Annals of the Missouri Botanical Garden Volume XLVIII 1961 Published quarterly at Galesburg, Illinois, by the Board of Trustees of the Missouri Botanical Garden, St. Louis, Mo. Entered as 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 Garden. 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. Beginning with Volume 45, 1958: Subscription Рпсе_______ $12.00 per volume Single Numbers... 3.00 each Contents of previous issues of the ANNALS OF THE Missour! BOTANICAL GARDEN are listed іл the Agricultural Index, published by the H. W. Wilson Company. TABLE OF CONTENTS Flora of Panama. Part IV, Fascicle 4 (Chenopodiaceae to Caryophyl- laceae) Robert E. Woodson, Jr., Robert W. Schery and Collaborators Morphology and Anatomy of the Saururaceae. I. Floral Anatomy and Embryology M. V. S. Raju Hepaticites devonicus, A New Fossil Liverwort Hep ka Devonian of New York cis M. Huebe Index of Orchid Names—1960 Robert L. Dressler Preliminary Studies in the Genus Stanhopea (Orchidaceae) C. H. Dodson and G. P. Frymire Preliminary Revision of the Genus Drymaria 2. James A. Duke Sanango: New Amazonian Genus of Loganiaceae- George S. Bunting and James A. Duke Two New Members of the Genus Scolecopteris ...... R. Bradley Ewart Portraits of Plants. A Limited Study of the "Icones" rna Rice Eisendrath A Synopsis of Poinsettia (Euphorbiaceae) Robert L. Dressler General Index to Volume XLVIII 1-106 107-124 125-132 133-136 137-172 173-268 269-274 275-289 291—327 329—341 343—349 MISSOURI BOTANICAL GARDEN STAFF Director Frits W. WENT EDGAR ANDERSON Joun D. YER, Curator of Useful Plants Research tee Henry N. ANDREWS, RAYMOND FREEBORG, ie i Research Associate HucH С. Cuter, Norton Н. ecg, weal Executive Director Morphologis CARROLL м6 "Юорсе, TRIFON VON ScHRE Е Ааййан Carator Sousa of CaLaw TH Потвом, |, omic Plants Taxonomist and arator о Owen J. SEXT — — Research Ecologist е" L. мету лед GroncE B. VAN SCHAACK, NNALS Curato и of ag asses and AMES A. DUKE J Assistant Curator. Коввкт E. Woopson, Jr. о of the Herbarium Curator of Herbarium BOARD OF TRUSTEES President Ковевт BROOKINGS SMITH Vice-President LEICESTER B. FAUST Second Liegen Henry B. PFLAG Howarp Е. BAER Henry Ниснсоск DUDLEY FRENCH Warren McK. bg cen EX-OFFICIO MEMBERS GEORGE L. Capica STRATF LEE Mor — 2.2. of the "cei of ада Ay the озған ‘of Science of St. Louis DANIEL ScHLAFL CARL TOLMA President, Board of Education Chancellor o "Washington of St. Lou University Каумомр К. Tucker, Mayor a av City of St. Louis C. RANLET LINCOLN, Secretary ee ee ам Volume ХЕУНЕ Number 1 Annals of the Missouri Botanical | a uu Journal. РА Scientific Contributions from the ‚ Garde and тубай Shaw School е Botany of. m BOTANICAL GARDEN appears four times May, Se - ptember, and Nov ember. Four FLORA OF PANAMA Vol XLVIII BY ROBERT E. WOODSON, Jr. AND ROBERT W. SCHERY AND COLLABORATORS PART IV Fascicle 4 CHENOPODIACEAE (J. A. Duke) AMARANTHACEAE (J. А. Duke) NYCTAGINACEAE BATIDACEAE PHYTOLACCACEAE (K. Raeder) AIZOACEAE (L. I. Nevling, Jr.) PORTULACACEAE (L. I. Nevling, Jr.) CARYOPHYLLACEAE (J. A. Duke) ANNALS OF THE Missouni BOTANICAL GARDEN FEBRUARY, 1961 FLORA OF PANAMA Part IV. Fascicle 4* CHENOPODIACEAE By JAMES A. DUKE Flowers perfect or unisexual, monoecious, polygamous or dioecious, sessile or shortly pedicellate, often bracteate and bibracteolate, the bracts mostly herbaceous. Perianth uniseriate, of (0-) 2-5 lobes, usually hypogynous, the lobes discrete or basally connate, usually greenish. Corolla absent. Stamens as many as or fewer than the sepals, the filaments mostly discrete, the anthers 2—4-locellate, usually introrse and dorso-medially attached. Ovary superior, unilocular, uniovulate, the ovule erect on a short funicle or pendulous from an elongate funicle; styles 1-3, the stigmata capitate or elongate or the stigmata 2-5 and elongate. Fruit a 1-seeded indehiscent or circumscissile utricle, the perianth often and the sepals occasionally adherent to the seed; seeds erect, horizontal or inverted, lenticular, cochleate, subglobose or ellipsoid; embryos circular or hippocrepiform and more or less encircling the endosperm or spirally coiled and nearly filling the seed. Herbs, rarely shrubs or small trees, with simple alternate or opposite exstipulate leaves, these often farinose or glandular. Flowers glomerulate in axillary or terminal spikes, racemes, panicles or cymes, or rarely solitary and axillary, or embedded in the strobiloid axis of the inflorescence. Embracing about a hundred genera and more than a thousand species, the chenopods are a rather cosmopolitan family containing many weeds and halophytes in addition to a few vegetables such as the beet, spinach and swiss chard, the first of which is sometimes grown around the Canal Zone. ee weedy species of Chenopodium occur in Panama, one probably native, the other two known only as weedy advents on San Jose Is 1. CHENOPODIUM Г. CHENOPODIUM L. Sp. РІ. 218. 1753. Morocarpus Adans. Fam. 2:261. 1763. Anserina Dumort. Fl. Belg. 21. 1827. Teloxys Moq. in Ann. Sci. Nat. 21:289. 1834. * Assisted by a grant from The National Science Foundation. Issued April 3, 1961. (1) (343) [ Vor. 48 2 ANNALS OF THE MISSOURI BOTANICAL GARDEN Agathophytum Мод. loc. cit. 21: a 1834. Roubieva Мод. loc. cit. 21:292. Oligandra Less. in Linnaea 9: a n non ч = 1832. Orthosporum Т. . Ness, Gen. Fl. Germ. "id 1: pl. 57. Oligantbera Endl. Gen. 1377. 1841. Oxybasis Kar. & Kir. in Bull. Soc. Nat. Mosc. 1841:738. 1841. Ortbospermum Opiz, Seznam 70. 1852 Vulvaria Bubani, Fl. Pyren. 1:174. 1897. Botrys Nieuwl. in Am. Midl. Nat. 3:274. 1914. Flowers perfect or rarely unisexual, sessile or subsessile, ebracteate. Sepals (3-) 5, hypogynous, free or basally united, herbaceous, subequal, often strongly 1-ribbed and cucullate. Stamens 5 or fewer, occasionally varying in number in different flowers of the same inflorescence, the flattened filaments free or basally connate, the anthers mostly suborbicular, introrse, dorso-medially attached. Ovary sub- globose, the stigmata 2 (—5), filiform or subulate, mostly sessile or subsessile. Fruit an indehiscent utricle, ovoid to subglobose, the pericarp membranaceous to carnose, free or adherent to the single seed; seeds mostly cochleate-lenticular, smooth to roughened, vertically or horizontally oriented. Annual or perennial often strong- scented herbs. Leaves alternate, entire to pinnatifid, the lowermost at least usually petiolate, frequently glandular or farinose. Inflorescences of terminal or axillary glomerules, the glomerules variously arranged. Consisting of about a hundred weedy species, Chenopodium is a rather cosmo- politan assemblage of meager economic importance. Various species are used in Central America and elsewhere as potherbs, condiments and vermifuges. Three easily differentiated species are presently known to occur in Panama, but several others occur in Central America north of Panama. a. Plants farinose, never with oil glands on the leaves and inflorescence; seeds Ses sante dull or lustrous, > 1,5 mm. broad, the circular embryo completely encircling the endosper b. Leaf bl rhombic, E А ees сені seeds "cx tuberculate, 1.2— = 5 mm. broad, the gins acute; perica acm to the seeds. ‚ C. MURALE sepals carinate; seeds bb. Leaf blades — to > oer entire; 1.2 . broad, the margins obtuse; pericarp lustrous, — free of the seeds....... 2. C. PRATERICOLA аа. зь — with resinous dots on the leaves and inflorescence; hori and vertical, heck 0.6—0 .8 mm. vam the hippo- preci din incompletely encircling the епдоврегт,...............-........ . C. AMBROSIOIDES 1. CHENOPODIUM MURALE L. Sp. Pl. 219. 1753. Atriplex muralis Crantz, Inst. 1:206. ~ ense Zuccagni, in Roem. Coll. 133. 1806. rri erue murale 8 pote Мод. Chenopod. Enum. 32. 1840. Chenopodium murale ү cart bagenense Мод. loc. cit. 32. 1840. (344) 1961] FLORA OF PANAMA (Chenopodiaceae) 3 А садағы trachiosperma Bubani, Fl. Pyren. 1:177. 18 enopodium murale spissidentatum Murr, in Mur boe Lap. 2:11. 1903. Erect or ascending annuals to as much as 6 dm. tall, the branches ascending, often deeply sulcate, occasionally farinose, especially toward the extremities. Leaves somewhat lustrous and occasionally bullate above, glabrous to rather densely farinose below, deltoid to rhomb-ovate, marginally irregularly and acutely dentate, apically acuminate to attenuate and often mucronate, basally acute to truncate, 2.5-8 cm. long, 2-5 cm. broad, the petioles mostly 3-70 mm. long, sometimes quite as long as the blades. Flowers irregularly disposed in glomerules, the glomerules sessile along the dichotomously branching rhachises, these axillary and terminal, distally farinose. Sepals 5, subequal, ovate, obtuse, cucullate, l-ribbed, slightly if at all carinate, farinose, basally connate, 1-1.5 mm. long; stamens 5, caducous, the filaments flattened, the anthers orbicular, exserted; ovary subglobose, the style quite short, the 2 (-3) stigmata ca. twice as long, spreading, irregularly glandular. ІЛгісіе partially enclosed by the sepals, the seed lenticular-cochleate, minutely tuberculate, dull reddish-brown, 1.2-1.5 mm. broad, horizontal, the pericarp adherent. Presumably a native of Europe, Asia and Africa, now widely adventive in America as far north as Canada; in Panama known only as a lawn weed in San Jose Island. PANAMA: main camp, San José Island, Perlas Archipelago, Gulf of Panama, ca. 55 mi. sse. of Balboa, Johnston 1206. 2. CHENOPODIUM PRATERICOLA Rydb. in Bull. Torr. Club 39:310. 1912. Chenopodium cen var. leptophylloides Murr. in Bull. Herb. Boiss. 24:994. 1904 Chenopodium leptophyllum var. leptophylloides Thellung & Aellen, in Fedde, Rep. Spec. Nu. 2614. 192, var. leptopbylloides Aellen, in Ostenia 100. 1933 Chenopodium pratericola ssp. pratericola var. leptophylloides Aellen, i in ће Mid]. Nat. 30: 64. 1943. Erect or ascending annuals to as much as 3 dm. high, the branches ascending, usually multisulcate, bicolored, bullate or farinose, especially distally. Leaves yellowish-green above, whitish and densely farinose below, narrowly oblong to lanceolate, entire or rarely subhastatulate, apically obtuse to acute and mucronu- late, basally acute to attenuate, 10-45 mm. long, 2-10 mm. broad, the petioles 2-10 mm. long. Flowers sessile in dense glomerules, the glomerules in panicled interrupted spikes, the rhachises densely farinose. Sepals 5, subequal, ovate, sub- acute, cucullate, strongly carinate, farinose, basally connate, ca. 1 mm. long; stamens 5, the flattened filaments included, the anthers orbicular; ovary subglobose, the 2 stigmata sessile or subsessile, spreading, irregularly glandular. Utricle com- pletely embraced by the sepals, the seed lenticular-cochleate, smooth, lustrous, reddish-brown, 1-1.2 mm. broad, horizontal, loosely embraced by the pericarp. Western United States to Mexico; Argentina; probably adventive on San Jose Island as it is in the eastern United States. (345) ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 110. Chenopodium pratericola (346) [Vor. 1961) FLORA OF PANAMA (Chenopodiaceae) 5 ANAMA: abandoned corral, = кәне Island, Perlas Archipelago, Gulf of Panama, ca. 55 mi. sse. of Balboa, Johnston 12 The occurrence of this species as a weed on San Jose Island would tend to sub- stantiate I. M. Johnston’s claim (in Sargentia 8:30. 1949) that the weeds were introduced from Texas in unclean hay. The two related taxa [C. pratericola ssp. pratericola var. thellungianum Aellen, with linear leaves, and C. pratericola ssp. desiccatum (A. Nelson) Aellen, a semi-prostrate form] were unknown in Texas at the time of Aellen’s monograph (in Fedde, Rep. Spec. Nov. 26:119. 1929). It must be noted that Aellen (in Am. Midl. Nat. 30:64. 1943) has reduced C. dessi- catum to a subspecies of C. pratericola, a very peculiar mishap since C. desiccatum antedates C. pratericola by 10 years and seems not to be homonymous. If they do indeed prove to be mere subspecies of the same species, several new combinations will be required, but I will leave these for the monographer. 3. CHENOPODIUM AMBROSIOIDES L. Sp. Pl. 219. 1753. Chenopodium anthelminticum L. Sp. Pl. 220. 1753. opodium : Chenopodium chilense Schrad. Ind. Sem. Hort. Gotting. 1832:2. 1832. non Pers. 1805. Ortbosporum ambrosioides Kostel. Allg. Med.-Pharm. Fl. 1433. 1835. m Ambrina antbelmintica Spach, loc. cit. 5:298. 1836. Roubieva antbelmintica H. & A. Bot. Beech. Voy. 387. 1840. Cbenopodium retusum Juss. ex CMT in DC. Prodr. 13?:75. 1849. Chenopodium obovatum Moq. sh cit. 132:73. 1849 27. ambrosioides ni қобы т» А. Gray, Manual 2:364. 1856. Ambrina тыс Phil. in Anal. Univ. Sant. 91:442. 1895. Chenopodium querciforme Murr, т Mag. Bot. Lap. 3:37. 19 Blitum ambrosioides G. Beck, in ааб. Іс. Fl. Germ. 24: iff. 1908. Botrys ambrosioides Nieuwl. in Am. Midl. Маг. 3; 272. 1914. Botrys anthelmintica Nieuwl. loc. cit. 3:275. Ch есте vagans Standl. іп М. Am. ЕІ. 21 ced 1916. ous other infraspecific sites; see Aellen (in Am. Midl. Nat. 30:51. 1943). Erect or ascending ill-scented perennials to as much as 15 dm. high, the branches ascending, usually multisulcate, bicolored, often lignescent, glabrous to tomentulose or villose about the inflorescence. Leaves yellowish green, puberulent, villosulous or glabrate, glandular, with amber-colored secretions, lanceolate to rhomb-elliptic, entire to coarsely and irregularly sinuate-dentate or sinuate- pinnatifid, apically acute, basally attenuate, 2-12.5 cm. long, 0.5—5.5 cm. broad. Flowers sessile, usually in dense glomerules, these contiguous or not and paniculately disposed, interspersed with reduced or rather large leaves, the rhachises glandular and villosulous to glabrate. Sepals (3-) 5, subequal, narrowly ovate, cucullate, very slightly carinate if at all, connate for about one third their length, (347) [Vor. 48 6 ANNALS OF THE MISSOURI BOTANICAL GARDEN са. 1 mm. long; stamens (3-) 5, the flattened filaments about as long as the sepals, the anthers orbicular, slightly exserted; ovary subglobose, the sessile or subsessile stigmata spreading, irregularly glandular. Utricle completely embraced by the sepals, the seed lenticular-cochleate, smooth, lustrous, reddish-brown, 0.6-0.8 mm. broad, horizontal and vertical, loosely embraced by the pericarp. A cosmopolitan weed, perhaps indigenous in Mexico and Central America. снікюоі: Alto Lino, 4200 ft., Bro. Maurice 882. PANAMA: Juan Diaz, Standley 39529. Polymorphically perplexing, this species, or some of its subspecies, varieties or forms, is quite cosmopolitan. Colloquial names applied to the wormseed in Central America are almost as numerous as the Latin names. In Panama only the name “paico” has been encountered. The plant is used medicinally as a vermifuge, a poultice and reputedly even as a soporific; it is also used as a culinary flavoring. AMARANTHACEAE By JAMES A. DUKE Flowers perfect or unisexual, monoecious, polygamous or dioecious, mostly pentamerous, usually bracteate and bibracteolate, sessile or short-pedicellate. Calyx uniseriate or biseriate, commonly hypogynous; sepals (2-) 5, discrete or partially connate, scariose, whitish or variously colored. Corolla absent. Stamens (2-) 5, rarely more, hypogynous or perigynous, the filaments discrete or flattened and united below into a tube, the tube often with filamentous pseudostaminodia inserted between the filaments; anthers 2- to 4-locellate, usually introrse and dorso-medially attached. Ovary superior, unilocular, uniovulate (multiovulate in Pleuropetalum, Celosia and allied genera); ovules campylotropous on mostly elongate flattened funicles; styles 1-8; stigmata capitate or filiform. Fruit a 1-seeded utricle or rarely a several-seeded capsule, often circumscissile; seeds usually cochleate-orbiculate, the embryo excentric in а mealy endosperm. Herbs, shrubs, small trees or clambering vines with alternate or opposite mostly entire exstipulate leaves. Inflorescences of solitary flowers, spikes, glomes, glomerules, or thyrses simply, racemosely or corymbosely disposed. This largely tropical and subtropical family, of some 50 genera and 500 species, is represented in Panama by eleven of the twelve genera thus far reported for Central America. The twelfth genus, represented in Central America by Froe- lichia interrupta, has not yet been reported between Guatemala and Colombia. Many of the Panama species are cultivants or anthropochorous weeds of disturbed habitats. Several species in several genera are employed as potherbs and species of Amaranthus are a source of grain. Species of Iresine, Celosia, Amaranthus and Gomphrena are planted as ornamentals in Central America and other places. (348) 1961] FLORA OF PANAMA (Amaranthaceae) a. Leaves alternate; fruit a enden capsule or а l-seeded utricle; stigmata 2-8; anthers 4-locellat · Fruit a mapaa ca а ; stamens united D "is nearly half their length; stigmata 2-8, usually exceeding the c c. 0; n han the sulcate аа inflorescence paniculate or corymbose; leaves usually ovat woodland plant . PLEUROPETALUM cc. Capsules included б hy the erect sepals; seeds less than 10, deccm —3, much shorter than the tere ete style; inflorescences of mostly i ed infl i th Я а. Abed: monoecious or amo-monoecious (in Panama spp.) ; ments discrete; seeds oe mp reddish brown, less than 1.5 oad; erect or ascending herbs dd. Flower prius filaments basally connate; len. eure black, 1.5—2.5 m. broad; clambering suffruticose herbs or shrubs........... 4, CHAMISSOA aa. Leaves opposite; ciel а I-seeded indehiscent udi ана 1-3; anthers 2- to 4-locellat е. reco of elongate spikes, the constituent flowers or glomerules deflexed in fruit; bracts and ning Ари ска: anthers 4-locellate; style 1, idi 2 capitate s f. Flowers not амер рег dens spines straight or slightly a te; seeds 0 m. broad, reddi sh brown; sepals glabrate or 3. AMARANTHUS 5. ACHYRANTHES ff. Flowers glomerulate, some ко and highly modified; spines uncinate; seeds 1-2.5 mm. broad, succineous; sepals pilose, with г 6. CYATHULA anicle rules or short congested spikes, the constituent flowers e десен deflexed in fruit; bracts = spinescent (Alternanthera spp.); а = 2-locellate: styles 1-3, the stigmata тане олан or capita ‚ Inflorescences of spikes in much branched panicles (in Panama); flowers perfect, polygamous or dioe often with an obvious tuft of hairs ing betw: the "bes с“ тіні эж? da — 1-3, if 1, bilabiates erect herb» shrubs, trees or pg, 58 h. геном perfect; stigma bilabiate; oute pals ch oader than the i inner, wich the basal hairs borne crate inside de outer sepals; clam suffruticose perennials....................... PFAFFIA hh. Flowers i or ee stigmata 2-3, fllifor orm to deltoid; als subequal, the basal bars ing mostly outside the sepals; erect =; r shrubs (in Pan $. Inflorescences of spikes or Sint is solitary or in trichoto- emp ©. fev branched corymbs; flowers perfect; tufts of intrafloral E t obvious to the naked eye; stigmata 1—2, if 1, capitate; Sarton or ec herbs (Alternanthera spp. may be shrubby се ѕ). i. с Баѕеѕ amplexicaul, with a tuft of short hairs around the ode; stigmata 2; pseudostaminodia absent, th filament e short or absent; flowers short-pedicellate within e bract 9. PHILOXERUS ii. » ues no t amplexicaul, thus not forming a cup with a tuft ss | the node; pseudostaminodia well-developed (in Panama), бе filament tube обе flowers sessile өсі the bra j- Same ES Seas Р Secrest но E with an obvious the filam ted b lacerate or dentate pseudo- vaminodia; 555 “shorter than the b, not conspicu- ously c e (in 10. ALTERNANTHERA ji- ipii = sa. peris sessile at the summit of the filament tube between DUM я nodia; bracteoles ualing or exceeding the sepals, cristat (349 қ 11. GOMPHRENA [Уох. 48 ANNALS OF THE MISSOURI BOTANICAL GARDEN «==: pa ЕЕ Е > и Е E 7 ===> = Fig. 111. 1961] FLORA OF PANAMA (Amaranthaceae) 9 1. PLEUROPETALUM Hook. f. PLEUROPETALUM Hook. f. in Proc. Linn. Soc. 1:278, and in Lond. Jour. Bot. 5: 108. 1846 Allochlamys Мод. in DC. Prodr. 137:463. Melanocarpum Hook. f. in Benth. & Hook. ox 3:24. 1880. Flowers perfect, subsessile to pedicellate, unibracteate and bibracteolate. Sepals а нь hypogynous, concave, subequal, many-ribbed. Petals absent. Stamens ypogynous, connate about half their length forming an exappendiculate пе shorter than the ovary; filaments flattened; anthers 4-locellate, introrse, medially attached. Ovary subglobose, 1-locular, multi-ovulate; ovules cam- pylotropous; funicles flattened or filiform, exarillate, with free central placenta- tion; style 1; stigmata mostly 3—6, at anthesis scarcely distinguishable, ultimately reflexed. Fruits at first baccate, later developing into circumscissile or irregularly dehiscent capsules; seeds cochleate-orbiculate, black, lustrous, reticulate. Glabrate sparingly branched erect suffruticose herbs or shrubs. Leaves alternate, entire, petiolate, lanceolate to rhombic-ovate, glabrous to minutely strigillose, deciduous. Inflorescences of terminal and axillary racemes, panicles or corymbs, with or without foliar leaves. Inhabiting moist forests of low elevations, this genus, of perhaps three species, ranges from Jalisco to Peru and the Galapagos Islands. The type species, P. darwinii „ endemic to the Galapagos Islands, was reluctantly assigned to the Portulacaceae by Bentham & Hooker (Gen. 1:157. 1862.). The two other species of this economically unimportant genus are both found in Panama. a. Inflorescence corymbose; sepals with 7-17 ribs; stigmata mostly 3-4; broad capsule irregularly dehiscent; seeds T3122 mum. М 1. P. sPRUCEI aa. Inflorescence racemose or paniculate; sepals I бый ribs; stigmata ostly 5-7; capsule ил си seeds 1-1.5 т . 2. P. PLEIOGYNUM 1. PLEUROPETALUM SPRUCEI (Hook. f.) Standl. in N. Am. Fl. 21:96. 1917. Melanocarpum sprucei Hook. f. in Benth. & Hook. Gen. 3:24. 1880 Pleuropetalum costaricense Hort. Kew, ex Hemsl. Biol. бен. Ат. noc 3:12. 1882. etalum tucurriquense Donn. Smith, in Bot. Gaz 387 16. еже calospermum Standl. in Jour. "Wash. ront Sci 13:368. 1923. Erect sparingly branched subglabrous suffruticose herbs or shrubs to 3 m. high. Leaves glabrous to minutely strigillose on the veins below, broadly lanceolate to thombic-ovate, often falcate, apically attenuate, basally acute to subrounded, 3-15 cm. long, 1-6 cm. broad; petioles 5-20 mm. long. Inflorescences of mostly terminal corymbose racemes, the rhachises usually glandular. Flowers perfect, on pedicels 1-10 mm. long; bracts and bracteoles subequal, ovate to orbicular, carinate, 0.5-1 mm. long; sepals 5, subequal, ovate, concave, glabrate or scurfy, rounded to subacute, ultimately spreading, 2-4 mm. long, 1-2 mm. broad, with 7-13 (-17) ribs; stamens 5-8; filaments 2-3 mm. long; filament tube 1-2 mm. long; anthers oblong; ovary globose; style 1, about 0.5 mm. long; stigmata mostly 3—4, longer than the style, ultimately reflexed. Fruit an irregularly dehiscent capsule exceeding (351) [Vor. 48 10 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN the sepals, 4-7 mm. long, 3-6 mm. broad; seeds cochleate-orbiculate, black, lustrous, reticulate, 1.5-2.2 mm. long, on filiform or flattened funicles becoming 1-3 mm. long. OCAS токо: vicinity of Chiriquí Lagoon, von Wedel 1028. cuirigui: Вајо Mono, Boquete ио А 4500 ft., Davidson 481; К. Chiriqui Viejo Valley, on island a River, V. Whi te 150; valley of the upper Rio Chiriqui Viejo, 1300-1900 m., White This woodland species ranges from Jalisco to Peru. The Jalisco specimen (Mexia 1800) is atypical in having a circumscissile capsule, in that respect resem- bling the Galapagos species P. darwinii Hook. f. Standley (in Jour. Wash. Acad. 13:368. 1923.) describes P. calospermum from Salvador, and ascribes to it larger seeds, sepals and capsules than occur in P. sprucei. The larger seed size is not always correlated with the larger sepals. Variations in sepal and capsule size are frequently great enough in single specimens to embrace the descriptions of both species, so they are here concluded to be conspecific. - 2. PLEUROPETALUM PLEIOGYNUM (О. Ktze.) Standl. in Jour. Wash. Acad. 13: 369. 1923 Celosia pleiogyna O. Ktze. Rev. Gen. Pleuropetalum standleyi Canin in ix ii Nov. 44:41. 1938. Erect sparingly branched subglabrous suffruticose herbs or shrubs to 3 m. high. Leaves glabrous to minutely strigillose on the veins below, lanceolate to rhombic- ovate, often falcate, apically attenuate, basally acute to subrounded, 3—15 cm. long, 1-5 cm. broad; petioles 5-25 mm. long. Inflorescences of mostly terminal racemes or panicles, the rhachises usually glandular. Flowers perfect, on pedicels 1-5 mm. long; bracts and bracteoles subequal, broadly ovate, carinate, 0.5-2 mm. long; sepals 5 (-6), subequal, ovate, concave, glabrate or scurfy, rounded to acute, with 19—27 ribs, 2.5—6 mm. long, 1—4 mm. broad; stamens 6—8; filaments 2-3 mm. long; filament tube 1-2 mm. long; anthers oblong; ovary globose to ellipsoid; styles 1, sulcate, about 0.5 mm. long; stigmata 5—6, rarely more or less, usually longer than the style, ultimately reflexed. Fruit a circumscissile capsule equaling or exceeding the sepals, red becoming black, 5-7 mm. long, 4-7 mm. broad; seeds cochleate-orbiculate, black, lustrous, reticulate, 1-1.2 (-1.5) mm. broad, on filiform or flattened funicles becoming 1-3 mm. long. L ZONE: vicinity of = зе Culebra Dodge & ‘Alen Seas tees і А ч 15 rm o. Durs and Cafion of R. Chagres, Dodge & Allen 17405; drowned forest of Quebrada Bonita and en s. n. teyerma i == цоо а. даван Қы Giles os y p рате ме 727. PANAMA: Cerro Campana, alt. 800 m., Allen 4024. This species is apparently confined to Costa Rica, the type locality, and Panama. уз occurs in rather deep woods, especially along streams in ravines and drowned orests. (352) 1961] FLORA OF PANAMA (Amaranthaceae ) 2. CELOSIA L. CzLosiA І. Sp. Pl. 205. 1753. a Adans. Fam. Pl. 2:269. 1763. not L. das ans. loc. cit. 269. 1763 Lestibudesia 'Thouars, Hist. Vég. Illes Afr. 53. Lophoxera Raf. Fl. Tell. 3:42. 1837. Gonufas Raf. Sylva Tell. 124. 1838. Flowers perfect, the uppermost occa- sionally sterile, subsessile or short pedicel- late, unibracteate and bibracteolate. Sepals 5, discrete, hypogynous, concave, variously colored, not strongly ribbed. Petals absent. Stamens 5, hypogynous, connate less than half their length forming an appendiculate or exappendiculate tube shorter than the ovary; filaments flattened; anthers 4- locellate, introrse, medially attached. Ovary subglobose to ellipsoid, 1-locular, multi-ovulate; ovules campylotropous; funicles flattened, exarillate, with free central placentation; styles 1; stigmata 2- 3. Fruit a 2- to many-seeded circumscissile capsule; seeds cochleate-orbiculate, dark reddish brown to black, reticulate. Gla- brate or pubescent herbaceous of sufíruti- cose perennials or annuals. Leaves alternate, entire, subsessile to petiolate, glabrous or pubescent, lanceolate to rhombic-elliptic, deciduous. Inflorescences of terminal an axillary panicles or spikes, with or without foliar leaves. A large genus of some fifty species, this seems to have reached its best development in subtropical regions of America, Africa and Asia. Only one species, the cultivated cockscomb, is so far reported from Panama. (353) NI y Fig. 112. Celosia argentea 11 (Мог. 48 12 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1. CELOSIA ARGENTEA L. Sp. РІ. 205. 1753. Celosia cristata L. loc. cit. 205. 1753. $ О < ч, $ К 8r! э "d ы © а ~ ~ -” N a N Nn w Celosia coccinea L. loc. cit. 297. 763. Celosia pyramidalis Burm. Fl. Ind. 65. 1768. Celosia marilandica Retz. Obs. Bot. 3:27. 1783. Celosia pallida Salisb. Prodr. 145. 1796 Celosi > : Amaranthus purpureus Nieuwl. in Amer. Midl. (s 3:279. 1914. Erect glabrous simple or much branched annuals to 1 m. high. Leaves glabrous, linear-lanceolate to rhombic or ovate, apically acuminate to acute, 3-12 cm. long, 0.5—6 cm. broad; petioles 1-30 mm. long. Inflorescence of simple (quite complex, often fasciated, in cultivated varieties) pedunculate cylindric spikes 2-20 cm. long, 1-2 cm. broad. Flowers perfect, the uppermost occasionally sterile, sessile; bracts and bracteoles subequal, ovate, mucronate, 2-7 mm. long; sepals 5, subequal, ovate, concave, mucronate, white to pinkish (variously colored in cultivated varieties), erect in fruit, 6-10 mm. long; stamens 5, 3-5 mm. long, the tube shorter than the free portions of the filaments; pseudostaminodia minute and deltoid or absent; anthers oblong; ovary ellipsoid; style 1, 3—6 mm. long, usually exceeding the sepals; stigmata 2 (-3), minute. Fruit a circumscissile capsule, shorter than the calyx, 3-4 mm. long; seeds (1-) 3-6 (-9), cochleate-orbiculate, dark reddish brown, about 1.5 mm. broad Cultivated in many regions, possibly natives of Africa, the cockscombs are reported by Standley (in Contr. U. $. Nat. Herb. 27:172. 1928) to be cultivated in gardens in Panama, where they are called abanico. The cultivants often escape and morphologically approach the natural variety. Backer (in Fl. Mal. 47:74. 1949.) notes that in some escapes, one branch may bear the mark of cultivation with another resembling the wild form. He informs us that the seeds are used by the Chinese for poultices and for adorning cakes, while the leaves furnish an inferior vegetable. In India, where it is used medicinally and as a vegetable, it often appears spontaneously in paddy and ragi fields. Among vernacular names reported from Mexico and Central America are moño, san josé, cresta de gallo, amor seco, mano de león, flor de mano, amaranto and boria. 3. AMARANTHUS L. AMARANTHUS L. Sp. РІ. 989. 1753. Bliton Adans. Fam. РІ. 2:506. 1763. Baj < 1763. Dimeiandra Raf. Neogen. 2. SEMI Schrad. Ind. Sem. Hort. G = 1835.; in Li Ў Dimeianthus Raf. Fl. Tell. 3:41. + "i in Linnaea 11: Litt.-ber. 89. 1837. rius Raf. loc. cit. 42. 1837, (354) 1961] FLORA OF PANAMA (Amaranthaceae) 13 Albersia Kunth, Fl. Berol. ed. 2. 144. 1838 Mengea Schauer, in Meyer, Nov. Acta Acad. Leop. 19: Supply. 1:405. 1843. 55-56 ag а in DC. Prodr. 132:262. 1849. Sarr cit. 262. 1849. Calan Bubani, HL. Pyren. 1:184. 1897. Flowers monoecious, dioecious or polygamous, sessile or subsessile, unibracteate and bibracteolate. Sepals 3—5, rarely 1, discrete, hypogynous, concave, occasionally basally clawed, equal or subequal, membranaceous, erect and persistent in fruit. Stamens 3—5, rarely 1, hypogynous, discrete; filaments filiform; anthers 4-locellate, introrse, medially attached, oblong. Ovary ovoid to lenticular, 1-locular, 1- ovulate, the ovule campylotropous on a short exarillate funicle, the placentation basal; styles 1-3; stigmata 2-3, usually exceeding the styles. Fruit a utricle, indehiscent or circumscissile; seeds cochleate-orbiculate, smooth to minutely verrucose. Leaves alternate, entire to minutely crenulate, long-petiolate. Erect or prostrate, glabrous to pubescent annual herbs. Inflorescences of terminal and/or axillary spicate or paniculate thyrses, in Panama monoecious or polygamous with female flowers basally, hermaphroditic flowers medially, and male flowers distally, the latter tending to have longer, narrower sepals. Widespread in both temperate and tropical regions throughout the world, Amaranthus consists of perhaps fifty species. Many of them are inhabitants of open disturbed areas where they may become rather annoying weeds cultivated for their edible seeds and leaves, and others, with brightly colored inflorescences, are planted as ornamentals. Sauer (Ann. Missouri Bot. Gard. 37: 561-632. 1950) reports on the detailed ethnological history of the grain-amaranths. Seven species have been reported in Panama, but one, adventive on San Jose Island, apparently no longer persists. а. Stamens a 1-3, the sepals mostly less than 1.5 mm. long; un rug зек terminal and axillary; sepals v ed pistillate flowers 5, the bracts about equaling them in length; sta . A. VIRIDIS bb. Inflorescences all axillary; sepals of the тише [o mostly 1-2, about half as long as the bracts; stamens mostly 2 2. A. CALIFORNICUS aa. Stamens and sepals 5, the sepals ue. more than 1.5 mm. long; utricle smooth or rugulose. с: а subglobose; оен circular in cross section; sepals red 3 purple; plants unarm . A. CRUENTUS сс. селек compress најма 1 2—3-lobulate in cross section; sepals g h; plants armed о d. Pa dcus анна ахШагу 5-7 =н cylindric е. Thyrses less than 6 mm. broad; bracts of ғ pistillate flowers mostly shorter than the ағар, some of the sepals spatulate........... ee. Thyrses more than 6 mm. broad; bracts of the pistillate flowers делі зе гімен than the sepals, giving the spikes a bristly appear- ance; of the bracts spatul 5 НУВ dd. Plants sark axillary аи globose. 6. А. SPINOSUS In addition to the aforementioned, Johnston (in Sargentia 8:124. 1949) adds that A. retroflexus L. was collected on San Jose Island in 1945 (Erlanson 188), but was unsuccessfully searched for in 1946. I have not seen the specimen. A > . DUBIUS (355) [Vor. 48 14 ANNALS OF THE MISSOURI BOTANICAL GARDEN species is separated from the closely allied species A. bybridus by the obtuse sepals of the pistillate flowers. 1. AMARANTHUS VIRIDIS L. Sp. Pl. 2:1405. 1763. Chenopodium caudatum Jacq. Coll. 2:325. 1788. Amaranthus gracilis Desf. Tabl. Bot. 43. 1804 Albersia gracilis Webb. & Berth. Phyt. Canar. 3:287. 1836. Euxolus caudatus Мод. in DC. Prodr. 13? :274. 1849. Euxolus caudatus B gracilis Moq. loc. cit. 13?:274. 1849. Euxolus caudatus y maximus Мод. loc. cit. 13?:274. 1849. Rather delicate glabrous annuals to as much as 1 m. high, the branches ascend- ing. Leaves glabrous; entire to minutely crenulate, deltoid- to rhombic-ovate, apically emarginate to rounded and mucronate, basally truncate to subacute, 1.5—6 cm. long, 1-4 cm. broad, the petioles 0.5—4 cm. long. Inflorescences of thyrses racemosely disposed, the terminal and axillary thyrses cylindric, mostly less than 7 mm. broad. Flowers polygamo-monoecious or monoecious; bracts and bracteoles subequal, lanceolate to ovate, ca. 1 mm. long; sepals 3, linear-oblong to obovate, rounded, mucronate, 1-1.5 mm. long, the midrib dark green, the margins scariose; stamens 3, discrete, 1-1.5 mm. long; ovary compressed-globose; style 1, minute, stigmata 3, longer than the style. Fruit an indehiscent, strongly rugose utricle, 1-1.5 mm. long, 1—1.5 broad; seeds cochleate-orbiculate, dark red to black, dull, minutely pebbled, 1-1.2 mm. broad. Presumably a native of the Old World tropics, this weedy species is adventive around Balboa (fide Standley, in Contr. U. 5. Nat. Herb. 27:173. 1928). It is also adventive in the eastern United States. The names bledo and calalú are applied to all the species of Panamanian Amaranthus, any of which may serve as potherbs. 2. AMARANTHUS CALIFORNICUS (Moq.) S. Wats. Bot. Calif. 2:42. 1880. Mengea си оне Мод. in = Prodr. 137:270. 1849. Amaranthus carneus Greene, in Pittonia 2:105. У Amaranthus boue ciet Uline & Bray, in Bot. Gaz. 19:318. 1894. Prostrate, radially spreading delicate annuals, the branches to as much as 5 dm. long, often tinged with anthocyanins. Leaves glabrous, bullate, entire, often white-margined, ovate to obovate and spatulate, apically emarginate to roun and mucronate, basally attenuate, 3-25 mm. long, 2-12 mm. broad, the petioles 3-25 mm. long. Flowers in axillary, often leafy, few-flowered clusters, with every node often bearing an inflorescence, these ultimately shorter than the petioles. Flowers monoecious; bracts and bracteoles lanceolate, apically aristate, 1.5-2.5 mm. long; sepals 3 and quite unequal, ог 1-2 in the pistillate flowers, linear to lanceolate, acute to obtuse, 0.4-1.0 mm. long; sepals of the staminate flowers 3, subequal; stamens 2, discrete, ca. 1 mm. long; ovary ellipsoid, the stigmata 3, ca. as long as the ovary at anthesis. Fruit a utricle, circumscissile about the middle, rugose 1-1.5 mm. long, 1-1.5 mm. broad; seeds cochleate-orbiculate, dark reddish- Ca lustrous, minutely pebbled, 0.8—1.1 mm. broad. (356) 1961] FLORA OF PANAMA (Amaranthaceae) 15 PANAMA: abandoned corral, San José Island, Perlas Archipelago, Gulf of Panama, са. 55 mi. sse. of Balboa, Johnston 1151 Although the cited specimen was labeled as A. albus L., the prostrate habit, few stamens, and reduced sepals in the pistillate flowers clearly separate it from that species. The small seeds and rugulose utricles also serve to separate it from A. blitoides $. Wats., a very closely related prostrate species. 3. AMARANTHUS CRUENTUS L. Syst. Veg. 10:1269. 1759. Amaranthus paniculatus L. Sp. РІ; 2: W^ 1763. Amaranthus flavus L. loc. cit. 2:1406. 1763. Amaranthus sanguineus L. loc. cit. 1407. 1763. Amaranthus parisiensis Schkuhr, Handb. 3:249. ee Amaranthus paniculatus а Mrivonce Moq. in DC. Prods 13?:257. 1849. a 18 Amarantbus bybridus paniculatus Ulin & Bray, à in ла Torr. hr 5:145. 1894. rantbus dussii Veg OA in Bull. Soc. Tosc. Ort. 3!:178. Galliaria patula Bubani, Fl. Pyren. 1: 187. 1 Amarantbus bybridus хы cruentus Thellung, Fl. Adv. Montp. 205. 1912. usually pubescent annuals to 2 m. high, simple or with ascending branches. Leaves glabrous above, pubescent or glabrate below, entire to minutely crenulate, ovate to rhombic, apically rounded to acute and mucronate, basally acute to cuneate, 3-15 cm. long, 1-6 cm. broad; petioles 1-8 cm. long. Inflores- cences of thyrses racemosely disposed, the terminal and axillary thyrses cylindric, often drooping, 5-15 mm. broad. Flowers polygamo-monoecious or monoecious; bracts and bracteoles subequal, lanceolate to ovate, 1.5-2.5 mm. long; sepals 5, oblong to ovate, rounded to mucronate apically, 1.5-2 mm. long, the midribs dark green, the margins scariose and tinted with anthocyanins; stamens 5, discrete, 1-2 mm. long; ovary compressed globose, capped by a circular stylopodium; stigmata 3 (—4), longer than the stylopodium. Fruit a smooth to rugulose subglobose utricle, circumscissile near the middle, about 2 mm. long, usually exceeding the sepals; seeds cochleate-orbiculate, dark reddish brown to black, minutely reticulate, 1-1.3 mm. broa This is the commonly cultivated grain amaranth of Central America. In Panama it is called abanico chino and calalí, and it is cultivated both as an ornamental and as a potherb. In Mexico and Guatemala the seeds are used in making sweetbreads and mushes. Standley & Steyermark (in Field Mus. Bot. 24*:153. 1946) point out that А. caudatus L., A. leucosperma Wats. and А. cruentus L. are best treated as one species "rather doubtfully distinct from A. bybridus." Sauer (Ann. Missouri Bot. Gard. 37:561-632. 1950) on the other hand contends that there are four major grain amaranth species in the Americas, A. leucocarpus Wats., chiefly in Mexico; A. cruentus L., chiefly in Guatemala; А. caudatus L., chiefly in the Andes; and A. edulis Spegazzini, chiefly in Argentina. Of the cultivated amaranths, whether they comprise several or one species, he cites only A. cruentus from Panama. (357) (Мог. 48 16 ANNALS OF THE MISSOURI BOTANICAL GARDEN 4. AMARANTHUS DUBIUS Mart. РІ. Hort. Erlang. 197. 1814. Amaranthus tristis Willd. Hist. Amaranth. 21. 1790. not А. tristis Amaranthus incomptus Willd. Enum. Hort. Berol. бірі: 64. 1813. byponymn. Amaranthus tristis 8 xanthostachys Moq. in DC. Prodr. 13?:260. 1849 Amaranthus tristis ^y flexuosus Moq. loc. cit. 260. 1849. Amaranthus tristis ô leptostacbys Мод. loc. cit. 260. 184 Amarant. iem ла us B xanthostachys Thellung, in Asch. x Graebn. Syn. Mittel-Eur. Fl. 5: ее y» ubius C flexuosus Thellung, loc. cit. 266. 1914. Amaranthus dubius D leptostachys 'Thellung, loc. cit. 266. 1914. Rather delicate mostly glabrous annuals to 1 m. high with ascending branches. Leaves glabrous or glabrate, entire or minutely crenulate, deltoid- to rhombic-ovate, apically rounded to acute, often mucronate, basally rounded to acute, 2-8 cm. long, 1-6 cm. broad; petioles 1-6 cm. long. Inflorescences of thyrses racemosely disposed, the terminal and axillary thyrses mostly cylindric and less than 6 mm. broad. Flowers polygamo-monoecious or monoecious; bracts and bracteoles sub- equal, lanceolate to ovate or obovate, acute to acuminate, 1.5-2 mm. long, the midribs dark green, the margins scariose; stamens discrete, 1-3 mm. long; ovary ovoid; styles (2-) 3, fimbrillate. Fruit a slightly rugose compressed ovoid utricle, circumscissile about the middle, 1.5-2.5 mm. long; seeds cochleate-orbiculate, reddish brown to black, reticulate, 0.8-1.1 mm. broad. AS DEL TORO: Isla Colón, vicinity of Chiriqui Lagoon, von Wedel 2854. CANAL near Gorgas Memorial Laboratory, vicinity of Miraflores, G. White 119; in govern- ment dons эче ng Las Cruces Trail, Hunter & aoe 712; Bohio Soldado, Cowell 235. DARIEN: Vi of B e Сире, ca. 40 m., Allen 874. РАМАМА: Agricultural Exp. Sta. at Matias унио Pittier 68 Reputedly the common weed amaranth of the Caribbean, this species extends from Mexico through tropical South America and is adventive in Europe. In Panama, where the leaves and young shoots are employed as potherbs, the names bledo and са are applied. The Mayas call it xetz and chactez; Mexicans call the amaranths quelite, a name derived from an Aztec word and used for any pot- herb. The specific epithet is strongly suggestive of its specific status. It is weakly separated from А. bybridus L., from which it is frequently keyed by the length of the sepals in proportion to the length of the utricles. In individuals of both species, however, the utricles vary from shorter to longer than the sepals. 5. AMARANTHUS HYBRIDUS L. Sp. РІ. 990. 1753. Amaranthus bypocondriacus L. Sp. Pl. 991. 1753. Amaranthus hecticus Willd. Hist. Amaranth. 25. 1790. i 790. Amaranti tus Willd. loc. cit. 28 Amaranthus chlorostachys Willd за "s 34 Amarantbus retroflexus var. dou. Man. ed. 5 5:412. we ari е а oir riacus obinson, in Rhodos 10:32. 1908. Amarantbus bybridus 2 chlorostachys G. Beck, in Reich in > à 08. Galliaria bybrida Nieuwl. in Am. Midl. Nat. 3: 4224 DH АА ad Ы (358) 17 FLORA OF PANAMA (Amaranthaceae) 19611 МА t М Ж MAY vé? WAyiz2 ~ ee. Ау; We W AV mS oy eA SSB wv VE Deh 5; + i at. LA “УАЙ L ) W \ AS E <%2 Se Tz n чу, за Sha ИК ey 4 We D NA УУ ДХ) ғ 7) ңе NAN 274 AX, ATE “4 ~ 7, S S EN Ж ny Amarenthus dubius (359) Fig. 113. [Vor. 48 18 ANNALS OF THE MISSOURI BOTANICAL GARDEN Rather coarse often pubescent annuals to 2.5 m. high with ascending branches. Leaves pubescent or glabrous, entire to minutely crenulate, deltoid- to rhombic- ovate, apically rounded to acute, often mucronate, basally acute to rounded, 2—15 cm. long, 1-7 cm. broad; petioles 1-8 cm. long. Inflorescences of congested thyrses racemosely disposed, the terminal and axillary thyrses mostly cylindric, 6-12 mm. broad. Flowers polygamo-monoecious or monoecious; bracts and bracteoles subequal, lanceolate to ovate, 2-4 mm. long, often conspicuously longer than the flowers; sepals lanceolate to ovate or obovate, acute to acuminate, 1.5-2.5 mm. long, the midribs dark green, the margins scariose, rarely tinged with anthocyanins; stamens 5, discrete, 1-3 mm. long; ovary ovoid; styles (2-) 3, conical; stigmata (2-) 3, fimbrillate. Fruit a slightly rugose compressed ovoid utricle, circum- scissile near the middle, 2–2.5 mm. long; seeds cochleate-orbiculate, reddish brown to black, minutely reticulate, 1-1.3 mm. broad. CHIRIQUÍ: vicinity of Boquete, 1200-1500 m., Woodson & Schery 722. This weedy species, frequently forming large stands in old fields, is found throughout temperate and tropical North and South America, and is adventive in many parts of the Old World. Vernacular names reported in Mexico and Central America аге quintoniles, calete, ses, buisquilete and quiec tes, bledo, xtez, quelite and buisquelite. The leaves and young shoots are often cooked as potherbs. 6. AMARANTHUS SPINOSUS L. Sp. Pl. 991. 1753. Amaranthus diacanthus Raf, Fl. Ludov. 31. 1817 marant hus caracasanus H.B.K. Nov. Gen. & Sp. 2:195. 1817. an rubricaulis Hassk. Flora 25; litt. 20. 1842. Amarantbus spinosus B purpurascens Moq. in DC. Prodr. 132:260. 1849. Amaranthus spinosus f. inermis Schum. & Laut. Fl, Deuts. Schutzg. Südsee 305. 1900 Бере бі йі. spinosus 1 circumscissus Thellung іп Asch. & Graebn. Syn. Mittel-Eur. Fl. 5: 9. 1914 Amarantbus spinosus II basiscissus Thellung, loc. cit. 269. 1914. Amaranthus spinosus Ш е сет; Thellung, loc. zr 269. 1914. Galliaria spinosa Nieuwl. i Midl. Nat. 3:278. Rather coarse, M m B^ armed Xen to 2 m. high, usually with erect branches. Leaves glabrous or pubescent below, narrowly rhombic-ovate, apically rounded to acute, basally acute to cuneate, 1-12 cm. long, 0.5-5 cm. broad; petioles 0.5-8 cm. long, with 2 thorns in their axils. Inflorescences of terminal and axillary thyrses racemosely disposed, the lower axillary thyrses globose and up to 15 mm. broad. Flowers polygamo-monoecious or monoecious, the bracts and мастона rather variable, lanceolate to acicular, 1-5 mm. long, often resembling thorns, some conspicuously longer than the flowers; sepals 5, lanceolate to oblong 1-2.5 mm. long, the midribs dark-green, the margins scariose; stamens crete, 1-2 mm. long; ovary ovoid; styles 3, conical; stigmata 3, fimbrillate. Fruit a slightly rugose compressed ovoid utricle, irregularly to regularly circum- scissile, 1.5—2.5 mm. long; seeds cochleate-orbiculate, reddish brown to black, minutely reticulate, 0.7—1.0 mm. broad (360) 1961] FLORA OF PANAMA (Amaranthaceae) 19 CANAL ZONE: Monkey Hill and vicinity, Cowe Il 19. cHrriguf: vicinity of Boquete, 1200-1500 m., Woodson & Schery 723; vicinity of Puerto Armuelles, 0-75 m., Woodson & Schery 8 33. COLON: и Cowell 2 214. HERRERA: vicinity of Ocu, 100 m., Allen 4086. Presumably a native of America, this anthropochorous species now has a circumpolar distribution in temperate and tropical countries. In Panama it is probably called bledo and сами. Other Mexican and Central American names are xtez, Кіх-хіет, buisquelite, bledo macho, ixtez, tsetz, labtzetz and nigua. In spite of the rather formidable thorns this species is also utilized as a potherb. 4. CHAMISSOA HBK. CuHamissoa НВК. Nov. Gen. & Sp. 2:196. 1817. Kokera Adans. Fam. Pl. 2:269. 1765. hyponym. Flowers perfect, subsessile, with 1—3 bracts subtending 1-5 bracteate fertile and 0—4 bracteate sterile flowers. Sepals 5, discrete, hypogynous, concave, greenish white, with conspicuous midribs and membranaceous margins. Stamens 5, hypogynous or perigynous, basally connate forming an exappendiculate tube shorter than, but occasionally basally fused with, the ovary; free portions of the filaments longer than the tube, usually somewhat flattened; anthers 4-locellate, introrse, medially attached. Ovary globose or ovoid, 1-locular, 1-ovulate; ovules campylotropous, the filiform arillate funicles with central basal placentae; styles 1, stigmata 2 (-3). Fruit a utricle circumscissile near the middle; seeds cochleate- orbiculate, black. Erect or clambering glabrate or pubescent herbs or shrubs. ves alternate, entire, petiolate or subsessile, linear-elliptic to broadly ovate, attenuate, deciduous. Inflorescences of axillary and terminal glomerules, these racemosely or paniculately disposed, with or without foliar leaves. This genus of perhaps five species, is chiefly a South and Central American genus with only one species ranging as far north as Mexico. The closely related oriental genus Allmania has occasionally been treated as a subgenus but morpho- logically seems to deserve generic status. Two species of Chamissoa occur in Panama. a. inflorescences = ee the glomerules paniculately disposed; oe less than 1.5 as long as the pesi ovary at anthesis tympaniform, нам d base of the style; sepals ovate, the -= more than 1 mm. broad; aril completely enveloping the seeds; punctate, 1. С. ALTISSIMA aa. Terminal inflorescences cylindric, the — racemosely disposed; styles 2 tim fig as the stigmata; ovary at anthesis doleiform, ла s to the base of the il iub tid the outer less than m. broad; aril minute; seeds verrucose 2. C. MAXIMILIANA 1. CHAMISSOA ALTISSIMA ( Jacq.) HBK. Nov. Gen. & Sp. 2:197. 1817. Celosia baniculata L. Sp. Pl. ed. 2:298. 1762. "e Pih anre altissima Jacq. Enum. PI. Carib. 1 soa macrocarpa НВК. Nov. Gen. & Sp. A i» таў: (361) [ Vor.48 ANNALS OF THE MISSOURI BOTANICAL GARDEN 20 Fig. 114. Chamissoa altissima (362) 1961] FLORA OF PANAMA (Amaranthaceae) 21 Celosia tomentosa Willd. in Roem. & Schult. Syst. Veg. 5: МЕ 1819. Achyranthes linkiana Roem & Schult. Syst. Уер. 5:545. 9 amis ti in DC. Prodr. 132:252 hamissoa altissima В laxiflora Мод. loc. cit. 251 D amissoa altissima y densiflora Moq = 251. Chamissoa altissima var. В glabrata Seub. in Mart. " 2 51:242. 1875 Kok e. Rev. Gen. 542 t Chamissoa altissima var. rubella Suesseng. in Rep. Spec. Nov. 35:306. 1934 Chamissoa altissima subsp. albo-grisea Suesseng. in Rep. Sp. Nov. 39:6. 1935. Chamissoa altissim a var. grandispicata Suesseng. loc. cit Chamissoa altissima var. grandispicata f. semispicata Suesse s. loc. cit. 6. 1935. Chamissoa altissima var. densi-paniculata Suesseng. in Lilloa 4 4:129. 1939. Clambering subglabrous shrubs or vines to 3 m. high. Leaves glabrous or slightly pilose, broadly lanceolate to ovate, apically acute to acuminate and mucronate, basally truncate to acute, 3-14 cm. long, 1—6 cm. broad; petioles 0.5— 3.5 cm. long. Inflorescences of glomerules paniculately disposed, the pubescent primary rhachis usually visible at intervals. Flowers perfect, some usually sterile, subsessile; bracts 1—3, broadly deltoid, carinate, mucronate, with membranaceous margins, 1—1.5 mm. long, 1-1.5 mm. broad; sepals 5, subequal, ovate, concave, acute to mucronate, 5-nerved, greenish white, 2.5—4 mm. long, 1-1.5 mm. broad; stamens 5; filaments 1.5—3 mm. long; filament tube 0.5-1 mm. long, often basally adnate to the ovary; anthers globose to oblong; ovary at anthesis tympaniform to globose, usually as broad as long and truncate or flanged at the summit; style 1, 0.5-1 mm. long, usually shorter than the stigmata; stigmata 2 (-3), erect at anthesis, reflexed in fruit. Fruit a utricle, exserted, circumscissile near the middle, compressed ovoid, the summit conspicuously operculate and truncate to incon- spicuously operculate and rounded, 3-5 mm. long, 2-3.5 mm. broad, aril bivalvate, becoming wrinkled, brownish white, completely investing the seeds; seeds smooth but punctate, black, lustrous, 2—2.5 mm. broad. BOCAS DEL TORO: Changuinola Meng Dunlop 309; vicinity of Chiriqui Lagoon, von Wedel 1380 9 1700; Water Valley, vicinity of Chiriqui 4... ер Wedel 634A, 840 8 1641; Old Bank Island, vicinity of Chiriquí Lagoon, von Wedel 1896 & 2141; Garay ved vicinity of Chiriqui Lagoon, von Wedel 2 37. CANAL ZONE: px Summit and G a, Greenman t$ oc Сс OCLÉ: E of El zu 600—1000 m., Allen 1205; El Valle de Antón, trail near Finca Tom rias, 600 Tucutí, Chepigana, Terry 8 Terry I ака жамай ead sod, on ка к less pe slopes, MacBride 2823. This most commonly collected species in the genus ranges from Mexico to Argentina, usually in thickets where it tends to clamber over adjacent plants. Of limited occurrence in amazonian South America is a rather distinct type often regarded as a separate species, C. macrocarpa HBK., with rounded exoperculate, frequently 3-stylate utricles much longer than the sepals. Kuntze (Rev. Gen 542. 1891), perhaps the first lumper, regards C. macrocarpa as conspecific with C. altissima. Annotation labels reveal that Schinz at one time considered C. “macrocarpa as only a variety of C. altissima, and finally Suessenguth (in Rep. Sp. Nov. 35:306. 1934) admits that a sharp distinction between the two is impossible. (363) [Vor. 48 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN 2. CHAMISSOA MAXIMILIANA Mart. ex Мод. in DC. Prodr. 137:251. 1849. Chamissoa celosioides Griseb. in Goett. Abh. 19:79. 1874 Chamissoa maximiliana var. В procumbens Seub. in Mart. Fl Bras. 51:243. 1875. Kokera acuminata (Mart.) O. пио loc. cit. 54 Chamissoa maximilian па маг . pubesc ns Chod. in xr Hi Boiss. 7:63. 1899. Chamissoa maximiliana Е. де (Griseb.) Suesseng. in Rep. Sp. Nov. 35:306. 1934. Clambering subglabrous herbs, shrubs or vines to 2 m. high. Leaves glabrous or slightly pilose, narrowly to broadly ovate, apically acuminate to mucronate, basally acute to rounded, 2.5—10 cm. long, 1-4 cm. broad; petioles 1-2.5 cm. long. Inflorescences of glomerules racemosely disposed, the pubescent primary rhachis usually obscured. (In the typical form the glomerules are not all closely approximated and the rhachis is visible at intervals.) Flowers perfect, some usually sterile, or tending to monoecism, subsessile; bracts 1-3, narrowly deltoid to lanceolate, carinate, mucronate, with membranaceous margins, 1.5-2 mm. long, 1-1.5 mm. broad; sepals 5, subequal, lance-oblong, concave, acuminate to mucro- nate, obscurely 3-7 nerved, greenish white, 3-4 mm. long, 0.5-1 mm. broad; stamens 5; filaments 1.5-3 mm. long; filament tube Бе 0.5 mm. long; ovary at anthesis doleiform, 1-2 times as long as broad, attenuate to the base of the style, often lobulate or flanged near the summit; style 1, 1-1.5 mm. long, longer than the stigmata; stigmata 2, erect at anthesis, reflexed in fruit. Fruit a utricle, usually included, circumscissile near the middle, globose to ovoid, the summit operculate and usually truncate, 2-3.5 mm. long, about 2 mm. broad; aril minute; seeds verrucose, black, often pleiochroistic, 1.5-2 mm. broad. BOCAS DEL ТОКО: Water Valley vicinity of Chiriqui Lagoon, von Wedel 1441; vicin- of Chiriqui Lagoon, von Wedel 1695. DARIÉN: vicinity of Cana, 1750 ft., Stern, Chambers et al. 677. Inhabiting thickets and forest clearings, this species ranges from Costa Rica to Argentina. The Panama specimens seem referable to the forma celosioides (Griseb.) Suesseng. which differs from the typical form in possessing compact inflorescences. The type of C. maximiliana has lax inflorescences with subglobose operculate utricles. Three other species with minute arils occur in South America. C. acuminata Mart., with smooth seeds and petiolate, narrowly ovate falcate leaves, occurs in Brazil and northern Argentina. С. blanchetii Moq., with smooth but punctate seeds and subsessile linear-elliptic leaves, occurs in Brazil. C. brasiliana (Moq.) К. E. Fr., native to Brazil and Argentina, differs from all other species in having axillary verticillate glomerules. 5. ACHYRANTHES L. ACHYRANTHES L. Sp. Pl. 20. 1753. Centrostachys Wall. in Roxb. a und 2:497. Cadelaria Raf. Fl. Tell. 3:39. и, Stachyarpagophora Vaill. i а Misa Yl Haban. 92. 1897. Flowers perfect, bracteate and bibracteolate, subsessile in elongate spikes. (364) 23 FLORA OF PANAMA (Amaranthaceae) 19611 Sis Achyranthes aspersa Fig. 115. (365) (Мог. 48 24 ANNALS OF THE MISSOURI BOTANICAL GARDEN Sepals 4—5, discrete, hypogynous, concave, subequal, obscurely 3—5 nerved, occa- sionally pungent. Stamens 2-5, the filaments flattened and united below forming an appendiculate tube; anthers 4-locellate, introrse, medially attached. Ovary obovoid, 1-locular, 1-ovulate; ovules campylotropous on elongate flattened funicles; style 1, filiform; stigma 1, capitate. Fruit a membranaceous turbinate indehiscent utricle; seeds cochleate-orbiculate, reddish brown. Glabrous to pubescent erect or decumbent annual or perennial herbs. Leaves opposite, short- petiolate, scantily appressed strigose to sericeous, deciduous. Inflorescences of terminal and axillary spikes, the flowers deflexed in age, the spinose tips of the bracts not uncinate. This weedy genus, of about five species, mostly of the Old World, is repre- sented by one species in Central America. The spines of the inflorescences are often a source of discomfort. Two varieties of one species occur in Panama. 1. ACHYRANTHES ASPERSA L. Sp. PI. 204. 1753. Achyranthes aspersa B indica L. loc. cit. 204. 1753. Achyranthes indica Mill. Gar. Dict. ed. 8: no. 2. 1768. Achyranthes argentea Lam. Encycl. 1:545. 1785. Achyranthes obtusifolia Lam. loc. cit. 545. 1785. Achyranthes sicula Roth, Cat. Bot. 1:39. 1797. Cadelaria sicula Raf. Е]. Tell. 3:39. 1837. Cadelaria indica Raf. loc. cit. 39. 1837. Stachyarpagophora aspersa Maza, Fl. Haban. 93. 1897. Achyranthes aspersa simplex Millsp. in Field. Mus. Bot. 2:36. 1900. Centrostachys indica (L.) Standl, in Jour. Wash. Acad. 5:75. 1915. Centrostachys aspersa (L.) Standl. loc. cit. 75. 1915. Erect or procumbent pubescent annuals or perennials, the usually branched stems to 2 m. long. Leaves scantily to densely pubescent, ovate to orbicular, apically acuminate to rounded, basally cuneate to rounded, 2-25 cm. long, 2-9 cm. broad; petioles 2~25 mm. long. Inflorescences of terminal and axillary pedunculate spikes, 3-40 cm. long, 6-12 mm. broad, the flowers deflexed. Flowers perfect; bracts and bracteoles subequal, 2-3.5 mm. long, ovate, spinescent, the tips rarely arcuate but not uncinate; sepals 4—5, greenish white, subequal, hypogy- nous, obscurely nerved, 4-7 mm. long; stamens 5, united below into a short tube; pseudostaminodia lacerate, exceeded by the 4-locellate anthers; ovary obovoid; style 1, filiform, at anthesis longer than the ovary; stigma 1, capitate. Fruit an indehiscent turbinate utricle; seeds cochleate-orbiculate, reddish brown, about 1 mm. broad. ZONE: Gatun, Cowell 300; near Frijoles, Sutton BOCAS DEL Toro: H. Wedel 418. CANAL т ier 4068. PROVINCE UNKNOWN: 3 DEL Pittier 6834; around Culebra, 50-150 m., Pitt Hayes 320. A rather ubiquitous pantropical weed, this species is perhaps adventive in America. In India the dried inflorescences are used by orthodox Hindus in sacred pyres. Standley & Steyermark (in Field Mus. Bot. 24*:145. 1946), who maintain A. indica as specifically distinct from A. aspersa, say that А. indica is scarce in Central America and known only from the Atlantic Coast, in contrast to the (366) 1961] FLORA OF PANAMA (Amaranthaceae) 25 widely distributed A. aspersa. The variety indica seems to be rather regularly distinct from the variety aspersa in having rounded leaves and shorter sepals. All the cited specimens except Hayes 320 seem referable to the variety indica. Many vernacular names are reported; zacpaiché and zorillo blanco in the Yucatan, abrojo in Salvador, mozote in Salvador and Honduras, mozotillo and rabo de chanco in Costa Rica, cola de armado, penegato, pije de gato and chile de perro in Guatemala where the typical variety is apparently distinguished by the natives from the variety indica to which the names pegapega, goincilla and mozotlexc are applied. 6. CYATHULA Lour. CxATHULA Lour. Fl. Cochinch. 1:101. 1790. Polyscalis Wall. Cat. no. 6939. 1832. Flowers perfect, some sterile and highly modified, bracteate and bibracteolate, subsessile in short-pedunculate glomerules. Sepals 5, discrete, hypogynous, concave, subequal, 1-3 ribbed, occasionally pungent. Stamens 5, the filaments flattened and united below into an appendiculate tube; anthers 4-locellate, introrse, medially attached. Ovary ovoid, 1-locular, 1-ovulate; ovules campylotropous on elongate flattened funicles; style 1, filiform; stigma 1, capitate. Fruit an ovoid indehiscent utricle; seeds cochleate-orbiculate, succineous. Pubescent branched ascending to prostrate occasionally frutescent herbs. Leaves opposite, entire, petiolate, strigose, deciduous. Inflorescences spicate, composed of terminal and axillary thyrses of complex glomerules containing some highly modified flowers with some of the bracts and sepals uncinate-spinescent. Of this rather weedy genus of some twenty species, most of the species are centered in Africa and the Orient. Only two occur in Central America, usually in disturbed habitats of low elevations, where they may become annoying weeds. . Terminal spikes 2-10 times as long as the uppermost leaves, interrupted over өте their аага lower oer with mostly 3 fer Bh ye an mm. broad; coals d mm. iss рибама по 3-toothe 1. C. PROSTRATA . Terminal spikes as long as the uppermost leaves, interrupted - than half nd = ү ты slomerules des 1 fertile | Sore and round 12 uncinate structures 1-4 m ong; seeds 1.5—2 . broad; codi 2.5—4 mm. long : леона ей а-аа 2. С. ACHYRANTHOIDES 1. CYATHULA PROSTRATA (L.) Blume, Bijdr. Ned. Ind. 549. 1826. Achyranthes prostrata L. Sp. Pl. ed. 2:296. 1762. — geniculata Lour. Fl. Cochinch. 1:102. vie ocbaeta prostrata DC. Cat. Hort. Monsp. 10 Pala prostrata Mart. Nova Acta Acad. Leop. Cael de 321. 1826. Erect or decumbent geniculate pubescent perennials to 1 m. high, often rooting at the lower nodes. Leaves strigose on both surfaces, entire or undulate, ovate to obovate, apically acute, basally acute to rounded, 2-6 cm. long, 1-4 cm. broad; petioles 2-8 mm. long. Inflorescences of terminal and axillary spikes of glomerules, the mature glomerule usually consisting of a central terminal perfect flower and 2 ы СЯ (367) [Vor. 48 26 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 116. Cyatbula prostrata lateral perfect flowers, each of the lateral flowers subtended on each side by a cluster of about 20 hooks representing 3 modified flowers, the hooks not conspicu- ously longer than the glomerule, the peduncle of the glomerule articulated near the pubescent primary rhachis; spikes 5- 30 cm. long, 4-7 mm. broad, with only flowers perfect; bracts and bracteoles subequal, ovate, mucronate or aristate, 1- 1.5 mm. long; sepals 5, subequal, ovate, concave, scariose, pubescent throughout their length, with 1-3 ribs strongly ex- serted dorsally, 2-3 mm. long; stamens 5, 1-2 mm. long, united below to form a tube bearing 5 intercalated 3-dentate pseudostaminodia; ovary at first tympani- form, becoming ovoid; style 1, stigma 1, at anthesis about equaling the stamens. Utricle ovoid, inconspicuously operculate, 1-2 mm. long; seeds cochleate-orbiculate, succineous, smooth 1-1.5 mm. long. BOCAS TORO: vicinity of Chiriqu La agoon, vom ‚ Wedel 1357; Water Vale, vicinty of 2 qui Lagoon, von I astimentos, ај yO Creek, itor of Chitiqui ы: von We 2808. Probably adventive in Ea Americas where it has a rather spotty distribution, this species is presumed to be native in tropical Asia and Africa. The glomerules are similar in complexity to the spikelets of some grasses, and they probably cling rather tenaciously to fur and clothing (368) 1961] FLORA OF PANAMA (Amaranthaceae) 27 2. CYATHULA ACHYRANTHOIDEs (HBK.) Мод. in DC. Prodr. 13::326. 1849. €— е НВК. Nov. ені : LS 2:210. 1817. Desmocbaeta densiflora HBK. loc. cit. 211. iua ыды bie sert R. & S. Syst hs 5 по 1819 Pupalia densiflora Mart. Nov. 1. 1826 : n.& Cyathula achyranthoides B Наса: Mod: in DC. Prodr. 13?:327. 1849. Cyantbula acbyrantboides ү densiflora M oq. loc. cit. 327. 1849. Achyranthes birtiflora А. Rich. іп Sagra, Hist. Cuba 11: b 1850. Cyatbula prostrata B acbyrantboides O. Ktze. Rev. Gen. 542. 1891. Erect or decumbent geniculate pubescent or glabrate annuals or perennials to 1 m. high, often rooting at the lower nodes. Leaves usually strigose on bot surfaces, entire or slightly undulate, ovate to rhombic-ovate, apically acuminate to attenuate, basally acute to cuneate, 4-15 cm. long, 1.5-5 cm. broad; petioles 2-10 mm. long. Inflorescences of terminal and axillary spikes of glomerules, the mature glomerule usually consisting of a fertile lower flower and a rudimentary, rarely perfect, terminal flower with 2 of its sepals uucinate-aristate, the terminal flower subtended on each side by a cluster of 3-7 hooks representing 1 modified flower, some of the hooks conspicuously longer than the glomerule; spikes 3-15 ст. long, 5-7 mm. broad, interrupted less than half their length; bracts and bracteoles subequal, ovate, concave, mucronate or aristate, 1.5-3 mm. long; sepals 5, subequal, ovate, concave, scariose, usually pubescent only at the extremities, with 1-3 ribs occasionally strongly exserted dorsally, 2.5-4 mm. long, 1-1.5 mm. broad; stamens 5, 1-2 mm. long, united below forming a tube bearing 5 inter- calated irregularly lacerate pseudostaminodia; ovary at first tympaniform, becoming ovoid; style 1, stigma 1, at anthesis about equaling the stamens. Utricle ovoid, rather conspicuously operculate, 1.5-2.5 mm. long; seeds cochleate-orbiculate, succineous, smooth, 1.5-2.5 mm. long. L TORO: vicinity of erect ca. 0-50 m., Woodson, Allen & Seibert 1806; Little B fece. vicinity of Chiriqui Lagoon, von Wedel 2525: Peach Creek, vicinity of ar a bars von Wedel 2650; Water "Valley, vicinity of Chiriqui Lagoon, von Wedel 610 CANAL ZONE: shrubby grassy ipe aes MacBride & Featherstone 2779; Balboa Heights Greenman & Greenman 5038. COLON: Juan Mina plantation, Rio Chagres, уы, 25 m., Allen 4110. basiti: trail ae Paya and Pucro, Stern, Chiba ји al. A; vicinity of Boca de Cupe, ca. 40 m., Allen 801. УЕКАСУА8: be- tween К. Tabasará aid Soná on ну“ W oodson, Seibert & Fera 504. Extending from Mexico and the West Indies to Brazil, this species is rather common in low-elevation disturbed habitats of tropical America. In Guatemala it is called cola de armado and in Honduras, mozote. The stouter hooks of the glomerules of this species may perhaps make them more painful and tenacious clingers than those of the preceeding species. 7. PFAFFIA Mart. PrarriA Mart. Nov. Gen. & Sp. 2:20. 1826. Serturnera Mart. loc. cit. 36. 1826. Hebanthe Mart. loc. cit. 42. 1826. (369) [Vor. 48 28 ANNALS OF THE MISSOURI BOTANICAL GARDEN Flowers normally perfect, bracteate and bibracteolate, sessile, often with an unpleasant odor. Sepals 5, discrete, hypogyneus, concave, subequal, the outer 3 slightly larger, often 3-ribbed, greenish white, becoming scariose, with a con- spicuous tuft of intertwined whitish hairs arising mostly within and longer than the outer sepals. Stamens 5, the filaments flattened and united below forming an entire or appendiculate tube; anthers 2-locellate, introrse, medially attached. Ovary ovoid to obovoid, 1-locular, 1-ovulate; ovules campylotropous on elongate flattened funicles; style 1, very short; stigma 1, bilabiate, exceeding the style. Fruit an indehiscent utricle; seeds cochleate-orbiculate, reddish brown. Subglabrous or copiously pubescent clambering vines or erect shrubs. Leaves opposite, entire to undulate, subsessile to short petiolate, glabrate to densely pubescent. Inflorescences paniculate with opposite branches or verticillate branches ultimately bearing spikes. (In some South American species, the inflorescence consists of condensed terminal heads.) Of this predominantly South American genus of some twenty species, two members of the clambering section HEBANTHE are found in Panama, one apparently reaching its northernmost station here. Stützer in a generic monograph (in Rep. Spec. Nov. Beih. 88:2. 1935) has noted that the section HEBANTHE is closer related to Iresine 6 TROMMsDORFFIA than to the other two sections of the genus Pfaffia. а. Мате sepals 2-3.5 mm. long, the ribs often not obvious to the е naked стары and lower leaf surfaces densely rufous-pubescent; stamen 1. Р. GRANDIFLORA ы >» È эбак зА 1.5—2.5 mm. long, the ribs obvious to the naked eye; rhachises and lower leaf surfaces slightly cinereous-pubescent; stamen tube vid 2 rounded teeth between adjacent 61атетіѕ............................ 2. P. PANICULATA 1. PFAFFIA GRANDIFLORA (Hook.) К. Е. Fr. in Ark. Bot. 16!2:10. 1921. Iresine grandiflora Hook. in Icon. Pl. 2: tab. 102. 1837. Gomphrena paniculata var. у bookeriana Seub. in Mart. Fl. Bras. 51:192. 1875. Hebanthe decipiens Hook. f. in Benth. & Hook. f. Gen. РІ. 3:41. 1880 Pfaffia bookeriana (Hems.) Greenm. in Field Mos. Bot: 2:330. 1912 Pfaffia grandiflora (Hook.) К. E. Fr. in Stützer, in Rep. е Nov. Већ. 88:9. 1935. Pfaffia grandiflora var. bookeriana Stützer, loc. cit. 10. Suffrutescent, usually pubescent, clambering eee to about 3 m. long. Leaves rufous-strigose, at least below, lanceolate to broadly ovate, apically acute to attenuate, basally rounded to acute, 2-10 cm. long, 1-5 cm. broad; petioles 2-10 mm. long. Inflorescence a terminal panicle of pedunculate, opposite or verticillate, simple or compound spikes, the lower often subtended by reduced leaves, the rhachises and peduncles densely rufous-pubescent. Flowers perfect, often infertile, sessile; bracts and bracteoles subequal, broadly ovate, rufous- tomentose, 1-2 mm. long, 1-2 mm. broad; sepals 5, subequal, elliptic, the ribs obscured by the pubescence, 2-3.5 mm. long, 1-2 mm. broad; stamens 5, 1.5-2.5 mm. long, united below into an exappendiculate tube about 0.5 mm. long; ovary (370) 19611 FLORA OF PANAMA (Amaranthaceae) 29 at anthesis ovoid; stigma 1, bilabiate, subsessile. Fruit an included indehiscent utricle to 2 mm. long; seeds cochleate-orbiculate, reddish brown, about 1.5 mm. long, on a funicle up to 1 mm. long. DARIEN: Cana and vicinity, 2000—6500 ft., Williams 828. This species ranges from Mexico to British Guiana and Peru. Standley (in Contr. U. S. Nat. Herb. 27:174. 1928) reports this species, as P. keriana (Hems.) Greenm., to have been collected at Gatuncillo. His identification is questionable however since he keys the species as having lobed stamen tubes. е reliability of pseudostaminodial characters is thrown open to suspicion by examina- tion of Williams 828 from Darién. In this specimen, possibly a hybrid between P. grandiflora and P. paniculata, although leaning strongly toward the former in other characters, the stamen tube in a single flower has pseudostaminodia between some filaments and none between others. Both Fries (in Ark. Bot. 16!?:10. 1921) and Standley (in Field Mus. Bot. 13:490. 1936) have pmi d associated the name P. grandiflora (Hook.) R. E. Fr. with P. brachiata Chod. (incl. P. bangii R. E. Fr.), a species differing by its striking dendroid pubescence. 2. PFAFFIA PANICULATA (Mart.) O. Ktze. Rev. Gen. 542. 1891. Iresine erianthos Poir. in Lam. 3: suppl. 180. Heban а Iresine paniculata Sine ve Veg. бс Post. 103. 1827. Gomphrena paniculata Мод. in DC. Prodr. 137:385. 1849. 5. 1891. Pfaffia eriantha (Poir.) О. Eo loc. cit. 543. 1891. Suffrutescent subglabrous clambering perennials to about 3 m. long. Leaves glabrous to appressed strigose below, lanceolate to ovate, apically acute, basally rounded to cuneate, 2-8 cm. long, 1—4 cm. broad; petioles 2-7 mm. long. Inflores- cence a terminal panicle of pedunculate, opposite or verticillate, simple or compound spikes, the lowermost often subtended by reduced leaves, the rhachises and peduncles slightly cinereous-pubescent. Flowers perfect, often infertile, sessile, bracts and bracteoles subequal, broadly ovate, translucent except for the carinae, 1-1.5 mm. long, 1-2 mm. broad; sepals 5, subequal, elliptic, the outer 3 con- spicuously 3-ribbed, 1.5—2.5 mm. long, 1—1.5 mm. broad; stamens 5, 1.5-2 mm. long, united below into a tube with 2 rounded lobes between adjacent filaments, the tube about 0.5 mm. long; ovary at anthesis obovoid, occasionally distally 4-lobate; stigma 1, bilabiate, subsessile. Fruit an included indehiscent utricle about 1 mm. long; submature seeds cochleate-orbiculate, reddish brown, about 0.5 mm. broad, on funicles about 1 mm. long. сосіЁ: El Valle, floor, 1800 ft., Allen 4475. This species, far from its reported range of Brazil to Peru and Paraguay, is possibly a recent introduction in Panama. The conspicuous cottony mass of hairs are certainly an aid to fruit dispersal. Three recent works on Pfaffia (Fries, in Ark. Bot. 1612:1, 1921; Suessenguth in Rep. Sp. Nov. 35:325. 1934; and Stützer, in (371) [Vor. 48 30 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 117. Pfaffia paniculata Rep. Sp. Nov. Beih. 88:1. 1935) all recognize that P. eriantha was described from a monstrosity. This monstrosity, which occurs in at least three species, has many or all of the flowers reduced to small sterile clusters of short stout hairs. However any flowers that may develop normally are adequate for identification. Moquin (in ОС. Prodr. 13*:386. 1849), who examined the type of P. eriantha, gives a description adequate to identify it with P. paniculata. International rules of nomenclature recommend that names based on monstrosities be rejected; therefore the name P. paniculata is upheld. (372) 19611 FLORA OF PANAMA (Amaranthaceae) 31 8. IRESINE P. Br. ТВЕЯМЕ Р. Br. Hist. Jam. 358. 1756. Trommsdorffia Mart. eH ~ & Sp. 2:40. 1826. ‚ 58. Xerandra Raf. Fl. Tell. 3 8. or 7. Ireneis Мод. in DC. Ргодг. 132:349. 1849. Е perfect, polygamous or dioecious, bracteate and bibracteolate, subsessile in spikes or glomes. Sepals 5, discrete, hypogynous, concave, subequal, becoming scariose, often with conspicuous tufts of hairs between the sepals and the bracteoles. Stamens 5, the filaments flattened and united below into an entire or appendiculate tube; anthers bilocellate, oblong, introrse, medially attached. Ovary 1-locular, 1- ovulate; ovule campylotropous on an elongate flattened funicle; stigmata 2-3, filiform or deltoid. Fruit an indehiscent utricle; seeds cochleate-orbiculate, whitish or reddish brown, smooth. Glabrous to copiously pubescent herbs, shrubs, trees or clambering vines. Leaves opposite, entire to denticulate, petiolate, glabrous to sericeous, herbaceous to coriaceous. Inflorescences of spikes or glomes, paniculately disposed, the branching alternate, opposite or verticillate. This heterogeneous genus, of some forty species, is largely if not wholly native to the Americas. I have seen no specimens, except of the cultivant Iresine herbstii Hook., from outside the Americas and the West Indies. Iresine herbstii is culti- vated in many places for its attractive variegated red and yellow foliage, but I have seen no specimens from Panama. The genus, and especially the polygamous and dioecious species, are sorely in need of revision. Four species are known to occur in Panama. a. Flowers perfect or polygamous; spikes ovoid or globose, pedunculate, alternate, opposite or verticillate on the rhachis; seeds whitish or reddish brown; style about half as long as the stigmata; herbaceous or suffruticose perennials, fe clamberin Leaves near the so sige Laien or ПРЕ their petioles amplexi- caul; silts mostly орров or ticillate; rhachises obviously cinereous-pubescent; реа Р cid, scarcely reflexed; seeds whitish. c. Leaves densely pubescent below; intrafloral bin a about ipie the calyx; pseu до машаны bilobate; РЕКУ 1- із long сс 1. I, HASSLERIANA сс; Leaves sparsely strigose below; intrafloral RE sos din as long Sar x yx; pseudostaminodia deltoid or absent; utricle 1.5-2 2. І. COMPLETA bb. им пеа Сай inflorescence linear or lanceolate, their petioles mostly not аай бакын se mostly alternate; rhachises glabrate or obscurely puberulent; stigmata filiform, ultimately reflexed; seeds reddish brown 3. 1. ANGUSTIFOLIA aa. Flowers strictly ANT spikes filiform or narrowly pyramidal, mostl нече Sunt on the rhachis; seeds reddish brown; style obsolete, les than rA ong as the filiform stigmata; herbaceous upright о ing ae . I. СЕКОЧА 1. IRESINE HASSLERIANA Chod. in Bull. Herb. Boiss. 23:390. 1903. Iresine macrophylla К. E. Fr. in Ark. Bot. 161?:41. 1921. Erect or clambering shrubs, the younger branches cinereous-pubescent. Leaves (373) [Vor. 48 32 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 118. Iresine hassleriana herbaceous to coriaceous, appressed pubescent below, glabrate above, narrowly elliptic to ovate, apically acute to attenuate and mucronate, basally rounded to acute, 2-15 cm. long, 1-8 cm. broad; petioles 3-15 mm. long. Inflorescence a panicle of opposite or verticillate pedunculate glomes, the rhachises cinereous- pubescent. Flowers perfect, the bracts and bracteoles subequal, orbicular, apically rounded, concave, entire, transparent, 1-1.5 mm. long, with a conspicuous tuft of hairs about equaling the sepals in length and arising between the bracts and the sepals; sepals 5, subequal, oblong-ovate, acute, obscurely 3—5 nerved, 1.5-2 mm. long; stamens 5, united below into a shallow tube bearing two rounded lobes tween adjacent filaments; ovary subglobose; style 1, shorter than the 2 deltoid stigmata. Fruit an indehiscent ovoid utricle, 1-1.3 mm. long; seeds whitish, cochleate-orbiculate, about 1 mm. long. PROVINCE UNKNOWN: Sutton Hayes 034. Apparently reaching its northern limits in Panama, this species is also found in (374) 1961] FLORA OF PANAMA (Amaranthaceae) 33 Peru, Bolivia, Brazil and Colombia. Specimens from the northern half of the range tend to have smaller less coriaceous leaves. Iresine domingensis Urban, described from Haiti, is a very closely related species, which Suessenguth (in Rep. Sp. Nov. 35:320. 1934) reports from Trinidad and Venezuela. 2. IRESINE COMPLETA Uline & Bray, in Bot. Gaz. 21:349. 1896. Erect or clambering shrubs, the younger branches cinereous-pubescent. Leaves herbaceous, glabrate or scantily strigose below, ovate, apically attenuate and mucronate, basally acute, 1.5-11 cm. long, 1-5.5 ст. broad; petioles 4-10 mm. long. Inflorescence a panicle of opposite or verticillate pedunculate glomes, the rhachises cinereous-pubescent. Flowers perfect (ог polygamous?); bracts and bracteoles subequal, broadly ovate, acute, concave, entire, transparent, about 1 mm. long, with a conspicuous tuft of septate hairs about twice as long as the sepals arising between the bracts and the sepals; sepals 5, subequal, oblong-ovate, acute, obscurely 3-nerved, 2-2.5 mm. long; stamens 5, united below into a shallow tube bearing obscure broadly deltoid pseudostaminodia between adjacent filaments; ovary subglobose; style 1, slightly shorter than the 2 deltoid stigmata. Fruit an indehiscent ovoid utricle, 1.5-2 mm. long; seeds whitish, cochleate-orbiculate, about 1.5 mm. long. DARIEN: above Paca and near Cana, Williams 706. Ranging from Panama to Guatemala, this species is nowhere commonly col- lected. The cited specimen, annotated by Standley, agrees with the original descriptions in all respects except that the pseudostaminodia are not prominent. Uline & Bray (in Bot. Gaz. 21:349. 1896) have emphasized that the type, Thieme 338, collected in 1888, is completely hermaphroditic. Donnell Smith (Enum. lant. Guat. 5:72. 1899) cites two of Thieme’s collections, omitting Thieme’s numbers, and using his own numbers, 5443 collected in 1889 and 5447 collected in 1890, as Iresine completa. Donnell Smith 5447 is however the staminate plant of a dioecious or polygamous species and has obvious pistillodes. It is obviously not the specimen used in the original description. Nevertheless Suessenguth (in Кер. Sp. Nov. 35:320. 1935) offers us his “Ergänzung zur Originaldiagnose" citing Donnell Smith 5447 as the type. This cannot be the holotype because the flowers are not perfect, they lack the basal tuft of hairs and they lack the orbicular utricles. It is possible, but presently strictly conjectural, that I. completa is polygamous with the pistillate flowers perfect and the staminate flowers bearing pistillodes. 3. IRESINE ANGUSTIFOLIA Euphr. Beskr. St. Barth. 165. 1795. Iresine elatior Rich. in Willd. Sp. Pl. 4:766. Iresine racemosa Poir. in Lam. ar Барра. B am. 1813. Rosea elatior Mart. Nov. Gen. & Sp. 2:59. Xerandra elatior Raf. Fl. Tell. 3:43. 1837. Alternanthera linearis Bello, in Anal. Soc. Esp. Hist. Nat. 12:107. 1883. (375) (Мог. 48 34 ANNALS OF THE MISSOURI BOTANICAL GARDEN Erect or reclining suffruticose perennials to 1 m. high, the younger branches glabrous or scantily strigose below, lanceolate to narrowly ovate, apically attenuate and mucronate, basally acute to attenuate, 5-10 cm. long, 0.5-3 ст. broad; petioles 0.5-2 cm. long. Inflorescence a panicle of alternate pedunculate ovoid spikes, these occasionally subtended by reduced linear leaves, the rhachises glabrate or puberulent. Flowers perfect, the bracts and bracteoles subequal, broadly ovate, mucronate, concave, entire, transparent, 0.5—1 mm. long, with a conspicuous mass of tawny hairs arising between the bracts and the sepals; sepals 5, subequal, oblong- ovate, acute 3-nerved, 1-1.5 mm. long; stamens 5, united below into a shallow tube with obsolete pseudostaminodia; ovary obovoid; style 1, slightly shorter than the 2 papillose filiform stigmata. Fruit an indehiscent utricle, 0.5—1 mm. long; seeds reddish brown, cochleate-orbiculate, 0.6-0.8 mm. broad. DARIEN: Rio Sabana, Leopold 131; Boca de Cupe, Williams 670. This lochmocolous species extends from Mexico through Central America to Brazil and Ecuador. Standley (in Contr. U. S. Nat. Herb. 27:174. 1928) reports that the name cadillo, a name applied to many plants with bur-like fruits, is used on Taboga Island. 4. IRESINE CELOSIA L. Syst. ed. 10:1291. 1759. Celosia paniculata L. Sp. Pl. 206. 1753 Iresine diffusa Humb. & Bonpl. in Willd. Sp. pl. 4:765. 1805. 7 805. Iresine bolym or bbs у verticillata Mart. loc. cit. 56. 1826. Xerandra celosioides Raf. Fl. Tell. 3:43. қ Iresine floribunda Mart. & Gal. in Bull. Acad. Brux. 101:347. 1843. Iresine celosioides var. eriobbylle Benth. Be Хоу. Sulph. 156. 1844. Iresine bookeri oq. in DC. Prodr. 13?:34 resine d. a in Iresine coim B pubescens Moq. loc. cit. 347. 1849. Iresine eriopbylla Moq. loc. cit. 347. 1849. Iresine gossypiantba А. Rich. in Sagra, ote Сұра 11:177. 1850. Iresine eriophora Peyr. in Linnaea 30:21. Alternanthera paniculata зае іп Апа]. Pe ЕЗ Зеро Nat. 12: 106. 1883. Iresine paniculata var. floridana Uline & Bray, in Bot. Gaz. 21:353. 1896. Erect or clambering herbaceous annuals or perennials, to 3 m. high, the younger branches glabrous or pubescent. Leaves herbaceous, glabrate to densely pubescent, ovate, apically acute to attenuate and mucronate, basally rounded to cuneate, 4— 15 cm. long, 1-7 cm. broad; petioles 0.5—6 cm. long. Inflorescence a panicle of subsessile filiform to pyramidal spikes, the rhachises glabrate to densely pubescent. (376) 19611 FLORA OF PANAMA (Amaranthaceae) 35 Flowers dioecious; bracts and bracteoles subequal, broadly ovate, mucronate, concave, often auriculate, transparent, 0.5-1 mm. long, with a conspicuous mass of hairs longer than the sepals arising between the bracts and the sepals of the female flowers; stamens 5, united below into an entire or minutely denticulate tube; ovary obovoid; style minute, much shorter than the 2 (-3) filiform stigmata. Fruit an indehiscent utricle 0.5-1 mm. long; seeds reddish brown, cochleate- orbiculate, 0.5—0.6 mm. broad. BOCAS DEL TORO: Water ie vicinity of Chiriqui Lagoon, von Wedel 1762 : ен CANAL ZONE: Colon to Empire, Panama R nep. ита 453; between Mt. Santa Rita trail, Cowell 81; с Isthmus of Рап nama, Fendler 260; between ies and Gamboa, Greenman t$ Greenman 5235. быға Росгего Muleto to Summit, Vol- can de Chiriqui, 3500-4000 m., Woodson & Schery 466; Rio Chiriqui to Remedios, 15-50 m., Woodson, Allen & Seibert 1102; Finca Lérida to Boquete, са. 1300-1700 m., Wood- son, Allen 8 Seibert 1131; Bajo Mono, mouth of Quebrada Chiquero, along Río Caldera, 155-2000 m., Woodson, Allen 8 Seibert 1012; Rio ME Viejo Valley between El Volcán and Cerro Punta, G. W bite 21; rain ем т, Bajo Chorro, Boquete, 6000 ft., Davidson 320; Volcan de Chiriqui, Boquete Distr., 6 ft., Пени 032; vicinity of New Switzerland, central valley of Rio Chiriqui Vido, ace m., Allen 1411. СОСТЕ: no specified locality, Macbride 2717. PANAMA: Taboga Island, up to 300 m., Allen 117; near mouth of R. Chagres, Allen 875; hills between Capire and Potrero, Dodge & Hunter 8645. PROVINCE UNKNOWN: Sutton Hayes 722, 928 8 020. This species is rather common in the warmer regions of North, Central and South America. Standley & Steyermark (in Field Mus. Bot. 24*:170. 1946) report that the sap is employed as a remedy for erysipelas around Cobán in Guatemala. Maya names for this plant are zactezxiu and zacxiu. Other vernacular names reported are pie de paloma, velo de princesa, adorno de nino, chancanil, tabudo, mosquito, hierba de gato, siete pellejos, coyontura, coyontura de pollo, taba de güegüecbo and camarón. Iresine spiculigera Seub. and Iresine acicularis Standl. were first lumped by Suessenguth (in Rep. Sp. Nov. 35:319. 1934), but later separated again (іп Rep. Sp. Nov. 39:13. 1935). Standley (т Field Mus. Bot. 132:517. 1937) apparently accepts the lumping and further states that 1. spiculigera "is doubtfully distinct from I. celosiz". Analysis of I. orgasm with cinereous spicular pubescence, I. acicularis, with ochraceous spicular pubescence, and I. frutescens Мод., with small flowers, reveals that they are best а conspecific with I. celosia L. 9. PHILOXERUS R. Br. Рнпохево$ К. Br. Prodr. 416. 1810. Caraxeron Vaill. in Raf. Fl. Tell. 3: = 1837. Blutaparon Raf. New Fl. 4:45. 18 Flowers perfect, bracteate im bibracteolate, subsessile in spikes. Sepals 5, discrete or basally slightly connate, hypogynous, the outer 3 flat and broader, the inner 2 conduplicate and narrower. Stamens 5, the filaments flattened and unite below to form a very shallow tube; pseudostaminodia absent; anthers 2-locellate, introrse, medially attached. Ovary ovoid to orbiculate, 1-locular, 1-ovulate; (377) (Мог. 48 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN ovules on elongate flattened funicles; stigmata 2, on 1 short style, often erect at anthesis and recurving in fruit. Fruit an orbiculate indehiscent utricle; seeds cochleate-orbiculate, reddish brown. Prostrate or decumbent glabrous to pubes- cent herbaceous perennials. Leaves opposite, entire, often succulent, the sessile bases usually amplexicaul. Inflorescences of terminal and axillary pedunculate congested spikes or glomes often subtended by slightly reduced foliar leaves. is small genus of two or three species seems to be largely restricted to beaches and salt flats in tropical America and west Africa. Schinz (in Engler & Prantl, Pflanzenf. 16c:83. 1934) has suggested that many Australian species formerly referred to this genus perhaps belong to Gomphrena. Only one species reaches North America. 1. PHILOXERUS VERMICULARIS (L.) R. Br. Prodr. 416. 1810. ompbrena vermicularis L. Sp. Pl. 224. 1753. peer vermiculatum L. Sp. Pl. ed. 2:300. 1762. E phrena aggregata Willd. Enum. 29 Philoxerus aggregatus HBK. Nov. Gen. & E 2:203. 1817. Philoxerus crassifoli BK. loc. cit. 203. Ac vermicularis Ell. Bot. .& Са. h 310. 1821. Gomphrena crassifolia Spreng. Syst. 1:824. 18 25. Caraxeron vermicularis Raf. Fl. Tell. 3: = 1837. Blutaparon brevifolium Raf. New. Fl. 4:45. 1838. Blutaparon repens Raf. loc. cit. 46. 1 Iresine aggregata Мод. loc. cit. 340. ы. Cruzeta crassifolia Maza, Fl. Haban. 94. Lithophila vermiculata Uline, in Field Mus. dm 2:39. 1900. Prostrate or decumbent perennials, the much branched stems glabrous except at the nodes. Leaves glabrous, succulent, linear to narrowly oblanceolate, apically acute to rounded, often mucronate, basally cuneate, 1-5.5 cm. long, 2-10 mm. broad, the leaf bases encircling the swollen nodes, with an axillary tuft of hairs. Inflorescences of terminal and axillary pedunculate bracteate white to stramineous spikes or glomes, 5-30 mm. long, 5-10 mm. broad, solitary or aggregated, the us. Flowers perfect; bracts and bracteoles subequal, ovate, acute to acuminate, concave, translucent, 2—3 mm. long, slightly shorter than or equaling the sepals; sepals 5, the outer 3 almost flat, usually glabrous, apically rounded, 3—4 mm. long, 1-1.5 mm. broad, the inner 2 conduplicate, pubescent, apically acute, 3-4 mm. long, 1 mm. broad; stamens 5, 1-2.5 mm. long, united below into a very shallow or obsolete tube; pseudostaminodia absent; ovary orbiculate to elliptic, style 1, about as long as the 2 papillose stigmata. Fruit an indehiscent utricle, about 2 mm. long; seeds cochleate-orbiculate, reddish brown, 0.8-1.2 mm. broad. BOCAS DEL ТОКО: Old Bank Island, vicinity of Chiriqui Lagoon, von Wedel 1948. A rather characteristic sea beach plant, this species ranges from Florida and Texas to Mexico and British Honduras, and from Panama to Brazil in the Amer- icas; it also occurs on the west coast of Africa. As the name implies, the plant (378) 1961) (A "4ranthacea ) 37 FLORA OF PANAMA Fig. 119, Philoxerys vermicularis (379) [Vor. 48 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN reputedly possesses vermifuge qualities. Р. portulacoides St.-Hil. differing in having glabrous white sepals, and P. littoralis Suesseng., differing in having glabrous white inner sepals and pilose foliar midribs are weakly defined species occurring in South America. 10. ALTERNANTHERA Forskal ALTERNANTHERA Forskal, Fl. Aeg.-Arab. 28. 1775. Flowers perfect, bracteate and bibracteolate, sessile or subsessile in spikes or glomes. Sepals 5, discrete, hypogynous, concave, subequal or strongly dissimilar, white to purple, the bases often indurate. Stamens 3—5, the filaments flattened and united below into an entire or appendiculate tube; anthers 2-locellate, oblong, introrse, medially attached. Ovary globose to obovoid, often compressed, 1-locular, l-ovulate; ovule campylotropous on an elongate funicle; stigma 1, capitate or obscurely bilabiate, on a short or elongate style. Fruit a membranaceous utricle, indehiscent; seeds cochleate-orbicular, smooth. Glabrous to copiously pubescent erect, scandent or prostrate annuals or perennials. Leaves opposite, sessile or petiolate, entire to denticulate, glabrous to densely pubescent, mostly herbaceous. Inflorescences of axillary and terminal sessile or pedunculate spikes or glomes. This genus of over 100 species is best developed in the Americas, although many species are ubiquitous weeds. Some species are cultivated for their ornamental inflorescences and others are planted as tufted border plants and soil binders. Alternanthera axillaris Hornem. and A. repens (L.) Gmel. are rather obnoxious weeds with pungent sepals. Working under the American Code of nomenclature, Standley (in Jour. Wash. Acad. 5:72. 1915) explains the rather complicated reasons which —led him to transfer the Alternanthera species to Achyranthes, a treatment which he later : abandoned. Hitchcock & Green (in Brittonia 6:114. 1947) have proposed that Alternanthera “repens (L.) Gmel. be the lectotype of the genus. Eleven species are reported from Panama. a. — sessile in the axils of small а s (mostly 1-4 cm. long); г and inner г sepals vn. pd ~~ the .5—2 times as broad а г (вера ч. subequal but exceeded by eg а in A. sessilis) ; сени or decumbent e b. Utricle lon: oe the sepals; ы subequal acute, 1-пегуей glabrous; ај of simple hairs bb. hence shorter than the sepals; sepals dimorphic or trimorphic, minate to ст 3-5 nerved, pubescent; pubescence of stellate 1. A. SESSILIS or ұзарта e келіні ibis e => pseudostaminodia; stigmata as long as or ger than the styles, attaining or exceeding the anthers; sepals ен ог анн prostrate sans with the leaves often less = 1.5 cm. long. d. Inflorescences subglobose to cylindric; sepals dimorphic, the subequal with flexible awns if any, whitish; bracteoles 0. = as long as бе sepals; stigmata longer than the styles; seeds rin 2 mm. broad. . Inflorescences mostly cylindric; sepals trimorphic, with 2 of the outer 3 aristate with rigid awns, stramineous, the third sepal 0.5—1 mm. shorter. , merely ac uminate; bracteoles more than half as long as cis sepas; stigmata about м long as the styles; seeds mm. 1-1.5 (380) 2. A. POLYGONOIDES с. с. 3. А. REPENS 1961] FLORA OF PANAMA (Amaranthaceae) 39 cc. Filaments shorter than the pseudostaminodia; stigmata shorter than the => exceeded by the anthers; sepals = orphic; prostrate ог decumbent herbs, the leaves often over 1.5 ong. е. Leaves apically acute or acuminate, nicks or sparsely strigose 4. A. FICOIDEA ee. ий apically rounded, densely cinereous-pubescent below with stellate hairs 5. A. HALIMIFOLIA aa. At least some of ve inflorescences pedunculate in the axils of larger leaves (mostly 4— m. long); outer ж. inner sepals subequal; erect or clambering rhe Se or decumbent aqua E Sepals di mm. "e 3-4 t as € s the bracts; glomes white, 10-18 . broad; pedunc узы “unbranched, mostly а than 5 ст. long; subglabrous Pesta aquatic pla 6. A. PHILOXEROIDES ff. Sep als 2-6 . long, ore than ise м long Ж iei bracts; ықы wile to тл 4—12 тт. 4; pedun branched or unbranched, 1-9 cm ‚ long; ica to сулый wane. suffruti- h. Peduncles mostly simple and exceeded by the subtending leaves; leaves ovate, short-strigose or glabrate; ek pe мець the hairs mostly less than 2 mm. long; styles 1-2 t as long as the stigmata; pseudostaminodia almost as ње as m sepals alec 7. А. WILLIAMSII about half as long as the sepals 8. А. LAGUROIDES IM б. g 5 о 2 a A ~“ м 3 RE m e 25 я о Nes Gd 2 R ~ 2 P. 2 ‚ STENOPHYLLA ii. Leaves ovate to elliptic, 2.5-10 i 6—50 mm. won peduncle branched or unbranched; "ean ~ stramineous о purple. j. Peduncles анара раа flowers white to stramineous; seeds reddish brown; styles about twice as long as the stigmata....... 10. A. MEXICANA jj. Peduncles Ei um flowers eer seeds reddish brown to black; styles about as long as the stigmata 11. А. PANAMENSIS 1. ALTERNANTHERA SESSILIS (L.) К. Br. Prodr. 417. 1810. Gomphrena sessilis L. Sp. Pl. 225. 1753. Illecebrum sessile L. Sp. Pl. ed. 2:300. 1762. Illecebrum indicum Houtt. Nat. Hist. 2:7. 1777. Alternantbera рабая А Lam. Encycl. 1:95. 1783. Ite era nodiflora К. Br. loc. cit. 41 Allaganthera forskalli Mart. Pl. Hort. Erlang. 69. 1814 on esf. Tabl. Bot. ed. 2: Mena amma opposi sitifolia Hassk. in Flora 31:754. 1851. Alternantbera tenuissima Suesseng. in Bot. Arch. 39:382. 1939. Prostrate or decumbent puberulent or glabrous annuals or perennials, the sparsely branched stems to 60 cm. long. Leaves glabrous or sparsely villous below, (381) (Мог. 48 40 ANNALS OF THE MISSOURI BOTANICAL GARDEN elliptic, apically acute and mucronate, basally cuneate to attenuate, 1—4 cm. long, 5 cm. broad; petioles 1-5 mm. long, often winged. Inflorescences of white or pinkish globose heads, 2-4 mm. long, 2-4 mm. broad, sessile in the axils. Flowers perfect, the bracts and bracteoles subequal, acute, 1 mm. long or less; sepals 5, subequal, transparent, not indurate, 1-nerved, slightly concave, 1-1.5 mm. long; stamens 5, united below into a short tube, the pseudostaminodia entire, exceeded by the anthers, about equaling the filaments; ovary obovoid, style 1, 3—4 times as long as the minute capitate stigma. Fruit an indehiscent obcordate yellowish utricle, 1.5-2 mm. long, usually slightly exceeding the sepals; seeds yel- lowish or reddish brown, cochleate-orbicular, 1-1.2 mm. broad. This pan-tropical weed seems to range from Honduras to Brazil in the Amer- icas. It inhabits open moist areas and may grow as an aquatic, in which case the vegetative portions are unusually large. 2. ALTERNANTHERA POLYGONOIDES (L.) К. Br. Prodr. 417. 1810. ompbrena polygonoides L. = Pi225. 1753. Шина gli iet oides 1. РІ. ed. 2:300. 1762. bes polygonoides ne Lam. Encycl. 1: +A 1785. не polygonoides Mart. Noy. Сеп. & Sp. 2 1826. Alternanthera paronychioides St. Hil. Vm Distr. Dia 2:439. 1833. eiremis repens Raf. Fl. Tell. 3:41. Telantbera polygonoides Moq. in DC. Toir 137:363. 1849. Alternanthera ficoidea Griseb. Fl. Brit. W. Ind. 67. 11 not Gomphrena ficoidea L. Procumbent villous or glabrate perennials, the much branched stems to 80 cm. long. Leaves villous or glabrate, elliptic to ovate or obovate, apically acute to rounded and mucronate, basally cuneate to attenuate, 1-3 cm. long, 0.5-1.5 cm. broad; petioles 2-10 mm. long, often winged and with basal tufts of dentate hairs. Inflorescences of white globose to cylindric heads, 4-10 mm. long, 3-6 mm. broad, ile in the axils. Flowers perfect, the bracts and bracteoles subequal, acute, ovate, 1-1.5 mm. long; sepals 5, the outer 3 broader, 3-nerved, slightly indurate, pilose below, acuminate, 3—4 mm. long; stamens 5, united below into a short tube, the pseudostaminodia dentate, much exceeded by the anthers; ovary obovoid, style 1, about half as long as the capitate stigma. Fruit an indehiscent obcordate to orbicular utricle, 1-1.5 mm. long; seeds reddish brown, often strongly reticulate, 1-1.2 mm. broad. This species, occasionally adventive in eastern United States, ranges from Mexico to Argentina in open dry to moist habitats. Standley (in Contr. U. S. Nat. Herb. 27:174. 1928) reports it from Panama but since he keys it as having simple pubescence, the identity of the specimen or specimens is questionable. Backer (in Fl. Mal. 15:93. 1949) seems to have lumped this species with A. ficoidea (L.) R. Br., but the staminodial differences between the two species appear to be con- stant. Standley (in Field Mus. Bot. = :419. 1937) reports the vernacular name golondrina in Costa Rica. The South American species Alternanthera boliviana Rusby is a very closely allied species, wd acute sepals and unusually long spikes. (382) 1961] FLORA OF PANAMA (Amaranthaceae) 41 3. ALTERNANTHERA REPENS (L.) Gmel. Linn. Syst. Nat. ed. 13?:106. 1791. Achyranthes repens L. Sp. Pl. 205. 1753. Alternantbera achyranth. Forsk. Fl. Aeg.-Arab. 28. 1755. Illecebrum achyrantha L. Sp. Pl. ed. 2:299. 1762. ranthus cras ть villiflore Scheele, in Linnaea 22:149. 1849. Alternanthera parviflora Fawc. & Rendle, Fl. Tio. 9: 139. 1914. Procumbent villous or glabrate perennials, the much branched stems up to 50 cm. long. Leaves appressed villous or glabrate, ovate to obovate, apically acute to rounded and mucronate, basally rounded to attenuate, 5-25 mm. long, 3-15 mm. broad; petioles 2-10 mm. long, often winged. Inflorescences of yellowish or stra- mineous usually cylindric heads, 5-12 mm. long, 4-8 mm. broad, sessile in axils. Flowers perfect, the bracts and bracteoles subequal, 3—4 mm. long; sepals 5, trimorphic, the outer 2 broader, strongly 3-nerved, indurate, pilose below and with rigid awns, the third sepal narrower, shorter and merely acuminate, the inner 2 conduplicate; stamens 5, united below into a shallow tube, the pseudostaminodia entire or dentate, exceeded by the anthers; ovary obovoid, style 1, about as long as the minute capitate stigma. Fruit ап indehiscent obcordate utricle, 1.2-1.5 mm. long; seeds reddish brown, 1-1.5 mm. broad, cochleate-orbicular. Presumably a native of tropical America, this weedy species is now also well established in Asia, Europe and the East Indies. Standley’s recount of the distribu- tion (in N. Am. Flora 21:136. 1916) indicates that the species has been collected in Panama. It certainly is to be expected there. In the Yucatan the Mayas apply the name cabalxtez; in Guatemala the names sanguinaria, sacachiquim and bierba de toro are reported. This species is doubtfully distinct from A. pungens HBK., which differs in having subrotund leaves and longer less villous sepals. 4. ALTERNANTHERA FICOIDEA (L.) В. Br. Prodr. 1:417. 1810. Gomphrena ficoidea L. i^ Pl. 225. 1753. lloc bres ficoideum L. Sp. Pl. ed. 2:300. 1762. Paronychia ficoidea Desf. Кы. Bot. ed. 2:14. 1815. Telanthera снаи зата В brachiata Мод. loc. cit. 364. 1849. Alternanthera polygonoides Griseb. Fl. Brit. W. Ind. 67. 1859. not Gomphrena polygono- ides Alternantbera polygonoides В герту С Quum loc. = ys 1859. Alternantbera versicolor Hort. ex Regel, Gartenfl. 10 uo Alternanthera bettzickianna Nich. Gard. Dict. ed. I: 59: Alternanthera amoena Back. & Sloot. Handb. Thee. 108. no. (383) [Vor. 48 42 ANNALS OF THE MISSOURI BOTANICAL GARDEN Procumbent or decumbent hispidulous or glabrate perennials, the branching stems to 100 cm. long. Leaves slightly pubescent but soon glabrate, elliptic to broadly ovate or obovate, apically acuminate to acute and mucronate, basally rounded to cuneate, 2-6 cm. long, 0.5—2 cm. wide; petioles 2-10 mm. long, usually winged. Inflorescences of stramineous to white globose to ovoid heads, 3-10 mm long, 3—6 mm. broad, sessile in the axils. Flowers perfect, bracts and bracteoles subequal, ovate, aristate, 1.5—3 mm. long; sepals 5, the outer 3 broader, 3-ribbed, basally indurate and hispidulous, 3-5 mm. long; stamens 5, united below into a tube, the pseudostaminodia lacerate and exceeding the filaments; ovary obovoid; style 1, 2-3 times as long as the capitate stigma. Fruit ап indehiscent suborbicular utricle, 1-1.5 mm. long; seeds reddish brown, cochleate-orbiculate, 0.8-1.2 mm. broad. L ZONE: vicinity of Monkey Hill, Cowell 33; Gatun, Sutton Hayes 18 5 near ада 50—200 m., Pittier а Chaerés, Isthmus of Panamá, Fendler 261; Ancón, Greenman & Greenman 5018; low w ground, outskirts of Ancón, Gree 8 Gr eenman nman 5047; near Corrosion Laboratory, vicinity of Miraflores Locks, Stern, Chambers et al. 65. CHIRIQUÍ: vicinity of San Bartolomé, Peninsula de Burica, 0-50 m., Woodson 9 Scbery 041. PROVINCE UNKNOWN: Sutton Hayes 9 e native range of this species is адыр from Mexico to Argentina, but various forms are cultivated elsewhere as ornamentals and soil binders. 5. ALTERNANTHERA HALIMIFOLIA (Lam.) Standl. in Pittier, Pl. Us. Venez. 145. 1926 анге balimifolia Lam. Encycl. 1:547. 1785. 5 elanthera frutescens B acutifolia Moq. loc. cit. 366. 184 Alternanthera ficoidea y balimifolia (Lam.) О. Кате. Rev. cH 539. 1891 зе flavogrisea Urban n, Symb. Ant. 1:300. 1899. ernanthera nero Uline, in Field. Mus. Bot. 1:419. 1899. Telanthera halimifolia A. сбыта ia Proc: Calif. уљу 41; foe: 1911 Alternanthera ficoidea var. flavogrisea Pinos & Rendle, Fl. Jam. 3:140. 1914. Procumbent or decumbent sericeous perennials, the sparingly branched stems to 20 cm. long. Leaves densely pubescent with stellate hairs, rarely glabrate, elliptic to ovate, apically rounded and mucronate, basally rounded to cuneate, 1-5 ст. long, 0.5-2.5 cm. broad; petioles 2-10 mm. long. Inflorescences of stramineous ovoid heads, 4-12 mm. long, 3-5 mm. broad, sessile in the axils. Flowers perfect; bracts and bracteoles subequal, ovate, acuminate to aristate, 1-3 mm. long; sepals 5, the outer 3 broader, 3-ribbed, basally indurate and hispidulous, 3-5 mm. long; stamens 5, united below into a tube, the pseuidostaminodia lacerate and exceeding the filaments; ovary globose to obovoid; style 1, about twice as long as the capitate stigma. Fruit an indehiscent globose utricle 1-1.5 mm. long; seeds reddish brown, cochleate-orbicular, 0.8—1.1 mm. broad. (384) 1961) FLORA OF PANAMA (Amaranthaceae) 43 Inhabiting woods, thickets, pond borders and sea beaches, mostly at low eleva- tions, this species extends from Mexico and the West Indies to Colombia, Chile an the Galapagos Islands. Standley (in М. Am. Flora 21:140. 1917) indicates that it occurs in Panama. The obvious silvery pubescence usually separates this from the preceding species with which it reputedly intergrades. 6. ALTERNANTHERA PHILOXEROIDES (Mart.) Griseb. Abh. Ges. Wiss. Gött. 24: 1879 Bucholzia philoxeroides Mart. Nova Acta Acad. Leop. = 131:315. 1826. Telanthera philoxeroides Мод. т DC. Prodr. 132:362. Telanthera pbiloxeroid. ifolia Мод. 1 t. 363. Telantbera philoxeroides var. 6 phyllantha Seub. in Mart. FL Teas 51;169. 1875. Telanthera philoxeroides var. y denticu . loc. си. 170. в Achyranthes philoxeroides (Mart.) Standl. in Jour. Wash. р Sci 5:74. 1915. Decumbent or ascending glabrate aquatic perennials, the simple or branched, often fistulose stems to 100 cm. long. Leaves glabrous or glabrate, lanceolate to narrowly obovate, apically rounded to acute, basally cuneate, rarely denticulate, 2-10 cm. long, 0.5-2 cm. broad; petioles 1-3 mm. long. Inflorescences of terminal and occasionally axillary white glomes, 10-18 mm. long, 10-18 mm. broad, the usually unbranched peduncles 1-5 cm. long. Flowers perfect, bracts and bracteoles subequal, ovate, acuminate, 1-2 mm. long; sepals 5, subequal, oblong, apically acute and occasionally denticulate, neither indurate nor ribbed, 5-6 mm. long, 1.5-2.5 mm. broad; stamens 5, united below into a tube, the pseudostaminodia lacerate and exceeding the anthers; ovary reniform, the style about twice as long as the globose capitate stigma. Fruit an indehiscent reniform utricle 1 mm. long, 1-1.5 mm. broad; mature seeds not see Standley (іп М. Am. Fl. 21:142. Ut ) gives the range of this species as from Colombia to Brazil and Argentina, adventive in the southeastern United States. I have seen no specimens from Panama but it could easily be adventive there. Suessenguth (in Rep. Spec. Nov. 39:4. 1935) reports a teratological specimen from Pearl Islands in San José in which the stamens are replaced by carpels. ALTERNANTHERA WILLIAMSII (Standl) Standl. in Jour. Wash. Acad. Sci. 15: 458. 1925 Acbyrantbes williamsii Standl. in Contr. U. S. Nat. Herb. 18:89. 1916. Erect or clambering pubescent perennials, the sparingly branched stems to 5 m. long. Leaves appressed pubescent, sometimes variegated with purple, ovate, apically acute to acuminate, basally rounded to acute, 2.5-10 cm. long, 1-4 cm. broad; petioles 2-8 mm. long. Inflorescences of terminal and axillary white to stramineous, globose to ovoid heads, 10—30 mm. long, 10-12 mm. broad, the usually unbranched peduncles 1-7 cm. long. Flowers perfect; bracts and bracteoles subequal, ovate, carinate, aristate, 2-4 mm. 1008; sepals 5, subequal, oblong, 3-ribbed and indurate basally, 4-6 mm. long, 1-1.5 mm. broad; stamens 5, unit below into a tube, the pseudostaminodia lacerate and exceeding the anthers; ovary (385) [Vor. 48 44 ANNALS OF THE MISSOURI BOTANICAL GARDEN globose; style 1, 1-2 times as long as the globose stigma. Fruit an indehiscent obovoid utricle, the seeds reddish brown, cochleate-orbicular, 1.2-1.5 mm. long. CANAL ZONE: between Summit and Gamboa, Greenman & Greenman 5244. COLON: Ahorca Lagarto, Cowell 255. DARIEN: vicinity Yape, Rio Tuira, Allen 4302; Tucuti, Chepigana, Terry & Terry 1388; Cituro, Williams 675. PROVINCE UNKNOWN: Sutton Hayes 949; Halsted in 1850. An inhabitant of dry thickets, this species ranges from Panama to southern Mexico. It is very closely related to the Mexican species A. gracilis (Mart. & Gal.) Standl. which supposedly differs in having globose heads, recurved sepals, trichoto- mous peduncles and black seeds. Intermediate forms do occur however. A. williamsii Е. purpurea Standl., with purplish red leaves, is cultivated in the Canal Zone as an ornamental. 8. ALTERNANTHERA LAGUROIDES (Standl.) Standl. in Standl. & Cald. Lista Pl. Salvador 74. 1925. Achyranthes laguroides Standl. in Contr. U. S. Nat. Herb. 18:90. 1916. Erect or clambering pubescent perennials, the much branched stems to 4 m. long. Leaves densely sericeous, especially below, lanceolate to narrowly ovate, apically attenuate, basally attenuate, 2-10 cm. long, 1-3.5 cm. wide; petioles 1-4 mm. long. Inflorescences of terminal and axillary stramineous to whitish ovoid heads, 5—15 mm. long, 4-12 mm. broad, the peduncles to 9 cm. long, branched or unbranched, naked or leafy (when leafy, the heads may appear sessile). Flowers perfect; bracts and bracteoles subequal, ovate, acuminate to aristate, 2-3.5 mm. long; sepals 5, narrowly oblong, 1-3 nerved, scarcely indurate, 4-5.5 mm. long, with basal hairs almost exceeding the sepals; stamens 5, united below into a tube, the lacerate pseudostaminodia exceeding the anthers; ovary obovoid to globose; style 1, 2-3 times as long as the globose stigma. Fruit an obovoid utricle to 1 mm. long; mature seeds not seen. According to Standley & Steyermark (in Field Mus. Bot. 24:149. 1946) this species ranges from Guatemala to Panama in dry thickets of the Pacific slope. In Guatemala it is called botoncito. Possibly Moquin was using this species in his description of Telanthera pubiflora a monocephala, the type of which was collected in Panama. I have not seen the type, but believe the description fits A. laguroides more closely than any of the other Central American species. 9. ALTERNANTHERA STENOPHYLLA (Standl.) Standl in Field Mus. Bot. 8:9. 1930. Achyranthes stenopbylla Standl. in Contr. U. $. Nat. Herb. 18:90. 1916. қ Ascending or erect sparsely strigose or glabrate perennials. Leaves sparsely pilose or glabrate, linear to linear-elliptic, apically acute, basally acuminate, 2.5-5 cm. long, 3-6 mm. broad; petioles 2-6 mm. long. Inflorescences of axillary and terminal stramineous cylindric heads, 6-10 mm. long, 5-6 mm. broad; peduncles unbranched, 2-5 cm. long. Flowers perfect, the bracts and bracteoles subequal, ovate, aristate, 1-2 mm. long; sepals 5, oblong, obscurely 3-nerved and indurate (386) 1961] FLORA OF PANAMA (Amaranthaceae) 45 basally, 2-2.5 mm. long; stamens 5, united below into a tube, the lacerate pseudo- staminodia exceeding the anthers; ovary aia style 1, 2-3 times as long as the globose stigma. Mature fruit not seen PROVINCE UNKNOWN: Sutton Hayes 941. The type for this species was collected by Sutton Hayes in the year 1859-1860 and the only given locality was Panama. Apparently it is known only from the type collection. Except for the linear leaves, this species very strongly resembles the following species. 10. ALTERNANTHERA MEXICANA (Schlecht.) Hieron. Bot. Jahrb. 20: Beibl. 49:8. Brandesia mexicana Schlecht. in Linnaea 7:392. 1832. Telantbera microcepbala Moq. in DC. Me и :371. 1849. ber. LM Ci Alternantbera lebmannii Hieron. in Bot Jed. 20: Beibl. 49:8. 1895. Acbyrantbes lebmannii (Hieron.) Standl. in Jour. Tub. Acad. 5:74. 1915. cbyrantbes mexicana (Schlecht.) Standl. тд cit. 915 Ascending or erect pilose or glabrate si aS ы scantily branched stems to 1 m. long. Leaves appressed pilose or glabrate, elliptic to ovate, apically attenu- ate, basally acute to cuneate, 2.5—10 cm. long, 1-5 cm. broad; petioles 2-15 mm. long. Inflorescences of axillary and terminal white to stramineous subglobose to cylindric heads, 5-10 mm. long, 4-7 mm. broad; peduncles mostly unbranched, to 9 ст. long. Flowers perfect, the bracts and bracteoles subequal, ovate, acuminate to aristate, 1-2 mm. long; sepals 5, oblong, obscurely 3-nerved and slightly indurate basally, 2-3.5 mm. long, 1 mm. broad; stamens 5, united below into a tube, the lacerate pseudostaminodia exceeding the filaments; ovary globose to obovoid; style 1, about twice as long as the globose stigma. Fruit an indehiscent obovoid utricle, 1-1.5 mm. long; seeds reddish brown, cochleate-orbiculate, about 1 mm. broad. PROVINCE UNKNOWN: Sutton Hayes 945. Inhabiting mesic forests, this species extends from Mexico south locally to Colombia. It is of more than casual interest that Sutton Hayes 045 is A. mexicana, Sutton Hayes 044 is the type of A. panamensis and Sutton Hayes 041 is the type of A. stenophylla. The flowers of all these specimens are very similar indeed and would suggest that the latter two were mere vegetative variants of A. mexicana. 11. ALTERNANTHERA PANAMENSIS (Standl) Standl in Field Mus. Bot. 8:9. 1930. Achyranthes panamensis Standl. in Contr. 1). S. Nat. Herb. 18:89. 1916. Erect or clambering pilose perennials, the stems much branched. Leaves appressed pilose, ovate to elliptic, apically acute to acuminate, basally subacute, 2-5.5 cm. long, 0.5-2 cm. wide; petioles 1-4 mm. long. Inflorescences of terminal and axillary stramineous to purplish globose to cylindric heads, 7-11 mm. long, about 7 mm. broad; peduncles mostly branched, 1-6 cm. long, villous. (387) [Vor. 48 46 ANNALS OF THE MISSOURI BOTANICAL GARDEN „+ > агае Сул ЖУРУ Fig. 120. Alternanthera panamensis (habit and lower details) and A. mexicana (upper details) (388) 1961] FLORA OF PANAMA (Amaranthaceae) 47 Flowers perfect; bracts and bracteoles subequal, ovate, acuminate to aristate, 1-2 mm. long; sepals 5, oblong, strongly 3-nerved and indurate below, 2.5-3.5 mm. long; stamens 5, united below into a tube, the lacerate pseudostaminodia exceeding the filaments; ovary obovoid to subglobose; style 1, about as long as the globose stigma. Fruit an indehiscent subglobose utricle; seeds dark reddish brown to black, cochleate-orbiculate, 1 mm. broad. PROVINCE UNKNOWN: Sutton Hayes 044. This species, represented solely by the type collection, is perhaps just a casual variant of A. mexicana (Schlecht.) Hieron. It seems to bear the same relation to А. mexicana that the branched-pedunculate A. gracilis (Mart. & Gal.) Standl. bears to the unbranched-pedunculate A. williamsii (Standl.) Standl. 11. GOMPHRENA L. GOMPHRENA І. Sp. Pl. 224. 1753. Coluppa Adans. Fam. Fl. 2:268. 1763. W 1” Turcz. in Bull. Soc. Nat. Mosc. 16:55. 1843. Xeraea О. Ktze. Rev. Gen. 545. 1891. Amarantoides i Fl. Haban. 94. 1897. Flowers perfect, bracteate and bibracteolate, sessile in spikes or heads. Sepals 5, basally connate or discrete, hypogynous, concave, subequal, colorless or brightly colored. Stamens 5, hypogynous, the filaments connate forming a deep stamen tube, the anthers sessile at the summit of the tube; pseudostaminodia 2-lobate; anthers 2-locellate, introrse, medially attached. Ovary globose to obovoid, 1- locular, 1-ovulate, the ovule campylotropous on an elongate flattened funicle; style 1, filiform, about as long as the 2 filiform stigmata. Fruit an indehiscent utricle; seeds cochleate-orbiculate. Pubescent erect or decumbent annual or perennial herbs. Leaves opposite, sessile or petiolate, entire, usually pubescent, apparently deciduous. Inflorescences of terminal and occasionally axillary globose to cylindric solitary or clustered heads, usually subtended by foliar leaves. This genus of nearly a hundred species, with some native to tropical America and others to tropical Austral-Asia, seem to reach its best development in South America. Several species are widely cultivated, as is the "bachelor's button", G. globosa, a rather handsome ornamental with edible leaves. Four species are known from Central America but only two are reported from Panama. a. Heads ru mm. in diameter; bracteoles 4—6 mm. long; leaves 1—5 lon cm. 1. G. DECUMBENS 2. G. GLOBOSA aa. Heads 15-25 mm. in diameter; bracteoles 7-12 mm. long; leaves 2- 10 cm. lon = 1. GOMPHRENA DECUMBENS Jacq. Hort. Schoenbr. 4:41. 1804. Xeraea decumbens O. Ktze. Rev. Gen. 545. с Gomphrena decumbens f. albiflora Chod. му in Bull. Herb. Boiss. 23:389. 1903. Gomphrena decumbens £. roseiflora Chod. & Hassl. loc. cit. 389. 1903. (389) [Vor. 48 48 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 121. Gomphrena decumbens Gomphrena ixiamensis Rusby, in Bull. М. Y. Bot. Gard. 6:502. 1910 Gomphrena cumbens var. genuina Stuchlik, in Rep. Spec. Nov. 11: :156. 1912. Gomphrena decumbens var. grandifolia Stuchlík, bie cit. 157. 1912. Gompbrena perennis subsp. pseudodecumbens Stuchlik, mee cit. 153. 1912. Gomphrena perennis f. simplex Stuchlik, loc. cit. 153. 1912. Gomphrena d ispersa Standl. in Contr. U. S. Nat. Herb. eS 91. 1916 Gom тепа 1, Gomphrena decumbens var. carinata Suesseng. in Rep. Sp. Nov. 39: 8. 1935. Prostrate or decumbent pilose annual ог perennial herbs, the much branching stems to 50 cm. long, often rooting at the nodes and forming mats. Leaves seri- ceous to strigose, occasionally glabrate, oblong to ovate or obovate, apically acute to rounded and mucronate, basally acute to attenuate, 1-5 cm. long, 0.5—2.5 cm. broad; petioles 1-5 mm. long. Inflorescences of terminal and axillary ovoid to cylindric whitish or purplish heads, occasionally clustered, 4-20 mm. long, 5-14 mm. broad, subtended by ovate leaves 0.5-2 cm. long. Flowers perfect; bracts (390) 1961] FLORA OF PANAMA (Amaranthaceae) 49 persistent, ovate, acuminate, 1.5-3 mm. long; bracteoles narrowly to broadly cristate, equaling or exceeding the sepals, 4—6 mm. long; sepals 5, subequal, basally connate, cinereous- to rufous-lanate, ultimately indurate and closely embracing the fruit; stamens 5, subequal, the anthers sessile between the 5 2-lobate pseudo- staminodia; stamen tube 3—6 mm. long; ovary globose; style 1, 1-1.5 mm. long; stigmata 2, 1-1.5 mm. long. Fruit ап indehiscent areolate utricle 1-2 mm. long; seeds cochleate-orbiculate, yellowish to reddish brown, ocellate, 1.5-1.7 mm. broad. BOCAS DEL TORO: Research grounds, region of Almirante, Cooper 138. CANAL ZONE: е . 08; Апсбп епта Greenm. White & White 102; Pedro Miguel, Bro. Heriberto 19. cmHiRiQuí: vicinity of Puerto Armuelles, 0—75 m., Woodson & Schery 819. This species ranges from the West Indies and Mexico to Panama and from Brazil and Bolivia to Argentina. Common names in Guatemala are bofoncillo, sangrinaria, sanguinaria, siempreviva and siempreviva de monte; other names іп the Yucatan and adjoining regions are chacmol, tmuul, amor seco and secicante. Standley (in Contr. Т). $. Nat. Herb. 18:91. 1916) has separated a variant, with the crests of the bracteoles conspicuously widest at the apex, as G. dispersa, which seems to occur throughout the range of G. decumbens. No correlated differences exist and transitions occur, so G. dispersa is reduced to synonymy. As Suessenguth (in Rep. Spec. Nov. 39:8. 1935) has noted Greenman & Greenman 5015 is intermediate between G. decumbens and G. nitida Rothr., a questionably distinct species supposed to differ in having lacinate-dentate crests, larger rarely purple heads and an erect habit. Palmer gor from Mexico has crests both dentate and laciniate-dentate, or even absent; the specimen also has both included and exserted stamen tubes. This particularly illuminating example serves to demonstrate that С. nitida is hardly more deserving of specific status than is С. dispersa. 2. GOMPHRENA GLOBOSA L. Sp. Pl. 224. 1753. Gomphrena globosa В carnea Мод. in DC. Prodr. 132:409. 1849. 1849, Amarantoides globosus Maza, Fl. Haban. 94. 1897. а ис globosus albiflorus Maza, loc. cit. 95. 1897 Сотрђтепа globosa var. aureiflora Stuchlik, in Rep. be Nov. 12:340. 1915. Fils or ascending pilose annual herbs, the branching stems to 60 cm. high. Leaves sericeous, occasionally glabrate, oblong to ovate or obovate, apically acute and mucronate, basally rounded to attenuate, 2-10 cm. long, 0.5-5 cm. broad; petioles 1-15 mm. long. Inflorescences of terminal and axillary reniform to cylindric variously colored heads or spikes, occasionally clustered, 10-25 mm. long, 15-25 mm. broad, subtended by ovate leaves 5-20 mm. long. Flowers perfect; bracts persistent, ovate, acuminate, 3—4 mm. long; bracteoles dentate-cristate, exceeding the sepals, 7-12 mm. long, 2—3.5 mm. broad; sepals 5, subequal, basally connate, cinereous- to rufous-lanate, ultimately indurate and closely embracing the fruit, 5-6.5 mm. long; stamens 5, the anthers sessile between the 5 2-lobate (391) у [Vor. 48 50 ANNALS OF THE MISSOURI BOTANICAL GARDEN pseudostaminodia; stamen tube 4-8 mm. long; ovary globose; style 1, 1-1.5 mm. long; stigmata 2, 1-1.5 mm. long. Fruit an indehiscent areolate utricle 1.5-2.5 mm. long; seeds cochleate-orbiculate, reddish brown, ocellate, 1.5-2 mm. broad. Cultivated and escaped in many regions of the world, this species may be a native of America although it was originally described from India. Standley (in Contr. U. S. Nat. Herb. 27:173. 1928) reports that it occurs in Panama as a cultivant and an escape. In Panama it is called siempreviva and suspiro. The names amor seco, inmortal and botón are also used in Guatemala. Some of these vernacular names suggest the persistence of the flowerss which makes them popular in wreaths and dry floral arrangements. That the differences between this and the preceding species are quantitative may suggest a polyploid derivation. (392) 1961] FLORA OF PANAMA (Nyctaginaceae) 51 NYCTAGINACEAE * Herbs, shrubs, or trees, sometimes scandent, the stems frequently swollen at the nodes, particularly in the herbaceous genera, and sometimes armed with axillary spines. Leaves alternate, opposite, whorled, or approximate, simple, estipulate, entire. Flowers monochlamydeous, perfect or unisexual, in various cymose inflorescences, occasionally solitary, the bracteoles often enlarged and involucrate, herbaceous or petalaceous and enclosing one or more flowers. Perianth monochlamydeous, herbaceous or petalaceous, tubular to infundibuliform, cam- panulate, urceolate, or salverform, involute-plicate in aestivation, the base of the tube persistent in fruit, the limb persistent or deciduous. Stamens 1 to many, hypogynous, the filaments unequal, usually united at the base, the anthers included or exserted, dorsifixed near the base. Ovary 1-celled, containing a solitary basal ovule; style elongate, short, or lacking. Fruit a fleshy, coriaceous, or woody anthocarp, occasionally bearing viscid stipitate glands. Nyctaginaceae are an essentially tropical and subtropical family, негу іп the New World. The Four-o’clock Family is represented in Panama by indigenous genera, but more familiar to the general public are the нача genera Bougainvillea and Mirabilis. Bougainvillea, almost ubiquitously planted as an ornamental woody vine because of its handsome clusters of three showy magenta, red, pink, white, or yellow bracts subtending the inconspicuous flowers, is a Sout American genus which does not appear to naturalize readily in Panama. The plants variously are referred to B. glabra Choisy and B. spectabilis Willd. It usually is known by its Latin name, or by the vernacular veranera and flor de verano because of its habit of blooming in the dry season. Mirabilis jalapa L., the common four-o'clock of the English, is a herb with showy red, white, yellow, or variegated flowers set in individual green calyx-like involucres. It is cultivated in gardens and escapes as a vigorous weed in waste ground and roadsides. It is known locally as maravilla and buenas tardes because of its vespertine blooms. The taxonomy of the Nyctaginaceae, particularly of the woody dioecious genera, is very confused. It seems entirely possible that interspecific hybridization may be one factor responsible for obscuring the lines of the species. The species bibliographies which follow are frankly provisional; to some, additional synonyms will have to be added from the South American literature and some such names possibly will have to take precedence over the Central American which are adopted here. The detailed study essential to a competent treatment unfortunately is beyond the scope of this flora. A. Herbs; flowers bisexual 1. BOERHAVIA AA. Trees, aier and woody lianas; flowers unisexual, dioecious e flowers with deeply „ар onm зуны раце seta ok Si se e ovary sessile and free fro es; eoles borne ее Қатық the poe кебек unarm C. Stamens 25—30; stamin nths broadly Е: anthocarps woody, not вааще she perianth limb persistent. 2. CEPHALOTOMANDRA * With the collaboration of Harold J. Kidd. (393) (Мог. 48 52 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN CC. Stamens 5-10, usually 8; staminate perianths tubular ьн anthocarps ius. the perianth limb withering = t length abscis . NEEA BB. Rol te бе wers Shen te, with widely exserted stamens; pistillate flowers with the ovary stipitate. C nts unarmed; anthocarps fleshy, not кессе. the perianth mb pe rsistent; bracteoles borne immediately beneath the peri- ашы; же же е flowers with the stam inodes about as long as the 2 bei ring conspicuous sterile 4. GUAPIRA СЕ: Plans Sealy armed with stout гаад spines; anthoc iaceous, ris with co: — век glands; сме по e born mediately beneat e perianths; pistillate flowers with th сны: — c ed to a low, occasionally minutely kar Берић 45 5. PISONIA 1. BOERHAVIA L. Borrnavia L. Sp. РІ. 3. 1753. Boheravia Parodi, in Anal. Soc. Cient. Argent. 5:210. 1878. Boer id auct. Annual or perennial herbs. Leaves opposite or subopposite, petiolate, frequently unequal at the nodes, the blade undulate or sinuate, conspicuously raphidulous. Inflorescence terminal, paniculiform to racemiform, many-flowered. Flowers small, perfect, subtended Бу 1-3 minute bracteoles, the perianth tube persistent, more or less herbaceous, frequently glandular, the perianth limb deciduous, petalaceous, obscurely 5-lobed; stamens 1-5, included or exserted; ovary shortly stipitate, the stigma capitate, more ог less exserted. Anthocarp obpyramidal to obovoid or subellipsoid, glabrous to glandular, 3- to 5-angulate Numerous species in the tropics and рее of both hemispheres. Both species in Panama are weeds and possibly not indigenous. a. Annuals; stems erect or ae бе or sini за = igi reddish-punc tate beneat h; infloresc not lea e flow 3-flowered dichasia, distinctly pedicellate; х0 бет е truncate, glabrous, conspicu aly рефсе 1. В. ЕВЕСТА а 8 "d И i Ih Ww Е d bent, more or = ise re not punctate vds $ orescences leafy-bracted, the flowers in small glomerulate s, sessile or жебесі Ды nthocarps obovoid to subellipsoid, obtuse clus or camels beaked, и. sessile or subsessile 2. B. DIFFUSA 1. BOERHAVIA ERECTA L. Sp. 3. 1753. Boerhavia elongata Salisb. Rane 56. 1796. Boerbaavia virgata НВК. Nov. Gen. 2:215. 1817. Boerbaavia dis е HBK baavia atomaria Raf. Aut Ман latifolia Mart. & Gal. in Bull Kad Brux. 111:124. 1844. paniculata B subacu ig in DC. Prodr. 137:451. 1849. oka thornberi М. Е. Jones, Contr. West. Bot. 12:72. 1908. — erecta thornberi (М. E. jon Standl. in Contr. U. S. Nat. Herb. 12:381. Б = age d herbs; stems erect or ascending, 2—8 dm. tall, rather slender, glabrous or essentially so. Leaves opposite or subopposite, often unequal at the nodes, petiolate; blade rather irregularly ovate-rhombic, obtuse to rounded, or infrequently (394) 1961] FLORA OF PANAMA (Nyctaginaceae) Fig. 122. Boerbevie diffusa; insert: fruit of B. erecta (395) 53 [Vor. 48 54 ANNALS OF THE MISSOURI BOTANICAL GARDEN acute, at the apex, broadly obtuse to rounded at the base, undulate or sinuate, 2-9 cm. long, 1-4 cm. broad, glabrous or essentially so, yellowish green above, paler and more or less conspicuously bracteate, glabrous or essentially so. Flowers in pedunculate 2- to 3-flowered dichasia, distinctly pedicellate, the perianth tube 1.0-1.5 mm. long, about 0.5 mm. broad, herbaceous, glabrous, the perianth limb campanulate, about 0.5-0.7 mm. long, white or pink; stamens 2-3, slightly exsetted. Anthocarps obpyramidal, truncate, glabrous, 3-4 mm. long, 1.0-1.5 mm. broad, borne upon accrescent pedicels of about equal length. small weed of beaches, open thickets, fields, roadsides, and wasteland; southern United States, Antilles, Central and South America; a rare sporadic ruderal in tropical Africa. AL ZONE: Gamboa, Bro. Heriberto 28, Standley 28461; Monte сава Махоп 6860; Obispo, Standley 31775; Gatün, Cowell 296; Frijoles, Piper 5829; New Frijoles, Piffier 6830; Balboa, Bro. Celestine 10; Paraiso, Pittier 2520; Summit, Standley 30133; кк Hill, Cowell 28. снікюоі: Puerto Armuelles, Woodson & Schery 838. сотом: Colón Macbride 8 Featherstone т, Piper 5892, Козе 220 The glabrous anthocarps and Mia есен of В. erecta have made this species less widely dispersed as a ruderal than B. diffusa. 2. ВОЕВНАМА DIFFUSA L. Sp. Pl. 3. 1753. Boerhaavia coccinea Mill. Gard. гы с. ed. 8 P no. 4. 1768. Boerbavia caribaea Jacq. Obs. Bot. ыз ets Boerbavia diandra Aubl. Hist. РІ. EDS 1:4. Boerbaavia paniculata A. Rich. Act. Soc. Hist. Nac. Paris 1:105. 1792. Boerhaavia polymorpha A. Rich. loc. cit. 185. Boerhavia hirsuta Willd. Phytog. 1:1. 1794. Boerhaavia adscendens Willd. Sp. Pl. 1:19. 1798. Boerhaavia viscosa Lag. & Rodr. in Anal. Ci. Nat. 4:256. 1801. Boerhaavia decumbens Vahl, Enum. Pl. 1:284. 1805. Boerbaavia laxa Pers. Syn. Pl. 1:36. 2 Boer] avia squamata Raf. Aut. s 40. B eR E 5; Net Herb. 1:111. 1891. Бы diffusa var. hirsuta Qi O. Ktze. Rev. Gen. 533. E [pK cas diffusa var. paniculata (Rich.) O. Ktze. loc. cit. 189 Boerbaavia diffusa var. viscosa (Lag. & Rodr.) Heimerl, Beitr. А Nyct. 27. 1897. Ез viscosa UE (anil Se eg in Ann. А n 5” Genéve 5:189. 1901. Boerhavia ramulosa М. Е. Jone West. Bot. 10:40. Boerhaavia viscosa ен E л in Contr. U. $. Nat. Heb. 12:383. 1909. Boerbaavia ixodes Standley, loc. cit. 13:423 Boerbaavia coccinea forma parcebirsuta Heimerl, in n Urb. Symb. Ant. 7:212. 1912. Perennial herbs; stems decumbent to ascending, 4-12 dm. long, rather slender, more or less viscid-puberulent. Leaves opposite or subopposite, often unequal at the nodes, petiolate; blade orbicular to rhombic-ovate below, frequently grading to lanceolate above, obtuse to rounded or acute at the apex, obtuse to rounded at the base, undulate or sinuate, 2-6 cm. long, 2-5 cm. broad, more or less puberulent to glabrate, yellowish green above, paler beneath, not punctate; petiole 1-4 cm. ong. Inflorescences paniculiform, lax and many-flowered, leafy-bracteate at least below, viscid-puberulent to glabrate. Flowers in pedunculate glomerulate clusters, sessile or subsessile, the perianth tube 0.7-1.0 mm. long, somewhat less than 0.5 (396) ч 1961] FLORA OF PANAMA (Nyctaginaceae) 55 mm. broad, herbaceous, minutely glandular, the perianth limb campanulate, about 0.5 mm. long, red to purple, rarely white; stamens 3, slightly exserted. Anthocarps obovoid to subellipsoid, obtuse to obscurely beaked, 3—4 mm. long, 1 mm. broad, glandular, sessile or subsessile. A weed widely dispersed throughout the tropics and subtropics of America, Africa, Asia, and Oceania. Carasola (Taboga Isl.). BOCAS DEL TORO: Lincoln Creek, Chaguinola ас Dunlap 377. CANAL ZONE: Fri- joles, Wilson 51; Empire, Hunter & Allen 786; A , Greenman & Greenman 5042; between Corozal and Апсбп, Pittier 2188; P^ pte Du to poe Cowell 369; Balboa, Macbride & Featherstone 39; Monte Lirio, Maxon 6847; Gamboa, Bro. Heriberto 29, Standley 28344, Piper 5649; Gatün, Standley 27321; Darién, Standley 31520, Mac- ride 2684. CHIRIQUÍ: Puerto Armuelles, Woodson t$ Schery 820. PANAMA: Taboga Island, Pittier 13 534, eg 3177, AN a 101, Е Ани 27845; San José Island, Pearl Archipelago, Harlow 47. : per Boerbavia erecta m B. pw Т grow together, and hybridization must occur occasionally but insufficiently to blur the outline of the species. In Standley's treatments of Boerbavia for the North American Flora and the Flora of the Panama Canal Zone, the parental species would be called B. erecta and B. caribaea, respectively, and the putative hybrids B. coccinea. Such plants will usually key readily to B. diffusa in the present treatment. A rather extensive examination of herbarium specimens has revealed no tangible differences between B. caribaea and B. coccinea of the New World and B. diffusa of the Old World. The species appears plainly to be an effusive pantropic weed, the dispersal of which has been greatly facilitated by the viscid anthocarps. It is probably, but not certainly, of American origin. 2. CEPHALOTOMANDRA Karst. & Triana CEPHALOTOMANDRA Karst. & Triana, ex Karst. in Linnaea 28:429. 1856. Dioecious trees. Leaves opposite to irregularly approximate. Inflorescences terminal, corymbosely thyrsiform. Flowers small, immediately subtended by 1-3 minute bracteoles. Staminate flowers broadly campanulate; stamens 25-30, unequal, included, the short tube of the filaments hypogynous and free from the sessile pistillode. РізеШате flowers unknown. Anthocarps woody, eglandular, longi- tudinally costate, the limb of the perianth persistent. A single species, Panama (?) and Colombia. 1. CEPHALOTOMANDRA FRAGRANS Karst. & Triana, ex Karst. in Linnaea 28:429. 1856. Cepbalotomandra panamensis Standl. in North Am. Flora 21:179. 1918. Dioecious trees to 25 m. tall. Branches rather stout, glabrous, with a longi- tudinally striate, scarcely lenticellate, yellow periderm. Leaves rather sparse, long- petiolate, the blade broadly elliptic to ovate-elliptic, Obtusely acuminate to acute, base obtuse or rounded, 8—25 cm. long, 2.5-15.0 cm. broad, glabrous, firmly membranaceous or subcoriaceous, yellowish green, venation prominent but rather (397) [Vor. 48 56 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 123. Cephalotomandra fragrans distant; petiole 1-12 cm. long. Inflorescence 10-20 cm. long, long-pedunculate, many-flowered, minutely puberulent, the ultimate cymules typically dichasial with the terminal flower sessile and 1-bracteolate and the lateral definitely pedicellate and 3-bracteolate. Staminate perianth broadly campanulate, yellowish, about 4 mm. long and 3 mm. broad at the conspicuously inflated, slightly spreading limb, minutely puberulent-papillate. Anthocarps oblong-fusiform, crowned by the persistent perianth limb, about 1 cm. long and 3 mm. broad, glabrous. CANAL ZONE (?): locality unknown, Sutton Hayes 908. The holotype of C. panamensis, cited above from the herbarium of the New York Botanical Garden, is suspiciously similar in all essential details to an isotype (398) 1961] FLORA OF PANAMA (Nyctaginaceae) 57 of С. fragrans (Triana 008) deposited in the herbarium of the Missouri Botanical Garden and originally distributed by the British Museum. It may be merely a peculiar coincidence, but it seems more than strange that both sheets bear a small fruiting twig and a small twig with a staminate inflorescence in identical stages of development, and that both the Triana specimen and that ascribed to Sutton Hayes bear the collection number 098. Karsten's description of Cephalotomandra is extremely vague in details of the pistillate inflorescence, and the account of the genus in Bentham & Hooker’s Genera Plantarum, although describing the fruit and the staminate flowers, expressly states that the pistillate flowers were not seen by them. I think that it is altogether probable that the New York sheet is actually a second duplicate of Triana’s original collection of C. fragrans and that its ascription to Sutton Hayes is in error. It is doubtful, therefore, that the genus Cephalotomandra can be included in the flora of Panama without serious question. 3 МЕГА R & P Neea К. & Р. Fl. Per. Prodr. 52. 1794. Neeania Raf. Princ. Som. 3 Mitscherlichia Kunth, in Abi. "Berl. Akad. еў 209. 1831. Eggersia Hook. f. in Hook. Ic. 15:1. Ё. 1401. 1883. Dioecious trees and shrubs. Leaves opposite to irregularly approximate. In- florescences terminal, thyrsiform. Flowers small, immediately subtended by 1-3 minute bracteoles. Staminate flowers urceolate; stamens 5-10, usually 8, unequal, included, the short tube of the filaments surrounding the sessile pistillode. Pistillate flowers urceolate or tubular-urceolate, the limb of the perianth erect or spreading, the staminodes usually somewhat surpassing the ovary and with enlarged sterile anthers, the pistil sessile. Anthocarps fleshy, eglandular, the limb of the perianth withering and at length abscissing. Numerous species in the Antilles, and Central and South America; southern peninsular Florida. a. Cymules strikingly dichasial, the terminal flowers sessile, the lateral with "iy long slender pedicels; leaves predominantly rather small or of m ewhat pubescent beneath. rate size, and only very rarely somew b. Mesi usually quite Leer the pedicels t lateral р, Сы long as the perianths or som t longer; staminate perianths o urceolate; anthocarps breathe ellipsoid 1. N. DELICATULA bb. Inflorescences minutely puberulent, the pedicels as long as perianths or somewhat shorter; staminate perianths oblong-urceolate; рр narrowly ellipsoid 2. М. LAETEVIRENS aa. Cymules obscurely dichasial, both terminal and lateral flowers pines or with similar short stout — radi predominantly large and frequently more or less pubescen 3. N. AMPLIFOLIA 1. NEEA DELICATULA Standl. in Contr. U. S. Nat. Herb. 18:98. 1916. Torrubia panamensis Standl. in North Am. Fl. 21:185. 1918. Dioecious shrubs or small trees to 6 m. tall. Branches rather slender, glabrous or minutely puberulent when very young, inconspicuously lenticellate at maturity. (399) (Уот. 48 58 ANNALS OF THE MISSOURI BOTANICAL GARDEN Leaves petiolate, the blade predominantly obovate to obovate-elliptic, acuminate, obtusely cuneate at the base, 4-12 cm. long, 2—4 cm. broad, firmly membranaceous, glabrous, the petiole 2-15 mm. long. Inflorescences 4-7 cm. long, usually glabrous, rarely minutely puberulent, rather few-flowered, the peduncles slender, the cymules strikingly dichasial, the terminal flowers sessile, the lateral with slender pedicels as long as the flowers or somewhat longer. Staminate perianths ovoid-urceolate, about 6 mm. long and 3 mm. broad, glabrous. Pistillate perianths tubular, about 3 mm. long and somewhat less than 1 mm. broad, minutely puberulent to glabrous. Anthocarps broadly ellipsoid or ellipsoid-obovoid, 8-10 mm. long, 4-5 mm. broad, pink or red. Panama, in forests and thickets near sea level to 100 m. alt. NAL ZONE: Balboa, Standley 32114. PANAMA: around Alajuela, Chagres peril Pittier 3472; Tumba Muerto Road, near Panama, Standley 29805; near Matias Hernandez, Standley 28880; Rio Tapia, Standley 28201; TOM eds ари = аре bland, Perlas Islands, Allen 2422. PROVINCE UNKNOWN: Seemann s.n. pro par I believe it quite possible that N. ds a N. карб Ernst, of Colombia and Venezuela (= N. wiesneri Heimerl) may be conspecific. The leaves of the latter, however, appear to be somewhat larger than those of the former, as well as of a somewhat different pattern of shape; the staminate perianths, also, seem to be somewhat narrower than those from Panama The type specimen of Torrubia panamensis definitely is Neea and differs from the other specimens cited only in the thin minute pubescence of the peduncles. Comments on the Seemann collection are included in the discussion of Guapira standleyana. Seemann’s illustration of Pisonia pacurero (in Bot. Voy. Herald, pl. 34. 1854) may have been prepared from a specimen of Neea delicatula, but is not definitely diagnostic. 2. NEEA LAETEVIRENS Standl. in Field Mus. Publ. Bot. 4:204. 1929. Neea xanthina Standl. in Ann. Missouri cis ата 30:86. 1943. Neea psychotrioides auct. pro parte, поп D Dioecious shrubs or small trees to 7 m. jet Branches rather slender, minutely puberulent to glabrate, inconspicuously lenticellate at maturity. Leaves petiolate, the blade elliptic to obovate- or oblong-elliptic, rarely lanceolate, acuminate, obtuse to cuneate, 5-25 cm. long, 3-9 cm. broad, glabrous or rarely very inconspicuously puberulent beneath particularly on the midrib, the petiole 0.5—4.0 cm. long. In- florescences 8-20 cm. long, minutely puberulent, many-flowered, the cymules strikingly dichasial with the terminal flowers sessile and the lateral with slender pedicels as long as the flowers or somewhat shorter. Staminate flowers oblong- urceolate, 6-9 mm. long, 2-3 mm. broad, thinly and minutely puberulent, white, yellow, or red. Pistillate flowers tubular, the lobes erect or suberect, conspicuously thickened at the orifice of the tube, 4-5 mm. long, 1.0-1.5 mm. broad, densely and minutely puberulent. Anthocarps narrowly ellipsoid, 7-10 mm. long, 2-3 mm. broad, г Panama and Costa Rica, in moist forests and thickets near sea level. (400) 1961] FLORA OF PANAMA (Nyctaginaceae) 59 i № №, 4 Fig. 124. Neea laetevirens BOCAS DEL TORO: Old Bank Island, Von Wedel 1885, 1970; La Bocas, Von Wedel 2533; Fish Creek Hills, Von Wedel 2433; Shepherd Island, Von Wedel 2695; Rio Crica- mola, Woodson, Allen 8 Seibert t 1014; region of Almirante, кабине 340, 544; ow а SAN BLAS: Permé, Cooper 2o This is the species which has been determined almost invariably as Neea psycho- trioides in Panama. The specimens enumerated above are relatively uniform; but the species apparently is capable of hybridization with №. am plifolia, although such (401) (Мог. 48 60 ANNALS OF THE MISSOURI BOTANICAL GARDEN Mte will usually key to the latter under the discussion of which they will be no This probably are scarcely more than a half dozen valid species of Neea in Central America. These seem to form two vicarious series upon the Pacific and the Atlantic slopes, recognizable by the staminate perianths which are ovoid- or oblongoid-urceolate respectively. True М. psychotrioides is а representative of the former series, as is N. delicatula of Panama. Both Panamanian species are distinguishable from their related species to the north by their longer-pedicelled flowers and proportionally longer lobes of the pistillate perianths. True N. psycho- trioides extends on the Pacific slope from southern Mexico to Nicaragua and is not yet known to occur in Costa Rica, although it might be expected in Guanacaste. In Costa Rica, N. laetevirens has been collected at Puerto Limón, and up the Reventazón and Parismina watersheds to the highlands of the provinces of Alajuela and Cartago. 3. NEEA AMPLIFOLIA Donn. Sm. in Bot. Gaz. 61:386. 1916. Neea pittieri Standl. in Contr. О. S. Nat. Herb. 13:383. 1911. Neea orosiana Standl. in Journ. Wash. Acad. Sci. 15:473. 1925. Neea urophylla Standl. in Field Mus. Publ. Bot. 4:203. 1929. Neea pycnantha Standl. in Ann. Missouri E Gard. 30:85. 1943. Neea psychotrioides auct. pro parte, non Donn. Sm. Dioecious shrubs and small trees to Үз 7 m. tall. Branches moderately stout, densely ferruginous-pubescent to glabrate. Leaves petiolate, the blade ovate- to obovate-elliptic, usually more or less caudate-acuminate, obtuse to cuneate at the base, 7-36 cm. long, 3-15 cm. broad, firmly membranaceous, glabrous above but frequently more ог less puberulent beneath, the petiole 0.5-5.0 cm. long. Inflorescences 4-15 cm. long, essentially glabrous to densely ferruginous-pubescent, rather few-flowered to many-flowered and occasionally greatly proliferous and the flowers and bracts becoming phylloid; cymules obscurely dichasial, both terminal and lateral flowers subsessile or with essentially similar short stout pedicels. Stami- nate perianths tubular-urceolate, 5-10 mm. long, 2-3 mm. broad, essentially glabrous to minutely puberulent to essentially glabrous. Anthocarps ellipsoid to ovoid, 7-12 mm. long, glabrous to minutely puberulent, red Panama and Costa Rica, in forests and moist thickets, from near sea level to about 700 m. ос. region, Cooper & Slater =. ; ин valley, "tra 9 Slater бо, 140; easels Cooper 491; Fish 5466 Mts., Von Wedel 2267; vicinity of Chiriqui Lagoon, Von Wedel 1011, 1017; Water Valley, Von Wedel 1574, 1609, 1503, 1433, 027. CANAL ZONE: hills north of я Standley 27427, 27500; Margarita Swamp, south of France Field, Maxon 8 Уа е 70. atun, Sutton Hayes 626; railroad berti between Gorgona and Gatün, Mie 2265, 2271; Вагго Colorado ада. Frost 123, Kenoyer 491, 638, Aviles 30, Bailey 8 Bailey 120, 522, Wetmore & Abbe 22, Standley 31310, 31336, 40905, 932 41023, 41015. CHIRIQUÍ: Progreso, Cooper & Slater 184. сосіЖ: region north of E Valle de Antón, Allen п 373 2. COLON: between France Field and Catival, Standley 301 г along Rio Fató, Pittier 3912. DARIEN: trail between Pinogana and Yavisa, Allen 268; poor around Pinogana, Pittier 6547. SAN BLAS: јони ы Puerto Obaldia, Pittier (402) 1961] FLORA OF PANAMA (Nyctaginaceae) 61 Neea amplifolia usually can be keyed without difficulty from the two preceding species, particularly by means of the inflorescence characters, the predominantly larger leaves and staminate perianths, and the more frequent and abundant indu- ment. In all respects intermediates occur, nevertheless, and the relative abundance of disparate specimens, particularly from Bocas del Toro, where N. laetevirens also is abundant and variable, provides grounds for strongly suspected hybridization. The proliferation and phyllody of the inflorescence is a frequent and puzzling propensity of this species, the agent of which is obscure. In addition, both in N. amplifolia and in N. laetevirens, the terminal staminate flowers occasionally develop at the expense of the lateral. 4. GUAPIRA Aubl. Guapma Aubl. Hist. Pl. Санап. 1:308; 3: Ё. 110. 1775. Gynastrum Neck. Elem. 1:224. 1790. Torrubia Vell. Fl. Flum. 139. 1825; Icon. 3: #. 150. 1827, non Tul. Dioecious trees and shrubs. Leaves opposite to irregularly approximate. In- florescences terminal, frequently at the tips of short lateral branches, corymbosely thyrsiform. Flowers small, immediately subtended by 1—3 minute bracteoles. Staminate flowers campanulate; stamens usually 6-8, unequal, widely exserted, the short tube of the filaments adnate to the stipe of the pistillode. Pistillate flowers tubular to tubular-campanulate, the limb of the perianth erect or spreading, the staminodes about as long as the ovary and with enlarged sterile anthers, the short tube of the filaments adnate to the stipe of the pistil. Anthocarps fleshy, eglandular, the limb of the perianth persistent and rather fleshy. Numerous species in the Antilles, and Central and South America. There can be little doubt that Спарта Aubl. is congeneric with Torrubia Vell. as suggested by Heimerl (in Engl. Nat. Pflanzenfam. ed. 2. 16°:127. 1934) and not an indecipherable verbenacea as previously held by many (although questioned by Bentham & Hooker). The specimen illustrated by Aublet obviously is pistillate and, with the understandable error of the four to five minute "leaflets of the calyx [i.e. bracteoles]", is a good representation of Torrubia for its time, particularly with respect to the fruit with its persistent perianth limb (“Drupa monosperma . . coronata denticulis corollae, quae evadit pulposa") , and the persistent style crowned by a penicillate stigma. With the exception of the supernumary bracteoles, the figure of the flower also is good, showing the spreading perianth limb characteristic of so many South American Torrubias, and the widely exserted style and penicillate stigma. ‘The single species described by Aublet, С. guianensis, perhaps is con- specific with Torrubia eggersiana (Heimerl) Standl. Perhaps too many transfers from Pisonia to Torrubia already have been made by Britton and by Standley, and one might be inclined to submit Torrubia as a candidate for conservation over Guapira in order to avoid additional transfers to the latter. I am inclined not to do so, however, for the following reasons. Tor- rubia has not generally been considered as distinct from Pisonia, and consequently (403) [Vor. 48 62 ANNALS OF THE MISSOURI BOTANICAL GARDEN the transfers to the former are not in wide use; transfer to Guapira therefore would entail little mental adjustment and inconvenience at this date. Both Torrubia and Guapira are based upon rather inadequate antique illustrations, but those of the latter are definitely more capable of critical interpretation than the former; it urthermore will be much easier to associate G. guianensis with a modern species than T. opposita Vell. a. Anthocarps globose or subglobose; pistillate perianths about m. lon the limb spreadin ing; stamens about half exserted; leaves predominantly elliptic and acum . С. COSTARICANA 22. KERREN blong fusiform pistillate perianths about 3 mm. long, lim about one-third exserted; leaves sr eris Pace UE aed бс 2. С. STANDLEYANA 1. СОАРЩА costaricana (Standl.) Woodson, comb. nov. Torrubia costaricana Standl. in Contr. U. S. Nat. Herb. 13:385. 1911. Small dioecious trees to 10 m. tall. Branches moderately stout, glabrous, prominently lenticellate at maturity. Leaves rather crowded, petiolate, the blade predominantly elliptic to oblong-elliptic, acuminate, obtuse at the base, 2.5—10.0 cm. long, 1.5—4.5 cm. broad, subcoriaceous, glabrous, the petiole 0.5—1.0 cm. long. Inflorescences 3—6 cm. long, rather shortly pedunculate, many-flowered, minutely puberulent, conspicuously accrescent in fruit, the flowers sessile. Staminate peri- anth campanulate, about 3 mm. long, about 0.5 mm. in diameter at the base and 2 mm. at the orifice, minutely puberulent, greenish white or yellow, the stamens about half exserted. Pistillate flowers tubular-campanulate, the limb slightly spreading, about 2 mm. long and 0.5 mm. in diameter, minutely puberulent- papillate. Anthocarps globose ог subglobose, about 7-8 mm. long, black or purple when ripe. Costa Rica and Panama, in forests and savannas, sea level to 1000 m. CHIRIQUÍ: Boquete, Davidson 689; David, Pittier 3370. состЕ: Valle de Antón, Bro. Maurice 777; north rim of El zum Alston 8 Allen 1847; region north of El Valle de Antón, trail to Las Minas, Allen 2. Brother Maurice reports the d dd names of llanto and mala sombra. Guapira costaricana is widely distributed also on the Pacific slope of Costa Rica. In both countries the leaves of plants growing at higher elevations seem broader and more elliptic than those of lower elevations. The fruit, however, is relatively constant. 2. Guapma standleyana Woodson, spec. nov. Arbor dioica usque са. 10 m. alta. Кати! graciliusculi indistincte lenticellati juventate tenue puberuli mox glabrati. Folia densiuscula sed mox decidua petiolata lamina praecipue obovata vel obovato-elliptica brevissime acuminata vel obtusa basi latiuscule cuneata 3-15 cm. longa 1-6 cm. lata firmiter membranacea glabra petiolo 0.5-1.0 cm. longo. Inflorescentiae 4-8 cm. longae minute puberulae in fructu paulo accrescentes floribus sessilibus vel subsessilibus. Perianthium mas- culum campanulatum ca. 3.5 mm. longum faucibus ca. 2 mm. diam. minute (404) 1961] FLORA OF PANAMA (Nyctaginaceae) 63 Fig. 125. Guepira costaricana puberulo-papillatum staminibus ca. tertio exsertis. Perianthium femineum tubu- losum limbo erecto vel fere erecto ca. 3 mm. longum et 0.5 mm. diam. minute puberulum. Anthocarpia oblongo-ellipsoidea 10—13 mm. longa ca. 6-7 mm. lata rubra (aetate immatura?). Panama, in savannas and bush, near sea level. CANAL ZONE: near mars Memorial бесеу в [Cocoli], Р. White че bank 2: Río о 73 oli, е 104. РАМА за ama City, Bro. Paul aboga Island, Pittier 54 (US, тия Е); vicinity of леде Егапсо Касе тас пеаг чана ma, Standley 27747; between nd Matias Hernandez, Standley 31823. 5АМ pro parte. (405) (Мог. 48 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN Sharply distinguished from G. costaricana because of the rather narrow oblong- ellipsoid fruit. Two dupla of the Seemann collection seen in the herbarium of the New York Botanical Garden and the Gray Herbarium consist of mixtures; the former of Guapira standleyana and Меса delicatula, and the latter of Спарта standleyana, Neea delicatula, and Pisonia aculeata. This mixture may be responsible for Seemann’s remarks on the “hermaphrodite” flowers of his collection, which he cited under Pisonia pacurero HBK., and the difficulty of distinguishing Neea from Pisonia (Seemann, in Bot. Voy. Herald, p. 192. 1854). Seemann’s plate of P. pacurero clearly is a pistillate Neea, although not definitely N. delicatula. Seemann cited his collection from "dark forests, near Cruces.” 5. PISONIA L. Ріѕомта Г. Sp. РІ. 1026. 1753. Bessera Vell. Fl. Flum. 147. 1825; Icon. 4: Ё. 12. 1841, nec Schult, пес ced Columella Vell. loc. cit. 155. 1825; Icon. 4: Ё. 17. 1841, пес Comm. пес Lour. Pallavia Vell. loc. cit. 151. 1825 ; Ico n. 4: 2. I2. 1841. Shrubs or small trees, сені scandent and armed with stout axillary spines in our species. Leaves opposite to irregularly approximate. Inflorescences usually terminal on highly modified short shoots, umbelliform or densely corymbiform- thyrsiform. Flowers small, the subtending bracteoles in a more or less contracted spiral upon the pedicel. Staminate flowers campanulate; stamens usually 6-8, unequal, widely exserted, the short tube of the filaments adnate to the stipe of the pistillode. Pistillate flowers tubular, the limb of the perianth apparently always erect, the staminodes reduced to a low, occasionally glandular-dentate disc adnate to the stipe of the pistillode. Anthocarps pentagonal-clavate, coriaceous, armed with longitudinal rows of stipitate glands upon the angles. Numerous species in the Antilles, Central and South America; southern penin- sular Florida; western Africa, southeastern Asia and Oceania. A single species is known from Panama at present. 1. Pisonta ACULEATA L. Sp. Pl. 1026. 1753. Pisonia villosa Poir. in Lam. Dict. 5:347. 1804. Pisonia sieberi Schlecht. in Linnaea 12:876. 1822. Pisonia loranthoides HBK. Nov. Gen. & Sp. 7:197. Pisonia monotaxadenia Wr ight, ex Sauy. in Anal. жей x. Habana 7:199. 1870. Pisonia grandifolia Standl. in Contr. Џ. 8. Nat. Herb. 13:391. 1911 Divaricately branched shrub or woody liana, usually Вы with stout recurved axillary spines, glabrous to densely pubescent. Leaves opposite to irregularly approximate, the blade very variable in shape and size, usually obovate to obovate- elliptic, acute to acuminate, 2-10 cm. long, 1-5 cm. broad, the petiole 1-3 cm. long. Inflorescences usually borne at the tips of greatly condensed short shoots, umbelliform or densely corymbiform-thyrsiform, rather shortly pedunculate, 3-6 cm. long, greatly accrescent and expanded in fruit, many-flowered. Staminate flowers campanulate, greenish yellow, about 3 mm. long, about 2 mm. in diameter at the orifice of the limb, minutely puberulent-papillate without, the stamens (406) 1961] FLORA OF PANAMA (Nyctaginaceae) 65 Fig. 126. Pisonia aculeata widely exserted. Pistillate flowers tubular to somewhat urceolate, about 2.5 mm. long and 1 mm. in diameter, the limb erect, the stigma and style widely exserted. Anthocarps borne upon greatly elongate pedicels, pentagonal-clavate, 7-9 mm. long, 3—4 mm. broad, green, coriaceous, the stipitate glands uniseriate upon the five angles, the sides minutely and densely puberulent. Very widely distributed in tropical America, and apparently a ruderal introduc- tion in western Africa and southeastern Asia; in rain forests and moist thickets from sea level to about 1000 m. Barro Colorado Island, SP ‚4093 3, Standley 40018. сосіЕ: hills Е va EA Valie S Antón, Allen 2835. Juan Diaz, Standley Ж. Rio Tapia, Standley 28251. PROVINCE UNKNOW it: "Servii s. n. pro parte. This species must be very much more frequent than the records would indicate. It is popularly known as ийа de gato, in allusion to the stout recurved thorns. (407) (Мог. 48 66 ANNALS OF THE MISSOURI BOTANICAL GARDEN BATIDACEAE Batis maritima L. is to be expected in Panama. This dioecious herbaceous to fruticose plant with opposite, simple, entire and sessile leaves is the sole species of the family. It is an inhabitant of the beaches of subtropical and tropical America and the Hawaiian Islands. PHYTOLACCACEAE By KATHERINE RAEDER Herbs, shrubs or trees, often vines. Leaves simple, entire, alternate, petiolate or subpetiolate, with or without stipules. Inflorescence in terminal or extra-axillary racemes, although occasionally axillary. Flowers bisexual or unisexual by reduction, actinomorphic; perianth monochlamydeous; tepals generally 4—5, free or slightly connate at the base, usually persistent in fruit; stamens 3—many, often varying in number within the same species, frequently borne on a hypogynous disc in 1 or 2 cycles opposite or alternate with the tepals, the filaments free or basally connate; ovary superior, rarely inferior, 1- to 16-carpellate, the carpels free to partly or entirely connate, the styles when present as many as the carpels, usually terminal, sometimes subterminal, generally free but occasionally united, the stigma capitate, or sessile and penicillate; ovules basal, campylotropous, solitary. Fruit a berry, ре, or achene, the seeds 1-many. Embryo coiled ог bent around the perisperm. a. Flowers subsessile in uu or axillary racemes; tepals persistent in fruit, erect; fruit an achen . PETIVERIA aa. Flowers obviously на tepals persistent in fruit (except in Phytolacca rivinoides), reflexed, erect, or spreading; fruit a berry o ru b. Tepals 5; ovary 2- to many-carpellate c. Ovary 6- to 15 ан кује free but connivent; stamens 6-33; fruit a many-seeded ber 2. PHYTOLACCA сс. pet Рене one 2, “fe and recurved; stamens 3-9, ually t a 1-seeded d 3. MICROTEA bb. udi 4; ovary ee d. Tepals erect in fruit; stamens 4; fruit a berry. 4. RIVINA dd. Tepals spreading or reflexed in fruit; stamens 8 (-25); fruit а drupe 5. TRICHOSTIGMA 1. PETIVERIA L. Petena L. Sp. Pl. 342. 1753. Tall herbs, sometimes woody at the base. Leaves alternate, simple, entire, petiolate, minutely stipulate. Inflorescence an elongate terminal or axillary raceme, the flowers small, subsessile, bracteate and bracteolate. Tepals 4, united into a short tube, the lobes subequal, linear, persistent and erect in fruit. Stamens 8, inserted irregularly on a hypogynous disc at the base of the ovary, the filaments filiform and of varying lengths, the anther linear, dorsifixed. Ovary 1-carpellate, (408) 1961] FLORA OF PANAMA ( Phytolaccaceae ) | 14 (409) [Vor. 48 68 ANNALS OF THE MISSOURI BOTANICAL GARDEN oblong, tomentose, 4- to 6-uncinate, the stigma 1, sessile, penicillate, decurrent along the ventral margin of the ovary, the ovule 1, basifixed. Fruit a linear achene; pericarp coriaceous and adherent to the seed. Seed 1, linear, the testa membranous, the albumen scanty and mealy, the cotyledons foliaceous. Two species in the Western Hemisphere. 1. РЕТІУЕМІА ALLIACEA L. Sp. РІ. 342. 1753. Petiveria octandra L. Sp. Pl. 486. 1762. Petivera foetida Salisb. Prodr. 214. 1796. Petiveria alliacea В grandifolia (L.) Мод. in DC. Pis 13?:9. 1849. Petiveria alliacea y octandra (L.) Mog. loc. cit. 18 Petiveria ochroleuca Moq. loc. cit. 1849. Petiveria paraguayensis id in Anal. Soc. pl Gore 160. 1878. Petiveria hexandria Sesse & Moc. Fl. Mex. ed. 2. 894. Tall herbs, often tem at the base, ма pe sometimes angled stems, to 5-10 dm. tall, with a strong odor of garlic. Leaves elliptic to obovate, the apex acute or acuminate, often mucronate, the base narrowed, 5.5—16.0 cm. long, about 2—6 cm. broad, slightly pubescent; petioles 1.5-2.0 cm. long. Inflorescence in slender usually sparsely flowered racemes 15-40 cm. long; pedicels up to 1 mm. long. Tepals white or greenish-white, sometimes pale pink, often basally pubescent, about 3—5 mm. long. Stamens 8, free, up to 3 mm. long. Ovary 4-uncinate, the hooks becoming elongate and quite prominent in fruit. Fruit linear, 8 mm. long. United States, Florida to Texas; throughout the West Indies and Central Amer- ica; South America, Colombia to Argentina; common in dry or moist fields and forests, often found near habitations, especially on waste ground, up to 1500 meters. Some of the common names are: Garlic-weed, Obeah-bush, Anomu, and Guinea-hen weed. $ DEL тово: Chiriqui Lagoon, Von Wedel 1323; Water Valley, Von Wedel 636. — Puerto Armuelles, Woodson & Schery 841, 900; between Remedios and David, P. White 308; Río Dupi, Pittier 5222. CANAL ZONE: Ancón Hill, Standley 26336; Las Cruces Trail, Government forest, Hunter & Allen 731, 694. PANAMA: Taboga Island, к 1201; бенен Ын 27025; Woodson, Allen 8 Seibert 1470; Macbride 2791. N: Pin x n quibu alliacea closely resembles P. tetrandra, a species limited in geographical distribution to Brazil. However, the two may be separated fairly readily by t presence in P. tetrandra of 6 hook-like processes in the ovary and fruit rather than the 4 found in P. alliacea. Furthermore, the flower pedicels in P. tetrandra are about 5 mm. long, whereas those of P. alliacea are so short that the flowers appear to be sessile. 2. PHYTOLACCA L. РНУТОГАССА L. Sp. Pl. 441. 1753. Sarcoca Raf. Tell. 3:55. 18 Pircunia Moq. in DC. Prodr. D :29. 1849. Tall perennial herbs or shrubs. Leaves alternate, simple, entire, petiolate, ex- stipulate, generally glabrous. Inflorescence a terminal, or extra-axillary raceme; (410) 1961] FLORA OF PANAMA (Phytolaccaceae) Fig. 128. Phytolacca rugosa (411) 69 (Мог. 48 70 ANNALS OF THE MISSOURI BOTANICAL GARDEN the pedicels basally bracteate, often with 1 or more bracteoles above. Tepals 5, equal, usually glabrous, persistent or deciduous in fruit. Stamens 8-22, inserted at the base of the calyx usually on a hypogynous disc, in 1 or 2 cycles, the filaments free, the anthers dorsifixed. Ovary subglobose, 5- to 16-carpellate, the carpels completely or only partly united, the styles equal in number to the carpels, terminal, generally connivent, often recurved, the ovule solitary in each cavity. Fruit a globose 5- to 16-celled fleshy berry. Seed 1 in each cavity, the endosperm mealy. About 26 species, tropical and subtropical, mostly in the Americas, some in Africa, Asia, and Asia Minor. a. —— connate at the base, the apices free; styles free; tepals reflexed 1. P. RUGOSA aa. — completely p styles с but connivent; tepals either erect or spreading in fruit, or deciduo b. ср 6—10; stamens 2 Же) erect or spreading in fruit; raceme up to 20 cm. lon 2. P. ICOSANDRA bb. mu ав л: stamens 9-22; tepals deciduous in fruit; raceme up cm. lon 3. P. RIVINOIDES 1. РНУТОГАССА RUGOSA Вг. & Bouché, Ind. Sem. Hort. Berol. 13. 1851. Linnaea 25:297. 1852 Woody herbs up to 2 У; m. tall, the glabrous branches erect and angled. Leaves lanceolate-elliptic, the apex acuminate, the base attenuate, 2-5 cm. broad, 4-14 cm. long. Racemes suberect, terminal or extra-axillary, 4-15 cm. long, the flowers fairly crowded; pedicels about 3.5—4.0 mm. long, the bracts about 4 mm. long, the bracteoles about 1 mm. long. Tepals oblong-elliptic, the apex rounded, white or pink, reflexed in fruit, 2-3 (-4) mm. long. Stamens 6-12 in 1 cycle, inserted оп а subhypogynous disc, about 1.5-2.0 mm. long. Ovary subglobose, 6- to 8-carpellate, the carpels free at the apex, the styles equal to the number of carpels and free. Fruit a purple berry, 6-8 mm. in diameter. In Panama, appearing ас 1200-2000 meters; generally somewhat higher altitudes throughout the rest of the range, Mexico southwards to Colombia. The common name is jaboncillo. $ DEL ToRO: Robalo тЫ nude slopes of Cerro Horqueta, Allen 4008. CHI- RIQUÍ: end from Paso Ancho to M Lirio, рағы valley of Rio Chiriqui Viejo, Allen I 511; Seibert 302; foot of Sierra del Воо» чеге, Mauric ; vicinity of Casita Alta, Volcan iriquí, Woodson, Allen & Seibert 975; north о face о о Copete, eastern spur of Volcán de Chiri riqui, Allen 4871; vicinity of Callejón Мо Volcan de Chiriqui, oodson & Schery 480; trail from Bambito to Cerro Punta, P. rugosa is distinguished from P. icosandra by the КЕ union of the carpels, the shorter racemes, and the tendency towards fewer stamens and carpels. The free apices of the carpels may be observed equally well in flower or in fruit. The styles of the flower are fairly close together, but are definitely free and not connivent; in fruit they are pulled farther apart by the expansion of the seeds within the carpels. The staminal cycle is generally considered to be the inner one, the outer cycle having been suppressed. As in most species of Phytolacca the number of stamens is variable, fluctuating even between flowers of the same inflorescence. (412) 1961] FLORA OF PANAMA (Phytolaccaceae) 71 2. PHYTOLACCA ICOSANDRA L. Syst. ed. 10. 1040. 1849. Phytolacca malabarica Crantz, Inst. 2:484. 1769. Phytolacca mexicana Crantz, loc. cit. 1769. Phytolacca mexicana Gaertn. Fruct. 1:377. 1788. Phytolacca triquetra Moench, Meth. Suppl. 107. Phytolacca bogotensis HBK. Nov. Gen. et Spec. 2:1 Phytolacca ри Kunth & Bouché, Ind. Sem. ‘Hort. Saal 15. 1848 Ann. Sci. Nat. Phytolacca 5... Hort. ex Мод. in DC. Prodr. 13?:33. 1849. Phytolacca longespica Мод. loc. си. m deinde a purpurascens A. Br. & Bouche, Ind. Sem. Hort. Berol. 13. 1851. Linnaea 25: 1 "Pisas macrostachys Wilid. ex J. A. Schmidt, in Mart. Fl. Bras. 142:344. 1872. Phytolacca nova-bispania Millsp. in Field Mus. Pu bl. Bot. 2:41. 0. Phytolacca icosandra var. angustitepala Н. Walt. in ган. "eau 483:61. 1909. Phytolacca icosandra var. sessiliflora H. Walt. loc. cit. Stout herbs 1—2 meters tall, sometimes rather еен with sharply angled and often pubescent branches. Leaves narrowly elliptic or sometimes ovate-elliptic, the apex acute, the base narrowed and decurrent along the petiole, 6.5—15 ст. long, 2.3—5.0 cm. broad. Inflorescence an elongate pubescent raceme of densely crowded flowers, 16-30 cm. long; pedicels 1-1.5 mm. long; bracts lanceolate- linear, 3—4 mm. long, the bracteoles about 0.8 mm. long. Tepals broadly elliptic or obovate, the apex rotund, pink, white, or sometimes greenish, occasionally pubescent, 2.5—3.5 mm. long. Stamens 8-20, inserted іп 2 cycles on а disc at the base of the perianth, about 3 mm. long. Ovary 6-8 carpellate, the carpels com- pletely united, the styles free but connivent, often recurved. Fruit a subglobose purple berry 6-8 mm. in diameter. Seed shiny black. Mexico, the West Indies, Central America um to Ecuador. Most commonly found up to 1450 meters in Panama. The common name is jaboncillo. BOCAS DEL TORO: Changuinola Valley, Dunlap gor. The taxonomic controversy over this perplexing species has probably arisen from the variability of the distinguishing characters. The single most distinctive ше which separates it from Р. rugosa is that of whether or not the carpels е ovary are entirely connate. To the unpracticed eye this is a tricky character, Bur it is the most reliable. The connivent styles are the best clue to the unite carpels, as even in fruit they are closely clustered at the center of the berry in spite of the expansion of the ovary which would tend to pull them outwards. The remaining separative characters are best expressed as strong tendencies. The pedicels vary in length but in the Panamanian representatives are usually about 1 mm. long. The number of carpels may vary from flower to flower in the same inflorescence, but 8- to 9- carpellate ovaries are the most common. The racemes wsd to longer than those of P. řügosa, and the flowers are often more crow ` Walter recognizes two varieties of which one, P. icosandra var. sessili fri occurs in a limited range. The subsessile flowers he considers to be important enough to afford varietal status. By “subsessile” it is intended that the pedicel is about 1 mm. in length, as no specimens with a shorter pedicel were found. Nearly all individuals (413) (Мог. 48 72 ANNALS OF THE MISSOURI BOTANICAL GARDEN examined had pedicels of 1 mm. which would place them in this category, although the typical variety with a pedicel of about 3 mm. is reportedly widespread through- out Panama. Herbarium sheets from adjoining countries show a gradual tendency toward longer pedicels, and it is probable that this character is expressed by gradual Variation, as are so many others. 3. РНУТОГАССА RIVINOIDES Kunth & Bouché, Ind. Sem. Hort. Berol. 15. 1848. Ann. Sci. Nat. Ш. 11:231. 1849, Phytolacca icosandra Wright, Mem. 268. 1828, non Linn. (1759). diens polystigma Benth. ex к Moq: loc. cit. 184 м аж polystyla Schomb. ex Мод. loc. cit. 1849. Woody herbs or weak shrubs up to 5 m. tall, with angled branches. Leaves ovate, elliptic, or even lanceolate, the apex mucronate, sometimes only acuminate, the base tending to be decurrent, 4.5-7.0 cm. broad, 9-17 cm. long, the petioles 0.9—4.5 cm. long. Racemes terminal or extra-axillary, the flowers uncrowded along the axis, 30-55 (-70) cm. long; pedicels 7-12 mm. long, the basal bract subulate, about 1.5 mm. long with 2 bracteoles above. Tepals white to red, some- times cream, elliptic, about 2 mm. long, deciduous in fruit. Stamens 9-14 (-22), inserted on a hypogynous disc, about 2 mm. long. Ovary globose, 12- to 16- carpellate, the carpels united throughout their length, the styles as many as carpels, connivent, and recurved. Fruit a purple berry 5-6 mm. in diameter. Throughout the West Indies; Mexico southwards to Bolivia. Found as high as 1680 meters, but generally below 1500 meters. The common name is jaboncillo. BOCAS DEL ТОКО: Water Valley, Von Wedel 833, 706; Chiriqui Lagoon, Von Wedel 2079, 2631, 1246, 2460, 2704; Fish Creek Hills, Von Wedel 2460; Gar her Creek, 263. CHIRIQUÍ: Вајо Chorro, Boquete district, Davidson 153. = forests near Arraijan, Woodson, Allen t$ Seibert 1392. COLON: und Dos Bocas, Rio Fató Valley, Pittier 4205. сост.Е: ЕІ Valle de Anton, along Rio Indio vet Hunter & Allen 313. Although P. rivinoides appears quite distinct from P. icosandra by virtue of its longer pedicels, deciduous tepals, more numerous carpels, smaller fruit and strik- ingly elongate racemes, nevertheless some specimens were examined which are suspiciously a mixture of greater or lesser degree of the two species. The most easily recognized clue to a questionable individual is a tendency for the pedicels to be longer than normal for P. icosandra and shorter than P. rivinoides. In com- bination with this character are usually found intermediate tendencies for raceme length, fruit size, and carpel number. It is interesting to note that even in plants showing extreme tendencies towards P. rivinoides the tepals are always persistent. Furthermore, the leaves of many of these intermediates are sometimes larger than found in either of the two species, although this is not always true. (414) 1961] FLORA OF PANAMA (Phytolaccaceae) 73 3. MICROTEA Swartz MicROTEA Swartz, Prodr. 53. 1788. Schollera Rohr, in Skirvt, Naturhist. Selsk. Kjoeb. 2:210. 1792. Ancistrocarpus НВК. Nov. Gen. et Sp. 2:186. #. 122. 1817. Potamophila Schrank, Pl. Rar. Hort. Nom. 2. t. 63. 1819. Ceratococca Willd. in Roem. et Schult. Syst. 6:800. 1820. Aphanathe Link, Enum. Hort. Berol. 1:383. 1821. Sprawling, decumbent or sometimes spreading annual herbs. Leaves alternate, simple, entire, petiolate or subpetiolate, exstipulate. Inflorescence extra-axillary or terminal, racemose, shortly pedicellate, bracteate. Flowers minute. Tepals 5, oblong, persistent and erect in fruit. Stamens 5, inserted at the base of the perianth, alternate with the tepals, the filament filiform, the anther dorsifixed. Fig. 129. Microtea debilis (415) [Vor. 48 74 ANNALS OF THE MISSOURI BOTANICAL GARDEN Ovary globose, 2-carpellate, unilocular, the styles 2, basally connate, the ovule 1, basifixed. Fruit a minute drupe, tuberculate, the pericarp adherent to the seed. Seed 1, the testa crustaceous, the embryo bent, the albumen scanty. About 9 species in the American tropics. 1. MicROoTEA DEBILIS Sw. Prodr. 53. 1788. Scbollera debilis Rohr, in Skirvt, Naturh. ж Kjoeb. 2:210. Microtea ovata Palle " Hore: жазаңы” ~ x Мод. in DC. ве p :17. 1849. Microtea debilis vi Microtea debilis 8 оваа Mid. is cit. 1849. Decumbent herbs to 50 cm., the stems sharply angled. Leaves elliptic to ovate, the apex acute, sometimes mucronate, the base attenuate, 1.0—2.3 cm. wide, 1.3-3.6 cm. long, glabrous. Inflorescence a many-flowered raceme 1.5—3.5 cm. long; bracts membranaceous, persistent, about 1 mm. long; pedicels about 1.0 mm. long. Tepals 5, lanceolate, white, about 0.5—0.7 mm. long. Stamens about 0.4 mm. long. Ovary globose, about 0.5 mm. in diameter. Fruit with the tubercles united into a honeycomb like pattern, 1.0-1.5 mm. in diameter. Throughout the West Indies; from Guatemala southward to Peru and Brazil. CANAL ZONE: Chagres, Fendler 109; hills between Río Grande and Pedro Miguel on road to Arraiján, Pittier 2709; Frijoles, Б е 31470; к of oo Standley 30145, 26968; пака. Standley 25826; ‚ Вто. Heriberto 115. BOCAS DEL TORO: Changuinola Valley, Dunlap 402. гура и Che epo, Pittier т Рапата City, Bro. Paul 166; Taboga Island, Standley 27088; Ехровісіба, Bro. Heriberto 277. DARIEN: Boca de Cupe, Allen 883. Microtea is a striking genus not only because of the extremely minute flowers, but also because the flowers seem to demonstrate a very much simplified flower plan. The relationship of Microtea to the rest of the family is obvious but it also suggests close affinity to some members of the Chenopodiaceae, particularly to the genus Chenopodium. Although the number of stamens are usually equal to and alternate with the petals, occasionally 8 stamens will be found irregularly inserted. The tubercles vary in length, and although usually rather short, may be as long as 0.5 mm. 4. КІУІМА Г. Rivina Г. Sp. Pl. 121. 1753. Tithona L. Syst. ed. 1. 1735. Rivinia L. Gen. Pl. = ‘ 57. 3754. ee: Moench, Meth. 307. Tithonia L. ex О. Kuntze, Rev. 2: 552. 1891, sphalm. Tall a woody herbs, frequently shrub-like, erect or straggling. Leaves alternate, simple, entire, petiolate, exstipulate. Inflorescence a suberect many- flowered terminal or axillary raceme. Flowers small, pedicellate. Tepals 4, sub- equal, obovate-oblong, rounded or pointed at the apex, persistent and erect or spreading in fruit. Stamens 4, inserted at the base of the perianth in 1 cycle alternate with the tepals, the filaments free, the anther dorsifixed. Ovary (416) 1961] FLORA OF PANAMA (Phytolaccaceae) 75 Fig. 130. Rivina humilis 1-carpellate, the style subterminal, short, curved, the stigma capitate, the ovule 1. Fruit a globose red berry, the pericarp adherent to the seed, the testa crustaceous. Three species in the American tropics and subtropics, introduced into Asia and ustrali (417) [Vor. 48 76 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1. RiviNA ном Г. Sp. Pl. 122. 1753. Rivina humilis а canescens L. Sp. Pl. ыў 175$, Rivina humilis 9 glabra E loc. cit. 17 Rivina laevis L. Mant. 4 67 Piercea glabra Mill. Gard. Dict. ed. 8. Piercea no. 1. Rivina lanceolata Willd. Enum. Hort. Berol. E T 1813. Rivina tetrandra Desfl. Tabl. ed. 2. 49. 1815. ин» puberula HBK Sans Gen. & Sp. 2:184. 1817. a tinctoria Ham. ex G. Don. in Loud. Hort. Brit. Suppl. 1:598. 1832. Rivina portulaccoides Tec. in Trans. Am. Phil. Soc. П. 5:167. 1837. Rivina acuminata Raf. New Fl. 4:13. 1838, поп НВК. (1817). Piercea acuminata Raf. loc. cit. 1838. Rivina obliquata Raf. loc. cit. 1838. Piercea obliquata Raf. лос. cit. 1838. Rivina canescens G. Don. in Steud. Nom. Bot. ed. 2. 2:460. 1841. Rivina laevis В acuminata Мод. in DC. Prodr. 132:12. 1849. Rivina viridiflora Bel, Anal. Soc. Esp. Hist. Nat. 12:105. 1883 Rivina paraguayensis Parodi, in Anal. Soc. Cient. ee 5:206. 1878. Tithonia humilis О. Kuntze, Rev. Gen. 552. 18 Tithonia humilis var. canescens f. albiflora О. ids loc. cit. 1891. — humilis var. glabra O. Kuntze, loc. cit. ; ina humilis laevis Millsp. in Field Mus. Publ. Bot. 2:4 кы humilis var. orientalis (Moq.) Н. Walt. in Engl. ile 453.105. 1909. Straggling shrubs or tall herbs sometimes woody at the base, up to 10 meters high. Leaves elliptic to ovate or lanceolate, the apex acute to acuminate, the base rounded or truncate, 2-6 cm. wide, 4-12 cm. long; petioles 0.6-11.0 cm. long. Flowers small, drooping; pedicels 3 mm. in flower, elongating to about 7 mm. in fruit; bracts lanceolate, about 2 mm. long; tepals 4, white or pinkish white, 2.0- 3.5 mm. long; stamens 4, inserted at the base of the perianth, about 1.5 mm. long. Fruit a scarlet or red berry, about 4.0—4.5 mm. in diameter. Florida to Oklahoma and Texas; throughout Mexico and southward to Argen- tina; West Indies. In Costa Rica found up to 1040 meters elevation. Particularly common on waste ground, often on coastal rocks. Among the common names are Wild Tomato, Bloodberry, and carmin. BOCAS DEL TORO: region of Almirante, Cooper 77; location not stated, Carleton 52; vicinity of "лесе Woodson & Schery 1023; Woodson, Allen t$ Seibert 1807; Water (418) 1961] FLORA OF PANAMA (Phytolaccaceae) 77 Valley, Von Wedel 609; Chiriqui Lagoon, Sarg of Little Bocas, Von Wedel 2526; vicinity of Chiriqui Lagoon, Big pate 5 бы = 882. снікюші: valley of upper Rio Chiriqui аан P. 5 С. White 88. mountains beyond La Pintada, Hunter & Allen 579. DARIÉN: near mouth of Rio Yapé, "Allen pe vicinity of pe de Cupe, Allen 880. Wilke recognizes three varieties of Rivina humilis based on glabrity or degree of pubescence. К. humilis var. orientalis is in Asia, presumably escaped from culti- vation as the genus is probably not indigenous to the region. К. bumilis var. canescens is recognized by the tomentose stems and densely pilose leaves and petioles, and is cited as found in Brazil and the lesser Antilles. К. bumilis var. glabra is, as the names implies, completely glabrous, and is widely distributed from Texas to Argentina. Use of pubescence as a distinguishing varietal character is unconvinc- ing; consequently the specimens examined have been treated as a single species with a wide range of variation in the degree of relationship between pubescence and glabrity of any part of a plant. The synonymy has likewise been regarded as pertaining to a single species. 5. TRICHOSTIGMA A. Rich. TricHosTicMa A. Rich. in Sagra, Hist. Cuba 10:306. 1845. Villamilla R. & B. ex Benth. & Hook. Gen. Pl. 3:81. 1880. Woody vines, shrubs. Leaves alternate, simple, entire, petiolate; stipules minute and deciduous. Inflorescence a many-flowered terminal or extra-axillary raceme, the bracts deciduous. Tepals 4, subequal, persistent and reflexed in fruit. Stamens 8-10 (-12), inserted on a hypogynous disc, the filaments free, the anther dorsi- fixed. Ovary 1-carpellate, unilocular, subglobose, the stigma sessile or subsessile, often penicillate, the ovule 1. Fruit a globose drupe, the pericarp adherent to the seed. Seed 1, the testa crustaceous. Three species in tropical America. a. Flowers in crowded condensed racemes; stamens 8-12; pedicels 7-9 mm. long 1. T. OCTANDRUM aa. Flowers in elongate lax racemes; stamens 20—25; pedicels 8-16 mm. long 2. Т. POLYANDRUM 1. TRICHOSTIGMA OCTANDRUM (L.) Н. Walt. in Engl. Pflanzenr. 45*:109. 1909. Rivina humilis В scandens L. Эр. РІ. eg 1753. Rivina освапата L. Cent. РІ. 2 17 Кита dodecandra Jacq. Obs. ж м >. 1764. Rivina scandens МИ. Gard. Dict. ed. 8. Rivinia по. 2. 1768. Rivina Mutisii Willd. ex Schult. Mant. 3:305. 1827. Rivina americana Raf. Fl. Tell. 3:56. 1837. Trichostigma rivinoides A. Rich. in Sagra. Hist. Cuba. 10:306. 1845. Rivina octandra В obtusifolia Мод. in DC. Prodr. 13:11. 1849. = ў 49. tziana Klotzsch, ex Мод. loc. cit. 184 мы ahuda Hook. f. in Benth. & Hook. Gen. Pl. 3:81. 1880. Decumbent or suberect shrubs or woody vines up to 10 meters. Leaves oblong to elliptic, the apex acute to acuminate, the base rounded or acute, 3.8-5.5 cm. broad, 12.4-14.7 cm. long; petioles 2.4—3.7 cm. long, glabrous. Inflorescence a (419) Fig. 131. Trichostigma polyandrum (420) [Vor. 48 1961] FLORA OF PANAMA (Phytolaccaceae) 79 fairly dense many-flowered raceme, about 5.0-6.5 (-11.0) cm. long; pedicels 3-9 mm. long; bracts lanceolate, 2 mm. long, deciduous, the bractlets triangular, 0.5 mm. long. Tepals white to whitish-green, 2-3 mm. long. Stamens 8—12, inserted irregularly at the base of the perianth, 3—4 mm. long. Fruit a subglobose drupe, black, 5.5—6.0 mm. in diameter. United States (southern peninsular Florida and the Keys); West Indies; throughout Mexico and Central America; South America (Venezuela to Argen- tina); common in abandoned fields and moist woods. NAL ZONE: drowned forest of upper Rio Pequeni between Salamanca cwm ырыс Station and Rio Boquerón, Allen 17270. РАМЕМ: Tucuti, M. Е. 8 К. A. Terry 1390. The Terrys have stated on the specimen label that T. octandrum is a tree, but there are no other indications that trees are even found in this genus. T. octandrum occurs as a woody shrub or liana sprawling over adjacent shrubs and trees. 2. TRICHOSTIGMA POLYANDRUM (Loesener) Н. Walt. in Engl. Pflanzenr. 45?:112. 1909 Rivina polyandra Loesener, in Engl. Bot. Jahrb. 23:123. 1896. Villamilla polyandra Н. Walt. loc. cit. 37: Beibl. 83:24. 1906. Weak shrubs up to 5 meters, or woody vines climbing on shrubs. Leaf blades elliptic to ovate, the apex acute to attenuate, the base rounded or acute, 7.0-16.8 cm. long, 3.5-6.6 cm. broad, the petioles 1.0-3.0 cm. long, glabrous. Inflorescence a long uncrowded raceme, about 8.0-21.0 cm. long; pedicels 8-16 mm. long; bracts lanceolate, 1.5-2.0 mm. long, deciduous, the bractlets about 0.5 mm. long. Tepals white in flower, red, purple or purplish-pink in fruit, 4.0-9.0 mm. long. Stamens 20-25 inserted at the base of the perianth. Ovary subglobose, the style very short. Fruit a drupe, purple ог red, 4.5-5.5 mm. in diameter. Central America (Nicaragua, Costa Rica and Panama). BOCAS DEL TORO: Pumpkin River near Chiriqui Lagoon, Von Wedel 2571; Water Valley, 2. Wedel 926, 1439, 714, 1547, 940; Woodson, Allen 8 Seibert 1832; Almirante, Cooper 133. PANAMA: Rio Juan Diaz above Juan Diaz, Allen 944. Trichostigma appears to have close affinities with Rivina. However the two may easily be distinguished by the more numerous stamens, the tepals reflexed in fruit, the terminal stigma, the deciduous stipules, and the drupaceous fruit of Trichostigma. The genera have been treated as one genus by some authors, but recently they have been regarded as separate. (421) (Мог. 48 80 ANNALS OF THE MISSOURI BOTANICAL GARDEN AIZOACEAE By LORIN I. NEVLING, Jr. Annual or perennial herbaceous or suffruticose plants, prostrate or upright, often succulent. Leaves usually simple, alternate, opposite or pseudoverticellate, often fleshy, sometimes reduced to scales, entire; stipules scarious or absent. In- florescences axillary or terminal, modified cymes or the flowers solitary. Flowers bisexual, polygamodioecious or unisexual, actinomorphic, often small. Perianth monochlamydeous, usually 4- or 5-parted, free or connate, sometimes appendaged on the outer surface, often persistent in fruit. Stamens (3), 4, 5 or many, the outermost often sterile and petaloid (but not in our species), the filaments free or variously connate at the base into fascicles or into a monadelphous sheath, free or adnate to the perianth, the anthers oblong or linear, small, dehiscing longi- tudinally. Pistil 1, the ovary superior to inferior, 1-5 (—20) loculate, the placenta- tion axile, parietal or basal, the ovules solitary to many per locule, anatropous or campylotropous, the styles as many as the locules. Fruit a loculicidal or circum- scissile capsule or indehiscent and either baccate or nut-like; seed with mealy endosperm, sometimes strophiolate, the embryo curved. A large weedy family particularly well-developed in South Africa. Three genera are represented in Panama, each by a single species. A fourth genus, Glinus, with a single species [G. radiatus (Ruiz & Pav.) Rohrb.] has been reported from Panama. I have been unable to locate a voucher for this record but have included the genus in the key to genera. This study is based primarily on the work of F. Pax and K. Hoffman (in Natur- Pflanzenf. 16°:179-233. 1934) and P. Wilson (in North Amer. Fl. 21:267-277. 1932) a. Leaves basal or pseudoverticellate, not fleshy; эр се to the base or nearly so, with or without appendages the outer surface ea beneath th х; ind a loculicidal енді бае to 5. seeds strophiolate or estrophio ate. b. 52% — leaves generally linear; flowers pedicellate; tepals distin the base, without appendages; seeds estrophiolat Е 1. Morruco bb. ees toment RR lea obovate, oblanceolate ог ed- spatulate; flowers sic Pai or · борова tepals et to the base or nearly so, with appendages; seeds strophiolate...............-....- GLINUS aa. Leaves opposite, fleshy; tepals conna te, with ap nicis on the outer surface beneath the apex; fruit a circ uio m capsule, 1- to 5- ate; te. c. Leaves ны то насаб gu strikingly unequal; ovary 1- or 2-loculat vides seeds few cc. S орун = рис or ane obovate, a or less equal; vary 3- to ве styles 3 to 5; seeds n rn! МИА ДУВАНА 3. SESUVIUM 2. TRIANTHEMA 1. МОШ СО L Morruco Г. [Gen. 356. 1737]; Sp. Pl. 89. 1753. Galiastrum Heist. ex Fabricius, Enum. Pl. Hort. Helmst. 108. 1759. Lam petia Raf. Fl. Tellur. 3:34. 1836. (422) 1961] FLORA OF PANAMA (Aizoaceae) 81 Fig. 132. Mollugo verticellata Herbaceous or suffruticose plants, glabrous throughout. Leaves simple, basal or pseudoverticellate, generally linear; stipules deciduous. Inflorescences axillary, cymose. Flowers bisexual, pedicellate. Perianth 5-parted, distinct to the base, imbricate, scarious at the margins, lacking appendages. Stamens 3-10, united at the base. Ovary superior, 3- to 5-loculate, the ovules campylotropous, numerous on the axile placenta. Capsule loculicidal; seeds generally numerous, small, estrophiolate. About 25 species in the temperate and tropical regions of both hemispheres. 1. MOLLUGO vERTICELLATA L. Sp. РІ. 89. 1753. Pharnaceum hoffmannseggianum Roem. & Schult. e f 692. 1820. Mollugo dichotoma Schrank, Pl. Rar. Hort. Monac. # Mollugo arenaria HBK. Nov. Gen. & Sp. 6:20. 1823. Mollugo парата; (Roem. & Schult.) Ser. in DC. Prod. 1:393. 1824. Mollugo schrankii Ser. loc. cit. 391. Pharnaceum verticellatum (L.) Spreng. Syst. 1:949. 1825. Pharnaceum arenarium (HBK.) Spreng. loc. cit. 1825. i i 715278 мыт іле Schlecht. ех Rohrb. loc. cit. 242. 1872, nom. nud. in syn. (423) [Vor. 48 82 ANNALS OF THE MISSOURI BOTANICAL GARDEN Annuals, prostrate to ascending, often falsely dichotomously branched. Leaves pseudoverticellate, 3-6 per node, unequal, generally oblong-linear, 1-3 cm. long, 1—2 mm. broad, more or less acute at the apex, gradually tapered to the base; sessile. Inflorescence axillary, modified cymose; pedicel 3-5 mm. long. Flowers with the tepals elliptic, about 2.0-2.5 mm. long, 0.5-0.75 mm. broad, persistent and subtending the fruit, glabrous; stamens 3, the filaments filiform, to 2 mm. long, connate at the very base, the anthers ovoid, about 0.25 mm. long and broad; ovary ovoid, 3-loculate, 1.0—1.5 mm. long, the styles 3, 0.5-0.75 mm. long, spreading. Capsule ovoid, 2-3 mm. long, about 1.5 mm. in diameter; seeds numerous, reni- form, about 0.5 mm. long and broad, the testa with several distinct ridges along the back and sides, brown. Cosmopolitan weeds in the temperate and tropical regions of the world. This species has a rather large and complex synonomy involving numerous infraspecific categories which are not included here in the synonymy. CANAL ZONE: Chagres, Fendler ІІ. PANAMA: San José Island, Jobnston 1232. 2. TRIANTHEMA L. TRIANTHEMA L. Sp. Pl. 223. 1753. Reme Adans. Fam. 2:245. 1763. Zaleya Burm. f. Fl. Ind. 110, Ё. 37. 1768. Papularia Forsk. Fl. Аевург. e 69. 1775. 2775: Portulacastrum Juss. ex Medi m Bot. 1:99. 1789. Zallia Roxb. Fl. Ind. 3:74. cers Fenzl, in je Wien Mus. 2:293. 1839. а Willd. ex Steud. Nom. ed. 2. 2:429. 1841. Zaleia Steud. loc. cit. 795. 1841. Pomatotbeca F. Müll. Fragm. 10:72. 1876. Herbaceous or suffruticose plants, branched, upright or prostrate. Leaves simple, opposite, strikingly unequal, fleshy, obovate to rounded-spatulate, the petioles connate into a petiolar sheath surrounding the stem; stipulate. Inflores- cences axillary, cymose or the flowers solitary. Flowers bisexual, sessile or pedicel- late. Perianth 5-parted, connate below, the lobes appendaged on the outer surface beneath the apex, imbricate. Stamens 5 or more, inserted on the perianth tube or free and monadelphic. Ovary superior, 1- or 2-loculate, the ovules campylo- tropous, few on the axile placenta, the styles 1 ог 2. Capsule circumscissile; seeds few, reniform, the testa often wrinkled, estrophiolate. About 15 species of the tropics and subtropics. 1. TRIANTHEMA PORTULACASTRUM L. Sp. Pl. 223. 1753. Trianthema monogynum L. Mant. 69. Trianthema procumbens Mill. Gard. Diet a 8. no 1768 нача monogynum (L.) Medic. Phil. Bot. "i 99. 1789. rianthema flexuosa Schum. & Thonn. Beskr. Guian. Pl. 241. 1828. (424) 1961] FLORA OF PANAMA (Aizoaceae) 83 Fig. 133. Triantbema portulacastrum Herbaceous annuals, erect or prostrate, the young stems sometimes with a decurrent line of hairs from the interpetiolar stipules to the node below and glabrescent, often alternately branched by the development of the axillary bud subtended by the smaller leaf of the nodal pair. Leaves of any pair strikingly unequal, the larger at least twice as large as the smaller, obovate to rounded- spatulate, 1-3 cm. long, 0.5-2.0 cm. broad, acute, obtuse, retuse and often mucronulate at the apex, cuneate at the base, glabrous above and below; petiole 0.3-1.5 cm. long, at the base connate into a sheath surrounding the stem, the interpetiolar stipule deltoid, 1-2 mm. long, remotely serrate or entire. Inflorescence with the flowers solitary; sessile. Flowers with the perianth tube campanulate, often intimately associated with the petiolar sheath of the subtending leaves, the lobes lanceolate, about 1.5 mm. long, 1 mm. broad, glabrous, appendage horn-like, barbed about 0.75 mm. long; stamens 10, inserted at the orifice of the perianth tube, the filaments filiform, about 1.5 mm. long, glabrous, the anthers ovoid, about 0.5 mm. long and broad; ovary turbinate, about 1 mm. long, 1 mm. in diameter, truncate and irregularly fleshy-lobed at the apex, glabrous, the style 1, about 1 mm. long. Capsule almost enclosed in the petiolar sheath, turbinate, about (425) [Vor. 48 84 ANNALS OF THE MISSOURI BOTANICAL GARDEN 4.5 mm. long, 3.5 mm. in diameter, sessile, truncate and crested at the apex, circumscissile at about the middle; seeds 2—5, reniform, 1.5—1.75 mm. in diameter, the testa wrinkled, reddish-brown to black. А pantropic species. CANAL ZONE: Fort Clayton, Standley 29003. 3. SESUVIUM L. SEsuviuM Г. Syst. ed. 10. 1058. 1759. Squibbia Raf. New Fl. 4:16. Diplocbonium fee Nov. Stirp. Dec. $5 Н. Pyxipoma Fenzl, in Ann. Wien Mus. 2:293. 1839. Psammantbe Hence, i in Walp. Ann. 2:659. 1852. Halimus O. Ktze. Rev. Gen. 1:263. Herbaceous or suffruticose plants, upright or prostrate, sparsely branched. Leaves simple, opposite, more or less equal, fleshy, generally linear, elliptic or narrowly obovate, the petioles clasping or connate and surrounding the stem; estipulate. Inflorescence axillary or rarely terminal, glomerules or the flowers solitary. Flowers bisexual, sessile or pedicellate. Perianth 5-parted, connate below, the lobes generally appendaged on the outer surface beneath the apex, imbricate. Stamens 5, free and alternitepalous, or numerous, and inserted on the perianth tube. Ovary superior, 3- to 5-loculate, the ovules campylotropous, many on the axile placenta, the styles 3 to 5. Capsule circumscissile; seeds numerous, reniform, the ta smooth, estrophiolate. Six to eight species, tropical and subtropical strand plants of both hemispheres. 1. SESUVIUM PORTULACASTRUM (L.) L. Syst. Nat. ed. 10. 1058. 1759. Portulaca portulacastrum L. Sp. Pl. 446. 1753. Sesuvium revolutifolium Ortega, Dec. 19. 1797. Sesuvium ortegae Spreng. Bot. Gart. Halle кезе 1:36. 1801. Sesuvium pedunculatum Pers. Syn. Pl. 2:39. Sesuvium sessile Pers. loc. cit. 1806. Sesuvium sessiliflorum Domb. ex Rohrb. in Mart. Ен Bras. 142:310. 1872, nom. nud. in syn. Halimus portulacastrum (L.) O. Ktze. Rev. Gen. 1:263. 1891. Herbaceous perennials, glabrous throughout, the branches trailing and often rooting at the nodes, sometimes rather stout. Leaves linear, elliptic or narrowly obovate, 1-5 cm. long, 0.2-1.0 cm. broad, subacute to acute at the apex, gradually tapered to the base; petiole 1-5 mm. long, dilated at the base and clasping the stem but the sheaths not connate. Inflorescence axillary, the flowers solitary, pink; pedicel 2-11 mm. long. Flowers with the perianth tube obconic to subcampanu- late, 1.5-3.0 mm. long, the lobes ovate, 4-7 mm. long, 3.0-4.5 mm. broad, some- what auriculate at the perianth tube orifice, persistent and somewhat reflexed in fruit, appendage horn-like, 1.0—1.5 mm. long; stamens numerous, inserted ас the orifice of the perianth tube, free or subconnate, the filaments 1.5—3.5 mm. long, gradually tapering to the apex, the anthers oblong, 0.5—0.75 mm. long, 0.25-0.5 mm. broad; ovary ovoid to subglobose, 3.0—3.5 mm. long, 2.5—3.0 mm. in diameter, (426) 1961] FLORA OF PANAMA (Portulacaceae) 85 Fig. 134. Sesuvium portulacastrum the styles 3 or 4, 1.5-3.5 mm. long spreading or erect. Capsule ovoid or obovoid, about 6.5 mm. long, 3 mm. in diameter, circumscissile below the middle; seeds 8-30, lenticular-reniform, about 1.0-1.5 mm. in diameter, the testa smooth, black. Circumtropical. BOCAS DEL ToRO: bar mouth, Changuinola Valley, Dunlap 139, Stork 139. CAN ZONE: Cristóbal, Artamanoff s. n.; Balboa, Standley 25625, 30882; railroad and dock yards 1 ity о tandley 31180. COLON: vicinity of Palenque, Pittier 4118. PANAMA: Pearl Islands, Johansen ІІІ; Saboga Island, Allen 2634, Miller 1905: near Punta Paitilla Military Reservation pent of Panama City, Bro. Maurice 754; San José Island, Erlamsom 173. sAN Bras: forests around Puerto Obaldia, Pittier a PORTULACACEAE By LORIN I. NEVLING, Jr. Herbs, subshrubs or shrubs, more or less succulent, often glabrous. Leaves alternate, opposite or in basal rosettes, simple, terete, subterete or plane, often fleshy, pinnately veined; stipules scarious, fimbriate, tufted or rarely absent. In- orescence terminal or axillary, paniculiform, racemose, cymose or the flowers solitary. Flowers bisexual, often insignificant. Sepals (involucral bracts) 2 or (427) ГУот. 48 86 ANNALS OF THE MISSOURI BOTANICAL GARDEN rarely 4—8, connate at the base or free, persistent or caducous, scarious or herba- ceous, imbricate. Petals (tepals) 4—5 or rarely fewer or more, connate at the base or free, deciduous or rarely calyptrate, imbricate. Stamens often as many as the petals and opposite them, sometimes fewer or more, free or basally adnate to the corolla, the filaments filiform, the anthers 2-celled, dehiscing longitudinally, introrse. Pistil 1; ovary superior to inferior, becoming uniloculate, the placentation central, basal, the ovules (1-) 2-тапу, campylotropous; styles and stigmas 1-9, united below or free. Fruit capsular, circumscissily or loculicidally dehiscent, rarely indehiscent and nut-like; seeds 1—many, generally reniform-round, com- pressed, the embryo curved, endosperm mealy. mily of weedy plants composed of 15-25 genera. Only 2 genera are represented in Panama. a. Leaves estipulate; ovary superior; styles 3; capsule loculicidal, 3-valved, splitting from the apex нь the base 1. TALINUM aa. Leaves stipulate or estipulate, the stipules scarious, fimbriate or tufted; ovary half to wholly inferior; styles 1-9; capsule circum- scissile 2. PORTULACA 1. TALINUM Adans. TALINUM Adans. Fam. 2:245. 1763. Heliantbemoides Medik. Phil. Bot. 1:95. 1789. Phemera о af. in Med. Repos. М. У. 5:350. 1808. Litanum Nivel in Amer тт, Nat. 4:90, 1915. Herbs, subshrubs or shrubs, often succulent. Leaves alternate or approximate, terete to plane, fleshy; estipulate. Inflorescence terminal or rarely axillary, paniculi- form, cymose or the flowers solitary. Sepals 2, opposite, deciduous or rarely persistent. Petals 5, rarely more, connate at the base or free, deciduous. Stamens 5-30, in antipetalous fascicles. Ovary superior, sessile or short-stipitate, the ovules numerous, the styles 3, more or less united, filiform. Capsule loculicidal, 3-valved, splitting from the apex to the base, chartaceous; seeds reniform-round, numerous, the embryo incompletely annular, the testa smooth, striate or tuberculate, distinctly or indistinctly strophiolate. A genus of about 50 species of which only 1 is found in Panama. 1. TALINUM PANICULATUM (Jacq.) Gaertn. Егист. 2:219. 1791. Portulaca paniculata Jacq. Enum. = Carib. 22. 1760. Portulaca patens L. Mant. 242. 1 Roelingia и (L.) xe T A 135. 1788. Talinum reflexum Cav. Іс. a (428) 1961] FLORA OF PANAMA (Portulacaceae) 87 Fig. 135. Talinum paniculatum Talinum moritziana Klotzsch, ex Rohrb. in Mart. Pi Bras. 142:295. 1872. Talinum roseum Klotzsch, ex Rohrb. loc. cit. Talinum patens var. sarmentosum (Englm.) lige in Proc. Amer. Acad, 22:275. 1887. Claytonia patens (L.) O. Ktze. Rev. Gen. 1:56. 1891. т paniculata (Jacq.) O. Ktze. loc. cit. 57. 1891. Claytonia reflexa (Cav.) O. Ktze. loc. cit. 1891. Clin sarmentosa (Englm.) O. Ktze. loc. cit. Talinum reflexum £. sarmentosum (Englm.) Small, EL. "SE. U. S. 415. 1903. Talinum chrysanthum Rose & Standl. in Contr. 1). S. Nat. Herb. 13:288. 1911. Talinum paniculatum var. sarmentosum (Englm.) v. Poellnitz, in Deut. Bot. Gessel. Berichte 54:123. 1953, Herbaceous plants, erect, slender, glabrous throughout, the root often thick and fleshy. Leaves alternate, broadly elliptic to obovoid, rarely oblanceolate, 4—13 cm. long, 1.5-5.5 cm. broad, acute to obtuse at the apex, cuneate or attenuate at the base, plane, petiole 3-15 mm. long. Inflorescence terminal, compound, lax, 10—60 cm. long, 6-20 cm. broad, the lateral simple or compound cymes racemosely arranged on the primary rhachis, bracteate; pedicel 6-25 mm. long, dilated distally. Sepals ovate, 1.5—3.0 mm. long and broad, connate at the base, slightly keeled, reflexing and deciduous; petals elliptic, 3.0—4.5 mm. long, about 1 mm. broad; stamens 15 or more, the filaments 1.5—2.0 mm. long, free, the anthers ovoid, to 0.5 mm. long and broad; ovary sessile, globose, about 1 mm. long, 1 mm. in diameter. Capsule ovoid to globose, 3-5 mm. long, 3-5 mm. in diameter; seeds (429) (Мог. 48 88 ANNALS OF THE MISSOURI BOTANICAL GARDEN strongly reniform, 0.75—1.0 mm. in diameter, the testa minutely striate, black, indistinctly strophiolate. Southern United States, Central and South America, West Indies, southern Asia and the East Indies. PANAMA: Isla Taboga, Woodson, Allen & Seibert 1491. 2. PORTULACA L. PonTULACA І. [Syst. ed. 1. 1735]; Sp. Pl. 445. 1753. Meridiana L. #. Suppl. 248. 1781 Portulacca Haw. Syn Lamia Vandelli ex Endl. бе. ыы. 1840. Annual ог perennial herbs, succulent. Leaves alternate ог approximate, terete to plane, often whorled about the flowers; stipules scarious, fimbriate or tufted, sometimes very small, rarely absent. Inflorescence near the stem apex, crowded, the flowers sometimes solitary, yellow to red. Sepals 2, opposite, connate below, the anterior larger than the posterior, usually persistent. Petals 4—6, connate at the base or free, somewhat auto-deliquescent. Stamens 4 to many, inserted at the base of the petals, the filaments often hairy below. Ovary half to wholly inferior, the ovules numerous, the styles 3- to 9-parted, rarely simple. Capsule circumscissile, chartaceous; seeds reniform to cochleate, numerous, the testa smooth or minutely tuberculate. A cosmopolitan genus in tropical and subtropical areas of the world, about 125 species, 1. PoRTULACA OLERACEA L. Sp. Pl. 445. 1753. Portulaca parvifolia Haw. Syn. Pl. Succ. 122. 1812. Portulaca marginata HBK. Nov. Gen. & Sp. 6:58. 1823. Portulaca oleracea B parviflora (Haw.) Griseb. Fl. Brit. W. дыр: 57. 1864. Portulaca oleracea а macrantha Eggers, Fl. St. Croix 27. 18 Portulaca oleracea В micrantha Eggers, loc. cit. 1879 Plants herbaceous, radially spreading are prostrate or somewhat ascending, glabrous throughout, the roots fibrous. Leaves alternate, elliptic to obovoid, 1-3 ста. long, 0.5—1.0 cm. broad, generally obtuse at the apex, cuneate at the Базе, plane; petiole 1-8 mm. long, the stipules fimbriate, inconspicuous. Inflorescence with the flowers crowded, the flowers sessile or essentially so, yellow. Perianth tube crateriform, 1 mm. long, 1.5 mm. in diameter; sepals ovate, 3.0-4.5 mm. long, 3-4 mm. broad, strongly keeled, connate at the base, generally persistent and enclosing the operculum of the fruit; petals 4, 3.0-4.5 mm. long, 1.5-3.0 mm. road; stamens 6-15, the filaments 1.5—1.75 mm. long, the anthers globose, to 0.5 mm. long and broad; ovary half-inferior, short-conical, the style lobes 4—6. Capsule ovoid to fusiform, 6-9 mm. long, about 2.5 mm. in diameter, circumscissile (430) 1961] FLORA OF PANAMA (Portulacaceae) 89 slightly below the middle; seeds almost cochleate, about 0.5—0.75 mm. in diameter, the testa minutely tuberculate, almost black. Known as verdolaga in Panama. According to Standley (Field Mus. Nat. Hist. Bot. 18:429. 1937) the plant is used as a pot herb throughout Central America. BOCAS DEL TORO: vicinity of Nievecita, Woodson, Allen & Seibert 1862. cumiquí: Boquete, Davidson 705. DARIEN: vicinity of Boca de Cupe, Allen 875; Río Chucunaque, Leopold 758. PANAMA: San José Island, Johnston 001. Fig. 136. Portulaca oleracea (431) [Vor. 48 90 ANNALS OF THE MISSOURI BOTANICAL GARDEN CARYOPHYLLACEAE By JAMES A. DUKE Annual or perennial herbs or subshrubs, often with swollen nodes. Leaves opposite and decussate or verticillate, the petioles often amplexicaul, some with conspicuous stipules. Flowers commonly bisexual, actinomorphic, 5- or 4-merous, in dichasial cymes, or solitary in the axils, or solitary and terminal (Githago). Sepals 5—4, separate or connate, often scarious, persistent. Petals as many as the sepals, rarely absent, hypogynous or slightly perigynous, entire, bifid or lacerate, usually white, pink or red. Stamens 2-10, mostly 5 ог 10; filaments filiform or flattened, occasionally united into a short tube below; anthers usually versatile, 2-celled, and longitudinally dehiscent. Carpels 2-5, the ovary 1-locular, with 1- many campylotropous ovules on basal, central or free central placentae, styles 2-5, free or united Fruit a utricle or a capsule longitudinally dehiscing into as many entire or deeply emarginate valves as there are styles; seeds 1—many, strophiolate or estrophiolate, smooth or tuberculate, the embryo curved about the perisperm. This family, with about 75 genera, mostly in temperate areas, is of little economic interest except for a number of ornamentals, such as Lychnis and Dianthus, which may ultimately be found in Panama gardens. Several weedy genera, of more or less cosmopolitan distribution, despairingly need monographic research, worldwide in scope. Five such weedy genera occur in Panama. a. Leaves longer than broad; stipules absent; stamens usually 10, rarely 4 ог 5; capsule splitting into 2-5 usually 2-cleft valves; styles not united. b. Leaves linear to elliptic; petals entire or shallowly emarginate............. 1. ARENARIA bb. Leaves ovate to cordate; petals dee ifid c. Leaves dpi capsules cylindric, arcuate, with 10 teeth; styles usuall cc. Leaves ovate or Voir capsules ovoid, straight, with 6-8 (—10) teeth; чэк s usually aa. Leaves shorter or ap dus broad; stipules present, occasionally fugaceous; stamen psules splitting into 3 entire valves; styles usually united at i hi КН p nee length. d. Leaves ped be long, opposite, the ко» stipules often fugaceous; seeds 1-50, tuberculate in lines (in Рапата)...................... 4. DRYMARIA dd. Leaves Mine than cee verticillate, the argentate stipules per- sistent; seeds 2—6, merely corrugated ~ . CERASTIUM . STELLARIA w 5. PoLYCARPAEA 1. ARENARIA L. ARENARIA І. [Rupp. ex І. Syst. ed. 1. 1735] Sp. Pl. 423. 1753. Moebringia L. [Syst. "* 2: a 1740] Sp. Pl. 359. 1753. Minuartia L. Sp. Pl. 8 Gypsopbytum А Tui s. 256. 17 Ammodenia Patrin, ex Gmel. Fl. ‘ibis, * 160. 1769. Alsine Scop. Fl. Carn. ed. 1:496. 1772. not L. 1753. Honkenya Ehrh. Beitr. 2:180. 178 Leptophyllum Ehrh. loc. cit. 4:147. 1789. (432) 1961) FLORA OF PANAMA (Caryophyllaceae) 91 Spergulastrum Michx. Fl. Bor. Am. 1:275. set in part. Gouffeia Robill. & Cast. in Lam. & DC. Fl. Fr T 609. 1815. irse Fries, Fl. Holland 75. 18 17. Adenarium Raf. in Desv. Jour. Phys. 89:249. 1818. Bigelovia Raf. in Jour. Phys. 89:289. 1819. Merckia Reichb. ax Cham. & SE in чел :59. 1826. Bracbystemma D. Don, Prodr. Fl. Мер. 2 Plinthine Reichb, lo loc. cit. к. 298. 1837, Stropbium Dulac. Fl. Hautes-Pyr. 247. 1867. Xeralsine Fourr. in Ann. Soc. Linn. п. 5. 16:347. 1868. tn 55 SE ЗЕ & x >5 is ezl T c 6 > wn кә > 22; Bre A. Gray, in Proc. Am. Acad. 8:620. 1869. Alienate Salt Fl. S. Е. Ц. 5. 419 & 1330. 1903. Annual, biennial or perennial herbs, glabrous or pubescent, prostrate and cespitose to diffuse and spreading, the stems somewhat resilient and occasionally suffruticose below. Leaves opposite, petiolate to sessile and slightly amplexicaul, exstipulate, subulate and coriaceous to broadly elliptic and membranaceous. Flowers axillary or terminal, solitary or in dichasial cymes. Sepals 5, connate only at the base if at all. Petals 5, rarely lacking, white or occasionally red, entire or shallowly emarginate. Stamens 10, rarely 5; anthers versatile; 2-celled, longitudinally dehis- cent; filaments flattened and usually united below to form a slightly perigynous disk. Ovary superior, 3- (2-5-) carpellate with 3 (2-5) filiform styles free to their bases; ovules numerous on basal placentae. Capsule dehiscing into as many entire to deeply emarginate valves as there are styles; seeds numerous, cochleate, smooth to tuberculate, estrophiolate, the embryo curved about the perisperm. A weedy genus containing nearly 200 species, this nearly cosmopolitan genus is in the tropics largely restricted to high altitudes. Fernald (in Rhodora 21:1. 1919) presents adequate reasons for retaining Arenaria sensu lato instead of main- taining several poorly separated generic segregates. Three intergrading units, best treated as subspecies of one polymorphic species, A. lanuginosa, are found in anama. (433) ГУог. 48 92 ANNALS OF THE MISSOURI BOTANICAL GARDEN a. puse oblong, lanceolate or e the gd les than 20 mm. long pals тен or pubesc in lines, mm tion the petals e Se sules longer or s авын атра маа ог diffus b. Plants diffuse, с spreading; sepals a dt 2-4 mm. long, the petals 2-4.5 . long; Ben. vid much longer than be. Pipe ta: leaves; leaf Лали usually р 1. ssp. LANUGINOSA bb. Plants cespitose, much bra o near the base, some of the branches occasionally ideis pony labro ous, 3-5 mm. long, the petals 4—6 mm. long; cels shorter or longer than the subtending leaves; af margins n volute 2. ssp. SAXOSA aa. Leaves ae e = о over 20 mm. long; sepals conspicuously pubescent on the с Међу m. long, the petals and capsules longer; plants diffuse and жаныл 3. ssp. GUATEMALENSIS 1. ARENARIA LANUGINOSA ssp. LANUGINOSA Maguire, in Am. Midl. Nat. 48:498. 193]. Spergulastrum lanuginosum Michx. Fl. Am. Bor. 1:275. 1803. Micropetalum lanuginosum (Michx.) Pers. Syn. Pl. 1:509 Arenaria alsinoides Willd. ex Schlecht. in Е Gesells. Nat. Freund. Berl. 7:201. 11816. Stellaria elongata Nutt. Gen. 1:289. Are enaria өндіре Ell. Sketch. Bot. S. С. с = 1:539. 1821. 17 К. Nov. Gen. & Sp. 6:35. 1823. Arenaria 1 nemorosa а quitensis DC. Prodr. 1:409. 1824. renaria nemorosa B novogranatensis DC. loc. cit. 1:409. 1824. renaria jussiaei Camb. ex St. Hilaire, Fl. Bras. Mer. 2:126. 1829. Arenaria scabra Vahl, ex St. Hilaire, loc. cit. 2:126. 1829 Arenaria baradoxa Bart. Rel. Haenk. 2:15. Stellaria lanuginosa (Michx. ) Torr. & sii Fl. Amer. Bor. 1:187. 1840. Arenaria lanuginosa a genuina Rohr Saee = V im 1872. Arennaria lanuginosa B diffusa Rohrb. bs 872. Arenaria lanuginosa diffusa (ЕП.) Macloskie, Rep. нА Univ. Exped. Patag. 8:394. 1905. Stellaria laxa Muschl. in Bot. Jahrb. 45:443. 1911. Arenaria lanuginosa var. longipedunculata Duncan, in Phytologia 3:282. 1950. Herbaceous puberulent laxly spreading perennials. Leaves opposite, lanceolate, oblanceolate or narrowly elliptic, apically attenuate and mucronulate, basally sub- sessile and attenuate, 5—20 mm. long, 2-5 mm. broad, scantily to densely puberulent with a whitish indumentum, often ciliate on the margins. Flowers solitary in the axils, the puberulent pedicels much longer than the subtending leaves. Sepals lanceolate to ovate, attenuate, puberulent 2-4,5 mm. long; petals 5, occasionally absent, obovate, entire, 2-4.5 mm. long; stamens 10, the filaments 1-4 mm. long; ovary sessile or short-stipitate, ovoid; styles 2-4. Capsule ovoid, 3-5 mm. long, the 2-4 valves deeply emarginate; seeds numerous, on basal placentae, cochleate, dark reddish brown to black, smooth or minutely tuberculate, ca. 0.75 mm. broad. Southeastern U. S. to Peru and Bolivia, in Central America only at moderate elevations. CHIRIQUÍ: vicinity of "New а iss -— valley of Rio Chiriqui = 1800- 2000 m., Allen 1414; vicinity of Boquete, 1 0 m., Woodson & Schery 802 The typical species was originally ный as apetalous, as are most specimens from the southeastern United States, the type locality. In Mexico and Central America however, where the ranges of several subspecies overlap, the large majority (434) 1961) FLORA OF PANAMA (Caryophyllaceae) 93 of specimens, obviously referable to this "apetalous" subspecies, have petals nearly or quite as long as the sepals. MacBride (in Field Mus. Bot. 13°:601. 1936-8) reports that in Peru, where the plant is called celedonia and teuchchalli, it is the source of an astringent used for hemorrhages of the uterus. 2. ARENARIA LANUGINOSA ssp. SAXOSA (A. Gray) Maguire, in Am. Midl. Nat. 48: 498. 1951. Arenaria saxosa A. Gray, Pl. Wright. 11: 22 1853. Arenaria saxosa var. cinerascens Robinson, in Proc. Am. Acad. 29:293. 1894. Arenaria confusa Rydb. in Bull. Torr. Club 28:275. 1901. Arenaria polycaulos Rydb. loc. cit. 31:406. Arenaria mearnsii Woot. & Standl. in Contr. U. s. Nat. Herb. 16:121. 9; Arenaria saxosa var. mearnsii (Woot. & Standl.) Kearney, & Peebles, in Jour. Wash. Acad. 9:475. 1939. Herbaceous (occasionally subligneous at the base) puberulent cespitose perennials, some of the branches occasionally proliferating and spreading. Leaves opposite, mostly linear-lanceolate, apically attenuate and mucronulate, basally subsessile and rounded to acute, 5-18 mm. long, 1-3 mm. broad, scantily to densely puberulent with a whitish indumentum, often ciliate on the involute margins. Flowers solitary in the axils, the puberulent pedicels longer or occasion- ally shorter than the subtending leaves. Sepals lanceolate to ovate, attenuate, glabrous, 3—5 mm. long; petals 5, obovate, entire, 4—6 mm. long; stamens 10, the filaments 1-4 mm. long; ovary sessile or short-stipitate, ovoid; styles 2-4. Capsule ovoid, 3-5 mm. long, the 2-4 valves deeply emarginate; seeds numerous, on basal placentae, cochleate, dark reddish brown, usually tuberculate, ca. 0.75 mm. broad. Southwestern U. S. to Mexico; Panama to northwestern South America, at rather high elevations. cHIRIQUÍ: Potrero Muleto, Volcan Chiriqui, Boquete District, 10,400 ft., Davidson 1041 & ш; Potrero Muleto to summit, Volcan Chiriqui, 3500-4000 m., Woodson & Schery 405 9 434; Loma Larga to summit, Volcan Chitiqui, 2500-3380 m., Woodson, ‘Allen & Seibert 1073. 3. ARENARIA LANUGINOSA ssp. guatemalensis (Standl. & Steyerm.) J. Duke comb. & stat. nov. Arenaria guatemalensis Standl. & Steyerm. in Field Mus. Bot. 23:50. 1944. Herbaceous puberulent laxly spreading perennials. Leaves opposite, lanceolate, apically attenuate and mucronulate, basally subsessile and attenuate, 10—40 mm. long, 2-6 mm. broad, irregularly puberulent with a whitish indumentum, often ciliate on the margins. Flowers solitary in the axils, the puberulent ebracteate pedicels 10—40 mm. long, usually exceeding the subtending leaves. Sepals lanceolate to ovate, attenuate, strongly puberulent on the costa, 4-6 mm. long; petals 5, obovate, entire, 5-8 mm. long; stamens 10, the filaments 3—6 mm. long; ovary sessile or short-stipitate, globose or ovoid; styles 2-4. Capsule ovoid, 5-8 mm long, the 3 valves deeply emarginate; seeds numerous, on basal placentae, cochleate, (435) [Vor. 48 94 ANNALS OF THE MISSOURI BOTANICAL GARDEN Ж 2223 ФР i Ў 2 MEN 24 kl 5% к= № Re 272 YL Fig. 137. Arenaria lanuginosa ssp. guatemalensis dark reddish brown to black, smooth or minutely tuberculate, ca. 0.75 mm. broad. Mexico to Panama, at moderate elevations. CHIRIQUÍ: rain forest, Bajo Chorro, Boquete гы 6000 ft., Davidson 256; vicinity of Finca Lérida, 1750 m., Woodson & Schery 214; vicinity of Casita Alta, Volcan Chiri- qui, ca. 1500-2000 m., Woodson, Allen & Seibert Sto, valley of the upper Rio Chiriqui Viejo, vicinity of Monte Lirio, 1300-1900 m., Seibert 297; in grassy sunny places, vicinity of Cerro Punta, 1500-2000 m., Seibert 260. Other variants of the lanuginosa complex are found in Mexico and Central America and could conceivably occur in Panama. In the subspecies guatemalensis, the variety ensifolia, with linear leaves and exerted petals, is so far reported only from southern Mexico. Characterized by ovate leaves, cuneate to cordate leaf bases and exerted petals, the variant currently passing as A. megalantha (Rohrb.) (436) 1961] FLORA OF PANAMA (Caryophyllaceae) 95 Е. М. Williams would probably better be treated as a lamuginosa subspecies, ranging from southern Mexico to Guatemala. The original description of A. guatemalensis would have been more edifying had it been contrasted with A. lanuginosa instead of A. megalantha, for although intergrades occur between all three, they are more frequent and perplexing between the first two. The relative constancy of these three taxa, where their ranges are distinct, coupled with limited intergradations near the commissures of their ranges, seems to justify their relegation to the subspecific level, in accord with the heirarchy established by Maguire (in Am. idl. Nat. 46:493. 1951) in his monograph of the North American species of Arenaria. 2. CERASTIUM Г. CERASTIUM L. Sp. Pl. 437. 1753. Prevotia Adans. Fam. 2:256. 1763. Centunculus Adans. loc. cit. 2:256. 1763. Pentaple Reichb. Ic. Fl. Germ. 5: 37. f 227; СЕ Dichodon Bartl. ex Reichb. Nom. 205. 184 Leucodonium Opiz, Seznam 59. 1852 Annual or perennial, usually pubescent, erect or decumbent herbs. Leaves opposite, usually sessile or subsessile, exstipulate, often viscid. Flowers few in dichasial cymes, white. Sepals 5, rarely 4, not connate. Petals as many as the sepals, rarely absent, emarginate or 2-cleft, white. Stamens 10, occasionally less; anthers versatile, 2-celled, longitudinally dehiscent; filaments flattened, scarcely if at all connate to form an obscurely glandular annulus. Ovary superior; carpels 5, rarely fewer, with as many distinct styles, the many campylotropous ovules arising from basal or central placentae. Capsule often arcuate (hence the generic name), dehiscing longitudinally into as many deeply emarginate valves as there are styles; seeds numerous, cochleate, usually tuberculate, estrophiolate, the embryo coiled about the perisperm. Of this weedy cosmopolitan, temperate genus, only one of about fifty species occurs in Panama. 1. CERASTIUM viscosuM L. Sp. РІ. 437. 1753. Cerastium glomeratum Thuill. Fl. chor 225. 1799. Cerastium consanguineum Wedd. in Ann. Sci. Nat. 9*: Cerastium viscosum B consanguineum » (Wedd. ) Rebeb. in жұ талан 37:284. 1872. Herbaceous viscid-villous freely branching annuals, prostrate, spreading or erect. Leaves opposite, viscid-villous, elliptic to ovate or obovate, apically rounded (437) ГУог. 48 96 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 138. Cerastium viscosum and usually mucronulate, basally attenuate, 5-25 mm. long, 3-15 mm. broad, the winged petioles less than 5 mm. long. Flowers few in terminal dichasial cymes with reduced foliaceous bracts at the major dichotomies, the lanate to villous pedicels becoming 5-10 mm. long, exceeding the subtending bracts. Sepals 5, ovate to lance-oblong, acute to attenuate, villous, 3-5 mm. long; petals 5, rarely none, oblong, deeply 2-cleft, 3-5.5 mm. long; stamens 10, the flattened filaments 2-4 mm. long; ovary sessile or short-stipitate, ovoid; styles usually 5, 1.5-2 mm. long, glandular throughout; ovules many, campylotropous on central placentae. Capsule subcylindric, arcuate, 6-8 mm. long, the 5 valves deeply 2-cleft; seeds numerous, cochleate, light brown, minutely tuberculate, ca. 1 mm. broad. Naturalized from Europe, now rather common throughout temperate and tropical America. 1051: Potrero Muleto to summit, Volcán Chiriquí, 3500-4000 m., Woodson & > Зар pus vicinity of Boquete, 1200-1500 m., Woodson 9 Schery 804. 3. STELLARIA L. STELLARIA L. Sp. Pl. 421. 1753. Myosoton Moench. Meth. 225. 17 Spergulastrum Michx. Fl. Am. Bor. % 275. 1803. In part. (438) 1961] FLORA OF PANAMA (Caryophyllaceae) 97 Micropetalon Pers. Syn. 1:509. 1805. Alsinella Swartz, Summa Veg. Scand. 17. 1814. ед PE Larbrea St. Hilaire, in Mém. Mus. Par. 2:287. 18 a Adenonema Bunge, in Mem. Sav. Etr. Petersb. 2: 448. an Schisbfacbivm Reichb. Nom. 205. 1841. Hylebia Fourr. in Ann. Soc. Linn. Lyon n. s. 16:347. 1868. Myosantbus Four i bc cit. n. s. 16:348. 1868. Annual or perennial herbs, often much branched and clambering, the stems somewhat resilient and occasionally suffruticose below. Leaves opposite, petiolate or sessile and somewhat perfoliate, exstipulate, in Central America rather broad and membranaceous. Flowers in few-flowered dichasial cymes or less frequently solitary in the axils of slightly reduced leaves. Sepals 4—5, not connate. Petals 4—5, rarely absent, white, deeply 2-cleft. Stamens 4—10; anthers versatile, 2-celled, the flattened hypogynous filaments connate below forming a brief glandular disk. Ovary superior, carpels 2—4, with as many filiform styles free to the base; ovules 4—many, campylotropous on basal, central or free central placentae. Capsule deeply or shallowly dehiscing into as many 2-cleft or rarely entire valves as there are styles; seeds 4—many, cochleate, smooth or tuberculate, estrophiolate, the embryo curved about the perisperm. Composed of about a hundred species, this weedy genus is nearly cosmopolitan, but is, as are most caryophyllaceous genera, largely restricted to high altitudes in the tropics. Three of the six Central American species are found in Panama. a. к solitary in the axils or in few-flowered leafy cymes; sepals petals 5; stamens 8-10; seeds numerous; pedicels not glandular- тна: leaves ovate ог cordate. b. Leaves ај petals 8-10 mm. long, 1.5-2 am as long as the sepals; sp: the m отт Мй аз . 5. NEMORUM bb. rata binc petals m. long, 1—1.5 times as "ya as the sepals; the seeds ай ten k broad 2. S. OVATA aa. ная rs many in dichasial cymes with minute bracts; sepals and petals 4; stamens 4-6 (-8); seeds 4-6; pedicels glandular-villous, = cordate. 3. 5. IRAZUENSIS 1. STELLARIA NEMORUM L. Sp. Pl. 421. 1753. Cerastium nemorum Crantz, Inst. 2:401. 1766. Stellaria nemoralis nn. Prodr. 301. 1796. tellaria cuspidata Willd. ex Schlecht. in Ges. Natur. Freund. Berlin Mag. 7:196. 1816. Pi ile reichenbachii Мы. in Reichb. Ic. Fl. Germ. 5:34. 1841. ia baldwini F зіп паеа 30:57. -60. Stellaria ciliata ta В cuspidata (Willd. ) Rohrb. in а 37:278. 1872. var. japonica Franch. & Sav. Enum. Pl. Jap. 21:295. 1876. nives montana Pierrat, in Soc. Bot. Roche AE 1880. Stellaria nemorum subsp. glocbidosperma Murb. in Act. Univ. Lunds. 27:156. 1892. St tellaria nemorum к — 65 Миф. Bot. ides 201. 1899. ~ orma reichenbachii (Wierzb.) Murb. loc. cit. 201. 1899. < ‹ % ет ellaria subsp. Stellaria у и в. a, NE gor Tokyo 43: ат 1929. Stellaria limitanea Standl. in Field Mus. Bot. 22:74. 1940. Herbaceous glabrous to pubescent у branching perennials, prostrate, (439) [Vor. 48 98 ANNALS OF THE MISSOURI BOTANICAL GARDEN spreading or clambering. Leaves opposite, glabrous to viscid-villous, ovate, apically acute to attenuate, basally truncate to cordate, 10-50 mm. long, 5-35 mm. broad, all except the uppermost petiolate, the petioles 5-40 mm. long. Flowers solitary, axillary or terminal, or in terminal few-flowered cymes, the glabrous to villous pedicels 5—45 mm. long, often exceeding the subtending leaves. Sepals 5, lanceolate to ovate, acute to attenuate, glabrous to villous, 4-6 mm. long; petals 5, obovoid, deeply 2-cleft, 6-12 mm. long; stamens 10, the filaments flattened, 3-8 mm. long; ovary sessile, globose to ovoid; styles usually 3, 2-3 mm. long, the numerous ovules campylotropous on basal or free central placentae. Capsule ovoid, 4-7 mm. long, the 3 valves deeply 2-cleft; seeds numerous, ca. 1 mm. broad, cochleate, dark reddish brown, tuberculate, the spines usually broader than long. In the New World, extending from Mexico to Patagonia. $ DEL TORO: Robalo trail, northern slopes of Cerro Horqueta, 6000-7000 ft., XA ipao cHIRIQUÍ: Potrero He to summit, Volcán Chiriqui, 3500—4000 m., Woodson & Scbery 393. The New World material seems indistinguishable from the Old World material except that seeds tend to be more uniformly tuberculate and leaves do not tend to be so large in New World specimens. Phytogeographically it might seem that New World specimens might at least deserve subspecific rank, but I am unable to detect any constant differentia therein. The non-papillate tubercles on the seeds of Panama representatives more closely resemble those of the European subspecies montana (Pierrat) Murb. than those of the subspecies glochidosperma Murb. 2. STELLARIA ОУАТА Willd. ex Schlecht. in Ges. Naturf. Freund. Berl. Mag. 7:196. Herbaceous mostly glabrous, branching perennials, prostrate or spreading. Leaves opposite, glabrous or subglabrous, ovate to broadly ovate, apically rounded and mucronulate, basally rounded but slightly attenuate, 10-40 mm. long, 5-30 mm. broad, the petioles 3-8 mm. long. Flowers solitary in the axils, the glabrous to villous pedicels 10—40 mm. long, usually exceeding the subtending leaves. Sepals 5, ovate to obovate, obtuse to acute, glabrous to villous, 2.5-5 mm. long; petals 5, obovate, deeply 2-cleft, 3-6 mm. long; stamens 10, the flattened filaments 3—4.5 mm. long; ovary sessile, ovoid; styles usually 3, 1-1.5 mm. long, the numerous ovules campylotropous on basal placentae. Capsule ovoid, 3-5 mm. long, the 3 valves deeply 2-cleft, occasionally entire; seeds numerous, ca. 1 mm. broad, cochleate, dark reddish brown, tuberculate, the mature spines usually longer than road. Mexico through Central America to northern and western South America. HiRIQUÍ: valley of the upper Río Chiriquí Viejo, 1300-1900 m., White 8 White 3; nis of the upper Río Chiriquí, vicinity of Monte wa 1300-1900 m., Seibert 275; Palo Alto, just e. of Boquete, 5000 ft., Stern, Chambers et Standley and Steyermark (in Field Mus. den 24*:238. 1946) report the following vernacular names are applied in Guatemala: /ripa de pollo, culantro de monte and cuartillera. (440) _——-——-—— in 19611 Fig. 139. Stellasia irazuensis (441) 99 (Мог. 48 100 ANNALS OF THE MISSOURI BOTANICAL GARDEN 3. STELLARIA IRAZUENSIS Donn. Smith, in Bot. Gaz. 25:236. 1897. Herbaceous, mostly glabrous, much branching perennials, prostrate or spread- ing. Leaves glabrous, ovate to deltoid, apically attenuate, basally cordate, 10-20 mm. long, 5-15 mm. broad, the scantily villous petioles 5-15 mm. long. Flowers several in dichotomously branching cymes 10-25 cm. long, with minute bracts at the major dichotomies, the glandular-villous pedicels 5-10 mm. long. Sepals 4, narrowly ovate to elliptic, obtuse to acute, glabrous or glabrate, 1.5-2.5 mm. long; petals 4, deeply 2-cleft, the segments linear, 1.5—3.0 mm. long; stamens 4—6 (-8), the flattened filaments 1.5-2.5 mm. long; ovary sessile, ellipsoid; styles usually 2, ca. 1 mm. long, the 4—6 ovules campylotropous on basal placentae. Capsule flattened, ellipsoid, ca. 2 mm. long, the 2 valves emarginate; seeds 6 or less, ca. 1 mm. broad, cochleate, dark reddish brown, tuberculate. Guatemala to Panama. cHIRIQUÍ: vicinity of Bajo Chorro, 1900 m., Woodson & Schery 640. Further study of this distinctive tetramerous species may well prove it to be conspecific with 8. venezuelana Steyerm., which Steyermark rightly separated from the heterogeneous assemblage described as S. micrantha Spruce ex Rohrb. Rohr- bach’s description is not at all applicable to Fendler 47 which was designated as cotype of S. micrantba, and which is quite apparently referable to S. venezuelana. 4. DRYMARIA Willd. ex Roem. & Schult. Drymaria Willd. ex Roem. & Schult. Syst. 5:31. 1819. Pinosia Urban, in Arkiv dor 2345:70. pl. 2. 19 Mollugopbytum M. E. Jones, in Extr. Contr. West. "Вог. 18:35. 1933. Annual ог рту glabrous to pubescent herbs, occasionally subligneous below, prostrate, spreading or erect. Leaves opposite, with persistent or fugaceous small stipules, sessile or petiolate, glabrous to villose, the hairs often glandular. Flowers in few-flowered racemes or in dichasial cymes, rarely solitary in the axils. Sepals 5, not connate. Petals (0-) 5, white, usually 2-cleft, with the sinus of the cleft occasionally laciniate. Stamens 2-5, the anthers versatile, 2-celled, the flat- tened filaments slightly connate at the base, rarely with prominent staminodia. Ovary superior, slightly stipitate; carpels 3, the 3 styles united below; ovules few—many, campylotropous on free central placentae. Capsule ovoid to spheroid, dehiscing into 3 entire valves; seeds 1-тапу, cochleate, foetiform or hippocrepiform, usually tuberculate, the embryo curved about the perisperm. Exceptional for the Caryophyllaceae in being an almost exclusively tropical genus, Drymaria is represented in Panama by only two species. D. glandulosa Presl. and D. palustris, both ranging from North to South America, have not yet been collected in Panama. The genus consists of perhaps sixty species, all but three confined to tropical and subtropical America. А peculiar group of species with foetiform seeds centers about D. holosteoides Benth. which is a seriously toxic range plant of the Sonoran Desert (Little, in Ecology 18:416. 1937). In the only (442) 1961) FLORA OF PANAMA (Caryophyllaceae) 101 existing revisionary treatment of the genus, Wiggins (in Proc. Calif. Acad. Sci. 425:189. 1944), dealing with species on and near the Sonoran Desert, recognizes that the seeds are distinctive but lays little emphasis thereon. In my soon forth- coming revision of the whole genus, the seeds are one of the most important diagnostic characters, and the petal and leaf shapes are next respectively in impor- tance to an understanding of the relationships that exist in the genus a. Leave 4 pedicels — = densely villose with long, spreading non-g гү ве йч мадар. чат of the petals acute to emarginate, 2—4-nerved, еб prod Y E г auricles; seeds 0.5—0.9 mm. broad, with 1 ste. tella te tubercl aa. ип шеше or puberulent; pedicels locally girdled with dense bands of glandular s lobes of the petals acute, 1-nerved, basally exauriculate; seeds 1 mm. broad, with domical tubercles............... . D. VILLOSA 2. D. coRDATA 1. DryMaria VILLOSA Cham. & Schlecht. in Linnaea 5:232. 1830. Drymaria birsuta Bartl. in Presl, Rel. Haenk. 2:8. 1831. Drymaria cubensis Regel, in Otto & Dietr. Allg. Garten. 8:298. 1840. Fig. 140. Drymaria villosa (443) [Vor. 48 102 ANNALS OF THE MISSOURI BOTANICAL GARDEN Drymaria cordata var. pilosa Schlecht. in Linnaea 26:374. 1853. Drymaria cordata var. 9 villosa (Cham. & Schlecht.) Rohrb. in Mart. Fl. Bras. 142:260. 8 Bonon stylosa Backer, in Bull. Jard. Bot. Buitenz. 21:15. 1913. Drymaria tepicana M. E. Jones, in Contr. West. Bot. 15:124. 1929. Drymaria barrancae M. E. Jones, loc. cit. 18:65. 1931 Prostrate or ascending herbaceous annuals to as much as 45 cm. long, the inter- nodes mostly longer than the leaves, villose to hirsute with septate hairs to as much as2 mm. long. Leaves opposite, scantily to densely villose or hirsute with cinereous or ochraceous hairs, orbicular to reniform, apically rounded to acute and apiculate, basally cordate to truncate, weakly 3—7-veined, 5-15 mm. long, 5-15 mm. broad; petioles 1-10 mm. long, the stipules mostly entire, scarcely distinguishable from the villosity, 0.5-1.5 mm. long. Inflorescences of terminal 5-many-flowered cymes, the ultimate branches often tending to be racemose; peduncles 1-5 cm. long; bracts 0.5-1.5 mm. long, the pedicels 2-20 mm. long, villose. Sepals 5, 2-3.6 mm. long, 1-2 mm. broad, narrowly to broadly ovate or elliptic, apically acute to obtuse, villose to glabrous, weakly 3-nerved, with translucent borders; petals 5 (absent or drastically reduced in forms passing as D. Zepicana) , 2-3.6 mm. long, bifid for half their length or more, the lobes apically acute to deeply emarginate, 2—4-nerved, basally provided with linear auricles, these very variable in number and orientation; stamens usually 5, 2—3.5 mm. long, the oblong anthers 0.3—0.5 mm. long, the filaments shallowly connate, devoid of staminodia; ovary at anthesis ovoid to globose; styles 3, united for half their length or more, 1—1.5 mm. long; ovules numerous, campylotropous on free central placentae. Capsule ovoid to ellipsoid, 2-3.5 mm. long, many-seeded (rarely as few as two in D. fepicana), the seeds cochleate, 0.5—0.9 mm. broad, the dorsal tubercles filiform to capitate or subpinnate, longer than broad, the facial tubercles stellate. Central Mexico through Central America along the western coast of South America to Peru; apparently introduced and widespread in the East Indies. CHIRIQUÍ: Finca Lérida to Boquete, ca. 1300-1700 m., hoe Allen & Seibert Mi Bajo Chorro, Boquete District, 6000 ft., Davidson 294; vicinity of Boquete, 1200-15 Woodson 9 Scbery bot COCLE: lower portion of valley d. marshes along R. peo "El Valle de Antón, ca. 500 m., Hunter t$ Allen 379; between Las Margaritas and El Valle, Woodson, Allen & Seibert 1771. Examination of adequate material from Mexico, home of D. villosa, from Peru, home of D. hirsuta, and Java, home of D. stylosa, clearly shows these are con- specific, closely resembling the horticulturally propagated type of D. villosa, with the petal lobes spatulate, retuse and basally provided with downwardly directed linear auricles. The Indonesian material is marked, although inconstantly, by glabrous sepals and stiff erect hairs, i.e. they are truly hirsute as contrasted with the predominantly villose Mexican specimens. I have seen no specimens repre- senting this species from the West Indies and believe that the epithet cubensis stems from some error in transposition of labels. (444) 1961] FLORA OF PANAMA (Caryophyllaceae) 103 Forms comparable to D. tepicana M. E. Jones аге not infrequent. In these, the petals are reduced to mere vestiges or are completely absent, and the seeds are often fewer in a capsule than is typical, but I am strongly inclined to doubt that these forms warrant even formal recognition. It may be here appropriately mentioned that similar reductions in the petals occur in the closely related species Drymaria palustris Cham. & Schlecht., and such variants have been described as Drymaria townsendii Robinson. Standley and Steyermark (in Field Mus. Bot. 24*:232. 1946) list poleo, millón and llovizna blanca as Guatemalan names for this species. 2. Drymaria СОКРАТА (L.) Willd. ex Roem. & Schult. Syst. Veg. 5:406. 1819. Holosteum cordatum L. Sp. Pl. 88. 1753. Holosteum diandrum Sw. Prodr. 27. 1788. Drymaria та (Sw.) Macfadyen Fl. Jam. 1:52. 1837. non eee 1825. Drymaria cordata B diandra (Sw.) Griseb. Fl. Brit. W. Ind. 56. 1859 а Drymaria procumbens М. Е. Rose, in Contr. 1). 5. Nat. Herb. 1:304. 1895. Drymaria adenophora Urban, in Fedde, R pert. Sp. Nov. 21:213. 1925. Drymaria cordata var. pacifica Ида, in Jour. Jap. Bot. 32:78. 1957. Glandular-puberulent to glabrate ramifying annuals, prostrate and spreading or erect, the internodes mostly longer than the nodes, glabrous to densely glandular, often rooting at the nodes. Leaves glabrous to scantily pubescent, orbiculate to reniform, apically rounded and occasionally mucronulate, basally rounded to cordate, 5-25 mm. long, 5-30 mm. broad, the petioles 2-15 mm. long; stipules mostly polylacerate, rather persistent, to 2 mm. long. Inflorescences of terminal or axillary few—many-flowered dichasial cymes (flowers rarely solitary in the axils), the bracteate pedicels locally girdled with a dense band of glandular pubescence, rarely subglabrous, 2-15 mm. long, equaling to much exceeding the subtending bracts. Sepals 5, lanceolate to ovate, acute, glandular-puberulent to glabrous, obscurely to strongly 3-nerved, 2.5—4 mm. long, the borders translucent; petals 5, 2-3 mm. long, deeply bifid, the lobes linear, acute, rarely obtuse, 1-nerved, basally exauriculate but rarely subdentate; stamens 2-3 (-5), the flattened fila- ments 2-2.5 mm. long, the anthers suborbicular, 0.2-0.3 mm. long; styles 3, free nearly to their bases or united for half their length, 0.5—1 mm. long; ovules few to many, campylotropous on free central placentae. Capsule ovoid, 1.5-2.5 mm. long, the 3 valves entire; seeds 1-12, 1.0-1.5 mm. broad, cochleate, dark reddish brown, tuberculate in lines, all the tubercles low and domical, closely approximated or contiguous. A nearly pantropical species, in America ranging from Florida and Mexico through the West Indies and Central America along both coasts of South America to Argentina. DEL TORO: vicinity of Chiriquí Lagoon, von Wedel 1240. CANAL ZONE: du] Isthmus of Panamá, Fendler 9. Fendler 9 was cited in the original description of D. cordata B puberula Tr. & Planch. but differs little, if any, from typical cordate. Mizushima (in Jour. Jap. (445) (Мог. 48 104 ANNALS OF THE MISSOURI BOTANICAL GARDEN Bot. 32:69. 1957), studying the varietal potentials of this polymorphic species, described var. pacifica, which differs from typical cordata in having glabrous sepals, more seeds, and a tendency to have a greater number of stamens and the styles divided completely to the ovary. This rather striking variety seems to be lacking in continental North America. Occurring in both var. pacifica and var. puberula, for those who wish to recognize them, are variants in which the flowers are subsolitary in the axils. Surprisingly and fittingly these have as yet received no taxonomic status. Drymaria adenophora Urban represents a form with diminutive leaves and flowers and apparently is to be expected throughout the range of the species. Mizushima resurrected D. diandra Blume from a long submergence in the synonymy of D. cordata. D. diandra seems to be confined to the Old World and differs in having pyriform flowers with the outer sepals strongly carinate and three-ribbed, the ribs interconnected. The petal lobes are broader and more obtuse and the seeds tend to be fewer and larger than in D. cordata. mong the many colloquial Central American names for this species reported by Standley and Steyermark (in Field Mus. Bot. 24*:228. 1946) are palitaria or pelitaria, betatillo, comida de canario, trencilla, comapa, compona and mervillo. Because of its lush mat-forming properties, the plant is often cultivated as a ground cover. According to Dickson (in Tea Quarterly 18:84. 1960) however, the beneficial effects are nullified after a few years, the plant becoming so aggressive as to notably reduce the yield of a tea plantation. 5. POLYCARPAEA Lam. PorvcARPAEA Lam. in Jour. Hist. Nat. Paris 2:8. Ё. 25. 1792. Nomen con- servandum cte Adans. Fam. 2 271. 1763. In part Lour. Fl. Cochinch. 164. 1790. Ronis rejiciendum. Puedes Lam. in Jour. Hist. ШЕ Paris 2:1. 1792. Polycarpoea Lam. loc. cit. 25. 1792. Hagaea Vent. Tabl. 3:240. Sa Hagea Pers. Syn. 1:262. Mollia Willd. Hort. Berol. 11. #. 11. 1806. Lahaya Roem. & Schult. Syst. " 402. 1819. Aylmeria Mart. in Nov. Act. Nat. Cur. 13:276. 1826. Hyala L'Herit ex DC. Prodr. 35375. 1826; efi Webb, ex Berthel. Ayr Nat. Isles Canary 3?:156. 1836-50. Plancbonia J. Gay, ex Benth. & ue тыры 1:154. 1862. Robbairea Boiss. Fl. Orient. 1 Dis 18 Polycarpea Pomel, Nouv. Mat. Fl. Atl. D: 4. Polycarpus O. Ktze. in Post, & O. Ktze. Lexicon 453. 1904. In part. Annual or perennial, usually pubescent, erect or prostrate herbs. Leaves opposite or verticillate, usually linear and sessile at the swollen nodes, with con- spicuous scarious stipules. Flowers in terminal dichasial cymes, white or reddish. Sepals 4-5, occasionally biseriate. Petals 4—5, shorter than the sepals, entire or thed. Stamens 4—5, the anthers versatile, the flattened hypogynous filaments (446) 1961] FLORA OF PANAMA (Caryophyllaceae) 105 | ie > T ћи Fig. 141. Polycarpaea corymbosa slightly connate below. Ovary superior, unilocular, carpels usually 3; styles united below, the stigmata subcapitate; ovules few, campylotropous on basal placentae. Capsule ovoid to ellipsoid, dehiscing longitudinally into 2-3 entire valves; seeds I-few, cochleate, tuberculate, the embryo curved about the perisperm. Primarily a tropical genus of some thirty species, with only one American species reaching as far north as Panama. 1. PorvcaRPAEA CORYMBOSA (L.) Lam. Illustr. Gen. 2:129. 1793. Acbyrantbes corymbosa L. Sp. Pl. 205. 1753. Polycarpaea spadicea Lam. Illustr. Gen. 2:129. 1793. P olycarpaea indic а Lam. Encycl. 5:483. e M H Labaya corymbosa Roem. & Schult. Syst. >> ч. 1819. Р flora Wall. Cat. ycar paea densiflora 1. n. 1513. P ycarpaea b il ік Camb. in St. Hilaire, u Во 2:132. 1829. Рој ycarpaea brasilensis var. В ramosissima Camb. in St. Hilaire, loc. си. 2:132. 1829. Polycarpaea subulata Wight, & Arn. Prodr. 358. е Polycarpaea atherophora Steud. т Flora 26:763. 1843. Po Seer pna Феу? ifolia ses in Rep. Babb. ies ^ 1858. Р усатра torym Poss O.K deed Gen. 1: "n Polycarpa brevifolia O. Ktze ; Cit. 1:51. Poi ycarbaea filifolia Muschl. i > саң Jahrb. n 963 1911. Polycar. corymbosa var. typica Domin, in Biblioth. Bot. Heft 89?:655. 1925. P ycarpaea corymbosa var. . brevifolia Domin, loc. cit. 89?:655. 1925. (447) [Vor. 48 106 ANNALS OF THE MISSOURI BOTANICAL GARDEN Herbaceous, argentate-pubescent, erect virgate or tufted annuals or perennials to 1 m. high. Leaves verticillate, linear, aristate, 3-15 mm. long, mostly less than 1 mm. broad, villous or glabrate, the aristate, often lacerate, argentate stipules 2—5 mm. long. Flowers many in congested dichasial cymes, the puberulent pedicels 2-5 mm. long, mostly exceeded by the subtending bracts. Sepals usually 5, occasionally biseriate, lanceolate to ovate, attenuate to aristate, argentate or rarely rufous, often with a darker triangle at the base, 2-4 mm. long; petals 5, obovate, apically undulate, drying yellowish or brown, 0.5-1 mm. long; stamens 5, the flattened filaments 0.5—1 mm. long; ovary short-stipitate, obovoid, the style 0.25- 0.50 mm. long, the stigmata minute. Capsule ellipsoid, 1-2.5 mm. long, dehiscing longitudinally into 3 entire valves; seeds 2—6, ca. 0.5 mm. broad, cochleate, minutely corrugated, light brown, on basal placentae. Usually in arenaceous savannas, cosmopolitan in the tropics, in the Americas reaching its northern limits in Panama. COCLÉ: in savannas near sea level, Aguadulce, Pittier 4953. (448) MISSOURI BOTANICAL GARDEN TS ANDERSON, urator of Useful Plants г “шкы к. ANDREWS, Norron H. NICKERSON, „Ооћсв S. Buntine, У Taxonomis _ Носи С. CUTLER, Executive Director ы *- Donpee, Mycoloms: ' “РБ E p Р еи т ist and Editor of MES A. Duke. 2 “Assistant Curator of Herbarium Volume XLVIII Number 2 Annals of the Missouri Botanical Garden MAY, 1961 Morphology e Anatomy of the Saururaceae. І. а Anatom and Embryology M. V. S. Raju 107-124 Hepaticites devonicus, A New Fossil Erena iom the Devonian f > · . . . Егапсі M. Hueber 125-132 Index of Orchid Манар 1960 >. . . Robert L. Dressler 133-136 Preliminary Studies іп the Genus Stanhopea (Orchidaceae) . . £o си о 40. се СН. Dodion and G. P. Frymire 137-172 ED QUARTERLY AT GALESBURG, ILLINOIS | BY THE BOARD C pe Қ сарқын OF THE MISSOU. RI BOTANICAL GARDEN т. LOUIS, MISSOURI a Entered салш Illinois, | ame the Act at Mach 5, 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 Garden. sts Information The ANNALS оғ THE Missouki BOTANICAL GARDEN appears four times during the calendar year: February, May, September, and November. Four a volume. i Beginning with Volume 45, 1958: 54 509 ‘Subscription Price $12.00 per volume Single Numbers... 300 each 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. : Annals of the Missouri Botanical Garden Vol. XLVIII MAY, 1961 No. 2 MORPHOLOGY AND ANATOMY OF THE SAURURACEAE. I. FLORAL ANATOMY AND EMBRYOLOGY! М. У. $. RAJU ABSTRACT The present paper deals with certain aspects of — and anatomy of the Saururaceae. ned ie atomy and embryology trii a dns relationship among the genera. In floral anatomy the us Saururus is the least specialize nemopsis the most 5 highly specialized. Structure and те ogy of inflor sare? ce and De: Pda suggest the manifestation of certain evolutionary trends, Presa = by red cess cohesion and adnation. Origin of inferior ovary in the Saururaceae is disc сі лады nd nature of placentation show certain primitive features. Origin and evolution fro а | to syncarpous condition of carpels are briefly discussed. Further ме will | ы d in a separate paper. М. V. S. Raju, Department of Biology, University of Saskat chewan, Saskatoon, Saskatchewan, Canada. INTRODUCTION The Saururaceae has been one of the long-neglected angiosperm families in the field of morphology and anatomy. A perusal of the literature shows that earlier investigations have been superficial. In several discussions of comparative mor- phology and anatomy of the families of the order Piperales and other allied orders, some of the old descriptions of the Saururaceae have been considered. Very often, the family Saururaceae and its composition and affinities have been discussed without much detailed work. In view of this lack of sufficient information, an attempt has been made to investigate in detail some of the salient morphological and anatomical features. The Saururaceae, according to present knowledge, consists of 6 species—Sauru- rus cernuus L., S. chinensis (Lour) Baill, Houttuynia cordata Thunb., Anemopsis californica Hook. & Arn., Gymnotheca chinensis Decaisne., and С. involucrata P'ei. No material of the last genus was available for this investigation. However, at various stages of description and discussion, previous reports on Gymnofbeca have been taken into consideration. The present paper deals with floral anatomy and embryology and other related features of the Saururaceae. Members of the family Saururaceae are perennial aromatic herbs inhabiting moist places. The spirally arranged simple leaves are stipulate (Figs. 1, 5, 8). Houttuynia the stipules are prominent, often enclosing a part of the internode 1 An investigation carried out in the бәйек laboratory of the Henry Shaw School of Botany of Washington University and submitted as part of a thesis in partial fulfillment of the requirements for the к of Doctor 4 Philosophy. (107) [Vor. 48 108 ANNALS OF THE MISSOURI BOTANICAL GARDEN above the node (Fig. 8). In the other genera, they are not prominently visible but are adnate to the petiole and form a sheath around the node as in monocots. The leaves of Saururus, Gymnotheca and Houttwynia are auriculate to lanceolate and palmately net-veined. In the case of Anemopsis, leaves are elliptic-oblong with a conspicuous mid-vein. INFLORESCENCE The inflorescence of the Saururaceae is either a terminal spike or a raceme (Figs. 1, 5, 8). In Saururus the naked bracteate flowers are often arranged in pairs, a feature which distorts their spiral arrangement. The flowers are adnate to the subtending bract in earlier stages of growth. The flowers of S. chinensis are much more sparsely arranged than those of S. cermuus, but in most other respects, the inflorescences of the two species are stmilar (Figs. 12, 13). e inflorescence of Houttuynia cordata has been described in detail by Nozeran (1955). It is a spike with 4 conspicuous bracts at the base (some flowers, however, have small pedicels). Bracts higher up are much reduced to either scales or simple enations. The flowers show various degree of adnation of their pedicels with the bracts. However, in most cases this adnation is not superficially recognizable (Figs. 9, 15). In this species, Nozeran (1955) has reported a progressive trans- formation of complete flowers into staminate and pistillate ones within the same spike. Similar transformation has been observed by the present author. Structures of both flower and inflorescence of Anemopsis have been described by Quibell (1941). The inflorescence is a highly condensed spike and is subtended by 6-8 petaloid bracts. The young inflorescence axis in early stages resembles a flower covered by bracts. The inflorescence axis is mainly made up of “sunken” pistils (Fig. 16). The bracts, which vary in number, are foliaceous and gradually decrease in size from base upwards (Fig. 5). Except for the lowermost bracts, the rest are adnate to naked flowers. The inflorescence of Gymnotheca chinensis is also a spike, if one follows the terminology of Nozeran (1955). The bracts are not so conspicuous as those of H. cordata and A. californica. However, the gradual reduction in size of bracts from base upward is similar to the situation in H. cordata. FLOWER Flowers of Saururus, particularly $. cernuus show considerable variation in structure and number. An average flower of S. cernuus is pedicellate with 4 simple pistils and 6 stamens. Other combinations like 4-5, 4-7, 4-8, 5-7, 5-8, 3-4, and 3—6 have been observed. Serial sections of flowers of $. cernuus show the spiral arrangements of carpels and stamens. Carpels are very closely approximated to one another giving an impression of syncarpy. The stamens are hypogynous and are opposite the carpels (Fig. 2). Pistils show extensive stigmatic crests on adaxi sides of styles (Figs. 2, 4). The stamens are much longer than the styles. In S. chinensis flowers show the same variation as S. cernuus, but the pistils have shorter styles. Stamens in $. chinensis are perigynous and do not exceed the pisti in height. In Houttuynia cordata, the flowers are naked and have small pedicels. These ra aam ie ned ai 1961] RAJU—MORPHOLOGY AND ANATOMY OF SAURURACEAE 109 pedicels toward the inflorescence apex are further shortened or the flowers are sessile. The compound pistil is made up of three carpels, and the styles are free with wide stigmatic crests on their adaxial sides (Figs. 9, 10). In the arrangement of flowers, Gymnotheca resembles Houttuynia. According to the diagram of Decaisne (1845), the flower of Gymnotheca chinensis has a 4-partite pistil and 6 epigynous stamens (Fig. 14). е inflorescence of Anemopsis californica has pistils buried in the "inflores- cence axis”. In these naked flowers, the compound pistil is made up of usually 3 carpels with free styles. The epigynous stamens are usually six in number. How- ever, this number is subject to variation. The petaloid bract appears to be situated Text Fig. 1. Figs. 1-4. Sa "uus d “stil arrangement of leaves and terminal inflorescences. Fig. 2. A flower with bract, stamens and pistils (note the warty structures on ovary). Fig. 3. Sectional view of pistil to show the position of — and vascular traces. Fig 4. Longisection of a carpe 5-7. Anemopsis californica. Fig. 5. small plant with roots, rhizome, rosette of leaves and an inflorescence. Fig. 6. Longisection of ovary showing placentation, flared margins of stigmatic crest and us sta ig. 7 Transection same showing pari place on and lete septa. Fi 11. Houttuynia cordata. Fig. 8. portion ch wi an ig low | pound pistil and epicarpellary stamens. Short pedicel of flower is adnate to scale-like bract. Fig. 10. Longisection of pistil showing placentation and stigmatic crest. Fig. 11. Transection of ovary showing parietal placentation. [Vor. 48 110 ANNALS OF THE MISSOURI BOTANICAL GARDEN at the neck of the compound pistil. Developmentally it is adnate to the pistil along its length (Fig. 49). It is also interesting to note that in certain inflorescences bracts show a gradual change from base upwards from a petaloid bract to a petaloid stamen. This also followed by a highly reticulate venation pattern in the lowest bract to a highly reduced “dichotomous pattern” in bracts at the tip of inflorescence. Figs. 12-16. Diagrammatic sketches of portions of infloresc Text escences of Saururus cernuus, As chia ensis, Gymnotheca involucrata, Houttuynia tendat and Anemopsis californica respectively. Figs. 17-22. rurus cernuus—Transverse sections of flower (stamens not included) at different levels to show spiral arrangement of car rpels a: на ы, » чё Figs. 23—27. ғ : nd ome outtuynia cordata—Transection of compound pistil at en ten levels (diagrammatic). Figs 28— 31. mde d californica—T ransection Zi flower (stamens not included) at rose led (dia- 1961] RAJU—MORPHOLOGY AND ANATOMY OF SAURURACEAE 111 ANATOMY OF THE FLOWER Saururus.—Anatomy of the flower has been recently described for S. cernuus by Murthy (1959). Carpellary traces* in the flower originate spirally. Serial sections also indicate the spiral arrangement of carpels. Although ventral margins of carpels are closely approximated or partially fused at the lowermost regions of ‘pistil’, the ventral traces remain free throughout their course. In certain cases the ventral and dorsal traces of each carpel merge at the apex and appear as though fused to form a common trace. It is also not uncommon to find ventral traces ending near the base of styles. Ventral traces give off branches to the two ovules present in each carpel; placentation in the carpel has been here interpreted as laminar (Figs. 3, 4, 17-22). The anatomy of the flower of Saururus chinensis is similar to that of S. cernuus with, however, a few deviations. Stamen traces arise as branches from the dorsal traces of carpels. The ventral trace invariably merges in the stigmatic region with the dorsal trace to form a thick common trace. Unlike S. cermuus, there is no conspicuous style in S. chinensis. Houttuynia.—The naked flowers are trimerous and bracteate (Fig. 9). The vascular bundle from the inflorescence axis extends into the cortex and becomes the trace for the flower. It branches to form two traces, one leading into the bract and the other into the flower proper (Figs. 32, 33). The bract trace may or may not ramify depending on whether the bract is "leaf-like" or scaly. The remaining trace branches at different levels to form three independent carpellary traces which show a spiral arrangement (Fig. 32). Each carpellary trace gives rise to a dorsal and two ventral traces (Figs. 32, 34-39). The ventral traces of adjacent carpels fuse together to form "compound ventral trace" or placental trace. Each placental trace gives off branches which supply the ovules. The placental trace when it reaches the style bases separates and extends into the respective styles. Ventral traces together with the dorsal trace go into the style and all three remain separate. No instance of their fusion at the tip of style was found. Placentation is parietal and ovules are present only in placental region where the ventral traces have fused to form the placental trace. Stamen traces are formed much later than the ventral traces and arise about the middle of the pistil from the carpellary dorsal trace (Figs. 32, 40-48). Anemopsis.—Quibell (1941) has given in sufficient detail the vascular anatomy of the flower. As in other genera, a single trace from the inflorescence axis diverges and branches to form a bract trace and a "flower trace" (Figs. 49-52). Although the bract is adnate to the pistil, the bract-trace is free throughout (Figs. 49—66). Each “flower trace" branches to form ventral and dorsal traces. Each carpel receives two ventral traces and a dorsal trace. As in Houttuynia, ventral traces of adjacent carpels immediately fuse to form the placental trace whose branches supply the ovules. At about the base of style, the placental traces separate to give ventral traces which pass into the respective styles. Each ventral trace, on its way * For the sake %- convenience the term ‘trace’ has been used іп describing the vasculature of different organs of flowers. [Vor. 48 36 | | xt Fig. + Нош шута cordata. Fig. 32. Diagrammatic sketch of a flower showing vascular Р pgs Figs. 33-48. Transections of flower at different levels to show vascular anatomy of = | flower (В, bract trace; C1, 2,3, traces of three carpels; D, dorsal trace of carpel; D1, 2 3, dorsa ca trace or common trace of dorsal trace of carpel and stamen trace; P, pedicel trace; 5, stamen; 51,2 d stamen traces; Vi Сі, M Сі, Уі Со, V2 Co, Уз Са, Vo Ca, ventral traces of respective carpets; 1961] RAJU—MORPHOLOGY AND ANATOMY OF SAURURACEAE 113 ъъ —- a 2 2. 49. Diagrammatic sketch of а flower to show vascular L Же ons at different levels of а flower showing vascular traces (В, bract; aces; D, D1, D2, Ds, dorsal traces; SB, axial bundle of inflorescence; S, stamen trace; У, ventral trace), Х 100. ext Fig. 4. Anemopsis californica. Fi 0-66. Transecti 85. Сі, 2; 8» carpel tr. [Vor. 48 114 ANNALS OF THE MISSOURI BOTANICAL GARDEN Же X Med E the base of the style, is connected with the dorsal trace. The dorsal trace with the other two ventral traces extend into the style (Figs. 58-66). At the base of style the dorsal bundle gives off branches to one or two stamens. Placentation is parietal as in Houttuynia. CARPELS AND PLACENTATION A considerable variation has been found in the structure of carpels and in the organization of pistils in the Saururaceae. Carpels in Saururus cernuus are free throughout except at the base, a little above the short pedicel where they show a lateral concrescence (Figs. 17-22). They arise in a spiral fashion, a fact clearly seen in serial sections. There is considerable elongation above the ovule-bearing region. In Houttuynia, Gymnotheca and Anemopsis, clearly recognizable stylar regions are found. Furthermore, in all these three genera, the pistil is “compound” (in the sense that the ovary is syncarpous and the styles are free) (Figs. 6, 7, 10, 11). In Saururus cernuus, the folded laminae are conspicuous and the ventral traces are present away from the carpellary margins. During organization of the pistil, these laminar margins of adjacent carpels undergo concrescence which is to a considerable extent mistaken for axile placentation; placentation here is actually laminar. The lateral concrescence of adjacent carpels may be attributed to a case of “incipient syncarpous tendency” (Bailey and Swamy, 1951). The compound pistil of Houttuynia and Anemopsis is formed as a result of lateral approximation and concrescence of carpellary margins of adjacent carpels. This concrescence is also accompanied by fusion of ventral traces of adjacent carpels. Only in this region of concrescence of carpellary margins and also of lateral or ventral traces is parietal placentation organized. The sterile margins of carpels also undergo concrescence to form septa; as a result the compound pistil appears to be three-loculed (Figs. 23-31). This kind of incomplete septation is found in the upper part of the pistil of Houttuynia cordata; it extends almost half the depth of the pistil in Anemopsis. EMBRYOLOGY Saururus cernuus.—Microsporogenesis and male gametophyte.— he wall of a young anther consists of four layers, an epidermis, hypodermal layer, middle layer and an innermost tapetal layer. The mature anther wall has an epidermis with cutinized outer wall, hypodermal layer with characteristic fibrous thickening and Text Fig. 5. Saururus cermuus. Fig. 67. A portion of anther wall showing epidermis, hypo- dermal layer with endothecial thickening, crushed middle layer, and a tepetal layer, X 333: Figs 68—71. Stages in the formation of microspore, Ж 833. Fig. 72. A young ape showing megaspore mother cell and dier parietal cell, Х 333. Fig. 73. Nucellus showing division of nucleus of , X 666. Fig. 74. Same at a later stage with a dyad, Х 666. Fig. 75. Shows division of lower Puer cell, X 666. Fig. 76. A linear row of three cells or a "triad", the lowermost being the megaspore, X 666. Fig. 77. P er dra of middle cell of a linear “triad”. Note vacu- olation in the lowermost cell, X 666. Fig. 78. The upper two E of a "triad" have degenerated and the lowermost has become the ой, megasporo X 666 Ре. 2 Mature embryo sac with the egg apparatus and secondary nucleus хи 6. i. 80. Lo ongisec n of a young seed ito lower c i Se a шығ | 1961) RAJU—MORPHOLOGY AND ANATOMY OF SAURURACEAE 115 [Vor. 48 116 ANNALS OF THE MISSOURI BOTANICAL GARDEN tapetal layer with binucleate cells (Fig. 67). The sporogenous cells differentiate themselves into microspore mother cells which undergo meiosis and form microspore tetrads and then microspores (Figs. 68—71). Megasporogenesis and female gametophyte.—The crassinucellate ovules are orthotropous-bitegmic and borne on laminar placentae (Figs. 3, 4, 19-21). micropyle is organized by both the integuments. The outer integument is two layered; the inner is three layered. A single hypodermal archesporial cell is differ- entiated in a young ovular primordium. It undergoes a transverse division to form primary sporogenous and parietal cells (Fig. 72). The primary parietal cell, by anticlinal and transverse divisions, forms a parietal tissue of about 8 superposed cells (Figs. 73-78). The primary sporogenous cell divides meiotically to form a dyad (Fig. 74). During further development, the lower dyad cell divides and the upper does not (Figs. 75-76). Thus, the dyad gives rise to a “triad” or a linear row of three cells (Figs. 76,77). Of these three cells, the upper two degenerate and the lower- most develops into the embryo-sac (Figs. 77-79). The antipodal cells are ephem- eral and seldom seen in a mature embryo sac. The egg apparatus is situated at the micropylar end and no case of fertilization was observed. Endosperm.—The endosperm nucleus divides transversely to form a micropylar chamber and a chalazal chamber. The chalazal chamber behaves as a haustorium and the nucleus becomes hypertrophied. The mature seed encloses massive peri- sperm and a little endosperm surrounding the developing embryo (Figs. 80-82). Houttuynia cordata.—Microsporogenesis and male gametophyte.—Microsporo- genesis and male gametophyte development follow the sequence given for Saururus cernuus, but with some differences. Division of the microspore mother cell shows various cytological abnormalities resulting in the formation of sterile pollen. Frequently, micronuclei are seen in microspore tetrads. Very rarely free micro- spores are observed (Figs. 83-88). Megasporogenesis and female gametophyte.—The bitegmic orthotropous ovules are borne on parietal placentae (Figs. 25, 26, 89-90, 99). The ovular primordium shows a hypodermal archesporium (Figs. 89, 90). Similar to S. cernuus, the outer integument is made up of two layers and inner of three layers (Fig. 99). Again, the micropyle is organized by both the integuments (Fig. 99). An archesporial cell of hypodermal origin functions directly as the megaspore mother cell (Figs. Text Fig. 6. Houttuynia cordata. n of anther showing anther wall and sporogenous tissue; ққ wall shows снае, Ыг аши Jive crushed middle layer аы tapetal layer, X 333. Fig. 84. Same at a later tage; note binucleate tapetal cells, Х 333. Fig. 85. Mature anther wall а dolci dick in hypodermal layer; note also microspores, x Е 86. Мега aphase I in microspore pus ther cell, Х 666. Fig. 87. Same showing anaphase I; note е ent of с i i 8 а. tj Fig. d cells, x 666. Fig. 97. А mature embry with e paratus, secondary nucleus and an- прода! cells, X 333. Fig. 98. A че showing $a; s гура ag division of endosperm nucleus has resulted in two cells, che lower one being a haustorium, Х 333 . 99. A young ovule showing the Te egaspore, massive себе, and two integuments, і inner У cn dere and outer two- layered, Х 1 1961] RAJU—MORPHOLOGY AND ANATOMY OF SAURURACEAE 117 [Vor. 48 118 ANNALS OF THE MISSOURI BOTANICAL GARDEN 91-98). 'The megaspore mother cell divides meiotically to give a linear row of three cells of which the lowermost is the megaspore that develops into the embryo sac (Figs. 91-98). Antipodal cells in the mature embryo sac are epheremal but in some cases these cells are seen in mature embryo sacs (Figs. 96-97). The egg apparatus in the micropylar end is made up of an egg cell and two hooked synergid cells (Fig. 96). Неге, also, fertilization stages were not observed. Endosperm.—The endosperm behaves as in S. cernuus (Fig. 98). In mature seeds often only endosperm and perisperm were observed. The embryo was not seen. Anemopsis californica.—Microsporogenesis and male gametophyte.—Micro- sporogenesis and development of male gametophyte simulate those of Saururus cernuus and Houttuynia cordata (Figs. 100-105). Megasporogenesis and female gametophyte.—The orthotropous bitegmic ovules are found on parietal placentae as in H. cordata (Figs. 6—7, 30-31). A hypodermal archesporial cell is differentiated and divides to form a parietal cell and primary sporogenous cell (Fig. 106). The primary sporogenous cell divides meiotically to give a row of three cells as in other genera (Fig. 107). The lowermost cell functions as the megaspore and develops into an embryo sac (Figs. 107-110). Stages in fertilization have not been observed. Endosperm.—The earlier stages of endosperm development are like those of Houttuynia and Saururus cernuus. In the mature seed, massive perisperm, degen- erated embryo and endosperm were observed. FRUIT In Saururus simple pistils organize the fruit. The pericarp becomes dry-fleshy enclosing a single seed. No case of fruit dehiscence has been observed. Further- more, there is no stony endocarp developed in the pericarp. On the other hand, seed coat becomes very hard and often closely abutting the inner wall of pericarp. The fruit is here interpreted as a berry In Houttaynia and Anemopsis the fruits are dehiscent capsules with many smaller seeds. The dehiscence of fruit occurs at the apical region by the disinte- gration of fused ventral margins in the middle region of three free styles. In a dehisced fruit a circular port is seen at the top of the ovary leaving the styles free. Discussion Observations on inflorescence of the six species of Saururaceae show an inter- esting evolutionary trend, probably guided by two processes: 1. reduction, and 2. cohesion and adnation. Of the four genera, Saururus shows the least change by the above processes. The “common stalk” of flower and bract shows various degree of reduction. In Saururus the “common stalk” is much elongated and shows stages of its reduc- tion. Further reduction of the same has resulted in a condition seen in Houttuynia in possessing a recognizable "common stalk” (Figs. 12, 13, 15). Amemopsis, on the other hand, shows the final culmination where the pistils appear embedde in the “inflorescence axis” and the bract appears to be situated at the top of the ovary (Figs. 16, 49). In all genera this process of reduction is associated with | | | | | 1961] RAJU—MORPHOLOGY AND ANATOMY ОЕ SAURURACEAE 119 $ TA Әуе A > К ои P es {е 1 T ip (x % О Tii a zs yy 108 ü HAM 642, ја ; HL NOUO J C TE WOW TI acu Q ХНА PEN НЕЙ ва! М dermal layer, crushed middle and tapetal layers surr 1. same at a later stage showing endothecial thickenings in the ermis; two-celled microspores are seen, X 333. Figs. 102-105. Stages in the formation of microspores, Х 666. Fig. 106. А young ovule showing megaspore mother cell and two integuments; note parietal cells, Ж 333. Fig. 107. A portion of nucellus showing a linear row of 3 cells of which the lowermost develops into an concomitant cohesion and adnation of floral parts. By a process of reduction in the number and cohesion of carpels, the polycarpellate condition seen in Saururus has resulted in the three or four carpelled syncarpous condition found in the other genera. This change has been accompanied by the adnation of stamens to carpels and is exhibited by Houttuynia, Gymnotheca and Anemopsis. The genus Anemop- sis has an inferior ovary, a feature that has resulted by their “sinking into” the inflorescence axis. (Мог. 48 120 ANNALS OF THE MISSOURI BOTANICAL GARDEN The phenomena mentioned above may also be explained in the light of floral anatomy. Although the bract appears to be a simple structure, it is very difficult to see where the bract begins and where it ends, It is morphologically intriguing to consider the trace of the common stalk” as a “compound” one of bract and flower pedicel. This condition has been interpreted in Saururus cernuus as a case of adnation of flower to bract, by Murthy (1959). Present observations show that only the pedicel of the flower is adnate to the bract; this feature is evidenced by vascular anatomy of Houttuynia and Gymno- theca. In Saururus, Houttuynia and Gymnotbeca, the bract tends to remain free from the flower proper. But, in the case of Anemopsis, however, the bract is adnate to the pistil. This feature is of secondary derivation for the bract trace is completely free from the flower. This secondary condition is, probably, due to "development sinking-in" of the pistil. It can also be interpreted as a case of lateral concrescence of compound pistils in the inflorescence. In Saururus cernuus, stamen traces are formed before the carpellary traces are differentiated. But in S. chinensis, vascular anatomy indicates that stamens have originated subsequently from the dorsal carpellary trace. This specialization has proceeded in increasing degree іп Нои иупіг, Gymnotbeca апа Anemopsis where the stamens are perigynous or epigynous. In Saururus ventral traces are as prominent as the dorsal traces; they remain free from one another throughout their course except at the stylar tip where they show signs of fusion. Similar behavior of ventral and dorsal traces is seen in Houttuynia and Anemopsis; their fusion in the stylar tip is seldom observed. Besides, in these two latter genera, ventral traces of adjacent carpels fuse to form a placental trace. Anemopsis shows a unique feature in that the ventral trace has independent connections with the dorsal trace (Fig. 49). The exact significance of these connections is not known. To begin with, the flower gets a single trace from the inflorescence axis in Saururus, Houttuynia and Anemopsis. Such a feature has been reported in many angiosperms. Puri (1951) in his review paper suggested that highly reduced flowers received just one trace. "Whether this condition is simple by primitiveness or by reduction remains to be critically examined. As far as the present observa- tions go, it seems that traces of different organs have undergone fusion to form a single composite trace. The Saururaceae show both hypogenous and epigynous conditions. The prob- lem then arises as to how the epigynous condition has been achieved. The problem of origin of the inferior ovary has been highly debated for a long time. Douglas (1944) in her comprehensive review has discussed different views regarding morphology of the inferior ovary. Of the many views postulated, only two have remained prominent; they are appendicular and axial views. It is beyond doubt that in all genera of Saururaceae except Saururus cermuus stamens have fused with the carpels. In Houttwynia and S. chinensis stamens аге situated half way up on the pistil. Anemopsis and Gymnotheca have their stamens on the ovary just below the styles. "These clearly show that concrescence of certain v ақынын 1961] RAJU—MORPHOLOGY AND ANATOMY ОЕ SAURURACEAE 121 floral parts (here stamens) with the pistils has given rise to epigyny. This is in support of the views expressed by van Tieghem (see Puri, 1951, 1952; Eames, 1931). In Anemopsis, besides the fusion of stamens with the ovary, there is develop- mentally a concrescence of pistils resulting in a massive spike. This condition has given an impression that pistils have “sunken”. Adnation of the bract, with its free trace, to the pistil is an additional evidence to show that the ovary wall is made up of tissues other than that of the ovary wall itself Prior to syncarpy and epigyny, profound morphological and anatomical changes have taken place in flowers of the Saururaceae. These changes have been well preserved in the genus Saururus. The carpels with their respective stamens are arranged “spirally”. The last-formed carpel does not possess a stamen or stamens. This detail suggests that the flowers of S. cernuus, adnate to bracts, are not actually simple; they are, probably, derived from a structure similar to an “inflorescence”. Absence of an axial tissue, development of a terminal normal or abortive carpel with or without a stamen or stamens, and close telescoping of simple pistils suggest that the flower is of the “sympodial type". Each flower then represented by a carpel with its own stamen or stamens. A similar analogy can be traced in the flowers of Chloranthaceae (Cardemoy, 1863; Hooker, 1890). However, this idea has been contradicted on the basis of the absence of supernumerary structures in the flowers of Chloranthaceae (Swamy, 1953). The successive arrangements of simple pistils with their associated stamen or stamens lead to two problems—nature of epigyny or perigyny in a simple pistil and same in compound pistil. In the case of Saururus chinensis the stamen or stamens аге perigynous. This can be interpreted as a case of adnation. Syncarpous gynoecium also shows perigynous and epigynous conditions. It is difficult to say whether the cohesion of carpels appeared first or adnation of stamens to carpels. Available evidences, however, show in the Saururaceae that adnation of stamens to carpels appeared first as exemplified by S. chinensis. So, the flowers of Hauttuynia and Anemopsis suggest that they are made up of three independent flowers which have undergone phylogenetic fusion. This is also amply evidenced by floral anatomy. The above interpretations further lead to the problem whether the pistil is appendicular or axial. Vascular anatomy shows that there is nowhere in the ovary a residual trace or an axis around which carpels are arranged. The vascular elements are completely used up by the differentiated carpels. It is on this basis that flowers of Saururaceae have been interpreted as condensed inflorescence of the "sympodial type" simulating to a considerable extent vegetative branches where sympodial growth prevails. Anatomical evidences do not, however, give a correct picture of inferior ovary. Developmental studies are necessary for the interpreta- tion of inferior ovary in some members of Saururaceae. The trends of specialization postulated by Bailey and Swamy (1951) for the ranalian carpel have recently been useful in interpreting some primitive carpels. Although carpels of Houttuynia and Anemopsis are quite specialized, the condupli- cate nature is still present in their styles which have flared-out stigmatic crests. [Vor. 48 122 ANNALS OF THE MISSOURI BOTANICAL GARDEN Although ontogenetic studies have not been made, available observations indicate that the ovule-bearing capacity of the carpel has brought several modifications in its train. The closure of carpels has occurred in the basal or ovule-bearing part leaving behind an extensive stigmatic crest. The stigmatic crest is supplied by both dorsal and ventral bundles. This feature is seen in Saururus, Houttwynia and Anemopsis. In some of the carpels, particularly of $. cernuus, retraction of ventral traces from stigmatic region toward the ovule-bearing part may be noticed. This retraction leaves a short style between ovary and stigma. In the case of S. chinen- sis, the stigmatic crest is supplied by both ventral and dorsal traces; it is situated just above the ovule-bearing part. This progressive phylogenetic specialization leading to the formation of a style between stigma and ovary is exhibited by the carpels of Saururus, Houttuynia and Anemopsis. Superimposed on the evolutionary specialization mentioned above there is con- comitant lateral concrescence of carpels resulting in a compound pistil. Further- more, the "spiral" arrangement of carpels becomes less and less clear. Thus, in the development of pistils, simple or compound, several independent trends of speciali- zation have occurred simultaneously in the Saururaceae. Placentation in the Saururaceae is of two types, laminar and parietal. Laminar placentation is found in Saururus and parietal placentation in НоиНиута and Anemopsis. Bailey and Swamy (1951) after an extensive survey of carpels in the Ranales have come to the conclusion that carpels with laminar placentation are primitive and show various trends of specialization leading toward other types of placentation. A detailed study of carpels of Saururus has shown not only a laminar placentation but also different degrees of incipient syncarpous tendencies. The carpellary margins undergo concrescence or close approximation. is con- crescence of carpellary margins does not occur at one level since they are arranged spirally. Axial tissue that is seen at the base of carpels is actually a composite of different carpellary margins. As a consequence, the pistil appears to be compound with more than one locule. In fact, this condition probably led several taxonomists to interpret this placentation as axile (Bailey, 1958; Lawrence, 1951; Benson, From this basic laminar placentation, parietal placentation of Houttwynia and Anemopsis can be derived. The presence of free styles in the pistils of the above two genera shows that lateral concrescence of adjacent carpels has occurred in the ovule-bearing parts of carpels. Closely approximated partitions appear in the locule of the ovary of Houttuynia and Anemopsis. These partitions are the f ventral carpellary margins. In the ovule bearing part, ventral parts of carpels have been retracted considerably apparently to accommodate the ovules developing inside, In the organization of syncarpous ovary from simple carpels in Saururaceae, the so-called classical “torus” is not involved. The evolutionary trend of specializa- tion from laminar to parietal placentation involves: firstly, the fusion of ventral margins of adjacent carpels; and secondly, lateral concrescence of the same carpels. These are further influenced greatly by the ovule bearing capacity of a syncarpous ovary. In Houttuynia and Anemopsis the ovary becomes “multilocular”. The 1961] RAJU—MORPHOLOGY AND ANATOMY OF SAURURACEAE 123 retraction of partitions within the ovary in the lower part may be due to the acquisition of ovule-bearing capacity of ovary. Normally, the ovaries of genera mentioned above show multilocular nature in the upper part of ovary and uni- locular in the lower part. Intermediate conditions showing various stages of phylogenetic retraction of partitions are not infrequent. Eichler (1875-78) interpreted the flowers of Saururus and Houttuynia as obdi- plostemonous; flowers which had two whorls of three stamens each and the outer whorl had aborted. In the present investigation the above-said feature has not been observed. In Anemopsis and Gymnotheca, each flower is usually associated with six stamens. Anatomy of flower has shown that traces for stamens arise from the dorsal bundle of carpels. In addition to the above feature, the first formed carpels of Saururus usually possess two stamens each, the last formed does not develop stamens. The absence of correlation between the number of carpels and stamens, negates the possibility of recognizing two whorls of stamens. A similar view has also been advanced by Nozeran (1955). In his comparative survey of floral structures Nozeran (1955) writes that "the spike of Houttuynia is not entirely an inflorescence and is not still a flower”. Furthermore, he hypothesized that the lowermost bracts transform into perianth lobes. Не also observed in inflorescence several instances of reduction in number of stamens and carpels. On the basis of above details, he derived the spike of Houttuynia from a homogeneous inflorescence which was composed probably of trimerous flowers. In the present investigation, however, all the details given by Nozeran (1955) were not found. There is no doubt that at the tip of the inflores- cence, an overy was often borne, but there has not been enough evidence at present to support Nozeran’s hypothesis. Embryological investigations show great similarities among the three species, Saururus cernuus, Houttuynia cordata and Anemopsis. Development of both male and female gametophytes are very similar. During development of female gameto- phyte the archesporial cell gives rise to parietal cells in Saururus and Anemopsis. These parietal cells are absent in Houttuynia. The development of the female gametophyte in all three species studied conforms to the monosporic type (as here interpreted). Some stages in the development of endosperm were observed in Houttuynia and Saururus cernuus. Іп both cases the endosperm is of the cellular type. The above two genera also show endosperm haustoria. The embryo in S. cernuus suggests that the first division of the zygote should be transverse. Seeds enclose a massive perisperm and a little endosperm surrounding the small embryo. Similar embryological features have been recorded by Johnson (1900) for S. cernuus. SUMMARY An account of floral anatomy and embryology suggests that the genera of Saururaceae are closely interrelated. Structure and morphology of inflorescence and flower are discussed in the light of anatomy. Anatomical details indicate that the inflorescence-type seen in Amemopsis is more specialized than the type seen іп Saururus or other genera. This has been [Vor. 48 124 ANNALS OF THE MISSOURI BOTANICAL GARDEN achieved, according to the evidences advanced, by processes of reduction, adnation and cohesion of various floral entities. Some of the morphological aspects of ovary, carpels and placentation have been discussed. Embryological details are similar in all the genera investigated here. ACKNOWLEDGEMENTS I am greatly indebted to Dr. N. H. Nickerson, for his kindness and generosity and valuable guidance during the course of this investigation. I am also highly thankful to Dr. R. E. Woodson, Jr., for generously placing some of the material used here at my disposal and for helpful suggestions. LITERATURE CITED ema I. W., and Swamy, В. С. 1. (1951). е а carpel of dicotyledons and its initial ends of speciali nee Amer. J. Bot. 38:3 Bailey, H. (1958). ual of ар i planes” 5 Revised ачан 2 Сол М. X: Benson, L. (1957). Tint "Classifica ath a Cardemoy, J. de. (1863). Monogra shard du panes с ти Chlorate. и 3:280—310. Decaisne, J. (1845). Gymnotbeca chi inensis Decne. Ann. Sci. Eames, A. J. (1931). The vascular anatomy of de flower d и of the theory of carpel lymorphism. Amer. J. Bot. 18:147-188. Hooker, J. D. (1890). “Flora of British India”. Vol. 5., London Johnson, D. S. (1900). On the development of уна cernuus 1. Bull. Torrey Bot. Club 27: 365—372. Lawrence, С. Н. М. (1951). ЧЇ еседен of Vascular Plants”. MacMillan Murthy, Y. 5, (1959). Studies run К“ 5 іре ales. VII. A contribution to vag Fee of Saururus cernuus L. Jour. Indian Bot . 38:1 heey René. (1955). Catia 3 a У AH de quelques structures florales. Ann. Des. Sc. Nat., 6:1—224. Pun, V. m The role of floral anatomy in the solution of morphological problems. Bot. Rev. 17:471—553. 952). Placentation in Angiosperms. Bot. Rev. 18:6 , (1 Quibell, с с. H. (184) Floral anatomy and morphology of рН " californica Н. & A. Вог. az. 102:749— Swamy, B. А $ ets The morphology and relationships of the Chloranthaceae. Jour. Arnold Arb. 34:375—408. HEPATICITES DEVONICUS A NEW FOSSIL LIVERWORT FROM THE DEVONIAN OF NEW YORK? FRANCIS M. HUEBER ABSTRACT Hepaticites не is described from the lowermost Upper Devonian of New York. Мапу fragments were obtained by macerating shale This is the oldest ій меле reported to = It is tentatively My MR to be a member of the Anacrogynae, and is compared with the genera Pallavicinia and Metzgeria. Francis M. HUEBER, Department of Бану; Cornell Univerdey, Ithaca, New York. INTRODUCTION The record of the Hepaticae in the Paleozoic is very meager. One may readily expect this to be the case owing to the small size and the delicate nature of the members of this group of plants. Up until now the oldest occurrence of undoubted hepatics was that of four species from the Upper Carboniferous of Shropshire, England (Walton, 1925, 1928). They are Hepaticites Kidstoni, H. lobatus, Н Langi (Walton, 1925), and H. Metzgerioides (Walton, 1928). These four fossil forms have been compared with the living genera Treubia, Fossombronia, Riccardia (Aneura) and Metzgeria respectively. In all cases the remains are of sterile shoots only and on the basis of their morphology it has been assumed that all of the forms are of the anacrogynous series of the Jungermanniales (Walton, 1925, 571). Definite conclusions can not be made until reproductive organs are found. MaTERIALS AND METHODS The plant remains which are described here occur as compressions associated with other plant remains in a fine-grained, dark-gray shale. The specimens were collected from a small quarry located on the northwest slope of South Mountain, Schoharie County, New York, 1.1 miles west of the Schoharie-Greene County line Livingston 7' 30" map at 74? 16' 30" E., 42? 23' 55" N. and at an elevation of 2180 feet. The horizon at which the quarry is located is very close to the base of the Onteora "red beds" which are the continental equivalent to the lowermost two-thirds of the marine Finger Lakes stage, Senecan series (Frasnian) of central and western New York. e beds are therefore lowermost Upper Devonian. The plants with which the specimens of Hepaticites are associated, that is in the same beds and at the same quarry, are Psilophyton princeps Dawson 1871, Schizopodium Harris 1929, Cladoxylon Unger 1856, Pseudosporochnus Potonie and Bernard 1904, Eospermatopteris (Dawson) Goldring 1924, Callixylon Zalessky 1911, and two new genera all of which are to be described in future papers. While preparing transfers of other plant remains, thin fragments of plant material were seen in the sludge derived from the dissolution of the shale. The fragments at first were thought to be epidermal layers from plants contained in the shale. However, after they were washed, dehydrated and mounted, they were This paper is a part of a thesis presented to = Graduate School of Cornell University in partial flies rs the epe for the ER. D. deg cated by Hen drew (125) (Мог. 48 126 ANNALS OF THE MISSOURI BOTANICAL GARDEN found to possess the characteristics of the genus Hepaticites. Additional prepara- tions were made using the bulk maceration technique (Harris, 1926). Pieces of rock in which large quantities of plant remains are evident are placed into poly- ethylene or polystyrene plastic containers. Concentrated hydrofluoric acid (48%) is poured over the specimens until they are completely immersed in the acid. From ten to twenty-four hours are generally required for dissolution of the rock; the time is of course dependent upon the size of the specimen being dissolved. Frequent observations of the progress of the reaction are necessary in order that any freed plant material which is either floating on the surface of the acid or lying in the sludge can be removed. The remains are easily removed by slipping a flat, wooden rod under the specimen and then carefully raising the rod out of the solution. The specimen is "floated" onto the rod in the way that one floats specimens of algae onto paper for the preparation of herbarium sheets. The writer has found that the wooden labels used for flower pots are very satisfactory for lifting the specimens from the solution. The plant material is floated off of the rod into a container of water and washed through three or four changes of water. The wash water is either decanted or siphoned off. After the specimen is washed, it is floated onto a glass slide and positioned on the slide by the use of brushes cut down to a single bristle. The slide is gradually raised from the water, keeping the slide inclined. A drop or two of liquid detergent added to the water will help reduce surface tension and facilitate the removal of the specimen. Any water remaining on the slide an around the specimen is carefully removed with blotting paper. After the specimen has partially dried, it is saturated with a 1% aqueous solution of gum arabic which when dried serves to hold the specimen in place on the surface of the slide. The gum arabic solution is applied with a fine-tipped brush. When the specimen is fully dried, a drop of Harleco Synthetic Mounting Medium or Canada balsam dissolved in xylene is placed on top of it and a coverslip lowered over the surface. At this point, if the specimen were not fixed to the surface of the slide with gum arabic, it would migrate to the edge of the coverslip or be broken by the flow of the viscous mounting medium. The mounting medium is dried and the prepara- tion is ready for study. Nearly eighty fragments were prepared in this way. Genus HEPATICITES Walton 1925 НЕРАТІСІТЕЅ devonicus sp. nov.—Description The thallus is very simply differentiated into two parts, a rhizomatous portion and a thallose region. The latter is differentiated into two parts, an axial or costal region composed of elongate parenchymatous cells and marginal wings or lamellae composed of polygonal parenchymatous cells (figs. 1-5). The axial region aver- ages 0.56 mm. in width. It is two to several cells thick; the actual number is impossible to determine because of the complex reticulum formed by the outlines of the cells. A midrib composed of very elongate cells and averaging 0.22 mm. in width forms the central portion of the axial region. е midrib is in turn sur- rounded by a zone of elongate cells which are transitional in size between the midrib and the cells of the wings. The cells are elongate parallel to the long axis of the thallus and аге 14 to 26 и in diameter. Because of the incomplete preserva- 1961] HUEBER—HEPATICITES DEVONICUS 127 tion of the end walls, the full length of any one of the cells of the midrib is unknown. The cells which form the transition zone between the midrib and the wings average 140 p in length. Мо rhizoids are evident along the axial region. he wings are only one cell thick and vary from 1.3 to 1.75 mm. in width. cells are 46.8 to 73.2 p in diameter and are polygonal in outline. Scattered through the tissue of the wings are dark cells (fig. 5). The contents may indicate that some of them were storage or secretory cells. Some, however, may be darkened due to the activity of fungi, for fungal hyphae are commonly observed adhering to the surfaces of the thallus. The margins of the wings are serrate, the teeth are spaced at intervals of 0.18 mm. (figs. 6, 7), but otherwise the margins show no well pronounced lobing. Dichotomous branching of the thallus is evidenced in three specimens, two of which are illustrated in figs. 2 and 3. Fragments of the wings are still attached to the axial regions beyond the dichotomy. Growth of e plant was apparently the result of the divisions of a single apical cell borne in a notch at the apex of the thallus. A fragment of the thallus illustrating what may be interpreted as an example of the apical notch is shown in fig. 4 The rhizomatous portion of the plant (fig. 8) varies from 0.58 to 1.7 mm. in width and is composed wholly of elongate parenchymatous cells, The midrib varies from 0.15 to 0.28 mm. in width; the cells of which it is composed vary from 14 to 29 и in diameter and as in the midrib of the thallus portion their length is unknown. The cells which surround the midrib are approximately the same diameter as those in the midrib and they vary from 136 to 220 y in length. Numerous rhizoids arise from the cells on the surface of the rhizome (figs. 8 and 9). They аге nonseptate and average 21 p in diameter just above the broadened base. The tips of the rhizoids are broken off and therefore the full length of any one individual is not known. The largest fragment is 0.56 mm. in length. It must be stated here that no rhizomatous portion has been found in direct organic connection with the thallose portion of the plant. Both parts were isolated from the same matrix and since the part which is here referred to as the rhizome has the same general form and structure of the axial region of the thallose segment, the writer feels justified in considering them parts of the same plant. Discussion A comparsion of the illustrations of Hepaticites devonicus with those for the species Н. Mefzgerioides (Walton, 1928, Plate XII, Figs. 1-6) will show how similar the thallose portion of H. devonicus is to one which has already been included in the genus Hepaticites. The finely serrate margin, the lack of rhizoids on the thallose portion of the plant, and the differentiation of a rhizome bearing rhizoids, all characteristics of H. devonicus, are the evident morphological differ- ences between the two species. The difference in the gross size between the two (H. devonicus is perhaps four to five times larger than H. Metzgerioides) may be of specific value, but the similar size of the constituent cells in the two plants suggests that the difference is probably one of growth form. As in H. Metzgerioides, H. devonicus shows a clear distinction between midrib and wing which is also characteristic of the modern genera Pallavicinia Gray, corr. Carruthers and Metzgeria Raddi. Hepaticites devonicus combines characteristics (Мог. 48 128 ANNALS OF THE MISSOURI BOTANICAL GARDEN of both of the modern genera. It is similar to Pallavicinia in the width of the thallus (3 to 4 mm.) and may represent a plant like Pallavicinia Zollingeri (Gottsch.) Schiffn. (See Smith, 1938, especially Fig. 32 and Campbell and Williams, 1914, especially Fig. 4) in which there is differentiation of a prostrate portion bearing rhizoids and an erect, branched portion on which rhizoids are not present. The small marginal teeth of the thallus in H. devonicus is another char- acter which is shared with Pallavicinia Zollingeri. Hepaticites devonicus differs from Pallavicinia in having a less differentiated central strand and larger central cells in the strand. Hepaticites devonicus and Metzgeria are similar in their slightly differentiated central strands, in the size of the cells in the central strand, the size of the cells in the wings, and the possibility that the teeth on H. devonicus are homologous to the hairs on the margins of the thallus in Metzgeria. They differ in that Metzgeria is smaller (1 to 2 mm. wide) and bears rhizoids along its midrib. Ftepaticites devonicus resembles Metzgeria in cellular details but resembles Palla- vicinia in size and form. Reproductive structures are not known for H. devonicus but on the basis of its similarity in habit and the many details of structure which it shares with mem- bers of the Anacrogynae the writer suggests that it be classed with this group until such a time as further evidence of its relationship is obtained. The occurence in Devonian sediments of remains assignable to the genus Hepaticites extends the range of the Hepaticae even further back in geologic time, and it emphasizes that this class of plants had evolved to a high order of organiza- tion far earlier than heretofore recorded. НЕРАТІСІТЕЅ devonicus Hueber sp. nov. Diagnosis: Dichotomously branching thallus differentiated into rhizomatous por- tion bearing rhizoids and thallose portion composed of axial region averaging 0.56 mm. in diameter and wings one cell in thickness ranging from 1.3 to 1.75 mm. in width. Cells of the axial region elongate, 14 to 26 и in diameter, length of central cells undetermined, length of transitional cells averaging 140 и. Cells of the wings polygonal, 46.8 to 73.2 и in diameter. Margins of wings finely serrate, lobing indefinite. Rhizomatous portion 0.58 to 1.7 mm. in diameter, cells elongate as in axial region of thallose portion, numerous non septate rhizoids averaging 21 р in diameter borne on surface of rhizome. Holotype number 23, specimen 1579-294, slides 6-12, paleobotanical collections of the Department of Botany, Cornell University, New York. Locality: Northwest slope South Mountain, Schoharie County, New York. Horizon: Base of the Onteora “red beds” lowermost Upper Devonian. SUMMARY A new species of Hepaticites, H. devonicus, is described from the lowermost Upper Devonian of New York. The plant is differentiated into two parts, 4 rhizomatous portion and a thallose region. The latter is differentiated into two parts, an axial region composed of elongate parenchymatous cells and marginal wings which are one cell thick and composed of polygonal parenchymatous cells. 1961] HUEBER—HEPATICITES DEVONICUS 129 The margins of the wings are finely serrate and show no lobing. There are no rhizoids borne on the thallose portion of the plant. The rhizome is composed wholly of elongate parenchymatous cells and resembles the axial region of the thallose portion of the plant. Rhizoids occur over the entire surface of the rhizome. No reproductive structures are known. The plant is compared with Pallavicinia Zollingeri and Metzgeria and placed for the time being with the fossil forms of the Anacrogynae. The report constitutes the earliest record of the Hepaticae in the geologic column. ACKNOWLEDGEMENTS The writer wishes to express his sincere appreciation to Dr. Harlan P. Banks for his supervision and guidance during the course of the research which resulted in the discovery of a diverse flora in the Devonian of eastern New York. Special acknowledgement is extended to the National Science Foundation for the support afforded by graduate fellowships for the years 1957-1958, 1958-1959 and 1959- 1960 during which time the research was carried to completion. Thanks are extended to Dr. Rudolph Schuster for his suggestions concerning the specimens of Hepaticites and to Mrs. J. Douglas Grierson for the photographic prints used to illustrate the present paper. LITERATURE CITED Campbell, D. H. and Williams, F. ЖА А Morphological Study of Some Members of the Genus ord Pellevicinia. Stan Univ., v. Publications, Univ. Қазы is, T. M. (1926). poe опа New Method for the еси of Fossil Plants. New Phytol. Smith, с. (1938). Cryptgamic Botany, Vol. II: Bryophytes and Pteridophytes. (First Ed. ird Impresion). McGraw-Hill, New York. Walton, J. (1925). Carboniferous Bryophyta I. Hepaticae. Ann. Bot. 39:56 ——— (1928). Carboniferous Bryophyta II. Hepaticae and Musci. a. Ba. 42: 707-716. (Мог. 48 130 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE I Hepaticites devonicus Fig. 1. Portion of the thallose region ~ A = showing the differentiation of the axial region and the marginal wings. Slide Fig. Portion of the thallose region 5 E a showing a dichotomy of the axis. Note A e elongate cells of the axial region and the polygonal cells of the marginal wings. Slide 8 Fig. 3. Portion of the thallose region of the plant showing a dichotomy of the axis. x megil wings are folded over the midrib in upper portions of the specimen. Slide 9, Te 4. жаз of the thallose portion of the plant. Slide 11, Х 58. tail of the marginal wing of the thallose portion of the plant. Note the масы ый irem may have been storage or secretory cells. Slide 10, X 78. PLATE I ANN. Мо. Bor. Garb., Vor. 48, 1961 HUEBER—HEPATICITES DEVONICUS ANN. Mo. Вот. Ganp., Vor. 48, 1961 PLATE П HUEBER—HEPATICITES DEVONICUS 1961) HUEBER—HEPATICITES DEVONICUS 131 EXPLANATION OF PLATE PLATE II Hebpaticites devonicus Fig. 6. Portion of the marginal та = the thallose region of the plant illustrating the serrate margin of the wing. Slide = Y Detail of one of the раса d shown on the specimen in Fig. 6. Slide 2: Fig. 8. Fragment of the pee portion of the plant. Note the dense covering of ete rhizoids. Slide 6, X 16. . 9. Detail of the ae of the rhizomatous portion of the plant ming the быа ats of the rhizoids. The tips of the rhizoids are broken off. Slide 6, X 190. INDEX OF ORCHID NAMES — 1960 ROBERT L. DRESSLER As an aid to ourselves and others studying the orchids, we have started an index of new orchid names, to be published annually. It is felt that such an index may be of value to the many botanists and others interested in this large and complex family. For new taxa we have indicated the country or origin and the location of type material (using the standard abbreviations), if this is known. For new combinations, new names or changes in rank, we cite the basionym with the new name. We have abbreviated the authors’ names less than is customary, as we feel that a saving of only one or two spaces does not justify the loss of clarity. A few names which were published in mimeographed news letters are purposely omitted. Such publication is of dubious validity under the rules of nomenclature, and certainly should be avoided. We would be very glad to receive corrections or additions to the index, to be appended to the Index for 1961. Subfamily Cypripedioideae Garay, Bot. Mus. Leafl. 19:86. Туре genus: Cypripedium L. Subfamily Epidendroideae Dressler, Taxon 9:213. Based on Kerosphaeroideae Garay. Subfamily Kerosphaeroideae Garay, Bot. Mus. Leafl. 19:87. Туре genus: Epidendrum L. This name is not legitimate under the Rules of Nomenclature. Subfamily Neottioideae Garay, Bot. Mus. Leafl. 19:86. Type genus: Neottia Sw. Subfamily Ophrydoideae Garay, Bot. Mus. Leafl. 19:87. Type genus: Orchis L. is name is not legitimate under the Rules of Nomenclature; this subfamily must be known as the Orchidoideae. Aeranthes albidiflora Toilliez-Genoud, Ursch & Bosser, Not. Syst. Paris 16: 205. Madagascar (TAN). Aeranthes ambrensis Toilliez-Genoud, Ursch & Bosser, Not. Syst. Paris 16: 209. Madagascar (TAN). Aeranthes carnosa Toilliez-Genoud, Ursch & Bosser, Not. Syst. Paris 16: 209. Madagascar (TAN). Aeranthes denticulata Toilliez-Genoud, Ursch & Bosser, Not. Syst. Paris 16: 207. Madagascar (TAN). Aeranthes neoperrieri Toilliez-Genoud, Ursch & Bosser, Not. Syst. Paris 16: 212. Madagascar (TAN). Aeranthes orthopoda Toilliez-Genoud, Ursch & Bosser, Not. Syst. Paris 16: 212: Madagascar (TAN). Bletia purpurea var. alba Ariza-Julia & Jiménez, Rhodora 62:256. Dominican Republic. Bulbophyllum jacquetii var. rosea Guillaumin, Bull. Mus. Hist. Nat. Paris П 32: 189. Annam (P?). Bulbophyllum rugosibulbum Summerh., Kew Bull 14:138. Northern Rhodesia (K). Cephalantheropsis lateriscapa Guillaumin, Bull. Mus. Hist. Nat. Paris II 32: 188—189. Annam (P?). (133) (Мог. 48 134 ANNALS OF THE MISSOURI BOTANICAL GARDEN Chroniochilus virescens (Ridley) Holttum, Kew Bull. 14:273. Sarcochilus vires- cens Ridley Cleisostoma tixieri Guillaumin, Bull. Mus. Hist. Nat. Paris П 32:369. Annam (P?). Cypripedium X andrewsii nm. favillianum (J. T. Curtis) Boivin, Naturaliste Canadien 87:32. Cypripedium Х favillianum J. Т. Curtis. Cypripedium X andrewsii nm. landonii (Garay) Boivin, Naturaliste Canadien 87: 32. Cypripedium X landonii Garay. Cyrtorchis arcuata ssp. leonensis Summerh., Kew Bull. 14:149. Sierra Leone js Cyrtorchis arcuata ssp. variabilis Summerh., Kew Bull. 14:148. Tanganyika Terr. (K). Cyrtorchis arcuata ssp. whytei (Rolfe) Summerh., Kew Bull 14:147. Lisfrosta- chys whytei Rolfe. Cyrtorchis neglecta Summerh., Kew Bull. 14:149. Tanganyika Terr. (K.) Dactylorbiza—Soó (Ann. Univ. Scient. Budapest. Eötvös Biol. 3:335—357) gives a synopsis of Dactylorhiza (Dactylorchis) which includes many apparent new combinations for species, subspecies and hybrid populations. We do not list these here because there is not “а full and direct reference to its author and original publication” with the basionyms. Further, it is quite unclear just which names Soó intends to publish as new combinations. It seems unlikely that one would intentionally publish so many new combinations in Dactylorhiza while recommending that Dactylorchis be considered for conservation. Diaphananthe section Rhipidoglossum (Schltr.) Summerh., Kew Bull. 14:141. Rhipidoglossum Schltr. Diaphananthe brevifolia (Summerh.) Summerh., Kew Bull. 14:143. Rhipido- glossum brevifolium Summerh. Diaphananthe cuneata Summerh., Kew Bull. 14:141. Uganda (K). Diaphananthe densiflora (Summerh.) Summerh., Kew Bull. 14:143. Rhipido- glossum densiflorum Summerh. Diaphananthe erecto-calcarata (De Wild.) Summerh., Kew Bull. 14:145. Angraecum erecto-calcaratum De Wild. Diaphananthe $loboso-calcarata (De Wild.) Summerh., Kew Bull. 14:142. An- graecum globoso-calcaratum De Wild. Diaphananthe laxiflora (Summerh.) Summerh., Kew Bull. 14:141. Rhipidoglos- sum laxiflorum Summerh. Diaphananthe longicalcar (Summerh.) Summerh., Kew Bull. 14:142. Rhipido- glossum longicalcar Summerh. Diaphananthe microphylla (Summerh.) Summerh., Kew Bull. 14:142. Rhipido- glossum microphyllum Summerh. Diaphananthe obanensis (Rendle) Summerh., Kew Bull. 14:142. Angraecum obanense Rendle. Diaphananthe рейегае (Bolus) Summerh., Kew Bull 14:143. Mystacidium peglerae Bolus Diaphananthe агй (Reichb. f.) Summerh., Kew Bull 14:140. Angraecum тођти Reichb. f 1961] DRESSLER—INDEX OF ORCHID МАМЕ$— 1960 135 Diaphananthe rutila (Reichb. f.) Summerh., Kew Bull 14:143. Aérantbus rutilus Reichb. f. Diaphananthe xanthopollinia (Reichb. f.) Summerh., Kew Bull. 14:143. Aéran- thus xantbopollinius Reichb. f. Dinklageélla minor Summerh., Kew Bull. 14:156. Liberia (K). Epidendrum hioramii Асића & Alain, Mem. Soc. Cubana Hist. Nat. 24:110. Cuba (SV). Epipactis tbunbergii forma subconformis Sakata, Jour. Jap. Bot. 35:224. Japan (Herb. T. Sakata). Eulophia laurentii (De Wild.) Summerh, Kew Bull 14:139. Lissocbilus laurentii De Wild. Habenaria section Kryptostoma Summerh., Kew Bull 14:135. Type species: H. tentaculigera Reichb. f. Habenaria $onatosiphon Summerh., Kew Bull. 14:134. Tanganyika Terr. (К). Habenaria magnirostris Summerh., Kew Bull. 14:132. Tanganyika Terr. (K). Habenaria richardsiae Summerh., Kew Bull. 14:131. Tanganyika Terr. (К). Holotbrix brachycheira Summerh., Kew Bull. 14:127. Northern Rhodesia (K). Holotbrix papillosa Summerh., Kew Bull. 14:128. Tanganyika Terr. (K). Holothrix pentadactyla (Summerh.) Summerh., Kew Bull. 14:129. Deroemera pentadactyla Summerh. Laeliopsis domingensis var. alba Ariza-Julia & Jiménez, Rhodora 62:236. Domini- can Republic. Luisia acutilabris Guillaumin, Bull. Mus. Hist. Nat. Paris II 32:188. Laos (P?). Masdevallia horrida Teuscher & Garay, Amer. Orchid Soc. Bull 29:23. Costa Rica (AMES). Maxillaria imbraicata var. carinata (Rodr.) Pabst, Orquidea (Brazil) 22: 134. Maxillaria carinata Rodr. Maxillaria imbricata var. i$uapensis (Hoehne & Schltr.) Pabst, Orquidea (Bra- zil) 22: 134. Maxillaria iguapensis Hoehne & Schltr. Oberonia attenuata Dockrill, No. Queensland Naturalist 29:4. Australia (BRI). Oberonia recurva var. lin$malensis (Blatt. & McC.) Sant. & Kapadia, Jour. Bombay Nat. Hist. Soc. 57:259. Oberonia lingmalensis Blatt. & McC. Oberonia santapaui Kapadia, Jour. Bombay Nat. Hist. Soc. 57:265. Oberonia lindleyana Wight, 1851, not O. lindleyana Duperr., 1834. Oberonia trullaelabris Guillaumin, Bull. Mus. Hist. Nat. Paris II 32:115. Annam (P?). Orchis rotundifolia forma beckettii Boivin, Naturaliste Canadien 87:42. Canada DAO ). . polars densus (Lindley) Santapau & Kapadia, Jour. Bombay Nat. Hist. Soc. 57:128. Coeloglossum densum Lindley. Polystachya bella Summerh., Kew Bull. 14:137. Kenya Colony (K). Polystachya testuana Summerh., Kew Bull. 14:136. Gabon (Herb. Le Testu). Pteroceras alatum (Holttum) Holttum, Kew Bull. 14:269. Sarcochilus alatum Holttum. Pteroceras appendiculatum (Bl) Holttum, Kew Bull. 14:269. Dendrocolla appendiculata ВІ. (Мог. 48 136 ANNALS OF THE MISSOURI BOTANICAL GARDEN Pteroceras berkeleyi (Reichb. f.) Holttum, Kew Bull. 14:269. Thrixspermum berkleyi Reichb. f. Pteroceras biserratum (Ridley) Holttum, Kew Bull. 14:269. Sarcochilus bi- serratus Ridley. Pteroceras caligare (Ridley) Holttum, Kew Bull. 14:269. Sarcochilus caligaris Ridley. Pteroceras carrii (L. Wms.) Holttum, Kew Bull. 14:269. Sarcochilus carrii . Wm Pteroceras Ridley. Pteroceras compressum (Bl) Holttum, Kew Bull. 14:270. Dendrocolla com- pressa Bl. Pteroceras ecalcaratum (Holttum) Holttum, Kew Bull. 14:270. Sarcochilus ecalcaratus Holttum. Pteroceras emarginatum (ВІ.) Holttum, Kew Bull. 14:270. Dendrocolla emar- ginata ВІ. Pteroceras fasciculatum (Carr) Holttum, Kew Bull. 14:270. Sarcochilus fas- ciculatus Carr. Pteroceras hirsutum (Hooker f.) Holttum, Kew Bull. 14:270. Sarcochilus hirsutus Hooker f. Pteroceras johorense (Holttum) Holttum, Kew Bull. 14:270. Sarcochilus johorensis Holttum. Pteroceras minutiflorum (Ridley) Holttum, Kew Bull. 14:270. Asochilus minutiflorus Ridley. Pteroceras pallidum (ВІ.) Holttum, Kew Bull. 14: 270. Dendrocolla pallida ВІ. Pteroceras siamense (Ridley) Holttum, Kew Bull. 14:271. Asochilus siamensis 5. ciliatum (Ridley) Носит, Kew Bull. 14:269. Dendrocolla ciliata ey. Pteroceras stenoglottis (Hooker f.) Holttum, Kew Bull. 14:271. Sarcochilus stenoglottis Hooker f. Pteroceras sauveolens (Roxb.) Holttum, Kew Bull. 14:271. Aérides suaveolens Roxb. Pteroceras tanyphyllum (Ridley) Holttum, Kew Bull 14:271 Sarcochilus tany- phyllus Ridley. Pteroceras teres (Bl.) Носит, Kew Bull. 14:271. Dendrocolla teres Bl. Pteroceras teysmannii (Miq.) Holttum, Kew Bull. 14:271. Aérides teysmannii Miq. Pteroceras violaceum (Ridley) Holttum, Kew Bull. 14:272. Sarcochilus viola- ceus Ridley. Pteroceras viridiflorum (Thw.) Holttum, Kew Bull. 14:272. Aérides viridiflora w. Pteroceras zollingeri (Reichb. f.) Holttum, Kew Bull. 14:272. Dendrocolla zollingeri Reichb. f. Schizochilus lepidus Summerh., Kew Bull. 14:130. Mozambique (Southern Rho- desia Gov. Herb.). Schomburgkia X parkinsoniana Н. С. Jones, Jour. Barbados Mus. Hist. Soc. 28: 4. Barbados (parent species introduced). Трихзреттит pygmaeum (King & Prantl.) Holttum, Kew Bull 14:275. Sarcochilus pygmaeus King & Prantl. PRELIMINARY STUDIES IN THE GENUS STANHOPEA (ORCHIDACEAE) С. Н. DODSON anp С. P. FRYMIRE ABSTRACT This preliminary study is an attempt to bring tr ie knowledge of мечт OA i eei and Pollination of the species preparatory t mal revision of the mation, gathered by population study in the natural сея appears to be highly price to aye сабаа ng of this complex gr oup. he distribution of the species is mapped. The phylogenetic relationships of cue іле аге соп sidered and it is believed that Stanbopee i is derived from plants similar to mem of the genus Sievekingia. e development of the advanced and Miei plex species from the p» species is i i uc idered. о ain ип Observations of pollination of S. £ricornis by Eulaema meriana and of S. Machos by Poterne bomboides are sos eas and pollination relationships are indicated as the major means of g „еы path см резин е were studied in Ecuador. The advanced and variable complexes in воме TA е very suc о а area of range and density of popu ulation. Ф, is felt chat this population structure iation as a process, oP wantin variability i is maintai Various nomenclatural өлігі im are ойый вм nd a key to the species зки, A list of the recognized species is given with gel synonyms, ~ а list is appa ded E жеу Pes species which Ех аѕ уег 7% properly plac Caraway Н. Dopson, Missouri аи Garden, 2315 Tower Grove Ave., St. Louis 10, Missouri. To say that the genus Stanhopea is poorly understood is an understatement. Rather it is in a state of taxonomic chaos. Before we can say that a genus is under- stood it is expected that an individual plant, taken at random from a population of similar plants, can be fitted into a species concept. That is to say, it should be possible for a trained botanist to determine whether an individual belongs to a known species or is new and unknown. At the present time the only way to deal with most individual plants of this genus is to name them as new species, thus defeating the whole precept of taxonomy. Throughout its range of distribution Stanhopea is characterized by populations which are extremely variable. The vari- ability within a single population can be so extreme as to appear as a hybrid swarm in which are represented many kinds of Stanhopea hitherto considered as different species. Because of the plant to plant variability within a single population, a great number of species have been described from single specimens growing in European greenhouses, Often the origin of these plants was not even known. Only the fact that they were different from the other known specimens was taken into consider- ation and that difference was considered sufficient reason for a new species descrip- tion. It has been estimated that at least 500 names have been proposed in Stanhopea (Ames and Correll, 1953). This estimate is high, actually only slightly over 100 names have been published. Recent works have reduced that number to between 25 and 50 which are considered as even reasonably vali One of the problems involved in the classification of this group is that the flowers are large and fleshy and of intricate structure. They do not preserve well as herbarium specimens and many of the structures of the lip in particular are destroyed or altered in drying. The flowers are not long lasting and seldom persist on the plant for more than three or four days. They are therefore difficult to study (137) [Vor. 48 138 ANNALS OF THE MISSOURI BOTANICAL GARDEN in the field since not many flowers can be seen at any one time within a population. Their period of flowering is often extended over a period of from five to seven months and a plant which is not large will generally flower only once in a given year. Since they are difficult to study in the herbarium it has been suggested that the only way to straighten out the taxonomic confusion in the genus would be by persistent field work. Our work bears out this suggestion and we do not as yet feel qualified to publish a revision of the genus even though we have studied a considerable number of large populations. We have also studied most of the available herbarium material, but find it hopelessly unrewarding. Such large fleshy flowers as are found in this group do not, as a general rule, make good specimens when crushed and dried. Even after boiling, the critical characters are often lost and determination is difficult. Our experience indicates that dried specimens made from the same plants on different occasions are so altered in drying that they can easily be given separate species designations. This situation is particularly true in certain of the critical species groups which are separated on small differences in structure of the fleshy parts of the lip. Perhaps all is not as hopeless as it appears on the surface. A number of workers have made limited regional studies for orchid floras. A survey of their work reveals that though they do not all agree, they do not seriously disagree in respect to the majority of the species. In fact, some species appear to be remarkably distinct and do not demonstrate significant variation within their populations. Therefore they are given recognition by each of these workers when they occur in regions treated by them en the species in which the problems occur are separated from the ones which are distinct we find that the latter group is not a difficult one. Each species has a distinct distribution and to a certain degree a distinct habitat. Within the problem group the species vary tremendously yet they are in many respects remark- ably alike, particularly in morphological characteristics. We feel that this is the key to the problem. They are not only remarkably alike, they are alike and only represent variations of a pattern If these complexes were treated as representing fewer species each of which may vary considerably within a particular population, then the genus probably could be treated taxonomically and could be understood. MORPHOLOGY Let us consider just what combination of characteristics place a plant in the genus Stanhopea. The vegetative characters are surprisingly uniform, so much so that it is nearly impossible to distinguish species without flowers. All are epiphytic or lithophytic. The pseudobulbs are usually large, ovoid, ribbed and unifoliate The leaves are broad, elliptic-lanceolate, contracted into a petiole at the base and are strongly veined and plicate. The inflorescence consists of a short pendulous raceme with several broad, papery bracts. The flowers are large, long pedicellate and shielded by a papery bract, this often brightly colored and generally similar in aspect to the sepals and petals. Nearly all species produce a strong fragrance, some being agreeable and others not. The sepals are membranaceous and concave, the dorsal sepal is free and erect and the lateral sepals broad, reflexed and connate at Зар тте — 1961] DODSON & FRYMIRE—THE GENUS STANHOPEA 139 y 4 y у (у Й kn s : 2 4 m > уә с> ^ W 7 Ж ` 2 „ Ж 7774 /А 2 4 ? Fig. 1. Distribution of the genus Stanhopea. the base. The petals in most species are membranaceous, subequal to the dorsal sepal and recurved. The principal characters which separate the species are to be found in the labellum. In general, the lip consists of three parts: the hypochile or globular, concave part nearest the base of the flowers; the mesochile, which is short and often projected on each side to form horns; and the articulated epichile, which is variously shaped but usually ovate. Certain of the species, such as S. ecornuta Lehm. and S. pulla Rchb. f., simply have a saccate, entire lip which would corre- spond to the hypocile of the more complex forms. These should perhaps be con- sidered as the more primitive forms and later discussion will attempt to show why. The column is rather similar in most of the species and is elongate and rather slender with or without broad lateral wings. The anther is ventral, incumbent and two-celled. The pollen apparatus consists of a lanceolate viscidium which rests on an elongate spine-like rostellum, a stipe which is long and narrow, and the two pollinia which are elongate, quite thin, and recurved. The stigma forms a pocket under the rostellum. DISTRIBUTION The genus is distributed from central Mexico through Central America and most of northern South America into southern Brazil The plants are generally found in localities where at least some shade is afforded and where they will rarely [Vor. 48 140 ANNALS OF THE MISSOURI BOTANICAL GARDEN HOULLETIA )( NEOMOOREA ) STANHOPEA LUEDDEMANNIA POLYCYCNIS KEGELIELLA PERISTERIA LYCOMORMIUM ACENITA LACAENA SCHLIMIA CORYANTHES GONGORA ENDRESIELLA Fig. 2. А diagram of suggested generic relationships within the tribe Stanhopeinae. COELIOPSIS become dry. They can withstand prolonged periods of moderate drouth but need considerable water during their growing season. Figure 1 shows the distribution of the genus. Stanbopea lewisae Ames & Correll occurs in Guatemala and Costa Rica. Stanhopea ecornuta Lindl. occurs in Hon- duras, Guatemala, Nicaragua and Costa Rica and S. cirrhata Lindl. is found in Nicaragua and Costa Rica. Stanhopea pulla Rchb. f. is found in Costa Rica and Panama. Stanhopea grandiflora Lindl.* extends from Trinidad through Venezuela, northern Brazil and the Amazon drainage region of Colombia, Ecuador and Peru. Stanhopea rodigasiana Hort. and S. reichenbachiana Roezl. ex Rchb. f. are little known species which occur in Colombia. Stanhopea connata КІ. occurs in eastern Ecuador and Peru and S. fricornis Lindl. is restricted to coastal Ecuador and southern Colombia. The remaining species in the genus fall into two major com- plexes. The first of these groups include $. insignis Frost, $. tigrina Batem., 5. martiana Batem. ex Lindl. and $. saccata Lindl. The second group is composed of S. oculata Lindl., $. bucephalus Lindl., S. wardii. Lodd. ex Lindl., and a considerable number of lesser known forms which are quite similar. These two groups occur throughout the range of the genus from Mexico to southern Brazil; however, certain forms are restricted to geographic regions. PHYLOGENY The genus Stanhopea was taken by Bentham (1881) as the type of the subtribe and thus the name Stanhopeinae should be used in place of Gongorinae Schltr. (See Dressler and Dodson, 1960). As close allies Stanhopea has the following genera: Sievekingia, Polycycnis, Paphinia, Cirrhea and Gongora and perhaps Peristeria and Houlletia. All of the species within these genera have a labellum which is more See the section on Taxonomy for the reason for using this name in place of the commonly used S. eburnea Lindl. — Қ” а фр" =“ ай 1961] DODSON & FRYMIRE—THE GENUS STANHOPEA 141 Stanhopes oculate CGS — eS Stonhopeo tigrino ZU у : XA Stanhopes mertione Stonhopeo „с / Stonhopes soccete Stanhopea тог N, Stonhopea insignis 2, T s Stomhopeg rodigasiona ata aoa Stonbopew grocdiiore Stonhopee connate / Stonhopea lewisae м `с®— Stanhopea reichenbochlane tricornis Staphopes cirrhete \ и \ M Stonhopes ecorauto | i x, \ | n : \ | ^6 м Stonhopes puta Sitvehingio trottil Sievehingio suavis Sievekingio shepheordil A diagram of the and column of the — of ни and the closely ae VEM E suggesting о between the species. Solid lin м гос close relat species of S ships, while dashed lines represent relationships which are more "лаи ог less с ог less divided into а hypochile and an epichile. Figure 2 illustrates а possible phylogenetic series demonstrating the relationships of the genera within the Stanhopeinae. We do not feel that a discussion of the phylogenetic relationships of the other members of the Stanhopeinae would fall within the purpose of this paper and therefore will only discuss those genera which seem to be directly involved with Stanhopea in a stepwise progression from the simple to the complex. The genera placed at the center of the diagram in Figure 2 have a rather simple labellum and as the list radiates outward toward the more advanced genera the lip becomes more complicated. Figure 3 demonstrates our concepts of the origin and development of the Stanhopeas from the poorly known genus Sievekingia, a genus of about seven species. In general the plants are very similar to small immature Stanhopeas and for this reason are often neglected by collectors. Very few specimens of these plants are in herbaria and in several instances the type is the only specimen in existence. Members of this genus were not discovered until late in the 19th Century and thus did not receive so many generic names as some of the earlier discovered members of the subtribe. Without doubt certain of the species placed in Sievekingia show fewer morphological relationships to each other than do the following genera to each other: Acineta, Lycomormium, Peristeria, Coeliopsis, [Vor. 48 142 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 4. A i arcem я of the vig tas rd = Te than пред ли Lindl. by Eulaema meriana Bee the flower. EN Los Fe ckin of the fl with the spine of the rostellum e қайы i the metat thor icr eic lower w with ' th ө: viscidióm attached to the pein? чайы D. n flight with Че a Mirum in place. бА entering an Е other flower, showing the pollinia ме in the stigmatic cleft. Е, Bee а the 2nd flower, showing the pollinia left i in the stigmatic cleft. Neomoorea, Lacaena and Lueddemania. The flowers of the Sievekingias are for the most part less complex than the Stanhopeas. Sievekingia sbepbeardii Rolfe has a semierect inflorescence with rather simple flowers which have only a slightly lobed, truncate lip with a simple fimbriate callus in the center. This is certainly the most primitive known species at least in respect to the labellum. Sievekingia suavis Rchb. f. and S. peruviana Rolfe have somewhat larger, erect side lobes which form a concave hypochile at the base of the lip. From this point the close relationships of the more primitive Stanhopeas are obvious. It is a simple step from Sievekingia to Stanhopea ecornuta and S. pulla in which the lip is saccate and without the complicated mesochile and epichile of the more advanced Stanhopeas. Stanhopea cirrbata (Fig. 8) is probably an advancement from a plant similar to S. ecornuta (Fig. 6) or S. pulla (Fig. 7) through the development of lobes along the upper edge of the hypochile. In other respects S. cirrhata is rather similar to the two primitive species except for the narrow, caudate wings formed along each 19611 DODSON % ЕКҮМІКЕ--ТНЕ GENUS STANHOPEA 143 Fig 5. A diagrammatic ee of the ut prim of Stanhopea bucepbalus Lindl. by ГЕ дый bomboides Friese. A. Bee attempting to land on the waxy surface of the hypochile. B. Bee falling down through the Pissing side of the column. This may be the basic form from which the two major branches of Stanhopea have developed. On one side, S. reichenbachiana (Fig. 10) could have been derived from S. cirrhata and could well have been the form from which Stanhopea grandiflora (Fig. 11) developed. Stanhopea grandiflora has a fairly complex lip with short horns produced from the hypochile rather than from the mesochile as in many species. Stanhopea cirrhata could also be considered as similar to the ancestral form from which S. lewisae (Fig. 13) developed. Stanhopea lewisae has very small, slightly developed horns on the mesochile. Most of the advanced species with well developed horns on the mesochile may well have been derived from S. lewisae or a similar form. Stanbopea tricornis (Fig. 9) shows little relationship to the other species in the genus. The petals are thick and fleshy and are held parallel to the lip and column. The lip has a hypochile which is only slightly saccate, no true mesochile and a very thick and fleshy epichile. The horns are produced as an extension of the hypochile and a sharp spine-like callus forms a third horn between the two lateral horns in the place of a mesochile. Two possibilities can be postulated for the origin of this unique species. First, it could have developed through evolution from the other known species of primitive Stanhopea as an isolated form. Мо intermediates are known, however, between this species and the other known Stanhopeas. The second hypothesis may seem a bit far-fetched, but there is some morphological evidence for derivation of S. tricornis from a species of Sievekingia, S. trollii Mansf., recently encountered as sympatric with this S/anbopea. This Sievekingia has petals similar to S. ¢ricornis. The plant is so similar that it doubtless has been passed over by previous collectors (including ourselves) as a young Stanhopea. A colored slide taken of this species, without a scale to indicate size, was at first (Мог. 48 144 ANNALS OF THE MISSOURI BOTANICAL GARDEN determined as a highly floriferous form of $. £ricornis. The lip of this Sievekingia is saccate at the base, has two side lobes which flare outward and an epichile very similar to S. £ricornis. The callus of the lip is in some respects similar to the center horn of S. ¢tricornis. It seems feasible that S. £ricormis may have been derived by way of this species of Sievekingia. Stanhopea tricornis may possibly have been derived as a result of hybridization between Sievekingia trollii or a similar form and some Stanhopea sympatric with it in distribution. Stanhopea bucephalus, $. tricornis and Sievekingia trollii are sympatric along the western slopes of the Andes in Ecuador. If chance pollination of Stanhopea bucephalus by the Sievekingia should occur the resulting intermediate hybrid might well be visited by a pollinator not utilized by either parental species and thereby be stabilized as a species. When an attempt is made to draw an intermediate between the morpho- logical characteristics of the Sievekingia and Stanhopea bucephalus the result is surprisingly similar to the labellum of S. £ricornis. Stanbopea rodigasiana (Fig. 12) is unique in the genus. The strictly one- flowered inflorescence, ovoid, smooth pseudobulbs, bifid rostellum, round viscidium and unique horns on the lip are characters not found in any other Stanhopea. In some respects it suggests a possible relationship to Paphinia, but it is certainly close to Stanhopea. It is surprising to us that this species was never placed in a mono- турс genus. Although it is obviously closely related to Stanhopea it is sufficiently different from the other species in the genus to allow any taxonomist with a narrow generic concept to separate it without qualms of conscience. The reason for its not having been so separated is very likely that it was not discovered until after the turn of the century, when more care was being taken in such matters. If it had been discovered 50 years earlier when Peristeria, Acineta, Lycomormium, and other closely allied genera, were being separated, it would surely have been considered as generically distinct. We do not believe that it warrants special distinction nor do we believe that so many genera should be recognized in the Peristeria complex. We hope that through more careful study of the entire subtribe Stanhopeinae we will be able to develop a useful revision of its members. The above mentioned species of Stanhopea are for the most part easily recog- nized, have distinct morphological characters and distributional patterns. They present no serious problems to the taxonomist. The remaining species within the genus form the difficult portion. All of them have a more complex labellum than the species previously discussed. The hypochile is saccate and of various sizes but it follows a similar pattern in all of the species. The mesochile is short and inserted on the apex of the hypochile with two lateral, elongate horns of various sizes. The epichile is articulated to the apex of the mesochile and while it may vary in size and proportions, they are all similar. Two major groups can be separated and while we are not yet prepared to present a formal revision of the genus we feel that some designation is necessary in order to be able to group the excessively large number of species names in such a way that they can be discussed with relative ease. We will use the terms “oculata complex” for those plants similar to S. oculata, and "insignis complex" for the plants allied to S. insignis. The "oculata complex” will therefore comprise those concepts which have a rather long narrow or rectangular 1961] DODSON & FRYMIRE—THE GENUS STANHOPEA 145 . 6. Stanbopea ecornuta Lem. A. Inflorescence (X V5). B. Plant habit (X 16). C. Side view 52 the lip and column (Х 12). D. Dorsal view of the lip (X 1). Е. Pollinarium (Х 3). hypochile, large mesochile horns and always have an entire, acute or apiculate epichile. The “oculata complex” comprises such taxa as: $. wardii Lodd. ex Lin (Fig. 21), $. oculata (Lodd.) Lindl. (Fig. 19), $. bucephalus Lindl, (Fig. 20), S. haseloviana Rchb. f., S. platyceras Rchb. f., etc. The “insignis complex” will consist of those forms which have a saccate or globular hypochile, large mesochile horns and either entire, acute or three-lobed epichiles. The “insignis complex” comprises such species as $. tigrina Batem. ех Lindl. (Fig. 16), $. martiana Batem. ex Lindl. (Fig. 17), S. insignis Frost (Fig. 15), S. saccata Batem. (Fig. 18) and S. devoniensis Lindl. Stanbopea quadricornis Lindl., if such a species exists, has a lip which is within the variation pattern of S. oculata with the exception that the base of the thicken- ings of the margin of the hypochile are projected into short horns. Apparently only one specimen was collected and was named by Lindley. It is quite possible that it was only an aberrant specimen or a hybrid and was not representative of any population. It seems apparent that S/anbopea is a rather terminal group in evolution, with Coryanthes, a genus often said to be closely allied to Stanhopea, having a derivation [Vor. 48 146 ANNALS OF THE MISSOURI BOTANICAL GARDEN .7. Stanhopea pulla Rchb. f. А. Inflorescence (X А ). В. Plant habit m Мо). C. Side view x the lip (X 1). D. Dorsal view of the lip ( 1). E. Pollinarium (Ж from some similar ancestor, but one probably more closely allied to Peristeria than to Stanhopea. Cirrhea and Gongora appear to be the termination of a separate line. In general, the Orchidaceae appears to be a rather recent but rapidly evolving family in which less extinction is evident than in older families. Such a situation often makes classification difficult due to the intergrading forms, but on the other hand, it makes the tracing of phylogeny easier since closely related groups are still present. We do feel that the Stanhopeas were derived from ancestors very muc like some of the species of Sievekingia which are known today. The species of Sievekingia are not particularly successful, as is evident from their extreme rarity, but may be considered as relict forms locally preserved in relatively hospitable habitats. Such forms would very likely become extinct and be removed from the record if they were members of plant families subjected to more stringent environ- mental conditions. POLLINATION This paper is perhaps somewhat premature in one respect, that is, the pollinators and pollination mechanisms are not as yet known for all the species. However it may be many years before all the information can be gathered. We feel that the major worth of the paper lies in its discussion of the problems within the genus. The pollination mechanisms and pollinators of two species have been well worked out and photographed. Pollination studies of the other species will be described when they are known. We have witnessed the pollination of both Stanbopea tricornis and S. bucepbalus in Ecuador. A future revision of this genus may well be based in large part upon the ideas gained from observations of pollination. Polymorphic genera are often difficult to understand and when confusion is compounded by having highly variable popula- tions, the naming of which has been multiplied on an unrealistic basis, such a genus —" ВА S." 1961] DODSON & FRYMIRE—THE GENUS STANHOPEA 147 . 8. Stanbopea cirrhata Lindl. A. Inflorescence (X V4). B. Plant habit (X Ит). C. Side view e the lip (X 24). D. Dorsal view of the lip (X 24). becomes chaotic. Occasionally new techniques or observations from a different standpoint, can provide flashes of insight for the investigator. The totally different pollination mechanisms of these kinds of Stanhopeas, when observed, immediately and clearly made obvious certain of the reasons for the polymorphism within the genus. The first system is typical of most orchids in which the pollinator simply enters the flower after landing on the expanded labellum. The transfer of pollinia Occurs as a result of proper positioning of the column and its sexual apparatus in relation to the size and conformity of the particular pollinator. The majority of the species of Stanhopea are of this type. These species are not extremely variable and are easily recognized. The second is a highly complex system in which the flight characteristics of the bee are used in the pollination of the flower. After landing, the bee is clumsy and has difficulty in regaining balance in flight. The bee, unable to grasp the slick surfaces of the pendant labellum, falls through the flower. This characteristic flight habit of the bee probably has contributed to the evolution of an entirely different system of pollination. Though this kind of flower is distinct, it apparently has been unable to preclude occasional hybrids within the related forms but has simply led to more variation. Color and morphology of the unessential flower parts appear to have been subject to little selection pressure for uniformity, and may vary considerably from plant to plant within a given population. The resulting confusion, caused by attempts to name every variant in cultivation, has not only increased out of all proportion but has tended to make the genus nearly impossible taxonomically. Stanhopea tricornis occurs frequently in old epiphyte covered cacao trees in the region of Quevedo in central, coastal Ecuador. The species generally succeeds best in fairly dense shade and on a large plant the pendant two-flowered racemes are pro- duced throughout the year. Several racemes may be produced by one pseudobulb. (Мог. 48 148 ANNALS OF THE MISSOURI BOTANICAL GARDEN In late January of 1960 we succeeded in finding a plant which had two open flowers (both had been pollinated) and two mature buds which would open the following day. We removed the plant and carefully carried it to our hotel room. Before dawn of the next morning we found the flowers to be open and emitting a heavy perfume very similar to "Diorissimo" manufactured by the Christian Dior Company of Paris, France. We returned to the cacao plantation at dawn and set the plant in a position where it could be easily photographed. At about 9 a.m. a large male bee, Eulaema meriana (Oliv.) (tribe Euglossini, family Apidae, E. dimidiata is a syno- nym) came to the flowers, which he proceeded to investigate, apparently searching for the source of the strong fragrance. Meanwhile a second bee of the same species arrived; neither bee seemed familiar with the flower and each searched for some time before discovering the tunnel-like opening, formed by the petals, column and lip, and entering the flower. It might be noted that this is the only species of Stan- hopea in which the petals are fleshy and held closely appressed to the column. The bee forced upward past the rostellar spine and entered the inner portion of the flower to the hypochile (see Fig. 4). The bee did not attempt to extract food. In backing out the bee encountered the rostellar spine once again. This time the rostellar spine slid under the posterior projection of the metathorax of the bee, and his struggles dislodged the lance-shaped viscidium of the pollinia, which immedi- ately became cemented to the underside of the metathorax. In further backing out he pulled the pollinia from the anther cap. The bee then flew away. Even if the bee had entered the other flower, pollination could not have been accomplished immediately for the stigmatic pocket of Stanhopea does not open sufficiently for insertion of pollinia until several hours after the pollinia have been removed. The following morning what appeared to be a different bee but one with pollinia attached to his metathorax visited the flowers. The pollinia became lodged in the stigma and were removed from the back of the bee, thus effecting pollination. In total, five instances of pollination of Stanhopea tricornis were observed with six bees, all E. meriana, involved. Other species of Eulaema were noted in the immediate vicinity and were quite curious about the camera, approaching to examine it, but showed no interest in the Stanhopea tricornis flowers only а foot or so away. These species were E. fropica (Linné) and E. cingulata (Fab.)* both of which are responsible for pollinating numerous other orchids including several species of Catasetum, Cycnoches, Sobralia violacea Lindl, Maxillaria grandiflora Lindl., Aspasia epidendroides Lindl. and Pescatoria wallisii Linden & Rchb From these observations we can glean the following important points: 1. Eulaema meriana appears to be the specific pollinator of 5. tricornis. 2. Eulaema tropica and E. cingulata are sympatric but showed no interest in S. tricornis. j 3. Eulaema meriana is attracted to the fragrance emitted by S. tricornis and without prior experience with the flower must search to find the entrance. * 'These identifications have been recently supplied by Dr. Krombein of the Smithsonian Institution. These two species were previously known as E. musitans and E. fasciata respectively. 1961] DODSON & ЕКҮМІКЕ--ТНЕ GENUS STANHOPEA 149 Fig. 9. Stanhopea tricornis Lindl. a. Inflorescence (X V4). b. Plant habit (X Мә). Side view a the lip (X уз). d. Ventral view of the column (X У). е. Dorsal view of iri lip 1$). F. Pollinaríam Ox IE Flower color and structure are certainly secondary if even of importance in the actual attraction of the Flowers of S. tricornis were carefully examined for nectaries or other anatomical structures which would offer food for the bee. None was found. The fragrance is produced in the hypochile; the other parts, when removed from the flower, have little or no fragrance. The inside of the hypochile is covered with long fleshy hairs which the bee might eat but in the flowers entered by E. meriana these hairs did not appear to have been disturbed. It appears then, that the bee is attracted strictly on the basis of fragrance and does not receive food. He apparently does not expect to receive food for he does not lower his tongue, which is so long that it must be lowered and swung forward before a narrow flower is entered. This pattern has been observed in other members of the Catasetinae and Stanhopeinae in which male Eulaemas are attracted by the fragrance and the bees do not appear to receive food of any kind. In all of these orchids the flowers are generally not long lasting but produce their strong, heady, spicy fragrance in such quantity that they can be detected by (Мог. 48 150 ANNALS OF THE MISSOURI BOTANICAL GARDEN the human nose, a notably weak olfactory organ, at distances of more than thirty feet. The bees are usually quite wary when they first come to the flowers but after being around them for some time seem to become drugged for they fly erratically and can be approached easily. Only the male bees of Eulaema appear to be attracted to this type of flower; the females, which are busy with nest building and food gathering to provision the nests, do not approach these flowers. "Тһе male bees have never been observed taking part in nest building or care of the nest and seem to leave the nest as soon as possible and live a vagabond life. They secure food from many of the same flowers as the female bees but also visit Stanhopeas, Cata- setums, Cycnoches and other flowers which offer only the heady and seemingly intoxicating fragrances. It is possible that the female bees are not attracted to these flowers because they cannot sense them. It is known that female bees of Apis mellifera, the domestic bee, have fewer olfactory pits in their antennae than do the males. The only orchids in which both male and female bees were noted were Sobralia violacea and Maxillaria grandiflora. Sobralia is a rather primitive orchid which does provide nectar for the bee and which is visited by several bees of different families. Maxillaria grandiflora provides starchy hairs at the base of the lip. One morphological feature found in the male bees of the Euglossini which is not present in the females is the presence of chemoreceptive hairs on the tarsi of the front legs. The male bees brush these pads through the scratched surface of the labellum of the orchids which they visit. Stanbopea bucephalus is pollinated іп a somewhat different manner. This vari- able species is distributed along the western slopes of the Andes from elevations of about 400 meters up to 1200 meters and is to be found on the summits of the coastal mountains where they rise to these elevations. It is sympatric with $. £ri- cornis in the region of Quevedo and the two species appear to be sympatric for at least 60 kilometers to the south. Pollination of this species was observed during the month of March at Olimpo on the railroad from Guayaquil to Quito. Unlike S. tricornis this species has thin, narrow petals which became retracted and curl up around the dorsal sepal shortly after the flower opens. The horns are not produced as an extension of the hypochile as in S. fricornis but are larger and developed from a definite mesochile. Thus there is left a large open space between the lip and the column at the level of the hypo- chile. The lower part of the column is effectively surrounded, to form a tunnel, by the horns of the mesochile, the epichile and the wings of the column. The pollinator of this species is also a male Eulaema, E. bomboides Friese, but this one is only about two thirds as large as E. meriana. Eulaema bomboides is a little larger than either E. cingulata or E. tropica. Eulaema bomboides is attracted by the fragrance of this Stanhopea which is honey-sweet, quite unlike S. £ricornis. The bees come to the flower of S. bucephalus but immediately fly into the space between the column and the hypochile and attempt to land on the hypochile, or they land on the side of the hypochile and attempt to crawl around and enter the hypochile to reach the source of the fra- grance. The surfaces of all parts of the lip are waxy and extremely slick, so that the bee can not successfully land and hold fast. He falls down and is guided 1961] DODSON & FRYMIRE—THE GENUS STANHOPEA 151 Fig. 10. Stanhopea reichenbachiana Roezl. ex Rchb. f. a. Inflorescence CX у). b. Plant habit (X Мо). c. Side view of lip (X 14). d. Dorsal view of the lip (X %). by the column and its wings at his back, the horns of the mesochile at his sides and the slick epichile beneath him so that he falls directly through the tunnel. At the lower end of the tunnel, almost at its exit, the rostellar spine with the viscidum of the pollinia on its outer surface projects into the path of the falling bee and is caught under the posterior projection of his thorax as in S. fricornis. The bee hits the rostellar spine with such force that it gives way and allows the bee to fall through. In doing so the viscidium of the pollinia is cemented firmly into place under the thorax (see Fig. 6). The bee is apparently disturbed and tends to fly away but he repeats the process when he encounters another flower. If the flower is older with an open stigma the pollinia are this time placed in the stigmatic pocket during his fall and pollination is effected. The pollination of S. bucephalus is in many respects quite similar to the pollina- tion of Gongora maculata Lindl. as reported by Allen (1956), with the exception that the Gongora is pollinated by species of Euglossa. Euglossa is a genus of bees which is very closely related to Eulaema and is also responsible for the pollination of Coryanthes (Allen 1950). In Gongora the bee, hanging upside down loses its hold on the slick surfaces of the lip, falls and is guided down the pendant column during his fall with the result that he also makes contact with the viscidium of the pollinia much in the same manner as Eulaema in S. bucephalus. Several plants of this Stanhopea were brought back to the garden of the Insti- tuto Botánico where quantities of Eulaema tropica and E. cingulata are common and at no time did any of these bees show interest in the flowers. Even when flowering plants of Catasetum were placed in the immediate proximity the Eulaemas ignored the Stanhopea flowers completely. (Мог. 48 152 ANNALS OF THE MISSOURI BOTANICAL GARDEN From these observations we can add the following facts: — . Ещаета bomboides appears to be the specific pollinator of $. bucephalus in the region studied. ~ . Stanhopea bucephalus is pollinated on ап entirely different principle from 5. tricornis. > . Eulaema bomboides is attracted by the fragrance of 5. bucephalus which is different from the fragrance of S. £ricornis. ^ If E. bomboides does not visit S. tricornis then S. tricornis and S. bucephalus are effectively isolated genetically. The last point is by far the most important. Eulaema meriana is too large to fall through S. bucephalus. If E. bomboides is not attracted by the fragrance of S. tricornis then the possibilities of the two species of Stanhopea forming hybrids are effectively nil. This appeared to be the case. Stanhopea tricornis shows very little variability throughout its populations. The sympatric species $. bucephalus is quite variable but shows no characters indicating introgression from S. fricornis. If $. tricornis evolved in this region with E. meriana as its pollinator and S. bucephalus was dependent upon a different insect pollinator and a different pollina- tion procedure the two would not hybridize. If by accident a bee of E. bomboides did enter S. tricornis and effect a hybridization between the two species of Stan- hopea, and it is feasible that this might occur, then the resulting hybrids would probably find no effective pollinator. If such a pollinator did exist then the chances might be better for development of a completely different, stable species with its own characteristics and its own structurally isolated population. Interestingly enough, perhaps not all Stanhopeas are pollinated by members of the genus Eulaema. The type locality of Euglossa viridissima Friese is given as "at Stanhopea tigrina at Cordoba and Orizaba in Mexico”. If this report is correct, and there is no reason to doubt its validity, it indicates that perhaps some of the species are even more widely separated ecologically than are S. fricornis and S. bucephalus. Our observations indicate, at least in Ecuador, that Euglossas and Eulaemas are not mutually attracted to the same orchid flowers. Where they are attracted to orchid species of the same genus (i.e., Eulaema cingulata is attracted to Cycnoches lebmanii and C. ventricosum while Euglossa viridissima is attracted to C. egertonianum) the flowers are extremely distinct morphologically. The prin- cipal morphological difference between S. tigrina and S. bucephalus which would be of importance in pollination procedure lies in the extreme closeness of the column and terminal portions of the lip in the former species whereas in the latter these structures are rather open. А large bee like a Eulaema would not be able to fall through S. tigrina due to this constriction of parts. On the other hand a Euglossa would fall through $. bucephalus without touching any of the parts and would be ineffectual in pollination. We feel that the significant factor in this situation does not lie in the morphological form being a deterrent to effective pollination, but rather that the fragrance of the flower does not attract a pollinator which would be ineffective. More likely the morphological structures of the flower have been modified to accommodate the type of pollinator which was attracted to the flower. E 1961] DODSON & FRYMIRE—THE GENUS STANHOPEA 153 view of Fig. 11. Stanhopea ОСМ (Lodd.) Lindl. a. Inflorescence (X Уз). b. Plant habit (X %). с. Side E of thelip (X 14). d. Dorsal view of the column (X x. e. Dorsal v the lip (X 14). ин (3) [Vor. 48 154 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 12. Stanbopea rodigasiana Claes. ex Cogn. Inflorescence (Ж 14). b. Plant habit (X c. Side view of the lip (X М). d. Doral « view of the lip (X 14). e. Ventral view of the columns а). 3 Pollinarium Ox 2. VARIATION We have pointed out that populations of species in Stanhopea are often ex- tremely variable. Unfortunately no extensive analysis of specific populations are yet available; however, we did superficially study two large populations in Ecuador. e first occurs along the western slopes of the Andes from Macuchi in north- western Ecuador to Paccha in the south near the Peruvian border. This population is approximately 400 miles long and ranges from 400 to 1,200 meters in elevation. ear Macuchi some of the plants correspond to a species named by Rolfe as S. anfracia. This species is very close to S. wardii and had been considered by Lindley as S. wardii var. venusta. Schweinfurth (1958) has placed it in synonymy with S. wardii. Other plants are similar to what has been known as S. bucephalus. The population was sampled at four locations through its central portion and was rather uniformly S. bucephalus with only occasional plants occurring which could be considered as referable to S. wardii. However, at Santa Isabela near the southern extension of the population the situation changed considerably. Plants were so 1961] DODSON & FRYMIRE—THE GENUS STANHOPEA 155 , Fig. 13. Stanhopea lewisae Ames & Correll. a. Inflorescence (X 15). b. Plant habit (X lig). c. Side view of the lip (X 24). d. Dorsal view of the lip (X 34). c. Ventral view of the column (X 24). variable that specimens were found which corresponded precisely to $. anfracta, S. bucepbalus, S. graveolens, S. oculata, S. wardii, S. baseloviana, S. guttulata, S. ornatissima, S. platyceras, and S. peruviana. Many plants also occurred in the population which were intermediate between these concepts. Most of these species are actually color forms and many have been relegated to synonymy by other authors. Some of them have been maintained, largely because no botanist knew much about them, for they were described from individual specimens cultivated in Europe and their origins were unknown. Plants from the population at Paccha, about 20 miles south of Santa Isabela were sent to Europe by a number of early collectors. Kruth named a plant which Humboldt and Bonpland had collected here as Epidendrum grandiflorum. Lindley later named another plant from the same locality as S. bucepbalus and it was illustrated in the Botanical Register, plate 24, in 1845. Reichenbach changed the name to $. grandiflora, based on Kunth's name. Another illustration of a plant in the Botanical Magazine, Plate 5278 given as S. bucephalus is so inaccurate that four (Мог. 48 156 ANNALS OF THE MISSOURI BOTANICAL GARDEN "species" can be identified from the same plate. Plate 8517 in the Botanical Maga- zine is actually a plant much nearer S. wardii. More recently S. bucephalus has been reduced to synonymy under S. oculata. This confusion has pointed up the results of describing species from individual specimens without knowledge of the variation within populations. We feel that this situation is not uncommon and clearly demonstrates the need for careful and extensive population studies of the genus. A second variable population of Stanhopea occurs in the valley formed by the headwaters of the Zamora river on the eastern slopes of the Andes in southern Ecuador. The population is extensive and we feel that it demonstrates another of the kinds of phenomena involved in variation of the genus. The upper end of the population reaches a point near Km. 45 on the road from Zamora to Loja, at an elevation of about 1800 meters. The valley is narrow, dense fogs and constant rains are the rule and temperatures are usually low, around 60° F. Normally the sun breaks through only for an hour or two each day but occa- sional days are sunny. When the sun is out for several hours the temperature rises to as high as 75? Е. The elevation decreases rapidly to the east and at Zamora, 62 km. from Loja, is about 1000 meters. At Zamora the valley levels out and becomes broader, the normal daily temperature is around 75° F. and occasionally reaches 85° F. The weather conditions change little during the year but less rainfall occurs in November and December. The population was studied as far as Yansasa, about 60 km. northeast of Zamora. Here the elevation is near 800 meters and weather conditions are similar to Zamora. In all, the portion of the population studied extended 85 km. and varied from 1 km. in width at the upper end to 3 km. wide at the lower end. Elevations run from 1800 meters at the upper end to 800 meters at the lower end. Though they are variable in color at the lower end of the population the flowers correspond to Stanhopea anfracta Rolfe. The flowers of plants at the upper end are similar to the species described by Schlechter as $. peruviana and demonstrate little variation. Both of these species have been reduced to synonymy under $. шағай. A cline occurs from the upper end of the population to the lower, the plants in the center having flowers essentially intermediate between the extremes. Variation increases considerably toward the lower end of the population. Vege- tatively the plants are indistinguishable throughout the population. The flowers of the plants at the upper end of the population are morphologically nearly identical to the flowers at the opposite extremes but are colored quite differ- ently and are much thicker in substance. The flowers are completely tangerine- yellow with no other color involved. The sepals and petals have heavy texture and are not reflexed. The flowers usually last for eight days. At the lower end of the population the flowers are thin in substance and have their sepals and petals strongly reflexed. The basal part of the lip, sepals and petals are yellow-orange flecked and spotted with maroon-red, and the extremities of the sepals, petals and lip are white flecked with red. The flowers last only one or two days. Plants from around Zamora had flowers more or less intermediate between the two extremes in all characters. 1961] DODSON & FRYMIRE—THE GENUS STANHOPEA 157 . Stanhopea connata Rchb. f. Inflorescence (X 14). b. Plant habit (X 1$). c. Side view of the lip (X 24). d. Dou 9 view эт the lip ^" V2). e. Dorsal view of the column (X №). Е. Pollinarium (X 2). From Zamora to Yansasa bees of the genus Eulaema were very common. One species in particular was quite common and appeared much like E boides. They were of the same size but slightly different in color pattern ad probably represent E. nivofasciata Friese. Though none was actually observed pollinating Stanhopeas it is probable that they аге the pollinators. Eulaema cingulata was also common and while they visited other flowers placed in proximity they showed no interest in Stanhopeas and would not enter them. The bees of this genus are partial to the warmer tropical climates. They nest in hollow limbs or in holes in embankments and their nests have not been encoun- tered at elevations above 1,000 meters in Ecuador. From Zamora to Yansasa they are commonly observed visiting Costus, Inga and Bixa. They are fast flying bees and are capable of long flights. They are seldom seen in the upper end of the valley and then only on the rare, sunny, warm days. It would appear that they do not nest in the upper extensions of the valley. Plants in the lower regions com- monly produce one or two seed capsules indicating considerable visitation by the in- sects. In the upper regions, on the other hand, seed capsules are rarely encountered. [Vor. 18 158 ANNALS OF THE MISSOURI BOTANICAL GARDEN , Fig. 15. Stanhopea insignis Frost. a. Inflorescence (X V4). b. Plant habit (X Иг). c. Side pid 5 the lip (X 1). d. Dorsal view of the lip (X У). е. Dorsal view of the column 2). This population in some respects appears rather typical of the populations of Stanhopea in Ecuador. It is not as complex as the population of Stanhopea on the western slope of the Andes which is extremely variable in the direction of $. wardii in its northern portion, rather stable in its central portion corresponding to S. bucephalus and mixed with typical $. oculata in its southern extension. It is our opinion that this type of variation is typical of $. oculata and allied species through- out their range. It would appear that the relatively stable, heavy substanced, long lasting type of flower at the upper extension of the Zamora population may be due to strong natural selection for that type of flower. In the upper elevations where pollinators are not abundant it would be of more value to the population to have long lived flowers which would have a greater possibility of visitation and pollina- tion. Such flowers may have to wait several days for weather conditions to be satisfactory for bees to reach them. Short lived flowers would stand little chance of being visited and therefore would rarely reproduce. Long life in these flowers depends on thickness of their parts. A selection pressure is therefore exerted for 1961] DODSON & FRYMIRE—THE GENUS STANHOPEA 159 Fig. 16. Stanhopea tigrina Валет. a. Inflorescence (X 14). b. Plant habit (X 1$). c. Side view S the lip (X V2). d. Dorsal view of the lip (X М3). е. Dorsal view of the column X »2). flowers with heavy substance. In the lower regions flowers are normally visited shortly after opening and long lived flowers would offer no selective advantage. Some botanists would attempt to attribute such extreme population variability as is found in Stanhopea to hybridization and introgression. Perhaps this is a partial answer but in many cases the variable populations are so widespread as to preclude hybridization. One might also suggest that the variation is due to high mutation rate. Perhaps this is also a partial answer. It is apparent from what has been pointed out above in the pollination section of this paper that the bees which accomplish the pollination are attracted by the strong fragrance of the flowers. The color, and to a degree, the form of the flower appears to be of minor impor- tance in its pollination. This is also true in the genera related to Stanhopea, such as Gongora and Coryantbes. [Vor. 48 160 ANNALS OF THE MISSOURI BOTANICAL GARDEN = 17. Stanhopea пати Batem. ex Lindl. a. Inflorescence ( x pond of the Пр (X V2). d. Dorsal view of the lip (X 14 ) e. Dorsal view o L4). b. Plant ee + ). ). Min CX 1 If the attraction of the fragrance is so strong, color may be a minor factor in selection and the bees will still visit the flowers even if color varies widely. Also form could vary widely in parts which are not essential to the actual mechanisms of pollination, allowing variants due to mutation to remain within the population as fully effective breeding members contributing to the gene pool as a whole. In S. oculata and its allied species the characters which vary the most are the form of the hypochile of the lip and the color of the lip, sepals and petals. These are char- acters which are not actually involved in the mechanisms of pollination, and may not be important in the actual attraction of the pollinator. Selection would not be strong for these characters and a plant which deviated from the norm to a consid- erable extent could still remain a breeding member of the population as long as it retained the character of fragrance to attract the pollinator and the correct propor- 1961] DODSON & FRYMIRE—THE GENUS STANHOPEA 161 Fig. 18. S/anbopea saccata Batem. a. Inflorescence (X 14). b. ze = AG %). с. Side view ol the lip (X 12). d. она я view of the ір (X $. e. of the column (X уз). tions of the tunnel of the lip, formed by the mesochile and column, to properly place the pollinia for transport by the bee to another flower. The fact that cross-pollination is nearly obligate in these genera may be of importance also. Certainly more variation would be produced, maintained and ispersed in a population of obligate cross-pollinating plants than in a population where self-pollination is possible and often of regular occurrence. Certain of the genera in the Catasetinae, і.е., Catasetum and Cycnoches, are characterized by unisexual flowers which make cross-pollination obligate. These are genera in which the species are notoriously variable. In many of the other genera self- [Уог. 48 162 ANNALS OF THE MISSOURI BOTANICAL GARDEN incompatibility is the rule. Not a great deal of information is available on this phenomenon but the authors have made numerous attempts to self-pollinate Stanhopea bucephalus and Gongora maculata with no success. The resulting seed pods dropped after a few weeks. In most of these variable genera the flowers are highly fragrant but extremely short lived, seldom lasting more than two or three days. The fragrance ceases very soon after the flower has been pollinated. It would appear that there is a correla- tion involved between the characteristics of powerful fragrance and short lived flowers. These species seem to depend on their fragrance to attract pollinators and are usually very successful, for a considerable portion of the flowers are pollinated. They have developed in the direction of stronger and more attractive fragrances but offer no food or nectar to the pollinator. Selection may even favor short-lived flowers which do not attract bees long after their pollination (i.e., do not compete with the younger flowers). Неге there seems to be a strong selection pressure for fragrance for pollinator attraction with lower selection pressure for constancy of morphological characters. Many of the orchids which have long lasting flowers are either not fragrant or are weakly fragrant. These orchids seem to depend on numbers of flowers few of which are ever pollinated. These species for the most part are not as variable as the fragrant types and have developed other mechanisms for attraction of the pol- linator. Where populations of these species are extremely variable, it is usually in a relatively limited geographical area and their variability can often be attributed to introgressive hybridization. SUCCESSFULNESS OF THE GENUS The objective measurement of success in a group of plants is difficult, as too many factors are involved. Obviously, so called weedy species would rate high on the scale. However, to attempt to rate other plants as being highly successful or only fairly successful is scarcely possible. We do feel, however, that we can sub- jectively rate the success of a group of orchids in relation to other orchids. In such a subjective sense we can use extent of distribution and density of populations as measuring tools. Many orchids are highly successful in respect to density of population but are limited in distribution. Others have wide distributions but are seldom encountered in quantity in a given area. The majority of the species of Stanhopea are in one or the other of these categories. The “insignis and oculata complexes” on this basis are highly successful in that they are commonly found in quantity (in undisturbed habitats) and have broad ranges. They are to be encountered in any region of the American tropics which provides the basic needs of the plants. They are found from sea level to 7000 feet and grow as well in either location provided there exists sufficient humidity and shade and the temper- atures are not too low. This very success, we feel, contributes to their variability, but on the other hand, may have suppressed speciation. The extremes of the range may be expected to produce forms which are unlike due simply to spatial relations. However isolation is difficult to achieve in a dense and essentially continuous population. In 1961] DODSON & FRYMIRE—THE GENUS STANHOPEA 163 ПРОЗУ > К di nam D “Њу соба 2 = МЕЈ Қз = . 19. Stanbopea oculata (Lodd.) Lindl. a. с. Side view of the lip (X 24). d. Dorsal view of the lip (X L2). e. Pollinarium Inflorescence (X 14). b. Plant habit (X Vis). i (X 2). this respect "success" and taxonomic diversity may be mutually incompatible. This situation is by no means limited to S/anbopea. The hooded types of Cata- setum reflect the same phenomena, though not quite to the same degree. In Mexico and middle Central America a form known as C. integerrimum Hook. is common and variable. Further south in Costa Rica and Panama intergrading populations known as С. viridiflavum Hook., C. oerstedii Rchb. f. and C. maculatum Kunth occur. From Panama extending through Venezuela, the Guianas and northern Brazil the form known as C. macrocarpum L. C. Rich. is found. In Ecuador C. macroglossum Rchb. f. is common. All of these populations are highly variable and in Ecuador alone, specimens corresponding to all of these "species" may found. There is little reason to recognize these species when they are now known to be only representatives of one widely distributed and highly successful popula- (Мог. 48 164 ANNALS OF THE MISSOURI BOTANICAL GARDEN tion. Here, as in Stanhopea, speciation seems to be inhibited by the otherwise highly successful population structure. Gongora maculata Lindl. demonstrates a close parallel. Several species of Cycnoches, Mormodes, Coryanthes, Epidendrum and Oncidium have similar pat- terns. Many other instances of highly successful, widely distributed orchids can be given in which the most striking chracteristic is their extreme variability. We feel that speciation has taken a back seat to success in the sense that development of distinct taxa is inhibited where gene flow is unrestricted and selection is weak. TAXONOMY Perhaps the most successful method of handling the great variability in Stan- hopea will be to follow that used by Allen (1952) in his particularly workable and useful revision of Cycnoches. Cycnoches reflects the same s of problems encountered in Stanhopea and by placing several closely related, intergrading populations, which had been previously known as species, as subspecies under one inclusive species, Allen was able to indicate the close relationships of these popula- tions. In our opinion this is the most practicable manner to handle the problems of such extreme variability in obviously closely related populations. Certainly, it is folly to attempt to apply species or even subspecies names to all of the variants found; the number would be endless. There may be some difference of opinion as to how many subspecies should be recognized, some authors suggesting more and others less. When numerous well authenticated population studies are accomplished in Stanhopea it is quite possible that certain of the dubious species within the “insignis and oculata complexes” will be found to exist as discrete populations and will be sufficiently distinct to be recognized, others will probably be reduced. Recent correspondence with a rather well informed amateur orchid enthusiast indicates that he has encountered a stable population of a Stamhopea. Specimens which he has kindly forwarded appear near the type form of $. oculata. He main- tains that these are different from typical $. oculata, are stable within what he knows of the population and should therefore be recognized as a new species. Perhaps with exhaustive study this population might be found to be distinct throughout its extent and should therefore be considered as a valid species. Our experience with populations of Stanhopea in Mexico, Guatemala, Costa Rica, Panama, Ecuador and Peru would indicate that this population, throughout its entire range, would very doubtfully represent even a valid subspecies of S. oculata. is case, however, is quite representative of the problems which S/anbofea offers and indicates the necessity of thorough knowledge of large percentages of over-all populations before rash action is taken in applying more epithets in the group. The nomenclature of Stanbopea is badly confused and the efforts made by modern workers to clarify the situation have not always been particularly successful. Many of the early descriptions of the species were not accompanied by illustrations and were at best brief. Several of them are nearly impossible to identify with any particular population. One of the unfortunate pitfalls the modern taxonomist encounters is the tendency to make one of these early descriptions fit his concept of a given species or vice versa. Normally the taxonomist has the type specimen 1961] DODSON & FRYMIRE—THE GENUS STANHOPEA 165 . 20. Stanhopea bucephalus Lindl. Inflorescence (X 14). b. Plant habit ( A Ji). =. Ha view of the lip (X 24). d. Desi view of the lip (X 34). e. Pollinarium (X 2 to fall back on and this specimen, when carefully examined, supplies the necessary information to avoid error. Unfortunately, herbarium specimens of Stanhopeas do not always fulfill this function even when they exist. The critical characters which separate species in this genus are for the most part destroyed in the preparation of specimens. Many times the type specimen does not reflect the actual structure of the popu- lation. An example is to be found in S. wardii in which the type description mentions two teeth produced at the base of the hypochile by the extension of the falcate lateral margins. This character is common in the specimens from southern Central America and perhaps to a lesser extent in Guatemala, but actually does not occur in many of the specimens from Mexico, Nicaragua and northern South America, which are certainly referable to this species on every other criterion. To further complicate the situation, this character does occur occasionally in other species such as S. oculata and S. bucephalus. Doubtless, the type specimen of S. wardii did have this character, but the presence or absence of this one character, (Vor. 48 166 ANNALS OF THE MISSOURI BOTANICAL GARDEN since it is not diagnostic of the total population and is found in other species, should not be considered as a criterion for determination. Several other characters help to make this species distinct from $. oculata and S. bucephalus. Allen (1949) has suggested that Lindley was in error in his description of S. oculata and S. bucepbalus. Allen placed S. bucephalus in synonymy under S. oculata and re-erected S. grave- olens which had been considered as synonymous with S. bucephalus. We cannot, however, agree with Allen and feel that the epithet, S. bucephalus, should be used in place of S. graveolens. Since Lindley's descriptions are extremely brief and the later illustrations which he used of his species correspond rather well with his descriptions and with actual populations, we feel that Allen has allowed his splendid knowledge of the Central American species to overshadow the variation of these species in other areas. Since the type specimens yield little information it will always be difficult to resolve this situation. Another of the confusing situations involved in the taxonomy of Stanhopea is the use of the name “grandiflora”. This name was first used by Kunth when he described and illustrated Epidendrum grandiflorum in 1808. In 1828 Loddiges described another plant and named it Ceratochilus grandiflorus but the name Cera- tochilus had previously been used by Blume for an East Indian orchid. In 1832 Lindley transferred the Loddiges plant to the genus S/anbopea of Frost and described a similar species as Stanhopea eburnea. Later these two plants were found to be conspecific but have been known as S. eburnea to the present time. Lindley's name, S. grandiflora, is valid and must be used in place of S. eburnea. In 1832 Lindley also named a plant which was essentially identical to Kunth's plant as Stanbopea bucephalus. The type specimen was a Ruíz and Pavón collection which, as Rolfe (1912) has pointed out, is marked as being from Mexico. It appears that many of Ruíz and Pavón's specimens were mixed with the Sessé and Мосійо Mexican collections and a great deal of confusion has been caused by this situation. The probable origin of the Ruíz and Pavón specimen was Peru, as it was then called, or what is now Ecuador. It must have come from very near the same locality as Kunth's specimen since the trails of access to the Andes were very few during this period. Reichenbach apparently attempted to clear up the situation by transferring Kunth's plant to Stanhopea and reducing Lindley's S. bucephalus to synonymy under S. grandiflora Rchb. f. Unfortunately, Reichenbach's S. grandiflora is invalid because Lindley's combination precedes it. Another attempt to clarify the situation in the “insignis complex” has been the reduction of the name $. tigrina Batem. to synonymy under $. devoniensis Lindl. by Williams (1951). However, the concept, early descriptions, and illustrations of S. devoniensis and S. tigrina appear to us to be clearly distinct. The plate of S. devoniensis іп Lindley’s Sertum Orchidacearum has little resemblance to Bateman's plate of S. tigrina in his Orchids of Mexico and Guatemala and very likely repre- sents either a rarely encountered species or a hybrid form between S. saccata and S. tigrina. The figure very definitely resembles S. saccata in the features of the lip more closely than it does S. tigrina. Williams may have been misled by the color, and was certainly handicapped by the lack of good type material Occasionally 1961] DODSON & FRYMIRE—THE GENUS STANHOPEA 167 . 21. Stanhopea wardii Lindl. a. Inflorescence (Ж V4). b. Plant habit (X М). с. Side veiw бе the lip (X 14). d. Dorsal view of the lip (X №). [Vor. 48 168 ANNALS OF THE MISSOURI BOTANICAL GARDEN plants with flowers somewhat like the description and plate (discounting the three- lobed epichile which is lacking in the plate but mentioned in the description) turn up, but are quite rare in comparison to the number of typical $. tigrina found in the Mexican populations. CONCLUSIONS This paper has been written as a discussion of Stanhopea preliminary to a formal revision of the genus. Our original intention was to include the revision of Stanbopea with this paper but after examination of herbarium material proved to be disappointing we have concluded that only with further field study of living populations can we accomplish a truly meaningful treatment of the genus. We have also found that to merely revise Stanbopea and leave the other members of the subtribe Stanhopeinae in the condition which they are at the present time would be to do a less than meaningful work. The members of the Stanhopeinae are so closely intermeshed with Stanhopea ie revision of the whole group is mandatory. We hope that in the not too distant future such a revision can be accomplished using classical methods of taxonomy combined with modern biosystematic procedures. In reality then, this paper can only point up the problems involved in the group; it brings up to date the information that is available from the literature and adds our own limited observations of the natural phenomena occurring in the group. These field observations have pointed out the path, so to speak, that should be followed in the study of such a complex and difficult genus as Stanhopea. It may well be true that similar procedures will need to be used in treating other problem groups in the Orchidaceae. Our experience indicates that future studies of the difficult subtribe Catasetinae can only be approached through field study. RS parts of the Sobraliinae, Epidendrinae and Pleurothallidinae will prove to s APPENDIX Key ro STANHOPEA a. Lip simple, composed of a accu hypochile with cbe midlobe forming a no articulated epichile, mesochile absent; column thick, wingless or narrowly winged rich the wings caudate and extended beyond the anther. b. Hypochile entire, not projected into side lobes on its upper edges; column wingless с. те; in center of the lip tongue-like, apex of the lip кр thickened; 1% 3.8 . lon $. ECORNUTA се. Calle in center of the lip absent, apex of the lip thick and fleshy; lip 1.8-2.5 cm. long 2. $. PULLA bb. ыы. projected into fleshy lobes on its upper edges; ime narrowly wines! with the apex of the wings €— and extended beyond the ant S. CIRRHATA aa. Lip ae composed of 2 or 3 distinct pos epichile дената to the — or to a fleshy m mé that is phuc ided with lateral horns; column slender and arcuate with narrow or broad wings which are not Sadan at their apex and extended er cal as anther. b. Муз! erect Find fleshy, paralleling the lip and column; mesochile of the lip а of e large lanceolate horn on each side and a third horn produced the center Нов а пије plate between and extending over the ерїсһїїе......................... . TRICORNIS bb. Petals thin in substance and reflexed; mesochile of the lip with 2 horns on its margins or none. 1961] — . Bo мл . ^ М е DODSON % ЕКҮМІКЕ--ТНЕ GENUS STANHOPEA 169 c. Hypochile of the lip provided with n net horns or protuberences, mesochile relatively simple and without lateral hor: d. сие ска provided with fleshy lobes on each side of = и column rowly winged REICHENBACHIANA dd. Hypochile relatively енг ыр ғы ас base slender and recurved, ae * йе apex not fleshy; column broadly w S. GRANDIFLORA се; okt te ib the lip Meg the what — thickened, not provided ath Vim or tuberences, mesochile with 2 distinct hor A “Hor of the mesochile divided at thei gea to produce a falcate, retrorse Бате р as well as a slender Red. projecting spine; flowers alw P solitary. S. RODIGASIANA dd. Horns of the mesochile acicular or flat, not тое Љиља normally 2 ог more. е. Horns of the mesochile very short, free for less than 1 сепштетег...........-. 8. S. LEWISAE ee. Horns of the mesochile long, free for more tun 2 centimeters f. Mesochile obsolete, very short, hypochile conspicuously adnate to the e: 9. S. CONNATA ff. Mesochile conspicuous, well developed, hypochile not adnate to the colum g. Hypochile globose or saccate 10. insignis dk gg. Hypochile more or less elongate or quadrate 11. oculata complex List or $РЕСЕ$* STANHOPEA ECORNUTA Lem. in Fl. des Serres 2: #. 181. 1846. (Fig. 6). Stanbopea calceolus Hort. ex Rchb. f. Xen. Orch. 1:117. 1858. Stanbopeastrum ecornutum Rchb. f. Xen. Orch. 1:124. 1858. STANHOPEA PULLA Rchb. f. in Gard. Chron. 7:810. 1877. (Fig. 7). STANHOPEA CIRRHATA Lindl. in Journ. Hort. Soc. 5:37. 1850. (Fig. 8). STANHOPEA TRICORNIS Lindl. in Journ. Hort. Soc. 4:236. 1849. (Fig. 9). TStanbopea stenocbila Lehm. & Kranzl. in Gard. Chron. 369. 1900 неон langlasseana Cogn. in Ga rd. Chron. 426. 1901. anhopea convulata Rolfe, in Kew Bull. 366. 1909. а. REICHENBACHIANA Roezl. ex Rchb. f. in Gard. Chron. 2:40. 1879. ( а is hopea amesiana Hort. ex Gard. Chron. 352. 1893. Речи Јошт Rolfe, іп Kew Bull. 63. 1893. Stanhopea suavis Hort. in Ospina, Orquidas Colomb. STANHOPEA GRANDIFLORA (Lodd.) Lindl. Gen. & Sp. Orch. 158. 1832. (Fig. 11). Ceratochilus grandiflorus Lodd. in Bot. Cab. £. 1414. 1828. Stanbopea eburnea Lindl. in Bot. Reg. 18: t. 1520. 1832. à SZ anbopea calceolata Dep Lenq. Hort. Univ. 2: 127 & 2 fStnbope candida Barb. Rodr. Gen. Spec. Orch. Nov. 1: "e 1877. tanbopea randii ima in Kew Bull. 363. 1894. eee RODIGASIANA Claes. ex Cogn. in Gard. Chron. 14. Fig. 9. 1898. (Fig. 12). STANHOPEA LEWISAE Ames & Correll, in Bot. Mus. Leafl. Harv. Univ. 10*:86. Pl. 10. 1942. (Fig. 13). STANHOPEA СОММАТА Kl. in Otto & Deitz. Alleg. Gartenz. 22:226. 1854. (Fig. 14). Stanbopea graveolens Kl. ex Rchb. f. Xen. Orch. 1:118. 1879 enymy in this list have been previously reduced by other authors with * 'The nam the exception ја" устани и thusly: f. 170 10. = — M ANNALS OF THE MISSOURI BOTANICAL GARDEN "insignis complex" STANHOPEA INSIGNIS Frost ex Hook. іп Bot. Mag. #. 1048 8 1040. (Fig. 15). Stanhopea flava Lodd. ex Beer. Prakt. Stud. Fam. Orch. 312. 1854. Stanbopea atropurpurea Lodd. ex Planch. Hort. Donat. Orch. 215. 1858. с. 1859. STANHOPEA TIGRINA Batem. ex Lindl. ‘Sere: Ord. 1. 1838. (Fig. 16). Stanhopea cavendishii Lindl. ex Batem. Lond. Hort. ae сани. У, 643. Stanhopea maculosa Knowles & Westc. Flor. Cab. 3 Stanhopea lyncea Р. М. Don. Hort. Cantab. 13:608. 1845. STANHOPEA DEVONIENSIS Lindl. Sert. Orch. 2. Т. 1838. STANHOPEA MARTIANA Batem. ex Lindl. in Bot. Reg. 26: Misc. 50. (Fig. 17). Stanhopea velata Morren, in Ann. Soc. Gand. 3. Ё. 153. noes SACCATA Batem. Orch. Мех. & Guatem. 7. а ә 1839. hopea implicata Westc. ex Lindl. њи = Ee 6. 1852. Son MS radiosa Lem. Illustr. Hort. 4: Misc Stanbopea marsbii Rchb. f. Xen. Or N 1: 120. fo Stanbopea paste Rolfe, in Kew Bull. 161. STANHOPEA INTERMEDIA КІ. in Act. Hort. р: 17:142. 1898. "oculata complex” STANHOPEA OCULATA (Lodd.) Lindl. Gen. & Sp. Orch. Pl. 158. 1832. 19). Ceratochilus oculatus Lodd. in Bot. Cab. t. 1764. Stanhopea lindleyi Zuccar, Abh. Akad. Muenich 2: ТЕ 1831-33. Stanhopea n Lindl. in Bot. Reg. 29: Misc. 75. 1843. Stanbopea graveolens Morren in Ann. de Gard. 2:55. #. 54. 1846. Stanbopea guttata Kich. Berl. Allg. Gartenz. 364. 1858 Stanhopea ornatissima Lem. in Illustr. Hort. 9:325. 1862. Penes in cymbiformis Rchb. f. Xen. Orch. 2:84. Ё. 124. 1865. anbopea minor Schltr. Notizbl. i Gart. Mus. Berlin "E 485. 1917. [Vor. 48 1829. 1840. (Fig. a BUCEPHALUS Lindl. Gen. & Sp. Orch. Pl. 157. 1832. (Fig. 20). Stanhopea graveolens Lindl. in Bot. Reg. 3: Misc. 59. 1840. Stanhopea aurantia Lodd. ex P. N. Don in Hort. Cantab. 608. 1845. Stanhopea jenishiana а ex Rchb. f. Bot. Zeit. 10:934. 1852. Stanhopea warscewicziana Kl. in Allg. Сое 20:214. 1852. Stanbopea guttata Beer. Prakt. Orch. 3 854. Stan a Rchb. f. Xen. Gai. $i 157. 1858. Stanbopea eurala Hort. ex Planch. Hort. Donat. Orch. 104. 1858. Stanbopea aurata Beer, Prakt. Stud. Fam. Orch. 310. 1858. Stanbopea costaricensis Rchb. f. in Hamb. Gartenz. 16:424. 1860. Stani opea кн =. + Walp. Ann. 6:587. Stanhopea oculata var. constricta Kl. in Acta Hort, Petrop. 17:15. 1898. Stanhopea lietzei Schltr, in Fedde Rep. Sp. Nov. 16:248. 1919. Stanhopea remota Hort. STANHOPEA WARDI Lodd. ex Lindl. Sert. Orch. 7. 20. 1838. (Fig. 21). Stanhopea aurea Lodd. ex Lindl. in Bot. Reg. 4: Misc. 11. 1841. Stanhopea venusta а Lindl. in Bot. Re eg. 4: Misc. 11. 1841. Stan] one inodora Lodd. ex Lindl. in Bot. Reg. #. 65. 1845. tan 3. 852. Stanhopea ритризи Schltr. in Orchis 10: 186. 1916. Stanhopea peruviana Rolfe, in Bot. Mag. Ё. 8417. 1912. 1961] DODSON & FRYMIRE—THE GENUS STANHOPEA 171 SPECIES TOO POORLY KNOWN TO PLACE AT PRESENT STANHOPEA ANNULATA Mansf. in Orchis 16:19. 1938. STANHOPEA BICOLOR C. Koch. Berl. Allg. Gartenz. 209. 1857. STANHOPEA DELTOIDEA Lem. in Illustr. Hort. 9:340. 1862. STANHOPEA FLORIDA Rchb. f. in Gard. Chron. 615. 1879. STANHOPEA FREGEANA Rchb. f. in Otto & Dietr. Allg. Gartenz. 23:313. 1855. STANHOPEA FUERSTENBERGIAE Kranzl. in Gard. Chron. 161. 1899 STANHOPEA GIBBOSA Rchb. f. in Gard. Chron. 1254. 1869. STANHOPEA HASELOVIANA Rchb. f. in Otto & Dietr. Alg. Gartenz. 23:322. 1855 STANHOPEA НОРРИ Schltr. in Fedde Rep. Sp. Nov. 27:82. 1924. STANHOPEA IMPRESSA Rolfe, in Kew Bull. 196. 1898. STANHOPEA MADOUXIANA Cogn. in Gard. Chron. 134. 1898. STANHOPEA MOLIANA Rolfe, in Lindenia 7:89. /. 331. 1891. STANHOPEA NIGRIPES Rolfe, in Kew Bull. 364. 1894. STANHOPEA PLATYCERAS Rchb. f. in Gard. Chron. 27. 1868. STANHOPEA RUCKERI Lindl. in Bot. Reg. subt. 44. 1843. STANHOPEA SCHILLERIANA Rchb. f. Xen. Orch. 2:158. 1858. STANHOPEA SHUTTLEWORTHII Rchb. f. in Gard. Chron. 795. 1876. STANHOPEA UNCINATA Drap. Hort. Univ. 4:65. 1843. STANHOPEA XYTRIOPHORA Rchb. f. in Gard. Chron. 842. 1868. DUBIOUS SPECIES Stanhopea quadricornis Lindl. in Bot. Reg. 24: 7. 5. 1838. OBSCURE SPECIES Stanbopea russelliana Lodd. ex P. N. Don in Donn. Hort. Cantab. 13:721. 1845. Stanbopea violacea Hort. ex Beer. Beitr. Morph. и Biol. Orch. 7. 4 8 8. 1863. Stanbopea wallisii Rchb. f. in Linnaea 41:109. 1877. Stanbopea bernandezii (Kunth) Schlechter, Beih. Bot. Centr. 2:490. 1918. LITERATURE CITED Allen, P. А, 1949. Flora of Panama (Orchidaecae). Ann. = Bot. Gard. 36:53-59 ------. 1951. Pollination in Coryanthes Let osa. Amer. Orch. Soc. Bull. 19:528. = E The Swan Orchids, A Revision of the Genus Cycnoches. Orch. Journ. 14-8:226. ------. 1956. Pollination in Gongora spe pon Amer. Orch. Soc. Bull. 24:230. Ames, O. & D. S. Correll. 1952. Flora of rig TE (Orchidaceae). Fieldiana: Bot. 262:528—536. ideae. Jou i . Bot. 18:281—360. Dressler, R. L. & C. H. DU 1960. Classification and Phylogeny i in "ба Orchidacese. Ann. Mo. Bot. Gard. 47:25-6 Rolfe, R. A. 1912. жне» үч and S. bucephalus. Orch. Rev. 20:172. Schweinfurth, C. 1958. Orchids of Peru. Fieldiana: Bot. m :606-612. Williams, L. O. 1951. The ры of Mexico. Ceiba 2:242-249. [Vor. 48 172 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE Ш Pollination in Stanbopea B. Bee falling Vine the flower and engaging the spine of the rostellum under his mete thorax. C. Pollinia being inserted into the cack cleft while vu bee is falling through the flower. D. Eulaema moriana approaching the flower of S. tricornis. ANN. Mo. Bor. Garp., Vor. 48, 1961 Pirate Ш DODSON & FRYMIRE—THE GENUS STANHOPEA MISSOURI BOTANICAL GARDEN STAFF Director Fairs W. WENT Ербак AN Joun D. Dwyer, Curator of Useful Plants Research Associate HENRY 45 Е "^ Мовтом H. „өн, Paleobotanis: Morphologi: cons par ке TRIFON VON ES Taxonomist Assistant Curator o ini of Economic Plants Носн C. CUTLER, xecutive Director ey x ]. Jute PE. Tu Дорсе, кек лове, Мусо из ке nm SCHAACE, | | ul. X Da of Grasses and Librarian beg egre mist st and 3 Editor of ed ат Е, ,oopsom, ж. кен Herbarium James A. Dike, елене Са йок £ Herbarium BOARD OF TRUSTEES. President 1 Rosrar Brookes SMITH Vice-President CESTER В. Faust DM Second Vice-President M UE “Henry B. РЕТАСЕК _ Howarp Е. BAER | 4 Hz Danret К. Сатым | | Jonn S. LEHMANN ЗАМ. C. Davis 0/0 _ КовЕвт У. Отто — E. Duprey FRENCH — S otl WARREN Ma SHAPLEIGH EX-OFFICIO MEMBERS оков. Um ка И x Eu окр Lez Monto Bishop of the Dioc SR nod ш E "President ofthe Academy of зан -- dul ‚ Seience оғ St. Louis NNELLY RD | Етан A H. mem o Kao wp R. Tin d e _ Mayr ofthe Cay oF $ St. Louis PE e Қамыт Linco, p Secretary Volume XLVIII Number 3 Annals of the Missouri Botanical Garden SEPTEMBER, 1961 Preliminary Revision of the Genus Drymaria . James A. Duke 173-268 Sanango: New Amazonian Genus of Loganiaceae с. ч . . George S. Bunting and Julie А. Duke 269-274 PUBLISHED QUARTERLY АТ GALESBURG, ILLINOIS BY THE BOARD OF gae nso OF THE ee BOTANICAL GARDEN Т. LOUIS, MISSOU Entered as second-class matter at the ee gaa = a Illinois, under the aes of March 3. Annals of the Missouri Botanical Garden А quarterly journal containing scientific contributions from the i Botanical Garden and the Henry Shaw School of Botany of елді University in affiliation with the Missouri Botanical Garden. Information The ANNALS OF THE MISSOURI BOTANICAL GARDEN appears four times during the calendar year: February, May, September, and November. Four numbers constitute а volume. Beginning with Volume 45, 1958: Subscription Price $12.00 per volume Single Numbers 3.00 each Contents of Tier issues of the ANNALS OF THE Missourr BOTANICAL GARDEN are listed in the Agricultural Index, published by the H. W. Wilson Company. Annals of the Missouri Botanical Garden Vol. XLVIII SEPTEMBER, 1961 No. 3 PRELIMINARY REVISION OF THE GENUS DRYMARIA JAMES A. DUKE ABSTRACT or the first time since the days of DeCandolle, a revision te all the known species of Drymaria is Ness The species are arranged in seventeen series, and keys are provided to the forty-eight species recognized. Illustrations, mostly based on type mater ial, and morphological descriptions are provided for each species. Two new species, D. barkleyi and D. conzattii, are described, as well as several new in DENM. taxa. JAMES А, DUKE, Mision Sod Garden, 2315 Tower Grove Ave., St. Louis 10, Missour INTRODUCTION The genus Drymaria comprised four species at the time of its establishment by Willdenow (in Roem. & Schult. Syst. Veg. 5:406. 1819). Less than 150 years later the number of names has multiplied more than thirty-fold, with most of the novelties being described before the turn of the present century. No complete revision has been attempted since this prodigious multiplication of epithets. Wiggins (in Proc. Cal. Acad. 475:189. 1944) has completed an able revision of the genus as represented in the vicinity of the Sonoran Desert. His conclusions were quite sound, and the only consequential differences expressed in the present paper are mere matters of heirarchic reduction. In addition to Wiggins, the most important recent American studies on Dry- maria have been pursued by Fosberg, who has clarified many of the problems created by M. E. Jones, proposed a few new taxa, and made several appropriate reductions, both in the literature and in various herbaria. Macbride's account in the Flora of Peru, although promulgating a few errors, has been a forward step toward under- standing the genus in an unusually complicated region, where speciation in Dry- maria has run rampant. I. M. Johnston's studies in the equally perplexing Coahuilan Desert have resulted in the description of some of the most bizarre species of Drymaria. The major contributor from Europe has been the late Dr. Mattfeld whose discerning work has been responsible for the recognition of several new South American species, all of them herein maintained. He has also clarified some of the errors stemming from the work of his predecessor Briquet, who described several new Mexican species, two of which were probably based upon Pavon's Peruvian collections. (173) [Vor. 48 174 ANNALS OF THE MISSOURI BOTANICAL GARDEN Gross MORPHOLOGY An amazing array of characters is exhibited in the genus, and will doubtless tempt the description of minor variants. The petals, for example, are nearly as variable as those of Silene. Almost every portion of the plant has been utilized as a taxonomic tool at one place or another in the genus. Various habits are assumed by members of the genus. Perhaps most common is the clambering and sprawling habit exhibited by D. cordata, which occasionally roots at the nodes. Erect suffrutescent perennials are not uncommon, and may raise an eyebrow on those not intimately familiar with the genus. Depressed sub- cespitose forms such as occur in some varieties of D. effusa recall some species of Arenaria, while delicate subvirigate annuals such as D. leptophylla may suggest Stipulicida. Too often, the duration has been used as a key character, but the duration of a plant is frequently a function of the environment. Plants grown in more xeric habitats often exhibit more attributes of a perennial. Although the leaves of most species are of an ovate type and often cordate at the base, every extreme, from linear and subsucculent to reniform and membrana- ceous, is found in one species or another. The leaves are usually entire, but undula- tions occur in the leaf margins of some South American species. The opposite arrangement prevails, but in a few series pseudoverticillate leaves are common. Venation of the leaves is usually obscure and rarely of taxonomic significance, but the texture and pigmentation occasionally aid in identification. The petioles may be as long as the blades or completely obsolete. Most species have stipules which may be entire, bifid, or irregularly lacerate, and the nature of the division is of occasional importance as a taxonomic criterion. Indumental variations are often called upon for specific delimitations. Septate or moniliform hairs occur in several species. Forms of D. villosa may be hirsute or villose in nature; forms of D. ovata are best described as lanose; stipitate or sessile glands are present in many species. In D. cordata the glands commonly take the form of a conspicuous farinose girdle on the pedicels. The indument is usually more or less translucent, but is not uncommonly white, or even reddish or yellowish. Completely glabrous species occur, and some of the xeromorphic and halophytic species are decidedly glaucous. Predominating in the genus is a terminal cymose inflorescence, but here again, numerous departures from the norm exist. In most species with the leaves pseudo- verticillate, the flowers reflect this arrangement. Rarely the flowers may be solitary or clustered in the axils of barely modified leaves, but more commonly each flower is subtended by more or less scariose bracts. Many inflorescences show racemose tendencies; in D. anomala, the first and sometimes the second branches of the inflorescence are dichasial, but subsequent branching tends to be monopodial. Pro- portionate lengths of bracts, pedicels, peduncles and flowers, as well as relative variations in their induments, have been used in distinguishing species. Almost invariably there are five sepals in a flower, but aberrant flowers are known with four or six sepals. The sepals are most commonly lanceolate or ovate but in some they are nearly orbicular and in others nearly deltoid. One to nine nerves may traverse each sepal, the 3-nerved sepal being most prevalent. In some 1961) DUKE—REVISION OF DRYMARIA 175 few species the venation is more or less dendritic and in others so obscure as to be difficult of interpretation. Occasionally the sepals are conspicuously carinate or cucullate. Shape, apex, nervation, indument and comparison of the sepals are frequent taxonomic tools. Petals are usually five in number, but three is characteristic of certain species, and the petals are absent or drastically reduced in D. apetala and in forms of D. villosa and D. xerophylla. Typically the petal is bifid for about half its length, but this basic construction is hard to visualize in some of the members of the first few series whose ornate petals are so bizarre as to defy terse terminology to describe them. The lobes in some more easily described forms are merely basally auriculate, the nature of the auricles being quite variable, as are the venation and apices of the lobes. Most of the bilobed petals are tapered directly to a narrow claw, but in some members (e.g. in Series FASCICULATAE) there is a longer or shorter trunk interpolated between the lobes and the claw. In some forms (e.g. Series HOLO- STEOIDES) the trunk never narrows to a claw; such forms are herein referred to as exunguiculate. The lobes of the bilobed petals are 1-тапу-пегуе4, the 1-nerved condition being characteristic of the Series СОВРАТАЕ, and certain species in other series. Less constant in number than the petals are the stamens, which are occasionally reduced to two in number. In species with the reduced stamen number, the anthers are often abortive, and tend to be more orbicular than in those species possessing the higher number of stamens; in pentandrous flowers the anthers are usually oblong. Not infrequently the filaments are shallowly connate into a cup, and there are occasionally minute staminodial flaps alternating with the filaments on such a сир. Rarely, as in D. stipitata, there are conspicuous saccate staminodia. In the Series FRUTESCENTES occasionally floral dimorphism suggests that monoecism is in the early stages of evolution. Offering little in the way of systematic clues is the gynoecium. In a few species the ovary is borne at the summit of a rather prominent stipe, the presence of which does not seem to be necessarily constant. Some species, which are also characterized by the development of staminodia, have a rather prominent stylopodium atop the ovary. In most species, there are three styles, but occasionally there may be two or four. Such numerical variations have been observed in a single inflorescence. The degree of fusion of the styles on the other hand seems to be a more reliable taxonomic criterion, the styles being fused to a greater or lesser degree in most species. Only with hesitancy do I use the word fused; it would seem to be just as well to speak of one style with three branches as to speak of three styles partially fused. For the sake of tersity, I have often described the style as being bifid or trifid half its length, when it might have been more correct morphologically to say that there were two or three styles fused for half their length. Interestingly the two or three styles are almost completely free in D. cordata, a species which has the most reduced androecium, and a species which I would judge to be one of the most advanced in the genus. The capsule may be three-valved or two- or four-valved, correlated with the number of styles. Rarely the size of the capsule may be diagnostic. Inside the (Мог. 48 176 ANNALS OF THE MISSOURI BOTANICAL GARDEN capsule, 1-many of the campylotropous ovules may mature into seeds, and the seeds have proven very valuable in the taxonomy of the group. The number, shape, color and sculpture of the seeds all play some part in species determination. Most species are characterized by many-seeded capsules, but in the Series CORDATAE, some occasionally mature only one seed per capsule. The seeds may assume any of several shapes. Perhaps most common is the cochleate, or snail-shaped seed, characteristic of Series CORDATAE. Foetiform seeds, shaped much like the human foetus, are found in the Series HoLosTEoIDES, and the vestiges of this shape are retained in some subsequent groups. Ampulliform, or retort-shaped, seeds occur in D. debilis and this shape is approached by a few species of other sections. The seeds of D. viscosa are unusual not only in being lacrimiform, but also in being smooth, dull and yellowish. Many members of the earlier series have hippocrepiform seeds and species of the Series LYROPETALA have hairs on the hump of the horseshoe. Seeds of most species are characterized by various kinds of sculpture. In the earlier series the seeds are merely granular, if we exclude the hairs of Series LYROPETALA from the realm of sculpture. Later sections, especially in the Series vILLOSAE are endowed with species whose seeds are ornate indeed. For example the tubercles in D. villosa are stellate in outline, those in its subsp. palustris are often capitate, those in D. divaricata var. stricta are secondarily tuberculate. On the other hand, forms of D. ovata have smooth and lustrous black seeds. Usually the seeds are reddish-brown, but in D. viscosa they are yellow; occasionally seeds of some species in the Series ARENARIOIDES emit a purple substance into the dissection medium. Seeds have turned out to be rather critical indicators of a species’ systematic position іп Drymaria. It should be pointed out however, that in working with Spergula, J. K. New (in Ann. Bot. 23:23. 1960) has found variations in the number of papillae on the seed coats to be environmentally induced. Furthermore she has found no genetic correlation between the variations in seed-coats and in hairiness. Species of Drymaria, whose seeds are highly viable a year after collection and germinate rapidly in the greenhouse, should prove interesting subjects for similar studies. GEOGRAPHY Departing somewhat from the “typical” picture of the Caryophyllaceae, Dry- maria is largely a subtropical genus. Of the forty-eight species herein maintained, all but two are strictly American, and only one subspecies does not occur in America. Аза whole in America the genus ranges from the western United States through Central and South America to Argentina and Chile, the large majority of the species occuring on the western halves of the continents. Florida, Surinam and Brazil, on the eastern shore, seem to possess only the pantropical D. cordata. For a geographical tabulation of species, the reader is referred to the citation of specimens which terminates this paper. Of particularly geographical interest is D. cordata, whose varietal potential and geographical distribution has been studied in some detail by Mizushima (in Jour. Jap. Bot. 32:69. 1957). Almost all material from continental Asia belongs to the subspecies diandra (maintained as a species by Mizushima), a subspecies completely 1961] DUKE—REVISION OF DRYMARIA 177 lacking in America (excl. Hawaii). On the Pacific Islands both subspecies and the rather distinct variety pacifica occur and appear to maintain their integrity by and large. In Africa however, there are about as many intermediates as there are pure representatives of the two subspecies. A second species occurs in Malaysia and, to judge from the literature, has been introduced there only within the present century. Of course there is the more remote possibility that it has merely been overlooked until this century. The species, D. villosa, is so distinct that I prefer to believe that it is a recent introduc- tion from the Americas. Several interesting disjunctions are apparent in the genus. Most striking is that illustrated by D. ladewii, known only from two collections in Bolivia, but repre- sented by bountiful material from Guatemala and southern Mexico. More easy of interpretation are several vicarious disjunctions between Baja California and main- land Mexico, e.g. D. holosteoides & D. pachyphylla; D. arenarioides ssp. peninsularis & D. arenarioides ssp. arenarioides; D. gracilis ssp. carinata & D. gracilis ssp. gracilis. Economics Often a conspicuous element in the herbaceous vegetation, Drymaria probably plays in the subtropics the insignificant role played by the chickweed in temperate North America, fairly well known, but hardly a topic for conversation. Drymaria cordata, and perhaps others, which spread radially and root at the nodes forming rather lush mats, have been planted as ground covers. In Ecuador various species are used as curatives for liver and kidney ailments. In Mexico, species are reputedly used to cure “yaza” and in the Orient, infusions are purportedly used to alleviate headaches. One should be careful of the plant, especially as an oral administration. Drymaria pachyphylla (and probably related species) is seriously toxic to livestock, and all measures should be taken to prevent its spread in the arid West, where palatable fodder is at a minimum. Fortunately the seeds alone of D. pachyphylla are so distinctive (see Fig. 1C) as to separate it from any other species. ACKNOWLEDGMENTS In the course of this study I have been fortunate enough to study all or most of the material of the Gray Herbarium, the University of California, the New York Botanical Garden, the Field Museum, the Pomona College Herbarium, the Missouri Botanical Garden, the Rijksherbarium, the Leningrad Herbarium, the Herbarium of the University of Glasgow, the Botanisches Museum of Berlin, the Botanische Staatssammlung at Munich, the Museum National d’Histoire Naturelle at Paris, and the University of Michigan. Critical material has also been loaned by the curators of the herbaria at Kew, Prague, Stockholm, Lija, Lund, Geneva, Göttingen, the University of Brazil, and the California Academy of Sciences. Dr. Ramon Ferreyra has forwarded many interesting Peruvian specimens for determination. To him and to the curators and my correspondents at all the aforementioned institutions, I express my sincere gratitude for g their time and efforts avail- able to me. To my colleagues here at the Missouri Botanical Garden, I am indebted (Мог. 48 178 ANNALS OF THE MISSOURI BOTANICAL GARDEN for their suggestions and comments.* Special thanks are extended to my wife, Peggy, who has been kind enough to prepare the illustrations. DRYMARIA Willd. Drymaria Willd. ex Roem. & Schult. т 5:31. 1819. Pinosia Urban, in Arkiv. Bot. 23А5:70. pl. 2. Mollugophytum M. E. Jones, in Extr. Contr. West. Bot. 18:35. 1933. Annual or perennial, glabrous or pubescent herbs, occasionally subligneous below, prostrate, spreading or erect. Leaves opposite or pseudoverticillate, glabrous to villose or glandular, sessile to long-petiolate, usually with persistent or fugaceous small stipules. Flowers few to many in dichasial cymes, rarely in pseudoverticils, racemes, or solitary or clustered in the leaf axils. Sepals 5, not connate. Petals 0-) 3-5, white, usually 2-cleft, occasionally appendiculate in the sinus, often auriculate. Stamens 2—5, the anthers versatile, 2-celled, the flattened filaments slightly connate at the base, rarely alternating with prominent staminodia. Ovary superior, slightly stipitate; carpels mostly 3, the 3 styles more or less united below; ovules few to many, campylotropous on free central placentae. Capsule ovoid to spheroid, usually dehiscing into 3 entire valves; seeds 1 to many, cochleate, foeti- form, hippocrepiform, or ampulliform, usually tuberculate, the embryo curved about the perisperm. As here conceived the genus consists of 48 species, all but two of them exclu- sively American. Since the genus has never been monographed in its entirety, and no supraspecific classification has been elaborated, I would like to propose the following informal series, most of which seem to be rather homogeneous. Follow- ing a tabular presentation of the series is an artificial analytic key to all species. Following this is a systematic key to the series. The latter key, although not assuredly phylogenetic, is so set up that species with the more elaborate floral characters, often correlated with certain vegetative factors, come first, and the species with simpler floral construction come later. I am not prepared to say whether evolution in the genus has been a matter of reduction or amplification or SuPRASPECIFIC CLASSIFICATION A. Series HOLOSTEOIDES D. Series ARENARIOIDES . D. bolosteoides 8. D. mollugin 2. D. pacbypbylla 9. D. arenarioides қ 10. D. axillaris B. iw о 1 her за kleyi 12. D. pol: ide. е р. noraidot ы т . D. suffru E. Series УІЅСОЅАЕ ^ D. redes gm 13. D. viscosa C. Series STIPITATAE F. Series oRTEGIOIDES 7. D. stipitata 14. D. ortegioides *Dr. F. Raymond Fosberg has — reviewed an early draft of this paper and made many — suggestions; unfor EN r. Fosberg's comments were received as this went to press, and be used to full adva 1961] DUKE—REVISION OF DRYMARIA 179 G. Series EXCISAE М. Series vILLOSAE 15; D. hypericifolia 31. D. multiflora 16. D. excisa 32. D. conzattii 17. D. longepedunculata 33. D. malachioides 34. D. villosa H. Series LEPTOPHYLLA 18. D. effusa O. Series GRANDIFLORES 19. D. leptophylla 35. D. firmula I. Series TENUE 36. D. ovate : 37. D. apetala 20. D. anomala 38. D. glaberrima 21. D. tenuis 39. D. montico . 40. D. grandiflora J. Series FRUTESCENTES 41. D. paposa 22. D. stellarioides 42. D. rotundiflora 25. D. stereophylla 24. D. auriculipetala P. Series DIVARICATAE 25. D. frutescens 43. D. divaricata K. Series FASCICULATAE је D Тағыны Q. Series CORDATAE 27. 5 ah toe 44. D. gracilis 28. D. prae 45. D. glandulosa 46. D. xerophylla Г. Series DEBILES 47. D. ladewii 29. D. debilis 48. D. cordata M. Series LAXIFLORES 1. 1. 30. D. laxiflora ARTIFICIAL КЕХ To SPECIES AND SUBSPECIFIC TAXA Petals absent or reduced, shorter than the stamens: 2. Se 3. Plants villose. 34а. D. VILLOSA f. TEPICANA 23. Plants not villose: 4. Sepals 3—4 mm. long; Mexico 46. D. XEROPHYLLA 4. Sepals 7.5—9.0 mm. long; Peru 37 D. APETALA Is obtuse 34b. D. vittosa f. TOWNSENDII Petals present; longer than or as long as the stamens 5. Petals laterally lacerate, alternating with clavate staminodia 7. D. STIPITATA 5. Petals not laterally lacerate; stamodia not clavate 6. Leaves pseudoverticillate; petals often distally lacerate; North America: 7. Seeds dorsally hispidulous; petals bifid or polylacerate apically. 8. Inflorescence compact, the bracts subcontinguous -5 mm. long 3. D. ELATA 9. Sepals 5-6 mm. long 4. D. SUBUMBELLATA 8. Inflorescence lax, the bracts remote. 10. Trunk of the petals nearly as broad as long; plants glabrous......... 5. D. SUFFRUTICOSA glandular 10. Trunk of the petals about twice as long as broad; plants QUE os - 6. D. LYROPETALA 7. Seeds not dorsally hispidulous; petals bifid or polylacerate apically: 11. Petals apically 3—many-fid. [Vor. 48 180 ANNALS OF THE MISSOURI BOTANICAL GARDEN 12. Leaves linear to oblong; seeds not foetiform. 13. Flowers in cymes, the bracts clearly different from the foliage leaves........... 8. D. MOLLUGINEA 13. Flowers more or less racemose, the bracts little different from the foliage leaves. 14. Plants not glandular; жеті trifid and bifid 11. О. BARKLEYI 14. Plants glandular; petals 4- 15. Sepals and petals 4-7 mm. long........... . D. ARENARIOIDES ssp. ARENARIOIDES 15. Sepals and petals 2.8-4.0 mm. long; ы Californi D. ARENARIOIDES ssp. PENINSULARIS 12. Leaves more or less elliptic, glaucous; seeds о 16. es with 5—10 lobes; sepals 3-5 mm. long; Baja California. es mostly more than 4 cm. apart, међа branchin a. D. HOLOSTEOIDES var. HOLOSTEOIDES 17. Nodes mostly less than 4 cm. apart, dy Sek. D. HOLOSTEOIDES Var. CRASSIFOLIA 16. Petals mostly with 4 lobes; sepals 2.0-3.5 mm. long; си Nort ica D. PACHYPHYLLA 11. Petals merely bifid. 18. Sepals obtuse, not strongly 3-ribbed; flowers racemose or cymose. 19. Sepals stipitate-glandular; Baja Californi = Seeds yellow, smooth 13. D. viscosa 0. Seeds brown, tuberculate or granular....... 9b. D. ARENARIOIDES ssp. PENINSULARIS 19. pes withont stipitate glands; continental North America: 21. Leaves linear $. D. MOLLUGINEA 11. D. BARKLEYI 21. Leaves elliptic 18. Sepals acute or strongly 3-ribbed, if obtuse, the flowers in cymes. 22. Sepals obtuse (D. effusa): 23. Bracts mostly shorter than the pedicels: 24. Axes of the inflorescence stipitate-glandular 18a. D. EFFUSA var. EFFUSA 24. Axes of the inflorescence not stipitate-glandular 18b. D. EFFUSA var. CONFUSA 23. Bracts mostly longer than the pedicels; plants minute. 18c EFFUSA Var. DEPRESSA 22. Some or all the sepals in a flower acute to acuminate (D. vod cidit 25. Peduncles not stipitate-glandular; — унесе id the bracts........... YLLA Var. pasen МА 25. oe stipitate-glandular; данак: y tassi анал than racts, 26. Sepals lanceolate, not stipitate-glandular...19b. D. LEPTOPHYLLA var. COGNATA 26. Sepals ovate, stipitate-glandular................... c. D. LEPTOPHYLLA var. NODOSA 6. Leaves opposite; petals bifid, the lobes themselves sometimes dichotomizing; North and South America, 2 spp. more or less pantropical. 27. Petals 2-many-times dichotomous; Mexico; West Indies: 28. Leaves Movil es 14. D. ORTEGIOIDES 28. Leaves 29. Mont dein een 1-2 times жылы С, 17. D. LONGEPEDUNCULATA 29. Plants not villose; 2—4 times dichotomous: 3 ves deltoid-ovate, rather nar 15. D. HYPERICIFOLIA XCISA 30. Leav r 27. T merely bifid, the lobes occasionally emargina - Leaves linear to elliptic: (cf. also D. pleads, D. axillaris.) = Plants rarely suffruticose; North Am 33. Leaves linear to spatulate; РАЙ 34. Sepals all obtuse, often cucullate (D. s 4): 35. D more than 5 cm. tall; many of the pedicels longer than the ts. —— 2 €—— 1961] DUKE—REVISION OF DRYMARIA 181 36. Peduncles and pedicels stipitate-glandul 18a. D. EFFUSA var. EFFUSA 36. Peduncles and pedicels not stipitate-glandular 18b. D. EFFUSA var. CONFUSA 35. a mostly less than 5 cm. tall; bracts — — than cels D. A var. ражи 34. Кан or all of the sepals acute ог acuminate (D. TING 37. — not stipitate-glandular; — — shorter than the D. LEPTOPHYLLA var. LEPTOPHYLLA 37. M Rm stipitate-glandular; — ade D - Sepals lanceolate, not stipitate-glan 19b. D. LEPTOPHYLLA var. COGNATA 38. Sepals ovate, stipitate-glandul 19c. D. LEPTOPHYLLA var. NODOSA 33. кейі lanceolate, the petioles үзген? th Sepals 3-ribbed 20. D. ANOMALA pals 1-ribbed D. TENUIS 32. ak suffrutescent; Peru and Ecuador: 40. Leaves neither imbricate nor pung 41. Sepals usually glandular, ha umi not excurrent. 42. Leaves subpe umm eous, stipitate; Ecuador 22. D. STELLARIOIDES 42. Lea em anaceous, often estipulate; Peru 23. D. STEREOPHYLLA 41. Sepals bau de mi excurrent 24. D. AURICULIPETALA 40. Leaves imbricate, pungent 25. D. FRUTESCENS 31. Leaves narrowly ovate to renifor 43. PM и, the bro ad trunk not narrowed to a claw, the trunk abou wide as the combined widths of the lobes. 44. ih -lobes emarginate 30. D. LAXIFLORA 44. Petal-lobes acute to emarginate. 45. Cyanic glaucous perennials of the Coahuilan Desert. 46. Flowers axillary, the sepals stipitate-glandular 10. D. AXILLARIS 46. Flowers cymose, the sepals glabrous 12. D. POLYCARPOIDES ee Vern dina or perennials of South America б асыуы more than 4.5 mm. long, ог the bracts not stipulate.. . Inflorescences mostly ер lax, the pedicels obvious...38. D. GLABERRIMA 48. Inflorescences terminal, dense, the pedicels obscured....... 26. D. FASCICULATA 47. Sepals less than 4.5 mm. long, the bracts often stipulate. 49. Sepals acute, n E DE 50. Pedicels mostly longer than the bracts 28. D. PRAECOX 50. Pedicels mostly shorter than the brach a. D. ENGLERIANA var. ENGLERIANA 49. 15 acuminate, glabrous 27b. D. ENGLERIANA var. DEVIA 43. Petals vid a definite claw, the two lobes basally tapered or truncate to the claw 51. Pet is with ciliate есы near the summit of the claw; seeds with sub- re or capitate or cylindric tubercles . Leaves broadest at or above the middle, " stipules ne Ро рат E vx widest quite near the D. MULTIFLORA 52. Lea t below the middle, зе sti — if entire, ue or —— e К to orbicular; w е bove the base. 53. s 4.5 mm. long or longer; 2 cordate, often broader than 1.5 cm. 54. Stipules longer than the petioles 32. D. CONZATTI 54. Stipules shorter than the petioles. 33. D. MALACHIOIDES 53. Sepals mostly less than 4.5 mm. long; leaves if rarely cordate, mostly less than 1.5 cm. broad: 55. Plants villose; mostly clambering annuals. 56. Sepals elliptic, acute; tubercles of the seeds stellate. 34a. D. VILLOSA ssp. VILLOSA (Vor. 48 ANNALS OF THE MISSOURI BOTANICAL GARDEN 56. кере oblong to ici obtuse; tubercles of the seeds stellate, capitate, cylindric p ions sae ornate ааа petal lobes not чат 2 epals usually glabro 34b. р OSA ssp. PALUSTRIS 7: маан ith simple т. petal lobes deeply ini cma sepals dol i нана Н, 55. Plants glabrous, к ves жамыла ct p lea ч, ville iei о plants sprawling annuals or erec nnial 34b. D. viLLOSA ssp. PALUSTRIS 51. Petals without ciliate auricles; seeds with simple or ornate tubercles, rarely uberculate: 58. Plants villose: 59. Seeds deco stamens of two sizes M California 55; 5 ochleate; stamens more or less equa 60 29. D. DEBILIS "ah with ornate tubercles; sepals no more than 4.5 mm. бі. Sepals glabrous, obtuse. 34b. 61. Sepals villose or villosulous, obtuse or acute . VILLOSA ssp. PALUSTRIS 62. Sepals obtuse; Venezuela and Colombia 62. Sepals acute; Pe c. D. VILLOSA ssp. PARAMORUM 28 60. Seeds smooth o D. PRAECOX or the tubercles not ornate; sepals 4—10 mm. long: 63. ena bifid about 34 their length, the lobes broad, Яз a ed; . GRANDIFLORA 63. ex hee out. на Тин length, the lobes narrow, Sona. Andea uth A 58. Plants not a D. ОУАТА 64. а ыч or shorter chin the sepals, the lobes mostly 1-nerved; often fewer than five eM pd about ibas the locals in length: 66. Sepals 3-nerved, not carinate; tubercles — 67. Sepals o to orbicular, obtuse, petals ics with ciliate aei tubercles of the seeds Ми ог sapis 34b. %-.--.--..... OSA em PALUSTRIS 67. Sepals се acute, the petals scarcely el a ез of the seeds domical or conical, ner ily tuberculat D: багета var. STRICTA 66. Sepals 1-nerved, carinate; tubercles not ornate (D. gracilis). 68. Stipules lacerate; continental Mexico and кам America a. D. GRACILIS ssp. GRACILIS Stipules entire; Baja California............. се Ра GRACILIS ssp. CARINATA 65. a — = the se 69. Flowers с ered in axi 7 subsessile leaves, the inflorescence, if rar arely. эчене ih supernumerary bracts.....46. D. XEROPHYLLA 69. Flowers in lax мл ая axillary cymes, if rarely solitary, the su eaves obviously petiolat 70. Leaves more or са невин ‚ trinerved, with the secondary v forming an y raised, reticulum, very А, sti Вә а. ду... 47. D. LADEWII reniform or cordate, not obviously trinerved or reticulate; бебче lacerate: 71 6–0.8 mm. long, се ға rectanguloid, remote; plants а erect and per 45a. D. GLANDULOSA Var. GLANDULOSA 71. .8-2.0 mm. broad, the аны domical, contiguous; plants дода. and usually pros 72. Flowers broadest at or етік Gg Jr eh sepals n inflexed; seeds 0 .8-1.5 mm. broad... ATA ssp. jonas 25 eh broadest above the "t. hé eme usually rved; seeds 1.5—2.0 mm. broa 48b. D. cORDATA ssp. DIANDRA LI 1961] DUKE—REVISION OF DRYMARIA 183 64. Petals — or —— the — the lobes with dendritic or late venation; stam ostly five 73. Sepals obtuse, often nail за America: 74. Flowers in lax ax illary ер хаза glabrous: 75. Petals subexunguiculate 38. D. GLABERRIMA 75. Petals tapered to a long ЧЕ claw; Galapagos er ERES 74. Flowers in dense to lax, mostly terminal, cymes: Sepals glandular or villosulous: 7. Sepals carinate, the inflorescences capitat 22 ng clearly exerted, the lobes truncate to the claw 41Ь. D. РА NA Var. WEBERBAUERI 78. "i — HA at all exerted, the эш орай to the claw; northern ANA Var. PAPOSANA 77. Sepals not or ranger carinate, the ли Е. 40. D. GRANDIFLORA 76. Sepals glabrous or with scanty sessile glands: 79. Sepals ellipsoid to orbicular, quite obtuse, often with sessile mandi a Leti 42a. D. ROTUNDIFOLIA Var, ROTUNDIFOLIA 79. Sepals narrowly to broadly ovate, тшк АБО. ные 42b. D. A cerise ei ex var. NITIDA = gs acute; North and South America: . Seeds lustrous, often smooth; leaves often pergameneous: 81. Stipules as long as the petioles; seeds tuberculate; ber or: ра D. FIRMULA 81. Petioles equaling to much longer than the stipules; = corru- gated or smooth; Colombia to Argentina along the Andes......... 36 80. Seeds dull, tuberculate; leaves membranaceous: 82. Seeds with low remote и Mexico. Ь. 10. GLANDULOSA var. GALEOTTIANA 82. — я prominent, as contiguous, tubercles; South 83. de carinate, the carinae often serrulate 84. Inflorescence capitate, the "m sd e to the ANA Var. Jona 84. Inflorescence lax, the petal be — to ен da. ANA Var. SERRULATA 3. Sepals rarely carinate, the carina not ee 85. Inflorescence capitate, the Fur lobes truncate to the claw. b. D. PAPOSANA var. WEBERBAUERI 85. Inflorescence not capitate, the petal lobes tapered to the claw but often auriculate: 86. Petals and sepals about equal in length: 87. € auriculate: lic cem weakly nerved, basally saccate: ы long- eyonu: tubercles of the seeds ammiliform. ...43а. D. DIVARICATA var. DIVARICATA 9. тте pe алалы ; tubercles he seeds secondarily tuberculate. 43b. D. DIVARICATA Var. STRICTA 8. Sepals strongly nerved, not basally saccate: 90. Leaves elliptic to еа short-petiolate................. e. D. DIVARICATA Var. REFLEXIFLORA 90. Leaves, at least some on a given plant, reniform, long-petiolate: 91. мо. - herbage densely viscid-glandular; t anthesis ca. twice as long as broad, [Vor. 48 184 ANNALS OF THE MISSOURI BOTANICAL GARDEN rather square in cross т alpine plant....... 4 . DIVARICATA Var. VISCIDULA 91. дан pos herbage Нена to villosulous; flow anthesis М pr e r, terete; loma plants. . DIVARICATA Var. DIVERGENS 87. Petals exauriculate: 92. Minute plants, the pedicels about as long as the bracts; tubercles of the seeds simple........... 28. D. PRAECOX 92. Large plants, the pedicels much lon eral = bracts; tubercles of the seeds ‘secondarily tu late ip Sei var. STRICTA 86. Tass much = than the e . Petals exauriculat eeds Vue 36. D. ovATA br . Leaves long-pevolate ed ае saccate..... TE CATA var. DIVARICATA 96. Leaves subsessile; pen not saccate: 97. Sepals strongly nerved, carinate; leaves ften crenulate 41с. D. PAPOSANA Var. SERRULATA 97. Sepals weakly рео, ecarinate; leaves entire 42b ROTUNDIFOLIA Var. NITIDA 95. Plants villose or шо a 0. D. GRANDIFLORA 93. Petals auriculate. 8. Plants not villose or villosulous: 99. Leaves subsessile, often crenulat 41с. D. PAPOSANA Var. SERRULATA 99. Leaves long-petiolate, entire. 5a. D. DIVARICATA var. DIVARICATA 98. Plants villose or villosulous. 100. Leaves sessile or subsessile............... 40. D. GRANDIFLORA 100. Leaves long-petiolate 43a. D. DIVARICATA Var. DIVARICATA KEY TO THE SERIES OF DRYMARIA a. Leaves estipulate, pc as the Mum petals са Бы; gic lobes; seeds foetiform or hippocrepiform, rsally his s A: HOLOSTEOIDES (p. 186) a. Leaves, if gibts dni defen: seeds, H erse dorsally e pid: b. Leaves Ванг) P amps upright perennials; petals with many distal aS seeds foetifo dorsally hisp Series B: LYROPETALA i to. 189) b. Leaves x to reniform; plants annual or perennial; petals entire to асое seeds, if rarely foetiform, not dorsally hispid: . Leaves elliptic, o opposite, жари. pau. apically and ев polylacest, alter- nating with conspicuous clavat C: sTIPITATAE (р. 193) c. Leaves and petals various; stamin sd nei ae conspicuous nor clav 4. Seeds distally 3—8-lobed, «к if merely bifid, the trunk ‘much qe than the claw, laterally донео јаве: ва mooth to granular; Мо m America ries D: ARENARIOIDES (p. 193) d. 2. а bifid, бана к» the lobes sometimes emarginate or dichoto- ; seeds tuberculate (ex. D. viscosa and D. ovata): aves lin RT cillate or г is opposite, viscous; seeds smooth; dorsally roe facially transparent, а. «a yellowish embryo ы сти rnia....... E: OSAE х Р 200) е. Leaves linear to reniform; opposite, glabrous to ен ог ар в. culate (except D. ovata var.) reddish brown to blac 1961] DUKE—REVISION OF DRYMARIA 185 f. Petals with four equal lobes or with two dichotomizing lobes. Cuba and Mex xico. g. Stamens м Sec елы the petals with four — lobes; bracts more or less imbric ries F: ORTEGIOIDES ha 200) в. Stamens d ped Fs the petals bifid, the a охота; brac es С: EXCISAE a 201) remote, E om aciei bifid, the lobes rarely emarginate: eaves r to lance-elliptic (ovate in D. fasciculata), stipulate or si jen айке. тагоб often exunguiculate; seeds without ornate tubercles i, Petals unguiculate; delicate p wer or erect virgate annuals, ж. spreading and perennial. North Am $ ne Раме to oblong or spain? i gue рее plants laxly Н: LEPTOP reed Е 204) ГА с снна to elliptic, the petioles ni Plants m spending. ENUES (p. 210) i. Petals exunguiculate or subexunguiculate; suffrutescent erect oe or delicate annuals. Peru and Ecuador. К. Large-flowered suffrutescent ee the leaves occasionally esti late; flowers occasionally dimorphic.................. Series J: FRUTESC deris "ik 212) k. 2 flowered delicate аслу (exc. D. fascic dae cd which may ga to Ла arge-flowered and ре аан leaves s £3 ч е Series етич y^ 216) + ovate to reniform, stipulate; petals (rarely келгін ак often h. Leaves auriculate; seeds occasionally with ornate tubercles, rarely smooth and lustrous L Villose и bei long-petiolate — bt simply bifid, exceeding the sepals; sta all fertile, unequal. a California...Series L: DEBILES (p. 220) Е reg re to егей еч with sessile to rie petiolate leaves; petals bi ate or pone stamens 2—5, if unequal, the the lobes sometimes emargin andis cs usually p to sterilization. m. Tr of the petals longer ЕЯ the а ge Me deeply emarginate, exauriculate. Western U. S. to Central Ame зе М: LAXIFLORES (р. 222) m. Trunk of the petals shorter than the claw (except D. верена), the lobes acute to с" occasionally basally auricu n. Lobes of the petals mostly with more than one vein, often basally dentate or —Ó the auricles in t: Is (except in apetalous forms) equaling or exceeding ve Mod: seeds usually numerous, with ма cles, o егі с ornate tuber the tubercles (absent in D. ta); stamens 5 with Е chine: the styles mostly three, men half their length or more; plants often villose with jointed esse o. Petals basally р малй — €—— (except in forms o . villosa with obtu cullat их кое of С see code selle cylindrie а or spe “Зең уе uth America; introduced i Seri x VILLOSAE (р. 223) о. Petals — or d auricles not ciliate; tubercles of the seeds nical, or rarely secondarily tubercululate; western p. Inflorescences often subcapitate, the pedicels if longer ond - s not divar rs sep. - often rig not bas ~ inate; о wit trace of leaves often ae “occasionally crenulate, а. PU densely villose; Colombia to Argent dit O: GRANDIFLORES (p. 230) p. Inflorescences lax, the pe: доде and divaricate; ` sepals acute or basally saccate, not carinate; pet ls with minute deltoid h -petiolat i ог mamm аан ricles; lea tire i doa not uae villose......... Series P: DIVARICATAE (p. 240) n. Lobes o diesen 1-n Me (except D. vosque и аралдан А екеніне, £ ostly shorter than the s (ex rice f 9 e petals ilis ek cpi has deltoid atone ds 1 ЕВ. the tubercles low and domical or уза ве Eius [Vor. 48 186 ANNALS OF THE MISSOURI BOTANICAL GARDEN remote; stamens and styles often reduced to 2, the anthers в. tending о be orbicular, and the styles free mer to their base; plants not villose Series О: CORDATAE (р. 245) A. Series HOLOSTEOIDES Leaves elliptic, pseudoverticillate, estipulate, often somewhat glaucous and succulent. Flowers in pseudoverticils. Sepals obtuse, often somewhat cucullate, the venation subdendritic. Seeds foetiform or hippocrepiform, merely granular, occasionally gibbose and umbonate. Two halophytic species of western Unite States and Mexico, both probably toxic to cattle. Fig. 1. a. Sepals 3-5 mm. long; petals 5-10-lobed; seeds 0.6-1.0 mm. long, about as broad as long; зыч glaucous, glabrous or glandular; Baja California: b. rada and stems often meh erar many, z not m the меская longer than 4 cm., with branching at mo nodes; petals 2.8—3.5 mm. long, often equaling or exceeding the sepals; seeds vun diae umbonate e рые D. HOLOSTEOIDES var. HOLOSTEOIDES b. Leaves and stems glaucous, rarely stipitate-glandular, often tinted with anthocyanins; most of the internodes shorter than 4 cm., with Ж anching at gt ew of the nodes; petals 2.0-3.5 mm. long, the sepals usually longer; seeds rarely g sige b. D. HOLOSTEOIDES Var. CRASSIFOLIA a. Sepals 2.0—3.5 mm. long; petals 4-lobed; seeds 1.8 mm. long, about twice as long a broad; herbage glabrous; continental Mexico us 5. г Stateb.os ы 2. D. ра АЙ la. DRYMARIA HOLOSTEOIDES var. HOLOSTEOIDES үнөн ao Benth. Bot. Voy. Sulph. 16. 1844. (HOLOTYPE: Hinds 5. п. т 1841 Drymaria veatebil Curran, e өмір Cal. Acad. 21:227. 1888. (HOLOTYPE: Veatch іп Майне» bolosteoides M. E. Jones, Extr. Contr. West. Bot. 18:35. 1933. Radially spreading herbaceous to lignescent annuals to as much as 40 cm. long, branching at nearly every node, stipitate-glandular to glabrescent, the internodes mostly much longer than the leaves, with a caducous rosette of spatulate leaves. Cauline leaves pseudoverticillate, the blades glabrous or scantily glandular, occasion- ally somewhat glaucous, narrowly to broadly elliptic, apically rounded or acutish, basally acute, 4-12 mm. long, 2-7 mm. broad, the petioles glabrous to stipitate glandular, 2-8 mm. long; stipules absent. Inflorescences of terminal and axillary contracted umbelloid cymes, subtended by both normal foliage leaves and scariose ovate bracts 1-2 mm. long; pedicels 2—6 mm. long, glandular or stipitate-glandular. Sepals 5, glandular or stipitate-glandular, broadly oblong, obovate or suborbicular, apically obtuse and often cucullate, 3-4 mm. long, 1.6—3.0 mm. broad, the venation obscure, dendritic; petals 5, 2.8-3.5 mm. long, bifid for about half their length, with 4—6 laciniae in the cleft, these about half as long as the oblong lobes, the trunk suborbicular, denticulate or crenulate, cordate or truncate to the minute claw; stamens 5, ca. 3 mm. long, the oblong anthers ca. 0.6 mm. long; ovary at anthesis globose, the styles attaining the anthers, bifid or trifid less than a third of its length. Capsule subglobose, 1.5-3.0 mm. long, 6—20-seeded, the seeds foeti- form, basally umbonate, laterally gibbose and often dorsally flanged, granular, 0.5— - 19611 DUKE—REVISION OF DRYMARIA 187 ACD bingy sete ote var. bolosteoides (holotype of D. veatcbii); gno 77; Ж; sepal, : их wed | 5 D. holosteoides var. crass: pi (holorype); petal, 10X; sepal, му ; seed, р. л (holotype) petal, 10Х; sepal, 10Х; stamen, 10X; ке 10X; habit, 1.0 mm. long, nearly or quite as broad as long. Fig. 1A (Holotype of D. veatchii). Sandy soils of Baja California, Mexico, mostly north of Cape San Lucas. The isotype at Leningrad has unusually long styles to 2 mm. long which are divided less than one-third their length, a character more frequent in the var. crassifolia. Indeed the type is in some other respects (as well as aspect) inter- mediate between these closely related varieties but by far the majority of its characteristics are those of the more northern populations. Among other inter- mediate specimens which forced me to reduce the following species to a variety may be cited Constance 3185, which in its strongly gibbose seeds and stipitate- glandular stems approaches the typical variety. However the specimen seems to possess more characteristics of var. crassifolia, e. g. the petals are much shorter than the sepals, the internodes are relatively short and the plant is obviously a perennial. 1b. DryMaria HOLOSTEOIDES var. crassifolia (Benth.) J. Duke comb. & stat. nov. p sce етеш ойе Бан | Bot. Voy. Sulph. 16. 1844. (HOLOTYPE: Hinds s.m. іп 1841; s;K! pes at GH, LE МОНЫ кенеді М. LE. Jones, M Contr. West. Bot. 18:35. 1933. (Мог. 48 188 ANNALS OF THE MISSOURI BOTANICAL GARDEN Subcespitose herbaceous to lignescent perennials verticillately branching but rarely branching at every node, glabrous or very rarely stipitate-glandular, to as much as 20 cm. long, some of the internodes occasionally shorter than the leaves, with a caducous rosette of spatulate leaves, the taproot to 8 mm. broad. Cauline leaves pseudoverticillate, the blades glabrous, carnose, glaucous, often cyanic, narrowly to broadly elliptic, 4-12 mm. long, 3-8 mm. broad, the petioles 4—12 mm. long, glabrous or glandular; stipules absent. Inflorescences of terminal and axillary contracted umbelloid verticils subtended by normal foliage leaves and by bracts 1-2 mm. long; pedicels 5-12 mm. long, glabrous to stipitate-glandular. Sepals 5, glabrous to scantily sessile-glandular, broadly oblong to elliptic, apically obtuse and often cucullate, 3.0-4.5 mm. long, 1.5-2.5 mm. broad, the venation obscure, dendritic; petals 5, 2.0-3.5 mm. long, bifid for about half their length, with 4—6 laciniae in the cleft, these about half as long as the main lobes; trunks ovate, denticulate, cordate or truncate to the minute claw; stamens 5, ca. 3 mm. long, the oblong anthers ca. 0.6 mm. long; ovary at anthesis subglobose, the styles attain- ing the anthers, bifid or trifid less than a third of their length. Capsule subglobose to ovoid or ellipsoid, 3—5 mm. long, 7—20-seeded, the seeds hippocrepiform, granular, scarcely umbonate or gibbose, 0.8-1.0 mm. broad, nearly or quite as long as broad. Fig. 1B. (Holotype). Saline soils around Cape San Lucas, Baja California, Mexico. = DryMARIA PACHYPHYLLA Wooton & Standl. in Contr. U. S. Nat Herb. 16: 121. 1913. (HOLOTYPE: Wooton 405, dry plains south of the White Sands, N. Mex., US; isotypes at F, GH, MO, UC, etc.!) Glaucous subsucculent annuals, the vegetative branching largely confined to radially diverging branches from a slender yellowish rootstock or to terminal pseudoverticillate branch systems, the elongate spreading internodes much exceed- ing the terminally crowded leaves. Leaves pseudoverticillate, glaucous, subsuccu- lent, wrinkling in drying, broadly elliptic to suborbicular, apically obtuse or acutish, basally tapering to the petiole, 4-14 mm. long, 4-12 mm. broad, the glabrous petioles 4—8 mm. long; stipules absent, the petioles clasping. Inflorescences of terminal and axillary contracted umbelloid verticils subtended by normal foliage leaves and bracts 0.5-1.5 mm. long, the pedicels 1-5 mm. long, the bracts ovate, obtuse, scariose and almost nerveless. Sepals 5, subequal, glabrous, glaucous, ellipsoid, obtuse, 2.5—3.3 mm. long, 1.5-2.0 mm. broad, obscurely 3—5-nerved, the central portion green, the margins scariose; petals 5, 2.5-3.0 mm. long, bifid about half their length, with 2 oblong laciniae in the cleft, the trunk serrulate, tapered to the base, the claw not clearly delineated; stamens 5, 1.0-1.5 mm. long, the oblong anthers ca. 0.5 mm. long; ovary at anthesis subglobose, the short style bifid or trifid more than half its length, slightly exceeding the anthers. Capsule sub- globose, 3-4 mm. long, mostly exceeding the sepals, 15—25-seeded, the seeds vermiculiform, facially gibbose, more or less tessellate, 0.8-1.8 mm. long, ca. twice as long as broad. Fig. 1C (Holotype). Heavy saline soils in denudated or ruderal sites, often as a pioneer on bare areas, western U.S. A. and northern Mexico. „ЦРНА кк анан 19611 DUKE—REVISION OF DRYMARIA 189 Although certain authorities have regarded this species as synonymous with D. holosteoides, there seems to be no justification for such a conclusion, as the mainland species differs constantly in many respects from the preceding species of Baja California. The seed alone would suffice to distinguish this species from all other species of Drymaria. Little (in Ecology 18:416. 1937), who regards D. pachyphylla as a synonym of D. holosteoides, presents some very interesting information about the ecology and toxicity of this serious weed. All the aerial portions of the plant are toxic to cattle and sheep. To complicate matters, the xeromorphic nature of the leaves is responsible for a degree of resistance to drought and to chemical sprays. Campbell (in Journ. Agr. Res. 43:1027. 1931) has shown the species to be a pioneer in plant succession on clay soils such as the adobe soils of portions of our western ranges. B. Series LYROPETALAE Leaves linear, pseudoverticillate, estipulate or with entire caducous stipules. Flowers in pseudoverticils or in cymes. Sepals obtuse or acute, the venation sub- dendritic. Petals apically 6—14-lobed, laterally denticulate, minutely unguiculate. Seeds hippocrepiform or foetiform, dorsally hispidulous, etuberculate. Four gypso- phytic species of the Coahuilan Desert of northeastern Mexico. Fig. 2. a. Inflorescence pseudoverticillate, compact, the bracts prd pap dere petals жал 6-8- lobed, the lobes lance-deltoid, attenuate, about half as long as “1 runk, the trunk осса- — bili inguate; | staminodia ligular or obsolescent; labs shop Basa leaves 10-70 mm. long, without true Nar m (suppressed branches or AS inside the leaf bases may ко confused торо stipules) : b. Sepals 3.0 . long, often glandular; petals 2.5—3.5 mm. long; stamens 2-3 mm. pue the шор е staminodia 0.2-0.5 mm. long, 0.5-0.8 mm. broad; bracts 0.5-1.5 mm. 3 X ELATA b. Sei = mm. long, glabrous or bullate; petals 4.5—6.0 mm. long; stamens 3- mm. ‚ the мы obsolete or absent; bracts 2.5-4.0 mm. long.......4. D. SUB енді 2. Inforscence cymose, lax, the bracts cider anie petals г pc the lobes linear, obtuse, outer nearly or quite as long as the trunk, the tru guate; staminodia cupular; sty паза conspicuous іп кета анан Задна gen mm. long, with minute EE setae stipules: c. Hairs on the seeds dense, erect, 0.3—0.5 mm. long; trunk of the petals nearly “ я as long, ве ыр и the pod the ук laciniae subequal іп length; S long; ts 3-5 mm. long; glabro D ишу РВА с. Hairs on т seeds scanty, о 0.2-0.3 (-0.5) mm. long; epa the petal nearly ce as long as broad, basally cordate or truncate, the apical айза нес эг на. bracts 0.5—3.5 mm. long; plants glandular. 6. D. LYROPETALA 3. Drymaria ELATA І. M. Johnston, in Jour. Arn. Arb. 21:68. 1940. (ного- : I. M. Johnston 7823; 10 km. s. of Laguna del Rey, locally abundant in gypsum silt; Coahuila. GH!) Upright lignescent perennials, the branches opposite or pseudoverticillate, glabrous or glaucous to stipitate-glandular, the internodes mostly longer than the leaves, the tap root to 1 cm. thick. Leaves opposite or pseudoverticillate, glabrous or glaucescent, succulent, obscurely veined, linear, apically obtuse, 1-7 cm. long, 0.5—1.5 mm. broad. Inflorescences of terminal contracted 5—13-flowered umbelloid cymes, the peduncles mostly 5-8 cm. long, glabrous; bracts 0.5-1.0 mm. long, [Уот.. 48 190 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 2: А. D. ae ote nr m sepal, 10 Х ; seed 15X. н, (holo- type); petal, 6X; sep: С. D. suffruticosa (holo Ж d 5X3 stamen, 5X; seed, “Bx hcc D. ed petala var. lyropetala сину ine ia 745 ла * sepal, 7V2X; stamen, 72 Х d, 10X. Е. D. lyropetala var. coahuilana (holotype); petal, 712X; sepal, 7 V2 X ; seed, 15x; ‘habit, YX. 1961) DUKE—REVISION OF DRYMARIA 191 deltoid-ovate, the mature pedicels 4-7 mm. long, puberulent or stipitate-glandular, the outer ones spreading or deflexed. Sepals 5, 3.0—4.5 mm. long, 2-3 mm. broad, stipitate-glandular or glabrate, broadly ellipsoid to suborbicular, apically obtuse, the venation obscure, with 7-9 veins arising from the base, all but the midrib dichotomizing; petals 5, 2.5—3.5 mm. long, with about 6 subequal apical laciniae, these lance-deltoid, falcate, attenuate, and denticulate, the trunk longer than the laciniae, apically bilinguate, laterally lacerate, ca. 1.5 mm. broad, basally cordate, the claw binute; stamens 5, 2-3 mm. long, the oblong anthers 0.5-0.8 mm. long; staminodia conspicuous, flap-shaped, 0.4 mm. high, 0.8 mm. broad; ovary at anthesis ovoid, the style slightly exceeding the anthers, trifid less than one-fourth of its length. Capsule subglobose, exceeding the sepals, 3.0-4.5 mm. long, 10- 20-seeded, the seeds hippocrepiform, 1.0-1.5 mm. long, densely hispidulous dorsally, ventrally slightly umbonate, the facies merely puncticulate. Fig. 2A. (Holotype). Known only from saline soils in the vicinity of Sierra del Rey and Laguna del Rey in Coahuila, Mexico. The petals are completely unlike those of any other species and suggest the dorsal profile of a beetle. This peculiar aspect is approached, but only slightly, by the petals of the following species. 4. DRYMARIA SUBUMBELLATA I. М. Johnston, in Jour. Arn. Arb. 31:188. 1950. (HOLOTYPE: I. M. Jobnston 8480, s. end of Cañada Oscuro near Tanque Ја Luz, confined to gypsum beds on the escarpment, western Coahuila; GH!) Upright or ascending lignescent perennials to as much as 25 cm. high, the branches mostly opposite, the internodes longer or shorter than the leaves, glabrous or glaucescent, the taproot to 1 cm. thick. Leaves opposite or pseudoverticillate, the blades 1.5—4.0 cm. long, 0.5—1.0 mm. broad, glabrous or glaucescent, succulent, linear, apically obtuse, attenuate to the sessile base; true stipules absent. Inflores- cences of terminal subumbelloid 3—9-flowered cymes, the peduncles 4—8 cm. long, the bracts mostly imbricate, lance-ovate, 2.5—4.0 mm. long; mature pedicels 3-7 mm. long, glabrous, the outer ones spreading. Sepals 5, 5—6 mm. long, 2—3 mm. broad, glabrous or bullate, broadly ovate, apically obtuse or acutish, the margins involute, the venation obscure, with 7-9 veins arising from the base, all but the midrib dichotomizing; petals 5, 4.5—6.0 mm. long, with 6-8 subequal apical laciniae, these lance-deltoid, attenuate, falcate, denticulate, the trunk longer the laciniae, laterally lacerate, basally truncate to rounded, 1.0-2.0 mm. broad; stamens 5, 3.5—4.0 mm. long, the oblong anthers 0.8-1.0 mm. long; staminodia obsolescent; ovary at anthesis ovoid, the style slightly exceeding the anthers, bifid or trifid less than one-fourth its length, nearly as long as the ovary. Capsule subglobose, 5-6 mm. long, exceeding the sepals, 10—20-seeded, the seeds hippo- crepiform, 1.2-1.5 mm. long, dorsally provided with dense white hairs to 0.5 mm. long, ventrally umbonulate, the facies smooth or puncticulate. Fig. 2B. (Holotype) Known only from the type collection. In the nature of the inflorescence and the petals, this bizarre species is inter- mediate between D. elata and the other two species in this homogeneous series. [Vor. 48 192 ANNALS OF THE MISSOURI BOTANICAL GARDEN мл . DRYMARIA sUFFRUTICOSA A. Gray ех S. Watson, in Proc. Am. Acad. 17:328. 1882. (HOLOTYPE: Palmer 74; San Lorenzo de Laguna, Coahuila; GH!; isotypes at US, MO, UC, etc.!) Upright suffrutescent glabrous perennials to as much as 30 cm. tall, the leaves often fascicled at the geniculate nodes, the taproot to as much as 8 mm. thick. Leaves pseudoverticillate, glabrous or glaucescent, somewhat succulent, linear, obtuse, 10-25 mm. long, са. 1 mm. broad, basally tapering to the stem; stipules apparently lacking. Flowers globose in few-flowered terminal cymes, the peduncles and pedicels glabrous; bracts lanceolate, 3—5 mm. long, about equaling the pedicels. Sepals 5, 5—6 mm. long, 3—4 mm. broad, broadly ellipsoid, apically rounded but with a minute acumen, strongly venose throughout, the small scariose margins involute; petals 5, 4.5-5.5 mm. long, са. 2 mm. broad, with 10-14 subequal apical laciniae, laterally lacerate, basally truncate or tapered to the minute claw; stamens 5, 4—5 mm. long, the oblong anthers ca. 1 mm. long; staminodia cupular, minute; ovary at anthesis renoid, the minute bifid style slightly exceeded by the anthers. Capsule broadly ellipsoid, ca. 6 mm. long, often capped by a stylopodium, many-seeded, the seeds hippocrepiform, ca. 0.9 mm. broad, dorsally provided with a row of stiff hairs to 0.5 mm. long. Fig. 2C. (Holotype). Known only from the Coahuilan Desert of Mexico. 6. DRYMARIA LYROPETALA I. M. Johnston, in Journ. Arn. Arb. 21:68. 1940. (HOLOTYPE: І. М. Johnston 7504; 3.5 km. s. of Cedral, gypsum plain, locally common, GH!) Drymaria вн var. coabuilana 1. M. Johnston, in Jour. Arn. Ar b. 189. 1950. ER tewart wid icis as Jobnston 567 in original ИОВ 2 km. s. of а lius, Sierra de las Cruces, Gypsum Ridge, Coahuila, GH!) Upright or ascending glandular or glabrous perennials to as much as 50 cm. high, branching mostly at the base, the internodes shorter to longer than the leaves, apparently elongating belatedly, the taproot to 1 cm. thick. Leaves opposite or pseudoverticillate, 3-15 mm. long, 0.5-1.2 mm. broad, glabrous or glandular, glaucescent, succulent, linear, apically obtuse, attenuate to the sessile base, the upper leaves minutely stipulate. Inflorescences of terminal lax 3- 9-flowered cymes, the peduncles 4—60 mm. long, glabrous to stipitate-glandular, the bracts 0.5-3.5 mm. long, remote, the mature pedicels 3-6 mm. long, glabrous or glandular, not recurved. Sepals 5, 3-5 mm. long, 1.5-3.0 mm. broad, glabrous to stipitate-glandular, ovate, apically obtuse, the venation obscure, with 7-9 veins arising from the base, all but the midrib dichotomizing; petals 5, 3.2-5.2 mm. long, with 10-12 unequal apical laciniae, these linear, obtuse, crenulate, the outer- most as long as or longer than the trunk, the trunk laterally lacerate, basally cordate to truncate, 1-2 mm. broad; stamens 5, 3-6 mm. long, the oblong anthers 0.7-1.0 mm. long; staminodia cupular, to 0.4 mm. high; ovary at anthesis sub- cylindric, the style about as long, 1.5—2.5 mm. long, trifid about one-fourth its length; stylopodium well-developed. Capsule globose, 3.0-3.5 mm. long, 3- 24-seeded, the seeds hippocrepiform, 1.0-1.5 mm. broad, ventrally umbonulate, 1961] DUKE—REVISION OF DRYMARIA 193 dorsally hispid, the hairs 0.2-0.3 (-0.5) mm. long, scanty and spreading. Fig. 1D. (Holotype of var. lyropetala) ; Fig. 1E (Holotype of var. coabuilana). Local on gypsum soils of the Coahuilan Desert of Mexico. The two varieties of this species may be separated by the following key: a. Bracts 2-3.5 mm. long; leaves 5-15 mm. long, shorter than the internodes; me. арени to stipitate-gla OAHUILANA а. Bracts 0.5—1.5 mm. long; leaves 3-6 mm. long, shorter to longer than the internodes plants glabrous or with a few sessile glands var. LYROPETALUM Johnston erred very slightly in his description of the more southern var. lyropetala, stating that it was glabrous, the glandulosity occurring only in the northern variety. The type of var. lyropetala has a few scattered glands. It should be noted here that the UC isotype has leaves which are mostly longer than the internodes while the converse is true of the holotype, which incidentally is less conspicuously glandular. C. Series STIPITATAE M DRYMARIA STIPITATA Fosberg, in Lloydia 4:281. 1941. (HOLOTYPE: С. Н. Muller 3301; Mun. de Sierra Mojada, Sierra Mojada, Cafion de San Salvador; common in high oak chaparral; Coahuila; US!) Upright suffrutescent perennials to as much as 20 cm. high, glabrous below, becoming stipitate-glandular above. Leaves opposite, the blades 5-15 mm. long, 1.5—5.5 mm. broad, glabrous, somewhat carnose, almost veinless, narrowly ovate to elliptic, apically acute, basally acutely tapering to the petiole, the petiole 1-2 mm. long; stipules apparently lacking. Inflorescences of few-flowered terminal cymes, the axes glandular-puberulent, locally densely so; bracts lance-ovate, 1.5-2.5 mm. long, apically acuminate; pedicels mostly 3-5 mm. long. pals 5, subequal, lanceolate to ovate, apically acuminate, basally umbonate, 5-6 mm. long, 1.5-2.5 mm. broad, transparent save for the 3 ribs; petals 5, 5.0-5.5 mm. long, apically and laterally lacerate, the laciniae linear, often dichotomous, some of the lateral laciniae directed downwardly; stamens 5, 5.0-5.5 mm. long, the broadly oblong anthers ca. 0.8 mm. long, alternating with 5 clavate staminodia to as much as 2.2 mm. long; ovary at anthesis ellipsoid, much shorter than the style, the style trifid for ca. one-fifth its length, exceeding the anthers. Capsule са. 4 mm. long, 6—8-seeded, the seeds 0.8—1.0 mm. long, symmetrically hippocrepiform, the surfaces irregularly corrugated. Fig. 3A (Holotype). Coahuila, Tamaulipas and probably Nuevo Leon, Mexico. D. Series ARENARIOIDES Leaves linear to narrowly ovate, opposite or pseudoverticillate, subsessile, stipu- late. Flowers in cymes, racemes or solitary in the axils of slightly reduced foliage leaves. Sepals mostly obtuse and weakly nerved. Petals 2-8-ІоБед, the trunk much longer than the claw, entire or denticulate. Seeds hippocrepiform and dorsally sulcate or lacrimiform, nearly smooth or granular. Five species in western U. S., northern Mexico and Baja California. Figs. 3B, 3C, 4. (Vor. 48 194 ANNALS OF THE MISSOURI BOTANICAL GARDEN a. Leaves ы: рид ин, glabrous; flowers cymose, the bracts clearly ee ies the foliag D. MOLLUGINEA a. Leaves linear to ovate, opposite or aea p e sqq glabrous to densely Е bra ee tending to be racem the often not different from foliage leaves. . Plants НЕ. Pid in the ns of scarcely reduced foliage leaves or racemose: c. Flowers racemose, the petals mostly 4—6 lobed; leaves linear to oblon d. Petals 4.0—6.2 mm. long, mostly 6-lobed, = middle lobes dd as long as outer; sepals 47 m POR yles 1.0-2.5 mm. long, united more than half Bes length; continental Mex 9a. D. SR ssp. ARENARIOIDES d. Petals 2.8—4.0 mm. long, 2—6-lobed, the pu lobes about half as long as the outer; sepals 2.8—4.0 mm. long; geras 2.8—4.5 mm. long, united 5 their ње or less Baja California 9b. D. p. PENINSULARIS c. Flowers axillary, the petals merely bifid; leaves ovate 10. D. AXILLARIS b. Plants without га glands; flowers іп сут е. ee = r to oblong; petals occasionally enti, briefly unguiculate; seeds ее tuber D. BARKLEY! е. йан 4. to ovate; petals bifid, exunguiculate; seeds yellowish, aid мй “кз D. POLYCARPOIDES 8. DRYMARIA MOLLUGINEA (Lag.) Didr. in Linnaea 29:738. 1859. (ното- TYPE: "Lagasca 1806” annotated by De Candolle in the De Candolle Her- barium; С. я. v.; photograph and drawing of type at СН!) Alsine molluginea Lag. Gen. & Sp. 13. Arenaria ? molluginea Ser. ex DC. Prodr. 1: Eu: Drymaria sperguloides A. Gray, Pl. Fendler Hn 1849. (HoLorvpr: Fendler 55; New Mexico, GH!; isotypes at US, UG; F, Mdb Lepigonum оба inea Fries, Ind. Hort. Sem. Mollugophytum sperguloides (A. Gray) M. E. Је: Extr. Contr, West. Bot. 18:35. 1933. Slender erect virgate or dichotomously branched annuals to as much as 20 cm. high, the internodes shorter to longer than the leaves, glabrous or with sessile glands, the tap root ca. 1 mm. thick. Leaves mostly pseudoverticillate, glabrous or scantily glandular, 9-25 mm. long, 0.5—2.0 mm. broad, linear, apically obtuse, attenuate to the sessile base, the stipules lance-deltoid, 1-2 mm. long. Inflores- cences of terminal cymose racemes, only the first 1 or 2 branches cymose, subse- quent branching tending to be racemose, the peduncle 8-30 mm. long; bracts 1.0-2.5 mm. long, the pedicels 2-5 mm. long, glabrous to minutely stipitate- glandular. Sepals 5, 2.5—3.5 mm. long, glabrous or minutely glandular, oblong, apically obtuse and often cucullate, only the midrib prominent; petals 5, 1.7-2.5 mm. long, 0.4-0.7 mm. broad, apically provided with usually 4 lacinae, the outer one-third to one-half as long as the trunk, the inner occasionally absent or reduced to mere dentations, the trunk subdeltoid, laterally denticulate, truncate or tapered to the minute claw; stamens 5, 1.5-2.0 mm. long, the oblong to suborbicular anthers 0.4—0.5 mm. long; staminodia absent; ovary at anthesis globose, са. 2 mm. long, the style 2-3-cleft nearly to the base, less than 1 mm. long; stylopodium absent. Capsule subglobose, 2-3 mm. long, exceeding the sepals, 3—17-seeded, the seeds 0.9-1.4 mm. broad, dark brown or purplish in age, hippocrepiform, dorsally flat, scarcely umbonate ventrally, minutely corrugated or tuberculate, the tubercles broader than long. Fig. 3B (Holotype of D. sperguloides) ; Fig. 3C ("”. molluginea Fenzl” at LE). 1961] DUKE—REVISION OF DRYMARIA 195 Sandy soils, Arizona to western Texas, U. S. А.; south to Puebla, Mexico. The specimen from Leningrad (Fig. 3C) apparently from the Ledebour her- barium, and labeled “Drymaria molluginea Fenzl” lacks the inner laciniae of the petals. I doubt that this feature has any taxonomic significance. A close parallel is exhibited by D. arenarioides ssp. peninsularis where the form which lacks the two central laciniae has been described as D. johnstonii. In both D. molluginea and D. arenarioides ssp. peninsularis specimens have been examined in which 1 or 2 of many flowers dissected had the inner laciniae reduced to mere dentations or apparently completely lacking. oe et tata (hol habit, ШХ; petal, 6X; 6X; stamen, 6X ; staminode, 6X; тан a oe | areas Мт э и P ulate): "m 1214 X, seed, 15X; sepal, вх. C. D. кй Г ‘D. molluginea ex Fenzl.” at LE); habit, УХ. (Мог. 48 196 ANNALS OF THE MISSOURI BOTANICAL GARDEN 9a. DRYMARIA ARENARIOIDES ssp. ARENARIOIDES Drymaria arenarioides Humb. & Bonpl. ex Roem. & Schult. Syst. Veg. 5:406. 1819. LoTYPE: Humb. & Bonpl. 4070 ad Pachuca, probably destroyed at Berlin; isotype ӘСЕ! Pies. frankenioides HBK. Nov. Gen. & Sp. 6:21. pl. 515. 1823. Prostrate radially spreading herbaceous or lignescent perennials to as much as 20 cm. long, branching mostly at the base, the internodes shorter to longer than the leaves, stipitate-glandular, the taproot to as much as 5 mm. thick. Leaves opposite or pseudoverticillate, stipitate-glandular, the blades linear-oblong to narrowly elliptic, apically obtuse, obscurely 1-пегуед, 5—15 mm. long, 1-3 mm. broad, basally tapered to the petiole, the petiole ca. 1 mm. long; stipules lance-deltoid, 1.0-2.5 mm. long, entire. Flowers, except the first formed, solitary in the axils of slightly and gradually reduced foliage leaves, the mature pedicels 3-15 mm. long, stipitate-glandular, the subtending leaves stipulate. Sepals 5, 4-7 mm. long, 1-3 mm. broad, stipitate-glandular, oblong to broadly elliptic, the venation obscure, subdendritic, the inner sepals shorter, broader and less glandular; petals 5, 4.0-6.5 mm. long, apically (4-) 6-8-ІоБаге, the outer lobes slightly longer and broader, about as long as the trunk, the inner lobes linear, equal, the trunk laterally denticu- late, 0.8-1.5 mm. broad; stamens 5, ultimately 3-5 mm. long, the oblong anthers 0.8—1.2 mm. long; ovary at anthesis ellipsoid, the style bifid or trifid less than half its length, slightly exceeding the stamens; stylopodium often conspicuous. Capsule ovoid, 3-5 mm. long, 15-25-seeded, the seeds hippocrepiform, dorsally flattened or sulcate, exumbonate, minutely tuberculate, 0.8—1.2 mm Sonora, Chihuahua, Zacatecas, San Luis Potosi, Guanajuato and Hidalgo, Mexico, usually in sandy soils. It would appear that Humboldt and Bonpland or Kunth had some reason for changing the name D. arenarioides to D. frankenioides as Roemer and Schultes cited the name as follows: “D. arenarioides Humb. et Bonpl. . . . Reliqu. Willd. MS". I would judge that Willdenow had written the name "D. arenarioides Humb. & Bonpl." on his herbarium specimen and that Roemer and Schultes were the first to publish the name, probably furnished earlier by Humboldt and Bonpland. Two other names in Drymaria seem to have the same history, i.e. D. ovata and D. stel- lericides., The question then arises whether the name should bear the authority umb. & Bonpl in Roem. & Schult.", "Humb. & Bonpl: Willd. ex Roem. & Schult.” or “Willd. ex Roem. & Schult.” Most publications appear to make use of the latter, but I feel it more proper to credit the name to Humboldt and Bon- pland as did Willdenow in his herbarium. At any rate, D. arenarioides and D. frankenioides are names based on the same specimen (typonyms). In its diminished leaves, sepals and petals, Parry & Palmer 49 from San Luis Potosi, approaches D. arenarioides subsp. peninsularis, but the petals, which seem to possess more constant characters, are like those of typical arenarioides, i.e. two larger outer lobes and two basally dichotomous inner lobes, the linear divisions of the inner lobes being nearly as long as the oblong outer lobes. The very long style of this specimen further corroborates its determination as D. arenarioides ssp. arenarioides nee ere ae RMSE >= Сы ise, ар аны s cose, ми ава felit dtd a La m 5; 197 19611 DUKE—REVISION OF DRYMARIA ЫҚ АР рар I Se M f. ` ы sep: A > pe eed); peta tal, 10 Ж 16 pal, 10X; ; seed 1214 a 5 Ик жж А pus Aes Р, 4 20x. E. і (holotype, ex haba, » 5; Е. D. жынына (holotype, dpt фест petal, 10X; ub 10X ; seed, 2 [Vor. 48 198 ANNALS OF THE MISSOURI BOTANICAL GARDEN 9b. DRYMARIA ARENARIOIDES ssp. peninsularis (S. F. Blake) J. Duke comb. & stat. nov. Drymaria peninsularis S. F. Blake, in Jour. Wash. Acad. 14:285. 1924. (HOLOTYPE: Purpus 423; Cape Region, Baja California, US!; isotypes at UC, F, GH, MO Drymaria ET KE in Proc. Cal. Acad. 425:203. 1944. (HOLOTYPE: I. M. Jobnston 3972; in crevices of rock on mesa near crest of island, the isthmus, Espiritu Santo Mind! Baja California, CAS!) Radially spreading or ascending annuals or suffrutescent perennials to as much as 25 cm. long, branching mostly at the base, the internodes shorter to longer than the leaves, stipitate-glandular, the taproot to as much as 1 cm. thick. Leaves opposite or pseudoverticillate, stipitate-glandular, linear, obtuse, obscurely veined, 4—25 mm. long, 0.5—1.0 mm. broad, attenuate to the sessile base, the stipules lance- deltoid, entire, 0.5—1.5 mm. long. Flowers, except the first formed, solitary in the axils of slightly and gradually reduced foliage leaves, the mature pedicels 4-12 mm. long, stipitate-glandular, the subtending leaves stipulate. Sepals 5, 2.8-4.5 mm. long, 1.0-2.2 mm. broad, stipitate-glandular, oblong to broadly elliptic, obtuse, the venation obscure, subdendritic, the inner sepals shorter, broader and less glandular; petals 5, 2.8-4.0 mm. long, apically 4(—8)-lobate, the outer lobes longer, about as long as the trunk, the inner about half as long as the outer, the trunk laterally denticulate, basally truncate or tapered to the minute claw; stamens 5, ultimately 2-3 mm. long, the oblong anthers 0.5-0.8 mm. long; ovary at anthesis obovoid, stipitate, the minute style slightly exceeded by the anthers. Capsule globose, 2.5—4.0 mm. long, the style less than 1 mm. long, trifid about half its length, the seeds 10-20, hippocrepiform, dorsally flattened or sulcate, scarcely umbonate ventrally, reticulate or minutely tuberculate, 0.6-1.0 mm. long. Fig 4B (Holotype of D. peninsularis) ; Fig. 4D (Holotype of D. johnstonii). Sandy soils; Cape Region of Baja California, Mexico. Unquestionably this subspecies is siete related to ssp. arenarioides. The two subspecies are nicely separated by the Gulf of California in addition to the diagnostic characters listed in the key. d have seen many specimens of both subspecies, and intermediates are few indeed. Craig 738 and Howell 10576 appear to represent intermediates with many of the petals similar to those of the typical subspecies. Other features, coupled with the geography, seem to justify their relegation to ssp. peninsularis. 10. DRYMARIA AXILLARIS Brandegee, in Univ. Cal. Publ. Bot. 4:178. 1911. (HOLOTYPE: Purpus 4526; Sierra del Rey, Coahuila, UC!; isotypes at GH, US, F, MO, etc.) Upright glandular-pubescent, cyanic glaucous perennials to 10 cm. high with many suffrutescent branches arising near the base, the taproot to 6 mm. in diameter. Leaves opposite, cyanic, glaucous, densely glandular-pubescent with capitate hairs to 0.7 mm. long, broadly ellipsoid, apically obtuse to acutish, basally subcordate or tapered to the petiole, 3-12 mm. long, 2.5-8.0 mm. broad; veins not apparent; petioles 0.5-2.0 mm. long, the stipules entire, broadly deltoid, 0.5—1.0 mm. long. Flowers solitary in the axils of reduced leaves, the glandular-pubescent pedicels 1961) DUKE—REVISION OF DRYMARIA 199 ebracteate, 1.5—3.0 mm. long. Sepals 5, subequal, subsucculent, glandular- pubescent, broadly lanceolate to ovate, slightly concavo-convex, obtuse to acutish, 4-5 mm. long, 1.5-2.0 mm. broad; venation obscure, the margins scariose; petals 5, 3.5-4.0 mm. long, bifid about half their length, the lobes obtuse, са. 0.6 mm broad, exappendiculate, the claw 0.5—1.0 mm. broad; stamens 5, 3.0-3.5 mm. long, the oblong anthers ca. 0.8 mm. long; ovary at anthesis broadly ellipsoid, the style bifid about one-third its length, slightly exceeding the anthers. Capsule bivalved, ca. 2.5 mm. long, many-seeded, the seeds hippocrepiform, merely reticu- late or puncticulate, ca. 0.7 mm. broad. Fig. 4C (Isotype). Known only from the type locality in Coahuila, Mexico. 11. Drymaria barkleyi J. Duke and Steyermark sp. nov. Plantae annuae graciles ramis paucis internodis glaucis quam foliis longioribus radice non viso. Folia opposita laminis glabris glaucis bullatisque 4-10 mm. longis, 1.0-2.5 mm. latis, ellipticis apice obtusis subsessilibus stipulis minutis deltoideis caducis apice saepe bifidis. Flores solitarii in axillis foliorum vix reductorum pedi- cellis glabris 4-8 mm. longis. Sepala (4—) 5, 3.0—4.5 mm. longa ca. 1 mm. lata glabra oblongo-elliptica obscuriter venosa; petala 5, 2.5—3.5 mm. longa ultra medium bifida vel rariter trifida lobis oblongis obtusis emarginatisve denticulatis basi acutis unguibus minutis; stamina (4—) 5, 3 mm. longa antheris oblongis 0.8— 1.0 mm. longis; ovarium oblongum, 1-2 mm. longum saepe 10-ovulatum stylis ca. 1.5 mm. longis (3-) 4-lobis minutis. Capsulae non visae. Fig. 4E (Holotype [except seed] ). СОАНИША: low pointed hill, almost Dn of soil, apparently a scie shale, 25 mi. sw. of Monterrey, pope 6 Barkley 148 26 М. (НОГОТУРЕ: Е, isotype Fortunately another specimen with mature capsules is referable to the new species. Hernandez, Rowell & Barkley 16M531, collected on talc-like limy soil and shaly limestone, (in Nuevo Leon), 11 mi. w. of Santa Catarina differs only in having a few sessile glands оп the herbage. The capsule is yellow, 3—4-toothed, 2.5-3.0 mm. long, 4—8 seeded. The seeds are lacrimiform, са. 0.7 mm. in diameter, minutely and remotely tuberculate. Apparently the closest related species is the preceding with which the new species shares the cyanic tinting, solitary axillary flowers, and minute deltoid stipules. The differences far outnumber the strictly superficial similarities however. The sharply pointed long-beaked seed is slightly reminiscent of D. debilis. 12. DRYMARIA POLYCARPOIDES A. Gray, РІ. Fendl 12. 1849. (HOLOTYPE: Gregg s. n.; valley of Bolsón de Mapimí, Mexico; GH!) Prostrate or ascending profusely branching perennials to as much as 10 cm. long, branching mostly from the base, the nodes geniculate, fragile, the internodes mostly shorter than the leaves, glabrous or quite glaucous, the taproot to as much as 1 cm. thick. Leaves opposite, carnose, glaucous, often cyanic, elliptic, apically acute to obtuse, basally tapered, 5-12 mm. long, 2-5 mm. broad, weakly if at all nerved; petioles 0.5—1.5 mm. long, exceeding the minute, entire, lance-deltoid [Vor. 48 200 ANNALS OF THE MISSOURI BOTANICAL GARDEN stipules. Flowers in few-flowered strongly dichotomous cymes, the pedicels 1—4 mm. long, the bracts deltoid, obtuse, not scariose, 0.8-1.5 mm. long. Sepals 5, 3.5-5.0 mm. long, 1.5-2.0 mm. broad, glaucous, oblong to ovate, apically obtuse and often cucullate, the scariose margin as broad as or broader than the central 3-nerved portion; petals 5, 3.5—4.5 mm. long, bifid little more than half their length, the lobes broadly oblong, apically obtuse, 1—3-nerved, merging imper- ceptibly with the trunk, the trunk 1.0-1.5 mm. long, nearly as broad, exunguicu- late or subexunguiculate; stamens 5, the anthers 0.8—1.0 mm. long, the filaments 1.5-2.5 mm. long, devoid of staminodia; ovary at anthesis ovoid, equaled or exceeded by the style, the style trifid for less than one-fourth its length. Capsule ellipsoid to oblongoid, 2.0-2.5 mm. long, few-seeded, the seeds lacrimiform, not circinate, umbonulate ventrally, smooth, yellowish-brown. Fig. 4F (Holotype [except seed]). Known only from the Coahuilan Desert of Mexico. E. Series vISCOSAE 13. Drymaria viscosa 5. Watson ex Orcutt, in West. Am. Sci. 2:57. 1886. nomen nudum. S. Watson in Proc. Am. Acad. 22:469. 1887. (HOLOTYPE: Orcutt 1330, Socorro (“Зосопо”), n. Baja California, GH!; isotypes at UC, D, F, US, MO, NY, etc.) Prostrate radially spreading or ascending diffusely branched viscid annuals, the branching below the inflorescence mostly dichotomous, the taproot to as much as 5 mm. thick. Leaves pseudoverticillate or opposite, glandular-puberulent, linear, apically obtuse, basally tapering to the clasping petiole, 3—15 mm. long, 0.5—1.5 mm. broad, the petiole 1-3 mm. long; stipules acicular, caducous, 1.0-1.5 mm. long. Flowers campanulate іп 1—5-flowered cymes, the peduncles mostly glabrous; bracts deltoid, marginally ciliolate, 0.5—1.0 mm. long, the pedicels 0—3 mm. long, all but the central equaled or exceeded by the bracts. Sepals 5, 2.3-3.0 mm. long, 1-1.5 mm. broad, glandular-puberulent, oblong, obtuse, obscurely 3-nerved, the midrib subapically excurrent; petals 5, 1.8-2.2 mm. long, bifid about two-thirds their length, the lobes oblong, obtuse, tapered to the unguiculate base, exappendicu- late; stamens 5, ca. 2 mm. long, the broadly oblong anthers ca. 0.4 mm. long; ovary at anthesis ellipsoid, about equaled by the style, the style trifid about: half its length, barely exceeding the anthers. Capsule 2.0-2.5 mm. long, equaling or exceeded by the sepals, 5—15-seeded, the seeds cochleate, nearly smooth, 0.5-0.6 mm. broad, dorsally brown, facially transparent and colorless, the yellowish embryo visible. Fig. 5A (Isotype). Sandy soils of Baja California, Mexico. In habit, this plant is not unlike D. molluginea as pictured in Fig. 3D. F. Series ORTEGIOIDES 14. DRYMARIA ORTEGIOIDES Griseb. Cat. Pl. Cub. 21. 1866. (HOLOTYPE: Wright 2019; Cuba occ., GOET!; isotypes at F, GH, MO, US, etc.) Pinosia ortegioides (Griseb.) Urban, in Ark. Bot. 23A5:71. 1930. 1961] DUKE—REVISION OF DRYMARIA 201 Upright sparsely dichotomizing glabrous or glandular perennials to as much as 20 cm. high, the internodes much longer than the leaves, the tap root to 5 mm. thick. Leaves opposite, glabrous, linear to elliptic, apically acute to obtuse, attenu- ate to the subsessile base, 3-12 mm. long, 1.0-3.5 mm. broad, the stipules lance- deltoid, 0.3—0.8 mm. long, entire. Inflorescences of terminal condensed 5-21- flowered dichotomous cymes; peduncles 1.5-3.5 cm. long, the bracts 1-2 mm. long, imbricate or subimbricate, the glabrous pedicels 1.5-4.0 mm. long. Sepals 5, 2.5- 3.5 mm. long, glabrous, ovate to obovate, acute to obtuse, the venation obscure, dichotomous and reticulate; petals 3, 2.3-3.2 mm. long, twice dichotomous, the lobes oblong, obtuse, nearly as long as the trunk, exappendiculate, the trunk merging gradually with the claw; stamens 3, the oblong anthers ca. 0.5 mm. long; ovary subglobose, the styles 0.5-1 mm. long. Capsule globose, 1.5—2.0 mm. long, shorter than the sepals, 1—4-seeded, the seeds hippocrepiform, scarcely if at all umbonulate, closely tuberculate, 1.0-1.5 mm. long, dark brown. Fig. 5B, 5C. Sandy savannas of Cuba. G. Series EXCISAE Leaves deltoid-ovate to reniform, opposite, stipulate. Flowers long-pedicellate in lax cymes. Sepals acute or obtuse, trinerved. Petals bifid, the lobes themselves dichotomizing, ciliate-auriculate at their bases. Seeds cochleate, tuberculate, the tubercles domical or substellate. 3 species in Mexico. Fig 5 D-F a. cae PE. Kaore niis to glabrous; leaves deltoid-ovate to reniform, rounded to acute at petals apically 2—4-times dichotomous. b. meld гумене шей and petals са. 5 mm. long, the petals 4 — womens D. soc ce RUN b. Leaves Rico. to кем ог reniform; sepals 5.0—10.0 mm. long; n 6-12 m long, 2- ichot 16 D. EXCISA a. Plants ipie таваа to "silos gio ovate to reniform, езы сл at the base; pet tals 1— $ dichot if only once dichotomous, the vei ually the apex giving lee Е the ае dichotom - D. LONGEPEDUNCULATA 15. DRYMARIA HYPERICIFOLIA Briq. in Ann. Cons. & Jard. Geneva 14:369. 1911. (HOLOTYPE: Jurgenson 38, de Lecambre de Yolotepeque à Juquilla, au sud- ouest d'Oaxaca; Mexicum G!; isotype at K) Erect or ascending perennials (?) to as much as 25 cm. high, the internodes about as long as the leaves, stipitate-glandular. Leaves opposite, 8-22 mm. long, 5—14 mm. broad, narrowly deltoid-ovate, glabrous or minutely glandular-puberulent below, apically acute, marginally somewhat undulate, basally obtuse to acute, plinerved; petioles 1-2 mm. long, the stipules entire or bifid, the divisions lance- deltoid, 0.5—1.2 mm. long. Inflorescences of terminal 5—11-flowered cymes, the peduncles 2-3 cm. long, the bracts lanceolate, 1.5-2.5 mm. long, the pedicels stipitate-glandular, 4—6 mm. long. Sepals 5, 4.5—5.5 mm. long, lanceolate to narrowly ovate, apically acute, the outer sepals scantily stipitate-glandular, tri- nerved; petals 5, ca. 6 mm. long, four times dichotomous, the ultimate segments emarginate, the trunk laterally provided with deciduous ciliate auricles; stamens 5, ca. 4 mm. long, the anthers oblong, ca. 0.7 mm. long; ovary at anthesis ellipsoid, about equaled by the partially trifid style, the style slightly exceeded by the anthers. [Vor. 48 202 ANNALS OF THE MISSOURI BOTANICAL GARDEN Capsule ellipsoid, ca. 5 mm. long, 3-valved, apically constricted, many-seeded, the seeds cochleate, 0.7—1.0 mm. long, irregularly corrugated or tuberculate; seeds immature. Fig. 5D (Holotype). Known only from two collections near Juquila in Oaxaca, Mexico. North American authors have confused Drymaria ladewii with this species, probably because of certain vegetative resemblances. The flowers of the two species however contrast vividly. 16. Drymaria ЕХСІЅА Standl. in Field Mus. Bot. 8:11. 1930. (HOLOTYPE: Mexia 1748; Real Alto, trail to El Tajo de Santiago, 2500 m., Jalisco, F!; iso- types at UC, GH, etc.) rige iem grandis Bullock, in Kew Bull. 1936:389. 1936. к. Hinton 5427, Los Hornos, Temascaltepec, 2500 m., Mexico, K!; isotypes at GH, MO, etc.) a spreading perennials, the procumbent or erect branches to 3 m. long (fide Bullock), the internodes mostly longer than the leaves, glandular-puberulent, glabrescent. Leaves opposite, 5-25 mm. long, 5-30 mm. broad, glabrous or minutely glandular, the blades orbicular to obcordate, apically emarginate or apiculate, marginally entire, basally obtuse to truncate, 3-pli-nerved; petioles 5-25 mm, long, the stipules entire or apically lacerate, lanceolate, 2-4 mm. long. Inflorescences of terminal lax 3-many-flowered cymes, the peduncles 3-6 (-15) cm. long; bracts lanceolate to ovate, 3.0-4.5 mm. long, the pedicels 3-15 mm. long, stipitate-glandular, glabrescent. Sepals 5, 5.0-10.0 mm. long, 2.0-2.8 road, ovate, apically acute, weakly 3-ribbed; petals 5, 6-12 mm. long, 2—3-times dichotomous, the ultimate segments entire or emarginate, the trunk laterally provided with several deciduous ciliate auricles; stamens 5, 5.0-6.5 mm. long, the oblong anthers ca. 1 mm. long, the staminodia minute, semicircular; ovary at anthesis ovoid, about equaling the style, the style 1.5—3.0 mm. long, trifid about one-third its length. Capsule ellipsoid, apically constricted, 3.0-5.5 mm. long, many-seeded, the seeds tightly cochleate, 0.9-1.2 mm. broad, evenly tuber- culate, the dorsal tubercles somewhat longer than broad, the facial tubercles polygonal (rarely substellate) in outline. Fig 5E (Holotype). Jalisco, Michoacan, Mexico, and Morelos, Mexico, usually above 2000 m. It is interesting to note that in none of the original descriptions of taxa refer- able to this species (or series) was any mention made of the polydichotomous nature of the petals. This may be due to the fact that the foliage is so outstanding that the species can usually be determined without resorting to the tedium of dissection. 17. DRYMARIA LONGEPEDUNCULATA $. Watson, in Proc. Ат. Acad. 25:142. 1890. (HOLOTYPE: Pringle 2121; under ledge of the barranca near Guadala- jara, GH!; isotypes at UC, US, etc.) Laxly spreading or rarely erect annuals, the internodes mostly longer than the leaves, villose with spreading jointed hairs or stipitate-glandular. Leaves opposite, 5-14 mm. long, 5-20 mm. broad, villose to glabrous, reniform to deltoid-ovate, apically rounded to acute and apiculate, marginally entire, basally truncate or 1961] DUKE—REVISION OF DRYMARIA 203 d А. D. ; petal, 10; sepal, 10%; stamen, 10X; 1536. С. ae: а athe Se ag inner pi ig “i 10X Areria holo otype); se peed rs 0 E ЖЗ cisa (holo $ aL Бы o nito d 3 MUN UT зоте ОВ УА асуре); habit, У; X ; petal, 5X ; sepal, 5X. [Vor. 48 204 ANNALS OF THE MISSOURI BOTANICAL GARDEN cordate, 3—7-рН-пегуе4; petioles 1-6 mm. long, stipules entire or bifid, the divisions filiform to narrowly lanceolate, 1.0—1.5 mm. long. Inflorescences of lax terminal 3-many-flowered cymes, the peduncles 1-10 cm. long; bracts lanceolate, 1.5-2.5 mm. long, the pedicels 5-25 mm. long, sparingly villose with glandular hairs occasionally admixed. Sepals 5, 4.2-5.5 mm. long, 1.7-2.7 mm. wide, glabrous to villose, narrowly to broadly ovate, the outer acute, villose, and weakly 3-nerved, the inner rounded and glabrous; petals 5, 5.6-7.0 mm. long, once or twice dichotomous, the ultimate segments emarginate, the trunk laterally provided with several ciliate auricles; stamens 5, 4-5 mm. long, the oblong anthers 0.8-1.0 mm long; ovary at anthesis subglobose, 1-2 mm. long, the style 2.0-2.5 mm. long, trifid about one-third its length, exceeded by the anthers. Capsule 3.5-5.0 mm. long, many-seeded, the seeds 0.7-0.9 mm. long, cochleate, evenly tuberculate, the dorsal tubercles conical, the facial tubercles elongate-stellate. Fig. 5F (Holotype). Jalisco, Michoacan, and Mexico, Mexico. Here the tendency toward a second dichotomy in the petals is often suppressed to the point that the petal is merely bilobed, but the lobes are emarginate, and the twice-dichotomous nature is still reflected in the venation. This species affords a good transition to the series vitLosaE. McVaugb & Кое 807 from Jalisco makes a marked approach toward D. multiflora, and may represent a hybrid. Indicative of hybrid tendencies are the broadly reniform leaves, the absence of villosity, and the weakly emarginate petals. The floral parts in general are a bit small for D. longepedunculata and come closer to the range of dimension exhibited by the flowers of D. multiflora. H. Series LEPTOPHYLLA Leaves linear, oblong or narrowly spatulate, subsessile, stipulate, the stipules usually entire. Flowers in bracteate cymes, the pedicels longer or shorter than the bracts. Sepals obtuse to acuminate, trinerved. Petals bilobed, unguiculate, exauriculate, the lobes 1(—3)-nerved. Seeds cochleate, tuberculate, the tubercles not ornate. Two polymorphic intergrading delicate annual (or rarely perennial) species; western U.S. A. to Guatemala. Fig. 6. a. All the sepals obtuse, often cucullate; seeds laxly cochleate, the anterior end not strongly recoiled: b. = usually taller than 5 cm., subvirgate, branching above the base; bracts mostly ualed or exceeded by the dobandi pedicels; lobes of the petals 1—3-nerved. e ксы and pedicels stipitate-glandular; lobes of the petals eral -nerved............. EFFUSA Var. EFFUSA с on = ен glabrous or with sessile glands; lobes = E * ted obovate, FUSA Var. CONFUSA b. Plants салчы и less than 5 cm. tall; branching mostly from the base and in the in жені е; bracts mostly exceeding the subtended pedicels; nee м the petals USA Var. DEPRESSA a. Some or all of the sepals acute to acuminate; seeds tightly ae me anterior end strongly recoiled. d. Peduncles, pedicels and sepals devoid of stipitate-glands, the pedicels mostly shorter than the subtending bracts; petals bifid half their length or чау и. т the sepals........... OPHYLLA Var. LEPTOPHYLLA "чу 1961) DUKE—REVISION OF DRYMARIA 205 d. Peduncles and often the pedicels and sepals stipitate-glandular, the pedicels mostly longer than the subtending bracts; petals bifid for more than half their length, included by or exerted from the calyx: e. Sepals lanceolate, exceeding the petals, not рр; plants са pags D. ar. COGNATA e. Sepals — equaled or exceeded by the petals, usually — -glandular; ios not suffrutesce c. D. LEPTOPHYLLA var. NODOSA 18a. DRYMARIA EFFUSA var. EFFUSA ii effusa A. Gray, Pl. Wright. 2:19. 1853. (ноготуре: Wright 869, mountains of Santa Cruz, Sonora, GH!) Erect, delicate, sparsely branched annuals to as much as 25 cm. high, the inter- nodes mostly longer than the leaves, minutely glandular-puberulent or glabrate, the roots to ca. 1 mm. thick. Basal leaves pseudoverticillate, spatulate to orbicular; cauline leaves opposite, rarely pseudoverticillate, 5-25 mm. long, 0.5-1.2 mm. broad, minutely puberulent or glabrous, linear, apically ‘icone or apiculate, sessile, the stipules entire, acicular, caducous, 0.5—1.0 mm. long. Inflorescences of terminal multifloriferous cymes, the ultimate branches showing tendencies to become monopodial; peduncles 2—5 cm. long, stipitate-glandular; bracts ovate, 0.5—1.8 mm. long, the mature pedicels 1-5 mm. long, stipitate-glandular, usually exceeding the bracts. Sepals 5, 1.4-2.2 mm. long, the outer narrower, shorter, more obtuse and often cucullate, glandular, the inner more ovate, often acutish, all trinerved, the veins subapically confluent; petals 5, 2—4 mm. long, bifid half their length or more, the lobes linear to oblong, 1-nerved, obtuse, tapering gradually to the claw, exap- pendiculate; stamens 5, 1.2—1.6 mm. long, the oblong anthers са. 0.3 mm. long; ovary at anthesis ellipsoid, the style rather elongate, trifid about half its length, about attaining the anthers. Capsule ellipsoid, nearly equaling the sepals, the stigmata exerted; seeds 2-9, cochleate to lacrimiform, coarsely tuberculate, 0.5— 0.8 mm. broad. Fig. 6B (Holotype). Arizona, U.S.A.; Sonora and Sinaloa, Mexico, the Sinaloa specimens approach- ing D. leptophylla. 18b. DryMaria EFFUSA var. confusa Ы; М. Rose) J. Duke comb. & stat. nov. D confusa se, in Contr. U. S. Nat. Herb. 5: Ape 1897. (HOLOTYPE: Palmer 59, nc od ТЗ Баайа, US!, Sarti at GH, MO, etc.) Delicate sparsely branching glabrous or subglabrous annuals to as much as 7.5 cm. high, scarcely branching at the base. Basal leaves glabrous, orbicular to broadly elliptic, 2.5—3.5 mm. long, 2.5-3.0 mm. broad, the petioles 1.5-2.5 mm long; cauline leaves linear to oblong, obtuse, slightly narrowed to the clasping base, 10-20 mm. long, са. 1 mm. broad, the stipules entire, acicular, 0.5-1.0 mm. long. Inflorescences of terminal diffuse several-flowered cymes, the glabrous peduncles 10-25 mm. long; bracts ovate to narrowly lance-deltoid, 0.5—0.8 mm. long, scariose and transparent save for the darkened midrib; pedicels glabrous, 2-8 mm. long, at maturity usually longer than the flowers. Sepals 5, 1.5—2.2 mm. long, 0.5— 7 mm. broad, minutely glandular, broadly oblong, apically obtuse, transparent save for the 3 ribs; petals 5, 2.0-2.5 mm. long, bifid about half their length, the lobes spatulate to narrowly oblong, trinerved, exappendiculate, merging imper- [Vor. 48 206 ANNALS OF THE MISSOURI BOTANICAL GARDEN ceptibly with the narrower claw; stamens (3?—) 5, са. 2 mm. long, the oblong anthers ca. 0.4 mm. long; ovary at anthesis ellipsoid, slightly exceeded by the anthers, the slightly trifid style exceeding the anthers. Capsule ca. 1.8 mm. long, few seeded, the seeds cochleate, minutely tuberculate, ca. 0.6 mm. long. Fig. 6C (Holotype). Chihuahua and Durango, Mexico. Nelson 4813 is determined as this variety only with some misgivings. All the sepals are obtuse and cucullatg and many of the pedicels are exceeded by the sub- tending bracts, in these respects approaching var. depressa. Although suggestive of var. depressa, this plant’s upright habit and obovate 3-nerved petal-lobes deny this possibility. 18c. DRYMARIA EFFUSA var. depressa (Greene) J. Duke comb. $ stat. nov. Aste depressa Greene, Leafl. Bot. Obs. 1:153. 1905. (HOLOTYPE: Metcalf "o ps an the Black Range, alt. 9,500 ft., N. Mex., ND; isotypes at F, GH, UC, US, Drymaria кенен Standl. & Steyerm. іп Field Mus. Bot. 23:52. 1944. (HOLOTYPE: Serene 50243; on rocky limestone outcrops under Juniperus standleyi, alt. 3700 . Vicinity » émal, summit of Sierra de los Cuchumatanes, dept. Huehuetenango, cse ala, F Minute, subcespitose, glabrous to minutely puberulent annuals rarely to 5 cm. high, the branching largely confined to the base and the inflorescence. Rosette leaves orbicular to spatulate, 1.5—10.0 mm. long, 1-3 mm. broad; cauline leaves when present opposite, the blades oblong, 3-10 mm. long, 1—4 mm. broad, apically obtuse, basally tapered, subsessile, the stipules entire, 0.5—1.2 mm. long, acicular, caducous. Inflorescences of terminal lax (rarely condensed) several-flowered cymes, the bracts 1.5-2.0 mm. long, mostly longer than the pedicels, the pedicels and peduncles glabrous or glandular. Sepals 5, 1.5—2.5 mm. long, ca. 1 mm. broad, glabrous or glandular, oblong to ovate, apically obtuse, cucullate, 3-ribbed, the ribs subapically confluent; petals 5, 1.5—2.8 mm. long, bifid about half their length, the lobes linear to narrowly oblong, apically obtuse, usually 1-nerved, gradually tapering to the claw; stamens 5, 0.7-1.7 mm. long, the oblong anthers 0.2-0.3 mm. long; ovary at anthesis globose to obturbinate, the styles 2—3-fid about half their length, about equaling the anthers. Capsule ellipsoid to subglobose, about equaling the sepals, 6—12-seeded, the seed cochleate to lacrimiform, 0.5-0.7 mm. long, dorsally sulcate, minutely tuberculate. Fig 6A. (Isotype). High altitudes, New Mexico, Colorado and Arizona, U.S. A., south to Guate- mala. As a rule this is quite a distinctive variety but it is probable that it hybridizes with other members of the series. Metcalfe 1428 strongly suggests that hybridiza- tion has occurred with D. leptophylla or one of its varieties. Except rarely, D leptophylla and D. effusa var. depressa are quite distinct, the former being char- acterized by an elongate subvirgate habit, the capsules shorter than the inner acute sepals, and the deeply cleft style, the latter being differentiated by the dwarf subcespitose habit, the capsules about equaling the obtuse, cucullate inner sepals, and the shallowly cleft style. 1961] DUKE—REVISION OF DRYMARIA 207 flower, p. 6: A. D. . depressa (isotype); petal, 10%; sepal, 10%; stamen, 10X; қ ре Б de YX. i D. effusa var. effusa (holotype); petal, 10X; sepal, 10X; x. le . n seed, 15X. Е. D. leptophylla var. nodosa ensi o petal, 10 Х ; sepal, 10X ; stamen, 10X; leptophyll . nodosa of D. gentryi); petal, 15 Х; sepal 15X; stamen, Bu mx sa "© sY D. tebe ag tae var. Мені, (isotype of D. gracillima); petal, 10X; sepal, 10; ЈЕ Ра seed, 15 [Vor. 48 208 ANNALS OF THE MISSOURI BOTANICAL GARDEN In correspondence with Dr. Steyermark, I have suggested to him that D. minus- cula probably represents an alpine reduction of what I here reduce to the var. depressa, and he tends to agree with this disposition of his diminutive species. The type from Guatemala has somewhat reduced nodes, leaves, sepals, etc., but such reductions also occur at the other end of the range of the variety, e.g. Colorado, and it seems doubtful that such diminutions are worthy of formal status. 19a. DRYMARIA LEPTOPHYLLA var. LEPTOPHYLLA Drymaria leptopbylia d & Schlecht.) Fenzl ex Rohrb. in Dur 37:195. 1871. v.) : $c ерре 511; ad radices montis Orizabae, B; Arenaria deptophylls Chan. & Schlecht. in Linnaea 5:233. 1830 Drymaria tenella A. Gray, Pl. Fen A 12. 1849. (HOLOTYPE: Fendler 56; eight miles west 25. New Mibo: Dora nodosa var. ? gracillima Неон. Diag. Pl. Nov. 2:22. 1879. (HOLOTYPE: Parry almer 60; in regione San Luis Potosi, alt. 6000-8000 ped., Mexico, K 7. v.; isotypes MO, GH Drymaria plegii (Hema) J. М. Rose, in Contr. О. S. Nat. Herb. 5:132. 1897. Erect delicate, usually sparsely branched annuals to as much as 20 cm. high, the internodes much longer than the leaves, glabrous to minutely glandular, the taproot ca. 1 mm. thick. Leaves opposite, rarely pseudoverticillate, glabrous, linear to narrowly oblong, 5-25 mm. long, 0.5—1.0 mm. broad, often involute and circinate, apically obtuse or acute, tapered to the subsessile base; stipules entire, 0.2-0.8 mm. long, acicular, caducous. Inflorescences of terminal many-flowered dichasial cymes, the peduncles 1—5 cm. long, glabrous to minutely glandular; bracts ovate, 0.5—1.0 (—1.5) mm. long, equaling or exceeding the pedicels. Sepals 5, ovate, 1.5-3.5 mm. long, narrowly ovate, apically acute and often somewhat reflexed (the outer sepals obtuse and cucullate in some northern material), 3-ribbed, glabrous or with a few sessile glands; petals 5, 1.2-2.4 mm. long, bifid about half their length, the lobes linear, obtuse or acute; stamens 5, 1.0-1.5 mm. long, the suborbicular anthers ca. 0.2 mm. long, the filaments basally and briefly connate; ovary at anthesis subglobose, the style about attaining the anthers, divided nearly to the base. Capsule 1.5-2.0 mm. long, 5—20-seeded, the seeds cochleate, 0.5— 0.7 mm. long, evenly tuberculate with domical tubercles. Fig 6G (Isotype of D. gracillima). Arizona to Colorado, U.S. A., south to southern Mexico, and in Baja California. Toward the northern end of its range, the specimens belonging to the typical variety show frequent, but inconsistent, tendencies toward more obtuse, often cucullate, outer sepals and more pronouncedly coiled and contorted leaves. In the northern material the outer sepals are more frequently shorter than the inner, while the converse condition is more frequent toward the south. Although there is not enough material to justify any rigid hypothesis, I would guess that clinal variation is at work in this complex variety. 19b. DRYMARIA LEPTOPHYLLA var. cognata (S. F. Blake) J. Duke comb. & stat. nov. pisei cni S. Е. Blake, in Journ. Wash. Acad. 14:285. 1924. (нототурЕ: Palmer 912, vicinity of Durango, Durango, US!) 1961] DUKE—REVISION ОЕ DRYMARIA 209 Upright, diffusely branching, locally ае puberulent annual to as much as 15 cm. high. Leaves opposite, the blades 6-20 mm. long, 0.5—1.5 mm. broad, glabrous, lance-linear, apically acute, fas aie tapering to the subsessile somewhat clasping base, veinless or obscurely 3-nerved; stipules mostly entire, acicular, 0.5— 0.8 mm. long. Inflorescences of several-flowered, diffuse, trichotomous cymes, the axes locally glandular-puberulent; bracts lance-ovate to ovate, aristate, scariose save or the green excurrent midrib, 1-2 mm. long, usually shorter than the subtended pedicels, the pedicels 0.5—4.0 mm. long. Sepals 5, 3.3-4.5 mm. long, 0.8-1.3 mm. broad, glabrous or very sparsely glandular, lance-ovate, apically attenuate, 3-ribbed, transparent save for the green ribbed area; petals 5, 3.0-3.5 mm. long, bifid slightly more than half their length, the sinus broad, the lobes narrowly oblong, obtuse, the trunk flaring before tapering rather abruptly to the claw; stamens 5, 2.0-2.5 mm. long, the oblong anthers са. 0.5 mm. long; ovary at anthesis sub- globose, the style slightly shorter, trifid about half its length, slightly exceeding the anthers. Capsule ellipsoid, 2-3 mm. long, са. 5-seeded, the seeds cochleate, са. 0.8 mm. broad, minutely and regularly tuberculate. Fig. 6D (Holotype). Durango and Chihuahua, Mexico. 19c. DRYMARIA LEPTOPHYLLA var. nodosa (Engelm.) J. Duke comb. nov. Drymaria nodosa Engelm. in A. Gray, Pl. Fendl. 12. 1849. (HOLOTYPE: reg grown ari 1944. Drymaria gentryi Fo sberg, in Proc. Biol. Soc. Wash. 62:147. 1949. (HOLOTYPE: Gentry + 60; ay ition pine slope, los Cascarones, Rio Mayo; Chihuahua; US!; isotypes at MO, , et Erect delicate, sparsely branched annuals (or perennials?) to as much as 25 cm. high, the internodes mostly longer than the leaves, usually glandular-puberulent, the taproot to 2 mm. in diameter. Basal leaves spatulate to orbicular, opposite or pseudoverticillate, caducous; cauline leaves opposite, glabrous or minutely glandu- lar, 5-30 mm. long, 0.5-2.5 mm. broad, 1-nerved, linear to narrowly elliptic, apically obtuse to apiculate, sessile; stipules entire, acicular, caducous, 0.5-1.5 mm long. Inflorescences of terminal many-flowered dichasial cymes, the peduncles 2-5 cm. long, stipitate-glandular; bracts lance-deltoid to ovate, acuminate, 0.7-2.5 mm. long, mostly shorter than the pedicels, the pedicels stipitate-glandular, 0.5— 5.0 mm. long. Sepals 5, subequal, 2.3-3.5 mm. long, 0.5-1.5 mm. broad, lanceolate to ovate, acuminate, basally or entirely stipitate-glandular, 3-ribbed, the ribs distally confluent; petals 5, 2.0-3.5 mm. long, bifid slightly more than half their length, the lobes linear to oblong or narrowly spatulate, obtuse, mostly 1-nerved, exappendiculate; stamens 5, 1.5-2.5 mm. long, the oblong anthers ca. 0.4 mm. в the filaments basally connate, often with inconspicuous staminodial flaps; ovary at anthesis ellipsoid to subglobose, the styles elongate, nearly as long as the ovary, bifid or trifid less than half their length, equaling or exceeding the anthers. Capsule ellipsoid, 1.5-2.0 mm. long, shorter than the sepals, 1-9- seeded, the seeds cochleate, the anterior end strongly recurved, minutely and regularly reticulate, 0.6-1.0 mm. broad. Fig. 6E (Holotype of D. nodosa), Fig. 6F (Holotype of D. gentryi). [Vor. 48 210 ANNALS OF THE MISSOURI BOTANICAL GARDEN Chihuahua, Sonora, Durango, San Luis Potosi and Hidalgo, Mexico. Gentry 2669, type of D. gentryi, stands out with its rather broad leaves, but examination of other specimens e.g. LeSueur 623, also from Chihuahua, shows that Fosberg’s species cannot stand as distinct from D. leptophylla var. nodosa. Equally aberrant are S. S. White 4185 and 4776 from Sonora. In these speci- mens, the inflorescence is strikingly congested and the petal-lobes are broader than those of most specimens of the variety nodosa. In spite of the bizarre inflorescence, it seems best to retain them within the variety, with which they coincide in other fundamental characteristics. I. Series TENUES Leaves opposite, lanceolate, petiolate, stipulate, the stipules mostly lacerate. Flowers in monopodial or dichasial cymes, the pedicels longer or shorter than the bracts. Sepals acute, 1—3-nerved. Petals bifid, the lobes 1—5-nerved, unguiculate, exauriculate. Stamens 2—5. Seeds cochleate, tuberculate, the tubercles not ornate. Two diffusely spreading annual or perennial Mexican species. Fig. 7 a. ас 3- ate ds ri res cnim ultimate branches of the cymes sitter petioles D. ANOMALA а. Srl ілмей je or with sessile glands; cymes predominantly dichasial; -— 1-6 D. TENUIS 20. DRYMARIA ANOMALA 5. Watson, in Proc. Am. Acad. 25:143. 1889. (ного- TYPE: Pringle 2847, Carneros Pass, Coahuila, US!; isotype at GH) Upright or spreading annuals or perennials, much branched from below, the internodes mostly longer than the leaves, stipitate-glandular, the taproot to 5 mm. thick. Cauline leaves opposite, 3—8 mm. long, 2-4 mm broad, glabrous to glandu- lar or puberulent, the blades lance-elliptic, obscurely veined, apically acute, basally cuneate to acute, the petioles mostly less than 1 mm. long; stipules entire or bifid, lance-deltoid to acicular, persistent, 0.5—1.5 mm. long. Inflorescences of terminal many-flowered cymes, the first branches dichasial, subsequent branches becoming monopodial; peduncles 2-5 cm. long, glabrous to stipitate-glandular; bracts 0.5— 2.0 mm. long, exceeding the pedicels, the pedicels stipitate-glandular or glabrescent. Sepals 5, 2-3 mm. long, 1.0-1.5 mm. broad, stipitate-glandular, broadly elliptic, apically acute, with 3 prominent veins, the outer submarginal, there being little or no scariose margin; petals 5, 1.5-2.0 mm. long, bifid a little over half their length, the lobes oblong, obtuse; stamens (2-) 3-5, 1.0-1.4 mm. long, the anthers oblong, ca. 0.3 mm. long; ovary at anthesis obturbinate, the style trifid about half its length, much shorter than the ovary, about attaining the anthers. Capsule about equaling the sepals, 5—9-seeded, the seeds cochleate, closely tuberculate, 0.6-0.9 mm. broad. Fig. 7A (details from Holotype). Coahuila, Zacatecas, San Luis Potosi and Hidalgo, Mexico. 21. DryMaria TENUIS S. Watson, in Proc. Am. Acad. 25:142. 1889. (HoLo- ТУРЕ: Pringle 2120; under ledges of the barranca near Guadalajara, Jalisco, US!; isotype at GH) apasand d 1961] DUKE—REVISION OF DRYMARIA 211 Bier. „ара Robinson & Seaton, in Proc. Am. Acad. 28:117. 1893. not Benth. LOTYPE: Seaton prs Виа slopes, Ме. Orizaba, 9000 ft. GH!; isotypes at is ad ay etc. Drymaria tenuis a var. genuina Briq. in Ann. Cons. & Jard. Bot. Gen. 13:374. 1911. Drymaria tenuis В var. jaliscana Briq. loc. си. 374. 1911. (HOLOTYPE: Pringle 4536; Civitas Jalisco, in declivibus altis prope Guadalajara, С; м. v.; isotypes at MO, US etel Erect or clambering delicate annuals or perennials to as much as 50 cm. long, the internodes much longer than the leaves, glabrous or pilosulous, the root some- times lignescent, to as much as 6 mm. thick. Leaves opposite, glabrous, glandular or pilosulous, the blades lance-elliptic to ovate, apically and basally acute, 2.5—10.0 mm. long, 1-7 mm. broad, the petioles 0.5—5.0 mm. long; stipules lacerate, the divisions acicular, 0.5—2.0 mm. long. Inflorescences of terminal dichasial cymes, the peduncles very elongate, glabrous; bracts ovate, scariose, 1-2 mm. long, the Fi A. D. anomala n from holotype); inflorescence, 1 Х; flower, 10Х; petal, 10X ; EN ES seed, 15 Х. В. D. tenuis (holotype of D. filiformis); petal, 10 Х ; sepal, 10X; stamen, ins seed, 22% Хх. С, D. 4 feuda (holotype); inflorescence, 4X; flower and бирте ы 15X; petal, 20X. [Уот.. 48 212 ANNALS OF THE MISSOURI BOTANICAL GARDEN pedicels 2-7 mm. long, glabrous, usually much longer than the subtending bracts. Sepals 5, the outer ovate, apically obtuse, 2.0-2.5 mm. long, са. 1 mm. broad, the one rib occasionally excurrent; inner sepals narrowly ovate to ovate, 2.5—3.0 mm. long; petals 5, 1-2 mm. long, bifid about two-thirds their length, the lobes oblong, apically obtuse or acutish, exappendiculate, gradually tapering to the linear claw; stamens 5, 1.0—1.3 mm. long, the oblong anthers са. 0.2 mm. long, the filaments basally connate; ovary at anthesis subglobose, ca. 0.8 mm. long; style bifid or trifid about half its length, about attaining the anthers. Capsule ovoid, about equaling the sepals, 5—15-seeded, the seeds cochleate, 0.5-0.7 mm. broad, evenly tuberculate, the dorsal tubercles mostly longer than broad. Fig. 7B (Holotype of D. filiformis Robinson & Seaton) ; Fig. 7C (Holotype of D. tenuis). Michoacan, Jalisco, Guanajuato, Zacatecas and Veracruz, Mexico. The type of D. filiformis from Mt. Orizaba differs only in having shorter petioles and smaller, more obtuse petal lobes, differences possibly resulting from the alpine environment and probably of no taxonomic consequence. Unfortunately Briquet relied too heavily on Watson’s description of D. tenuis as glabrous, and published the variety jaliscana, believing it to differ in being pubescent. Examina- tion of the holotype of D. tenuis reveals that Watson’s description was erroneous; the type is locally villosulous with hairs up to 0.5 mm. long. Briquet's pubescent variety, collected very near the type locality of the typical variety, is probably no more pubescent than the typical variety. J. Series FRUTESCENTES Leaves linear to lanceolate, opposite, occasionally pergameneous, sessile or briefly petiolate, stipulate or estipulate. Flowers, showing tendencies toward monoecism, in dichasial bracteate cymes. Sepals acute, 3—5-ribbed. Petals bifid, the lobes 3- 9-nerved, exauriculate, unguiculate or exunguiculate, the trunk entire or denticu- te. Sepals cochleate, tuberculate, the tubercles acute or obtuse. Four more or less erect suffrutescent perennial species of Peru and Ecuador. Fig. 8. a. Leaves neither imbricate nor — pedicels and sepals often glandular, the sepals 3-4- nerved; petals not tapered to the claw Ь. Pedicels and sepals usually Шей: the midrib of the Lia not et seeds with domical or conical tubercles; stipules present or absent. Peru and Ecu c. Leaves край) trunk of the petals conspicuously constricted at thi c commissure with the lobes, claw shorter than the trunk; seeds with acute — => Ecuador. ‚ STELLARIOIDES c. Leaves often estipulate; trunk of the petals more or less continuous wi = gi lobes, хы claw occasionally longer than the trunk; seeds with blunt, pow иис» eru b. Pedicels aec sepals glabrous, = midrib of the sepals excurrent; а aie "e ical tuber ir pules present. ru Р. алон . Leaves құқ pungently scutes pedicels and sepals glabrous, the sepals — 5-nerved; Satie а to the claw. D. FRUTESCENS TEREOPHYLLA 22. DRYMARIA STELLARIOIDES Humb. & Bonpl. ex Roem. & Schult. Syst. Veg. 5: 406. 1819. (HOLOTYPE: Humb. & Bonpl. s. n.; crescit prope Hambato, alt. 1380 hex. [Regno Quitensi]; probably destroyed; fragment and photograph at F! illustr. HBK. Nov. Gen. & Sp. 6; pl. 516. 1823.) 1961] DUKE— REVISION OF DRYMARIA 213 У i f plant, 12X Fig D. stellarioides; petal, 3 Х; sepal, 3X; seed, 15X; upper Jets о В. noe Seal var. срба e (isotype) ; pe bie i cans 5 vi š iei гр sereop bil і І > 4 5 see 2 н ур stipulata pp aon ru P amen, ЗХ se eed, 74x. E. D. frutescens y оер habit, 4X; реш, AX: ы. rA stamen, 4% Х [Уот. 48 214 ANNALS OF THE MISSOURI BOTANICAL GARDEN Upright or ascending dichotomous lignescent perennials from a stout tap- root to 15 mm. thick, the internodes mostly longer than the leaves except on condensed lateral branches, glabrous to glandular-pubescent. Leaves opposite, glabrous or rarely glandular-puberulent, lanceolate to oblanceolate, apically acute, marginally entire, 3-ribbed, 5-20 mm. long, 2.5-7.5 mm. broad, tapered to the sessile, clasping base; stipules entire, apparently fused or occasionally absent, 0.0- 5 mm. long. Inflorescences of lax to crowded 5—many-flowered cymes, the peduncles 1-8 cm. long; bracts lanceolate, 3-5 mm. long, intergrading with the foliage leaves; pedicels glabrous to densely glandular-puberulent, all but the central pedicel usually exceeded by the bracts. Sepals 5, narrowly to broadly lanceolate, glabrous to densely glandular-puberulent, strongly 3-ribbed, apically acute, (4—) 5-9 mm. long, 2-3 mm. broad; petals 5, 5-10 mm. long, bifid about half their length, the lobes oblong, obtuse, 4—8-nerved, exauriculate, constricted slightly toward the trunk, the trunk rhombic to rectangular, truncate to the short claw; stamens 5, 3-6 mm. long, those of fertile flowers (destined to mature seeds) usually much shorter than those of infertile flowers; anthers 0.7-1.1 mm. long; filaments basally connate into a cup ca. 1 mm. deep; ovary (occasionally absent in flowers with long stamens) ellipsoid, multiovulate, those of fertile flowers about equaling the anthers; styles 1.5—3.0 mm. long, trifid for about one-fourth their length. Capsule ellipsoid, substylopodiate, 3-5 mm. long, 2—many-seeded, the seeds laxly circinnate, 0.8—1.2 mm. broad, tuberculate, the dorsal tubercles subspinulose, the facial tubercles tending to be polygonal. Fig. 8A. Ecuador. Pachano 95 (Fig. 8A) and the isotype seem to differ from other representatives of this species in that the sepals and petals are unusually long and the petal lobes have more numerous veins. Seeds of this species, as illustrated in HBK. Nov. Gen. & Sp. 6: pl. 516. 1823, appear to be constantly subspinulose-tuberculate. There is no indication in the aforementioned illustration however of the polygamous tendency which I believe characterizes this species. Although the material available for dissection has not been too copious, I feel that certain correlates exist in differ- ent types of flowers. Many of the ovaries bear aberrant ovules and in some flowers the ovary is completely lacking. In flowers which mature seed, the anthers rarely surpass the body of the capsule, while in those flowers with aberrant ovules, the filaments seem to have elongated so that the anthers equal or surpass the style branches. In some unnumbered specimens collected by Jameson near Cuenca and Loxa, there are flowers with long glandular sepals and others with shorter glabrous sepals; those with the shorter glabrous sepals proved to be fertile while some of those with longer glandular sepals were completely devoid of ovaries. 23. DRYMARIA STEREOPHYLLA Mattf. in Notizbl. Bot. Gart. Berl. 13:436. 1936. (HOLOTYPE: Raimondi 2502; Peru. Dep. Junin: Prov. Jauja; Abhang am Fusse _ des Chacapalpa zwischen Huari und Jauja, В, и. v.; probably destroyed) Drymaria stereophylla Mattf. ex MacBride, in Field Mus. Bot. 132:626. 1937. (HOLOTYPE: MacBride 962; Junin; in crevices of limestone cliffs, 4000 m., La Oroya, Peru; F!; iso- 1961] DUKE—REVISION OF DRYMARIA 215 че а var. sr eit Mattf. in Notizbl. Bot. Gart, Berl. 13:437. 1936. (HO W auer 6600; Peru; do ,Junin; nordóstlich von и unter- halb. der 1 Hiciende Acopales, geogr. Br. 5' S., an Felsen, 3600-3700 m ü. M.; B, 7. v.; isotypes at Е, MO, US!) Laxly spreading or ascending lignescent perennials to as much as 35 cm. tall, the internodes shorter to longer than the leaves, glandular-puberulent, the taproot to as much as 1 cm. thick. Leaves opposite, 4-12 mm. long, 2-6 mm. broad, glandular-puberulent, broadly lanceolate to narrowly ovate, apically acute to aristately acuminate, subsessile, the brief petioles clasping; stipules entire, lance- linear, caducous, 0.5-1.0 mm. long, absent іп the variety. Inflorescences of lax occasionally congested terminal 3—many-flowered cymes, the peduncles 1-6 cm. long; bracts lanceolate to lance-ovate, 3-5 mm. long, the pedicels 2-10 mm. long, stipitate-glandular. Sepals 5, 5.0-7.5 mm. long, 2.0-4.5 mm. broad, glandular, ovate, 3—4-nerved, the midrib not excurrent, the outer acute, the inner obtuse; petals 5, 5-9 mm. long, deeply bifid, the lobes oblong, obtuse, 5-8-nerved, exap- pendiculate, not constricted at the junction with the trunk, the trunk truncate to the claw; the claw minute, or, in the variety, as long as the trunk; stamens 5, 4-6 mm. long, the oblong anthers са. 1 mm. long, the filaments basally connate forming a cup about 1 mm. deep; ovary at anthesis turbinate to ellipsoid, the style 1.0-2.5 mm. long, bifid or trifid about one-third its length. Capsule ovoid, 2.5-3.5 mm. long, 4—27-seeded, the seeds loosely circinnate, 0.9—1.3 mm. broad, tuberculate, the tubercles low and contiguous. Fig. 8B (Isotype of var. stereophylla); Fig. 8C (Isotype of var. exstipulata). Departments Junin, Cuzco and Lima, Peru. Mattfield commented on the homogeneity of the specimens which he cited in the original description of D. stereophylla var. exstipulata. Material collected sub- sequently has further substantiated the differences in the varieties and other differ- ences have been uncovered. The shape of the petals, frequent telescoping of the internodes, and the absence of stipules in the var. exstipulata are characteristics which point toward the bizarre D. frutescens. The two varieties of D. stereophylla may be keyed as follows: a. Stipules к the иңе seid, glabrous or with a few sessile glands; trunk of the petals much longer than the claw, the claw occasionally not even apparent................... var. STEREOPHYLLA a. Stipules 2. the өбден: not rigid, ам» often densely stipitate-glandular; авг of the petals about as long as the trunk, EXSTIPULATA 24. DRYMARIA AURICULIPETALA Mattf. in Notizbl Bot. Gard. Berl 13:439. 1936. (HOLOTYPE: MacBride & Featherstone 2264; Peru; Llata, von Fels- bándern ћегађћапрепа, ca. 2100 m ü. M.; F!; isotypes ас GH, MO, US) Upright or ascending glabrous suffruticose annuals or perennials to as much as 60 cm. long. Leaves opposite, glabrous, 5-15 mm. long, 1.5—3.0 mm. broad, the blades narrowly lanceolate, slightly falcate, apically attenuate, aristate, basally clasping, the stipules entire, acicular, сага у deciduous, 1.0-1.5 mm. long. res- cences of lax terminal 1—7-flowered cymes, the glabrous peduncles 5-40 mm. long; bracts broadly ovate, strongly 1-ribbed, 2.5—3.5 mm. long, exceeding the pedicels. Sepals 5, subequal, 7-8 mm. long, 2-3 mm. broad, glabrous or minutely glandular, [Vor. 48 216 ANNALS OF THE MISSOURI BOTANICAL GARDEN lanceolate, apically attenuate and aristate, basally unguiculate or umbonulate, only the midrib conspicuous; petals 5, 6.5—7.5 mm. long, bifid about half their length, the lobes broadly oblong, apically rounded, 1.0—1.3 mm. broad, 4—6-nerved, ex- apendiculate, hardly constricted at the junction with the trunk, the trunk 1.2-1.6 mm. broad, the claw minute; stamens 5, ca. 5.5 mm. long, the oblong anthers ca. 0.8 mm. long, often somewhat twisted; ovary at anthesis obovoid, slightly exceeded by the anthers, the style trifid about half its length, about equaling the anthers. Capsule ca. 4 mm. long, many-seeded, the seeds cochleate, ca. 1 mm. broad, dorsally flattened or sulcate, tuberculate, the tubercles low and contiguous. Fig. 8D (Holotype). Known only from the type locality; Llata, Peru. 25. DRYMARIA FRUTESCENS Mattf. іп Notizbl. Bot. Gart. Berl. 13:439. 1936. (ноготүре: Weberbauer 7203; Peru; Dep. Libertad, Prov. Santiago de Chuco, über der Hacienda Angasmarca; Grasssteppe mit zahlreichen eingestreuten Sträuchern, an felsigen Abhingen, 3650 m й. M.; В, ә. v.; isotypes at Е, US!) Ascending frutescent glabrous perennials to 30 cm. tall, the leaves closely appressed to the stems. Leaves 2—6 mm. long, 1.0—1.5 mm. broad, imbricate in two ranks, lance-ovate, apically aristate-acuminate, basally clasping, 3-ribbed, margin- ally transparent; stipules entire, narrowly lance-deltoid, ca. 0.5 mm. long. Flowers solitary at the ends of the branches, the bracts similar to the foliage leaves, the glabrous pedicels 2-3 mm. long. Sepals 5, subequal, 5.5-7.0 mm. long, 1.5-2.0 mm. broad, glabrous concavo-convex, ovate, apically acute or shortly acuminate, transparent save for the 7 nerves; petals 5, 5-7 mm. long, bifid for about two- thirds their length, the lobes oblong to spatulate, obtuse or emarginate, exappen- diculate, the trunk and claw merging almost imperceptibly; stamens 5, 2.5—4.5 mm. long, the oblong anthers ca. 0.8 mm. long, often spirally contorted in drying; ovary at anthesis ellipsoid, exceeded by the anthers, the style trifid about half its length, about equaling the anthers. Capsule 2.5—3.0 mm. long; mature seeds not seen. Fig. 8E (Isotype). Known only from the type locality in Dept. Libertad, Peru. K. Series FASCICULATAE Leaves opposite, linear-elliptic to narrowly ovate, subsessile, stipulate, the stipules entire or bifid. Flowers in rather dense bracteate cymes, the bracts often stipulate. Sepals acute, 1–3-пегуед. Petals bifid, the lobes 1—3-nerved, exauricu- late, the claw minute or completely absent (except in D. praecox), the trunk as wide as the combined width of the lobes. Stamens 3—5. Seeds loosely or tightly cochleate, tuberculate. Three species in Peru, two rather delicate and minute annuals and the more or less suffrutescent D. fasciculata. Fig. 9. a. Plants upright, often suffrutescent; "e 224 mm. long; leaves mostly larger er 6m long and 4 mm. broad; bracts not stipul | PASCICULATA a. MN дерге емей ог acd cue = 2.0-4.5 mm. long, glabrous, пије pe villosulous; leaves mostly less than 6 mm. long and 4 mm. broad; bracts often етй 1961] DUKE—REVISION OF DRYMARIA 217 b. зере, acuminate, glabrous, с пина reflexed in age; herbage occasionally glandular but not villose. Dept. Lima, Huancavelica and Puno: Peru............. 7b. D. ENGLERIANA Var. DEVIA b. Sepals — — dde, — Ó apically inflexed; herbage villose or es lous. Dept. Ancash and Cuzco; P . Petals ih a ужа d trunk, not tap 2. % а claw, ке — 1-3-пегуеЯ; pedicels of the central flowers about as long as e bracts. Dep a ENGLERIANA var. ENGLERIANA с. phe gradually tapered to a definite claw, the ides 1-күн) pedicels of the tral flowers much longer than their bracts. Dept. C 28. D. PRAECOX 26. DRYMARIA FASCICULATA A. Gray, Bot. 1). S. Expl. Exped. 125. 1854. (ного- TYPE: Wilkes exped. s. n.; Andes of Peru, near Obrajillo, US!) Upright dichotomous, paucifoliate annuals to as much as 25 cm. high, sub- glabrous except for the glandular axes of the inflorescence, the taproot to as much as 3 mm. broad. Leaves opposite, the blades 8-18 mm. long, 4-10 mm. broad, ovate (the lowermost occasionally spatulate), apically acute or acuminate, margin- ally somewhat undulate, basally clasping, only the midrib obvious, the stipules entire, linear-lanceolate, caducous, ca. 1 mm. long. Inflorescences of termina fasciculate, strongly dichotomous, 3-many-flowered cymes, the glabrous to densely glandular peduncles 2-6 cm. long, the bracts lanceolate to ovate, acute ог mucronate, transparent save for the midrib, 4-6 mm. long, 2-4 mm. broad, obscuring the short pedicels. Sepals 5, subequal, 4-7 mm. long, 1.0-1.8 mm. broad, lance-oblong to narrowly elliptic, acuminate, rather strongly 3-ribbed, often with а scariose margin; petals 5, 2.5-7.0 mm. long, bifid about half their length, the lobes oblong, apically rounded to emarginate, 2—3-nerved, their outer margins forming straight lines with the margins of the trunk, exappendiculate, the trunk exunguiculate or subexunguiculate; stamens 5, 2-6 mm. long, the oblong anthers 0.4-1.2 mm. long; ovary at anthesis subcylindric, slightly exceeded by the anthers, the barely trifid style slightly exceeding the anthers. Capsule narrowly ellipsoid, included, 3—4 mm. long, several-seeded, the seeds laxly circinnate, minutely and rather evenly tuberculate, the anterior end usually long and pointed. Fig. 9A (details from Holotype). | Andes of Peru. 27a. DRYMARIA ENGLERIANA Var. ENGLERIANA crei englerianum Muschler, in Bot. Jahrb. 45:452. 1911. о ҮРЕ: отет r 3101; Peru; Ancash; Huaraz, 4300 m.; probably destroyed ; С!) : роуа engleriana Ра Baehni & MacBride, in Field Mus. Вог. 137:621. 1937. -Minute prostrate annuals to 5 cm. high, the internodes mostly longer than the leaves, hirsutulous. Leaves opposite, the blades 2-8 mm. long, 1.0-4.5 mm. broad, ovate, apically acute, marginally entire, basally attenuate to the alate clasping petioles, weakly 3-nerved, scantily hirsutulous, the stipules entire or bifid, the divisions filiform to lance-deltoid. Inflorescences of terminal, rather lax, several- flowered cymes, the axes hirsutulous, the bracts 1.5—3.0 mm. mostly longer than the pedicels. Sepals 5, 2.0-3.6 mm. long, 0.6-1.2 mm. broad, narrowly ovate, apically acute or briefly acuminate, often with sessile or stipitate glands, 3- ribbed, only the midrib prominent, the apex not reflexed, often inflexed; petals 5, 1.0-2.4 mm. long, (Vor. 48 218 ANNALS OF THE MISSOURI BOTANICAL GARDEN M. „А. ТУ. асы де» from holotype); petal, 4X ; sepal, 4X ; stamen, 7% Х; seed, 124X: eye NX X iana var. devia (holotype); pen, 10X; sepal, 10X; seed, 20X- eriana var. ыы pes pe); poe, 10X; sepal, 10X. D. D. praecox (holotype); da 1x; petal, 716 X ; sepal, 72 X ; seed aX. 1961] DUKE—REVISION OF DRYMARIA 219 bifid about half their length, the lobes obtuse to acute, oblong, 1-nerved, the outer margin forming more or less a straight line with the margin of the trunk, the claw not apparent; stamens 3-5, 1.0-1.5 mm. long, the suborbicular to oblong anthers ca. 0.2 mm. long, the filaments basally connate into a minute carnose column; ovary renoid, the style shorter than the ovary, ca. 0.5 mm, long, trifid about half its length. Capsule globose to ellipsoid, included, 1.5-2.0 mm. long, 2-8-seeded, the seeds laxly сігсіппате, minutely and evenly tuberculate, 0.7-0.9 mm. broad. Fig. 9C (Isotype). High altitudes, Dept. Ancash, Peru. Although Baehni and MacBride (in Field Mus. Bot. 137:621. 1937) correctly transferred this species to Drymaria from Polycarpon, MacBride (р. 627) also retains it as a species of Polycarpon, curiously stating that it is "probably only a densely cespitose form of P. apurense.’ 27b. DRYMARIA ENGLERIANA var. devia (Baehni & MacBride) J. Duke comb. & stat. nov. Drymaria devia Baehni & MacBride, in Field Mus. Bot. 13?:620. 1937. (HOLOTYPE: Pen- nell 14655; Lima; open rocky slopes near Canta, 4000 m., F!) Delicate prostrate or ascending subglabrous annuals usually less than 5 cm. tall, branching mostly from the base. Leaves opposite, the blades 2.5-5.0 mm. long, 2-3 mm. broad, subcoriaceous, ovate, occasionally glandular, apically acute or attenuate and apiculate, basally clasping, the uppermost rather strongly 3-ribbed; stipules entire or bifid, the divisions narrowly lance-deltoid, ca. 1 mm. long. Inflorescences of dense terminal 1—11-flowered cymes, the glabrous peduncles 4-8 mm. long, the bracts 2-3 mm. long, ca. 1 mm. broad, ovate, aristate-acuminate, the midrib often cyanic; pedicels mostly less than 1 mm. long, concealed by the bracts. Sepals 5, 4.0-4.5 mm. long, 1.5-2.0 mm. broad, narrowly ovate, aristate-acuminate, strongly 3-ribbed, glabrous to minutely glandular, transparent save for the ribs; petals 5, 2.5-3.0 mm. long, bifid about half their length, the lobes oblong, obtuse, 1-nerved, curving outwardly to the inflated trunk, the trunk ca. 1.2 mm. broad, exappen- diculate, the claw not apparent; stamens 5, ca. 2.2 mm. long, the oblong anthers ca. 0.4 mm. long; ovary at anthesis slightly exceeding the anthers, the style trifid about half its length. Capsule 2.5-3.0 mm. long, са. 8-seeded, the seeds dorsally flattened, minutely tuberculate, ca. 0.6 mm. broad, dark brown. Fig. 9B (Holotype). At high altitudes in Depts. Lima, Puno and Huancavelica, Peru. This diminutive variety is closely related to the preceding and seems sufficiently distinct to warrant varietal recognition. The acuminate often reflexed, strongly ribbed sepals and the lack of jointed hairs seem to characterize the few specimens of the variety examined. 28. DRYMARIA PRAECOX Baehni & MacBride, in Field Mus. Вог. 13?:625. 1937. (ноготүре: Weberbauer 6016; between Pisac & Paucartambo, prov. Paucar- tambo, Dept. Cuzco, 4100 m., Peru, F!; isotypes at MO, US, etc.) Delicate, virgate, glandular-puberulent annuals to as much as 5 cm. high, with few nodes below the inflorescence, the basal rosette often persistent. Basal leaves [Vor. 48 220 ANNALS OF THE MISSOURI BOTANICAL GARDEN spatulate; cauline leaves opposite, the blades 2-4 mm. ling, 1.5—3.0 mm. broad; villosulous, ovate, apically acute, basally clasping, obscurely trinerved, the stipules entire, acicular, 0.5—0.8 mm. long. Inflorescences of diffuse 1—9-flowered cymes, the axes densely glandular-puberulent; bracts ovate to lanceolate, stipitate, 1-ribbed, the rib often excurrent, 1.0-1.5 mm. long, the pedicels usually much longer. Sepals 5, 2.5-3.5 mm. long, 1.0-1.4 mm. broad, glandular-pubescent, narrowly ovate, apically acute or obtuse, often inflexed, 3-ribbed, transparent save for the ribs; petals 5, ca. 3 mm. long, bifid half their length or more, the lobes oblong, obtuse, exappendiculate, 3-nerved, gradually tapering to the base, the trunk and claw not discernibly differentiated; stamens 5, 1.2-1.5 mm. long, the suborbicular anthers ca. 0.3 mm. long; ovary at anthesis compressed-globose, exceeded by the anthers, about as long as the style; style trifid about half its length, slightly exceed- ing the anthers. Capsule included, ca. 2.2 mm. long, ca. 5-seeded, the seeds cochleate, ca. 0.7 mm. broad, tuberculate, the tubercles rather prominent, nearly contiguous. Fig. 9D (Holotype). Known only from the type collection from Cuzco, Peru. Perhaps a misfit in this series, this species may be of hybrid origin, and in some respects it recalls D. glandulosa var. galeottiana, perhaps also of hybrid origin. L. Series DEBILES 29. DRYMARIA DEBILIS Brandegee, in Proc. Cal. Acad. 27:131. 1889. (HOLOTYPE: Brandegee s. n.; Purisima, Feb. 13, 1889; UC!) Воч polystachya Brandegee, in Zoe 2:70. 1891. ee quae 35; cliffs near San José del Cabo, Baja California; UC!; isotypes at D, US, kieres diffusa J. N. Rose, in Contr. U. S. Nat. Herb. 1:130. "pl. ia. 1892. (HOLOTYPE: Palmer $19; Carmen Island; US!; isotypes at F, MO, US, NY, etc.) Раны polystachya var. diffusa (Rose) Wiggins, in Proc. Cal. head 4?5:198. 1944. Ascending or clambering, occasionally subscandent annuals or perennials, the tenuous, villose or glandular internodes shorter to longer than the leaves, the tap- root to as much as 15 mm. thick. Leaves opposite, the blades 3-20 mm. long, 3—20 mm. broad, villose or glandular, broadly ovate to reniform, apically acutish and apiculate, basally cordate, often drying with a yellowish tinge, the petioles nearly as long as the blades, the stipules mostly entire, filiform, 1-3 mm. long. Inflores- cences of terminal lax 3-many-flowered cymes, the axes villose or glandular puberulent, the peduncles 1-10 mm. long; bracts narrowly to broadly ovate, scariose, 1.52.5 mm. long, much exceeded by the pedicels, the villose or glandular pedicels to as much as 25 mm. long. Sepals 5, 2.5—5.5 mm. long, 1.5—2.4 mm. broad, glandular to villose, oblong to elliptic or ovate, apically obtuse, emarginate or with the midrib occasionally excurrent, the venation obscurely dichotomous and reticulate; petals 5, 3.0—6.5 mm. long, bifid about half their length, the lobes oblong, rounded to truncate at the apex, 1—2-nerved, gradually tapered to the claw; stamens 5, 1.0—3.5 mm. long, the filaments usually dimorphic, two or three being distinctly longer than the others, the anthers oblong 0.5—1.0 mm. long; ovary at anthesis ellipsoid, often stipitate, exceeded by the anthers, the style 1-2 mm. long, trifid to half its length. Capsule ellipsoid to ovoid, 1.0-3.5 mm. long, 6– 1961) DUKE—REVISION OF DRYMARIA 221 Fig. 10: А. D. debilis (holotype of D. polystach habit, ИЖ. В. D. debilis (holotype); petal, 5X; D. chibuabuensis) ; habit, 2X 5X ; sepals, 5X ; seed, 20X. ya); petal, 7%Х; бергі, 7УХ; seed, 20X; debilis (holotype of D. diffusa); petal, 6X; sepal, 6X; seed, 20X. - sepal, 5X; stamen, 5X; seed, 10X. D. D. laxiflora (isotype of ; petal, 6X ; sepal, 6X ; seed, 20X. Е. D. laxiflora (isotype); petal, (Мог. 48 222 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN 15-seeded, the seeds ampulliform, 0.6-1.0 mm. broad, the pointed anterior ends not strongly recoiled, the tubercles low, polygonal, rather remote. Fig. 10A (Holotype of D. polystachya); Fig. 108 (Holotype of D. diffusa); Fig. 10C (Holotype of D. debilis). Baja California, Mexico. This polymorphic species has generally been regarded as three separate species, the more or less sympatric differences purported to reside in the duration, pubes- cence, and the length and shape of the sepals and petals. I am more impressed by the homogeneity of certain other characters (e.g. the peculiar retort-shaped seeds, the inequality of the stamens, the venation of the sepals) than by the differences supposed to separate the three species. Particularly suspicious to me are keys in which the primary dichotomy is annual vs. perennial Many Drymaria species, originally described as annual, have turned out to have perennial representatives. I prefer to think of D. debilis as a species embracing many contrasting character- istics, but with few, if any, correlated contrasts. M. Series LAXIFLORES 30. DRYMARIA LAXIFLORA Benth. Pl. Hartw. 73. 1839. (НОТОТУРЕ: Hartweg 523; Guatemala; in rupibus Sunilo, BM? x. v.; isotypes at GL, Е, LE, P, etc!) они Vb ur Briq. in Ann. Cons. & Jard. Bot. Gen. 13: 370. 1911. (Horo- ngle 331; Mexicum: Civitas Chihuahua i in rupibus umbrosis prope Chihua- тыд ср п. V.; isotypes ас Е, GH, MO, NY, Р, US, etc.!) Glabrous to densely stipitate-glandular rr to subcespitose perennials to as much as 30 cm. high, the internodes mostly longer than the leaf-blades, the taproot to as much as 6 mm. thick. Leaves opposite, the blades glabrous to densely stipitate-glandular, ovate to reniform, 4-12 mm. long, 3-14 mm. broad, trinerved, apically obtuse to acute and mucronulate, marginally entire, basally acute to subcordate, the petioles 1-5 mm. long, the stipules bifid or trifid, rarely entire, the divisions setaceous, 0.5—3.5 mm. long. Inflorescences of lax few-flowered cymes, the peduncles 5-40 mm. long; bracts ovate, strongly 1-ribbed, scariose, apiculate, 1.5—3.0 mm. long, transparent save for the midrib, mostly shorter than the pedicels, the pedicels subglabrous to densely stipitate-glandular, 2—6 mm. long. Sepals 5, 2.5-6.0 mm. long, 1.5-2.2 mm. broad, glabrous to densely stipitate- glandular, lanceolate to narrowly ovate, trinerved, acute, the midrib occasionally excurrent, the laterals often subapically dissipating; petals 5, 2.5—6.0 mm. long, bifid one-half to two-thirds their length, the lobes oblong, apically emarginate, with one dichotomous vein, exappendiculate, continuous with the trunk, the trunk as long as or longer than the claw; stamens 4—5, 2-4 mm. long, the oblong anthers 0.3—0.8 mm. long; ovary at anthesis ellipsoid, exceeded by the anthers, the elongate style trifid one-third to one-half its length, slightly exceeding the anthers. Capsule 3-4-valved, 2-4 mm. long, many-seeded, the seeds 0.5-0.7 mm. broad, evenly tuberculate, the dorsal tubercles conical, the facial tubercles substellate. Fig 10D (Isotype of D. chihuahuensis); Fig. 10E (Isotype of D. laxiflora). Texas through continental Mexico to Guatemala, to the south becoming less glandular and tending to have larger floral parts. 1961) DUKE—REVISION OF DRYMARIA 223 Examination of several isotypes of Hartweg 523 from Guatemala reveals that the plant, although described as glabrous, possesses a few scattered stipitate glands. Bernoulli & Cario 3255 at Leningrad, collected in Quezaltenango, matches the types of D. laxiflora in all respects except that it is more glandular. Toward the northern end of the range, the species has some very glandular representatives whose seeds are nearly black, pleiochroistic and very ornately sculptured. In spite of the extreme variation, some of which appears to be along a north-south cline, only one species seems to be involved. Bourgeau 2046 and Rose & Hay 5676 from Mt. Orizaba have petals, the lobes of which are so deeply emarginate as to recall the twice dichotomous petals of D. longepedunculata, and it is here suggested that the Series LAxIFLORES is genetically closely related to the Series EXCISAE. М. Series УПЛОЗАЕ Leaves opposite, sp ӘЛДЕН mostly long-petiolate, and ovate to orbicular, often villose. Flowers іп few-many-flowered cymes, rarely subsolitary in the axils. Sepals acute or obtuse, чой Petals (occasionally absent) merely bifid, the lobes usually ciliate-auriculate, 1—7-nerved, unguiculate. Stamens 3-5. Seeds cochleate, tuberculate, the tubercles stellate, cylindric or capitate. Four species of continental Mexico, Central and South America, one introduced in Malaysia. Figs. 11 and 12. a. Sepals сећа glabrous, unctuous; petals with 2-4 filiform auricles on either side; leaves owly to 36 oadly orbiculate, not basally cordate; stipules porns “lanceolate Mexico E oed Ric MULTIFLORA a. Sepals lanceolate to ovate or be to piesne glabrous, rarei or villose, occasion- ally unctuous; petals with 0—6 filiform auricles on either side; leaves ovate to reniform, rarely манаа occasionally ge: tae бригу entire or агас, vir divisions filiform to lanc b. su. and petals 3.5—7.0 mm. long; leaves cordate, 10—30 mm. broad. Mexico. e. d vie than the peti ioles; ee аи midrib = A sd s exc rrent; petals with 1 ciliate auricle on either sid CONZATTII e. isis longer than the ле 351 villose; midrib of the outer s apically ed petals with 3-6 ciliate auricles on either side............. 33. e nma b. Sepals 1.6-3 m. long, the uh rarely to 4 mm. long, but then divided seer to their ale fake ovate to reniform, occasionally with и ies 5-15 mm. Mexico to South America; Malaysia. d. Sepals elliptic, acute (rarely obtuse, but then rather strongly trinerved) ; petals, when present, divided about half their length, with 1-4 linear auri icles on either side; capsule not or к ber slightly longer than the sepals; plants villose ~ hirsute. Mexico to Peru; Malay: a. D. VILLOSA ssp. VILLOSA d. Sepa " еткін to orbicular, obtuse; petals, when present, divided halfwa ay or almost completely to their base, the auricles, when present, deltoid t ар orm, if БИ ог the petals divided nearly to the base; capsule often twice as long A = Pan glabrous to villose. Mexico to Peru, lacking in southern Centra уби е. Sepals glabrous; petals occasionally auriculate, often divided My to the base; tubercles of сна зе = ада or capitate, the dorsal seers а longer than broad. Mexico VILLOSA ssp. PALUSTRIS е. Sepals v ign exauriculate, divided about half their d tubercles of the seeds е the dorsal tubercles rarely longer than broad. zuela, ска mbia. 34с. 5^ асы . PARAMORUM (Мог. 48 224 ANNALS OF THE MISSOURI BOTANICAL GARDEN 31. DRYMARIA MULTIFLORA Brandegee, in Zoe 5:232. 1906. (HOLOTYPE: Purpus 1653; dry hills, Salto de Agua, Mexico, UC!; isotypes at F, GH, MO, US, etc.) Erect or ascending perennials to as much as 50 cm. tall, much branched, the internodes mostly longer than the leaves, glabrous to stipitate-glandular, rarely villosulous. Leaves opposite, the blades glabrous, 4-22 mm. long, 4-25 mm. broad, broadly deltoid-ovate, orbicular or reniform, apically rounded to deeply emarginate, marginally entire, basally truncate to acute, not cordate, weakly 3—5-nerved; peti- oles 2-20 mm. long, the stipules entire, lanceolate, 1.0-2.5 mm. long. Inflores- cences of terminal 3-many-flowered cymes, the peduncles to as much as 10 cm. long, often stipitate-glandular; bracts ovate, 2.5-5.0 mm. long, the pedicels mostly 5-22 mm. long, glabrous or stipitate-glandular. Sepals 5, glabrous, unctuous, rarely with sessile glands, lance-deltoid to deltoid-ovate, apically acute, at least by extension of the midrib, 3-6 mm. long, 1.2-2.0 mm. broad, often saccate basally; petals 5, 2.5-7.0 mm. long, usually equaling or exceeding the sepals, bifid about two-thirds their length, the lobes apically rounded, oblong 2—5-пегуе4, basally provided with ca. 3 ciliate auricles; stamens 3—5, 2.0-3.5 mm. long, the oblong anthers 0.3-0.5 mm. long; ovary at anthesis globose to ovoid, the style 1-2 mm. long, bifid or trifid about half its length. Capsule 2.0-3.5 mm. long, ovoid, 4—26-seeded, the seeds ca. 1 mm. broad, the dorsal tubercles longer than broad, the facial tubercles substellate. Fig. 11A (Isotype). Sinaloa, Mexico, to Costa Rica. Standley 83478 from Quezaltenango is highly aberrant. Many flowers are apetalous; others have entire liguliform petals; two flowers had twice-dichotomous petals suggestive of D. longepedunculata. 32. Drymaria conzattii J. Duke sp. nov. Plantae annuae erectae vel diffusae glanduloso-villosae usque ad 45 cm. altae internodis saepe quam foliis longioribus. Folia opposita glanduloso-villosa reni- forma apice vix acuminata basi cordata leviter 5—7 nervata, 5-20 mm. longa, 5-30 mm. lata, petiolis brevibus ad 2.5 mm. longis. Stipulae integrae vel laceratae lobis filiformibus quam petioli longioribus. Inflorescentiae terminales axillaresque in confertas 3-11-floriferes cymas dispositae rhachide glanduloso-villoso bracteis ovatis 3.0—4.5 mm. longis costis excurrentibus quam pedicellis longioribus. Sepala 5 externa minuta glanduloso-puberula angusto-ovata acuta conspicue 3-costata, 5.0—5.6 mm. longa, 1.8-2.0 mm. lata, interna paulo breviora paene glabrescentia; petala 5, 4.0—4.5 mm. longa 3/4-bifida lobis oblongis obtusis gradatim ad angusto- oblongum unguem constrictis basi ciliatis; stamina 5, 3.0—3.5 mm. longa antheris oblongis ca. 0.6 mm. longis anthesi subglobosis ca. 0.6 mm. longis; ovarium anthesi subglobosum ca. 1 mm. longum stylo subaequale, stylo vix divisio quam antheris breviore. Capsulae maturae non visae. Fig. 11B (Holotype). Mexico: Oaxaca: de Almoloya à Sta. Catarina, 1000 m., С. Conzatti 1688. ното- TYPE MO; ~ US. Known only from the type collection. 1961] DUKE—REVISION OF DRYMARIA 225 33. DRYMARIA MALACHIOIDES Briq. in Ann. Cons. & Jard. Bot. Gen. 13:372. 1911. (HOLOTYPE: Galeotti 4415; Mexicum: Cordillére d'Ario, 4000'; G!; isotype at P!) Clambering or ascending herbaceous annuals to as much as 45 cm. high, the internodes longer than the leaves, villose with jointed hairs to as much as 1 ong. Leaves opposite, the blades reniform to deltoid-ovate, 1-3 cm. long, 1-3 cm broad, apically rounded, marginally entire, obtuse to cordate basally, weakly . D. multiflora Grotype): petal, 7 ig gr 2 ee seed, onzattii Кас. Seria of plant, и Х; petal, 6X; $ пар ме Ио іа масы “ower, о petal, тя pal, 6X os pa Tec ад Ра УЯ 6: ancae); petal, 10X; sep: ; „2036: iei cad of D. hirsuta); portion of vos 1%Х; petal, 10%; = 10x; seed, Peg (Мог. 48 226 ANNALS OF THE MISSOURI BOTANICAL GARDEN 5-nerved, scantily villose; petioles 1-20 mm. long, the stipules entire or bifid, the divisions lanceolate to filiform, 1-2 mm. long. Inflorescences of terminal 3—many- flowered cymes, ultimately lax, the peduncles 1-6 cm. long; bracts 1.5-2.5 mm. long, the villose pedicels to as much as 12 mm. long. Sepals 5, 4.5-5.0 mm. long, 1.5-2.0 mm. broad, elliptic, apically acute, obscurely 3-nerved, with a few sessile glands; petals 5, 5-6 mm. long, bifid about two-thirds their length, the lobes spatulate, apically rounded to apiculate, several-nerved, basally provided with 4—6 ciliate auricles; stamens 5, 2-4 mm. long, the oblong anthers 0.5—0.8 mm. long; ovary at anthesis ovoid, the styles longer than the ovary, trifid about half their length. Capsule ellipsoid, ca. 4 mm. long, several-seeded, the seeds ca. 0.6 mm. broad, cochleate, tuberculate, the dorsal tubercles filiform, the facial substellate. Fig. 11C (Holotype). nown only from the type collection, probably in Michoacan, and a few other collections near Tancitaro, Michoacan. Certain characteristics of the plant suggest that it might have been derived from D. villosa, perhaps through polyploidy, since most of the plant parts are proportionately larger than their equivalents in D. villosa 34a. DRYMARIA VILLOSA subsp. VILLOSA Drymaria villosa Cham. & Schlecht. іп Linnaea 5:232, 1830. (нототурЕ: Schiede 8 bbe 505; in aquosis prope Jalapam, B, probably ed isotype at LE! Drymaria hirsuta Bartl. in Presl, Rel. Haenk. i 831. (noLorvPr: Haenke s.n. Habitat in Расим montanis huanoccensibus, PR 5i Drymaria cubensis Regel, in Otto & Dietr. Allg. Garten. 8:298. 1840. (No type indi- ed; horticultural) Drymaria ciliaris Hort. Berol. ex C. A. Mey. in Ind. Sem. Hort. Petrop. 9:71. 1843. (Hort.; no type indicated) Drymaria rg Hort. Berol. ex C. A. Mey. loc. cit. 9:71. 1843. (Hort; no type indica Drymaria dite var. pilosa Schlecht. in Linnaea 26:374. 1853. (no type indicated) dicioni ria cordata var. 8 villosa (Cham. & Schlecht.) Rohrb. in Mart. Fl. Bras. 14?:260. 872. а пазе Backer, in Bull. Jard. Bot. Buitenz. 21?:15. 1913. (no type s ange a a M. E. oo abe Jones, in Contr. West, Bot. 15:124. 1929. (HOLOTYPE: Jon пар Tepic, Abe dea POM, 1) Dios barrancae ones, loc. cit. 18:65. 1931. (HOLOTYPE: М. E. Jones 270514; arranca, Guadalajara, POM!) Prostrate or ascending annuals to as much as 45 cm. long, the internodes mostly longer than the leaves, villose to hirsute with septate hairs to as much as 2 mm. ng. Leaves opposite, the blades scantily to densely villose or hirsute with cinereous or ochraceous hairs, orbicular to reniform, apically rounded to acute and apiculate, basally cordate to truncate, weakly 3-7-хеіпей, 5-15 mm. long, 5-15 mm. broad; petioles 1-10 mm. long, the stipules mostly entire, 0.5-1.5 mm. long, scarcely distinguishable from the indument. Inflorescences of terminal and axillary 5—many-flowered cymes, the ultimate branches often tending to be racemose; peduncles 1-5 cm. long; bracts 0.5-1.5 mm. long, the pedicels 2-20 mm. long, villose. Sepals 5, 2.0-3.6 mm. long, 1-2 mm. broad, narrowly to broadly ovate or elliptic, apically acute to obtuse, villose, occasionally glabrescent, weakly C ОИ 1961] DUKE— REVISION OF DRYMARIA 227 3-nerved, with translucent borders; petals 5 [absent or drastically reduced in D. villosa forma tepicana (M. E. Jones) J. Duke com. & stat. nov.], 2.0-3.6 mm. long, bifid for half their length or more, the lobes apically acute to deeply emarginate, 4-nerved, basally provided with caducous filiform auricles, these variable in number and orientation; stamens usually 5, 2.0-3.5 mm. long, the oblong anthers 0.3-0.5 mm. long, the filaments shallowly connate, devoid of staminodia; ovary at anthesis ovoid to globose, the style 1.0-1.5 mm. long, trifid to as much as half its length. Capsules ovoid to ellipsoid, 2.0-3.5 mm. long, equaling or exceeding the sepals, many-seeded (rarely as few as two in D. bar- rancae), the seeds cochleate, 0.5—0.9 mm. broad, tuberculate, the dorsal tubercles cylindric to capitate, the facial tubercles stellate. Fig. 11D (Holotype of D. barrancae) ; Fig. 11E (Isotype of D. hirsuta). Central Mexico through Central America along the western coast of South America to Peru; apparently introduced and rather widespread in the East Indies. Examination of adequate material from Mexico, type locality of D. villosa, from Peru, type locality of D. hirsuta, and from Java, type locality of D. stylosa, clearly shows that these are conspecific. Indonesian material is marked although incon- stantly, by glabrous sepals and hirsute indument, in contrast to Mexican material, which is predominantly villose. I have seen no specimen representing this species from the West Indies, and believe very strongly that the epithet cubensis results from error or misconception. The contention seems to be supported by the following quote (from Ind. Sem. Hort. Petrop. 9:71. 1843): “1093. DRYMARIA CILIATA. Eadem planta, jam diu sub D. ciliaris et D. ciliatae H. Berol. nomine culta, nunc sub nomine D. cubensis H. Berol. in hort. occurit. Num sit D. cubensis species a D. ciliata distincta? Ignoramus. — Nostra planta D. villosae est proxima et vix nisi pilis in caule et foliis rarioribus ad illa differt. M." I have seen numerous horticultural specimens under these names from various European herbaria and they were all referable to D. villosa. 36b. DRYMARIA УПЛОЗА ssp. palustris (Cham. & Schlecht.) J. Duke comb. & stat. nov. Drymaria balustris Cham. & Schlecht. in Linnaea 5:232. 1830. (HOLOTYPE: Schiede 8 Deppe 504; in paludosis prope Jalapam, B, probably destroyed; Lectotype Schiede 8 Deppe 405, 1. Drymaria pauciflora Bartl. in Presl, Rel. Haenk. 2:8. 1831. (HOLOTYPE: Haenke $. п.; Habitat in Peruviae montanis huanoccensibus; PR!) Drymaria cordata var. В palustris (Cham. & Schlecht.) Rohrb. in Mart. Fl. Bras. 14?:260. 1872. Drymaria townsendii Robinson, in Bot. Gaz. 30:58. 1900. (HOLOTYPE: Townsend 8 Barber 231; on the Sierra Madre 8 km. southeast of Colonia Garcia, Chihushus, GH!; isotypes at F, MO, US, Drymaria аны Briq. in Кой Conserv. & Jard. Bot. Gen. 13:371. 4t (HOLO- ТУРЕ: Galeotti 4416; Мезгш: een ae 4000’, G!; isotype Drymaria subsessilis M. E. Jones, in Cont: №: 131125; 1929. НИИ М. Е. ones 22848; in rivulets at Ixtlan, Nayarit, “PO !) a олз нн Baehni & MacBride, in Field Mus. Bot. 137:625. 1937. (HOLO- Bride 1542; sunny sp. hansen bog, Mito, Huánuco, Peru; Е!) (Мог. 48 228 ANNALS OF THE MISSOURI BOTANICAL GARDEN Prostrate annuals or erect perennials (in var. perennis J. Duke var. поу.“) shorter to longer than the leaves, glabrous, glandular, or rarely villose and glabres- cent, the prostrate forms often rooting at the nodes, the erect forms tending to become suffrutescent. Leaves opposite, the blades 2-15 mm. long, 2~15 mm. broad, deltoid-ovate to reniform, apically rounded to apiculate, marginally entire, basally obtuse to cordate, glabrous, glandular, or villose and usually glabrescent, weakly 3~7-nerved; petioles 0.5-6.0 mm. long, the stipules entire, (rarely lacerate in var. perennis) filiform to Lareidiloid, 0.5—2.0 mm. long. Inflorescences of terminal and axillary 1-many-flowered cymes, the peduncles 0-10 cm. long; bracts ovate, to elliptic, acute to obtuse, 0.5—2.0 mm. long, the pedicels glabrous, rarely glandular or villosulous, 1-15 mm. long. Sepals 5, 1.0-3.0 mm. long, 1-2 mm. broad, glabrous or with sessile glands, oblong to orbicular, apically obtuse and often cucullate, very obscurely trinerved, the midrib subapically dissipating, often distally saccate; petals 5 [absent in subsp. palustris forma townsendii (Robinson) Duke comb. & stat. nov.], 1-4 mm. long, bifid from half to nearly all their length, the lobes oblanceolate to oblong, apically acute to emarginate, 1- 2-nerved, the shallowly cleft petals exauriculate or with mere dentations, the deeply cleft petals usually with ciliate, downwardly directed auricles; stamens 2-5, 1.0—2.5 mm. long, the oblong to suborbicular anthers 0.2-0.4 mm. long; ovary at anthesis ovoid to globose, the styles 1.0-1.5 mm. long, bifid or trifid half their length or more, usually at least as long as the ovary. Capsule 2.0-3.5 mm. long, usually exceeding the sepals, occasionally twice as long, 5—24-seeded, the seeds cochleate, 0.6-1.1 mm. broad, densely tuberculate, the dorsal tubercles longer than broad, occasionally capitate, the facial tubercles usually stellate. Fig. 12A (Holotype of D. villosa ssp. palustris var. perennis) ; Fig. 12B (Isotype of D. nummularia) ; Fig. 12C (Isotype of D. pauciflora); Fig. 12D (Holotype of D. sphagnophila) ; Fig. 12E (Holotype of D. townsendii) ; Fig. 12F (Holotype of D. subsessilis). Mexico through Central America at high altitudes down western South Amer- ica to Peru. Bolivia. Three rather easily discernible elements may be distinguished in this subspecies. Firstly, there is typical ssp. palustris, which (if I am correct in assuming that Schiede & Deppe 405 at Leningrad is the same as Schiede 8 Deppe 504, the holotype) has large petals cleft nearly to the base and provided with ciliate auricles. This is an annual plant, usually prostrate and often rooting at the nodes. I have seen no specimens from outside Mexico. Among Mexican specimens of this element are M. E. Jones 22848, Bell & Duke 16624, Rose & Hay 4800, H. E. Moore 2801, Hinton et al. 12233, and J. H. Maysilles 7782. By far the most common element is that prostrate form whose petals are not so deeply cleft and not so ornate, i.e. there are no ciliate auricles. In habit this is similar to the typical form and no other characters are discernably correlated. *D. vILLOSA ssp. PALUSTRIS var. panna J. Duke var. nov. Subspecie typica differt habitu erecto perenne, sepalis quoque angusto-ovatis vel oblongis ы glandulosis apice lividis uai, orden , rire жы fissis ны еді ег HOLOTYPE: Palmer 187; Alvarez, San L otosí, Mexico, 1961] DUKE—REVISION OF DRYMARIA 229 - 12: D. villosa ssp. palustris var. perennis (holotype); seed, 20; petal, 1212 X ; бергі, 124%; potion of plant, 34X. В. D. villosa ssp. ој (isotype of D. nummularia); — 10Х; sepal, 1 ох; seed, 20X. С. D. villosa ssp. palustris (isotype of D. pauciflora) ; petal, 15Х; бергі, 15X. D. D. villosa ssp. 5 sat of D. vente ‘petal, 10X; sepal, 10X; seed, 20X. villosa ssp. palustris forma townsendii (holotype); sepal, 10X; Sx. . D. villosa m deri боре 1 D. subsessilis) ; ae 15X; ы, 15Ж. С. D. villosa ssp. ратетотит (holotype seed); petal, 5X ; sepal, 5X ; stamen, 5X ; seed, 20 X. (Мог, 48 230 ANNALS OF THE MISSOURI BOTANICAL GARDEN To this more prevalent portion of the population, the following names have been applied: D. pauciflora, D. nummularia, D. sphagnophila. The third element seems to have received no name previously. To this belong erect perennial relatives which have little else in the way of differentia, except a more pronounced tendency to have subacute, somewhat cucullate sepals and glandular herbage. Representing specimens of the var. perennis, which may have resulted from gene exchange with D. laxiflora or one of its relatives, are Palmer 356, and 187, Pringle 3265, I. M. Johnston 8965, M. T. Edwards 804B, H. E. Moore 1669, and Salinas, Rowell 9 Barkley 16M375. In Ecuador where the plants are considered medicinal, this, the “Drimaria llamba" is reputed to be the most potently therapeutic of the local species. 34c. DRYMARIA VILLOSA subsp. paramorum (S. F. Blake) J. Duke comb. & stat. nov. Drymaria paramorum S. F. Blake, in Contr. U. S. Nat. Herb. 20:521. 1924. (HOLOTYPE: . Jabn 111; Paramo de la Cristalina, Trujillo, Venez.; alt. 2900 m.; US!) Prostrate or ascending annuals to as much as 20 cm. long, the internodes mostly longer than the leaves, sparsely to densely villose with caducous septate hairs. Leaves opposite, the blades 5-10 mm. long, 4-8 mm. broad, broadly ovate, apically acutish, basally rounded to cordate, trinerved, sparsely to densely villose; petioles 0.5—3.0 mm. long, the stipules entire, scariose, lance-deltoid, ca. 1 mm. ong. Inflorescences of lax 3—many-flowered cymes, the glabrous to villose peduncles to 4 cm. long; bracts scariose, ovate, acute, mucronulate, 1.0-2.5 mm. long, the glabrous to villose pedicels 2-6 mm. long. Sepals 4—5, subequal, 3.5—4.2 mm. long, 1.2-2.0 mm. broad, the outer villose, broadly oblong, obtuse, weakly trinerved, marginally scariose, the inner glabrous, almost nerveless, broader; petals 4-5, 3.5—5.0, bifid about half their length, the lobes oblong, 3—7-nerved, obtuse, exauriculate or with minute deltoid auricles at the base, tapered abruptly to the linear claw; stamens 4—5, 2-3 mm. long, the oblong anthers са. 0.5 mm. long; ovary at anthesis subglobose, the style trifid about half its length, slightly exceeded by the anthers. Capsule equaling or exceeding the sepals, са. 3.5 mm. long, 15- 34-seeded, the seeds 0.8—1.2 mm. broad, tuberculate, the tubercles stellate. Fig. 12G (Holotype; [seed from Pittier 131701). Trujillo and Merida, Venezuela; also from the Cordillera Oriental in Colombia. Certain specimens of D. ovata from Ecuador (e.g. Fosberg & Giles 23225 and Rose & Rose 23225) seem to share some of the characteristics of this subspecies. O. Series GRANDIFLORES Leaves opposite, ovate to reniform, sessile or petiolate, entire or crenulate, stipulate, the stipules entire or lacerate. Flowers in lax or subcapitate terminal or axillary many-flowered cymes. Sepals acute or obtuse, weakly to strongly 3-7- nerved, the midrib often carinate. Petals bifid (absent in D. apetala), unguiculate (subexunguiculate in D. glaberimma), exauriculate or with deltoid auricles, the lobes 2—15-пегуед. Stamens 5. Seeds cochleate, tuberculate (smooth in forms of D. ovata), the tubercles domical or conical, not stellate nor capitate nor cylindric. 1961] DUKE—REVISION OF DRYMARIA 231 8 species from Colombia to Argentina, along western South America. Figs. 13 and 14. . Seeds lustrous, smooth to papa a petit at anthesis obconical; petals about equaling the sepals, bifid ca. half their length, e culate. b. Seeds КУ leaves glabrous r pergameneous, the stipules ca. as long P the petioles D. FIRMULA b. Seeds s ен to ранне leaves glabrous to villose, Mri ies to pergameneous, the orsa usually longer than the stipules; Colombia to г” LENEE IMS URN 36. D. ОУАТА . Seeds not lustrous, variously tuberculate; flowe anthes any pyriform; petals minute ei twice as long as “А sepals, bifid one- "half to nag ore their length, occasion- ally auriculate. c. Petals vestigial or absent; sepals 6-10 mm. long 37. D. APETALA c. Petals present and obvious; br 2.5—8.0 mm. long. . Sepals acute, glabrou escences often pei hr gas in the leaf axils; petals occasionally su aka eda the lobes oblo: е. eae me hg the seeds secondarily а trunk of the petal за amr claw D. GLABERRIMA e. Tele “of the seeds simple; trunk of the petals shorter than the "mis Galapagos 39. D. MONTICOLA a 5 4 acute to obtuse, if acute, carinate or glandular; inflorescences terminal; petals uicu E -— bad to narrowly obovate: ^ Ted the lobes oblong, tapered to the claw; leaves — to ovate, dba n to л. often strongly ped: за cate of the seeds obtu g. Sepals more or йети obtuse, shorter than е petals, not carinate, —_ glandular or villose; leaves not crenulate; alpine Per GRANDIFLORA 5. Sepals am de as long as rw es the outer carinate, de carinae ofte serrulate; es often crenulate; lomas of Peru........... 41c. D. PAPOSANA var. она ДНА Ж. а ps than m. long, if rarely that long, with narrowly ov iub. truncate Пана ovate, glabrous to glandular, not idle ува ri of the агч ан Һ. Sepals она carinate, 4.0-6.5 mm. long, usually glandular; loma plants of Peru and Chile: a а па pow wig. truncate to the ет, conspicuously exserted; rescence capitat . РАРОЗАМА Var. WEBERBAUERI t Pel: = = сре to the claw, slightly * = ш иеЫ наш capit escence var. PAPOSANA h. Sepals obtuse Е carinate, then obtus рив вно ог Slightly glan duh je nd the Galapagos blande, - edam 2d Bolivi j- Sepals ellipsoid to orbicular, very obtuse, — mi slandular аА 42а. var. ROTUNDIFOLIA і. Sepals narrowly to broadly ovate, acutish, rant fen carinate. aeree b. D. ROTUNDIFOLIA var. NITIDA 35. DRYMARIA FIRMULA Steyermark, in Field Mus. Bot. 28:227. 1951. (ного- TYPE: Steyermark 56536; Venezuela; Merida; Páramo de los Colorados, between El Molino & San Isidro Alto, 2745—2955 m. F!) Ascending or erect herbs to as much as 25 cm. high, the internodes mostly longer than the leaves, often cyanic. Leaves opposite, the blades glabrous, sub- pergameneous, 3.0-8.5 mm. long, 2.5-7.0 mm., apically acute, marginally involute, basally cordate, subsessile or on petioles to 1.5 mm. long, the stipules lacerate, the divisions linear, caducous, equaling or exceeding the petioles. Inflorescences of lax terminal 3-7-flowered cymes, the axes glabrous to minutely glandular; pr ovate, l-nerved, scariose, 1.5—2.5 mm. long, mostly equaling or exceeding subtended pedicels. Sepals 5, subequal, glabrous, 3.8—4.8 mm. long, 1.4-2.0 mm., [Vor. 48 232 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 13: А. D. firmula нур) petal, 6X; sepal, 6X; stamen, 6X ; seed, 17\%Ж. В. D. ovata; petal, 6X; sepal, „© ; seed, 15X. С. D. apetala; androecium, 7:6 Х; “petal”, са. 20X; км 15 Х ; sepal, 5X. C» glaberrima (isotype); petal, 7% х; sepals, 712X; seed, 20X; tuber- es of seed, ca. 60X. E Da onticola (holotype); petal, 5X; sepals, 5X; seed, 15X; portion of е ER, D, dive petal, 3X ; sepal, 3 Х ; seed, 10X. 1961) DUKE—REVISION OF DRYMARIA 233 narrowly ovate to oblong, obtuse, trinerved, the midrib of the outer, somewhat narrower, sepals often excurrent; petals 5, 4.5-5.0 mm. long, bifid slightly more than half their length, the lobes broadly oblong, obtuse, 1—4-nerved, са. 1 mm. broad, gradually tapered to the claw, exappendiculate or unidentate; stamens 5, 3.0-3.5 mm. long, the oblong anthers ca. 0.6 mm. long; ovary at anthesis ellipsoid, the long style trifid for about one-third its length, about attaining the apex of the sepals. Capsule ovoid, 3-valved, 2 mm. long, 2-seeded, the seed cochleate, 1.2-1.5 mm. broad, dorsally tuberculate, the tubercles low and domical, facially merely reticulate, sublustrous. Fig. 13A (Holotype). Known only from the type collection. The lacerate stipules, small domical tubercles on the large seeds and the occa- sionally 1-nerved petal-lobes suggest a close affinity with D. cordata. On the other hand, the size and shape of the petals, coupled with the subsessile, subpergameneous leaves, suggest a closer alliance with D. ovata, D. monticola and D. glaberrima, and these four poorly known species seem to form a closely knit group. 36. ПкүмамА ovata Humb. & Bonpl. ех Roem. & Schultes, Syst. Veg. 5:406. 1819. (HOLOTYPE: Humb. & Bonpl. s.n.: crescit prope Quito, alt. 1460 hex. n.v.) Prostrate to rigidly ascending annuals or perennials to as much as 50 cm. high, the internodes mostly longer than the leaves, glabrous, farinose, glandular- puberulent or densely villose. Leaves opposite, the blades glabrous to villose, membranaceous to subpergameneous, deltoid-ovate to reniform, apically acute and apiculate to rounded, basally truncate to subcordate, 5-40 mm. long, 5-30 mm. broad, subsessile or the petioles to as much as 8 mm. long, the stipules mostly lacerate, the divisions filiform to lanceolate. Flowers broadly obconic, 1-тапу in lax terminal, rarely axillary, cymes, the peduncles to as much as 12 cm. long; bracts lanceolate, 2-5 mm. long, usually shorter than the glabrous to villose pedicels. Sepals 5, glabrous to villose, elliptic to ovate, 4.0-6.5 mm. long, apically acute to obtuse, marginally entire, 3-8-nerved, the inner broader, more obtuse and glabrous; petals 5, 4-7 mm. long, bifid one-half to three-fourths their length, the lobes narrowly oblong, 2—5-nerved, apically obtuse or emarginate, basally truncate or rarely tapered to the narrow claw, usually exappendiculate; stamens 5 (—6), the oblong anthers 0.4—1.0 mm., the filaments 2-5 mm. long; ovary at anthesis ovoid, the style 1-2 mm. long, bifid or trifid about half its length. Capsules 2.0—4.5 mm. long, 2-15-seeded, the seeds tightly circinnate, subdiscoid, 0.9—1.5 mm. broad, smooth or provided with low domical tubercles, often nearly black, lustrous. Fig. 13B (Lehmann 6112). Along the Andes from Venezuela to northern Argentina. Exhibiting a confusing array of variation, this polymorphic species is difficult of definition, but a rather constant correlation of flower- and petal-shape seems to prevail. The lustre of the seeds, equaled nowhere else in the genus, is usually evident, even in those forms which have tuberculate seeds. The following con- trasting characteristics may be found in representative specimens: ovate vs. reni- [Vor. 48 234 ANNALS OF THE MISSOURI BOTANICAL GARDEN form leaves, sessile vs. petiolate leaves, membranaceous vs. subpergameneous leaves, petals tapered vs. truncate to the claw, and smooth vs. tuberculate seeds. Wit this number of variables, and undoubtedly more exist, many combinations are possible, and I have seen many of them, but am unable to detect any correlations among these characteristics except that, south of Peru, all specimens have had smooth seeds and large leaves. Weberbauer 6543 from Peru differs in having subpergameneous leaves, 1—2-nerved petal-lobes, and a farinose girdle on the pedicels, characteristics suggesting that it might have resulted from hybridization with D. cordata. From Huanuco, Macbride & Featherstone 1714 has most of the features of D. cordata but the seeds are nearly smooth and lustrous, and the veins in the petal-lobes have several branches. Collections made by Prieto for W. W. Camp in Prov. Cafiar and Azuay of Ecuador possessed the colloquial names "Drimaria del cerco" and “Огітагіа macho” and the plants are utilized as treatments for liver and kidney ailments. 37. DRYMARIA APETALA Bartl. in Presl, Rel. Haenk. 2:7. 1831. (HOLOTYPE: Haenke s.n. Habitat in Chile; PR!) Stellaria virgata ipo in DC. Prodr. 1:396. 1824. po D. virgata Briq., 1911. (ного- TYPE: patr. “Alsine virgata Deless. herb.": Drymaria cda A. Gray, Bot. U. S. Expl. Eu. 126. 1854. (HOLOTYPE: Wilkes exped. s. n.; Obrajillo; US!) Drymaria virgata Виа. in Ann. Cons. & Jard. Bot. 13:370. 1911. Virgate or sparingly branched suffrutescent perennials (?) to as much as 30 cm. high, the internodes mostly longer than the leaves, locally farinose. Leaves opposite, the blades glabrous to minutely glandular, narrowly to broadly ovate, apically acute, basally cordate and somewhat clasping, 5-15 mm. long, 3-9 mm. broad; stipules mostly entire, deltoid, acuminate, caducous, 0.5—1.5 mm. long. Inflorescences of rather dense few-flowered cymes, the axes often densely glandular; bracts lanceolate, acuminate, strongly 1-ribbed, 2.5—4.0 mm. long; pedicels 2-10 mm. long. Sepals 5, subequal, rather densely glandular, 7.5-9.0 mm. long, narrowly ovate, apically acute with the midrib often excurrent as an apiculum, basally rather strongly trinerved; petals 5, reduced to mere vestiges or completely absent; stamens 5, 5.5-6.5 mm. long, the oblong anthers 1.8-2.0 mm. long; ovary at anthesis narrowly ellipsoid, the trifid style exceeded by the anthers. Capsule included, 5.5-6.5 mm. long, many-seeded, the submature seeds cochleate, rather sharply pointed at the hypocotylar end, dorsally sulcate, weakly tuberculate, the tubercles low and subhemispherical. Fig. 13C Although attributed to Mexico, erroneously, and questionably to Chile, I have seen no specimens that were collected outside of Peru. e Pavon specimens from “Mexique” and from Peru, probably are duplicates of one collection. 38. DRYMARIA GLABERRIMA Bartl. in Presl, Rel. Haenk. 2:7. 1831. (HOLOTYPE: aenke s.n.; habitat in Peruviae montanis huanoccensibus; PR!; isotype at GOET) Procumbent or ascending lignescent annuals or perennials to as much at 45 cm. high, the internodes mostly longer than the leaves. Leaves opposite, the blades 1961] DUKE—REVISION OF DRYMARIA 235 glabrous, narrowly to broadly ovate or elliptic, apically acute, marginally entire or undulate, basally obtuse to truncate, 3-5-nerved, subsessile, the stipules filiform, caducous, usually exceeding the petioles. Inflorescences of mostly axillary 3-11 flowered cymes, the peduncles 1-8 cm. long; bracts lanceolate to ovate, 1.5-2.5 mm., all except those of the central flowers equaling or exceeding the glabrous subtended pedicels. Sepals 5, glabrous, narrowly ovate, apically acute to obtuse, weakly 3—5-nerved, 4.0-5.0 mm. long, the inner obtuse, with a broad hyaline margin, са. 2 mm. broad; petals 5, 3.0-4.5 mm. long, bifid halfway or slightly more, the lobes oblong, obtuse, 3-7-пегуей, са. 0.5 mm. broad, the trunk ca. 1 mm broad, truncate to a minute claw, or completely exunguiculate; stamens 5, ca. 3 mm. long, the oblong anthers ca. 0.6 mm. long, the filaments basally connate into a very shallow cup; ovary broadly oblongoid, the style ca. 1.5 mm. long, trifid са. half its length. Capsule oblongoid, 2—10-seeded, the seeds tightly circin- nate, tuberculate, the dorsal tubercles longer than broad, often themselves second- arily tubercululate, the facial tubercles low and domical or slightly elongated. Fig. 13D (Isotype). Known from several stations, all in Peru. The material that I am citing for this species is heterogeneous indeed and it is possible that more than one taxon is involved. Intensive collecting and field studies will be needed to clarify this whole series, which is still full of problems. 39 DRYMARIA MONTICOLA Howell in Proc. Cal. Acad. 421:329. 1935. (HOLO- TYPE: J. T. Howell 9243; Mt. Crocker, Indefatigable Island; CAS!) Decumbent or ascending perennials, to as much as 40 cm. high, the glabrous internodes exceeding the leaves, the roots to 4 mm. thick. Leaves opposite, the blades glabrous, 3—5-nerved, ovate, acute, marginally undulate, basally truncate to subcordate, subsessile; the stipules mostly bifid, 1-2 mm. long, exceeding the petioles. Flowers in rather dense 5—9-flowered cymes, the peduncles 2-7 mm. long, glabrous, the bracts ovate, 1.0-1.5 mm. long, shorter or slightly longer than the glabrous pedicels. Sepals 4—5, glabrous, narrowly to broadly ovate, weakly 3-ribbed 4.5-6.0 mm. long, 1.5-2.5 mm. broad, apically acute, the midrib — dissipating, the lateral ribs weak, with 2 intercalary veins; petals 4-5, 4-5 m long, bifid about half their length, the lobes oblong, obtuse, 2—4-nerved, bene basally auriculate, tapered to the linear claw; stamens 4—5, 3.5—4.5 mm. long, the oblong anthers 0.7-0.9 mm. long, the filaments basally connate into a cup to as much as 0.7 mm. deep; ovary globose to ovoid, stipitate, the styles 1.5-2.0 mm. long, trifid a little more than half their length. Capsule 2.0-3.5 mm. long, 8- 16-seeded, the seeds tightly cochleate, ca. 1 mm. broad, tuberculate, the dorsal tubercles domical, slightly higher than broad, grading into the polygonal facial tubercles. Fig. 13E (Holotype). Known only from the Galapagos Islands. This species was likened by its author to D. apetala (D. macrantha) and in its sessile leaves it does somewhat resemble that species. A closer relative appears to be D. firmula from Venezuela which differs somewhat in its strongly three-ribbed sepals and exappendiculate petals. D. glaberrima also belongs in this relationship but differs in its tuberculate tubercles and obtuse inner sepals. [Vor. 48 236 ANNALS OF THE MISSOURI BOTANICAL GARDEN 40. DRYMARIA GRANDIFLORA Bartl. in Presl, Rel. Haenk. 2:7. 1831. (HOLOTYPE: Haenke s. п. Habitat in Chile, et in Peruviae montanis huanoccensibus; PR!) Prostrate or clambering rarely ascending perennials, the internodes mostly longer than the leaves, villose to glabrescent. Leaves opposite, the blades glandular- villosulous, rarely glabrous, narrowly to broadly ovate, weakly to strongly 3-nerved, apically acute, marginally entire, basally rounded to truncate, usually clasping; the main leaves 10-30 mm. long and 5-20 mm. broad, those of the branches much smaller; stipules entire to lacerate 1.0-2.5 mm. long, usually longer than the petioles Flowers narrowly to broadly campanulate in terminal, lax, few-flowered cymes, the peduncles 1-15 cm. long; bracts 2-5 mm. long, usually much exceeded by the pedicels, the pedicels usually densely glandular-villosulous, 3-25 mm. long. Sepals 5, glandular-puberulent to glabrous, ovate, weakly nerved, 6-8 mm. long, the outer sometimes carinate and acute, the inner flatter and often obtuse; petals 5, 7-12 mm. long, bifid about two-thirds their length, the lobes linear to oblong, apically obtuse, many nerved, to as much as 3 mm. broad, basally gradually merging with the poorly differentiated trunk and claw, exauriculate or rarely unilaterally unidenticulate; stamens 5, the anthers oblong 1.0-1.5 mm. long, the filaments 4-6 mm. long, connate into a cup as much as 1.5 mm. deep; ovary at anthesis ellipsoid, usually longer than the style, the style 2-3 mm. long, trifid about one-third its length. Capsule ellipsoid, 4-5 mm. long, 5—many-seeded, the seeds tightly circin- nate 1.2-1.6 mm. broad, coarsely tuberculate, the tubercles sine conical, some- what flattened, grading facially into polygonal tubercles. Fig. 1 Peru; departments Huanuco, Cajamarca, and Lima; iam GAS Chile, but perhaps erroneously. 41a. DRYMARIA PAPOSANA Vat. PAPOSANA е Lowe Phil. Fl. Atac. 10, 1860. (ноготуре: Philippi s. n.; prope Paposo іп tal alisque locis ejusdem regionis crescit, ОСО? n.v. photograph of isotype buen T ыйы Austrian herbarium; МО!) Upright or ascending annuals to as much as 25 cm. high, the internodes mostly longer than the leaves, glandular-puberulent, the taproot to as much as 2 mm. thick. Leaves opposite, the blades narrowly to broadly ovate, apically acute, basally cuneate to subcordate, glabrous or scantily glandular-puberulent, the upper leaves sessile, 5-20 mm. long, 4-18 mm. broad, the stipules lacerate, the divisions iform, mostly exceeding the petioles. Inflorescences of terminal subcapitate or monopodial cymes, the peduncles 1-30 mm. long, glandular-puberulent, the bracts ovate, usually longer than the glandular pedicels. Sepals 5, 3.5-5.0 mm. long, ellipsoid, glandular to glabrous, apically obtuse, weakly 3-nerved, the midrib sub- apically dissipating, the outermost sepals occasionally carinate; petals 5, 3.5-4.5 mm. long, bifid about half their length, the lobes oblong, rarely to 1 mm. broad, apically obtuse or emarginate, 2—3-nerved, tapered gently to the claw, exauriculate; stamens 5, 2.7-3.0 mm. long, the oblong anthers са. 0.6 mm. long; ovary at anthesis ovoid, the style ca. 1.2 mm. long, trifid about half its length. Capsules 2.6-3.0 mm. long, many-seeded, the seeds cochleate, 0.6-0.8 mm. broad, the tubercles conical, acute. Fig. 14D 1961] Fig. us Vox D. papos PERE var. we type) petal, 7%х; sepa ШАР sepal, 7% Х; = DUKE—REVISION OF DRYMARIA 237 ana var. serrulata; petal, 412 X ; viae 44X P nd var. we eberbau (holotype of D. fenzliana) ; eberba ectotype); petal, 5х; " s, dies var. paposana; petal, ЖАҒЫ sepals, Tu imd 25У: В, ; seed, 20X; portion of A sae ке C-D petal, 6X; sepal, seed, 20 x. paposana undifolia var. dais As (holo- Бе их. Во 3 иу var. nifida (isotype?); petal, 15X. [Vor. 48 238 ANNALS OF THE MISSOURI BOTANICAL GARDEN Northern Chile. Although I. M. Johnston lumped this with D. cordata, it is quite distinct from that species in many respects. 41b. DmvMARIA PAPOSANA var. weberbaueri (Muschl.) J. Duke comb. & stat. nov. Drymaria weberbaueri Muschl. in Engl. Bot. Jahrb. 45:451. July 25, dis «ооо Weberbauer 1662; Peru, Barranca prope Lima in formatione "Loma" 2 $ m; В, п.9., probably destroyed; as lectotype may be designated Wiberbesir 1699 power мын sd in Ann. Cons. & Jard. Bot. Gen. 13:375. May 10, 1911. (ного- -» “Mexique”, С!) d or 2. annuals to as much as 30 cm. tall, the few internodes mostly longer than the leaves, glabrous to glandular-pubescent. Leaves opposite, the blades glabrous to glandular-puberulent, narrowly to broadly ovate, 5-20 mm. long, 4-15 mm. broad, apically acute, basally cuneate to truncate or subcordate, usually sub- sessile, the lacerate stipules mostly longer than the petioles. Inflorescences of terminal, usually subcapitate, 3-many-flowered cymes, the peduncles 1-4 cm. long, the pedicels 0.5-3.0 (-5.0) mm. long, glandular-puberulent, usually exceeded by the ovate, occasionally dentate, bracts. Sepals 5, 3.5-7.0 mm. long, ovate, usually glandular, the outer acute and often carinate, 1—3-nerved, the inner often broader, more obtuse, less glandular; petals 5; 5-7 mm. long, usually exceeding the sepals, bifid little more than half their length, the lobes broadly oblong to spatulate, to 2 mm. broad, 3—many-nerved, apically obtuse or emarginate, basally auriculate, often asymmetrically so, more or less truncate to the linear claw; stamens 5, 3.0— 5.0 mm. long, the oblong anthers 0.5—1.0 mm. long, the filaments shallowly connate; ovary at anthesis ovoid to ellipsoid, the style 1.0-2.5 mm. long, trifid for about one-third its length. Capsule ovoid, 3-valved, often stipitate, 2.5-4.0 mm. long, many-seeded, the seeds cochleate, 0.5-1.0 mm. broad, tuberculate, the tubercles mostly domical and obtuse. Fig. 148 (Holotype of D. fenzliana); Fig. 14C (lectotype of D. weberbaueri). | Lomas of Peru from the departments of Lima to Arequip With its rather dense clover-like heads, this is perhaps the most handsome Drymaria. A nice photograph may be seen in Goodspeed and Stork (in Univ. Cal. Publ. Bot. 28: pl. 8. 1955). Although Briquet in his description of D. fenzliana believed it to be a Mexican species, it becomes apparent that the label on the Pavon specimen had suffered the same fate as those on some of his specimens of D. apetala, some of which were labeled as being from Mexico and others from Peru, although obviously belonging to the same collection. Toward the southern end of its range, this variety intergrades with the var. paposana and the newly described var. serrulata. 41с. DRYMARIA PAPOSANA var. serrulata J. Duke var. nov. Plantae perennes erectae vel glabrae sublignosae internodis quam foliis saepe longioribus, radice usque 1 cm. in diametro. Folio opposita laminis glabris anguste vel late ovatis apice acutis saepe mucronatisque basim tuncatis usque cordatis, saepe 1961] DUKE—REVISION OF DRYMARIA 239 marginibus conspicue crenulatis 5-nervatisque subsessilibus, stipulis laceratis quam petiolis longioribus. Flores 3 usque multi cymis terminalibus laxis aut vix con- fertis pedunculis 1-8 cm. longis, ie agp minus 1 ст. longis rare divaricatis. Sepala 5, 5-8 mm. longa, 2.5—3.5 mm. lata minute granulosa angusto-ovata vel elliptica, apice acuta (externissimo carinato cum carinis saepe serrulatis) ; petala 5, 6-8 mm. longa, 2/3-bifida lobis oblongis 4—6-nervatis exauriculatis vel auriculatis parvis vix ungue constricto; stamina 5, 4.5—6.5 mm. longa, antheris oblongis ca. 1.5 mm. longis; ovaria ovoidea vel ellipsoidea, stylo ca. 2 mm. longo, 3/4-trifido. Capsulae inclusae seminibus multis ca. 0.8 mm. latis crasse tuberculatis opacis. ye 14A. v: Dept. "neq Prov. Caraveli; Lomas de Анасын Penes io falda arcillosa- „С 500—550 amon Ferreyra 13490. (HO О). Not uncommon іп the lomas from the наана ins Arequipa and Moquegua, this unusual variety seems largely to replace var. weberbaueri, and although it resembles that variety in habit, the differences in their flowers are quite striking. Although the epithet applies to the carina of the outer sepals, the leaves themselves are often distinctively crenulate, so much so as to appear serrulate. See Fig. 14A. 42a. DRYMARIA ROTUNDIFOLIA var. ROTUNDIFOLIA Drymaria dear ge A. Gray, Bot. U. S. Expl. Exped. 123. 1854. (ноготуре: Wilkes Expedition s. п. Obrajillo, Peru US!) Upright iu to as much as 20 cm. high, the internodes mostly longer than the leaves, glabrous to glandular-puberulent. Leaves opposite, the blades glabrous to minutely glandular, reniform, apically rounded or apiculate, basally clasping, 4—10 mm. long, 5-15 mm. broad; the stipules lacerate, caducous. Inflorescence of rather congested 3-11-flowered cymes, the peduncles glandular, 1-45 mm. long; bracts scarious, ovate, acute, 0.5—2.0 mm. long; pedicels mostly 1.5—3.0 mm. long, those of the central flowers to 8 mm. long; glandular. Sepals 5, glabrous or sessile- glandular, broadly ellipsoid, obtuse or rounded, subcucullate, weakly 3-nerved, the midvein subapically dissipating, often carinate, marginally scarious and transparent, 2.5-3.5 mm. long, 1.2-2.8 mm. broad; petals 5, 3.2-4.5 mm. long, bifid about half their length, the lobes oblong, ca. 0.5 mm. broad, exappendiculate but somewhat flared before narrowing into the linear claw; stamens 5, about 3 mm. long, the oblong anthers about 0.6 mm. long; ovary at anthesis turbinate to ellipsoid, the style trifid about half its length, ca. 1.5 mm. long, exceeded by the anthers. Mature capsule ellipsoid, 2.5—3.5 mm. long, several-seeded, the seeds cochleate, often dorsally sulcate, 0.6-0.8 mm. broad, tuberculate, the tubercles conical, obtuse or rarely acute. Fig. 14E (Holotype). Peru. Stewart 1512 from the Galapagos Islands might just as well be included here as in var. nitida. The specimen has some of the characters of both varieties, and perhaps represents a new entity. 42b. DRYMARIA ROTUNDIFOLIA var. nitida (J. Ball) J. Duke comb. & stat. nov. Drymaria nitida J. Ball, in Jour. Linn. Soc. 22:31. 1887. (HOLOTYPE: Peru 2 ЖК "ie ndium Peruviae juxta pagum Chicla, 12-13000” s. т. J. Ball s. n. К, n. v., isotype !) [Vor. 48 240 ANNALS OF THE MISSOURI BOTANICAL GARDEN Minute to rather large ascending annuals to as much as 15 cm. high, the inter- nodes glabrous, mostly longer than the leaves. Leaves opposite, the blades glabrous, often somewhat fleshy, narrowly ovate to elliptic, 3-10 mm. long, 2-8 mm. broad, glabrous, apically acute to cuspidate, marginally hyaline, basally attenuate and clasping, the venation obscure; stipules entire, very fugaceous, acicular, less than 1 mm. long. Flowers campanulate, in rather dense few-flowered terminal cymes, the lowermost bracts merging with the leaves, often stipulate, ovate, ca. 1.5 mm. long; peduncles 4-7 mm. long, the pedicels glabrous, exceeded by or slightly exceeding the bracts. Sepals 5, glabrous, 2.5—3.0 mm. long, the outer broadly oblong and weakly carinate, the inner suborbicular, obtuse, to 2 mm. broad, obscurely nerved, the midnerve often subapically dissipating; petals 5, 3.2-4.0 mm. long, bifid about two-thirds their length, the lobes narrowly spatulate, apically obtuse, 2—3-nerved, exauriculate, acutely tapered to the claw; stamens 3—5, 2.5—3.5 mm. long, the anthers oblong; ovary at anthesis cordate, са. 8-ovulate, the style deeply trifid. Capsules subglobose, 3—8-seeded, са. 2 mm. long, the seeds са. 1 mm. broad, tuberculate, the tubercles subdendritic. Fig. 14F (Isotype). Peru, Ecuador and the Galapagos Islands. Although the type is a diminutive plant, other specimens that I refer to this variety attain a good size and in some respects approach D. glaberrima: e.g. Mandon 948 from Bolivia. Characters in the flowers however suggest that it is closer allied to D. rotundifolia and D. paposana, especially its var. weberbaueri. P. Series DIVARICATAE Leaves opposite, petiolate, elliptic to reniform, stipulate. Flowers in cymes, the pedicels usually long and divaricate, exceeding the bracts. Sepals trinerved, acute, occasionally saccate basally. Petals bifid about half their length, with mamilliform auricles (except D. divaricata forms), the lobes (1-) 2—5-nerved. Stamens 4-5. Seeds cochleate, tuberculate, the tubercles domical conical, occasionally subdendritic or mamilliform. Five poorly defined varieties in Peru and Bolivia. Fig. 15. a. Leaves b m ovate to reniform, short- to астамын. pates to villosulous; pedicels erect to strongly divaricate, mostly longer than s, if shorter, viscid; sepals usually dakie or villosulous; loma or alpine pla b. Sepals weakly nerved, basally saccate; petals ot auricled, чн auricles deltoid when present; tubercles of seeds mamilliform or secondarily tuberculat c. Leaves ры pea as en to twice as long as che pe n with mamilli- orm or coarsely domical tu D. р CATA Var. DIVARICATA c. Leaves hoyle ds about as long as the sepals; seeds with the tubercles secondarily tuberc 43b. D. DIVARICATA var. STRICTA b. Sepals strongly nerved, rather evenly keeled, not ba niis site ee petals strongly auricu- late, the auricles oblong; tubercles of the seeds domical or с d. Sepals and herbage densely viscid-glandular; pedicels not divine siping plants......... р. у> ла var. VISCIDULA d. Sepals and herbage subglabrous to villosulous; pedicels divaricate; dme plants............. 43d. D. DIVARICATA var. DIVERGENS a. ves elliptic to broadly ovate, short-petiolate, glabrous; Aena por divaricating, ону, if any longer than the flowers, glabrous; sepals p n an D. DIVARICATA var. ‚ REFLEXIPLORA 1961] DUKE—REVISION OF DRYMARIA 241 g.15: A. D. divaricata are Bu seis qe Rig Лат 4d petal, 7 алдай sepal, 7 1 X ; seed, 20%. B. n. divaricata var. al, 4X; T — € poe aad type of D. agapatensis) ; petal, 22 $ ач ў x seed, 15 Vois има уаг. ee rpm petal, 712 X ; sepal, 712 Х ; seed, 15X. E. D. divaricata var. жу secs tour, gray 7 AX sepal, 712 X ; seed, 20%; habit, 12 X. (Мог. 48 242 ANNALS OF THE MISSOURI BOTANICAL GARDEN 43а. DRYMARIA DIVARICATA Var. DIVARICATA 2 Danes me НВК. Nov. Gen. & Sp. 6:24. 1832. (НОТОТУРЕ: Humb. & Bonpl. scit ad litora Oceani Pacifici, prope Lima Peruvianorum, P n. v.) LA xg annuals, the internodes much longer than the са to as much as 10 cm. long, minutely glandular-puberulent. Leaves opposite, the blades glabrous or minutely puberulent or villosulous, narrowly to broadly deltoid-ovate, 6-12 mm. long, 6-15 mm. broad, apically acute, marginally entire, basally trun- cate; the distally alate petioles 2-5 mm. long, the stipules filiform, caducous, mostly entire, 1.0-1.5 mm. long. Inflorescence of very lax terminal cymes, the flowers many, squarely campanulate; the glandular-puberulent peduncles to 10 cm. long; bracts oblong to lance-linear, ca. 1 mm. long; the pedicels glandular- puberulent, to as much as 35 mm. long. Sepals 5, glandular-puberulent to glabrous, unctuous, triangular-ovate, apically obtuse to acutish, and often cucullate, basally saccate, 3.0—4.5 mm. long; petals 5, 5-9 mm. long, bifid about two-thirds their length, the lobes narrowly spatulate, weakly 2—5-veined, apically obtuse, basally merging gradually with the linear claw, exauriculate to minutely denticulate; stamens 5, 3.5-5.0 mm. long, the anthers oblong 0.3-0.6 mm. long. Ovary at anthesis ovoid, the style trifid one-fourth to one-half its length 1.5-2.5 mm. long. Capsule ovoid, 2.5—3.5 mm. long, са. 8-seeded, the seeds tightly circinnate, 0.8—1.5 mm. broad, coarsely tuberculate, the dorsal tubercles longer than broad, sub- dendritic. Fig. 15B (Pennell 14560). Province of Lima, Peru. Since I have not seen the type of D. divaricata, I sent specimens of all the varieties herein recognized to Dra. Lourteig for comparison with the Humboldt and Bonpland specimen in the Historical Herbarium at Paris. After comparison with the various specimens sent, she selected Ferreyra 3460 as the best match for D. divaricata HBK. Also representing the typical variety is Pennell 14560 which is illustrated in Fig. 158. This has quite long petals proportionately much longer than those of Ferreyra 3460. Both specimens have peculiar dwarf flowers in the axils of some of the unusually small bracts. Both were collected at altitudes higher than 2700 meters, which might seem incompatible with the type locality, the esie of which would at least suggest that D. divaricata might be a loma plant. 43b. DRYMARIA DIVARICATA var. stricta (Rusby) J. Duke comb. & stat. nov. Drymaria stricta Rusby in Phytologia 1:55. 1934. (нототурЕ: С. Н. H. Tate 160. Pongo de Quime Bolivia, alt. 12,000 ft. Drymaria agapatensis Baehni & Macbride, in Field Mus. Bot. 13: ad 1937. (HOLOTYPE: Lechler 1947; Puno, Agapata, G!; isotype at UPS, Е, MO, Р, е Upright or ascending lignescent annuals or perennials to as much as 50 cm. high, the internodes mostly longer than the leaves, minutely but obviously glandular-puberulent. Leaves opposite, glandular-puberulent, the blades ovate to reniform, apically acute, marginally entire or undulate, basally truncate to cordate, 5-14 mm. long, 5-20 mm. broad, the petioles often alate, 2-6 mm. long, stipules lacerate, the laciniae filiform to acicular, 1.5-2.0 mm. long. Inflorescence of 1961] DUKE—REVISION OF DRYMARIA 243 mostly terminal lax, several-flowered cymes, the peduncles 1-6 cm. long, glandular- puberulent, bracts oblong to lance-deltoid, 1-2 mm. long, mostly exceeded by the pedicels, these 1-10 mm. long, sepals 5, glandular-puberulent, ovate, basally saccate but not carinate, apically acute to shortly acuminate, subcucullate, 4.5-5.5 mm long, weakly 3—4-nerved; petals 5, 4,5-5.5 mm. long, bifid for 2-3 mm., the lobes linear, obtuse to emarginate, narrowed to the linear claw, 1-nerved, usually auriculate; stamens 5, the anthers often aberrant, oblong to orbicular, ca. 0.3 mm. long; ovary at anthesis ovoid, the style about as long, 1.5-2.5 mm. long, trifid about half its length. Capsule 2.5—3.5 mm. long 2—8-seeded, the seeds tightly circinnate, 0.8—1.0 mm. broad, densely tuberculate, the dorsal tubercles secondarily tubercululate, the facial irregularly polygonal. Fig. 15C (Isotype of D. agapatensis). Peru and Bolivia. The seeds of this variety are among the most bizarre in Drymaria. Some speci- mens are referred here only reluctantly as they approach D. glaberrima, a species which shares the unusual tuberculation of the seeds 43c. DRYMARIA DIVARICATA var. viscidula (A. Gray) J. Duke comb. & stat. nov. Home poe A. Gray, Bot. О. S. Expl. Exped. 1:124. 1854. (ноготүрЕ: Wilkes x ped. s. n.; Obrajillo, Peru US!) ce annuals to as much as 35 cm., the internodes mostly longer than the leaves. Leaves opposite, the blades glabrescent, broadly ovate, apically acute and mucronulate, basally subcordate, 5-8 mm. long, 6-10 mm. broad, the veins very obscure; petioles 1-2 mm. long, glabrescent; stipules entire or bifid, the divisions linear, deciduous, 1.0—1.5 mm. long. Inflorescences of 3—9-flowered cymes, the peduncles rather densely glandular-pubescent; bracts linear, 1.0-2.5 mm. long, gradually reduced; pedicels 1.0-2.5 mm. long. Sepals 5, subequal, the outer glandular, broadly lanceolate, acute, strongly 3-nerved, 3—4 mm. long, ca. 1 mm. broad, the inner flatter, obtuse, 1.0—1.5 mm. broad, with scariose margins outside the 5 ribs; petals 5, 3.5—4.0 mm. long, bifid about half their length, the lobes oblong, ca. 0.5 mm. broad, basally auriculate, the auricles obtusely deltoid, the claw gradually tapering to the auricle; stamens 5, ca. 3.2 mm. long, the anthers oblong, ca. 0.5 mm. long; ovary at anthesis subglobose, exceeded by the anthers, the style trifid more than half its length, slightly exceeding the anthers. Capsule ovoid, 2.8—3.5 mm. long, maturing ca. 6 seeds, the seeds cochleate, ca. 0.7 mm. broad, dorsally tuberculate with low rounded tubercles broader than long, facially flattened or sulcate, merely lineolate. Fig. 15D (Holotype). Peru, at rather high elevations. Although in some respects (especially the seeds) suggestive of D. glandulosa, I think a perusal of the plate depicting the Series РГУАКЈСАТАЕ will convince one that the var. viscidula is more properly aligned here. 43d. DRYMARIA DIVARICATA var. divergens J. Duke var. nov. Herbae annuae graciles diffusae glanduloso-pilosulosae internodis saepe quam foliis longioribus. Folia opposita laminis reniformibus apice rotundatis vel obtusis [Vor. 48 244 ANNALS OF THE MISSOURI BOTANICAL GARDEN mucronatisque basi truncatis aut rotundatis, 5-12 mm. longis, ea іп ramis lateral- ibus saepe reducta petioli saepe in longitudine laminae foliorum aequales stipulis lanceolatis frequenter quam petiolis brevioribus. Flores in cymas terminales pleures dispositi pedicellis saepe divergentibus frequenter 2-4 X in longitudine pluribus quam floribus bracteis lanceolato-linearibus usque ad 3 mm. longis. Sepala 5, 3.5—4.5 mm. longa ovata acuta 3-nervata vix stipitato-glandulosa; petala 5, sepalis aequalia ut vix plus, bifida lobis angusto-spatulatis obtusis vel truncatis 3—5- nervatis basi auriculatis ungue lineari; stamina 5 quam petala brevioria, antheris oblongis ca. 0.6 mm. latis. Capsulae inclusae seminis cochleatis. Ca. 0.8 mm. latis tuberculatis tuberculis dorsalibus latis quam longioribus. Fig. 15E (Holotype). Peru: Dept. Lima, prov. Pativilca; Lomas de Pacar, alt. 150-700 m. O. V. Nunez 2320 (HOLOTYPE, US). If I have erred in my interpretation of the typification of D. divaricata НВК, this variety of the Peruvian lomas would be my second choice, and indeed were it not for the kind assistance of Dra. Lourteig, I believe this new variety would be masquerading under the name D. divaricata HBK. The similarities of the two concepts are strong, but the slight morphological differences plus the ecologic separation seem to warrant varietal segregation. 43e. DRYMARIA DIVARICATA var. reflexiflora J. Duke var. nov. Drymaria divaricata Auct. Am. non HBK. Plantae parvae ascendentes ad 25 cm. altae internodis quam foliis saepe longi- oribus glabris. Folia opposita glabris, eis in rostellam aggregatis spatulatis longe petiolatisque gradatim foliis caulis anguste ad late ovatis vel elliptico-ovatis acutis basi truncatis saepe undulatis (in marginibus) mixtis, eisdem etiam secundum breves alatos petiolos attenuatis, petiolis saepe quam stipulis brevioribus stipulis saepe trifidis lobis filiformibus, 1.0-2.5 mm. longis. Flores campanulata in laxas evidenter trichotomas cymas foliis veris ovatis bracteis conmixtis; bractea 2—4 mm. longae, acutae vel acuminatae, prope aequales vel praestantes divaricatis glabris pedicellis. Sepala 5, glabra ovata vel elliptica, 3.0—4.5 (—5) mm. longa evidenter 3-costata basim umbonata vix saccata exteriora acuta, interiora obtusa vel acuta; petala 5, vix exserta 3.0—4.5 mm. longa, 2/3-bifida lobis angusto-spatulata apice obtusa 3—7-nervata, basi auriculata vel biauriculata tum in unguem linearem con- stricta; stamina 5 antheris fertilibus saepe reductione 3 oblongis filamentis 3.0-3.5 mm. longis; ovarium ovoideum, 2.5—3.5 mm. longum, stylo 1.0-1.8 mm. longo, 1/2-fido trilobato. Capsule ovoideae, 15—25-seminiferes seminibus constricte circinatis ca. 0.6 mm. latis tuberculis dorsalibus latis quam longioribus scabris superficie tuberculis conicis vel ellipsoideis munitis. Fig: 15A (Holotype): Peru: Dept. Lima, Prov. Chancay; Lomas de Chancay, 500-600 m.; Ferreyra 11522. (HOLOTYPE, MO). Although long confused with typical D. divaricata HBK., also reputed to be a loma plant, there is little doubt that this variety is distinct, morphologically and possibly ecologically. Plants representing this new variety bear little resemblance to var. divaricata, as depicted in a photograph of the type. {Кын кнши л енене ИЩ: ООО Чы ето 1961] DUKE—REVISION ОЕ DRYMARIA 245 Q. Series CORDATAE Leaves opposite, ovate to reniform, sessile to petiolate, entire, stipulate, the stipules entire or lacerate. Flowers in terminal cymes, rarely solitary in the axils. Sepals mostly acute, 1—3-nerved. Petals bifid (occasionally ligular in D. xero- phylla), unguiculate, exauriculate (poorly defined auricles occasionally present іп D. gracilis), the lobes 1-nerved (except D. galeottiana). Stamens 2—5, the anthers orbicular to oblong. Styles 2-3, free or basally united. Seeds 1-many, cochleate, tuberculate, the tubercles domical or rectanguloid, contiguous to remote. Five species, 1 more or less pantropical, 1 ranging from western U. S. A. to Argentina, the others confined to Mexico and vicinity. Figs. 16, 17, 18. a. Petals € equaling the i ж? the lobes income. es, „1-nerved and carinate or 3-nerved and subcarinate; seeds numerous, less than b. Leaves dls d, long-petiolate, ‘the stipules lacerate or €— Рение than the petioles; petals divided more than half their mae the lobes 1-nerved: с. Stipules lacerate; Р miel Vr ipao the er ind approximate; continental and northern Central A 44a. D. GRACILIS sp. GRACILIS с: ^s entire; ега merely EA ACA with remote diee уя Califo: 44b. D. GRACILIS irs Pium . Leaves ovate to reniform, subsessile, the stipules eee .. - the реза petals divided about half their length, the lobes 2—4-nerved..... ‚ GALEOTTIANA a. Petals shorter than the виа cen lobes oe hec сое ecarinate oe D. cordata ssp. к any, 0.5-2.0 т d. Petals much shorter than ae sepals, bilobe + ог des = ia 5-0.8 mm. broad, the facial tubercles маја or lineolate, remote; stipules lacera e. Flowers in terminal c ; leaves obviously petiolate; petals ‘ifs stamens 3—5; seeds 5—many; plants vali ебе glandulat: ш 45а. D. GLANDULOSA var. нео does clustered in axils al subsessile leaves; petals bifid or ligular; stamens eeds 1-3; plants a glabro 46 D. x XEROPHYLLA 4. Petals little to Қақ 5һогге РЕМ the sepals, bilobed; seeds 0.7-2.0 mm. broad, the tubercles more or less dotis] and subcontiguous; stipules entire or AN ate: f. Leaves pet ы the stipules "yu. зз наас ary veins forming a аср. inter- rupted reticulum; stamens 4—5, seeds 3—5; styles united for half their length or more; southern Mexico, Guatemala and во 47 D. LADEWII niform, the stipules lacerate; secondary veins obscu —5; seeds 1-12; styles He Pica зе free or united to half their athe ~ чы Id p < 2 с. о —— et o к а. о 2 > et о et o mplex: $. Flowers campanulate, the pedicels usually with a conspicuous glandular girdle; sepals ecarinate or subecarinate, glabrous to нат чеп о not peed — oad; Ө; © stamens EH seeds 1—12, 0.8—1.5 mm. broad; pantropical..... 48a. A SSp. CORDATA g. Flow the pedicels without a су pos girdle; Fani — densely Бапа apically incurved; stamens 2 (-4); seeds 1-2 (-4), 1.5-2.0 broad; Old World Корк» 48b. D. CORDATA ssp. DIANDRA 44a. DRYMARIA GRACILIS ssp. GRACILIS Drymaria gracilis Cham. & Schlecht. in Linnaea 5:232. 1830. (HOLOTYPE: Schiede & Deppe 503; prope Jalapa. В, м. v. isotype at ) Бочни cordata var. y gracilis (Cham. & Schlecht.) Rohrb. in Mart. Fl. Bras. 142:260. 72. Erect or ascending perennials to as much as 45 cm. tall; the internodes mostly longer than the leaves, glabrous or minutely stipitate-glandular; leaves opposite, the blades glabrous, rarely villosulous or punctate, deltoid-ovate (narrower upwards), 5-15 mm. long, 3-15 mm. broad, apically acute to rounded and apiculate, mar- ginally entire, basally truncate 3-5-пегуед, the petioles 1-8 mm. long; the stipules [Vor. 48 246 ANNALS OF THE MISSOURI BOTANICAL GARDEN bifid or trifid, 1.0-2.5 mm. long, the divisions filiform. Inflorescences of lax terminal, many-flowered cymes, the peduncles 2—6 cm. long; bracts ovate, carinate, 0.5-1.5 mm. long; pedicels 3-6 mm. long, glabrous, divaricate. Sepals 5, ovate, glabrous, 2.0-2.7 mm. long, apically acute, 1-пегуед, carinate; petals 5, 2.0-2.7 mm. long, bifid at least two-thirds their length, the lobes obtuse to emarginate, basally entire to auriculate, divaricate, tapered or truncate to the minute claw; stamens 5, rarely fewer, 1.8-2.2 mm. long, the anthers oblong, 0.4-0.5 mm. long, briefly connate into a shallow cup; ovary at anthesis globose, the styles elongate, bifid or trifid less than half their length, exceeding the ovary; capsule globose, equaling or exceeding the sepals, 1.8-2.5 mm. long, many-seeded, the seeds cochleate, 0.5—0.6 mm. broad, coarsely tuberculate, the dorsal tubercles domical, the facial tubercles polygonal, approximate. Fig. 16B D. glandulosa var. ee i eos e); petal, 10 1, 10 d, 22 B. D gos uj gracilis; petal, 10; 2 d | 2% ue x: e gat raci uus ssp. carinata (isotype); petal, 10X; sepal, 10x; 22. D. glandulosa өйы of D. lepto- n een XN sepal, 10X. E, D. р ув т, ма сү он of D. fendleri); petal, 10X; sepal, 1961] DUKE—REVISION OF DRYMARIA 247 Hidalgo to Vera Cruz, Mexico. Specimens in the Lund Herbarium, and the herbaria at Leningrad and the Field Museum, although labeled as grown from seeds of D. divaricata, are clearly refer- able to D. gracilis. Somewhere a mixup in labels must have occurred. For that reason, I sent a specimen of this species to Dra. Lourteig for comparison with the type of D. divaricata. As mentioned in the discussion of D. divaricata previously, she cited the Peruvian Ferreyra 3460, very different from the Mexican D. gracilis, as comparable to D. divaricata HBK. 44b. DRYMARIA GRACILIS ssp. carinata (Brandegee) J. Duke comb. & stat. nov. Drymaria carinata та j in Zoe 2:70. 1891. (HOLOTYPE: Brandegee 34; Sierra de la Laguna, Baja Calif. UC xb rains carinata var. we Wiggins, in Proc. Cal. Acad. Sci. 4?5:197. 1944. (ното- TYPE: Gentry 4415; La Laguna, Sierra Laguna UC!) Prostrate to ascending annuals or perennials to as much as 40 cm. high, the internodes mostly longer than the leaves, glabrous. Leaves opposite, the blades glabrous, narrowly to broadly deltoid-ovate, 5-20 mm. long, 2-15 mm. broad, apically rounded to acute, marginally entire, basally truncate, 3-nerved, petioles 1-10 mm. long, the stipules entire, 1-2 mm. long, filiform. Inflorescences of terminal lax 3—many-flowered cymes, ‘the peduncles 2—6 cm. long; bracts ovate, carinate, 1.0-1.5 mm. long, the pedicels 2-12 mm. long, glabrous. Sepals 5, ovate, glabrous, 1.5-2.5 mm. long, apically acute to acuminate, 1-пегуе and carinate; petals 5, 1.5—3.0 mm. long, bifid at least two-thirds their length, the lobes apically obtuse to emarginate, 1-пегуед, divaricate, basally auriculate to entire, abruptly truncate or tapered to the claw; stamens 3-5, 1.5-2.0 mm. long, anthers oblong 0.3—0.4 mm. long, shallowly connate into a cup; ovary at anthesis globose, the style trifid one-half to five-sixths its length. Capsule globose, 1.5-2.0 mm. long, many-seeded, the seeds cochleate, 0.4-0.6 mm. long, minutely and remotely tuber- culate. Fig. 16C (Isotype). Baja California, Mexico. This subspecies shares with the typical subspecies the tendency toward peren- niality. Probably separated for some time, yet obviously closely related, these subspecies exhibit an interesting disjunction. 45a. DRYMARIA GLANDULOSA var. GLANDULOSA Drymaria d idend Presl, Rel. Haenk. 2:9. 1831. (нототурЕ: Haenke s. п.; habitat in Mexico. ш ramodsim 4 Schlecht. in HM 12:206. 1838. urbem Marien, GOET (?) n Drymaria leptoclados Hemsl. Diag. РІ. Nov. 2. 1878. (HOLOTYPE: Bernoulli 240, Guate- Camino del Sapote; K!; isotype at NY) ж. Die fendleri S. Wats. in Proc. Am. Acad. 17:328. 1882. (HOLOTYPE: Fendler New Mexico, GH!) Drymaria leptoclados var. Lee jana J. Ball, in Journ. Proc. ие. бос. 22:32. 1885. HOLOTYPE: J. Ball 5. п.; Chicla, Peru, K?; isotype (?) at GH Drymaria blasiana M. E. Зай дез; Contr. West. Bot. Es: 125. oie (НОТОТУРЕ: М. E. ones ; in open places, San les: Sinaloa; PO Drymaria “ene de pueris . Е. Jones, Extr. du West. Bot. 18:65. 1933. (но T E. Jones 27050, Laguna Mtns., Baja Cal. POM!; isotypes at UC, etc.) (HOLOTYPE: Hegewisch s.n., [Vor. 48 248 ANNALS OF THE MISSOURI BOTANICAL GARDEN Drymaria glandulosa var. fendleri (S. Wats.) Fosberg, in Morton, in Contr. U. S. Nat. H 1945. Drymaria glandulosa var. perennis (M. E. Jones) Fosberg, loc. cit. 29:96. 1945. Sparingly to densely stipitate glandular upright or ascending annuals or peren- nials, the internodes mostly longer than the leaves, the taproot to 1 cm. thick. Leaves opposite, the blades sparingly to densely glandular-puberulent, ovate to reniform, 5-12 mm. long, 5-20 mm. broad, apically rounded and apiculate, basally cordate, marginally entire, the petioles 1-8 mm. long; stipules lacerate, the divisions lance-linear to filiform. Inflorescences of terminal and rarely axillary lax to dense 5—-many-flowered cymes, the peduncles (0-) 2-8 cm. long (the peduncles often axillary and abbreviated in var. fendleri); bracts lanceolate-aristate 2.5—5.0 mm. long, often exceeding the pedicels (especially in var. fendleri), lanceolate, weakly to strongly 3-ribbed, the ribs subapically confluent, the mid-rib often excurrent, the outer sepals frequently conspicuously shorter than the inner and more attenuate at the apex, 3.0-4.6 mm. long; petals 3—5, 1.2-3.0 mm. long, bifid about half their length, the lobes linear, apically acute to emarginate, 1-nerved, basally auriculate or entire; stamens 2—5, the anthers oblong, 0.2—0.4 mm. long, the filaments 1-2 mm. long; ovary at anthesis subglobose, longer than the style, the style bifid or trifid about half its length; capsule 1.5—2.8 mm. long (2-) 4—20-seeded, the seeds cochleate, 0.5—0.8 mm. broad, minutely but regularly tuberculate, the dorsal tuber- cles domical, the facial tubercles rectanguloid or lineolate, remote. Fig. 16D (Holotype D. leptoclados) ; Fig. 16E (Holotype D. fendleri). Baja California; Arizona through western South America along the Andes to Argentina; apparently skipping southern Central America. Fosberg (in Contr. U. S. Nat. Herb. 29:96. 1945) recognizes three varieties of D. glandulosa, a variable species indeed. The varieties are separated on basis of varying vegetative characters, var. fendleri having subglabrous leaves and var. perennis being perennial. Var. fendleri also tends to have cinereous less cordate leaves and more dense inflorescences. Although the type of D. glandulosa was described as being an annual, the root is apparently perennial and nearly 5 mm. thick. Since the perennial habit occurs in both the subglabrous and the densely pubescent forms and all possible combinations of the characters can be found throughout the range of the species I see little reason for their recognition. Some of the forms are only with difficulty separated from D. cordata, but the seeds of the two have markedly different sculpture. Drymaria glandulosa is also marked by longer anthers and proportionately longer inner sepals, and the inflores- cence is more obviously cymose with a conspicuous central leader. The perennial habit, frequently encountered in D. glandulosa, is rarely if ever encountered in Mandon 950 from the Andes of Bolivia (2600-2700 m.) dept: Larecaja, referred tentatively by Ball (Kew Bull. 22:32) to D. leptocladus var. peruviana is obviously D. glandulosa, and in its tendency toward glabrescence it does resemble D. leptocladus. Berlandier 999 and 1122 from Mexico are distinctive in that the petals have pli-nerved veins and subcylindric tubercles thus suggesting intergression with a member of the Series viLLosa. 19611 DUKE—REVISION ОЕ DRYMARIA 249 45b. DryMARIA GLANDULOSA var. galeottiana (Briq.) J. Duke comb. & stat. nov. Drymaria galeottiana Briq. in Ann, Conserv. & Jard. Bot. Gen. 13:373. 1911. (HOLOTYPE: Galeotti 4408; near Oaxaca, Mexico, G!; isotypes at K [sub 4416] and L.) Rather delicate erect annuals to as much as 20 cm. high, the internodes mostly longer than the leaves, glandular-puberulent, the roots fusiform. Leaves opposite, the blades glandular-puberulent and glabrescent, broadly ovate to reniform, apically obtuse and apiculate, marginally entire, basally truncate to subcordate, subsessile, the stipules bifid or trifid, 1-2 mm. long, about equaling the petioles. Flowers obconic, in rather dense terminal cymes, the peduncles 1-2 cm. long, glandular- puberulent; bracts ovate, aristate to acuminate, 1.5-2.0 mm. long, about equaling all but the central pedicels which may be up to 6 mm. long. Sepals 5, subequal, glandular-puberulent, 3.0-3.6 mm. long, 1.5-2.8 mm. broad, ovate to broadly oblong, 3-nerved, the midrib sometimes prominent and excurrent, the inner sepals obtuse, the outer acute, transparent save for the nerves; petals 5, 2.5—3.2 mm. long, bifid ca. half their length, the lobes oblong, obtuse, 2—4-nerved, tapering gradually to the claw, exappendiculate; stamens 3—5, 1.5-2.2 mm. long, the anthers sub- orbicular, 0.2-0.3 mm. long; ovary at anthesis subglobose, the style ca. 1 mm. long, trifid ca. half its length. Mature capsule subglobose, 2.5-3.0 mm. long, many-seeded, the seeds cochleate, about as long as broad, ca. 0.6 mm. broad, the dorsal tubercles low and domical, the facial tubercles rectangular, somewhat remote. Fig. 16A (Holotype). Known only from few scattered collections in Oaxaca and Hidalgo, Mexico. At Kew, the specimen labeled 4416 is called D. galeottiana and that labeled 4408 is called D. nummularia, so apparently the labels have been crossed. This variety is a misfit in this series because of the petal nervation and it may prove to represent a mere hybrid. The seeds, sepals, and pubescence suggest D. glandulosa whereas the nervation of the petals and to some extent their shape and size, recall D. laxiflora. These two are the most likely candidates for the parents if indeed the plant is a hybrid. 46. DRYMARIA XEROPHYLLA A. Gray, Pl. Wright. 2:18. 1852. (HOLOTYPE: Coulter 722). Ascending perennials profusely branched from the base, the internodes longer or shorter than the leaves, glabrous. Leaves opposite, the blades broadly to nar- rowly ovate, glabrous, strongly 3—7-nerved, 5-10 mm. long, 3-10 mm. broad, apically acute and apiculate, marginally entire, basally rounded to obtuse, subsessile; the stipules lacerate, the divisions lance-linear to filiform, exceeding the petioles. Inflorescences of dense axillary cymes, or fascicles, the lanceolate acuminate bracts longer than the glabrous peduncles and pedicels, 1.5-2.5 mm. long. Sepals 5, oblong, apically acute to obtuse, strongly 3-nerved with the nerves subapically confluent, basally umbonate, 3—4 mm. long, glabrous to scantily stipitate- glandular; petals 3—5, entire and ligulate or bifid for half their length, the lobes apically [Vor. 48 250 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 17: D. xerophylla; sepal, 15 <; petal, 15 <; seed, 15 Х ; habit, “YX. denticulate or entire, 1-nerved, 1.0-1.8 mm. long; stamens 3, ca. 1 mm. long, the anthers oblong, the filaments basally connate into a shallow cup; ovary at anthesis subglobose, the style about as long, bifid or trifid about half its length. Capsule ellipsoid, 3-valved, 1—3-seeded, the seeds cochleate, 0.7—0.8 mm. broad, minutely tuberculate or merely reticulate on the facies. Fig. 17. San Luis Potosí to Oaxaca, Mexico. 47. DryMaria LADEWII Rusby, in Phytologia 1:54. 1934. (НОТОТУРЕ: С. Н. H. Tate 652: Bolivia; Nequejahuira; alt. $000 ft. NY!) Drymaria bypericifolia Auct. Am. non Briq. Diffusely spreading annuals or perennials, the internodes much longer than the leaves, strongly divaricate, viscid-puberulent. Leaves opposite, the blades glabrous, 1961] DUKE—REVISION OF DRYMARIA 231 deltoid-ovate, apically rounded to acute and apiculate, marginally entire, basally more or less truncate; stipules entire, more or less lanceolate; larger leaves to 2.8 cm. long, 2.5 cm. broad, trinerved, with the secondary veins forming an obvious but interrupted reticulum, the leaves as the rather long petioles strongly reduced upwardly. Inflorescences of terminal and axillary 5-many-flowered cymes, the axes glandular-puberulent, the bracts lanceolate, 1.5—3.0 mm. long, exceeded by the long, divaricate pedicels, the pedicels to 12 mm. long. Sepals 5, lance-oblong, 2-3 mm. long, apically obtuse or acute by extension of the midrib, weakly 1-3 nerved, the midrib often distally subcyanic; petals 5, about equaling the sepals in length, bifid about three-fourths their length, the lobes obtuse, exauriculate, subdivaricate, weakly 2—5-nerved, the claw сагпове, subcylindric; stamens 5; 1.5-2.5 mm. long, the oblong anthers ca. 0.5 mm. long, the filaments basally connate into a more or less carnose ring; ovary at anthesis turbinate, the style trifid ca. half its length, about attaining the anthers. Capsule turbinate, 3-valved, included (seeds rarely mature in the specimens at hand), 1—5-seeded, the seeds similar to those o D. cordata. Fig. 18A. Southern Mexico, Guatemala, Bolivia. After scrutiny of the copious Guatemalan material and the scanty Bolivian material (2 specimens, one from the O. Kuntze herbarium, the other the type; [the second specimen cited by Rusby, in his description of the species, apparently is a hybrid with D. cordata or some related species and must be excluded from D. ladewii]), І am unable to find the slightest differences between them. Asserting rather strongly that there is no obvious difference between the Guatemalan and Bolivian material, I am quite at a loss to explain the peculiar disjunction exhibited by this species. 48а. DRYMARIA СОКРАТА (L.) Willd. ex Roem. & Schult. Syst. Veg. 5:406. 1819. (LECTOTYPE after Mizushima: Linnaean Herbarium specimen no. 109-1; LINN т. V.) 53. - Holosteum diandrum Sw. Prodr. 27. 1788. (HOLOTYPE: habitat in Jamaica, 5!) Triana s. n.; Bogota alt. 2600 m., Р, n. v.; isotype at NY!) Drymaria procumbens J. N. Rose, in Contr. U. S. Nat. Herb. 1:304. 1895. (HOLOTYPE: Palmer 1165, near water about Colima, Mex., US!; isotype at GH) Drymaria adenopbora Urban, in Fedde, Repert. Sp. Nov. 21:213. 1925. eri man 12978, Cuba; prov. Pinar del Río prope Cortez, B?; isotype at G, S, NY!) Stellaria adenopbora (Urb.) León in León & Alain, Flora de Cuba 2:154. 1950. Drymaria cordata var. pacifica Mizushima, in Jour. Jap. Bot. 32:78. 195 (HOLOTYPE: Svenson 65; Academy Bay, Indefatigable Island, Galapagos Islands, 100 ft. alt.; GH!) Glandular-puberulent to glabrate ramifying annuals, prostrate and spreading or erect, the internodes mostly longer than the leaves, glabrous to densely glandular, often rooting at the nodes. Leaves glabrous to scantily puberulent, orbicular to reniform, apically rounded and occasionally mucronulate, basally rounded to cordate, 5-25 mm. long, 5-30 mm. broad, the petioles 2-15 mm. long; stipules mostly lacerate, rather persistent, to 2 mm. long. Inflorescences of terminal or (Мог. 48 252 ANNALS OF THE MISSOURI BOTANICAL GARDEN axillary few—many-flowered dichasial cymes, (flowers rarely solitary in the axils) the bracteate pedicels locally girdled with a dense band of glandular pubescence, rarely subglabrous, 2-15 mm. long, much exceeding the subtending bracts. Sepals 5, lanceolate to ovate, acute, glandular-puberulent to glabrous, obscurely to strongly 3-nerved, 2.5-4.0 (-5.0) mm. long, the borders translucent; petals 5, 2-3 mm. long, deeply bifid, the lobes linear, acute, rarely obtuse, 1-nerved, basally exauricu- late but rarely subdentate; stamens 2-3 (-5), the flattened filaments 2.0-2.5 mm. long, the anthers suborbicular, 0.2-0.3 mm. long; style 0.5—1.0 mm. long, bifid or ||| INS . procumben yx; jum petal, 714 х; p 7V2X ; seed, ordata ssp. diandra (type of D. retusa ? а 12 1 12% Х Е. D. cordata ssp. diendra- (lect totype); petal, 10 Х ; sepals, 10. hacer 7% da 1961] DUKE—REVISION ОЕ DRYMARIA 253 trifid from half to its entire length; ovules few to many, campylotropous on free central placentae. Capsule ovoid, 1.5-2.5 mm. long, the 3 valves entire; seeds 1-тапу, 1.0-1.5 mm. broad, cochleate, dark reddish brown, tuberculate in lines, all the tubercles low and domical, closely approximated or contiguous. Fig. 18B (Holotype of var. pacifica) ; Fig. 18C (Holotype of D. procumbens). A nearly pantropical species, in America ranging from Florida and Mexico through the West Indies and Central America to Argentina. Mizushima (in Jour. Jap. Bot. 32:69. 1957), studying the varietal potentials of this polymorphic species, described var. pacifica, which differs from typical cordata in having glabrous sepals, more seeds, and a tendency to have a greater number o stamens and the styles divided completely to the ovary. This rather striking variety seems to be lacking in continental North America. Occuring in both var. pacifica and var. puberula, for those who wish to recognize them, are variants in which the flowers are subsolitary in the axils (e.g. Holton 733 from Colombia). Surprisingly and fittingly these have as yet received no taxonomic status. Drymaria adenopbora Urban represents a more or less viscid form with diminutive leaves and flowers and apparently is very similar to specimen 109-1 in the Linnaean herbarium. A similar reduction is exhibited by the type of Holosteum diandrum Sw. generously loaned by Stockholm. Mosen 405 at Lund from Brazil with the sepals to as much as 5.2 mm. long represents the opposite extreme of D. adenopbora, which I believe is typical D. cordata. Perhaps the strangest aberration is that represented by the flowers of the type of D. procumbens, here considered to represent only a monstrosity. The petals are bilobed but one of the lobes is clearly staminal in nature (see Fig. 18C), the other lobe being normal and petaloid. I have seen a similar aberration in one specimen of D. paposana var. weberbaueri, perhaps lending weight to Pax's interpretation of the petals as staminodia. 48b. DRYMARIA СОКРАТА ssp. diandra (Blume) J. Duke comb. & stat. nov. Drymaria diandra Blume, Bidjr. Fl. Nederl. Ind. 62. 1825. (LECTOTYPE: after Mizu- shima, sect. 99, 143—199 at L!) Drymaria retusa Wallich, ex Wight & Arnott, Prodr. Fl. Ind. Or. 359. 1834. as synonym. (? түре?: Wight 152, С, L, !) Drymaria cordata forma indica Miquel, in Junghuhn, РІ. Jungh. 395. 1855. Drymaria extensa Wallich, ex Edgeworth, & Hook.f. in Hook. 4. Fl. Brit. Ind. 12:244. 74. (? түре?: Wallich 647, С!) Drymaria gerontogea F. Muller, Descr. Papuan Pl. 1:87. 1877. Prostrate or ascending annuals, the internodes longer than the leaves, glabrous to stipitate-glandular. Leaves opposite, the blades glabrous, rarely minutely puberulent, deltoid-ovate to orbicular or subreniform, apically rounded and often apiculate, marginally entire, basally truncate to obtuse, rarely cordate, 5-25 mm. long, 3-20 mm. broad, 3—7-nerved; the petioles 2-8 mm. long usually exceeding the stipules; the stipules lacerate, the segments filiform, 1-3 mm. long. Inflorescence a terminal lax to dense 3—many-flowered cyme, the peduncles 1-12 cm. long, glabrous or glandular-puberulent; bracts lanceolate, 2-5 mm. long; pedicels evenly glandular-puberulent; 1-8 mm. long. Sepals 5, narrowly obovate, 2.04.5 mm. long, the outer glandular-puberulent, strongly carinate, strongly three-ribbed, the [Vor. 48 254 ANNALS OF THE MISSOURI BOTANICAL GARDEN ribs protrusive and often connected by cross-veins; petals 3—5, 1.5-3.0 mm. long, bifid one-half to two-thirds their length, the lobes oblong, 1-nerved, apically obtuse to emarginate, basally tapered to the linear claw. Stamens 2-3 (-4), 1.6-2.2 mm. long, the anthers suborbicular, the filaments basally connate into a shallow cup (the cup often carnose in African forms). Ovary at anthesis globose, the styles short, bifid or trifid from half to all their length. Capsule 2-3-valved, 1.5-2.5 mm. long, 1—2-seeded, the seeds cochleate, 1.4-2.0 mm. broad, rather densely tuberculate. Fig. 18D (type of D. retusa ?); Fig. 18E (lectotype of D. diandra). Tropical Africa, Australia, and Tropical Asia (China, $. & W.; India, Indo- china, Malaysia, Formosa, Ryukyu Archipelago), Oceania, Hawaii, Philippines, New Guinea, Celebes. Mizushima (in Journ. Jap. Bot. 32:69. 1957), after a detailed study of the variation of D. cordata, reinstated D. diandra, long lost in synonymy under D. cordata, to specific status. Intermediates do occur between the two, especially in rica. The absence of the taxon from the New World when weighed with the characters enumerated in the key, none of them invariably constant, suggest that reduction to a subspecies more truly reflects the hierarchial relationships. African forms are very perplexing and differ from the typical material in the large sepals and petals, higher number of stamens, with prominent, often carnose, staminodial cups. The flowers often lack the pyriform shape which so frequently differentiates ssp. diandra from ssp. cordata. Description of these variants as formal taxa would seem to serve no purpose. SPECIES EXCLUDENDAE D. filiformis Benth. = Stellaria filiformis (Benth.) Mattf. D. idiopoda S. Е. Blake = Stellaria sp. D. mairei Leville = Cerastium sp. D. palmeri Hemsl. = Stellaria sp. D. rotundifolia Harriot = Stellaria rotundifolia Poir. fide Kew Index SPECIES NON SATIS. NOTAE D. adiantoides Muschl. (туре: Sodiro 1291) Ex снак. = D. villosa? D. grandiflora Schlecht. non Bartl. May be Stellaria? D. megalantha Steud. “ = D. oxalidea Pax (түре: Schaffner 827) Ex CHAR. VEG. — D. excisa??? D. peruviana Muschl. (түре: Weberbauer 2601) Ех cuar. vec. = D. divaricata? D. sessiliflora Fiori (Descr. non viso) ЕХ СЕОСЕ. = D. cordata COMBINATIONES ЕТ Taxa NOVAE D. bolosteoides var. crassifolia (Benth.) J. Duke D. arenarioides ssp. peninsularis (S. F. Blake) J. Duke D. barkleyi J. Duke & Steyermark D. conzatti J. Duke D. effusa var. confusa (J. N. Rose) J. Duke 1961) DUKE—REVISION OF DRYMARIA effusa var. depressa (E. L. Greene) J. Duke lebtopbylla var. cognata (S. F. Blake) J. Duke eptophylla var. nodosa (Engelm.) J. Duke engleriana var. devia (Baehni & MacBride) J. Duke villosa ssp. palustris (Cham. & Schlecht.) J. Duke villosa ssp. palustris forma townsendii (Robins.) J. Duke villosa ssp. palustris var. perennis J. Duke villosa ssp. villosa forma tepicana (M. E. Jones) J. Duke villosa ssp. paramorum (5. Е. Blake) J. Duke D. paposana var. serrulata J. Duke D. paposana var. weberbaueri (Muschl.) J. Duke D. rotundifolia var. nitida (J. Ball) J. Duke D. divaricata var. stricta (Rusby) J. Duke D. divaricata var. viscidula (A. Gray) J. Duke D. divaricata var. divergens J. Duke D. divaricata var. reflexiflora J. Duke D. gracilis ssp. carinata (Brandegee) J. Duke D. glandulosa var. galeottiana (Briq.) J. Duke D. cordata ssp. diandra (Blume) J. Duke оррророоо CITATION OF SPECIMENS 255 01.5. A. D. cordata (L.) Willd. Gould oo 4050 Fredholm 6487 Green Godfrey, R. K. 58025 Hanson Ж а А962 McFarlin, J. В. 3192 Harrison & Kearney 6104 eese, 1506 Hinckley Schallert, Р. О. 7500 Jones, M. E. 96; 28677 Small, J. K. 12761 Metcalfe 793, 1378 Small, Britton & DeWinkeler 9310 Mulford 11 mall, Mosier & DeWinkeler 10836 Purpus 8147 Steyermark, J. 63326 Rose & Fitch 17647 Rus 2 О. EFFUSA var. EFFUSA Shreve 7768 Blumer 1639; 3368 (appr. leptophylla) Standley уай Harrison & Kea Tu ie hs emmon 509; 2644 rig Rothrock 169; 619 24 Wrig t 869 (TYPE) D. LAXIFLORA Benth. Cory 30590 О. EFFUSA var. DEPRESSA (Greene) Hinckley dde Tl 2847 J. Duke Innes & Моо: Gould & Robin Moore & Styermarh 3044 Metcalfe 1428 губио or (TYPE) Warnock W8 c" D. rEPTOPHYLLA (Cham. & Schlecht) D. GLANDULOSA Presl Fenzl ex Rohrb. Arséne 19280; 21665 Arséne 19375 Barkley 14NM715 Blumer 1411 Earle & Earle 460 Co е Eggleston 16381; 16469а Darro Fendler 60 (түре: D. fendleri) ee et ey 1485 [Vor. 48 256 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fendler 56 Greene 332; 12778 Harrison t гене 6187 Hinckley Metcalfe 27 ве ејјиза) “> Ач usb SANT 13932 Wright, C. 868 D. MorLUGINEA (Lag.) Didr. Arséne 18596; 19073 Blumer 1433; 2100; 3583 me 10945; 15989 Ellis 4 Fender 53 pes D. sperguloides) Gould Gould & Porn 4049 Greene 12 Hanson & Hanson 952 Harrison, Kearney & Hastings 6228; 8092 Kearney & Peebles 10042; 12723 Lemmon 3002 Metcalfe d 1424 Mulford 8 Parker & ce 7509 Purpus 8148 b Standley 14212 Wooton 593 Wright 867 D. PACHYPHYLLA Woot. & Standl. Cory 30793 Eggleston 19345; 19353 8 5 Wooton 405 (TYPE) Mexico (Continental) D. ANOMALA = Watson Bell & Duke 16581 Johnston & Maller 622A Palmer 424 Pringle 2 У (түр Е); Stanford, uta & fede 485 D. AnENARIOIDES Humb. & Bonpl. ex Roem. & Schult. Bonpland 4070 (түре?) Coulter 698 Parry & Palmer 49 Pringle 236, doe: 715, 35393 7589; 7927 112 Purpus 133 Rose, Painter z Row 8719 Schaffner 13 Shreve 6673; debe 9205 Thurber 815 White, 3. 22 2477 Wiggins 7235 D. AXILLARIS Brandegee Purpus 4526 (TYPE) D. BARKLEY! J. Duke & Steyermark Hernandez, Rowell & Barkley 16M531 Warnock & Barkley 14826M (ТУРЕ) D. coNzarTH J. Duke Conzatti 1688 (TYPE) D. сократа (L.) Willd. Berlandier 1217 Ervendberg 228 Hinton 7442 ЛЕ pacifica) Karwin 2; Kerber 70, Martinez- “Calderon 408 D. EFFUSA var. EFFUSA Thurber 995 D. EFFUSA var. DEPRESSA (Greene) J. Duke EE pore & Westlund 77 Nelso сене и Co ee & Barkley 16M699 Pringle Reiche 8 Rose МА (appr. var. confusa) Rose & Н Rose & Painter 7037; 7206; 7914 D. EFFUSA var. CONFUSA (Rose) J. Duke Knobloch 1298 Maysilles 7773-A, 7441 3 Palmer 59 (ТУРЕ) D. ЕтАТА I. M. Johnston J Johnston 7823 (ТУРЕ) 19611 DUKE—REVISION OF DRYMARIA 237 Purpus 4496 D. HYPERICIFOLIA Briq. Stewart 2651 Jurgenson 38 (ТУРЕ) D. excisa Standl. D. raewn Rusby Arséne 8450 Matuda 2008; 2283; 2994; 4000 Hinton 508; 4919; ЕАН 5427 (TYPE: Р”. grandis) ; 5429 D. LaxiFLora Benth. McVaugh 10153; MN 10: Arséne 1905; 5129 Mexía 1748 (TYPE) Balls B4472 D. GLANDULOsA Presl ell & p bh Aguilar 782 Mica dapat 2 Arséne si dd 2295 (appr. laxiflora) Doa M 706 boot: gracilis) ; 710 Bartlett 1031 4 Berlandier 677; 341; 1122; 1308 Fisher 44293 Bourgesu 285; 5922 Johnston 44293 Chase 7216; 7954 а” Drouet & 2. 3907 = Edwards Lundell 5095 SENT 133 У Ме 5 &R 2493; 2526 es, M. 2845, 27050 (түре: D. M ida i: aera) 533 кемдік var. с 27051 моћ БР od Roa 7 кү Orcutt 3910, 3915, 3929 McVaugh 16790 Ortenburger, Paxtod & Barkley 16M682; Moore 1471; 3064 16M693; 16M747 = Palmer 253; 26 о 2 4337 Parry & Palmer 5 Palm ы : қ Parry ай Р a in Pringle ee (түре: D. chihuahuensis) Pringle 5089; 7122 Purpus 3310 Purpus 430; 3311 Rose 2789 Hose Мау 3311 Rose & Hay 5510, 5676, 5973 se & Painter 6617; 6775; 7083 (appr. Rose & Painter 7148; 7207 var. galeottiana) 773 Rose, Painter & Rose 8449; 8488; 8773; Rose, Standley & Russell 8882 Rowell, Webster & Barkley 17M470 Rose, Standley & Russell 12796 Schaffner 1 прси 332b " қ Sent, Ме Мос |. 446, 447, 1394? Schumann [did өк, Mocibo et al. 4 Sessé, Mocifio et al. 449; 468; 983 Shreve 9039; 9302 D. GLANDULOSA var. GALEOTTIANA (Briq.) Stewart 2545 . Duke Stewart & ecc с Galeotti 4408 (TYPE); 4418 Wynd & Mueller 6 D. скаспле Cham. & Schlecht. р. орна oe & Schlecht) Barnes & Land 665 = Botteri 349 Barlow 30 Botteri & Sumichrast 1567 Coulter 697 Bourgeau Gentry 8474 Fischer 642, 643 McVaugh 13105 Loesener 138 Palmer 497; 794 Matuda 0905 Parry & Palmer 60 re 2965 Pringle 581; 1194; 6482 Müller 720, 775; 879 Purpus 3734 Rose & Hay 6114 ч 2690 Rose & Hough 4939 & Painter 7132 Smith, J. С. 360 tius 140; 388 (Мог. 48 258 ANNALS OF THE MISSOURI BOTANICAL GARDEN LEPTOPHYLLA var. COGNATA (S. F. Blake) J. Duke Maysilles 7545 Schaffner 542 p. p. D. LEPTOPHYLLA var. NODOSA (Engelm.) J. Duke Gentry 571M; 1814; 2669 (түре: D. gentryi Hartman 138 Knobloch 5522; 5615 LeSueur n 623 (sensu gentryi) Muller 34 Pringle с 716; LM 13506 Rose E Painter 666 Scha Townsend З Barber 201 (sensu gentryi) Tucker White үз 2557; 2733; 4185; 4403; aber.) 4776 ( D. LONGEPEDUNCULATA S. Watson Iz Pringle 2121 irs] po 4348 D. LYROPETALA 1. M. Johnston € е 7513; 7594 (TYPE): 8352; Johnson к Malle 241 arsh 1 aera S6 (TYPE: var. coabuilana) D. MALACHIOIDEs Briq. Leavenworth 595 (?) D. MOULEINEA (Lag.) Didr. Gain D Mearns 2248 Pringle 1192; 1571; 6481 Sessé, Mocifio et al. 984 White 970 D. MuLTIFLORA Brandegee Arséne 5309; 6710 Gentry 6479 Hinton 1549; е = 11538; 12167; 12489; 15050; 1526 McVsugh 12962; 134413 13442; 14066; MeVaush ~. ae 1264 Nelson 1 Purpus yt i (TYP Fin E) he Painter 6517; 6773 p. p.; 6774; тш; 1; 367 Sessé, Mociño et al. 5005 D. PACHYPHYLLA Woot. & Stand! Johnston 7684; 7790; 8009; 8925; 522 x Mur 787; 1108; 1234 Nelson Orcutt í у 5 Pringle Purpus 47 2 be Stewart 1161; 1745; 2956 Waterfall 13281 D. РОГУСАКРОШЕ5 A. Gray Банда 7344; 7 41 poses е Maller 335; 761 Palmer Purpus x x Stewart 814; 845; 1102; 1152; 1753 D. өтірітата Fosberg Stewart 1093 D. suBUMBELLATA I. M. Johnston Johnston 8489 (TYPE) D. surrruticosa A. Gray ех S. Watson Palmer 74 (TYPE Stewart 2813; 2916 D. теміл S. Watson éne ig 6651 112 McVaugh wee Pringle 2120 е qum ч (ТҮРЕ: var. jaliscana) 7648; Schaffner 363; idm Seaton 267 (түре: D. filiform Sessé, Мосійо et 448, 466; pot 5184 D. уплоза Cham. & Schlecht. Arséne 5129 p. p.; 5214; p. р.; 5281; 6810 Bell & Duke 16648 Botteri 350; 944 Botteri & Sumichrast 568 Feddema 1095 Galeotti 1726; 4414 inton 1384; 5152; 9471 Jones, M. E. 22847 (type: D. tepica- n4); 27051a нен D. er ae) we л Hoogstraal 129 Matuda 4 19611 DUKE—REVISION ОБ DRYMARIA 259 пика ris 575a Palmer 616 ` d (both f. townsendii) Orcu 614 Parry & Palm Palmer ш. e ise 1602 Pringle 1501; P Pringle 1196 Rose & Hay Purpus 4440, Hs eg 10229 Schiede & Dep 405 (prob. trans- Rose 2078 numeration of TYPE number) Rose & Painter 6967; 7436 wnsend & Barber 231 (түре: D. Rose, Standley & Russel 14145 & 14386 townsendii) (bo Schiede арор 505 (TYPE) D. VILLOSA ssp. PALUSTRIS var. PERENNIS Duke Seat Sez кез ш & еген 16M833 Edwards ыы жалы: 1568; Johnston Kenoyer de (appr. laxiflora) D. VILLOSA ssp. PALUSTRIS (Cham. & Schlecht.) J. Duke Arséne 16; 5191; Aide: 5416; 5911 Palace г v 7 356 Pringle 3 E XEROPHYLLA A. Gray e 46 Galeotti 4416 (түре: D. nummularia) conce 722 (TYPE) h 793 Hinton 5304; dud 12233 cVaugh 167 ones, M. E. 90 PE: D. subsessilis) Parry & Palmer 50 LeSueur 1322 (f. pesi dii) Pringle 6910 25 Moore 2801; 2955 Sessé, Mociño et al. 5003 Baja CALIFORNIA D. ARENARIOIDES 8р. PENINSULARIS (S. F. D. GRACILIS ssp. CARINATA (Brandegee) J. Duke Blake) J. Duk Carter et al. 2 Brandegee 34 (TYPE) Craig 738 Gentry 4415 (түре: D. carinata var. Dawson 1189 Howell 105 ennis) : м Jones, М. Е. 24193 Johnston 3972 (туре: D. johnstonii) Purpus 216 Purpus 337; 423 (TYPE); 496 Rose 16357 D. ногоѕтеоіреѕ Benth. Bryant 1888 (TYPE: D. veatchii) Carter et al. 1935; 2474 Carter & Kellogg 3089; 3221 ber: Xantus 5 D. ревил5 Brandegee ие 35 (түре: D. отаи ; n. Feb. 13, 1889 UC (TYPE Cham сары 2578 у арене ; 2 Carter & Ferris 3709; 3812; 4093 Gentry 4059; 7516 Carter & у 2922 ѕ. п. (ТҮРЕ) С Қ от & Kempton 157; 211 Johnston p MM 4263 Daw gae. те Е. Gantry А Маѕоп 1908 Palmer 4; М 5 (түре: D. diffusa) Pati: 52% Rose 16609 Pu Wiggins 11409 D. GLANDULOSA Presl Carter et al. 2379; 2447 Carter ai Ferris 3774; 390 Jones, 24149; A 24191; 27050 vrbe D. fendleri var. perennis) urpus 469 Reed 6156; 6255 Rose 16268; 16528 Shreve 7053; 70 Wiggins 5456; 5531; 7898; 11350 D. HOLOSTEOIDES var. CRASSIFOLIA (Benth.) J. Duke Carter et al. 2255 260 Constance 3185 raig inds s.n. (TYPE) Howell 10572 Jones, M. E. 27046 Moran 4168 Xantus 6 D. LEPTOPHYLLA (Cham. & Schlecht.) Fenzl. ex Rohrb. Wiggins & Demaree 4915 Мог. 48 ANNALS OF THE MISSOURI BOTANICAL GARDEN D. viscosa S. Watson Dressler 599 Gentry 7359 Orcutt 51; 1330 (TYPE) 767 Wiggins 7862 CENTRAL AMERICA D. сократа (L.) Willd. Aguilar 193; 978 26 6 Glassman 1805 Grant 766 Mitchell 47 Muenscher 12352; 12475 Oersted 3640; 3649 Orozco 434; 436 Peck 519 Rodriguez, J. V. Rowlee & Rowlee 345; 500 Standley 1720; 2300; 7691; 8588; 8921; 5175451; 19236; 20256; 21557; 22502; 22650; 22714; 23493; 24022; ; 53336; 66994; 72489; 75828; Stork 8; 1065 von Wedel 1249A Watso: Wilson с. Yuncker 4770 О. EFFUSA var. DEPRESSA (Greene) J. Duke Steyermark 50243 (түре: D. minuscula) D. crtANDULOsA Presl Bernoulli 240 (түрк: D. leptoclados) tandley 58053; 59111; 59852; 60343; 60837; 60874; 60895; 61235; 62818; 62897; 63790; 64317 .; 64427; 66095; 66170; 67287; 77390; 80085; 80810; 81211; 81909; 82454; 83002; 83193; 83247; 83512; 83884; 84704; Steyermark 32299; 33991 D. Lapewn Rusby Stanley 58398; 59961; 61978; 65477; 67942; 68554; 80175; 80683; 83275; 83294; 83823; 85147; 85599; 85827; 86254 Steyermark 30047; 32349; 33964; 35653; 36187 D. LAXIFLORA Benth. 4. & Cario 3255 artweg 523 a ud 35768 D. MULTIFLORA Brandegee Bell & Duke 16975 Skutch 2992 Standley 59446; 65172; 79941; 80208 Steyermark 30500 D. упл.о5А Т” & Schlecht. Baker Bell & Duke 16971 Bernou Bree. ve. 17202; 17415 Carlso 91 hn Kellerman 7625; 8013 Molina En Morales ы jo» 36515 3652 289 Rowlee & Коз, 132a Skutch 532; 1961) DUKE— REVISION OF DRYMARIA 261 Standley 1897; 11000; 11873; 12155; 12330; 13284; 13324; 15488; 21136 . p.; 20257; 21730; 21786; 21827; 64283; 64426; 65793; 69598: 71912; 76602; cur 80809; 81050; 82240; 83258; Standley, Allen е al. 919 orb 29615; че е а р. Ра; 5. т. уоп Тана 684 (appr. D. malachi- oides) 20 & Molina 10076; 12016 Vos. Allen & Seibert 1162; 1771 Woods on & Schery 7 Yuncker, Dawson & 2и 5856 D. VILLOSA ssp. PALUSTRIS (Cham. & Schlecht.) J. Duke Muenscher 12428 Skutch Standley 61036; 61054; 61553; 62663; 62855; 62969; 64316; 64697; 64709; 65915; 67816; 82677; 83746; 85203; 85816; Standley & ч fede | 3649 Standley & Jaim Steyermark e. yd р. р.; 35597; 35735; 50126; 50314 WEST INDIES D. сократа (L.) Willd. Baker 1375 Baker & van Hermann 4246 Bolfiagh 1670; 2306 x 985 Britton 2287; Britton & сл Du 12790 Bree & Hazen Britton & Hess ТЕ Britton & Shafer 1702 Britton & Wilson 4958 rown 111 B ant 79 Clement 4088 Cowell 776 Crawford 793 Curtiss 28 Duss 1779; Eggers 436b; 3538; 4972c; 6232a Ekman 612; Sod 203 (түре: Р. nini Fuertes 1323; Hahn 236 Harris чазы 11704; 12760 Heller Heller ~ Није 175; 526A Hodge 438; 439; 440; 2460 Howard & Howard 8440; 8615 Howard & Proctor 13403 Hunnewell 15267; 19453 ma 834 1166 Orcutt 908; 2598; 3208; 3463; 3707; 3721 Otero 41 Palmer 370,371 Proctor 17087 Ricksecker 490 Rose, Fitch & Russell 3492 Shafer "s 208; 523; 13737 Sieber 270; dr w 6487 Smith & Smith 162 Stahl 779 Taylor 202 Tho Wright Wright, Parry & Brummell 2 D. ORTEGIOIDES Griseb. усон ig & Wilson 14279 Curt: Wright pet (TYPE) 262 [Vor. 48 ANNALS OF THE MISSOURI BOTANICAL GARDEN VENEZUELA D. corbata (L.) Willd. 5/5 и e 665 (var. pacifica) Fendler түлей el uery 964 Pittior ta 9827; 11431 Vogl 253; 3 D. FIRMULA Steyermark Steyermark 56536 (TYPE) D. уплоза Cham. & Schlecht. Farenholtz 795 Pittier 12672 D. VILLOSA ssp. PARAMORUM ($. Е. Blake) J. Duke Jahn 111 (TYPE) Pittier 13170 COLOMBIA D. corpata (L.) Willd. André 747 (P. р.) Allen, С Сааты me 12904 Cuatrecasas, Schultes & Smith 12217 Fosberg 19276; 19491, 20367 Ten S Barkley 17С529 Killip 5 Killip & nik 17425 ( aber.) n d. duet Pring 65 Rusby & Pennell 283; 501 Schiefer 625 oro Es Triana s.n. (TYPE: var. puberula) von ышна 2 2256 D. GLANDULOSA Presl Apollinaire 24 D. ovata Humb. & Bonpl. ex Roem. & Schult. André 747 p. p.; 2778; 5865; 6112; Arbelaez & Cuatrecasas 6001 Pennell 7133; 7139; 7621; 8914 Rusby & Pennell 622 D. уплоза Cham. & Schlecht. André 4289 (appr. ssp. palustris) Arbeláez & Cuatrecasas 5 Killip & Smith 16142 p. p.; 17073 Pennell 1735; 3244 Smith, H. H. 556 (appr. ssp. palustris) ; 2850 D. VILLOSA ssp. PALUSTRIS (Cham. & Schlecht.) J. Duke Killip 5358 p. p.; 6732 Killip & Smith 16142 p. p. Rusby & Pennell 644 D. VILLOSA ssp. PARAMORUM (S. F. Blake) J. Duke Fosberg 21491 ECUADOR D. сократа (L.) Willd. reda үз 15170 (var. pacifica) Jameson 171 D. ovata Humb. & Bonpl. ex Roem. & Schult. Benoist De a eus ak 2620 Espino Родена РА 22528 Fosberg & Gila a 23225 (appr. D. villosa ssp. paramoru Fosberg & Prieto 22689 р. р.; 22789 Jameson Penland 46 Rose, Расћапо & Rose 23875 p. p. ose & Rose 22227; 23542 Spruce 6539 Tate 559; 595; 610 Wagner 13 D. ROTUNDIFOLIA var. NITIDA (J. Ball) J. Duke André 3978 (appr. D. glaberrima) D. 5ТЕТТАКТОЕ5 Humb. & Bonpl. ex Roem. & Schult. Camp E-1862 Fosberg 23203 Hitchcock 21406 ano 95 Rose, Pachano = 22952; 23183 p. p. Козе & Козе 2 D. упљоза Cham, & Schlecht. André 198 D. VILLOSA ssp. PALUSTRIS (Cham. & Schlecht.) J. Duke Benoist 77 Camp E-1929; E-2624 Fosberg & Prieto 22689 p. p. Rose, Расћапо & Rose 23183 р. р.; 23875 p. p. 1961] DUKE— REVISION OF DRYMARIA 263 GALAPAGOS IsLANDS О. сократа (L.) Willd. Howell 9158; 9671 Stewart 1513; 1514; 1515; 1516; 1517 Svenson 65 (TYPE: var. pacifica); (var. pacifica) D. APETALA Bartl. : D. virgata) 2823 Weberbaver 5814; 5827 Wi ө d О. AURICULIPETALA Mattf. MacBride & Featherstone 2264 (TYPE) D. corpata (L.) Willd. y 899 Ferreyra 255; 1064; 1758; 11305; 12726; 12753 Hunnewell 15972 nina ДЫ 391 E. dm теі ed D. prvaricata НВК? Pennell 14560 D. DIVARICATA var. DIVERGENS J. Duke 724 Милет 5320, 4189 p. p. D. DIVARICATA var. REFLEXIFLORA J. Duke Ferreyra 3855 p. р.; 3958; 86952; 11522 ufiez 2261; 3579; 3754; 3752; 3755; 3757; 3758; 4198; 4291 D. DIVARICATA var. STRICTA (Rusby) J. Duke Ferreyra 763 (appr. D. glaberrima); Lechler 1947 (type: D. agapatensis) Weberbauer 5327 (appr. D. glaberrima) D. DIVARICATA var. VISCIDULA (A. Gray) J. Duke Ferreyra 6102 MacBride & Featherstone 189? Pennell eg 14516 Stork 11456 websted: 10143 Wilkes Exped. s.n. (TYPE) 134 Exped. s.n. (TYPE: D. macran- D. MONTICOLA Howell Bowman 77 Howell 9243 (түре) D. ROTUNDIFOLIA A. Gra Stewart 1512 (var. ч, 222) РЕКЏ О. ENGLERIANA (Muschl.) Baehni & Мас- Bride Cerrate 1583; Weberbauer ы a D. ENGLERIANA ы DEVIA (Baehni & Мас- Br ie. ты Мас 3 & эе tone 566 Pennell 13462; 14655 (TYPE) Tovar 1277B D. FascicuLata A. Gray Wilkes 5 у 5. п. (ТУРЕ) D. FRUTESCENS Mattf. Weberbauer 7203 (TYPE) D. SUMA UNE Bartl. enke s. Б ae MacBr ide Wortkowiki us (???) D. GRANDIFLORA Bartl. Cerrate 1101 ны s. 8587 (appr. D. glaber- iud 14724 Vargas 1903 D. ovata Humb. & Bonpl. ex Roem. & Schult.) Espinos MacBride & Featherstone 1714 D. PAPOSANA Phil. Nuñez 4189 p. р.; 2430 (appr. var. weber- baueri) D. PAPOSANA Var. WEBERBAUERI (Muschl.) exe Cerrate Ferreyra sem 2435; 3855; 6328 p. p.; 8025; 8695; 9580 Grant 7441,7468 264 м. э, 5904 (aber.) 5963 Miranda Pennell илз Soukup 2111 Stork, Mo gees 9241 Stork fae h & Morrison 15741 (appr. var. paposana) D. PAPOSANA Var. SERRULATA J. Duke Ferreyra 1497; 1515; 7234; 8816; 11616; 13428; 13490; 13522 Matthews 997 Worth & Morrison 15677 D. РКАЕСОХ Baehni & MacBride Weberbauer 6916 (TYPE) D. ROTUNDIFOLIA A. Gra Ferreyra 647; Қы 7048; 8301 Ae : Smith 2 Mac пуче E ds 261 Pennell 14390; 14516 p P4 Wilkes Exped. s. n. (TYPE) D. ROTUNDIFOLIA var. NITIDA (J. Ball) J. me ke E | n. in аа (ТҮРЕ) Pall e 678 са n e ; 1376 Gilbert 615; 638 Fue 3030 (appr. D. glaberrima); 74 8 Killip & Smith 21810 [Уог. 48 ANNALS OF THE MISSOURI BOTANICAL GARDEN MacBride 3506 (appr. D. glaberrima) MacBride & Featherstone 188 (appr. D. data) cor. Nufez 131 Savatier, L. 372 Stafford s.n. (closely resembles Ball s. n. from Chicla) D. sTEREOPHYLLA Mattf. Killip & Smith 22132 MacBride & Рале ‚ыы 6600 (TYPE: var. exstipu- lata) D. уплоза Cham. & Schlecht. Haenke $. п. (TY e = hirsuta) Killip & gee 232 MacBride Mac digi 5 Featherstone 1543 Poeppig Vargas zi D. viLLOSA ssp. PALUSTR (Cham. & ib talcs J. Duke Allard 550 MacBride MacBride E T nna 1542 (TYPE: D. spbagnopbila) Stork & Horton 10479 BorrviA D. corbata (L.) Willd. 233 Bang Buchtien es P p. 255; 5405 ici Rusby Seal E 5442; 6216; 6269 Weddell 4 Williams ig D. DIVARICATA var. STRICTA (Rusby) J. e Tate 160 (TYPE) D. GLANDULOSsA Presl Bang 92 (appr. D. cordata) ; 1136 Buchtien 5 Cárdenas predi 2424 Eyerdam 24941; 24957 Mandon 950 d'Orbigny 1187 Steinbach 4011 D. тарЕУП Rusby Tate 652 (TYPE) р. ek. Humb. & Вопр!. ex Roem. & eee 2332; 3379 Herzog 2390 D. ROTUNDIFOLIA var. NITIDA (J. Ball) J. Duke Mandon 948 (appr. D. ovata) D. уплоза Cham. & Schlecht. Buchtien 183 p. p. Mandon 949 Tate 928 D. VILLOSA ssp. PALU (Cham. & Schlecht.) J. Duke Rusby 1183 1961] DUKE—REVISION OF DRYMARIA 265 CHILE О. РАРОЗАМА Phil. Jaffuel 1025; 1033 (appr. D. rotundi- 7 254 ); Каным 3620; 5213 PARAGUAY AND URUGUAY D. сократа (L.) Willd. Jorgenson 4016 THE GUIANAS D. corpata (L.) Willd. Berthaud-Caulon 538 Broadw Focke Hering = Dieperink 113 Hostmann 714 Maguire & Fanshawe 23562 Mell 232 Persaud 165 Schomburgk 930 Sieber 392 BRAZIL D. CORDATA ud Willd. n 4 Leite 523; 3937 Mosen 409 Muller 249 Netto 124 Pickel 2737 Piers & Black 632 Segadas-Vianna 2642 aen сардар 216 Spru Smith. & А 8799 ARGENTINA D. сократа (L) Willd. ntz & Hieronymus 1123 Parodi 9080 nturi 5104; (var. pacifica); 5911; D. cLANDULOsA Presl Eyerdam & =» 22360 Hieronymus Jorgensen 1172 97 Venturi 3685; 9430 D. ovata Humb. & Bonpl. ex Roem. & Schult. Venturi 2900; 4421 ASIA AND OCEANIA D. сократа (L.) Willd. 32379, 32380 be var. (appr. D. diandra) Bakhuizen 28a (var. pacifica Buchtien 695, dae (var. pacifica) unnemeyer 1107 Cid i 2e pein) Dam Degener 12345 (var. pacifica) Едаћо Dee 11845 (both ма: pacifica) Flecker 3 Heller Peri Hochreutiner 106, 1090 Kakah 53 (var. pacifica) Koorders снаја 16540b; 43293b (all var. pacifica [Vor. 48 266 ANNALS OF THE MISSOURI BOTANICAL GARDEN Meebold 2% i pacifica) Nodi & SUN Nur, Md. Опол ees (var. pacifica appr. ssp. d Posthumus 3459 (var. pacifica) Purseglove 4322 (var. pacifica) Schiffner 1956; 1958; 1960; 1961 (var. pac ee St. John 9 St John & d 17850 Steiner 54787 Words. Ps p^ Royen 5911 (var. aci Womersley, Hoogland & Taylor 4909 (var. pacifica) D. СОКРАТА ssp, DIANDRA (Blume) J. Duke ыг 570 Biswas 4445 Bicecibersen 3371 Bodinier 361 eea 8482; 8653; 8921 Bon 5837 Boswezen 3065; Brass 3784; AR uas Carr I boni 29323 Chun & Tso 44268 Clarke 4547; 35833 Clemens 17642; 41187 Dickason 3123 Elbert 20; 813; 834; 881; 1447; 1675; 9 Forbes 3 Kooders bene 23195b Lau 616; 2118 (appr. ssp. cordata) Loher 1625 Lérzing 12455; 14385 McClure 8422; 9215 McKee 1288 Merrill 992; 4341 Peekel 9 Маи тм. 21; 15604; 25973 Pulle eae So Robinson 18233 Rao 4689 Rock 6419 Savinierre 283 067 Tsang 21658, 25780 Tsang, Tang & Fung 226 Tsiang 922 Van Steenis 6214 Vanoverbergh 1455 Wallich 647 (түрЕ?: D. extensa) Wang T 75334; 77314; 29325. 80900 Wight 108; 152 (туре: ? D. retusa) Wright 28 Yu 16813 Zollinger 892 D. уплоза Cham. & Schlecht. Backer 5240; 5619; 10665; 12940 (TYPE: ? D. stylosa) ; 13632; 15815; 21530; 92 Bakhuizen 6603 Koorders 41287b Lorzing 80a; 1456 Main 2 Meijer 1599 Sapin Schiffner 1952; 24937 Van Ooststrom 13059; 13136 Van Steenis 10916; 11584 79 1961] DUKE——REVISION OF DRYMARIA AFRICA AND MADAGASCAR 267 D. сократа (L.) Willd. Bogden B39 Exell 87 EN 5656; 5761 (aber.) Gille лімен 7640 95% „= ssp. diandra) бесін 2577 Stolz 629 Burchell зе (appr. ssp. cordata) ummer 351 Granvik 7 am ssp. cordata) Hakanson Te 2 eod 6 vicinia 2987; 12858 Stopp 97 оен 192; 674 Schlieben 1839 ; 458 3? Wild 4518 Viguier & Humbet 442 Viguier & Humbert 1236 (appr. ssp. diandra) SYSTEMATIC INDEX D. СОКРАТА ssp. DIANDRA (Blume) J. Duke RUM нај (appr. ssp. cordata) n type indicates accepted, о names; italics indicates synonyms; bold face tu Rom PP ent new names, combinations or statu Alsine molluginea ____________ _ 194 dev 25219 Arenaria leptopbylla .... 208 diandra Ae 251 molluginea __ 194 andra Blum 253 Drymaria adenophora ____________ 251 ren 220 айат ойе; ______ 254 ана 241, 242 agapatensis __ 242 var. diverge 241, 243 попа — 210; 277 ar. reflexiflora 241, 244 crm 232, 234 var. Strict 241, 242 196, 10 var. viscidula 241, 243 soiculipetal 213, 215 Жаза 205, 207 axillaris - 197, 198 var. confusa 205, 207 а : I97, 199 var. depressa 206, 207 barrancae 226 elata 189, 100 blasiana 247 engleriana 217, 218 ME 24 ar. devia 218, 219 r. perennis 247 excisa 202, 203 chibuabuens sis 222 ensa 253 ciliar 226 fasciculata 217, 218 ие — is 226 endleri 248 Сорпа. 20 var. perennis 248 confusa . 205 fenzl е 238 conzattii 224, 225 filiformis Benth. 254 filiformis Robins. & Seaton 2... 211 cordata aes 251, 292 she 231, 232 мр, diandra 252, e frankenioides 96 В онага ча 251 frutescens 213, 216 var. y gracilis 245, 246 ilies 249 f. indica сри 8 туі 209 Var. егілсе ee ese 251 gerontogea 253 уат. В palustris мед ое INST ug oe S 227 laberri .232. 234 glaberrima 3 var. pilosa 226 246, 247 glandulosa 40, а мс м ш 25 var. fendleri 248 vars y villow осы = 226 var. faleottiana 246, 249 CONDE oes со re 187 var. perennis 248 9s. abe xum 246 БЕРИК м. 245, 246 debilis 220, 221 р. carinata 246, 247 206 А сс с. _ 208 depressa (Мог. 48 268 ANNALS OF THE MISSOURI BOTANICAL GARDEN grandiflora Вагі. 232, 236 оныи 247 ни а СЕНЕ 254 retus 255 202 rotundifolia AK ONE 22225, 139 mt 226 var. nitida 277; 239 cec 186, 187 йен Harriot 254 sifolia 187 sessiliflor 254 Пајк тазы Auct. 250 sperguloides 194 hypericifolia Briq. 201, 203 sphagnophila 227 idiopoda 254 stellarioides 412, 213 johnstonii 198 stereophyll I3, 214 ladewii 250, 252 ar. — жыр 213. 215 ra 221; 222 stipit 193, 105 leptoclados 7 ee aris ar. peruviana 7 los. 226 leptophylla 207, 208 subsessili 7 cognat 207, 208 subumbellata 100, 191 var. nodosa 207, 209 suffruticosa 190, 192 longepedunculata _____________ 202, 203 tenella 208 lyropetala 190, 192 var. nodosa __ 209 var. coahuilana 190, 192 tenuis 210, 211 macrantha 234 а var. зенита 211 Аы 22222 254 B var. |а сапа Е malachioides ..... 225 tepicana 226 megalantba 254 townsendii 227 minuscula 206 veatchii 186 molluginea 194, 105 villosa 225, 226 monticola 232, 235 ssp. palustris 227, 229 multiflora ... 224, 225 ssp. рагатогшт 229, 2 nitida 239 var. perennis 228, 220 nod 209 f. tepicana 2 var, gracillima is 208 f. townsendii 228, 229 НИ ос з er osi coe 24 virgata 234 ое. Еки 200, 203 viscidula 243 о 232, 233 viscosa 200, 203 oxalidea 54 weberbaueri 238 pachyphylla 187, 188 xerophylla 249, 250 pelmeri a a Holosieum cordatum 251 papos 3 236, 237 diandrum 251 var. serrulata 237, 238 Key to Series 184 var, weberbauer 237, 238 Key to species ___ 179 palustris 0... 227 Lepigonum mollugin 194 enorm 5 12-230 Mollugophy tm Prat eb сс sf pauciflora |... 227 boloste 186 peninsularis _ 198 sperguloides —_ 194 peruviana _. 254 Pinosia ndn 200 polycarpoides 197, 199 Polycarpon еп епапит ___________- 217 polystachya 220 Stellaria adenophora 251 var. diffusa 220 filiformis 254 praecox 218, 219 rotundifolia 254 procumbens 252 virgata — 234 SANANGO: NEW AMAZONIAN GENUS OF LOGANIACEAE GEORGE $. BUNTING лмо JAMES A. DUKE ABSTRACT The new genus, Sanango, shows affinity to the genera Buddleja, Nuxia and Peltanthera, of the Loganiaceae, сое h the five-lobed corolla ме a nly = stamens is more characteristic of other Vari families as the Scrophulariaceae. arious features of Sanango durum are discussed and реон үсте Е $. BUNTING, І. Н. В Мне: тои Corn i University, Ithaca, New York; JAMES А. DUKE, Ма Botanical Garden, 2315 Tower Grove Ave., St. Louis 10, Мек ri Among recent collections of Felix Woytkowski from Amazonian Peru is a speci- men of a plant that appears to be intermediate between the families Loganiaceae and Scrophulariaceae! It is a fine example of the situation which has led to the wide- spread view that the Loganiaceae are an artificial group constituted primarily of genera excluded for one reason or another from other well-marked, natural families. There is still ample cause to argue this case, for there is no obvious “family resemblance” to guide us, and often the circumscription of the family is inadequate to positively include or exclude an unknown plant. A century ago George Bentham (Jour. Linn. Soc. Bot. 1:52-114. 1854) wrote: “There can be no doubt that Rubiaceae, Apocyneae, Gentianeae, and Scrophulari- neae are large independent orders indicated in nature, yet those genera now amalga- mated under the name of Loganiaceae bind them so firmly together, that some of these genera will be found even more closely allied to certain others of each of the above orders respectively than they are to each other.” And he continues: “The Loganiaceae lie very near to some part or other of the vast family of Rubiaceae, but by their free ovary, they are absolutely, and by very few exceptions, clearly separated. The connecting genera with the Apocyneae, Gentianeae, and Scrophu- larineae are on the other hand, much fewer, but their union is much closer. With Scrophularineae in particular, although the general affinity is more remote, the few intermediate genera and species are intermediate in every respect, in habit as in technical characters. The main distinction, the presence of stipules in Loganiaceae, disappears very gradually, and the difficulty of drawing the line is the greater from there being no habit or family resemblance to unite the several members of the Loganiaceae.” It is in this paper that Bentham discussed the similarity of the genus Buddleja (previously included in Scrophulariaceae) to Logania, and concluded the necessity to place them together, along with the genus Polypremum, in the family Loganiaceae. Now we have discovered another plant that seems to stand nearly between the Loganiaceae and Scrophulariaceae. The fruiting specimen has a distinct aspect of Buddleja, but several technical features prevent its inclusion in that genus. It is equally, if remotely, allied to the genus Peltanthera. After thorough study of this plant, and consultation with N. Y. Sandwith at Kew, and J. J. Wurdack at Wash- ington, we are convinced that it represents an unnamed genus, and describe it as new, Issued December 28, 1961 (269) (Мог. 48 270 ANNALS OF THE MISSOURI BOTANICAL GARDEN Sanango Bunting et Duke, gen. nov. Arbores parvae ligno valde duro, foliis oppositis petiolatis binis lineis conjunctis. Flores in thyrsis bracteolatis, calyce ad medium lobato, segmentis lanceolatis sub- aequalibus, corolla 5-lobata, = hypocrateriformi, gibbosa, segmentis rotundatis subaequalibus, aestivatione cochleari, staminibus 4 (!) sub sinibus prope tubi basim affixis, filamentis inflatis, antheris subrotundis singulatim 2-thecas ferentibus, introrsis versatilibus in linea + hippocrepiformi aperientibus, staminodio parvulo, ovario globoso leviter 4-lobato biloculari disco cupuliformi circumdato placentis axillaribus multiovulatis, stylo brevissimo, stigmate capitato bilobato. Capsula polysperma septicidalis, utroque carpello iterum scisso. Sanango durum Bunting et Duke, sp. nov. Arbor 16 m. alta ramulis lenticellatis. Foliorum lamina 10-25 cm. longa 4-7 cm. lata elliptica vel anguste obovata ad apicem acuminata, ad basim cuneata vel attenuata, parce et inaequaliter denticulata praecipue in dimidio superiore, venis lateralibus utrinque 5-7 arcuatis ascendentibus, infra pallida venis strigulosis instructa; petiolus 0.5-1.0 cm. longus, ad basim latus, facie adaxiali velutina ferruginea. Inflorescentia 5-12 cm. longa terminalis vel axillaris + secunda, pedunculo et pedicellis strigulosis, pedicello ca. 1 mm. longo; florum calyces ca. 3.5 mm. longi puberuli lobis 5 acutis 1.5-2 mm. longis; corollae albae aliquantum carnosae, ca. 1.2 cm. longae extus puberulae et glanduloso-puberulae, intus glanduloso-puberulae et etiam pilosae in tubi basi, lobis ca. 3-4 mm. longis; fila- mentorum staminalium partes liberae pilulosae vix 2.5 mm. longae antheris ca. 0.5 mm. latis brevioribus quam latis; staminodium parvulum vix 0.2 mm. longum; ovarium globosum ca. 1 mm. longum, disco cupuliformi tam procero atque ovario; stylus crassus persistens, una cum stigmate magno ca. 1 mm. longus. Capsula ellipsoidea vel anguste obovoidea ca. 3.5—4.0 mm. lata parietibus osseis; semina orthotropa + elongata fusiformia ca. 0.7 mm. longa recta vel sigmoidea. Type collection: Felix Woytkowski 5610, "tree 12 m. tall; no latex; flowers white; in forest, elev. 300 m.; Aramango, s. of Nazaret,” depto. Amazonas, Peru; 2 April 1960 (HOLOTYPE MO, isotype K, US). Distribution: Forests between 300 and 750 m. elevation, west central Ama- zonas and central Junín, Peru. These localities are separated by nearly 500 air iles. It seems plausible that the distribution may be more or less continuous between these points. Peru: Junín: Satipo, Aug. 1960, Woytkowski 5025 (MO). Vernacular name: "Sanango" (Amazonas). The generic name Sanango is the colloquial name furnished by the olive of this plant. It is a corruption of а Quechuan word, "sanangu", an Indian word applied also to species of Tabernaemontana, Rauwolfia, and Malpighia. The specific epithet refers to the hard wood to which Woytkowski called attention in his field notes. Untreated, the wood disfigures a razor blade when one attempts to cut a hand section. 19611 BUNTING & DUKE—SANANGO 271 ЖУР ыле” t с ез. 5227” со ПД 7 i} age Л. ULES: Z ма Ату Figure 1. Sanango durum Bunting & Duke. а. flowering branch b. flower с. flower with corolla cut open d. stamen from young flower bud, with adaxial view of young anther (center) and anther after dehiscence (top) e. enlargement of pistil, ovary nearly covered by cup-like disk f. diagrams of a single flower bud from opposite sides illustrating aestivation. g. seed [Vor. 48 272 ANNALS OF THE MISSOURI BOTANICAL GAKDEN Sanango belongs to the subfamily Buddlejoideae. Its nearest relatives appear to be Buddleja, Nuxia, and Peltantbera. 1 should be noted that Sanango has been collected only in localities that lie in the middle of the known range of Pel£antbera.! Although the affinity is not strong, these latter two genera share the following characteristics: hippocrepiform dehiscence of the anthers, a disk between the ovary and corolla, multicellular hairs, and a lack of stipules. The combination of characters occurring in Sanango is unusual in Loganiaceae, but most of them occur singly in some other genus of the family. The hippo- crepiform mode of dehiscence of the stamens occurs also in the African genus Nuxia. There is a very well developed annular disk of firm texture in the flower of Sanango. In Peltantbera, a very short disk is present, conspicuous only in young flowers. In Nuxia, however, the disk is annular, and though membranous, very similar in form to that in Sanango.? Notwithstanding, the style and stigma of each of these three genera are extremely different. An approach to the Samango-type pistil occurs in Buddleja, and in the herba- ceous genera Mitreola and Polypremum. In Mitreola, the style is short, the stigma smaller than in Sanango, and the ovary splits in two at the top as it develops. Even more similar is the fruit of Polypremum which splits into four segments at maturity as it does also in Buddleja. Multicellular hairs similar to the dendroid or branched type in Peltanthera have not been observed in Sanango, nor have we encountered the glandular, peltate type similar to those described by Solereder (Pflanzenfam. 4'2):23, fig. 12. 1892) and by Hunziker and Fulvio (op. cit. in footnote). Certain types of multicellular hairs are, however, common in Sanango on both vegetative and floral parts. The corolla is 5-lobed in some genera of Loganiaceae and 4-lobed in others. Indeed both conditions are known to exist in a single species of Buddleja. The loganiaceous flower typically has an equal number of stamens and corolla lobes. Sanango is unique? in the Loganiaceae in possessing a five-lobed corolla with only four stamens, the fifth one merely suggested by a very small appendage not apparent in all flowers. This characteristic is typical of such families as Verbenaceae and Scrophulariaceae. e feel, nonetheless, that Sanango is more closely alied to Loganiaceae than to any other. Such an opinion is dictated by the presence of a stipular line between the leaf bases of each leaf pair, and the strong similarity of the pistil and the fruits to Buddleja, which clearly is related to Logania. The more or less irregular nature of the corolla in aestivation, and the reduction of one stamen in Sanango decidedly suggest Scrophulariaceae. There is, In their excellent paper on Peltanthera, Hunziker and Fulvio (Bol. Acad. Nac. Ci. Cordoba 40: 217-228. 1957) посед that the genus is known from Costa Rica, Peru, and Bolivia. During the € this study we have encountered the collection of A. E. Lawrence (no. 432; 100 mi. Colombi i a y Hunziker and Fulvio for Costa Ric n specimens with the exception of the инее which seem to extend at least half the length of de ovary as is characteristic of the Bolivian specim 2 The ovary of Nuxia floribunda -— $ = ly « eel hich, disk, is very suggestive of сабада and о À pee wake іы v ан € the annular 3 [n the African genus Usteria, three of € gar: jeu байы abort. 1961] BUNTING & DUKE—SANANGO r 273 however, no tendency toward didynamy of the stamens, as is characteristic of the latter group. „ AESTIVATION . Aestivation of the 4 of this genus is TE i.e., one corolla lobe is interior, one exterior, and the other three have one edge overlapping and exposed with the opposite edge hidden by the adjacent overlapping lobe (Fig. 1f). The corolla tube is gibbous near the base, and the flower appears to be oriented so that the interior corolla lobe and the gibbous side of the tube are uppermost (adaxial). The vestigial fifth stamen arises on the gibbous side, below one of the two sinuses adjacent to the interior corolla lobe (Fig. 1c). The bud is distinctly zygomorphic, more so than in Peltanthera. It is impossible to ascertain from the dried material to what degree the open flower may show this character. We are unaware of an identical type of aestivation among the other Loganiaceae, though it is not uncommon among the dicots. The usual pattern in five-lobed corollas of this family is with two lobes external, two internal, and one imbricate. VESTURE At least four types of multicellular hairs occur in Sanango. The largest are present on the adaxial face of the basal part of the petiole. These are composed of 3—5 elongate, hyaline cells, the basal one inflated, the terminal one attenuate (Plate IVa). Very young stems and leaves are clothed with hairs of a similar type, a few of which persist on the veins on the lower surface of the mature leaves. The calyx bears hairs of similar structure, but very much smaller than those described above. In some, however, it appears that the terminal cell is slightly inflated (Plate IVb). Both surfaces of the corolla are very hairy (Plate IVc). The outside is closely covered by very small, mostly 3-celled attenuate hairs similar in form to those of the leaves, and intermixed with these are some still shorter gland-tipped hairs (Plate IVe). The glandular tip cell 15: opaque, borne upon a greatly inflated, hyaline basal cell. а The inside of the corolla bears 3—4-celled, gland-tipped hairs about twice as long as those on the outside of the corolla (Plate IVc). [The glandular tip may be composed of several cells arranged radially, but the small size of the structure has prevented us from ascertaining its exact nature. Some lobing is apparent in certain preparations.] In addition, thére are many very long, slender hairs in the throat of the corolla that appear to have no cross walls dividing them into cells (Plate IVd). ; . ae чине ' à | à PoLLEN A pollen sample was sent to Dr. G. Erdtman for study. He reports that “the grains are small, uninteresting from a morphological point of view, and similar to those in Peltanthera...” His diagnosis follows: “Pollen grains small, 3-colporate (about 15 Х 12.5). Colpi about 12 u long. Ora about 2.5-3 р high. Exine thin. Sexine of about the same thickness as nexine. [No pattern or traces of bacula etc. could be seen even by means of an apochromatic objective with the numerical (Мог. 48 274 ANNALS OF THE MISSOURI BOTANICAL GARDEN aperture 1.40. The pollen grains in Peltanthera costaricensis are equally 3-colporate and measure about 15 X 11 y; the exine seems to be very minutely reticulate.]” ANATOMICAL CONSIDERATIONS Dr. Stern, U. $. National Museum, plans to do a detailed anatomical study of the wood of this genus. We wish, however, to point out that there appears to be no included phloem in this species, which is the typical condition in the Buddlejoideae. This contrasts strongly with the intraxylary phloem present in members of the Loganioideae (Metcalfe and Chalk 2:931. 1950). The wood does not appear to differ strikingly from that of Peltanthera. EXPLANATION OF PLATE PLATE IV Types of trichomes in Sanango from adaxial face of petiole b. on calyx c. on corolla, «рок (left) and interior (ем) [e oS at ery scale] xt nipa of long hair on interior of corolla, a gle s tipped one at its base ( е 2 t lobi f e. idet of a single gland-tipped hair on te the apparent | lla өш | ANN. Mo. Вот. GARD., Vor. 48, 1961 Pirate IV BUNTING & DUKE—SANANGO MISSOURI BOTANICAL GARDEN STAFF Director Fars W. WENT Epcar AND юнч Di D. Оттек, Curator of кегі Plants esearch Associate Henry М. Амрак Palecbotahiat » пена pret Нисн С. CUTLER, ти Dire КЕ 4 NICKERSON, ae У. Dooce, "uso volt сия муља Assistant Cu v Mond» ef | Caraway Н. DobpsoN, Economic Plants Mg ey той =i Curator of OVEN J. SEXTON, Roserr L. DRESSLER OPER л Ты ай мы BUE Сковсь B. VAN SCHAACK, of the ANNALS Curator of Grasses and Librarian James A. DUKE, Assistant Curator практ 2 Е. соци, је. of the Herbarium Curator of Herbarium BOARD OF TRUSTEES | President — Rosrar Brooxine SMITH Vice-President | | Leicester В. Faust Second Vice-President Henry B. РЕТАСЕК Howaro Е. Вак 00 22222 Немаү Нітснсоск Daner К. CATLIN јонм $. LEHMANN AM'L. C. Davis | КОВЕЋТ W. Отто Duprey FRENCH i Warren McK. SHAPLEIGH | : EX-OFFICIO MEMBERS анак | __ Sraarrorp Lee Morro Bishop of the Diocese of President of the Academy of _ Science of St. Louis DANIEL. Ee Beard dt 4 Cart TOLMAN Eie Б President, Board ucation Chancellor o angton ‘of St. Louis ыы к R. TUCKER, Mayor of the City of st. Louis Oscar E. GLAESSNER, Secretary Volume XLVIII Number 4 Annals of the Missouri Botanical Garden NOVEMBER, 1961 | M New Members of the Genus shears idm scu. Bradley жн 275—289 “ЕУ of Plants. А Limited Study of њег “Тсопез” Erna Rice Eisendrath 291-327 A Synopsis of Poinsettia (Euphorbiaceae) Robert + Dressler 329-341 саа ludex to M bises хуш а M 99 LISHED QUARTERLY АТ GALESBURG, ILLINOIS BY iis BOARD OF TRUSTEES OF THE Mene ны BOTANICAL GARDEN ST. · LOUIS, MISSO Entered as ы matter 2 me, etait ні P patrie Тов, under Annals of the Missouri Botanical Garden А quarterly journal containing scientific contributions from the . Missouri Botanical Garden and the Henry Shaw School of Botany of — Washington University in affiliation with the Missouri Botanical Garden. Information _ The ANNALS оғ THE MissOURI BOTANICAL GARDEN appears four times during the calendar year: February, May, September, and November. Four numbers constitute a volume. Beginning with Volume 45, 195%: : Subscription Price _______ $12.00 per volume Single Numbers 300 each 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. | | | 1 | ү | E : Annals of the Missouri Botanical Garden Vol. XLVIII NOVEMBER, 1961 No. 4 TWO NEW MEMBERS OF THE GENUS SCOLECOPTERIS! R. BRADLEY EWART ABSTRACT The genera of Carboniferous Marattiaceae are А with special reference to Scolecopteris, and two new species of 2 аге described: 5. — and S. illinoensis. Scolecopteris monotbrix is ani in e long multicellular hair Sree ne by each sporangium, while S. illinoensis is оса by having и ог six sporangia per synangium, Ағат spores and 22 eee multicellular hairs on the lower Ат ли of the rachis and pinnules. К. BRApLEY Ewart, 228 Ridge Terrace, Park Ridge, "Illinois. INTRODUCTION The genus Scolecopteris, founded by Zenker in 1837, is based upon silicified fructifications from the Lower Permian of Saxony. To date the following species have been described: S. elegans Zenker, Linnaea 11:509. 1837. (A more detailed description was given by Strassburger in 1874.) S. subelegans Grand'Eury, Acad. Sci. Inst. France, Mém. 24:72-73. 1877. S. ripageriensis Grand'Eury, ibid., p. 73. S. minor Hoskins, Bot. Gaz. 82:427-436. 1926. S. oliveri Scott, Journ. Linnean Soc. London, Botany, 49:1-12. 1932, and рр. 309— 1953 S. latifolia Graham, Bot. Gaz. 95:456—458. 1934. Emend Mamay, Ann. Missouri t. Gard. 37:425—428. 1950. S. radforthii Andrews, Ann. Missouri Bot. Gard. 30:435—437. 1947. S. iowensis Mamay, Ann. Missouri Bot. Gard. 37:429—431. 1950. S. minor Hoskins, var. parvifolia Mamay, ibid., pp. 431—432. S. incisifolia Mamay, ibid., pp. 433—436 S. major Mamay, ibid., pp. 436—439. Scolecopteris is characterized by pecopterid pinnules bearing sporangia in sori on the abaxial surfaces. The sporangia composing a sorus are attached to a common pedicel, but are free above, and lack a common synangial sheath. The following account deals with two additional species that fit these charac- teristics and thus have been assigned to the genus Scolecopteris. Further evidence for placing them in this genus will be recorded under the description of each species. 1 An investigation carried out in the graduate laboratory of the Henry Shaw Schoool of Botany of E Washington University and s itted as part of a thesis in partial fulfillment of the requirements for the Шы of Master of Art Issued April 19, 1962. (275) (Мо. 48 276 ANNALS OF THE MISSOURI BOTANICAL GARDEN MATERIALS Both species are based on fertile pinnules found in coal balls that were collected by Dr. Henry N. Andrews from a site two miles northeast of Berryville, Illinois. The outcrop lies in the upper part of the McLeansboro group of the Pennsylvanian of Illinois. THE CARBONIFEROUS MARATTIACEAE Fructifications referable to the Marattiaceae are borne mostly on Pecopteris type foliage. They were borne on the under (abaxial) surface of the leaves, situated over a veinlet or between the veinlet tip and the margin, and unlike the living genera the majority of the fossils have radially symmetrical sori. Mamay (1950) included the following genera in the Carboniferous Maratti- aceae: 1. Cyathotrachus Watson, emend. Mamay: Synangia pedicellate; sporangia enclosed in a continuous synangial sheath; basal portion of synangium with central column; apical portion hollow and cup-like. Based on petrifactions. 2. Ptychocarpus Weiss: Synangia pedicellate; sporangia enclosed in a continu- ous synangial sheath, and fused to a central column extending through the entire length of the synangium. Based mainly on compressions. 3. Scolecopteris Zenker: Synangia pedicellate; sporangia attached to a central receptacle but free distally. Based mainly on petrifactions. 4. Sturiella Weiss: Synangia pedicellate; sporangia basally fused but free above; annulus present. Based mainly on petrifactions. 5. Acitheca Schimper: Synangia sessile; sporangia attached at their bases to a central column, with distal portions free; sporangial apices long and bristle-like. Based mainly on petrifactions. 6. Asterotheca Presl: Synangia sessile; sporangia attached to a receptacle, but free distally, their long axes parallel to the plane of the pinnule. Based mainly on compressions. 7. Eoangiopteris Mamay: Sori linear; sporangia arranged in two rows along fleshy receptacle; sori located along lateral veins; sporangia free. Monotypic. Based on petrifactions. us, the accepted criteria for separating the fossil genera are the following: 1. Sori linear or radial. 2. Synangia pedicellate or sessile. 3. Sporangia free above or enclosed in a continuous synangial sheath. 4. Central column present or absent in synangia. (It is important whether the central column extends through the full length of the synangium or not, as in distinguishing Cyathotrachus from Ptychocar pus.) Since the discovery of the pteridosperms, some paleobotanists have regarded certain of the supposed marattiaceous fructifications as more correctly referable to Pteridospermae. Kidston (1925) expressed this view in finding resemblances be- tween Acitheca and Telangium. However, Mamay (1950) concluded that upon the basis of the sizes of fructifications and spores, the arrangement of sporangia in sori, and the superficial position of sori, this fossil group more closely resembles the living Marattiaceae than the pteridosperms. 1961] EWART—TWO NEW MEMBERS OF SCOLECOPTERIS 277 Mamay (1950) considered the coenopterid fructification Chorionopteris to best fill the qualifications of a progenitor of the Marattiaceae. This evolution supposedly came about by means of a “phyletic slide” from marginal or terminal sori to superficial ones. He further proposed that those fossil genera with fused sporangia are most closely allied to the living genera Danaea and Christensenia, while those with free sporangia are most closely allied to the living genera, Angiopteris, Arch- angiopteris, and Macroglossum. DESCRIPTION OF New SPECIES ЗСОТЕСОРТЕВ!$ monothrix К. B. Ewart, sp. nov. (figs. 1-12). This fructification is distinguished by long slender sporangia each of which terminates in a single, unbranched, multicellular hair. These were borne on pecopterid pinnules with strongly inflexed and deeply incised margins in synangia consisting of four, sometimes five, or rarely three sporangia. One well-preserved specimen was discovered in coal ball WCB 1203. It con- sists of an ultimate rachis with about 15 attached pinnules. Several detached pinnules and synangia were found in the same coal ball. A second intact specimen was found in WCB 1238 along with more fragments. Nearly all synangia were found attached to pinnules and seem to have been preserved in their natural position. The morphology of the ultimate rachis and attached pinnules may be seen in the reconstruction in text-fig. 1. The average dimension of the pinnules and penultimate mid-rib are as follows: Pinnule length (normal) 6.0 mm. Pinnule length if unrolled 12.0 mm Pinnule width (normal) 2.5 mm Pinnule width if unrolled 10.5 mm Penultimate midrib diameter 0.85 mm. The penultimate or rachis midrib (fig. 7) shows no cellular detail except for the vascular supply which consists of a U-shaped strand with the concavity adaxial. The basal part is two or three tracheids in width while the arms narrow to one tracheid. Short multicellular hairs are found on the lower side. Figure 11 (A) shows some of these hairs on the longitudinally sectioned midrib. Some of them are as long as the diameter of the midrib. Figure 7 shows that these hairs are several cells in width and are irregularly spaced in contrast to the hairs found in the sporangia. As seen in fig. 11, the pinnules are joined laterally where attached to the rachis but become separate a short distance out from the rachis. They are of the pecopterid outline characteristic of the genus. The margins are so strongly flexed downward that they parallel and finally almost enclose the synangia as may be seen in fig. 10 (B). The pinnule is devoid of synangia along the distal third of its length and this portion is strongly flexed downward and inward toward the rachis midrib almost completely enclosing the synangia and the space directly below them. This space appears to be loosely filled with sporangial hairs which are oriented in the direction of the midrib. These hairs may be seen, mostly in transverse section, in [Vor. 48 278 ANNALS OF THE MISSOURI BOTANICAL GARDEN fig. 10 (A). The pinnule margins are deeply incised and may be compared in structure to those of $. latifolia and S. incisifolia. Figure 12 (B) shows one of the marginal lobes in vertical section. A row of these lobes in transverse section may be seen in figs. 8 and 11 (B). In fig. 8 the cellular contents of these lobes may be seen. The lobes join laterally at about the level of the sporangial apices, but their vascular strands continue as discrete bundles. The only other cellular struc- ture of the lamina that is preserved is a layer of hypodermal cells below the upper epidermis; it is one or two cells in thickness and may be seen at the top of fig. 4 and in figs. 10, 11, and 12. The synangia form a single row of nine or ten sori on either side of the midrib of each pinnule. A synangium usually contains four sporangia, but those with five, such as the one in fig. 9, are also common and one has been found with three. The sporangia composing a synangium are arranged radially on a common pedicel. Due to the long slender nature of the synangia, it was not possible to find a good median longi- tudinal section showing both the pedicellate attachment and the apex bearing a portion of a sporangial hair. Thus, fig. 4 was prepared from three photomicro- graphs showing these features in their proper proportions: the upper shows the pedicellate attachment and the proximal half of two sporangia; the middle shows the distal half of two sporangia and their apices; and the third shows simply the basal portion of a hair characteristic of the species. The sporangia are long, slender and roughly cylindrical but are slightly wider near the base than at the apex; they are tightly appressed to one another and the synangia themselves are quite crowded. The apex of each sporangium, prior to dehiscence, is curved inward toward the center of the synangium and fits closely against the apices of the other sporangia (figs. 2, 4). Thus a central synangial cavity is formed which is completely enclosed. Dehiscence was accomplished by a longitudinal slit along the inner face of the sporangium (fig. 1); in fig. 8 a mass of spores may be seen discharging through the slit. This leads to the question of how the spores escaped from the enclosed synangial cavity. Figure 6 shows a synangium of S. monothrix at a later stage and is from a different specimen than that shown in most of the figures. The preservation is poor but it may be seen here that the sporangia have separated except where joined to the pedicel. This synangium, along with several empty ones, was attached to a fragment of a pinnule. In all probability the strongly curved pinnules to which these fructifications are attached straightened somewhat at maturity, decreasing the crowding of the synangia, and allowing the component sporangia to separate. In figs. 3 and 4 it may be seen that a single, long multicellular hair is attached on the outer periphery of a sporangium near the apex. It is likely that such a hair grew on most if not all sporangia of S. monothrix, but no sporangium has been observed to have more than one hair. No branching has been observed in these hairs nor has any deviation in position or structure been noted. In fig. 3, the “bulb” at the distal end of the hair is merely the point at which the hair leaves the plane of the section and is not the termination of the hair; the total length is much greater than is shown. By following the course of a cluster of hairs it 1961] EWART—TWO NEW MEMBERS OF SCOLECOPTERIS 279 Text-figure 1. Reconstruction showing abaxial view of part of a pinna of Scolecopteris mono- thrix. Portions of some pinnules are cut away to reveal arrangement of the fructifications. X 9. is possible to estimate their length. The hairs in such a cluster all tend to point in one direction, toward the rachis midrib. In fig. 10 (A), a cluster of hairs may be seen in transverse section in the space below the sporangia. In fig. 12, clusters of hairs may be seen on either side of and pointing toward the rachis midrib (A) where they terminate. It is almost certain that these hairs originate on the sporan- gia, as no trace of an attachment has been observed elsewhere. It is thus concluded that the sporangial hairs are 2 or 5 times the length of the sporangia or 4 to 6 mm. in length. The average dimensions of a synangium are as follows: Total length of synangium (exclusive of hairs) 2.0 mm. Total length of sporangium (exclusive of hair) 1.8 mm. Maximum diameter of sporangium 0.35 mm. Diameter of pedice 0.35 mm. The spores as shown іп fig. 5 are monolete and bilateral with a length of 12 п and a width of 9 д. The smooth walls appear to be thick, but lack any markings of the exine, except where obviously due to damage. The only member of the genus with smaller spores 15 S. elegans with spores 10 р in diameter. Using the method described by Radforth (1939), the average number of spores per sporan- gium is estimated at 9,000. From the above description it may appear that S. monothrix has some rather striking characteristics for this genus. Some thought was given to the possibility of establishing a new genus bur it was decided that this would not be necessary. The following are the reasons for placing this species in the genus Scolecopteris: 1. It seems most appropriate to refer it to the Marattiaceae on the basis of the nonindusial sori borne on the abaxial surface of pecopterid leaves. 2. The pedicellate, radial synangia limit it to Scolecopteris, Ptychocarpus, or Cyathotrachus. [Vor. 48 280 ANNALS OF THE MISSOURI BOTANICAL GARDEN 3. Lack of a continuous synangial sheath and central column limit it to Scolecopteris. . It is borne on pinnules with a morphological structure similar to those of S. latifolia and S. incisifolia and is believed to be closely related to these species. 5. Although it has a closed synangial cavity at maturity, this opens at a later stage causing the synangia to assume a more scolecopterid appearance. Other members of the genus, such as $. incisifolia, also have closely appressed sporangia. 6. Although hairs of the S. monotbrix type are unique for the genus, S. major is known to have multicellular hairs on the sporangia. 7. Although S. monothrix has small spores and large sporangia, they do not represent the extremes of the genus. As mentioned above, S. elegans has smaller spores, while S. major has sporangia from 0.3 mm. to 1.1 mm. longer than those of S. monotbrix Scolecopteris monotbrix may be differentiated from other members of the genus by the following characteristics: 1. No other member of the genus has a single, long, multicellular hair in a fixed position on the outer periphery of the apex of the sporangium. The synangial hairs of S. major are quite different. 2. S. oliveri and S. incisifolia are the only other species with bilateral spores. Of these two, only S. incisifolia compares to S. monotbrix in having a smooth exine but the spores of the former аге 22 X 14 и while those of the latter are only 12 СЫ 3. In sporangial length, 5. monothrix is intermediate between 5. major, which is 0.3 mm. to 1.1 mm. longer, and S. oliveri, which is 0.48 mm. to 0.40 mm. shorter. However, both of these species have a greater sporangial width than $. monotbrix. The ratio of sporangial length to width is greater іп $. monthrix than in any other member of the genus, being about 6:1. 4. The blunt connivent apices of 5. monothrix further distinguish it from S. major, which has very acute apices. The specific name $. monothrix is proposed in reference to the unique sporangial hairs. Specific diagnosis: Synangia consisting of four, sometimes five, rarely three ‘sporangia basally attached to a common pedicel. Sporangia 1.8 mm. in length and 0.35 mm. in width; apices blunt, connivent in synangia but separating after dehiscence; dehiscence by longitudinal clefts along inner sporangial wall. A single, unbranched, multicellular hair one cell in diameter located on the outer periphery of the apex of each sporangium, reaching a length two or three times the length of a sporangium. Pinnules averaging 6 mm. in length and 2.5 mm. in width (exclusive of inflexed margins), margins strongly inflexed and deeply incised, ultimate rachis with U-shaped vascular strand an ick multicellular hairs aver- aging 0.85 mm. in length. Spores monolete, averaging 12 м long and 9 y wide, with thick walls and smooth exines. Age: McLeansboro Group, Upper Pennsylvanian. Locality: Berryville, Illinois. Holotype slides: Slides 2987-2995 (from WCB 1203). 1961] EWART—TWO NEW MEMBERS OF SCOLECOPTERIS 281 ScoLecopTERis illinoensis К. B. Ewart, sp. nov. (figs. 13 and 15-26). This description is based upon specimens from WCB 1204 and WCB 1239, particularly the latter from which the type slides were chosen. The fructifications in these coal balls were found to be in an excellent state of preservation. They were found attached to pinnules which were in turn attached to pinnae midribs. By cutting WCB 1239 in two planes, it was possible to obtain peels which revealed the morphology and something of the anatomy of the specimens. Five or six sporangia are radially grouped on a short common pedicel to form а synangium. Of the 14 synangia seen in fig. 16, four contain six sporangia, seven contain five, while it is difficult to determine the numbers of the other three. Examination of other synangia has shown that those synangia with six sporangia are almost as numerous as those with five. As may be seen in fig. 16, the synangia form a single row on either side of the pinnule midrib. There are generally six or seven synangia per row. A vascular bundle branching from the pinnule midrib is found at the base of each synangium as seen in fig. 23 (C). The outer sporangial walls are thick and curved as seen in fig. 16, but are com- posed of but one layer of cells. The lateral walls are thinner and straight, but do not touch one another, except in synangia which, like the one at the bottom left of fig. 16, contain spores and are probably less mature than the others. It is likely that the sporangia in a synangium separate as they mature, like those in $. mono- thrix. Dehiscence occurs by means of a longitudinal slit along the inner wall of each sporangium. Most of the sporangia seen in this study had already dehisced but a few may be seen to contain spores (fig. 22). The sporangia are roughly ovate in vertical section with acute apices that curve inward toward the center of the synangium as seen in fig. 23 (B). The sporangial cavity does not extend into the distal part of the apex. Figure 13 shows a transverse section through the apices of a synangium. The average dimensions of the fructifications are as follows: Total length of synangium 0.66 mm. Total length of sporangium 0.65 mm. Maximum diameter of sporangium 0.31 mm. Diameter of pedicel 0.16 mm. The spores are spherical in shape and have a crescent-shaped indentation in the exine at one point as indicated by the arrow in fig. 18. Examination of well pre- served spores leads to the conclusion that this may be regarded as a natural and constant character. The diameter of the spores averages 15 p. The penultimate midrib may be seen in transverse section in fig. 23 (A). The vascular supply consists of a U-shaped line of tracheids, and extending from the free end on the right hand side is a small bundle of tracheids representing a pinnule trace just departing. Figure 24 represents an oblique longitudinal section of the vascular supply in an ultimate pinnule. The arrow indicates a branch of this trace. Following the trace distally, it may be seen that this is one of six such branches each of which passes into the pedicel of a synangium. One of these traces may be seen in trans- verse section in fig. 23 (C). [Vor. 48 282 ANNALS OF THE MISSOURI BOTANICAL GARDEN The pecopterid pinnules are alternately spaced along the rachis midrib as may be seen in fig. 15. e pinnule margins are minutely toothed (fig. 17) and reflexed downward (figs. 19-22). Following are the average dimensions of the rachis midribs and pinnules in these specimens: Pinnule length 5.5 mm. Pinnule width 2.2 mm. Rachis midrib diameter 1.0 mm. Figure 19 shows a row of pinnules, each with a pair of sori, arranged along a rachis midrib and sectioned transversely. At the extreme left is the rachis midrib in oblique section. The anatomy of the pinnules is well shown in fig. 20. The upper epidermis is not preserved, but the underlying, large hypodermal cells (A), form a layer one or two cells thick. Below this layer is a zone of palisade parenchyma (B), consisting of one or two layers of narrow, tightly-packed, elongated cells. Underlying this is the spongy parenchyma (C) of loosely-massed rounded cells, and numerous air chambers. Bounding this is the lower epidermis (D). A notable feature of Scolecopteris illinoensis is the abundance of hairs on the lower surfaces of the pinnule midribs (figs. 24, 26). They are multicellular, un- branched, and one cell in diameter. Figure 21 shows that they extend beyond the apices of the sporangia. The hairs in transverse section may be seen between the two rows of synangia in fig. 16. Similar hairs have been observed attached to the synangial pedicel. In fig. 22, the arrow indicates a small outgrowth of the sporangial wall which may be the base of a hair, but these have not been seen on other sporangia. S. illinoensis clearly belongs in this genus but differs from other members of the genus as follows: 1. The only other species in which it is not rare to find six sporangia in a synangium is S. jowensis. S. minor rarely has six, but a comparison of that species in fig. 14 with 5. illinoensis shows a distinct difference in numbers of sporangia. 2. This species differs from S. iowensis which has spores measuring 65-80 шіп diameter, four times the diameter of those in S. illinoensis. $. iowensis sometimes has a double series of synangia on each side of the pinnule midrib, while this has not been observed in the present species. 3. Although multicellular hairs are sparingly present on the lower side of rachis and pinnules of S. minor, no species has hairs on the pinnules that are as conspicuous as in S. illinoensis. Specific diagnosis: Synangia consisting of five or six sporangia basally attached to a common short pedicel. Sporangia 0.65 mm. in length and 0.31 mm. in width; apices acute and incurved; dehiscence by longitudinal cleft along the inner sporan- gial walls. Pinnules averaging 5.5 mm. in length and 2.2 mm. in width, with inflexed minutely toothed margins. Ultimate rachis with U-shaped vascular strand. Unbranched, multicellular hairs, one cell in diameter and reaching 0.7 mm. in length produced on pinnule midribs and synangial pedicels. Spores spherical, 15 и in diameter with crescent-shaped mark on exines. 1961] EWART—TWO NEW MEMBERS OF SCOLECOPTERIS 283 Age: McLeansboro и 2e Pennsylvanian. Locality: Berryville, Illino Holotype slides: Slides nij 3005 (from WCB 1239). SUMMARY Two new species of Scolecopteris are recorded both of which were found in the Upper Pennsylvanian of Illinois. S. monothrix is distinguished by its long sporangia each of which has a distally attached long, multicellular hair. The tips of the com- ponent sporangia were tightly appressed when immature. The spores are very small and their number per sporangium high. S. illinoensis is characterized by having five or six sporangia per synangium, small spores, and conspicuous multicellular hairs on the underside of rachis and pinnules. ACKNOWLEDGMENT This study was carried out in the laboratories of Washington University under the guidance of Dr. Henry N. Andrews, to whom the writer wishes to express his sincere thanks; it is a part of a paleobotanical research program aided largely by the National Science Foundation. Thanks are also extended to Dr. Tom L. Phillips for assistance with paleobotanical techniques and for aid in other ways. LITERATURE CITED Andrews, H. N. (1943). Contributions to our БЕА edge of American Carboniferous floras. VI. о 2 Bower, Е. О. (1926). The ferns (Filicales), vol. "i 344 pp. Cathe idge University Pre: Graham, К. (1934). Pennsylvanian flora of Illinois as revealed in coal balls, І. Bot. сга 95:453- Grand'Eury, F. C. (1877). Flore vut du Départment de la Loire et du centre de la France. cad. sci. inst. France Mém., 24:1 Heine: J. H. (1926). Structure of и. plants from Illinois. Bot. сены 82:427-436. Kidston, К. (1925). Fossil plants of tee Carboniferous rocks of Great Britain. . Geol. Survey Great Brit., Palaeontology II(6):5 Mamay, S. H. (1950). Some hoes Ciiboniferóné fern fructifications. Ann. Missouri Bot. 76 Radforth, N. W. (1939) . Further contributions to our p of the fossil Schizaeaceae. "Trans. I Roy. S $ Scott, D. H. (1932 On a Scolecopteris (S. Oliveri, s n.) from = Perino- Carboniferous of utun. I. The fructification. London Journ., Botany 4 _ кє D. 2 г Н. 5. Holden пети "On tee Oliveri. II. The v vegetative organs. Ibid. Senor t TE Über есен". elegans, pss Jen. Zeitschr. Naturwiss. 8: Zenker (1837). poe edid elegans Zenk. Ein neues fossiles Farrngewachs mit Fructifica- Linnaea 11:50 (Мог. 48 284 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE V. Scolecopteris monotbrix Fig Transverse section of a synangium sectioned in the plane 1-1 of fig. 4. Note spores раак into synangial cavity. Slide 2994. Х 85. Transverse section of apex of synangium sectioned in the plane 2-2 of fig. Note that all four ен fit closely together. A te the E celled base of a seen. Slide 2 Fig. 3. 2. section of sporangial apex showing hair. jet gee at distal end is x point at which the hair leaves the plane of the peel. Slide 2988. Fig gitudinal sections of synangium and synangial hair ное character- istic folk an 4 өшеді attachment to the pinnule. Three separate sections from slides 2988, 2989, and 2990. 50. Fig. 5. Spores. Slide 2987. Х 2250. ANN. Mo. Вот. Garb., Vor. 48, 1961 PLATE У EWART—TWO NEW MEMBERS OF SCOLECOPTERIS ANN. Мо. Bor. GARD., VOL. 48, 1961 PLATE VI м” ames ж E у» g 08 EWART—TWO NEW MEMBERS OF SCOLECOPTERIS 1961] EWART—TWO NEW MEMBERS OF SCOLECOPTERIS 285 EXPLANATION OF PLATE PLATE VI Scolecopteris monothrix Fig. 6. Longitudinal section of open synangium of more mature specimen. Slide 2992. Fig. 7. Transverse Mp oe midrib of rachis. Note U-shaped vascular strand Ка) and hairs. Slide X 55. Fig. 8. Transverse section E rough synangium, sporangial hairs, "S n та margin. Note spore mass passing into synangial cavity (arrow). Slide 2 Fig. 9. Transverse section of а synangium consisting of five sporangia. ся 2994. (Мог. 48 286 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE VII. Scolecopteris monothrix 10. Transverse section of pinnules in sores 10-10 of fig. 11. Note sporangial E (А) me space below synangia. Slide 29 713. ction of pinnules in plane of а a dure on rachis midrib (A) and transverse section of pinnule margins (B). Slide 2993. Section at right angle to rachis midrib in de 12-12 of гас k Note rachis midrib (A) and strongly inflexed pinnule margin (B). Slide 2991. ANN. Мо. Вот. Garp., Vor. 48, 1961 Рі ҮП ATE EWART—TWO NEW MEMBERS OF SCOLECOPTERIS Pratre УШ ANN. Мо. Вот. Garp., VoL. 48, 1961 F SCOLECOPTERIS MEMBERS O УУ МЕ EWART—TWO — Ham 1961] EWART—TWO NEW MEMBERS OF SCOLECOPTERIS 287 EXPLANATION OF PLATE PLATE ҮНІ Figs. 13, 15-17. Scolecopteris eee Fig. 14. S. minor Hoski Fig. 13. Transverse section of a synangium in the region near the apex. Slide 3004. 2355 Fig. 14. Transverse section of а row of synangia. Compare with 5. illinoensis in fig. 16. Slide 3004. X 30. Fig. 15. Longitudinal section of rachis midrib with attached fertile pinnules. Slide $005.03 5, Fig. 16 Single ee 25 0: 15 enlarged: Note hairs іп transverse section between rows о synangia. Slide x35 Fig. 17. Section though an margin showing toothed nature of the margin (arrows). Slide 2998. X 35 [Уот.. 48 288 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE IX. Scolecopteris illinoensis Fig. 18. Spores. Note crescent-shaped mark on exine. Slide 2996. X 1033. 219: Ориат. transverse section of fertile pinnules. Note rachis midrib at extreme left. Slide 300 6 Fig. 20. Transverse section of half of a pinnule showing anatomical detail. A. Hypo- dermal cells Bia € is poorly preserved); В. Palisade parenchyma; С. Spongy nchyma; D. Lower epidermis; E. Pinnule midrib. Slide 3000. 62. т 21. Transverse section of a fertile pinnule. Note long hairs on midrib. Slide ЗОО X54. Fig. 22. Transverse section of a fertile аа Note hair оп sporangium and sporangium with spore mass, Slide 3001. x 5 ANN. Mo. Вот. Сакр., Vor. 48, 1961 Pirate ІХ EWART—TWO NEW MEMBERS OF SCOLECOPTERIS ANN. Mo. Bor. GARD., Vor. 48, 1961 PLATE X EWART—Two NEW ME MBERS OF SCOLECOPTERIS 1961] EWART—TWO NEW MEMBERS OF SCOLECOPTERIS 289 EXPLANATION OF PLATE PLATE X. Scolecopteris illinoensis {зил section МЕ kae Put ki pinnule. A. Transverse section of Бір; 23. pinna midrib. B. Inc apex C. Vascular bundle at base of synangial pedicel. D. Spore mass in а. “slide 2997. E Fig. 24. Longitudinal section of pinnule midrib. Note vascular strands branching from midrib and hairs on midrib. Slide 2997. Fig. 25. Section showing pinna midrib with pinnules on either side. Slide 3003. X 11. Fig. 26. Hairs on pinnule midrib. Slide 2999. X 90. PORTRAITS OF PLANTS. A LIMITED STUDY OF THE “ICONES”! ? ERNA RICE EISENDRATH ABSTRACT A set of four — for defining the word "icones" is established: that the illustrations be full page in si né that y include some portrayal of the habit of growth of the plant; that they be botanically secure "Vat they give aesthetic satisfactio Each of the "icones" must fulfill each requireme ree, except lant portrayals of the 16th and 17th century herb cept in the plan The history of куз cuts is traced throughout their eredi use, and their exception to the e li i i general rule discussed in the light of their great botani a our genera of the Cucurbitaceae (Fevillea, Cucurbi v. Lage е ) are chosen as limiting factors in the study, and illustrations of these plants Кыша: їп ба. Г. ibrar of the Missouri Botanical Garden discussed in relation to the criteria previously set up. The uo dy is concluded with a chronological index of “i of these key species. ems Rice ErENDRATH, Henry Shaw School of Botany, Washington University, St. Louis 30, Missour BOTANICAL USE or THE WorD “ICONES” The word “icones” was taken into botanical parlance by Otto Brunfels in 1530. Its spelling in the title of his Herbarum vivae eicones still conveys a hint of orientalism just strong enough to tie in with its common meaning today; to the layman, an icon is (as it was, to the publishers of Webster’s New International Dictionary of 1918) “а religious image in the Eastern church”. For botanists, however, it is not the gilt on these distorted saints that glows when they speak reverently of “icones”. Their гы is evoked by а kind of floral illustration which they пасо tarip as “icones” in special sections of libraries; to whic they readily fete as "icones" in conversation; of which they will confidently name names as true "icones"; but for which they hesitate to pronounce a definition. So, though everybody knows the special meaning of the word, there remains the prob- lem of establishing criteria by which to implement its definition; a later problem will be to list the books containing illustrations that satisfy the criteria. To start from an authoritative beginning I give you Benjamin Daydon Jackson, who in 1900 first published in London his most useful Glossary of Botanic Terms. To him, “icones”? are merely “pictorial representations of plants; botanic figures". In this sense which includes all botanical illustrations it was certainly used by Pritzel* in his Iconum Botanicarum of 1855 and is also so used in the Latin title page of the Index Londinensis. For the purposes of this paper, however, I prefer basing my criteria on the last word of the last definition of "icon" given in the Oxford English Dictionary (Oxford, 1933): "esp. applied to 'figures' of animals, plants, etc. in books of Natural History. Obs." The use of “icon” to include all natural history illustra- tions is now obsolete; when botanists speak today of the "icones" it is with a of Wing University and sinit етае татам нра ee for the degree of Ma Ar n express her ертн for their = in preparing this manuscript to Dr. 2 белинен E ler and Dr. George Van S s And "De Hash Curie: an Sc У Jackson ОЕ В is given for the acr giia more ых used by botanists than the “The task which Pritzel set himself in the eager of his Index was to record in the simplest f the Phanerogams and Ferns, excluding, with certain exceptions, those o C E t tes as ЊЕ Editor of а py ex Londinensis, in Introduction to the 1929 edition. Issued April 19, 1962. (291 ) (Мог. 48 292 ANNALS OF THE MISSOURI BOTANICAL GARDEN different connotation, not specifically stated in any source that I can find. It implies more than scientifically accurate illustrations, and more than attractive plant portrayal, falling some place between the two and partaking of both but, with a kind of hybrid vigor, better than either. Dr. Hugh Cutler bravely ventured to make the following brief but telling statement. ‘The ‘icones’,” he said, “may have nothing to do with the church exactly, but a great many botanists worship them.” That is, apparently, about as far as educated angels dare to tread; this fool will now step in to define "icones" by establishing the criteria which distinguish them from other illustrations in botanical books. CRITERIA DISTINGUISHING “ICONES” After examining every lead to illustrations of the key® genera of Cucurbi- taceae available at the Missouri Botanical Garden library (some 157 volumes), I have set up four arbitrary criteria for “icones”. The first is that the illustrations be full page® in size; the second, that they include portrayal of the habit of growth of the plant. The third is botanical accuracy, and the fourth artistic presentation. In many instances my first criterion is necessitated in order to eliminate illustra- tions used simply as elaborations of a text. According to Nissen (1951), “‘the documental botanical illustration should, for the co-investigator, replace the plant itself, be he distant in time or рјасе“. So, too, must true "icones" be visual portraits of plants, dependent in no sense upon the written word. Р. J. Е. Turpin, the great French botanical artist, quite naturally believed that the brush was greater than the pen as а naturalist’s tool, and wrote that he “who possesses only the latter loses perhaps the more significant” (Turpin, 1820). But the brush used to depict the plant must do so in considerable detail, and the portrait must be extremely accurate—so accurate as to compete with the botanist’s dried herbarium specimen as a source material for study. (The possibility of this is recognized in the official acceptance, though rare, of such representations as type specimens.) Small figures of plants cannot possibly fulfill all of these requirements; hence the first criterion for true "icones": that they be full page illustrations. My second criterion for defining true "icones" is that they represent the habit of plants, and include both vegetative and reproductive organs. It is possible to object that this is unnecessary, since one can assume that generic characters are "given", and that only identifying specific characters need be depicted in plant portraiture. This objection I liken to a statement that a man’s fingerprints identify him; certainly they will differentiate him from all other men, as certain characters of a species of plants will also differentiate it. Neither fingerprints nor selected characters, however, will give us portraits of the individual or the species, and, in defining today's usage of the word "icones", I insist that portrait is the proper synonym (if any is truly proper). "Icones", then, must portray habit of growth, and here is another example of the need to pin a word to exact definition. What is "habit"? Webster (New : See p. 296. a many 16th and 17th century herbals this criterion must be ignored for reasons explained later in the text. 7 Translated by Mr. Alfred Sadler. 1961] EISENDRATH—A STUDY OF THE ICONES 293 Intern. Dict., 1918) gives as its eighth definition of the word, “Nat. Hist. Char- acteristic form or mode of occurrence or growth; as, elms have a spreading habit.” ackson, deriving “habit” from the Latin, gives, “Тһе general appearance of a plant, whether erect, prostrate, climbing, etc.” Sir Joseph Dalton Hooker, the great systematist, Director of Kew Gardens, and editor of volumes 91-130 of Curtis’ Botanical Magazine, wrote, "Habit of a plant, of a species, a genus, etc., consists of such general characters as strike the eye at first sight, such as size, colour, ramifica- tion, arrangement of the leaves, inflorescence, etc., and are chiefly derived from the organs of vegetation” (Hooker, 1875-97). To this I append the requirement that either flower or fruit must be depicted; however, it is not always easy to decide where “habit” is illustrated and where it is not. I have, for instance, not listed among the “icones” pertinent plates (see plate ХІ) іп Tournefort’s Institutiones rei herbariae (1700), even though they are cited by Linnaeus (Species Plantarum, Stockholm, 1753, vol. 2, рр. 1010-1011) in his original description of the genus Cucurbita. My reasons are that Aubriet illustrated for Tournefort only parts of plants—fruits (and with dissections, too!) , seeds, flow- ers, peduncles, etc., but no idea of the appearance of the plant itself is given. Again, L. H. Bailey, in his Garden of Gourds (1937), includes page after page of drawings, almost all of which show sections of the stem, flowers, fruits and leaves. Since they also show tendrils where tendrils exist, it is apparent that such plants would climb if given a chance to do so, and I have therefore listed a number of these illustrations among the "icones"; but I have eliminated such as that of the “Mock Orange” variety of Cucurbita Pepo, which depicts a pistillate flower, a fruit with peduncle attached, a tendril, a stem in cross-section, and enlarged dissections of the style and stigma, as well as of the anther column; but habit, no!—so it is not listed. In this connection it is interesting to note that only the earliest "icones" illustrate the root. This is undoubtedly a loss to plant portrayal, and probably came about when interest in plants shifted to a taxonomic basis from the pragmatic approach of the rhizotomists and their followers. These predecessors of the herbal- ists performed magic and cures based on lore often dating from the Greek period, and generally associated, in popular folk ritual, with properties inherent in plant roots (Singer, 1928). Also worth noting is the rarity of portrayal of habitat along with habit. This is so important a factor in the systematics of botany, and was used so wonderfully by Audubon in his portrayals of birds and animals, that one wishes it were more often used as backdrop to "icones" of our Йога. Nissen's (1951) statement, “Plant sociology... 15 of recent date, and representation of it is prepared more and more with the camera,”® very possibly explains this lack. If the portrait of a plant belongs at all within the field of botany, it must be scientifically accurate, my third criterion for "icones". The degree of precise detail will vary in each plant portrait, as it does in human portraiture. There is, however, a most important difference in the requirements of the two. A portrait of a man shows all the artist can tell us about a single individual, be he godlike or sickly earthling; but, to quote from Arber (1938), “the drawing which is ideal from the standpoint of systematic botany, avoids the accidental peculiarities of any indi- 8 Translated by Mr. Alfred Sadler. (Мог. 48 294 ANNALS OF THE MISSOURI BOTANICAL GARDEN vidual specimen, seeking rather to portray the characters fully typical for the species.” This distinction is well illustrated in the very book that introduced the word we are trying to define, Brunfels’ Herbarum vivae eicones: here we find that some of Weiditz’ magnificent “living portraits” exhibit such “accidental peculiari- ties”, probably as a result of an overly enthusiastic return to direct observation from nature. Note figure 1, of Weiditz’ Arctium lappa L., in which the plant’s poor wilted leaves add nothing except the information that it had been pulled from the earth too long before Weiditz got around to drawing it. He apparently did not itch, as Tournefort felt the true botanical artist should, to get his observations on to paper. “It frets а man", Tournefort wrote (quoted from Blunt, 1951) "to see fine Objects and not be able to take Draughts of them; for without this help of Drawing, 'tis impossible any account thereof would be perfectly intelligible.” The requirements of accurate botanical illustration are that the characters of a species be accurately represented; fortuitous accident in the life story of the individual, such as its being attacked by voracious insects, is of no interest to anything or anybody but the insect and the plant. It is almost pedantic to add that accuracy is not impaired by the additional quality of aesthetic satisfaction, the fourth of my criteria. Among the "icones" the two are found in variable proportions, the balance being struck not only on the basis of the purpose for which the pictures were made, but also depending upon the date of their publication. The demands of fashion in taste and interest can be followed clearly throughout the centuries. Many flower illustrations of great decorative quality have no botanical value whatsoever, and quite correctly end their usefulness as lamp shades; this kind of work reached its nadir in the 19th century, when spinster ladies sentimentally painted charming bouquets on china, as well as any other available surface. When these illustrations appeared in books, they were usually accompanied by saccharine verse, under such titles as The Moral of Flowers (Marquand, 1947). "Today's floral illustrations represent the opposite extreme. We tend to file scientific data in compartments well isolated from all aesthetic consideration; for this reason we must beware of eyeing with suspicion the accuracy of figures that are pleasing. It is important to remember, as is subsequently discussed, that the "icones" include the work of Turpin, Fitch and many other extremely gifted artists, whose talents were exhibited in a field that demanded far more of them than simple art for art's sake. An aesthetic quality, then, is the last of four criteria set up for true "icones"; full page illustration, portrayal of habit and botanical accuracy have all been di above. Extremely few illustrations will pass all these tests; but who 9 The sentimental (ог — I ен гүне approach cannot а limited entirely, however, to spinster ladies or to the 19th c Linnaeus himself effused most colorfully: ‘The petals of the flower contribute nothing to T c ion but serve only as bridal A locate лені 5 the great Creator, who has adorne ed them MB de such noble bed-curtains and perfumed them m any sweet perfumes that the bridegroom may celebrate his параи with all ¢ the greater sacs "When the bed is thus и it is Жеген for the bridegroom to embrace his beloved bride oa ае ар his gifts to her." And E us Darwin, Кыйыны: of iade: versified "With honey'd lips enamoured woodbines meet, asp with fond arms, and mix their ees sweet." (Both quotes are from Krutch, Joseph Wood, Tbe Gardener's World, New York, 1959; p. 24.) 1961] 295 EISENDRATH—a STUDY OF THE ICONES | ре: ZZ, Ре = А ad 1 2 Қ | Ж. | у ұз € ЕЕ Sii ER 8 За 2 с 8? а v emu foe v BM u^ om oe: № —— oan "ES ЫС mse § © з б, ғ.з a << [Vor. 48 296 ANNALS OF THE MISSOURI BOTANICAL GARDEN knows better than a botanist that taxa must be elastic, and that no set of characters is invariably true for all the units that the taxon must include! “Icones” may be found in regional floras, de luxe tomes concerned only with the plants growing in a single garden, learned treatises on classification, serials pub- lished by scientific institutions or nurserymen’s catalogues. All of these fall within the enormous body of illustrated botanical literature, of which the Index Londinensis lists well over 3000 titles, and for the purposes of this paper it was necessary to establish a limited approach. One could set up chronological barriers, choosing, as did Sitwell and Blunt (1956), two centuries as a high point in botan- ical illustration. One could profitably discuss only the work of one great artist, such as Turpin, whose name appears several times on these pages. One could trace any of several aspects of the subject through the pages of Curtis’ Botanical Magazine during its almost two hundred years of publication. Or, one can approach this loose aggregate of illustrated books with a particular subject as the tool by which to cut it down to workable proportions. This method has been used, with four genera of the Cucurbitaceae the limiting factor. Key SPECIES OF THE CUCURBITACEAE AS LIMITING FacTors IN THIS STUDY Various reasons may be cited for pursuing the study of “Чсопез” through illus- trations of members of the Cucurbitaceae. As is later discussed, the family is generally distinguished by a fleshy fruit, a structure known as a “pepo”; but, whereas in many plant groups the fruits may be used to delimit various taxa, it cannot be said of the cucurbits that “by their fruits ye shall know them”. There аге “реров” and “рероз”, and the close relatives, among many cultivated forms, will produce offspring which exhibit tremendous differences. In size alone, the “реро” may be as small, let us say, as a tangerine, or absolutely enormous, as witness the Hubbard squash or the long-as-a-man-is-tall Lagenaria. It is never possible con- veniently to mount (and seldom even to preserve) such fruits as herbarium mate- rial; therefore, since it is for the edible, decorative and useable fruits that the family is cultivated, illustrations of them are of great botanical importance. The flowers, too, present a problem in survival. Though they may be showy, they collapse dismally after one day of glory in the sun, a sad truism for the taxonomist, since they present most interesting morphological variations. (It may be due to the same truism that the work of many of the best known floral artists does not appear among "icones" of the Cucurbitaceae; an illustrator who attempts to draw them painstakingly from life will find that their lives are too brief to enable him to create the illusion of prolonging them!) The choice of the key genera was made for several reasons: Fevillea because it is the primitive group in the family, serving as starting point for a number of phylogenetic series; Cucurbita, Lagenaria and Luffa because their distribution is so wide and varied as to insure their appear- ance in a variety of publications. The Cucurbitaceae include about one hundred genera (fide Lawrence, 1951). Not only the placement of the family in a natural system, but also the delimitation of generic characters have intrigued and baffled botanists over the years. This problem is, however, entirely beyond the scope of my study. 1961] EISENDRATH—A STUDY OF THE ICONES 297 The Cucurbitaceae are generally tropical plants, with some species hardy enough to extend into the temperate zones. All, however, are frost-tender herbs, and sur- vive to fruit in chilly climates only because they grow very rapidly. Cucurbita is native to the American tropics, and Luffa (with the exception of L. operculata, also New World) to the Old World tropics, whence Lagenaria probably also came, but this genus has been for a very long time universally distributed throughout the warm parts of the world. The same three genera are of interestingly varied economic importance. Cucur- bita is cultivated for its edible and decorative fruits, the pumpkins and squashes which traditionally are staples of the American diet, and archeologically are known to have been important long before the advent of the white man to either Amer- ican continent. Luffe’s fruit has a fibrous interior when mature which, dried, becomes the "vegetable sponge" so widely used by masseurs; the young fruits (apparently less abrasive!) are edible. Lagenaria fruits in an incredible variety of orms; wherever it is grown, it teases the imagination to put the fruits to use as ladles, vases, pitchers, bird houses or merely decorations. A visitor to Georgia and Florida at the turn of the century (Odell, 1904—05) found that "hardly a domestic operation there is complete without the aid of the Gourd in some form". It was even used as a cradle for negro babies! Fevillea (tropical American) is neither widespread nor economic, but is included because it is not cultivated, which fact, along with its many primitive characters, presents interesting botanical contrasts to the other genera. In Fevillea, for instance, the corolla is distinctly polypetalous, as it is also in Lagenaria. In Luffa it is deeply lobed; in Cucurbita the five petals are joined in a floral tube. Again, in Fevillea five stamens with biloculate anthers of an unspecialized type stand perfectly free, alternating in position with the petals; іп Luffa, as in the other three genera, there are usually basically only three stamens (because of the coherence by their filaments of two pairs of the original five), but they are borne free on the calyx tube. In Lagenaria the anthers of the stamens cohere lightly, but are not truly connate; in Cucurbita the filaments are connate (except that sometimes they are free at the very base) and the anthers are twisted so inextricably together in a column as to be almost indistinguishable. In none of the last three genera are the anthers primitively biloculate; they dehisce by a single longitudinal split. Of the books listed below, the following contain "icones" of Fevillea: Marcgrave-Piso, Plumier, The Botanical Magazine, Dictionnaire des Sciences Natu- relles, Descourtilz, Martius, Velloso, Engler and Rendle. Of these, all but the serial, the dictionary and the systematists concern plants of the Caribbean and South America, the earliest of them being Marcgrave's first edition in 1648. Lagenaria is illustrated, among the books listed below, by Fuchs, Dodoens, L'Obel, Dalechamps, Tabernaemontanus, Gerarde, Parkinson, Bauhin, eede, Rumpf, Oskamp-Zorn, Descourtilz, Velloso, linuma, Duthie and Fuller, Bettfreund and Bailey, indicating an early European knowledge of the plant, as well as its wide spread in tropical areas of the world. Luffa, rather strangely, is not pictured by those earlier herbalists who illustrated (Мог. 48 298 ANNALS OF THE MISSOURI BOTANICAL GARDEN Lagenaria, though both genera are thought to have originated in the Old World tropics. Among the "icones" listed we find Luffa first in Vesling, illustrating Egyptian plants in 1638. It appears next in Commelin, illustrating the plants cultivated in the Physic Garden of Amsterdam in 1706; later, it is pictured by Rheede, Rumpf, Cavanilles, The Botanical Magazine, Velloso, Martius, Wight, Iinuma, Blanco, Duthie and Fuller, Lecomte, Engler and Bailey. This indicates а wide distribution indeed, although the plant was apparently not cultivated in urope as early as was Lagenaria. Of the four genera considered in this study, Cucurbita is not only the best known and most widely cultivated today, but archeologically it is of great interest, being found in diggings where it can be used to trace the agriculture of early man in the so-called New World. Because of its tremendously variable fruits, seeds and other organs, Cucurbita also challenges the systematist and intrigues the geneticist; because its floral parts are large, it is useful to morphologists and physiologists and, because the cucurbits are easily grown, their large fruits can be retained for study, while their indeterminate growth permits flowering from maturity to frost. As a matter of fact, Cucurbita presents satisfying study material for most botanical specialists, though geneticists, pleased with the large pollen mother-cells, find diffi- culties in their miniscule chromosomes (Whitaker and Bohn, 1950). However, it is probably because of its edible and decorative fruits that we find among the icones” more illustrations of Cucurbita than of the other three genera selected. The earliest “ісопев” of Cucurbita, the two appearing in the fabulous folio edition of Fuchs, 1542, are probably also the most beautiful. We find Cucurbita next in the 1552 Aztec herbal known as the Badianus Manuscript; and at short intervals from then on, mostly in pictures derived more or less from Fuchs, cucurbits appear in Matthiolus, Dodoens, L’Obel, Dalechamps, Tabernaemontanus, Gerarde, Parkinson and Bauhin. Original figures are found in Rheede, Rumpf, Buch’oz, Schkuhr, Vietz, Descourtilz, Chaumeton, Wright, linuma, Blanco, Duthie and Fuller, Bettfreund and Bailey. The list evidences a rapid and wide spread of knowledge about and interest in the plants. EARLIEST ILLUSTRATIONS OF KEY SPECIES None of the titles in the list of “icones” of the selected genera predates 1542, the year of publication of Fuchs’ De historia stirpium. ‘This fact is significant because the historical background to a discussion of plant illustration is strangely poor. Though all of man’s existence (along with that of other animals!) must of necessity have been ultimately dependent upon the flora of his world, relatively very few early examples of plant delineation are known, in comparison to those of fauna. Among the key genera, the earliest I find are chance reprints in modern books, all of Lagenaria. The first is printed by Behling (1957), and is taken from the manu- script Tacuinum Sanitatis in the Austrian National Library. It is described as a page from the амада der sepeti; of upper-Italian wai, from the turn of the На to the 15th cen sees the flas жері B ied (Cucurbita e а, L.) in an excellent example, banging i in a thicket of leaves. The te t below dec ; "that ay Bai is cold and m and most useful to be eaten fresh = green. ig RN shire 5 1961] EISENDRATH—A STUDY OF THE ICONES 299 The second reproduction is Arber's plate iii (1938). This is from Konrad von Megenberg’s Das Puch der Natur, Augsburg, 1475, which contains the first printed plant illustrations. It shows, in the center of a group of plants, an unmistakeable Lagenaria fruit, pendulous on a terminal branch, the whole plant arising from a basal rosette of heart-shaped leaves. The third is in a 1940 facsimile of an Aztec herbal of 1552 (Codex Barberini, Latin 241, in the Vatican Library). Familiarly known, from the name of its translator, as the Badianus Manuscript, it was put together by Martin de la Cruz, described in the text as an “Indian physician .. who is not theoretically learned, but is taught only by experience" (Arber, 1938). The colored illustration shows a Cucurbita foetidissima, so identified as the leaves and root seem to belong to a perennial plant. In the herbal it is called the “Ayonel- huatl”, translated as “calabash root”, and is described as an ingredient of a complex concoction recommended to relieve the pain of childbirth. Fucus’ DE HISTORIA STIRPIUM Fuchs’ beautiful De historia stirpium was published in Basle in 1542, twelve years after Brunfels introduced "icones" as a botanical word. Like Brunfels’, Fuchs’ book is illustrated with portraits of plants made by artists who had actually looked at them. This statement does not seem startling, but it has great significance. Plant illustration in the herbals, prior to Brunfels and Fuchs, consisted of constantly recopied figures, derived from ancient originals, which in each reproduction lost more of their freshness and validity. Strangely, the invention (circa 1450) of movable-type printing, with its tremendous impact on the dissemination of all inds of knowledge, did not immediately influence publishers to have their artists take a fresh look at plants. And this, despite the fact that illuminated manu- scripts of the period often include charmingly naturalistic flower paintings, and that such artists as the Van Eycks, Van der Goes, Jacopo Bellini and Botticelli painted flowers beautifully even before a magnificent high of superb flower por- trayal was reached іп the work of Leonardo!’ (1452-1519) and Dürer (1471- 1528). Brunfels and Fuchs let fresh air into the musty chambers of illustration of the herbals, Fuchs leaving for us fine woodcuts of both Cucurbita and Lagenaria; but it is from them (see below) that subsequent generations of herbalists were satisfied to derive their illustrations, thus slamming down the window once again. Fuchs shared with Brunfels the realization of the importance of drawing plants from life, but his artists, unlike Weiditz, (whose “living portraits”, you will remember, were sometimes limited portrayals of individual plants) also understood the necessity of depicting plants typical of a species. From his own statement we know that Fuchs worked closely with the artists. > he wrote іп his preface, “is positively delineated according to the fea k "Each (picture) . м should be mos зри s ind чыын, of the living plants; we have taken peculiar care that М” and, we у» е d ы depicted poem its s, stalks, leaves, flowers, seeds and frui ruits, Further егде "E have purposely and ых pti Re de Е са of the natural form of the plants by 10 Leonardo wrote, “The painter will produce pictures of жей merit if he takes the work of =; ly himself to learn from the objects of nature he will produce good жолата Я" Edwar d, "The N Notebooks of гавал da Vinci, New York, 1934; р. (Мог. 48 300 ANNALS OF THE MISSOURI BOTANICAL GARDEN shadows, and other less necessary зак by which the delineators sometimes try to win artistic glory.” (Quoted from Arber, 8.) Fuchs was himself a physician, famed throughout Europe for his successful treatment of an epidemic disease that hit Germany in 1529, but his love of plants was more than a professional interest. Again in his preface, he wrote, e who does not know that there is nothing in this Life pleasanter and mo ны than to wander over woods, mountains, plains, garlanded and а ig with flower- lets and Ма of various sorts хрв most ли to boot, pan to gaze xod n them. But it increase e pleasure and delig a little, 5 thee be added a E ates with the virtues eer! powers of these same dish (Оне d from Arber, 1938 ) Fuchs’ great work represents the high point of the Renaissance herbal (Singer, 1928) ; the text has the botanical limitations of all herbals, but the illustrations are superb. The figures, fitted gracefully into the frame of the full folio page, do not thereby lose their naturalism. The lack of shading that Fuchs mentions sometimes makes the outlines appear thin, but this is much relieved by the presence of color, which the artists probably intended should be applied by hand, after printing (Blunt, 1951). The large folio plates of the key genera include two of Cucurbita Pepo, called by Fuchs “Cucumis Turcicus" and "Cucumer Marinus", and three of Lagenaria vulgaris, labeled respectively "Cucurbita Maior", "Cucurbita Minor" and "Cu- curbita Oblonga" In 1545 an octavo edition of Fuchs appeared which contained the same illus- trations of the plants under discussion, with exactly the same names given them, but in very much reduced size and mirror images of the larger cuts.!! They were to have a long life indeed. These small-scale cuts from Fuchs, along with some from other sources, were used for over a hundred years by subsequent generations of herbalists. It would take a contortionist to squeeze these overused or derived plant illustrations into the criteria set up for "icones"; but I feel they must nonetheless be included in this study, and I claim justification from the excellent authority of Dr. William Trelease, first director of the Missouri Botanical Garden. Dr. Trelease wrote, RÉ study of the cultivated plants of the present time can be at all complete unless the ute and arc ad desee of the herbal are consulted, for, with so mutable a class of тч ts as our flowers, vegetables, ағар сг and fruits, the tracing of their — story — бај: is по ie с of their st Phat i is asserted of кыны plan . . is also true, though to a less dine of the native ESO. ." (Trelease, 1896). We shall see that a number of the plant figures of the herbalists are far from "painstaking records", but we shall also see that others which my definition would carry me by with the merest swish of the skirt, play important roles in Linnaean classification. 11 Illustrations can be mechanically changed in size by use of a bee ы дә This instrument, с structed о four bars, has a pencil placed at the juncture of tw them, another at the tip as a a a ( for much of this information to Finan, John J., Maize in don Great hein Ann. Missouri Bot. Gard., 1948; 35:8-186 $ 1961] EISENDRATH—A STUDY ОЕ THE ICONES 301 With Fuchs’ “icones” well in mind, we now embark upon an unbelievably com- plex task, tracing from his and other true portraits of plants the often inadequate thumb-nail sketches used and reused by successive herbalists. Woopcuts DERIVED FROM ONE SOURCE AND USED BY SEVERAL AUTHORS Bailey (1929) identified Fuchs’ “Cucumis Turcicus” as “undoubtedly a Cucurbita Pepo of the vegetable marrow kind”, and Whitaker (1947 ) agrees. In 1563, in his Cruydeboeck, Dodoens reproduced the smaller cut as "Pepones magni", using it again in 1616, in his Stirpium historiae pemptades, as “Pepo maior ob- longus”. In 1587, in his Historia generalis plantarum, Dalechamps, possibly using the same cut, called it “Cucurbita Indica longa”, and in 1636 Gerarde, in herball, described it as “Pepo maximus oblongus”. Тһе picture ends its career, as far as I know, as "Cucurbita foliis asperis sive Zucha, flore luteo, Ic. II" in Johannes Bauhin's Historia plantarum of 1650. is last derivative from a magnificent original has been cut down to diminutive measurements (812 X 414 cms.); the tendrils of the plant are so stylized as to be almost indistinguishable organs, the character of the peduncle at the point of attachment is entirely lost, and the illustration becomes as far removed as possible from a portrait of a living plant. This series presents a good example, however, of the botanical significance of many of the small derived wood-cut figures of the herbals. Linnaeus in his defini- tion of Cucurbita (Species plantarum, 1753, vol. II, p. 1010) cited a variety В with Caspar Bauhin’s Pinax as reference. Caspar, the son of John, did not illustrate his book, but referred to the pictures mentioned above, from Dodoens’ work of 1616 and Gerarde’s 1636 Herball, so Linnaeus’ description is indirectly des- cended from a picture which may be described as a foster child of Fuchs’. Again, Fuchs’ “Cucumer marinus" had a long and mostly unhappy history (see Figures 2, 3 and 4). Matthiolus (Commentarii in libros sex Pedacii Dioscoridis de materia medica) in 1558 printed аз "Zucche Indiane” and іп 1560 as “Cucurbita Indica” a plate which is certainly not a direct copy of Fuchs’, but which almost as certainly was copied from his. Dalechamps in 1587 used Matthiolus’ cut, in mirror image, as “Cucurbita Indica rotunda”, and Linnaeus (Species plantarum, 1753, vol. II, p. 1010) cites Dalechamps as a reference for his Cucurbita Pepo. Bailey (1929) says of this cut that it represents “а tendril-bearing plant; and the fruit stems are long, ck eta ridged “ not enlarged next to the fruit; Se: leaf shape also answers well for the species we know as Cucurbita Pepo. The iriness along the stem is undoubtedly hana d the рае ч to ike roots is con- Pur рак sri] as vell as the mode of fruiting.’ In 1563 Dodan printed a cut more exactly derived from Fuchs’ octavo “Cu- cumer marinus", calling it ран rotundi”, and іп 1616 Dodoens reused it, this time as “Реро rotundus minor”. By 1636, when Gerarde called it “Реро minor sylvestris”, the detail was сш быу coarsened, but it is very possible that the same plate was used. In 1650 Bauhin printed another poor, small, derivative cut, this one “Cucurbita foliis asperis sive Zucha flore luteo, Ic. У. Fuchs’ illustrations of Lagenaria, too, producea a motly swarm et progeny. His "Cucurbita oblonga" is in Dodoens, 1563, as "Cucurbita anguina", and in Pain, 1650, as “Cucurbita longa, folio molli, flore albo”. Fuchs’ “Cucurbita maior” [Vor. 48 302 ANNALS OF THE MISSOURI BOTANICAL GARDEN we 29 CN Figure 2. “Сиситег marinus” (Cucurbita Pepo), p. 699, De istoria stirpium, Leonhart Fuchs, Basle, 1542. CVCVMER MARINVS 1961] Figure 3. “Реропез rotundi" > (Cucurbita Pepo), р. 508, Стиу- : deboeck, Rembert Dodoens, Ant- eck, werp, 1563. EISENDRATH—A STUDY OF THE ICONES Peponesrotundi, Ronde Pepoenen, Pepones lati, Breede Pepocnen, 303 [Vor. 48 304 ANNALS OF THE MISSOURI BOTANICAL GARDEN reappears іп Dodoens in 1563 as “Cucurbita cameraria maior", which Dalechamps also called it in 1587. (His plate, however, is a reverse of Dodoens’, with slightly different measurements.) Gerarde (1636) seems to have used Dodoens' block, calling it “Cucurbita lagenaria", and in 1650 Bauhin brings it to a small, inglorious end, again, as “Cucurbita latior folio molli, flore albo". Fuchs’ third and last Lagenaria was called “Cucurbita minor", and Dodoens (1563) called it by the same name. In 1587 it became “Cucurbita cameraria minor" in Dalechamps, and in 1650 "Cucurbita lagenaria" in Bauhin. Obviously, there was some confusion in nomenclature! All of the Dodoens illustrations are approximately the same size as the cuts of Fuchs’ 1545 edition, and possibly from the same blocks. Dalechamps' cuts are slightly smaller and all but the "Cucumer marinus" derivative in reverse of the above, so he cannot have used the originals; Gerarde's print is clearer than Dodoens', but may be from the same cuts. The cuts used by Bauhin are all smaller and far coarser than any of the others. However, not all the wood cuts used in the herbals of the period after Fuchs derive from him. А cut called “Pepones lati" in 1563 and “Реро latus" in 1616 by Dodoens is used again by Gerarde in 1636 as “Реро Indicus angulosus”, and identified by Whitaker (1947) as "C. Pepo, possibly var. White Bush Scallop”. Again, Dalechamps in 1587 published a charming decorative print of “Cucurbita verrucosa" with a good deal of fine, accurate detail, identified by Whitaker as "evidently a warted variety of C. Pepo". The same material, cut down in size, in mirror image, with far less grace and detail, was published by Bauhin (1650) as "Cucurbita verrucosa" and cited by Linnaeus, in 1753, in Species plantarum in his definition of "Cucurbita verrucosa". (But Linnaeus added, in later editions, that the plant bore marked resemblances, in flowers and seeds especially, to C. Pepo.) This is another example of the enormous botanical value of even such a coarse, much-worried hand-me-down as Bauhin's figure, especially as, according to Bailey ee) the Linnaean herbarium contains no specimen named “Cucurbita verrucosa” by Linnaeus.” In 1576 (Plantarum seu stirpium bistoria) and 1591 (Icones stirpium) L'Obel published as “Pepo maximus Indicus compressus" a cut identified by Whitaker (1947) as the first illustration of Cucurbita maxima. In 1587 Dalechamps used the picture in mirror image, as “Pepo maximus Indicus, L'Obel"; his cut measures 12 X 7 cms., a bit smaller than the one used by L'Obel, by Dodoens as “Реро rotundus maior" in 1616, and again in 1636 by Gerarde, as “Pepo maximus com- pressus". This cut (referred to L'Obel) is cited by Lamarck in his Encyclopédie, ii, 151 (1786). Bauhin is also cited by Lamarck, for his portrait of “Cucurbita aspera, folio non fisso, fructu maximo, albo sessili". In 1591 Tabernaemontanus published in his Мени’ Kreuterbuch as “Pepo Indicus minor angulosus" a cut used by Gerarde in 1597 as "Pepo Indicus angulo- sus”. print is poor in Gerarde, but the detail in the cut is excellent and is identified by Whitaker (1947) as "probably C. Pepo, var. "Table Queen’ ”. As before, Bauhin has a mirror image of the same cut, in smaller size, and names it as 12 In section 1151 of his Catalogue of the Linnaean Herbarium, S 1945, Spencer Savage lists five specimens of the genus, but none is labeled Cucurbita verrucos — соғы; —— MET LO EUNT И АЙ аа ыык S o анык он жыл зынан 19611 EISENDRATH—A STUDY ОЕ THE ICONES 305 did Tabernaemontanus. The cut is un- usual among those of the period in that it does not show a root. The same authors use another cut, originally described by Tabernaemontanus as “Реро Indicus minor clypeatus” and by Gerarde (1597) as “Pepo Indicus fungi- formis". Again Bauhin's use of it is in mirror image on a smaller scale; he calls it “Cucurbita clypeiformis cortice molli et ramosa" and Bailey (1929) says it is “un- doubtedly one of the forms of Cucurbita Pepo var. Melopepo” although tendrils are a conspicuous part of this figure, and Bailey himself (1954) describes var. Melopepo as "Plant covering small space, compact, not running or tendril-bearing". However, this last, negative character is not cited by all authorities. Again, Tabernaemontanus’ “Cucur- bita capitata” appears in Gerarde (1597 and 1636) as “Cucurbita sylvestris fungi- formis” and in Bauhin (smaller and reversed) as “Cucurbita capitata Taber- nomontani sive clypeiformis”. This, which bears no tendrils, Bailey (1929) again iden- tifies as Cucurbita Pepo var. Melope po. кез № Tabernaemontanus’ "Melopepo clype- ides cla uei qu atus" becomes “Реро maximus clypeatus” Zach fons hier светим Жі) т vol. II, p. 219, Ic. V, Historia plantarum universalis, in Gerarde (1537) and (diminished in Johannes Bauhin, Yverdon, 1650-51. size and reversed in image) “Cucurbita clypeiformis, sive Siciliana Melopepon latus a nonnullis vocata” in Bauhin. This cut is again without roots; it does, however, show exceptionally good detail of flower and fruit structure and is without tendrils. It is particularly important as it тергее another definite citation by Linnaeus in his description of Cucurbita, this time “С. Melopepo”. Bailey (1929) says "the plant is undoubtedly what we know as Bush Scallop squash”. Bauhin used Tabernaemontanus’ “Cucurbita longa” as his “Pepo Indicus minor, Tabern.” and Gerarde in 1597 = Tabernaemontanus’ “Меореро сони. аѕ his “Реро maximus compressus”; Tabernaemontanus’ “Ме|орерохегез” as his “Pepo maximus rotundus"; and Tabernaemontanus’ “Реро maximus oblongus", as (for once not complicating the problem) "Pepo maximus oblongus”. Gerarde, in 1636, had access to cuts used by L’Obel in 1576 and 1591. L’Obel’s “Pepo oblongus vulgatissimus” (1576) became “Pepo oblongus” in 1591, but [Vor. 48 306 ANNALS OF THE MISSOURI BOTANICAL GARDEN Gerarde called it “Реро maior sylvestris". L’Obel’s "Pepo rotundus compressus Melonis effigie" retains its name in both his editions, but Gerarde calls it “Реро Indicus minor rotundus", and Parkinson (Theatrum botanicum) in a very coarse 1640 reprint, "Melo Indicus parvus". The last of the cuts of Cucurbita, in Bauhin, makes a very poor print indeed, and must, like the others, have been cut down from another wood block, although I have not found it printed elsewhere. Ho r, it is another important, thoug indirect, Linnaean reference. Bauhin called it “Cucurbita foliis asperis sive Zucha flore luteo, Ic. I.” Linnaeus refers in his Hortus Cliffortianus!? and his Hortus U psaliensis to Ray's Historia plantarum“ which in turn refers to this cut in Bauhin. Bailey (1929) says, "This figure is again C. Pepo as we know it”. Gerarde in 1597 used three cuts of Lagenaria which had been used previously by Tabernaemontanus. The latter author's "Cucurbita Indica minor" became “Cucurbita anguina" in Gerarde; his “Cucurbita lagenaria minor", “Cucurbita lagenaria sylvestris”. In 1636 Gerarde used cuts as “Cucurbita anguina” and "Cu- curbita lagenaria sylvestris” which had also been used before and were to be used again. L’Obel (1576 and 1591) had used the first as “Cucurbita sive Zuccha omnium maxima anguina” and Dodoens as “Cucurbita longior". Parkinson appar- ently used the same plate, in 1640, as "Cucurbita longa." L’Obel also in both his editions used the second of these as, simply, “Cucurbita lagenaria" and Dodoens as “Cucurbita prior". Again, Parkinson apparently used the same plate, this time as "Cucurbita lagenaria maior". The constant and confusing repetition of these figures is largely due to “pools” of botanical cuts made by the publishers of botanical books. Of these the best known is Christopher Plantin, the great printer of Antwerp. Plantin’s firm con- tinued publishing for three hundred years, his place of business becoming a museum in 1876, when it was purchased by the city of Antwerp from the eighth generation of the family Moretus, direct descendants of Plantin’s son-in-law, who was also his chief assistant. Plantin published L’Obel’s work as well as the later writing of Dodoens. These two men, along with Charles de L’Ecluse (who apparently used no cuts of the key genera) were friends who worked so closely together that it is not possible to tell which or who was originally responsible for material or figures in their publications. In 1576 Plantin published Plantarum seu stirpium icones using L’Obel’s name although many of the cuts had also been used to illustrate the herbals of Dodoens and de L’Ecluse. It is thought that it was this selection of woodcuts which Johnson bought to use in his editions of Gerarde’s herbal in 1633 and 1636 (Arber, 1938). Arber (1938) believes this to be the last time the Plantin collection of blocks was used. Jacques Dalechamps, whose Historia generalis plantarum appeared at Lyons in cia syngenesia Cucurbita-g. C. seminum margine integro tumido Pepo vip dus Ж 639.” = Ciera, а, 1737; p. 452, Tse Pepo; 1. Cucurbita foliis asperis sive Zuc cha e luteo ыы ой Ран ane м. = кестеде; The Melon.” (Ray, John, Historia okie, ее 1686-1704; р. 639.) 1961} EISENDRATH—A STUDY OF THE ICONES 307 1587, and Jakob Tabernaemontanus, whose Мени’ Kreuterbuch was published in Frankfurt a few years later, both seem somehow to have had access to the Plantin "pool" of cuts, as is evident from the above tracing of relationships to the same figures found in Dodoens and L'Obel; or they used, as did so many others, the octavo blocks from Fuchs’ 1545 edition. Arber (1938) says that Tabernae- montanus made a collection of blocks for himself, and that they were acquired from him by John Norton to be used in his 1597 edition (the first) of Gerarde's Herball. According to Bartlett (1949) this was based upon a translation of Dodoens’ second herbal, begun by a Dr. Priest; Gerarde took over the work for Norton, the publisher, after the death of Dr. Priest, so the material is hardly original. Gerarde is particu- larly remembered as a horticulturalist, having supervised the care of several famous English gardens and having himself cultivated for twenty years a fine garden of his own in Holborn Although John Рава also had a famous London garden, and was known as "Herbarist to Charles I”, the examples of illustrations of my key genera found in his Theatrum botanicum leave much to be desired. OTHER Woopcurs IN HERBALS OF SIXTEENTH AND SEVENTEENTH CENTURIES The search for "icones" of the key genera takes us now to Padua, where іп 1638, appeared a little book on the plants of Egypt. The woodcuts of Luffa used by Vesling (De Plantis Aegyptiis, 1638) are highly stylized and poorly printed, and are not included as illustrations in Alpino’s original (1592) edition of the book to which Vesling is supposedly only adding "observationes et notae". However, Sims, who edited the 1814 volume of the Botanical Magazine, thought rather highly of Veling s cut. “Professor Cavanilles”, Sims wrote in connection with Plate 1638, “supposes that ‘Momordica Luffa’ of Linnaeus may probably belong to the same genus (Luffa). Indeed these two plants appear to be extremely similar, as we judge from the excellent figures of Veslingius [sic], in his observations on Prosper Alpino”. Most of the seventeenth century botanical books, however, differ from Vesling in that they reflect the stimulus which all the natural sciences received from the discovery of the New World. The earliest of these that falls within the limits of this paper appeared in Amsterdam іп 1648, the Historia Naturalis Brasiliae. The author of the section entitled Historia Rerum Naturalium was Georg Marcgrave, who had accompanied a Dutch expedition which conquered Brazil and took it briefly from the Portuguese. According to Wm. Swainson (1834), "Marcgrave's work abounds with a vastness of new and original information, very different from what was to be found in the crude and verbose compilations of this period.” He has left for us a small wood-cut of Fevillea, and several other cuts which are apparently of members of the Cucurbitaceae, but are too crude and rough to be recognizable. The cut of Fevillea most certainly does not fit my definition of “icones”, but like those of many of the herbalists it has value because of the time and place it was made. The same cut is found again in Piso’s De Indiae utriusque, Amsterdam, 1658, actually the second edition of the volume listed above. The work of Marcgrave and Piso is so inextricably entangled that it is impossible to (Мог. 48 308 ANNALS OF THE MISSOURI BOTANICAL GARDEN know who is responsible for what. The editor of the 1648 edition gave credit to both (after Marcgrave’s death), but Piso took all the credit for the second edition unto self. The credit for the illustrations, however, is of dubious value, as Nissen (1951) writes, in his inimitable Teutonic verbosity, that ‘ ‘the cuts are in no way suitable to verify a doubt-free determination of a specific object”. ENGRAVINGS The decadence into which the botanical woodcut sank after reaching a high in Fuchs’ herbal has been traced through a devious history; with the introduction of metal plate incision comes another breath of good fresh air. This introduction presents a second of the strange anachronisms in the history of botanical illustra- tion, comparable to the generally poor plant portrayals in the printed herbals during a period when beautiful flower paintings were being made. The technique of engraving on metal stems from the great goldsmith tradition of the artisans and was translated to the graphic arts sufficiently early to have reached a high point in the works of Martin Schoengauer, 1445-91. However, the earliest strictly botanical book containing illustrations printed from metal plates did not appear until a century after Schoengauer’s death (Van Schaack, 1959). This was Fabius Columna’s Phyfobasanos; it contained no pictures of my selected Cucurbitaceae, nor have I found such pictures in books printed until a hundred years later still. This fact, however, gives a falsely exaggerated impression of delayed adaptation of the new technique to botanical illustration. After the publication of Columna’s etchings, the makers of botanical prints finally recognized that the incision of metal plates permits a far more delicate line than does wood- cutting, and the genus flourished through the seventeenth century. Between 1678 and 1703 appeared the 12 folio volumes of Hortus indicus mala- baricus, copiously illustrated with double page copper engravings, by Heinrich Adrian van Rheede tot Draakestein, and containing "icones" of Cucurbita mos- chata, Lagenaria vulgaris (see plate XIV), Luffa acutangula and L. cylindrica. Malabar lies on the west coast of southern India, and Rheede was governor of the province. Blunt (1951) feels that the author may himself have had a part in making the drawings of plants, though the plates were engraved by an Italian monk, Father Mathieu. The illustrations are sometimes of immature plants and sometimes (like those of Weiditz) picture in close detail the flaws of individual specimens, but their large size and strong calligraphic quality give them a vitality which many other plant illustrations lack. This is one of the best known of the pre-Linnean botanical books which reflect the stimulus of the introduction into Europe of new plants found during voyages of exploration or scientific inquiry. The works of Commelin, Rumpf and Plumier (see below) fit into this same category, Rheede, Commelin and Rumpf particularly pointing up, as Blunt (1951) reminds us, "the interest shown by the Dutch in the flora of their colonial possessions." Commelin’s book, Horti medici Amstelaedamensis, concerns the rare and exotic plants in this Physic Garden, with illustrations drawn from life. These include a Luffa operculata, typical of the stiff engravings taken from paintings by Johan 1961] EISENDRATH—A STUDY OF THE ICONES 309 and Maria Moninckx, to which originals Blunt (1951) claims that justice is not done. Caspar Commelin, the author, was a professor of botany and the nephew of Jan Commelin, who had been director of the Physic Garden. The next illustrations of the key genera appear in the 6 volumes of Georg Eberhard Rumpf’s Herbarium Amboinense published, again in Amsterdam, between 1741 and 1750. The story of the author’s life is a series of mishaps so catastrophic that one can only wonder that his work was ever published. Rumpf became an employee of the Dutch East India Company in 1652, and settled at Amboina! where he indulged his love of nature until he died іп 1702. However, fire, ship- wreck and blindness all tested Rumpf’s patience and courage, so that one cringes from criticism of the illustrations of Cucurbita (C. Pepo and C. moschata), Lagenaria and Luffa (L. acutangula and L. cylindrica) in his book. Let me only say that their quality is extremely variable, due to the fact that Rumpf’s drawings from which his descriptions were made, were burned and he became blind before he could make more. Hence several people, including his son, made the new set, in many of which the botanical detail is poor. However, the books remain im- portant because later authors have used his descriptions and figures as the types of many binomials of plants of the Malay flora (Merrill, 1917). Plantarum americanarum listed hereafter as containing "icones" was published in Amsterdam between 1755 and 1760, and was edited by Burmann, a well-known professor of botany in that city. Burmann’s "last labor" (General Biographical Dictionary, 1813) was to procure "engravings to be executed for the drawings of American plants left by Plumier, to which he added descriptions". Plumier, a member of a French religious order, had been sent to explore the French settlements in the West Indies, and succeeded so well that he was subsequently appointed botanist to the king (General Biograpbical Dictionary, 1813). He published many of his drawings in Paris in 1693, but these did not include the Fevilleas that give him entrée to the list. His plant figures were drawn to large scale in bare, cold outline, and have little artistic merit; but they were often referred to by Linnaeus, who had access to a set of copies at the University of Groningen (Sitwell and Blunt, 1956). The next book (Histoire universelle du regne végétal by Pierre Joseph Buch'oz, 1775-80) containing an illustration (Cucurbita Pepo) that falls within the limits of this paper is the work of a botanical artist of mixed repute. Buch’oz’ works are listed by Sitwell and Blunt, by Dunthorne and in most of the catalogues of exhibits of fine flower books, but the opinion in which he was held by contemporary botanists needs no elaboration beyond the specific epithet "foetida" which L'Héri- tier applied to a plant of the genus Buchozia. Nissen (1951) quotes Pritzel in a brief statement which shows no evidence of Buch’oz’ reputation having soared posthumously. ‘Miserimus compilator”, Pritzel called him, "fraude ac ignorantia aeque emineus”. Pierre Joseph Buch’oz is credited with having written on all branches of natural science though understanding none, and with compiling some 300 volumes though remaining undistinguished himself! But his illustrations are very attractive indeed and Blunt (1951) grants him a few kind words because he intro- 15 Amboina is one of the Moluccas in Indonesia. (Мо. 48 310 ANNALS OF THE MISSOURI BOTANICAL GARDEN duced into floral illustration “the decorative qualities which we always associate with Far Eastern art". Antonio José Cavanilles is quite something else again. Of great botanical inter- est, his 6 volume Icones et descriptiones plantarum, concerning the plants that grew in Spain, either “spontaneously or cultivated in gardens”, during his time, contain rather stiff engravings with no oriental charm whatsoever. These ks were published at Madrid between 1791 and 1801, and among them is a clean cut of a | Luffa acutangula. The engravings of Rheede, Rumpf, Plumier, Buch'oz and Cavanilles were all printed and published in fine, folio volumes, representing "the apogee of the art of botanical illustration, reached in the 18th century" (Mongan, 1952). Next on my list is a group of smaller volumes, all published as handbooks to medical, economic or systematic botany. THE INFLUENCE OF THE LINNAEAN SYSTEM Christian Schkuhr published at Wittenberg, in 1791, 6 octavo volumes of his Botaniscbes Handbucb, the last 5 volumes consisting of an atlas of colored (in the Missouri Bot. Gard. copy) engravings. Nobody but Nissen and I seem to consider him worth mentioning, but I find the engravings both charming and carefully done, and Nissen gives Schkuhr credit for both the drawings and the plates. Schkuhr's Handbucb, post-Linnean, is the first of my list to exemplify the new spirit which entered the study of botany after the publication of the Species Plantarum in 1753. Linnaeus’ binomial system stimulated revision of older means of plant description and classification, and "called for a new and different type of illustration which emphasized scientific accuracy in the delineation of flower, fruit, leaf and stalk” (Dunthorne, 1938). The wealth of diagnostic detail included in Schkuhr’s plate of Cucurbita Pepo is an excellent example of this new and different type, and also illustrates Dunthorne's point that "the new scientific artists first appeared in Germany". The particular care taken in dissections of the androecium and gynoecium were of course important, because Linnaeus'!9 sexual system was based on the number of parts included in each of these floral organs. Mr. Nissen and I are joined by Sitwell and Blunt, and Dunthorne in listing the next entry, Johannes Zorn's Icones Plantarum Medicinalium, which also appears in Dutch translation by Dheoderich Leonard Oskamp as Afbeeldingin der Artseny- gewassen. The colored engravings of Lagenaria are identical in the two editions, and are quite charmi The next series chronologically on my list is mentioned by Nissen and Sitwell 16 Linnaeus’ sexual system was introduced in his Systema Naturae of 1735; the кон were grouped pt into жөке: according to e т mber of stamens, into orders according to their number of С ita comes under Clas: ‚ Monoecia, described as "Mares с сит feminis in eadem | dom o, sed div erso ay an о бан masculi et feminei in eadem planta sunt.” Besides being monoecio curbita is described as Syngenesia, or “Мапа genitalibus foedus анма ice Sta- mini athe (raro генине in cylindrum coalita English translations of Linnaeus’ Latin were made in 1783 by “a Botanical Society” with thanks for the help of Je Samuel Johnson. Thes Ба appear in the Ray Society’s 1957 facsimile 6 of Vol. ый E ies ies Plantarum in naeu xual о ssification", and one to suspect the learned doctor of baiting with tongue in cheek those cuit who had вена described Linnaeus’ mathematical system as “lewd” and “licentious” e 1961) EISENDRATH—A STUDY OF THE ICONES 311 and Blunt. Published by Ferdinand Bernhard Vietz, 10 quarto volumes including 1086 hand-colored engraved plates, Icones plantarum medico-oeconomico-techno- logicarum, appeared at Vienna between 1800 and 1820. The plate of Cucurbita Pepo is large, printed on a folded double page, and has a great deal of charm. A Lagenaria plate is also ascribed to Vietz, but it is not included among the volumes (1-3) in the Missouri Botanical Garden library. I bring in Curtis’ Botanical Magazine now because its earliest plate of one of the key genera fits chronologically into the list of "icones" at this point. Actually, publication of this splendid serial began in 1787 and has continued almost uninter- ruptedly until the present day, practically all of its plates being hand-colored until as late as 1948! William Curtis, Praefectus Horti and Demonstrator to the Society of Apothe- caries at Chelsea, founded the magazine to describe and illustrate “the most Ornamental FOREIGN PLANTS”, the illustrations to be “always drawn from the living plant”. When Curtis died the editorship was taken over by his friend, J. Sims. Sims’ plate 1638, published in 1814, gives (as mentioned above) Cavanilles as a reference for his Luffa acutangula, a lovely copper engraving, prob- ably by Sansom, from a drawing by Sydenham Edwards. Edwards is described by Blunt (1951) as “Curtis own creation", as it was Curtis who recognized talent in the son of a Welsh schoolmaster, and brought him to London for instruction. Edwards remained practically the sole illustrator of the Botanical Magazine from the publication of the second volume in 1788 until 1815, some years after Curtis’ death; he is credited with about 1650 plates! The only other of the key genera illustrated among the nearly 10,000 plates in the Botanical Magazine is a Fevillea cordifolia, plate 6356, published in 1878. It presents an interesting contrast to the Luffa, as this was a period of low ebb in the printing arts; it is lithographed on poor quality paper, and has none of the charm or calligraphic quality of either earlier or later plates. At this time J. D. Hooker was editor, and it was under him that “the last traces of the Linnaean System vanished from the classification of plants in the Botanical Magazine” (Blunt, 1951). Sir Joseph Hooker is best known as an early champion of Darwin and for his botanical explorations, which contributed greatly to the knowledge of plant geography. W. Fitch, artist of the Fevillea plate, was "discovered" by William Hooker, father of Sir Joseph, while the to-be-Director of Kew Gardens was a professor of botany and the to-be-artist an apprentice designer of calico, both in Glasgow (Blunt, 1951). Sir Joseph Hooker, after inheriting this enormously prolific drafts- man, described him as an “incomparable botanical artist”, with “unrivaled skill in seizing the natural character of a plant”. Blunt (1951) finds his work too facile and insensitive, but adds that Fitch “remains the most outstanding botanical artist of his day in Europe”, despite the fact that his drawings were made from та plants, Blunt gives him additional credit as the first artist to be able to " ^ is satisfactorily. Otto Stapf, editor of vols. 148-156 of the Botanical Magazine, has given a satisfying explanation of why this was possible: (Мог. 48 312 ANNALS OF THE MISSOURI BOTANICAL GARDEN oe ms a marvelous power of visualizing plants as they lived ies retaining ег image іп his т This emboldened him rather to treat his originals as sketches than to work them into finished кее € the result that when finally drawn on табы iy Gilde ent a certain generalization which the type of the species ope to life and took the place of a ао чу с T (Quoted from Blunt, 1951.) This statement concerning Fitch's method!" is of interest in relation to my earlier discussion of "icones" as essentially portraits of species rather than of individuals as were the woodcuts of Weiditz. Frangois Pierre Chaumeton, "docteur en médecine", well understood the value of fine botanical illustration. "There аге few sciences", he wrote іп the preface to his Flore Médicale, 1814-20, “which demand more imperiously than botany, the help of painting. One would have to try very hard indeed to replace it with the most precise description". A doctor with such an opinion would also try very hard indeed to illustrate his book with the most precise drawings, and Chaumeton did just that. His two pictures of Cucurbita Pepo were made by P. J. F. Turpin, one of the great botanical draughtsmen of the day. Turpin worked with botanists of the highest caliber, Poiteau and de Candolle, as well as illustrating many of the American plants pictured in Humboldt, Bonpland and Kunth's publication con- cerning the expedition to the "régions équinoxiales du Nouveau Continent, 1799— 1804". Turpin is described as being largely self-taught, but Blunt (1951) sees in his work “great indebtedness to the tradition of van Spaendonck and Redouté", both artists from whom no follower need feel shame to borrow. Elizabeth Mongan (1952) gives an excellent explanation for this: rance became pre-eminent in the art of botanical flower prints at the beginning of the nine- teenth century vid such artists as Redouté, Prévost, » Turpin, Poit ame Vincent. all the pupils of van Spaendonck, and the skill of these artists can be attributed sia entirely to the th h training th d had with s , but е nother reason for their ascen A This was their use of se eng raving, a ies that made possible color printing à Ia poupée. Such excellence these artists attained could not have been achiev ed by the use of the etched or engraved iise that necessi- ох coloring of the print by hand.” Two years after Chaumeton began publishing the various editions of his work (folio, quarto and octavo), publication began, under the general direction of Cuvier,” of the Dictionnaire des Sciences Naturelles. This was a most ambitious work, appearing in Paris between 1816-30 in 60 volumes of text, with 14 of atlas, and planned (according to the title page) to “treat methodically the various natural beings, from the point of view of our knowledge of them and also relative ™ Fitch also published eight је articles on the subject of n drawing, whic appeared in The Gardeners’ Chronicle 1869. a se elaborate his method i grea t det ail, but contribute cite new t шер sion о: of ure onstitutes living portraits of planes or “ісопев” 18 Among these was a so-called Fevillea, but O relegate d it to p. icydium, so the great Nova oe et species Мен (better known as “Н.В.К.”) of 1 15-25 | is not included in this discussio 19 “That is, the plate was daubed in various enar with тың knotted У resembling rag dolls ( d d then the whole ching ы. nce." (Van Schaack, 19 éopold M a Егедег ert, Bed ron Sige Рене ip was a Frenc 17 ist Pod among other high piti КАЙ. ЊЕ post 4 Professor of Comparative ‘Anatomy at the Janie des Plan 1961] EISENDRATH—A STUDY OF THE ICONES 313 to their use in medicine, agriculture, business and the arts". DeJussieu?! was originally in charge of the botanical section; Turpin of the execution of the draw- ings and the general direction of the engraving. Unfortunately Turpin made only two plates among the key genera, both of Fevillea cordifolia (see plates XII and XIII). Turpin's drawings of botanical detail are generally acknowledged to be exceptionally fine, and these plates well illustrate the fact. One of the plates, for instance, depicts not only a "sterile" (male) flower in natural size, but a view of it from below: the androecium; a single stamen; and a view of i£ from the rear. Equally detailed dissections are made of a "fertile" (female) flower; and, need I add, the habit of the plant is also shown, all in one beautiful, colored, octavo page. In 1821, M. E. Descourtilz published in Paris his 8 volumes on the Flore Médi- cale des Antilles, an assemblage made while he, a doctor, served in official capacity at Santo Domingo. Descourtilz used illustrations of several of the key genera (Cucurbita moscbata, C. Pepo—both of which are pictured twice under different names; Fevillea cordifolia and Lagenaria vulgaris) ; these pictures have a good deal of charm, but are poor specimens of botanical illustration, despite the fact that the author claims to have drawn them from nature and to have watched his son cut them down in size before painting them with the greatest care. Though the father claims that his son Théodore was a pupil of the "célébre Wanspaendonck", the slipshod attempt at portrayal of detail is only a sort of casual acknowledgment of its desirability. In contrast to the works of Turpin, these are sad indeed. LrrHocRAPHS The eleven large folio volumes of Velloso's Florae fluminensis icones, published in Paris in 1827—32, illustrate the flora of the region around Rio de Janeiro. Vel- loso was a native Brazilian who became one of the most zealous students of Domenico Vandelli, the Italian botanical explorer, and is thought to have supplied most of the plants described in Vandelli’s works (Nissen, 1951). Velloso must have been an ardent botanist to collect so many plants, but the art work in his book leaves much to be desired. The de Candolles (1878) wrote that "the figures [of Florae fluminensis] like those of Plumier, are very bad. The safest thing is to class the species of these authors...in a very doubtful category, which one can call inextricables”. The figures do, however, play some part in the history of the graphic arts as they were lithographed by Senefelder, who discovered the process "almost accidentally" (Blunt, 1951) in 1797. The advantage of lithography, the technique employed in all the remaining "icones" on my list except those of Iinuma and Bailey, is that it permits subtle gradations of line; аз is readily seen in his Fevillea trilobata, Luffa cylindrica and two Lagenarias, however, Velloso did not profit from this opportunity, but drew in bold outline only. The next "icones" on the list also illustrate Brazilian plants. Karl Friederich Philipp von Martius, a German botanist and traveler, was sent to Brazil in 1817 by the king of Bavaria. He returned to positions at the botanical garden and the university in Munich, where he began work on his Flora Brasiliensis in 1840. 1748—1836) is known as the first taxonomist to present a natura se Ps Біле, actually за improved | vein of M. месті bu 774 published, by NM шесі nti 1795, the younger toine d'Histoire Naturelle de Paris (Lawrence, 1951). [Vor. 48 314 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fifteen volumes in 40 folio parts were published between that date and 1906 in Munich, Vienna and Leipzig, the later volumes, of course, edited by younger men. The 3000 lithographs in the series were made by a number of artists, and include a Fevillea trilobata and a Luffa operculata, both of which are drawn in unusually fine detail. Robert Wight’s Icones plantarum Indiae orientalis, 6 quarto volumes of litho- graphs published in Madras between 1840 and 1856, contains some of the botan- ically finest "icones" on my list. The two-page spread of Cucurbita maxima includes an extraordinarily complete floral analysis; the Luffa cylindrica is done in such detail that one sees even the fine webbing at the edge of the leaf. The two plates, drawn by the Indian artist Rungiah and lithographed by Dumphy, are typical of the work of several generations of talented native artists who worked in India at about this time (Nissen, 1951). Wight, however, was not too happy about the way they were printed, especially in the early volumes. He apologizes in his preface with the explanation that lithography, still in its infancy in Europe, was even more of a fledgling in Madras, where inexperienced printers had not yet learned to compensate for the atmospheric difficulties which affected the success or failure of the plates. Hooker and Thomson (1855) in their Introductory Essay to the Flora Indica call Wight’s works "the most important contributions not only to Botany, but to Natural Science, which have ever been published in India”. LATER WORK IN VARIOUS TECHNIQUES When discussing the botanical woodcuts of the 16th and 17th centuries, I expressed admiration for the manner in which Brunfels’ and Fuchs’ artists handled them, but welcomed warmly the increased grace and delicacy made possible by the technique of engraving. Now, in the last quarter of the 19th century, we find a series of woodcuts so beautiful and finely made that one is forced to realize (as one always is!) that a great artist in any field surmounts the limitations of his medium. These effusions are aroused by Yokusai Iinuma’s Somoku-dzusetsu ог "Illustrated Flora of Тарап”. The set at the Missouri Botanical Garden is the second edition, printed at Ogaki in 1874 on rice paper of the highest quality. The set consists of 20 quarto volumes in which the illustrations (Cucurbita Pepo, 2 Lagenaria vulgaris and Luffa cylindrica) of my key genera extend across two pages. In discussing the beautiful woodcuts of Fuchs, I pointed out that, in black and white, they sometimes appeared to be drawn in too thin outline, having obviously been designed with the intent that they be colored by hand. In these Japanese cuts the opposite is often true; the body of the figure is inked and the line, remaining white, is so fine as to seem almost impossible of achievement. In several of the plates I cite, the enlargements of the anther column and gynoecium are printed in color. Nissen, the verbose and usually all-seeing, only notes the Somoku-dzusetsu in a list for which he expresses gratitude to others; and, to = surprise, Blunt (1951) says only that it is illustrated with “effective woodcu Manuel Blanco, in the third edition of whose Flora de жайы we find a Cucurbita Pepo and two Luffas (L. acutangula and L. cylindrica), was an Augus- tinian monk with no training whatsoever in botany, and a botanical library con- 1961] EISENDRATH—A STUDY OF THE ICONES 315 sisting only of one volume of Linnaeus. Hooker and Thomson (1855) in their Introductory Essay call his first edition of 1837 a “botanical curiosity”, a state- ment which covers its reception in Europe, though it met with great local acclaim. Neither the first nor the second edition was illustrated. The third is the work of Fathers Celestino Fernandez Villar and Andres Naves, whose only contribution, according to Merrill (1918), is their Latin translations which made Blanco’s Spanish descriptions more generally accessible. The identifications of the plates (drawn by F. Domingo and lithographed by C. Verdaguer) show many errors, a large number of them referring to species which Blanco had never seen. None of the plates of the key genera falls within this category, however. Nissen (1951) goes so far as to damn the illustrations of this deluxe edition by praise so faint it hardly fits the definition. Neither the drawings nor their lithographic reproductions, he felt, compare with the quality of “the old work”, but must be compared with their contemporaries, all, by inference, very bad indeed. Field and Garden Crops of Duthie and Fuller, published in Roorkee in 1882-93, is not mentioned by Blunt or Nissen, nor do its illustrations appear in any of the exhibition catalogues, but the lithographs of Cucurbita moschata, Lagenaria vulgaris and Luffa cylindrica, albeit utterly lacking in artistic pretension or value, merit place in my list through their other qualifications. The artist of the plates, H. Hormusji, a Parsee imported from the Bombay School of Art, drew the botanical details with great care, and, though they are not pretty, his pictures are surely portraits of living plants. J. F. Duthie was Superintendent of the Saharanpur Botanical Gardens; J. B. Fuller, Director of Agriculture and Commerce for the Northwestern Provinces and Oudh. Carlos Bettfreund’s 3 octavo volumes, Flora Argentina, published in Buenos Aires between 1898 and 1901, deal with the “plantas vivas” of Argentina, and are illustrated in the manner of the period, with poorly colored lithographs of no artistic, but presumably some botanical value. These include a Cucurbita Pepo and a Lagenaria vulgaris. The same criticism may be leveled at F. Gagnepain’s section on the Cucurbitaceae іп Lecomte’s Flore Générale de l’Indo-Chine, published in Paris in 1921. This contains a Luffa cylindrica drawn, according to the plate, by Mlles. Vesque. I find no mention of this name elsewhere, presumably because the quality of work does not redound to the glory of the artists. My criteria for "icones" have had to be as much contorted to include the last two publications, as they were also to stretch across the post-Fuchsian herbals. Now they can return to their original shape, though they must, perhaps, still retain a fair amount of elasticity. Engler’s Das Pflanzenreich would not be found on library shelves under the heading of "icones", nor would its many volumes find their way into the collection of a bibliophile. They do contain, however, besides the most universally accepted system of plant classification, a number of fine draw- ings of plants. Of the key genera, only a Fevillea trilobata appears on a full page and includes the habit of the plant, but I am grateful for it, because it is the best of its type of illustration, apparently the closest we will come, in our time, to fulfilling the criteria for "icones". The Fevillea plate contains precise and accurate botanical detail and, though not showy in the slightest lampshadish way, gives a [Vor. 48 316 ANNALS OF THE MISSOURI BOTANICAL GARDEN very pleasant impression of being a portrait of a living plant. This opinion is blessed by Blunt (1951), who points out that many of the illustrations used by Engler, the "greatest of German taxonomists” were unsigned works of Joseph Pohl, whose drawings were “conscientious, accurate, useful and in great quantity, but not on a high artistic level". Of even more significance in these, our troubled times, is the fact that they were made from specimens in the Berlin Herbarium, subsequently destroyed during World War П. ull-page illustration of Fevillea cordifolia in Rendle's Classification of Flow- ering Plants squeezes in among the "icones" by the same reasoning as does Engler's F. trilobata. The picture is in Rendle's second volume, which did not appear until 1925, although the first was published in 1904, at which time the material for the second was "almost finished" (Burkill, 1938). It is impossible, therefore, to know if I have placed the book in proper chronological sequence. For our purposes, however, it is only important that we have here again a carefully drawn plant portrait, containing details acceptable to a distinguished taxonomist. Rendle joined the Department of Botany at the British Museum at the age of twenty-three, in 1888, and remained there for fifty years, pursuing, according to his “Кееп concep- tion of his duty", his work in systematics. Burkill (1938) further describes him as one of those "ships which, as they leave port, have their course set for the whole voyage". He held many botanical posts of honor and responsibility, but his interest was always directed toward taxonomy. L. H. Bailey's Garden of Gourds, a quarto volume issued in New York in 1937, is illustrated with a number of zincograph black and white plates mechanically reproduced from pen and ink drawings. Again, they are of no artistic value, but are pleasing and contain fine botanical detail acceptable to the author, an acknowl- edged authority on cultivated plants. Of the many illustrations of key genera printed in Bailey's book, I have listed as "icones" only those which best give an impression of habit: “Wild Texas Gourd”, "Egg Gourd”, "Apple Gourd”, “Big Bell Gourd” and “White Pear Gourd”, all belonging to the species Cucurbita Pepo, var. ovifera; Lagenaria vulgaris or Luffa cylindrica. CONCLUSION Bailey's book contains the last of my "icones", representing well a type of illustration with which it seems that we must learn in these latter days to satisfy ourselves. Botanically they are, of course, authoritative, but they grade very low on the artistic scale, as do all the last group listed. Actually there have been very few on my list that rate high both artistically and botanically, but it must be remembered that this is probably because the key plants do not appeal as floral décor, and have not posed for most of the best-known botanical artists. Of the publications listed, only Fuchs in the 16th century and Rheede in the 17th contain "icones" that completely fulfill the requirements of my definition— size, habit, botanical accuracy and artistic value; Rumpf, Plumier, Buch'oz and Cavanilles in the 18th century meet the criteria in varying degree, all being at the very least large, fine books from the bibliophile's point of view. Schkuhr, Zorn, Vietz, Curtis’ Botanical Magazine, Chaumeton, the Diction- 1961] EISENDRATH—A STUDY OF THE ICONES 317 naire des Sciences Naturelles and Descourtilz are less pretentious publications of the late 18th and early 19th centuries, with the happy artistic addition of color, again complying with my requirements to some degree. Velloso and Martius return to the grand scale, but are more pretentious than beautiful, and poor Velloso is also not held in very high botanical esteem. From these two on, except for the Japanese work of linuma, the artistic measure must be dropped entirely, or my list would have to end. I cannot describe Bailey’s book (more attractive at least than the others) by even so slight a word of praise as charming, and there is no more temptation to use his illustrations than Engler’s or Rendle’s on my lamps. This may be a good thing; if the paper is of decent quality, the bookworms stay away, and if anybody supports our libraries, the books may be preserved, as books, for posterity. Though beauty is no longer important in these publications, botanical accuracy is a sine qua non; for this reason and with the message of Dr. Trelease still in mind, we can feel sure that they will serve the purposes of mankind for a very long time to come. For the sake of consistency in the following indices I have employed the spelling of authors’ names used by Nissen (1951), because his bibliography is the most complete I have found. This has led to one or two unfamiliar spellings, as “Karl von Linné" and "Rumpf". Consequently, I have used the name "Linnaeus" in the text, as it is the familiar form in American botany. LITERATURE CITED (* indicates works containing "icones" of key species) „Arber, A. Herbals. 2d ed. enia i The st nus manuscript СЕ peak ана 241, ME Library); an Aztec aig of 1552. imile in color; translat "is E. W. Emmart. Baltimor , 1940. Small paintin, Bailey, 1. Hk The domesticated odere I. Gentes herbarum 2:63-115. Ithaca, 1929. *__—__—.. The dudar a gourds. New York, 1937. Full-page 2 zincographs. M: l of culti ора о а. New Y 949. É if Sie dak Do = ае library of Mrs. Roy Arthur Hunt. у es an rler, Heinr. iE plan ntarum universalis. deed eid Dominicus Chabraeus. Yverdon, 1650-51. 3 v. спыну a ; Nisen 103) Sm Sm ede woodcu ehling, L. Die Pflanze in der ер ipsias Tafelmalerei. eim. свашта, аб Flora Argentina. Buenos Aires [1898-1901]. 5 5 NÉ 162) Colored lith- *Blanco, Машо. Flora de Filipinas. [34 ed.]. Manila, 1877-80. 4 v. in 6 (4 of text and 2 of P pu (Nissen 170) Colored lithographs. Blunt, V. The art of botanical illustration. 2d ed. London, 195 И? Brunfl, Otto. Herbarum vivae eicones. Strassburg, 1530. rived 1283; Nissen 257 Ia) Wood- étal. Paris, 1775—80. T 4 Ile du régne végé 12 v. of text and Buch’ са Pierre Joseph. Histoire I e g ы: E NE Qui ul um ro plates А7; 1-9, 11 of plates only at 7 :De thorn 5 ) Engravings Bot. 76:65—68. 1938 kill. 7 Н. Alfr ed B rton Rendle. Jour. : : у Coni es rege de, dd Candolle, Casimir de, editors. Monographiae phanerogamarum. Paris, cdd . н Cavanilles, ее піо, José. nes ес descriptiones plantarum. Madrid, 1791-1801. 6 v. (Pritzel 616; Nissen 341; Dun ca ne 75) Engra dicio : *Chaumeton, Francois Pierre, Poiret, J. ee J. В. Flore médicale. Paris, 1814-18. у. (Nissen 349; Dunthorne 78) Coil engravings [Vor. 48 318 ANNALS OF THE MISSOURI BOTANICAL GARDEN *Cogniaux, A. поранено ес Melothrieae in Engler, A., Pflanzenreich IV. 275. I. Шери, ine. Lithog arms, н. Cuecusbitaceae-Cucurbitese-Cucumesinas i in Engler, A., Pflanzenreich IV. 275; п" Leipzig, 1924. Lithog *Commelin, Caspar. Ноги medici "Amstledamenss plantae rariores et exoticae. Leyden, 1706. (Pritzel 1837; Nissen eui Engravings. * Curtis's Botanic al magazin: с. ondon, 1787— (vols. 1-14 as Botanical magazine, by rtis; У. ie ued by J. Sims and later edited by a series = Ки botanists) (Pritzel 2007; dies 23: 50; Қазыны 88) Colored engravings and li *[Dalechamps, Jacques]. His tops generalis plantarum. Lyons, 1587, 1586. 2 v. ннн 2035; 2171; Nissen 471; Dunthorne 90) Colored engravin “Dictionnaire des sciences aede par plusieurs professeurs du Jardin du Roi, et des principales coles de Paris. Paris, 1816-30. 60 v. of text and 15 v. of atlas. (Nissen 2239) Colored mall woodcu — Michel Etienne. Flor re (pittoresque et) médicale Ум Antilles. Paris, 1821-29. 87. (Pritzel gravings. “Dodoens, қалы. Cruydeboeck. Antwerp, 1563. (Pritzel 2345; Nissen 512) Small woodcuts. *——————, Stirpium мина pemptades sex. Antwerp, 1616. (Pritzel 2350); Nissen 517) Small cuts. Dunthorne, G. Flower and fruit prints of the 18th and early 19th centuries. London, 1938. *Duthie, J. ЈЕ. and Fuller, J. B. Field and garden crops of the Northwest Province and Oudh. aphs. Encyclopédie méthodiq Botanique. By Jean B. P. A. de Monet = Lamarck. Continued by . L. M. Poiret. Paris, 1789-1823. 13 У. of ek ik 4 v. of plat Engler, A. See under Cogniaux, A. Finan, J. J. Maize in the great herbals. Ann. Missouri Bot. = 4. 55:149-186. *Flora Brasiliensis. Edited Бу К. Е. Р. von Martius (later b G. Eichler and I. Urban) Munich, 1840—1906. 15 v. in 40. (Pritzel 5902; Nissen 2248) ы 8, *Flore générale de l'Indo-Chine. Edited Ьу Н. се у by H. Humbert). Paris, 1907-51. art. colored i in one of two copies at Missouri Bot. Gar Мата Primi де stirpium historia comment € vécue vivae imagines. Basle, 1545. (Nissen Woodeut oils reduced from the work of 1 eo + W., editor. The gardener’s world. New York 1 The general аы dictionary. А new edition by жыласа Chalmers. London, 1812-17. “Gerard, Je The herball, or generall historie of plantes. London, 1597. (Pritzel 3282; Nissen mall exe cuts. M The erball, or C historie of plantes. ет and amended by Thomas Johnson. AC (Pritzel 3282; Nissen 698) Small w Mas 3 B. (and ea Allen and c nough's New Tuin grammar. Boston, 1903. Hooker, J. D. The flora of -— India. London, 1875-97. — and Thomson, T. a Indica. London, d *Tinuma, Yokusai. Somo. pice P (“Illustrated flora of Japan”). 2d ed. Ogaki, 1874. 20 v. Woodcuts, partially colored. Index Kewensis plantarum phanerogamarum. Compiled by B. D. Jackson [and others]. Oxford, 1895. 2 v. Supplements. Oxford, ее 95 Index Londinensis to illustrations of flowering plants, ferns. and fern allies, = i Otto prapt [and others] for the Royal SA Society. Oxford, 1920-31. ; Supp Oxford, 1941. 2 v. Jackson, B. D. Glossary of botanic terms. 4th ed. London, 1928. Lawrence, G. H. M. Taxonomy of v ueni plants. New York, 1951. Lecomte, M. H. See эр Flore générale de l'Indo-Chine. Linné, Karl von. Hortus Cliffortianus. Amsterdam, 1737. ——————. Hortus seni Stockho rm 1748 Species plantarum. Stockholm ‚ 1753. ------. Species plantarum. А PERS of th е first ат dois With an introduction and appendix ыы У. Т. Stearn and J. L. Heller. Агер 957—5 ————, stema п: 7% Matthias de. Icones stir ‚ 1591; re 5549; Nissen 1220; although more orrectly attributed to Свара шш) Small w seu pium historia. Antwerp, Kr 2 У. in 1. (Pritzel 5548; Nissen 1218) Sm all. wanda The MacCurdy, E. paisa of Leonardo da Vinci. New York, 1939. Am See under Piso, Willem Marq , E. C. Flowers of ten centu си an address. In се nee Morgan Library. Flowers of oam centurion: catalogue of an exhibition. New York, 1 Martius, К. Е. P. von. See under Flora Brasiliensis. Te а В 1961] EISENDRATH—A STUDY OF THE ICONES 319 *Mattioli, Pier Andrea. Commentarii secundo aucti in libro: Pedacii Dioscoridis n de dica materia. Venice, 1558 and 1560. (Pritzel 5985; ge 1305) Small woodcu Merrill, E An interpretation of Rumphius’s Herbarium Amboinense. Manila, 1917. ^ (Manila. 2 of Science, Publ. 9). ------. Species Blancoanae. Manila, 1918. (Manila. Bureau of Science. Publ. 12). Mongan, E. dendi in Botanical books, prints and drawings from the collection of Mrs. Roy Arthur Hunt. tsburgh, 19 Nissen, C. ре Botanische Buchillustration. Stuttgart, 1951. 2 v. in 1. Odell, J. W. = and cucurbits. Jour. Roy. Hort. Soc. 29: 450-461, 1904-05, Oskamp, D. L under Zork оар ‘English T being a corrected re-issue. Edited by Sir J. A. Н. Murray. Oxford, 1933 “Parkinson, Joho Theatrum botanicum. London, 1640. (Pritzel 6934; Nissen 1490) Small * Piso, "Villen. and Marcgrave, Georg. Historia naturalis Brasiliae. terdam, 1648. (Pritzel 57; Nissen deg уте woodcuts — in one of two copies at тутта Вог. Саг *——, ———, ee ob. De Indiae utriusque. Amsterdam, 1658. (Pritzel 7157; Nissen 1533) Small | voden *Plumier, Charles. Plantar americanarum. Edited by J. Burmann. Amsterdam, 1755-60. (Pritzel 7217; a 1547) Е ngravin, Pritzel, С. А. Iconum botanicarum index locupletissimus. Berlin, 1855. ------. Thes за literatura botanicae. Leipzig, ~ Бау ы e Historia “ње rum. London, 1686-1704. 3 v. Rendle А. В. The P ree of flowering plants. Cambridge, ен 2 v. Lithographs. ik A Hor woe tot Draakestein, Hendrik Adrian van. Indicus Malabaricus. Amsterdam, 1678- 1703. 12 v. in 6. (Prit 467545; Nissen 1625) ie pisi жш. *Rumpf, Georg Eberhard. Herbarium Amboinense. Edited by J. Burmann. Amsterdam, 1741—50. 6 v. (Pritzel 7908; Nissen 1700) —— Savage, S. A catalogue of the Linnaean herbariu London. *Schkuhr, Christian. Botanisches Handbuch. Vittenberg, ње 3 у. of text and 3 у. of atlas. (Pritzel 8202; Nissen 17 261,5 салмас engravi Singer, C. From magic to science. #1925: Sitwell, S. and Blunt, W. Great flower gers 4 1700-1900. London, Swainson, Wm. A preliminary pese se on the study of natural ‘cen London, 1834. (The abinet cycl sera natural hist *Tabernaemontanus, Jakob Theodor. Nerd Kreuterbuch. Frankfurt, 1588-91. 2 v. (Nissen 1930) mall woodcuts. Tournefort, Joseph Pitton de. Institutiones rei herbariae. 2d ed. Paris, 1700. D Wm. [Catalogue of] The Sturtevant Prelinnean library. Annual dn of the Missouri Gard. 7:123 22, 1896. Supplement (by С. Е. Hutchings) Annual report of the ord Bot. ом! :233—316. 1903. Turpin, P. J. S. Essai did iconographie élémentaire et de philosophique des paren „е 1820. Van Schaack, С Flower prints of five centuries. Missouri Bot. Gard. Bull. 47:93-9 The Vegetables * New Y ork. I. Part. IV. The cucurbits. Report of the демі ҮогЕ Ade ки tural Experiment Senon for the year ending June 30, 1935. Albany, 1937. у *[Velloso, Jose Marianno da Conceicao]. Florae fluminensis icones. Pur xt: 11 v. (Pritzel 9727; Nissen 2046) Lithographs. * Vesling, Johannes De plantis Acgyptis observationes et notae ad Prosperam Alpinum. Padua, 1638. (Pritzel 9745; ME 2057) Full-page w = "Hes Ferdinand B rnhard. Icones plantaru mico-technologicarum. Vienna, 1800— e at Missouri Bot. Gard.) Core 9764; gnet 2062) Colored engravings. Webstes’s ace 2 dictionary. Spr Vil ker, T. W. American origin of the ене oc Mb Ann. Missouri Bot. Gard. 34:101- — and Bohn, G. V. Th ance i genetics, production and uses of the cultivated species of Cucurbita. Econ. Bot. 4:52—81. es —— — ——, Cutler, Н. C. and MacNeish, К 25. Cucurbit materials from three caves near Ocampo, Tama е i ? : *Wight, Robert. Icones plantarum Indiae orantes. Madras, 1840—53. 6 v. (Pritzel 10246; Nissen 2139) Lithographs. *[Zorn, Рана зат 51. nee der epe ое. Translated into Dutch by D. L. Oskamp. sicud к бу. (М dice Colored engravings. ecu es plan и ed. Nürnberg, 1784-90. 6 v. (Nissen 2202; Dun- thorne 35) pg va engravings. [Vor. 48 320 ANNALS OF THE MISSOURI BOTANICAL GARDEN CHRONOLOGICAL INDEX or “ICONES” OF KEY SPECIES Cucurbita foetidissima: 1552, Badianus Manuscript: "Ayonelhuatl", p. 193, pl. 109. —— maxim 1576, sieh Pepo maximus Indicus compressus", p. Tos Arve “Реро maximus Indicus, д v i, р. 626. 1591, L'Obel: “Реро maximus Indicus compressus" 1616 : otu s D: q 1636, Gerarde: “Реро maximus compressus", p. 920. 1650, Bauhin: “Cucurbita aspera, folio n an fno fructu maximo, albo sessili", vol. IL, p. 221. 1800, Vietz: ‘Cucurbita Pepo", vol. II, p. 80, pl. 148 (double page). 1840, Wight: C. maxima, vol. IL pl. 507 (debe page). Heo moschata Rheede: Ои ‚ vol. VIII, pl. 2 Pi Rumpf: Magen Wm Bonnie Pipoam Soe vol. V, p. 399, pl. 1 1821, Descourtilz: "Cour à Verrues", vol. V, p. 74, pl. 322; d феи Girau- um 5 рей 1882, Duthie and Е ullar ae es icy Pty ul. Il, pls. 58 and 59; and C. moscbata, var., vol. ЇЇ, and H E 3 б < Cucurbita Pepo: 1542, Fuchs: “Cucumis tea ‚р. 698; and "Cucumer marinus" P 699. 1545, Fuchs: “Cucumis Turcicu: э. p. 402; and "Cucumer marinus", p. 403. i sp. 292: 55292: 1563, Dodoens: “Реропез lati”, “Реропев rotundi" and “Реропев magni", p. 50 1576, L'Obel: “Реро bu! vulgatissimus" and “Реро rotundus ef ы ' Melonis effigie", oth on p. 365. 1587, Ойы: “Сис urbita sais bua vol. I, p. 616; “Cucurbita Indica longa" and Cucurbita verrucosa", vol. I 1591, Tabernaemontanus: “Реро maximus oblongus” ‚ “Melopepo compressus" and “Мејореро- teres”, vol. II, x 178; “Pepo Indicus minor angulosus”, vol. II, р. 179; "Cucurbit 2. longa”, vol. II, 1591, L'Obel: “Реро oblongus”, р. 641; “Реро rotundus compressus, Melonis efigie” ‚р. 642. 1597, Gerarde: “Pepo maximus rot endure and “Реро maximus oblongus", р. 773 Рио тахі- mus ——M€ апі “Реро Indicus uocat : 4 1616, Dodoens: “Реро maior oblongus", p. "Pepo rotundus minor" and “Реро latus", p. 666. 1636, Gera rde: “Реро maximus oblongus” ‚р. 919; “Pepo Indicus angulosus” and ‘ T De minor rotundus", р. 920; “Pepo ті пог sylvestris” and “Реро maior sylvestris”, р. 1640, Parkinson: “Мејо Indicus parvus? > P- 1650, Bauhin: ‘Cucurbita foliis asperis sive Tocha flore luteo, је Ё IE HE IV, V vol I p. 218—219; "Pepo Indicus minor angulosus, Tabern." д қы . 226; “Реро Indica mabe Tabern.”, vol. II, p. 227; “Cucurbita verrucosa”, vol. II 21741, Rumpfi. qo en Indict wali, Pepones Labo”, vol. V. р. 399, pl. 145. 1775, Buch’oz: С. Реро, D г 1814, chen: “Cy cans А, ш, pl. 123 ме 123 bis. 1821, Descourtilz: "Courge patisson”, vol. V, p. 82, pl. 324; "Courge Pepon”, vol. V, p. 89, 1874, Берл С. tns vol. II, P RE ae 3 bis. 1877, Blanco: “Cucurbita р axima", а or, P 320. 1898, petii ен ашса". vol. II, C. Pepo, var. ovifera: 1791, Schkuhr: “Cucurbita ovifera", 2- VI, ты 1937, ee “Wild Texas Gourd”, p. 5; “E iq s : and 11; "Apple Gourd", p. 15; g Bell Gourd”, p. 29; and hse Rd с С. gaa var. Melopepo: es с “Меореро clypeatus” > vol. П, а 178; “Реро Indicus minor clype- vol. JUL p де. = ee ah р + 1597, Gerar de : Pid m 2% Dr тейік fungiformis", р. 775; "Cu- urbita Беде байта 75, = Te TU 1961) EISENDRATH—A STUDY OF THE ICONES 321 1636, Gerarde: “Cucurbita санету fungiformis", 924. 1650, э» uhin: “Cucurbita clypeiformis cortice d et ramosa", “Cucurbita capitata Taberno- ntani siv кр жы, eee » vol. 225; “Cucurbita clypeiformis sive Siciliana, 224. г oe a nonnullis vocata", ig ТІ, р. Fevillea cordifolia: 1755, Plumier: “ЕеуШеа foliis cordatis”, р. 203, pl. 2 1816, += onnaire des Sciences Naturelles: “Perilea ce is Poir.", vol. LXIII, pl. 210 and 1821, е "Nandhirobe à Feuilles р m , vol. II, p. 21 1925, Rendle: Fevillea cordifolia, vol. II, p. Fevillea cordifolia, var. bederac 1755, Plumier: “Fevillea folis trilobis", p. 204, pl. 210. Fevillea Moor 1878, cu Botanical Magazine: F. Moorei Hook, 3rd series, vol. Fevillea trilobata: 1648, ecd ded des “Ghandiroba ЈЕ DAR Lib. I, p. 46. 1658, “МҺапйгоҺа”, Ca 1827, тд. “Fevillea са. lia” $ i qs 4: 102. 1878, Martius: F. trilobata, vol. VI, pt. 4 А 37. 1916, Engler: Е. trilobata, vol. IV, 275, Lagenaria vulgaris: 1542, 1545, Fuchs: “Cucurbita maior", рр. 368 and 209, resp.; and 210, resp.; “Cucurbita oblonga", pp. 570 and 211, resp. 6, pl. 198. XXXIV, 1878, pl. 6356. "Cucurbita minor", pp. 369 1563, Ге ова "Cucu rbita cameraria maior" and "Cucurbita minor", p. 511; "Cucurbita anguina”, p. 512. 1576, а "x "Cucurbita sive zuccha, omnium maxima anguina" , and "Cucurbita lagenaria", 1587, Pacis “Cucurbita cameraria minor" and “Cucurbita cameraria maior", vol. I, p. 1591, ee "Cucurbita sive zuccha omnium maxima anguina" 644. 1591, Tabernaemontanus: “Cucurbita lagenaria maior”, ed II, ' and “Cucurbita lagenaria minor", vol. II, 1597, Gere do "Cucurbita anguina" and "Cucurbita ны", sylvestris, р. and “Cucurbita lagenaria", p. 181; “Cucurbita Indica р. 777; “Cucurbita lagenaria 1616 н. аа а ргіог”, P. 668; "Cucurbita longior" ES pee latior", 1636, arde: ben rbita lagenaria" nd "Cucurbita anguina", estris", p. 9 а 1640, Parkin nson: Уве ifbita lagenaria maior" and "Cucurbita longa", p. 7 1650, Bauhin: “Сис as ae folio molli, flore albo", vol. II, p. molli, flore albo 215; “Cucurbita lagenaria", есі. 1678, Rheede: “Веја- Ке $ n VIII, pl. 1; "Caca Palam", vol. vol. VIII, pl. 5 ҮШ, р pf: circ lagenaria", vol. V, p. 398, pl. 1 p. 669. ; "Cucurbita Аней 214; "Сша latior folio II, p ҮШ, x г. “Сара Schora", 1741, uae 1779, н Zorn- Aere p: мены кена" E "vol. VL E 597 and 598 and vol. IV, pl. and 321, 1821, абе “Cou rge calebasse", vol. ne pl. 3 1827, Velloso: “Си curbita lagena у vol. X pl. > Аы Реро”, vol. X, рі. 1 1874, apr M agenaria Dasistem а Ма." , vol. XX, pl. 44; 'Lapenatis асобн Ser" , vol. 1882, odis d Fuller: Lagenaria, vol. II, pl. 4 1898, Bettfreund: Lagina ер nain vol. I, p. i 1937, Bailey: Lagenaria, p. 9 Luffa acutangula: 1678, Rheede: "Picinna", vol. VIII, pl. 7 1741, Rumpf: "Petola y bengalensis” 2 E v, р. = а 149. 1791, Cavanilles: “Luffa foetida”, vol. I ыы 1814, 4. Botanical Манба : "Luffa о: “Cucumis acutangulis", међа II, pl. 2 un agnis and Fuller: L. acutangula, vol. II, pl. is foetida", is series, vol. XXXIX, 1814, pl. 1638. Уо, 48 322 ANNALS OF THE MISSOURI BOTANICAL GARDEN € cylindrica: 638, Vesling: "Luffa Arabum, seu Cucumis охе latus Aegyptius", рр. 50 and 51. E Rheede: “саға рісіппа”, vol. VIII, р 1741, ӨТ “Ресо!а sylvestris", vol. ү i 410, pl. 150; "Petola Tschina", vol. V, р. 406, 1827, Velloso: “Momordica carinata” „жо. X. pl. 97. 1840, Wight: "Luffa decree ra" > vol. П, pl. 499. 1874, Iinuma: “Luft tola, , vol. хх, P 36 1877, Blanco: таба rota", pcs. II, pl. 3 1882, Duthie ey Fu ler: "Luffa Ae аура" К ud П, pl. 63. 1907, Lecomte: Г. cylindrica, vol. П, р. 1924, Engler: Г. ааа» val. IV 275. "i р. 66. 1937, Bailey: L. cylindrica, p. 105. Luffa echinata: 1924, Engler: Г. echinata, vol. IV. 275. П, р. 70. Luffa operculata: 1706, Commelin: “Momordica Americana Fructu Reticulata Sicco”, fig. 22. 1878, Martius: L. operculata, vol. VI, pt. 4, p 1924, Engler: L. operculata, vol. IV. 275. un. " 70. INDEX, BY Key Species, oF Woopcuts DERIVED FROM ONE SOURCE AND USED BY SEVERAL AUTHORS (“Mirror image” prints are reverse of earliest listed use of figure.) Cucurbita maxima: 1576, L'Obel: “Реро maximus Indicus compressus"; 7 X 1 1587, Dalechamps: “Pepo maximus Indicu s, L'Obdl"; TX Be cms.; mirror image. 1591, L'Obel: “Реро maximus Indicus compress" T OXCI 1616, Dodoens: “Реро една maior"; 7 Х 13 cms. 1636, Gerarde: "Pepo maximus comp reni": 2 УЕ 1650, Bauhin: кеме аврега, folio non бег; кыйы: йү ий, albo sessili"; 412 X 812 cms.; mirror imag Cucurbita Pepo: 1. 1542, Fuchs: CD marinus" Е са рав 1545, Fuchs: “Сисит tavo к, mirror image. 1558, Matthiolus: “Zuc cc ie fa 32 эр X 6% cms. 1560, Matthiolus: A acini Indici 12 X 615 cms. 1563, oens: "Pepones rotundi"; 12 X 7 cms.; mirror image. 1587, Dalechamps: “ vidus ы rotunda”; 12 X 7 cms.; mirror image. 1516 Dodoens: “Реро rotundus minor"; 12 X, 9 cms.; mirror ima, ner 2: 1542, Fuchs: "Cu cumis Turcicus"; folio p 1545, Fuchs: "Cucumis Turcicus"; ; deed әкей mirror image. 1563, Dodoens: “Реропез magni"; 12 cms.; mirror image. 6 1616, oens: = maior lb cages 12 Х 7 cms.; mirror im 1636, Gerarde: ongus"; 12 X 7 cms.; mirror 1650, Bauhin: “Cosa foliis asperis sive Zucha flore bit», їс. E nmm X 4% cms. X 156, Dodoens: “Реропев lati"; 13 X 712 cms. 1616, Vets 4. latus”; 15 X 716 cms. 1636, Gerarde: Seu з angulosa”; 13 X 7 6 cms. 4. 1576, = : “Pepo oblongus Wesce тренді 15 Х 7 1591, L’Obel: “Реро тутын) З УСТ 636, Gerarde: “Реро maior sylvestris” $ {3 5X7 5. 1576, L'Obel: "Pepo rotundus compressus па ну ж 113 X 7 сви 1591, L'Obel: as above 1636, Gerarde: “оеро Indicus minor vy 3:15 #22 7 cms. 1640, Parkinson: “Melo Indicus parvus"; 3x7 6. 1587, Dalechamps: “Cucurbita verrucosa”; 11 X ы cms. 1650, Bauhin: “Cucurbita varrucosa"; 812 414 cms.; mirror image. 1961] EISENDRATH—A STUDY ОЕ THE ICONES 323 ің, 1591, Tabernaemontanus: “Cucurbita NE ; 12 Х 7% cms. 1650, Bauhin: “Pepo Indicus minor, Tabern.”; 812 X 41% cms. el ee 1591, Tabernaemontanus: 1597, Gerarde: “Pepo maximus compressus"; 12 X 7% cms. 1591, аас “Melopepoteres” 12 X 7 cms, 1597, Gerarde: “Реро maximus rotundus”; 12 10. 1591, Tabernaemontanus: “Реро maximus oblongus”; 11 UA Х7 cms. 1597, Gerarde: as abov 1591, ии “Реро Indicus minor ered 12 X 7% cms. 1597, Gerarde: “Реро Indicu у арии $; 12 Х7% 1650, Bauhin: “Реро Тайсон minor angulonis, Табан”; + 29 X 416 cms.; mirror image. a ке а Cucurbita Реро, var. Melopepo: 1. 1591, Tabernaemontanus: “Реро Indicus minor clypeatus”; EM X 7% cms. 1597, ес “Реро ћи fungiformis”; 1214 X714 с 1650, Bauhin: “Cucurbita чыр formis cortice тый, et ramosa"; 815 X 412 cms.; mirror image. 2. 1591, überaus tanus: “Cucurbita capitata"; 12 X 7 У; cms. 1597, Gerarde: Berd: а hace fungiformis”; 12 X 71 cms. 1636, Ghi de: 1650, cens "Cucurbita capitata Tabernomontani sive clypeiformis"; 812 X 41 cms.; mirror 3. 15 , Tabernaemontanus “Ме[ореро сурени" 312% X 7 cms. 1650, Beuhiné- “Cucurbita clypei formis sive Siciliana, Моня latus a nonnullis vocata"; 8% 41 cms.; mirror image. Fevillea trilobata: 1648, Marcgrave-Piso: "Ghandiroba vel оженат, 8 X 1416 cms. 1658, Piso: "Nhandiroba"; 8 X 1416 c Lagenaria vulgaris: 1542, Fuchs: “Cucurbita p ; folio page. 1545, Fuchs: “Сиси жас; € oe жі mirror E 1563, Dodoens: “Сис 2X 1650, Ba bius Tiros s longs, ae ЈЕ. ka p e $5» X p^ cms. 2. 1542, Fuchs: “Cucurbita maior"; folio p pan 1545, Fuchs: o ins maior"; octavo page; mirror ima 1565, Dodoe о bi ч cameraria maior"; 12 X 615 eel .; mirror image. 1587; пије ee meraria maior"; 12 iie 6 Di: cms. 1636, Gerarde: “Сс bita peers 12 X ву с г image. 1650, Bauhin: “Cucurbita latior folio moli flore albo"; p РА У 5 cms. 3. 1542, Fuchs: “Cucurbita minor"; folio page. 5 2 1545, Fuchs: “Cucurbita minor"; octavo posit mirror image. e іпог”; 6 : Cu 1587, Dalechamps: “Сис cameraria minor"; 1115 Х cis с cms. 1650, p : спине pedore 815 X 416 cms 4. 1576, L'Obel: Vie sive zuccha omnium maxima anguina"; 13 X 7 cms. 1591, L'Obel: as abov oens: "Cua Site ster ; 13 Х 7 ems. 1636, dne “Cucurbita an ; 13 X 7 cms. 1640, Parkinson: “Cucurbita longa" ; D X 7 cms. 5. 1576, L’Obel: “Cucurbita lagenaria”; 15 X 7 cms. 1591, L’Obel: as above. 1616, Dodoens: “Cucurbita prior”; 13 X 7 1636, Gerarde: “Cucurbita шаг: ын; ; `13 Х 7 cms. 1640, Parkinson: “Cucurbita lagenaria maior”; 13 di^ 7 cms. 6. 1591, Tabernaemontanus: “Cucur rbita => minor”; 12 Х 7% cms. 1597, Gerarde: “Cucurbita а. 2 X 716 cms. 2, 1891; M ntanus: "Cucurbita lagenaria Luce ; 12% X 7 cms. 1597, Gerarde: "Cucurbita venis их cms. 8. 1591; а ве anus: ‘Cucurbita lagenari: т, 12 ge 2% cms. 1597, Gerarde: є "басын lagenaria sylvestris”; 12 Х 7 к nA — о “ [Vor. 48 324 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE XI “Реро” о a vol. II, pl. 33, Institutiones rei berbariae, Joseph Pitton de Tournefort, Paris, 1 ХІ PLATE Tab .33- 1961 ANN. Mo. Bor. GARD., Vor. 48, pe E NON Г ne eee z 2233 dudit HU РАТНЕ ЕНЕН n I } 1 i LM PLATE XII ANN. Mo. Вот. Gard., Vor. 48, 1961 BOTAN IQI E. DICOTYLEDONES. ЊЕ Nandirobees. 777. У | { 1 * № 4 d | ы ыс — 5 ие ЖЕР 2% — % i ж 4 ^ - re. 7h Urpin pina Гор drew 7 Zorgeaht sendy NANDIROBE а feuilles de lierre. 1961] EISENDRATH—A STUDY OF THE ICONES 325 EXPLANATION OF PLATE PLATE XII Nandirobe à feuilles de lierre" TM cordifolia), vol. LXIII, pl. 210, Dictionnaire s Мана: Naturelles, Paris, 1816-3 [Vor. 48 326 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE XIII “Nandirobe a feuilles de lierre” L Fevillea cordifolia), vol. LXII, pl. 211, Dictionnaire des Sciences Naturelles, Paris, 1816- ANN. Мо. Bor. Ganp., Vor. 48, 1961 РАТЕ XIII : BO'TANIQUE. DICOTYLEDONES. Nandirobees 27 Б i, о i RERUM = 2... Гогутай “еее d à feuilles de herre. ea : NANDIROBE EISENDRATH—A STUDY OF THE ICONES ANN. Mo. Вот. Garb., Vor. 48, 1961 РРАТЕ XIV EISENDRATH—A STUDY OF THE ICONES 19611 EISENDRATH——A STUDY OF THE ICONES 327 EXPLANATION OF PLATE PLATE XIV "Caipa Schora" (Lagenaria vulgaris), vol. VIII, pl. 5, Hortus Indicus Malabaricus, Hendrik Adrian van Rheede tot Draakestein, petra dan 1678—1703. A SYNOPSIS OF POINSETTIA (EUPHORBIACEAE) ROBERT L. DRESSLER ABSTRACT The рее i: pM ttia in the tribe Euphorbieae is медет 52 а key is given for the Am can gen me 7 e opbylla is found to be ppl cable weedy tropical plant, and the н between this and the North American Р. суа? anrea are + бу Gametic chro some numbers of 14 and 28 are reported for both Р. A ah and P. dentata, iie le P. betero- ША dich only n — The geographic variati f th North American annual ipods. is briefly discussed. A key is pide for ds 11 species sind à, of which two are аа as new: P. coccinea and P. bonis a, while new a are published in Poinsettia for P. colorata, P. pentadactyla and P. restiacea. Ковевт L. Dresser, Missouri Botanical Garden, 2315 Tower Grove Ave., St. Louis 10, Missouri. There are not many species in the genus Poinsettia, but their bewildering foliar polymorphism has caused a great deal of confusion. While studying the genus Pedilanthus (Dressler 1957), some observations were made on other genera of the tribe Euphorbieae. In the summer of 1957 I was able to observe most of the North American species of Poinsettia in Mexico and the southwestern United States.* Many collections made at that time have been grown and studied at the Missouri Botanical Garden. These observations have permitted considerable clarification of the annual species, especially. While I am no longer actively studying the Euphor- bieae, it is felt that these notes on Poinsettia may be of some value. GENERIC CONCEPT The Linnaean genus Euphorbia, as it is frequently used, includes most of the tribe Euphorbieae, and is surely the most broadly inclusive generic concept in modern usage (among flowering plants). I much prefer to recognize about four- teen genera in the tribe Euphorbieae, six of these being included in the genus Euphorbia by many authors. I have no quarrel with those who prefer the inclusive concept of Euphorbia, but feel that the same arguments could be used to reduce most Compositae to Aster, with about equal profit. In the narrower sense, Ен- phorbia is a distinctive group of about 80 Old World species which are woody or succulent, and usually have two spines (rarely 1) near each leaf base. These spines are frequently considered to be stipular, but true stipules appear to be present in addition to the spines. The large group of south African succulents which are spineless or have spines formed from axillary branches are not directly related to the above group. The earliest generic name for the south African group is Medusea Haw. Neither of these groups is represented in the New World, the American succulents being members of Agaloma or Tithymalus. The largest and most difficult of the segregate genera is ee which is largely an Old World group, but includes three major American groups: (1) Primitive woody species, such as Euphorbia (Tithymalus) fulva Stapf and Tithymalus calyculatus (HBK.) Kl. & Gke., of the West Indies, Central America and the Andes. (2) Herbaceous * Т am especially grateful to the Gray 225 and the М ational Science — д for their support »r he field work in northeastern co. Special thanks are also due Dr. Rogers McVaugh, who kindly lent material of P. restiacea rens he collected, and Dr. John D. Duo for aid with the Latin diagnoses Issued April 19, 1962. (329) [Vor. 48 330 ANNALS OF THE MISSOURI BOTANICAL GARDEN perennials related to Tithymalus chrysophyllus Kl. & Ске. and Eupborbia (Titby- malus) floridana Chapm. This group includes two or three species of the south- eastern United States and a number of southern South America, all of them with a conspicuous pistillate calyx. (3) Herbaceous species which are closely related to Old World sorts and are surely geologically late invaders of the New World. These include Tithymalus commutatus (Engelm.) Kl. & Gke. and T. campester (Cham. & Schlecht.) Kl. & Gke., and are largely montane forms. There are also a few anomolous species which do not fall into these groups, such as Tithymalus trichotomus (HBK.) Kl. & Gke. and Eupborbia (Titbymalus) pteroneura Berger. Most New World Euphorbieae belong to the distinctive group made up of Agaloma, Poinsettia and Chamaesyce. Of the three, Chamaesyce is most frequently treated as a distinct genus, while Agaloma generally has been kept in Euphorbia. ese two groups are closely related, but can be distinguished by habit and by the distinctive leaf structure of Chamaesyce. While Agaloma can almost always be distinguished from Tithymalus by the appendaged glands, this feature is not in- fallible. е diminutive Euphorbia misella Watson, of alpine -meadows in central and southern Mexico appears to lack gland appendages, but close scrutiny shows rudimentary appendages beneath the glands, indicating that it is a reduced member of Agaloma. Similarly, Euphorbia heterophylla var. eriocarpa Millsp. may belong in Agaloma, but lacks the gland appendages characteristic of that group (see excluded species, p. 340). Agaloma is greatly in need of monographic study, and there would be very little profit in wholesale new combinations made without such For this reason, I propose to cite the species of Agaloma as, for example, "Bupborbia (Agaloma) corollata L.," with the understanding that this group is quite distinct from true Euphorbia. The same system, of course, may be used for Medusea, or even Tithymalus. А tentative key to the New World genera of Euphorbieae is given below. 1. Leaves opposite or (rarely) ternate; the branching dichasial throughout; the veins of the pipa sheathed with gue d (usually visible | by transmitted Ae CHAMAESYCE S. F. Gray le 1. Leaves various; at the base of the plant monopodial; leaf veins without distinct chlorenchyma dad 2. Glands n the involucre раја or entire, often with two lateral horns, but without distinct petaloid appendag Glands of the НЕ тежа 4 or 5, not cup-like or bilabiate.............................. TirHYMALUS Trew 3. Glands usually 1 or 2, de Hike, Ми ог зен 98 The involucral lobes с their аріс united into a lateral glan shield-like structure; c5 pst ааны: he каны flowers —€— be: мтниз Millsp. 4. The involucral lobes apically distinct; the gland cup-like or едені nforeenc а ЫҚ еті pleiochasium ог dichasium, the branches becom ; the e flower naked ны NSETTIA ж 2. Glands of ee involucre with distinct petaloid appendages, these often laciniate..........--.------------ 5. The involucral lobes equal; the gland appendages not forming a single hood-like mir- r ALOMA R . The involucre highly zygomorphic, with 2 large lobes and 3 very “о ones; the ых. appendages forming a spur оп one side of the involucre.................--- EDILANTHUS Neck. GENERIC RELATIONSHIPS As noted above, Poinsettia is closely allied to Agaloma, though readily distin- guished by the lack of petaloid appendages on the glands. Euphorbia (Agaloma) lancifolia Schlecht. is especially similar to Poinsettia, and resembles that genus in 1961] DRESSLER—A SYNOPSIS OF POINSETTIA 331 е 1. Cyathia and seed о rth American annual poinsettias. А. P. cyathophora T 2430), B. P. бита | (Dressler 2254), C. P. dentata (Dressler 2198). Above: side view of cyathium, X5; middle: top view of cyathium, Ж5; below: dorsal view of seed, Х7.5. All drawn from fresh, living сабое habit, in the tendency toward reduced gland number, in the form of the glands, in the reduced petaloid appendages and in the thick-walled, coarsely reticulate pollen grains. The seed of E. (A.) lancifolia is very different from that of Poinsettia, which serves to reinforce the generic distinction. Several other species of Agaloma have been confused with Poinsettia, though they are less closely related (see ex- cluded species, p. 340). Poinsettia is evidently derived from Agaloma by the loss of petaloid appendages and reduction in number of glands. While the involucre of Poinsettia generally has only a single gland, the central involucre of a pleio- chasium, especially, is likely to have a full complement of five glan THE IDENTITY оғ EUPHORBIA HETEROPHYLLA L. Much of the confusion concerning the annual poinsettias has centered about the identity of Linnaeus’ Euphorbia heterophylla. The epithet seems very descriptive of the familiar North American plant with red and green floral bracts, and most botanists seem to have assumed that it applied to this plant. One finds, though, that Linnaeus’ epithet refers to the shape of the leaves, rather than to their color. [Vor. 48 332 ANNALS OF THE MISSOURI BOTANICAL GARDEN There is no specimen of E. heterophylla іп the Linnaean herbarium, so one must turn to Tithymalus curassavicus, salicis 9 atriplicis foliis varius, caulibus viridantibus of Plukenet's Almagestum Botanicum (1696). Plukenet's figure shows a plant with both lanceolate and pandurate leaves, quite characteristic of the weedy tropical species with green floral bracts. Leaf shape is none too dependable in this group, but the entire lanceolate form shown is rarely encountered in the North American P. cyatbopbora. It is conceivable, of course, that the North American ornamental had been introduced into Curacao by that date, but it is hardly likely that Plukenet would have failed to mention the red floral bracts of that species. It should not be concluded that Linnaeus' epithet has been consistently and discerningly misapplied to the North American species. Rather, most specimens of either species from North and Central America have been determined as E. hetero- phylla, while South American specimens of true E. heterophylla have more often been labeled as E. geniculata or E. elliptica. The consistent confusion of these two species is based on very superficial observation, for the two differ from each other much more than either differs from P. dentata, which is generally recognized as distinct. A tabular comparison of the two species is given herewith. Poinsettia cyathophora Poinsettia heterophylla Stems and petioles glabrous or Stems and petioles often markedly sparsely pilose Leaves glossy green. Leaves dull green. Floral bracts usually basally red. Floral bracts green or basally pale, never basally red (often purple potted). Gland broad, more or less "DEN Gland narrow, substipitate, the the opening narrowly oblon opening circular. Seed gsi Be and sharply Scere Seed coarsely and bluntly tubercu- not angular. late, angular. CHROMOSOME NUMBER A number of chromosome counts have been made from material cultivated at Missouri Botanical Garden. These are given below, with the counts previously published by Perry (1943). Voucher specimens are available for all those which have collection numbers. ameti matic Poinsettia cyathophora ~ No. Я aros Dressler 1821 (Tamaulipas) . 14 Dressler 2271 (Tamaulipas) 14 Dressler 2430 (Nuevo León) 14 $. n. ts но, oe: 14 n. Gre t, Misso 14 Drala 2373 о) 28 Cultivated (Rex Pearce) 28 Cultivated (Parke) 28 Perry (cult. as Euphorbia heterophylla) 56 19611 DRESSLER—A SYNOPSIS OF POINSETTIA 333 Gametic Somatic Poinsettia dentata No. No. ме эн 2268 (Оахаса) 14 ummit, Missouri 14 Diss 2198 (Texas) 28 mpalme Jue a Guanajuato 28 Perry, лы Со., UN 14, 28 Dav s, Okla hom 28 dul Hill, Mies 56 Gametic Somatic Poinsettia heterophylla No. No. Dressler 2136 (Tamaulipas) 14 Dressler 2194 (Arizona) 14 Mec 2243 (Vera С 14 ward & Proctor 13883 (Jamaica) 14 Ma nning & Manning 531217 (Jalisco) 14 s. n. Soledad, Cuba 14 Moyer (cited by Perry) as E. geniculata 28 Somatic Poinsettia pulcherrima No. Perry 28 From the above counts, one could consider 14 to be the base number for the genus, if it were not for Perry’s record of a somatic number of 14 for one plant of P. dentata. It is at least conceivable that the plant studied by Perry was a chance haploid, but more counts are needed from the eastern United States, before this can be evaluated properly. There is no indication of polyploidy in P. hetero- bbylla, but P. cyathophora and P. dentata both contain polyploids. These аге discussed further in the next section. PATTERNS OF VARIATION The most outstanding feature of the annual species of Poinsettia is their extreme polymorphism. Linear-lanceolate, ovate and deeply pandurate leaves may be found not only in the same population, but on the same individual. More frequently, however, extremely different leaf types are found on different individuals or on different stages of one individual. It would appear that there is an interaction of genetic and environmental control in such cases. The perennial species are less markedly polymorphic, but all of the adequately sampled species show some degree of foliar polymorphism. Even seed structure shows а good deal of variation in Poinsettia, though this feature is usually dependably stable in the Euphorbieae. In spite of the extreme polymorphism, some patterns of geographic or genetic variation are evident, and merit more detailed study. 1. Poinsettia cyathophora. This species is perhaps the most highly variable of the genus. The cultivated polyploids (п = 28) are coarse, somewhat weedy plants that show little polymorphism, and have somewhat coriaceous leaves. When col- lected in a dry, rocky arroyo, the plants of Dressler 2373 were relatively small, but when cultivated in St. Louis, they became large and coarse, very closely resembling the cultivated plants of the same chromosome number. Many of the more delicate [Vor. 48 334 ANNALS OF THE MISSOURI BOTANICAL GARDEN plants of lower chromosome number would seem to have greater ornamental potential than the polyploids now available. Collections from Oaxaca and Nuevo Гебп have behaved as distinctly short-day plants, while one collection from Tamaulipas (Rancho del Cielo) has proven to be relatively day-neutral. The characteristic red bracts of this species are a surpris- ingly dependable key feature, but some plants of the Missouri Ozarks lack the red coloring. 2. Poinsettia dentata. This species shows a good deal of geographic variation, which is complicated by polyploidy. The plants with n = 14 of southern Mexico are much branched and often decumbent, strongly pilose plants with very con- densed, nearly capitate inflorescence and cream or yellowish floral bracts. This is apparently the form described as Poinsettia schiedeana. These features become less marked to the north, suggesting a clinal pattern. One also finds erect, rather weedy plants with little or no bract coloration in Mexico. These are frequent in disturbed areas of central Mexico, and closely resemble the forms of this species which grow in Arizona and western Texas. It is probable that these weedy forms have entered Mexico from the north and are genetically isolated from the native form by their higher chromosome number. Some of the semi-desert forms from northern Mexico are striking because of their small, coriaceous leaves. The type of Engelmann’s var. rigidia is not the extreme in this feature. 3. Poinsettia heterophylla. This tropical species shows less variation than its temperate counterparts. Occasional populations with whitish floral bracts, strongly dentate leaves or very pilose stems suggest introgression from Р. dentata, and 1 suspect that these two do hybridize to a certain degree where P. heterophylla grows with a population of P. dentata having the same chromosome number. TAXONOMY РогчзЕттіл Graham, Edinb. New Philos. Jour. 20:412. 1836. Type: Euphorbia pulcherrima Willd. Pleuradena Raf., Ач. Локи, 1(6):182. 1833, not Pleuradenia Raf., 1825. Туре: Pleura- Girone Fl. Tell. b Cyatbopbora Ra ell. 4:117. 1838, not Cyatbopbora S. F. Gray, 1821. Type: Eu- bborbia heterophylla L : Woody or herbaceous, often perennial from a fleshy storage root; leaves alter- nate or opposite, stipules inconspicuous or absent, appearing glandular; inflorescence a condensed pleiochasium or dichasium, the branches becoming monochasial; involucral glands more or less cup-like, usually reduced in number (commonly 1); pollen grains thick-walled and coarsely reticulate; pistillate flower naked. . KEY то SPECIES 1. Plants woody, or perennial from thickened stora age roo 2 2, еп plants каал cid J meters in paca; involucre usually = than 5 . in diameter; seeds c P. PULCHERRIMA У obs plants, less ranks i meter nl involucre usually less than 5 mm. in diameter; OO лек лек m р. В 1961] DRESSLER—A SYNOPSIS ОЕ POINSETTIA 333 3. Plant ca. 1 dm. tall, ee = бон, leafless stems, the vegetative stems и а later, usually сая г less Р. RADIANS 3. Plants 2—5 . tall еб зейін EE wers borne on same stem, stems 4. ранч, bracts whites glands 6 with the outer rim flaring қысу ii а А yee ie id (w n Mex: . RESTIACEA 4. Floral, ганне a or red and green; glands red, yellow or green, the outer rim not Led and petaloid 5 . Leaves entire, markedly agen 2 (northwestern Мехісо)................... 5. P. COLORATA * Leaves more or less serrate, n arkedly paler beneat 6 6. ipee ирке scabrous-strigose; sand broadly campanulate, the inner and rims similar; seed 3—4 тт. long (western Mexico) 4. Р. sTRIGOSA 6. Foliage ИЕН gland narrow, the outer rim much thicker than wid inner eed 2—2.5 mm. long (п Bec Mexico) P. СОССТЧЕА 1. jane annual, without thickened stora 7 Без es (3) entire, about one Salf c connate, much longer than the "t (премали ay ntina) PENTADACTYLA 2. Sere рта divided, not longer than the о 8 Seeds smooth, with an equatorial row of iur tubercles and one or two era rows; E spreading, delicate plant (Peru) 0. P. INORNATA 9 8. Seeds strongly tuberculate; ge enerally соагѕег, erect p 9. Glands more or less funnel-like, the opening anro coarse tropical weed............... 11. Р. HETEROPHYLLA 10 9. Glands more or less bilabiate, the opening narrowly 10. зандаа leaves usually alternate; floral bracts ae ‘beni red; FE withou cle (North America) CYATHOPHORA 10. All | гана usually ane floral bracts green or cream at йад ор wit purple spots, but never basally red; seeds often carunculat 1 11. Seeds finely and sharply tuberculate (North America) 8. Р. DENTATA 11. Seeds coarsely and bluntly tuberculate (Argentina) P. species* 1. POINSETTIA PULCHERRIMA (Willd.) Grah., Edinb. New Philos. Jour. 20:412. 1836. Euphorbi = eiiis Willd. ex Klotzsch, Allg. Gartenz. 2:27. 1834. Pleuradena coccinea Raf., Atl. Jour. 1(6):182. 1833, not Poinsettia coccinea Dressler Euphorbia bur hd Bertol., Nov. Comment. Acad. Scient. Inst. Bonon. 4:419, t. 41. Kawa fastuosa Sessé & Mocifio, Pl. Nov. Hisp. 81. 1888. Distribution. The familiar cultivated poinsettia occurs as a wild plant in the rocky canyons of western Mexico, in Nayarit, Jalisco and Colima. Dr. Rzedowski suggests (personal communication) that it may also occur further south, in Michoacan and Guerrero. The habit of the plant, with a fleshy storage root especially well developed in the seedling stage, is characteristic of the tropical deciduous forest of this area. 2. POINSETTIA RADIANS (Benth.) Kl. & Gke., Monatsb. Akad. Berlin 1859:253. 1859. Eupborbia radians Benth., Pl. Hartw. 8. 1839. Eupborbia stormiae Croke. Rev. Seda Bat 6:13. 1939 [Type: Storm s. n. АНП. Distribution. Arizona through central Mexico to Oaxaca. represent an undescribed species. Some Seis de eno as теті бно Croiz., but that epithet is seii ra T ва E? a pran A of P. heterophylla. The plants in question are very similar to P. dentat 2 but differ 50 = in seed structure, and possi 7 iras proportions pie ah spin Mor terial and stu у needed to fix the инә of thes cis so they shall r ess for is а They аг Castellano 5. п. Dec. 5, 1927 (GH), ‘Castillon 1851 (GH), арт ва ари (GH) = Schreiter Roo [Vor. 48 336 ANNALS OF THE MISSOURI BOTANICAL GARDEN 3. PorwsETTIA restiacea (Benth.) comb. nov. Euphorbia restiacea Benth., Bot. Voy. Sulph. 162. 1844. Distribution. Western Mexico (Nayarit and Durango). Superficially the plants of P. restiacea resemble some species of Agaloma, such as Euphorbia spbaerorbiza Benth., because of the slender stems, narrow white bracts and conspicuous white glands. Though the outer rim of the gland is flattened and somewhat petaloid, there is no sharp distinction between gland and appendage, such as characterizes the members of Agaloma. The type specimen (K) is very scrappy, but a photograph (MICH) indicates that it is probably the same species as the excellent material which Dr. McVaugh has collected in Nayarit. 4. PorNsETTIA stricosa (Hook. & Arn.) Arthur, ae 11:260. 1912. Euphorbia strigosa Hook. & Arn., Bot. Voy. Beech. 310. 1840. Poinsettia bedunculata Klotzsch, Seem, Bot. Voy. Herald ee 1856. Distribution. Western Mexico (Nayarit and Jalisco). 5. РотчѕеттіА colorata (Engelm.) comb. nov. Euphorbia _ Engelm., Rep. U. 5. & Mex. Bound. Surv. 2:190. 1859 [Type: Thurber 5» Euphorbia с Rose, Contr. U. 5. Nat. Herb. 1:111. 1891, пос Е, tuberosa L., 1753. Distribution. Northwestern Mexico (Sinaloa, Sonora and Zacatecas). 6. PorNsETTIA coccinea sp. nov. Figure 2 erennis, erecta, 40—70 cm. alta, radice tuberosa oriens, caulibus basi lignes- centibus, glabris; folia alterna, linearia vel anguste lanceolata, 1.5—6 mm. lata, 40— 120 mm. longa, subtus hispidula, ad basin barbellata, apice attenuata, basi attenuata vel breviter petiolata, marginibus serratis vel subintegris; cymarum bracteae coloratae, lineari-lanceolatae vel elliptico-lanceolatae, 3.5-7 mm. latae, 30-80 mm. longae; inflorescentiae compactae; involucra anguste campanulata, 3—3.5 mm. lata, 3.5—4 mm. longa, glandulis involucrum excedentibus, 1.5-2 mm. latis, marginibus externis crassioribus altioribusque; ovarium glabrum; styli ca. 1 mm. longi, basi breve connati, fere ad medium bipartiti; semina oblongo-ovoidea, ecarunculata, tuberculata, ca. 2.5 mm. longa. Mexico. TAMAULIPAS, Municipio de Aldama, Sierra de Tamaulipas, above Juan Tomás, east of Rancho Las Yucas (ca. 40 km. nnw. of Aldama). Oak scrub and rocky arroyo, October 13, 1957, ares basally red, roots with series of thickened "tubers," R. L. Dressler 2395 (Ту , isotypes to be distributed). Other collections from the same region are: Las о Oak scrub (chaparral), peak of Cerro de Las Yucas, July 20, 1957, June 22, 1937, “Pico de perico,” М. T. Edwards 313 (MO, Абақты Y Pur crema this species). This species is related to P. colorata, but may be distinguished by the usually serrate leaves which are not markedly paler beneath and by the smaller green (rather than red) cyathia; the gland also is smaller and differently shaped. Poin- settia coccinea will more often be confused with the annual P. cyatbopbora. Under favorable conditions P. cyathophora may survive for two or more seasons, but it is 1961] DRESSLER—A SYNOPSIS OF POINSETTIA 337 Figu side view o ге 2. Poinsettia coccinea, drawn from living qe ipe f cyathium X5, top view X 6, dorsal view of see of the type collection. Habit X 1⁄2, Pi [Vor. 48 338 ANNALS OF THE MISSOURI BOTANICAL GARDEN not known to form the fleshy "sausage-string" roots of P. coccinea. Crutchfield & Johnston 5611A, from near Jaumave, Tamaulipas, is definitely a perennial, but appears to be P. cyatbopbora (no roots were present on the specimen seen). Poinsettia coccinea is frequent in rocky oak scrub and open oak forest in the region of the type locality. In this area it is quite distinct from P. cyatbopbora, which was found only in the polyploid form (n = 28). To the north of the Sierra de Tamaulipas other forms of P. cyatbopbora occur (Crutchfield, Johnston 8 McMillan 6082, Graham & Jobnston 4695, both near Padilla, Tamaulipas), and it is quite possible that introgression may occur between the annual and perennial species in this region. No satisfactory chromosome counts have been obtained for this species, but the slides which were made indicated approximately 12 to 15 chromosomes, so this probably has n — 14. 7. PorNsETTIA CYATHOPHORA (Murr.) Kl. & Gke., Monatsb. Akad. Berlin 1859: 253. 1859. Ка cyathophora Murr., Comm. Gótting. 7:81. 1786. Tithymalus cyathophorus (Murr.) Moench, Meth. 667. 1794. Euphorbia heterophylla 8 cyathophora (Murr.) Bois DC. Prod. 15(2):72. 1862. Euphorbia heterophylla forma cyathophora (Маге. Voss, Vilmorin, Blumengiártn. (ed. 3) 1:898 5: ао graminifolia Michz., Fl. Bor.-Am. 2:210. 1803. Poinsettia graminifolia (Michx.) Millsp., Field Mus. Pub. Bot. 2:304. 1909. Euphorbia barbellata Engelm., Rep. U. S. Mex. Bound. Surv. 2:190. 1859 [Type: ight s.n., GH!]. Exphorbia ОЈ var. barbellata (Engelm.) Holzinger, са Џ. $. Nat. Mer 1:216. 1892. Poinsettia barbellata (Engelm.) Small, Fl. theastern U. S. 722, 1334. 5 Вора беше а var. graminifolia Engelm., Rep. U. S. & Mex. Bound. Surv. 2:190. Poinsettia edwardsii Kl. & Gke., Abh. Akad. Berlin dn 1859:104. 1860. Poinsettia pinetorum Small, Fl. Mismi 111, 200. 19 Distribution. The eastern United States south through eastern Mexico to the Isthmus of Tehuantepec; west to Colima in Mexico. Possibly native in the Greater Antilles. Widely cultivated and often naturalized. There is a specimen from the Géttingen Botanical Garden (1794) in the Missouri Botanical Garden herbarium. While this was prepared after the descrip- tion of Euphorbia cyathophora, it probably represents the population described by ufray. Though this species is usually an annual, clonal material is easy to maintain by cuttings, which should facilitate the experimental study of its polymorphism. Under favorable conditions it may be a facultative perennial. I have seen specimens from northeastern Mexico and southern Florida which had over-wintered at least one season before collection. 8. PorNsETTIA DENTATA (Michx.) КІ. & Gke., Monatsb. Akad. Berlin 1859:253. Euphorbia dentata Michx., Fl. Bor.-Am. 2:211. 1803. Anisophyllum dentatum (Michx.) aw., Syn. РІ. Succ. 162. 1812. Euphorbia herronii Riddell, Syn. Fl. W. States 32. 1835. 1961] DRESSLER—A SYNOPSIS OF POINSETTIA 339 Euphorbia dé bac В rigidia Engelm., Rep. Џ. 8. & Mex. Bound. Surv. 2:190. 1859 [Type: Wrig 1 Euphorbia Рафа y cuphosperma Engelm., Кер. Џ. S. & Mex. Bound. Surv. 2:190. 1859 [Type: Wright 1834, МО!]. Euphorbia ақ sadly (Engelm.) Boiss, DC. Prod. 15(2):73. 1862. Poinsettia cupbosperma (Engelm.) Small, Fl. Southeastern U. S. 721, 1334. 1903. Euphorbia dentata forma cuphosperma (Engelm.) Fern., Rhodora 50:148. > Poinsettia schiedeana К]. & Gke., Abh. Akad. о жемей 1859:102. 1860. Euphorbia dentata В lasiocarpa Boiss., д ee 15(2):72. 1862. Euphorbia dentata у linearis Engelm. ex. Boiss., DC. Prod. 15(2) :72. 1862. Euphorbia dentata var. gracillima Millsp. » үте 2:90. 1 Euphorbia dentata var. lancifolia Farwell, Am. Midl. Nat. 8: 273, 1923. Distribution. The eastern United States west to Arizona and south through Mexico possibly to Guatemala. To be expected as a weed elsewhere. 9. Ротмветтіл pentadactyla (Griseb.) comb. nov. Euphorbia pentadactyla Griseb., Gött. Abh. 24:63. 1879. Distribution. Paraguay and northern Argentina. 10. Рогмзеттіл inornata sp. nov. Figure 3 Annua, ramosa, 8—14 cm. alta, ramisque procumbens ad 12 cm. longa, caulibus sparse pilosis; folia alterna vel ea basalia opposita, ovata vel elliptica, 8-16 mm. lata, 12-32 mm. longa, subtus sparse hispidula, apice acuta, basi cuneata, mar- ginibus leviter serratis et hispidulis, petiolis 2-6 mm. longis; cymae dichotomae, involucris late campanulatis, са. 2 mm. longis, 1.2-2 mm. latis, glandulis 0.6-0.8 mm. latis, ostio oblongo; ovarium glabrum styli ca. 1 mm. longi, basi breve connati, fere ad basin bipartiti; semina quadrato-ovoidea, minute carunculata, sub-laevia, paucituberculata, 2.5 mm. longa. + ШЕ Бері & Prov. Lima, Dist. Pachacamac; Atacongo; among rocks in arid valley, alt. m., common, scattered, October 14, 1935, Ynes Mexia 04044 (Type: MO, isotype GH, и. also at UC). This is apparently a deep-rooted, spreading annual. The plant would pass superficially for a small delicate P. heterophylla, but the gland is quite different in shape, and the nearly smooth seed with an equatorial row of small, blunt tubercles, and fainter basal and distal rows, is unlike that of any other species of Poinsettia. 11. PorNsETTIA HETEROPHYLLA (L.) КІ. & Gke., Monatsb. Akad. Berlin 1859: 253. 1859 Euphorbi terophylla L., Sp. Pl. 453. 1753. Tithymalus heterophyllus (L.) Haw., р T т са 141 1811. " Cyathophiid heterophylla (L.) Raf., Fl. Tell. 4:117. 1838. и сы Lam., Епсус. Meth. Вог. aia 1788. Euphorbia heterophylla var. elliptica (Lam.) O. Қал. Кеу. Сеп. 2:605. 189 тыны қане) Vahl, Symb. Bot. 2:53. 1791. "вео heterophylla var. linifolia (Vahl) O. Ktze., Rev. Gen. 2:605. 1891. А Ei , Hort. Mat. Dec. 18. 1797. Poinsettia geniculata (Ort.) Kl. и есеи sie Berlin 1859:253. 1859. вика heterophylla е geniculata (Ort.) Gómez, Anal. Hist. Nat. Madrid 25:46. : fots l Hort. Schoenb. 3:15 . 1798. Tithymalus prunijoli prid I TUN Suc " с 143: 1512: Poinset bia jaan (Jacq.) Ki. & Gke., Monatsb. Akad. Berlin 19592755: 1859. (Мог. 48 340 ANNALS OF THE MISSOURI BOTANICAL GARDEN M frangulaefolia HBK., Nov. Gen. et Sp. 2:62. 1817. Poinsettia frangulaefolia & Gke., Monatsb. Akad. Berlin 1859:253. 1859 debe packet Ж Rich., La Sagra, Hist. Cuba 9:198. 1850. Euphorbia morisoniana Kl., Seem. Bot. Voy. Herald 100. 1853. Poinsettia morisoniana Ў 5% Monatsb. Akad. Berlin 1859:253. 1859. Poinsettia ruizian Kl.& Gke., Abh. Akad. Berlin Phys. 1859:102. 1860. ?Euphorbia be wein minor Boiss., DC. Prod. 15(2):73. 1862. Euphorbia heterophylla var. brasiliensis Müll. Arg., Mart. Fl. Bras. 11(2):695. 1874. Euphorbia prunifolia var. a repanda Müll. Arg., Mart. Fl. Bras. 11(2):694. 1874. Euphorbia prunifolia var. y angustifolia Müll. Arg., Mart. Fl. em o 695. 1874. Poinsettia havanensis ваше Fl. Southeastern U. S. 7 1334. 903. Eupborbia aureocincta Croiz., Jour. Arnold Arb. un Ht 1943 [Type: Rojas 3379, AH!]. Distribution. From Arizona and Tamaulipas southward throughout the Amer- ican tropics. As a weed throughout the tropics. Naturalized in Louisiana and Texas and to be expected elsewhere. EXCLUDED SPECIES Poinsettia eriantha (Benth.) Rose & Standley, Contrib. U. S. Nat. Herb. 16:13. = Euphorbia (Agaloma) eriantha Benth. This species, E. (A.) exstipulata Engelm., E. (A.) jaliscensis Rob. & Greenm., and E. (A.) lacera Boiss. are all closely related. All of these species have small but distinct petaloid appendages on the glands and have seeds which are unlike those of Poinsettia. In E. (A.) eriantha, at least, the pollen grains are very unlike those of Poinsettia. Euphorbia heterophylla var. eriocarpa Millsp., Proc. Cal. Acad. П 2:230. 1889. The type (F!) is not a Poinsettia, but appears to be an Agaloma with reduced gland appendages, or possibly a Tithymalus. It is probably unnamed as a species. Poinsettia insulana (Vell.) Kl. & Gke., Monatsb. Akad. Berlin 1859:253. 1859. = Euphorbia (Agaloma) insulana Vell. This and the following two species are closely related, if not conspecific. Poinsettia lancifolia (Schlecht.) Kl. & Gke., Monatsb. Akad. Berlin 1859:253. 1859. = Euphorbia (Agaloma) lancifolia (Schlecht.) Poinsettia oerstediana Kl. & Gke., Abh. Akad. Berlin Phys. 1859:103. 1860. = Euphorbia (Agaloma) oerstediana (Kl. & Gke.) Boiss. Poinsettia punicea (Sw.) Kl. & Gke., Monatsb. Akad. Berlin 1859:253. 1859. = Tithymalus puniceus (Sw.) Haw. Poinsettia xalapensis (HBK.) Kl. & Gke., Monatsb. Akad. Berlin 1859:253. 1859. = Euphorbia (Agaloma) xalapensis HBK. REFERENCES Dressler, К. 1. 1957. The Genus Pedilanthus (Euphorbiaceae). Contrib. Gray Herb. 182:1-188 Perry, В. A. 1943. Chromosome Number and Phylogenetic Relationships in the Euphorbiacese. Amer. Jour. Bot. 30:527-543. Plukenet, L. 1696. Almagestu: ondon. Wheeler, L. C. 1943. The Genera of the Li li ыы "Amer. Мі, Nat. 30:456—503. 1961] DRESSLER—A SYNOPSIS OF POINSETTIA 1 Figure 3. Poinsettia inornata, drawn from the type specimen. Нађи Х%, cyathium X10, dorsal view of seed X 8. GENERAL INDEX TO VOLUME XLVIII New scientific names of бан and the final members of new combinations аге printed in bold face type; synonym italics; and all other 5 іп ordinary type and page numbers having reference to figures and plates, in Achyranthes, 22; altissima, 19; argentea, = aspersa, 23, 24, В indica, 24, [var implex, 24; corymbosa, 105; halimifolia, ree birtiflora, 27; indica, 24; la 4; le ii 5 43; polygonoides, 40; Prost ар = radi- cans, 41; repens, 41; sicula, steno- ta 44; гиены ВЕ 36 ; и теме Ааа 2 276 Adena ,9 Adenosramma a eas 39 Adenon oon zoaceae of E em 80 bersia, 13; gracilis, 14 lezentbera forskalli, 39 lochlamys, > > 91 sine, 90; м 194 = , 97 РЕКЕ ORE ы = > M сан ес тық 38; achyranth., 41; amoena, 41; asterotricha, 42; bettzickianna, 41; crucis, 42; denticulata, 39; ficoidea, 41; ficoidea, 40, var , flavogrisea, 42, у halimi- folia, 42; flevogrisea, 42; јан lia, 42; laguroides, 44; lebmannii, 45; linearis, 33; 40; parvifl 41; philoxeroides, 43; polygonoides, 40, l ide E-B glabrescens, epens, 41; sessilis, 39; steno ; tenuissima, 39; triandra, phylla, 39; LT 41; villiflora, 41; william- sii, 4 Amaranthaceae of Panama, 6 Amarantbus, 11 Amaranthus, 12; I MMC mé cali- fornicus, 14; caracasanus, ; carneus, 14; chlorostachys, 16, var. өзін 16; > 6 acus 16, 2 мени 16, [var.] pani- culatus, 15; bypocondriacus, 16, incomp- 5; ; laetus, 16; paniculatus, 15, P йк оны: 15, а рит sis, 15; purpureus, 12; retroflexus var chlorostachys, 16, var. bybridus, 16; sanguineu eciosus, ; Spinosus, у flex sta че 16, В боена 16; viridis, 14 2. 47; globosus, 49, [уаг.] albi- Рени а, Ambrina, 2; а 5; andicola, 5; antbelmintice; 5; chilensis, denudata, 5 Ammodenia, 90 Ancistrocar pus, z Ancistrostigma Anemopsis porr A 107, 109, 110, 113, Anisophyllum dentatum, 338 92; confusa, 93; diffusa, 92; qi ogee 93; jussiaei, 92; саан subsp. guatem alensis, 93, i sp. saxosa, [ 93; va i diffusa, 92, В diffusa, 92, а genuina, 92; leptophylla, 208; mearn 95; m ginea, 194; nemorosa, novograna- var. cineras- ға Atriplex, ambrosioides, 5, anthelmintica, 5; muralis, Aylmeria, 104 Bajan, 12 Ballarion, 97 Batidaceae of Panama, 66 y 12 Blitum ambrosioides, 5 (343) 344 eee Ка ие, 36; repens, 36 ; adscendens, па, ече” 52; caribaea, 54; diandra, 54; diffusa, 53, 54; elongata, 52; erecta, 52, 53; hirsuta, 54; ramulosa, 54; squamata, Botrydiu Botrys, 2; и 5; anthelmintica, 5 кы Bragantia, 47 ла. техісапа, 45 Brewerin Bucbolzia ficoidea, 41; philoxeroides, 43; polygonoides, M B diffu usa 41, а erecta, 1, Y relitent; Macon George S., z James A. Duke: San- ango: New Amazonian genus of Logani- aceae, 269 C нин, з е» 24, sicula, 24 aipa Schor R одын етоп, 35; ermicularis, 36 не ве айы of Panama, 90 Celosia, 11; argentea, 11, 12; coccinea, 12; cristata, 12; huttonii, 12; жер 12; marilandica, 12; pallida, 12; pan lata, M s _bleiogyna, 10; Я: би BA aig 2 aspersa, 24; indica, 24 Centunculus Cephalo tomandra, 55; fragrans, 55, 50; фапатеп Cerastium, 95; dosis 95; glomer- atum, s h nemorum, 97; viscosum, 95, ‚ B consanguineum, 95 MA. grandiflorus, 169; oculatus, 170 Ceratococca, 7 Chamissoa, 19; “alte ssima, 19, 20, subsp. albo- p 21, var. deniipaniculata, 21, var. B = 4 semispicata, 21, var. rubella, flora, 21, B ире zu celosioides, 22; macrocarpa, 19; 21; maximiliana, 22, var. B procu Moreni 22, var. pubescens, 22, f. celosioides, 22 Салор of = Chenopodi 1; banca a 5, [var.] anthelminticum, 5; anthelminti 5; carthagenense, 2; caudatum, 14; chilense, [Vor. 48 ANNALS OF THE MISSOURI BOTANICAL GARDEN 5; guineense, 2; leptophyllum var. lepto- phylloides, 3; murale, 2 albescens, 2, y HAT 2, [var.] spissidentatum, 3; obovatum, 5; ; petiolare var. leptophy l- 5; retusum, Cruzeta crassifolia, 36 "Cucumer marinus," Cucurbita "foliis asperis size Zucha flore luteo," 305; pepo, 302, 303, 305, 324 Cucurbitaceae, 2 Cyatbopbora, 2t ciliata, 338; heterophylla, 339; picta, неба — ҚЫ schyranthoies 27, Ү densi- flora, 27, B glabrescens, 27; geniculata, 25; prostrata, 25, Ж В achyranthoides, 27 Desmochaeta achyranthoides, 274 densiflora, 27; prostrata, 25; uncinata, . P. Frymire: Prelimi- nary studies in the genus Stanhopea (Orchidaceae), 137 Doerriena, 95 Doerriera, 95 Dolophragma, 91 Dressler, ге L.: A Synopsis of Poinset- 329; Index of Drymaria, Preliminary = of, 47 gern citation о 17 "di ser. arenarioides, 178, ; ser. сона 179, 245; ser. debiles, 179, 220; ser. divaricatae, 179, 240; ser. excisae, 179, 201; ser. fasciculatae, 179, 16; ser. Eruceicentes, 179, 212; ser. grandiflores, 179, 230; ser. holosteoides, stipitatae, 178, 193; ; ser. viscosae, 178, 200; ser. villosae, 179, 223; adenophora, 103, 251; adiantoides, 1961] INDEX 345 254; agapatensis, 242; anomala, 210 211; 241, 244 cidula, 247, 243; elata, 198, 100; e effusa var. бой fusi; 205, 207, var. depressa, 206, 207, var. effusa, 205, 207; engleriana glaber 5 4; nulosa var. йен, im vai: сс 246, 249, var. gla loni; 246, 24 perennis, var. crassifolia, 187, var. holo- ње 186, 187; hypercifolia, dua 203; bypercifolia, 250; еі жеш ді; ; Tobis- stonii, 198; ladewii, 2 252; Ae 221, 245; leptoclados, 2 var. ‘peruviana, u 194, $2; multiflora, 224, 225; nitida, ii ee acillima, 208; MM a, v ortegioides, 200, 203; yes 233; oxalidea, 254; pachyphylla, 787, 188; palmeri, 254 ; palustris, 227; papo peninsularis, 198; peruviana, 254; poly- carpoides, 197, 199; polystachya, 220, var. diffusa, 220; praecox, 218, 219; pro- cumbens, 103, 251; ramosissima, 247; 2r 3; Я а, 214, var. ин 213, 15: tepicana, 102, 226; townsendii, 227; veatcbii, 186; vilióis, IOI, subsp. palus- tris, 227, 220, var. perennis, 228, 220, townsendi, 228, 220, subsp. para- morum, 229, 230, subsp. villosa, 225, 246, +; tepicana, 227; virgata, 234; vis- cidula, 243; viscosa, 200, 203; weber- baueri, 238; xerophylla, 249, 250 Dufourea, 91 Duke, James A.: Amaranthaceae of Pana- nary Revision of the Genus Drymaria, 173; —— & George S. Bunting: Sanan- go: New Amazonian Genus of Logani- aceae, 269 E Eggersia, 57 Eisendrath, Erna Rice: Portraits of Plants. A Limited Study of the "Icones," 291 one, 9 Euphorbia aureocincta, 340; barbellata, 338; colorata, 336; cupbosberma, 339; cyat tho- phora, 338; dentata, з уаг. оне 339, уаг. lancifolia, 339, у cuphosperma, graminifolia, 338; herronii, 338; hetero- phylla, 331, 334, 339, var. barbellata, 338, var. caius 340, var. elliptica, 339, var. erioca шқ май gramini- folia, 338, var. limifolia a, 339, B cyatba- fora, 538, B elliptica f. ты 340, Е geniculate, 339, 6 minor, 340, athophora, 338; insulana , 340; jaliscensis ia, ini- 340; кын. 339; prunifolia, $e var 'y angustifolia, 340, var. a repan 340; pulcberrima, 334, 335; radians, 3551 restiacea, 336; stormiae, 335; strigosa, 346 336; trachyphylla, 340; xalapensis, 340; zonosperma, 340 Euphorbieae: 330 Euthalia, 91 Euxolus, 12; caudatus, 14, В gracilis, 14, Y maxim Ewart, R. Bradley: Two ped Members of the Genus Scolecopteris, 27 Eoangiopteris, 276 F Faccbinia, 91 Fevillea сова оци, A des Frymire, G. P., & C. H. Dodson: Prelimi- nary Studies in cs Genus Stanhopea (Orchidaceae), 137 Fuchs' De Historia stirpium, 299 G Galiastrum, 8 — 13; и 16; patula, 15; spi- оза, ба + ка Generic 2 in the subtribe Stan- hopeinae, I Glinus ifo: 80 Glomeraria, 12 Gomphrena, 47; aggregata, 36; crassifolia, decumbens, 48, f. simplex, 48; polygo- noides, 40, 41; sessilis, 39, vermicularis, ~ pa Se decipiens, 28, hookerianus, 28 Gouffeia, 91 Grenier. aN 9 1 Guapira, 61; costaricana, 62, 63; standley- ‚ 62 ana ле chinensis, 107; involucrata, 107, ІІО Gynastrum значана 90 Hagaea, 104 Hagea, 10 Надан thus, Halimus, i pe ortulacastrum, 84 Hebantbe, 27; decipiens, 28; hookeriana, 28; paniculaie, 29; virgata, 29 Helianthemoides, Hepaticites, 126; devonicus, 126, 128, 120, 131 tuberosa, 336; key to New World genera, [Vor. 48 ANNALS OF THE MISSOURI BOTANICAL GARDEN Holosteum cordatum, 103, 251; diandrum, > Honkenya, 90 Houttuynia cordata, 107, 100, IIO, 112, Hueber, Francis М.: Hepaticites devonicus, A New Fossil зен from the De- vonian of New York, 12 Hyala, 104 Hylebia, 97 Icones: A limited study, 291; botanical use f the word index of icone 5 of key u ence 3 species of the REER as limiting factors in study, 296; later work in vari- ous techniques, 314; lithographs, 313; m ове source and used by several uthors, lecebrun LE tha, 41; ficoideum, 41; frutescens, 42; indicum, 39; limense, 42; bo olygonoides 40; sessile, 39; vermi iculs- ,36 М2 or Orchid Names—1960, 133 36; Ed 33; Mp cria 34; celosioides, 3 ana, 34; fered re, 34; Баланы 34. a ЈЕ оре- сито 4, В effusa, 34, у verticillata, 34; калышы, 33; venil де 36; verti- сари. 34 Kokera, 19, acuminata, 22, celosioides, 22, panicu ulata, 2 Krascheninikovia, 97 x i ms SNR Labay Pagina 105 Lemia, 88 Шы ди E ennai sce —— И 1961] INDEX 347 onum se ысын 194 um Lithophila vermiculata, 36 Loganiaceae: € ew Genus, 269 Lopboxera, 1 M Malachium пы 9; sprucei, 9 Mengea, 13; californica, 14 Merckia, Micropetalon, 9 Mictopetalums A Microtea, 73; debilis, 73, 7 "fg а ovata, 74, В rhombifo lia, 74; ovata, 74 Mollie à ve wd 05 ollugo, 80; arenaria, 81; axillaris, 81; icbotoma, 81; diffusa, 81; пена. 81; hoffmannseggiana, 81; juncea, 81; schrankii, ; spergulaefolia, 81; tri- lla, 81; Vern Mollugophytum, 100; crassifolium, 187; офа 186: ойга, 194 Morocar pus Morphology aa Anatomy of the Sauru- Sun Floral papa a and Embry sie BY; MIR. 97 Myosotis, 95 Myosoton, 96 “Мапа ође à feuilles de lierre,” 325, 32 26 laetevirens, 58, 50; orosiana, 60; pittieri, 60; psychotrioides, 58, 60; pycnantha, 60; urophylla, 60; xanthina, 58 Nevling, Lorin I., Jr.: Aizoaceae of Pana- ma, 80; Portulacaceds of Panama, 85 бананы of Panama, 51 a 91 Oligan. ie Oligant. Orchid дай Index, 133 Orthospermum, 2 d —— 2; ambrosioides, 5; suffruti- Viris 2 Pallavia, ~ Papularia, Раја. achyrantha, ha ficoidea, 41; frutescens, 42; sessilis, 3 » 324 “Реропев rotundi,” Petiveria, 66; ДЕ 67, 68, B EH 68, у oct nudos. 68; foet ids, 68; dria, 68; ochroleuca, 68; наиме 68, paraguayensis, 68 Pettera, 91 en 27; rever 29; grandiflora, 28, ookeriana р 2 hookeriana, 28; paniculata, 29, Phar m arenarium, 81; cervia о 81; нения аа ғанда, 35; "Mendes 36; crassifolius, 6; vermicularis, 36, 37 Proh 68; acuminata, 71, 72; bogoten- ici hove: spade 71; рне 72; ‘poly styla, 72; purpurascens, 71; rivinoides, 72; rugosa, 60,70; sessiliflora, 71; trique- га tra, Phytolaccaceae of Panama, 66 Piercea, 74; acuminata, 76; glabra, 76; obliquata, 76; tomentosa, 76 Pinosia, 100, 178; ortegioides, 200 Mud 68 a, 64; кен <> 65; Джек руй 4; lorant siei onotaxadenia sie Aes i, 64; o ш НЫ нуға Еа 41 Planchonia, 1 Pleuradena, 334; coccinea, 334, 335 Pleuradenia, 3 Pleuropetalum, 9; calospermum, 9; costa- ricense, 9; pleiogynum, 8, 10; sprucei, 9; standleyi, 10; tucurriquense, 91 to species, 334; patterns of 0 ariation, 3 Poinsettia, 334; barbellata, 338; coccinea, 348 36, 337; colorata, 336; cupbosperma, 339; ee 337, 332, 335, 338; dentata, 331, 334, 338; edwardsii, 338 eriantha, 340; ам 340; genic- ; bavanensis, 3353 340; schiedeana, 339; strigosa, 336; xalapensis, Polia, uie Polycarpa Tak 105; corymbosa, 105 Polycarpaea, 104; atherophora, 105; brasil- var. ramosissima, 105; brevi- folia, 105; с» > 7 05; densiflora, 105; filifolia, p Ms 105; spadicea, 10 P Portraits of Pans. A Limited Study of the "Icones," 291 Portulaca, ^3 marginata, 88; oleracea, 88, 80, а macrant 88; patens, 86; portulacastrum, 84; 86 flexa, Pártulicacede of Panama, 8 Portulacastrum, 82; monogynum, 82 Portulacca. е. Potamophila, 7 Prelim inary Revision of the Genus Dry- ma Prelimin: ry Studies in the Genus Stanhopea ( Orchidaceae), 137 Prevoi сване $ M p^ Ptychocarp: Pupalia demon, 27; prostrata, 25 Pyxidium, Pyxipoma, У Quaternella, 95 R Racom Raeder, a Phytolaccaceae of Pana- fis. М. V. S.: Morphology and Anatomy [Vor. 48 ANNALS OF THE MISSOURI BOTANICAL GARDEN of the Sauruaceae. I. Floral Anatomy and Embryology, 107 eme, 82 Rhodalsine, 91 Rivina, 74; acuminata, 76; americana, 77; eurantiscá, 76; brasiliensis, 76; canescens, 76, var. orientalis, 76, а canescens, 76, У 77 mutisii, 77; peer 76; a 77, 76; pal ere 79; Pede 76; viridiflora, 76; viridis, 76 Rivinia, 74 Robbairea, 104 Rocam поен patens, 86 Roemeria Rosea, 31; elati Rouhieve, 2; estis S Sabulina, 9 Sanango: New Amazonian Genus of Logani- aceae, 26 Sanango: жй ion, 273; anatomical con- siderations, 174; pollen, 273; trichomes, 274; vesture, 273 Sanango durum, 270, 271 Sarcoca, 68 Sarratia, 13 Saururaceae: Floral Anatomy and Embry- , Saururaceae: anatomy of the flower, 111; carpels and сыы 114; yes 118; embryology, 114; flower, 108; fru 118; inflorescence; 108; ае 123 кнн сетив ID F ее 107, A 110, 115 Зена еб Schollera, A bl 74 Scleropus, 1 баари Two new members of the EZ eiie er 275, 276; elegans, 275; illino- ensis, 281, 28 ‚ 288, 289; incisifolia, 275; iowensis, 275; latifolia, 275; major, 275; 75; subelegans, 275. Serturnera, 27 suvium, 84; ortegae, 84; pedunculatum, 84; portulacast trum, 84, 85; revoluti- 19611 INDEX 349 folium 84; sessile, 84; sessiliflorum, 84 Siebera, 91 Solanoides, 74; laevis, 76; pubescens, 76; = 6 a, 91 Sperguletram, 91, 96; lanuginosum, 92 Squi Stacbyarpagopbora, 22; aspersa, 24 the genus, 162; taxonomy, 164; variation, ee amesiana, 169; amoena, 170 anfrac 170; annulata, 171; atropur- у еј 1 aurata, 170; aurantia, 170; aurea, 170; bicolor, 171; bucephalus, 743, 5, 170; calceolata, 169; calceolus, 169; candida, 1 cavendishii, 7a; cirrhata, 146, 169; connata, 157, voluta, 1 169; costaricensis, 170; eee 170, deltoides, 171; dorn. 17 > A 5; > ~ fregeana, 17 1; fuerstenber gine, 171; gib- bosa, 171; grandiflora, 153, 169; grandi- flora, 170; graveolens, 169, 170; guttata, 170; guttulata, 170; haseloviana, 171; hernandezii, 171; koppi. 171; implicata, 70; langlasseana, 169; lewisae, 155, 169; lietzei, 170; lindleyi, 170; lowii, 169; упсеа, 170; сын, 170; maculosa, 170; жайыш: 171; жәні; 170; 171; nigripes, 171; migroviolacea, 170; шаға, 103, 170, var. constricta, 170; odoratissima, 170; ornatissima, 170; jan 149, 1 150, 170; uncinata, 171; velata, 170; venusta, 170; violacea, 171; «wallisii, 171; wardii, 167, 170; warscewicziana, 170; xytriophora, 171 oo crepe 169 Steiremis repen Stellaria, 96; y Rd ER 251; baldwini, 97; ciliata В cuspidata, z cuspidata, 97; ‚ 254; francheti, nemoralis, 97; nemorum, 97, subsp. glo- chidosperma, 97, subsp. montana, 97, f. reichenbachii, 97, var. japonica, 97; ovata, 98; reichenbachii, 97; rotundifolia, 254; virgata, 234 Strophium, 91 Sturiella, 276 Sukana, 11 T Talinium, 86 T. 2n chrysanthum, 87; dicboto- mum, 86; fruticosum, 86; moritziana, 87; wn Pid i 6, var. sarmentosum 87; patens, 86, var. sarmentosum, 87; bur- putem. 86; seflexum, 9 , f. sarmento- sum, 87; т 87; о pis 86; а latu 6 Telanthera ers, 4 m 4 22 ; flavogrisea, 42; fru- acutifolia, 42; halimifolia, crocebbale, 45; philoxeroides, 43, у“ aeri ote 43, 7 den- ticulata, 45, tusifolia, 43, var. pbyl- € А polygonoides, 40, B bracbiata, usd, „о, ыы ИЙ, 41 Tithonia, 74; humilis, 76, var. canescens f. albiflora, 76, var. glabra, Tithymalus cyatbobborus,. 338; heterophyl- lus, 339; prunifolius, 339; puniceus, 340 gin 61; costaricana, 62, 63; banamen- Trianthema, 82; kae 82; monogynum 2: portilacastrum, % 2, 83; procumbens, oe 77; йет бр 77; polyan- m, 78, 79; rivinoide 5, Tr ние 91 Tryphane, 91 = Valeriana latifolia, 5 Villamilla, 77; ade. 77; a 79 Vulvaria, 2; tracbios sperma, W Wadapus, 47 Wrerzbickia, 91 Wilhelmsia, 91 X Xeraea, vi decumbens, 47; globosa, 49 Xeralsine, X erondra, i ны 34; elatior, 33 Xerea pani iculata, Xerosiphon, 47 Zaleia, 82 Zaleya, 82 Zallia, 82 MISSOURI BOTANICAL GARDEN STAFF Director .Furrs W. WENT AR ANDER юным D. DWYER, ve Кыш of setal Plants Research Associate Y М. ANDREYS, RAYMOND FREEBORG, ; та 5 Research Associate Новы C. CUTLER, | Norton Н. NICKERSON, Executive Director 5 Меде logist 4 CARROLL hs Бойс, ; TRIFON SCHRENK. a а еа Curator Museum oF CALAWAY DSONy қате КӨН Taxonomist and Ситни of Jur OWEN: % SEXTON, . МУ iig Plants Research Ecologist 5 OBERT L. DRESSLER ? R о леви Ма Editor GEORGE es Хам x бондлок, PEE di ALS £5 ; х ee ; 5% “Шы ; james А. Du KE Assistant Curator of the Herbarium BOARD [5 TRUSTEES : ; President. 3. | Vice-President | _ LEICESTER В. алыс fe Second Vice-President |