series it transverse pees described on pages 480-484. 1, peel Хо. 1; 2, peel No. 3; 3, peel No. 8; 4, pee 2055; Мо. 38 eel No. 53; 7, и No. 64; 8, peel No. 66; 9, peel No. 69. 4% жы агеа, epidermis: st tipple, о outer parenc chyma; cross-hatch, inner sclerotic tissue; inner stipple, inner paren- о rane. Іп 1 only t i |" де ме о ы < D og ы [^] 3 B о 8 — о B. о „ЕВ 873. с. w oO ч B = ы в a "n P, 172 et la ю 5 с. m ”" % р ап г chyma, central г. and beginnings of the inner sclerotic tissue; 4-6 Present the complete sequence of tissues with the traces shown at either end of the long transverse axis; 7 is taken above the distal limit of the megaspore [Уог. 36 486 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN Before leaving this description of the normal seeds I feel compelled to add а comment on Mr. Darrah's recent paper in which he sums up the distinctive features of these fossils: "Thus far in the development of the sporangium there are no structures or tissues which are unusual. Externally, there is the sporangium wall of usual lepidodendrid construction innermost the megaspore, which can be removed easily by maceration. The tissues between are sterile sporogenous tissues." (1949, p. 3). Ido not understand the last sentence but to state that the sporangium wall with its thickness, its complex series of tissues, its vascularization, and its mode of attachment is “of usual lepidodendrid construction" certainly displays a taxonomic freedom that might allow the inclusion of anything within the genus Lepidocar pon. Darrah notes further that "My preference for broad rather than narrow inter- pretations of genera is well known." (p. 12). But surely somewhere there must be limitations. I feel certain that there is nothing in the literature of lycopod sporangia, living or fossil, which presents a close comparison with this fossil. It is clearly far beyond the bounds of Scott's generic description for Lepidocarpon (Scott, 1901) and bears no resemblance to L. lomaxi, the type of the genus. It is perhaps obvious that the principal problem that is involved in correctly interpreting the morphology of this fossil is whether we are dealing with a spor- angium or whether it is a true зеед. That is, whether the structure described above is a sporangium ("nucellus") enclosing a single fertile megaspore, or whether it is an integument enclosing the remains of a nucellus and the megaspore. I am of course following Darrah's interpretation in accepting the former choice. The reasons for this are as follows: There is no break in the continuity of the four tissues composing the wall of the fossil. They are all clearly in organic connection, there is no delimiting epidermal layer on the inside, and between this innermost parenchymatous layer and the megaspore there is no structure that might be in- terpreted as the remnants of а nucellus. Furthermore, there is no evidence that a micropylar opening existed at the distal end of the fossil. The tissue appears to be continuous here, allowing access of microspores only by a dehiscence of the sporangium, presumably along the lateral ridges. Although the outer epidermal layer is very resistant, the shape of the cells and their alignment are as closely comparable to the prismatic epidermis of many cryptogamic sporangia as they are to the epidermis of seed integuments. It has long been recognized that the nucellus, in fact, is a modified sporangium and an epidermal layer so strongly suggestive of its sporangial homology is not surprising in a form that, as far as we know, probably lacks integuments completely. The “proliferated” seeds.— Associated with the above-described normal seeds in the Iowa coal balls from the Urbandale mine are other fossils of an even more problematical nature. 1 believe that they present, as Darrah indicates, a different growth stage than that of the normal specimens. It is freely admitted by the present writer that he is certain of neither their natural affinities nor their morphology but evidence will be 1949] ANDREWS—NUCELLANGIUM 487 offered to support the contention that the structures described as "gametophytes" and "sporelings" are morphologically one and the same and that they constitute proliferations of sporangial wall tissue. In the Urbandale coal balls that have passed through my hands some two or three dozens of these proliferated seeds have been observed but, as in the normal ones, a single particularly well-preserved specimen was selected for detailed con- sideration. However, casual study of the other less well-preserved ones clearly indicates that we are dealing with a typical specimen. In view of the unique nature of the fossils the reader is referred to figs. 18 and 19 as an aid in following the description. These are representative peels taken from specimen No. 519. The over-all dimensions as illustrated in fig. 19 are 13 X 10 mm. Extending about half way around the specimen (the lower half as it is oriented in figs. 18 and 19) is an epidermis of heavily thickened palisade-like cells which, allowing for some variation among the individuals, agree exactly in size and shape with those of the epidermis of the normal seeds. Within this epidermis there is a parenchymatous tissue which composes the remainder of the fossil. This tissue consists of rather thin-walled cells; it is organically connected with the epidermis; it is vascularized; and it proliferates out into a central area in the form of branches of varying size. Each of these branches contains a delicate vascular strand and is bordered by a well-defined, thin-walled epidermis (fig. 13) which is consequently quite different from the outer epidermis of the fossil as a whole. It does not seem necessary to comment on the outer thick-walled epidermal layer but a more detailed consideration of the parenchymatous tissue within is very much in order. This consists of rather irregularly shaped cells (fig. 3) in the peripheral region although in the central proliferating arms of tissue (fig. 13) the cells show some tendency to be elongated parallel to the long axis of the arms. It may be noted also in fig. 13 that the epidermis is only slightly differentiated from the interior parenchyma. It is pertinent to indicate at this point the reasons for correlating these fossils with the previously described normal seeds. The former are, as noted above, some- what larger and the epidermal layer is split and does not include the entire structure which would be expected if the normal seeds "germinated" to produce the dis- tinctive proliferations shown in figs. 18 and 19. То me, it would seem most likely that the normal seeds or sporangia opened longitudinally along the ridged lateral edges. However, it has not been possible to determine the mechanics of germination from the available specimens, and I find it difficult to glean satisfactory informa- tion from Darrah's brief description of this point. Like the epidermis in the two supposed growth forms, the outer parenchyma of the normal seeds agrees precisely with that of the proliferated seeds. It is clear that the parenchyma is in organic connection with the epidermis just as the epidermis and outer parenchyma of the normal seeds are organically connected, and it is equally clear, as shown in figs. 18 and 19, that there is no break in this parenchymatous tissue from the epidermis to the inner extremity of the arms. The [Vor. 36 488 ANNALS OF THE MISSOURI BOTANICAL GARDEN latter vary considerably in size, some being apparently simple unbranched structures while others branch rather profusely. In the specimen shown in figs. 18 and 19 there may be noted a rather massive central "clump" which gives rise to numerous branches. It will also be noted that many branches appear unconnected with the peripheral tissue but in following the series of peels many of these are readily ob- served to be connected and I believe that in view of the very close similarity of all of these central islands of tissue (as they appear in an individual peel) there is no reason to doubt that all are so connected. Finally it is important to note that all of these arms are vascularized by a delicate central strand of tracheids similar to those composing the traces of the normal seeds. Although the vascular strands of these are small and composed of few tracheids (fig. 14) the system as a whole is rather extensive. Іп fig. 18 a tracheidal strand may Бе noted at 7 and from this lateral strands branch out into the central arms. Discussion.— In the opening paragraph of his recent contribution Darrah states that “The discovery of well-preserved fossil embryos in a known plant group is therefore an event of considerable interest." There can be no doubt that such a discovery would be enthusiastically welcomed by botanists їп general and paleobotanists in particular, and it is one that may be expected with justification due to the present interest in the coal-ball petrifactions. It is, however, my belief that satisfactory proof of this discovery has not been offered to date. It is not a pleasant task to have to refute the work of a colleague but in view of the seeming importance of these fossils no other course seems feasible. It is very possible that the restorations presented in Darrah's figs. 14—17 and fig. 45 (1949) might well be taken up by writers of text-books and without a first hand knowledge of the fossils it must be admitted that his descriptions are fairly convincing. It is my contention that these restorations showing "Lepidodendroid embryos" within the sporangia are en- tirely unjustified from the anatomical evidence, that the succession of tissues con- tained within the fossils has been misinterpreted, and that the evidence does not support the view that they are of lycopod affinities. I wish to admit freely that satisfactory conclusions regarding the natural rela- tionships of this fossil have not been reached yet. For nearly three years I have pondered over their morphology and affinities and have discussed them with numerous paleobotanists and morphologists. Sincere thanks are due to many of my colleagues for consoling suggestions. These fossils remain as the most prob- lematical ones that I have had occasion to study, but in view of the above- mentioned publications it seemed necessary to present the results of my own observations to date. If future investigations are able to improve on the ad- mittedly vague suggestions offered here they will be received cheerfully. Darrah (1949) has interpreted the conspicuous peripheral parenchymatous tissue as a gametophyte and certain of the central patches of tissue as portions of 1949] ANDREWS—NUCELLANGIUM 489 an embryo sporophyte. The basis for his differentiation of sporangial wall (or nucellar) tissue from gametophyte is not apparent in his illustrations or from his description. On page 3 he notes: "Close examination shows that a gametophyte has developed within the megaspore rupturing it and pressing it against the com- pressed sterile sporangial tissue (remnants of the megaspore membrane can in nearly all cases, be recognized)." And he also notes оп this page that "The game- tophyte is relatively undifferentiated.” And on later pages reference is made to the “тоге or less disintegrated” gametophyte in more mature specimens in which the embryo sporophyte has developed at the expense of the nucellus and game- tophyte. Thus there is an essential conflict in our descriptions because my sections show clearly that the “gametophyte” and "nucellar" tissues are continuous and the same. Furthermore, no remnants of the megaspore membrane can be defined in any of the proliferated seeds I have examined. In the well preserved specimen which serves as the basis for the present descrip- tion there is no evidence of any disorganization of tissue in the peripheral region of the parenchyma adjacent to the epidermis. On the contrary, these two tissues are, as noted above, clearly in organic connection. Тһе general organization and degree of maturity appear to be essentially identical with those described by Darrah. It will be helpful in this respect to compare figs. 18 and 19 with Darrah's figs. 3 and 4. It would seem, therefore, that there is no justification for referring to a tissue organically connected with the epidermis of a sporangium as gametophyte and sporophyte. I can find no evidence for its alleged development within a mega- spore wall. From the prominence of the yellow membrane in the normal seeds there is little question that it would be visible if it were present. It seems especially significant to note that in my specimen the peripheral tissue, that would be termed gametophyte in accordance with Darrah's interpretation, is clearly vascularized. There is no mention of such vascularization in his description, and this oversight may be, in part, responsible for the confusion. In his fig. 6 Darrah shows what is claimed to be a megaspore membrane in the lower half of his seed and adds that "the tissue outside being in large part, if not entirely, sterile sporogenous tissue." (p. 7). This is a very critical point, and if it is “пос entirely, sterile sporogenous tissue" (presumably this means tissue of the sporangium wall) some explanation of what it might be is certainly in order. In view of my own observations there also is doubt regarding the presence of a mega- spore membrane in the section Darrah has illustrated. The definition of detail in Darrah's fig. 6 is so inadequate that the reader is afforded no basis for reliable interpretation, and the illustration in no way lends objective support to his con- clusions. Perhaps the most serious criticism that I find necessary to make is one pertain- ing to the reconstruction (fig. 45) of what is apparently a seed containing a [Vor. 36 490 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN mature embryo. Оп page 7 Darrah notes four qualifications relative to this re- construction. It is stated: 1. That the gametophyte is not shown since it would be more or less disintegrated. Yet no specimen is described in which this stage of develop- ment is in any way discernible. 2. That "the sporangium, with an embryo of this degree of develop- ment, would be ruptured, probably with the embryonic shoot considerably exserted." But the embryo is shown neatly curled within the unbroken sporangium epidermis. That the "embryo would have a much greater number of leaves, particularly at the growing tip." But the description and illustrations in no way bear this out; and if there were many more leaves present it would seem that this point could have been readily shown in the drawing. 4. That "the orientation of the embryo is variable.” I certainly agree that these parenchymatous proliferations are variable but it seems equally evident that they do not represent an embryo. Following the enumeration of these four qualifications he concludes: "Never- theless this sketch shows the zones of the embryo in their proper relation, and despite the rather unnatural aspect portrays the characteristics faithfully." Accurate or even tentative paleobotanical restorations are certainly very much со be desired. Mr. Darrah deserves commendation for taking the trouble to sum- marize his findings in this form so that those who are not familiar with these fossils may gain a clearer concept of their life form, but to contend that the res- toration “portrays the characteristics faithfully" with the observable facts. seems to be very much at variance We may now return to the normal seeds to consider the supposed correlation of their contained tissues with those of the proliferated seeds. It is probably apparent that the most critical phase of this correlation lies in a determination of the origin of the parenchymatous tissue of the proliferated seeds. If Darrah's contentions are correct one would expect to find some remnants of the sporangial wall (nucellus) tissue, the megaspore membrane, and the gametophyte in those specimens contain- ing immature embryos or even embryos in a rather advanced stage of development. Since no such sequence of disintegrated tissues is in evidence, and since tissues of the supposed gametophyte and sporeling are continuously traceable and connect with the sporangium wall epidermis, some other solution is necessary. At this point it is pertinent to refer to the inner sclerotic layer of the normal seeds which is shown in figs. 12, 16, and 17. И Darrah’s concepts are correct it is hardly possible that this tissue would have been completely disintegrated, yet he makes no mention of it in the germinating seeds. It is not clearly described in his 1941 paper (page 98) but the characteristic sclerenchyma is evident in the top figure of his plate П. Мо mention is made of the three dimensional aspect and pitting of these cells, so different from any other tissue in either the proliferated or normal seeds. My own observations on the elongate nature and pitting of these 1949] ANDREWS—NUCELLANGIUM 491 cells were made largely on Darrah's own slides (preserved in the collections of the Harvard Botanical Museum) and it is difficult to understand why it is not given more prominence at least in the specific description, in view of the fact that, like the vascular strands, it is a tissue quite foreign to Lepidocar pon. In his original paper (1941) on L. glabrum, following his discussion of the sporangium wall, Darrah notes that within this "The seed megaspore is always present” and in a later sentence adds “The gametophyte is extensive, nearly filling the whole cavity.” Мо further description of the gametophyte is given nor do I find it possible to understand the caption to the lower figure of plate П (1941) in which the gametophyte is said to be present. One might overlook this lack on the assumption that the 1941 paper was a preliminary account but I do not feel that his description of the so-called gametophyte in the 1949 paper can be correlated with its organization as presented in the earlier contribution. In view of the supplementary factual evidence presented here, the divergent descriptions, based, in part, on the same material, and the conclusion that this fossil cannot be referred to the genus Lepidocarpon the following emended diagnosis is given: Nucellangium glabrum (Darrah) emend. Andrews. Ovoid seed-like bodies approximately 12 mm. long and 6 X 9.5 mm. in diam- eter with a small circular hilum scar at proximal end. Presumably a sporangium with a wall consisting of the following sequence of tissues: a thick-walled, colum- nar epidermis; broad parenchymatous tissue of isodiametric cells; semi-sclerotic, longitudinally elongate, pitted cells; and a narrow thin-walled inner parenchyma. A single large megaspore contained within. Outer parenchyma traversed from base to apex by two delicate vascular strands. The supposed proliferated form of the fossil somewhat larger, split longitud- inally, consisting of the epidermis and outer parenchyma, the latter with numerous proliferating arms that extend approximately into the former area of the central cavity; peripheral region of this parenchyma as well as arms are vascularized. Locality: Urbandale coal mine, Des Moines, Iowa, and other localities as given by W. C. Darrah, 1941. Age: Middle Pennsylvanian, Des Moines series. Specimens on which the present emended description is based are No. 677 and No. 519 in the paleobotanical collections of the Henry Shaw School of Botany. In the previous accounts no specimens are specifically designated as the type or types. I have, therefore selected the following from Darrah's papers (1941, 1949) for this purpose: As the type for the normal seeds the specimen illustrated in the top figure of plate II (1941), No. 44103 in the collections of the Harvard Botan- ical Museum. As the type for the proliferated seeds the specimen illustrated in fig. 2 (1949). Affinities of the fossil.— I am keenly aware that the following remarks are inadequate as an explanation of the morphology and affinities of this fossil. It is quite evidently an instance in [Vor. 36 492 ANNALS OF THE MISSOURI BOTANICAL GARDEN which it is easier to destroy than to build, and if too much of the former has ap- peared in the preceding pages at the expense of a constructive treatment it is due partly to a lack of sufficient information and partly to an admittedly inadequate interpretation. The suggestions that are given below are based, first, on the belief that Nucellangium is not a lycopod. Ас the expense of repetition the reasons for this belief may be briefly reviewed: The complex sporangium wall with its internal "sclerotic" conducting tissue, the vascularization, and the circular hilum scar. These seem to be of fundamental importance and are not in accord with previously described species of Lepidocar pon. Upon the suggestions of at least two competent morphologists, which, inci- dentally, were offered independently, the possibility has been entertained that Nucellangiuwm represents a hydropterid sporocarp. This possibility was supported by the general shape of the fossils, which is not unlike that of a Marsilea sporocarp, the mode of attachment, the thick-walled epidermal layer, and the vascularization of the peripheral parenchymatous tissue. However, certain features of the wall of the normal seeds, notably the inner sclerotic tissue and the single large megaspore, are not in accord with such a relationship, and the fertile specimens show no evi- dence of having borne sporangia after the manner of Marsilea. Furthermore, no associated remains are known which present hydropterid affinities. The possibility of such an affinity has, therefore, been abandoned. Of the pteridophytic groups, other than the lycopods, which are known from the Upper Carboniferous there seem to be none which present a likely comparison. The remaining alternatives are the pteridosperms or cordaites as seed plant groups, or the possibility that we are dealing with an entirely distinct group of fossils, the affinities of which cannot be conjectured. Since the rest of the plant is not known, speculation in the latter direction at present seems useless. For reasons which will be given below it is, therefore, tentatively proposed that Nucellangium be con- sidered as a primitive cordaite seed, or, if more noncommittal terminology seems preferable, a cordaite reproductive organ. In searching for a lead that might suggest relationships with previously de- scribed fossils comparisons have been made with some of the many seed compres- sions. Of these, certain species assigned to the genus Cardiocarpus offer at least provocative suggestions. In examining the specimens of Cardiocarpus in the Lacoe Collection of the U. S. Geological Survey a few have been noted which correspond very closely to the expected appearance of a compression specimen of Nucellanginm. For example Cardiocarpus minor Newberry (Lacoe coll, U. S. National Museum No. 25421) presents an aspect virtually identical with the profile of the broad face of the Nucellangium fossils. A compression of the latter would almost certainly produce a fossil that would be difficult or impossible to distinguish from this species of Cardiocar pus. Although it is somewhat larger, Cardiocarpus injens Lesquereux (Lacoe coll., U. 5. N. М. No. 25425) may also be mentioned since it displays an epidermal con- 1949] ANDREWS—NUCELLANGIUM 493 figuration that compares closely with the type of epidermis of Nucellangium. Cardiocarpus bicuspidatus (Sternberg) Lesquereux, another compression species, also is closely comparable in size and shape with Nucellangium. Since it may have some bearing on the present problem it seems significant to note that the many species assigned to Cardiocarpus (approximately 125 species are recorded in the U. S. Geological Survey's Compendium Index of Paleobotany) present an amazing variety of form. Were they better known I believe it is con- servative to estimate that a few dozens of natural genera are included in this "compression dumping ground." For want of the necessary anatomical informa- tion that might allow a different disposition, these fossils are regarded as "seeds" and at present it seems most plausible that they have their alliance with plants of cordaitean affinities. In tentatively considering the identity of Nucellangium with a species of Cardiocarpus, such as C. minor, a suggestion of the way in which they may have been borne is presented іп Lesquereux's figure of Cordaianthus spicatus in the “Coal Flora? (Lesquereux, 1884, III, Pl. 109, бе. 1). Неге are seeds of the Cardiocar pus type arranged pinnately in two rows on an elongate axis 5 mm. broad. А com- parable organization is illustrated by Renault and Zeiller in their ‘Flora of Com- mentry’ (Renault and Zeiller, 1888) in figs. 30 and 31 of plate 73. Other authors have figured similar Cordaitalean inflorescences showing seeds of the Cardiocar pus type borne apparently terminally on the appendages of short branches. At this point it is perhaps significant to note that the most abundant plant remains in the coal balls from which Nucellangium has been obtained are the inflorescences, stems, and leaves of the cordaitales. I am aware that the evidence afforded by association is hardly conclusive, yet in view of the abundance of these cordaitean remains, and the МисеПапдінт fossils which compare closely with compressions known to have been borne on Cordaianthus inflorescences, some significance may be attached to this association. In attempting to postulate a satisfactory explanation of the morphology and affinities of Nucellangium it is clear that the unintegumented nature of the fossil is particularly perplexing. I feel quite certain that the normal seeds as described in the earlier portion of this paper represent a nucellus or sporangium wall and in this one respect I seem to be in accord with Mr. Darrah. Is it plausible that we are dealing with an aberrant cordaitean stock whose presence has not been previously suspected? The possibility may exist, of course, as Darrah suggested, that the seeds were shed from their integuments but at present there appears utterly no evidence that would serve even for conjecture. It is clear that we are dealing with a sporangium possessing a wall that is specialized as a protective device to a very high degree. Certainly the epidermal layer would have served most effectively against the attacks of fungi or small animals and equally well to prevent the loss of water from within. Can it be that we are dealing with a plant in which this protective function of the integument was developed by the sporangium wall, that is, the tissue that would normally have evolved into a nucellus of the more usual type? [VoL. 36 494 ANNALS OF THE MISSOURI BOTANICAL GARDEN With reference to the morphology of the proliferated seeds the problem becomes much more involved. Having discarded the gametophyte-sporeling nature of these bodies some other explanation is clearly in order. It is tentatively suggested that the proliferated specimens could represent either aposporus growths of the outer parenchymatous layer of the nucellus, or a gemma-type reproductive tissue. If any weight can be placed on the suggestion that the proliferations represent an aposporus tissue it would seem likely that archegonia should be found in some abundance, but none have been observed in the specimen described here. It is, moreover, strange that sporelings, as described by Darrah, are so abundant and yet no trace of their earlier stages is present, and only one archegonium has been reported. The fact that the parenchymatous tissues of the proliferated specimen, both the peripheral region and the internal "arms," are vascularized, is indicative of sporophytic rather than gametophytic tissue. It would seem, therefore, that the most likely function this structure served was as a purely vegetative reproductive organ—that is, a gemma in the broad sense. It is assumed, following this interpre- tation, that the proliferations developed directly into a new sporophyte plant. Such being the case, it seems likely that the central proliferated "clump" shown in figs. 18 and 19 represents the initial apical meristematic region of the new sporophyte. The question of course will arise as to the disappearance of the inner sclerotic layer which is so conspicuous in the normal seeds and the only explanation that I am able to offer is that the characteristic development of the proliferated seeds originated before the normal maturation of the internal tissue layers. Acknowled g ment.— The abundant Urbandale coal balls, containing a wealth of well preserved, unique plants have been gathered by Mr. Frederick O. Thompson of Des Moines. Unfortunately it has not been possible to obtain more of this material in recent years and although we may reasonably hope that other Iowa localities will eventually contribute toward a more satisfactory solution of this and other paleobotanical problems there appears to be little chance of obtaining more Urbandale coal balls. It is partly on this account that the present writer has decided to submit his study of the available specimens of the fossil described above. It is a pleasure to acknowledge again with gratitude the contributions that Mr. Thompson has made in aid of our investigations of American Carboniferous plants. Sincere thanks are also due Dr. James M. Schopf for his many helpful sug- gestions during various stages of this study. The author, however, assumes all responsibility for such criticism, theories, and conclusions as may be found herein. Bibliography.— Andrews, H. N. and Jules Kernen (1946). ea sawaspa to our knowledge of реден СагБоп- iferous floras. VIII. Another Мей ова from Iowa. . Мо. Bot. Gard. 33:1 Darrah, W. С. (1941). The fossil flora of Iowa coal balls IV. Lepidocerbon. ыы Üniv. Bot. Миз. Leafl 47, -------, (1949). ee Notices II. Paleozoic Lepidodendroid embryos. Medford, Mass. publis hed by the author. cu L. (188 4). Coal flora of the Carboniferous formation. 2nd Geol. Surv. of Penna Renault, and R. Zeiller (1888). Études sur le terrain houiller de Commentry, Atlas. Soc. del г. Min., St. 4. Scott, D. H. (1901). On the structure and affinities of fossil plants from the Palaeozoic rocks. IV. The seed-like initi of Lepidocarpon, a genus n Tue odiaceous cones from the Carbon- iferous formation. Phil. Trans. Roy. Soc. Lond. 194:29 [Vor. 36, 1949] ANNALS OF THE MISSOURI BOTANICAI. GARDEN EXPLANATION Or PLATE PLATE 35 OTE: А considerable ы of the following figures are taken from the series of peels de- scribed on pages 481—485 e peel number refers to the respective position as indicated in text-fig. 1 The slide number is the ы опе assigned in the paleobotanical slide collection of the Непгу Shaw School of Botany. Nucellangium glabrum Figs. 1, 2. Specimens isolated whole from coal balls. The one in fig. 1 displays the hilum scar at the base; this specimen was used in preparing фа series of peels described оп pages 48 5 Epidermis and outer parenchyma showing the two in organic connection in Ее seed. Slide No. 1497. 5 Fig Cells, showing pitting, of the inner sclerotic tissue of a sterile seed in longi- tudinal view. From slide No, 50896, collections of the Botanical Museum of Harvard University, 225. Fig. 5. Hilum scar of the specimen show Photograph of peel No. 9 (slide No. 1643) showing central vascular strand ‚ and departing trace at left. X 110. n in fig. l. X 20, g. 6. (near base of seed) in right center NN. Mo. Вот. Сакр., Vor. 36, 1949 PLATE 35 > [Vor. 36, 1949] 498 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 36 Nucellangium glabrum Fig. 7. Part of transverse section through the median region of a seed showing the vascular trace on that side. From slide No. 50897, collections of the 4. al Museum of Harvard University. Fig. 8. The central vascular strand in the base of the seed, from peel No. 13 (slide No. 1647). X 220 Fig. 9. Section through epidermis of normal seed, from peel No. 34 (slide No. 1656). x 280. Fig. 10. A nearly median longitudinal section through the basal portion of a seed. t, tracheidal tissue in “ar chesporial pad.” From slide No. 50895, collections of the Botani- cal Museum of Harvard University. 6 ll. A highly on view of a branch trace of the normal seed, from рее! . 8 (slide No. 1642). X 50 36 PLATI 1949 GARD., Vor. 36, Bor. ANN. Mo. [VoL. 36, 1949] 500 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 37 Nucellangium glabrum g. ee section through the wall of the normal seed, from peel No. 34 (slide No. 1656): е, epidermis; ор, outer ка is, inner sclerotic layer; ip, inner parenchyma; m, megaspore membrane. lig. 13. One of the central arms, or branch proliferations, from the specimen in fig. 18. Slide No. 1496. Fig. 14. A vascular strand of the proliferated specimen (fig. 18). Slide No. 1498. X 440. A section in the transverse plane through а normal seed taken at one side of the central basal strand. The dark tissue represents the lowermost extension of the inner sclerotic tissue. This is associated at this level with the departing traces and is shown in text-fig. 3 (3) as the small central cross-hatched areas. From peel No. 11 (slide No. 1645). X 80. ) - 7 ; А 3 ATE 3 ANN. Мо. Вот. Garp., Vor. 36, 1949 PLAT 7 Не: LN [Vor. 36, 1949] 502 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION Or РЕАТЕ PLATE 38 Nucellangium glabrum Fig. 16. Photograph iere half of the transverse section of the normal seed, рее! No. 38 (slide No. 1658): epidermis; ор, outer parenchyma; А, inner sclerotic layer; ip, inner parenchyma; m, ее membrane; /, vascular trace. Х 20. Fig. 17. С и transverse section of the сз seed, peel No. 20 (slide No. 1653): f, position of traces; is, inner sclerotic layer. Note interruptions of the inner sclerotic layer on the two 415 compare with text-fig. 3 (4). ANN. Mo. Вот. Сакр., Vor. 36, 1949 PLATE 38 [Vor. 36, 1949] 504 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 39 Nuce'langium glabrum Figs. 18, 19. Photographs of sections of the proliferated specimen described on pages 486-488, /', vascular strand іп proliferated vascular strand in peripheral parenchymatous zone; e, epidermis. ag. 18. From peel No. 519,77 (slide No. 1674): arm (see fig. 14); /”, X 12. Fig. 19. From peel No. 519,729 (slide No. 1675): v, epidermis; р, peripheral par- enchymatous zone which is in organic connection with the epidermis (е), and from which the proliferated arms arise; i, by following through successive peels these scattered "islands" of tissue may be observed to be proliferated from the peripheral parenchyma zone. X 16 ANN. Мо. Вот. Сакр., Vor. 36, 1949 PLATE 39 А REVISION OF THE GENUS HELIOCARPUS 1." КО КО LAY INTRODUCTION Heliocarbus has received considerable attention from plant systematists prob- ably because of the perplexing variation found in the genus and because of the few constant characters of taxonomic value. Furthermore, in the herbarium, the speci- mens are either in fruit or in flower, never both; and when in flower are either hermaphrodite or pistilate. Thus, assigning them to any particular species be- comes extremely difficult. Despite a recent taxonomic study of the genus,” there is still considerable confusion regarding many species both in the literature and in the herbarium. More than fifty species and varieties have been named thus far, and with the prevalent vagueness in the concept of speciation, there appears super- ficially to be but two alternatives: either to split the genus into innumerable in- distinct and undefinable "species" or to lump them indiscriminately into few cate- gories of scarcely greater reality. In my study of the genus an attempt has been made to escape this dilemma by clarifying the concept of speciation. However, as this study has been confined entirely to herbarium specimens which represent only very small portions of the woody plants, no definite idea or suggestion as to the individual variations of single plants has been obtained. I have been fortunate enough in being able to study specimens from nearly all the major herbaria both in the United States and in Europe. The standard method of the herbarium taxonomist has been used for the interpretation of the species, and an attempt has been mede to identify the fruiting specimens with the flowering ones. As far as possible, no intergrading forms have been considered as worthy of specific rank, and I have tried to group the "species" into fewer categories of perhaps greater biological reality, in the hope that they will be satisfactory both from a taxonomic and from a practical standpoint. The key has been so prepared that it should be usable for both the fruiting and the flowering specimens. GENERIC RELATIONSHIPS Heliocarpus 1.3 commonly is referred to the tribe Grewieae* of the family Tiliaceae and usually is recognized by its characteristic fruits. The genus is dis- tinct from the other genera of Grewieae except Triumfetta. There is no difficulty in distinguishing the two genera when both are in fruit, as the fruits are very dis- pnis чан кае out ас the м; Botanical Garden and submitted as а thesis іп partial x. of the requirements for the degree of Master of Arts іп the Henry Shaw School о Botany of Жо, University. " Watson, Е. E. The genus Heliocarpus. Bull. Torrey Bot. Club 50:109. 1923. 35р. Pl. ed. 1. 448. 1753. *K. Sch. in 00 & Prantl, Nat. Pflanzenfam. 16:29. 1895. (507) [Vor. 36 508 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN tinct. А very good description of the fruit of Heliocarpus was given by Linnaeus’, who asked, in naming the genus, "Who could ever behold an almost rounded fruit, bordered with a halo of rays, without thinking of the sun as conceived by the painters?” In Triumfetta the fruit is a burr, with many bristles all over the surface. It is slightly more difficult to differentiate between the two genera in the case of flowering specimens. Both have alternate, palmately reticulated leaves with long, slender petioles and stellate pubescence. The aspect of the specimens also is very similar and the superficial resemblance rather striking. The chief differ- entiating characters may be summarized as follows: (1) The inflorescence—In Heliocarpus usually terminal, and when axillary large and spreading; in Triumfetta generally axillary, rarely large and spreading. (2) The cymes—In Heliocarpus disposed in nodose clusters of about 12-20 flowers each; in Triumfetta generally not in nodose clusters, and if in nodose clusters usually 6- to 12-flowered. (3) The flowering peduncles—In Heliocarpus nearly always 3-flowered (rarely 2-flowered), and usually ebracteolate; in Triumfetta 1- or 2-flowered (rarely 3- flowered) and conspicuously bracteolate. (4) The mature floral buds—In Heliocarpus as long as 6—7 mm., the sepals appendaged or unappendaged at the tips, the petals valvate; in Triumfetta as long as 2-3 cm., the sepals always with apical appendages, the petals with twisted aestivation. (5) The number of stamens—In Heliocarpus usually 12-40; іп Triumfetta usually more numerous. (6) Tbe gonopbore—Simple in Heliocarpus; іп Triumfetta with a ciliate saucer-shaped margin (urceolus) surrounding the stamens. (7) Tbe ovary—In Heliocarpus either borne upon a gynophore or sessile upon the gonophore, 2-celled, laterally compressed and ciliate about the margins; in Triumfetta always sessile upon the gonophore, 3- to 5-celled, not laterally com- pressed, and generally pubescent. (8) The style—In Heliocarpus not more than three to four times the length of the ovary, usually much shorter, always bifid at the tip with the stigma lobes spreading; іп Triumfetta usually longer than three to four times the length of the ovary, simple throughout, the stigma flattened or capitate. Heliocarpus can be divided into two major groups of species based on the presence or absence of a gynophore in flower or fruit; further, the presence of ap- pendages at the tips of the sepals is a character which is correlated in the majority of species (except іп H. mexicanus and Н. nodiflorus) with the absence of the gynophore. 5 Сгігіса Botanica (transl. by Hort and Green). Roy. Society, London. p. 79. 1938. 1949] КО КО LAY—REVISION ОҒ HELIOCARPUS 509 GEOGRAPHICAL DISTRIBUTION The geographical range of Heliocarpus embraces nearly the whole of the tropical Americas. With the exception of the predominantly South American H. popayanensis, the species are indigenous entirely to Mexico and Central America. The presence of some plants of H. popayanensis in Hawaii apparently is due to recent introduction for reforestation in the foothills. Possibly in the same category is the presence of some plants of H. Donnell-Smithii in Martinique. The species can be divided into five groups with respect to their geographical distributions: (1) Northwestern Мехісо--Н. attenuatus and H. Palmeri. (2) Southwestern Mexico—H. terebinthinaceus, H. pallidus, and H. occi- dentalis. (3) South-central Mexico and Central America—H. mexicanus, H. appendicu- latus, H. Donnell-Smithii, and H. nodiflorus. (4) Southeastern Mexico—H. americanus. (5) Southern Central America and South America—H. popayanensis. species of the sessile-fruited group are confined for the most part to the northern range of the distribution; while those of the stipitate-fruited group, where the fruits are borne upon bristly gynophores, are mostly in the middle and southern = 8 areas of distribution. The northernmost limit of the genus is in subtropical Sonora and Chihuahua, Mexico, where the plants grow in canyon forests or oak flats; the southernmost limit is in central Argentina, where lower elevations along river banks are inhabited. Except for shrubby plants (H. occidentalis and H. pallidus), which sometimes grow on barren hill slopes, the majority of the species, consisting of small trees, grow only in moist places at higher elevations, usually at about 1000 m. or more, in rain- or cloud-forests, along roadways or river banks in sheltered places, or on edges of forests (usually on cut-over lands in second growth, where they some- times form a pure stand). ECONOMIC VALUE In so far as is known from collectors’ notes and from the published accounts of Standley® and Martinez’, the economic importance of Heliocarpus is primarily іп its bark, which produces a very strong and durable fibre. The bark of the young branches yields a fibre from which a strong but coarse rope is made. It is used also for weaving mats and baskets. Тһе principal component of the Mexican fabric belem is the cordage extracted from the barks. Resistant paper formerly was made from the wood in Mexico; in Brazil it is still so used to some extent. The wood of the trees is soft and light and is used for floats and bottle-stoppers. Because of its lightness it has been used for rafts. The bark is used for mecates іп Guate- $Contr. О. S. Nat. Herb. 23:739. 1923. "Plantas utiles de Mexico, p. 253. 1936. [VoL. 36 510 ANNALS OF THE MISSOURI BOTANICAL GARDEN mala, and a decoction is used in sickness of cattle, and sometimes is applied on sores. In Hawaii the trees have been planted on the foothills for reforestation. VERNACULAR NAMES H. americanus: janote (Vera Cruz). H. appendiculatus: burio, Бито blanco, and burio colorado (Costa Васа); cajeton (Guatemala); balsa and pastano mula (Nicaragua); calagua (El Salvador); jonote, jonote blanco, jonote colorado (Vera Cruz) ; majao (Honduras). H. attenuatus: samo baboso (Sinaloa). H. Donnell-Smitbii: bolol (Quintana Roo); jolocin (Tabasco); jonote (Vera Cruz); majao (Honduras) ; торо, “broad-leaved mobo”, and "mountain торо” (British Honduras). H. mexicanus: antigua, cajeto, and majagua (Guatemala); mobo (British Hon- duras). Н. nodiflorus: "broad-leaf торе” (British Honduras) ; cajeto (Guatemala) ; majao (Honduras). H. occidentalis: pulmonilla, guasima, and panigua (Chiapas and Jalisco). H. pallidus: guasima and tilia (Guerrero, Jalisco, and Mexico). H. Palmeri: coche (Sinaloa) ; тата kowusamo, and palo chinu (Sonora). Н. popayanensis: balsa (Colombia); majagua (Venezuela); palo de balsa (Peru); sanpan (Ecuador). H. terebinthinaceus: cuabualagua or quaubalagua (Morelos and Puebla); guasima, jolotzin, majorbua, and jonote (Morelos, Oaxaca, and Guerrero). ACKNOWLEDGMENTS During the course of this study, materials from the following herbaria have been examined, for which I wish to acknowledge my indebtedness: Jardin Principal de l'Etat, Bruxelles, Belgium. Conservatoire Botanique, Genéve, Switzerland. Chicago Natural History Museum (Field), Chicago, Illinois. Gray Herbarium of Harvard University, Cambridge, Massachusetts. Royal Botanic Gardens, Kew, England. Missouri Botanical Garden, St. Louis, Missouri. New York Botanical Garden, New York, N. Y. Royal Botanic Gardens of Trinidad and Tobago, Port-of-Spain, Trinidad, B.W.I. University of California, Berkeley, California. University of Michigan, Ann Arbor, Michigan. United States National Herbarium, Washington, D. C. I am greatly indebted to the Missouri Botanical Garden and to its director, Dr. G. T. Moore, for the use of its library and herbarium facilities during the course of this study. Particular thanks are due to Dr. R. E. Woodson, Jr., for his advice, guidance, and constructive criticism. 1949] KO KO LAY— REVISION ОЕ HELIOCARPUS 311 TAXONOMY HELIOCARPUS L. Sp. Pl. ed. 1. 448. 1753; Gen. РІ. ед.5. n. 606. 1754. Adenodiscus Turcz. in Bull. Soc. Nat. Moscou 19?:504. 1846. Woody trees or shrubs; older branches cream to brown, nearly glabrate, the younger branches usually stellate-pubescent, dark brown, fibrous, with many mucilage canals. Leaves alternate, petiolate, stipulate, the stipules usually large, early deciduous, very rarely persistent, the blade 3-lobed to undivided, venation palmate and either 5- to 7-costate or 3-costate at the base, irregularly serrated, the basal serrations usually glandular, slightly to densely stellate-pubescent, acute to тал. JF Aetoli Illustrations of taxonomic criteria of the genus Heliocarpus: 1, leaf of r gem show- ing the 3-costate venation at the base; 2, leaf of H. рот 8а showin tate venation at the base and the 2 leafy auricles at th к sinus; 3, емер fruit of H. кыл showing the ІН tufts E Mays hairs; 4, M md fruit of H. Donnell Sibi borne on a bristly gynophore; ral bud o puris 07 the relativ: Mes large, т ќи ps trigonal appendages on the tips of os sepals; 6, 1” al pcd Н. appendiculatus (note the absence of apical Р ages); fig. 7, 3- radiate flowering peduncle EA ) of H. Palmeri viti a pair of bracteoles [Vor. 36 512 ANNALS OF THE MISSOURI BOTANICAL GARDEN acuminate, the base rounded or cordate. Inflorescence gynodioecious or hermaph- rodite, usually terminal (axillary in some species), large and spreading, consisting of numerous aggregate dichasia, the main axis branching sympodially and at each node bearing a cluster (cyme) of some 12—20 flowers, the terminal branches with 5—7 cymes, usually more or less secund, each cyme consisting of reduced sympo- dially branched axes terminating in 3- (rarely 2-) flowered cymules. Flowers either hermaphrodite or pistillate, regular, hypogynous, 4- to 5-merous; the sepals valvate, usually stellate-pubescent without, glabrate within, sometimes with small appendages at the tips; the petals 4-5 in hermaphrodite flowers, absent in the pistillate, free, valvate, 1- to 3-nerved, shorter than the sepals, usually ciliate at the base, sometimes slightly so at the apex; the stamens 12—40 in hermaphrodite flowers, staminodial or absent in the pistillates, borne cyclically on the enlarged gonophore, with 2-lobed, 4-celled, introrse, longitudinally dehiscent anthers; the ovary wholly superior, sessile upon the gonophore or with a manifest gynophore, ellipsoid to orbicular, laterally compressed, ciliate and with shorter pubescence on the faces, 2-celled, falsely 4-celled at the base, each cell with 2 anatropous ovules, the style filiform, bifid at the tip with the stigma lobes spreading and simple or slightly 3- lobed. Fruit dry, indehiscent, 2-celled, 2-seeded, laterally compressed, ellipsoid to orbicular, slightly rugose, sessile or stipitate upon the accrescent gynophore, ciliate with 2 rows of plumose bristles decurrent upon the gynophore when present, the faces pubescent to glabrate; the seed compressed-ovoid or -pyrifrom, with a median funicular groove, endosperm oily. Standard species: Heliocarpus americanus L. KEY TO THE SPECIES A. Fruit sessile; ovary sessile on the gonophor B. Leaves 3- p veins 3-costate at de buds buds with or without appendages at the tips ы the Ç m with dense coarse pubescence on both surfaces, cordate, 3-lobed, рена E . long a cm. wide; buds sometimes ж smal appendages at vy tip s жори rss anie tomentose; southwestern Mexi H. As Ao PE CC. Leaves with p smooth to um benea i gm го the D d to slightly yir ана 5 -lobed to лавот so, about 10 ст. long and 8 ст. wide; buds with conspi dpi erect appendages at the tips of the sepals; p ovoid, glabrate; southwestern Mex z, Н, ‘pallidus BB. ~~ not lobed, veins 5- to 7-costate at the base; buds with conspicuous appendages t the tips of the sepals ë Leaves cordate eaves өт glabrate оп both surfaces, about 14 cm. long and 10 cm. I seuminate; appendages on the sepals trigonal, large and reflexed; stamens abou 40 style about 4 times longer RR ~~ ovary; fruit spherical, е. а e гы groove in the body, slightly tomentose; northern Mexico . Palmeri DD. Leaves glabrescent above, tomentose ийем small, about 8 ст. ед E 5 cm. wide, caudate-acuminate; appendages on the sepals тирен erect; stamens 16-20; style about twice the length of the ovary; fruit дона, tomentose; northern exico 4. H. attenuatus СС: den rounded or cuneate at the base, Кя ог nearly so on both surfaces; endages on the sepals short, erect; sta about 40; style about twice the length gh d цар fruit ovoid with ша: реди tufts of stellate өзі south H. calmi 1949] KO KO LAY—REVISION OF HELIOCARPUS 515 АА. Fruit borne on a bristly gynophore; ovary borne оп a gynophore upon the gonophore. B. vano perl leafy; leaves not lobed, the base obtuse, glabrate above, slightly = den ath; buds with conspicuous appendages at the tips of the sepals; stamens about 30; fruit ellipsoid. glabrous; southern central Mexico аа Central . mexicanus ВВ. Inflorescence generally еке, пос leafy. C. Lea wo conspicuous auricles at the basal sinus, usually not lobed, the base roun одеа or subcordate, bud above, tomentose beneath; buds without түрен аре ас об ps of os "pe stamens теді 30; fruit пБ е amy? souther tral ME xico and Central Am y. и СС: quum without ош: ШЕ: ас the База! sinus. D m ves not lobed, rounded at the base, not longer than 14 cm. or wider than E. Lied very densely tomentose beneath, glabrate above; stamens i etm 20; style very briefly bifid; fruit ee tomentose; southeastern Мехісо...... . americanus EE. Leaves glabrate on both surfaces; stamens about 12-16; style bifid em t ha | S Е fruit ellipsoid, glabrate; southern central Mexico те Cent 9 Plon Smithii DD. Leve рога 3-lobed, cordate ас the base, about 17 cm. long i. over 14 cm. E. Du glabrate above, the lower рш puberulent with eri stellate and h u ag of the sepals; stamens about 12; style briefly bifid, each ict with 3 acute lobes; fruit e slightly tomentose to nearly glabrate; “р Central America and Sou . popayanensis EF. Leaves 5 on Y ы veins not pubescent on the lower Ее е; buds sometimes wi pass short appendages at the tips of the sepals; stamens about 24; icis bifid about half its length, phis -lobes simple, pe [pu ih pn slightly BE southern Mexico and Central Americ H. nodiflorus 1. HELIOCARPUS TEREBINTHINACEUS (DC.) Hochr. in Ann. Conserv. et Jard. Bot. Genéve 18:125. 1914. Grewia terebinthinaceus DC. Cat. Hort. Monsp. 114. 1813. (Т.: Berlandier 1064). sonii Rose, in Contr. U. 5. Nat. Herb. 6:128. 1897. (T.: Nelson 1485) Н. reticulatus Rose, loc. cit. 128. 1897. (Т.: Pringle 1701 H. microcarpus Rose, loc. cit. 8:316. 1905. (T.: Pringle 8719). Small trees or shrubs 4—5 m. high; older branches cream to brownish, glabrate, slightly longitudinally ridged, irregularly punctate with white lenticels; younger branches and inflorescence axes rough, scurfy, ferruginous-tomentose with stellate hairs. Leaves 3-lobed, acuminate, about 13-15 cm. long and 10-12 cm. wide, 3- costate, the mid-costa arising independently of the lateral costae, and each lateral costa consisting of 2 or 3 veins united at the basal sinus, base cordate, irregularly and bluntly serrated, both surfaces light green or pale brownish with thick, matted, stellate pubescence, the veins more pubescent with both stellate and long simple hairs; petioles slender, about 8 cm. long, covered with ferruginous tomentum. Inflorescences gynodioecious; the hermaphrodite usually axillary, rarely terminal, large and leafy, usually about 20 cm. long and 15 cm. wide, sometimes about 25 cm. long and 20 cm. wide, the cymes of about 15—18 flowers, rather compact in nodose clusters, the flowering peduncles 3-radiate, about 2 mm. long, the pedicels about 2 mm. long, the buds obovoid, constricted towards the base, sometimes with small erect appendages on the tips of the sepals, the sepals 4, spatulate, about 4—5 [Vor. 36 514 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 1. H. terebinthinaceus mm. long, densely stellate-tomentose without, glabrate within, the petals 4, linear, 1-пегуед, about 3-4 mm. long, the ovary ellipsoid, about 1.5 mm. long, sessile, with a style about 3 mm. long, bifid about one quarter of its length and with stigma lobes acute, spreading; the pistillate much smaller than the hermaphrodite, about 10—15 cm. long and 8—10 cm. wide, the cymes of about 18 flowers, very much crowded in compact nodose clusters, the flowering peduncles 3-radiate, very short, less than 1 mm. long, the pedicels about 1-2 mm. long, the buds obovoid, not appendaged at the tips of the sepals, the sepals 4, linear, about 2 mm. long, densely stellate-tomentose without, glabrate within, the petals none, the staminodes about 10—12, the ovary ellipsoid, minute, sessile, with a style about twice the length of the ovary, briefly bifid at the tip, and with the stigma lobes acute. Fruit suborbicular, 4—5 mm. in diameter, rather densely tomentose, sessile, the fringe of two rows of plumose bristles about 5—7 mm. long; the seed ovoid, about 2 mm. long, with a very slight groove in the middle. Distribution: On the highlands of central Mexico, especially on open hill slopes, altitude above 1000 m. Flowers throughout September and October, and fruits from late October to December. МЕХ CHIAPAS: Tuxtla Guttierez, и 1461. GUERRERO: above Chil- pancingo, p ea 70403 ич Madre, alt. 1250 т., Langlassé 572. JALISCO: road to бап omingo mine, alt. 0 ft., Etzatlan, Barnes @ Land 285; barranca, Guadalajara, alt. 4500 ft., Pringle ens ав. Pringle 1701. MEXICO: Tejupilco, Temascaltepec, 1949) KO КО LAY—REVISION OF HELIOCARPUS 515 Map 1. Distribution of H. terebinthinaceus Hinton 0736; Acatitlan, Temascaltepec, alt. 1130 m., Hinton 5110. МОК vaca, alt. 0 ft., Pringle 8227; lava field near Yantepec, alt. 4000 ft., po» 9692; vicinity Ж TE 35220 8 lo s. .; Cuernavaca, Atlacomulco, alt. 0 m. Woronow 2333; Cuernavaca, Bird 1064; Cuernavaca, Bourgeau т: about Cuernavaca, alt. 5000 ft., Pringle 8710. NAYARIT: El Trapiche, s. е. of Yxtian, alt. 1200 m., Mexia 800. олхаса: Valley of Oaxaca, alt. 5100-5800 ft., Nelson 1243; Valley of Oaxaca, alt. 5500—7500 ft., Nel 1485; on slopes ЕНА of Cincatlan, alt. 2500- 4000 ft., Nelson 1818; Rio Blanco, San Juan del Estado, alt. 0f L: ^ Smitb 797; eie Ай, alt. 6200 ft., L. С. Smitb 030; Hacienda ш. alt. 1600 m., Conzatti s. ль; Tomellin Cañon, near city of OE Rose & Hough 4505. DATA INCOMPLETE: Ahualulco ?, Barcena, Urbina 630. This species 15 very distinct and сап Бе recognized easily Бу its 3-lobed leaves with the characteristic venation, and also by its sessile fruits. Hochreutiner pointed out that Berlandier 1064, collected from Cuernavaca, which Rose cited for H. Nelsoni, is identical with the specimen which de Candolle, on the assumption of a 4-locular ovary, described as Grewia terebinthinaceus. Rose differentiated his H. Nelsoni (type: Nelson 1243 & 1485), H. reticulatus (type: Pringle 1791) and H. microcarpus (type: Pringle 8719) primarily on the degree of pubescence on the fruits. This character is extremely variable. In matching the three types, it is noticeable that H. microcarpus has slightly smaller fruits and that the flowers have minute erect appendages. The size of the fruits varies slightly from plant to plant and sometimes on the same plant. Furthermore, the apical appendages on the sepals, when present, are extremely minute and not clearly seen. As they stood, it was difficult to tell the three species apart and there were many misdeterminations. 2. НЕШОСАКРЏ5 PALLIDUS Rose, in Contr. U. S. Nat. Herb. 5:128. 1897. (T.: Е. Palmer 157). H. velutinus Rose, loc. cit. 8:317. 1905. (T.: Pringle 8604). [Vor. 36 516 ANNALS OF THE MISSOURI BOTANICAL GARDEN Small trees or shrubs about 2-4 m. high; older branches yellowish brown, glabrate, slight- ly rugose, punctate with few white lenticels; younger branches and inflorescence axes scurfy, slightly ferruginous-tomentose. Leaves small, younger ones generally not lobed or obscurely so, older ones 3-lobed, about 8-10 cm. long and 7—8 cm. broad, 3-costate, with the mid- costa arising independently of the two lateral costae, and each lateral costa consisting of 2 or 3 veins united at the sinus, base usually cordate or subcordate, in younger leaves slightly rounded, acuminate, irregularly serrated, upper surface dark green, glabrate, lower surface fi \ | whitish, densely stellate-tomentose with ap- for ! pressed hairs; petioles long and slender, 6-7 cm. long. Inflorescence hermaphrodite, usually axil- lary, large and spreading, leafy, about 14-20 cm. long and 12-20 cm. wide; the cymes of about 12-15 flowers borne loosely in nodose clusters, the flowering peduncles 3-radiate, about 2-3 cm. long, the pedicels about 4—5 mm. long, the buds obovoid, constricted at the base, with small linear, acute, erect appendages at the tips of the sepals, the sepals 4, linear, 4—6 mm. long, light green and slightly stellate-pubescent without, glabrate and yellowish within, the petals 4, linear, about 3-4 mm. long, the ovary ovoid, small, 0.5—1.0 mm. long, sessile, with a style about twice the length of the ovary, briefly bifid at the tip, and with the stigma lobes acute. Fruit ovoid, usually in compact clusters, very tomentose when immature, becoming glabrate at maturity, about 5 mm. long and 3 mm. wide, sessile, the fringe of two rows of plumose bristles about 4—5 mm. long; the seed obliquely ovoid, about 3 mm. long and 2 mm. wide, with a very deep groove in the middle Distribution: On the highlands of central and southwestern Mexico, especially on open hill slopes, altitude above 500 m. Flowers from October to November, and fruits in December and January. Usually leafless іп January. Fig. 2. H. pallidus O: GUERRERO: Paso de las 275 Nelson 6070; Pungarabato, Coyuca, Hinton "o ae. and vicinity, E. Palmer 157. Jatisco: Puerto Vallarta, El Real, Mexia 1100. MEXICO: Chorrera, а. alt. 1350 т., Hinton 5416; Ocotepec, Temascal- tepec, Hinton 7036; Tenayac, Temascaltepec, Hinton 5109; —' Temascaltepec, Hin- ton 7236; Ixtapan, Temascaltepec, Hinton 7155. MORELOS: — alt. 4500 ft., Pringle 8604; NO Berlandier 071. DATA INCOMPLETE: i Hore The types of the two species H. pallidus (E. Palmer 157) and H. velutinus (Pringle 8604) have leaves which apparently look distinct, those of H. pallidus being small, obscurely 3-lobed, with rounded base, and of H. velutinus relatively 1949] КО КО LAY—REVISION ОЕ HELIOCARPUS 917 Мар 2. Distribution of H. pallidus small but definitely 3-lobed with subcordate base. It is apparent that the type of H. pallidus represents the immature and that of H. velutinus the mature state of the leaves. On the same branch (cf. Hinton 5410 and Mexia 1100) the leaves at the base of the specimens may correspond to those of H. velutinus and those near the tip to H. pallidus. The fruits on the type of H. velutinus are slightly larger and more glabrate than those on the type of Н. pallidus. There are many intermediates between these two extremes, in specimens that answer to the above descriptions. Presumably the fruit, which is slightly tomentose when young, becomes nearly glabrate at maturity. 3. НЕШОСАКРО5 PALMERI S. Wats. in Proc. Amer. Acad. 21:420. 1886. (Т.: Е. Palmer ТОТ). H. polyandrus S. Wats. loc. cit. 1886. (T.: E. Palmer 100). H. glaber Brandegee, in Zoe 5:207. 1804. (T.: Brandegee s. n. in Herb. Calif.). Diffusely spreading shrubs about 2-3 m. high; older branches dark brown, glabrate with few short tufts of stellate hairs mostly around the white lenticels; younger branches and inflorescence axes covered lightly with short stellate tomen- 518 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 3. H. Palmeri - но Lr \ \ I *, % 9 Са т ¿ \ ам ` `® „ e ° ES - š 4, М Q- a ` / " I ^ М РЕ I # w | РА À A <р \ am! di \ f us P LAE | [ № | `. à < š % \ ' PR Ж ' sh i Ы, Ж Қ. iin oeie ыы „Б-ке нид, Мы N, ж» «а ми iid [Vor. Map 3. Distribution of H. Palmeri 36 1949] KO KO LAY—REVISION ОЕ HELIOCARPUS 519 tum. Leaves broadly ovate, 12-14 cm. long and 8-10 cm. wide, not lobed, 5- costate, narrowly acuminate, base cordate or subcordate, thin, unequally and bluntly serrated, basal serrations glandular, upper surface dark green, nearly glabrous to glabrate, rough, with few short suppressed, scattered tufts of stellate hairs, some- times slightly scabrous on the veins, lower surface lighter green, slightly more pubescent than the upper, nearly glabrate, generally coarse, with short suppressed stellate hairs, veins scabrous, prominent; petioles slender, 5-7 cm. long, covered very lightly with short suppressed yellowish stellate pubescence. Inflorescence hermaphrodite, terminal, rarely axillary, slightly leafy, about 14 cm. long and 12 cm. wide, rarely larger, the cymes of about 12 flowers rather loose, the flowering peduncles 3-radiate, about 2 mm. long, the pedicels 3-4 mm. long, the buds obo- void, constricted towards the base, with conspicuous appendages at the tips of the sepals, the sepals 5, spatulate, 5—6 mm. long, 3-nerved, with large, about 2 mm. long, reflexed, trigonal appendages at the tips, light green with short suppressed tufts of stellate hairs without, glabrate, yellow to yellowish brown within, the petals 5, linear, slightly shorter than the sepals, about 4 mm. long, 1- to 3-nerved, yellowish brown, the stamens about 40, with filaments about 4—5 mm. long, the ovary orbicular, 1-2 mm. long, sessile, with a slender style about 4 times the length of the ovary, briefly bifid at the tip, and with small acute stigma lobes. Fruit orbicular, sessile, densely stellate-pubescent, 5 mm. in diameter, with a distinct groove in the middle of the fruit body, the fringe dense, of 2-3 rows of plumose bristles shorter than the diameter of the fruit body, about 3 mm. long; the seed plump, ovoid, very slightly grooved, about 2 mm. long. Distribution: On shady canyon slopes or oak flats in northwestern Mexico, altitude about 500 m. Flowers in September-October and fruits in late October— December. Mexico: CHIHUAHUA: Hacienda San Miguel, near Batopilas, Е. Palmer 97, 100, 101; Алтын Río Mayo, Gentry 2440, 2000; La Mesa, Gentry ббоў; Almaden, Le Sueur SINALOA: Cerro Colorado, alt. 2000 ft., Gentry 5060; vicinity of Culican, Кый; s. n. (type of H. glaber). SONORA: Соб Sapopa, Río Mayo, с. 1644, 1709. А very distinct species with nearly glabrous leaves and sessile fruits character- istically grooved in the middle and with flowers bearing relatively large trigonal apical appendages on the sepals. The leaves in the type of H. Palmeri (E. Palmer IQI) are slightly more pubes- cent than those of H. polyandrus (E. Palmer 100), but in both they are nearly glabrous and the variation is only one of slight degree. S. Watson mentioned that in H. Palmeri the number of stamens is about 20, but I have not been able to find any specimen that answers to the description of H. Palmeri with 20 stamens, either in the specimens cited by S. Watson or in others. | VoL. 36 520 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN 4. HELIOCARPUS ATTENUATUS S. Wats. in Proc. Amer. Acad. 21:420. 1886. (Т.: E. Palmer 99). = s E. E. Wats. in Bull. Torr. Bot. Club 50:120. 1923. (T.: Rose, Standley 8 ine 12828), Small shrubs about 2-3 т. high; older branches dark brown, rather slender, slightly pubescent, smooth, irregularly punctate with few white lenticels; younger branches and inflorescence axes softly tomentose with relatively long, light yellowish brown, stellate hairs, smooth. Leaves ovate to very obscurely 3-lobed, 5-costate at the base and gradually tapering to the caudate-acuminate tip, base cor- date or subcordate, irregularly and bluntly serrated, upper surface slight- ly pubescent, coarse, with many sup- pressed stellate hairs, scabrous on the veins, lower surface densely stellate- pubescent especially on the veins. In- florescence hermaphrodite, usually axillary, small, leafy, about 8 cm. long and 10 ст. wide, rarely larger, the cymes of about 16—18 flowers borne loosely in nodose clusters, the lowering peduncles mostly 3-radiate, rarely 2-radiate, about 5—6 cm. long, usually subtended by two short brac- teoles, the pedicels 2—3 mm. long, the buds obovoid, very slightly con- stricted towards the base, with short appendages on the tips of the sepals, the sepals 3 or 4, linear, 4—5 mm. long, 1- to Fig. 4. H. attenuatus 3-nerved, the short, erect appendages on the tips 1 mm. long, light green with fine tomentum without, glabrate, yellowish brown within, the petals 3 or 4, linear, shorter than the sepals, 2-3 mm. long, the stamens about 16—20, with the filaments as long as the petals, the ovary ovoid, 1.0-1.5 mm. long, sessile, with a style about twice the length of the ovary, bifid about one quarter of its length and with the stigma lobes acute, spreading. Fruit ovoid, sessile, densely tomentose, 3 mm. long and 2 mm. wide, the fringe of two rows of slender plumose bristles, 6—7 mm. long: the seed ovoid, 1-2 mm. long. Distribution: Confined entirely to northwestern Mexico, apparently rare, usual- ly growing at altitudes of about 300-500 m. Flowers from August to September and fruits from October to December. MEXICO: CHIHUAHUA: southwestern Chihuahua, E. Palmer 00; Rio Benito, Le Sueur 661, 1158. SINALOA: Choix, Tasfera, alt. 420 m., Gonzalez 860. SONORA: Alamos, Е. Palmer 647, 733; Sierra de Ales: Rose, Standley 8 Russell 12828, 1949] КО КО LAY—REVISION OF HELIOCARPUS 521 Map 4. Distribution of H. attenuatus This is a very distinct species, with the caudate-acuminate leaves the smallest known for the genus. It does not resemble any other species in its vegetative characters. The leaves of the type of H. viridis (Rose, Standley & Russell 12828) are small but not caudate-acuminate. The type is from a relatively young plant and rather poorly preserved. I have not been able to find any other specimen which matches the type, and I am doubtful about the type itself. Except for its small wrinkled leaves, there is nothing which would prevent its identification with Н. attenuatus (type: E. Palmer 00). 5. HELIOCARPUS OCCIDENTALIS Rose, in Contr. U. $. Nat. Herb. 5:127. 1897. (T.: Palmer 440). H. laevis Rose, loc. cit. 8:317. 1905. (T.: Rose 2860). Small trees or shrubs about 5—6 т. high; older branches generally glabrate, cream to brownish, slightly rugose, irregularly punctate with white lenticels; younger branches and inflorescence axes covered with rather dense tomentum of short yellowish stellate hairs. Leaves broadly ovate, sometimes rather obscurely 3- lobed, 10—13 cm. long and 8-11 cm. wide, 5- to 7-costate at the base, narrowly acuminate, base rounded or cuneate, rarely subcordate, somewhat regularly and doubly serrated, upper surface generally dark green, glabrate, coarse, with sup- pressed stellate hairs, veins and veinlets more pubescent with short tufts of yellow- ish stellate hairs, lower surface slightly more pubescent than the upper, becoming f VoL. 36 222 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN nearly glabrate when mature, yellowish green, the veins and veinlets covered with dense tufts of yellow stellate hairs; petiole about 6 cm. long, slender, covered lightly with yellow stellate hairs. Inflorescences hermaphrodite, usually axillary, rarely terminal, rather leafy, about 14 cm. long and 10 cm. wide, some- times much smaller, the cymes of about 12 flowers, rather loose in nodose clusters, the flowering peduncles 3-radiate, about 2 mm. long, subtended by two small bracteoles, the pedicels 2-3 mm. long, the buds obovoid, slightly constricted towards the base, with conspicuous appendages at the tips of the Fig. 5. Н. occidentalis sepals, the sepals 5, rarely 4, linear-spatulate, 4-5 mm. long, 3- to 5-nerved, with small, about 1 mm. long, erect, linear appendages at the tips, light green with short stellate hairs without, glabrate, cream or light brown within, the petals 5, rarely 4, linear, nearly as long as the sepals, 1- to 3-nerved, the stamens about 40, with the filaments about 4-5 mm. long, the ovary ovoid, 1.0-1.5 mm. long, sessile, with a short style about twice the length of the ovary, briefly bifid at the tips and with small acute stigma lobes. Fruit orbicular, sessile, punctate with short separate tufts of stellate hairs, about 3 mm. in diameter, the fringe of relatively stout plumose bristles about 8—9 mm. long; the seed ovoid, about 2 mm. long. Distribution: Plants of central Mexico, growing on hill slopes and chaparrals, between 500 and 1000 m. Flowering in September and October and fruiting from October to December. Mexico: СНІАРАЅ: Manzanillo, E. Palmer 086. GUERRERO: Acapulco, E. Palmer 440; La Providencia, Acapulco, Hancock 12; Iguala Canyon, alt. 2500 ft., Pringle тообо; Arecelia-Fraguas, Coyuca, Hinton 6605; Coyuca, Hinton 6901, 6906; Pungarabato, Coyuca, Hinton 6876; Atoyac, Galeana, Hinton 10001. yatisco: Bolanos, Rose 2860a, 2860b; Guadalajara, Rose & Painter 7380. мехісо: Limones, Temascaltepec, alt. 910 m., Hinton 6724; Paso del Rio, Emrick 102. MicHoacaN: Apatzingan, alt. 2000 ft., Leaven- worth 9 Hoogstraal 1731; between Acahuato and Apatzingan, Leavenworth t$ Hoogstraal 1525; El Ocote, Langlassé 623. The fruits of this species are very characteristic, being nearly orbicular, with short separate tufts of stellate hairs. The small erect sepal appendages and the glabrate leaves make this species an easy one to identify. Н. laevis (type: Rose 2860) which has been identified with many other species actually is conspecific with this. 1949] KO КО LAY—REVISION OF HELIOCARPUS 523 105 100 Мар 5. Distribution of H. occidentalis 6. HELIOCARPUS MEXICANUS (Turcz.) Sprague, in Kew Bull. 272. 1921. (Т.: Galeotti 4154). Adenodiscus mexicanus 'Turcz. in Bull. Soc. Nat. Du 19?:504. 1846. ое mexicana 'Turcz. in D. cit. 311:230. 1858 H. glanduliferus Robinson, in Contr. U. S. Nat. Herb. 5:127. 1897. (Т.: Heyde 281). H. glabrescens Hochr. in Ann. NR et Jard. Bot. Genéve 18:122. 1914. (T.: Galeotti 4154). H. costaricensis Sprague, in Kew Bull. 349. 1923. (Т. РИ ет 13022). H. belizensis Lundell, in ri 2:2. 1941. (T.: Gentle 2273) H. yucatenensis Millsp. in herb. Small trees or shrubs 5—6 m. high; older branches dark brown, glabrate, slightly rugose, punctate irregularly with minute white lenticels; younger branches and inflorescence axes very lightly scurfy-pubescent, with small clusters of simple and stellate hairs, nearly glabrate. Leaves broadly ovate to ovate-lanceolate, the more mature ones broadly ovate, the younger ones around the inflorescence usually ovate- lanceolate, 5-costate at the base, not lobed, 12—14 cm. long and 6-10 cm. wide, acute to acuminate, base rounded or slightly obtuse, irregularly and bluntly serrated, the basal serrations glandular, upper surface dark green, glabrate with few sup- pressed stellate hairs, lower surface more pubescent with fine, weak, crisped, stellate [Vor. 36 524 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 6. Н. mexicanus 95 90 55 Map 6. Distribution of H. mexicanus 1949] КО КО LAY—REVISION OF HELIOCARPUS 225 hairs, becoming nearly glabrate at maturity. Inflorescences gynodioecious, axillary, large and very leafy; the hermaphrodite large and spreading, very leafy, about 20 cm. long and 15 cm. wide, the cymes of about 20 flowers, rather loose, the flowering peduncle mostly 3-radiate, about 2-3 mm. long, usually subtended by a pair of small blunt bracteoles, the pedicel about 6—8 mm. long, the buds obovoid, slightly constricted in the middle towards the base, rusty brown, with small appendages at the tips of the sepals, the sepals usually 5, rarely 4, spatulate, about 5—6 mm. long, 1-2 mm. wide, rusty brown and glabrate without, lighter brown and glabrate within, with linear, about 1.5 mm. long, erect appendages on the tips, the petals 5, rarely 4, linear, 1-nerved, 3-4 mm. long, the stamens 24-30, with the filaments as long as the petals, the ovary ellipsoid, 1-2 mm. long, borne on a very short gyno- phore, with a style about 3 times the length of the ovary, briefly bifid, and with acute, slightly spreading stigma lobes; the pistillate nearly as large as the herrnaph- rodite, the cymes of about 20 flowers, greatly condensed and crowded in nodose clusters, the flowering peduncles 3-radiate, short, less than 1 mm. long, the pedicels about 1 mm. long, the buds about 2-3 mm. long, appendaged at the tips of the sepals, the sepals 5, linear, about 3 mm. long, appendaged at the tips, rusty brown and glabrate without and within, the petals none, the staminodes about 20-30, the ovary ovoid, less than 1 mm. long, borne on a very short gynophore, with a short style, briefly bifid and with acute stigma lobes. The fruit ellipsoid, glabrate, sur- face red-glandular, about 6—8 mm. long and 3 mm. wide, with few ridges, borne on a bristly gynophore 2—8 mm. long, the fringe usually of one row of plumose bristles 6—8 mm. long; the seed pyriform, blunt-pointed, about 3—4 mm. long. Distribution: Plants of southeastern Mexico and northern Central America, growing along roadways, river banks, or on hill slopes; in thickets of bushy slopes or on hill tops in open places. Flowers from September to October and sometimes as late as December, and fruits from November to February. MEXICO: CAMPECHE: us Lundell 050, 1102. ОАХА айту пос mentioned, Ghiesbreght 51. VERA CRUZ: Zacuapan, Linden 808; Cordillera, LEE 456 Zacuapan and vicinity, Purpus 2227; Mirador, Ізгілік 523; Mirador, Linden 858; rocky barranca near Gaucho, Viejo, Purpus 157044. YUCATAN: Pocoboch, a tolol, E ей San Anselmo, Gaumer 1234; Calotmul, Gaumer 2302; Chichankanab, Gaumer 2275; Xnocao, Gaumer 234 24177. RITISH noon El Cayo District, Vaca, Gentle 2273; Jacinto Hills, alt. 600 ft., Schipp S-589. GUATEMALA: ALTA VERAPAZ: southwest of Lanquin, alt. 600-1000 m., Steyermark 44076. CHIMALTENANGO: along Río Guacalate, southwest of Chimaltenango, alt. about 1700 m., Standley 80064, 81067; appro жуа Johnston 1106, 1332. CHIQUIMULA: Volcan Ipala, near Amatillo, alt. 900-1510 m., Steyermark 30377. GUATEMALA: exact кн lacking Aguilar 61; alt: n m., » Morales 1.122. JUTIAPA: Volcán Suchitan, Be est ncion Mita, alt. 600—205 Steyermark 31012. SACATEPEQUEZ: near Das ie 1560- n m. m^ 50888; near ой alt. 1500—1600 m., Standley 64273. SANTA ROSA e Cerro Redondo, Steyermark 52218; near Cuilapilla, и фо 900 m., Standley UM Void der alt. 5000 рр., Heyde 9 Lux 3056. DATA INCOM- PLETE: Hey e : HONDURAS: EL PARAISO: Guinope, alt. 1430 m., Rodriguez 1679. [VoL. 36 526 ANNALS OF THE MISSOURI BOTANICAL GARDEN EL SALVADOR: gms of San Salvador, Remson 61; vicinity of San Salvador, alt. 650-850 m., „ Standley 9660 CosrA RICA: sAN me n o Torres, alt. 1100 m., Tonduz 7451; bords de Río Torres prés San Francisco de немој alt. 1100 m., Tonduz 8253. This species has been described by most authors as having sessile fruits. Actually the fruits are borne on a gynophore the length of which varies from 2 to 8 mm. Many species have been named because of the extreme variability of the length of the gynophore. The correlation of the flowering with the fruiting specimens has been greatly confused. H. mexicanus (type: Galeotti 4154) can be recognized easily by its brown glabrate fruits borne either on a very short or on a slightly longer gynophore. In flower, this is one of the easiest species to identify because of the large erect ap- pendages on the tips of the sepals and the nearly glabrate ovate-lanceolate leaves, as well as by the leafy axillary inflorescence. This species actually is a link between the sessile-fruited and the stipitate groups. Іс has the characteristics of both the groups. 7. HELIOCARPUS APPENDICUI ATUS Turcz. in Bull. Soc. Nat. Moscou 31!:226. 1858. (T.: Linden 2065). H. chontalensis Sprague, in Kew Bull. 350. 1923. (Т. Lévy 483). Trees 12-14 m. high; older branches dark brown, scurfy, ferruginous-tomentose with both simple and stellate hairs, rarely glabrate, ridged, punctate irregularly with small, white lenticels; younger branches and inflorescence axes covered with thick, scurfy, ferruginous tomentum. Leaves broadly ovate, sometimes obscurely 3-lobed, 14—16 cm. long and 12-14 cm. wide, 5- to 7-costate at the base, acum- inate, base subcordate or rounded, with two conspicuous leafy auricles at the basal sinus, each about 5—7 mm. wide and 4—5 mm. long, unequally and bluntly serrated, the basal serrations glandular, upper surface dark green, glabrate, with few sup- pressed, compact tufts of stellate hairs, lower surface whitish, with dense stellate tomentum, petioles relatively stout, about 6—8 cm. long, covered with dense scurfy, ferruginous tomentum. Inflorescences hermaphrodite, usually terminal, rarely axillary, slightly leafy, about 15 cm. long and 12-14 cm. wide, the cymes of about 16—18 flowers, rather loose, the flowering peduncles 3-radiate, about 2-3 mm. long, the pedicels 3—4 mm. long, the buds obovoid, slightly constricted in the middle, rather large for the genus, about 6 mm. long, not appendaged at the tips of the sepals, the sepals 4, linear, 6—7 mm. long, light green with short separate tufts of stellate hairs without, glabrate, yellow-brown within, the petals 4, spatulate, 3- nerved, 3—4 mm. long, the stamens about 30, with filaments about 4—5 mm. long, the ovary ellipsoid, 1-2 mm. long, borne upon a gynophore about the length of the ovary itself, with a style about twice the length of the ovary, bifid about half its length and with small, acute, spreading stigma lobes. The fruit suborbicular, about 5 mm. long and 3-4 mm. wide, densely tomentose, borne upon a gynophore about 5-8 mm. long, with 2—3 pairs of plumose bristles, the fringe of 1 or 2 rows of plumose bristles about 6—8 mm. long; the seed pyriform with a shallow groove, slightly stellate-pubescent, about 2-3 mm. long. 1949] КО KO LAY—REVISION ОҒ HELIOCARPUS 9 o Map 7. Distribution of H. appendiculatus 527 [Vor. 36 528 ANNALS OF THE MISSOURI BOTANICAL GARDEN Distribution: Plants of central Mexico and Central America: a species of rela- tively wide distribution, usually growing along river banks or roadways in sec- ondary growth subjected to seasonal floods; at times forming a pure stand in moist places, usually in heavy red clay loam, at various altitudes but most abundant above 1000 m. Flowers from December to about the middle of March and fruits per- sistent up to about the middle of June. MEXICO: SAN LUIS POTOSI: near Tancanhuitz, Nelson 4393. TABASCO: locality lack- ing, alt. 300 m., Linden 1600; Teapas, Linden 2 VERA CRUZ: Fortuno, Coatzacoalcos River, alt. 30- 50 m., L. Williams 8273, 8288, 8280, Sa, 8526, 8527; Rio Seco prope Cordoba, Woronow 2063; Mirador, Linden 469; near . Rose 9 Hougb 4314. GUATEMALA: ALTA VERAPAZ: Pansamala, alt. 3800 pp., J. D. Smith 1723; 2% miles west of Cubilquitz, alt. 250-300 m., Steyermark на ай from Senahu (о Actala, M 6 Hay 3305, 3306, 1344 Cubilquitz, alt. 350 m., Tuerckbeim 7828. HUEHUE- NGO: between Maxbal and Xoxlac, Sierra de los Cochusq quan, alt. 1500-1600 m., Steyermark 48087; pr о Chible, between Finca San Rafael and Ixcan, Sierra de los atanes, alt. 1200-2000 m., Steyermark 401 а ATLANTIDA: некелі Valley, near Tela, alt. 20-600 m., Standley 54006. NICARAGUA: SEGOVIA: region of Braggman’s Bluff, Englesing 136, 140; Chontales, alt. 600 m. dod td 483; Chontales, Tate 31 (384). Costa Rica: cARTAGO: Rubber Plant se ae ини Station, Turrialba, Aguilar 07100. GUANACASTE: upper slopes of Cerro San José de Libano, alt. 500—900 m., Dodge, Hanckel 9 Thomas 6370. san JOSÉ: vicinity of El General, alt. 700 m., Skutch 3022; vicinity of El General, alt. 670 m., Skutch 4226. БАТА INCOMPLETE: Rowlee 9 Rowlee 212; Tibarcia, alt. 980 m., Solis R. pop Santiago, Brenez 655. An extremely easy species to identify due to the presence of two conspicuous auricles at the basal sinus of the relatively large leaf. Тһе fruits are suborbicular and tomentose. 8. HrLiOCARPUS AMERICANUS L. Sp. Pl. ed. 1, 448. 1753. (Т. in Br. Mus.).® ° H. tomentosus Turcz. in Bull. Soc. Nat. Moscou 31:255. 1858. I Tia 857). H. americanus var. typica К. Schum, in Martius, Fl. Bras. 12?:141 Small trees 3—5 m. high; older branches light brown, I4 smooth, irreg- ularly punctate with many small white lenticels; younger branches and inflorescence axes lightly tomentose with both stellate and simple hairs. Leaves ovate, not lobed, 10-12 cm. long and 6-8 cm. wide, 5-costate at the base, acuminate, base rounded or obtuse, never cordate, irregularly and bluntly serrated, upper surface dark green, glabrate with few highly suppressed stellate hairs, lower surface whitish, very densely tomentose with both long stellate and simple hairs; petioles relatively stout, about 4 cm. long, very densely tomentose. Inflorescences hermaphrodite, terminal, about 15 cm. long and 12 cm. wide, sometimes very slightly leafy, the cymes of about 18—20 flowers, densely crowded in nodose clusters, the flowering peduncles 3-radiate, short, about 1 mm. long, the pedicels about 1-2 mm. long, the buds obovoid, slightly constricted towards the base, without appendages at the tips of the sepals, the sepals 4, spatulate, 4—5 mm. long, light green and densely tomentose without, glabrate, yellow to yellow-brown within, the petals 4, linear, 1-nerved, 8]Jour. Bot. 36:131. 1898. 9Bot. Gaz. 61:256. 1923. 1949] KO КО LAY—REVISION OF HELIOCARPUS 529 about 2-3 mm. long, the edges slightly ciliate, the stamens about 20, with the filaments about 3 mm. long, the ovary ovoid, about 1 mm. long, borne on a short gynophore, with a style about 2—3 times the length of the ovary, bifid about one quarter of its length and with acute, spreading stigma lobes. Fruit ovoid, densely tomentose, becoming slightly less so with age, about 3—4 mm. long and 2-3 mm. wide, borne on a long bristly gynophore about 5-8 mm. long, with 2-3 pairs of plumose bristles, the fringe of two rows of plumose bristles about 4—6 mm. long; the seed ovoid, about 1.5 mm. long and 1.0 mm. wide, with a distinct groove in the middle, slightly stellate-pubescent. о Fig. 8. H. americanus Мар 8. Distribution of H. americanus Distribution: Plants of central and southeastern Mexico, growing at altitudes between 400-1200 m. Flowers in January and February and the fruits are per- sistent up to the middle of May. Mexico: oaxaca: locality not mentioned, Galeotti 4162, 4162b. PuEBLA: Huauchi- nango, convalles тен, alt. 1100 m., Fróderstróm & Hulten $60. VERA CRUZ: Zacuapan and vicinity, Purpus 2226, 2387; Zacuapan, Purpus 8275; Mirador, Liebmann 475; Orizaba, niit 340, 341, 882, 888, 922; Vallee de Cordova, Bourgeau 1810, 1974; Mirador, alt. m., Linden 857; Cordova, Ј. С. Smitb 284; near Jalapa, Rose 9 Hough 4304; Estacion El 1. Orizaba, Conzatti 1604; near Orizaba, alt. 4000 ft., Pringle 6106; Sanborn Orcutt 3068; circa Cordoba, Woronow 2047; Río xo is Cordoba, Woronow 3036; Cordoba, Greenman 161; barranca of Chevastl near Huatusco, Rozynski 782. DATA INCOMPLETE: Sumichrast 882. The identity of H. americanus is based on the interpretations by E. G. Baker? and T. A. Sprague.® This species has very characteristic ovate-lanceolate leaves, which are densely tomentose only on the lower surface. [Vor. 36 530 ANNALS OF THE MISSOURI BOTANICAL GARDEN 9. HELIOCARPUS DONNELL-SMITHU Rose, in Bot. Gaz. 31:110. 1901. (Т. J. D. Smitb 1722). H. Caeciliae Loesener, in Fedde, Rep. Spec. Nov. 12:227. 1913. (T.: Seler 4976). H. borridus Lundell, in Bull. Torr. Bot. Club 66:597. 1939. (T.: Lundell 8 Lundell 7821). H. cuspidatus Lundell, in Phytologia 2:2. 1941. (T.: Gentle 2297). H. floribundus Lundell, loc. cit. 1941. (T.: Gentle 1534). Small trees about 10 m. high; older branches glabrous, smooth, yellow to yellowish brown, irregularly punctate with white lenticels; younger branches and inflorescence axes slightly pubescent with short stellate and simple hairs. Leaves broadly ovate, 12—14 cm. long and 8—10 cm. wide, not lobed, 5-costate at the base, shortly acuminate, base rounded or obtuse, irregularly serrated, lower serrations glandular, upper surface dark green, essentially glabrous with short greatly sup- pressed stellate hairs, lower surface lighter green, slightly more pubescent than the upper, nearly glabrate, with short stellate pubescence; petioles glabrous, smooth, 6—8 cm. long. Inflorescences gynodioecious, usually terminal, rarely axillary; the hermaphrodite large and spreading, about 18 cm. long and 15 cm. wide, the cymes of about 20 flowers, rather loose, the flowering peduncle mostly 3-radiate, rarely 2-radiate, 2 mm. long, the pedicels 4-5 mm. long, the buds obovoid, slightly con- stricted in the middle, not appendaged at the tips of the sepals, the sepals 4, linear, 4—6 mm. long and 1-2 mm. wide, light green with stellate pubescence without, glabrate, yellow to yellowish brown within, the petals 4, linear-spatulate, 3-nerved, slightly shorter than the sepals, about 4 mm. long, the stamens 12—16, with fila- ments about as long as the petals, the ovary ellipsoid, about 1-2 mm. long, borne upon a gynophore nearly as long or very slightly shorter than the ovary itself, with a style 3—4 mm. long, bifid about one quarter of its length and with stigma lobes acute, spreading; the pistillate greatly condensed and crowded, about 5 cm. long and 5 cm. wide, the cymes of about 20 flowers, crowded in nodose clusters, the flowering peduncles very short, 3-radiate, bearing 2 small bracteoles, the pedicels 1-2 mm. long, the buds about 2-3 mm. long, not appendaged at the tips of the sepals, the sepals 4, linear, 2-3 mm. long, light green and slightly pubescent with- out, generally brownish and glabrate within, the petals none, the staminodes none, the ovary suborbicular, about 1 mm. long, borne upon a very short gynophore shorter than the length of the ovary, with a style nearly twice the length of the ovary, very briefly bifid, the stigma lobes acute. The fruit ellipsoid, nearly glabrate at maturity, slightly rugose, 5 mm. long and 3 mm. wide, borne on a gynophore 8—12 mm. long, bearing 2—4 pairs of plumose bristles, the fringe of two rows of plumose bristles 5—7 mm. long; the seed obliquely ovoid, 2—3 mm. long and about 2 mm. wide, with a distinct groove in the middle. Distribution: Plants of southern Mexico and Central America. This species is of rather wide continental distribution but its presence in Martinique is difficult to explain, unless through introduction. It is abundant оп the edges of forest, in secondary growth, and on mountain slopes at altitudes from 100 to 1500 m. Flow- ers from December to February, and the fruits are persistent up to June. 221 H. Donnell-Smitbii Бір. 9. ------=--| КО КО LAY—REVISION OF HELIOCARPUS 1949] 3 ` in J ~ Vt Distribution of H. Donnell-Smithii Map 9. [Vor. 36 532 ANNALS OF THE MISSOURI BOTANICAL GARDEN XICO: OAXACA: Tuxtepec, Chiltepec ну а, Calderón 452. QUINTANA ROO: и Lundell 9 Lundell 7821. TABASCO: Rovirosa 120. VERA CRUZ: Mont- zorongo, J. G. Smith 234; Wartenberg, near ‚ине ај rov. Huasteca, Ervendberg 225; Santa Packie. Mell 670; Coatzacoalcoa, isthmus of аке е C. L. Smith 1002; district San Andres Tuxtla, Salto de Chilapan, Seler 4976. Витон Номрокав: Belize District, Gracie Rock, Sibun River, Gentle 1534; El Cayo District, Vaca, Gentle 2207; El Cayo District near Camp 6, Gentle 2355; Inacahual, Stann Creek District, Middlesex, Gentle 2788; Middlesex, тойи 3336; 22 Mile, alt. 200 ft., Schipp 872. GUATEMALA: ALTA VERAPAZ: Arenal, alt. 3000 pp., J. D. Smitb 1722; Savanna, north of Concepcion, = —5 жиз southeast of Finca Yalpemech, near Alta Verapaz-Peten boundary line, alt. 100—11 » Steyermark 45220; southwest of Lanquin, alt. 600-1000 m., Steyer- mark. 44007, pes елей trail to Panzas, Goll 255; road to Sepaciute, Wilson 1078; mount slopes above Finca, Semay, Wilson 108. IZABAL: between Los Amates and P Kellerman 6 6448. SUCHITEPEQUEZ: Mazatenango, alt. 300 m., Kellerman 6068. RAS: ATLANTIDA: vicinity of Tela, alt. 20-600 m., Standley 55518. Ка UA: MANAGUA: vicinity of Managua, Garnier 560. GRANADA: Volcán кыы. “Baker 2490. MARTINIQUE: bois près Fort de France, Habn 1340; St. Pierre, Curran » This species is characterized by its ellipsoid, nearly glabrate fruits borne on a long bristly gynophore, and by its ovate leaves, which are nearly blabrous to glabrate. 10. HELIOCARPUS POPAYANENSIS НВК. Nov. Gen. Sp. 5:341. 1821. (T.: Hum- boldt & Bonpland, in Herb. Paris). ма Turcz. in Bull. Soc. Nat. Moscou 311:226. 1858. (T.: Funck & Schlim americanus var. bopayanensis (HBK.) K. Schum., in Martius, Fl. Bras. 12?:141. 1886. bopayanensis var. Schumanni E. С. Baker, in Jour. Bot. 36:132. 1898, popayanensis var. Purdiei E. G. Baker, loc. cit. bopayanensis var. trichopoda (Turcz.) E. G. Baker, loc. cit. 189 bobayanensis var. fum Hochr. in Ann. Conserv. et Jard. Su Genéve 18:116. 1914. (T.: Ва diclinus Hochr. b oc, cit, 117. 1914. (T.: H. H. Smitb 1008). boliviensis Hochr. loc. cit. 118. 1914. (T.: Bang 1401). stipulatus Hochr. loc. cit. 121. 1914. (T.: Poeppig 3102). Rosei Hochr. loc. cit. e 1914. (T.: Bang 2305). americanus acc. to E. E. Wats. in Bull. Torr. Bot. Club 50:123. 1925, not 1. australis E. E. дей loc. cit. 124. 1923. (T.: Hassler 557) udi ts. loc. cit. 126. 1923. (T.: Pittier 3082 subtrilobus Б. іп Вот. Gaz. 61:257. 1923. (Т.: Fendler I277B). mmm m m soria = Trees usually about 8-10 m. high or more; older branches glabrate, slightly rugose, cream to brown, irregularly punctate with few, rather small, white lenticels; younger branches and sometimes a few of the older branches covered lightly with ferruginous tomentum of both stellate and long coarse simple hairs. Leaves usually 3-lobed with acuminate apices, but sometimes only obscurely so, about 16—20 cm. long and 14—18 cm. wide, base markedly cordate on mature leaves, younger leaves usually with rounded bases, irregularly serrated, upper surface generally dark green, glabrate with short ferruginous stellate tomentum, especially on the veins and veinlets, lower surface light green, slightly more pubescent than the upper, the 1949] КО КО LAY—REVISION ОЕ HELIOCARPUS 235 pubescence varying from densely stellate to thinly suppressed stellate, the principal veins usually with simple hairs about 2 mm. long; petioles 6-8 cm. long, usually densely covered with both stellate and simple hairs. Inflores- cences gynodioecious, usually terminal; the hermaphrodite about 12 cm. long and 14 cm. wide, the cymes of about 12-16 flowers, the flowering peduncles 3-radiate, about 1-2 mm. long, the pedicels about 2-3 mm. long, the buds obovoid, slightly constricted towards the base, without appendages at the tips of the sepals, the sepals 4, spatulate, generally 3- nerved, about 5 mm. long, light green and densely tomentose without, glabrate and yel- lowish brown within, the petals 4, spatulate, about 3—4 mm. long, slightly ciliate, the sta- mens about 12-16, with the filaments about Fig. 10. Н. popayanensis 3-4 mm. long, the ovary suborbicular, laterally compressed, about 1.0—1.25 mm. long, borne on a gynophore about 0.75 mm. long, with a style about twice the length of the ovary, bifid about one quarter of its length and with the stigma lobes spreading, each with 3 acute lobes; the pistillate rather large, much larger than the hermaphrodite, about 14 cm. long and 20 cm. wide, the cymes of 12-16 flowers, rather condensed in nodose clusters, the flowering peduncles 3-radiate, less than 1 mm. long, the pedicels about 1 mm. long, the buds about 3 mm. long, the sepals 4, linear, about 3 mm. long, light green, stellate-tomentose without, glabrate, cream to light brown within, the petals none, the staminodes about 12, the ovary suborbicular, about 1 mm. long, borne on a short gynophore, with a style about twice the length of the ovary, bifid about one quarter of its length and with spreading stigma lobes, each with 3 acute lobes. The fruit ellipsoid to ovoid, with many short tufts of stellate hairs, slightly pubescent, about 5-6 mm. long and 2—3 mm. wide, borne on a bristly gynophore about 1.0-1.5 mm. long with 2-3 pairs of plumose bristles, the fringe of two rows of plumose bristles about 8—12 mm. long; the seed ovoid, about 2 mm. long, with a shallow depression in the middle. Distribution: A species which extends from Costa Rica to northern Argentina. It is primarily South American and is the only species known there. In southern Central America and northern South America it is found in highlands above 1000 m. altitude, while in Paraguay and Argentina it is usually at lower elevations. H. popayanensis frequents steep slopes in forested river valleys, thickets, or in sunny bushy slopes, usually in second growth in cut-over forests. It is abundant in rain or cloud forests or on the edges of forests at stream sides. Flowers in its northern [ VoL. 36 534 ANNALS OF THE MISSOURI BOTANICAL GARDEN Map 10. Distribution of H. popayanensis range from December to January and in its southern range from May to June. Fruits persistent in the north until about the middle of March and in the south to about the end of September. Costa Rica: ALAJUELA: hills west of Zapate, San Carlos, alt. 1550 m., A. Smith NY 1263; Zacero, upper continental divide, alt. 1800 m., А. Smith H1358. HEREDIA: Vera Blanca de Sapiqui, near slope of Central Cordillera, bire Poas and Barba volcanoes, alt. 1680 m., Skutch 3346. saN ЈОЗЕ: La Palma de San Ramon, Brenez 6360; эы a: alt. 5700 ft., Stork 1158, DATA INCOMPLETE: Charco, Goicochea, alt. 1400 m., Jimen 993. PANAMA: CANAL € Cerro Gordo, near Culebra, alt. ve m., Pittier 2305; Barro Colorado Island, 218; Canal Zone and vicinity, C.16, back of Curu Harvey 5258; Barro rf rd Island, Shattuck 669, 754; пеаг lab, Barro Colorado ene 1949] КО КО LAY—REVISION OF HELIOCARPUS 299 Woodworth & Vestal 730, 748; Barro Colorado Island, Wetmore 9 Abbe 108; Empire Station, Hayes 437. cHiRIQUÍ: valley of the upper Rio Chiriqui Viejo, W bite 29; around © vicinity of "New Switzerland" central valley of Río Chiriquí Viejo, alt. 1800-2000 m., Allen 1398; vicinity of Cerro Punta, alt. 2000 m., Allen 1519. cocrÉ: hills s ç El Valle de Antón, alt. 600-800 m., Allen 2849, 2012; El Valle de Antón, alt. 600- m., Hunter & Allen 304. DARIEN: locality not mentioned, Macbride 2713. TRINIDAD: St. Ann's Cascade, Broadway 6569; Santa Cruz, Simmonds 281; Lady eia did Road, St. Ann’s, кшш 0200; Knagg's Hill, R. O. William: 12671; St. Cruz, collector unknown 649, COLOMBIA: ANTIOQUIA: да of Medellin, Toro 206. Boyac 130 m. n. of Bogota, El Umbo, alt. 3700 ft., Lawrance 500. cauca: El Tambo in b. alt. 1700 m., Sneidern 434; highlands of Рорауап, alt. 1400-2000 т., Lebmann 5502; Rio Toribio region, alt. 100-1200 m., Espiña 9 Giacometto 20861; cerca de Popayan, matorrales en Río Blanco, alt. 1800 т., Arbelaez 9 Cuatrecasas 5708. MAGDALENA: Santa Marta, alt. > 46 20 ‚ Н. Н. Smith 1908; Pita, André ARINO: frontera Colombo-ecuatoriana, selva higrofila del Río San Miguel, junto a la desembo-cadure del Río Conejo, alt. 3 4 Cuatrecasas IO NORTE DE SANTANDER: Cordillera Oriental, region del Sarare, Que- entre Chorro Colorado y Bata, alt. 1300 m., Cuatrecasas, Schultes © Smith 12237; Cordillera Oriental, region del Sarare, La Cabuya, alt. 1300 m., Cuatrecasas, Schultes 9 Smitb 12590; Cordillera Oriental, region del Sarare, entre el Alto del сады el Alto де Santa Ines, bosques, alt. 1800-2000 m., реді жеріні Schultes & Smith I VENEZUELA: DISTRITO FEDERAL: Caracas, alt. 3000 pp., Funck © S 150; Carteza, San Pablo, Pittier E DATA INCOMPLETE: Las Trincheras, Warming 313; al sur de Río AME n. 1360 m., Saer 778 Е: near mouth of Río Macauhan (tributary of Río Yaco), lat. 9.20" S., NS: jm p "ұу. Krukoff 5261. Ecuapor: AZUAY: along Rio Patul pr же; Hacienda Yubay and Hacienda San José de Caimotan, in region of Sanaguin, alt. 85 ., Steyermark 52754; Hacienda Yubay, at Sanaguin, on south side of Rio Patul, alt. 1» m., Steyermark 52604; Rio Norcay between Río Gamolotal and Río Norcay, alt. 1095— -1370 m., Steyermark 52878. CHIMBORAZO: Chimbo, alt. 1000 m., Rimbach 288. DATA INCOMPLETE: vicinity of Ventura, Rose & Rose 23516. PERU: CAJAMARCA: Guerocotillo, Prov. Contervo, alt. 2000 m., Weberbauer 7124. cUzco: Río Chaupimayo, Convencion, Soukup 700. HUANUCO: Манон, alt. 900-1000 m., Weberbauer 3432; Cueva Grande, estacion near Pozuzo, alt. 3500 ft., Macbride 4765; Ушуўаси, Sawada IOI. JUNIN: Chanchamayo Valley, alt. 1500 m., Schunke 200, 203; Pichis Trail, Dos de Mayo, alt. 1700-1900 т., Killip & Smith 25864. LoRETO: Muna, Macbride 4065; lower Río Huallaga, alt. 155-210 m., Klug 3076; west of Tarapota, Spruce 4558. SAN MARTIN: Juan Jui, alto Río азза, ile 400-800 m., Klug 4305. DATA INCOMPLETE: Poeppig 1804, 3102; Ruiz © Pavon 39/5, 30/9. BOLIVIA: COCHABAMBA: Incachaca—S. Antonio, alt. 1500 m., Werdermann 212 paz: Calapampa, Coroico, Bang 2305; Mapiri, Bang 1401; Солай Tipuani, Bang 1455; Guanai, alt. 2000 ft., Rusby 1492; Beni River, Rusby Ы miles: Region von Cori- pata, Hacienda "El Choro", Buchtien 8125; Province of S. Yungas, basin of Río Bopi, San Bartoloma (near Co). alt. 750-900 m., Krukoff 4020, psc 10400. SANTA CRUZ: E 2. of Tecahuasi, alt. 1600 m., Cardena МА: CATAMARCA: Ре! Alto, Bategua, alt. Eo m., Venturi 7074. MISIONES: Bs of чалын Aguirre, alt. 100 m., Curran 0, I7; vicinity of P uerto Leon, alt. 75-100 m., Curran 700, 718. TUCUMAN: Бета $ап Pablo, alt. 600 m., Venturi 1002, 1002С; [Vor. 36 536 ANNALS OF THE MISSOURI BOTANICAL GARDEN Famailla, San Rafael, Venturi 9205; locality not mentioned, Hauman 9206. Hawan: Oahu, planted in foothills for reforestation, Degener 10843. The plants of this species have leaves whose diversity of shape and size is unparalleled in the whole genus. А general tendency is for the plants in the cen- tral range of the species to have leaves more definitely and acutely 3-lobed than those towards either the northern or the southern limits, where the leaves fre- quently may be quite undivided. Тһе pubescence varies from slightly tomentose to nearly glabrate. The constant characters throughout its wide distribution are the hirsute induments on the lower nerves, the sepals without appendages, the 3- parted stigma lobes, and the fruits, which are ellipsoid to ovoid and slightly tomentose. 11. НЕШОСАКРЏ5 NODIFLORUS (Donn. Sm.) Donn. Sm. & Rose, in Contr. U. S. Nat. Herb. 5:126. 1897. H. polyandrus var. nodiflorus Donn. Sm. in Bot. Gaz. 23:240. 1897. (Т.: Heyde 9 Lux 4329). H. excelsior Morton, in Jour. Wash. Acad. Sci. 27:307. 1937. (T.: Skutch 2250). Н. Gentlei Lundell, in Phytologia 2:2. 1941. (T.: Gentle 1787). mee Fig. 11. Н. nodiflorus Trees about 10-18 т. high; older branches glabrate, longitudinally ridged, light brown, punctate irregularly with few white lenticels; younger branches and inflorescence axes scurfy, slightly ferruginous-pubescent with both simple and stellate hairs. Leaves 3-lobed, sometimes obscurely so, generally medianly oblique, large, 16—18 cm. long and 15—18 cm. wide, 5- to 7-costate at the base, acute to 1945] KO KO LAY—REVISION OF HELIOCARPUS 3537 Map 11. Distribution of Н. nodiflorus acuminate, base usually cordate, sometimes slightly rounded, rather regularly and doubly serrated, lower serrations slightly glandular, upper surface dark green, glabrate, with few tufts of short stellate hairs in the nerve axes, lower surface light green, slightly more pubescent than the upper, especially in the nerve axes; petioles large and stout, about 10 cm. long, lightly ferruginous-stellate-pubescent, scurfy; stipules sometimes persistent. Inflorescences gynodioecious, usually terminal, large and spreading; the hermaphrodite about 15 cm. long and 18 cm. wide, the cymes of about 15—20 flowers borne in compact nodose clusters, the flowering peduncles 3-radiate, about 2 mm. long, the pedicels 1—3 mm. long; the buds obovoid, slightly constricted in the middle towards the base, sometimes with small acute appendages on the tips of the sepals, the sepals 4, spatulate, 4-5 mm. long, light green with stellate tomentum without, glabrate and yellowish brown within, the petals 4, linear, 1-nerved, about 4 mm. long, the stamens about 20-24, with the filaments about 4 mm. long, the ovary ovoid, slightly compressed, about 1.5 mm. long, borne оп а short gynophore less than 1 mm. long, with a style about 3-4 mm. long, bifid about half its length at anthesis and with the stigma lobes acute and spreading; the [VoL. 36 538 ANNALS OF THE MISSOURI BOTANICAL GARDEN pistillate about as large as the hermaphrodite, the cymes of about 20 flowers, rather compressed and crowded in nodose clusters, the flowering peduncles 3-radiate, less than 1 mm. long, the pedicels about 1-2 cm. long, the buds about 2-3 mm. long, usually not appendaged at the tips of the sepals, the sepals 4, linear, about 2-3 mm. long, light green and stellate-pubescent without, glabrate and yellowish brown within, the petals none, the staminodes about 12—16 or none, the ovary ovoid, small, about 1 mm. long, slightly compressed, borne on a short gynophore, with a style slightly longer than the ovary, briefly bifid and with the stigma lobes acute. Fruit orbicular, slightly tomentose with short separate tufts of stellate hairs, ut 5-6 mm. in diameter, borne on a bristly gynophore 10-15 mm. long, bearing 2-3 pairs of plumose bristles, the fringe of 1 or 2 rows of plumose bristles 6-8 mm. long; the seed obliquely ovoid, about 2-3 mm. long, with a distinct groove in the middle. Distribution: A species primarily of Central America, extending into south- eastern Mexico. It is common in clearings and in wet thickets, nearly always in secondary growth besides rivers or in sheltered valleys, usually at altitudes from 1000 to 2500 m. Flowers from January to February, the fruits persisting until about the middle of April. 5н HoNDURAs: Gracie Rock, Sibun River, Gentle 1787; Stann Creek Valley, 17 ш. Сене 3202, 3203, 3204; Stann Creek Valley; Big Eddy Creek, Gentle 3470; Little Cocquericot, — bits ира 4177. НІМА NGO: lower апа middle southwestern slopes of Volcán Fuego, above Finca Кш soq along ыбы Espinazo and tributary of Río Pantaleon, alt. 1200-1600 m., Steyermark 52113. EL PROGRESO: Montana Canahu ui, between Finca San Miguel and м of mountain, near upper limits of Finca Caieta, alt. 1600—2300 m., Steyermark 43825. кзсілті.А: between Río Jute and Río Pantaleon, on road between Escuintla and Santa Lucia Cotz, alt. 540—720 m., Standley 63443. QUEZALTENANGO: Pueblo Nuevo, alt. 500—600 m., Steyermark 35467; Colomba, alt. 3000 ft., Skutch 2037. QUICHE: Nebaj, alt. up to 6200 ft., Skutch 1763. SAN MARCOS: between Rodeo and Malacate, alt. 1400-3500 ft., Nelson 3742. SANTA ROsA: Río Pinula, alt. 4000 pp., "m E Ж © 4329; Магапјо, alt. 1100 m., Неу4е 9 Lux 4177. $: ATLANTIDA: Lancetilla Valley, near Tela. alt. 20-600 m., Standley 53854, 55791, 55793, 56640. NICARAGUA: GRANADA: Mombacho, Oersted 14820. Costa Rica: SAN Jost: vicinity of El General, alt. 640-1100 m., Skutch 2250, 2266, 2. It is unfortunate that the type of Н. nodiflorus (Heyde 8 Lux 4329) has flowers with small apical appendages. Тһе appendages in this species actually are rarely present, and if present are extremely small. This species is characterized by its large leaves, the largest known for the genus, which are nearly glabrous on both the surfaces. The fruit is like that of H. Donnell-Smithii, but is more spheroid, slightly more tomentose, and larger in size. It can be distinguished from Н. popayanensis by its large orbicular fruits, as well as by its glabrous leaves. 1949] KO KO LAY—REVISION OF HELIOCARPUS 539 INDEX TO EXSICCATAE Italicized numerals refer to collector’s number s. n. (sine numero) to unnumbered collections; parent thetical numerals refer to the d ed of taxonomic entities conserved in this revision Aguilar, I. 2-2 Ар” Allart, А. 273, 516(10 АПеп, Р. Н. 641, 1308, 1510, 2840, 2012 (10 André, E. 407, 800, К. 024, К. ГАРА & Arbelaez, Cuatrecasas, J. (10). Ariste-Joseph, Bro. А. 370(10). Baker, C 2490 (10). Balansa, B. 2205(1 ). 1455, 1491, 2305(10). Barnes, С. В. & Land, W. С. 285(1). 2. 971(2). Т. п.(3). Вгепеѕ, А. 5:50; 6369 (10). 340, 341, 882, 888, 992 (8). 1200 (1) ; 1574, 1819, 1875, Britton, N. L. & Broadway, W. E. 467 10 Broadway: W. E. 5100, 6560, 9200 (10). 25(10). Conzatti, E 4( Cook, O. F. & Griggs, R. F. 4, 7220). Cowell, J. E 270, 271(10). Cuatrecasas, Ј. 10925, 13262, 25421 а J., Schultes, В. E. mith, E. 2237, 12471, 12590 (10). "scd H. га Curran, Н. М. 7(9);0, 17, 700, 718 (10). Daniel, Bro. 1024(10). Dayton, W. А. 3156(7) Degener, О. 10843(10). Dodge, , Hanckel, R. & Thomas, W. 5. 637900). Edwards, J. В. 465(11). Emrick, G. ТОЛ (5). 136, 140 (7). Ervendberg, L. С. 225(9) Espifia, —. & Giacometto, —. 20861(10). Fendler, А. 1277, 1277В(10). Ferris, R. 5. 6025(5). Fiebrig, К. 2а(10). Fróderstróm, H. & Hulten, E. 860(8); 780(9). ProsG:S M: 0). Funck & Schlim 150(10). Galeotti, H. 415446); 4162, 4162В(8). 560(9). Garnier, A. Gaumer, С. F. 1234, 1315, 2275, 2302, a с. 24177 (6). entle, P. H. 2273(6); 1534, 2207, 2355, “2781, 2203 (9): ТЫҢ E 347001). Gentry, H. S. 1644, 1709, 2440, 5069. 6607 (3); 4792 (4). Ghiesbreght, M. 51(6). Goldman, E. A. 226(3); 70(8); 37(9). Goll, G. Р. 255(9). Gonzalez, J. 860(4). Haha, L. Harvey, D. R. 5258(1 0). Hassler, E. 557, 557a, 11856(10). Hauman, —. 0 Науев, 5. 437, 047, 1015(10). Heyde, E. Т. 381(6); 637, 658(11). E. T. & Lux, E. 3956(6); 4177, 2723, 4053, 5119, 6736, 7077, 12449, 12477, 13246, 14831(1); 2843, 5100, 5360, 5416, 7036, 7155, 7236, 7243, 13308 (2) ; 4265, 6605, 6621, 6724, 7, 6901, 6905, 6006, 9801, 0802, 10001(5). Holton, I. Е. 770(1 Hunter, А. А 304(10). 1106, 1332(6). Jones, M. E. 27153, 07400(1). Jorgenson, P. 4500(10). Juzepezuk, —. 1461(1). Kellerman, W. А. 6068, 6448 (9). ' Kerber, E. 276, 483(8). Killip, E. P. 8248(10). Killip, E. P. & Smith, A. C. 10461, 25864(10). Klug, C. di 3627, 4395 (10 Krukoff, B. А. 5261, 10249, 10283, 10400 (10 15000, 15518, щий. Е. 572(1); 623(5). ). Lawrance, А. E. 540 Leavenworth, W. C. & Hoogstraal, H. I731, 1525(5). Lehmann, Е. C. В.Т. 553, 5502 (10). т J. G. & Lemmon, —. s.n., 251 “P MAN H. 1404(3); 661, 1158 (4). ‚Р. 483(7). . 4) 522, 523(6); 477(7); 471, P £A (8 Linden, J. $58, 808(6); 1609(7); 857 Lundell, C. L. 959, 1102(6); 4177, 0516 Lundell С. L. & А. A. Lundell 7821(9). Lyonnet, E ); 1100, E 2) MacBride, J. F. 2713, 4065, 4765(10) Martinez, M. 5 С H 222(8) Maxon, №. В. 4751 Maxon, W. R. & RATS `2156(6); 3305, 3306, 3322(7). n R. 92(8). Mell, C. D. 670(9)- e Y. 809(1); i 7141(10). i 5 (10). Morales, ]. 1.122(6). 3634(4). 884, 1037, 1040(8). Mutis, J. С. 1474, 1478, 3921, 4383 (10). Narvaez-Montes, M. & Salazar, А. Е. 860 (4). Nelson 3, 1485, 1818, 3520, E. 124 7046(1); 2208 6070 (2); 3400 (6); 4383 (7); 70, 110(8). Orcutt, С. E 4463, 4465(5) ; 3068 (8). 3057 (31 3130, 4012 (4). 10). 7 (2); 97, 100, тот, 620 (3) ; 99, 647, 732, 7330) 440, 086(5). Pennell, F. ХУ, & Killip, E. P. 6337, 8007 10 Piper, С. V. 5532, 5612(10). Pittier, Н. 13022(6); 273, 2305, 3062, 5302, 11075, 12605, 13022(10). Poeppig, E. 1904, 3102, ar Pringle, 1791, 8 719, 9602, 0603 (1); 8604(2): D 2. Purpus, 7524(2); 2227 . А. 3062 15704а(6); 2226, 2357 а ). Кесога, 5. ]. & „КЕ Н.25, 6-00 (6); 0058 (7); 4. ). HI 285(10). Reko, B. P. 4448, 4571(1). Renson, C. б1(6). Rimbach, А. 288(10). Rodriguez, J. У. 1670(6). [ Vor. 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN “pun S. т A STONE "y 2660a, + ose, J. М. & ai da `V. 4335, 4565, 48280); 4314(7); 4304(8). Rose, J. N. & Painter, J. H. 7390, 7537, 7640 (1); 7389 (5 5). Rose, J. М. & Rose, С. Rose, А a Seley, P Pe 4. 7 N. 120(9). .W.& v: H. E. Kozy I K. 782 ( i & Pavon, v 30/8, 30/9 (10). 1402, 1493 (10). е 3090(1); 1694 23518(10). . C. & Russell, Р. С. 212(7). ; 872(9). 10). . 4106, 4820(1); 5361(2); ; 4976 (9). Shattuck, О. 662, 754(10). Simmonds, М. W. 2ér(10). Skutch, A. F. 3922, 4226(7); 3346(10); 1445, 1763, 2037, 2250, 2266, 3820, 3852 (11). Smith, А. H 1358, Ny. 1263 (10). mith, C. L. 1 (9). Smith, Н. Н. 1908 (10). Smith, J. D. 1723(7); 1722(9). Smith, J. С. 284(8); 234(9). Smith, L. С. 797, 930(1). Sneidern, K. von. 434, 1431 (10). 4(7). 700 (10). Spruce, К. 4558(10). Standley, 10660, 20304, . С. 1930. , 50888, 64273, 78068, 80064, 810576); 54096(7); 55140, 55154, 18(9); 31552(10); 53854, 55701, EA 56640, 63443, 88541 1(11). Standley, P. C. & Valerio, J. 44633, 45148 (7); 44147 (10). 336(9). 7, 31912, 44076, (7); 44068, 45220(9); 52604, 52754, 25/2 (0); 35407, 43825, 52113 (11). Stork, —. (10). Sumichrast, —. 8&82( Tonduz, А. 7451, 8453, 14828 (6) ; 8561 (7). 1949] KO КО LAY—REVISION ОЕ HELIOCARPUS 541 Того, В. А. 206, 784, 846, 887 (10). White, P. 29(10). Triana, J. s.n., 5367 (10). Williams, L. 8273, $288, 8280, 8400, Tuerckheim, H. von 7828, 8500(7). 8520, 8527 (7); 4207, 9957, 10602(10). Urbina, —. 639(1). Williams, R. O. 649, 650, 12671(10). Velasco, L. У. &997(6). Williams, R. 5. 827(10). Venturi, S. 1002, 1002c, 7074, 9205(10). Wilson, C. L. 107, 108(9). Vogl, P. С. 50«(10). Woodworth, R. H. & Vestal, P. A. 730, Warmings, E. 313(10). 746 (10). Weberbauer, A. 3432, 7124(10). Woronow, С. 2333(1); 2963(7); 2047, Werdermann, E. 2126(1 3036 (8). 0). Wetmore, R. Н. & Abbe, E. С. 108(10). NEW APOCYNACEAE ОЕ SOUTH AMERICA DAVID DE AZUMBUJA! AND ROBERT E. WOODSON, JR. Зесохрлттл Adolphii Azambuja, spec. nov. Frutex altissime scandens lactes- cens; ramulis crassiusculis glabris in maturitate conspicue lenticellatis. Folia ovato- elliptica vel lanceolata basi rotundata vel obtusa apice anguste acuminata saepe mucronata 5-8 cm. longa 2.0—4.5 cm. lata firme membranacea vel leviter coriacea glabra supra nitidula subtus opaca ibique albida et tenuissime reticulata; petiolis 0.6—0.7 cm. longis. Inflorescentia terminalis laxe cymosa pauciflora folia sub- tendentia paulo superans; pedicellis 0.7—1.1 cm. longis glabris; bracteis minimis. Calycis laciniae ovato-triangulares acutae 2.5-3.0 mm. longae intus cum squamis parvis longis. Corolla salverformis tubo 6—7 mm. longo in regione staminium insertionis leviter dilatato intus puberulente lobis oblongo-linearibus vel lineari- lanceolatis obliquis 15-17 mm. longis glabris patentibus. Stamina prope basim tubi corollae inserta; antheris oblongo-sagittatis 3.5—4.0 mm. longis dorso puberu- lentis. Ovarium rotundatum 1.2-1.5 mm. altum minute puberulentum; stigmate 1.2 mm. alto subsessili basi dilatato ubique duos annulos superpositos efformat; disco annulare ovario multo minore. Folliculi fusiformi compressi et prope basim maxime dilatati apicem versus gradatim angustati lignosi striati 19—20 cm. longi ca. 3.5 cm. lati; seminibus oblongo-ellipticis coma e fasciculis pilorum opacorum argentorum reflexorum.—BRAZIL: AMAZONAs: Manaus, matas de terras altas a nordeste de Flores, Sept. 14, 1945, A. Ducke 1,758 (fl.); same locality, Aug. 29, 1947, A. Ducke 2,105 (fr.), in Herb. Jard. Rio de Janeiro zo. 60,224. This species is dedicated to my friend Dr. Adolpho Ducke. Although similar to S. Duckei in habit, the flowers of S. Adolphii are shorter than those of the latter because of the difference in size of the corolla lobes. LACMELLEA speciosa Woodson, spec. nov. Arbores. Folia opposita breviter petiolata late oblonga acuta basi obtusa 14—18 cm. longa 5.5-8.0 cm. lata coriacea glabra; petiolis 7-8 mm. longis. Inflorescentia ut videtur terminalis sed versimiliter in axillis foliorum superiorum lateralis; pedunculo 5—7 mm. longo, bracteis ovatis minimis; floribus speciosis albidis subsessilibus. Corollae salverformis tubum ca. 3 cm. longum, faucibus paulo ampliatis ca. 5 mm. longis; lobis ca. 1.3 cm. longis 4 mm. latis. Calycis laciniae late subreniformes rotundatae ca. 2 mm. longae piloso-ciliatae. Antherae lineares 4 mm. longae. Pistillum 1 cm. longum in ovario gradatim ampliatum; stigmate ca. 2 mm. longo. Fructus ca. 3.5 cm. diam. lutei; seminibus 2.—COLOMBIA: DEL VALLE: Río Calima (region del Choco), La Trojita, 5-50 m. alt., Feb. 19—March 10, 1944, J. Cuatrecasas 16568, in Herb. Missouri Bot. Gard., HOLOTYPE. 1 Agrónomo Silvicultor, Forest Service, Brazil. Issued November 30, 1949. (543) [Vor. 36 544 ANNALS OF THE MISSOURI BOTANICAL GARDEN This species is particularly noteworthy because of the corolla lobes which are much longer than those of any other known species of the genus. Since the lobes are nearly thrice the length of the corolla throat, a redefinition of Markgraf's (Notizbl. 15:618—619. 1941) sections EULACMELLEA and ZscHOKKEA is required. Less satisfactory would be the erection of a third section to accommodate L. speciosa, merely on the basis of corolla proportions. МАГОЏЕТТА parvifolia Woodson, spec. nov. Arbores graciles vel fructices lactescentes ca. 4—5 m. alti; ramulis gracilibus cortice nigris. Folia opposita brevis- sime petiolata late ovato-elliptica 4.5—5.5 cm. longa 2-3 cm. lata late obtusa basi obtusa vel rotundata subcoriacea glabra subtus cum domatiis parvis; petiolis 0.2—0.3 cm. longa. Inflorescentiae axillares pauciflorae subsessiles. Flores parvi albi; pedi- cellis cum pedunculo petiolos subaequantibus. Corollae salverformes tubo cylindrico 5-gono ca. 0.4 cm. longo basi ca. 0.1 cm. diam. extus glabro, lobis oblique lanceo- latis acuminatis 0.8—0.9 cm. longis valde patulis intus papillatis. Calycis laciniae valde imbricatae late ovatae scariaceae ca. 0.2 cm. longae cum squamellis internis marginalibus solitariis. Antherae paene inclusae anguste lanceolato-sagittatae ca. 0.3 cm. longae dorso apicem versus sparse pilosulae. Ovaria са. 0.5 mm. longa puberulo-papillata cum nectariis 5 fere aequilongis; stigmate fusiformi ca. 1 mm. longo cum stylo aequilongo. Folliculi lineari-fusiformi 6—9 cm. longi ca. 0.3 cm. crassi glabri; seminibus 2.5 cm. longis valde canaliculatis sparse pilosis —VENEZU- ELA: AMAZONAS: aquatico, en las margenes abiertas y completamente inundadas cerca de la boca del Río Sanariapo, arriba de Raudal de Maipures, alt. 120 m., Feb. 7, 1942, L. Williams 15066, in Herb. Missouri Bot. Gard., HOLOTYPE. This species, like that immediately preceding, was received from the U. S. National Herbarium a few years ago in an exchange of unnamed duplicates. M. parvifolia apparently is most closely allied to M. cestroides, of the Robbia-complex, but differs from it quantitatively in almost every detail. Marourria grandiflora Woodson, spec. nov. Arbores lactescentes ca. 8 m. altae; trunco ca. 12 cm. diam., ramulis gracilibus cortice tenui fere nigro. Folia opposita subsessilia late elliptica acuminata basi acuta 20—25 cm. longa 7—9 cm. lata glabra subcoriacea subtus sine domatiis. Inflorescentia terminalis umbelliformis pauciflora, pedunculo ca. 0.5 cm. longo, bracteis scariaceis minimis. Flores speci- osissimi albi; pedicellis gracillimis са. 2.5 cm. longis; calycis laciniis aequalibus plus-minus imbricatis ovato-trigonalibus acutis 0.4 cm. longis scariaceis papillatis intus cum squamellis alternatis solitariis bifidis; corolla infundibuliformi extus glabra vel indistincte papillata, tubo proprio valde arcuato ca. 3.5 cm. longo basi ca. 0.3 cm. diam. apicem versus sensim attentuato, faucibus conicis ca. 1.5 cm. longis ostio ca. 1 cm. diam., limbi lobis oblique ovatis late acutis ca. 2 cm. longis basi 1.5 cm. latis paulo patulis; ovariis late oblongoideis ca. 0.2 cm. longis dense puberulo-papillatis cum nectariis 5 compressis ca. 0.7 cm. longis, stylo gracili ca. 1.2 cm. longo, stigmate oblongo-fusiformi ca. 0.2 cm. longi; antheris anguste lanceo- 1949] AZUMBUJA & WOODSON—NEW APOCYNACEAE 545 lato-sagittatis paene inclusis ca. 0.6 cm. longis dorso apicem versus sparse pilosulis. Fructus ignoti.—VENEZUELA: AMAZONAS: Maroa, Río Guainía, alt. 127 m., Feb. 10, 1942, L. Williams 14237, in Herb. Missouri Bot. Garden, HOLOTYPE. For a Malouetia, this species is almost alarming, with its relatively gigantic, in- fundibuliform corollas so alien to its cogeners. Surely it will be made the type of a distinct subgenus by the next monographer of the genus. І am persuaded to forbear the erection of a new genus because the affinities of M. grandiflora so obviously are with Malouetia, as seen іп the habit of the plants, the rather distinctive bark, the leaves (although our specimen is without the domatia found with more or less regularity in the other species), and the structure of the inflorescence and reproductive organs. НЕ А i ЌЕ. [Vor. 36, 194] ANNALS OF THE MISSOURI BOTANICAL GARDEN : EXPLANATION OF PLATE PLATE 40 Habit and floral dissections of Secondatia Adolpbii Azumbuja. ANN. Mo. Вот. Сакр., Vor. 36, 1949 PLATE 40 AZUMBUJA & WOODSON—NEW APOCYNACEAE [Vor. 3¢ 548 ANNALS OF THE MISSOURI BOTANICAI. GARDEN EXPLANATION OF PLATE PLATE 41 Fruit and seed of Secondatia Adolphii Azumbuja. dy 1949] ANN. Мо. Вот. Ganp., Vor. 36, 1949 Р1.АТЕ 41 bs Í c е е. | a pi АД с» d > 2 > ! ° | т“ > k 10 cm. : AZUMBUJA & WOODSON—NEW APOCYNACEAE GENERAL INDEX TO VOLUME XXXVI New scientific names of plants and the final members of new combinations are printed in bold-face type; synonyms and page italics; and all other matter іп ie ae type numbers having reference to figures and plates, in A Adenodiscus, 511; mexicanus, Allen, P. p Окс of Panam (third part), ш part), 1 а di some uses of Me a the Sierra 05 Anderson, Edgar: Charles M. Rick and. uses of maize in the Sierra of са. 405; Stoner, С. R., and. Maize among the Hill Бев of Ао, 355 Andrews, epee ә Jr. Nucellangium, a new genus of fossil = рн 25- signed to Lepi idocarpon, Aneurophyton, 291 iie ot South America, New, 543 Asia: maize in, 389; distribution of major races in, 391 Assam, Maize among the Hill Peoples of, Assam hills: qi in the, 375; history in the, 374; of, 357; tribes of, 356; varieties of d among the, 359 UE David de, and Rater N Apocynaceae E. Wood- 4 South rer 543 B Baxter, Robert W. Some Pteridosperm stems and fructifications with particular ref- erence to the Medullosae, 287 Beer prepared from maize: in Assam, 377, in Peru, 410 Bhutan, maize in, 372 Burma, maize in, 373 С Calamopitys, 291 California, Assamese varieties of maize grown in, 272 СаПејоп de Huailas, uses of maize іп the, ын ncha, preparation of, from maize, 409 чаи d plants, 287, 479 Cardiocarpus, 492; EE HI. 493; injens, 92; minor, 492 Chicha, preparation of, from maize, 410; chicha de jora, 410; chicha morada, 411 Chochaca, preparation of, from maize, 4 Chondrorhyncha aromatica, 85; calo жы 85; жалк 86; Чї scolor, 87; mar- £inata, 9 Chromosomes in Paphiopedilum: compara- B length of, in P. Lawrenceanum an P. «ЕГЕР ku". 443; numbers of, 445; terminal, 453, 4 Coal balls, foil; found in, 287, 479 Coix, uses of, by the Assamese е. 358 Cordaianthus spicatus, 493 Cusqueño pr kg Cypripedilum Cypripedium, 2. callosum, 434; H yeanum, 435; Lawrenceanum var. Hyeanum, 435 Cytology: I Paphiopedilum callosum, 447; of P audiae Hort., 433, 449; of P. B vu 448; analysis of root-tips, 440; of yeasts, 260 D Dafla tribe of Assam, maize culture by the, Darrah hd H., description of coal-ball fossil by, Dolerotheca, 315; formosa, 316, 344, 348; Reedana, 317; Schopfii, 317, 348, 350, 352; sclerotica, 315. 316, 346, 346, 352; villosa, 317 E Economics in the Assam hills, 375 2 Euspermopteris, 294 Ethnology in relation to maize, 355, 405 F Ferns, fossil, plants e the Hill Peoples of As- 3125: ром: seeds, stems, fructifications, 479; 8 G Galactose, fermentation of, 260, 265 Grewia terebintbinaceus, 513 (551) 552 H Heliocarpus, А revision of the genus, 507; generic relationships, 507; economic value of, 509; geographic distribution, 509; taxonomy, 511; vernacular names, 510 Heliocarpus, 511; atus, boliviensis, 532; сөзіегісетій, 523; Саеса- P ae, 530; bontelens sis, 526; cuspidatus, 30; diclinus, 532; Donnell- Smithii, 530; 521; pallidus, 515; Palmeri, 517; p andrus, 517, var. nodiflorus, 536; pop- ayanensis, 532; popayanensis var. gresdi- folius, 532, var. Schumanni, 532, var. trichopoda, 532; reticulatus, 513; Rosei, 532; rudis, 532; stipulatus, 532; subtri- lobus, 532; киынба, 513; tomen- tosus, 528; trichopodus, 532; velutinus, 2 Heterangium Grievii, 204; Kukuki, 294; minimum, Hybrids: of d дашы акы. 433; of бассһа- romyces Inheritance in yeast, 260 gai ^: gram d a variety of Assamese ба dosis found in coal-balls from, 287, 479 J ob's tears, use of, by the Assamese tribes, K Kaloxylon, 293 Б Lacmellea speciosa, 543 Lay, Ko др revision of the genus Helio- carpus L., NADA. on glabrum, 479, 491; lomaxi, 486 PIA New species of, from Panama, [Vor. 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN Lonchocarpus atropurpureus, 282; calcara- tus, 282; densiflorus, 284; glabrescens, ath ‘oliganthus, 281; и. 282; са. 5 М McQuade, Henry A. ШИНИ of Paphio- і 33 поп of, 405; history of, 405; іп hill areas outside Assam, 372; in the Orient, 355; morphological survey of, grown by the Nagas, 379; On some uses of, in the Sierra of Amas 405; races of, in pun "dag and Asia, 390; uses of, in Assam, 377, in Peru, 405; varieties ki 380, 383 Maize: "Caribbean", 385; "Drooping Waxy", 381, 386, 402; “Early Slender”, 383, 400; “Early Upright”, 384, 404; “Late Side- wise", 381, 385, pe "Late Upright", 4, 404. БТ cestroides, 544; grandiflora, 544; уїйога, 54 Manioc, vele и іп the Assam Hills, 374 Manipur State: maize in, 373; tribes of, 357 Marsilea, 4 Medicine, use of corn preparations in, 409 Medullosa, 301; endocentrica, 303; Leuckarti, 298; Noei, 298; primaeva, 298; Thompsonii, 298 Medullosae, Some Pteridosperm stems and m with particular reference tot 8 Melibiose, fermentation of, , 26 geen ian, Non-, чарна ај in baki 2 йс аный aphyllum, 289, 322, 324; partial reconstruction of, 382 Millet, use of, by the Assamese tribes, 358 Monba tribe of bees 358, 306; енй ing maize, 306; maize nacked for ripening, Mote, preparation of, к= maize, 411 Mundkur, Balaji mutations, ois so possible causes of Medie сы inheri- tance in Saccharomyces, 259 N Naga tribes of Assam, 357, 9, 308; corner of a “jhum” field, 308; varieties of maize 1949] INDEX grown by, 379, characteristics of, 384 73 segregations іп бассһаго- s, Evidence as УРШЫ causes of, 259 Nucellus, fossil, 480, 486 oes ngium, a new genus of fossil seeds eviously assigned to Lepidocarpon 479 Кыса glabrum, 4 О Olla canchera, a prehistoric vessel used to parch maize, , 409 55 of Panama (third part), 1, Index (part III), Pleonandrae of, 434; Rolfe's key to suborder Diandrae of, 434 р Paleobotany studies, 2 а: Е of, - ІШ, Fascicle 4 Pabhiopedilum Mae Hort., The cytology Paphiopedilum, baad albino forms of, 437 mbers in, 445; felon of P. Plo “447, of P. Lawrenceanum, 448, of P. Maudiae, 449; key to riu ed meiosis іп, 447; root-tips, 440; ‚ 43 Rui “Alma Gavaert”, 438; bella- tulum, var 37; callosum, 434, 437, 441, 447, 466, list of crosses in which sed as parent, 459, var. Sanderae, 433, 434, 437, 459; Charlesworthii var. album, 437; Curtisii, 437; "Emerald", 438; “Еп- chantress", 438; Holdenii, 438; ; insigne var. Sande 38, Sanderianum, h used as parent, 462, var. Hyeanum, 433, 435, 437; чудат in 433, 435, analysis of, 448, Hat var. ouse", 436, var. "D 436, var. magnificum, 436, 4 убиено, Vds E 437; Rosettii, ; "Warden" ; Behe ies B Peru, uses of maize in ин Dept. of, 405 Psilophyton princeps, Photographing Aa ж Selaginella, 414 Phragmopedilum, 4 Phragmopedium, 44 553 Phytolaccaceae, 475 Pollen grains: of Paphiopedilum callosum, 447, 470; of P. Lawrenceanum, 449, 470; of P. Maudiae, 450, 470 . 5 transfer of crops by, 2 Popcorns: in Assam, 379; Oriental, Em in Peru, 40 Poo Pm ш, іп the Assam Hills, 374 Pteridosperm stems and fructifications, Some, with par США diis to the Medul- losae, 287 Q Qualea, A first dope for the ee (Vochy- siaceae), in North America Qualea cymulosa, 285; rupicola, Ao R Rhetinangium, 290 Rice in the Assam Hills, Rick, Charles M., and = fa sad: On some uses of maize in the Sierra of An- , 405 Rogers, David J. Stegnosperma: A new species and a generic commentary, 475 S Saccharomyces: Evidence excluding muta- tions, polysomy and polyploidy as possible cau n i i i 1 ӛсһегу, Robert W.: А first genus Qualea (Vochysiaceae) from North Те (Panama), 285; Robert E. Wood- collaborators. Cu ih Part ІП, Fasc. 4, 1, Fasc. 5, 133 Schopfiastrum, 291 Secondatia Айрып, 543, 546, 549; Duckei, 5 43 Ыг ч 288; oes ai characters of m genera of, 29 Seeds, го ші. сед", “normal” 480, "prolif- Segregacions in yeast, Non-Mendelian, 259, 71 зімі, Soe of the ares in North rth of Mexico, 4 Ыы. "413; acanthonota, 419, 426; da, 423, 431; arenicola, 418, 425; "o ue. 422, 430; armata, 413, 414, 422, 431; asprella, 420, 421, 420; Bige- lovii, 417, 425; cinerascens, 419, 427; Ser. Circinatae, 423; Coryi, 418; 421, 429; Douglasii, 422, 431; eremophila, 422, 430; subg. Euselaginella, 415, 417; 554 floridana, 419, 426; funiformis, 419, 426; Напвепі, 421, 422, 430; lepidophylla, 423, 431; leucobryoides, 42 0, 428; mutica, 419, 427; mexicana, ; огерапа, 419, 426; Paridi, 422; pilifera, 415, 423, 431; Riddellii, , 425; rupestris, 413, 417, 420, 421, уме т 417, 418, 425; scopulorum, 420, 421, 429; id noides, 42 Standleyi, 420, 428; tortipila, 414, 417, 424; Underwoodii, 419, 427; Wallocei, Watsoni, 420, 427; E maize ни 3 т: relation between maize апа, 393; е Assamese tribes, 358 росупасеае оў, 543; Sporangium, fusil 480, 486 Spores of the genus Selaginella in North America es = Mexico, 413 oe species of Medullosa, 312, Ж Stegnosperma: А чий Кеса and а generic - commentary, 4 cubense, 476; balimi- folia, 477 ; НИЕ 475, 477; balimi- М” т, 476; scandens, 476; Watso onii, 47 Бы. genera of seed-ferns, 290 Stems and fructifications, Some Pterido- sperm, with particular reference to the Medullosae, 287 [Vor. 36, 1949] ANNALS OF THE MISSOURI BOTANICAL GARDEN ге 291 Stonor, С. К., and Edgar Anderson. Maize — the Hill Peoples of Assam, 355 Sutcliffia, 291 г Tercio pelo, 4 Tetrad mies " yeast hybrids, 260, 268 Tiestos, 4 OCOS, Nas UR of, from maize, 406 Trichilia scandens, 476 Spores o ү? the genus Selag- inella in North America north of Mexico, 413 V Valcárcel's illustration of Olla canchera, 409 Vitamin synthesizing abilities in veasts, seg- regations of, 271 Vochysiaceae, the genus Lonchocarpus of , 285 W Woodson, Robert E., Jr., David de Azumbuja. New Acci of South III, Fasc. 4, 1, Fas A Yeast hybrids, tetrad m of, 260 Yeast, see Saccharom Z Zea Mays—see Maize j STAFF OF THE MISSOURI BOTANICAL GARDEN Director GEORGE T. MOORE HERMANN VON SCHRENK, RoBERT ХУ, SCHERY, Pathologist Research Associate eg M. GRE Gustav А. L. MEHLQUIST, urator [aae ізн M the Herbarium Research Horticulturist CARROLL ХУ, DODGE, Коал М, Try Mycologist ени ибн Gannon of the EDGAR ANDERSON, ' GEORGE B. VAN SCHAACK, Genetici Honorary Curator of Grasses ROBERT E. WOODSON, Jr. P JULIAN А. STEYERMARK, > of the Herbarium ç "Honorary Research Associate Henry N. ANDREWS, NELL C. Hor Paleobotanist : Librarian cat "ditor of Publications GERALD ULRICI, Business Manager BOARD OF TRUSTEES OF THE MISSOURI BOTANICAL GARDEN President RICHARD J. Lock woop Vice-President DANIEL К, CATLIN Second бу шош EUGENE PETTUS L. RAY CARTER Јонм S. LEHMANN рорівү FRENCH GEORGE T. Moore CHCOCK А, WESSEL SHAPLEIGH ETHAN А. H, SHEPLEY i е EX-OFFICIO MEMBERS Автнов H. Сомрто Wir. GR LIAM P. UNER, Marra nd of тежа ЈА President of the Academy of rsity Science of St. Louis M. DARST, WILLIAM SCARLETT, е of the City of Bishop «de the Diocesé of t. Louis Missou НЕКВЕВТ C. WIN President of the Board of feted y* St. Louis GERALD UrnicL Secretary