K/ dese -5 ig f D9VEF GRAY HERBARIUM OF HARVARD UNIVERSITY CONTRIBUTIONS Nos. 179-184 1956-58 ouN! BoTANicaAly GARDEN LIBRARY oster er Fe Cc. re rs oF Aa ; 4 ; ey ; yet Sy \ and . % ae ONS FROM + ; 4 UTI OF REVISION A Py ot fi " ¥ a i> . f ae %. =! ss vs : ste : aS oan teh Be t pa os ; S 5 7 c ae > 4 ; : Seen 4 ee ° aoe ieee Bib g sy " ’ % eae CONTRIBUTIONS FROM THE GRAY HERBARIUM OF HARVARD UNIVERSITY Edited by Reed C. Rollins and Robert C. Foster NO. CLXXIX A REVISION OF THE AMERICAN SPECIES OF NOTHOLAENA By Rotita TRYON MISSOURI BOTANICAL GARDEN ILLUSTRATED BY BERNADETTE VELICK Published by THE GRAY HERBARIUM OF HARVARD UNIVERSITY CAMBRIDGE, MASS., U.S.A. 1956 Issued October Ist. SG. A; Ballard). Weatherby working on Notholaena at Kew in July, 1939 (photo F. PREFATORY NOTE MR. C. A. WEATHERBY’S WORK ON NOTHOLAENA. Una F. WEATHERBY In 1935 we visited some of the larger herbaria in Holland and Belgium and also the ones in Paris and London, studying and photographing the type specimens of various ferns Mr. Weatherby was interested in, especially some of the Notholaena group. After our return a more careful study was made of the specimens of this group as found in our American herbaria. In 1937 we returned to Europe for more study of the types and this time we added those in the herbaria of Berlin, Vienna and Prague. By 1939 he had decided upon a complete revision of the Notholaena group, saying many species were masquerading under that name but did not belong there. Though there was much talk of war we decided to go back to Europe and get as many notes and pictures as possible for fear that if there was a war the herbaria might be destroyed. We spent some time in London and in Paris where we were when the war began. Trunks and baggage were left behind but the notes and photographs we carried in our hands and got them safely home with us. During the war years he felt it was his duty to devote his time to any war work he could do. He put aside his work on No- tholaena though he made numerous notes on it and charts of each species or related groups for comparison. After the war the work was again interrupted while he did some nomenclatorial work for the International Botanical Congress. He disliked purely nomenclatorial work, but having been given the job he felt it his duty to do it. He said that plants like people must have names if one was to talk about them intelligently and it was necessary for the name to be accurate to prevent confusion. So at his sudden death he had a large pile of notes, charts, etc. concerning his Notholaena monograph but many matters were still undecided. These I turned over to Dr. Tryon who came from St. Louis to help me place some of Mr. Weatherby’s botanical possessions. After several years he has succeeded in working these up for publication. All taxonomic decisions are purely his. He has done a marvelous job in bringing order out of the mass of notes and the excellent drawings prepared by Mrs. Velick add greatly to their value.—Cambridge, Massachusetts, March 18, 1956. 4 ROLLA TRYON INTRODUCTION Some time after the death of C. A. Weatherby in 1949, Mrs. Weatherby turned over to me his manuscript and notes on Notho- laena. A careful perusal revealed that the treatment was fairly well along toward completion, to the extent, at least, that it was certainly not worthwhile for one to begin a similar study de novo, and accordingly it was decided, in consultation with Mrs. Weatherby, that the study should be completed. However, Mr. Weatherby’s own wishes did not allow the monograph to be published under his authorship, since he had not finished it to his own satisfaction. So I have accepted the difficult position of pub- lishing what is principally the work of another, appreciating on the one hand the reasons for this, and, on the other, the necessity of making what would have been Mr. Weatherby’s major publi- cation in taxonomy available to fern students and floristic bot- anists. In a number of smaller papers, Mr. Weatherby had already used portions of the manuscript and in these seventeen species were fully treated and fifteen more partially so. The appropriate reference is given after each species to his own publications on it. The remainder of the manuscript I have checked and elaborated when necessary, the keys have often been recast, two new species that Mr. Weatherby did not see material of are described and finally the treatments of the last two species have been added, since they clearly belong with the ones immediately preceding. In the taxonomic treatment I have been guided in difficult cases by Mr. Weatherby’s annotations at the Gray Herbarium and by the general mode of treatment in his publications. In all cases the judgment is my own although I have made every effort to follow the manuscript and in the cases where a species was dealt with in: one of his papers his treatment has been essentially copied. Maps and illustrations have been furnished for the species and it is hoped that these will increase the usefulness of the paper. The meaning of the different symbols used on the distribution maps is as follows: a solid dot is used for a specimen that is from a particular locality, a circle is used for a specimen with only general locality data and an X is used when the record is taken from the literature. In the latter case the source of the report is given in the discussion of the species concerned. My work has proceeded more slowly than I would have liked but most of it was completed in 1954. The necessary time needed THE AMERICAN SPECIES OF NOTHOLAENA 5 to put the final touches on the treatment has been supported by the National Science Foundation in relation to work on the Ferns of Peru; some twenty-five percent of the species occur in that country. Mrs. Una F. Weatherby has generously supported the cost of preparation of the illustrations and of the plates. I am indebted to Mrs. Bernadette Velick for her accurate and artistic work on the illustrations. Tue Genus NOTHOLAENA There need be no apology for a lack of a definition of the genus Notholaena. The difficulties of such a definition are well known to fern students and are in fact perhaps the principal reason for the present study. It is not likely that an adequate classification of the genera of the Cheilantheae, Cheilanthes, Notholaena, Pellaea, Doryopteris and related smaller genera, can be achieved without a previous knowledge of the species concerned and it is in this manner that this paper is a contribution to an understand- ing of the genus. It seems that the basic problem will be to determine the lines of evolution so that the meaning of characters is sufficiently clear, which characters are of generic value, which are convergent and which are only of specific expression. The group of species, included here in Notholaena, grades sus- piciously into Cheilanthes on the one hand (via N. peruviana, N. brachypus and a few others among species nos. 1-18) and into Pellaea on the other (via species nos. 44-58). The first mentioned species are similar to Cheilanthes in having a slightly to moder- ately developed margin-indusium and the others are similar to Pellaea in rhizome scales and their imparipinnate lamina. The last two species in this treatment, N. parvifolia (Pellaea micro- _phylla) and N. formosa, have not been placed in Notholaena previously, usually residing in Pellaea, but their obvious affinity with the species preceding them necessitates their transfer. It is unfortunate to make new names under Notholaena when the generic situation is in such doubt but if they were maintained in Pellaea other species that would necessarily go with them would require new names and these would have to be made with equal doubt. The uncertainty of the definition of Notholaena is not all of the difficulty, for if Cheilanthes and Pellaea were well defined the task might be more certain, but the problems in those genera are, in their own way, as perplexing as those in Notholaena. Characteristically, Notholaena has a flat unmodified margin 6 ROLLA TRYON although sometimes it is revolute and sometimes it is modified into a marginal indusium. The species with wax may be distin- guished from Pellaea by that character and from Cheilanthes by the unmodified margin. The species without any indument (N. Jonesii, N. Lumholtzii, N. nivea var. tenera, N. parvifolia and N. formosa) may be distinguished from both genera by their un- modified margin, although they may resemble closely some species of Pellaea. The fact that N. nivea has a variety lacking indument demonstrates that this character alone is not sufficient to place a species in a genus other than Notholaena and such species are included here on the basis of their close relation in other characters to wax-bearing species. The species with hairs or scales (some also have wax) may be distinguished from Pellaea by their indument as well as by their usually unmodified margin. The latter character separates them from Cheilanthes. The few species with a slightly to moderately developed marginal indu- sium are transitional to Cheilanthes but retained here because of apparent relations to species without a modified margin. Although Mr. Weatherby had paid considerable attention to the grouping of species within Notholaena, it has not been pos- sible for me to define clearly any segregates or sections, or to place individual species with certainty elsewhere, to the end that the genus would have a better organization and circumscription. The synopsis of the genus presents such groups of species as have certain characters in common, but lacking sufficient understand- ing of the meaning of the characters I have not tried to do more. The American element of Notholaena comprises 59 species, 7 of which are divided into 17 varieties. The United States has 19 species, with 13 in Texas; Mexico has 34 species; Central America 7; the Greater Antilles 8; and South America 20, with 9 species and 4 varieties in Peru. SYSTEMATIC TREATMENT Notholaena R. Br. Prod. Fl. Nov. Holl. 145. 1810. Hook. Sp. Fil. 5: 107-121. 1864; Hook. & Baker, Syn. Fil. 370-374. 1868; Christ, Farnkr. Erde, 150-153. 1897; Diels, in Engl. & Prantl, Nat. Pflanzen- fam. 14; 272-274, 1899; C. Chr. Ind. Fil. 459-464. 1906; C. Chr. in . ? THE AMERICAN SPECIES OF NOTHOLAENA 7 Fern Journ. 36: 7-9. 1946; Weath. “Notholaena” in Brazil, Journ. Arn. Arb. 27: 361-370. 1946. Cincinalis Desv. Berl. Mag 5: 311. 1811, not Gled. 1764. Aleuritopteris Fée, Gen. Fil. 153. 1850-52, in part. Gymnogramma Desy. sensu Mett. Cheil. footnotes 6-13 & text (Abhandl. Senckenb. nat. Ges. 3: 5-7). 1859, in par Cheilanthes Sw. sensu Mett. Cheil. no. | et seq. (Abhandl. Senckenb. nat. Ges. 3: 19 et seq.). 1859, in part; Domin, Bibl. Bot. 20 (Heft 85): 133-136. 1913, in part; Copel. Gen. Fil. 65. 1947, in part. Pellaea Link sensu Prantl, in Engl. Bot. Jahrb. 3: 417. 1882, in art part. Chrysochosma Kiimmerle, Mag. Bot. Lapok. 13: 35. 1914. Type species (chosen by C. Chr. Ind. Fil. xli. 1906): Acrostichum Marantae L. The generic name has been spelled Nothochlaena by various authors. SYNOPSIS OF AMERICAN NOTHOLAENA a. Indument of lamina of scales or hairs, or both, never ceraceous. b. b. Indument of scales, at least in part. 1. N. peruviana, 2. N. are- quipensis, 3. N. squamosa, 4. N. lonchophylla, 5. N. sinuata, a. var. sinuata, 5b. var. integerrima, 5c. var. cochisensis, 6. N. brachypus, 7. N. Hassleri. b. Indument of hairs only. c. c. Lamina deltoid or pentagonal, basal pinnae much the largest, usually strongly inequilateral. 8. N. venusta, 9. N. eriophora, 10. N. goyazensis, 11. N. geraniifolia. c. Lamina ovate to linear, basal pinnae not much enlarged nor strongly inequilateral. 12. N. Pohliana, 13. N. cinnamomea, 14. N. obducta, 15. N. aurea, 16. N. tomentosa, 17. N. Fra- seri, 18. N. Buchtienii, 18a. var. Buchtienii, 18b. var. ven- tanensis, 19. N. Parryi, 20. N. Newberryi, 21. N. mollis. 29b. var. Nealleyji. e. Scales of rhizome not spinescent-ciliate, or if so then the lamina not narrowed at the base and not longer than the stipe. g. 8 ROLLA TRYON g. Basal pinnae not markedly larger than those immediately above, not strongly inequilateral, lamina commonly longer than broad. 30. N. Grayi, 31. N. aliena, 31A. N. Weather- biana, 32. N. Lemmonii, 32a. var. Lemmonii 32b. var. australis, 33. N. Rosei, 34. N. leon g. Basal pinnae much the largest, deltoid and strongly inequi- lateral, lamina often as broad as long, or broader. 35. N. galapagensis, 36. N. candida, 36a. var. candida, 36b. var. Copelandii, 37. N. sul hurea, 38. N. sae 39. N. aurantiaca, 40. N. californica, 41. N. ne d. Scales of rhizome lax, concolorous, not Gite (partially so in no. h. h. Ultimate segments non-articulate, ae and the base of uni- form color, or petiolulate and ark color of the petiolule passing into the base of the Soniene 42. N. Greggii, 43. N. brt ee 44, N. Palmeri, 45. N. pallens, 46. N. pilifera, 47. N. chilensis, 48. N. peninsularis, 49. N. Jonesii, 50. N. Lum- holt 51. N. limitanea, 51a. var. pies 51b. var. mexi- a, 52. N. dealbata, 53. N. Fendler h. Ultimate segments articulate, the dark “ail of the petiolule stopping abruptly in a transverse line. 54. N. delicatula, 55. N. incana, 56. N. nivea, 56a. var. nivea, 56b. var. oblongata, 6c, var. tenera, 56d. var. flava, 57. N. parvifolia, 58. N. formosa. ° KEY TO SPECIES OF AMERICAN NOTHOLAENA 1, Indument of lamina, including upper and lower — but excluding rachis, of scales or of hairs, or both, not ceraceous. 4 Paes on: of scales, at least in part. 3. e 4. Lamina lanceolate or broader, bipinnate, ced ae few; stipe commonly at least half as long as lamina, with one vascular bundle. 5. 5. Scales of rhizome entire or minutely serrulate with very short ascending teeth; scales of upper surface of lamina soon de- ciduous, lance-subulate, flat, or if narrow and subpiliform en 1 mm. or less long, scales of the lower surface erose- serrulate with mostly deltoid teeth. 6. 6. Lamina lanceolate, mostly 12-18 cm. tall; rhizome scales entire or nearly so; scales of lamina dark chestnut brown; dilated soriferous vein-ends projecting into small, hyaline, i. DIAPGITAE JODON oie ons ys nig bern hs sg We peruviana. 6. Lamina deltoid-ovate, acd 2-4.5 cm. tall; rhizome scales minutely serrulate; scales of the lower surface of lamina peor or et chek wn; aie tous vein-ends in Brin un- ns of the margin...... 2. N. arequipensis. ed 5. Reales of atta pret antrorse-serrulate; shin i upper THE AMERICAN SPECIES OF NOTHOLAENA 9 surface of lamina piliform, more or less persistent, more than 1 mm. long, scales of the lower surface ee with often piliform inp 8 angers — 7. Lamina lanceolate, much longer than the east scales of the upper surface of the lamina whitish, those of the lower surface lance- or “inearsubulat, up to 2 mm Weige Pera hy 68 ee N. lonchophylla. 4. Lamina linear with numerous merely lobed hs sometimes en- — pinnae; stipe much shorter than lamina, with two cine bundles. of curses fee ee ae ee 3. Indument of both hairs and scales; fertile margin slightly modified, ora rugose. 8. Stipe very short; rhizome scales siapmne eit or nearly so, not sclerotic; hairs of lamina slender, not moniliform, lanate, on both surfaces; aay and Central America... "6. N. achypus. 8. Stipe nearly as long as lamina; rhizome scales with sharply con- trasting dark videcauie central band and pale margins; hairs of Ree thick, moniliform, on upper surface only; Para- ROU PO OTS OUP PT ae 7. N. Hassleri. 2. siidainahe of hairs only. 9. Lamina deltoid or pentagonal (subpedate), as broad as long or nearly so, usually much shorter than the stipe (sometimes not in no. 9), lowest pinnae much the le eel strongly in- equilateral; Brazil, no. 9 also in Colombia Stipe thinly villous with straight, distin moniliform, often gland-tipped, hairs; lamina with 1-3 wholly free puget mc divided 10. Stipe lanate with long, t Salmon matted, slender, ah Posten hairs, or ae usually none of the primary segments of the lawniria wholly 11. Indument of eth sirtaces of lamina of long, slender, tortuous, tobe articulate hairs; only the basal segments deep- 9. N. eriophora. 11. Indument of lower surface of lamina of fine, short, densely felted tomentum ‘genta by long, straightish, conepionoesly pinngeetoes hairs; median segments, as well as the bas lobed. 12. £2: porn of the upper surface of lamina of = whitish, tortuous, more or less matte airs, m segments simply eaten with entire lobes; e ssng scales bright yg Pe. warawios 10. N. goyazensis. 42. Indument of upper surface of lamina of coarse, Straightish, olden to whitish, not much matted hairs; median segments o bbipinnatifd, their lobes more or less cut; rhizome a 9. Lamina ovate to linear, equalling or exceeding the sti pinnae not greatly enlarged nor strongly inequilatera 13. Indument of simple or pectinately branched hairs, not rane 14. 10 ROLLA TRYON 14. iogament of lower surface of lamina of ne loosely matted in, not concealing the leaf-tissu 15. ‘Hairs of rachis short, spreading or eee unlike those of a; lamina tapering evenly from base to ong-attenuate ONES SRN IA ho Fria ceeere Sie ER oe: 2. N. Pohliana. 15. Hairs of rachis long, like those of lamina; lamina not conspicuously broader at base; Guatemala................ Pek ~ gh esis: 5 eee amen al 13. N. cinnamomea 14. Indument ot "elas surface of lamina dense. concealing the leaf-tissue. 16. Lamina i ae much longer than the on 2 appressed-ascending-pubescent beneath .. 14. obducta. 16. Lamina not much longer than the stipe, or if so, Se ae uni- formly mera ge ifid. 17. 17. Leming gradually reduced at base, usually much longer the stipe, uniformly pinnate-pinnatifid; rhizome at ascending, knotted............... urea. 17. Lamina not conspicuously reduced at base Gometines with a single pair of a reduced pinnae); rhizom i t 18. Stipe and rachis sulcate, glabrous; upper surface of lamina glabrous or glabrate: Chile.. 16. N. tomentosa. 18. Stipe and rachis terete, déciduou sly or persistently —— upper surface of lamina thinly to densely rere - Rhizome scales pale, or with dark center and light margins; lamina pinnate-pinnatifid to quadripinnate, if more than 2-pinnate, then lanate-pubescent above. 20+ 20. iad usually pinnate-pinnatifid, upper surface arachnoid-tomentose. 21. 21. Rhizome with numerous pale scales with con- aacoonely tortuous capi ary tips, giving a tomentose appe rance; no scales on rachis or up- per part of stipe; median pinnae mostly deltoid, veins not visible on the upper — Ecuador . N. Fras i BOuVig? es eee 21. Rhizome usually with few eae abel ci pale, capillary-tipped i not g tomentose; a few deciduous scales among the ac an deltoid- Lanoaclate, e veins usually « subim- pressed and visible on the upper surface; Bolivia and Argentina. .... 18. N. Buchtienii. 20. Lamina usually tri uadripinnate, upper surface lanate; sw. United States and Baja Calif......... Carlen: eticsie FAG reo tits: 10. N. Parryi. 19, ae scales dark, hardly, if at all, margined; lamina _tri-quadripinnate, ultimate segments small, roundish, upper surface “arachnoid pubescent s. Call. & Dae Come 3. . N. Newberryi. THE AMERICAN SPECIES OF NOTHOLAENA ll 13. joan of stellate hairs; lamina tri- -quadripinnate, -~ ulti- ate divisions small, suborbicular: s. Peru & Chile. .21. N. mollis. ke ieee of lamina, including upper and lower surfaces but excluding rachis, lacking, or wholly, or at least in part, ceraceous (in nos. 26-28 the wax often concealed by scales, in no. 31A, by hairs). 2 22. Indument of lamina ceraceous and of scales or hairs (or both in no. 25. Ultimate se ments ‘non-aiticulate, the dark color of = stalk passing into at least the basal part of the segment. 25 25.55 ee and rachis castaneous to brownish; Utah and Ariz. to 49. N. Jonesii. Stipe aad rachis blackish; Sonora......... 50. N. Lumholetzii. 24. Ultimate segments articulate, the dark color of the stalk stop- ping abruptly in a trans e line at its apex. 26 e viscid; lamina 3-4-pinnate, ultimate segments broadly ovate to eibdclieds. the base mostly equilateral; Texas and N. Mex. s to: Tana inessics clone. wee ih cn cha ise ais 57. N. parvifolia. 26. 2 Eps, aided of rachis rounded. 27. of ultimate segments equilateral, may cuneate to vole rhizome scales not v6 cid; lamina 2-3-pinnate; ery. Argenting: a5 ici =33 6c. N. cies var. tenera. 27. one of ultimate segments pao inequilaterally ss rhiz, me scales viscid; lamina 3-4-pinnate, ultimat ee | ment oe to oblong-ovate; Nuevo Leén and Tamau to, Chianas 2 ss ie, Cis 58. N. formosa. pas pa a es oF sient or hairs, or both, in addition to wax. 28 . Indument on lower surface of ‘Hast ceraceous and of large 29. trichomes. 29. Lower surface of the lamina and the rachis with =e brown, stiff trichomes; rhizome scales pire eer rigi strongly pectinate-ciliate; sporangia borne at the ve ein-ends; s. Texas Puen, lamaicn oo. sc ee eeen 29. N. Schaffneri. 29. wer surface of the lamina and the rachis with filiform, weakish, whitish trichomes; rhizome scales thin, bright brown, entire; sporangia borne on the outer % * 14 of the Vemna; Momeiog $3) isc ao. ai iol asc k ee oe 46. AB gid ha 28. Indument on lower surface of lamina — and of scales s oO and sometimes a. tomentose (in no. 31A the tomentum more 30. Scales on lower — of lanting long-ciliate or dissected. 31. 3 1, Lamina linear, its scales with a narro tral portion and long, slender ale or stellate; pinnae entire or very shallowly (rarely deeply) lobed, lateral veins very oblique; Wes Wee 26. N. trichomanoides. tt 31. Lamina broader, the — cut nearly to their rachis, o: lateral veins at a broad an 32. Upper surface of lamina oth stellate trichomes; Texas to s. Ariz. and Mexico............. 27. N. Aschenborniana. 32. Upper surface of lamina with sparse, weak, simple seed Jalisco and San Luis Potosi s. to Oaxaca. .28. N. G Galeo 12 ROLLA TRYON 30. Scales on lower surface of lamina entire or nearly so. 33. Wax orange; scales on lower surface of bette large, broadly ovate; lamina entagonal, the basal pin- nae much the babes and _ strongly ee a Ja- MOR) ee. Cl ela yl Setemang: 39. rantiaca. 33. Wax white to yellow; scales on lower surface “of lamina Oeeenar or narrower; lamina much | r than — e basal pinnae not much the deiegdal nor strongly sepillececk 34, . Upper surface of lamina somewhat ceraceous .30. N. Grayj. 34. Upper surface of lamina with sparse hairs, slightly or not ceraceous, 35. 35. Lower surface of lamina scaly (many scales wel be pili- form but they do not form a tomentum e upper thinly pubescent and sparingly ceraceous; rachis scaly, with the few hairs not forming a tomentu ae ees Seger 31A rN. Weniierbie 29, Indument of ts sara ceraceous. 36. All ultimate segments iy rt for ae, sie breadth; rhizome scales dark-sclerotic, with or without pale margins. = 7. Lamina commonly longer than broad, the lowest pinnae not markedly larger than those immediately eee nor strongly in- equilateral. tipe and rachis sulcate on the upper side. 39. 39. Scales of the rhizome spinescent-ciliate, gid, ie... ... rr ee Ge ee N. Ekmanii. 39. Scales of the rhizome entire. 40. 0. Stipe and rachis blackish or castaneous, glabrate; lamina eae > with a reduced pair of pinnae at base; s. Ariz. and Oey ee ee er ee ey S2.N: Lemmondi. 40. Stipe (except at base) and rachis stramineous, gone puberulent above; ane region, Jalisc Re Os Ta en See ga Ea nica . Rosei. id. 42. Lamina linear to ae the serie deltoid to oblong pinnae with five (or fewer) pairs of lobes. 43. 43. Stipe usually beating sowie base lance-deltoid, long- acuminate scales; rachis a 4 rt, weak, brown, non- glandular trichomes, at least on the upper side; both stipe and rachis blackish; coal and Central America. : N. ee ee N. cubensis. 42. Lamina oblanceolate, the deltoid-lanceolate, acute pinnae with 8-10 pairs of lobes; Nuevo Leén and {Sagan — 41. Rhizome scales _ entire, glandular-margined; Monterrey, THE AMERICAN SPECIES OF NOTHOLAENA 13 ain SORE Shae Sei eee oN: leo ina. 46. Hhsicdcie re bright manta to brownish, they and scales the t the base of the stipe with ve pale maces stipe isecoldens indument of lamina pale yellow to whitish; Galapagos Is e 46. Rhizome scales dark onset to blackish, they and espe- cially those at the of the stipe with broad pale mar- a — blackish to a indument of lamina white; central:Mexico: 03... 665.3. 36. N. candida. 45. Seles of iene and rae Of stipe with erose-denticulate, — glandular margins; lamina broadly pentagonal, pan more than a third as long as the i Mexico eee. Ws ee ag 37. N. sulphurea. 44. Lamina de lowest pinnae usually ae he. lower _ segments usually shortened; indument of lamina com- ellow; sw. United States and Mexico. 38. N. Standley. 36. At ak 6 some of the ultimate segments constricted and yee at the base, not adnate for their full breadth; rhizome scales various but most often thin, bright brown, concolorous. 47. pinnules glandular on upper s shikai their margins sisgacléts and rather slightly revolute; sw. Ariz., s. Calif. and Baja Calif. rn Fn SO) ee Se a sore 40. N. californica. 48. Lamina narrowly deltoid, long-acuminate; stipe and rachis blackish; pinnules glabrate on cia gat surface, their margins entire, strongly revolute; se. Ariz., s. Texas ~ Sprspeog OMICO 6 ee OE EA 41. iain 49. sre and rachis eee pinnae arhianlsite ‘with numerous, approximate, small, cor oat mostly simple pinnules with 0. Rhizome scales subulate, si sclerotic central band and pale, entire margins, those of the stipe lance-ovate; lamina about equalling stipe22.5.. 6 42. N. Greggii. 50. Rhizome scales linear, bright brown and subconcolorous with subsclerotic central band: those of the stipe subulate; lamina longer thas sting: 20. iui ro ks = N. enkiseors 49. Stipe and rachis terete; pinnae acute or obtuse, with m ompound and not crowded pinnules with seat or slightly revolute margins; rhizome scales non-sclerotic (rarely sub- 14 ROLLA TRYON sclerotic), bright brown and concolorous. . Ultimate segments non-articulate, the cmt color of the stalk passing at least — the basal part of the segment, or the segments sessile. 52. Rachis and ae pee! flexuous, the branch- e pinnae an upper part of oh . mina oe pearing dichotomous; s. Wyo. to s. central N. Mex Sa he ba es ns le: Caeieanaiea eee DS; Fendleri. 52. Rachis and pinna-rachises ce or Ai so, the branching obviou usly pinna 53. Most primary pinnules supetilulate or sessile. 54. 54. Stipe and rachis blacki 5 55. Rachis glabrate, xia scales; scales of the stipe ovate, relatively short-acuminate; San Luis Potosi. Vat opereuins ry pa pics ila . N. Palmeri. 55. Rachis ceraceous-glandular, with a few sma all, scales; scales of the stipe lance- or linear-attenuate; s. Baja Calif.......... Hes eo, Vice, aie ae 48. N. peninsularis. 54. Stipe and rachis castaneous to dark castaneous. 56. . Ra ichis ceraceous-glandular, with a few small, red- scales; upper surface of segments ceraceous- g oO 56. Rachis slightly sos mana. mare to glabrous, a scales; upper surface of segments gla- brous; Juan Fernandez Is......... ye N. chilensis. 53. Most primary pinnules a 57. Stipe comparatively stout, it and the wren — brown to blackish; wenaite coriaceous, the v not visible; Utah s. to n. central Mexico _.. ip oe ee gee eee aa ttt cs 51. N. lin vitanea. Ne 51. Ultimate casas articulate, the dark color of the stalk stop- castaneous to dark reddish-br ibe veins nti TE 58. Ultimate . usually ears com oe 6 ails e oe at the base, glabrous on the upper sur- 59. “Stipe blackish; veins usually sporangiferous for their whole length; spores minutely and incons icuously roughened; Mexico and Central America, a niola. siResoi lh AMEN + waas ee ade os 5. N. incana. 59. Stipe light to dark noes: veins usually sporan ngifer- ous only on their outer third or half; — strongly rugose; Colombia Ht pont and se THE AMERICAN SPECIES OF NOTHOLAENA 15 1. Notholaena peruviana Desy. Prod. 220. 1827. Weath. Lilloa 6: 273. 1941. Type: Peru, Dombey P!, photo cul, fragment Bl. Fic. 1. Map 2. Cheilanthes peruviana (Desv.) Moore, Ind. Fil. 250. 1861. Notholaena sinuata var. bipinnata Hook. Sp. Fil. 5: 108. 1864. TyPE: Bafios, Andes of Peru, Brackenridge (Wilkes Exped.), x!, photo cul. Notholaena Brackenridgei Baker, Syn. Fil. 371. 1868. Type: Peru, Brackenridge (evidently the same specimen as above). Rhizome erect, stout; scales linear-ligulate, long-attenuate to a capil- lary tip, tortuous and crisped in drying, with the margins entire or we TToemeeny ee. x “rg “es, LH, Rey me Seats ‘ a ef ~ an) © ooo, (Oe% , Se a Maps 5-7. Map 5, N. sinuata (all in South America is var. sinuata). Map 6, var. oa (north of South America). Map 7, var. integerrima. Map 8, var. co- chisensis. THE AMERICAN SPECIES OF NOTHOLAENA 23 basis of this phase and some later authors misapplied the name N. laevis to it. Texas to Arizona, south to northwestern Argentina; Hispaniola. Some specimens labeled as collected in Chile were evidently not taken within the present boundaries of that country. Looser ° does not report it. Representative specimens: United States. Texas: E. J. Palmer 11235 (BM, GH, MO); Lindheimer 505 (cH, Mo), Exsicc. 1277 (cH, Mo); Wright 814 (BM, GH, MO); Moore & Steyermark 3017 (cu, Mo); Jermy 344 (mo); Mueller 8263 (cu); Waterfall 4571, 4806 (GH, Mo). NEw Mexico: Metcalfe 953 (mo); Maguire et al. 11926 (cH); Wooton 101 (B, GH, MO). ARIzonA: W. W. Brown 19 (cH, Mo); Goodding 2387a (cH, MO); Palmer 274 (mo); M. E. Jones 24681 (cH, MO); Blumer 1527, 2142, 3272 (cH). Mexico. Baya Ca.irornia: Carter et al. 2047 (mo); Shreve 7048 (GH, Mo). Sonora: Hartman 81, 130 (cH); Gentry 1195 (Gu, k, Mo); Shreve 6747 (mo). Cutmuanua: Palmer 82 (cH). Coanumta: Wynd & Mueller 510 (cH, Mo); Johnston 9097 (cu, MO); Palmer 1400 (cu, Mo); Stanford et al. 105, 203 (cH, Mo). Tamau.ipas: Stanford et al. 663 (GH, Mo), 2616 (mo); Palmer 190 (GH, Mo). SmnaLoa: Gentry 5844 (GH, Mo). Duranco: Palmer 462 (s, GH, MO), 883 (B, BM, GH, MO, us). Zacatecas: Lloyd & Kirkwood 23 (mo), 136 (cH). San Luis Porost: Schaffner 947 (cu); Parry & Palmer 995, 996 (cH, Mo), 997 (Mo). QuERéTARO: Arséne 10646 (GH, Mo). Hmatco: Pringle 11269 (8, GH, MO, P, us). Distrito FEDE- RAL: Lyonnet 94 (Gu, Mo), 1582 (GH). Pursta: Rosenstock Fil. Mex. Exsicc. 35 (Mo). VeRAcruz: Purpus 9043 (cH, Mo, us); Bourgeau 2770 (GH). Gurerrero: Hinton 9724 (Gu, Mo). Oaxaca: C. L. Smith 2027a (mo); Conzatti 837 (cu, Pp). Cutapas: Ghiesbreght 212 (3M, GH, ¥); Schumann 1880 (P). Guatemala: Skutch 1149, 1964 (cH); Seler 3039 (B). Costa Rica: Bernoulli & Cario 239 (B, Pp); Warszewicz 48 (B). Haiti: Leonard 7831 (cH, us). Dominican Republic: H. von Schrenk 46 (Mo). Venezuela: Tamayo 263 (us); Engel 139 (B); Mocquerys 1115 (Pp); Linden 511 (8M, Pp). Colombia: Stiibel 222 (B); Mutis 2779 (us). Ecuador: Mille 269 (Pp); ee 728 (B); Jameson 6 (Bm); Rose & Rose 23968 (cu, Us). Macbride 3190 (¥, us); Macbride ¢ Featherstone oD (F, GH, US). Junin: Killip & Smith 21808 (us). Lima: Goodspeed 11307 (cu, MO); Macbride ¢& Featherstone 282 (¥F, us). Cuzco: Stork, Horton & Vargas 10449 (mo). Arequipa: Pennell 13071 (r, cu). Bolivia: Buchtien 3114, 3115 (vs); Steinbach 5045 (cH); Bang 786 (B, cH, MO, us); Julio 97 (cu, us); Herzog 1223 (B). Argentina. Juyuy Burkart & Troncoso 11461 (mo). Sata: Hieronymus & Lorentz 1 95 (B). TucumAn: Lorentz 217 (), 218 (s, us); Venturi 838 (cH, LIL, us). Caramarca: Castillén 11622 (cu). * Looser, G., “El ‘Notholaena’ en Chile,” Darwiniana 7: 62. 1945. 94 ROLLA TRYON 5b. Notholaena sinuata var. integerrima Hook. Sp. Fil. 5: 108. 1864. Weath. a Arn. Arb, 24: 313, 1943. Type: Mexico, Liebmann x}, photo cH Fic. 5b. Map C. A. Weatherby (l.c.) has discussed the general misapplica- tion of Hooker’s name to what is now known as var. cochisensis, the proper choice of type and his redefinition of the variety. Southern Oklahoma to Arizona south to Veracruz. Representative specimens: United States. OxLtanoma: 1925, F. C. Greene (cH); April 20, 1928, Featherly (mo). Texas: Lindheimer 414, Exsicc. 1278 (cH, Mo); Cory 869, 29692, 43284 (cH); E. J. Palmer 11791 (cu, Mo); Wright 815 (cH, Mo, us); Waterfall 4838 (cH, MO); Reverchon 1184 (mo); Tracy 8051, 8321 (mo); Jermy 674 (mo). New Mexico: Earle & Earle 564 (mo); Eastwood 8268 (cH); Goodman 2329 (cH, Mo). Arizona: Palmer 275 (Mo); Blumer 1530 (cH, MO); Goodding 373 (cH); Eggleston 10972, 10982 (cH); W. W. Brown 18 (GH, MO). Mexico. Sonora: S. S. White 4585 (GH). Cui- HuAHUA: Pringle 464 (GH, Mo). CoanurLa: Gregg 297 (mo), 365 (cH, MO); file’ 1406 (cH), ‘1407, 1409 (cH, Mo); Wynd & Mueller 321 ae ; Stanford et al. 429a (GH, Mo). Nurvo LEON: Mueller & Mueller 1071 (cH); Palmer 1404 (cH, Mo). Duranco: Gentry 8323 cH). Zacatecas: Lloyd & Kirkwood "24 (mo), 137 (cH). San Luts Potosi: Johnston 7573 (cH); Palmer 447 (BM, GH, MO). H1pa.co: M. T. Edwards 712 (Mo). Veracruz: Aug. 9, 1924, Fisher (Mo). 5c. Notholaena sinuata var. cochisensis — Weath. Journ. Arn. Arb. 24: 314. 1943. Fic. 5c. Map 8. Notholaena a var. crenata Lemmon, Ferns Pacific Slope 7. 1882, nomen nu Notholaena bbohdecnals Goodd. Muhlenbergia 8: 93. 1912. TyPE: Montezuma canyon, Huachuca Mts., Arizona, Aug. 1910, Goodding Notholaena sinuata f£. crenata (Lemmon) Clute, Fern Bull. 15: 46. 1915. This variety was long called var. integerrima; C. A. Weatherby (l.c.) has discussed the reapplication of the name. This has been reported as poisonous to stock and this property might reinforce its morphological characters sufficiently for specific status were it not for its intergradation with var. integerrima. Texas to southern California, south to Aguascalientes. Representative specimens: United States. Texas: Waterfall 3909 (cH), 4447 (cH, Mo), 4543, 4741 (GH), 4794 (cH, Mo), 4969 (cH), 5043 (GH, MO); Reverchon 1184 (cu); E. J. Palmer 34167 (cu, Mo); Moore & Steyermark 3240 (cu, Mo); M. E. Jones 5133 (cu); Cory as SarA Hib JH Ne 1 ‘Ny a A \ }} \ , . Fic. 5-7. Fic. 5. N. sinvaTa: 5a. var. stnuata: A, frond, X 0.5; B, segment, X 4.5; \, sorus, enlarged; D, scale from upper surface of pinna, enlarged; all fro in 1895 (mo). 5b. var. INTEGERRIMA: scale from underside of pinna, X 9, from Cory (Mo). 5c. var. cocHIsENsis: A, frond, 5; B, scale from underside of Pinna, X 18; C, scale from upper surface of pinna, enlarged; all from Stanford et al. 355 (mo). Fic. 6. N. BRACHYPUS: A, frond, X 0.5, from Pringle 1787 (Mo); B, : t, 5, from Pringle 1787 (mo). Fic. 7. N. Hasstert: A, frond, X 0.5; B, upper surface of Segment, X 4.5; C, segment, X 4.5; all from Hassler 10996 (mo). 26 ROLLA TRYON 871, 873, 874 (cH). New Mexico: Metcalfe 954 (mo); eee 3733 (cH, Mo). Arizona: Eastwood 5971 (cH); Blumer 1522 (B, G Mo, us); Goodding 373 (cH); Thornber 2051 (cu); Phillips 3613 (cu); CB incr 276 (mo). Nevapa: April, 1924, Hewett (us). Catt- FORNIA: Munz et al. 3550, 4224 (us). Mexico. Sonora: Hartman 25 (cH). Camauanua: Johnston 7853, 7880a (GH); Palmer 357 (GH, MO); Harvey 1358 (Gu, Mo). Coanurta: Wynd & Mueller 659 (cH, Mo); Gregg 297 (cH); Stanford et al. 42, 194, 355 (cH, Mo); Wynd 745 (cH, Mo); Johnston 9096 (cH, Mo); Palmer 359 (GH, MO), 1401 (cH), 1405 (cu, Pp, us), 1408, 1410 (GH). Duranco: Johnston 7792 (cH). ZacaTEcas: Lloyd & Kirkwood 138 (cH, Mo); Stanford et al. 548 (GH, Mo). AGUASCALIENTES: Rose & Painter 7720 (cu, us). 6. Notholaena brachypus (Kze.) J. Sm. Ferns Brit. & For. 172. Fic. 6. Map 9. Cheilanthes squamosa var. brachypus Kze. Linnaea 18: 340. 1844. TYPE: Mexico, Leibold 52, s!, photo of specimen at k, Gul. Cheilanthes brachypus Kze. Linnaea 18: 341. 1844, nomen pro- visorium; Linnaea 23: 307. 1850. Rhizome short, thick, erect; scales linear-attenuate, brown or castaneous, capillary-tipped, entire or somewhat pectinate-serrulate. Fronds 4-28 cm. tall; stipe usually very short or obsolete, with one vascular bundle. Lamina narrowly elliptic or lance-elliptic, gradually narrowed below, pinnate-pinnatifid with 10-21 pairs of pinnae, under surface with dense, rusty, lanate tomentum beneath the imbricate, ovate-lanceolate, long-attenuate, entire or somewhat serrulate scales, upper surface with thin tomentum of similar hairs, rachis brown, covered with scales similar to those of the lamina. Pinnae lanceolate, obtuse or acutish, the median deeply cut into 5-10 pairs of oblong, obtuse lobes. Veins simple or once-forked, sporangia (with 32 spores) borne on their dilated ends, margin modified into a hyaline band some- what incurved over the sporangia. The short stipe, dense tomentum beneath the scales on the under | tie and the modified margin are characteristic of this specie Diep shaded banks, rocky places and on rocks, 200-1600 m., central Mexico, Sinaloa to Veracruz; Guatemala to Costa Rica. Representative specimens: Mexico. Leibold 52 (B, Kk). SINALOA: Oct. 31 obi sb wei ae (us). San Luts eee Parry & Palmer 998 (GH, K, US, y). c (Nayarit): Mexia 511 (BM, us), 769 (c); Pennell 19940 fen. as a Pringle 1787 a BM, F, G, GH, K, MO, Pp, us, y), 11784 (8, F, Gu, y : feed Palmer, all in 1886, 57 (cH, US, Y), BM, GH, K, P, Us, Y), 733 (BM, K, P, US, x); R. S. Ferris 5814 (cu, us); Rose & Painter ay (us). Cotrma: Palmer 1230 (cu, xk, THE AMERICAN SPECIES OF NOTHOLAENA 27 us, ¥). MicHoacan: Arséne 42 (B, BM, P). More tos: Pringle 11785 (GH, P, us); Deam 31 (cH, us); Rose, Painter & Rose 6875 (cH). VERACRUZ: Purpus 5693, 7870 (us). Guatemala: Muenscher 12120 (us); Standley 58224 (¥); Lehmann 1691 (3, BM, K, us); Salvin ¢ Godman 1862 (x); Heyde & Lux 3200 (n, cu, k, us), 4091 (cH, Mo, Pp, us); Tuerckheim 8825 (p). Salvador: Padilla 168 (us); Standley 20948, 21328 (cH, us), 22536 (us); Calderén 1785 (cH, us). Nica- ragua: Lévy 1131 (B, F, Pp); Jimenez 349 (us); Chaves 287 (us); Maxon 7656 (cu, us), 7676 (us); Maxon, Harvey & Valentine 7622 (us). Costa Rica: C. Hoffmann 666 (B); Brenes 15699 (F); Brade 359 (Ny). 7. Notholaena Hassleri Weath. Lilloa 6: 274, pl. iv. 1941. TyPE: Paraguay, Hassler 10996 x!, photo cu!. Fic. 7. Map 10. Rhizome horizontal or ascending, short; scales linear-subulate, ca. 2.5 mm. long, at least the older with shining dark brown or blackish sclerotic central band and well-developed, pale brown, minutely and closely serrulate hyaline margin. Fronds densely and _ irregularly clustered, 8-20 cm. tall; stipes stout, about as long as or somewhat shorter than the lamina, terete, blackish-castaneous, with one vascular bundle, densely beset with lance-ovate, acuminate, cordate-based, pectinate-serrulate, thin, brownish to whitish scales with large trans- lucent thin-walled oblong cells. Lamina lanceolate or oblong-lance- olate, not narrowed at base, above rather abruptly contracted to an acute pinnatifid apex, bipinnate-pinnatifid or subtripinnate at base; rachis on both surfaces and costae and costulae beneath densely beset with scales like those of the stipe, leaf-tissue otherwise glabrous on the lower surface, on the upper rather sparsely beset with short, thick, whitish, moniliform trichomes with very short cells. Pinnae 8-10 pairs, the lower rather distant, oblong, or linear-oblong (or the lowest narrowly subsessile) , obtuse to acutish, pinnules deltoid-ovate, obtuse. Veins simple, sporangia borne on the outer third of the slightly or not at all thickened veins which stop just short of the somewhat modi- fied (with short, rounded, hyaline lobes) margin. Specimens examined: Paraguay: Hassler 10996 (8, BM, G, K, MO), 12980 (Pp). 8. Notholaena venusta Brade, Anais Prim. Reun. Sul-Amer. Bot. 2: 7, t. 4, figs. 1, 2. 1940. Weath. Journ. Arn. Arb. 27: 363. 1946. TYPE: Diamantina, Minas Geraes, Brazil, June, 1934, Brade 13494 np, not seen. Fic. 8. Map II. Notholaena capillus St. Hilaire in herb.; ex Christ, Bull. Herb. Boiss. II. 2: 381. 1902, pro syn. based on St. Hilaire B’ 1220. _ Rhizome about 2 mm. in diameter; scales about 2 mm. long, some- times serrulate toward the apex with narrow, ascending teeth. Fronds 3-20 cm. tall; stipe terete, 0.4-0.5 mm. in diameter, 6.5-9 cm. 28 ROLLA TRYON -— ~ i) ~ Maps 9-11. Map 9, N. brachypus. Map 10, N. Hassleri: Map 11, N. venusta. noe Shee oe shining, with one vascular bundle. aoe pentag- onal t what elongate-deltoid, 3-4 cm. long, cm. wide, rote “fully pinnate to about the third pair of is from base; tomentum of the upper surface grayish, of slender, tortuous and matted long hairs, that of the lower surface rufous, of similar hairs underlaid by a dense felt of shorter and finer hairs, rachis castaneous. Basal pair of pinnae deeply pinnatifid and more or less inequilateral by the elongation of the basal segment on the lower side, their divi- sions oblong, obtuse, entire or the developed basal segment shallowly undulate-lobed; median pinnae oblong or linear-oblong, entire or shal- lowly lobed, rather distant, decreasing rather gradually to a somewhat prolonged, obtuse, narrow, pinnatifid lamina-apex. Veins immersed, ultimate veinlets 1-2-forked, sporangia (with 64 spores) borne on the rather abruptly dilated, somewhat flabellate tips at or very near the unmodified margin. Eastern Brazil. Specimens seen: Brazil. Minas Gerags: St. Hilaire B’ 1220 (Pp, photo cH); Magalhdes Gomes 1099 (Pr); Schwacke 14250 (cu, P). Piauny: 1836, Gardner 2392, in part (P). . Notholaena eriophora Fée, Gen. Fil. 159, t. 13, fig. 3. 1850-52. Weath. Journ. Arn. Arb. 27: 365. 1946. Type: Shady cliffs on the hills near Oeiras, Piauhy, Brazil, March, 1839, Gardner 2390, at rB?, not seen, isotypes! (see below). Fic. 9. Map i. Notholaena palmatifida Kze. Farnkr. 1: 148. 1844, nomen nudum. Based on Gardner 2390. Polypodium eriophorum (Fée) Hook. Ic. Pl. 10: t. 991 (Cent. Ferns t. 91). Cheilanthes eriophora (Fée) Mett. Cheil. no. 14. 1859. upper surfac e of pinna, X 2.5; all tg Clean 14408 (cH). 30 ROLLA TRYON Rhizome 2-3 mm. in diameter. Fronds ca. 5-12 cm. tall; stipe terete, 4-11 cm. long, 0.4-0.5 mm. in diameter, with one vascular bundle. Lamina pedate-pinnatifid, 2-3.5 cm. long and as broad or somewhat broader, rather thin, indument of both surfaces of whitish to rufescent hairs. Basal pair of segments usually much produced on the lower side and strongly inequilateral, their basal divisions on the lower side themselves lobed and somewhat inequilateral, all divisions broadly obtuse; upper part of lamina cut to near rachis into 3-4 oblong, obtuse entire segments, tapering evenly into a broad, short, 3-4-lobed obtuse apex. Veins 1-2-forked, sporangia (with 64 spores) borne on their apices a little back from the unmodified margin. Northeastern Brazil and Colombia (Macarena Mts.), some- times on sandstone rocks and perhaps confined to them. The report from Colombia is taken from Alston in Mutisia 7: 8. 1952; it is based on Philipson 2293. Specimens seen: Brazil. Glaziou 14409 (B, xk, Pp, us); Herb. Kew 1037 (¥). Prauny: Ule 46, 7423 (Bp); Gardner 2390 (BM, G, GH, K, Ny, P, us). CrarA: Schwacke 2545 (B). 10. Notholaena goyazensis Taubert, in Engl. Bot. Jahrb. 21: 421. 1896. Weath. Journ. Arn. Arb. 27: 366. 1946. Type: Serra Dourada, Goyaz, Brazil, Jan. 1893, Ule 3222, whereabouts unknown. ic. 10. Map 13. Cheilanthes goyazensis (Taubert) Domin, Bibl. Bot. 20 (Heft 85): 133. 1913. Rhizome about 4 mm. in diameter; scales 3-4 mm. long, 0.1-0.2 mm. wide at base, remotely serrulate toward apex, those of the young growth bright brown and concolorous, the older with castane- ous, sclerotic central band. Fronds ca. 8-15 cm. tall; stipe 3-10 cm. long, with one vascular bundle, with the tomentum about 1 mm. in diameter, in age glabrate and blackish castaneous. Lamina 4.5-6 cm. long, usually about as wide, pinnate-pinnatifid or, in the basal pinnae only, sub-bipinnatifid; tomentum of upper surface whitish or gray- ish, that of the lower surface at first whitish, in age dull brown or pale ferrugineous. Basal pinnae usually connected with those above by a narrow wing along the rachis, inequilaterally elongate-deltoid, the basal segments on the lower side much produced and pinnatifid with oblong, obtuse lobes, median segments of lamina _linear-oblong, equilateral, pinnatifid with oblong, obtuse entire lobes, the basal obes adnate to the rachis and forming a broad, basally narrowed wing along it, the 2-4 upper segments rather abruptly contracted into an obtusish, short or sometimes produced, pinnatifid lamina-apex. Veins at an acute angle to the costule, simple or 1-forked, the sporangia (with 64 spores) borne on their subflabellate tips at the margin which is bordered by a very narrow hyaline band. THE AMERICAN SPECIES OF NOTHOLAENA 31 Eastern Brazil, on sandstone and perhaps other rocks. pecimens seen: Brazil. Minas Gerars: Glaziou 16643 (B, GH, K, p); April, 1905, Silveira (e); L. O. Williams 6871 (cu, us). Goyaz: Glaziou 22625 (B, G, p); Damazio 1859 (B, Ny, P, US). 11. Notholaena ee’ St. Hil. ex Weath. Journ. Arn. Arb. 27: 367. 1946. TyPE: prope S. _— da Ti[Je]quitinhonha, Minas Geraes, Brazil, St. Hilaire B’ 1489 p!, photo cu!. Fic. 11. Map 14. Mars 12-17. Map 12, eriophora. Map 13, N. goyazensis. Map 14, N. geraniifolia. Map 15, N. Pohliana. Map 16, N. cinnamomea. Map 17, N. obducta. i ROLLA TRYON Rhizome horizontal, ca. 2 mm. in diameter; scales sub-appressed, narrowly linear with dark brown subsclerotic central band and nar- row paler hyaline, remotely serrulate margins, and capillary tip (ca. 3-4 mm. long, 0.2 mm. wide). Fronds subapproximate ca. 10-22 cm. tall; stipe slender, somewhat angled and subterete, deep castaneous, shining, with an inconspicuous, sparse, thin, pale, scarcely matted tomentum or nearly glabrous, much longer (3-5 times) than the lamina, with one vascular bundle. Lamina pentagonal, almost as broad as long 5 cm. or less each way, deeply bipinnatifid (the lower pinnae) or at base tripinnatifid, upper surface densely subappressed hirsutulous with comparatively coarse golden-brown shining, almost straight and little entangled hairs, the lower with dense, close, lanate tomentum of very fine, tortuous and closely matted, strongly fer- ruginous hairs. Lower pinnae enlarged, inequilaterally deltoid, the basal pinnules on the lower side much elongated and deeply pinnatifid, upper pinnae (about 4 pairs) pinnatifid, abruptly reduced to a short, pinnatifid acutish lamina-apex, ultimate lobes linear-oblong. Veins once- or twice-forked, the few sporangia (with 64 spores) on a some- what elevated punctiform receptacle at their tips, margins with a small hyaline lobe immediately outside of the vein-end. Eastern Brazil, on rocks. Specimens seen: Brazil. Minas Gerass: St. Hilaire B’ 1489 (P); A. Maublanc 584 (pr); Glaziou 14408 (8, G, GH, K, P). Banta (?): bei Calderao, Ule 7239 (s) 12. Notholaena Pohliana Kze. Farnkr. 1: 45. 1840. Weath. Journ. Arn. Arb, 27: 369. 1946. Type: Goyaz, Brazil, Pohl, at w?, not seen. A specimen of Gardner 3554 at c! det. by Kze. may be taken as authentic. Fic. 12. Map 15. Cheilanthes Pohliana (Kze.) Mett. Cheil. no. 13. 1859. Fronds ca. 30 em. tall; stipe slender, terete, blackish, dull, glabrous or beset with short, pale, retrorse, sometimes branched trichomes. Lamina tapering regularly from base to long-attenuate apex, 8-17 cm. long, cm. wide, w up to 18 pairs of pinae, texture herbaceous, both surfaces loosely beset with long, simple, distinctly articulate, pale brownish hairs; rachis like stipe. Lower pinnae with 3-5 pairs of free, oblong or deltoid-ovate, obtuse, entire or sometimes lobed pinnules 6 mm. or less long, and 3-5-lobed deltoid, obtuse terminal segments. Veins 3-4-forked, sporangia borne on the only slightly clavate vein-ends, margin unmodified Glaziou 15735 from “Environs de Rio de Janeiro et dOuro Preto” was probably not actually collected at both localities; I am considering Ouro Preto as the correct one. Eastern Brazil, presumably on rocks. Abs WARS raat VO oh ON ads NY Fe a MONS IUR: ai UD YINY ua ( 2 TRS Ss YAS h Aes NA) WARTS nA MTA NS VA typ pag A DS * eNO \ i if}, ii} Fic. 12-15. Fic : lee tee eS om Gard . Pontiana: A, frond, ‘ ok tachia writers (us). Fic. 13. N. pe er ters hes se oes X 4.5; both A, frond, X 05: ; C, segment, X 4.5; all from Sal eee > 5; B, portion 1 5686 Gao). me a dee of ray Day ge og eye Leggs a . . . AUREA: t ie > Fs ojas segment, x 9; all from E. z, P Moe PE bor underside of pinna, bs 45; 34 ROLLA TRYON Specimens seen: Brazil. Minas Gerars; Glaziou 15735 (B, k). Goyaz: Gardner 3554 (B, BM, F as 3551, G, K, P, US). 13. Notholaena cinnamomea Baker, Syn. Fil. ed. 2, 515. 1874. TYPE: Motagua (“Montagua”), Guatemala, 1862, Salvin & Godman K!, photo cul. Fic. 13. Map 16. Cheilanthes cinnamomea (Baker) Domin, Bibl. Bot. 20 (Heft 85): 133. 1913. remotely toothed scales and hairs like those of lamina. Lamina herbaceous, ovate or lanceolate 6-12 cm. long, 2.5-5 cm. wide, bipinnate-pinnatifid, nearly glabrous on the upper surface, beneath loosely beset with long weak pluricellular (but not obviously articulate) brownish hairs pinnately branched at base. Pinnae 8-9 pairs, lance-oblong, obtuse 0.8-1.3 cm. wide, the lowest not or but slightly smaller, pinnules, 5 to 7 pairs, deltoid-ovate or oblong, very obtuse, inequilateral at base, distal side much wider. Veins 1-2- forked, sporangia (with 64 spores) borne on their terminal portions which are slightly or not at all thickened, margin unmodified. Guatemala, moist shaded bank, 450 m. Specimens seen: Guatemala: 1862, Salvin & Godman (BM, K, US); Volcan de Fuego, Salvin (cx). 14. Notholaena obducta (Kuhn) Baker, Syn. Fil, ed. 2, 515. 1874. Weath. Lilloa 6: 266. 1941. Fic. 14. Map 17. Cheilanthes obducta Mett. ex Kuhn, Linnaea 36: 83. 1869. TYPE: Bolivia, d’Orbigny 386 s!, photo cul. Notholaena Balansae Baker, Journ. Bot. 16: 301. 1878. TyPE: Para- guay, Balansa 330 x!, photo cul. Cheilanthes Balansae (Baker) Domin, Bibl. Bot. 20 (Heft 85): 133. 1913 Notholaena Herzogii Rosenst. in Fedde, Rep. Sp. Nov. 6: 175. 1905. TYPE: Bolivia, Th. Herzog 117, probably at s-pa, not seen, isotype, Bl, photo cul. ? single, vascular bundle, stoutish, castaneous, covered with subap- pressed deciduous trichomes similar to those of lamina. Lamina THE AMERICAN SPECIES OF NOTHOLAENA 35 lanceolate, bipinnate, with numerous (15-25) pairs of oblong-lanceo- late pinnae, upper surface glabrous or nearly so, lower surface densely clothed with long, straightish, subappressed, relatively thick, brownish, pluricellular, compound trichomes, with 3-6 elongate lower cells attached at their bases only, the remainder of each cell free, more or less reflexed and imbricate; rachis similar to stipe. Pinnae simply pinnate, with 8-10 pairs of oblong, broadly obtuse, entire pinnules, which are sessile or nearly so, subcordate and nearly equilateral at the broad base. Veins 1-, or the basal, 2-forked, sporangia (with 32 spores) borne on the abruptly dilated vein-ends a little within the unmodified margin which is narrowly revolute over them. The peculiar trichomes make this an exceptionally well-marked species Colombia (upper Magdalena valley); Bolivia, northern Argen- tina and Paraguay, dry or shaded rocky places or in sandy soil, mostly at 1000 m. or less. Representative specimens: Colombia: Lehmann 6056 (8, GH, P, US). Bolivia: R. S. Williams 1374 (us); dOrbigny 386 (B, Pp). Paraguay: Balansa 330 (xk, Pp); Rojas 3347 (p); Anisits 2147 (Ny, us). Argen- tina. Juyuy: Schreiter 5129 (cu, tL); Venturi 5111 (cu, us). SAvTa: Hieronymus & Lorentz 269 (B); Eyerdam A ee oe ie TucuMAN: Venturi 846, 1871 (GH, LIL, Us). 1910, Castillon (pe), 714 (GH, LiL). ENTRE fee "Shuckert 6582 Khas . Notholaena aurea (Poir.) Desv. Mém. Soc. Linn. Paris 6: 219. 1897, Weath. Lilloa 6: 260. 1941; Journ. Arn. Arb. 24: 312. 1943. Fic. 15. Mar 18. Pteris aurea Poir. Encycl. Meth. 5: 710. 1804. type: Peru, Joseph de Jussieu (sheet 1333 in hb. Jussieu) P!, photo cul. Acrostichum eps Willd. Sp. Pl. 5: 114. 1810. type: “Bonaria,” B!, photo cu!. Cheilanthes fetes Willd. ex Link, Enum. Pl. Berol. 2: 463. 182 “America meridionali,” B!, ae cH!. Not Cincinalis Pica ‘Dev. which is N. trichomanoi Notholaena rufa and vars. minor & major ‘Presi, Rel. Haenk. 1: 19. 1825, the species-name superfluous. TyPE: Mexico, Haenke, pr! (sheets 78539 and 78540), photo cu!. Pellaea ferruginea (Link) Nees, Linnaea 19: 684. 1847. Notholaena ferruginea (Link) Hook., Second Century Ferns, sub t. 52. 1861; not Desv., 1813, which is N. trichomanoides. Notholaena ferruginea var. canescens Kze. ex Fourn. Mex. Pl. 1: 120. 1872, nomen nudum (Schaffner 47 P!, is so labeled by Fourn.). ronetholaena bonariensis (Willd.) C. Chr., Ind. Fil. 6. 1905; 459. 36 ROLLA TRYON Notholaena chiapensis Rovirosa, Pteridographia Mex. 229, t. 48, fig. 1-6. 1909. TYPE: Chiapas, Mexico, Rovirosa 1077, location un- known; isotype, GH!, pH—photo mol, vs!. Rhizome short-repent; scales lance-linear, with shining, castaneous, sclerotic central band and narrow or relatively broad, pale brown, entire, hyaline margin. Fronds ca. 20-60 cm. tall; stipe % as long as blade or less, terete, castaneous or blackish, with a single vascular bundle, clothed, as is the rachis, with coarse, straightish, subap- pressed, whitish, septate hairs. Lamina linear-elliptic, long-attenuate at base, pinnate-pinnatifid almost to the gradually or abruptly nar- rowed obtusish apex, upper surface rather sparsely clothed with straightish, rather coarse, whitish or golden tinged pluricellular hairs, the lower covered with dense tawny tomentum (white when young) of fine matted hairs. Pinnae numerous (up to 40 pairs), oblong or deltoid-oblong, obtuse or acutish, cut %-% to the costa into oblong or linear-oblong, entire obtuse lobes. Veins oblique, 1-2-forked, sporangia (with 32 spores) borne on their clavate to subpunctiform ends, margin modified with a very narrow subhyaline band. Notholaena aurea is a distinctive, and, considering its broad distribution, a remarkably uniform species. C. A. Weatherby ° has discussed the proper application of Poiret’s long-overlooked Pteris aurea. The literature records mapped are from Morton in Kearney & Peebles, Arizona Flora, 1951, Correll in Lundell, Flora of Texas 11, 1955 and Hicken, Cat. Polipod. Argent. in Rev. Mus. La Plata 15. Southwestern United States, Mexico, Central America, West Indies and through the Andean region to northern Chile and Argentina; growing in a variety of moist or dry, usually rocky, habitats, ca. 700-4000 m. Representative specimens: United States. Texas: C. H. Mueller 8261 (cu); Havard 1219 (P); Moore & Steyermark 3056 (cu, vs), 3238 (cu, us); E. J. Palmer 30608 (cu, p, us), 31975 (us), 34101 (cH). New Mexico: Wooton 102 (cu, Pp, us), in 1902 (us); June 9, 1906, Standley (us). Arizona: Goodding 2574 (cu); M. E. Jones 4329 (Gu, p, us); Rothrock 612 (cu, x), 613 (GH, us, ¥); Blumer 2122 (cu), 2141 (us); Parish 267 (us); Shreve 5066 (us); Peebles et al. 1338, 1345 (us); Eggleston 14224 (us). Mexico. Baja CALiroRNIA: Brandegee 658 (cu, us); 1930, M. E. Jones (us). Sonora: Hartman 171 (cH), 238 (cH, us), 253 (GH), 328 (cH, us). Cumauanua: Gold- man 152 (cu, us); Pringle 462 (¥, GH, MO, P, y), 899 (GH, P, US). * Weatherby, C. A. Contrib. Gray Herb. 124: 21. 1939. THE AMERICAN SPECIES OF NOTHOLAENA 87 CoanuiLa: Gregg 219 (B, cH, Mo); Wynd & onbtiot 511 (cu, - Palmer 369 (cH, us), 1399 (cH, P, Us, ¥). Nuevo LEOn: C. H. M. T. Mueller 332 (cu), 955 (F, cH). Pantha: Bartlett Pod (us). Duranco: Palmer 240 (¥F, Gu, us), 354 (Gu, us), 463 (GH, Us), 464 (Bb, F, GH, Us), 508 (Gu, us), 890 (F, GH, us). ZacaTEcas: Rose 2747 (Gu, us), M. E. Jones 541 (us). SAN Luis Potosi: Schaffner 35 (GH, us), 946 (Gu, k, us, y); Parry & Palmer 993, 994 (¥F, GH, P, US, Y). Jauisco: Palmer 551 (Pp, us, y), 632 (Gu, P, US). Guanajuato: Dugeés 5 (GH). QueRETARO: Arséne 10648 (us). Hipaxco: Rose & Painter 6681 (us). MicuHoacan: Seler 1243 (B, GH, us); Sint 3382 (F, GH, MO). Mexico: stein bint 34 (B, BM, P, US), 35 (B, F, P, US); Bourgeau 257 (B, Ny, P, us); Rose & Painter 6474, 6613, 6800, 6990, 7126, 7858 (us). Distrito Feperat: Pringle 11268 (B, F, GH, Mo, P, US), 11788 (3, cu, P, us); Hinton 3309 (cu, K); Seler 3525 (B, GH, Us). More os: Rose dr Rose 11059, 11097 (us). Pursta: Pittier 445 (us); J. G. Smith 56 (us). Veracruz: Seaton 121 (cx, us); Bourgeau 3153 B, GH, K, NY, P, y). GuERRERO: Hinton 9486 (F). Oaxaca: Purpus 1134] (us). Cutapas: Ghiesbreght 275 (cH, x). Guatemala: Ber- noulli & Cario 243 (B, k, Pp); Skutch 799, 807 (cH); Heyde & Lux 6284 (B, GH, US); Lehmann 1527 (B, BM, K, us); Standley 58588, 65566, 65765, ‘66479, 82388 (F); Tonduz 756 (F, K, us). Honduras: T. J. Dyer A280 (us). Costa Rica: Brade 136 (8, P). Jamaica: Watt H11944 (n, F, k, us), 13875 (B, us); Fuertes 1913 (cu, P, vs). Marion Moritz 250 (x), 350 (B, BM, K, P); Linden 514 (8, BM, K, P); Gehriger 239 (us). Colombia: Mutis 3088 (us); Killip & Smith 16386, 16486, 16857, (cu, us); Hartweg 1514 (B, BM, us); Lindig 130 (BM, x, p); Lehmann 4446 (B, K, P, US), 7588 (B, GH, K, US). Ecuador: Jameson 31, 32 (B), 46 (x), 218 (BM, P); Lehmann 631 (B); André 564 (k, Ny); Spruce 5325 (BM, GH, NY, P, ¥); Mexia 7429 (us). Peru. AMAzonas: Matthews 205 (BM, K), 3289 (B, BM, Us). HuANuco: Macbride & Featherstone 1921, 2050 (us). JuNin: Safford 991 (cH, P, us); Killip & Smith 21809 (us). Ayacucno: Killip & Smith 22256 (us). Cuzco: Cook & Gilbert 361, 695 (us); Hitchcock 22548 (us). Bolivia: D’Orbigny 166, 372, 404 (Pp); Buchtien 261 (x, GH, P, US), 471 (vp, us), 3113 (us), 5007 (us); Bang 205 (x, Pp, us), 752 (GH, P, us); Mandon 1572 (BM, GH, K, NY, P); Fiebrig 2860 (B, GH, P, PR, US); Rusby 332 (cu, p, us), 336 (n, cu, P, us); Herzog 1717, 2496 (B, us). Chile: Johnston 545 (cH, us). Argentina. Juyuy: Parodi 9794 (cH); Venturi 7397, 9006 (us). Satta: Venturi 6838 (us); Lorentz & Hieronymus 158 (x, Pp, us), 160 (B, Pp). TucuMANn: Rodriguez 547 (GH, LIL); Venturi 1719 (LIL, Us), 4190 (cu, tit, us); Lillo 1043 (cu, ut), 5061 (cH, LIL). 88 ROLLA TRYON | aes. a pase Ae Ties a com irae a ane eo | Maps 18-22. Map 18, N. aurea. Map 19, N. tomentosa. Map 20, N. Fraseri. Map 21, N. Bichuend, large dots var. Buchtienii, small dot var. ventanensis. Map 22, N. Parryi. THE AMERICAN SPECIES OF NOTHOLAENA 39 16. Notholaena tomentosa Desv. Journ. Bot. Appl. 1: 92. 1813; not (Link) Keys. 1873, which is Cheilanthes tomentosa. Weath. Lilloa 6: 274. 1941 Fic. 16. Map 19. Cincinalis tomentosa Desv. Berl. Mag. 5: 312. 1811. type: Con- cepcién, Chile, 1782, Dombey r!, photo cu! (see Weath. in Contrib. Gray Herb. 114: 22. 1936). Notholaena hypoleuca Kze. Linnaea 9: 54. 1834; not Goodding, 1912, which is st Grayi. TyPE: near Valparaiso, Chile, Poeppig I 262 (Diar. 41), not seen; isotypes: B!, photo cu, P!, photo cul. : Cheilanthes liopelolcl (Kze.) Mett. Cheil. no. 11. 1859. Rhizome slender 2-3 mm. diam., short-creeping, branched; scales subulate, dark castaneous, selebatic. shining, without pale bow er. Fronds ca. 10-30 cm. tall, approximate; stipe slender, castaneous, glabrous, strongly sulcate as is the rachis, mostly longer than lamina, with one vascular bundle. Lamina narrowly lanceolate, pinnate-pin- natifid or bipinnate (rarely subtripinnate) at base, not or little nar- rowed below, 2-13 cm. long, 1.5-3 cm. wide, upper surface wi thin, lower with dense, ‘lanate tomentum of slender weak much en- tangled hairs, whitish or pale brown. Pinnae 8-12 pairs, rather distant, oblong- lanceolate, narrowed to blunt apex, with about 5 pairs of oblong, obtuse segments or pinnules, the basal only really separated and then often with 2-3 pairs of deltoid obtuse lobes cut % to mid- vein. Veins 1-2-forked, sporangia (with 32 spores) borne on their terminal, subclavate portions, margin not or very slightly modified. Edwyn Reed (x) from Juan Fernandez Islands and Rusby 334 (Ny) from Bolivia are not included on the map, awaiting con- firmation of these range extensions. The record from Antofagasta is taken from Looser in Darwiniana 7: 65. 1945. Chile, in rocky places. Representative specimens: Chile. Bertero 564 (c, Pp); Cuming 200 (B, BM); Macrae 1825 (pm, x). Coguimpo: Rose 19457, 19459 (us). Aconcacua: Cuming 490 (BM, GH, K); Bertero 1249 (c, cu, Pp), 1950 (P); Gaudichaud 30 (3, F, Ps Lechler ae (B); Looser 44, 717 (cx); Rose 19124 (us); Poeppig I 262 (B, BM, G, P). Santiaco: Skottsberg 968 (Bm); Looser 732 (cH). MAULE: ? Stibel 1192 (B). 17. Notholaena wi gens tg Baker, Syn. Fil. be 2, 514. 1874. Weath. Lilloa 6: 273. Fic. 17. Map 20. Cheilanthes Fraseri ‘Male ex Kuhn, Linnaea 34: 83. 1869. TyPE: Ecuador, Fraser Wagner and Peru, Ruiz & Pavon. A sheet at B! (photo cu!) containing both collections, cited and labelled by Mettenius, is taken as authentic. e short or moderately long-repent, often branched and intertangled, 2-3 mm. diam.; scales narrowly linear-subulate with 40 ROLLA TRYON castaneous, subsclerotic, shining central band, narrow paler entire or remotely serrulate margins or pale and hyaline throughout, all with very long capillary, pale brown apex, crisped and contorted in drying (no pectinate teeth seen), excluding capillary apex 1.5-2 mm. long, cells of central band elongate, very small almost without visible lumina. Fronds ca. 20-40 cm. tall, stipes comparatively stout, ap- proximate and forming a loose cluster; terete, mostly about half as long as lamina, with one vascular bundle, dark-castaneous to blackish tomentose with somewhat matted hairs like those of lamina or glabrate. Lamina narrowly linear, about the same width to near apex, not nar- rowed, or not much, at base, acute at apex, pinnate-pinnatifid or bipinnatifid_at base, the pinnatifid tip relatively very short with 2-3 pairs of lobes, lower surface densely tomentose with comparatively coarse much matted pale brown hairs, the upper with a thin covering of fine white hairs or glabrate, texture coriaceous, veins not visible on the upper surface; rachis like stipe, permanently tomentose or rarely glabrate. Pinnae up to 25 pairs, deltoid-ovate, petiolulate, up to 2.5 x 1.3 cm., somewhat articulate, tapering from the commonly inequilateral base to the blunt or acutish apex, pinnatifid 12-% to rachis into 3-5 pairs of deltoid-oblong to narrowly oblong, obtuse lobes, these entire, crenate, or the basal on the lower side of the pinnae more or less elongate and themselves lobed. Veins at acute angle to costa, 1-2-forked, sporangia (with 32 spores) borne at the dilated tips, margin slightly modified. A critical species, separated from the next, N. Buchtienii, by the numerous and tortuous capillary tips of the rhizome scales giving the rhizome a tomentose appearance, the lack of scales on the rachis and upper part of the stipe, the deltoid shape of the median pinnae and the immersed veins. Ecuador to Bolivia, in rocky places, frequently shaded, from 900 to 3600 m. Representative specimens: Ecuador: April 6, 1921, Popenoe (us). Peru. CajaAMarca: Weberbauer 4139 (Bp). HuAnuco: Macbride & Featherstone 2328 (Gc, GH, us). Cuzco: F. L. Herrera 3295 (cu, us); Cook & Gilbert 553 (us); Mexia 8055 (F, cu, us); Vargas 3150, 10449 (F, GH). Bolivia: Stiibel 1233 (8); Bang 1093a, in part (cH). 18. Notholaena Buchtienii Rosenst. in Fedde, Rep. Sp. Nov. 5: 238. 1908. Weath. Lilloa 6: 263. 1941. rypE; Sirupaya, Suryungas, Bolivia, Buchtien 472, presumably at s-pa, not seen; isotype P!, photo cul. Map 21. Rhizome repent, about 3 mm. in diameter, short or up to 10 cm. long, scales narrowly linear, about 3 mm. long, with relatively broad, castaneous, shining, sclerotic central portion and narrow, pale hyaline, 42 ROLLA TRYON subentire or serrulate margin, tipped with a long, weak hair. Fronds approximate, up to 40 em. long; stipe about equalling or shorter than lamina, with a single vascular bundle, castaneous, deciduously tomentose, usually with a few small scales mingled with the tomentum on its upper part. Lamina linear-lanceolate, not or only a little nar- rowed at base, obtuse or acutish at the short pinnatifid apex, below this pinnate-pinnatifid, upper surface more or less arachnoid-tomen- tose with fine white hairs or glabrate, lower surface densely rufescent- tomentose, texture comparatively thin, the veins somewhat im- pressed and usually visible on the upper surface (except as hidden y the tomentum). Pinnae 8-20 pairs, lanceolate or deltoid-lanceo- late, tapering from near the base to the obtuse apex, deeply pinnatifid or the lower pinnate at base, segments oblong or oblong-lanceolate, obtuse, entire or cut not more than % to costule into 2-4 pairs of broad, obtuse lobes. Veins oblique, 1-2-forked, sporangia (with 32 spores in var. Buchtienii), borne on their clavate to flabellate-dilated tips, margin slightly modified. This species is close to the previous one, N. Fraseri. It differs in the few capillary-tipped rhizome scales, the presence of a few (although deciduous) scales on the upper part of the stipe and on the rachis, the lanceolate or deltoid-lanceolate median pinnae and the subimpressed, visible veins. Bolivia and Argentina, in rocky places and on walls, 600 to m. KEY TO VARIETIES Well-developed fronds usually more than 15 cm. tall, indument of the lower surf. ) en a Well-developed fronds to 15 cm. tall, indument of the lower surface pale brown, of the upper densely and persistently whitish-tomento oe eee ona fe le ey ee er ee 18b. var. ventanensis. 18a. Notholaena Buchtienii var. Buchtienii. Fic. 18. Map 21. Notholaena Fraseri var. robusta Hieron. in Engl. Bot. Jahrb. 22: 400. 1896. type: Argentina (Cérdoba), Galander, B! Representative specimens: Bolivia: Pearce (x); R. S. Williams 1374 (us); Cardenas 829 (cu); Buchtien 472 (B, BM, G, P, us), 4225 (F, G, Ny, us). Argentina. Juyvy: Eyerdam & Beetle 22229 (GH). Satta: Venturi 9984 (cu); Lillo 3860 (cu, LIL). TucuMAN: Venturi 750 (GH, LiL, us), 4194 (GH, LiL); Monetti 1661 (GH, LIL). CATAMARCA: 18b. Notholaena Buchtienii var. ventanensis Weath. Amer. Fern Journ. 36: 7. 1946. type: Partido de Tornquist, Sierra de la Ventana, THE AMERICAN SPECIES OF NOTHOLAENA 43 Estancia Funke, Cerros Grietas, November 14, 1943, Cabrera 8101 GH!. Map 21. A local variant, known only from Sierra de Ja Ventana; the following specimen, in addition to the type, belongs here: Lorentz 60 (B, cu, P, PR, US). 19. Notholaena Parryi D. C. Eaton, Amer. Nat. 9: 351. 1875. TYPE: St. George, Utah, Parry 263 y!. Fic. 19. Map 22. se Parryi (D. C. Eaton) Domin, Bibl. Bot. 20 (Heft 85): 133. 191 Rhizome short, erect, stout, often branching and forming dense clumps; scales lance-linear with narrow shining blackish sclerotic cen- tral band and broad entire deep brown thin margins. Fronds clustered, ca. 8-15 cm. tall; stipe slender, castaneous with a few brownish scales toward base, beset somewhat sparsely with weak, divergent, articu- late, often gland-tipped hairs, as long as lamina or nearly so, = one vascular bundle. Lamina mostly lanceolate or narrowly ova 3-12 cm. long, 1.6-4 cm. wide, bipinnate-pinnatifid, beset ppacily above and densely below with long, relatively coarse, loosely curled and intricate, white to pale rufescent hairs which are few aie, the cells very long; rachis similar to stipe. Pinnae 5-8 pairs, the lower distant but not reduced, segments broad-oblong to ovate, very obtuse, the lower deeply 2- to 3-lobed. Veins simple to 2-forked, sporangia (with 64 spores) borne on their clavate terminal portions, margin unmodified. Southwestern Utah to New Mexico, California and Baja Cali- fornia, pi slice or igneous rock crevices and among rocks, 200-28 Representative specimens: United States. UTAH: pilin: a, = (Ge. vs eli 263 (, GH, MO, P, us, ¥); Cottam 4235 (us); M. E. Jones 06 (B, BM, F, GH, P, us), 5110 (us). Nevapa: Coville & Funston 367 (us, x); tomes 613Z (us). Arizona: Shreve 7630 (F); Gold- man 2823 (us); Palmer 555 (Fr, x); Peebles 6486 (us); Harrison & Kearney 6575 (us); Goodding 2130 (cu). Cairornta: Parish 504 B, P, PR, US), 6113 (Pr); Johnston 1046 (us), 2321 (us); Mearns 2805 (us); Eastwood 3019 (us); Howell 3985 (F); Palmer 431 (BM), 432 (BM, F, GH, P). Mexico. Baja Cauirornia: Shreve 6857 (F). 20. Notholaena Newberryi D. C. Eaton, Bull. Torr. Bot. Cl. 4: 12. 1873. Type: San Diego, California, Nov. 8, 1851, Dr. Newberry yl. Fic. 20. Map 23. Cheilanthes =e hit (D. C. Eaton) Domin, Bibl. Bot. 20 (Heft 85): 133. 1913 % So Segunerie < AS frond; x05, gre Pringle 3880 mane = scale hoy rhizom we x 9, from Pringle 5410 (mo). 29 Ob. var. NEALLEYI: A, ide of pinnule and venation, X 4.5; B, scale from Pease *X 9; both from 0. M. Clark 4129 (mo). 56 ROLLA TRYON Tetele, Puebla, Mexico, Arséne 2046 pP!; paratype: same locality, Arséne 1686, not seen. Notholaena hyalina Maxon, Amer. Fern Journ. 5: 4. 1915. TYPE: San José Pass, San Luis Potosi, Mexico, Pringle 3297 (us). Rhizome subhorizontal, short-creeping; scales subulate, black, shining, ca. 3 mm. long. Fronds ca. 15-40 cm. tall; stipe much shorter than lamina, stout, terete, with one vascular bundle, almost black, densely beset with rigid brown spinose-ciliate scales, those of the lower part with pale centers. Lamina bipinnate, to bipinnate- pinnatifid, lance-elliptic, acute or subacuminate, somewhat narrowed at base, broadest about the middle, texture coriaceous, upper surface sparsely beset with weak, simple, white hairs, or glabrate, lower densely covered with stiff-ciliate scales like those of rachis, entirely obscuring the ceraceous indument; rachis like stipe. Pinnae about 20 pairs in well-developed fronds, lanceolate, acute, slightly in- equilateral, the pinnules on the lower side longer, with 7-10 pairs of oblong, obtuse or acutish pinnules mostly cut half-way to midrib into 2-4 pairs of shallow, broadly rounded lobes. Veins simple to 1-forked, sporangia Soe 32 spores) borne on the slightly dilated tips, margin unmodified N. Galeottii is easily distinguished from the related N. Aschen- borniana by the sparse, weak, simple trichomes on the upper surface of the segments; in N. Aschenborniana there are stellate trichomes. Jalisco and San Luis Potosi south to Oaxaca, shaded or exposed rocks and rocky banks, 1500-2200 m. Representative specimens: Mexico. San Luis Porosi: Orcutt 1843 ss Se 5406 ie inet 43 (P); Pringle 3297 (Bs, F, G, GH, , MO, P, y). Jauisco: M. E. Jones 533 (us). Hmatco: Coulter 1 679 (xk); Vv. H. Chase 7199, 7412 (¥); Distriro FEDERAL: Schaffner 84 (B, Ny). Moretos: Arséne (Rosenst. Fil. Mex. 10) (3B, BM, PR); Copeland 98a (us). PureBia: Arséne 2046 (P); Endlich 1980 (8); Copeland 98 (us); Rose, Painter & Rose 10126 (us). GuERRERO: Pringle 13848 (p); Rose, Painter & Rose 9392 (us). Oaxaca: Galeotti 6565 (BM, BR, K); Purpus 4018 (us); Conzatti 475 (cu, Pp), 2003 (F), 4234 (us); Pringle & Conzatti 1392 (cH). 29. Notholaena Schaffneri (Fourn.) Underw. ex Davenp. Gard. & Forest 4; 519. 1891. Weath. Journ. Arn. Arb. 24; 315. 1943. Map 32. Rhizome short, branched, ascending; scales narrow-subulate, al- most black, rigid and shining, strongly pectinate-ciliate. Fronds ca. 5-25 cm. tall, approximate; stipe much shorter than lamina, terete, with one vascular bundle, castaneous to blackish, densely beset with THE AMERICAN SPECIES OF NOTHOLAENA 57 scales similar to those of rhizome but much thinner, brown and with long, stiff shining trichomes. Lamina elliptic-lanceolate, narrowed to acute apex and at base to 2-3 pairs of reduced lower pinnae, bipinnate- pinnatifid throughout, texture coriaceous; upper surface sparsely ceraceous-glandular, lower densely white-ceraceous and with long brown trichomes on the rachillae and midnerves; rachis like stipe. Pinnae up to 30 pairs, oblong- -lanceolate, slightly inequilateral, taper- ing to a rather blunt apex, with up to 15 pairs of linear-oblong obtuse pinnules with 2-5 pairs of suborbicular, broadly rounded lobes. Veins simple to 2-forked, sporangia (with 64 spores) borne on the some- what to distinctly flabellately dilated tips, margin unmodified. Southern Texas south to Puebla, Jamaica; rocky places and shaded ledges, 100-1700 m. The following specimens are intermediate between the two varieties: Veracruz, Mexico, Purpus 6199 (GH, Mo, Us) and Ja- maica, Wilson, Webster & Rogers 428 (Mo, us). TO VARIETIE Rhizome scales narrowly SRR ah 6 densely and conspicuously pec- tinate-ciliate; median pinnae with 4-6 pairs of free epee pe ee eat 29a. ag a en i io gl eal r. Schaffneri Rhizome scales linear-subulate, sparsely and inconspicuously ectioate aa median pinnae usually with 1-3 pairs of free pinn 29b. var. Nealle: 29a. Notholaena Schaffneri var. Schaffneri. Fic. 29a. Map 32. Aleuritopteris Schaffneri Fourn. Bull. Soc. Bot. France 27: 328. 1880. type: San Miguelito Mts., San Luis Potosi, Mexico, Schaffner 2 P!, photo cn!; isotype GH!. ‘Notholaena Nealleyi var. mexicana Davenp. Bot. Gaz. 16: 54. 1891. TYPE: near Guadalajara, Jalisco, Mexico, Pringle 1864 cul. Notholaena Schaffneri var. mexicana Davenp. Gard. & Forest 4: 519. 91. Cheilanthes Schaffneri (Fourn.) Domin, Bibl. Bot. 20 (Heft 85): 133. 1913. Nuevo Leén south to Jalisco and Puebla. Representative specimens: Mexico. Nuevo Leon: Orcutt 1304 (us). Tamautipas: Bartlett 10731, 11007 (us); Fisher 3318 (us). Zaca- TECAS: Rose 2662 (cu, xk, us). San Luis Porosi: Schaffner 962 (cu, K, y). Jauisco: Pringle 1864 (cH, k, Mo, P, y), 3880 (B, BM, F, G, GH, K, MO, P, us, ¥), 11789 (B, F, GH, K, P, US); Palmer 555 (BM, GH, K, NY, P, us, y). Distrrro FEDERAL: Schaffner 30 (B, GH, Ny, P). PUEBLA: Purpus 3146 (BM, F, GH), 4029 (B, us). 29b. Notholaena Schaffneri var. Nealleyi ere Weath. Journ. Arn. Arb. 24; 315. 1943. G. 29b. Map 32. Notholaena Nealleyi Seaton ex Coulter, eee U. S. Nat. Herb. 58 ROLLA TRYON 1: 61. 1890. Type: Limpia Canyon, Jeff Davis Co., Texas, Nealley 560 us!, photo cx! (as published, the collection was said to be 894 from the Chinati Mts. ). oe Nealleyi (Coulter) Domin, Bibl. Bot. 20 (Heft 85): 133. 1 Southern Texas and Coahuila. The literature record in Texas is from Correll in Lundell Flora of Texas 114, 1955 Representative specimens: United States. Texas: Nealley 123 (cu, us), 560 (us); Fisher 6, 305 (us). Mexico. Coanuita: Johnston 7167 (ce); Johnston & Muller 205 (cx). 30. Notholaena Grayi Davenp. Bull. Torr. Bot. Cl. 7: 50, pl. 4. 1880. Weath. Journ. Arn. Arb. 24: 316. 1943. eres Mts. of se. Arizona, Feb. and March, 1880, Courtis cu!, photo m Fic. 30. Map 33. Notholaena hypoleuca Goodding, pieces 8: 94. 1912; not Kze. 1834, which is N. tome ees bei Mule Mts., Arizona, Aug. 1911, Goodding 1004 wus!, photo Cheilanthes Grayi (Davenp.) sda Bibl. Bot. 20 (Heft 85): 133. 1913 Rhizome ascending, branched, forming rather dense clumps; scales subulate, blackish-castaneous, sclerotic, narrow-celled, with filiform but hard, stiff, sparsely pectinate-serrulate apex and below with very but smaller and paler and somewhat glandular, ceraceous, about as long as lamina, with one vascular bundle. Lamina pinnate-pinnatifid, or bipinnate-pinnatifid at base of pinnae, linear-lanceolate, with rather abruptly narrowed obtuse apex, not reduced below, texture coriaceous, pper surface gray-green and sparsely ceraceous, lower with dense ceraceous indument and with pale red-brown, thin, entire-margined scales on midribs and midnerves; rachis like stipe but rather densely ceraceous-glandular and slightly ‘channeled, with scales like those of Jamina. Pinnae somewhat distant, 6-10 pairs, deltoid-ovate to deltoid- oblong, often as wide as long, the basal pirinules longest, obtuse, with about 4 pairs of segments or pinnules below the broad, lobed apex, these oblong, obtuse, be but the lowest oblique and broadly adnate, entire or shallowly, or the lowest somewhat deeply lobed. Veins 1-2-forked, sporangia \(ovith 32 spores) borne on the flabellately dilated tips, margin more or less revolute, unmodifi Southern Texas to Arizona, south to Jalisco, cliffs and rocky slopes, ca. 1000-1500 m. The literature records in Texas are from Correll in Lundell, Flora of Texas 1 1, 1955. THE AMERICAN SPECIES OF NOTHOLAENA 59 Fic. 30-31. Fic. 30. N. Grayi: A, frond, X 0.5; B, upper surface of pinnule, X 4.5; C, underside of pinnule, X 4.5; D, venation of pinnule, X 4.5; all from W.W. Brown 16 (Mo). Fie, 31. N. avrena: A, frond, X 0.5; B, upper surface of pinna, X 4.5; C, underside of segments (one with scales removed, one wit scales removed), X 4.5; all from Pringle 463, in part (cH). Representative specimens: United States. Texas: Ingram 2392 (Pp, us); E. J. Palmer 13504 (xB, us); Nealley 551 S (¥, Ny, us). New Mexico: Feb. 1881, Rusby (cu, x); Feb. 1886, Rusby (B). ARIZONA: April 7, 1884, Pringle (c, cu, Pp, us); July 17, 1881, Pringle (c, cu, P, us, y); Feb. 1881, Rusby (F, p, us, xy); M. E. Jones 4250 (F, G, P, us); 1880, Courtis (Gu, k, y). Mexico. Sonora: Kennedy 7027 (us); Wig- gins 7400 (us). Curmuanua: Pringle 463 (¥, GH in part, Mo, US); Johnston 7880, 7902, 7984 (cu). Coanuita: Palmer 1388 (cH, x, us, y); Johnston ¢> Muller 926 (cu). Stnatoa: Oct. 28, 1904, T. S. Brandegee (cu). Jauisco: Pringle 1864, 5373 (cH). 31. Notholaena aliena Maxon, Contrib. U.S. Nat. Herb. 17: 605. 1916. Weath. Journ. Arn. Arb. 24: 316. 1943. TyPE: Soledad, sw. of Monclova, Coahuila, Mexico, 1880, E. Palmer 1389 vus!, photo cul. Fic. 31. Map 34. Rhizome short-creeping, multicipital; scales subulate, blackish castaneous, sclerotic, narrow-celled, with filiform but stiff apex, the margins freely ciliate with weak hairs, with very narrow pale band below apex. Fronds 8-12 cm. tall; stipe comparatively stout, brown 60 ROLLA TRYON to dark brown, terete, beset with weak, often gland-tipped, hairs and also scales similar to those of the rhizome but paler and less sclerotic, sparingly ceraceous, about as long as lamina, with one vascular bundle. Lamina pinnate-pinnatifid, or bipinnate-pinnatifid at base of pinnae, linear-lanceolate, slightly or not reduced below, texture coriaceous, upper surface slightly ceraceous, with weak whitish hairs (subarachnoid), lower densely whitish ceraceous and midribs and midveins with pale brown, thin, mostly ciliate scales, rachis similar to stipe but slightly channeled. Pinnae ca. 7-10 pairs, lance-deltoid, the basal pinnules longest or nearly so, rather oblong, entire or pin- natifid into broad lobes. Veins simple to 1-forked, sporangia (with 16 spores) borne on the subpunctiform tips, margins slightly revolute, unmodified. This species differs from the closely allied N. Grayi by the whitish hairs on the upper surface of the lamina and the ciliate scales beneath. In N. Grayi the upper surface is ceraceous and the scales beneath are entire. Southern Texas (one collection) and adjacent Mexico, rocky places. Specimens examined: United States. Texas: Cutler 661 (Mo). Mexico. Curmuanua: Pringle 463 in part (cH). Coanura: Palmer 1389 (GH, K, us). TAamau.ipas: Bartlett 10999 (us). 31A. Notholaena Weatherbiana Tryon, sp. nov. Rhizoma breve horizontale, paleis subulatis brunneis vel nigrescen- tibus plerumque scleroticis marginibus longe ciliatis apice filiformi. Frondes 6-15 cm. altae caespitosae, stipes teres rubelle castaneus vel saturate brunneus quam lamina brevior vel aequalis fasciculo vasculari uno tomentosus paleaceus paleis eis rhizomatis similibus nisi plerum- que pallidioribus et rion scleroticis. Lamina anguste lanceolata vel. lanceolata bipinnato-pinnatifida coriacea pagina superiore tomentosa pagina inferiore dense tomentosa ceraceo-glandulosa pallide flava paleis paucis, rhachide eis stipitis similibus nisi leviter canaliculatis. Pinnae 6-11-jugae deltoideae vel oblongo-deltoideae plus minusve subaequilaterales 4-6-jugis loborum obtusorum. Venulae 1-2-furcatae porangia 32-sporis in apicibus gerentes, margem immutatus leviter revolutus. Typus: Southwestern Chihuahua, Mexico, Aug.-Nov. 1885, E. Palmer 215 mo!. This species was not recognized until the manuscript was too advanced to allow a map and drawings to be completed. The figure of the leaf of N. aliena, its closest relative, may be taken as representing the general aspect of this species. From N. aliena THE AMERICAN SPECIES OF NOTHOLAENA 61 it is sufficiently distinguished by the tomentum on the rachis and upper and lower surfaces of the lamina and the lack of ceraceous glands on the upper surface. In addition to the tomentum, the rachis is moderately scaly and the lower surface of the lamina has a few scales. The single specimen seen bears no information as to the habitat or the exact locality. Specimens at B, k and us of this collection are N. incana. Specimen seen: Mexico. Cummuanua: Palmer 215 (Mo). 32. Notholaena Lemmonii D. C. Eaton, Bull. Torr. Bot. Cl. 7: 63. 1880. Type: Santa enim ie near Ft. Lowell, Arizona, Pits 18, 1880, Lemmon yY!, photo AP 35. Rhizome short, subhorizontal or ascending; scales lanceolate or subulate, dark-castaneous sclerotic and shining, cells not visible, with stiff capillary tip and narrow pale margin with weak cilia. Fronds 10-35 tall, clustered; stipe rather stout, shallowly and broadly silcath shee Boars to blackish, glabrous except for a few ovate, acuminate, cordate-based scales with brown, sclerotic center and broad, pale, ithe; “thi margins, shorter than lamina, with o vascular bundle. Lamina lanceolate or linear-lanceolate or hia: with 10-15 seis of pinnae, Lisgeete or partially bipinnate, not narrowed at base or with o fan ir of somewhat reduced pinnae, rather abruptly Gacremed § ute apex, coriaceous, upper surface glabrous, the lower densely ahae or pale yellowish ceraceous; rachis like stipe, glabrous, strongly sulcate above. Pinnae deltoid to lance- olate, somewhat inequilateral with the lower side broader, broadest near base, acute or subacuminate, with 5 to 7 pairs of linear-oblong, obtuse segments which are broadly adnate and sometimes dilated at base and shallowly lobed, the lobes broad and rounded and connected by a narrow wing along the costa, or the lowest pair separate. Veins 1-2-forked, sporan a (with 64 spores in var. Leer borne on the flabellately dilated tips, margin revolute, unmodified Southeastern Arizona to southern Baja California; south-central Mexico; on slopes and cliffs of igneous rocks, ca. 1000-1500 m. KEY TO VARIETIES Mature stipe and rachis blackish or dark castaneous, rhizome scales lance- Be subulate, 2-mm..or more long. +). .)siv7:. sc oe see. mmonii. Mature stipe and rachis pale castaneous, rhizome scales lanceolate, scarcely Bae es oe a ca Lee i ee ne 32b. var. australis. 32a. Notholaena Lemmonii var. Lemmonii Fic. 32. Map 35. Cheilanthes Lemmonii (D. C. Eaton) Domin, Bibl. Bot. 20 (Heft 85): 133. 1913. 62 ROLLA TRYON Southeastern Arizona to southern Baja California. Representative ee gi States. Arizona: Spring of 1880, Lemmon (¥F, GH, K, MO, US, n 1881 (BM, G, GH, K, P, us); 1881, Pringle (GH, K, Mo, P, US). Moxics. Baya Catirornia: Brandegee 6 (y), 659 (cH, us); Johnston 4019 (us). Sonora: Palmer 266 (3M, GH, K, us, ¥); Drouet & Richards 3773 (x), 3809 (vy); Wiggins 6160 (us), 6479 (F, GH, us), 7063 (us), 7337 (cH, Us). Crmauanua: Palmer 84 (BM, GH, K, MO, US, Y). 32b. Notholaena Lemmonii var. australis Tryon, var. nov. Map 35. A varietate typica differt stipitibus thachidibusque maturis pallide weet paleis rhizomatis lanceolatis vix 1 mm. longis. pus: Coxcatlan, Puebla, Sept. 1909, Purpus 4178 us!. Other aia examined: Mexico. Puresia: Schenck 161 (B); Purpus 4026 (B). Oaxaca: Rose, Painter & Rose 10093 (us); Aug. 14, 1910, Hitchcock (us); Kenoyer 1620 (cu). 33. Notholaena Rosei Maxon, Contrib. U.S. Nat. Herb. 16: 59. 1912. type: Chapala, Jalisco, Mexico, Oct. 5, 1903, Rose & Painter 7665 us!, photo cul. Fic. 33. Map 36. Notholaena Lemmonii var. straminea Davenp. Gard. & Forest 4: 519. 1891. Type: near Guadalajara, Jalisco, Mexico, Pringle 2830 cul. Rhizome _ short-horizontal; scales lanceolate, acuminate, dark brown or blackish, shining, sclerotic with very narrow irregular, pale, m en 10-50 cm. tall; stipes stoutish, castaneous at base, stramineous above, somewhat sulcate, with short, stiff gland-tipped hairs on upper side, glabrous on other faces, shorter than lamina, with one vascular bundle. Lamina lanceolate, broadest at base, coriaceous, upper surface spar- ingly and deciduously whitish ceraceous, the lower densely so; rachis glandular above, like stipe. Pinnae petiolulate 10 or more pairs, the lower 2 or 3 pairs reduced, the basal greatly so, lanceolate or deltoid- lanceolate, somewhat inequilateral, the lower side widest, fully pin- nate toward base, with pinnatifid apex, lower pinnules linear-lanceo- late, deeply pinnatifid below into 3 or 4 pairs of obliquely deltoid- ovate, obtuse lobes with entire obtuse tip like that of pinna. Veins simple to 1-forked, sporangia (with 64 spores) borne on the clavate tips, margin revolute, slightly modified. Rocky hillsides near Guadalajara, Jalisco. Specimens examined: Mexico. Jatisco: Pringle 2830 (cu); Rose U Painter 7665 (Ny, us); Palmer 701 (8M, GH, K, MO, NY, P, US, Y 32 . a os a ae ’ | {RS Dyan 32-D 33-A 35°C 34°A G. 32-35. Fic. 32. y: LemmMont var. Lemmonu: A, frond, X 0.5, from Pringle in 188i (mo); B, portion of upper side of scare en nlarged, from Wiggins & Rollins 303 (mo); C, ae of pinnule a, ve nyo 40; D, — from rhizome, X 18; the last two from Pringle in 1881 tha Fic. 33. N. Roser: A, frond, X 0.5; B, portion of = enlarged; — portion of upper side of rachis enlarged; D, underside of segmen ~ — X 4.5; all from Palmer 701 (mo). Fic. 34. - LEONINA: A, fro ; B, goareers - pinnule nih X 4.5; both from Palmer 1381, r* ay as B (mo). Fic. 35. N. GaLapacensis: A, frond, X ; B, underside of segment and venation, X 4. 5; C, scale fet rhizome, X 9; all from Stewart 930 (cx). 7 ea ROLLA TRYON . Notholaena leonina. Maxon, Contrib. U.S. Nat. Herb. 16: 58. 19d TYPE: oe Nuevo Leon, Mexico, Feb. 1880, Palmer 1381 us!, photo c Fic. 34. Map Rhizome short, ascending or horizontal, branched; scales lance- subulate with central castaneous, sclerotic, shining, central band and paler, a glandular-ciliate or papillate, thin margin, tipped wi weak, deciduous gland-tipped hair, the glands apparently pret wax. Fronds 5-10 cm. tall, numerous and clustered; stipes slender, blackish with a few ovate brown scales, pgs 9 ohm like those of rhizome and somewhat ceraceous-glandular, longer than lamina, with one vascular bundle. Lamina deltoid to deltoid-oblong, -— Maps 35-39. Map 35, N. Lemmonii, large dots var. Lemmonii, small dots var. = ete Map 36, N. Rosei. Map 37, N. leonina. Map 38, N. aban Map 39, N. a large dots var. candida, small dots and x’s var. Copelandii, half dot intermediate THE AMERICAN SPECIES OF NOTHOLAENA 65 narrowed rather evenly and shortly to the blunt tip, bipinnate to bipinnate-pinnatifid or nearly tripinnate at base, texture coriaceous, upper surface thinly, lower densely, ceraceous. Pinnae about 5 pairs, the upper ovate, obtusish with 2-3 pairs of oblong or ovate-oblong, obtuse, entire adnate segments, the lowest distant with the basiscopic pinnules somewhat produced and deeply lobed or pinnate. n 1-forked, sporangia (with 32 spores) borne on the flabellately dilated tips, margin strongly revolute, unmodified. Monterrey, Nuevo Leon. Specimens examined: Mexico. Nuevo Leon: Palmer 1381 (cu, x, MO, P, US, Y). 35. Notholaena galapagensis Weath. & Svenson, Bull. Torr. Bot. Cl. 65: 319, t. 9, fig. 2. 1938. rypE: Iguana Cove, Albemarle Is., Galapagos Islands, Dec. 30, 1899, Snodgrass & Heller 23 cu!. Fic. 35. Map 38. Rhizome short, oblique; scales bright-castaneous to brownish, shin- ing, lance-subulate, acuminate, with a very narrow, almost obsolete, paler, serrulate margin which sometimes bears a few cilia, a weak, short, articulate, early deciduous trichome at tip, texture sclerotic, cells narrow, without lumina, their walls thick and some- what prominent. Fronds 7-15 cm. tall, clustered; stipes slender, castaneous, somewhat shining, terete, near the base with lance-ovate, cordate-based, acuminate, thin, slightly serrulate, nearly concolorous scales their cells elongate-linear except at base, with thin walls, longer than lamina, with one vascular bundle. Lamina ovate or deltoid- equilateral with 1 or 2 basal pairs of pinnules free and 2 or 3 on the ower side at base somewhat elongate, linear-oblong and deeply lobed, the apex produced into a narrow, shallowly pinnatifid tip, blunt or acutish at the extreme apex. Veins simple or 1-forked, sporangia (with 64 spores) borne on the clavate to flabellately dilated tips, margin slightly revolute, unmodified. The lighter colored scales and stipe, the narrowly margined scales and the yellowish indument separate this species from the related N. candida. Galapagos Islands, in rocky places up to 500 m. Representative specimens: Galapagos Islands. Cuming 110 (cH, K). 66 ROLLA TRYON ALBEMARLE: Snodgrass & Heller 23 (cu); Stewart 930 (cu, us), 931, 932 (us). cg Gare Svenson 79 (us). JAMEs: Scouler (Kk). NARBOROUGH: Stewart (Cas). 36. Notholaena candida (Mart. & Gal.) Hook. Sp. Fil. 5: 110. 1864. Map 39. Rhizome short, horizontal, branched; scales lanceolate or lance- subulate, 4-5 mm. long, central dark brown to blackish, sclerotic band rather narrow, paler margin entire to ciliate. Fronds rather few, mostly 10-30 cm. tall; stipe stoutish, blackish to black, terete, rather dull, its scales at base ovate-lanceolate, with a brown center and pale margins, cordate, somewhat longer than the lamina, with one vascular bundle. Lamina ovate-pentagonal, except for the lowest pinnae, bipin- nate-pinnatifid, acute, the apex strongly produced, or not and acuminate, coriaceous, upper surface glabrous, the lower densely white ceraceous. Pinnae 5-10 pairs, lanceolate or linear,.acute or acuminate, with 10-12 pairs of linear-oblong, subacute, adnate seg- Sri, these entire, crenulate or often broadly and obtusely lobed % midribs. Veins 1-2-forked, sporangia (with 64 wot borne on ‘ks flabellate tips, margin somewhat revolute, unmodified Southern Texas to Sonora, south to Honduras. O VARIETIES Rhizome scales slightly toothed or entire; basal pinnae with two to five elongated and pinnatifid lower ee the other pinnae rather pat tapering from base to apex; apex of the lamina gradually and evenly con- ein RE Boe 5 CE a OE ee ETE var. candida. Rhizome scales ciliate; basal pinnae with a single greatly ribodipined and eeply pinnatifid inner lower pinnule, the other pinnae with more or less parallel sles to near the apex; apex of the lamina abruptly contracted to a more or less well defined terminal pinna............ 36b. var. Copelandii. 36a. Notholaena candida var. candida. Fic. 36a. Map 39. Cheilanthes candida Mart. & Gal. Mém. Acad. Brux. 15: 73, t. 20, fig. 1b. 1842. TYPE: eager Jalisco, Mexico, Galeotti 6442 sri. Another specimen of the same number from “Jalisco” is at Br! and others from “Oaxaca and Jalisco” at BR! and x!. Aleuritopteris candida (Mart. & Gal.) Fée, Gen. Fil. 154. 1850-52. Ceropteris monosticha Fée, Mém. Fam. Foug. 7: 44, t. 22, fig. 2. 1857. type: Orizaba, Schaffner 155, not seen. Cheilanthes furfuracea Presl ex Koch, Wochen. Gartn. Pflanzenkr. 1: 3. 1858, based on Cheilanthes candida Mart. & Gal. Allosorus ei bat ial Presl ex Koch, Wochen. Girtn. Pflanzenkr. 1: 3. 1858, i Cheilanthes monostiche (Fée) Mett. Cheil. no. 4. 1859. Notholaena monosticha (Fée) Moore, Ind. Fil. 231. 1861. Notholaena sulphurea var. alba Mett. ex Kuhn, Abhandl. Ges. Halle YY Ds ~~ e a6a-A Fic. 36-38. Fic. 36. N. canpma: 36a. var. canpipa: A, frond, X 0.5; B, scale from rhizome, X 9; both from Palmer 82 (mo). 36b. var. CopeLanpu: A, frond, 5; B, segment, X 4.5; C,-scale from rhizome, X 9; all from Meyer & Rogers 2496 (mo). Fic. 37. N. sucpHurea: A, frond, X 0.5; B, underside of segment and venation, X 4.5; C, scale from rhizome, X 9; 0) Fic. 38. N. SranpLeyi: A, frond, X 0.5; B. underside of segment, X 4.5; C, pinna showing venation, X 2.5; all from Cutler 741 (Mo). 68 ROLLA TRYON 11: 32. 1869, ex synon. princip., “Mexi sora erage candidum (Mart. & Gal. re Kiimm. Magyar Bot. Lapok. 13: 42. 1914. The relation of var. candida and var. Copelandii is discussed under the latter variety. Western Mexico from northern Sonora to Jalisco, then to Hidalgo and south to Map 56 , N. Fendleri. Map 59, N. delicatula. Map 55, var. limitanea. » N.. limitanea. » N. dealbata. Map 58 Maps 54-60. Map 54 mexicana. Map 57 60, N. incana. THE AMERICAN SPECIES OF NOTHOLAENA 89 Rhizome short, slender; scales linear, pale brown, entire, thin and delicate, concolorous, ca. 3 mm. long. Fronds clustered, numerous, mostly 5-15 cm. tall: stipe ise slender, terete, glabrous, shining, bright chestnut, as long as or longer than the lamina, with one vascular eee Lamina broadly to narrowly deltoid, often nearly as broad as long, 4-5-pinnate, with 3-5 pairs of pinnae, upper surface glabrous or very slightly white-ceraceous, glaucous, the lower white or rarely yellowish ceraceous, usually densely so. Pinnae deltoid, long-petiolulate, segments small, ovate to oblong, entire, non-articu- late, dark color of the stalk sient into the segment. Veins 1-2-forked, sporangia (with 64 spores) borne (one or two of them) back of the vein-tip, margin unmodified. Nebraska and Missouri south to central Texas, exposed or shaded calcareous cliffs and ledges; the literature records in Missouri are from Steyermark (ms.). Representative specimens: United States. Missourt: Bush 132 (F, us), 798 (B), 819 (cu, K), 850, 7706 (¥F, cu, us); E. J. Palmer 30421, 34733, 39290 (Fr, us); Steyermark 8371 (mo), 5749, 22502, 22966 (F). Arkansas: E. J. Palmer 4538 (us), 5951 (F, us), 17200 (x); Harvey 22, 39 (us, y). Nepraska: Nov. 10, 1895, Williams & Wilcox - K, us). Kansas: Fendler 1017b (8, Mo); Hitchcock 963 (cu, Mo, Y); E. J. Palmer 41806 (us). OxLAHoMa: Stevens 48 (G, GH, K, US); Cartton — ips Texas: Lindheimer 608 (B, Mo), 1279 (MBG dist.) » G, us); Reverchon 1183 (Fr, us); E. J. Palmer 10377 (vs), i114 ‘esi "11805 (cu, us), 14200 (8); Jermy 670 (us); Tharp 2162 (vs). 53. Notholaena Fendleri Kze. Farnkr. 2: 87, t. 136. 1851. TYPE: Trapp formations of the Rio del Norte near Santa Fe, New Mexico, May, 1847, Fendler 1017a, a originally = Lz and now de- stroyed; isotypes: B!, BM!, cu!, K!, Mol. Fic. 53. Map 58. Gymnogramma F endleri (Kze. ) Mott Cheil. footnote 13. 1859. Pellaea Fendleri (Kze.) Prantl, in Engl. Bot. Jahrb. 3: 417. 1882. Cincinalis Fendleri (Kze.) Fée, Gen. Fil. 160. 1850-52. AE agit stout, often horizontal or oblique; scales linear, chestnut brown, entire, thin, concolorous, 5 mm. or more long. ronds numer- ous, a clustered, ca. 10-30 cm. long; stipe relatively stout, terete, castaneous, glabrous, as long as or longer than the lamina, with one vascular bundle. Lamina broadly deltoid, often broader than long, 4-6-pinnate, upper surface glabrous or ceraceous-glandular, the lower densely sus Phare rachis (and pinna-rachises and costae) markedly flexuous, the divisions of the next order arising om the angles and seit divergent so as to give the snopes of dichotomous branching. Pinnae alternate, up to 11 of them, 90 ROLLA TRYON broadly deltoid, both they and the pinnules long-petiolulate, ulti- mate segments ovate to oblong, obtuse, broadly cuneate at base, non-articulate, dark color of stalk saree into the base. Veins 1-2- forked, sporangia (with 64 spores) borne (usually one or two of them) back of the vein-tip, margin unmodified. The strongly flexuous Wd and other axes make this an excep- tionally well-marked specie Southeastern Wyoming ak to New Mexico, dry rocky bluffs and cliffs, ca. 1750-2500 m. Representative specimens: United States. Wyominc: Nelson 2020 (mo, Ny). Cotorapo: M. E. Jones 12 (s, Bm, Pp); Hall & Harbour 693 (F, GH, MO, x); Clokey 3287 (¥, GH, Mo, us); Clements 73.2 (3, GH, Mo, us); Patterson 163 (8, F, GH, K, MO, P, US). NEW MEeExIco: Fendler 1017a (B, BM, GH, K, Mo); Ragliston 16960 (us); Arséne & Benedict 15433 (Fr, us); O. M. Clark 9669, 9704, 10035 (mo); Griffiths 553 (us). 54. ee delicatula Maxon & Weath. Contrib. Gray Herb. 127: 7. 1939. Weath. Journ. Arn. Arb. 24: 312. 1943. Tyre: Lerios, 45 miles east of Saltillo; July, 1880, Palmer 1387 us!. Fic. 54. Map 59. Rhizome short, erect or oblique; scales slender, brown, concolorous, linear-subulate, long-acuminate, about 4 mm. long, entire. Fronds clustered, 10-35 cm. tall; stipe castaneous, slender, terete, glabrous, mewhat shining, nearly equal to the lamina, with one vascular eae Lamina mostly deltoid, tripinnate or toward base nearly quadripinnate, herbaceous, the upper surface minutely ceraceous- Pinnae about 4-8 pairs, deltoid or narrowly so, terminal segments rhombic or flabelliform, others roundish, oblong or subdeltoid, seg- ments articulate where the dark color of the stalk abruptly paseo at the base of the segment. Veins 1-2-forked, sporangia (with 64 spores) borne on nearly the whole length, or on the apical % of the ed. vein, margin unmo As Maxon and Weatherby mentioned in their publication of this species, it is not entirely distinct from N. incana, although from the few collections available it seems sufficiently so. Addi- tional material may indicate it should be considered a geographic variety of N. incana, or may confirm its specific status. Coahuila and Nuevo Leon, in exposed or shaded, rocky places, 1200-3400 m. Specimens examined: Mexico. Coanurta: Palmer 1385 (GH, Mo, US, xy), 1387 (cu, k, P, us, y); Johnston 9046 (GH). Nurvo Leon: Palmer 1386 (cH, P, US); Pringle 2581 (B, BM, GH, K, MO, P, Us, Y). Oh oD a —\~ aw BV eR yee Q ho [eLo, SO OG awy7 / ORY ODP > q WS ¢) oO QO MH IOg () Sw% = Sol cA AD 9 OOS? PEROT =. TY ORED ON AVS NGS ZR Ss £K) 5 Noe dy 4 ee. We My a we aS aN YY & Qo es aN a YOU TS | ? FG Se e PARR 4 p Me AABN a 07 Ds / II & CIN OB 5 of ay MS TOG ‘ eI “Fag <> i UES VY apt la = - TELL Oy | So “OM 5 TIS O/ &»* AS ae 9 DP 2 CV ae = KY HOS 54-4 53-D 55°A 55°C G. 54- 553 Pid; es the DELICATULA: A, frond, X 0.5, from Pringle “hed ets , X 7, from P Palmer 1385 ( 0). Fic. 55. N. frond, X 0.5; B, u scile of segment, X 4.5; C, segment, X 4.5; D, hte “iat rhizome, Gy & all sis Lyonnet 35 (Mo). 92 ROLLA TRYON 55. Notholaena incana Presl, Rel. Haenk. 1: 19, t. 1, fig. 2. 1825. Maxon & Weath. Contrib. Gray Herb. 127: 5. 1939. Lectotype: (by Maxon & Weath. l.c.): Mexico, Haenke pr!, photo cul. F I 60 Gymnogramma candida Mett. Cheil. footnote 9. ace LECTOTYPE: Valle de Mexico, Schmitz 231 B!, photos BM!, cul, Pellaea candida (Mett.) Prantl, in Engl. Bot. Jahrb. 8: 417. 1882. Rhizome short, thick, erect or oblique; sr chestnut posal con- colorous, linear-ligulate, long-attenuate, entire, up to long. Fronds clustered, mostly 15-40 cm. tall; rise: rather stout, lacks shining, terete, glabrous, usually about equalling the lamina, with one vascular bundle. Lamina rather narrowly deltoid, tripinnate, at least toward base, with 6-9 pairs of subopposite pinnae, coriaceous, glabrous and gray-green above, densely white-ceraceous beneath. Pinnae deltoid to ovate, long-petiolulate, ultimate segments broadly oblong to ovate, broadly ‘obtuse, truncate or subcordate at base, articu- late where the dark color of the stalk abruptly terminates at the base of the segment. Veins 1-2-forked, sporangia (with 64 spores) borne on the whole length, or on the apical % of the vein, margin un- modified. Maxon and Weatherby (I.c.) have discussed the type-specimen of N. incana and the application of the name. Dr. Maxon was prepared to name this segregate of N. nivea for Mettenius and such a name appears on a number of labels but the identity of Presl’s name was established and the manuscript name never published. Sonora and Chihuahua south to Guatemala; Hispaniola; on cliffs or rocky banks, or in clay soil, ca. 1800-3000 m. A specimen of this species at Kew: Santa Rita, Arizona, March, 1881, Lemmon is provisionally questioned as to locality and not mapped; the species is not reported by Morton in Kearney & Peebles, Arizona Flora, 1951. Representative specimens: Mexico. Sonora: Drouet et al. 3667 (F, MO); Shreve 6735 (mo); Schott 16 (¥). Curauanua: Palmer 115 (c, GH, xy), 215 (BM, k, us). San Luts Potosi: Schaffner 961 (c, GH, K); Virlet d’Aoust 21 (p). Jatisco: Barnes & Land 132, 231 (F). Micuoacan: Arséne 5341 (Pp), 5756 (us). Mexico: Pringle 5248 (Mo, P), 11267 (B, G, GH, K, us), 15021 (c, us); Bourgeau 754 (c, K, p); Distrito FEDERAL: Schaffner 234 (B, K, P), 262 (Kk); Hinton 4195 (F, GH, MO), 6244 (F, Mo). Moretos: Pringle 15698 (us). PuEBLA: Arséne 1993, 2145 (ep). Guerrero: Mexia 9014 (F, GH). Oaxaca: Ghiesbreght 415 (re); Conzatti & Gonzales 463 (G, GH). CHIAPAS: Ghiesbreght 226 (c, cu, K, ¥). Guatemala: Salvin 238 (c, cu); Ber- THE AMERICAN SPECIES OF NOTHOLAENA 93 noulli & Cario 251 (B, Kk, Pp). Dominican Republic: Ekman 13770 (us); Ri AS tr ES. Habird 9113 (us). 56. Notholaena nivea (Poir.) Desv. Journ. Bot. Appl. 1: 93. 1813. Maxon & Weath. Contrib. Gray Herb. 127: 8. 1939. Map 61. Rhizome short, thick, erect or sgn onlioes scales concolorous, castaneous, linear- subulate 2.5-3 mm. long, and delicate, often much crisped in drying, margins entire or with the walls of the marginal cells craieceing, Fronds crowded, mostly 10-30 cm. tall; stipe rather slender, bright to dark castaneous, terete, glabrous, usually dull, shorter than or about equalling the lamina a, with one vascular bundle. Lamina lanceolate or deltoid- lanceolate to ovate, sub- tripinnate, with up to 12 subopposite pairs of pinnae, coriaceous, the upper surface glabrous, the lower densely yellow- or white-ceraceous or glabrous; rachis like stipe. Pinnae petiolulate, ovate to lanceolate, pinnules rather long, petiolulate, ultimate segments oblong to sub- orbicular, obtuse, subpetiolulate at the truncate to subcordate base, entire, articulate where the dark color of the stalk abruptly sera at the base of the segment. Veins 1-2-forked, sporangia (with 3 spores in all varieties), borne on the apical % to % of the vein, ieiga unmodified. Maxon and Weatherby (I.c.) have discussed the variability of the varieties and some of the intermediates between them. It is evident that none of the variations are entitled to specific rank and equally so that they are entitled to recognition of some kind. Considering the number and kinds of characters involved and the fact that each variety has a reasonable range, the varietal treatment seems justified. Colombia to Argentina and eastern Brazil, in crevices and on pe of cliffs, on rocky banks, less often in clay soil, ca. 1300- KEY TO VARIETIES a. Indument white b Lamina more or r less completely tripinnate, ultimate segments s sub- Orisped eres ee ee PT peer es 56b. var. oblongata. a. fag yellow or none, ultimate segments oe to elliptic-oblong. c. Indument none, lamina Sully ony Lapeer e except toward base, ter- minal segments entire to | obed, s e castaneous to ei gr eg rhizome scales not strongly crisped..............-.. c. var. tenera. c. Indument yellow, lamina fully tripinnate when Sales dooedl terminal segments usually entire, stipe dark-castaneous, rhizome scales not Creep es pe er ee 56d. var. flava. 94 ROLLA TRYON Maps 61-65. Map 61, N. nivea. = 62, var. nivea. Map 63, var. oblongata. Map 64, var. tenera. Map 65, var. flav THE AMERICAN SPECIES OF NOTHOLAENA 95 56a. Notholaena nivea var. nivea. Maxon & Weath. Contrib. Gray Herb. 127: 8. 1939 (as N. nivea). Fic. 56a. Map 62. Pteris nivea Poir. Encycl. 5: 718. 1804. Type: Peru, Jos. de Jussieu (Hb. Jussieu sheet no. 1047) v!, photo cul. Acrostichum albidulum Sw. Syn. Fil. 16, 205, t. 1, fig. 2. 1806. TYPE: Née, specimen so named, “Cav. misit” in Hb. Swartz, s-pal. Cincinalis nivea (Poir.) Desv. Berl. Mag. 5: 313. 1811. Gymnogramma nivea (Poir.) Mett. Cheil. footnote 11. 1859. Pellaea nivea (Poir.) Prantl, in Engl. Bot. Jahrb. 3: 417. 1882. Cincinalis tarapacana Phil. Anal. Mus. Nac. Chile Bot. 1891: 91. 1891. Type: Sibaya, Tarapaca, Chile, Philippi, not seen; isotypes B!, x!. Pellaea nivea f. nivea (Poir.) Hieron. in Engl. Bot. Jahrb. 22: 390. 896. Notholaena albidula C. Chr. Ind. Fil. 459. 482. 1906, nomen in synonymy; basonym A. albidulum Sw. implied but combination not made by Sturm as cited and juvstionabity made by C. Chr. Ecuador to western Argentina. Representative specimens: Ecuador: Spruce 5632 (B, BM, GH, K, NY, P); Heinrichs 36 (B, Ny); Mille 171 (ep). Peru. ANCACHS Hs: Webe rbauer 3009, 3061 (8). HuAnuco: Ruiz 45 (B). Junin: Killip & Smith 21812 (Ny, us), 22161 (us). Lima: Macbride 3207 (Fr); Matthews 755 (x, P); Safford 990 (cu, Pp, Ny, us); Killip & Smith 21561 (cu, Ny, us); Weberbauer 135 (8). Huancave ica: Stork & Horton 10823 (F). Cuzco: Weberbauer 4866 (B); Hitchcock 22528, 22549 (us); Herrera 144 (us); Cook & Gilbert 196, 221 (us). Apurntmac: Stork & Horton 10718 (¥). Puno: Mexia 7788 (¥F, GH, MO). Bolivia: Shepard 181 tig NY, P, us); Buchtien 600 (Ny, us), 1142 (us); Bang 19 (B, BM, GH, K, NY, P, us), 2600 (cu, Ny, us); Fiebrig 3024 (B, BM, GH, K, P, US). Chile: Sibaya, Philippi (8, Kk); Werdermann 1068 (B, K, NY, us). Ar- gentina. Los ANDES: Catalano 2 (cH); Budin 7 (cH). Satta: Lorentz & Hieronymus 53 (B, us). TucuMAn: Lorentz & Hieronymus 955 (3). Catamarca: Castellanos 30/313 (cH). La Rioja: Hieronymus & Niederlein 555 (B). Corvosa: Stuckert 1764 (Pp). San JUAN: Jan. 1876, Echegaray (B). 56b. Notholaena nivea var. oblongata Griseb. Abh. Konig. Ges Wiss. Gott. 24 (Symb. FI. Argent.): 342. 1879. Maxon & Weath. Contrib. Gray Herb. 127: 10. 1939; Weath. Journ. Arn. Arb. 27: 369. 1946. type: said to be from Salta, not seen; a specimen at k! “comm. Grisebach 1878” is taken as authentic. Fic. 56b. Map 63. Southern Peru; Argentina; Santa Catharina, Brazil. Representative specimens: Peru. Cuzco: Soukup 76 (cH). ApuRI- MAC: Herrera 1498, in part (GH). Puno: Lechler 1830 (8). Argentina. Tucuman: Burkart 5168 (cu); Venturi 1649, in part (GH, us). Cata- 96 ROLLA TRYON marca: April 12, 1910, Castillon (cu, p). Corposa: Lossen 242 (PH); Osten 10510 (s-pa). Brazil. Santa CaTHARINA: Spannagel 172 (Ny, PH). 56c. Notholaena nivea var. tenera (Hook.) Griseb. Abh. Konig. Ges. Wiss. Gott. 24 (Symb. FI. ees 342. 1879. Maxon & Weath. Contrib. Gray Herb. 127: 11. Fic. 56c. Map 64. Notholaena tenera Gill. ex Hook, in Curtis, Bot. Mag. t. 3055 & text. 1831. TYPE: a Sunes specimen from spores sent by Gillies from Mendoza, Argentina, k!, photo cul. Cincinalis eae, (Hook) Fée, Gen. Fil. 160. 1850-52. Pellaea tenera (Hook.) Prantl, in Engl. Bot. Jahrb. 3: 417. 1882. Pellaea nivea f. tenera (Hook.) Hieron. in Engl. Bot. Jahrb. 22: 390. 1896. Notholaena tenera var. major Christ ex , Stuckert, Anal. Mus. Nac. Buenos Aires s. 3, 1: 300. 1902. TyPE: Rio pee vicinity of Cérdoba, Stuckert Hb. Argent. 5971, not seen; isotype Pellaea peruviana Copel. Univ. Cal. Bot. Publ. 19: 302, 't. 59. 1941. Type: Abancay region, Apurimac, Peru, Oct. 1935, pupils of V. Santander C. uc!, photo uc! Central Peru to Argentina. Representative specimens: Peru. haha Killip & Smith 22162 (ny, us). Lia: Savatier 589 (x); Macbride & Featherstone 82 (¥, GH); Safford 992 (us); Killip & Smith 21578 (ny). Cuzco: Herrera 717 (us), 3156, 3174 (¥). Bolivia: Mandon 1550 (3M, GH, K, Ny, P), 1863 (8); Rusby 326 (cH, Ny, us), 327 (Ny, us); Buchtien 1141 (vs), 3112 (cu, us). Argentina. Juyuy: Venturi 4931 (cu, us); Schulz 974 (cH). TucumAn: Lorentz 888 (B); Venturi 1064 (cu, us), 10369 (Ny). CatTaMArca: Schickendantz 44 (p), 364 (Bn); Jorgensen 1237 (cH La Rioja: Hieronymus & Niederlein 439, 732 (Bp). CorDoBA: Stuckert 5971 (Pp); Lossen 242, in part (Mo). MeEnpoza: Wilcheck 2 (Pp). Buenos Arres: Bettfreund F006 (B). 56d. Notholaena nivea var. flava Hook. Sp. Fil. 5: 112. 1855. Maxon & Weath. Contrib. Gray Herb. 127: 12. 1939; Weath. Journ. Arn. Arb. 27: 370. 1946. Type: uncertain, but identity clear. Map 65. Acrostichum flavens Sw. Syn. Fil. 16, 204. 1806. type: South Amer- ica, Née, presumably at s-pa, not seen, but identity not doubtful. Acrostichum tereticaulon Desv. Berl. Mag. 5: 310. 1811. type: (per- haps Peru, Dombey) P!; see Weath. Contrib. Gray Herb. 124: 14. Cincinalis (?) flavens (Sw.) Desv. Berl. Mag. 5: oe sie Gymnogramma flavens (Sw.) Kaulf. Enum. 77. Notholaena chrysophylla Kl. Allg. Gartenzeit. 23: ‘965. 1855. TYPE: cultivated — said to have been originally collected in Peru by Warszewicz, Machado: pie (Sw.) Moore, Ind. Fil. Ixx. 1857. . 56-58. Fic. 56. N. EA: 56a. var. NIvEA: A, frond, X 0.5; B, sao ae se Polat X 4.5; both “ea fice 600 (mo). 56b. var. OBLONGATA: under of ens X 4.5, from Lossen 242, in part (Mo). 56c. var. TENERA: codekle ge segment, X 4.5, fron Vena 10369 (m Ps Fic. 57. N. parviroiia: A, frond, X 0.5; B, Sere n of rachis, enlarged; =a underside of segment, X 4.5; all fr rom Tryon & Tryon 5036 (m 0). Fic. 58. N. reerny a nd, X 0.5, from C. L. Smith 2080 (mo); B, underside of segment, CX re from Meyer & Rogers 2609 (mo). 98 ROLLA TRYON Pellaea nivea f. flavens (Sw.) Hieron. in Engl. Bot. Jahrb. 22: 390. 1896 Pellaca flavens (Sw.) C. Chr. Ind. Fil. 480. 1906. Pellaea nivea var. flavens (Sw.) [incorrectly attributed to Hieron. by] Hicken, Apunt. Hist. Nat. Buenos Aires 1: 117. 1909. Colombia to western Argentina; Minas Geraes, Brazil. Representative specimens: Colombia: Mutis 2102, 2104 (us). Ecua- dor: 1896, Sodiro (rp); Seemann (xk). Peru. HuAnuco: 1863, Pearce (k). Lima: Macbride 3706 (¥, us). Bolivia: White (Mulford Exped.) 619 (xk, Ny, us); Rusby 143 (ny); Weddell eee (p); Stiibel 1228 (B); Cardenas 141 (Gu). Argentina. Juyuy: Pearce (BM, K). SALTA: Lorentz & Hieronymus 209 (B, us). TuCUMAN: Vestn 1386 (GH, US), 5138 (us). A ena 1910, Castillén (cu, Pp). La Rioya: 1906, Uniche (B). Cérposa: Stuckert 11714 (pr); Hieronymus 270 (8). Brazil. Minas oars Schwacke 12764 (p); Damazio 1728 (xy, us). 57. Notholaena parvifolia Tryon, nom. no Weath. Amer. Fern Journ. 8: 107. 1918, and Journ. Arn. Arb. 34: 311. 1943 (as Pellaea microphylla) . Fic. 57. Map 66. Pellaea microphylla Mett. ex Kuhn, Linnaea 36: 86. 1869 (prior to Sept. 10); not Pellaea microphylla Fée, Crypt. Vasc. Brésil 1: 43. 1869 (after Nov. 7); not Notholaena microphylla (Sw.) Keys., 1873 which is Cheilanthes microphylla, nor Notholaena microphylla Bolle, 1858, which is Cheilanthes persica, according to C. Chr. Ind. Fil. Type: New Mexico, C. Wright 825 B!, photo cul. Rhizome compact, short-creeping; scales linear, chestnut brown, entire, thin, concolorous. Fronds ca. 10-25 cm. tall; stipe slightly grooved on the upper surface, reddish-brown, darker at base, glabrous vase bundle. Lamina quadripinnate below to bipinnate near a eltoid or ovate-deltoid, with some 5-9 pairs of alternate or satiipposite pinnae, rather abruptly contracted to the apex, upper and lower surfaces glabrous, glaucous; rachis lighter than stipe in color, grooved or channeled-flattened on the upper side, it and the pinna-rachises or slightly flexuous. Pinnae strongly patent to somewhat ascending, the basal much enlarged, the segments small, numerous, ay roundish to oblong or deltoid-oblong, petiolulate, articulate where the dark color of the stalk abruptly terminates. Veins 1-2-forked, sporangia (with 64 sod eg on the apical %4 of the vein, margin strongly revolute, unmodified. This is the only species in which the treatment of this alliance of species as Notholaena causes any serious nomenclatural con- cern. In the next species a new combination is needed but the THE AMERICAN SPECIES OF NOTHOLAENA 99 Maps 66-67. Map 66, N. parvifolia. Map 67, N. formosa. epithet remains the same. It is hoped that the choice of the new epithet will be sufficiently suggestive of the species-name under Pellaea. The distinguishing features of this species from the next, N. formosa, are discussed under the treatment of the latter. Texas and southern New Mexico, south to Tamaulipas, lime- stone cliffs and ledges and rocky calcareous slopes, up to ca. 2200 m. The literature records in Texas are from Correll, in Lundell, Flora of Texas 11, 1955 and in New Mexico from Ditt- mer, Castetter and Clark, The Ferns and Fern Allies of New Mexico (Univ. New Mex. Publ. Biol. 6), 1954. Representative specimens: United States. Texas: Waterfall 4522 (cH); Cory 885, 8403, 26675 (cu); Moore & Steyermark 3514 (cu, MO); E. J. Palmer 11554, 12991 (cu, Mo); R. M. & A. F. Tryon 5036 (Mo). New Mexico: Standley 40757 (cu); C. Wright 825 (cn, —). . COAHUILA: Hinton 16510 (cu); Stanford et al. 416 (GH, Mo); Palmer 404, 1423, 1424 (cu, mo); Johnston 7206 (cH), 8389, 8930 (cH, Mo); Stewart 271, 346, 2767 (cH). Nuevo Leon: C. H. & M. T. Muller 956, 1070 cH). Zacatecas: Stanford et al. 47 (GH, Mo). TaMautipas: Stanford et al. 882 (mo). gl? 58. Notholaena formosa (Liebm.) Tryon, comb. nov. Weath. Amer. Fern Journ. 8: 107. 1918 (as Pellaea pulche Fic. 58. Map 67. 024 Allosorus pulchellus Mart. & Gal. Mém. Acad. Sci. Brux. 15: 47, 100 ROLLA TRYON t. 10, fig. 1. 1842, not Allosorus pulchellus Presl, 1836. “type: south of Sola, Oaxaca, Mexico, Galeotti 6352 Br!. Purpus 4882a cu! is anno- tated by C. A. Weatherby as Naege cee with type at Brussels, more glaucous but in essential agreement Allosorus formosus Liebm. Vid. Selsk. Skr. s. 5, 1: 220 (Mex. Bregn., reprint 68). 1849, based on Allosorus pulchellus Mart. & Gal. Pellaea pulchella (Mart. & Gal.) Fée, Gen. Fil. 129. 1850-52. Platyloma pulchella (Mart. & Gal.) Moore, Ind. Fil. 45. 1857. Cincinalis pulchella (Mart. & Gal.) J. Sm. Ferns Brit. & For. 178. 1866 Pellaca blateneg (Liebm.) Maxon, Contrib. U.S. Nat. Herb. 24: 61. 192 Cassebeer pulchella (Mart. & Gal.) Farwell, Amer. Midl. Nat. 12: 281. Rhizome compact, short-creeping; scales linear, chestnut brown to brown, entire, thin, concolorus, oily-viscid. Fronds ca. 15-40 cm. tall; stipe terete, reddish-brown to atropurpureous, often glaucous, glabrous, or with some scales similar to those on the rhizome, definitely shorter than to longer than the lamina, with one vascular bundle. Lamina mostly tripinnate, to quadripinnate at the base, ovate to long- deltoid to ovate-lanceolate, with 6-9 subopposite to alternate pairs of pinnae, apex acute, upper and lower surfaces glabrous, usually glaucous; rachis similar to stipe. Pinnae somewhat ascending, basal largest, segments numerous, spaced, deltoid-oblong to elliptic, or narrowly so, petiolulate, articulate where the dark color of =e stalk abruptly terminates. Veins 1-2-forked, sporangia (wi spore: es) borne on the apical % of the vein, margin strongly vey unmodified. Distinguished from the related N. parvifolia, which, like N. formosa, lacks indument, by its rounded rachis, its oily-viscid rhizome scales and its inequilaterally based segments. In N. par- vifolia the rachis is grooved or channeled-flattened, the rhizome scales are dry and the segments equilateral at the base. Tamaulipas and Nuevo Leén south to Oaxaca and perhaps Chiapas, cliffs es ledges of limestone, perhaps confined to it, ca. 1500-2700 Representative specimens: Mexico. Nuevo LEOn: C. H. & M. T. Muller 971 (cH); Meyer & Rogers 2609 (mo). TaMau.tpas: Stanford et al. 882 (Gu). SAN Luts Potosi: Pringle 3401 (cu, Mo); Orcutt 5405 (Mo); Purpus 4882a (cu, Mo). Hatco: T. C. _ E. M. Frye 2558 (cH, Mo); M. T. Edwards 717 Nagi Veracruz: Spence 78 (cH); Bourgeau 2894 (cH). Oaxaca: Pringle & Conzatti 1395 (cH); C. L. Smith 2080 (mo). Cxtapas: “Chiapas etc.,” GNesbvephe 227 (cH). THE AMERICAN SPECIES OF NOTHOLAENA 101 Dusious AND ExcLupED NAMES Notholaena andromedaefolia (Kaulf.) ne pe Cyath. Hb. Bung. 29. = PELLAEA ANDROMEDAEFOLIA ( Kaulf.) F Notholaena argentea Moore & Houlst. e,. Mag. Bot. 3: 20. 1851, is prob- bl ILANTHES FARINOSA (Forsk.) Kaulf. pero He argyrostigma J. Sm. in Hook. naa Bot. 4: 50. 1841, nomen 1 Nuthohions eee per ri in Engl. Bot. Jahrb. 24: 132. 1897. Described from the island of G oupe which is not within the range of Notho- laena; it is per aps a ge eae eteans Notholaena atropurpurea or oe Polyp. Cyath. Hb. Bung. 30. 1873 = A ATROPURPUREA ink. Metkcocns candida var. aurea Pe Sp. Fil. 5: 111. 1864, nomen nudum. It is perhaps N. nivea var. flava, but I cannot place the name with cer- tainty ies oe. of the specimens cited: Ruiz & Pavon, Peru and Seemann 946, Ecu Nosunibandce redacahalee Spreng. Nova Acta 10: 227, t. 17, figs. 3-4. 1821 = CHEILANTHES MICROPHYLLA Sw. Notholaena cordata (Cav.) Keys. Polyp. ee Hb. ri 29. 1873, not (Thbg.) Desv. 1813 ; = PELLAEA SAGITTATA (Cav.) Link. Notholaena ferruginea (Kze.) D. C. Eaton, — Amer. pas d.: 28.82 201. 1860, based on SS er sores ferruginea Kze.; not Notholaena ferruginea pew: ey, He is N. trichomanoides, nor (Link) Hook. 1861, which s N. aurea = PIryROGRAMMA FERRUGINEA (Kze.) Maxon. Notholiend Filarszk yi 2 xteneanel, Mag. Bot. Lapok. 13: 38. 1914. I cannot place this name from the description although it suggests Cheilanthes . pilosa Goldm. The type is given as Warscewicz 12 from “Cord. Ecuad. & Nov. G Notholaena lanuginosa (Die: seg nos Xow noe. Cyath. Hb. Bung. 28. 1873, not (Desv.) Poir. 1816 = HES FEE1 Moore. Evidently the first valid publication of Nuttall’s hate name, Cheilanthes lanuginosa, was by Eaton in Gray’s — Addenda ci. 1863. There it — appear be a superfluous name but of the two synonyms cited, C. vestita Hook. a sensu name, not a nice aa: synonym, and C. gracilis Mett is a lite hom Notholaena lehildors (Cav.) J. Sm. in Curtis, Bot. rit 72 (s. 3, 2) Comp. = CHEILANTHES LENDIGERA (Cav Nene “lentigera (Sw.)” J. Sm. in Hook. poe. Bot. 4: 50. ie evi- dently an error for lendigera = CHEILANTHES LENDIGERA (Cav. Notholaena lutea (Desv.) Moore, Ind. Fil. 11. 1857 = aenedbetl ae aurea (Willd.) C. Chr., see Weath. Contrib. Gray Herb. 124: 14. 1939. Notholaena ec ppt (Kze.) Griseb. Abhandl. Ges. Wiss. Gott. 19: 276. 187 A Kau Notholaena Ficeophules (Sw.) Keys. Polyp. we Hb. Bung. 28. 1873, not Bolle, 1858 — CHEmANTHES MICROPHYLLA Meena micropteris (Sw.) Loe Polyp. Gyath. Hb. Bung. 28. 1873 = HEILANTHES MICROPTERIS Sw. Nithalacns mollis Hort. Gard. Chro 1893, not Kze. 1834, which is Notholaena mollis. The pe es, sn green, silvery powdered on the underside; habit dense, — e the divisi of the fronds.” is not sufficient for me to place the name Notholaena myriophylla (Desv.) J. Sm. in Hook. jeors, Bot. 4: 50. 1841, 102 ROLLA TRYON ag te on eiaeiceaaeoe a Hook. which is presumably the same esv. = MYRIOPHYLLA Desv. The combination was prop- pe ma as: by sad Palyp. Cyath. Hb. Bung. 28. Notholaena pulveracea (Presl) Kze. Linnaea 13: 135. 1839 — CHEraNTHES FARINOSA (Forsk.) Kau Notholaena rufidula Desv. Mém. Soc. Linn, Paris 6: 221. 1827, is probably ENSIS Dr: sintcorcio scariosa (Se. ) Baker, in Mart. Fl. Bras. 17: 540. 1870 = CuHeEr- LANTHES SCARIOSA (Sw.) Presl, see Weath. Contrib. Gray Herb. 124: 19. 1939. Notholaena squamata Moore & Houlst. Gard. Mag. Bot. 3: 20, 1851. I can- not place this name from the description. Said to be from Mexico and Peru, the former eon babe prereset place it with N. brachypus, the latter country perh N. peruv ~ teria (Cae) "Keys. neh hg “Gyatk. Hb. Bung. 30. 1873 = ERNIFOLIA (Cav. in Wotholedan tomentosa " Link) Keys. Polyp. Cyath. Hb. Bung. 28. 1873, not ee sv. 3, which is Notholaena tomentosa = CHEILANTHES TOMENTOSA Natholaen vestita (Spreng.) Desv. eur oem Appl. 1: 93. 1813 = CHEI- NTHES LANOSA (Michx.) D. C. INDEX TO NAMES Acrostichum albidulum 95 onariense 35 vens 96 Marantae 7 tereticaulon 96 Aleuritopteris 7 affinis candida 66 cretacea 69 ~ gulphurea 69 Allosorus formosus 100 furfuraceus 66 Cassebeera pulchella 100 il tabi Ponte 66 Cheilant affinis Aah Sinise 54 Balansae 34 cretacea 69 Davenportii 76 dealbata 87 deltoidea 21 emmonii 61 lepida 6 sis ylla 98 li Schaffneri 57 sinuata 21 squamosa 18 92 Seitbate 87 Fendleri 89 vens 96 nivea 95 sinuata 21 var. integra 21 104 es 6, 15 (Map 1) affini 46 (Fig. 23), 47, 48 la 9 _ 7 “(Map 34), 59 (Fig. 31), ait cladachots arequipensis 15 (Map 3), 16, 17 (Fig. 2) argentea 101 dle 101 Arse ecabeeae 53 Oise 30), 54, 55 (Fig. 27), 56 asplenioides ve atropurpurea 1 aurantiaca 71 (Map 42), 73, 75 Fig aurea 33 (Fig. 15), 35, 36, 38 (Map 18) Balansae 34 bipinnata 54 bonariensis a brachypoda 7 —— 25 ‘ig 6), 26, 28 Map 9 Brackenridgei bir taser 75 (Fe. 43), 77, 80 (Map 46) Buchtienii 40 var. Buchtienii 38 (Map 21), 41 (Fig. 18), 42 var. ventanensis 38 (Map 21), 42 californica ie (Map 43), 73, 40) ssp. nigrescens — 65, 66, ey “(Fie 36), var. accessita 73 var. aurea 101 var. candida 64 (Map 39), 66, 67 (Fig. 36a) var. Copelandii 64 (Map INDEX ) Sif ee 36b), 68 var Nedcte pe 70 pie es 27 chalcolepis 21 oe 101 chiapensi chilensis i, *6 (Map 50), 81, 82 (Fig. 47) chrysophylla 96 cinnamomea a (Map 16), 33 (Fig. 13), 8 cochisensis igs a cubensis prs (Map 27), 49, 50 (Fig. gers 85 (Fig. 52), 87, 88 57 delicattta 88 (Map 88), 90, 91 (Fig. 54) deltoidea a Doradilla 4 re 6. (Fig. 22), 47, 48 eiphora 2 28, 29 (Fig. 9), 31 (Map 12 Fendleri 85 (Fig. 53), 88 (Map 58); 89 ferruginea 35, 51, 101 var. Canescens 35 oe 101 flavens 96 eel 97 (Fig. 58), 99 (Map 67 Siege mA (Map 20), 39, 41 (Fig. var. ae galapagensis rs ( Fig. 35), 64 (Map 38), 6 Galeotti = (Map 31), 54, 55 (Fig Ph Be 29 (Fig. 11), 31 (Map 14) INDEX Gilliesii 18 ——_ 29 (Fig. 10), 30, 31 Map 13 Grayi 53 Ng 33), 58, 59 (Fig. 3 Greggii 75 (Fig. 42), 76, 80 45 Hassleri 25 (Fig. 7), 7, 28 (Map 10) Herzogii 34 Hookeri 70, 81 hyalina 56 hypoleuca 39, 58 incana 61, ne = (Map 60), 91 (Fig. 5 Jonesii 208 (Q ae A 82 (Fig. 9 64 var. australis 62, 64 (Map 35) var. Lemmonii 61, 63 (Fig. 32), 64 (M ap 35) 64 (Map etl: 69 Lilloi 18 eve 85 (Fig. 51), 86, 88 (Map 54) var. mere 8 (Fig. 51a), 86, 88 (Map 55) ssp. inesicane 87 var. mexicana S (Fig. 51b), 87, 88 (Ma lonchophylla 17 (Fig 4), we Lumholtzii 2s (Map 53), 85 (Fig. 50) lutea 101 Matthewsii 101 microphylla hi 101 micropte pteris mollis 44 (Fig. 21), 45, 48 (Map 24) mollis Fol monosticha 66 myriophylla 101 Nealleyi 53, 55, 57 var. mexicana 57 neglecta 71 (Map 44), 74, 75 Fig. 4 ( Newnan 43, 44 (Fig. 20) nivea 92, 93, 94 lag 61), 97 (Fig. 56 var. dealbata 87 var. flava 94 oe 64), 96 var. mexicana 8 var. oe A (Map 62), 95, 97 (Fig. 56a) var. eae ed 94 ‘(Map 63), 95, 97 (Fig. 56b) var. fase 94 (Map 64), 96, 97 (Fig. obducta = hie 17), 33 (Fig. 14),3 pallens ‘ (Fig. 45), 80 (Map 48 palmatifida 28 Palmeri 77, Soe (Fig. 44), 79, 80 (Map 47) Parryi ie (Map 22), 48, 44 (Fig. sai rh 97 (Fig. 57), 98, 99 (Map 66), 100 peninsularis 80 hae 2 51), 82 (Fig. 48), ap stg 15 (Map 2), 16, 17 (Fig. 1-4 pilifera 79, 80 (Map 49), 82 (Fi tid 31 (Map 15), 32, 33 (Fig. 12) 1 pulchella ee pulveracea 1 ae? = (Mep 28), 49, 50 (Fig. 25) 106 Rosei 62, 63 (Fig. 33), 64 (Fig. 36) rufa 35 var. major 35 var. minor 35 rufidula ie scariosa 19, sro gat 3. (Map 32), 55, var. mexicana var. Nealleyi 55 ig 29b) 57 var. Schaffmeri 55 (Fig. 29a), 57 sinuata 19, 22 (Map 5) f. crenata 24 f. pinnatifida 21 var. bipinnata 15 var. Tine ss 22 (Map 8) 24, 25 (Fig. 5c) var. ‘bids 24 var. integerrima 22 (Map 7), , 25 (Fig. 5b) var. integra 21 var. laevis 21 var. pinnatifida 21 ar. pruinosa 21 var. sinuata 21, 22 (Map 5 in its sie Ar 25 (Fig. 5a) squam ei iconhor re (Map 4), 17 (Fig. iS < Standleyi, 67 (Fig. 38), 70, 71 (Map 4 ys stellapilis sulphurea er ie 37), 69, 70, 71 (Map 40) var. accessita 73 var. alba 66 var. Borsigiana 69 var. a 73 a 69 var. quin nausea 72 Tectaria 21 tenera 96 var. major 96 ternifolia 102 INDEX tomentosa 38 (Map 19), 39, 41 Fig. 16 tomentosa 58, 102 trichomanoides 50 (Fig. 26), 51, 53 (Map 29) venta o7, 28 Otis 11), 29 . 8) oeulle 102 Weatherbiana 60 Pellaea 7 candida 92 chilensis 81 dealbata 87 var. limitanea 86 var. Stubeliana 87 Fendleri 89 ferruginea 35 flavens 98 formosa 100 Greggii 76 Jonesii 83, 84 Lilloi 18 limitanea 86 var. mexicana 87 microphylla 98 nivea 95 f. nivea 95 f. flavens 98 f. tenera 96 var. flavens peruviana 96 Ichella 99, 100 Stubeliana 87 tenera 96 Platyloma pulchella 100 Polypodium a 28 Pteris aurea 35, Trichomanes nivea 51 — 2 Coe ie eee ead CONTRIBUTIONS FROM THE GRAY HERBARIUM OF HARVARD UNIVERSITY Edited by Reed C. Rollins and Robert C. Foster NO. CLXXX STUDIES IN SIDA (MALVACEAE) I. A Review of the Genus and Monograph of the Sections Malacroideae, Physalodes, Pseudomalvastrum, Incanifolia, Oligandrae, Pseudonapaea, Hookeria and Steninda. By I. D. CLEMENT PUBLISHED BY THE GRAY HERBARIUM OF HARVARD UNIVERSITY CAMBRIDGE, MASS., U.S.A. 1957 ISSUED APRIL 8th Ane, pasate - INTRODUCTION This paper is the first in a series of detailed systematic studies of the genus Sida. It reviews the previous work on the genus and the changing concepts of its limits and evaluates the twenty section names which have been published. Of the nearly twelve hundred published specific and subspecific names, 112 are here individually assigned to five of the previously established sections or to the three new ones proposed here, and these, involving 28 species and 9 varieties, are monographed. The remainder are reserved for consideration in later papers in this series. The genus is one of the warm-temperate and tropical zones, with the primary center in the New World tropics and a second- ary center in. Australia. About one hundred and twenty species are confined to the New World and about thirty-five to Australia. One species is found only in Europe and Asia Minor; there are several endemic species in Africa, and a few others are found in Asia and the Pacific Islands. Eleven species of weedy behavior are shared by the New World and the Old. Of these, S$. rhom- bifolia, S. spinosa and S. acuta are aggressive weeds found nearly throughout the range of the genus. S. leprosa of the New World has the greatest, although a discontinuous, latitudinal range, from nearly 48° north to nearly 48° south. SuRVEY OF PREVIOUS STUDIES Linnaeus originally described 10 species in Sida, which he based on Malvinda Dill. and Abutilon of Tournefort and of Dillenius. Of these, only the first four remain in Sida, S. spinosa, S. rhombifolia, §. alnifolia and S. cordifolia; two species are now in Wissadula, and one each in Abutilon, Bogenhardia, Anoda and Malachra. The genus as he defined it would include the presently accepted genera of subtribes Abutilinae and Sidinae of tribe Malveae, and Malachra of tribe Urenae, which he separated from Sida in 1767. Adanson (1763) proposed the substitution of the name Abutilon for Sida, apparently because it was the classically correct one. He cited Malvinda Dill., Abutilon Dill., and Sida L. as synonyms of Abutilon, which was thus superfluous when published. Miller (1754) had already used Abutilon as a generic name, and is 6 STUDIES IN SIDA (MALVACEAE) sometimes cited as the author of the genus In the first work in which he consistently used the binomial system (1768), under the heading “‘Abutilon’’, he merely said “see Sida’. Under “Sida”, he indicated that he thought Abutilon a separate genus and gave the distinguishing characters. He did not list nor describe any species in Abutilon, and one of the species of Sida he described is now considered an Abutilon. I have considerable doubts that this publication of Abutilon in the abridged 4th edition of his Dictionary (1754) should be considered as valid, but if it is acceptable, and there is nothing in the present Rules specifically against it, then the problem created by the Adanson use of the name, invalidating it for use in its presently accepted context, is avoided. Cavanilles (1785) began his extensive writings on the Linnaean Monadelphia with a monographic dissertation on Sida, which he added to and corrected in subsequent publications. He described Anoda (1785) and Cristaria (1799) as segregate genera, but his concept of the genus was still wide enough to include Abutilon and other pluriovulate genera. In his various publications, he attributed a total of 109 specific names to Sida. Medicus (1787), whose attitude toward Linnaeus’ work is summarized by his comment on the Genera Plantarum (p. 147), “der irrigste Wegweiser.... den ich kenne”, divided Sida into the pre-Linnaean genera on which it was based, Malvinda and Abutilon, and reserved Sida for species not now considered as belonging in the genus. He also established the new genus Wissa- dula, which was represented in Sida of Linnaeus by S. periploci- folia. In 1789, he described a new genus, Lamarkia, for the taxon now called S. humilis Cav. His use of Sida, Malvinda and Abutilon was followed by Gaertner (1791) and Moench (1794), but by few others, as the authoritative stature of the work of Linnaeus increased. Kunth (1822) maintained Abutilon as a separate genus and is often credited with being its author. In addition, he described two new genera, Gaya and Bastardia, for species formerly in Sida or new in this work. De Candolle (1824) in the Prodromus made the species of Gaya and Bastardia the constituents of a new section, Abutiloides, and reduced Abutilon Kunth to a section. Rather inconsistently, he described Periptera as a new genus separated from Sida. Only a STUDIES IN SIDA (MALVACEAE) rs few subsequent authors followed de Candolle in regard to Gaya and Bastardia, although many continued to consider Abutilon as a section of Sida containing the pluriovulate species. De Candolle’s third section, Malvinda, based on Medicus’ genus of that name, was almost exclusively composed of species which are still retained in Sida. This section has undergone progressive limitation since its founding and is narrowed still further here. Despite this, it remains by far the largest section of the genus, with about a hundred species, and is the one which contains the typical element, S. rhombifolia L., and most of the common and weedy species. Under the present Rules, it must be called section Sida. Sprengel (1826) treated 176 species, returning or newly trans- ferring to Sida a number of species of Anoda, Bastardia, Cristaria and Gaya. G. Don (1831) distributed 180 species into three sections: Malvinda DC., Abutiloides DC. and his new Malacroi- deae, the last of which I have accepted here. Rafinesque (1836), declaring that certain species, still kept in Sida, should be consid- ered distinct, proposed Diadesma as the name of a new genus to accommodate them, without ever describing it. Wight (1837) described a new genus, Dictyocarpus, for Sida cuneifolia Roxb., which has won no acceptance. The confused nomenclatorial history of this species, which includes its misidentification as a species of Riedleia and thus to the occasional citation of that genus of the Sterculiaceae as a synonym of Sida, is discussed under Sida Grayana (S. cuneifolia Gray). Steudel proposed a new monotypic genus, Fleischeria, for a species soon identified with and here called S. calyxhymenia J. Gay. D. Dietrich (1847) arranged 358 species in the following groups whose category was not specified: Malvinda, Abutiloides, Mala- croideae, Abutilon, Cristaria, Anoda and one unnamed. He restored its Linnaean extension to Sida, including in it everything then known in the genera Sida, Gaya, Bastardia, Abutilon, Perip- tera and Cristaria. The next subdivisions of the genus were made by Gray (1849) when he published the section Pseudonapaea for two species now known as S. hermaphrodita (L.) Rusby and Abutilon Hulsea- num (Gray) Torr. & Gray. In the same paper Gray established the section Psexdomalvastrum for the two involucellate species S. hederacea and S. sulphurea, here considered as synonyms 0 S. leprosa. He and Baker separately added other species, two of 8 STUDIES IN SIDA (MALVACEAE) which, S. cuneifolia Gray and S. Helleri Rose, are here considered to belong in a new section. In the Plantae Wrightianae (1852), Gray described a third section, Physalodes, for certain species with inflated fruiting calyces, including his own S. physocalyx. The aberrant Sida Sherardiana was made the basis of a new genus Malvella by Jaubert & Spach (1855) but its separation from Sida is now maintained by very few botanists. I have included it in section Psendomalvastrum. Grisebach, in his Flora of the British West Indian Islands (1859), divided the genus into three sections, Malvinda, Steninda and Wissada, without indicating that de Candolle was the author of the first. This led Schumann and, later, Baker to cite Grisebach as the author of that section. Section Steninda was established with only one species, S. linifolia. Although Baker (1892) greatly modified the section-concept, the section is restored here to Grise- bach’s definition of it and includes only S. linifolia and _ its immediate allies. Section Wissada was established to accommodate S. divergens Benth., which is usually placed as an anomalous element in Wissadula. Baker (1892) transferred the section-name with the species to Wissadula and added a new species, W. Balansae, the latter now considered synonymous with Briquetia denudata (Nees & Mart.) Chod. & Hass., based on Sida denudata Nees & Mart. Miquel (1859) used the conventional sign for section in con- junction with four Latin adjectival plural names, each followed by a brief phrase descriptive of the leaves, to divide the species of Sida in his Flora van Nederlandsch Indié. Since he did not name the category of these divisions and obviously did not intend to describe formal sections, I have not listed these. All of the species in his work are usually placed in section Sida. In 1860-62, Mueller in his Plants Indigenous to the Colony of Victoria described three new sections, Eusida, Notho-Plagian- thus and Lawrencia. Section Eusida was intended for ‘typical’ species of the genus, as opposed to those of the other two sections, but as specified it included only certain aberrant Australian species which are not in section Sida as I understand that section. The species of section Notho-Plagianthus, together with the syno- nyms given, are now treated as Plagianthus pulchellus Gray, thus removing the sectional name from the genus. The section Law- rencia is now ireated as a section of Plagianthus. In this same STUDIES IN SIDA (MALVACEAE) 9 work, Mueller included a fourth and last section, Abutilon, with two species, both now referred to the genus Abutilon. Grisebach in his Plantae Lorentzianae (1874) added a fourth section Sidastrum, to the three he previously recognized and in this he included several species usually referred to Malvastrum; none of these is here considered as belonging in Sida. Baker (1892) used the name Sidastrum for a monotypic genus based on S. quinquenervia Duchass. which has no relation to this section and which Kearney (1954) and others have retained in Sida. In 1887 Gray published two new sectional names, Abutilastrum and Calyxhymenia. The first was a nomen nudum, Gray merely indicating that it was appropriate for a new section which included §. Lindeniana, now referred to Robinsonella, while the second was superfluously published for his own section Physalodes (1852), since it included the same group of species. A certain amount of confusion in the nomenclature of the sections was created by Schumann. In his treatment of the Mal- vaceae in Engler and Prantl (1890) he listed the names, without descriptions, of 5 sections of Sida (as he gave them): Pseudo- malachra K. Schum., Physalodes A. Gray, Steninda Griseb., Malvinda Griseb. and Pseudomalvastrum K. Schum. The first was a nomen nudum, the fourth should have been attributed to de Candolle and the last to A. Gray. In 1891, in the Flora Brasiliensis, he listed and described seven sections, adding to the previous list two new ones, Bastardiopsis K. Schum. and Thyr- sinda K. Schum. He still attributed Malvinda to Grisebach and Pseudomalvastrum to himself and changed the attribution of Physalodes, wrongly, from Gray to Grisebach. His section Pseu- domalachra, based on the same taxa which Don had made the basis for his section Malacroideae, was superfluous when pub- lished. He also published here a new section-name without de- scription, in his observations on S. myriantha, using the following words, “Hae tres species, nempe S. myriantha Pl. & Lind., S. Lindeniana Turcz. et indescripta ex Insula Tabago probabiliter sectionem bene distinctam efformant, cui apte nomen Dendrinda attribuendum est”. Thus another nomen nudum in addition to Gray’s Abutilastrum was published for a section including S. Lindeniana. The last survey of the whole genus was that by E. G. Baker (1892). He indexed a total of 634 specific and subspecific names, 10 STUDIES IN SIDA (MALVACEAE) excluded 340 of them and distributed the rest in 108 species divided among 6 sections, or listed them as “dub. vel. ignot.” or “incert. sect.”. Descriptions were given only for the sections, and the synoptic key in the text was only to groups of species. This synopsis has been the basis for most of the recent work in the genus and even his nomenclature has been perpetuated, despite changes made necessary by the changes in the Rules. The six sections which he recognized were, as he gave them: Pseudomalvastrum Gray (4 spp.), Abutilastrum Gray (5 spp.), Calyxhymenia Gray (10 spp.), Steninda Griseb. (12 spp.), Pseu- domalachra K. Schum. (2 spp.) and Malvinda Griseb. (75 spp.). He took up Gray’s name Abutilastrum for a section including S. Lindeniana, S$. myriantha, S. lepida of Australia, S. Eggersti (with reservations as to its place in the section, since the carpels were very little inflated) and S. densiflora. As was his custom in this work, he provided a Latin diagnosis of the section, thus validating Gray’s name. Since he included S. densiflora, which had been made the basis for the monotypic section Bastardiopsis by Schumann the previous year, section Abutilastrum was super- fluous when published. Section Bastardiopsis was later trans- ferred to the genus Bastardia and then raised to generic rank by Hassler (1910). Section Calyxhymenia Gray, as previously noted, was superfluous for section Physalodes Gray, and section Pseu- domalachra K. Schum. for section Malacroideae G. Don. Section Malvinda should have been attributed to de Candolle. Baker, as already mentioned, greatly altered the original concept of Steninda and placed in it a number of Australian species which have nothing in common with the typical element, S. linifolia, save terete or subterete calyces. He did not accept Schumann’s section Thyrsinda, returning the two species Schumann included to section, Malvinda. The list of sections in Dalla Torre and Harms follows Schumann in incorrectly attributing section Mal- vinda to Grisebach and Pseudomalvastrum to Schumann and uses Calyxhymenia A. Gray instead of the earlier Physalodes A. Gray. Greene (1906) separated several species, here considered as be- longing in sections Psexdomalvastrum or Incanifolia, as the ele- ments of a new genus, Disella. For a time, some botanists of the western United States followed Greene in this, but few if any do so now. STUDIES IN SIDA (MALVACEAE) 11 In addition to those studies of the genus in several geographic areas which have already been mentioned because of their im- portance in the delimitation of the genus and its sections, certain others are of interest. The first of these was St. Hilaire and Naudin’s survey (1842) of the species then known from Brasil, 11 of them new. This was followed by Schumann’s work in Martius’ Flora Brasiliensis (1891) already referred to, and by Monteiro’s preliminary survey (1935) consisting mainly of keys, which is the only published fascicle of a proposed monograph of the Brasilian Malvaceae. Baker (1937) reviewed Monteiro’s paper, calling attention to some of the errors an isolated worker inevitably commits when he does not have access to types, to material from outside his immediate area, nor to a good library. Monteiro later (1949) published an account of the Brasilian, Uruguayan and Argentine species of section Sida, in which he provided keys to a series of 4 named but not formally established subsections and to the 48 species of these which he recognized from the area. He depended to a great extent on Rodrigo’s (1944) paper for the species found in Argentina and Uruguay. Although his paper shows some of the same inadequacies, it is considerably more accurate and useful than his earlier one. Rodrigo’s paper also shows the effects of a lack of references and authentic material. In attempting to make up the latter deficiency, she assembled photographs of “types” of all the species treated. These are excellently reproduced in her paper, but un- fortunately some of them do not illustrate the actual types. The plate illustrating the “type” of S. ciliaris L. actually shows the type of S. muricata Cav. in the Madrid herbarium; that of S. linifolia Cav. shows a Boldo collection from Cuba, whereas the type was collected by A. de Jussieu; and that of S. prostrata Cav. shows a Née collection purportedly from Chile, whereas the type is a Commerson collection from Uruguay The first treatment covering all the truly North American species of the genus was that in Gray’ s Synoptical Flora (1897), which considers 21 species in 4 sections. Since then, Small has added 2 and Fernald 1 new species to those known from the United States, all 3 of which are in close relationship to the complex centering on S. Elliottii Torr. & Gray of section Sida. The Mexican species were treated in the 2 editions of Sessé and 12 | STUDIES IN SIDA (MALVACEAE) Mocifio’s Flora Mexicana (1889, 1894) and by Hemsley (1879), but the first organized treatment was that of Standley (1924). Hemsley added only a varietal nomen nudum, while the 2 editions of Sessé & Mociiio’s work contained a number of new names and overly brief descriptions. Baker, who made a special trip to Madrid to examine the types of certain Cavanilles’ species, men- tioned in a brief article on his trip the availability of the Sessé and Mociiio specimens there but completely ignored many of their species in his Synopsis. Standley keyed out 28 species and gave summaries of their characters, bibliography and designation of type-specimens. Kearney (1954) has published a tentative key, with brief notes and partial synonymy, for 55 North American, Mexican, Central American and West Indian species. The paper contains very few of the defects which might reasonably be expected in a geographi- cally restricted treatment of a large, variable and wide-ranging genus and is a much-needed and valuable contribution. The species of Australia were last summarized in Bentham & Mueller’s Flora Australiensis (1863). The many species described and the changes made in the classification of others since then are only partly covered by regional floras. The species are mostly endemics, only 6 of them occurring outside Australia. For other parts of the Old World, only conventional treatments in floristic works are available, with the exception of Gagnepain’s (1909) brief and inadequate “Essai d’une classification des Sida asiati- ques” in which he keys a mere 10 species and confesses his inability to separate S$. rhombifolia from S. spinosa. Hu (1955) in her study of the Malvaceae of China has published a number of new combinations and taxa in Sida, all of them in section Sida. Special studies of the morphology, anatomy and cytology of the genus are almost completely lacking. Hochreutiner has pub- lished some observations, his (1920) paper on carpellary structures in the family being of special interest. Kearney (1951) in his useful survey of the American genera of Malvaceae has provided an excellent summary and discussion of the taxonomically useful characters of these genera, with a fairly complete bibliography. Metcalfe and Chalk (1950) summarize the slight amount of in- formation on the anatomy of the genus, while the literature of economic botany has innumerable references, particularly to the use of a number of species as locally important fiber plants. STUDIES IN SIDA (MALVACEAE) 13 ACKNOWLEDGEMENTS Studies preliminary to this paper were begun in 1946 under the late Prof. M. L. Fernald and completed as a doctoral thesis in 1947—1948 under Prof. Lincoln Constance while he was Acting Director of the Gray Herbarium following Prof. Fernald’s retirement. During that period, I received considerable assistance, which I here gratefully ack- nowledge, not only from Professor Constance but also from members of the staff of the herbarium, particularly Drs. Lyman Smith, Bernice Schubert and Robert Foster. Dr. Schubert rendered me a great service by making critical notes and photographs of certain Miller, Baker and Linnaeus types while studying types of American species represented in various British herbaria. This paper was completed at the United States National Herbarium in 1956 while on sabbatical leave from the Atkins Garden and Research Laboratory. I wish to thank the Head Curator, Mr. Jason Swallen, for permitting me to work there and to acknowledge once again the helpful assistance of Dr. Lyman Smith, now of that herbarium. The Directors and Curators of the following herbaria have generously permitted me to borrow from the collections in their care (the herbaria designated in the text by the symbols given here): Chicago Natural History Museum (formely Field Museum) (F); Dudley Her- barium, Stanford University (DS); Gray Herbarium, Harvard University (GH); Missouri Botanical Garden, St. Louis (MO); Muséum d'Histoire gag Paris (P); New York Botanic Garden (NY); Pomona College (POM); Rocky Mt. Her- barium, University of Wyoming (WYO); Royal Botanic Garden, “i England (K); Southern Methodist University, Dallas (SMU); United States National Her- barium (US); pacar of Arizona (ARIZ); ee of California (UC); Uni- pares of Mexico, Herbario Nacional (MEXU); University of New Mexico (UNM); University of Utah (UAC); Vegetation Type Map Herbarium, at Uni- versity of California Other citations, peiindesly: to indicate the location of type, use the symbols of Index Herbariorum (ed. 3) TAXONOMIC TREATMENT Sipa L, Species Plantarum ed. 1, 2: 683-686. 1753. Abutilon Adans., Fam. 2: 283. 1763. Malvinda Medic., Malv. 23. 1787. Lamarkia Medic., Phil. Bot. 1: 28. 1789. Diadesma Raf., New Fl. Am. 1: 41. 1836. (nom. nud.). Dictyocarpus Wight, Madras Jour. Sc. 5: 309. 1837. Fleischeria Steudel in Lehmann, Pl. Preiss. 1: 236. 1845, non Steudel and Hochstetter 1838. 14 STUDIES IN SIDA (MALVACEAE) Malvella Jaub. & Spach, Illustr. Pl. Or. 5: 47 t. 446. 1855. Sidastrum E. G. Baker, Brit. Journ. Bot. 30: 137-8. 1892. Disella Greene, Lflts. Bot. Obs. & Crit. 1: 209. 1906. Herbs, shrubs or small trees, annual to perennial. Stems fibrous to woody, pubescent or glabrous, simple or freely branching. Stipules pre- sent, free, sometimes deciduous. Leaves alternate, distichous or spiralled, from simple-linear and entire to palmately lobate or orbicular-reniform and serrate. Inflorescence solitary and axillary to paniculate and termi- nal; individual flowers never large and showy. Epicalyx none or, in a few species, of 1—5 bracteoles at the base of the calyx (a false involucel of 6—8 linear bracts on the upper pedicel in one species). Calyx valvate, 5-lobed, terete or angled, campanulate or urceolate, in some species greatly accrescent after anthesis. Petals five, adnate at the base to the lower part of the stamen-tube. Stamens fused in a column, the free portion of the filaments terminal or some of them subterminal, the anthers 5 to 40 or more, unilocular, usually everted in anthesis. Ovary superior, enclosed by the stamen-tube and the fused bases of the petals. Styles free at least in the upper part, isomerous with the carpels. Stig- mas terminal, capitellate or capitate, sometimes oblique, never decur- rent. Mature fruit a single whorl of schizocarps usually separating freely from the axis and each other, in an open, closed or greatly ex- panded calyx. Schizocarps uniovulate, indehiscent or dehiscent along the mid-dorsal line, rarely along the inner margin or by breakdown of the lateral or basal walls; apex muticous or variously rostrate, two-beaked or biaristate (in some specimens of S. rhombifolia a single arista by fusion), the aristae sometimes subapical; apical wings absent; basal hooks absent; lateral walls persistent, membranous, reticulate, rugose or clathrate; endoglossum absent. Seed rounded, somewhat cordiform, with- out an arilloid envelope, separating freely in maturity or adherent to the lateral walls of the carpel, pendulous from the upper inner angle, the funicle, raphe and radicle thus superior. Columella (receptacle) trun- cate, usually slender and slightly conical, the base occasionally flared; remnant vascular traces from the base to the individual carpels persisting in a few species. Cotyledons variously plicate or contortuplicate. The name used by pre-Linnaean authors in many applications, by Theophrastus for a Nymphaea, and by other classical authors supposedly for some “marsh-mallow” or water plant; from the Greek 2167. Type species: Sida rhombifolia L. Britton and Wilson (1924) named S. alnifolia L. as the type species without giving any reasons for their choice. Hitchcock and Green (1935) named S. rhombifolia L. Their list of Linnaean STUDIES IN SIDA (MALVACEAE) 15 generic lectotypes having been published as a supplement to the third edition of the International Rules led Rodrigo (1944) to assume that it carried the authority of the Rules, and she cited these as the authority for her designation of S. rhombifolia. Hu (1955) also designated S. rhombifolia. Neither S. spinosa nor S. cordifolia has been selected; they are the first and last of the four species originally named by Linnaeus which are still con- sidered to be in Sida. Both are represented by at least one specimen in the Linnaean Herbarium, according to Savage (1945 Although there is some question of the reliability of Jackson’s (1912) dating of the presence of the various specimens in ‘this herbarium, in all probability the single specimen representing S. rhombifolia was there before the publication of the Species Plantarum ed. 1. The species is well known and widely distrib- uted, nearly throughout the natural range of the genus and beyond it as a weed or waif. It is in every respect a typical species of the genus, and for these several reasons, I have accepted Hitchcock and Green’s selection of it as the generic type. THE SECTIONS The following sections of the genus have been named: Malvinda (Medic.) DC., Prod. 1: 459. 1824. (The typical section, Sida). Abutiloides DC., op. cit. 467 = Gaya & Bastardia spp. Abutilon (Kunth) DC., 1. c. = Abutilon spp. Malacroideae G. Don, Gen. Syst. 1: 498. 1831. Pseudomalvastrum A. Gray, Mem. Am. Acad. 4: 23. 1849. Pseudonapaea A. Gray, l.c. Physa- lodes A. Gray, Smithson. Contr. Knowl. 3, art. 5: 20. 1852. Steninda Gris., Fl. Brit. W. Ind. 76. 1859. Wissada Gris., op. cit. 77 = Wissadula sp. Notho-Plagianthus F. Muell., Pl. Vict. 160. 1860-62 = Plagianthus sp. Lawrencia F. Muell., op. cit. 162 = Plagianthus, sect. Lawrencia. Eusida F. Muell., op. cit. 163. Sidastrum Gris., Pl. Lorentz. 42. 1874 = Malvastrum spp. Calyxhymenia A. Gray, Proc. Am. Acad. 22: 294. 1887 = sect. Physalodes. Abutilastrum A. Gray, op. cit. 295 (nom. nud.). Dendrinda K. Sch., Mart. Fl. Bras. 12 (3): 347. 1891. (nom. nud.). Bastardiopsis K. Sch., op. cit. 280 = Bastardiopsis sp. Pseudo- malachra K. Sch., |. c. = sect. Malacroideae. Thyrsinda K, Sch., op. cit. 281. Abanilastoem A. Gray, valid. Bak. f., Jour. Bot. 30: 38. 1892. 16 STUDIES IN SIDA (MALVACEAE) Those which I am accepting are in bold-face. Reasons for the rejection of the others are either summarized by reference above to present disposition of the contained species or were given in the survey part of this paper. In addition to those accepted, three new sections are proposed here for species formerly in section Pseudomalvastrum or section Sida. KEY TO THE SECTIONS > Pedicels of the inflorescence adnate to the petioles of foliaceous bracts; florescence often arrcrecmrc aE at a apices of padchll; calyx terete, not accrescent . . Malacroideae (p. ie Pedicels of the inflorescence re Calyx inflated, sear anous, greatly accrescent steal anthesis, stipulate: gif cate; carpels mutic : raghsig 3B Page (p. 30) yx not or only sLighOY i accrescent seas baiics Ses Epicalys of 1—5 narrow bracteoles; — muticous, »-glightl - inflaved: in- t S Ill. Pseudoz o> quingquenervia) D Leaves suborbicular, to 14 mm. long, a stellate-velutinous; carpels muticous, the apex obtuse IV. Incanifolia (p. 60) Leaves linear to cordate or palmately I lobed, or a small-suborbicular, then not velutin E Leaves sible lo be -d Carpels two-beaked or bideiecate, hie 4 aristae peenodeely haibed: anthacs 5—20 Oligandrae (p. 64) Carpels muticous, the rostrum neatly horhabeal or prolonged upwards, nev: G Shorwees subumbellate-conymboe, rostrum of ‘the carpels ascendent: ‘anthers numerous VI. Pseudonapaea 5 Flowers axillary. Soles or peised ‘rayely teversil from solitary or several peduncles; rostrum of the 5 a Be ge horizontal; anthers +—20 VII. Hookeria (p. ay oo: FS ise! ao} _ ao z J * ~ om 7 oO =] ct oe NA fo eh 5-8) a mr 5 < = = a 2. ° aad ie.) > ad pP ia) co n ™ tri 7 teases simple, coldaee: or ebeprdat Leaves entire, linear to elliptic; tive culsionbies: carps ‘bluntly diel: inflorescence corymbose . : VIII. Sestate (p. 82) Leaves variously dentate, at lee apically. . . . Thyrsinda, Sida et al. I ta I. Section MatacrorEaE G. Don, Gen. Syst. 1: 498. 1831. Section Tsend Omang K. Schumann, Engl. & Prantl, Nat. Pfl. 3, Abt. 6: 890, Subshrubs or pa Pedicels adnate to the petioles of foliaceous bracts, flowers often subumbellate-congested at the apices of the STUDIES IN SIDA (MALVACEAE) 17 branches. Calyx not accrescent, epicalyx absent. Carpels often stellate-pubescent, strongly muricate to nearly smooth, dehiscent mostly along the inner margin. Don established this as the first new section since those of the Prodromus, with the brief characterization “heads of the flowers involucrated”, and remarked that “this section of Sida agrees with Malachra as it is now constituted. There is much difference of opinion about the character of that genus”. He included the elements represented by the names S. fulva, S. anomala, and S. plumosa, and these three elements are also represented in K. Schumann’s section Pseudomalachra. It might seem that the very brief diagnostic phrase given by Don was insufficient, yet this section contains the only species of the genus in which the pedicels are regularly adnate to the petioles of foliaceous bracts which, with their stipules, give the appearance of involucres. Schumann’s characterization of the section in Mart. Fl. Bras. (1891) is more exact and is the basis for the one used here. Don’s name has been nearly completely ignored and Schumann’s used, particularly in recent works. Type species: Sida ciliaris L. KEY TO THE SPECIES > Stem-leaves long-hirsute on both surfaces; dorsum of the carpels densely puberulent, the outer margin a 3-peaked ridge; petals 12—13 mm. long; anthers 15—20; stamen-tube with pe ine he strigose a plants pro- 1. S. Brittonit. > cumbent Stem-leaves slabrcud or i aed pubescent on ibe upper ae etals 18—22 mm. long, 10—14 mm. wide; anthers 30—40; dorsum of the carpels Ce verrucate and rugose; plants — eae ee . 8. centuriata. Petals 1; mm. tong, 2 —11 mm. wide: anthers 10—30 Cc Stamen-tube heavily patent-stellate-pilose; petals 13—-16 mm. long, ca. 5 mm. wide; carpels 7—8, verruculate, submuricate; plants procumbent 3. S. surumuensis. A ow Stamen-tube Giahious to > anes pilose : mm. lo —11 mm. aide stem- ae caneite-oblons, rarely more than 1 long, crowded and clustered; anthers ca. 20; carpels 5; plants emidecumbent 4. S. paradoxa. on ro g “a _ os oe) 7 fed 3 a BY ae wn 3} ee . “ao, 5 a. tone = co ° ce — B 5. 2: ae aL) 4 oO 49 ae £2 w wn c o o igs bicular to elliptic to ‘oie not crowded aad eae’: shiek 5—8 muticous to short-aristate, the dorsum “ee eesiape often mur- icate; plants prostrate to erect . eee ies SES 18 STUDIES IN SIDA (MALVACEAE) I. Sida Brittonii Leédn, Torreya 19: 172. 1919. Perennial prostrate herb. Stems hirsute-strigose, diffusely branched at the base, 3—4 dm. long, tips of the branches sometimes arching. Leaves oblong to elliptic or obovate, rounded at the apex, serrate above the middle, 1—2 cm. long, 4—9 mm. wide, with long, scattered, villous or hirsute hairs on the upper surface and margins, and with that plus densely stellate-strigose hairs on the prominent midnerve of the lower surface. Stipules linear or somewhat spatulate, long-ciliate. Petioles 4—7 mm. long. Calyx 5 mm. long; lobes ovate, acute, long-ciliate, divided slightly more than halfway to the base, densely hirsute within. Petals yellow, 12—13 mm. long, ca. 5 mm. wide, obliquely spatulate, the margins minutely ciliate. Column terete, stout, 2—2.5 mm. from base of the petals to the filaments, with appressed, ascending, strigose hairs; filaments 1—2 mm. long. Anthers 15—20, dotted. Styles five, ca. 7.5 mm. long, ca. 4 mm. free; stigmas capitellate. Carpels five, strongly tetrahedral, 2.5—2.9 mm. high, 1.3—1.5 mm. radially, biapiculate, the apiculations ca. 1 mm. high; indument of upper dorsum rather densely multicellular-puberulent; outer dorsal spines reduced to a sharp 3-peaked ridge; lateral walls coarsely reticulate-rugose. TYPE: Leén & Roca 7466, “dry savanna, Chirigota, Pinar del Rio Prov., Cuba”, (Colegio de La ‘Salle, Cuba). DISTRIBUTION: Pine and oak woods and palm savannas of Pinar del Rio Prov., w. Cu This species, “ee only from Pinar del Rio Province, Cuba, is remarkable for the long-hirsute indument of the leaves and the pubescence of the carpels. There are specimens of S. ciliaris var. ciliaris from Brasil which are superficially similar to specimens of S. Brittonii. Schumann based S. ciliaris var. guianensis on speci- mens of this kind, but his variety and the specimens represent no more than a tendency in that species and have no close relation to this one. Cuba: Pinar veEL Rio: sitiagel et al. 6342, near Herradura (Ny). Britton et al. 6996, near Coloma (ny). Earle 748, Herradura (Ny). Ekman 10710, Herradura (Ny). Ekman 11054, Mantua (Fr). Ekman 18244, between Caldura and Pinar del Rio (ny). Herman hat near Herradura (pom, F, us). Killip 32370, nr. El Payuco, bet. Guane and Remates (us). Leédn & Roca 7466, La Chirigota (isoTyPE, NY). Palmer & Riley 56 nr. Coloma (us). Wright 2046, Asiento Viejo de San Julian (Ny, Gu, Us). 2. Sida centuriata Clement, sp. nov. Herbae perennes procumbentes. Caules 25 cm. vel breviores, cum pilis parvulis stellatis strigosis vestiti. Stipulae 5—8 mm., linearilanceo- STUDIES IN SIDA (MALVACEAE) 19 latae. Petioli 5—8 mm. Folia (7—) 10—23 mm. long., (4—) 8—12 mm. lat., elliptica vel anguste ovata, basi cordata, apice subacuta, trun- cata. Pedicelli 3 mm. Calyx 4.5—5.0 mm. long., cupuliformi-campanu- latus, lobis lanceolatis ca. 4 mm. long., 2.0—2.5 mm. lat. ad basem. Petala 18—22 mm. long., 10—14 mm. lat., obovata, conspicue obliquo- gibbosa, cum partibus paginae exteriore in alabastro expositis puberu- lentibus. Columna staminalis subangulata pilis sparsis stellatis patentibus, Filamenta 2—3-seriata. Antherae 30—40. Styli 11 mm., perfecte liberi. Carpella 7—8, a dorso conspicue verrucata et rugosa, omnino minute stellato-pubescentia et pulverulentia, a latere rugulosa, caeco-membrana- cea, nec cancellata. Perennial procumbent herbs from a woody root. Stems terete, 1—1.3 mm. diam. at base, to 25 cm. long, covered with small, stellate-strigose, appressed hairs. Stipules linear-lanceolate, 5—8 mm. long, ca. 0.7 mm. broad with scattered, marginal, pilose hairs, stellate-pubescent on the lower surface, glabrescent or glabrous on the upper. Petioles 5—8 mm. long, channeled adaxially, indument like that of the stem. Leaves (7—) 10—23 mm. long, (4—) 8—12 mm. broad, elliptic, or long-ovate, cor- date at the base, palmately 3-nerved, the nerves conspicuous on the lower surface; indument of small, yellowish, appressed, 6—7-rayed, stellate hairs, denser on the lower surface; margins serrate on the upper third to half; apex subacute to truncate. Pedicels 3 mm. long. Calyx cupuliform-campanulate, 4.5—5 mm. long, lobes ca. 4 mm. long, 2—2.5 mm. wide at the base, lanceolate, acute, tube with many fine veins, lobes with a single midvein and a double set of intramarginal veins separately confluent below the sinuses; exterior densely stellate-hirsutul- ous with a few long marginal and apical pilose hairs, interior of the tube glabrous, of the lobes hirsutulous. Petals obovate, strongly oblique, 18—22 mm. long, 10—14 mm. wide, 1.5 mm. wide at the claw, the margins with regularly scattered, short, fimbriate hairs, outer surface puberulent where exposed in bud. Stamen-tube subangulate, 3 mm. high from claw to level of filaments, with scattered, patent, stellate hairs. Filaments in 2—3 series, the lowest ca. 1.5 mm. long, longest 3 mm. long, flattened, tapering to the apex, recurved in anthesis. Anthers dotted, 30—40, ca. 0.8 mm. wide, 0.7 mm. high, globose, the sinus deep and constricted. Styles 7—8, free to the base, 11 mm. long, recurved in anthesis. Stigmas capitellate, the apex of the style dilated below them. Ovary 1.5 mm. in diam., 1.0 mm. high, densely stellate-puberulent. Car- pels 7—8; the dorsum strongly verrucate and rugose, the spines chiefly on the outer angle of the dorsum, everywhere finely-stellate-pubescent and pulverulent; lateral walls rugulose, opaquely membranous, not can- cellate. Seed plump, brownish-black, densely yellowish-stellate-puberulent 20 STUDIES IN SIDA (MALVACEAE) except the funicle; hilar depression rather shallow. Columella ca. 2 mm. high, subcylindrical, truncate, angulate TYPE: K. Fiebrig 4051, “N. Paniniay: Zwischen Rio Apa und Rio Aquidaban. Centurion, Okt. 1908.” (GH DISTRIBUTION: Known only from the type locality. S. centuriata is obviously in close relationship to S. ciliaris, differing from it primarily in having much larger flowers, with minor differences in the indument of the leaves and the carpels. The smallest open flowers on the three sheets cited have the petals 18 mm. long, the largest 22 mm. long. Paraguay: Fiebrig 4051, Oct. 1908, Centurion, between R. Apa and R. Aquidaban, n. Paraguay. (TYPE, GH). Fiebrig 4213, Nov. 1908, same locality. (AA, Us). 3, Sida surumuensis Ulbrich, Notizbl. 6: 320. 1915. Procumbent perennial herb. Stems few-branched, terete, tips ascen- dent, fulvid-scabrous to glabrescent, to 30 cm. long. Stipules 4—5 mm. long, linear, setose-fimbriate, persistent. Petioles 2—3 mm. long, chan- neled adaxially. Leaves narrowly linear, 10—20 mm. long, 1—2 mm wide, cordate at the base, subacute, the single nerve prominent beneath: indument of small, vitreous, stellate hairs on both surfaces or scattered on the lower surface only. Pedicels 1—2.5 mm. long, densely stellate- strigose. Calyx 5—6 mm. long, cupuliform-campanulate, densely and finely stellate-tomentose, the margins pilose; lobes 3—3.5 mm. long, 2—2.5 mm. wide at the base, acute or acuminate, pi iio Petals orange, yellow, or white, 13—16 mm. long, 4.5—5.5 mm. wide, obli- quely obovate, the inner margin concave, the outer convex; apex obtuse or subtruncate, margins with scattered very short ciliate hairs, the small portion exposed in bud with scattered stubby puberulence. Stamen-tube 2.5 mm. long from base of petal to level of filaments, heavily patent-stellate-pilose; filaments multiseriate, 1—2 mm. long. Anthers 25, dotted. Styles 7 mm. long, all but 1 mm. free; stigmas flat- tened-capitate. Carpels 7—8, verruculate, biapiculate, submuricate, with scattered, small, stellate hairs; ca. 2 mm. high; lateral walls rugulose. TYPE: Ule 8227, “auf trockenen rs bei der Serra do Mel, Rio Branco, Surumu. Brasil Aug. 1909.” DISTRIBUTION: Northern Rio Sac State, Brasil, and adjacent British uiana. This species is distinguishable from S. ciliaris by its somewhat verticillate linear leaves and large flowers. Superficially, it resembles S. ciliaris var. mexicana, differing from it, however, in STUDIES IN SIDA (MALVACEAE) 21 the longer and broader petals as well as in minor features of the habit, indument, and androecium. A certain amount of variation in the indument is found in the three localities from which the species is known, the type being rather heavily stellate-pubescent on both surfaces of the leaves, while that of British Guiana has the leaves glabrous or glabrescent on the upper surfaces. One of Tate’s two collection from Brasil resembles the type, the other the Guiana material. British Guiana: A. C. Smith 2340, basin of Rupununi R., nr. mouth of Brasil: Rio Branco: E. Ule 8227, nr. Serra do Mel, Surumi, (ISOTYPES: NY, UC, US; TYPE photographs: Gu, F, us). G. H. H. Tate 85, Limao (ny). G. H. H. Tate 125, Limao, (ny). 4. Sida paradoxa A. P. Rodrigo, par: Mus. La Plata 2: 105, figs. 3,5, P14. 1957. Perennial semidecumbent herb. Stems leafy, many-branched, slightly woody, 20—25 cm. high, the indument of whitish stellate hairs. Roots strong, woody, deep. Leaves numerous, and clustered at the nodes, emer- ald-green in ‘life: stipules lanceolate, 5—-7 mm. long; petioles as long as or shorter than the blades; blades oblong or slightly cuneate, truncate, 4—8, rarely to 11 mm. long, 1.5—3.5 mm. wide, the base obtuse or narrowly subcordate, the apex tridentate, the lateral teeth larger, the central one shorter, usually no more than an apiculation; upper surface glabrous, lower surface with scattered to dense white stellate hairs, the midnerve prominent. Flowers solitary or usually in clusters of two to four, rarely five, at the ends of the branches, the foliaceous bracts of the inflorescence linear, slightly spatulate, sparsely stellate-pubescent. Peduncles 2—4 mm. long. Calyx angulate, 8 mm. high, 3 mm. wide, covered with fine, white, stellate hairs, 5-nerved; lobes cordate-acumi- Mate, two-thirds the height of the calyx, margins fine-ciliate; inner sur- face of the lobes with ascending, appressed, fine hairs; inner surface of the tube glabrous. Petals yellowish pink, veined in red, with a tuft of white ciliate hairs at the base, strongly oblique, obtuse, ‘widest near the apex, 14 to 17 mm. long and 8—11 mm. wide, the corolla to 25 mm. in diameter. Stamen-tube glabrous, very short, 1—1.5 mm. long, the filaments 1.5—2.0 mm. long; stamens ca. 20. Styles five, pink, 6—7 mm. long; stigmas apie ovary conoid. Mature carpels five, 3 mm. high, biaristate, the aristae 1—1.5 mm. high, they and the dorsum with numerous small, stellate, ese hairs; dorsum rugose, the outer margins few-spined; lateral walls reticulate; pulverulent; dehiscence along inner margin, nearly complete. Columella 3 mm. high, knobbed at apex, flared 22 STUDIES IN SIDA (MALVACEAE) at base. Seeds 2 mm. high, 1.8 mm. wide, with minute, scattered, cin- ereous stellate hairs. TYPE: A. P. Rodrigo 605, Mercedes, Arroyo La Garza, center of Cor- rientes Prov., Argentina (Museo La Plata) DISTRIBUTION: Known only from the type-area. I have seen only one sheet of this species, but this and Rodrigo’s original and subsequent descriptions and illustrations (1937, 1944) convince me that S. paradoxa is a distinct species. It probably arose as a local segregate of S. ciliaris and differs from it mainly in having numerous, small, clustered leaves and large flowers. The specimen cited does not have the long-aristate carpels des- cribed and figured by Rodrigo and its leaves are shorter. Argentina: Pedersen 121, Corrientes, Estancia Sta. Maria (us). 5. Sida ciliaris L., Syst. ed. 10, 1145. 1758. Perennial herbs or subshrubs. Stems sublignescent to herbaceous, nu- merous, prostrate to ascending or erect from a branching or single stout woody root, clothed with appressed, rigid, stellate or submalpighiaceous hairs, often becoming hirsute to tomentose toward the apices. Leaves extremely variable, linear to linear-oblong to oblong to ovate to sub- orbicular; the base truncate to rounded to narrowly cordate; the apex truncate to obtuse to acute; the margins variably serrate, only at and near the apex or merely tridentate, to slightly more than halfway toward stellate or submalpighiaceous hairs, scabrous to softly oe the mat- gins occ asionally short-ciliate. Petioles 3—9 mm. long, ca. mm. lower portion flattened Sanpack upper portion terete or subterete, geniculate at the apex; the indument variable, of flavid, cinereous of white, stellate or submalpighiaceous and ciliate hairs. Stipules linear to spatulate, 6—12 mm. long, the apex acuminate to bluntly acute, ciliate; stipules of the involucre somewhat longer and longer-ciliate, the hairs flavid, cinereous or white. Pedicels from less than 1 to ca. 2 mm. long, involucrated in two- to eight-flowered heads at the apices of oe stems, rarely solitary in lower axils. Calyx 5—7 mm. long, campanulate, terete, lobes ovate or triangular to lanceolate, 2—4 mm. long, acute; indument villous, of simple hairs, to subtomentose, of stellate hairs, or ‘teived: the hairs or individual rays occasionally to 2.5 mm. long, white, cinereous , or ferrugineous-flavid, the lobes puberulent near the margins within, STUDIES IN SIDA (MALVACEAE) 23 the tube glabrous within. Petals 3—15 mm. long, 2—7 (—10) mm. wide, white, yellow, or variously rose or purple, often darker at the base, ovate-spatulate or broadly rotund-triangular, slightly to markedly obli- que, the apex obtuse or retuse, the margins ciliolate, the base glabrous or sparsely pilosulous. Stamen-tube slender, terete, 3—5 mm. high, gla- brous or sparsely pilose with spreading, simple, hyaline hairs; filaments 2—3 (—4) mm. long, slender, reflexed in anthesis; anthers 10—40, minutely dotted or plain. Ovary five- to eight-lobed, glabrous or min- utely hairy, conoid or subglobose, the apex truncate to apiculate. Styles 4—8 mm. long, 2—5 mm. free; stigmas capitellate. Carpels 5—8, ex- tremely variable in the sculpturing, muticous or rostellate or biaristate, the dorsum muricate or reticulate or rugose, glabrous to densely stellate- pilosulous, thick or fragile; the lateral walls finely to coarsely reticulate, membranous between the reticulations or rugosities or cancellate; 2— mm. high, 2.5—3 mm. radially. Seeds usually with scattered stellate pu- berulence, denser near the hilum, 2—2.5 mm. radially, light brown, yellowish or nearly white in maturity, rarely purpurascent. KEY TO THE VARIETIES A. eit: suborbicular to ovate, obtuse or truncate (becoming lanceolate and e in nerd intermediate between vars. ciliaris and anomala); petals ai 0 mm. long, 2—4 mm. wide; plants prostrate to ascending B. Carpels 5k briefly biaristate, muricate to verruculate, the lateral wills ment of the involucre sparse to peace, white cinereous; solitary axillary flowers sometimes prese . . Sa. S. ciliaris var. ciliaris. B. Carpels 5, rarely 6, muticous or sbi the oie walls membranous, weakly reticulate; indument of the involucre dense, flavid or fulvous; flowers numerous in the involucres, never solitary 5b. S. ciliaris var. fulva. A. Leaves linear to narrowly oblong to elliptic or Sacechaiks the margins ser- rate usually in the upper part only or merely tridentate; petals 5—15 mm. long, 2—10 mm. wi SS ge Pe a an gr ak ae ae, Leaves narrowly oblong, often cuneate, truncate to acute; petals 8—15 mm. long, 7—10 mm. wide, ea ed oblique; boas 5—8, muricate, biaristate Sc. S. ciliaris var. Leaves linear to finear-lanceclate or dilipcic a acute; eae slightly oblique, 2—4 mm. wide D Carpels 5, rarely 6, biapicalate, muriculate: leaves linearlliptic to elliptic to 17 mm. long and 4.5 mm. wide; petals 5—6 mm. long. 3 mm wide; stamen-tube glabrous; age seein nd 5d. 2) S. ciliaris var. involucrata. D. Carpels ‘ieoally 7, shortarisate, muricate; leaves linear to linear-lanceolate, 21 1—14 mm. long, 3.5 mm. wide; stamen-tube with ao hairs; plants ascending to erect. . ‘Se. S. ciliaris var. mexicana. 24 STUDIES IN SIDA (MALVACEAE) 5a. Sida ciliaris var. ciliaris. S. muricata Cav., Ic. 6: 78, f. 2. 1801. S. erosa Salzm. ms. in Tr. and Pl., Prod. Fl. Nov. Gran. 1: 176. 1862. (nomen nudum). S. ciliaris var. guianensis K. Schum., Mart. FI. Bras. 12 (3): 284. 1891. S. ciliaris f. flava O. Ktze., Rev. Gen. 3 (2): 22. 1898. S. longistipula Merr., Phil. Journ. Sci., Bot. 13: 30 1918. TYPE: Browne, Jamaica (Linnaean Herbarium). In publishing this species, Linnaeus cited a plate in Sloane’s Natural History of Jamaica. Unfortunately, the plate shows a plant which is very doubtfully of this species, since it much more resembles S. spinosa. The phrase in the type-description “semini- bus muricatis”, however, can refer only to S. ciliaris, and the additional phrase “fol. ....retusis” certainly does not refer to the acute leaves of the cited figure, which finds no match in the Jamaican material of the species which I have seen. Linnaeus had purchased Browne’s Jamaica collections before the species was described, and undoubtedly his original description was based on the specimen which he labelled “Sida ciliaris’ and “Br.”, for Patrick Browne (Savage 1945). DISTRIBUTION: Florida Keys and Texas; Bahamas and Antilles (except Puerto Rico); Mexico to Bolivia and Argentina; Philippines (introd.). 5b. Sida ciliaris var. fulva (St. Hil.) K. Schum., Mart. Fl. Bras. 12 (3): 284. 1891. S. plumosa Cav., Diss. 1: 7. t. 12, f. 4. 1785. Malachra plumosa (Cav.) Desr., Lam. Encyc. 3: 363. 1791. S. fulva St. Hil., Fl. Bras. Mer. 1: 176. 1827. TYPE: The type of S. plumosa, the oldest name for the taxon, is a Commerson specimen from Brasil loaned by Thouin to Cavanilles in Paris. There is a probable isotype of this collection in Madrid. DISTRIBUTION: Southern Brasil. 5c. Sida ciliaris var. anomala (St. Hil.) Hochr., Ann. Cons. Jard. Bot. Gen. 6: 35. 1902. S. anomala St. Hil., Fl. Bras. Mer. 1: 177. 1827. S. ciliaris var. anomala K. Schum., in Engl. & Prantl, Nat. Pfl. STUDIES IN SIDA (MALVACEAE) Fe 3, Abt. 6: 43. 1890. (nom. nud.). S. anomala f. albiflora Chod. & Hass., Bull. Herb. Boiss. ser. 2, 5: 290. 1905. (including subforms). S. anomala f. roseiflora Chod. & Hass., l.c. (incl. subforms). S. anomala f. violaceiflora Chod. & Hass., op. cit. 291. TYPE: St. Hilaire C2/2470 “prés le village de Sando” Paisandu, Uru- guay (P, specimen in F pei “Santos” probably of this collection; St. Hilaire was never in Sant ee Colombia, certs Brasil, Uruguay, Paraguay and Argen 5d. Sida ciliaris yar. involucrata (A. Rich.) Clement, stat. nov. S. involucrata A. Rich., Ess. Fl. Cub. 162. 1845. ? S. tridentata Cav., Teadies at 6: 1797 TYPE: de la Sagra “crescit in incultis ‘ent Cuba” (p, fragment in F). DISTRIBUTION: Bahamas, Cuba, Hispaniola and Lesser Antilles; Yuca- tan peninsula of Mexico 5e. Sida ciliaris yar. mexicana (Moric.) Shinners, Field and Laboratory 21: 94. 1953. S. anomala var. mexicana Moricand, Pl. Nouv. Am. 36, t. 24. 1837. S. fasciculata Torrey & Gray, Fl. N. Am. 1: 231. 1838, non Willd. Malvastrum linearifolium Buckley, Proc. Acad. Sci. Phila. 1861: 442. 1862. S. ciliaris var. fasciculata (Torr. & Gray) Gray, Proc. Am. Acad. 22: 294. 1887 TYPE: Berlandier exsicc. 66, “circa Tampico de Tamaulipas” (G). DISTRIBUTION: Texas and Mexico to Panama. The species is variable nearly throughout its range and is especially so in South America. A major problem arises in the satisfactory disposition of the element usually identified as S. anomala. The type-specimen of S$. anomala, from Uruguay, is not what most authors consider it to be. Its leaves are oblong, slightly cuneate and mostly truncate, rather than linear and acute; its carpels muricate only on the dorsal margins rather than on the whole dorsum and on the lateral walls; and its petals 5—8 mm. long rather than around 10 mm. It is, as it is usually described, 26 STUDIES IN SIDA (MALVACEAE) few-stemmed and apparently erect, and is, in sum, intermediate between the extremes usually separated as S. ciliaris and anomala. There is no truly linear-leaved material in South America, although this character has appeared in most keys because Schumann, whose key has usually been taken over out- right by later authors, included S$. anomala var. mexicana, here considered a variety of S. ciliaris, as a synonym of S. anomala. I do not think that S. ciliaris var. mexicana and S. ciliaris var. anomala have any close relationship but are, instead, two separate tendencies in the species, both having narrow leaves and large flowers. Both intergrade with the typical smaller-flowered, broader-leaved material, and in the South American material, such combinations as large flowers and small obovate and truncate leaves occasionally occur, as in Swallen 9507 (us). Despite the number of intermediates, I have attempted here to define these tendencies as separate varieties, but it should be understood that they are only tendencies. Only those specimens representing the non-typical varieties in or near their maximum expression have been listed under them in the specimen-citation. Other tendencies in the species in South America have been named. Of these, only S. ciliaris var. fulua has any constancy, although a few intergrades between it and the typical state are present. S. ciliaris var. guianensis, even in the specimens cited by Schumann, varies continuously from the large-leaved, six-carpelled extreme to the typical, and I have not retained it. The color- forms and pubescence-subforms of Kuntze and Chodat & Hassler, described under both S. anomala and S. ciliaris, are of no real worth in a species where color-variation is common, even in local populations, and pubescence-variation is still more common. The species appears to be one of hybrid origin which is still in the process of segregation and recombination in much of its area. Only in a few areas such as Jamaica is there any constancy in the nature of the population. In the southern part of its range, as in the northern, a tendency to narrow leaves and large flowers is strongly evident, and to a lesser extent in the intervening area. Centering in Cuba, there is a third narrow-leaved element, here named as var. involucrata, but the flowers here are small, about the size of the typical. The other species of this section are un- doubtedly segregates from this complex which have achieved genetic or geographic isolation, and I can think of no group in STUDIES IN SIDA (MALVACEAE) 27 the genus where genetic and cytological studies of inter- and intra-specific variation might be more rewarding. 5a. Sida ciliaris var. ciliaris United States: pee Blodgett, Key West (Gu, us 2). Chapman, Key West (GH). Curtiss 5445, Long Key (GH 2, vec, us). Hitchcock, March 1906 Small 10108, Big Pine Key (Gu); Small 10217, Key West (us). Texas: Cory 5910, w. of Menthalia, Gonzales Co. (GH). Heller 1567, near Regie San Patricio Pes (Ny, us). Palmer 91, Sutherland Springs, Mere (GH, us) : Baya Cauirornia: Jones 24127, Todos Santos (us). Joe oder Todos Satine (POM). CAMPECHE: poke 1812, Chanter (F). LIMA: Ferris 6095, vicinity of Manzanillo, Colima (ps, GH, us). GUERRERO: pear 6490, Coyuca, Coyuca (GH, Ny, us). Palmer 30, mais and vicinity (GH, MO, NY, us). JaLisco: Mexia 1155, Puerto Vallarta (GH, Mo, 2: POM, UC). Pringle 4497, near Guadalajara (MExU 2, MO, UC, NY, GH). Rose 2581, between Huajuquilla and Mosquitac (us). Rose 3696, Bolanos (GH, us). eee Hinton 1469, Dist. of Temascaltepec, Calera (F, MEXU 2, Us). MicHOACAN: Hinton 15181, Apatzingan (GH, NY). Oaxaca: Conzatti 1600, Oaxaca (Us). Gonzales 288, El Forties (cu). Nelson 1231, Valley of Oaxaca (us). Orcutt 5028, Puerto Angle (ps). Smith, L. C. 510, Cuicatlan (GH). StnaLoa: Brandegee, 8 Oct. 1893, Mazatlan rae Brandegee, 13 Sept. 1904, vicinity of Culiacan (Gu, uc). Brandegee, 1 Nov. vicinity of Culiacin (us). Mexia 64, hill “La Neveria’, near Mazatlan (ey Mexia 168, “La Noria’ (mo, uc). Mexia 350, “La Noria” (Pom, MO, UC). Narvaez Montes & Salazar 666, San Ignacio (us). Ortega 4361, Gerke: de la Manatee Lagoon (cu). Espat 35, Bakers, near Belize R. 18 miles from Belize (F). Guatemala: Heyde 175, Guatemala (us). Heyde & Lux 3948, Carrizal, Dept. Santa Rosa (GH, NY, MO, US). Standley 24379, vicinity of Los Amates, Dept. de Izabal (Ny, GH, us). Standley 59629, near Fiscal, Dept. Guatemala (Ny, F). Standley 73600, vicinity of Zacapa, Dept. Zacapa (F). Standley 73928, Dept. capa: Bafios de Santa Marta, north of Zacapa (F). Standley 74373, Dept. Chiquimula: Quebrada Shusho, above Chiquimula (F). Standley 75038, Dept. Jutiapa: vicinity of Jutiapa (F). Standley 78805, Dept. Santa Rosa: southeast of Chiquimulilla (F). Standley 80467, Dept. Guatemala: near Fiscal (r). Steyer- mark 29077, Dept. Zacapa: near Estanzuela (F). El Salvador: Standley 19835, nr. Ahuachapan, Dep. Ahuachapan (GH, us). Honduras: Rodriguez 312, re Dept. Morazién (Fr). Yuncker, Dawson & Youse 5812, Siguatepeque (Mo, GH Nicaragua: Maxon et al. 7584, near ‘Granade (us). Costa Rica: Brenes 22814, “Los Loros’’, Pacific coast (F). ama: Pittier 4843, Aguadulce, Prov. of Coclé (Ny, GH, us). Sta sere Francisco, Prov. of Panama (us). Standley 31863, between Las ‘Sabanas a Matias Hernandez, Prov. of Panama (us). 28 STUDIES IN SIDA (MALVACEAE) Bahamas: Brace 132, New Providence (ny). Hitchcock, 12/3/90, Inagua (mo). Millspaugh & Millspaugh 9099, East Caicos, Jacksonville and vicinity (us, Ny, F). Wilson 7820, Great Ragged Island (cH). Cuba: Ekman 8243, Prov. Oriente, Maria Pilar, on Rio Baconao (ny). Bro. Hiram 2274, Boca del Jaibo (ny). Shafer 2482, Cayo Romano, Camagiey (GH, NY, us Jamaica: eae 1850 (ny). Britton 785, vicinity of Kingston (Ny, F). rey 6023 (ny). Harris 9768, Long Acre Point, west of Black River (ny, us). Harris 12044, near Spanish Town, St. Catherine (GH, mo, Ny, us). McNab (GH, NY). Haiti: Ekman H8396, St. Michel, towards I'Halaye (us). Leonard 2855, vicinity of St. Marc, (us, GH, Ny). Leonard 7312, vicinity of St. Michel de I'Halaye, Dept. du Nord (us). Leonard 7307, vicinity of St. Michel de I’Halaye, Dept. du Nord (cu, ny, us). Leonard 12610, vicinity of Jean Rabel (GH, Ny, US). Leonard 9855, vicinity of Gros Morne, Dept. de I’Artibonite (uc, us). aoe & Leonard 15107, vicinity of Bassin Bleu (us). Leonard & Leonard 11052, vicinity of Port de Paix (us, Mo). Nash 938, Cap Haitien (nx). Potter 5026, Port au Prince (GH). minican Republic: Abbott 866, Guayubin, Prov. de Monte Cristi (us). Ekman H15996, Valle del Cibao, Prov. de Santiago, Hato del Yague (us). Fuertes 20, Prov. Barahona (us, ny). Fuertes 1018, ae: Barahona (ny). Valeur 246, Moncion, Prov. of Monte Cristo (mo, vs, Y, POM, Ds, uc). Wright, Parry & Brummel 46, Santiago i: los Caballeros fay Bertero, Sento Domingo” St. Croix: Haunen 159 (ny). Ricksecker, A. E. 3, Bassin yard (us, NY, GH, MO). Richierker, J. J. 260, Bassin yard (us, Mo). Thompson 42, Anna's (GH). Thompson 333, Coakley Bay (cu). St. Jam: Britton & Shafer 329, Betheania (us, ny). Virgin Gorda: Fishlock 163, “valley” (ny). Anegada: Britton and Fishlock 1022, “rocky plain near the settlement’ (ny). Fishlock 33, “near a small pond” (cu). Tortola: Britton & Shafer 687, road, Town to Sea Cow Bay (us, ny). St. Thomas: Britton, Britton & Shafer 36, Belgian Road (us, F, NY). Kuntze 139 (Ny). Holton (ny). Eggers, 16 May 1876 (mo). Eggers, July 1887 (us). Eggers a ee (us). St. Martin: Britton & Cowell 87, élemy: Questel 187, Gustavia (ny). Questel 298, Gustavia (Ny). Questel ote fo le Fort (ny). St. Eustatius: Boldingh 1153 B (ny). Antigua: Box 1073, High Point (cu, us). Guadeloupe: Duss 2790 (us). Duss 2790.4116, “St. Francis, Désirade. Les Saintes. 1892” (ny). Duss 4141, along the Marne (ny). Stehlé 148, Marie-Galante, sands of Grand Bourg (us). Stehlé 194, Désirade (ny). Stehlé & Quentin 5294, Désirade, “prés du Phare” (us). Martinique: Duss 862, St. Anne (ny). Steblé 5443, near Madiana (us). Barbados: Gooding 583, Gibbon’s Christ Church (ny). British Guiana: Schomburgk 405, Pirara (Isotype of S. - var. guianensis, us). Brasil: Allemao 96, Ceara (us). Dahlgren 854, Ceara, Fortaleza region, Kagado (Fr). Drouet 2317, State of Cearé, Municipio de Zoitelens (cH, F). Drouet ote Ceara, Fortaleza (F, GH, NY). Gardner 2052, Prov. de Piauhy (GH, US, F, NY 2). Kuntze 1799, La Guayra (ny). Pickel 3991, Pernambuco, Russinha rte it F). Swallen 9507, Bonito, Mato Grosso (us). STUDIES IN SIDA (MALVACEAE) . 29 Paraguay: Hassler 1238, “Duarte” (ny). Hassler 2491, Cerros de Paraguary (cu). Hassler 7508, near eee (cH). Hassler 12660, Cordillera de Altos (Us, MO, GH). Jérgensen 4387 (F, NY 2). Kuntze, Sept. 1892, Concepcidn de Paraguay a Kunize, Sept. 1892, «Sud. Paraguay”’ tee Morong 240, Asuncién Argentina: Aguilar 480, Chaco, Dep. Resistencia Margarita Belén (us). Jorgensen 2312, Formosa, in Territorio de Formosa Bolivia: Bang 934, vicinity Cochabamba sr us). ie 4115, nr. Serrano, ea (us). Fiebrig 2551, Padcaya (us uador: Asplund 5619, Prov. Guayis, clan (us). Eggers 14813 (F). PN hi 580, environs of Guayaquil (us). Svenson 11238, Prov. Guayas, Salinas (Ny). Philippines: Ramos 27311 and 32704, Burgos, Luzon (us). Ramos 27492, Bangui, Luzon (us). 5b. S. ciliaris var. fulva Brasil: Barreto 5245, Cruz das Almas, Andrelandia (Fr). Burchell 958 (cu). Commerson (photograph of probable type, Pp: F). Cummerson (photo. of 1soTyPE in herb. Cav., Madrid: us). Gardner 12, Rio Comprido, Rio de Janeiro US, GH). Hoses 19516, Canelleiras, Ouro Fino (Ny). Lutzelburg 339, Bahi (ny). Martius Aug. ’70, “in aridis ad vias Sebastianopolis’, Rio de oe (photo. us). Riedel 1332, Rio de Janeiro (cu, us). Riedel 2026, near Yta (us). Rose 19688, vicinity of Bahia (us). Schwartze 14684, “east of Rio” (us). Smith, , 1637, Rio de Janeiro, vicinity Monte Serrat, Mt. Itatiaya re us). Usteri, 18 March 1906 (ny). 5c. S. ciliaris var. anomala Colombia: Le/mann 2276, Tolima (us). Lehmann 6051, Neiva (F). Lehmann 8719, El Gigante and Altamira, Rio Magdalena (ny, cH). Venezuela: Holt and Gehriger 37, Anzoategui, Soledad (us). Brasil: Dias da Rocha 4, Ceara (us). (TYPE, P, specimen in F labelled “Santos’’ probably of this collection). Paraguay: Fiebrig 910, without locality (Fr, GH). Hassler 6388, Cerros de Tobaty (cH) Hassler 6615, Central Cordillera, Rio Y-aca (Gu). Jérgensen 4287, Villa Rica (us). Argentina: Jérgensen 2315, Chaco Territory, Las Palmas (GH, Us Lorentz 1762, Dec. 1877, Concepcién del Uruguay, Entre Rios Prov. (GH * Venturi 9120, Formosa (us). 5d. S. ciliaris var. involucrata ico: YUCATAN: Gaumer 779, Izamal (GH, DS, US, NY, UC, MO). Gaumer Brace 131, New Providence (F). Brace 4909, Ruaria Hill, Long Cay (ny). Brace 4257, Acklin’s Island, Spring Point (Ny, us, F). Britton 3369, Tea House, New Providence (Fr, us, GH, MO, NY). Britton & Brace 201, Farringdon Road, New 30 STUDIES IN SIDA (MALVACEAE) Providence (Ny, F). Britton & Millspaugh 2248, Little Harbor Cay, Berry Islands NY, us). Britton & Millspaugh 5467, Eleuthera, Governor's Harbor and vicinity agua, Mathew Town (ny). Wight 22, near Fort Charlette, Nassau (GH, NY). Aa 98 7376, Atwood Cay po Wilson 7820, Great Ragged Island (Ny, MO). Cuba: Baker & Wilson 2342, Batabandé, Prov. Habana (Fr, pom, us). Britton & Earle 7566, Prov. of Pinar del Rio, Bay of Mariel (ny). Curtiss, March 1904, north of Marianao, Habana (ny). Palmer & Riley 1050, near Nueva Gerona, Isle of Pines (ny). ‘Haiti: Leonard & Leonard 15852, vicinity of Port au Prince US). 5e. S. ciliaris var. mexicana United States: Texas: Berlandier 3035, without locality (us). Bigelow, San Antonio (Ny). Bush 293, Brazos R., Columbia (GH, Ny, us). Cory 14187, Queen City Falfurrias (GH). Cory 55767, 6 mi. WSW of Giddings, Lee Co. Drummond 37 (Type of S. fasciculata, ny). Drummond 47 (Gu). Drummond 50 Willacy Co. (ps). Lundell & Lundell 10036, nr. San Manuel, Hidalgo Co. (ps). McDonald, nr. Bastrop (us). Metz 745, 20 mi. south of San Antonio, Bexar Co. (uc). Nealley, Nov. 1884, Houston (cH). Ramyon 2441, Yturria, Willacy Co. (ps, uc). Snyder, 1905, Cypress (uc). Tharp 1170, Brownsville (us). Tharp 1392, Austin (us). Tharp et al. 49172, 18 mi. southeast of Yoakum, Lavaca Co. (us). Thuron 12, Hockley (us). Wright, Eagle Lake (us). Mexico: Berlandier 66, nr. Tampico, Tamaulipas (1soryre, Mo). Haenke, Mexico (mo). Mel] 2108, San Gerénimo, siege (ny, us). Mexia 963, trail from Los Labrados to La Marisma, Sinaloa (ps, GH, MO, NY, UC, US 2). Ortega Honduras: Molina R. 278, Maraita (us). Rodriguez 510, road to Tatumbla, Dep. Morazan (F). Rodriguez 3297, San Lorenzo, Dep. Valle (Fr). Rodriguez 3438, San Lorenzo (F). Williams and Molina 10211, nr. Galeras, Dep. El Paraiso (uc). Costa Rica: Shannon 5006, Volcan Orosi, Prov. Guanacaste (US). Panama: Allen 817, Nata (F, GH, MO, NY, US) II. Section PHysatopes A. Gray, Smiths. Contr. Knowl. 3, art. 3: 20. 1852. Section Calyxhymenia A. Gray, Proc. Am. Acad. 22: 294. 1887. Calyx much inflated after anthesis, the lobes deeply divided, the margins plicate; carpels muticous, lateral walls persistent. The section was founded by Gray in the following words: STUDIES IN SIDA (MALVACEAE) 31 “this species [S. physocalyx Gray], S. hastata, St. Hil., and S. physalodes, [sic] Presl, form a section remarkable for the bladdery inflated calyx, which may be distinguished by the name of Physalodes”. In the later paper, he described section Calyx- hymenia as follows: “To the peculiar sections named in PI. Fendlerianae.... a third may be added, Calyxhymenia, for species which have the ebracteolate calyx much accrescent around or under the fruit, and membranaceous or scarious, — the name taken from S. calyxhymenia, Gay, of Australia, .... Our species, S. physocalyx, Gray, Pl. Lindh. 2, 163, in which the 5-parted and angulate bladdery fruiting calyx imitates that of Nicandra, has rather peculiar and very thin-walled reticulated indehiscent carpels with a beak-like apex. The homonymous S. physocalyx F. Mueller from Australia is much later, and will find another name”’, Schumann later showed that S. physocalyx Gray and S. hastata St. Hil. were synonymous, the first described from Texas, the other from Brasil. He apparently did not realize that St. Hilaire’s name was a later homonym for S. hastata (Cav.) Willd. (1801), and for S. bastata Sims (1811), since he placed Gray’s name in the synonymy, which has been done by almost all workers since. The nomenclatorial type of the section is, then, S. physocalyx A. Gray, while the type-collection of that species and thus of the section is St. Hilaire’s “In provincia Cisplatina” Uruguay, in the Paris herbarium. A photograph of this is reproduced in Rodrigo (1944). All of the previously known American species of the section are represented in the flora of Argentina and Uruguay, and Rodrigo’s analysis of them (1944) is satisfactory in most respects. She added S$. decumbens St. Hil. & Naud. to the section on the grounds that it has the calyx inflated after anthesis and muticous carpels. Hochreutiner had transferred the species to Anoda because of the disappearance of the lateral walls of the carpels in maturity, but Kearney (1954) retains it in Sida. Even if it be ept in Sida, it does not belong in the section. I have here added to the section a new species from Mexico. oO Baker placed a number of Australian species in the section, and of these, S. calyxhymenia Gay ex DC., and S. physocalyx F. Muell. non Gray definitely belong here. Study of types, authentic material and published descriptions of the other species assures 32 STUDIES IN SIDA (MALVACEAE) me that they do not belong here: S. Kingii F. Muell. has a some- what accrescent calyx, but the spinulous fruits are quite unlike those of any species of this section. The calyx of S. cleisocalyx is not at all accrescent, while that of S. inclusa and of S. platy- calyx is of a very different type with numerous ribs. S. inclusa, furthermore, has a tremendous globose fruit, beset with spines clothed in stellate pubescence. S$. Clementii Domin was related to S. calyxhymenia by its author, but the material I have seen of it differs in a number of details from that species and the calyx is only slightly accrescent. Type species: Sida physocalyx ray. KEY TO THE SPECIES A. Carpels ten to seventee Be pe ene ee ee B. Carpels ten to fourteen. "ee World) . ~ ow a G. Si physocalyx. B. Carpels fifteen to seventeen. (Australia) . : . 7. §. Brownis. A. Carpels five or eight, rarely nine ar aay Gee ee. C. Cerpels: eight or tercly. nine... 2 Bc Sere, C. Carpels five . hone ae nem tes D. Flowers uieeted fous to sixteen in the upper axils. . 9. S. urticaefolia. D. Flowers solitary or paired in the axils, rarely in threes . E E. Plants erect, leaves rounded at the base, flowers usually solitary. ‘in the axils. (Australia) 1G. 5, calyxhymenia. E. Plants prostrate, lekvex aidan at he bie: Howenk usually paired in the axils. (New World Hes F F. Petals 5 mm. long, stamens ca. cwensy: Gkeiace ok Leas is 5. Standleyi. F. Petals 8—9 mm. long, stamens 30—40. (S. America) 12. S. flavescens. 6. Sida physocalyx A. Gray, Bost. J. Nat. Hist. 6: 163. 1850. S. hastata St. Hil., Fl. Bras. Mer. 1: 190. 1827, non Willd., Sp. Pl. 3: 763. 1801. S. hastata var. tomentosa R. E. Fries, K. Sv. Vets. Handl. 42, £2: 39. 1908. S. hastata var. glabriuscula R. E. Fries, oe S. inflata Larrafiaga, Escritos 2: 218. Perennial herb from a thick, fusiform, ee taproot; stems decum- bent or prostrate, 10—30 cm. long, few- to many-branched, dark green, covered more or less densely with 4-rayed submalpighiaceous hairs, hya- line or yellowish. Leaves suborbicular, ovate, or oblong, or long-elliptic, to 6 cm. long and 5 cm. wide, the base cuneate to truncate, sometimes oblique; the apex rounded to acute; the margins serrate, sometimes irregularly; nerves 5, palmate from the base, prominent beneath; the indument variable, that of the upper surface from glabrescent to mo- derately stellate-tomentellous, the hairs fine or coarse, sometimes like STUDIES IN SIDA (MALVACEAE) 33 those of the stem; indument of the lower surface like that of the upper, usually much denser; occasionally both surfaces with simple, appressed, pilose hairs, intermingled, of various lengths. Petioles 1.0—2.2 cm. long, always shorter than their leaves, the indument like that of the stem. Stipules linear-lanceolate, persistent, yellowish-ciliate, ca. 3 mm. long, ca. 0.8 mm. wide. Pedicels solitary and axillary, 12—20 mm. long, filiform, finely stellate-hirsute, articulated in the upper fourth. Calyx greatly accrescent during the whole period of growth, strongly plicate. the lobes and subjacent tube flat or concave, during anthesis ca. 5 mm. high, 4 mm. wide, the base of the tube rounded, later impressed; the lobes soft-awned, the lobal face of the calyx almost cordate in appear- ance; the indument like that of the leaves. Petals white, yellow or rose, veins and base darker, gibbous, the outer portion rotund, one margin concave, the other convex; 1.2—2.0 cm. high, 5—9 mm. wide, the claw 1.0—1.5 mm. wide, entirely glabrous. Stamen-column 5 mm. high, terete, with five pairs of strongly marked traces; the base of the column broadly dilated, glabrous; the filaments 1—2 mm. long. Anthers ca. 40, pale, not dotted. Ovary 1.3 mm. high, ca. 2.5 mm. wide, the lower third impressed, the apex subapplanate. Styles 5—G mm. long, 2.5—3 mm. free; the stigmas capitellate. Carpels 9—14, ca. 5 mm. radially, 3 mm. high, forming a flat-topped fruit, blackish, the rostrum prolonged as a long horizontal beak over the apex of the columella; the dorsum many- nerved, impressed transversely over the hilar depression of the seed, sharply impressed radially on the outer surface, the outer margins crest- ed; the lateral walls horizontally compressed-rugose; dehiscence by the tardy disintegration of the apex. Seed black, with minute, rather thickly scattered, stellate, white, appressed puberulence, 2—3 mm. high and wide, the funicle black, glabrous, flattened. Columella capstan-shaped, 1.0 mm. high, 1 mm. wide at the top, 2 mm. at the base, the sides concave in outline, the top stellate in vertical view, black. TYPE: St. Hilaire “Assez commun sur les bords des chemins dans la province Cisplatine”. Uruguay (P). DISTRIBUTION: Texas, New Mexico, Arizona; northern Mexico, Bra- sil?, Uruguay, Argentina, Bolivia. The problem of typification of the section rests on the typi- fication of S, physocalyx and has been discussed previously. The distribution of the species is unusual but not unprecedented; Schumann (1891) pointed out that S. leprosa has a rather similar bicentric range, as do species of the genera Mentzelia, Daucus, Cuphea and Bouchetia. Two closely allied species of this same section also show this pattern when their ranges are mapped together. The presence of the species in Bolivia is based on a 34 STUDIES IN SIDA (MALVACEAE) single O. Kuntze collection which is not in mature fruit, but it occurs in adjacent Jujuy province, Argentina, and will probably be found again. Fries’ two varieties, based almost solely on the leaf-pubescence, were supposedly geographic, his var. glabriuscula found through- out the range and var. tomentosa confined to the South American part of it. In actuality, both are seen to occur at random through- out the range when a long series of specimens is studied, and there are so many intergrades that they are not even worthy of being considered forms. Larrafaga’s field notes, unfortunately published, named as S. inflata a plant he had observed on April Fool’s Day 1814 which had solitary flowers in the axils, an inflated calyx and eleven carpels and which was undoubtedly S. physocalyx. The extraordinary carpel-shape in this species finds its parallel in that of S$. aprica Domin of Australia. Examination of authentic material of that species from Kew shows that it does not belong to this section, the calyx being very little inflated, nor does that species resemble S$. physocalyx very closely in any other details. United States: (only enough U.S.A. specimens to indicate the range are listed here). OKLAHOMA: Harmon Co.: Stevens 1103, near Hollis (Ny, GH). TEXAS: Baytor Co.: erodes 104 (us). Bett Co.: Wolff 992, northwest of Belton (us). Bexar Co.: Clemens 601, San Antonio (Mo, POM). BREWSTER Co.: Warnock W99, Altuda Pass, Glass Mts. (GH). Brown Co.: Palmer 26804, seein ig (mo). Cameron Co.: Hanson 8, Brownsville (Mo, NY, GH, US). . May 1900, Comanche (Mo 2). CULBERSON CO.: Waterfall. 5345, 9 miles east of Van Horn (mo, GH). Epwarps Co.: Cory 37900, Substation No. 14 (GH). Et Paso Co.: Ferris & Duncan 2369, east of Ft. Bliss on Alamagordo Road (mo, ny). Garza Co.: Ruth 1318 (us). Hipatco Co.: Camaron 109, Mc Allen (F). Hupspetu Co.: Waterfall 4966, east side of Eagle Mts. (mo, GH). Jerr Davis Co.: Hinckley 1200, Davis Mts. (F, ny). Waterfall 4727, 2 miles southwest of Chispa (GH). Kerr Co.: Heller, 12-19 June 1894, Kerrville (ariz, GH, UC, NY 2, MO). KLEBERG Co.: Sinclair 175, Kingsville (GH). Kinney Co.: Mearns 1455, Fort Clark (us, ps). MipLtanp Co.: Tracy 7815, Lake Lytle (uc). TerreLt Co.: Palmer 33510, near Feodora (Ny). ToM GREEN Co.: Tweedy 220, Dove Creek (us). Travis Co.: Taylor 3054, Austin (uc). Hall 60, Coronado R. (MO, PoM, NY, F). UvaLpe Co.: Munz 13302, 7 miles Co: DEFINITE LOCALITY: Wright, 1848 (ti MO, NY). Type-sheet of S. physocalyx STUDIES IN SIDA (MALVACEAE) 35 Gray has following: Cult., Harvard Botanic Garden, from Texas seed sent by Wright; Gregg, valley near San Pablo, Mexico; Lindheimer, \lano, west Texas; and Wright, 1848, Texas. NEW MEXICO: Cuaves Co.: Earle 352, 20 miles south of Roswell (us, Mo 2, ny). Dofia Ana Co.: Standley 3414, near Tortugas Mountain (us). Wooton 560, in the Organ Mountains (Ny, US, MO, ARIZ). WITHOUT DEFINITE LOCALITY: Wright, 1851 (Ny, us). ARIZONA: CocutsE Co.: Wolf 2543, 5 miles north of Douglas (ps, cH). Peebles, Harrison & Kearney 3514, near Ft. Huachuca (us). Pima Co.: Abrams 12678, Tucson (Ds, POM, NY). Kearney & Peebles 14971, Baboquivari Canyon, Baboquivari Mts. (GH). SANTA Cruz Co.: Beattie 4, Tubac (mo). Yavapat Co.: Purpus 57, Beaver Creek (MO, US, UC). Mexico: Sonora: Thurber 419, Fronteras (GH). Cuinuanua: LeSueur 743, Meoqui Pass (Gu, F). Stewart 675, 20 ——— northwest Jaco along road to Victoria (GH). pone ae Carretas (GH). White 2164, 3 miles north Cd. Jiminez (cH). : Johnston & Muller 852, 1 mile east Bufido (GH). Marsh 1800, Bar ee Palmer 87, Parras, 112 miles west Saltillo (Gu, us). Stewart 551, 4 kilometers west Castillon (GH). Stewart 1164, Llano de Guaje, near Tanque de La India (GH). Stewart 1766, 16 kilometers west San Guillermo GH). Nurvo Lrén: Heard, Webster & Tk 14565, 2 miles north of Sabinas Hidalgo (cH). Thurber 851, Parras Brasil: Se//o, without number or ically (us ruguay: Rosengurtt B-4710, Durazno eat Mo, us). St. Hilaire, “sur les bords des chemins dans la province Cisplatine” (TYPE, P Argentina: Bridarolli 3210, Carapari, Dept. de Oren (cH). Eyerdam & Beetle 23092, Campana, Prov. Buenos Aires (GH, uc). Hauthal, Ventana, Prov. Buenos Aires 1892 (ny). Jorgensen 1825, Catamarca, Dept. Andalgal4, El Guacho (GH). Palmer, 1853-1856, La Plata (us). Parodi 11328, Soe me Prov. de Buenos Aires (GH). Parodi 11278, Vera, Prov. de Santa Fé (F). er 4840, Dept. of Tumbaya, Prov. de Jujuy, Volcan (us). Venturi 1722, Tucum Trancas (us Bolivia: Katee 1892, Parotani (Ny). 7. Sida Brownii Clement, nom. nov. Sida physocalyx F. Muell., Fragm. 3: 3. 1862, non A. Gray, Bost. Journ. Nat. Hist. 6: 163. 1850. Erect subshrub to 4.5 dm. high, stems few, densely soft-stellate tomen- tose, the rays pale yellow, very fine and short. Leaves orbicular or ovate, subcordate at the base, 2.5—5 cm. long, to 3.2 cm. broad; the apex rounded or acute; the margins irregularly crenate; the indument dense, fulvid or canescent-tomentose; petioles densely tomentose, like the stem, terete, 1.0—1.5 mm. diam. Stipules filiform, 1.3—1.5 cm. long. Pedicels solitary in the axils, somewhat crowded in the upper leaf- axils, arti- culated above the middle, 2.4—2.7 cm. long, sharply bent at the arti- culation ca. 1 cm. below the fruiting calyx and flexuous above it, erect below. Calyx plicate, fulvid-tomentose externally, the lobes cordate, sparsely stellate-pilose internally, the tube glabrous internally; lobes con- nivent above the corolla, Petals yellow, obcordate, scarcely 6 mm. long, 36 STUDIES IN SIDA (MALVACEAE) glabrous or sparsely ciliate at the base. Stamen-tube short, glabrous. Filaments ca. 2 mm. long. Styles yellow, free nearly to the base, capil- lary, ca. 6 mm. long; stigmas terminal, depressed-capitate. Fruiting calyx ca. 2 cm. high, 6.5—7 cm. in diameter when split and flattened, penta- gonal in section, the angles strongly winged by the plicate edges of the lobes and subjacent tube, the lobes broadly cordate, 3.5 cm. broad, the whole reticulate-veined, yellow, the external indument reduced, glabrous within. Mature carpels 15—17, forming a depressed discoid fruit ca. 1.3 cm. in diameter; individual carpels 45—5 mm. radially, 3 mm. high; dorsum thick, pale brown, glabrous, coarsely verrucate on the outer surface and margins; lateral walls lighter in color and thinner; the rostrum prolonged as a sharp-pointed horizontal beak. Seed 2 mm. high and wide, red brown, smooth, dull, glabrous; the funicle greatly prolonged, 1.5 mm. long, extending 0.8 mm. beyond the seed; dehiscent apparently by the separation of the top of the dorsum from the lateral walls. Columella 2.5 mm. high, 3 mm. broad at the base, 4 mm. in diam. across the middle with acuminate flanges from the domed upper half projecting between the carpels, constricted below this and dilating again to the base; edge of the base rounded, its lower surface with many radial grooves. TYPE: “In locis rupestribus montium Hammersly Range legit N. Brown in expeditione Fr. Gregorii.” (Fragment in k). DISTRIBUTION: Western Australia. The only material which I have seen of this species has been two terminal scraps about 15 cm. and 10 cm. long, sent by Mueller to Hooker at Kew and to Gray at Harvard. The description given above has been based on Mueller’s original diagnosis, supplemented wherever possible by details from the fragments. The species must have a new name, as Gray pointed out, and it seems fitting to name it for the collector. The extraordinary inflation of the calyx in this species makes it perhaps the most striking of all the Sidas when in fruit. It does not closely resemble any other species of the genus in any of its characters, and is especially distinct in the ornate structure of the columella, the extremely slender and elongate stipules and the accrescence of the calyx. Australia: Collector unknown, “Hb. Mueller’, Gascoyne River, Western Australia (GH). Brown, Hammersley Range, Western Australia (type fragment, K). 8. Sida macrodon DC., Prodr., 1: 464. 1824. S. cymbalaria Hochr., Bull. Herb. Boiss., ser. 2, 5: 291. 1905, as to description, in part as to specs. cited. STUDIES IN SIDA (MALVACEAE) 37 S. macrodon var. cymbalaria (Hochr.) Hassl., lc. 7: 729. 1907. S. physaloides Presl, Rel. Haenk. 2: 105. 1835. Perennial herbs or subshrubs, prostrate or semidecumbent, freely branching from the base; stems (S—) 10—20 (—60) cm. long, lax, terete, the tips ascendent, 1—2 mm. diameter, the indument of minute, shining, white, simple hairs, tomentulose ca. 2 mm. long, and pulver- ulent. Leaves suborbicular to rotund-ovate, the base cordate, the apex obtuse, the margins coarsely serrate, (7—) 12—20 (+45) mm. long, (7—) 10—20 (—30) mm. wide, broadest below the middle; discolorous, gteen above, yellowish or cinereous below, appressed-tomentose on both surfaces; the younger leaves with setose-villous margins; seven-nerved, Prominent beneath; petioles terete, (3—) 8—20 (—25) mm. long, ca. 0.5 mm. diameter, about as long as the leaves or shorter. Stipules linear to linear-lanceolate, 3—4 mm. long, 0.5—1.0 mm. wide, acute, indu- ment like that of stems and more or less ciliate. Pedicels axillary, solitary or rarely paired, 2—3 cm. long, 0.5 mm. in diameter, terete, gracile, tomentellous, articulated in the upper fourth. Calyx broadly campanu- late, subpentagonal, 6—8 mm. high; the lobes erect, 3—4 mm. high, acute; the indument subtomentose, the margins short-ciliate. Petals white, rose, or salmon pink, 8—12 mm. long, 6—9 mm. wide, symme- trical or slightly oblique, rotund, emarginate, tapering quickly to the narrow claw, the margins short-ciliate. Stamen-tube 2—3.5 (—4) mm. high, terete, with ten paired traces obvious externally at the base; gla- tous or scattered-pilose (rarely heavily spreading-short-pilose). Fila- ments 1—2 mm. long. Anthers 20, pale, not dotted. Ovary 0.7 mm. high, 1 mm. in diam., eight-lobed, glabrous. Styles 8—10 mm. long, 4—5 mm. free; stigmas capitate. Fruiting calyx open, greatly inflated, the base of the tube impressed, the lobes erect, not plicate, to 1.2 cm. long. Carpels 7—9 rarely 6, muticous, united by a web around the outer dorsum, subrugulose when fully mature, indehiscent; the apex fragile, prolonged upward; dorsum with a distinct midnerve, cinereous; 2.5 mm. high, 2 mm. wide; the exocarp firmly adherent to the seed at the base, occasionally on the sides. Seed plump, glabrous, subrotund, filling all but the apex, blackish, paler near the hilum, 2 mm. in diameter. Colu- mella conical, the sides concave, angulate in section, woody, persistent, 2 mm. high. TYPE: “Mr. Stevens 1820. Bresil’” (G). DISTRIBUTION: Southern Brasil, Paraguay, northern Argentina, Bolivia, Peru. Although de Candolle described the species as having ten carpels, and the type-sheet bears a note that there are ten muticous carpels, I have seen no material which has more than nine and Rodrigo (1944) reported no more than eight. The species 38 STUDIES IN SIDA (MALVACEAE) is variable enough in its other characters, although no discrete varieties are recognizable. S. physaloides Presl is customarily placed in the synonymy of S. macrodon although it is described as having only five carpels. It was based on a Haenke collection from Huanuco, Peru, and I have seen a photograph of a probable isotype which is in the Vienna Herbarium. It is undoubtedly very close to S. macrodon, and, in the absence of any material from Peru, I am retaining it as a synonym of S. macrodon. Bang 2164 from Bolivia is nearly identical with the Haenke specimen in gross appearance. I have seen four sheets of it from U.S. herbaria, while Fries (1907) reported on duplicates in European herbaria, and we are in agreement that the material belongs to S. macrodon. The sheet in the U.S. National Herbarium has seven carpels; other sheets were too scrappy to permit carpel-counts. Hochreutiner (1905) cited three collections from Paraguay for his new species S. cymbalaria: Hassler 4325, with pink flowers, Hassler 6806, with white flowers, and Hassler 7047, which Chodat and Hassler in the same paper made the type of S. cymbalaria f. viridescens. Hassler (1907) reduced Hochreutiner’s species to a variety of S. macrodon, pointing out that Hoch- reutiner’s description was based on only one of the three collec- tions cited: Hassler 4325. The other two collections Hassler then cited as the basis for his S. rubifolia ssp. pseudocymbalaria of section Sida. The differences which Hochreutiner claimed existed between his S$. cymbalaria and other species of the section are actually no more than those often found between individuals of S. macrodon. S. macrodon var. juidrmepes Griseb. (1874) is a synonym of S. flavescens Cav. Brasil: Archer 4143, Bututan Séo Paulo (us). Bornmiiller 595, Panambi (Neu Wiirttemberg), Rio Grande do Sul (Gu). Burchell AA18-2, without locality (cH). Hatschbach 2697, Rio do Salto, Parana (us). Hoehne & Gehrt 36563, Suzanno, Sao Paulo (us). Lund, near Lagoa Santa (us). Novaes 490, Campinas, Jundiahy (us). Sello, without locality (us). Stevens, 1820, without locality (Photograph of type, G: F, US). Warming, Lagoa Santa (us). Widgren, 1845, Minas Geraes (us). Paraguay: Hassler 9377, near Caaguazi (Gu). Hassler 5085, Sierra de Maracayu (Gu). Hauthal 38, Rio Ibitimi (ny). Argentina: Ekman 133, 1908, “Posadas, Bonpland” (ny). Bolivia: Bang 2164, “Yungas, Coripati’’, La Paz (GH, MO, US 2). STUDIES IN SIDA (MALVACEAE) 39 Peru: Haenke, “ad Huanacco’ (Photograph of Isotype? of S. physaloides Pres], Vienna: F 9. Sida urticaefolia St. Hil., Fl. Bras. Merid. 1: 148, t. 37. 1827. S. flavescens Cav. sensu Schum., Mart. Fl. Bras. 12, pt. 3: 290. 1891. Erect perennial, to 40 cm. high, the stems few-branched, covered with a double indument of dense, minute, stellate pubescence and scattered- spreading, pilose-stellate hairs, the rays ca. 1 mm. long. Leaves 3.5—7.0 cm. long, 2.0—4.3 cm. wide, nerves prominent on the lower surface, the lower suborbicular and obtuse the upper oblong-ovate, often oblique, the base truncate to shallowly cordate, the apex obtuse to acute, the margins coarsely serrate, the serrations irregular; the upper surface scabrid with a thick scattering of appressed, yellowish or golden, 5—7- rayed, fine, stellate hairs; the lower surface densely covered with minute, stellate, cinereous or yellowish hairs and slightly larger, many-rayed, yel- low, stellate hairs, the bases of the rays often bulbous; the petioles 1.4—2.3 cm. long, terete, the indument like that of the upper portions of the stem, ascending, straight, to 1.0 mm. thick. Stipules 4—7 mm. long, 0.7—1 mm. wide, lanceolate or falcate, entire or with one or two blunt lobes on the inner curve, the upper ones occasionally linear, al- ways acuminate. Pedicels 4—16 in the axils, rarely fewer in the lower axils, 2—12 mm. long, mostly ca. 7 mm. long, slender, the indument like that of the upper stems but the long-rayed stellate-pilose hairs more numerous. Calyx campanulate, densely stellate-hirtellous to pilose, strongly 5-angled, ca. 4 mm. high in early anthesis, rapidly accrescent to ca. 9 mm, in late anthesis; the lobes changing shape and dimension during the accrescence, from broadly deltoid to cordate, mucronate, ca. 2 mm. wide at the base, elongating from 1 to 2 or more mm.; interior markedly reticulate, shining, the tube glabrous, the margins finely stellate-hirtellous. Petals white to salmon pink or rose, strongly oblique- Ovate, the margins somewhat irregular, the apex often retuse, the mar- gins with regularly scattered short-ciliate hairs, increasingly dense to- wards the claw; the claw ca. 1.5 mm. wide. Stamen-column 2.0—2.5 mm. long, terete, glabrous, thick, the lower third dilated; filaments 1—1.5 mm. long, slender, not paired; anthers 20—30, verruculate, not dotted, ca. 0.5 mm. wide; the valves usually not reflexing in age, the septum prominent. Styles five, ca. 4 mm. long, 3 mm. free; the stigmas depressed-capitate, dark red. Ovary 1 mm. high and wide, light brown, five-lobed, ovate, glabrous. Fruiting calyx 7—8 mm. long; the lobes 3—3.5 mm. long, 3.5—4 mm. wide; reticulated, membranous, translu- cent, grayish-tan, persistent after the fall of the fruits, rarely abscissing at the inconspicuous articulation ca. 3 mm. below the calyx. Carpels 40 STUDIES IN SIDA (MALVACEAE) five, obtuse, muticous, dark reddish-brown, ca. 2 mm. high by 1.8 mm. wide, the entire dorsum convex, reticulate, with a distinct midnerve, the margins with the lateral walls slightly ridged; lateral walls membra- nous. Seeds glabrous, black, nearly spherical, ca. 1.5 mm. in diameter. TYPE: St. Hilaire, C2/2683, Feb. 1821, “Paturages de la province des Missiones, prés du village de S. Borga”, western Rio Grande do Sul state, Brasil (P). DISTRIBUTION: Southern Brasil, Paraguay, and northern Argentina. Study of St. Hilaire’s type and of a photograph of material he sent to Berlin (labelled as Isotype) has given me some certainty about the application of the name to the cited material, despite uncertainties resulting from a study of his description alone. Schumann confused this species with S. flavescens Cav., and some later authors have followed his example. Rodrigo’s key (1944), unfortunately, does not sufficiently emphasize the diagnostic characters of the plant, although her description is quite clear; she makes a main key-heading of the habit of the plant, a character which is not easy to recognize on most specimens. The map of collectors’ travels in vol. 1 of Martius’ Flora Brasiliensis shows St. Hilaire’s route extending into what is now the Gobernacién de Misiones of Argentina. It is clear from St. Hilaire’s own account (1946) that he collected only on the Brasilian side of the Uruguay River in an area around San Borja once under the influence of the Missions of Paraguay, seven of which were on that side of the river. Brasil: Bornmiiller 183 and 423, near Panambi (Neu Wiirttemberg), Rio Grande do Sul (cu). Rambo 51611, Passo do Socorro, Rio Grande do Sul (us). St. Hilaire “Brasil (photo. of fragment sent 2 him to Kunth, B: Gu, vs). St. Hilaire, near San Borja, Rio Grande do Sul TYPE Paraguay: Hassler 6943, Cordillera Central, near the Y- -aca (GH, US). Kunize, Sept. 1892. “Siid-Paraguay”’ ; ntina: Kuntze, Dec. 1891, Dique, near Cérdoba (nx). Montes 3376, Puerto Rico, Dep. Cainguas, Gob. Misiones (us). Parodi 11183, Reconquista, Prov. Santa Fé. (GH). Rodrigo 2498, Enrique Urion, Dept. Tapenaga (Ny). Schwindt 916, Cainguas, Gob. Misiones (us). 10. Sida calyxhymenia J. Gay ex DC., Prodromus 1: 462. 1824. Perennial shrub 6—15 dm. high, the stems erect, terete, branching, densely stellate- incanous-tomentose, the branches numerous and crowd- mm. wide, the upper leaves narrower in proportion, 8—12 mm. long, ca. 2 mm. wide, rounded at the base, obtuse to subacute at che apex, STUDIES IN SIDA (MALVACEAE) 41 the margins repand-undulate to subentire, flavescent from the dense short-stellate tomentum, above, velutinous-incanous beneath and _ less dense; three-nerved from the base, the midnerve lightly impressed above, prominent beneath, the petioles terete, their indument like that of the stems, 6—8 mm. long, the upper ones shorter. Stipules subulate, stellate- tomentose, glabrescent towards the apex, deciduous, 2—3 mm. lon 8 Pedicels solitary and axillary or rarely paired, terete, short and erect in flower, elongating later and flexuous, densely incanous-tomentose, 12— 16 mm, long, articulated 2—4 mm. above the base in fruit, not visible beneath the tomentum. Calyx 6—8 mm. high in anthesis; lobes 4—5 mm. high, 4—5 mm. wide slightly below the middle; cordate in out- line, acute, greatly accrescent, flavid-velutinous; lower margins of the lobes plicate; sinuses closed; accrescent to 1 cm. in height, becoming membranous; sapeeae! veined, with three main veins per lobe. Petals erect, ca. 6 mm. high and wide, obovate, the claw ca. 0.8 mm. wide, dens- ely white- flexuous- fimbriate, the margins with scattered very slender cilia, the remainder glabrous. Stamen-tube glabrous, with 5 conspicuous pairs of traces, slender, tapering, ca. 4 mm. high; filaments 1.5—2.0 mm. long, filiform; anthers 10—20, ferrugineous, not dotted, reniform- globose, ca. 0.5 mm. wide. Styles five, 6 mm. long, 2—3 mm. free re- flexed, dark; stigmas capitellate, dark red. Carpels five, 3 mm. high, 1.5 mm. radially; the rostrum prolonged vertically, ca. 1.5 mm. long; dorsum wrinkled, scattered-puberulent, transversely impressed in the hilar de- pression of the seed; lateral walls rugose-reticulate, paler; dehiscence along the midline of the dorsum and the inner margin. Seed glabrous, 2—3 mm. high, dark reddish-brown, the hilar depression rather deep, the rostrum prolonged briefly upward; funicle flattened. Columella ca. 3 mm. high, crowned with the remnant style-base, 5-angled; the angles subalate; the base dilated into a very small crispate disk. KEY TO THE VARIETIES Plants cinereous-tomentose; calyx flavid- Lesan Para leaves 23—43 mm. lon yxhymenia vat. calyxhymenta. ng . Plants _ferrugineous-tomentos calyx ¢ tomentose a bit leaves shorter ee aes 10b. S. calyxhymenta var. ferruginea. 10a. Sida calyxhymenia var. calyxhymenia. Fleischeria pubens Steudel in Lehmann, Plantae Preissianae 1: 236. 1845 and emend. Steetz, op. cit. 2: 365. 1848. Type: Leschenault et al. 1795, “cueillie 4 la Riviere des Cygnes, sur la céte occidentale de la heeclle Hollande” (K). DISTRIBUTION: Western Australia. 42 STUDIES IN SIDA (MALVACEAE) 10b. Sida calyxhymenia var. ferruginea Pritzel, Engl. Bot. Jahrb. 35: 362. 1904. COTYPES: George 1902, “e. gr. in. distr. ae pr. Murrin- murrin” and Diels 3269, “pr. Cue, fl. m. DISTRIBUTION: Known only from the sah The variety is known to me only from the description, there being no material of the species which I have examined which can accurately be termed “‘ferrugineous” in the color of the indument. Domin has tentatively identified an E. Clement collec- tion (Kk) from between the Ashburton and Yule rivers of Western Australia as possibly belonging to this variety. While it has a reddish cast and leaves shorter than the typical, it cannot be identified certainly since it has no fruit. In addition, the stamen- column is shorter and not so markedly nerved, although the petals are very similar to those of the typical. Fleischeria was established by Steudel (1845) as a new genus, with the single species F. pubens, distinct from Sida in having a “petaloid” calyx and the corolla shorter than the calyx. It was Steetz’s opinion, expressed at the time he emended the generic and specific diagnoses (1848), that Sida was too vast and diverse, and that since F. pubens was so markedly distinct from all the other Australian species which might be placed in Sida, it was best to treat it as generically distinct. There is no indication that either Steudel or Steetz recognized the conspecificity of F. pubens with S. calyxhymenia. Oddly, the name Fleischeria had been used in 1838 by Steudel and Hochstetter in Endlicher’s Genera Plantarum for species usually referred to Scorzonera of the Compositae. Australia: ES ahesitey 02 Blake 19195, Mt. Howitt Station, Gregory South District (k). WesTreRN AustraLia: Burbidge 265, Glenorn Station, Malcolm (x). Clement, n.w. Australia, between the Ashburton and Yule Rivers (x) Cl n.w. Australia, betwe the Ashburton and Gray Rivers (Kk) Drummond, 1839, Swan River (Kk 2). Drummond 52, 1843, Swan River (Kk). n. of Broken Hill (x). Oldfield, Murchison River (Gu, K). Preiss wae Mount Mathilde, York District (Isotype of Fleische eria pubens Steud. kK). SOUTHERN AustrRALia: Holms, “Arco-cillinna Well.” (xk). WrirnHour DEFINITE LOCALITY: Anketell, “Trans Australian Railway Survey, 4-8-1910" (x). Amketell, “Trans Australian Railway Survey, 12-8-1901"" (x). STUDIES IN SIDA (MALVACEAE) 43 11. Sida Standleyi Clement, sp. nov. Herbae perennes prostratae 40 cm. vel minores; folia 1.3—3.4 cm. long., 1.4—2.8 cm. lat., plurima ovata caetera suborbicularia, apice acuta, basi cordata, marginibus grosse serratis. Petioli 1.0—1.8 cm. Stipulae filiformes sive lineari-lanceolatae. Pedicelli 5—20 mm., filiformes arti- culati prope apicem bini et in axillis superioribus soli. Calyx accrescens conspicue 5-gonus, lobis acuminatis nervo expresso. Petala ca. 5 mm., rotunda gibbosa, margine exteriore subretuso. Columna staminales teres, pilis subulatis patentibus sparsesque munita. Antherae 20. Styli 3.5 mm., 5 in ovario, Calyx marcescens, aetate fructus maturandi membranaceus et ad 1.0 cm. alt. accrescens. Carpella 5 in fructu, 2.5—2.8 mm. alt., ca. 1.7 mm. lat. secundum radium, a dorso convexa glabra, parietibus Jater- alibus planis albo-pulverulentibus. Semen cum parietibus adnatum nisi ad aspicem. Perennial prostrate herb. Stems 20—40 cm. long, slender, 1—2 mm. in diam., terete, fibrous-woody near the base, densely clothed in minute, greenish, soft, many-rayed, stellate hairs, appressed, and scattered longer- rayed stellate hairs, the rays yellowish, spreading, and simple, spreading, pilose hairs. Leaves 1.3—5.4 cm. long, 1.4—2.8 cm. wide, mostly ovate; some of the lower leaves suborbicular; the base cordate, symmetrical; the apex acute; nerves 7—9, palmate from the base, the midnerve with 2—+3 lateral nerves subopposite or alternate, all prominent beneath, yel- lowish; the margins coarsely serrate, the serrations ovate, acute or blunt- ly mucronate; indument of the upper surface minutely stellate-toment- ellous, somewhat scabrous; that of the lower surface similar, the hairs softer, slightly longer-rayed, denser. Petioles 1.0—1.8 cm. long, 0.5—0.8 mm. diam., terete, spreading or ascendent; indument like that of the upper stem, often with more numerous, long, pilose hairs. Pedicels filiform, 5—20 mm. long, articulated 1.5—4 mm. below the calyx, soli- tary or usually paired in the axils. Calyx accrescent, strongly 5-angled, ca. 7 mm. high; the base of the tube impressed; the lobes acuminate, with a prominent mid-nerve, 3 mm. high, 4 mm. wide, fine, white, stellate-hirtellous outside, the same inside but the lower part of the tube glabrous. Petals 5.0 mm. long and broad, rotund, gibbous; the outer margin slightly retuse; the claw ca. 0.6 mm. wide, densely ciliate, the rest of the petal glabrous, 5—6-veined. Stamen-tube 1.5 mm. high from the base of petal to level of filaments, terete, with scattered, spreading, hyaline, subulate hairs; the interior with 5 pairs of traces; filaments ca. 8 mm. long, slender, flexuous. Anthers 20, ca. 0.3 mm. wide, the sur- face subpapillose. Styles five, 2.8 mm. free, 3.5 mm. long; stigmas de- pressed-capitate. Ovary 0.5 mm. high and wide, ovate, pale, glabrous, strongly five-lobed. Fruiting calyx tan, membranous, to 1 cm. high; in- dument persistent. Carpels five, 2.5—2.8 mm. high, 1.7 mm. radially; 44 STUDIES IN SIDA (MALVACEAE) the dorsum convex, light brown, glabrous, rough, dull, with an impress- ed midline; lateral walls flat, ca. 1 mm. wide, as high as the carpel, white-pulverulent, indehiscent; the walls adnate to the seed except at the apex. Seed dark red-brown, glabrous, paler near the hilum; the funicle prominent, reddish. Columella 2 mm. high, the lower fourth dilated, woody, ‘engeuite, truncate. TYPE: H. H. Bartlett 10167, “Cerro de los Armadillos, vicinity of San José, Sierra de San Carlos, Tamaulipas, Mexico. 3100 ft. 8 July 1930” (F). DISTRIBUTION: Mexico: Sierra de San Carlos, Tamaulipas and Sierra Madre, nr. Monterrey, Nuevo Leén, S. Standleyi is closely allied to S. flavescens of Argentina, Paraguay and Uruguay, the carpels being almost indistinguish- able, although the petal-size and stamen-number are quite different. It is curious that S. physocalyx, also of section Physalodes, has a distribution similar to that of S$. Standleyi and S. flavescens combined. The specimens cited were given an herbarium name by Dr. Paul C. Standley, but I think it more appropriate to name the species for him. Mexico: ats Bartlett 10167, Cerro de los Armadillos, vic. of San José, (TYPE, F). oO Leén: Mueller & Mueller 79, Diente Canyon, Sierra Madre, nr. ae ie MEXU 2). 12. Sida flavescens Cav., Diss. 1: 14, t. 13, f. q. 1785. S: ‘prostrata Cay, op, cit. 12°13, te: 13;-t2:3: S. intermedia St. Hil, Fl. Bras. Mer. 1: 188. 1827. S. macrodon var. ttermedia (St. Hil.) Griseb., Pl. Lorentz. 43. 1874. S. prostrata var. flavescens Bak. f., Jour. Bot. 30: 294. 1892. S. prostrata var. genuina Hassl., bee S51. 1920 S. prostrata var. flavescens Hassl., Prostrate perennial herbs, many- co fatten freely lamang deep, woody taproot; the stems 25—40 cm. long, terete, 1—2 mm. in diam., densely stellate-velutinous, the hairs minute, whitish or earothithe and with scattered to dense, longer, slender-rayed, spreading or somewhat appressed, white or flavid hairs. Leaves 1.5—3.2 cm. long, 1.0—3.1 cm. wide, cordate, sometimes oblique; the apex subacute or acute; the mar- gins irregularly or fairly regularly serrate, the serrations obtuse, 2.5—6 mm. wide at the base; indument of the upper surface greenish, densely stellate-puberulent; that of the under surface the same, plus scattered to STUDIES IN SIDA (MALVACEAE) 45 dense longer-rayed stellate hairs, the rays flavid, slender, soft; nerves 7—9, palmate from the base, prominent beneath; petioles 10—15 mm long, usually about half as long as their leaves, channeled adaxially; the indument like that of the stem. Stipules to 5 mm. long, 1 mm. wide, linear, lanceolate, or falcate, occasionally with a small lateral lobe, vary- ing even within one pair; indument like that of the stems. Pedicels axillary; solitary, paired or in threes, usually paired, 5—8 mm. long, articulated 1—3 mm. below the calyx. Calyx five-angled, the tube round- ed at the base; the sides and lobes parallel; 3—4 mm. high during anthesis, accrescent to 7—9 mm. in fruit; external indument greenish to flavid, thickly to densely stellate-puberulent, with longer-rayed stellate hairs on the prominent midnerves; the lobes erect, ovate, acuminate, ca. 2 mm. high in flower, to 5 mm. high in fruit. Petals rose, 8—9 mm. long, 5—7 (—9) mm. wide, obliquely obovate-retuse, the margins short-ciliate all around; the cilia longer and denser towards the base; claw ca. 2 mm. wide. Stamen-column ca. 1.5 mm. high, terete, stout, glabrous or with few,’ short, scattered bristles; filaments ca. 1 mm. Jong, not paired, flexuous; anthers 30—40, ca. 0.5 mm. wide, not dotted. Ovary five-lobed, truncate-pyramidal, ca. 1 mm. high and wide, glabrous, light brown. Styles 3.2—4.5 mm. long, 3.5 mm. free; stigmas capitate. Fruiting calyx pale tan, membranous-reticulated; the indument persist- ent. Carpels five, muticous, light brown, forming a pyriform fruit, 3 mm. high, 2 mm. radially, glabrous; the lateral walls flattened, white- pulverulent; midline of the dorsum visible only near the apex; partially dehiscent by the inner margin, the walls separating with difficulty from the base and lower dorsum. Seed reddish brown, the surface dull, densely and minutely pitted, 2 mm. high, 1.5 mm. radially; funicle lighter in color, corded. Columella 2 mm. high, tapering slightly, finely ribbed, pale; the apex irregular, crowned by the short persistent style-base. TYPE: Commerson, “in rupibus Montevideo”, Uruguay. (P). DISTRIBUTION: Southern Brasil, Uruguay, northern and central Argen- tina, perhaps Bolivia and Chile. S. flavescens Cav. and S. prostrata Cav. were first combined by Willdenow (1801) in the following words, extracted from the intercalated description, “Sida flavescens .... S. (prostrata) .... Cav. Diss. 1 p. 13.... B S. (flavescens) .... Cav. Diss. 1. p. 4....”. De Candolle (1824) gave Willdenow as the author of the combined species, citing both of Cavanilles’ species in the synonymy. Under the present rules, Willdenow’s combination of the elements under the name S. flavescens must be followed. As is evident from the synonymy, later authors have apparently gone on the basis of page priority or have followed Garcke (1896). 46 STUDIES IN SIDA (MALVACEAE) Schumann, however, used S. flavescens, with a synonym-citation including S. prostrata and referring to Willdenow and de Can- dolle. He also included S. urticaefolia in the species, although it is quite distinct. The types of both of Cavanilles’ species were Commerson collections from Montevideo, both originally sent to Thouin. That of S. prostrata was lent by de Jussieu to Cavanilles for study, that of S. flavescens by Thouin. The specimens in the Madrid herbarium, photographs of which are reproduced in Rodrigo’s paper (1944), are certainly not the types, that ostensibly of S. prostrata being a Née collection from Chile, although accompanied by a drawing which closely resembles that of the Dissertationes, while that of S. flavescens gives no clue as to its origin and is accompanied by a sheet bearing a handwritten diagnosis, in what looks like Cavanilles’ hand, which is worded quite differently from the published diagnosis. There can be no doubt that the actual types are in Paris, where the bulk of the Commerson and half the Dombey collections were sent. I have - studied the types of both S. flavescens Cav. and S. prostrata Cav. and the brief descriptions on the original labels are repeated in, Cavanilles’ published description and attributed to Commerson. Cavanilles apparently took some material to Madrid with him when he left Paris in 1789, and distributed duplicates to Berlin, Copenhagen and other herbaria, but the types themselves, usually loaned to Cavanilles from the herbaria of such men as de Jussieu, Thouin, and de Lamarck or grown from seed in the Jardin des Plantes, remained in Paris. The type of St. Hilaire’s S. intermedia, so named because it was supposedly intermediate between S$. macrodon and S. urticae- folia, was collected by its author from fields near the Salto Grande in Uruguay. It is a small plant which is a ratoon or sprout-growth from an old base, and there can be no doubt that it is conspecific with S. flavescens. Baker’s treatment of this species-complex was perhaps the most original, for he took up the name S. prostrata for his assemblage of entities, with S$. urticaefolia as an outright synonym of the typical element, and with S. flavescens reduced to the rank of a variety. The distribution of the species given above is based in part on specimens seen, which are from Uruguay and Argentina only, in STUDIES IN SIDA (MALVACEAE) 47 part on that given by Rodrigo (1944). This seems reasonably safe, since her concept of the species is the same as mine, despite the difference of name. Uruguay: Commerson, Montevideo (type, Pp; fragment, F). Commerson, Kesslooties (Photograph of probable Isotype, in Madrid: Fr, us). Commerson, Montevideo (type of S. prostrata Cav., Pp). Herter 96563, Biller: Dept. San José (mo). Kuntze, Nov. 1892, Sierra de Locy (F, Ny). Rosengurit B 467, Rio Yi and Matanzas, Dept. Flores (Gu). Rosengurtt B 725, Cerro Colorado, Dept. Florida (us). Rosengurtt et al. PE 596 1/2, Juan Jackson, Dept. Soriano (us). St. Hilaire, “Paturages prés de Salto Grande” Uruguay (type of S. intermedia; P). Argentina: Bartlett 20183, Rosario, Sierra de Cérdoba, Prov. Cérdoba (us). Krapovickas 2919, Tandil, Prov. Buenos Aires (mo). Kuntze, Dec. 1891, Cérdoba (ny). Montero 522, Loreto, Dep. Candelaria, Gob. Misiones (us) Pedersen 1221, Est. Herradura, Dep. Boca del Bermejo, Gob. Formosa (us). WirHouT DEFINITE LocaLity: Née; “Chile” (Photograph of specimen in p: F). Née, “Colonia de Sacramento’ (Fragment from Madrid: F). III. Section PssuDOoMALVAsTRUM Gray, Mem. Am. Acad. 4: 23, 1849 (Pl. Fendl.). Low or prostrate herbs or subshrubs. Leaves usually oblique, reniform to elongate-triangular. Indument of scattered to dense lepidote scales or appressed stellate hairs. Peduncles commonly recurving in fruit, mostly solitary in the axils. Epicalyx present, of 1—3 filiform to setaceous bracteoles; calyx terete. Carpels tumescent, the dorsum stellate-puber- ulent to hirsute, indehiscent or dehiscent irregularly along the mid- dorsal line. Funicle curved over the rostrum of the seed, usually dilated towards the columella. The section was originally described by Gray for only two species, S. hederacea and S. sulphurea, with the comment “S. sulphurea .. .. from Mendoza [Argentina] is a nearly related species [to $. hederacea]|; and some others from the same region usually referred to Malva probably belong here.” There is a marginal note in his hand on the type- -sheet of S. hederacea (GH) © “cf. Malva leprosa Orteg. ex Hook.”, but that species was not transferred into the genus or the secon until 1890, when chumann made the transfer in his discussion of the section. Meanwhile, in 1852 Gray had described S. lepidota and its varieties depauperata and sagittaefolia, commenting that S. le pi- ota was “a well-marked though somewhat polymorphous species of the Section Pseudomalvastrum.” In Schumann’s treatment of the section mentioned above, he recognized only one other species, S. Sherardiana, as belonging 48 STUDIES IN SIDA (MALVACEAE) to the section, apparently considering S. hederacea and S. lepidota as subspecific entities under S. leprosa, without, however, mentioning either. Two years later Baker reshuffled the names and entities in the section, selecting S. hederacea as the species-name for the leprosa- hederacea complex, and adding S. cuneifolia Gray to the section, although it has no bracteoles and is sufficiently unlike the rest of the section to form a new section with S. Helleri. Hoch- reutiner (1902) attempted to unscramble the nomenclatorial snarl created by Baker with only partial success, falling into the trap of following Baker’s acceptance of S. hederacea var.? parvifolia Hemsley as legitimate, although it was published without descrip- tion, and transferring it to S. /eprosa as a still illegitimate variety. In 1906 Greene transferred S. cuneifolia Gray, S. hederacea, S. lepidota and S. lepidota var. sagittaefolia to the new genus Disella. The genus was based on the presence of an epicalyx, despite the fact that the first-named species never has one. Very few botanists have accepted Disella. Kearney (1954) includes the section in his key, lists and keys out S. hederacea, S. lepidota, S. Grayana and S. Helleri, and comments in his notes that the section is heterogeneous and poorly defined. Type species: Sida leprosa (Ort.) K. Schum. KEY TO THE SPECIES Leaves mostly symmetrical, reniform, with a deep and narrow sinus. Caf heavily hirsute, with a pr oe rostrum noone over the apex of the columella 13. S. Sherardiana. Leaves reniioan' to sbrnieagctate). esualy ‘ophee chen un so. Carpels moderately hirsute or hirtellous or puberulent, the rostrum not prolonged, never more than slightly hooked over the apex of the columella 14. S. leprosa. 13. Sida Sherardiana (L.) Benth., Jour. Linn. Soc. 6: 101. 1862. Malva Scherardiana L., Sp. Pl. ed. 2, 1675. 1763. -Malva Sherardiana L., Syst. ed. 12 and later. Malva cymbalarifolia Desr. in Lam., Encyc. 3: 753. 1791. Malva cymbalariaefolia Desr. (orth. var. in DC. Prod. 1: 431, 1824). Malvella Sherardiana (L.) Jaub. & Spach, Illustr. 5: 47. 1853—57. STUDIES IN SIDA (MALVACEAE) 49 Malvella cymbalariaefolia (Desr.) B. D. Jackson, Index Kew. 3: 156. 1894, erroneously attrib. to Boiss. Fl. Or. 1: 835. 1867. Prostrate or decumbent perennial herbs. Stems numerous from a woody taproot, the entire plant covered with a dense, soft, whitish, long-rayed, stellate indument. Stipules oblong or ovate, acutish, 2—3 mm. long, ca. 0.8 mm. wide, persistent, yellowing in age, spreading. Petioles to 27 mm. long, subcomplanate, assurgent. Leaves reniform, variably crenate or crenate-serrate except in the deep rather narrow sinus, (10O—) 12—25 (—28) mm. wide, (S—) 9—14 (—23) mm. long; indument occasionally thin on the upper surface of large-leaved speci- mens; nerves 6—7, palmate, prominent beneath. Pedicels solitary in the axils, slender, sinuate, terete, not articulated, 10—5O mm. Jong, ac- crescent after anthesis; indument double, of long- and short-rayed stellate hairs. Calyx subcampanulate, plicate in specimens with ovate lobes, in- vaginated around the receptacle in fruit, 6—8 mm. high; lobes 3—5 mm. long, 3—5 mm. wide, broadly lanceolate or ovate, acute, with four main nerves on each, folding over the fruit, somewhat accrescent after anthesis; inner surface glabrous at the base, finely pubescent on the lobes, not dotted, Epicalyx of 2—3 thin, lanceolate, spreading bracteoles ca. 2 mm. long. Petals rose, pink to yellowish-brown on drying, broadly spatulate, obtuse, 5—8 mm. wide, 7—12 mm. long, sparsely hirsute near the claw, inserted slightly above the base of the column. Stamen- tube 2 mm. high from the base of the ovary to the level of the fila- ments, short-obpyriform, sparsely hirsute, strongly geniculate at the base; filaments ca. 2 mm. long, not paired. Anthers numerous, ca. 0.8 mm. wide when open, pale, not dotted. Ovary ca. 1 mm. high and wide, densely hirsute. Styles deeply inserted between the apices of carpels, free almost to the base, connate, subclavate apically, S—6 mm. high; stigmas strongly oblique, obtuse, only slightly thicker than the subjacent portion of the styles. Carpels 7—16, 3—4 mm. high, 4—} mm. radially, forming a flat-topped fruit; dorsum pale yellowish-brown, muticous, inflated; rostrum prolonged into a vertical flange hooking downward to the top of the very short columella; lateral walls nearly white, rugose- reticulate. Seeds plump, grayish, glabrous, minutely verruculate, 2 mm. high and radially; funicle thin, flat; hilar depression shallow. TYPE: Specimen in Linnaean Herbarium, listed in 1767 enumeration. DISTRIBUTION: Central Spain, Greece, Asia Minor and the Caucasus. The species was named by Linnaeus for John Sherard, an English botanist of his time, but was misspelled in the original publication. In Linnaeus’ later works, however, the name was consistently spelled as it is used here. e€ species was first transferred from Malva by Jaubert and . 50 STUDIES IN SIDA (MALVACEAE) Spach as the basis for a new genus Malvella, distinguished primarily by the characters of the carpel; the genus has found little acceptance, whereas Bentham’s transferral of the species to Sida, because of the pendulous ovule and despite the presence of bracteoles, has been generally considered as correct. S. Sherar- diana is closely allied to the S. leprosa-hederacea complex of the western hemisphere in the inflation of its carpels and in general features of leaf-shape, indument and habit. It is the only species of the genus native in Europe. Spain: Bourgeau, Cerro Negro near Madrid (Gu, uc). Graelsius, Cerro Negro near Madrid (GH). Greece: “Triv.”, “Macedonia’” (Mo). Turkey: Sintenis 677, Dardanelles (GH). U.S.S.R.: Bohenacker, May 1836, “Cauc. ...... prope Sheki Geor.” (Mo). Newodowakij, 12 June 1911, Prov. Tiflie, Karajazy (cu). Turkey in Asia: Balansa 724, “Village de Tchaousli, prés de Mersina (Cilicia)”’ (uc). Hildreich, Oct. 1845, near Timboukchan (Gu). Pichler, Sept. 1875, “circa Brufsa” (mo). Szovits, “Armenia” (us, Gu, Ny). Syria: Boissier, June 1846, near Aleppo (uc). Kotschy?, near Ain el Ombarek in vicinity Aleppo (mo 2). Post 460, Ramoth (us). Ssvtenis 1418, Kiredjik near Zambur (mo). Palestine: E/g & Grizi 357, el-Qubab (GH). Meyers 790, Jerusalem (Fr). Without collector or date, Arimathea (Mo). Iraq: Rolland 114, “Mesopotamia” (cH). 14. Sida leprosa (Ort.) K. Schum., ns & Prantl, Nat. Pflf. 3, Abt. 6: 43. 1890 Low perennial herbs from branching woody roots. Stems 5—40 cm. long, often assurgent at the tips, woody, terete basally, complanate or terete above, densely clothed with silvery, lepidote to free-rayed .or appressed, stellate hairs; branches few to many from any part of the axis. Stipules lanceolate, linear-subulate or subfalcate; indument like that of the stem; 1—2 mm. long, to 0.5 mm. wide, deciduous, appressed. Petioles 4—30 mm. long, channeled on the adaxial surface; indument like that of the stem, dense. Leaves extremely polymorphic, reniform to ovate-subcordate with the margins nearly entire to serrate, or obliquely triangular and semicordate, with the margins irregular, or narrowly trfi- angular with the upper margins entire and the semisagittate base laci- niate; leaves 8—34 mm. long, 2—50 mm. wide; indument variable, like that of the stem, scattered to dense, scabrous, denser on the younger leaves; nerves mostly 5, palmate from the base, prominent beneath, usually paler than the surface. Pedicels solitary in the axils or occasion- ally from the axils of much shortened branchlets, terete or subcompla- nate, flexuous to recurved, 10—50 mm. long, not noticeably articulated; indument like that of the stem, dense, the hairs silvery or yellowish or cinereous. Epicalyx of (O—) 1—3 filiform to subulate, appressed or spreading bracteoles 1—3 mm. long, often hidden in the indument of STUDIES IN SIDA (MALVACEAE) 51 the calyx, deciduous, occasionally at the apex of the pedicel, usually on the base of the calyx; indument like that of the calyx. Calyx campanu- late, terete, 3—8 mm. high, deeply divided into deltoid or ovate, acute or acuminate lobes 2—6 mm. long; indument dense, like that of the stem, the interior glabrous except the lobes. Petals 10—16 mm. long, 7—11 mm. wide, white to rose to sulfur yellow, varying within one colony at times, often with the portion exposed in bud more deeply colored and with scattered to dense, minute, stellate hairs, obliquely obovate, the inner margin often auriculate, the outer margin concave, the apex obtuse, rounded; veins fine, numerous; claw glabrescent to softly ciliate, inserted on the stamen-tube above the level of the ovary. Stamen-tube 3—4.5 mm. high, glabrous, 10-striate, somewhat 5-ribbed, ca. 2 mm. diam. at the claws; filaments numerous, crowded, falsely multiseriate, often paired, 1—4 long, recurved in anthesis. Anthers 40—50, ca. 0.6 mm. wide when open, not dotted. Styles erect, 10—12 mm. long, slightly clavate at the apex; stigmas capitellate, sometimes oblique. Ovary densely stellate-puberulent, occupying about two-thirds of the cavity, ca. 1 mm. high, 1.5 mm. diam. Carpels 6—10 (rarely more, and the additional ones usually aborted), muticous, tumescent, 2.7—3.2 mm. high, indehiscent, or dehiscent irregularly along the mid- dorsal line; dorsum dull red-brown, moderately to densely stellate-pu- berulent; lateral walls membranaceous to thick, smooth, straw-colored, white-pulverulent; outer margins sporadically umbonate; outer dorsum tumescent or somewhat invaginated; apex nearly horizontal or some- what ascendent; inner margin vertical or somewhat concave. Seed 2.2— 2.5 mm. high, ca. 2 mm. radially, dark red-brown, plump, with scattered puberulence; funicle paler, terete inwardly, prominent; hilar depression shallow, abrupt. Columella variable, slender to broadly conical, truncate, ca. 2 mm. high, flanged, pale straw-colored. KEY TO THE VARIETIES A. Leaves reniform to broadly and eed ovate and acute, mostly broader than Jong; indument stellate or lepido’ pimp rarely ‘longer; cnlyx Said mm. high; flowers sulfur ae ethane tinged with rose 14a. S. leprosa var. leprosa. lee} — 5 a. £ B 5 ® 5 4 a 4s) o. 4 g < o un“ io) B. tndavune siéllate, ieee to 40 mm. “beg: tasely less than 10 mm.; calyx 4 . hi sh; flowers cream-colored or light yellow, often tinged with red, rarel 14b a var. hederacea. A. Leaves obliquely cringe aod acute, =, mostly ae than broad; indument lepi # C. Leaves delt and: procter shictly 10—30 mm. bisa he poet iilegins irregularly notched and toothe 14c. S. leprosa var. ee C. Leaves narrowly elongate-triangular, sem isagittate, mostly 2 0 mm. long, angular the upper margins entire and sttaiahe 14d. S. leprosa var. Pamuioue oF STUDIES IN SIDA (MALVACEAE) 14a. Sida leprosa var. leprosa. Malva leprosa Ortega, Dec. 8: 95. 1798. Malva sulphurea Gillies, Hook. Bot. Misc. 3: 149. 1833. S. sulphurea (Gillies) Gray, Mem. Am. Acad. 4: 23. 1849. Malvastrum sulphureum (Gillies) Griseb., Symbol. Fl. Argent. 43. 1879. S. hederacea var.? parvifolia Hemsl., Biol. Centr. Amer. 1: 104. 1879. (nom. nud.). S. hederacea var. saipearen (Gillies) Bak. f., Jour. Bot. 30: 138. 1892. S. leprosa var. ee (Gillies) Hochr., Ann. Cons. Jard. Bot. Gen. 6: 33. 1902 S. leprosa var. parvifolia (Hemsl.) Hochr., l.c. (nom. nud.). Indument usually lepidote, occasionally semi-lepidote, the inner por- tions of the rays fused, the outer portions free. Leaves to 10 mm. long, 12 mm. wide, reniform. Calyx 3—4 mm. high. Petals often greenish- black on drying, sulfur yellow or yellow flushed with rose in life. TYPE: Collector unknown, “Hab. in insula Cuba” (Madrid). The species is not known from Cuba and a photograph of a probable isotype in Geneva sent by Cavanilles is very much like the Argentina material. DISTRIBUTION: Uruguay, Paraguay, Argentina and Chile; Mexico. 14b. Sida leprosa var. hederacea (Dougl.) K. Schum., Mart. Fl. Bras. 12(3): 342. 1891. Malva hederacea Dougl. ex Hook., Fl. Bor. Am. 1: 107. 1831. Malva californica Presl, Rel. Haenk. 2: 121. 1835. Malva plicata Nutt. ex Torr. & Gray, Fl. N. Am. 227. 1838. Malva obliqua Nutt. mss. cited by Torr. & Gray, l.c., as basonym for following: Sida? obliqua (Nutt. mss.) Torr. & Gray, L.c. S. hederacea (Dougl.) Torr. ex Gray, Mem. Am. Acad. 4: 23. 1849 (Pl. Fendl.). igee hederacea (Dougl.) Greene, Lflts. Bot. Obs. & Crit. 1: 209. ae of free-rayed stellate hairs, becoming partially or completely lepidote in intermediates to var. depauperata. Leaves to 40 mm. long, 50 mm. wide, rotund-reniform to ovate-subcordate, the margins vari- ably and irregularly crenate to acute-serrate. Calyx 4—7 mm. high. Petals usually pale brown or pale brown and rose on drying, white, cream-colored or rose in life. STUDIES IN SIDA (MALVACEAE) 53 TYPE: Douglas, “in the interior districts of the Columbia [River}.” DISTRIBUTION: Washington and western Idaho, U.S.A. to Baja Cali- fornia, Mexico (eastward from Arizona to Texas, Oklahoma and Kan- sas, U.S.A. and south to Querétaro, Mexico, mostly as intermediates to var. depauperata). Reported as a weed in South Australia by Black Chern 14c, Sida leprosa var. depauperata (Gray) Clement, comb. nov. S. lepidota, A. Gray, Smithson. Contrib. Knowl. 3, art. 5: 18. 1852 (Pl. Wright. 1). S. lepidota var. depauperata Gray, l.c. Disella lepidota (Gray) Greene, Lflts. Bot. Obs. & Crit. 1: 209. 1906. Indument of silvery many-rayed scales, like those of var. leprosa, be- coming mixed with stellate hairs, some of which in intermediates to var. hederacea have the bases of the rays fused. Leaves mostly 1—3 cm. long, rarely to 4.5 cm. long, obliquely deltoid, acute, semicordate, occa- sionally yellow, often tinged with various shades of red. TYPE: Wright 45, “Hillsides between El Paso and the Mountains, Aug. 1848.” (Gu). DISTRIBUTION: Arizona, New Mexico and Texas, U.S.A.; Baja Cali- fornia (1 coll.), Sonora, Chihuahua and Coahuila, Mexico. 14d. Sida leprosa var. sagittaefolia (Gray) Clement, comb. nov. S. lepidota var. sagittaefolia A. Gray, Smithson. Contr. Knowl. 3, art. 5: 18. 1852. S. sagittaefolia (Gray) Rydb., Bull. Torr. Club 33: 145. 1906. Disella sagittaefolia (Gray) Greene, Lflts. Bot. Obs. & Crit. 1: 209. 1906. Indument like that of var. depauperata, always lepidote, scattered to dense. Leaves to 54 mm. long, 2—6, rarely to 10, mm. wide, elongate- triangular to nearly linear; base strongly oblique, coarsely laciniate; upper margins entire, usually straight; apex acute, blunt. Petals (9—) 12—15 (—17) mm. long, 8—10 mm. broad, yellow or white, often suffused with red, the depth of color strongest on the portion exposed in bud, or that portion specked with red, or only the base reddened. TYPE: Wright 47, “Mountain valleys, 60 miles west of the Pecos; Aug.” 1848 (GH). The only collection listed in Prof. I. M. Johnston's annotated list of Wright’s field notes and numbers which could possibly be this distribution number is no. 869, Aug. 21, 1849, collected 80 miles west of the Pecos. 54 STUDIES IN SIDA (MALVACEAE) DISTRIBUTION: Arizona, New Mexico and Texas, U.S.A.; Sonora, Chi- huahua, Coahuila and Durango, Mexico. The present organization of this species-complex into a single species with four varieties may seem at first to be the ultimate in reduction, an oversimplification of the situation, yet it has been impossible to find absolute criteria of any kind which would effectively distinguish the entities as species. The host of inter- mediates between S. leprosa var. hederacea and S. leprosa var. depauperata in the area of overlap of their ranges bears uncontest- able witness to the fact that genetic divergence of the types cannot have proceeded very far, although typical examples of each variety are amply distinct from one another in general appearance. In the case of S. leprosa var. depauperata and S. leprosa var. sagittaefolia, there are few intermediates between them, and they apparently do not occur together in the field. Yet their ranges are strikingly similar and they have no differential characters save those pertaining to leaf-shape. Apparently a genetic barrier exists but morphological differences, save of leaf- shape, do not. These specimens have the margins of the leaf above the base straight and entire, but the proportions of the leaf have changed, broadening towards that of var. depauperata. Wooton 402 from Doria Ana Co., New Mexico, of which I have seen seven specimens, represents the transition from one variety to the other quite well, having all but the extremes represented on each of several sheets. The UC and ARIZ specimens are close to var. depauperata, while the NY and US sheets are close to var. sagittaefolia, Other transitional specimens are Warnock 549 and Sperry 1465 from the Dog Flats of Brewster Co., Texas, Havard 134, Presidio Co., Texas, and Mulford 1149, Hudson’s Hot Springs, New Mexico (which I have been unable to locate on any map). Johnston and Mueller 1224 from western Coahuila represents an absolute extreme of leaf-size in this complex. The proportions are those of an intermediate between the varieties, the leaves measuring up to 4.5 cm. long and 1.2 cm. broa (measured as before) or 2.5 cm. broad, measured along the axis of the base. I have seen no other specimens which approach the size of this. The stems are to 40 cm. long, the petals white, tinged with maroon, and the plant is in every respect, save leaf-size, a good specimen of var. depauperata, the inappropriateness of the name never more evident than here, where its largest leaf is 4.5 STUDIES IN SIDA (MALVACEAE) 55 times as long as the largest leaf of an Argentina specimen of the typical variety. The typical variety of the species is found in Chile, Argentina and Uruguay and is rather variable in leaf-form, even as var. hederacea is in North America. It appears again in the region around Mexico City, near San Luis Potosi (Parry & Palmer 75), these very scrappy specimens the basis for the nomen nudum S. hederacea var.? parvifolia Hemsl., and in Coahuila (Fisher 44105), collected along the railroad, near cultivated fields, which may account for its presence so far north. It has been impossible to distinguish this material from the South American, and the parallel distribution of S. physocalyx immediately comes to mind. Other material gradually intermediate between var. /eprosa and var. depauperata or var. hederacea (and one cannot be certain which) appears in a chain from near Zaragoza and Mohovano, Durango, Mexico (Johnston 7791 and Purpus 4538), through eastern Chihuahua (Stewart 682) and northern Chihuahua, Mexico (Rose & Hough 4203, Pringle 7742), to Presidio Co., Texas (Hinckley 1860) and Cochise Co., Arizona (Blumer 1689), U.S.A. The last most nearly resembles var. depauperata, but the leaves are obtuse at the apex rather than acute, and the base and upper margins shallowly and minutely toothed rather than laciniate and sharply toothed as in var. depauperata. 14a. S. leprosa var. leprosa. Chile: Atacama: Gigoux, March 1886, vicinity a Dept. Copiapé (cH). Pop stead 4984, vicinity Copiapé, Dept. Copiap6 (GH). Cogurmso: Joseph 5439, a Serena (us). Werdermann 130, Coquimbo (uc, F, GH, Mo). Worth & Morrison 1 ee east Fray Jorge Forest, Dept. Ovalle (GH, MO, Sites Sastiaalay i 969, aries (us). Joseph 1641, Santiago (us). W oo 1216, 4 Nov. 1920 (us). Behn, 4 Feb. 1934, Pinan (uc). ie “Prov, aes " (cn ga EE a Buenos Arres: Alvares 446, “Km. 54 — F.C.S.” (us). Burkart 8510, Baradero, Rio Parana (ny, F). Eyerdam et al. 23605, 7 kilometers east Mar del Plata (GH, uc). Eyerdam et al. 23731, Dept. Coronel Dorego, Sauce Chico Rio, 120 kilometers west Bahia Blanca (Gu, uc). Parker 3, Buenos Aires (GH). Juyuy: Lorentz & Hieronymus, 13 May 1873, El Volcan (nx). Venturi sa Tilcara, Dept. Tilcara (us). Venturi 10075, Tres Cruces, Dept. Humahuaca MENDoza: Gillies 105, Mendoza, (1soType or TyPE fragment of Malva alpbare Gillies: GH). Rio Necro: Fischer 210, vicinity General Roca (ny, MO, US, F, GH). Meyer 7242, Salcheta (us). Wilkes Exped. 1838-42, “Rio Negro, north tes (cu, us). SALTA: Venturi 6911, San Carlos, Dept. San Carlos (us). TucumAn: Venturi 445, Chanar Pazo, Dept. Leales (us). WrrHout 56 STUDIES IN SIDA (MALVACEAE) DEFINITE Lity: Hicken 738, La Plata? (ny). Moreno & Tonini 353, fe s673" (Ny). Lorentz 168, Feb.-April 1881 “Sierras Pampeanas’’ (us, GH Uruguay: CANELONES: Rosengurtt B-3073, Canelones (Ny). “Cuba”? (see discussion of Type): Collector unknown, (photograph of TYPE in Madrid Herbarium: us). Collector unknown, (photograph of IsoTYPE? in Delessert “ag SS m: F). Mexico: Coanutta: Fisher 44105, Viesca (GH). SAN Luis Porosi: Parry & Palmer - region of San Luis Potosi (GH, Mo). StaTE oF Mexico: Pringle 11934, Tlalnepantla (Gu). Rose 8398, Tlalnepantls (GH). Schaffner 187, “Vallée de Mexico” (GH). FepERAL District: Patoni-Ochotorena 4998 (MExU). Pringle 9415, Mexico City (GH 2). Bourgeau 32, jee of fields near Mexico City (GH). S. leprosa var. leprosa [intermediates to var. phasic or var. hederacea. | Mexico: Cummuanua: Pringle 7742, Rancheria (MExu 2). Rose & Hough 4203, Rancheria (GH, uc). Stewart 682, n.w. of Jaco (GH). CoanuiLa: Purpus 4538, “Movano, Coahuila” (uc). Duranco: Johnston 7791, road from Bermejillo to Zaragoza and north to Mohavano (cu). United States: Arizona: Blumer 1689, Paradise, Chiricahua Mts., Cochise Co. (GH 2, ARIZ, F, NY 2, MO, DS, US). Texas: Hinckley 1860, Sierra Tierra Vieja (GH). 14b. S. leprosa var. hederacea. United States: WASHINGTON: Oxanocan Co.: Fiker 1424, between Nuttall, Oregon plains (GH 2). IDAHO: Wasnincton Co.: Davis 3006, 5 miles south Weiser ay ps). CALIFORNIA: (selected citation sufficient to et range): ALAMEDA Co.: Brewer 849, Corral Hollow (GH, Mo, uc, Us). AMADO Co.: Braunton ats near Ione (ny 2, uc, MO 2, us). CoLusa Co.: Heller mpers near Colusa (GH, Ny, MO, Ds). Stinchfield 294, College City, Tule levee (POM, NY, DS). Contra Costa Co.: Rose 38310, Byron (uc, mo). Abrams 5728, Concord Road between Walnut Creek and Martinez (ps). Inyo Co.: Coville & Funston 951, 2 miles south Lone Pine (us). Kern Co.: Bauer 8 Oct. ed near Bakersfield (ps). Kincs Co.: Kearney 24, near Hanford (Mo, us Co.: Blankinship 1 Sept. 1926, Kelseyville (mo). Lassen Co.: Burtt Dav 3330, Noney Lake Valley (uc). Los ANGELES Co.: Abrams 2959, near Mesmer (UC, NY, POM, MO 2, DS, us). Manz 6622a, San Clemente Island (pom). Trask 258, west end San Clemente Island (us). Mercep Co.: Wolf 725, 9 miles east Los Abrams Man Pajaro River on Los Boba. road (ps). STUDIES IN SIDA (MALVACEAE) 57 SAN BERNARDINO Co.; Wheeler 2079, Chino Creek (GH, NY, » MO). SAN Dieco Co.: Chandler 5396, Escondido (uc, ny, ps). Orcutt on San Diego (GH, UC, Us 2). Youngherg 11 July 1934, Lindo Lake, Lakeside (pom, uc, ps). San JoaQuin Co.: Burtt Davy 826, Stockton (uc 2). Michener & Bioletti 12, San Joaquin Bridge (mo, uc, GH, us 2). SAN Luis Optspo Co.: Brandegee 2 Oct. 1912, San Miguel (uc, us). SoLano Co.: Jepson Sept. 1891, Vacaville (Ny, Us 2). Baker 3223, Suisun (GH, NY, POM, MO). SANTA CLARA Co.: Dudley, 4 Oct. 1895, Lake — seveghing Univ. (ps). STaNnisLAus Co.: Hoover 108, Crow’s Landing (uc). Ventura Co.: Feudge 1156, Upper Ojai Valley (pom). Yoto Co.: Keck eee 6 ne snith Woodland (ps). WirHouT DEFINITE 1767, Carson Sink region (pom, ps). Tidestrom 10780, near Carson Lake (cu, NY, US). HumBotpr Co.: Torrey 56, “sterile saline plains. 1865” (Ny, MO, GH). LANDER Co.: Hitchcock 598, Battle Mountain (us). Lyon Co.: Allen 247, 6 Co.: Peebles & Fulton 9551, near Leupp (us). Ward, 20 June 1901, Woodruff (us, ny). Pinan Co.: Peebles 9409, near Sacaton (uc, us, POM). YUMA Co. County Agriculture Agent, June 1925 (Ariz). Peebles, Harrison & pace? 4935, near Dome (us). Swingle S$. 258, Yuma (ariz). Thornber, 26 Sept. 1912, Yuma (aniz). Wilkinson, 27 Aug. 1905, Yuma (Ariz). WITHOUT DEFINITE LOCALITY: Millspaugh 151, to the Grand Cafion (Fr). NEW MEXICO: BERNALILLO Co.: Herrick, 7 Sept. 1894, Albuquerque (us). Dofia ANA Co.: Archer 445, 5 miles north of Mesilla Park (smu). Child 524, Archer Ranch, Brazito (mo). Eggleston 20221, Mesilla Park (us). Ferris 1147, 5 miles from Las, Cruces (ps). Fosberg 53457, Mesilla Valley, Lower Sonoran Zone (uc, pom). Wooton, June 4 1893, ae ae mm 2, MO). Wooton 16, Mesilla (wyo, POM, MO, ARIZ, UC, US, Ds, . Wooton, May, 1899, Mesilla Valley (ariz, wyo). Wooton, 26 May, 1899, "Mesilla Valley (ps, uc). Wooton, 15 Oct. 1901, Mesilla Valley (pom). Wooton & Standley 3283, Mesilla Valley (F, ps 2, NY, Wyo, MO 2). SOCORRO “banks of the Brazos’ (cH, Mo). EasTLanp Co.: Oyster, 28 May 1883, vicinity Eastland (ny). Ex Paso Co.: Barlow, 1 July 1911, Rio Grande Valley at Canutillo (Fr). Cory 2064 (GH). Cory 52932, 7 miles east El Paso (GH, smu). ewey, 15 June 1891, west of El Paso (us). Earle 474, El Paso (ny 2). El Paso (vac, us, Ny, uC 2, F, POM, Ds). Mearns 677, El Paso (us). Mearns 1522, Ft. Hancock (us). Palmer 31097, near El Paso (Mo). Shinners 8926, south east side of El Paso (smu). Stearns 12, vicinity of El Paso (us). Vasey, May 1881, El Paso (us 2). Waterfall 3959, 31/2 miles southeast of Clint (GH). W bitehouse 8398, El Paso (Fr). Howarp Co.: Palmer 12477, Big Springs (uc 2 58 STUDIES IN SIDA (MALVACEAE) MO). Hupspetu Co.: Waterfall 3985, Rio Grande Valley, near Ft. Quitman (cH). Waterfall 4590, along Rio Grande, McNary-Ft. Quitman levee road (ARIZ, GH, MO, sMU). Jerr Davis Co.: Waterfall 5328, 3 miles west Chispa 3824, Amarillo (Mo 2, POM, NY, GH, US). REEves Co.: Nealley 715, 42, Pecos Flat, mear Pecos City (us, Fr). Taytor Co.: Tolstead 7545, 18 miles southwest of View, on Edwards Plateau. (GH 2, smu 2). YounG Co.: Reverchon, 29 Oct. 1902, banks of Brazos, near Graham (Mo). WirHouT DEFINITE LOCALITY: Wright 44, “western Texas” (GH 2, uc, Ny). Wright 45, “western Texas” (us, UC, GH 2). Young, 12 Sept. 1917, Llano Estacado, near Paloduro Canyon (Mo). ico: GuapALuPE Istanp: Thoburn, Green & Wing, July 1897, near northeast landing (ps). REVILLAGIGEDO IsLANDs: Howell 8359, Sulphur Bay, Clarion Island (ps). Baya Ca.irorniA: Brandegee, 5 April 1889, San Gregorio (uc). Orcutt, 31 Aug. 1889, Ensenada (mo 3). Brandegee 50, San Jose del Cabo (uc). Sonora: Palmer (ps). Thurber 424, Fronteras (GH). CHIHUAHUA: Pringle 5278, Paso del Norte (mexu). Stearns 12, “state of Chihuahua” (F). CoanuiLa: Palmer 92, Lorenzo de Laguna (GH). QuERETARO: Deam 95, Querétaro (GH). WirHouT DEFINITE LocaLity: Thurber, “1852” (cu). 14c. S. leprosa var. depauperata. United States: ARIZONA: CocuisE Co.: Darrow, Phillips & Pultz 1321, Apache Pass (Gu, uc). Shreve 6361, south of Rodeo (F). Thornber 2562, Sulphur Spring Valley (ariz). Thornber 260, et Pedro Valley, Benson (ARIzZ).: anaes 4642, Bowie (ariz). Monave Co.: Lemmon 3255, Kingman (GH). A Co.: Lemmon 14, Tucson (GH). YAVAPAI ig Mearns 191, Ft. Verde (Ny 2). stort DEFINITE LOCALITY: Rothrock 188, Deer Springs (F, Gu, us). NEW MEXICO: Catron Co.: Metcalfe, Aug. 1901, Mangas Springs (ARIZ, US). Dofia ANA Co.: Standley, 17 June 1906, Mesilla Valley, College Farm (cH). Wooton, 22 April 1894, Mesilla Valley (ps, aniz, uc, Mo, POM). Wooton & Standley 3236, Mesilla Valley (us, Fr, ps). Granr Co.: Hevlinon 16509, along Gila river and on mesa above Cliff (F). Eggleston 19992, Shelley “Poison apres Cliff (GH). Holzinger, 31 July 1911, Hanover Mt. (us). Holzinger, 27 Little Florida Mts. (Ds, POM). i 16261 and 16262, Nutt (Mo, F). Mulford 1024, Deming (mo, ny). RA Co.: Beals, Sept. 1914, Lake Valley (us). WITHOUT DEFINITE LOCALITY: Nae 10764, Meenjani Road (mo). Wooton, 5 July 1906, Cactus Flat (us 2). Wright 889, sandy soil between the Rio Grande and Mimbres, June (GH, TYPE & ISOTYPE, NY, UC, MO, US). TEXAS: Brewster Co.: Sperry 1335, Dog Canyon Wash (us). CuLBERSON Co.: Earle & Tracy 406, R. near Kent (Ny, us). Ex Paso Co.: Wright, Oct. 1904, El Paso (ps). Howarp Co.: Tracy 7817, Big Springs (mo, Ny 3, GH, US). (cH, smu). Hanson, 17 Dec. 1918, Pecos (ny). Hanson 45, Pecos (GH, MO): STUDIES IN SIDA (MALVACEAE) 59 TayLor Co.: Tolstead 7241, Camp Barkeley (mo, uc). W1THoUT DEFIN LOCALITY: Havard, 1881 (us). Wright 45, “west Texas’, (Type no. of S. lenidee var. Sander (UC, GH, MO, Ny, US). Mexico: Baja CALIFORNIA: phere 3 Sept. 1893, San José del Cabo (GH). ee Schott, 25 June 1855, El Potrero (Fr). Cumuanua: Stearns, 1911, valleys, near Ciudad Juarez (Ny 2). WAste 2613, Carretas (GH). COAHUILA: Aguirre & Reko 116, Paila (Ny). Johnston & Muller 1224, north end of Bolson de los Lipanes (cu). S. leprosa var. depauperata [intermediates to var. sagittaefolia.] United States: New Mexico: Maualford 1149, “Hudson’s Hot Springs” (Mo, Ny). ao eae nig plains south of the white Sands, Dofia Ana Co. (ny, ARIZ, UC, POM, ps). Texas: Sperry 1465, Dog Flats, Brewster Co. (GH). Warnock esib same pean (us). Havard 134, Presidio Co. (GH). 14d. S. leprosa var. sagittaefolia. United chic ARIZONA: Cocutse Co.: Parish, 2 May 1884, — (NY, UC, Ds, Us 2). Lemmon & wife, 11, 1880 Ft. Lowell Aah ie A Co.: Lemmon Herberiune 11, Aug. 1880, nr. Gila Bend (GH). Mouave Co.: Boiwoad 18455, road from Kingman to Peach Springs (us). McKelvey. 2267, between Hackberry and Peach Springs (pom). Prima Co.: Carter, 1 Oct. 1933, Tucson (anIz). Gould 3062, 10 miles east of Tucson on Nogales Road (ARIz, UAC, UC, MO, US, GH). Griffiths 8962, Baboquivari Valley (mo). Jones, 25 Aug., 1903, Tucson (pom). S. B. & W. F. Parish, April 1884, Tucson (uc). Peebles, Harrison, Kearney 3849, Papago Reservation (us). Pringle, 29 June 1881, (Mo, 13635), Santa Cruz Valley near Tucson (Ny, GH). Shreve 7376, near Tucson (Mo, F, ps). Spalding, 21 March 1906, Altar Valley (east side of Baboquivari Mts.), (artz). Pra Co.: Thornber 54, Santa Cruz Valley, Tucson (ariz, us). Thornber 453, Santa Cruz bottoms (Ariz 2, MO, UC, DS, US). Peebles 7390, near Casa Grande (us). Thornber 7347, Casa Grande (Ariz). YAVAPAI Coues & Palmer 524, Rio Verde, Ft. Whipple (mo, Gu 2). Purpus 8268, Plains, Beaver Creek (Mo, uc, US). WITHOUT DEFINITE LOCALITY: Engelmann, 23 Sept. 1880, Mesa, Arizona Desert (mo). Potts, southern Arizona (Mo). Purpus 72, Black Mesa (Mo, uc, US). ae MEXICO Co.: Rusby 51, alluvial flats, north of Socorro (uc, F). WITHOUT DEFINITE LocaLity: Rusby 51, dry plains near Alamillo ete an. eee 51, Mogollon Mts. (ny). Rusby 51, dry plains, Jornada del Muerto (mo). sucess 1848 (Ny, GH). Rusby, 1881 (ny). Wright 1331 (Gu 2, uc, NY 3, MO, us). TEXAS RD : Pohl 4424, 4 miles north of Murphy School (smu). BrewsTEeR Co.: Cory 53130, 5 3/4 miles northwest of Alpine (smu). Cory 53131, 5 3/4 miles northwest of Alpine (GH, sMU). Ingram 2829, 16 miles east Alpine (us). Palmer 34031a, Alpine (mo). Schulz 11 Oct. 1931 (r). Tharp 18 Aug. 1935, Stockton, Alpine (mo, GH, uc, smu). Tharp 175 (Mo, vc, cn). Tharp 3581, 60 STUDIES IN SIDA (MALVACEAE) Marathon (us). Crossy Co.: Erlanson 1166, 20 miles southwest Spur (sMv). Howarpb Co.: Letterman 104, Big Springs (Mo, us). Kinney Co.: Cory 16781, 9 miles south Spafford (GH). La SALLE Co.: Cory 28518, 73/4 miles west Fowlerton (GH). Lusspock Co.: Reed 3137, Lubbock (us). Mr anp Co.: Tracy 7814, Midland (mo, ny 3, GH, US): Pecos Co.: Cory 17513, 41/2 miles northeast Hovey (GH). Presipio Co.: Drushel 11159, near Marfa (Ny, F). Hinckley 701, 12 m.s. Marfa (ny, F). Hinckley July 1936, San Esteban Lake, Marfa (GH). Hinckley 713, Marfa (Fr). Nealley 43, “The Chenatic’ (F). ZAVALLA Co.: Palmer pple ae City (mo). Tharp 6037, Crystal City (us). WrrHouT DEFINITE LOCALITY: V. Havard 188, Mts., We estern Texas (us). Nealley 788 (ps). Wright 47, Ate valleys west of the Pec UC, MO, GH TYPE 2, NY, US). Mexico: Sonora: Wiggins 6370, 1/2 ioe west Punta Piedras, 15 miles west of Empalme (ps). IV ee eee Pee pte west Sonoyta (Ds). eg A: LeSueur 383, El Carmen (F, GH, sMU). Palmer 220, near Chihuahua, (Ny, G MO, US). Pringle 27, Llanos, Escalén (MEXU). ee wart 2366, 1 kilometer ane Escobillas. (GH). Stewart 2589, near Trinidad, road from Guimbalete west he south end of Sierra see sig (cH). Thurber 846, El Gallote (GH 2, NY 2). White 2030, 6 miles t Guimbalete, road to Escalén (GH). W hite ae Escal6n (cH). Wilkiwson, ghoes Santa Eulalia Plains (Ny). Coanumia: Fisher 44136, 3 miles northeast Torreén (Gu). Fisher 44152, 3 miles east Torreén Encantada Ranch Hq. and east to escarpment of the Sierra del Carmen, northwest of Muzquiz (GH, F). Marsh 1530, vicinity Encantada Ranch Hq. and east to escarpment of the Sierra del Carmen, northwest of Muzquiz (Gu, F). Stewart 432, 18 miles south Castillon, road to Santa Elena Mines, via Jesus Maria India (GH). Stewart 2777, Rancho Parritas, 5 kilometers south base of mountains along east margin of Valley de Acatita (GH). Duranco: Palmer 536, Mapimi (GH, NY, UC 2, Mo 2, MEXU). Patoni-Ochotorena 324, near San Juan de Guadalupe, south of Sierra de Ramirez (MExU). WirHouT DEFINITE LocaLity: Schott 9 July 1855, Escopeta Valley (F). IV. Section INCANIFOLIA Clement, sect. nov. Herbae prostratae, omnino molle stellato-canescentes; folia symmetrica, Crassius griseo-viridia; flores axillares subsessiles; lobis calycis angusto- ovatis; carpella 5, paullum inflata et ad modum carpellorum Abutilonts acuminata. Prostrate herbs, everywhere softly stellate-canescent; leaves symme- trical, dark gray-green; flowers subsessile in the axils; calyx lobes long- ovate; carpels five, slightly inflated, peaked. The two species of the section have customarily been placed in section Pseudomalvastrum, although they are never bracteolate and have symmetrical leaves which are softly canescent, rather STUDIES IN SIDA (MALVACEAE) 61 than asymmetrical leaves which are scabrous. The carpels are always five in number here and always more than five in section Pseudomalvastrum. Type species: Sida Grayana Clement in Kearney. KEY TO THE SPECIES Carpels 5.5 mm. high, with two protracted blunt apical peaks from the middle of the upper dorsum (similar to those of Abutilon); stamen-column 3. a; high, the lower half with coarse, scattered, stellate hairs; petioles 3—6 mm. long; stipules 5—6 mm. long, oblanceolate, acute. . . . 15. S. Grayana. Carpels 5 mm. high, the apex peaked but not protracted; stamen-column 3.0 mm. high, glabrous; petioles 6—12 mm. long; stipules 7 mm. long, ovate, Obrase eos Re ae Sa ee ee eee eee eR ls. 15. Sida Grayana Clement in Kearney, Lflts. West. Bot. 7: 140. 1954. S. cuneifolia A. Gray, Bost. Journ. Nat. Hist. 6: 165. 1850, non Roxb., Fl. Ind. 3: 170. 1832. Disella cuneifolia (Gray) Greene, Lflts. Bot. Obs. 1: 209. 1906. Perennial assurgent subshrub. Stems many from a slender, tapering, pale, woody taproot, freely branching, up to 45 cm. long, woody; the whole plant covered with a canescent, soft, fine-rayed, stellate pubes- cence with a characteristically soft feel; the wood slightly yellowish, with a high waxy luster when cut; leaves suborbicular to rotund-cune- ate, 3 (—5)-nerved, the nerves prominent beneath, variably crenate to serrate, except on the base, concolorous, 6—14 mm. long, 6—16 wide; stipules oblanceolate, acute, spreading, slightly folded on the median nerve, 5—6 mm. long, ca. 1 mm. wide, persistent; petiole terete, 3—6 mm. long; flowers solitary or clustered on much shortened branch- lets in the axils; pedicels 2—3 mm. long, terete, erect, articulated near the base, elongating slightly in fruit; calyx campanulate, terete, 5—7 mm. long, lobes ovate to lanceolate, acute, 3—4.5 mm. long, 1—1.8 mm. wide, erect, closing slightly over the fruit, with one main nerve and several fine small ones, stellate-pubescent within except the tube; petals yellow, obovate, subtruncate, slightly oblique, 5—6 mm. long, 3 mm. wide, the claw 1.5 mm. wide, rather coarsely veined, inserted just below the level of the top of the ovary; stamen-column slender, terete, 3.5 mm. long from base of ovary to level of the filaments, with scattered, Coarse, stellate hairs on the lower part and tufted stellate hairs between the claws of the petals. Stamens 18—20, reniform with a deep, narrow sinus, pale yellow, with many fine red-brown dots, 0.5 mm. wide open and closed; filaments 1—1.5 mm. long, not paired; pollen 0.03 mm. 62 STUDIES IN SIDA (MALVACEAE) diam. Ovary globose, densely stellate-hirsute apically, thinning basally, ca, 1 mm. high, 1 mm. wide, almost filling the cavity. Styles 4.5—5.5 mm. long, 4—4.5 mm. free, subconnate in bud, protruding in early anthesis, reflexing later; stigmas capitellate, 0.1 mm. in diameter; carpels 5.5 mm. high, 2 mm. radially; apex peaked, with two blunt, stellate- pubescent, slightly introrse aristae, dehiscence between them by a well- marked suture, the dehiscence often continuing to the base; lateral walls membranaceous, pale, smooth, 2 mm. high, 1 mm. wide, basal; dorsum stellate-hirsute, denser and longer-rayed apically, nearly glabrous basally; inner surfaces straw-colored, shiny, perfectly glabrous except — on the strigose margins of the suture. Seed globose, glabrous, dark red- brown, minutely verruculate, filling all but the apex of the carpel; the funicle broad and flat, with a few setulose hairs of the same color. Co- lumella slender, narrow-flanged, 2.5 mm. high, pale, base of the styles persistent. TYPE: Wright, 1848, “35 miles northeast of Eagle Pass, Rio Grande, Texas. Place overgrown with plants indicative of salt, Sept.” (Gu, frag- ment US). DISTRIBUTION: Southern Texas; Tamaulipas and Nuevo Leon, Mexico. The name Sida cuneifolia was first used by Roxburgh (1814), without accompanying description, for a plant growing in the Calcutta garden. He later (1832) provided a description and validated the name. In 1834, Wight and Arnott identified this plant with Riedleia truncata (Willd.) DC. (1824), based on Melochia truncata Willd. (1800), but in 1837, Wight, with better material available, showed that Willdenow’s plant and Rox- burgh’s were actually of the Malvaceae rather than the Sterculia- ceae and proposed a new genus, Dictyocarpus, closely related to Sida, for the species. Bentham and all others since have agreed that it belongs in Sida. Sida Schimperiana Hochst. ex A. Rich. (1847) is apparently the same thing, described from African material, and this name is often applied to the Indian material. Sida cuneifolia Roxb. is the first available name, there already being a Sida truncata, of Cavanilles (1785), and should be used. It is earlier than Gray’s Sida cuneifolia by 28 years, and I have renamed the latter species for its author. Hochreutiner (1902, p. 33) remarks under S. cuneifolia Gray, “Nous voudrions signaler ici une plante de Lemue, provenant de Litakoun, Afrique australe, et qui nous parait étre identique a la plante américaine”. I have seen no African material which resembles this species nor have I read of any anywhere else. STUDIES IN SIDA (MALVACEAE) ae United Rentee: TEXAS: Bexar Co.: Parks 4819, Near San Antonio (mo). CAMERON Co.: Cory 51383, Loma Alto, 8 airline miles northeast of Brownsville (SMU, US, GH). Hare 19 May 1919, 10 miles east Brownsville (GH). Hunt 16, Brownsville (ps). Runyon 455, Brownsville (us). Runyon 654, Brownsville (us). Shiller M-115, Brownsville (mo, GH). Dimmrr Co.: Jones 28239, Carriso Spring (uc, POM, MO, Ds). Tharp 593, 10 miles south of Big Wells (vs). Jim Hocc Co.: Hanson 30, 25 miles south of Hebronville (mo, us). La SALLE Co.: Cory 14980, 2 miles south of Los Angeles (GH). MAVERICK eu Wright, 1848, 35 miles northeast of Eagle Pass, Rio Grande (GH, US). EBB Co.: sperints 26 April, Te valley, 7 miles east of Laredo (ny). WITHOUT DEFINITE ocaLity: Parks M-115 (mo). Wright 48, = westert Mae 1849” (GH 3, US). Berlei 645 & 2055, Rio Frio (GH 0: Nuevo Lrén: Berlandier 3104, (GH, NY 2, MO, US). TAMAULIPAS: aide pee vicinity El Mulato (F). 16. Sida Helleri Rose, ee pe Franklin & Marshall Coll. Perennial prostrate shrub, . cm. or see high, to 60 cm, in diameter. Stems woody, numerous from near the base, only slightly branching; flowering shoots prostrate to erect, herbaceous, clothed with a soft, very slender-rayed, stellate indument, the older stems becoming woody and glabrous. Leaves suborbicular, crenate to obtusely serrate except the rounded, subcuneate or truncate bases, finely stellate-pubescent, scattered above, dense beneath, 8—13 mm. long, 1 10—16 mm. wide, three-nerved, the nerves eS above, prominent and straw-colored beneath. Petioles 6—12 mm. long, terete or slightly ribbed under the dense in- dument. Stipules obtuse, ovate, foliaceous, usually persistent, appearing as bracts at the base of the pedicels, 7 mm. long, 3 mm. wide, 1-nerved, slightly folded, erect. Flowers subsessile, solitary in the axils. Calyx campanulate, terete, accrescent after anthesis, 4—5 mm. high in flower, 5—8 mm. high in fruit; the lobes ovate, without apparent nerves, re- maining erect in hae obtuse or subacute, densely stellate except the inside of the tube proper, 3—3.5 mm. long, 1—1.5 mm. wide; epicalyx none. Petals “pale copper-colored”, broadly obovate or obliquely sub- orbicular, broadly and shallowly emarginate, glabrous throughout, 4—5 mm. long, 3—4 mm. broad, inserted on the stamen-column above the top of the ovary, venation very fine, Stamen-column slender, faintly ribbed, glabrous, 3 mm. high from the base of the ovary to the level of the filaments; filaments all from the same level, not paired, ca. 1 mm. long. Anthers 20, ca. 0.8 mm. wide open, ca. 0.5 mm. wide closed, minutely dark-verruculate; pollen ca. 0.05 mm. diam. Styles five, 5 mm. long, 3 mm. free, reflexed in anthesis; stigmas capitellate. Ovary 1.5 mm. high and wide, subspherical, densely strigose laterally, densely hirsute apically; the styles inserted high within the ring of ovules. Fruit deeply five-lobed; carpels five, 5 mm. high, 3 mm. radially, inflated 64 STUDIES IN SIDA (MALVACEAE) apically, partially dehiscent by a medio-dorsal suture, the margins thick- ened; dorsum obtusely peaked apically, the peak hirsute to the funicular opening, remainder of the dorsum short-strigose; lateral walls chartace- ous, smooth, 2 mm. wide radially, 3 mm. high. Seed plump, filling the lower two thirds of the carpel, puberulent; the wall woody, hard, dark red-brown; funicle flattened, puberulent, lighter red-brown. Columella 3 mm. inti 5-flanged, the basal portion of the stylar column often persist TYPE: Hole 1533, shores of Corpus Christi Bay, Nueces Co., Texas S). DISTRIBUTION: Southern Gulf coast of Texas, United States. United States: Texas: Heller 1533, shores of Corpus Christi Bay, Nueces Co. (us 2, NY, UC, ARIZ, GH, F). Shiller 350, near Brownsville, Cameron Co. (us). V. Section OLIGANDRAE Clement, sect. nov. Herbae annuae; folia palmatilobata; carpella aristis elongatis retrorse barbatis; antherae 5—20; carpella 5—10; calyx accrescens non tamen inflatus; epicalyx deest; flores paniculati (flores inflorescentia foliosa paniculata); parietes laterales carpellorum ad semina firmiter adheren- tes. Annual herbs. Leaves palmately lobed. Calyx somewhat accrescent but not inflated; epicalyx absent. Stamens 5—20. Carpels 5—10, long- aristate, the aristae retrorsely barbed, the lateral walls firmly adherent to the seed. A well-defined natural section of western Bolivia, Peru and Ecuador, section Oligandrae is composed of species previously placed in section Sida. Fries (1947) treated the species as an “Artgruppe” with this name but did not formally establish a new section. The treatment here presented is at best preliminary because of the lack of material of certain species and the relatively few specimens of others. In addition, it is felt that were collections available from intervening parts of the disjunct ranges of certain species, the distinctions between them might easily be removed. Still another barrier to satisfactory treatment has been the un- availability of type or authentic material of two species which are, then, maintained “de fide” only. Type species: Sida palmata Cay; KEY TO THE SPECIES A. Carpels 7—9; stamens 10— B. Leaves 3—5-lobed, lobes id, or slightly narrowed at t the base ‘margin regularly dentate. 17. S. palmata. STUDIES IN SIDA (MALVACEAE) 65 B. wegtiess mostly 7—9-lobed, lobes lanceolate, shes meta narrowed to- ward the base and deeply divided . . S. jatrophoides. A. Carpels : ; Ree RO asa ROR eME eat C. Stamens 10, stamen- abe Villon Se eh Re a D. Leaves mostly 3-lobed; aristae of the carpe a 12—14 mm. long 19. S. decandra. , Sa apres: 5—7-lobed; hes Ca; DS Ini. lous os gt 20 FeO C. Stam 5, stamen-tube gla Pe, Se ae Sr oe E. pny ‘eluate lobed; aristae ney mm. lone cA 2h Sh omaegeion E. Leaves deeply lobed AS oe a ag ee ee F, Aristae of the carpels ca. 14 t mm. longs ea icin RZ ema F. Aristae of the carpels Aas m. long . G G. Stamen-column 1.5—2 m uu petals with a tuft of stellate irs in ‘the outer eras visible in the bud . 23. S. patuliloba. G. Stamen-column ca. 3.5 mm. high; pet tis eiemone caatpinal tuft but with scattered, ome stellate baits on the outer surface 24. S. oligandra. 17. Sida palmata Cav., Diss. 1: 20, t. 2, f. 3. 1785. S. ricinoides ’Hérit., Stirp. Nov. 115, t. 55. 1788. Malvinda palmata Moench, Suppl. 203. 1802. S. Ricini Spreng., Syst. 3: 116. 1826. Erect annual herb, 10—18, dm. high. Stems terete, covered with a white tomentum of spreading, stiff, unicellular hairs 1—3 mm. long, seen, possibly absent. Petioles of the upper leaves 5—15 cm. long, faintly ribbed, with a tomentum like that of the stems. Leaves 3—5 Jobed, reniform in circumscription; upper leaves to 10 cm. long and 15 cm. wide, outer pair of lobes smaller, deltoid or ovate, acuminate, divided about halfway to the base or slightly more; base cordate, the sinus deep pressed, thin, hyaline, unicellular hairs of two sizes: to 2.5 mm. long and to 0.5 mm. long, more prominent on the veins; indument of the lower surface densely tomentose, the stellate hairs of two types: scatter- ed, mostly 3-rayed hairs, the rays ca. 0.5 mm. long, and denser, much smaller, finer, many-rayed hairs. Lower leaves caducous. Inflorescence of few-flowered terminal panicles, additional solitary flowers from the lower axils rarely present. Pedicels terete, ca. 3 mm. thick, conspicuously articulated G6—9 mm. below the calyx, flexed above or around the arti- culation, elongating in fruit to as much as 5 cm.; pedicels of the upper- most part of the panicle much abbreviated, ca. 0. 5 cm. long in anthesis; tomentum like that of the stem. Calyx green, cupuliform-campanulate, 5—6 mm. long in anthesis; lobes deltoid, acute, ca. 3 mm. long, 2.5 mm. wide at the base, without prominent nerves; tomentum of the ex- 66 STUDIES IN SIDA (MALVACEAE) terior like that of the stem with additional scattered, mostly 3-rayed, stellate hairs; margins finely ciliate; interior finely puberulent, becoming sericeous-hirsute on the lobes. Petals obovate, truncate, 5—6 mm. long, ca. 4 mm. wide, finely veined; the outer upper surface sprinkled with fine, multicellular, twisted, simple hairs; claw 1 mm. wide, finely villous on the margins. Stamen-tube 1.5 mm. high from base of the petals to the filaments, conoid, assurgent-villous on the lower portion, with ten paired traces; filaments erect, reflexing after anthesis, 1.5 mm. long, somewhat paired. Stamens ten, fifteen or twenty, ca. 0.5 mm. wide when open, yellow, not dotted. Ovary obturbinate, subangulate, glabrous, olivaceous, less than 1 mm. high and wide, 5-beaked, each beak split. Styles 3—4 mm. long, 1.5—2.5 mm. free, slender, reflexed in anthesis; stigmas subcapitellate, minutely papillose. Fruiting calyx accrescent after anthesis to 1 cm. high, remaining green; the lobes 5—6 mm. long and wide, erect, acute or acuminate; the midnerve of each lobe becoming prominent. Carpels seven to nine, 4 mm. high, subquadrate viewed. laterally; dorsum finely downy-pubescent, the outer surface invaginated and with a conspicuous midnerve; lateral walls membranous or scarious, subrugose, not or slightly adherent to the seed; aristae 5 mm. high, erect, retrorsely setose, slightly divergent, never reflexed. Columella conical, truncate, 3.5 mm. high, occasionally splitting or flattening on drying and pressing; base of the styles persistent; remnant traces to the lateral walls of the carpels sometimes persistent, curving upward and outward from the base of the columella. Seed dark brown, 3 mm. high, 2 mm. tadially, dull, pulverulent; the rostrum assurgent, thickly covered with collapsed, obtuse, hyaline cells; funicle flattened at the hilum, becoming terete along the rostrum. TYPE: Specimen cultivated in the Jardin des Plantes, Paris, from seed collected by Dombey, presumably near Lima, Peru (P). DISTRIBUTION: Ecuador and Peru. The nomenclatorial confusion surrounding the application of the name S. palmata is discussed under S. jatrophoides, with whic this species has often been confused, even by its author. Cavanilles’ original description and illustration were sufficiently precise so that the name can be applied only to the taxon which has 7— carpels, ten or more stamens, and the leaves divided into three or five lobes, the lobes about half as long as the leaves and triangular or at most slightly narrowed at the base. In all the material of this section available to me, the only specimens which fit the original description of S. palmata are from near Huigra, Ecuador, although the species was said by Cavanilles to have been observed by Commerson near Lima. It was originally described STUDIES IN SIDA (MALVACEAE) 67 from living material grown in the Jardin des Plantes, presumably from seed collected by Dombey in Peru. I have seen no modern collections of the species from that country, and all the material from around Lima which has more than five carpels is unquestion- ably S. jatrophoides. Ecuador: Cuimporazo: Camp E-2999, near Huigra (us). Hitchcock 20335, Huigra (Ny, us, GH). Hitchcock 20620, Huigra (us, GH). Rose 22176, vicinity of Huigra (GH, Ny, US). Peru: Dombey, (Photograph of specimen in G: ¥F, GH). Dombey, Lima (Photograph of specimen in M: F). 18. Sida jatrophoides |’Héritier, Stirp. Nov. 117, t. 56. 1788. S. palmata Cav., Diss. 2: 48. 1786 and Diss. 5: 274, t. 131, f. 3. 1790 in part, non Cav. Diss. 1: 20. 1785. Erect annual herb, 5—6 dm. high. Stems woody, glabrescent or short- hirsute, hairs of the indument of two lengths, spreading, white. Stipules early deciduous, not seen. Petioles subangulate, to 7 cm. in length, the indument like that of the stem. Leaves cordate in circumscription, the sinus deep and narrow, palmately 7—9-lobed; lobes acute, ca. 6 cm. long or less, divided almost to the base, at least two-thirds of the way, often with secondary lobules or sublaciniate, lanceolate, narrowing be- low the middle; nerves 7—9, prominent on both surfaces, pale; upper surface with few, scattered, appressed, simple or few-rayed stellate hairs, these flavid golden or brownish mostly pointing apically; margins ci- liate; lower surface paler than the upper, with scattered, mostly 3—4- rayed, yellowish, stellate, appressed hairs. Inflorescence of few-flowered terminal and subterminal glomerules elongating to form lax panicles, a few flowers occasionally solitary in the axils as well; pedicels very short, to 1.5 cm. long, slender, densely tomentose, terete, articulate near the apex. Calyx campanulate, densely long-sericeous, the hairs occasionally to 4 mm. long, yellowish; flowering calyx 4—5 mm. high, the lobes lanceolate, acute to subacuminate, ca. 3 mm. high, 1—1.5.mm. wide; indument of appressed, sericeous, apically pointing hairs. Petals broadly obovate, slightly oblique, 3.5—5 mm. long, 2.5—3 mm. wide, contorted after anthesis, scattered-pulverulent; claw ca. 1 mm. wide, the margins few-ciliate; veins not prominent. Stamen-tube conical, 1.5—2 mm. long from base of the petals to filaments; traces ten, paired, prominent on the interior; filaments ca. 0.5 mm. long, reflexed, slender. Anthers 10—15, ca. 0.6 mm. long, verruculate, pale yellow. Ovary short- turbinate, multiapiculate, ca. 1 mm. high and wide. Styles ca. 2.5 mm. long, the upper half free; stigmas subcapitellate. Fruiting calyx 5—6 mm. high; lobes erect or spreading, acuminate, ca. 2 mm. wide at the base; green. Carpel 2 mm. high, 1.5 mm. radially, muticous, awned, or 68 STUDIES IN SIDA (MALVACEAE) briefly apiculate; dorsum yellowish, with rather dense dark brown ver- ruculations, invaginated slightly on the outer surface; lateral margins rugose; lateral walls membranous, pulverulent, firmly adherent to the partly visible seed near the base, a vascular trace from the columella paralleling the lower and outer margin of the wall; apex of the dorsum prolonged into two yellowish, laterally flattened, pyramidal projections from which the aristae may develop; the latter, when present, ca. 6 mm. long, yellowish, flattened, dilated basally, with few retrorse hairs; inner margin not differentiated from the lateral walls in texture; all walls except the apex adherent to the seed. Seed almost black, plump, the hilar sinus shallow, abruptly ridged on the outward edge; funicle flat- tened at the hilum, dark. Columella conoid-truncate, ca. 2 mm. high, flanged; remnant traces to. the lateral walls of the carpels sometimes present. TYPE: Specimen cultivated in the Jardin des Plantes, Paris, from seed collected by Dombey in Chancay, Peru (P). DISTRIBUTION: Western Peru. There are two distinct elements in the S. palmata group, one with 5-lobed leaves, the lobes cut approximately half-way to the base, the margins crenate-serrate, S. palmata Cav. 1785 only; and a second with 7—9 lobes, the lobes divided almost to the base, the lobes markedly narrowed toward the base, S. jatrophoides PHérit. Cavanilles and many later authors have consistently confused the two entities, taxonomically and nomenclatorially. Cavanilles emended his original description of S$. palmata, the 5-lobed element, in two subsequent Dissertations. His long feud with l’Héritier seems to have affected his usual good judgment for he sank the latter’s S. ricinoides and S. jatrophoides in his own S. palmata. It is clear from a comparison of the descriptions, the plates, and photographs of authentic material in various European herbaria (Paris, Geneva, Copenhagen, Berlin and Madrid) that S. ricinoides is identical with S. palmata. The original descriptions of both were based on material grown in the Jardin des Plantes from seed sent by Dombey. S. jatrophoides is the same as the taxon described by Cavanilles under the name S. palmata in the first Mantissa. Although Cavanilles said that his material was grown “ex seminibus Mexicanis”’, it was probably from more Dombey seed from Peru, as was the type-material of S. jatropho- ides. In the second Mantissa, Cavanilles added to the original description of S. palmata still further information from living STUDIES IN SIDA (MALVACEAE) 69 material and published a plate which is a very good match for l’Héritier’s plate of S. jatrophoides. Since Cavanilles thought both elements belonged to S. palmata and did not therefore name the second, S. jatrophoides must be used for the element with deeply 7—9-lobed leaves. Peru: Domdey, “670” (Pp), Dombey “Perot’’ ex herb. de Franqueville (pr). Dombey, ex herbs. Cosson, Maire (re). Dombey, “Chile et Peroi” (Fr). Dombey, (Photograph of ?isorypr, B: F, GH). Dombey, (Photograph of ?1soryPE, M: F). Rose & Rose 18619, vicinity of Santa Clara (us). Stork 11470, between Ambar and Huacho, Prov. Cajatambo, Lima (uc, GH). Weberbauer 7427, mountains between Moquegua and Terata, Prov. Moquegua. (Mo, US, GH, F, Ny). Weber- descr. of S. rupo). Collector unknown, (photograph of ?1soTYPE, C: GH, F). Collector unknown, (photograph of spec. B: GH, F). Collector unknown, cult. Paris (GH, ex Hb. J. Gay). 19. Sida decandra R. E. Fries, K. Sv. Vet. Akad. Hand. ser. 3, 24(2): 16. 1947. Erect annual herb. Young branches clothed in multicellular, pilose, stellate-pilose and rigid-setaceous hairs. Petioles to 7 cm. long, stellate- hirsute and setose; lamina to 10 cm. long and broad, membranaceous, concolorous; upper surface with numerous long appressed-pilose and smaller, deciduous, stellate-pilose hairs; lower surface stellate-hirsute; circumscription more or less 5-angled; base cordate; lobes 3, divided mm. long. Flowers very numerous, aggregated into rather dense oblong panicles; pedicels flexuous, erect, 3—4 mm. long, accrescent in fruiting to 15 mm. Calyx 5 mm. long, campanulate, hirsute without, divided about halfway into lanceolate lobes about 2 mm. wide at the base. Co- rolla equaling the calyx, violaceous on drying, red in life (according to the collector). Androecium 3 mm. long; stamen-tube 2—2.5 mm. long, cylindric, slightly dilated towards the base, 0.5 mm. thick at the apex, ca. 1 mm. thick at the base, with sparse, short, rigid, pilose hairs below the middle. Anthers ten. Ovary glabrous, conical. Styles ca. 3 mm. long, divided to the middle; stigmas capitate, papillose. Carpels five, truncate at the base, ovoid-trigonous, 3.5 mm. long; dorsum flattened and briefly hirsute; margins somewhat muricate; aristae 2, slender, customarily di- vergent, 12—14 mm. long, violaceous, with minute, yellowish, lax, re- trorse hairs. Seed castaneous, orbicular, flattened (not mature), 2 mm. in diameter, _ TYPE: Asplund 6797, June 1939, Ecuador, Prov, Chimborazo, Alausi, 70 STUDIES IN SIDA (MALVACEAE) Alt. 2300 m. (Herb. Regnell., Stockholm.) DISTRIBUTION: Known only from the type. I have seen no material of this species, but it is obvious from the description that it is close to S. rupo as that species is under- stood here, differing from it chiefly in the length of the aristae, the sparseness of the pubescence of the stamen column, and the shape of the leaves. With additional material, it may prove to be no more than a variety. 20. Sida rupo Ulbr., Engl. Bot. Jahrb. 54, Bbl. 117: 75. 1916. Annual herb, few-branched, erect, S—12 dm. high. Stems 10 mm. thick at the base, terete, with long-pilose patent’ hairs; the younget portions subtomentose and hirsute. Stipules not seen. Petioles 4—10 cm. long, longer than the leaves; pubescence like that of the stems. Leaves deeply 5—7-lobed, 4—5 to 8 cm. long, almost equally broad; lobes ovate, acute or acuminate, narrowing towards the base; the mar- gins irregularly subcrenate-serrate; both surfaces sparsely pubescent, with simple and stellate hairs; young leaves subtomentose; nerves 5—7, prfo- minent on both surfaces. Flowers in the upper axils and terminating the branches, forming lax panicles; pedicels 2—7 mm. long, articulated 1—3 mm. below the calyx, elongating to as much as 14 mm. after anthesis, fulvid-tomentose and sparsely villous. Calyx campanulate, ca. 5 mm. long, divided almost halfway into ovate-lanceolate lobes; fuscid- tomentose and villous on the outside. Corolla purple, campanulate or cylindric; petals obovate, glabrous, 6—8 mm. long, obtuse. Sta- men-tube conical, almost 3 mm. high, rather densely stellate-pilose, ca. 1.5 mm. long. Anthers ca. 10, ovoid, pale yellow. Ovary subconoid- globose, glabrous, subangulate. Styles five, white, 4 mm. long, 1.5 mm. free, filiform; stigmas capitate, small, glabrous. Fruiting calyx cupuli- form, enfolding the fruit. Carpels five, obliquely pyriform, ca. 3 mm. high, 2 mm. thick, glabrous, brown, rugulose-loriform, the apex with 2 aristae ca. 5 mm. long, yellow, armed with retrorse pilose hairs. Seeds obliquely ovoid, ca. 2.5 mm. high, 2 mm. thick, black, verruculose, glabrous, pulverulent at the base; the funicle indurate. TYPE: Weberbauer 5392, May 1910, between Pampano and Huaytara, on the Rio Huaytara, ca. 50 miles inland from Pisco, Peru. On stony slopes, 17—1800 m. (B). DISTRIBUTION: Known only from the type. No material which matches the description of this species has been seen. Ulbrich listed two collections, Weberbauer 5392 and 5247. Examination of several sheets of the latter number shows STUDIES IN SIDA (MALVACEAE) 71 clearly that they cannot belong to the species as described, since they have 7—10 carpels, rather than 5, and a carpel number of either 5 or 7—10 in the same species is without parellel in the genus. The specimens match S. jatrophoides well in all other particulars and are referred here to that species. If Weberbauer 5392 has 5 carpels as described, it alone should be considered the type. 21. Sida lomageiton Ulbr., Engl. Bot. Jahrb. 54, Bbl. 117: 73. 1916. Annual erect herb. Stems terete, simple or few-branched, hirsute, with spreading or retrorse bristles, double-coated above, the longer to 1.5 mm. long, stiff. Stipules linear-lanceolate, 1.5—2 mm. long, hirsute, deciduous. Petioles 0.7—3 cm. long, ca. 0.5 mm. diam., hirsute and tomentose, shorter than the blades. Leaves broadly subcordate in cir- cumscription, 3-lobed, to 5 cm. long and wide, lobed halfway; lobes Ovate-acute, irregularly subcrenate-serrate; nerves 5, palmate from very near the base, inconspicuous above, prominent beneath; the upper sur- face with scattered single and fewer few-rayed stellate hairs; the under surface densely covered with mostly stellate hairs, many of them 3-rayed, golden; the veins with dense setaceous hairs. Flowers subpaniculate, axillary and in shortened branches, appearing subglomerate when young; pedicels 2—5 mm. long, elongating after anthesis to 10 mm. or longer; articulated ca. 2 mm. below the calyx; pubescence like that of the stem. Calyx campanulate, 6 mm. long, divided halfway into acute, ovate- lanceolate lobes ca. 3 mm. Jong, 2 mm. wide at the base, three-nerved, densely pubescent, with branched and single hairs; accrescent, but not inflated in fruit, glabrous within. Petals 4—5 mm. long, ca. 2.5 mm. wide, as long as the calyx, obovate-subspatulate, purple, obtuse; apex clothed on the outside with large, scattered, stellate hairs. Stamen-tube subcylindric, glabrous, conically dilated towards the base, ca. 3 mm. high, whitish; filaments 5, very short. Anthers five, ovoid, yellowish, not dotted, 0.5 mm. long and wide. Ovary obpyriform, subangulate, glabrous. Styles five, 3.5—4 mm. long, white, the upper 3 mm. free stigmas depressed-capitate, glabrous, yellow. Fruiting calyx only slightly enlarged, enfolding the fruit except the aristae. Carpels “subclavate- pyriform”, glabrous, ca. 4.5 mm. long; the apex divided into two corni- culate aristae ca. 4 mm. long; these divergent, attenuate, retrohispid; lateral walls firmly adherent to the seed, verruculate; margins strongly transverse-rugose. Seeds filling the carpel except the apex, ca. 3 mm high, 2 mm. radially, very short-pilose on the funicle, pulverulent, dark red-brown 72 STUDIES IN SIDA (MALVACEAE) TYPE: Weberbauer 5240, Sept. 1909, vt San Augustin, nr. Lima, Peru. In the Loma formation, 300—400 DISTRIBUTION: Known only from the ie -locality. Ulbrich pointed out no distinguishing features between this and its nearest relative, S. oligandra of Bolivia, and in gross aspect there are few. The habital difference is very striking, however, and the leaf-shape of the isotype is rather distinct. Peru: Weberbauer 5240, Lima, Mt. San Augustin, Loma formation, Sept. 26, 1909, adders purple. (Photograph of TyPE, B: GH, F), (ISOTYPES: F, US, MO, GH). 22. Sida lomana Bruns, Mitteil. Inst. Bot. Hamburg 8: book. S124. Annual herb, simple or few-branched, 30—50 cm. tall. Stems terete, branching above, densely covered with a double coat of stellate and simple hairs, the latter 2 mm. long. Stipules persistent, subulate, linear, 5 mm. long, 1 mm. wide, pilose. Petioles 4—10 cm. long, densely pilose, shorter than the blades. Leaves cordate in circumscription, deeply 3—5- lobed, 10 cm. or more long, equally broad; the margins irregularly sub- crenate-serrate, 5—-7-nerved, sparsely covered everywhere with simple hairs; lobes of about equal length, ovate, acute or acuminate, narrowed toward the base. Flowers crowded in the upper axils forming dense panicles; pedicels 2 cm. long, elongating after anthesis to 5 cm., arti- culated near the apex, densely pilose. Calyx campanulate, 4 mm. long, accrescent after anthesis to 8 mm.; lobes 3 mm. long, 2 mm. wide at the base, one-nerved, hispid. Corolla pale purple, subcampanulate, ca. 6 mm. long; petals obovate, obtuse, 6 mm. long, 5 mm. wide, covered near and to the apex with scattered stellate hairs. Stamen-tube conoid, 5 mm. long, glabrous; filaments filiform, 1.5 mm. long. Anthers five, subglobose, 0.5 mm. long and wide. Ovary subglobose, glabrous. Styles five, white, ca. 4 mm. long, connate at the base; stigmas subcapitellate. Fruiting calyx accrescent around the fruit, but not inflated. Carpels five, obliquely pyriform, 5 mm. long and wide, glabrous, rugose, brown; the apex divided into two divergent and reflexed aristae ca. 14 mm. long (f. Bruns), 4—6 mm. on specimens cited; the basal portion of the aristae conoid, glabrous, 3 mm. long (f. Bruns), ca. 1 mm. on specimens; the terminal portion 11 mm. long (f. Bruns), 3-5 mm. on specimens; with yellow retrorse hairs. Seed ellipsoid, 2.5 mm. long. SYNTYPES: Giinther & Buchtien 193, 194b, Nov., 1923, Posco, Peru; 194, Aug. 1923, Mejia, Peru; 1944, Mejia, Hacienda Challascapa, Peru, Oct., 1923 (Hamburg). DISTRIBUTION: Known only from the cotype localities. STUDIES IN SIDA (MALVACEAE) 73 This species as described is very close to S. oligandra and S. lomageiton. Not having seen any authentic material, I must accept Bruns’ statements that the aristae are ca. 14 mm. long although the figures he published to illustrate the species indicate a length of 5—6 mm. Cook & Gilbert 49, collected in the same general area and matching the original description quite well in some respects, has the aristae about 5—6 mm. long. Similarly, a photograph of one of the syntypes in the Hamburg herbarium (Giinther & Buchtien 194) shows the aristae on that specimen to be never longer than ca. 6 mm. In a number of other species, full development of the aristae is not reached until very late in the maturation of fruit, and this may be the explanation of the discrepancy in measurements noted here. Peru: Cook & Gilbert 49, Posco between Mollendo and Arequipa (us). Guenther & Buchtien 194, Mejia, Lomas (photograph of sYNTYPE, H: F) 23. Sida patuliloba R. E. Fries, K. Sv. Vets. Akad. Handl. ser. 3, 24, n. 2: 15, t. 1, ff. 4—6. 1947. Erect annual herb from a slender, branching, woody root. Stems and branches terete, wrinkling longitudinally on drying, their indument of dense, soft, spreading, simple, white hairs and dense or scattered seta- ceous-stellate hairs to 1.5 mm. long. Stipules linear or linear-subulate, 2—4 mm. long. Petioles ascendent, channeled adaxially, to 5 cm. long, those of the upper leaves shorter, their indument like that of the stem, often denser, occasionally on one surface only. Leaves cordate or trian- gular-cordate in circumscription, 2—8 cm. long, and about as wide, concolorous or somewhat paler beneath, deeply three- or five-lobed; the lobes three-quarters or more the length of the leaf, obtuse to acute, nar- rowed towards the base, rather coarsely serrate; the lateral lobes of the © younger leaves at nearly right angles to the midlobe; the upper surface with many, fine, nearly appressed, bifurcate or simple hirsute hairs pointing apically; the lower surface with many, fine, mostly 3-rayed, stellate hairs, the rays whitish and nearly appressed; the base cordate; the nerves 5—7, prominent beneath, yellowish, lightly impressed above. Calyx campanulate, densely hirsute, 3—4 mm. high, accrescent after anthesis to 7—9 mm.; the lobes erect, lanceolate or long-triangular, 2.5—3 mm. long, ca. 1 mm. wide at the base. Petals obovate, slightly oblique, often purple on drying, red in life, membranaceous, fine-nerved, with a tuft of stellate bristles on the outer portion of the margin ex- posed in bud, ca. 3.5 mm. long, 2.0—2.5 mm. wide; the claw broad. Stamen-column 1.5—2 mm. high, terete, pale, translucent, with five traces, glabrous; the lower portion markedly dilated over the apex of 74 STUDIES IN SIDA (MALVACEAE) the ovary; the petals inserted about halfway up the height of the ovary; the filaments less than 0.5 mm. long, reflexed. Anthers five, 0.4 mm., pale yellow, not dotted; the valves reflexing in maturity. Styles five, 5 mm. long, ca. 4.5 mm. free; the stigmas capitate. Ovary pale brown, glabrous, five-beaked around the style-base. Carpels five, glabrous, sub- trigonous, 3 mm. high; the dorsum tuberculate on the outer surface; the apex protracted into two brief, vertical, triangular wings each bear- ing a long, extrorse, tapered arista 4—5 mm. long furnished with minute, retrorse, subulate, stiff hairs; lateral walls flat, pulverulent; dehiscence incomplete between the aristae; the walls firmly adherent to the seed elsewhere. Seed 3 mm. high, plump, dark reddish brown; the hilar depression deep; the entire surface with a thickly scattered, brown- _ ish, stellate Ameen Columella 2 mm. high, pale, usually flattened, not apparently winged TYPE: Asplund 11 242, 30 May 1940, “Peru: Dep. Lima, Matucana, 2400 m. ti d. M.” (R). DISTRIBUTION: Near Huigra, Ecuador and Lima, Peru. S. patuliloba is in many respects similar to S. lomageiton, and much of the material here cited was originally identified as that species. A study of a series of isotypes of S. lomageiton indicates that that species is quite distinct in a number of ways, the leaves being shallowly lobed, the lobes of different outline and toothing, while the slightly larger flowers are in crowded, heavily hirsute panicles rather than in the lax panicles of S$. patuliloba, Further collections from the area around Lima, Peru, may show an inter- gradation of the characters here used to separate the two as species. Macbride and Featherstone 102 from Matucana, Peru, is a well-marked leaf-form with 5 distinct lobes to the leaf. Ecuador: Rose 22414, vicinity of Huigra (us, GH, Ny). Peru: Lima: Machride & Featherstone 161, Matucana (Gu, F, Us). Pennell 14474, along Rio Chillon, near Viscas (Ny, Us, GH, F). Pennell 14597, Canta (r, GH). Machride & Featherstone 102, Matucana (Fr, GH), LEAF FORM. LA Lipertap: Lépez Miranda 392, Cerro Campana (us). Avacucno: Killip & Smith 23343, Huanta (us). 24. Sida oligandra K. Schum., Mart. Fl. Bras. 13, pt. 3: 321; 2891. Annual herbs (possibly perennial at times). Stems 7—40 dm. high, woody at the base in larger specimens, terete or slightly angled, in- dument of intermixed stellate and long-pilose hairs, becoming glabres- cent. Petioles about as long as the leaves or less, 1. ers - long, mostly 3—4 cm. long. Stipules filiform, caducous, pilose, mm. long, ca 0.4 mm. wide at the base. Leaves 3—5-lobed, pee ‘cae sae half- STUDIES IN SIDA (MALVACEAE) 75 way to the base, to 6 cm. long and wide on usual specimen; lower leaves (seldom found on specimens) much larger, to 18 cm. long and wide; lobes usually 5, variable, divided about halfway to the base to almost all the way, the latter especially in young upper leaves, lanceolate-oblong in outline, sometimes deltoid, occasionally narrowed towards the base, anthesis, divided halfway to the base into lanceolate-acute lobes; lobes 3-nerved, the marginal nerves confluent below the sinuses, but incon- spicuous under the fairly dense tomentum of 5-—6-rayed stellate hairs; the margins of the lobes clothed with stiff, tufted, 2—4-rayed, stellate hairs. Petals as long as the calyx, 5—-6 mm. long, 2.5 mm. wide, obovate, slightly oblique, glabrous at the base; the upper portion sprinkled with stellate hairs. Stamen-tube terete, glabrous, 3.5 mm. long; filaments ca. 0.5 mm. long. Anthers five, minute, 0.3 mm. diam. Ovary conical, 5-peaked, glabrous or the beaks sometimes pilosulose. Styles five, 4 mm. long, 2.5 mm. free; stigmas subcapitellate. Fruiting calyx to 8 mm. long, accrescent but not inflated, enfolding the fruit except the aristae. Carpels ive, 3—4 mm. long, trigonous; the dorsum rotund, muriculate-verru- culate; the apex eee into two divergent, reflexed, retrorsely pilose aristae, ca. 6 mm. long; base of the latter dilated. Seed ca. 2 mm. long and wide, cakescaline dark brown, almost black, adhering to the lateral walls of the carpel. Columella slender, tapering very slightly, 3.5 mm. high; flanges not prominent. TYPE: Mandon 818, near Mt. Sorata, Prov. Larecaja, Bolivia. “in se- pibus et graminosis” (B). DISTRIBUTION: Type locality only. The name S. oligandra has been used indiscriminately for any material of the section which was not of the extreme “jatrophoid” leaf-type or which had only 5 carpels. As far as I can judge from the scanty material in this section, it can be applied only to the type-material. Bolivia: Mandon 818, Prov. Larecaja, near Sorata. (Photograph of Trp, B: F, GH), (ISOTYPES: F, GH, NY 2). VI. Section Le Gray, Mem. Am. aoa 4: . 1849 (PI. Fendler.). Calyx terete ; flowers see corymbose, white; leaves palmately divi- ded; carpels gibbous, the rostrum ascendent. 76 STUDIES IN SIDA (MALVACEAE) As founded, the section was composed of two species, Sida hermaphrodita (L.) Rusby and S. Hulseana Torr. & Gray. The latter species was referred to Abutilon as A. Hulseanum Torr. by Baker in his list of species excluded from Sida, although he nowhere treats that species in his synopsis of Abutilon. The section has no close affinities with any other in the genus, nor with Napaea save in habit. Type species: Sida hermaphrodita (L.) Rusby, the only species. 25. Sida hermaphrodita (L.) Rusby, Mem. Torrey Club 5: 223. 1894. Napaea hermaphrodita L., Sp. Pl. 1: 685. 1753. Sida Napaea Cav., Diss. 5: 277, t. 132, f. 1. 1788. Erect perennial herb. Stems few or single from a woody root, terete, 1—5 m. high, glabrous or with scattered, small, stellate hairs. Stipules linear-lanceolate, deciduous, 7—9 mm. long, densely short-stellate- tomentose at the base, 3-nerved, erect or slightly spreading. Petioles terete or slightly channeled on the adaxial surface, 0.5—2 mm. diam., to 8.5 cm. long, mostly glabrous. Leaves palmately (3—) 5 (—7)-lobed, 5—7-nerved, broadly cordate in circumscription with a wide and shal- low sinus, acute or acuminate, to 24 cm. in length; lobes to 18 cm. in length, coarsely incised-dentate. Inflorescense of axillary few- to many- flowered corymbs or terminal corymbose panicles, to 5.5 cm. long; pe- duncles terete, 15—30 mm. long to point of branching, glabrous or with scattered stellate hairs, terete; pedicels 3—15 mm. long, terete, inconspicuously articulated ca. 4 mm. or less below the calyx. Calyx short- campanulate, terete, 5 mm. high, densely stellate-velutinous, without pro- minent veins, glabrous within; lobes erect, deltoid, acute, 1.5 mm. high, 2.5 mm. broad. Petals white, concave, obliquely obovate, obtuse, 8—9 mm. long, 5 mm. broad, occasionally retuse; claw moderately narrow, densely stellate-hirsute on both surfaces. Stamen-tube cylindrical above the ovary, 5 mm. long to level of first filaments, slightly 10-ribbed; the lower two-thirds with scattered to dense, spreading, flattened (col- lapsed?), lanceolate, white hairs, with a few stellate hairs at the base of the column; filaments in several series, ca. 50, 0.5—2 mm. long, the lower longer, paired or single, slender, tapering. Anthers pale, not dot- ted, 0.5 mm. wide. Styles connate, or reflexing slightly after anthesis, 7 mm. long, 5 mm. free; stigmas subcapitellate or merely rounde papillose apices of the styles. Ovary short-pyriform, 1.5 mm. high, 2 mm. diam., filling the cavity, somewhat angulate, with stellate flattened hairs on the upper surface of the angles. Carpels 8—10, one-beaked from an upward prolongation of the rostrum becoming vertical in ripe- STUDIES IN SIDA (MALVACEAE) ¥7 ning; dorsum and beak subglabrous to moderately stellate-pubescent, the hairs with fine assurgent rays; base of rostrum hooked over the top of the columella; the entire carpel 6—7 mm. high, 3 mm. radially, the beak 2.5 mm. high; lateral walls membranous, pulverulent, 3 mm. high, 2 mm. wide; dorsum thinning radially and downward; carpel dehiscent by the disintegration of the lateral walls. Seed plump, gla- brous, red-brown; rostrum ascendent; hilar sinus shallow; funicle per- sistent, flattened at the hilum. Columella conical, truncate, flanged, ca. 3.5 mm. high, usually with the base of the style persistent. TYPE: Probably specimen from Hortus Cliffortianus or Hortus Up- saliensis. (Linn. Hb., 1753). DISTRIBUTION: Lower Susquehanna R. Valley of Pennsylvania and Maryland, flats of the Potomac (escape?), valley of the Kanawha R. an tributaries, W. Va., and upper Eastern Tennessee. Cultivated as “River Mallow”, escaping but apparently not persisting. United States: PENNsyLvania: Crawford, shores of the Susquehanna R., York Furnace, York Co. (ny). Diffenbaugh, Aug. 1861, banks of Susquehanna, York Co. (Fr, GH). Durand, “Penns” (ny). Durand, 1863, banks of the Schuylkill below Philadelphia (GH). Garber, banks of the Susquehanna, York Co. (Ny 2, MO 2, F). Palmer, 1883, on the Susquehanna R., Rockville, York Co. (us, Ny). Porter, 2 Sept. 1863, banks of the Susquehanna, opposite Safe Harbor, eee. i (Ny, UC, GH). Porter, Sept. 1867, rocky bank of Susquehanna R., York Co. nate unknown, Lancaster (MO). MaryLanp: Killip 32667, Phushenecs Island, along river, , Montgom mery Co. (GH). Tatnall 4554, seas R., 0.1 mile below MO, NY 2). Vasey, 1887 (cn): West VirGinia: Brown, “Kanawha Co. fee House” (Gu). Eby, Aug. 1890, Wild Cat. (ies 2); Holton, “Virginia” (Ny). Millspaugh 616, Great Kanawha R., Kanawha Co., 1890 (ny). Maillspaugh, 1891 (ny). Millspaugh, 1892, banks of the Great Kanawha R. from Nuttallburg to the mouth (F). Tosh 1158, in drain to New River 3 miles below McCreery, 15 miles east Beckley (uc). ue unknown, “Virginia’’ (photograph of specimen of Sida Napaea Cav., M: F, US). TENNESSEE: Gattinger, June 1883, Cumberland Mts. near Kentucky line (us). paras ats July 1884, Wolf Creek near Hot Springs, Cocke Co. (us). Gattinger, July 1884, “upper east Tennessee’’ (NY). Gattinger 14 July 1884 (Ny). Gattinger, July ‘eas “upper east Tennessee” (GH). Gattinger, “upper east Tennessee” (GH). VII. Section Hooxeria Clement, sect. nov. Calyx teres urceolatus nec accrescens, lobis patentibus aetate matu- randi; epicalyx deest; folia palmatilobata; antherae 10—20; carpella mutica, indehiscentia; flores ee in axilla vel bini raro plures ex pedunculis solitaribus vel pluri Calyx terete, urceolate, not nope lobes patent i in fruit, epicalyx 78 STUDIES IN SIDA (MALVACEAE) none. Leaves palmately lobed. Anthers 10—20. Carpels muticous, inde- hiscent. Flowers axillary solitary or paired (rarely several) from solitary or several peduncles. Two species. Eastern Africa from Ethiopia to Cape of Good Hope, and southwestern Australia. The species of this section have customarily been placed in section Sida with other species having palmately lobed leaves. The combination of this character with those of the fruit and inflorescence is sufficiently striking and unlike that found in any other section of the genus to warrant the recognition of a distinct section, which is most closely allied to section Psexdonapaea of eastern North America. Type species: Sida Hookeriana Miq. ex Lehm. KEY TO THE SPECIES Annual herbs 3—6 dm. sand anthers 10—15; funicle lateral on the uppef surface of the seed; Austra 26. S. Hookeriana. Perennial subshrubs 6—12 has hia ashics 20; Honicls cpntered on the upper surface of the seed; African 27. S. ternata. 26. Sida Hookeriana Miquel ex Lehmann, PI. Preiss. 1: 242. 1844. S. leiophloia Miq. ex Lehm., op. cit. 241. S. rupestris Miq. ex Lehm., op cit. 242. Erect or decumbent annual herbs, 3—6 dm. high, essentially glabrous or with scattered small hairs, the younger parts sometimes. stellate- pubescent. Stems single or few from the base, simple or weakly branch- ing. Leaves cordate in circumscription, 3—3.5 cm. long, about as broaa, deeply divided into 3 or 5 ovate or cuneate deeply toothed lobes; upper surface with few, scattered, stellate hairs; lower surface glabrous; mat- gitis with short, subulate, contour hairs. Stipules lanceolate, 3—3.5 mm. long, 1 mm. wide, 1-nerved, erect, herbaceous, becoming patent in age, straw-colored, at length deciduous. Petioles terete, 1.5—5.6 cm. long, ca. 0.5 mm. diam. Flowers solitary or several from the axils, or 2 of more fet one peduncle, the point of branching articulated and marked by a pair of straw-colored stipular bracts; peduncles terete, 2—4 cm. long, filiform, erect; pedicels when solitary to 4.8 cm. long, when paired from a peduncle, one 1.5—3.5 cm. long, the other 0.2—1.0 cm. long. Calyx campanulate or somewhat urceolate in flower, 10-veined, 5—6.5 mm. high, pulverulent within; lobes 3—4 mm. long, 3 mm. wide, ovate or broadly lanceolate, acute to acuminate, slightly accrescent after an- thesis; intramarginal veins of the lobes abruptly confluent below the sinus, the midvein prominent to the tip; lobes patent in fruit, becoming STUDIES IN SIDA (MALVACEAE) 79 straw-colored, the veins becoming more prominent. Petals white, 6 mm. long, 3 mm. wide, obovate, glabrous except for a slight tuft at the base of the claw, inrolling after anthesis; claws narrow, separated from one another at the point of insertion on the column. Stamen-tube 2—2.5 mm. long from base of claw to filaments, glabrous, or sparsely hirtel- lous, subterete, pale; filaments very short. Anthers 10—15, 0 ‘ wide when open, ca. 3 mm. long. Styles 3 mm. long, 0.5 mm. free, re- curved at anthesis; stigmas capitellate. Ovary 0.5 mm. high, filling the cavity, 10-angled, glabrous. Carpels ten, ca. 2 mm. high and wide, form- ing a flat-topped fruit with an annular depression sunk in the hilar depressions of the seeds, muticous, rostellate; upper dorsum thick, the Outer portion thinning; lateral walls very thin, pulverulent, cancellate or rugulose, becoming rugose along the outer margins; all but the upper dorsum firmly or loosely adherent to the seed. Seed strongly trigonous, the outer surface invaginated, 1.5—2 mm. high and wide, dark brown; funicle lateral. Columella ca. 2 mm. high, truncate-conical; midveins of the carpels often persistent from its base. TYPE: Preiss 1894, Rottnest Island, near Perth, Western Australia (Hamburg). DISTRIBUTION: Extreme southwestern Australia. Australia: Western AvusTRALIA: Andrews 13, Darlington, 15 mi. e. of Perth. (k). R. Brown 5109, without locality (x). Collies, Swan River (x). Drummond, Swan River (x). Koch 1820, Wooruloo (x). Koch 1890, Lowden, Warren District (GH). Koch 1986, Lowden (x). Oldfield, Vasse River (x, 2). Preiss 1894, Rottnest Island (1soryPE, MO). 27. Sida ternata L. f., Suppl. 307. 1781. S. triloba Cav., Diss. 1: 11, t. 1, f. 11. 1785. S. permutata Hochst. ex A. Rich., Tent. Fl. Abyss. 1: 67. 1847. Subshrubs 6—12 dm. high from a woody branching root. Stems few, slender, woody at the base, subsimple or branching. Leaves concolorous, palmately 3-lobed, rarely 5-lobed, 3—7 cm. long, 3. 7? cm. — extremely variable in form, cordate in circumscription nus broadly U-shaped; nerves 5—7; margins serrate to crenate, divided foun a third to almost to the base, depth of lobing usually greater on the upper leaves; lobes deltoid to lanceolate, obtuse to acuminate, the cen- tral lobe longest, 3—7 cm. long, the lateral lobes 2.5—5 cm. long. Petioles 1.5—4.0 (—7.0) mm. long, genes channeled adaxially, occasionally deflexed near the apex, ca. 1 mm. in diameter. Stipules lanceolate, 4—5 mm. long, 1 mm. broad, geaaieed herbaceous, thin, tardily deciduous, usually glabrous. Pedicels solitary and axillary, fili- form, 15—55 mm. long, not articulated, glabrous or occasionally finely stellate-hairy when young, arching at the tip in flower. Calyx campa- 80 STUDIES IN SIDA (MALVACEAE) nulate, terete, 5.5—8 mm. long, glabrous or sparsely puberulent, gla- brous within, dull; lobes ovate or acuminate, 2.5—-5 mm. broad, 3.5—5 mm. long, erect in flower, patent in fruit, 3s nerved, on adjacent intra- marginal nerves confluent below the sinus. Petals pale yellow or whitish, barely exceeding the calyx, 5—6 mm. long, 3—5 mm. wide, suborbicu- lar to oblong, rarely retuse; claw narrow, rarely hirtellous. Stamen-tube terete, glabrous, 1.5—2 high from base of claw to filaments; fila- dotted, 1 mm. wide when open. Ovary subglobose, glabrous, ca. 1.5 mm. in diameter, occupying bie fourths of the cavity. Styles slender, 3.5— 4.5 mm. long, 2—2.5 mm. free, recurving in anthesis; stigmas dark, capitellate. Carpels (7—) ao (—13), muticous, glabrous, rostrate, forming a flat-topped fruit 3—3.5 mm. high, 3.5—4 mm. in radius; dorsum thick above, straw-colored, becoming light brown in age, thin- ning from the distal margin to the base, marked with a fine median line; lateral walls membranous, pulverulent, rarely partly adherent to the seed; dehiscent by disintegration of the thinner walls. Seed glabrous, dark dull brown; funicle flattened; hilar sinus shallow. Columella coarsely ribbed, truncate, 1.5—2 mm. high, pale. TYPE: Thunberg, Cape of Good Hope, (Linn. Herb.). DISTRIBUTION: Upper reaches of the Nile in Ethiopia; Somaliland, Kenya, ad Batter Rhodesia, to Cape of Good Hope in Union of South Afric The name se for this species. was known to Cavanilles at the time he described S. triloba, for he indicated in the descrip- tion that his might be the same plant as S. ternata, if by “foliis ternatis” the younger Linnaeus meant the type of division illustrated by fig. 64 of the elder Linnaeus’ Philosophia Botanica (p. 290), but a different plant if he meant that shown by fig. 65, in which the lobes are divided all the way to the base and petiolate. Since the lobes, though often divided to the base, are never petiolate, most authors have taken up Cavanilles’ names, although Thunberg listed both species in his Flora Capensis. I have restored the name given by Linnaeus f. despite the brevity of his description: “foliis ternatis, foliolis serratis. Habitat in Cap. bonae spei. Thunb.” since it unmistakably applies to this species. Specimens distributed by Hochstetter from Schimper’s Abyssin- ian collection bore label names, one of which was validated by A. Richard in his Tent. Fl. Abyss. as S. permutata Hochst. ex A. Rich., with the other in synonymy. I can find no constant STUDIES IN SIDA (MALVACEAE) 81 characters to separate these plants of Ethiopia from S. ternata. They appear to be only the terminals in a series of very slight geographic variations, having a greater number of 5-lobed leaves, the carpels somewhat more numerous, and the petals (reportedly) rose-purple. The specimens cited by Richard for S. permutata were Schim- per 1911, Schimper 7, and Dillon & Petit (without number). I have seen examples of the first two, and Schimper 7 fits the description, whereas Schimper 1911 (Gu) is a Pavonia. Apparently Schimper 364, the type no. of Pavonia Kraussiana Hochst. (fide Ulbrich in his Monograph of the African species of Pavonia, p. 126), became mixed with Schimper 1911 in some cases, for Ulbrich in the same work, (p. 180) refers Schimper 364 to S. triloba. To complete the confusion, Schimper 364 was distributed with the label name Urena mollis. The Kew specimen of Schimper 1911 bears a note in Hochreutiner’s hand that the labels should be exchanged, and a note in another hand that they had been, although what the original label was is not stated. There is a form of this species in Kenya which has 9—13 carpels, the walls of which are rugulose and the seeds with a deeper hilar sinus. Since I know it only from one specimen (Curtis 721, 10 Jul. 1932, “Buffalo Country”, Kenya Colony, British East Africa, in GH), I refrain from naming it. In all other characters, it comes within the range of variation of the species. Ethiopia: Gillett 5163, Galla Pass (x). Schimper 7, Tut-Adegi, Scholoda, (uc, K, GH). Schimper 401, Scholoda (us, uc). Schimper 1911, Scholoda, 4 Oct. 1842 (k). Roth 101, Ankober (x). Eritrea: Schweinfurth & Riva 911, Gorge de Degorra (K). Kenya: Baily 7699, Chyulu South (xk). Curtis 721, in Buffalo country (GH). Fries & Fries 695, “occid.: in siloc montana, loco aperto, 2300 m.” (x). Hill 649, upper Baca | (x). van Someren 2458, Nairobi Dist. (xk). Wyte, Oct. 1898, Eldama ravine (x). Uganda: Maitland 1002, (x). Chandler-Hawok 2610, Kirwa Kigezi (xk). Rogers 353, Kigezi District, edge of Lake Chibapi (x). Tanganyika: Burtt 4364, Olomoti Volcano, gorge of Munge River (x). South Africa: Transvaa.: Rogers 23817, Irene (us). Gulfin 994, Rimer’s Creek, Barberton (k). Nata: Rogers 504, Port Shepstone (Kk). ORANGE FREE Kaffraria (x). Cooper 563, Dist. Drege 7331, Somerset East Div. (x). Gill, Fish River (x). Kuntze 94 (x). Long 978, Addo Heights (x). MacOwan 301, near Grahamstown (cu 2). Prior, Nov. 1847, on Uitenhage Karroo (x). Thode A2601, Enon, (x). 82 STUDIES IN SIDA (MALVACEAE) VIII. Section Strenrnpa Grisebach, Fl. Brit. W. Ind. 76. 1859. Calyx-tube subterete; carpels briefly two-beaked, indehiscent; flowers in terminal corymbs; leaves entire, linear to lanceolate. One species: S. linifolia Cav. The section was founded by Grisebach (1859) for the single species in the following words, “Calyx 5-fid, tube subterete. Carpids bluntish, indehiscent. — Flowers in terminal corymbs. Leaves quite entire.” Schumann (1890) maintained the section as founded, but Baker (1892) changed the concept completely, omitting the essential characteristics “leaves quite entire” and “flowers in terminal corymbs”. In their place, he established a series of negative criteria: “calyx non angulatus post anthesin non accrescens.... carpella indehiscentia vel fere indehiscentia.... nunquam aristata.... bracteolae O.... flores petiolis haud adnati.” The result of this series of exclusions was that the section, sensu Baker, was now available for any number of species other- wise difficult to place in the genus. While he did not take full advantage of the opportunity thus afforded, he did include ten species from Australia and one from New Caledonia. None o these has the leaves entire, eight have the flowers solitary or subsolitary, and all have only generic affinity with S. linifolia. The closest species to S. linifolia in the morphology of its flowers and carpels seems to be S. blepharoprion Ulbr. in which, however, the flowers are strictly solitary from the upper leaf- axils and the calyx strongly angulate. It is most satisfactorily placed in section Sida. 28. Sida linifolia Cav., Diss. 1: 14, t. 2, f. 1. 1785. Annual or biennial herb, erect from a strong fibrous root-system. Stems terete, 2—10 dm. high, 2—5 mm. diam. at the base, few-to many- branched; branches virgate; entire stem covered with simple, forked or stellate hairs, glabrescent, hispidulous or long-villous. Stipules linear or narrowly lanceolate, 4—7 mm. long, 0.5—1.0 mm. wide, entire or sef- rulate at the bases of the regularly spaced, patent or subpatent, pilose hairs, these present also on the prominent midnerve. Petioles 3—6 mm. long, flattened adaxially, sparsely pilose. Leaves (2—) 4—6 (—11) cm. long, (2—) 3—10 (—15) mm. wide, linear to lanceolate, long-acumi- nate, usually broader below the middle, minutely mucronulate, quite entire, rounded or truncate at the base or rarely narrowly cordate, 3- nerved, the lateral nerves marginal, joined by the secondary veins from STUDIES IN SIDA (MALVACEAE) 83. the midnerve; indument like that of the stem, usually denser beneath, appressed or partially so. Inflorescence of 8—12 (—20)-flowered co- tymbs, the upper ones usually congested, the lower laxer; occasionally solitary from the upper axils in addition. Peduncles terete, 6—12 cm. long, patent-pilose, articulated 2—3 mm. below the calyx, appressed- pilose above the articulation; bracts subulate, acute, caducous. Calyx campanulate, subterete, ten-nerved, 4.88—7.0 mm. high; indument hir- sute; lobes erect in flower, partly enclosing the fruit, triangular, acute, 2—4 mm. high, 2.5—3 mm. broad at the base, with a prominent mid- nerve; interior glabrous except the margins, straw-colored in fruit, shining. Petals 8—15 mm. long, 4—5 mm. wide, obovate, white or yellow; base reddish, purple, or nearly black; quite glabrous. Stamen- tube subterete, ca. 2 mm. long, glabrous near the base, becoming histri- cine near the summit, the hairs subulate, hyaline, simple; base broadly dilated; filaments slender, paired, 0.5—1.0 mm. long, pale. Anthers numerous, small, 0.5 mm. long or less, pale, not dotted. Styles 5—6 mm. long, ca. 4 mm. free, slender, reddish; stigmas capitellate, dark red, minutely hispidulous. Ovary 1 mm. diam., 0.6—0.8 mm. high, some- what lobed; upper surface with an annular depression; apex multiden- ticulate. Carpels five to nine, 2.5—3.5 mm. high, 1.6—2.0 mm. radially, rotund-trigonous, dull brownish black, papyraceous; apex with two tri- angular, blunt, divergent beaks; lateral walls flat, very thin, fragmenting in 7 the seed thus escaping, cinereous-pulverulent. Seed 2 mm high, ca. 1.7 mm. radially, dark brown, dull, the sides subimpressed, the hilar depression rather deep. Columella 2 mm. high, persistent, stellate in section, tapering slightly, pale. KEY TO THE VARIETIES Stems glabrescent to hispidulous; leaves linear to narrowly lanceolate, 30—110 mm. long, 2—12 mm. wide; petals 8.0—9.2 mm. “rs calyx-lobes 2.0—2.5 m. high; carpels 5—9 . 28a. S. linifolia var. linifolia. Sem ead rest of plant ville: leaves 25—60 mm. long, 7—15 mm. ie lower es broadly lanceolate; ete ca. 15 mm. hie calyx lobes ca. 4.0 mm. high; we usually G6. . vi tere 28b. S. linifolia var. brevis. 28a. Sida linifolia var. linifolia. S. graminifolia A. Rich., Act. Soc. Hist. Nat. Paris 1: 111. 1792. S. viminea Fisch. ex Link, Enum. Hort. Berol. 2: 202. 1822. S. campi Vell., Fl. Flum. 278. 1825. S. linearifolia Thonn. in Schum. & Thonn., Beskr. Guin. Pl. 2: 77 (303). 1827. S. linifolia var. angustifolia St. Hil., Fl. Bras. Merid. 1: 181. 1827. 84 STUDIES IN SIDA (MALVACEAE) S. angustissima Migq., Stirp. Surin. Sel. 102. 1850, non St. Hil. S. longifolia Brandegee, Zoé 5: 212. 1905. S. linifolia f. flaviflora Chod. & Hass.; Bull. Herb. Boiss. ser. 2, B92: 1905: S. Fiebrigii Ulbr., Engl. Bot. Jahrb. 54, Bbl. 117: 72. 1916. TYPE: J. de Jussieu, “In insula Caienae & in Peru.” (P). DISTRIBUTION: Antilles; Mexico to Brasil, Paraguay and Bolivia; Sierra Leone to nothern Angola and Tanganyika; Fiji. 4s 28b. Sida linifolia var. brevis (Hochr.) Clement, comb. nov. S. Hassleri Hochr., Ann. Cons. & Jard. Bot. Gen. 6: 33. 1902. S. Hassleri var. brevis Hochr., lic. S. linifolia var. Hassleri (Hochr.) Hassl., Add. Pl. Hassl. 15. 1917, TYPE: Hassler 4728, “near Igatimi’, Sierra de Maracayu, Paraguay DISTRIBUTION. Northeastern Paraguay. On the basis of two Hassler collections from Paraguay, Hoch- reutiner described S. Hassleri and its variety brevis. Hassler later considered these as no more than specimens of a rather broad- leaved variety of S. linifolia which he called S. linifolia var. Hassleri (Hochr.) Hassler. The earliest available varietal name in S. linifolia for this taxon is brevis, however, and I have made the combination here. This variety, found at the southern limit of the species, is connected to the typical by such intermediate specimens as Jorgensen 4586 and Fiebrig 572, both of Paraguay and the latter the type-collection of S. Fiebrigii Ulbrich. The characters given by Ulbrich to distinguish it from S. linifolia var. brevis (which he recognized as S. Hassleri) merely place it as intermediate between that and the typical variety. His statement that the calyx-lobes are only 2 mm. high is not confirmed by actual measurements on the isotypes or on the photograph of the type. The species is remarkably consistent in the remainder of its very extensive range, even the African material being very little different from that of eastern South America. Schumann and Arechavaleta have stated that the species occurs in Uruguay and was collected there by Tweedie. No material from that country was known to Rodrigo (1944), nor have I seen any. The only Tweedie specimen I have seen is from Rio de Janeiro, Brasil. STUDIES IN SIDA (MALVACEAE) 85 The citation of the author of the species as “Cavanilles” rather than “Jussieu” is the result of a survey of the general practice of the former in the Dissertationes. He almost invariably cited the person who loaned him the material upon which he described his new species, and the citation is an indication of the type- specimen rather than of authorship. Cavanilles’ practice was to use the herbarium or field notes attached to specimens in framing his descriptions, but these, even when quoted and attributed, as in the case of the specimen loaned by de Jussieu, are never the first and decisive paragraph, which is always Cavanilles’ own description and is the one on which the species rests. 28a. Sida linifolia var. linifolia. (About one-half of the New World material I have seen is listed here. Because of the great uniformity of the species throughout most of its range, only enough specimens to define the distribution have been listed for Mexico, Central America, Colombia and Venezu Mexico: Cuiapas: Matuda 3777, Aeviciie Palenque, (Ny, F, GH, MO). GUER- RERO: Hinton 11397, Vallecitos, Dist. Montes de Oca (Ny, GH). Palmer 279, Acapulco and vicinity (us). Jatisco: Pringle, 6 Sept. 1893, Guadalajara (mExv). Pringle 5440, Tequila (mexu 2, GH). Mexico: Hinton 2239, Ixtapan, Dist. of Mo). Vera Cruz: Matuda 1442, Cuitléboad (x60); Mell, 28 Nov. 1928, Minatitlén (us). Purpus 1947, Zacuapan (uC, NY, GH, US, MO). Guatemala: Heyde & Lux 3949, Chiapas, serie Santa Rosa (Mo, US, NY, GH) Standley 24519, vicinity of Quirigua Dept. de Izabal (cu, ny, us). Standley 74930, Dept. Jutiapa, vicinity of Jutiapa (F). rev 76798, Dept. Jalapa, northeast of Jalapa (F). Standley 80471, Dept. Guatemala, near Fiscal (F). Steyermark 30610, Chiquimula, Volcan Ipala, near Amatillo (Fr). Steyermark 42194 F British Honduras: O'Neil] 8770, Sibun Road, 9 miles southwest of Belize (GH). Peck 243, Pine ridge near Manatee Lagoon (cH). Lundell 11, Honey Camp, Orange Walk (F). Gentle 1362, Maskall Pine Ridge (mo, Us, NY 3 kilometers west of Guinope, Dept. El Paraiso (uc). Yuncker, cla & Youse 6034, near Siguatepeque, Dept. af Comayagua (NY, US, GH, MO. El Salvador: Calderén 978, near Chalchuapa (Ny, US, GH). ‘Caleta 2472, El Roblar, Dept. eneciva (ny, US, F). Standley pith Serb of Santa Ana, Dept. de Santa Ana (us, Costa Rica: ac prec so ecm de Poas” (Fr). Brenes 17302, “entre los rios Pilas " Tacaros” (Fr). Ridley 7098, near Saymateo (us). ‘anama: Davidson 752, sas uete, Chiriqui Prov. (F). Erlanson 288, San José Island, sais Archipelago (us). Pittier 3595, Taboga Island, Gulf of Panama 86 _ STUDIES IN SIDA (MALVACEAE) (us). Pittier 4596, near Cheop, Prov. of Panama (us, ny). Standley 26298, near Punta Paitilla, Prov. of Panama (us). Standley 26320, Ancon Hill, Canal Zone (us). Standley 28177, Rio Tapia, Prov. of Panama (us). Standley 29161, between Fort Clayton and Corozal (us). Cuba: Shafer 11707, Prov. of Pinar del Rio, Los Palacios to Herradura (Ny, MO, Us). Baker 2110, Herradura, Prov. Habana (pom). Combs 595, Prov. of Santa Clara, Dist. oe seats (Ny, MO, GH). Curtiss 279, near Nueva Gerona, Isla de Pinos (Ny, Mo, Gu, US). Ekman 9958, Prov. Oriente, Sierra de Vije, on the Lojo trail ae Shafer 542, Sierra Cubitas to Santa Rosa, Camagiey Prov. Ekman H. 9417, St. Michel de I'Halaye (us). Holdridge 1862, La Victoire (us). Leonard 8644, vicinity of Dondon, Dept. du Nord (us, ny, GH). Dominican Republic: Abbott 676, Pimentel, Hios. Pacificador (us). lard 13066, — of Piedra Blanca, Prov. of La Vega (us). Jiménez 1055, El Rubio, Prov Santiago (us). Valeur 510, Dist. iy Exbeteta Prov. of Monte Cristy (Mo 2, ee F). Martinique: Ste+/é 472, Monte-Bello, Petit Bourg (ny). Tobago: Broadway 9052, “Botanic Station, wild” (mo). Eggers 5474, near Bacolet (us). Trinidad: Broadway, 12 June 1908, St. Margarete (ny). Broadway 2565, Yoco, roadside to Seis Point (Mo 2 mbia: Daniel 4170, Medellin, parionnis (us). Killip 35598, Dept. El Valle, Cisneros (us). Killip & Smith 16304, Dept. Santander, northwest of Bucaramanga (GH, US). er 3464, Dept. of Tolima, west of San Lorenzo MO, GH, US, NY). Sneidern 245, Dist. Cauca, El Tam bo (us). Veebic ke Killip 37581, “Bolivar: La Paragua” (us) Pittier 12485, Mesa de El Sombrero, Guarico (us, ny). Pittier tee. Carabobo, Carabobo (us). Pittier 9818, Federal Dist. aed Caracas (Gu, NY). Pittier 14529, State of Anzodtegui, Pariaguan (us) uiana: Jenman 22, Ipelemanta, Atapoo R. (us). Sandwith 1227, Mazaruni Station (ny). Schomburgk 131, near Anna-y (us). Dutch Guiana: Hostmann 816 (cu). French. Guiana: Broadway 191, vicinity of Cayenne NY, US, GH). Melinon 217, Maroni, 1876 (ny, F, US) rasil: Barreto 8045, Capitio Eduardo, Belo Horizonte (Fr). Barreto 8056, Lagoa Santa, Santa Luzia (F). Carvalho 652, ss Sao Paulo (cu). Dahlgren, Ceara in. Drouet 2036, Belém, Para (us). Drouet 2216, Fortaleza (Mo, F, ; . Hoehne 286, Butantan, Sao Paulo nate Mexia 5594, Corinto, Mings eae, (us, MO, GH, NY, F). Monteiro C. 1002, State of Para, Belém, Souza (us, F). Pickel, Sept. 1931, a ig asc (ps, us, Gu, F). Pehl 317, Mt. Corcovada (GH, F). Smith, L. 7108, Araruama, Rio de Janeiro (us). Tate 128, Limao (Ny). Tweedie 131, ge A leewind (GH). Paraguay: Fiebrig 572, Cordillera de Altos (photograph of type of S. Fiebrigii, B: GH, F, US), (IsoTYPES, GH, F). Fiebrig 4721, “N. Paraguay. Zwischen Rio Apa und Rio Aquidaban” (Gu). Jérgensen 4586, Santa Barbara (Ny, Us, DS). Peru: Williams 7493, San Roque, Dept. San Martin (r). Williams 5847, Dept. San Martin near Tarapoto (Fr, us). Bolivia: Cardenas 2699, Reg. Oriente, Rio Grande (F). Steinbach 5331, Buena bi tien Sara, Dept. Santa Cruz fm, F). Rusby 1314, San Pedro, vicinity Reyes Sierra Leone: Brown & Brown 64, iFivciows (us). \Deiphiow D4?) ebetordn < STUDIES IN SIDA (MALVACEAE) 87 (Kk). Glanville 145, Makump (x). Thomas 3804, Bumbuna (x). Gold Coast: Brown 380, Accra (x). Kitson 1166, Tafo (x). Togo: Warnecke 267, rear Lomé (xk). Nigeria: Le/y 279, Naraguat (x). Newberry 98, Ibadan (x). Vogel, 1843, Secorra (k). French Equatorial Africa: Debeaux 186, Libreville, Gabon (x). Le Cestre 4218, Yalinga (Ht. Oubangui) (x). Angola: Monteiro, Jan. 1873, Boma (xk). Fiji Islands: Wilkes Exped., 1838-42, Ovalau (Gu, Us). 28b. Sida linifolia var. brevis. Paraguay: Hassler 5738, “In mits Sete Yeruti”, Sierra de Maracayi. Kiidioatanh of type of S. Hassleri var. genuina Hochr. G: F), (ISOTYPE, GH). Hassler 4728, near Igatimi, Sierra de Maracayts (isotype of S. Hassleri var. brevis Hochr., GH). Jérgensen 4852, “in the campo, Estancia Primera (us). LITERATURE CITED The following bibliography completes the year-references given in the text. The dates of publication cited, whenever in disagreement with the title-page dates, have been taken from many sources. Of particular help have been the many articles on bibliography in the British Journal of Botany by Britten, Woodward and others, and the system of entering references to pertinent bibliographic literature on the fly-leaves or title- pages of books in the Gray Herbarium. Specific references to the liter- ature which appear after scientific names are not repeated here. ADANSON. 1763. Familles des Plarites 2: 398. Baker, E. G. 1892. Synopsis of Genera and Species of Malveae; XVIII. Sida. Jour. Bot. Brit. & For. 30: 137—142, 235—240, 290—296, 324—332. ——. 1937. (Review of Monteiro’s mondgraph of Brasilian Sida). op. cit. 75: 116, BentHaM, G. & F. MUELLER. 1863. Flora Australiensis 1: 197. Biack, J. M. 1952. Flora of South Australia, ed. 2, 3: 557—5 Britron, N. L. & P. Witson. 1924. Botany of Porto Rico a the Virgin Islands 1: 550. DOLLE, A. P. 1824. Prodromus 1: 459—474 CavaniLtes, A. J. 1785. Dissertatio Botanica de ‘Sida (Monadelphia Classis Dissertationes 1; 1—47, t. 1—13 ——. 1799. Icones et Descriptiones Plantarum 5: 10. Drerricu, D. 1847. Synopsis Plantarum 4: 841—859. Don, G. 1831. General System of Gardening and Botany 1: 498. Fries, R. E. 1907. Einige neue phanerogamen aus der Siid- und Central- Amerikanischen Flora. Bull. Herb. Boiss. ser. 2, 7: - 1947. Zur kenntniss der Siid- und Zentral- Acalaschen Malvaceen- flora. Kgl. Sv. Vetensk. Handi. ser. 3, 24(2): 1—37. GAERTNER, J. 1791. De fructibus et seminibus plantarum 2: 249—251 GaGnepain, F. 1909. Essai d'une classification des Sida asiatiques. Notulae Systematicae 1: 27—32. Garcxe, A. 1896. Uber einige Malvaceengattungen. 1. Sida. Engl. Botan. Jahrbiich. 21: 379—386. ' me AEE og 88 STUDIES IN SIDA (MALVACEAE) Gray, A. 1849. Plantae Fendlerianae. Mem. Am. Acad. 2(4): 23. 1852. Plantae Wrightianae I. Smithson. Contr. Kn. he ye 1887. Revision of some polypetalous genera ...... Pro sly hea 22: 294—295. 1897. Synoptical Flora of North America (ed. B. L. Robinson) 1: 26. GREENE, E. L. 1906. Certain nee types. Lfl. Bot. Obs. and Crit. 1: 209. GRISEBACH, A. H. R. 1859. Flora of the British West Indian Islands 76—77. —. 1874. Plantae teas a Hassuer, E. 1907. antae paraguarienses. Bull. Herb. Boiss. ser 1) 120. . 1910. Novitates paraguarienses ex herb. Hassler., Malvaceae ot das Rep. Spec. Nov. 8: 28—31 HeMsLeEy, W. B. 1879. Pn Centr.-Amer., Bot. 1: 104. Hircucock, A. S. & M. L. Green. 1935. Species lectotypicae generum Linnaei. Int. Rules Bot. hana ed. 3, Suppl. II. 1: 139—143. HocurevTINer, B. P. G. 1902. Malvaceae nov. vel minus cognitae. Ann. Cons. & Jard. Bot. Gen. 6: 33. ———. 1920. Organes carpiques nouveaux ou méconnues chez les Malvacées. op cit. 21: 347—387. Hu, S.-Y. 1955. Flora of China, Malvaceae. 13— Hurcuinson, J. B. and DauzreLit. 1928. Flora e ‘ia Tropical Africa 1(3): 261. Jacxson, B. D. 1912. Index of = Linnaean Herbarium. Proc. Linn. Soc. London, sess. 1911-1912, Suppl., JauBERT, H. F. & E. Spacu. 1855. fi tr. Pl. Orient. 5: 47. eo - H. 1951. The American Genera of Kiifvaceds. Am. Midl. Nat. -~—-. ey Tentative site to the North American Species of Sida. Leafl. West. Bot. 7(6): 138— Kuntu, K. S. ae ae pS: et eR Plantarum: 254—270. Mepicus, F. C. 1787. Malvenfamilien. 158 pp. 1789. Phil. Bot. 1: 28. Mercatre, C. R. and L. Cuaik. 1950. Anatomy of the Dicotyledons. Miiier, P. 1754. Gardener's satay Abr. 4th ed., vol. 2. . 1768. Gardener's Dictionary, 8th ed. MigueL, F. A. W. 1859. Flora van NEL Indié (Fl. Ind. Bat.) 1(2): 138—143. Moencu, K. 1794. Methodus plantas horti ...... Marburgensis 618—621 oo Fituo, H. pa C. 1935. Monographia das Malvaceas Brasileiras, Fue. . O Genero Sida. 56 pp. . 1949. As especies mrpeonaes ssa e uruguayas da seccio Malvinda do genero Sida. Lilloa 19: 501—52 MUELLER, F. 1860-62. Plants poms to the Colony of Victoria 160—163. RAFINESQUE, os S. 1836. New Flora of North America 1: 41. Roprico, A. 1937. mor nueva especie de Malvacea Argentina. Not. Mus. de La Plata % ‘Bot. 14: 1944. Las. pete Argentinas y Uruguayas del genero Sida. Rev. Mus. ie La Plata ser. 2, 6: 81— ~ STUDIES IN SIDA (MALVACEAE) 89 RoxsurGH, W. 1814. Hortus — 50. 1832. Flora Indica 3: St: Pa ashe! A. DE 1827. og ne Merid. 1: 173—191. 19 (repr. of 1824). Esquisse de mes voyages. Chronica Botanica 10. ST: Hinvarnr, A. & Cu. Naupin. 1842. Revue de la flore du Brésil méridional. . Nat. Paris ser. 2, 182 51. bev ace. e 1945. Catalogue of the Linnaean Herbarium. Linn. Soc. London, special publ. p. 120. ScHUMANN, K. 1890. Sida. Engl. & Prantl, Nat. Pflzfam. 3, Abt. 6: 30. 1891. Sida. Martius, Flora oo Ran 12(3): 284—336 Sxssi and Mocimo. 1889. Plantae ei e Hispaniae 108—110. 1894. Flora Mexicana ed. 2: Naar ee P. C. 1924. Trees and Pate of Mexico. Contr. U.S.. Nat. Herb. 23: 746—786. SPRENGEL, K. 1826. Systema Vegetabilium 3: 109—123. STEUDEL, E. G. 1845. Fleischeria. Lehmann, eal Preissianae 1: 236. Ursgan, I. 1920. Symbolae Antillanae 8: 413—418. Wicut, R. 1837. Dictyocarpus. Madras J. Lit. Sci. 5: 309. 1cHT, R. & G. Arnott. 1834. Prodromus FI. Ind. Or. 66. WiILLpENow, C. L. 1801. Species Plantarum 3(1): 755. News NaAMEs AND TAXA Hookeria, sect. nov.; Incanifolia, sect. nov.; Oligandrae, sect. nov.; Sida centuriata, sp. nov.; Sida ciliaris var. involucrata, stat. nov.; Sida Brownii, nom. nov.; Sida Grayana, nom. nov.; Sida leprosa var. depau erata, comb. nov.; Sida leprosa var. sagittaefolia, panes nov.; Sida saiteleg var. brevis, comb. nov.; Sida Standleyi, sp. nov. INDEX TO NAMES Abutilastrum (section) 9, 10, 15 Dendrinda (section) o Abutilinae 5 Diadesma 7 Abutiloides (section) 6, 7, 15 Dictyocarpus io Ge Abutilon (genus and_ section) Disella 10, 14, 48 6, 7, 8; 15; 15 cuneifolia 61 Hulseanum 7 hederacea 52 noda . 6 7 lepidota 53 astardia 6 7, 1035 sagittaefolia 53 Bastardiopsis (genus and section) Eusida (section) 8, 15 5 40, 15 Fleischeria 7,43; 82 Bogenhardia 5 pubens 41, 42 Bouchetia 33 Gaya 6, % 15 Briquetia denudata 8 Hookeria (section) 16, 77 Calyxhymenia 9. 10, 15, 30, 31. Incanifolia (section), 10, 16, 60 Cristaria 9 Lamarkia 6&1 Cuphea 33 Lawrencia (section) a 15 Daucus 33 Malachra, S, 27,24 90: Malacroideae (section) 7, 9, 15, 16 californica 52 cymbalariaefolia 48 hederacea 52 leprosa 52 obliqua 52 licata a2 Sherardiana 48 sulphurea 52 Malveae 5 Malvastrum 9 linearifolium 25 sulphureum 52 Malvella 8, 14 = area mneaEs 49 Sherardi 48 Malvinda ra and section) 5 3,9; 10) 15. 35 palmata 5 Mentzelia 33 Melochia truncata 62 Napaea hermaphrodita 76 Notho-Plagianthus (section) 8, 15 Nymphaea 14 Oligandree ces 16, 64 Pavonia Kraussi 81 Periptera 6 Physalodes 8, 9, 10, 15, 16, 30, 31 Plagian 15 pulchellus 8 Pseudomalachra (section) 9, 10515; 16.,247 Pseudomalvastrum (section) dy 8)"9, 40,45, 16:47, 60 Pseudonapaea (section) 7, 15, 16, 75 Riedleia 7 truncata 62 Robinsonella 9 Scorzonera 42 ida 13 acuta 5 alnifolia 5, 14 angustissima 84 nomala 17, 24, 230 26. f. albiflora 25 f. roseiflora 25 « INDEX TO NAMES f. violaceiflora blepharoprion Brittonii Lis Brownii 32; calyxhymenia 7, 31, 32, 40, var. ferruginea campi centuriata Lars iliaris ti17; 18, 20;-22, var. anomala 23, 24, var. fasciculata var. fulva 23, 24, 26, var. guianensis 18, 24, var. involucrata 23, 25, 26, var. mexicana 20, 25, 26, f. flava cleisocalyx Clementii cordifolia cuneifolia 5,2, 48; 61, cymbalaria 36, f. viridescens decandra 65, decumbens densiflora denudata divergens Eggersii Elliottii arosa fasciculata Fiebrigii falvescens 32, 38, 39, 40, 44, 45, ulva ty graminifolia Gray 7 Hassleri var. brevis hastata 31, var. —— 32, var. tomentosa 32, ARG 7, 41,248; var.? parvifolia 48, var. sulphurea Helleri 8, 48, rocrucmttaai ay Hooker Hulseana humilis sce 65, leiophloia lepida lepidota var. depauperata var. sagittaefolia leprosa var. depauperata var. cymbalaria var. intermedia patuliloba INDEX TO NAMES 66, 67, 68, 64, 65, sesincclios perm 13; eee a 37, physocalyx 8, 31, 32, 33, 34, 35, platycalyx pl 17, pros 11, 44, 45, var. flavescens var. genui quinquenervia rhombifolia 75 425-14, Ricini ricinoides rubifolia ssp. pseudocymbalaria rupestris rupo sagittaefolia Schimperiana _ Sherardiana 8, 47, spinosa % 1245; Standleyi 32, 43, sulphurea 7, AT; surumuensis rs; ternata 79, tridentata ilob 79, 80, truncata urticaefolia 32, 39, nea Sidastrum (genus) 9, Sidastrum (section) 9; Sidinae Steninda (section) 8, 9, 10, 15, 16, Thyrsinda (section) 9:10, 15; Urena mollis Urenae Wissada (section) 8, Wissadula J. Oy Bs Balansae CONTRIBUTIONS FROM THE GRAY OF HARVARD UNIVERSITY Edited by Reed C. Rollins and Robert C. Foster NO. CLXXXI INTERSPECIFIC HYBRIDIZATION IN LESQUERELLA (CRUCIFERAE) Se Reep C. RoLins A REVISIONAL STUDY OF THE GENUS ‘MAR E ATTY eee By RB. Cuannece THE GRAY HERBARIUM OF HARVARD _ CAMBRIDGE, MASS., USA. CONTRIBUTIONS FROM THE GRAY HERBARIUM OF HARVARD UNIVERSITY Edited by Reed C. Rollins and Robert C. Foster NO. CLXXXI INTERSPECIFIC HYBRIDIZATION IN LESQUERELLA (CRUCIFERAE) By Reep C. ROLLINS A REVISIONAL STUDY OF THE GENUS MARSHALLIA (COMPOSITAE) By R. B. CHANNELL PUBLISHED BY THE GRAY HERBARIUM OF HARVARD UNIVERSITY CAMBRIDGE, MASS., U.S.A, 1957 Issued April Ist. vs INTERSPECIFIC HYBRIDIZATION IN LESQUERELLA (CRUCIFERAE) ReEep C, ROoLLins The results of initial studies on naturally occurring populations of hybrid plants of Lesquerella were presented before a session of the Eighth International Botanical Congress in Paris (Rollins, 1954). At that time, data had been accumulated that pointed unmistakably to the involvement of L. densipila and L. Lescurii in the hybrid origin of the several large populations of plants found along the flood plain of the Harpeth River in middle Ten- nessee. However, experimental proof of the compatibility of these species had not been fully obtained and there were a num- ber of questions about the geographical relationships both of the species and the hybrid populations that required further field work and further study. In screening measurement data froin samples of two populations of plants from the Stones River flood plain, previously identified as L. densipila var. maxima (Rollins, 1952), I was impressed by the wide range of variation present. By using Fisher's discriminant function and an analysis of vari- ance,’ it was readily shown that these populations, .if they were the result of interspecific hybridization, could not have been de- rived from any pair of known species from central Tennessee. However, the probability of L. densipila being one of the parental Species was demonstrated. Thus, questions regarding the possi- bility of a second series of hybrid populations along Stones River remained open and a further complication was introduced, name- ly, the question as to whether there was yet another undetected species of Lesquerella in the Central Basin of Tennessee. A second puzzle arose from the failure in 1953 to find Lesque- rella densipila and L. Lescurii coming completely together in the area where their ranges appeared to approach each other most closely. At that time, there had been certain deterrents, such as inaccessibility by roads and the lack of time, that left open the question as to how and where cross-pollination takes place be- tween these two species. It was highly desirable to conduct a further search of the upper Harpeth River and Arrington Creek areas for additional populations of L. densipila, of L. Lescurii and of their hybrids. In the spring of 1955, I returned to middle Tennessee with ne am indebted to Dr. R. C. Foster for having carried out these statistical pro- cedures, 4 REED C. ROLLINS several tasks in view. The first and major objective was to locate the exact place where L. densipila and L. Lescurii come together, or, alternatively, to rule out the possibility of an actual point of contact of the ranges of the two species. A second objective was to find out what insects were pollinating Lesquerella in the area. Such information would be of importance especially if the species were found not to come into contact areally. Thirdly, it was needful to determine more accurately the geographical distribu- tion of both L. densipila and L. Lescurii, especially in certain critical areas such as that mentioned above and in the upper drainage area of Stones River. Furthermore, there was a need to search for an unknown species of Lesquerella. If such a spe- cies existed, as there was reason to believe, it was most likely to be found on the upper part of Stones River. Answers to several of the questions remaining from previous studies were provided by materials gathered and information obtained as a result of the 1955 field work. MATERIALS AND METHODS The present study combines an examination and evaluation of field populations of Lesquerella occurring in the Central Basin of Tennessee and the results of experiments with samples of these same populations in the greenhouse in Cambridge, Mass. Our concentrated effort has been toward determining the extent and nature of interspecific hybridization in the wild material and to offer as much in the way of explanation for the existing situations as has become evident. Much of the analytical work in connection with hybrid indices has been carried out using mature siliques. An important aspect of the silique has been shape. Therefore, it has been necessary to protect carefully all samples so that the natural shapes of the individual siliques were maintained. This was done by placing the samples loosely in envelopes with the infructescences project- ing above, where they would dry without receiving any kind of pressure. Once fully dry, the shape of the silique is permanent. In sampling field populations, a single infructescence only was taken from a given plant. Each infructescence provided ten to twenty siliques for analyses. The plants sampled were picked at random by arbitrarily taking from the plant nearest the toe of one foot every other step as one walked across the population. The largest populations were extensive enough to provide several hundred samples from one trip in a straight walk across them. ~ HYBRIDIZATION IN LESQUERELLA o Others required several trips taken at different angles to avoid sampling from the same place twice. Every wild population sampled consisted of thousands of plants. No attempt was made to estimate the total number of plants or to estimate the mini- mum number of infructescences required to give an adequate sample of a given population. However, as will be seen from the graphs, a relatively large number of infructescences were taken in each instance, probably several times the minimum require- ment in most cases. Seeds were collected from many of the wild populations for use in germination tests and to establish living collections for cytogenetic studies. In a number of instances, young buds were fixed for cytological studies. At the time of the initial flowering in the greenhouse of each of the species, L. densipila, L. Lescurii, and L. perforata, and of the hybrids, potted plants were isolated and self-pollinated, or bagged and self-pollinated. The flowers so treated failed to set seed, showing that a high degree of self-incompatibility was present. The results of these preliminary trials led to the use of the plants in various combinations of crosses without resorting to the emasculation of the individual flowers. The particular combinations of pollinations made are given in tables 2 and 4. The plants grew vigorously in the greenhouse, responding well to artificial conditions and not suffering noticeably from manipula- tive procedures. The plants began flowering ten to twelve weeks after planting and the pollinations were carried out over a period of two months in March and April, 1954. The progenies were grown during the early months of 1955. There were approxi- mately two thousand progeny plants, each grown individually in a separate three inch pot. These were grown to maturity and data were taken periodically during their period of growth. Mature infructescences were harvested from each plant for final detailed analyses. THE WILD SPECIES POPULATIONS The species and hybrids of Lesquerella under consideration are endemic to the Central Basin, as shown in Map 1. All four species shown are extremely abundant at given locations and a popula- tion may consist of hundreds of thousands of individuals. Les- querella densipila occupies cedar glade-like situations (Quarter- man, 1950) in southeastern Williamson County, in Maury County and southeast of Murfreesboro in Rutherford County, but it bas 6 REED C. ROLLINS . ‘ a nog et wae é x Bee : . @ ' 1 CHEATHAM COUNTY ' DAVIDSON COUNTY WILSON COUNTY (53128 c {85130 53129\_ ae bd ’ 53134 —y a z ‘ x2 WILLIAMSON COUNTY | a eG cg “ 7 : } moe Y j y A a : ' ij b ' ~ z { Maury COUNTY / ORT rape RUTHERFORD COUNTY / i rr / ‘ ‘ . : ' c- ane a) f ' ‘ ' ‘ ' H \ i ' \ ‘ ' ' i ° Rope 2 L LEWISBURG ; me t \ BEDFORD COUNTY : y ' e& L. perforota ‘- \ aoe ; MARSHALL COUNTY 4 “aut ' sie e , Ds sirens 3 ae eo) deistle 3 cai E cs ; ‘ pave’ we see ' ; : - ee © wie Xone re : ‘ i eae ” a =) ¢ ra Ra NS ene RENAL. Map |. The pidbing nade of the auriculate- — Lesquerellas and of their hybrids in pede Tenness each case, this is tal presently known distribution. The Central Basin covers Shek of the area shown been found most frequently near streams in level fields that are part of the flood plain of the stream. It is particularly abundant along the Duck River and its tributaries and along the upper Pp HYBRIDIZATION IN LESQUERELLA 7 of the Harpeth River. In the drainage area of the West Fork of Stones River, this species appears to be less abundant than in the other two river systems mentioned. Lesquerella Lescurii occurs in cedar glade-like situations at many stations, but also occupies many flood-plain sites on the Cumberland River, Arrington Creek in Williamson County, the Little Harpeth River in Davidson County and Stewarts Creek in Rutherford County. In general, the distribution of L. Lescurii is in the northern portion of the Central Basin and is distinct from that of L. densipila (cf. Map 1). Lesquerella stonensis is known at present only from the area of the East Fork of Stones River. Here it occurs on the flood plain except at one location on an upland limestone knoll less than a mile from the river. The presence of this recently dis- covered species on the upper Stones River was presumed on the basis of comparative data derived from the hybrids of it and L. densipila. L. stonensis was sought and found in 1955. L. per- forata is known only from three stations near Lebanon, where it is abundant. All three localities are flood plain or low level field-like areas. It is completely isolated from the other species. The detailed distribution of all four species has been documented with herbarium specimens and many of these have been cited in previous papers (Rollins, 1952, 1955). So far as I have been able to discover, these four species are the only ones of the auriculate-leaved group of Lesquerellas found in central Ten- nessee, A fifth, unrelated species, L. globosa (cf. Rollins, 1952) found along the Cumberland and in Maury County belongs to another section of the genus and need not receive attention in the present connection. THE WILD HYBRID POPULATIONS Three species in two combinations have been involved in producing wild hybrids, L. densipila x L. Lescurii and L. den- ipila x L. stonensis. The most extensive hybrid populations are those of L. densipila x L. Lescurii, found along the Harpeth River. These populations extend from just below the junction of Arrington Creek with the Harpeth River in Williamson County to about four miles above the point where the Harpeth enters the Cumberland River in Cheatham County ( Map 1). It is hard to ascertain the full extent of flood plain area occupied by Lesquerella hybrids, for there are stretches of the river that are difficult to get at. I have searched the most accessible parts 8 REED C. ROLLINS of the river flood plain from Arrington Creek to the Cumberland River, and from the hybrid population farthest down stream to the Cumberland, I have searched the area thoroughly. Apparent- ly the hybrids have not reached the Cumberland, for only pure L. Lescurii has been found there up to the present. There are long stretches of the river banks and flood plain where no hybrids are to be found. The largest populations appear to be on low lying land on the upper side of long river bends. This suggests that the seeds tend to settle-out in areas of quiet water at a time when the river is at flood stage. The river follows a tortuous course, providing many places which seem ideal for the develop- ment of hybrid populations, but all such places are not occupied. The largest hybrid population I have seen was sampled as collection 55115 and is found below the junction of Arrington Creek, along the Harpeth River (cf. Map 1). It was estimated that over 600 acres were well sprinkled with hybrid plants and many of these acres held hundreds of thousands of plants each. This hybrid population, barely down river from where L. den- sipila and L. Lescurii come together, is just below the area where these species become intermixed. A graph depicting the results of an analysis of this population is given on page 13. Most of the plants studied were found to be intermediate to some degree between the species and only a few had the combination of char- acters found in either L. densipila or L. Lescurii. The more or less bimodal nature of graph 55115, as compared to the graphs for the other hybrid populations, probably reflects the continuing entrance through migration of each of the species directly into this population. The hybrid plants are found almost exclusively on the flood plain but there are two known locations where they have begun to move off it. At one of these, the hybrids are found along the margin of the grade of U. S. Highway 70. Here, plants were seen on three different years, 1953, 1955, and 1956. The other location is at the edge of a field near U. S. Highway 70, approxi- mately one half mile from the Harpeth River toward White Bluff. In the latter instance, it is probable that seed was moved from the flood plain area to the higher ground by farming operations. LESQUERELLA DENSIPILA X L, LESCURII The Hybrid Index In developing a hybrid index, characters of the fruit were used. The assignment of values to these characters was arbitrary, HYBRIDIZATION IN LESQUERELLA 9 except that I was guided somewhat in my stress on characters of the fruit by the knowledge that these are the most important classificatory characters in the Cruciferae. The object was to set up a system of analysis that would reflect the pattern of changes actually seen in the hybrids, leaving aside the matter of an ideal model. For example, it was easily observed that the complete range of variation from globose siliques, characteristic of L. den- sipila, to the strongly flattened ones of L. Lescurii, was present among the hybrids. Various silique shapes were combined in what appeared to be a random way, with trichome characters and other silique features differing radically in the two species. e same index has been used throughout the study of L. den- sipila x L. Lescurii hybrids, the essential features of which are repeated from my previous publication (1954). L. Lescurn L. DENSIPILA A. Nature and Size of Trichomes on Exterior of Siliques. lsib-0 m si b- 1 ] si-2 msi -3 s si-4 misi -5 sbr -0 She, f+ 1. ete s s f, s si - : mi, mi si - 5 b=bulbous; br=branched; f=forked; 1=long; m=medium length; mi=minute; s=short; si=simple. B. Silique Shape. Strongly Flattened Slightly Subglobose Globose — Slightly depressed- flattened flattened globose 6 8 C. Trichomes on Style. None At base only Extending up style 0 bs 2 D. Trichomes on Replum Margin. None ew Numerous 0 1 E. Shape of Replum Apex. rel Definitely Rounded projecting projecting 0 1 2 F. Trichomes on Interior of Valves. Many to Few to None abundant scarce 0 2 As pointed out above, each of the values was arbitrarily as- signed to a model. The characteristics of L. Lescurii were NUMBERS OF PLANTS I i 16 Pié e« rs 53128 ty rome o123 45 6 7 8 9 10 It 12 13 14 18 16 17 18 19 20 21 22 23 24 25 26 27 28 ” p24 16a ris 8a a c 5 53133 m 2 eo sea | ° nv" 2 p2ae z “4 oa 166 rié §3135 84 rs SoS ert 24 L.LESCURII L. DENSIPILA $3131 53140 16 16 53129 O91 23 4 § 6 7 B 8 10 It 12 13 1415 1617 18 19 20 21 22 23 24 25 26 27 28 INDEX NUMBERS Figure | HYBRIDIZATION IN LESQUERELLA ll assigned zero throughout and those of L. densipila were assigned the highest number in each category A through F. Intermediates from one of the hybrid populations were selected to represent all of the different types that could be recognized. Each type was given a value and a model of it was selected for use as a standard. The samples from every hybrid population and from the hybrid cultures were actually compared with the same set of models to obtain the ultimate index number. The results of the analyses are presented in the graphs of fig. 1-3. This method of presenting the results is not a highly critical or an analytical one because a particular index number can be reached by a variety of combinations of the assigned values. The exact nature of the graph, particularly if it were converted into a curve, could be markedly affected by the arbitrary values assigned. However, the method adequately shows the intermediacy of the hybrid plants which combine as individuals the characters scored. This is the crux of the present study; not an examination of the way in which each trait is inherited. Wild Hybrids The overall picture, with considerable added data, is the same as that presented previously (1954). Fig. 1 is the same as that of the previous paper with the numbering of the graphs changed by placing the year date 53 before each of them. Other graph numbers beginning with 55 signify that the collections were made in 1955. Some populations were sampled in 1953 and again in 1955. Graphs for two such populations are given in fig. 2. Graphs 53128 and 55130 were made from the scores of samples taken in 1953 and 1955 respectively, from the same population. Graphs 53130 and 55131 similarly were made from samples of the two different years from the same population. Graphs 53128 and 55130 are quite comparable. Evidently the general character combinations remained roughly the same in the population, even though the second group of samples was of plants two generations away from the first. Graphs 53130 and 55131 are roughly comparable, although the latter tends slightly in the direction of L. Lescurii and, in this respect, is similar to Fic. 1. Graphs prepared from hybrid index numbers derived from scoring plants of four different hybrid populations and one population each of L. Lescuri and L. densipila. The number of plants scored to produce each graph is as follows: 53128—129 plants; 53129—95 plants; 53131—50_ plants: 93133--157 plants; 53135—257 plants; 53140—50 plants; Map | shows the location of each population. 12 REED C. ROLLINS graphs 53128 and 55130. This may indicate that plants of L. Les- curii, which are known to be abundant on the Little Harpeth 16 r 16 8 +8 | 55127 8 8 55131 | NUMBERS OF PLANTS @ uo ul uo ° os @ SINW1d 30 SHYSEBWNN @ a ; s a i) n @ r r hed o 16 ¢ 84 + 8 55130 o1 2 34 $ 67 8 9 10 tt 12 13 14 15 16 I7 18 19 20 21 22 23 24 25 26 27 28 INDEX NUMBERS . 2. Graphs prepared from gents index sri ge sie scoring plants - het > hybrid ge swore taken wo. differ years and third population ampled i ; ion. Graphs 53130 aa $5131 are from the same population. The number of plants scored to produce each graph is as follows: 53128—129 aga 53130—80 plants; 55127— 110 plants; 55130 165 plants; 55131—91 plants. Map 1 shows the location of each population, wi a?) 7) wi Ww _— nN me © fo] a5 VA ne Wig — we i © od o lanl 5 2 co aa o nr i: a) a. as) ee n> ow so) a HYBRIDIZATION IN LESQUERELLA 13 River (cf. population 53134, Map 1), only a few miles distant, are actually contributing secondarily to the genetical makeup of these populations. It will be noted that graphs 55129, 55127, 53133 and 53135, all showing an intermediacy between the 16 16 22 - 8 ~ 16 ~ 53138 53137 ° 8 53134 12) 20 21 22 23 24 25 26 27 28 20 21 22 23 24 25 26 27 28 hed z 2 53139 = = = a s 16 5 16 5 & a 3 ° nn ra we bar = > = z 2 4 * o 55129 O = 165 16 8 - 8 55115 ) Aaa | © 1 23 4 5 6 7 8 9 10 II 1213 1415 16 17 18 19 20 21 22 23 24 25 26 27 28 INDEX NUMBERS . Graphs prepared from hybrid cin numbers derived fer stig plants ficient two hybrid populations, one population of L. Lescurii, one lation of pure densipila, and two itp ns be L acai ‘pila showing idan of iutripeconon The number of plants oe oduce Kee graph is as follows: L. Lescuru, graph 53134—S50 plants; L. den ‘bile, pire 53139—25 plants; introgressed L. densipila, graph 53137—50 plants, sre 53138—50 plants; hybrid populations, graph 55115 3138 —185 plants, graph 55129—106 plants. Map | shows the location of each population. graphs of the species, are from populations above the entrance of the Little Harpeth River. On the other hand, graphs 53129 and 53130 are also Sincasateite and they are from populations below the mouth of the Little Harpeth. Graph 55115 is of special significance because it reflects the situation in the first hybrid population below pure L. densipila 14 REED C. ROLLINS on the Harpeth River and the junction of Arrington Creek where pure stands of L. Lescurii are found. Evidently, plants of both species come together at the junction of Arrington Creek with the Harpeth River, producing the hybrid population less than half-a-mile below that point from which the samples for graph 55115 were taken. This graph indicates that neither species had been fully assimilated into the hybrid complex as completely as apparently occurs farther downstream on the Harpeth. Experimental Hybrids The graphs of fig. 4 were produced from families of plants rown from seed produced by controlled pollinations. Graphs 54-5 and 54-12 were prepared from families of F, hybrids of the species cross L. densipila * L. Lescurii. In general, the F, plants were relatively uniform and somewhat intermediate between the species. Deviations from a uniform pattern were slight and scarcely noticeable in the families as a whole upon casual in- spection, but the study of individual characters did reveal the variations reflected in the graphs. Plants of Family 54-5 were usually golden yellow flowered, the siliques subglobose, the trichomes on the silique medium length and simple with an understory of short branched and short forked types; there were no trichomes on the style and few on the replum margin; the apex of the replum was barely projecting and the trichomes on the interior of the valves were few to scarce. Plants of family 54-12 differed rather consistently in having a few trichomes at the base of the style and in the absence of trichomes on the in- terior of the valves. Otherwise they were similar to the plants of family 54-5. The variation found in these interspecific F, families is appar- ently attributable to the heterozygosity present in the parent plants. Both L. densipila and L. Lescurii are self incompatible, which means that each plant is obligately outcrossed within its own natural population. Thus a fair amount of heterozygosity is certain to be present. It is interesting to note that the variation pattern can readily be pushed toward one species or the other by selecting the hybrids to be crossed. Graph 54-25 was prepared from the index numbers of a progeny produced by crossing an intermediate hybrid plant and one showing marked tendencies in the direction of L. Lescurii, both from the same hybrid population. The graph shows a definite trend toward L. Lescurii. On the other hand, HYBRIDIZATION IN LESQUERELLA 15 graph 54-26 shows an opposite trend toward L. densipila. This progeny was produced by crossing an intermediate hybrid plant with one showing marked tendencies in the direction of L. den- 8 - 54-5 n 12 13 14 15 16 17 18 19 20 z 4 c z z 2 8 8 © a m a wu oa vs 54-26 ef es | £4 : ° : 9 10 11 1213 14 15 16 17 18 19 2021 22 Ww v 5 > = > z od a 8 54-25 i ES ke 456 7 8 9 10 II 12 13 1415 16 54-12 16 17 18 19 ee Hh 12 13 14 15 a 8 8 54-10 54-11 9 '123 45 67 8 9 1011 12 13 1415 16 17 1819 2021 22 23 24 25 26 27 28 INDEX NUMBERS 4. Graphs Prepared from hybrid index numbers derived from scoring plants pisthast? by crossing as follows: L. poeeaey (53140) xX L. Lescuru (53134), graph (fe 5, plants scored 25; L. pillars eet i x L. Lescuri (53137), graph 54-12, plants scored 30; L. Lescurii (53134) intermediate hybrid (53129), graph at plants scored 74; = denip ($314 0) at intermediate hybrid (53129), graph 54-11, plants sco 6; intermediate hybrid (53125) > _ hybrid wing L. Lescurii tendencies (33135), sine 54-25, plants sedan 41; intermediate hybrid (53129) x hybrid showing L. densipila tendencies (53129), graph 54-26, Plants scored 30. sipila. Again, the plants selected were from the same hybrid population The reouh of backcrossing intermediate hybrids to each of 16 REED C. ROLLINS the species are shown in graphs 54-10 and 54-11. Of course, it is impossible to select an exactly intermediate hybrid from a group of highly variable individuals but an effort was made to select plants as nearly intermediate to the species as possible for backcrossing purposes. Index numbers of the backcross progeny to L. Lescurii are shown in graph 54-10. In graph 54-11 are shown scores from plants of the backcross to L. densipila. Seeds and Pollen The size of the hybrid population varies with the size of the available flood plain land. The largest population covered an TABLE 1. le quality in species oi ee hybrids of Lesquerella om wild sources in Tenn Name & Coll. No. No, seeds ae germinated % germination L. densipila 50 49 98.0 53138 50 37 74.0 53139 50 42 84.0 53140 46 92.0 Total 2 174 87.0 L. Lescurii 5312 20 7 35.0 53131 50 47 94.0 53134 50 38 76.0 53136 38 28 13.6 Total 158 120 75.94 L. densipila L. Lescurii* 53129-53 30 24 80.0 53130-79 29 25 86.2 - § ao 85.7 53130-23 23 14 60.8 53130-15 45 18 40.0 53130-73 3 10 30.3 53130- 9 28 8 8.5 53135-248 63 48 76.1 53135-204 29 11 37.9 Total 315 188 59.68 L. densipila < L. stonensis 53141 50 39 78.0 53142 50 42 84.0 Total TOO "BL 81.0 L. perforata 53145 50 43 86.0 *Each number under this combination represents an individual plant. HYBRIDIZATION IN LESQUERELLA 17 estimated 600 acres of land and the smallest was of several hun- dred individuals. However, most populations were large, ranging upward from several hundred thousand of plants. The aie success of the hybrids as colonizers is apparent from the vigor of the plants and the numbers produced. The capacity to reproduce is excellent. Table 1 gives the germinability of the seeds of wild hybrids of L. densipila x Lescurii and of wild kaa of L. densipila x stonensis, as compared with seeds of L. densipila, L. Lescurii and L. perforata. Although the per- centage of germinable seeds in one hybrid plant was 28.5, in another it was as high as 86.2. This is ‘much better than would ordinarily be expected in interspecific hybrids and certainly would easily permit the maintenance of the hybrid plants. In table 2 the germinability of seeds produced from various com- binations of crosses is shown. The controls were different, hand TABLE 2. cine CCE of seed produced by reciprocal pollinations be- twe lants of the same and different species and including various interspecific hybrid combinations in Lesquerella. Name & Coil. No.* No. seeds No. germinated % germination L. densipila * L. densipila 53140 X 53140 298 278 93.2 53140 X 53139 464 399 85.9 Total 762 677 88.84 : densipila * L. Lescurii 53140 X 53127 393 359 91.3 33140 53134 804 747 92.9 593138 X 53134 411 387 94.1 53137 X 53136 545 531 97.4 Total 2153 2024 94.00 I. densipila wild hybric 53140 X 53129-5-3 295 292 98.9 L. ioe “scurii wild hybrid soihe X 53129-5-3 355 350 98.5 Wild hybrid x wild hybrid 53135-248 X°53135-248 661 658 99.5 53129-53 xX 53129-53 224 208 92.8 Total 385 866 97.85 18 REED C. ROLLINS TaBLe 2. (continued) Name & Coll. No.* No. seeds No. germinated % germination L. densipila x L. perforata 53140 X 53145 402 394 98.0 L. densipila x L. X maxima 53140 X 53141 44] 385 87.3 L. Lescurii L. perforata 53131 X 53145 888 786 88.5 53134 xX 53145 370 349 94.3 Total 1258 1135 90.59 L. Lescurii L. X maxima 50142 -< 50134 657 635 96.6 L. perforata x L. perforata 53145 x 5ai45 495 469 94.7 L. perforata x L. X maxima 53145. x. b3142 668 636 95.2 L. * maxima pe iy maxima 53142 ® 53142 50 45 90.0 L. X maxima X wild hybrid 53142 % 53126-5-5 366 B57 92.0 *Reciprocal pollinations were made for all combinations. Since there were no instances where significant differences in the data from reciprocal combinations 0c- curred, these data were merged. pollinated plants of L. densipila. The high percentages of germinable seeds from all of the combinations shows a igh degree of compatibility between the species, between the hybrids and between the hybrids and species. A high degree of compatibility is further demonstrated in tables 3 and 4. In table 3, the pollen quality of wild plants 1s shown and in table 4, the pollen quality of plants produced by a variety of pollinations is given. Here, also, is overwhelming evidence in confirmation of the high compatibility present. Again, L. densipila was used as a control. HYBRIDIZATION IN LESQUERELLA 19 Cytological Observations * Seeds were gathered in the field from a number of individual hybrid plants which were open pollinated. Several families were grown from this seed source and meiosis was studied in the plants produced. The results of these studies are given in table 5 together with observations on the pollen of the same plants. To be scored as regular, no deviations from normal, how- TasLe 3. Pollen quality in plants of species and hybrids of Lesquerella S. grown from seed obtained from wild source Name & No. No. of plants No. filled grains No. unfilled grains % filled grains L. densipila DSLSL ES 1363 137 90.8 53138 1D 1335 165 89.0 53139 15 1403 97 93.5 53140 14 1340 60 95.7 Total 59 5441 459 92.22 L. Lescurii 53127 1 99 1 99.9 53131 5 494 6 98.8 53134 5 400 10 97.5 53136 5 303 17 94.6 Total 16 1296 34 97.44 L. densipila x Lescurii 531 3 18 Tiis 87 95.1 53130-8 17 1561 139 91.8 53130-9 4 360 40 99.0 53130-15 10 984 16 98.4 53130-23 9 847 53 94.1 53130-73 8 lt 27 96.6 53130-79 14 1166 234 83.2 53135-204 6 548 52 91.3 53135-248 24 2126 974 88.5 Total 110 10,078 922 91.61 L. densipila x L. stonensis 53141 15 1204 296 80.2 53142 15 1444 56 96.2 Total 30 2648 352 88.26 L. perforata ke: 53145 5 479 i 95.8 a am greatly indebted to and wish to thank Dr. L. O. Gaiser for producing the slides and making many of the observations recorded here. Many of the pollen counts given in tables 3-5 were made by Dr. John H. Beaman, to whom I wish to express my appreciation. 20 REED C. ROLLINS Taste 4. Pollen quality in plants grown from seed produced by reciprocal pollinations of different species and interspecific hybrid combina- tions in Lesquerella. Name & No. No. of plants No. filled grains No. unfilled grains 9% filled grains L. densi densipila sie x 53140 2 196 4 98.0 1 a Ie a X 53127 2 182 18 91.0 53140 X 53134 Z 196 4 98.0 50138 *. 53134 3 219 89 veal 53137 X 53136 2 149 5 T4h5 Total 9 746 162 82.15 L. densipila wild hybrid 53140 X 53129-5-3 2 196 4 98.0 L. Lescurii * wild hybrid 53134 X 53129-5-3 2 191 9 95.5 Wild hybrid * wild hybrid 53135-248 X 53135-248 2 198 24 99.0 53129-53 xX 53129-53 2 147 53 isn Total 4 345 55 86.25 L. densipila L. perforata 53140 X 53145 Dd 180 20 90.0 ie ar sue 738 maxima a x 53141 2 200 0 100.0 L. Lescurii x L. perforata S3I31L X 53145 3 218 82 72.6 53134 X 53145 ?- 197 3 98.5 Total 5 415 85 83.00 L. Lescurii et maxima 53134 X 53142 2 197 3 98.5 55; geen . maxima satus x 53142 2 198 2 99.0 L. X maxima * wild hybrid 53142 X 53129-5-3 2 187 13 93.5 *Reciprocal Depron were made and offspring grown from all of the com- binations indicated belo HYBRIDIZATION IN LESQUERELLA rd ever slight, were permitted. Thus, if a chromosome lagged ever so slightly, the figure was scored as slightly irregular and the plant was classed as having slightly irregular meiosis. In this, we were perhaps too cautious, for it is obvious from the pollen and on the evidence given above, that very little meiotic irregu- larity of sufficient gravity to produce sterility is present in the hybrids. Perhaps it would give a more accurate reading if the columns under regular and slightly irregular were put together under regular. As shown in table 5, in a progeny of 14 plants of accession Tasie 5. Observations on the Meiotic Chromosomes and Pollen of Fam- ilies Grown from Seed of Individual Wild Hybrid Plants. Population Family Number MEIOSIS % FILLED POLLEN & Acc. Number of Plants Slightly Number Regular IrregularIrregular low high all plants 53129 53 14 7 6 1 67 100 53130 8 11 2 8 1 57 100 89.1 53130 9 3 2 1 0 67 99 87.3 130 23 7 5 2 0 86 99 93.2 53130 73 5 1 2 v/ 97 100 98.8 5313 79 5 3 2 0 56 99 77.0 531 204 3 1 1 1 95 100 97.6 53135 248 20 6 11 3 59 100 87.0 Totals 68 27 33 8 TaBLe 6. Observations on the Meiotic Chromosomes of L. densipila and L. Lescurii. Les Accession Number MEIOSIS Spectes Number of Plants Regular Slightly Irregular Irregular L. densipila 53137 1 1 0 53138 0 1 0 53139 3 1 2 0 53140 1 1 0 7) L. Lescurii 53131 2 1 1 0 53136 2 2 0 0 53129, 6 showed some slight irregularities. The greatest irregu- larity was found in plant 6, classed as irregular, where, in the first division, chromosomes were in advance at metaphase, bridg- ing was seen at anaphase and stragglers were observed at telo- phase. Some plants, such as no. 4 and 12, showed only a slightly uneven line from a lateral view at first metaphase. Nevertheless oF REED C. ROLLINS these were classed as slightly irregular. Irregularities observed in accessions 53129, 53130 and 53135 included the following:— (1) at first metaphase, chromosomes off-plate from polar view; from later view, chromosomes uneven or some in advance; (2) at first anaphase, occasional bridges or pairs of chromosomes in contact; (3) at first telophase, stragglers present or an occa- sional bridge; (4) at second metaphase, uneven at plate or separation in advance; (5) at second anaphase, occasional chromosomes slow to divide. Second telophase was remarkably free of any irregularities even in those plants where bridges were fairly frequent at the earlier stages of meiosis. The relatively high quality of the pollen in the same plants suggests that many of the irregularities of meiosis observed were of a very minor nature and that the majority of dividing p.m.c.’s must complete their divisions without failure. As shown in table 6, a few plants of L. densipila and L. Lescurit were studied for comparisons with the hybrid progenies scored in table 5. The amount of meiotic irregularity among the hybrid plants is not greatly different from that observed in the species. In the few plants of the species examined, we did not see any highly irregular figures although those plants classed as slightly irregular were much the same as the hybrid plants similarly classified. Certainly, the hybrid plants do not show the amount or the extent of meiotic irregularity that is usually thought of as being characteristic of interspecific hybrids. Evidence of Introgression By and large, there is remarkabiy little evidence of intro- gression in the field involving L. densipila and L. Lescurit, especially in view of the ease with which these species cross and the very large populations of hybrids found. Population after population throughout the range of each species has been care- fully examined for evidence of introgression and in only one area has this been found. In an area in southeastern Williamson County, L. densipila on the south approaches L. Lescurii, which appears to be spreading from the north in a southerly direction. There is a heavy concentration of L. densipila southward and of L. Lescurii northward, as shown on Map 1. Graphs prepared from 50 samples each from populations 53137 and 53138 are given in fig. 3. Comparing these with graph 53139, which repre- sents a population of L. densipila to the south and remote enough from L. Lescurii to escape any chance of cross pollination, HYBRIDIZATION IN LESQUERELLA 23 shows that there has been some slight introgression from L. Les- curii. Indeed, the greater variability of populations in the area of 53137 and 53138 was recognized (cf. Rollins, 1952) before we were in possession of any real evidence that interspecific hybridization might be involved in Lesquerella. At that time, the relatively longer trichomes on the exterior of the siliques of these introgressed plants over those of pure L. densipila was noted. Although it is clear that introgression has occurred, it should not be assumed that these are hybrid populations in the sense of those so designated on Map 1. In fact, the genetic influence from L. Lescurii has been very slight, the shape of the siliques be- ing relatively unaffected. The difference between an average length for the trichomes of 81 microns for pure L. densipila and of 100 microns for population 53138 is not great but does show a slight influence from L. Lescurii, which has much longer tri- chomes. This point is in addition to those scored to produce the index figures of the graphs. LESQUERELLA DENSIPILA X L. STONENSIS [L. X MAXIMA] Wild populations of hybrids produced by the crossing of these two species were found before L. stonensis was discovered and before L. densipila was definitely known to be present in the drainage area of Stones River. These plants were originally described as L. densipila var. maxima in 1952 (1. c.) and now may be designated L. pappus scales with very short, triangular bases, little or not a all batsicnts abruptly pres he into long, orm acumina- tions: subanstate 240 hs i ak ek 1. Marshallia trinervia. C. Leaves thick and firm in texture, en in age, the middle and upper rapidly decreasing in : ower cauline (usually intact) being distinctly longer than those owt cera usually 4-12 i mber, successively reduced in len wards; plants with short caudices surrounded by fiber of old leaf bases; pappus scales with ovate bases, usually more or sie imbricate, gradually nar- rowed into acuminate apices, not at all aristate ................ D D. Stem simple, bearing only one head; icles te usually copiously pubescent near the summit; phyllaries usually with conspicuous white-hyaline autres at least below the middle, the adaxial surfaces smooth or at least not raised-veiny; corolla-tube regu- larly dilated sie into a distinct throat 2-3 mm. long; sur- face of achene between the ribs without resin atoms; pa scales regularly serrulate, the marge usually unbroken bi ieregdlar pointe) 5 i Marshallia grandiflora. D. Stem cymosely branched above the Bes bearing 2-10 heads; peduncle perf only sparingly pubescens near the summit; y ribs bearing minute resin atoms; pa pps irregularly serrulate, the margins broken by a few sharp points . .3. Mar shall: Mohrii. B. avai, and pales of the receptacle obtuse, be pales linear-s “naar clavate, usually conspicuously dilated at ag cir gi plants as e spring and early summer ...... . Marshallia o ta E. Plants caulescent, the 16 afy portion ears on less than % ac length of the peduncles, sometimes as long or longer; lateral nerves of the leaves usually prominent to the naked eye; upper leaf a (0.5-) 1-1.5 mm. long, crten regularly serrulate, sometimes somewhat fons nk eer WA i and Mountains fro ia to Alabama, extending into the eet Plain and ania Florida along the drainage area of the Chatt. ee ee rgia and Alabama . 4a. var obova E. beg strictly scapose, all the leaves basal, o r subscapose aves, but wi e | portion nelly less than % the Legh of the peduncle; lateral nerves of the leaves (eipectilly rom ace) usually obscure to the na eye; upper leaf surfaces, outer (abaxial) surfaces of the phyllaries and pales usually w cattered resin glands a set with minute resin droplets; achenes turbinate at turity, broadest at the truncate summit, the hairs of the ribs usually longer than the distance be- tween adjacent ribs; pappus scales (1-) 1.5-2.5 (-3) mm. long, 68 R. B.. GHANNELL sear irregularly aaa sometimes regularly serrulate; plants ically of sandy — the Coastal Plain from North Carolina Bidtida and“ Adabama ’ 200805005 sates ie var. scaposa A. Leaf blades, except the amet st, of a narrow, linear or linear-attenuate type, usualy 4-many times longer than broad y F. Phyllaries and pales of the receptacle merely acute, sometimes obtuse and perenne mucronate, but not subulate-tipped (or rarely 380 8 E 8 ~~ ga — pa) oR =| ° S R64 | = Be “< = fe) a i") wn 5 fo) o a _— 5 om io") - a jay =) So a a = = a fo] i=} -p ) 4 6 ° 5 orgia . Behe Sapa PSA ER TE sap mse eens .M ee ramosa. G. Rappus scales mainly over 2 mm. long, usually as long or longer ene; Tg, A 12 mm. ie ; inv volucres soa ) r. sig F, Phyllaries ae pales of the pe es gg sap oe senninate or 1 subulate- ed, pubescent with scattered monili S usually sis at least below the middle, oo say and of a rhomboidal form; plants flowering in the late summer and CONEY FOU oy ses a na oe ea Aa I. Hasek and lower cauline leaves with elongate, ascending, linear- lanceolate, prominently 3-nerved blades of relatively firm, thick te wi ; xture; vascular fiber d leaf bases we intact at the base oF the plant; plants of the igo Plain of N d oot GCaranne coo eS shallia cémnifolle I. Basal and lower cauline leaves sraisaate onion @ with spatulate, elliptic to elliptic-ovate, oblon: oblong-ovate, obtuse, 3-nerved blades of relatively thin-membranous texture, the lateral nerves not prominent; the base of the plant without the tin dae vascular fiber of old lea epg "alate of the aa is Plain from Geo Fe to Pilorida ana Terme 9 8. Marshallia " sonaslfolll 1. Marshallia trinervia (Walt.) Trel. ex Branner & Cov. Athanasia trinervia Walt. Fl. Carol. 201. 1788. (cHarL-Neotype: marked “Marshallia latifolia, St. John’s [Parish], Berkley, So. Calr]., June [18] °41,” Cranmore Wallace Herbarium. ) % Persoonia latifolia Michx. Fl. Bor.-Am. 2: 105, Er 43. 1803. “Hab. THE GENUS MARSHALLIA 69 in montosis Carolinae”; A. trinervia Walt. cited as a questionable syno- nym. (Lectotype-tab. 43, Michx. Fl. Bor.-Am. 2: 105. Trattenikia latifolia (Michx.) Pers. Syn. Pl. 2: 403. 1807. Marshallia latifolia (Michx.) Pursh, Fl. Amer. Sept. 2: 519, 520. 1814. “In the Carolina mountains . . . v. s. in Herb. Walter.”; A. tri- nervia Walt. cited as a synonym, Marshallia Schreberi J. F. Gmel. sensu Tratt. Arch. Gew. 1: 108. 1814 Phyteumopsis latifolia (Michx.) Poir. Encyc. Suppl. 4: 406. 1516. s. in Herb. Michx.”; A. trinervia Walt. cited as a synonym. Marshallia trinervia (Walt.) Trel. ex Branner & Cov. Rep. Geol. Surv. Ark. 4: 196. 1889. Basionym cited and M. latifolia |Michx.] Pursh as a synonym. Credit for the earliest publication of this combi- nation has been erroneously attributed to Porter, Mem. Torr. Bot. Club 5: 377. A, Perennial herb (3-)4-7(-8) dm. in height, stems usually arising singly from a thickish, sometimes slender, short-creeping, horizontal or ascending rhizome with coarse fibrous roots; the plants sometimes apparently solitary but usually occurring in small to large clump-like colonies with some of the stems sterile. Stem leafy, glabrous, striate, usually obtusely 4-5-angled and often purplish below, becoming dis- tinctly grooved and greenish near the peduncle. Peduncles (4-) 10- 20(-24) em. long, terminating the stem (and branches when present) , suleate, glabrous to glabrate below, becoming distinctly pubescent below the head. Leaves glabrous, entire, prominently 3-nerved, thin and membranous in texture, the lower narrowed into broad, petiole- “<< (4-)5-9(-10) cm. long, sessile or somewhat clasping, elliptic, oblong, ovate, ovate-lanceolate or lanceolate in outline, acute, usually wit strongly acuminate apices. Upper leaves sessile, acuminate, not greatly reduced in size. Involucres hemispherical to broadly campanulate, (10-)15-20(-25) mm. wide, (7-)8-10(-12) mm. high, rarely sub- tended by a reduced leaf-bract. Phyllaries glabrous, entire, little if at all imbricated at anthesis, subequal, marginless or with narrow white-hyaline margins below the middle, the outer usually (1-)1.5-3 mm. wide, (7-)9-11(-13) mm. long, linear, linear-lanceolate or lanceo- 70 R. B. CHANNELL Puare 3. Marshallia trinervia: Channell 2857, Tuscaloosa County, Alabama. THE GENUS MARSHALLIA 71 late, acuminate or attenuate, the midrib prominent, the veins obscure; abaxial (outer) surfaces pale green; the adaxial (inner) darker green, the whitish midrib prominent, the lateral veins obscure if visible at all. Pales of the receptacle narrowly linear, acute or acuminate-tipped, 8-9 mm. long, persisting beyond the shedding of the achenes, finally deciduous. Receptacle plane or slightly convex, conical in age. Flowers lilac purple. Dried corollas 10-15 (-17) mm. long, the tubes 6-10 mm. long, conspicuously expanded above into a well-defined throat about 2-3 mm. long. Anthers purple, (2.5-)3-3.1(-3.5) mm. long. Stigmas and sometimes the style exserted beyond the anther- column. Pappus scales shorter than the mature achenes, 2-3 mm. long, subaristate, with very short, triangular bases, not at all imbricate, abruptly contracted into long, filiform acuminations, regularly serru- late, minutely scabrous on the abaxial surfaces. Achenes 3-4.5 mm. ong at maturity, short-hairy on the ribs, the surfaces between the ribs with scattered resin atoms. Flowering from late May to the middle of July; fruiting in July and August. (Plate 3.) DISTRIBUTION. Occurring very locally along rocky stream banks and cliffs and in damp deciduous woods, often on calcareous clayey soils; Virginia (?), western North Carolina, central Tennessee, southward throughout Alabama and southeastern Mississippi into Louisiana (?). Map 1.) lanes EXAMINED. From cultivation: New York Botanical Gar- den, Moldenke 4940 (vps); Biltmore, North Carolina, Biltmore Her- barium (Collectors) 4215!. Without locality: Collector unknown, April 1840 (P); Steinhauer (pH). Virginia. Without definite locality: Stebbins (Ny). North Carolina. Without definite locality: Durand [?] (PH). Macon Co.—Highlands: Reinkes Place, Forest, 14 July 1938 (ncu); Harbison, 25 July 1904 (cx). South Carolina. Berkeley Co.— St. John’s, ex Cranmore Wallace Herbarium, June 1841 (cHARL- Neotype). Georgia. Without definite locality: Nuttall, 1815-30 (BM). Barton Co.—deciduous woods, sandy soil of natural levee, along road on e. side of Allatoona Creek, about 1 mi. from Etowah River, 5 June 1948, Duncan 8274 (axu, CU, FLAS, GA, NA, NCSC, SMU). Tennessee. Coffee Co.—Tullahoma, Biltmore Herbarium (Collectors) 4215° (us); oak barrens s. of Manchester, Svenson 10540 (BK, cu). Davidson Co.—Eggert, 13 July 1897 (mo). Lewis Co.—Little Swan Creek, near Meriwether Lewis National Monument, Natchez Trace Parkway: King 147 (vps), McDougall 1366, 1676 (us). Robertson Co.—Ridge Top, Eggert, 13 July 1897 (mo, ny). Alabama. County Uncertain: Sand Mountain, Biltmore Herbarium (collectors) 42154 (us), 4215° (Ncu, Us); mountains of Alabama, Ashe [?], May 1902 (Ncu). Without definite locality: Buckley, Sept. 1841 (mo), (BM, GH, ny). Bibb Co.— limestone cliffs, banks of Little Cahaba River, Pratts Ferry, Mohr, 3 June 1883 (F, us). Cullman Co.—Cullman, Mohr, 1 June 1882 (UARK). De Kalb Co.—De Soto Falls, e. of Valley Head, Wherry, 11 72 R. B. CHANNELL June 1933 (NA, PENN). Franklin Co.—Vicinity of Russellville, James 21 (mo). Lee Co.—Auburn: Earle & Baker, 29 June 1892 (ALU), 1347 [?], 29 June 1897 (mun, Mo, Ny), 28 May 1898 (¥F, GH, ILL, NY, poM, us). Mobile Co.—Dukes 88 (ny). Shelby Co.—Colena, Everts, 1879 (us); Shipman (F). Tuscaloosa Co.—Windham Springs: Chan- nell 2857 (to be distributed) ; Smith, 23 June 1875 (atu, us). Wilcox Co.—Buckley, June 1840 (au, cu, pwc). Mississippi. County un- known: south Mississippi, Hilgard, May 1859 (mo); north Missis- barium (iLL); Stewart (GH). Clarke Co.—Pachuta, Tracy 3299 (Ny). Forrest Co.—Hattiesburg, Channell 2610 (to be distributed). Greene Co.—1 mi. w. of Leaksville, Bailey, 19 July 1915 (massu). Perry Co.— Augusta, Tracy 4384 (ny). Scott Co.—Forest, Cook, 25 May 1925 (us); between Morton and Forest, Ripley & Barneby 11224 (ny); near Forest, Channell 2114, 2115 (to be distributed). Louisiana. Without definite localitv: Hartmann, 1877 [?] (Ft). The synonomy of this species has already been covered in con- siderable detail under the historical account of the genus. The correct combination, Marshallia trinervia, was first validly pub- lished in a checklist of the plants of Arkansas by Branner and Coville, who acknowledged in the introduction the assistance of Trelease in revising the nomenclature. Trelease, therefore, is cited as the author of the transfer. The type specimen of Athanasia trinervia Walt. is apparently nonexistent, for there is no specimen of the species in the Walter Herbarium now preserved at the British Museum. The last record of a Walter specimen of this species is in the reference given by Pursh in 1814 to the effect that he had seen the dried specimen in the Walter Herbarium (“v.s. in Herb. Walter.”). Consequently, there is no indication beyond the general infor- mation in the introduction of Flora Caroliniana as to the precise source of the Walter specimen. One would presume, in the ab- sence of information to the contrary, that it was collected in the vicinity of Walter’s home in South Carolina. Harper (Bull. Torr. Bot. Club, 38; 232. 1911), however, has pointed out: “But Walter's Flora Caroliniana contains the names of a considerable number of plants that probably do not grow within many miles of his home (which was near the center of the coastal plain of South Carolina), and a few that have not even been seen in South Carolina at all, in modern times at least; and it is reasonable to assume that he had some specimens from the two adjoining states in addition to those from his own.” The list of examples enumerated by Harper is headed by Mar- shallia trinervia. THE GENUS MARSHALLIA 73 The herbarium collections examined during the course of this revision have yielded, however, a single specimen of Marshallia trinervia from South Carolina. This specimen is located at the Charleston Museum and bears the data: “St. John’s [Parish], Berkley, So[uth]. Ca[rolina]., June [18] “41, Cranmore Wallace Herbarium.” These collection data have been carefully checked and verified through the assistance of Mr. Albert Swartz (at the time associated with the Charleston Museum), who has also kindly furnished information concerning the Cranmore Wallace Collection.'4 In view of the fact that the Wallace specimen is the only one of the species known to be in existence from South Carolina and since it was collected in Berkeley County, presumably near the type locality of Athanasia trinervia Walt., it has been designated as the neotype. The name Marshallia Schreberi |. F. Gmel. (Syst. Nat. 2: 1207. 1791) has been erroneously equated by various authors with Persoonia latifolia Michx. and thus with the earlier synonym Athanasia trinervia Walt. Although M. Schreberi was the first combination to be made under the genus and first appeared without a description, reference was clearly made at the time of publication to the generic description of Schreber. This name must consequently be equated rather with the name of the single plant upon which the generic description is known to be based, M. obovata (Walt.) Beadle & F. E. Boynt. [M. lanceolata (Michx.) Pursh]. Marshallia trinervia is the most distinctive species of the genus, being unique in having very thin-textured, ovate-lanceolate leaves, usually with strongly acuminate apices. The pappus scales (Plate 2, fig. 1) are also unique, being triangular at the very short bases, abruptly contracted into long, filiform acumina- tions and being somewhat subaristate in form. This species is apparently related to M. grandiflora which it sometimes ap- proaches in habit and with which it has in common the dilated corolla-throats. This species was reported from Arkansas by Lesquereux (Cat. Plants Ark.: Bot. & Paleont. Rep. Ark. Geol. Surv. p. 370) in 1860, but there is apparently no specimen to substantiate the record. On this basis, the species was also included by Branner and Coville (Rep. Ark. Geol. Surv. 1888. 4: 196) in their checklist 1 . . ad oii : , * Bulletin of the Charleston Museum, vol. 8, no. 5, May 1912, edited by Paul - Rea, 74 R. B. CHANNELL of the plants of Arkansas. There are, however, no modern col- lections of the species from Arkansas at all. Other than the fragmentary, although easily identifiable, speci- men in the herbarium of the New York Botanical Garden labeled “Va., Mr. Stebbins,” no specimen is known to support the Vir- ginia record for Marshallia trinervia. Although this specimen was apparently unknown to Fernald, the species was included in his enumeration of Virginia records (Rhodora 39: 473. 1937) on the basis of the erroneous interpretation of the Gronovian polynomial “Erigeron caule simplicissimo, saepius bifloro, folio caulino semi- amplexicaule .. .” (Fl. Virg. 122. 1762). According to Fernald, this entry, based by Gronovius on Clayton’s specimen number 375, was annotated by Asa Gray as Marshallia in the latter’s copy of Flora Virginica when he examined the Gronovian herba- rium in 1839. Photographs of the actual specimen kindly fur- nished by the British Museum show, however, in the hand of Asa Gray, the annotation “E[rigeron]. bellidifolius” only. The explanation for the discrepancy in the two annotations is not nown. In any case, the photographs clearly indicate that the specimen in question is not one of any species of Marshallia. The specimens of the species upon which the Louisiana re- ports are based are without precise locality data. These records, as well as those of South Carolina, Arkansas and Virginia, need verification by modern collections. Marshallia trinervia grows well and flowers in peaty soil under ordinary greenhouse conditions. Isolated plants, however, fail to set fertile achenes. 2. Marshallia grandiflora Beadle & F. E. Boynt. Marshallia grandiflora Beadle & F. E. Boynt. Bilt. Bot. Stud. 1: 7, pl. 1. 1901. “Dry soil near Saluda, Polk County, North Carolina”; Biltmore Herbarium (Collectors) 42154, 22 July 1898 (us-966506- Type, ex Biltmore Herbarium, presented by Mrs. George W. Vander- bilt, 1917, marked “Type Sheet” by C. D. Beadle and F. E. Boynton; Isotypes—GH, Ny, us-335500). Perennial unbranched herb bearing only one, long-pedunculate head, the stem usually (2-)3-5(-8.5) dm. in height, from a_ short caudex with coarse fibrous roots and, in age, with the tenacious fiber of old leaf bases, sometimes bearing leafy offsets; plants often somewhat clustered, the stems rarely with leafy proliferations from the lower leaf axils and rooting from the lower nodes. Stem glabrous, obtusely angled below, striate, becoming more distinctly grooved above, light green, buff or stramineous in drying, rarely if ever reddish, leafy to the peduncle. Peduncle sulcate, glabrous to glabrate below, becoming THE GENUS MARSHALLIA 15 or rounded apices, gradually narrowed below into broad, somewhat sheathing, petiole-like bases. Middle cauline leaves sessile or becomin glabrous, entire, with usually conspicuous white-hyaline (rarely purplish) borders to the middle and above, thin in texture at flowering time, becoming thick, firm and rigid in age, the outer usually 2-3 (-4.5) mm. wide and 7-12 mm. long, ovate, ovate-lanceolate, lanceolate or oblong in outline, usually acute, rarely obtusish; the abaxial (outer) surfaces green, becoming buff in age, at flowering time copiously minutely punctate with resin glands (visible under 10 x magnifica- tion), superficial resin atoms absent, the midrib prominent, obscurely reticulate-veiny, in age becoming thick and smooth with neither glands hor veins apparent; adaxial (inner) surfaces green, usually smooth, in rying sometimes rugose but not raised-veiny. Pales of the receptacle linear, acute, sometimes purple-tipped, 8-15 mm. long. Receptacle convex, conical in age. Flowers purple. Dried corollas 10-15 mm. long, the tubes about 10 mm. long, the upper third regularly and con- spicuously dilated into a distinct throat about 2-3 mm. long. Anthers purple, 3-3.5(-4) mm. long, the column exserted beyond the corolla throat. Stigmas and often the style also exserted beyond the anther- column. Pappus scales shorter than the achene, about 2 mm. long, acuminate from a broad base, regularly serrulate, minutely scabrous on the abaxial surfaces. Achenes 4-5 mm. ong at maturity, hairy on the ribs, the surface between without resin atoms. Flowering from mid June through July; fruiting in August and September. (Plate 4.) 76 R. B. CHANNELL PLate 4. Marshallia grandiflora: Bright 16904, Fayette County, Pennsylvania. THE GENUS MARSHALLIA ear g DISTRIBUTION. Occurring along rocky river and lake shores, creek banks, bluffs and flood plains in moist to wet sandy soil and in medi- acid soil of Coastal Plain-like “bogs,” uplands from southwestern Pennsylvania southward through West Virginia into southeastern Ken- tucky, east-central Tennessee and southwestern North Carolina. (Map 1.) SPECIMENS EXAMINED. From cultivation: New York Botanical Garden, Hartling, 24 June 1918 (m1). Pennsylvania. Allegheny Co.— near Pittsburgh, Leighton, Sept. 1915 (cu, PENN). Fayette Co.— Younghiogheny River, Ohiopyle: Bartholomew, 22 June 1940 (wva); Bright 283, 1884 (inp), 3062, 9417 (min), 6711, 13515 (PENN), 9416 (Na), 16900 (TEx), 16902 (LLC), 16904 (oxL), 21 June 1923 (cM, wva), 20 June 1936 (rex); Brown, Crawford & Van Pell 95 (pH); Clark, 30 June 1894 (cH); Fogg 18445 (PENN); Fretz, 3 July 1905 (pu); Henry, 24 June 1939 (cm); Jennings 9859 (pac), 9882 (cm), 9883 (PENN); Patterson, 3 July 1905 (cu); Ricker 1217 (us); Shafer 44 (¥, us), 444/11 (PENN), 1479 (cH, PH), 2195 (cm), 1 July 1900 (xy), 29 June 1902 (ny); Shafer & Medsgar 44 (cu, No, PENN), 44a (pu), 1 July 1900 (Ny, pH); Smith 6966 (PENN, pH, Us); Welton, July 1891 (pH). West Virginia. Without definite localitv: Millspaugh, 1892 (¥). Barbour Co.—Tygart Junction, Greenman 217 (¥F, GH); Tygart River, Moore 2556, 2568 (cH). Monongalia Co.—Quarry Run, Brooks 4547 (wva); Cheat River, Davis 5892 (wva). Preston Co.— Big Sandy Creek, Davis 5853 (wva); Cheat River, Albright, Smith & Carroll, 29 June 1947 (wva). Randolph Co.—Cheat River: Core, 20 July 1935 (wva); Hutton & Whitlach, 12 July 1941 (FLas, cH, LCU, NA, NY, OKL, PENN, TAES, WVA); Strausbough, 1 Aug. 1933 (Ny, OKLA). Stuart Camp, Orton, 10 July 1943 (wva); Shaver’s Fork, Cheat Mt., Sharp, 9 July 1941 (TENN); West Virginia Biological Ex- pedition (Collectors), 9 July 1941 (wva). Summers Co.—Greenbrier River below cliff at Bacon’s Falls, Fox 2479 (Ga, NA, NCSC, SMU, WVA). Upshur Co.—Sago: Grose & Grose, 25 July 1946 (wva); Polloch, 24 June 1895 (F, GH, ILL, MIN, MO, PH, POM, US, VHC), 4 July 1896 KL, CU, GH, LCU, MIN, MO, NCU, NY, US). Kentucky. McCreary Co.— south fork of Cumberland River, Braun, 18 June 1935 (GH). Tennes- see. Roane Co.—Emory River, Harrimon: Sharp 2007 (TENN); Under- wood ¢> Sharp 1144 ( BKL); Underwood 2767 (pH). South Carolina. Vithout definite locality; source questionable: Smith, 1881 (us). North Carolina. Without definite locality: western part of state, Hyams (xy). Henderson Co.—Ashe Herbarium, May 1902 (ncv); Biltmore Forest, Hendersonville, Clement, 12 Aug. 1936 (puKE); near Hendersonville, Biltmore Herbarium (Collectors) 4215” (us); Flat Rock, Crayton, Aug. 1919 (LAs, Ny); Muddy Creek, Smith, 19-26 Aug. 1881 (BKL, DUKE, F, GH, PHIL, us, YU); bog near Flat Rock Sta- tion, Wherry, 29 July 1933 (PENN). Polk Co.—Saluda, Biltmore Herbarium (Collectors) 4215 (Holotype—us; Isotvpes—cu, Ny, us). 78 R. B. CHANNELL Before, and even after, the recognition of Marshallia grandi- flora as a distinct species, specimens of this taxon were identified as M. trinervia and M. obovata var. platyphylla (var. obovata of this treatment) and for many years passed unnoticed under these names. Although Marshallia grandiflora was described in 1901, speci- mens clearly referable to the species were distributed by the Southern Appalachian Botanical Club in 1941, under the name M. obovata var. platyphylla. This illustrates the taxonomic con- fusion existing between these taxa. As pointed out elsewhere, M. grandiflora was considered to be only a robust form or state of M. obovata var. platyphylla in the seventh edition of Gray's Manual. The similarity of Marshallia grandiflora and M. trinervia has already been pointed out in the discussion of the latter species. In addition, both these species are usually simple-stemmed. Al- though M. grandiflora is rarely branched, when it is, the branches arise from the basal nodes. Marshallia trinervia, on the contrary, is branched from the upper leaf axils, if it is branched at all. Marshallia grandiflora may be distinguished from both M. trinervia and M. obovata var. obovata by the gradual reduction in the size of the leaves from the base of the plant upwards. The leaves of M. trinervia and M. obovata var. obovata extend up- ward without much reduction in size. The phyllaries of the latter are decidedly obtuse, unlike the acute or merely obtusish phyllaries of M. grandiflora. Moreover, the achene-surface of Marshallia grandiflora apparently lacks the superficial resin atoms characteristic of all other species of the genus. In western North Carolina, the only area in which the ranges of M. grandiflora and M. obovata var. obovata overlap (see Maps 1 & 2), the general flowering period of M. obovata var. obovata is usually from two weeks to a month or more in advance of that of M. grandiflora. In general, on an annual basis, a similar dif- ference in flowering period obtains throughout their ranges. Thus, these two taxa are effectively isolated both geographically and seasonally. Marshallia grandiflora specimens transplanted to the green- house from Henderson County, North Carolina and Randolph County, West Virginia, did not flower during the two-year period in which they were under observation. The plants survived but showed very poor growth. Specimens of M. trinervia an M. obovata var. obovata, however, grew very well in the greenhouse THE GENUS MARSHALLIA 79 under similar conditions and flowered two consecutive years. Having the highest altitudinal range and most northern distribu- tion of any other species of the genus, M. grandiflora may require a physiological cold treatment for successful growth and flower- ing in the greenhouse. 3. Marshallia Mohrii Beadle & F. E. Boynt. Marshallia mohri 1° Beadle & F. E. Boynt. Bilt. Bot. Stud. 1: 8, pl. 3. 1901. “. . . collected by Dr. Charles Mohr . . . at Cullman, Cullman County Alabama, June 24, 1893 . . .” (us-Type, ex Herb. Chapman of Biltmore Herbarium Collection, presented by Mrs. George soneeepilt 1917, marked “Type Sheet” by C. D. Beadle and F. E. oynton ** The original spelling with the single “i” is treated in accordance with Recom- mendation 73c of the International Code of Botanical Nomenclature, 1956. R. B. CHANNELL tito Sie a. Bie 897. Cullman County, Alabama. 21 June | Pirate 5. Marshallia Mohri: Eggert, THE GENUS MARSHALLIA 81 linear or falcate bract), usually (6-) 10-22(-25) mm. in width and (5-)7-10(-15) mm. in height including the pales. Phyllaries glabrous, without resin atoms, entire, with or without conspicuous hyaline bor- ders to the middle and aboye, thin to firm in texture at flowering time, thick and rigid, at least in age, usually (1-)2-3(-3.5) mm. wide, (5-)6-8(-13) mm. long, ovate-lanceolate, lanceolate, elliptic or elliptic- oblong in outline, usually acute, sometimes attenuate, rarely obtusish and minutely mucronate; the abaxial (outer) surfaces light brown, tan to pale greenish in color, usually smooth to smoothish under 10 x magnification, the midrib evident but not prominent, the lateral veins obscure within the tissue if evident at all; the adaxial (inner) surfaces usually greenish to dark green in color, distinctly raised-veiny, the mid- rib and vein-reticulum light green to whitish. Pales of the receptacle linear, acute. Receptacle convex, especially in age. Florets pale lilac- purple in color. Dried corollas (8-)10-15 mm. long, the tubes about 5 mm. long, not regularly dilated above into well-defined throats (if occasionally at all abruptly expanded, this portion only about 1 mm. in length or less). Anthers 3-3.5(-4) mm. long. Stigmas, and sometimes the style, exserted beyond the anther-column. Pappus scales 1-3 mm. long, shorter than the mature achene, acute to strongly acuminate, the margins irregularly broken by a few sharp points. Achenes to about 4 mm. in length at maturity, prominently hairy on the ribs, the inter- vening achene-surface beset with minute resin atoms, sometimes copiously so. Flowering from mid May through June; fruiting in July and August. (Plate 5.) DISTRIBUTION. Springy places and grassy glades (according to Mohr, Plant Life of Alabama), damp places in woodlands, grassy woodlands, low damp openings in flatwoods and in sandy soil of open meadows; known only from northern Alabama and northwestern Georgia (Lookout Mountain). (Map 1.) SPECIMENS EXAMINED. Without locality: Wray [?], ex Herb. Short broadly campanulate (rarely subtended on the peduncles by a reduced U 1891 (us); Eight Miles Creek, Eggert, 21 June 1897 (Mo); sandy loam, flat open woods near CCC Camp, Morgan, 17 June 1941 (xcv). Walker Co.—Without definite locality: pine woods, Earle, 20 June 1899 (F, ny). Georgia. County uncertain: Lookout Mountain: Ruth, July 1898 (m1, ny), 262 (vnc), 638 (Ny), Rock Creek, 662 (us), 692 (mo). The collection data (low pine barrens, Beaufort, Carteret 82 R. B. CHANNELL County, North Carolina, 10 August 1902) borne by an Albert Ruth specimen of this species (GH) are apparently erroneous. The specimen is mentioned here because it is thought to belong to Ruth’s collection of July 1898, from Lookout Mountain Geor- gia. The specimen resembles duplicates of the 1898 collection so closely that it seems safe to suppose that the specimen was originally a part of that series and that, inadvertently, it received the data intended for a different collection. Prior to the original description of Marshallia Mohrii in 1901, specimens of the species were labeled M. trinervia. Since that date specimens of the species have frequently been labeled M. obovata var. platyphylla (var. obovata of the present treatment). These incorrect determinations are suggestive of the general similarity of the three taxa. Marshallia Mohrii may be distin- guished from all other broad-leaved Marshallias, however, by the cymosely branched inflorescence of 2-10 heads, a feature unique in this species group. Unlike M. trinervia and M. grandiflora, the pappus scales of M. Mohrii have irregular, broken margins (Plate 2, fig. 3). Marshallia Mohrii (Plate 5) is considered to be very closely related to M. grandiflora (Plate 4). Specimens of these two spe- cies are quite similar in general appearance, as well as in finer details of structure. In fact, the morphological similarities and geographical distribution are such that M. Mohrii may be re- garded as a southern counterpart of M. grandiflora. There is also an apparent relationship between Marshallia Mohrii and M. trinervia. The phyllaries of M. Mohrii for example, are very similar in quality and form to the maturing phyllaries of M. trinervia. In addition, certain specimens representing ex- treme elements in the two species are very similar in general appearance. Marshallia Mohrii is apparently very rare and local in distri- bution. As far as is known it was last collected in 1941 by Morgan (apparently connected at the time with St. Bernard Abbey, St. Bernard, Alabama) in the vicinity of the type locality, Cullman, Alabama. Trips by the writer to the area in search of the species in the spring of two consecutive years (on one occasion assisted by Drs. Roland M. Harper and E. Gibbes Patton, University of Alabama), were not successful in terms of locating the plant. If located in the future it should be collected very sparingly an every effort should be made to protect it so as to avoid its extinction. THE GENUS MARSHALLIA 83 4a. Marshallia obovata (Walt.) Beadle & F. E. Boynt. var. obovata Athanasia obovata Walt. F]. Carol. 201. 1788. (BM—Type, on page 16 Herb. Walt., analvsis of type supplied by Mr. W. T. Stearn; GH— photograph examined.) South Carolina, Walter. Persoonia lanceolata Michx. Fl. Bor.-Am. 2: 105. 1803. “Hab. in montosis Carolinae”; A. obovata Walt. cited as a questionable syno- nym. (Pp—Lectotype, Herb. Richard, ex Herb. Drake. Trattenikia lanceolata (Michx.) Pers. Syn. Pl. 2: 403. 1807. Marshallia lanceolata (Michx.) Pursh, Fl. Amer. Sept. 2: 519. 1814. “In the Carolina mountains . . . v. s. i . Walt.” Marshallia lanceolata (Michx.) Tratt. Arch. Gew. 1: 109. 1814. Phyteumopsis lanceolata (Michx.) Poir. Encye. Suppl. 4: 405, 406. 1816. “vy. s. in Herb. Michx.”; A. obovata Walt. cited as a questionable synonym. Marshallia lanceolata (Michx.) Pursh var. platyphylla M. A. Curtis ex Chapm. Fl. Southern U. S., ed. 1, 241. 1860. ( cH—Lectotvpe, col- lected by M. A. Curtis at Hillsborough, North Carolina. ) Marshallia obovata (Walt.) Beadle & F. E. Boynt. var. platyphylla M. A. Curtis ex Chapm.) Beadle & F. E. Boynt. Bilt. Bot. Stud. 1: 6, pl. 6. 1901. striate, somewhat angled, becoming distinctly grooved above, green or greenish at flowering time, becoming stramineous in age, typically leafy to the middle or above, the leafy portion (5-)8-20(-30) cm. long, often as long or longer than the peduncle, occasional genoty es (ap- parently depauperate individuals) leafy below the middle only, inter- node length and leaf size not greatly reduced upwards, the leaves more or less abruptly discontinuous upwards (without the gradual ultimate reduction in size typical of M. Mohrii and M. grandiflora). Peduncles (8-) 15-35 (-45) cm. long, sulcate, glabrous or glabrate below, becom- ing copiously white-pubescent with ascending-incurving moniliform hairs, the pubescence usually abruptly discontinuous below the phyl- 84 R. B. CHANNELL laries. Leaves glabrous, rarely punctate with a few resin glands, entire, prominently 3-nerved, the radical and lower cauline leaves contracte below the middle into broad, somewhat sheathing, petiole-like bases, the upper becoming sessile with narrowed, clasping bases; radical leaves obovate, spatulate or oblong-spatulate, (2-)4-12(-15) mm. wide, (2-)3-6(-8) cm. long, with obtuse, sometimes emarginate blades; cauline leaves with oblanceolate, elliptic-lanceolate or elliptic, obtuse, rarely acutish blades, (3-)5-20(-30) mm. wide, (3-)5-12(-16) cm. long. Involucres hemispherical or broadly campanulate, 12- 25(-30) mm. wide, (5-)7-12(-15) mm. high including the pales. Phyllaries subequal in about 2 rows, the outer (5-)6-10(-13) mm. long, 1-2.5(-3.5) mm. wide, typically oblong in outline with obtuse and often minutely mucronate apices, sometimes lanceolate, elliptic or linear-elliptic but rarely with acutish apices, glabrous, usually without resin atoms, bordered to the middle and above by white-hyaline mar- gins, entire or occasionally remotely toothed, light green on the outer (adaxial) surfaces with the midrib thickened and somewhat raised above the adjacent tissue, the inner surfaces dark green except the white midrib, phyllaries becoming stramineous and firm in age. Pales of the receptacle glabrous, usually without resin atoms, 5-8 mm. long, linear-spatulate or spatulate-clavate, usually distinctly dilated upwards, with obtuse or minutely mucronate apices, gradually narrowed below. Receptacle convex, dome-shaped or conical in age, the pales persistent beyond the sheading of the achenes, finally deciduous. Flowers pale lavender to purplish, cream-colored or even white. Dried corollas (8-)10-12(-13) mm. long, the tube 5-7 mm. long, sometimes dilated colored or white, (2.5-)3-3.5 mm. long, the column exserted beyond the corolla-throat. Stigmas divergent and recurved in age, usually exserted beyond the anther-column. Pappus scales (0.5-)1-1.5 mm. ally lacerate, shorter than the achene. Achene at maturity broadest below the summit, (2.5-)3-4 mm. long, 10-ribbed, the ribs short- pubescent, the hairs of the achene shorter than the distance between adjacent ribs, the concave achene-surface between ribs covered with minute resin atoms, sometimes copiously so. Flowering from April through June; fruiting from May into July. (Plate 6.) DISTRIBUTION. Occurring on heavy clayey soils in meadows, woods, along rocky stream banks and road shoulders from Mecklenburg County, Virginia, southward through the Piedmont and upper districts of North and South Carolina, Georgia and Alabama, extending into western Florida along the drainage area of the Chattahoochee. (Map 2.) REPRESENTATIVE SPECIMENS. Without definite locality: Ameriq. Sept.: Baldwin (pH-258, pm); LeConte, Herb. Richard (Pp-ex Herb. Drake); Michaux [?] (v); Carolina, Fraser (pm—photograph). Mon- THE GENUS MARSHALLIA 85 Pare 6. Marshallia obovata var. obovata: Channell 2820, Lee County, North — Carolina 86 R. B. CHANNELL tosis Carolinae, Herb. Richard (p-ex Herb. Drake—Lectotype of Persoonia lanceolata). Elliott (cHarL, Mo-3636, mMo-Herb. Bern- hardi). Montibus Carolinae et Georgiae, Buckley (cu, Bm-photo- graph-Herb. Shuttleworth). Louisiana [?]. Drummond (x); Hale (pH). Virginia. Mecklenburg Co.—Near Soudan, Channell 2841 (to be distributed). Tennessee. Territorio Tennessee (probably western North Carolina), Herb. Richard (e—Herb. Drake). North Carolina. Without definite locality: Schweinitz (k-Herb. Bentham, ny); chiefly of the mountains and upper country, Vasey, 1878 (F); region of the Yadkin River, Herb. Britton 14341 (mo); Curtis (GH, MO, PH). Chatham Co.—Merry Oaks, Freeman, 22 May 1925 (Na); Sanford, Rad- ford & Radford 2004 (Ncsc, Ncu). Cleveland Co.—Kings Mt., Biltmore Herbarium (Collectors) 55219 (Ny, us). Davidson Co.—Yadkin River, Radford & Haesloop 7063 (Ncu). Durham Co.—Braggtown, near Eno River, Godfrey, Fox Anderson 40391 (GA, GH, MIN, NCSC, SMU); Channell 1936, 1938, 1939, 1946, 1957, 2817, 2819, 2832 (to be dis- tributed). Forsyth Co.—Salem, Schweinitz (GH); vicinity of Winston- Salem, Schallert, variously dated (Cu, DUKE, GH, LLC, MO , NY, TEX, UARK). Granville Co.—near Soudan, Virginia, Channell 2844 (to be distributed). Guilford Co.—Greensboro, Correll 10619 (DUKE, NA). Henderson Co.—swamps along Muddy Creek, Smith, 24 Aug. 1881 (mun). Iredell Co.—Gray, Sargent, Redfield & Canby, 24 Aug. 1881 (pH); vicinity of Statesville, Hyams, variously dated (NY, MIN, MO, pHiL-photograph—Herb. Martindale, pom). Lee Co.—near Sanford, Biltmore Herbarium (Collectors) 5521°, 55214 (us); Channell 2820 (to be distributed). Montgomery Co.—Troy, Fox & O'Connell 4706 (ncsc). Moore Co.—Hillsborough: Curtis, 1848 (GH-Lectotype of Marshallia lanceolata var. platyphylla) ; Palmer 42241 (GH, MO). Duke Forest, Rhododendron Bluff, Channell 728 (to be distributed). Person Co.—near Roxboro, Channell 1932, 2836 (to be distributed ). Polk Co. —Tryon: Townsend, 24 May 1897 (cu, us); Churchill, May 1899 (GH, MO, TENN). Randolph Co.—vicinity of Asheboro: Wiegand & Manning 3452 (cu, GH); Oosting 2456 (puKE, MIN). Rowan Co.—Gold Hill, Small & Heller 485 (¥, 1LL, MIN, MO, PH). Vance Co.—2 mi. S.€. of Fairport, Ahles 12697 (cu, Ncu, duplicates to be distributed). Wake Co.—between Norrisville and Cary, Channell 1941 (to be distributed). Warren Co.—Little Shocco Creek, Bell 2910, 2920 (cu, Ncu, duplicates to be distributed). South Carolina. Without definite locality: Buckley (cu); McCarthy, 1888 (us); Ravenel (r1); Herb. Walter (am—Type).- Abbeville Co.—Abbeville, Gibbes, 1855 (Ny). Anderson Co.—Blecklev Woods near Anderson, Davis, 24 May 1920 (cu, Lcu, MIN, MO TEX). Cherokee Co.—Broad Bank River near Gaffney, Wright & Munz 1314 (cu, Pom). Georgetown Co.—Georgetown (considered to be erroneous source of collection), Foster, 5 March 1859 (ny). Greenville Co.— Laurel Creek, Radford 91 (Ga). McCormick Co.—vicinity of Clark Hill Dam, Duncan 9442 (UALA, CU, FLAS, GA, NCSC, smu). Oconee Co.— THE GENUS MARSHALLIA 87 Seneca, McCarthy, June 1888 (us); Keowee, House 2198 (Ny, us); Old Stone Church, near Clemson, Morrison, 10 May 1933 (cLEMs). Pickens Co.—Anderson 1245 (min); dry woods near Clemson, House 3356, 3362 (ny). Spartanburg Co.—Spartanburg, Wheeler, 1891 (px). Georgia. Without definite locality: near Flint River, Herb. Muhlen- berg, 1812 (pu); Nuttall (pu); Elliott (xk); north Georgia, Vasey, 1878 (Ny); Chapman (¥, Ny, us); Darlington (¥1); Herb. Mann (cv). Baldwin Co.—Milledgeville, Harbison, 5 May 1914 (Ncu). Cherokee Co.—bottomland of small stream near Etowah River, Duncan 8315 (Ga). Clay Co.—near ravine, 1 mi. s. of Fort Gaines, Thorne ¢ Thorne 3790 (cu, GA, GH). Cobb Co.—Marietta, Allmendinger (Ny). Early Co.—near Hilton, Thorne 3616 (cu, Ga). Floyd Co.—Mount Berry, Jones 16 (NA). Fulton Co.—McPherson Park, Atlanta, Schneck, 20 May 1904 (iLL); Oakland City, Spratt, 28 May 1933 (ca); vicinity of Atlanta, Darby 60 (Na). Guinnett Co.—McGuire’s Mill, Biltmore Her- barium (Collectors) 1248” (BKL, F, ILL, MIN, NY, RM); Thompsons Mills and vicinity, Allard 262 (Mo, ny, us). Haralson Co.—Tallapoosa, Way 19 (us). Meriwether Co.—Pine Mt., Harper 1269 (GH, Mo, NY, us). Randolph Co.—Grier’s Cave, n. of Cuthbert, Thorne & Muenscher 7918 (cu). Richmond Co.—Augusta: Biltmore Herbarium (Collectors) 5521 (us); Dudgeon (ny); Cuthbert, variously dated (CONN, FLAS, Mohr, 20 May 1887 (Fr); Mohr, 16 May 1881 (vs). Etowah Co.— Mitchell, Harper 4 (GH, MO, Ny, PH, US). Florida. Without definite locality: Herb. Buckley (mo); Chapman (cu, Ny, us); Herb. Darling- ton (pwc) ; Jackson, ex Herb. Swarthmore College (pH). Jackson Co.— Marianna, Knight & Arnold, 5 April 1944 (FLAs). This taxon was first described by Walter in 1788 as Athanasia obovata. In 1803, Michaux unnecessarily renamed the species Persoonia lanceolata, the epithet of which was subsequently used to the exclusion of Walter’s obovata. The specific epithet lanceo- lata was preserved through a series of transfers which followed and was first combined with Marshallia by Pursh in 1814. Marshallia lanceolata var. platyphylla was proposed as new in Chapman’s Flora of the Southeastern States (page 241, 1860). The name was attributed by Chapman to M. A. Curtis who, judging from the labels on herbarium specimens examined during 88 R. B. CHANNELL the course of this revision, is clearly responsible for the taxon. The variety was said to differ from typical M. lanceolata in hav- ing stems leafy to the middle. One would infer, therefore, that M. lanceolata at this time was interpreted as having mostly basal or radical leaves. In the revision of the species of Marshallia by Beadle and Boynton (Bilt. Bot. Stud. J: 3-10. 1991) var. platyphylla was up- held and applied to the leafy-stemmed plant. The authors stated that, while the original description of Athanasia obovata was very brief, they had no hesitancy in applying the Walter epithet to the species, i.e. the scapose plant. On this basis the following transfers and combinations were made: M. obovata ( Walt.) Beadle & F. E. Boynt. and M. obovata (Walt.) Beadle & F. E. Boynt. var. platyphylla (M. A. Curtis ex Chapm.) Beadle & F. E. Boynt. By so defining these taxa Beadle and Boynton are largely responsible for the concepts that have subsequently come into general usage, i.e. interpreting M. obovata as comprising the scapose and subscapose plants and var. platyphylla as including the caulescent plants with larger leaves. Investigations conducted during the present revision, however, have indicated that the type of Walter’s Athanasia obovata (now preserved at the British Museum) consists of two specimens both of which are themselves leafy-stemmed. This fact was first sus- pected when the Gray Herbarium photograph of the type was examined. The brief original description was then re-examined in the light of this information. The entire passage referring to foliage is: “foliis radicalibus obovato-oblongis subnervosis . . On the basis of this statement it cannot be concluded whether the plants are scapose or caulescent, for only the radical leaves are described. Additional information concerning the type of Athanasia obovata, however, has been kindly supplied by Mr. W. T. Stearn of the British Museum. Mr. Stearn has furnished diagrams of the relative position of the individual leaves of the type specimens. On the basis of these diagrams there is no doubt that the type- material belongs to the leafy-stemmed taxon. Mr. Stearn arrived at a similar conclusion after comparing the type-material with representatives of both the leafy-stemmed and scapose variants. Mr. Stearn’s analyses indicate that, on the basis of leaf-characters, the Walter type-material belongs to the leafy-stemmed taxon with conspicuously 3-nerved leaves, as figured by Beadle and Boynton (1. c., pl. 6) for Marshallia obovata var. platyphylla. THE GENUS MARSHALLIA 89 The type-material, according to Mr. Stearn, essentially matches several specimens in the British Museum under the name Mar- shallia lanceolata. 1 have examined photographs or duplicate specimens of these collections and concur in this opinion. Mr. Stearn was also kind enough to compare the Walter type- material with a specimen in the British Museum collected by R. K. Godfrey (no. 49184, Brunswick County, North Carolina ) and labeled Marshallia obovata. This is a collection of strictly scapose plants of which several duplicates have been examined during the course of the present revision. Mr. Stearn concurs in the belief that this collection differs from the Walter type and that it corresponds to Marshallia obovata as figured by Beadle and Boynton (1. c., pl. 5) with apparently 1-nerved leaves. On the basis of these data it has been concluded that Walter’s Athanasia obovata has been incorrectly interpreted in the past. It is the same taxon as that described as var. platyphylla. On taxonomic grounds, however, two varieties are here recognized as comprising the species. For reasons of priority, the leafy- stemmed variety must be called Marshallia obovata ( Walt.) Beadle & F. C. Boynt. var. obovata and the scapose one must be named. Despite the fact that the new combination Marshallia obovata was misapplied to the scapose variety, that name must be retained for the leafy-stemmed taxon upon which the epithet was originally based. Both varieties of Marshallia obovata flower in the late spring, in general earlier than the three preceding broad-leaved species. Their phyllaries (Plate 1, figs. 4 & 5) are elliptic, linear-elliptic or oblong in outline, usually with obtuse, sometimes minutely mucronate apices. Occasionally the phyllaries may be abruptly pointed but they always have a marked tendency toward the obtuse condition. The pales are unique in being linear-spatulate- clavate and are also often minutely mucronate at the apices. In general, the obtuse phyllaries and the clavate-spatulate pales serve to distinguish M. obovata from all other species of the genus. Occasionally the phyllaries of M. grandiflora are abruptly pointed or even obtusish, but the acute apex is nevertheless the general rule in this species. The occasional striking similarity of the phyllaries of M. obovata var. obovata and M. grandiflora, the simple habit and similar leaf-form are suggestive of relation- ship. Indeed, the two taxa have often been confused taxonom- ically as already pointed out. The relationships of the two varieties of Marshallia obovata to 90 - BB. CHANNELL one another and further discussion of their relationships to M. grandiflora are presented under var. scaposa. Marshallia obovata var. obovata occurs almost exclusively on the geologically older Piedmont and upper districts on very clayey, highly plastic soils (Map 2). The southern extension of the range of this variety into western Florida is along the drain- age area of the Chattahoochee where the sandy soil has appar- ently been eroded away, exposing clay, thus providing suitable edaphic accommodations for the spread of the plants into the Coastal Plain. The geographical range of Marshallia obovata var. obovata (as var. platyphylla), according to the eighth edition of Gray's Manual, extends from Florida to southern Missouri and North Carolina. The Missouri record, credited to the authority of Palmer and Steyermark, is apparently based on Palmer's collec- tion number 34811 from Ozark County. This collection, of which three duplicates have been examined, proved to represent un- commonly broad-leaved specimens of M. caespitosa var. signata. The inclusion of M. obovata var. obovata in the Gray’s Manual range on this basis is thus incorrect. According to the present study, however, the natural range of this variety extends from North Carolina into Mecklenburg County, southern Virginia. This is the first authentic report of the variety from the territory covered by Gray’s Manual. Marshallia obovata var. obovata grows well and flowers in the greenhouse in ordinary garden-type soil. Under these conditions the leafy-stemmed nature of this variety is emphasized. 4b. Marshallia obovata (Walt.) Beadle & F. E. Boynt. var. scaposa 16 Var. nov. Marshallia obovata (Walt.) Beadle & F. E. Boynt. sensu Beadle & F. E. Boynt. Bilt. Bot. Stud. 1: 5, pl. 5. 1901. Not Athanasia obovata Walt. Fl. Carol. 201. ; Perennial scapose or subscapose herb (0.5-)1.5-3.5(-5.0) dm. tall, the stem from a short caudex wtih coarse fibrous roots, the vascular Planta acaulis arte scaposa vel subscaposa cum caule breve (minusquam “4 longitudo pedunculi), foliis caulinis paucis; superficies foliorum cum nervis laterali- bus obscuris sine amplificationem; foliis, squamulis involucri ac_paleis receptaculi plerumque supra glanduliferis, et sparse glandulari-atomiferis; acheniis ad maturl- tatem turbinatis, ad apicem late truncatis, 10-costatis, villosis, trichomis longioribus spatium inter ¢ tes is pappi (1-) 1.5-2.5 (-3) mm. longs, 5. ype, coarse sand, plowed fire lane, 3 miles west of Leland along U. S. Rt. 76, Brunswick County; North Carolina, R. K. Godfrey 49184, 21 May 1949; Isotypes-BM, DUKE, G, GA, NA NCSC; NCU). c THE GENUS MARSHALLIA 91 fiber of old leaf bases and several to many radical leaves; plants usually simple with a single scape but occasionally with 1-5 intact leafy offsets, each bearing a peduncle. Inflorescence a single head, typically borne on a simple (true) scape but sometimes apparently long-pedunculate from a short stem with a few leaves, the leafy portion % the length o the peduncle or less. Peduncles (5-)15-30(-40) cm. long, naked or sometimes with a remote linear-oblong, obtuse bract, sulcate, glabrous or glabrate below, becoming copiously white-pubescent beneath the head with ascending-incurving, moniliform hairs, the pubescence abruptly discontinuous below the phyllaries. Leaves radical or nearly so, glabrous, 3-nerved or apparently 1-nerved with the laterals of the upper (adaxial) surface obscure to the naked eye; lowermost (radical! ) leaves (1-)2-5(-10) cm. long, 4-10(-15) mm. wide, with obovate, spatulate or oblong-spatulate, obtuse, sometimes emarginate blades, narrowed below into broad petioles; subradical (the lower cauline or upper rosette) leaves (3-)5-10(-12) cm. long, (3-)5-10(-15) mm. wide, with lanceolate, oblanceolate or linear-oblong, obtuse or acutish blades, narrowed below into sheathing petioles usually shorter than the blades. Heads hemispherical or broadly campanulate, (10-) 15-20 (-25) mm. wide, (5-)7-12(-15) mm. high including the pales. Phyllaries usually oblong with obtuse and often minutely mucronate apices, some- times lanceolate, elliptic or linear-elliptic with obtuse or abruptly pointed apices, (5-)6-8(-10) mm. long, (1.5-)2-2.5(-3) mm. wide, glabrous, often with scattered resin atoms, usually green (rarely red- dish) and thin in texture at anthesis, buff to stramineous and firm in age, bordered to the middle and above by narrow to quite conspicuous white-hyaline margins, usually entire, sometimes remotely dentate or erose, the midribs slightly thickened. Pales of the receptacle glabrous, usually with scattered resin atoms, 5-8(-10) mm. long, spatulate- clavate or linear-spatulate, gradually narrowed below, the summit usually distinctly dilated, the apices obtuse and minutely mucronate or abruptly pointed. Receptacle convex, dome-shaped or conical in age, the pales usually persistent beyond the spreading of the achenes. Flowers white, whitish or pale lavender in color, externally pubescent. Dried corollas 10-12 mm. long, the tube 5-6 mm. long, expanded above into a shallow throat, the limbs spirally twisted. Anthers cream-colored, white or bluish, 2.5-3.5 mm. long, the column exserted beyond the corolla throat. Stigmas exserted beyond the anther-column, divergent and reflexed in age. Pappus scales white or castaneous in color, usu- ally imbricate at the base, ovate, strongly acuminate, usually with at least the upper half somewhat irregularly lacerate, sometimes regularly serrulate, (1-)1.5-2.5(-3) mm. long, as long or shorter than the achene. Achenes obconic, 10-ribbed, about 3 mm. long, the ribs copi- ously pubescent, the hairs as long or longer than the distance between adjacent ribs, the concave achene-surface between ribs usually covered with minute resin atoms. Flowering from April through the first week of June; fruiting from May to mid July. (Plate 7.) 92 R. B. CHANNELL Warshellia obovate (Malt. Beadle & F. E. ys. eeapome ver. nor. in ed. Bxamincd in revisional sandy of tlorshelfia AREER. saa ano & 4 ae e & & PES SLC He eee BEER Oe LGM OF MORMETH 1 AROLINA Comnty ak Marchallis obovate (Walt) 8, & 8, Coarse ennd, ploved fire lane, 3 miles went of Leland along 1. 3. Rt, 76. ' mn vy * an, 4 ® eh . Cadllected by BK. Godfrey awe Ne. ys May 21, TE 7. Marshallia obovata var. scaposa: Fig. 1, Godfrey 49184-Type, agree’ ees North Carolina (to scale); Fig. 2, specimen from Harnett Count Carolina grown in the gna a and 1 (approx. 4 scale). Ys showing the conspicuous rosette ne THE GENUS MARSHALLIA 93 DISTRIBUTION. Occurring on light sandy soils in dry pinelands, edges of sandhills, dry savannas and oak woods in the Coastal Plain from North Carolina southward into Florida and Alabama. (Map 2.) REPRESENTATIVE SPECIMENS. Without locality: Herb. Wallace 55 (cHARL); Herb. Rugel (rpum—Herb. Martindale). Am. Sept.: Le Conte, Herb. Richard (p—Herb. Drake); Fraser (r1-Herb. Webb). North Carolina. Bladen Co.—Biltmore Herbarium (Collectors) 1258 (conn, CU, RM, Us); vicinity of White Lake: Blomquist 5280 (puke); Channell 2816 (to be distributed) ; Oosting 34109 (PENN, pH). Brunswick Co.— 3 mi. w. of Leland, Godfrey 49184 (GH-Tvpe; DUKE, G, GA, NA, NCSC, ncu—Isotypes). Columbus Co.—Bolton, Schallert 11112 (11, Min, TAES). Cumberland Co.—Vicinity of Fayetteville: Biltmore Herbarium (Collectors) 12584 (us), 1258¢ (ncsc, ny); Channell 1906, 1910 (to be distributed) ; Godfrey & Fox 49450 (Ncsc). Durham Co.—Radford & Radford 3036 (ncu). Hartnett Co.—Channell 2821, 2822, 2823, 2830, 2831 (to be distributed). Johnson Co.—Princeton, Mitchell, 19 May 1936 (puKE). Montgomery Co.—near Eldorado, Correll 810 (DUKE). Moore Co.—near Aberdeen, Biltmore Herbarium (Collectors) 1258/ (ILL, MIN, Ny, us); near Pinebluff, Palmer 42370 (cH, Mo, NY). New Hanover Co.—Wilmington, Bartram & Long 1054 (rH). Rich- mond Co.—Hamlet, Williamson (pH). Scotland Co.—Laurinburg, Rad- ford & Stewart 441 (Ncsc, Ncu). Wake Co.—near Raleigh, Godfrey 77 (GH, Ncsc); between Cary and Morrisville, Channell 1940 (to be dis- tributed). South Carolina. Without definite locality; McCarthy, 1888 NY, P (CHARL, DWC, F, us), 3 May 1867 (GH); Smith (PENN). Barnwell Co.— (Collectors) 1258° (us); Bartram 2283 (rH). Horry Co.—2.8 mi. s.e. of county line, S. C. Rt. 9, Radford & Stewart 987 (Ncu). Lexington Co.—Swansea, Weatherby 6126 (GH, NY, Us). Richland Co.—vicinity of Columbia: Bartram, 16 May 1912 (pH); Blackman (Fr); Canby with Sargent 57 (cH); Philson, 12 May 1936 (puKE, OKLA); Taylor, May 1890 (BKL, F). Georgia. Bibb Co.—Macon: Greenman (pH); Shaw, May 1888 (yu). Burke Co.— Alexander, Ellis (pom). Calhoun Co.— Arlington, Thorne & Muencher 2879 (ca, us), 8033 (cu, Ga, GH); Leary, Thorne 3331 (cu). Lee Co.—3 mi. e. of Smithville, Duncan 2314 (Ga). Macon Co.—Marshallville, Earle 3140 (ny). Peach Co.— near Byron, J. C. D. 28, 23 April 1927 (uark). Florida. Without defi- nite locality: Buckley (mo); Chapman (ny); Elliott (x); Rugel 74 (F, GH, Mo, Ny, us). Alabama. Barbour Co.—15 mi s.w. of Eufaula, 94 R. B. CHANNELL Wherry, 12 April 1897 (Na). Lee Co.—Auburn, Earle & Baker (Fr, ILL, MIN, MO, NY, RM, US). Pike Co.—Troy, Leland, 17 April 1891 (GH). As indicated in the discussion of the preceding variety (obovata), the taxon treated here as var. scaposa has never before had a legitimate name, although it has previously been recognized taxonomically by implication. This situation devel- oped as a result of the description of var. platyphylla as a leaty- stemmed variant of M. obovata and with the subsequent erroneous restriction of the application of the epithet obovata to its scapose counterpart. Thus the taxon treated by Beadle and Boynton as M. obovata represents a new variety, different from Athanasia obovata Walt. Marshallia obovata var. scaposa occurs almost exclusively on the geologically more recent, light sandy soils of the Coastal Plain, along or outside the Fall Line. Although the two varieties of M. obovata flower over the same general period and although their ranges abut, they are nevertheless effectively isolated ecologically. Variety scaposa occupies more specialized, xeric habitats on the edges of sand hills beneath pines or amongst scrub oaks, ete., while var. obovata is more widely distributed over the Piedmont in more mesic habitats and even extends into the mountains. While the Fall Line between the Piedmont and Coastal Plain forms the more or less sharp line of demarcation between the ranges of the two varieties (Map 2), the apparent overlapping of ranges at certain points is due to the uneven nature of the Fall Line in certain areas where there are inter- fingerings of geologically older clayey soils with younger sandy soils. At certain points along the Fall Line where there are abrupt changes in the physical nature and geologic history of the soils, the two varieties may occur in close proximity to one another. Yet morphological distinction is maintained along the line of contact with very little if any intergradation. This suggests that the varieties may in fact be isolated genetically. Experi- mental attempts to hybridize the two varieties in the greenhouse were unsuccessful. There is no question but that the two taxa are very closely related morphologically. Genetically the evidence is inconclusive for additional experiments are required to determine whether or not they will actually hybridize. From the standpoint of geo- graphical distribution the two taxa are strongly allopatric. His- torically the two elements have been treated as geographical THE GENUS MARSHALLIA 95 varieties. Despite the fact that there are indications that the two taxa may probably be completely isolated genetically, and have certain qualities of distinct species, it seems best to retain the varietal status until sufficient experimental information is accumu- lated to justify fully their elevation to specific rank. In considering the probable direction of evolution within Mar- shallia obovata there are two indications which suggest that var. scaposa is probably the younger, derived member. The first in- dication is the fact that the plants are scapose or subscapose in habit, a growth form which morphologically may be considered to have been derived from the caulescent through a telescopic type of reduction of the stem. The second indication concerns the distribution of the varieties, respectively, over areas with en- tirely different geological histories. It is significant that the dis- tribution of the scapose variety, presumed on morphological grounds to be the more recently evolved, occupies the area of geologically more recent soils and which has only in geologically recent times become available for plant occupancy. It is perhaps significant further that the scapose variety has not been collected in any of the disjunct Coastal Plain-like habitats isolated in the Piedmont and mountains which are floristically notorious for their richness in Coastal Plain species. This may be due to the relatively recent origin of var. scaposa somewhere along, at or near the Fall Line and its subsequent spread into the adjacent coastal plain only. If the actual course of evolution in Marshallia obovata has been in the direction supposed here, it may be concluded that var. obovata is the older representative and that it apparently has Appalachian affinities. Turning, then, to those species with an Appalachian distribution, it can be readily seen from Map | that M. grandiflora is conspicuous in this respect. Moreover, this is the one species which on a morphological basis is more closely related to M. obovata var. obovata, as already pointed out. Marshallia obovata var. scaposa is apparently adapted geneti- cally to the more rigorous xeric habitat of the Coastal Plain. Representatives of the two varieties maintained their respective differences in growth habit under similar greenhouse conditions when grown in a garden-soil mixture. In fact, the scapose nature of var. scaposa was actually emphasized as shown in Plate 7, fig. 2. y the-production of more numerous radical leaves and a more striking rosette habit of growth. Variety obovata responded in a similar manner as under natural field conditions. 96 R. B. CHANNELL 5. Marshallia ramosa Beadle & F. E. Boynt. Marshallia ramosa Beadle & F. E. Boynt. Bilt. Bot. Stud. 1: 8, 9, pm Soul, ows in moist, sandy pine-lands at Eastman, Georgia (type locality), where it was collected in full flower June 5, 1900, by Mr. C. L. Boynton, of the Biltmore Herbarium.” (us—Type, ex Biltmore Herbarium, presented by Mrs. George W. Vanderbilt, 1917, marked “Type Sheet” by C. D. Beadle and F. E. Boynton.) Perennial linear-leaved herb (1-)2.5-5(-6) dm. tall, the stem from a thickish crown (caudex) with coarse fibrous roots, usually bearing wiry fiber of old leaf bases, freely cymosely branched about the middle and above, the alternate branches ascending and producing a more or less flat-topped, compound inflorescence of (2-)4-12(-20 or even more) small, long-pedunculate heads, the peduncles sometimes approximate, giving an umbellate appearance; stems sometimes solitary but usually clustered with several arising from the same caudex, and often with sterile, leafy offsets at the base. Stems annual, glabrous below, becom- ing very sparingly pubescent on the branches, striate or angled below (sometimes squarish) , becoming more distinctly sulcate above, usually green or greenish in drying, stramineous in age and often purple at the base, leafy to the inflorescence branches and above. Peduncles slender, 0.5-1 mm. in diameter, sulcate, glabrate below, becoming sparsely pubescent near the summit, naked or leafy-bracted below the middle by the reduced, linear, obtuse leaves of the upper stem. Leaves gla- brous, entire, firm in texture, the lower prominently 3-nerved, gradually diminishing in size upwards and usually extending well upon the branches to the bases of the peduncles as reduced linear bracts; the lower leaves contracted into winged, petiole-like bases as long or longer than the blades; the middle and upper cauline leaves becoming sessile and only 1-nerved. Radical leaves rarely intact or well developed, when present 5-10(-15) mm. wide, (2.5-)4-8(-10) cm. long including the petioles, with oblong, oblong-spatulate or obovate, obtuse blades about as long as the petiolar portion; lower cauline leaves (3-)5-8(-10) mm. wide, (5-)8-20(-25 or more) cm. long including the prolonged petioles, linear, linear-attenuate or linear-lanceolate in outline, wi obtuse apices; middle cauline leaves gradually reduced in size up- wards, usually 3-8(-10) mm. wide, 3-10 cm. long; the upper leaves greatly reduced (1-)2-3(-4) mm. wide, (1-)2-4 cm. long, the upper- most narrowly linear, linear-oblong or linear-lanceolate, with obtuse or acutish apices. Involucres hemispherical or broadly campanulate, tyP- ically small, usually (7-)8-10(-15) mm. wide and (5-)6-8(-10) mm. high including the pales, sometimes subtended by a remote leaf-bract. Phyllaries herbaceous, thin in texture at flowering time, the midri slightly thickened, stramineous to buff and rigid in age, glabrous or THE GENUS MARSHALLIA 97 series, the outer (0.8-) 1-2(-2.5) mm. wide, 4-6(-7) mm. long, shorter than the pales, quite variable in outline: oblong with rounded or obtuse and minutely mucronate apices, linear-lanceolate with acute apices, elliptical with acute apices and white-hyaline borders to the middle and above or rhomboidal, broadest above the middle by conspicuous white- hyaline shoulders, with acute apices or occasionally subulate-tipped. Pales of the receptacle linear, usually merely acute, sometimes subu- late-tipped, 5-7 mm. long, typically glabrous but occasionally sparingly pubescent with scattered hairs, without resin atoms, rigid in age, per- sistent beyond the shedding of the achenes. Receptacle convex and dome-shaped or weakly conical in age, 3-4 mm. wide, about 3 mm. high. Flowers predominantly white or whitish, occasionally very pale lavender. Dried corollas 8-10 mm. long, the tube about 5 mm. long, usually expanded at the junction of the corolla-lobes into a shallow throat 1-2 mm. in length. Anthers usually white or cream-colored, of south-central Georgia (known only from Dodge, Johnson, Tattnall and Telfair Counties); occurring on dry rocky outcrops, dry wire-grass pine-barrens, bases of post-oak sandhills and somewhat moist sandy pinelands. (Map 3 SPECIMENS EXAMINED. Without locality: Muhlenberg Herbarium 599 (pH). Georgia. Dodge Co.—vicinity of Eastman: Ashe Herbarium (ncu); Biltmore Herbarium (Collectors) 2370 (us), 8708 (mun, vs), 8708* (us). Johnson Co.—near Wrightsville: Channell 1963 (to be distributed) ; Harper 1342 (BKL, BM, F, GH, MO, NY, US, wva); Pyron & McVaugh 3081 (ca, Na). Tattnall Co.—flat rocks near Ohoopee River, Harper 1855 (aniz, F, GH, MO, NY, US). Telfair Co.—vicinity of Lumber City: Biltmore Herbarium (Collectors) 8708” (us); Wilbur 3449 (DUKE, Ncsc). Marshallia ramosa is a very localized endemic in south-central Georgia (Map 3), known only from Johnson, Telfair, Dodge and Tattnall Counties. It is isolated geographically from M. caespitosa var. signata, the taxon which it most closely resembles morpho- logically. In fact, both varieties of M. caespitosa occur west of the Mississippi River only. Pratre 8. Marshallia ramosa: Channell 1963, Johnson County, Georgia. THE GENUS MARSHALLIA 99 The relationships of Marshallia ramosa to M. caespitosa var. signata were pointed out by Beadle and Boynton (Bilt. Bot. Stud. 1: 8, 9) in 1901, when the two taxa were first described. According to Beadle and Boynton, M. ramosa differs from both varieties of M. caespitosa in the extremely floriferous habit, the shorter and more obtuse phyllaries and in the smaller pappus scales. These same characters are used in the present treatment to distinguish these species. As originally described and as represented in the type, the phyllaries of M. ramosa are “. . . oblong, rounded at the apex and minutely mucronate, smooth, 1-2 mm. wide, thin in texture at flowering time, the midrib slightly thickened:” The phyllaries of the species as a unit, however, are quite variable in form, in fact, considerably more variable than indicated in the original description. They are sometimes pubescent with scattered monili- form hairs, are frequently rhomboidal in form due to the posses- sion of broad shoulders or wings and are occasionally acute or even subulate at the apices. While some modification of the “glabrous-oblong-obtuse” form represented in the type might be expected to accompany the maturation of the phyllaries, the extreme forms encountered are not easily explainable on this basis. The kinds of variation just mentioned are exemplified by specimens of R. L. Wilbur’s collection number 3449 from Lumber City, Telfair County. It should be emphasized at this point that the element of this species represented by the type is encountered exclusively in certain populations. Living plants were taken from such a popu- lation near Wrightsville, Johnson County (R. B. Channell’s col- lection number 1963), and transplanted to the greenhouse. It is significant that the “glabrous-oblong-obtuse” character of the phyllaries was maintained over a two-year period in the green- house during which time some of the plants flowered twice. As suggested elsewhere in this work, the spring-flowering species Marshallia ramosa with the varieties of M. caespitosa form a closely related species group which is morphologically distinct from the late summer or fall-flowering species group comprised of M. tenuifolia and M. graminifolia, despite the fact that they have in common the narrow, linear-lanceolate leaf form. Moreover, as groups, the two are in large measure isolated geo- graphically. It would appear, however, in contradistinction to the stated divergencies, that certain members of these two groups still retain genetic compatibility. The basis for this reasoning 100 R. B. CHANNELL involves the extreme, atypical morphological element of M ramosa, exemplified by Wilbur's number 3449, already discussed. This element may have been derived through hybridization with M. tenuifolia, that is, from introgression of the characters of M. tenuifolia into M. ramosa. Marshallia tenuifolia possesses all of the characters considered to comprise the atypical element in M. ramosa and hybridization between these species would form a logical explanation for this peculiar variation pattern. The present study indicates that the geographical ranges of the two species (Maps 3 & 4) overlap in Telfair County, the source of the Wilbur collection, and at other points, according to the literature. In general, however, these species are isolated seasonally. Mar- shallia ramosa with M. caespitosa comprise a spring-flowering species-group, while M. tenuifolia with M. graminifolia comprise a late summer or fall-flowering group. Yet, of these species, M. vamosa has a relatively long flowering period which may have in some years overlapped that of M. tenuifolia, especially if the flowering of the latter were occasionally earlier than usual. This would help to explain the localized population-wise nature of the presence of characters of M. tenuifolia in M. ramosa which is clearly not a general feature of the latter as a unit. Otherwise, one must regard the introgression of the characters of M. tenui- folia into M. ramosa as antedating the evolutionary adaptation to the different flowering seasons of the two species, respectively. This, however, probably would have allowed the characters of M. tenuifolia to be considerably more general or widespread in M. ramosa than is now the case. Marshallia caespitosa (Nutt! in litt. 1825) DC. Prod. 5: 680. 1836. was very probably in what is now southeastern Oklahoma. (c—Type, photograph examined; Isotypes—sm, photograph examined; K, PH, specimens examined. Marshallia spiralis Raf. New Fl. 4: 77. 1838. “. . . in Arkansas and west Louisiana found by Binder . . .” (Type not located. THE GENUS MARSHALLIA 101 Perennial scapose or subscapose herb (1-)2-4(-6) dm. in height, the stem arising from a short caudex with coarse fibrous roots, the caudex somewhat fleshy in age and often bearing the vascular fiber of old leaf bases; plants sometimes singly disposed but usually somewhat tufte or cespitose in habit, usually with several offsets borne on the same caudex, each bearing a simple scape or peduncle. Stem very short, with a few reduced leaves at the base of the peduncle, or plants appar- ently stemless, with all the leaves crowded at the base. Peduncle usu- ally more or less stout, 1-4 mm. wide, (8-)15-35(-50) cm. long, sulcate, glabrate to sparingly pubescent below, becoming densely pubescent beneath the with white or tawny, ascending-incurving, moniliform hairs, frequently with 1 or 2 remote, linear leaf-bracts. 102 R. B. CHANNELL e a i ys Plants al (8 bho Herbarinne of CREO A A Mon. Prate 9. Marshallia caespitosa var. caespitosa: Waterfall 11391, Pushmataha County, Oklahoma. THE GENUS MARSHALLIA 103 DISTRIBUTION. Prairies, open woods, fields and rocky hills; occurring locally in Jasper County, Missouri, extending southward through west- ern Oklahoma into northeastern Texas; distributed on sandy soils over much of southeastern Texas and locally in western Louisiana. REPRESENTATIVE SPECIMENS. Missouri. Jasper Co.—High prairies: Alba, Palmer 26946 (mo); Jasper, Palmer 3407 (cu, Mo), 21565 (Ny); Prosperity, Palmer 2153 (GH, MIN, MO, Ny, US), 2153A (GH, NY, Us, WVA); Webb City, Palmer 2015 (cH, ILL, MO, Ny, US). Arkan- sas. Without definite locality: Nuttall (¥, GH, k, PH). Oklahoma. Red River, Nuttall, 1825 (cu—photograph of Type; BM, K, pH—Isotypes). Choctaw Co.—Hugo, Demaree 12591 (oKL, Ny). Creek Co.—Bristow, Williams 1, 24 (pH). Cravin Co.—Stratford, Hudson 22 (oxi). Haskell Co.—Bebb 5468, 8 June 1940 (oKL); 5 mi. s. of Kinta, Hopkins 5127 (OKLA). Pottawatomie Co.—Wanette, Faulkner 10, 140 (oxi). Sem- inole Co.—2 mi. n. of Seminole, Nelson 66 (ox). Sequoyha Co.—Fort Salissa and Illinois River, Engelmann 582 (mo). Texas. Without defi- nite locality: Lindley, 1843 (xk); Jordan, 1869 (P); Hildebrands (vs); Milligan (ariz); Drummond 174 (Ft, G, GH, K, P, PH); Mill Creek, Lindheimer 47 (mo). Austin Co.—Industry, Wurzlow, 1891 (11). Bastrop Co.—Smithville, Morrs, 22 April 1930 (TEx); Bastrop, Duval 5 (us). Brazos Co.—Brazos, Lindheimer, May 1844 (Mo); White Creek, 2 mi. s.e. of College Station, Cory 51569 (cu, smu); College Station, Parks, April 1946 (Tars, TEX). Burleson Co.—9.7 mi. n.e. of Lyons, Cory 5164 (xm, smu, us). Caldwell Co.—near Luling, Perkins & Hall 2863 (cv). Chambers Co.—14.5 mi. n.w. of White Ranch Depot, Parks & Cory 22504 (ras), 22506 (Na). Fayette Co.— Matthes 63 (G). Galveston Co.—Kemah, Fisher, 21 April 1924 (DUKE, us); Nelson, 14 April 1942 (TEx). Gonzales Co.—Gonzales, Schulz 2318 (icv). Grayson Co.—Liberty High School, Gentry 50-267 (OK, PENN, TEX). Hardin Co.—2 mi. n.w. of Sour Lake, Rosier, 24 Mare 1952 (smu). Harris Co.—Houston: Hall, May 1872 (BKL), 365 (F, 6G, GH, ILL, K, MO, NY, PHIL, POM, US), 366 (GH); Lindheimer 23, 32 (mo), 110 (BM, GH, Mo, Ny, P, PH, SMU). Jackson Co.—Ganado, Palmer 9221 (Mo, us). Jefferson Co.—Hamshire, Stuckman 7 (rex); Nome, Fisher 38034 (F, vac). Lamar Co.—Paris, McMullen, 3 May 1926 (TEX). Navarro Co.—Corsicana, Williams 18 (TEx); 3 mi. s.e. of Dawson, 104 R. B. CHANNELL Cory 55630 (us). Orange Co.—Orange, Warren, 4 May 1930 (NA). Polk Co.—Corrigan, Fisher 3486 (Fr). Refugio Co.—Refugio, Benke 5441 (¥, G, GH). Louisiana. Without definite locality: Drummond (Kk); Leavenworth (¥1, GH); Hale (cu, ep). Caddo Parish.—Shreveport, Cocks, 20 April 1916 (mo). Calcasieu Parish.—English Bayou area, Cooley & Brass 4104 (cu, duplicates to be distributed) ; Sulphur, Pal- mer 7700 (mo, Ny, us); Parkins, Pennell 10215 (Ny, pH). Rapides Parish.—Alexandria, Hale, April 19?? (pu). Vernon Parish.—7 mi. n.e. of junction of Haddenburg Ferry, Cooley & Brass 4044 (cu, duplicates to be distributed). In the original description of Marshallia caespitosa, De Candolle (Prod. 5: 680. 1836) ascribed the name to Nuttall in the reference “(Nutt.: in litt. 1825).” On the basis of this refer- ence, however, it is not clear whether Nuttall supplied the name alone, or both the name and description published by De Candolle. The Nuttall type of Marshallia caespitosa, preserved at the Conservatoire et Jardin botaniques, Genéve, has thus far furnished the only clue to the meaning of this citation. The Geneve specimen is the only one of the series of Nuttall isotypes which bears the inscription “Mr. Nuttall 1825,” in addition to “Marshallia * caespitosa, Red River.” Only the latter inscription appears on the remaining isotypes. The inscription “1825" is presumed to indicate the year that Nuttall probably sent the specimen and perhaps information “in the form of a letter to De Candolle. It could not refer to the year of collection, for Nuttall is known to have traveled in the Red River area of the Arkansas Territory (now southeastern Oklahoma) only in the year 1819.'* This is the area from which the isotypes were appat- ently collected. In the absence of information to the effect that Nuttall actually supplied the description of Marshallia caespitosa for publication by De Candolle, the authorities’ names are separated by the word ex. While this is in keeping with the citation as written by Shin- ners (Field and Lab. 17: 174. 1949), it is contrary to the citation currently in general use in which Nuttall alone is acknowledged. Even if Nuttall were definitely known to be the author of the description, the name of the publishing author, being the more important, should be retained (Recommendation 46A, Interna- tional Code of Botanical Nomenclature, 1956). ** A Journal of Travels into the Arkansas Territory during the Year 1819. Philadelphia: printed and published by Thos. H. Palmer. 1821. Reprinted as Volume 13 of “Early Western Travels,” edited by G. H. Thwaites. THE GENUS MARSHALLIA 105 In any case, the Genéve specimen has been selected as the lectotype. Because this sheet bears the inscription “1825,” cited by De Candolle, it is believed to be the one used in drawing up the description. Marshallia spiralis Raf. is clearly a synonym of Marshallia caespitosa Nutt. ex DC. However, this binomial has not previ- ously been interpreted. It seems desirable, therefore, to clarify the matter in some detail. The original description (New FI. 4: 77. 1836) is as follows: “Marshallia spiralis Raf. smooth subscapose, leaves radical linear lanceo- late obtuse uninerve, scape elongate spiraly grooved, pubescent above, LD ie) 3 inches long, scape twisted as some Xurides (not so in M. lanceolata) pedal, flower incarnate.” Inasmuch as no Binder collections have been located during the course of this revision, it has been necessary to rely solely upon the original description for the interpretation of the species. The original description clearly defines the plant as subscapose, with linear-lanceolate, obtuse leaves. The possibilities are thus considered narrowed, for there is only one subscapose Marshallia with linear-lanceolate, obtuse leaves. This species, M. caespitosa, is definitely known to occur in Arkansas and Louisiana and is the only species which is not excluded by Rafinesque’s circumscrip- tion of M. spiralis. On the basis of this information it seems safe to conclude that Marshallia spiralis Raf. represents the same species as that pub- lished by De Candolle as M. caespitosa. Moreover, Rafinesque was apparently unaware of the previous description of M. caespitosa, for, in summarizing the Marshallia species known to him in 1836 (New FI. 4: 77), the latter was not taken into account. The geographical distribution of Marshallia caespitosa is given by Rydberg (Flora of the prairies and plains of central North America, page 839. 1932) as “Ark.-Kans.-Tex.-La.” No specimens were located during the present study, however, which sub- stantiate the report of either variety of this species from Kansas. The Arkansas record for the species is supported by Nuttall col- lections which lack definite locality data. These reports need verification by modern collections. The relationships of the two varieties of Marshallia caespitosa are discussed under var. signata. 106 R. B. CHANNELL 6b. Marshallia caespitosa Nutt. ex DC. var. signata Beadle & F. E. Boynt. Marshallia caespitosa Nutt. var. signata Beadle & F. E. Boynt. Bilt. Bot. Stud. 1: 9, 10, pl. 8. 1901. “Based on A. A. Heller’s no. 1618 from Kerrville, Kerr County, Texas, April 19-25, 1894.” (us—966507— Type, ex Biltmore Herbarium, presented by Mrs. George W. Vander- bilt, 1917, marked “Type Sheet” by C. D. Beadle and F. E. Boynton; Isotypes—ARniIZ—25260 & 58432, BkKL—5917 & 016055, cu, F, GH, ILL, MIN, MO, NY, PH, POM, RM, US.) Perennial, caulescent, linear-leaved herb, (1.5-)2-5(-6.5) dm. in height, the stem from a short crown or caudex with coarse fibrous roots and often the vascular fiber of old leaf bases, the caudex becoming fleshy in age, sometimes 1 cm. thick and 2 cm. long, with numerous short internodes. Stems tufted or solitary, usually cymosely branched below or above the middle and bearing 2-5(-12 or even more) heads, but often simple and monocephalous, the stems striate, somewhat angled and glabrous or glabrate below, becoming more distinctly sul- cate and pubescent near the middle and on the branches, pale green to olivaceous in color, becoming stramineous in age. Peduncles (3-)5- 20(-30) cm. long, suleate, pubescent below, often becoming copiously so beneath the phyllaries, naked or often leafy-bracted at the base. Leaves glabrous, entire, more or less 3-nerved, sometimes apparently 1-nerved, the radical 4-10(-20) mm. wide, (2-)4-8(-16) cm. long with obovate, spatulate or oblanceolate, obtuse blades, contracted below into broad, sheathing, petiole-like bases. The lower cauline leaves usually crowded at the base, 2-10(-12) mm. wide, (2-)5-15(-18) cm. long, with linear-attenuate, linear-oblong or linear-oblong or linear- lanceolate, obtuse, rarely acutish, blades, narrowed below into sheath- ing petiole-like bases about '% as long as the blades. Middle and upper cauline leaves similar to the latter in outline, becoming 1-nerved and sessile above, gradually reduced in size upwards, extending at least to the middle of the stem and usually well up on the branches as reduce leaf-bracts at the bases of the peduncles. Involucres broadly campanu- late or hemispherical, (10-)15-25(-32) mm. wide, 5-15 mm. high including the pales, often subtended by a remote, linear leaf-bract. Phyllaries glabrous, entire, herbaceous, green in color, except for the white-hyaline borders to the middle and above, thin in texture at anthesis, firm and somewhat rigid in age, subequal in about 2 series, the outer linear, linear-lanceolate or lanceolate in outline, 1-2(-3) mm. wide, (5-)7-12(-15) mm. long, usually imbricate at the base but some- times merely approximate, scarcely at all imbricate. Receptacle dome- shaped, conical in age. Pales of the receptacle linear, acute, 6-8 mm. long, persistent beyond the shedding of the achenes. Florets white or cream-colored, rarely pale lavender. Dried corollas 10-12 mm. long, the tube 5-7 mm. long, sometimes slightly dilated above into a shallow throat. Pappus scales (1.5-)2-3(-4) mm. long, white, tawny or ferru- THE GENUS MARSHALLIA 107 gineous in color, imbricate at the base, regularly serrulate or lacerate, acuminate, acute to scarcely acute, minutely scabrous, usually shorter than the mature achene. Achene 3-4 mm. long, obconic or obpyram- idal, 10-ribbed, the ribs hairy with minutely bifurcate trichomes, achene-surface between the ribs concave, beset with minute resin atoms. Flowering in April and May, occasionally into June; fruiting in May and June. (Plate 10.) DISTRIBUTION. Sandy prairies, limestone bluffs, dry hillsides; a com- mon and characteristic plant of chalk and limestone outcrops an calcareous soils throughout central Texas; occurring locally in Ozark County, Missouri, on rocky ground of the “bald knobs.” (Map 3.) REPRESENTATIVE SPECIMENS. From cultivation: Harvard University Botanic Garden, 1884 (GH). Missouri. Ozark Co.—Tecumseh, Palmer 34811 (mo, pH, us). Texas. Without definite locality: Wright (cx); Lindheimer, April 1842 (pH), 1849-1851 (Ny, pom). Rocky soil on the upper Guadalupe, Lindheimer 53 (cH, Mo), Pierdenales, Lind- heimer, distributed under 647, Fasc. IV (cH, kK— hotograph, Mo, PH); Phantom Hill, Hayes 494 (¥, G, Ny); Oak Cliff, Reverchon 532 (Mo). Bell Co.—Crawford, Browning, 9 April 1947 (BayLu); Normand, 21 April 1928 (mo, Ny, TAES, TEX). Bexar Co.—19 mi. n. of San Antonio, Metz 635 (Lcvu, LLC, Pom), 3000 (Ny, pom). Blanco Co.—Blanco, Palmer 12153 (mo, vs). Bosque Co.—Clifton, Albers 47014 (TEX). Burnet Co.—Schaupp, Aug. 1892 (Ny, us). Collin Co.—White Rock School, McCort 1672 (n1sc, TEx). Comal Co.—Comanche Spring [?], Lindheimer 948 (aniz, F, GH, MO, NY, OKL, PENN, PH, TEX, US). Cooke 0.—4.5 mi. n. of Gainesville, Shinners 12456 (smu). Coryell Co.— 6.4 mi. e. of Evant, Parks & Cory 13079 (Taxs); Gatesville, Whitley 502 (ntsc). Dallas Co.—Dallas: Biltmore Herbarium (Collectors) 9301 (MIN, Ny, RM, US); Bush 676 (muN, NCU, NY, US); Lundell & Lundell 9134 (cH, LUNDELL, Ncsc); Reverchon, distributed under Curtiss’ 1525 (F, GH, MIN, NY, PH, UARK, US), 532 (F, MO, PENN, us). Denton Co.—Harris, spring 1926 (TEx). De Witt Co.—Yorktown, Riedel, 20 April 1942 (mo, smu). Ellis Co.—Waxahachie, Turrell, 11 May 1898 (ariz). Erath Co.—Dublin, Maxwell 41 (F); Gough 6512 (TEx, NA). Gillespie Co.—F redericksburg, Bray 269a (us); Otto Mt., Jermy 306 (Fr, Mo). Grayson Co.—Whitewright, Gentry 51-1777 (TEx). Hamilton Co.—13.4 mi. s. of Hamilton, Cory 53782 (GH, Ncsc). Hays Co.—San Marcos, Stanfield (Ny). Hill Co.—Itasca, Smith 22 (TEX); 4.25 mi. s.w. of Milford, Cory 23411 (na). Hood Co.—Gran- bury, Bogusch 46248 (tars); Center Mills, Blackwell 29 (Nrsc, NY). Johnson Co.—13.5 mi. s.w. of Cleburne, Shinners 11259 (Ncsc). Ken- dall Co.—Boerne, Palmer 9850 (man, us); Lindendale, Palmer 9906 (Mo); Waring, Parks ¢ Cory 12981 (cu, TaEs). Kerr Co.—Kerrville, Heller 1618 (us—Type; ARIZ, BKL, CU, F, L, MIN, Y, PH POM, RM, us—Isotypes). McLennan Co.—Crawford-Valley Mills Road, Smith 546 (TEx, varK, us). Menard Co.—Menard: Palmer 11873 108 R. B. CHANNELL (mo, RM); McVaugh & Harvill 8286 (GH, Na, TEX). Parker Co.—5.5 mi. e. of Weatherford, McVaugh 8369 (GH, NA, TEX). Real Co.— Thousand Springs, Parks & Cory 8573, 12660 (TaxEs). Sansaba Co.— Nealley, collector not indicated, 1890 (F). Sutton Co.—29 mi. s.e. of Sonora, Cory, 2 June 1931 (cH). Tarrant Co.—Fort Worth, Eggert, 4 May 1900 (mo); Ruth 475, variously dated (cu, F, GH, ILL, NTSC, NY, PH, RM, US), 983 (Ny). Taylor Co.—Sayles, Parks & Cory 8792 (TAES); near Camp Barkeley on north rocky slope of Edwards Escarp- ment, Tolstead 7182 (Mo, TEX). Travis Co.—Austin: Berlandier 1566 (BM, FI, G, GH, K, P); Hall 366 (BKL, F, ILL, MO, NY, POM, us); Tharp 44061 (CU, DUKE, IND, LLC, MIN, NCU, NY, NTSC, OKL, OKLA, RM, TEX, UARK, VDB, WvA), 46036 (CU, DUKE, GH, IND, MIN, MISS 10, OKL, OKLA, RM, TAES, TENN, TEX, UARK, VDB); York 46037 (DUKE, ILL, MIN , NTSC, OKL, TAES, TEX, UARK). Val Verde Co.—Devil's , MO, NCS River, Orcuff 6207 (mo). Wise Co.—Fouts’ Ranch, McCart 1698 (NTSC, NY). Walker Co.—Hempstead, Hall, 1872 (us). Although Marshallia caespitosa var. signata was first described by Beadle and Boynton in 1901, it had been previously recog- nized taxonomically by Lindheimer. Lindheimer collections dated as early as 1846, for example, carry such inscriptions as “M. caespitosa Var. caule folioso,’ “Variety” and “caule folioso! These specimens were collected in Texas from rocky soil on the Upper Guadaloupe and were distributed under number 647. According to Gray’s “Plantae Lindheimerianae” (Boston Jour- nal Nat. Hist. 6: 231. 1850), those specimens distributed under number 647, Fasc. IV, belong to Lindheimer’s collection of 1847- 1848. Prior to the description of var. signata, the caulescent or branched specimens for the most part passed as Marshallia caespitosa Nutt. e two varieties of Marshallia caespitosa appear to bear the same relationship to one another as far as growth habit is con- cerned as the varieties of M. obovata. Marshallia caespitosa var. caespitosa (Plate 9), which is scapose or subscapose in growth abit, may be considered to have been derived from its caulescent counterpart, var. signata (Plate 10). The differences in growth habit, which admittedly are no more absolute in this case than emphasized for M. obovata, represent the only means thus far discovered for distinguishing the varieties of M. caespitosa mor- phologically. Only by classifying the relatively small percentage of subscapose individuals with the strictly scapose, however, will the distributions of the varieties correspond to that of Map 3 which roughly reflects the geologic picture of the area covered. The vast majority of the specimens examined could easily be THE GENUS MARSHALLIA 109 ee Piet PLANTS OF OKHUTHERN TERA CORO TED 80 ARMOUR) RM BOUNTY, fe Aier Mse Re R 1618 Marghaihe cacspitosa, Nott. emerom son me one Prate 10. Marshallia caespitosa var. signata: Heller, 19-25 April 1894-Isotype, Kerr County, Texas 110 R. B. CHANNELL placed without question in either the caulescent or scapose cate- gory, only a relatively small proportion of individuals represent- ing intermediates. Inadequate or obviously depauperate individ- uals were not taken into account unless they could with certainty be categorized on the basis of the morphological characters they exhibited, apart from any knowledge of the source of such speci- mens. By assembling all of the duplicates of a particular collec- tion, a better picture of the local range in variation was pre- sented and by this procedure, the distinction between the two varieties was established as precisely stated in the key. Shinners (Field and Laboratory 17: 174, 175. 1949), seeking to differentiate between the two varieties on a local basis, utilized the relative difference in the number of cauline leaves below the first branch and the number of heads, even though in the original description of M. caespitosa the plant is patently “. . . caespitosa, caule simplici aphyllo 1-cephalo . . .” While this highly arbitrary, artificial system correctly resulted in the concentration of the range of var. signata in the “Hill Country” of central Texas, a dissected portion of the Edwards Peateau and one of the most notable areas of plant endemism in Texas, the procedure is nevertheless untenable. According to Shinners’ approach a con- siderable proportion of var. signata would be classified with var. caespitosa, with the result that the range of the former is unduly restricted. More natural geographical varieties may be segregated on the basis of distinctions used in the present treatment, dis- tinctions in large part corresponding to those of the original circumscriptions, Precise habitat data are not available for all of the various collections of Marshallia caespitosa examined but the existing data suggest that the respective distributions of the two varieties may be explained on the basis of edaphic adaptation. M arshallia caespitosa var. signata (Map 3), according to Shinners ( Field and Laboratory 17: 174. 1949, as M. caespitosa), is a common and characteristic plant of chalk and limestone outcrops and calcareous soils throughout central Texas. Marshallia caespitosa var. caespitosa (Map 3) occurs on apparently noncalcareous, sandy soils in southeastern Texas, Louisiana and Oklahoma. Al- though southeastern Texas is mostly sandy, there are rather nar- row belts of calcareous clay and also small, scattered outcrops of such soil. This may permit plants of the caulescent variety to appear within the range of the scapose variety. This may be the explanation for the more or less isolated record of M. caespitosa THE GENUS MARSHALLIA Lit var. signata in Walker County Texas, within the range of var. caespitosa. Other examples of this same sort of interfingering of ranges are likely to be discovered with more extensive collecting of the varieties. According to Shinners (personal communication), limestone beds often overlie sandstone in northern Texas and Oklahoma and in eroded country there may be mesas topped by limestone in a predominantly sandy area. The sand, however, is more calcareous than where limestone caps have been entirely re- moved. This would apparently permit Marshallia caespitosa var. signata to occur within the range of var. caespitosa in Oklahoma. Additional collections may substantiate this view also. The dis- junct occurrence of M. caespitosa var. signata in Ozark County, southern Missouri suggests a relict population. The specimens upon which this record is based were originally identified as M. obovata var. platyphylla (var. obovata of this treatment). The specimens, as stated elsewhere, are exceptionally broad- leaved, exhibiting an even closer resemblance in habit and ap- pearance to certain phases of M. Mohrii than to M. obovata. The disjunction between M. caespitosa var. signata populations in Texas and Missouri is very similar to that of the calciphile Juniperus Ashei (J. mexicana of Gray’s Manual). It should be emphasized in connection with the results of the analysis of variation within this complex that the Oklahoma pop- ulation of Marshallia caespitosa var. caespitosa contains a larger proportion of subscapose (as opposed to strictly scapose) in- dividuals than the populations of southeastern Texas, at least as indicated by herbarium collections. The explanation for this is not known. It is a matter of conjecture as to whether or not and, if so, to what extent an established growth habit is conditioned by the influence of the concentration of alkali in the soil. Mar- shallia caespitosa is particularly adaptable to transplant experi- ments by reason of its perennial nature and cespitose habit. Reciprocal transplant studies would help to clarify the genetical and environmental relationships which may exist. It would appear from these considerations that there is a definite correlation between the geologic history of the areas and the present distribution of the two varieties, respectively. The fact that Marshallia caespitosa var. caespitosa in general occupies the geologically more recent area lends support to the supposition that the scapose variety may have been derived from the caulescent. Marshallia caespitosa appears to have had an 112 R. B. CHANNELL Ozarkian origin but may at the same time be related to the Appalachian, broad-leaved species in that prior to the Mississippi Embayment a continuous distribution pattern of the genus may have existed between these two regions. Marshallia caespitosa var. signata survives under ordinary greenhouse conditions on a non-calcareous, garden-type soil mixture and flowers satisfactorily. Individual specimens, how- ever, fail to set fertile achenes. The caudices of plants persisting in the greenhouse over a two-year period become quite thickened and evidently rhizomatous. Greenhouse growth of M. caespitosa var. caespitosa was never well established and the specimens did not constitute a sample. 7. Marshallia graminifolia (Walt.) Small Athanasia graminifolia Walt. Fl. Carol. 200. 1788. (smM—Type, on page 16 Herb. Walt.; Ga—photograph of type examined.) South Caro- lina, Walter. é Persoonia angustifolia Michx. Fl. Bor.-Am. 2: 106. 1803. “Hab. in territorio Tennessee.” (p—Type, Herb. Michx., subsequently labeled “Phyteumopsis angustifolia”; Ga—photograph of type examined. Trattenikia angustifolia (Michx.) Pers. Syn. Pl. 2: 403. 1807. Marshallia angustifolia (Michx.) Pursh, Fl. Amer. Sept. 2: 520. 1814. “In swamps, near Wilmington, North Carolina . . . v. V: Marshallia angustifolia (Michx.) Tratt. Arch. Gew. 1: 109. 1814. Phyteumopsis angustifolia (Michx.) Poir. Encycl. Suppl. 4: 406. 1816. Marshallia graminifolia (Walt.) Small, Bull. Torr. Bot. Club 25: 482. 1898. As to basionym only, the binomial at least in part misapplied. Marshallia lacinarioides Small, Bull. Torr. Bot. Club 25: 482. 1898. “The original specimens were collected at Macon, Georgia, by Mr. [|H. B.] Croom, and in North Carolina (eastern) by Mr. M. A. Curtis. (wy—Syntypes, ex Torrey Herbarium.) The sin le sheet carries two specimens both of which were originally labeled Marshallia angustt- folia. One is marked “(N. Carol. Rev. M. A. Curtis)”; the other orig} nally carried only the collector's name—“Croom,” to which “Macon, Georgia” (thought to be erroneous) was subsequently added in a dif- ferent hand, with different ink. Marshallia graminifolia (Walt.) Small var. lacinarioides (Small) Beadle & F. E. Boynt. Bilt. Bot. Stud. 1: 5, pl. 11. 1901. Marshallia Williamsonii Small, Fl. Southeastern U. S. 1284, 1340. 1903. (Ny—Type, collected by C. S$. Williamson at Wilmington, North Carolina, September 2, 1900.) The sheet is marked “(Type) Mar- shallia Williamsonii Small.” Leafy-stemmed perennial herb, (1.5-)4-8(-10) dm. tall, the stem THE GENUS MARSHALLIA 113 from a short crown or caudex, with coarse fibrous roots more or less clustered at the base and usually bearing conspicuous, coarse vascular fiber of decaying leaf bases, the elongated basal leaves firm-textured and more or less erect. Plants propagating by basal offsets, these usu- ally not present at flowering time. Stem sometimes simple but usually freely cvmosely branched near the middle, the slender, erect, somewhat spreading branches each terminated by a single, long-pedunculate head, the lower portion of the stem glabrous, longitudinally striate, more or less angled, reddish in color near the base, the upper stem stem and branches. Leaves numerous, linear, linear-attenuate or linear- lanceolate in outline, entire, glabrous, asal and lower cauline leaves erect, 1-2 dm. in length, 1-1.5 dm. broad, prominently 3-nerved, attenuate at the apices, with obtuse or acutish tips, gradually narrowed below into broad-margined, petiole-like bases, the cauline leaves sessile, gradually and successively reduced in size upwards to the mere linear, acute, leaf-bracts of the inflorescence branches and peduncles, finally becoming only l-nerved, the upper reduced leaves sometimes with a few moniliform hairs. Involucres hemispherical or broadly campanu- late, (10-) 15-25(-40) mm. wide, (8-)10-15(-20) mm. high including the pales. Receptacle dome-shaped, conical in age. Phyllaries herba- ceous, often reddish in color especially at the apices, sparingly pubes- cent, thick in texture, rigid at least in age, frequently with scattered resin glands, (4-)6-8(-10) mm. long, (1-)1.5-2.5(-3) mm. wide, rhomboidal in outline, broadest above or near the middle, with attenu- 8-10(-15) mm. long, the tube (4-)5-6(-8) mm. long, usually dilated above into a shallow throat. Stigmas divergent, recurved in age, ex- serted beyond the anther-column. Anthers purple or pale lavender, (2-)2.5-3(-4) mm. long, the column and also the filaments exserted bevond the corolla-throat. Pappus scales (1-)1.5-2 mm. long, ovate, acuminate, regularly serrulate, white-hvaline in color, sometimes tawny. Achenes 2.5-3(-4) mm. in length, turbinate, truncate at the summit, 10-ribbed, the ribs pubescent, the achene-surface between 114 R. B. CHANNELL ej ae Gol dotcbeubel inlay iui ina Z wow ye ‘ erat RA EAS Prare 11. Marshallia graminifolia: Channell 1783, Pender County, North Carolina. THE GENUS MARSHALLIA 115 the ribs beset with minute resin atoms. Flowering from July through mid October; fruiting from August into October. (Plate 11.) DISTRIBUTION. Low pine barrens, pitcher plant bogs, edges of shrub bogs, wet savannas, pocosins, sandy ditches and road shoulders, sphagnous bogs; distributed throughout the C ata Plain of North and South Carolina, reaching the Piedmont only in very local Coastal Plain-like habitats. RESENTATIVE SPECIMENS. North Carolina. Without definite lo- cality: McCarthy 112 (¥F, us), 1884 (F, Ny, us), 1885 (alug, ¥, ILL, MISSA, NCU, US); McCarthy of Bless (min). County uncertain: Rebel Retreat, Hyams, June 1880 (¥); near Kirkland, Rhoades, ANE: 1932 (GH); Pineway, Schallert, “4 Aug. 1933 (Ny). Beaufort Co.— mi. s.e. of Chocowinity, Fox 3186 (GH, IND, NCSC, SMU); Nashines Freeman, 18 Oct. 1930 (Na). Bladen Co.—n. of White Lake, Correll & Blomquist 2569 (puKE); 2.1 mi. n.e. of Columbus, Fox & Whitford 1869 (ncsc). Brunswick Co.—2.5 mi. s. of Crissettown, Godfrey & Fox DUKE, GA, GH 5 SM e tributed); near Supply, Channel 1778, 1780 (to be distributed). Car- teret Co.—Newport, Godfrey & Blomquist 49795 (GH, Na, NCSC); between Beaufort and Atlantic, Blomquist 11299 (DUKE, MO); Aiea Channell 1668 (to be distributed); Beaufort-Moorehead City Ai Channell 1693 Se - Meese eas § hpi ee Channell 1898, 1899 ie be distributed). n Co.—Rt. 211, 2.1 mi. n.e. of Columbus Co. line, Fox ¢& dteenh pi 369 (LAs, smu). Columbus Co.—Delco, God- frey & Fox 49755 (puKE, MIN, Ncsc); Nakina, Schallert, 26 Aug. 1934 (Np, NY). Craven Co.—Newbern, Croom (GH); Newbern Airport, Blomquist 16429 (puxE). Cumberland Co. —Favetteville, Biltmore ig (Collectors) 14848 (us); 3 mi. w. of Stedman, Fox 1785 (Ncsc). Duplin Co.—8 mi. w. of Richland, Godfrey 5864 (cH, us); between hd oso and Warsaw, Channell 1793, 1794 (to be distri- buted); between Bunnlevel and Richland, Channell 1796 (to be sibel) Guilford Co.—High Point, Shave, 1886 [P] (cH). Har- nett Co.—between Erwin and Dunn, Correll & Blomquist 2555 (DUKE); Pineview, Channell 2863 ue be sip nee Iredell Co.—States- "ie oe mi. n.w. ok Ric badd Boma 15357 (nein, PENN). Peiilex Co. —Burga aw, Hyams, Aug. 187 MO, NY, PENN, us); Rt. 421, | mi. from Sampson Co. line, ape y ee Fox 48715 (FLAS, GA, NCSC, 116 R. B. CHANNELL ncu, sMuU); 10 mi. s. of Holly Ridge, Channell 1722 (to be distri- buted); between Burgaw and Wallace, Channell 1783 (to be distri- buted). Pitt Co.—3 mi. e. of Grimesland, Fox 3181 (GH, MIN, NCSC, smu). Richmond Co.—Smith, 30 July 1933 (puke). Sampson Co.— Roseboro, Godfrey 5710 (puKE, GH). South Carolina. Without defi- nite locality: Curtis (mo). Allendale Co.—Ulmers, Bell 5190 (Gu, Ncu, duplicates to be distributed). Berkeley Co.—Cooper River, Wallace (CHARL); Santee Canal, Ravenel, Aug. 1??? (cHARL); 6 mi. ne. ot Summerville, Wilbur & Webster 2848 (Ncsc, smu); Monks Corner, (CU, DUKE, F, GH, MO, NY, TENN, us). Horry Co.—Conway, Schallert, 1 Aug. 1940 (uark); Coker & Coker, 25 July 1946 (Ncu). Lexington Co.—Gaston, Wherry, 19 July 1932 (us); 5 mi. s. of Columbia, Godfrey & Tryon 1267 (cu, DUKE, F, GH, MO, NY, PENN, TENN, US). Orangeburg Co.—Orangeburg, Hitchcock, Aug. 1905 (1). Richland Co.—Colum- bia, Harhbemock [?] (cHart—Herb. EIl.). This species was first described from South Carolina by Walter in 1788 as Athanasia graminifolia. Historically, the epithet was almost completely displaced for over a century. From the brief original description, supplemented by a photograph of the type. it has been possible to confirm the identity of the species. It is, so far as the present study indicates, the only narrow-leaved Marshallia occurring in South Carolina. Moreover, the present study has shown this species to be restricted in distribution to the Carolinas, where it is confined largely to the Coastal Plain proper, but occurs also in the Piedmont in very localized, Coastal Plain-like habitats. In 1803, Persoonia angustifolia was described by Michaux from Tennessee, an area in which no narrow-leaved Marshallia has subsequently been found. Insofar as the photograph of the type reveals, it is clearly the same at Athanasia graminifolia of Walter. (The type sheet, incidentally, did not originally carry the Michaux binomial but was subsequently labeled “Phyteumopsts angustifolia.” ) Moreover, the original description of Persoome angustifolia compares very closely with that of A. graminifolia and could apply to no other species. In equating the names, it 1s suggested that the apparent discrepancy “Territorio Tennessee, THE GENUS MARSHALLIA LT given by Michaux as the “habitat” of P. angustifolia, may have resulted through an error in labeling. The original collection upon which P. angustifolia was based probably came from North Caro- lina where the species is well represented and where Michaux is known to have collected. The adoption of the Michaux epithet for the species up to 1898 is of considerable importance, for its application was not con- fined to the narrow-leaved species of the Carolinas. It came to apply also to the coastal, narrow-leaved plant farther southward, a species recognized here as Marshallia tenuifolia Raf. At this point Small (1898) made the correct nomenclatural combination, M. graminifolia, erroneously applying it, however, to a more southern species with obovate basal leaves, the range of which was reputed at that time to extend northward into the Carolinas. At the same time (1898) Small described Marshallia lacinario- ides as new. This species was based on the two syntypes indicated in the synonymy above. One of the specimens was collected by Curtis in North Carolina; the other by Croom at Macon, Georgia. Apparently on the basis of these two specimens alone, Small gave the range of the species as extending from North Carolina, along the middle country, to Georgia. The species was reputed to differ from M. graminifolia on the basis of the smaller heads and a growth habit suggestive of certain “Lacinaria” species of Liatris, whence the specific name. The erect, elongate, basal leaves, with their prominently 3-nerved blades emphasized by Small, while imparting a distinctive aspect to certain individual specimens, represents a character which has not been found to segregate this so-called species, nor, indeed, has the smaller sized heads. In fact, all intergradations of the “graminifolia” and “lac- inarioides” types were found to exist in each of several different populations. Marshallia lacinarioides is thus considered to be taxonomically inseparable from M. graminifolia, a species already shown to be confined to the Carolinas. This casts serious doubt, therefore, upon the validity of the collection data “Macon, Georgia” included on the label of the Croom specimen. In fact, this information appears to have been added to the label at a time subsequent to the collector's name, for these data appear in distinctly different handwritings and in different inks. In view of the fact that Croom is known to have collected the same spe- cies at New Bern, North Carolina, it seems much more likely that this syn specimen was itself collected in North Carolina and subsequently mislabeled, especially since no other such 118 R. B. CHANNELL specimens have been reported from Georgia. This view is sup- ported by an authentic Croom specimen in the Gray Herbarium labeled “New Bern” and by the inclusion of the species in “A Catalogue of the Plants of New-Bern, North Carolina” (with remarks and synonyms) by H. B. Croom (John Torrey, Edi- tor), New York, 1831. In the only revision of the genus, Beadle and Boynton (1901) upheld Marshallia lacinarioides but reduced it to varietal status under M. graminifolia. This change illustrated the type of con- fusion which has surrounded M. lacinarioides since its descrip- tion and which has persisted up to the time of the present treat- ment. It illustrates further that Beadle and Boynton did not have the correct concept of Walter’s Athanasia graminifolia, for they erroneously applied that epithet to a species not even represented in the Carolinas. In 1903, Small described still another species, Marshallia Williamsonii, from Wilmington, North Carolina. The range of this species was given by Small as eastern North and South Carolina, the very area of the range of M. graminifolia. The minor quantitative characters by which the author distinguished this new species from M. lacinarioides are not considered to be of sufficient magnitude to warrant taxonomic recognition in any status. The present study has shown, especially in the areas near and around Wilmington, that specimens of M. graminifolia are oftentimes considerably taller, more branched and larger-leaved. This is particularly true of specimens growing in sphagnous situa- tions in the edges of shrub bogs. Such specimens usually possess more heads and floral parts of correspondingly larger size, the very characters used by Small to distinguish his M. lacinarioides from M. Williamsonii. The shape of the pappus scales used by Small to distinguish these species has proved too variable in this com- plex for any taxonomic reliance. Thus, Persoonia angustifolia, Marshallia lacinarioides and M. Williamsonii are treated as synonyms of Athanasia graminifolia. The combination M. graminifolia (Walt.) Small is in no way applicable to the species M. tenuifolia Raf. of this treatment. It is therefore restored in the present treatment to the taxon upon which the epithet was originally based. Finally, it is not surpris- ing that practical difficulties have arisen in attempting: to apply three names to the component variation of a single species exhib- iting a more or less random assortment of quantitative variation. It will be noted upon referring to Maps 1-4 that most of the THE GENUS MARSHALLIA 119 species of Marshallia exhibit allopatric distributional patterns. In North and South Carolina, however, M. graminifolia (Map 4) occurs over the same general geographical area as M. obovata (Map 2). Well-defined species both morphologically and in season of flowering, they occupy distinctly different ecological habitats within the area in question. The source of two specimens of Marshallia graminifolia (Po, MIN) collected by M. E. Hyams and labeled “Statesville” has been questioned. Statesville, Iredell County, in the Piedmont of North Carolina, was Hyams’ home and place of business as a seedsman and druggist and served as headquarters for rather extensive plant collecting. Although the locality notation may have been intended to indicate merely the collector's address, it Suggests that the specimens were collected in that vicinity. If accepted as authentic, these records indicate the disjunct occur- rence of the species outside its normal range on the Coastal Plain, well within the Piedmont, a distribution which is not entirely unlikely. As a matter of fact, pockets of Coastal Plain vegetation, including species of other genera as disjuncts, are known to persist in the vicinity of Statesville at the present time. On the basis of other collections, these habitats were known to Hyams, doubtless in what may be assumed to have been a less ravaged or disturbed state. Judging from the more or less isolated existence of M. graminifolia in other such areas, as, for example, Nash County, North Carolina, it is not inconceivable that the species at one time existed as disjunct colonies around Statesville also, but subsequently disappeared. In the absence of other col- lections or data to substantiate it, the Statesville record has not, however, been included as a part of the geographical range of the species. It is nevertheless intriguing and should not be com- pletely ignored. It is unfortunate that the specimens were not accompanied by additional data. Marshallia graminifolia grows well and flowers profusely under humid greenhouse conditions in sandy soil of a high organic content, when supplied with plenty of moisture. One of the most handsome species because of the purple or lavender heads, it grows naturally under moist acid conditions in pineland savan- nas as well as in the partial shade at the edges of shrub-bogs and pocosins. Variation in the size of leaves, heads and other parts of the plant is evidently controlled to a large extent by ecological factors. Neither narrow-leaved forms nor those with relatively small heads could be segregated satisfactorily as geo- 120 R. B. CHANNELL graphical varieties. The perennial duration of these plants may account for a certain amount of the quantitative variation pre- vailing. Older plants are often very robust while seedlings or relatively young plants are correspondingly smaller in size. 8. Marshallia tenuifolia Raf. Marshallia angustifolia (Michx.) Pursh var. cyananthera EIl. Sk. 2: 317. 1823. “. . . collected by Dr. Baldwin, near St. Mary's, Georgia.” (CcHAaRL—Type, Herb. Ell.) Marshallia tenuifolia Raf. New Fl. 4: 77, 78. 1838. “—found by Baldwin on the sea shore of Florida and Georgia . . .”_ (pwce—Lecto- type, labeled “Georgia, Dr. Baldwin”.) 18 Leafy-stemmed herb, strict in habit, (2-)4-10(-12) dm. in height, the stem from a perennial crown or caudex with coarse fibrous roots and spreading, thin-textured, obovate or spatulate, obtuse radical leaves. Plants solitary or propagating by basal offsets and thus often somewhat clustered in disposition. Stem sometimes simple but usually branched near the middle, the branches erect and scarcely at all spreading, each terminated by a single long-pedunculate head, primary stem furrowed with longitudinal channels or grooves, somewhat angled, glabrous, reddish at the base, greenish or olivaceous above in drving, sulcate and sparingly pubescent with jointed (moniliform), incurved, purple-colored hairs, stems leafy throughout, the leaves numerous, gradually reduced upwards. Peduncles erect, long and_ slender, the mere linear-subulate leaf-bracts of the peduncles. Involucres hemi- spherical or broadly campanulate, globose in age, (0.5-) 1.5-2.0(-2.5) em. wide, (0.5-)1-1.5(-2) em. high including the pales. esis Sree dome-shaped, strongly conical in age. Phyllaries herbaceous, subequa in about two series, often reddish in color especially at the apices, . a «3% . ‘ . c discussion under M. graminifolia for the erroneous application of th ; ; “oli 18 ee i epithets “angustifolia” and “graminifolia” to M. tenuifolia. THE GENUS MARSHALLIA 121 sparsely pubescent, frequently with scattered resin glands over the outer surtace, rigid, usually winged below the middle with white- hyaline or somewhat scarious margins, these sometimes remotely toothed or erose and occasionally with a few marginal trichomes, linear- lanceolate, lanceolate or somewhat rhomboidal in outline, with strongly subulate apices, (2-)4-5(-6) mm. long, (1-)1.5-2(-2.5) mm. wide. Pales of the receptacle linear-attenuate, strongly subulate-tipped, (4-)5-6(-7) mm. long, longer than the phyllaries, green in color, often with reddish apices, frequently with scattered resin glands on the outer surfaces, persistent beyond the shedding of the achenes. Florets usu- ally purple or pale lavender in color. Dried corollas (6-)8-12(-14) mm. long, the tubes (3-)4-5(-6) mm. long, sometimes dilated upwards into a shallow throat. Stigmas divergent, recurved, exserted beyond the anther-column; the style usually included. Anthers purple, “blue” or pale lavender in color, 2-2.5 mm. long, exserted beyond the corolla- tube. Pappus scales (0.7-)1-2(-2.5) mm. long, shorter than the achene, ovate-acuminate or merely acuminate from a broad base, regularly serrulate, occasionally lacerate, white-hyaline in color, some- times tawny in age. Achenes (1.5-)2-2.5(-3) mm. long, turbinate, truncate at the summit, 10-ribbed, the ribs pubescent with hairs shorter than the distance between adjacent ribs, the achene-surface between the ribs beset with minute resin atoms, sometimes copiously so. Flow- ering from July through September, occasionally in January and June; fruiting from August into October. (Plate 12.) DISTRIBUTION. Wire grass-longleaf pine savannas, moist pine barrens, flatwoods, edges of bavheads and shrub bogs, low swampy areas and pitcher plant bogs; occurring in southern Georgia and Florida, extend- ing westward along the Gulf Coast into eastern Texas. (Map 4.) REPRESENTATIVE SPECIMENS. Georgia. Without definite locality: Baldwin (wwe); Cleveland (ras); Harris, 5 Sept. 1930 (ntsc); Le Conte (pH); Nuttall (pu). Appling Co.—Baxley, Biltmore Her- barium ( Collectors) 2807" (us, RM). Bullock Co.—Statesboro, Jones, summer 1923 (cu). Camden Co.—St. Marys, Baldwin (cHart—Type of Marshallia angustifolia var. cyananthera Ell.). Charleston Co.— Trader’s Hill, Small, 24-26 July 1895 (ny, us). Clinch Co.—Homer- ville, McVaugh 5288 (Na). Colquitt Co.—Moultrie: Svenson 6941 (ND, BKL); Channell 3276 (to be distributed). Glynn Co.—3 mi. n. of Brookman, Wiegand & Manning 3448 (cu, GH, pom). Irwin Co.— Oscilla, Harper 1416 (¥, GH, Mo, Ny, Us). Coffee Co.—Harper—photo- graph, mounted on Irwin Co. collection 1416 (GH). Lowndes Co.— 123. R. B. CHANNELL 5100 (Ny), 5095 (GH, Mo, Ny). Wayne Co.—Gessup, Biltmore Her- barium (Collectors) 2807‘ (Ny, us). Worth Co.—8 mi. s. of Sylvester, Svenson 7268 (BKL, ND); Poulan, Pollard & Maxon 550 (GH, Ny, US). Florida. Without definite locality: Baldwin (cHarL), Chapman (Fr, GH, MO, NY, OKL, us); Buckley (GH, Mo); Torrey (pH), 3119 (GH); Curtiss (F), 1875 (us). County uncertain: Fort Craine, Rugel 244 (F, MO, Ny, us). Alachua Co.—Gainesville, Murrill 136 (us, Mo); Alachua-Putnam Co. line, Channell 2098, 2099 (to be distributed). Baker Co.—5 mi. n. of McClenny, O’Neill, 12 Aug. 1929 (us, Lcu). Bay Co.—Panama City Beach, Godfrey & Kral 55056 (cu, duplicates to be distributed). Bradford Co.—Lawtey, Murrill, 9 Sept. 1940 (FLAs). Brevard Co.—Okeechobee region, Fredholm 5878 (GH, MO, us). Calhoun Co.—Altha, West, 9 Aug. 1931 (FLas, TEX). Clay Co.— Doctors Inlet, Murrill 302 (mo, us). Columbia Co.—Lake City: Rolfs 370 (FLAS, F, ILL, MO), 406 (F), 973 (FLAS); Nash 2221 (CONN, PH, us); Hitchcook, June-July 1898 (r, Mo). Duval Co.—Jacksonville: Drummond, Jan. 1835 (cH); Curtiss, Aug. 1876 (pac), 1525 (BKL, CU, F, FLAS, GH, LCU, MIN, MISSA, MO, NY, PH, SMU, US, YU), 4493 (MO, ny, uS), 5109 (cu, F, GH, MIN, Ny, Us). Baldwin: Mohr, 29 July 1880 (pH, us); Nash 2318 (¥, FLAS, GH, MIN, MO, NY, PH, POM, US). EsScam- bia Co.—5 mi. w. of Pensacola, Morgan, 12 Aug. 1941 (GH, LCU, US); near Fla.-Ala. state line, Channell 3262 (to be distributed). Flagler Co.—St. John’s Park, West & Arnold, 10 Oct. 1940 (FLAs, pac). Frank- lin Co.—Apalachicola, Biltmore Herbarium (Collectors) 2807 * (GH, MO, Ny, uS), 2807 (GH, MIN, Mo, Ny, us). Gadsden Co.—10 mi. w. 0 Quincy, Arnold, 20 Aug. 1940 (rias). Highlands Co.—4 mi. s.w. of Childs, Correll 6189 (nuke, GH, IND, NA); Highlands Hammock, Channell 2031, 2035 (to be distributed). Indian River Co.—8 mi. w of Vero Beach, Channell 2077 (to be distributed); 8 mi. e. of Yeehaw Junction, Channell 2082, 2083 (to be distributed). Jackson Co.—Mari- anna: Weigand & Manning 3450 (cu, cH); Channell 3273 (to be dis- tributed). 11 mi. w. of Chattahoochee, Thorne 5962 (cu, GA, GH). Liberty Co.—Bristol, Biltmore Herbarium (Collectors) 2807! (MIN, US). Nassau Co.—Callahan, Knight, 28 July 1941 (Fxas). Okaloosa Co.— Crestview, Hood 3138 (Frias). Osceola Co.—Kissimmee: Singeltary 496 (DUKE); Schallert 120 (FLas). Vicinity of Yeehaw Junction, Chan- nell 2085, 2089 (to be distributed). Putman Co.—between Palatka and Interlaken, Correll 6425 (pUKE, NA, NCSC); Palatka, Channell 2094 (to be distributed). Santa Rosa Co.—Milton, Ford 4071 (cx). St. Lucie Co.—between Okeechobee City and Fort Pierce, Brass 20569 (GH). Union Co.—Lake Butler, O’Neill, 2 Aug. 1929 (Frias). Volusia Co- National Gardens, Butts, 22 July 1943 (cH). Walton Co.—Argyle, Curtiss 5932 (BKL, CONN, CU, F, FLAS, GH, ILL, MIN, MO, NY, PAC, us). Washington Co.—Rock Hill, Kral 3532 (cx). Alabama. Baldwin Co.— Southport, Webster & Wilbur 3555 (Gu, Ncsc, smu); 8 mi. west of Ala.-Fla. state line, Rt. 90, Channell 2111 (to be distributed). Coving- THE GENUS MARSHALLIA 123 ton Co.—McRae, Pennell 4610 (PENN). Escambia Co.—Flomaton, Biltmore Herbarium (Collectors), 15 Aug. 1899 (us). Houston Co.— Dothan, Blanton 58 (GH, Lcu, NA, ND, Ny, RM). Mobile Co.—Mohr, Aug. 1870 (stu); Mobile, Mohr, 15 July 1875 (atu), 1879 (us), 1880 (PH); Williams, 28 Aug. 1951 (uaLA). Mississippi. Jackson Co.—4 mi. e. of Moss Point, Webster & Wilbur 3464 (cu, Ncsc, smu); Channell 2142, 2258, 2286, 2314, 2432, 2437, 2444, 3260 (to be distributed). Louisiana. Without definite locality: Hale (GH); Mead (F). Beau- regard Parish.—10 mi. s. of De Ridder, Correll ¢ Correll 9670 ( DUKE). Calcasieu Parish.—Lake Charles, Cocks, Sept. 1914 (Np). Natchi- toches Parish.—Kisatchie, Correll ¢ Correll 9787 (DUKE, GH, NA). Rapides Parish.—Alexandria, Hale (No), 1840 (us). St. Tammany Parish.—St. Tammany, Cocks, Sept. 1900 (No). Texas. Anderson Co.—Grapeland Bog, Le Sueur & Smith, 7 July 1935 (Tex). Hardin Co.—Kountze-Saratoga Road, Tharp, Follansbee & Thompson 51-1502 (MIN, PENN, TAES, TEX); 2.2 mi. w. of Silsbee, Cory 49895 (cH, smu, us). Jefferson Co.—Beaumont, Smith, 23 Oct. 1922 (Tex). Newton 0.—5 mi. e. of Kirbyville, Cory 49802 (cu, RM, sMu, Us). Orange Co. —Vidor, Tharp 2592 (TEx, us). Trinity Co.—Trinity, Tharp 937 (us). Tyler Co.—17 mi. s. of Woodville, Cory 49959 (GH, us). This species was first described by Elliott in 1823 as Marshallia angustifolia (Michx.) Pursh var. cyananthera. The original de- scription was based upon material collected by Baldwin near St. Marys, Georgia and, in an appended note, Elliott states that the new variety may represent a distinct species. The leaves of the plant are described as linear-lanceolate and conspicuously 3- nerved. The type (cHARL), however, has cauline leaves which are linear and 1-nerved, with only the basal leaves conspicuously 3-nerved. In 1836 Rafinesque described Marshallia tenuifolia. This spe- cies, like var. cyananthera above, was based upon material col- lected by Baldwin on the coast of Florida and Georgia. Accord- ing to Rafinesque, it is more closely related to M. cyananthera (M. angustifolia (Michx.) Pursh var. cyananthera] than to M. angustifolia [M. graminifolia (Walt.) Small of this treatment]. Although no Baldwin specimen has been located which Rafines- que is definitely known to have examined, it seems safe on the basis of the close similarity of the original descriptions to con- clude that the Baldwin specimens examined respectively by Elliott and Rafinesque were either duplicates of the same col- lection or otherwise very similar. The avowed close relationship of Rafinesque’s M. tenuifolia to M. cyananthera suggests that this interpretation is correct. The close similarity of two other 124 R. B. CHANNELL Baldwin specimens to the Elliott type, marked “Georgia” (pwc) and “Florida” (cHarL), further supports this conclusion. In any case, the two descriptions do not differ in essential details. The only character of M. tenuifolia which Rafinesque considered to be diagnostic is “leaves quite slender uninerve,” a feature which is shared as shown above by the Elliott type. The two names, var. cyananthera and M. tenuifolia, are thus treated as synonyms. The taxon in question is recognized in the present study as a species, highly variable as to quantitative characters, distinct from Marshallia graminifolia. As indicated in the discussion of the later species, the names M. angustifolia and M. graminifolia were erroneously employed up to the time of the present study to the species in question. It is clear, however, that that name has no application outside of the Carolinas so far as present distributional records indicate. The selection of the correct name for this species presents cer- tain problems which are not clearly covered by the International Code. Specifically, the present case is complicated in that Rafines- que, in the original description of Marshallia tenuifolia, acknowl- edged the Elliott epithet cyananthera in the following phrase in pointing out the affinities of the new species: “... nearer to M. cyananthera than M. angustifolia.” It is impossible from. this statement to determine whether or not Rafinesque intended to introduce the new combination Marshallia cyananthera, although one might assume that he did not. The problem is not clarified by Article 33 which states that “A name is not validly published . . when it is merely mentioned incidentally.” According to this article, “By ‘incidental mention’ of a new name or combination is meant mention by an author who does not intend to introduce the new name or combination concerned.” It is hardly possible, therefore, to determine whether the combination “M. cyanan- thera” as used by Rafinesque is validly published or not. If this reference constitutes valid publication, the two specific epithets date simultaneously from the same published article and the matter is thus one of choice by the author first uniting the taxa; if it does not, then tenuifolia, being the earliest available epithet in the rank of species, automatically becomes the correct name. The present recourse has been the more or less arbitrary selec- tion of Marshallia tenuifolia Raf. for the species. Although diffi- cult to typify, this name, in the opinion of the present writer, 1S to be preferred because it has not been confused through erro- neous application to any other taxon. This is not true of the epithet THE GENUS MARSHALLIA 125 cyananthera, however, as the following paragraph illustrates. Marshallia angustifolia (Michx.) Pursh var. cyananthera EIL, as erroneously interpreted by Fernald (Bot. Gaz. 24: 435. 1897) and various subsequent authors, including Beadle and Boynton, refers only to a portion of the species treated here as M. tenuifolia Raf. The name was inadvertently applied to the very slender plants of the species first found by A. H. Curtiss (no. 5932) in “seepy pitcher-plant bogs” in Liberty and Walton Counties, west Florida, which are not typified by any Baldwin specimen. This was a very logical mistake in view of the fact that Elliott's de- scription of var. cyananthera was naturally restricted, evidently based solely upon the Baldwin material, and did not circumscribe the species as such. It is an even more logical mistake when one recalls from the discussion of the preceding species that the name M. angustifolia for many years erroneously included the larger portion of the species under consideration here (M. tenuifolia Raf.). It is not surprising, therefore, that the slender, bog Marshallia discovered by Curtiss completely usurped the name and description of Elliott’s var. cyananthera to the present time. Nomenclaturally, it is clear that if the unit of variation exemplified by Curtiss’ collections is ever recognized taxonomi- cally, the epithet cyananthera cannot be employed for it. The Marshallias with attenuate or subulate-tipped phyllaries and grass-like leaves form a complex distributed over the outer Coastal Plain from North Carolina southward into Florida and westward into Texas. Considerable confusion has resulted from the taxonomic interpretation of the component elements. Taxon- omists have perhaps been unduly impressed by morphological variation as expressed by local populations. Disregarding minor quantitative variation, however, the complex, when viewed geo- graphically from north to south, is seen to consist of only two, but highly variable species. Marshallia graminifolia is the north- ernmost species, restricted, so far as is known, to the Carolinas. Marshallia tenuifolia comprises the remainder of the complex. Herbarium specimens of these two species are sometimes dif- ficult to distinguish, especially when they do not provide evi- dence as to the form of the basal leaves or are otherwise inadequate. In nature, however, Marshallia tenuifolia (Plate 12) consistently possesses spreading basal leaves with obovate, spatu- ate or oblong, obtuse blades, distinctly unlike those of the stem proper. Such leaves are never produced by M. graminifolia (Plate 11), all the leaves of which have blades of an elongate, 126 R. B. CHANNELL (ere fi faye 2 12. Marshallia or Fig. 1, ering 2098, Alachua County, Florida: Fig. oy Channell 3273, Jackson County, Florida; . 3, the so-called ‘“cyanz anthera”’ variant, Channell 2111, pare. Collie, Al abe THE GENUS MARSHALLIA 197 linear-attenuate type. It is largely because of the absolute nature of this morphological discontinuity that the two taxa are recog- nized here as distinct species. Insofar as the present distributional records indicate, the Savannah River forms a sharp line of de- marcation between the ranges of the two species. It remains to be seen, however, if the ranges actually overlap in that area and whether or not the Savannah River serves as an effective barrier. Whether or not the history of the Savannah River watershed is in any way to be considered correlated with or the effective cause of the successful differentiation of the two species as they now exist, is purely speculative. Of the two species, Marshallia tenuifolia exhibits considerably more diversity in growth-habit. Local variation of this sort, the nature and range of which are shown in part in Plate 12, is not surprising for a species distributed more or less continuously from Georgia southward into Florida and westward into eastern Texas. The so-called “cyananthera” type of variant, collected in Baldwin County, Alabama (Channell 2111), is represented by the two specimens of Plate 12, Figure 3. These represent the same identical type of specimens as those collected by Curtiss in Walton County (Curtiss 5932) and Liberty County, Florida, reported by Fernald. This extreme form is evidently restricted to open, bog-like terraces and slopes in the pinelands of southern Alabama, extreme southern Georgia and western Florida, where the seepy coarse sandy soil conditions produce habitats which support Sarracenias, Droseras, Tofieldias, Xyris species, etc. Al- though local, these distinctive habitats are oftentimes quite ex- tensive, in some cases covering several acres. As may be expected, therefore, the Marshallia in question occurs within these areas as relatively uniform populations of hundreds of individuals. us, the extreme “cyananthera” type of variation is morpho- logically and ecologically distinct from the more widespread, sympatric element illustrated in Figure 1. Despite this, however, it has been impossible on a morphological basis to delimit the element satisfactorily as a separate taxonomic unit. Taking all of the available herbarium material into account, the “cyanan- thera” variant loses its identity as such. It appears to be con- nected to the remainder of the species by individuals such as that shown in F igure 2, to form a more or less complete intergra- dation series, no segment of which can be delimited taxonomically or segregated geographically. 128 R. B. CHANNELL Greenhouse studies involving the growth of sample specimens of two contrasting elements of the complex, although inadequate for establishment of general conclusions, indicate that the two elements tested are at least to some extent genetically adapted to their respective habitats. Specimens of the “cyananthera” ele- ment (Channell 2111) were collected from an open pitcherplant bog in Baldwin County, Alabama. Tall, robust specimens, rep- resenting the other, more widespread element (Channell 2031), were collected in the moist, shrubby flatwoods of Highlands Hammock, Highlands County, Florida. Transplanted on sods to the greenhouse and grown under moist, humid conditions of partial shade, favoring the Highlands type habitat, marked dif- ferences in the shape, size and quality of the basal leaves soon became evident, differences which persisted over the three-year period they were observed. The Highlands specimens produced spreading basal leaves of an oblong-spatulate shape and of such thin texture as to render them subject to prompt wilting under conditions of low water supply. The “cyananthera” specimens produced basal leaves of a shorter, more ovate-spatulate shape, and of a distinctly firm, glossy texture, features apparently re- lated to the resistance of comparatively long periods of low water supply without wilting. The spindly, single-stemmed, monoceph- alous habit, typical of “cyananthera” specimens in nature, was lost by the specimens tested during the period under which they were grown and observed. At the end of the period they had assumed essentially the same status and general appearance of the specimen shown in Plate 12, Figure 2, but with the production of a branched stem and two to four heads instead of only one. The heads were considerably larger than those shown in Figure 3, but did not attain the size of the larger heads of the specimen shown in Figure 1. The Highlands specimens retained in the greenhouse the more or less strict, typically monocephalous habit of growth as produced under natural conditions, but they did not attain maximum field height. It is safe to conclude from these observations that the majority of the “cyananthera” specimens, occurring as more or less uni- form populations under field conditions, do not attain their potential maximum size. This would appear to be due to en- vironmental conditions imposed by the local habitats to which the “cyananthera” variant is apparently restricted. It is possible, therefore, that individuals dispersed as seeds to different habitats could become considerably larger, branched and several-headed. THE GENUS MARSHALLIA 129 Essentially no field data are available which would indicate the nature of variation encountered over the area between the typical “cyananthera” habitats, usually occurring as terraces on gentle slopes, and the low areas near streams, in relatively close proxim- ity, where considerably larger, much-branched specimens occur. Nor are field data available on the variation encountered be- tween these and still other habitats supporting the plants, such as moist, wire-grass savannas, moist, shrubby drainage draws within turkey-oak sand-hills, well-drained road-shoulders, ete. Collectively, however, the herbarium material indicates that there is no sharp line of distinction on any basis between the typical “cyananthera” element and the remainder of the complex. Qualitative features observable in the living “cyananthera” speci- mens are obscured in herbarium material and are of little or no assistance as distinguishing criteria. No quantitative or qualita- tive floral or vegetative characters have been found to segregate the “cyananthera” element or any other element of the species as a taxonomic unit. Despite this, however, the “cyananthera” element exists, as stated previously, in distinctive habitats as relatively uniform populations, recognizable in the field. Further study of this problem would most likely clarify the relationship of these plants to Marshallia tenuifolia proper as ecotypic. The best recourse, with the information available at the present time, seems to be that of grouping all the variation together as a single species. It is interesting to note that, for certain years, herbarium re- cords indicate two flowering periods, annually, for the popula- tions of Marshallia growing in the moist flatwoods of Highlands Hammock State Park, Highlands County, Florida. The flowering of these plants around the first of January as well as around the first of August was verified by the park naturalist, Miss Carol Beck. The significance of this is not known. Persisting over the colder months as rosettes, however, it would not be surprising, especially after a relatively warm winter, for such specimens to respond to favorable conditions by flowering. The ability of cer- tain populations of this species to flower at a time other than in the fall is probably restricted to those populations of the warmer parts of Florida. It is mentioned here because of its bearing upon the problem of a possible means of contact with M. ramosa, a spring-flowering species of Georgia, with which it is believed to have hybridized. If vernal flowering of M. tenuifolia ever occurs as far north as south-central Georgia, where M. ramosa grows, it 130 R. B. CHANNELL is probably later than the first of January and may thus coincide with the flowering of M. ramosa. Putative hybrids of M. ramosa and M. tenuifolia are discussed under the former species. DovuBTFUL AND EXCLUDED SPECIES Marshallia Lagunae (Scop.) Gmel. Syst. Nat. 2: 836. 1791.=Homal- ium Jacq. See (Kew) Bulletin Misc. Information, Nos. 6, 7, 8 & 9, page 432, 1935. Marshallia bulbosa Raf. F|. Ludov. 61. 1817=Sarrette bulbeuse Robin, Vog. Louis. 3: 430. 1807. Collected in Louisiana. De Candolle (Prod. 5: 680. 1836) equates the Rafinesque name with Liatris (spec.?) which appears correct except for the scaly pappus which is unknown in that genus. Definitely not Marshallia. Therolepta pumila Raf. Neogenyton 3: 1825. Collected in Kentucky by Rafinesque. The only clue to the identity of this binomial is that given by Rafinesque (New Fl. N. Amer. 4: 77. 1836) himself, to the effect that it may be Marshallia pumila (Raf.) Raf. Marshallia aliena Spreng. Syst. Veg. 3: 446. 1826. Collected in Monte Video by Sello. De Candolle (Prod. 5: 680. 1836) equates the name with Heterothalamus brunioides Less. Marshallia australis Spreng.! in h. Bald. 1822 sed in ejus oper. desid. Valde affinis Calea prunifoliae. (v.s.) This reference is included under Calea Berteriana DC. (Prod. 5: 680. 1836). Definitely not Marshallia. Marshallia dentata Bertol. (Misc. Bot. 15) Mem. Accad. Sc. Instit. Bologna 5: 422, tab. 21. 1854 (collected in Alabama) =rayless plants of Gaillardia aestivalis (Walt.) H. Rock. GENERAL REFERENCES BEADLE, C. D. anv F. E. Boynton. 1901. A revision of the species of Marshallia. Biltmore Botanical Studies 1: 3-10. De CaNbOoLLe, AuGustTrIn P. 1836. Marshallia in Prodromus systematis naturalis regni vegetabilis 5: 680. Paris. Gray, Asa. 1884. Marshallia in Synoptical Flora of North America 1: 303. New York. MALL, J. K. 1901. Marshallia in Flora of the southeastern United States. 1283-1285. New York. aa ——. 1933. Marshallia in Manual of the southeastern Flora. 1455-1457. New York. TorREY, JOHN AND AsA Gray. 1842. Marshallia in Flora of North America 2: 390, 391. New York. S 131 INDEX Athanasia 44, 46 graminifolia 44, 48, 49, 112, 116, 118 obovata 44, 46, 47, 48, 83, 87, 88, 89, 90, 94 trinervia 44, 46, 47, 48, 68, 69, 12. 74 Balduina 59, 60 Bebbia 60 Blepharipappus 59, 60 9 Erigeron ae aha 74 Eupatorieae Flacourtiace na Gaillardia ks 130 5 ae 59 Stata “on wigs 130 Homalium 65, crossing diagr cytological observations 19 densipila x Lescurii 8 densipila x ak 23 discussion 3 distribution map 6 experimental hybrids 14 general observations 26 graphs 10, 12, 13, 15 hy brid index 8, 9 introgression, evidence of 22 literature cited 40 meiotic chromosomes 21 pollen quality 19, 20, 21 seed quality 16, 17 seeds and pollen 16 species question 28 summary and conclusions 38 wild hybrid io ears 7 wild hybrids 1 wild species pioneer 5 Liatris 117, 130 Marshallia 64 aliena 130 egies 47, 112, 123, 124, var. cyananthera 48, 120, 123, 124, 125 australis 130 bulbosa 130 mate 51, 54; 56, 59; 62, , 68, 97, 99, 100, 104, 105, ri 110, 111 var. caespitosa 50, 55, 58, 62, 63, 68, 100, 102, 108, tO. Jit, 12 var. signata 55, 58, 62, 63, 64, 68, 90, 97, 99, 105, 106, 108, 109, 110, 111, 112 oo ee 123, 124 dentata 1 ae 49, 51, 52, 53, 54, 55, 58, 59, 62, 63, 68, 99, 100, 112, 114, 117, 118, 119, 120, 123, 124, 125 var. lacinarioides 112 var. cyananthera 48 grandiflora 50, 52, 53, 54, 55, 56, 57, 58, 61, 62, 63, 67, 73, 74, 76, 78, 79, 82, 83, 89, 90, 5 lacinarioides 48, 49, 112, 117, 132 Lagunae 130 lanceolata 45, 47, 65, 73, 83, 88, 89, 10! var. aes gy feo 48, 83, 86, 87, 89, latifolia i a 68, 69 Mohrii 50, 52. 54, 55, 57, 58, 61, 62, 63, 64, 67, 79, 80, 82, &3, 11] obovata 45, 51, 53, 54, 56, 57, 59, 61, 62, 64, 67, 73, 83, 88, 89, 94, 95, 108, 111, 119 var. obovata 50, 53, 55, ! re) | oo Se 8 o> & [o>] oo (onli galt De os ~] H G0 § Oo Ul NS 89, 90, 94, 95, 111 vat. sinkypbylis 48, 78, 82, 83, 88, 90, 111 var. scaposa 50, 55, 56, 58, 62, 63, 68, 83, 90, 92, 94, 95 pumila 130 ramosa 50, 52.°53, 64, 35, 36, 58, 59, 62, 63, 64, 68, 96, 97, 98, 99, 100, 129, 130 Schreberi 46, 69, 73 spiralis 100, 105 tenuifolia 52, 53, 54, 55, 56, 59, : , 64, 68, 99, 100, 17, 18,1200, 123, 124. 195, 126, 127, 128; 130 INDEX trinervia 50, 51, 69, 70, 72, 75, 14: loree Williamsonii 112, 118 Mutisieae 6 Pebecsisin 46, 47, 65 angustifolia 46, 47, 112, 116, 117, 118 reboot 46, 65, 83, 87 i 13 angustifolia 112, 116 lanceolata 65, Sarrette ewe 130 Therolepta 47, 48 umila 47, 130 Trattenikia 46, 47, 65 angustifolia 112 lanceolata 65, 83 latifolia 69 Tridax 59 Verbesineae 59, 60 Mains Xyris 1 52, 54, 55, 56, 57, 58, 59, 61, 62, 63, 67, 68 CONTRIBUTIONS FROM THE GRAY HERBARIUM — OF HARVARD UNIVERSITY Edited by ; ' Po Reed C. Rollins and Robert C. Foster gk R “ ¥ ‘THE GRAY HERBARIUM O CAMB F HAR RIDGE, MASS, USA. CONTRIBUTIONS FROM THE GRAY HERBARIUM OF HARVARD UNIVERSITY Edited by Reed C. Rollins and Robert C. Foster NO. CLXXXII 3? THE GENUS PEDILANTHUS (EUPHORBIACEAE) BY RoBeRT L. DRESSLER Published by THE GRAY HERBARIUM OF HARVARD UN IVERSITY CAMBRIDGE, MASS., U.S.A. 1957 Issued July 26th PART I. GENERAL INTRODUCTION Pedilanthus, a Mexican and circum-Caribbean genus of Euphorbiaceae-Euphorbieae, has long attracted the atten- tion of botanists because of its strangely formed inflores- cence, and the curious, succulent plants are often cultivated in greenhouses and tropical gardens. Nevertheless, the genus is botanically poorly known; identification by the use of herbarium material or published keys has been nearly impossible, and most of the morphological studies have been both superficial and obscure. The present study was initiated at the University of Southern California, after brief field acquaintance with the genus in western Mexico, and con- tinued at the Gray Herbarium, with short periods of field work in other areas of Mexico. The taxonomy of the genus is herein revised, studies of several aspects of the morphology are given, and the probable evolution and relationships of the genus are treated. The tribe Euphorbieae, of which this genus is a member, is characterized by its inflorescence, the ‘“cyathium.” This inflorescence, with its central (terminal) pistillate flower, surrounded by clusters of small staminate flowers, which, in turn, are subtended by a calyx-like whorl of bracts, is in many cases one of the most flower-like “pseudanthia” known. The tribe is especially interesting to students of phylogeny. It is very well defined, and the reduction and modification of the inflorescence seem to provide evidence of clear evolutionary trends. There is rarely need to resort to hypoth- esis or the fossil record for unsatisfactory “missing links’’ of a major sort. This is emphasized by the fact that the genus Euphorbia, which includes over a thousand diverse Species, has so long resisted rational “‘splitting’’ on a world- wide basis. Pedilanthus is distinctive, within this tribe, by the possession of a zygomorphic, spurred involucre. Several other Euphorbieae (Cubanthus, Monadenium, Euphorbia subgenus Poinsettia) have developed some degree of zy go- morphy, but only in Pedilanthus are the glands concealed in a spur. While it is hoped that the taxonomy of the genus has been clarified by the present study, and although certain morpho- 4 ROBERT L. DRESSLER logical features are treated here in some detail, Pedilanthus still deserves careful study in several aspects. We have, within a single, relatively small genus, a series of species with widely varying degrees of xeric adaptation. This genus would be appropriate for an intensive comparative study of anatomical and physiological adaptation to xeric conditions. The cytology of Pedilanthus is very inadequately treated in the present work. Slow-growing perennials are poor material for cytogenetic studies, but the group is one which appears to be undergoing rapid, radiative evolution, and may warrant careful cytological study for this reason. A careful field study of variation in P. macrocarpus should be rewarding, and the Antillean taxa need extensive field and laboratory study. The natural history of this interesting group, and especially the pollinating agents (presumably humming- birds), remains essentially unknown. ACKNOWLEDGEMENTS The author is deeply indebted to the many persons whose aid has made the present work possible. Dr. Reed C. Rollins has kindly guided and advised the author through the greater part of the research, and generously gave of his time in personally searching for critical material in European herbaria. Dr. Louis C. Wheeler first sparked the author's interest in the Euphorbieae (and, indeed, in botany) ; his nomenclatorial work, published and manuscript, has contrib- uted much to the present study. Professor I. W. Bailey has greatly aided in the morphological studies by his interest and suggestions. Professor Ernst Mayr and Dr. Ernest Williams have read the manuscript and given valuable sug- gestions. Dr. Sherwin Carlquist has shown a sincere interest in the work, and his wide knowledge of techniques for anatomical and cytological study has been an invaluable source of help. Dr. Grady L. Webster, too, has taken an en- couraging interest nearly throughout the present study. Living plants and other special materials have been con- tributed by a number of persons, including Dr. Lucas Calpouzos, Mr. Ladislaus Cutak, Dr. Charles DuQuesnay, Dr. Howard S. Gentry, Dr. Richard A. Howard, Mrs. Ruth Oberg, Dr. George R. Proctor and Dr. Grady L. Webster. Valuable photographs have been contributed by Dr. J. W. THE GENUS PEDILANTHUS (EUPHORBIACEAE) 5 Carmichael, Dr. Carl Partanen and Mrs. Anstis H. Wagner. Dr. Robert E. McLaughlin and Mr. Donald R. Whitehead have kindly prepared a number of pollen samples by aceto- lysis. The diagnostic drawings of involucres in Plates XVI to XXI were prepared by Miss Joan Ericksen; her pa- tience and skill are deeply appreciated. Dr. Ding Hou helped greatly in preparing the habit sketches of the new species. Most of the distribution maps are based on Goode’s Series of Base Maps, which are used by permission of the Uni- versity of Chicago Press. The outline map on which Map 6 is based was made available through the kindness of Dr. Ira L. Wiggins. Drs. W. L. Brown, Jr., and E. O. Wilson have identified some of the ants which were found on plants of Pedilanthus, and the thrips were identified by Dr. Lewis J. Stannard. Much of the field work was made possible by grants from the Fernald Fund, for which I am indebted to Mr. F. W. Hunnewell. National Science Foundation pre- doctoral grants have given generous support during the later portion of this study. The author is indebted to the curators who have kindly lent material for study, as well as to those who have searched their herbaria for critical specimens. Material from the following herbaria has been studied: Arnold Arboretum (A) Allan Hancock Foundation, University of Southern California University of er ce (ARIZ) Botanical Mus and nie eee Copenhagen (C) Chicago pein ans Museu Conservatoire et Jardin fictions Genéve (G) ray Herbarium (GH) Botanische Anstalten der Martin Liither Universitat, Halle (HAL) oyal Botanic Gardens, Kew ( 9 dan Staatssammlung, gag oe (M) Instituto de Biologia, Mexico (MEXU) ; ) ew York Botanical Garden (NY) Muséum National d’Histoire Naturelle, Paris (P) Academy of Natural Beene Philadelphia (PH) Naturhistoriska Pia eum, Stockholm ( Botanical Museu and Hebetiee, Utrecht (U) Naturhistorisches Museum, Wien ) 6 ROBERT L. DRESSLER HISTORY Pedilanthus was known to the pre-Columbian botanists of Mexico, and used there as a medicinal plant. A species, ap- parently P. tithymaloides, is described by Francisco Her- nandez (1651) as chapolxochitl, or grasshopper flower, and its reputed medicinal uses and native synonymy are given. Cuttings of P. tithymaloides were early carried to Europe, and the plant was known to several pre-Linnaean writers. Linnaeus described this plant as Euphorbia tithymaloides in his Species Plantarum (1753), and two varieties were as- signed to the species. Miller, in 1754, and Gomez Ortega, in 1783, considered this plant and its varieties to form a distinct genus, for which they took up pre-Linnaean names. The first, Tithymalus, is a homonym of the contemporaneous Tithymalus of Trew, used for a different segregate of the Linnaean Euphorbia; and the second, Tithymaloides, has remained in nearly total disuse. The now well-known name, Pedilanthus, was proposed by Necker (1790) and taken up by Poiteau (1812), who recognized three species (all here treated as subspecies of P. tithymaloides). Necker’s generic name became a nomen genericum conservandum at the most recent International Botanical Congress (1954). All of the earlier “species” which were described are very similar to Euphorbia tithymaloides L. in the inflorescence, and they are here considered to be conspecific with that taxon. The first clearly distinct species was described by Jac- quin as Euphorbia bracteata (1798). Klotszch and Garcke, in their hasty and superficial revision of the Euphorbieae (1859), recognized three quite unnatural genera within Pedilanthus, and named several supposedly new species, with inadequate descriptions. Boissier revised the group for DeCandolle’s “Prodromus” (1866), but accepted most earlier species and descriptions uncritically. At this time he created the subgenus Cubanthus, for P. linearifolius, described by Grisebach in 1860. Millspaugh (1913) presented a “re- vision” of Pedilanthus, with a number of putative new species, and raised Cubanthus to generic rank. The change in status for this group proves to be correct, but his paper otherwise contains numerous errors and is based on an entirely uncritical species concept. In all, about 57 epithets have been ascribed to the genus, and only a few are referable THE GENUS PEDILANTHUS (EUPHORBIACEAE) 7 to other genera (three to Cubanthus, two to Euphorbia). Most of these species have been described without the study of previous type specimens, and the descriptions are often diagnostic only for the striking generic characters. It is largely because of this sort of confusion that Pedilanthus has been known as a “difficult” genus. ECONOMIC VALUE Aside from widespread cultivation as a somewhat orna- mental and indestructible hedge plants in tropical and sub-tropical gardens, Pedilanthus is of little economic impor- tance. Nearly everywhere, the latex is reputed to have dras- tically purgative action, and other medicinal qualities are sometimes ascribed to the plants, but their medicinal uses are now limited to folk-medicine. The stems of Pedilanthus, especially of P. cymbiferus, bear a wax similar to that of Euphorbia antisyphilitica, and the latices appear to have some rubber content, but neither of these has been exploited commercially. References to the commercial extraction of wax from ‘“‘Pedilanthus Pavonis” stem from an early writer’s confusion between Pedilanthus and the above-mentioned Euphorbia (both being called candelilla). Pedilanthus tithymaloides is widely cultivated in tropical areas. It is readily propagated by cuttings, drought-resistant, and relatively unpalatable to animals, and the stem and leaves remain green and presentable, if not attractive, even after considerable mistreatment. During the winter months, the leaves are often shed, but the bright red involucres are then borne in abundance. This species and P. macrocarpus are frequently found in greenhouse culture in temperate latitudes. Other species which might prove useful as green- house or garden plants are P. Palmeri, P. Finkii, P. pulchel- lus, P. cymbiferus and P. bracteatus (especially the form from Iguala, Guerrero). Common names are listed for each taxon, but a few of the names are, as with a majority of common names, generic rather than specific, and merit brief comment here. Ditamo real, from some fancied resemblance to the European Dic- tamnus, is widely used in Caribbean America, usually for the different forms of P. tithymaloides. Candelilla is applied to various Mexican species (especially the succulent ones) 8 ROBERT L. DRESSLER as well as to Euphorbia. The colorful and oddly formed involucres of the genus may be compared to birds or shoes and have resulted in numerous names such as pie de nino (child’s foot), zapato del diablo (devil’s shoe), pericos (par- rots), gallitos (little chickens), redbird cactus, slipper spurge, etc. PART II. MORPHOLOGY MATERIALS AND METHODS The specimens used in the morphological studies have been largely from the author’s own collections, either collected in the field, or taken from plants grown in the greenhouse. In a few cases, samples (especially of stems) have been removed from herbarium specimens. Table I lists the collections used (materials used in pollen and chromosome studies are listed separately). Materials were preserved in Formalin-Acetic acid-Alcohol (specimens from herbarium collections were softened in hot water and stored in FAA, with a little glycerine added). Samples of roots, stems, leaves and inflorescence parts were imbedded in paraffin, sectioned with a rotary microtome and stained with safranin and fast green (when not otherwise specified, the techniques are those given by Johansen, 1940). Older stems and wood samples were cut with a sliding microtome, without imbedding, and similarly stained. Wood macerations were prepared using Jeffrey’s method. Involucres and other parts were cleared, for study of the vascularization, in 85% lactic acid. The parts used were generally de-colored in hot 6% sodium hydroxide (in a paraffin oven) until translucent, washed in water, and then heated in lactic acid for ten minutes in a boiling water bath. Very small structures clear adequately without de-coloring. Materials containing tannins and similar substances do not de-color well with sodium hydroxide. Too long a period in sodium hydroxide results in very clear material, but this is too fragile for dissection. When stored in lactic acid, the samples become clearer and tougher with time, but only very slowly. Parts cleared by this technique may be con- veniently mounted in Hoyer’s medium (Beeks, 1955). Since pollen cannot be stored indefinitely in lactic acid mounts, a modification of the lactic acid treatment was developed ; THE GENUS PEDILANTHUS (EUPHORBIACEAE) 9 TABLE 1. Materials used in morphological studies. When an her- barium specimen is not now available to document the study, the locality of origin is given for the collection. Data on the remaining collections will be found in the specimen oe in Part IV. The initial “D” refers to the author’s collectio PART STUDIED ce re) © = SPECIES AND COLLECTION 4 = = SB See Oe 3 2 o > om WM 2) a =| 0 P. Palmeri, D 1064 x ee x Palmer 1995 PS « P. Finkii, Thompson, s.n. x x P. calearatus, D 1067 = ae 6 > WES: See 380, ca. Tuxtla Gutiérrez, Chiapas ae oe D 1766 oe P. gracilis, Hinton 10973 x P. pulchellus, Conzatti, Reko and Makrinius 3106 x x P. coalcomanensis, Hinton 12685 x x P. cymbiferus, D 13 Sx x P. macrocarpus, D 1090, northern Sinaloa eee tee: Gentry 7677a x x P. bracteatus, D 979 x 2 athe D 1803 » ee x P. tomentellus, D 1375, Oaxaca, Oaxaca Ss oe P. tehuacanus, D 1795, Tehuacan, Puebla x x P. nodiflorus, D 1362 x D 18 x x x x P. tithymaloides ssp. tithymaloides, D 1376, Cuicatlan, Oaxaca ae Abie Ei x 1252, cult., Cuba ee Se ssp. parasiticus, D 1364 x ssp. angustifolius, Cutak, s.n Maricao Mts., P. R., cult at M B.G x Ssp. jgamaicensis, R. A. ae Four Mile Wood, Ja x x DuQuesnay, Negril, Janiiick pelts this is given in full in the section on pollen morphology. Pollen samples were also prepared by acetolysis (Erdtmann, 1952). All chromosome counts were obtained by using smears of young staminate flowers. Material collected late in the after- 10 ROBERT L. DRESSLER noon of a sunny day generally proved most satisfactory. The involucres were opened and removed from the flowers under a dissecting microscope and the flowers placed in ferric propionate fixative (Hyde and Gardella, 1953) for about 24 hours. The anthers were then crushed and heated in aceto-carmine. These slides were later made permanent, without removing the cover slips, by washing out the aceto- carmine with 40% acetic acid (applying the acid to one side of the cover slip and blotting up the stain on the other), leaving the slides overnight in a moist chamber with absolute alchohol, and then ringing with Diaphane. The slides may be counterstained by adding a drop of Fast Green (0.1% in 40% acetic acid) when removing the aceto-carmine. VEGETATIVE MORPHOLOGY Habit. — The range in habit found within Pedilanthus is unusual in so small a genus. Pedilanthus calcaratus and P. coaleomanensis are usually small trees, with distinct trunk and extensive wood formation, sometimes reaching seven meters in height. Both are deciduous and bear large leaves (some ecotypes of P. calcaratus are only facultatively deciduous and may retain their leaves through the winter in moist sites). Pedilanthus Palmeri is distinctly woody and often reaches two meters in height, but branches extensively from the base. Pedilanthus bracteatus and P. tithymaloides and their immediate allies are distinctly shrubby, generally ranging from one to two meters in height and branching from the base. These plants often have a roughly obconic form when undisturbed. Pedilanthus cymbiferus and P. macrocarpus are clump-forming desert shrubs with no trace of a trunk or main stem (see Plate XVI). The former pro- duces numerous rush-like stems from the ground and forms modified underground stems which serve to spread the colo- nies over short distances. Pedilanthus macrocarpus is larger and markedly succulent; a habit resembling that of P. cym- biferus is achieved by the formation of adventitious buds on the roots, giving rise to extensive clonal colonies. Pedilan- thus Finkwi, P. gracilis, P. pulchellus and P. tehuacanus are all species for which exact information concerning the habit is unavailable; the first is apparently less woody than P. Palmeri (its closest ally), the second and third are woody THE GENUS PEDILANTHUS (EUPHORBIACEAE) 11 and perhaps small trees, and P. tehuacanus appears to be a succulent plant similar to P. bracteatus or of smaller stature. Although P. cymbiferus is a desert plant, it is scarcely more succulent than E’phedra, some species of which it re- sembles in growth form. Some populations of P. calcaratus have rather thick twigs, but true succulence is limited to P. bracteatus and P. tithymaloides and their allies (spp. 8-13). These species are predominantly stem succulents. The leaves of P. macrocarpus and P. nodiflorus may be some- what fleshy, but they are small and soon shed. Pedilanthus tithymaloides subsp. parasiticus, on the other hand, includes some populations with quite fleshy leaves that are retained for some time. Roots. — The roots of Pedilanthus are generally woody and much branched. In P. calcaratus, and possibly in some other species, the primary root and later some secondary roots form characteristic thickened storage organs. In these roots the pattern of the primary xylem is completely ob- scured, and the greater portion of the xylem consists of large, starch-bearing parenchyma cells, with irregularly scattered vessels, usually in small groups. The parenchyma cells near the vessel groups are smaller than the ground parenchyma cells and may occasionally be lignified or be accompanied by a few gelatinous fibers. The storage xylem of these roots increases in girth by. cambial activity, and the cells in the outer portions are in more clearly radial alignment. In one storage root of Dressler 1380, the outer- most mature xylem forms a lignified band 0.2 - 0.5 mm. thick. In this band some fibers have weakly lignified gelatinous secondary walls, but the majority are septate, without gelatinous walls, and include starch grains. The longer parenchyma cells are also occasionally septate, making the distinction between fiber and parenchyma difficult. The septae often remain non-lignified, but it is apparent that these late-formed septae may become lignified, and often tend to be formed at the same level in adjacent cells; the series of cells from a single cambial initial thus simulating, at times, series of cells from separate initials. Storage roots similar to those here described are found in Cubanthus Brittonii (in this species the parenchyma has a much more regular geometrical arrangement, but the 12 ROBERT L. DRESSLER structure is otherwise very similar) and in Euphorbia radians (Reiche, 1923). Structures which may be similar are found in many other species of Euphorbia, Manihot, Cnidoscolus, and Jatropha, and in seedlings of Hevea and Joannesia. Older roots form a homogeneous cork of thin walled cells. The root cortex consists of relatively large, isodiametric parenchyma cells, which often contain starch grains, and frequent laticifers (very young roots have not been ex- amined). In the cortex of P. macrocarpus, there are found, in addition to starch grains, numerous spheroid bodies of varying size, which stain dull red with safranin. In the cortex of the storage roots of P. calcaratus there may be occasional gelatinous fibers. In large roots, and especially in storage roots, the outer cortical parenchyma includes tangential series of cells within the outlines of their mother cells, indicating that series of anticlinal divisions have taken place in the growth of the root. The primary vascular system varies from triarch to pen- tarch, with a conspicuous pith remaining in the center. With secondary growth, broad, large-celled rays (1-)2-5 cells wide (wider near branch roots) are formed at the protoxylem points, and weakly lignified wood develops be- tween these. This xylem contains numerous rays which are homogeneous (upright) and largely uniseriate; two- and three-seriate rays also occur, especially in older roots. The rays may be somewhat lignified. The vessels, which are In scattered radial groups, are scalariformly or occasionally (in P. calearatus, often) alternately pitted. Wood paren- chyma is mostly paratracheal. There are numerous fibers, which in smaller roots are scarcely or not at all lignified and have a markedly gelatinous secondary wall. In P. macro- carpus and P. nodiflorus the gelatinous fibers are very thick walled and stand out strongly in cross section; in the first species they are found in the primary body as well as in the secondary xylem. In P. tithymaloides and P. calcaratus the fibers are usually (in the material examined) thinner walled and less conspicuous, but still markedly gelatinous. Narrow bands of well-lignified wood may be found in the central portion of the root, and, in larger roots, the outer portion is largely well-lignified. The fibers in such areas have thinner walls and may occasionally be septate. THE GENUS PEDILANTHUS (EUPHORBIACEAE) 13 Stems. — The stems of Pedilanthus show an interesting range of variation. The stems of the arboreous species are woody and may produce extensive xylem, the trunk of P. calearatus, for example, reaching 6-8 cm. in diameter. In the nearly leafless extreme xerophytes, such as P. macrocar- pus and P. cymbiferus, the stem may be thick and fleshy or thin and woody, but has very limited secondary growth. In intermediate types, such as P. bracteatus, there is no distinct trunk, but old stems develop a good deal of secondary xylem and may reach 2-3 cm. in diameter at the base. The stem epidermis is moderately to heavily cutinized, with the cutin extending between the epidermal cells for about half their thickness. In the markedly xerophytic species, and especially in P. cymbiferus, there is a conspic- uous wax on the older stems. This wax, like that of Euphor- bia antisyphilitica, may be removed by boiling water. The stem of P. cymbiferus is irregularly pitted, with the tri- chomes occurring on the ridges and tubercles, and the stomata in the pits. A tendency toward this pattern is seen in other more or less xerophytic types, but it is much less marked. The stomata are more or less flush with the stem surface in P. pulchellus and a collection of P. calearatus from a relatively moist habitat (Dressler 1380), but they are slightly to markedly sunken below the surface in the other plants examined, including a more xerophytic form of P. calearatus (Dressler 1067). The cutin is thin over the stomatal guard and accessory cells, but in some cases an especially thick cutin surrounds these (see fig. 9). The basal cell of each trichome is heavily cutinized on the basal and lateral walls. The stomata and trichomes wil] be described in greater detail under leaf structure. When much secondary growth occurs, a smooth, grey, homogeneous bark of thin-walled cells is formed. Bark for- mation takes place immediately beneath the epidermis. On P. calcaratus, at least, small, reddish lenticels are formed. The cortex consists of relatively spongy parenchyma and numerous branching laticifers. No cortical collenchyma has been observed, though this is reported for P. tithymaloides by Assailly (1954). Chloroplasts are found in the cortex of all species examined (not certain for P. pulchellus and P. coalcomanensis, where only herbarium material was avail- able). In the large-leaved types only the outer one half to 14 ROBERT L. DRESSLER cortical cells of P. cymbiferus bear chloroplasts. In these species the outer half of the cortical parenchyma has a dis- tinctly “palisade” arrangement. The same arrangement is present in P. bracteatus, P. tithymaloides ssp. angustifolius and P. t. ssp. jamaicensis, and (but less marked) in P. tithy- maloides ssp. parasiticus. Substomatal chambers are usually distinct in the stems of Pedilanthus, so that the stomata are macroscopically visible as pale flecks. In P. cymbiferus and forms of P. macrocarpus (Dressler 1090) the substomatal chambers are quite large. In another collection of P. macro- carpus (Gentry 7677a), however, the substomatal chambers are relatively small, and some of them are actually cutinized within. Starch grains are frequent in the cortical paren- chyma, and small spheroidal bodies which stain dark red with safranin are frequent, especially in the chlorenchyma. The laticifers are best developed in the cortex. Several to many laticifers are found in the pith, and laticifers oc- casionally extend through the secondary wood by way of small multiseriate rays. The most conspicuous concentra- tion of laticifers is found in the inner cortex, where they are mostly longitudinal and of large diameter. These large laticifers are often thick-walled, and on one specimen of P. tithymaloides ssp. tithymaloides the walls of some laticifers seemed to be weakly lignified. In this same species (Dressler 1376), afew markedly constricted laticifers were noted, per- haps supporting Milanez, suggestion that the ‘“nonarticula- ted” laticifers of the Euphorbieae actually develop by the fusion of distinct cells (1952). Laticifers of medium size are frequently seen to extend across the cortex more or less radially, and a network of very tiny laticifers is found im- mediately beneath the epidermis. These subepidermal lati- cifers, which are so small as to pass unnoticed or to be mistaken for intercellular spaces under low magnification, account for the ready “bleeding” of the stems at the slightest scratch. The laticifers are multinucleate (but the nuclei are difficult to distinguish except under very favorable circum- stances) and contain numerous very elongate starch grains (fig. 5). Short or isodiametric starch grains are infrequent in the laticifers. THE GENUS PEDILANTHUS (EUPHORBIACEAE) 15 The primary vascular tissues of the stem are eustelic, with numerous protoxylem points surrounding the pith, and a similar number of groups of non-lignified primary phloem fibers. These phloem fibers are well developed in the woody species, including P. cymbiferus, moderately developed in P. bracteatus, P. tomentellus and P. nodiflorus, moderately to poorly developed in P. tithymaloides, and very poorly de- veloped in P. macroca The pith is largely composed of relatively isodiametric parenchyma cells, which often include starch grains. In P calcaratus, one or two tubes which appear to be laticifers are found in the outer pith and are distinctive in being sur- rounded by a layer of small non-lignified cells, and a partial layer of two to three rows of lignified or partly lignified cells ; these latter are partly gelatinous fibers, in which only the primary wall is lignified (see fig. 8). Similar structures are found in the pith of P. coalcomanensis and P. pulchellus, but have been seen in no other species. The pith cells of P. coalcomanensis each contain a single dark-centered, many- pointed druse, and smaller, more irregular crystals are found in the pith of P. Finkii. Pedilanthus cymbiferus is interest- ing for the occurrence of lignified parenchyma cells in the outer pith, touching and near the protoxylem. Pedilanthus cymbiferus spreads locally by thick under- ground stems. The aerial stems of this species are 2.5-3.5 mm. in diameter, with strongly lignified xylem. The under- ground stems are about 5 mm. in diameter, produce cork externally, and have abundantly starch-bearing parenchyma in the cortex and pith (without chloroplasts, palisade ar- rangement, or lignified pith cells). The phloem fibers are well developed, as in the aerial stem. The secondary xylem of the “stolon” is weakly lignified, and, except for an incom- plete (in the material seen) band of lignification, the fibers are moderately thin-walled and gelatinous. Some of the ray parenchyma is lignified in the band of lignification, and some of the fibers are septate. Wood Structure. — The secondary xylem of Pedilanthus has been relatively well sampled, and the major trends of variation can be indicated, but better sampling of each spe- cies is needed for a detailed treatment. No growth rings are present. The pores are commonly aggregated into radial multiples. This trend is much less ROBERT L. DRESSLER 16 x xe x x o A EY Ay) aR are Z9S AeusanOnc ‘sisuaovwwoyl ‘dss xx x x xX @ oS: Gor = Set T68 9LET xx x x x @ bo SLY '- Fee 9SP SSSI “Cd ‘Saproypwhyyy “dss saproppuhiyiy *d x xz ase: 62OV: GS: Gor SBt 80s. gas 608T Cd ‘snsoyipou *q xx @ xexx 08S OF 802 999 0°62 G6LT “CT “SnunDononya} *d xx % xx @ ve Glo 696 v'vS GLET C ‘snpjazuamo? “qd xx % aw SCG Of oto see S&P GO8T Cd ‘snzVa}0D4q “qd xx Kx S00F VS S8r GL Tré L9E 060T C ‘srdimo0u9nM *q x xX xX x xX 6rss OC LO MBL OLB FOE 89ST Cd ‘snwafiquihia “gq (x)xX x x x xX OL6S Fh OPT TI9 9E% 6ZE9OTE SNIULLYRW WY oxy ‘WQezZU0D ‘snpjayoynd *q (xX)xX x x xuux VISE. SP SL “ss O'eS Ss "PS G89ZT UOJUIH{ ‘szsuaupwos)p09 *q Xx @XX X ®% X(xX) L’¥SP TLP SL60T UOT ‘syn *q O'T9P L'T9 99LT A xO XX X & x(x) Sy ro gZ "69 LOOT a OFS - #8" BRL util eee ee yuna 19yno ‘OgET Cd Ge tl - er 3 9P yuna} teuul “oggt d x@xXxX XxX 2% x(x) oy. 2) L'2S 9°6F SIM4 ‘OSET C ‘snzpLpI7D9 * J XUKE XBEX 66cr TS 6P COG = =L'8Z ‘u's uosduoyL, “yur “d xUxXxX %XUUX GOrS: OF OVE Ge 0°69 G66T Teed “Wwaupngd ‘d Sr eos beg s % Sc-. if “S838 ee Saran sf Foe > Ee a3 5a 55 a se age i eee ee ee Se ee ee B 8g B oS oy 8 ® g 8 88 § NOILOATION GNV Saloads ae ee a DNILLIG DNILLIG 6 % = BE o5 5 VWAHONGYVd 9 ‘THSSHA : a Z e —T1assaA —TASSaA ‘yuonbeajur AraA ynq “uaserd—,,(x),, ‘yueurmopeard—,.2,, ‘yuesa1d—,.x « YUNIY Iayno ‘QgeT C,, Ul ydaoxa ‘poom Jo Ta,OUTL] [IU JSOULIBUUT BY} WAT UdYR} 91B SJUNWIAAINSBOUT [TY “SUOTOTU UT oe SUOISUBULIP [[@ :S}UIUIAINSvEW 0Y9-0G JO Sasvr9ae Juasaideat SIeqUINN “snYyZUD/Ipag UI SaInyeay JossoA °Z ATAV], THE GENUS PEDILANTHUS (EUPHORBIACEAE) 17 marked in the P. bracteatus and P. tithymaloides groups and P. Palmeri than in the other species (see Table 2). in #. Finkii and P. pulchellus the vessels tend to be in radial multiples, but the multiples are not so clearly delineated as in the species with larger vessels. The number of vessels per square millimeter is highly variable and, with the vessel diameter, seems to correlate to some degree with the habitat and the relative need for rapid water conduction. This re- lationship is better shown by the vessel diameters. The smallest diameters are found in the most mesophytic species, P. Finkii and P. pulchellus, and in some markedly xerophytic ones, P. cymbiferus and P. tehuacanus. In both the shaded mesophytes and the leafless xerophytes the stress of heavy transpiration, with the resultant need for rapid water trans- port, would be unlikely or infrequent, though for different reasons. The largest vessel diameters are found in those species which are large leaved deciduous trees of the tropical deciduous forest; P. calcaratus and P. coalcomanensis. It is reasonable to assume that these trees are frequently sub- jected to heavy transpiration and that selection for efficient water conduction has been correspondingly effective. It is interesting that Dressler 1067 and Dressler 1766, collections of P. calcaratus from relatively xeric tropical deciduous forest, have very large vessels (av. 61-63 microns) in their inner wood, while Dressler 1380, from a much wetter site, has markedly smaller vessels (av. 46-49 microns). ood from the outer trunk of the last collection has, however, much larger vessels. A similar increase in vessel size with ontogeny would doubtless be found in the other species, if they were adequately sampled (Bailey and Tupper, 1918). The discrepancy between vessel diameter as measured in section and as measured from macerated wood, in this col- lection, is probably due to the presence of elements from one or two exceptionally large vessels in the sample chosen for maceration. One vessel element in that preparation showed an extreme diameter of 162 microns. The moderately succulent xerophytic species, such as P. tithymaloides and P. bracteatus, and the moderately mesophytic types, such as P. Palmeri and P. gracilis, tend to have vessel diameters intermediate between the extremes given above. The pattern of vessel element length is less clear; the relatively primitive, mesophytic P. pulchellus shows the longest vessel elements 18 ROBERT L. DRESSLER _ - . - : n toe Sys atts e. oe! Mati @t:. ? anh, es e a | «ag eect 4:8 te i te 78 Ge @ a: te . at? @. 8 ges g ei? He "% & al aoe? o & Ady a = a : apa heel oe, * am & @ het T. i ae ™, & Fd O6I20 0 6ben, ee r 415% & é eT og ae Gee aes *@ 2 a SP aap 4 7 @ ‘iy % é.3'* g@ 0° A eee4 t ; as, ha Me 3 ‘ 42. @ ees? eh ae é 9e,8i8 * atte . -& a at Atlee ‘ al “tae we te ‘gf *@8e56 UT ig Saas & 7 bee Se ee ee at re = Oe Cd - ** A ot Wepeneses ; Ct / Lael, a tsoas ee? a ee THY 4 as je at ote ie Sq: a Astle, ee é RG 10 ni Fos eabs “pn@a (oor = Peas, | Sig @e,t ts GS elpaneaes & i on aeah Peru 47) ae ea" ‘* te @ oan tyahs ee cht ty Mee ¢ “5 >A 746% & ry) oe Ae Lae tee? Se a —. 7+ a? se. ni “i Shy aat® tS TL} se et ae © Soe a >» os . Ld ae * itz a 9 VIZ. 3 & aGs:" quiets tage . a ve By ee rj - eurt (K,: “ib we past’ 2 < veo .. s eo. * it SMe eho eSaeseeiiiy e 2 & em a bee! LS as €2 ey 8) GP & ‘ne @ > TR r . ss . ° . > PLATE I. Wood Structure. Fig. 1. Pedilanthus t. ssp. tithymaloides, Dressler 1376, cross section of old stem, showing small, scattered vessels, magnifi ion ca. 155 ay ae : : s Fig.-2. P. coalcomanensis, cross section of twig, magnification ea. 155 19 THE GENUS PEDILANTHUS (EUPHORBIACEAE) ‘eee ee tate ere + ae eee SEL etme oo emma AIRS SontA gE S&S ose Craw nem asta attrsesaameecemeen sit il ssler 1376, Dre vides, sp. tithymalc e. Fig. 3. Pedilanthus t. s PLATE II. Wood Structur tangential section of old stem, note the biseriate rays, magnification ca. 155. > P. calearatus, Dr Fig. 4. essler 1067, tangential section of small stem (ca. 1.5 em. diam.) note the rays are uniseriate and scarcely distinguishable from the surrounding cells, magnification ca. 155. 20 ROBERT L. DRESSLER ® * & PLATE III. Stem St ructure. Fie. 5. Pedilanthus calearatus, Dressler 1380, laticifer in longitudinal seetion, note the bone-sh: : alo 450. F haped starch grains with yngate hila, magnifi- eation ca. ig. 6. P. calearatus, Dressler 1380, vessel, from tange gene section, showing sirouker — pits in alternate arrangement, magnification ca. 2 Fig. I vesse wwential section, showing sealariform and some transi- Ca. 200. hig. 8. PB: even nie Jressler 1067, cross * pith, showing sheath of lignifiied a magnification ¢a. 230. Fig. 9. P. calearatus, Dressler 1380, ca. 230. cS PS) = o a cross section of stoma on twig, mag nifice ation THE GENUS PEDILANTHUS (EUPHORBIACEAE) 21 and the extreme xerophyte, P. cymbiferus, the shortest, but the other species depart widely from this seeming tendency. Better sampling is needed of this feature, with careful selection of wood from comparable regions of the stem. Perforation plates are always simple. The end wall inclina- tion is quite variable, but tends to be greater in narrow vessels. The “tails”? projecting beyond the perforation plate are also quite variable in length. Interesting trends are to be found in the pitting of the vessels. The pitting is quite variable within the individual sample; but the range of variation and the dominant type of pitting, so far as has been observed, are characteristic for species, and often for species groups. These ranges and trends, subjectively determined, are shown in Table 2. Though “transitional” customarily refers to the transition between scalariform and opposite, a second transition is shown between opposite and alternate; this should not cause in the vessels, but are not lignified. The imperforate tracheary elements consist largely of cells with only moderately thickened walls and very small pits. In P. pulchellus and P. cymbiferus, these fiber-tracheids 22 ROBERT L. DRESSLER have thicker walls than in the other species, and the pits are distinctly bordered, with narrow, crossed apertures. Crossed apertures are apparently present in some other species, but the borders are somewhat reduced, and the pits are so tiny that it is difficult to distinguish fiber-tracheids and libriform fibers, though both may be technically present. Gelatinous fibers are common and often form wide bands. They have been observed in all species, but are absent from the aerial stems of P. cymbiferus and from some stems of P. macrocarpus. Septate fibers, with thin, non-lignified cross walls, have been observed in P. Palmeri, P. Finkii, P. calcaratus and P. tithymaloides, and may be present in other species. Ray and wood parenchyma cells with such non- lignified cross walls have been observed in the above species and in P. macrocarpus. The rays in most species of Pedilanthus are uniseriate and composed entirely of erect cells. In any species, an occasional multiseriate ray will be found containing a transverse laticifer. In P. bracteatus and P. tithymaloides there is a distinct tendency to multiseriate, slightly heterogeneous rays (see Plate II). In these species the body of the ray is often 2-3 cells in width, and, though the cells are not truly “pro- cumbent,” they are more nearly isodiametric than the erect cells of the “wings.” In P. macrocarpus and P. tomentellus, the rays are predominately uniseriate, but the cells of the middle portion are often shorter than those of the extensions. Both apotracheal and paratracheal wood parenchyma are found in Pedilanthus. Apotracheal parenchyma is diffuse, scanty, and very inconspicuous except in the bands of thick- walled gelatinous fibers. Paratracheal parenchyma is simi- larly quite scanty. In summary, the vessel diameter is rather well correlated with the habitat and the relative need for efticient water transport ; and the moderately succulent groups, such as P. bracteatus and P. tithymaloides, seem to have retained the most primitive features in their wood structure: many solitary vessels, scalariform intervascular pitting and slight- ly heterogeneous rays. Pedilanthus pulchellus, a mesophyte with narrow, very long vessel elements and indistinct radial multiples, may show the most primitive features among the truly woody species. While the evolutionary trends in wood structure are well established and, in major features, ir- THE GENUS PEDILANTHUS (EUPHORBIACEAE) 23 reversible (Bailey, 1944), it is possible that these trends are reversible in finer details; that is, within the framework of ontogenetic variability. Leaf Structure. — The nodal anatomy of Pedilanthus is simply trilacunar, the three traces remaining distinct through the petiole and for some distance in the midrib. A small amount of collenchyma (of the “lacunar pattern,” with intercellular spaces) is sometimes found in the petiole and midrib, especially if these are keeled. The stipules, which are borne on the stem, adjacent to the petiole base, vary from rudimentary or absent, as in P. macrocarpus, to nearly a millimeter in length, as in P. tomentellus and some forms of P. tithymaloides. They may be rounded and hemis- pheric in form, but are more commonly conic or spur-like. In P. tomentellus, which has been sectioned and examined microscopically, the stipules are definitely associated with the petiole base (though not shed with it) and are lacking in vascular tissue. The epidermis of the stipule is of the narrow palisade type, very similar to that of the inflorescence glands. In those species of Pedilanthus with large foliage leaves, the leaves are bifacial, with 2-3 layers of palisade paren- chyma beneath the upper epidermis, occupying 1/4 to 1/3 of the leaf thickness. The remainder of the leaf is made up of spongy parenchyma. In P. macrocarpus, the very small leaves are scarcely thicker than those of the mesophytic species, but there is no clearly defined palisade layer, the entire leaf chlorenchyma being made up of an open spongy tissue, the cells of which are somewhat perpendicular to the epidermis. Only a single type of trichome is to be found in the genus. All trichomes are compound and uniseriate; the basal cell of each is very thick-walled and heavily cutinized throughout. The protoplasts of the trichome cells are thus cut off from the epidermal cells, and they die at an early stage. The second cell of the trichome is also frequently thick-walled, but less so than the basal cell. The stomata of Pedilanthus are of the “Rubiaceous”’ pat- tern, with the subsidiary cells paralleling the guard cells. Stomata are found on both upper and lower leaf surfaces, but are more numerous on the lower surface. They are also numerous on the stems, which, in many species, take over 24 ROBERT L. DRESSLER the photosynthetic role of the plant. The subsidiary cells are commonly deeper than either the guard cells or the surrounding epidermal cells (see fig. 9), and often partially enclose the guard cells above and beneath. The guard cells have both inner and outer ledges well developed. The stomata of the leaves which have been examined are only slightly sunken beneath the leaf surface, but the stem stomata of the more xerophytic species are markedly sunken. In the stem of one collection of P. macrocarpus (Gentry 7677a), a number of stomata were observed in which the substomatal chambers were partially filled by proliferation of the cortical cells, and the reduced chambers heavily cutinized within over nearly or quite the entire surface. Such stomata are apparently non-functional. Markowski (1912) describes rudimentary stomata from the leaf of Pedilanthus (either P. cymbiferus or P. tithy- maloides ssp. parasiticus?). Such non-functional stomata have, in the present study, been found to be frequent on the leaf of P. Palmeri, but very few have been found on the examined material of P. tithymaloides (all grown in the greenhouse). The subsidiary cells of the rudimentary stomata appear to be normal, but the guard cells are very narrow and shrunken in appearance. The opening between the guard cells is greatly reduced or, more frequently, absent. In some cases, two distinct cells cannot be recognized. These peculiar structures would appear to be the result of atrophy at an early stage of development, but more study is needed. Abscission Layers. — The efficient removal of non- functional parts can be of some adaptive value to xerophytes, and mechanisms of abscission are well developed in Pedilan- thus. The leaves are cut off by an abscission layer, very early in the extreme xerophytes, much later in the meso- phytes. If the pistillate flower of a cyathium is not pollinated, the cyathium soon abscisses at the base of the peduncle. If pollination does occur, a ring of corky tissue is formed about the base of the involucre, cutting off the staminate flowers and the involucral bracts, which soon dry, but do not normal- ly fall off. When the cyathia and fruits have all dried, the entire inflorescence is cut off by an abscission layer formed above the uppermost functional axillary bud. Of greater interest is the formation of such supra-axillary abscission layers anywhere in the plant, in the event of injury or un- THE GENUS PEDILANTHUS (EUPHORBIACEAE) 25 favorable conditions. Yellowing and death set in promptly just above the abscission layer, though the terminal portion of an internode may remain alive and green for some time after the onset of abscission. If little secondary growth has occurred, the stem falls off unaided, leaving a clean, corky layer on the living portion. If the stem is woody, the dead portion persists, but is very sharply delimited. The abscis- sion layer is often not horizontal, but slanted or curved from a point above the axillary bud to a point nearly opposite the bud. This suggests that its formation is governed by growth substances formed in the axillary buds. Abscission layers were not observed on stems which still bore leaves. Oc- casionally one may find a moribund stem in which there is an abscission layer above each node and each internode is dying from the base upwards. The abscission of the axillary bud, as reported by Mar- kowski (1912), has not been observed in the present study. It is possible that the bud of the specimen sectioned by Markowski had started growth before collection, and that abscission of the young shoot started after the material was gathered and before it was preserved. REPRODUCTIVE MORPHOLOGY _ Inflorescence. — The complete inflorescence of Pedilanthus is a compound dichasium of flower-like involucres. These compound dichasia may be either terminal or axillary, or both. The basic unit in this inflorescence is a terminal involucre subtended by two bracts and their axillary buds, each of which can develop into a similar unit (see fig. 31). Occasionally (P. coalcomanensis and P. tehuacanus) the inflorescence may, by abortion, be monochasial for several successive nodes. The subtending bracts, which are sessile and clasping, conceal the developing involucre for a greater or lesser period (very briefly in P. macrocarpus and P. nodiflorus; nearly until anthesis in P. coalcomanensis and P. bracteatus). The three vascular traces which supply each bract branch before passing through the abscission layer, so that several to many veins enter the broad base of the bract. As noted by Markowski (1912), the bracts of P. tithy- maloides (and P. nodiflorus) are not quite opposite, but 26 ROBERT L. DRESSLER nearly so, the abaxial angle (ventral, relative to the in- volucre) being a little less than 180 degrees, so that, as the dichotomous, axillary cluster of cyathia increases in size, none of the involucres faces back upon the main stem. This divergence may occur in other species, as well, but it is inconspicuous and not readily determined from herbarium material. In P. bracteatus, P. tomentellus, P. tehuacanus, and apparently in P. coalcomanensis, all species with very large bracts, the bracts are opposite, or, if anything, the lower (abaxial, in axillary dichasia) angle is a bit the larger. In these species, however, each successive pair of bracts is borne in the same plane as the preceding pair, so that the successive dichotomies are in the same plane, and the cyathia are all oriented more or less side by side, facing outward from the main stem (in axillary dichasia). The Involucre of Euphorbia. — Before attempting to dis- cuss the involucre of Pedilanthus, a brief discussion of the in- florescence of the related and better known genus Euphorbia is required. Lamarck first suggested (1788) that the ‘‘flow- er,” with its central pistil, clusters of simple, jointed stamens and subtending “perianth,” might possibly be an inflores- cence, a condensed cluster of many tiny flowers. Since then, many conflicting interpretations have been given for this remarkable structure, the “cyathium,” but there is now gen- eral agreement on most of the main features (a more complete summary of the literature may be found in the works of Schoute, 1937; Haber, 1925; and Schmidt, 1907). The Euphorbia cyathium normally consists of a terminal pistillate flower, which is naked or very nearly so, subtended by five condensed clusters of quite naked, one-anthered staminate flowers. The whole structure is enclosed by a more or less campanulate involucre of five bracts which closely subtend the staminate flower clusters. Each of these bracts shows a normal three-trace foliar vascularization pattern (see Plate IV). From one to five (commonly four or five) nectariferous glands are borne in the sinuses between the involucral lobes. Each gland is a double structure, receiving half of its vascular supply from the lateral trace of the involucral glandular tissue is much more heavily vascularized than the 27 @ PLATE IV. Vascularization of Euphorbia involucre. Fig. 10. —— Tirucalli (from cultivated plant), involucre cleared, split aan involueral brs acts and spread tar 3376, two open, magnification ca. 10. Fig. 11. apecaien tie Xantii, Carter cleared involucral bracts, magnification ca. A. petaloid ae iia of gland; involucral lobe (not evident in E. Xantii) ; i gland 28 ROBERT L. DRESSLER intervening foliar structure, the entire lateral trace of the involucral bract, and commonly a few branches from the median trace, go into the vascular supply of the gland. In Euphorbia, subgenera Agaloma and Chamaesyce (which are apparently related) and in the section Medusea of the subgenus Euphorbia (here, apparently developed quite in- dependently), and in the genus Pedilanthus, the glands are differentiated into a basal, glandular portion and a flattened, distal, non-glandular portion, which is commonly white or red, and, in Euphorbia, is quite petal-like in appearance. These “petaloid appendages” account for the most startling- ly flower-like cyathia, notably those of Euphorbia fulgens and EF. Xantti. In the subgenus Agaloma (and in Cham- aesyce, as well), a common pattern involves the development of only four glands and their appendages, the pistillate flower, after anthesis, recurving through the gap where no gland develops. It is from this slight asymmetry that the highly zygomorphic pattern of the genus Pedilanthus ap- pears to have developed (see fig. 28). The structures commonly referred to as “bracteoles” are found between the clusters of staminate flowers. These are continuous basally, and often abaxially, with the involucre, and may occur as irregular rows of trichome-like structures, as solid partitions separating or surrounding the clusters of staminate flowers, or, commonly, as somewhat intermedi- ate structures; partitions which are laciniate above. Since these bracteoles are usually completely without vascular tissue, their exact morphological nature and relationship to the other parts of the inflorescence are often difficult to determine. It appears that two distinct structures may have been involved in the evolution of the bracteoles. (1) The infolded and connate margins of the involucral bracts. The structure of Anthostema, Dichostemma, and, apparently, of Neoguillauminia, as well as the course of the lateral bract traces in some species of Euphorbia (especially E. pulcher- rima) suggest that the involucral bracts primitively enfolded their respective staminate flower clusters. A portion of the bracteole-partition may, thus, be derived from the involucral bracts. This suggestion is supported by the developmental patterns described by Schmidt (1907). (2) The bracts which accompany the condensed branchings of the staminate flower cluster. The bracteoles are customarily interpreted as rep- THE GENUS PEDILANTHUS (EUPHORBIACEAE) 29 SPUR PARTITION LATERAL iz i GLANDS pp MEDIAL SPUR LOBE ACCESSORY cea INVOLUCRAL - (Opes 0 ue SPUR, INTERNAL MENTUM PEDUNCLE INVOLUCRAL TUBE LATERAL VIEW SPUR LOBES angie INVOLUCRAL LOBES (ACCESSORY) DORSAL, DORSAL VIEW ANTERIOR POSTERIOR VENTRAL TERMINOLOGY 1G. 12. Terminology of Pedilanthus Involucre. On the left, schematic dorsal views of P. cymbiferus involucres, showing parts, the upper showing the interior of the Spur; on the right lateral outlines of P. calearatus. 30 ROBERT L. DRESSLER rele tes flowers but one removed. bee tions: ct., bracteoles; part., partition; lat., lateral; med., median or phi i oh. ak: inv., involucral iinet sp., spur lobe; pist., pistillate; stam., staminate; fl., flower. Fic. 14. Diagrammatic sections through base of cyathium. A. pedun eS showing five larger vascular bundles to bracts and staminate flower clusters and fi (often only three) smaller, inner traces, which s ly the pistillate flower. B. r bundles branch, each giving rise to three bract da ce the staminate fl cluster. C he ree bract traces trace to the staminate flower cluster are distinct The traces to the staminate flower cluster each form two branche da small trace which s li he central flower. E. e s to each staminate flower are shown. F. section begirrsss cyathium of Euphorbia. 1. gland; 2. involueral lobe; 3. bracteole-partition; 4. stamin ae flower cluster; 5. pistillate pedicel. This — section is highly schematic, as little space is actually found in the involucre, ee ¢ the glands and bracteole-partitions are ae actually found at the same level. THE GENUS PEDILANTHUS (EUPHORBIACEAE) 31 resenting such bracts, and part or most of the bracteolar structure is surely of this nature (see the developmental studies by Schmidt, 1907, and Michaelis, 1924). It is prob- able that the bracts subtending the first two branchings of the staminate flower cluster primitively form the greater part of the bracteoles (see the structure of Anthostema, Michaelis 1924; Calycopeplus and Euphorbia Tessmannii, Mansfeld 1929; and Neoguillauwminia, Croizat 1938). The nature of the staminate flower cluster is still a subject of some controversy, and will be discussed at greater length under the morphology of Pedilanthus. Terminology of the Involucre in Pedilanthus. — In some earlier descriptions of Pedilanthus, where the inflorescence is discussed in terms of inner and outer sepals or upper and lower lip, and even in some more recent descriptions, the discussion is almost impossible of interpretation, even with the plant in hand, because of difficulties in terminology. For the description of so complex a structure as the cyathium of Pedilanthus, a clear and exact terminology is necessary, yet, for so small a portion of the plant kingdom, a highly spe- cialized and esoteric nomenclature is not justified. It is hoped that the accompanying illustrations, especially figures 12 and 13, will aid in understanding the terms used herein. The terminology of Millspaugh (1913) has been followed, where practicable. The bilateral symmetry of the Pedilanthus cyathium is so marked, and the normal position of the involucre so indeed, necessary. Their usage, which is straightforward, is indicated in fig. 12. The most characteristic feature o Pedilanthus is the hood-like or pocket-like structure in which the glands are concealed. This has often been termed the “appendix,” but the older and more descriptive term, “spur,” ; is here taken up. The German Driisentasche (Markowski, 1912) is also appropriate and readily intelligible. This structure is, in effect, an involucral spur, and is usually markedly spur-like in form. The term appendix is awkward when the homologies with Euphorbia, with its petaloid ap- pendages, are considered, and is not so evident in meaning as spur. “Apex of the spur,” as here used, refers to the tip of the spur, not to the tip or tips of the spur lobes. The 32 ROBERT L. DRESSLER “base” of the spur is less easily defined, and, for that reason, alternative measurements are given (see dotted and solid brackets in fig. 12, and discussion under measurement, p. 94). The number of spur lobes is always considered to be four, homologous with four petaloid appendages of Euphor- bia, though the lateral lobes are always adnate to the in- volucral tube, and the two medial lobes may be completely connate. The glands within the spur are conveniently desig- nated as lateral and medial, with the medial glands sometimes split into “gland-pairs.” The spur may be partially divided into three chambers within; the parenchymatous dividing walls being “spur partitions.” The two large and con- spicuous involucral lobes are simply known as the “main” involucral lobes, and the three smaller, dorsal lobes are collectively termed the “accessory” lobes, being further designated as lateral or median. There is sometimes a which, analogous to that in some flowers, may be called the mentum. When the bracteolar structures form partitions between the groups of staminate flowers, they are referred to as bracteole-partitions, and when these are continued dorsally or anteriorly in the involucral tube as distinct ridges, these are the “bracteole-ridges.” Small projections between and beneath the accessory involucral lobes are re- ferred to as “roof lobules.” The terms used here in discussion of the flowers, pedicel, filament, calyx, etc., are used in their usual senses ; special terms such as androphore and calyculus being thought superfluous where the homologies are clear. “Peduncle,” in the Euphorbieae, is used to refer to the stem bearing the individual involucre. Where confusion may occur between the involucral bracts and the opposite, free bracts subtending the involucre (and peduncle), the latter may be distinguished as “cyme bracts.” The Involucre of Pedilanthus. — The involucral homolo- gies between Euphorbia and Pedilanthus have been correctly interpreted, on the basis of external form, by Ridola (1903) and Troll (1928), and well indicated by Michaelis’ work on development (1924), yet the morphology of Pedilanthus has remained poorly known, and a detailed comparison with Euphorbia is quite lacking. The five involucral lobes of Pedilanthus correspond to those of Euphorbia, and the spur lobes (including the adnate, lateral lobes) correspond to four THE GENUS PEDILANTHUS (EUPHORBIACEAE) 33 petaloid appendages of the subgenus Agaloma. The striking differences between Pedilanthus and Euphorbia result from the very unequal size of the involucral lobes, above the level of the glands, and the great anterior-posterior and dorso- ventral developmental deformation which places the four gids on one side of the involucre and forms the characteris- A. a bract of the . lateral view of TE V. b ggueuiee rote of — calearatus involucre. dorsa PP seg air; u la 1 bract: C. 1 (median) ¢ dorsal bract Sieben position; Lnaet e of involucre, showing relationship of the involueral brace lia sory involucral lobe; 2. spur lobe; 3. glan only the half-gland cananlnica this bract is shown the right, but the gt 1 from n gland, half of which is vascularized from the upper lateral bract (B), is shown o 6. left. Magnification ea 34 ROBERT L. DRESSLER tic spur, and a greater degree of adnation of parts than is usual in Euphorbia. This deformation and adnation of parts makes the interpretation of the involucre from external features, or from microtome sections, very difficult, but clearing and partial dissection show the relationships, at least of the vascularized parts, very well (Plates V and VI The basal part of the involucre shows a radial symmetr (or nearly so) and agrees closely with the basal structure E VI. ae Geek of Pedilanthus nodiflorus involucre. A. a bract of the lower pair; B. r lateral bract; C. dorsal (median) bract; ¢. lateral view of dorsal bract in sei ¢ ae inset, outline of involucre, showing relationship of the wero bracts (dorsal bract — visible from side). 1. accessory involucral lobe; 2 lobe; 3. gland. Magnification ea. 7. THE GENUS PEDILANTHUS (EUPHORBIACEAE) 35 of the Euphorbia cyathium. Each of the five bracts has three main vascular traces, and, in the basal portions, the five bracts are of about equal width. In the material examined of P. calcaratus, a curious crimp or zig-zag is usually found at the base of each bract trace (Plate V). This may be re- lated to the abscission layer which forms in that region after anthesis (if pollination occurs). The base of the bract trace is often a weak point, and breakage during dissection is all too easy, especially in the more weakly vascularized lower bracts. The involucral bracts of Pedilanthus fall into three types: (1) two lower bracts, which include the main involucral lobes (A in Plates V and VI) ; (2) two upper lateral bracts, which bear the lateral accessory lobes (B in Plates V and VI); and (3) a single, median dorsal bract, which bears the median accessory lobe and is bilaterally symmetrical (C in Plates V and VI). These three types are described in greater detail below. (1) The lower pair of bracts are the least complicated in structure. In those species with a scarcely elongate spur, such as P. tithymaloides or P. nodiflorus, the lower bracts show a quite simple, three-trace, foliar pattern of vasculari- zation. In those involucres with elongate spurs, however, the upper trace, or a large part of it, bends back and up into the spur, before bending sharply forward to vascularize part of the lateral spur lobe, which is not readily distin- guished from the main involucral lobe. This upper trace May occasionally contribute to the lateral gland (when present) or, if this is small, as in P. nodiflorus, may even form the entire vascular supply for the gland. More com- monly, however, no glandular tissue is directly connected with the lower bracts. There are frequently weak cross- connections between the lower bracts and the adjacent, upper lateral bracts in the spur, possibly representing, in P. cal- caratus, the area of the non-developed lateral gland. Similar- ly, cross-connections may occur ventrally between the lower racts. (2) The upper lateral bracts are involved to a much greater extent in the formation of the spur, and are conse- quently more complicated. The lower vascular trace mostly goes into the lateral gland, when this is present, the re- mainder of the trace supplying the upper part of the lateral _ < nen 36 ROBERT L. DRESSLER spur lobes. The central trace of the bract extends to a point near the glands, where a single, central branch (rarely two small branches) bends more or less sharply forward to supply the lateral accessory involucral lobe. The branches on each side of this small, central branch continue to the apex of the spur and forward into the spur lobes. The upper trace of this bract follows a pattern similar to that of the lower trace, supplying the outer half of a medial gland, with small branches often continuing into the corresponding medial spur lobe. (3) The median, dorsal bract resembles the upper por- tions of the adjacent bracts, but is bilaterally symmetrical. The lateral traces supply the inner halves of the medial glands, with small branches usually contributing to the inner halves of the medial spur lobes. The median trace roduces a small central branch to the median accessory involucral lobe, and the remaining branches go to the medial spur lobes. Cross-connections between this and the adjacent bract-systems are absent, except at and just below the glands. Interesting variation is to be found in the form and structure of the accessory involucral lobes. In P. bracteatus and its immediate allies, P. tehuacanus and P. tomentellus, and in P. nodiflorus, the accessory lobes are markedly thickened above near the base, and the thickening has the appearance of representing the up-folded and connate mar- gins of widened lobes. In P. nodiflorus this appearance is found in the vascular pattern (Plate VI; here the two margins are separated and spread apart). The central part of the vascular supply runs forward near the ceiling of the involucral tube and lateral branches bend upward to supply the upturned margins. In P. bracteatus, which presents a similar external appearance, the dorsal portions of the lobes are without vascularization, only small lateral branches, if any, extending upward. In P. tithymaloides, the accessory lobes are greatly thickened basally, but there is much less appearance of folding. The vascular supply in this species curves abruptly upward and follows the upper surface, leaving the lower part of the thickened area without vas- cularization. Small lateral branches, which resemble the larger ones of P. nodiflorus, are sometimes seen. Pedilanthus cymbiferus also has thickened accessory lobes, but the curva- ture of the vascular traces, which are similar to those of P. THE GENUS PEDILANTHUS (EUPHORBIACEAE) 37 tithymaloides, is much less pronounced. Pedilanthus pulchel- lus and P. coalcomanensis are externally similar in this feature. In the other species which have been examined, the accessory lobes curve smoothly forward and downward from their origin in the spur. The spur, it is evident, is a compound structure, involving all five involucral bracts, though not in equal degree. The spur, in any measurable sense, is not exactly homologous with the petaloid appendages of Euphorbia subgenus Agaloma. Only the spur lobes, including their usually con- nate bases, correspond to these appendages. The dotted line in fig. 11 is intended to indicate the approximate portion of the involucre which is homologous with the petaloid ap- pendages. A considerable portion of the spur (where this is elongate) represents a lateral extension of the involucral tube. This extension is always solid and parenchymatous, though Millspaugh (1913) was led to consider the peduncle of P. macrocarpus as “centrally affixed to the tube;” an impression clearly obtained from dried material in which the parenchymatous core of the tube extension had been crushed. The lobing of the spur apex in some species cor- responds, not to the number of petaloid appendages involved in its structure, but to the number of involucral bracts (three) involved at that point. Glands. — The double nature of the glands, already clearly indicated for Euphorbia, is reflected in the vascular pattern of the involucre, and, in some species, the medial glands have, phylogenetically, each split again into two parts (that is, the primordia remain distinct). In other species, the pri- mordia occasionally remain distinct as a slight anomaly. In Pedilanthus, none of the lateral branches of the median bract trace supplies the glands themselves, reflecting the proportionate increase in the gland appendages (spur lobes) and, at least in proportion to the involucre, a decrease in the amount of nectariferous tissue. The glands show a typical palisade epidermis, with thickened cuticle, which appears to slough off at maturity. The cuticle shows knob- like thickenings on the lower surface, which follow the lateral contacts between epidermal cells. The cells immedi- ately beneath the epidermis may themselves be radially elongate and somewhat palisade-like, but the remainder of the cells are relatively small and isodiametric, with densely 38 ROBERT L. DRESSLER staining cytoplasm. The gland structure of P. tithymaloides has been illustrated by Banerji (1951) and Sperlich (1939). Millspaugh described the glands of some species as “stip- itate”; since, however, the “stipes” (spur partitions) are never free, the term is inappropriate. The glands produce an abundant supply of sweet nectar. In greenhouses, where no animals can remove the nectar, the gland chamber is soon filled to overflowing (see fig. 31). The glands of the Euphorbieae have often been interpreted as stipular. Their lateral position and vascularization are such that they could be of stipular origin. As indicated by Warming (1879), however, the occurrence of clearly non- appearance. Serial sections show, for the first two genera, that the vascular pattern is basically that of Euphorbia. The glands are between the clusters of staminate flowers, but the vascularization is entirely from the lateral traces of the adjacent involucral bracts. In these genera, and probably in Neoguillauminia as well, the glands are lateral on the involucral bracts, which basally enfold the staminate flower clusters, so that the marginal glands are between these clusters, and even adjacent to the central pistillate flower The spur lobes are so formed as to enclose the gland chamber above and laterally, leaving an opening or groove between the median spur lobes and the accessory involucral lobes, where a slender beak or tongue may be inserted (P. tehuacanus is exceptional in this respect ; the spur lobes do not always fit closely together). The spur lobes are, thus, elongate and thickened, as compared with the petaloid ap- pendages of Euphorbia. The medial lobes are frequently thickened near their apices. In P. calcaratus and P. gracilis the medial lobes are each folded upon themselves, and the folded edges coalesced, the terminal portion forming a recurved thickened point, reminiscent of a plow blade in form (see Plate XVI). In several species of Pedilanthus, the color differences between the spur lobes (petaloid ap- THE GENUS PEDILANTHUS (EUPHORBIACEAE) 39 pendages) and the involucral lobes, which are usual in Euphorbia, are maintained, even though the structural boundaries are much blurred. In other species, the red color of the spur lobes may extend over part or all of the involucral tube, while the red color is quite lacking in the involucres of P. bracteatus and P. tomentellus, and nearly so in P. tehua- canus. Partitions and Bracteoles. — The spur partitions form in- teresting and curious structures, whose exact significance is difficult to determine. In P. coalcomanensis and P. macro- carpus these partitions are especially well developed and di- vide the gland chamber into three compartments terminally. In other species the spur is not so greatly lengthened beyond the glands, and the spur partitions are conspicuous only as ridges running down and forward from the medial glands to insert between the accessory involucral lobes. Other species, such as P. calcaratus and P. nodiflorus, have three terminal compartments in the gland chamber, but the lateral ones are SO small as to escape notice except in section. The glands, in P. macrocarpus at least, are partially vascularized through the spur partitions. Part of the vascular supply goes nearly to the tip of the spur and enters the glands from the dorsal side, as is shown for P. calcaratus (Plate V). The remainder of the vascular supply, particularly that of the medial glands, enters more directly from behind and beneath the glands, through the spur partition. The indirect route of the vascu- lar traces in P. calcaratus and partly in P. macrocarpus Suggests that the portion of the spur distal to the glands represents an “everted” portion of the involucral tube, with the spur partitions representing the bracteole-partitions. Position alone suggests that there may be a genetic and de- velopmental relationship between the two sets of partitions. Since the partitions are largely parenchymatous, their exact structural relationships are not easily determined. However, the ridge descending from the medial gland and inserting between the accessory involucral lobes does sometimes have a weak vascular system of its own. In P. bracteatus, a branch may run forward from just beneath the gland; this passes between the accessory involucral lobes, spreading out some- what laterally beneath the vascular systems of the accessory lobes (fig. 15). In that species, this weak vascular trace runs about half the length of the accessory lobes. A similar ney: e VIL. Fig. 15. Pedilanthus bracteatus, Dressler 1808, portion of to the n frivelns ® owing vascular supply tissues between and beneath the accessory involucral lobes (A); B. gland; C. lateral accessory lobe; ian accessory lobe. obes are separated, and th graph is slightly retouched to show per tive, magnificatio a. 10, Fig. lmeri, b of staminate flower cluster, cleared and with the pedicels separated abaxially and spread apart. The central flower is indicated a "ae a rows indicat r of flowering. The older, central flowers have already formed a ¢ bseission layer near the b The bract traces are sho eo : L. lateral brace ace; M. ian bract trace. ee AT: eri, the central staminate flower and on ne ‘alm seen from the side, iledeg trees cn labelled as in Fig. 16. Magnification ca. 17. THE GENUS PEDILANTHUS (EUPHORBIACEAE) Al branch is sometimes seen in P. macrocarpus, coming either from beneath the gland or from an adjacent branch of the lateral bract trace (which continues into the spur lobe). Much weaker branches of the same sort are seen in nodiflorus. None was observed in P. cymbiferus, P. cal- caratus or P. tithymaloides. If the spur partitions are of bracteolar nature, as suggested, this vascular pattern may represent a nearly complete fusion of the bracteolar traces with the lateral bract traces. Patterns found in Euphorbia pulcherrima suggest that such a fusion has taken place. The bracteolar tissues within the involucral tube typically form partitions which partially separate the clusters of staminate flowers basally. These partitions are more or less laciniate above into slender, filamentous, or occasionally ligulate, segments, which are tangled among the staminate flowers when well developed. These partitions are connate with the involucral tube basally, and, even in those species without bracteoles, basal ridges are present within the in- volucral tube. Such a ridge commonly continues upward beyond the point at which the bracteoles become free from the involucral wall, especially in the two dorsal bracteole groups (see fig. 13). In P. coalcomanensis a distinct ridge or keel runs the length of the involucral tube and terminates in small lobules (“roof lobules’) in the sulci between the accessory involucral lobes. These lobules are frequently found in other species, where the dorsal ridges are indistinct, such as P. macrocarpus, P. bracteatus and P. nodiflorus. The parenchymatous ceiling of the involucral tube may be partly bracteolar in all species. A single collection of P. macrocarpus (Wiggins 5345) has been seen in which some of the bracteoles contain vascular tissue. An involucre from this collection was carefully cleared and dissected, only to show that the vascular tissue did not connect with either the vascular system of the in- volucre or that of the staminate flowers. One vascularized bracteole occurred between each two adjacent staminate flower groups. One of these bracteoles had only about a milli- meter of vascular tissue in the mid-portion; the other four were vascularized throughout their lengths, but the strands ended blindly at the bases. Development of the Involucre. — The development of the cyathium of Pedilanthus has been investigated by Baillon 42 ROBERT L. DRESSLER (1858), Beille (1902), Markowski (1912) and Michaelis (1924), in all cases using P. tithymaloides ssp. tithymaloides. The development of Euphorbia has been the subject of many P.LatTe VIII. Early development of the cyathium, upper row fresh material of calcaratus, lower row preserved specimens of P. cymbiferus. gz 1 and the five staminate flower cluster pri- mordia, with the involueral bract primordia differentiating about these. The order aa" i i ia is: lower left, upper, lower right, upper left, upper right (a 2/5 s yiral), magnification ca. 50. B € nd the 2 Ps ~ 3 x a @ =| > A) 20. . t gland primordia (3) are just beginni ; . 33. C, the involucre has overtopped the staminate flowers; the gland primordia (3) are differentiated and the medial ones are differentiat ing into gland and spur lobe primordia (the lateral ones form only spur lobes in this i d t 2 Ale e : as o < =} 5 _ < sor} Vv S$ ar different; the distinction between lateral spur lobe and main involucral lobe is clear at this stage; the three carpels of the pistillate flower are seen in the center, magnifica- tion ca. 35. D. lateral view of the same apex. E. an older involuere, in which the adult form is approached, magnification ca. 30. F. the same involucre as in E, lateral view; note the cyme primordium at the base; magnification ea. 20. 1. main involucral lobe; 2. accessory involueral lobe: 3. gland (or spur lobe) primordium THE GENUS PEDILANTHUS (EUPHORBIACEAE) 43 at ae alt ak il HILL PLate IX. Fig Pedilanthus pepe rus, later stages, showing the development of the spur ate involucral tube, scale — mm. Fig. 19. Developmental anomalies in P. cumbiferus., A and B. involucres ick all bracts cortege on the ‘accessory lobe pattern,” the second with a slight Bn of zygomorphy. C. an involucre with four : lo ree free spur lobes) and only one main lobe. D. an in dos with y one accessory lobe (no free spur tobias} and four main lobes. Seale 1 44 ROBERT L. DRESSLER investigations, those of Schmidt (1907) and Michaelis (1924) being especially useful. The early developmental stages in these genera agree closely. The dichasial primor- dium differentiates the two protecting cyme bracts at an early stage, and the developing cyathium is completely con- cealed by these bracts during the earlier stages. The first steps of the external differentiation of the young cyathium are the formation of five smaller primordia around the cen- tral portion which becomes the pistillate flower. Those five primordia, which represent the staminate flower clusters and their subtending involucral bracts, develop in a 2/5 phyllotactic order, numbers 1 and 3 being abaxial (ventral), number 2 being adaxial (dorsal) and numbers 4 and 5 de- veloping into the lateral bracts. These primordia differenti- ate into distinct upper portions, the primordia of the staminate flower clusters, and a lower 5-lobed ring, the united primordia of the gamophyllous involucre (Plate Vuil A). This involucral ring soon overtops the developing staminate flowers, but before these latter are completely concealed, the first signs of marked zygomorphy appear. quite concealing the accessory involucral lobes), giving a structure like that shown in Plate VIII E and F. At this stage, though the glands and their appendages (spur lobes) are displaced dorsally and the ventral involucral lobes are disproportionately large, the involucre is readily comparable with that of many species of Euphorbia. It 1s the intercalary growth of later stages which brings about the bizarre forms of the mature involucre. In P. cymbiferus (fig. 18) and similar forms, this is seen to involve dispro- portionate growth in two areas: (1) the two ventral (main) involucral lobes, which elongate and make up the greater part of the involucral tube above the base; and (2) the involucral tube and the spur lobes in the immediate vicinity of the glands, which undergo great differential growth to form the spur. In P. macrocarpus and P. Palmeri, the spur THE GENUS PEDILANTHUS (EUPHORBIACEAE) 45 growth greatly exceeds the enlargement of the involucral lobes, so that the spur forms the greater and most con- spicuous part of the mature cyathium. In P. tehuacanus, P. tithymaloides and P. nodiflorus, on the contrary, spur elongation is quite limited. Pedilanthus tithymaloides (and, doubtless, P. coalcomanensis) differs in its earlier stages from the development outlined above, in that the two medial spur lobes are connate throughout. Developmental Anomalies. — In many species of Euphor- bia, the occurrence of abnormal cyathia is frequent, especial- ly involving the cyathium terminating a main stem, the first and central cyathium of a pleiochasium (E. capitulata Reichb. is noteworthy in that the normal axillary cyathia abort and the single, terminal cyathium is regularly “ab- normal,” Schmidt, 1907). Similar phenomena are to be found in Pedilanthus, chiefly involving imbalance among the different patterns of bract development. The most com- mon pattern of abnormal development is that in which all five bracts of an involucre develop along the accessory lobe pattern (fig. 19, A and B). In this case, the spur cannot elongate, but the gland chamber develops as a ring about the involucre. The five connate accessory involucral lobes leave such a small opening that only the pistillate flower is exserted, most of the staminate flowers remaining packed in the base of the involucre. The opposite case, in which all five bracts develop on the main lobe pattern, has only been seen once, on a small cutting of P. bracteatus. This involucre, essentially that of a Euphorbia without glands, failed to reach maturity, doubtless because of the small size of the plant. Intermediate patterns of abnormal growth are oc- casionally found, two of which are shown in fig. 19. In both of these cases, the normal number of five bracts is found, but in one case, four of the bracts form “main” involucral lobes, the fifth being a bilaterally symmetrical “lateral accessory lobe” (each half forming part of a lateral spur lobe) ; in the other, only one “main” lobe is formed, there being two “median accessory lobes.” Since the photograph was taken, the remaining combination, an involucre with 2 accessory lobes and 3 main lobes, has been observed. There may be, basically, two patterns of development involved: the spur pattern and the non-spur (main lobe) pattern. If this is the case, the upper lateral bracts normally represent 46 ROBERT L. DRESSLER intermediates, with the upper portions developing on the spur pattern and the lower portions approaching the main lobe pattern. a zig-zag manner. It is thus not surprising that Wydler (1845) and many subsequent students have regarded each cluster of staminate flowers as a simple cincinnus or scorploid cyme. This view has been defended more recently by Schoute (1937). Schmidt (1907), on the basis of de- velopment and vascular pattern, considered each staminate flower cluster to represent a dichasium, with each branch sort of dichasium found in the primitive genera Anthostema and Dichostemma, as diagrammatically shown in fig. 20. Schmidt (1907) has reported that the primordia of both the second and third flowers of a cluster originate from the primordium of the first flower, and that the fourth is derived from the second, the fifth from the third, the sixth from the fourth, etc. This is the primary basis for his THE GENUS PEDILANTHUS (EUPHORBIACEAE) 47 hypothesis that the cluster represents a dichasium. Michaelis (1924) examined Pedilanthus tithymaloides and several species of Euphorbia, and reports that these do not show the order of development reported by Schmidt, but that the third primordium is derived from the second, the fourth from the third, etc. In my own work, I have examined a number of early stages of Pedilanthus (several species) and must report that the staminate flower primordia are so compacted that, on the basis of external features, at least, neither pattern of development can be proven to the exclusion of the other. The vascularization of the staminate flower cluster is basically two-parted (in Euphorbia Xantii the two-parted nature is so marked that the two halves are independently inserted on the lateral bract traces; the primary staminate flower is aborted, or, at least, not centrally located, as in most Euphorbieae). The first staminate flower of each cluster is most commonly between the two halves, as illustrated by Schmidt (1907) and by Haber (1925), though it is often a @) = ~ ‘ ‘ ‘ eae ) a ~~ Circles The two hypothetical cesar: of the staminate flower cluster. e indicating order of flowering. The crescents ed circles indicate suppressed flowers; 3. eses. diagrams of the monochasial (above) and dichasial (below) hypot 48 ROBERT L. DRESSLER little more closely associated with one half or the other (as the primary flower in fig. 16 is markedly associated with the half on the left side of the illustration). The two branches from this primary staminate flower form, or closely simulate, much condensed monochasia. It is possible, as Schoute (1937) suggests, that the two separate sympodia of floral traces are merely the result of extreme condensa- tion of a simple cincinnus. The vascular pattern alone, however, strongly supports the hypothesis of a dichasial nature for the staminate flower cluster of Euphorbia and Pedilanthus. Studies of Neoguillauminia, Calycopeplus and primitive species of Euphorbia will probably serve to clarify this controversy. As Markowski (1912) has observed, the zygomorphy of Pedilanthus extends even to the staminate flower clusters, in that each of the two ventral clusters frequently has one less flower than does each of the three dorsal clusters. This tendency is especially striking in P. nodiflorus, in which the pistillate flower is markedly excentric and the ventral clus- ters of staminate flowers often have 2-3 less flowers each than the dorsal clusters. The staminate pedicels elongate greatly in development, so that the staminate flowers are exserted from the orifice of the involucral tube, as the an- thers mature. The pedicels of the lower clusters curve and twist in such a way that each flower emerges from the top of the orifice, under the accessory involucral lobes, just before anthesis, pushing down the older flowers and being displaced itself after the pollen is shed. All anthers are oriented with the locules opening dorsally (with respect to the involucre). The pedicels are separated from the fila- ments by a transverse articulation and abscission layer, without trace of perianth, and with only slight thickening. The filaments are often differentiated from the pedicels by pubescence or color, but are structurally similar. It should be noted that the articulation in Anthostema and Dichostemma is directly beneath the rudimentary calyx, unlike the struc- ture described by Nozeran (1953) for Dalechampia and some members of the Hippomaneae. There is surely no “ peri- cladium” involved in the Euphorbieae, which do not fit well into the evolutionary (?) scheme offered by Nozeran (p. 114). The vascular trace of the filament ends with a club- like thickening in the connective, between the two halves THE GENUS PEDILANTHUS (EUPHORBIACEAE) 49 of the anther. The anther is of a normal four-locular type, becoming two-celled at maturity. Dehiscence is longitudinal and abaxial. The development of the anther in P. tithy- maloides has been investigated by Banerji (1951). The tapetum is glandular, and the tapetal cells largely become binucleate, though quadrinucleate, and rarely trinucleate, cells are found. Meiosis is normal, the tetrads show a tetra- hedral arrangement, and cytokinesis takes place by furrow- ing. The pollen grains are reportedly binucleate when shed. Pollen Structure. — While Erdtman (1952) has devoted special attention to the pollen of the Euphorbiaceae, this large and interesting family is still imperfectly sampled, and the tribe Euphorbieae, in particular, has received only cur- sory attention from pollen students. The great interest in the identification of fossil pollen has led to the frequent adoption of methods designed to make pollen samples from living plants as much like the fossil pollen grains as possible. While this is necessary for the paleontologist, such methods obscure or destroy features of great interest to the botanist studying recent plants. In the present study, methods have been developed, largely by accident, which have proven quite satisfactory for Euphorbiaceous pollen. Details of exine structure are well shown by grains which are rendered transparent and expanded by treatment with sodium hy- droxide and lactic acid. Since the latter has not proven satisfactory for permanent slides, Hoyer’s mounting medium (Beeks, 1955) has been used. Raphide-like crystals are often formed in this medium, if it is improperly prepared, but pollen mounts are usable even when this occurs. The proto- plast and intine are destroyed by sodium hydroxide, but their features may be seen in pollen preserved in FAA and treated with lactic acid. The protoplasts of fresh or dry pollen, or of pollen freshly killed in FAA, tend to swell ex- cessively and obscure the features of the intine. Moderately good restoration of pollen from herbarium specimens may be obtained by storing the anthers in FAA. In practice, the following schedule has given good results: 1. Two or more, usually six to ten, mature but unopened anthers are removed from preserved material and half of the sample set aside in a small vial of FAA. 2. The remain- ing anthers are placed in 6% NaOH and heated in a paraffin oven for about six hours (a little longer may be required 50 - ROBERT L. DRESSLER for any inaperturate pollen types, and the large, inapertur- ate, “croton pattern” grains of Jatropha require about eight hours). 3. The material from NaOH is washed in water, and the two portions of the sample combined and heated in 85% lactic acid in a boiling water bath for about ten minutes. 4. All the anthers are broken open on a slide in a large drop of Hoyer’s medium. The larger fragments of the anthers may be removed under a dissecting microscope, if desired, before the coverslip is applied. The slide should be placed on the microscope lamp or other heat source for a few min- utes, until the mounting medium has spread evenly under the coverslip. The size of a pollen grain depends to a very great degree on the treatment which it has received. For this reason, measurements were taken from samples of pollen of P. Palmeri (all from one plant of Dressler 1064) which had been given a number of the different treatments used in the present study (Table 3). Similarly, at least one sample from each species, treated with NaOH and lactic acid, has been measured, and the average dimensions given in Table 4. Even though all samples were given similar treatment, some of the size differences may be due to differences in degree of expansion, and the size measurements, from so few samples, should only be taken to indicate the major trends in pollen grain size. In both tables, the averages represent thirty measurements where possible, and obviously shrunken or abnormal grains were not measured. The pollen grains of most of the species are very much of the same type, though differing in details of wall thickness, coarseness of sculpturing and dimensions of furrows, furrow margins and pores. The predominant pollen type will be described first, and then the few markedly divergent species (P. Finkii, P. tehuacanus and P. tomentellus) will be dis- priser The terminology used is that of Faegri and Iverson 50). The grains are tricolporate (three furrows, each contain- ing a pore) and, when shed, the three furrows are strongly plicate and folded inward, so that the grain is a football- shaped ellipsoid (Plate X-F), strongly three-lobed in equato- rial section. In this form, very little of the structure can be observed. When the grain contacts an aqueous medium, the entire structure swells strongly, expanding the wall and THE GENUS PEDILANTHUS (EUPHORBIACEAE) 51 TABLE 3. Measurements of pollen from Pedilanthus Palmeri (Dress- ler 1064) after various methods of preparation. All measurements in microns. TREATMENT POLAR AXIS EQUATORIAL DIAMETER range average range average NaOH ae lactic acid 72.5 - 89.8 79.3 89.8 -100.1 94.4 Lactic acid 69.1 - 76.0 73.3 85.2 - 93.2 89.7 NaOH 71.4 - 80.6 75.1 86.3 - 99.0 91.6 Mounted in Hoyer’s 68.0 - 73.7 70.5 81.7 - 87.5 85.3 from FAA (no treatment) Acetylated, dry 66.8 - 81.7 76.1 57.6 - 74.8 66.4 Freshly shed pollen mounted in oil 74.8 - 81.7 19.1 51.8 - 61.0 57.6 BLE 4. Average dimensions (in microns) of pollen erame treated nkii rep with lactic acid. measurem re- sents the distance from furrow to opposite angle; the distance aig angles averages 104 microns. In the inaperturate grain of P. u- acanus poles and equator cannot be distinguished. SPECIES AND COLLECTION POLAR EQUATORIAL AXIS DIAMETER P. Palmeri, D 1064 79.3 94.4 P, Finkii, Thompson, s. n. 64.7 96.6 P. calearatus, D 1067 59.6 71.5 P. gracilis, Hinton 1097 60.1 68.1 P. pulchellus, Conzatti, Reko & Makrinius 3106 72.6 88.9 P. coaleomanensis, Hinton 12685 68.7 79.8 P. cymbiferus, D 1368 ca. 92.0 105.1 P. macrocarpus, D 1090 63.7 78.7 ntry 767 65.2 73.5 P. bracteatus, D 978, northern Sinaloa 19.4 96.7 D 979 74.9 95.6 803 97.0 112.3 P. tomentellus, Pringle 4912 Tao 82.7 P. tehuacanus, D 1795 74. P. nodiflorus, D 1362 72.8 91.4 Pr ghee aghe soc 2 ssp. tithymaloides, D 1376, Cuicatlan, Oaxaca 77.6 87.5 D 13878 79.1 94.8 Cutak, Isthmus of Tehuantepec 82.6 103.8 ssp. parasiticus, D 1 San Andrés Tuxtla, Veracruz 71.0 82.4 ssp. jamaicensis, How Four Mile Wood, Jamaica 67.0 77.7 ssp. angustifolius, Britton - Cowell & Brown 4607 55.0 67.6 THE GENUS PEDILANTHUS (EUPHORBIACEAE) 53 exposing the furrows and pores. In the fully expanded state the equatorial diameter of the pollen grain somewhat exceeds the polar axis, giving the grain a more or less flat- tened or oblate form. The expanded grains vary from large to very large (see Table 4). Each furrow (colpus) is fusi- form in outline, tapering abruptly at each end of the pore, and attenuate to the blunt, or occasionally slightly bifid apex (Plate X-G). The furrows have distinct, narrow margins which are thinner than the remainder of the exine and have little or no sculpturing. These margins fold inward in the dry grain. The pores are distinct, large and oblong, occupy- ing about one third to one half the length of the furrow, parallel to, and completely contained within, the furrow. Small irregular bits of exinous material or “opercula’” some- times occur on the pore, especially near the ends (usually absent after NaOH treatment). The exine varies from about 3-3.5 microns in thickness (P. calcaratus) to about 5 microns (P. cymbiferus), or even 6-7 microns (P. bracteatus, Dressler 1803), in the thickest portion, midway between the furrows. The exine thins gradually and then abruptly to the finely or scarcely sculp- tured furrow margin. The inner,non-sculptured portion of the exine (endexine) is quite thin and inconspicuous. Thin sections (Plate X-H) show the sculptured portion (ektexine) to be composed of narrow, radial rods (columellae), whose uppermost portions are fused into a perforate reticulum (tectum), which gives the grains their characteristic surface texture. The intine of Pedilanthus pollen shows distinct longitu- dinal thickenings, which, though characteristic of many Euphorbieae, seem never to have been described. These thickenings occur in three pairs which alternate with (or flank) the pores. The thickenings parallel the furrows, and are of about the same length as the furrows. In material treated with lactic acid, the thickenings are visible as hyaline depressions in the protoplast (Plate X-C). The protoplasts of grains treated as above retain their form when the exine is broken away; such a protoplast is shown in Plate X-B (this rigidity of the protoplast may only be the result of thorough fixation in FAA). The thickenings stain with fast green (as does the entire intine) and, in fresh material, stain rapidly and strongly with ruthenium red, probably 54 ROBERT L. DRESSLER The most aberrant pollen type within Pedilanthus is that of P. Finkii (Plate X-E). This is markedly triangular in equatorial outline, even when fully expanded, the furrows occurring on the sides of the triangle, between the blunt angles. The distance between adjacent angles (average THE GENUS PEDILANTHUS (EUPHORBIACEAE) 55 104 microns) is nearly a tenth greater than the distance between an angle and the opposite pore (average 96.6 mi- crons). The grains are very markedly flattened in the polar dimension. The furrows of P. Finkii are narrowly elliptic, with no trace of a pore, and bear distinct, though irregular, “opercula”’ similar to those of other species, but better de- veloped. The furrow margins, in strong contrast to those of the other species, are markedly thickened. These thick mar- gins, unlike the ‘‘costa colpi” described by Faegri and Iver- sen (1950, p. 22) and Erdtman (1952, ‘“crassimarginate grains’), are largely composed of ektexine and show distinct sculpturing similar to that of the remainder of the grain, though the endexine, too, appears to be thickened about the furrow. From the structure of these grains, it appears un- likely that any great volume changes could occur, such as regularly take place in the porate grains of other species. Intine thickenings occur, but are small and inconspicuous. The pollen grains of P. tehuacanus are very strikingly different, in that they are completely inaperturate (Plate X-D). There are slight irregularities in the thickness of the exine, but no pores or furrows. Distinct intine thickenings occur, but these appear to follow no set pattern in their dis- tribution. In texture and size, the grains of P. tehuacanus resemble those of P. tomentellus, which, also, are somewhat aberrant. A large percentage of the grains examined of this last species are shriveled, and evidently nonviable. Of those grains which expand normally, however, a large number lack pores, the furrows in these grains remaining narrow and unbroken. The appearance of the pollen strongly sug- gests that P. tomentellus is an abnormal, and perhaps hy- brid, population. Pistillate Flower. — The pistillate flower, which termi- nates the axis of the cyathium, is similar to that of Euphorbia. In several species, the terete pedicel elongates markedly and curves downward between the main involucral lobes, after anthesis, so that the ovary and styles are beneath the involucre and directed toward the peduncle when the staminate flowers mature. Self-pollination of a cyathium is thus effectively prevented. This does not occur, however, In P. pulchellus or P. coalcomanensis (and is variable in P. tithymaloides) ; in these two species the fruit develops with- in the involucral tube. A ring-like or very slightly lobed 56 ROBERT L. DRESSLER “disk” occurs at the base of the ovary. This structure, totally lacking vascular tissue, is the only trace of a ‘‘calyx” to be found in the genus. Such a well-developed rudimentary perianth as that illustrated by Baillon (1858) or Michaelis (1924) has never been observed in the present study. Their drawings suggest material distorted by drying or rather crudely drawn. References to a distinct calyx in generic descriptions appear to stem from the inclusion in the genus Pedilanthus of Cubanthus, which does have a small but dis- tinct calyx. ovary. The three to five small central traces of the peduncle divide so as to form nine traces in the upper pedicel, three outer single strands and three alternating, inner pairs. The outer traces form the dorsal carpel traces, which are con- tinuous through the style. These traces bear small lateral branches which anastomose through the carpel wall and converge between the dorsal traces at about the level of the placentae, where they pass inward through the septae to join with the ventral carpel traces. These lateral branches form a network which becomes more prominent as the fruit enlarges. The paired ventral traces from adjacent carpels pass upward through the axis of the ovary to just below the level of the placentae, where the pairs diverge. The diverg- ing ventral trace pairs each fuse and terminate in the upper septae, where they merge with the lateral branches from the dorsal traces. One or two small strands may (but often do not) arise from each such plexus and pass through the style, alternating with the dorsal carpel traces, and usually closer to the center of the style. A small vascular strand arises from each ventral trace of a carpel, shortly above the point where the trace pairs diverge and the two fuse to form the vascular supply of the anatropous, pendulous ovule. This strand terminates in a radiating, spoke-like disk at the chalaza. The styles are elongate and nearly completely connate, and the three stigmas are each bifid. The dorsal carpel traces each divide into two to vascularize the stigmas, and this division may, in some cases, occur near the base of the style. The compound style is solid, with a central core or dense, collenchyma-like “stigmatoid tissue.” Irregular THE GENUS PEDILANTHUS (EUPHORBIACEAE) 57 vascular bundles are often formed about this central core. These bundles may connect with the outer traces below or may end blindly. The embryology of P. tithymaloides has been studied by Arnoldi (1912), Markowski (1912) and Banerji (1951). The ovules are bitegmic, the inner integument being formed first; the outer integument is formed soon after the inner and grows rapidly, covering the nucellus by the megaspore mother-cell stage. The outer integument is about four cells thick, except about the micropyle, where it is thicker. The Inner integument is relatively thick, being about 8-10 cell layers in thickness. Both integuments take part in the formation of the micropyle, which is relatively wide. The nucellus is relatively massive and more or less pointed; the beak projects into, but not through, the micropyle. The outer- most layer of beak cells is somewhat elongated (markedly so in P. tithymaloides and P. nodiflorus, less so in P. cym- biferus and P. calcaratus). There is a single archesporial megaspore mother-cell. Reduction division is normal, result- ing in a linear tetrad (only three cells are formed, according to Markowski). The embryo-sac development follows the normal or “Polygonum” pattern. The antipodals are small and ephemeral, as in a number of Euphorbiaceae. The de- velopment of the embryo has not been studied. The early development of the endosperm follows the “nuclear” pattern. Obturator and Caruncle. — The obturator, a characteristic feature of the family, arises from the placenta, above the Ovule, at an early stage in ovule development. At the time of pollination, it forms a prominent mass of matted, elongate, almost mycelium-like cells which cap the micropylar end of the ovule. The cells of the obturator penetrate the micropyle and contact the beak of the nucellus. The obturator is continuous above with the stigmatoid tissue of the style and is thought to provide a pathway to the nucellus for the pollen tube. The obturator is representative of the type “I” de- scribed by Schweiger (1905, p. 375), thus agreeing with Euphorbia. The cells of the obturator often contain starch grains. Soon after pollination, the obturator declines and is supplanted from beneath by a proliferation of the funiculus, the caruncle. This parenchymatous structure caps the seed (much as the obturator caps the ovule) until very late in 58 ROBERT L. DRESSLER seed development. In Pedilanthus the caruncle remains firm- ly associated with the placental region, and, at maturity, shrivels and remains on the columella when the seed is shed. Here, the caruncle appears to have a nutritive function in seed development. In those species of Euphorbia with a well developed caruncle on the mature seed, the caruncle is re- ported to develop from the outer integument, rather than the funiculus (Schweiger, 1905) ; in Pedilanthus, the outer rim of the integument, at least, makes at most a negligible contribution to the caruncle, if the structure here described may be included under that term. It is interesting to note that the caruncle is often (perhaps always) well developed in all three locules, even when one ovule aborts from non- fertilization or other reason. Fruit and Seed. — The fruit of most species of Pedilanthus is a capsule of the sort characteristic for the family: a septicidal and loculicidal capsule which dehisces explosively, throwing the seeds with surprising force, and leaving a three-angled or three-winged, more or less clavate, central axis, the columella. The endocarp of the fruit wall becomes woody with age and is so constructed that strong, twisting tensions are set up in drying. At maturity the largely parenchymatous exocarp shrinks and finally ruptures along the lines of dehiscence. The carpels split dorsally along the mid-vein and ventrally near the lateral traces, which remain in the columella. The septae split down to the columella. The columella is basally slender and slightly three-winged or angled (from the ventral portions of the three septae), and distally abruptly expanded (paralleling the diverging ventral carpel traces), where the three septal wings alternate with the persistent funicular “caruncles.” A careful study of the anatomy and mechanism of dehiscence in the Euphorbia fruit would be of very great interest. Pedilanthus macro- carpus is anomalous in that the fruit is corky and indehiscent. The endocarp is thin and papery and the exocarp thick and spongy. What the function of this structure may be — to float during flash-floods, to hold moisture during germina- tion, or some other — is not known. The seeds of Pedilanthus are more or less ovoid in form. The hilum is subterminal and roughly elliptical. The raphe, extending ventrally from the hilum to the chalaza, is quite prominent. No caruncle is present on the mature seed. The THE GENUS PEDILANTHUS (EUPHORBIACEAE) 59 seed coat, whose structure has not been investigated, is either smooth or finely tuberculate. Endosperm is abundant and the embryo is straight, with plane cotyledons which are somewhat fleshy. Cytology. — The meiotic chromosomes of several taxa have been examined, and the haploid chromosome number determined. The chromosomes are relatively small (see Perry, 1943, for comparison with other Euphorbiaceae) , but can often be counted with some confidence at diakinesis or metaphase I. At other stages they are much more difficult to count. The chromosome numbers, so far as determined, are given in Table 5. TABLE 5. CHROMOSOME NUMBERS IN PEDILANTHUS. Species and collection Haploid number P, eri, 64 16 P. calearatus, D 1067 16 P. macrocarpus, D 1090 17? P. bracteatus, D 978 17 979 17 P. nodiflorus, D 1809 7? P. tithymaloides ssp. tithymaloides, Cutak, Isthmus of Ps a ar Be 17 D 1366, cult., San Andrés oni Verac 17 D 1378, Tomellin Canyon, Oax 17 ssp. Smallii, cult., Harvard Biological Laboratories 17 “P. tithym aloides, “ cult., West Bengal, India 18 ( Renee “P. tithy patois, ” cult., Florida (Perry, 1943) (18) The data in the above table indicate that there is an aneuploid series within Pedilanthus. Perry (1943) has shown that the subgenera Agaloma, Chamaesyce and Poinsettia, of Euphorbia, all have a basic chromosome number of seven (without aneuploidy, though this is clearly found in the Subgenus E'sula). It may be suggested that Pedilanthus was derived from a tetraploid member of Euphorbia, subgenus Agaloma, with n=14, but more counts are needed in both genera, especially from the woody members. Whatever the primitive base number, there is clearly no basis for consider- ing it to be 9, as did Darlington and Wylie (1955), on the basis of the counts of Perry and Banerji. There appears to be variation within P. tithymaloides, and it is unfortunate that the exact origin and nature of the plants studied by Perry 60 ROBERT L. DRESSLER and Banerji are not known (they might be P. t. ssp. Smallii, or possibly P. t. ssp. parasiticus). In one preparation of P. t. ssp. tithymaloides (Dressler 1366) a single telophase II was found in which the chromosomes of all four daughter cells were easily counted. In this instance, unequal division had oc- curred, and while two of the cells each contained 17 chromo- somes, the remaining cells showed 16 and 18, respectively. “= () : a is @e, ae . e *'» ‘, 308 eo @ &. 2% 4a A 21, Chromosomes of Pedilanthus. A. P. t. subsp. vibe NEE (Cutak, ee of Tehuantepec), diakinesis, showing 17 pairs of chro n: nucleolus; B. P. bracteatus, Dressler 979, at metaphase I, note that in ain ¢ case one chromosome pair is markedly larger than all others, magnification ca. 1440 This isolated case of imbalance may, however, be without great significance, as it is thought that simple reduplication of a chromosome pair is unlikely to result in a stable, in- creased basic number (Stebbins, 1950). No mitotic chromo- somes have been examined in the present study, and little can be said concerning chromosome morphology. In the meiotic preparations, however, there appears to be some size varia- tion and one chromosome pair usually appears much larger than the others, in all the species studied. PART III. DISTRIBUTION AND EVOLUTION LOCAL DISTRIBUTION Most populations of Pedilanthus are characteristically local and discontinuous. Though the plants are numerous, where found, they are rarely frequent over an extensive area. THE GENUS PEDILANTHUS (EUPHORBIACEAE) 61 The only exception, which I have observed, is the extensive occurrence of P. tithymaloides in the Isthmus of Tehuan- tepec, Oaxaca, Mexico. There the plants are frequent along the highway for many kilometers, in arid tropical scrub. Elsewhere, Pedilanthus appears to be local in distribution, even when the vegetation type is relatively constant over a large area. While an adequate explanation of this tendency to local distribution is not available, the significance of such a discontinuous and localized population structure for evo- lutionary processes is generally recognized. Soil type may be important in determining the occurrence of Pedilanthus. Several of the species, at least, are markedly calciphilous. Pedilanthus cymbiferus, P. nodiflorus and probably P. tehuacanus, seem to be restricted to calcareous regions; P. calcaratus, P. bracteatus and P. tithymaloides have all been observed on limestone outcroppings, but are not clearly limited to such areas. The last species (P. t. ssp. angustifolius) has been reported on serpentine soil in Puerto Rico (Wheeler, Cutak and White, 1944). Observations are lacking or inconclusive for the remaining species. The different species of Pedilanthus are to be found in rather diverse habitats, as might be expected from the varied vegetative features. A few species are mesophytic or include mesophytic populations. Several are more or less markedly xerophytic. The degree of xeric adaptation rather closely parallels the environment or environments in which a species is to be found. In Table 6 the vegetation types in which Pedilanthus is found are considered as a spectrum, ranging from tropical evergreen forest to cactus desert. The termi- nology is that of Leopold (1950), though in the present paper applied to broad vegetation types, rather than to geographic zones. These vegetation types do, in fact, intergrade, and the distinctions may at times be difficult or meaningless. It will be seen that the species do have characteristic ranges in habitat, the more widespread and variable species often ranging into two vegetation types, and the very widespread P. tithymaloides occurring in a wide range of habitats. Pedi- lanthus calearatus occurs on rocky, well-drained sites when in the more mesic forest types ; P. macrocarpus is to be found only in the drier, more open types of thorn scrub; and P. bracteatus in those types which approach thorn forest. Some of the habitats are shown in Plates XI and XII. ROBERT L. DRESSLER PLATE XI. Habitats of Pe dilanthus. Fig, sutiérrez, Chia tithymalo oce a 1,300 m. elevation. Fig. 23. m a, July; arid thorn forest, approaching cactus desert. Hihepnialoibes is locally abundant. THE GENUS PEDILANTHUS (EUPHORBIACEAE) 63 fucatan, - Tehuackn, AS = PLATE XII. Ha abitats of Pedilanthus. Fig. 24. Thorn forest near September, near sea level; the habitat of P. aisdihorus. Fig. 25. Des Pr extensive colonies in the stedinenelt (arrow). 1ebla, September; P. cymbiferus forms 64 Po SON. Pe Ne ROBERT L. DRESSLER TABLE 6. Habitats occupied by different species of Pedilanthus. Information is scanty or circumstantial for those species marked with a query Palmeri Finkii calcaratus gracilis coaleomanensis pulchellus cymbiferus Macrocarpus bracteatus tomentellus tehuacanus nodiflorus tithymaloides Tropical Arid deciduous Thorn tropical Desert forest forest serub a, a as | lan —— oe ae oe we | PHOTOPERIOD Most species of Pedilanthus flower in the winter and are markedly “short-day” plants, but there are several excep- tions to this generality. As far as known, P. cymbiferus, P. tehuacanus and P. nodiflorus are all summer flowering spe- cies. Some populations of P. bracteatus are summer flower- ing (Taxco, Guerrero, but not the nearby Iguala collection), THE GENUS PEDILANTHUS (EUPHORBIACEAE) 65 and plants from other areas (northern Sinaloa) flower irregularly in cultivation. Pedilanthus macrocarpus appears to be largely winter flowering in nature, but flowers in spring and summer when cultivated in Massachusetts. The north- ernmost and westernmost Mexican populations of P. tithy- maloides flower in summer, and the plants in Tomellin Canyon, Oaxaca, flower sporadically during August (but much less, apparently, than in the winter). This species is also variable as to flowering season in other areas. There seems to be a tendency for the plants of more arid habitats (as P. cymbiferus, etc.) to flower in the summer. Pedilan- thus macrocarpus is an exception to this trend. The areas in which summer flowering is predominant may be those in which hummingbirds are of seasonal occurrence. The photoperiodic response of Pedilanthus was unex- pectedly demonstrated in the winter of 1955-56, when cultivated plants of P. Palmeri, P. calearatus and P. tithy- maloides were beginning to flower. The lighting system in the greenhouse was (anonymously and apparently inad- vertently) reset for long-day. This was first evidenced by a complete cessation of flowering in the first two species and continued abortion of young cyathia of the last. When the condition was discovered and corrected, P. tithymaloides proceeded to flower normally, but no flower primordia re- mained on the other species, and the expected cytological involucre primordia continued to differentiate, but the young cyathia aborted. GEOGRAPHIC DISTRIBUTION Pedilanthus is characteristically a Mexican genus. Only one species, the problematical P. Millspaughii, is found en- tirely outside of Mexico; two species, P. calearatus and P. nodiflorus, range south to Guatemala and Honduras, respec- tively; and P. tithymaloides ranges widely in Caribbean America (with one poorly known form in the Amazon basin). Several species appear to have very restricted ranges, but this is by no means always the case. Pedilanthus tithymaloi- des is, as noted above, very wide-ranging ; P. calearatus and 66 ROBERT L. DRESSLER P. bracteatus have wide ranges and P. Palmeri and P. macrocarpus also occur over relatively large areas. Pedilan- thus Finkii, P. cymbiferus, P. tomentellus and P. nodiflorus, on the other hand, seem to have rather restricted ranges, and the remaining species are known only from the type locali- ties. It is probable that further collecting will show all of the species to be less local than present knowledge implies. Note, for example, the wide range of P. Palmeri, which was described as recently as 1913. Its range is largely known through the collections of George B. Hinton. Though the species of Pedilanthus are usually quite dis- tinctive, sympatric species are unusual and seem to occur only very rarely within the species groups. Pedilanthus cal- caratus and P. tithymaloides, which are in quite different species groups, have been found together in tropical decid- uous forest in Chiapas (see fig. 22). Pedilanthus Palmeri and P. calcaratus, P. Finkii and P. calcaratus, P. calcaratus and P. pulchellus (found together, to judge from the mixture in Conzatti, Reko and Makrinius 3106), P. cymbiferus and P. tehuacanus; P. bracteatus and P. tithymaloides, and (if P. tithymaloides is native in coastal Yucatan) P. nodiflorus and P. tithymaloides are all pairs of species that occur in the same regions, but are not known to share the same habitats. Similarly, the ends of the P. tithymaloides circle of subspecies overlap in the West Indies, but there is no evidence that these broadly sympatric taxa actually occur together in the same habitat. As noted above, the genus is essentially Mexican. There is no reason to doubt that the genus evolved in tropical Mexico, its present center of diversity. While its “center of origin” cannot be more exactly determined, it is interest- ing that the greatest number of species is to be found in western Mexico. Furthermore, in four of the five species groups, the least specialized species are to be found in western Mexico: P. Palmeri, P. gracilis, P. coaleomanensis, P. pulchellus and P. bracteatus. In the remaining species group, both species are rather specialized. These two species, P. nodiflorus and P. tithymaloides, seem to be more markedly Central American in distribution, though the latter is wide- spread in Mexico, as well. Pedilanthus is probably a re- cent invader in South America, having arrived in the late Tertiary, with the reestablishment of land connections with THE GENUS PEDILANTHUS (EUPHORBIACEAE) 67 North America. The development of the semi-arid corridor across northern South America probably facilitated the spread of P. tithymaloides in that continent. The dispersal and evolution of this species will be discussed in greater de- tail in later paragraphs. BIOTIC RELATIONSHIPS The ecological relationships between Pedilanthus and other organisms are scarcely known, though one phase of these relationships, the pollination pattern, is probably of basic importance to the evolution of the genus. Several cases have been observed of arthropods living on or in Pedi- lanthus. The population of P. bracteatus near Taxco, Guer- rero, is extensively affected by a lepidopteran whose larvae feed in the cyathia and developing fruit. When a larva in- fests an involucre, the staminate flowers, and sometimes the glands of the spur, are eaten. Many involucres are thus emasculated ; nevertheless, a large proportion of the capsules develop. The damage caused by destroying the developing seed is probably more directly deleterious to the Pedilanthus population. Damage from similar infestations of the in- volucre has been noted occasionally in herbarium material of P. macrocarpus and rather frequently in P. tithymaloides ssp. padifolius. In P. nodiflorus one case was observed in which a lepidopteran larva (Phycitidae) had bound several involucres together with a web and was feeding on these involucres. Smaller organisms, notably thrips (Thysanura) and mites (Acarina), often find the involucres both a secure haven and an abundant source of food. Both are occasionally found as dried and broken bodies in herbarium specimens, and they are probably to be expected in all or most species. They are usually in the involucral tube, among the staminate flowers and bracteoles, and only rarely in the spur. Thrips of the genus Frankliniella (in the tritici-cephalica complex) were found to be abundant in P. cymbiferus (Dressler 1368), P. bracteatus (Dressler 1803) and P. tithymaloides (Dressler 1378), while mites (Phytoseiidae) were recovered from P. bracteatus (Dressler 1803), P. nodiflorus (Dressler 1362) and P. tithymaloides (Dressler 1378). Ants visit the involu- cres of Pedilanthus, apparently seeking the nectar exuding 68 ROBERT L. DRESSLER from the spurs. In a few cases dried ants (Monomorium and Solenopsis) have been found in the spurs of herbarium speci- mens of P. calcaratus and P. tithymaloides. Two or more genera of ants were observed on the involucres of P. cym- biferus at Tehuacan (Dressler 1368; unfortunately the col- lected specimens found their way into the museum without their identity having been recorded). Pedilanthus nodiflorus, berculatum Olivier were collected on the involucres of this species near Progreso, Yucatan (on Dressler 1362), and a fifth ant, seen but not collected, appeared to be a Pseudomyr- mex. The involucres, especially of P. calcaratus, are fre- quently chewed between and at the bases of the spur lobes or at the spur apex, indicating that short-tongued and chew- ing insects often seek the protected nectar. An infectious disease of Pedilanthus has appeared in the author’s greenhouse collections. This is manifested as a localized stem-rot in P. tithymaloides ssp. Smallii and causes severe damage to plants of that taxon. Similar but less severe symptoms occur in other forms of P. tithymaloides, P. bracteatus and P. macrocarpus. A deformation of young leaves, which may be due to the same infection, has been noted in P. calearatus and P. bracteatus. Nothing is known of the origin or causative agent of this infection. POLLINATION Although direct evidence is scanty, Pedilanthus is general- ly considered to be a hummingbird “flower.”’ Porsch (1923) discusses this at some length, and cites the observation by Cammerloher of an Old World honeybird visiting cultivated P. bracteatus. Porsch gives the color (red in either brac or involucre, of nearly all species), lack of scent, abundance of nectar, lack of “Sitzfliche” or landing surface for insects, and the strength of the spur lobes (only a relatively large animal could insert its mouthparts into the spur), as eVl- dence that Pedilanthus is adapted for bird pollination. It has, in recent years, been shown that hummingbirds seem to have no “preference” for red over other colors (Wagner, 1946; Bené, 1947), in spite of the observations of Porsch, THE GENUS PEDILANTHUS (EUPHORBIACEAE) 69 Pickens (1951) and others that red is the predominant color among bird flowers. The two groups of data are by no means in conflict. Red is complementary to green, contrasts strong- ly with the green background of vegetation, and should give a strong selective advantage in a species pollinated by an organism capable of detecting the red wave lengths, even though the pollinating agent may have no true color pref- erence. There are now available two records of hummingbirds visiting Pedilanthus under natural conditions. George R. Proctor has noted (Proctor 3418, in collection data) that P. tithymaloides is visited by hummingbirds on Isla de Provi- dencia (in the Caribbean, ca. 270 km. east of Nicaragua). The only hummingbird reported from this island is Anthra- cothorax prevostii hendersoni (Cory), and is said to be com- mon there (Bond, 1950). Ivan M. Johnston has observed hummingbirds visiting P. macrocarpus in Baja California (personal communication). Even with these two records, the greater part of the evidence is morphological and cireum- stantial. Pedilanthus fails to set seed in greenhouse culture unless hand pollinated, and it is difficult to see how any organisms other than birds (or possibly hawk moths) could account for the high percentage of seed-set in nature. Bees and other fairly large insects might profitably visit the involucres when the gland-chambers overflow, or bite into the spur, but pollination would only occasionally be effected. Hummingbirds, on the other hand, would have to approach the involucre from the front to insert their beaks into the gland-chamber, thus assuring that either the chin or the lower mandible would touch the stigmas or anthers, which Successively occupy the same position relative to the in- volucre. For most species of the genus it is a reasonable, if not inescapable, hypothesis that they are pollinated by birds and that this relationship has strongly oriented the evolution of the genus. The poorly known P. tehuacanus is an exception to the generalities of size, color and form which hold for the re- mainder of the genus. The bracts and involucre are green, the spur lobes do not snugly enclose the gland-chamber, the style is short and bent back toward the gland-chamber, and the staminate flowers are only shortly exserted. It seems most likely that it is pollinated by short-tongued insects. 70 ROBERT L. DRESSLER The structure of P. nodiflorus is somewhat puzzling as to its adaptive significance. Its form is such that it could be quite efficiently pollinated by hummingbirds, but the small, exposed lateral glands would seem to have no direct part in this relationship. They might possible serve to attract short- tongued insects, such as bees and flies (or to keep them from biting through the spur), that could provide a secondary and less dependable means of pollination, which might nevertheless have some selective advantage. The small ants that have been observed on the cyathia are not likely to be effective in pollination. It would seem that such small in- sects could scarcely serve any function for the plant species. GENERIC RELATIONSHIPS From its first botanical recognition, Pedilanthus has been considered a close ally, if not a member, of the Linnaean genus Euphorbia, but its relationships within that large and complex assemblage have received little consideration. There is no evidence to support the imaginative hypothesis of Croizat (1940-1942), which would derive the Euphorbieae from Dalechampia or a similar type, through Pedilanthus. Since a convenient summary of the Euphorbieae is not avail- able, it may be useful to present such a summary here, in so far as this is presently possible. SYNOPSIS OF THE EUPHORBIEAE 1. Dichostemma Pierre: Eihee: epoven? 1 sp.; woody, involucre 4- merous, glands large, between the staminate flower clusters, bracts ithin the involucre well Seca and aimee both staminate and pistillate flowers calyculate, cyathia often unisexual. 2. Anthostema A. Jussieu: West Africa and Madagascar, ca. 3S similar to no. 1, but involucre open on one side (hewaih a giant), isexual. 3 pease illauminia Croizat: New Caledonia, 1 sp.; woody, involucre 5-merous, involucral lobes enlarged and petaloid, glands large, in pairs between the staminate flower estar , bracts within involucre bi developed and distinct, pistillate flower caly culate. . Calycopeplus Planchon: Australia, 3-4 sp.; ephedroid shrubs es small, caducous leaves, involucre small, mostly ihe glands between and slightly within involucral lobes, bracts within involucre moderately well developed, pistillate flower calyculate. 5. Euphorbia L.: Cosmopolitan, A poorly developed in cooler a over 1,000 species; woody, succulent or eiaetet s, involucre ually 5-merous, glands between ieolneral lobes: usually distinct, THE GENUS PEDILANTHUS (EUPHORBIACEAE) 71 bracts within involucre (bracteoles) reduced, pistillate calyx small or absent. The numbers of species given for the subgenera (which a. subgenus Esula Pers.: largely Old World, ca. 375 spp.; diverse woody and herbaceous species without petaloid appendages on the glands, net iiate calyx sometimes present; includes the most primitive members of the genus (section Balsamis Webb. and Berthelot and § Launfoliag B oiss. ) 5b. subgenus pe ieee (Boiss.) L. C. Wheeler: African shrubs, ca. 10 spp 5c. subgenus Eremophyton (Boiss.) L. C. Wheeler: Old World shrubs or herbs, ca. 10 spp. 5d. subgenus Euphorbia: Old World succulents, ca. 275 spp., a polyphyletic group of at least ts Med ane 5e. subgenus Agaloma (Raf.) H : American, ca. 130 spp.; woody, cachaonitn and a few succulent sey ys ide with petaloid appendages, these sometimes reduced, pistillate calyx sometimes present. 5f. subgenus Chamaesyce Raf.: Cosmopolitan, probably of American origin, ca. 220 spp.; a characteristic sympodial habit of growth in most species, glands with petaloid appendages, or these vcaiieed: close- ly related to subgenus A galoma 5g. subgenus Poinsettia (Grah. ) House: American, 8-10 spp.; woody or orhasabal a aeA cup-like, often only one on each ht i diol re- — to dr es oma. Mebehertia Stapf: Africa, ca. 4 spp.; woody or semi-succulent docs ovary sessile and fruit a drupe; similar to some members of Euphorbia, subgenus Suihevtin (sensu s 7. Sy nium Boissier: Africa, ca. 15 SP woody or semi-succulent shrubs, involucral glands united into a 8. M ium Pax: Africa, ca. 30 “SPP; ; succulent or subherba- ceous, glands united into an incomplete ring, which overtops the in- volucral lobes; apparently related to Synadenium. 9. Cubanthus (Boiss.) Millsp.: West Indies, 5 s woody, glands two, ntited into a _ “like By ahintigge patie Acie calyculate. Pedilanthus Nec rican, oody or succulent, — appendaged, the pee sip Hon deg ity ai involucral lobes unequal. The petaloid appendages of the glands (spur lobes) of Pedilanthus, alone, clearly indicate that the Euphorbia an- cestor of the genus would, if living, be assigned to the Subgenus Agaloma. The woody habit of Pedilanthus, gland- like stipules, ecarunculate seeds (when mature), and small chromosomes are further characters shared with Euphorbia Subgenus Agaloma. These features are also shared, in part, with the subgenera Poinsettia and Chamaesyce, but these subgenera could not, on morphological grounds, be as closely related or ancestral to Pedilanthus. The features which are clearly primitive or universal in Pedilanthus and might be 72 ROBERT L. DRESSLER expected in its immediate ancestors are: woody habit; al- ternate, and possibly distichous, leaves; small, conic, gland- like stipules; dichotomous, terminal or upper axillary, inflorescence ; four red petaloid appendages on the involucre; primitive pollen with “euphorbioid” pattern of intine thick- enings and moderately thick, reticulate exine. These features are not to be found in combination in any living Euphorbia, but all may be found in different species of the subgenus Agaloma. TABLE 7. A comparison of Pedilanthus and Cubanthus. 16, 17, 18. Pedilanthus Cubanthus 1. Vessels never strictly scalari- 1. Vessels strictly print form pitted with slit-like pits. pitted with slit-like 2. Leaves distichous and distant. 2. Leaves closely intel 3. Stipules small, gland-like. 3. madness none. 4. Cymes co aamrionads terminal 4. Cy pseudo - umbellate, or axillar doves coated at stem apex. 5. Involucral lobes unequal, 5. Involucral lobes equal, com- partly free pletely connate. 6. Glands bas to six, free, ap- 6. Glands two, completely con- pendaged and concealed by nate into a shield-like ges the appendages (spur). ture, without appendag 7. Pollen: Intine thickenings 7. Intine thickenings peat be- closer Sige ei furrows than neath furrows than between beneath furrows; furrows furrows: furrows narrow, not opening Widely when grains opening widely when grains are expanded, exposing pores. are expanded. 8. Pistillate flower naked 8. Pistillate flower with a small calyx. 9. Haploid chromosome numbers 9. Haploid chromosome number 20. Certain infrequently collected West Indian plants with spiral leaves, urn-shaped involucres, shield-like glands, and other distinctive features have been placed in Pedilanthus, as members of the section Cubanthus Boissier. Millspaugh elevated this group to generic rank in 1913, and a careful examination of the material shows this action to be well justified. The members of Cubanthus are not directly related to Pedilanthus, but have been derived from Euphorbia subgenus Esula (§ Laurifoliae), mainly through the con- nation of the non-appendaged glands. Relatively closely related species of Euphorbia, especially E. gymnonota Ur- THE GENUS PEDILANTHUS (EUPHORBIACEAE) 73 ban, are to be found in the West Indies. A partial comparison of Pedilanthus and Cubanthus is given in Table 7. In most of the features enumerated, Cubanthus agrees closely with the woody Antillean members of Euphorbia subgenus Fsula. SPECIES RELATIONSHIPS The species of Pedilanthus fall into five fairly discrete species groups. These groups and their affinities are diagram- matically shown in figure 26. Some of the characters that vary within the genus and their specific distribution are shown in Table Palmeri group (species 1-2). These two species are closely allied by habit, inflorescence, prominent lateral spur- lobes, and involucral texture. They are very similar in most a” 2 . . @ od oo Wh bed wo oe. ww Qes3s ea 3 ade - aie) — 2 SS 5 O° ae 5s = 2a Co® ecg = E — cae = £ x o °V o- 2 ao 2s rg eee i i ee) Bo ges Sea = o oa so oo o = a. oo ge Sg ess ey gee e= Species relationships in een The reticulate relationships between Fig. 26. the species groups are shown by dotted | 74 ROBERT L. DRESSLER features, but strikingly different in spur form. The group seems to be allied with the next group, because of broad spur lobes, habital and seed resemblances, and some degree of interfertility (see under Hybridization), but shows no close resemblances to the other groups. . P. calearatus group (species 3-4). These species are obviously quite closely related, and, as now known, differ mainly in proportion and aspect. They are related to the above group and to the P. cymbiferus group on the basis of seed and internal indument of the involucre. The keeled spur lobe suture, the thickened spur lobes and the aspect of the involucre suggest a less close relationship to the P. bracteatus group, especially to P. macrocarpus 3. P. cymbiferus group (species 5-7). The form and struc- ture of the involucre, as well as the non-elongating pistillate pedicel (of 2 species), and anatomical features tie these species into a very “natural” group. Aside from the resem- blances to the P. calearatus group (above), this assemblage shows affinity with the P. bracteatus group (especially through P. coalcomanensis) and to the P. tithymaloides group (especially through P. pulchellus). Features indicat- ing an alliance with the first group are: wide cyme bracts, the branching of the cymes (nearly in one plane), the pilose staminate flowers, the texture and indument of the plant (these features approach P. bracteatus especially), and the spur partitions (like those of P. macrocarpus). The non-elongating pistillate pedicel, partial spur partitions, and truncate spurs sometimes found in P. tithymaloides, the indument (external and internal) of the P. nodiflorus m- volucre, as well as less definable textural resemblances, seem to indicate a relationship between the P. cymbiferus and P. tithymaloides groups. 4. P. bracteatus group (species 8-11). Pedilanthus brac- teatus, P. tomentellus and P. tehuacanus are closely allied by resemblances of the plant, cyme, involucre and seed, as well as the unique yellow latex. If only the Sonoran and Sinaloan populations of P. macrocarpus were known, it probably would be placed in a separate species “group” of puzzling affinity. The peripheral, and presumably primitive, popula- tions of Baja California closely resemble P. bracteatus In nearly all features. The apparent interfertility between the two species (see Hybridization) confirms this relationship. THE GENUS PEDILANTHUS (EUPHORBIACEAE) 75 ABLE 8. The distribution of certain features in Pedilanthus. x, present throughout the species; -, present in part of the species. Hey oe g 4 - 3 cee DS gae Bpe mn — o gis 330 843 eo ae o 2 $y 2s7 soSgg ts~e 8s SaBea 2a as ae 5 eolpdcas P38 Sats SRLS SLE PPR SESS Ene Fy 95 3 SYISsS suas > tana pam ATanmnwmodgasn _ = sons e122 2S SS so Sy ene by BB eee BREE SSSS SSS Ee SEER EREE ESS § BRPL SERRE SSRSSRE EIRBTAES P. Palmeri Se cies 5 x x's P. Finkti 5 GEE ene. x P. calcaratus oe ee x x > o que P. gracilis Tots = x x xxx x P. pulche xox x. oe oe xx P. coaleomanensis eee sip Gee Cli See x »& P. cymbiferus x x x xX oe - P, carpus Swe x Sie P. bracteatus > dy Gia a> aie eee ae 4 Etc ux P. tomentellus ree Ses - x x P. tehwacanus x x = es —- x x P. nodiflorus x" £2 7h" ERR x P. tithymaloides joa ere tox OX --- The moderate resemblance between P. tehuacanus and P. nodiflorus is thought to be superficial, resulting from the parallel development of very short spurs and reduced leaves ; in most characters they are quite different. There seems, however, to be a distinct affinity between the two species groups, especially because of the similarly formed accessory involucral lobes. These two species groups also share stem succulence, similar anatomical features (especially in the wood) and the same chromosome number. 5. P. tithymaloides group (species 12-13). This group is apparently related to groups 3 and 4, as discussed above. It cannot be definitely shown that these two species are properly placed together. The involucres, however, are very similar in form, and anatomical, cytological and other observed features do not contradict this relationship. HYBRIDIZATION Only a limited number of cross-pollinations have been attempted with the greenhouse cultures of Pedilanthus. Dif- 76 ROBERT L. DRESSLER ferences in flowering time and the small number of cyathia produced have limited the possibilities. The genus is further unsatisfactory for experimental hybridization in that only three ovules can develop from one pollination. Self-pollina- tion and cross-pollination within a species usually meet with easy success. Pedilanthus tithymaloides has given somewhat erratic results and little can be said concerning its fertility relationships without more material and further tests. At present, one successful cross within a species group and one cross between closely allied species groups have been made in the greenhouse, and a putative hybrid between the same two species groups is represented by very poor herbarium material. Crosses which have been attempted without success are: P. Palmeri x bracteatus, P. calcaratus x bracteatus, P. calcaratus x tithymaloides, and P. nodiflorus x tithymaloides. In each case, only very few pollinations have been attempted (with reciprocal pollination, when possible), and these nega- tive results cannot be given much weight. Known and putative hybrids are discussed below. 1. P. Palmeri x calcaratus. Pedilanthus Palmeri (Dressler 1064) set seed readily when pollinated with pollen of P. calearatus (Dressler 1067), though the reciprocal failed. The seeds germinated well and four seedlings have now completed two seasons of normal growth. The leaves of the seedlings have a soft pubescence on the underside, which is lacking in the maternal parent, but are not otherwise markedly dif- ferent from those of either parent. Two of the seedlings have recently flowered, and the proportions of the involucres are quite intermediate between those of the parent types, leaving no doubt of their hybrid nature. Even the position of the cyathium is intermediate: the peduncles twist about 90° so that the spur projects horizontally. Microscopic exam!- nation shows about 45 percent of the pollen to be abortive. No crosses involving these hybrids have yet been attempted. 2. P. macrocarpus x bracteatus. A single cyathium of P. macrocarpus (Dressler 1090) was pollinated with pollen of P. bracteatus (Dressler 978). All three ovules developed and the seed germinated normally, the cotyledons being inter- mediate in width between the parent types. Two of the seedlings were lost through damping-off and slug damage. The remaining plant has grown for five years, and showed signs of flower initiation in the fall of 1956, though the THE GENUS PEDILANTHUS (EUPHORBIACEAE) Th primordia all aborted. Its features are clearly intermediate between the parent species. The plant is of much slower growth than P. bracteatus, but Se far outgrown P. macro- carpus seedlings of greater age. The leaves are flat and narrowly ovate, about 1-1.8 cm. wide and 2-3.5 cm. long; even the stem color is intermediate between the parent types. Several leaves beneath each developing inflorescence are small and bract-like, about 1 cm. long. 3. P. calearatus x Finkii? Among the several sheets of Liebmann 5721 (all at C), all badly decomposed and dam- aged by insects, there are numerous leaves (mostly unat- tached) and a few involucres of typical P. calcaratus. In a packet of one of the sheets, however, there is an involucre that differs markedly from the known species of Pedilanthus and suggests either an interspecific hybrid or an undescribed Species. It is tentatively considered a hybrid between P. calearatus and P. Finkii. A description, as far as one can be written for such a damaged specimen, is given herewith: Involucre glabrate without, thick in texture like that of P. calcaratus, tube ca. 11 mm. long, tomentulose within, main lobes obtuse, accessory lobes ca. 1.2 mm. wide, obtuse ; spur arising ca. 1.5 mm. from peduncle, bending abruptly forward at ca. 2 mm. from tube and sub-parallel with the tube, terminal (sub-parallel) portion flattened, 5.5 mm. long, 7 mm. wide, broadly rounded, spur lobes arising beneath the flattened portion, lance-ligulate, ca. 3 and 4 mm. long (un- equal), ca. 2mm. wide basally and 0.7-0.8 mm. wide terminal- ly, obliquely truncate and somewhat folded and thickened terminally, glands four, 2 lateral and 2 folded about short terminal partitions; anther lobes slightly convergent ter- Fic. 27. An involucre from Liebmann 5721, which is thought to represent a hybrid between P. calcaratus and P. Finkii (the remainder of the collection is P. calear tus) This involucre has been badly damaged by insects, and the sketch is a reconstruction. Note the long, free spur lobes (arrow). Magnification ca. 4. 78 ROBERT L. DRESSLER minally (similar to P. Finkii), pollen averaging 76.6 microns in polar dimension and 93.8 in equatorial diameter ; pistillate pedicel glabrous, ca. 8 mm. long, ovary ovoid, ca. 2 mm. long, style 5.5 mm. long; branches 1 mm. long, spreading, suben- tire. The bent and flattened spur, as well as the appearance of the anthers and style, point strongly to the distinctive P. Finkii, while the texture, the folding and thickening of the spur lobes, and the associated materials suggest P. cal- caratus. These and the very different P. tithymaloides are the only species known from Veracruz. The pollen from the two half-eaten anthers does not show the distinctive thick- ened furrow-margins of P. Finkii, but does have relatively narrow pores and a sub-triangular equatorial outline sug- gestive of that species. The pollen neither confirms nor contradicts its hybrid nature, but there is not a high per- centage of abortive pollen. The elongate, narrow spur lobes of this involucre are of special interest, since free, narrow spur lobes are not a feature of either putative parent. It 1s not impossible that these species are descended from types with narrow spur lobes like those of the P. cymbiferus group, and that hybridization in this case has broken down the modifying and inhibiting systems present in each parent species. Until material of P. Finkii is again available in cultivation, the nature of this interesting fragment cannot be experimentally checked. The putative inter-subspecific hybrids of P. tithymaloides, which have been observed, are described with that species. There is some evidence of hybridization between P. t. ssp. tithymaloides and the subspp. Smallii and parasiticus, where cultivation has brought them together. Intermediates be- tween P. t. ssp. angustifolius and P. t. ssp. padifolius are also known, but there is no indication of hybridization between P. t. ssp. angustifolius or its immediate allies and subspp. tithymaloides or parasiticus. ORIGIN OF THE GENUS The genus Pedilanthus probably arose in tropical Mexico, possibly on the Pacific slopes. It clearly developed from a member of Euphorbia subgenus Agaloma, a woody. plant whose involucres bore four appendaged glands, the ap- pendages probably red. The differentiation of the distinctive THE GENUS PEDILANTHUS (EUPHORBIACEAE) 13 features of the genus must not have commenced until there were hummingbirds in tropical North America. This, un- fortunately, cannot be dated. These distinctive South Ameri- can birds have probably occupied their present ecological niche nearly or quite throughout the Tertiary. Wagner (1946) has suggested that the hummingbirds did not reach North America until the late Tertiary, but the Central American water gap can scarcely have confined them (Mayr, 1946), and we cannot agree that all the ornithophilous plants of Mexico are, without exception, recent immigrants from South America. More suggestive is the distribution of the Euphorbieae. The truly primitive genera of this tribe, with markedly calyculate flowers, are all found in the Old World, where they occur largely in peripheral areas: New Caledonia, Australia, Madagascar and western tropical Africa. The most primitive groups of the genus Euphorbia, the woody members of the subgenus EHsula, also show relict, discon- tinuous distribution. There are in the New World several species in the West Indies and Middle America, but the few South American species are strictly Andean and rather closely related to the Middle American species, suggesting that they are recent invaders of South America (a few more advanced types of Euphorbia doubtless reached South Ameri- ca in the Tertiary, by “island hopping” from North America, and perhaps elsewhere). As a whole, the genus is poorly developed in South America. This distributional pattern is taken to indicate that the Euphorbieae invaded North Ameri- ca in the early Tertiary, when North and South America were already isolated. These two lines of evidence serve only to suggest that Pedilanthus has developed since the beginning of the Tertiary. One cannot be more exact. The ancestral Euphorbia was probably a local, not too successful sort, as a Euphorbia. Its closest approach, as an adaptive type (but taxonomically very distinct), EF. fulgens, appears to be quite local, and no other close counterpart 1s found in the modern flora. The pattern by which this Euphorbia became a Pedilanthus can be rather readily visual'zed ; the author’s ideas of this pattern are sketched in fig. 28. The cyathia of Euphorbia are often slightly zygomor- phic, in the lack of a fifth gland and the recurving of the pistillate pedicel to one side, through the gap between the petaloid appendages. This slight degree of zygomorphy is 80 ROBERT L. DRESSLER apparently the basis from which the spurred, bent involucre of Pedilanthus has developed. The stigmas are receptive to pollen and fertilization is still possible while the pistillate pedicel recurves. A tendency for the pistillate flower to remain partly deflexed, coupled with a tendency for the gland appendages to remain partly folded (fig. 28b), as they do in E. lancifolia and some of its allies, would be enough to convert an appropriate Euphorbia into a hummingbird “flower,” though an inefficient one. Once this stage was reached, it is easy to see that selection pressure toward the further stages shown in fig. 28 would be high. A further folding (or rather an inhibition of the unfolding) of the gland appendages would narrow the possible approach of the pollinating agent, and a lateral deformation leading to a spur and a bent, elongated involucral tube would require the bird to probe more deeply, both greatly improving the efficiency of the now zygomorphic flower cluster. This hypo- thetical pattern of evolution would lead to a type with an involucre of much the same general proportions as P. brac- teatus, and all of the modern involucres could be readily derived from such a type. The intermediate types shown in Fic. 28. Hypothetical derivatio above, top view; below, lateral A intermediate in which bilateral deformation has started to form an involucral tube anteriorly and a spur posteriorly. D. A generalized Pedilanthus. THE GENUS PEDILANTHUS (EUPHORBIACEAE) 81 fig. 28b and ¢ would seem to be in an “adaptive valley” (Simpson, 1953): inefficient either as insect or humming- bird “flowers.” As soon, however, as 2 moderate degree of zygomorphy was achieved, the species would be on the slope of a new “adaptive peak,” and selection pressure would be especially great, with a correspondingly rapid change toward the new peak. PRIMITIVE AND ADVANCED FEATURES No one species can be picked out as the primitive species of the genus. Pedilanthus has almost certainly arisen from Euphorbia subgenus Agaloma; the unique inflorescence of the tribe can only be derived through reduction from a cer- tain type, and the evolutionary pattern of certain anatomical features is especially well known (Bailey, 1944). For these reasons, we can, with some certainty, enumerate primitive and advanced features for the genus (Table 9), but these me primitive and advanced features in the genus Pedilanthus. Some of the preety ri pam are also primitive for the ancestral genus, Euphorbia Primitive Advanced ia succulent or 1. Tree or large shrub, woody. 1. Small subherbaceo 2. Vessels narrow 2. Vessels wi 3. Many solitary vessels. 3. Most vessels = distinct ra- dial multip 4. Vessels mostly scalariform 4, Vesse mei Paya or itt alternate pittin 5. Rays _multiseriate, hetero- 5. Rays Saleueate. homogen- geneo eous. 6. Lata 6. Latex & -f Eaaven ¥ races large (pos- 7. Leaves small, caducous. sibly evergreen 8. Cyme bracts of moderate or 8. Cyme bracts large. small size. 9 9. Cyathia solid red or solid oO green. 10. Numerous staminate flowers. 10. Few staminate flowers 11. Well developed paler ae 11. Bracteoles small or few. 12. Spur and i ral 12. Spur sessile or very elongate, ] both moderately Son caied or involucral tube greatly . boty green, with red spur elongate. Medial spur lobes connate. 13. Free medial spur lobes. 13, 14. Glands 4. 14. Glan . 15. Pollen colporate. 15. Pollen colpate or inapertur- ate. 82 ROBERT L. DRESSLER features show a checker-board distribution among the species (Table 10). Pedilanthus tehuacanus deviates most widely from the hypothetical primitive type; P. tithymaloides, in spite of its involucral specializations, has a large number of primitive features, especially in wood structure. Most of the features in Table 9 are self-explanatory. In keeping with the evolutionary pattern given above for the genus, an involucre with both spur and involucral tube moderately elongated is taken as the primitive type. It would be possible to think of the primary differentiation of the genus as effect- ing only one of these structures, but such a pattern is less easily visualized and is not supported by the modern varia- tion pattern of the genus. Wide, blunt spur lobes and proportionately wide accessory involucral lobes should both be primitive features for the genus. There is doubt, however, that they are primitive where now found. The species with wide spur lobes may very well be secondarily derived from ancestors with more narrow ones( see P. Finkii x calcara- tus?, under Hybridization). The proportionately wide acces- sory involucral lobes, in some species of the P. bracteatus and P. tithymaloides species groups, are thought to be the result of a certain degree of neoteny. TABLE 10. The distribution of some primitive features in Pedi- lanthus. The numbering of features follows that in Table 9. An asterisk indicates that the feature checked is present in the species, but not necessarily in all plants. 1.2 S$ 42 5 6-7: 8 8:30 11 19 13 1616 oe P. Palmeri * * * * * & * xe TO P. Finkii * * * * * * 6 P. calearatus . eee x * * * 9 P. gracilis * * *# * x *& # * 9 P. pulchellus ——* x ok & 2 +s & 8 8 10 . coal- comanensis * * * * *« * * * 8 3g cymbiferus besa * * #8 uke 9 P. macrocarpus * * & * * * * * * 9 P. bracteatus * oe 4 * * * * 7 P. tomentellus * * * x * * * 7 P. tehuacanus * * * x * 5 P. nodiflorus * * * * x « eRe Be P. tithymaloides ie, eet oe ee ee Gee eee ee ee || THE GENUS PEDILANTHUS (EUPHORBIACEAE) 83 TRENDS IN THE INVOLUCRE The distinctive features of the genus are to be found in the involucre, and in this structure are to be found some of the most interesting modifications of the genus, if not of the tribe. Striking variation is to be seen in the relative develop- ment of the spur and involucral tube. In P. macrocarpus and P. Palmeri the involucral tube is relatively short, while the spur forms the greater part of the inflorescence. Similarly, the spur of P. Finkii overtops the involucral tube in size, though its form is very different. In P. nodiflorus and P. tithymaloides, on the other hand, the spur elongates only slightly and the involucral tube comprises the greater part of the cyathium. Pedilanthus tehuacanus has a very short spur, but here the development of the involucral tube is also arrested. One of the themes of the adaptation of the involucre, or rather of the entire inflorescence, has been that of “Schau- apparat,” or the device to heighten the visibility of the in- florescence. This has been accomplished in various ways. Pedilanthus coaleomanensis, P. bracteatus and P. tomentellus have large, colored cyme bracts; in the last two species, indeed, the involucre itself is green. In most species the involucre, and especially the spur, is the most conspicuous feature. In P. macrocarpus and P. cymbiferus, leafless small- bracted desert species, the spur is widened and flattened to provide a more efficient “sign-board.” That this is the main adaptive significance is especially evident in P. macrocarpus, for the flattened terminal part of the spur is solid and parenchymatous. Pedilanthus nodiflorus and P. tithyma- loides have short spurs and relatively small involucres, as Well as small inconspicuous cyme bracts. In these species, however, numerous cyathia are produced and these are borne in relatively compact clusters, so that the total inflorescence may be quite as showy as a few larger cyathia. Quite aside from size and flattening of the spur, there is great diversity in the form of the spur, the spur lobes and the glands. With- out better information on the pollinating agents it would be futile to speculate further on the adaptive significance of these variations. While the primary features of the genus result from an elaboration of the later stages of inflorescence ontogeny, the opposite trend also occurs. At least two groups of popula- 84 ROBERT L. DRESSLER tions seem to owe their distinctive features largely to ‘“‘paedomorphosis,” or neoteny (see deBeer, 1954), of the cyathium. Pedilanthus tehuacanus and P. tithymaloides ssp. angustifolius differ from their respective allies by features which are best explained as arrested development of the entire involucre, coupled with normal maturation of the anthers, ovules and stigmatic tissues. The aspect of the P. tehuacanus involucre is sometimes approximated by in- volucres borne on cuttings of P. bracteatus when these fail to develop normally. The partial connation of the cyme bracts and the sessile involucre of this species also are best explained as arrested development. The pistillate flower, with its short, blunt style, looks distinctly “immature” in both P. tehuacanus and P. tithymaloides ssp. angustifolius. The short spurs of P. nodiflorus and P. tithymaloides, too, may be explained as a paedomorphic, or partly allometric, pattern. The young involucre of P. calcaratus, P. cymbiferus (see the first stages in fig. 18) and similar species is quite similar to the mature involucre of P. tithymaloides. If spur development be greatly arrested, relative to the development of the involucral tube, in any of the moderately long-spurred species, something very similar in proportion to P. tithy- maloides would result. XERIC ADAPTATION One of the striking features of Pedilanthus, as of the entire tribe Euphorbieae, is the adaptation of several lines for survival in dry habitats. The ancestral “proto-Pedilan- thus’? may have been adapted to a seasonally dry habitat, or it may have occupied a rather mesic niche. In any case, it possessed the morphological and physiological features which predispose the Euphorbieae to successful evolution as desert plants. The woody P. cymbiferus seems to have adapted to a truly desert habitat quite independently of the other xerophytes of the genus. The P. bracteatus and P. tithyma- loides species groups both show true stem succulence, and this feature may have arisen only once within the genus. Essentially leafless xerophytes, however, have evolved sev- eral times among these few species. Both P. macrocarpus and P. tehuacanus appear to have evolved from a large- leaved ancestor similar to P. bracteatus. There are large- leaved populations of P. tithymaloides, which suggests that THE GENUS PEDILANTHUS (EUPHORBIACEAE) 85 its ally, P. nodiflorus, has taken up the microphyllous habit quite independently. Small or caducous leaved populations are to be found in P. tithymaloides in Oaxaca, Mexico, in Venezuela, and in the Greater Antilles. The patterns of variation within the genus suggest that there may have been two or more ways in which the small- leaved xerophytic habit was achieved. In Pedilanthus nodiflorus and P. tithymaloides subsp. angustifolius and its close allies, leaf reduction has involved primarily reduction in width, resulting in a narrow, more or less lanceolate leaf form. In some other populations of P. tithymaloides, how- ever, there is a certain degree of foliar dimorphism, suggest- ing a different pattern for the evolution of reduced leaf size. In P. tithymaloides ssp. parasiticus and some populations of P. t. ssp. tithymaloides (Tomellin Canyon, see Plate XXI) there is a strong tendency for the upper branches to develop several to many small bract-like leaves, especially beneath the inflorescence. Such a pattern is not unusual among plants with distinctive bracts, and may suggest a pattern for the evolution of microphylly in a number of different xerophytes. These populations with a tendency towards dimorphic leaves are thought to represent a change in the threshold for some of the features of bract development, so that the upper leaves develop along a pattern intermediate between bracts and foliage leaves. In an isolated arid habitat selection for small- leaved variants might shift this threshold so that a greater proportion of the leaves develop on the ‘‘semi-bract” pattern, and with continued selection the normal foliage pattern might be entirely lost. Partial support for this interpreta- tion is provided by the hybrid between P. macrocarpus and P. bracteatus; the first parent is a plant with only very small bract-like leaves, and the second a large-leaved plant with little or no reduction in the size of the upper leaves. The hybrid has several small ovate, bract-like leaves beneath the inflorescence; leaves very like the wide leaves of primitive P. macrocarpus (the parent plant of this species is an ad- vanced type from Sinaloa, with nearly linear leaves). Pedilanthus cymbiferus and P. macrocarpus are species which may have evolved through such a pattern of foliar dimorphism. The populations of P. tithymaloides which show foliar dimorphism are probably prevented from further progress in this line by gene exchange with populations of 86 ROBERT L. DRESSLER less xeric habitats. Isolation would seem to be necessary for successful adaptation to an extreme habitat. A certain degree of genetic correlation between leaf and bract is evident. All of the truly microphyllous xerophytes have very small bracts. Pedilanthus tehuacanus is a partial exception in having relatively large bracts and relatively small leaves, but its leaves are markedly larger than those of the other xerophytes. With the involucres tightly enclosed by the bracts at anthesis, a further reduction in the bracts would be quite impossible. Selection for smaller leaves and selection for the same or larger bract size may be an uneasy balance in this species. Further reduction in leaf size will require mutations which do not affect bract size or ones which increase bract size relative to leaf size. ADAPTIVE RADIATION Once the new adaptive type of a relatively efficient hum- mingbird “flower” was achieved by the ancestral Pedilan- thus, something very like adaptive radiation occurred, and seems still to be occurring. This pattern, the relatively rapid (“tachytelic”) evolution of a population (probably small) into a new adaptive type, followed by adaptive radiation in the new niche, is a well recognized phenomenon among anl- mal groups, and is often connected with the origin of new families and other higher categories (Simpson, 1953). It may be argued that, as to pollination relationships, Pedilan- thus (except for P. tehuacanus) occupies only one niche, and the scarcity of truly sympatric species pairs tends to support this. Nevertheless, the genus has adapted to that niche In diverse ways (see above, under Trends in the Involucre). Simpson and others have rightly emphasized the primary role of environment and selection in adaptive radiation. Certain features of the genetic systems may, however, con- tribute to the rapid radiation of “new” groups. It sometimes appears that a new structure or developmental pattern 1S especially subject to variation in its phylogenetic youth, and such may, to a degree, be true. A strong selection pressure is required to develop a new adaptive type from a gene com- plex already well balanced about a different adaptive peak. With time the new pattern will become strongly “buffered” by the development of polygenic systems which affect its THE GENUS PEDILANTHUS (EUPHORBIACEAE) 87 ontogeny, but, until this buffering has occurred, mutations affecting the new structure will have a high degree of penetrance. According to this concept, a structure or de- velopmental pattern might, indeed, have a stage of “youth,” when it is not more mutable than in later stages, but when individual mutations may have a more pronounced effect and be more readily acted upon by selection. The occurrence of Pedilanthus in a variety of habitats may also be considered a sort of adaptive radiation. This, however, has largely involved the evolution of similar xerophytic adaptations in several lines. Such parallelism in- volves no mystic “orthogenesis.’’ When a species is already adapted to withstand some degree of drought, it will usually be found in a habitat where selection favors the evolution of further xeric adaptation. PARALLELISM The interaction of similar environments and similar gene complexes often results in parallel patterns of evolution. This has been the case in several instances in the present genus. Probably the most striking case of parallelism is to be found in P. cymbiferus and P. macrocarpus. Both are grey-green, essentially leafless, clump-forming desert shrubs with small, caducous bracts and large, bright red involucres with wide conspicuous spurs. They are strikingly similar in aspect, yet they do not seem to be closely related: one is woody, the other succulent; one spreads by under- ground stems, the other by forming adventitious buds on the roots; the details of wood structure, leaf, involucre, fruit and seed are all markedly different. Their similarities seem clearly to result from the development of parallel features in similar habitats. Pedilanthus nodiflorus and P. tithyma- loides ssp. angustifolius provide a less striking case of parallelism. The two species are thought to be closely re- lated, yet the narrow leaves and proportional similarities of the involucre seem to have been derived quite independ- ently in the two taxa. A more general case of parallelism is the development of small or caducous leaves in dry habitats in about seven separate populations within the genus (three different centers of xeric adaptation being counted for P. tithymaloides). 88 ROBERT L. DRESSLER SOME SPECIFIC PATTERNS OF EVOLUTION Having considered some of the general aspects of evolu- tion in Pedilanthus, we will now consider three of the species in greater detail. 1. Pedilanthus macrocarpus. This Sonoran Desert species has apparently evolved from an ancestor very like the pres- ent day P. bracteatus. The common ancestor of these two species must have had white latex and retained red spur lobes, as well as the full complement of four glands, but in other features it may have differed very little from the modern P. bracteatus. The isolation of the ancestral P. macrocarpus population probably occurred somewhere on the present peninsula of Baja California. This isolation could have occurred as early as the middle Miocene, when the Gulf of California was being formed and the peninsula apparently became an archipelago (Gentry, 1949). This isolation could, of course, have occurred much more recently. In any case, the isolated population found itself in an arid or semi-arid habitat, and selection for reduced leaf and bract size and increased succulence was effective. In late Pliocene and Pleistocene the California Gulf region acquired its modern character, with some fluctuation in sea level during the Pleistocene, and large areas of arid coastal plain de- veloped. In these geologically recent times, P. macrocarpus has occupied much of the peninsula and invaded the main- land of Sonora, where it is quite successful in the cactus desert of the coastal plain and has spread southward into northern Sinaloa. These mainland populations have a nar- row, fleshy, almost sub-terete leaf very different from the ovate leaves of the peninsular populations (but leaves are known for very few collections from either area). In Sinaloa and southern Sonora there is clearly strong selection pres- sure for a larger, more readily visible spur and for more connate spur lobes. Whether this is due to a difference in pollinating agents or to other differences cannot be de- termined; the denser vegetation of the thorn scrub in this area might place a premium on larger, more visible spurs. There is a distinct cline in both involucre and fruit charac- ters, from the highly specialized populations of the southern mainland, running north along the coast and then south on the peninsula to the primitive bracteatus-like populations of THE GENUS PEDILANTHUS (EUPHORBIACEAE) 89 the southern peninsula. The population of San Pedro Nolasco Island does not fit in the cline, but is very similar to the Cape population. It is not yet clear whether the cline represents a gradient of selection pressures (with some gene exchange, of course) or a relatively efficient dispersal and gene exchange between the northern populations across the gulf. The primitive nature of the San Pedro Nolasco popu- lation casts doubt on trans-gulf gene exchange. A relatively recent continuity about the northern end of the gulf is con- ceivable, but the gulf extended much further to the north in truly recent times. It should be noted that the bracteatus- like population is, geographically, the farthest removed from the range of P. bracteatus. There is no evidence that the ranges of P. bracteatus and P. macrocarpus actually over- lap. Thus both introgression and character displacement Seem to be excluded as causes of the variation in P. macro- carpus. Pedilanthus macrocarpus has the appearance of a population complex which is now undergoing rapid and obvious evolution. An intensive study of this species would undoubtedly be rewarding. At the very least, it shows us how strikingly different species, species as different as the five species groups of Pedilanthus, can and do arise from a common ancestry. 2. Pedilanthus tehuacanus. This species is known only from fragmentary material, yet its features are so striking that it well deserves special attention. Pedilanthus tehua- canus has arisen from the same ancestral stock as the pre- ceding species, but not until this stock had acquired yellow latex and a tendency to reduce the red color of the involucre (perhaps also large cyme bracts, but the author is not pre- pared to state that the ancestor of P. macrocarpus did not also have large bracts like those of P. bracteatus). It is known only from the area of local aridity near Tehuacan, Puebla, a dry basin that may date, as such, from a late Tertiary orogeny. Whatever the original isolation, or its habitat, the population has become more markedly xerophy- tic than P. bracteatus, though this trend has not gone as far as in P. macrocarpus. It would appear that hummingbird pollination ceased to be adequate for the species; possibly because the birds were scarce in that habitat or because the trend to bract and leaf reduction had too greatly reduced the size of the bracts and the visibility of the inflorescence. 90 ROBERT L. DRESSLER With a return to small insects as the major agents of pollina- tion there was a very strong selection pressure for reduction in the size of the cyathium, so that the anthers and styles were closer to the nectaries. This was most easily achieved by neoteny, or a retardation of involucral development rela- tive to the flowers. This same retardation further increased its efficiency as an insect “flower,” by partial connation of the cyme bracts, which forces the insects to approach the involucre from one side, near the flowers. This effect has been further heightened by a shortening of the staminate flowers and a bending of the style so that both the stigma and the anthers are borne quite near the orifice of the bract hood. This one species, thus, has again left the adaptive type of its congeners and entered a new one. It is, to be sure, insect pollinated like most Euphorbias, but its structure is very different from that of any Euphorbia. 3. Pedilanthus tithymaloides. This species, the best known of the slipper spurges, occurs naturally over a wide area and in a variety of habitats. Its pattern of variation is especially interesting and informative, as to the origins and relationships of the West Indian populations. The exact geographic origin of such a widespread complex can rarely be determined, but distribution and occurrence of variabili- ty and primitive features suggest Central America as the center of dispersal for this species. The present discussion will deal largely with the Antillean populations, and little will be said of subspp. parasiticus and Smallii, except that they appear to have a common ancestry and to have reached the West Indies from Central America. The Floridian population of Smallii must be of very recent dispersal, as are most of the West Indian plants in the geologically recent tip of Florida. As noted previously, P. tithymaloides is thought to have reached South America in the late Pliocene, when the Tertiary water-gap between the divergent North and South American biotas was closed (Simpson, 1950; Mayr, 1946). Since the species can and does occasionally “island-hop,” it may be argued that the water-gap would not be a serious barrier. If dispersal antedated the land connec- tion, it probably did not occur until mid-Pliocene times, when the gap was being narrowed (had it occurred much earlier, one might expect specifically distinct populations). Once the species invaded South America, its dispersal along the THE GENUS PEDILANTHUS (EUPHORBIACEAE) 91 semi-arid north coast may have been relatively rapid. The history of P. tithymaloides in the West Indies, thus, would appear to concern the Pleistocene, the Recent and possibly the latest Pliocene. Since Pedilanthus has so successfully spread through the West Indies, one may inquire as to its means of dispersal. The seeds are moderately large (ovoid, 3-5 mm. long) and without evident special means of dis- persal other than the elastic dehiscence of the capsule, which could, at most, throw the seed only two or three meters. The island-hopping of this species is probably to be explained by the dispersal of parts of the plant itself in times of storm. The xeromorphic stems could probably survive brief im- mersion in salt water, and, if swept beyond the strand line, become established, even under relatively unfavorable con- ditions. The population that invaded the West Indies, or soon evolved there, probably had a moderately large involucre with projecting, slightly three-lobed spur (such as is found in the Grenadines, western Jamaica and the western Ba- hamas), glabrous staminate flowers, and blunt, obovate leaves. Such a plant is very closely approached, in all its features, by the peripheral western population of P. tithy- maloides ssp. jamaicensis. The principal trend in the Greater Antilles has been the development of a population complex with narrow leaves and very small involucres, probably the result of paedomorphic retardation of involucral develop- ment relative to the maturation of the flowers. The trend to narrower leaves is probably an adaptation to relatively dry habitats. This trend may have fluctuated markedly throughout the Pleistocene. The selective basis for reduction in involucre size is not clear. It is to be noted, though, that, except for Anthracothorax, the hummingbirds of the Greater Antilles are all quite small or tiny, which may be related to the size reduction in Pedilanthus. That there has been a selective basis for this trend in supported by the parellel size reduction of P. t. ssp. parasiticus involucres on Isle de la Gonave, Haiti. The populations of the Lesser Antilles were affected to some degree by these trends, es- pecially in involucral form; but these populations have been isolated from those of the Greater Antilles for some time, and later gene exchange from the continent, as evidenced by pilose staminate pedicels, and probably wider leaves, has 92 ROBERT L. DRESSLER not gone beyond St. Croix (where it is probably very recent). The populations of the Bahamas and Jamaica are very closely related, and were probably derived from a His- paniolan common ancester. The eastern Jamaican ja- maicensis is somewhat closer to the modern angustifolius Map 1. The probable routes of dispersal and gene exchange in Pedilanthus ened cenkaldeg: thi sak angustifolius and its immediate allies in the West Indies. Solid line: routes of primary dispersal, dashed lines: gene exchange, dotted lines: major eaves to gene exchange. A. the center of dispersal for the angustifolius type with narro leaves and short involucres, B. the dispersal of genes for pilose staminate flowers into the Lesser Antilles the dispersal of coriaceous, obtus af type of subsp padifolius into Trinidad. D. the center of dispersal of the small, pilose involucres and other res of the “P. Fendleri” ty of Andros, which eserves a primitive, long involucre type in the Ba 2 ater gap between Hispaniol 1 Jamaica. e r gap between Hispaniola and Bahamas. 4 e water gap be n the Greater a wesser Antilles, which has apparently per- mitted limited gene ci from maroon) to Aap oth age but little if any in, reverse: limited gene flow each way. THE GENUS PEDILANTHUS (EUPHORBIACEAE) 93 the Greater Antilles in size and form (or, again, this may be a matter of varying selection pressures). The Bahamas were not available for colonization until sometime in the Pleistocene (Clench, 1938), but their origin is not yet more exactly dated. Plants with quite narrow leaves are found in Puerto Rico, but the modern populations of Hispaniola and Cuba have moderately wide, oblanceolate leaves, sug- gesting that selection pressures may have changed in this area, if bahamensis and the eastern population of ja- maicensis were derived from a similar Hispaniolan ancestor. Both angustifolius and padifolius, as well as plants inter- mediate between the two, have been collected on St. Croix. One may suspect that padifolius has recently invaded this island, probably as an escape from cultivation, but field study is needed to clarify the matter. The Pleistocene fluctuations of sea level in the West Indies have undoubtedly affected the evolution of the biota by uniting some island groups and facilitating dispersal between others. The present distribution and variation of Pedilanthus apparently reflects the last major lowering of the sea in the area, when the Bahamas, the Greater Antilles (except Jamaica?) and the lesser Antilles each formed a more closely interconnected archipelago. How else could the existence of distinctive population complexes in each island group be explained? The somewhat clinal variation in the Bahamas may possibly represent fragments of an actual cline which existed when the Bahamas were partially united in the late Pleistocene. The present distribution and varia- tion of P. tithymaloides may best be shown by mapping cen- ters for certain characters and character complexes, routes of gene exchange, and partial barriers to gene exchange (Map 1). PART IV. TAXONOMY MEASUREMENT The terminology used in the taxonomic descriptions and discussion is that given under Morphology (p. 29), but some of the procedures used in measurement need explanation, as it is not always feasible to follow strict morphological boundaries for descriptive purposes. The involucral tube is measured from the top of the peduncle (a fairly definable point) to the apex of the main lobes, as shown in fig. 12. 94 ROBERT L. DRESSLER The shapes given for the main lobes, however, refer to the anterior portions thereof, and, even then, are rather arbi- trary, because the main lobes grade imperceptibly into the spur parts dorsally and posteriorly. The accessory involucral lobes, strictly speaking, should be measured from the glands, or, in some species, from fairly definable bases anterior to the glands. This procedure, is, in most forms, difficult to follow without demolishing the involucre. This difficulty is here avoided by measuring the differences in length be- tween the main lobes and the accessory lobes (which are shorter) and giving the distances for which the lateral ac- cessory lobes are free laterally and medially. The spur offers similar difficulties in measurement; the external boundary between involucral tube and spur is often obscure (and in P. Palmeri, its appearance varies with the condition of the involucre), so the length is given both from the peduncle (dotted bracket in fig. 12) and from the involucral tube (solid bracket in fig. 12). In those forms where the spur is scarcely elongate, its thickness is given and the dis- tance between the peduncle and the base of the spur (in these forms this is clearly definable). The lateral spur lobes are difficult to measure, for much the same reasons which cause trouble with the main involucral lobes, and little at- tempt has been made to measure these structures, except where they are especially conspicuous. The measurements and descriptions given for the medial spur lobes refer only to the free portions. Where these lobes are markedly con- nate with each other and with the lateral spur lobes, the distance between the apex of the spur and the free portion of the spur lobes is given (“spur lobes arising at’). Where the lateral and medial (free) margins are markedly dif- ferent, this is indicated. Bracteole-ridge refers to a marked keel on the inner wall of the involucre, which appears to represent bracteolar tissue completely adnate to the 1n- volucral wall. The measurements given for seeds refer to the length and the greatest diameter. TAXONOMIC CONCEPTS In the present work, the number of species recognized is less than half that listed by the last revision, that of Mill- spaugh (1913). In such a case the reader may reasonably THE GENUS PEDILANTHUS (EUPHORBIACEAE) 95 wish to examine critically the concepts used. A number of the changes in the present revision are due not to any un- usual species concept, but merely to the great need for revision, and are changes which would have been made by any botanist with adequate material. In recent years, much has been accomplished in the at- tempt to reach relatively objective species criteria which reflect accurately and conveniently the “natural” situation, as nearly as this can be done. The cornerstone of the most adequate species concepts has been the sympatric coexistence of distinct populations which retain their individuality. In spite of the complications of apomixis and polyploidy, these concepts are as usefully applicable to botany as to zoology. This “biological species concept” is used in the present paper. A species is considered to be a population or group of populations which share a common pool of genetic and morphological characters, which are interconnected by actual or potential gene exchange, and are separated from other such populations by barriers to gene exchange and by resultant gaps in morphological features. The barriers to gene exchange may be entirely “intrinsic” or may result from interactions between the intrinsic features of the or- ganism and the “extrinsic” environmental factors. Some may object to the emphasis on the gaps between species, yet this is the one approach which allows a high degree of objectivity in the treatment of species, and the highest possible degree of objectivity should be sought at all levels of taxonomy. It has been well demonstrated that strong barriers to interbreeding do not always curtail all gene exchange (introgression), especially where the environ- ment has been drastically altered. Nevertheless, the presence of any appreciable amount of hybridization places the bur- den of proof upon those who would recognize distinct species. The magic word, “introgression,” is not enough; it must be shown, by careful and critical analysis, that there are bar- riers, other than purely geographic factors, which separate the taxa. The demonstration of such gaps does not neces- Sarily require genetic or embryological data, which cannot be obtained without lengthy experimental work. Critical taxonomy, based entirely on morphological features, was practiced before any alliance between genetics, evolution and taxonomy had been considered. 96 ROBERT L. DRESSLER The biological species concept is most directly applicable when one is dealing with sympatric species. An arbitrary and subjective element is almost inevitable in the treatment of disjunct groups. Even extensive evidence of genetic compatibility cannot indicate what would occur if the popu- lations were naturally sympatric, for they are not. When dealing with disjunct groups one must use a certain degree of “taxonomic judgment,” and this is best acquired by the study of related, sympatric groups. The “species standard” concept ( Rollins, 1952), devised primarily to deal with the complexities of apomixis and polyploidy, can be applied equally well to allopatric populations. A note of caution has recently been brought out by Brown and Wilson (1956), in their re-emphasis of character displacement. Where the features used to distinguish species are those affected by competition, or those involved in any way in the inter- specific barriers, related species will be more sharply dif- ferentiated where they are sympatric. It is to be hoped, however, that this concept will not be used to justify the naming of many disjunct, putative species, merely because some of the populations might be distinct species. Subspecies are used in the almost universal sense of geographic groups which are taxonomically recognizable, but which intergrade where their ranges are in contact. The term, “subspecies,” is taken up in preference to “varietas” with considerable reluctance, for the latter term has been used in this same sense by many botanists, and simple “name-changing” should be kept at a minimum. There has been little uniformity and a good deal of chaos in the botanical usage of infra-specific taxa, and it is hoped that greater clarity may be achieved by the growing use of the more clearly defined term, “subspecies.”” However, this term, too, can easily be thrown into disrepute by careless usage. The existence of broadly sympatric populations which are partially isolated by ecological factors and quite correct- ly treated as subspecies is, in theory, possible. However, “sympatric subspecies” is something of a contradiction in terms, and they must not be proposed without critical study. Similarly, the description as subspecies of segments in a cline, populations which can only be recognized on a statis- tical basis, or (worst of all) phenotypes without population status, can only serve to devalue the concept. THE GENUS PEDILANTHUS (EUPHORBIACEAE) 97 While these concepts associated with the biological species concept and developed through the study of evolution can lead to a high degree of objectivity and a much greater uniformity in taxonomic treatments, most revisions must fall short of the ideal goal. This is not due to inadequate concepts or methods, or to inability of the taxonomist, but to inadequate sampling of the plants. By far the greater part of the world’s vegetation is quite inadequately rep- resented in the herbaria. With poorly known plants we must simply do the best we can and admit that our treatments may be modified when more material and information are available. Another source of inconsistency, the inconsistency of biological nature, is, of course, always with us, and cau- tions against overly narrow or rigid concepts. Borderline cases between subspecies and species, between species groups and genera, are to be expected, and must be recognized and studied as such. Indeed, our whole structure of evolutionary thought demands that such borderline cases occur. Pedilanthus Necker, Elem. Bot. 2:354. 1790. Nomen conservandum. TYPE: Euphorbia tithymaloides L. Tithymalus Mill., Gard. Dict. ed. 4, vol. 8. 1754. Type: T. my[r] ti- folius sm not Tithyma lus Trew, nomen conservandum. Tithymaloides Crees, Tab. Bot. ed. 2. 28. 1783. Type: Tithymalus wjtsfoleas Mill. vlicens ir aaay Tratt., Gen. Pl. 86. 1802. Type: Euphorbia bracteata c idaria Haw., Syn. Pl. Succ. 136. 1812. Type: Tithymalus myrtifolius: Mill. Tirucalia Raf., Fl. Tell. 4th part, 112. 1838. Based in part on Pedilanthus Diadenaria Kl. and Gke., Monatsb. Akad. Berl. 1859:254. 1859. Type: D. Pavonis Kl. and Gke. exadenia Kl, and Gke., Monatsb. Akad. Berl. 1859:253. Type: Pal twthas macrocarpus Benth. For a detailed treatment of the generic and infra-generic synonyms of Hoogigtent ga and rs typification, see Wheeler (1939, 1943). Trees o oody or succulent; leaves entire, alternate, distichous, Wapules nat blunt or spur-like; cymes terminal or axil- lary, dichasial or partly monochasial by abortion, cyme bracts op- 98 ROBERT L. DRESSLER (lateral accessory involucral lobes) above the glands; and a dorsal aoa bilaterally symmetrical, bearing glands on each edge and form- an inconspicuous, more or less ligulate lobe (median accessory in- voloetal lobe) above the glands; the involucral tube distorted, with the glands all on the dorsal side, and often borne on a more cr less elongate, solid, spur-like, dorsal extension of the tube; glands 2, 4 or (by di- vision of the medial glands) 6, appendiculate, the appendages (spur lobes) partly connate with each o her and with the main involucral lobes, forming a more or less el aay spur, the lateral spur lobes 3, bifid, largely connate; fruit capsular or indehiscent; mature seed ecarunculate, smooth or tuberculate. KEY Note that P. Millspaughii (species no. 14), of Costa rela is not ineluded in the key because of inadequate description and lack of mater A. Spur Seen projecting te greed from ge involucral tube for more than m. or, if shorter, the bracts mostly more than 15 mm. long and carta eee the involucre at anthesis .......... B A. Spur short, cucullate, projecting less m. fr the in- 0 volucral tube, bracts usually less than 15 mm. long, never panei Sr the involucre at anthesi -L B. Lateral spur lobes well developed, enfolding and partially con- cealing the medial lobes or completely connate with the medial lobes Cc B. Lateral spur lobes relatively small, not enfolding nor connate with the medial lobes D C. Medial spur lobes free, enfolded by the lateral etd dence project- ing obliquely from the involucral tube (western Mexico) ..... gee osdesnnes alm Medial spur lobes completely connate with each dee ‘and with “the lateral lobes, leaving a small semi-circular orifice over the ac- cessory involucral lobes; spur inflated and flattened parallel a the is involucral tube (Vera acruz kit. D. Spur lobes obtuse, with a thickened, recurved point beneath, mot ptberuate sc ee E D. Spur ‘ches ne truncate, thickened beneath, but without S recurved point. F pol . Cymes sonniabe d; internodes mostly less than 4 mm. long; bracts usually pubescent without; glands ovate or oblong; leaves — or broadly acute (Sinaloa to Guatemala) ou... 3. P. calearatus. ymes lax, internodes ca. 10 mm. long; bracts glabrous tga’ ‘funds orbicular or nearly so; leaves acuminate (coastal Guerre voi F. Tip of spur truncate, sometimes 2-notched and thus 8-lobulate eeeeereeoee Tip of spur rounded, if 3-lobulate then central lobe projecting frien Nag Wa het eee Aa oe mean Bare inacksiy toodsue teers: Be. SRECTHEN I THE GENUS PEDILANTHUS (EUPHORBIACEAE) 99 G. Bracts large (ca. 2 em. avee?s persistent, medial spur lobes com- pletely connate (Michoaca 6. P. coalcomanensis. G. Bracts small (less than 1 cm. long), caducous, medial spur lobes not connate H Tree or large shrub with large leaves (more than 4 cm. long), pistillate pedicel not elongating, capsule developing within the in- volucral tube (Oaxaca 5. P. pulchellus. ow desert shrub, leaves less than 1.5 cm. long, elvan pistil- late pedicel elongating and exposing the developing capsule ( Puebla and a see cent Oaxaca 7. P. cymbiferus. s 6, spur flattened terminally, strongly partitioned within, fruit Pabst: cornuate, indehiscent; latex white (Sonoran Desert) ocarpus I. Glands 2 or 4, spur not terminally flattened or partitioned; fruit capsular, dehiscent; latex yellow J. Cyathia pedunculate; art more than 6 mm. long, style branches usually spreadin K J. Cyathia Giciaites style ca. 4 mm. long, style soem connivent teh (Puebla) nie co uacanus. ‘ vary densely puberulent; peduncles and mais rufescent- tomentose (Oaxaca 10. P. tomentellus. vary glabrous; peduncles and petioles not Rab estar (Sonora to Oaxaca 9. P. bracteatus. L. Medial spur lobes free, attenuate, each with a pronounced tooth the inner margin; lateral glands exposed (Yucatan and Hon- uras) Medial spur lobes truncate, usually connate, without lateral teeth; lateral glands concealed (widespre OM) anes 13. P. tithymaloides. 1. Pedilanthus Palmeri Millspaugh Plate XVI, 2 Al, F!, GH!, s!, uc!, us!). -— Tithymalus koilopremnos Croiz., Am. Jour. Bot. 24: 704, “ae based on P. Palmeri Millsp., not T. Palmeri (Engelm.) Abram P. peritro eae Millsp., Field Mus. Nat. Hist. Pub. Bot. 2: 369. 1913. Type: Hacienda Coahuayula, Michoacan, ye Nov. 1906, ick 80 (F no. 200, 416!). — Tithymalus peritropoides (Millsp.) Croiz., Am. Jour, Bot. 24:704. 1 Evergreen shrub 1-2 m. tall; tending to branch from the base, stems Woody, glabrous; leaves thin, glabrous, eliptieabiong to oblanceolate- obovate or oblong-o vate, 4 m,. wi g, base cuneate long ess; infloresence termina - OL 8 inal cyme and few (1-3) subterminal, abarag ta pedunculate c ;: es (3- ) 5-13 mm at pu lent; bracts pink to red, puberulent gre seeaely so), 100 ROBERT L. DRESSLER 8-15 mm. wide; peduncle slender, puberulent, (12-) 20-30 (-45) mm. long, apex expanded into an oblique receptacle; involucral tube green, glabrate to densely puberulent throughout, usually basally glabrate s 1, involucral lobes i or 0. mm. ' storall and 1.5-2.5 mm. medially, c ru : peduncle), indented essay eal ca. 1 mm. from base in fresh ma- i d mm. fr ase in dried specimens, tip of spur broad, mm. wide; tateral spur lobes well developed, narrowly Shoable-cilintic, extending from n near tip of spur to tip of involucral e, 1 a © < e. . = :O¢ La} < ie) ® -: point, thus arching over and partially concealing the medial lobes nd the accessory involucral lobes; the medial spur lobes arising at or ne pe he spur, rectangular-deltoid, 5-8 apering evenly to tru neate apex 1.5- wide, anterior to the glands; glands smooth, orbicular-oblong, lateral glands -1 wide, 1.5-2 mm. long, sometimes auricular twisted, medial gla nds in pairs, each pair placed back to back on the partition and — subquadrate, 0.8-1.5 mm. wide, 1.2-2 mm. long; bracteoles absent r poorly developed, bracteole-ridges up to - or, ae and ew fila- in outline, base obtu use, terminally pelea raphe dark and flanked by lighter bands, seeds 4.5-5.5 mm. long, 4-4.25 mm, uy Specimens seen: Mexico. Nayarit: 7 mi. E. of San Blas, Dressler 1082 (GH); 29 mi. So. of Compostela, ge 1064 (GH). Jalisco. as cinity of San Sebastian, 3850-5000 ft., E. W. Nelson 4090 (GH, US ie Cruz de Vallarta, 300 m., Mexia iis (A, F, GH, MICH, MO, NY, US); upper slopes of barranca above summit of pass 9 mi. S. W. of "Autlén, toward Manzanillo, UcVeuck 10235 (GH, MICH) ; eantale summits 9- 10 mi. S. W. of Pray 1300-1600 m., MeVaugh 14199 (MICH); along highway 15-18 mi. S. W. of Autlén, 900-1200 m., MeVaugh 11938 (GH, rearre _Michoacén. Huizontla, Coaleoman, 600 on et al. 3 (AHFH, F, HY U, UC, US). Guerrero. Guayameo- Sant Teresa, Mina, 1040 m., G. eee 9955 (AHFH, THE GENUS PEDILANTHUS (EUPHORBIACEAE) 101 F, G, K, NY, U, US); by the trail to Cundan-Grande, San Antonio, Montes. de Oca, G. B. Hinton et al. 11678 (AHFH, F, GH, K, NY, U, UC); San Antonio,Montes de Oca, G. B. Hinton et al. 14004 (AHFH, F, GH, ARIZ, GH, K, NY, US); Plan gs Carrizo, Galeana, 900 m., G. a noes 00 y; E. 08 (GH, K). Without plete locality Nueva Espafia, Sessé, “Mociio, Castillo and Maldonado 1768 © P Pacmeri 3 : : @® P. Finkn 10s 100 DISTRIBUTION AND HABITAT. (Map 2) From near sea level to 1,500 m. alt. along the western edge of the Sierra de Volcanes and the Sierra Madre del Sur, from Nayarit to north-central Guerrero. In Nayarit, this species was found along small streams in Orbignya palm forest (the palm forest is broken by scattered dicot trees along the streams) and in tall mixed forest. Data on collectors’ labels, “dense shade,” “forest by the river,” “moist woods,” “cafion bot- tom,” “ravines,” etc., indicate that these habitats may be fairly characteristic for the species. Flowering in winter, November to April. COMMON NAMES. Jumete, western Jalisco. VARIATION. Leaf shape varies from narrowly elliptic (cuneate base and acute apex) to larger and broadly obovate 102 ROBERT L. DRESSLER (obtuse base and apex) and is fairly constant for a given collection, but shows no clear-cut geographic pattern. Size of cyathium also seems to show no distinct geographic pat- tern, but this is obscured by differences in season of collec- tion, the first formed cyathia usually being the largest on a given plant. The fruits and seeds (though these are poorly sampled) are apparently larger in southern plants than in the northern ones. The bracteoles tend to be better developed and the glands to be more nearly terminal in the spur (the spur partitions less well developed) in the southern collec- tions. Langlassé 808 is distinctive in its relatively small leaves and its compact inflorescence (short internodes and peduncles) with relatively large bracts. It probably repre- sents a population which has not been sampled by more recent collectors. Cultivated material of Dressler 1064 (Nayarit) consistently bears the cyathia upside-down as compared with all other species of Pedilanthus (in Plate XVI the cyathia are drawn in the upright position for better comparison with other species) ; the long peduncles, in effect, undergo resupination, and the large spurs are borne beneath the involucral tube, which, like the peduncle, is horizontal. The spur would seemingly serve its normal function quite well in this position, the anthers and stigma lobes touching the visiting hummingbird on the forehead rather than the chin. It is difficult to determine the posture of the cyathia from herbarium material; the cyathia of the Langlasse col- lection discussed above, at least, seem not to be resupinate, and it seems improbable that its relatively short peduncles could twist in the same manner as the longer ones more characteristic of the species. This resupination, thus, may not be universal in P. Palmeri. RELATIONSHIP. Pedilanthus Palmeri is very distinctive in the position and great development of the lateral spur lobes and in the origin of the medial spur lobes near the apex of the spur. The unusual posture of the cyathia of at least some plants (discussed under Variation) is also unique. In aspect of the plant and structure of the cyathium, Palmeri resembles P. Finkii, and the two are doubtless close- ly related. They do not seem to be very close, in morphology, to any other species. The well developed and distinct spur lobes, the many staminate flowers, the short involucral tube, THE GENUS PEDILANTHUS (EUPHORBIACEAE) 103 and the drrecreenay eure may be interpreted as primitive features in P. Palm NOTES. The ake alberta seen of Emrick 80 the type of P. peritropoides, is badly moulded and quite lacking in cyathia or complete bracts. The lateral view of the cyathium as sketched by Millspaugh (in herb. F) is distinctly that of P. Palmeri, and the available foliar scraps do not differ in any way from that species. The other drawing made by Millspaugh shows the spur lobes and indicates that he dis- _Sected out the median accessory lobe of the involucral tube and interpreted this as a third spur lobe. The other species described by Millspaugh as having a “tripartite appendix lobe” is discussed under P. tithymaloides ssp. parasiticus, and here, too, the cyathium was apparently discarded after being sketched and described. 2. Pedilanthus Finkii Boissier Plates XIII & XVI, Map 2 P. Finkii Boiss., DC. Prod. 15(2): 1261. 1866. Type: Found in a ng eas at the source ee the Atoyac River in a very damp situation, Hanbury IV/1865, from H. Finck, Cérdova, Mexico (K!). — ail Finkii Bike ) O. Kuntze, Rev. Gen. Pl. 2: 620. 1891. terminal and often | or 2 pe edune ulate, upper-axillary cymes, inter- — (3-)5-30 mm. long, nage mae rey elliptic-oblong to aliptie . long, 2 mm., accessory lobes ligulate, densely appressed-puberulent, obtuse, 0.8-1.5 mm. wide, lateral lobes slightly shorter (to 0. than main lobes, free 1.5-2 mm. laterally, 2-3 mm. medially, median lobe ca. 3.5-4 mm. from me tube ( - mm. from ‘peduncle) etal ated ere abruptly for ward and exp ng into an inflated, semicircular flattened — hay the involueral tube, ie hood 7-8 mm. in width and 8-12 mm. long, the an r edge rea or surpassing the tip Seals" of the eetoeest tube; Se “redial and Saas spur lobes cael 104 ROBERT L. DRESSLER connate, leaving a scncines ia gy ca. 1.5 mm. wide over the ac- rest involucral lobes at ca. . from the apex of the involucral tube, the apices of the mig neg tt es tis argin of orifice) thickened and densely puberulent within; iahide sessile or slightly stipitate, flattened the spur anterior to the curve, the medial glands anterior to the lateral glands; lateral glands semi-orbicular, 1-1.2 mm. in diameter, medial glands each single, sub-rectangular, ca. 0.8 x 1.2 mm.; parti- tion none; bracteoles 1-several in a cluster, filamentous, 3-6 mm. long, cluster united basally; staminate flowers 30-37,, glabrous, pedicels glabrous, pedicel 7-8 mm. long, ovary spheric-ovoid, ca. 1.5 mm. long, style wih ph ca. 7-10 mm. long, trifid, branches attenuate, c 2 g, entire or slightly bifid; fruit 6-7 mm. long, sub-orbicular in outline, deeply trilobed in cross " section ; mature seed kissin Specimens seen: Mexico. Veracruz. Potrero, Vallée de Cérdov pO pa 1802 (F, P); Cérdoba, Galeotti 875 (F, G). Cultivated a ‘the Missouri Botanical Garden, from the Mexican Commission of Louisiana Purchase Exposition, 1904, geographic source ids Jan. 21, 1907, C. H. Thompson (A, F, MO, photos of living plant F, MO). DISTRIBUTION AND HABITAT. (Map 2) Known only from the vicinity of Cérdoba, Veracruz. The data accompanying the type collection indicate a rather moist habitat, as does the presence of epiphyllous bryophytes on the same collection, and a fragment of Peperomia (P. Deppeana or P. quadri- folia) pressed with Bourgeau 1802. From the scanty data, the locality and the texture of the plant, this may occupy the most mesic habitat of the genus. VARIATION. There is little variation in the small sample available. The leaves of the plant cultivated at Missouri Botanical Garden are narrower and more acute than most leaves of the other collections, and the glands of Bourgeau 1802 are much further forward in the saccate spur than are those of the other collections. The type collection has quite large involucres. RELATIONSHIP. This species is remarkable for the complete connation of the spur lobes and the hood-like, saccate spur parallel with the involucral tube. It is too similar to af Palmeri to warrant the maintenance of section Calceolas- trum. 3. Pedilanthus calearatus Schlechtendal Plates XIII & XVI, Map 3 P. calcaratus Schlecht., Linnaea 19: 255. 1847. Neotype (here chosen): Mexico, Veracruz, Barranca near Corral de Piedras, THE GENUS PEDILANTHUS (EUPHORBIACEAE) 105 a ch 1916, C. A. Purpus 7544 (UC 185411:, isotypes: A!, GH — Tithymalus calcaratus (Schlecht. ) Croiz., Am. 37 < Eas P. er eoesaideetsioe Baill., Adansonia 1: 340. 1861. Type: A plant grown in Paris from material collected by Ghiesbreght in Mexico, n herbarium material has been located. — Tithym alol sides Ghies- 5M yeaa (Baill.) O. Kuntze, Rev. Gen. Pl. 2: 620. cradenius Donnell Smith: Bot. Gaz. 19: 263. ior Type: Guhink|a: Canibal, Dept. Huehuetenango, alt. 3,180 ft., Dec., 1891, W. C. Shannon 412 (us 931112!). — Tithymalus macradeni qs (Donn. ery Croiz., Am. Jour. Bot. 24: 704. 1937. P. Purpusii Brandegee, Univ. Calif. Pub. Bot. 4: 377. 1913. Type: Mex xico, Veracruz, Barranca de Santa Maria y Ternera, a an, Pei; 19182°C.-A. ’ Pur urpus 5759 (UC ieeseat isotypes: A!, GH!, MO!). — ® 2 _ co 3 = *. 3 = e Cel a. ry eo EY me and ltd, tapering to a Hputute ened ca. 1.5-2.5 mm. long and 1-1 material), “sto 1 glands at the base of the spur, oblong-reniform to elliptic. oblong, 1.5-2 mm. long, 0.8-1 mm. wide, lateral glands ca. 2 mm. aileron to the medial glands, externally visible between the lisersd si medial spur lobes, oblong-elliptic to elliptic-ovoid, 0.8-1.5 mm. long, 0.4-1 mm. wide, spur partitions not developed; bracteoles Several to numerous, ca. 8-10 mm. long, filamentous, twisted (oc- casionally a few ligulate and pilose), basally connate, adnate to the involucral tube for ca. 3-4 mm , the dorsal bracteole-ridge beg cern terminating in a pilose, linguiform lobe ca. 1 mm, yoo Ti 2 a les mm glabrous, 20-34, t vabieal. clusters tending to on e 2-3 few rsa than the lateral and dorsal clusters, pedicels 8-13 long, filame long; pistillate glabrous or terminally in diameter, glabrous to densely erisped- tothe rulent, style 7-10 mm. al fo puberulent throughout, style branch 05- 1 long, bifid, con- nivent or spreadi ing; fruit vohaeiaesik dee a "3-lobed. ca. 6 mm. long, glabrous to crisped-puberulent; seed ovate-subquadrate, truncate es Bopeosy acute apically, very weakly keeled, ca. 3 mm. long, 2-2.5 wide, slate-grey or brownish grey, vane dark, hilum flanked by 2 slong dark areas, each about equal to the hilum in area. Specimens seen: Mexico. rice About 6 km. from Sisal "4 eprint road), Dressler 1810 (GH); km. 28, Mérida-Progreso road, undell 7978 (A, US); re 25, Mérida-Progreso road, Drsvelee 1362 (Gu); hx ees, Millspaugh 1667 (F); Mina de Oro, north coast, G. Gaumer and sons 23324 (A, F, GH, MO, NY, US); Silam, G. Gaumer s. n. (F). Honduras. Dept. Comayagua, ca. 600 m., Standley and J. Chacén P. 6034 (F); El Banco, Dept. Comayagua, 640 m. Valerio Rotrtpies 2385 (F). DISTRIBUTION AND HABITAT (Map 4). North coastal Yuca- tan and the Comayagua Valley in Honduras. In Yucatan this species is found in the open thorn scrub which is typical of the northern coast (see fig. 24). Near Sisal and Progreso, 136 ROBERT L. DRESSLER the species occurs near the coastal swamps, in areas where the water table is very near the surface; at Progreso it is not limited to this narrow zone, but occurs for two or three kilometers along the Mérida-Progreso road. Like sev- eral other species of this genus, its local distribution is spotty and erratic, but it is quite abundant in some sites. Its habitat in Honduras is scarcely coastal, but appears to be an open thorn scrub similar to that of Yucatan. The distribution, as now known, is quite anomalous. Little is known of the phytogeography of the arid region of Honduras; it is clear, though, that the present distribution in Yucatan must be relatively recent, since the limestone which makes up the northern part of the peninsula is of late Pleistocene and Recent origin. Flowering in the spring and summer (Feb.- Aug., flowering sparsely in September). COMMON NAMES. Pie de nifo, ditamo real (Honduras) ; Yaxhalalché (Yucatan, Mayan “green-stem-tree”’). VARIATION. The Yucatan population of P. nodiflorus is variable in indument and color. The cyathia of the plants near Sisal and Progreso are all densely puberulent without and green beneath, the involucral tube being red only near the lateral spur lobes. The Silam collections vary from densely to rather sparsely puberulent, and the Mina de Oro collection from densely puberulent to nearly glabrous with- out. The indument of the ovaries varies correspondingly. Some of the cyathia from Mina de Oro, at least, are uniform- ly red (color cannot always be determined from dried ma- terial). Leaf size is highly variable, as is often the case with species which are essentially leafless. Honduran material differs somewhat from the Yucatan material in a number of details, but is, on the whole, very similar. The cyathia of Honduran plants average a little longer; the indument of the involucral tube is moderately dense without an somewhat shorter within than in Yucatan plants; the 1n- volucral lobes are wider terminally and the free portions of the accessory lobes longer; the medial spur lobes are a little more slender and the terminal lobules more strongly curved ; the lateral glands tend to be smaller and less conspicuous ; and the styles and staminate pedicels tend to be longer. The leaves, fruits and seeds of the Honduran plants are unknown. RELATIONSHIP. Reference to Miss Ericksen’s accurate drawings (Plate XXI) will show that this, in the form of THE GENUS PEDILANTHUS (EUPHORBIACEAE) 137 its involucre, is one of the strangest of a bizarre genus. The eye-like, exposed lateral glands are unique and give the structure a quite animate appearance. Though P. nodiflorus superficially resembles P. tehuacanus in some respects, their juxtaposition is coincidental, and not meant to indicate close relationship. In the form of the involucral lobes and the shortness of the spur, the two species are similar, but they differ widely in most other characters, and their re- semblances are probably largely parallel developments. In the cucullate spur, the “palate” at the base of the accessory involucral lobes, the reduction of the ventral clusters of staminate flowers, the folded lateral spur lobes, and the in- florescence, seed and fruit characters, P. nodiflorus resem- bles the P. tithymaloides complex, and they are thought to be close relatives. In inflorescence, indument, seed and in- volucral texture, this species somewhat resembles P. pulchel- lus, and, to a lesser degree, P. calcaratus. These resem- blances are taken to represent a more distant common ancestry. Notes. The Honduran plants of this species are, in gen- eral, very close to those from Yucatan; the gap in the known distribution is, itself, far more striking than the slight morphological differences which are apparent, and, at least until its range and variation are much better known, there is little advantage in subspecific recognition of the southern population. 13. Pedilanthus tithymaloides (L.) Poit. This, the type species of the genus, is by far the most widely distributed and variable of the slipper spurges. It is the only species in which clear geographic populations of subspecific rank are found. The allopatric West Indian taxa have generally been treated as distinct species, but they are all very closely allied to P. tithymaloides, sensu strictiore. There is actual intergradation in some cases, and in others, the subspecies appear to hybridize freely, where their ranges have been changed by the influence o man. While angustifolius and parasiticus are broadly sympatric and appear to behave as distinct species (there is no evidence that they actually occur together, in the same habitat), each is closely tied in to the mainland population by a different chain of taxa. The best arrangement, at the present, is to 138 ROBERT L. DRESSLER consider all of these geographically replacing populations as subspecies of the somewhat variable P. tithymaloides. It is interesting that there are no clear subspecies in the main- land population complex which ranges from northern Mexico to Surinam. Plants which appear to be hybrids between different subspecies are discussed on pages 167-170; such plants would be difficult to identify by the use of the follow- ing key. KEY TO THE SUBSPECIES OF P. TITHYMALOIDES A. Leaves wide, more than half as wide as long A. Leaves narrow, less than half as wide as long; staminate pedicels always gla B. Leaf f apices pees or, if obtuse, leaves ovate and somewhat at- tenuate to a C Leaf in obtuse or retuse, if obtuse, then leaves usually obovate C. Leaf bases cuneate, leaves not markedly glos D C. Leaf bases obtuse to subcordate, leaves gabrate. “0g glossy (Yucatan and Greater Antilles). 13b. P. t. ssp. parasitic D. Stem straight or nearly so (Mexico to northern South ; Acerien) 13a. P. t. ssp. tithymaloides. D. Stem zig-zag, usually markedly so ( Cuba and Florida). ........-+++++ 3c. P. t. ssp. Smalli. E. Leaves retuse, widest below the middle (Amazonia) ........+++s+++ ida. f. retusus. Leaves obtuse, obovate or elliptic, widest above the middle (Les- ser Antilles 18e. P. t. ssp. padifolius. Styles short, 3-5 mm. tid alice tube usually short, 6-9 mm. long, glabrous without or rsely pilose with stiffly erect hairs; ovary glabrous (Greater Antilles Except JAMAICA) ........seseeerersersrerees 13¢. P. t. ssp. angustifolius. F. Styles longer, 5-10 mm. long (if short, ovary densely tomentu- lose); involucral tube longer, 7.5-11.5 mm. long, glabrous without OF MOTE OF less pubeErulent .........0cccccecsrserrccsssecssscssevssseeeeseersonsssssnscossncess G. Involucres minutely crisped-puberulent Vege the trichomes appressed (Bahamas) 13h. P. t. ssp. bahamensis. G. Involucres glabrous or puberulent without, bat not minute ly crisped-puberulent (Jamaica) .........:008 13f. P. t. ssp. jamaicensis. 13a. Pedilanthus tithymaloides subsp. tithymaloides Plate XXI, Map 7 Euphorbia tithymaloides L., Sp. Pl. 453. 1753. Neotype (here desig- nated): Plate 92, Jacquin, Sel. Stirp. Amer. Hist., 1763. — Tt ithymalus tithymaloides (L.) Croiz., Am. Jour. Bot. 24: 704. 1937. Euphorbia tithyma loides [a] myrtifolia L., Sp. Pl. 453. 1753. Type (here designated): Plate 16, Commelin, Hort. Med. Amst. vol. 1, THE GENUS PEDILANTHUS (EUPHORBIACEAE) 139 1697. — Tithymalo[i]des myrtifolium (L.) O. Kuntze, Rev. Gen. Pl. 2: 620. 1891 Pithymatus my[r]tifolius Mill., Gard. Dict. ed. 8, Tithymaius no. 1. 1768. Type: a plant sent from Cartagena by Robert Millar and culti- : . — Crepidari myrtifolia (Mill.) Haw., Syn. Pl. Suce., 136. 1812. — Pedilanthus myrtifolius (Mill.) Link, Enum. ad Berol. 2: 18. 1822. Euphorbia canaliculata Lodd., Bot. Cab. 727. 1823. Type: Plate 727, Bot. Cab. — Pedilanthus canaliculatus tread" Sweet, Hort. Brit. (e P. Houlletii Baill., Adansonia 1: 431. 1861. Type: a sre grown in the garden of M. Ho llet. in Paris; a sterile specimen c hort. par., 1859”) may be from this plant. — Tith ennlol tides Houlletii (Baill.) 0. Kuntze, Rev. Gen. Pl. : : P. endleri Boiss., DC Prod. 15 (2): 5. 1866. Type: — ) ithy es Fend (Boiss.) 0. Kuntze, Rev. Gen. Pl. 2: 620. 1891. — Tithymalus Fendleri biaaede Croiz., Am. Jour. Bot. 24: 704. 1937, not T. Fendleri Kl. and Sen lat 5107 aa isotype: F!). — Tithymalus Pringlei (Robins.) P. gritensis Zahlbr., Ann. K. K. Nat. Hofmus. Wien 12: 104. 1897. nf rah Venezuela, La Grita, Mérida, pp s. n. (W! fragment and pho F!). P. Deamii Millsp., Field Mus. Pub. Bot. 2: 356. 1913. Type: Guate- mala, Fiscal, elev. 3,700 ft., May 31, si ravines, C. C. Deam 6081 (F 247,005!, isotypes: a!, GH!, MIcH!, us!). — Tithymalus Deamii (Miilsp.) Croiz., son Jour. Bot. 24: 704. 1937. a campester Brdgee., Univ. Calif. Publ. Bot. 6: 56. 1914. Type: Mexico, Oaxaca, Picacho — San Gerénimo, Oct., 1913, dry, rocky Plains, C. A. Purpus 6885 (UC 172,985!, isotypes: - Fl, = mo}, NY!, vs!). — Tithymalus villicus Croiz., Am, Jour. Bot. 24: 704, based on P. campester, not T. campester (Cham. ie Sehlacht.) Kl. Gke. P. petraeus sl Univ. Calif. Publ. Bot. 10: 411. 1924. Type: Mexico, Veracr nes April, 1923, on rocks, C. A. Purpus 8885 (ue 218 ,896!, pagar : A). — Tithy malus petraeus (Brdgee.) Croiz., Am. Jour. Bot. 24: 704. "1987, P. ierensis Britt., Bull. Torr. Bot. Cl. 53: 468. 1926. Type: Trinidad, B. W. I., Penal Rock Road, March 28, 1920, woodlands, 1.5 m. high, - Tithymalus ierensis (Britt.) Croiz., Am. ton ur. Bot. Ps 704, 1937. camporum, Standl. and Steyerm., heap rate foe tosh 23: 124. about 1 stall, stems dark g een, involucre. rose-red, comm Standley 3 87781 (F 1,109,145!, asus A!). 140 ROBERT L. DRESSLER Shrub 0.4-3 m. tall, deciduous or evergreen, stems slender and woody or thick and markedly succulent, puberulent when young, becoming glabrate with age; leaves ovate, elliptic or ovate-lanceolate, glabrous in, 4-12 (-14) mm. long, 2-5 mm. wide, ovate, ovate-lanceolate or obovate, attenuate, acute or obtuse, peduncle glabrous to puberulent, 3-8 mm. long; involucral tube red above, basally green or is yellow, glabrous or pilosulous without, glabrous, pilosulous (especially dorsally), or sericeous within 7-14.5 mm, long, main lobes oblong or cuneate-oblong, obtuse, usually ciliolate or hispidulous- -ciliolate on margins, free beneath for 2-4.5 mm., accessory lobes ligulate or te) 3.5-8 mm. medially, median lobe 1-5 mm. shorter than lateral Tee 2-6 mm. long; spur red, glabrous or pilosulous without, glabrous or hispidulous within, Reet arising at 0.5-3.5 mm. from peduncle, -4 mm. thick, projecting 1-3 mm. posteriorly, lateral spur lobes pectin: folded in over accessory involucral lobes, medial lobes con- gether narrowly deltoid, ee ae or ligulate from a acid gety 3-6 mm. long, 3-4.5 mm. wide basally, 0.6-1.2 mm. wide at tr uncate or fetus apex, often thickened beneath at apex or some- nd sometimes hispidulous beneat apex; medial glands basal or (in markedly projecting spurs) eter es reniform-oblon sub- pice oblong or quadrate-oblong, 0.8-1.6 mm. long, 0.6-1 mm. wide, what ent about partitions, rarely more or less split into gland- ace lateral glands 0.5-1.5 mm. anterior to medial glands, oblong, ovate-oblong, elliptic-oblong or lanceolate, 0.3-2.5 mm. long, 0.2-0.9 m. wide pidulous to villous for 1-7 mm., filaments glabrous or puberulent, m. long, anthers glabrous or sparsely pilosulous; pistillate pedicel 4-14 mm. long, glabrous to tomentulose, ovary oblong or oblong-obovate, trigonous, ca. 1.5-1.8 mm. long and 1.2 mm. wide, glabrous, puberulent or tomentulose, style glabrous or sparsely pilosulous, 5-11 mm. long, branches 0.5-1 mm. long, sub-entire to % bifid; capsule subquadrate, 5-6 mm. in diameter, deeply 3-lobed in cross section; seeds ovoid or subglobose to quadrate-oblong, faintly keeled or not at all, 3-4.5 mm. long, 2.5-3.2 mm. wide, ashen grey, THE GENUS PEDILANTHUS (EUPHORBIACEAE) 141 grey-brown or red-brown, usually mottled, base truncate or obtuse, apex acute but low. Re resentative specimens seen: Mexico. Tamaulipas. 7 mi. So. of Padilla, June 25, 1951, C. M. Rowell 2350 (MICH) ; Chamal Hacienda, June 25, 1919, E. O. Wooton s.n. (US). Veracruz: Km. 368, Jalapa to L a Remudadero and Puente Nacional, March 1935, C. A. Purpus 1624 (Cc); Puente Nacional, 1928, C. A. Purpus 10981 (F). Guerrero. Petat- lan, Montes de Oca, alt. O m., G. B. Hinton 10331 (A, AHFH, GH, K, MICH, NY, UC, US). Oaxaca. Vicinity of Cuicatlan, E. W. Nelson 1682 (F, GH, US); dry hills bordering the valley of Cuicatlan, 2,000 ft., Pringle 6043 (A, GH, MO, NY, PH, 8, UC, US); Tomellin Canon, J. N. and J. S. Rose 11360 (NY, US); canyon above Tomellin, Gentry 6998 (AHFH) pec, Seler 1682 (GH); Salina Cruz, Deam 119 (GH, MICH); Picacho, Purpus 7352 (uc). Chiapas. Copainalé, Wonderly 15 (MICH); Pan- American highway 24 mi. S. E. of Comitan, Carlson 1942 (F); Trapi- chito, Comitan, 1,350 m., Matuda 5670 (F). Guatemala. Retalhuleu. Plains between Nueva Linda and Champerico, 120 m. or less, Standley 87775 (F). Escuintla. Escuintla, 1,100 ft., J. Donnell Smith 2072 (Us). Guatemala. Near Fiscal, about 1,100 m., Standley 80399 (F); Concua Ana, 370 m., Standley and Padilla V. 3213 (F). Honduras. Near Pro- greso, Dept. Yoro, 30 m., Standley 55106 (A, F, US); valley near dam for water supply of Progreso, Atlantida, Bangham 350 (A, F); San Pedro Sula, Dept. Santa Barbara, 350 m., Thieme 5467 (US); vicinity of Coyoles, Aguan River Valley, Dept. Yoro, Yuncker, Koepper and Wagner 8639 (F, GH, MICH, MO, NY, S, us) ; Wispernini Camp, Guarunta region, Colén, 75-100 ft., C. and W. von Hagen 1335 (NY). Isla de 23’ W., Proctor acienda Animas, Prov. Guanacaste, 500 r tral, headwaters of the Rio Cafazas, 300-600 m., Prov., Veraguas, ‘ttier 5000 (US); Penonomé Allen 196 (F); Aguadulce, Prov. Coclé, Pi and vicinity, Williams 229 (NY, US); Taboga Island, Macbride 2804 (F, US); near Madden Dam at Alahuela, 70 m., Dodge, Steyermark and Allen 16824 (G, 8); Tabernilla, Cowell 276 (Ny). Colombia. Valle del Cauca. La Paila, 1853, J. F. Holton s. n. (ny); El Valle, near Cali, 142 ROBERT L. DRESSLER m 4 millo 38265 (US). Huila. Entre Gigante y Rioloro, 820-860 m. Pérez Arbelaez and Cuatrecasas 8339 (US). Meta. Salinas de Macapay, H. Garcia B. 5115 (us). Bolivar. Tierrabomba Island, Cartagena Bay, Killip and Smith 14127 (GH, NY, US). Atlantico. Alrededores de Puerto Colombia, altura aproximada 200 m., Barkley and Gutiérrez V. 1841 (GH, NY, US); entre Puerto Colombia y Salgar, Dugand and Jaramillo 3223 (US); cerca de Ponedera, 5-10 m., Dugand and Jaramillo 2752 (us). Magdalena. Santa Marta, H. H. Smith 1295 (¥F, GH, MO, NY, PH, S, U, US); common on road from La Paz towards Manaure, ca. 200 m., Haught 3871 (A, US). Goajira. 9 km. So. of Carraipia, 200 m., Haught Paraguana, 700-800 ft., Curran and Haman 717 (GH, NY). Carabobo. Hacienda Taborda near El Palito, 0-200 m., Pittier 7674 (GH, US); vicinity of Puerto Cabello, Rose and Rose 21862 (us). Aragua. Ocumare e la Costa, 0-15 m., Steyermark 54938 (F, NY). Distrito Federal. La Guaira, Robinson and Lyo n. (US); near Maiquetia, Pittier flowers, Curran and Haman 204 (GH, NY). Trinidad. B. W. I. Western end of Monos Island, Britton, Britton and Brown 2737 (Ny); Little Gasparee, Britton 2789 (GH, NY, US); Manzanilla, N. L. and E. G. Britton 2194 (GH, NY, US); North Manzanilla, headland to E. of bay, 40 ft., Baker and Simmonds (K); Trinidad, Fendler 699 (k). British Guiana. Waini River, Northwest Dist., lat. 8°20’ N., long. 59°40’ W., De La Cruz 3771 (F, GH, MO, NY, PH, US, US). Surinam. Commew1jne River near plant. Ma. Retraite, Soeprata 39B (U). DISTRIBUTION AND HABITAT (Map 7). From Tamaulipas and Guerrero in Mexico south through Central America (ex- cept Yucatan Peninsula) to southern Colombia, and along the northern coast of South America (including Curacao, Isla de Margarita and Trinidad) to Surinam. In northern Oaxaca, this species is frequent in the open, arid thorn forest of Tomellin Canyon (see fig. 23) ; on the Isthmus of Tehuantepec, it is found in low thorn scrub, and, here too, it is quite frequent. Near Tuxtla Gutiérrez, Chia- pas, P. tithymaloides is occasional and local in tropical deciduous forest on limestone (see fig. 22). On the Pacific slope of Guatemala the species is found in “brushy, rocky slopes,” in Petén and Honduras, its habitats range from “semi-arid woodland” to “jungle,” “logwood swamp,” and “humid tropical forest.” The vegetation of rocky sites (pedregal) occupied by this species at Uaxactun, Guate- mala, is briefly described by Bartlett (1935, p. 20). In THE GENUS PEDILANTHUS (EUPHORBIACEAE) 143 northern South America, similarly, the sites in which this species is found range from “arid littoral scrub” to “cliffs in rather humid forest.” It is evident that this species has P ritHymacoioes | @ SUBSP TITHYMALOIDES * SUBSP PARASITICUS 7 iN ea pid 4 Swacun aes 5S SE = Ai ay cry vo as so @ susse TITHYMALOIDES MAP 8 # SuBSR RETUSUS Ps ritHymavoiwes ee SE be ae a = SUBSP) BAHAMENSIS 9 @ suese JamaiceNsis @ SUBSP ANGUSTIFOLIUS % ® SUBSP PADIFOLIUS 8 g ® ee Maps 7 aNp 8, The paar at Pedilanthus tithymaloides. A single a of subsp, preialess aby from Su is not shown in Map 7, but is shown in the inset of Map 8. Some or all of ae localities for subsp. parasitieus outside of stiaiianlichs y represent escapes from cultivati 144 ROBERT L. DRESSLER a much wider range of habitats than any other in the genus, but that, in the wetter areas, it is usually to be found in rocky sites, and frequently on limestone. Flowering especial- ly in the winter, but found in flower at all seasons of the year in different areas. COMMON NAMES. Candelilla (Tamaulipas, Mexico), saca espinas sin dolor (Puebla, Mexico, “remove spines without pain”), comalpahtli (central Mexico, Nahuatl: “griddle medicine,” referring apparently to its preparation as medi- cine). The other Nahuatl names reported by Hernandez (1651), chapolxochitl (grasshopper flower), mincapahtli (arrow medicine), tenapalitl (live-forever), and pinipiniche (?, probably not Nahuatl), seem not to have been reported more recently, but some may still be in use. Suelda con suelda (Guatemala, referring to use in treating broken bones?), pie de nino (Central America), ditamo real and variants (bitamo, pitamo, itamo, etc.) (Central and South America and West Indies), hierba ipecacuana (South America and West Indies, doubtless referring to its purga- tive qualities), espuela de gallo, clavo de Cristo (Colombia), pinopinita (Venezuela), Melksapblad (Surinam). VARIATION. The vegetative features of Mexican P. t. ssp. tithymaloides reflect the relatively xeric habitat of these northern populations. The stems tend to be thick and suc- culent and the leaves to be puberulent and of moderate size (3-6 cm. long) ; the leaves of Mexican plants are generally markedly elliptic. In northern Oaxaca, at least, the leaves on some shoots are quite small and there seems to be a dis- tinct tendency toward foliar dimorphism (see Plate XXI). The cyathia of Mexican plants are of relatively large size (12-14 mm. long) and nearly or quite glabrous. The few collections from Veracruz bear cyathia which are sparsely hispidulous within and the cymes are mostly axillary. In southern and western Guatemala (including Zacapa and Chiquimula), the plants bear foliage similar to that of the Mexican plants, but the stems are mostly very slender. The involucres are of moderate size (10-11 mm. long) and nearly glabrous, except for the two collections from Re- talhuleu, the basis of P. camporum. In these collections, the cyathia are pilosulous without (the hairs rather thin), the ovaries tomentulose and the anthers sparsely pilosulous, the staminate pedicels and filaments glabrous. The bracts of THE GENUS PEDILANTHUS (EUPHORBIACEAE) 145 these plants are small (4-5 mm. long) and densely puberu- lent. No leaves are present in either collection; more ma- terial is needed from northwestern Guatemala. In Petén and Izabal, on the wetter Atlantic slopes of Guatemala, and in British Honduras, the plants of P. tithymaloides are most- ly markedly different; they show the very slender stems of the other Guatemalan collections, while the leaves are larger (up to 8-12 cm. long), glabrous or glabrate, more ovate (proportionally much wider near the base) and sometimes obtuse. The involucres of these plants are fairly large (ca. 11-12.5 mm. long) and glabrous or nearly so. Honduran collections are similar to those from Izabal and Petén in vegetative features, but some are remarkable for their large bracts and involucres and short, non-elongating pistillate pedicels; in Standley 55106, the bracts are 12-14 mm. long, attenuate and bright red, the involucres are 13.5- 14.5 mm. long, and the pistillate pedicel 4-5 mm. long. Most other Honduran collections are similar but less extreme in their features; Yuncker, Koepper and Wagner 1335 has only a few cyathia, but these have normally elongating pistil- late pedicels. In the short pistillate pedicel, the Honduran P. tithymaloides resembles P. pulchellus and P. coalco- manensis. Proctor 3418, from Isla de Providencia, resembles the plants of eastern Guatemala, but has a more succulent stem. The rest of Central America is too poorly collected to permit any discussion of variation. The plants from southern Colombia and some of the Panamanian collections (Cowell 276 and Williams 229) re- semble the sort of plant which was described and figured as P. canaliculatus; these plants have widely lanceolate leaves which are strongly attenuate and have the midrib markedly winged beneath. The staminate pedicels of these plants are puberulent. This sort of plant is often cultivated in the West Indies and could well have been introduced from Panama. South American collections of P. tithymaloides are quite variable as to leaf shape (probably to be correlated in large part with dryness of habitat), but show a very interesting pattern of variation in the features of the involucre. In western Colombia, the involucres are of medium size (10-12 mm. long) and glabrous or somewhat pilosulous, the stami- nate pedicels are puberulent or pilosulous, and the spurs 146 ROBERT L. DRESSLER are similar to that figured in Plate XXI (a Mexican plant), rounded and somewhat projecting, with the median spur lobe partially connate to the lateral spur lobes. In eastern Venezuela, and especially on the islands of Margarita and Curacao, the involucres are much smaller (7-9 mm. long) and markedly pilosulous without, the staminate pedicels are villous, the spurs are only slightly projecting, and the median spur lobes are free (from the lateral lobes) nearly or quite to the base. This last feature permits the spur to gape widely when lateral pressure is applied, as in pressed specimens (in this, the specimens have somewhat the aspect of subspp. bahamensis or angustifolius). It should not be thought that these two types are distinct populations; they are merely the two ends of a complex series of clines. The plants of eastern Colombia and western Venezuela bridge the gap as nearly as can be expected of the available collec- tions (see Map 7). The type of P. gritensis is such an 1n- termediate plant as to size and form of the involucre, but has an unusually dense indument in all parts, and propor- tionately very wide leaves. Pedilanthus tithymaloides from Trinidad does not greatly resemble the Venezuelan plants (there is a considerable geographic gap, bridged only by one sterile collection), but the Trinidad populations are rather variable, and some plants show a marked resemblance to subsp. padifolius of the Lesser Antilles. Britton and Britton 2194, Britton 2789, and Baker and Simmonds s n. are especially similar to padifolius in leaf shape and texture. Fendler 699 and Brit- ton, Hazen and Mendelson 1093 (the type of P. ierensis) have thin ovate leaves quite unlike those of padifolius; the type of P. ierensis has unusually large leaves (up to 16 cm. long). : The single collection from British Guiana is unusual In its sub-lanceolate leaves, which give it a quite distinctive aspect. The cyathia of this collection are similar to those of Mexican and Central American collections ; they are medium large (11-11.5 mm. long) glabrous or nearly so, and have spurs which project posteriorly and arise quite near the peduncle (at ca. 0.5-0.7 mm.). The staminate pedicels of this collection are apically puberulent, but not strongly so. The single collection seen from Surinam is without flowers, THE GENUS PEDILANTHUS (EUPHORBIACEAE) 147 and the leaves are badly broken, but it resembles the above collection vegetatively. The above discussion is manifestly incomplete and sub- jective. An adequate treatment of this taxon would require many different measurements in the cyathium alone and careful statistical treatment. Even then, the major gaps in material from Central and South America and the rather poor material from some North American localities would make such a study only preliminary. Nevertheless, a study of this sort would be of great value, especially if augmented by field work in South America. RELATIONSHIP. Within the species, the subspecies is closely allied to subspp. Smallii, parasiticus, padifolius and retusus; all of these are probably derived from subsp. tithy- maloides, or ancestral types which would fall within its rather wide range of variability. Pedilanthus tithymaloides seems clearly allied to P. nodiflorus in a number of features, and the two show some resemblances to the P. pulchellus alliance. NoTEs. The typification of Euphorbia tithymaloides L. is a troublesome matter, since there are no specimens of this plant in the Linnean Herbarium and no plates are cited by Linnaeus under typical E. tithymaloides. Croizat (1943) has taken Poiteau’s Plate 19 (1812) as neotype for the spe- cies. Unfortunately, this choice must be reconsidered. Pedi- lanthus tithymaloides has been traditionally applied to a plant with more or less cuneate leaf bases and puberulent staminate pedicels; the sort of plant which may be found in Central America or Colombia. From Dr. Croizat’s dis- cussion of Dugand and Jaramillo 2752 (Colombia), it would appear that he had no intention of changing the customary concept of P. tithymaloides. Poiteau’s figure rather clearly represents the plant which has been variously known as P. parasiticus, P. ramosissimus, P. itzaeus and P. latifolius (here treated as P. tithymaloides subsp. parasiticus). The Subcordate leaf bases, the small bract-like leaves below the inflorescence, the shortly exserted styles with widely spread- ing stigmas, and the locality, Cap Francais, St.-Domingue, all point strongly to this conclusion. Rather than take this poorly known (though easily recognizable) entity as type, T have chosen Jacquin’s plate 92 (1763), which is, historical- ly, a good choice, and which clearly represents the sort of 148 ROBERT L. DRESSLER plant which has been consistently considered to be P. tithy- maloides. To state that the Euphorbia tithymaloides of Lin- naeus represented ‘‘a mixture of all the species in this group known to Linnaeus and his predecessors” (Croizat 1943) is to color the issue somewhat; there is no evidence that any Pedilanthus other than P. tithymaloides, including subsp. padifolius, were known to Linnaeus or the earlier European writers This plant is widely cultivated as a hedge and medicinal plant in tropical and sub-tropical America. It is the sub- species most commonly encountered in botanical gardens and greenhouses. Several forms in which the leaves are varie- gated with white or white and pink are widely grown. The physiological or genetic basis for this pattern is not known. The leaves of variegated plants are unusually narrow, but branches of these plants occasionally “sport” to the normal green-leaved sorts, and these normal branches (in those which have been seen to “sport”) bear wider leaves which are typical of subsp. tithymaloides. 13b. Pedilanthus pees subsp. parasiticus (Kl. and Gke.) comb. nov. P. parasiticus Kl. and Gke., Abh. Akad. Berl., Phys. 1859: 105. 1860. Type: Nueva Espana, Herb. Pavon (G!, photo and tracing: FI, Sessé, Mocino, Castillo and goonies 1 if F! is able an isotype). P. ramosissimus Boiss., DC. Prod. 15(2): 5. 1866. — Tithymalo[i]des parasiticum (Kl. and Gke.) O. Kuntze uty Gen. Pl. 2: 620. 1891. — Tithymalus parasiticus (Kl. and Gke.) Croiz., Amer. Jour. Bot. 24: 704. 1937. P. itzaeus Millsp., Field Mus. Pub. Bot. 1: 305, Pl. 18. 1896. Type: Mexico, Yucatan, on cultivated lands, port of Silam, March to June, 1895 oe F.. Gaumer 452 (¥F 36255!). — Tithymalus itzaeus (Mills sp.) Croiz,, Amer. Jour. Bot. 24: 704. 1937. P. latifolius Millsp. and Britton, Ann. Mo. Bot. Gard. 2: 43. 1915. Type: Bermuda, Castle Point, waste grounds, Aug. 27 - Sept. 21, 1912, m. high, mi idrib not winged beneath, flowers salmon, S. Bro ast ai . L. Britton 820 (Ny!, isotypes: F!, PH!). — Tithymalus peta phyllus Croiz., pee Jour. Bot. 24: 704, 1937. Based on P. latifolius, not T. latifolius (C. A. Meyer) KI. and Gke. Shrub 0.8-2 m. tall, succulent; stem thick, sparsely crisped-puberu- lent when young, becoming glabrate, straight or slightly zig-zag; broadly ovate, rarely elliptic-ovate, apex acute or narrowly 0 obtu sot usually acuminate, base truncate or subcordate, 3-9 em. long, 2.5- THE GENUS PEDILANTHUS (EUPHORBIACEAE) 149 em. wide, branches with small bract-like leaves 0.7-2 cm. long and 0.5-1 em. wide often occur, midrib weakly keeled, petiole 2-4 mm. long, pad ag ngs ag stipules brown, blunt and spur-like or depressed, 0.5 ide and 0.2-0.5 mm. high; inflorescence terminal or eulary. “branches often partly monochasial by abortion, internodes 1-2.5 mm. long, glabrate or sparsely puberulent, bracts red, oblong long, 3-4 mm. wide, glab us ovate, te or acuminate, 4-7 mm sparsely puberulent within and usually sparsely puberulent with- ut, especially along midvein and margins, peduncle 3-4 mm g, glabrous; involucre pink, entirely glabrous except on margins, in volucral tube 7.5-11 mm. long, main lobes oblong, obtuse, fre r 2.5-3 mm. beneath, ciliolate on margins, accessory lobes ligulate, 1-2.5 mm. 0.6-1.1 mm. wide, obtuse, margins ciliolate, lateral lobes shorter than main lobes, free 1-2 mm. laterally and 3-5 mm. medially, . shorter than ponies lobes ; bcd cucullate, median lobe m arising en aeeed at 2-3 mm. from pedun 5- thick, pro- jecting 1-2 posteriorly and poneainns ‘faintly 3. Jobulate near g m. base, lateral lobes narrow, folded over accessory involucral lobes, edial completely connate, together attenuate-deltoid, m. wide at retuse apex, rior, ovate or stig 0. 8-1.2 mm. long, 0.5 glands ca. m. mm. wide; panties sh long), but distinct between medial glands and bases of accessory involucral lobes, ca. 0.4 mm. high; bracteoles absent or very few, filamentous, to 3 long, dorsal le-ri long, terminating in sho cels glabrous, 8-1 long, filaments 1-2 mm. long, gla Sparsely oe anthers glabrous; erate flower glabrous, pedisel 4.5-11 appa ovary i) ren — not elongating after pipes n diam ey style thick, 5.5- . long Representative specimens seen. pee Yucatan. Mérida, Schott 552 (F); Yucatan, Valdez 85 (F). ruz. San Andrés Tuxtla, cultivated as hedge, Dressler 1807 Pe British spouitenyiee Corozal Dist., Gentle 360 (micu). Jamaica. Parish of St. Thomas, Bank near Bath, Britton 3491 (Ny). 9 Arid mountains northwest of Bom- bardopolis, 610 m., E. C. and G. M. Leonard 13456 (us); La oat vicinity of Pikmi, Gonave Teland, Leonard 5196 (NY, US); Dept. l’Artibonite, Puilboreau road, vicinity of Ennery, 325-900 m., Léonawd 9711 (GH, US); Massif de la Hotte, western group, Jérémie, at Testas, escaped, Ekman 10787 (s). Santo Domingo. Sanchez, Rose, Fitch and Russell 4397 (us); Feb. 6, 1891, Wright, Parry and Brummel s. 1. (US). Puerto Rico. Local ng Lega (Rio oes to Fajardo), Britton, Britton and Brown 0 (NY); near house, Ubero, Mona Island, Britton, Cowell and ag ? 786 (NY); St. fake Midland, Mrs. J.J. Ricksecker 287 (F). 150 ROBERT L. DRESSLER DISTRIBUTION AND HABITAT (Map 7). Cultivated and per- haps native in the Yucatan Peninsula, apparently native in the Greater Antilles, in Jamaica and Hispaniola, and per- haps in Puerto Rico and St. Croix. Widely cultivated in Mexico and apparently cultivated in Bermuda and China as well as in the Greater Antilles. If native in Yucatan, the plant occurs in coastal thorn scrub similar to that occupied by P. nodiflorus (see fig. 24). The scant data from the An- tilles indicate dry, and in some cases rocky, sites. Apparently flowering mainly in winter (Dec.- April), but flowering sporadically in summer (July - Sept.) as well. COMMON NAMES. Mayorga, zapatito de la reina (Sierra de Tuxtla, Mexico, the second name, “the queen’s slipper,” is clearly a poetic name and not the “common name” in the area); zapatito de la Virgen, “the Virgin’s slipper,” vela de sebo, “tallow candle,” in reference to the thick, glossy, greyish stems (Guerrero, Mexico); yaxhalalché, Mayan: “green stem tree” (Yucatan peninsula) ; pie de nino, monkey fiddle (British Honduras) ; ipecacuana and variants (His- paniola). VARIATION. The variation pattern of this mysterious en- tity cannot yet be satisfactorily handled. The features of the involucre are relatively uniform except in size; the collection from Ile de la Gonave, Haiti, has very small in- volucres (7.5-8 mm. long), suggesting a size reduction parallel to that of subsp. angustifolius and its allies in the Greater Antilles. Faintly three-lobed spurs were most clear- ly noted in the material from Bermuda (where it is doubt- less introduced); the same feature is present, but less marked, in the specimens from Veracruz. Subspecies para- siticus is interesting for the frequent occurrence of free medial spur lobes, presumably, like the lobed spur apex, 4 primitive character within the species. This feature is of little taxonomic value, however, since it varies on a single plant. Plants of this subspecies are readily recognized not so much from leaf shape as from the succulent, glossy oF almost glaucous leaves and stems. As cultivated in Veracruz and Yucatan, the plants show two very different aspects, the Veracruz plants having dark green, nearly plane leaves and the Yucatan plants bearing leaves which are muc lighter in color and somewhat ruffled or crisped. _ plants from these two sources are grown together In the THE GENUS PEDILANTHUS (EUPHORBIACEAE) 151 greenhouse, they are vegetatively indistinguishable. The tendency to produce brarithes with small bract-like leaves has been noted only in cultivated material in the mainland of Mexico and in some specimens from Hispaniola. Whether or not other populations have this potentiality is not known. RELATIONSHIP. This subspecies is closely related to subsp. Smallii, and both are notable for being nearly glabrous in most parts. This feature appears to point to the Atlantic slope of northern Central America. The non-elongating pistillate pedicel of parasiticus finds a parallel in Honduran material of P. t. ssp. tithymaloides. It is quite possible that parasiticus and Smallii were derived from plants which reached the West Indies from Honduras or Nicaragua and that parasiticus has only recently been introduced into the Yucatan Peninsula by man. Notes. Though three times described as a new species, this plant has not been collected often, and adequate flower- ing specimens are few, indeed. Pedilanthus latifolius was clearly based on cyathia with free medial spur lobes, but this character is not constant, even for the type collection. The basis of P. itzaeus is less clear; if a normal plant were found which fit the description given by Millspaugh, it would well merit the “§ Anomalus” which Millspaugh used and later discarded. The type specimen is completely with- out cyathia, though the confusing description and even more in which most of the involucral bracts had developed on the accessory lobe pattern. The subspecific epithet is apparently that applied by Pavon as an herbarium annotation. It was wrongly credited, by Klotzsch and Garcke, to Boissier, who later re-named the species, commenting: “Nomen specificum Pavonii prob- abilissime improprium.” In the isolated Sierra de los Tuxtlas of southern Vera- cruz, this Pedilanthus is widely cultivated as a hedge and ornamental, the subsp. tithymaloides being seen only in its variegated form or an occasional green “‘sport.” The varie- gated Pedilanthus was referred to by one housewife as mayorga de lucir, or “fancy mayorga,” as contrasted with the ordinary mayorga of hedges. The common hedge plant 152 ROBERT L. DRESSLER of los Tuxtlas agrees well with the type specimen of P. parasiticus, as far as can be determined; it is quite possible that the type was collected there by Mocino, in 1793 or 1794, when he visited the then active volcano near San Andrés Tuxtla and spent some time in the town. Pedilanthus tithymaloides ssp. parasiticus has been seen in cultivation in Cuernavaca, Morelos and Iguala and Atoyac in Guerrero; cuttings from these areas failed to survive in the green- house. The collection from British Honduras is assigned to this taxon with some reservations; the plant, which is sterile, was rather damaged before collection, and the leaves were much crumpled in drying. 13c. Pedilanthus tithymaloides subsp. Smallii (Millsp.) comb. nov. P. Smallii cane Field Mus. Pub. Bot. 2: 358. 1913. Type: Florida, in pinelands, Miami, Nov. 1-30, 1904, J. K. Small 2286 (NyY!, photo and fragment: F!). — Tithymalus Smallit (Millsp.) Small, Man. S. E. Flora, 804. 1933. Shrub 0. ‘i 2 m. tall, stems puberulent when young, sige glabrate with age, each node ‘usually bent sharply in an obtuse angle, so that the stem te conspicuously zig-zag; leaves sparsely "plone pubernlent beneath and glabrate or very sparsely puberulent above, generally nd 1.38-3. i but b i long and 1.38- , but branches with sma ract- e leaves, mostly 0.8-1.5 em. long and 0.35-0.5 em. wide often formed, bleiisa elliptic-ovate e-ovate, e , apex acuminate, cute or narrowly obtuse, midrib keeled or winged, petiole puberulent, 1.5-5 mm. long, stipules dark brown, spur-like, blunt, ca. 0.5- in diameter and height; cymes terminal and axillary, congested, internodes puberulent, 1-2 mm. long, bracts red, oblong-lanceolate to oblong-ovate, cute or soot ae puberulent without and more or less sericeous within, 5-7 mm , 2- wide, peduncle glabrous, 6-9 mm, long; cyathia pink or red, ’ involucral tube glabrous through- ut, 8.5-12 mm. peters main lobes oblong or lance-oblong, obtuse, sparse- ly ciliolate on margins, free beneath for ca. 2.5 mm., accessory lobes ligulate or sukepatalate: ee SS ciliolate, margins spar rsely hispidulous above near base, 0.8-1 mm. wide, lateral lobe ca , 2-3 mm. shorter than main lobe, oa 1-1.5 mm. laterally, 5-6 mm. “medially, medial el 1.5-2 mm. shorter than lateral lobes; spur cucullate, sigs’ at ca. 1.5 mm. from peduncle, ca. 2.5-3 mm. thick, projecting m. _Posteriony, glabrous without, lobes basally hispidulous on and n argins within, lateral lobes folded in over accessory in- volucral eka, medial lobes connate, together 3.5-5 mm. long, basally 3.5-4.5 mm, wide, subligulate from a broadly a hase the ligulate portion 2-3 mm. long, slightly thickened, 0.8-1.1 mm. wide retuse apex, medial glands basal, thick, slightly ae cabquadirate to THE GENUS PEDILANTHUS (EUPHORBIACEAE) 153 ca. 20- glabrous, pedicels 9-11 mm. long, filaments 1.7-2.5 mm. long; pistillate flower glabrous, pedicel 6-10 mm. long, protruding Scarcely or not at all from the involucral tube, ovary oblong-ovoid, trigonous, ca. 1.5 mm. in diameter, style thick, 5-6 mm. long, branches 0.7-1 mm. long, ca. 1/3 bifid, spreading or connivent; fruit unknown; Seed grey, ovoid-subglobose, apiculate, base rounded, faintly keeled dorsally, 3.3 mm. long, 2.7 mm. wide, raphe dark brown. ¥,8 s K of Msticins playa, Britton, Britton and Shafer 156 (F, NY); Santiago de las Vegas, Prov. Havana, cultivated in gardens, van ann 507 (F, NY); Antilla, Oriente, hotel garden, Britton, Britton and Cowell 12512 (¥, NY). DISTRIBUTION AND HABITAT (Map 7). Southern Florida, from near Lake Okeechobee south to Meigs Key, and north- ern Cuba (Matanzas). Widely cultivated; specimens have been seen from Florida, Cuba, Mexico, El Salvador, Costa Rica, the Philippines, Java and China. The plants were probably carried from Cuba at an early date by Spanish Ships. The habitat in Florida is given as pinelands and ham- mocks. Flowering in winter (Dec. - March). COMMON NAMES. “Jacob’s ladder” (Florida), pie de nino, bitamo real, zapatillo, lechilla (Central America). VARIATION. As now sampled, this subspecies is relatively uniform in all its features. The stems are sometimes nearly Straight and the leaves vary in size, but no conspicuous variation has been noted in other characters. RELATIONSHIP. Though this subspecies is very different in aspect from parasiticus, these two entities are closely re- lated and differ mainly in leaf shape and texture, and in the more markedly zig-zag stems of Smallii. The involucral characters of the two are very similar except for the long-attenuate spur lobe of Smallii. This feature is also found in subsp. tithymaloides (Von Hagen and Von Hagen 1335) from Honduras, which further suggests that the an- cestor of Smallii and parasiticus had its origin in Central America. 154 ROBERT L. DRESSLER NoTEs. Plants of cultivated origin were transplanted to the greenhouse for further study, but all were destroyed by an infectious stem-rot, which also attacked some plants of parasiticus. The type specimen has suffered serious attrition since the photograph at Chicago Natural History Museum was taken ; the specimen is now without cyathia and nearly leaf- less. Some of the variegated forms of P. tithymaloides which are cultivated are referable to this taxon, but, since the variegation distorts the leaves to some degree, it is often difficult to assign these forms to a subspecies. 13d. Pedilanthus tithymaloides subsp. retusus (Benth.) comb. nov P. retusus Benth., Hook. shia Bot. and Kew Misc. 6: 321. (1854. Type: e Spruce 1469 (Herb. Benth. at K, sotypes Herb. Hook. at kK, G!, N P!, fragment F!). — Tithymaloti}Aes retusum (Benth.) O. Kuntze, Rev. Gen. Pl. 2: 620. 1891. Tithymalus melanopotamicus Croiz Am. Jour. Bot. 24: 703. 1937, vader on P. retusus, not T. re tusus (Cav.) Kl. and Gke. Shrub 0.7-1.7 m. tall; stem glabrate, apparently sparsely puberulent when young; leaves glabrous above, glabrate or sparsely puberulent _— cm. wide, obtuse and usually retuse, base cuneate, midrib carinate, petiole 3-5 mm. long, — dark, low, sub-hemispheric or transversely oblong, 1-1.7 m Sie 5-1 mm, wide; inflorescence terminal, on lateral branche xilla ary, internodes 3-5 mm. long, sparsely puberulent, bracts oblong orate “glabrate within, ee ——, especially along margins and mid-vein, 5-9 mm. lon 5-3 m. wide, sore pes) 4-7 mm. long, cchiesieni involucre clabions without, tube 8 mm. long, pilosulous within, ‘especially dorsally, main lobes tide. rer ciliolate, free eneath for ca. 2 mm., ac- cessory lobes thickened at ca. 2 mm. from medial glands, glabrous above except for aida margins, henints or Neulate. -spatulate, 1-1.2 mm. wide, obtuse, ciliate, lateral lobes 1-2 mm. shorter than main lobes, free ca. 1 mm. lat terally, 2-4 mm. medially, — lobe 1.4-1.5 mm. shorter than lateral lobes, free ail wis ee 5 ong; spur cucullate, arising at 1.5-2 mm. from tobe cle, -2 mm. thick, measurements given), owe deltoid, truncate, 3.5-4.5 mm. lo te 3-5 mm. wide basally, 1.2-1.4 mm. wide at apex, slightly thicken apically, sericeous-puberulent or hispidulous-puberulent beneath, THE GENUS PEDILANTHUS (EUPHORBIACEAE) 155 somewhat grooved medially on underside, medial glands basal, oblong or reniform-oblong, straight or slightly bent, 1-1.3 mm. lon 6-0.8 pilose terminally for 2-4 mm., filaments ca. 1.5 mm. long, glabrous; pistillate pedicel 7-9 mm. long, puberulent, ovary glabrous, oblong- void, ca. 2 mm. long and 1.5 mm. wide, trigonous, style glabrate, 5-5.5 mm. long, branches 0.5-1 mm. long, shallowly bifid; capsule subquad- rate, deeply 3-lobed, ca. long; seed ashen grey, oblong-ovoid Searcely keeled, basally truncate, apically acute but low, apex brown, raphe light brown on dark brown band, seeds ca. 4 mm. long and 2.5-3 wide Specimens seen. Peru. Rio Mazan near Iquitos, Dept. Loreto, 125 m., L. Williams 8133 (F); Iquitos, Dept. Loreto, 120 m., L. Williams 3567 (F). Brazil. Rio Negro, Martius s. n. (M); Para, sylvis fl. Amazon. conterminis, Maio, Martius s. n. (M); Para, ad Breves in ins. Marajo, Aug., Martius 2654 (M); Pernambuco (Tapera), cult., D. Bento Pickel 2129 (US DISTRIBUTION AND HABITAT (Map 8). Known only from scattered localities in Amazonian South America where it from Portuguese) : “Sapatinho: . . plant with caustic latex, sometimes used in folk medicine, not infrequent on farms [quintais], in Para and Amazonas. Origin not known with certainty, possibly from the northern part of tropical Ameri- ca.” Apparently flowering in autumn and spring (Sept., Oct., and April). COMMON NAMES. Sapatinho, sapatinho do diabo (Brazil) ; zapatito del nino (Peru). VARIATION. Little variation has been noted aside from the unusual, free medial spur lobes of Bento Pickel 2129. In this collection the spur lobes are uniformly free nearly to the ase. This same condition occurs sporadically in subsp. parasiticus, but is rarely consistent throughout a collection. Bento Pickel 2129 also has somewhat more oblong leaves than do the other collections seen. The type collection, which is brittle and perhaps shrunken, seems to have slightly smaller involucres than do the other specimens at hand. RELATIONSHIP. In some respects this taxon is similar to subsp. padifolius. It is not certain that the leaf shape indi- cates actual relationship, and the other features are also Shared with Venezuelan populations of subsp. tithymaloides. The geographically closer (as the ranges are now known) 156 ROBERT L. DRESSLER subsp. tithymaloides of Colombia and of the Guianas do not approach retusus so closely as do the Venezuelan plants, either in indument of the involucre or in leaf shape. 13e. Pedilanthus cai tea arte subsp. padifolius (L.) mb. nov agar piri rested loides B. eee L,, Sp. Pl. 1:. 458. 1763, af 9 A specimen in the Dillenian Herbarium (OxF, photo GH!) Pehamatsis baitocariantols ill., Gard. Dict. e — P padifolius (L.) Poit., s. Paris 19: 393 padifolia (L.) Haw., l. Suce. 136. 1812. Tithymalo[i]des padifolium (L.) O. Kuntze, Rev ozs 62 maloides var adr (L.) Griseb., Fl. Br. W. I. 52. 1859. — Tithy- malus (L.) Croiz., Am. t. 24s ifolius ur. Bo 703. laurocerasifolius (Mill.) L. C. Wheeler, Contr. Gray Herb. No. 124: 42. 1939. Euphorbia anacampseroides Lam., Encye. Meth. Bot. 2:420. 1786. Shrub 1-3 m. aa stems g labrous s, slender to mode — pers 5 em. 5 stipules spur-like, blunt, sometimes transversely oblong, in diameter; inflorescence terminal and upper fede occasionally i r 7 cm, ac on lateral branches, becoming rather large (to ross), internodes glabrate or sparsely puberulent, 4-10 m iota. bracts red, often tending to sist, glabrous or glabra cept on margins (and sometimes mid-veins) which are puberulent, ovate, oblong-ovate or obovate, obtuse, broadly acute or occasionally te, sometimes apiculate, 8-15 mm. long, 4.5-8 mm. wide; peduncle sparsely puberu- lent, 8-11 mm. long, involucral tube yellow- reen h or Tre n g throughout, glabrous without except on margins, internally sparsely pilose above and otherwise glabrous or (rarely) sparsely hispidulous- itne, 9.5-12 mm, egg ain lobes oblong or delto sg oblong, obtuse, ciliate or hispid-ciliate on margins, free beneat r 2.5-5 mm., ac- cessory lobes siealae lance-ligulate or pis ats 0.5-1 mm, wide, glabrous or basally sparsely puberulent govt: except along margins, i m. from the me : glands, terminally obtuse, ciliate or ciliolate, lateral lobes 1-2 mm shorter than main lobes, free laterally for 0.7-1.5 mm. and medially ian secethies ‘atttiiaale deleaad or deltoid, 3-5 mm. long, (3-)4-5 mm. wide, THE GENUS PEDILANTHUS (EUPHORBIACEAE) 157 apex truncate or slightly retuse, 0.9-1.2 mm. wide, thickened beneath for 1.5-2 -, occasionally sparsely hispidulous beneath behind the thickening, medial glands sub-basal, oblong, subquadrate or reniform, more or less bent about the partitions, 1-1.3 mm. long, 0.7-1 mm rca, 1 mm. posterior to the medial glands; bracteoles ew, filamen- tous or occasionally ligulate, glabrous or pilose, 4-8 mm. long, adnate to the involucral tube for ca. 1.5-3 mm. basally; roof lobules obscure occasionally small seta-like lobules; staminate flowers -2 ° ac f the ovary, ovary glabrous or very sparsely pilosulous, ca. 2 mm. long and 1.5 mm. wide, oblong-ovoid, style 5-9 mm. long, the branches 0.3-1 mm. long, shallowly or deeply bifid; capsule subquadrate, ca. 7 mm. long; seed ashen grey to mottled dark brown, sometimes darker about apex, 4-5 mm. long, 2.5-3.2 mm. wide, ovoid-oblong to oblong, truncate basally, acute but low apically, weakly keeled, the keels Sometimes terminating basally in low tubercles, raphe brown. Representative specimens seen. ser Antilles. St. Croix. Big Princess, Mrs. J. J. Ricksecker 165 (F, MO); Signal Hill, A. #. Rick- secker 181 (F, GH, MO, NY, US); East End, April 24, 1896, A. E. from medium altitudes to summit of Mt. Royal, Howard ee ae a DISTRIBUTION AND HABITAT (Map 8). The Lesser Antilles, including St. Croix, south to the Isle of Ronde and probably Grenada. Collections from Jamaica and St. Thomas are apparently cultivated or escapes from cultivation. An in- habitant of xeric and often rocky scrub and thorn forest. Howard (1952) writes, of its occurrence in the Grenadines: “Occasional . . . on the Isle of Ronde and Union where they grew in the Opuntia thickets or on rocky outcrops. These 158 ROBERT L. DRESSLER plants were less than 3 ft. tall. On Cannouan, however, this was a very common plant in the dry woodlands on the slopes of Mt. Royal. Here the plants reached ten feet in height. The “flowers” were usually red and yellow although occasionally pure red stands were observed. The leaves are succulent when fresh and commonly a bright red color.” Flowering mostly from mid-winter through mid-summer (Dec. - July). MON NAMES. Lady’s slipper, dumb-cane (Barbados) ; Bois damoiselle (Martinique) . VARIATION. Relatively uniform except for intergradation with angustifolius in the north (see fig. 29). and subsp. tithymaloides in the south. The variation in pubescence of the pistillate pedicel seems to show no geographic pattern, but there is a slight increase in indument of most parts of the involucre and flowers from north to south. Style length and seed size show increases from north to south. Of special interest are the projecting spurs of the collections from the Grenadines (slightly projecting on Union and markedly so on Cannouan and Ronde). Distinctly projecting spurs are found also in jamaicensis and bahamensis, suggesting a primitive character for this phyletic group, which has per- sisted only on the periphery of the Antilles. Here it might also be interpreted as gene flow from subsp. tithymaloides of Trinidad. RELATIONSHIP. It is intermediate in most features between subspp. tithymaloides and angustifolius, with which it in- tergrades to a surprising degree (considering the insular distribution) ; related also to jamaicensis. Notes. The specimen taken as type is annotated by Dil- lenius (as Tithymaloides laurocerasifolio non crenato [sic] H. Elth.), so that it may reasonably be considered the type for Tithymaloides laurocerasifolio non serrato, the pre- Linnaean basis for Euphorbia tithymaloides B padifolia L. and Tithymalus laurocerasifolius Mill. I have not seen the plate of Plumier which is the type of Euphorbia anacampseroides, but the locality, description and synonymy indicate that it is this plant. ; is is apparently a rather attractive shrub, but little cultivated outside of its native area. THE GENUS PEDILANTHUS (EUPHORBIACEAE) 159 13f. Pedilanthus paar g ay _ Sa Er ay (Millsp. and Britt.) comb. F. jamaicensis Millsp. and Britt., Field Mus. Pub. Bot. 2: 356. 1913. Type: Jamaica, Negril and vicinity, March 9-12, 1908, rocky woods, 4 dm. high, N. L. "Britton and A. Hollick 2067 (NyY!, isotype F!). — Tithy- malus jamaicensis (Millsp. and Britt.) Croiz., Am. Jour, Bot. 24: 704, 1937. P. Grisebachii Millsp. and Britt., Field Mus. Pub. Bot. 2: 361. 1913. Type: Jamaica, Kingston to Bath, Bull Bay, ote 14, 1908, bank, shoots 1 m., N. L. Britton 3677 (xy!). malus Grisebachii (Millsp. and Britt.) Croiz., Am. Jour. Bot. 24: Bb ih rub 0.4-2 (-4) m. tall; stems slender, puberulent when young, l lobes ligulate-spatulate, obtuse, ciliate, glabrous ‘to puberu rulent above, thickened at about 2-3 mm. from the medial glands, lateral lobes 0.5-1.5 pierre: t 1.7-2.5 mm. from ast .7-3 mm. in uiicknehs, isis elated hn near margins, lateral lobes , Ided in over accessory involucral lobes, medial lobes completely connate, together attenuate-deltoid, 2.7-4 mm. long, 2.5-3 mm. wide, apex sub-ligulate, -7-1 mm. wide, thickened and densely puberulent panaatk for ca. 2 mm., apex truncate or retuse, sometimes upturned, medial glands basal or ca. 1 mm. from base, oblong or oblong-ovoid, 1-1.1 mm. long, 0.6-0.9 mm. wide, ‘ieral glands ca. 0.5-1 mm. anterior to the medial tL esis or ees pateraibne: ca. 6 mm. lo ong; seed un nown Specimens seen. Jamaica. Negril, eotdi near lighthouse, March 1 ii, 1908, W. Harris 10,238 (F, NY, US); Negril, March 24, 1955, M. L. 160 ROBERT L. DRESSLER Farr s. n. (INSTITUTE OF JAMAICA); Manchester Parish, near Great Bay, 3 miles southeast of Alligator Pond, Proctor 11,957 (GH, INSTI- TUTE OF JAMAICA); Manchester Parish, 1 mile E.S.E. of Gods Well, north side of Round Hill, Proctor 11,940 (GH, INSTITUTE OF JAMAICA) ; near Bull Bay, Harris 9,645 (NY); near Bull Bay, Harris 12,117 (NY); without exact data, Distin s. n. (K) DISTRIBUTION AND HABITAT (Map 8). This plant is found along the south coast of Jamaica in rocky, arid thickets in areas of limestone outcrop. Flowering in spring and sum- mer (March, April, June and Sept.). VARIATION. The Jamaican populations of Pedilanthus tithymaloides show striking variation in several features. When only the Negril and Bull Bay populations were sampled (nearly the opposite ends of the island), there ap- peared to be two very distinct and isolated populations in Jamaica. Recent collections from intermediate localities in- dicate that there is a single, more or less continuous popula- tion, with strong clinal variation. The Bull Bay collections — have very small, narrowly lanceolate leaves, which are 10- 25 mm. long, and 2-3.5 mm. wide, and apparently caducous. The leaves of Proctor 11,940, from Manchester Parish, are similar but somewhat larger, ranging from 15-30 mm. long and 3.5-7 mm. wide. The plants from Negril have very much larger leaves, which are obtuse and broadly oblan- ceolate or sometimes nearly obovate; these are mostly 5-9 em. long and 1-4 cm. wide. The smallest leaves from these collections tend to be more nearly lanceolate. Farr s. n. has somewhat narrower leaves than the other collections from Negril (mostly about 2 cm. wide), and Distin s. n. also has relatively narrow leaves (1-2.5 cm. wide) which are mostly oblanceolate. The leaves of these western collections are relatively fleshy and persistent. The involucral tubes and styles of the Bull Bay popula- tion are somewhat shortened as in subsp. angustifolius (involucral tube 7.5-9 mm. long, style 4-6 mm. long) ; the pistillate flowers are markedly puberulent, the ovaries dense- ly so; and the involucres are puberulent externally. The indument of the involucres is intermediate in character be- tween that of angustifolius and bahamensis, being neither stiffly erect nor strongly crisped and appressed. The other collections of this subspecies have larger involucres and styles (involucral tube 9.5-11.5 mm. long, styles 6-8.5 mm. THE GENUS PEDILANTHUS (EUPHORBIACEAE) 161 long) ; the pistillate flowers of these are glabrous and the involucre externally glabrous or nearly so. In these features Farr s.n. (western end of the island) approaches the eastern samples a little more closely than do collections from the middle of the south coast. The involucres of this collection are sparsely puberulent without and the styles 6-6.5 mm. in length. It is evident that the clinal variation in leaf form is primarily found in the western end of the range, while the variation in involucral features is much the greatest in oo eastern end. A much larger sample of this plant is needed. RELATIONSHIP. This interesting and variable subspecies shows close relationship to subspp. padifolius, bahamensis and angustifolius. The westernmost population (Negril) resembles padifolius in its vegetative features. These plants may well be peripheral relicts which retain many features of the population ancestral to all four subspecies. In subsp. jamaicensis the indument of the involucre is intermediate in character between angustifolius and bahamensis, but in other features it is closer to bahamensis. The parallel east- west variation in involucre size in the Jamaican and Baha- man plants is striking. Notes. As noted by Wheeler, Cutak and White (1944), the original description of P. Grisebachii was drawn in large part from Puerto Rican material of angustifolius. 13g. Pedilanthus tithymaloides subsp. angustifolius (Poiteau) comb. nov. Plates XIV & XXI, Map 8 des Péres, au cap francais,” Poiteau s. n. (P!, fra P tithymaloides var. angustifolius (Poit.) Griseb., Fl W. I. 62. 59. — Tithymalo[i]des angustifolium (Poit.) O. Kuntze, Rev. Gen Pl. 2: 620. 1891 us sarissophyllus Croiz., A r. Bot 162 ROBERT L. DRESSLER dark brown, sub-hemispheric, ca. 0.5 mm. wide; cymes terminal and on short axillary branches, fa eee a 1.3-8(-15) mm. long, glabrate to puberulent, bracts puberulent or sparsely puberulent without, glabrate within, elliptic-ovate to oblong, acute to rane 5-10 mm. long, 2-6 mm, wide, peduncle appressed See ee long, involucral tube green beneath, 6- on vilbue neberuiene (rarely glabrate) within, glabrous or sparsely nilesent becca with lo: oblong-ligulate ¢ or sub- -spatulate, 0.7-0.9 mm. wide, obtuse, the lateral lobes 0.5-1 mm. shorter than main iabes, free 0.5-0.8 mm, laterally and 1-1.7 mm. medially, medial lobe ca. 1 mm. shorter than lateral lobes (rarely subequal); spur red, cucullate, at ca. 2.5-4 mm. from the peduncle, 1.5-2 mm. thick, glabrous or glabrate without, usually hispid- puberulent within, at least along lobe margins, lateral lobes folded in over accessory involucral lobes, medial lobes completely connate, together attenuate-deltoid, 3-4 mm. long, basally 2.8-4 mm. wide, apex truncate or shallowly retuse, 0. 9-1. 3 mm. wide, somewhat thickened, medial glands basal, oblong to subquadrate, slightly bent, 0.8-1 mm. long, 0.5-0.8 mm. wide, decaatonalty split into gland-pairs, lateral glands deltoid-ovate, ca. 0.5-1 mm. in diameter, near the media glands or 1-1.3 mm. reclaimed partition none or very slight; bracteoles ew or none, igure othe 3-5 mm, long, adnate to the involucral tube for 1.5-2 m roof lo bules minute, sometimes seta-like; niger flowers 20- oT, ‘pales pedicels 6- 8 mm. long, filaments ca long; pistillate pedicels sparsely to densely appressed pilose- -puberulent, 8- mm. long, 2 mm. wide, glabrous, style thick, 3.2-5 mm. long, the branches 0.35-0.5 mm. long, 1/3 to Stage bifid; capsule sub- oer 3-lobed in cross section, 6-7 mm. in diameter; seed oblong- oid to conic-ovoid, 3.5-5 mm. long, 2.5-3 mm. wide, weakly keeled dorsally and tiateraliy. basally mead apically acute but low, mottled brown-tan to grey-brown (ashen), raphe bro Representative specimens seen. Cuba. La Perla, Oriente, Aug. 6, 1913, (flowered in Vedado in April), Hno. Leén 5025 (NY); Maisi, Oriente, valley of Yumuri River, (coll. P. Matos) Hno. Leén 17935 (NY); La Laja, C. Wright s. n. (GH). Hispaniola. Haiti. Presqu’ile os Nord-Ouest, Mole St.-Nicolas, Morne Rouge, fifth terrace, Ekma 4477 (S); arid west banks of Mole River, 2 mi. So. of Mole St. Nice- las, E. C. and G. M. Leonard 13093 (NY, US); Presqu’ile du Nord- Ouest, re Port-de-Paix to Jean Rabel, Morne Palm'ste, Ekman 3556 (s, us) ; Aguin [Aquin?], Arndt 13 (PH) ; Port-au-Prince, Ekman 2108 (Ss). Dominican Republic. pina shy de Santiago, alt. 100-300 m., Abbott 1031 (NY, US); La Romana, Prov. of Sere ibe? 525 (NY); La Romana, Prov. Seibo, ay costero, La Caleta, Ekman 12080 (S, US). Puerto Rico. Salinas de Boquerén, Britton, Cowell and Brown 4607 (F, MO, NY, US); prope Salinas de Cabo Rojo, P. Santon 769 (GH, K, US); Buena Vista, Carolina, Johnston an venson 1397 (NY); Guanica and vicinity, Britton and Shafer 1865 (F, NY); Yauco, aA nderwood and Griggs 637 (Us); prope Pefiuelas, Stahl 10 ot (Us); 2 mi. E. of Ponce, Heller 6192 (A, F, GH, MO, NY, PH, US); E con Coamo River, Britton, Britton and Brown 6001 (F, sh THE GENUS PEDILANTHUS (EUPHORBIACEAE) 163 gunda, Shafer 2410 (F, Ny, US); Culebra Island, Britton and Wheeler 121 (NY, US). St. John. Lamosure, Britton and Shafer 619 (NY, us), St. Croix. Sandy Point, Paulsen 322 (c); Borgesen 133 (Cc); Raunkiaer 2578 (Cc) DISTRIBUTION AND HABITAT. From eastern Cuba through the Greater Antilles to St. John and St. Croix, largely coastal. Collector’s labels, “dry jungle,” “dense scrub,” “coastal thicket,” “rocky woods,” etc., suggest a dry scrub forest as the usual habitat for this subspecies; the Cuban localities are in relatively wetter parts of the Oriente, and may be less arid sites than those of most other collections. Though frequently reported from limestone soils, angustifolius ap- parently occurs in volcanic sites near Coamo Springs, P. R., (“eruptive rocks’) and is found in lateritic soil in western Puerto Rico. For a photograph of the last habitat, see Wheeler, Cutak and White (1944). Flowering in late winter and spring (Dec.-June). COMMON NAME. ipecacuana (Puerto Rico). VARIATION. Subspecies angustifolius is a relatively uni- form entity, in spite of its necessarily discontinuous range. Leaf shape varies considerably, being generally oblanceolate in Cuba and Hispaniola, but tending to be narrowly lanceo- late in Puerto Rico and to the east. The degree of pubescence is variable in most parts, but shows no clear-cut pattern, though the involucres of the eastern populations tend to be more nearly glabrous without. The only Cuban collection with seed (Leén 17935) shows seed about a millimeter longer than any other collection (ca. 5 mm. long as compared to 3.5-4 mm. long). The Cuban material is otherwise quite comparable to that from Hispaniola. RELATIONSHIP. Closely related to bahamensis and jamai- censis, as discussed under those taxa. The relationship of this entity to padifolius is especially interesting and critical to any classification of this alliance. In two features, leaf Shape and pubescence of staminate flowers, there is a sharp break between these two taxa (discounting the two collec- tions which are so exactly intermediate as to suggest F, hybrids). In involucral shape, these populations approach each other rather closely, and there seems to be a geographic trend within each one, suggesting gene flow or continuing 164 ROBERT L. DRESSLER differentiation. In angustifolius, the most striking features of the involucre are the small size and the proportionately long spur lobe. This plant strongly suggests a sort of “organ- neoteny,” like that described for P. tehuacanus, in which the development of the involucral tube, especially, is arrested. In order to compare the involucral forms of different popu- lations in a mathematical manner, measurements were made of main involucral lobes and spur lobes, both measurements being made from the apex of the spur (in retrospect, the base of the spur would provide a less subjective point). These measurements were made on 1-3 involucres from each herbarium sheet, depending on the material available. The ratios between these two measurements and the leaf length/ width ratios were averaged for each collection, and these ratios are graphed against distribution in fig. 29. The geo- graphic trend in involucral proportion is seen rather clearly, ot ; ee eee i : oo © .% " o¢ @ $ %» $ x LEAF LENGTH/ WIDTH 7 .. qT =f i SPUR LOBE/ INV. LOBE > Bere, » ¢ 0% & * ha * * | Lo ae: Fic. 29. Correlation of some characters of P. tithymaloides aoeret me in the West Indies. The abscissa represents geographic occurrence, wh mitt linear. The ordinates represent ratios as en Cae — collection is fro dia eral its locality by symbols: diamonds, ngustifolius; dots, P. t Regan ifol X. putative hybrids between senile peey padifolius; T, P. t. sickobs grrcenewed THE GENUS PEDILANTHUS (EUPHORBIACEAE) 165 and the intermediate nature of the two collections discussed on p. 169 is shown. 13h. Pedilanthus tithymaloides subsp. bahamensis (Millsp.) comb. nov. P. bahamensis Millsp., Field Mus. Pub. Bot. 2: 359. 1913. Type: Bahama Islands, Grand Turk Island, Waterloo and vicinity, rocky plain (pitted rock) south of town, where it is one of the common shrubs, C. F. and C. M. Millspaugh 9030 (F 287, 782!, isotypes: GH!, BME — i de inate bahamensis (Millsp.) Croiz., Am. Jour. Bot. 24: 4 Breet kaa 0.7-1.5 m. tall, branching from the base, the branches somewhat arching; stems succulent, glabrate; leaves r ree and finely puberulent above, eparpely © crisped- ar gee e long, 3.5-6.5 mm. wide, obtuse or acute (2), ¢ neate to an indis- tinct petiole ca. 1-1.5 mm. long, midrib strongly keeled (winged), stipules dark brown, sub- hemispheric c, ea. 0. in diameter; cymes terminal and on short axillary branches, intern cnr glabrate or ‘sparse- ly crisped-puberulent, bracts ovate or deltoid-ovate, obtuse or acute, glabrate or finely puberulent on both surfaces, 2.5-3 mm. long, 1.8-2 free beneath for ca. 1.5-2 m ciara lobes ligulate to ligulate- ic a obtuse and ciliate or ciliolate, lateral lobes 0.6 mm. shorter than m lobes to subequal, free laterally 0.7-1.7 mm., medially 2-3.5 mm., Median lol .o- mm. shorter than lateral ‘lobes; spur cucullate it c from peduncle, 2-2. hick, finely —_— within, lateral lobes more or long, 0.6- nag mm, wide, occasionally split into gland pairs, lateral glands ca. 0.5-1 mm. anterior to medial glands, oblong to oblong-ovate or deltoid-ovate, vag 1.2 mm. long, oo mm. wide, partitions variable, absent to ca. 0.7 m posterior to glands; brac- teoles none or few, filamentous, 3- 6 mm. ot distinct bracteole-ridges ca. 5 mm. tage dorsally in some (Andros), roof lobu 2 none cot i icels long and 1.5 mm. wide, trigonous, style glabrous or sparsely puberu- lent, 6.5-9.5 mm. long, branches 0.7-1 mm. long, shallowly to deeply bifid; capsule eg ca. 6 mm. long; seed grey-brown to 166 ROBERT L. DRESSLER dish brown, turgid sub-spheric to ovoid, obtuse basally, apex acute but low, weakly 3-keeled, raphe brown. Specimens seen. Bahama Islands. Deep Creek, Andros Island, Brace 5145 (F, NY); Deep Creek, Andros, I. J. and A. R. Northrop 693 (F, G, GH); Spring Point, Acklin’s Island, Brace 4244 (F, NY); Atwood [Sa- mana] Cay, Wilson 7410 (F, NY); Salt Cay, Turk’s Islands, C. F and C. M. Millspaugh 9392 (F); Grand Turk Island, Nash and Taylor 3761 (F, NY); Grand Turk Island, Waterloo and vicinity, C. F. an C. M. Millspaugh 9390 (F, NY). DISTRIBUTION AND HABITAT (Map 8). The Bahama Islands, collected or reported from southern Andros Island, Acklin’s Island, Samana Cay, South Caicos Island, and the Turks Islands. The habitat of this subspecies in the Turks Islands is described as “rocky plain;” a photograph taken by Mills- paugh (in Herb. F and NY) shows the plant to be locally abundant in open, xeric scrub with Opuntia and other low shrubs. On Andros, bahamensis was collected in “coppet,” a hardwood thicket or low forest. In both areas Pedilanthus evidently occurs on the eroded limestone which is typical of the Bahamas. Flowering in both winter and summer (Dec.- March, July and August). COMMON NAME. “Monkey-fiddle,” the stems being rubbed together by children to produce a high squeak, according to Millspaugh. VARIATION. This subspecies shows a very interesting pat- tern of variation ; the plants from Turks Islands and Acklin’s Island bear small involucres with very short tubes, very like angustifolius in form, while the Andros Island collections show large involucres with long involucral tubes, the form being comparable to that of subsp. tithymaloides or jamat- censis. The collection from Samana Cay is intermediate. The Andros collections have relatively slender stems, more evenly tapering spur lobes, ligulate accessory involucral lobes, and, correlated with the larger involucres, longer staminate flowers and styles. Turks Island collections have abruptly attenuate spur lobes and sub-spatulate accessory involucral lobes. Average involucre lengths for the small samples from each island, from east to west, follow: Grand Turk — 8.3 mm., Salt Cay — 9 mm., Samana Cay — 10 mm., Acklin’s — 8.5 mm., Andros — 11.5 mm. RELATIONSHIP. In leaf size and shape (as far as known) this subspecies is similar to jamaicensis (eastern popula- THE GENUS PEDILANTHUS (EUPHORBIACEAE) 167 tion), which it also resembles in the characters of the in- volucre; it is also quite close to angustifolius (especially the eastern populations). The Andros Island form closely ap- proximates western jamaicensis or subspecies tithymaloides in involucral size and shape. Both this subspecies and the western population of jamaicensis probably represent the peripheral persistence of primitive characters within the West Indian complex related to angustifolius. Notes. As described by Millspaugh, this would be a most remarkable plant. No evidence can be found, however, of the ‘‘siliceous transverse ridges” of the stem (the waxy cutin might well cause a high pitched squeak when the stems are rubbed together), the glands are 4 as in all the P. tithy- maloides complex, and small caducous leaves are clearly produced by normal growth, as shown by the fragments ac- companying Brace 4244 and Brace 5145, and Brace’s partly illegible notes (with Brace 5145). Pedilanthus tithymaloides, subsp. tithymaloides < subsp. Smallii Two collections have been seen, whose characters suggest that the plants are the result of hybridization between Smallii and different forms of subsp. tithymaloides (culti- vated or escaped). Each collection is briefly described below. A. Stem straight, succulent, glabrate; leaves glabrous above, puberulent beneath, ovate, long-attenuate, basally obtuse, 5-6.8 cm. long, 2.5-3.8 em. wide, midrib strongly keeled beneath; cymes terminal and axillary; involucral tube ca. 12-13 mm, long, glabrous ae glabrate, with a few hairs apically, filaments sparsely pilosulous; bracteoles few, filamentous, ca. 5-6 mm. long; pistillate flower glabrous, style 9 mm. long. Cuba, Havana, inter Marianao et la Lisa, May 4, 1914, in fruticetum (ad domum), Ekman 713 (s). While the glabrate involucres with attenuate spur lobes suggest Smallii, the straight stem, wide leaves, and the slightly greater indument within the involucre indicate that this plant may include subsp. tithymaloides in its ancestry, probably the “P. canaliculatus” form, widely cultivated in the Antilles. 168 ROBERT L. DRESSLER B. Stem moderately zig-zag, sparsely Babee leaves elliptic- ovate, puberulent above and beneath, 3-4.5 cm. long, 1.5-2.3 em. wide, roadly cuneate basally, acute but not ateiuate apically, midrib sérinicly keeled or winged; cymes terminal, peduncles sparsely puberu- lent, involucral tube ca. 11 mm. long, glabrous without and sparsely pilose within, spur lobe attenuate-deltoid, 3.5-4 mm. long, 3-3.5 mm. wide, apex thickened and hispidulous beneath, margins of spur lobes a bracteoles few, ca. 6 mm. long, pilosulous; staminate pedicels sparsely pilose apically, filaments glabrate; pistillate pedicel pilosulous-puberu- ent, style ca. 7 mm. long. uba, Santa Clara Prov., Milpa, Cienfuegos Bay, July 1941, Howard 5441 (GH, NY). In vegetative features this collection diverges from Smallii less than the first, except for the too strongly pu- berulent leaves and less tangible textural features. The hispidulous spur lobe, the puberulent pedicels, and the pilose involucral tube all indicate the influence of subsp. tithyma- loid es. These plants were collected in cultivated or disturbed areas, and a hybrid nature for both of them seems probable. Nothing is known, however, of the Pedilanthus which is probably native to the eastern two thirds of Cuba. It is reasonable to expect that this area should be the home of Smallii or of forms intermediate between that and para- siticus. As the greater part of lowland Cuba has been much disturbed over a long period, it is possible that unquestion- ably native Pedilanthus will never be found there. Pedilanthus tithymaloides, subsp. tithymaloides < subsp. parasiticus Though no herbarium specimens showing this parentage have been seen, there are some indications that hybridiza- tion between these subspecies may have occurred in the West Indies, since the introduction of subsp. tithymaloides. Some plants sent from Jamaica by Dr. DuQuesnay resem- bled the “P. canaliculatus” form of subsp. tithymaloides, but had shorter, broader leaves. The first few cyathia formed in each cyme of these plants bore free medial spur lobes, as are frequently found in parasiticus; cyathia formed later had connate medial spur lobes. Truncate spur lobes, his- pidulous both above and beneath, hispid-pilose staminate pedicels and sparse pilosity within the involucral tube are THE GENUS PEDILANTHUS (EUPHORBIACEAE) 169 all features of subsp. tithymaloides, which were shown by these plants. Unfortunately, all of the plants were severely attacked by the stem-rot which killed the plants of Smallii in greenhouse culture (in itself evidence of parasiticus in- fluence) before herbarium specimens had been made, and no vegetative material has been saved. The plant illustrated in Leén’s “Flora de Cuba” (3:126) resembles the plants discussed above; its leaf shape is nearly that of subsp. tithymaloides (“P. canaliculatus” form), but the bract-like upper leaves strongly indicate the influence of parasiticus. Pedilanthus tithymaloides, subsp. padifolius subsp. angustifolius The description of two similar specimens from St. Croix, which are evidently intermediate between subsp. padifolius and angustifolius, follows. Features which are similar or identical in both putative parents may be omitted. Stem glabbrate (apparently sparsely crisped-puberulent when young) ; leaves sparsely crisped- puberulent beneath and very sparsely a na i cm. n, ca. n nal, internodes 3- ng, § parsely aaa rulent, bracts red, semi-persistent, oblong or ot bane oblong, obtuse, apiculate, 8-24 mm. long, 5-9 mm. wide; peduncle 12-14 mm. long, sparsely crisped- uberulent, involucral tube 9-10 mm. long, glabrous without, Nici em pilosulous to sericeous within, accessory g eerste lobes hispidulous above near base, median lobe ca. 1.5 mm. ae eigen = 3 mm. thick, median lobe attenuate-deltoid, 4- 14.5 mm. wide, 0.6-0.7 mm. bel terminally, truncate, slightly ieee Shige for ca, 2 m nally, "iid ulous be- neath (behind ‘the thickened portion) ; Weaciaokes filame rs pilosu- lous, 3.5-7 mm. long, staminate flower rs ca. 24, pedice -10 mm. long, pilosulous terminally for ca. 3 mm aments sata but less densely so than wediets: pistillate pedicel ca. 10 mm. long, puberu- lent, o ovary glabrous, style 6.5-7 mm. long, branches ca. 0.8 mm. long; ca psule ca. 6 mm. long; seed oblong-ovad, grey, brown about apiculate tip, ca, 4, 5 mm. ‘tone and 2.8 mm. wide. These intermediate pie td vs are: St. Croix. Near East End, May, 1896, A. E. Ricksecker s. n. (F); Bassin, “aig 1896, A. E. Ricksecker 8... ( MO). The East End collection has somewhat larger leaves than the other and markedly larger bracts; the accompanying packet contains two nearly mature seeds, and the plant had 170 ROBERT L. DRESSLER clearly produced a number of capsules. The Bassin collection is in a younger stage, but shows one young capsule and a quantity of apparently normal pollen. Both plants are inter- mediate between the supposed parent types in nearly every character. The bracts in the East End collection are un- usually large for either subspecies. 14. Pedilanthus Millspaughii Pax and K. Hoffm. P. Mi Sia Pax and K. Hoffm., Fedde Rep. Spec. Nov. 19: 174. 1923. Type: Costa Rica, Miravalles, Brade 2302 ta gnaneiay aeneyee by war at Ber a — Tithymalus Millspaughii (Pax and K. Hoffm.) iz Eye “Stem thick, fleshy; leaves thin (when dry), glabrous, ovate, acute, 9-11 em. long, 4-7 cm. wide, attenuate to the petiole; cymes short, apical, bracts small, shorter than the peduncles, phe ca. 5 mm. long, glabrous; cyathium ay mm. long, tube glabrous without, ubescent within, long fiss ; spur 4 the length or the tube, spur hin red; lobes lightly re al the ‘lateral teeth larger; glands 2; staminate and pistillate pedicels pilos No material of Sua 2302 has been located, nor has any other collection which might be referable to this species been seen. It is probable that it is a distinct species, but it cannot be placed with any assurance without specimens. Most of the features, as described by Pax and Hoffmann, are strongly reminiscent of P. calcaratus; this species, however, has pilose peduncles and glabrous staminate pedicels. Whether or not the median spur lobes are connate is not stated, and what is meant by “lightly tridentate” spur lobe is unclear. Perhaps the spur lobes are similar to those of P. nodiflorus, but with larger lateral teeth ; or possibly there are two completely connate, retuse lobes. The authors indi- cate that this species may be related to P. Oerstedii, but the basis for this cannot now be determined; there may have been authentic material of that species at Berlin. EXCLUDED oR DuBiouS SPECIES Diadenaria oe Kl. and Gke., Monatsb. Akad. Berl. 1859: 254. 1859, nomen nud Pedilanthus Brittonii (Millsp.) Pax and K. Hoffm., Engl. and Prantl, Nat. Pflanzenfam. 2, 19c: 223. 1931. = = Cubanthus Brittonti Millsp., Field Mus. Pub. Bot. 2: 372. 1913. Pedilanthus cordatus one Linn. Syst. Veg. ed. 16, 3: 802. 1826. Based on the follow thus secihitnatice tices Steud., Nomenclator Botanicus ed. 2, 2: 282. — Euphorbia cordellata Haw., Misc. Nat. 185. 1803. — THE GENUS PEDILANTHUS (EUPHORBIACEAE) 171 Crepidaria ? cordellata Haw., Syn. Pl. Suec. 136. 1812. Assigned to Crepidaria with a query by Ha worth; there is nothing in the descrip- tion to indicate that it is a Pedilanthus Pedilanthus linearifolius Griseb., Men m. Acad. n. s, 8: 161. ea! = Cub oe linearifolius (Griseb.) Miléon ., Field Mus. Pub. Bot. 372. 191 = Euphorbia portinetholdes Denis, toate ‘Ties at str. Afriq ue 6. 1921. The latter is a somewhat unfortunate choice of epithet, since the plant does not resemble Pedilanthus. ri ,n and the description is hopelessly inadequate. The plant illustrated by “tga (1912) is P. eymbiferus and, though the seca origin is not given, was probably collected by Karsten at Tehua The status of P. Oerstedii cannot be decided until authentic or pina material is at hand. edilanthus pectinatus Baker, Jour. Linn. Soc. Bot. 25: 343. 1890. Doubtless a Euphorbia; the type could not be found by Denis, Euph. Iles Austr. Afrique 120. 1921. Pedilanthus subcarinatus Haw. ex Steudel, Nomenclator Botanicus ed. 2, 2: 282. 1841, nomen nudum SUMMARY Pedilanthus differs from Euphorbia by the strongly bi- lateral symmetry of the involucre. Fourteen species are recognized in the genus, including the questionable P. Mills- paughii, of which no specimens are available. The most widely known species, P. tithymaloides, is widespread and variable, with a number of geographic populations distin- guishable as subspecies, especially in the West Indies. In the Greater Antilles two of these subspecies (angustifolius and parasiticus) seem to form the two ends of an overlap- Ping circle of subspecies and interact as distinct species, but there is no evidence that the two are narrowly sympatric. The thirteen species which are represented by herbarium Specimens are arranged in five interrelated species groups. Sympatric species pairs are few, even between species groups. The West Indian plants of the “section Cubanthus”’ are to be excluded from Pedilanthus, being derived from a different ancestral group (Euphorbia subgenus Esula). All species (except Pedilanthus Millspaughii, of Costa Rica) are Mexican or Mexican and Central American, and P. tithymaloides ranges into peninsular Florida, the West Indies and South America. The plants are usually local and 172 ROBERT L. DRESSLER discontinuous in distribution and often occur on areas of limestone outcrop. While a few species are mesophytic trees with large foliage leaves, there is a wide range of habitat and vegetative specialization, some species being low, suc- culent desert shrubs with small caducous leaves. Morphological features indicate that Pedilanthus is de- rived from a member of Euphorbia subgenus Agaloma, which is characterized by petal-like appendages on the in- volucral glands. The spur, largely formed of these gland appendages, is the most distinctive feature of Pedilanthus. This structure is thought to have developed as an adaptation to hummingbird pollination, and its possible evolution is sketched on p. 80 The distribution of primitive Ewphorbieae and the prob- able geologic occurrence of hummingbirds suggest that the evolution of Pedilanthus has largely occurred during the Tertiary, probably in Mexico. The dispersal of the genus into South America and the West Indies is thought to be not older than late Pliocene. The relatively few species of Pedilanthus are highly diverse in involucral structure, and it is suggested that a relatively rapid adaptive radiation has occurred since the development of a spurred involucre. In certain cases it appears that the evolution of the involucre has occurred through neoteny, or the developmental retarda- tion of certain characters. Adaptation to xeric environment has occurred in several lines within the genus, and a mech- anism is suggested through which relatively rapid reduc- tion in leaf size might be accomplished. An aneuploid series of chromosome numbers is found in the genus, involving haploid numbers of 16, 17 and 18. It is suggested that this genus evolved from a Euphorbia with a haploid number of 14 (a base number of 7 is characteristic of Euphorbia sub- genus dgaloma). The wide range in habit shown by Pedilanthus is paral- leled by its anatomical features. The more marked xero- phytes, especially, possess a number of the structural adap- tations which are usual in desert plants: small, caducous leaves; heavy cutin; a waxy coating over the cutin; sunken stomata ; thick chlorenchyma in the stem cortex, often with a palisade arrangement; and, in one species, a thickened storage root. A system of abscission layers at the nodes THE GENUS PEDILANTHUS (EUPHORBIACEAE) 173 LATE XIII. Fig. 30. Pedilanthus ceils a plant cultivated at Missouri Botanical rifi Garden, cyathia, showing the inflated spur and small without free s spur lobes, twice life — photo by C. H. Thompson, vides 1907, courtesy of souri Botanical Garden. Fig. 31. P. calearatus, Dressler 1067, in eultivation, cy thi a. 2.5 S e spur orifice (two drops have also life size, note the drop of nectar exuding from by J. W. Carmichael. flowed down to the base of the staminate flowers), photo ROBERT L. DRESSLER XIV. Fig. 32. Pedilanthus tithymaloides subsp. angustifolius, near Villa Is: pang Snag! Cristi Prov., Dominican Republic arch 1956, slightly enlarged, from Mrs. Antis H. Wagner. P. t. subsp. jamaicensis (eastern a g ubst Jamaica, January 1956, about 2.5 times life size, from 7 Wood pe ‘os Dr. R. A. Howard. THE GENUS PEDILANTHUS (EUPHORBIACEAE) 175 permits ready shedding of injured or non-functional parts. The wood structure is especially interesting and has been relatively well sampled. Extensive secondary xylem is formed by some species, but wood formation is markedly limited in the extreme xerophytes. The number of vessels per unit area and, especially, the vessel diameter are closely related to the habitat and the presumed water loss through transpiration. The large-leaved trees of tropical deciduous forest possess many large vessels, while both the true meso- phytes and the xerophytes with reduced leaf area show smaller and usually fewer vessels. The moderately succu- lent plants, such as P. bracteatus and P. tithymaloides, ap- pear to retain the most primitive features in wood structure: many solitary vessels, scalariform intervascular pitting and Slightly heterogeneous rays. Pedilanthus pulchellus, a meso- phyte with narrow, very long vessel elements and indistinct radial multiples may show the most primitive features among the truly woody species. The anatomical features, as far as observed, do not contradict the classification of- fered, and often support the relationships hypothesized. Since many of the anatomical features are markedly adap- tive, anatomical similarities between plants of similar habit and habitat, as between P. bracteatus and P. tithymaloides, aL not necessarily indicative of close phylogenetic relation- ship. The inflorescence of Pedilanthus, as of most Euphorbieae, is a compound dichasium of involucres. The involucre has the same basic structure as that of Euphorbia. There are five involucral bracts, each subtending a cluster of stam- inate flowers. Each bract shows a normal foliar three-trace pattern of vascularization, with the lateral traces vascular- izing the glands, when these are present. The glands are double structures, and occasionally develop as separate gland-halves. The glands bear petaloid appendages which, in Pedilanthus, are highly modified to form part of the dorsal spur. The strong zygomorphy of the genus involves the development of three types of involucral bracts: two ventral “main lobe” bracts which form the greater part of the invo- lucral tube above the level of the glands, and three ‘‘acces- sory lobe” bracts, of which the median member is bilaterally symmetrical and the two lateral ones are not. The “acces- 176 ROBERT L. DRESSLER sory lobe” bracts contribute largely to the formation of the spur and form inconspicuous ligulate lobes above the level of the glands. Within the involucre, the five clusters of staminate flowers are usually separated by more or less laciniate bracteoles. These bracteoles are thought to be derived from two types of structure: the bracts which sub- tend the branchings of the staminate flower cluster and the incurved and connate margins of the involucral bracts. These bracteoles often form solid partitions basally, and the partitions which divide the gland chamber of the spur in some species are thought to be developmentally related to those of the involucral tube. The clusters of staminate flowers are extremely condensed, and there is still con- troversy concerning the branching of these structures. The vascular pattern strongly suggests that each cluster is basically dichasial with each branch forming a monochas- ium throughout. The primitive type of inflorescence for the Euphorbieae was probably that still found in a few relict genera: one in which the involucral bracts strongly enfold their respective dichasial clusters of staminate flowers and the pairs of marginal glands are borne between these clusters. The development of the involucre of Pedilanthus agrees closely with that of Euphorbia in its earlier stages. The pollen of most species of Pedilanthus is three-colpate, with large oblong pores and reticulate exine. The pollen of P. tehuacanus is completely inaperturate, while that of P. Finkii has no pores, and narrow furrows with greatly thickened margins. A pattern of six longitudinal intine thickenings is found in Pedilanthus and appears to be characteristic of most woody Euphorbieae. ; While no caruncle is present on the mature seed of Pedi- lanthus, a caruncle or caruncle-like structure develops from the funiculus and appears to serve a nutritive function dur- ing seed development. This structure withers and stays on the columella when the seed matures. PLATES XVI - XXI The drawings are uniformly numbered as follows. 1. cy- athium, lateral view; 2. cyathium, dorsal view; 3. dorsal view of spur, cut open to show gland chamber; 4. leaf out- line; 5. bract outline; 6. as indicated. THE GENUS PEDILANTHUS (EUPHORBIACEAE) 177 Jeti, ty PLaTE XVI. A. P. Palmeri, based on a 1064, living materi B. P. Fin based on C. H, T son s. n. C. P. calcaratus, based on Dressler poten living material, hom 6. longitudinal section of spur 178 ROBERT L. DRESSLER P. gracilis PLATE XVII. P. gracilis, based on Hinton 10973, 6. habit sketch. THE GENUS PEDILANTHUS (EUPHORBIACEAE) 179 ie A opetetutlaihee 2202 PLate XVIII. P. pulchellus, based on Conzatti, Reko & Makrinius 3106, 6. habit etch, sk 180 ROBERT L. DRESSLER = Ee cvmbiferus.. \ SY ae Sy LS LY ce NY SN fh facet SENS _ om (es “2 NS C 6 fe 4 Gt P macrocarpus _| Pate XIX. A. P coalcomanensis, 1-8 & 5 based on Hinton 12685, 4 on Hinton 15130. B. P. eymbiferus, based on Dressler 1368, material preserved in liquid. C. P. macrocarpus, based on Dressler 1090, living material and specimens preserved in liquid. 4’, a leaf from Gentry 7767a; 6, cross section of leaf. THE GENUS PEDILANTHUS (EUPHORBIACEAE) 181 e P tehuacanus PLate XX. A. P. bracteatus, based on Dressler 979, living material. B. P. tomentellus, 1-8 & 5 based on Pringle 4912, 4 based on Dressler 1375. C. P. tehuacanus, 1-3, 5 5 & based on Dressler 1795, pres oe in liquid, 4 based on Purpus 7065. 6, lateral view of cyathium, with eyme rents in place. 182 ROBERT L. DRESSLER yt + P. tithymaloides subsp. tithymaloides 5 P tithymaloides — C subsp. angustifolius LATE XXI. A. P. nodiflorus, based on Dressler 1362, living material, B. P. t. SSP tithymaloides, based on Dressler 1376, preserved in liquid, 4’ a small, bract-like leaf from the same plant. C. P. t. ssp. angustifolius, 1-3 & 5 based on Britton, Cowell Brown 4607, 4 based on Leén 5025. THE GENUS PEDILANTHUS (EUPHORBIACEAE) 183 LITERATURE CITED ma ae W. 1912. Zur Embryologie einiger Euphorbiaceen. Trav. s. Bot. Acad., St. Pét tersbourg 9: 136-154. propa A. 1954. Contribution 4 la détermination des Euphorbiacées a méthode anatomique. Bull. Soc. d’Hist. Nat. Toulouse 89: 157-194. BalLey, I. W. 1944. The development of vessels in angiosperms and its significance in morpho logical research, Am. Jour. Bot. 31: 421-428. ——. 1956. The relationship between Spaomivenaa of New Caledonia and ae of New Guinea. Jour. Arn. Arb. 37: 360-365. ae PPER. 1918. Size variation in tracheary cells: I. A aaa Hise between the secondary xylems of vascular ecryptogams, Symnosperms and angiosperms. Proc. Am. Acad. Arts and Sci. 54: 149-204. BAILLon, E. H. 1858. Etude Générale du Groupe des Euphorbiacées. Paris, 684 pp. BANERJI, I. 1951. Pollen and embryo-sac of two Euphorbiaceae. Proc. Ind. Acad. Sci. B 34: 172-1 BarTLetTt, H. H. 1935. A method of procedure for field work in tropical American phytogeography based upon otanical reconnaissance in parts of British ee eee and the Peten forest of Guatemala. Carn. Inst. Publ. No. 461, 1- BEEKs, R. M. 1955. Improvements in the squash technique for plant chromosomes, El Aliso 3: 131- BEILLE, L. 1902. Recherches sur de ‘développement floral des Disciflores. Thesis, Bordeaux, 177 pp. BENE, F. 1947, The feeding and Agr behavior of hummingbirds. Mem. Boston Soe. Nat. Hist. 9: 399-4 Bonp, J. 1950. Results of the Gathered. Chaplin West Indies Ex- pedition, 1948. Part II. Birds of Cayo Largo (Cuba), San Andrés and Providencia, Proc. Acad. Nat. Sci. Phila. 102: 43-68. Brown, W. L., JR., AND E. O. WILSON. 1956. Character displacement. Syst. Zool. 5: 49-64, oe A. A. 1938. Pedilanthus versus Tithymalus. Kew Bull. 1938: 68-470. CLENCH, W. J. 1938. Origin of the land and freshwater sia fauna of the Bahamas, with a list of the species occurring on and Little San Salvador Islands. Bull. Mus. Comp. Zool., Harv. 80: 481-5 CROIzAT, a. 1937. Tithymalus or Pedilanthus? Nomenclatural con- siderations, notes, new names and combinations. Am. Jour. Bot. 24: 2-704. 38a. Notes on the Se with a new genus of the Euphorbieae. Philippine Jour. Sci. 6 7-412. ee 1938b. Glands of ott ecirtris and of Euphorbia. Chron. Bot. 12-514 MEA A. 40, On the phylogeny of the eh emer and some of their presumed allies. Rev. Univ., Chile 25: 205-22 ———~, 1943; On thé guatarsaile position of Pianta and its allies. Lingnan Sci. Jour. 20: 79-103. 184 ROBERT L. DRESSLER 942. Peculiarities of the inflorescence in the Euphorbiaceae. oe, rig 103: 771-779. 43a. Euphorbiaceae cactaceaeque novae vel criticae colom- bianae. — I. Caldasia 2: 123-139. —. 19 otes on American Euphorbiaceae with descriptions of 11 new species. Jour. ig h. Acad. Sci. 33: 11-20. DARLINGTON, C. D., AND A. P. ioe 1955. Chromosome Atlas of Flowering Plants. fae 519 DUCKE, 1946, ntas de eeticte precolombiana na Amazonia Braz ileira. Notas sdbre as espécies ou formas espontaneas que supostamente thes teriam dado origem. Bol. téc. Inst. Agron. norte ERDTMAN, G. 1952. Pollen Morphology and Plant Taxonomy. Stock- holm, 539 9 pp. Esau, K. 1953. Plant Anatomy. New York, 735 FAEGRI, K., AND J. rea . 1950. Textbook of irodarn Pollen Analysis. Copenhagen, 168 GAUCHER, L. 1902. "Recherches anatomiques sur les Euphorbiacées. GENTRY, H. S. 1942. 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Mex. 14: 37-67. Sa AND F,, MIRANDA, 1942 2, Nombres qclan de plantas en el S. e estado de Puebla. An. Inst. Biol. Mex. 13: 493-498. oe Rouuins, R. C. 1952. Taxonomy today and tomorrow. Rhodora 54: 1 Rose, J. N. 1895. Report on a collection of plants made in the rope of Sonora and Colima, Mexico by E. Palmer in the years 1890 an 1891. Contr. U. S. Nat. Herb. 1: 293-366. 186 ROBERT L. DRESSLER Scumipt, H. 1907. Uber die Entwicklung der Bliiten und Bliitenstande von Euphorbia und Diplocyathium. Beih. z. bot. Centralbl. 22: 21-84. SCHOUTE, J. C. 1937. On Be aestivation in the cyathium of Euphorbia fulgens, with some remarks on the morphological ee Bos of the cyathium in general. Rec. Trav. Bot. Néerl. 34: its 938. The cyathium glands of Euphorbia. Chron. 3 5 4: 30-32. SCHWEIGER, J. 1905. Beitrige zur Kenntnis der Samenentwicklung der Euphorbiaceen. Flora 94: 339-379. SHREVE, F. 1951. Vegetation of the Sonoran Desert. Carn. Inst. Wash. Publ. No. 591, 1-192. Smmpson, G. G. 1950. 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The subspecies concept sar tet applicatio a oye Zool. 2: 97-111 Weotek H. 1845, Miahaicecks Beitrage. 2. Infloreszenz von Euphor- bia. Flora 28: 452-454. 187 INDEX abscission layer, 24-25 adaptive radiation, 86-87 biotic relationships, 67-68 common names, 7-8, 101, 108, 114, AIT, 1225 127, 131, 136, 144, 150, (153, 155, 158, 168, 166 Rractucta: myrtifolia, pedilanthoides, 171 ti aloides, 1 7 ome hati 119 hist hybriiizetion between species, 75- Baap rin es within P. tithyma- 67-170 feliceecmce 25-26 involucre, development of, 41-45 mbna peas in, 83-84 ru 3 3-24 materials and methods, 8-10 measurement, 23-24 m, 8 rtitions and bracteoles, 39-41 Pedilanthus Finkii, boa hihg a 105 er Greg Grisebachii, laurocerasifoling, 156 linearifoliu av 1 dairener 171 peritropoides, 99 188 INDEX personatus, 134 Pavonis, 125 petraeus, 139 usum, 154 Pringlei, 13 Tithymalus, 97 pulchellus, 111 aztec 125 Purpusii, bahamensis, 165 ramosissimus, 148 bracteatus, 125 retusus, 15 calcaratus, 105 pe ssp. ja ssp. ssp. ssp. 8 ssp. 152 ssp. tithymaloides, 138 tomentellus, 130 photoperiod, 64- 65 pistillate flower, 55-59 pollen, 49-55 pollination, 68-70 ri e and advanced features, 81- 82 root structure, 11- specific seintecnalites, 73-75 staminate inflorescence, 46-49 — structure, 13-15 xonomic oe 94-97 Tithymaloides 03 GN age 105 Houlletii, 139 involucratum, 125 parasiticum, 148 Gre ae Grisebachii, 159 ierensis, 139 itzaeus, 148 jamaicensis, 159 koilopremnos laurocerasifolius, 156 acradenius, macrocarpus, melanopotamicus, 154 orus, 134 Olsson-Sefferi, 126 padifolius, 156 = parasiticus, 148 105 ee 161 Smallii, 152 spectabilis, 125 subpavonianus, 125 tehuacanus, 132 rama 138 tomentellus, 1 wood structure, 15-23 xeric adaptation, 84-86 CONTRIBUTIONS FROM THE GRAY HERBARIUM OF HARVARD UNIVERSITY Edited by Reed C. Rollins and Robert C. Foster NO. CLXXXIII THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA (COMPOSITAE) By Joun H. BEaAMAN PUBLISHED BY THE GRAY HERBARIUM OF HARVARD UNIVERSITY CAMBRIDGE, MASS., U.S.A. Issued 31 Oct., 1957 JOHN H. BEAMAN ABSTRACT e genus Townsendia (tribe Astereae) consists of perennial, biennial, ur in the western United States, Mexico, an and embryological study. Meiosis in both micro- and megasporogenesis in diploid, sexual plants = 9) is regular with the formation of tetrads of micro- and spores. At anthesis the pollen grains are 3-nucleate, and the percentage of aborte s is very lo evelopment of the female Mei oth micro- and megasporogenesis in polyploid, apomictic plants (2n = 27-36) is irregular with many or all of the chromosomes occurrin i II of normal meiosis, an yad with unreduced e tw halves of the dyad develop separately into 3-nucleate pollen grains that are conspicuously larger than those produced by di hesis ins i required to initiate development of the embryo or endosperm. No diploid plants are known to be apomictic; similarly, no polyploid plants are known apomicts have not been given special taxonomic recognition. They have been referred to species known also from one or more sexual geome tions. Apomictic plants are concentrated at the higher elevations an higher latitudes relative to whole range of the species in whic cars occur. Apomixis is found most commonly in species with the perennial habit; however, it occurs also in some biennials. : ybridization apparently has been extensive in the genus. Intermediate i i av n found. Certain apomictic races in six species exhibit morphological evidence of hybridity, but other apomicts THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 3 divergence. Polyploidy apparently has not been a mechanism of evolution, as it seems to be closely associated with obligate apomixis. Several examples of parallel or convergent evolution have been discussed. INTRODUCTION A cytotaxonomic study begun with several seed collections was the forerunner of the present paper. Early in the study it became evident that apomixis might be present in Townsendia. In the limited herbarium material then examined, several of the specimens seemed difficult to determine. Thus, it appeared that Townsendia was deserving of a full-scale systematic investigation, even though it had been revised by Larsen (1927). From several standpoints Townsendia has proved to be an ideal genus in which to employ some of the more recently de- veloped taxonomic techniques. The western North American distribution of its 21 species offers certain practical advantages for a biosystematic investigation. Field studies within the ranges of all the species have been made. Some of these have been necessarily brief, but the plants are within a convenient distance for further population studies of a more detailed nature. The con- centration of herbarium material of Townsendia is in American herbaria and a large percentage of this has been examined. With the recent interest in collecting in the western United States, most of the species have become well represented by herbarium speci- mens. A knowledge of intraspecific variation is therefore pos- sible, and, where feasible in the present systematic treatment, an attempt has been made to note the type of population diversity that has been encountered. The small size of the plants of most of the species frequently has permitted collectors to include sev- eral plants on a single herbarium sheet. Some herbarium speci- mens approach being “mass” collections. Abundant plants in single collections have facilitated the understanding of intra- population variation. The position of Townsendia in the Astereae is a fortunate one from the comparative cytological standpoint. Relatively few in- vestigations of a cytological nature have been made in this tribe of the Compositae. Erigeron and Aster are the only genera of the tribe other than Townsendia that have received more than cursory cytological attention, while genera in some of the other tribes of the Compositae, especially in the Cichorieae, have been the sub- jects of considerable cytotaxonomic and cytogenetic interest. The chromosomes of Townsendia are large and easily prepared for observation. Although chromosome morphology has proved of 4 JOHN H. BEAMAN Townsendia. In Townsendia, studies on the evolutionary significance of polyploidy appear to have little promise because of the strong correlation of polyploidy with obligate apomixis. However, the opportunities in Townsendia for research on the causal aspects of polyploidy and apomixis have not been exhausted. The present investigation embodies an attempt to bring to- gether as much information about Townsendia as is now avall- able. It is hoped that the systematic treatment will provide a solid foundation upon which future investigations may be built. Obviously, all of the problems associated with Townsendia have not been solved. But if this study can serve as a reference point from which problems in plant migration, population variability, and phylogenetic modification can be attacked, then a primary aim will have been fulfilled. ACKNOWLEDGMENTS The inspiration from Dr. Reed C. Rollins has contributed much to which aided funds, facilities, and material for the investigation has been pone" td Appreciation is extended also to Dr. Marion Ownbey for his advic when the study was in its early stages. THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 5 I wish to thank the curators of the herbaria from which specimens have been used. These herbaria are listed at the beginning of the systematic treatment. Most of the field investigations were financed by a generous grant from the Fernald Fund of Harvard University. To Mr. F. W. Hunne- well of Wellesley, Massachusetts, a large measure of appreciation is due for providing the Fernald Fund. I am indebted to Mr. Irwin Lane and Mr. Henry Andreozzi for handling the collections of living plants which were sent to them while the field studies were in progress. A grant from Sigma Xi has helped cover the expense of the illus- trations. Dr. R. E. Schultes and Dr. R. C. Foster have edited the Latin diagnoses and Mrs. L. Schwarten has given assistance in the library. Dr. S. J. Preece provided fixed material from the experimental garden at the State College of Washington which has formed an im- portant part of the embryological investigations. The assistance in the the field studies given by Dr. Preece in 1952 is also gratefully ac- knowledged. Seed collections have been received from Mr. J. T. Howell, Dr. Quentin Jones, Dr. Marion Ownbey, Dr. C. L. Porter, and Dr. B. L. Turner. I am indebted to Dr. W. A. Weber for guiding me to an alpine habitat of the rare Townsendia Rothrockii. Mr. and Mrs. W. L. Bar- tholomew have provided detailed field notes and abundant seed and ee material of the species of Townsendia in the Uinta Basin of Utah Dr. R. B. Channell, Mr. E. Hernandez X., and Mr. D. R. Whitehead gave much-needed help with the multitude of details which were attended to in the final preparatory stages of the paper. The devoted assistance of my sister, Mary Lou Beaman, in the summer of 1956 made possible a timely completion of the study. HistoricAL ACCOUNT Some details in the history of the genus Townsendia have been recorded in the revision by Larsen (1927). In addition to two new species in the present treatment, three species (T. de- jecta, T. minima, and T. anomala) have been described since the time of Larsen’s treatment. One of these, T. minima, is recognized as T. montana var. minima in this study. The other two are placed in synonymy. Botanists who have done the most work with Townsendia have generally considered the genus a difficult one. Thus Gray (1880) stated: “Ever since the discovery of a considerable number of species, this very characteristic genus of the Rocky Mountain region has been particularly difficult.” And Marcus Jones (1893) in his Notes on Townsendia stated: “This genus has always been 6 JOHN H. BEAMAN a trying one to me because the descriptions have not fitted the plants as they grow.” Heiser (1948) noted that “A number of problems in the genus Townsendia call for extensive field work and experimental studies.” The recognition of apomixis and its effects in the genus has been a great aid in solving problems which confronted and to some extent confounded previous in- vestigators. The taxonomic treatment of the present paper corresponds more closely to that by Gray than to the revision of Larsen. Im- provements on Gray’s treatment have been possible mainly be- cause many more specimens are now available. The present study indicates that only three of the 17 species recognized by Gray cannot be maintained. Generally, the varieties he described are not recognized in this treatment, but one of his varieties, T. seri- cea var. leptotes, was elevated to specific rank by Osterhout and is here recognized as such. The revision by Larsen, although based on more specimens than were available to Gray, shows a lack of understanding of Townsendia. Larsen’s confusion of T. mensana with T. Hookeri (T. sericea in her treatment), of T. condensata with T. spathulata, and of T. Fendleri and T. annua with T. strigosa has caused considerable difficulty for botanists working with the flora of the western United States. F urther- more, her key is not satisfactory for determining many of the species. One helpful contribution of Larsen’s (1927) study of Town- sendia was her demonstration that Hooker’s “species,” T. sericea, was based on two specifically distinct elements. She indicated that plants collected in the “Rocky Mountains” by Drummond accord with Hooker’s description of T. sericea in his “Flora Boreali-Americana,” and that those collected by Richardson (which are the type of Aster ? exscapus = T. exscapa [Richards.] Porter) present some discrepancies with Hooker's description, particularly in the character of the leaf, pubescence, and pappus: She consequently interpreted the Drummond plants as the type of T. sericea Hook. and the type of the genus. “en Two specifically distinct elements were represented in the plants collected by Richardson and Drummond. However, T. ex- scapa must be taken as the type of the genus because its type Was included in the original species of Townsendia. doshas te Hooker's species, T. sericea, cannot be maintained to include the element of the original material which is not T. exscapa because it included the type of the earlier Aster P exscapus. This other element is referred to a newly described species, T. Hookeri. THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 7 Tue Locauities or Nutrait’s TOWNSENDIA COLLECTIONS The largest number of species recognized in this study which has been described by a single botanist is four. Both Asa Gray and Thomas Nuttall described that number. The specimens upon which Gray's species are based were obtained by several col- lectors and their data are fairly precise. The specimens upon which Nuttall’s species are based were collected by himself dur- ing his journey with the Wyeth expedition from St. Louis, Mis- souri, to Fort Vancouver, Oregon, in 1834. In general the local- ities Nuttall listed for his specimens are almost impossibly vague. The localities for his specimens of Townsendia are no exception. No journal made by Nuttall during this trip is known (Graustein, 1951), so most of the information about his localities must come from his published account (1840, 1841) and the label data on his specimens. In addition to the papers of Jepson (1934) and Pennell (1936) which consider Nuttall’s journey across the West, the problem has recently been approached by Goodman (1943), Barneby (1947), and Rollins (1950) in regard to Parthenium alpinum. One of the species of Townsendia which Nuttall described (T. spathulata) has range limitations somewhat similar to Parthenium alpinum. The other species of Townsendia which Nuttall de- scribed also have relatively small and well-defined ranges. There- fore, it has been considered worthwhile to bring together the available data on Nuttall’s Townsendia collections and make a critical analysis of them. For recording the label information on the specimens in the British Museum, I am indebted to Dr. Rollins. The four species Nuttall described and the one additional species he collected are listed below with the data which Nuttall published and the data which accompany his specimens. Townsendia grandiflora. . ; Published locality: “Hab. With the preceding, . . .” [the preceding is T. strigosa]. ] Label localities: Hills of the Platte and into the R. Mts. (BM). Townsendia Hookeri Beaman (the determination by Nuttall was T. sericea Hook.). : : Published locality: “Hab. On the Black Hills, (an alpine chain 8 JOHN H. BEAMAN toward the sources of the Platte). Flowering probably in April.” None of Nuttall’s collection of this species is in American herbaria and through an oversight of mine the data of the specimen in the British Museum were not recorded by Dr. Rollins. Townsendia incana. Published locality: “Hab. With the above [T. Hookeri]. Flowering in June.” Label localities: Black Hills of the Platte, June (pm). Black Hills of the Platte, grassy plains (pm). Black Hills of the Platte (PH). Platte plains (Gu, ex Herb. John A. Lowell). Rocky Mts. lat. 42° (cu, in Gray’s handwriting). Townsendia spgthulata. Published locality: “Hab. With the above.” [T. Hookeri and T. incana]. Label localities: Black Hills of the Platte, June (sm). Black Hills of the Platte in “[?, nearly illegible] campis etato rocky” (BM). Black Hills of Platte (pH). Black Hills [of, illegible] Platte (cu, ex Elias Durand). Townsendia strigosa. Published locality: “Hab. On the Black Hills, (or eastern chain of the Rocky Mountains,) near the banks of the Platte.—Flowering June.” Label localities: Black Hills of the Platte (pm). Black Hills of the Platte, June (pm). Black Hills (cu, in Gray’s handwriting). R. Mts. Platte (pH). The most confused and inconsistent of the sets of data above is associated with T. grandiflora. It seems unlikely that Nuttall could have collected T. strigosa and T. grandiflora together as he stated. The well-defined ranges of the two species are not known to be less than 100 miles apart at their closest points. T ownsendia grandiflora is probably the eastern-most species of the genus collected by Nuttall. It appears likely that the locality “Plains of Platte” as is on the Philadelphia Academy specimen is more accurate than the published locality or that on the specimens of the British Museum. It is possible that Nuttall obtained the species more than once, which may account for some of the di- versity of his label information. Evidence is presented in the systematic treatment under the discussion of T. spathulata which suggests that Nuttall obtained this species from more than one locality. Nuttall’s inference that he obtained T. Hookeri, T. incana, and T. spathulata together seems likely. I have obtained these THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 9 three species from a single locality near Alcova, Natrona County, Wyoming (T. Hookeri, Beaman 877, T. incana, Beaman 875, and T. spathulata, Beaman 878 ). Townsendia incana and T. spathulata have been obtained at this locality by other collectors also. It is in- teresting that Parthenium alpinum has recently been obtained by Ripley and Barneby and by Porter at this same locality after be- ing known only from the type collection for more than 100 years. Ironically, Nuttall’s locality data for Parthenium and those for the Townsendia species are entirely different. It seems probable that Nuttall’s term “Black Hills of the Platte” embraces a rather large area in central and western Wy- oming. His “Black Hills” are possibly the Wind River Mountains. These mountains are an alpine chain and are at one source of the Platte (actually Sweetwater) River. They would have been the ighest mountains which Nuttall saw in Wyoming, as he followed the Oregon Trail. Aside from his Townsendia collections, the “Black Hills” locality was not frequently used by Nuttall. He used it for Gnaphalium dimorphum ( = Antennaria dimorpha), for Antennaria plantaginea ( = ?, I have not seen specimens of this Nuttall collection), and for A. parvifolia. Neither Antennaria dimorpha nor A. parvifolia has a very limited range as do the Townsendia species, and therefore are of little value for locating the “Black Hills.” One feature about Nuttall’s locality data throughout his paper (1840, 1841) is that he frequently used the same or similar locality information for each species or most of the species of a genus. This makes one wonder if the similarity always resulted from a coincidence of collecting several species of a genus in one locality, or if some other factor accounts for the similarity. In Townsendia, with the possible exception of T. grandiflora, Nuttall’s locality data are vague enough to escape being classed as incorrect. GEOGRAPHICAL DISTRIBUTION, CLIMATIC AND EpapHic REQUIREMENTS Townsendia is primarily a genus of the Rocky Mountains of the United States. The ranges of apomictic populations of three species extend into Canada. One of these species, T. exscapa, is found also in Mexico. Only one other species, a Mexican en- demic, is now known from that country. The easternmost range of a member of the genus is that of T. exscapa, which occurs as far east on the Great Plains as northeastern Nebraska. The west- ern limit of the genus is on the Columbia River plateau in central 10 JOHN H. BEAMAN Oregon where T. florifer occurs. Townsendia is mostly found above 3500 to 4000 ft., but along the Columbia River in Oregon and Washington T. florifer is at less than 1000 ft. The upper elevational limit of the genus is marked by apomictic popula- tions of T. Rothrockii and T. leptotes which occur at least as high as 13,000 ft. in Colorado. Ten species have been collected in both Colorado and Wyo- ming; nine occur in Utah and seven are found in New Mexico. Idaho and Nevada each have six; Montana has five, and Arizona has four species. Several of the western states have two or three species each. It is unusual for a species of Townsendia to be found in more than four states, and ten of the 21 species occupy only small areas in one or two states. Narrow endemism may be regarded as a common feature of the genus. It may be inferred from the distributions outlined above that both latitudinal and altitudinal climatic influences are important in governing the distribution of the species of Townsendia. Townsendia exscapa, T. Hookeri, and T. leptotes are the only species which do not occur within a relatively narrow climatic province. Even in the latter two of these species, the sexual forms are restricted to small areas with nearly uniform climatic conditions. Thus it seems probable that climate has had an im- portant role in the evolution of the members of the genus. n many instances edaphic factors appear to be even more important than climatic factors in regulating the distribution of the species of Townsendia. Especially within the range of a species, the occurrence of plants on very localized edaphic situ- ations is frequently striking, As a generalization, with not too many exceptions, it may be stated that the acaulescent, rosulate forms require more highly specialized edaphic situations than do the caulescent types. The former mostly require open, finely divided limestone, sandstone, or shale rubble with a sparse vege- tation. Edaphic sites of this type are illustrated by the habitat photographs in Plate XV, figs. 1 and 2 and in Plate XVI, fig. 2. These edaphic conditions might be characterized as stable in a short-time sense and unstable in a longer- ( geological ) time sense. Even though the caulescent forms are less strictly limited edaphically, their requirements are somewhat parallel to those of the acaulescent types. , Soil samples have been collected at habitats of all the camer but time limitations have not permitted an analysis of these. me H analysis is planned as a future study. Tentatively it may inferred from the general character of the soils of the western THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA ll United States that the species in the more arid habitats occur in soils on the basic side of the pH range, while those in the moister habitats are probably in neutral or only slightly acid soils. GENERIC RELATIONSHIPS In a genus with species as closely related to one another as are those of Townsendia, it would seem.that relationships of the genus to other genera might also be evident. However, few in- vestigators have committed themselves on the generic relation- ships of Townsendia. The limits of the genus never have been questioned. There has been but a single species placed in Town- sendia which belongs in a different genus, and this was first ex- cluded by Gray, its author. Likewise, there have been no species transferred from other genera into Townsendia (with the excep- tions of T. exscapa and T. florifer which were described original- ly with doubt as to their generic position ). Consequently, the generic relatives of Townsendia are not revealed by its nomen- clatorial history. The genera conventionally assigned close to Townsendia have not had a similarly tranquil history. The many changes in Aster are well known. Astranthium has been with Bellis; Dichaetophora has been with Boltonia; and Keerlia is now in Chaeto These examples by no means exhaust the list of changes, but the reader is referred to Shinners (1946a) for further details. Although the uncomplicated nomenclatorial history of Town- sendia suggests for it a distinct phyletic position, its features were not considered unique by Bentham (1873). In his comprehensive “Notes on the Compositae” he wrote (p. 408): “Thus we have the Asiatic and North-American Boltonia, the Asiatic Heteropappus, and the N.-American Townsendia, Monoptilon, and Psilactis dit- fering from the typical Asters of the same regions in scarcely any thing but the pappus.” Gray (1888) in his “Synoptical Flora in the section Megalastrum of Aster tersely wrote: related to subsection Xylorrhiza and to Townsendia.” And Larsen (1927) noted that “the heads [of Townsendia] resemble those of the closely related genus Aster.” Otherwise, ideas of the generic re- lationship of Townsendia have been indicated only by its system- atic position in works by such authors as Gray (1888) and Hoff- mann (1897). The rationale of the statement by Gray (quoted above) is evident. Section Megalastrum includes Aster Wrightii which Gray had once put in Townsendia. From the present study, it appears that Aster Wrightii could be considered to have 12 JOHN H. BEAMAN only a rather distant relationship with Townsendia. Species of several other genera have a considerably closer relationship. In Shinners’ (1946b) study of some of the Astereae he noted that in the American genera of the heterochromous Astereae an arrangement on the basis of the form of the receptacle was better than that based on the pappus. Those conical-receptacled genera which he treated are Egletes, Erigeron (in part), Aphanostephus, Astranthium, Dichaetophora, and Boltonia. In his key to separate the above genera, Astranthium, Dichaetophora, and Boltonia fall into one group. An examination I have made of these three genera indicates that they have a number of morphological simi- larities. Also, they have characteristics in common with Town- sendia, which was not included in Shinners’ treatment because it was not at that time known to have a conical receptacle. In the present study, Townsendia formosa has been found to have a conical receptacle. Through this species it is possible to recognize the relationship of Townsendia with Dichaetophora, Astranthium, and Boltonia. Another feature which indicates their relationship is the stem pubescence, which is consistently of simple, strigose trichomes. The leaves, especially of Dichaetophora and some of the Astranthium species, are very similar in shape, texture, and pubescence to those of T. formosa. The pappus of T. formosa is very similar to that of Astranthium, while the pappus of Boltonia and Dichaetophora is somewhat similar to that of T. eximia. The achenes, which are compressed, ovate in shape, and with two conspicuous margins (extended into wings 10 Dichaetophora), are very similar in all four genera. Dichaetoph- ora also has duplex, glochidiate achenial hairs (“peculiar gland- tipped hairs” according to Shinners) on the central portion an margins of the achenes. These hairs are indistinguishable from those of several species of Townsendia. The achenes of Dichae- tophora also are papillose in a manner very similar to those of some of the members of Townsendia. The four genera also have a close phytogeographic relation- ship. The center of diversity of Townsendia is in the southern cordillera of the United States. The centers of Dichaetophora and Astranthium are not far to the south, and that of Boltonia 1s to the east. Townsendia and Dichaetophora are probably the two closest members of this four-genus alliance, but Astranthium is not too distantly related. Boltonia seems somewhat more widely set art. Within Townsendia, the characters I regard as primitive are: THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 13 a perennial, fibrous-rooted, rhizomatous and stoloniferous habit; erect, monocephalous stems; large, thin, glabrate, spatulate leaves; lightly pubescent stems; large heads; a conical receptacle; several rows of broad phyllaries with narrowly scarious and minutely ciliate margins; large, obovate achenes; and a small, unelaborated pappus. With the exception of the phyllaries, about which I am not sure, these characters may be considered to be primitive (when present) also in the three other genera under discussion. Townsendia formosa appears to be a more unspecial- ized form than the other Townsendia species, and also the mem- bers of the other three genera. The concept of a small, squamellate, unelaborated pappus as a primitive character requires explanation. Under most circum- stances it would be looked upon as a reduced feature. But in this case, T. formosa and three closely related genera have the small pappus. Within Townsendia, the species most closely related to T. formosa (T. eximia) has a larger but similar pappus. The de- rived relatives of T. eximia in turn have a pappus which is even more highly elaborated. It seems, therefore, that the evolutionary trend has been in the elaboration rather than the reduction of the pappus. The selective advantage of a large pappus as an agent of dispersal seems evident. The alternative explanation, that the pappus of Townsendia formosa is a reduced feature, would require the postulate that the genera related to Townsendia have been more recently segre- gated from T. formosa stock than have the rest of the Townsendia species. This explanation seems improbable. Although many similarities between T. formosa and the three other genera have been indicated above, the latter still have greater differences from T. formosa than do the other species of Townsendia. For example, Dichaetophora, which seems to be the genus most closely related to Townsendia, is comprised of a single annual species. This life- form is surely a long phylogenetic distance from the rhizomatous, long-lived T. formosa. : It is beyond the scope of the present work to consider the position of Townsendia and its close generic relatives in regard to their position with the rest of the Astereae. However, it should be noted that the embryological similarities between Townsendia and Erigeron (considered in the section on reproduction ) do not necessarily suggest a close relationship for the two genera. These peculiar embryological features may eventually be found in still other members of the Astereae. 14 JOHN H. BEAMAN SPECIES CONCEPT It is hoped that the present systematic treatment approaches an ideal which has been expressed by Cain (1944). He stated (p- 7): “Taxonomy attains a logical basis when the data of com- parative morphology can be arranged in a geographical pattern that coincides with the probable phylogeny of the group and the history of the floras . . . in which it has been involved.” In this study an attempt has been made to designate the taxa by a system which applies insofar as possible a traditional nomenclature, yet emphasizes the morphologically recognizable units on a geo- graphic basis. There is a marked tendency in Townsendia for species with the same flowering periods to have allopatric ranges. A criterion of strict genetic isolation cannot be used in Townsendia. Such an approach would result in the “lumping” of most or all of the species into one. Geographic barriers are most important In isolating the species of Townsendia. If genetic barriers do exist in the genus, they are probably not strongly expressed in the first hybrid generation in most instances. An understanding of the mechanism of reproduction in Town- sendia has been a necessary basis for explaining peculiarities of geographic distribution of some of the species. Cytogenetic and embryological data have therefore been of value in facilitating an understanding of the comparative morphology of the group on a geographic basis. But these data have been of little value in the determination of specific boundaries. ; Most of the species are nearly parallel in their degree of dis- tinctness. However, T. formosa possesses a considerably larger set of unique characteristics than does any other species. Those taxa at the varietal level were placed there as geographic segre- gants with fewer characters distinguishing them from their near- est relatives than were usually available to distinguish species. No complete intergrading series is known between the varieties, in the two species where they were designated. The term va- riety” rather than the term “subspecies” is used for the infra- specific units primarily because its use in this sense has long been accepted in botanical classification. According to the Inter- national Code of Botanical Nomenclature (Lanjouw et al., 1956, Article 4) either of the terms might have been used. In the present taxonomic interpretation of Townsendia, it would not have been possible to use both of these infraspecific categories. THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 15 REPRODUCTION Apomixis in Townsendia previously was suggested by five indirect lines of evidence reported in a preliminary cytogenetic investigation in the genus (Beaman, 1954). In the present study the problem of apomixis was attacked directly by comparative embryological examinations of mega- and microsporogenesis and subsequent development in sexual and apomictic populations. Knowledge of the mechanism of apomixis in Townsendia has been considered indispensable to a formulation of taxonomic con- cepts in the genus. Different types of apomixis may have different effects on evolutionary trends. For example, pseudogamous apomicts are more dependent on environmental factors, such as pollinating agents, than are autogamous apomicts. Facultative apomicts may lack some of the efficiency in reproduction which characterizes obligate apomicts. But obligate apomicts may be less plastic under the action of selective forces than are facultative apomicts. Therefore, this study was designed to expose the morphological and cytological features of apomixis in Townsendia and to evaluate the significance of apomixis to evolution in the genus. MATERIALS AND METHODS The plant material which has been used in this study is listed in Table 1. Voucher specimens are filed in either the Gray Herbarium of Harvard University or in the Herbarium of the State College of Washington. The plants were grown in the ex- perimental gardens at Harvard University in Cambridge, Massa- chusetts or at the State College of Washington in Pullman, Wash- ington. In some cases buds used in the study were taken from plants growing in their native habitat. Studies of microsporogenesis and male gametophyte develop- ment were made from florets fixed and smeared according to a method previously outlined (Beaman, 1954). A few observations of microsporogenesis were made in embedded and sectioned anthers prepared along with the florets used in the study of mega- sporogenesis. However, this material was unsatisfactory for critical examinations. Studies of pollen were made from unopened (i.¢., pre- anthesis) florets from herbarium specimens. The corollas of these florets were macerated in a drop of aceto-carmine on a slide and observed immediately under the microscope. Pollen size and 16 JOHN H. BEAMAN degree of abortion are readily apparent by the use of this rapid technique, which was employed in the taxonomic study to deter- mine whether the herbarium specimens represented sexual or apomictic populations. The pollen nuclei become darkly stained after being exposed to aceto-carmine for from one to ten minutes. Pollen may be satisfactorily photographed for about 45 minutes after being placed in aceto-carmine. If the staining period is longer, the cytoplasm becomes darkly stained, and a satisfactory contrast between the nuclei and the cytoplasm cannot be ob- tained. Studies of megasporogenesis, female gametophyte develop- ment, and embryogeny were made from buds fixed either in Belling’s modified Navashin’s fluid or in a Carnoy’s fluid (6 parts absolute alcohol: 3 parts chloroform: 1 part glacial acetic acid). When Carnoy’s fluid was used, the buds were transferred to 70 per cent alcohol after being in the fixative for about one hour. With Navashin’s fluid, aspiration was sometimes employed. The best fixation was obtained by removal of the top portion of the achene before fixation. Aspiration was unnecessary when ovaries were opened in this manner. The period of fixation lasted from a few days to over two years. As might be expected, the material used shortly after fixation yielded the best results. Also, material fixed in Navashin’s fluid gave much better results with the paraffin-embedded technique than did that fixed in Carnoys fluid. Material fixed in the latter was used only to supplement the material fixed in Navashin’s fluid. Material fixed in Navashin’s fluid was washed in water and dehydrated in a tertiary butyl alcohol series. Material fixed in Carnoy’s fluid was transferred directly from 70 per cent alcohol to the tertiary butyl alcohol series. The material was embedded in Fisher Tissuemat and sectioned at from 10 to 15 microns. Even in old florets, 10 microns proved to be the most satisfying thickness. A ferric ammonium sulphate-haematoxylin staining schedule, described by Esau (1944), gave good results. The slide preparations are in the personal collection of the gar e photomicrographs were made with Bausch and Lom photomicrographic equipment and a Zeiss microscope with apo- chromatic 20X, 40X, and 90X objectives. Ansco Isopan film was used. THE MALE REPRODUCTIVE PHASE In a previous paper (Beaman, 1954) studies of microsporo- genesis in ten species (13 according to the species delimitations THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA pigs then used) of Townsendia were reported. At that time the basic chromosome number x = 9 was postulated for the genus. In diploid, sexual plants almost no meiotic irregularities were found, but a high degree of meiotic irregularity was encountered in polyploid, apomictic plants. The specific names of some of the material used in the previous study (1.c.) should be emended as follows: T. anomala (culture 33)=T. condensata; T. arizonica (cultures 4 and 5) = T. incana; T. Hookeri (culture 38) was TABLE I. Material of Townsendia used in the es and cytological investigations.1,? Taxon boa fees Culture 2n Place grown Date of Chromosome fixation number T. condensata Beaman 893 — ? Native aa Aug. 1954 T. condensata Witt 1845 47 ca. 36 Pullm Spring 1955 T. eximia Beaman 699 699 18 Native Sake June 1954 an Cambridge, Mass. Nov. 1955 T. eximia Beaman 704 — 18 Native habitat une 1954 T. eximia Beaman 719 seis 18 Native habitat une 1954 T. eximia Beaman 740 — 18 & 20 Native habitat une 1954 T. eximia Beaman 745 oo 18 Native itat uly 1954 T. formosa Beaman 994 994 18 Cambridge, Mass. T. glabella Beaman 761 761 18 Cambridge, Mass. June 1956 T. grandiflora Beaman & 39 18 Pullman, Wn. spring 1955 Preece 509 -- T. grandiflora) Beaman 744 — 18 Native habitat July 1954 T. incana Preece & 5 18 Pullman, Wn. Spring 1953 Turner 261 - incana Beaman 802 802 18 Native aye July 1954 T. incana Beaman 971 971 18 Cambridge, Mass. Nov. 1955 T. incana Preece & 6 ca. 27 Canmnge: 7 Nov. 1955 Turner 2975 (variable) and Pian Wn. Spring 1955 T. incana Preece & 17 ca. 30 Pullman, Wn. Springs of : Turner 2510 — 1953 & 1955 T. incana Beaman’801 bail: 29 Native habitat July 1954 T. incana Beaman 875 875 36 Native habitat Aug. 19 T. mexicana Beaman 1002 sh 18 Native habitat Sept. 1954 T. montana Beaman & 35 18 Pullman, Wn. cee 1955 var. montana Preece 505 T. Parryi Ownbey s.n. 8 36 Pullman, Wn. Spring 1953 in 1948 T. Parryi Roeder 280 27 36 Pullman Spring 1955 T. Rothrockii Beaman & 798 ? Native on July 1954 Barclay 798 T. texensis Seana Fo — 18 Native habitat June 1954 Original localities for this material may be found in the Systematic Treatment in the citations of specimens under the various species. Additional chromosome counts in Townsendia will be found in a previous paper (pedal 1954). 18 JOHN H. BEAMAN misidentified as T. mensana; T. minima (culture 3) = T. montana var. minima; T. condensata (culture 47) was misidentified as T. spathulata. The chromosome numbers of five species not in- cluded in the previous study are reported in Table I in the present paper. Microsporogenesis in diploid, sexual plants In all but one collection of the diploid material examined in this study, microsporogenesis was regular and nine bivalents were observed at metaphase I. In the exceptional collection (Beaman 945) at least one plant was characterized by 10 bivalents. Ap- parently the pairing attraction of the tenth chromosome pair was not very strong, as some cells were found with nine bivalents and two univalents instead of ten bivalents. Other material collected from this same population is characterized by nine bivalents at metaphase I and no meiotic irregularities. It seems probable that the ten-chromosome material is a rare abnormality. An examination of the meiotic karyotype of T. mexicana re- vealed meiotic chromosomes of a smaller size than has been found in any other species of Townsendia. A metaphase plate from a preparation of T. mexicana is shown in Plate I, fig. 3. This figure may be compared with Plate I, fig. 2, which illustrates larger metaphase chromosomes typical in the genus. The karo- type of T. annua, the species morphologically most similar to T. mexicana, has not been examined. Microsporogenesis in polyploid, apomictic plants In the present study no special attempts have been made to obtain additional chromosome counts in apomictic material. In- stead, investigations have been concentrated on material from apomictic populations of two species. These were chosen pat- ticularly because one is triploid and the other tetraploid. Both bivalents and univalents are found at metaphase I in the triploid. This stage in the tetraploid is characterized by 36 univalent chromosomes. The triploid material is from a population (cul- ture 6) of T. incana. This species, especially this population, has been used extensively also in the investigation of megasporo- genesis. Thus, comparative data of the male and female repro- ductive phases in the same population have been obtained. The tetraploid material is from a population (culture 8) of T. Parry! Megasporogenesis in this population also has received attention. Townsendia incana (culture 6). Meiosis in the anthers of plants of this population is characterized by a considerable THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 19 amount of chromosome pairings, as may be seen from Plate I, fig. 4. The number of bivalents in relation to the number of uni- valents is not constant. Mostly there are between five to ten bivalents and six to 15 univalents. No pollen mother cells with all univalents have been observed. Meiosis in the ovules, on the other hand, frequently is characterized by all univalent chromo- somes at late prophase and metaphase (see Table III). Some variation of the chromosome number occurs within the population. In addition to the chromosome count of 2n = 28 which was previ- ously reported (Beaman, 1954), the count 2n = 27 also has been obtained. From some of the examinations of microsporogenesis it appears that the number 2n = 26 might occur in the population, but chromosome counts from microsporogenesis are difficult t obtain and often are unreliable. A variable chromosome number could be expected in the population, however, as a result of meiotic irregularities in megasporogenesis. Micronuclei, which have been observed with dyads of megaspores, probably repre- sent chromosomes or chromosome fragments excluded during megasporogenesis. In this triploid population the loss or gain of a small amount of chromosomal material might not be lethal. At anaphase I in culture 6, lagging univalents are frequently seen (Plate II, fig. 1). The halves of the bivalents go directly to the poles during anaphase I, but univalents seem to be much less strongly attracted to the poles. Usually the univalents eventu- ally reach one of the poles, but sometimes they are excluded from the main nuclei and form micronuclei. Chromosome counts at metaphase II indicate that in the first division the triploid num- ber of chromosomes is reduced by about half in the two resulting nuclei. Usually 12 to 15 chromosomes are seen at metaphase II. The distribution of the univalents at the first anaphase is unequal, and the two poles do not receive exactly the same number of chromosomes. Tetrads of microspores in culture 6 (Plate II, fig. 2) have a fairly normal appearance. Their subsequent development, how- ever, is not normal. Cell walls usually do not form between the microspore nuclei, and all four nuclei are surrounded by a wall which develops the structural characteristics of an ordinary pollen wall (Plate II, fig. 3). The ultimate “male gametophyte” which results from this type of development is a giant, several-nucleate pollen grain (Plate IV, fig. 2). At anthesis, in addition to the large grains, there are many aborted grains of various sizes. The determination, from herbarium specimens, of plants from apo- mictic populations is an easy matter when the pollen has been 20 JOHN H. BEAMAN formed in the manner outlined above. A reference to this peculiar method of pollen development was made by Holmgren (1919) in his study of Erigeron. He noted, in sexual material, that pollen of florets in a transitional position on the head between pistillate ray-florets and hermaphroditic disk-florets developed by wall formation around the four nuclei of the tetrad. He stated that this abnormality was not infrequently encountered in apomicts and hybrids. A few other references to this type of pollen de- velopment occur in the embryological literature. Microsporogenesis and pollen development in culture 6 are of interest in regard to the recognition of apomixis in populations from herbarium specimens. However, since reduction occurs in this material, little information is gained on the mechanism of meiosis on the female side where reduction must be avoided. Tetraploid culture 8 of T. Parryi differs in its microsporogenesis from culture 6 of T. incana and possibly may have similar proc- esses in both the male and female phases. Townsendia Parryi (culture 8). Gustafsson (1934, 1935) has further considered. ; No anaphase restitution nuclei have been positively identified in Townsendia in mega- or microsporogenesis. It might seem, therefore, that the pseudohomeotypic division occurs in the genus. The univalent chromosomes observed in megasporogenesis 11 the apomicts might also be suggestive that the pseudohomeotypt¢ division occurs in Townsendia. Especially, the occurrence, at diakinesis or prometaphase in megasporogenesis, of univalents with visible chromatids (Plate I, fig. 5) might indicate this type of division. as Fagerlind recently has examined megasporogenesis In so mictic material of Rudbeckia (1946), Erigeron (1947a), ae Taraxacum (1947c), and microsporogenesis in apomictic Hierac- THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 21 Prare I. e biv mosomes genus genesis in or prometaphé 2g in apom rictic, triploid tome blade bisected this cell, ad the yi ig iyi Fig. CC hesencuiiees at metaphase II in 1 oid T. Nine rt ery et ee I in microsporogen wie Fig. a _are smaller in T. mexicana t - Fig. 4. Bivalent an nd univalent chromosomes at metaphase I in mic ian 5 6 es in ps msendia (all X 1200). xual, diploid T. incana. is T. eximia. Selrccrie id — of meiotic chromosomes valen diakinesis in microsporog x 7 is in T. mexicana. The o- hose ponte in any other species of Fike -rosporo- lents at metaphase I in mi esi triploid T. incana. Fig. ivalent chromosomes at diakinesis T. incana. The micro- portion shown contains about half of the nicrosporogenesis in sexual, =) inca 2a. 22 JOHN H. BEAMAN Piate II. Some aspects = toro ace in riper populations of Tow sendia. Fig. 1. Anaphase I agging ivalent chromosomes in triploi incana (X 1200). The vein of the nage bey x0 dissetty to * poles at gre : one the les but while the univalents lag. Usually the univalents reac y arate m times they are incorporated into separat saree ig °. Tetrads in ‘riploid T. incana (X 200). These — the appearance of normal tetrads, but the microspores do not separate, and all four nuclei are a obese a single po le gral A; Fig. 3. Young pollen grains of triploid T. coe a forn y the inclusion of tetrad of microspores within one wa all (xX 47( Fig. 4, ole mother cells with al univalent chromosomes in tetrap _— Si hh & ras se . 5. A more highly mag- nified view than fig. 4 of 36 u chro a ts taphase I in T. et ae 1200). Fig. 6. Anaphase I in ceo “AN 7, : Berens: (x 1200). This is an extre naphase. Usually hele: univalent chromosomes do not have a strong tendency ‘to eae e to the two poles. THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 23 Prare III. Some phases of i hPa TY in — ctic b popiesions of Town- sendia (continued from Plate Il). Fig. 1. A metaphase restitution or contraction nucleus composed of u ala serra in pio T. Parryi (X 1175). Fig. A restitution or contraction nucleus in tetraploid T. nial ( a slightly later ome number h | when all er aa »ymosomes clump at metaphase I instead of anerenns » the oe poles and remaining there at anaphase I. Fig. 4. Side view of n See : tetraploid r. Weve: (< 1175). Fig. 5. Side view of anaphase II in ieee Sa Pairs (X 1175). Reduction apt not occur at anaphase I and thus a dyad ot med at ana- phase II. Fig. 6. Dyads apom mictic polyploi id a age Oe )). The two m5 ee develop Lag into pollen grains. Apomicts which have all or nearly wate valent chromosomes at metap os I form dyads athe than tetrads of micro- 24 JOHN H. BEAMAN Photomicrogr sol . pollen in sexual and apomictic poptane” of Townernie (all & 330). . Pollen of sexual, diploid T. incana. Fig. 2. Pollen - Laine: ao vid T. inca A: ioe giant grains result feces the aes of e four spores of a tetrad into a single -_ Non-abortive grains contain more fan three nuclei but these are so contorted t is not possible to determine xactly Boa: ny are present. Fig. 3. Pollen Ry apomicti, tetraploid T. yi ese very eae grains have sath is om dyads s eyes to the ones sho in Plate IIT, fig. 6. The nuclei in these pollen grains bis the unreduced, tetraploid chromosome number. Fig i a = ies a randomly selected herbarium 6 ) ally of T. i a (Walker 206). T centage of paras grain in apc as Eis as or higher than that gt sero here by the grains ahaut nuclei. THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 25 ium (1947b). His observations in these genera have led him to doubt that the pseudohomeotypic division occurs frequently or that it even occurs at all. He emphasizes that restitution nuclei are not necessarily formed only at anaphase but may occur also at metaphase, prometaphase, diakinesis, or even earlier in pro- phase. Fagerlind especially stresses the fact that the “contraction nuclei” (a term coined by Rosenberg for the clumped meiotic chromosomes of Hieracium) he observed in Erigeron, Hieracium, Taraxacum, and Rudbeckia might be restitution nuclei which are formed at metaphase or earlier. A clumping of meiotic chromosomes in several apomictic collections of Townsendia was reported in the previous cyto- genetic study in this genus (Beaman, 1954). Clumped chromo- somes in meiosis appear to be a regular feature in many apomictic forms. In Townsendia, the clumping in microsporogenesis is so prevalent in material from some populations that it defies analy- sis. In culture 8 of T. Parryi, however, it has been possible to analyze the various processes of the apomictic meiosis in micro- sporogenesis, and to determine the probable sequence of the meiotic stages. Smears have been used in this study in order to render visible as much as possible of the chromosomal detail. Fagerlind’s studies apparently have all been made with sectioned material. Also, other investigators of the mechanism of the meiotic non-reduction phenomena have used mostly sectioned material. Sectioned microsporocytes of Townsendia are unsatis- factory for critical chromosomal analyses. Bivalents have never been found in microsporogenesis in culture 8 of T. Parryi. When chromosomes become clearly evi- dent in this material at diakinesis or prometaphase they are uni- valents. Whether the chromosomes were paired at an earlier stage and have come unpaired, or whether they were never paired, was not determined in this study. More refined cytologi- cal techniques will be required for the solution of that problem. Typical univalent chromosomes in this material are shown at two different magnifications in Plate Il, figs. 4 and 5. Infrequently, a slight tendency is manifested for a migration of the univalents to the two poles (Plate II, fig. 6) at anaphase I. More frequently, a single clump of chromosomes is found at the center of the cell (Plate III, fig. 1). In some clumps, in cells which are apparently at slightly later stages (Plate III, fig. 2), the chromatids become evident. Seen in polar view at a still later stage, the clump is loosened and the chromatids are very conspicuous (Plate III, fig. 3). A count of the chromosomes at this stage indicates that 26 JOHN H. BEAMAN the number has not been reduced. After the anaphase that fol- lows, the chromosomes in the resulting dyad have a telophase II or slightly mitotic aspect (Plate III, fig. 5). These are like telo- phase I chromosomes. A wall is formed after this division and an unreduced dyad results. Perhaps this dyad is homologous to those found after megasporogenesis! The “microspore dyads” which result after non-reductional meiosis soon separate into their component “microspores” and these may develop into normal- appearing (except for their large size) pollen grains (Plate IV, fig. 3). At anthesis many aborted grains of various smaller sizes are present also. It is therefore possible to detect these apomictic forms by herbarium specimens. It appears, according to the events outlined above, that the univalents at metaphase I may undergo a migration toward the poles. The migration is weak, however, and two nuclei and a corresponding reduction in the chromosome number do not occur. Rather, the chromosomes are oriented into a single clump (con- traction nucleus). While thus clumped, chromatid separation becomes evident. By the time the clump loosens, the chromatids are nearly separated. Anaphase II begins immediately and two unreduced nuclei result. Discussion of microsporogenesis in polyploid, apomictic plants. Fagerlind has stressed the point that at the time of formation of the contraction nucleus, a nuclear membrane _ re-encloses the chromosomal mass. No evidence of a nuclear membrane around the chromosome clumps has been detected in the smears of Townsendia microsporocytes. In sectioned prepara- tions, the nuclear material is rather diffuse at this stage, but such figures might be interpreted as having a nuclear membrane around the chromosome clump. No indications of twisted or varl- ously contorted chromosomal masses have been observed in smears of anthers of Townsendia. However, misshapen nuclei were seen at corresponding stages in sectioned material. It is possible that some of the details of Fagerlind’s description of the metaphase restitution nucleus (contraction nucleus) are based on artifacts in his sectioned material. Smears are most useful for chromosoma’ analysis, and sectioned material is mainly valuable as a chec for determining the proper sequences in meiotic development which are more difficult to determine in smears. The evidence from smears of material of Townsendia — to corroborate Fagerlind’s idea of the significance of the nienne meiotic chromosomes. It seems probable that the genetic controls THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA ya of the rhythm of meiosis are such that, even in apomicts, univalent meiotic chromosomes cannot behave as ordinary mitotic chromo- valents is actually equivalent to telophase I in normal, sexual material. In some of the apomicts meiosis may be impossible to analyze because the contraction nucleus or chromosome clump is formed earlier than metaphase. It appears, therefore, that the apomicts of Townsendia have a specialized type of meiosis (the semiheterotypic division ) which avoids reduction of the chromo- somes during microsporogenesis. From the present studies of megasporogenesis it seems probable that the type of meiosis out- lined above for the male reproductive phase of culture 8 of T. Parryi may occur also in the female phase of this and the other apomictic populations of the genus. It should be noted that Avers (1954) has produced evidence for the occurrence of a functional tertiary split of meiotic uni- valents in the microsporogenesis of triploid Aster hybrids. In her material univalent division appears to occur frequently at ana- phase I in addition to the normal anaphase II division. No similar mechanism has been observed in Townsendia. The peculiar alterations of microsporogenesis in the polyploid apomicts of Townsendia suggest a lack of functionality of the pollen which results. Also, it is unusual in these plants for the percentage of nucleated pollen to run higher than 20 per cent. Frequently this percentage is lower than five per cent, and some- times no pollen is produced. Thus, the evidence from the male reproductive phase in the apomicts argues strongly for obligate apomixis in this genus. Evidence from the female reproductive phases reinforces this conclusion. THE FEMALE REPRODUCTIVE PHASE Investigations of the female reproductive phase in Town- sendia have not been made previously. However, studies in other members of the Compositae, especially in Erigeron, are of signif- icance to this study. Harling (1950, 1951a, and 1951b ) recently has been making comparative embryological studies in certain tribes of the Compositae. His paper on embryology in the Astereae (1951b) includes investigations in 14 genera of this 28 JOHN H. BEAMAN tribe. Except for one genus, Harling found a single, constant type of embryo-sac development in the Astereae. This develop- ment is of the “normal,” monosporic type with the production of an eight-nucleate embryo sac. The exceptional genus, Erigeron, has embryo-sac development of the normal type in some of its members, but also bisporic and tetrasporic developmental-types are found in the genus. Variation occurs both within and between species. In the Astereae the only genus other than Townsendia now known to have apomictic forms is Erigeron. In the latter, asexual reproduction was known from embryo-sac studies as early as 1915 from a preliminary report by Tahara. Several investigators have made subsequent studies of apomixis in Erigeron. Fagerlind (1947a) has reviewed these investigations and given a new inter- pretation regarding the mechanism of non-reduction in the mei- osis of apomicts of Erigeron. His ideas have been considered in detail in this paper under the section on microsporogenesis in apomictic plants. The features of both sexual and apomictic reproduction in Townsendia are very similar to those in Erigeron. The processes in apomictic reproduction in these two genera are not known to be exactly duplicated in any other genus. Thus, the results of the investigations in Erigeron have significance in the interpretation of the embryological phenomena in Townsendia. The female reproductive phase in sexual plants Material of Townsendia incana has been more extensively used that that of any other species in the investigation of sexua and apomictic reproduction in the genus. Several populations of both sexual and apomictic types have been available for study. Comparisons of the two types of reproduction are therefore pos- sible within a single species. Sufficient material of other species also has been available to indicate that most of the reproductive phenomena encountered in T. incana occur in other members of the genus. Townsendia incana (culture 971). This species, as is charac- teristic for the genus, has a single archesporial cell in each ovule. The early prophase stages of meiosis are morphologically similar in all the material thus far examined, including the apomicts. It does not seem possible with the present techniques to determine at early prophase whether the chromonematal strands are un- aired, pairing, or paired. Therefore, the terms leptotene, zyg0- tene, and pachytene are not used. A frequently encountered fea- THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 29 ture at early prophase is the so-called synezesis which consists of clumped chromonematal material and an associated, distinct nu- phase stages were encountered in culture 971. In the one dyad observed in culture 971 no cell plate was present. At the tetrad stage and at subsequent stages certain peculiarities become evident. Twenty-one ovules were observed in which no nuclei of the tetrad were separated by cell walls. These, with the exception of one tetrad which still possessed a phragmoplast between two of the nuclei and was thus too young for determination, may be considered as 4-nucleate coenomega- spores. The megaspore nuclei are arranged in a linear fashion, and, usually, the nucleus nearest the micropyle is separated by a considerable distance from the other three nuclei. Ten tetrads were observed which had cell walls between the micropylar and submicropylar megaspore nuclei (the micropylar megaspore nucleus is the one nearest the micropylar end of the ovule, etc. ). Eight tetrads were found with a wall between the submicropylar and subchalazal megaspore nuclei. In one ovule the cell wa had formed on the micropylar side of the micropylar megaspore nucleus. In Harling’s (1951b) study of Erigeron, he noted variations in wall formation in tetrads. Some of his material was character- ized by no walls; some had wall formation apparently after meta- phase I, some apparently after metaphase II, and some by wall formation after both meiotic divisions. He inferred the time of wall formation from the position of the walls relative to the nuclei of the tetrad. In Erigeron glabellus he found all of these conditions in a single plant. In Townsendia incana the occurrence of a wall outside of the tetrad, separating none of its nuclei, sug- gests that wall formation between the nuclei of tetrads has no perfect synchronization with the meiotic divisions. Furthermore, in much of the sexual Townsendia material, the abundance of walls between only the micropylar and submicropylar megaspore nuclei, without corresponding walls between the chalazal and subchalazal megaspore nuclei, suggests some lack of correlation of wall formation with meiotic division. In culture 971, tetrasporic embryo sac development is most 30 JOHN H. BEAMAN common, but also one of the megaspore nuclei may degenerate and the other three germinate to form a three-nucleate embryo sac. This fact is indicated by the observation of two three- nucleate and two six-nucleate embryo sacs. However, tetrasporic development must be more frequent than other types in this population (cf. Table II). The embryo sac becomes morphologi- cally organized after two series of mitotic divisions. Sufficient observations in culture 971 have not yet been made to permit a full understanding of the great amount of variability in nuclear number in embryo sacs above the eight-nucleate level. This type of variability has been observed also in several other composite genera, particularly in Erigeron (cf. Harling, 1951b). In regard to several species of Erigeron, Harling states that in the development of the female gametophyte, one or more of the chalazal nuclei “strike” (i.c., cease dividing while others con- tinue). Nuclear fusions in the embryo sacs of some plants, in- cluding Erigeron, have been noted. However, nothing which might indicate nuclear fusion has been observed in any of the material of Townsendia. Thus, it seems plausible that the vari- able number of nuclei in the embryo sac in culture 971 results from the “strike” of some of the chalazal nuclei at either or both of the postmeiotic divisions. A statement is necessary in regard to the rather large amount of aborted and undetermined ovules of culture 971 (recorded in Table II). Some of this material was fixed after two or three nights with frosts, and it appears probable that freezing had adverse affects. Material from the same plant fixed before the frosts was not affected. : Townsendia incana (cultures 5 and 802). No material at a stage later than the beginning of female gametophyte develop- ment has been available from these populations. Fewer meiotic stages, also, were examined in these ovules than in those of are 971, but no significant differences in meiosis in the three popula- tions were noted. A difference is apparent at the tetrad stage. however. In cultures 5 and 802 a wall and a space regularly separate the micropylar nucleus from the other three es of the tetrad (Plate VI, fig. 4). A sufficient number of later stage has not been seen to permit conclusions regarding further “i velopment of the embryo sac in these two populations, ce . emale gametophyte is initially three-nucleate from the develop ment of the three chalazal megaspore nuclei and the abortion of the micropylar nucleus. : ble A difference in genetic constitution probably is respons! THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 31 TABLE II. parative classification of ovule development in five seach ochepliclon populations of three species of Townsendin? Developmental stage Numbers of ovules observed in the e populations (cultures) 971 5 S6r°""'55- S04 39 Saclg cecil cell 2 a 1 1 — Early prophase 6 3 4 10 oe 2 ivaltaake. 2 — s i — 1 Diakinesis ? — _— 1 —_— —_ —_ Metaphase I 3 2 a —_ — apha — — 1 — _ — Kei e 1 2, Z 6 — 1 Metaphase II 1 1 — —_ = 1 Te 43 86 44 31 —_ 20 Tetrad — Z — — —_— = Z-nucleate embryo sat .......s.scscseseseeeore —_ —_ ae aR: 255. 3 3-nucleate 46 POA ecg cer 2 + es 6 — 48 3-nucleate ss adhe Cae easton — 1 1 1 3 4-nucleate iy susan ten Pee a ee 18 —_ _— 92 = os 4-nucleate td uO dah cover aenne neo 1 — — 2 SES = 5-nucleate x webiste OT SS OSE RE. i — —_ _ sees — 2 6-nucleate ‘3 Wek; Maa eaten eA 2 _ 2 — 17 6-nucleate —_” Sey en Re er — — — 1 ~~ a 7-nucleate 4 (Ge REE eh —_— — — 1 — ae 8-nucleate ” apes gut iba emia ante P —- _ 18 — 1 8-nucleate ” gle es Ree ee — — —_ 1 — = 9-nu € “4 LAR Ce ine ee 2 — — - eit = 10-nucleate ” lege CER RE ECie — — —_ 3 — — 11-nucleate ” piibgeas iste aed eet 1 a _ 2 — rae 12-nucleate ” Ot aan a 3 _ —_ 31 ae 5 12-nucleate ” air gave, oe Sane — —_— oe l cig rae 13-nucleate ” Rog usar care varweays 1 aa _ a ar 100). This part of meiosis in the apomicts differs IN no visible way from the corresponding stage in sexual plants. It is not possible to determine in this preparation whether or not the chromatin threads paire Fig. 3. Early prophase of meiosis in sexual diploid = incana (X 400). Synezesis apparently results during fixation at this stage. Fig. 4. Early proph 1ase of meiosis in apomictic, triploid T. incana (X 400). Synezesis is a prophase feature in both sexual and apomictic material. | r aw PLaTE VI. Phot of longitudinal sections of ovules in Townsendia (all X 400). Fig. 1. aces I with clumped and scattered univalents in apomictic, triploid T. incana. Fig. 2. Anaphase II in apomictic, triploid T. incana with three extra gro me of chromosomes excluded from sd two main nuclei a anaphase 7} 3 Fig. 3. A tetrad of 24, acne nuclei (4-nucled 1 apomictc triploid T. incana. Fig. 4. A tetrad of Beeston pacies with the nuclev the — end of the t trad sania ed by a space and a wall from en sie r hee megaspore nuclei, in ial, diploid T. incana. Embryo sac development is trisporic from pies of this type. THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 37 Plate VII. Phot of longitudinal sections of ovules and ovaries in Townsendia, Fig. 1. Dy: ad of sexual, diploid T. montana var. montana, seal a cell plate (X Mgt sl Fig. “3 bie of sexual, diploid Tincana, without cell plate (X 400). ‘his is the st frec sntly enc -ountered type of dyad in sexual mat ih Tir. a7 Dyad of ment scbepined T. Rothrockii (X 400). This is the only population of Townsendia known to have walls separating the dyad nuclei. Fig. 4. Ovule and ovary a the dyad stage in stennetc triploid T. incana (xX 185). Pirate VIII. Photomicrographs of longitudinal sections of ovules in apomictic, triploid T. incana (all & 400). Fig. 1. Dy: ad. Fig. 2. A very young 2-nucleate embryo sac which has — oe from an unreduced dyad. At the time of fixation the nucellar epidermis wa the process of disintegration and the embryo sac was expanding, —— the cavity a by ot i of the eee Fig. 3. A 2-nucleate mbryo sac with the nuclei a rophase. Fig. 4. A young 4- nucleate ‘dues sac This embryo sac is the result of wena si ea ae. a 4-nucleate coenanieel spore. Photomicrographs of longitudinal sections of ovules Townsen , the result of trisporic de- i ella IX. ATE (all & 400). y young 3-nucleate embryo sac velopment, in pina "diploid T. grandiflora the time of fixat the nucellar epidermis wa the process of disintegration e remnants of the fourth megaspore nucleus and part of the llar epidermis are evident the micropylar end of this embryo sac. Fig. 2 . ate embryo sa ich has resulted from trisporic randiflora - 4-n ryo sac diploid 7 ana. develop ment in sexual, - diploi dd: é ‘ hich has resulted fro om _tetrasporic development ual, - 4. 6-nucleate embryo sac in sexual, diploid "T gina flora. The six — e ‘the result of one mitotic division-series in a 3-nucleate embryo sac of the type re shown 40 ATE X. Photomicrograpl triploid T. incana (all & 4 at metaphase. Figs. 2 e synergids (fi the antipodals ha g ve alread JOHN H. BEAMAN ae g * ‘ ~~ ) 00) he egg cell (fig n serial sections thre y disintegrated. 1s of longitudinal s 0). Fig. 1. A 4-nuc ect ea ae SU i“ BY: uugh a ions of embryo sacs in apomictic, ate embryo sac with all four nuclei Pig ~ > 4 ~~ the unfused polars (fig. 3), anc single embryo sac. At this stage LATE X hot icr of longitudinal sections of embrvo sacs in Townsendia (all & 400). ae 1 A 12-nucle: er embryo sac in sexual, diploid 7. montana var. montana. Three large antipod: ul cells ar : Nadal in our sms iller antip odal cells farther toward the chalazal end were excludec 1 n ”, = = 3 = z - ao) O & yz > 5 or = 72 = a g : scies of t a § ies nani adjacent sec- have -wellideveloned ‘entipedal ee The egg cell was included in - ait on tion. The two conical-shaped synergid cells are immediately below the fusing =) i is figure. Fig The egg and fusing polar nuclei in apomictic, Arh sory : ; . Saale . ee i ‘ — C T. incana. Portions of the synergids are at the bottom of the figure. ba Bohne jacent sections of an embryo sac in apomictic, triploid «I. sncana. " : proembryo (fig. 3) is associated with fusing polar nuclei (fig. 4). BEAMAN Pirate XII. Photomicrographs of longitudinal sections of embryo sacs, with embryos, in apomictic, triploid Townsendia incana. Figs. 1-4 from florets at pr e-anthesis stages. Fig. 1. A 4-celled pe (x 350) scronepaiies (in another section) ‘ fusing polar nuclei similar to those shown in Plate fig. 4. One of the intact, conical- — synergid cells is at the lower right oF “a embryo. Figs. 2-3. Adjacent sec- in an embryo sac showing an 8-nucleate proembryo, accompanied by 4-nucleate parva (three of the endosperm nuclei were not included in this section) (X 350). At the time of papacy the three apical nuclei of this a had not been Sadan by walls. Fig. 4. A non- a section showing 3 of the 8 endosperm nuclei in this mbryo sac at metaphase ( 350). Fig. 5. A portion a the ovary and its contents from a floret just past anthesis (« 90). THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 43 been used most extensively, was grouped into five series accord- ing to the stages selected for examination and the conditions under which it was grown. Series I-IV were grown in Cambridge, Massachusetts and fixed in November, 1955. Series V was grown in Pullman, Washington and fixed in the spring of 1955. Townsendia incana (culture 6-1). The material of this series was taken from five plants. The developmental stages of the florets ranged from pre-meiotic to pre-anthesis. Early prophase of megasporogenesis is similar in appearance in sexual and apo- mictic forms (Plate V, figs. 2 and 4). Synezesis is as frequent in the early prophase stages in the apomicts as in the sexual material. Diakinesis is the earliest stage at which differences be- tween the sexual and the apomictic types become apparent. At this stage both megaspore mother cells with all the chromosomes unpaired and those with some pairing have been observed. In sectioned material it is often difficult to determine how many paper on the male reproductive phase. Megaspore tetrads are rare in this apomictic population of T. incana (cf. Table IV). The most normal-appearing tetrad which has yet been found is shown in Plate VI, fig. 3. It should be noted that no walls separate the megaspore nuclei. When re- duction occurs, embryo sac development is tetrasporic ( Plate VIII, fig. 4). Tetrads indicate reduction in the chromosome number. This fact is evident from the study of microsporogenesis in Townsendia. In this triploid, apomictic material, nuclei with a reduced chromosome number probably cannot function in the production of new embryos. It appears improbable that the few eggs which might develop with the reduced chromosome number would be fertilized by functional sperms, especially since the 44 JOHN H. BEAMAN TABLE III. Comparative classification of ovule development in si apomictically reproducing populations of three species of Fomusabae Developmental stage Numbers of ovules observed in the in ovule populations (cultures) 6-Ia? 6-II 6-Va 17-1 875 R27 798 a rial = VAI eine eee — 11 — 2 2 — 6 arly pret ata ie ice 14 53 10 28 5 x few rein Pall univalents) .... 1 — 1 — et Diakinesis (some pairing) ........ =~ 1 a — — eo ee Diakinesis (pairing not det.) ... — 3 6 9 4 1 3 3 Prometaphase (all univalents) 2 — oo — 1 eee Metaphase I (all univalents) ... 1 1 — — — Ee Metaphase I (s iring) .... 2 1 — — = ee Metaphase I (pairing not det.) a 1 3 4 —_ 1 2 1 Anaphase I — _ if 5 1 mere ig Me - 3-96 47 23 69 32 64 8 9 Dyad — micronuclei .......... ; 5 1 2 — — ae ee CE Dyad 3 a neue ue i Sarin cee SANE uc I PODER reo) Aree Youre 1 — — — os eS ee Anapnase Te ee 1 1 — — — —_ = Tetrad 4 5 2k 4 — ll Tetrad haba micronuclei. ........ 2 — — } — or eyes ee Double tetrad 1 ey pa — — Lo 2 1 _ — ee ree er 1-nucleate embryo oe eR Fe 6 —_ — — — short (ia eee ieee 1-nucleate Picak gies 12 — a _ — ae ee 2-nucleate ” OOS hae 110 ~— 30 Pee oo 2-nucleate éf eS ae — 1 —_ ON an ee (with micronuclei) 2-nucleate embryo sac ? ........ 19 3 — —_ a4 eons: 3-nucleate Z ee ie _— —_ — 1 seein eS om 4-nucleate 6h Ege rk etre 38 4 — — 3 — le aaa 4-nucleate de hss GBs cade ys _ —_— — — — —_ OS cam 6-nucleate “f gE SS — — — — ee ee 8-nucleate mm Wee ae oe 1 — — — —_ cosh mean ek al 8-nucleate a EER de ey er 2 — — — er Sooeste 2 — — — _ erases Genera ree eR Egg and seas pre Pe aN 83 = ound ci a SS ME i ae Egg and unfused polars ? ........ 1 — — a me Skee Egg and acae POMPE oc a — — _— — eae er ena g and fusing polars ? ........ y ae ten oe —_— — | et vo Embryo and fusing polars <:..;. cane pa _— — Se ert er Embryo and endosperm ............ 2 — — — a ee ORS pe we and endosperm ? ........ — —_— — a se AE gee Gs ssale Wea aa eet ts 3 — 1 _ 4 eas Not Ndbieodbais ie Sy Mats eS 100 2 2 17 : 8 4 Gs AO bViets eh vadaea edna aunciees 636 =174 56.435. 206 7 102.2 54 sendia incana (cultures 6, 17, and 875), T. Parryi (cultures 8 and 27), E: Rothrock (culture 798). ? Explanation of the culture-series symbols will be found in the text, pp. 43-50. nN ® i] a. THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 45 pollen develops in an anomalous manner by the formation of a wall around the four members of the microspore tetrad. The consequence of much chromosomal pairing in apomictic megasporogenesis probably is sterility. Asexual reproduction is possible only when all or nearly all the chromosomes occur as univalents. One expression of apomictic “meiotic irregularity” as a result of pairing in an apomict is shown in Plate VI, fig. 2. In this photograph of anaphase II, three small groups of dividing chromosomes may be seen in addition to the two main dividing groups. Each of the five groups of chromosomes has its own spindle mechanism. The three small groups of chromosomes would probably have become micronuclei. Both dyads and tetrads with micronuclei have been observed. Such irregularities must often prevent the formation of viable seeds, but the variable chromosome number in this apomictic population suggests that slightly aberrant chromosome races may survive. The great abundance of dyads in this apomictic form in con- trast to their rarity in sexual material (cf. Table IV) is one cri- terion of apomixis. In addition to the difference in numerical ratio of dyads to tetrads between apomictic and sexual forms, there are considerable morphological differences in the dyads of the two types. The dyad stage of the apomicts is one of long dura- tion during which nuclear-size increase, vacuolation, and a grad- ual breaking down of the nucellus occur. The dyad stage of the apomicts corresponds to the entire period in sexual forms from the dyad to the early female gametophyte. The development of dyads directly into two-nucleate em- bryo sacs was demonstrated by Holmgren (1919) in apomictic forms of two species of Erigeron. Townsendia apparently is the only other genus in which unreduced female gametophytes are now known to develop regularly in the same manner. It should be noted that the process by which a dyad becomes a two- nucleate embryo sac is a gradual one with no very sharply marked steps. Therefore, assignment of the material at intermediate stages to either the dyad or the two-nucleate categories is some- times difficult. In this study an arbitrary distinction, based on the condition of the nucellar epidermis, has been made between the two stages. When this structure is intact the material is con- sidered to be at the dyad stage. When the nucellar epidermis disintegrates, the two-nucleate embryo sac stage has been reache A few uninucleate embryo sacs have been found in this apo- mictic form of T. incana. They are rare, however, and the present 46 JOHN H. BEAMAN material is insufficient to permit an interpretation of their signifi- cance. Three- and six-nucleate embryo sacs present in this culture could be the result of trisporic embryo sac development. Most of the four-nucleate embryo sacs (Table III) are merely in inter- mediate stages between two- and eight-nucleate female gameto- phytes. However, a few are the direct result of tetrasporic de- velopment from the megaspore tetrad. A four-nucleate embryo sac of the former type is shown in Plate X, fig. 1. One of the latter type is shown in Plate VIII, fig. 4. The eight-nucleate stage of the embryo sac is short. The antipodal nuclei disintegrate almost as soon as they are formed. Hence, eight-nucleate embryo sacs are found only at the earliest stages of morphological organization of the female gametophyte. When the eight-nucleate stage is reached and the antipodals disintegrate, further developmental processes can no longer be designated as the eight-nucleate stage. Only five nuclei, the egg, the two synergids, and the two polars, are involved in subsequent developmental processes. But two major phases are distinguish- able during morphological organization of the female gameto- phyte. Before the polar nuclei have come together, a stage “unfused polars” may be designated (Plate X, figs. 2, 3, and 4 and Tables II] and IV). After they have come together, the stage “polars fusing” has been reached (Plate XI, fig. 2 and Tables III and V). Some less conspicuous changes of the egg and synergids occur during the migration and fusing process of the polar nuclei, but these changes are not obvious enough to use in designating developmental stages. In the apomicts the polar fusion process is not completed until after the first division of the egg or a few subsequent nuclear divisions in the proembryo (Plate XI, figs. 3 and 4 and Table VI). The fairly large amount of abortive and undetermined ma- terial in culture 6-I should be noted. This material was fixed after three nights with frost. Freezing may have adverse affects on megasporogenesis and female gametophyte development 1 apomictic as well as in sexual plants of T. incana. Abortion in sexual material, probably caused by freezing weather, was noted previously. Townsendia incana (culture 6-II). An examination of mega- sporogenesis in a single plant (6HF) of T. incana was made as a check against the other investigations which employed several plants. It is evident from this examination that both normal apomictic development and meiotic irregularities indicating fea- tures “abnormal” in apomictic reproduction may occur in a single THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 47 TABLE IV. Megasporogenesis in sexual and apomictic populations of Townsendia. Percentages of ovules with ssue at the fol- Chromo- No. of Arche- Dividing dyads tetrads some no., ovules sporial megaspore 2n examined cells mother cells Taxon Culture 1 Sexual T. incana 971 18 55 3.6 16.4 1.8 78.2 T. incana 5 18 97 — hi2 21 90.7 T. incana 802 18 58 1.7 19.0 39 75.8 T. montana 35 18 55 1.8 30.9 10.9 56.4 var. montana T. grandiflora 39 18 25 — 16.0 4.0 80.0 Apomictic Te incana 6-Ia 28 135 16.3 77.0 6.7 Ee incana 6-Il 28 126 8.7 48.5 38.1 4.7 T. incana 6-Va 28 45 44.4 55.6 — ap incana 17-I ca. 30 119 — 39.5 58.0 ZO T. incana 875 45 4.5 24.5 71.0 ea T. Parryi 8 36 80 peg Li 80.0 7 T. Parryi 27 36 — 46.7 53.3 — T. Rothrockii 798 ? 52 TES 50.0 17.3 21 1 Explanation of the culture-series symbols will be found in the text, pp. 28-50. plant. Diakinesis and metaphase figures were found in which some of the chromosomes were paired, and one metaphase was observed in which all the chromosomes were unpaired. The dyad- tetrad numerical ratio (Table IV) in this plant approximates that of the other plants from culture 6. Micronuclei were found asso- ciated with a dyad in one preparation of this material, and the same feature was observed several times in the plants of the mass collection. The study of a single plant suggests, therefore, that all the deviations from the usual apomictic reproductive cycle in the mass collection (culture 6-1) did not come from a single plant. Rather, in this apomictic population, various plants may have irregularities which result primarily from occasional chromo- some pairing. Total univalent formation, on the other hand, per- mits a regular apomictic “meiosis.” Townsendia incana (culture 6-IIT). The achenes used in this series were from florets in which the corollas had not opened. More advanced florets of the same heads had already reached anthesis. The use of heads with both open and unopen florets made it possible to obtain achenes from florets which would have very soon reached anthesis. The florets which had already opened on these heads were put in the next series to be discussed. Culture 6-III provides strong direct evidence for apomixis in 48 JOHN H. BEAMAN Townsendia. A very high proportion of precociously developed embryos were found in the ovules of these florets with unopened corollas (Table V). Some details of early embryo development have been listed in Table VI. These data indicate that the division of the egg ordinarily precedes the completion of the fusion process of the polar nuclei. In angiosperms the division of the primary endosperm nucleus normally occurs before the division of the zygote. This is not true in apomictic T. incana. Holmgren (1919) indicated that as far as he could discern, embryo and endosperm development proceeded concurrently in apomictic Erigeron. It may be found, however, that the same division sequence, with embryo development before that of the endosperm, will be found to be similar in apomicts of Erigeron and Townsendia. When the process of fusion of the polar nuclei is completed the first ensuing division apparently occurs immediately. In this material no completely fused polars, i.e., primary endosperm nuclei, were found. Wall formation in the endosperm probably occurs after the first or second mitotic division series. The nuclear number in the embryo at the earliest stages of development ex- ceeds that of the endosperm, but at slightly later stages the nu- clear number of the endosperm is greater than that of the embryo. Very little endosperm remains when the embryo has attained its maximum size. Embryo development appears to be in accordance with the asterad type. The early development of the embryo must be recorded by nuclear number rather than by cell number be- cause wall formation does not always immediately follow mitosis (cf. Plate XII, fig. 3). Mitotic divisions in the young embryo are not synchronous, but those in the early development of the endo- sperm are. The latter feature is illustrated in Plate XII, fig. 4. Townsendia -incana (culture 6-IV). This series was made from opened florets on the same heads from which unopened florets for the last series were taken. In most of the material at this stage of development the embryos and endosperm had at- tained a fairly advanced degree of development (Plate XII, fig. 5). The few ovules without embryos and endosperm already developed may have resulted from megaspores with the reduced chromosome number and probably would have aborted at slightly later stages. The ovules with embryos without endosperm an those with endosperm without embryos were rare and abnormal. Townsendia incana (culture 6-V). As a check on the repro- ductive behavior of this apomictic population of T. incana In a different environment, material grown in Pullman, Washington and fixed during spring rather than fall growing conditions has THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 49 TABLE V. pr on of ovule development just before anthesis and at or just r anthesis in apomictic populations of two species of Townsen net Percentages of embryo sacs with the contents listed below Taxon Culture 1 plac of Egg and Embryo Embryo Abor- Not mbryo fusing and and tive hn to 4 yon polars fusing endo- ined examined polars sperm Just before anthesis incana 6-IlI 202 11.9 6.9 68.3 9 3.0 T. incana -Vb 5 6 46.8 26.6 T. incana 17-II 8 12.5 — 25.0 1 se, 50.0 T. condensata 47 SZ 6.3 3.1 593 6.3 25.0 At or just after anthesis . incana 6-IV 144 21 1.4 82.6 10.4 E Ho T. incana 6-Vec 22 — ae 90.9 9.1 — * Explanation of the culture-series symbols will be found in the text, pp. 47-51. TABLE VI. eee a oe stages of embryos and endosperm just before anthesis in apomictic aioli incana, Culture 6-III.* Developmental stage * Number observed 2-nucleate embryo and fusing polars 4-nucleate embryo and fusing polars 3 2-nucleate embryo and 2-nucleate endosperm l 4-nucleate embryo and 4-nucleate endosperm : 1 1 4-nucleate embryo and 16-nucleate endosperm 5-nucleate embryo and 4-nucleate endosperm 7-nucleate embryo and 4-nucleate endosperm dosperm 8-nucleate embryo and 4-nucleate en osper 1 i al and endosperm at various later stages at } Explanation of the culture-series symbols will be found in the text, p. 47. * Embryos are listed by nuclear ders than — number because walls do not always develop immediately after the mitotic divisi been utilized. The series was divided into the subseries a, b, and ture-series 6-I, 6-II, 6-III, a 6-IV with 6-V in Tables III, IV, ad V) suggests that environmental factors cause no significant differ- ences in apomictic reproduction in a single bio Townsendia incana (cultures 17 and 975). Aecanttic popula- tions from the northern portion, the southern portion, and about the middle of the range of the species were mie in this ed to determine what intraspecific variation occ apomictic reproduction. A comparison of the data (Tables Tl and 1V) from these different populations of apomictic T. incana suggests 50 JOHN H. BEAMAN that no major and very few minor differences in the apomictic process occur between populations. Culture 17 was divided into two series, I and II, according to developmental stage, for con- venience in tabulation. Townsendia Parryi (cultures 8 and 27). Most of the study of apomixis in Townsendia has been made in material with a triploid or triploid-derived chromosome complement, but it was con- sidered essential also to employ some material with the tetraploid chromosome number. Therefore, investigations have been made in two tetraploid populations of T. Parryi. The apomictic process in these plants is very similar to that observed in triploid and tetraploid populations of T. incana. In T. Parryi, as in T. incana, the number of univalent chromosomes at diakinesis or metaphase in megasporogenesis is difficult to determine, but numbers ap- proaching 36 may be counted. A double tetrad in one ovule of this material may have resulted from the functioning of two archesporial cells instead of the usual single cell. Some of the integumentary cells in these ovules have a more sporogenous appearance than those in any other Townsendia species which has been examined. Townsendia Rothrockii (culture 798). Microsporogenesis in this population is so irregular that no approximation of the chromosome number has been obtained (seeds for root-tip chro- mosome counts were not available). Most features of megasporo- genesis in these plants are similar to those in apomicts of the other species. However, the dyads and tetrads are somewhat different from those encountered in other material. The dyads are charac- terized by a wall between the two nuclei. No other population of Townsendia is known which has wall formation between the dyad nuclei. Wall formation in the megaspore tetrads of this species also is peculiar. The nuclei of the tetrad have been num- bered arbitrarily 1, 2, 3, and 4, in order, proceeding from the micropylar to the chalazal end of the ovule. A single wall has been found in some tetrads between nuclei 1 and 2, in some be- tween 2 and 3, and in some between 3 and 4. In other instances two walls are present, between nuclei 1 and 2 and between 3 and 4. For an apomictic population, the percentage of tetrads formed by these plants is very high. It seems possible that the structures which appear to be tetrads actually are not. Instead, two adjacent archesporial cells may have given rise to dyads and these merely appear to be tetrads because of their adjacent position. More material must be examined before this problem can be satisfac- torily solved. THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 51 Townsendia condensata (culture 47). A very limited amount of this material was available. The present investigation has demonstrated only that embryos and endosperm are formed in ovules before the corollas open, and evidence of apomixis in the population is thereby obtained. THE CAUSAL ASPECTS OF APOMIXIS IN TOWNSENDIA No data on the genetic control of apomixis in Townsendia have been obtained. Therefore, the only information on the causes of apomixis in this genus must now come from inference. The special conditions under which apomixis is expressed in Townsendia are suggested by the geographical distribution pat- terns of the sexual and apomictic forms. Without exception, the apomicts of Townsendia are concentrated at the higher eleva- tions and higher latitudes relative to the total range of the species in which they occur. It seems scarcely deniable that a cold, rigorous climate is a very important factor in permitting the ex- pression of apomixis in Townsendia. A correlation is found in Townsendia (as apparently in all groups with apomicts) of the perennial habit and apomixis. This correlation is shown in Chart 1. Gustafsson (1948) and Stebbins (1950) have reviewed the current ideas on the relationship of the perennial habit with polyploidy. Stebbins’ interpretation from the available evidence is that the longer growth period of peren- nials, compared with annuals or biennials, permits a longer time during which a sterility “bottleneck” may be overcome after the initial polyploidization. No experimental attempts have : made to induce apomixis in plants of Townsendia by increasing the chromosome number, and it seems futile, therefore, to specu- late on the importance of the increase in the chromosome number per se in permitting apomixis. However, the lack of correlation of apomixis with hybridization in the genus and the apparently frequent independent recurrence of apomixis in many popula- tions suggest that little more than polyploidy may be required to release the potential apomictic mechanisms. If apomixis is immediately expressable when the polyploid state has been at- tained, the perennial habit would hardly be necessary for the prevention of a sterility “bottleneck.” It is generally recognized that the perennial habit, rather than the annual or biennial habit, is correlated with cold, rigorous climates. Apomixis in T ownsendia also is associated with this pe of climate. Perhaps the correlation of apomixis in Town- sendia with the perennial habit is merely coincidental in that both 52 JOHN H. BEAMAN features are more or less independently correlated with a cold, rigorous environment. The two features cannot be entirely inde- pendent, of course, because apomixis must be of some adaptive advantage to the plants in which it occurs. However, its origin could be merely incidental to the perennial habit. The fact that apomixis occurs in some biennial plants of Townsendia as well as in perennial plants supports to some extent this view. TOWNSENDIA AS AN AGAMIC COMPLEX From the investigation of Babcock and Stebbins (1938), the agamic complex (a term defined by these authors) in the Amer- ican species of Crepis has become well known. Their work may therefore serve as a standard of comparison for evaluation of the agamic complex in Townsendia. Stebbins (1950) has pointed out that from the systematic and phytogeographic points of view, Crepis is easier to understand than most of the other known agamic complexes. American Crepis is confined to one geo- graphic and climatic zone, and is so recent that the diploid species which gave rise to the polyploids are still growing in the same area. These features are exactly duplicated in Townsendia, but similarities extend little farther. n Townsendia it appears that hybridization may occur en- tirely independent of polyploidy. No polyploid system is neces- sary to permit the exchange of genes between the species. Likewise, in Townsendia hybridization seems to be an entirely unnecessary feature for setting in action apomictic reproduction. In spite of frequent hybridization between the diploid forms of the species of Townsendia there are as many, if not more, sus- pected populations of autoploid apomicts as alloploid apomicts. All of the presently known apomicts of T. Rothrockii, T. scapigera, THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 53 F i strigosa, and T. grandiflora are apparently autoploid. Town- sendia Parryi and T. Hookeri have a very high percentage of auto- ploid apomicts, and even T. leptotes, T. exscapa, and T. montana seem to have numerous autoploid apomicts. In none of the species do all of the apomicts appear to be alloploid. Thus, apomixis in Townsendia, apparently unlike that in Crepis, can be viewed independently from hybridization. The only reason that hybrid- ization appears to be involved in the apomicts of Townsendia is because some of the diploid, hybrid-derived types have become polyploid apomicts. Probably some of the gene recombinations which result from hybridization are more vigorous than the non- hybrid types, and when these are fixed by apomixis they have an advantage over the non-hybrid types and thus become wide- spread. Otherwise, hybrid apomicts in Townsendia might not even be known. montana, T. Parryi, and T. Rothrockii, do the sexual forms have a very limited and possibly relict distribution. Even T. Parryi probably should not be included in the above group because its sexual forms have both a considerabie geographic range and interpopulation variability. In the other species of Townsendia which have apomixis, the sexual forms have much morphological diversity and often are nearly as widespread as, or even more widespread than, the apomictic forms. Thus, even though the American species of Crepis are similar to the species of Town- sendia in occupying a single geographic and climatic realm, and in having the sexual precursors of all the species known, the two agamic complexes have few other common features. A limited evolutionary future for the apomicts of Townsendia may be postulated, as the embryological data indicate that apomixis is obligate in the genus. The statement by Stebbins (1950, p..417) that agamic complexes are “blind alleys” is prob- ably better applicable to the Townsendia apomicts than to the facultative apomicts of many other genera. The best evidence that the apomicts of Townsendia have not given rise to any new apomictic forms lies in the fact that no species of the genus is represented only by apomictic types. Sex- 54 JOHN H. BEAMAN -ual precursors of all the apomictic forms are known, which is positive evidence that the apomicts have undergone little if any evolution. Although their evolutionary future may be limited, the suc- cess of the apomicts in colonizing wide areas is evident. Apomixis occurs in 12 of the 21 species of Townsendia; it occurs abundantly in nine species. The geographic range of the apomicts in some of these species exceeds considerably that of the sexual forms. Therefore, apomixis must have potent adaptive advantages. TRENDS OF EVOLUTION DIVERGENT EVOLUTION The patterns of speciation in Townsendia seem explainable mainly on a geographic, climatic, and edaphic basis. A correlation of the morphological traits of the taxa of Townsendia with their geographic distributional-patterns makes possible a suggestion of their phylogenetic relationship. A tentative phylogenetic dia- gram is presented in Chart 1. The center of this diagram may be imagined as the time when diversification in Townsendia began. 4 bal T. INCANA T FENDLERI z T MEXICANA 2 T. STRIGOSA 3 R 3 T ANNUA T. FORMOSA 3. CAULESCENT ANNUALS, T TEXENSIS BIENNIALS, OR PERENNIALS 2. T GRANDIFLORA -— ve ade & — T GLABELLA _— Z Py aa T ROTHROCKII h one T. PARRY! m4 ste eee T MONTAI 5 ae VAR. MONTANA T FLORIFER eaten ‘ T MONTANA 2 ee VAR. MINIMA > areal T. MENSANA 3. 21 SCAPIGERA VAR. MENSANA -— T MENSANA 4 VAR. JONE SII ACAULESCENT, MOSTLY : t LONG-LIVED, PERENNIALS T LEPTOTES T CONDENSATA T HOOKER! ‘ d T. SPATHULATA 4 T EXSCAPA 4 APOMIXIS FREQUENT 2 APOMIXIS INFREQUENT 3. APOMIXIS UNKNOWN Cuart 1. Phylogenetic relationships of the species of Townsendia and a summary of “the habit and the presence of apomixis in the species. THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 55 veloped from the others. At the periphery of the diagram the species have been placed, in so far as feasible, closest to the species which their development parallels. The broken line through the center of the figure separates the caulescent from PARRY! ~~~~. 7 os a3 / CONDENSATA tn ‘ / ? ‘ , ‘ FLORIFER ng i SPATHULATA \ ‘ D \ / STRIGOSA . SCAPIGERA i MONT. vor as 4 i Papin HOOKER! iN ! MONTANA wen o> ‘ ' MEN LEPTOTES H | MENSANA, SANA . i vor. INC ANA SRANDIFLORA 5 | JONESi! : : FEN H ‘ ROTH LER! ' ' ROCKII H ' | ; \ MONTANA var, GLA ’ MINIMA BELLA o u ee ee au es EXIMIA . . TEXENSIS ANNUA \ ? ‘ EXSCAPA| “o> ahi. .S FORMOSA MEXICANA Map 1, bas Phylogenetic relationships of the species of Townsendia on a geographic is. 56 JOHN H. BEAMAN the acaulescent types. The relative amount of apomixis in the various species has been indicated. The phylogenetic relationships of the species on a geographic basis are suggested by Map 1. The arrow-points indicate derived species, and where there are no arrows terminating the lines, the possible derivations are not suggested. The broken lines indicate indistinct relationships between more primitive types. The extensive climatic variations of the Pleistocene must be acknowledged as important factors in the phylogenetic history of Townsendia. It seems probable that some of the species as they are now known were not in existence during much of the Pleisto- cene. The high level of correlation of some species with special- ized edaphic and climatic conditions suggests that climatic changes could have easily taken their toll of various evolutionary experiments. On the other hand, the relative abundance of Townsendia species now in habitats which were unavailable dur- ing the last glacial advance could indicate both the recent evolu- tion of some new types and the ability for rapid migration of some of the older forms. An understanding of the evolutionary trends in Townsendia requires an understanding of the primitive characteristics in the genus. Townsendia formosa seems to qualify in all respects as the most primitive member. The probable primitive generic characters are outlined in the section on generic relationships. The species with the greatest morphological similarity to T. formosa is T. eximia. The latter species most conspicuously differs from the former in having a shorter life-span, in develop- ing a branched stem, and in having a taproot rather than a fibrous- root system. These species occur allopatrically in adjacent moun- tain masses mostly in New Mexico. Townsendia eximia has close relatives on all geographic sides of its range. To the east and north, respectively, are the very close cognates T. texensis and T. grandiflora. Townsendia glabella, which occurs only a short distance to the northwest, is similarly closely related to T. eximia. A continuation of the reduction trend which has produced T. glabella is found a short distance to the north in T. leptotes. These two species occur together in southwestern Colorado, but T. leptotes is more diversified to the north of the range 0 T. glabella. Differentiation through isolation probably occurred while the two populations were isolated from each other by the mountain masses in western Colorado, and the populations of T. leptotes within the range of T. glabella in southwestern Colo- rado are probably recent migrants there. THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 57 Populations of the T. leptotes type which migrated to the east side of the Rocky Mountains in Colorado have been geographi- cally isolated there and the resulting evolutionary divergence has produced T. Hookeri. Both of these species are advanced types; one obvious feature indicating their advanced nature is the ex- treme reduction of the aerial portions of the plant body. It should be noted that the sexual forms of T. Hookeri and T. leptotes are entirely allopatric, but the apomicts of T. Hookeri occur in the range of the sexual forms of T. leptotes. The apomicts of both species have wide geographic ranges. Speciation from the T. eximia-T. glabella stock can also be traced geographically to the west of the range of that evolution- ary complex. Townsendia montana (both varieties) exhibits a fairly close morphological relationship to the T. eximia-T. glabella stock and probably has resulted after geographical isolation in the mountainous areas of central and north-central Utah. Differ- entiation may have occurred in isolated populations at both high and low elevations; the high-elevation type has become T. mon- tana, while evolution in the populations at lower elevations has resulted in T. mensana. The evolutionary trends in specialization from a more general- ized T. eximia stock have occurred in the mountainous areas to the northeast, north, and northwest of the range of T. eximia. Such a pattern is probably attributable mostly to the fact that a large number of habitats suitable for occupancy by Townsendia populations occur in that area. Because of the mountainous terrain, the populations were easily isolated from each other, and climatic changes probably have facilitated the isolation. The divergences apparently have been recent enough that the morpho- logical trends they followed are still evident. Many connecting “links” may not yet have become extinct in spite of numerous Pleistocene climatic fluctuations. Speciation with a different geographic pattern from that out- lined above has had an orientation from the northern rather than 58 JOHN H. BEAMAN the last ice advance in the area which T. florifer and T. scapigera now occupy, it must be inferred that these two species have ha a recent origin. Townsendia Parryji, their closest relative, is in a geographic position which further indicates that this evolution must have been recent. The geographic trend in evolutionary specialization along this line is from the northeast in Montana to the southwest in Nevada. It can best be interpreted as a morphological reduction series which occurred only in one direc- tion. The advanced forms probably have spread since the last glaciation into more arid regions from an area relatively near the present range of T. Parryi. A different evolutionary trend from T. Parryi stock appears to have occurred in a southeastern direction from the present range of that species. Townsendia condensata and T. spathulata are the specialized members along this line of development. They appear to have become strongly adapted to some of the peculiar edaphic situations of western Wyoming. Townsendia condensata may be a relict type of a former colder and more moist period, and it is now represented mostly by alpine apomicts. Edaphically, T. spathulata is one of the most highly specialized species of Townsendia. Its unusual habitat requirements probably are a reflection of its reduced morphological features. Very likely it could not withstand the competition from larger plants on sites more than sparsely vegetated. Part of the evolutionary line which may have diverged from a T. Parryi prototype is not so closely related to that complex as the two trends just outlined. However, the ancestral forms which gave rise to T. strigosa and T. incana probably occupied the Colo- rado Plateau and Green River basin area and may have been somewhat similar to T. Parryi. The T. strigosa-T. incana stock probably migrated into the central highlands of Mexico and, wit isolation, T. mexicana has arisen. The separation of T. strigosa and T. annua into distinct taxa appears to have been a recent occurrence. Townsendia incana and T. Fendleri similarly are cognate species. tks The clearest patterns of speciation in Townsendia indicate that geographic isolation has been of prime importance In the evolution of the members of the genus. A position of importance for polyploidy in the evolutional pattern seems to be ruled out, as no polyploid sexually reproducing species are known to exist. The role of hybridization in the development of new forms m Townsendia seems relatively unimportant, but the process does occur and cannot be entirely excluded as a possible evolutionary THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 59 agent. However, none of the present species is suspected to be of hybrid origin. The importance of geographic isolation in speciation in Townsendia is further emphasized by the unique case of T. ex- scapa. It is pointed out in greater detail in the systematic treat- ment that this species occurs in an area where Pleistocene climatic fluctuations probably would not break up its range into geo- graphically isolated populations. Townsendia exscapa has there- fore not become differentiated into segregate species but rather is a single large species showing clinal variation. From both the negative approach (with T. exscapa) and the positive approach (with the other species ), the role of geographic isolation in the development of the species of Townsendia i striking. Other factors, such as polyploidy and hybridization, may be important evolutional factors after a group has become some- what differentiated through geographic isolation. In the young and actively evolving species of Townsendia the primary impor- tance of geographic isolation in permitting the fixation of inde- pendent gene mutations seems amply clear. CONVERGENT EVOLUTION Convergent or parallel evolution is a feature of some of the species of Townsendia. This phenomenon is forcefully illustrated by mixed collections which are often unwittingly made by collec- tors when two species grow side by side. For example, the apo- mictic plants of T. montana var. montana and those of T. leptotes which occur in the high mountains of northern Utah and north- central Idaho are so similar that they have been confused in the field by five collectors (including myself). A careful examination of the plants and a knowledge of the total range of variation to be expected in each of the species made it possible to show that two specific elements were involved. Perhaps the instance of convergent evolution in Townsendia which has misled the most botanists is between T. exscapa and T. Hookeri. When Hooker established the genus he did not real- ize that his material included two species. Ninety-three years later Larsen (1927) first recognized that Hooker had based his “species” on two different specific elements. Specimens from the central portions of the ranges of the sexual forms of these species are easily distinguishable. In the cases of parallel evolution in T. montana and T. leptotes and in T. exscapa and T. Hookeri, hybridization may have been the factor which permitted convergent evolution. Strong selec- 60 JOHN H. BEAMAN tive forces in the rigorous habitats these plants occupy probably permitted only a very limited number of morphological (and physiological) recombinants to survive. These happened to be nearly intermediate forms between the two species. Successful recombinations between the species are now fixed by apomixis. An instance of convergent evolution which has probably in- volved hybridization but not apomixis is found between T. Fend- leri and T. annua. These species were similar enough that Larsen was confused and treated them as a single species in her revision of the genus. A mixed collection of the two species has occurred at least once (cf. discussion of T. Fendleri in the systematic treat- ment). Two cases of convergent evolution which appear not to have involved hybridization are between T. mensana var. Jonesii and T. scapigera on the one hand and T. mensana var. mensana and T. Hookeri on the other. In the former case, the populations of T. mensana var. Jonesii in the Charleston Mountains of Nevada are superficially very similar to some of the plants of T. scapigera in the Inyo Mountains of California. There is no evidence that these populations of the two species have ever hybridized. Rather, it seems probable that the very similar climatic regimes in the two areas have been responsible for the selection of life- forms which have a considerable degree of morphological similar- ity. One is led to assume that the genetic complement of the genus is such that it will permit the expression of only a limited number of morphological types under a given set of environ- mental conditions. In the case of the convergent evolution of T. mensana var. mensana and T. Hookeri, the possibility of hybridization seems to be very clearly ruled out. In the Uinta Basin of Utah where this convergence occurs, the population of T. Hookeri is apomictic. Sexual forms of T. Hookeri are known only on the eastern side of the Rocky Mountains and it seems probable that the popula- tion of this species in the Uinta Basin migrated there as an apo- mict (this postulate is given fuller consideration under the discussion of T. Hookeri in the systematic treatment). These apo- mictic plants have been able to survive because their basic geno- type was already suited for the cold winters and dry summers and peculiar edaphic conditions which occur in the Uinta Basin. The narrowly endemic T. mensana var. mensana, in contrast, 1s a sexual form which apparently has developed under the influence of the factors of natural selection in the Uinta Basin. The similar- THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 61 plasticity must have made possible the rapid divergent evolution in Townsendia. HysripizATION Experimental F, hybrids have been obtained between the seven species of Townsendia which are indicated on Chart 2. All crosses attempted were successful. One plant of the cross T. florifer < T. incana and one of the reciprocal were grown to maturity. These had neither meiotic irregularities nor a greatly reduced pollen fertility (cf. Beaman, 1954). Conditions at the time the F, plants were available did not permit securing the F, crosses, so none have been made. Plants of the other F, crosses were grown for some time under greenhouse conditions in Cam- bridge, Massachusetts, but growth of both hybrids and non- hybrids was unsatisfactory. From the few crosses which have been obtained, it appears that genetic barriers are lacking be- tween those species involved, but further experimental evidence is needed. Most of the crosses attempted were between relatively closely related species (compare experimental crosses, Chart 2, with phylogenetic relationships, Chart 1). A greater number of cross- ing attempts, with special emphasis on obtaining hybrids between the more distantly related species, is contemplated as a future project. Such crosses might provide evidence in addition to that of comparative morphology for inferring evolutionary relation- ships. Also, extensive crossing experiments could provide im- portant data on the nature of genetic barriers between species. With a closely related series of species, such as those of Town- sendia, it might be possible to correlate levels of morphological differentiation with the development of genetic isolating mecha- nisms. Thus a better insight might be gained on the nature of this important factor in speciation. ae Evidence for natural interspecific hybridization has been out- lined in the systematic treatment under the discussions of the species. The species between which natural hybridization is sus- 62 JOHN H. BEAMAN HYBRIDIZATION IN TOWNSENDIA PUTATIVE NATURAL FENDLERI| ————————_ ANNUA GRANDIFLORA STRIGOSA sex op. EXIMIA EXPERIMENTAL INCANA wo CONDENSATA _ MONTANA INCANA FLORIFER var, MONTANA, CONDENSATA HOOKER} PARRY}! op. sex. op. PARRY! EXSCAPA op. sex. op. FLORIFER LEPTOTES sex. sex. MONTANA var. MONTANA sex. ap. MENSANA vor, JONESI! SPATHULATA Cuart 2. Hybridization in Townsendia THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 63 bers should be involved in hybridization. To understand the basis of this peculiarity, it is natural to inquire why the other species are not involved also. Hybridization is not known in T. formosa, T. glabella, T. mexicana, T. Rothrockii, T. scapigera, and T. texensis. Most of these species are geographically isolated from all the other species. Townsendia glabella and T. scapigera are the only two members of this group which, on a geographic basis, might be suspected of being involved in hybridization with other species. Neither of these is well enough known to permit a sufficient understanding of its population diversity. The dis- tribution of natural hybridization among the species suggests that the principal requirements for its occurrence are an overlapping or adjoining of ranges of the species and a coincidence of flower- ing time. Sexual hybrids occur in a greater number of species than do hybrids in which apomixis is involved (Chart 2). From their dis- tribution within the species, it appears that in Townsendia hy- bridization and apomixis are independent phenomena. Hybrid- ization probably is not necessary to permit the expression of apomixis. However, certain hybrid derivatives which become apomictic probably have a greater survival value than some non- hybrid apomicts. It seems probable that in Townsendia hybridization is not a function of intermediate edaphic conditions. Most of the species which occur together have similar, rather specialized, edaphic requirements. Thus it appears that soil conditions figure prin- cipally in permitting species to grow together where they can hybridize. Although hybridization occurs in numerous species, there is little or no evidence to indicate that it is an important factor in speciation in Townsendia. It seems mainly to have increased the morphologica! diversity of certain species where it is coupled with apomixis. Sexual hybrids are most abundant between T. ex- scapa and T. Hookeri, and may be abundant between T. florifer and T. mensana var. Jonesii. Otherwise they are not frequently encountered. The diversity in the species as a result of hybrid- ization seems slight in comparison to that which has resulted from geographic isolation. MORPHOLOGY Townsendia is similar to many other genera of the Compositae in that its members have considerable microscopic homogeneity. Gross morphology is more useful than minute structure In deter- 64 JOHN H. BEAMAN mining specific limits. Proper determination of the taxa usually must be based on several characters from various aspects of the plant. One species of Townsendia has been the subject of an ana- tomical study by Bunton (1910). Her investigation was directed at determining in T. exscapa the anatomical adaptations to xero- phytism. HABIT. Two trends occur in the genus in the modification of the habit. Erect, monocephalous stems and a rhizomatous basal mat, a primitive and unspecialized habit, are found only in T. for- mosa. One reduction trend has resulted in low, densely rosulate, matted or tufted perennials, as in T. Hookeri. Another trend in- volves less shortening of the stem, but the life-span of the plant is reduced, as in T. annua. roots. All but one of the species of Townsendia have tap- roots; the exception is T. formosa which has a fibrous root-system. The taproots of young plants are slender and light in color; in older plants they become thick and woody but never develop a thick, rough bark. A gradual transition zone is usually found between the root and stem, and leaf-scars best indicate the change from root to stem. The biennials are characterized by an enlarged root-stem junction which seems to develop during the rosette stage of the first year of growth; this swollen portion may be hol- low or have a chambered pith. steMS. The perennial species usually have a branched, woody caudex which forms a crown at the ground surface. Herbaceous stems may develop from the caudex branches, as in T. glabella, or they may be terminated only by tufts of leaves and flowering heads, as in T. Hookeri. The caudex branches of the short-lived perennials are slender; those of the longer-lived species become correspondingly thicker. In most of the species, the stem is ex- panded just below its junction, with the head. This condition 1s seen most clearly in T. Parryi. Some members of the genera closely related to Townsendia exhibit the same feature. A few 0 the species of Townsendia are characterized by peduncles, but this character is not prevalent in the genus. Usually the heads terminate more or less leafy stems. eaves. External leaf morphology in Townsendia is remark- ably uniform. Throughout the genus leaf modifications are minor ones in shape, pubescence, and texture. The insertion is alternate and there is no strong differentiation between blade and petiole. The leaves are expanded at the junction with the stem, narrowe into a petiole-like portion, and expanded again into the blade. THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 65 With but two minor exceptions, the margins are always entire. Two minute notches infrequently occur at the apices of the leaves of T. Parryi, and in T. mexicana one or two small lobes are some- times found near the apices. In the acaulescent species all the leaves on a single plant are essentially the same size and shape. Caulescent plants have a basal rosette of leaves longer and broader than those of the upper portions of the stem. There is little differentiation between upper and lower leaf surfaces in any of the species. Thickened leaves are found mostly in plants growing at moderate to high altitudes, as in T. Rothrockii, but some species frequenting the higher elevations do not have thick- ened leaves. Involute leaves occur primarily in the narrow-leaved forms. PUBESCENCE. Both the stems and foliage in the members of Townsendia are characterized by a strigose pubescence. The trichomes are usually few-celled (3-8) and have a somewhat enlarged basal portion and a sharp-pointed apex. Taxonomically, the most useful pubescence variation is in density. Excluding the achenial hairs, there are only two variations in trichomes in the genus which are not merely quantitative. Simple, multicellular trichomes, conspicuous because of their very short cells, are found on the upper portion of the stem in T. formosa. In T. condensata and T. spathulata the trichomes are very long, with elongate cells which have conspicuous end walls; these trichomes may be very abundant, giving the plant a woolly aspect. The phyllaries may be strigose, pilose-strigose, or glabrous. In several species the trichomes of the phyllaries tend not to be appressed in a single, apical direction while stem and leaf tri- chomes almost invariably are apically appressed. INVOLUCRE AND RECEPTACLE. Generally, the involucre in Town- sendia is hemispheric or campanulate, but in T. montana and T. mensana it may be somewhat obconical. Phyllary morphology comprises a useful set of taxonomic criteria in the genus. In addition to the number of series in which the phyllaries occur, overall phyllary shape, form of the apex, and nature of the margin are useful characters. In species with very broad phyllaries, the apices tend to be obtuse, while in those with narrower phyllaries, the apices are acute, but acuminate apices occur both in species with wide and with narrow phyllaries. Only two species, T. eximia and T. grandiflora, are characterized by conspicuously bristly phyllaries. The margins always are at least somewhat scarious and ciliate or lacerate-ciliate and may serve as a useful character to someone familiar with the group, 66 JOHN H. BEAMAN but characters of the margin are too intangible to be useful in the key. Characteristics of the receptacle have proved useful primarily in the demonstration of the generic relationships of Townsendia. A conical receptacle is found only in T. formosa, the most primi- tive member of the genus; all the other species have nearly flat or slightly convex receptacles. The close generic relatives of Townsendia have conical receptacles. FLOWERS. The pistillate ray-florets occur in a single series. As is characteristic in the heterochromous members of the As- tereae, the species of Townsendia have only cyanic pigments in the ray-corollas, and ray color is often a useful taxonomic charac- ter. Uniformly blue or bluish-purple rays occur in four species and in some populations of two others. In most of the members of the genus, the rays are white on the adaxial surface and have a darker abaxial surface, often with a median stripe which may be pinkish, lavender, or mauve-purple. In old or dried rays the color usually is darker. The adaxial surface of the ray-corollas is generally glabrous, but the abaxial surface in several species is beset with few to numerous gland-like hairs, which apparently are under the control of very few genes. Care must be observed in their taxonomic use, but they are a constant feature of a few species. The length of the ray-corollas is sometimes helpful in delimiting species. The width, on the other hand is of little use. In some of the apomicts the rays never fully expand. The only character of the disk-corollas which has been used in this study is their length. A comparison of the length of the pappus to that of the disk-corollas has been helpful in several instances. The style-branches of Townsendia are typical for a member of the Astereae. The disk-styles have the upper portion of the branches modified with projecting hairlike cells, while the ray- styles have marginal stigmatic surfaces extending nearly to the apices of the branches. There are slight differences in shape of the style-branches between some of the species, but these are of no use for distinguishing closely related or nearly similar species. Neither anther structure nor morphological characteristics of the pollen has been employed as taxonomic criteria in this in- vestigation. However, pollen studies have been of great value in distinguishing specimens from sexual populations from those rep- resenting apomictic populations. The peculiarities of pollen formation in the apomicts are considered in the section on repro- duction. Pollen of the sexual plants (see Plate IV, fig. 1) is THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 67 spherical, three-pored, and uniformly spinescent. It is three- nucleate at anthesis. At maturity its mean diameter ranges be- tween 23 and 30 microns. ACHENES AND PAPPUS. There is considerable diversity in size and shape of the achenes of the species of Townsendia. Within the species the achenial morphology is relatively uniform, and in some instances provides useful taxonomic characters. Amon the more problematical species the achenes are so similar that they are of little systematic value. Achene shape varies from narrowly oblanceolate to obovate, and the achenes are always somewhat compressed. They usually have ribbed or callous- thickened margins. Gray (1880) in his synopsis of the genus placed considerable reliance on the achenial hairs in delimiting species and groups of species. In the present study also the character of these hairs has been found useful. Macloskie (1883) examined the achenial hairs of all the species of Townsendia known at the time of his study, and the descriptive terms which he and Gray used have been employed in this paper. The achenial hairs are always du- plex; two connate cells project from the surface of the achene. One cell originates at the surface of the achene, while the other joins an enlarged or unenlarged basal cell. A second basal cell might occasionally be present but apparently is mostly reduced or absent. The distal portions of the two principal cells may re- main connate and be acutely terminated or one cell may exceed the other and have an acute apex. Hairs with acute apices are termed “entire” in this treatment. Hitchcock and Thompson (1945) noted that the hairs of T. condensata (called T. spathulata in their paper) are simple. However, the duplex condition is retained in the hairs of this species, and the term “simple” is misleading. In some species of Townsendia the achenial hairs are ifurcate or emarginate, but glochidiate hairs are commonest in the genus. In the latter type, the distal ends of the two elongate cells are separate and recurved. There is no evidence in Town- sendia of a glandular nature of the achenial hairs. The pappus has been a much-used taxonomic character, and within limits it has great value. It cannot be used, however, to the exclusion of other characters for distinguishing species. The pappus is always uniseriate and is made up of connate setae which are usually arranged in well-developed, rather stiff bristles. At frequent (sometimes very frequent) intervals along the bristles, individual setae project out as non-recurved or rarely recurved barbs or barbulae. In T. exscapa the bristles may reach a length 68 JOHN H. BEAMAN of 13 mm. while in T. formosa the longest never attain a length of 1.5 mm. The pappus of T. eximia usually consists of two bristles and a crown of squamellae, but as many as 50 bristles are found in T. exscapa. The ray-pappus of several species is much shorter than the disk-pappus; the former sometimes is so short that the connate setae merely form a ring of coroniform squamellae around the apex of the achene. The short ray-pappus is a constant feature in some species but mostly there are excep- tions which make it an unreliable character. In T. mexicana the ray-pappus appears to have been reduced mostly to single, sepa- rate setae which frequently are somewhat glochidiate, resembling the achenial hairs. MEASUREMENTS. In this study the measurements have been based on dried material. Boiling was employed only in the width measurements of the ray-corollas. Small objects, such as phyl- laries, were measured under 9 X magnification with a millimeter rule. Very small parts, such as ray-pappus squamellae, were measured with the same rule under 54 X magnification. Pollen measurements were obtained with a compound microscope and a.calibrated ocular micrometer. SYSTEMATIC TREATMENT In the citation of specimens, herbarium abbreviations in the third edition of “Index Herbariorum” (Lanjouw and Stafleu, 1956) have Royal Botanic Gardens, Kew (xk); Herbario Nacional del Instituto de Biologia de la Universidad Nacional de México (Mexu); Uni- versity of Michigan (micu); Missouri Botanical Garden (MO); Montana State College (mMonT); Montana State University (MONTU) ; termountain Herbarium of the Utah State Universi c); Unive sity of Wisconsin (wis); State College of Washington (ws); University THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 69 specimens were examined in this s : The symbol “*” before an herbarium abbreviation signifies that of Washington (wru); Yale University (yu). Approximately 4500 h d specimen probably represents : sexual, diploid population, but the advanced flowering stage did not permit a positive determination. Townsendia Hook. Fl. Bor.-Am. 2: 16. 1834. Annual, or most frequently biennial or perennial, caulescent or acaulescent herbs with taproots (fibrous-rooted in one perennial ray-florets pistillate, fertile, uniseriate, ca. 10-100, mostly 20-40; ray- corollas yellow and frequently pink- or purple-tipped or tinged, gla- brous or lightly glandular; style-branches lanceolate, oblanceolate or oblong-linear, 0.8-2.5 mm. long, the upper portion (shorter or absent to glochidiate setae). Type-species ( originally the only species): Townsendia exscapa (Richards.) Porter. ARTIFICIAL KEY TO THE SPECIES AND VARIETIES A. Disk-pappus of very short bristles and squamellae less than 1.5 mm. long; plants fibrous-rooted and rhizomatous or stoloniferous; receptacle 70 ' JOHN H. BEAMAN 1. T. formosa. Disk-pe appus of bristles longer than 2 mm. (rarely with squamellae or short bristles also); plants ig eam not chisainitote or: stoloniferous; receptacle. merely. convex: or; flat... ., . cvsi.c2is . iad. canal. erin B. Disk-pappus of short pasillen or bristles and 2-4 see Lh to 8) longer, coarse bristles mia. B. Disk-pappus of 12 or more plurisetose bristles ................... C. Phyllaries bristly-stiff, apices attenuate-acuminate .............,..- Les, Catia Ein ke ee os is eee ee a 3. i anion. C. Phyllaries not bristly-stiff, apices acuminate, acute, or obtuse . Cee i Shee Bia Sue ON es eer ree tay ee feet eee D. D. Achenes at or near maturity with es deciduous pappus; plants villous-woolly with long trichomes ................ E. E. Phyllaries acuminate; involucres of Mecele ne gh more than 17 id a We i ae a T. condensata. E. nied msiye acute; involucres of all heads ~ less than 16 mm. ee a, ae See ee 16. 'T; er D. iabeaek with persistent pappus; plants villous-woolly | . F, Achenial hairs bifurcate or unevenly forked with one prong exceeding the other (sometimes pas simple), glochidiate by ike eR EES ee an eee G. Plants rosulate or ig aa mesh heads pedunculate or sessile among the tufts of leaves .................. . Ray-corollas Riboud or ie glandular on the abaxial — ace; achenes with delicate ili —_ near = Pr ere montana va H. raat ok emer chai on the dioeiial surface achenes pubescent from base to apex with straight Reay MEME DAE ol). iat eee I. Phyllaries in . (rarely 4) series, acute, pilose-strigose on the outer ace 14. T. scapigera I. Phyllaries in 4 (rarely 5) series, obtuse or acute, nearly glabrous or the outer lightly Lapeecmimcor ae near ae pe ees G. Plants with erect or spreading-suberect shapes heads terminatin leaf ON i eee J. Rays blue — whitish or pinkish); ere acumi- 2. T. Parryi. ae in 4- Oy Se a J. Rays white or plekighs phyllaries acute, in 3 (rarely OO elie opens cake ve cos oy qT. agi F. Achenial hairs glochidiate or achenes glabrous _....... K. Phyllaries linear to narrowly lanceolate ‘Chroader in some plants with the disk-corollas longer than 6 mm. and in some plants with very narrow, glabrous leaves), apices acuminate or acute, in 5-7 s L. L. Phyllaries with a ‘tuft of tangled cilia at rg linear, nate . Hoo keri. ke Phyllaries without a tuft of tangled a 4 ‘the apex, narrowly lanceolate, acute M. M. Disk-pappus more than 6.5 mm. long; leaf mid-veins conspicuous T. exsca mney. Pp M. Disk-pappus less than 6.5 mm. . long (if longer, the THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 71 ray- peppus less than half the length of the ee a pus); leaf mid-veins not conspicuous ............ N. veces — densely woo eee pubescent with long mensana pi acetal Senk Aicieaie. Sc acecnrg eta peemh hit ptotes. K. ye broadly ne to ovate or pe apices SCAR TE ON ep ahreel O. O. Leaves or ae or ‘both WOE ag ok cnet r. P. Ray-pappus nearly as ve i“ the disk-pappus ....... montana var. montana P. Ray-pappus less than ‘i as hee as the disk-pappus Q. Leaves conspicuously thickened; heads nearly sessile or short-pedunculate; phyllaries obovate, ovate, or broadly oblanceolate, mostly obtuse .... ......... ee aah Unite eek re ee a sai Leaves not cocaee “age thicke ned; pao gs ong- pedunculate; phyllaries lanceolate, acute .......... be ae a age ce 5. T. glabella. O. Leaves and achenes conspicuously pubescent ........ R. R. Disk-pappus shorter than the disk-corollas _....... S. S. Phyllaries in 2 series; plants of lesics eee nee po) en a 4. IAA PA GT ESE T. Plants nee phyllaries mostly in 4 series, the SCOR GON Se ce aes» endleri. R. Disk-pappus as long as or longer than the disk-corollas nials Ray-corollas eer pee? of the Texas ; Peskolidie exensis. Texas of Onrsnome 8 os ss. ee es oe W. Stems eraycokits with a dense ates (Ome? 17. ana. W. Stems merely lightly to moderately strigos SER MEG TRS strigo. U. —_ reget long-lived oan nials X. ves with dense tufts of “a0 simple pai ie oe Me NG et gn: antic: bay tacos eee . mensana var, mensana. X. Leaves oblanc eolate Oe Weer cs. z. Y. Leaves conspicuously thickened .............. De iie iver eh ery § ontana var. minima, Y. Leaves not conspicuously thickened ........ Z. Z. Phyllaries glabrous or the outer ones only htly pubescent; ray-corollas densely — GRE ick 5 aes . mensana var. cuous] “age trigose; ray-coolas Z. Phyllaries 8 siiacts to 5 lig tly glandular ..17. T. incana. 72 JOHN H. BEAMAN 1. Townsendia formosa Greene wnsendia formosa Greene, Leafl. Bot. Obs. & Crit. 1: 213. 1906. es Metcalfe 1434, Sawyer’s Peak, ca. 9000 ft. alt., pein Co., New Mexico, 30 Sept. 1904 (cas, lectotype; COLO, MoO, OC, isotypes ownsendia pinetorum Greene ex Nels. Coult. & Nels. Man. Cent. nécky 3 Mts. 508. 1909 (in synonymy, nom. nud. plsnkint ore 3 perennial, with a basal rhizomatous or stoloniferous mat, cing few t ny erect, monocephalous (or very rarely weakly pated. leafy stems; aerial stems striate, Bint strigose, becoming more heavily so near the head with both few- and many-celled, simple trichomes, inn coming lanceolate, apex acute, acuminate or obtuse, broadly scarious- margined and minutely ciliate, nearly glabrous or lightly pubescent along the midrib, 4-14 mm. long, 1.5-5.6 mm. wide; ray-florets ca. 20-35; ray- corollas white on fhe adaxial surface and bie urple or mauve abaxially except for the white margins, inconspicuous! glandular on the oe - m. surface, 14-26 mm. long, 2- e; disk-corollas yellow long, lightly gender. outside; achenes broadly oblanceolate to broadly obovate, compressed, 2-rib those e ray sometimes , gla- brous or minutely dandolar. 3-4.5 mm. , 2-2. oe mm. wide; ray-pappus of minute coroniform squamellae, less oe 0. long; disk-pappus of one to Heed (mostly 2) short, stiff, plurisetose ‘ites ssn less than Se mm. long and of short squamellae similar to those of the ray-pappus. = ae Re — oe Mountains of Baa a w Mexico and 73), White pe Ferris 10113 (ps, Ke Thompson aed Black River, White Mountains, Goodding 561 (coLo, GH, NY, °RM, US); Phelps Botanical Area, White Mountains, 9500 ft. alt., Phillips d> Phil- lips 3169 (cas). New Mexico. Catron Co.: 0.3 mi. east of Willow Creek a Ground, 18 mi. northeast of Mogollon, Beaman 994 (*cxH); Gilita Camp Ground, 20 mi. northeast of oa ge ca. 8000 ft. alt., Hitchcock et al. 4403 (cas, GH, NA, UC, UTC, Tu); Mogollon Moun tains, on or near the West Fork of the Gila saver b ca. 8500 ft. alt. “9 THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 73 Co.: Cloudcroft, Slater s.n., Aug. 1914 (us); Sacramento Mountains, Wooten s.n., 23 July 1899 (coLo, ps, oc, POM, °RM, UC, US Townsendia formosa is the most sharply defined of the species of Townsendia. Its rhizomatous or stoloniferous, fibrous-rooted habit is unique in the genus. None of the taprooted species occupies as mesic a habitat as does this fibrous-rooted species (the evolutionary trend of the genus is toward drier conditions ). Townsendia formosa is the only species with a conical receptacle. Monocephalous stems are found elsewhere only in T. Parryi. The peculiar small pappus also is not duplicated in any other species. All of the foregoing characteristics are interpreted to be primitive features in this genus. These are discussed in greater detail in the sections on generic relationships and trends of evo- lution. The primitive characters of T. formosa do not obscure the fact that it is related to T. eximia. Some degree of similarity can be seen in almost every aspect of the two. Their habits are similar; they have a similar indument; their phyllaries are (slightly ) similar; their achenes are very similar; and their pappus is similar. The edaphic requirements and geographical distribution of the two species fit a pattern which, when correlated with the morpho- logical similarities, indicates a close relationship. 2. Townsendia eximia A. Gray Townsendia eximia A. Gray, Mem. Am. Acad. 4 (PI. Fendl.): 70. 1849. Type: Fendler 353, sides of high mountains, Santa Fé Creek, Santa Fé Co., N. Mex., June 28, 1847 (cu, holotype; Mo, pH? [not numbered], yu, isotypes). Townsendia Vreelandii Rydb., Bull. Torr. Bot. Club 28: 22. 1901. Type: Vreeland 639, side of Veta Mountain, 8500 ft. alt., Huerfano Co., Colo. (xy, holotype; CAN, RM, isotypes). Caulescent, taprooted, biennial or short-lived perennial, erect or nearly so; stems few to numerous, branching from the base or above or both —— or villous, few-celled trichomes, up eaves oblanceolate to slightly spatulate, entire, mucronate, sometimes i Bide to lightly strigose especially along the 13 cm. | 1.1 cm. wide; upper cauline leaves similar, be smaller and more apiculate near the head; involucre 1.2-4 cm. wide; phyl- laries in 4-6 series, lanceolate to ovate-lanceolate with bristly-stiff, acuminate apices, glabrous or very lightly strigose, ciliate and scarious-margined, 5-15 mm. long, 1.5-4.2 mm. broad, those of the inner series longest and narrowest; ray-florets ca. 15-55; ray-corollas blue, 8-20 mm. long, 1.5-2.5 mm. wide; 74 JOHN H. BEAMAN disk-corollas yellow, sometimes red-purple-tinged, 3.5-5 mm. long; achenes obovate to broadly oblanceolate, truncate, compressed, those of the disk- florets 2-ribbed, those of the ray-flo orets 2-3-ribbed, sparsely pubescent with short, thick, duplex, glochidiate or bifurcate trichomes, —_ denser on the ray than the disk; achenes often papillose, 3.2-4.4 mm. long; pappus of the ray-florets o stiff, coroniform, basally connate pete fe than 0 length of the disk-corollas. 2n = 18. Reproduction sexual. Mountainous areas from south-central Colorado to Hon acne 1 New Mexico. Map 2. Plate XVIII, fig. 2. Representative specimens. Colorado. Conejos Co.: Cumbres, Fer- ril s.n., 2 Aug. 1902 (cs). Costilla Co.: east side of La Veta Pass, 9000 ft. alt., Hitchcock et al. 4160 (Na, wru). Huerfano Co.: south side of Mt. Mestas (formerly Veta Mountain), Beaman 745 .(*GH); 4 mi. northeast of Cucharas Pass, San Isabel National Forest, west of La Veta, 10,000 ft. alt., Rollins 1301 (Gu, Mo, Na, Ny); side of Veta Moun- tain, 8500 ft. alt. , Vree land 639 (CAN, Ny, RM). Las Animas Co.: Stonewall, 8300-9000 ft. alt., Beckwith 233 (*ny); near head of Ber- wind Canyon on road east of Delagua, 7500 ft. alt., Robbins 518 (uc). New Mexico. Bernalillo Co.: crest of Sandia Mts., 10,670 ft. alt., Bea- man 699 (*cuH); hillsides, Balsam Park, Sandia Mts., 8200 ft. alt., Ellis * Me NY, US); Albuquerque, Sandia Mts., Jones 4157 (cas, F, GH, SC, NY, OC, PH, POM, *RM, US, UTC). Colfa x Co.: Cimarron, Berg 3135 (cs); Colfax Co. (2): Cimarron Pass in Raton Mts., McKelvey 2419 (GH, POM); vicinity of Ute Park, 2200-2900 m. alt., Standley 14490 (*ny, us). Mora Co.: 10 mi. northwest of Mora, Beaman 719 (°on). Sandoval Co.: 11 mi. northeast of Jemez Pueblo on U. S. Forest Service road, Beaman 704 (°eH); aactcath Field Secgp area, i Springs, Nelson 11640 (ps, GH, *Mo, NY, RM, UC, UTC); Sandia Mts., Wooton s.n., 4 Aug. 1910 ( a fan Mi igu a Co: vicinity of Las Vegas, Anect 41 (cas, GH, us); hillside along Pecos River, Drouet & Richards 3316 (ps, GH, MICH, *Mo). Santa Fé Co.: % mi. west of Glorieta in Glorieta Pass, Beaman 717 (*cH); sides of high mountains, Santa Fé Creek, Fendler 353 (cu, Mo, PH? [not numbered], xu); Canoncito, Loch ft. alt., Heller & josie ee (DAO, DS, GH, ILL, *MO, MSC, ND, POM, RM, RSA, Tu); canyon wall 20 mi. wis 3 of Golden, ca. 7000 ft. sy " Hizehoook et al. 4229 (ps, NA, UC, WTU). Taos Co.: 18 mi. south of Taos, Beaman 723 (*cH); 9 mi. east of esta, Beaman 740, (*cHu); 5 mi. south of Questa, ca. 7000 ft. alt., Hitchcock et al. 4175 (cas, ps, GH, NA, UC, WTU). Plants of the southernmost geographical segment of this species have only slightly bristly-acuminate phyllaries that have a very sharp sofa thas etween the ciliate margin and her- baceous inner portion. The rest of the species is characterized by strongly Letaite ierainabe phyllaries with a transitional scarious THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 75 @T EXIMIA a1 FORMOSA T. GRANDIFLORA @ SEXUAL O APOMICTIC a T. TEXENSIS Map Z. Geographic yi of Townsendia formosa and T. eximia. 3. Geographic distribution of Townsendia texensis and sexual and apomictic T. ndiflora 76 JOHN H. BEAMAN area between the ciliate margin and herbaceous inner portion. The southern populations of T. eximia tend to approach their more southern neighbor, T. formosa, but there is no evidence which suggests hybridization between the two. Possibly this re- semblance is merely an indication of relatedness of the two species. Further evidence of their relationship was outlined in the discussion of T. formosa. Under most conditions T. eximia is a biennial, but in high- elevation populations, such as those near the crest of the Sandia Mountains and in the Raton Mountains, the perennial habit is well-developed. It appears that life-duration is not a strongly- fixed genetic character in the species. A parallel variation occurs in T. scapigera in eastern California where it occupies low- and high-elevation habitats. Likewise, several other members of the genus may be biennial or perennial, depending upon the habitat which they occupy. In the southern portion of its range T. eximia is usually an erect, sparsely-branched plant; to the north, especially in southern Colorado, it is suberect and more abundantly branched. In this feature T. eximia approaches T. grandiflora. Two collections, Standley 13289 and Standley 6357, from the vicinity of Raton, New Mexico, strongly suggest hybridization between these species. The plants in question have the widely branched habit of T. grandiflora. The disk-pappus, of several bristles, is inter- mediate between the two. The very bristly-acuminate phyllaries are suggestive ot T. grandiflora. Most of the other characters are typical of T. eximia, and these plants have been referred to that species. Field studies might reveal the relative importance of whatever gene interchange occurs between T. eximia and T. grandiflora. 3. Townsendia grandiflora Nutt. Townsendia grandiflora Nutt. Trans. Am. Phil. Soc. VII: 306. 1840. Type: Nuttall s.n., “On the Black Hills, (or eastern chain of the Rocky Mountains,) near the banks of the Platte,” 1834 (Bm, holotype, exam- ined by Dr. R. C. Rollins; cu!, pu!, isotypes). ubescence, ca. 1.5-30 cm. long; basal leaves spatulate, entire, obtuse, some- times slightly mucronate, lightly strigose above, nearly glabrous below, up to 5 cm. lon ide, mostly deciduous by flowering time; cauline leaves spatulate to oblanceolate, entire, acute or slightly acuminate or obtuse, mucronate, with a fairly conspicuous median nerve, lightly strigose especial- THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 77 ly along the nerve and margins, up to 9 cm. long (averaging ca. 4 cm.) and 1 cm. wide (averaging ca. 4 mm.); heads borne at the ends of the stems, often. subtended by one or a few leaves; involucres 15-30 mm. wide, 10-18 mm. high; hyllaries in 4-7 series, ovate-lanceolate or coe eer eto bristly-acuminate apices, ciliate and broadly scarious-margin median green or meine? eee. baa Pr: ae or lightly ths the outer surface, 4-1 ee series usually longest; eat ni i Soy te peice white above and usua e Mita: a median pink or purplish stripe below, 12-23 mm. long, 1-2.5 ; disk-corollas yellow, rarely pink-purplish tipped, 4.6 mm. long; shines oblancedlts! compressed, 2-ribbed, lightly to densely pubescent (denser on the disk than the ray) ‘with vather short, thick, duplex, glochidi- i 8 mm. wi short, coroniform squamellae or plurisetose bristles, not more than long; Mg: | ge of ca. 15-30 stiff, Seiigetnas: barbellate bristles, 3.8-6 mm. lon 18 in srcigh plants, a uae 2n = 27-36? in apomictic plants (the latter a en measurements). Reproduction mostly sexual, rarely apomi solic. yr noe estern South Dakota south to northeastern New Mexico. Map 3. Plate XVIII, fig. 3. Representative specimens. Colorado. Boulder Co.: southeast-facing slope near summit of Flagstaff Mtn., Beaman & Preece 509 hy: EI- dorado Springs, 5300 ft. alt., Clokey 2810 (CAN, CAS, DS, F, *NY, RM, uc, us). Custer Co.: southern slope, Hardscrabble Creek, shove Wet- more, 6000 ft. alt., Sn the 1 Sept. 1951 (*cs). Douglas Co.: hill- side 10 mi. north of Castle R Rock, 6000 ft. alt., ae 684 (*RM, ws). El Paso Co.: mountain side, Manitou, Shear 3687 (ny, *us). Fremont Co.: 11 mi. east of Canon City, Waterfall 11490 (*uc). Jefferson Co.: Mount aes yg sei of Denver, 6100 ft. alt., Constance & Rol- lins 1932 (NA ws). Gilpin Co.: Central City, Scovell s.n., 1869 (*micH). parties! Co.: hill p mi. west of Bellvue, Preece & Turner 2858 (*ws); Owl i Cae ca. 18 mi. northwest of Fort Collins, Weber 4873 (*coto). Park Co.: 7 mi. south of F airplay, 9000 ft. alt., Ripley & Barneby 10371 (#cas). Pueblo Co.: 2% mi. northwest of Rye, Beaman 744 Be ge Weld Co.: Grover, Johnston 228 (*RM). ico. Colfax Co.; Raton Mountains, Bell s.n., Aug. 887 (*pn); Gai Mountain, Raton, Cockerell & Cockerell s.n., 26 Aug. 1900 (us). Mora or Colfax Cu: prairie between Ocate Creek ‘and Rio Colorado, Fendler 533 (*mo); Mora River prairie, Fendler 157 (*mo). South Dakota. Shannon Co.: Cedar Draw, 8 mi. northeast of Rocky Ford, McIntosh s.n., (*pAo); sandbars, White River, Over 2399 (us). Washabaugh Co.: summit of Snake Butte, Over 2069 ( *us). ean, rue Co.: South Sibylee, Nelson 7373 (CM, COLO, “GH, ILL , NY, , RM, us). Goshen Co.: Fort Laramie, Nelson 8312 (*cH, MO, RM, us): "Lar- 78 JOHN H. BEAMAN amie Co.: Cheyenne, 1850 m. alt., Eggleston 12563 (*us). Niobrara Co.: Mexican Mines (Spanish Diggings), Nelson 578 (GH, ILL, MSC, ND, °PH, RM, ws). Platte Co.: hillsides 6 mi. north of Chugwater, Porter 3391 (Ds, GH, *MO, NY, RM, UC, US, WTU); slopes on the Oregon Trail near Warm Springs, 3 mi. west of Guernsey, 4400 ft. alt., Porter 4910 (COLO, DAO, *GH, MONT, OKL, PH, RM, RSA, UC, US, UTC, WTU). The bristly-acuminate phyllaries, erect or suberect, biennial habit, allopatric ranges, and same flowering times suggest a close relationship between T. grandiflora and T. eximia. ‘ownsendia grandiflora is adapted to the relatively arid foothills east of the Front Range in Colorado while T. eximia ocupies slightly more moist habitats at higher elevations in mountains to the south. A T. eximia stock may have migrated north along the foothills area and become isolated from the central concentration of T. eximia. During that isolation the two species as they are now known probably developed. Townsendia grandiflora is the more specialized of the two. An interesting parallel in the occurrence of apomixis in T. grandiflora and in another biennial, T. scapigera, has been men- tioned in the discussion of the latter species. Unlike apomictic T. scapigera, the apomicts of T. grandiflora appear to have re- tained the biennial nature of their sexual precursors (the apomicts of T. scapigera are short-lived perennials, although most plants of the species are biennial). It should be noted that the locality of Ripley & Barneby 10371 is the highest-elevation station now known for this species, and is the only station from which the species is known to be apomictic. 4. Townsendia texensis Larsen Townsendia texensis Larsen, Ann. Mo. Bot. Gard. 14: 15, pl. 3. 1927. Type: Eggert s.n., rocky bluffs of the Red River, Randall Co. (?), Texas, 13 Aug. 1900 (mo 121021, holotype, missing; cul, PoM!, iso- types). Caulescent, taprooted biennial; root-stem junction enlarged or elongated; stems branched mostly at the base, few-branched above, the central stem erect and the laterals spreading-suberect, striate, with a moderate to dense strigose-pilose pubescence, ca. 0.5-2.5 cm. long; basal leaves mostly decid- ong an mm. wide; heads borne at the ends of the stems, often subtended and surpassed by a few leaves; involucres 1.3-2 cm. wide, 0.7-1.1 cm. high; phyllaries in 4-6 series, ovate-lanceolate or lanceolate, acute or slightly acu- minate, ciliate and broadly scarious-margined, with a darker median streak, strigose on the outer surface or sometimes nearly glabrous, 3.5-10 mm. long, 0.5-2.4 mm. wide, the inner series usually longest; ray-florets ca. 25-40; THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 79 ray-corollas blue or very Ago white, sometimes pubescent on the —_ surface, 9.9-1.8 cm. long, 1.5-2.5 m m, wide; disk-corollas yellow, 3.5-5 m reece achenes oblanceolate, com siete ssed, 2-ribbed, lightly to stotleektely rie eee with rather short, thick, duplex, glochidiate hairs, 3-3.8 mm. long, EAL 7 wide; ray-pappus of Goconiborni concreted squamellae or sien plurisetose bristles, usually not more than 1 mm. long but rarely up to -pappus of ca. 16-25 plurisetose, a batetles, odin or since tha an the disk-corollas. 2n = 18. Reproduction sexual. Texas Panhandle and adjacent western Oklahoma. Map 3. Plate XVIII, fig. 4. Representative specimens. Oklahoma. Roger Mills Co.: side of one of the Antelope Hills, in northwestern yp of et Beaman 691 Sth Antelope Hills, Goodman 2614 ~~ WwTU Texas. Briscoe Co.: 5.6 mi. northwest o i Ouidaue: iets 17306 (Na). Carson Co.: ca. 4 mi. south of Borger, Beaman 696 (cH). Hansford Co.: near Paloduro Creek, 6 mi. southeast of Gruver, Beaman 694 (*cH); 5 mi. southeast of Gruver, Shinners 8227 (*nM, uc). Hartley Co.: 13.6 mi. west of Channing, Cory 16458 (Na). Hemphill Co.: near the Canadian River, 5 mi. south of Canadian, Beaman 693 (cH). Hutchinson Co.: 10 mi. southeast of Stinnett, Cory 16352 (Na). Mot- ley Co.: hillside 3 mi. west of Matador, Waterfall 7839 (cH, °Mo). Oldham Co.: 4 mi. north and 14% mi. east of Vega, 4000 ft. alt., Howard 159 (Na). Randall Co.: bluffs, 1 mi. north of Canyon, Beaman 698 (GH). In most of its morphological anapalie T. texensis is inter- mediate between T. eximia and T. grandiflora. > is closely re- lated to but distinct from both. Toinniandiat xensis and T. grandiflora probably are about equally derived species from a . eximia ancestral stock. The non-cuspidate (rarely minutely od ages phyllaries distinguish T. texensis from the other two s ue field observations of T. texensis in Roger Mills County, Oklahoma and several counties in the Texas Panhandle indicate that it is abundant in that area. It grows on Cretaceous limestones Fre Ati by stream erosion of the deep, black soils of the high plain 5. Townsendia glabella A. ue Townsendia glabella A. Gray, Proc. Am 86. 1880. Type: Newberry s.n., La Pagosa (Pagosa sone): ahisbel Co., Colo., July 29, 1859 (cn, holotype; Ny, Yu, isotypes). Townse ndia Bakeri Greene, Pittonia 4: 157. 1900. Type: Baker 727, Los Pinos (Bayfield), La Plata oe Colo. (Np, lectotype; F, GH, °MO, NY, POM, RM, UC, US, isotypes). Long-lived, decumbent-cespitose, taprooted perennial; caudex widely branched, becoming woody and subterranean; young stems at ends of the 80 JOHN H. BEAMAN caudex branches leafy, more densely so near the tips, up to 5.5 cm. long, often with one or a few peduncles in the axils of the upper leaves; leaves spatulate to oblanceolate, entire, the apex sometimes apiculate and mucro- er n 4 mm. wide, 7.8-12.3 mm. high, often broadly expanded at maturity; phyllaries in 3-6, mostly 4, series, broadly lanceolate or oblanceolate, acute or rarely slightly acuminate, glabrous or nearly so, scarious-margined, ciliate, 1 5m i i : Representative specimens. Colorado. Archuleta Co.: 9 mi. south- east of Pagosa Springs, Beaman 761 (*c1); hills, Pagosa Springs, 2160 m. alt., Bethel, Willey & Clokey 4340 (CAN, CAs, COLO, DS, F, MICH, °MO, MONTU, NY, PH, POM, RM, UC, US, UTC, Ws, WTU); 1.5 mi. south- east of Pagosa Springs, Turner 2892 (ws); mesa slopes ca. 12 mi. no of Arboles, 6400 ft. alt., Weber & Livingston 6238 (COLO, DAO, “RM, RSA, Ws, wTuU). La Plata Co.: hillside near Dix, 8500 ft. alt., Baker, Co.: Mesa Verde National Park, Nelson 10419 (GH, *MO, MONTU, NY, RM, UC). This species is closely related to T. eximia. Such features as the similar involucral bracts, similar vesture, and similar habit (of the high-elevation types of T. eximia and T. glabella) suggest the relationship. Townsendia glabella has the more specialized form of the two, and appears also to have much more specialized habitat requirements. It is abundant only on the Mancos shales of southwestern Colorado. Its endemism may be an expression of precise edaphic and climatic requirements that are satisfied only in that area. Even though its range is restricted, T. glabella is somewhat variable. Phyllary variation is especially noticeable. For example, in Baker 727 some plants have slightly bristly-acuminate phyl- laries; others have acute, and in one plant (Pom) broadly obtuse, THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 81 phyllaries. The plants of this collection also have shorter pe- duncles than do plants of the other collections. It is perhaps the minor divergence of Baker 727 from typical T. glabella which led Greene to describe T. Bakeri. One of the plants which Gray had at hand when he described T. glabella is a teratological form. Its ray-corollas, with long tubes and two conspicuous lateral lobes, are intermediate between ray- and disk-corollas. The ray-pappus is half as long as the disk- pappus. I have not considered the abnormal features of this par- ticular plant in the key. 6. Townsendia Rothrockii A. Gray ex Rothrock Townsendia Rothrockii A. Gray ex Rothrock, Wheeler Rept. U. S. Geograph. Surv. 6 (Bot.): 148. Pl. VII, A. 1878. Type: Rothrock 875, South Park, Park Co., Colorado, 13,500 ft. alt., July 1873 (¥F, holotype; GH, tNy, isotypes). Rosulate perennial with taproot; caudex short-branched, with: tufts of leaves at the ends, becoming woody, often subterranean; leaves spatulate- oblanceolate, entire, acute or obtuse, sometimes mucronulate, conspicuously thickened, shiny, glabrous or very lightly strigose, 10-35 mm. long and 2-7 mm. wide; heads sometimes nearly sessile, but usually on short pedun- cles up to 2.7 cm. long; peduncles naked or with a phyllary-like bract near the involucre, striate, villous; involucres 12-28 mm. wide, 8-12 mm. high; phyllaries in 4-6 series, elliptical, ovate, obovate, broadly lanceolate, or oblanceolate, acute, sometimes rounded at the tip, glabrous, scarious- margined, ciliate, red-tinged near the apex, 6-9.5 mm. long, 2-4 mm. wide; rets m. achenes broadly oblanceolate, compressed, 2-ri , those of the ray-florets rarely 3-ribbed; ray-achenes moderately pubescent with lon i and 2n = 36 in apomictic plants. Mountains of southwestern Colorado. Map 6. Plate XIX, fig. 2. Representative specimens. Colorado. Archuleta Co. (?): mts. east of Bayfield, 10,000 ft. alt., Bequaert s.n., July 1929 (*cH). Gunnison Co.: crest of ridge between Avery Mtn. and Virginia Mtn., near Gothic, Beaman & Barclay 798 (+cH); Taylor Pass, 12,400 ft. alt., Langenheim 442 (tcoxo); foot of Matchless Mtn., 11,000 ft. alt., Langenheim 1310 (tcoto). Hinsdale Co.: Crystal Lake near Lake City, 11,000 ft. alt., Pease s.n., 28 June 1878 (fru). Mesa Co.: 6 mi. above Taylor Ranch, 82 JOHN H. BEAMAN COLORADO e T. GLABELLA COLORADO T. ROTHROCKII © APOMICTIC T. MONTANA VAR. MONTANA @ SEXUAL © APOMICTIC fe) re) fe) o re) Ceo oO DO ely [ MONTANA e VAR. MINIMA ® @& SEXUAL & APOMICTic Sa a s 4-7. 4. Geographic distribution of Noa sg Fendleri. 5. Geographic dis- ibeibe of T. glabella. 6. Geographic distribution of sexual and apomi Rothrockii. po. ic fi romaphae distribution of sexual ed apomictic “Tr montana var. montana and T. montana var. minima. THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 83 sier Ridge, ocak ft. alt., Weber & Rollins 7166 (cas, coo, DAO, a +RM, RSA, ws, wru); South Park, Wolf 417 (Ny, us). San Juan ” Bapneezs Peak, 12,000 ft. alt., Purpus s.n., 27 July 1893 (F, Soa MICH, MSC). Townsendia Rothrockii appears specialized for existence in a very cold climate. Its morphological specialization obscures its relationship with other species, but possibly its nearest, although not too close, relationship may be with T. glabella. In Park and Gunnison Counties, Colorado, two apomictic populations of the species have very unusual habitats. At eleva- tions over 12,000 ft. they grow in red sandstone fragments under snowbanks, flowering as the snow melts away. Such peculiar conditions perhaps are not required by the species but may be optimum, thus helping to explain the narrow endemism of T. Rothrockii. 7. Townsendia montana M. E. Jones Rosulate-pulvinate, taprooted perennial; caudex becoming much- ss often partly su esis the old seaglonag often persisting oe veral years, sometimes becoming much elongate in seth bearing the Palgyat scars of old leaves and ee oO nie pa a roots; leaves spatulate, sometimes thickish, entire, obtuse, sometimes mucro- nate, rarely emarginate, nearly glabrous to ate onan strigose, up to 40 mm. lon d ide; heads gaan fey sessile); peduncles naked or nearly so, were at junction with the head, with pilose-strigose pu- bescence, i Ae o ca. 6. . lon ng; involucres obconical at base, 8-15 mm. wide, 6- ec ies phyllaries in 3-6, mostly 4, series, oblong, obovate, 12 m oblanceolate “ Tan nce ve obtuse and siacied at the apex or rarely _ 3-ribbed, ig rd or sparse ely pubesce nt with rather pola duplex bifurcate hairs, some times papillose, 37-52 mm. sod 1.0-1.6 mm. ; pa ch of the ray- and disk-florets similar, of ca. cee oe be acid bristles sewed KEY TO THE VARIETIES Heads at — slightly nr Pe rays blue or white; iad _ _ not conspicuously thickened 052... 6.0 ee Hoeisls sessile; rays pink; a involute, conspicuously ilies 7b. var. minima. 84 JOHN H. BEAMAN 7a. Townsendia montana M. E. Jones var. montana Townsendia montana M. E. Jones, Zoe 4: 262. 1893. Type: Jones s.n., above the Flagstaff Mine, Alta, 9500 ft. alt., Salt Lake Co., Utah, 7 August 1879 (rom 40755, holotype). Townsendia alpigena Piper, Bull. Torr. Bot. Club 27: 394. 1900. Type: Cusick 2294, subalpine ridges of the Wallowa Mts., 7000 ft. alt., Wallowa Co., Oregon, July 31, 1899 (tws, lectotype; F, GH, MO, MSC, UC, US, isotypes). Townsendia dejecta A. Nels. Bot. Gaz. 37: 267. 1904. Type: Goodding 1238, Dyer Mine, Uinta Mts., ca. 3000 m. alt., Uintah Co., Utah, July 3, 1902 (*?xM, holotype; Mo, us, isotypes). 2n = 18 in sexual plants and 2n = 36 in apomictic plants. Reproduction predominantly apomictic. Mountains of western Montana to eastern. Ore- gon and south to central Utah. Map 7. Plate XIX, fig. 3. Fremont Co.: east side of Mt. Jefferson, southwest of Henry’s Lake, 9500 ft. alt., Cronquist 1924 (ips, }Mo, UTC, Wru); mountains north- east of Henry Lake, 8700 ft. alt., Payson & Payson 1986 (cas, tcM, GH, MO, Ny, RM, uc). Lemhi Co.: Liberty Range, west of Gilmore, Christ & Ward 14886 (tNny p.p., with 5 plants of T. leptotes). Teton Co.: Teton Peaks, Davis s.n., July 1930 (fips). Montana. Beaverhead Co.: ridge connecting Sheep and Black Lion Mts., Pioneer Range, ca. 9300 ft. alt., Hitchcock & Muhlick 13003 (ps, +mMo, RM, RSA, UC, WS, wtu). Gallatin Co.: without definite locality, 8500 ft. alt., Tweedy s.n., Aug. 1886 (tv). Madison Co.: % mi. north of Kock Peak, Taylor Mts., Hitchcock & Muhlick 15176 (+Ny, wru); head of Cottonwood Creek, Tobacco Root Range, 9000 ft. alt., Blankinship s.n., 10 Aug. 1902 (¢mMonT, UTC, wru). Oregon. Wallowa Co.: subalpine ridges of THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 85 east of East Fork of Bear River, Uinta Mts., 11,000 ft. alt., Goodman & Payson 452 (oi, tRM); divide between East Fork of Bear River and Black’s Fork, 10,500 ft. alt., Goodman & Hitchcock 1517 (¥; cu P. ps, with 4 plants of T. leptotes; ty p.p., with 3 plants of T. leptot es). Uintah Co.: Dyer Mine, Uinta Mts., Goodding 1238 (mo, *? RM, vs); rocky crest of east side ‘of Whiterocks chen between Red Pine and Paradise Creeks, 10,000 ft. alt., Graham 10065 (*cm, *NA). Wyo- Big Horn or Sheridan Co.: summit of Big Horn Mts., Blankin- (tuc). Lincoln Co.: ridge near Cottonw Lake, east of Smoot, 10,400 ft. alt., Payson & Armstrong 3706 fox tmo, Msc, RM); Dead Man Peak, 9500 ft. alt., Williams 1301 ( CAS, Mo, RM, UTC). ee Pass Mts., east of Victor, Idaho, 9200 ft. alt., Payson ¢& Payson 2078 (CAS, CM, GH, MO, *Ny, RM); Teton Pass, 9000 ft. alt., Ripley & Bar- neby 8898 (®cas); Sheep Mtn., alpine, 10,200 ft. alt., Tweedy 532 baa uy vicinity of Teton Pass, 9500 ft. alt., Williams 786 (*cas, NY, RM, UTC). Yellowstone National Park: Mammoth Hot seria Oleson 116 (tRM). 7b. Townsendia montana M. E. Jones var. minima (Eastwood) Beaman, comb. no Townsendia minima Eastwood, Leafl. West. Bot. 1: 206. 1936. Type: Eastwood & Howell 727, Bryce Canyon, Garfield Co., Utah, June 19, 1933 aki holotype). 2n=18? xual plants (inferred from pollen measurements) and 2n = 27 in ison ee eh Southwestern Utah. Map 7. Plate XIX, fig. 4. Representative specimens. Utah. Garfield Co.: Red Canyon near Bryce, 7000 ft. alt., Cottam 9691 ee Red Canyon, Eastwood 785 (cas); Red pean 10 mi. from Brvce Park entrance, Hitchcock 2962 o a); AN, “UTC, wTu); liffs, 4 mi. east of Pine Lake, Table Cliff Plateau, Powell National Forest, 9000 ft. alt., Maguire 19120 (GH, NY, UTC). Kane Co.: near summit of Pink Cliffs, headwaters, left fork Virgin River, 15 mi. northwest of Orderville, Maguire 18807 (utc); 5 mi. west of Long Valley Junction on Highways 14 and 89, Preece & Turner 2462 (ten, tws). Townsendia montana appears to be a continuation of the T. eximia-T. glabella evolutionary line. Its relationship with this 86 JOHN H. BEAMAN group is indicated by the glabrate leaves, lightly pubescent or glabrous achenes, and broad, glabrate phyllaries. Townsendia montana may the evolutionary result of isolation of this (T. eximia-T. glabella) stock in the mountains of Utah away from the central core of the genus in the Rocky Mountains. Variety montana is mostly a high-elevation type, occurring at about timberline. Variety minima is found at somewhat lower elevations, but probably does not occur below the ponderosa pine belt. It has been collected at elevations as high as 9000 ft. Although var. montana is known from very few collections that represent sexually reproducing populations, each of these is slightly different morphologically from the others. At the south- ern extremity of its range, the San Pete County population is a fairly close morphological approach to var. minima. The most problematical portion of T. montana is in the Uinta Mountains. A collection (Goodding 1238) described by Nelson as T. dejecta is somewhat distinct from the other collections of T. montana var. montana. Variety montana usually has peduncu- late heads, while the heads of this collection are sessile. Typical var. montana has glabrate, spatulate leaves, and this material has moderately pubescent, oblanceolate leaves. In these unusual fea- tures Goodding 1238 seems to approach T. leptotes. Another collection (Graham 10065) from the same area represents the only known population of T. montana with a short ray-pappus. It seems significant that T. leptotes in this area also has a short ray- pappus. Only apomictic T. leptotes is now known in the Uinta Mountains, but sexual material could be there now or recently may have been there. Sexual T. leptotes of the long ray-pappus form is nearby in western Moffatt County, Colorado ( Wolf dy Dever 5193). The type of variation in the collections Goodding 1238 and Graham 10065 seems most readily explainable on the basis that hybridization between T. montana var. montana and T. leptotes has occurred. It should be noted also in this connec- tion that the collection Goodman & Hitchcock 1517, from the Uinta Mountains, is a mixture of T. montana var. montana and T. leptotes. These specimens of the two species are very difficult to distinguish from each other and appear to be from apomictic populations of hybrid derivation. The sexual population of T. montana var. montana in Wyo- ming at Teton Pass, Teton County, is of interest because of its con- siderable intra-population variation. Four collections, Beaman & Preece 505, Payson & Payson 2078, Ripley ¢ Barneby 8898, and ‘Williams 786, each consisting of a fairly large number of plants, THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 87 have been made at or near the same spot. Plant-to-plant variation in these collections is evident in at least three features. Some plants have nearly glabrous ray-corollas; in others the ray-corol- las are moderately or densely glandular. Some plants have nearly glabrous phyllaries; in others the phyllaries, especially the outer ones, are moderately or densely strigose. The apex of the phyl- laries likewise is variable. In some plants the apices are broadly obtuse; in others they are acute, and in still others they are inter- mediate. Variety montana is ordinarily characterized by glabrous ray-corollas, and nearly glabrous or only lightly pubescent, obtuse phyllaries. The unusual characters in this population probably belong to T. florifer. The latter species has densely glandular ray-corollas and moderately strigose, acute phyllaries. It seems possible that pollen occasionally has been carried up to the T. montana population by insects which have visited T. florifer plants in the lower areas of adjacent Idaho. Apomixis in var. montana is concentrated in the high-eleva- tion, northernmost populations. This pattern is no different from that of the other species in the genus with abundant apomixis. Variety minima is so restricted that the geographic and altitudinal pattern of the range of its apomictic forms is not at present demonstrable. It should be noted that in one area (Red Canyon) apomicts and sexual plants of var. minima occur together. 8. Townsendia mensana M. E. Jones Rosulate-pulvinate perennial with well-developed taproot; caudex de- veloping few to numerous short, often subterranean, branches; leaves spatu- late, oblanceolate or linear, entire, acute or~slightly. acuminate, — i g and 1- ate, mostly acute, infrequently obtuse, with scarious or ciliate margins, glabrous to lightly pappus of the ray-florets variously eastern Utah to southeastern Nevada. KEY TO THE VARIETIES Leaves narrowly oblanceolate to linear, abaxial surface of leaf-bases densely woolly with long, white, multicellular trichomes; phyllaries lanceolate; 88 JOHN H. BEAMAN heads sessile, almost hidden in the tufts of leaves; ay of the Uinta Basin 0 sheer Ripe Fi Gand Ue. se homen ey ey tee var. mensana. Ehecien heads usually short-pedunculate, or if appearing a not itiritege va —— tufts of leaves; plants of western sig south- astern Nev 8b. var. Jonesii. 8a. Townsendia mensana M. E. Jones var. mensana Townsendia mensana M. E. Jones, Contr. West. Bot. 13: 15. 1910. Type: Jones s.n., benches of the Uintas, Theodore (now Duchesne), 7,500 ft. alt., Duchesne Co., Utah, 14 M ci 1908 (rom, holotype). Uinta Basin, Utah. Map 8. Plate XX, fig. Representative specimens. Utah. Du . Co.: 26 mi. southwest of Myton, 7000 ft. alt., Parols 12701 (*cas); 15 mi. west of Du- chesne, Beaman 865 (cH); “bench” of the Uintas, 11 mi. north of Duchesne, Beaman 868 (cH); 3 mi. west of Duchesne, 5500 ft. alt., Ripley & Barneby 4677 (cas). Uintah Co.: between Hill Creek and Green River, ca. 20 mi. south of Ouray, Bartholomew & Bartholomew s.n., June 1955 (cH). 8b. Townsendia mensana M. E. Jones var. Jonesii Beaman, var. nov. Type: Jones s.n., Mammoth, 7000 ft. alt., Juab Co., Utah, 10 May 1910 (rom 39891, ‘holotype). Leaves oblanceolate or spatulate, acuminate, the bases strigose on the abaxial surface; heads ee! ge eee a Western Utah and south- eastern Nevada. Ma Plate XX Foliis tibet vel spathulats, am basi_ strigosis, capitulis Boise breviter pedunculat Representative specimens. Nevada. Clark Co.: Deer Creek, Charleston Mountain, 8250 ft. alt., Alexander 774 (*uc); ridge south of Deer Creek, 2670 m. alt., Clokey 7772 (uc); Lee Canyon, 2650 m. alt., Clokey 7773 (uc); Hidden Forest Canyon, Sheep Range, 9500 ft. alt., Munz 16836 (pom, uc). Lincoln Co.: Mt. Irish, 6000 ft. alt., Jaeger s.n., 19 June 1938 (pom). White Pine Co.: Schellbourne, Jones s.n., 13 July 1891 (pom). Utah. Juab Co.: McIntyre’s Ranch, Jones s.n., 18 May 1891 (pom); Silver City, Jones s.n., 27 March 1896 (pom): Tintic Junction, 5500 ft. alt., Jones s.n., 9 May 1910 (pom). Millard Co.: Leamington, 5000 ft. alt., Jones s.n., 8 May 1911 (pom, atypical, hybrid with T. florifer ?, cf. discussion); San Pete Co.: 2 mi. southeast of Ephraim, Beaman 820 (cH); Gunnison, 5000 ft. alt., Jones s.n., 18 April 1911 (rom). Sevier Co.: ridge between Cottonwood and Willow Creeks, 8000 ft. alt., Robinette 105 L. R. (*Na). Tooele Co.: Deep Creek, Jones s.n., 6 June 1891 (pom). Tooele or Utah Co. Mercur, 5500 ft. alt., Jones s.n., 6 June 1896 (°Mo, Ny, POM, US, miss), Wayne Co.: 10 mi. scuthenat of Teasdale, Beaman 822 (*cxH). THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 89 This nearly unknown species has recently figured in a nomen- clatorial confusion. It was placed in synonomy under T. sericea (=T. Hookeri) by Larsen (1927). Its resemblance to that species is superficial. Cronquist first recognized that T. sericea is invalid as it included the type of the earlier Aster P exscapus [=Town- sendia exscapa (Richards.) Porter]. With T. mensana residing in synonomy under T. sericea in Larsen’s treatment, it appeared that the epithet “mensana” should be taken up for the latter species. Both Cronquist (1955, pp. 326-327) and I (Beaman, 1954, p. 172) fell into a trap by not having seen the type of T. mensana. Critical study of the type and other material indi- cates that T. mensana must be recognized as a species in its own right Townsendia mensana var. mensana is endemic to the Uinta Basin of Utah. From my field observations, it appears that the variety has its best development on the pinon- and juniper- covered “benches” of the Uintas to the north and west of Du- chesne. It occurs also at slightly lower elevations, with a host of other specialized little xerophytes, on the white shales of the Green River formation. Townsendia mensana var. Jonesii occurs over a wider area and is represented by many more specimens than is var. mensana. Marcus E. Jones, however, has been the principal collector of var. Jonesii, and the variety is named in his honor. The diversity of T. mensana var. Jonesii is considerable. The the phyllaries are usually slightly obtuse. The variation between plants of the two areas is continuous, however, and it seems best to consider the material as a single taxon. The variation trend of the Utah material of var. Jonesii is in the direction of T. florifer. For the most part this variation is subtle, and the present material is not adequate to permit a good understanding of its cause. Only two collections, Jones s.n., 8 May 1911 (Millard Co., Utah) and Jones s.n., 6 June 1891 (Tooele Co., Utah), show strongly intermediate characters between T. florifer and T. mensana var. Jonesii. Unfortunately both collections are scanty, one consisting of one small plant, the other of two. In the Millard County collection (2 plants), one plant seems nearer 1It should be noted that T. Hookeri has been collected by Marcus Jones near the type-locality of T. mensana var. mensana. JOHN H. BEAMAN a T. MENS VAR MENSANA @ T MENSANA VAR JONESII oe 8 e cm %e e e e ® T LEPTOTES oO SEXUAL O © APomictic ° © UNDETERMINED 8 ae .@) ore) - Qs ea Maps 8-9. 8. mari date PP Pp wits T. mensana var. Jonesii. leptotes. ion of Townsendia mensana yar. mensan graphic distribution of sexual and apomictic T. THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 91 T. florifer, the other nearer T. mensana var. Jonesii. One plant has mostly glochidiate achenial hairs; the other has mostly bifurcate hairs. The plant with the bifurcate hairs appears to be annual or biennial; the other could be perennial. The phyllaries of both plants are in three series and similar to those of T. florifer. In the Tooele County specimen the achenial hairs are bifurcate, the phyllaries are in four series and have a strigose pubescence intermediate between typical T. florifer and typical T. mensana var. Jonesii. In general aspect this plant seems closer to typical T. mensana var. Jonesii than do the two Millard County plants. This evidence of hybridization between T. mensana var. Jonesii and T. florifer is meager. However, in view of the general situation in Townsendia as regards interspecific hybridization, it would be surprising if crossing did not occur between the two. They occur sympatrically over a fairly large area in the sagebrush lands of west-central Utah. They probably occupy the same or at least very similar habitats. Marcus Jones (1893, p. 262) states that T. mensana var. Jonesii (det. by him as T. scapigera) is rare. Very likely it is, since he is the only collector of it in the area of probable hybridization. But even though it is rare, an intensive field investigation might be expected to demonstrate considerable gene interchange between T. florifer and T. mensana var. Jonesii. The closest relationship of T. mensana appears to be with T. montana. The ancestral stock in the Wasatch region may have differentiated into low-elevation T. mensana and high-elevation T. montana. Some relationship between the two species is shown by the similar habit, the similar involucral bracts (especially be- tween T. montana and the southern Nevada populations of T. mensana var. Jonesii which have broad, obtuse phyllaries ), and the similar light pubescence of the phyllaries. The differences between the two varieties of T. mensana are minor, although constant. The two taxa have been placed at the varietal level to indicate their very close morphological relation- ship. Variety mensana has probably been isolated in the Uinta Basin for sufficient time to become somewhat differentiated from a more primitive var. Jonesii stock. 9. Townsendia leptotes (A. Gray) Osterhout 92 JOHN H. BEAMAN Colorado, July-August 1864 (*cH, holotype; F, MO, NY, PH, UC, YU, isotypes; us, probable isotype but dated 1862). Rosulate-pulvinate perennial with well-developed taproot; caudex of ew to numerous, short, subterranean branches terminated by tufts of leaves; leaves linear to oblanceolate or narrowly spatulate, entire, mucro- nate, thickish, usually involute, glabrous to rather heavily strigose-sericeous, up to 60 mm. long and 3.5 mm. wide; heads sessile or pedunculate, the peduncles not more than 2.8 cm. long; involucres campanulate, 0.8-2.3 cm. wide, 9.5-1.5 cm. high; phyllaries in 4-7 (rarely 3) series, lanceolate to li brous or very lightly strigose on the outer surface, 3.5-12 mm. long, 0.5-2 m. wide; ray-florets ca. 15-40; ray-corollas whitish, cream, pink, or blue glabrous or nearly so, 8-14 mm. long, 1-2.5 mm. wide; disk-corollas yellow, similar, of plurisetose barbellate bristles, 3-8 ong, or us of th ray-florets variously shortened or absent. 2n = 18 in sexual plants, and - in apomictic plants (inferred from pollen measurements ) central Nevada and east-central California. Map 9. Plate XX, figs. 3 and 4. Representative specimens. California. Mono Co.: R. 34 E, T. 5 S. Sect. 2, White Mountains U. S. G. S. Map, 11,500 ft. alt., Duran 1661 (tuc); north slopes of Sheep Mountain, White Mountains, 12,000 ft. alt., Maguire & Holmgren 26109A (uc, turc). Colorado. Archu- leta Co.: Pagosa Springs, Bethel s.n., 23 May 1917 (*cs); hillsides, southeast of Pagosa Springs, 7200 ft. alt., Ripley & Barneby 7589 (*cas); Pagosa Springs, Smith s.n., 12 May 1894 (pH). Grand Co.: 4.5 mi. west of Kremmling, Beaman & Preece 513 (*ws); Kremmling, Osterhout 5221 (*oxKL, POM, RM); 1 mi. east of Grandby, 7800 ft. alt., Ripley & Barneby 10586 (*cas). Gunnison Co.: crest of ridge be- tween Avery Mountain and Virginia Mountain, near Gothic, Beaman & Barclay 799 (+c); North Station Ridge near Gothic, 12,000 ft. alt., Langenheim 725 (coo, cs); Fairview Peak, 13,200 ft. alt., Langenheim 1292 (tcoLo); Matchless Mountain, 12,000 ft. alt., Langenheim 1309 (coo). Lake Co.: near Leadville, Schedin & Schedin 478 (*nm). La Plata Co.: 5 mi. northwest of Hesperus, Pennell 21464 (*ny, pH). Moffat Co.: saddle just west of fire look- out, summit of Roundtop Mountain, Dinosaur Nationa Monument, 2800 m. alt., Wolf & Dever 5193 (*coLo). Montezuma Co.: Mancos, Eastwood s.n., June 1891 (*pom); Cortez, Nelson 10437 (GH, MO, NY, *RM, UC). Park Co.: South Park, Wolf & Rothrock 418 (+GH, NY, tus). Saguache Co.: Marshall Pass., Ferrill s.n., 13 June 1903 (*cs); Cochetopa Park, just south of summit of Cochetopa Pass, ca. 96 ft. alt., Weber 9421 (coro, *cH). Idaho. Blaine Co.: Mt. Hyndman, THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 93 Sawtooth Range, 8000 ft. alt., Thompson 13626 (mo p.p., with 3 plants of T. montana). Clark Co.: 3 mi. from head of Medicine Lodge Creek, Davis 3148 (tips; tna p-p-, with 1 plant of T. Hookeri). Custer Co.: Mt. Borah, Chilly, Christ & Ward 10424 (tny); south- east of Double Springs Summit, 8 mi. northeast of Dickey, 8500 ft. alt., Cronquist 3197 (+cH, ws, Mo, all .p., with plants of T. montana) ; pass in mts. 7 mi. north of Dickey, 8500 ft. alt., Hitchcock et al. 3790 (twru); mouth of Rock Creek, north slope, Lost River Mountains, Hitchcock 15725 (twrv). Lemhi Co.: Liberty Mountain, Lemhi Range, west of Gilmore, Christ & Ward 14886 (tNny p.p., with 3 plants of T. montana); near head of Spring Canyon ca. 8 mi. southeast of Gilmore, Lemhi Mountains, Hitchcock & Muhlick 9317 (Ny, twrv). Montana. Madison Co.: without definite locality, 9500 ft. alt., Dome, 10,500 ft. alt., Maguire & Holmgren 25944 (cas, tNy, uC, utc). New Mexico. Rio Arriba Co.: hills south of village, Tierra = Amarilla, 2300 m. alt., Eggleston 6444 (¥, *cH, *Mo, Ny, us). U San Pete Co.: Spring Hollow Drainage above Great Basin Experi- : : 1 River, Uinta Mountains, 11,400 ft. alt., Goodman & Payson 450 (oxi, tro); divide between East Fork of Bear River and Black’s Fork, Uinta Mountains, 10,500 ft. alt., Goodman & Hitchcock 1517 (Ds, F; GH p.p., with 1 plant of T. montana; micu, +MO, MONTU, p-p., with 2 plants of T. montana; pH, RM, UC). Wyoming. Fremont Co. (?): gravelly hills, Wind River Mts., Hayden s.n., 15 May 1860 (tmo). Yellowstone National Park: Saddle Mountain, 10,000 ft. alt., Tweedy 697 ({cH, Ny). Townsendia leptotes exhibits the greatest interpopulation variation of any species in the genus. No two collections from sexual populations are entirely similar. Variation occurs even between populations which are only a short distance apart. The species is most diversified in western Colorado. Sexual populations of T. leptotes appear to be confusing and difficult to determine, mostly as a result of hybridization with T. exscapa. The populations of southwestern Colorado and ad- jacent Rio Arriba County, New Mexico exhibit a polymorphy which seems attributable only to the influence of T. exscapa. In southwestern Colorado T. leptotes becomes a large plant with a much-branched caudex; it has relatively large, sessile heads with involucres averaging, ca. 2.0-2.5 cm. wide; the disk-corollas are unusually long, averaging 5.5-6 mm., and the disk-pappus is 94 JOHN H. BEAMAN correspondingly long. These atypical features of T. leptotes are characteristic of T. exscapa. An unusual character, not from T. exscapa, in the southwestern Colorado material of T. leptotes is the short ray-pappus. Town- sendia glabella, which occurs in that area also, has a short ray- pappus. However, no other features definitely suggest gene ex- change between these two species. This character could have had an independent development in the southwestern Colorado popu- lations of T. leptotes. As the ray-pappus is a variable character in several other Townsendia species, such an explanation might apply here. The polymorphy between populations of the apomicts of T. leptotes indicates that apomixis must have occurred independently in several different sexual populations. For this reason the apo- micts are interesting from a phytogeographic point of view. Those of the Uinta Mountains (e.g., Goodman & Hitchcock 1517) are very similar to the plants of southwestern Colorado. They have the short ray-pappus and the habit almost exactly duplicates that of the purer (relatively undiluted with T. exscapa) sexual plants of southwestern Colorado (e.g., Ripley & Barneby 7589). The apomicts of Park County, Colorado (Wolf & Rothrock 418) are very similar to sexual T. leptotes from Marshall Pass, Saguache County, Colorado (Ferril s.n., 13 June 1902). The apomicts of San Pete County, Utah, Nye County, Nevada, and Mono County, California (e.g., Holmgren & Shaw 7640, Maguire & Holmgren 25944 and Maguire & Holmgren 26109A, respectively ) are more difficult than the preceding examples to relate to a known sexual type but they are morphologically close to some of the alpine apomicts from Gunnison County, Colorado (e.g., Beaman & Bar- clay 799). The Idaho apomicts are closest morphologically to the apomicts of the Uinta Mountains of Utah, but in the Idaho ma- terial (in contrast to the Uinta material with the short ray- pappus) the ray-pappus is about as long as that of the disk. In Wyoming T. leptotes is known only from apomicts of two old collections (Tweedy 697 and Hayden s.n., 15 May 1860). These are perplexing because of their resemblance to T. spathu- lata. Their narrow phyllaries and glabrous achenes are characters of T. leptotes. Their pubescence is nearly as dense as that of T. spathulata, and their pappus is deciduous like that of T. spa- thulata. The oblanceolate leaves are about intermediate between the two species. These plants could be apomictic hybrid deriva- tives of T. leptotes and T. spathulata. Further field studies of the THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 95 populations in western Wyoming are needed to help clarify the relationship that seems to exist between these two species. Putative hybridization between T. montana var. montana and Cronquist 3197, and Thompson 13626). 10. Townsendia Hookeri Beaman, sp. nov. Type: Clokey 4338, dry hills, Mt. Vernon Canon, 1730 m. alt., Jefferson Co., Colorado, 13 April 1920 (*coto 12061, holotype; CAN, CAS, DS, F, GH, MICH, MO, MONTU, NA, PH, POM, *RM, UC, US, UTC, WS, WTu, isotypes). Densely rosulate-pulvinate perennial with well-developed taproot; cau- dex of few to numerous short, subterranean branches, appearing smoo g head; inear to narrowly oblanceolate, entire, acute, mucronate, involute, thickish, densely strigose-sericeous, up to 45 mm. long and 2 mm. wide rarely u m passed by the tufts of leaves; involucres obconical-campanulate, 8-18 mm wide, 9-15 mm. high; phyllaries in 5-7 (rarely 3-4) series, linear or nearly so, acute or acuminate, mostly terminated by a tuft of tangled cilia, ciliate i e a purplish upward, 5-13 mm. long, 0.6-1.8 mm. wide; ray-florets ca. 15-30; ray-corollas white on the adaxial surface, the abaxial surface cream or pink- ish, glabrous or nearly so, 8.5-14 mm. long, 1-2.5 mm. wide; isk-corollas > i?) S 3 = o ya 72) i9°) a ie) ~ = e. o a ie) ® = or & y = we & ~ Ue a. e ys ) s aa oF io} © ie 24 » or ® ee io = pollen measurements). Northeastern British Columbia south to southern Colorado and central Utah. Map 10. Plate XVII and Plate XX, figs. 5 and 6. erba rmis; numerosis; folia linearia ad suboblanceolata, integerrima, strigoso-sericea, a 4.5 cm. longa et 2 mm. lata. Capituli in summo caudicis sessiles; involucrum 8-18 mm. latum, 9-15 mm. altum; involucri squamis plerumque 5-7 seriatis, lineari-subulatis, acutis vel acuminatis, 5-13 mm. longis, 0.6-1.8 mm. latis. 96 - JOHN H. BEAMAN Corollae radiorum 8.5-14 mm. longae, 1-2.5 mm. latae; corollae disci 4.0-6.5 mm. longae. Achaenia oblanceolata, compressa, pubescentia cum _ pilis glochidiatis, 3.5-4.7 mm. longa, 1.0-1.4 mm. lata; radiorum pappo ca. 1 mm. longo vel usque 5.5 mm. longo; disci pappo plurisetoso, 5-8.2 mm. longo. Representative specimens. CANADA. Alberta. Rocky Mountains, Drummond 573 (}tcan); Sweet Grass Hills, Macoun 10897 (can, ND); Waterton Lakes National Park, hilltop near north park entrance, 4200 ft. alt., Breitung 17067 (Ny); Banff National Park, upper drain- age of N. Saskatchewan River, Valley of Mistaya River, between Saskatchewan Crossing and Waterfowl Lakes, 4500 ft. alt., Porsild & Breitung 14758 (cAN, cH); Medicine Hat, Spreadborough 5027 _(cAN, GH, MO, Ny, US). British Columbia. Fort McLeod, Cowdry 7709 (can). Saskatchewan. Cypress Hills, summit of hills east of Boyd’s Ranch, Breitung 5357 (pao); Maple Creek, Campbell 129 (tpao p.p., with 1 plant of T. exscapa). uNrTeD sTaTEs. Colorado. Boulder Co.: near Boulder, Ramaley 654 (coLo, *RM); % mi. east of Hi-Way Mine, 3 mi. south of LaFayette, Weber 3713 (COL, p.p., with 1 plant of T. exscapa). Clear Creek Co.: Georgetown, Wolf & Rothrock 416 (¥, }cH; us p.p., with 1 plant of T. leptotes). Custer Co.: high mountain valley, Cusack s.n., 1888 (F); Denver Goa.: Denver, 5000 ft. alt., Bethel s.n., 1 May 1894 (*Ny; us p.p., with 1 plant of T. exscapa). Fremont Co.: Canon City, Brandegee 41 (*? PH p-p., with 2 plants of T. exscapa). Gilpin Co.: Central City, Scovell s.n., 1869 (*micH). Gunnison Co.: Sapinero, Wheeler 598 (coLo: tr p.p., with 2 plants of T. exscapa). Larimer Co.: F oothills (Fort Collins), Crandall 305 (*vus p.p., with 3 plants of T. exscapa) ; Horse- tooth Gulch, Crandall 3131 (cs, *Ny, RM, YU). Mineral Co.: the “Mesa”, Creede, Rio Grande National Forest, 8900 ft. alt., Murdock Lewis & Clark Co.: ‘Helena, Kelsey s.n., April 1888 (ps, trom, UC). Liberty Co.: Cottonwood Coulee, ca. 20 mi. southeast of Chester, Marks s.n., 4 May 1946 (wis). Madison Co.: Silver Star, Fitch s.n., 20 April 1919 (+mMonr). Sweetgrass Co.: northeast corner of county, Anderson & Scharff s.n., 21 April 1952 ({mMonT). Wibaux THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 97 Co.: Wibaux, Blankinship 1503 (tmont). Yellowstone Co.: 7 mi. south of Custer Station, Northern Pacific Railway, Big Horn River, Blankinship 147 (mo, MONT, us). South Dakota. Custer Co.: Pringle, Black Hills National Forest, 4900 ft. alt., Murdock 3510 Forwood 93 (tus). Utah. Carbon Co.: Scofield, Jones s-n., 24 June 1904 (pom). Duchesne Co.: Theodore (now Duchesne), benches of the Uintas, 8000 ft. alt., Jones s.n., 13 May 1908 (trom). Wy- oming. Albany Co.: ca. 12 mi. southeast of Laramie on U. S. Rt. 30, ca. 8800 ft. alt., Beaman & Preece 508 (*ws); Laramie, Nelson 7055 (MO, Ny, POM, *RM, Us). Crook Co.: hillsides near Hulett, 4000 ft. alt., Ownbey 526 (coLo, GH, IDs, }MO, MONTU, NY, RM, UC west end of Bate’s Hole, Payson & Payson 4738 (nm). Park Co.: Shoshone Mountains, Hapeman s.n., May 1907 (+mo). Teton Co.: 11 mi. north of Jackson, ca. 6200 ft. alt., Beaman & Preece 506 (ws); Black Tail Butte, Moose, Craighead s.n., 25 May 1947 (tips). Uinta Co.: near Lyman, 6500 ft. alt., Rollins 1619 (cu, tNy). Yellowstone National Park: Hills near Mammoth Hot Springs, 6200 ft. alt., Bur- glehaus s.n., May 1893 (tus, wrv). Larsen (1927) recognized that Hooker had included two spe- cific elements in his species, Townsendia sericea, upon which he based the genus. She did not recognize, however, that T. sericea was invalid because it included the type of the earlier Aster ? exscapus (cf. Beaman, 1954, Cronquist, 1955, and the discussion of T. mensana in the present paper). Thus, the element which was not Aster P exscapus [Townsendia exscapa ( Richards.) Por- ter] was not described by her. The “new” species, T. Hookeri, is named in honor of the founder of the genus Townsendia. To the layman, T. Hookeri is perhaps as well known as any species of Townsendia. In Colorado it is abundant at the base of the Front Range and is one of the first flowers of the year; hence, its name Easter Daisy (applied also to T. exscapa). The botanists’ confusion of T. Hookeri and T. exscapa is a result of the great similarity of the two, which may have resulted from extensive hybridization. In addition, over most of their ranges from Wyoming and Nebraska northward, both species reproduce only apomictically. Some specimens are easy to distinguish, but others are more intermediate and may be distinguished only with difficulty. 98 JOHN H. BEAMAN T. PARRY! @ SEXUAL O APOMICTIC Maps 10-11. 10. Geographic distribution of sexual and apomictic _Townsendia i. Hookeri. 11, Geographic distribution of sexual and apomictic T. Parry In southeastern Wyoming and the Front Range of Colorado sexual T. Hookeri and sexual T. exscapa occur together. I have not seen the two growing side by side, but several mixed collec- tions such as Bethel s.n., 1 May 1894, Brandegee 41, Crandall 305, Rydberg & Vreeland 5408, and Weber 3713, indicate that they must grow very close together. Their flowering periods at least partly coincide (both species of the plants of the mixed collections are in flower ), and thus conditions to permit hybridization appear favorable. The nature of variation of plants of both species strongly suggests that hybridization has occurred. Field studies should be made to permit a better understanding of the frequency of hybridization and introgression, which, from herbarium ma- terial, appear to be important in these two species. As an aid to field population studies, the important characteristics of the pure T. Hookeri and T. exscapa populations are compared in the table below. Although the dates of collection of the mixed collections of T. Hookeri and T. exscapa from the Front Range of Colorado indi- cate that the two do flower at the same time, they may be at least partly isolated by differences in flowering period. Dr. Marion Ownbey (in conversation) has indicated that he found T. Hookeri (Ownbey 526, apomictic) in flower on April 27 and noted that THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 99 T. exscapa T. Hookeri branches very short, or in old plants becoming longer and thicker, appear- ing smooth to the unaided eye with the matted pubescence of old leaf bases. Leaves linear or very narrowly oblanceo- late, involute, sericeous-canescent. s relatively small, sessile, the involu- 1.5 cm. wide. Puywvaries linear-subulate, with a tuft of tangled cilia at the apex CaupEx branches _ short, longer in old plants, but not or becoming conspicuously thickened, the | an in T. Hookeri, th the ithout the perficially smooth texture of T. Hookeri Leaves oblanceolate to narrowly spatulate, not involute, strigose. Heaps relatively large, frequently pedunculate, the involucres ca. 2.2 cm PHyYLLaRies mostly narrowly without a tuft of cilia at the ape short- . wide. lanceolate, X. ACHENES moderately to densely pubescent. ACHENES moderately to densely pubescent. DisK-COROLLAS Ca. mm. | Disk-coroLLas ca. 9-10 mm DisK-PAPPUSs conspicuously the disk-corollas. 5 ong. . long. Disk-pappus barely exceeding the length longer than of the disk-corollas. other rather similar plants at the same locality were still in bud. He returned to this locality on May 6 and collected the later- flowering plants which proved to be T. exscapa (Ownbey 534, apomictic ). The behavior of these apomicts may indicate that the owering periods of their sexual precursors are slightly different. ownsendia Hookeri is closely related to T. leptotes. This relationship is evident from a number of characters common to both species. Both have linear or at least very narrowly lance- olate leaves; in both, the phyllaries are very narrow. The habit of both is similar, and the two have a flowering period closely correlated with the beginning of spring. Townsendia Hookeri may have differentiated from a stock close to the present T. lep- totes, as a result of isolation of populations on the two sides of the Front Range in Colorado. The center of diversity of T. exscapa is to the south of Colorado. It probably was well-differentiated from T. Hookeri or the proto- e of T. Hookeri before the two jointly occupied the area east of the Front Range in Colorado. It is possible, although entirely speculative, that the adaptation of T. exscapa to the Great Plains area has been coupled, through hybridization and apomixis, with the specialization of T. Hookeri to a colder environment. A com- bination of these two characteristics in the apomicts of both spe- cies may have permitted their migration far into the plains of Canada. It must be pointed out, of course, that some of the apo- micts of the northern regions seem to have most of the character- istics of the pure species. This is particularly true of the apomicts of T. Hookeri in Montana. In that area they are more easily dis- 100 JOHN H. BEAMAN tinguished from T. exscapa than are some sexual T. Hookeri plants in Colorado. Of the two species, T. exscapa appears to be the one - more markedly affected by hybridization. Townsendia Hookeri is especially difficult to culture in the _ greenhouse as it merely develops buds which abort before they reach the meiotic stage. A cold treatment of at least two or three months is required to induce preformed buds to continue develop- ment and flower. When this species was first collected by Drum- mond he noted this oe In so far as I know, it has never since been mentioned. Drummond stated (1829, p. 206): “It has a very singular habit, little like that of the genus Aster [he considered it to be Richardson’s Aster ? exscapus]; the flower buds are formed in Autumn, and bear an exact similarity to those of Globularia vulgaris.” Drummond was disappointed in the small, inconspicuous rays. Many apomicts, and his was one, have rays which do not fully expand. The geographic pattern of sexual-apomictic distribution of T. Hookeri is a nearly diagrammatic one. The sexual forms oc- cupy a limited area at the southern portion of the range. The apomicts, on the other hand, seem to be excluded from the im- mediate area where the sexual plants occur (except for high- elevation apomicts ) but are widely distributed to the north. 11. Townsendia exscapa (Richards.) Porter Townsendia exscapa (Richards.) Porter, Mem. Torr. Bot. Club 5: 321. 1894. Aster ? exscapus Richards. Frankl. Journ. Bot. App. 748. 1823. Townsendia sericea Hook. Fl. Bor.-Am. 2: 16. 1834. Type: Richardson s.n., Carleton House, Saskatchewan (+k, holotype). Townsendia sericea Hook. 8 papposa A. Gray, Mem. Am. Acad. 4 (Pl. Fend.): 69. 1849. Type: Fendler 349, ‘arid hillsides, less frequent in grassy places, Santa Fé, Santa Fé Co., N. Mex., April-May 1847 (*cu, holotype; Ny, PH, UC, Us, isotypes). Townsendia Wilcoxiana Wood, Bull. Torr. Bot. Club 6: 163. 1877. scone: exscapa Wilcoxiana (Wood) Nels. in Coult. & Nels. Man. Bot. Rocky Mts. 510. 1909. Type: Wilcox s.n., Camp sappy, Indian ir (Oklahoma, Woodward Co.), March 1877 (us, type). nsendia intermedia Rydb. in Britt. Man. 944. 1901. Type: Rich 718. Prairie, Trego Co., Kansas, 1896 (*Nny, holotype; GH, MO, RM, US, isotypes). osulate, taprooted perennial; caudex developing several short (or a ppt ed), often subterranean, branches, becoming woody; leaves o anceolate, sometimes only narrowly so, atte, mucronate, eh ‘ose a trigose-sericeous, up to 8 cm. long, averaging ca. 3-3.5 cm 0.6 cm. wide, rarely with the widest less than 2 mm. wide; fiat a THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 101 or short-pedunculate (in the southern sporting of the range), the peduncles rarely more than 3 cm. long; involucres 1.3-3.7 cm. wide, 1-2.2 cm. high; phyllaries in 4-7 series, Tanceolate or linear, acute or Laren obtuse, args long, 1 _ wi e; ray-florets ca. 20-40; tay Lund 3 (utc); 20 mi. west of Goldhill, Utah, 4900 ft. alt., Maguire & Becraft 2828 (utc); Silver Zone Pass, 5200 ft. alt., Ripley & Barneby 4613 (cas). White Pine Co.: 8 mi. west of Baker, 5800-6100 ft. alt., Ripley & Barneby 3569 (cas). Oregon. Baker Co.: Huntington, Sweetser (?) s.n., (uc). Crook Co.: just west of Prine ville, Twp. 14 S., R. 15 E., S. 36, Ramee Be sei riage 6960 (*cH, ws); Cline Falls, Nelson 81 5 (cM M, uC). Deschutes Co.: 15 mi. north of Bend, ca. 3500 rs a t Hécheook ¢> Martin 4906 (Ds, NA, *NY, POM, UC, UTC, WS, WTU). Gilliam Co.: west slope along John Day River, 1 mi. above the alae of Rock Creek, Twp. 1 N., R. 19 E., Cronquist 6209 (ws). Harney Co.: western slopes north of Squaw Butte, Squaw Butte Experiment Station, Maguire & Holmgren 26510 (ny, uc, utc). Jefferson Co.: flats at = Palisades, 13 mi southwest of Madras, Twp. 12 S., R. 12 E.. 5. 2. 1300 ft. alt, Crom quist 6946 (CAs, DS, ILL, RSA, UC, UTC, ws). Lake Co.: hills west of Silver Creek, Cusick 2616 vg MO, MSC, ND, “NY, POM, RM, UC, US, : ; .: Frisco on Highway 21, ca. 30 mi. west of Milford, 5000 ft. alt., Cottam 4636 (uc); Milford, Coodding 1046 (cH, RM). Box Elder = 20 mi. north of Lucien, Maguire & Maguire 21518 (CAN, GH, *MO, NY, PH, UTC, ws, wtu). Garfield Co.: 10 mi. south of Coyote, 6500 ft. alt., Jones 114 JOHN H. BEAMAN 5315b (us). Juab Co.: 2 mi. east of Troutcreek, 5000 ft. alt., Maguire & Becraft 2829 (cH, utc). Millard Co.: 37 mi. west of Delta, 5200 ft. alt., Maguire & Becraft 2830 (pom, *RM, uc, UTC); Warm Point Ridge, west end Pine Valley, 5 mi. west Hdqts. Desert Range Experi- ment Station, Maguire 20880 (cu, *Ny, utc). Piute Co.: Marysvale, 6000 ft. alt., Jones 5323 (¥, Mo, Msc, POM, us). Sevier Co.: Joseph City, 5500 ft. alt., Jones 6379 (*mo, pom, us). Tooele Co.: Gold Hill, Jones s.n., 6 June 1917 (pom); Dugway, Jones s.n., 30 May 1891 (cs, GH, POM); Stansbury Island, Watson 520 (Gu, us, yu). Tooele or Utah Co.: Mercur, 5500 ft. alt., Jones s.n., 6 June 1896 (pom). Utah Co.: Goshen, Garrett 3955 (ur). Wayne Co.: Rabbit Valley, 6800 ft. alt., Ward 523 (cu, us). Washington. Adams Co.: Ritzville, 488 m. alt., Cantelow s.n., (wru). Franklin Co.: hillside, Kahlotus, Cotton 1013 us, ws). Grant Co.: slopes north of Soap Lake in Grand Coulee, Thompson 11498 (cas, DAO, Ds, *"GH, MO, MONT, MONTU, NA, NY, PH, POM, RSA, US, WS, WTU). Kittitas Co.: near Vantage, Thompson 1 3707 (¥, GH, MO, Ny, US). Klickitat Co.: Columbia River opposite Willows, Howell s.n., 18 April 1880 (cH). Lincoln Co.: slopes at Almira, Thompson 11666 (cas, ps, MO, Ny, POM, wTu). Walla Walla Co.: hills, Wallula, Cotton 1034 (us, ws). Yakima Co.: ravine near High- way 97, on Logy Creek, Hitchcock & Martin 3386 (ps, NA, POM, RM, RSA, UC, WS, WTU). Townsendia florifer is a sharply defined species and may be determined with ease. It has been taxonomically troublesome, however, when attempts have been made to distinguish T. Wat- soni A. Gray from T. florifer. The type of T. Watsoni (Watson 520) is from near the center of the geographic range of T. florifer and is intermediate in the gamut of morphological variation of T. florifer. It cannot, therefore, be maintained at the specific or varietal level. Townsendia florifer is closely related to T. Parryi. The two species occasionally have been confused, apparently because of hybridization between them. This is considered in the discussion of T. Parryi. The close phylogenetic relationship of T. florifer and T. scapigera is considered in the discussion of the latter species. Townsendia florifer is common and has been abundantly col- lected along the Snake River Plains of Idaho and in central Ore- gon and western Utah. It flowers mostly in May and June. Be- tween Hinkley, Utah and the Utah-Nevada boundary, I made arbitrary stops every 10 miles for 70 miles, during the summer of 1954. A plant or two of T. florifer was found at three of these stops. At two non-arbitrary stops plants were also found. It THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 115 appears that the species is ubiquitous in that area. It probably is abundant only in moist seasons. @T. FLORIFER. T INCANA Maps 13-14. 13. Geographic distribution of Townsendia florifer and sexual = apomictic T. scapigera. 14. Geographic distribution of sexual and apomictic T. incan 14. Townsendia scapigera D. C. Eat. in S. Wats. wnsendia scapigera D. C. Eat. in S$. Wats. Bot. Expl. Fortieth Parallel, 145, pl. 17. 1871. Type: Watson 518, Trinity Mts., i ft. alt., Pershing Co., Nevada, May 1868 (xu, holotype; us, iso Ff og scapigera var. caulescens D. C. Eat. in S. Wats. ‘ibid. p- 145. Type: Watson 519, Monitor Mas 5000 ft. alt., Nye Co., Nevada (xu, holotype; GH, Ny, US, isotype Cespitose, taprooted biennial or short-lived sities much-branched and densely leafy at the base, sometimes with a well-developed woody caudex with Seabank leaf scars; leaves spatulate, 3 "rors sometimes mucronate, infrequently emarginate, strigose, up to long (averaging long; achenes oblanceolate, compressed, 2-ri the ray-florets rarely 3-ribbed, pubescent with long, thin, duplex, “oso hairs, papillose, 116 JOHN H. BEAMAN 3.8-5.6 mm. long, 1.1-2.0 mm. wide; pappus of ca. 20-30 plurisetose a the bristles awl-shaped, obcompressed, densely barbellate or ciliate; ray- pappus 3.1-6.2 mm. long; disk- -pappus slightly longer than the ray- pipiet. 5.0-7.2 mm. long, slightly exceeding the length of the disk-corollas. Repro- duction mostly sexual, rarely apomictic. Nevada and eastern California. Map 13. Plate XXI, fig. 4. Representative specimens. California. Inyo Co.: Teufel Canyon, Inyo Mts., 25 mi. northwest of Darwin, 4600 ft. alt., Jaeger s.n., 28 May 1938 (pom); Teufel Canyon, southern Inyo Mts., 5200 ft. alt., Jaeger s.n., 27 May 1939 (ps); Seep Hole Spring, Inyo Mts., Kerr s.n., May 1940 (*cas); ae Flat SS a ae south of Reed Flat, White Mts., 10,000 ft. alt., Roos & Roos 5120 (ps, *Rsa, uC); Westgard Pass, 7000 ft. alt., Peet de Kellogg 2492 (uc, ws); Westgard ass, summit of ri ridge a above small canyon, 7300 ft. alt., Alexander & Kellogg 2492a (ps, uc); Mazourka Canyon, 7800 ft. alt., Alexander & Kellogg 2993 (mo, *rM, uc, us, UTC); Cerro Gordo Peak, crest of divide above Cerro Gordo Mines, 8000 ft. alt., Alexander & Kellogg 3020 (uc). Modoc Co.: Ft. Bidwell, Austin iss April 1878 (cH); Buffalo Ravine, near Surprise Valley, Lemmon 29 (° Mo, yu); Warner's Ranch, Lemmon s.n., May 1879 (uc). Mono Co.: western side of Sweetwater Canyon, 10 500 ft. alt., Alexander & Kellogg 3959 (tuc); top of divide east of Deep role, ll, 200 ft. alt., Alexander ¢& Kellogg 4053 (GH, MO, RM, tUC, U' c, wTu); divide, 1 mi. north of Mt. Pat- terson, 11,000 ft. alt., ficomer & Kellogg 4556 (mo, uc); east of Swamp Meadows, 9500 ft. alt., Alexander & Kellogg 4561 1 (cH, *Mo, uc, utc); 15 mi. north of Mono Lake P. O., on Conway Grade (to Bridgeport), ca. 7500 ft. alt., Blake 11831 (ps). Plumas Co.: with- out definite locality, Austin s.n., 1880 (Ny). Nevada. Elko Co.: north of Deeth, upper Humboldt Valley, 5350 ft. alt., Hall 10365 {*uc); Cobre, 6000 ft. alt., Jones s.n., 16 June 1906 ( POM); mesas and hills, near Cave Creek Post Office, Ruby Valley, Mason 4694 (uc). Esmeralda Co.: between Pinchot Creek and Pinon Hill on road to B. & B. Mine, 6700 ft. alt., Duran 2758 (*uc); Magruder Mtn. south end of the Silver Peak Range, 8200 ft. alt., Maguire & Holmgren 25640 (cu, ny, uc, *utc). Eureka Co.: Palisade, Jones s.n., 14 June 1882 (pom). Humboldt Co.: 5 mi. north of Farnham Hot Springs, Quinn River Valley, 5000 ft. alt., Train TC4 (us). Lander Co.: Victory Highway, 20 mi. east of Battle Mountain, East- wood & Howell 175 (cas). Mineral Co.: top of ridge, north, over- looking Hawthorne, 9500 ft. alt., Alexander & Kellogg 4440 (*uc); Mount Grant, Wassuk Range, 10 000 ft. alt., Archer 7130 (uc); road west to Laphan Canyon rim along north basin, edge of Laphan Meadows, west slope of Mt. Grant, Wassuk Range, 9400 ft. alt., Train 4178 (rsa, uc). Nye Co.: 4 mi. south of Millett, 5500 ft. alt., Lindsdale & Linsdale 690 (cas, uTC); west slope, Toquima Range, mouth of Mariposa Canyon on Manhattan-Round Mountain road, 5700 ft. alt., Train 2738 (Ny, uc, utc, wru). Pershing Co.: Rochester, THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA LIZ mtn. slopes, 4000 ft. alt., Train R30 (us); Trinity Mts., 5500 ft. alt., Watson 518 (Ny); Pah-Ute Mts., 5500 ft. alt., Watson 518 (cH). White Pine Co.: Warm Springs, King s.n., May 1918 (cas). Townsendia scapigera is morphologically closest to the central Oregon populations of T. florifer. In that area the stem leaves of T. florifer are few and the base of the plant is very leafy. Only a slight change from that condition would be required to produce T. scapigera. In northeastern Nevada, on the other hand, where a varietal distinction between them. More collecting in northern Nevada and southern Oregon will be required to elucidate com- pletely the status of the two species. Townsendia scapigera attains a considerable degree of diver- sity in the mountains of Inyo and Mono Counties in California. This diversity has already been noted by Heiser (1948). He sug- gested that apomixis might be involved. It is involved, but prob- ably is not important in increasing the diversity of these popula- tions. The large plants in the mountains of Mono County probably have become differentiated from the smaller plants in the mountains of Inyo County largely as a result of geographic semi-isolation. In Inyo County especially, one might expect the development of different ecotypes. In that county, plants have been collected as low as 4600 ft. elev. (Jaeger s.n., 28 May 1938) and as high as 10,000 ft. (Roos ¢ Roos 5120). This is the greatest elevational range in a species of Townsendia. If ecotypes have developed, they are not very apparent in herbarium material, however. The only obvious feature is that the low-elevation plants are smaller and shorter-lived than those from higher elevations. Apomixis in this primarily biennial species parallels the cor- respondingly small amount in another biennial, T. grandiflora. Apomixis in these species is rare, and occurs only in plants of high elevations. In the Sweetwater Mountains of California, plants from the summit or very near the summit of the mountains are apomicts. Morphologically very similar sexual plants (Alexander é> Kellogg 4561) occur on a protected knoll about 1000 ft. below the summit of the mountains. The apomicts must have some adaptational qualities, not present in the sexual plants, which better suit them to the tops of these mountains. 118 JOHN H. BEAMAN 15. Townsendia condensata D. C. Eat. in Parry Townsendia condensata D. C. Eat. in Parry, Am. Nat. 8: [106 nom. nud.] 213. 1874. Type: Putnam s.n., Washakies Needles, Owl Creek Range, Hot Springs Co., Wyoming, 1873 (yu, holotype; ten, isotype). Townsendia anomala Heiser, Madrofio 9: 240. 1948. Type: von Schrenk s.n., Holm Lodge, about 40 mi. west of Cody, Park Co., Wyoming, 26 Aug. 1922 (mo, holotype). Ros eet RCO taprooted, short-lived perennial; caudex becoming r few and very hoe or peracid nearly absent in alpine age or sometimes long, 2-4 mm. no et in ae cas cao sessile si em opener in the tufts of leaves, in plants of lower elevations the first-formed head usually essile, the later, lateral heads terminating the stems in a corymbose manner; mm. wi 8 involucres 1 ide, 8-18 mm. high; phyllaries in 35 series, lanceo- ometimes linear, acuminate or less often acute, scarious- and long- sdlinteshareined. ‘abendadis pubescent with long several-celled trichomes spicuo d-walls, 4-14 mm. long, 0.5-2.8 mm. wide; ray-florets ca mp: ri , those of the ray florets sometimes 3-ribbed, moderately pabescent with bifurcate or — duplex hairs, cat aang 3.2- 4.5 mm. weg i 0. 8-1. 3 plants. Reproduction predominantly apomictic, rarely sexual. Mostly at high elevations, western Montana and Wyoming to east-central California. Map 15. Plate XXIII, figs. 2 and 3 Representative specimens. California. Co.: western side of Sweetwater Canyon, 10,500 ft. alt., ne & Kellogg 3953 (tuc); top of divide east of Deep Creek, 11,200 ft. alt., Alexander t Fakes 4061 (tuc); R. 34 E., T. 5 S., Sec. 2, White Mt. U.S.G.S. Map, 11,500 ft. alt., Duran 1662 (tuc); north slopes of Sheep Mt., White Mts., Maguire & Holmgren 26109 (tutc). Idaho. Custer Co.: saddle near head of Rock Creek, ca. 1 mi. northwest of Mt. Borah, Beaman 893 (cH); Lost River Mts., 1 mi. west of Borah, at summit of Saddle between Rock and Mahogany Creeks, 10,800 ft. alt., Hitchcock & Muhlick 10977 (cas, ps, tNy, RM, UC, UTC wre). Montana. Glacier Co.: Aberkuny [Appekunny Bovine Glacier ft. Witt 1884 ade Wyoming. Fremont Co. (?): Wind River, Parry 143 (cH). Hot Springs Co.: Washakies Needles, Owl Creek Range, THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 119 Putnam s.n., 1873 (cH, yu). Park Co.: south-facing slope near Holm Lodge on Crossed-Sabre Ranch, 8 mi. east of the east entrance of Yellowstone National Park, 7000 ft. alt., Beaman & Preece 503 (cu, *“ws); Holm Lodge, about 40 mi. west of Cody, von Schrenk s.n., 26 Aug. 1922 (mo). This species has long been problematical to the few botanists who have seen it. Some of its peculiar characters are shared with the related and almost equally rare species, T. spathulata. This similarity led Larsen (1927) to consider T. condensata to be the same as T. spathulata. She stated (p. 23) that the head size is of no value for specific demarcation between the two species. This notion was occasioned by her confusion of material of a thir species, T. Parryi, with these two. She reduced T. condensata and T. Parryi var. alpina (= T. Parryi) to T. spathulata apparently because the head size of the plants of T. Parryi was intermediate between the other two species. Townsendia condensata and T. spathulata may be distinguished by head size and by several other characters as well. These are given in tabular form below. Townsendia condensata Short-lived perennial. Wi r without conspicuous caudex and often with conspicuous lateral stems. Phyllaries acuminate. Primary heads large, involucres (of the primary heads, at least) s more than 1.7 cm. wide. Townsendia spathulata Long-lived perennial. With caudex densely matted with the old and new leaves. Phyllaries acute or only slightly acuminate. All heads small, the involucres less than 1.6 cm. wide Townsendia condensata is known almost entirely from alpine, apomictic material. Fortunately, attention was called to a sexual population of the species in Park County, Wyoming when Heiser (1948) described T. anomala. The material on which Heiser’s species is based is a little different from the alpine apomicts of T. condensata. However, the differences are only in the degree of pubescence and length of stems, and both of these characters vary from population to population in the apomicts. If T. spathu- lata had not been confused with T. condensata at the time of Heiser’s investigation, he probably would have been able to asso- ciate his material with the latter species. Sexual material of T. condensata probably will be found also in Fremont County, Wyoming. Parry 143, from the Wind River 120 JOHN H. BEAMAN T. CONDENSATA @& SEXUAL a ~~ & apomictic a & . T. SPATHULATA @ = sexuar © APOMICTIC _ @ UNDETERMINED T. MEXICANA gh I6 f Maps 15-16. 15. Geographic distribution of sexual and apomictic Townsendia pues — sexual and apomictic T. spathulata. 16. Geographic distribution of T. mexic THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 121 Valley, is a single dried-up plant, but it is much like the presently- known sexual population in Park County. The Wind River Valley has a number of habitats similar to that in Park County where the sexual population is now known to occur. Further collections in western Wyoming will be required to solve the problems of loca- tion and diversity of sexual plants of this peculiar little species. A feature not apparent in sexual material of T. condensata grown in its native habitat becomes apparent in greenhouse cul- tures. Under these conditions it does not remain a pulvinate little mass, but instead develops a single, erect, leafy stem which later branches at the base. This habit reveals a close, but otherwise not too obvious, relationship with T. Parryi. Once the similarity in habit has been noted, other indications of this relationship be- come more evident. Additional features the two have in common include the long-acuminate phyllaries, large heads, and bifurcate achenial hairs. 16. Townsendia spathulata Nutt. Townsendia spathulata Nutt. Trans. Am. Phil. Soc. 7: 305. 1840. Type: Nuttall s.n., “On the Black Hills, (an alpine chain toward the sources of the Platte.)”, 1834. (BM, holotype, examined by Dr. R. C. Rollins; cu!, px!, isotypes). Densely rosulate-pulvinate, taprooted perennial; caudex becoming several-branched, obscur: the dense tufts of new leaves and the per- sistent dead leaves; leaves spatulate, entire, acute or obtuse, mucronulate (usually hidden by the pubescence ), densely pubescent with a_villous- or sericeous pubescence of ge multicellular trichomes with m mm over 10 mm. lo in 3-4 series, mostly lanceolate, sometimes narrowly ovate or elliptical, acute i i cilia multicellular trichomes, 3-9 mm. long, 0.8-2.5 mm. wide; ray-florets ca. 12- 30; ray-corollas pinkish, brownish orange, or lavender, often glandular on ed ong, 1.5-3 mm. wide; disk- corollas yellow and usually pinkish or reddish tinged, 3.8-5.3 mm. long; achenes_ oblanc compressed, 2-ri , moderately pubescent with long, 1-1.5 mm. wide; pappus of the ray- a a. 14-30 plurisetose, barbellate (sometimes with recurved barbs ) bristles, de- ciduous in a ring around the apex of the achene, 4.5-6.2 mm. long. Repro- duction sexual and apomictic. Fremont, Natrona, and Sweetwater Counties, Wyoming. Map 15. Plate XXIII, fig. 4. Representative specimens. Wyoming. Fremont Co.: Beaver Hill, 32.5 mi. southeast of Lander, Beaman 880 (cH); 1 mi. south of Pa- 122 JOHN H. BEAMAN cific Creek, Beaman 883 (cH); open ridges, 1 mi. south of Pacific Springs, Porter 4544 (+cH, tRM, RSA, wru); South Pass, Wind River Mts., 7650 ft. alt., Ripley & Barneby 7965 (* and tcas). Fremont Co. (?): Wind River, Parry 142 (¥, GH, Mo, Ny, *PH, yu). Natrona Co.: 4 mi. west of Alcova, Beaman 878 (cH); on the Satanka Forma- tion ca. 4 mi. west of Alcova, 5400 ft. alt., Porter 4428 (*pao, GH, °RM, RSA, WTU). Sweetwater Co.: Bush Ranch (near Steamboat Mtn.), Nelson 7094 (tro). Although T. spathulata and T. condensata are amply distinct species, they are closely related. The confused history of the two gives one indication of that relationship. The following similari- ties between the two give further indication: spatulate leaves, densely pubescent with long, simple, multicellular trichomes with conspicuous end-walls; bifurcate or entire achienial hairs (al- though some populations of T. spathulata have glochidiate hairs ) ; very deciduous pappus; conspicuously papillose achenes. Even though sexual T. condensata seems to be rarer than T. spathulata, the former species is the more generalized type. Townsendia spathulata appears to have become specialized to conditions on the relatively arid plains of central Wyoming. The type-localities of this and the other Nuttall species have been considered with the history of the genus. An additional feature, which concerns only this species, was merely mentioned there. In the specimen Ripley & Barneby 7956, from South Pass, Fremont County, Wyoming, two plants on the sheet differ in cer- tain minor features from a third. A pollen examination revealed that the one represents apomictic and the other two sexual ma- terial. In examining the Nuttall type-material of this species in the British Museum, Dr. Rollins noted (on a slip of paper now with the Gray Herbarium material of T. spathulata): “I would say that there is the possibility of 3 of the 4 plants belonging to one thing and the other to another. The three are less dense, less wooly ete.” In these same features the plants of Ripley & Barneby 7956 differ. The sexual plants are less dense and less woolly. In addition, the ray-corollas of the sexual plants are densely glandu- lar and those of the apomicts are nearly glabrous. Also, the ray- corollas of the sexual plants are more fully expanded than are those of the apomicts. Is it possible that Ripley and Barneby have sampled the same populations of sexual and apomictic plants which yielded Nuttall’s type-collection of this species? It seems likely that Nuttall collected this species in more than one locality. The plants of the type collection in the British Mu- seum are in flower. Nuttall’s specimens in the Gray and Phila- THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 123 delphia Academy Herbaria are well past flowering. Ripley & Barneby 7956 in flower was collected on June 14 and Porter 4428 in flower was collected on May 9. Dr. Porter’s material from near Alcova, Natrona County, Wyoming, grew at an elevation about 2000 ft. lower than South Pass where Ripley and Barneby made their collection. Possibly Nuttall first collected T. spathulata at a low elevation after it had flowered. As he proceeded west he may have obtained flowering material from some higher station. This higher station could be South Pass, through which his ex- pedition traveled. 17. Townsendia incana Nutt. Townsendia incana Nutt., Trans. Phil. Soc. 7: 305. 1840. Type: Nuttall s.n., “On the Black Hills, ei an alpine chain ee the sources of the Platte. )”, 1834 (BM, holotype, examined by Dr. R. C. Rollins; Gx!, px!, isotypes). Townsendia Fremontii T. & G., Bost. Journ. Nat. Hist. 5: 106. 1845. Original material: apparently collected in Wyoming by Fre- mont on his first expedition i in ee ine ownsendia arizonica A. Gra cad. 16: 85. 1880. Type: Palmer 204 in 1877, wiecbull gars Ca, Arizona (*GH, lectotype; *Mo, Ny, US, syntypes). Townsenda arizonica X incana M. E. Jones, Zoe 2: 248. 1891. Type: Jones s.n., south Little Colorado, Navajo or Apache Co., Ariz., 9 June 1890 (*pom, holotype). Townsendia incana Nutt. var. ambigua M. E. Jones, Zoe 4: 264. 1893. A variety based on several ig: designated collections. Townsendia diversa Osterhout, Bull. Torr. Bot. Club 55: 75. 1928. Type: Osterhout 6116, hills south of Grand Junction, Mesa Co., Colo. (*RmM, holotype). Pulvinate or suberect, taprooted, perennial (rarely biennial); caudex becoming much-branched, sometimes subterranean; stems mostly ao ously canescent, sometimes long and much-branched; leaves narrow spat late or oblanceolate, entire, acute, mucronate, moderately to densely os gose, up to 45 mm. long and 5 mm. wide; heads mostly terminal on stems, spe ge edunculate; oe campanulate, 8-19 mm. wide, 7-14 mm. high; hyllar aries in 3-4 (rarely 5) series, lanceolate, she scari- ous- and laceraclatemargined, pa de on the outer surface, 3-12 mm. long, 1-3.6 mm. wide; ray-florets ca. 10-30; ray-corollas mostly white on Hong, 1.5-3 nee dove: disk-corollas yellow and often pink-tinged, 3.7-6.5 oblanceolate, ha OR 2-ribbed, pubescent with dupe 3 loci hidiate ghen 3.0-4.7 mm. long, 1.0-1.8 mm. ond pee of c -39 apt barbellate bristles 0. if 6. : mm ee ¢ n of differ 124 JOHN H. BEAMAN and 36 in apomictic plants. Central Wyoming southwest to Nevada and central Arizona. Map 14. Plate XXII, fig. 1. Representative specimens. Arizona. Apache Co.: Water Lily Can- yon, 35 mi. northeast of Kayenta, Burton s.n., Aug. 1934 (tNa); 5 mi. west of Rock Point, Cutler 2196 (cas, GH, {Mo, us); Navajo Indian Reservation, about the north end of the Carrizo Mts., Standley 7332 (tny, us). Coconino Co.: ca. 24 mi. north of Flagstaff, Beaman 971 (*cH); 6 mi. west of Grand Canyon Bridge, Marble Canyon, 3800 ft. alt., Benson 129 (tuc); Moqui Wash, 8 mi. west of Winslow, 5000 ft. alt., Darrow 2699 (*cas) ; 6 mi. south of Kaibito, Navajo Indian Reser- vation, 6300 ft. alt., Darrow 2730 (tcas); U. S. Highway 89 along Vermilion Cliffs about 19 mi. east of Jacob Lake, Ferris 10262 (tps); between Winslow and Flagstaff, McKelvey 4507 (*cH); 15 mi. north- east of Tuba City, 5450 ft. alt., Kearney & Peebles 12889 (+ MICH, PpoM); 2 mi. east of Ashfork, Preece & Turner 2617 (*ws). Mohave Co.: brink of Toroweap Fault, Kanab Plateau, Grand Canyon National Monument, Toroweap, 6300 ft. alt., Cottam 14026 (*uT); Hackberry, cs, #RSA, UTC, ws). Gunnison Co.: hillside 15.6 mi. west of Gunnison on Highway 50, Preece &Turner 2795 (tws). Grand Co.: Kremmling, Osterhout 3497 (*nm). Mesa Co.: 1.4 mi. inside Colorado National Monument on road from Grand Junction, Beaman 801 (tcH); 1 mi. outside Colorado National Monument toward Glade Park, Beaman 802 (*cH); Grand Junction, Eastwood s.n., May 1891 (teom, us); 5 mi. west of Gateway in southern part of county, 4300 ft. alt., Harrington 4362 (*coto); Grand Junction, Jones s.n., 15 April 1891 (trpom); 4 mi. south of Mesa, 7800 ft. alt., Rollins 2190 (ps, cH, *NA, °US, UTC). Moffat Co.: Craig, Osterhout 5117 (RM); canyon near confluence of Green and Yampa Rivers, ca. 7000 ft. alt., Porter 3620 (tps, GH, RM, uc, us, wru). Montezuma Co.: McElmo Creek, Eastwood s.n., June 1892 (trom); entrance of Mesa Verde National Park, Nelson 10419a (GH, MO, tNy, RM, UC). Montrose Co.: Naturita, 5400 ft. alt., Payson 242 (cM, COLO, GH, MO, MONT, ¢RM, ws); hills near Montrose, 5800 ft. alt., Payson 657 (+nm); Paradox, 5400 ft. alt., Walker 90 (ps, GH, MO, NY, POM, , us); bench above San Miguel River, 24 mi. northwest of Naturita, Weber 3566 (COLO, Cs, DS, MONTU, OKL, PH, {RM, RSA, UC, THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 125 uTC, ws). Rio Blanco Co.: south side of White River, 2 mi. southeast of mouth of Wolf Creek, 6000 ft. alt., Graham 9039 (cm, tna); bluff of Gypsum Gap, T 44 N, R 16 W, east of Gladel, Weber 4725 (COLO, RSA, UC, Ws, WrU). Nevada. Lincoln Co.: Panaca, Jones s.n. 6 Sept. 1912 (trom); Cathedral Gorge, north of Panaca, 5000 ft. alt., Ripley & Barneby 6344 (tcas). N 5628 (tcas). Sandoval Co.: Jemez Biological Camp, Jemez Mts., Castetter 1170 (trm). San Juan Co.: Aztec, Baker 729 (Mo, Ny, POM, RM; US p.p., with 1 plant of T. exscapa). Utah. Beaver Co.: Frisco, Jones s.n., 1880 (*cH); Milford, 5000 ft. alt., Jones 1794 (cM, MO, “NY, PH, POM, US, UTC, “ws, yu). Carbon Co.: Price, Flowers F18- 30 (*ur); Sunnyside, Jones s.n., 15 Nov. 1907 (trom); 4 mi. north of Price, Maguire & Maguire 18342 (can, {Ny, urc). Daggett Co.: base of cliff, Sheep Creek, 12 mi. south of Manila, 6000 ft. alt., Hitchcock et al. 3913 (ps, NA, t}wru). Duchesne Co.: Myton, 5000 ft. alt., Jones s.n., 20 May 1908 (trom). Emery Co.: % mi. north of the San Rafael River crossing along the Hanksville-Greenriver Road, Holmgren, Boyle, & Will 7768 (tutc); Orangeville, 6000 ft. alt., Jones 5464c (pom, tus); first fork of Calf Springs Canyon, San Rafael Swell, Maguire & Maguire 18303 (cu, tNy, us, uTC, ws). Garfield Co.: wash east of Escalante, 5500 ft. alt., Cottam 4402 (tuc, ur); west slopes of Henry ts., 5500 ft. alt., Cottam 5524 (tur); open forest, northwest rim of Bryce Canyon, 8000 ft. alt., Degener & Peiler 16533 (tmMo); 7 mi. east of Escalante, Holmgren & Nielson 7736 (+ uc, uTC); 25 mi. south- east of Hanksville, Colorado River Rim, Parry s.n., 6 June 1941 (ny, futc); 5 mi. north of Hatch, 7000 ft. alt., Ripley & Barneby 8537 (tcas). Grand Co.: Thompson, Jones s.n., 7 May 1891 (*pom); West- water, 4400 ft. alt., Jones s.n., 28 June 1898 (tpom); Moab and vicin- ity, 1200-1500 m. alt., Rydberg & Garrett 8444 (tNy, RM, uc). Iron Co:. Cedar City, Parry 94 (cu, tmo, yu). Kane Co.: 8 mi. north of Kanab, Eastwood & Howell 9270 (tcas); 2 mi. east of Zion Park, Bridge north of Montecello, Clover & Jotter 2007 (tmicH); without edge of Green River Desert about 5 mi. northeast of Hanksville, ca. 1500 m. alt., McVaugh 14563 (cas, try); Fruita Arch Canyon, Ma- 126 JOHN H. BEAMAN guire & Maguire 18123 (cu, Ny, us, fuTc). Wyoming. Fremont Co.: canyon side above Wind River, 16 mi. southeast of Dubois, Beaman & Preece 507 (tws); hillside near Black Buttes Mill Site, 20 mi. north of Shoshoni, 5200 ft. alt., Freytag 78 (trm); Badwater (probably Badwater Creek near Lysite), Nelson s.n., 26 June 1910 (RM); River- ton, Nelson s.n., May 1920 (trm). Natrona Co.: 4 mi. west of Alcova, Beaman 875 (tcH); Alcova, Goodding 166 (Mo, RM, us). Sweet- water Co.: 6 mi. north of McKinnon, Goodman 3105 (mo, Ny); hills near Rock Springs, 6200 ft. alt., Larsen 17 (trm); Granger, Nelson 4622 (cH, tMo, Ny, RM, us). Uinta Co.: Ft. Bridger, Nelson 4599 (CM, CS, MONT, NY, POM, RM). Apomicts of T. incana usually are easy to determine as that species; sexual populations sometimes are more difficult. A most- ly-sexual geographic segment of this species which has long caused taxonomic difficulties is located in northern Arizona. Gray described these plants as T. arizonica and distinguished them from T. incana by their long ray-pappus. Subsequently, long-ray- pappus forms of T. incana from many parts of its range were referred to T. arizonica. It is evident from the material now available that the ray-pappus length is much too variable to be a useful taxonomic character in T. incana. Larsen (1927) accepted T. arizonica and referred some of the western Colorado specimens of T. incana to T. arizonica. Her key was designed to separate the two by leaf length and shape. How- ever, the vigor of the plant influences these characters which are not constant, even in obligate apomicts. In addition, Larsen did not distinguish T. annua and T. Fendleri from T. strigosa, and her key does not satisfactorily separate many specimens of T. strigosa (as she understood it) from T. incana. The most atypical portion of T. incana is represented by three sexual collections, Rollins 2190 and Osterhout 2761 and 3474, from northwestern Colorado. The habit of these plants is unlike that of typical T. incana. The former have short, thick caudices, not unlike T. exscapa. The heads, borne on short peduncles, are somewhat embedded in the tufts of leaves. In their other features these plants are characteristic of T. incana. They are from rela- tively high elevations and possibly represent elevational and geo- graphic polymorphy in T. incana. Another sexual population which diverges from typical T. in- cana is Jones 1794 from Beaver County, Utah. These plants are distinctive because of their reduced size and very heavy, ca- nescent pubescence. The apomicts in this area are morphologi- cally more typical of T. incana. THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 127 A small amount of variation in T. incana is not attributable to geographic differentiation. In San Juan County, Utah two collec- tions, Cottam 5816 and Holmgren 3183, are intermediate between T. annua and T. incana. Although these plants seem to be short- lived perennials, their general aspect is that of T. annua. The disk-pappus of Holmgren 3183 is shorter than the disk-corollas, a characteristic of T. annua. The stems are canescent, and the phyllaries are in 4-5 series, characteristic of T. incana. Hybridiza- tion of the two species in this area therefore seems probable. The ranges of T. incana and T. annua overlap in an extensive area, and their flowering periods coincide. Probably they are at least partly interfertile. Herbarium material, however, suggests that hybridization is very rare. The great abundance of apomixis, or more directly, the rarity of sexual material of T. incana prob- ably restricts the frequency of hybridization between these two species. It is possible that some of the T. incana populations in Mesa County, Colorado and Grand County, Utah have been affected through hybridization with T. strigosa. Several collections from this area are of suberect plants, characteristic of T. strigosa but not of T. incana. However, since some of the best characters for distinguishing these two species involve the habit, it is very diffi- cult, with as variable a species as T. incana, to be reasonably sure, on the basis of herbarium specimens, that hybridization has oc- curred. Jones s.n., 7 May 1891 (Grand Co., Utah) is an example of possible crossing. Five plants of this collection (Pom 39664) are fairly typical T. incana. A sixth plant has a swollen root-stem junction with persistent, broad basal leaves, resembling T. stri- gosa. This plant retains the canescent pubescence of T. incana.. Townsendia incana occurs at elevations from approximately 4000 to 8000 ft. It is especially abundant on the Colorado Plateau in sandy soils amidst junipers and pifions. It flowers in the spring and summer at times when sufficient moisture is available, usually in May and early June. Some flowering material, however, has been collected in April, July, August, and September. All ma- terial which has been available for greenhouse and garden cul- tures exhibits a more or less indeterminate flowering period. Some plants lived and flowered in the greenhouse for as long as two years. Townsendia incana is most closely related to T. F endleri. The latter was probably derived from ancestral stock nearly or quite like the former species. No other species are very closely related to these two. 128 JOHN H. BEAMAN 18. Townsendia Fendleri A. Gray Townsendia Fendleri A. Gray, Mem. Am. Acad. 4 (PI. Fendl.): 70. 1849. Type: Fendler 350, gravelly hillsides, Santa Fé, Santa Fé Co., New Mexico, May-July 1847 (cx, holotype; Mo, uc, isotypes). Caulescent, decumbent or suberect, or infrequently rosulate, taprooted perennial (or sometimes biennial ?); caudex becoming much-branched, often subterranean; stems much-branched, short or becoming limes, stri- gose or pilose-strigose, sometimes canescent, up to 3 dm. long; leaves nar- i stems; involucres campanulate, 6.4-13 mm. wide, 5.0-8.5 mm. high; phyl- laries in 4-5 series, lanceolate or ovate-lanceolate, acute, ciliate and broadly scarious-margined, strigose on the outer surface, 2.7-8 mm. long, 1-3 mm. , pu t with du os zoshidints hairs; ray-achenes more densely duction sexual. South-central Colorado and north-central New Mexico. Map 4. Plate XXII, fig. 2. Representative specimens. Colorado. County unknown: valley of the Arkansas, Wolf 517 (¥, pom, us). Chaffee Co.: Salida, Baker, Earle & Tracy 1016 (¥, Mo, NY, POM, us); Salida, Osterhout 3424 (Ds, GH, NY, POM, *RM); Arkansas River near Poncha Pass, Jones 767 (CAS, CM, COLO, DS, GH, "MO, NY, OC, POM, UTC); hills southeast of ridge along the Arkansas ee & mi. northwest of Salida, Waterfall 11511 (OKL, RsA, *uc). t Co.: Texas Creek, Brandegee 951 (MO, PH, uc); sandbar, tiie City, 5280 ft. alt., Brandegee s.n., June 1876 (*uc). fiedand Creek, Brandegee 1294 ( GH, MO, UC). Pueblo Co.: Pueblo, Bethel s.n., 2 July 1897 (*cs). New Mexico. Rio Arriba vie ig Ranch, Rio Chama drainage, Goodwin s.n., 24 Aug. 1932 ; 13 mi. south of Cebolla, Ripley & Barneby 1 0294 (cas); Ojo Cidenthe 6000 ft. alt., Smith s.n., 27 Aug. 1894 (pH). Sandoval Co.: 6 mi. west of San Ysidro, 5500 ft. alt., Ripley & Barneby 8344 (*cas); San Ysidro, Nelson 11619 (*nm, uc). Santa Fé Co.: Santa Fé, Degener 4689 (Ny); near Espafola, 5600 ft. alt., Heller & Heller 3547 (np, NY; US p.p., with 1 plant of T. annua); 2 mi. north of Santa Fé, ca. 7000 ft. alt., Hitchcock et al. 4186 (Na, *wtv). Since Larsen (1927) made no distinction between T. annua and T. strigosa, and considered T. Fendleri a worthless taxon, she reduced the latter to T. strigosa. Her confusion, of course, was between T. annua and T. Fendleri. No modern material of these THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 129 species is difficult to determine. However, some of the specimens she had, especially Brandegee 951 and Jones 767 have consider- ably longer stems than is usual for T. Fendleri. In addition, on one sheet she examined (mo 121087) Brandegee 951 was accom- panied by the following undated scribbled note in Gray’s hand- writing: “951 fits much of my Townsendia strigosa well. I dare say Fendleri will not be very Im’pt.!” This note, and the poor specimens mentioned above, must have influenced Larsen to reduce T. Fendleri. In the confusion, Brandegee 1094 further complicates matters; Mo 121070 with the number 1094 is clearly T. Fendleri. Brandegee 1094 in the New York Botanical Garden Herbarium and the Philadelphia Academy Herbarium is clearly T. annua. A Brandegee collection in the Gray Herbarium without a number but with essentially the same label data “Adobe plains, San Juan R.” is also T. annua. Brandegee 1294 (cu, Mo 121088, uc) Huerfano Creek, Huerfano Co., Colo., is T. Fendleri. It is probable that Brandegee 1094 is not actually a mixed collection of the two species. Very likely, because of the similarity of the num- bers 1094 and 1294, the label 1094, with its accompanying data, was erroneously transferred to specimen Mo 121070 which should be Brandegee 1294. The high degree of similarity of the two plants on this sheet to Brandegee 1294 supports this idea. More- over, it is improbable that T. Fendleri occurs on the “adobe plains” of the San Juan River valley. In most areas, and in most collections, T. Fendleri and T. annua are easily distinguishable. But around Santa Fé, New Mexico, where their ranges overlap, the two species closely re- semble each other. Townsendia Fendleri especially seems to ap- proach T. annua. The mixed collection, Heller & Heller 3547 from near Santa Fé, indicates a considerable similarity of the two.2, Townsendia Fendleri is probably not a perennial in that area, while it is in the other portions of its range. In northern New Mexico it may have obtained genes for a shorter life-cycle from T. annua. Except in that area the two species probably are fairly well isolated geographically. Townsendia Fendleri occurs at moderate elevations mostly within the pifion-juniper belt while T. annua occurs at lower elevations below the pifons and junipers. Although T. annua and T. Fendleri have been confused be- cause of their similarity, the closest relationship of T. Fendleri seems to be with T. incana. These two species are allopatric (considering the range of the sexual forms of T. incana only) ® Heller & Heller 3547 in Mo, Msc, and vs p.p. is of T. annua; in Np, ny, and us p.p., this collection is made up of plants of T. Fendleri. 130 JOHN H. BEAMAN and T. Fendleri probably is a specialized, reduced form from more generalized T. incana stock. The habit of the two species (at least of some forms of T. incana) is almost identical. The differ- ences are mainly quantitative ones, paralleling those between T. strigosa and T. annua. 19. Townsendia strigosa Nutt. Townsendia. strigosa Nutt. Trans. Am. Phil. Soc. 7: 306. 1840. Type: Nuttall s.n., “On the Black Hills, (or eastern chain of the Rocky Mountains,) near ‘the banks of the Platte.” , 1834 (Bm, holotype, ex- amined by Dr. R. C. Rollins; cu!, pul, isotypes Townsendia incana Nutt. var. prolixa M. E. Jones, Contrib. West. Bot. 13: 15. 1910. Type: Jones s.n., Chepeta Well, Uintah Co. (fF), Utah, 6000 ft. alt., 23 May 1908 (*po, holotype). Caulescent, taprooted biennial; root-stem junction enlarged; stems branched at the base, few to moderately numerous, branched again just below the heads in a cymose manner, mostly spreading-suberect, usually red with cyanic pigments, striate, strigose-pilose, up to m. long; basal tion mostly sexual, rarely apomictic. No eastern U Utah and ait ares probably also in cae northwestern Colorado. Map 17. Plate XII, fig. 3 Representative specimens. Utah. Carbon Co.: 9-mile Canyon, 5000 ft. alt., Jones s.n., 22 May 1896 (*pom).. Duchesne Co.: north Duchesne along the Victory Highony, Osterhout 6196 (*Rm). Uintah Co.: 5 mi. northwest of Dinosaur National Monument, 5500 ft. alt., Graham 7671 = .p., with 2 fragments of T. incana); bench west of Green River, ah A of mouth of Sand Wash, 5000 ft. alt., Graham 7946 (cm, ai paertos slope of Big Pack Mountain, west of Willow THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 131 Creek, Thorne’s Ranch, Uinta Basin, 5500 ft. alt., Rollins 1705 (*cn, Ny). Wyoming. Carbon Co.: Solon, Williams s.n., July 1897 (*Rm). Sublette Co.: roadside, Big Piney, Payson & Payson 4350 (tGH, Mo, MSC, PH, tRM, ws). Sweetwater Co.: Green River, Nelson 3031 (*cn, ILL, MO, NY, *RM, us); Green River, Nelson 4724 (*cm, cs, MONT, POM, WS); mi. west of Green River, Payson & Armstrong 3205 (GH, ILL, MO, 4X PH, POM, *RM); desert about 40 mi. south of Rock Springs, on ery Hiawatha case Porter 4573 (DAO, GH, *RM, RSA, WTU). Uinta Co.: 6 mi. out of Mountainview toward Lonetree, H olmgre no Tillett 9474 (*uc, utc); hillside near Lyman, 6500 ft. alt., Rollins 1636 (ps, *NA, US, WS). Townsendia strigosa is not restricted to the Green River shales, but it occurs mostly within the bounds of the Green River forma- tion in northeastern Utah and southwestern Wyoming. From the Ns T STRIGOSA ye & SEXUAL <& APOMICTIC 7 @ T. ANNUA j N AP Aa Pipi isie ns distribution of Townsendia annua and sexual and apomictic r. strig 132 JOHN H. BEAMAN observations of Mr. and Mrs. W. L. Bartholomew (personal com- munication ), residents of the Uinta Basin of Utah, I have learned that the species is abundant in that area. It flowers in May and early June, and often occurs in fairly dense, conspicuous stands. It sometimes is mixed among apomictic plants of T. incana and sometimes occurs in pure stands. Sheep browse the plants, and by the end of June scarcely any standing plants are to be found. Some variation in this species, especially near Green River, Wyoming, suggests hybridization with T. annua. Most of the plants of Nelson 3031 and 4724 have the disk-pappus shorter than the disk-corollas. Some plants have ovate phyllaries, in others the phyllaries are merely broadly lanceolate. The short disk-pappus and ovate phyllaries are characters of T. annua. One other speci- men, Jones s.n., 22 May 1896, from Carbon County, Utah, very close to the range of T. annua, also has ovate phyllaries. Although T. strigosa and T. annua are allopatric, hybridization could be expected between them. Their ranges adjoin and there is no geographic barrier. They are very similar and probably have no strong genetic barriers to interbreeding. Townsendia strigosa is closely related to T. annua, which is an evolutionarily more advanced species. The two have no other very close relatives. Townsendia strigosa superficially resembles T. florifer but is readily distinguished from that species by its glochidiate rather than bifurcate achenial hairs. It has been con- fused with T. incana; however, these two species are distinct enough so that no confusion is necessary. As might be expected i in a biennial, apomixis is unimportant in T. strigosa. It is interesting, but of minor significance, that the only known apomictic collection is from the northernmost station in the range of the species. 20. Townsendia annua Beaman, sp. nov. Type: Maguire 13509, sand bed of Cottonwood Wash, vicinity of Wayland’s Ranch, 1% mi. north of Bluff, San Juan Co., Utah, April 19, 1936 (*cH, holotype; CAN, PH, UC, UTC, WTU, isotypes). ahre = ranching at the base an nd a , minutely striat e, pin .2-2.5 dm. long; basal leaves short-lived, pedir or spatulate, entire, often mucronate, oe strigose on both surfaces, ie to 3 cm. long; cauline clustered on young branches below unexpanded heads, usually not surpass- "the leafy stems, or pedunculate on inconspicuous peduncles; involucres 8-16 mm. wide, 4.5-7.5 mm. high, THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 133 broadly expanded at maturity; phyllaries in 3, rarely 2-4, series, elliptical, obovate or ovate, obtuse, acute, or rarely slightly ae the margins scarious and ciliate at least above, 2-6.7 mm. lon ng, 1-2.6 . broad, the outer shorter than the inner, lightly or rarely moderately Sard -pilose on the outer surface, the inner series nearly or entirely Poa ray-florets ray-corollas white to pink or light ae often arker longitudinal streak on the abaxial surface, 5-9 mm. long, 1.5- 27 mm, i isk-corollas yellow, sometimes pink- to erMintipodds ng; achenes oblanceolate to obova a e, compressed, 2- _— those of t dhe ay-florets rarely oiainet oy to moderately pubescent duplex, glochidiate hairs, those o ray-florets o— slightly more simacen nt than those of the disk-florets, “ae eat those of the ra aye , often papil- lose, 2.0-2.8 mm. lon mm. e; pappus of the ray- ay ae ne of short, plurisetose, a brite ae exceeding 1 mm. ength; ee of the disk-achenes of ca. 15-28 caren barbellate ue 1.8- m. long, very rarely siete the length of the disk-corollas. Repro- risen pale East-central Utah south to southern Arizona, New Mexico, arg Texas, probably in adjacent northern Mexico. Map 17. Plate XXII, g. 4 Herba annua. Caules decumbentes, pauci ad numerosi, striati, strigosi, 2.0-2.8 mm. 16 . latis; radii semper breve, squam lato (squamellis conforms) disci pappo etestens: quam disci Satok lae plerumque breviore Representative specimens. Arizona. Apache : Four Corners, 5000 ft. alt., Deaver 4037 (cas); oe ae just inthe of Petrified Forest, near the Rio Puerco, Nelson & Nelson 2144 (*RmM). Coconino Co.: (*urc). Navajo Co.: between Kayenta and Betatakin, Eastwood & Howell 6575 (cas); between Winslow and Holbrook, McKelvey 4561 (cH); Holbrook, Ward s.n., 15 June 1901 (ny, us). Pinal Co.: near Dudleyville, Griffiths 3673 (wa). Yavapai Co.: Beaver Creek, Purpus 8300 (mo, uc, us). Colorado. Delta Co.: between Delta mtn Grand Junction, 4800 ft. alt., Penland 1763 (cas, RSA). Montezuma Co.: Mancos, southwestern Colo., Eastwood s.n., June 1890 ( copa gg “Mont. rose Co.: Paradox, 5400 ft. alt., Walker 93 (GH, ILL, MO, NY, POM, *RM, US, Ws, \ wtu); Naturita, 5400 ft. alt., Payson 321 (CM, COLO, Ds, F, MO, MONT, *RM, Ws). New Mexico. Bernalillo Co.: plains near Albuquer- que, Meus 31206 (mo, pH). Dona Ana Co.: Mesilla Valley, near Mesilla, Standley s.n., 23 May fea (*us); Mesilla Valley, 3850 ft. alt., 134 JOHN H. BEAMAN Wooton & Standley s.n., 2 April 1907 (ps, Fr, Mo, Ny, *RM, us). Grant Co.: Mangas Springs, 18 mi. oranda of Silver ees 4300 fe ale Metcalfe 15 (cas, Ds, GH, ILL, MO, POM, “RM, UC, US). Lindoin Co.: Carrizoza, Earle 607 (ny). McKinley Co.: along Highway 66, 15 mi. east of G Gallup, Nelson & Nelson 2174 (Mo, NY, *RM, UTC). Otero Co.: desert, just outside entrance to White Sands N ational Monu- ment, Goodman & Waterfall 4987 (okL). San Juan Co.: Aztec, Baker 728 (F, GH, MO, ND, NY, POM, *RM, US); 2 mi. northwest of Waterflow, Cutler 3331 (®mo, NA, vs). Sandoval Co.: Torreones arroyo, Keesecker 38 (oKL). Santa Fé C55 Santa Fé, Fendler 351 (GH, Mo, PH); near Espanola; 5600 ft. alt., Heller & Heller 3547 (*Mo, MSC; Us p.p., with 2 plants of T. F endleri). Sierra Co.: Hillsboro, north of Percha, 5500 ft. alt., Metcalfe 1510 ( cas, F, GH, MO, NY, POM, UC, US). Seoorre Co.: Datil F orest, above Walter Medley’ s Ranch, ca. 9 mi. north of Magda- lena, Eggleston 16197 (mo). Taos Co.: Taos, 6900 ft. alt., Castetter 1302 (*nm). Valencia Co.: Mount Taylor, between coal mine and Grants, 7000 ft. alt., Parker 2318 (rsa, uc). Texas. El Paso Co:: El Paso, 3700 ft. alt., Ripley & Barneby 4211 (cas). Hudspeth Co.: in a basin between hi Ils, 4 mi. east of Hueco, Waterfall 3867 (GH). Utah. Emery Co.: Green River, Jones s.n., 9 May 1890 (*Mo, msc, POM, UC, US); Mounds, Jones s.n., 5 Sept. 1901 (pom); Red Knoll enclosure, vic. Buckhorn Reservoir, San Rafael Swell, Maguire 18493 Sa NY, US, UTC, WS; WTU p.p., with 1 plant of T. incana). Garfield Co.: Henry Mts., 5000 ft. alt., ‘beat 1059 (ur). Grand Co.: 5 mi. south of Crescent cgpatcag Holm mgren & Hansen 3295 (CAN, GH, IDS °MO, NY, UC, US, UTC, Ws, WTU); Westwater, Jones, s.n., 20 May 1901 (PoM); Moab a vicinity, 1200-1500 m. alt., Rydberg dr Garrett 8441 (Ny). San Juan Co.: Cottonwood Wash, vicinity of Wayland’s Ranch, 1% mi. ame of Bluff, Mase 13509 (can, "GH, PH, UC, UTC, WTU). Previously this species has been identified as T. strigosa.* It is indeed closely related to that species. The present material, however, is separable into two clear-cut groups, each with its own range of variation which hardly overlaps that of the other. Townsendia annua and T. strigosa may be distinguished by the characters tabulated below. In addition to the differences indicated in the table above, there are others which do not lend themselves to tabulation. Townsendia annua. is more delicate in appearance and has very slender branches.. On herbarium specimens it usually has no anthocyanin in the stems while T. strigosa usually does. The hyllaries of T. annua usually are shorter and broader with more conspicuously lacerate margins than are those of T. strigosa. * Nuttall’s type material of T. strigosa in American herbaria is rather depauperate. Superficially, his plants do closely resemble T. annua. It is not surprising, therefore, € two species previously have not been distinguished. THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 135 Townsendia annua Townsendia strigosa Annual. Basal leaves soon deciduous. Root-stem junction not enlarged. Phyllaries mostly in 3, rarely 4 Biennial. Basal leaves persistent. Root-stem junction enlarged. es mostly in 4, rarely 3, eri series. Disk-corollas 2.3-3.7 mm. long. Dike 3.3-5.0 mm. long. — 2.0-2.8 mm. long. Achenes 3.0-4.0 mm. long. Disk-pappus 1.8-3.0 mm. long, Disk-pappus 3.3-5.5 mm. long, esa shorter than the disk- about equal to or longer i. the corollas disk-corollas. Aside from the purely morphological evidence for this species being a true annual, the flowering time also gives that indication. Specimens in flower collected in every month from February to October are included in the present material. May and June collections are most frequent. Townsendia annua appears to be derived from a prototype nearly or entirely similar to T. strigosa. The distinguishing char- acteristics of T. annua are mostly quantitative, reduced features of T. strigosa. The comparative measurements in the table above suggest this fact. Townsendia annua probably has become spe- cialized for a somewhat more xeric existence than that of T. stri- gosa. This adaptation has come about primarily by development of the annual habit and correlation of the flowering season with the availability of water. Putative hybridization of T. annua with T. strigosa is consid- ered in the discussion of the latter species; that with T. incana under the discussion of T. incana, and that with T. Fendleri under the discussion of that species. 21. Townsendia mexicana A. Gray Townsendia mexicana A. Gray, Mem. Am. Acad. 4 (PI. Fendl.): 70. 1849. Type: Gregg s.n., Saltillo, Coahuila, Mexico, March 1847 (cu, holotype; Mo, Ny, isotypes). ow, caulescent, taprooted perennial; stems often slightly woody at the be loosely branched from the base upward, usually nisconbent some- times terminally ascending and suberect, strigose-sericeous, up to 2.0 dm. long; leaves more or less evenly distributed on the stems, linear to narrowh oblanceolate, acute, mucronate, entire, 0 1-2 small lateral lobes near the tip, strigose, 5-25 mm. long, 642% si mm. wide; pedun = rarely 2-3 at the ends of the seem strigose, den involucres 6-12 mm. wide, 3. mm. high, broadly pan sey at jiathtey ehyllaries in 2 (rarely 3)" series, * elliptiocobowate, scarious-margined, ciliate 136 JOHN H. BEAMAN near the obtuse-rounded (rarely acute) apices, the outer 2.5-4.5 mm. long, 0.5-2.2 mm. wide, lightly to moderately strigose on the abaxial surface, the inner 3.5-6.0 mm. long, 0.8-2.4 mm. wide, lightly strigose to nearly glabrous; ray-florets ca. 15- 35; ray-corollas white on the adaxial surface and some- to lightly pubescent with duplex, ee hairs, Tay- een nes more pu- ego than those of re disk, both or only the ray-achenes papillose, 1.8- mm. long, BL. m. wide. 2n = 18. Reproduction sexual. Southern Seahorse south to he D. F., Mexico. Map 16.. Plate XXIII, fig. 1. Representative specimens. Mexico. Coahuila. Ca. 3 mi. southwest of Saltillo, Beaman 1002 (*cx); Saltillo, Palmer 499 in 1880 (*mo, , Us, yu); valley 15 kms. west (?) of Conception del Oro just within Coah. border, 2300 m. alt., Stanford, Retherford & Northcraft 509 (Ds, GH, *MO, NY, UC, WTU). Distrito Federal. Vicinity of Rancho de Pics near San Pablo, 15 mi. south of Mexico City, 7350-7600 ft. alt., Happ 296 (*mo). Hidalgo. EI Arenal, slopes at Km. 100 Pan Amer- ican (Laredo) highway south of Actopan , 2500 m. alt., Moore 1395 (*cu, uc); bluffs near Tula, 6800 ft. alt., Pringle 6573 (CAN, Cas, OM, F, CH, MEXU, MO, MSC, ND, “Ny, PH, UC, us). Nuevo Leén. 17 southeast of Galeana, Schneider I 115 (F). Querétaro. Near Higueril. las, Rose et al. 9791 GH, us). San Luis Potosi. Estacion Catorce, Sierra Madre Oriental, 2000- 2100 m. alt., Pennell 17575 (pu, us). Tamaulipas. Near reservoir of Miquihuana at base of hills, Stanford, Taylor & Lauber 2384 ( uc, wru). Zacatecas. Between La Honda and Santa Rita via Pinos, jomata 7472 oo cide near Concepcion del Oro, Palmer 252 in 1904 (¥F, GH, Mo, This Mexican endemic probably is iis most highly evolved species of Townsendia. Its diminutive size, two series of phyl- Jaries, very short ray- and disk-corollas, short pappus, and small achenes all are features of reduction. It is the only member of the genus with lobed leaves. All of its meiotic chromosomes are distinctly smaller than those known in any other species of Town- sendia (cf. Plate I, fig. 3).4 Townsendia mexicana has no very close relatives but ‘belongs with the group of species which in- clude T. incana, T. Fendleri i ag strigosa, and T. annua. Town- sendia Fendleri, T. annua, and T. mexicana exhibit a very strong parallel development. The range of T. mexicana probably coincides with the occur- rence of alkaline soils in central Mexico between the elevations of ca. 5000-8000 ft. It has been collected in flower from March to September. ‘Unfortunately, the chromosomes of T. annua, a species which shares many reduced features with T. mexicana, have not yet been examine THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 137 Species ExCLUDED FROM TOWNSENDIA Townsendia Wrightii A. Gray, Bot. Mex. Bound. Surv. p-. 78. 1859. = Aster Wrightii A. Gray, Smithson. Contrib. Knowl. 5 (Art. 6): 75. 1853. LITERATURE CITED Avers, Cuartotre J. 1954. Chromosome behavior in fertile triploid Aster hybrids. iy a 39: ei en Baneeee Ben: . L. Sressrns. 19338. The American species of Crepis: their reetonaee and disteibiiek as affected by polyploidy and apomixis. Carnegie Inst. Washington, Publ. No. 504. 200 Barnesy, R. C. 1947. A new riiceaadaes Parthenium. Leafl. West. Bot. 5: 19-22. Beaman, J. H. 1954. Chromosome numbers, apomixis, and interspecific hy- bridization i in the genus Townsendia. Madrofio 12: 169-1 BENTHAM, G. 1873. Nowe on the classification, er and geographical dis- tribution of Compositae. Journ. Linnean Bunton, Lian. 910, Histology of Townsendia ‘ppcabd and Lesquerella spathulata. Kans. Univ. Sci. Bull. 5: 183- Cain, S. A. 1944, Fundtion of plant geography. Harper & Brothers. New York and eee 556 Cronguist, A. 1955. Vaseuise plants of the Pacific Northwest, pt. 5. Univ. of Washington Press. Seattle. Drummonp, T. 1829. Sketch of a journey to the Rocky Mountains and to the Columbia River in North America. Hook. Bot. Miscell. 1: 178-219. Esau, ang ae 1944, oe and cytological studies on beet mosaic. Journ. Agr. Res. 69: 95-1 Facknunp, F. 1941. Die Embryosackenvcklung bei Tanacetum a und einigen aoe eagles & Svensk Bot. Tidskr. 35: 157-176 ee ee orogenesis, ‘ict aaeeioasebinca und pseudogame Samenbildung be Rudbeckia laciniata L. Acta Horti Bergiani 14: 39- ————. 1947 a. Macrogametophyte formation ~ two agamospermous Eri- geron — Acta Horti Bergiani 14: 221 one ae . Die Restitutions- und K Kontationker der Hieracium- Mikrosporogenese. Svensk Bot. Tidskr. 41: 247- ———. 1947 c. Makrosporogenese und Fmnbrostelidung bei agamo- spermischen et ong gh biotypen. Svensk Bot. Tidskr. 41: 365-3 Goopman, G. J. 1943. The story of Parthenium olpiee um. Madrono 7: 115-1 GRAUSTEIN, JEANNETTE E. 1951. Nuttall’s travels into the old ee , an unpublished 1810 diary. Chronica Botanica Gray, A. 1880. Contributions to North American botany. I. Notes on some Compositae. Proc. Am. Acad. 16: 78-102. 1888. Synoptical flora of North leaage oe ii Gamopetalae. Second ed. Smithsonian Institution. Washing Gustarsson, A. 1934. Die Pacicaiaitoce oe Foralsparsdkes, Hereditas 19: ; 1935 Studies on the mechanism of parthenogenesis. Hereditas 21: 1-112. 1948. Polyploidy, life form, and vegetative reproduction. Here- ditas 34: 122; 138 JOHN H. BEAMAN agree G,..1950. Embryological studies in the Compositae. I. Anthemideae- Salen a Acta Horti Bergiani 15: 135-168. 195 Embryological studies in the rae ee II. Anthemideae- Cheyeaiihieyainae: Acta Horti Bergiani 16: 1951 b. Embryological studies in te Compositae. III. Astereae. a Horti Bergiani 16: 73-120. eae C. B. 1948. Notes on the genus Townsendia in western North Amer- ica. Madrofo 9: tea 241. Hitcucock, C, L. anp J. W. THompson. 1945. Noteworthy plants of Idaho. eafl. West. Bot. i’ 197-206 Horrmann, O. 1897. in A. Engler and K. Prantl, Die Natiirlichen Pflanzen- f n 4 ‘er 2. Hormeren, I. 1919. ‘Tbainli Studien tber die Fortpflanzung bei den Gattungen Erigeron und Eupatorium. Kungl. Svenska Vet.-Akad. Handl. Bd. 59, No. 7. 118 Hooker, W. J. 1834. FI. Bor.-Am. II, pt. 7. 1-48. London. Henry G. Bohn. Jepson, W. L. 1934. The overland journey of Thomas Nuttall. Madrofio 2: 143-147 Jones, M. E. 1893. Contributions to western botany. 5. Notes on Townsendia. rs : 260-265. Lanjouw, J. al. oe go Code of Botanical Nomenclature. egnum Weceabite 8: p. Lanjouw, J. anp F. A. Seat 1956. Index Herbariorum. Pt. 1. The heterts of the world. Ed. 3. Regnum Wocecanile 6: 224 pp. Larsen, EstHer L. 1927. A revision of the genus Towmnsendia. Ann. Mo. ‘Al, MacLoskIE, G. 1883. The achenial hairs of Townsendia. Am. Nat. 17: 1102- Mavartzon, J. 1933. Studien on die Embryologie der Familien Crassulaceae psk de d. 152 pp. Saxifragaceae. Gleer a Univ.-Bokhandeln. Lun Nevnce T. 1840, 1841. Picstvis tiaus of new species ane genera of plants in the natural order of the Compositae, etc. Trans. il. Soc. 7: 283-453. Ricuarpson, J. 1823. Bot. App. in J. Franklin, Maruive ms a journey to the shores of the Polar Sea. 729-768. London. John Murr Rouuins, R. C. 1950. The guayule rubber plant and fis ralouvek Contrib. Gray Herb. 172: 73 pp. Saou L. H. 1946 a. Revision of the genus Chaetopappa DC. Wrightia 1: 3.8 —. 1946b. The genus Dichaetophora A. Gray and its relationships. Wrightia 1: 90-94. Sressins, G. L. 1950. Variation and evolution in plants. Columbia Univ. Press. Tanara, M. 1915, Parthenogenesis i in Erigeron annuus Pers. Bot. Mag. Tokyo 29:. 245-254. THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA ATE XIII. Habitats of Tow one Fig. 1. The iota Hills in Roger Mills County, Oklahoma. Townsen nsis is an endemic of this area, restricted imestones IOSEC y am erosion of the deep, black ait of the high plains ‘ig. 2. Near Gothic, Gunnison County, Colorado. Apomicts of the eager : Rothr ockii grow under snowbanks 1 sandstone soils at the tops of mountains. They flower as oO the snow melts off. Apomicts of T. leptotes are “elles the mountain lee also, but on drier site S. 139 140 JOHN H. BEAMAN ~ a s a ™. oS Bros eS Pirate XIV. stn its of T oeopesenra: Fig. 1. In the Colorado he apa Hou near Grand fais ‘tion, Mesa County, Colorado. A typical locality for T. incana. Both sexual and apomictic oli ints are ieee here. Fig. 2. The type- ton iy of T. montana var. montana, above the Flagstaff Mine at Alta, Salt Lake County, Utah. The plants are restricted t she. crest of this subalpine limestone ridg THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 141 Pratre XV. Habitats of Townsendia. Fig. 1. A white airy outcrop of the Green River formation in the Uinta Basin of Utah. The paler =mic mensana var. mensana ee ; 2. . is found amidst ~ shale fragments. Fig. 2. Fairylanc Sasi near Bryce in Garfield Cou ah. The endemic T. montana var. minima grows i ffs. Ut fragmented "ial at the tops of these cli =) =] ~ a oO Sy as 2 = S 142 JOHN H. BEAMAN PrateE XVI. Mt. Borah, the highest peak in Idaho. Apomicts of T. condensata, his T. leptotes if montana var. montana and T. Parryi grow on the limestones of mountain Fi 2. The Satanka formation, near Alcova, Natrona County, Wyoming. Nuttall m lected T. spatht ta here in 4, e de ] ht little plants of this species are abundant in localized areas among the shale fragments. 143 THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 2 Town clean dete Feed. fof : es th Gale * Cestafewt gee SRN e We i ‘ oe a pe ¢ : A eatin, Ce, se Pec a Aa gerne “ ‘a A idee mo eek wn. DR a. ROMANE. cco Sle = Pirate XVII. eee of ay original collections of Townsendia (see explanation in the discussion o exsca 144 JOHN H. BEAMAN Ff OEE OE: rl sts Paes papapipied Oe fret say Se a 3 de Sete since, 49 4 * 6 RE ow $8, Pirate XVIII. asi gp a hedge ay ets of species of 4 wanes: Fig. = The lectotyp T. for . Fig. 2. A representative specimen of T. Fig. 3. A Ria resentative ec rok i riediiioes. Fig. 4, A representative epson of om tex THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 145 FLORA of MONTANA Set Me Cinch a CUMS EY?S TOWSON LA Meera ocr: om TAU OF mouNTAIR Shy. CATH OF FOCM Puak, Tavien brs. 2 hewuat 1 teae 3 4 : PLATE i ti of representative specimens of species of Tow — Fi glabella ve ner 2892). Fig. 2. Townsendia Rothrocku (Bea We b v; +o Protie S14). Fig. Tow cade ot mneenene var. montana. Fig. 4 I sendia montana var. minima (Hite hAcock 2962). JOHN H. BEAMAN p ON Sey 4 PLATE XX. pceiaers of representative —— of species of Townsendia. Fig. 1. The holotype of T. mensana var. mensana. Fig. 2. The holotype of mensana var. Jonesii Fig. 3. A representativ xual specimen of T. / neb 7589). I 4 ned’ . A representative — s specimen of T. leptotes (Hol @ Shaw 7640). Fig. 5. holotype Hookeri (Clokey 3 12 061 Fig. e ro ae! 6. Part of Drummond’s collection (Drummond 573) of T. Hookert from the Rocky Mountains. THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 147 Loeergcdineerngtnee ae Ra ee SARE PLANTS OP AREZODSA 4 Pirate XXI. Pho gt of representative specimens of spec cies of Townsendia. Fig. 1. Town dns exs “apa. a g. 2. Townsendia Parry. Fig. 3. Tow erie Pony. Fig. 4. Townsendia pcaniat 148 JOHN H. BEAMAN Pirate XXII. Photographs of representative specimens et species of Townsendia. ~ 1. A representative specimen of T. incana. Fig. 2. A a rgnggrents biome T. Fendleri. Fig. 3. A representative specimen of T. strigosa (Holmgren @ Tillet 9474). Fig. 4. The holotype of T. annua (cu). THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 149 BA Pama) ‘ se . , soar Pirate XXIII. Photographs of representative specimens of species of Townsendia. } ; i : T. condensata. Fig. 3. Apomictic T. gr Fig. 1. Townsendia mexicana. Fig. 2. Sexua condensata. Fig. 4. Townsendia spathulata. THE SYSTEMATICS AND EVOLUTION OF TOWNSENDIA 151 INDEX Aplopappus florifer 112 apomixis, causal aspects of, 51-52 divergent evolution 54-59 Erigeron florifer 111 generic relationships 11-13 geographical distribution 9-11 hybridization 61-63 megasporogenesis in apomicts 34-51 in sexual plants 28-34 microsporogenesis in diploids 18 in polyploid apomicts 18-27 morphology 63-6 Nuttall, localities of specimens 7 species concept | Stenotus florifer 112 arizonica 123 arizonica X incana 123 Bakeri 79 condensata 118, map 120 Fendleri 128, map florifer 111, map 115 var iedicats 135, map 120 minima 85 montana 83, map 82 pinetoru othvoke 81, map 82 scapigera 115 var. ambigua 112 var. | 115 sericea var. on 91 Wrightii 137 CONTRIBUTIONS FROM THE GRAY HERBARIUM OF HARVARD UNIVERSITY Edited by Reed C, Rollins and Robert C. Foster NO. CLXXXIV Qj + (313 A CATALOGUE OF THE FERNS AND FLOWERING PLANTS OF BOLIVIA By RoBerT C. FOSTER Published by THE GRAY HERBARIUM OF HARVARD UNIVERSITY CAMBRIDGE, MASS., U.S.A. 1958 Issued June 27th INTRODUCTORY NOTE When I first began to work on Bolivian plants, I hoped to produce a Flora of Bolivia. Unfortunately, inevitable delays and the slowness with which the work has proceeded make the chance of its completion within my lifetime extremely doubtful. Consequently, it seems advisable to publish this catalogue of the ferns and flowering plants of Bolivia. Al- though a complete listing is too much to be hoped for, it is the most nearly complete catalogue of Bolivian plants yet to appear. Additional records are being accumulated and supplements will be published from time to time, whenever the number of additions makes this advisable. Plant families are arranged in the Engler and Prantl sequence. Within families, genera are arranged alphabeti- cally ; within genera, species are cited alphabetically. Italics mean that the name is considered to be a synonym. A ? before a name indicates considerable doubt as to the cor- rectness of the identification. Some idea of the richness of the Bolivian flora can be gained from the fact that 196 families of pteridophytes and phanerogams are included in this list. Although quite con- servative generic concepts have been followed (with few ex- ceptions), 1874 genera are listed here. They are distributed as follows: Pteridophytes 68 genera Gymnosperms 3 genera Monocotyledons 355 genera Dicotyledons 1252 genera Whenever possible, I have followed critical monographs and partial treatments of plant-groups. In some cases, I have been able to make critical judgments based on my own studies. A number of specialists have been kind enough to go over the lists of their special interests. To the following I am much indebted for their assistance: Dr. H. Horn Rantzien (aquatic monocotyledons); Dr. H. K. a (Cyperaceae); Dr. L. B. Smith (Bromeliaceae; Begonia) ; Mr. Irwin F. Lane (Heliconia) ; Mr. Charles Schweinfurth (Orchidaceae) ; Dr. C. E. Wood, Jr. (Chloranthaceae ; Teph- rosia) ; Dr. Alicia Lourteig (Ranunculaceae ; Mayacaceae ‘ pr. °C. Rollins (Cruciferae) ; Dr. B. G. Schubert (Desmo- dium; Begonia); Dr. G. L. Webster (Phyllanthus) ; Dr. C. E. Smith, Jr. (Sloanea) ; Dr. C. E. Kobuski (Theaceae) ; Dr. Lincoln Constance (Umbelliferae); Dr. Rogers Mc- Vaugh (Campanulaceae). In addition, I must express my appreciation to Dr. Martin Cardenas and Dr. H. C. Cutler for specimens sent to me. PTERIDOPHYTA HYMENOPHYLLACEAE Hymenophyllum apicale v. d. B. in Ned. Kr. Arch. 4: 397 (1859). H. Buchtienii Rosenst. in Fedde, Repert. Spec. Nov. 5: 229 (1908) = H. ele- gantulum. H. crispum HBK. Nov. Gen. & Spec. 1: 26 (1816). H. crispum var. brasilianum Fée, Crypt. Vasc. Brés. 1: 195, t. 71, fig. 2 (1869). H. dendritis Rosenst. in Fedde, Repert. apes Nov. 6: 308 (1909). H. elegans Spreng. Syst. Veg. 4: 133 (1827). H. elegantulum v. d. B. in Ned. Kr. Arch. 4: 408 (1859). H. fragile (Hedw.) Morton in Contrib. U. S. Nat. Herb. 29(3): 172 (1947). H. fucoides Sw. in Schrad. Journ. 1800 (2): 99 a 1). H. helicoideum Sod. Crypt. Vasc. Quit. 20 (189 Herzogii Rosenst. in Meded. Rijks Herbar. Ae 5 (1913). hirsutum (L.) Sw. in Schrad. Journ. 1800 (2): 99 (1801). interruptum Kunze in Linnaea, 9: 107 (1834). microcarpum Desv. in Mém. Soe: Linn. Paris, 6: 333 (1827). multiflorum Rosenst. in Meded. Rijks Herbar. on 3 (1913). myriocarpum Hook. Sp. Fil. 1: 106, t. 37D (184 nigrescens var. gracile Rosenst. in Meded. sis Berbae. 19: 4 (1913). Orbignianum v. d. B. in Ned. Kr. Arch. 5(3): 191 (1863) = H. crispum. peltatum (Poir.) Desv. in Mém. Soc. Linn. Paris, 6: 333 (1827). ruvianum Hook. & Grev. Ic. Fil. 2: t. 208 (1831). plumosum Kaulf. Enum. Fil 267 (1824). polyanthos Sw. in Schrad. Journ. 1800 (2): We (1801). protrusum Hook. Sp. Fil. 1: 104, t. 37B (18 pyramidatum Desv. in Mém. Soe. Linn. oe 6: 832 (1827). Ruizianum (K1.) Kunze in Bot. Zeit. 5: 199 (1847). speciosum v. d. B. in Ned. Kr. Arch. 5(3): 181 (1863). spectabile Mett. in Linnaea, 35: 392 (1867-1868) — H. speciosum. tenerrimum v. d. B. in Ned. Kr. Arch. 5(3): 185 (1863). Trianae Hieron. in Engler, Bot. Jahrb. 34: 429 er trichophyllum HBK. Nov. Gen. & Spec. 1: 27 (1816). undulatum var. regenerans C. Chr. in Ark. Bot. 20A (7): 8 (1926). valvatum Hook. & Grev. Ic. Fil. 2: t. 219 (18381). verecundum Morton in Contrib. U. S. Nat. Herb. 29: 183 (1947). richomanes angustatum Carm. in Trans. Linn. Soc. 12: 513 (1818). angustatum var. subexsertum (v. d. B.) C. Chr. in Ark. Bot. 20A (7): 9 (1926). Ankersii Parker ex Hook. & Grev. Ic. Fil. 2: t. 201 Longe crispum L. Sp. Pl. 2: 1097 (1758). cristatum Kaulf. Enum. Fil. 265 (1824). delicatum v. d. B. in Ned. Kr. Arch. 5(2): 145 (1861). diversifrons (Bory) Mett. ex Sadeb. in Engl. & Prantl, Nat. Pflanzenfam. 1(4): 108 (1899 Herzogii Bose. in Meded. Rijks Herbar. rie 5 (1913). Krausii Hook. & Grev. Ic. Fil. 2: t. 149 (18 Kunzeanum Hook. Sp. Fil. 1: 127, t. 39D See membranaceum L. Sp. Pl. 2: 1097 (1753). montanum Hook. Ic. Pl. 2: t. 187 (1837). pinnatum Hedw. Fil. Gen. & Spec. t. 4, fig. 1 (1799). plumosum Kunze in Linnaea, 9: 104 (1834). polypodioides L. Sp. Pl. 2: 1098 (1753). . pyxidiferum L. Sp. Pl. 2: 1098 (1753). aalalatalalatal eS at ad at af tS et af att tt . Rew eee Saas 6S T. radicans Sw. in Schrad. Journ. 1800 (2): 97 (1801). T. rigidum Sw. Prodr. 137 (1788). T. rupestre (Raddi) v. d. B. in Ned. Kr. Arch. 4: 370 (1859). T. Sellowianum Presl, Hym. 15, 37 (1843). T. Trollii Bergdolt in Flora (N. S.) 127: 263 (1933). CYATHEACEAE Alsophila armata (Sw.) Presl, Tent. 62 (1836). A. bulligera Rosenst. in Fedde, Repert. Spec. Nov. 25: 57 (1928). A. lasiosora Mett, ex Kuhn in Linnaea, 36: 157 (1869). A. Lechleri Mett. Fil. Lechl. 2:28 (1859). A. mapiriensis Rosenst. in Fedde, Repert. Spec. Nov. 25: 57 (1928). A. pubescens Baker, Synop. Fil. (ed. 1) 449 (1868). A. quadripinnata (Gmel.) C. Chr. Ind. Fil. 47 (1905). A. rostrata (HBK.) Mart. Ic. Crypt. Bras. 64, t. 39 (1834). A. rufa Fée, Crypt. Vasc. Brés. 166, t. 59, fig. 1 (1869). A. villosa (Humb. & Bonpl.) Desv. in Mém. Soc. Linn. Paris, 6: 319 (1827). Cyathea castanea Baker, Synop. Fil. (ed. 2) 451 (1874). C. cuspidata Kunze in Linnaea, 9: 101 (1834). C. cuspidata var. rigida Rosenst. in Meded. Rijks Herbar. 19: 6 (1913). C. furfuracea Baker, Synop. Fil. (ed. 2) 450 (1874). C. Herzogii Rosenst. in Meded. Rijks Herbar. 19: 7 (1913). C. mexicana var. boliviensis Rosenst. in Fedde, Repert. Spec. Nov. 25: 56 28 ? C. pilosa Baker, Synop. Fil. (ed. 2) 19 (1874). C. Schanschin Mart. Ic. Crypt. Bras. 77, t. 29, fig. 3-4, t. 54 (1834). C. vestita Mart. in Denkschr. Bot. Ges. Regensb. 2: 146 (1822). C. yungensis C. Chr. in Ark. Bot. 20A (7): 10 (1926). Hemitelia grandifolia (Willd.) Spreng. Syst. Veg. 4: 125 (1827). H. subincisa Kunze in Bot. Zeit. 2: 296 (1844). POLYPODIACEAE Adiantopsis radiata (L.) Fée, Gen. 145 (1850-1852). A. ternata Prantl in Gartenfl. 32: 101 (1883). Adiantum Baenitzii Rosenst. in Fedde, Repert. Spec. Nov. 5: 230 (1908). A. boliviense Christ & Rosenst. in Fedde, Repert. Spec. Nov. 5: 230 (1908). : te cuneatum Langsd. & Fisch. Ic. Fil. 23, t 26 (1810). decorum var. quadripinnatum Rosenst. in Meded. Rijks Herbar. 19: 8 (1918). digitatum Presl ex Hook. Sp. Fil. 2: 38: (1851). latifolium Lam. Encye. 1: 43 (1783). Lorentzii Hieron. in Engler, Bot. Jahrb. 22: 393 (1896). macrophyllum Sw. Prodr. 135 (1788). Moorei Baker in Gard. Chron. 1878: 811. obliquum Willd. Sp. Pl. 5: 429 (1810). Orbignyanum Mett. ex Kuhn in Linnaea, 36: 78 (1869). peruvianum Kl. in Linnaea, 18: 555 (1844). petiolatum Desv. in Ges. Naturforsch. Berl. Mag. 5: 326 (1811). platyphyllum Sw. in Kgl. Vet. Akad. Handl. 1817: 74, t. 3, fig. 6. Poiretii Wikstr. in Kgl. Vet. Akad. Handl. 1825: 443 (1826). polyphyllum Willd. Sp. Pl. 5: 454 (1810). pulverulentum L. Sp. Pl. 2: 1096 (1753). A. serrato-dentatum Willd. Sp. Pl. 5: 445 (1810). A. tinctum Moore in Gard. Chron. 1862: 932. Perr rr rr rr rrr Rel 6 Anetium citrifolium (L.) Splitg. in Tijdschr. Nat. Gesch. 7: 895 —_— Antrophyum discoideum Kunze in Bot. Zeit. 6: 702 (1848). A. abscissum var. subaequilaterale Rosenst. in Fedde, Repert. Spec. Nov. 12: 470 (1913) alatum Humb. & Bonpl. ex Willd. Sp. Pl. 5: 319 (1810). auricularium var. acutidens Rosenst. in Fedde, Repert. Spec. Nov. 12: 469 (1913) auricularium var. subintegerrimum (Hieron.) Rosenst. in Fedde, Repert. Spec. Nov. 12: 469 (1913) auriculatum Sw. in Kel. Vet. Akad, Handl. 1817: 68. auritum Sw. in Schrad. Journ. 1800 (2): 52 (1801). auritum var. davallioides Rosenst. in Fedde, Repert. Spec. Nov. 5: 232 (1908). auritum var. davallioides forma diversifolium Rosenst. in Fedde, Repert. Spec. Nov. 5: 282 (1908 Ballivianii Rosenst. in Fedde, Repert. Spec. Nov. 11: 55 (1912). Bangii Hieron. in Hedwigia, 60: 245 (1918). . Bangii Gandoger in Bull. Soc. Bot. France, 66: 305 (1920). castaneum Schlechtd. & Cham. in Linnaea, 5: 611 (1830). cirrhatum Richard ex Willd. Sp. Pl. 5: 321 (1810). Claussenii Hieron. in Hedwigia, 60: 241 (1919). cristatum Lam. Encye. 2: 310 (1786). debile Fée, Mém. 10: 28, t. 35, fig. 2 (1866). depauperatum Fée, Mém. 7: 52, t. 15, fig. 3 (1857). dimidiatum var. boliviense Rosenst. in Fedde, Repert. Spec. Nov. 12: 470 (1913). discrepans Rosenst. in Fedde, Repert. Spec. Nov. 12: 469 (1913). divergens Mett. ex Baker in Mart. Fl. Bras. 1(2): 445 (1870). erosum L. Syst. (ed. 10) 2: 13824 (1759). fluminense Hieron. in Hedwigia, 61: 17 (1919). formosum Willd. Sp. Pl. 5: 329 (1810). fragile Presl, Tent. 108 (1836). fragrans Sw. Prodr. 130 (1788). fragrans var. foeniculaceum (HBK.) Hook. Sp. Fil. 3: 181 (1860). Gilliesii Hook. Exot. Fl. 3: ad t. 208 (1827). harpeodes Kunze in Linnaea, 18: 329 (1844). harpeodes var. incisum (Rosenst.) Hieron. in Hedwigia, 60: 238 (1918). hastatum Kl. ex Kunze in Linnaea, 23: 235, 305 (1850). Herzogii Rosenst. in Meded. Rijks Herbar. 19: 12 (1913). integerrimum Spreng. in Nov. Act. Acad. Caes. Leop. 10: 231 (1821). laetum Sw. Syn. 79, 271 (1806). . Lorentzii Hieron. in Engler, Bot. Jahrb. 22: 375 (1896). . lunulatum Sw. in Schrad. Journ. 1800 (2): 52 (1801). - monanthes L. Mant. 130 (1767 monanthes var. yungense Rosenst, in Fedde, Repert. Spec. Nov. 11: 54 12). RAGABAEADE AAAS ties cate > pb>> p> b> obtusifolium L. Sp. Pl. 2: 1080 (1753). partitum (KI.) C. Chr. Ind. Fil. 125 (1905). poloénse Rosenst. in Fedde, Repert. Spec. Nov. 12: 469 (1918). praemorsum Sw. Prodr. 130 (1788). radicans L. Syst. (ed. 10) 2: 1823 (1759). recumbens Gandoger in Bull. Soc. Bot. France; 66: 305 (1920). resiliens Kunze in Linnaea, 18: 331 (1844). PPP bb b> A. rigidum Sw. in Kgl. Vet. Akad. Handl. 1817: 68. A. Rusbyanum Domin, Pterid. Domin. 171 (1929). A. rutaceum (Willd.) Mett. Aspl. 129, no. 98 (1859). A. sanguinolentum Kunze ex Mett. Aspl. 98, no. S t. 4, fig. 10 (1859). A. serra Langsd. & Fisch. Ic. Fil. 16, t. 19 (1810 A. serratum L. Sp. Pl. 2: 1079 (1753). A. serratum var. caudatum Rosenst. in Fedde, Repert. Spec. Nov. 6: 310 (1909). A. sessilifolium Desv. in Ges. Naturforsch. Berl. Mag. 5: 322 (1811). A. sessilifolium var. minus Hieron. in Engler, Bot. Jahrb. 34: 461 (1904). A. spinescens Mett. ex Kuhn in Linnaea, 36: 98 (1869). . A. squamosum L, Sp. Pl. 2: 1082 (1753). A. tocoraniense Rosenst. in Meded. Rijks Herbar. 19: 11 (1913). A. tricholepis Rosenst. in Fedde, Repert. Spec. Nov. 12: 468 (1913). A. trilobatum C. Chr. in Ark. Bot. 20A(7): 15, fig. 1 (1926). A. triphyllum Presl, Rel. Haenk. 1: 45 (1825). A. triphyllum var. compactum Hook. Sp. Fil. 3: 203 (1860). A. uniseriale Raddi, Opusc. Sci. Bologn. 3: 291 (1819). ? Athyrium Dombeyi Desv. in Mém. Soc. Linn. Paris, 6: 266 (1827). Blechnum auriculatum Cav. Descr. 262 (1802). B. blechnoides (Lag.) C. Chr. Ind. Fil. 151 (1905). B. blechnoides var. gracilipes Rosenst. in Fedde, Repert. Spec. Nov. 9: 343 B. brasiliense Desv. in Ges. Naturforsch. Berl. Mag. 5: 330 (1811). B. Buchtienii Rosenst. in Fedde, ag Spec. Nov. 5: 231 (1908). B. caudatum Cav. Descr. 262 (1802) B. chilense (Kaulf.) Mett. Fil. Bee, 1: 14 (1856). B. delicatum Maxon & Morton in Bull. Torr. Bot. Club, 66: 41 (1939). B. fraxineum Willd. Sp. Pl. 5: 413 (1810). B. glandulosum Link, Enum. Alt. 2: 462 (1822). B. glandulosum var. distans (Presl) C. Chr. in Ark. Bot. 20A (7): 17 (1926). B. gracile Kaulf. Enum. Fil. 158 (1824). B. Kunthianum C, Chr. Ind. Fil. Suppl. 1906-1912: 16 (1913). B. lanceola Sw. in Kgl. Vet. Akad. Handl. 1817: 71, t. 3, fig. 2. B. lima Rosenst. in Fedde, Repert. Spec. Nov. 11: 53 Soop B. loxense Hook. ex Salomon, Nomencl. Gefasskr. 117 (18 B. malacothrix Maxon & Morton in Bull. Torr. Bot. Club, < “40 (1939). B. nigro-squamatum Gilbert in Bull. Torr. Bot. Club, 24: 258 (1897). B. occidentale L. Sp. Pl. 2: 1077 (1758). B. penna-marina (Poir.) Kuhn, Fil. Afr. 92 (1868). B. penna-marina var. bolivianum Rosenst. ex Looser in Rev. Univ. Catol. Chile, 32: 77 (1948). ? B. Raddianum Rosenst. in ra 46: 91 (1906). B. Sprucei C. Chr. Ind. Fil. 160 (1905). B. subtile Rosenst. in Fedde, Repert. Spec. Nov. 11: 54 (1912). B. unilaterale Sw. in Ges. Naturforsch. Berl. Mag. 4: 79, t. 3, (1810). B. volubile Kaulf, Enum. Fil. 159 (1824). Bolbitis crenata (Presl) C. Chr. Ind. Fil. Suppl. 3: 47 (1934). B. serratifolia (Martens) Schott, Gen. Fil. t. 13 (1834). B. Stuebelii (Hieron.) C. Chr. Ind. Fil. Suppl. 3: 50 (1934). Cheilanthes boliviana C. Chr. in Ark. Bot. 20A (7): 19 (1926). elegans Desv. in Ges. Naturforsch. Berl. Mag. 5: 328 (1811). Hieronymi Herter in Anal. Mus. Nac. Montevideo (ser. 2) 1: 360 (1925). marginata HBK. Nov. Gen. & Spec. 1: 22 (1816). myriophylla Desv. in Ges. Naturforsch. Berl. hog 5: 328 (1811). obducta Mett. ex Kuhn in Linnaea, 36: 83 (1869). Pilosa Goldm. in Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 455 (1843). AAARAAA 8 C. Poeppigiana Mett. ex Kuhn in Linnaea, 36: 84 (1869). C. pruinata Kaulf. Enum. Fil. 210 (1824). C. rufo-punctata Rosenst. in Meded. Rijks Herbar. 19: 9 (1913). Cyclopeltis semicordata (Sw.) J. Sm. in Bot. Mag. 72: Comp. 36 (1846). Cystopteris fragilis (L.) Bernh. in Schrad. Neu. Journ. 1(2): 27, t. 2 fig. 4 1806 Dennstaedtia adiantoides (Humb. & Bonpl.) Moore, Ind. XCVII (1857). D. bipinnata (Cav.) Maxon in Proc. Biol. Soc. Wash. 51: 39 (1938). D. cornuta (Kaulf.) Mett. in Ann. Sci. Nat. (ser. 5) 2: 260 (1864). D. D’Orbignyana Kuhn in Linnaea, 36: 146 (1869). D. glauca (Cav.) C. Chr. ex Looser in Rev. Hist. Geog. Chile, 69: 184 (1982). D. Mathewsii (Hook.) C. Chr. Ind. Fil. 218 (1905). D. rubiginosa (Kaulf.) Moore, Ind. XCVII (1857). Didymochlaena truncatula (Sw.) J. Sm. in Journ. Bot. 4: 196 (1841). Diplazium ambiguum Raddi, Opusce. Sci. Bologn. 3: 292 (1819). D. Ballivianii Rosenst. in Fedde, Repert. Spec. Nov. 6: 311 (1909). D. Buchtienii Rosenst. in Fedde, Repert. Spec. Nov. 6: 312 (1909). D. cristatum (Desr.) Alston in Journ. Bot. 74: 173 (1936). D. cuneifolium Rosenst. in Fedde, Repert. Spec. Nov. 12: 470 (1918). D. delitescens Maxon in Contrib. U. S. Nat. Herb. 10: 497 (1908). D. divergens Rosenst. in Fedde, Repert. Spec. Nov. 12: 471 (1913). D. Lindbergii (Mett.) Christ in Pittier, Primit. F]. Costar. 3: 27 (1901). D. mapiriense Rosenst. in Fedde, Repert. Spec. Nov. 6: 310 (1909). D. marginatum (L.) Diels in Engl. & Prantl, Nat. Pflanzenfam. 1(4): 229 1899). D. pinnatifidum Kunze in gee) 9: 72 (1834). D. plantaginifolium (L.) Urb. Symb. Ant. 4: 31 (1908). D. yungense Christ & Bosensé, in Fedde, Repert. Spec. Nov. 5: 233 (1908). Doryopteris concolor (Langsd. & Fisch.) Kuhn in v. Deck. Reis. 3(3): Bot. 19 crenulans (Fée) Christ in Bull. Herb. Boiss. (ser. 2) 2: 549 (1902). Lorentzii (Hieron.) Diels in Engl. & Prantl, Nat. Pflanzenfam. 1(4): 270 (1899). nobilis (Moore) Baker, Synop. Fil. (ed. 2) 167 (1874), in synon. ornithopus (Mett.) J. Sm. Hist. Fil. 289 (1875). pedata var, multipartita (Fée) Tryon in Contrib. Gray Herb. 143: 38 (1942). pedata var. palmata (Willd.) Hicken in Rev. Mus. La Plata, 15: 253 (1908). ryopteris ampla (Humb. & Bonpl.) O. Ktze. Rev. Gen. 2: 812 (1891). amplissima (Presl) O. Ktze. Rev. Gen. 2: 812 (1891). ina Morton in Journ. Wash. Acad. Sci. 28: 526 (1988). angustifolia (Willd.) Urb. Symb. Ant, 4: 21 (1903) argentina (Hieron.) C. Chr. Ind. Fil. 253 (1905). aspidioides (Willd.) C. Chr. Ind. Fil. 253 (1905). Bangii C. Chr. in Dansk. Vidensk. Selsk. Skr. (ser. 7) 4: 333 (1907). boliviensis Morton in Journ. Wash. Acad. Sci. 28: 527 (1938). Canadasii (Sod.) C. Chr. Ind. Fil. 256 (1905). catocarpa (Kunze) O. Ktze. Rev. Gen. 2: 812 (1891) — D. nemophila. cheilanthoides (Fée) C. Chr. Ind. Fil. 257 (1905). coarctata (Kunze) C. Chr. Ind. Fil. 258 (1905). concinna (Willd.) O. Ktze. Rev. Gen. 2: 812 (1891). contermina ( Willd.) O. Ktze. Rev. Gen. 2: 812 (1891). dentata (Forsk.) C. Chr. in Dansk. Vidensk. Selsk. Skr. (ser. 8) 6: 24 (1920). denticulata forma boliviensis C. Chr. in Dansk. Vidensk. Selsk. Skr. (ser. 8) 6: 116 (1920). Desvauxii Maxon & Morton in Bull. Torr. Bot. Club, 65: 369 (1938). Ye ee SP SSSSSsssyssssssgsgys o AAA nd P Pewee’ FP Pe. SF. PP bP SPyeoepeeD os SSose . 9 "ers tie forma glandulosa Maxon & Morton in Bull. Torr. Bot. Club, 65: 372 (1938). diplazioides (Desv.) Urb. Symb. Ant. 4: 21 (1903 diplazioides var. chacoénsis C. Chr. in Re Bot. Fi (7): 12 (1926). effusa (Sw.) Urb. Symb. Ant. 4: 16 (1903). Guentheri Rosenst. in Fedde, Repert. Spec. Nov. 25: . (1928). Herzogii Rosenst. in Meded. "Rijks Herbar. 19: 15 (1913). honesta (Kunze) C. Chr. Ind. Fil. 271 (1905). Jurgensenii (Fée) Maxon & Morton in Bull. Torr. Bot. Club, 65: 360 (1938). Leprieuri (Hook.) O. Ktze. Rev. Gen. 2: 813 (1891). leucothrix C. Chr. in Smithsonian Misc. Coll. (new series) 5: 377 (1909). leucothrix oe glanduligera C. Chr. & Rosenst. in Fedde, Repert. Spec. Nov. 12: 471 (19 macrotis var. eas Sica Rosenst. in Fedde, Repert. Spec. Nov. 7: 298 (1909). mapiriensis Rosenst. in Fedde, Repert. Spec. Nov. 6: 313 (1909). meniscioides var. conferta (Kaulf.) Morton in Bull. Torr. Bot. Club, 66: 51 1939). nemophila (Kunze) C. Chr. in Dansk. Vidensk. Selsk. Skr. (Ser. 8) 6: 57 (1920). nephrodioides var. glandulosa C. Chr. & Rosenst. in Fedde, Repert. Spec. Nov. 12: 473 (19138) nervosa (Kl.) C. Chr. Ind. Fil. 279 (1905). oligocarpa (Humb. & Bonpl.) O. Ktze. Rev. Gen. 3(2): 378 (1898). oligophylla var. aequatorialis C. Chr. in Dansk. Vidensk. Selsk. Skr. (ser. 7) 10(2): 189 (1913). oligophylla var. pallescens C. Chr. in Dansk. Vidensk. Selsk. Skr. (ser. 7) 10(2): 188 (1913). opposita var. furcativenia Rosenst. in Fedde, Repert. Spec. Nov. 6: 313 ). paleacea (Sw.) C. Chr. in Amer. Fern pees 1: 94 (1911). patens (Sw.) O. Ktze. Rev. Gen. 2: 813 (1891). patula (Sw.) Underw. Our Native Ferns (ed. 4), 117 (1893). Pavoniana (KI.) C. Chr. Ind. Fil. 283 (1905). permollis Maxon & Morton in Bull. Torr. Bot. Club, 65: 372 (1938). phacelothrix C. Chr. & Rosenst. in Fedde, Repert. Spec. Nov. 11: 56 (1912). piloso-hispida (Hook.) C. Chr. in Dansk. Vidensk. Selsk. Skr. (ser. 7) 10(2): 148 (1913). ptarmiciformis C. Chr. & Rosenst. in Fedde, Repert. Spec. Nov. 12: 472 (1918). pterifolia (Mett.) O. Ktze. Rev. Gen. 2: 813 (1891). resinosofoetida (Hook.) O. Ktze. Rev. Gen. 2: 813 (1891). rivulariformis Rosenst. in Fedde, Repert. Spec. Nov. 6: 316 (1909). rivularioides (Fée) C. Chr. in Hedwigia, 46: 125 (1906). Rosei Maxon in Smithsonian Misc. Coll. 65(8): 10 (1915). rotundata (Willd.) C. Chr. Ind. Fil. 289 a. 905). rudis (Kunze) C, Chr. Ind, Fil. 289 (190 Ruiziana (Kl.) C. Chr. in Dansk. ata Selsk. Skr. (ser. 7) 10(2): 152 913). Rusbyi C. Chr. in Smithsonian Misc. Coll. (new series) 5: 390 (1909). Rusbyi var. major Rosenst. in Fedde, Repert. Spec. Nov. 25: 59 (1928). Salzmannii (Fée) Maxon & Morton in Bull. Torr. Bot. Club, 65: 357 (1938). serrata (Cav.) C. Chr. Ind. Fil. 291 (1905). Standleyi Maxon & Morton in Bull. Torr. Bot. Club, 65: 368 (1938). stenophylla Rosenst. in Fedde, Repert. Spec. Nov. 5: 233 (1908) = D. rivulariformis. subandina C. Chr. & Rosenst. in Fedde, Repert. Spec. Nov. 12: 472 (1913). subincisa (Willd.) Urb. Symb. Ant. 4: 19 (1903) submarginalis (Langsd. & Fisch.) C. Chr. Ind. Fil. 296 (1905). Tatei Maxon & Morton in Journ, Wash. Acad. Sci. 28: 529 (1938) tristis (Kunze) O. Ktze. Rev. Gen. 2: 814 (1891). tristis va auriculata C. Chr. & Rosenst. in Fedde, Repert. Spec. Nov. 12: 471 (19 vasta aca Hieron. in Hedwigia, 46: 347 (1907). villosa var. inaequalis Gilbert in Bull. Torr. Bot. Club, 24: 259 (1897). villosula C. Chr. in Dansk. Vidensk. Selsk. Skr. (ser. 8) 6: 89 (1920). yungensis Christ & Rosenst. in Fedde, Repert. Spec. Nov. 5: 234 (1908). aphoglossum accedens (Mett.) Christ, Monog. Elaph. 91 (1899). actinotrichum (Mart.) Moore, Ind. 4 (1857). acutifolium Rosenst. in Fedde, Repert. Spec. Nov. 25: 60 (1928). angustissimum (Fée) C. Chr. in Ark. Bot. 20A(7): 25 (1926). angustissimum var. minus C. Chr. in Ark. Bot. 20A(7): 25 (1926). . Ballivian Rosenst. in Fedde, Repert. Spec. Nov. 12: 474 (1918) [as E. Bol- livian . Bangii ‘Ghiiet; Monog. Elaph. 99 (1899). . barbatum (Karst.) Hieron. in Engler, Bot. Jahrb. 34: 553 (1904). . blandum Rosenst. in Fedde, Repert. Spec. Nov. 12: 476 (1913). . Brausei Rosenst. in Fedde, Repert. Spec. Nov. 12: 475 (1913). . Buchtienii Rosenst. in Fedde, Repert. Spec. Nov. 12: 475 (1913). . Cardenasii Wagner in Bull. Torr. Bot. Club, 81: 62 (1954). casanense Rosenst. in Fedde, Repert. Spec. Nov. 25: 64 (1928). cinctum Rosenst. in Fedde, Repert. Spec. Nov. 25: 63 (1928). cordifolium Rosenst. in Fedde, Repert. Spec. Nov. 25: 62 (1928). erassipes (Hieron.) Diels in Engl. & Prantl, Nat. Pflanzenfam. 1(4): 334 (1899). cuspidatum (Willd.) Moore, Ind. XVI (1857). . decoratum (Kunze) Moore, Ind. 8 (1857). . didymoglossoides C. Chr. in Bot. Tidsskr. 26: 299 (1904). -. Dombeyanum (Fée) Mocca’ & Houlst. in Gard. Mag. Bot. 3: 95 ( 1851). . Eatonianum (E. G. Britt.) C. Chr. Ind. Fil. 306 (1905). . Engelii (Karst.) Christ, Monog. Elaph. 81 (1899). . Engelii var. subnudum Rosenst. in Fedde, Repert. Spec. Nov. 11: 59 (1912). erinaceum var. boliviense Rosenst. in Fedde, Repert. Spec. Nov. 12: 476 (19138). . filipes Rosenst. in Fedde, Repert. Spec. Nov. 25: 62 (1928). . Gayanum (Fée) Moore, Ind. 10 (1857). . glabellum J. Sm. in Hook. London Journ. Bot. 1: 197 (1842). . glaucescens Rosenst. in Fedde, Repert. Spec. Nov. 25: 61 (1928). . Guentheri Rosenst. in Fedde, Repert. Spec. Nov. 25: 61 (1928). . horridulum (Kaulf.) J. Sm. Bot. Voy. Herald, 232 (1854). . Huaessaro (Ruiz) Christ, Monog. Elaph. 96 (1899). interruptum Rosenst. in Fedde, Repert. Spec. on 12: 474 (1913). Jamesonii (Hook. & Grev.) Moore, Ind. 10 (1857). . laxepaleaceum Rosenst. in Fedde, Repert. Spec. Nov. 11: 59 (1912). oe var. truncatum Rosenst. in Fedde, Repert. Spec. Nov. 25: 63 linguaeforme (Cav.) Moore, Ind. 11 (1857). - Mandonii (Mett.) Christ, Motiog. ia 128 (1899). Mathewsii (Fée) Moore, Ind. 12 (1857). meridense var. boliviense Rosenst: in Fedde, Repert. Spec. Nov. 25: 63 ( 1928). micropus Rosenst. in Fedde, Repert. Spec. Nov. 6: 316 (1909). micropus var. majus Rosenst. in Fedde, Repert. Spec. Nov. 6: 316 (1909). apenas aa aaaaialslals oa ed ad PPYSPY & 2) be bed ted bet ed bd ad dt ds tt notatum (Fée) Moore, Ind. 12 (1857). Orbignyanum (Fée) Moore, Ind. XVI (1857). Orbignyanum var. tectiforme Rosenst. in Fedde, Repert. Spec. Nov. 12: 477 ovalifolium (Fée) Christ in Bull. Herb. Boiss. (ser. 2) 2: 361 (1902). E. perelegans (Fée) Moore, Ind. XVI (1857). petiolatum (Sw.) Urb. Symb. Ant. 4: 61 (1903). piloselloides (Presl) Moore, Ind. 13 (1857). praelongum (Fée) C. Chr. in Dansk Bot. Ark. 9(3): 8 (1987). productum Rosenst. in Fedde. Repert. Spec. Nov. 6: 315 (1909). pseudohirtum Rosenst. in Meded. Rijks Herbar. 19: 23 (1913). scolopendrifolium (Raddi) J. Sm. in Bot. Mag. 72: Comp. 17 (1846). squamipes (Hook.) Moore, Ind. 15 (1857). subarborescens var. bolivianum Rosenst. in Meded. Rijks Herbar. 19: 23 (1913). tectum (Humb. & Bonpl.) Moore, Ind. 15 (1857). tenuiculum (Fée) Moore, Ind. 15 (1857). unduaviense Rosenst. in Fedde, Repert. Spec. Nov. 11: 58 (1912). unduaviense var. leptophylloides Rosenst. in Fedde, Repert. Spec. Nov. 11: 58 (1912). . Wacketii Rosenst. in Hedwigia, 46: 151 (1907). Eschatogramme Desvauxii (Kl.) C. Chr. in Dansk Bot. Ark. 6(3): 37 werent E. E. furcata var. subnuda C. Chr. in Dansk Bot. Ark. 6(3): 36 (1929). panamensis C. Chr. in Dansk Bot. Ark. 6(3): 37 (1929). Gymnogramma Ballivianii Rosenst. in Fedde, Repert. Spec. Nov. 6: 314 (1909). QAARANAA aH elongata Hook. & Grev. in Journ. Bot. 1: 61, t. 119 (1834). flexuosa (Humb. & Bonpl.) Desv. in Ges. Naturforsch. Berl. Mag. 5: 306 (1811). Herzogii Rosenst. in Meded. Rijks Herbar. 19: 21 (1913). Hookeri J. Sm. ex Baker, Synop. Fil. (ed. 1) 381 (1868). Orbignyana Mett. ex Kuhn in Linnaea, 36: 70 (1869). prehensibilis Baker, Synop. Fil. (ed. 2) 517 (1874). . scandens (Fée) Baker in Journ. Linn. Soc. 14: 25 (1873). Gymnopteris tomentosa (Lam.) Underw. in Bull. Torr. Bot. Club, 29: 627 (1902). Hemionitis palmata L. Sp. Pl. 2: 1077 (1753). Histiopteris incisa (Thunb.) J. Sm. Hist. Fil. 295 (1875). Hypolepis Buchtienii Rosenst. in Fedde, Repert. Spec. Nov. 25: 58 (1928). H. viscosa Karst. Fl. Colomb. 2: 89, tt. 145, 146 (1869). ? Jamesonia glutinosa Karst. Fl. Colomb, 2: 85, t. 143 (1869). J. J. scalaris Kunze in Bot. Zeit. 2: 739 (1844). verticalis var. humilis Karst. Fl. Colomb. 2: 86 (1869). Lindsaea horizontalis Hook. Sp. Fil. 1: 214, t. 62B (1846). L. lancea (L.) Bedd. Ferns Brit. India, Suppl. 6 (1876). L. rigescens Willd. Sp. Pl. 5: 421 (1810). L. stricta (Sw.) Dry. in Trans. Linn. aoe 8: 42 (1797). Lonchitis hirsuta L. Sp. Pl. 2: 1078 (175 Microlepia speluncae (L.) pric Ind. emt (1857). Microstaphyla Moorei (E. G. Britt.) Underw. in Torreya, 5: 88 (1905). Nephrolepis biserrata (Sw.) Schott, Gen. Fil. ad t. 3 (1834). N. N. mH cordifolia L.) Presl, Tent. 79 (1836). exaltata (L.) Schott, Gen. Fil. t. 3 (1834). pectinata (Willd.) Schott, Gen. Fil. sub t. 3 (1834). N. pendula (Raddi) J. Sm. in Journ. Bot. 4: 197 (1841). Notholsens aurea (Poir.) Desv. in Mém. Soe. Linn. Paris, 6: 219 (1827). N. N. bonariensis (Willd.) C. Chr. Ind. Fil. 6 (1905). Buchtienii Rosenst. in Fedde, Repert. Spec. Nov. 5: 238 (1908). N. Fraseri (Mett.) Baker, Synop. Fil. (ed. 2) 514 (1874). N. Herzogii Rosenst. in Fedde, Repert. Spec. Nov. 6: 175 (1908) = N. obducta. N. nivea (Poir.) Desv. in Journ. de Bot. tay a 93 (1813). N. nivea var. flava Hook. Sp. Fil. 5: 112 (186 N. obducta (Mett.) Baker, Synop. Fil. (ed. “ ie (1874). N. sinuata (Lag.) Kaulf. ‘Enum. Fil. 185 (1824). N. squamosa (Gill.) Lowe, Ferns, 1: t. 17B (1856). Oleandr. Paesia sp. Palinied sagittata (Cav.) Link, Fil. Sp. 60 (1841). P. tenera (Gill.) Prantl] in Engler; Bot. Jahrb. 3: 417 (1882). P. ternifolia (Cav.) Link, Fil. Sp. 59 (1841). P. ternifolia var. petiolulata C. Chr. Ark. Bot. 20A (7): 18 (1926). P. Weddelliana Fée, Mém. 8: 74 (185 Pityrogramma adiantoides ( Karst.) Sis in Publ. Fac. Sci. Univ. Charles, 88: (1928). P. austroamericana Domin in Publ. Fac. Sci. Univ. Charles, 88: 7 (1928). P. Ballivianii (Rosenst.) Domin in Publ. Fac. Sci. Univ. Charles, 88: 10 (1928). P. calomelanos (L.) Link, Hand. Gewachs. 3: 20 (1833 P. calomelanos var. aureo-flava (Hook.) Weath. ex Bailey, Man. Cult. Pls. 64 1924). P. ochracea (Presl) Domin in Publ. Fac. Sci. Univ. Univ. Charles, 88: 8 (1928). P. Ornithopteris (K].) Maxon ex Knuth in Fedde, Repert. Spec. Nov. Beih. 43: (1926). P. tartarea (Cav.) Maxon in Contrib. U. S. Nat. Herb. 17: 173 (1913). Plagiogyria denticulata Copel. in Philipp. Journ. Sci. 38: 412 (1929). P. semicordata (Presl) Christ, Farnkr. 176 (1897). Platycerium andinum Baker in Ann. Bot. 5: 496 (1891). Polybotrya caudata Kunze in Linnaea, 9: 23 (1834). P. cervina (L.) Kaulf. Enum. Fil. 55 (1824). Polypodium allosuroides Rosenst. in Meded. Rijks Herbar. 19: 16 (1913). P. angustifolium Sw. Prodr. 130 (1788) P. angustipaleatum Alston in Journ. Bot. 77: 346 (1989). P. adpressum Copeland in Univ. Calif. Publ. Bot. 19: 305 (1941). P. Asplundii C. Chr. in Ark. Bot. 20A(7): 24 (126). P. aureum L. Sp. Pl. 2: 1087 (17538). P. aureum ea areolatum (Humb. & Bonpl.) Baker in Mart. Fl. Bras. 1(2): 528 (1870). P. Ballivianii Rosenst. in Fedde, Repert. Spec. Nov. 9: 344 (1911). P. Bangii Baker in Kew Bull. 1901: 145. P. bolivianum Rosenst. in Fedde, Repert. Spec. Nov. 5: 236 (1908). P; bolivianum var. brevipes Rosenst. in Fedde, Repert. Spec. Nov. 12: 478 (1913). P. brevifolium Link, Hort. Berol. 2: 90 (1833). bryopodum Maxon in Contrib. U. S. Nat. Herb. 17: 568 (1916). Buchtienii C. Chr. & Rosenst. in Fedde. Repert. Spec. Nov. 5: 287 (1908). Caceresii Sod. Crypt. Vasc. Quit. 360 (1893). choquetangense Rosenst. in Meded. Rijks Herbar. 19: 18 (1913). chrysolepis Hook. Ic. Pl. 8: t. 721 (1848). ciliatum Willd. Sp. Pl. 5: 144 (1810). ciliolepis C. Chr. in Ark. Bot. 20A(7): 21 (1926). cinerascens Lindm. in Ark. Bot. 1: 238 (1903). ircinatum Sod. Crypt. Vasc. Quit. 333 (1893). crassifolium L. Sp. Pl. 2: 1083 (1758). crassifolium var. longipes Rosenst. in Fedde, Repert. Spec. Nov. 11: 58 (1912). Pow MUU ic) DOU UUM NNN See Te OOO es tO-t0 es DOU UU UNO oD x=) crystalloneuron Rosenst. in Fedde, Repert. Spec. Nov. 11; 57 (1912). curvatum Sw. in Schrad. Journ. 1800(2): 24 (1801). decumanum Willd. Sp. Pl. 5: 170 (1810). duale Maxon in Contrib. U. S. Nat. Herb. 16: 61 (1912). filicula Kaulf. Enum. Fil. 275 (1824). fraxinifolium Jacq. Collect. 3: 187 (1789). fulgens Hieron. in Hedwigia, 48: 268 (1909). fuscopunctatum Hook. Sp. Fil. 5: 69, t. 285A (1864). Gilliesii C. Chr. Ind, Fil. 529 (1906). glaucophyllum Kunze ex KI. in Linnaea, 20: 393 (1847). Herzogii Rosenst. in Fedde, Repert. Spec. Nov. 6: es (1908). hirsutissimum Raddi, Opusc. Se. Bologn. 3: 286 (1819). lachniferum var. glabrescens Rosenst. in Fedde, Repert. Spec. Nov. 11: 57 (1912). lanceolatum L. Sp. Pl. 2: 1082 (1758). lanigerum Desv. in Ges. Naturforsch. Berl. Mag. 5: 316 (18 latevagans Maxon & C. Chr. in Proc. Biol. Soc. Wash. aa ge (1939). latipes Langsd. & Fisch. Ic. Fil. 10, t. 10 (1810). latum (Moore) Sod. Crypt. Vasc. Quit. 371 (1893). P. laxifrons Liebm. in Dansk. Vidensk. Selsk. Skr. (ser. 5) 1: 204 (1849). laxum Presl, Rel. Haenk. 1: 23 (1825). leptophyllum L. Sp. Pl. 2: 1092 (1753). leucatomos Poir. in Lam. Encyc. 5: 516 (1804). leuconeuron (Fée) Christ in Schwacke, Pl. Nov. Mineiras, 2: 22 (1900). leuconeuron var. augustifolium Rosenst. in Fedde, Repert. Spec. Nov. 11: 58 (1912 leuconeuron var. latifolium Rosenst. in Fedde, Repert. Spec. Nov. 11: 58 1912) leucosticton Kunze ex KI]. in Linnaea, 20: 380 (1847). longum C. Chr. Ind. Fil. 541 (1906). Lorentzii Hieron. in Engler, Bot. Jahrb. 22: 406 (1896). loriceum L. Sp. Pl. 2: 1086 (1758). loriceum var. hirto-pubescens Hieron. in Hedwigia, 48: 263 (1909). lycopodioides L. Sp. Pl. 2: 1082 (1753). Mathewsii Kunze ex Mett. Pol. 74, no. 112 (1857). megalolepis Maxon & Morton in Bull. Torr. Bot. Club, 66: 39 (1939). meniscifolium Langsd. & Fisch. Ic. Fil. 11, t. 12 (1810). moniliforme Lag. ex Sw. Syn. 33 (1806) myriophyllum Mett. Fil. Lechl. 1: 6 (1856). nigrolimbatum (Spruce) Jenm. in Bull. Bot. Dept. Jamaica, 4: 69 (1897). nitidissimum var. latius Rosenst. in Fedde, Repert. Spec. Nov. 12: 474 (1913). paradiseae Langsd. & Fisch. Ic. Fil. 11, t. 11 (1810). pectinatum L. Sp. Pl. 2: 1085 (1753). pendulum Sw. Prodr, 131 (1788). endulum var. boliviense Rosenst. in Fedde, Repert. Spec. Nov. 25: 60 (1928). percussum Cav. Descr. 243 (1802). persicariifolium Schrad. in Gétt. Gelehrt. Anz. 1824: 867. peruvianum Desv. in Mém. Soc. Linn. Paris, 6: 231 (1827). peruvianum var. subgibbosum Rosenst. in Meded. Rijks Herbar. 19: 16 (1913). phyllitidis L. Sp. Pl. 2: 1083 (1753). P. piloselloides L. Sp. Pl. 2: 1083 (1753). plumula Humb. & Bonpl. ex Willd. Sp. Pl. 5: 178 (1810). poloénse Rosenst. in Fedde, Repert. Spec. Nov. 12: 473 (1913). . polypodioides (L.) Watt in Canad. Nat. (new ser.) 3: 158 (1866). 14 P. polypodioides var. Burchellii (Baker) Weath. in Contrib. Gray Herb. 124: 29 (1939). P. Preslianum var. immersum Rosenst. in Fedde, Repert. Spec. Nov. 6: 314 ? P. prominulum C. Chr. Ind. Fil. Suppl. 1906-1912: 61 (1913). P. pseudocapillare Rosenst. in Meded. Rijks Herbar. 19: 17 (1913). P. pyenocarpum C. Chr. Ind. Fil. 557 (1906). P. repens Aubl. Hist. Pl. Guian. 2: 962 (1775). P. rhizocaulon var. hirsutulum Rosenst. in Fedde, Repert. Spec. Nov. 12: 473 (1913). P. Rusbyi Maxon in Contrib. U. S. Nat. Herb. 17: 570 (1916). P. semihirsutum K1. in Linnaea, 20: 379 (1847 P. senile var. minus Rosenst. in Meded. err i aahinas 19: 19 (1913). P. sericeo-lanatum Hook. Sp. Fil. 4: 221 (1864). P. simacense Rosenst. in Fedde, Repert. Spec. Nov. but 60 (1928). P. sphenodes Kunze ex Kl. in Linnaea, 20: 402 (1847). P. squamulosum Kaulf. Enum. Fil. 89 (1824). P. subflabelliforme Rosenst. in Fedde, Repert. Spec. Nov. 7: 306 (1909). P. subvestitum Maxon in Contrib. U. S. Nat. Herb. 17: 566 (1916). P. tectum Kaulf. Enum. Fil. 87 (1824). P. thyssanolepis A. Br. ex Kl. in Linnaea, 20: 392 (1847). P. triseriale Sw. in Schrad. Journ. 1800(2): 26 (1801). P. truncatulum Rosenst. in Fedde, Repert. Spec. Nov. 9: 343 (1911). P, Tweedianum Hook. Ic. Pl. 1: t. 86 (1887). P. vacciniifolium Langsd. & Fisch. Ic. Fil. 8, t. 7 (1810). P. vittariiforme Rosenst. in Fedde, Repert. Spec. Nov. 5: 235 (1908). P. Williamsii Maxon in Contrib. U. S. Nat. Herb. 17: 547 (1916). P. xanthotrichium Kl. in Linnaea, 20: 376 (1847). P. yungense Rosenst. in Fedde, Repert. Spec. Nov. 5: 236 a, Polystichum adiantiforme (Forst.) J. Sm. Hist. Fil. 220 (187 dubium (Karst.) Diels in Engl. & Prantl, Nat. Lesbeiedbenia 1(4): 194 (1899). gelidum (Kunze) Fée, Gen. 278 (1850-1852). montevidense (Spreng.) Rosenst. in Hedwigia, 46: 111 (1906). muricatum Fée, Gen. 278 (1850-1852). nudicaule Rosenst. in Fedde, Repert. Spec. Nov. 11: 56 (1912). orbiculatum (Desv.) Gay, Fl. Chile, 6: 515 (1853). Pflanzii Hieron. in Engler, Bot. Jahrb. 49: 179 (1913). platyphyllum (Willd.) Presl, Tent. 84 (1836). polyphyllum Presl, Tent. 83 (1836). pycnolepis (Kunze) Hieron. in Engler, Bot. Jahrb. 34: 452 (1904). pygmaeum Copeland in Univ. Calif. Publ. Bot. 19: 300 (1941). rigidum (Hook. & Grev.) C. Chr. in Ark. Bot. 20A(7): 13 (1926). Stuebelii Hieron. in Hedwigia, 46: 355 (1907). Wolfii Hieron. in Hedwigia, 46: 356 (1907). yungense Rosenst. in Fedde, Repert. Spec. Nov. 11: 55 (1912). Pteridium aquilinum var. arachnoideum (Kaulf.) Brade in Zeitsch. Deut. Ver. Wissensch. Kunst, Sao Paulo, 1: 56 (1920). Pteris altissima Poir. in Lam. Encyc. 5: 722 (1804). P. Buchtienii Rosenst. in Fedde, Repert. Spec. Nov. 6: 309 (1909). P. deflexa Link, Hort. Berol. 2: 30 (1883). P. denticulata Sw. Prodr. 129 (1788). P. Haenkeana Presl, Rel. Haenk. 1: 55 (1825). P. Herne var. adaucta Rosenst. in Fedde, Repert. Spec. Nov. 12: 468 (191 ¥, Rea var. inermis Rosenst. in Fedde, Repert. Spec. Nov. 11: 53 (1912). P. podophylla Sw. in Schrad. Journ. 1800(2): 67 (1801). 1950 IO ee to ee ee 15 P. propinqua Agardh, Recens. 65 (1839). P. pungens Willd. Sp. Pl. 5: 387 (1810). P. quadriaurita Retz. Obs. 6: 38 (1791). Saccoloma brasiliense (Presl) Mett. in Ann. Sci. Nat. (ser. 4) 15: 80 (1861). S. elegans Kaulf. in Berl. Jahrb. Pharm. 1820: 51. S. Guentheri Rosenst. in Fedde, Repert. Spec. Nov. 25: 58 (1928). S. inaequale var. caudatum Hieron. in Hedwigia, 47: 207 (1908). Stenochlaena angusta Underw. in Bull. Torr. Bot. Club, 33: 594 (1906). S. erythrodes (Kunze) Underw. in Bull. Torr. Bot. Club, 33: 595 (1906). S. japurensis (Mart.) Griseb. Fl. Brit. W. Ind. 674 (1864). S. marginata (Schrad.) C. Chr. Ind. Fil. 624 (1905). S. sorbifolia (L.) J. Sm. in Journ. Bot. 4: 149 (1841). Tectaria Buchtienii (Rosenst.) Maxon in Proc. Biol. Soc. Wash. 46: 148 (1933). T. incisa Cav. Deser. 249 (1802). Trismeria trifoliata (L.) Diels in Engl. & Prantl, Nat. Pflanzenfam. 1(4): 265 (1899). Vittaria angustifolia (Sw.) Baker in Mart. Fl. Bras. 1(2): 544 (1870). filifolia Fée, Mém. 3: 20, t. 3, fig. 6 (1851-1852). latifolia Benedict in Bull. Torr. Bot. Club, 41: 403 (1914). lineata (L.) J. Sm. in Mém. Acad. Turin. 5: 421, t. foc fig. 5 (1793). Orbignyana Mett. ex Kuhn in Linnaea, 36: 66 (1869). remota Fée, Mém. 7: 26, t. 20, fig. 1 (1857). Ruiziana Fée, Mém. 3: 16, t. 3, fig. 3 aoe 1852). stipitata Kunze in Linnaea, 9: 77 (1834). Williamsii Benedict in Bull. Torr. Bot. Club, 41: 407 (1914). Woodsia crenata (Kunze) Hieron. in Engler, Bot. Jahrb. 34: 440 (1904). W. crenata var. pallidipes Hieron. in Engler, Bot. Jahrb. 34: 440 (1904). W. montevidensis (Spreng.) Hieron. in Engler, Bot. Jahrb. 22: 363 (1896). W. montevidensis var. fuscipes Hieron. in Hedwigia, 46: 322 (1907). eee GLEICHENIACEAE Dicranopteris boliviensis Maxon & Morton in Bull. Torr. Bot. Club, 66: 44 (1939). Gleichenia Bancroftii Hook. Sp. Fil. 1: 5, t. 4A (1844). bifida (Willd.) Spreng. Syst. Veg. 4: 27 (1827). Buchtienii Christ & Rosenst. in Fedde, Repert. Spec. Nov. 5: 229 (1908). flexuosa (Schrad.) Mett. in Ann. Lugd. Bat. 1: 50 (1863). nervosa (Kaulf.) Spreng. Syst. Veg. 4: 25 (1827). pectinata (Willd.) Presl, Rel. Haenk. 1: 71 (1825). tomentosa (Sw.) Spreng. Syst. Veg. 4: 27 (1827). velata (Kunze) Mett. Fil. Lips. 113 (1856). yungensis Rosenst. in Fedde, Repert. Spec. Nov. 5: 228 (1908). ANNAN AMA SCHIZAEACEAE Anemia anthriscifolia Schrad. in Gott. Gelehrt. Anz. 1824: 865. anthriscifolia forma nana Lindm. in Ark. Bot. 1: 258 (1903). flexuosa (Sav.) Sw. Syn. 156 (1806). Herzogii Rosenst. in Meded. Rijks Herbar. 19: 24 (1913). myriophylla Christ in Bull. Herb. Boiss. (ser. 2) 7: 793 (1907). phyllitidis (L.) Sw. Syn. 155 (1806). Presliana Prantl, Schiz. 104 (1881). rotundifolia Schrad. in Gétt. Gelehrt. Anz. 1824: 865. tomentosa (Sav.) Sw. Syn. 157 (1806). trichorhiza Gardn. in Hook. Ic. Pl. 9: t. 876 (1852). Wettsteinii Christ in Denkschr. Akad. Wiss. Wien, 79: 54, t. 9 (1908). ah oft il al all a at 16 Lygodium polymorphum (Cav.) HBK. Nov. Gen. & Spec. 1: 31 (1816). L. venustum Sw. in Schrad. Journ. 1801(2): 303 (1803). SALVINIACEAE Azolla filiculoides Lam. Encyc, 1: 343 (1783). A. microphylla Kaulf. Enum, Fil. 273 (1824). Salvinia auriculata Aubl. Hist. Pl. Guian. 2: 969 t. 367 (1775). MARSILIACEAE Pilularia Mandonii A. Br. in Monatsber. Akad. Berl. 1870: 752. MARATTIACEAE Danaea Moritziana Pres] in Abh. Bohm. Ges. Wiss. (ser. 5) 4: 295 (1847). Marattia alata Sw. Prodr, 128 (1788). M. Kaulfussii J. Sm. in Hook. Gen. sub t. 26 (1838). OPHIOGLOSSACEAE Botrychium cicutarium (Sav.) Sw. Syn. 171 (180 B. Schaffneri Underw. in Bull. le Bot. Club, = 61 (1903). Ophioglossum coriaceum A. Cunn. in Hook. Compan. Bot. Mag. 2: 361 (1837). O. crotalophoroides Walt. FI. Gar 256 (1788). ? QO. ellipticum Hook. & Grev. Ic. Fil. 1: t. 40A (1827). O. nudicaule var. tenerum (Mett.) Clausen in Mem. Torr. Bot. Club, 19(2): 146 O. reticulatum L. Sp. Pl. 2: 1063 (1753). O. scariosum Clausen in Mem. Torr. Bot. Club, 19(2): 153 (1938). EQUISETACEAE Equisetum bogotense HBK. Nov. Gen. & Spec. 1: 42 (1816). E. giganteum L. Sp. Pl. (ed. 2) 2: 1517 (1763). E. pyramidale J. E. Goldm. in Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 469 LYCOPODIACEAE Lycopodium andinum Rosenst. in Fedde, Repert. Spec. Nov. 5: 239 (1908). L. aristatum Humb. & Bonpl. ex Willd. Sp. Pl. 5: 17 (1810). L. bolivianum Rosenst. in Fedde, Repert. Spec. Nov. 11: 59 (1912). L. bolivianum var. teretiusculum Rosenst. in Fedde, Repert. Spec. Nov. 11: 60 (1912). L. Brongniartii Spring, Monog. 1: 33 (1842). L. carolinianum L. Sp. Pl. 2: 1104 (1758). L, cernuum L., Sp. Pl, 2: 1103 (1753). L. clavatum L. Sp. Pl. 2: 1101 (1758). L. complanatum L. Sp. Pl. 2: 1104 (1753). L. contiguum Kl. in Linnaea, 18: 519 (1844). L. dichotomum Jacq. Enum. Stirp. Vindob. 314 (1762). L, Jussiaei Desv. ex Poir. in Lam. Encyc. Suppl. 3: 543 (1813). L. Lechleri Hieron. in Engler, Bot. Jahrb. 34: 571 (1905). L. linifolium L. Sp. Pl. 2: 1100 (1753). L. longipes Hook. & Grev. in Hook. Bot. Misc. 2: 372 (1831). L. myrsinites Lam. Encyc. 3: 654 (1789). L. nubigenum Herzog in Meded. Rijks Herbar. 27: 2 (1915). L. paniculatum Desv. ex Poir. in Lam. Encyc. Suppl. 3: 543 (1813). L. Pearcei Baker, Handbk. eee 14 (1887). L. reflexum Lam. Encye. 3: 653 (178 L. Saururus Lam. Encyc. 3: 658 NN L. subulatum Desyv. ex Poir. in Lam. Encye. Suppl. 3: 544 (1813). L. taxifolium Sw. Fl. Ind. Oce. 3: 1573 (1806). ? L. tetragonum Hook. & Grev. Ic. Fil. 1: t. 109 (1828). L. verticillatum L. f. Suppl. 448 (1781). L. Williamsii Underw. & Lloyd in Bull. Torr. Bot. Club, 33: 112 (1906). Urostachys Pflanzii Nessel in Fedde, Repert Spec. Nov. 36: 181 (1934) = Lycopodium sp. PSILOTACEAE Psilotum nudum (L.) Beauv. Prodr. Cing. & Six. Fam. L’Aethéogamie, etc. 112 (1805). SELAGINELLACEAE Selaginella anceps Presl in Abh. Bohm. Ges. Wiss. (ser. 5) 3: 581 (1845). - Galeottii Spring in Bull. Acad. Roy. Bruxelles, 10(3) : 230 aa . haematodes (Kunze) Spring i in Mart. Fl. Bras. 1(1): 126 (184 leptoblepharis A. Br. in Planch. & Triana in Ann. Sci. Nat. "ise 5) 3: 279 18 nn tn in (1865). macrophylla A. Br. in Planch. & Triana in Ann. Sci. Nat. (ser. 5) 3: 302 (1865 eguholig (HBK.) Spring in Bull. Acad. Roy. Bruxelles, aoe ae (1843). Mildei Hieron. in Engl. & Prantl, Nat. Pflanzenfam. 1(4): 671 01). peruviana (Milde) Hieron. in Hedwigia, 39: 307 (1900). peruviana var. Dombeyana Hieron. in npn Se 39: 308 (1900). polycephala Baker in Journ. Bot. 21: 332 (1883 radiata (Aubl.) Spring, Monog. 2: 120 elon: rubescens Hieron. in Hedwigia, 43: 236 (1904). silvestris Asplund in Ark. Bot. 20A(7): 30, fig. 3-5 (1926). trisuleata Asplund in Ark. Bot. 20A(7): 34, fig. 6 (1926). ISOETACEAE Isoétes boliviensis Weber in Hedwigia, 63: 247 (1922). I. glacialis Asplund in Ark. Bot. 20A(7): 35, fig. 7 (1926). I. Herzogii Weber in Hedwigia, 63: 250 (1922). GYMNOSPERMAE CYCADACEAE Zamia boliviana (Brongn.) A. DC. Prodr. 16(2): 540 (1868). PODOCARPACEAE Podocarpus Cardenasii Buchholz & Gray in Journ. Arnold Arb. st 142 (1948). P. magnifolius Buchholz & Gray in Journ. Arnold Arb. 29: 133 (1948). montanus sensu Britton in Bull. Torr. Bot. Club, 16: 13 (1889) = P. utilior. P. oleifolius Don in Lambert, Pinus, 2: 20 (1824). P. Parlatorei Pilger in Engler, Pflanzenr. IV, 5: 86 (1903). 18 P. Rusbyi Buchholz & Gray in Journ. Arnold Arb. 29: 134 (1948). Sa oe sensu Britton in Bull. Torr. Bot. Club, 16: 18 (1889) = P. sbyi. P. utilior Pilger in Fedde, Repert. Spec. Nov. 1: 189 (1905). GNETACEAE Ephedra americana var. andina (Poepp.) Stapf in Denkschr. Akad. Wiss Math- Natu Ki. 56: 86 (1889). . americana var. Humboldtii Stapf in Denkschr. Akad. Wiss. Math.-Naturw. K1. 56: 85 (1889). americana var. rupestris (Benth.) Stapf in Denkschr. Akad. Math.-Naturw. Kl. 56: 86 (1889 andina var. kaveilic (Wedd.) estesaats in DC. Prodr. 16(2): 353 (1868). = E, americana var. rupestri: breana Phil. in Anal. Univ. Chile, 91: 519 (1895). . triandra Tul. in Ann. Sci. Nat. ((ser. 4) 10: 125 (1858). ANGIOSPERMAE MONOCOTYLEDONEAE TYPHACEAE Typha domingensis Pers. Synops. Pl. 2: 532 (1807). POTAMOGETONACEAE Potamogeton filiformis Pers. Synops. Pl. 1: 152 (1805). P. pectinatus L, Sp. Pl. 127 (1758). P. pectinatus var. striatus (R. & P.) Hagstr. in Kgl. Sv. Vetensk. Akad. Handl. 55(5): 51 (1916). Ruppia filifolia (Phil.) Skottsb. in Kgl. Sv. Vetensk. Akad. Handl. 56(5): 171 (1916). Zannichellia palustris L. Sp. Pl. 969 (1753). NAJADACEAE Najas —— (Spreng.) Morong in Mem. Torr. Bot. Club, 3(2): 60, t. 67 (1893). SCHEUCHZERIACEAE Lilaea subulata Humb. & Bonpl. Pl. Aequin. 1: 222, t. 63 (1808). Triglochin concinna var. deserticola (Phil.) J. T. Howell in Leafl. West. Bot. 5(1): 19 (1947). ALISMACEAE Alisma bolivianum Rusby in Mem. N. Y. Bot. Gard. 7: 208 (1927). A. tenellum Mart. ex Schult. f. Syst. Veg. 7: 1600 (1830). Echinodorus grandiflorus var. ovatus Micheli in DC Monog. Phan. 3: 58 (1881): E. subulatus Engelm. in Gray, Man. (ed. 1) 460 (1848). E. tenellus (Mart.) Buchenau in Abh. Naturw. Ver. Bremen, 2: 21 (1868). E. tenellus var. latifolius (Seubert) Fassett in Rhodora, 57: 202 (1955). Lophotocarpus guyanensis (HBK.) Dur. & Schinz, Consp. Fl. Afr. 5: 487 (1894). Sagittaria lancifolia L. Syst. (ed. 10) 2: 1270 (1759). S. montevidensis Cham. & Schlechtd. in Linnaea, 2: 156 (1827). 19 BUTOMACEAE Hydrocleis cryptopetala R. E. Fries in Ark. Bot. 8(8): 47, t. 2, fig. 1-3 (1908). H. nymphoides (Willd.) Buchenau in Bremen Abh. 2: 7 (1869). HYDROCHARITACEAE Elodea chilensis (Planch.) Casp. in Monatsber. Berl. Akad. 1857: 47. GRAMINEAE Aciachne pulvinata Benth. in Hook. Ic. Pl. 14: 44, t. 1862 (1881). A. uniflora Baill. in Bull. Mens. Soc. Linn. Paris, 2: 1073 (1893). Aegopogon bryophilus Doell in Mart. Fl. Bras. 2(3): 289 (1880). Ae. cenchroides Humb. & Bonpl. ex Willd. Sp. Pl. 4(2): 899 (1806). Ae. Fiebrigii Mez in Fedde, Repert. Spec. Nov. 17: 145 (1921) = Ae. bryo- philus, Ae. geminiflorus var muticus Pilger in Engler, Bot. Jahrb. 27: 25 (1899) = Ae. bryophilus. Agenium villosum (Nees) Pilger in Fedde, Repert. Spec. Nov. 43: 82 (1938). Agropyron attenuatum (HBK.) Roem. & Schult. Syst. Veg. 2: 751 (1817). A. boliviacum P. Cand. Tribu des Hordées, 25, 46 (1901). A. breviaristatum Hitche. in Contrib. U. S. Nat. Herb. 24: 353 (1927). Agrostis araucana Phil. in Anal, Univ. Chile, 94: 14 (1896). boliviana Mez in Fedde, Repert. Spec. Nov. 18: 1 (1922). breviculmis Hitche. in U. S. Dept. Agric. Bur. Pl. Ind. Bull. 68:36, t. 18 (1905). exasperata Trin. in Mém. Acad. St. Pétersb. (ser. 6) 4(1): 352 (1841). gelida Trin. in Mém. Acad. St. Pétersb. (ser. 6) 4(1): 343 (1841). Haenkeana Hitchc. in Contrib. U. S. Nat. Herb. 24: 381 (1927). montevidensis Spreng. ex Nees, Agrost. Bras. 403 ( 1829). nana (Presl) Kunth, Rév. Gram. 3: 596 (1829) = A. breviculmis. perennans (Walt.) Tuckerm. in Amer. Journ. Sci. 45: 44 (1843). stolonifera L. Sp. Pl. 62 (1753). tolucensis HBK. Nov. Gen. & Spec. 1: 135 (1816). verticillata Vill. Prosp. Pl. Dauph. 16 (1779). Aira conferta (Pilg.) Hitche. in Contrib. U. S. Nat. Herb. 24: 361 (1927). Alopecurus aequalis Sobol. Fl. Petrop. 16 (1799). A. alpinus var. aristatus Hook. f. Fl. Antarct. 2: 370 (1847). A. bracteatus Phil. in Anal. Univ. Chile, 94: 6 (1896). A. Hitchcockii Parodi in Rev. Fac. Agron. Vet. Buenos Aires, 7: 366 (1931). Andropogon altus Hitche. in Contrib. U. S. Nat. Herb. 17: 208 (1913). A. bicornis L. Sp. Pl. 1046 (1753). cirratus Hack. in Flora, 68: 119 (1885). condensatus HBK. Nov. Gen. & Spec. 1: 188 (1816). cordatus Swallen in Contrib. U. S. Nat. Herb. 29(6): 274 (1948). Fabricii Thunb. ex Henr. in Meded. Rijks Herbar. 40: 44 (1921). Hassleri Hack. in Bull. Herb. Boiss. (ser. 2) 4(3): 266 (1904). hirtiflorus (Nees) Kunth, Rév. Gram. 2: 569 (1832). lateralis Nees, Agrost. Bras. 329 (1829). leptocladus Hack. in Flora, 68: 122 ( 1885). leucostachyus HBK. Nov. Gen. & Spec. 1: 187 (1816). nutans var. agrostoides (Speg.) Hack. in DC. Monog. Phan. 6: 952 (1889). nutans var. stipoides (HBK.) Hack. in DC. Monog. Phan. 6: 530 (1889). Riedelii Trin. in Mém. Acad. St. Pétersb. (ser. 6) 2: 263 (1833). saccharoides Sw. Prodr. 26 (1788). PPP PRP >>> pp PP PP > >>> >>> b> saccharoides subsp. genuinus var. barbinodis (Lag.) Hack. in DC. Monog. Phan. 6: 494 (1889). saccharoides var. laguroides (DC.) Hack. in Mart. Fl. Bras. 2(8):293 (1883). saccharoides var. parvispiculus Hitche. in Contrib. U. S. Nat. Herb. 24: 497 (1927). saccharoides var. perforatus (Trin.) Hack. in DC. Monog. Phan. 6: 496 (1889). Selloanus (Hack.) Hack. in Bull. Herb. Boiss. (ser. 2) 4: 266 (1904). tener (Nees) Kunth, Rév. Gram. 2: 565 (1832). tener var. genuinus subvar. hirtiglumis Henr. in Meded. Rijks Herbar. 40: 42 (1920). ternatus subsp. macrothrix (Trin.) Hack. in Mart. Fl. Bras. 2(3): 287 1883 i‘. villosus (Nees) Ekman in Ark. Bot. 11(4): 9 (1912). virgatus Desv. ex Hamilt. Prodr. Pl. Ind. Occ. 9 (1825). Anthaenantia gigantea (O. Ktze.) K. Schum. in Just, Bot. Jahresber. 26(1): 829 (1900). Anthaenantiopsis Fiebrigii Mez in Engler, Bot. Jahrb. 56, Beibl. 125: 11 (1921). Anthochloa lepidula Nees & Meyen in Meyen, Reise, 2: 14 (1835). A. rupestris Remy in Ann. Sci. Nat. (ser. 3) 6:347 (1846) = A. lepidula. Aphanelytrum procumbens Hack. in Oesterr. Bot. Zeitschr. 52: 18 (1902). Aristida adscensionis L. Sp. Pl. 82 (1758). A. adscensionis var. coerulescens subvar. condensata Hack. apud Stuckert, Contrib. Conoc. Gram. Argent. 91 (1904). ae or Peer oP Ue UP A. adscensionis var. humilis forma viridis O. Ktze. Rev. Gen. 3(2):339 (1898). A. poste oe var. normalis forma violascens O. Ktze. Rev. Gen. 3(2): 339 (189. A. Asplundii aes in Meded. Rijks Herbar. 54: 42 (1926). A. capillacea Lam, Encyc. Tabl. 1: 156 (1791). A. circinalis Lindm. in Kgl. Sv. Vet. Akad. Hand]. 34(6): 18, t. 7a (1900). A. complanata Trin. in Mém. Acad. St. Pétersb. (ser. 6) 1: 853 (1880). A. ecuadoriensis Henr. in Meded. Rijks Herbar. 58A: 307 (1932). A. enodis Hack. in Fedde, Repert. Spec. Nov. 11: 21 (1912). A. Friesii Hack. ex Henr. in Meded. Rijks Herbar. 54: 186 (192 A. implexa Trin. in Mém. Acad. St. Pétersb. (ser. 6) 2(1): 48 (188) A. mnversa Hack. in Ark. Bot. 8(8): 37 (1909) == A. mendoc A. longiramea var. boliviana Henr. in Meded. Rijks Herbar. 40: 56 (1921) = A. complanata. A. powcr var. robusta Merrill in Cire. U. S. Div. Agrost. 34: 5 (1901). A. Mandoniana Henr. in Meded. Rijks Herbar. 40: 55 (1921) — A. adscen- sionis, A. mendocina R? A. Phil. in Anal. Univ. Chile, 36: 205 (1870). A. mendocina var. macrantha (Parodi) Henri, in Meded. Rijks Herbar. 54A: 267 (1927). A. Pflanzii Mez in Fedde, Repert. Spec. Nov. 17: 151 (1921) = A. enodis. A. riparia Trin. in Mém. Acad. St. Pétersb. (ser. 6) 2(1): 48 (1836). A. riparia var, andina Henr. in Meded. Rijks Herbar. 58A: 185 (1932). Arthrostylidium racemiflorum Steud. Syn. Pl. Gram. 336 (1854). Arundinaria Herzogiana Henr. in Meded. Rijks Herbar. 40: 75 (1921). Arundinella Berteroniana (Schult.) Hitchc. & Chase in Contrib. U. S. Nat. Herb. 18: 290 (1917). A. confinis (Schult.) Hitche. & Chase in Contrib. U. S. Nat. Herb. 18: 290 (1917). Arundo donax L. Sp. Pl. 81 (1753). Avena barbata Brot. Fl. Lusit. 1: 108 (1804). A. fatua L. Sp. Pl. 80 (1758). A. scabrivalvis Trin. in Mém. Acad. St. Pétersb. (ser. 6) 2: 28 (1836). Axonopus aureus Beauv. Ess. Agrost. 12 (1812). barbigerus (Kunth) Hitche. in Contrib. U. S. Nat. Herb. 24: 483 (1927). capillaris (Lam.) Chase in Proc. Biol. Soc. Wash. 24: 183 (1911). chrysoblepharis (Lag.) Chase in Proc. Biol. Soc. Wash. 24: 184 (1911). compressus (Sw.) Beauv. ex Steud. Nomencl. (ed. 2) 1: 174 (1840). elegantulus (Presl) Hitche. in Contrib. U. S. Nat. Herb. 24: 483 (1927). fissifolius (Raddi) Kuhlm. in Comm, Linkas Telegr. Matto Grosso, 67: 87 22). Herzogii (Hack.) Hitch. in Contrib. U. S. Nat. Herb. 24: 431 (1927). marginatus (Trin.) Chase in Contrib. U. S. Nat. Herb. 17: 226 (1918). Purpusii (Mez) Chase in Journ. Wash. Acad. Sci. 17: 144 (1927). siccus (Nees) Kuhlm. in Comm. Linkas Telegr. Matto Grosso, 67: 87 (1922). scoparius (Fliigge) Hitchc. in Contrib. U. S. Nat. Herb. 22: 471 (1922). uteloua aristidoides (HBK.) Griseb. Fl. Brit. W. Ind. 537 (1864). curtipendula Torr. in Emory, Milit. Reconn. 154 (1848), nomen provisoriwm. megapotamica (Spreng.) O. Ktze. Rev. Gen. 3(2): 341 (1898). prostrata Lag. in Varied. Cienc. 2(4): 141 (1805). racemosa Lag. in Varied. Cienc. 2(4): 141 (1805). simplex Lag. in Varied. Cienc. 2(4): 141 (1805). simplex var. actinochloides Henr. in Meded. Rijks Herbar. 40: 65 (1921) DWH E b> >>> bbb bb> . simplex. Brachiaria plantaginea (Link) Hitche. in Contrib. U. S. Nat Herb. 12: 212 (1909). Brachypodium mexicanum (Lag.) Link, Hort. Berol. 1: 41 (1827). Briza Mandoniana (Griseb.) Henr. in Meded. Rijks Herbar. 40: 70 (1921). B. Mandoniana var. Herzogiana Henr. in Meded. Rijks Herbar. 40: 71 (1921) = B. Mandoniana. B. Mandoniana var. tuberculata Henr. in Meded. Rijks Herbar. 40: 71 (1921) = B. Mandoniana. B. Mandoniana var. vallegrandensis Henr. in Meded. Rijks Herbar. 40: 71 (1921) = B. Mandoniana. - Spicigera (Presl]) Steud. Nomencl. (ed. 2) 1: 225 (1840). stricta (Hook. & Arn.) Steud. Nomencl. (ed. 2) 1: 225 (1840). Bromidium hygrometricum var. breviaristatum O. Ktze. Rev. Gen. 3(2): 342 (1898) = Calamagrostis sp. ? Bromus angustatus Pilg. in Engler, Bot. Jahrb. 25: 719 (1898) = B. unioloides. Buchtienii Hack. in Fedde, Repert. Spec. Nov. 11: 30 (1912) = B. pitensis. lanatus HBK. Nov. Gen. & Spec. 1: 150 (1816). Mandonianus Henr. in Fedde, Repert. Spec. Nov. 23: 177 (1926) = Dielsi- ochloa floribunda, Pflanzii Pilg. in Engler, Bot. Jahrb. 49: 189 (1913) = B. lanatus. pitensis HBK. Nov. Gen. & Spec. 1: 152 (1816). segetum HBK. Nov. Gen. & Spec. 1: 151 (1816). Trinii Desv. in Gay, Fl. Chile, 6: 441 (1853). unioloides HBK. Nov. Gen. & Spec. 1: 151 (1816). amagrostis amoena (Pilg.) Pilg. in Engler, Bot. Jahrb. 42: 60 (1908). Antoniana (Griseb.) Steud. ex Hitche. in Contrib. U. S. Nat. Herb. 24: 378 (1927). Beyrichiana Ness ex Doell in Mart. Fl. Bras. 2(3): 53, t. 16 (1878). boliviensis Hack. in Fedde, Repert. Spec. Nov. 6: 156 (1908) = C. hetero- hylla. brevifolia (Presl) Steud. Nomencl. (ed. 2) 1: 249 (1840). calderillensis Pilg. in Engler, Bot. Jahrb. 42: 72 (1908). cephalantha Pilg. in Engler, Bot. Jahrb. 42: 61 (1908). oo oo 222 Af AO PEP wHb bo bo chrysantha (Presl) Steud. Nomencl. (ed. 2) 1: 250 (1840). curta (Wedd.) Hitche. in Contrib. U. 8S. Nat. Herb. 24: 376 (1927). curvula (Wedd.) Pilg. in Engler, Bot. Jahrb. 42: 60 (1908). elegans (Wedd.) Henr. in Meded. Rijks Herbar. 40: 61 (1921) = eminens, eminens = big Steud. Nomencl. (ed. 2) 1: 250 (1840). emt r. sordida O. Ktze. Rev. Gen. 3(2): 344 (1898) = C. eminens. eminens var. tunariensis O. Ktze. Rev. Gen. 3(2). 344 (1898) = C. eminens. Fiebrigii Pilg. in Engler, Bot. Jahrb. 42: 68 (1909). filifolia (Wedd.) Henr. in Meded. Rijks Herbar. 40: 61 (1921) — C. amoena. glacialis (Wedd.) Hitche. in Contrib. U. S. Nat. Herb. 24: 375 (1927). gracilis (Wedd.) Henr. in Meded. Rijks Herbar. 50: 61 (1921). heterophylla (Wedd.) Pilg. in Engler, Bot. Jahrb. 42: 64 (1908). Humboldtiana Steud. Nomencl. (ed. 2) 1: 250 (1840). Jamesonii Steud. Syn. Pl. Glum. 1: 191 (1854). leiophylla (Wedd.) Hitche. in Contrib. U. S. Nat. Herb. 24: 367 (1927). Lilloi Hack. in Stuckert in Anal. Mus. Nac. Buenos Aires, 13: 477 (1906). longearistata (Wedd.) Hack. ex Sodiro in Anal. Univ. Quit. 3: 481 (1889). montevidensis Nees, Agrost. Bras. 401 (1829 } linearis Hack. in Bada: Repert. Spec. Nov. 6: 157 (1908) = C. montevidensis. oye tg is ae eee

D. mutica Wedd. in Bull. Soc. Bot. France, 22: 177 (1875) — Calamagros- tis sp. ? D. nematophylla Wedd. in Bull. Soc. Bot. France, 22: 179 (1875) = Calama- grostis sp. ? D. nivalis Wedd. in Bull. Soc. Bot. France, 22: 156 (1875) — Calamagrostis ovata. D. obtusata Wedd. in Bull. Soc. Bot. France, 22: 177 (1875) — Calamagros- is sp. ? D. phalaroides Wedd. in Bull. Soc. Bot. France, 22: 177 (1875) = Calama- wi rigescens D. pic oo in Bull. Soc. Bot. France, 22: 177 (1875) — Calamagros- ? D. pia Wedd. in Bull. Soc. Bot. France, 22: 178 (1875) — Calamagros- is sp. ? D. subsimilis Wedd. in Bull. Sce. Bot. France, 22: 178 (1875) — Calamagros- tS SD. 2 A coe Wea in Bull. Soc. Bot. France, 22: 178 (1875) — Calamagros- Sv bcuicnce floribunda (Pilg.) Pilg. in Rev. Argent. Agron. 11: 257 (1944). Digitaria Gerdesii var. boliviensis Henr. Monog. Digit. 287 (1950). D. Lehmanniana Henr. in Blumea, 1(1): 107 (1934). Dissanthelium calycinum (Presl) Hitche. in Journ. Wash. Acad. Sci. 13: 224 1923). D. minimum Pilg. in Engler, Bot. Jahrb. 56, Beibl. 123: 28 (1920). D. peruvianum (Nees & Mey.) Pilg. in ee Bot. Jahrb. 37: 378 (1906). D. Trollii Pilg. in Notizbl. 11: 778 (193 Distichlis humilis Phil. in Anal. Mus. m5 Chile, 8: 86 (1891). D. spicata (L.) Greene in Bull. Calif. Acad. 2: 415 (1887). Echinochloa crusgalli var. crus-pavonis (HBK.) Hitche. in Contrib, U. S. Nat. Herb. 22: 148 (1920). Eleusine indica (L.) Gaertn. Fruct. 1: 8 (1788). Elymus angulatus Presl, Rel. Haenk. 1: 264 (1830). Elyonurus adustus (Trin.) Ekm. in Ark. Bot. 13(10): 6 (1918). E. tripsacoides Humb. & Bonpl. ex Willd. Sp. Pl. 4(2): 941 (1806). E. tripsacoides var. ciliaris (Kunth) Hack. in DC. Monog. Phan. 6: 333 (1889) = E. tripsacoides. Eragrostis articulata (Schrsak) Nees, Agrost. Bras. 502 (1829) articulata var. glabrescens Henr. in Meded. Rijks Herbar. 40: 69 (1921). . atrovirens (Desf.) Trin. in Steud. Nomencl. (ed. 2) 1: 562 (1840). . bahiensis Schrad. ex Roem. & Schult. Mant. 2: 318 (1824). iensis var. boliviensis Henr, in Meded. Rijks Herbar. 40: 68 (1921) = = E. lurida. . boliviensis Jedw. in Bot. Archiv Mez, 5: 205 (1924) = E. ality . Buchtienii Hack. in Fedde, Repert. Spec. Nov. 6: 157 (1908) — E. mon- f tufari. cilianensis (All.) Link ex Vign. Lut. in Malpighia, 18: 386 (1904). ciliaris (L.) Link, Hort. Berol. 1: 192 (1827). contristata Nees & Meyen in Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 163 (1843) = E. lurida. expansa Link, Hort. Berol. 1: 190 (1827). glomerata (Walt.) L. H. Dewey in Contrib. U. S. Nat. Herb. 2: 543 (1894). hypnoides (Lam.) BSP. Prelim. Cat. N. Y. 69 (1888). lugens Nees, Agrost. Bras. 505 (1829). lugens subsp. flaccida (Lindm.) Hack. in Stuckert in Anal. Mus. Nac. Buenos Aires, 21: 134 (1911). lurida Presl, Rel. Haenk. 1: 276 (1830). 25 maypurensis (HBK.) Steud. Syn. Pl. Glum. 1: 276 (1854). mexicana (Lag.) Link. Hort. Berol. 1: 190 (1827). montufari (HBK.) Steud. Nomencl. (ed. 2) 1: 563 (1840). nigricans (HBK.) Steud. Nomencl. (ed. 2) 1: 563 (1840). pilosa (L.) Beauv. Ess. Agrost. 71 (1812), combination implied but not actually made. polytricha Nees, Agrost. Bras. 507 (1829). reptans (Michx.) Nees, Agrost. Bras. 514 (1829). solida Nees, Agrost. Bras. 501 (1829 soratensis Jedw. in Bot. Archiv Mez, 5: 213 (1924) = E. lugens. . subatra Jedw. in Bot Archiv Mez, 5: 202 (1924) = E. nigricans. . tristis Jedw. in Bot. Archiv Mez, 5: 205 (1924) = E. nigricans. . villamontana Jedw. in Bot. Archiv Mez, 5: 197 (1924) = ? Erianthus angustifolius Nees, Agrost. Bras. 316 (1829). E. Trinii (Hack.) Hack. in DC. Monog. Phan. 6: 135 (1889). Eriochloa distachya HBK. Nov. Gen. & Spec. 1: 95, t. 30 (1816). E. punctata (L.) Desv. ex Hamilt. Prodr. Pl. Ind. Occ. 5 (1825). Eriochrysis cayennensis Beauv. Ess. Agrost. 8, t. 4, fig. 11 (1812). E. Warmingiana (Hack.) Kuhlm. in Comm. Linkas Telegr. Matto Grosso, 67: 29 (1922). Festuca bromoides L. Sp. Pl. 75 (1753). - Buchtienii Hack. in Fedde, Repert. Spec. Nov. 6: 160 (1908) —F. dolicho- ylla. dissitiflora Steud. ex Griseb. in Goett. Abh. 24: 287 (1879) = F. Saeed dissitiflora subsp. sa Srieraeseday var. trachyphylla Hack. ex St.-Yves Candollea, 3: 246 (1927) = dissitiflora subsp. loricata var. villipalea St.-Yves in Candollea, 3: 250 (1927) =? SSSA Se %y dolichophylla Presl, Rel. Haenk. 1: 258 (1830). Fiebrigii Pilg. in Engler, Bot. Jahrb. 37: 510 (1906) = F. procera laetiviridis Pilg. in Engler, Bot. Jahrb. 87: 510 (1906) = F. dolichophylla. megalura Nutt. in Journ. Acad. Philad. (ser. 2) 1: 3). (1848). orthophylla Pilg. in Engler, Bot. Jahrb. 25: 717 (18 phylla var. boliviana Pilg. in Engler, Bot. nee 37: 508 (1906) = F. orthophylla. Pflanzii Pilg. in Engler, Bot. Jahrb. 49: 188 (1913) = F. dolichophylla. procera HBK. Nov. Gen. & Spec. 1: 154 (1816). rigescens (Presl) Kunth, Rév. Gram. 3: 609 (1829). scirpifolia (Presl) Kunth, Rév. Gram. 3: 609 (1829) = F. dolichophylla. Stuebelii Pilg. in Engler, Bot. Jahrb. 25: 717 (1898) — F. rigescens. sublimis Pilg. in Engler, Bot. Jahrb. 25. 718 (1898). tectoria subsp. Mandoniana St.-Yves in Candollea, 3: 242 (1927). tectoria subsp. Mandoniana var. mutica St.-Yves in Candollea, 3: 243 (1927). ulochaeta Steud. Syn. Pl. Glum. 1: 305 (1854). Gouinia latifolia (Griseb.) Vasey in Contrib. U. S. Nat. Herb. 1: 365 (1895). G. virgata (Presl) Scribn. in U. S. Dept. Agric. Div. Agrost. Bull. 4: 10 (1897). Gymnopogon jubiflorus Hitche. in Contrib. U. S. ver Herb, 24: 412 (1927). G. spicatus (Spreng.) O. Ktze. Rev. Gen. 3(2): 354 (1898). G. spicatus var. longiaristatus O. Ktze. Rev. Gen. sa): 354 (1898). Gynerium atacamense Phil. in Linnaea, 33: 289 (1864-1865). G. sagittatum (Aubl.) Beauv. Ess. Agrost. 138, t. 24, fig. 6 (1812). Hackelochloa granularis (L.) O. Ktze. Rev. Gen. 2: 776 (1891). Heteropogon contortus (L.) Beauv. ex Roem. & Schult. Syst. Veg. 2: 836 (1817). H. melanocarpus (EIlI.) Benth. in Journ. Linn. Soc. 19: 71 (1882). ow eee ee ey 26 H. villosus Nees, Agrost. Bras. 362 (1829). Holcus halepensis L. Sp. Pl. 1047 (1753). H. lanatus L. Sp. Pl. 1048 (1753). H. Sorghum L. Sp. Pl. 1047 (1753). Homolepis aturensis (HBK.) Chase in Proc. Biol. Soc. Wash. 24: 146 (1911). Hordeum halophilum Griseb. in Goett. Abh. 19: 249 (1874). . muticum Presl, Rel. Haenk. 1: 327 (1830). . muticum var. andicola (Griseb.) Thell. Fl. Advent. Montpellier, 157 (1912). . nodosum L. Sp. Pl. (ed. 2) 1: 126 (1762). H. nodosum var. parviflorum (Hack.) Henr. & Thell. in Meded. Rijks Herbar. 40: 75 (1921). Hyparrhenia bracteata (Humb. & Bonpl.) Stapf in Fl. Trop. Afr. 9: 360 (1919). Ichnanthus breviscrobs Doell in Mart. Fl. Bras. 2(2): 294 (1877). calvescens (Nees) Doell in Mart. Fl. Bras. 2(2): 285 (1877). candicans (Nees) Doell in Mart, Fl. Bras. 2(2): 291 (1877). - Minarum (Nees) Doell in Mart. Fl. Bras. 2(2): 294 (1877). pallens (Sw.) Munro in Benth. Fl. Hongk. 414 (1861). peruvianus Mez in Fedde, Repert. Spec. Nov. 15: 129 (1918). I. Ruprechtii Doell in Mart, Fl. Bras. 2(2): 293 (1877). Imperata brasiliensis Trin. Mém. Acad. St. Pétersb. (ser. 6) 2: 331 (1832). I. contracta (HBK.) Hitche. in Rept. Missouri Bot. Gard. 4: 146 (1893). I. minutifiora Hack. in DC. Monog. Phan. 6: ee pore I. tenuis Hack. in DC. Monog. Phan. 6: 689 (18 Isachne arundinacea (Sw.) Griseb. Fl. Brit. W. 4 c (1864). Koeleria cristata (L.) Pers. Synops. Pl. 1: 97 (1805). K. gracilis Pers. Synops. Pl. 1: 97 (1805). K. gracilis subsp. boliviensis (Domin) Domin in Biblioth. Bot. 65: 237 (1907). K. pseudocristata var. andicola Domin in Fedde, Repert. Spec. Nov. 2: 94 ne _ . Pee Lamprothyrsus Hieronymi (O. Ktze.) Pilg. in Engler, Bot. Jahrb. 37, Beibl. 85: 58 (1906). L. Hieronymi var. Ae hy Pilg. in Engler, Bot. Jahrb. 37, Beibl. 85: 59 (1906) = L. Hieron L. Hieronymi var. tincta Pig, in Engler, Bot. Jahrb. 37, Beibl. 85: 59 (1906) = L. Hieron Lasiacis divaricata (L.) Hitche. in Contrib. U. S. Nat. Herb. 15: 16 (1910). L. ligulata Hitche. & Chase in Contrib. U. S. Nat. Herb. 18: 337 (1917). L. sorghoides (Desv.) Hitchc. & Chase in Contrib. U. S. Nat. Herb. 18: 338 (1917). Leersia hexandra Sw. Prodr. 21 (1788). Leptochloa domingensis (Jacq.) Trin. Fund. Agrost. 133 (1820). L. dubia (HBK.) Nees in Syllog. Pl. Ratisb. 1: 4 (1824). L. uninervia (Presl) Hitchc. & Chase in Contrib. U. S. Nat Herb. 18: 3383 7). L. virgata (L.) Beauv. Ess. Agrost. 71, t. 15, fig. 1 (1812). Leptocoryphium lanatum (HBK.) Nees, Agrost. Bras. 84 (1829). Lolium multiflorum Lam. Fl. Franc. 3: 621 (1778). L. perenne L. Sp. Pl. 83 (1753). L. temulentum L. Sp. Pl. 83 (1753). L. temulentum var. arvense (With.) Bab. Man. Brit. Bot. 377 (1843). Luziola peruviana Gmel. Syst. Nat. 1: 637 (1791). Lycurus phalaroides HBK. Nov. Gen. & Spec. 1: 142 (1816). L. phleoides HBK. Nov. Gen. & Spec. 1: 142, t. 45 (1816). Manisuris aurita (Steud.) Hitche. & Chase in Contrib. U. S. Nat. Herb. 18: 6 (1917). M. fasciculata (Lam.) Hitche. in Amer. Journ. Bot. 2: 299 (1915). Melica adhaerens Hack. in Fedde, Repert. Spec. Nov. 6: 158 (1908). M. adhaerens var. tenuis (Papp) Papp in Notizbl. 10: 412 (1928). Mandonii Papp in Notizbl. 10: 356 (1928). . secabra HBK. Nov. Gen. & Spec. 1: 164 (1816). . scabra var. glabra Papp in Fedde, Repert. Spec. Nov. 25: 145 (1928). violacea Cav. Ic. 5: 47, t. 472, fig. 2 (1799). Mesosetum rottboellioides (HBK.) Hitche. in Contrib. U. S. Nat. Herb. 12: 211 (1909). SRE S Microchloa indica (L. f.) O. Ktze. Rev. Gen. 3(2): 356 (1898). Muhlenbergia angustata (Presl) Kunth, Rév. Gram. 3: 594 (1829). elegans var. atroviolacea O. Ktze. Rev. Gen. 3(2): 357 (1898). M. elegans var. subviridis O. Ktze. Rev. Gen. 3(2): 357 (1898). fastigiata (Presl) Henr. in Meded. Rijks Herbar. 40: 59 (1921). Herzogiana Henr. in Meded. Rijks Herbar. 40: 58 (1921) — M. peruviana. Holwayorum Hitche. in Contrib. U. S. Nat. Herb. 24: 389 (1927). ligularis (Hack.) Hitchc. in Contrib. U. S. Nat. Herb. 24: 388 (1927). peruviana (Beauv.) Steud. Nomencl. (ed. 2) 1: 41 (1840). phragmitoides Griseb. in Goett. Abh. 19: 255 (1874). quitensis (HBK.) Hitche. in Contrib. U. S. a. Herb. 17: 292 (19138). rigida (HBK) Kunth, Rév. Gram. 1: 63 (1829 . tenuissima (Presl) Kunth, Rév. Gram. 3: 594 tay Munroa andina Phil. in Anal. Mus. Nac. Chile, 8: 90 (1891). M. andina var. breviseta Hack. ex Stuckert in Ann. Conserv. & Jard. Bot. Genéve, 17: 294 (1914). M. argentina Griseb. in Goett. Abh. 24: 300 (1879). M. decumbens R. A. Phil. in Anal. Mus. Nac. Chile, 8: 90 (1891). Nassella Asplundii Hitche. in Contrib. U. S. Nat. Herb. 24: 394 (1927). N. corniculata Hack. in Fedde, Repert. Spec. Nov. 6: 155 (1908) = N. pubi- Al latlal aba flora. N. deltoidea Hack. in Fedde, Repert. Spec. Nov. 11: 23 (1912) = N. pubi- flora. N. flaccidula Hack. in Fedde, Repert. Spec. Nov. 6: 154 (1908). = Stipa in- conspicua. N. flaccidula var. humilior Hack. in Fedde, Repert. Spec. Nov. 6: 155 (1908) = N. pubiflora. N. Meyeniana (Trin. & Rupr.) Parodi in Darwiniana, 7: 379 (1947). N. pubiflora (Trin. & Rupr.) Desv. in Gay, Fl. Chile, 6: 264 (1853). N. trachyphylla Henr. Meded. Rijks Herbar. 40: 57 (1921) — N. pubiflora. Olyra Buchtienti Hack. in Fedde, Repert. Spec. Nov. 11: 20 (1912). O. ciliatifolia Raddi, Agrost. Bras. 19 (1823)). O. ecaudata Doell in Mart. Fl. Bras. 2(2): 326 (1877). O. Heliconia Lindm. in Kgl. Sv. Vet. Akad. Handl. 34 (6): 11, t. 6 (1900). O. lateralis (Presl) Chase in Proc. Biol. Soc. Wash. 21: 179 (1908). O. latifolia L. Syst. Nat. (ed. 10) 2: 1261 (1759). O. micrantha HBK. Nov. Gen. & Spec. 1: 199 (1816). QO. pauciflora var. atrocarpa O. Ktze. Rev. Gen. 3(2): 357 (1898). O. pauciflora var. leucocarpa O. Ktze. Rev. Gen. 3(2): 357 (1898). Oplismenus hirtellus (L.) Beauv. Ess. Agrost. 54, 168 (1812). Orthoclada laxa (Rich.) Beauv. ex Nees, Agrost. pg ‘vi (1829). Oryzopsis florulenta Pilg. in Engler, Bot. Jahrb. 27: 26 (1899). O. Neesii Pilg. in Engler, Bot. Jahrb. 56, Beibl. Ps 26 (1920) = Stipa obtusa Panicum siusiat var. leianthum Hack. in Fedde, Repert. Spec. Nov. 6: 342 (19 P. aquaticum Poir. in Lam. Encyc. Suppl. 4: 281 (1816). 34(6): 10 (1900). boliviense Hack. in Fedde, Repert. Spec. Nov. 11: 19 (1912). caaguazense Henr. in Meded. Rijks Herbar. 47: 2 (1922). cayennense Lam. Encyc. Tabl. 1: 173 (1791). chloroticum Nees ex Trin. Gram. Pan. 236 (1826). cordovense Fourn. Mex. Pl. 2: 26 (1886 cyanescens Nees, Agrost. Bras. 220 (1829). echinulatum Mez in Notizbl. 7: 62 (1917). echinulatum var. boliviense Henr. in Meded. Rijks Herbar. 40: 50 (1921). (1905) = Trichachne saccharata. frondescens G.F.W. Meyer, Primit. Fl. Esseq. 56 (1818). Gerdesii Hack. in Oesterr. Bot. Zeitschr. 51: 333 (1901). Ghiesbreghtii Fourn, Mex. Pl. 2: 29 (1886) glutinosum Sw. Prodr. 24 (1788). granuliferum var. longifolium O. Ktze. Rev. Gen. 3(2): 361 (1898). hebotes Trin. in Mém. Acad. St. Pétersb. (ser. 6) 1: 30 (1834). helobium Mez ex Henr. in Meded. Rijks Herbar. 40: 52 (1921). hirticaule J. & C. Pres]. Rel. Haenk. 1: 308 0). laxum forma minus Hack. ex Stuckert, Terc. Contrib. 39 (1911). maximum Jacq. Collect. 1: 76 (1786). megiston Schult. Mant. 2: 248 (1824). nemorosum var. uncinatum O. Ktze. Rev. Gen. 3(2): 363 (1898). olyroides HBK. Nov. Gen. & Spec. 1: 102 (1816). pantrichum Hack. in Verh. Zool.—Bot. Ges. Wien, 65: 72 (1915). parvifolium Lam. Encyc. Tabl. 1: 173 (1791). paucispicatum Morong in Ann. N. Y. Acad. Sci. 7: 262 (1893). peladoénse Henr. in eae 4(3): 504 (1941). pilosum Sw. Prodr. 22 (178 polygonatum Schrad. ex a eats Mant. 2: 256 (1824). procurrens Nees ex Trin. Gram. Pan. 183 (1826). pulchellum Raddi, Agrost. Bras. 42 (1823). pygmaeum var. glabrescens Hack. in Fedde, Repert. Spec. Nov. 11: 18 (1912). quadriglume (Doell) Hitche. in Contrib. U. S. Nat. Herb. 24: 460 (1927). Rudgei Roem. & Schult. Syst. Veg. 2: 444 (1817). sciurotis Trin. Gram. Pan. 228 (1826). Sellowii Nees, Agrost. Bras. 153 (1829). (?) sempervirens O. Ktze. Rev. Gen. 3(2): 364 (1898). stoloniferum Poir. in Lam. Encyc. Suppl. 4: 274 (1816). stramineum Hitche. & Chase in Contrib. U. he Nat. Herb. 15: 67 (1910). trichanthum Nees, Agrost. Bras. 210 (1829 tricholaenoides Steud. Syn. Pl. Glum. ‘3 = (1854). uncinatum Raddi, Agrost. Bras. 41 (182 versicolor Doell in Mart. Fl. Bras. ee 254 (1877). zizanioides HBK. Nov. Gen. & Spec. 1: 100 (1816). WN NWNTs MO ION I Pariana bicolor Tutin in Journ. Linn. Soc. 50: 355 (1936). P. gracilis Doell in Mart. Fl. Bras. 2(2): 377 (1877). P. lunata Nees, Agrost. Bras. 295 (1829). P. zingiberina Doell in Mart. Fl. Bras. 2(2): 337 (1877). Paspalum attenuatum Presl, Rel. Haenk. 1: 212 (1830). P. barbatum Nees, Agrost. Bras. 27 (1829). P. boliviense Chase in Hitche. in Contrib. U. S. Nat. Herb. 24: 454 (1927). Bergii var. leiophyllum Hack. & Lindm. in Kgl. Sv. Vet. Akad. Handl. Friesiti Hack. ex Fries in Nov. Act. Soc. Sci. Upsal. (ser. 4) 1(1): 170 29 . Buchtienii Hack. in Fedde, Repert. Spec. Nov. 6: 153 (1908). candidum (Humb. & Bonpl.) Kunth in Mém. Mus. Hist. Nat. 2: 68 (1815). capillare Lam. Encyc. Tabl. 1: 176 (1791). castaneum Remy in Ann. Sci. Nat. (ser. 3) 6: 348 (1846). ceresia (O. Ktze.) Chase in Niles in Contrib. U. S. Nat Herb. 24: 153 (1925). collinum Chase in Hitche. in Contrib. U. S. Nat. Herb. 24: 451 (1927). conjugatum Berg. in Act. Helv. Phys. Math. 7: 129 (1772) conspersum Schrad. ex Schult. Mant. 2: 174 (1824). decumbens Sw. Prodr. 22 (1788). depauperatum Presl, Rel. Haenk. 1: 215 (1830). densum Poir. in Lam, Encye. 5: 32 (1804). distichum L. Syst. Nat. (ed. 10) 2: 855 (1759). Ekmanianum Henr. in Meded. Rijks Herbar. 40: 49 (1921). Humboldtianum Fluegge, Monog. 1: 67 (1810). inaequivalve Raddi, Agrost. Bras. 228 (1823). intermedium Munro ex Morong & Britton in Ann. N. Y. Acad. Sci. 7: 258 (1893). inconstans Chase in Hitche. in Contrib. U. S. Nat. Herb. 24: 446 (1927). iridifolium Poepp. Reise, 2: 324 (1836). Juergensii Hack. in Fedde, Repert. Spec. Nov. 7: 312 (1909). lepidum Chase in Hitche. in Contrib. U. S. Nat. Herb. 24: 447 (1927). lineispathum Mez in Fedde, Repert. Spec. Nov. 15: 27 (1917). macrophyllum HBK. Nov. Gen. & Spec. 1: 92 (1816). malacophyllum Trin. Sp. Gram. 3: t. 271 (1836). marginatum Remy in Ann. Sci. Nat. (ser. 3) 6:348 (1846). melanospermum Desv. in Lam. Encyc. Suppl. 4: 315 (1816). membranaceum Lam. Encyc. Tabl. 1: 177 (1791). membranaceum var. aequiglume Doell in Mart. Fl. Bras. 2(2): 94 (1877). millegrana Schrad. ex Schult. Mant. 2: 175 (1824). minus Fourn. Mex. Pl. 2:6 (1886). multicaule Poir. in Lam. Encyc. Suppl. 4: 309 (1816). notatum Fluegge, Monog. 1: 106 (1810). paniculatum L. Syst. Nat. (ed. 10) 2: 855 (1759). penicillatum Hook. f. in Trans. Linn. Soc. 20: 171 (1847). pictum Ekm. in Ark. Bot. 10(17): 11, t. 1, fig. 6 (1911). pilosum Lam. Encyce. Tabl. 1: 175 (1791). plicatulum Michx. Fl. Bor.-Am. 1: 45 (1803). polyphyllum Nees in Trin. Gram. Pan. 114 (1826). prostratum Scribn. & Merr. in U. S. Dept. Agric. Div. Agrost. Bull. 24: 9 (1901). pygmaeum Hack. in Fedde, Repert. Spec. Nov. 11: 18 (1912). pygmaeum var. glabrescens Hack. in Fedde, Repert. Spec. Nov. 11: 18 (1912) =P. pygmaeum. Regnellii Mez in Fedde, Repert. Spec. Nov. 15: 75 (1917). remotum Remy in Ann. Sci. Nat. (ser. 3) 6: 349 (1846). saccharoides Nees in Trin. Sp. Gram. 1: t. 107 (1828). stellatum Humb. & Bonpl. ex Fluegge, Monog. 1: 62 (1810). tripinnatum Mez in Fedde, Repert. Spec. Nov. 15: 64 (1917). Urvillei Steud. Syn. Pl. Glum. 1: 24 (1854). vinosum Mez in Fedde, Repert. Spec. Nov. 15: 28 (1917) = P. depaup- eratum. virgatum L. Syst. Nat. (ed. 10) 2: 855 (1759). Pennisetum bambusiforme (Fourn.) Hemsl. Biol. Centr. Amer. Bot. 3: 506 (1885). P. chilense (Desv.) Jacks. ex R. E. Fries in Nov. Act. Soc. Sci. Upsal. (ser. 4) 1: 172 (1905). em TOO he foe te i a . ~~ OE ul pl gee RRR! . eo wee . latifolium Spreng. Syst. Veg. 1: 302 (1825). . mutilatum (O. Ktze.) Hack. in O. Ktze. Rev. Gen. 3(2): 347 (1898), i . setosum (Sw.) Pers. Synops. Pl. 1: 72 (18 . tristachyum subsp. boliviense Chase in Hitche. in Contrib. U. S. Nat. Herb. 24: 486 (1927). P. villosum R. Br. in Fresen. in Mus. Senckenb. 2: 134 (1837). Phalaris angusta Nees ex Trin. Sp Gram. 1: t. 78 (1827). Pharus glaber HBK. Nov. Gen. & Spec. 1: 196 (1816). P. latifolius L. Syst. Nat. (ed. 10) 2: 1269 (1759). P. parvifolius Nash in Bull. Torr. Bot. Club, 35: 301 (1908). Phragmites communis (L.) Trin. Fund. Agrost. 134 (1820). Piptochaetium laeve (Nees) Pilg. in Engler, Bot. Jahrb. 56, Beibl. 123: 26 (1920) = Nassella pubiflora. P. setifolium Presl, Rel. Haenk. 1: 222 (1880). P. tuberculatum Desv. in Gay, Fl. Chile, 6: 272 (1853). Poa androgyna Hack. in Fedde, Repert. Spec. Nov. 6: 159 (1908) = P. hor- ie Ee synon ¥. sagittatam Henr. in Blumea, Suppl. 1: 229, a 16 (1937). P 05). $ ula. annua L. Sp. Pl. 68 (1753). asperiflora Hack. in Fedde, Repert. Spec. Nov. 11: 28 (1912). boliviensis Hack. in Fedde, Renert. Spec. Nov. 11: 25 (1912) — P. pra- tensis. bonariensis (Lam.) Kunth, Rév. Gram. 1: 115 (1829). Buchtienii Hack. in Fedde, Repert. Spec. Nov. 11: 29 (1912). Buchtienii var. subacuminata Hack. in Fedde, Repert. Spec. Nov. 11: 30 (1912) = P. Buchtienii. candamoana Pilg. in Engler, Bot. Jahrb. 37: 381 (1906). chamaeclinos Pilg. in Engler, Bot. Jahrb. 37: 379 (1906). denticulata Hack. in Fedde, Repert. Spec. Nov. 11: 27 (1912). dumetorum Hack. in Fedde, Repert. Spec. Nov. 11: 26 (1912) = P. hor- ridula. dumetorum var. unduavensis Hack. in Fedde, Repert. Spec. Nov. 11: 27 (1912) = P. horridula. gymnantha Pilg. in Engler, Bot. Jahrb. 56, Beibl. 123: 28 (1920). gymnantha var. aperta Pilg. in Notizbl. 11: 780 (1933). horridula Pilg. in Engler, Bot. Jahrb. 37: 506 (1906). humillima Pilg. in Engler, Bot. Jahrb. 37: 378 (1906). Lilloi Hack. ex Stuckert in Anal. Mus. Nac. Buenos Aires, 21: 153 (1911). myriantha Hack. ex Stuckert, Segunda Contrib. 517, t. 3 (1906). perligulata Pilg. in Notizbl. 11: 779 (1933 Pflanzii Pilg. in Engler, Bot. Jahrb. 49: 187 (1913) = P. asperiflora. pratensis L. Sp. Pl. 67 (1753). scaberula Hook. f. Fl. Antarct. 2: 378 (1846). subspicata (Presl) Kunth, Rév. Gram. 3: 606 (1829). Ve Cee eee oe ee of et Puccinellia oresigena (Phil.) Hitche. in Contrib. U. S. Nat. Herb. 24: 326 (1927). P. parvula Hitche. in Contrib. U. S. Nat. Herb. 24: 325 (1927). Saccharum cayennense (P. Br.) Benth. in Journ. Linn. Soc. 19: 66 (1881). S. officinarum L. Sp. Pl. 54 (1753). Sacciolepis myuros (Lam.) Chase in Proc. Biol. Soc. Wash. 21: 7 (1908). Setaria gracilis HBK. Nov. Gen. & Spec. 1: 109 (1816). Ss. Ss. S. i 31 gracilis forma brevispica — Hack. in Stuckert in Anal. Mus. Nac. Buenoes Aires, 21: 48 (1911). gracilis forma penicillata (Willd.) Mez ex Ekm. in Ark. Bot. 13(10): 33 (1910). leiocarpa Herrm. in Beitr. Biol. Pflanz. 10: 62 (1910) = Chaetochloa ob- ongata. Sorghastrum minarum (Nees) Hitche. in Contrib. U. S. Nat. Herb. 24: 501 (1927). Ss. Ss. parviflorum (Desv.) Hitche. & Chase in Contrib. U. S. Nat. Herb. 18: 287 (1917). stipoides (HBK).) Nash in N. Am, Fl. 17: 129 (1912). Sporobolus aeneus (Trin.) Kunth, Rév. Gram. 3: 595 (1829). AM Mnnn th - argutus (Nees) Kunth, Enum. Pl. 1: 215 (1833). ) asperifolius Nees & Meyen in Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 141 (1841 + Berteroanus (Trin.) Hitch. & Chase in Contrib. U. S. Nat. Herb. 18: 370 (1917). eximius (Nees) Ekm, in Ark. Bot. 13(10): 41 (1913). indicus (L.) R. Br. Prodr. Fl. Nov. Holl. 1: 170 (1810). Poiretii (Roem. & Schult.) Hitche. in Bartonia, 14: 32 (19382). gee (Mart. & Schrank) Hack. in Bull. Herb. Boiss. (ser. 2) 4(3): 278 (19 Stipa boliviensi Hack. in Fedde, Repert. Spec. Nov. 11: 21 (1912) = obtus py Pen 2 MPp PP pep peep seni siete Hitche. in Contrib. U. S. Nat. Herb. 24: 275 (1925). capilliseta Hitche. in Contrib. U. S. Nat. Herb. 24: 271 (1925). curviseta Hitche. in Contrib. U. S. Nat. Herb. 24: 282 (1925). dasycarpa Hitche. in Contrib. U. S. Nat. Herb. 24: 281 (1925). depauperata Pilg. in Engler, Bot. Jahrb. 56, Beibl. 123: 23 (1920). Hans-Meyeri Pilg. in Engler, Bot. Jahrb. 56, Beibl. 123: 24 (1920). Holwayi Hitche. in Contrib. U. S. Nat. Herb. hk 287 (1927). Ichu (R. & P.) Kunth, Rév. Gram. 1: 60 (18 illimanica Hack. in Fedde, Repert. Spec. hoc oan 22 (1912). inconspicua Presl, Rel. Haenk. 1: 227 (1830). eptothera var. atroviolacea Hack. in Fedde, Repert. Spec. Nov. 6: 154 (1908) = S. depauperata. mucronata HBK. Nov. Gen. & Spec. 1: 125 (1816). nardoides (R. A. Phil.) Hack. ex Hitche. in Contrib. U. S. Nat. Herb. 24: 271 (1925). Neesiana Trin. & Rupr. in Mém. Acad. St. Pétersb. (ser. 6) 5(1): 27 (1842). obtusa (Trin & Rupr.) Hitche. in Contrib. U. S. Nat. Herb. 24: 284 (1925). pampagrandensis Spegazz. in Anal. Mus. Nac. Montevideo, 4: 158, fig. 48 (1901). Pflanzii Mez in Fedde, Repert. Spec. Nov. 17: 206 (1921) — Nassella pubiflora. plumosa Trin. in Mém. Acad. St. Pétersb. (ser. 8 vee 87 (1836). plumosula Nees in Steud. Syn. Pl. Gram. 127 (18 polyclada Hack. ex Stuckert in Anal. Mus. meats Buenos Aires, 21: 80 11). pumila Mez in Fedde, Repert. Spec. Nov. 17: 205 (1921) = S. depauperata. . speciosa Trin. & Rupr. in Mém. Acad. St. Pétersb. (ser. 6) 5(1): 45 (1842). . Trollii Pilg. in Notizbl. 11: 777 (1933). Stylagrostis leiopoda (Wedd.) Mez in Bot. Archiv Mez, 1: 20 (1922 Thrasya campylostochya (Hack.) Chase in Proc. Biol. Soc. ane ‘24: 115 (1911). Trachypogon canescens Nees, Agrost. Bras. 343 (1829). 32 T. montufari (HKB.) Nees, Agrost. Bras. 342 (1829) T. montufari var. bolivianus (Pilg.) Pilg. in eet 11 T1T. 41988) == T. montufari. T. plumosus (Humb. & Bonpl.) Nees, Agrost. Bras. 344 (1829). Tragus alienus (Spreng.) Schult. in Roem. & Schult. Syst. Veg. Mant. 2: 205 (1824). Trichachne saccharata (Buckl.) Nash ex Small, Fl. SE. U. S. 83 (1903). T. sacchariflora (Raddi) Nees, Agrost. Bras. 87 (1829). Trichloris mendozina (Phil.) Kurtz in Mem. Fac. Cienc. Exact. Univ. Cordoba, 1896: 37 (1897). T. plurifiora Fourn. Mex. Pl. 2: 142 (1886). Tricholena rosea Nees, Fl. Afr. Austr. 1: 17 (1841). Trichopteryx flammida (Trin.) Benth. in Journ. Linn. Soc. 19: 59 (1882), combination implied but not actually made. Triniochloa stipoides (HBK.) Hitche. in Contrib. U. S. Nat. Herb. 17: 303 (1913). Triodia avenacea HBK. Nov. Gen. & Spec. 1: 156, t. 48 (1816). Tripogon spicatus (Nees) Ekm. in Ark. Bot, 11(4): 36 (1912). Tripsacum dactyloides (L.) L. Syst. Nat. (ed. 10) 2: 1261 (1759). Trisetum oreophilum Louis-Marie in Rhodora, 30: 221 (1928). T. spicatum (L.) Richt. Pl. Eur. 1: 59 (1890 T. subspicatum (L.) Beauv. Ess. Agrost. 88 (1812). Tristachya chrysothrix Nees, Agrost. Bras. 460 (1829). Zea Mays L. Sp. Pl. 971 (1758). Zeugites mexicana (Kunth) Trin. ex Steud. Nomencl. (ed. 2) 2: 798 case). Z. mexicana var. glandulosa Hack. in Fedde, Repert. Spec. Nov. 6: (1908). CYPERACEAE Ascolepis brasiliensis (Kunth) Benth. ex C. B. Clarke in Dur. & Schinz, Consp. Fl. Afr. 5: 651 (1894). Becquerelia glomerulata Brongn. in Duperrey, Bot. Voy. Coq. 163 (1829). Bulbostylis boliviana Palla in Oesterr. Bot. Zeitschr. 59: 191 (1909). B. capillaris (L.) C. B. Clarke in Hook. Fl. Brit. India, 6: 652 (1893). hirtella (Schrad.) Urb. Symb. Ant. 2: 166 (1900). junciformis (HBK.) C. B. Clarke in Trans. Linn, Soc. (ser. 2) 4: 512 (1895). juncoides (Vahl) Kiikenth. ex Osten in Anal. Mus. Hist. Nat. Montevideo (ser. 2), 3: 187 (1931). juncoides var. ampliceps Kiikenth. ex Osten in Anal. Mus. Hist. Nat. Monte- video (ser. 2), 3: 188 (1931). papillosa Kiikenth. in Fedde, Repert. Spec. Nov. 23: 198 (1926). sphaerocephala (Boeck.) C. B. Clarke in Bull. Herb. Boiss. (ser. 2) 3: 1018 (1903). sphaerocephala var. macrocephala Kiikenth. ex Osten in Anal. Mus. Hist. Nat. Montevideo (ser. 2), 3: 186 (1981). sphaerolepis (Boeck.) Beetle in Amer. Midl. Nat. 41: 487 (1949). tenuifolia (Rudge) Macbride in Field Mus. Publ. Bot. 11: 5 (1931). Calyptrocarya fragifera (Rudge) Nees in Linnaea, 9: 304 (1834). Carex boliviensis v. Heurck & Muell. Arg. in v. Heurck, Obs. Bot. 32 (1870). C. Bonplandii Kunth, Enum, Pl. 2: 380 (18387). C. cladostachya var. maxima Kiikenth. in Engler, Pflanzenr. IV, 20(38): 268 (1909). C. decidua var. Brehmeri (Boeck.) Kiikenth. in Engler, Pflanzenr. IV, 20(38): 307 (1909). C. fecunda Steud. Syn. Cyp. 194 (1855). yo se USS = mh Aga aagagaaa 33 Goodenowii var. stolonifera (Hoppe) Aschers. Fl. Brandenb. 1: 777 (1864). Jamesonii var. subfulva Kiikenth. in Fedde, Repert. Spec. Nov. 8: 7 (1910). lapazensis C. B. Clarke in Kew Bull. Add. Ser. 8: 76 (1908). Mandoniana Boeck. in Allg. Bot. Zeit. no. 11: 174 (1896). nebularum Phil. in Anal. Univ. Chile, 93: 492 (1896). perprava C. B. Clarke in Kew Bull. Add. Ser. 8: 76 (1908). phalaroides var. moesta (Kunth) Kiikenth. in Verh. Bot. Ver. Brandenb. 47: 208 (1905). pichinchensis HBK. Nov. Gen. & Spec. 1: 233 (1816). pichinchensis var. obtusisquama R. Gross in Notizbl. 14: 193 (1938). pinetorum Liebm. in Vidensk. Selsk Skr. Kjobenh. (ser. 2) 5: 263 (1851). Cladium Mariscus R. Br. Prodr. 236 (1810) C. Mariscus subsp. jamaicense var. ferruginescens Kiikenth. in Fedde, Repert. Spec. Nov. 51: 192 (1942). Cyperus albomarginatus Mart. & Schrad. ex Nees in Mart. Fl. Bras. 2(1): 9 em A 2 228A 8Ae 6 6 & Be a6 4 29 A GS a0 (1842) [this is the usual reference given, but actually the name pub- lished was Pycreus albo-marginatus]. . andinus Palla ex Kiikenth. in Engler, Pflanzenr. IV, 20 (101): 310 (1936). Andreanus var. capitinduensis (Maury) Kiikenth. in Engler, Pflanzenr. IV, 20(101): 212 (1936) aristatus var. inflexus (Muehl.) Boeck. in Linnaea, 35: 500 (1868), combin- ation implied but not actually made. articulatus var. nodosus ( Willd.) Kiikenth. in Engler, Pflanzenr. IV, 20(101): 79 (1935). Bangianus Gandoger in Bull. Soc. Bot. France, 66: 297 (1920). pen var. boliviensis Kiikenth. in Engler, Pflanzenr. IV, 20(101): 422 (19 Potbesen “Lam.) Britton in Bull. Dept. Agric. Jamaica, 5, Suppl. 1: 8 (1907). compressus L, Sp. Pl. 46 (1753). corymbosus var. subnodosus (Nees & Mey.) Kiikenth. ex Osten in Anal. Mus. Hist. Nat. Montevideo (ser. 2), 3: 147 (1931). densicaespitosus Mattf. & Kiikenth. in Engler, Pflanzenr. IV, 20(101): 597 (1936). usus subsp. chalaranthus (Presl) Kiikenth. in Engler, Pflanzenr. IV, 20(101): 210 (1936). diffusus subsp. chalaranthus var. tolucensis (HBK.) Kiikenth. in Engler, Pflanzenr. IV, 20(101): 211 (1936). diffusus subsp. chalaranthus var. umbrosus (Lindl. & Nees) Kiikenth. in Engler. Pflanzenr. IV, 20(101): 210 (1936). digitatus var. obtusifructus Kiikenth. in Engler, Pflanzenr. IV, 20(101): 56 (1935). Eggersii Boeck. Cyper. Nov. 1: 53 (1888). elegans L. Sp. Pl. 45 (1753). entrerianus var. parvicapitulus Kiikenth. in Engler, Pflanzenr. [IV( 20(101): 170 (1936). Eragrostis Lam. Encyc. Tabl. 1: 146 (1791). esculentus var. leptostachyus Boeck. in Linnaea, 36: 290 (1870). ferax L. C. Rich. in Act. Soc. Hist. Nat. Paris, 1: 106 (1792). flavus var. aggregatus (Willd.) Kiikenth. in Engler, Pflanzenr, IV, 20(101): 582 (1936). flavus var. gigas (Lindm.) Kiikenth. in Engler, Pflanzenr. IV, 20(101): 532 (1936). flavus subsp. redolens (Maury) Osten in Anal. Mus. Hist. Nat. Montevideo (ser. 2), 3: 152 (19381) friburgensis Boeck. Cyper. Nov. 2: 2 (1890). eo c= 2-2 2 2 A RAH B9S G2 AH 6 AA: AnAgaaa ann e ae a ang AAD giganteus Vahl, Enum. 2: 364 (1806). aspan var. americanus Boeck. in Linnaea, 35: 575 (1868). Haspan se juncoides (Lam.) Kiikenth. in Fedde, Repert. Spec. Nov. 23: 184 (1926). hermaphroditus (Jacq.) Standl. in Contrib. U. S. Nat. Herb. 18: 88 G58); sera ao a var. condensatus Kiikenth. in Fedde, Repert. Spec. Nov. 2 251 (192 soa var. Fiebrigii (Kiikenth.) Kiikenth. in Engler, Pflanzenr. IV, 20(101): 184 (1936). incomtus Kunth, Enum. Pl. 2: 39 (1837). incomtus var. dissolutior Kiikenth. in Engler, Pflanzenr. IV, 20(101): 184 (1936). incomtus var. Miguelii Kiikenth. in Engler, Pflanzenr. IV, 20(101): (1936). laevigatus L. Mant. 2: 179 (1771). laevigatus var. distachyus (All.) Coss. & Durieu, Fl. Algér. 2: 251 (1854). laevigatus var. reptans (Boeck.) C. B. Clarke in Journ. Linn. Soc. 21: 78 lanceolatus Poir. in Lam. Encyc. 7: 245 (1806). lanceolatus var. compositus Presl, Rel. Haenk, 1: 167 (1830). ligularis L. Pl. Jamaic. Pugill. 3 (1759). luzulae (L.) Retz, Obs. 4:11 (1786). megapotamicus Kunth, Enum. Pl. 2: 10 (1837). Meyenianus Kunth, Enum. Pl. 2: 88 (1837) miliifolius var. saturatus (Donn. Sm.) Kiikenth. in Engler, Pflanzenr. IV, 20(101): 221 (1936). Mutisii (HBK.) Griseb. Fl. Brit. W. Ind. 567 (1864). niger R. & P. Fl. Peruv. 1: 47 (1789). niger var. intricato-ramosus (Boeck.) Kiikenth. in Engler, Pflanzenr. IV, 20(101): 345 (1936). niger var. Lorentzianus (Boeck.) Kiikenth. in Engler, Pflanzenr. IV, 20(101): 345 (1936). obtusatus (Presl) Mattf. & Kiikenth. in Engler, Pflanzenr. IV, 20(101): 585 (1936). ochraceus var. excelsior Kiikenth. in Engler, Pflanzenr. IV, 20(101): 182 (1936). Pearcei C. B. Clarke in Kew Bull. Add. Ser. 8: 7 ie phaeocephalus Griseb. in Goett. Abh. 19: 214 (187 phaeocephalus var. major Kiikenth. in Engler, Ebon IV, 20(101): 464 (1936). piceus Liebm. in Vidensk. Selsk. Skr. Kjobenh. 5(2): 200 (1851). prolixus HBK. Nov. Gen, & Spec. 1: 206 (1816). reflexus Vahl, Pian: 2: 299 (1806). ae var. australis (Lindm.) Kiikenth. in Engler, Pflanzenr. IV, 20(101): (1936). "252 (102 var. rufidulus (Steud.) Kiikenth. in Fedde, Repert. Spec. Nov. 26: 29). peer = tarijensis Kikenth. in Engler, Pflanzenr. IV, 20(101): 444 rivularis subsp. lagunetto (Steud.) Kiikenth. in Engler, Pflanzenr. IV, 20(101): 383 (1936). seslerioides HBK. Nov. Gen. & Spec. 1: 209 (1816). sesquiflorus (Torr.) Mattf. & Kiikenth. in Engler, Pflanzenr. IV, 20(101): 591 (1936). simplex HBK. Nov. Gen. & Spec. 1: 209 (1816). sphacelatus Rottb. Descr. & Icon. 26 (1773). surinamensis Rottb. Descr. & Icon. 35, t. 6, fig. 5 (1773). surinamensis var. lutescens Boeck. in Linnaea, = 555 (1868). tabina Steud. ex Boeck. in Linnaea, 35: 559 (18 uncinulatus Schrad. ex Nees in Mart. Fl. Bras. eh 23 (1842). unioloides R. Br. Prodr. Fl. Nov. Holl. 216 (1810). vegetus var. obtusangulus O. Ktze. Rev. Gen. 3(2): 334 (1898) = C. Era- agaaana grostis. Dichiumena ciliata Vahl, Enum. 2: 240 (1806). D. Kuntzei (Clarke) Macbride in Field Mus. Publ. Bot. 11:42 (1931). D. Mandonii (Clarke) Macbride in Field Mus. Publ. Bot. 4: 167 (1929). ‘ D. monostachya (Boeck.) C. B. Clarke in Kew Bull. Add. Ser. 8: 32 (1908). D. nervosa Vahl, Enum. 2: 241 (1806). Diplacrum longifolium (Griseb.) C. B. Clarke in Dur. & Schinz, Consp. FI. Afr. 5: 669 (1894 Eleocharis sii thinctlele Nees & Meyen ex Kunth, Enum. Pl. 2: 143 (1837). Brehmeriana Boeck. in Allg. Bot. Zeit. no, 11: 33 (1896). capitata (L) R. Br. Prodr. Fl. Nov. Holl. 225 (1810). caribaea (Rottb.) Blake in Rhodora, 20: 24 (1918). costulata Nees & Meyen ex Kunth, Enum. Pl. 2: 142 (1887). crinalis (Griseb.) C. B. Clarke in Kew Bull. Add. Ser. 8: 23 (1908). Dombeyana Kunth, Enum. Pl. 2: 145 (1937). exigua (HBK.) Roem. & Schult. Syst. Veg. 2: 154 (1817). filiculmis Kunth, Enum. Pl. 2: 144 (1837). fistulosa (Poir.) Schult. Mant. 2: 89 (182 geniculata (L.) Roem. & Schult. Syst. Veg. ‘2: 150 (1817). interstincta (Vahl) Roem. & Schult. Syst. Veg. 2: 149 (1817). maculosa R. Br. Prodr. Fl. Nov. Holl. 224 (1810). minima Kunth, Enum. Pl. 2: 139 (18387). nodulosa (Roth) Schult. Mant. 2: 87 (1824). nubigena C. B. Clarke in Engler, Bot. Jahrb. 37: 518 (1906). ochreata var. flaccida (Spreng.) Boeck. in Urb. Symb. Ant. 2: 63 (1900). . Sellowiana Kunth, Enum. Pl. 2: 149 (1837). suleata (Roth) Nees in Linnaea, 9: 294 (1834) [combination was not actu- ally made here]. Fimbristylis aestivalis Vahl, Enum, 2: 288 (1806). F. annua (All.) Roem. & Schult. Syst. Veg. 2: 95 (1817). F. monostachya (L.) Hassk. Pl. Jav. Rar. 61 (1848). Fuirena umbellata Rottb. Descr. & Icon. 70, t. 19, fig. 3 (1773). Lipocarpha Sellowiana Kunth, Enum. Pl, 2: 267 (1837). Pleurostachys puberula ope Buchtienii (Kiikenth.) Kiikenth. in Fedde, Repert. Spec. Nov. 26: 254 (1929). Shrachiisers andina Kiikenth. in Fedde, Repert. Spec. Nov. 53: 73 (1944). aristata Boeck. in Flora, 40: 36 (1857). aristata var. latilaminata Kiikenth. in Fedde, Repert. Spec. Nov. 26: 253 (1929). boliviensis C. B. Clarke in Kew Bull. Add. Ser. 8: 37 (1908). confinis (Nees) Clarke in Kew Bull. Add. Ser. 8: 40, 119 (1908). corymbosa (L.) Britton in Trans. N. Y. Acad. 11: 84 (1892). cyperoides (Sw.) Mart. in Denkschr. Akad. Muench. 6: 149 (1820). emaciata (Nees) Boeck. in Vidensk. Meddel. Kjoebenh. 1869: 149 (1870). exaltata var. cophalophora (Nees) Kiikenth. in Engler, Bot. Jahrb. 74: 440 (1949). exaltata var. ovalis Kiikenth. in res Bot. Jahrb. 74: 440 (1949). glauca Vahl, Enum. 2: 233 (18 globosa (HBK.) Roem & ee roe Veg. 2. 89 (1817). PRR Et Bt Bt bt et BB a Pir WR A WP 36 R. globosa var. Loefgrenii (Boeck.) Kiikenth. in Engler, Bot. Jahrb. 74: 465 1949). R. hirta (Nees) Boeck. in Vidensk. Meddel. Kjoebenh. 1869: 146 (1870). R. Lundii Boeck. in Vidensk. Meddel. Kjoebenh. 1869: 147 (1870). luzuliformis Boeck. in Linnaea, 37: 632 (1873). macrochaeta Steud. ex Boeck. in Linnaea, 37: 632 (1878). macrochaeta var. colombiensis forma condensata Kiikenth. in Engler, Bot. Jahrb. 74: 394 (1949 macrochaeta var. Guthiniedatiate (Boeck.) Kiikenth. in Engler, Bot. Jahrb. 74: 394 aceaita a Ruiziana (Boeck.) Kiikenth. in Engler, Bot. Jahrb. 74: 393 (1949). Marisculus Nees in Linnaea, 9: 297 (1834). R. polyphila Vahl, Enum. 2: 230 (1806). R. polyphylla var. longispiculosa Kiikenth. in Engler, Bot. Jahrb. 74: 406 94 nm 2 Fee (1 R. radicans (Schlechtd. & Cham.) H. Pfeiff. in Fedde, Repert. Spec. Nov. 38: 3 (19 R. honed (Schlechtd.) Kunth, Enum, Pl. 2: 300 (1837). R. setacea (Berg.) Boeck. in Vidensk. Meddel. Kjoebenh. 1869: 159 (1870). R. tenerrima Nees ex Spreng. Syst. Veg. 4: Cur. Post. 26 (1827 <8 R. tenuis Link in Sprengel, Schrader & Link, Jahrb. 1(3): 76 (182 R. tenuis var. emaciata (Boeck.) Lindm. in Bihang Kgl. Sv. eile Handl. 26, Afd. 3(9): 28 (1900). R i var. Kuntzei (C. B. Clarke) Kiikenth. in Engler, Bot. Jahrb. 74: 391 (1949). R. velutina var. Sellowiana (Kunth) Boeck. in Linnaea, 37: 614 (1873). irpus americanus Pers. Synops. Pl. . 68 (1805). asper Presl, Rel. Haenk. 1: 194 (1830). atacamensis (Phil.) O. Ktze. Rev. Gen. 2: 757 (1891). cernuus Vahl, Enum. 2: 245 (1806). deserticola Phil .F1. hee 53 (1860). inundatus Poir. in Lam, Encyc. — 5: 103 (1817). micranthus Vahl, Enum. 2: 254 (18 rigidus Boeck. in Linnaea, 36: 492 (1869-1870). riparius Presl, Rel. Haenk. 1: 193 (1830). Scleria bracteata Cav. Icon. 5: 34, t. 457 (1799). Ss. por forma simplicior Kiikenth. in Fedde, Repert. Spec. Nov. 26: 253 (1929 bres Core in Brittonia, 1: 239 (1934). cyperina var. dentata H. Pfeiff. in Fedde, Repert. Spec. Nov. 52: 169 (1943). hirtella Sw. Prodr. 19 (1788). lithosperma (L.) Sw. Prodr. 18 (1788). macrogyne C. B. Clarke in Kew Bull.. Add. Ser. 8: 59 (1908). macrophylla Presl, Rel. Haenk. 1: 200 (1830). microcarpa Nees in Linnaea, 9: 302 (1834), nomen. nud mitis Berg. in Vet. Akad. Handl. Stockh. 27: 145, t. 5 (1765). obtusa Core in Brittonia, 1: 240 (1934). paludosa Poepp. & Endl. ex Kunth, Enum. Pl. 2: 344 (1837). pleostachya Kunth, Enum. Pl. 2: 355 (1837). reflexa HBK. Nov. Gen. & Spec. 1: 232 (1816). secabra Willd. Sp. Pl. 4(1): 315 (1805). stipularis Nees in Hook. Journ. Bot. 2: bm (1840). vaginata Steud. Syn. Pl. Cyp. 179 (185 Uncinia hamata (Sw.) Urb. Symb. Ant. 2: a (1900). U. phleoides (Cav.) Pers. Synops. Pl. 2: 534 (1806). . oho hohe poecrrree ee PALMAE Acrocomia Totai Mart. Palmet. Orbign. 78, t. 9, fig. 1, t. 29B (1847). Aiphanes caryotifolia (HBK.) Wendl. Kerch. Palm. 230 (1878). A. truncata (Brongn. ex Mart.) Wendl. Kerch. Palm 230 (1878). Astrocaryum Chonta Mart. Palmet. Orbign. 84, t. 4, fig. 1, t. 29C (1847). A. Huaimi Mart. Palmet. Orbign. 86, t. 18, fig. 3, t. 30A (1847). Bactris faucium Mart. Palmet. Orbign. 60, t. 6, fig. 2, t. 28B (1847). B. inundata Mart. Palmet. Orbign. 58, t. 7, fig. 3, t. 27B (1847). Ceroxylon pityrophyllum Mart. ex Wendl. Kerch. Palm. 239 (1878). Chamaedorea boliviensis Dammer in Notizbl. 6: 262 (1915). C. conocarpa Mart. Palmet. Orbign. 6, t. 6, fig. 1, t. 16B (1847 ); C. fragrans (R. & P.) Mart. Hist. Nat. Palm. 2: 4, t. 3, fig. 1-2 (1823). C. lanceolata (R. & P.) Kunth, Enum. Pl. 3: 172 (1841). C. macroloba Burret in Notizbl. 11: 757 (1933). Copernicia australis Becc. in Webbia, 2: 158 (1907). Desmoncus Kuhlmannii Burrett in Notizbl. 14: 267 (1938). D. latisectus Burret in Fedde, Repert. Spec. Nov. 36: 215 (1934). D. rudentum Mart. Palmet. Orbign. 48, t. 14, fig. 3, t. 26A (1847). Dictyocaryum Lamarckianum (Mart.) Wendl. in Bot. Zeit. 21: 131 (1863). Diplothenium maritimum Mart. Hist. Nat. Palm. 2: 108, t. 75, t. 77, fig. 3 26). Euterpe andicola Brongn. ex Mart. Palmet. Orbign. 8, t. 2, fig. 2, t. 17A (1847). E. Haenkeana Brongn. ex Mart. Palmet. Orbign. 9, t. 2, fig. 3, t. 17B (1847). G. Desmarestii Mart. Palmet. Orbign. 23, t. 11, fig. 3, t. 22B (1847). G. Jussieuana Mart. Palmet. Orbign. 24, t. 12, fig. 2, t. 23A ( 1847). Pyrenoglyphis Brongniartii (Mart.) Burret in Fedde, Repert. Spec. Nov. 34: 251 (1934). P. infesta (Mart.) Burret in Fedde, Repert. Spec. Nov. 34: 248 (1934). P. socialis (Mart.) Burret in Fedde, Repert. Spec. Nov. 34: 246 (1934). Scheelea blepharopus (Mart.) Burret in Notizbl. 10: 674 (1929). S. princeps (Mart.) Karst. in Linnaea, 28: 269 (1856). Socratea Orbignyana (Mart.) Karst. in Linnaea, 28: 264 (1856). Syagrus petraea (Mart.) Becc.in L’Agric. Colon. 10(2): 467 (1916). Taenianthera macrostachys (Mart.) Burret in Engler, Bot. Jahrb. 63: 268 (1930). Tessmanniophoenix Chuco (Mart.) Burret in Notizbl. 10: 400 (1928). Trithrinax schizophylla Drude in Mart. Fl. Bras. 3(2): 551, t. 130 (1882). 38 CYCLANTHACEAE Carludovica palmata R. & P. Syst. 291 (1798). ARACEAE Anthurium apiculatum Krause in Engler, Bot. Jahrb. 44, Beibl. 101: 9 (1910). Buchtienii Krause in Engler, Bot. Jahrb. 44, Beibl. 101: 10 (1910). she bate Krause in Notizbl. 11: 612 (1932). oripatense Engl. in Engler, Pflanzenr. ae 23b: 255 (1905) — A. Miguelii. Sracile Lindl. in Bot. Reg. 19: t. 1635 (1833). grande N. E. Br. in Engler, Pflanzenr. IV, 23b: 204 (1905). indecorum Schott in Oesterr. Bot. Zeitschr. = 350 (1858). Lechlerianum Schott, Prodr. Aroid. 534 (18 mapiriense Krause in Engler, Bot. Jahrb. - fan 101: 10 (1910). Miguelii Engl. in Engler, Pflanzenr. IV, 23b: 126 (1905). Ottonis Krause in Notizbl. 11: 609 (1932). paraguayense Engler, Bot. Jahrb. 25: 361 (1898). parile N. E. Br. in Engler, Pflanzner. IV, 23b: 151 (1905). rusticum N. E. Br. in Engler, Pflanzenr. IV, 23b: 82 (1905). scandens Engl. in Mart. Fl. Bras. 3(2): 78 (1878). scandens var. violaceum Engl. in Mart. Fl. Bras. 3(2): 78, t. 7 (1878). trinerve Miq. in Linnaea, 17: 66 (1843). trinerve var. angustifolium Krause in Engler, Bot. Jahrb. 54, Beibl. 118: 123 (1916). triphyllum Brongn. ex Schott, Prodr. Aroid. 548 (1860). violaceum (Sw.) Schott, Prodr. Aroid. 437 (1860). Asterostigma Pavonii Schott, Prodr. Aroid. 339 (1860). Caladium sororium Schott in Oesterr. Bot. Zeitschr. 9: 38 (1859). Heteropsis boliviana Rusby in Bull. N. Y. Bot. Gard. 6: 493 (1910). Monstera boliviana Rusby in Bull. N. Y. Bot. Gard. 6: 494 (1910). M. falcifolia Engl. in Engler, Bot. Jahrb. 37: 117 (1905). M. unilatera Rusby in Bull. N. Y. Bot. Gard. 6: 494 (1910). Philodendron arcuatum Krause in Engler, Pflanzenr. IV, 23Db: 72 (1913). Brandtianum Krause in Engler, Pflanzenr. IV, 23Db: 46 (1913). Buchtienii Krause in Engler, Pflanzenr. IV, 23Db: 30 (1913). caudatum Krause in Engler, Pflanzenr. IV, aavie 13 (1918). Lechlerianum Schott, Broly. Aroid. 250 (1860). . Paxianum Krause in Engler, Pflanzenr. IV, 23Db: 31 (1913). rubens Schott, Syn. Aroid. 84 (1856 ? P tripartitum (Jacq.) Schott in Wien. Zeitschr. 3: 780 (1829). Pistia Stratiotes L. Sp. Pl. 963 (175 Rhodospatha boliviensis Engl. & sa in Engler, Bot. Jahrb. 44, Beibl. 101: 13 (1910). Seaphispatha gracilis Brongn. ex Schott, Prodr. Aroid. 214 (1860). Spathanthium heterandrum (Baker) N. E. Br. in Gard. Chron. 20: 70 (1883). S. Orbignyanum Schott in Bonplandia, 7: 165 (1859). Spathicarpa hastifolia Hook. Bot. Misc. 2: 147 (1831). Stenospermation Mathewsii Schott, Gen. Aroid. 70 (1858). S. Rusbyi N. E. Br. in Bull. N. Y. Bot Gard. 4: 461 (1907). Synandrospadix vermitoxicum (Griseb.) Engl. in Engler, Bot. Jahrb. 4: 62 1883). Ge ee re OO Taccarum caudatum Rusby in Mem. N. Y. Bot. Gard. 7: 210 (1927). T. Weddellianum Brongn. ex Schott, Gen. Aroid. t. 65 (1858). Xanthosoma Buchtienii Engl. in Engler, Pflanzenr. IV, 23E: 54 (1920). X. hylaeae Engl. & Krause in Notizbl. 6: 115 (1914). 39 X. mafaffa var. Poeppigii (Schott) Engl. in Mart. Fl. Bras. 3(2): 193 (1878). X. roseum Schott in Oesterr. Bot. Zeitschr. 8: 178 (1858). X. syngoniifolium Rusby in Mem. N. Y. Bot. Gard. 7: 209 (1927). LEMNACEAE Lemna gibba L. Sp. Pl. 970 (1753). L. valdiviana Phil. in Linnaea, 33: 239 (1864). CENTROLEPIDACEAE Gaimardia boliviana Pax in Fedde, Repert. Spec. Nov. 5: 225 (1908). MAYACACEAE Mayaca boliviana Rusby in Mem. N. Y. Bot. Gard. 7: 211 (1927) = M. Sello- wiana. M. Sellowiana Kunth, Enum. Pl. 4: 32 (1848). XYRIDACEAE Xyris caroliniana Walt. Fl. Carol. 69 (1788). . macrocephala Vahl, Enum. 'Pl. 2: 204 (1806). savanensis Mig. in Linnaea, 18: 605 (1844). simulans Alb. Nilsson in Kgl. Sv. Vet. Akad. Handl. 24 (14): 47 (1892). subulata R. & P. FI. Peruv. 1: 46, t. 7, fig. 6 (1798). 4 4 Pd ERIOCAULACEAE Eriocaulon Steinbachii (Moldenke) Moldenke in Phytologia, 2(9): 364 (1947). Paepalanthus chiquitensis Herzog in Fedde, Repert. Spec. Nov. 20: 86 (1924). P. manicatus var. pulvinatus Herzog in Fedde, Repert. Spec. Nov. 20: 86 (1924). P. muscosus Koern. in Mart. Fl. Bras. 3(1): 348 (1863). Syngonanthus caulescens (Poir.) Ruhl. in Engler, Pflanzenr. IV, 30: 267 (1903). S. Fischerianus (Bong.) Ruhl. in Engler, Pflanzenz. IV, 30: 256 (1903). S. gracilis var. bolivianus Ruhl. in Engler, Pflanzenr. IV, 30: 252 (1903). BROMELIACEAE Abromeitiella brevifolia (Griseb.) Castellanos in Anal. Mus. Nac. Hist. Nat. Buenos Aires, 36: 371, tt. 2, 3 (1931). Aechmea angustifolia Poepp. & Endl. Nov. Gen. & Spec. 2: 43, t. 159 (1838). Ae. boliviana Rusby in Bull. N. Y. Bot. Gard. 4: 456 (1907) — Ae. angusti- folia. Ae. brachyclada Baker, Handbk. Bromel. 87 (1889). Ae. bromeliaefolia (Rudge) Baker ex Benth. & Hook. f. Gen. Pl. 3: 664 (1883). Ae Castelnavii Baker, Handbk. Bromel. 39 (1889). Ae. distichantha var. Schlumbergeri E. Morr. ex Mez in Mart. Fl. Bras. 3(3): 843 (1892). Ae. ellipsoidea Rusby in Mem. N. Y. Bot. Gard. 7: 212 (1927) — Ae. brom- eliaefolia. Ae. inconspicua Harms in Notizbl. 10: 786 (1929) — Ae. angustifolia. Ae. involucrata Rusby in Bull. N. Y. Bot. Gard. 4: 456 (1907) = Ae. distich- antha var. Schlumbergeri. Ae. involucrifera Mez in Enler, Pflanzenr. IV, 32(1): 157 (1934) = Ae. dis- tichantha var. Schlumbergeri. 40 Ae. Kuntzeana Mez in DC. Monog. Phan. 9: 208 (1896). Ae. tocantina Baker, Handbk. Bromel. 39 (1889 Ananas ananassoides (Baker) L. B. Smith in Bot. Mus. Leafl. Harvard Univ. 7: 79 (1939). Billbergia boliviensis Baker, Handbk. Bromel, 81 (1889) — B. decora. B. Cardenasii L. B. Smith in Phytologia, 4(6): 382 (1953). B. decora Poepp. & Endl. Nov. Gen. & Spec. 2: 42, t. 157 (1888). Bromelia Hieronymi Mez. in Mart. Fl. Bras. 3(3): 199 (1891). B. serra Griseb. in Goett. Abh. 24: 328 (1879). Cottendorfia Rusbyi Baker in Bull. Torr. Bot. Club, 29: 697 (1902) = Lind- mania villosula. Deuterocohnia strobilifera Mez in Fedde, Repert. Spec. Nov. 3: 15 (1906). D. strobilifera var. inermis L. B. Smith in Contrib. U. S. Nat. Herb. 29: 535 (1954). Dyckia boliviensis Mez in DC. Monog. Phan. 9: 524 (1896). D. gracilis Mez in DC. Monog. Phan. 9: 516 (1896). ? D. leptostachya Baker in Gard. Chron. 22: 198 (1884). D. Meziana O, Ktze. Rev. Gen. 3(2): 302 (1898). D. pulquinensis Wittm. in Meded. Rijks Herbar. 29: 88 (1916). Guzmania calothyrsa (Beer) Mez in DC. Monog. Phan. 9: 910 (1896). complanata Wittm. in Meded. Rijks Herbar. 29: 92 (1916) — Tillandsia disticha. lingulata (L.) Mez in DC. Monog. Phan. 9: 899 (1896). Melinonis Regel in Gartenfl. 34: 116 (1885). obtusa Rusby in Mem. N. Y. Bot. Gard. 7: 212 (1927) = Vriesia heliconi- Q 2 Qa oides Roezlii (Morr.) Mez in DC. Monog. Phan. 9: 948 (1896). Lindmania gracilis (Rusby) L. B. Smith in Contrib. Gray Herb. 104: 78 (1934). L. graminea L. B. Smith in oe = 93 (1948). L. Pearcei (Baker) Mez in DC. Monog. Phan. 9: 537 (1896). L. pendulifiora (C. H. Wright) Stan i in Bot. Mag. 150: - 9029 (1924). L. Rusbyi Mez in Engler, Bot. Jahrb. 30, Beibl. 67 :6 (1901). L. villosula Harms in Notizbl. 10: 794 (1929 L. Weberbaueri Mez in Fedde, Repert. Spec. Nas: 12: 417 (1913). L. Weddelliana (Brongn.) Mez in DC. Monog. Phan. 9: 538 (1896). Pitcairnia Bangii Baker in Mem. Torr. Bot. Club, 6: 124 (1896) — Puya stenothyrsa. P. biattenuata Rusby in Bull. N. Y. Bot. Gard. 4: 457 (1907) = P. paniculata. P. Brittoniana Mez in DC. Monog. Phan. 9: 451 (1896). P. consimilis Baker in Journ. Bot. 19: 266 (1881) = P. ferruginea. P. crassa L. B. Smith in Lilloa, 14: 94 (1948). P. divaricata Wittm. in Meded. Rijks Herbar. 29: 81 (1916). P. ferruginea R. & P. Fl. Peruv. 3: 36 (1802). P. grandiflora Mez. in Fedde, Repert. Spec. Nov. 3: 5 (1906). P. mirabilis Mez in Fedde, Repert. Spec. Nov. 3: 6 (1906). P. multiramosa Mez in DC. Monog. Phan. 9: 419 (1896). P. odontopoda Baker, Handbk. Bromel. 93 (1889). P. paniculata R. & P. Fl. Peruv. 3: 36, t. 260 (1802). P. platystemon Mez in DC. Monog. Phan. 9: 421 (1896). rigida Mez in Bull. Herb. Boiss. (ser. 2) 4: 625 (1904). P. robusta Rusby in Bull. N. Y. Bot. Gard. 6: 488 (1910) — Puya sanctae- crucis. P. sessiliflora Rusby in Bull. N. Y. Bot. Gard. 4: 457 (1907) = P. subpeti- olata. P. subpetiolata Baker in Journ. Bot. 19: 267 (1881). 41 Pseudananas sagenarius (Arruda) Camargo in Rev. Agric. Piracicaba, 14 (7, 8): reprint p. 4 (1939). Puya alba L. B. Smith in Phytologia, 5: 49 (1954). P. atra L. B. Smith in Contrib. U. S. Nat. Herb. 29: 536 (1954). P. boliviensis Baker, Handbk. Bromel. 126 (1889). P. Brittoniana Baker, Handbk. Bromel. 124 (1889). P. Cardenasii L. B. Smith in Lilloa, 14: 94 (1948). P. cristata L. B. Smith in Lilloa, 14: 95 (1948). P. ctenorhyncha L. B. Smith in Phytologia, 5: 49 (1954). P. dyckioides (Baker) Mez in DC. Monog. Phan. 9: 486 (1896). P. Fiebrigii Mez. in Fedde, Repert. Spec. Nov. 3: 9 (1906). P. Fosteriana L. B. Smith in Journ.. Wash. Acad. Sci. 40: 216 (1950). P. glabrescens L. B. Smith in Contrib. U. S. Nat. Herb. 29: 537 (1954). P. glareosa L. B. Smith in Lilloa, 14: 96 (1948). P. Hauthalii Mez in Fedde, Repert. Spec. Nov. 16: 65 (1919). P. Herzogii Wittm. in Meded. Rijks Herbar. 29:86 (1916). P. Hofstenii Mez. in Fedde, Repert. Spec. Nov. 3: 8 (1906). P. humilis Mez in DC. Monog. Phan. 9: 498 (1896). P. Kuntzeana Mez in DC. Monog. Phan. 9: 490 (1896). P. lasiopoda L. B. Smith in Proc. Amer. Acad. 70: 153 (1935). P. leptostachya L. B. Smith in Lilloa, 14: 97 (1948). P. Meziana Wittm. in Meded. Rijks Herbar. 29: 85 (1916). P. micrantha Mez in Fedde, Repert. Spec. Nov. 3: 8 (1906). P. mollis Baker ex Mez in DC. Monog. Phan. 9: 488 (1896). P. nana Wittm. in Meded. Rijks Herbar. 29: 85 (1916). P. olivacea Wittm. in Meded. Rijks Herbar. 29: 83 (1916). P. paupera Mez in Fedde, Repert. Spec. Nov. 3: 14 (1906). P. Pearcei (Baker) Mez in DC. Monog. Phan. 9: 480 (1896). P. penduliflora L. B. Smith in Contrib. Gray Herb. 98: 12 (1932). P. Raimondii Harms in Notizbl. 10: 213 (1928). P. Rusbyi (Baker) Mez in DC Monog. Phan. 9: 482 (1896). P. sanctae-crucis (Baker) L. B. Smith in Proc. Amer. Acad. 70: 154 (1935). P. spathacea (Griseb.) Mez in DC. Monog. Phan. 9: 481 (1896). P. stenothyrsa (Baker) Mez in DC. Monog. Phan. 9: 481 (1896). P. tuberosa Mez in DC. Monog. Phan. 9: 483 (1896). P. tunarensis Mez in DC. Monog. Phan. 9: 498 (1896). P. ultima L. B. Smith in Contrib. U. S. Nat. Herb. 29: 540 (1954). P. Weddelliana (Baker) Mez in DC. Monog. Phan. 9: 475 (1896). P. Werdermannii Harms in Notizbl. 10: 793 (1929). Tillandsia rappel a Tonduziana (Mez) L. B. Smith in Contrib. Gray Herb. 89: 8 (19 T. apoloénsis aes in Bull. N. Y. Bot. Gard. 6: 489 (1910) = T. cha bala T. australis Mez in Fedde, Repert. Spec. Nov. 16: 75 (1919) = T. maxima. T. Bakeri L. B. Smith in Contrib. Gray Herb. 95: 45 (1931). T. Bangii Baker in Mem. Torr. Bot. Club, 6: 124 (1896) = T. Deppeana. T. biflora R. & P. Fl. Peruv. 3:41, t. 268 (1802). T. boliviana Mez in Bull. Herb. Boiss. (ser. 2) 4: 1130 (1904). T. boliviensis Baker in Mem. Torr. Bot. Club, 4(3): 267 (1895). T. Brittoniana Baker, Handbk. Bromel. 195 ~— T. bryoides Griseb. in Goett. Abh. 24: 334 (1879 T. Buchtienti H. Winkl. in Fedde, Repert. Ah ov. 7: 207. (1909) = T. Rusbyi zy! aliakals Wittm. in Meded. Rijks Herbar. 29: 90 (1916). T. capillaris forma cordobensis (Hieron.) L. B. Smith in Proc. Amer. Acad. 70: 211 (1935). rs bo a 7’. Q capillaris forma Siceoriant (Mez) L. B. Smith in Proc. Amer. Acad. 70: 213 (1935). capillaris forma incana (Gill.) L. B. Smith in Proc. Amer. Acad. 70: 211 935). capillaris forma virescens (R. & P.) L. B. Smith in Proc. Amer. Acad. 70: 212 (1935). Cardenasii L. B. Smith in Proc. Amer. Acad. 70: 154 (1985). caulescens Brongn. ex Baker, Handbk. Bromel. 168 (1889). complanata Benth. Bot. Voy. Sulphur, 173 (1846). condensata Baker, Handbk. Bromel. 164 (1889). crocata (E. Morr.) Baker in Journ. Bot. 25: 214 (1887). decomposita Baker, Handbk. Bromel. 168 (1889). Deppeana Steud. Nomencl. (ed. 2) 2: 688 (1841). didisticha Baker in Journ. Bot. 26: 16 (1888). Duratii Vis. in Nuov. Sagg. Padov. 5: 271 (1841). flexuosa (Baker) Mez in DC. Monog, Phan. 9: 774 (1896) = T. Bakeri. Friesii Mez in Fedde, Repert. Spec. Nov. 3: 37 (1906). funebris Castellanos in Anal. Mus. Nac. Hist. Nat. Buenos Aires, 37: 502 (1933). fusco-guttata Mez in Bull. Herb. Boiss. (ser. 2) 5: 101 (1905). Gilliesii Baker in Journ. Bot. 16: 240 (1878 entheri Harms in Notizbl. 10: 794 (1929) — = T. Rusb Herzogii Wittm. in Meded. Rijks Herbar. 29: 89 (agi6) = = T. Reichen- bachii. incurvata Griseb. in Goett. Nachr. 1864: 15. ixioides Griseb. in Goett. Abh. 24: 333 (1879). juncea (R. & P.) Poir. in Lam. Encyc. Suppl. 5: 309 (1818). Krukoffiana L. B. Smith in Contrib. Gray Herb. 154: 36 (1945). Kuntzeana Mez in DC. Monog. 9: 790 (1896). laxissima Mez in Bull. Herb. Boiss. (ser. 2) 5:108 (1905). loliacea Mart. ex Schult. f. Syst. Veg. 7: 1204 (1830). longifolia Baker, Handbk. Bromel. 185 (1889). Lorentziana Griseb. in Goett. Abh. 19: 217 (1874). Mamdonii Morr. ex Mez in DC. Monog. Phan. 9: 871 (1896) = T. crocata. marantoidea Rusby, Descr. S. Amer. Pls. 4(1 920). maxima Lillo & Hauman in Anal. Mus. Nac. Buenos Aires, 29: 416 (1917). maxima var. densior L. B. Smith in Lilloa, 14: 97 (1948). Maxoniana L. B. Smith in Contrib. Gray Herb. 124: 11 (1939). micrantha Baker in Bull. Torr. Bot. Club, 29: 698 (1902) — T. spiculosa. monticola Mez & Sodiro in Bull, Herb. Boiss. (ser. 2) 4: 1185 (1904). myosura Griseb. in Goett. Abh. 24: 333 (187 9). paleacea Presl, Rel. Haenk. 1: 125 (1827). paraénsis Mez in Mart. FI. Bras. 3(3): 586 (1894). pardina L. B. Smith in Lillloa, 14: 98 (1 948). parviflora R. & P. Fl. Peruv. 3: 41, t. 259 (1802). pendulispica Mez in DC. Monog. Phan. 9: 745 (1896). polyphylla Baker, Handbk. Bromel. 178 (1889) = T. vernicosa. polystachya L. Sp. Pl. (ed. 2) 1: 410 (1762). pulchella Hook. Exot. Fl. 2: t. 154 (1825). pulchella var. rosea (Lindl.) Mez. in Mart. Fl. Bras. 3(3): 603 (1894). quadriflora ag Handbk. Bromel. 163 (1889) — T. loliacea. recurvata L. Sp. Pl. (ed. 2) 1: 410 (1762). Reichenbachii Baker, Handbk. seopweee a (1889). rubella Baker in Journ. Bot. 26: 44 Rusbyi Baker, Handbk. Bromel. 176 Vian scalarifolia Baker in Journ. Bot. 25: 235 (1887) = T. paleacea. Seemannii (Baker) Mez in DC. Monog. Phan. 9: 737 (1896). soratensis Baker in Journ. Bot. 25: 235 (1887). sphaerocephala Baker in Journ. Bot. 26: 141 (1888). spiculosa Griseb. in Goett. Nachr. 1864: 17. streptocarpa Baker in Journ. Bot. 25. 241 (1887). triangularis Rusby, Descr. S. Amer. Pls. 3 (1920) = T. pendulispica. tricholepis Baker in Journ. Bot. 25: 234 (1887) . tricholepis var. macrophylla L. B. Smith in Lilloa, 14: 98 (1948). triticea Burchell ex Baker in Journ. Bot. 26: 42 (1888). tucumanensis Mez in DC. Monog. Phan. 9: 853 (1896) = T. Reichenbachii. usneoides L. Sp. Pl. (ed. 2) 1: 411 (1762). Valenzuelana A. Rich. Fl. Cub. Fanerog. 2: 267 (1853). vernicosa Baker in Journ. Bot. 25: 241 (1887). violascens Mez in DC. Monog. Phan. 9: 797 (1896). Walteri Mez in Fedde, Repert. Spec. Nov. 3: 43 (1906). Weddellii Baker, Handbk. Bromel. 181 (1889) = T. decomposita. Williamsii Rusby in Bull. N. Y. Bot. Gard. 6: 489 (1910) = T. capil- lari ; aris. Vriesia heliconioides (Kunth) Lindl. in Bot. Reg. 29: t. 10, in obs. (1843), combination implied, not made. V. heterandra (E. André) L. B. Smith in Contrib. U. S. Nat. Herb. 29: 443 V. icterica Castellanos in Lilloa, 11: 150 (1945). V. rubra (R. & P.) Beer, Bromel. 98 (1857). COMMELINACEAE Aneilema ovato-oblongum Beauv. Fl. Owar. 2: 71, t. 104 (1807). Athyrocarpus rufipes (Seub.) Standl. in Standley & Calderén, Lista Prelim. Pl. El Salvador, 47 (1925). Callisia repens L. Sp. Pl. (ed. 2) 1: 62 (1762). C. repens var. Mandonii C. B. Clarke in DC. Monog. Phan. 3: 311 (1881). Campelia Zanonia (L.) HBK. Nov. Gen. & Spec. 1: 264 (1816). Cochliostema odoratissimum Lem. in Illustr. Hortic. 6: Misc. 70, t. 217 (1859). Commelina Bangii Rusby in Bull. N. Y. Bot. Gard. 6: 490 (1910). elliptica HBK. Nov. Gen. & Spec. 1: 259 (1816). fasciculata R. & P. Fl. Peruv. 1: 44, t. 72 (1798). gracilis R. & P. Fl. Peruv. 1: 44, t. 72, fig. a (1798). hispida R. & P. Fl. Peruv. 1: 43, t. 73 (1798). hispida var. Gaudichaudii C. B. Clarke in DC. Monog. Phan. 3: 156 (1881). nudiflora L. Sp. Pl. 41 (1753). platyphylla var. Balansae C. B. Clarke in DC. Monog. Phan. 3: 177 (1881). quitensis Benth. Pl. Hartw. 258 (1846). quitensis var. Mandonii C. B. Clarke in DC. Monog. Phan. 3: 156 (1881). reflexa Rusby in Phytologia, 1(2): 50 (1934). robusta Kunth, Enum. Pl. 4: 52 (1843). virginica L. Sp. Pl. (ed. 2) 1: 61 (1762). Descantaria Pflanzii Briickn. in Notizbl. 10: 57 (1927). Dichorisandra Aubletiana Schult. f. Syst. Veg. 7: 1181 (1830). Gaudichaudiana Kunth, Enum. Pl. 4: 113 (1843). D. hexandra (Aubl.) Standl. in Standley & Calderén, Lista Prelim. Pl. El Salvador, 48 (1925). D. inaequalis Presl, Rel. Haenk. 1: 140 (1828). D. pubescens Mart. ex Schult. f. Syst. Veg. 7: 1186 (1830). D. villosula Mart. ex Schult. f. Syst. Veg. 7: 1185 (1830). loscopa perforans Rusby in Bull. N. Y. Bot. Gard. 6: 490 (1910). anaanaasaaaaa o 62 | 44 Mandonia boliviana Hassk. in Flora, 54: 260 (1871) = Tradescantia sp. ? Phaeospherion persicariaefolium (Delile) C. B. Clarke in DC. Monog. Phan. 3: 187 (1881). Tinantia fugax Scheidw. in Otto & Dietr. Allg. Gartenzeit. 7: 365 (1839). T. fugax var. erecta (Drummond) C. B. Clarke in DC Monog. Phan. 3: 286 Tradescantia ambigua Mart. ex Schult. f. Syst. Veg. 7: 1170 (1830). cymbispatha C. B. Clarke in DC. Monog. Phan. 3: 296 (1881). elongata G. F. W. Meyer, Prim. FI. a 146 (1818). geniculata Jacq. Enum. Pl. Carib. 18 (1760). . multiflora Sw. Prodr. 57 (1788) : sesame var. parviflora (R. & P.) C. B. Clarke in DC. Monog. Phan. 3: 306 (1881). HHas8H PONTEDERIACEAE Eichhornia azurea (Sw.) Kunth, Enum. Pl. 4: 129 (1848). E. crassipes (Mart.) Solms-Laubach in DC. Monog. Phan. 4: 527 (1883). E. pauciflora Seub. in Mart. Fl. Bras. 3(1): 91 (1847). Heteranthera limosa (Sw.) Vahl, Enum. 2: 44 (1806). H. reniformis R. & P. Fl. Peruv. 1: 43, t. 71 (1798). H. zosteraefolia Mart. Nov. Gen. & Spec. 1: 7, t. 3 (1823). Pontederia rotundifolia L. f. Suppl. 192 (1781). Reussia subovata (Seub.) Solms-Laubach in DC. Monog. Phan. 4: 534 (1883). JUNCACEAE Distichia filamentosa (Buch.) Griseb. in Abh. Nat. Ver. Bremen, 6: 369 (1879). D. muscoides Nees & Meyen in Nov. Act. Acad, Caes. Leop. 19, Suppl. 1: 129 (18438). Juncus andicola Hook. Ic. Pl. 8: t. 714 (1848). J. brunneus Buch. in Abh. Nat. Ver. Bremen, 6: 403 (1879). J. dichotomus Ell. Sketch, 1: 406 (1821). J. imbricatus var. Chamissonis (Kunth) Buch. in Engler, Pflanzenr. IV, 36: 122 (1906). involucratus Steud. ex Buch. in Abh. Nat. Ver. Bremen, 4: *g ecto . Mandonii Buch. in Abh. Nat. Ver. Bremen, 4: 121, t. 3 (187 microcephalus HBK. Nov. Gen. & Spec. 1: 237 (1816). microcephalus var. floribundus (HBK.) Kunth, Enum. Pl. 3: 324 (1841). Luzula meions Buch. in Abh, Nat. Ver. Bremen, 4: 128, t. 4 (1874) = L. peruvian L. excelsa Buch. in Abh. Nat. Ver. Bremen, 4: 126 (1874). L. gigantea Desv. in Journ. Bot. 1: 145 (1808). L. peruviana Desv. in Journ. Bot. 1: 160 (1808). L. racemosa Desv. in Journ. Bot. 1: 162 (1808). Oxychloé andina Phil. Fl. Atac. 52, t. 6 (1860). ahah: LILIACEAE Allium bivalve var. Bangit O. Ktze. Rev. Gen. 3(2): 313 (1898) = Notho- scordum sp. ? Anthericum boliviense v. Poelln. in Rev. Sudam. Bot. 7: 99 (1942). A. eccremorrhizum R. & P. Fl. Peruv. 3: 67, t. 301 (1802). A. glaucum R. & P. Fl. Peruv. 3: 68 (18 02). A. glaucum var. andinum v. Poelln, in Rev. Sudam. Bot. 7: 157 (1943). A. peruvianum Willd. ex Kunth, Enum. Pl. 4: 596 (1843), in synon. Asagraea longiflora Rusby in Bull. N. Y. Bot. Gard. 6: 491 (1910). Asparagus officinalis L. Sp. Pl. 313 apes Asphodelus fistulosus L. Sp. Pl. 309 (1753). Cordyline dracaenoides Kunth in Abh. Acad. Berl. 1842: 30. Dianella boliviana Schlittler in Mitteil. Bot. Mus. Univ. Ziirich, 163: 253 (1940). Excremis coarctata (R. & P.) Baker in Journ. Linn. Soc. 15: 320 (1876). E. scabra O. Ktze. Rev. Gen. 3(2): 316 (1898). Herreria montevidensis Klotzsch ex Griseb. in Mart. Fl. Bras. 3(1): 24 (1847). Milla andicola (Kunth) Baker in Journ. Linn. Soc. 11: 381 (1870 Nothoscordum flavescens Poepp. in Kunth, Enum. Pl. 4: 459 (1843). N. bivalve (L.) Britton in Britton & Brown, Illustr. Fl. 1: 415 (1896). Smilax argyraea L. Lind. & Rod. in Illustr. Hortic. 39: te t. 152 (1892). flavicaulis Rusby in Mem. N. Y. Bot. Gard. 7: 213 (192 Kunthii Killip & Morton in Carnegie Inst. Wash. Publ. pit 269 (1936). irrorata Mart. ex Griseb. in Mart. Fl. Bras. 3(1): 10 (1842). mexicana Griseb. ex Kunth, Enum. Pl. 5: 167 (1850). phylloloba Mart. ex Griseb. in Mart. Fl. Bras. aie 21 (1842). tomentosa HBK. Nov. Gen. & Spec. 1: 272 (1816). HAEMODORACEAE Xiphidium coeruleum Aubl. Pl. Guian. France. 1: 33, t. 11 (1775). AMARYLLIDACEAE Alstroemeria Fiebrigiana Krinzl. in Engler, Bot. Jahrb. 40: 237 (1908). A. pygmaea Herb. Amaryll. 100, t. 8, fig. 4-13 (1837). ? Amaryllis Aglaiae Castellanos in Hesbertia, 7: 68 (1941). Belladonna L. Sp. Pl. 293 (1753). Belladonna var. Haywardii (Traub & Uphof) Traub & Moldenke, Amarylli- dac. 123 (1949). crociflora (Rusby) Traub & Uphof in Herbertia, 5: 128 (1988). Cybister (Herb.) Planch. in Fl. des Serres, 5: t. 455 (1849). elegans Spreng. Pl. Min. Cognit. Pugill. 2: 59 (1815). Mandonii (Baker) Traub & Uphof in Herbertia, 5: 129 (1938). . scopulorum (Baker) Traub & Uphof in Herbertia, 5: 129 (1938). soratensis (Baker) Traub & Uphof in Herbertia, 5: 123 (1938). viridiflora (Rusby) Traub & Uphof in Herbertia, 5: 124 (1938). tamosco microcarpa Rusby in Mem. N. Y. Bot. Gard. 7: 213 (1927). Bomarea acutifolia (Lk. & Otto) Herb. Amaryll. 112 (1837). aurantiaca Herb. Amaryll. 399, t. 46, fig. 1 (1837). boliviensis Baker in Rusby in Bull. Torr. Bot. Club, 29: 700 (1902). brevis (Herb.) Baker in Journ. Bot. 20: 202 (1882). Bridgesiana Beauverd in Bull. Soc. Bot. Genéve (ser. 2), 14: 173 (1922). calyculata Kranzl. in Kew Bull. 1913: 189. coccinea (R. & P.) Baker in Journ. Bot. 20: 202 (1882). crocea (R. & P.) Herb. Amaryll. 119 (1887). distichophylla (Herb.) Baker in Journ. Bot. 20: 202 (1882). dulcis (Hook.) Beauverd in Bull. Soc. Bot. Genéve (ser. 2), 14: 172 (1922). edulis (Tussac) Herb. Amaryll. 111 (1837). Fiebrigiana Krinzl. in Engler, Bot. Jahrb. ~ 230 (1908). fimbriata (R. & P.) Herb. Amaryll. 116 (1837). flava Baker in Rusby in Bull. N. Y. Bot. Gard. 4: 459 (1907). formosissima (R. & P.) Griseb. ex Baker, Handbk. a. 153 (1888). glaucescens (Kunth) Baker in Journ. Bot. 20: 201 (1882). Herbertiana Baker, Handbk. Amaryll. 155 (1888). aes Pe: ~* Spb ias faa na for) involucrosa (Herb.) Baker in Journ. Bot. 20: 201 (1882). multiflora (L. f.) Mirbel in Hist. Nat. Pl. 9: 72 (1804). ovata (Cav.) Mirbel in Hist. Nat. Pl. 9: 72 (1804). petiolata Rusby in Mem. N. Y. Bot. Gard. 7: 216 (1927). petraea Kranzl. in Engler, Bot. Jahrb. 40: 229 (1908). polyphylla Kranzl. in Ann. Naturh. Hofmus. Wien, 27: 158 (1913). puberula (Herb.) Kranzl. in Engler, Bot. Jahrb. 49: 192 (1913). salcilla Mirbel in Hist. Nat. Pl. 9:71 (1804). ee Kranzl. in Engler, Bot. Jahrb. 50, Beibl. 112: 6 (1913). ns Kranzl. in Engler, Bot. Jahrb. 40: 231 (1908). metros Killip in Journ. Wash. Acad. Sci. 25: 373 (1935). superba Herb. Amaryll. 117, t. 6, fig. 1 (1837 tomentosa (R. & P.) Herb. Amaryll. 117 (1837). trachypetala Kranzl. in Engler, Bot. Jahrb. 50, Beibl. 112: 5 (1913). uniflora (Mathews) Killip in Journ. Wash. Acad. Sci. 25: 372 (1935). Chlidanthus fragrans Herb. App. 46 (1821). Collania guadelupensis Kranzl. in Ann. Naturh. Hofmus. Wien, 27: 157 (1913). C. Herzogiana Kranzl. in Engler, Bot. Jahrb. 50, Beibl. 111: 3 (1913). C. subverticillata Kranzl. in Ann. Naturh. Hofmus. Wien, 27: 152 (19138). is fulgens Pax in Engler, Bot. Jahrb. 11: 324 (1889). eka scorzoneraefolia (Lam.) Baker in Journ. Linn. Soc. 17: 124 (1878). Furcraea sp. Habaiithes brachyandrus (Baker) Sealy in Journ. Roy. Hort. Soc. 62: 208 937). gee ieee Ber ee Haylockia andina R. E. Fries in Nov. Act. Soc. Sci. Upsal. (ser. 4) 1(1): 160, t. 9, fig. 1-2 (1905). H. pseudo-colchicum (Kranzl.) Hume in Proc. Florida Acad. Sci. 2: 91 (1938). H. pseudocrocus Solms-Laubach in Bot. Zeit. 65(1): 135 (1907). Hymenocallis narcissiflora (Jacq.) Macbride in Field Mus. Publ. Bot. 11: 11 (1931 H. Guitotnsis Herb. App. 45 (1821). Hypoxis breviscapa HBK. Nov. Gen. & Spec. x 286 (1816). H. humilis HBK. Nov. Gen. & Spec. 1: 286 (1816). Lepidopharynx defliexa Rusby in Mem. N. Y. Bot. Gard. 7: 214, fig. 1 (1927). Pyrolirion boliviense (Baker) Sealy in Journ. Roy. Hort. Soc. 62: 207 (1937). P. xiphopetalum (Baker) Sealy in Journ. Roy. Hort. Soc. 62: 208 (1937). Stenomesson incarnatum (HBK.) Baker in Saunder, Ref. Bot. 5: sub t. 308 (1873). S. Pearcei Baker in Saunder, Ref. Bot. 5: t. 308 (1873). S. recurvatum (R. & P.) Baker in Saunder, Ref. Bot. 5: sub t. 308 (1873). Urceolina peruviana (Presl) Macbride in Field Mus. Publ. Bot. 11: 11 (1931). Wichuraea acicularis (Herb.) M. J. Roem. Syn. Ensat. 280 (1847) — Bomarea dulcis. Zephyranthes tubispatha Herb. App. 36 (1821). Z. viridi-lutea Kranzl. in Fedde, Repert. Spec. Nov. 13: 118 (1914). VELLOZIACEAE Barbacenia boliviensis (Baker) Hauman in Anal. Mus. Nac. Hist. Nat. Buenos Aires, 29: 429 (1917). B. Castillonii Hauman in Anal. Mus. Nac. Hist. Nat. Buenos Aires, 29: 426, t. 4 (1917). DIOSCOREACEAE D. angustifolia Rusby in Bull. Torr. Bot. Club, 29: 701 (1902). Dioscorea acanthogyne Rusby in Bull. N. Y. Bot. Gard. 6: 492 (1910). DUDOSPYUSSEPSOEESOUOEyESyOySoY PoSdoUyOyy Sedo yHydUd 47 arcuata Rusby in Bull. N. Y. Bot. Gard. 4: 460 (1907) = D. Mandonii. Bangii R. Knuth in Engler, Pflanzenr. IV, 43: 324 (1924). 2 bermejensis R. Knuth in Notizbl. 7: 199 (1917). boliviensis R. Knuth in Notizbl. 7: 188 (1917). calderillensis R. Knuth in Notizbl. 7: (1917). chacoénsis R. Knuth in Fedde, Repert. Spec. Nov. 21: 77 (1925). chiquiacensis R. Knuth in Notizbl. 7: 196 (1917). convolvulacea Schlechtd. & Cham. in Linnaea, 6: 49 (1831). coripatensis Macbride in Candollea, 6: 2 (1984). cubijensis R. Knuth in Notizbl. 7: 202 (1917) = Disciphania cubijensis [ Menispermaceae]. cymosula Hemsl. Biol. Centr. Am. Bot. 3: 355, t. 90 (1884). ferruginicaulis Rusby in Mem. N. Y. Bot. Gard. 7: 217 (1927). Friesii R. Knuth in Fedde, Repert. Spec. Nov. 21: 77 (1925). fuliginosa R. Knuth in Fedde, Repert. Spec. Nov. 21: 78 (1925). furcata Griseb. in Mart. Fl. Bras. 3(1): 45 (1842). Galeottiana Kunth, Enum. Pl. 5: 409 (1850). glauca Rusby in Bull. N. Y. Bot. Gard, 4: 459 (1907) = D. coripatensis. glomerulata var. Mandonii (Uline) R. Knuth in Engler, Pflanzenr. IV, 43: 59 (1924). guanaiensis R. Knuth in Engler, Pflanzenr. IV, 43: 78 (1924). hastiformis R. Knuth in Notizbl. 7: 197 (1917). Herzogii R. Knuth in Meded. Rijks Herbar. 29: 56 (1916). irupanensis R. Knuth in Fedde, Repert. Spec. Nov. 30: 160 (19382). Kuntzéi Uline in O. Ktze. Rev. Gen. 3(2): 311 (1898). lagoa-santa Uline ex R. Knuth in Notizbl. 7: 201 (1917) = D. monadelpha. larecajensis Uline ex R. Knuth in Notizbl. 7: 195 (1917). Lorentzii var. Mandonii Uline ex R. Knuth in Notizbl. 7: 187 (1917). macrantha Uline ex R. Knuth in Notizbl. 7: 198 (1917 yi Mandonii Rusby in Bull. Torr. Bot. Club, 29: 701 (1902). monadelpha (Kunth) Griseb .in Vidensk. Medd. Kjoebenh. 1875: 164. multiflora Griseb. in Mart. Fl. Bras. 3(1): 35 (1842). multispicata R. Knuth in Meded. Rijks Herbar. 29: 55 (1916). nodosa R. Knuth in Notizbl. 7: 187 (1917). oblongifolia Rusby in Bull. N. Y. Bot. Gard. 6: 492 (1901). orthoneura var. acutissima Uline ex R. Knuth in Notizbl. 7: 216 (1917). ovalifolia R. Knuth in Notizbl. 7: 200 (1917). polygonoides Humb. & Bonpl. ex Willd. Sp. Pl. 4(2): 795 (1805). racemosa Rusby in Bull. N. Y. Bot. Gard. 4: 459 (1907) = D. Bangii. Rusbyi Uline in Engl. & Prantl, Nat. Pflanzenfam. Nachtr. I1(5): 86 (1897). sinuata Vell. Fl. Flum. 10: t. 129 (1827). spectabilis R. Knuth in Meded. Rijks Herbar. 29: 55 (1916). tarijensis R. Knuth in Notizbl. 7: 212 (1917). toldosensis R. Knuth in Engler, Pflanzenr. IV, 43: 351 (1924). Trollii R. Knuth in Fedde, Repert. Spec. Nov. 30: 161 (1932). violacea R. Knuth in Notizbl. 7: 200 (1917) = D. toldosensis. IRIDACEAE Alophia tigridioides Hicken in Darwiniana, 1: 116 (1926). Calydorea approximata R. C. Foster in Contrib. Gray Herb. 155: 46 (1945). C. azurea Klatt in Abh. Naturf. Ges. Halle, 15: 387 (1882). C. campestris (Klatt) Baker in Journ, Bot. 14: 187 (1876). Cardenanthus boliviensis R. C. Foster in Contrib. Gray Herb. 155: 4 (1945). C. longitubus R. C. Foster in Contrib. Gray Herb. 171: 23 (1950). C. orurensis R. C. Foster in Contrib. Gray Herb. 161: 14 (1946). 48 C. tunariensis R. C. Foster in Contrib. Gray Herb. 155: 5 (1945). Cipura major Rusby in Bull. N .Y. Bot. Gard. 6: 493 (1910). C. paludosa Aubl. Pl. Guian. Fr. 1: 38, t. 18 (1775). Cypella linearis (HBK.) Baker, Handbk. Irid. 65 (1892). C. peruviana Baker in Bot. Mag. 102: t. 6213 (1876). Eleutherine bulbosa (Mill.) Urb. in Fedde, Repert. Spec. Nov. 15: 305 (1918). Libertia boliviana R. C. Foster in Contrib. Gray Herb. 161: 4 (1946). Mastigostyle brevicaulis (Baker) R. C, Foster in Contrib. Gray Herb. 161: 16 946). M. Cardenasii R. C. Foster in Contrib. Gray Herb. 155: 23.(1945). Orthrosanthus nigrorhynchus Rusby in Mem. Torr. Bot. Club, 6: 126 (1896). O ; Rev. Gen. 3(2): 309 (1898) = O. nigrorhynchus. Phaiophleps acaulis (Klatt) R. C. Foster in Contrib. Gray Herb. 127: 43 (1939). Sphenostigma boliviense Baker, Handbk. Irid. 107 (1892). S. Goodspeedianum R. C. Foster in Contrib. Gray Herb. 171: 27 (1950). S. Mandonii (Rusby) R. C. Foster in Contrib. Gray Herb. 161: 10 (1946). S. Mandonii var. bulbilliferum R. C. Foster in Contrib. Gray Herb. 161: 11 (1946). S. umbellatum (Klatt) Klatt in Abh. Naturf. Ges. Halle, 15: 363 (1882). Sisyrinchium alatum var. minus Rusby in Bull, N. Y. Bot. Gard. 6: 493 (1910). S. azureum Phil. Fl. Atac. 50 (1860). S. Bakeri Klatt in Abh. Naturf. Ges. Halle, 15: 378 (1882) — S. trinerve. S. brevipes Baker, Handbk. Irid. 130 (1882). S. eryptocarpum Rusby in Mem. Torr. Bot. Club, 6: 126 (1896) — S. trinerve.. S. hypsophilum I. M. Johnst. ex R. C. Foster in Contrib. Gray Herb. 166: 30 (1948). Ivanii R. C. Foster in Contrib. Gray Herb. 166: 30 (1948). junceum E. Mey. ex Presl, Rel. Haenk. 1: 118 (1827). laterale Baker in Journ. Bot. 14: 269 (1876). macrocephalum R. Graham in Edinb. New Philos. Journ. 176 (Jan. 1833). Mandonii Baker in Journ. Bot. 14: 269 (1876). . Marchio (Vell.) Steud. Nomencl. (ed. 2) 2: 596 (1841). micranthum Cav. Diss. 2: 345, t. 191, fig. 2 (1788). . pictum Kranzl. in Fedde, Repert. Spec. Nov. 13: 119 (1914) = S. lat- erale ? rigidifolium Baker, Handbk. Irid. 131 (1892). . tinctorium HBK. Nov. Gen. & Spec. 1: 824 (1816). trinerve Baker in Journ. Bot. 14: 267 (1876). eae —o in Linnaea, 34: 737 (1866). . Syst. Veg. 1: 166 (1825). Tord bearteliale (Klatt) Macbride in Candollea, 5: 348 (1934). RANA aiasasas MUSACEAE Heliconia acuminata A. Rich. in Nov. Act. Acad. Caes. Leop. 15, Suppl.: 26, t. 11 (1831) H. Bihai sensu auctt. non L. H. cannoides sensu auctt. non A. Rich. H. Cardenasii L. B. Smith in Contrib. Gray naa 124: 5 (1939). H. episcopalis Vell. Fl. Flum. 3: 107, t. 22 (1825). H. Pearcei Rusby in Bull. N. Y. Bot. Gard. 6: 494 (1910) = H. acuminata. H. psittacorum L. f. Suppl. 158 (1781). H. robusta Pax in Fedde, Repert. Spec. Nov. 7: 107 (1909). H. rostrata R. & P. Fl. Peruv. 3: 71, t. 305 (1802). 49 ZINGIBERACEAE Costus cylindricus Jacq. Fragm. 34, t. 77 (18 cylindricus var. pulcherrimus (O. Ktze.) c Sehain. in Engler, Pflanzenr. IV, 46: 406 (1904). guanaiensis Rusby in Bull. Torr. Bot. ry Fa 694 (1902). longifolius Rusby in Phytologia, 1(2): 1934). Mooreanus Rusby in Bull. N. Y. Bot. ek 4: 454 (1907). rurrenabaqueanus Rusby in Mem. N. Y. Bot. Gard. 7: ae (1927). sinningiaeflorus Rusby in Mem. N. Y. Bot. Gard. 7: 219 (1927). spicatus (Jacq.) Sw. Prodr. 11 (1788). Steinbachii Loes. in Notizbl. 10: 714 (1929). . Tatei Rusby in Phytologia, 1(2): 51 (1934). Dimerocostus bolivianus (Rusby) Loes. in Notizbl. 10: 716 (1929). D. Gutierrezii O. Ktze. Rev. Gen. 3(2): 301 (1898). Renealmia Cardenasii Rusby i in Mem. N. Y. Bot. Club, 7: 219 (1927). R. dermatopetala K. Schum. in Engler, Pflanzenr. IV, 46: 304 (1904). icrantha K. Schum. in Engler, Pflanzenr. IV, 46: 298 (1904). R. Ruiziana (Steud.) Horan. Prodr. Monog. Scitam. 33 (1862). R. spectabilis Rusby in Mem. N. Y. Bot. Gard. 7: 218 (1927). saqQaaaaaa CANNACEAE Canna Bangii Kranzl. in Engler, Pflanzenr. IV, 47: 38 (1912). Brittonii Rusby in Bull. Torr. Bot. Club, 29: 695 (1902). coccinea Mill. Dict. (ed. 8) no. 3 (1768). edulis Ker in Bot. Reg. 9: t. 775 (1824). glauca L. Sp. Pl. 1 (1753). lanuginosa Rose. Scit. t. 16 (1828). pedunculatus Sims in Bot. Mag. 49: t. 2323 (1822). agaaaa MARANTACEAE Calathea bracteosa Rusby in Mem. N. Y. Bot, Gard. 7: 220 (1927). Buchtienii Pax in Fedde, Repert. Spec. Nov. 7: 107 (1909). capitata (R. & P.) Lindl. in Bot. Reg. 14: sub t. 1210 (1829). Cardenasii Rusby in Mem. N. Y. Bot. Gard. 7: 222 (1927). divaricata Rusby in Bull. Torr. Bott. Club, 29: 695 (1902). exserta Rusby in Bull. N. Y. Bot. Gard. 6: 495 (1910). gigas Gagnep. in Bull. Soc. Bot. France, 50: 589 (1904). Mansonis Koern. in Bull. Soc. Nat. Mosc. 35(1): 119 (1862). nigricans Rusby in Bull. N. Y. Bot. Gard. 6: 496 (1910) —C. Rusbyi. nodosa Rusby in Bull. N. Y. Bot. Gard. 4: 455 (1907). Pearcei Rusby in Mem. Torr. Bot. Club, 6: 123 (1896). pilosa Rusby in Bull. N. Y. Bot. Gard. 6: 496 (1910). Rusbyi Loes, in Engl. & Prantl, Nat. Pflanzenfam. (ed. 2) 15a: 678 (1930). Sprucei Rusby in Bull. N. Y. Bot. Gard. 6: 495 (1910). stenostachys Rusby in Mem. N. Y. Bot. Gard. 7: 221 (1 stromanthifolia Rusby in Bull. N. Y. Bot. Gard. 4: 456 (1907). Whitei Rusby in Mem, N. Y. Bot. Gard. 7: 221 (1927). Ethanium jamaicense (Gaertn.) O. Ktze. Rev. Gen. 2: 689 (1891). Ischnosiphon Baenitzii Pax in Fedde, Repert. Spec. Nov. 7: 108 (1909). I. bolivianus Loes. in Notizbl. 6: 280 (1915). I. gracilis (Rudge) Koern. in Bull. Soc. Nat. Mose. 35(1): 94 (1862). I. lasiocoleus var. bolivianoides Loes. in Notizbl. 6: 280 (1915). I. Parkeri (Rosc.) Koern. in Bull. Soc. Nat. Mosc. 35(1): 81 (1862). MQAARAAAHRHARARAARAAARA 50 Maranta amplifolia K. Schum. in Engler, Pfianzenr. IV, 48: 128 (1902). M. Tonckat Aubl. Pl. Guian. Fr. 1: 3 (1775) Myrosma boliviana Loes. in Notizbl. 6: 270 (1915). M. boliviana var. acreana Loes. in Notizbl. 6: 270 (1915). Stromanthe angustifolia Rusby in Mem. Torr. Bot. Club, sek 266 (1895). S. boliviana K. Schum. in Engler, Pflanzenr. IV, 48: 151 (1902). S. confusa K. Schum. in Engler, Pflanzenr. IV, 48: 150 (1902). S. Porteana Griseb. in Ann. Sci. Nat. (ser. 4) 9: 185 (1858). BURMANNIACEAE Apteria aphylla (Nutt.) Barnh. ex Small, Fl. SE. U. S. 309 (1903). A. boliviana Rusby in Bull. N. Y. Bot. Gard. 4: 447 (1907) = A. aphylla. A. gentianoides Jonker, Monog. Burmann, 211 (1938). Burmannia capitata (Walt.) Mart. Nov. Gen. 1: 12 (1823). B. flava Mart. Nov. Gen. 1: 11, t. 5 (1823). B. tenella Benth. in Journ. Bot. 7: 12 (1855). Dictyostega orobanchioides Miers in Proc. Linn. Soc. 1: 61 (1840). ORCHIDACEAE Aa microtidis Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 35 (1922) Altensteinia sp. ? A. sphaeroglossa Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 36 (1922) Altensteinia sp. ? A. trilobulata Schltr. Pa Fedde, Repert. Spec. Nov. Beih. 10: 37 (1922) Altensteinia sp. Altensteinia nea Reichb. f. Xen. Orch. 3: 19 (18 | 78). A. boliviensis Rolfe in Mem. Torr. Bot. Club, 4(3): 265 (1895) = A. fim- briata. ; — (Schitr.) C. Schweinf. in Bot. Mus. Leafl. Harvard Univ. 16(1): 1953 A A. LB oa Schltr. in Fedde, Repert. Spec. Nov. 10: 445 (1912). A. fimbriata HBK. Nov. Gen. & Spec. 1: 333, t. 72 (1816). gymnandra Reichb. f. Xen. Orch. 3: 18 (1878). inaequalis Reichb. f. Xen. Orch. 3: 19 (1878). Mandonii Reichb. f. Xen. Orch. 3: 19 (1878). . marginata Reichb. f. Xen. Orch. 3: 19 (1878). . Matthewsii Reichb. f. Xen. Orch. 3: 19 (1878). paludosa Reichb. f. Xen. Orch. 3: 19 (1878). Weddelliana Reichb. f. Xen. Orch. 3: 19 (1878). PP bbb bb Amblostoma cernuum Scheidw. in Otto & Dietr. Gartenzeit. 6: 383 (1838). A. densum Reichb. f. Xen. Orch. 3: 22 (1878). Beloglottis boliviensis Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: (1922) = (probably) Spiranthes costaricensis. Bletia catenulata R. & P. Syst. 229 (1798). B. Mandonii Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 48 (1922). B. Wageneri Reichb. f. in Bonplandia, 2: 22 (1854). Brachionidium concolor Lindl. Fol. Orch. Brach. 1 (1859). Brassia boliviensis Schltr. in Fedde, Repert. Spec. Nov. 12: 494 (1913). B. caudata (L.) Lindl. in Bot. Reg. 10: t. 832 (1824). ' B. Lanceana Lindl. in Bot. Reg. 21: t. 1754 (1835). B. thyrsodes Reichb. f. in Gard. Chron. 1868: 342. Buchtienia boliviensis Schltr. in Fedde, Repert. Spec. Nov. 27: 34 (1929). Bulbophyllum amazonicum L. O. Wms. in Lilloa, 5: 7 (1940). B. bolivianum Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 49 (1922). 38 51 B. tricolor Smith & Harris in Contrib. Gray Herb. 114: 11 (1936). B. Weberbauerianum var. angustius C. Schweinf. in Bot. Mus. Leafl. Harvard Univ. 11: 189 (1944). Camaridium Meg Rolfe in Mem, Torr. Bot. Club, 4(3): 264 (1895) = Maxillaria sp C. ecbae aekiix! in : Weadé: Repert. Spec. Nov. 27: 76 (1929) = Maxillaria sp. C. vagans Schltr. in Fedde, Repert. Spec. Nov. 27: 77 (1929) = Maxillaria ey Ganpeidecacs apiculatum Schltr. in Fedde, Repert. Spec. Nov. 27: 84 (1929). C. Kuntzei Cogn. in O. Ktze. Rev. Gen. 3(2): 298 (1898). C. neglectum (Reichb. f. & Warm.) Cogn. ex Chod. in Bull. Herb. Boiss. (ser. 2) 1: 425 (1901). Catasetum Buchtienii Kranzl. in Fedde, Repert. aa Nov. 25: 27 (1928). C. cristatum Lindl. in Bot. Reg. 10: sub t. 840 (18 C. fimbriatum (C. Morr.) Lindl. in Paxt. Flow. se § 1: 124 (1850). C. fimbriatum var. me Sg a (Morr.) Mansf. in Fedde, Repert. Spec. Nov. 31: 108 (1932). C. Pflanzii Schltr. in dacs: Repert. Spec. Nov. 11: 45 (1912) = C. fimbria- . Morrenianu C. Wredeanum Schltr. in “Geet 9:14 (1915), nomen nudum = C. fimbria- ar. renianum. Cattleya luteola Lindl. in Gard. Chron. 1853: 774. C. Walkeriana Gardn. in Hook. Lond. Journ. Bot. 2: 662 (1843). Centrogenium roseoalbum (Reichb. f.) Schltr. in Beih. Bot. Centralbl. 37(2): 453 (1920). Chloraea boliviana (Reichb. f.) Kranzl. Orch. Gen. & Spec. 2: 139, t. 16 (1904). C. calantha Kranzl. in Engler, Bot. Jahrb. 37: 395 (1906). C. Fiebrigiana Krianzl. in Engler, Bot. Jahrb. 37: 396 (1906). C. ignea Krianzl. in Engler, Bot. Jahrb. 37: 396 (1906). C. reticulata Schltr. in Fedde, Repert. Spec. Nov. 15: 210 (1918). Cochlioda Noezliana Rolfe in Lindenia, 6:55, t. 266 (1890). Comparettia falcata Poepp. & Endl. Nov. Gen. 1: 42, t. 73 (1855). C. macroplectron Reichb. f. & Triana in Gard. Chron. 1878 (II): 524. C. splendens Schltr. in Fedde, Repert, Spec. Nov. Beih. 10: 51 (1922). Corymborchis flava (Sw.) O. Ktze. Rev. Gen. 2: 658 (18 etl Cranichis ciliata (HBK.) Kunth, Syn. Pl. Aeq. 1: 324 (1822 C. Mandonii Schltr. in Fedde, Repert. Spec. Nov. Beih. i 38 (1922) =— C. ciliata. Cyclopogon casanaénsis Schltr. in Fedde, Repert. Spec. Nov. 27: 32 (1929) = Spiranthes sp. ? Cycnoches Buchtienii Kranzl. in Fedde, Repert. Spec. Nov. 25: 26 (1928). ? C. Haagei Rodr. Orch. Nov. 2: 221 (1882). Cryrtopodium Buchtienii Schltr. in Fedde, Repert. Spec. Nov. 25: 26 (1928). C. Pflanzii Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 49 (1922). C. punctatum (L.) Lindl. Gen. & Spec. Orch. 188 (1833 Diadenium micranthum Poepp. & Endl. Nov. Gen. 1: 41, t. 71 (1835). Dichaea anguina Schltr. in Fedde, Repert. Spec. Nov. 27: 82 (1929): D. Buchtienii Schitr. in Fedde, Repert. Spec. Nov. Beih. 10: 53 (1922). D. echinocarpa (Sw.) Lindl. Gen. & Spec. Orch. 208 (1833). D. hamata Rolfe in Mem. Torr. Bot. Club, 4(3): 264 (1895). D. laxa (R. & P.) Poepp. & Endl. Nov. Gen. 2: 4, t. 105 (1838). D. longa Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 54 (1922). ? D. muricata (Sw.) Lindl. Gen. & Spec. Orch. 209 (1833). D. robusta Schltr. in Fedde, Repert. Spec. Nov. 27: 83 (1929). 52 D. stenophylla Schltr. in Fedde, Repert. Spec. Nov. 27: 84 (1929). ? Elleanthus brasiliensis Reichb. f. in Walp. Ann. 6: 475 (1861). . linifolius Presl, Rel. Haenk. 1: 97 (1827). maculatus (Lindl.) Reichb. f. in Walp. Ann. 6: 482 (1861). scopula Schltr. in Fedde, Repert. Spec. Nov. 10: 457 (1912). setosus Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 34 (1922). yungasensis Rolfe in Mem. Tomm, Bot. Club. 4(3): 262 (1895). Encyclia Buchtienii Schltr. in Fedde, Repert. Spec. Nov. 27: 62 (1929) idendrum sp., not Epidendrum Buchtienii. E. Pflanzii Schitr. in Fedde, Repert. Spec. Nov. Beih. 10: 46 (1922) = Epidendrum sp. ? E. Steinbachii Schitr. in Fedde, Repert. Spec. Nov. Beih. 10: 47 (1922) = Epidendrum sp. ? Epidendrum albifiorum Schitr. in Fedde, Repert. Spec. Nov. 12: 488 (1913). alopecurum Schltr. in Fedde, Repert. Spec. Nov. 27: 58 (1929). armeniacum Lindl. in Bot. Reg. 22: t. 1867 (1836). Bangii Rolfe in Bull. N. Y. Bot Gard. 4: 451 (1907) = E. rostrigerum. bolivianum Schltr. in Fedde, Repert. Spec. Nov. 10: 456 (1912). brachycladium Lindl. Fol. Orch. Epid. 60 (1853) . brachyglossum Lindl. Fol. Orch. Epid. 72 (1853). Buchtienii Schltr. in Fedde, Repert. Spec. Nov. 10: 455 (1912). buenavistae Kranzl. in Fedde, Repert. Spec. Nov. 6: 19 (1908). cartilaginiflorum Reichb. f. Xen. Orch. 3: 24 (1878). Christyanum Reichb. f. in Gard. Chron. 22: 38 (1884). coroicoénse Schltr. in Fedde, Repert. Spec. Nov. 27: 60 (1929). corymbosum Lindl. Fol. Orch. Epid. 61 (1853). crassinervium Kranzl. in Fedde, Repert. Spec. Nov. 1: 182 (1905). cristatum R. & P. Syst. 248 (1798). cuneatum Schltr. in Fedde, Repert. Spec. Nov. 10: 456 (1912). difforme Jacq. Enum. Pl. Carib. 29 (1760). E. elongatum Jacq. Ic. Pl. Rar. 3:17, t. 604 (1793). Evelynae Reichb. f. Xen. Orch. 3: 23 (1878). exaltatum Krinzl. in Engler, Bot. Jahrb. 54, Beibl. 117: 26 (1916). E. fallax Lindl. Orch. Linden. 9 (1846). fimbriatum var. rhomboglossum (Krinzl.) C. Schweinf. in Bot. Mus. Leafl. Harvard Univ. 11: 224 (1844). fragrans Sw. Prodr. 123 (1788). Friderici-Guglielmi Warsc. ex Reichb. f. in Bonplandia, 2: 110 (1854). gladiatum Lindl. Gen. & Spec. Orch. 106 (1831). Guentherianum Kranzl. in Fedde, Repert. Spec. Nov. 25: 20 (1928). Harrisoniae Hook. in Bot. Mag. 60: t. 3209 (1833). Herzogii Schltr. in Fedde, Repert. Spec. Nov. 12: 489 (1913). humidicola Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 44 (1922) = E. paniculatum. ibaguense HBK. Nov. Gen. & Spec. 1: 352 (1816). insectiferum Lindl. Fol. Orch. Epid. 87 (1853). integrilabium Ames & Schweinf. Sched. Orch. 8: 46 (1925) = E. fimbriatum var. rhomboglossum. lanioides Schltr. in Fedde, Repert. Spec. Nov. 12: 389 (1913). lanipes Lindl. Fol. Orch. Epid. 91 (1853). mapiriense Kranzl. in Fedde, Repert. Spec. Nov. 25: 21 (1928). mesomicron Lindl. Fol. Orch. Epid. 51 (1853). Miguelii Schltr. in Fedde, Repert. Spec. Nov. 21: 337 (1925). nigricans Schitr. in Fedde, Repert. Spec. Nov. 12: 490 (1913). nocturnum Jacq. Enum. Pl. Carib. 29 (1760). obliquum Schltr. in Fedde, Repert. Spec. Nov. 10: 457 (1912). bs be el td bt | BY ed bed ed eed ty th yt td et bd tbat ete to Bad ed at db at td ty odontospathum Reichb. f. Xen. Orch. 3: 23 (1878). oreonastes Reichb. f. Xen. Orch. 3: 22 (1878). paniculatum R. & P. Syst. 243 (1789). Peperomia Reichb. f. in Bonplandia, 2: 20 (1854). physophorum Schltr. in Fedde, Repert, sath Nov. 12: 490 (1918). purum Lindl. in Bot. Reg. 30: Misc. 75 (1844). pygmaeum Hook. in Bot. Mag. 60: t. 3233 (1833). quinquepartitum Schltr. in Fedde, Repert. Spec. Nov. Beih. 10:45 (1922). ramosum Jacq. Select. Stirp. Amer. 221 (1763). rigidum Jacq. Enum. Pl. Carib. 29 (1760). rostrigerum Reichb. f. in Linnaea, 41: 38 (1877). scopulorum Reichb. f. Xen. Orch. 3: 23 (1873). soratae Reichb. f. Xen. Orch. 3: 24 (1878). Steinbachii Ames, Sched. Orch. 1: 20 (1922) = E. obliquum. syringodes Schltr. in Fedde, Repert. Spec. Nov. 27: 61 (1929). syringothyrsus Reichb. f. Xen. Orch. 3: 22 (1878). . Theodori Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 46 (1922). trichopetalum Schltr. in Fedde, Repert. Spec. Nov. 12: 491 (1931). trichorhizum Kranz]. in Fedde, Repert. Spec. Nov. 25: 21 (1928). yungasense Rolfe in Mem. Torr. Bot. Club, 4(3): 262 (1895). Epistephium amplexicaule ((R. & P.) Poepp. & Endl. Nov. Gen. 1: 52, t. 91 E. Herzogianum Kranzl. in Fedde, Repert. Spec. Nov. 6: 21 (1908). E. sclerophyllum Lindl. Gen. & Spec. Orch. 433 (1840). Eulophidium maculatum (Lindl.) Pfitz. in Engl. & Prantl, Nat. Pflanzenfam. 2(6): 188 (1889). Galeandra Fiebrigii Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 47 (1922). ? G. lagoénsis Reichb. f. & Warsc. in Otia Bot. Hamb. fasc. 1: 88 (1881). Gomphichis longifolia (Rolfe) Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 60 (1922). G. plantaginifolia C. Schweinf. in Bot. Mus. Leafl. Harvard Univ. 11: 217 (1944). G. valida Reichb. f. Xen. Orch. 3: 20 (1878). Govenia boliviensis Rolfe in Mem. Torr. Bot. Club, ia 263 (1895). G. tingens Poepp. & Endl. Nov. Gen.2: 5, t. 107 (1838) — G. utriculata. G. utriculata (Sw.) Lindl. in Bot. Reg. 25: Misc. 47 (1839) Habenaria Bangii Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 30 (1922). H. bermejoénsis Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 30 (1922). boliviana Reichb. f. in Bonplandia, 3: 213 (1885). bractescens Lindl. Gen. & Spec. Orch. 308 (1835). Buchtienii Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 31 (1922). flexa Reichb. f. ex Kranzl. in Engler, Bot. Jahrb. 16: 97 (1892). Guentheriana Krinzl. in Fedde, Repert. Spec. Nov. 25: 18 (1928). Herzogii Schltr. in Fedde, Repert. Spec. Nov. 12: 48 (1913). leptantha Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 31 (1922). leucosantha Rodr. Orch. Nov. 1: 151 (1877). ligulata C. Schweinf. in Bot. Mus. Leafl. Harvard Univ. 9: 47 (1941). maxillaris Lindl. in Hook. Journ. Bot. 1: 5 (1834). microstylina Reichb. f. Xen. Orch. 3: 17 (1878). Miguelii Schltr. in Fedde, Repert. Spec. Beih. 10: 32 (1922). monorrhiza (Sw.) Reichb. f. in Ber. Deut. Bot. Ges. 3: 274 (1885). Ottonis Schltr. in Fedde, Repert. Spec. Nov. 27: 28 (1929) — H. Guenth- eriana. Paiveana Reichb. f. Xen. Orch. 3: 17 (1878). petrogeiton Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 33 (1922). pseudorepens Schltr. in Fedde, Repert. Spec. Nov. 12: 481 (1913). SO RRR ERT eee pratensis (Lindl.) Reichb. f. in Linnaea, 22: 813 (1849). pungens Cogn. in O. Ktze. Rev. Gen. 3(2): 299 (1898). repens Nutt. Gen. Am. Pl. 2: 190 (1818). sartor Lindl. in Hook. Journ. Bot. 2: 662 (1843). simillima Reichb. f. Xen. Orch. 3: 18 (1878). speciosa Poepp. & Endl. Nov. Gen. 1: 44 (1835). subandina Schltr. in Fedde, Repert. Spec. Nov. 27: 29 (1929). Theodori Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 33 (1922). Williamsii Schltr. in Fedde, Repert. Spec. Nov. 10: 445 (1912). yungasensis Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 34 (1922). Hexadesmia boliviensis Rolfe in Mem. Torr. Bot. Club, 6: 122 (1896) = Sca- phyglottis Summersii, at least in part. Houlletia boliviana Schltr. in Fedde, Repert. Spec. Nov. 27: 65 (1929). H. Buchtienii Kranzl. in Fedde, Repert. Spec. Nov. 25: 28 (1928) H. odoratissima Lindl. in Paxt. Flow. Gard. 3: 172 (1853). H. Sanderi Rolfe in Gard. Chron. (ser. 3) 47: 206 (1910). Ionopsis utricularioides (Sw.) Lindl. Coll. Bot. t. 39A (1821-24). Isochilus linearis (Jacq.) R. Br. in Ait. Hort. Kew. (ed. 2) 5:209 (1813). Kefersteinia pulchella Schltr. in Fedde, Repert. Spec. Nov. 27: 68 (1929). Koellensteinia boliviensis (Rolfe) Schltr. in Orchis, 12: 32 (1918). Lacaena grandis Kranzl. in Fedde, Repert. Spec. Nov. 25: 25 ae Laelia undulata (Lindl.) L. O. Wms. in Darwiniana, 5: 76 (1941). Lepanthes Koehleri Schltr. in Fedde, Repert. Spec. Nov. 10: 386 (1912). L. Paiveana Reichb. f. Xen. Orch. 3: 26 (1878). L. rupicola Schltr. in Fedde, Repert. Spec. Nov. 12: 485 (1913). L. sillarensis Schltr. in Fedde, Repert. Spec. Nov. 12: 486 (1913). Liparis elata Lindl. in Bot. Reg. 14: t. 1175 (1828). L. neuroglossa Reichb. f. Xen. Orch. 3: 26 (1878). L. otophyllon Schitr. in Fedde, Repert. Spec. Nov. Beih. 10: 41 (1922). L. ramosa Poepp. & Endl. Nov. Gen. 2: 9, t. 112 (1837). L. retusa Fawe. & Rendl. in Journ. Bot. 47: 128 (1909). L. Rusbyi Rolfe in Bull. N. Y. Bot. Gard. 4: 454 (1907). L. vexillifera (Llave & Lex.) Cogn. in Mart. Fl. Bras. 3(4): 289 (1895). ? Lockhartia lunifera (Lindl.) Reichb. f. in Bot. Zeit. 10: 767 (1852). L. parthenocomos (Reichb. f.) Reichb. f. in Bot, Zeit. 10: 767 (1852). Lycaste costata (Lindl.) Lindl. in Bot. Reg. 29: Misc. 15 (1848). L. macrophylla (Poepp. & Endl.) Lindl. in Bot. Reg. 28: Misc. 85 (1842). L. neglecta Schltr. in Fedde, Repert. Spec. Nov. 27: 66 (1929). Macradenia Buchtienii Schltr. in Fedde, Repert. Spec. Nov. 27: 79 (1929). Malaxis boliviana (Schltr.) Ames, Sched. Orch. 9: 18 (1925). M. excavata (Lindl.) O. Ktze. Rev. Gen. 2: 673 (1891). fastigiata (Reichb. f.) O. Ktze. Rev. Gen. 2: 673 (1891). hastilabia (Reichb. f.) O. Ktze. Rev. Gen. 2: 673 (1891). Hieronymi (Cogn.) L. O. Wms. in Lilloa, 4: 363 (1939). major (Reichb. f.) Leon ex A. D. Hawkes in Phytologia, 3: 260 (1950). Reichenbachiana (Schlitr.) L. O. Wms. in Bot. Mus. Leafl. Harvard Univ. 5: 148 (1938). Masdevallia aspera Reichb. f. ex Kranzl. in Fedde, Repert. Spec. Nov. Beih. 34: 82 (1925). auropurpurea Reichb. f. & Warsc. in Bonplandia, 2: 115 (1854). Bangii Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 41 (1922). boliviensis Schltr. in Fedde, Repert. Spec. Nov. 12: 483 (1913) = d ol Jaahapanabala ¢ SR RSS scandens. brachyantha Schltr. in Fedde, Repert. Spec. Nov. 27: 87 (1929). . Buchtienii Schitr. in Fedde, Repert. Spec. Nov. 10: 450 (1912) = M. scan- dens. RR EEE 55 ees Sabie in Fedde, Repert. Spec. Nov. Beih, 10: 42 (1922) = M. auropurpurea, ionocharis Reichb. f. in Gard. Chron. — (II): 388. . Paiveana Reichb. f. Xen. Orch. 3: 25 (1878). scandens Rolfe in Bull. N. Y. Bot. Gard. 4: 453 (1907). setipes Schltr. in Fedde, Repert. Spec. Nov. 27: 38 (1929). tubata Schltr. in Fedde, Repert. Spec. Nov. 27: 39 (1929). xanthura Schitr. in Fedde, Repert. Spec. Nov. 27: 39 (1929) = M. auro- = Apibslal = purpurea, axillaria aurea var. “eeere? (Lindl.) C. Schweinf. in Bot. Mus. Leafil. Har- vard Univ. 11: 263 (1945). boliviensis Schltr. in Fedde, Repert. Spec. Nov. Beith. 10: 51 (1922). M. breviscapa Poepp. & Endl. Nov. Gen. 1: 36 (1835). . Buchtienii Schltr. in Fedde, Repert. Spec. Nov. 27: 68 (1929). casta Kranzl. in Fedde, Repert. Spec. Nov. 25: 30 (1928). compressibulba Schltr. in Fedde, Repert. Spec. Nov. 27: 69 (1929). . densifolia (Poepp. & Endl.) Reichb. f. in Walp. Ann. 6: 534 (1861). . divaricata (Barb. Rodr.) Cogn. in Mart. Fl. Bras. 3(6): 49, t. 4, fig. 1 (1904). . dolichophylla Schltr. in Fedde, Repert. Spec. Nov. 10: 458 (1912). . fallax Schltr. in Fedde, Repert. Spec. Nov. 27: 70 (1929) = M. divaricata. . gracilipes Schltr. in Fedde, Repert. Spec. Nov. 27: 71 (1929). . grandiflora (HBK.) Lindl. Gen. & Spec. Orch. 147 (1832). . Guentheriana Kranzl. in Fedde, Repert. Spec. Nov. 25: 31 (1928). . Herzogiana Kranzl. in Fedde, Repert. Spec. Nov. 6: 20 (1908). . leucantha Schltr. in Fedde, Repert. Spec. Nov. 27: 72 (1929). . longibracteata (Lindl.) Reichb. f. in Walp. Ann. 6: 540 (1861). . longicaulis Schltr. in Fedde, Repert. Spec. Nov. 27: 72 (1929). mapiriensis (Kranzl.) L. O. Wms. in Caldasia, 5: 16 (1942). nervosa Rolfe in Mem. Torr. Bot. Club, 4 (3): 263 (1895). oxysepala Schltr. in Fedde, Repert. Spec. Nov. 27: 73 (1929). . poaefolia Schltr. in Fedde, Repert. Spec. Nov. 27: 74 (1929). . polybulbon Krinzl. in Fedde, Repert. Spec. Nov. 6: 19 (1908). simacoana Schltr. in Fedde, Repert. Spec. Nov. 27: 75 (1929). splendens Poepp. & Endl. Nov.. Gen. 1: 38, t. 66 (1835). xylobiiflora Schltr. in Fedde, Repert. Spec. Nov. 27: 76 (1929). Microstylis sures Schltr. in Fedde, Repert. Spec. Nov. 10: 449 (1912) = Malaxis M. Mandonii "Reich. f. in Flora, 71: 152 (1888) = Malaxis sp. ? M. mixta Schltr. in Fedde, Repert. Spec. Nov. 10: 449 (1912) = Malaxis sp. ? M. nasuta Schltr. in Fedde, Repert. Spec. Nov. 27: 35 (1929) — Malaxis sp. ? M. Ottonis Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 39 (1922) = Malaxis hastilabia. M. tridentula Schltr. in Fedde, Repert. Spec. Nov. 27: 36 (1929) = Malaxis sp. ? M. Weddellii Finet in Bull. Soc. Bot. France, 54: 532 (1907) = Malaxis sp. ? Mormodes Guentherianum (Krinzl.) Mansf. in Fedde, Repert. Spec. Nov. 31: 112 (1932). Neodryas Herzogii Schltr. in Fedde, Repert. Spec. Nov. 12: 493 (1913). latilabia hus & Harris in Contrib. Gray Herb. 117: 39 (19837) = N rhodoneu N. Mandonii Reich. f. Xen. Orch. 3: 21 (1878) = N. rhodon ; N. stan Smith & Harris in Contrib. Gray Herb. 117: 39 (4987) oe NN. N. N. Se PET PPE RERAERES EERE oF z rhodoneu ce Reichb. f. in Bot. Zeit. 10: 835 : 1852). Sacciana Lind. Cogn. in Journ. Orch. 4: 73 (1893). Notylia arachnites Reichb. f. in Nederl. Kruidk. Arch. 4: 326 (1859). 56 N. boliviensis Schltr. in Repert. Spec. Nov. Beih. 10: 53 (1922). N. Buchtienii Schltr. in Fedde, Repert. Spec. Nov. 10: 458 (1912). N. lilacina Kranzl. in Fedde, Repert. Spec. Nov. 25: 29 (1928). ? N. micrantha Lindl. in Bot. Reg. 24: Misc. 91 (1838). Octomeria boliviensis Rolfe in Bull. N. Y. Bot. Gard. 4: 452 (1907). O. Buchtienii Schltr. in Fedde, Repert. Spec. Nov. 27: 57 (1929). O. grandiflora Lindl. in Bot. Reg. 28: Misc. 64 (1842). O. Guentheriana Kranzl. in Fedde, Repert. Spec. Nov. 25: 19 (1928). O. tenuis Schltr. in Fedde, Repert. Spec. Nov. 10: 455 (1912). ? Odontoglossum chiriquense Reichb. f. in Bot. Zeit. 10: 692 (1852). O. coronarium Lindl. Fol. Orch. Odont. 21 (1852). O. mapiriense Mansf. in Fedde, Repert. Spec. Nov. 36: 62 (1934). O. mystacinum Lindl. Fol. Orch. Odont. 6 (1852). O. rigidum Lindl. in Benth. Pl. Hartw. 152 (1844) = O. mystacinum. Oncidium aurarium Reichb. f. in Gard. Chron. 22: 394 (1884). aureum Lindl. Sert. Orch. sub t. 25 (1838). barbatum Lindl. Coll. Bot. t. 27 (1821-24). Batemannianum Parm. ex Knowl. & Westc. Flor. te 3: 183, t. 187 (1840). Baueri Lindl. Illustr. Gen. & Spec. Orch. t. 7 (18 Blanchetii Reichb. f. in Linnaea, 22: 845 (1849). bolivianense Oppenheim in Orchis, 10: 93 (1916). bolivianum Schltr. in Fedde, Repert. Spec. Nov, 10: 459 (1912) = O. Baueri. ete Rolfe in Bull. N. Y. Bot. Gard. 4: 452 (1907) = O. Batemannia- Schchrsaelll Kranzl. in Engler, Pflanzenr. IV, 50: 229 (1922). Buchtienii Schltr. in Fedde, Repert. Spec. Nov. 27: 81 (1929). disciferum Lindl. Fol. Orch. Oncid. 24 (1855). globuliferum HBK. Nov. Gen. & Spec. 1: 347 (1816). glossomystax Reichb. f. in Bot. Zeit. 10: 696 (1852). Herzogii Schltr. in Fedde, Repert. Spec. Nov. 12: 494 (1913) — O. barbatum. heteranthum Poepp. & Endl. Nov. Gen. 1: 34, t. 60 (1835). Jonesianum Reichb. f. in Gard. Chron. 1883 (II): 781. lepturum Reichb. f. in Gard. Chron. 1886 (I): 41. macropetalum Lindl. Sert. Orch. sub t. 48 (1841). Mandonii Reichb. f. Xen. Orch. 3: 21 (1878). Methonica Reichb. f. Linnaea, 41: 21 (1877). microxiphium Reichb. f. in Linnaea, 41: 21 (1877). otometon Kranzl. in Engler, Pflanzenr. IV, 50: 190 (1922). pusillum (L.) Reichb. f. in Walp. Ann. 6: 714 (1861). reductum Kranzl. in Fedde, Repert. Spec. Nov. 26: 344 (1929). retusum Lindl. in Bot. Reg. 25: sub t. 1920 (1837). Rusbyi Rolfe in Mem. Torr. Bot. Club, 4(3): 265 (1895). subulifolium Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 79 (1922) = O. bolivianense. superbiens Reichb. f. in Linnaea, 22: 848 (1849). trilingue Lindl. & Paxt. Flow. Gard. 1: 42 (1850). varicosum Lindl. in Bot. Reg. 25: sub. t. 1920 (1837). Weddellii Lindl. Fol. Orch. Oncid. 39 (1855). Williamsii Schltr. in Fedde, Repert. Spec. N~v. 10: 459 (1912). Wittii Oppenheim in Orchis, 10: 94 (1916). rnithidium bolivianum Schltr. in Fedde, Repert. Spec. Nov. 27: 78 (1929) = Maxillaria aurea var. gigantea. ae te Kranzl. in Fedde, Repert. Spec. Nov. 25:31 (1928) = Maxillaria So PES SIEOOR SSS PPS er ere Seat ors g999909 oO. rhombogloscum Schltr. in Fedde, Repert. Spec. Nov. 27: 78 (1929) = Max- illaria sp. Ornithocephalus gladiatus Hook. Exot. Fl. 2: 127 (1825). O. myrticola Lindl. in Ann, Mag. Nat. Hist. 4: 383 (1840). Pachyphyllum Cardenasii Smith & Harris in Contrib. Gray Herb. 114: 12 (1936). confusum Kranzl. in Engler, Pflanzenr. IV, 50: 22 (1923) = P. cyrtophyllum. cyrtophyllum Schltr. in Fedde, Repert. Spec. Nov. 15: 217 (1918). falcifolium Schltr. in Fedde, Repert. Spec. Nov. 10: 460 (1912) — P. cyrto- phyllum. Herzogii Schltr. in Meded. Rijks Herbar. 29: 80 (1916). minus Schltr. in Fedde, Repert. Spec. Nov. 10: 460 (1912). pectinatum Reichb. f. Xen. Orch. 3: 22 (1878). . pseudo-dichaea Reichb. f. Xen. Orch. 3: 22 (1878). Pelexia bonariensis (Lindl.) Schltr. in Beih. Bot. Centralbl. 37(2): 400 (1920). P. Fiebrigii Schltr. in Beih. Bot. Centralbl. 37(2): 402 (1920) Phragmipedium caricinum (Lindl.) Rolfe in Orch. Rev. 4: 331 (1896). Physosiphon andinum Schltr. in Fedde, Repert. Spec. Nov. 10: 451 (1912). P. Herzogii Schltr. in Fedde, Repert. Spec. Nov. 12: 485 (1913). Physurus anchoriferus Schltr. in Fedde, Repert. Spec. Nov. 12: 482 (1913) = Erythrodes sp. ? boliviensis Cogn. in O. Ktze. Rev. Gen. 3(2) : 300 (1898) = Erythrodes sp. ? Buchtienii Schltr. in Fedde, Repert. Spec. Nov. 27: 34 (1929) = Erythrodes ? Wet ty idty ae sp. ? Herzogii Schltr. in Fedde, Repert. Spec. Nov. 12: 482 (1913) — Erythrodes a Pleurothallis affinis Lindl. in Hook. Compan. Bot. Mag. 2: 554 (1837). agathophylla Reichb. f. Xen. Orch. 3: 25 (1878). amblyopetala Schltr. in Fedde, Repert. Spec. Nov. 12: 486 (1913). boliviana Reichb. f. in Bonplandia, 3: 224 (1855). Brittonii Rolfe in Bull. N. Y. Bot. Gard. 4: 449 (1907). Buchtienii Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 66 (1922). bulbophylloides Schltr. in Fedde, Repert. Spec. Nov. 27: 50 (1929). chamensis Lindl. Orch. Lindl. Orch. Linden. 2 (1846). coffeicola Schltr. in Fedde, Repert. Spec. Nov. 27: 50 (1929). complicata Rolfe in Mem. Torr. Bot. Club, 6: 121 (1896) = P. semipellucida. densifolia Rolfe in Mem. Torr. Bot. Club, 4(3): 260 (1895). diffusa Poepp. & Endl. Nov. Gen. 1: 49, t. 86 (1835). dolichocaulon Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 48 pisendd fraterna Lindl. Fol. Orch. Pleur. 3 (1859). frutex Schltr. in Fedde, Repert. Spec. Nov. 10: 454 (1912). Guentheri Schltr. in Fedde, Repert. Spec. Nov. 27: 51 (1929). herpethophyton Schltr. in Fedde, Repert. Spec. Nov. ak 52 (1929). Herzogii Schltr. in Fedde, Repert. Spec. Nov. 12: 487 (1913). lamellaris Lindl. Fol. Orch. Pleur. 8 (1859). linguifera Lindl. Fol. Orch. Pleur. 10 (1859). Mandonii Reichb. f. Xen. Orch. 3: 24 (1878). obovata (Lindl.) Lindl. in Bot. Reg. 28: Mise. 75 (1842). Ottonis Schltr. in Fedde, Repert. Spec. Nov. 27: 52 (1929). papuligera Schltr. in Fedde, Repert. Spec. Nov. 10: 453 (1912). papuligera var. macra Schltr. in Fedde, Repert. Spec. Nov. 10: 454 (1912). plumosa Lindl. in Bot. Reg. 28: Misc. 72 (1842). rhopalocarpa Schltr. in Fedde, Repert. Spec. “a Beih. 10: 43 (1922). P. riograndensis Rodr. Orch. Nov. 2: 28 (188 ruscifolia (Jacq.) R. Br. in Ait. Hort. Kew. aL 2) 5: 211 (18138). saltatoria Lindl. Fol. Orch. Pleur. 30 (1859 sanjanae Schltr. in Fedde, Repert. Spec. Nov. 12: 487 (19138). scabrata Lindl. Fol. Orch. Pleur. 30 (185 scabridula Rolfe in Mem. Torr. Bot. Club, sn 260 (1895). Oe de eaea, eee o1 ie. ¢) scleropus Schltr. in Fedde, Repert. Spec. Nov. 27: 53 (1929). semipellucida Reichb. f. in Linnaea, 22: 823 (1855). serrisepala Kranzl. in Fedde, Repert. Spec. Nov. 6: 18 (1908). simacoana Schltr. in Fedde, Repert. Spec. Nov. 27: 54 (1929). soratana Reichb. f. Xen. Orch. 3: 25 (1878). spathata Schltr. in Fedde, Repert. Spec. Nov. 27: 54 (1929). stenopetala Lindl. in Bot. Reg. 24: Misc. 95 (1838). tenuiflora Schltr. in Fedde, Repert. Spec. Nov. 12: 488 (1913). trialata Rolfe in Bull. N. Y. Bot. Gard. 4: 449 (1907) = P. triptera. tricarinata Poepp. & Endl. Nov. Gen. 1: 49, t. 87 (1835). triptera Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 68 (1922). tripterocarpa Schltr. in Fedde, Repert. Spec. Nov. 27: 55 (1929). triquetra Schltr. in Fedde, Repert. Spec. Nov. 12: 488 (1913). umbraticola Schltr. in Fedde, Repert. Spec. Nov. 27: 56 (1929) = vaginata. . vaginata Schltr. in Fedde, Repert. Spec. Nov. aa 19: 197 (1923). . Weddelliana Reichb. f. Xen. Orch. 3: 24 (1878). yungasensis Rolfe in Mem. Torr. Bot. Club, 4(3): 259 (1895). Polystachya boliviensis Schltr. in Fedde, Repert. Spec. Nov. 12: 483 (1918). P. simacoana Schltr. in Fedde, Repert. Spee, Nov. 27: 63 (1929) =P. bolivien- sis, at least in part Ponera mapiriensis Kiet in Fedde, Repert. Spec. Nov. 25: 22 (1928) = aphyglottis cuneata. Pinthiees cornuta Reichb. f. in Linnaea, 41: 18 (1877). elegans (Kranzl.) Schltr. in Fedde, Repert. Spec. Nov. 10: 447 (1912). Mandonii Reichb. f. Xen. Orch, 3: 18 (1878). Mandonii var. bilobipetala Hoehne in FI. Brasil. 12(2): 150 (1945). montana Lindl. in Benth. Pl. Hartw. 155 (1845). unguiculata Ames & Schweinf. in Ames, Sched. Orch. 8: 9 (1925). terichis Bangii Rolfe in Bull. N. Y. Bot. Gard. 4: 448 (1907). boliviana Schltr. in Fedde, Repert. Spec. Nov. 9: 436 (1911). galeata Lindl. Gen. & Spec. Orch. 445 (1840). Mandonii (Reichb. f.) Rolfe in Bull. N. Y. Bot. Gard. 4: 449 (1907). saxicola Schltr. in Fedde, Repert. Spec. Nov. 12: 482 (1913). silvestris Schltr. in Fedde, Repert. Spec. Nov. 10: 446 (1912). yungasensis Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 37 (1922). Rodriguezia corydaloides Krianzl in Fedde, Repert. Spec. Nov. 25: 24 (1928). Rusbyella caespitosa Rolfe in Mem. Torr. Bot. Club, 6: 122 (1896). Sarcoglottis Herzogii Schltr. in Fedde, Hacert Spec. Nov. 21: 332 (1925) = Spiranthes sp. ? S. picta (Anders.) Kl. in Allg. Gartenzeit. 10: 106 (1842) — Spiranthes sp. ? Scaphyglottis boliviana Schltr. in Fedde, Repert. Spec. Nov. Beih. 10: 44: (1922). S. cuneata Schltr. in Bot. Centralbl. Beih. 36(2): 398 (1918). S. Summersii L. O. Wms. in Bot. Mus. Leafl. Harvard Univ. 9: 14, t. 4 (1940). Sievekingia Trollii Mansf. in Fedde, Repert. Spec. Nov. 36: 61 (1984). Sigmatostalix Buchtienii Kranzl. in Fedde, Repert. Spec. Nov. 25: 24 (1928). S. graminea (Poepp. & Endl.) Reichb. f. in Bot. Zeit. 10: 769 (1852). Sobralia boliviensis Schltr. in Fedde, Repert. Spec. Nov. 12: 491 (1913). . Buchtienii Schltr. in Fedde, Repert. Spec. Nov. 27: 30 (1929). caloglossa Schltr. in Fedde, Repert. Spec. Nov. 27: 31 (1929). chrysoleuca Reichb. f. Xen. Orch. 2: 179 (1874). dichotoma R. & P. Syst. 232 (1798). D’Orbignyana Reichb. f. Xen. Orch. 2: 179 (1874). fruticetorum Schltr. in Fedde, Repert. Spec. Nov. 12: 492 (1913). Herzogii Schltr. in Fedde, Repert. Spec. Nov. 12: 492 (1913). Mandonii Reichb. f. Xen. Orch. 2: 175, t. 174 (1874) = S. dichotoma. eat oO ieee eee AT oe WaT Ie Se ee RAMAN Minin 59 parviflora L. O. Wms. in Lilloa, 3: 475 (1938). rupicola Krinzl. in Fedde, Repert. Spec. Nov. 6: 21 (1908). . Sancti-josephi Kranzl. in Fedde, Repert. Spec. aN 25: 22 (1928). scopulorum Reichb. f. Xen. Orch. 2: 176 (1874). semperfiorens Krinzl. in Viertelj. Naturf. Ges. Ziirich, 60: 428 (1915). setigera Poepp. & Endl. Nov. Gen. 1: 54 (1835). S. violacea Lindl. Orch. Linden. 26 (1846). Spiranthes acaulis (J. E. Sm.) Cogn. in Mart. Fl. Bras. Nl 221 (1895). S. Castillonii (Haumann) L, O. Wms. in Lilloa, 3: 478 (1938). S. chlorops Reichb. f. Xen. Orch. 3: 20 (1878 ? S. costaricensis Reichb, f. in Bonplandia, 3: 214 (1855). S. elata (Sw.) L. C. Rich. in Mém. Mus. Paris, 4: 59 (1818). S. goodyeroides Schltr. in Fedde, Repert. Spec. Nov. 10: 448 (1912). Ss. ptirs od (Rolfe) C. Schweinf. in Bot. Mus. Leafl. Harvard Univ. 10: 30 (194 Ss. ee Reichh. f. Xen. Orch. 3: 21 (1878). S. orchioides (Sw.) A. Rich. Fl. Cub. Fanerog. 2: 252 (1853). S. plantaginea Lindl. Gen. & Spec. Orch. 468 (1840). Ss. ee came Rolfe in Mem. Torr. Bot. Club, Pas 266 (1895). ? Stanhopea eburnea Lindl. in Bot. Reg. 18: t. 1529 (1832). Stelis atrobrunnea Schltr. in Fedde, Repert. Spec. Nov. 27: 40 (1929). S. Bangii Rolfe in Mem. Torr. Bot. Club, 4(8): 260 (1895). S. boliviensis Rolfe in Bull. N. Y. Bot. Gard. 4: 450 (1907). S. Brittoniana Rolfe in Mem. Torr. Bot. Club, 4(8): 261 (1895). S. Buchtienii Schltr. in Fedde, Repert. Spec. Nov. 10: 451 (1912). S. campanulifera Lindl. Fol. Orch. Stel. 3 (1858), S. casanaénsis Schltr. in Fedde, Repert. nag Nov. 27: 40 (1929). S. connata Presl, Rel. Haenk. 1: 103 (1827). S. discolor Reichb. f. in Bonplandia, 3: 240 (1855). S. euspatha Reichb. f. in Bonplandia, 3: 225 (1855). S. filiformis Lindl. Fol. Orch. Stel. 6 (1859). S. flexa Schltr. in Fedde, Repert. Spec. Nov. 27: 41 (1929). S. Herzogii Schltr. in Fedde, Repert. Spec. Nov. 12: 484 (1913). S. heterosepala Schitr. in Fedde, Repert. Spec. Nov. 27: 42 (1929). S. iminapensis Reichb. f. Xen. Orch. 3: 25 (1878) S. laxa Schltr. in Fedde, Repert. Spec. Nov. 10: 451 (1912). S. macrantha Rolfe in Bull. N. Y. Bot. Gard. 4: 450 (1907). S. Mandoniana Schltr. in Fedde, Repert. Spec. Nov. 27: 42 (1929). S. microtatantha Schltr. in Fedde, Repert. Spec. Nov. 27: 43 (1929). S. naviculigera Schltr. in Fedde, Repert. Spec. Nov. 27: 44 (1929). S. Ottonis Schltr. in Fedde, Repert. Spec. Nov. 27: 45 (1929). S. phaeomelana Schltr. in Fedde, Repert. Spec. Nov. 27: 45 (1929). S. polycarpa Schltr. in Fedde, Repert. Spec. Nov. 27: 46 (1929). S. Rusbyi Rolfe in Mem. Torr. Bot. Club, 4(3): 261 (1895). S. saxicola Schltr. in Fedde, Repert. Spec. Nov. 10: 452 (1912). S. scandens Rolfe in Bull. N. Y. Bot. Gard. 4: 451 (1907). S. setacea Lindl. Fol. Orch. Stel. 3 (1859). S. simacoénsis Schltr. in Fedde, Repert. Spec. Nov. 27: 47 (1929). S. tenuicaulis Lindl. Fol. Orch. Stel. 16 (1859). S. trianguliflora Schltr. in Fedde, Repert. Spec. Nov. 27: 48 (1929). ? S. tricardium Lindl. Fol. Orch. Stel. 14 (1859). S. triseta Lindl. Fol. Orch. Stel. 17 (1859). S. vagans Schltr. in Fedde, Repert. Snec. Nov. 27: 48 (1929). S. virens Schltr . in Fedde, Repert. Spec. Nov. 12: 484 (1913). S. xanthantha Schltr. in Fedde, Repert. Spec. Nov. 10: 452 (1912). S. yungasensis Schltr. in Fedde, Repert. Spec. Nov. 27: 49 (1929). DAMP TA Ip 60 Stenoptera acuta Lindl. Gen. & Spec. Orch. 447 (1840). S. Guentheriana Krinzl. in Fedde, Repert. Spec. Nov. 25: 19 (1928). S. plantaginea Schltr. in Fedde, Repert. Spec. Nov. 10: 446 (1912). Stenorrhynchus apetalus Kranzl. in Fedde, Repert. Spec. Nov. 6: 23 (1908). aphyllus Lindl. Gen. & Spec. Orch. 478 (1840). australis Lindl. Gen. & Spec. Orch. 477 (1840). comosus Cogn. in O. Ktze. Rev. Gen. 3(2): 299 (1898). macranthus (Reichb. f.) Cogn. in Mart. Fl. Bras. 3(4): 176, t. 40 (1895). . sancti-josephi Krinzl. in Fedde, Repert. Spec. Pe a 6: 22 (1908). Teliposon Benedicti Reichb. f. in Linviaea. 41: 3 (18 Trichoceros parviflorus HBK. Nov. Gen. & Spec. 1: ph t. 76 (1816). Trichopilia fragrans (Lindl.) Reichb. f. in Hamb. Gartenzeit. 14: 229 (1858). Trizeuxis andina Schitr. in Fedde, Repert. Spec. Nov. Beih. 10: 52 (1922) = ‘abeaiads . falcata. T. falcata Lindl. Coll. Bot. t. 2 (1821). Vanilla odorata Presl, Rel. Haenk. 1: 101 (1827). V. palmarum Lindl. Gen. & Spec. Orch. a a ei a V. pompona Schiede in Linnaea, 4: 573 (18 Xylobium Buchtienianum Kranz. in Orchis, 7 429 (1908). X. flavescens Schltr. in Fedde, Repert. Spec. Nov. 12: 493 (1913). X. latifolium Schltr. in Fedde, Repert. Spec. Nov. 27: 66 (1929) = X. pal- lidiflorum X. miliaceum (Reich. f.) Rolfe in Orch, Rev. 20: 43 ue X. pallidiflorum (Hook.) Nicholson, Dict. Gard. 4: 225 (1887). ». ria (Lindl.) Rolfe ex Gentil, Pl. Cult. Jard. Bot. Bruxelles, 194 05) = X. squalens > 2 sch (Lindl.) Lindl. in Bot. Reg. 11: sub t. 897 (1825 X. varicosum (Lindl.) Rolfe in Mem. Torr. Bot. Club, 4(3): 268 (1895). Zygopetalum bolivianum Schltr. in Fedde, Bore Spec. Nov. Beih. 10: 50 = Z. intermedium var. peruvian Z. intermedium var. peruvianum Rolfe in Lindentat 9: T1, t. 418 (1893). DICOTYLEDONEAE PIPERACEAE Peperomia aceramarcana Trel. in Bull. Torr. Bot. Club, 55: 169 (1928). P. aceramarcana var. variifolia Yuncker in Lilloa, 27: 199, t. 76 (1955). P. aceroana C. DC. in Bull. Torr. Bot. Club, 25: 572 (1898). P, adenocarpa C. DC. in Bull. Torr. Bot. Club, 25: 569 (1898). P. alata R. & P. Fl. Peruv. 1: 31, t. 48 (1798) P. alata var. angustifolia C. DC. Prodr. 16(1): 418 (1869) P. angustata HBK. Nov. Gen. & Spec. 1: 68 (1815). P. apodostachya Yuncker in Lilloa, 27: 251, t. 128 (1955). P. argyreia (Miq.) E. Morr. in Belg. Hort. 17: 2, t. 2 (1867). P. Balansana C. DC. in Mém. Soc. Phys. Hist. Nat. Genéve, 27(2): 313, t. 10 (1882). P. Bangii C DC. in Bull. Torr. Bot. Club. 19: 49 (1892). P. barbaranoides Yuncker in Lilloa, 27: 173, se 53 (1955). P. blanda HBK. Nov. Gen. & Spec. 1: 67 (18 P. blanda var. Langsdorfii (Miq.) AAS ce in Nov. Act. Soc. Sci. Upsal. (ser. 3) 8: 39 (1873). P. blanda var, pseudo-dindygulensis (C. DC.) Yuncker in Lilloa, 27: 218 (1955). P. boliviensis C. DC. in DC, Prodr. 16(1): 453 (1898) = P. reflexa ? ... ve oe Pe MNSANU Nhe wees eS bopiana Trel. in Mem, N. Y. Bot. Gard. 7: 227 (1927). brevispica C. DC. in Bull. Torr. Bot. Club, 25: 570 (1898). Brittonii C. DC. in Bull. Torr. Bot. Club, 19: 254 (1892). Buchtienii Yuncker in Lilloa, 27: 241, t. 118 (1955). canaminana Trel. in Mem. N. Y. Bot. Gard. 7: 228 (1927). Cardenasii Trel. in Mem. N. Y. Bot. Gard. 7: 227 (1927). carnifolia Yuncker in Lilloa, 27: 193, t. 70 (1955 chromatogena Yuncker in Lilloa, 27: 278, t. 159 (1955). chromatogena var. subpeltata Yuncker in Lilloa, 27: 279 (1955). circinnata Link in Bot. Jahrb. 1: 64 (1820). comarapana C. DC. in Meded. Rijks Herbar. 27: 8 (1915). coroicensis Yuncker in Lilloa, 27: 237, t. 114 (1955). cyclaminoides A. W. Hill in Ann. Bot. 21: 149, t. 15, on 16-17 (1907). cyclophylla Miq. ex Seeman, Bot. Voy. Herald, 198 (1854). divaricata Yuncker in Lilloa, 27. 203, t. 81 (1955 duidana Trel. in Bull. Torr. Bot. Club, 58: 354 (19 dumeticola. C. DC. in Bull. Torr. Bot. Club, a se ST il == P. trinervis ? effusa Yuncker in Lilloa, 27: 240, t. 117 (195 efimbriata Trel. in Mem. N. Y. Bot. Gard. 7: ase (1927). elegans C. DC. in DC. Prodr. 16(1): 430 (1869). elongata HBK. Nov. Gen. & Spec. 1: 62 (1815 elongata var. piliramea Trel. & Yuncker, Piperac. North. S. Amer. 659 (1950). emarginella (Sw.) C. DC. in DC. Prodr. 16(1): 437 (1869). emarginella forma glabrior C. DC. in Fedde, Repert. Spec. Nov. 9: 235 (1911) = P. emarginella. Fiebrigii C. DC. in Fedde, Repert. Spec. Nov. 14: 397 (1916). Fiebrigii forma glabrata Yuncker in Lilloa, 27: 185 (1955). galiifolia Trel. in Mem. N. Y. Bot. Gard. 7: 227 (1927) = P. galioides. galioides HBK. Nov. Gen. & Spec. 1: 71, t. 17 (1815). galioides var. longifolia C. DC. in DC. Prodr. 16(1): 464 (1869) = P. galioides. : glabella (Sw.) A. Dietr. Sp. Pl. 1: 156 (1831). glabella var. nervulosa (C. DC.) Yuncker in Ann. Missouri Bot. Gard. 37: 98 (1950). Herzogii C. DC. in Meded. Rijks Herbar. 27: 9 (1915). heterophylla Miq. in Hook. Lond. Journ. Bot. 4: 415 (1845). heterophylla var. grandis Yuncker in Lilloa, 27: 244, t. 121 (1955). hispidula (Sw.) A. Dietr. Sp. Pl. 1: 165 (1831). hispidula var. muscophila (C. DC.) C. DC. in Candollea, 1: 335, 392 (1923). hispiduliformis Trel. in Lilloa, 6: 295 (1941). imbracteata Yuncker in Lilloa, 27: t. 55 (1955). Klotzschiana Mig. Syst. Pip. 145 (184 Kuntzei C. DC. in O, Ktze. Rev. ae ete 272 (1898). lancifolia Hook. Ic. Pl. 4: t. 332 (184 larecajana C. DC. in DC. Prodr. a 406 (1869). larecajana var. angustifolia Yuncker in Lilloa, 27: 170 (1955). : Lorentzii var. boliviensis Yuncker in Lilloa, 27: 198, t. 75 (1955). macrostachya (Vahl) A. Dietr. Sp. Pl. 1: 149 (1831) Mandonii C. DC. in DC. Prodr. 16(1): 395 (1869). Mandonii var. excelsis C. DC. in Bull. Torr. Bot. Club, 21: 161 (1894) = P. Mandon sige Trel. in Macbr. in Field. Mus. Publ. Bot. 13(2): 61 (1936). - multifolia Yuncker in Lilloa, 27: 227, t. 103 (1955). multispica C. DC. in Bull. Torr. Bot. Club, 25: 571 (1898). for) bo WU WM COE DRRBE oO DE hyd ty hd ty ty nequejahuirana Trel. in Bull. Torr. Bot. Club, 55: 169 (1928). nudicaulis C, DC. in Bull. Torr. Bot. Club, 19: 48 (1892). obtusifolia (L.) A. Dietr. Sp. Pl. 1: 154 (1831). okarana Trel. in Bull. Torr. Bot. Club, 55: 170 (1928) = P. galioides. olens C. DC. in Fedde, Repert. Spec. Nov. 13: 307 (1914). Ottoniana forma boliviensis C. DC. in Fedde, Repert. Spec. Nov. 13: 308 (1914) = P. Buchtienii. parvifolia C. DC. in Journ. Bot. 4: 133 (1866). peltifolia C. DC. in Bull. Torr. Bot. Club, 25: 570 (1898). pentadactyla Yuncker in Lilloa, 27: 283, t. 164 (1955). perlongipes C. DC. in Fedde, Repert. Spec. Nov. 13: 308 (1914). perlongispica Yuncker in Lilloa, 27: 277, t. 157 (1955). peruviana (Miq.) Dahlst. in Kgl. Sv. Vet. Akad. Handl. 33(2): 32 (1900). peruviana var. major A. W. Hill in Ann. Bot. 21: 151 (1907). pitiguayana Trel. in Bull. Torr. Bot. Club, 55: 169 (1928). procumbens C. DC. in DC Prodr. 16(1): 435 (1869). pseudo-cobana Yuncker in Lilloa, 27: 256, t. 133 Sail pseudofureata C. DC. in DC. Prodr. 16(1): 399 (1869). pseudo-rufescens C. DC. in Bull. Torr. Bot. Club, 21: 160 (1894). pseudosilvarum Yuncker in Lilloa, 27: 200, t. 77 (1955). . pseudo-umbilicata Yuncker in Lilloa, 27: 275, t. 155 (1955). psilophylla C. DC. in Bull. Torr. Bot. Club, 21: 160 (1894). psilostachya C. DC. in Mém. Soe. Phys. Hist. Nat. Genéve, 32(pt. 1, no. 2): 9, t. 54, fig. 6-9 (1893). punctulatissima Trel. in Mem. N. Y. Bot. Gard. 7: 228 (1927). quaesita Trel. in Macbr. in Field Mus. Publ. Bot. 18(2): 83 (1936). reflexa (L. f.) A. Dietr. Sp. Pl. 1: 180 (1831). reflexa forma robustior C. DC. in Fedde, Repert. Spec. Nov. 13: 304 (1914). reflexa forma variipila Yuncker in Lilloa, 27: 192 (1955). retropuberula Yuncker in Lilloa, 27: 184, t. 64 (1955). rhombea R. & P. Fl. Peruv. 1: 31, t. 46, fig. b. (1798). rhombifolia Trel. in Mem. N. Y. Bot. Gard. 7: 226 (1927). rotundifolia HBK. Nov. Gen. & Spec. 1: 67, t. 12 (1815). rotundifolia var. pilosior (Miq.) C. DC. in Urb. Symb. Ant. 3: 230 (1902). rurrenabaqueana Trel. in Mem. N. Y. Bot. Gard. 7: 226 (1927 Rusbyi C. DC. in Bull. Torr. Bot. Club, 19: 49 (1892). saxicola C. DC. in Bull. Torr. Bot. Club, 25: 571 (1898). seutifolia C. DC. in Notizbl. 7: 497 (1917). scutilimba Yuncker in Lilloa, 27: 275, t. 154 (1955). se — C. DC. in Fedde, Repert. Spec. Nov. 13: 305 (1914) = P. angust serpens ope Loud. Hort. Brit. 13 (1830). silvarum C, DC. in Fedde, Repert. Spec. ee 13: 305 (1914). sirupayana C. DC. in Candollea, 1: 361 (1923). soratana C. DC. Bull. Torr. Bot. Club, 25: 569 (1898). Steinbachii Yuncker in Lilloa, 27: 234, t. 113 (1955). Stuebelii C. DC. in Bull. Torr. Bot. Club, 19: 255 (1892). Stuebelii var. glabrata Yuncker in Lilloa, 27: 181 (1955). suspensa C. DC, in Fedde, Repert. Spec. Nov. 13: 304 (1914). Swartziana Miq. Syst. Pip. 155 (1848). talinifolia HBK. Nov. Gen. & Spec. 1: 62, t. 8 (1815). talinifolia var. longipetiolata C. DC. in Bull. Torr. Bot. Club, 27: 182 (1900) = P. talinifolia. Tatei Yuncker in Lilloa, 27: 220, t. 95 (1955). tenuipeduncula C. DC. in Fedde, Repert. Spec. Nov. 13: 306 (1914). ternata C. DC. in Bull. Herb. Boiss. 6: 509 (1898). . tetragona R. & P. Fl. Peruv. 1: 31, t. 47 (1798). . Theodori Trel. in Lilloa, 5: 356 (1940). . Theodori var. glabricaulis Yuncker in Lilloa, 27: 189 (1955). . ticunhuayana Trel. in Bull. Torr. Bot. Club, 55: 170 (1928). . tominana C. DC. in Bull. Torr. Bot. Club, 25. 572 (1898). . tominana forma pubifolia Yuncker in Lilloa, 27: 186 (1955). . trinervis R. & P. Fl. Peruv. 1: 32, t. 48, fig. b (1798). unduavina C. DC, in Fedde, Repert. Spec. Nov. 13: 306 (1914). vestita C. DC. in Bull. Torr. Bot. Club, 25: 568 (1898). . Williamsii C. DC. in Ann. Conserv. & Jard. Bot. Genéve, 21: 248 (1920). yanacachiana Yuncker in Lilloa, 27: 228, t. 104 (1955). yungasana C. DC. in Fedde, Repert. Spec. Nov. 13: 306 (1914). Piper acutifolium R. & P. Fl. Peruv. 1: 38, t. 64, fig. a (1798). aduncum L. Sp. Pl. 1: 29 (1758). aduncum var. cordulatum (C. DC.) Yuncker in Lilloa, 27: 129 (1955). aduncum var. Garcia-Barrigae Trel. & Yuncker, Pip. North. S. Amer. 251 (1950 erhoseamn Aubl. Pl. Guian. Fr: 1: 23 (2715); . Bangii C. DC. in Bull. Torr. Bot. Club, 19: 254 (1892). . Bangii var. pubinervium Yuncker in Lilloa, 27: 105 (1955). . Bartlingianum (Miq.) C. DC. in DC. Prodr. 16(1): 257 (1869) . benianum Trel. in Mem. N. Y. Bot. Gard. 7: 222 (1927) = P. callosum. . bolivianum C. DC. in DC. Prodr. 16(1): 280 (1869). . bopianum Trel. in Mem. N. Y. Bot. Gard. 7: 223 (1927) = P. longesty- Dik eee ache erase se, ty . Buchtienii C. DC. in Fedde, Repert. Spec. Nov. 9: 230 (1911). Buchtienii var. charopampanum (C. DC.) Yuncker in Lilloa, 27: 161 (1955). callosum R. & P. FI. Peruv. 1: 34, t. 53, fig. a (1798). . Cardenasii Ca in Mem. N. Y. Bot. Gard. 7: 223 (1927) = P. aduncum var. cordulat cataractarum Trel. in Mem. N. Y. Bot. Gard. 7: 224 See = P. hispidum. cingens C. DC. in Fedde, Repert. Spec. Nov. 13: 311 (19 coriaceilimbum C. DC. in Fedde, Repert. Spec. Nov. . wai (1911). cuspidibracteatum Yuncker in Lilloa, 27: 107, t. 4, fig. 4 (1955). duidaénse var. bolivianum Yuncker in Lilloa, 27: 162, t. 42 (1955). elliptico-oblongifolium ih in ites N. Y. Bot. Gard. 7: 224 (1927) = P. Buchtienii var. charopampanu songatan Vahl, Enum. 1: "8 (4808). guanaianum C. DC. in Fedde, Repert. Spec. Nov. 9: 232 (1911) = P. adun- cum var. cordulatum. Herzogii C. DC. in Meded. Rijks Herbar. 27: 7 (1915) = P. aduncum var. Garcia-Barrigae. Hieronymi C. DC. in O. Ktze. Rev. Gen. 3(2): 273 (1898). hispidum Sw. Prodr. 15 (1788). . hispidum var. magnifolium (C. DC.) C. DC. in Urb. Symb. Ant. 3: 188 (1902). hispidum var. trachydermum (Trel.) Yuncker in Ann. Missouri Bot. Gard. 37:33 (1950). Kuntzeit C. DC. in O. Ktze. Rev. Gen. 3(2): 274 (1898) = P. aduncum. laevigatum HBK. Nov. Gen. & Spec. 1: 56 (1815). laevilimbum C. DC. in Fedde, Repert. Spec. Nov. 9: 232 (1911). lanceolatum R. & P. Fl. Peruv. 1:36, t. 61, fig. b (1798). longestylosum C. DC. in Bull. Torr. Bot. Club, 21: 161 (1894). longichaetum Yuncker in Lilloa, 27: 127, t. 15 (1955). mapirense C. DC. in Bull. Torr. Bot. Club, 19: 47 (1892). DUM UN MD BM WN UY Pe TO TY tO aD 1 ee SL agaga See eee es PPR. NSIS. Ss UN NS ee mapirense var. magnifolium C. DC. in Fedde, Repert. Spec. Nov. 9: 230 (1911) = P. mapirense. marequitense C. DC. in DC. Prodr. er: 290 (1869). medium Jacq. Ic. Pl. Rar. 1: 2, t. 8 (1781). medium var. perglabrum Yuncker in Lilloa, 27: 118 (1955). mercens Yuncker in Lilloa, 27: 135, t. 19, fig. 1 (1955 microtrichum C. DC. in Fedde, Repert. Spec. Nov. 13: 310 wore nigriconnectivum C. DC. in Meded. Rijks Herbar. 27: 7 (1915). nigro-granulatum Trel. in Mem. N. Y. Bot. Gard. 7: 224 (1927) — P. lanceolatum. obovatum var. bolivianum C. DC. in Ann. Conserv. & Jard. Bot. Genéve, 21: 237 (1920) = P. Buchtienii. oxyphyllum C. DC. in Bull. Torr. Bot. Club, 19: 48 (1892). pachyphloium C. DC. in O. Ktze. Rev. Gen. 3(2): 274 (1898). pachyphloium var. macrophyllum Yuncker in Lilloa, 27: 107, t. 4, io: 1955). peltilimbum Yuncker in Lilloa, 27: 155, t. 37 (1955). percostatum Yuncker in Lilloa, 27: 157, t. 39 (1955). perscabrifolium Yuncker in Lilloa, 27: 111, t. 7, fig. 1 (1955). pilirameum C. DC. in Fedde, Repert. Spec. Nov. 9: 229 (1911). pilirameum var. soca Yuncker in Lilloa, 27: 110 (1955). praeacutilimbum C. DC. in Fedde, Repert. Spec. Nov. 9: 231 (1911). pseudogrande Yuncker in Lilloa, 27: 146, t. 27, fig. 1 (1955). psilophyllum C. DC. in Bull. Torr. Bot. Club, 19: 47 (1892). psilophyllum forma longepedunculatum Yuncker in Lilloa, 27: 148, t. 28 (1955). puberulinerve C. DC. in Fedde, Reeprt. Spec. Nov. 9: 231 (1911). puberulinerve var. subarborescens (C. DC.) Yuncker in Lilloa, 27: 149 (1955). pubiovarium Yuncker in Lilloa, 27: t. 25, fig. 2 (1955). punctulantherum C. DC. in Fedde, Repert. Spec. Nov. 9: 233 (1911). rectispicum Trel. in Mem. N. Y. Bot. Gard. 7: 225 Oe = P. mapirense. rogaguanum Trel. in Mem. N. Y. Bot. Gard. 7: 225 (1927). rurrenabaqueanum Trel. in Mem. N. Y. Bot. Gard. 7: 225 (1927) = P. Buchtienii. Rusbyi C. DC. in Bull. Torr. Bot. Club, 19: 47 (1892). Rusbyi var. hirsutum Yuncker in Lilloa, 27: 139 (1955). sanctae-crucis C. DC. in O. Ktze. Rev. Gen. 3(2): 274 (1898) = P. mapi- rense. semimetrale C. DC. in Fedde, Repert. Spec. Nov. 9: 232 (1911). Steinbachii Yuncker in Lilloa, 27: 136, t. 186 (1955). subsilvulanum C. DC. in Notizbl. 6: 453 (1917). svidaefolium Trel in Mem. N. Y. Bot. Gard. 7: 225 (1927) = P. subsilvu- trichogynum C. DC. in Fedde, Repert. Spec. Nov. 9: 233 (1911). trichorhachis C. DC. in Bull. Torr. Bot. Club, 25: 566 (1898). trigoniastrifolium C. DC. in Bull. Torr. Bot. Club, 25: 567 (1898). tuberculatum Jacq. Collect. 2: 2, t. 211 (1788). tuberculatum var. minus C. DC. in DC. Prodr. ae ge 266 (1869). tumupasense Yuncker in Lilloa, 27: 106, t. 2, fig. 2 umbrigaudens C. DC. in Fedde, Repert. Spec. Nov. cy 310 (1614) == P: Rusbyi. unduavinum C. DC. in Fedde, aa tid Spec. Nov. 13: 309 (1914) = P. psilophyllum. Pothomorphe peltata (L.) Mig. Comm. Phyt. 87 (1840 Fi (1840). umbellata var. vestita (C. DC.) Yuncker in Lilloa, 27: 166 (1955). CHLORANTHACEAE Hedyosmum brasiliense Miq. in Mart. Fl. Bras. 4(1): 3 (1852). H. bolivianum Cordem. in Adansonia, 3: 306 (1863) = H. racemosum. H. Mandonii Solms-Laub. in DC. Prodr. 16(1): 480 (1869). H. maximum (0. Ktze.) K. Schum. in Just, Bot. Jahresber. 26(1): 345 (1900). H. racemosum (R. & P.) G. Don, Gen. Hist. 3: 434 (1834). SALICACEAE Salix Humboldtiana Willd. Sp. Pl. 4(2): 657 (1806). MYRICACEAE Myrica arguta HBK. Nov. Gen. & Spec. 2: 17, t. 98 (1817). M. costata Rusby, Descr. S. Am. Pls. 8 (1920). M. macrocarpa HBK. Nov. Gen. & Spec. 2: 16 (1817). M. xalapensis HBK. Nov. Gen. & Spec. 2: 16 (1817). J UGLANDACEAE Juglans boliviana Dode in Bull. Soc. Dendrol. France, 13: 211 (1909). CORYLACEAE Alnus acuminata HBK. Nov. Gen. & Spec. 2: 20 (1817). A. jorullensis var. ferruginea (HBK.) O. Ktze. Rev. Gen. 3(2): 295 (1898). A. jorullensis var. Mirbelii (Spach) H. Winkl. in Engler, Pflanzenr. IV, 61: 126 (1904). ULMACEAE Ampelocera sp mit A bliviana Planch. in Ann. Sci. Nat. (ser. 3) 10: 311 (1848) = C. pube- C. ‘iccusanan Planch. in Ann. Sci. Nat. (ser. 3) 810 (1848). iguanaea (Jacq.) Sarg. Silv. N. Am. 7: 64 (1895). morifolia Planch. in Ann. Sci. Nat. (ser. 3) 10: 311 (1848). pubescens (HBK.) Spreng. Syst. Veg. 1: 931 (1825). pubescens var. Chichape (Wedd.) Baehni in Candollea, 7: 201 (1936). spinosa Spreng. Syst. Veg. 1: 931 (1825). spinosa var. Weddelliana (Planch.) Baehni in Candollea, 7: 204 (1936). Tala var. Gilliesiana forma velutina Herzog in Meded. Rijks Herbar. 27: 72 (1915) = C. pubescens var. Chichape. . triflora (Klotzsch) Mig. in Mart. Fl. Bras. 4(1): 181 (1855). Williamsii Rusby in Bull. N. Y. Bot. Gard. 6: 497 (1910) = C. iguanaea. Lozanella permollis Killip & Morton in Journ. Wash. Acad. Sci. 21: 338 (1931). Momisia crenata Wedd. in Ann. Sci. Nat. (ser. 3) 18: 195 (1852) = Celtis An afgAnaaas pubesce M. in "Wedd. in Ann. Sci. Nat. (ser. 3) 18: 195 (1852) = Celtis pube- scens. M. tarijensis Wedd. in Ann. Sci. Nat. (ser. 3) 18: 194 (1852) = Celtis tri- flora. Phyllostylum orthopterum Hall. f. in Meded. Rijks Herbar. 27: 70 (1915). Trema affinis (Planch.) Blume in Mus. Bot. Lugd. Bat. 2: 58 (1852). T. micrantha (L.) Blume in Mus. Bot. Lugd. Bat. 2: 58 (1852) 66 MORACEAE Batocarpus amazonicus (Ducke) Fosberg in Proc. Biol. Soc. Wash. 55: 101 2). Brosimum Gaudichaudii Trécul in Ann. Sci. Nat. (ser.3) 8: 140 (1847). B. guianense (Aubl.) Huber ex Ducke in Bol. Mus. Goéldi, 7: 172 (1913). Cecropia angustifolia Trécul in Ann. Sci. Nat. (ser. 3) 8: 83 (1847). C. elongata Rusby in Bull. N. Y. Bot. Gard. 4: 446 (1907). C. ficifolia Warb. ex Snethlage in Notizbl. 8: 365 (1923). C. latiloba Miq. in Mart. Fl. Bras. 4(1): 147 (1853). C. leucocoma Migq. in Mart. Fl. Bras. 4(1): 142 (1853). C. obovata Rusby in Bull. N. Y. Bot. Gard. 6: 498 (1910). C. Tessmannii Mildbr. in Notizbl. 9: 260 (1925). Chlorophora reticulata Herzog in Meded. Rijks Herbar. 27: 73 (1915). C. tinctoria (L.) Gaud. in Freyc. Voy. Bot. 508 (1827), combination implied but not actually made. Clarisia biflora R. & P. Syst. 256 (1798). C. ilicifolia (Spreng.) Lanj. & Rossb. in Meded. Bot. Mus. Utrecht, 36: 717 (1936). Coussapoa asperifolia Trécul in Ann. Sci. Nat. (ser. 3) x 96 (1847). C. boliviana Standl. in Field Mus. Publ. Bot. 17: 159 (1937). C. ovalifolia Trécul in Ann. Sci. Nat. (ser. 3) 8: 95 (1847). Dorstenia brasiliensis Lam. Encyc. 2: 317 (1786). Duguetia glabra Britton in Bull. Torr. Bot. Club, 16: 14 (1889) == Ogcodeia Sandwithiana. D. montana Herzog in Meded. Rijks Herbar. 27: 73 (1915). Ficus bopiana Rusby in Mem. N. Y. Bot. Gard. 7: 230 (1927). caballina Standl. in Field Mus. Publ. Bot. 13: 301 (1937). coerulescens (Rusby) Rossberg in Fedde, Repert. Spec. Nov. 42: 61 (1937). — (Herzog) Herter in Rev. Sudam. Bot. 6: 151 (1940). a Ruiz ex Mig. in Mart. Fl. Bras. 4(1): 98 (1853), in synon. ee A. D. Hawkes in Phytologia, 3:32 (1948). Killipii Standl. in Field Mus. Publ. Bot. 13: 304 (1937). Macbridei Standl. in Field Mus. Publ. Bot. 13: 305 (1937). Matthewsii (Miq.) Miq. in Ann. Mus. Bot. Lugd. Bat. 3: 298 (1867). oblanceolata Rusby in Bull. N. Y. Bot. Gard. 4: 446 (1907) = F. Matthewsii. oblanceolata Rusby in Bull. N. Y. Bot. Gard. 6: 498 (1910) — F. Kath- erinae. paraénsis Mig. in Ann. Mus. Bot. Lugd. Bat. 3: 298 (1867). radula Willd. Sp. Pl. 4(2): 1144 (1806). regularis Standl. in Field Mus. Publ. Bot. 17: 175 (1937). Ruiziana Standl. in Field Mus. Publ. Bot. 13(2): 307 (1987). Whitei Rusby in Mem. N. Y. Bot. Gard. 7: 230 (1927). Williamsii (Rusby) Rossberg in Fedde, Repert. Spec. Nov. 42: 61 (1937). Wuiana Rossberg in Fedde, Repert. Spec. Nov. 42: 61 (1987). Helicostylis tomentosa (Poepp. & Endl.) Rusby in Mem. Torr. Bot. Club, 6: 1896). . Morus alba L. Sp. Pl. 986 (1753). Ogcodeia acreana Mildbr. in Notizbl. 11: 421 (19382). O. pseudo-naga Mildbr. in Notizbl. 11: 419 (1932). O. Sandwithiana Mildbr. in Notizbl. 11: 422 (1932). O. ternstroemiiflora Mildbr. in Notizbl. 10: 188 (1927). Olmedia aspera R. & P. Syst. 257 (1798). O. Habas Pax in Fedde, Repert. Spec. Nov. 7: 108 (1909). Pharmacosycea Brittonii Rusby in Bull. Torr. Bot. Club, 28: 309 (1901). Poulsenia armata (Miq.) Standl. in Trop. Woods, 33: 4 (1933). 67 Pourouma crassivenosa Mildbr. in Notizbl. 10: 419 (1928). P. scabra Rusby in Bull. N. Y. Bot. Gard. 6: 498 (1910). P. subtriloba Rusby in Mem. N. Y. Bot. Gard. 7: 232 (1927). P. uvifera Rusby in Mem. N. Y. Bot. Gard. 7: 231 (1927). Pseudolmedia alnifolia Rusby in Mem. N. Y. Bot. Gard. 7: 229 (1927). P. hirtellaefolia Rusby in Mem. N. Y. Bot. Gard. 7: 228 (1927) P. laevigata Trécul in Ann. Sci Nat. (ser. 3) 8: 131 (1847). P. laevis (R. & P.) Macbr. in Field Mus. Publ. Bot. 11: 16 (1981). P. Mildbraedii Macbr. in Field Mus. Publ. Bot. 11: 61 (1931). Sorocea ilicifolia Mig. in Mart. Fl. Bras. 4(1): 114 (1853). S. muriculata Miq. in Mart. Fl. Bras 4(1): 118 (1858). Trophis aurantiaca Herzog in Fedde, Repert. Spec. Nov. 7: 51 (1909). Urostigma costata Rusby in Bull. N. Y. Bot. Gard. 6: 499 (1910) = Ficus uiana URTICACEAE Boehmeria brevirostris Wedd. in Ann. Sci. Nat. (ser. 4) 1: 201 (1854). caudata Sw. Prodr. 34 (1788). fallax Wedd. in Arch. Mus. Paris, 9: 346 (1856-57). pallida (Rusby) Killip in Journ. Wash. Acad. Sci. ae 847 (1931). Pavonii Wedd. in Ann. Sci. Nat. (ser. or 1: 202 (18 sordida Rusby, Descr. S. Am. Pls. 9 (1920). tenuistachys Rusby in Bull. Torr. Bot. Club, 28: 312 (1901). Goethartia edentata (O. Ktze.) Herzog in Meded. Rijks Herbar. 27: 77 (1915). Myriocarpa densiflora Benth. Bot. Voy. Sulphur, 169 (1846) = M. stipitata. M. filiformis Rusby in Bull. N. Y. Bot. Gard. 6: 500 (1910). M. obscura Rusby, Descr. S. Am. Pls. 11 (1920). M. purpurascens Rusby in Mem. N. Y. Bot. Gard. 7: 233 (1927). M. stipitata Benth. Bot. Voy. Sulphur, 168, t. 55 (1846). M. Tatei Rusby in Phytologia, 1: 52 (1984). Parietaria debilis Forst. f. Fl. Ins. Austral. Prodr. 73 (1786). Phenax angustifolius (HBK.) Wedd. in ey Sci. Nat. (ser. 4) 1: 193 (1854). P. bullatus Rusby in Phytologia, 1: 52 (193 P. flavifolius Rusby in Phytologia, 1: 53 ee P. globuliferus Rusby in Mem. Torr. Bot. Club. 6: 120 (1896). P. laevigatus Wedd. in Ann. Sci. Nat. (ser. 4) 1: 192 (1854). P. laevigatus var. serratus Wedd. in DC. Prodr. 16(1): 23549 (1869). P. pallidus Rusby in Mem. Torr. Bot. Club, 4(3): 259 (1895) = Boehmeria Pavonii. P. pauciserratus (Wedd.) Rusby in Bull. Torr. Bot. Club, 28: 313 (1901). P. rugosus (Poir.) Wedd. in DC. Prodr. 16(1): 28538 (1869). P. rugosus var. Mandonii Wedd. in DC. Prodr. 16(1): 23535 (1869). P. Sonnerattii Wedd. in DC. Prodr. 16(1): 23537 (1869). Pilea anomala Wedd. in Ann. Sci. Nat. (ser. 3) 18: 217 (1852) = P. multiflora. P. Buchtienii Killip in Journ. Wash. Acad. Sci. 15: 297 (1925). capitellata Wedd. in Arch. Mus. Paris. 9: 220 (1856-57). cymbifolia Rusby in Bull. Torr. Bot. Club, 28: 311 (1901 dauciodora (R. & P.) Wedd. in Ann. Sci. Nat. (ser.3) 18: 238 (1852). filipes Rusby in Bull. Torr. Bot. Club, 28: 311 (1901). hyalina Fenzl] in Denkschr. Akad. Wiss. Math. Naturw. Wien, 1: 256 (1850). macrophylla Rusby, Descr. S. Am. Pls. 10 (1920) = P. picta. multiflora (Poir.) Wedd. in Ann. Sci. Nat. (ser. 3) 18: 217 (1852). nutans (Poepp.) Wedd. in Arch. Mus. Paris, 9: 196, t. 7 (1856-57). pauciserrata Killip in Journ. Wash. Acad. Sci. 15: 293 (1925). picta Herzog in Meded. Rijks Herbar. 27: 76 (1915). rotundata Griseb. Fl. Brit. W. Ind. 158 (1859). PW Www Pee we oY 20 V0 oe Rusbyi (Britton) Killip in Journ. Wash. Acad. Sci. 15: 293 (1925). serpyllacea (HBK.) Wedd. in Ann. Sci. Nat. (ser. 3) 18: 207 (1852). Spruceana Wedd. in DC. Prodr. 16(1): 161 (1869). strigosa Wedd. in Ann. Sci. Nat. (ser. 3) 18: 225 (1852). sublobata Rusby in Bull. Torr. Bot. Club, 28: 311 (1901). uncidens Wedd. in Ann. Sci. Nat. (ser. 3) 18: 224 (1852) — P. dauciodora. urerifolia Rusby in Bull. Torr. Bot. Club, 28: 312 (1901) = P. cymbifolia. Pouzolzia asper (Wedd.) Wedd. in DC. Prodr. 16(1): 233 (1869) — P. Poep- et oe ee ee pigiana. P. discolor Wedd. in Arch. Mus. Paris, 9: 408 (1856-57) — P. Poeppigiana. P. Poeppigiana (Wedd.) Killip in Journ. Wash. Acad. Sci. 21: 347 (1931). Urera alceaefolia Gaud. in Freyc. Voy. Bot. 407 (1830). U. baccifera (L.) Gaud. in Freye. Voy. Bot. 497 (1830), combination implied but not actually made. boliviensis Herzog in Meded. Rijks Herbar. 27: 75 (1915). capitata Wedd. in Ann. Sci. Nat. (ser. 3) 18: 201 (1852). ecaracasana (Jacq.) Griseb. Fl. Brit. W. Ind. 154 (1859). filiformis Rusby in Bull. Torr. Bot. Club, 28: 310 (1901). laciniata Wedd. in Ann. Sci. Nat. (ser. 3) 18: 203 (1952). rugosa Rusby in Bull. Torr. Bot. Club, 28: 310 (1901). sinuata Wedd. in Ann. Sci. Nat. (ser. 3) 18: 201 (1852). viridisetosa Rusby in Mem. N. Y. Bot. Gard. 7: 232 (1927). rtica ballotaefolia Wedd. in Ann. Sci. Nat. (ser. 3) 18: 197 (1852). echinata Benth. Pl. Hartw. 252 (1846). echinata var. trichantha Wedd. in DC. Prodr. 16(1): 42 (1869). flabellata HBK. Nov. Gen. & Spec. 2: 40 (1817). Seana Juss. ex Poir. in Lam. Encyc. Suppl. 4: 223 (1816). purp cens Nutt. in Trans. Amer. Phil. Soc. 5: 169 (1837). subincisa Benth. Pl. Hartw. 292 (1848). Trianae Rusby in Bull. N. Y. Bot. Gard. 4: 319 (1907). ORG BE “BE Ge Gig PROTEACEAE Orites Fiebrigii (Perkins) Sleumer in Engler, Bot. Jahrb. 76: 141 (1954). Pamopsis Pearcei Rusby in Bull. N. Y. Bot. Gard. 4: 439 (1907) = P. rube- scens var. Sprucei. P. rubescens var. Sprucei (Meisn. ex Rusby) Sleumer in Engler, Bot. Jahrb. 76: 180 (1954). Roupala Meisneri Sleumer in Engler, Bot. Jahrb. 76: 162 (1954). R. montana Aubl. Pl. Guian. Fr. 1: 83, t. 32 (1775). R. montana var. dentata (R. Br.) Sleumer in Engler, Bot. Jahrb. 76: 173 (1954). R. Steinbachii Sleumer in Engler, Bot. Jahrb. 76: 157 (1954). LORANTHACEAE Antidaphne viscoidea Poepp. & Endl. Nov. Gen. 2: 70, t. 199 (1838). Dendrophthora buxifolia (Lam). Eichl. in Mart. Fl. Bras. 5(2): 105 (1868). clavata (Benth.) Urb. in Ber. Deut. Bot. Ges. 14: 285 (1896). inaequidenatta (Rusby) Trel. Clinas Phorad. 218 (1916). mesembryanthemifolia (Griseb.) Urb. in Ber. Deut. Bot. Ges. 14: 285 (1896). Rusbyi (Britton) Trel. Genus siege 218 (1916). striata Rusby, Descr. S. Am. Pls. 920). subtrinervis Urb. in Ber. Deut. “ Ges, 14: 285 (1896). subtrinervis (Rusby) Trel. Genus Phorad. 218 (1916). PS9S SSD 69 Loranthus concinnus Mart. in Schult. Syst. Veg. 7: 150 (1829) = Phrygilan- thus sp. ? L. cubeboides Rusby in Bull. Torr. Bot. Club, 27: 135 (1900) = Phrygilan- thus sp. ? L. flexilis Rusby in Mem. Torr. Bot. Club, 4(3): 253 (1895) = Phrygilan- thus sp. ? Oryctanthus botryostachys Eichl. in Mart. Fl. Bras. 5(2): 89, t. 29 (1868) Phoradendron angustifolium (HBK.) Eichl. in Mart. Fl. Bras. 5(2): 115 (1868). argentinum Urb. in Engler, Bot. Jahrb. 23, Beibl. 57: 14 (1897). Balleanum (Seem.) Eichl. in Mart. Fl. Bras. 5(2): 184m (1868). bathyoryctorum Eichl. in Mart. Fl. Bras. 5(2): 123, t. 43 (1868). bolivianum Trel. Genus Phorad. 138 (1916). Brittonianum Rusby in Mem. Torr. Bot. Club, 4(3): 254 (1895). clavatum (Benth.) Eich]. in Mart. Fl. Bras. 5(2): 107 (1868). crassifolium (Pohl) Eichl. in Mart. Fl. Bras. 5(2): 125, t. 40 (1868). emarginatum Eichl. in Mart. Fl. Bras. 5(2): 118, t. 38 (1868). Kuntzei Urb. in Engler, Bot. Jahrb. 23, Beibl. 58: 11 (1897). latifolium (Sw.) Griseb. in Mem. Amer. Acad. (ser. 2) 8: 191 (1861). Liga (Gill.) Eichl. in Mart. Fl. Bras. 5(2): 184m (1868). Mandonii Eichl. in Mart. Fl. Bras. 5(2): 124 (1868). meliae Trel. Genus Phorad. 121 (1916). paucifolium Rusby in Bull. N. Y. Bot. Gard. 6: 501 (1910). Pearcei Rusby in Bull. Torr. Bot. Club, 27: 186 (1900). pennivenium (DC.) Eichl. in Mart. Fl. Bras. 5(2): 128 (1868). piperoides (HBK.) Trel. Genus Phorad. 145 (1916). Rusbyanum Trel. in Mem. N. Y. Bot. Gard. 7: 234 (1927). semiteres Trel. Genus Phorad. 69 (1916). tafallaeoides Rusby in Bull. N. Y. Bot. Gard. 4: 441 (1907). tucumanense Urb. in Engler, Bot. Jahrb. 23, Beibl. 57: 16 (1897). undulatum (Pohl) Eichl. in Mart. Fl. Bras. 5(2): 122, t. 39 (1868). rygilanthus bolivianus Engl. in Engl. & Prantl, Nat. Pflanzenfam. Nachtr. 183 (1897). cordifolius Rusby in Mem. N. Y. Bot. Gard. 7: 233 (1927). cuneifolius (R. & P.) Eichl. in Mart. Fl. Bras. 5(2): 49, t. 11 (1868). eugenioides (HBK.) Eichl. in Mart. Fl. Bras. 5(2): 50, t. 12 (1868). falcatus Rusby in Mem. N. Y. Bot. Gard. 7: 233 (1927). faleatus var. (?) macrocalyx Rusby in Mem. N. Y. Bot. Gard. 7: 234 (1927). mapirensis (Rusby) Engl. in Engl. & Prantl, Nat. Pflanzenfam. Nachtr. 184 (1897). punctatus (R. & P.) Hichl. in Mart. Fl. Bras. 5(2): 47 (1868). tetrandrus (R. & P.) Eichl. in Mart. Fl. Bras. 5(2): 47 (1868). P. verticillatus (R. & P.) Eichl. in Mart. Fl. Bras 5(2): 47 (1868). Phthirusa heterophylla Rusby in Bull. N. Y. Bot. Gard. 6: 500 (1910). Oe A Se NS UUM Se P. robusta Rusby in Bull. N. Y. Bot. Gard. 6: 501 (1910). Psittacanthus cupulifer (HBK.) G. Don, Gen. Hist. 3: 417 (1834). Struthanthus concinnus Mart. in Flora, 13( 1): 104 (1830). S. divaricatus Rusby, Descr. S. Am. Pls. 12 (1920). S. nudipes Rusby in Mem. Torr. Bot. Club, 6: 116 (1896). S. oblongifolius Rusby in Bull. N. Y. Bot. Gard. 4: 440 (1907). SANTALACEAE Acanthosyris falcata Griseb. in Goett. Abh. 24: 151 (1879). lodina rhombifolia Hook. & Arn. in Hook. Bot. Misc. 3: 172 (1833). 70 Quinchamalium chilense Lam. Illustr. 2: 125, t. 142 (1793). Q. gracile Brongn. in Duperr. Voy. Coq. Bot. 231, t. 52 (1829). Q. majus Brongn. in Duperr. Voy. Coq. Bot. 229, t. 51 (1829). Q. Stuebelii Hieron. in Engler, Bot. Jahrb. 21: 306 (1896). OPILIACEAE Agonandra brasiliensis Miers in Ann. & Mag. Nat. Hist. (ser. 2) 8: 172 (1851). OLACACEAE Heisteria biflora Rusby in Mem. Torr. Bot. Club, 6: 18 (1896) = H. brasiliensis. H. brasiliensis Engl. in Mart. Fl. Bras. 12(2): 19 (1872). ? H. caloneura Sleumer in Notizbl. 12: 66 (1934). H. ixiamensis Rusby in Mem. N. Y. Bot. Gard. 7: 234 (1927). H. scandens Ducke in Archiv. Jard. Bot. Rio Janeiro, 4: 9 (1925). H. yapacaniensis (O. Ktze.) K. Schum. in Just, Bot. Jahresber. 26(1): 360 (1900). Schoepfia obliquifolia Turcz. in Bull. Soc. Imp. Nat. Mosc. 31(1): 249 (1858). S. tetramera Herzog in Meded. Rijks Herbar. 29: 4 (1916) Ximenia americana L. Sp. Pl. 1197 (1753). BALANOPHORACEAE ? Helosis sp. Juelia subterranea Asplund in Sv. Bot. Tidskr. 22: 274 (1928). Lophophytum bolivianum Wedd. in Ann. Sci. Nat. (ser. 3) 14: 185, t. 10 (1850). Ombrophytum zamioides Wedd. in Ann. Sci. Nat. (ser. 3) 14: 184, t. 10 (1850). ARISTOLOCHIACEAE Aristolochia angustifolia Cham. in Linnaea, 7: 211, t. 5 (1832). A. apoloénsis Rusby in Bull. N. Y. Bot. Gard. 6: 502 (1910). A. asperifolia Ule in Verh. Bot. Ver. Brandenb. 47: 121 (1905). A. boliviensis O. Ktze. Rev. Gen. 3(2): 271 (1898). A. Buchtienii O. C. Schmidt in Fedde, Repert. Spec. ay. 27: 292 (1930). A. Burelae Herzog in Meded. Rijks Herbar. 40: 2 (19 A. Cardenasii Standl. in Field Mus. Publ. Bot. 17: aay tly A. ceresensis O. Ktze. Rev. Gen. 3(2): 272 (1898). A. chiquitensis Duchtr. in Ann. Sci. Nat. (ser. 4) 2: 49 (1854). A. cymbifera var. abbreviata Duchtr. in DC. Prodr. 15( 5 470 (1864). A. didyma Sp. Moore in Journ. Bot. 58: 7, t. 535, fig. 1 (1915). A. eriantha Mart. & Zucc. Nov. Gen. 1: 78, t. 53 (1824). A. esperanzae O. Ktze. Rev. Gen. 3(2): 272 (1898). A. Gibertii Hook. in Bot. Mag. 88: t. 5345 A priak A. Guentheri O. C. Schmidt in Fedde, Repert. Spec. Nov. 27: 292 (1980). A. Hoehneana O. C. Schmidt in Notizbl. 12: 390 (1935 A. Joergensenii Haum, in Anal. Mus. Nac. Buenos Aires: 82: 328, t. 1 (1923). A. Lindneri Berger in Notizbl. 10: 129 (1927). A. lingulata Ule in Verh. Bot. Ver. Brandenb. 47: 123 (1905). A. lingulata var. parviflora Herzog in Meded. Rijks Herbar. 40: 3 (1921). A. macroura Gomez in Act. Olyss. 77 (1812 A. odoratissima L. Sp, Pl. (ed. 2) 2: 1862 (1763). A. pandurata Jacq. Hort. Schoenbr. 4: 49, t. 497 (1804). ? A. peruviana O. C. Schmidt in Notizbl. 9: 186 (1924). A. pilosa HBK. Nov. Gen. & Spec. 2: 146, t. 113 (1817). A. prostrata Duchtr, in Ann. Sci. Nat. (ser. 4) 2: 68 (1854). 71 pseudo-triangularis O. C. Schmidt in Fedde, Repert. Spec. Nov. 38: 110 1935). Rimbachii O. C. Schmidt in Fedde, Repert. Spec. Nov. 23: 287 (1927). Rodriguesii Hoehne in Mem. Inst. Oswaldo Cruz, 20(1): 140 (1927). sessilifolia (Klotzsch) Duchtr. in DC. Prodr. 15(1): 464 (1864). stenocarpa O. Ktze. Rev. Gen. 3(2): 271 (1898) triangularis Cham. in Linnaea, 7: 209, t. 6 (1832) Trollii O. C. Schmidt in Fedde, Repert. Spec. gr a0: oa ee Weddellii Duchtr. in Ann. Sci. Nat. (ser. 4) 2: 62 (18 Werdermanniana O. C. Schmidt in Fedde, nee 9 — 30: 67 (1932). Williamsii Rusby in Bull. N. Y. Bot. Gard. 6: 501 (1910). yungasensis Rusby in Bull. N. Y. Bot. Gard. 4: 437 (1907). er PP Pe Pi aoe RAFFLESIACEAE Apodanthes tribracteata Rusby, Descr. S. Am. Pls. 15 (1920). ? Pilostyles sp POLYGONACEAE Coccoloba chacoénsis Standl. in Field Mus. Publ. Bot. 17: web (1937). illheénsis Wedd. in Ann. Sci. Nat. (ser. 3) 13: 258 (185 Meissneriana (Britton) K. Schum. in Just, Bot. See 28(1): 451 (1902). padifolia Rusby in Mem. N. Y. Bot. Gard. 7: 235 (1927). Persicaria Wedd. in Ann. Sci. Nat. (ser. 3) 13: 256 (1850). polystachya Wedd. in Ann. Sci. Nat. (ser. 3) 13: 261 (1850). strobilulifera Meissn. in Mart. Fl. Bras. 5(1): 25 (1855). tiliacea Lindau in Engler, Bot. Jahrb. 13: 198, t. 5 (1890). annoens M. fruticulosa (Walp.) Standl. in Field Mus. Publ. Bot. 13: 453 (1937). M. hastulata (Smith) I. M. Johnst. in Contrib. Gray Herb. 81: 88 (1928). M. rupestris Wedd. in Ann. Sci. Nat. (ser. 3) 13: 256 (1850) = M. aoaanaiap M. rupestris var. nivalis Wedd. in Ann. Sci. Nat. (ser. 3) 13: 256 (1850) = M. tamnifolia (HBK.) Meissn. Pl. Vase. Gen. Comm. 2: 227 (1843) M. tiliaefolia Wedd. in Ann. Sci. Nat. (ser. 3) 13: 255 (1850). M. vuleanica (Benth.) Endl. Gen, Suppl. 4, pt. 2: 51 (1847). Polygonum acuminatum HBK. Nov. Gen. & Spec. 2: 178 (1817). P. fallax Small in Bull. Torr. Bot. Club, 24: 46 (1897) P. lacerum HBK. Nov. Gen. & Spec. 2: 179 (1817). P. persicarioides HBK. Nov. Gen. & Spec. 2: 179 (1817). Rumex conglomeratus Murr. Prodr. Fl. Gotting. 52 (1770). R. crispus L. Sp. Pl. 335 (1753). R. cuneifolius Campd. Monog. Rum. 66, 95 (1819). Ruprechtia apetala Wedd. in Ann. Sci. Nat. (ser. 3) 13: 268 (1850). R. boliviensis Herzog in Meded. Rijks Herbar. 46: 5 (1922). R. mollis Wedd. in Ann. Sci. Nat. (ser. 3) 18: 268 (1850). R. scandens Rusby in Mem. N. Y. Bot. Gard. 7: 237 (1927). R. triflora Griseb. in Goett. Abh. 24: 89 (1879 ~reere boliviana Britton ex Rusby in Bull. Torr: Bot. Club, 27: 130 (1900) . Pavonii. . Caracasana Cham. in Linnaea, 8: 137 (1833). corylifolia var. intermedia O. Ktze. Rev. Gen. 3(2): 271 (1898). efistulifera Rusby in Bull. Torr. Bot. Club, 27: 129 (1900). estriata O. Ktze. Rev. Gen. 3(2): 271 (1898). “ce fag Ee 72 ke 7. tT. We ce die t. Ts guanaiensis Rusby in Mem. Torr. Bot. Club, 6: 111 (1896). hispida Britton ex Rusby in Mem. Torr. Bot. Club, 6: 111 (1896) — T. Poep- pigiana. Pavonii Meissn. in DC. Prodr. 14: 172 (1856). peruviana Fisch. & Mey. ex C. A. Mey. in Mém. Acad. St. Pétersb. (ser. 6) Sci. Nat. 4: 149 (1845). Poeppigiana Wedd. in Ann. Sci. Nat. (ser. 3) 13: 265 (1850). setosa Rusby in Mem. N. Y. Bot. Gard. 7: 287 (1927). vestita Rusby in Mem. N. Y. Bot. Gard. 7: 236 (1927). Williamsii Rusby in Mem. N. Y. Bot. Gard. 7: 235 (1927). CHENOPODIACEAE Atriplex Asplundii Standl. in Field Mus. Publ. Bot. 11: 125 (1931). A. A. A. A. A. A. a cristata Humb. & Bonpl. ex Willd. Sp. Pl. 4(2): 959 (1806). cristata var. depauperata O. Ktze. Rev. Gen. 3(2): 266 (1898) — A. Her- zogii. cristata var. pulvinata O. Ktze. Tev. Gen. 3(2): 266 (1898) — A. Herzogii. Herzogii Standl. in Field Mus. Publ. Bot. 11: 125 (1931). imbricata (Mogq.) D. Dietr. Syn. Pl. 5: 536 rte Lampa Gill. ex Mog. in DC. Prodr. 13(2): 110 (1849). Rusbyi Britton in Rusby in Mem. Torr. Bot. Club, 4(3): 250 (1895). A. semibaccata R. Br. Prodr. 406 (1810). A. serpyllifolia Herzog in Meded. Rijks Herbar. 27: 11 (1915) = A. Herzogii. Chenopodium ambrosioides L. Sp. Pl. 219 (1753). Q SHES AOE Sp © . So nanan & 4 ambrosioides subsp. chilense (Schrad.) Aellen in Fedde, Repert. Spec. Nov. 26: 36 (1929). ambrosioides subsp. page var. andicola (Phil.) Aellen in Fedde, Repert. Spec. Nov. 26: 37 (1929). . ambrosioides forma minus (Murr.) Aellen in Fedde, Repert. Spec. Nov. 26: 37 (1929) ambrosioides var. querciforme (Murr.) Aellen in Fedde, Repert. Spec. Nov. 26: 37 (1929). bolivianum Murr. in Magyar Bot. rms 1: 859 (1902) = C. petiolare. carnosulum Mog. in DC. Prodr. 13(2): 64 (1849 graveolens var. Bangii (Murr.) Aellen in Verh. Natur. Ges. Basel, 41: 107 (1931). hircinum Schrad. Ind. Sem. Hort. Goett. 1832: 2 (1838). hircinum var. rhombicum Aellen in Fedde, neler Spec. Nov. 26: 122 (1929). incisum Poir. in Lam. Encyc. Suppl. 1: 392 (1810). macrospermum var. halophilum (Phil.) Standl. in Field Mus. Publ. Bot. 11: 118 (1931). macrospermum subsp. halophilum forma angustius Thell. & Aellen in Fedde, Repert. Spec. Nov. 26: 44 (1929). macrospermum forma farinosum (S, Wats.) Aellen in Fedde, Repert. Spec. Nov. 26: 43 (1929). macrospermum forma nanum Aellen in — Repert. Spec. Nov. 26: 44 (1929) = C. macrospermum forma angust murale L. Sp. Pl. 219 (1753). pallidicaule Aellen in Fedde, Repert. Spec. Nov. 26: 126 (1929). paniculatum Hook. ex Mog. in DC. Prodr. 13(2): 65 (1849). paniculatum var. incanum (Wats.) Murr. in Allg. Bot. Zeitschr. 12: 54 (1906). petiolare HBK. Nov. Gen. & Spec. 2: 191 (1817). petiolare forma hastatum (Phil.) Aellen in Fedde, Repert. Spec. Nov. = 150 (1929) 73 petiolare forma incanum (Murr.) Aellen in Fedde, Repert. Spec. Nov. 26: 150 (1929). petiolare forma scutatum Aellen in Fedde, Repert. Spec. Nov. 26: 151 9). petiolare forma trilobum Aellen i Fedde,Repert. Spec. Nov. 26: 151 (1929). Quinoa Willd. Sp. Pl. 1: 1301 (1798). wae var. melanospermum Hunziker in Revist. Argent. Agron. 10: 317 (194 Quins Portas purpureum ) > Psa Lingelsh. in Fedde, Repert. Spec. Nov. 7: 241 (1909) = C. graveol- ens var. Bangii. Salicornia andina Phil. in Anal. Mus. Nac. Chile, Bot. 8: 75 (1891). S. fruticosa L. Sp. Pl. (ed. 2) 1: 5 (1762). Suaeda foliosa Mog. in DC. Prodr. 13(2): 156 (1849). S. fruticosa var. crassifolia Moq. in DC. Prodr. 13 (2): 157 (1849) = = S. foliosa. Teloxys Mandonii S. Wats. in Proc. Amer. Acad. 9: 91 (1874) —= Chenopodium graveolens var. Bangii. AMARANTHACEAE Achyranthes aspera L. Sp. Pl. 204 (1753). A. aspera var. indica L. Sp. Pl. 204 (17538). A. Bangii Standl. in Journ. Wash. Acad. Sci. 5: 74 (1915) = Alternanthera aniculata. Alternanthera altacruzensis Suesseng. in Mitteil. Bot. Staatss. Miinch. 1: 3 1950). boliviana (Rusby) Standl. in Journ. Wash. Acad. Sci. 5: 74 (1915). coriacea Herzog in Meded. Rijks Herbar. 46: 6 (1922). lanceolata (Benth.) Schinz in Engl. & Prantl, Nat. Pflanzenfam. (ed. 2) 16C: 75 (1934) mexicana forma lanuginosa Suesseng. in Fedde, Repert. Spec. Nov. 35: 301 (1934). microcephala (Moq.) Schinz in Engl. & Prantl, Nat. Pflanzenfam. (ed. 2) 16C: 75 (1934). microphylla R. E. Fries in Nov. Act. Soc. Sci. Upsal. (ser. 4) 1(1): 154 (1905). Moquinii (Webb) Dusén in Arch. Mus. Nac. Rio Janeiro, 13: 63 (1903). paniculata HBK. Nov. Gen. & Spec. 2: 208 (1818). paronychioides St. Hil. Voy. Brés. 2(2): 439 (1833). philoxeroides (Moq.) Griseb. in Goett. Abh. 24: 36 (1879). pilosa Moq. in DC. Prodr. 13(2): 357 (1849). polygonoides (L.) R. Br. Prodr. 417 (1810), combination implied but not actually made. hiss (Benth.) O. Ktze. Rev. Gen. 2: 538 (1891). pungens HBK. Nov. Gen. & Spec. 2: 206 (1818). repens (LD O. Ktze. Rev. Gen. 2: 536, 540 (1891). scandens Herzog in Meded. Rijks Herbar. 46: 7 (1922). tomentosa (Mogq.) Schinz in Engl. & Prantl, Nat. Pflanzenfam. (ed. 2) 16C: 76 (1934). maranthus affinis Thell. in Fedde, Repert. Spec. Nov. 21: 324 (1925). Asplundii Thell. in Fedde, Repert. Spec. Nov. 21: 322 (1925). Buchtienianus Thell. in Fedde, Repert. Spec. Nov. 21: 323 (1925). Cardenasianus Hunziker in Bol. Soc. Argent. Bot. 4: 186 (1952). caudatus L. Sp. Pl. 990 (1753). chlorostachys Willd. Amaranth. 34, t. 10, fig. 19 (1790). ee gl es ee a Perr rs PP PPP 74 A. muricatus Gill. ex Moq. in DC. Prodr. 13(2): 276 (1849). A. peruvianus (Schauer) Standl. in Field Mus. Publ. Bot. 13(2): 487 (1987). A. retroflexus L. Sp. Pl. 991 (1753). Chamissoa altissima (Sw.) HBK. Nov. Gen & Spec. 2: 197, t. 125 (1817). C. altissima subsp. albo-grisea Suesseng. in Fedde, Repert. Spec. Nov. 39: 6 7 C. altissima var. rubella Suesseng. in Fedde, Repert. Spec. Nov. 35: 306 (1934). C. celosioides Griseb. in Goett. Abh. 19: 79 (1874). Cyathula achyranthoides (HBK.) Mog. in DC. Prodr. 18(2): 326 (1849). — acaulis Remy in Ann. Sci. Nat. (ser. 3) 6: 350 (1846) = G. Meyeni- Reitaee Mog. in DC. Prodr. 13(2): 401 (1849). cinnabarina Suesseng. in Fedde, Repert. Spec. Nov. 35: 309 (1984). Conwayi Rusby in Bull. N. Y. Bot. Gard. 8: 89 (1912) — G. Meyeniana. decumbens Jacq. Hort. Schoenbr. 5: 41, t. 482 (1804). decumbens var. boliviana Stuchlik in Fedde, Repert. Spec. Nov. 12: 519 discolor R. E. Fries in Ark. Bot. 16(12): 34 (1920). elegans var. mandonioides Suesseng. in Fedde, Repert. Spec. Nov. 35: 310 (1934). Gardneri Mog. in DC. Prodr. 13(2): 404 (1849). glabrata (Mart.) Mog. in DC. Prodr. 13(2): 388 (1849). glutinosa R. E. Fries in Ark. Bot. 16(12): 28 (1920). hygrophila forma luteiflora Herzog in Meded. Rijks Herbar. 46: 8 (1922). hygrophila forma subecristata Herzog in Meded. Rijks Herbar. 46: 8 (1922). ARAMA AL prees ixtamensis Rusby in Bull. N. Y. Bot. Gard. 6: 502 (1910) — G. decumbens. lutea Rusby in Mem. N. Y. Bot. Gard, 7: 238 (1927). Mandonii R. E. Fries in Ark. Bot. 16(12): 22 (1920). Meyeniana Walp. in Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 404 (1843). oligocephala Remy in Ann. Sci. Nat. (ser. 3) 6: 350 (1846). oligocephala var. pallida Suesseng. in Fedde, Repert. Spec. Nov. 49: 9 ). perennis L. Sp. Pl. 224 (1753). perennis var. boliviana Stuchlik in Fedde, Repert. Spec. Nov. 12: 521 (1918). perennis var. saxatilis R. E. Fries in Ark. Bot. 16(12): 40 (1920). potosiana Suesseng. & Benl in Mitteil. Bot. Staatss. Miinch. 1: 6 (1950 pulchella forma nana Suesseng. in Fedde, Repert. Spec. Nov. 35: 315 (1934). pulchella var. rosea Stuchlik in Fedde, Repert. Spec. Nov. 12: 523 (1913). subalpina Herzog in Meded. Rijks Herbar. 46: 7 (1922) tarijensis R. E. Fries in Ark. Bot. 16(12): 27 (1920). Trollii Suesseng. in Fedde, Repert. Spec. Nov. 35: 315 (1934). umbellata Remy in Ann. Sci. Nat. (ser. 3) 6: 349 (1846). vaga Mart. Nov. Gen. 2: 17, t. 120 (1826). Guilleminia densa (Willd.) Mog. in DC. Prodr. 13(2): 338 (1849). Hebanthe holosericea Mart. in Flora, 21, II Beibl.: 65 (1838) = Pfaffia sp. ? Iresine angustifolia Euphr. Beskr. St. Barthel. 165 (1795). . Cardenasii Standl. in Field Mus, Publ. Bot. 17: 241 (1937). Celosia L. Syst. (ed. 10) 1291 (1759). celostoides L. Sp. Pl. (ed. 2) 2: 1456 (1763) = I. Celosia. celosioides var. nicotianoides Suesseng. in Fedde, Repert. Spec. Nov. 39: 11 ANAANAHAARAH ARRAAD et es et et Hassleriana Chod. in Bull. Herb. Boiss. (ser. 2) 3: 390 (1903). . Hassleriana var. guaranoides Suesseng. in Fedde, Repert. Spec. Nov. 35: 321 — . (1934). . paniculata (L.) O. Ktze. Rev. Gen. 2: 542 (1891). — 75 I. spiculigera forma pauciglandulosa Herzog in Meded. Rijks Herbar. 46: 9 (1922). Mogiphanes soratensis Rusby in Bull. N. Y. Bot. Gard. 6: 503 (1910) = Al- ternanthera lanceolata Pfaffia Bangii R. E. Fries in Ark. Bot. 16(12): 11 (1920). brachiata Chod. in Chod. & Rehf. in Bull. Soc. Bot. Genéve, 14: 285 (1927). brachiata var. grandiflora (Fries) Stiitzer in Fedde, Repert. Spec. Nov. Beih. 88: 8 (1935). Dunaliana (Mog.) Schinz in Engl. & Prantl, Nat. Pflanzenfam. (ed. 2) 16C: 68 (1934). Fiebrigii Suesseng. in Fedde, Repert.. Spec. Nov. 35: 329 (1934). fruticulosa Suesseng. in Fedde, Repert. Spec. Nov. 35: 330 (1934). glabrata Mart. Nov. Gen. 2: 21, t. 122 (1826). gnaphalioides (L. f.) Mart. Nov. Gen. 2: 24 (1826). gnaphalioides pet floccosa (Seub.) Stiitzer in Fedde, Repert. Spec. Nov. Beih. 88: 26 (1935). grandiflora var. prominulifera Stiitzer in Fedde, Repert. — Nov. Beih. 88: 10 (1935). holosericea (Moq.) Standl. in Field Mus. Publ. Bot. 18(2): 491 (19387). arr eid forma gracilis Stiitzer in Fedde, Repert. Spec. Nov. Beih. 88: 34 (193 soiree (Mart.) O. Ktze. Rev. Gen. 2: 543 (1891). soratensis Rusby in Bull. N .Y. Bot. Gard. 6: 502 (1910) = Gomphrena, aff. G. elegans var. mandonioides. stenophylla (Spreng.) Stuchlik in Fedde, Repert. Spec. Nov. ch io (1913). Pleuropetalum Sprucei (Hook. f.) Standl. in N. Am. Fl. 21: 96 (19 Telanthera Bangii Rusby in Mem. Torr. Bot. Club, 6: 110 ‘aan <= i Gudnan thera paniculata. WU WM Wo Ww Wh ry NYCTAGINACEAE Allionia incarnata L. Syst. ig at 2: 890 (1759). Boerhaavia caribaea Jacq. Obs. Bot. 4: . t. 84 (1771). (1903). spectabilis Willd. Sp. Pl. 2: 348 (1799). spinosa (Cav.) Heimerl in Engl. & Prantl, Nat. Pflanzenfam. 3 (lb): 27 (1889 B. coccinea Mill. Gard. Dict. (ed. a no. 4 (1768). B. coccinea var. leiocarpa (Heimer!) Standl. in Field Mus. Publ. Bot. 11: 108 (1931). B. erecta L. Sp. Pl. 3 (1753). B. Friesii Heimerl in Oesterr. Bot. Zeitschr. 56: 253 (1906) = B. coccinea var. leiocarpa. B. scandens L, Sp. Pl. 3 (1753). B. viscosa Lag. & Rod. in Anal. Cienc. Nat. 4: 256 (1801). Bougainvillea berberidifolia Heimer! in Denkschr. Akad. Wiss. Wien, 70: 121, t. 1 (1900). B. campanulata Heimerl in Meded. Rijks Herbar. 19: 33 (1913). B. Herzogiana Heimerl in Meded. Rijks Herbar. 27: 12 ( 1915). B. infesta Griseb. in Goett. Abh. 24: 40 (1879). B. longispinosa Rusby in Mem. Torr. Bot. Club, 6: 109 (1896). B. modesta Heimerl in Denkschr. Akad. Wiss. Wien, 70: 118 (1900). B. praecox Griseb. in Goett. Abh. 24: 40 (1879). B. praecox var. rhombifolia Heimerl in Verh. Zool. -Bot. Ges. Wien, 62: 4 (1912). B. praecox var. spinosa Chod. & Hassl. in Bull. Herb. Boiss. (ser. 2) 3: 415 B. B. B. stipitata Griseb. in Goett. Abh. 24: 39 (1879), nomen nudum B. stipitata var. Fiebrigii Heimerl in Engler, Bot. Jahrb. 42: 76 (1908). B. stipitata var. Grisebachiana Heimerl in Denkschr. Akad. Wiss. Wien, 70: 116 (1900). B. stipitata var. Kuntzeana Heimerl in Denksch. Akad. Wiss. Wien, 70: 117 (1900). B. Trollii Heimerl in Notizbl. 11: 464 (19382). Colignonia glomerata Griseb. in Goett. Abh. 19: 87 (1874). C. glomerata var. boliviana Heimerl in Denkschr. Akad. Wiss. Wien, 70: 136 1900). C. parvior ye Endl. Gen. 311 (1837), combination implied but not actu- ally m. C. ieee ious 0. Ktze. Rev. Gen. 3(2): 264 (1898). paar crassifolius Heimerl in Notizbl. 11: 461 (1932). Crypt us pyriformis HBK. Nov. Gen, & Spec. 2: 188, t. 124 (1817). Mirabilis ae i L. Sp. Pl. 177 (1758). M. micrantha (Chois.) Heimerl in Engl. & Prantl, Nat. Pflanzenfam. 3(lb): 24 (1889). M. prostrata (R. & P.) Heimerl, Beitr. Syst. Nyct. 21 (1897). Neea Bangii Rusby in Bull. N. Y. Bot. Gard. 4: 435 (1907). boliviana Standl. in Field Mus. Publ. Bot. 11: 81 (1981). Brittonii Standl. in Field Mus. Publ. Bot. 11: 80 (1931). dimorphophylla Standl. in Field Mus. Publ. Bot. 11: 78 (1931). divaricata Poepp. & Endl. Nov. Gen. 2: 45, t. 161 (1838). longipedunculata Britton in Rusby in Bull. Torr. Bot. Club, 27: 126 (1900). macrophylla Britton in Rusby in Bull. Torr. Bot. Club, 27: 126 (1900) = N. Brittonii. . mapirensis Standl. in Field Mus. Publ. Bot. 11: 78 (1931). Oxybaphus bracteosus Griseb. in Goett. Abh. 19: 86 (1874). Pisonia aculeata forma inermis O. Ktze. Rev. Gen. 3(2): 265 (1898) = P. Zapallo. P. hirtella HBK. Nov. Gen. & Spec. 2: 217 (1818). P. indecora Heimer! in Notizbl. 11: 469 (1932). P. Zapallo Griseb. in Goett. Abh. 24: 39 (1879). Pisoniella arborescens var. glabrata (Heimerl) Heimer! in Meded. Rijks Herbar. 19: 35 (1913). Torrubia boliviana (Britton). Standl. in Contrib. U. S. Nat. Herb. 18: 100 (1916). T. Hassleriana (Heimerl) Standl. in Contrib. U. S. Nat. Herb. 18: 100 (1916). T. Olfersiana (Lk. Kl. & Otto) Standl. in Contrib. U. S. Nat. Herb. 18: 101 2AmA2z vA (1916). T. suspensa (Heimerl) Standl. in Contrib. U. S. Nat. Herb. 18: 101 (1916). PHYTOLACCACEAE Achatocarpus microcarpus Schinz & Autran in Bull. Herb. Boiss. 1: 8 (1893). A. praecox Griseb. in Goett. Abh. 24: 32 (1879). pena Gorazema (Vell.) Mog. in DC. Prodr. 18(2): 8 (1849). crotea maypurensis (HBK.) G. Don in Loud. Hort. Brit. (ed. 2) 98 (1832). M. paniculata var. latifolia O. Ktze. Rev. Gen. 3(2): 268 (1898). M. scandens Rusby in Mem. N. Y. Bot. Gard. 7: 239 (1927). Mohlana nemoralis Mart. Nov. Gen. 3: 171, t. 290 (1829). M. secunda (R. & P.) Mart. Nov. Gen. 3: 172 (1829). Petiveria alliacea L. Sp. Pl. 342 (1753). Phytolacea australis Phil. in Anal. Univ. Chile, 1873: 536. P. icosandra L. Sp. Pl. (ed. 2) 1: 681 (1762) P. octandra L. Sp. Pl. (ed. 2) 1: 631 (1762). P. rivinoides Kunth & Bouché, Ind. Sem. Hort. Berol. 15 (1848). P. thyrsiflora Fenzl in Mart. Fl. Bras. 14(2): 343 (1872). Rivina humilis L. Sp. Pl. 121 (1758). R. humilis var. glabra L. Sp. Pl. 122 (1753). Schindleria densiflora (O. Ktze.) Monachino in Phytologia, 4: 39 (1952). S. mollis H. Walt. in Engler, Pflanzenr. IV, 83: 116 (1909). S. racemosa (Britton) H. Walt. in Engler, Bot. Jahrb. 37, Beibl. 85: 24 (1906). S. rivinoides (Rusby) H. Walt. in Engler, Bot. Jahrb. 37, Beibl. 85: 24 (1906). S. rosea H. Walt. in Engler, Bot. Jahrb. 37, Beibl. 85: 24 (1906). Seguieria brevithyrsa H. Walt. in Engler, Pflanzenr. IV, 83: 87 (1909). S. guaranitica Speg. in Anal. Soc. Cient. Argent. 16: 88 (1883). S. macrophylla Benth. in Trans. Linn. Soc. 18: 235 (1839). S. paraguayensis Morong in Ann. N. Y. Acad. Sci. 7: 210 (1898). S. parvifolia Benth. in Trans. Linn. Soc. 18: 235 (1839). Trichostigma octandrum (L.) H. Walt. in Engler, Pflanzenr. IV, 83: 109 (1909). AIZOACEAE Mollugo verticillata L. Sp. Pl. 89 (1758). PORTULACACEAE Calandrinia acaulis HBK. Nov. Gen. & Spec. 6: 78 (1823). C. chromantha Griseb. in Goett. Abh. 24: 30 (1879). C. ciliata (R. & P.) DC. Prodr. 3: 359 (1828). Portulaca elongata Rusby in Mem. Torr. Bot. Club, 6: 7 (1896). P. fragilis v. Poelln. in Fedde, Repert. Spec. Nov. 50: 61 (1941). Gilliesii var. pedicellata Legrand in Lilloa, 17: 351 (1949). gracilis v. Poelln. in Fedde, Repert. Spec. Nov. 50: 107 (1941). grandiflora var. macrophylla Rohrb. in Mart. Fl. Bras. 14(2): 302 (1872). longiusculo-tuberculata v. Poelln. in Fedde, Repert. Spec. Nov. 33: 161 33). marginata HBK. Nov. Gen. & Spec. 6: 72 (1823). pedicellata var. cochabambensis Legrand in Lilloa, 17: 356 (1949). perennis R. E. Fries in Nov. Act. Soc. Sci. Upsal. (ser. 4) 1(1): 151, t. 8 (1905). Philippii I. M. Johnst. in Contrib. Gray Herb. 85:39 (1929). pilosa L. Sp. Pl. 445 (1753). rotundifolia R. E. Fries in Nov. Act. Soc. Sci. Upsal. (ser. 4) 1(1): 149 By wet Wey ww simpiciuseula Mart. ex Rohrb. in Mart. Fl. Bras. 14(2): 301, t. 68 (1872). sien patens (L.) Willd. Sp. Pl. 2: 863 (1799). BASELLACEAE ? Boussingaultia baselloides HBK. Nov. Gen. & Spec. 7: 196, t. 645 bis (1825). B. boliviensis (Hauman) Macbr. in Field Mus. Publ. Bot. 13: 577 (1987). B. tucumanensis Lillo & Hauman ex Haum. in Anal. Mus, Hist. Nat. Buenos Aires, 33: 353 (1925). Ullucus tuberosus Caldas, Seman. Nuov. Granad. 185 (1809). CARYOPHYLLACEAE Acanthonychia polycnemoides Rohrb. in Mart. Fl. Bras. 14(2): 250 (1872). A. ramosissima (Weinm.) Rohrb. in Mart. Fl. Bras. 14(2): 249, t. 56 (1872). Alsine yungasensis Rusby in Phytologia, 1: 53 (1934) = ? 78 pear Ta (Michx.) Rohrb. in Mart. Fl. Bras. 14(2): 274 (1872), in A, aivithia ‘Rohrb. in Linnaea, 37: 255 (1871-738). A. boliviana Williams in Journ. Linn. Soc. 33: 425 (1898). catamarcensis Pax in Engler, Bot. Jahrb. 18: 29 (1893). conferta Wedd. in Ann. Sci. Nat. (ser. 5) 1: 293 (1864) — A. boliviana. conferta var. villosa Wedd. in Ann. Sci. Nat. (ser. 5) 1: 293 (1864) = digyna Schlechtd. in Ges. Naturf. Fr. Berl. Mag. 7: 201 (1813). lanuginosa (Michx.) Rohrb. in Mart. Fl. Bras. 14(2): 274 (1872). Mandoniana Wedd. in Ann. Sci. Nat. (ser. 5) 1: 294 (1864). Orbignyana Wedd. in Ann.Sci. Nat. (ser. 5) 1: 293 (1864). pedunculosa Wedd. in Ann. Sci. Nat. (ser. 5) 1: 294 (1864). A. pycnophylla Rohrb. in Linnaea, 37: 250 (1871-73). A. soratensis Rohrb. in Linnaea, 37: 266 (1871-73). A. Stuebelii Hieron. in Engler, Bot. Jahrb. 21: 307 (1895). Cerastium arvense L. Sp. Pl. 438 (1753). Sl ol of gl od og a C. arvense var. arvensiforme (Wedd.) Rohrb. in Linnaea, 37: 305 Ge 1- fh C. arvensiforme var. glandulosum Wedd. in Ann. Sci. Nat. (ser. 5) 1 (1864). C. Behmianum Muschl. in Engler, Bot. Jahrb. wt 446 (1911). C. brevicarpum Rusby in Phytologia,1: 54 (193 C. consanguineum Wedd. in Ann. Sci. Nat. oe 5) 1: 296 (1864). C. crassipes Bartl. in Presl, Rel. Haenk. 2: 18 (1831). C. humifusum Camb. ex St. Hil. Fl. Bras. Mer. 2: 166 (1829). C. imbricatum HBK. Nov. Gen. & Spec. 6: 28 (1823) C. imbricatum var. Mandonianum Rohrb. in Linnaea, ‘37: 293 (1871-73). C. mucronatum Wedd. in Ann. Sci. Nat. (ser. 5) 1: 294 (1864). C. nutans Raf. Préc. Découv. 36 (1814). C. orophilum Wedd. in Ann. Sci. Nat. (ser. 5) 1: 295 (1864). C. soratense Rohrb. in Linnaea, 37: 291 (1871-73). C. subspicatum Wedd. in Ann. Sci. Nat. (ser. 5) 1: 295 (1864). C. viscosum L, Sp. Pl. 437 (1753). C. vulgatum L. Fl. Suec. (ed. 2) 158 (1755). C. vulgatum var. peruvianum A. Gray in Bot. U. S. Explor. Exped. 1: 120 (1854 Colobanthus bolivianus Pax in Fedde, Repert. Spec. Nov. 7:.109 (1909). Drymaria cordata (L.) Willd. ex Roem. & Schult. Syst. Veg. 5: 406 (1819). glandulosa Bartl. in Presl, Rel. Haenk. 2: 9 (1831). hirsuta Bartl. in Presl, Rel. Haenk. 2: 8 (1831). Ladewii Rusby in Phytologia, 1: 54 (1934). - pauciflora Bartl. in Presl, Rel. Haenk. 2: 8 (1831). sperguloides A. Gray, Pl. Fendl. 11 (1849). . stricta Rusby in Phytologia, 1: 55 (1934). Herniaria setigera Gill. in Hook. Bot. Misc. 3: 337 (1833). Lychnis andicola (Gill.) Britton in Bull. Torr. Bot. Club, 16: 61 (1889). Melandrium cucubaloides Fenzl ex Rohrb. in Linnaea, 36: 223 (1869-70) = M. Mandonii Rohrb. in Linnaea, 36: 222 (1869-70) = Lychnis andicola ? Paronychia andina A. Gray in Bot. U. S. Explor. Exped. 1: 128 (1854). P. chilensis DC. Prodr. 3: 370 (1828). P. Mandoniana Rohrb. in Linnaea, 37: 208 (1871-73). P. microphylla Phil. in Anal. Mus. Nac. Chile, Bot. 8: 26 (1891). P. sessiliflora Nutt. Gen. Am. Pl. 1: 160 (1818). Pentacaena ramosissima (Weinm.) Hook. & Arn. in Hook. Bot. Misc. 3: 338 (1833). Polycarpaea Hassleriana Chod. in Bull. Herb. Boiss. (ser.2) 3: 791 (1903). DyOpUD 79 Pycnophyllopsis keraiopetala Mattf. in Schr. Ver. Naturf. Unterwes. 7: 22 (1934). Pycnophyllum dicranoides (HBK.) Muschl. in Engler, Bot. Jahrb. 45: 454 (191 kobalanthom Mattf. in Fedde, Repert. Spec. Nov. 36: 274 (1934). molle Remy in Ann. Sci. Nat. (ser. 3) 6: 355, t. 20 (1846). Pilgerianum Muschl. in Engler, Bot. Jahrb. 45: 454 (1911). spathulatum Mattf. in Fedde, Repert. Spec. Nov. 18: 175 (1922). Stuebelii Mattf. in Fedde, Repert. Spec. Nov. 18: 171 (1922). tetrastichum Remy in Ann. Sci. Nat. (ser. 3) 6: 356, t. 20 (1846). Sagina graminifolia Wedd. in Ann. Sci. Nat. (ser. 5) 1: 292 (1864). Silene gallica L. Sp. Pl. 417 (1753). Spergularia andina Rohrb. in Linnaea, 37: 234 (1871-73). S. firma Kze. ex Kindb. Monog. Lepig. 34 (1863), in synon. S. floribunda Rohrb. in Linnaea, 37: 230 (1871-73). Ss. ry (Rusby) R. P. Rossbach in Rhodora, 42: 205 (1940). ;. a Camb. in St. Hil. Fl. Bras. Mer. 2: 178 (1829). Stellaria lactupetela Benth. Pl. Hartw. 163 (1845). S. media (L.) Sowerby & Smith, English Bot. 8: t. 537 (1799). S. nemorum L. Sp. Pl. 42 1(1753). em NYMPHAEACEAE Cabomba piauhyensis Gardn. in Hook. Ic. Pl. 7: t. 641 (1844). RANUNCULACEAE Anemone decapetala var. foliolosa Eich]. in Mart. Fi. Bras. 13(1): 151, t. 35 (1864 ; Caltha alata A. W. Hill in Ann. Bot. a 428, fig. 3-4 (1918) — C. sagittata. C. sagittata Cav. Ic. 5: 8, t. 414 (179 Clematis alborosea Ulbr. in Notizbl. . "325 (1923). Bangii Rusby in Mem. Torr. Bot. Club, 3(3): 3: (1893) = C. Hilarii. cochabambensis Rusby in Mem. Torr. Bot. Club, 3(3): 8..(1803): =: G millefoliata dioica var. bresilians (DC.) Eich]. in Mart. Fl. Bras. 13(1): 148 (1864). Hilarii Spreng. Syst. Veg. 5: 177 (1828). millefoliata Eichl. in Mart. Fl. Bras. 18(1): 150 (1864). aaaa ana R. Cymbalaria Pursh, Fl. Am. Sept. 2: 392 (1814). R. Cymbalaria var. exilis R. E. Fries in Nov. Act. Soc. Sci. Upsal. (ser. 4) 1(1): 145 (1905). filamentosus Wedd. Chlor. And. 2: 301 (1861). flagelliformis Smith in Rees, Cycl. 29: sod 13 (1819). Guzmannii Humb. Tabl. Reg. Eq. 69 (1805). haemanthus Ulbr. in Engler, Bot. Jahrb. nt 404 (1906). Mandonianus Wedd. Chlor. And. 2: 299 (1861). minutiusculus Ulbr. in Engler, Bot. Jahrb. 37: 403 (1906) = R. Cymbalaria. var. exilis. praemorsus HBK. Nov. Gen. & Spec. 5:47 (1821). psychrophilus Wedd. Chlor. And. 2: 300 (1861). repens L. Sp. Pl. 554 (1753). tichaphyiive Chaix in Vill. Hist. Pl. Dauph. 1: 335 (1786), nomen nudum. Rhopalodium Gusmanni var. bolivianwm Ulbr. in Notizbl. 8: 259 (1922); = Ranunculus Guzmannii ? VAP AP NAPPA A 80 Thalictrum Cardenasianum Boivin in Bull. Torr. Bot. Club, 80: 136 (1953). T. cincinnatum Boivin in Rhodora, 46: 398 (1944). T. decipiens Boivin in Rhodora, 46: 405 (1944). T. imuncans Boivin in Rhodora, 46: 404 (1944) = T. decipiens . T. Steinbachit Boivin in Rhodora, 46: 397 (1944) = T. cincinnatum. T. Venturii Boivin in Rhodora, 46: 422 (1944). BERBERIDACEAE Berberis agapatensis Lechl. Berb. Amer. Austr. 44 (1857). B. bumeliaefolia C. K. Schneider in Bull. Herb. Boiss. (ser. 2) 5: 818 (1905). B. carinata Lechl. Berb. Amer, Austr. 41 (1857). B. chrysacantha C. K. Schneider in Engler, Bot. Jahrb. 42: 81 (1908). B. ciliaris Lindl. in Journ. Hort. Soc. 5: 5 (1850 B. commutata Eichl. in Mart. Fl. Bras. 13(1): 231 ‘(186 4). B. conferta var. boliviana (Lechl.) C. K. Schneider in Bull. Herb. Boiss. (ser. 2) 5: 393 (1905). B. densifolia Rusby, Descr. S. Am. Pls. 16 (1920). B. divaricata Rusby in Mem. Torr. Bot. Club, 4(3): 203 (1895) — B. commu- tata. B. edentata Rusby in Mem. N. Y. Bot. Gard. 7: 239 (1927). B. ferruginea Lechl. Berb, Amer. Austr. 9 (1857). B. Fiebrigii C. K. Schneider in Engler. Bot. Jahrb. 42: 85 (1908 B. Keissleriana C. K. Schneider in aie Herb. Boiss. (ser. 2) 5: "808 (1905). B. laurina Thunb. Pl. Bras, 1: 8 (18 B. Lobbiana (Schneid.) C. K. ei in gies Bot. Jahrb. 42: 83 (1908). B. ovalifolia Rusby, Descr. S. Am. Pls. 16 (192 B. paucidentata Rusby in Bull. N. Y. Bot. a #: 321 (1907). B. phyllacantha Rusby in Mem. Torr. Bot. Club, 6: 4 (1896). B. pseudo-spinulosa Job in Not. Mus. La. Plata, 8: 129 (1948) . B. rariflora Lechl. Berb. Amer. Austr. 33 (1857). B. Rechingeri C. K. Schneider in Bull. Herb. Boiss. (ser. 2) 6 808 (1905). B. rectinervia Rusby in Mem. Torr. Bot. Club, 3(3): 5 (1893 B. Trollii Diels in Notizbl. 11: 781 (1933). B. Weddellii Lechl. Berb. Amer. Austr. 21 (1857). B. Wettsteiniana C. K. Schneider in Bull. Herb. Boiss. (ser. 2) 5: 809 (1905). MENISPERMACEAE Abuta boliviana Rusby in Mem. N. Y. Bot. Gard. 7: 241 (1927) = Chondoden- dron tomentocarpum. A. Candollei Tri. & Pl. in Ann. Sci. Nat. (ser. 7 ane 47 (1862). A. concolor Poepp. & Endl. Nov. Gen. 2: 64, t. 8 (1838). A. grandifolia (Mart.) Sandw. in Kew Bull. ar 397. A. splendida Krukoff & Moldenke in Bull. Torr. Bot. ona 68: 241 (1941). A. trinervis (Rusby) Moldenke in Brittonia, 3: 59 (19 Anomospermum bolivianum Krukoff & Moldenke in ies 5: 234 (1940). A. Schomburgkii Miers, Contrib. Bot. 3: 71 (1871). Chondodendron tomentocarpum (Rusby) Moldenke in Brittonia, 3: 21 (1938). C. tomentosum R. & P. Syst. 261 (1798). Cissampelos ciliata Rusby in Mem. N. Y. Bot. Gard. 7: 240 (1927). C. ovalifolia DC. Syst. 1: 587 (1818). C. Pareira L. Sp. ‘Pl. 1031 (1753). C. Pareira var. Gardneri Diels in Engler, Pflanzenr. IV, 94: 294 (1910). C. Pareira var. Haenkeana (Presl) Diels in Engler, Palativenr. IV, 94: 292 (1910). 81 C. sympodialis var. grandifolia Britton in Bull. Torr. Bot. Club, 16: 15 (1889). C. tropaeolifolia DC. Syst. 1: 5382 (1818). C. violaefolia Rusby in Mem, N. Y. Bot. Gard. 7: 240 (1927). Disciphania clausa Diels in Engler, Pflanzenr. IV, 94: 176 (1910). D. cubijensis (R. Knuth) Sandw. in Kew Bull. 1954: 614 (1955). Hyperbaena domingensis (P. DC.) Benth. in Journ. Linn. Soc. 5, Suppl. 2: 50 (1861). H. Hassleri Diels in Engler, Pflanzenr. IV, 94: 201 (1910). ? Somphoxylon sp ANNONACEAE Annona Cherimola Mill. Gard. Dict. (ed. 4 no. 5 (1768). dioica St. Hil. Fl. Bras. Mer. 1: 34 (1825). . hypoglauca Mart. Fl. Bras. 13(1): 18 (1841). nano-fruticosa fsa in Fedde, Repert. Spec. Nov. 7: 51 (1909). nutans (R. E. es) R. E. Fries in Bull. Herb. Boiss. (ser. 2) 4: 1171 (1904). . reticulata L. Sp. Pl. 587 (1753). Cardiopetalum calophyllum Schlechtd. in Linnaea, 9: 328 (1835). Crematosperma leiophyllum (Diels) R. E. Fries in Act. Hort. Berg. 10(2): 328 (1931). C. monospermum (Rusby) R. E. Fries in Act. Hort. Berg. 10(2): 327 (19381). Cymbopetalum longipes Diels in Verh. Bot. Ver. Brandenb. 47: 132 (1906). C. parvifolium Rusby in Bull. N. Y. Bot. Gard. 6: 505 (1910) = Cardiopetalum calophyllum. Duguetia ibonensis Rusby in Mem. N. Y. Bot. Gard. 7: 246 (1927) = D. quitar- PPP > > ensis. D. quitarensis Benth. in Hook. Lond. Journ. Bot. 2: 361 (1843). Guatteria alutacea var. Steinbachii R. E. Fries in Act. Hort. Berg. 12(3): 862 (1939). G. boliviana H. Winkl. in Fedde, Repert. Spec. Nov. 7: 242 a Buchtienii R. E. Fries in Act. Hort. Berg. 12(3): 388 (1939). ta Rusby in Mem, N. Y. Bot. Gard. 7: 245 (1927) = ? (not Annona- ceae, fide R. E. Fries). Guentheriana Diels in Notizbl. 11: 75 (1931). lasiocalyx R. E. Fries in Act. Hort. Berg. 12(3): 388 (1939). lucida Rusby in Mem. N. Y. Bot. Gard. 7: 245 (1927) = Crematosperma onospermum. oblongifolia Rusby in Bull. N. Y. Bot. Gard. 4: 320 (1907). Rusbyi Macbr. in Field Mus. Publ. Bot. 4: 171 (1929) == Crematosperma monospermum. — setosa Rusby in Phytologia, 1: 55 (1934) — G. trichoclonia. tomentosa Rusby in Bull. N. Y. Bot. Gard. 6: 504 (1910). trichoclonia Diels in Notizbl. 11: 77 (1981). Oxandra Espintana (Spruce) Baill. Hist. Pl. 1: 207 (1868). O. ovata Rusby, Descr. S. Am. Pls. 19 (1920) = O. Espintana. Porcelia nitidifolia R. & P. Syst. 144 (1798). P. ponderosa (Rusby) Rusby in Mem. N. Y. Bot. Gard. 7: 245 (1927) = P. nitidifolia. P. Saffordiana Rusby in Mem. N, Y. Bot. Gard. 7: 242, fig. 2 (1927) = P. nitidifolia. P. Steinbachii (Diels) R. E. Fries in Act. Hort. Berg. 10(1): 35 (1930). Rollinia boliviana R. E. Fries in Act. Hort. Berg. 13(3): 115 (1941). R. Hassleriana var. vestita R. E. Fries in Act. Hort. Berg. 12(1): 171 (1934). R. Herzogii R. E. Fries in Act. Hort. Berg. 12(1): 177 (1934). AAA AR AAD ag: 82 R. Williamsii Rusby ex R. E. Fries in Act. Hort. Berg. 12(1): 186 (1934). Ruizodendron ovale (R. & P.) R. E. Fries in Ark. Bot. 28B (4): 3 (1936). Trigynaea Periquino Rusby in Mem. N. Y. Bot. Gard. 7: 247 (1927) = Un- onopsis sp. Unonopsis boliviensis (Britton) R. E. Fries in Kgl. Sv. Vet. Akad. Handl. 34(5): 28 (1900). U. Buchtienii R. E. Fries in Fedde, Repert. Spec. Nov. 24: 247 (1928). U. Guaraya Herzog in Fedde, Repert. Spec. Nov. 7: 52 (1909). Ui pegs nea R. E. Fries in Kgl. Sv. Vet. Akad. Handl. 34(5): 28 (19 MYRISTICACEAE Dialyanthera parvifolia Markgraf in Notizbl. 9: 964 (1926). Virola seen a Warb. in Nov. Act. Acad. Caes. Leop. 68: 184 (1897) = V. sebifer: V. flexuosa A. °o. Smith in Brittonia, 2: 151 (1936). V. sebifera Aubl. Pl. Guian. Fr. 2: 904, t. 345 (1775). MONIMIACEAE Mollinedia boliviensis A. DC. in Journ. Bot. 3: 220 (1865). M. caloneura Perk. in Engler, Bot. Jahrb, 27: 663 (1900). M. ovata R. & P. Syst. 143 (1798). M. Rusbyana Perk. in Engler, Bot. Jahrb. 27: 682 (1900). M. Steinbachiana Perk. in Notizbl.. 10: 160 (1927). Siparuna Apiosyce (Mart.) A. DC. in DC. Prodr. 16(2): 645 (1864). S. bifida (Poepp. & Endl.) A. DC. in DC. Prodr. 16(2): 652 (1864). S. boliviensis Herzog in Meded. Rijks Herbar. 40: 3 (1921). S. chrysantha Perk. in Engler, Bot. Jahrb. 28: 685 (1901). S. cinerea Perk. in Engler, Bot. Jahrb. at 695 (1901). S. dasyantha Perk. in Notizbl. 6: 134 (19 S. guianensis Aubl. Pl. Guian. Fr. 2: re t' 333 (1775). S. hispida A. DC. in Journ. Bot. 3: 219 (18 S. hypoglauca Perk. in Engler, Bot. Jahrb. Bs 691 (1901). S. limoniodora (R. & P.) A. DC. in DC. Prodr. 16(2): 646 (1864). S. macrophylla (HBK.) A. DC. in Prodr. 16(2): 646 (1864). S. muricato-alata Herzog in Meded. Rijks Herbar. 40: 4 (1921). S. nigra Rusby in Mem. Torr. Bot. Club, 4(3): 252 (1895). S. obovata (Gardn.) A. DC. in DC. Prodr. 16(2): 644 (1864). S. pellita (Tul.) A. DC. in DC. Prodr. 16(2): 645 (1864). S. polyantha (Tul.) A. DC. in DC. Prodr. 16(2): 646 (1864). S. pseudospectabilis Sleumer in Fedde, Repert. Spec. Nov. 39: 276 (1935). S. spectabilis Perk. in Engler, Bot. Jahrb. 28: 686 (1901) S. Sprucei A. DC. in Journ. Bot. 3: 219 (1865). S. tetradenia Perk. in Notizbl. 10: 163 (1927). S. tomentosa (R. & P.) Perk. in Engler, Bot. Jahrb. 28: 691 (1901). LAURACEAE Acrodiclidium benense Rusby in Bull. Torr. Bot. Club, 49: 262 (1922) = Licaria limbosa Aniba Coto (Rea) Kosterm. in Macbr. in Field Mus. Publ. Bot. 13: 863 (1937 A. gigvctitole O. C. Schmidt in Notizbl. 10: 225 (1928). A. Muca (R. & P.) Mez. in Jahrb. Bot. Gart. Berl. 5: 57 (1889). 83 A. perutilis Hemsl. in Kew Bull. 1894: 197. A. pseudo-coto (Rusby) Kosterm. in Rec. Trav. Bot. Néerl. 35: 872 (1938).. ; hury-minor (Mart.) Mez in Jahrb. Bot. Gart. Berl. 5: 70 (1889). Endlicheria boliviensis Kosterm. in Rec. Trav. Bot Néerl. 34: 553 (1937). dysodantha (R. & P.) Mez in Jahrb. Bot. Gart. Berl. 5: 119 (1889). . Lhotzkyi (Nees) Mez in Jahrb. Bot. Gart. Berl. 5: 122 (1889). pyriformis (Nees) Mez in Jahrb. Bot. Gart. Berl. 5: 116 (1889). Szyszylowiczii Mez in Jahrb. Bot. Gart. Berl. 5: 121 (1889). . tomentella Mez in Jahrb. Bot. Gart. Berl. 5:115 (1889). Licaria amara (Mez) Kosterm. in Rec, Trav. Bot. Néerl. 34: 583 (1937). L. limbosa (R. & P.) Kosterm. in Rec. Trav. Bot. Néerl. 34: 585 (1937). Nectandra acutifolia (R. & P.) Mez in Jahrb. Bot. Gart. Berl. 5: 409 (1889). . Brittonii Mez in Jahrb. Bot. Gart. Berl. 5: 485 (1889). . citrifolia Mez & Rusby in Mem. Torr. Bot. Club, 6: 115 (1896). cuspidata Nees & Mart. in Nees, Syst. Laur. 330 (1836). guanaiensis Rusby in Bull. N. Y. Bot. Gard. 6: 508 (1910). laevis Mez in Jahrb. Bot. Gart. Berl. 5: 451 (1889). lanceolata Nees & Mart. in Linnaea, 8: 47 (1833). Laurel Klotzsch ex Nees in Linnaea, 21: 505 (1848). Pichurim (HBK.) Mez in Jahrb. Bot. Gart. Berl. 5: 449 (1889). pisi Miq. Stirp. Surin. Sel. 199 (1850). pulverulenta Nees, Syst. Laurin. 283 (1836). reticulata (R. & P.) Mez in Jahrb. Bot. Gart. Berl. 5: 404 (1889). Steinbachii O. C. Schmidt in Fedde, Repert. Spec. Nov. 31: 186 (1933). armingii Meissn. in Warm. in Vid. Medd. 1870: 141. tea albida Mez & Rusby in Mem. Torr. Bot. Club, 6: 114 (1896). Bangii Mez & Rusby in Mem. Torr. Bot. Club, 6: 115 (1896) — Endlicheria bel be be bl rmis. cuneifolia (R. & P.) Mez in Jahrb. Bot. Gart. Berl. 5: 259 (1889). cuprea (Meissn.) Mez in Jahrb. Bot. Gart. Berl. 5: 299 (1889). guianensis Aubl. Pl. Guian. Fr. 2: 781, t. 310 (1775). illustris Rusby in Bull. N. Y. Bot. Gard. 6: 507 (1910). Jelskii Mez in Jahrb. Bot. Gart. Berl. 5: 261 (1889). laxiflora (Meissn.) Mez in Jahrb. Bot. Gart. Berl. 5: 371 (1889). Mandonii Mez in Jahrb. Bot. Gart. Berl. 5: 311 (1889). maranhana Rusby in Bull. N. Y. Bot. Gard. 6: 507 (1910), nomen nudum. marmellensis Mez in Bull. Herb. Boiss. (ser. 2) 3: 238 (1903). marowynensis Mez in Jahrb. Bot. Gart. Berl. 5: 380 (1889). oblanceolata Rusby in Bull. N. Y. Bot. Gard. 6: 507 (1910) = Aniba Muca. oblonga (Meissn.) Mez in Jahrb. Bot. Gart. Berl. 5: 367 (1889). proboscidea Rusby in Bull. N. Y. Bot. Gard. 4: 318 (1907). prunifolia Rusby in Bull. N. Y. Bot. Gard. 4: 439 (1907). reticulata Mez in Jahrb. Bot. Gart. Berl. 5: 303 (1889). rubrinervis Mez in Jahrb. Bot. Gart. Berl. 5: 351 (1889). Rusbyana Mez in Jahrb. Bot. Gart. Berl. 5: 303 (1889). O. suaveolens var. robusta Hassl. in Ann. Conserv. & Jard. Bot. Genéve, 21: 90 (1919). Persea boliviensis Mez & Rusby in Mem. Torr. Bot. Club, 6: 113 (1896). Buchtienii O. C. Schmidt in Fedde, Repert. Spec. Nov. 31: 179 (1933). caerulea (R. & P.) Mez in Jahrb. Bot. Gart. Berl. 5: 171 (1889). filipes Rusby in Bull. N. Y. Bot. Gard. 6: 505 (1910). gratissima Gaertn. Fruct. 3: 222, t. 221 (1805). laevigata HBK. Nov. Gen. & Spec. 2: 157 (1817). negracotensis O. C. Schmidt in Fedde, Repert. = Nov. 31: 180 (1933). scoparia Mez in Arb. Bot. Gart. Breslau, 1: 115 (1892). S99999999999999900 ORS RS RERRRE Re A A 84 ‘P. Trollii O. C. Schmidt in Fedde, Repert. Spec. Nov. 31: 180 (1933). P. vestita Mez in Jahrb. Bot. Gart. Berl. 5: 154 (1889). Phoebe porphyria (Griseb.) Mez in Jahrb. Bot. Gart. Berl. 5: 199 (1889). HERNANDIACEAE Sparattanthelium acreanum Pilger in Notizbl. 6: 295 (1915). S. botocudorum Mart. ex Meissn. in Mart. Fl. Bras. 5(2): 293 (1866). S. Burchellii Rusby in Bull. N. Y. Bot. Gard. 8: 109 (1912). _§. glabrum Rusby in Mem. Torr. Bot. Club, 6: 35 (1896). PAPAVERACEAE Argemone mexicana L. Sp. Pl. 508 (1753). Bocconia integrifolia Humb. & Bonpl. Pl. Aequin. 1: 119, t. 35 (1807). B. Pearcei Hutchinson in Kew Bull. 1920: 278. CAPPARIDACEAE Atamisquea emarginata Miers, Trav. Chile, 2: 529 (1826), nomen nudum. Capparis Bangii Rusby in Mem. Torr. Bot. Club, 6:5 (1896). C. cordata R. & P. ex DC. Prodr. 1: 251 (1824). C. retusa Griseb. in Goett. Abh. 24: 18 (1879). C. salicifolia Griseb. in Goett. Abh. 24: 17 (1879). C. Tweediana Eichl. in Mart. Fl. Bras. made oe (1865). Cleome aculeata L. Syst. (ed. 12) 3: 232 (17 Bangiana Gilg ex Heilborn in in Bot. tis ee 12 (1930). chilensis DC. Prodr. 1: 238 (18 consimilis A. Ernst in ae an 879 (1936). cordobensis Eichl. ex Griseb. in Goett. Abh. 19: 73 (1874). Eyerdamii Standl. & Barkl. in Madrofio, 9: 150 (1948). Friesii O. Heilborn in Ark. Bot. 23A(10): 7 (19380). gigantea L. Mant. 480 (1771). glandulosa R. & P. ex DC. Prodr. 1: 238 (1824). Trollii A. Ernst in Notizbl. 13: 380 (1936). tunarensis O. Ktze. Rev. Gen. 3(2): 7 (1898). Werdermannii A. Ernst in Notizbl. 18: 878 (1986). orisonia oblongifolia Britton in Bull. Torr. Bot. Club, 16: 17 (1889). BAAHHAHAAHAAAAH CRUCIFERAE Alyssum boliviense Muschl. in Engler, Bot. Jahrb. 40: 275 (1908) — Lesquer- ella mendocina. A. Pflanzii Muschl. in Engler, Bot. Jahrb. 49: 201 (1913) = Draba scopulorum. A. serene Muschl. in Engler, Bot. Jahrb. 40: 274 (1908) = Lesquerella doci Arabis Sractate Wedd. in Ann. Sci. Nat. (ser. 5) 1: 291 (1864). Aschersoniodoxa Mandoniana (Wedd.) Gilg & Muschl. in Engler, Bot. Jahrb. 42: 469 (1908). A. Rusbyi O. E. Schulz in Notizbl. 10: 115 (1927). Brassica juncea Coss. in Bull. Soc. Bot. France, 6: 609 (1859). Brayopsis calycina (Desv.) Gilg & Muschl. in Engler, Bot. Jahrb. 42: 484 (1908). B. calycina var. filiformis O. E. Schulz in Engler, Pflanzenr. IV, 105: 240 (1924). B. calycina var. filiformis forma leiophylla O. E. Schulz in Engler, Pflanzenr. IV, 105: 240 (1924). 85 B. — var. filiformis forma trichophylla O. E. Schulz in Engler, Pflan- nr. IV, 105: 240 (1924). B. fem be Gilg & Muschl. in Engler, Bot. Jahrb. 42: 484 (1909). Capsella Bursa-pastoris var. rubriflora Muschl. in Engler, Bot. Jahrb. 49: 199 (1918). Cardamine africana L. Sp. Pl. 655 (1758). axillaris Wedd. in Ann. Sci. Nat. (ser. 5) 1: 290 (1864). chenopodifolia Pers. Synops. Pl. 2: 195 (1807). flaccida subsp. minima (Steud.) O. E. Schulz in Engler, Bot. Jahrb. 32: 451 (1903). hispidula Phil. in Anal. Univ. Chile, 81: 79 (1892). ibaguensis Tri. & Pl. in Ann. Sci. Nat. (ser. 4) 17: 60 (1862). Jamesonii var. speciosa (Britton) O. E. Schulz in Engler, Bot. Jahrb. 32: 422 (1908). . ovata Benth Pl. Hartw. 158 (1845). ovata var. corymbosa Britton in Bull. Torr. Bot. Club, 16: 16 (1889). Cremaloins bolivianus Britton in Bull. Torr. Bot. Club, 16: 17 (1889). C. parviflorus Wedd. in Ann, Sci. Nat. (ser. 5) 1: 283 (1864). C. subscandens O. Ktze. Rev. Gen. 3(2): 4 (1898). Descurainia athroécarpa (A. Gray) O. E. Schulz in Engler, Pflanzenr. IV, 105: 340 (1924). . athrodcarpa var. Gilgiana (Muschl.) O. E. Schulz in Engler, Pflanzenr. IV, 105: 341 (1924). athroécarpa var. macrorrhiza O. E. Schulz in Engler, Pfianzenr: IV, 105: 341 (1924). depressa (Phil.) Prant] in Anal. Univ. Chile, 90: 148 (1895). depressa var. Pflanzii (Muschl.) O. E. Schulz in Engler, Pflanzenr. IV, 105: 339 i latisiliqua O. E. Schulz in Engler, Pflanzenr. IV, 105: 336 (1924). leptoclada Muschl. in Engler, Bot. Jahrb. 40: 272 (1908). Perkinsiana Muschl. in Engler, Bot. Jahrb. 49: 199 (1913). pulcherrima Muschl. in Engler, Bot. Jahrb 49: 200 (1913). Urbaniana Muschl. in Engler, Bot. Jahrb. 40: 271 (1908) = D. athrosécarpa. aba affinis Hook. f. Fl. Antarct. 2: 235 (1847). boliviana O. E. Schulz in Engler, Pflanzenr. IV, 105: 168 (1927). Brackenridgei A. Gray in Bot. U. S. Explor. Exped. 1: 53 (1854). ntha. Gilg in Engler, Bot. Jahrb. 42: 477 (1908) = D. Brackenridgei. discoidea Wedd. in Ann. Sci. Nat. (ser. 5) 1: 286 (1864). discoidea var. leiocarpa O. E. Schulz in Engler, Pflanzenr. IV, 105: 167 (1927). discoidea var. minor Wedd. ex O. E. Schulz in Engler, Pflanzenr. IV, 105: 167 (1927). Herzogii O. E. Schulz in Engler, Pflanzenr. IV, 105: 168 (1927). Macleanii Hook. f. Fl. Antarct. 2: 235 (1847). Pickeringii var. Pearcei O. E. Schulz in Engler, Pflanzenr. IV, 105: 135 (1927). scopulorum Wedd. in Ann. Sci. Nat. (ser. 5) 1: 286 (1864). scopulorum forma elongata Wedd. in Ann. Sci. Nat. (ser. 5) 1: 287 (1864). soratensis Wedd. in Ann. Sci. Nat. (ser. 5) 1: 287 (1864). Erysimum laxum Muschl. in Engler, Bot. Jahrb. 40: 273 (1908) = Sisymbrium pf Ape nae ~) PRS Ser Fr SPS ery ees. oy peruvianum. Eudema diapensioides (Wedd.) O. E. Schulz in Engler, Pflanzenr. IV, 105: 245 (1924). Halimolobos adpressa O. E. Schulz in Engler, Pflanzenr. IV, 105: 293 (1924). H. hispidula var. Weddellii (Fourn.) Rollins in Contrib. Dudley Herb. 3: 252 (1943). 86 H. montana (Griseb.) O. E. Schulz in Engler, Pflanzenr. IV, 105: 292 (1924). Heterothrix gracilis (Wedd.) O. E. Schulz in Engler, Pflanzént. IV, 105: 29 924). Hutchinsia pusillima Wedd. in Ann. Sci. Nat. (ser. 5) 1: 284 (1864). Lepidium abrotanifolium Turcz. in Bull. Soc. Nat. Mosc. 27(2): 308 (1854). abrotanifolium var. Steinmannii Thell. in Denkschr. Schweiz. Ges. Natur- wiss. 41(1): 247 (1906). affine Wedd. in Ann. Sci. Nat. (ser. 5) 1: 284 (1864) = L. Weddellii. angustifolium Rusby, Descr. S. Am, Pls. 23 (1920) pits not Lepidium]. bipinnatifidum Desv. in Journ. Bot. 3: 165 (1814), n ud Chichicara Desv. in Journ. Bot. 3: 165 (1814), nomen rages Chichicara var. rhombocarpum Thell. in Fedde, Repert. Spec. Now 11: 309 (1912). depressum Thell. in Denkschr. Schweiz. Ges. Naturwiss. 41 (1): 201 (1906). demissum C. L. Hitche. in Lilloa, 11: 121 , Meyenii subsp. gelidum (Wedd.) Thell. in Denkschr. Schweiz. Ges. Natur- wiss. 41(1): 203 (1906). Meyenii subsp. gelidum forma rhombicum Thell. in Mitteil. Bot. Mus. Univ. Ziirich, 28: 203 (1906). . Meyenii subsp. gelidum forma rotundatum Thell. in Mitteil. Bot. Mus. Univ. Ziirich, 28: 203 (1906). Philippianum (O. Ktze.) Thell. in Denkschr. Schweiz. Ges. Naturwiss. 41(1): 200 (1906). Philippianum var. boliviense Thell. in Denkschr. Schweiz. Ges. Naturwiss. 41(1): 201 (1906). quitense var. integrifolium Thell. in Denkschr. Schweiz. Ges. Naturwiss. 41(1): 213 (1906). reticulatum var. austro-americanum Thell, in Bull. Herb. Boiss. (ser. 2) 8: 914 (1908). seabrifructum C. L. Hitche. in Lilloa, 11: 119 (1945). Steinbachii O. E. Schulz in Notizbl. 9: 1037 (1926). Trianae Thell. in Denkschr. Schweiz. Ges. Naturwiss. 41(1): 214 (1906). ‘Walpersii Macbr. in Candollea, 5: 357 (1934) — L. depressum. L. Weddellii O. E. Schulz in Notizbl. 11: 391 (1932). Lesquerella mendocina (Phil.) Kurtz in Rev. Mus. LaPlata, 5: 286 (1893). Mancoa foliosa (Wedd.) O. E. Schulz in Engler , Bot. Jahrb. 66: 98 (1933). M. hispida Wedd. in Ann. Sci. Nat. (ser. 5) 1: 285 (1864). M. laevis Wedd. in Ann. Sci. Nat. (ser. 5) 1: 285 (1864). M. minima Rollins in Contrib. Dudley Herb. 3: 194 (1941). Mathewsia boliviana Gilg & Muschl. in Engler, Bot. Jahrb. 42: 464 (1908) = Rorippa nana. M. diffusa Rusby in Bull, N. Y. Bot. Gard. 4: 322 (1907) = Rorippa sp. ? Nasturtium bonariense var. erectum (Trev.) O. E. Schulz in Fedde, Repert. pec. Nov. 33: 283 (1934). N. clandestinum var. brevistylum O. E. Schulz in Fedde, Repert. Spec. Nov. 33: 283 (1934). % hee ore Mee ee She Pe Pia N. pubescens var. punense O. Ktze. Rev. Gen. 3(2): 6 (1898). N. pubescens var. serratifolium O. Ktze. Rev. Gen. 3(2): 6 (1898). Polypsecadium Harmsianum (Muschl.) O. E. Schulz in Engler, Pflanzenr. IV, 105: 177 (1924) P. Harmsianum var. dentatum (Muschl.) O. E. Schulz in Engler, Pflanzenr. IV, 105: 177 (1924). Radicula seabra Rusby, Descr. S. Am, Pls. 23 (1920) = Mancoa hispida. R. nana (Wedd.) Rusby, Deser. S. Am. Pls. 28 (1920) — Rorippa nana. Rorippa nana (Schlechtd.) Macbr. in Field Mus. Publ. Bot. 13(2): 965 (1938). R. Nasturtium-aquaticum (L.) G. Beck, Fl. Nied.—Oesterr. 1: 463 (1892). 87 Sarcodraba Herzogii O. E. Schulz in Notizbl. 10: 563 (1929). Sisymbrium anomalum Wedd. in Ann. Sci. Nat. (ser. 5) 1: 289 (1864) = S. peruvianum calycinum (Deer Wedd. in Ann. Sci. Nat. (ser. 5) 1: 289 (1864). fragile Wedd. in Ann. Sci. Nat. (ser. 5) 1: 288 (1864). lanatum (Walp.) O. E. Schulz in Notizbl. 11: 642 (1932). Mandonii Fourn. Thése Crucif. 109 ‘ . officinale (L.) Scopoli, Fl. Carn. (ed. 2) 2: 26 (1772). oliganthum Wedd. in Ann. Sci. Nat. (ser. S 1: 289 (1864) = Weberbauera pusilla. Orbignyanum Fourn. Thése Crucif. 107 (1865). orophilum Wedd. in Ann. Sci. Nat. (ser. 5) 1: 288 (1864). pazensis Rusby in Mem. Torr. Bot. Club, 8(3): 5 (1893) = Halimolobos Weddellii. . peruvianum DC. Syst. Nat. 2: 477 (1821). Rusbyi Britton in Bull. Torr. Bot. Club, 16: 16 apse . setaceum Wedd. in Ann. Sci. Nat. (ser. 5) 1: 289 (1864). Streptanthus boliviensis Muschl. in Engler, Bot. Jahrb. 40: 268 (1908) = Heterothrix gracilis. Thlaspi alpestre L. Sp. Pl. (ed. 2) 2: 903 (1768). Weberbauera pusilla (Gill.) O. E. Schulz in Engler, Pfianzenr. IV, 105: 194 (1924). Dnnn NH Nn wn TOVARIACEAE Tovaria pendula R. & P. Fl. Peruv. 3: 73, t. 306 (1802). DROSERACEAE Drosera montana St. Hil. Pl. Remarg. Brés. 260 (1824). PODOSTEMACEAE Apinagia boliviana v. Roy. in Meded. Bot. Mus. Utrecht, 107: 130 (1951). A. fluitans v. Roy. in Meded. Bot. Mus. Utrecht, 107: 128 (1951). CRASSULACEAE Cotyledon peruvianum Baker in Saund. Ref. Bot. 1: sub t. 58 (1869). Echeveria Buchtienii v. Poelln. in Fedde, Repert. Spec. Nov. 36: 193 (1934). E. chilonensis (O. Ktze.) E. Walther in Cact. & Succ. Journ. 7: 40 (1935). E. quitensis (HBK.) Lindl. in Journ. Hort. Soc. 7: 268 (1852). E. Whitei Rose in Addisonia, 10: 47, t. 344 (1925). Sedum cymatopetalum Fréd. in Act. Hort. Gotoberg. 10: 83 (1935). Tillaea connata R. & P. Fl. Peruv. 1: 70, t. 106 (1798). SAXIFRAGACEAE Escallonia aculeata O. Ktze. Rev. Gen. 3(2): 81 (1898). - ads Rusby in Mem. Torr. Bot. Club, 6: 32 (1896) = E. corymbosa. Bridgesii Rusby in Mem. Torr. Bot. Club. 6: 32 (1896). corymbosa (R. & P.) Pers. Synops. Pl. 1: 234 (1805). hypoglauca Herzog in Meded. Rijks Herbar. 27: 88 (1915). Mandonii Rusby in Mem. Torr. Bot. Club, 3(8): 25 (1893). Mandonii var. microphylla Herzog in Meded. Rijks Herbar. 27: 89 (1915). millegrana Griseb. in Goett. Abh. 24: 141 (1879). E. myrtilloides L. f. Suppl. 156 (1781). 88 E. paniculata var. acuminatissima O. Ktze. Rev. Gen. 3(2): 81 (1898). E. resinosa (R. & P.) Pers. Synops. Pl. 1: 235 (1805). Hieronymusia alchemilloides (Griseb.) Engl. in Notizbl. 7: 267 (1918). Hydrangea Bangii Engl. in Engl. & Prantl, Nat. Pflanzenfam. (ed. 2) 18A: 207 (1930) = H. tarapotensis H. scandens Poepp. ex DC. ea 4: 666 (1830). H. tarapotensis Briq. in Ann. Conserv. & Jard. Bot. Genéve, 20: 415 (1919). Phyllonoma integerrima (Turcz.) Britton in Bull. Torr. Bot. Club, 17: 11 P. ruscifolia Willd. ex Roem. & Schult. Syst. Veg. 6: 210 (1820). Ribes albifolium R, & P. Fl. Peruv. 3: 12, t. 232 (1802). R. bolivianum Jancx: in Bull. Acad. Sci. Cracov. 1905: 759. brachybotrys (Wedd.) Jancz. in Bull. Acad. Sci. Cracov. 1905: 759. glandulosum R. & P. Fl. Peruv. 3: 13, t. 233 (1802). . Pentlandii Britton fn? Rusby in Mem. Torr. Bot. Club, 3(3): 26 (1893). . punctatum R. & P. Fl. Peruv. 3:12, t. 233 (1802). sucheziense Jancz. in Bull. Acad. Sci. Cracov. 1906: 8 = R. brachybotrys. . viscosum R. & P. Fl. Peruv. 3: 13 (1802). Saxifraga cespitosa L. Sp. Pl. 404 (1753). S. cordillerarum Presl, Rel. Haenk. 2: 55 (1831). S. cordillerarum var. trigyna (Remy) Engler, Monog. Saxif. 184 (1869). S. magellanica Poir. in Lam. Encye. 6: 686 (1804). S. stylosa Remy in Ann. Sci. Nat. (ser. 3) 8: 236 (1847). ro RR PO PO BRUNELLIACEAE Brunellia boliviana Britton ex Rusby in Mem. Torr. Bot. Club, 3(3): 18 (1893) . Oliveri. B. Brittonii Rusby, Descr. S. Am. Pls. 25 (1920). B. crenata Engl. in Engl. & Prantl, Nat. Pflanzenfam. Nachtr. 184 (1897). B. Oliveri Britton in Bull. Torr. Bot. Club, 16: 160 (1889). B. rhoides Rusby in Bull. N. Y. Bot. Gard. 4: 310 (1907) = B. crenata. CUNONIACEAE basa Balbisiana var. Bangiana Cuatr. in Lloydia, 11: 202 (1949). B usby in Mem. Torr. Bot. Club, 3(3): 26 (1893). holivicas R. E. Fries in Ark. Bot. 8(8): 16, t. 11 (1909). Buchtienii Engl. in Engl. & Prantl, Nat. Pflanzenfam. (ed. 2) 18A: 252 1930). bullata Rusby in Bull. N. Y. Bot. Gard. 6: 508 (1910). cochabambensis Rusby in Mem. Torr. Bot. Club, 6:33 (1896). Cuatrecasasii Macbr. in Field Mus. Publ. Bot. 13(2): 1051 (1938). fagaroides HBK. Nov. Gen. & Spec. 6: 54, t. 524 (1823). geometrica Rusby in Mem. N. Y. Bot. Gard. 7: 248 (1927). hirtella HBK. Nov. Gen. & Spec. 6: 56 (1823). laurina HBK. Nov. Gen. & Spec. 6: 51 (1823). laxiflora Pampanini in Ann. di Bot. 2: 77 (1904). laxiflora var. polyphylla Pampanini in Ann. di Bot. 2: 78 (1904). lyrata Rusby, Descr. S. Am. Pls. 25 (1920). microphylla R. & P. Fl. Peruv. 4: t. 334 (1802). obtusifolia Rusby in Bull. N. Y. Bot. Gard. 6: 508 (1910). rhoifolia Rusby in Bull. N. Y. Bot. Gard. 4: 353 (1907). sessilifolia Rusby in Mem. Torr. Bot. Club, 6: 33 (1896). sessilifolia Rusby in Bull. N. Y. Bot. Gard. 6: 509 (1910). sorbifolia HBK. Nov. Gen. & Spec. 6: 57 (1823). PEP AAAAARAAA ARSE AAS 89 W. sorbifolia var. crenata (Presl) Cuatr. in Lloydia, 11: 204 (1949). W. spiciformis Engl. in Engl. & Prantl, Nat. Pflanzenfam. (ed. 2) 18A: 252 (1930). W. Trollii O. C. Schmidt in Fedde, Repert. Spec. Nov. 32: 97 (1933). ROSACEAE Acaena boliviana Gandog. in Bull. Soc. Bot. France, 59: 707 (1918). A. cylindrostachya R. & P. Fl. Peruv. 1: 68, t. 54 (1798). A. cylindrostachya var. nitidissima Bitter in Bibl. Bot. 17, Heft 74: 46 (1910). A. cylindrostachya var. nitidissima subvar. pusilla Bitter in Bibl. Bot. 17, Heft 74: 46 (1910). A. elongata L. Mant. 200 (1771). A. longiscapa Bitter in Bibl. Bot. 17, Heft 74: 195 (1910). A. ovalifolia R. & P. Fl. Peruv. 1: 67, t. 1 A. ovalifolia subsp. chamaephyllon var. calvescenticupula Bitter in Bibl. Bot. 17, Heft 74: 238 (1910). A. ovalifolia var. subsexjuga Bitter in Fedde, Repert. Spec. Nov. 10: 495 (1912). A. pinnata Citerne in Bull. Soc. Sci. Nat. Ouest, 7: 42 (1897). A. stricta var. gracilis Bitter in Bibl. Bot. 17, Heft 74: 36 (1910). A. stricta var. robusta Bitter in Bibl. Bot. 17, Heft 74: 36 (1910). Alchemilla aphanoides var. tripartita (R. & P.) Perry in Contrib. Gray Herb. 84: 40 (1929) = Lachemilla sp. A. appendiculata Wedd. ex Murbeck in Bot. Notis. 1915: 94 = Lachemilla diplophylla. A. hirsuta HBK. Nov. Gen. & Spec. 6: 224 (1824) — Lachemilla sp. A. pinnata forma rosulata Pilg. ex Rothm. in Trab. Mus. Nac. Cienc. Nat. 21: 33 (1935) = Lachemilla sp. Apopetalum pinnatum Pax in Fedde, Repert. Spec. Nov. 5: 226 (1908). Cotoneaster Baenitzii Pax in Fedde, Repert. Spec. Nov. 5: 226 (1908) = Hesperomeles cuneata. Eriobotrya japonica (Thunb.) Lindl. in Trans. tae Soc. 18: 102 (1822). Fragaria chiloénsis (L.) Ehrh. Beitr. 7: 26 (179 Geum boliviense Focke in Engler, Bot. Jahrb. a 540 (1906). Hesperomeles cuneata Lindl. in Bot. Reg. 23: sub t. 1956 (1837). H. ferruginea (HBK.) Benth, Pl. Hartw. 129 (1844) H. lanuginosa R. & P. ex Hook. Ic. Pl. 9: t. 846 (1852). H. incerta (Pittier) Maguire in Fieldiana, Bot. 28: 251 (1952). H. pernettyoides Wedd. Chlor. And. 2: 230 (1861). H. Weberbaueri C. K. Schneider in meee Bot. Jahrb. 42: 85 (1908). Hirtella acuminata Rusby in Mem. N. Y. Bot. Gard. 7: 248 (1927). americana Aubl. Pl. Guian. Fr. 1: 247 a bracteata Mart. & Zucc. in Abh. Akad. Miinch. 1: 384 (1832). bullata Benth. in Hook. Journ. Bot. 2: 216 (1840). Burchellii Britton in Bull. Torr. Bot. Club, 17: 10 (1890). lightioides Rusby in Bull. N. Y. Bot. Gard. 4: 350 (1907). Sprucei Benth. ex Hook. f. in Mart. Fl. Bras. 14(2): 30 (1867). stipitadenia Rusby in Mem. N. Y. Bot. Gard. 7: 249 (1927). triandra Sw. Prodr. 51 (1788). Kageneckia lanceolata R. & P. Syst. 290 (1798). Lachemilla achilleifolia (Remy) Rothm. in Fedde, Repert. Spec. Nov. 42: 169 (1937). L. andina (Perry) Rothm. in Fedde, Repert. Spec. Nov. 42: 169 (1937). L. aphanoides (Mutis ex L. f.) Rothm. in Fedde, Repert. Spec. Nov. 42: 170 (1937). Fu pp pt pt L. bipinnatifida (Perry) Rothm. in Fedde, Repert. Spec. Nov. 42: 171 (1937). L. diplophylla (Diels) Rothm. in Fedde, Repert. Spec. Nov. 42: 169 (1937). L. erodiifolia (Wedd.) Rothm. in Fedde, Repert. Spec. Nov. 42: 172 (1937). L. frigida (Wedd.) Rothm. in Kew Bull. 1938: 271. L. Mandoniana (Wedd.) Rothm. in Fedde, Repert. Spec. Nov. 42: 171 (1937). L. pectinata (HBK.) Rothm. in Fedde, Repert. Spec. Nov. 42: 171 (1937). L. pinnata (R. & P.) Rothm. in Kew Bull. 1938: 271. . ranunculoides (Perry) Rothm. in Fedde, Repert. Spec. Nov. 42: 172 (1987). Rusbyi (Perry) Rothm. in Fedde, Repert. Spec. Nov. 42: 171 (1937). L. sarmentosa (Perry) Rothm. in Fedde, Repert. Spec. Nov. 42: 171 (1937). L. Steinbachii Rothm. in Kew Bull. 1938: 271. L. vuleanica (Cham. & Schlechtd.) Rydb. in N. Am. Fl. 22: 382 (1908). L. Williamsii (Perry) Rothm. in Fedde, Repert. Spec. Nov. 42: 172 (1987). Licania Benthamii Hook. f. in Mart. Fl. Bras. 14(2): 12 (1867). L. pallida Britton in Bull. Torr. Bot. Club, 17: 9 (1890). Parinarium laxiflorum Ducke in Arch. Jard. Bot. Rio Janeiro, 3: 44 (1922). i ). Polylepis Besseri Hieron. in Engler, Bot. Jahrb. 21: 312 ( P. Besseri var. abbreviata Bitter in Engler, Bot. Jahrb. 45: 628 (1911). P. Besseri subsp. longipedicellata Bitter in Engler, Bot. Jahrb. 45: 629 (1911). P. crista-galli Bitter in Engler, Bot. Jahrb. 45: 633 (1911). P. crista-galli var. longiracemosa Bitter in Engler, Bot. Jahrb. 45: 634 (1911). P. Hieronymi Pilg. in Engler, Bot. Jahrb. 37: 534 (1906). P. incana subsp. brachypoda Bitter in Engler, Bot. Jahrb. 45: 644 (1911). P. incana subsp. incarum Bitter in Engler, Bot. Jahrb. 45: 643 (1911). P. incana es subtusalbida Bitter in to Bot. Jahrb. 45: 640 (1911). P. racemosa Wedd. Chlor. And. 2: 238 (186 P. racemosa var. lanata O. Ktze. Rev. i 3(2): 77 (1898). P. tarapacana var. sajamensis Bitter in Engler, 0 Jahrb. 45: 654 (1911). P. tomentella Wedd. Chlor. And. 2: 237, t. 78 (1861 P. tomentella subsp. dentatialata Bitter in Engler, a Jahrb. 45: 650 (1911). P. triacontandra Bitter in Engler, Bot. Jahrb. 45: 630 (1911). Potentilla lignipes Rusby in Bull. N. Y. Bot. Gard. 4: 352 (1907). Poterium Sanguisorba L. Sp. Pl. 994 (1753). Prunus Brittoniana Rusby in Mem. Torr. Bot. Club, — 24 (1893). . guanaiensis Rusby in Mem. Torr. Bot. Club, 6: 31 (18 P. guanaiensis var. micradenia Koehne in Engler, Bot. meee 52: 318 (1915). P. oleifolia var. Bangii Koehne in Engler, Bot. Jahrb. 52: 318 (1915). ¥, Pearcei Rusby in Mem. Torr. Bot. Club, 6: 30 (1896) tes a var. subintegra Koehne in Engler, Bot. ate 52: 316 (1915). Rosa repellens Rusby in Bull. N. Y. Bot. Gard. 6: 509 (1910). Rubus adenothallus Focke in Meded. Rijks Herbar. 19: 56 (1913). aenigmaticus Focke in Meded. Rijks Herbar. 19: 55 (1913). betonicifolius Focke in Bibl. Bot. 72: 33 (1910). bogotensis HBK. Nov. Gen. & Spec. 6: 220 (1824). boliviensis Focke in Abh. Nat. Ges. Bremen, 4: 158 (1875). Briareus Focke in Fedde, Repert. Spec. Nov. 9: 235 (1911). Briareus subsp. Herzogii (Focke) Focke in Bibl. Bot. 19, Heft 83: 56 (1914). . Buchtienii Focke in Fedde, Repert. Spec. Nov. 9: 237 (1911) bullatus Rusby in Bull. N. Y. Bot. Gard. 4: 351 (1907). conchyliatus Focke in Meded. Rijks Herbar. 19: 54 (1913). glaucus Benth. Pl. Hartw. 173 (1845). Holtonii O. Ktze. Rev. Gen. 3(2): 78 (1898). R. megalococcus Focke in Abh. Nat. Ges. Bremen, 4: 157 (1875). PAAR AP PA AAD R. nubigenus HBK. Nov. Gen. & Spec. 6: 220 (1824). R. ostrinus Focke in Bibl. Bot. 19, Heft. 83: 21 (1914). R. robustus Presl, Epim. Bot. 196 (1849) R. roseus Poir. in Lam. Encyc. 6: 245 (1804). R. roseus var. santarosensis (O. Ktze.) Macbr. in Field Mus. Publ. Bot. 8: 118 (1930). R. Rusbyi Britton in Bull. Torr. Bot. Club, 17: 10 (1890). Tetraglochin Ameghinoi (Speg.) Speg. in Anal. Mus. Buenos Aires, 7: 283 1902 T. cristatum (Britton) Rothm. in Darwiniana, 3: 433 (1939). T. strictum Poepp. Fragm. 26 (1833). T. Tragacantha Rothm. in Darwiniana, 3: 434 (1989). CONNARACEAE Connarus fulvus Planch. in Linnaea, 28: 434 (1850). C. Martii Schellenb. in Candollea, 2: 116 (1925). C. Patrisii (DC.) Planch. in Linnaea, 23: 432 (1850). Rourea Bakerana Britton in Bull. Toe Bot. Club, 16: 192 (1889) = Cnesti- dium Bakeranum [ Meliaceae]. camptoneura Radlk. in Sitzb. Bayer. Akad. Wiss. Miinch. 14: 375 (1886). glabra var. — Britton in Bull. Torr. Bot. Club, 16: 192 (1889) = R. campton laxiflora aay once S. Am. Pls. 28 (1920). puberula Baker in Mart. Fl. Bras. 14(2): 179 (1871). Sprucei Schellenb. in Engler, Pflanzenr. IV, 127: 205 (1938). eee NA LEGUMINOSAE Acacia albicorticata Burkart in Darwiniana, 7: 504 (1947). - ampeloclada Rusby in Mem. N. Y. Bot. Gard. 7: 256 (1927). boliviana Rusby in Bull. N. Y. Bot. Gard. 4: 348 (1907). bonariensis Gill. ex Hook. Bot. Misc. 3: 207 (1832). Farnesiana (L.) Willd. Sp. Pl. 4(2): 1083 (1806). Feddeana Harms in Fedde, Repert. Spec. Nov. 16: 450 (1920). Fiebrigii Harms in Fedde, Repert. Spec. Nov. 16: 351 (1920) = A. Fedde- DPD PP > ana. furcata Gill in Hook. Bot. Misc. 3: 206 (1832). lutea (Mill.) Britton in Bull. Torr. Bot. Club, 16: 327 (1889). Michelii Rusby in Mem. Torr. Bot. Club, 6: 28 (1896). paniculata Willd. Sp. Pl. 4(2): 1074 (1806). paraguariensis (D. Parodi) Burkart in Darwiniana, 10: 26 (1952). pedicellata Benth. in Hook. Lond. Journ. Bot. 1: 522 (1842). praecox Griseb. in Goett. Abh. 19: 136 (1874). riparia HBK. Nov. Gen. & Spec. 6: 276 (1824). riparia var. angustifolia O. Ktze. Rev. Gen. 3(2): 47 (1898). riparia var. media O. Ktze. Rev. Gen. 3(2): 47 (1898). rynchocarpa Rusby in Bull. N. Y. Bot. Gard. 8: 90 (1912). rurrenabaqueana Rusby in Mem. N. Y. Bot. Gard. 7: 255 (1927). esmia amblysepala Solms-Laub. in Bot. Zeit. 65(Abt. 1): 134 (1907). miraflorensis Remy in Ann. Sci. Nat. (ser. 3) 6: 357 (1846). muricata var. dentata (Lag.) Benth. in Mart. Fl. Bras. 15(1): 54 (1859). Nordenskioeldii (R. E. Fries) Hicken ex Cardenas, Pl. Potosinae, 13 (1932). patancana Ulbr. in Engler, Bot. Jahrb. 387: 554 (1906). pinifolia Gill. in Hook. Bot. Misc. 3: 192 (1832). A. polyacantha Wedd. Chlor. And. 2: 265 (1861). {AGRI SHS aR: 92 A. rupicola Wedd. Chlor. And. 2: 266, t. 79 bis (1861). A. Schickendantzii Griseb. in Goett. Abh. 24: 104 (1879). A. spinosissima Meyen, Reise, 2: 27 (1835).. Aeschynomene americana var. glandulosa (Poir.) Rudd in Contrib. U. S. Nat. Herb. 32: 26 (1955). Ae. apoloana Rusby in Bull. N. Y. Bot. Gard. 6: 511 (1910) = Ae. falcata. Ae. brasiliana (/Poir.) DC. Prodr. 2: 322 (1825). Ae. denticulata Rudd in Contrib. U. S. Nat. Herb. 32: 69 (1955). Ae. elegans Schlechtd. & Cham. in Linnaea, 5: 583 (1830). Ae. evenia Wright in Sauvalle in Anal. Acad. Cienc. Habana, 5: 334 (1868). Ae. falcata (Poir.) DC. Prodr. 2: 322 (1825). Ae. fluminensis Vell. Fl. Flum. 310 (1825). Ae. paniculata Willd. ex Vog. in Linnaea, 12: 95 (1838). Ae. parviflora Micheli in Kjobenh. Vid. Medd. 1875: 66. Ae. pratensis var. caribaea Rudd in Contrib. U. S. Nat. Herb. 32: 47 (1955). Ae. rudis Benth. Pl. Hartw. 116 (1843) Ae. sensitiva Sw. Prodr. 107 (1788). Affonsea trigyna (Rusby) Burkart in Darwiniana, 7: 513 (1947). Amburana acreana (Ducke) A. C. Smith in Trop. Woods, 62: 30 (1940). A. cearensis (Allem.) A. C. Smith in Trop. Woods, 62: 30 (1940). Am geay ovale Rusby in Mem. N. Y. Bot. Gard. 7: 265 (1927) = Dalbergia sp Amicie AR Harms ex O. Ktze. Rev. Gen. 3(2): 49 (1898). A. Lobbiana Benth. ex Rusby in Mem. Torr. Bot. Club, 3(3): 20 (1893). A. medicaginea Griseb. in Goett. Abh. 24: 105 (1879). A. micrantha Harms ex O. Ktze. Rev. Gen. 3(2): 49 (1898). A. parvula Rusby in Mem. Torr. Bot. Club, 6: 23 (1896). Apurimacia Michelii (Rusby) Harms in Fedde, Repert. Spec. Nov. 19: 10 923). Arachis hypogaea L. Sp. Pl. 741 (1753). A. prostrata Benth. in Trans. Linn. Soc. 18: 159 (1839). Astragalus arequipensis Vog. in Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 17 3 (1843). A. bolivianus Phil. Cat. Pl. Itin. Tarapaca, 15 (1891) — A. arequipensis. . capitellus Britton in Bull. Torr. Bot. Club, 16: 260 (1889) — A. micran- thellus colliontns Rusby in Mem. Torr. Bot. Club, 3(3): 19 (1893). == A. minimus. erymophilus I. M. Johnst. in Journ. Arnold Arb. 28: 400 (1947). cryptanthus Wedd. Chlor. And. 2: 259 (1861). deminutivus I. M. Johnst. in Journ. Arnold Arb. 28: 406 (1947). flavocreatus I. M. Johnst. in Journ. Arnold Arb. 28: 405 (1947). Garbancillo Cav. Ic. 1: 59, t. 85 (1791). gracilis Romero in Bol. Direc. Nac. Estad. y Estud. Geogr. La Paz, sec. Epoc. III, nos. 31-33: 45 (1920). Herzogti Ulbr. in Meded. Rijks Herbar. 27: 53 (1915) — A. uniflorus. Hieronymi Ulbr. in Engler, Bot. Jahrb. 37: 418 (1906) — A. tarijensis. hypsogenus I. M. Johnst. in Journ. Arnold Arb. 28: 399 (1947). Mandonii Rusby in Mem. Torr. Bot. Club, 3(3): 19 (1893) — A. Garban- pa cillo. micranthellus Wedd. Chlor. And. 2: 262 (1861). minimus Vog. in Nov. Act. Acad. Caes. ratiew 19, Suppl. 1: 18 (1843). minutissimus Wedd. Chlor. And. 2: 257 (186 modestus Wedd. Chlor. And. 2: 262 (1861) = _ Weddellianus. Orbignyanus Wedd. Chlor. And. 2: 260 (1861) — A. arequipensis. patancanus Ulbr. in Engler, Bot. Jahrb. 37: 417 (1906) = A. micran- thellus. SehP PP BES e Peer eS peruvianus Vog. in Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 18 (1843). pusillus Vog. in Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 19 (1848). Reichei Speg. in Anal. Mus. Nac. Buenos Aires (ser. 2), 4: 264 (1902). sinocarpus Rusby in Mem. Torr. Bot. Club, 3(3): 19 (1893) — A. arequip- ensis. tarijensis Wedd. Chlor. And. 2: 262 (1861). uniflorus (Dombey) DC. Astrag. 243 (1802). Urbanianus Ulbr. in Engler, Bot. Jahrb. 37: 422 (1906) Weddellianus (O. Ktze.) I. M. Johnst. in Journ. Arnold Azki 28: 395 (1947). Ateleia guaraya Herzog in Fedde, Repert. Spec. Nov. 7: 55 (1909). Barbiera pinnata (Pers.) Baill. Hist. Pl. 2: 263 (1870). uhinia acreana Harms in Notizbl. 6: 307 (1915). calliandroides Rusby in Bull. N. Y. Bot. Gard. 8: 92 (1912). Conwayi Rusby in Bull. N. Y. Bot. Gard. 8: 92 mae humilis Rusby in Mem. N. Y. Bot. Gard. 7: 256 (1927). Langsdorffiana Bong. in Mém. Acad. St. Pétersb. (ser. 4) 6: 109 (1836). longipetala Walp. Repert. 1: 852 (1842). inermis (Cav.) Pers. Synops. Pl. 1: 455 (1805). Rusbyi Britton in Bull. Torr. Bot. Club, 16: 326 (1889). Straussiana Harms in Notizbl. 6: 308 (1915). tumupasensis Rusby in Bull. N. Y. Bot. Gard. 8: 93 (1912). vulpina Rusby in Mem. N. Y. Bot. Gard. 7: 257 (1927). Bradburya cologanoides Rusby in Bull. N. Y. Bot. Gard. 6: 515 (1910) = osema sp. ? Brongniartia Ulbrichiana Harms in Engler, Bot. Jahrb. 42: 94 (1908). Caesalpinia argentina Burkart in Rev. Argent. Agron. 3: 105 (1936). Bangii Rusby in Mem. Torr. Bot. Club, 3(3): 22 (1893). coluteifolia Griseb. in Goett. Abh. 24: 111 (1879). coulterioides Griseb. in Goett. Abh. 24: 113 (1879). fimbriata Tul. in Arch. Mus. Paris, 4: 145 (1845). Fisheriana Rusby in Mem. Torr. Bot. Club, jh 23 (1898). floribunda Tul. in Arch. Mus. Paris, 4: 140 (1845). Herzogii Harms in Meded. Rijks Herbar. 27: re Jae gale = C. Stuckertii. mimosifolia Griseb. in Goett. Abh. 19: 128 (1874 pulcherrima (L.) Swartz, Obs. 166 (1791). rosulata Rusby in Mem. Torr. Bot. Club, 3(3): 23 (1893). oe >>> > bp >> . anaaagaaaaa Calliandra boliviana Britton in Bull. Torr. Bot. Club, 16: 327 (1889). C. formosa (Kunth) Benth. in Hook Lond. Journ. Bot. 3: 98 (1844). C. inaequilatera Rusby in Mem. Torr. Bot. Club, 6: 28 (1896). C. portoricensis (Jacq.) Benth. in Hook. Lond. Journ. Bot. 3: 99 (1844). C. stricta Rusby in Mem. N. Y. Bot. Gard. 7: 255 (1927). Calopogonium caeruleum (Benth.) Britton in Bull. Torr. Bot. Club, 16: 262 C. galactioides Benth. ex Hemsl. in Biol. Centr. Am. Bot. 1: 301 (1880). Canavalia boliviana Piper in Contrib. U. S. Nat. Herb. 20: 573 (1925). C. brasiliensis Mart. ex Benth. in Ann. Naturh. Hofmus. Wien, 2: 135 (1838). C. ensiformis (L.) DC. Prodr. 2: 404 (1825). C. lasiocalyx O. Ktze. Rev. Gen. 3(2 : 55 (1898). C. obtusifolia DC. Prodr. 2: 404 (1825). C. villosa Benth. in Ann. Naturh. Hofmus. Wien, 2: 135 (1838). Cardenasia setacea Rusby in Mem. N. Y. Bot. Gard. 7: 259, fig. 3 (1927). Cascaronia astragalina Griseb. in Goett. Abh. 24: 100 (1879). Cassia Absus L. Sp. Pl. 376 (1753). C. acinicarpa Rusby in Bull. N. Y. Bot. Gard. 4: 311 (1907). © r= affinis Benth. in Mart. Fl. Bras. 15(2): 98 (1870). atomaria L. Mant. 68 (1767). bacillaria L. f. Suppl. 231 (1781). birostris Domb. ex Vog. Syn. Cass. (18387). brachypoda Benth. in aoe FI. 15(2): 172 (1870). Chamaecrista L. Sp. ‘Pl. 379 (1753). Chamaecrista var. brasiliensis Vog. in Mart. Fl. Bras. 15(2): 172, t. 45 (1870). chloroclada Harms in Meded. Rijks Herbar. 27: 37 (1915). cochabambae Herzog in Fedde, Repert. Spec. Nov. 7: 55 (1909). conjugata R. & P. ex Benth. in Trans. Linn. Soc. 27: 540 (1871). corymbosa Lam. Encyc. 1: 644 (1785). crassiramea Benth. in Hook. Ic. Pl. 11: 50 (1870). flavicoma HBK. Nov. Gen. & Spec. 6: 366 (1824). flexuosa L. Sp. Pl. 379 (1753). fruticosa Mill. Gard. Dict. (ed. 8) no. 10 (1768). Herzogii Harms in Meded. Rijks Herbar. 27: 37 (1915). hirsuta L. Sp. Pl. 378 (1753). Hookeriana Gill. ex Hook. & Arn. in Hook. Bot. Misc. 3: 210 (1832). incarnata Pav. ex Benth. in Trans. Linn. Soc. 27: 545 (1871). latopetiolata Domb. ex Vog. Syn. Cass. 29 (1837). Mandonii Benth. in Trans. Linn. Soc. 27: 540 (1871). Morongii Britton in Ann. N. Y. Acad. Sci. 7: 91 (1892). multijuga Rich. in Act. Soc. ony Nat. Paris, 1: 108 (1792). occidentalis L. Sp. Pl. 377 (175 patellaria DC. in Collad. Hist. oor 125, t. 16 (1816). patellaria var. longifolia Benth. in Trans. Linn. Soc. 27: 579 (1871). pazensis Rusby in Bull. N. Y. Bot. Gard. 8: 94 (1912). Pearcei Benth. in Trans. Linn. Soc. 27: 552 (1871). pendula Humb. & Bonpl. ex Willd. Enum. Hort. Berol. 440 (1809). Pennelliana Amsh. in Meded. Bot. Mus. Utrecht, 52: 28 (1939). pilifera Vog. Syn. Cass. 23 (1837). racemosa Mill. Gard. Dict. (ed. 8) no. 19 (1768). rotundifolia Pers. Synops. Pl. 1: 456 (1805). serpens L. Syst. (ed. 10) 1018 (1759). spectabilis DC. Cat. Hort. Monsp. 90 (1813). splendida Vog. Syn. Cass. 17 (1837). subelliptica Rusby in Bull. N. Y. Bot. Gard. 8: 94 (1912). silvestris Vell. Fl. Flum. 169, t. 78 (1825). tomentosa L. f. Suppl. 231 (1781). tomentosa var. paucijuga O. Ktze. Rev. Gen. 3(2): 56 (1898). Tora L. Sp. Pl. 376 (1753). trachypus Mart. ex Benth. in Mart. Fl. Bras. 15(2): 122 (1870). versicolor Meyen ex Vog. Syn. Cass. 29 (1837). Centrolobium minus Presl, Bot. Bemerk. 61 (1844). Centrosema brasilianum Benth. in Ann. Naturh. Hofmus. Wien, 2: 118 (1838). C. pascuorum Mart. ex Benth. in Ann. Naturh. Hofmus. Wien, 2: 120 (1838). C. Plumieri (Turp.) Benth. in Mart. Fl. Bras. 15(1): 127 (1859). C. pubescens Benth. in Mart. Fl. Bras. 15(1): 131, t. 34 (1859). C. sagittatum (Humb. & Bonpl.) Brandg. in Zoé, 5: 202 (1905). C. virginianum (L.) Benth. in Mart. Fl. Bras. 15(1): 1382 (1859). Cercidium andicola Griseb. in Goett. Abh. 24: 114 (1879). C. praecox (R. & P.) Harms in Engler, Bot. Jahrb. 42: 91 (1908). Chaetocalyx brasiliensis (Vog.) Benth. in Mart. Fl. Bras. 15(1): 75, t. 18 (1859). Clitoria nervosa Herzog in Fedde, Repert. Spec. Nov. 7: 56 (1909). C. Poitaei DC. Prodr. 2: 234 (1825). Ae a lipek ee so Vee) | a ORE BR ee ARARAARAA Collaea speciosa DC. in Mém. Lég. 6: 245 (1825). Cologania ovalifolia HBK. Nov. Gen. & Spec. 6: 412 (1824). C. pulchella HBK. Nov. Gen. & Spec. 6: 413 (1824). Colutea frutescens L. Sp. Pl. 723 (1753). Copaiba Langsdorffii (Desf.) O. Ktze. Rev. Gen. 1: 172 (1891). Copaifera paupera (Herzog) Dwyer in Brittonia, 7: 169 (1950). C. reticulata Ducke in Arch. Jard. Bot. Rio Janeiro, 1: 22 (1915). Coursetia boliviana Britton in Bull .Torr. Bot. Club, 16: 260 (1889). C. brachyrhachis Harms in Meded. Rijks Herbar. 27: 52 (1915 C. grandiflora Benth. ex Oerst. in Kjobenh. Vid. Medd. 1853: 10 (1854). Cracca benensis Rusby in Mem. N. Y. Bot. Gard. 7: 262 (1927 C. heterantha (Griseb.) Harms in Fedde, Repert. Spec. Nov. 18: 237 (1922). C. Kuntzet Harms in O. Ktze. Rev. Gen. 3(2): 69 (1898) — C. heterantha. C. ochroleuca (Pers.) Benth. ex Oerst. in Kjobenh. Vid. Medd. 1853: 9 (1854). Cratylia nutans Herzog in Fedde, Repert. Spec. Nov. 7: 56 (1909 Crotalaria anagyroides HBK. Nov. Gen. & Spec. 6: 404 (1824). incana L. Sp. Pl. 716 (1753). nitens HBK. Nov. Gen. & Spec. 6: 399 (1824). Pohliana Benth. in Ann, Nat. Hist. 3: 428 (1839). pterocaula Desv. in Journ. Bot. 3: 76 (1814). pumila Ortega in Hort. Bot. Matrit. Dec. 2: 23 (1797). sagittalis L. Sp. Pl. 714 (1753). C. stipularia Desv. in Journ. Bot. 3: 76 (1814). Dalbergia frutescens (Vell.) Britton in Bull. Torr. Bot. Club, 16: 324 (1889). D. nitida (Radlk.) Ducke ex Hoehne in FI. Brasil. 25 (pt. 3, fase. 4): 22 (1941). D. oxphylla Harms ex O. Ktze. Rev. Gen. 3(2): 49 (1898), in synon. DPD. Spruceana Benth. in Mart. Fl. Bras. 15(1): 223 (1862). Dalea alopecuroides Willd. Sp. Pl. 3: 1836 (1799). boliviana Britton in Bull. Torr. Bot. Club, 16: 259 (1889). calliantha Ulbr. in Fedde, Repert. Spec. Nov. 2: 11 (1906). eosina (Macbr.) Macbr. in Candollea, 7: 222 (19387). Hofstenii R. E. Fries in Nov. Act. Soc. Sci. Upsal. a. 4) 1(1): 132 (1905). Kuntzei Harms ex O. Ktze. Rev. Gen. 3(2): 59 (1898). microphylla HBK. Nov. Gen. & Spec. 6: 482 (1824). pazensis Rusby in Mem. Torr. Bot. Club 3(3): 18 (1893). peruviana (Macbr.) Macbr. in Candollea, 7: 223 (1937). retusifolia Harms ex O. Ktze. Rev. Gen. 3(2): 59 (1898). . rubricaulis Ulbr. in Meded. Rijks Herbar. 27: 50 (1915). A — cariensis Harms ex O. Ktze. Rev. Gen. 3(2): 59 (1898 = D. bolivi- QAARQAAAHA SOSSSSosyss Derris Ladodiiea (Mia. ) Macbr. in Field Mus. Publ. Bot. ~~ 262 (1943). D. negrensis Benth. in Mart. Fl. Bras. 15(1): 289 (186 D. Pterocarpus (DC.) Killip in Journ. Wash. Acad. oe fo 360 (1936). D. Steinbachii Harms in Notizbl. 10: 346 (1928). Desmanthus depressus Humb. & Bonpl. ex Willd. a ey 4: 1046 (1806). Desmodium adscendens (Sw.) DC. Prodr. 2: 332 (1825). albiflorum Salzm. in Benth. in Mart. Fl. Bras. 15(1): 99 (1859). axillare (Sw.) DC. Prodr. 2: 333 (1825). barbatum (L.) Benth. in Mart. Fl. Bras. 15(1): 95 (1859). Bridgesii (Schindl.) Burkart in Darwiniana, 3: 194 (1939). cajanifolium (HBK.) DC. Prodr. 2: 331 (1825). canum (Gmel.) Schinz & Thell. in Mém. Soc. Neuchatel. Sci. Nat. 5: 371 (1913). cuneatum Hook. & Arn. in Hook. Bot. Misc. 3: 195 (1832). distortum (Aubl.) Macbr. in Field Mus. Publ. Bot. 8: 101 (1930). Hassleri (Schindl.) Burkart in Darwiniana, 3: 211 (1939). POS Sees © for) longiarticulatum (Rusby) Burkart in Darwiniana, 3: 198 (1939). Mandonii Britton in Bull. Torr. Bot. Club, 16: 261 (1889). molliculum (HBK.) DC. Prodr. 2: 331 (1825). neo-mexicanum A. Gray, Pl. Wright. 1: a (1852). pachyrhizum Vog. in Linnaea, 12: 97 (1838). sclerophyllum ae in Mart. Fl. Bras. sees 102 (1859). tortuosum (Sw.) DC. Prodr. 2: 3382 (18 uncinatum (Jacq.) a Prodr. 2: 331 ties yungasense Britton in Bull. Torr. Bot. Club, 16: 261 (1889). Dioclea argentea Desv. in Ann. Sci. Nat. (ser. 1) 9: 420 (1826). lasiocarpa Mart. in Benth. in Mart. Fl. Bras. 15(1): 166 (1859). lasiophylla Mart. ex Benth. in Ann. Naturh. Hofmus. Wien, 2: 134 (1838). ornithorhyncha Rusby in Phytologia, 1: 56 (1984). pauciflora Rusby in Mem. Torr. Bot. Club, 6: 25 (1896) — D. argentea. reflexa Hook. f. Niger Fl. 306 (1849 virgata (Rich.) Amsh. in Meded. Bot. Mus. Utrecht, 52: 69 (1939). Dolichos Lablab L. Sp. Pl. 725 (17538). Dolicholus ovatus Rusby in Bull. N. Y. Bot. Gard. 4: 346 (1907). D. phaseoloides (Sw.) in Bull. N. Y. Bot. Gard. 4: 346 (1907). Drepanocarpus lunatus (L. f.) G. F. W. Meyer, Primit Fl. Esseq. 238 (1818). Enterolobium Timbouva Mart. in Flora, 20, II Beibl.: 128 (1837). Eriosema canescens Rusby in Bull. N. Y. Bot. Gard. 4: 346 (1907). Conwayi Rusby in Bull. N. Y. Bot. Gard. 8: 90 (1912): crinitum (HBK.) E. Meyer, Comm. Pl. Afr. Austr. 128 (1836). fusiforme Rusby in Bull. N. Y. Bot. Gard. 6: 516 (1910). rufum (HBK.) E. Meyer, Comm. Pl. Afr. Austr. 128 (1836). rufum var. flexuosum O. Ktze. Rev. Gen. 3(2): 63 (1898). simplicifolium (HBK.) Walp. a te 2: 902 (1848). rythrina crista-galli L. Mant. 99 (176 Dominguezii Hassl. in Physis, 6: aot (1922). edulis Triana ex M. Micheli in Journ. Bot. 6: 145 (1892). faleata Benth. in Mart. Fl. Bras. 15(1): 172 (1859). flamma Herzog in Fedde, Repert. Spec. Nov. 7: 57 (1909). glauca Willd. in Ges. Nat. Freunde Berl. Neue Schr. 3: 428 (1801). Poeppigiana (Walp.) O. F. Cook in U. S. Dept. Agric. Div. Bot. Bull. 25: 57 (1901). rubrinervia HBK. Nov. Gen. & Spec. 6: 434 (1824). similis Krukoff in Brittonia, 3: 271 (1989). Ulei Harms in Verh. Bot Ver. Brandenb. 48: 172 (1907). Ferreirea spectabilis Allem. in Trab. Soc. Vell. 26 (1845). Fiebrigiella gracilis Harms in Engler, Bot. Jahrb. 42: 96 (1908). Galactia glaucescens HBK. Nov. Gen. & Spec. 6: 431 (1824). G. Jussiaeana Kunth, Mimos. 196, t. 55 (1824 G. montana Britton in Bull. Torr. Bot. Club, 16: 324 (1889). G. speciosa (DC.) Britton in Bull. Torr. Bot. Club, 16: 262 (1889). G. tenuiflora (Willd.) Wight & Arn. Prodr. 1: 206 (1834). Geoffroya striata (Willd.) Morong in Ann. N. Y. Acad. Sci. 7: 87 (1892). G. superba Humb. & Bonpl. Pl. Aequin. 2: 69, t. 100 (1809). Gleditsia amorphoides (Griseb.) Taub. in Ber. Deut. Bot. Ges. 10: 688, t. 32 SOSSSesyy Syd OUY te fed bt bed bd bt bo td bd Dt bs bd Gourliaea decorticans Gill. ex Hook. in Hook. Bot. Misc. 3: 208 (1833). Hoffmannseggia gracilis Hook. & Arn. in Hook. Bot. Misc. 3: 209 (18338). Hoita hirsuta Rusby in Mem. N. Y. Bot. Gard. 7: 261 (1927). H. versicolor Rusby Mem. N. Y. Bot. Gard. 7: 260 (1927) = Psoralea lasi- ostachys var. pot Indigofera Anil L. hue. “272 (1771). 97 asperifolia Bong. ex Benth. in Ann. Nat. Hist. 3: 431 (1839). lespedezioides HBK. Nov. Gen. & Spec. 6: 457 (1824). microcarpa Desv. in Journ. Bot. 8: 79 (1814). sabulicola Benth. in Mart. Fl. Bras. 15(1): 40 (1859). suffruticosa Mill. Gard. Dict. (ed. 8) no. 2 (1768). nga acreana Harms in Notizbl. 6: 298 (1915). adenophylla Pittier in Contrib. U. S. Nat. Herb. 18: 210 (1916). aggregata G. Don, Gen. Hist. 2: 391 (1832). apiculata Rusby in Mem. N. Y. Bot. Gard. 7: 250 (1927). Bangii Harms in Fedde, Repert. Spec. Nov. 18: 525 (1915). boliviana Britton in Bull. Torr. Bot. Club, 17: 9 (1890). Bourgonii (Aubl.) DC. Prodr. 2: 434 (1825). I. calophylla Harms in Notizbl. 6: 298 (1915). canaminensis Rusby in Mem. N. Y. Bot. Gard. rh 249 (1927). chartacea Poepp. & Endl. Nov. Gen. 3: 79 (1845). chrysotricha Pittier in Contrib. U. S. Nat. Herb. 18: 202 (1916) = I. hirsu- issima. ellipsoidea Rusby in Mem. N. Y. Bot. Gard. 7: 252 (1927). expansa Rusby in Bull. N. Y. Bot. Gard. 8: 90 (1912) =I. velutina. heterophylla Willd. Sp. Pl. 4(2): 1020 (1806). hirsutissima Rusby in Bull. N. Y. Bot. Gard. 4: 349 7). macrophylla Humb. & Bonpl. ex Willd. Sp. Pl. 4(2): saaee (1806). mapiriensis Pittier in Contrib. U. S. Nat. Herb. 18: 174 (1916). marginata Willd. Sp. Pl. 4(2): 1015 (1806). Mathewsiana Benth. in Hook. Lond. Journ. Bot. 4: 594 (1845). yriocephala Pittier in Contrib. U. S. Nat. Herb. 18: 184 (1916). nobilis Willd. Enum. 1047 (1809). pallida Rusby in Mem. Torr. Bot. Club, 6: 30 (1896). peltadenia Harms in Verh. Bot. Ver. Brandenb. 48: 160 (1906). punctata Willd. Sp. Pl. 4(2): 1016 (1806). radiata Rusby in Mem. N. Y. Bot. Gard. 7: 252 (1927). . rugosa Rusby in Bull. N. Y. Bot. Gard. 4: 350 (1907). I. Ruiziana G. Don, Gen. Hist. 2: 391 (1832). Rusbyi Pittier in Contrib. U. S. Nat. Herb. 18: 179 (1916). Steinbachii Harms in Notizbl. 9: 1037 (1926). stenopoda Pittier in Contrib. U. S. Nat. Herb. 18: 192 (1916). stipularis DC. Mém. Lég. 12: 440 (1825). striata Benth. in Hook. Lond. Journ. Bot. 4: 608 (1845). strigillosa Spruce ex Benth. in Trans. Linn. Soc. 30: 612 (1875). tenuifolia Salzm. ex Benth. in Hook. Lond. Journ. Bot. 4: 596 (1845). Thibaudiana DC. Prodr. ” 434 (1825). . tomentosa Benth. in Trans. Linn. Soc. 30: 609 (1875). . velutina Willd. Sp. PI. 4(2): 1014 (1806). rameria triandra R. & P. Fl. Peruv. 1: 61, t. 93 (1798). K. triandra var. Humboldtiana Chod. in Arch. Sci. Phys. & Nat. (ser. 3) 24: 498 (1890 Lathyrus tiagellanicus Lam. Encye. 2: 708 (1785). L. magellanicus var. tucumanensis Burkart in Rev. Fac. Agron. & Vet. 8: 108 (1935). L. pubescens Hook. & Arn. Bot. Beechey Voy. 21 (1830). L. pubescens var. monticola Burkart in Darwiniana, 6: 17 (1942). L. tropicalandinus Burkart in Darwiniana, 6: 16 (1942). Leucaena boliviana Rusby in Bull. N. Y. Bot. Gard. 8: 91 (1912). Lonchocarpus boliviensis Pittier in Contrib. U. S. Nat. Herb. 20: 93 (1917). L. macrocarpus Benth. in Journ. Linn. Soc. 4: Suppl. 91 (1860). L. pluvialis Rusby in Mem. N. Y. Bot. Gard. 7: 268 (1927). ra en i bt ps tome yet ge a el 9 ato oR eel pee | sll ge ga ms Bel ae ge on a la ns ele ed dv el 98 L. velutinus Benth. in Seem. Bot. Voy. Herald, 111 (1853). Lupinus alaristatus C. P. Smith, Spec. Lup. 461 (1945). altimontanus C. P. Smith, Spec. Lup. 152 (1940). altiplani C. P. Smith, Spec. Lup. 455 (1945). Asplundianus C. P. Smith, Spec. Lup. 454 Spemes Bandelierae C. P. Smith, ped. Lup. 214 (1940). Bangii Rusby in Mem. Torr. Bot. Club, 3(3): 17 (1893). bogotensis Benth. Pl. Hartw. 168 (18 1845 5). bolivianus Rusby ex C. P. Smith, Spec. Lup. 457 (1945). breviscapus Ulbr. in Meded. Rijks Herbar. 27: 46 (1915). Buchtienii Rusby in Phytologia, 1: 56 (1934). Cardenasianus C. P. Smith, Spec. Lup. 462 (1945). celsimontanus C. P. Smith, Spec. Lup. 153 (1940). chilensis C. P. Smith, Spec. Lup. 197 (1940). chrysanthus Ulbr. in Engler, Bot. Jahrb. 37: 543 (1906). cuspidatus Rusby in Bull. N. Y. Bot. Gard. 4: 342 (1907). erectifolius C. P. Smith, Spec. Lup. 351 (1944). Fiebrigianus Ulbr. in Engler, Bot. Jahrb. 37: 545 (1906). Guggenheimianus Rusby in Mem. N. Y. Bot. Gard. 7: 260 (1927). Herzogii Ulbr. in Meded. Rijks Herbar. 27: 44 (1915). hirsutus L. Sp. Pl. 721 (1753). humifusus Benth. Pl. Hartw. 169 (1845). imminutus C. P. Smith, Spec. ae 462 (194 macrostachys Rusby in Bull. N. Y. Bot. me vr 342 (1907) = L. bolivi- anus. i remnag var. sessiliflorus Rusby in Bull. N. Y. Bot. Gard. 4: 343 (19077 == Rint h Seay C. P. Smith, Spec. Lup. 464 (1945). microphyllus Desr. in Lam. Encyc. 3: 625 (1789). montanus var. austrovulcanicus C. P. Smith, Spec. Lup. 90 (1938). mutabilis Sweet, Brit. Fl. Gard. (ser. 1) 2: t. 130 (1825) nubilorum C. P. Smith, Spec. Lup. 458 (1945). Otto-Buchtienii C. P. Smith, Spec. Lup. 454 (1945). Otto-Kuntzeanus C. P. Smith, Spec. Lup. 453 (1945). paniculatus Desr. in Lam. Encyc. 3: 625 (1789). paniculatus var. argenteus Wedd. Chlor. And. 2: 252 (1861). Pearceanus C. P. Smith, Spec. Lup. 460 (1945). perissophytus C. P. Smith, Spec. Lup. ons Sayed podpoénsis C. P. Smith, Spec. Lup. 459 (19 pulvinaris Ulbr. in Engler, Bot. Jahrb. -. "te (1906). purosericeus C. P. Smith, Spec. Lup. 451 (1945). pycnostachys C. P. Smith, Spec. Lup. 248 (1941). ramosissimus Benth. Pl. Hartw. 170 (1845). Rusbyanus C. P. Smith, Spec. Lup. 157 (1940). soratensis Rusby in Bull. N. Y. Bot. Gard. 6: 510 (1910). sufferrugineus Rusby in Bull. N. Y. “ee hae 6: 510 (1910). tacitus C. P. Smith, Spec. Lup. 460 (194 tarijensis Ulbr. in Engler, Bot. noes 7 543 (1906). Tatei Rusby in Phytologia, 1: 57 (193 tominensis Wedd. Chlor. And. 2: 251 inigh Williamsianus C. P. Smith, Spec. Lup. 163 (1940). xenophytus C. P. Smith, Spec. Lup. 216 (1940). Lysiloma polyphylla (Clos) Benth. in Trans. Linn. Soc. 30: 535 (1875). Machaerium aculeatum Raddi in Mem. Soc. Ital. Moden. 18: 598 (1820). M. acutifolium Vog. in Linnaea, 11: 187 (1837). M. angustifolium Vog. in Linnaea, 11: 193 (1837). AE AR ORR b Prrr hrm r rrr rr ete Bangii Rusby in Bull. N. Y. Bot. Gard. 4: 347 (1907). bolivianum Gandog. in Bull. Soc. Bot. France, 60: 460 (1913). decorticans Ducke in Archiv. Jard. Bot. Rio Janeiro, 3: 150 (1922). eriocarpum Benth. in Ann. Naturh. Hofmus. Wien, 2: 98 (1838). foliosum Rusby in Mem. N. Y. Bot. Gard. 7: 266 (1927 jacarandifolium Rusby in Bull. N. Y. Bot. Gard. 6: 514 (1910). juglandifolium Rusby in Bull. N. Y. Bot. Gard. 6: 513 (1910). latifolium Rusby in Bull. N. Y. Bot. Gard. 6: 513 es longifolium Benth. in Journ. Linn. Soc. 4: Suppl. 55 (1860). longistipitatum Hoehne in Arq. Bot. Est. S. Paulo (n. s.) 1: 48, t. 59 (1938). Moritzianum Benth. in Journ. Linn. Soc. 4: Suppl. 58 (1860). paraguariense Hassl. in Bull. Herb. Boiss. (ser. 2) 7: 358 (1907). rogaguense Rusby in Mem. N. Y. Bot. Gard. 7: 266 (1927). sordidum Benth. in Mart. Fl. Bras. 15(1): 241 (1862). Steinbachianum Hoehne in Arq. Bot. Est. S. Paulo (n. s.) 1: 48, t. 60 (1938). villosum Vog. in Linnaea, 11: 189 (1837). Medicago arabica All. Flor. Pedem. 1: 315 (1785). M. denticulata Willd. Sp. Pl. 3(2): 1414 (1800). M. lupulina L. Sp. Pl. 779 (1758). M. sativa L. Sp. Pl. 778 (1753). Meibomia adhaerens Rusby in Phytologia, 1: 58 (1934) = Desmodium sp. M. adpressa Rusby in Phytologia, 1: 58 (1934) — Desmodium sp. M. andina Rusby in Mem. Torr. Bot. Club, 3(3): 21 (1893) = Desmodium axillare. ” M. Malmei Schindl. in Fedde, Repert. Spec. Nov. 20: 144 (1924) — Desmodium sp. M. microcarpa Rusby in Mem. N, Y. Bot. Gard. 7: 263 (1927) = Desmodium sp. M. nummularia Rusby in Phytologia, 1: 59 (1934) = Desmodium s M. ovalis Rusby in Mem. N. Y. Bot. Gard. 7: 264 (1927) = aapeliiens sp. M. parva Schindl. in Fede, Repert. Spec. Nov. 20: 15 (1924) = Desmodium M. rhmctoiota Rusby in Bull. N. Y. Bot. Gard. 6: 513 (1910) = Desmodium M. Sehipict hive Rusby in Phytologia, 1: 60 (1934) = Desmodium a M. tetrasperma Rusby in og pee 1: 60 (1934) = Desmodium s: M. variegata Rusby in Bull. N. Y. Bot. Gard. 4: 344 (1907) = beanadtans sp. Melilotus albus Desv. in Lam. Rueye: 4: 63 (1796). M. indicus (L.) All. Flor. Pedem. 1: 308 (1785 Mimosa albida Humb. & Bonpl. ex Willd. Sp. PI. ‘4(2): 1030 (1806). M. auriculata Benth. in Trans. Linn. Soc. 30: 430 (1875). M. eurycarpa Herzog in Fedde, Repert. Spec. Nov. 7: 53 (1909) = M. Herzogii. M. floribunda Willd. Sp. Pl. 4(2): 1031 (1806). M. glaucescens var. ramosa Benth. in Mart. FI. Bras. fon 308 (1876). M. gonoclada Benth. in Trans. Linn. Soc. 30: 427 (18 M. grandistipula Herzog in Fedde, Repert. Spec. Nov. se 58 (1909). M. Herzogii Macbr. in Contrib. Gray Herb. 59: 14 (1919). M. insidiosa Mart. Herb. Fl. Bras. 134 (1837). M. Kuntzei Harms in O. Ktze. Rev. Gen. 3(2): 67 (1898) = M. boliviana. M. lepidota Herzog in Fedde, Repert. Spec. Nov. 7: 53 (1909) M. Lorentzii Griseb. in Goett. Abh. 19: 134 (1874). M. neptunioides Harms ex O. Ktze. Rev. Gen. 3(2): 67 (1898). M. nuda Benth. in Hook. Journ. Bot. 4: 362 (1841). orthacantha Benth. in Hook. Journ. Bot. 4: 365 (1841). parapitiensis Burkart in Bol. Soc. Argent. Bot. 1(1): 36 (1945). pigra L. Cent. Pl. 1: 18 (1755). polycarpa Kunth, Mimos. 8, t. 3 (1819). So (Kunth) Benth. in Hook. Journ. Bot. 4: 409 (1841). a Mart. in Flora, 21 (II Beibl.): 57 (1838). AEP Saal. Benth. in Mart. Fl. Bras. 15(2): nse (1876). — Griseb. in Goett. Abh, 24: 119 (187 somnians Humb. & Bonpl. ex Willd. Sp. Pl. oe 1036 (1806). M. ie anaia Benth. in Trans. Linn. Soc. 30: 427 (1875). M. Steinbachii Harms in Notizbl. 9: 1038 (1926). M. stenoptera Benth. in Trans. Linn. Soc. 30: 395 (1875). M. subsericea Benth. in Hook. Journ. Bot. 4: 380 (1841). M. velascoénsis Harms ex O. Ktze. Rev. Gen. 3(2): 68 (1898). M. Velloziana Mart. in Flora, 22 (I Beibl.): 9 (1839). M. Williamsii Rusby in Bull. N. Y. Bot. Gard. 8: 91 (1912). Mucuna mapirensis (Rusby) Macbr. in Field Mus. Publ. Bot. 13(3): 315 (1943). M. rostrata Benth. in Mart. Fl. Bras. 15(1): 171, t. 47 (1859). Myroxylon peruiferum L. f. Suppl. 233 (1781). Neocracca Kuntzei O. Ktze. Rev. Gen. 3(2): 68 (1898) = Cracca heterantha. Nissolia fruticosa Jacq. Enum. Pl. Carib. 27 (1760). Ormosia bopiensis Pierce ex Macbr. in Field Mus. Publ. Bot. 13: 248 (1943). Pachyrhizus bulbosus (L.) Britton in Bull. Torr. Bot. Club, 16: 324 (1889). P. tuberosus (Lam.) Spreng. Syst. 4: ae Post. 281 (1827). Parkinsonia aculeata L. Sp. Pl. 375 (175 —— oblongifolia Rusby in Mem, N. “Bot. Gard. 7: 262 (1927) = Dalea 2 4 aapaiapage Yr. pr Ree Rusby in Mem. N. Y. Bot. Gard. 7: 261 (1927) = Dalea sp. saab incr pi Catieai Rusby in Bull. N. Y. Bot. Gard. 6: 512 (1910) = Adesm cue F. grandidentatum Rusby in Mem. Torr. Bot. Club, 6: 24 (1896) — Adesmia sp. Fr, et Rusby in Bull. N. Y. Bot. Gard. 6: 512 (1910) — Adesmia sp. ? Phaseolus adenanthus var. radicans (Benth.) Hassl. in Candollea, 1: 4438 (1923). Augustii Harms in Notizbl. 7: 503 (1921). P. bolivianus Piper in Contrib. U. S. Nat. Herb. 22: 694 (1926). bracteolatus Nees & Mart. in Nov. Act. Acad. Caes. Leop. 12: 27 (1824). campestris (Benth.) Mart. ex Benth. in Mart. Fl. Bras. 15(1): 188 (1859). caracalla L. Sp. Pl. 725 (1753). erythroloma (Benth.) Mart. ex Benth. in Mart. Fl. Bras. 15(1): 191 (1859). fraternus Piper in Contrib. U. S. Nat. Herb. 22: 679 (1926). lathyroides L. Sp. Pl. (ed. 2) 1018 (1763). ligulatus Piper in Contrib. U. S. Nat. Herb. 22: 676 (1926). eS var. linearifoliolates forma boliviensis Hassl. in Candol- lea, 1: 453 (19 ovatus pene as tl in Mart. Fl. Bras. 15(1): 188 (1859). peduncularis HBK. Nov. Gen. & Spec. 6: 447 (1824). prostratus Benth. in Mart. Fl. Bras. 15(1): 192 (1859). prostratus var. angustifolius forma flavoviridis O. Ktze. Rev. Gen. 3(2): 70 (1898). prostratus forma macrophyllus Hassl. in Candollea, 1: 459 (1923). rigidus Piper in Contrib. U. S. Nat. Herb. 22: 699 (1926) truxillensis HBK. Nov. Gen. & Spec. 6: 451 (1824). vignoides Rusby in Bull. N. Y. Bot. Gard. 4: 845 (1907) = Canavalia lasio- calyx. sy Pee wet BARRE oe Piptadenia colubrina (Vell.) Benth. in Hook. Journ. Bot. 4: 341 (1841). P. communis Benth. in Hook. Journ. Bot. 4: 337 (1841). P. macrocarpa Benth. in Hook. Journ. Bot. 4: 341 (1841). Pithecellobium angustifolium (Rusby) Rusby in Mem. N. Y. Bot. Gard. 7: 253 (1927) = P. sophorocarpum. bifoliolum Rusby in Mem. N. Y. Bot. Gard. 7: 254 (1927). dependens Rusby in Mem. N. Y. Bot. Gard. 7: 253 (1927). divaricatum Benth. in Hook. Lond. Journ. Bot. 3: 213 (1844). latifolium (L.) Benth. in Hook. Lond. Journ. Bot. 2: 214 (1844). laxiflorum Rusby in Mem. Torr. Bot. Club, 3(3): 24 (1893). niopoides Spruce ex Benth. in Mart. Fl. Bras. 15(2): 447 (1876). pedicellare Benth. in Hook. Lond. Journ. Bot. 3: 219 (1844). P. pendulum Lindm. in Bihang Kgl. Sv. Vet. Akad. Handl. 24 (Afd. 3, no. 7): 51 (1898). Saman (Jacq.) Benth. in Hook. Lond. Journ. Bot. 3: 216 (1844). scalare Griseb. in Goett. Abh. 24: 123 (1879). sophorocarpum Benth. in Benth. & Hook. f. Gen. Pl. 1: 598 (1865). trapezifolium (Vahl) Benth. in Hook. Lond. Journ. Bot. 3: 204 (1844). tortum Mart. Herb. Fl. Bras. 114 (1837). venosum Rusby in Mem. Torr. Bot. Club, 6: 29 (1896). Platymiscium cochabambense Rusby in Mem. Torr. Bot. Club, 6: 26 (1896). P. ellipticum Rusby in Bull. N. Y. Bot. Gard. 6: 514 (1910). P. fragrans Rusby in Mem. N. Y. Bot. Gard. 7: 267 (1927). Poiretia scandens Vent. Choix de Pl. t. 42 (1803). Prosopis Algarobillo Griseb. in Goett. Abh. 19: 131 (1874). ferox Griseb. in Goett. Abh. 24: 118 (1879). Herzogii Harms in Meded. Rijks Herbar. 27: 35 (1915). julifiora (Sw.) DC. Prodr. 2: 447 (1825). Kuntzei Harms ex O. Ktze. Rev. Gen. 3(2): 71 (1898). nigra (Griseb.) Hieron. in Bol. Acad. Nac. Cérdova, 4: 283 (1881). ruscifolia Griseb. in Goett. Abh. 19: 130 (1874). Psoralea glandulosa L. Sp. Pl. (ed. 2) 1075 (1763). lasiostachys Vog. in Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 13 (1843). lasiostachys var. potens (Macbr.) Macbr. in Field Mus. Publ. Bot. 8: 88 (1930). mexicana (L. f.) A. M. Vail in Bull. Torr. Bot. Club, 21: 119 (1894). Mutisii HBK. Nov. Gen. & Spec. 6: 487 (1824). timorata Macbr. in Field Mus. Publ. Bot. 8: 89 (1930). Pterocarpus Steinbachianus Harms in Notizbl. 9: 1039 (1926). P. violaceus Vog. in Linnaea, 11: 416 (1887). ee IU UU MMMM UN a ea a Samanea coripatensis (Rusby) Killip ex Record in Trop. Woods, 63: 6 (1940). Sclerolobium Radlkoferi Rusby in Mem. Torr. Bot. Club, 6: 26 (1896). Steinbachiella leptoclada Harms in Notizbl. 10: 345 (1928). Stylosanthes Bangii Taub. in Rusby in Mem. Torr. Bot. Club, 4(3): 206 (1895). S. gracilis var. vulgaris Burkart in Darwiniana, 3: 247 (1939). S. guianensis (Aubl.) Sw. in Vet. Acad. Stockh. 1789: 296. S. guianensis var. subviscosa Benth. in Mart. Fl. Bras. 15(1): 92 (1859). S. hamata (L.) Taub. in Abh. Bot. Ver. Brandenb. 32: 22, fig. 22 (1890). 102 S. juncea Micheli in Mém. Soc. Phys. Hist. Nat. Genéve, 28 (7): 19 (1883). S. montevidensis Vog. in Linnaea, 12: 67 (1838). S. scabra Vog. in Linnaea, 12: 69 (1838). Swartzia Jorori Harms in Meded. Rijks Herbar. 27: 39 (1915). S. Matthewsii Benth. in Hook. Ic. Pl. 11: 51, t. 1064 (1870). Tamarindus indica L. Sp. Pl. 34 (1753). Tephrosia adunca Benth. in Ann. Nat. Hist. (ser. 1) 3: 432 (1839). T. leptostachya DC. Prodr. 2: 251 (1825). T. toxicaria Pers. Synops. Pl. 2: 329 (1807). Teramnus uncinatus (L.) Sw. Fl. Ind. Occ. 3: 1239 (1806). Tipuana speciosa Benth. in Journ. Linn. Soc. 4: Suppl. 72 (1860). Tounatea arborescens (Aubl.) Britton in Bull. Torr. Bot. Club, 16: 325 (1889). T. costata Rusby, Descr. S. Am. Pls. 28 (1920). T. fugax (Spruce) Britton in Bull. Torr. Bot. Club, 16: 325 (1889). Tragacantha arequibensis var. minima O. Ktze. Rev. Gen. 3(2): 73 (1898) = Astragalus micranthellus. T. arequibensis var. tenuifolia O, Ktze. Rev. Gen. 3(2): 73 (1898) — As- tragalus micranthellus. Trifolium amabile HBK. Nov. Gen. & Spec. 6: 503, t. 593 (182 4). T. peruvianum Vog. in Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 12 (1848). Vicia acerosa Clos in Gay, Hist. Chile, 2: 187 (1846 V. graminea Smith in Rees, Cycl. 37: no. 27 (1819). V. montevidensis Vog. in Linnaea, 13: 34 (1839). V. setifolia HBK. Nov. Gen. & Spec. 6: 500 (1824). Vigna myrtifolia Piper in Contrib. U. S. Nat. Herb. 22: 666 (1926). Zornia diphylla (L.) Pers. Synops. Pl. 2: 318 (1807). Z. diphylla var. latifolia (DC.) Benth. in Mart. Fl. Bras. 15(1): 81 (1859). OXALIDACEAE Biophytum bolivianum R. Knuth in Engler, Pflanzenr. IV, 130: 403 (1930). . ferrugineum Rusby in Bull. N. Y. Bot. Gard. 8: 95 (1912). globuliflorum R. Knuth in Engler, Pflanzenr. IV, 130: 403 (1930). mapirense R. Knuth in Engler, Pflanzenr. IV, 130: 405 (1930). mapirense var. hirtum R. Knuth in Engler, Pflanzenr. IV, 180: 405 (1930). peruvianum R. Knuth in Engler, Pflanzenr. IV, 130: 402 (1930). pseocharis corydalifolia R. Knuth in Engler, Bot. Jahrb. 41: 173 (1908). Fiebrigii R. Knuth in Engler, Bot. Jahrb. 41: 173 (1908). malpasensis R. Knuth in Fedde, Repert. Spec. Nov. 29: 219 (1931). moschata R. Knuth in Meded. Rijks Herbar. 27: 67 (1915). pedicularifolia R. Knuth in Engler, Bot. Jahrb. 41: 173 (1908). . pimpinellifolia Remy in Ann. Sci. Nat. (ser. 3) 8: 239 (1847). . tridentata Griseb. in Goett. Nachr. 1877: 495. cnonelia canaminensis Rusby in Mem. N. Y. Bot. Gard. 7: 269 (1927) Oxalis Lotoxalis ech Rusby in Mem. N. Y. Bot. Gard. 7: 269 (1927) = Oxalis sepium. Oxalis aetheria Macbr. in Candollea, 6: 8 (19384) — O. pygmaea. . adpressa R. Knuth in Engler, Pflanzenr. IV, 130: 160 (1930). affinis R. Knuth in Meded. Rijks Herbar. 27: 61 (1915) — O. cognita. albicans var. sericea DC. Prodr. 1: 693 (1824). andina Britton in Bull. Torr. Bot. Club, 16: 159 (1889). aphylla Rusby in Bull. N. Y. Bot. Gard. 8: 95 (1912). arenaria Bert. in Merc. Chil. 16: 739 (1829). Asplundii R. Knuth in Fedde, Repert. Spec. Nov. 24: 53 (1927). azanaquensis R. Knuth in Fedde, Repert. Spec. Nov. 29: 217 (1931). eae eee S99999909 Poeoof SEES9SS99S9999999999 S9999999999999999 SOODD9000000 103 Bangii Rusby in Bull. N. Y. Bot. Gard. 4: 337 ( 1907). Barrelieri L. Sp. Pl. (ed. 2) 1: 624 (1762). bermejensis R. Knuth in Notizbl. 7: 312 (1919). bipartita St. Hil. Fl. Bras. Mer. 1: 125, t. 25 (1825). bisfracta Turcz. in Bull. Soc. Nat. Mose. 36(1): 595 (1863). boliviana Britton in Bull. Torr. Bot. Club, 16: 159 (1889). breviramulosa Rusby in Mem. Torr. Bot. Club, 3(3): 13 (1898). brunneo-pilosa R. Knuth in Fedde, Repert. Spec. Nov. 29: 215 (1981). Buchtienii (Rusby) R. Knuth in Fedde, Repert. Spec. Nov. 24: 55 (1927). bulbifera R. Knuth in Notizbl. 7: 310 (1919). calachaccensis R. Knuth in Meded. Rijks Herbar. 27: 64 (1915) = O. brevi- ramulosa. capitata R. Knuth in Meded. Rijks Herbar. 27: 61 (1915). carnosa Molina, Sagg. Stor. Nat. Chil. (ed. 2) 288 (1810). chacoénsis R. Knuth in Fedde, Repert. Spec. Nov. 23: 138 (1926), charaguensis R. Knuth in Meded. Rijks Herbar. 27: 62 (1915). cognita R. Knuth in Fedde, Repert. Spec. Nov. 23: 141 (1926). Commersonii Pers. Synops. Pl. 1: 519 (1805) corniculata L. Sp. Pl. 485 (1753). cotagaitensis R. Knuth in Fedde, Repert. Spec. Nov. 29: 215 (1931). cuzcensis R. Knuth in Notizbl. 7: 303 (1919). densissima Rusby in Phytologia, 1: 61 (1934). dolichopoda Diels in Engler, Bot. Jahrb. 37: 423 (1906). elegans HBK. Nov. Gen. & Spec. 5: 234, t. 466 (1822). eriolepis Wedd. Chlor. And. 2: 290 (1861). erythropoda Rusby in Mem. Torr. Bot. Club, 6: 16 (1896). filiformis HBK. Nov. Gen. & Spec. 5: 245, t. 469 (1822). glaberrima Norlind in Ark. Bot. 20A (4): 26, t. 3, fig. 4 (1926). guaquiensis R, Knuth in Meded. Rijks Herbar. 27: 63 (1915) = O. brevi- ramulosa. Harmsiana R. Knuth in Engler, Pflanzenr. IV, 130: 431 (1930). Hauthalii R. Knuth in Notizbl. 7: 305 (1919). Herzogii R. Knuth in Meded. Rijks Herbar. 27: 63 (1915). irregularis R. Knuth in Fedde, Repert. Spec. Nov. 23: 141 (1926). lepidocaulis Norlind in Ark. Bot. 20A (4): 22, t. 2, fig. 1 (1926). Lilloana R. Knuth in Fedde, Repert. Spec. Nov. 23: 140 (1926). longissima (O. Ktze.) K. Schum. in Just, Bot. Jahresber. 26(1): 360 (1898). lotoides HBK. Nov. Gen. & Spec. 5: 241 (1822). magellanica Forst. in Comm. Goett. 9: 33 (1789). manihotoides (Rusby) R. Knuth in Fedde, Repert. Spec. Nov. 23: 141 (1926). mapirensis R. Knuth in Engler, Pflanzenr. IV, 130: 174 (1930). artiana Zucc. in Denkschr. Akad. Miinch. 9: 144 (1825). medicaginea HBK. Nov. Gen. & Spec. 5: 242 (1822). megalorrhiza Jacq. Oxal. 33 (1794). microcarpa Benth. Pl. Hartw. 115 (1843). mollis HBK. Nov. Gen. & Spec. 5: 241 (1822). mollissima (Rusby) R. Knuth in Fedde, Repert. Spec. Nov. 23: 275 (1927). nubigena Walp. in Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 320 (1843). ollantaytambensis R. Knuth in Fedde, Repert. Spec. Nov. 28: 142 (1926) = O. cuzcensis. oxyptera Prog. in Mart. Fl. Bras. 12(2): 489, t. 103 (1877). parapitensis R. Knuth in Meded. Rijks Herbar. 27: 62 (1915). parvifolia DC. Prodr. 1: 693 (1824). parvifolia var. pluriflora R. Knuth in gma . 301 (1919). parvula Remy in Ann. Sci. Nat. (ser. 3) 6: pazensis (Rusby) R. Knuth in Fedde, ihe Spec. Nov, 24: 55 (1927). e i) _ phaseolifolia (Rusby) R. Knath in Fedde, Repert. Spec. Nov. 23: 140 (1926). Philippii R. Knuth in Meded. Rijks Herbar. 27: 67 (1915). pinguiculacea R. Knuth in Notizbl. 7: 311 (1919). pinoénsis R. Knuth in Fedde, Repert. Spec. Nov. 29: 213 (1931). eiatviedi Wedd. Chlor. And. 2: 290 (1861). pseudo-violacea R. Knuth in Fedde, Repert. Spec. Nov. 24: 55 (1927). pubescens HBK. Nov. Gen. & Spec. 5: 240 (1822). pycnophylla Wedd. Chlor. And. 2: 291 (1861). pygmaea A. Gray in Bot. U. S. Explor. Exped. 1: 322 (1854). Regnellii Mig. in Linnaea, 22: 545 (1849). renifolia R. Knuth in Notizbl. 7: 296 (1919). rubrovenosa Norlind in Ark. Bot. 20A (4): 21 (1926). scandens HBK. Nov. Gen. & Spec. 5: 242 (1822). soldanelliflora R. Knuth in Fedde, Repert. Spec. Nov. 23: 275 (1926). Steinbachii R. Knuth in Fedde, Repert. Spec. Nov. 24: 55 (1927). Steinmannii Solms-Laub. in Bot. Zeitung, 65(1): 182, t. 2, fig. 7 (1907). Tatei Rusby in Phytologia, 1: 62 (1934). teneriensis R. Knuth in Meded. Rijks Herbar. 27: 64 (1915). tenerrima R. Knuth in Fedde, Repert. Spec. Nov. 23: 277 (1926). tenuiscaposa R. Knuth in Meded. Rijks Herbar. 27: 66 (1915). tocoranensis R. Knuth in Meded. Rijks Herbar. 27: 65 (1915). Trollii R. Knuth in Fedde, Repert. Spec. Nov. 29: 218 (1931). tuberosa Molina, Sagg. Stor. Nat. Chile, 132 (1782). unduavensis (Rusby) R. Knuth in Fedde, Repert. Spec. Nov. 23: 142 (1926). violacea L. Sp. Pl. 434 (1753). virgata Rusby in Phytologia, 1: 62 (1934). yapacaniensis (O. Ktze.) K. Schum. in Just, Bot. Jahresber. 26(1): 360 (1898). peers Po. oP oemer ear ere es - yungasensis Rusby in Mem. Torr. Bot. Club, 3(3): 12 (18938). Xanthoxalis biflera Rusby, Descr. S. Am. Pls. 31 (1920) —= Oxalis breviramu- losa. X. flagellata Rusby, Deser. S. Am. Pls. 81 (1920) == Oxalis andina. GRERANTACKAR Balbisia integrifolia R. Knuth in Engler, Pflanzenr. IV, 129: 558 (1912). B. Meyeniana Klotzsch in Linnaea, 10: 432 (1836). Erodium cicutarium (L.) L’Hérit. ex Ait. Hort. Kew. 2: 414 (1789). E. moschatum L’Hérit. ex Ait. Hort. Kew. 2: 414 (1789). Geranium album R. Knuth in Engler, Bot. Jahrb. 37: 557 (1906). amoenum R. Knuth in Fedde, Repert. Spec. Nov. 40: 217 (1936). . Bangii Hieron. in Engler, Bot. Jahrb. 21: 314 (1895). bolivianum R. Knuth in Engler, Pflanzenr. IV, 129: 575 (1912). carolinianum L. Sp. Pl. 682 (1753). chaparense R. Knuth in Fedde, Repert. Spec. Nov. 28: 4 (1930). comarapense R. Knuth in Meded. Rijks Herbar. 27: 69 (1915). cuchillense R. Knuth in Fedde, Repert. Spec. Nov. 34: 144 (19383). diffusum HBK. Nov. Gen. & Spec. 5: 230 (1822). escalonense R. Knuth in Fedde, Repert. Spec. Nov. 34: 145 (1933). Fiebrigianum R. Knuth in Engler, Bot. Jahrb. 37: 560 (1906). fuscicaule R. Knuth in Fedde, Repert. Spec. Nov. 28: 7 (1930). Herzogii R. Knuth in Meded. Rijks Herbar. 27: 69 (1915). Lechleri R. Knuth in Engler, Pflanzenr. IV, 129: 80 (1912). malpasense R. Knuth in Fedde, Repert. Spec. Nov. 34: 146 (1933). mexicanum var. minoriflorum (Briq.) R. Knuth in Engler, Pflanzenr. IV, 129: 197 (1912). AAARAAARAAARAARANHAA 105 G. paleaénse R. Knuth in Meded. Rijks Herbar. 27: 68 (1915). G. pallidifolium R. Knuth in Fedde, Repert. Spec. ang 28: 3 (1930). G. patagonicum Hook. f. Fl. Antarct. 2: 252 (184 G. Pflanzii R. Knuth in Engler, Pflanzenr. IV, ms 576 (1912). G. rupicola Wedd. Chlor. And, 2: 285 (1861). G. sepalo-roseum Rusby in Mem. Torr. Bot. Club, 2 12 (1898). G. sessiliflorum Cav. Diss. 4: 198, t. 77, fig. 2 (1787). G. sessiliflorum var. compactum R. Knuth in Engler, Pflanzenr. IV, 129: 85 G. sessiliflorum var. lanatum R. Knuth in Engler, Bot. Jahrb. 37: 565 (1906). G. sessiliflorum var. microphyllum O. Ktze. Rev. Gen. 3(2): 33 (1898). G. soratae R. Knuth in Engler, Pflanzenr. IV, 129: 212 (1912). G. superbum R. Knuth in Engler, Bot. Jahrb. 37: 561 (1906). . tablasense R. Knuth in Meded. Rijks Herbar. 27: 68 (1915) = G. patagoni- cum. G. titicacaénse R. Knuth in Fedde, Repert. Spec. Nov. 45: 60 (1938) = G. patagonicum. G. totorense R. Knuth in Fedde, Repert. Spec. Nov. 28: 7 (1930). G. Weddellii Briq. in Ann. Conserv. & Jard. Bot. Genéve, 11-12: 183 (1908). TROPAEOLACEAE Tropaeolum boliviense Loes. in Engler, Bot. Jahrb. 45: 462 (1911). T. cochabambense Buchenau in Engler, Bot. Jahrb. 22: 161 (1895). T. cuspidatum Buchenau in Engler, Bot. Jahrb. 26: 581 (1899). T. infundibularum Rusby in Bull. N. Y. Bot. Gard. 4: 336 (1907) = T. cuspi- datum. T. Kuntzeanum Buchenau in Engler, Bot. Jahrb. 22: 163 (1895). T. maculatum Rusby in Mem. Torr. Bot. Club, 6: 15 (1896). T. pentaphyllum var. megapetalum Buchenau in Engler, Bot. Jahrb. 22: 169 T. peregrinum L. Sp. Pl. 345 (1753). T. rectangulum Buchenau in Engler, Bot. Jahrb. 22: 165 (1895). T. rectangulum var. bicolor O. Ktze. Rev. Gen. 3(2): 33 (1898). T. rectangulum var. pallidum O. Ktze. Rev. Gen. 3(2): 33 (1898). T. Seemannii Buchenau in Engler, Bot. Jahrb. 15: 226 (1892). T. Smithii DC. Prodr. 1: 684 (1824). T. tuberosum R. & P. Fl. Peruv. 3: 77, t. 314 (1802). T. umbellatum Hook. in Bot. Mag. 73: t. 4337 (1847). LINACEAE Linum filiforme Urb. in Linnaea, 41: 643 (1877). ? L. scoparium Griseb. in Goett. Abh. 19: 103 (1874). Roucheria laxiflora H. Winkl. in Fedde, Repert. Spec. Nov. 7: 109 (1909). ERYTHROXYLACEAE picts ene anguifugum Mart. in Abh. Akad. Wiss. Miinch. 1840: 361. E. Bangii Rusby in Mem. Torr. Bot. Club, 3(3): 11 (1893). bolivianum Burck. in Teysmannia, 1: 456, t. 3 (1890). Coca Lam. Encye. 2: 393 (1786). cuneifolium var. silvaticum O. E. Schulz in Engler, Pflanzenr. IV, 134: 122 1907). epee O. E. Schulz in Fedde, Repert. Spec. Nov. 7: 57 (1909). lucidum HBK. Nov. Gen. & Spec. 5: 179 (1822). macrophyllum Cav. Diss. 8: 401 (1789). bs et et ted bt opacum Rusby in Mem. N. Y. Bot. Gard. 7: 270 (1927). pauciflorum Rusby in Mem. Torr. Bot. Club, 6: 13 (1806). paraénse Peyr. in Mart. Fl. Bras, 12(1): 164, t. 30 (1878). Trollii O. E. Schulz in Notizbl. 11: 723 (1933). Ulei O. E. Schulz in Engler, Pflanzenr. IV, 134: 62 (1907). venosum Rusby in Mem. N. Y. Bot. Gard. 7: 270 (1927). ZYGOPHYLLACEAE Bulnesia bonariensis Griseb. in Goett. Abh. ee 105 (1874). B. foliosa Griseb. in Goett. Abh. 19: 106 (1874). Sarmienti Lor. ex Griseb. in Goett. Abh. 24: 75 (1879). Kallstroemia boliviana Standl. in Field Mus. Publ. Bot. 11: 161 (1936). K. maxima (L.) Torr. & Gray, N. Am. Fl. 1: 213 (1838). K. tribuloides (Mart.) Wight & Arn. Prodr. 145 (1834), combination implied but not actually made. Larrea divaricata Cav. in Anal. Hist. Nat. Madrid, 2: 122, t. 19, fig. 1 (1800). Porliera arida Rusby in Mem. Torr. Bot. Club, 6: 15 (1896). P. hygrometra R. & P. Fl. Peruv. 4: t. 343 (1802). P. Lorentzii Engl. in Engl. & Prantl, Nat. Pflanzenfam. 3(4): 84 (1890) = P. microphylla. P. microphylla (Baill.) Desc. O’Don. & Lourt. in Lilloa, 5: 329 (1940). P. Steinbachii Standl. in Field Mus. Publ. Bot. 11: 162 (1936) = P. micro- phylla. Tribulus cistoides L. Sp. Pl. te (1753). T. maximus var. roseus O. Ktze. Rev. Gen. 3(2): 30 (1898) = ? T. terrestris L. Sp. Pl. 387 esis. RUTACEAE Citrus acida Roxb. Fl. Ind. 3: 390 (1882). Cusparia pilocarpidia Rusby in Bull. N. Y. Bot. Gard. 8: 98 (1912). Dictyoloma peruvianum Planch. in Hook. Lond. Journ. Bot. 5: 583 (1846). Erythrochiton brasiliensis Nees & Mart. in Nov. Act. Acad. Caes. Leop. 11: 166 (1823). Esenbeckia lucida Rusby in Bull. N. Y. Bot. Gard. 8: 98 (1912). Fagara aculeatissima Engl. in Engl. & Prantl, Nat. Pflanzenfam. (ed. 2) 19A: 220 yi comosa Herzog in Meded. Rijks Herbar. 27: 80 (1915). cuiabensis var. axillaris O. Ktze. Rev. Gen. 3(2): 34 (1898). Naranjillo (Griseb.) Engl. in Engl. & Prantl, Nat. Pflanzenfam. 3(4): 117 (1896). nebuletorum Herzog in Meded. Rijks Herbar. 27: 79 (1915). nigrescens R. E. Fries in Ark. Bot. 8(8): 12, t. 1, fig. 4-5 (1908). Pterota L. Syst. (ed. 10) 897 (1759). rhoifolia (Lam.) Engl. in Engl. & Prantl, Nat. Pflanzenfam. 3(4): 118 897). rigidifolia Herzog in Meded. Rijks Herbar. 27: 78 (1915). . tenuifolia forma aculeata Herzog in Meded. Rijks Herbar. 27: 80 (1915). Ruta graveolens L. Sp. Pl. 383 (1753). Ticorea longiflora DC. in Mém. Mus. Paris, 9: 146, t. 9 (1822). Zanthoxylum annulatum Rusby in Mem. N. Y. Bot. Gard. 7: 277 (1927). Z. Cardenasii Rusby in Mem. N. Y. Bot. Gard. 7: 277 (1927). B. Coco Gill. ex Hook. & Arn. in Hook. Bot. Misc. 3: 168 (1833). Z. pubescens St. Hil. & Tul. in Ann. Sci. Nat. (ser. 2) 17: 141 (1842). Z. stipitatum Engl. in Mart. Fl. Bras. 12(2): 161, t. 34 (1874). meh Pea af Pat Pah a 107 SIMAROUBACEAE Alvaradoa amorphoides Liebm. in Vidensk. te for 1853: 101 (1854). A. subovata Cronquist in Brittonia, 5: 134 (1944). Icica rhynchophylla Rusby in Mem. N. Y. Bot. Gard. 7: 278 (1927) = Protium insigne. Picramnia Corallodendron Tul. in Ann. Sci. Nat. (ser. 3) 7: 259 (1847). magnifolia Macbr. in Candollea, 5: 376 (1934). monninaefolia Rusby in Mem. N. Y. Bot. Gard. 7: 278 (1927). pendula Herzog in Fedde, Repert. Spec. Nov. 7: 58 (1909). Sellowii Planch. in Hook. Lond. Journ. Bot. 5: 578 (1846). Spruceana Engl. in Mart. Fl. Bras, 12(2): 238 (1874). Simarouba amara Aubl. Pl. Guian. Fr. 2: 860, tt. 331-332 (1775). S. versicolor var. pallida Engl. in Mart. Fl. Bras. 12(2): 226 (1874). +: RRA dd BURSERACEAE Bursera amplifolia Rusby in Bull. N. Y. Bot. Gard. 4: 338 (1907). Protium Bangii Swart in Act. Bot. Neerland. 1: 244 (1952) bolivianum Britton in Bull. Torr. Bot. Club, 16: 189 (1899) = Mauria ferruginea. P. guianense (Aubl.) Planch. in Adansonia, 8: 52 (1867). heptaphyllum var. brasiliense Engl. in Mart. Fl. Bras. 12(2): 263 (1874). insigne (Tr. & Pl.) Engl. in DC. Monog. Phan. 4: 77 (1883). Llewelynii Macbr. in Candollea, 5: 878 (1934). meridionale Swart in Act. Bot. Neerland. 1: 246 (1952). montanum Swart in Act. Bot. Neerland. 1: 247 (1952). vatum Engl. in Mart. Fl. Bras. 12(2): 264, t. 52 (1874). ; ger otnoho Engl. in DC. Monog. Phan. 4: 76 (1883). S sp. ~ ry ote dG ° Tetragas Trattinickis Lawrancei var. boliviana Swart in Act. Bot. Neerland. 1: 249 952). T. rhoifolia subsp. Sprucei var. pubescens Swart in Act. Bot. Neerland. 1: 249 (1952). MELIACEAE Cedrela boliviana Rusby, Descr. S. Am. Pls. 36 (1920). co. brunellioides Rusby in Bull. N. Y. Bot. Gard. 8: 99 (1912). c. odorata L. Syst. (ed. 10) 940 (1759). C. Steinbachii Harms in Notizbl. 11: 381 (1932). Guarea alborosea Rusby in Bull. Torr. Bot. Club, 49: 263 (1922) = G. pen- dulispica. Bangii Rusby in Bull. Torr. Bot. Club, 49: 262 (1922). membranacea Rusby in Mem. N. Y. Bot. Gard. 7: 279 (1927). ovalis (Rusby) Rusby in Mem. Torr. Bot. Club, 4(3): 205 (1895). pendulispica C. DC. in Fedde, Repert. Spec. Nov. 7: 59 (1909). Steinbachii Harms in Notizbl. iy 848 (1928). iniecinona L. Mant. 228 (177 ana Briq. in Candollea, i. "20 (1935). Melia gran L. Sp. Pl. 384 (1753). pinnae dati viride Rusby in Mem. Torr. Bot. Club, 6: 18 (1896) = R ensis Harms in Notizbl. 10: 347 (1928) = Guarea sp. Swietenia A qegaeate Jacq. Enum. Pl. Carib. 20 (1760). Sycocarpus Rusbyi Britton in Bull. Torr. Bot. Club, 14: 143 (1887) = = Guarea ciation 108 Trichilia Buchtienii Harms in Notizbl. 11: 386 (1932). Cardenasii Rusby in Mem. N. Y. Bot. Gard. 7: 279 (1927). Catigua A. Juss. in St. Hil. Fl. Bras. Mer. 2: 77 (1829). Claussenii C. DC. in Mart. Fl. Bras. 11(1): 207 (1878). elegans A. Juss. in St. Hil. Fl. Bras. Mer. 2: 79, t. 98 (1829). guayaquilensis C. DC. in DC. Monog. Phan. 1: 682 (1878). guayaquilensis var. Candollei O. Ktze. Rev. Gen. 3(2): 36 (1898). Harmsii Rusby in Bull. N. Y. Bot. Gard. 4: 338 (1907). longifolia C. DC. in Meded. Rijks Herbar. 27: 82 bot multifoliola C. DC. in Meded. Rijks Herbar. 27: 82 (19 pachypoda (Rusby) C. DC. ex Harms in Engl. & ‘aid ee Pflanzenfam. (ed. 2) 19b: 114 (1940). . pauciflora Rusby in Mem. N. Y. Bot. Gard. 7: 280 (1927). sexanthera C. DC. in Notizbl. 6: 501 (1917). Steinbachii Harms in Notizbl. 9: 1156 (1927). stellato-tomentosa O. Ktze. Rev. Gen. 3(2): 36 (1898). stellipila C. DC. in Bull. Herb. Boiss. (ser. 2) 3: 412 (1903) = T. stellato- tomentosa. subarborescens C. DC. in Fedde, Repert. Spec. Nov. 7: 58 (1909). ensis C. DC. in Meded. Rijks Herbar. 27: 81 (1915). Trollii Harms in Notizbl. 11: 385 (1932). viridis var. puberula Lingelsh. in Fedde, Repert. Spec. Nov. 8: 1 (1910). MALPIGHIACEAE Aemanthera Radlkoferi O. Ktze. Rev. Gen. 3(2): 27 (1898) = Bunchosia Lin- deniana var. boliviensis. Aspicarpa argentea var. tenuifolia Niedenzu in Engler, Pflanzenr. IV, 141: 557 (1928). A. A. boliviensis Ndzu. in Meded. Rijks Herbar. 19: 72 (1913). lanata (Chod.) Ndzu. in Verz. Vorles. Akad. Braunsberg, 1912-13: 59 (1912). A. sericea Griseb. in Goett. Abh. 24: 68 (1879). Banisteria argentea var. acuminata forma eglandulosa Ndzu. in Engler, od m When fw BD bt ob oD Pflanzenr. IV, 141: 486 (1928 ) . argentea var. obtusiuscula forma glandulifera Ndzu. in Engler, Pflanzenr. IV, 141: 436 (1 “sigh . atrosanguinea Juss. in Ann. Sci. Nat. (ser.2) 13: 281 (1840). cinerascens var. slats alle Ndzu. in Engler, Pflanzenr. IV, 141: 406 (1928). cinerea Rusby in Bull. N. Y. Bot. Gard. 4: 335 (1907) = B. Gardneriana ? cristata Griseb. in Linnaea, 22: 16 (1849). crotonifolia Juss. in St. Hil. Fl. Bras. Mer. 3: 36 (1832). neriana Juss. in Arch. Mus. Paris, 3: 421 (1843). illustris (Rusby) Ndzu. in Engler, Pflanzenr. IV, 141: 451 (1928). lutea Griseb. in Linnaea, 22: 15 (1849 metallicolor var. aurea forma eglandulosa Ndzu. in Engler, Pflanzenr. IV, 141: 434 (1928). metallicolor var. aurea forma glandulifera Ndzu. in Engler, Pflanzenr. IV, 141: 434 (1928). metallicolor var. subrotunda forma glandulifera Ndzu. in Engler, Pflan- zenr. IV, 141: 433 (1928). metallicolor var. subsalicina forma eglandulosa Ndzu. in Engler, Pflan- zenr. IV, 141: 485 (1928). muricata Cav. Diss. 9: 423, t. 246 (1790). nigrescens Juss. in St. Hil. Fl. Bras. Mer. 3: 44 (1882). nitrosiodora Griseb. in Goett. Abh. 24: 65 (1879). 109 oxyclada Juss. in Arch. Mus. Paris, 3: 396 (1848). oxyclada forma brevialata Ndzu. De Gen. Banist. pt. 1: 26 (1900). ewetinite var. boliviensis Ndzu. in Fedde, Repert. Spec. Nov. 26: 3845 (1929). Pearcei Rusby in Bull. N. Y. Bot. Gard. 4: 884 (1907) = B. argentea. pubipetala Juss. in St. Hil. Fl. Bras. Mer. 3: 41, t. 169 (183 oe Rusbyana Ndzu. in Ind. Lect. Lye. Brawisbery, 1901: 19 (19 sanguinea Rusby in Bull. N. Y. Bot. Gard 4: 835 (1907) = sii le Beecheyana var. andina. Sellowiana var. Blanchetiana (Juss.) Ndzu. in Ind. Lect. Lyc. Braunsberg p. hiem. 1900-1901: 8 (1900). Sellowiana var. Blanchetiana forma grandifolia Ndzu. in Engler, Pflan- zenr. IV, 141: 409 (1928). Spruceana Griseb. in Mart. Fl. Bras. 12(1): 45 (1858). sublucida (Rusby) Ndzu. in Engler, Pflanzenr. IV, 141: 452 (1928). Whitei (Rusby) Ndzu. in Engler, Pflanzenr. IV, 141: 453 (1928). B. Williamsii (Rusby) Ndzu. in Engler, Pflanzenr. IV, 141: 458 (1928). Brittonella pilosa Rusby in Bull. Torr. Bot. Club, 20: 480 (1893) —= Mionandra camareoides. Bunchosia angustifolia forma parvifolia Ndzu. in Meded. Rijks Herbar. 19: 75 (19138). we ow wy SPO ee . armeniaca (Cav.) Rich. in A. Juss. in Ann. Mus. Paris, 18: 481 (1811), com- bination implied, but not actually made. armeniaca forma systyla Ndzu. in Arb. Bot. Inst. Akad. Braunsberg, 5: 42 (1914). lanceolata Turez. in Bull. Soc. Nat. Mosc. 36(1): 582 (1863). Lindeniana no boliviensis Ndzu. in Ind. Lect. Lye. Braunsb. p. hiem. 1898- 99: 10 (1898). pilocarpa Rusby in Bull. N. Y. Bot. Gard. 4: 333 (1907) = B. armeniaca. yrsonima biacuminata Rusby in Mem. Torr. Bot. Club, 6: 13 (1896) = B. coriacea var. spicata forma angustifolia. coccolobifolia HBK. Nov. Gen. & Spec. 5: 148 (1822). coriacea var. spicata (Cav.) Ndzu. in Engler, Pflanzenr. IV, 141: 700 (1928). coriacea var. spicata forma angustifolia (Benth.) Ndzu. in Engler, Pflan- zenr. IV, 141: 701 (1928). crassifolia (L.) HBK. Nov. Gen. & Spec. 5: 149 (1822). cydoniaefolia Juss. in St. Hil. Fl. Bras. Mer. 3: 77 (1832). cydoniaefolia var. chiquitensis Juss. in Arch. Mus. Paris, 3:281 (1843). laevigata (Poir.) DC. Prodr. 1: 580 (1824). Orbignyana Juss. in Ann. Sci. Nat. (ser. 2) 13:332 (1840). Poeppigiana var. velutina Ndzu. in Engler, Pflanzenr. IV, 141: 715 (1928). spicata (Cav.) Rich. ex Juss. in Ann. Mus. Paris, 18: 481 (1812), combina- tion implied, but not actually made. variabilis Juss. in St. Hil. Fl. Bras. Mer. 3: 78 (1832). Dicella Conwayi Rusby in Bull. N. Y. Bot. Gard. 8: 98 (1912). D. macroptera (Mart.) Juss. in Ann. Sci. Nat. (ser. 2) 18: 323 (1840). Galphimia brasiliensis (L.) Juss. in St. Hil. Fl. Bras. Mer. 3: 71, t. 178 (1832). Heteropterys aceroides forma Grisebachiana subf, elongata Ndzu. in Arb. Bot. Inst. Lyc. Hos. 2: 34 (1903). H. anoptera var. eglandulosa subvar. latifolia Ndzu. in Arb. Bot. Inst. Lyc. Hos. 2: 49 (1903). H. anoptera var. glandulifera subvar. ovata Ndzu. in Arb. Bot. Inst. Lyc. Hos. 2: 48 (1903). H. Beecheyana var. andina Ndzu. in Arb. Bot. Inst. Lyc. Hos. 2: 55 (1903). H. bopiana (Rusby) Ndzu. in Engler, Pflanzenr. IV, 141: 377 (1928). wh Dh wf DOP Wm we wm _ = =) H. campestris Juss. in St. Hil. Fl. Bras. Mer. 3: 33 (1832). H. canaminensis (Rusby) Ndzu. in Engler, Pflanzenr. IV, 141: 878 (1928). H. cochleosperma forma glandulifera Ndzu. in Arb. Bot. Inst. Lyc. Hos. 2: 11 (1903). H. dumetorum (Griseb.) Ndzu. in Engler, Pflanzenr. IV, 141: 336 (1928). H. falcifera Juss. in Ann. Sci. Nat. (ser. 2) 13: 274 (1840) H. hypericifolia Juss. in St. Hil. Fl. Bras. Mer. 3: 24 (1832). H. macrostachya A. Juss. in Arch. Mus. Paris, 3: 450 (1843). H. ovalifolia Rusby in Bull. N. Y. Bot. Gard. 4: 334 (1907) — H. falcifera. H. rufula forma hirta Ndzu. in Fedde, Repert. Spec. Nov. 26: 345 (1929). H. sphaerandra (Rusby) Ndzu. in Engler, Pflanzenr. IV, 141: 380 (1928). H. suberosa var. Candolleana (Juss.) Ndzu. in Arb. Bot. Inst. Lyc. Hos. 2: 51 (1903). suberosa var. Lessertiana (Juss.) Griseb. in Mart. Fl. Bras. 12(1): 69 (1858). sylvatica Juss. in Ann. Sci. Nat. (ser. 2) 13: 277 (1840). syringifolia Griseb. in Linnaea, 13: 224 (1830). syringifolia var. Pilgeri Ndzu. in Arb. Bot. Inst. Lyc. Hos. 2: 20 (1903). tenuifolia (Ndzu.) Ndzu. in Engler, Pflanzenr. IV, 141: 333 (1928). tomentosa Juss. in St. Hil. Fl. Bras. Mer. 3: 31 (1832). trichanthera A. Juss. in Arch. Mus. Paris, 3: 438 (1843). trigoniifolia forma glandulifera Ndzu. in Arb. Bot. Inst. Lyc. Hos. 2: 8 3) . umbellata Juss. in St. Hil. Fl. Bras. Mer. 3: 25, t. 166 (1832). iraea chlorocarpa A. Juss. in Ann. Sci, Nat. (ser. 2) 13: 259 (1840). fagifolia var. Blanchetiana forma longifolia Ndzu. in Verz. Vorles. Lyc. Hos. W.-S. 1906-07: 14 (1906). H. fagifolia var. Blanchetiana forma longifolia subf. parvifolia Ndzu. in ed. Rijks Herbar. 19: 71 (1918). H. Jussieana Mig. in Linnaea, 19: 142 (1847). H. Kunthiana Juss. in Arch. Mus. Paris, 3: 571 (1848). H. strigulosa Rusby in Bull. N. Y. Bot. Gard. 8: 95 (1912). H. transiens Ndzu. in Verz. Vorles. Lyc. Hos. W.-S. 1906-07: 8 (1906). Janusia guaranitica (St. Hil.) Juss. in Ann. Sci. Nat. (ser. 2) 13: 251 (1840). J. guaranitica var. vulgaris forma sericans Ndzu. in Verz. Vorles. Akad. Braunsberg im W.-S. 1912-18: 49 (1912). Jubistylis mollis Rusby in Mem. N. Y. Bot. Gard. 7: 274, fig. 4 (1927) = Banis- teria cristata. Mascagnia anisopetala (Juss.) Griseb. in Mart. Fl. Bras. 12(1): 95 (1858). M. brevifolia Griseb. in Goett. Abh. 24: 67 (1879). brevifolia var. paniculata Ndzu. in Arb. Bot. Inst. Lye. Braunsberg, 3: 15 (1908). . chlorocarpa var. cristata Ndzu. in Arb. Bot. Inst. Lyc. Braunsberg, 3: 21 Hm eee s 4 (1908). M. ixiamensis Rusby in Mem. N. Y. Bot. Gard. 7: 271 (1927) — M. ovatifolia. M. macrophylla Rusby in Mem, N. Y. Bot. Gard. 7: 272 (1927). M. ovatifolia (HBK.) Griseb. Fl. Brit. W. Ind. 121 (1864). M. ovatifolia forma cordata Ndzu. in Arb. Bot. Inst. Lye. Braunsberg, 3: 13 (1908). M. pachyptera Rusby in Mem. N. Y. Bot. Gard. 7: 271 (1927) = M. psilophylla. M. psilophylla (Juss.) Griseb. in Mart. Fl. Bras, 12(1): 94 (1858). M. rigida (Juss.) Griseb. in Mart. Fl. Bras. 12(1): 92 (1858). M. rigida subsp. coriacea (Griseb.) Ndzu. in Arb. Bot. Inst. Lyc. Braunsberg, 3: 19 (1908). M. sepium var. rufescens Griseb. in Mart. Fl. Bras. 12(1): 96 (1858). 111 M. sericans var. boliviensis Ndzu. in Arb. Bot. Inst. Lyc. Braunsberg, 3: 21 (1908). Mionandra camareoides Griseb. in Goett. Abh. 19: 102 (1874). Peixotoa reticulata Griseb. in Linnaea, 13: 213 (1839). Ptilochaeta nudipes Griseb. in Goett. Abh. 24: 66 (1879). Stigmatophyllum bogotense Tri. & Pl. in Ann. Sci. Nat. (ser. 4) 18: 320 1862). S. calearatum N. E. Brown in Trans. Bot. Soc. Edinb. 20: 48 (1894). S. coloratum Rusby in Mem. Torr. Bot. Club, 6: 14 (1896). S. tiliaefolium (HBK.) Ndzu. De Gen. Stigmat. pt. 2: 16 (1900). S. tomentosum Juss. in St. Hil. Fl. Bras. Mer. 3: 53 (1832). Tetrapterys acapulcensis HBK. Nov. Gen. & Spec. 5: 168 (1821). mbigua var. paraguayensis (Ndzu.) Ndzu. in Engler, Pflanzenr. IV, 141: 169 (1928). . boliviensis Ndzu. in Verz. Vorles. Lyc. W.-S. 1909-10: 17 (1909). calophylla var. boliviana Rusby in Mem. Torr. Bot. Club, 6: 14 (1896). calophylla var. glabrior Ndzu. in Verz. Vorles. Lyc. W.-S. 1909-10: 47 discolor (G. Meyer) DC. Prodr. 1: 587 (1824). discolor var. andina Ndzu. in Verz. Vorles. Lyc. W.-S. 1909-10: 42 (1909). elliptica Rusby in Bull. N. Y. Bot. Gard. 8: 96 (1912). mucronata var. boliviensis Ndzu. in Verz. Vorles. Lyc. W.-S. 1909-10: 55 (1909). multiglandulosa Juss. in Ann. Sci. Nat. (ser. 2) 13: 264 (1840). suaveolens Juss. in Ann. Sci. Nat. (ser. 2) 13: 263 (1840). ryallis latifolia var. ovatifolia (Ndzu.) Ndzu. in Arb. Bot. Inst. Akad. Braunsberg, 5: 12 (1914). HA Sxas Ans TRIGONIACEAE Trigonia bicolor Suesseng. & Overk. in Fedde, Repert. Spec. Nov. 51: 204 (1942). boliviana Warm. in Mart. Fl. Bras. 138(2): 134 (1875). echitifolia Rusby in Bull. N. Y. Bot. Gard. 4: 324 (1907). floccosa Rusby in Bull. N. Y. Bot. Gard. 4: 325 (1907). parviflora Benth. in Hook. Journ. Bot. 3: 163 (1851). . pubescens Camb. in St. Hil. Fl. Bras. Mer. 2: 114 (1829). simplex var. pilosula O. Ktze. Rev. Gen. 3(2): 12 (1898). HHaabH VOCHYSIACEAE Callisthene fasciculata Mart. Nov. Gen. & Spec. 1: 126, t. 100 (1824). Erisma calcaratum (Link) Warm. in Mart. FI. Bras. 13(2): 111 (1875). Qualea acuminata Spruce ex Warm. in Mart. Fl. Bras. 13(2): 40 (1875). . grandiflora Mart. Nov. Gen. & Spec. 1: 133, t. 79 (1826). multiflora subsp. pubescens (Mart.) Stafleu in Act. Bot. Neerland. 2: 196 (1953). . parviflora Mart. Nov. Gen. & Spec. 1: 135, t. 81 (1826). pilosa var. heterophylla O. Ktze. Rev. Gen. 3(2): 11 (1898). pilosa var. multinervia O. Ktze. Rev. Gen. aif 11 (1898). Tessmannii Mildbr. in Notizbl. 9: 141 (1924). virgata Rusby in Bull, N. Y. Bot. Gard. 8: 99 (1912) = Q. multiflora subsp. pubescens. Salvertia convallariodora St. Hil. in Mém. Mus. Paris, 6: 266 (1820). Vochysia boliviana Rusby in Mem. Torr. Bot. Club, 3(3): 7 (1898). V. caesia Stafleu in Rec. Trav. Bot. Néerl. 41: 488 (1948). 2000 OO divergens Pohl, Fl. Bras. 2: 19, t. 3 (18381). Haenkeana Mart. Nov. Gen. & Spec. 1: 147, t. 89 (1826). Leguiana Macbr. in Field Mus. Publ. Bot. 8: 121 (1930). mapirensis Rusby in Mem. Torr. Bot. Club, 6: 7 (1826). Radlkoferi O. Ktze. Rev. Gen. 3(2): 12 (1898). rufa Mart. Nov. Gen. & Spec. 1: 144, t. 86 (1826). A ASASS POLYGALACEAE Bredemeyera densiflora A. W. Bennett in Mart. Fl. Bras. 13(2): 52 (1875). B. floribunda Willd. in Ges. Naturf. Freunde Neue Schr. 3: 412 (1801). Monnina aestuans (L. f.) DC. Prodr. 1: 338 (1824). M. alatodrupacea O. Ktze. Rev. Gen. 3(2): 10 (1898). M. Arbutus Chod. in Meded. Rijks Herbar. 27: 30 (1915). M. Autraniana Chod. in Bull. Herb. Boiss. (ser.1) 3: 543 (1895). M. Bangii Chod. in Bull. Herb. Boiss. (ser. 1) 4: 243 (1896). M. boliviana Chod. in Bull. Herb. Boiss. (ser. 1) 3: 543 (1895). M. boliviensis A. W. Bennett in Bull. Torr. Bot. Club, 16: 20 (1889). M. brachystachya Griseb. in Goett. Abh. 19: 75 (1874). M. Bridgesii Chod. in Bull. Herb. Boiss. (ser. 1) 3: 542 (1895). M. Buchtienii Rusby, Descr. S. Am. Pls. 41 (1920). M. eriocarpa Chod. in Meded. Rijks Herbar. 27: 29 (1915). M. Franchetii Chod. in Bull. Herb. Boiss. (ser. 1) 4: 248 (1896). M. gracilis Chod. in Bull. Herb. Boiss. (ser. 1) 4: 244 (1896). M. Hassleri Chod. in Engler, Bot. Jahrb. 42: 100 (1908). M. herbacea DC. Prodr. 1 M. Herzogii Chod. in Meded. Rijks Herbar. 27: 31 (1915). M. Laureola Chod. in Bull. Herb. Boiss. (ser. 1) 4: 245 (1896). M. macroclada Chod. in Bull. Herb. Boiss. (ser. 1) 4: 246 (1896). M. macrostachya var. stenophylla O. Ktze. Rev. Gen. 3(2): 10 (1898). M. nigrescens Rusby in Bull. N. Y. Bot. Gard. 4: 324 (1907). M. Pearcei Chod. in Bull. Herb. Boiss (ser. 1) 4:243_ (1896). M. pseudostipulata Chod. in Bull. Herb. Boiss. (ser. 1) 4: 245 (1896). M. resedoides St. Hil. Fl. Bras. Mer. 2: 61 (1829). M. — HBK. Nov. Gen. & Spec. 5: 415 (1822). M. Rusbyi Chod. in Rusby in Mem. Torr. Bot. Club, 6: 6 (1896). M. nies R. & P. Syst. 172 (1798). M. stipulata Chod. in Bull. Herb. Boiss. (ser. 1) 2: 170 (1894). M. Weddelliana Chod. in Bull. Herb. Boiss. (ser. 1) 4: 244 (1896). Polygala acuminata (Willd.) Chod. Monog. Polyg. 2: 46 (1893). P. andina A. W. Bennett in Bull. Torr. Bot. Club, 16: 19 (1889). P. angustifolia HBK. Nov. Gen. & Spec. 5: 405, t. 511 (1822). P. Bangiana Chod. in Bull. Herb. Boiss. (ser. 1) 4: 234 (1896). P. boliviensis A. W. Bennett in Journ. Bot. 17: 171 (1879). P. cisandina Chod. in Meded. Rijks Herbar. 27: 28 (1915). P. filiformis St. Hil. Fl. Bras. Mer. 2: 7 (1829) P. formosa A. W. Bennett in Bull. Torr. Bot. Club, 16: 19 (1889). P. gymnosepala Chod. in Meded. Rijks Herbar. 27: 26 (1915). P. hebeclada DC. Prodr. 1: 331 (1824). P. macerrima Blake in Proc. Biol. Soc. Wash. 48: 5 (1930). P. Mandonii Chod. in Engler, Bot. Jahrb. 42: 98 (1908). P. molluginifolia St. Hil. Fl. Bras. Mer. 2: 25 (1829). P. monodonta Chod. in Meded. Rijks Herbar. 27: 28 (1915). P. myurus Chod. in Meded. Rijks Herbar. 27: 27 (1915). P. nemoralis A. W. Bennett in Journ. Bot. 17: 172 (1879). 1138 P. paludosa var. angustocarpa Chod. in Mém., Soc. Phys. Hist. Nat. Genéve, 30 (pt. 2, no. 8): 106 (1889). P. paniculata L. Sp. Pl. (ed. 2) 2: 987 (1763). P. Pearcei A. W. Bennett in Journ. Bot. 17: 201 (1879). P. spectabilis var. minor Chod. ex Rusby in Bull. N. Y. Bot. Gard. 4: 323 (1907). Securidaca volubilis L. Sp. Pl. 707 (1753). EUPHORBIACEAE Acalypha alchorneoides Rusby in Bull. N. Y. Bot. Gard. 8: 101 (1912). odonta var. villosa Muell. Arg. in Mart. Fl. Bras. 11(2): 366, t. 53 (1874 arvensis : Poepp. & Endl. Nov. Gen. 3: 21 (1845 Baenitzii Pax in Fedde, Repert. Spec. Nov. 5: 227 (1908) = A. stenoloba. benensis Britton ex Rusby in Bull. Torr. Bot. Club, 28: 304 (1901) boliviensis Muell. Arg. in Linnaea, 34: 162 (1865). bopiana Rusby in Mem. N. Y. Bot. Gard. 7: 287 (1927). Brittonii Rusby in Bull. Torr. Bot. Club, 28: 303 (1901). Buchtienii Pax in Fedde, Repert. Spec. Nov. 5: 227 (1908). callosa Benth. Pl. Hartw. 252 (1846). capillaris Rusby in Mem. Torr. Bot, Club, ohh 257 (1895) = A. stenoloba. communis Muell. Arg. in Linnaea, 34: 23 communis var. guaranitica Chod. & Hassl. in Bull. Herb. Boiss. (ser. 2) 5: 605 (1905). communis var. hirta (Spreng.) Muell. Arg. in Linnaea, 34: 24 (1865). communis var. tomentella Muell. Arg. in Linnaea, 34: 24 (1865). controversa (O. Ktze.) K. Schum. in Just, Bot. Jahresber. 26(1): 348 (1898). cuprea Herzog in Fedde, Repert. Spec. Nov. 7: 60 ( 1909) cuspidata Jacq. Hort. Schoenbr. 2: 63, t. 243 (1797). diversifolia Jacq. Hort. Schoenbr. 2: 63, t. 244 (1797). diversifolia var. carpinifolia (Poepp.) Muell. Arg. in DC. Prodr. 15(2): 854 (1866). Douilleana Rusby in Mem. N. Y. Bot. Gard. 7: 285 (1927). erosa Rusby in Bull. Torr. Bot. Club, 28: 305 (1901). eugenifolia Rusby in Bull. N. Y. Bot. Gard. 4: 443 (1907). flabellifera Rusby in Mem. Torr. Bot. Club, 6: 119 (1896) = A. plicata. . foliosa Rusby in Bull. N. Y. Bot. Gard. 4: 443 (1907). . grandispicata Britton ex Rusby in Bull. Torr. Bot. Club, 28: fs (1901). % ‘Herzogiana Pax & Hoffm. in Meded. Rijks Herbar. 40: 24 (1921). A. heteromorpha Rusby in Mem. N. Y. Bot. Gard. 7: 286 (1927). A. hibiscifolia Britton ex Rusby in Mem. Torr. Bot. Club, 4(3): 257 (1895). A. inaequalis Rusby in Bull. Torr. Bot. Club, 28: 303 (1901). jubifera Rusby, Descr. S. Am. Pls. 48 (1920). Lechleri Britton ex Rusby in Bull. Torr. Bot. Club, my 304 (1901). lucida Rusby in Bull. N. Y. Bot. Gard. 4: 444 (190 lycioides Pax. & Hoffm. in Meded. Rijks Herbar. ng “24 (1921). macrophylla Ule in Verh. Bot. Ver. Brandenb. 50: 79 (1908). macrostachya Jacq. Hort. Schoenbr. 2: 63, t. 245 (1797). macrostachya var. sidaefolia (HBK.) Muell. Arg. in Linnaea, 34: 11 (1865). macrostachya var. tristis Muell. Arg. in Mart. Fl. Bras. 11(2): 345 (1874). Mandonii Muell. Arg. in Linnaea, 34: 162 (1865-66). mapirensis Pax in Fedde, Repert. Spec. Nov. 7: 110 (1909). mapirensis var. pubescens Pax & Hoffm. in Engler, Pflanzenr. IV, 147 PPP PP pap PPP RPP PPPP PPh PPB: ihc ghp . Nitschkeana Pax & Hoffm. in Engler, Pflanzenr. IV, 147 (xvi): 88 (1924). ovata Pax & Hoffm. in Meded. Rijks Herbar. 40: 23 (1921) = A. stenoloba. D> PPP PD PD — ar ~~ paupercula Pax & Hoffm. in Meded. Rijks Herbar. 40: 24 (1921). plicata Muell. Arg. in DC. Prodr. 15(2): 855 (1866). Poiretii Spreng. Syst. Veg. 3: 879 (1826). soratensis Pax & Hoffm. in Engler, Pflanzenr. IV, 147 (xvi): 126 (1924). stachyura Pax in Fedde, Repert. Spec. Nov. 7: 110 (1909) = A. macro- ie Samed phylla. stenoloba Muell. Arg. in Flora, 55: 41 (1872). variegata Rusby in Mem. N. Y. Bot. Gard. 7: 285 (1927). vermifera Rusby in Mem. N. Y. Bot. Gard. 7: 286 (1927) = A. diversifolia. silteas Jacq. Enum. Pl. Carib. 32 (1760). villosa var. latiuscula Pax & Hoffm. in Engler, Pflanzenr. IV, 147 (xvi): 17 Williamsii Rusby in Bull. N. Y. Bot. Gard. 8: 101 (1912). Williamsii Rusby, Descr. S. Am. Pls. 47 (1920). Ichornea castaneifolia var. salicifolia (Baill.) Baill. in Adansonia, 5: 238 (1865). bie: fee iricurana forma pubescens (Britton) Pax & Hoffm. in Engler, Pflanzenr. IV, 147 (vii): 233 (1914). latifolia Sw. Prodr. 98 (1788). megalostylis Rusby in Phytologia, 1: 63 (1934). Pearcei Britton ex Rusby in Bull. Torr. Bot. Club, 28: 305 (1901). sclerophylla Pax in Fedde, Repert. Spec. Nov. 7: 242 (1909). triplinervia (Spreng.) Muell. Arg. in DC. Prodr. 15(2): 909 (1866). triplinervia var. boliviana Pax &,Hoffm. in Engler, Pflanzenr. IV, 147 (vii): 229 (1914). . triplinervia var. janeirensis (Casar.) Muell. Arg. in DC. Prodr. 15(2): 909 (1866). Amanoa muricata Rusby in Bull. N. Y. Bot. Gard. 8: 100 Npulel Aparisthmium cordatum (Juss.) Baill. in Adansonia, 5: 307 (1865). Apodandra Buchtienii (Pax) Pax in Engler, Pflanzenr. IV, 147 (ix): 21 (1919). Astrocasia sp. Bernardia paraguariensis var. orbiculata Chod. & Hassl. in Bull. Herb. Boiss. er. 2) 5: 504 (1905). B. rotundifolia Herzog in Fedde, Repert. Spec. Nov. 7: 59 (1909). Caperonia castaneifolia (L.) St. Hil. Pl. Remarg. Brésil, 245 (1824). Chaetocarpus Pearcei Rusby in Bull. N. Y. Bot. Gard. 8: 102 (1912). Chiropetalum boliviense (Muell. Arg.) Pax & Hoffm. in Engler, Pflanzenr. IV, 147 (vi): 94 (1912). Cleidion amazonicum Ule in Verh. Bot. Ver. Brandenb. 50: 76 (1908). C. tricoccum (Casar.) Baill. in Adansonia, 4: 370 (1863-64). Cnidoscolus tubulosus var. trilobus (Muell. Arg.) Lourt. & O’Don. in Lilloa, 9: 118 (1948). Conceveiba guianensis Aubl. Pl. Guian, Fr. 2: 924, t. 353 (1775). Croton abutilifolius Croiz. in Darwiniana, 6: 448 (1944). . andinus Muell. Arg. in Linnaea, 34: 126 (1865). . apicifolius Croiz. in Darwiniana, 6: 447 (1944). avulsus Croiz. in Darwiniana, 6: 450 44). Bangii Rusby in Mem. Torr. Bot. Club, 4(3): 256 (1895). boliviensis Muell. Arg. in Linnaea, 34: 91 (1895) Bonplandianus Baill. in Adansonia, 4: 339 (1864). Bridgesti Muell. Arg. in Linnaea, aE 115 (1865) = C, Orbignyanus. Buchtienii Rusby in Mem. N. Y. Bot. Gard. 7: 284 (1927). caladiifolius Croiz. in Journ. ena Arb. 21: 96 (1940) = C. peltophorus. callicarpifolius var. pubescens Muell. Arg. in Linnaea, 34: 85 (1865). Cardenasii Standl. in Field Mus. Publ. Bot. 22: 34 (1940). C. chamaedryfolius Lam. ex Griseb. Fl. Brit. W. Ind. 41 (1859). a ofl ue eka é > anmaagaeagaaaa charaguensis Standl. in Field Mus. Publ. Bot. 22: 35 (1940). densiflorus Pax & Hoffm. in Meded. Rijks Herbar. 40: 19 (1921). emporiorum Croiz. in Journ. Arnold Arb. 21: 97 (1940). erythrochyloides Croiz. in Darwiniana, 6: 449 (1944). Frieseanus Muell. Arg. in DC. Prodr. 15(2): 543 (1866). glandulosus L. Syst. (ed. 10) 1275 (1759). Hieronymi Griseb. in Goett. Abh. 24: 54 (1879). laeticapsulus Croiz. in Darwiniana, 6: 448 (1944). lobatus L. Sp. Pl. 1005 (1753). Mandonis Muell. Arg. in Linnaea, 34: 85 (1865). matourensis Aubl. Pl. Guian. Fr. 2: 879, t. 338 (1775). nudulus Croiz in Journ. Arnold Arb. 21: 98 (1940). Orbignyanus Muell. Arg. in Linnaea, 34: 123 (1865-66). pedicellatus HBK. Nov. Gen. & Spec. 2: 75 (1817). peltophorus Muell. Arg. in Mart. Fl. Bras. 11(2): 154 (1873). perintricatus Croiz. in Darwiniana, 6: 450 (1944). piluliferus Rusby in Mem. Torr. Bot. Club, 4(3): 256 (1895). pseudogracilipes Pax & Hoffm. in Meded. Rijks Herbar. 40: 20 (1921) = C. peltophorus. pungens Jacq. Ic. Pl. Rar. 3: 19, t. 622 (1794). rhamnifolius var. boliviensis Pax & Hoffm, in Meded. Rijks Herbar. 40: 21 (1921) = C. Orbignyanus. roborensis Standl. in Field Mus. Publ. Bot. 22: 36 (1940). Rusbyi Britton in Rusby in Mem. Torr. Bot. Club, 6: 119 (1898). sarcopetalus var. longipetiolatus Pax & Hoffm. in Meded. Riiks Herbaz, 40: 21 (1921). Sellowii Baill. in Adansonia, 4: 304 (1864). soratensis Muell. Arg. in Linnaea, 34: 115 (1865-66). soratensis var. intermedius Muell. Arg. in Linnaea, 34: 116 (1865-66). soratensis var. leptobotryus Muell. Arg. in Linnaea, 34: 116 (1865-66). soratensis var. pycnanthus Muell. Arg. in Linnaea, 34: 116 (1865-66). tartonrairoides Pax & Hoffm. in Meded. Rijks Herbar. 40: 20 (1921). triqueter Lam. Encye. 2: 214 (1786). - Urucurana Baill. in Adansonia, 4: 335 (1863-64). C. Williamsii Rusby in Bull. N. Y. Bot. Gard. 8: 100 (1912) = C. peltophorus. C. yungensis Croiz. in Journ. Arnold Arb. 21: 103 (1940). Dalechampia adscendens (Muell. Arg.) Muell. Arg. in Mart. Fl. Bras. 11(2): 640 (1874). albibracteosa Rusby in Mem. N. Y. Bot. Gard. 7: 287 (1927). Bangii Pax & Hoffm. in Engler, Pflanzenr. IV, 147 (xii): 24 (1919). boliviana Pax & Hoffm. in Engler, Pflanzenr. IV, 147 (xii): 50 (1919). boliviana Gandog. in Bull. Soc. Bot. France, 66: 286 (1920). Burchellii Muell. Arg. in Mart. Fl. Bras. 11(2): 649 (1874). cujabensis Muell. Arg. in Linnaea, 34: 222 (1865-66). Herzogiana Pax & Hoffm. in Engler, Pflanzenr. IV, 147 (xii): 36 (1919). humilis var. adscendens Muell. Arg. in Linnaea, 84: 224 (1865-66). scandens L. Sp. Pl. 2: 1054 (1753). scandens var. fimbriata Muell. Arg. in DC. Prodr. 15(2): 1244 (1866). Ditaxis breviramea (Muell. Arg.) Pax & Hoffm. in Engler, Pflanzenr. IV, 147(vi): 65 (1912). Euphorbia acerensis Boiss. in DC. Prodr. 15(2): 55 (1862). E. boerhaavioides Rusby in Bull. N. Y. Bot. Gard. 4: 441 (1907). E. boliviana Rusby in Bull. N. Y. Bot. Gard. 4: 442 (1907). E. caecorum Mart. ex Boiss. in DC. Prodr. 15 (2): 51 (1862). E. Chamaesyce L. Sp. Pl. 455 (1753). E. cymbiformis Rusby in Mem. Torr. Bot. Club, 4(3): 255 (1895). angaaaaa aan aan aannanaaaaaaanaaaagaa . cca dad dicelt nd my 116 PS SEE ROBB! i bb be dentata Michx. Fl. Bor. Am, 2: 211 (1803). duriuscula Pax & Hoffm. in Blumea, 5(3): 641 (1945). eanophylla Croiz. in Rev. Sudam. Bot. 6: 10 (19389). geniculata Orteg. in Hort. Matrit. Dec. 2: 18 (1797). Huanchahana var. peperomioides Croiz. in Journ. Arnold Arb. 24: 179 (1943). hypericifolia L. Sp. Pl. 454 (1753). hyssopifolia L. Syst. (ed. 10) 2: 1048 (1759). insulana subsp. tovarensis (Boiss.) Croiz. in Journ. Arnold Arb. 24: 178 (1948). lasiocarpa Klotzsch in Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 414 (1843). lasiocarpa var. subprostrata Boiss. in DC. Prodr. 15(2): 1262 (1866). Lathyrus L. Sp. Pl. 457 (1753) longipila Rusby in Bull. N. Y. Bot. Gard. 4: 442 (1907) = E. acerensis. Mandoniana Boiss. in DC. Prodr. 15(2): 1264 (1866). Meyeniana Klotzsch in Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 414 (1848). nutans Lag. Gen. & Spec. Nov. 17 (1816). orbiculata HBK. Nov. Gen. & Spec. 2: 52 (1817). Peplus L. Sp. Pl. 456 (1753). pilulifera L. Sp. Pl. 454 (1753). Poeppigii (Kl. & Gke.) Boiss. in DC. Prodr. 15(2): 56 (1862). portulacoides L. Sp. Pl. 456 (1753). Preslii var. andicola Danguy & Cherm. in Bull. Mus. Hist. Nat. Paris, 28: 438 (1922 prostrata Ait. Hort. Kew. 2: 188 (1789). prunifolia Jacq. Hort. Schoenbr. 3: 15, t. 277 (1798). . serpens HBK. Nov. Gen. & Spec. 2: 52 (1817) Hevea brasiliensis (Willd.) Muell. Arg. in Linnaea, 34: 204 (1865-66). H. Spruceana Muell. Arg. in Linnaea, 34: 204 (1865-66). Hieronyma andina Pax & Hoffm. in Engler, Pflanzenr. ou od (xv): 37 (1922). H. boliviana Pax in Fedde, Repert. Spec. Nov. 7: 109 (19 br mm . Buchtienii Pax & Hoffm. in Engler, Pflanzenr. IV, ¥ ee 33 (1922). Moritziana var. yungasensis Pax & Hoffm. in Bugler, Pflanzenr. IV, 147 (xv): 88 (1922). oblonga (Tul.) Muell. Arg. in Linnaea, 34: 66 (1865-66). reticulata (Pl.) Britton ex Rusby in Mem. Torr. Bot. Club, 4(3): 255 (1895). Hura crepitans L. Sp. Pl. 1008 (1753). Jatropha clavuligera Muell. Arg. in Linnaea, 34: 209 (1865-66). Sa Sy Sy ae not St Hon a J. . Curecas L. Sp. Pl. 1006 (1753). . elliptica (Pohl) Muell. Arg. in Mart. Fl. Bras. 11(2): 489 (1874). gossypifolia var. staphysagrifolia (Mill.) Muell. Arg. in DC. Prodr. 15(2): 1087 (1866 grossidentata Pax & Hoffm. in Engler, Pflanzenr. IV, 147(vii): 398 (1914). Hieronymi O. Ktze. Rev. Gen. 3(2): 287 (1898). intercedens Pax in Engler, Pflanzenr. IV, 147: 31 (1910). pachypoda Pax in Engler, Pflanzenr. IV, 147: 47 (1910). papyrifera Pax & Hoffm. in Notizbl. 10: 385 (1928). pedatipartita O. Ktze. Rev. Gen. 3(2): 287 (1898). thyrsantha Pax & Hoffm. in Engler, Pflanzenr. [V, 147 (vii): 897 (1914). tubulosa var. triloba Muell. Arg. in Linnaea, 34: 212 (1865-66). Re (194 paniculatus Pax & Hoffm. in "Medea. Rijks Herbar. 40: 22 (1921). 117 J. peruvianus var. flavispicatus (Rusby) Croiz. in Rev. Argent. Agron. 10: 136 (1948). Mabea anadena Pax & Hoffm. in Engler, Pflanzenr. IV, 147 (xiv): 55 (1919). M. elegans Rusby in Mem. N. Y. Bot. Gard. 7: 288 (1927). M. fistulifera Mart. in Spix & Mart. Reise Bras. 2: 479 (1828). M. longifolia (Britton) Pax & Hoffm. in Engler, Pflanzenr. IV, 147 (v): 80 12). M. paniculata Spruce ex Benth. in Hook. Journ. Bot. 6: 367 (1854). Manihot boliviana Pax & Hoffm. in Engler, Pflanzenr. IV, 147(vii): 402 (1914). M. glabrata (Chod. & Hassl.) Pax & Hoffm. in Engler, Pflanzenr. IV, 147 (ii): M. Pavoniana Muell. Arg. in Linnaea, 34: 205 (1865-66). M. Rusbyi Britton in Bull. Torr. Bot. Club, 28: 302 (1901). M. utilissima Pohl, Pl. Bras. 1: 32, t. 24 (1827). Maprounea guianensis Aubl. Pl. Guian. Fr. 2: 895, t. 342 (1775). Margaritaria nobilis L. f. Suppl. 428 (1781). Omphalea diandra L. Syst. (ed. 10) 1264 (1759). Pachystroma ilicifolium var. longifolium Muell. Arg. in Linnaea, 34: 178 (1865-66). Pera benensis Rusby, Descr. S. Am. Pls. 49 (1920). P. elliptica Rusby in Mem. N. Y. Bot. Gard. 7: 288 (1927). Phyllanthus acuminatus Vahl, Symb. 2: 95 (1791). biflorus Rusby in Mem. N. Y. Bot. Gard. 7: 282 (1927). bolivianus Pax & Hoffm. in Meded. Rijks oes 40: 18 (1921). brasiliensis (Aubl.) Poir. Encyc. 5: 296 (18 caroliniensis Walt. Fl. Car. 228 (1788). cassioides Rusby in Bull. N. Y. Bot. Gard. 8: 100 (1912). graveolens var. glaber Pax & Hoffm. in Meded. Rijks Herbar. 40: 18 (1921). ibonensis Rusby in Mem. N. Y. Bot. Gard. 7: 281 (1927) = Margaritaria nobilis ichthyomethius Rusby in Mem. N. Y. Bot. Gard. 7: 282 (1927). inaequalis Rusby in Mem. Torr. Bot. Club, 6: 118 (1896) — Astrocasia sp. microphyllus var. Orbignyanus Muell. Arg. in Linnaea, 32: 45 (1863). Niruri subsp. lathyroides (HBK.) Webster in Contrib. Gray Herb. 176: 52 orbiculatus L. C. Rich. in Act. Soc. Hist. Nat. Paris, 1: 113 (1792). prunifolius Rusby in Mem. N. Y. Bot. Gard. 7: 283 (1927 pseudo-nobilis Rusby in Mem. N. Y. Bot. Gard. 7: 281 (1927) = = Margari- taria nobilis. P. rupestris HBK. Nov. Gen. & Spec. 2: 110 (1817). Plukenetia volubilis L. Sp. Pl. 1192 (1753). Ricinus communis L. Sp. Pl. 1007 (1753). Sapium bolivianum Pax & Hoffm. in Engler, Pflanzenr. IV, 147 (v): 221 (1912). S. haematospermum Muell. Arg. in Linnaea, 34: 217 (1865-66). S. montevidense Klotzsch ex Baill. in Adansonia, 5: 320 (1865), nomen mudwm VU VN Oe ee in synon. Ss. main Pax & Hoffm. in Engler, Pflanzenr. IV, 147 (v): 210 (1912). S. rhombifolium Rusby in Bull. Torr. Bot. Club, 28: 307 (1901). Sebastiania boliviana Rusby, Descr. S. Am. Pls. 50 (1920). S. brasiliensis Spreng. in Neue Entdeck. 2: 118, t. 3 (1821). S. brasiliensis var. divaricata (Muell. Arg.) Muell. Arg. in DC. Prodr. 15(2): 1187 (1866 S. brasiliensis var. ramosissima (St. Hil.) Muell. Arg. in DC. Prodr. 15(2): 1187 (1866). S. Bridgesii (Muell. Arg.) Pax in Engler, Pflanzenr. IV, 147 (v): 143 (1912). S. Fiebrigii Pax in Engler, Pflanzenr. IV, 147 (v): 142 (1912). 118 S. hispida var. paraguayensis (Chod. & Hassl.) Pax & Hoffm. in Engler, Pflanzenr. IV, 147 (v): 111 (1912). S. nervosa (Muell. Arg.) Muell. Arg. in DC. Prodr. 15(2): 1183 (1866). Stillingia salpingadenia (Muell. Arg.) Huber in Bull. Herb. Boiss. (ser. 2) 6: 452 (1906). Tragia aurea Rusby in Bull. N. Y. Bot. Gard. 4: 444 (1907). Bangii Rusby in Bull. N. Y. Bot. Gard. 4: 445 (1907). Fendleri Muell. Arg. in Linnaea, 34: 179 (1865-66). Friesii Pax & Hoffm. in Engler, Pflanzenr. IV, 147 (xvii): 186 (1924). geraniifolia Klotzsch ex Baill. Etude générale Euphorb. 461 (1858). oligantha Pax & Hoffm. in Engler, Pflanzenr. IV, 147 (xvii): 187 (1924). Sellowiana var. glabrifolia Britton ex Rusby in Bull. Torr. Bot. Club, 28: 307 (1901). tristis Muell. Arg. in Mart. Fl. Bras. 11(2): 410 (1874). . volubilis L. Sp. Pl. 980 (1753). Ha SHH888 CALLITRICHACEAE Callitriche heteropoda Engelm. ex Hegelm. in Verh. Bot. Ver. Brandenb. 9: 40 (1867). BUXACEAE Styloceras columnare Muell. Arg. in DC. Prodr. 16(1): 10 (1869). ANACARDIACEAE Anacardium occidentale L. Sp. Pl. 383 (1753 Astronium fraxinifolium forma Pegaelie f Mattick in Notizbl. 11: 1005 A. fraxinifolium forma mollissimum Mattick in Notizbl. 11: 1003 (1934). A. fraxinifolium forma subglabrum Mattick in Notizbl. 11: 1004 (1934). A. urundueva (Allem.) Engl. in Engler, Bot. Jahrb. 1: 45 (1881). Cardenasiodendron brachypterum (Loesen.) Barkley in Lloydia, 17: 242 (1954). Lithraea molleoides (Vell.) Engl. in Mart. Fl. Bras. 12(2): 394, t. 83 (1876). L. Huasango Spruce ex Engl. in DC. Monog. Phan. 4: 461 (1883). Mauria Biringo Tul. in Ann. Sci. Nat. (ser. 3) 6: 365 (1846). M. boliviana Herzog in Fedde, Repert. Spec. Nov. 7: 60 (1909). M. ferruginea Tul. in Ann. Sci. Nat. (ser. 3) 6: 366 (1846). M. suaveolens Poepp. & Endl. Nov. Gen. & Spec. 3: 77 (1845). M. thaumatophylla Loesen. in Engler, Bot. Jahrb. 37: 573 (1906). Protium bolivianum Britton in Bull. Torr. Bot. Club, 16: 189 (1889) — Mauria erruginea. Schinopsis Balansae Engl. in Engler, Bot. Jahrb. 6: 286 goes S. cornuta Loesen. in Meded. Rijks Herbar. 27: 86 (1915 S. Lorentzii (Griseb.) Engl. in Engler, Bot. Jahrb. ve (1881). S. marginata Engl. in DC. Monog. Phan. 4: 464 (18 Schinus andinus (Engl.) I. M. Johnst. in Journ. singed ree 19: 259 (1938). S. andinus var. subtridentatus (O. Ktze.) Barkley in Brittonia, 5: 179 (1944). S. dependens var. andinus forma grandifolius Loesen. in Meded. Rijks Herbar. 27: 84 (1915) = S. andinus. S. dependens var. crenatus Engl. in Mart. Fl. Bras. 12(2): 388 (1876). S. dependens var. obovatus Engl. in Mart. Fl. Bras. 12(2): 387 (1876). S. dependens var. tomentosus R. E. Fries in Ark. Bot. 8(8): 11 (1908). S. diversifolius Rusby in Mem. Torr. Bot. Club, 4(3): 206 (1895) = S. s Pearcei. . Engleri Barkley in Brittonia, 5: 178 (1944). 119 S. fasciculatus var. boliviensis Barkley in Brittonia, 5: 177 (1944). S. ferox Hassl. in Fedde, Repert. Spec. Nov. 12: 373 (1913). S. maurioides Rusby in Bull. N. Y. Bot. Gard, 8: 102 (1912). S. Molle L. Sp. Pl. 388 (1753). S. myrtifolius (Griseb.) co in Inst. Mus. Univ. Nac. La Plata, Obra del Cincuentario, 2: 269 (19 S. Pearcei Engl. in Engler, ce Jahrb. 1: 423 (1881). S. polygamus (Cav.) Cabrera in Inst. Mus. Univ. Nac. La Plata, Obra del Cincuentario, 2: 269 (1937). S. polygamus forma parviflorus (March.) Cabrera in Rev. Mus. La Plata, secc. Bot. 2: 34 (1938). S. tomentosus Rusby in Bull. N. Y. Bot. Gard. 8: 102 (1912). S. Venturii Barkley in Brittonia, 5: 179 (1944). Tapirira guianensis Aubl. Pl. Guian. Fr. 1: 470, t. 188 (1775). T. Pearcei Rusby in Mem. Torr. Bot. Club, 6: 22 (1896). AQUIFOLIACEAE Ilex aggregata (R. & P.) Loesen. in Notizbl. 11: 95 (1931). I. amboroica Loesen. in Fedde, Repert. Spec. Nov. 7: 4" Aiea I. amplifolia Rusby in Mem. Torr. Bot. Club, 6: 20 (1896). is a Rusby in Mem. Torr. Bot. Club, nie 15 (1898) = I. ag- si ates dees in Nov. Act. Acad. Caes. Leop. 78: 161 (1901). I. boliviana Britton ex Rusby in Mem. Torr. Bot. Club, 3(3): 15 (1893). I. boliviana var. acutata Loesen. in Nov. Act. Acad. Caes. Leop. 78: 155 (1901). I. boliviana var. Brittoniana Loesen. in Nov. Act. Acad. Caes. Leop. 78: 155 (1901). I. boliviana var. Rusbyana Loesen. in Nov. Act. Acad. Caes. Leop. 78: 155 I. buxifolioides Loesen. in Nov. Act. Acad. Caes. Leop. 78: 225 (1901). I. buxifolioides var. fastigiata Loesen. in Nov. Act. Acad. Caes. Leop. 78: 226 (1901). I. Herzogii Loesen. in Meded. Rijks Herbar. 29: 3 (1916). I. imbricata Rusby, Descr. S. Am. Pls. 52 (1920). I. Mandonii Loesen. in Nov. Act. Acad. Caes. Leop. 78: 226 (1901). I. minimifolia Loesen. in Nov. Act. Acad. Caes. Leop. 78: 174 (1901). I. parviflora Benth. in Hook. Kew Journ. 4: 11 (1852). I. pseudoébenacea Loesen. in Nov. Act. Acad. Caes. Leop. 78: 386 (1901). I. sessiliflora Tr. & Pl. in Ann. Sci. Nat. (ser. 5) 16: 378 (1872). L. sessiliflora var. Pearcei Loesen. in Nov. Act. Acad. Caes. Leop. 78: 168 (1901). I. teratops Loesen. in Engl. & Prantl, Nat. Pflanzenfam. Nachtr. 3: 218 (1897). I. trichoclada Loesen. in Nov. Act. Acad. Caes. Leop. 78: 178 (1901). CELASTRACEAE Maytenus apurimacensis var. Trollii Loesen. in Notizbl. 18: 219 (1936). M. Cardenasii Rusby in Mem. N. Y. Bot. Gard. 7: 290 (1927). erythrocarpa Rusby in Mem. N. Y. Bot. Gard. 7: 290 (1927). flagellata Rusby in Mem. Torr. Bot. Club, 6: 20 (1896). ilicifolia var. boliviana Loesen. in Notizbl. 13: 218 (1936). meguillensis Rusby in Mem. N. Y. Bot. Gard. 7: 289 (1927). pseudoboaria Loesen. in Notizbl. 12: 29 (1934). pseudoboaria var. monantha Loesen. in Notizbl. 12: 29 (1934). M. subalata Reiss. in Mart. Fl. Bras. 11(1): 12, t. 2 (1861). M. tunarina Loesen. ex O. Ktze. Rev. Gen. 3(2): 37 (1898). Pre a 120 M. verticillata (R. & P.) DC. Prodr. 2: 10 (1825 ‘ M. Vitis-Idaea Griseb. in Goett. Abh. 19: 110 (1874). Moya boliviana (Loesen.) Loesen. in Engl. & Prantl, Nat. Pflanzenfam. (ed. 2) 20b: 147, 405 (1942). Plenckia integerrima (Lundell) Lundell in Phytologia, 1: 284 (1938). Schaefferia Dietheri Herter in Rev. Sudam. Bot. 6: 155 (1940). S. uruguayensis Speg. in Physis, 3: 346 (1917). HIPPOCRATEACEAE Anthodon decussatum R. & P. Fl. Peruv. 1: 45, t. 74 (1798). Cheiloclinium cognatum (Miers) A. C. Smith in, Brittonia, 3: 529 (1940). C. hippocrateoides (Peyr.) A. C. Smith in Brittonia, 3: 546 (1940). Hippocratea volubilis L. Sp. Pl. 1191 (1753) Prionostemma aspera (Lam.) Miers in Trans. Linn. Soc. 28: 355 (1872). Pristimera andina Miers in Trans. Linn. Soc. 28: 364 Pr ibe P. nervosa (Miers) A. C. Smith in Brittonia, 3: 370 (1940). alacia arborescens Rusby in Mem. N. Y. Bot, Gard. 7: 290 (1927) = Pristi- mera nervosa. S. impressifolia (Miers) A. C. Smith in Bull. Torr. Bot. Club, 66: 247 (1939). S. rotundifolia Rusby in Bull. N. Y. Bot. Gard. 4: 339 (1907) — Pristimera andina. Tontelea cuspidata A. C. Smith in Brittonia, 3: 490 (1940). T. fluminensis (Peyr.) A. C. Smith in Brittonia, 3: 477 (1940). T. glabra A. C. Smith in Brittonia, 3: 500 (1940). T. mauritioides (A. C. Smith) A. C. Smith in Brittonia, 3: 481 (1940). T. Ulei (Loesen.) A. C. Smith in Brittonia, 3: 498 (1940 STAPHYLEACEAE ? Turpinia sp. ICACINACEAE Citronella apogon (Griseb.) Howard in Contrib. Gray Herb. 142: 75 (1942). Dendrobangia boliviana Rusby in Mem. Torr. Bot. Club, 6: 19 (1896). SAPINDACEAE Allophylus angustatus (Tr. & Pl.) Radlk. in Engl. & Prantl, Nat. Pflanzenfam. 3(5): 312 (1895). cinnamomeus Radlk. in Bull. Torr. Bot. Club, 25: 336 (1898). edulis (St. Hil.) Niederlein in Bol. Mens. Mus. Prod. Argent. 3: 180 (1890). edulis var. gracilis Radlk. in Mart. Fl. Bras. 13(3): 385 (1900) leptostachys Radlk. in Mart. Fl. Bras. 13(3): 489 (1900). pauciflorus Radlk. in Meded. Rijks Herbar. 19: 61 (1913). petiolulatus var. pulverulentus Radlk. in Mart. Fl. Bras. 13(3): 491 (1900). punctatus (Poepp.) Radlk. in Mem. Torr. Bot. Club, 6: 21 (1896). strictus Radlk. in Engl. & Prantl, Nat. Pflanzenfam. 3(5): 312, fig. 162 (1895). Athyana weinmannifolia Syier} Radlk. in Durand, to Gen. 73 (1888). Cardiospermum corindum L. Sp. Pl. (ed. 2) 1: 526 (1762). C. corindum forma idacdethsaloncel (Griseb.) Radlk. in Engler, Pflanzenr. IV, 165(1): 406 (1932). C. corindum forma elongatum Radlk. in Engler, Pflanzenr. IV, 165(1): 406 1932). c: ince di forma loxense (HBK.) Radlk. in Engler, Pflanzenr. IV, 165(1): 402 (1932). oP oe ee ee 121 ae forma subsetulosum Radlk. in Engler, Pflanzenr. IV, 165(1): 404 1932) woe forma villosum (Mill.) Radlk. in Engler, Pflanzenr. IV, 165(1): 403 (1932). grandiflorum Sw. Prodr. 64 (1788). grandiflorum forma elegans (HBK.) Radlk. in Sitzber. Bayer. Akad. 8: 260 (1878). grandiflorum forma hirsutum (Willd.) Radlk. in Sitzber. Bayer. Akad. 8: 260 (1878). . Halicacabum L., Sp. Pl. 366 (1753). Cupania cinerea Poepp. & Endl. Nov. Gen. 3: 38 (1845). C. vernalis Camb. in St. Hil. Fl. Bras. Mer. 1: 387 (1827). Diatenopteryx sorbifolia Radlk. in Sitzber. Bayer. Akad. 8: 285 (1878). bipinnatum Radlk. in Sitzber. Math.-Phys. Akad. Muench. 8: 357 2 Ae Pe Q 78). Diplokeleba Herzogii Radlk. in Meded. Rijks Herbar. 40: 11 (1921). Dodonaea viscosa Jacq. Enum. PI. Carib. 19 (1760). D. viscosa var. vulgaris forma Burmanniana (DC.) Radlk. in Mart. Fl. Bras. 13(3): 646 (1900). Llagunoa Mandonti Rusby in Bull. N. Y. Bot. Gard. 4: 341 (1907) = L. nitida. L. nitida R. & P. Syst. 252 (1798). L. nitida var. mollis (HBK.) O. Ktze. Rev. Gen. 3(2): 43 (1898). Lophostigma plumosum Radlk. in Engl. & Prantl, Nat. Pflanzenfam. Nachtr. 228 (1897). Magonia glabrata St. Hil. Pl. Remarq. Brés. 241 (1824). M. pubescens St. Hil. Pl. Remarg. Brés. 239, t. 23, fig. A, t. 24, fig. A (1824). Matayba boliviana Radlk. in Bull. Torr. Bot. Club, 25: 336 (1898). M. scrobiculata (HBK.) Radlk. in Sitzber. Bayer. Akad. 9: 536 (1879). M. Steinbachii Melch. in Notizbl. 10: 349 (1928). Melicocca lepidopetala Radlk. in Sitzber. Bayer. Akad. 8: 344 (1878). Paullinia acutangula (R. & P.) Pers. Synops. Pl. 1: 443 (1805). bilobulata Radlk. in Engler, Bot. Jahrb. 37: 152 (1905). boliviana Radlk. in Mem. Torr. Bot. Club, 4(3): 206 (1895). boliviana forma glabrescens Radlk. in Mem. Torr. Bot. Club, 4(3): 206 5). cuneata Radlk. in Notizbl. 6: 150 (1914). dasystachya Radlk. in Abh. Math.-Phys. Bayer. Akad. Wiss. 19: 119, 270 (1896). dasystachya forma hirta Radlk. in Abh. Math.-Phys. Bayer. Akad. Wiss. 19: 271 (1896). elegans Camb. in St. Hil. Fl. Bras. Mer. 1: 370 (1827). ingaefolia Rusby in Mem. N. Y. Bot. Gard. 7: 291 (1927). neglecta Radlk. in Abh. Math.-Phys. Bayer. Akad. Wiss. 19: 167 (1896). pendulifolia Rusby in Mem. N. Y. Bot. Gard. 7: 291 (1927). pinnata L. Sp. Pl. 366 (1753). quercifolia Rusby in Mem. N. Y. Bot. Gard. 7: 292 (1927). rhizantha Poepp. & Endl. Nov. Gen. 3: 36, t. 243 (1845). ribesiaecarpa Rusby in Mem. N. Y. Bot. Gard. i 293 (1927). P. riparia HBK. Nov. Gen. & Spec. 5: 116 (1821). Tatei Rusby in Phytologia, 1: 64 (1934). Sapindus Saponaria L. Sp. Pl. 367 (1753). Serjania altissima (Poepp.) Radlk. Serj. Monog. 125 (1875). S. areolata Radlk. Monog. Serj. Suppl. 87 (1886 S. caracasana Willd. Sp. Pl. 2(1): 465 (1799). S. caracasana forma flavoviridis Radlk. in Fedde, Repert. Spec. Nov. 7: 356 (1909). sO ae, WU Sr We a8 1 122 ™M . caracasana forma puberula Radlk. in Trab. Mus. Farm. Fac. Cienc. Med. Buenos Aires, 21: 78 (1909). caracasana forma Radlkoferi O. Ktze. Rev. Gen. 3(2): 44 (1898). chaetocarpa Radlk. in Engl. & Prantl, Nat. Pflanzenfam. 3(5): 302 (1895). confertiflora Radlk. Consp. Serj. 4 (1874 ). confertiflora var. dasycephala Radlk. in Mem. Torr. Bot. Club, 6: 21 (1896). crassifolia Radlk. Serj. Monog. 225 (1875). dibotrya Poepp. & Endl. Nov. Gen. 3: 35, t. 242 (1844). didymadenia Radlk. in Bull. Herb. Boiss. (ser. 1) 1: 467 (1893). diffusa Radlk. Serj. Monog. 302 (1875) dumicola Radlk. Consp. Serj. 4 (1874). erecta Radlk. Consp. Serj. 8 (1874). glabrata HBK. Nov. Gen. & Spec. 5: 110 (1821). glabrata forma mollior Radlk. Serj. Monog. 169 (1875). glabrata forma mollissima Radlk. Serj. Monog. 168 (1875). . grandiceps Radlk. in Bull. Torr. Bot. Club, 25: 336 (1898). . humifusa Radlk. in Fedde, Repert. Spec. Nov. 7: 355 (1909). . leptocarpa Radlk. Serj. Monog. 112 (1875). lethalis St. Hil. Pl. Remarq. Brés. 235 (1825) leucocephala Radlk. in Meded. Rijks Herbar. 19: 59 (1913). lyrata Rusby in peel ceo 1: 64 (1934). Mansiana Mart. in Flora, 22, Beibl. 1: 9 (1839). marginata cana Decad. Stirp. Nov. Bras. 5: 44 (1848). marginata forma pluridentata Radlk. Serj. Monog. 160 (1875). meridionalis Camb. in St. Hil. Fl. Bras. Mer. 1: 366, t. 76 (1827). nutans Poepp. & Endl. ex Walp. Repert. 2: 813 (1843). ovalifolia Radlk. Serj. Monog. 218 (1875). pannifolia Radlk. in Engler, Bot. Jahrb. 37: 146 (1905). perulacea Radlk. Serj. Monog. 227 (1875). reticulata Camb. in St. Hil. Fl. Bras. Mer. 1: 359 (1827). reticulata forma platyptera Radlk. Serj. Monog. 159 (1875). rigida Radlk. Serj. Monog. 283 (1875). rubicaulis Benth. ex Radlk. Serj. Monog. 254 (1875). rubicunda Radlk. in Meded. Rijks Herbar. 19: 58 (1913). sphaerococca Radlk. Serj. Monog. 153 (1875). sufferruginea Radlk. Serj. Monog. 299 (1875). eripbtels Radlk. in Notizbl. 6: 149 (1914). Talisia esculenta (Camb.) Radlk. in Sitzber. Bayer. Akad. 8: 345 (1878). Thinouia coriacea Britton in Bull. Torr. Bot. Club, 16: 191 (1889). T. paraguariensis (Britton) Radlk. in Engl. & Prantl, Nat. Pflanzenfam. 3(5): 308 (1895). T. repanda Radlk. in Engl. & Prantl, Nat. Pflanzenfam. 3(5): 308 (1895). Urvillea filipes Radlk. in Fedde, Repert. Spec. Nov. 7: 354 9). U. laevis Radlk. in Att. Congr. Internat. Bot. Firenze, 1874: 63 (1876), nomen nudum. U. rufescens Camb. in St. Hil. Fl. Bras. Mer. 1: 354 (1827). U. ulmacea HBK. Noy. Gen. & Spec. 5: 106, t. 440 (1821). bee a a Ta bel las ae haga VITACEAE Cissus alata Jacq. Sel. Stirp. Amer. 23 (1763). C. erosa L. C. Rich. in Act. Soc. Hist. Nat. Paris, 1: 106 (1792). C. gongyloides (Baker) Planch. in DC. Monog. Phan. 5: 550 (1887). C. paraguayensis Planch. in DC. Monog. Phan. 5: 554 (1887). C. pruinosa Herzog in Meded. Rijks Herbar. 40: 30 (1921). ? C. salutaris (Baker) Herzog in Meded. Rijks Herbar. 40: 30 (1921). 123 C. sicyoides L. Sp. Pl. (ed. 2) 1: 170 (1762). C. sicyoides forma canescens (Lam.) Planch. in DC. Monog. Phan. 5: 531 (1887). C. sicyoides forma ovata (Lam.) Planch. in DC. Monog. Phan. 5: 526 (1887). C. trifoliolata L. Sp. Pl. (ed. 2) 1: 170 (1762). Vitis obliqua (R. & P.) O. Ktze. Rev. Gen. 3(2): 41 (1898). RHAMNACEAE Colletia foliosa var. microphylla 0. Ktze. Rev. Gen. 3(2): 38 (1898). C. spinosa Lam. Tabl. Encye. 2: 91, t. 129 (1793). Condalia Weberbaueri Perkins in Engler, Bot. Jahrb. 45: 463 (1911). Discaria Weddelliana (Miers) Escalante in Bol. Soc. Argent. Bot. 1: 223 (1946) Gouania Blanchetiana Miq. in Linnaea, 22: 797 ye colurnaefolia Reiss. in Mart. Fl. Bras. 11(1): 107 (1861). latifolia Reiss. in Mart. Fl. Bras. eave 103 (1861). polygama Urb. Symb. Ant. 4: 378 (191 sepiaria Mart. ex Britton in Bull. i ‘Bot. Club, 16: 189 (1889) = G. Blanchetiana. tomentosa Jacq. Sel. Stirp. Amer. 263 (1763). ursinicarpa Rusby in Bull. N. Y. Bot. Gard. 8: 103 (1912). Karwinskia oblongifolia Rusby in Mem. N. Y. Bot. Gard. 7: 293 (1927). Kentrothamnus foliosus (Rusby) Fscueebsit & Overk. in Fedde, Repert. Spec. Nov. 50: 327 (1941). K. penninervius Suesseng. & Overk. in Fedde, Repert. Spec. Nov. 50: 327 (1941). a Baeey Rhamnidium elaeocarpum Reiss. in Mart. Fl. Bras. 11(1): 94 (1861). R. glabrum Reiss. in Mart. Fl. Bras. 11(1): 95 (1861). Rhamnus boliviana Rusby in Mem. Torr. Bot. Club, cle 15 (1898). R. citrifolia Rusby in Bull. N. Y. Bot. Gard. 4: 340 (190 R. polymorpha (Reiss.) Weberb. in Eng]. & Prantl, Nat. stasis 3(5): 410 Sageretia elegans (HBK.) Brongn. in Ann. Sci. Nat. (ser. - 10: 360 (1827). Scutia Fiebrigii Perkins in Engler, Bot. Jahrb. 45: 464 (1911). Scypharia senticosa (HBK.) Miers in Ann. & Mag. Nat. Hist. (ser. 3) 6: 11 (1860). Zizyphus Mistol Griseb. in Goett. Abh. 19: 99 (1874). Z. piurensis Pilger in Engler, Bot. Jahrb. 54, Beibl. 117: 46 (1916). ELAEOCARPACEAE Crinodendron tucumanum Lillo in Act. 1 Reunién Nac. Soc. Argent. Cienc. Nat. Tucuman, 1916: 220 (1919). Muntingia Calabura L. Sp. Pl. 509 (1753). Sloanea fragrans Rusby in Mem. N. Y. Bot. Gard. 7: 294 (1927). S. obtusa (Splitg.) K. Schum. in Mart. Fl. Bras. 12(3): 181 (1886). S. arascee Radlk. in Sitzber. Math.-Phys. Akad. Muench. 12: 329 (1882). é rpa Rusby in Mem. N. Y. Bot, Gard. 7: 294 (1927) = S. pubescens. Vallea geckates Mutis ex L. f. Suppl. 266 (1781). V. stipularis var. pyrifolia (Turcz.) F. Ballard in Bot. Mag. 157: t. 9365 (1934). TILIACEAE Apeiba hispida Gaertn. Fruct. 2: 188, t. 121 (1791). A. membranacea Spruce ex Benth. in Journ. Linn. Soc. 5, Suppl. 2: 61 (1861). A. Tibourbou Aubl. Pl. Guian. Fr. 1: 538 (1775). Corchorus aquaticus Rusby in Mem. N. Y. Bot. Gard. 7: 295 (1927). 124 C. argutus var. longicarpus O. Ktze. Rev. Gen. 3(2): 26 (1898). C. hirtus L. Sp. Pl. (ed. 2) 1: 747 (1762). C. hirtus var. orinocensis (HBK.) K. Schum, in Mart. Fl. Bras. 12(3): 127 (1886). C. pilolobus Link, Enum. 2: 72 (1822). Curatella americana L. Syst. (ed. vm 1079 (1759). Heliocarpus americanus L. Sp. Pl. 448 (1753). ' H. boliviensis Hochr. in Ann. og od & Jard. Bot. eect 18-19: 118 (1914). H. popayanensis HBK. Nov. Gen. & Spec. 5: 341 (182 R. Rosei Hochr. in Ann. Conserv. & Jard. Bot. Genéve, = 19: 119 (1914). Luehea Fiebrigii Burret in Notizbl. 9: 829 (1926). L. Herzogiana R. E. Fries in Meded. Rijks Herbar. 19: 41 (1913). L. nobilis Tri. & Pl. ex Sprague in Kew Bull. 1926: 42 L. paniculata Mart. Nov. Gen. 1: 100, t. 62 (1826). L. speciosa Willd. in Neue Schrift. Ges. Naturf. Freunde, 3: 410 (1801). L. splendens Rusby in Mem. Torr. Bot. Club, 6: 12 (1896). L. Steinbachii Burret in Notizbl. 9: 827 (1926). L. tomentella Rusby in Bull. N. Y. Bot. Gard. 4: 382 (1907). Mollia boliviana Britton in Bull. Torr. Bot. Club, 16: 156 (1889). Triumfetta abutiloides St. Hil. Fl. Bras. Mer. 1: 283 (1827). . altheoides Lam. Encyc. 3: 420 (1789). grandiflora Vahl, Eclog. 2: 34 (1798). T. rhomboidea Jack: Sel. Stirp. Amer. 147 (1763). T. semitriloba L. Mant. 1: 73 (1767). T. semitriloba var. brasiliensis K. Schum. in Mart. Fl. Bras. 12(3): 135 (1886). T. semitriloba var. Martiana K. Schum. in Mart. Fl. Bras. 12(3): 135 (1886). a a MALVACEAE Abutilon amplissimum var. subpeltatum O. Ktze. Rev. Gen. 3(2): 17 (1898). A. arboreum (L.) Sweet, Hort. Brit. (ed. 1) 1: 53 (1827). A. Bakeri Rusby in Bull. N. Y. Bot. Gard. 4: 329 (1907). A. benense (Britton) Baker f. in Journ. Bot. 31: 338 (1893). A. Bridgesii Baker f. in Journ. Bot. 31: 338 (1893). A. cyclonervosum Hochr. in Ann. Conserv. & Jard. Bot. Genéve, 6: 27 (1902). A. cymosum Tri. & Pl. in Ann. Sci. Nat. (ser. 4) 17: 185 (1862) A. fuscicalyx Ulbr. in Fedde, Repert. Spec. Nov. 18: 498 (1915). A. Herzogianum R, E. Fries in Meded. Rijks Herbar. 19: 45 (1918). A. laxum Rusby in Mem. N. Y. Bot. Gard. 7: 296 (1927). A. mollissimum (Cav.) Sweet, Hort. Brit. (ed. 2) 65 (1830). A. paucifiorum St. Hil. Fl. Bras. Mer. 1: 206 (1827). A. ramiflorum St. Hil. Fl. Bras. Mer. 1: 199 (1827). A. stellatum (Cav.) O. Ktze. Rev. Gen. 3(2): 18 (1898 A. silvaticum (Cav.) K. Schum. in Mart. Fl. Bras. 12(3): 418 (1891). A. silvaticum subsp. Buchtienii R. E. Fries in Kgl. Sv. Vetensk. Handl. 24(2): 8 47). A. thyrsodendron Griseb. in Goett. Abh. 24: 48 (1879). A. tiubae K. Schum. in Mart. Fl. Bras. 12(3): 381 (1891). A. virgatum Sweet, Hort. Brit. (ed. 1) 1: 53 (1827). A. virgatum var. tomentosum K. Schum. in Mart. 12(3): 391 (1891). Anoda acerifolia var. minoriflora Hochr. in Ann. Conserv. & Jard. Bot. Genéve, 20: 51 (1916). A. cristata (L.) Schlechtd. in Linnaea, 11: 210 (1837). A. hastata (Willd.) Cav. Diss. 1: 39, t. 11, fig. 2 (1785). A. triangularis DC. Prodr. 1: 459 (1824). Bogenhardia crispa (L.) Kearney in Leafl. West. Bot. 7: 120 (1954). 125 Cienfuegosia argentina Giirke in Mart. Fl. Bras. 12(3): 579 fgg C. sulphurea (St. Hil.) Garcke in Bonplandia, 8: 150 (18 Gaya Gaudichaudiana St. Hil. Fl. Bras. Mer. 1: 191 (1827 oA G. Gaudichaudiana var. tarijensis (R. E. Fries) Hassl. in Fedde, Repert. Spec. Nov. 12: 366 (1913). G. gracilipes K. Schum. in Mart. Fl. Bras. 12(3): ba (1891). G. grandiflora Baker f. in Journ. Bot. 30: 136 (1892). G. hermannioides HBK. Nov. Gen. & Spec. 5: 268, t. 475 (1822). G. pilosa K. Schum. in Mart. Fl. Bras. 12(3): 355 (1891). G. rubricaulis Rusby, Descr. S. Am. Pls. 55 (1920). seca om maritimum var. polycarpum Todaro, Rel. Cult. Cott. 226, t. 8 (187 Hibiscus Janene Cav. Diss. 3: 146, t. 51, fig. 1 (1787). H. furcellatus Desr. in Lam. Encye. 3: 358 (1789 Lambertianus HBK. Nov. Gen. & Spec. 5: 291, * 478 (1822). rectiflorus Rusby in Mem. N. Y. Bot. Gard. 7: 300 (1927). rhomboideus Rusby in Mem. N. Y. Bot. Gard. 7: 301 (1927). sulphureus HBK. Nov. Gen. & Spec. 5: 289 (1822 Malachra radiata L. Syst. (ed. 12) 459 (1760). M. ruderalis Giirke in Mart. Fl. Bras. 12(3): 460 (1892). Malvastrum acaule (Cav.) A. Gray in Bot. U. S. Explor. sees 1: 150 (1854). amblyphyllum R. E. Fries in Ark. Bot. 6(2): 6, t. 2 (1906). bolivianum Baker f. in Journ. Bot. 29: 168 (18 91) = = Tarasa heterophylla. ome Pax in Fedde, Repert. Spec. Nov. 7: 243 (1909) = M. nubigen- mm mm mati (Cav.) Griseb. in Goett. Abh. 19: 90 (1874). dryadifolium Solms-Laub. in Bot. Zeit. 65(1): 131, fig. 6 (1907). Fiebrigit Ulbr. in Engler, Bot. Jahrb. 42: 113 (1908) = M. nubigenum. geranioides (Cham. & Schlechtd.) Hemsl. in Biol. Centr. Am. Bot. 1: 99 1879). er oe Ulbr. in Engler, Bot. Jahrb. 42: 114 (1908). icranthum Rusby, Descr. S. Am. Pls. 55 (1920) = seg tenella, esc maaan (Walp.) Baker f. in Journ. Bot. 29: 172 (189 nubigenum var. bipinnatifidum R. E. Fries in Kgl. Sv. wlio Handl. 24(2): 11 (1947). Oriastrum (Wedd.) Baker f. in Journ. Bot. 29: 172 (1891). parnassifolium (Hook.) A. Gray in Bot. U. S. Explor. Exped. 1: 150 (1854). peruvianum (L.) A. Gray in Bot. U. S. Explor. Exped. 1: 146 (1854). Rusbyi Britton in Bull. Torr. Bot. Club, 16: 64 (1889) — Tarasa Horn- schuchiana. tenellum Hieron. in Bol. Acad. Nac. Cienc. repre 4: 15 (1881). tricuspidatum (L.) A. Gray, Pl. Wright. 1: 16 (1852). waltheriifolium (Link) Ulbr. in Fedde, Repert. Spec. ae 13: 510 (1915). Modiola caroliniana (L.) G. Don, Gen. Syst. 1: 466 (1831 Nototriche acuminata A. W. Hill in Trans. Linn. Soc. 7: ee (1909). anthemidifolia (Wedd.) A. W. Hill in Engler, Bot. Jahrb. 37: 578 (1906). anthemidifolia var. sericea A. W. Hill in Kew Bull. 1932: 80. argyllioides A. W. Hill in Trans. Linn. Soc. 7: 245 (1909). bicolor Pax in Fedde, Repert. Spec. Nov. 7: 111 (1909). cinerea A. W. Hill in Trans. Linn. Soc. 7: 257 (1909). coactilis A. W. Hill in Kew Bull. 1932: 78. flabellata (Wedd.) A. W. Hill in Engler, Bot. Jahrb. all 579 (1906). glauca A. W. Hill in Engler, Bot. Jahrb. 37: 586 (1906). lanata A. W. Hill in Trans. Linn. Soc. 7: og (1909). leucosphaera A. W. Hill in Kew Bull. 1932: longirostris (Wedd.) A. W. Hill in Engler, ee Jahrb. 37: 579 (1906). dna S555 sade apa S55 eee cease 126 N. Mandoniana (Wedd.) A. W. Hill in Engler, Bot. Jahrb. 37: 579 (1906). N. nivea A. W. Hill in Kew Bull. 1932: 79. N. obcuneata (Baker f.) A. W. Hill in Engler, Bot. Jahrb. 37: 579 (1906). N. obcuneata var. cinerea A. W. Hill in Kew Bull. 1932: 80. N. Orbignyana (Wedd.) A. W. Hill in Trans. Linn. Soc. 7: 237 (1909). N. Pearcei (Baker f.) A. W. Hill in Engler, Bot. Jahrb. 37: 579 (1906). N. pedicularifolia (Remy) A. W. Hill in Engler, Bot. Jahrb. 37: 578 (1906). N. pseudoglabra A. W. Hill in Engler, Bot. Jahrb. 37: 586 (1906). N. pulverulenta Burtt & A. W. Hill in Kew Bull. 1948: 135. N. purpurascens A. W. Hill in Trans. Linn. Soc. 7: 249 (1909). N. pygmaea (Remy) A. W. Hill in Trans. Linn. Soc. 7: 218 (1909). N. sajamensis (Hieron.) A. W. Hill in Engler, Bot. Jahrb. 37: 579 (1906). N. Trollii Ulbr. in Notizbl. 11: 5382 (1932). N. violacea A. W. Hill in Kew Bull. 1932: 79. Pavonia ageratoides Rusby in Mem. N. Y. Bot. Gard. 7: 297 (1927). P. Bangii Ulbr. in Notizbl. 11: 543 (1932). P. canaminensis Rusby in Mem. N. Y. Bot. Gard. 7: 297 (1927). P. communis St. Hil. Fl. Bras. Mer. 1: 224 (1827). P. glechomoides A. Juss. in St. Hil. Fl. Bras. = 1: 227, t. 45 (1827). P. hastata Cav. Diss. 3: 138, t. 47, fig. 2 (178 P. humifusa A. Juss. in St. Hil. Fl. Bras. Mer. _ 235 (1827). P. leucantha Garcke in Jahrb. Bot. Gart. & Mus. Berlin, 1: 211 (1881). P. malacophylla (Nees & Mart.) Britton in Bull. Torr. Bot. Club, 16: 154 (1889). P. nigrobracteata (Rusby) Uittien “i Rec. Trav. Bot. Néerl. 33: 772 (1936). P. paniculata Cav. Diss. 3: 135 (178 P. Riedelii Giirke in Mart. Fl. Bras. core 493 (1892). P. sepium St. Hil. Fl. Bras. Mer. 1: 225 (1827). P. sidaefolia HBK. Nov. Gen. & Spec. 5: 283 (1822). P. sidaefolia var. diuretica (St. Hil.) Giirke in Mart. FI. Bras. 12(3): 509 (1892). P. speciosa subsp. polymorpha (St. Hil.) Giirke in Mart. Fl. Bras. 12(3): 495 (1892). P. spinifex (L.) Cav. Diss. 3: 183 (1787). P. subtriloba Rusby in Mem. N. Y. Bot. Gard. 7: 298 (1927). P. Typhalea (L.) Cav. Diss. 3: 184 (1787). P. umbrosa R. E. Fries in Kgl. Sv. Vetensk. Handl. 24(2): 25 (1947). Pseudabutilon callimorphum var. Friesii (Hassl.) R. E. Fries in Kgl. Sv. Vet- ensk. Handl. 43(4): 106 (1908) P. spicatum (HBK.) R. E. Fries in Kgl. Sv. Vetensk. Handl. 43(4): 98 (1908). Sida anomala St. Hil. Fl. Bras. Mer. 1: 177, t. 83 (1827) S. argentina K. Schum. in Mart. Fl. Bras. 12(3): 315 (1891). S. Bakeriana Rusby in Mem. Torr. Bot. Club, 6: 10 (1896). S. ciliaris L. Sp. Pl. (ed. 2) 2: 961 (1768). S. cordifolia L. Sp. Pl. (ed. 2) 2: 961 (1763). S. cordifolia var. serrata Baker f. in Journ. Bot. 30: 291 (1892). S. dictyocarpa var. esperanzae (R. E. Fries) Rodrigo in Rev. Mus. La Plata, 6: 151 (1944). S. diffusa HBK. Nov. Gen. & Spec. 5: 257 (1822). S. Glaziovii K. Schum. in Mart. Fl. Bras. 12(3): 322 (1891). S. glutinosa Cav. Diss. 1: 16, t. 2, fig. 8 (1785). S. gracilipes Rusby in Mem. Torr. Bot. Club, 6: 10 (1896). S. hastata St. Hil. Fl. Bras. Mer. 1: 190, t. 36, fig. 2 (1827). S. intermedia St. Hil. Fl. Bras. Mer. 1: 188, t. 36, fig. 1 (1827). S. linifolia Cav. Diss. 1: 14, t. 2, fig. 1 (1785). S. macrodon DC. Prodr. 1: 464 (1824). 127 oligandra K. Schum. in Engl. & Prantl, Nat. Pflanzenfam. 3(6): 43 (1890). paniculata L. Sp. Pl. (ed. 2) 2: 962 (1763). piauhyensis Ulbr. in Engler, Bot. Jahrb. 42: 226 (1908). potentilloides St. Hil. Fl. Bras. Mer. 1: 178 (1827). rhombifolia L. Sp. Pl. 684 (1753). rhombifolia var. canariensis (Willd.) Griseb. Fl. Brit. W. Ind. 74 (1859). rufescens St. Hil. Fl. Bras. Mer. 1: 185 (1827). supina L’Hérit. Stirp. 109 bis, t. 52 (1785). ns L. Sp. Pl. (ed. 2) 2: 963 (1763). . veronicifolia Lam. Encyc. 1: 5 (1783). Sphaeralcea miniata (Cav.) Spach, Hist. Vég. 3: 352 (1834). S. miniata var. inquilina Ulbr. in Fedde, Repert. Spec. Nov. 13: 504 (1915). Tarasa eo (Phil.) Krapovickas in Bol. Soc. Argent. Bot. 5: 130 (195 Ss vacates Krapovickas in Bol. Soc. Argent. Bot. 5: 131 (1954). T. heterophylla (Hook. & Arn.) Krapovickas in Bol. Soc. Argent. Bot. 5: 124 Dnnnnnnn n(n T. Hornschuchiana (Walp.) Krapovickas in Bol. Soc. Argent. Bot. 5: 189 (1954). T. Joergensenii (I. M. Johnst.) Krapovickas in Bol. Soc. Argent. Bot. 5: 138 T. Mandonii (Baker f.) Kearney in Leafl. West. Bot. 5: 190 (1949). T. O’Donellii Krapovickas in Bol. Soc. Argent. Bot. 5: 127 (1954). T. tarapacana (Phil.) Krapovickas in Bol. Soc. Argent. Bot. 5: 121 (1954). T. tenella (Cav.) Krapovickas in Bol. Soc. Argent. Bot. 5: 123 (1954). Urena lobata L. Sp. Pl. 692 (1753). Urocarpidium Shepardae (I. M. Johnst.) Krapovickas in Darwiniana, 10: 621 (1954). Wissadula andina Britton in Bull. Torr. Bot. Club, 16: 153 (1889). W. boliviana R. E. Fries in Kgl. Sv. Vetensk. Akad. Handl. 43(4): 40, t. 6 (1908). W. cruziana R. E. Fries in Kgl. Sv. Vetensk. Akad. Handl. 24(2): 10, t. 2 (1947). W. densiflora R. E. Fries in Kgl. Sv. Vetensk. Akad. Handl. 43(4): 64, t. 4, 6. W. excelsior (Cav.) Presl, Rel. Haenk. 2: 118 (1836). W. filipes Rusby in Mem. N. Y. Bot. Gard. 7: 296 (1927). W. grandifolia Baker f. in Bull. N. Y. Bot. Gard. 4: 328 (1907). W. Grisebachii R. E. Fries in Kgl. Sv. Vetensk. Akad. Handl. 43(4): 74 (1908). W. macrantha R. E. Fries in Kgl. Sv. Vetensk. Akad. Handl. 43(4): 67 (1908). W. paraguariensis Chod. in Bull. Herb. Boiss. (ser. 2) 1: 400 (1901). W. pedunculata R. E. Fries in Ark. Bot. 6(2): 12, t. 2, fig. 1-5 (1906). W. periplocifolia (L.) Griseb. Cat. Pl. Cubens. 25 (1866) W. periplocifolia var. hernandioides (L’Hérit.) Hochr. in Ann. Conserv. & Jard. Bot. Genéve, 6: 29 (1902). W. rostrata (Schum. & Thoun.) Hook. Niger Fl. 229 (1849). W. sordida Hochr. in Ann. Conserv. & Jard. Bot. or 6: 29 (1902). W. spicata (HBK.) Presl, Rel. Haenk. 2: 117 (1835). W. subpeltata (O. Ktze.) R. E. Fries in Kgl. Sv. Vetensk. Akad. Handl. 43(4): 56 (1908) BOMBACACEAE Bombax cumanense HBK. Nov. Gen. & Spec. 5: 300 (1822). B. gracilipes K. Schum. in Mart. Fl. Bras. 12(3): 211, t. 42 (1886). B. heteromorphum O. Ktze. Rev. Gen. 3(2): 18 (1898). B. marginatum (St. Hil.) K. Schum. in Mart. Fl. Bras. 12(3): 223, t. 44 (1886). B. Martianum K. Schum. in Mart. Fl. Bras. 12(3): 224 (1886). 128 B. rurrenabaqueanum Rusby in Mem. N. Y. Bot. Gard. 7: 301 (1927). B. Rusbyi Baker f. in Bull. N. Y. Bot. Gard. 4: 330 (1907). Cavanillesia hylogeiton Ulbr. in Notizbl. 6: 163 (1914). Ceiba boliviana Britton & Baker f. in Journ. Bot. 34: 174 (1896). C. Burchellii K. Schum. in Mart. Fl. Bras. 12(8): 211 (1886). C. Mandonii Britton & Baker f. in Journ. Bot. 34: 175 (1896). C. Samaiima (Mart. & Zucc.) K. Schum. in Mart. Fl. Bras. 12(3): 210 (1886). C. tunariense (O. Ktze.) K. Schum. in Just, Bot. Jahresber. 26(1): 343 (1900). Chorisia insignis HBK. Nov. Gen. & Spec. 5: 297, t. 485 (1821) C. speciosa St. Hil. Pl. Us. Bras. t. 63 (1828). C. ventricosa Nees & Mart. in Nov. Act. Acad. Caes. Leop. 11: 102, t. 9 (1823). Ochroma boliviana Rowlee in Journ. Wash. Acad. Sci. 9: 166 (1919) = O. Lagopus. O. Lagopus Sw. Prodr. 98 (1788). STERCULIACEAE Ayenia boliviana Rusby in Mem. Torr. Bot. Club, 3(3): 10 (1893). A. glabrescens K. Schum. in Mart. Fl. Bras. 12(3): 102 (1886). A. Schumanniana O. Ktze. Rev. Gen. 3(2): 24 (1898) A. tomentosa L. Syst (ed. 10) 1247 (1757). Byttneria asterotricha Mildbr. in Notizbl. 11: 143 (19381). benensis Britton in Bull. Torr. Bot. Club, 16: 155 (1889). boliviana Britton in Bull. Torr. Bot. Club, 16: 155 (1889). carthaginensis Jacq. in Sel. Stirp. Amer. 96 (1788). catalpaefolia Jacq. Hort. Schoenbr. 1: 21, t. 46 (1797). coriacea Britton in Bull. Torr. Bot. Club, 16: 156 (1889). filipes Mart. ex K. Schum. in Mart. Fl. Bras. 12(3): 95 (1886). hirsuta R. & P. Fl. Peruv. 3: 10 (1802). hirsuta var. parvifolia O. Ktze. Rev. Gen. — 24 (1898). lanceolata Sessé ex DC. Prodr. 1: 487 (1824). pescapraefolia Britton in Bull. Torr. Bot. Club, 16: 155 (1889). scabra L. Sp. Pl. (ed. 2) 1: 284 (1762). zuma coriacea Rusby in Bull. N. Y. Bot. Gard. 4: 332 (1907) = G. ulmifo- ay eA a lia G. tonentoen HBK. Nov. Gen. & Spec. 5: 320 (1823). G. ulmifolia Lam. Encyc. 3: 52 (1789). G. ulmifolia var. glabra K. Schum. in Mart. Fl. Bras. 12(3): 81 (1886). Helicteres amplifolia Rusby in Bull. N. Y. Bot. Gard. 4: 331 (1907). baruensis Jacq. Sel. Stirp. Amer. 236, t. 147 (1763) brevispica St. Hil. Fl. Bras. Mer. 1: 274 (1827). guanaiensis Rusby in Bull. N. Y. Bot. Gard. 4: 310 (1907). Lhotzkyana K. Schum. in Mart. Fl. Bras. 12(3): 19 (1886). Lhotzkyana var. pubinervis O. Ktze. Rev. Gen. 3(2): 25 (1898). Lhotzkyana var. tomentosa O. Ktze. Rev. Gen. 3(2): 25 (1898). pentandra L. Mant. 294 (1771). Rusbyi Britton in Bull. Torr. Bot. Club, 16: 154 (1889). Sacarolha A. Juss. in St. Hil. Pl. Us. Bras. t. 64 (1828). elochia anomala Griseb. in Goett. Abh. 19: 93 (1874). graminifolia St. Hil. Fl. Bras. Mer. 1: 160 (1825). hirsuta Cav. Diss. 6: 323 (1788). nervosa var. purpurea O. Ktze. Rev. Gen. 3(2): 25 (1898). nodiflora Sw. Prodr. 97 (1788). polystachya (HBK.) Tri. & Pl. in Ann. Sci. Nat. (ser. 4) g 341 (1862). pyramidata var. flava O. Ktze. Rev. Gen. 3(2): 25 (189 . pyramidata var. Grisebachii O. Ktze. Rev. Gen. 8(2): = (1898). eS SSS SS Bt tt at a 129 M. pyramidata var. Hieronymi K. Schum. in Mart. Fl. Bras. 12(3): 35 (1886). M. venosa Sw. Prodr. 97 (1788 M. venosa var. sericea (St. Hil.) K. Seraiic in Mart, Fl. Bras. 12(3): 38 (1886). M. yungasensis Rusby in Bull. N. Y. Bot. Gard. 4: 331 (1907). Sterculia excelsa Mart. in Flora, 24, II Beibl.: 40 (1841). S. laxiflora Rusby, Descr. S. Am. Pls. 56 (1920). S. pruriens (Aubl.) K. Schum. in Mart. Fl. Bras. 12(3): 8 (1886). S. striata St. Hil. & Naud. in Ann. Sci. Nat. (ser. 2) 18: 213 (1842), Theobroma Cacao L. Sp. Pl. 782 (1753). T. sylvestre Aubl. ex Mart. in Buchn. Repert. Pharm. 35: 24 (1830). Waltheria americana L. Sp. Pl. 673 (1753). W. communis St. Hil. Fl. Bras. Mer. 1: 155 (1825). W. Ladewii Rusby in Phytologia, 1: 65 (1934). DILLENIACEAE Davilla grandiflora St. Hil. & Tul. in Ann. Sci. Nat. (ser. 2) 17: 181 (1842). D. Lechleri Rusby in Mem. Torr. Bot. Club, 6: 2 (1896). D. microcalyx Herzog in Fedde, Repert. Spec. Nov. 7: 62 (1909). D. rugosa Poir. Encye. Suppl. 2: 457 (1811). D. rugosa var. capitata Rusby in Mem. Torr. Bot. Club, 6: 2 (1896). Doliocarpus dentatus (Aubl.) Standl. in Journ, Wash. Acad. Sci. 15: 286 (1925). D. ferrugineus Rusby in Mem. Torr. Bot. Club, 6: 3 (1896). D. magnificus Sleumer in Fedde, Repert. Spec. Nov. 39: 45 (1935). D. rufescens Sleumer in Fedde, Repert. Spec. Nov. 39: 46 (19385). D. semidentatus Garcke in Linnaea, 22: 48 (1849). Saurauia brevipes Rusby, Descr. S. Am. Pls. 57 (1920). S. coroicana Busc. in Malpighia, 26: t. 9, fig. 17 (1918). S. excelsa Willd. in Ges. Naturf. Freunde, Berlin, Neue Schr. 3: 407 (1801). S. parviflora Tri. & Pl. in Ann. Sci. Nat. (ser. 4) a 268 (1862). S. pseudoparviflora Busc. in Malpighia, 30: 158 (1927). S. pyramidata Sleumer in Notizbl. 12: 145 (1984). S. Rusbyi Britton in Bull. Torr. Bot. Club, 16: 64 (1889). S. scabra var. boliviana Busc. in Malpighia, 25: 11 (1912), nomen nudum. S. serrata DC. Prodr. 1: 526 (1824). S. Trolliana Sleumer in Notizbl. 12: 148 (1934). Tetracera aspera var. boliviana O, Ktze. Rev. Gen. 3(2): 2 (1898) = T. parvi- ora. T. parviflora (Rusby) Sleumer in Fedde, Repert. Spec. Nov. 39: 47 (1935). OCHNACEAE Cespedezia excelsa Rusby in Mem. Torr. Bot. Club, 6: 17 (1896). C. spathulata Planch. in Hook. Lond. Journ. Bot. 5: 647 (1846). Godoya oblonga R. & P. Fl. Peruv. Prodr. 58 (1794). Ouratea boliviana v. Tiegh. in Ann. Sci. Nat. (ser. 8) 16: 263 (1902). castaneaefolia (DC.) Engler in Mart. Fl. Bras. 12(2): 309 (1876). denudata v. Tiegh. in Ann. Sci. Nat. (ser. 8) 16: 263 (1902). flexuosa Rusby in Mem. N. Y. Bot. Gard. 7: 303 (1927). macrobotrys Rusby in Mem. N. Y. Bot. Gard. 7: 303 (1927). oblongifolia Rusby in Bull. N. Y. Bot. Gard. 8: 103 (1912). Trollii Sleumer in Notizbl. 18: 354 (1936). . Werdermannii Sleumer in Notizbl. 13: 356 (193 Seuvagesia deflexifolia Gardn. in Hook. Ic. Pl. a s 484 (1842). S. erecta L. Sp. Pl. 203 (1753). 9999909009 130 CARYOCARACEAE Caryocar glabrum (Aubl.) Pers. Synops. Pl. 2: 84 (1807). C. parviflorum A. C. Smith in Journ. Arnold Arb. 20: 298 (1939). MARCGRAVIACEAE Maregravia peduncularis Poepp. ex Rusby in Mem. Torr. Bot. Club, 3(3): 8 (1893) = Souroubea crassipes ? M. rectiflora Tri. & Pl. in Ann. Sci. Nat. (ser. 4) 17: 364 (1862). M. rectiflora var. macrophylla Wittm. in Mart. Fl. Bras. 12(1): 222, t. 40, fig. 2 (1878). Norantea anomala HBK. Nov. Gen. & Spec. 7: 218, t. 647 bis (1825). N. droseriformis Rusby ex Gilg & Werdermann in Engl. & Prantl, Nat. Pflan- zenfam. (ed. 2) 21: 101 (1925). N. macrostoma Gilg in Engler, Bot. Jahrb. x Beibl. 60: 30 (1898). N. oxystilis Baill. in Adansonia, 10: 243 (1872). N. Weddelliana Baill. in Adansonia, 10: 242 (1872). Souroubea brachystachya Rusby in Bull. N. Y. Bot. Gard. 8: 103 (1912). S. crassipes Wittm. in Mart. Fl. Bras. 12(1): 254 (1878). S. guianensis Aubl. Pl. Guian. Fr. 1: 244, t. 97 (1775). QUIINACEAE Quiina Blackii Pires in Bol. Técn. Inst. Agron. Norte, Para, no. 20: 44, t. 8 (1950). THEACEAE Freziera angulosa Tul. in Ann. Sci. Nat. (ser. 3) 8: 332 (1847) F. boliviensis Wawra in Mart. Fl. Bras. 12(1): 284 (1886) — F. lanata. F. caloneura Kobuski in Journ. Arnold Arb. 22: 477 (1941). F.. maequalifolia (Lingelsh.) Kobuski in Ann. Missouri Bot. Gard. 25: 354 (1938) = F. inaequilatera. F. inaequilatera Britton in Bull. Torr. Bot. Club, 16: 63 (1889). F. lanata (R. & P.) Tul. in Ann. Sci. Nat. (ser. 3) 8: 334 (1847). F. subintegrifolia (Rusby) Kobuski in Ann. Missouri Bot. Gard. 25: 355 (1938). F. yungasia Tul. in Ann. Sci. Nat. (ser. 3) 8: 333 (1847) = F. lanata. Laplacea fruticosa (Schreb.) Kobuski in Journ. Arnold Arb. 28: 487 (1947). L. pubescens Planch. & Lind. in Ann. Sci. Nat. (ser. 4) 18: 269 (1862). L. pubescens var. subcaudata Kobuski in Journ. Arnold Arb. 31: 427 (1950). L. symplocoides Planch. & Lind. in Ann. Sci. Nat. (ser. 4) 18: 269 (1862). Taonabo flavifolia Rusby in Bull. N. Y. Bot. Gard. 8: 104 (1912) = Tern- stroemia asymmetrica Ternstroemia aaymnetrion Rusby in Bull. N. Y. Bot. Gard. 4: 327 (1907). T. brasiliensis Camb. in St. Hil. Fl. Bras. Mer. 1: 298 (1827). T. circumscissilis Kobuski in Journ. Arnold Arb. 23: 304 (1942). T. polyandra Kobuski in Journ. Arnold Arb. 23: 333 (1942). T. subserrata (Rusby) Melchior in Engl. & Prantl, Nat. Pflanzenfam. (ed. 2) 21: 142 (1925). GUTTIFERAE Calophyllum ellipticum Rusby in Mem. N. Y. Bot. Gard. 7: 303 (1927). Caopia cordata Rusby in Bull. N. Y. Bot. Gard. 8: 105 (1912) = Vismia sp. ? C. crassa Rusby in Mem. Torr. Bot. Club, 4(3): 204 (1895) = Vismia sp. C. parvifolia Rusby in Phytologia, 1: 65 (1934) = Vismia sp. ? 131 Chrysochlamys macrophylla Pax in Fedde, Repert. Spec. Nov. 7:111 (1909). C. myrcioides Tri. & Pl. in Ann. Sci. Nat. (ser. 4) 14: 260 (1860). Clusia amazonica Tri. & Pl. in Ann. Sci. Nat. (ser. 4) 13: 358 (1860). criuva Camb. in St. Hil. Fl. Bras. Mer. 1: 317 (1827). elongata Rusby in Bull. N. Y. Bot. Gard. 8: 105 (1912). insignis Mart. Nov. Gen. 3: 164 (1829). latipes Tri. & Pl. in Ann. Sci. Nat. (ser. 4) 13: 365 (1860). Lechleri Rusby in Bull. N. Y. Bot. Gard. 8: 105 (1912). multiflora HBK. Nov. Gen. & Spec. 5: 200 (1822). pseudomangle Tri. & Pl. in Ann. Sci. Nat. (ser. 4) 13: 370 (1860). ramosa Rusby in Mem. Torr. Bot. Club, 4(3): 204 (1895). ternstroemioides Rusby, Descr. S. Am. Pls. 59 (1920). trochiformis Vesque, Epharmosis, 3: 4, t. 3 (1892). Havetia laurifolia HBK. Nov. Gen. & Spec. 5: 204, t. 462 (1822). Havetiopsis flavida (Benth.) Pl. & Tri. in Ann. Sci. Nat. (ser. 4) 14: 247 1860). aaaanaaaaa H. glauca Rusby in Bull. N. Y. Bot. Gard. 4: 309 (1907). gsi oes andinum Gleason in Torreya, 29: 137 (1929). H. bolivianum Keller in Bull. Herb. Boiss. (ser. 2) 8: 189 (1908). brasiliense Chois. ex DC. Prodr. 1: 547 (1824). brevistylum Chois. Prodr. Hyper. 51 (1821). connatum Lam. Encyc. 4: 168 (1797). connatum var. Fiebrigii Briq. in Ann. Conserv. & Jard. Bot. Genéve, 20: 390 (1919). connatum var. paraguariense Briq. in Ann. Conserv. & Jard. Bot. Genéve, 20: 390 (1919). laricifolium Juss. in Ann. Mus. Paris, 3: 160, t. 16 (1804). struthiolaefolium Juss. in Ann. Mus. Paris, 3: 160 (1804). struthiolaefolium var. parviflorum Keller in Bull. Herb. Boiss. (ser. zy 8s 182 (1908). stylosum Rusby in Bull. N. Y. Bot. Gard. 4: 326 (1907). thesiifolium HBK. Nov. Gen. & Spec. 5: 192 (1822). Kielmeyera paniculata Rusby in Mem. Torr. Bot. Club, 6: 9 (1896). Marila laxiflora Rusby in Mem. Torr. Bot. Club, 6: 9 (1896). Rengifa acuminata Tri. & Pl. in Ann. Sci. Nat. (ser. 4) 14: 243 (1860). Rheedia Achachairu Rusby in Mem. N. Y. Bot. Gard. 7: 304 (1927). R. floribunda (Miq.) Tri. & Pl. in Ann. Sci. Nat. (ser. 4) 14: 319 (1860). R. rogaguensis Rusby in Mem. N| Y. Bot. Gard. 7: 304 (1927). R. Spruceana Engl. in Mart. Fl. Bras. 12(1): 463 (1888). Symphonia globulifera L. f. Suppl. 302 (1781). Tovomita alatopetiolata O. Ktze. Rev. Gen. 3(2): 16 (1898). T. micrantha A. C. Smith in Phytologia, 1: 123 (1935). T. umbellata Benth. in Hook. Lond. Journ. Bot. 2: 367 (1843). T. Weddelliana Tri. & Pl. in Ann. Sci. Nat. (ser. 4) 14: 277 (1860). Vismia cayennensis (L.) Pers. Synops. Pl. 2: 86 (1807). . dealbata HBK. Nov. Gen. & Spec. 5: 184, t. 454 (1822). glabra R. & P. Syst. 183 (1798). guianensis (Aubl.) Pers. Synops. Pl. 2: 86 (1807). magnoliifolia Cham. & Schlechtd. in Linnaea, 3: 118 (1828). plicatifolia Hochr. in Ann. Conserv. & Jard. Bot. Genéve, 21: 54 (1919). tomentosa R. & P. Syst. 183 (1798). <<<<<< ELATINACEAE Elatine triandra Schkuhr, Bot. Handb. 1: 345, t. 109b (1791). 132 FRANKENIACEAE Anthobryum tetragonum Phil. in Anal. Mus. Nac. Chile, Bot. 1891: 51, t. 2 1891). A. triandrum (Remy) Surgis in Rev. Gén. Bot. 34: 455 (1922). Frankenia farinosa Remy in Ann. Sci. Nat. (ser. 3) 8: 236 (1847). BIXACEAE Bixa Orellana L. Sp. Pl. 512 (1753). COCHLOSPERMACEAE Amoureuxia unipora v. Tiegh. in Journ. de Bot. 14: 48 (1900). Cochlosperum hibiscioides Kunth, Syn. Pl. Aequin. 3: 214 (1824). C. insigne St. Hil. Pl. Us. Bras. t. 57 (1824-28). C. tetraporum Hall. f. in Meded. Rijks Herbar. 19: 39 (1913). C. trilobum Standl. in Field Mus. Publ. Bot. 22: 92 (1940). KOEBERLINIACEAE Koeberlinia spinosa Zucc. in Abh. Akad. Muench. 1: 359 (1832). VIOLACEAE Anchietea parviflora Hall. f. in Meded. Rijks ae 19: 64 (1913 la appendiculata Rusby in Mem. N. Y. Bot. Gard. 7: Hg (1927 e— Hybanthus sp. C. balaénsis Rusby in Mem. N. Y. Bot. Gard. 7: 306 (1927) = Hybanthus sp. C. sessiliflora O. Ktze. Rev. Gen. 3(2): 8 (1898) — Hybanthus sp. Hybanthus atropurpureus (St. Hil.) Taub. in Engl. & Prantl, Nat. Pflanzen- fam. 3(6): 333 (1895). H. biacuminatus (Rusby) G. K. Schulze in Notizbl. 12: 114 (1934). H. Calceolaria (L.) G. K. Schulze in Notizbl. 12: 114 (1934). H. communis (St. Hil.) Taub. in Engl. & Prantl, Nat. Pflanzenfam. 3(6): 333 (1895 iF oppositifolius (L.) Taub. in Engl. & Prantl, Nat. Pflanzenfam. 3(6): 333 1895 parviflorus (Mut.) Baill. Bot. Médic. 841 (1884). parviflorus var. Bangii (Rusby) Sparre in Lilloa, 23: 535 (1950). rviflorus var. glutinosus (Vent.) Hassl. in Bull. Soc. Bot. Genéve (ser. 2), 1: 214 (1909) Tonidium subglaucum Herzog in Meded. Rijks Herbar. 40: 15 (1921) = Hy- banthus sp. Leonia glycycarpa R. & P. Fl. Peruv. 2: 69, t. 222 (1799). Rinorea albicaulis (Turcz.) Blake in Contrib. U. S. Nat. Herb. 20: 517 (1924). R. gracilis Rusby in Bull. N. Y. Bot. Gard. 8: 106 (1912). R. juruana Ule in Verh. Bot. Ver. Brandenb. 47: 158 (1905). R. Lindeniana (Tul.) O. Ktze. Rev. Gen. 1: 42 (1891). R. ovalifolia (Britton) Blake in Contrib. U. S. Nat. Aas 20: 513 (1924). R. viridifolia Rusby in Mem. Torr. Bot. Club, 6: 5 (1896 Viola Bangiana W. Becker in Beih. Bot. Centralbl. aa: 89 (1907). V. Bangii Rusby in Mem. Torr. Bot. Club, 6: 5 (1896). V. boliviana W. Becker in Beih. Bot. Centralbl. 22(2): 88 (1907) = V. Stein- bachii. V. boliviana Britton in Bull. Torr. Bot. Club, 16: 18 (1889). pt it bet 133 Bridgesii Britton in Bull. Torr. Bot. Club, 16: 18 (1889). ii Becker in Beih. Bot. Centralbl. 22(2): 94 (1907). exigua W. Becker in Engler, Bot. Jahrb. 37: 590 (1906). flavicans Wedd. in Ann. Sci. Nat. a 5) 1: 292 (1864). Hillii W. Becker in Kew Bull. 1928: Humboldtii var. renifolia Britton in et Torr. Bot. Club, jo 18 (1889). Mandonii W. Becker in Beih. Bot. Centralbl. 22(2): 95 (19 micranthella Wedd. in Ann. Sci. Nat. (ser. 5) 1: 291 rth Orbignyana Remy in Ann. Sci. Nat. (ser. 3) 6: 353 (1846). producta W. Becker in Engler, Bot. Jahrb. 37: 591 (1906). pusillima Wedd. in Ann. Sci. Nat. (ser. 5) 1: 291 (1864). pygmaea Juss. ex Poir. in Lam. Encyc. 8: 630 (1808). scandens Humb. & Bonpl. ex Roem. & Schult. cab 5: 891 (1819). Steinbachii W. Becker in Notizbl. 9: 1040 (1926 thymifolia Britton in Bull. Torr. Bot. Club, 16: 18 (18 veronicaefolia Pl. & Lind. in Ann. Sci. Nat. (ser. 4) = ee (1862). SS SMe Pe MS Sta ssa FLACOURTIACEAE =~ boliviana (Mand. & Wedd.) Britton in Bull. Torr. Bot. Club, 17: 214 1890) = A. stellata A. stellata Lillo in Act. 1 Reunién Nac. Soc. Arg. Cienc. Nat. Tucuman, 1916: 223 (1919). Azara salicifolia Griseb. in Goett. Abh. 24: 20 (1879). Banara amazonica Sleumer in Notizbl. 12: 50 (1934). B. laxiflora Benth. in Journ. Linn. Soc. 5, Suppl. 2: 91 (1861). B. pyramidata Rusby in Mem. Torr. Bot. Club, 3(3): 33 (1893). Casearia aculeata Jacq. Enum. Pl, Carib. 21 (1760). C. acuminata DC. Prodr. 2: 50 (1825). C. albicaulis Rusby in Mem. N. Y. Bot. Gard. 7: 307 (1927). C. arborea (L. C. Rich.) Urb. Symb. Ant. 4: 421 (1910). C. attenuata Rusby in Mem. Torr. Bot. Club, 6: 41 (1896). . Bangii Rusby in Mem. Torr. Bot. Club, 3(3) : 34 (1893) —= C. Cambessedesii. berberoidia Rusby in Mem, N. Y. Bot. Gard. 7: 307 (1927) = C. aculeata. boliviana Briq. in Ann. Conserv. & Jard. Bot. Genéve, 2: 69 (1898). brasiliensis Eichl. in Mart. Fl. Bras. 13(1): 477 (1872). Cambessedesii Eichl. in Mart. Fl. Bras. 13(1): 475 (1872). combaymensis Tul. in Ann, Sci. Nat. (ser. 3) 7: 362 (1847). commutata Briq. in Ann. Conserv. & Jard. Bot. Genéve, 2: 65 (1898) membranacea Britton in Bull. Torr. Bot. Club, 17: 214 (1890) = C. combay- oblongifolia Camb, in St. Hil. Fl. Bras. Mer. 2: 234 (1829). obtusifolia Rusby in Bull. N. Y. Bot. Gard. 4: 362 (1907) = C. boliviana. punctata Spreng. Neue Entdeck. 2: 154 (1821). sylvestris Sw. Fl. Ind. Occ. 2: 752 (1800). sylvestris var. tomentella Rusby in Mem. Torr. Bot. Club, 6: 41 (1896). Prockia completa Hook. Ic. Pl. 1: t. 94 (1828). P. crucis L. Sp. Pl. (ed. 2) 1: 745 (1762). P. grandiflora Herzog in Fedde, Repert. Spec. Nov. 7: 62 (1909). P. septemnervia Spreng. Syst. 2: 609 (1825). Xylosma ellipticum (Clos) Hemsl. Biol. Centr. Am. Bot. 1: 57 (1879). X. ovatum Rusby in Bull. N. Y. Bot. Gard. 4: 323 (1907) = X. Rusbyanum. X. Rusbyanum Sleumer in Notizbl. 12: 56 (1934), X. venosum N. E. Br. in Trans. & Proc. Bot. Soc. Edinb. 20: 46 (1894). PAN ARARAHAARAAA one) 134 LACISTEMACEAE Lacistema aggregatum (Berg) Rusby in Bull. N. Y. Bot. Gard. 4: 447 (1907). L. bolivianum Gandog. in Bull. Soc. Bot. France, 66: 288 (1920) = L. aggregat- um. TURNERACEAE Piriqueta Duarteana var. grandifolia Urb. in Jahrb. Bot. Gart. Berlin, 2: 67 P. seticarpa Rusby in Mem. N. Y. Bot. Gard. 7: 308 (1927). Turnera melochioides var. oblongifolia Urb. in Jahrb. Bot. Gart. Berlin, 2: 116 4H 88 8 4 8 5H muricata Rusby in Mem. N. Y. Bot. Gard. 7: 308 (1927). sidoides var. lycopifolia (DC.) Urb. in Jahrb. Bot. Gart. Berlin, 2: 102 (1883). ulmifolia var. caerulea (DC.) Urb. in Jahrb. Bot. Gart. Berlin, 2: 144 (1883) ulmifolia var. elegans (Otto) Urb. in Jahrb. Bot. Gart. Berlin, 2: 139 (1883) ulmifolia var. grandidentata Urb. in Jahrb. Bot. Gart. Berlin, 2: 139 Weddelliana Urb. & Rolfe in Jahrb. Bot. Gart. Berlin, 2: 90 (1883). - Weddelliana var. brachyphylla Urb. in Engler, Bot. Jahrb. 25, Beibl. 60: 3 (1898). . Whitei Rusby in Mem. N. Y. Bot. Gard. 7: 309 (1927). PASSIFLORACEAE Passiflora auriculata HBK. Nov. Gen. & Spec. 2: 131 (1817). P. Bangii Masters in Rusby in Bull. N. Y. Bot. Gard. 4: 363 eines ace Sel Oy fy ty . OS ty ty violacea. Buchtienii Killip in Journ. Wash. Acad. Sci. 14: 115 (1924). callimorpha Harms in Fedde, Repert. Spec. Nov. 18: 295 (1922). So mi Tri. & Pl. in Ann. Sci. Nat. (ser. 5) 17: 161 (1873), nomen pro- visori capanentieles Rusby in Bull. N. Y. Bot. Gard. 8: 107 (1912) = P. auricu- ata, cincinnata Mast. in Gard. Chron. 1868: 966. coccinea Aubl. Pl. Guian. Fr. 2: 828, t. 8324 (1775). coriacea Juss. in Ann. Mus. Hist. Nat. Paris, 6: 109, t. 39, fig. 2 (1805). dalechampioides Killip in Journ. Wash. Acad. Sci. 17: 429 (1927). erosa Rusby in Bull. N. Y. Bot. Gard. 4: 863 (1907) = P. morifolia. . exoperculata Mast. in Mart. Fl. Bras. 18(1): 556 (1872). foetida L. Sp. Pl. 959 (17538). foetida var. gossypifolia (Desv.) Mast. in Trans. Linn. Soc. 27: 631 (1871). gracilens (A. Gray) Harms in Engl. & Prantl, Nat. Pflanzenfam. 3(6a): 91 893). Guentheri Harms in Notizbl. 10: 811 (1929). hastifolia Killip in Journ. Wash. Acad. Sci. 14: 115 (1924). ianthina Mast. in Journ. Bot. 21: 36 (1883) = P. umbilicata. ichthyura Mast. in Mart. Fl. Bras. 13(1): 587 (1872). ligularis Juss. in Ann. Mus. Hist. Nat. 6: 113, t. 40 (1805). Mandonii (Mast.) Killip in Journ. Wash. Acad. Sci. 14: 213 (1924). mapiriensis Harms in Notizbl. 10: 810 (1929). misera HBK. Nov. Gen. & Spec. 2: 136 (1817). SPU UU MUU RMN UNUM MUN 135 mixta L. f. Suppl. 408 (1781). mollissima (HBK.) Bailey in Rhodora, 18: 156 (1916). Mooreana Hook. f. in Bot. Mag. 66: t. 3773 (1840). morifolia Mast. in Mart. Fl. Bras. 13(1): 555 (1872). naviculata Griseb. in Goett. Abh. 19: 149 (1874). nephrodes Mast. in Bull. Torr. Bot. Club, 17: 282 (1890). nigradenia Rusby in Mem. N. Y. Bot. Gard. 7: 311 (1927). palmatisecta Mast. in Mart. Fl. Bras. 13(1): 564 (1872). pinnatistipula Cav. Icon. 5: 16, t. 428 (1799). Pohlii Mast. in Mart. Fl. Bras. 13(1): 586 (1872). punctata L. Sp. Pl. 957 (1753). quadrangularis L. Syst. (ed. 10) 1248 (1759). quadriglandulosa Rodschied, Med. Chir. Bemerk. Esseq. 77 (1796). Rojasii Hassl. ex Harms in Notizbl. 10: 812 (1929). rubra L. Sp. Pl. 956 (1753). rubrotincta Killip in Journ. Wash. Acad. Sci. 17: 429 (1929). Rusbyi Masters in Bull. Torr. Bot. a 17: 282 (1890). serrato-digitata L. Sp. Pl. 960 (175 Steinbachiit Harms in Notizbl. 10: Hee (1920) = P. Mandonii. suberosa L. Sp. Pl. 958 (1753). Tatei Killip & Rusby in Phytologia, 1: 66 (1934). tenuifila Killip in Journ. Wash. Acad. Sci. 17: 430 (1927). translinearis Rusby in Mem. N. Y. Bot. Gard. Uy ek (1927) = P. misera as to foliage and P. quadriglandulosa as to flow tricuspis Masters in Mart. Fl. Bras. 13(1): ey sey triloba R. & P. ex DC. Prodr. 3: 330 (1828). trisecta Masters in Mart. Fl. Bras. 13(1): 564 (1872). umbilicata (Griseb.) Harms in Engl. & Prantl, Nat. Pflanzenfam. 3 (pt. 6a): 91 (1893). urnaefolia Rusby in Mem. Torr. Bot. Club, 6: 42 (1896). venosa Rusby in Mem. Torr. Bot. Club, 6: 42 (1896). vespertilio L. Sp. Pl. 597 (1753). violacea Vell. Fl. Flum. 9: t. 84 (1827). Warmingii, subsp. chacoénsis R. E. Fries in Ark. Bot. 8 (no. 8): 4, t. 1, fig. 7-8 (1908) = P. morifolia. acumensis Rusby in Mem. N. Y. Bot. Gard. 7: 310 (1927) = P. quadrig- landulosa. Tacsonia boliviana Rusby in Mem. Torr. Bot. Club, 3(3): 37 (1893) = Pas- siflora gracilens. CARICACEAE Carica boliviana Rusby in Bull. N. Y. Bot. Gard. 4: 364 (1907). aaaagaaaaaa Fiebrigii Harms in Notizbl. 8: 99 (1921). gossypiifolia Griesb. in Goett. Abh. 24: 137 (1879). lanceolata (A. DC.) Solms-Laub. in Mart. Fl. Bras. 13(3): 179 (1889). microcarpa Jacq. Hort. Schoenbr. 3: 32, tt. 309, 310 (1798). Papaya L. Sp. Pl. 1036 (1753). pinnatifida O. Heilb. in Sv. Bot. Tidskr. 30: 224 (1936 quercifolia (St. Hil.) Solms-Laub. in Mart. Fl. Bras. . 178 (1889). stenocarpa O. Heilb. in Sv. Bot. Tidskr. 30: 221 (19386). triplisecta Herzog in Meded. Rijks Herbar. 27: 15 (1915). tunariensis (O. Ktze.) K. Schum. in Just, Bot. Jahresber. 26(1): 345 (1900). 912). Jacaratia boliviana Rusby in Bull. N. Y. Bot. Gard. 8: 107 J. Hassleriana Chod in Bull. Herb. Boiss. (ser. 2) 3: 546 (1903). 136 LOASACEAE Blumenbachia chuquitensis (Meyen) Hook. f. in Bot. Mag. 51: t. 6143 (1875). Cajophora andina Urb. & Gilg in Nov. Act. Acad. Caes. Leop. 76: 287 (1900). boliviana Urb. & Gilg in Nov. Act. Acad. Caes. Leop. 76: 282 (1900). Buraeavi Urb. & Gilg in Mem. Torr. Bot. Club, 3(3): 37 (1893). canarinoides (Len. & C. Koch) Urb. & Gilg in Engl. & Prantl, Nat. Pflan- zenfam. 3(6a): 119 (1894). cernua (Griseb.) Urb. & Gilg in Rev. Mus, La Plata, 5: 291 (1893). chuquisacana Urb. & Gilg in Nov. Act. Acad. Caes. Leop. 76: 322 (1900). cinerea Urb. & Gilg in Nov. Act. Acad. Caes. Leop. 76: 321 (1900). cirsiifolia Pres], Rel. Haenk. 2: 42, t. 56 (1831). contorta Presl, Rel. Haenk. 2: 42 (1831) = C. Preslii. coronata (Arn.) Hook. & Arn. in Hook. Bot. Misc. 3: 327 (1833). Fiebrigii Urb. & Gilg in Engler, Bot. Jahrb. 45: 470 (1911). heptamera (Wedd.) Urb. & Gilg in Engl. & Prantl, Nat. Pflanzenfam. 3(6a): 119 (1894). horrida (Britton) Urb. & Gilg in Mem. Torr. Bot. Club, 3(3): (1893). Kuntzei Urb. & Gilg in Nov. Act. Acad. Caes. oor 76: 314 (1900). macrophylla Rusby in Phytologia, 1: 67 (193 Mandoniana Urb. & Gilg in Nov. Act. Acad. ae Leop. 76: 299 (1900). Orbignyana Urb. & Gilg in Engl. & Prantl, Nat. Pflanzenfam. 3(6a): 119 (1894). pedicularifolia Killip in Journ. Wash. Acad. Sci. 18: 95 (1928). Preslii Urb. & Gilg in Nov. Act. Acad. Caes. Leop. 76: 306 (1900). rosulata (Wedd.) Urb. & Gilg in Engl. & Prantl, Nat. Pflanzenfam. 3(6a): 119 (1894). Rusbyana Urb. & Gilg in Mem. Torr. Bot. Club, 3(3): 35 (1893). sepiaria (G. Don) Macbr. in Candollea, 8: 23 (1940). sphaerocarpa Urb. & Gilg in Nov. Act. Acad. Caes. Leop. 76: 296 (1900). . superba R. A. Phil. in Anal. Mus. Nac. Chile, ‘ate 8: 23 (1891). Gronovia scandens L. Sp. Pl. 202 (1758). Klaprothia mentzelioides HBK. Nov. Gen. & Spec. 6: 123, t. 587 (1823). Loasa ferruginea Urb. & Gilg in Nov. Act. Acad. Caes. Leop. 76: 225 (1900). L. heptamera var. chelidonifolia Wedd. Chlor. And. 2: 218 (1861) == Cajo- phora superba. L. Herzogii Urb. & Gilg in Meded. Rijks Herbar. 29: 54 (1916). Mentzelia aspera L. Sp. Pl. 516 (1753). M. cordifolia Dombey ex Urb. & Gilg in Nov. Act. Acad. Caes. Leop. 76: 74 (1900). M. Fendleriana Urb. & Gilg in Mem. Torr. Bot. Club, 3(3): 34 (1898). M. parvifolia Urb. & Gilg in Rev. Mus. La Plata, 5: 289 (1893). M. soratensis Urb. & Gilg in Nov. Act. Acad. Caes. Leop. 76: 68 (1900). Sclerothrix fasciculata Pres], Symb. Bot. 2: 3, t. 53 (1833). QAgeaa aaa eanAaaa aagaaaaaa ana LYTHRACEAE Adenaria floribunda HBK. Nov. Gen. & Spec. 6: 188, t. 549 (1824). A. floribunda var. grisleoides (HBK.) Koehne in Mart. Fl. Bras. 13(2): 210 (1877). A. floribunda var. microphylla Koehne in Engler, Pflanzenr. IV, 216: 247 (1903) A. floribunda forma purpurata (HBK.) Koehne in Engler, Pflanzenr. IV, 216: 247 (1903). Ammannia auriculata Willd. Hort. Berol. 1: 7, t. 7 (1804). A. Friesii Koehne in Engler, Pflanzenr. IV, 216: 50 (1903). 137 Cuphea chiquitensis Herzog in Meded. Rijks Herbar. 27: 16 (1915). annaaaaaaaanaaaagaa anana . cordata R. & P. Syst. 119 (1798). glutinosa Cham. & Schlechtd. in Linnaea, 2: 369 (1827). ianthina Koehne in Mart. Fl. Bras. 13(2): 238 (1877). ianthina var. Bangii Koehne in Engler, Bot. Jahrb. 29: 163 (1900) = C. ianthina var. Rusbyana. ianthina var. Kuntzei Koehne in O. Ktze. Rev. Gen. 3(2): 96 (1898). ianthina var. Rusbyana Koehne in O. Ktze. Rev. Gen. 3(2): 96 (1898). ianthina var. Stuebelii Koehne in Engler, Bot. Jahrb. 29: 163 (1900). inaequalifolia Koehne in Mart. Fl. Bras. 13(2): 286, t. 51, fig. 6 (1877). longiflora Koehne in Mart. Fl. Bras. 13(2): 246, t. 42, fig. 2 (1877). mapiriensis Lourteig in Sv. Bot. Tidskr. 48: 83 (1954). Melvilla Lindl. in Bot. Reg. 10: t. 852 (1824). mesostemon Koehne in Mart. Fl. Bras. 18(2): 252, t. 43, fig. 5 (1877). pannoso-cortica Rusby in Bull. N. Y. Bot. Gard. 4: 361 (1907). petalosa Herzog in Meded. Rijks Herbar. 27: 17 (1915). Rusbyi Lourteig in Sv. Bot. Tidskr. 48: 84 (1954). scaberrima Koehne in Mart. Fl. Bras. 13(2): 238 (1877). setosa Koehne in Mart. FI. Bras, 13(2): 223 (1877). Spruceana var. Bangii Koehne in Mem. Torr. Bot. Club, 6: 39 (1896). thymoides Cham. & Schlechtd. in Linnaea, 2: 368 (1827). trichopetala Koehne in Mem. Torr. Bot. Club, 6: 40 (1896). verticillata HBK. Nov. Gen. & Spec. 6: 207, t. 552 (1824). Weddelliana Koehne in Engler, Bot. Jahrb. 4: 402 (1883). Heimia salicifolia (HBK.) Link, Enum. Pl. Hort. Berol. 2: 3 (1822). Lafoénsia punicaefolia DC. in Mém. Soc. Phys. Genéve, 3(2): 86, t. 1 (1826). L. speciosa (HBK.) DC. in Mém. Soc. Phys. Genéve, 3(2): 73 (1826). Lythrum maritimum HBK. Nov. Gen. & Spec. 6: 194 (1824). Parsonsia satureioides Rusby in Phytologia, 1: 69 (1934) — Cuphea sp. Physocalymma scaberrimum Pohl in Flora, 10: 153 (1827). Pleurophora annulosa Koehne in Engler, Pflanzenr. IV, 216: 181 (1903). is" anomala (St. Hil.) Koehne in Mart. Fl. Bras. 13(2): 307, t. 57 (1877). Rotala ramosior (L.) Koehne in Mart. FI. Bras. 13(2): 194, t. 39 (1877). BEGONIACEAE Begonia altoperuviana A. DC. in Ann. Sci. Nat. (ser. 4) 11: 123 (1859). 2). bo by bo eo De andina Rusby in Bull, N. Y. Bot. Gard. 8: 108 (191 Bangii O. Ktze. Rev. Gen. 3(2): 105 (1898). Baumannii Lemoine in Le Jardin, 1890: 273. boliviensis A. DC. in Ann. Sci. Nat. (ser. 4) 11: 122 (1859). Bridgesii A. DC. in Ann. Sci. Nat. (ser. 4) 11: 182 (1859). chaetocarpa O. Ktze. Rev. Gen. 3(2): 105 (1898). chaetocarpa var. glabriflora Smith & Schubert in Rev. Univ. Cuzco, 33(87): 76 (1944). juntasensis O. Ktze. Rev. Gen. 3(2): 106 (1898). lignosa Rusby, Descr. S. Am. Pls. 65 (1920) — B. unduavensis. micranthera var. fimbriata Smith & Schubert in Darwiniana, 5: 98 (1941). 138 micranthera var. foliosa Smith & Schubert in Darwiniana, 5: 92 (1941). micranthera var. Venturii Smith & Schubert in Darwiniana, 5: 97 (1941). tha Britton in Bull. Torr. Bot. Club, 18: 85 (1891) = B. parviflora. oblanceolata Rusby, Descr. S. Am. Pls. 65 (1920). octopetala L’Hérit. Stirp. Nov. 101 (1788). parviflora Poepp. & Endl. Nov. Gen. 1: 7, t. 12 (1835). Pearcei Hook. in Bot. Mag. 91: t. 5545 (1865). platanifolia var. acuminatissima O. Ktze. Rev. Gen. 3(2): 106 (1898). pleiopetala A. DC. in Ann. Sci. Nat. (ser. 4) 11: 121 (1859). santarosensis O. Ktze. Rev. Gen. 3(2): 106 (1898). subcaudata Rusby ex Smith & Schubert in Rev. Univ. Cuzco, 33(87): 81, t. 14 (1945) subrectangula Rusby in Phytologia, 1: 68 (1934) = B. juntasensis ? subvillosa Kl. Begon. 32 (1855). tovarensis Kl]. Begon. 31 (1855). unduavensis Rusby, Descr. S. Am. Pls. 64 (1920). unilateralis Rusby in Phytologia, 1: 68 (1934). varistyla Irmscher in Engler, Bot. Jahrb. 76: 80 (1953). Veitchii Hook. f. in Gard. Chron. 1867: 734. Weddelliana A. DC. in Ann. Sci. Nat. (ser. 4) 11: 122 (1859). Warburgiana Hieron. in Engler, Bot. Jahrb. 21: 325 (1895) = B. pleio- etala. Williamsii Rusby & Nash in Torreya, 6: 47 (1906). Wollneyi Herzog in Fedde, Repert. Spec. Nov. 7: 63 (1909) = B. Williamsii. DH DWP WWW Wee CACTACEAE Acanthorhipsalis crenata (Britton) Britt. & Rose, Cact. 4: 212 (1923). Aylostera Kupperiana (Béd.) Backebg. in Backebg. & Knuth, Kaktus-ABC, 275 (1935). ; Bolivicereus samaipatanus Cardenas in Cact. & Succ. Journ. 23: 91 (1951) B. samaipatanus var. divi-miseratus Cardenas in Nat. Cact. & Succ. Journ. 6 9 (1951). B. samaipatanus var. multiflorus Cardenas in Cact. & Succ. Journ. 23: 93 (1951). Castellanosia caineana Cardenas in Cact. & Succ. Journ. 23: 90 (1951). Cephalocereus Guentheri Kupp. in Monats. Deut. Kakt. Gesell. 3: 159 (1931). Cereus Baumannii Lem. in Hortic. Univ. 1844: 126, 315. C. dayami Speg. in Anal. Mus. Nac. Buenos Aires (ser. 3), 4: 480 (1905). C. peruvianus Mill. Gard. Dict. (ed. 8) no. 4 (1768). C. Spachianus Lem. Hort. Univ. 1: 225 (1839). C. Spegazzinii Weber in Monats. Kakteenk. 9: 102 (1899). C. stenogonus K. Schum. in Monats. Kakteenk. 9: 165 (1899). C. tacaquirensis Vaupel in Monats. Kakteenk. 26: 122 (1916). C. trigonodendron K. Schum. in Engler, Bot. Jahrb. 40: 413 (1908). ragga anu areolatus oe Riccob. in Bol. R. Ort. Bot. Giard. Col. Palermo, 8: 264 (1909). C. Srokeaoe 'Chrdenas in Cact. & Suce. Journ. 24: 144 (1952). C. Buchtienii Backebg. in Backebg. & Knuth, Kaktus-ABC, 189, 411 (1935). . Buchtienii var. flavispinus Cardenas in Cact. & Succ. Journ. 24: 182 (1952). . Candelilla Cardenas in Cact. & Suce. Journ. 24: 146 (1952). . Candelilla var. pojoénsis Cardenas in Cact. & Succ. Journ. 24: 147 (1952). . dependens Cardenas in Cact. & Succ. Journ. 24: 148 (1952). . Herzogianus Backebg. in Blatt. Kakteenf. 1934, 6: Genus 45: 2, fig. (1934). laniceps (K. Schum.) Goss. in Bull. Soc. Cent. Agric. Hort. Acclim. Nice, 44: 82 (1904). aaaaaa 139 . parapetiensis Cardenas in Cact. & Succ. Journ. 24: 182 (1952). . parviflorus (K. Schum.) Goss. in Bull. Soc, Cent. Agric. Hort. Acclim. Nice, 44: 32 (1904). . Roezlii (Hge. f.) Backebg. in Backebg. & Knuth, Kaktus-ABC, 190 (1935). 13). . sucrensis Cardenas in Cact. & Succ. Journ. 24: 148 (1952). . tominensis (Wegt.) Backebg. in Backebg. & Knuth, Kaktus-ABC, 190 (1935). . tupizensis (Vpl.) Backebg. in Backebg. & Knuth, Kaktus-ABC, 190 (1935). . Wendlandiorum Backebg. in Kakt. & Andere Sukkul. 6: 115 (1955). Corryocactus ayopayanus Cardenas in Revist. Agric. Cochabamba, 7: 21 (1952). C. melanotrichus (K. Schum.) Britt. & Rose, Cact. 2: 68 (1920). C. melanotrichus var. caulescens Cardenas in Revist. Agric. Cochabamba, 20 (1952). C. Perezianus Cardenas in Revist. Agric. Cochabamba, 7: 22 (1952). C. tarijensis Cardenas in Revist. Agric. Cochabamba, 7: 23 (1952). Cylindropuntia haematacantha Backebg. in Backebg. & Knuth, Kaktus-ABC, 118, 411 (1935). C. humahuacana Backebg. in Backebg. & Knuth, Kaktus-ABC, 119, 411 (1935). C. teres (Cels) Backebg. in Backebg. & Knuth, Kaktus-ABC, 119 (1935). C. Weingartiana (Backebg.) Backebg. & Knuth, Kaktus-ABC, 119 (1935). Echinocactus ambiguus Hildm. ex K. Schum. Gesamtb. Kakt. 311 (1898). E. pulverulentus Miihlenpf. in Allg. Gartenzeit. 16: 9 (1848). Echinopsis albispinosa K. Schum. in Monats. Kakteenk. 13: 154 (1903). Boedekeriana Hard. in Jahrb. Deut. Kakt.-Ges. 1: 70 (1936) Bridgesii Salm-Dyck, Cact. Hort. Dyck. 1849: 181 (1850). calochlora K. Schum. in Monats. Kakteenk. 13: 108 (1903). Fiebrigii Giirke in Notizbl. 4: 184 (1905). formosissima Lab. in Rev. Hort. (ser. 4) 4: 26 (1855). Hardeniana Béd. in Jahrb. Deut. Kakt.-Ges. 1: 24 (1935). Huottii (Cels) Lab. Monog. Cact. 301 (1853). mamillosa Giirke in Monats. Kakteenk. 17: 128 (1907). Mieckleyi R. Mey. in Monats. Kakteenk. 28: 122 (1918). obrepanda (Salm-Dyck) K. Schum. in Engl. & Prantl, Nat. Pflanzenfam. 3(6a): 184 (1894). potosina Werd. in Notizbl. 11: 267 (1981). Ritteri Bod. in Monats. Deut. Kakt.-Ges. 4: 141 (1932). Rojasii Cardenas in Revist. Agric. Cochabamba, 6: 31 (1951). Rojasii var. albiflora Cardenas in Revist. Agric. Cochabamba, 6: 33 (1951). Smrziana Backebg. in Backebg. & Knuth, Kaktus-ABC, 219, 412 (1935). Stollenwerkiana Bod. in Jahrb. Deut. Kakt.-Ges. 1: 59 (1936). Epiphyllum Bridgesii Lem. in Illustr. Hort. 8: Misc. 5 (1861). E. phyllanthus (L.) Haw. Syn. Pl. Suce. 197 (1812). Eriocereus tephracanthus (Lab.) Riccob. in Bol. R. Ort. Bot. Giard. Col. Pa- lermo, 8: 244 (1909). E. tephracanthus var. boliviensis (Web.) Marshall in Marshall & Bock, Cact. 98 (1941) MO) ©): ©: Ses wm ct ba) rt) c n 2 -o~ an) oO oO n co) _— : G. stellipilis Gleason in Amer. Journ. Bot. 19: 742 (1932). Heterotrichum octonum (Bonpl.) DG. Prodr. 3: 173 (1828). Henrietella sylvestris Gleason in Phytologia, 1: 108 (1934). Leandra aurea (Cham.) Cogn. in Mart. Fl. Bras. 14(4): 142 (1886). L. boliviensis Cogn. in Mem. Torr. Bot. Club, 3(3): 29 (1893). 145 carassana var. estrellensis (Raddi) Cogn. in Mart. Fl. Bras. 14(4): 120 (1886). crenata (D. Don) Cogn. in Mart. Fl. Bras. 14(4): 137 (1886). dichotoma (D. Don) Cogn. in Mart. Fl. Bras. 14(4): 200 (1886). erostrata DC.) Cogn. in Mart. Fl. Bras. 14(4): 139 (1886). hylophila Gleason in Bull. Torr. Bot. Club, 68: 247 (1941). reversa (DC.) Cogn. in Mart. Fl. Bras. 14(4): 198 (1886). stellulata Cogn. in Bull. Torr. Bot. Club, 17: 58 (1890). Maieta guianensis Aubl. Pl. Guian. Fr. 1: 443, t. 176 (1775). M. hispida Rusby in Mem. N. Y. Bot. Gard. 7: 315 (1927) = M. guianensis. Meriania axinaeoides Gleason in Amer. Journ. Bot. 19: 741 (19382). Peer hr oe M. amabilis Cogn. in Bull. N. Y. Bot. Gard. 4: 359 (1907). M. amplexans (Crueg.) Cogn. in Mart. Fl. Bras. 14(4): 256 (1887). M. amplexicaulis Naud. in Ann. Sci. Nat. (ser. 3) 16: 186 (1851). M. andina Naud. in Ann. Sci. Nat. (ser. 3) 16: 236 (1851). M. annulata (Naud.) Triana in Trans. Linn. Soc. 28: 121 (187 1). M. arborea Pav. ex Triana in Trans. Linn. Soc. 28: 108 (1871), in synon. M. argyrophylla DC. Prodr. 3: 181 (1828). M. axinaeoides Gleason in Bull. Torr. Bot. Club, 59: 368 (1932). M. Bangii Cogn. in Mem. Torr. Bot. Club, 3(3): 30 (1893). M. biacuta Cogn. in Mem. Torr. Bot. Club, 3(3): 31 (1893). M. biformis Cogn. in Meded. Rijks Herbar. 19: 80 (1913). M. biformis var. brevifolia Cogn. in Meded. Rijks Herbar. 19: 81 (1913). M. boliviensis Cogn. in DC. Monog. Phan. 7: 901 (1891). M. boliviensis var. glabra Cogn. in Mem. Torr. Bot. Club, 3(3): 30 (1893). M. Brittonii Cogn. in Bull. Torr. Bot. Club, 17: 92 (1890). M. Brittonii var. glabrata Cogn. in Bull. Torr. Bot. Club, 23: 277 (1896). M. Buchtienii Cogn. in Fedde, Repert. Spec. Nov. 8: 2 (1910). M. calvescens DC. Prodr. 3: 185 (1828). M. centrodesma Naud. in Ann. Sci. Nat. (ser. 3) 16: 164 (1851). M. cordata Triana in Trans. Linn. Soc. 28: 114 (1871). M. cremophylla Naud. in Ann. Sci. Nat. (ser. 3) 16: 228 (1851). M. cretacea Gleason in Amer. Journ. Bot. 19: 744 (1932). M. cuspidata Naud. in Ann. Sci. Nat. (ser. 3) 16: 163 (1871). M. cyanocarpa Naud. in Ann. Sci. Nat. (ser. 3) 16: 221 (1851). M. cyanocarpa var. hirsuta Cogn. in Bull. Torr. Bot. Club, 23: 277 (1896). M. cyanocarpa var. parvifolia Cogn. in Bull. Torr. Bot. Club, 23: 17 (1896). M. desmantha Benth. Pl. Hartw. 181 (1845). M. dipsacea Naud. in Ann. Sci. Nat. (ser. 3) 16: 138 (1851). ? M. discolor DC. Prodr. 3: 114 (1828). M. dispar Benth. in Hook. Kew Journ. 2: 241 (1850). M. dodecandra (Desr.) Cogn. in Engler, Bot. Jahrb. 8: 21 (1887). M. dolichorhyncha Naud. in Ann. Sci. Nat. (ser. 3) 16: 166 (1851). M. elaeagnoides Cogn. in Mart. Fi. Bras. 14(4): 390 (1887). M. elongata Cogn. in Bull. Torr. Bot. Club, 17: 91 (1890). M. erioclada Triana in Trans. Linn. Soc. 28: 111 (1871). M. erioneura Cogn. in Mem. Torr. Bot. Club, 6: 38 (1896). 146 M. flavescens Cogn. in Bull. Torr. Bot. Club, 17: 93 (1890). M. glandulifera var. boliviensis Cogn. in Mem. Torr. Bot. Club, 6: 38 (1896). . SERREEERERE Ss glomerulifera Cogn. in Bull. Torr. Bot. Club, 23: 15 (1896). granulosa (Bonpl.) Naud. in Ann. Sci. Nat. (ser. 3) 16: 218 (1851). guianensis (Aubl.) Cogn. in Jahrb. Bot. Gart. Berlin, 4: 280 (1886). Herrerae Gleason in Amer. Journ. Bot. 19: 745 (1932). Herzogii Cogn. in Meded. Rijks Herbar. 19: 80 (1913). irta Cogn. in Mem. Torr. Bot. Club, 3(3): 31 (1893). holosericea (L.) DC. Prodr. 3: 181 (1828). holosericea var. bracteata (DC.) Cogn. in Mart. Fl. Bras. 14(4): 237 (1887). hygrophila Naud. in Ann. Sci. Nat. (ser. 3) 16: 220 (1851). ibaguensis (Bonpl.) Triana in Trans. Linn. Soc. 28: 110 (1871). impetiolaris (Sw.) D. Don in Mem. Wern. Soc. 4: 316 (1828). juruensis Pilger in Verh. Bot. Ver. Brandenb. 47: 172 (1905). Kuntzei Cogn. in O. Ktze. Rev. Gen. 3(2): 93 (1898). lanata (DC.) Triana in Trans. Linn. Soc. 28: 113 (1871). lasiocalyx Cogn. in Bull. Torr. Bot. Club, 23: 278 (1896). latistigma Cogn. in Bull. N. Y. Bot. Gard. 4: 360 (1907). lepidota DC. Prodr. 3: 180 (1828). livida Triana in Trans. Linn. Soc. 28: 121 (1871). longicuspis Herzog in Fedde, Repert. Spec. Nov. 7: 64 (1909). longifolia (Aubl.) DC. Prodr. 3: 184 (1828). Mandonii Cogn. in DC. Monog. Phan. 7: 911 (189 mapirensis Gleason in Bull. Torr. Bot. Club, 68: 249 (1941). Matthaei Naud. in Ann. Sci. Nat. (ser. 3) 16: 176 (1851). megaphylla Gleason in Bull. Torr. Bot. Club, 59: 363 (1932). micrantha Cogn. in Bull. Torr. Bot. Club, 23: 16 (1896). minutiflora (Bonpl.) DC. Prodr. 3: 189 (1828). molybdea Naud. in Ann. Sci. Nat. (ser. 3) 16: 185 (1851). molybdea var. titanea (Naud.) Cogn. in DC. Monog. Phan. 7: 827 (1891). multiflora Cogn. in Bull. Torr. Bot. Club, 17: 60 (1890). myriantha Benth. in Hook. Journ. Bot. 2: 314 (1840). nervosa (Smith) Triana in Trans. Linn. Soc. 28: 111 (1871). novemnervia Naud. in Ann, Sci. Nat. (ser. 3) 16: 216 (1851). obliqua Gleason in Amer. Journ. Bot. 19: 743 (1932). obovalis Naud. in Ann. Sci. Nat . (ser. 3) 16: 183 (1851). papillosa (Desr.) Naud. in Ann. Sci. Nat. (ser. 3) 16: 216 (1851). Pentlandii Naud. in Ann. Sci. Nat. (ser. 3) 16: 182 (1851). persicariaefolia Cogn. in Bull. Torr. Bot. Club, 17: roi (1890). plumifera Triana in Trans. Linn. Soc. 28: 124 (1871). plumifera var. Bangii Cogn. in Bull. Torr. Bot. Club, 23: 16 (1896). Poeppigii Triana in Trans. Linn. Soc. 28: 107 (1871). polygama Cogn. in Bull. Torr. Bot. Club, 17: 92 (1890). prasina (Sw.) DC. Prodr. 3: 188 (1828). pteropoda Benth. in Hook. Journ. Bot. 2: 314 (1840). pubicalyx Gleason in Amer. Journ. Bot. 19: 742 (1932). punctata (Desr.) D. Don in Mem. Wern. Soc. 4: 316 (1828). resima Naud. in Ann, Sci. Nat. (ser. 8) 16: 226 (1851). rosea Gleason in Bull. Torr. Bot. Club, 68: 247 (1941). rubiginosa (Bonpl.) DC. Prodr. 3: 183 (1828). rufescens (Aubl.) DC. Prodr. 3: 180 (1828). Ruizii Naud. in Ann. Sci. Nat. (ser. 3) 16: 192 (1851). Rusbyana Cogn. in Bull. Torr. Bot. Club, 17: 93 (1890). seabriuscula Cogn. in Meded. Rijks Herbar. 19: 80 (1913). serialis DC. Prodr. 3: 182 (1828). serrulata (DC.) Naud. in Ann. Sci. Nat. (ser. 3) 16: 118 (1851). 147 sessilifolia Naud. in Ann. Sci. Nat. (ser. 3) 16: 181 (1851). spennerostachya Naud. in Ann. Sci. Nat. (ser. 3) 16: 187 (1851). spennerostachya var. angustifolia Cogn. in Bull. Torr. Bot. Club, 17: 59 staphidioides (Naud.) Triana in Trans. Linn. Soc. 28: 102 (1871). Steinbachii Markgraf in Notizbl. 9: 1158 (1927). stellipilis Cogn. in Bull. N. Y. Bot. Gard. 4: 360 (1907). stellulata Gleason in Bull. Torr. Bot. Club, 68: 247 (1941). stenocardia Cogn. in Meded. Rijks Herbar. 19: 79 (1918). stenostachya DC. Prodr. 3: 181 (1828). subnodosa Triana in Trans. Linn. Soc. 28: 106 (1871). ternatifolia Triana in Trans. Linn. Soc. 28: 118 (1871). tetrasperma Gleason in Bull. Torr. Bot. Club, 58: 237 (1931). theaezans var. glaberrima (DC.) Cogn. in Mart. Fl. Bras. 14(4): 420 (1888). eaezans var. subtriplinervia Cogn. in Mart. Fl. Bras. aby 421 (1888). tiliaefolia Naud. in Ann. Sci. Nat. (ser. 3) 16: 151 (185 tomentosa (L. C. Rich.) D. Don in Mem. Wern. Soc. 4: a (1823). triplinervia R. & P. Syst. 105 (1798). tunicata (Bonpl.) Naud. in Ann. Sci. Nat. (ser. 3) 16: 243 (1851). undata var. robusta Cogn. in Bull. Torr. Bot. Club, 23: 277 (1896). uvifera Naud. in Ann. Sci. Nat. (ser. 3) 16: 222 (1851 valida Cogn. in Bull. Torr. Bot. Club, 28: 278 (1896). ? M. Wagneri Macbr. in Field Mus. Publ. Bot. a 179 (1929). iE eee ee aoe DC. Prodr. 3: 120 (18 M. peruviana Cogn. in Mart. Fl. Bras. 14(3): is (1883). Manette parva Benth. Bot. Voy. Sulphur, 97, t. 36 (1845). M. princeps Naud. in Ann. Sci. Nat. (ser. 3) 18: 283 (1852). Ossaea boliviensis (Cogn.) Gleason in Bull. Torr. Bot. en 58: 259 (1931). O. Buchtienii Gleason in Amer. Journ. Bot. 19: 751 (19 O. petiolaris (Naud.) Triana in Trans. Linn. Soc. 28: re (1871). O. secundiflora Cogn. in Bull. N. Y. Bot. Gard. 4: 361 Nets Pterolepis boliviensis Cogn. in Fedde, Repert. Spec. Nov. 8: ; P. glomerata var. angustifolia Cogn. in Mart. Fl. Bras. err 275 (1885). P. glomerata var. peruviana Cogn. in Mart. Fl. Bras. 14(3): 276 (1885). P. pumila (DC.) Cogn. in Mart. FI. Hebe 14(3): 263 (1885). P P . eres SRRBSRSSSSS SSE . trichotoma (Rottb.) Cogn. in Mart. Fl. Bras, 14(3): 261 (1885). . Weddelliana Triana in Trans. Linn. Soc. 28:39 (1871 R. parvifolia Cogn. in O. Ktze. Rev. Gen. 3(2): 95 (1898). R. secundiflora Naud. in Ann. Sci. Nat. (ser. 3) 12: 207 (1849). Tibouchina adenophora Cogn. in Bull. N. Y. Bot. Gard. 4: 355 (1907). alpestris Cogn. in Meded. Rijks Herbar. 19: 78 (1913). amoena Herzog in Fedde, Repert. Spec. Nov. 7: 64 (1909). aurea Cogn. in Meded. Rijks Herbar. 19: 77 (1913). barbigera (Naud.) Baill. in Adansonia, 12: 75 (1877) bicolor (Triana) Cogn. in DC. Monog. Phan. 7: 265 (1891). brachyphylla Gleason in Ph vlolovia 1: 186 (1935). Brittoniana Cogn. in Bull. Torr. Bot. Club, 17: 55 (1890). calycina var. parvifolia Cogn. in Engler, Bot. Jahrb. 42: 135 (1908). capitata (Naud.) Cogn. in Bull. Torr. Bot. Club, 17: 54 (1890). citrina (Triana) Cogn. in DC. Monog. Phan. 7: 253 (1891). confertifiora (Triana) Cogn. in DC. Monog. Phan. 7: 253 (1891). cornuta Gleason in Bull. Torr. Bot. Club, 66: 415 (1939). excoriata Cogn. in Bull. N. Y. Bot. Gard. 4: 356 (1907). 148 gracilis Cogn. in Mart. Fl. Bras. 14(3): 386 (1885). granulosa (Desr.) Cogn. in Mart. Fl. Bras. 14(3): 340 (1885). granulosa var. angustifolia Britton in Bull. Torr. Bot. Club, 17: 54 (1890). grossa (L. f.) Cogn. in Mart. Fl. Bras. 14(3): 297 (1885). Herzogii Cogn. in Meded. Rijks Herbar. 19: 78 (1913). lanceolata Cogn. in Bull. Torr. Bot. Club, 17: 55 (1890). lancifolia Wurdack in hig eae . 54 (1954). latifolia (Naud.) Britton in Bull. Torr. Bot. Club, 17: 54 (1890). longifolia (Vahl) Baill. in RGR es 74 (1877). membranifolia Cogn. in Bull. Torr. Bot. Club, 23: 17 (1896). obtusifolia Cogn. in Bull. N. Y. Bot. Gard. 4: 355 (1907). octopetala Cogn. in Bull. Torr. Bot. Club, 17: 57 (1890). oligantha Gleason in Phytologia, 1: 134 (1935). Orbignyana (Naud.) Cogn. in DC. Monog. Phan. 7: 261 (1891). panicularis (Naud.) Britton in Bull. Torr. Bot. Club, 17: 54 (1890). paratropica (Griseb.) Cogn. in DC. Monog. Phan. 7: 257 (1891). pentamera (Ule) Macbr. in Field Mus. Publ. Bot. 13(4): 290 (1941). purpurascens Cogn. in Bull. Torr. Bot. Club, 17: 15 (1890) Rusbyi Cogn. in Bull. Torr. Bot. Club, 17: 54 (1890). stenocarpa Cogn. in Mart. Fl. Bras. 14(3): 344 (1885). stenocarpa var. boliviensis Cogn. in Mart. FI. wire 14(8): 345 (1885). . stenopetala Cogn. in DC. Monog. Phan. 7: 262 (1891). stenophylla Cogn. in Bull. Torr. Bot. Club, 17: 56 (1890). tetrapetala Cogn. in Bull. Torr. Bot. Club, 23: 276 (1896). venosa Gleason in Phytologia, 1: 134 (19385) Tococa carolensis Gleason in Amer. Journ. Bot. 19: 746 (1932). T. coronata Benth. in Hook. Journ. Bot. 2: 303 (1840). T. guianensis Aubl. Pl. Guian. Fr. 1: 488 (1775). T. occidentalis Naud. in Ann. Sci. Nat. (ser. 3) 16: 92 (1851). ei orcheophora Naud. in Ann. Sci. Nat. (ser. 3) 16: 89 (1851). Topobea boliviensis Cogn. in O. Ktze. Rev. Gen. 3(2): 94 (1898) = Blakea multiflora. ONAGRACEAE Epilobium andicola Hausskn. in Oesterr. Bot. Zeitschr. 29: 118 (1879). Asplundii Samuelsson in Sv. Bot. Tidskr. 17: 256 (1923). bolivianum Samuelsson in Sv, Bot. Tidskr. 17: 263 (1923). deminutum Samuelsson in Sv. Bot. Tidskr. 17: 264 (1923). denticulatum R. & P. Fl. Peruv. 8: 78, t. 314 (1802). denticulatum var. aberrans Samuelsson in Sv. Bot. Tidskr. 17: 253 (1923). denticulatum var. confertum Samuelsson in Sv. Bot. Tidskr. 17: 252 (1923). denticulatum var. macropetalum Samuelsson in Sv. Bot. Tidskr. 17: 253 (1923). fragile Samuelsson in Sy. Bot. Tidskr. 17: 291 (1923). Haenkeanum Hausskn. in Oesterr. Bot. Zeitschr. 29: 148 (1879). hirtum Samuelsson in Sv. Bot. Tidskr. 17: 266 (1923). Fuchsia boliviana Carr. in Rev. Hort. 48: 150 (1876). F. boliviana forma puberulenta Munz in Proc. Calif. Acad. Sci. (ser. 4) 25: 1943). pst Do bb bt F. — Britton in Bull. Torr. Bot. Club, 17: 214 (1890) — F. sanctae- F. Brittonii I. M. Johnst. in Contrib. Gray Herb. 75: 39 (1925) — F. sanctae- rosae. F. denticulata R. & P. Fl. Peruv. 3: 87, t. 325, fig. b (1802). F. filipes Rusby in Mem. N. Y. Bot. Gard. 7: 8317 (1927) = F. sanctae-rosae. F. furfuracea I. M. Johnst. in Contrib. Gray Herb. 75: 39 (1925). 149 Garleppiana O. Ktze. & Whittmack in Gartenfl. 47: 461, fig. 96 (1893). hirsuta Hemsl. in Journ. Bot. 14: 69 (1876). juntasensis O. Ktze. Rev. Gen. 3(2): 97 (1898). salicifolia Hemsl. in Journ. Bot. 14: 70 pha sanctae-rosae O. Ktze. Rev. Gen. 3(2): 98 (18 —— I. M. Johnst. in Contrib. Gray Bore he 38 (1925) — F. juntas- nsis. Meonsent O. Ktze. Rev. Gen. 3(2): 98 (1898). unduavensis Munz in Proc. Calif, Acad. Sci. (ser. 4) 25: 81 (1943). phe crs boliviana Rusby, Descr. S. Am. Pls. 72 (1920) = Oenothera multi- cauli Sunn affinis DC. Prodr. 3: 53 (1828). densiflora Micheli in ro 57: 301 (1874). erecta L. Sp. Pl. 388 (1753). ferruginea Rusby in Bull. N. Y. Bot. Gard. 8: 110 (1912) — J. affinis. leptocarpa forma biacuminata (Rusby) Munz in Darwiniana, 4: 257 (1942). lithospermifolia Kunth ex Micheli in Flora, 57: 300 (1874). longifolia DC. in Mém. Soc. Phys. es 2(2): 141 Siig natans Humb. & Bonpl. Pl. Aequin. 1: 16, t. 3, fig. b (1805). nervosa var. marginata (Rusby) Munz in Datwinians, 4: 210 (1942). peruviana L, Sp. Pl. 388 (1753). repens var. peploides (HBK.) Griseb. Cat. Pl. Cubens. 107 (1866). sedioides Humb. & Bonpl. Pl. Aequin. 1: 18, t. 3, fig. a (1805). suffruticosa var. ligustrifolia (HBK.) Griseb. in Mem. Amer. Acad. 8: 187 (1860). suffruticosa var. ligustrifolia forma linearifolia (Hassl.) Munz in Darwini- ana, 4: 243 (1942). . uruguayensis Camb. in St. Hil. Fl. Bras. Mer. 2: 264 (1829). acumensis Rusby in Mem. N. Y. Bot. Gard. 7: 316 (1927) = J. litho- spermifolia. Oenothera albicans Lam. Encycl. 4: a (1797). Oe. Barbeyana Lévl. Monog. 48 (1902) = Oe. multicaulis. Oe. campylocalyx Koch & Bouché, Ind. Sem. Hort. Berol. App. 17 (1855). Oe. coccinea Britton in Bull. Torr. Bot, Club, 17: 218 (1890) = Oe. campylo- cal HUSH * SSH alyx. Oe. Ebates Rusby in Mem. Torr. Bot. oe 8(3): 38 (1893). Oe. Mandonii Lévl. Monog. 43 (1902) = Oe. multicaulis R. & P. FI. Peruv. 3: 80, t. e (1802). Oe. nana Griseb, in Goett. Abh. 19: 143 (1874). Oe. punae O. Ktze. Rev. Gen. 3(2): 99 (1898) = Oe. nana. Oe. rosea Ait. Hort. Kew. 2: 3 (1789). Oe. rubida Rusby in Bull. N. Y. Bot. Gard. 8: 110 (1912). Oe. scabra Krause in Fedde, Repert. Spec. Nov. 1: 168 (1905). Oe. serratifolia Krause in Fedde, Repert. Spec. Nov. 1: 168 (1905) = Oe«e. elongata. HALORRHAGIDACEAE Gunnera apiculata Schindl. in Engler, Pflanzenr. IV, 225: 121 (1905). G. boliviana Morong ex Rusby in Mem. Torr. Bot. Club, 6: 33 (1896). G. magellanica Lam. Encycl. 3: 61 (1789). G. pilosa Kunth, Synops. Pl. Aequin. 1: 378 ( 1822). G. scabra R. & P. Fl. Peruv. 1: 29, t, 44, fig. a (1798). Myriophyllum elatinoides Gaudich. in Ann. Sci. Nat. (ser. 1) 5: 105 (1825). M. pallidum Rusby in Bull. N. Y. Bot. Gard. 8: 110 (1912). M. titikakense Remy in Ann. Sci. Nat. (ser. 3) 6: 352 (1846). 150 ARALIACEAE Aralia soratensis March. in Bull. Acad. Belg. (ser. 2) 47: 75 (1879). Dendropanax arboreum (L.) Decne. & Pl. in Rev. Hort. (ser. 4) 3: 107 (1854). D. bolivianus Gandog. in Bull. Soc. Bot. France, 65: 33 (1918). D. oblongifolium Rusby in Bull. N. Y. Bot. Gard. 4: 366 (1907 ya Didymopanax allocotanthus Harms in Notizbl. 11: 292 (19381). D. Kuntzei Harms in O. Ktze. Rev. Gen. 3 (2): 115 (1898). D. Morototoni (Aubl.) Decne. & Pl. in Rev. Hort. (ser. 4) 3: 109 (1854). D. nebularum Harms in Notizbl. 11: 298 (1931). Gilibertia macropoda Harms in Notizbl. 9: 980 (1926). G. Pavonii (Decne. & Pl.) March. in Mart. Fl. Bras. 11(1): 248 (1878). Oreopanax artocarpoides Standl. in Field Mus. Publ. Bot. 17: 205 (1937). atopanthum Harms in Notizbl. 9: 1040 (1926). . boliviense Seem. in Journ. Bot. 3: 272 (1865). fulvum March. in Mart. Fl. Bras. 11(1): 254 (1878). grosseserratum Rusby in Bull. N. Y. Bot. Gard. 4: 367 (1907). Herzogii Harms in Notizbl. 15: 686 (1942). Kuntzei Harms in O. Ktze. Rev. Gen. 3(2): 116 (1898). macrocephalum Decne. & Pl. ex Wedd. Chlor. And. 2: 207 (1861). membranaceum Rusby in Mem. Torr. Bot. Club, 3(3): 42 (1893). Pentlandianum Decne. & Pl. ex Wedd. Chlor. And. 2: 207 (1861). Rusbyi Britton in Bull. Torr. Bot. Club, 18: 38 (1891). Steinbachianum Harms in Notizbl. 9: 1041 (1926). thaumasiophyllum Harms in Notizbl. 11: 291 (1931). Trollii Harms in Notizbl. 11: 292 (1981). Pentapanax angelicifolium Griseb. in Goett. Abh. 24: 144 (1879). Schefflera Bangii Harms in Notizbl. 11: 285 (1931). meena Harms in Notizbl. 11: 286 (1931). gii Harms in Meded. Rijks Herbar. 29: 6 (1916). nephelophiti Harms in Notizbl. 11: 286 (1931). patula (Rusby) Harms in Engl. & Prantl, Nat. Pflanzenfam. 3(8): 37 2999999999999 Hy: tipuanica Harms in Notizbl. 11: 287 (1931). Trollii Harms in Notizbl. 11: 287 (1931). Sciadophyllum conicum Poir, in Lam. Encycl. 6: 746 (1804). S. paniculatum Britton in Bull. Torr. Bot. Club, 18: 37 (1891) — Didymopanax a wn orototoni. S. pentandrum (R. & P.) Poir. in Lam. Encycl. 6: 747 (1804). UMBELLIFERAE Apium leptophyllum (Pers.) F. Muell. in Benth. Fl. Austral. 3: 372 (1866). Arracacia andina Britton in Bull. Torr. Bot. Club, 18: 37 (1891). Azorella biloba Wedd. Chlor. And. 2: 195, t. 66B (1860). A. biloba var. strigosa Wedd. Chlor. And. 2: 196 (1860). columnaris Wolff in Engler, Bot. Jahrb. 40: 288 (1908). diapensioides A. Gray in Bot. U. S. Explor. Exped. 1: 702 (1854). diapensioides var. denticulata Wedd. Chlor. And. 2: 191 (1860). monanthos Clos in C. Gay, Fl. Chile, 3: 79 (1847). multifida (R. & P.) Pers. Synops. Pl. 1: 303 (1805). prismatoclada Domin in Fedde, Repert. Spec. Nov. 4: 297 (1907). pulvinata Wedd. Chlor. And. 2: 194, t. 66A (1869). Azorellopsis trisecta Wolff in Fedde, Repert. Spec. Nov. 19: 312 (1924). Bowlesia acutiloba Wolff in Engler, Bot. Jahrb. 40: 284 (1908). B. incana forma tenera (Spreng.) Urb. in Mart. Fl. Bras. 11(1): 292 (1879). lage ca ol ok ae B. lobata R. & P. Fl. Peruv. 3: 28, t. 251, fig. b (1802). B. Mandonii Rusby in Mem. Torr. Bot. Club, 3(3): 41 ee B. palmata R. & P. Fl. Peruv. 3: 28, t. 251, fig. a (1802 B. palmata var. acutangula (Benth.) O. Ktze. Rev. ld 3(2): 112 (1898). B. palmata var. lacerata O. Ktze. Rev. Gen. 3(2): 112 (1898). B. platanifolia Wolff in Engler, Bot. Jahrb. 40: 284 (1908). B. pulchella Wedd. Chlor. And. 2: 188, t. 67B (1860). Centella asiatica (L.) Urb. in Mart. Fl. Bras. 11(1): 287 (1879). Conium maculatum L. Sp. Pl. 243 (1753). Daucus montanus Willd. ex Schult. Syst. 6: 482 (1820). Eryngium andicola Wolff in Engler, Bot. Jahrb. 40: 296 (1908). Boissieuanum Wolff in Fedde, Repert. Spec. Nov. 7: 275 (1909). Buchtienii Wolff in Fedde, Repert. Spec. Nov. 6: 24 (1908). Delarocheanum Wolff in Engler, Pflanzenr. IV, 228: 251 (1913). ebracteatum Lam. Encycl. 4: 759 (1796). bracteatum var. poterioides (Griseb.) Urb. in sien 43: 297 (1882). elegans Cham. & Schlechtd. in Linnaea, 1: 348 (1826). elegans var. boliviense Urb. in Mart. Fl. Bras. 11(1): 311 (1879). foetidum L. Sp. Pl. 232 (1753). glossophyllum Wolff in Engler, Bot. Jahrb. 40: 301 (1908). nudicaule Lam. Encycl. 4: 759 (1796). nudicaule var. peruvianum Urb. in Mart. Fl. Bras. 11(1): 304 (1879). paniculatum Laroche Eryng. 59, t. 26 (1808). paniculatum var. atrocephalum O. Ktze. Rev. Gen. 3(2): 112 (1898). ydrocotyle acuminata Urb. in Mart. Fl. Bras. 11(1): 273, t. 72, fig. 4 (1879). acuminata var. pubescens Mathias & Constance in Bull. Torr. Bot. Club, 78: 300 (1951). boliviana (O. Ktze.) Mathias in eso 2: 210 (1936). bonariensis Lam. Encycl. 3: 153 (1789). eccentrica Britton in Bull. Torr. Bot. ane 18: 36 (1891) = H. acuminata. filipes Mathias in Brittonia, 2: 230 (1936). grossulariaefolia var. Buchtienii Mathias in Brittonia, 2: 227 (1936). incrassata R. & P. Fl. Peruv. 3: 26 (1802). leucocephala Cham. & Schlechtd. in Linnaea, 1: 364 (1826). peruviana Wolff in Engler, Bot. Jahrb. 40: 283 (1908). pusilla A. Rich. in Ann. Gén. Sci. Phys. 4: 167, t. 52, fig. 2 (1820). ranunculoides L. f. Suppl. 177 (1781). H. ranunculoides forma minima O. Ktze. Rev. Gen. 8(2): 114 (1898) = H. ranunculoides. H. tenerrima var. puberula Mathias in Brittonia, 2: 209 (1936). H. umbellata L. Sp. Pl. 234 (1753). Lilaeopsis andina A. W. Hill in Journ. Linn. Soc. 47: 540, t. 20, fig 3 (1927). L. andina var. inundata A. W. Hill in Journ. Linn. Soc. 47: 541 (19 L. andina forma subulata (Wedd.) Pérez-Moreau in Lilloa, 1: 292 (1937 ). Niphogeton dissecta (Benth.) Macbr. in Field Mus. Publ. Bot. 8: 125 (1930). Oreomyrrhis andicola (Lag.) Hook. f. Fl. Antarct. 2: 288 (1847). Oreosciadium andinum Rusby in Bull. N. Y. Bot. Gard. 4: 366 (1907) = Api- um leptophyllum Sanicula mexicana DC. Prodr. 4: 84 (1830). Spananthe paniculata Jacq. Collect. 3: 247 (1789). ES Ag 0 a kanae ties bo gat ted ed tet ed tet ed tt ttt CORNACEAE Cornus boliviana Macbr. in Trop. Woods, 19: 5 (1929) = C. peruviana. C. peruviana Macbr. in Trop. Woods, 19: 5 (1929). 152 ERICACEAE Befaria boliviensis Fedtsch. & Basil. in Bot. Gaz. 85: 318 (1928) — B. glauca var. setosa. B. denticulata Remy in Ann. Sci. Nat. (ser. 3) 8: 230 (1847) = B. hispida. B. glauca Humb. & Bonpl. Pl. Aequin. 2: 118, t. 117 (1809). B. glauca var. coarctata (Humb. & Bonpl.) Mansf. & Sleum. in Notizbl. 12: 253 (1935). B. glauca var. glandulosa Mansf. & Sleum. in Notizbl. 12: 257 (1935). B. glauca var. setosa Mansf. & Sleum. in Notizbl. 12: 256 (1935). B. glauca var. tomentella Mansf, & Sleum. in Notizbl. 12: 255 (1935). B. hispida Poepp. & Endl. Nov. Gen. 1: 24, t. 39 (1835). B. parvifolia Rusby in Bull. N. Y. Bot. Gard. 8: 110 (1912) = B. glau Cavendishia acuminata (Hook.) Hemsl. in Biol. Centr. Am. Bot. 2: 272 "(eaty, C. Beckmanniana Hoer. in Engler, Bot. Jahrb. 42: 329 (1909). C. Martii (Meissn.) A. C. Smith in Bull. Torr. Bot. Club, 63: 313 (1936). C. paniculata Rusby in Mem. Torr. Bot, Club 4(3): 215 (1895) = C. Martii. C. pubescens (HBK.) Hemsl. in Biol. Centr. Am. Bot. 2: 273 (1881). C. pubescens var. boliviensis Hoer. in Engler, Bot. Jahrb. 42: 332 (1909). C. sillarensis Herzog in Meded. Rijks Herbar. 27: 20 (1915). C. strobilifera (HBK.) Hoer. in Engler, Bot. Jahrb. 42: 279 (1909). Ceratostemma Hookeri Britton in Bull. Torr. Bot. Club, 20: 187 (1893) = Siphonandra elliptica. C. pee sid Britton in Bull. Torr. Bot. Club, 20: 137 (1893) = Thibaudia ocalyx Chapaton viriiftorum O. Ktze, Rev. Gen. 3(2): 190 (1898) — Cavendishia cea Sicsaakaa Cham. in Linnaea, 8: 510 (1853). Bridgesii B. Fedtsch. in Sv. Bot. Tidskr. 18: 488 (1924). cuneata Rusby in Bull. N. Y. Bot. Gard. 4: 314 (1907). elongata Rusby in Bull. N. Y. Bot. Gard. 4: 405 (1907). fimbriata HBK. Nov. Gen. & Spec. 3: 290, t. 264 (1819). laevigata Meissn. in Mart. Fl. Bras. 7: 168 (1863). micrantha Remy in Ann. Sci. Nat. (ser. 3) 8: 233 (1847). Demosthenesia fabulosa (Sleum.) A. C. Smith in Bull. Torr. Bot. Club, 63: 310 (1936). D. Graebneriana (Hoer.) A. C. Smith in Bull. Torr. Bot. Club, 63: 310 (1936). D. Mandonii (Britton) A. C. Smith in Bull. Torr. Bot. Club, 63: 311 (1936). D. Pearcei (Britton) A. C. Smith in Bull. Torr. Bot. Club, 63: 310 (1936). O. spectabilis (Rusby) A. C. Smith in Bull. Torr. Bot. Club, 63: 311 (1936). Disterigma alaternoides var. parvifolium (Benth.) A. C. eomrths in Brittonia, 1: 220 (1933). D. ovatum (Rusby) Blake in Journ. Wash. Acad. Sci. 16: 365 (1926). D. pallidum A. C. Smith in Brittonia, 1: 222 (1933). D. pernettyoides (Griseb.) Ndzu. in Engler, Bot. Jahrb. 11: 224 (1889). Eleutherostemon bolivianum (Britton) Herzog in Meded. Rijks Herbar. 27: 23 (1915). E. racemosum Herzog in Meded. Rijks Herbar. 27: 22 (1915). Gaultheria anastomosans (L. f.) HBK. Nov, Gen. & Spec. 3: 283 (1819). barosmoides Rusby in Bull. N. Y. Bot. Gard. 4: 313 (1907). brachybotrys DC. Prodr. 7: 595 (1839). conferta Benth. Pl. Hartw. 219 (1846). cordifolia HBK. Nov. Gen. & Spec. 3: 285, t. 261 (1819). formosa Remy in Ann, Sci. Nat. (ser. 3) 8: 231 (1847). glabra DC. Prodr. 7: 596 (1839). mucronata Remy in Ann. Sci. Nat. (ser. 3) 8: 282 (1847) = G. Remyana. anaaaa ANQAARLALA 153 odorata Willd. in Ges. Naturf. Freunde Neue Schr. 3: 425 (1801). pichinchensis Benth. Pl. Hartw. 225 (1846). Remyana A. C. Smith in Contrib. U. S. Nat. Herb. 29: 347 (1950). reticulata HBK. Nov. Gen. & Spec. 3: 284 (1819). rufescens DC. Prodr. 7: 595 (1839). saxicola Wedd. Chlor. And. 2: 174 (1860). secunda Remy in Ann. Sci. Nat. (ser. 3) 8: 231 (1847). serrulata Herzog in Meded. Rijks Herbar. 27: 19 (1915). tetriches Rusby, Descr. S. Am. Pls. 74 (1920). . tomentosa HBK. Nov. Gen. & Spec. 3: 287, t. 262 (1819). vaccinioides Griseb. ex Wedd. Chlor. And. 2: 176 (1860). Gaylussacia Cardenasii A. C. Smith in Journ. Wash. Acad. Sci. — 206 (1958). G. ee Cham. & Schlechtd. in Linnaea, 1: 535 (1826). nia boliviensis O. Ktze. Rev. Gen. 3(2): 191 (1898) — Thibaudia sei ata, Leucothoé boliviensis Sleum. in Notizbl. 12: 131 (19384). Orthaea boliviensis Fedtsch. & Basil. in Not. Syst. Herb. Hort. Bot. USSR, 6: 25 (1926). O. constans A. C. Smith in Contrib. U. S. Nat. Herb. 28: 512 (1932). O. Weberbaueri Hoer. in Engler, Bot. Jahrb. 42: 320 (1909). Pernettya densa Rusby in Phytologia, 1: 70 (1984) = P. prostrata. . phyllyraefolia (Pers.) DC. Prodr. 7: 587 (1839). prostrata (Cav.) Sleumer in Notizbl. 12: 289 (1935). prostrata var. Pentlandii (DC.) Sleumer in Notizbl. 12: 290 (1935). prostrata var. purpurea (Don) Sleumer in Notizbl. 12: 290 (1935). pret Rusby in Phytologia, 1: 70 (1934) = P. prostrata var. Pent- QAAANANHAAAAD ty td td db lan Polyclita ‘ell (O. Ktze.) A. C. Smith in Bull. Torr. Bot. Club, 63: 314 (1935). Psammisia elliptica (Rusby) A. C. Smith in Contrib. U. S. Nat. Herb. 28: 395 1932). P. guianensis Klotzsch in Linnaea, 24: 43 (1851). P. pauciflora Griseb. ex A. C. Smith in Contrib. U. S. Nat. Herb. 28: 394 (19382). Rusbya taxifolia Britton in Bull. Torr. Bot. Club, 20: 68 (1893). Satyria neglecta A. C. Smith in Contrib. U. S. Nat. Herb. 28: 520 (1932). Siphonandra elliptica (R & P.) Klotzsch in Linnaea, 24: 24 (1851). S. magnifica Sleum. in Notizbl. 12: 182 (1934). S. pilosa A. C. Smith in Contrib. U. S. Nat. Herb. 28: 355 (1932). Sophoclesia robusta Rusby, Descr. S. Am. Pls. 77 (1920) — Sphyrospermum cordifolium Sphvidsniesaak buxifolium Poepp. & Endl. Nov. Gen. 1: 4, t. 8 (1835). S. cordifolium Benth. Pl. Hartw. 222 (1846). Thibaudia axillaris Rusby in A. C. Smith in Contrib. U. S. Nat. Herb. 28: 417 (1932). . erenulata Remy in Ann. Sci. Nat. (ser. 3) 8: 234 (1847). densiflora (Herzog) A. C. Smith in Bull. Torr. Bot. Club, 60: 114 (1933). . macrocalyx Remy in Ann. Sci. Nat. (ser. 3) 8: 235 (1847). oblongifolia Remy in Ann. Sci. Nat. (ser. 3) 8: 233 (1847) — Cavendishia ubesce Vacciniopeis t tetrauers Rusby, Descr. S. Am. Pls. 77 (1920) — Disterigma alaternoides var. parvifolium. Vaccinium didymanthum Dun. in DC. Prodr. 7: 575 (1839). V. empetrifolium HBK. Nov. Gen. & Spec. 3: 268, t. 248 (1819). V. epacridifolium Benth. Pl. Hartw. 221 (1846). V. floribundum HBK. Nov. Gen. & Spec. 3: 226, t. 251 (1819). SHAH 154 V. floribundum var. ramosissimum (Dun.) Sleum. in Notizbl. 13: 131 (1936). V. floribundum var. Tatei (Rusby) Sleum. in Notizbl. 13: 132 (1936). V. marginatum Dun. in DC. Prodr. 7: 570 (1839). V. penaeoides HBK. Nov. Gen. & Spec.. 3: ta (1819). V. polystachyum Benth. Pl. Hartw. 140 (1844 V. oo sore Sleum. in Notizbl. 12: 140 : 1934) — = Eleutherostemon race- THEOPHRASTACEAE Clavija boliviensis Mez in Engler, Pflanzenr. IV, 236a: 26, hig C. Cardenasii Rusby in Mem. N. Y. Bot. Gard. 7: 318 C. formosa Remy in Ann. Sci. Nat. (ser. 3) 8: 227 a C. Hassleri Mez in Bull. Herb. Boiss. (ser. 2) 3: 238 (1903). C. spathulata R. & P. Syst. 285 (1798). C. tarapotana Spruce ex Rusby in Bull. N. Y. Bot. Gard. 4: 406 (1907). C. tarapotana Rusby in Bull. N. Y. Bot. Gard. 8: 111 (1912). MYRSINACEAE Ardisia acuminata Willd. Sp. Pl. 1(2): 1062 (1798). Conomorpha peruviana A. DC. in Ann. Sci. Nat. (ser. 2) 16: 92 (1841). Cybianthus glaucus Rusby in Mem. N. Y. Bot. Gard. 7: 318 (1927). C. lanceolatus Pax in Fedde, Repert. Spec. Nov. 7: 112 (1909). C. psychotriaefolius (Rusby) Rusby ex Mez in Engler, Pflanzenr. IV, 236: 227 ). Geissanthus Bangii Rusby in Mem. Torr. Bot. Club, 4(3): 217 (1895). G. bolivianus Britton in Bull. Torr. Bot. Club, 20: 140 (1893). G. Haenkeanus Mez in Engler, Pflanzenr. IV, 236: 236 (1902). G. multiflorus Mez in Engler, Pflanzenr. IV, 236: 239 (1902). G. Pentlandii Mez in Engler, Pflanzenr. IV, 236: 236 (1902). yrsine viridis Rusby in Mem. Torr. Bot. Club, 6: 74 (1896) —= Rapanea ferruginea. Parathesis macrophylla (Benth.) Britton in Mem. Torr. Bot. Club, 4(3): 217 1895). Peckia purpurea Rusby in Bull. N. Y. Bot. Gard. 4: 405 (1907). Rapanea denticulata ae in Phytologia, 1: 72 (1934). dependens (R. & P.) Mez in Engler, Pflanzenr. IV, 236: 377 (1902). ferruginea (R. & P.) Mez in Urb. Symb. Ant. 2: 429 (1901). ianensis Aubl. Pl. Guian. Fr. 1: 121 (1775). latifolia (R & P.) Mez in Engler, Pflanzenr. IV, 236: 387 (1902). lucida Herzog in Meded. Rijks Herbar. 27: 25 (1915) Mandonii Mez in Engler, Pflanzenr. IV, 236: 378 (1902). pseudocrenata Mez in Engler, Pflanzenr. IV, 236: 393 (1902). sessiliflora Mez in Fedde, Repert. Spec. Nov. 3: 103 (1906). Sprucei Rusby in Bull. N. Y. Bot. Gard. 8: 112 (1912). venosa (A. DC.) Mez in Engler, Pflanzenr. IV, 236: 386 (1902). Stylogyne Poeppigii Mez in Engler, Pflanzenr. IV, 236: 274 (1902). Weigeltia Buchtienii Pax in Fedde, Repert. Spec. Nov. 7: 112 (1909). PARA AAPA AA PRIMULACEAE Anagallis arvensis L. Sp. Pl. 148 (1753). Centunculus pumilus var. ovalis (R. & P.) O. Ktze. Rev. Gen. 3(2): 193 (1898) C. pumilus var. pentandrus (R. Br.) O. Ktze. Rev. Gen. 3(2): 193 (1898). Samolus Valerandi var. floribundus BSP. Prelim. Cat. N. Y. Pls. 34 (1888). 155 PLUMBAGINACEAE Plumbago coerulea HBK. Nov. Gen. & Spec. 2: 220 (1817). P. scandens L. Sp. Pl. (ed. 2) 1: 215 (1762). SAPOTACEAE Bumelia obtusifolia subsp. excelsa (A. DC.) Cronquist in Journ. Arnold Arb. 26: 458 45). B. sartorum Mart. Herb. Fl. Bras. 233 (1837). Chrysophyllum gonocarpum (Mart. & Eichl.) Engler in Engler, Bot. Jahrb. 12: 523 (1892). iicioides Rusby in Bull. N. Y. Bot. Gard. 4: 406 (1907) = C. gonocarpum. - marginatum (Hook. & Arn.) Radlk. in Act. Congr. Bot. Anvers, 170 (1887). - maytenoides Mart. in Flora, 22, I Beibl.: 1 (1839). C. ovale Rusby in Mem. N. Y. Bot. Gard. 7: 320 (1927). Lucuma reticulata Remy in Ann. Sci. Nat. (ser. 3) 8: 228 (1847). Mimusops surinamensis Miq. in Mart. Fl. Bras. 7: 44 (1863). Pouteria bilocularis (Winkl.) Baehni in Candollea, 9: 229 (1942). P. boliviana (Rusby) Baehni in Candollea, 9: 419 (1942). P. catoclantha (Eichl.) Baehni in Candollea, 9: 232 (1942). P. nemorosa Baehni in Candollea, 9: 348 (1942). P. polycarpa (Rusby) Baehni in Candollea, 9: 425 (1942). P. ramiflora (Mart.) Radlk. in Sitzb. Akad. Wiss. Miinch. 12: 333 (1882). Sideroxylon Bangii Rusby in Bull. N. Y. Bot. Gard. 4: 407 (1907). 88 EBENACEAE Diospyros boliviana Rusby in Mem. N. Y. Bot. Gard. 7: 320 (1927). D. tetramera Rusby in Mem. Torr. Bot. Club, 6: 75 (1896). D. velutina Hiern in Trans. Cambr. Phil. Soc. 12: 200 (1873). STYRACACEAE Styrax Buchtienii Sleum. in Fedde, Repert. Spec. Nov. a 125 (1936). S. Pearcei Perkins in Engler, Pflanzenr. IV, 241: 46 (190 S. Pearcei var. bolivianus Perkins in Engler, Pflanzenr. 1V, 241: 46 (1907). S. Pentlandianus Remy in Ann. Sci. Nat. (ser. 3) 8: 229 (1847). S. subheterotrichus Herzog in Meded. Rijks Herbar. 40: 31 (1921). S. tarapotensis Perkins in Engler, Bot. Jahrb. 31: 479 (1902). SYMPLOCACEAE Symplocos boliviana Brand in Engler, Pflanzenr. IV, 242: 89 (1901). castanea Brand in Fedde, Repert. Spec. Nov. 14: 324 (1916). colorata Brand in Engler, Pflanzenr. IV, 242: 74 (1901). flavescens Rusby in Bull. N. Y. Bot. Gard. 4: 407 (1907). hiemalis Lingelsh. in Fedde, Repert. Spec. Nov. 8: 3 (1910). mapiriensis Brand in Engler, Pflanzenr. IV, 242: 74 (1901). Matthewsii A. DC. Prodr. 8: 250 (1844). parvifolia var. subcuneata Herzog in Meded. Rijks Herbar. 40: 7 (1921). theaeformis (L. f.) Giirke in Engl. & Prantl, Nat. Pflanzenfam. 4(1): 172 NAAN INAD tn CQLEACEAE Jasminum grandiflorum L. Sp. Pl. (ed. 2) 1: 9 (1762). Mayepea implicata Rusby in Bull. N. Y. Bot. Gard. 4: 314 (1907). Menodora chlorargantha (Remy) Steyerm. in Ann. Missouri Bot. Gard. 19: 156 (1932). 156 M. helianthemoides Humb. & Bonpl. Pl. Aequin, 2: 98, t. 110 (1809). M. integrifolia (Cham. & Schlechtd.) Steud. Nomencl. (ed. 2) 2: 124 (1841). M. integrifolia var. trifida (Cham. & Schlechtd.) Steyerm. in Ann. Missouri Bot. Gard. 19: 131 (1932). M. pulchella Markgraf in Notizbl. 8: 219 (1922). LOGANIACEAE Buddleia americana L. Sp. Pl. 112 (1753). B. anchoénsis O. Ktze. Rev. Gen. 3(2): 200 (1898). B. andina Britton ex Rusby in Mem. Torr. Bot. Club, 4(3): 222 (1895). B. aromatica Remy in Ann. Sci. Nat. (ser. 3) 8: 227 (1847). B. Bangii Krinzl. in Ann. Naturh. Hofmus. Wien, 26: 395 (19 B. boliviana Pax in Fedde, Repert. Spec. Nov. 5: 227 dOny 2 a “Cordis bifur- B. brasiliensis Jacq. ex Spreng. Syst. 1: 430 (1825). B. canescens Rusby in Mem. Torr. Bot. Club, 6: 78 (1896). B. cochabambensis Rusby, Descr. S. Am. Pls. 81 (1920). B. coriacea Remy in Ann. Sci. Nat. (ser. 3) 8: 226 (1847). B. coroicensis Rusby in Bull. N. Y. Bot. Gard. 4: 412 (1907). B. hypoleuca Kranzl. in Engler, Bot. Jahrb. 50, Beibl. 111: 39 (1913). B. ignea Kranzl. in Engler, Bot. Jahrb. 40: 309 (1908). B. inconspicua Kranzl. in Engler, Bot. Jahrb. 40: 310 2. B. ledifolia Krianzl. in Engler, Bot. Jahrb. 50, Beibl. 111: 42 (1918). B. microcephala Rusby in Bull. N. Y. Bot. Gard. 8: 113 (1912). B. misera Kranzl. in Engler, Bot. Jahrb. 40: 308 (1908). B. monocephala Kranzl. in Engler, Bot. Jahrb. 40: 309 (1908). B. montana Britton ex Rusby in Bull. Torr. Bot. Club, 25: 544 (1898). B. oblongifolia Rusby in Bull. N. Y. Bot. Gard. 8: 112 (1912). B. rhododendroides Krinzl. in Ann. Naturh. Hofmus. Wien, 26: 395 (1912). B. soratae Kranzl. in Engler, Bot. Jahrb. 50, Beibl. 111: 47 (1913). B. tiraquiensis O. Ktze. Rev. Gen. 3(2): 201 (1898). B. tucumanensis Griseb. in Goett. Abh. 19: 213 (1874). B. Urbaniana Krianzl. in Engler, Bot. Jahrb. 40: 310 (1908). B. verbascifolia HBK. Nov. Gen. & Spec. 2: 351 (1818). Desfontainea parvifolia D. Don in Edinb. New Philos. Journ. 1831: 275. D. spinosa R. & P. Fl. Peruv. 2: 47, t. 186 (1799). Spigelia elongata Rusby in Bull. Torr. Bot. Club, 25: 542 (1898). S. Herzogiana Kranzl. in Fedde, Repert. Spec. Nov. 13: 117 (1914). S. sessilifolia Rusby in Bull. Torr. Bot. Club, 25: 543 (1898). Strychnos brachiata R. & P. Fl. Peruv. 2: 30, t. 157 (1799). S. darienensis Seem. Bot. Voy. Herald, 166 (1854). . Mitscherlichii Rich. Schomb. Reisen, 2: 451 (1848). parvifolia DC. Prodr. 9: 16 (1845). Peckii Robinson in Proc. Amer. Acad. 49: 504 (1913). rondeletioides Spruce ex Benth. in Journ. Linn. Soc. 1: 104 (1856). nn ta th GENTIANACEAE mmeorrsiery ae (R. & P.) Gilg in Engl. & Prantl, Nat. Planzenfam. 4(2): C. Whitei stir in Mem. N. Y. Bot. Gard. 7: 322 (1927). Curtia tenuifolia (Aubl.) Knobl. in Bot. Centralbl. 60: 323 (1894). Deianira chiquitana Herzog in Fedde, Repert. Spec. Nov. 7: 65 (1909). Gentiana albido-coerulea Gilg in Engler, Bot. evn: 22: 323 (1896). G. alticola R. C. Foster in Rhodora, 56: 103 (1954). G. anthosphaera Gilg in Fedde, Repert. Spec. Nov. 2: 46 (1906). AMNAARNARAR AAMPANAAAARAAAAAAA ANAAANAARAHAR ARALRAHAA Aan A 157 armerioides Griseb. ex Gilg in Engler, Bot. Jahrb. 22: 306 (1896), nomen udum. nudum. Bangiana Gilg in Engler, Bot. Jahrb. 22: 335 (1896). Bangii Gilg in Engler, Bot. Jahrb. 22: 324 (May 22, 1896). Bangii Rusby in Mem. Torr. Bot. Club, 6: 79 (Nov. 17, 1896) = G. albido- coerulea. bellatula Gilg in Engler, Bot. Jahrb. 50, Beibl. 111: 49 (1913). Benedictae Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 80 (1916). Bockii Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 34 (1916). boliviana Pax in Fedde, Repert. Spec. Nov. 7: 243 (1909). Bridgesii Gilg in Engler, Bot. Jahrb. 22: 316 (1896). Briquetiana Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 32 (1916). Buchtienii Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 66 (1916). cerastioides var. emarginata Gilg in Engler, Bot. Jahrb. 22: 328 (1896) = G. paleana. chrysantha Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 81 (1916). ambensis Rusby in Mem. Torr. Bot. Club, 6: 79 (1896) = G. Kuntzei. comarapana Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 82 (1916). dasythamna Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 63 (1916). Dielsiana Gilg in Engler, Bot. Jahrb. 22: 316 (1896). dolichantha Gilg in Torreya, 5: 109 (1905). erythrochrysea Gilg in Fedde, Repert. Spec. Nov. 2: 38 (1906). Fiebrigii Gilg in Fedde, Repert. Spec. Nov. 2: 45 (1906). florida Griseb. in Goett. Abh. 24: 236 (1879). gageoides Gilg in Engler, Bot. Jahrb. 22: 320 (1896). gynophora Gilg in Engler, Bot. Jahrb. 22: 305 (1896). Hauthalii Gilg ex Perkins in Engler, Bot. Jahrb. 49: 212 (19138) = G. Dielsiana. Herzogii Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 57 (1916). hygrophiloides Gilg in Fedde, Repert. Spec. Nov. 2: 40 (1906) = G. Kuntzei. inaequicalyx Gilg in Engler, Bot. Jahrb. 22: 324 (1896). incurva Hook. in Bot. Misc. 2: 228 (1831). Krauseana Gilg in Fedde, Repert. Spec. Nov. 2: 45 (1906). Kuntzei Gilg in Engler, Bot. Jahrb. 22: 326 (1896). Kusnezowii Gilg in Engler, Bot. Jahrb. 22: 825 (1896). lancifolia Gilg in Engler, Bot. Jahrb. 22: 326 (1896) = G. alticola. larecajensis Gilg in Engler, Bot. Jahrb. 54. Beibl. 118: 31 (1916). lilacino-flavescens Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 38 (1916). limoselloides HBK. Nov. Gen. & Spec. 3: 167, t. 220, fig. 1 (1819). lithophila Gilg in Engler, Bot. Jahrb, 54, Beibl. 118: 39 (1916). longipes Rusby in Mem. N. Y. Bot. Gard. 7: 321 (1927). lythroides Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 24 (1916). macroclada Gilg in Fedde, Repert. Spec. Nov. 2: 47 (1906). macrorrhiza Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 40 (1916). Mandonii Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 37 (1916) = G. neo- mandonii. Mandonii Rusby in Mem. Torr. Bot. Club, 6: 80 (1896) = G. inaequicalyx. myriantha Gilg in Fedde, Repert. Spec. Nov. 2: 43 (1906). narcissoides Gilg in Engler, Bot. J ahrb. 54, Beibl. 118: 65 (1916). neomandonii R. C. Foster in Rhodora, 56: 103 (1954). odontosepala Gilg in Fedde, Repert. Spec. Nov. 2: 48 (1906). orobanchoides Gilg in Engler, Bot. Jahrb. 22: 333 (1896). paleana Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 47 (1916). pallide-lilacina Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 58 (1916). Pilgeriana Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 42 (1916). pinifolia R. & P. ex G. Don, Gen. Syst. 4: 182 (1837). 158 praticola en in Engler, Bot. Jahrb. 54, Beibl. 118: 46 (1916) = G. sancti- matthae primulifolia Griseb. Gen. & Spec. Gent. 221 (1839). primuloides Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 31 (1916). prostrata Jacq. Collect. 2: 66, t. 17 (1788 prostrata var. podocarpa (Griseb.) Rams in Act. Hort. Petrop. 15: 373 904). pseudocrassula Gilg in Fedde, Repert. Spec. Nov. 2: 44 (1906) . pseudolycopodium Gilg in Fedde, Repert. Spec. Nov. 2: 38 (1906). punicea Wedd. Chlor. And. 2: 70 (1859 purpureiflora Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 65 (1916). rupicola HBK. Nov. Gen. & Spec. 3: 167, t. 220, fig. 2 (1819) sancti-matthaei R. C. Foster in Rhodora, 56: 103 (1954). sandiensis Gilg in Fedde, Repert. Spec. Nov. 2: 36 (1906). scopulorum Wedd. Chlor. And. 2: 67 (1859). sini HBK. Nov. Gen. & Spec. 3: 173, t. 225 (1819). a Rusby in Mem. Torr. Bot. Club, 6: 81 (1896) = G. gynophora. icnioides Gilg in Engler, Bot. Jahrb. 22: 319 (1896). soratensis Gilg in Engler, Bot. Jahrb. 22: 332 (1896). spectabilis Rusby in Mem. Torr. Bot. Club, 6: 80 (1896) = G. Dielsiana. stenocephala Gilg in Engler, Bot. Jahrb. 22: 331 (1896). striaticalyx Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 56 (1916). Tatei Rusby in Phytologia, 1: 72 (1934). thiosphaera Gilg in Fedde, Repert. Spec. Nov. 2: 46 (1906). . totorensis Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 84 (1916). . tradescantiifolia Britton ex Rusby in Bull. Torr. Bot. Club, 25: 545 (1898). virgata Rusby in Mem. Torr. Bot. Club, 6: 81 (1896) — G. Kusnezowii. alenia bifida Rusby & Allen in Ann. Missouri Bot. Gard. 20: 201 (1933). Dombeyana (Griseb.) Wedd. Chlor. And. 2: 76 (1859). - gracilis (HBK.) G. Don. Gen. Hist. 4: 177 (1887). Herzogii Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 118 (1916). Hieronymi Gilg in Fedde, Repert. Spec. Nov. 2: 52 (1906). pendulifiora Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 116 (1916). pusilla Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 107 (1916). robusta Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 110 (1916). Rusbyi Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 117 (1916). silenoides Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 113 (1916). valerianoides Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 106 (1916). vincetoxicoides Gilg in Engler, Bot. Jahrb. 54, Beibl. 118: 108 (1916). Leiphaimos ben dp (Jacq.) Gilg in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 104 (1895). Limnanthemum Humboldtianum (HBK.) Griseb. Gen. & Spec. Gent. 347 (1839). L. verrucosum R. E. Fries in Ark. Bot. 6(11): 25 (1906). Lisianthus calycogonus R. & P. Fl. Peruv. 2: 14, t. 126 (1799). L. ovalis R. & P. Fl. Peruv. 2: 18 (1799). Macrocarpaea Bangiana Gilg in Engler, Bot. Jahrb. 22: 335 (1896). M. cochabambensis Ch. Gilg in Notizbl. 13: 381 (1936). Microcala quadrangularis (Lam.) Griseb. in DC. Prodr. 9: 63 (1845). Rusbyanthus cinchonifolius (Britton) Gilg in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 95 (1895). Schultesia Pohliana Progel in Mart. Fl. Bras. 6(1): 205 (1865). Symbolanthus Brittonianus Gilg in Engler, Bot. Jahrb. 22: 342 (1896). S. Rusbyanus Gilg in Engler, Bot. Jahrb. 22: 344 (1896). Tachia sp. DE} RE AHA RARER S? CoRgpog agpoR Caeue e ARAA A 159 Tetragonanthus Whitei Rusby in Mem. N. Y. Bot. Gard. 7: 321 (1927) = Halenia vincetoxicoides. Zygostigma australe (Cham. & Schlechtd.) Griseb. Gen. & Spec. Gent. 150 (1839). APOCYNACEAE Aspidosperma australe Muell. Arg. in Mart. Fl. Bras. 6(1): 58 (1860). brevifolium Rusby in Bull. N. Y. Bot. Gard. 8: 113 (1912) = A. cylindro- carpon. cylindrocarpon Muell. Arg. in Mart. Fl. Bras. 6(1): 54 (1860). macrocarpon Mart. Nov. Gen. 1: 59 (1826). Marcgravianum Woodson in Ann. Missouri Bot. Gard. 38: 170 (1951). Quebracho-blanco Schlechtd. in Bot. Zeit. 19: 137 (1861). ramifiorum Muell. Arg. in Mart. Fl. Bras. 6(1): 55 (1860). rauwolfioides Markgraf in Notizbl. 12: 300 (19385) = A. rigidum. rigidum Rusby in Mem. N. Y. Bot. Gard. 7: 323 (1927). Steinbachii Markgraf in Notizbl. 9: 1158 (1927). ep > ee > Dipladenia Buchtienii Rusby, Descr. S. Am. Pls. 87 (1920) —= Mandevilla cuspidata. D. glabra Rusby, Descr. S. Am, Pls. 88 (1920) — Mandevilla pulchra. mollis Rusby in Bull. N. Y. Bot. Gard. 8: 114 (1912) = Mandevilla cuspi- piladenia Rusby, Descr. S. Am. Pls. 87 (1920) — Mandevilla cuspidata. rotundifolia Rusby in Mem. N. Y. Bot. Gard. 7: 326 (1927) = Mandevilla cuspidata . tetradenia Rusby, Descr. S. Am. Pls. 88 (1920) — Mandevilla cuspidata. Echites altescandens H. Winkl. in Fedde, Repert. Spec. Nov. 7: 243 (1909) = Mandevilla antennacea. . Bangii Rusby in Bull. N. Y. Bot. Gard. 4: 409 (1907) = Prestonia acuti- folia. E. boliviana Britton ex ag in Mem. Torr. Bot. Club, 4(3): 219 (1895) = Mandevilla antennace E. mapirensis H. Winkl. - in Fedde, Repert. Spec. Nov. 7: 113 (1909) = Mesechites acuminata ? E. rigida Rusby in Mem. N. Y. Bot. Gard. 7: 325 (1927) —= Mesechites trifida. E. — forma puberula Markgraf in Notizbl. 9: 80 (1924) = Mesechites acuminata. Hevetenuas amblybasis Blake in Journ. Wash. Acad. Sci. roe 292 (1924). F. glabrescens Muell. Arg. in Mart. Fl. Bras. 6(1): 102 (1860). F. mollis Rusby in Mem. Torr. Bot. Club, 4(3): 218 (1895). F. mollis var. foliosa (Rusby) Woodson in Ann. Missouri Bot. Gard. 22: 188 (1935). F.. obscura Rusby in Mem, Torr. Bot. Club, 4(3): 219 (1895) — F. thyrsoidea var. glabriuscula. F. pubescens A. DC. Prodr. 8: 436 (1844). F. Sellowii Muell. Arg. in Mart. Fl. Bras. 6(1): 101 (1860). F. thyrsoidea var. glabriuscula (A. DC.) Woodson in Ann. Missouri Bot. Gard. 22: 201 (1935). Himatanthus obovatus (Muell. Arg.) Woodson in Ann, Missouri Bot. Gard. 25: 201 (1938). H. Sucuuba (Spruce) Woodson in Ann. Missouri Bot. Gard. 25: 198 (1937). Landolphia boliviensis Markgraf in Notizbl. 9: 1041 (1926). Laseguea Mandonii Britton ex Rusby in Mem. Torr. Bot. Club, 4(3): 220 by 160 (1895) = Mandevilla Pentlandiana. Macropharynx fistulosa Rusby in Mem. N. Y. Bot. Gard. 7: 329, fig. 6 (1927) = M. spectabilis. M. spectabilis (Stadelm.) Woodson in Ann. Missouri Bot. Gard. 18: 552 (1931). Macrosiphonia longiflora (Desf.) Muell. Arg. in Mart. Fl. Bras. 6(1): 140, t. 43 (1860). scarce antennacea (A. DC.) K. Schum. in Engl. & Prantl, Nat. Pflanzen- 4(2): 171 (1895). 7] Poe Rusby in Mem. Torr. Bot. Club, 6: 76 (1896) — M. Bridgesii. M. boliviensis (Hook. f.) Woodson in Ann. Missouri Bot. Gard. 20: 716 (1933). rachyloba (Muell. Arg.) K. Schum. in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 171 (1895). bracteosa (Rusby) Woodson in Ann. Missouri Bot. Gard. 20: 742 prea eee (Muell. Arg.) Woodson in Ann. Missouri Bot. Gard. 19: (1932) cereola Mlaclacn in Ann. Missouri Bot. Gard. 20: 712 (1933). collium Woodson in Ann. Missouri Bot. Gard. 23: 380 (1936). cuspidata (Rusby) Woodson in Ann. Missouri Bot. Gard. 20: 730 (1933). erecta (Vell.) Woodson in Ann. Missouri Bot. Gard. 19: 62 (1932). . fragilis Woodson in Ann. Missouri Bot. Gard. 19: 59 (1932). hirsuta (A. Rich.) K. Schum. in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 171 (1895). laxa (R. & P.) Woodson in Ann. Missouri Bot. Gard. 19: 68 (1932). Mandonii Britton ex Rusby in Bull. Torr. Bot. Club, 25: 496 (1898) = BEESS 5S FE S& & M. Bridgesii oblongifolia (Woodson) Woodson in Ann. Missouri Bot. Gard. 20: 708 1933). Pentlandiana (A. DC.) Woodson in Ann. Missouri Bot. Gard. 19: 63 (1932). pulchra Woodson in Ann. Missouri Bot. Gard. 23: 376 (1936). Rusbyi Britton ex Rusby in Bull. N. Y. Bot. Gard. 4: 409 (1907) = M. hir- suta. . rutila Woodson in Ann. Missouri Bot. Gard. 19: 385 (1982). M. subcordata Rusby in Bull. N. Y. Bot. Gard. 4: 315 (1907). M. superba Herzog in Fedde, Repert. Spec. Nov. 7: 65 (1909). M. tenuicarpa Rusby in Bull. N. Y. Bot. Gard. 8: 114 (1912) = M. antennacea. Mesechites acuminata (R. & P.) Muell. Arg. in Linnaea, 30: 446 (1860). M. Mansoana (A. DC.) Woodson in Ann. Missouri Bot. Gard. 20: 636 (1936). M. trifida ( Jacq.) Muell. Arg. in Mart. Fl. Bras. 6(1): 151 (1860). Odontadenia boliviana Rusby in Mem. Torr. Bot. Club, 6: 76 (1896). O. laxiflora (Rusby) Woodson in Ann. Missouri Bot. Gard. 19: 386 (1932). Peltastes giganteus Woodson in Ann. Misouri Bot. Gard. 19: 873 (1982). Plumeria bracteata A. DC. Prodr. 8: 394 (1844). P. tarapotensis K. Schum. ex Markgraf in Notizbl. 11: 339 (1982). P. velutina var. boliviana O. Ktze. Rev. Gen. 3(2): 198 (1898). Prestonia acutifolia (Benth.) K. Schum. in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 188 (1895) cephalantha Rusby in Mem. N. Y. Bot. Gard. 7: 330 (1927). cornutisepala Rusby in Mem. N. Y. Bot. Gard. 7: 829 (1927). cyaniphylla (Rusby) Woodson in Ann. Missouri Bot. Gard. 23: 284 (1936). Muelleri Rusby in Mem. Torr. Bot. Club, 4(3): 217 (1895) — P. Riedelii. Phenax Woodson in Ann. Missouri Bot. Gard. 23: 314 (1936). Riedelii (Muell. Arg.) Markgraf in Fedde, Repert. Spec. Nov. 20: 26 (1924). robusta Rusby, Descr. S. Am. Pls. 91 (1920). sericocalyx Malme in Bih. Sv. Vet. Akad. Handl. 24(10): 29, t. 3 (1899). SS 5 & = iw ef ee 161 P. tomentosa R. Br. in Mem. Wern. Soc. 1: 70 (1811). Rauwolfia boliviana Markgraf in Fedde, Repert. Spec. Nov. 20: 122 (1924). R. ternifolia HBK. Nov. Gen. & Spec. 3: 232 (1819). Secondatia densiflora A. DC. Prodr. 8: 445 (1844). Stemmadenia obovata var. mollis (Benth.) Woodson in Ann. Missouri Bot. Gard. 15: 358 (1928). Tabernaemontana Buchtienii H. Winkl. in Fedde, Repert. Spec. Nov. 7: 244 (1909). T. laeta Mart. in Flora, 20, II Beibl.: 98 (1887). T. macrosiphon Herzog in Fedde, Repert. Spec. Nov. 7: 66 (1909). T. mapirensis Rusby in Bull. N. Y. Bot. Gard. 8: 115 (1912). T. myriantha Britton ex Rusby, Descr. S. Am. Pls. 84 (1920). T. undulata Mey. Primit. Fl. Esseq. 135 (1818). T. unguiculata Rusby in Mem. N. Y. Bot. Gard. 7: 324 (1927). Vallesia glabra (Cav.) Link, Enum. Hort. Berol. 1: 207 (1821). ASCLEPIADACEAE Amblystigma hypoleucum Benth. in Hook. Ic. Pl. 12: 76, t. 1188 (1876). A. pedunculare Benth. in Hook. Ic. Pl. 12: 77, sub t. 1188 (1876) — Sarcostem- ma Gilliesii. Amphistelma Pearcei Rusby in Bull. N. Y. Bot. Gard. 4: 411 (1907). Aphanostelma parviflorum (Malme) Maime in Ark. Bot. 25A(7): 11 (1932). Araujia grandiflora (Mart. & Zucc.) Morong in Ann. N. Y. Acad. Sci. 7: 161 (1893). A. plumosa Schltr. in Oesterr. Bot. Zeitschr. 45: 449 (1895). Asclepias barjoniaefolia Fourn. in Ann. Sci. Nat. (ser. 6) 14: 372 (1882). A. boliviensis Fourn. in Ann. Sci. Nat. (ser. 6) 14: 381 (1882). . brasiliensis (Fourn.) Schltr. in Meded. Rijks Herbar. 29: 12 (1916). . Bridgesii Fourn. in Ann. Sci. Nat. (ser. 6) 14: 383 (1882). campestris Decne. in DC. Prodr. 8: 566 (1844). cochabambensis Rusby in Mem. Torr. Bot. Club, 4(3): 221 (1895). curassavica L. Sp. Pl. 215 (1753). . Fiebrigii Schltr. in Engler, Bot. Jahrb. 37: 608 (1906). . Kuntzei Schitr. in Oesterr. Bot. Zeitschr. 49: 450 (1895). marginata Decne, in DC. Prodr. 8: 566 (1844). nervosa Decne. in DC. Prodr. 8: 568 (1844). . Pilgeriana Schltr. in Engler, Bot. Jahrb. 37: 608 (1906). ramosa Fourn. in Ann. Sci. Nat. (ser. 6) 14: 373 (1882). _ Weddellii Fourn. in Ann. Sci. Nat. (ser. 6) 14: 381 (1882). Blepharodon bolivianum Malme in Ark. Bot. 25A (7): 17 (1982). B. mucronatum (Schlechtd.) Decne. in DC. Prodr. 8: 603 (1844). B. philibertioides Schltr. in Fedde, Repert. Spec. Nov. 13: 440 (1914). B. Rahmeri Phil. in Anal. Mus. Nac. Chile, Bot. 8: 52, t. 2 (1891). Corollonema boliviense Schltr. in Fedde, Repert. Spec. Nov. 13: 441 ( 1914). Cynanchum Rusbyi Malme in Ark. Bot. 25A (7): 9 (1932). Dactylostelma boliviense Schltr. in Oesterr. Bot. Zeitschr. 45: 452 (1895). >>> > > > > Pbpb PP D. bonariensis Decne. in DC. Prodr. 8: 577 ( 1844). D. Fiebrigii Malme in Ark. Bot. 28A (5): 22 (1936). D. lanceolata Decne. in DC. Prodr. 8: 576 (1844). D. Mandonii Rusby, Descr. S. Am. Pls. 97 (1920). D. montana Schitr. in Fedde, Repert. Spec. Nov. 13: 489 (1914) = D. Schlech- teri. D. racemosa Britton ex Rusby in Bull. Torr. Bot. Club, 25: 498 (1898). D. Schlechteri Macbr. in Field Mus. Publ. Bot. 11: 84 (1931). 162 D. subalpina Schltr. in Fedde, Repert. Spec. Nov. 18: 489 (1914). Fischeria boliviana Blake in Journ. Wash. Acad. Sci. 14: 292 (1924). F. Martiana Decne. in DC. Prodr. 8: 601 (1844). Funastrum fragile Rusby in Mem. N. Y. Bot. Gard. 7: 332 (1927) = Sarcostem- lausum. F. lanceolatum Rusby in Mem. N. Y. Bot. Gard. 7: 332 (1927) — Sarcostem- ma clausum. Gonolobus andinus Malme in Ark. Bot. 25A a }: 9 (1932). G. Asplundii Malme in Ark. Bot. 25A (7): 20 (1932). G. attenuatus Rusby, Descr. S. Am. Pls. (1920). G. boliviensis (Schltr.) T. Meyer in Lilloa, 23: 57 (1950). G. Brittonii Rusby in Bull. Torr. Bot. Club, 25: 500 (1898). G. ellipticus Rusby in Mem. Torr. Bot. Club, 4(3): 222 (1895). G. Fiebrigii Schltr. in Engler, Bot. Jahrb. 37: 625 (1906). G. floribundus Malme in Ark. Bot. 25A (7): 19 (1982). G. hirsutissimus Schltr. in Engler, Bot. Jahrb. 37: 625 (1906). G. leucodermis Rusby, Descr. S. Am. Pls. 102 (1920). Gothofreda andina Rusby in Mem. Torr. Bot. Club, 4(3): 220 (1895) = Oxypetalum sp. ? G. apoloénsis Rusby in Bull. N. Y. Bot. Gard. 8: 113 (1912) = Oxypetalum ? sp. ? G. rete’ Rusby in Mem. N. Y. Bot. Gard. 7: 334 (1927) = Oxypetalum sp. G. macrapoes Rusby in Mem. N. Y. Bot. Gard. 7: 334 (1927) —= Oxypetalum G. Pearson Rusby in Mem, N. Y. Bot. Gard. 7: 335 (1927) — Oxypetalum ahs ts andinum Rusby, Descr. S. Am. a 92 (1920). H. Williamsii Rusby, Descr. 8S. Am. Pls. 92 (1920). Jobinia chlorantha (K. Schum.) Malme in Ark. Bot. 25A (7): 3 (1932). Macroditassa tassadioides (Schltr.) Malme in Ark. Bot. 28A (5): 6 (1936). Marsdenia Hilariana Fourn. in Mart. Fl. Bras. 6(4): 322 (1885). Melinia campanulata Schltr. in Engler, Bot. Jahrb. 37: 615 (1906). M. discolor Schltr. in Engler, Bot. Jahrb. 37: 615 (1906). Metastelma ditassoides Schltr. in Fedde, Repert. Spec. Nov. 18: 488 (1914). M. ditassoides Schltr. in Meded. Rijks Herbar. 29: 12 (1916) —= M. Schlechteri. Fiebrigii Schltr. in Engler, Bot. Jahrb. 37: 609 (1906). Herzogii Schltr. in Fedde, Repert. Spec. Nov. 13: 488 (1914). Mathewsii Rusby in Bull. Torr. Bot. Club, 25: 497 (1898). myrianthum Schltr. in Oesterr. Bot. Zeitschr. 45: 451 (1895). parviflorum (Sw.) R. Br. in Mem. Wern. Soc. 1: 52 (1811), combination implied but not actually made. retinaculatum Schltr. in Engler, Bot. Jahrb. 37: 610 (1906). Schlechteri Macbr. in Field Mus. Publ. Bot. 11: 34 (1931). Mitostigma boliviense Schltr. in Engler, Bot. Jahrb. 37: 601 (1906). M. Fiebrigii Schltr. in Engler, Bot. Jahrb. 37: 602 (1906). M. grandiflorum Schltr. in Engler, Bot. Jahrb. 37: 603 (1906). M. Herzogii Schltr. in Fedde, Repert. Spec. Nov. 13: 441 (1914). M. niveum Griseb. in Goett. Abh. 24: 266 (1879). M. subniveum Malme in Ark. Bot. 3(1): 9 (1904). Morrenia brachystephana Griseb, in Goett. Abh. 19: 205 (1874). M. *grandiflora Malme in Ark. Bot. 8(1): 21 (1909) [but called “subsp. nov.”]. M. Herzogii Schltr. in Fedde, Repert. Spec. Nov. 13: 440 (1914). M. odorata (Hook. & Arn.) Lindl. in Bot. Reg. 24: Mise. 71 (1838). Nephradenia pendula Rusby in Mem, N. Y. Bot. Gard. 7: 336 (1927). Oxypetalum albiflorum Schltr. in Engler, Bot. Jahrb. 37: 621 (1906). 163 attenuatum (Rusby) Malme in Ark. Bot. 25A (14): 15 (1933). Balansae Malme in Kgl. Sv. Vet. Akad. Handl. ra 51 (1900). . boliviense Schltr. in Engler, Bot. Jahrb. 37: 622 (1906). brachystemma Malme in Ark. Bot. 3(8): 2, t. 1, fig. 3 (1904). Schlechteri Malme in Ark. Bot. O5A (7): 15 ‘(1982 ). Phaeostemma grandifolium Rusby, Descr. S. Am. Pls. 101 (1920). Philibertia hypoleuca Schitr. in Oesterr, Bot. Zeitschr. 45: 450 (1895) = Sarcostemma campanulat P. picta Schltr. in Engler, Det: sdaheb. 37: 606 (1906) — Sarcostemma, prob- ably a hybrid Pseudibatia Herzogii Schltr. in Fedde, Repert. Spec. Nov. 13: 443 (1914). Roulinia Mannii Rusby in Mem. N. Y. Bot. Gard. 7: 333 (1927). Sarcostemma campanulatum Lindl, in Bot. Reg. 32: t. 86 (1846). clausum (Jacq.) Roem. & Schult. Syst. Veg. 6: 114 (1820). . Gilliesii (Hook. & Arn.) Decne. in DC. Prodr. 8: 542 (1844). . gracile Decne. in DC. Prodr. 8: 589 (1844). . lysimachioides (Wedd.) R. Holm in Ann. Missouri Bot. Gard. 37: 547 (1950). 20999 Dnmnmn solanoides (HBK.) Decne. in DC. Prodr. 8: 541 (1844). Vaileae (Rusby) R. Holm in Ann. Missouri Bot. Gard. 37: 543 (1950). Schistogyne boliviensis Schltr. in Fedde, Repert. Spec. Nov. 18: 442 (1914). S. Fiebrigii Malme in Ark. Bot. 25A (14): 14 (1938). S. Mandonii Malme in Ark. Bot. 25A (14): 12 (19383). S. oxypetaloides Schltr. in Fedde, Repert. Spec. Nov. 18: 442 (1914) = Oxy- petalum attenuatum. S. pentaseta Rusby, Descr. S. Am. Pls. 98 (1920). Steleostemma pulchellum Schltr. in Engler, Bot. Jahrb. 37: 604 (1906). Stelmatocodon Fiebrigii Schltr. in Engler, Bot. Jahrb. 37: 618 (1906). Tassadia Hutchisoniana Rusby in Mem. N. Y. Bot. Gard. 7: 331 (1927). T. rhombifolia Rusby in Mem. N. Y. Bot. Gard. 7: 331 (1927). T. Rusbyi Macbr. in Field Mus. Publ. Bot. 11: 34 (1981). Sprucei Rusby in Bull. Torr. Bot. Club, 25: 498 (1898) = T. Rusbyi. Tweedia brunonis Hook. & Arn. in Hook. Journ. Bot. 1: 292 (1934). Vailia mucronata Rusby in Bull. Torr. Bot. Club, 25: 542 (1898). Vincetoxicum cuspidatum Rusby in Mem. N. Y. Bot. Gard. 7: 333 (1927). V. umbellatum Rusby in Mem. Torr. Bot. Club, 6: 78 (1896) = Cynanchum Rusbyi. V. unguiculatum (R. & P.) Britton ex Rusby in Bull. Torr. Bot. Club, 25: 499 (1898). Rta CONVOLVULACEAE Bonamia boliviana O’Donell in Lilloa, 23: 458 (1950). Calonyction aculeatum (L.) House in Bull. Torr. Bot. Club, 31: 590 (1904). Convolvulus Blanchetii forma albiflorus O. Ktze. Rev. Gen. 3(2): 212 (1898). Blanchetii forma coeruleus O. Ktze. Rev. Gen. 3(2): 212 (1898). bogotensis forma albiflorus O. Ktze. Rev. Gen. 3(2): 212 (1898). bogotensis forma lilacinus O. Ktze. Rev. Gen. 3(2): 212 (1898). bonariensis Cav. Ic. 5: 54, t. 480, fig. 2 (1799). crenatifolius R. & P. Fl. Peruv. 2: 10, t. 118 (1799). crenatifolius var. argentinicus Hall. f. in Jahrb. Hamb. Wiss Anst. 16, 3 Beih.: 35 (1899). npertrg var. peruvianus Hall. f. in Jahrb. Hamb. Wiss. Anst. 16, 3 Beih.: 34 (18 caine Deis in Lam. Encycl. 3: 546 (1789). Ottonis (Chois.) Meissn. in Mart. Fl. Bras. 7: 311 (1869). aa a aaaaan 164 C. pauciflorus var. chilensis O. Ktze. Rev. Gen. 3(2): 214 (1898). ). fragrans Rusby in Mem. Torr. Bot. Club, 6: 85 (1896) — C. odorata. grandiflora HBK. Nov. Gen. & Spec. 3: 123, t. 213 (1819). insquamata Yuncker in Amer. Journ. Bot. 10: 12, t. 5, fig. 30a-d (1923). odorata R. & P. Fl. Peruv. 1: 69, t. 105, fig. a (1798) odorata var. Holwayana Yuncker in Amer. Journ. Bot. 9: 564, t. 4, fig. 21f (1922). partita Choisy in Mém. Soc. Phys. Hist. Nat. Genéve, 9: 284, t. 5 (1841). Rojasii Hunziker in Darwiniana, 7: 323 (1947). Dichondra argentea Willd. Enum. Hort. Berol. 297 (1809). D. evolvulacea var. villosa (Parodi) O. Ktze. Rev. Gen. 3(2): 216 (1898). D. repens Forst. Char. Gen. 40, t. 20 (1776). D. repens var. microcalyx Hall. f. in Engler, Bot. Jahrb. 18: 84 (1893). Evolvulus alsinoides var. debilis (HBK.) v. Ooststr. in Meded. Bot. Mus. Utrecht, 14: 33 (1934). argyreus Choisy in Mém. Soc. Phys. Hist. Nat. Genéve, 8: 75 (1837). boliviensis v. Ooststr. in Meded. Bot. Mus Utrecht, 14: 83 (1934). columbianus var. incanus Hall. f. in Meded. Rijks Herbar. 46: 13 (1922) = E. tenuis subsp. longifolius. corumbaénsis Hoehne in Anex. Mem. Inst. Butantan Bot. 1(6): 38, t. 1 (1922). elegans Moric. Pl. Nouv. Amér. 53, t. 36 (1838). frankenioides Moric Pl. Nouv. Amér. 49, t. 33 (1838). frankenioides var. subglaber v. Ooststr. in Meded. Bot. Mus. Utrecht, 14: 149 (1934). glaber Spreng. Syst. Veg. 1: 862 (1825). glomeratus subsp. eu-glomeratus forma echioides (Moric.) v. Ooststr. in Meded. Bot. Mus. Utrecht, 14: 228 (19384). linoides Moric. Pl. Nouv. Amér. 139, t. 83 (1844). nummularius (L.) L. Sp. Pl. (ed. 2) 1: 391 (1762). pterocaulon var. floccosus Meissn. in Mart. Fl. Bras. 7: 333 (1869). sericeus Sw. Prodr. 55 (1788). sericeus var. holosericeus (HBK.) v. Ooststr. in Meded. Bot. Mus. Utrecht, 14: 180 (1934). E. tenuis subsp. longifolius (Choisy) v. Ooststr. in Meded. Bot. Mus. Utrecht, 14: 61 (1934). Ipomoea alba L. Sp. Pl. 161 (1753). I. Batatas Poir. in Lam. Encycl. 6: 14 (1804). . bona-nox L. Sp. Pl. (ed. 2) 1: 228 (1762). carnea Jacq. Enum. Pl. Carib. 13 (1760). chiliantha Hall. f. in Bull. Herb. Boiss. (ser. 1) 7, App. 1: 50 (1899). . coccinea L. Sp. Pl. (ed. 2) 1: 228 (1762). densibracteata O’Donell in Lilloa, 23: 438 (1950). edulis (Choisy) Niederl. in Bol. Mens. Mus. Prod. Argent. 3(29): 190 (1890). fastigiata (Roxb.) Sweet, Hort. Brit. (ed. 1) 288 (1827). filipedunculata Rusby in Bull. Torr. Bot. Club, 26: 150 (1899). fistulosa Mart. ex Choisy in DC. Prodr. 9: 349 (1845). floribunda Moric. Pl. Nouv. Amér. 46, t. 31 (1838). ederacea Jacq. Collect. 1: 124 (1786). indivisa (Vell.) Hall. f. in Meded. Rijks Herbar. 46: 20 (1922). magnifolia Rusby in Mem. Torr. Bot. Club, 6: 84 (1896). neurocephala Hall. f. in Jahrb. Hamb. Wiss. Anst. 16, 3 Beih.: 40 (1899). Nil (L.) Roth, Catalect. Bot. 1: 36 (1797). IAA |egan by bl bt . . Se SS See: bb bl . ° oe a ee ee Sea 165 I. opulifolia Rusby in Bull. Torr. Bot. Club, 169 A. Sellowii (Brig.) Moldenke in Revist. Sudam. Bot. 4: 15 (1937). A. virgata (R. & P.) A. Juss. in Ann. Mus. Paris, 7: 73 (1806), combination implied but not actually made. Baillonia amabilis Bocq. in Adansonia, 2: 252, t. 7 (1862). B. amabilis var. longifolia O. Ktze. Rev. Gen. 3(2): 250 (1898). Bouchea boliviana (O. Ktze.) Moldenke in Phytologia, 1: 18 (1933). B. fluminensis (Vell.) Moldenke in Fedde, Repert. Spec. Nov. 49: 117 (1940). B. incisa Rusby in Bull. N. Y. Bot. Gard. 4: 482 (1907) = B. Rusbyi B. pseudogervao forma pilosa Herzog in Meded. Rijks Herbar. 29: 46 (1916) . bolivia B. Rusbyi Moldenke in Torreya, 34: 8 (1934). Callicarpa acuminata HBK. Nov. Gen. & Spec. 2: 252 (1818). C. minutiflora Rusby in Mem. N. Y. Bot. Gard. 7: 339 (1927). Castelia cuneato-obovata Cav. in Anal. Cienc. Nat. 3: 134 (180 Citharexylum andinum Moldenke in Fedde, Repert. Spec. Nov. a. eae (1934). ilicifolium HBK. Nov. Gen. & Spec. 2: 256 (1818). laurifolium Hayek in Engler, Bot. Jahrb. 42: 170 (1908). megacanthum Rusby in Bull. N. Y. Bot. Gard. 8: 115 (1912). mendocinum R. A. Phil. Sert. Mend. Alt. 193 (1870) Poeppigii var. margaritaceum Poepp. & Moldenke in | Fedde, Repert. Spec. Nov. 37: 2383 (1934). . punctatum Greenm. in Field Columb. Mus. Bot. Ser. 2: 189 (1907). Clerodendron bolivianum Britton ex Rusby in Bull. Torr. Bot. Club, 27: 82 (1900) = Aegiphila multiflora. C. fragrans var. pleniflorum Schauer in DC. Prodr. 11: 666 (1847). Duranta Benthamii Briq. in Bull. Herb. ime (ser. 1) 4: 343 (1896). Mandonii Moldenke in Lilloa, 5: 390 (19 Pearcei Rusby in Bull. N. Y. Bot. Ga oy 4: 433 (1907). recurvistachys Rusby in es hyrengioet 1: 73 (1934). repens L. Sp. Pl. 637 (175 rupestris Hayek in Engler, "ait. Jahrb, 42: 171 (1908). D. serratifolia (Griseb.) O. Ktze. Rev. Gen. 3(2): 251 (1898). Ghinia Cardenasii Moldenke in Bull. Torr. Bot. Club, 68: 504 (1941). Junellia aretioides (R. E. Fries) Moldenke in Lilloa, 5: 393 (1940). J. bisuleata (Hayek) Moldenke, Known Geogr. Distrib. Verb. & Avicenn. 77 1942). a aeeae Sooss J. minima (Meyen) Moldenke in Lilloa, 5: 398 (1940). Lantana achyranthifolia Desf. Cat. Hort. Paris (ed. 3) 392 (1829). L. aristata var. Cabrerae Moldenke in Lilloa, 5: 404 (1940). L. aristata var. pluripedunculata (O. Ktze.) Moldenke in Phytologia, 1: 170 (1935). L. Balansae Briq. in Ann. Conserv. & Jard. Bot. Genéve, 7-8: 300 (1904). L. Balansae var. peduncularis Briq. in Bull. Herb. Boiss. (ser. 2) 4: 1063 (1904 L. beseliy yes Hayek in Engler, Bot. Jahrb. 42: 167 (1908). L. Camara L. Sp. Pl. 627 (1753). L. Camara var. mista (L.) L. H. Bailey, Cycl. Am. Hort. 884 (1900). L. Camara var. nivea (Vent.) L. H. Bailey, Cycl. Am. Hort. 883 (1900). L. canescens HBK. Nov. Gen. & Spec. 2: 259 (1818). L. Chamissonis (D. Dietr.) Benth. & Hook. f. Gen. Pl. 2: 1142 (1876), com- bination implied but not actually made. L. cujabensis Schauer in DC. Prodr. 11: 599 (1847). L. Fiebrigii Hayek in Engler, Bot. Jahrb. 42: 169 (1908). L. foetida Rusby in Bull. N. Y. Bot. Gard. 4: 431 (1907). L. fucata Lindl. in Bot. Reg. 10: t. 798 (1824). L. hyptoides Rusby in Bull. N. Y. Bot. Gard. 4: 431 (1907). 170 micrantha Briq. in Ann. Conserv. & Jard. Bot. Genéve, 7-8: 299 (1904). micrantha var. armata Moldenke in Phytologia, 2: 468 (1948). montevidensis (Spreng.) Brig. in Ann. Conserv. & Jard. Bot. Genéve, 7-8: 301 (1904). ovata Hayek in Fedde, Repert. Spec. Nov. 2: 163 (1906). procurrens Schauer in DC, Prodr. 11: 596 (1847). rugulosa HBK. Nov. Gen. & Spec. 2: 261 (1818). Rusbyana Moldenke in Lilloa, 8: 422 (1942). tenuifolia Rusby in Phytologia, 1: 74 (1984) — L. cujabensis. trifolia L. Sp. Pl. 626 (1753 Lippia affinis Schauer in DC. Prone: 11::676: (1847). L. alba (L.) N. E. Br. in Britt. & Wils. Sci. Surv. Porto Rico, 6: 141 (1925). L. aristata Schauer in DC. Prodr. 11: 581 (1847). L. asperrima Cham. in Linnaea, 7: 215 (1832). L. boliviana Rusby in Mem. Torr. Bot. Club, 4(3): 248 (1895). L. chacensis Moldenke in Phytologia, 2: 414 ; L. dumetorum Herzog in Meded. Rijks Herbar. 29: 45 (1916), L. fimbriata Rusby in Mem. Torr. Bot. Club, 4(3): 244 (1895) = Lantana achyranthifolia. L. hirsuta var. ee (Cham.) O. Ktze. Rev. Gen. 3(2): 252 (1898). } rrr rr ee L. imbricata O. Ktze. Rev. Gen. 3(2): 252 (1898 L. lasiocalycina hie in Linnaea, 7: 231 (1832). L. lasiocalyx Herzog in Meded. Rijks Herbar, 29: 44 (1916). L. laxibracteata Herzog in Meded. Rijks Herbar. 29: 45 (1916). L. lupulina Cham. in Linnaea, 7: 222 (1832). L. origanoides HBK. Nov. Gen. & Spec. 2: 267 (1818). L. pendula Rusby in Bull. N. Y. Bot. Gard. : 116 (1912), L. salviaefolia Cham. in Linnaea, 7: 227 (1832). L, suffruticosa (Griseb.) O. Ktze. Rev. Gen. Ng 253 (1898). L. turnerifolia Cham. in wa : 217 (1832 L. venosa Rusby in Mem. N, Y. Gerd. 7: 338 (1927). Parodianthus ilicifolius Na — in Darwiniana, 5: 39 (1941). Petrea fragrantissima Rusby in Mem. N. Y. Bot. Gard. 7: 388 (1927). P. maynensis Huber in Bol. Mus. Goéldi, 4: 602 (1906). Phyla betulaefolia (HBK.) Greene in Pittonia, 4: 48 (1899). P. caespitosa (Rusby) Moldenke in Rev. Sudam. Bot. 5: 2 (1987). P. nodiflora (L) Greene in Pittonia, 4: 46 (1899). ig i dacs var. reptans (HBK.) Moldenke, Alph. List. Inval. Names Verb. 31 (1942). - P. nodiflora var. rosea (D. Don) Moldenke in Phytologia, 2: 22 (1941). Priva boliviana Moldenke in Phytologia, 3: 172 (1949). P. lappulacea (L.) Pers. Synops. Pl. 2: 189 (1806). Recordia boliviana Moldenke in Phytologia, 1: 99 (1934). Stachytarphaeta canescens HBK. Nov. Gen. & girs 2: 281 (1818). S. cajanensis (L. C. Rich.) Vahl, Enum. 1: 208 (1804). S. elatior Schrad. ex Schult. Mant. 1: 172 (1822). Timotocia Mansoi (Schauer) Moldenke in Fedde, Repert. Spec. Nov. 39: 142 (1936) Verbena Bangiana Moldenke in Phytologia, 3: 638 (1949). V. bonariensis L. Sp. Pl. 20 (1753 V. brasiliensis Vell. Fl. Flum. 17 (1825). V. cochabambensis Moldenke in Castanea, 10: 45 (1945). V. dissecta Willd. ex Spreng. Syst. Veg. 2: 750 (1825). V. gracilescens (Cham.) Herter in Rev. Sudam. Bot. 4: 186 (1937). V. hispida R. & P. FI. Peruv. 1: 22, t. 34, fig. a (1798). V. laciniata (L.) Brig. in Ann. Conserv. & Jard. Bot. Genéve, 7-8: 296 (1904). 171 . littoralis HBK. Nov. Gen. & Spec. 2: 276, t. 187 (1818). microphylla HBK. Nov. Gen. & Spec. 2: 272, t, 133 (1818). officinalis L. Sp. Pl. 20 (1753). parvula Hayek in Engler, Bot. Jahrb. 42: 162 (1908) peruviana (L.) Britton in Ann. N, Y. Ac ad. Sci. 1: 197 (1893). rigida Spreng. Syst.: Cur. Post. 230 (1827). seriphioides Gill. & Hook. in Bot. Misc. 1: 164 (1829). tenera Spreng. Syst. Veg. 2: 750 (1825). tenuisecta Briq. in Ann. Conserv. & Jard. Bot. Genéve, 7-8: 294 (1904). Vitex cymosa Bert. ex Spreng. Syst. Veg. 2: 757 (1925 V. pseudolea Rusby in Mem. N. Y. Bot. Gard. 7: 341 (1927). V. triflora Vahl, Symb. 2: 49 (1798). LABIATAE Bystropogon andinus var. hypoleucus Briq. in Bull. Herb. Boiss. (ser. 1) 4: 799 (1896) = Minthostachys ovata. B. minutus Briq. in Bull. Herb. Boiss. (ser. 1) 4: 803 (1896) = Satureja parvifolia B. uniflorus Rusby ex Brig. in Bull. Herb. Boiss. (ser. 1) 4: 802 (1896) = Satureja axillaris Ceratominthe Kiminenee Brig. in Bull. Herb. Boiss. (ser. 1) 4: 877 (1896) = Satureja odora. ee. Kuntzeanum O. Ktze. Rev. Gen. 3(2): 259 (1898) = Satureja bolivian Eriope Putas Benth. Lab. Gen. 144 (1833). Hedeoma adscendens Rusby in Phytologia, 1: fe (1934). H. Mandoniana Wedd. Chlor. And. 2: 148 (1860). Hyptis arborea Benth. in DC. Prodr. 12: 182 (1848). H. brevipes Poit. in Ann. Mus. Paris, 7: 465 (1806). H. canaminensis Rusby in Mem. N. Y. Bot. Gard. 7: 342 (1927) = H. mu- tabilis var. canescens. carpinifolia Benth. Lab. Gen. 115 (1833). compacta Rusby in Mem. N. Y. Bot. Gard. 7: 343 (1927). conferta var. angustifolia Benth. in DC. Prodr. 12: 112 (1848), nomen nudum. dumetorum Morong in Ann. N. Y. Acad. Sci. 7: 200 (1893). duplicato-dentata Pohl ex Benth. Lab. Gen. — ‘ses 3). eriocephala Benth. in DC. Prodr. 12: 124 (184 parece St. Hil. ex Benth. Lab. Gen. 92 seen [as H. goyavensis]. gymnodonta Briq. in Bull. Herb. Boiss. (ser. 1) 4: 795 (1896) = H. du- iddddd<<<< metorum hirsuta HBK. Nov. Gen. & Spec. 2: 318, t. 161 (1818). taioernele Briq. in Bull. Herb. Boiss. (ser. 1) 4: 792 (1896) = H. Lorent- zian ae Herzog in Fedde, Repert. Spec. Nov. 7: 66 (1909) = H. micro- phyla. Kuntzeana Brig. in Bull. Herb. Boiss. (ser. 1) 4: 787 (1896). lantanaefolia Poit. in Ann. Mus. Paris, 7: 468, t. 29, fig. 1 (1806). lappacea Benth. Lab. Gen. 103 (1833). lappulacea Mart. ex Benth. Lab. Gen. 104 (1833). longifrons Briq. in Bull. Herb. Boiss. (ser. 1) 4: 793 (1896) = H. velutina. Lorentziana O. Hoffm. in Linnaea, 43: 137 (1880). lutescens Pohl ex Benth. Lab. Gen. 109 pcre microphylla Pohl ex Benth. Lab. Gen. 82 (183 mutabilis (A. Rich.) Briq. in Bull. Herb. Boiss. a 1) 4: 788 (1896). mutabilis var. canescens (HBK.) Brig. in Bull. Herb. Boiss. (ser. 1) 4: 788 (1896). ee Rm Ame . Po eo Ee a “ = =~] bo mutabilis var. spicata (Poit.) Brig. in Bull. Herb. Boiss. (ser. 1) 4: 788 (1896). obtusiflora Pres] ex Benth. Lab. Gen. 107 (1833). odorata Benth. Lab. Gen. 81 (1833). pectinata Poit. in Ann. Mus. Paris, 7: 474, t. 30 (1806). recurvata Poit. in Ann. Mus. Paris, 7: 467, t. 28, fig. 2 (1806). recurvata var. megacephala Benth. in DC. Prodr. 12: 90 (1848). rugosa var. canescens (Briq.) Epl. in Fedde, Repert. Spec. Nov. Beih. 85: 275 (1937). sinuata Pohl ex Benth. Lab. Gen. 103 (1833). spicigera Lam. Encycl. 3: 185 (1789). suaveolens Poit. in Ann. Mus. Paris, 7: 472, t. 29, fig. 2 (1806). Tafallae Benth. Lab. Gen. 132 (1833). pao aiagl Briq. in Bull. Herb. Boiss. (ser. 1) 4: 790 (1896) = H. lutes- ie aap po at fat ft tect ; Subioa var. Mikanii Benth. in DC. Prodr. 12: 124 (1848). . uncinata Benth. Lab. Gen. 80 (1833). H. velascana Briq. in Bull. Herb. Boiss. (ser. 1) 4: 798 (1896) = H. virgata. H. velutina Pohl ex Benth. Lab. Gen. 90 (1833). H. virgata Benth. Lab. Gen. 77 (1833). Lepechinia aurifera (Rusby) Epl. in Fedde, Repert. Spec. Nov. Beih. 85: 25 (1935). L. bella Epl. in Fedde, Repert. Spec. Nov. Beih. 85: 21 (1935). L. confusa (Briq.) Epl. in Fedde, Repert. Spec. Nov. Beih. 85: 25 (1935). L. floribunda (Benth.) Epl. in Fedde, Repert. Spec. Nov. Beih. 85: 22 (1935). L. graveolens (Regel) Epl. in Fedde, Repert. Spec. Nov. Beih. 85: 25 (1935). L. heteromorpha (Briq.) Epl. in Fedde, Repert. Spec. Nov. Beih. 85: 26 L. inflata (Briq.) Epl. in Fedde, Repert. Spec. Nov. Beih. 85: 24 (1935). L. lancifolia (Rusby) Epl. in Fedde, Repert. Spec. Nov. Beih. 85: 23 (1935). L. Meyenii (Walp.) Epl. in Fedde, Repert. Spec. Nov. Beih. 85: 20 (1935). Marrubium vulgare L. Sp. Pl. 583 (1753). Marsypianthes Chamaedrys (Vahl) O. Ktze. Rev. Gen. 2: 524 (1891). Mentha aquatica L. Sp. Pl. 576 (1753). Mesosphaerum arboreum var. bracteosum Rusby in Mem. Torr. Bot. Club, 6: 107 (1896) = Hyptis arborea. M. —— var. albiflorum O. Ktze. Rev. Gen. 3(2): 260 (1898) — Hyptis Sp. M. —— var. subviolaceum O. Ktze. Rev. Gen. 3(2): 259 (1898) —= Hyptis M. praniforum Rusby in Bull. N. Y. Bot. Gard. 8: 116 (1912) = Hyptis Tafal M. pallidum ‘Rusby in Bull. N. Y. Bot. Gard. 4: 483 (1907) = Hyptis obtusi- flor M. anishdsinied Britton ex Rusby in Mem. Torr. Bot. Club, 4(3): 246 (1895) = Hyptis mutabilis var. canescens. Minthostachys acutifolia Epl. in Fedde, Repert. Spec. Nov. Beih. 85: 166 (1936). M. andina (Britton) Epl. in Fedde, Repert. Spec. Nov. Beih. 85: 166 (1936)). M. diffusa Epl. in Fedde, Repert. Spec. Nov. Beih. 85: 167 (1936). M. Mandoniana (Briq.) Epl. in Fedde, Repert. Spec. Nov. Beih. 85: 167 (1936). M. ovata (Briq.) Epl. in Fedde, Repert. Spec. Nov. Beih. 85: 167 (1936). M. setosa (Briq.) Epl. in Fedde, Repert. Spec. Nov. Beih. 85: 166 (1936). Ocimum micranthum Willd. Enum. Hort. Berol. 630 (1809). Perilomia ocimoides HBK. Nov. Gen. & Spec. 2: 828 (1818). Rosmarinus officinalis L. Sp. Pl. 23. (1753). Salvia amplifrons Briq. in Bull. Herb. Boiss. (ser. 1) 4: 863 (1896). ECECECEC TA Annnnn ae ERC REE np mw Manny tn 173 atrocyanea Epl. in Fedde, Repert. Spec. Nov. Beih. 85: 98 (1936). australis Epl. in Rev. Sudam. Bot. 4: 48 (1937 avicularis Brig. in Bull. Herb. Boiss. (ser. 1) 4: 866 (1896) = S. Haenkei. Bangii Rusby in Mem. Torr. Bot. Club, 4(3): 246 (1895). Bridgesii Britton ex Rusby in Bull. Torr. Bot. Club, 27: 124 (1900) = S. Haenkei. cardiophylla Benth. Lab. Gen. 721 (1835). ease antha Briq. in Bull. Herb. Boiss. (ser. 1) 4: 861 (1896) = S. Orbig- porereree Epl. in Fedde, Repert. Spec Nov. Beih. 85: 57 (1935). preteen Rusby in Mem. Torr. Bot. Club, 6: 108 (1896) = S. Orbig- Dombei Epl. in Rev. Sudam. Bot. 4: 47 (1937). erythradenia Briq. in Bull. Herb. Boiss. (ser. 1) 4: 853 (1896) = S. Bangii. Gilliesii Benth. Lab. Gen. 265 (1833). grewiaefolia S. Moore in Journ. Bot. 42: 109 (1904). Haenkei Benth. Lab. Gen. 283 (18338). aie Brig. in Bull. Herb. Boiss. (ser. 1) 4: 859 (1896) = S. Orbig- ated R. & P. Fl. Peruv. 1: 23, t. 40 (1798). occidentalis Sw. Prodr. 14 (1788). Orbignaei Benth. in DC. Prodr. 12: 338 (1848). oxyphora Briq. in Bull. Herb. Boiss. (ser. 1) 4: 864 (1896). personata Epl. in Fedde, Repert. Spec. Nov. Beih. 85: 73 (1935). platystoma Epl. in Fedde, Repert. Spec. Nov. Beih. 85: 81 (1935). praeclara Epl. in Fedde, Repert. Spec. Nov. Beih. 85: 121 (1936). pseudoavicularis Briq. in Bull. Herb. Boiss. (ser. 1) 4: 867 (1896) = S. Haenkei. retinervia Briq. in Bull. Herb. Boiss. (ser. 1) 4: 857 (1896). rhombifolia R. & P. Fl. Peruv. 1: 26, t. 36 (1798). Rusbyi Britton ex Rusby in Mem. Torr. Bot. Club, 4(3): 247 (1895). Briq. in Bull. Herb. Boiss. (ser. 1) 4: 850 (1896) = S. tiraquensis. sophrona Briq. in Bull. Herb. Boiss. (ser. 1) 4: 854 (1896). stachydifolia Benth. in DC. Prodr. 12: 311 (1848). tiliaefolia Vahl, Symb. 3: 7 (1794). . tiraquensis Bria. in Bull. Herb. Boiss. (ser. 1) 4: 851 (1896). es axillaris (Rusby) Epl. in Ann. Missouri Bot. Gard. 14: 82 (1927). boliviana (Benth.) Briq. in Engl. & Prantl, Nat. Pflanzenfam. 4(3a): 300 (1896). boliviana var. tarijensis (Wedd.) Ep]. in Ann. Missouri Bot. Gard. 14: 81 (1927). Kuntzeana Briq. in Bull. Herb. Boiss. (ser. 1) 4: 874 (1896) = S. boliviana. odora (Griseb.) Epl. in Fedde, Repert. Spec. Nov. Beih. 85: 158 (1936). pallida Epl. in Ann. Missouri Bot. Gard. 14: 65 (1927). parvifolia (Phil.) Epl. in Fedde, Repert. Spec. Nov. Beih. 85: 159 (1936). simulans Epl. in Ann. Missouri Bot. Gard. 14: 81 (1927). Scutellaria breviflora Benth. in Bot. Reg. 18: sub t. 1493, no. 7 (1832). Ss. Ss. S. racemosa Pers. Synops. Pl. 2: 136 (1806). uliginosa St. Hil. ex Benth. Lab. Gen. 427 (183 4). Sphacele cochabambana Briq. in Bull. Herb. Boiss. (ser. 1) 4: 807 (1896) = Lepechinia aurifera. Kuntzeana Briq. in Bull, Herb. Boiss. (ser. 1) 4: 805 (1896) = Lepechinia aurifera. Brig. in Ann. Conserv. & Jard. Bot. Genéve, 2: 176 (1898) —= Lepechinia inflata. 174 Stachys boliviana Briq. in Bull. Herb. Boiss. (ser. 1) 4: 871 (1896) = S. petio- S. Mandoniana Briq. in Ann. Conserv. & Jard. Bot. Genéve, 2: 118 (1898). S. petiolosa Briq. in Bull. Herb. Boiss. (ser. 1) 4: 868 (1896). S. pusilla (Wedd.) Brig. in Ann. Conserv. & Jard. Bot. Genéve, 2: 120 (1898). Teucrium vesicarium Mill. Gard. Dict. (ed. 8) no. 17 (1768). SOLANACEAE Acnistus oblongifolius Rusby in ses Torr. Bot. Club, 4(3): 2382 (1895) = Dunalia breviflora var. glabra Bassovia anceps (R. & P.) aay in n Bull, Torr. Bot. Club, 26: 197 (1899). B. Fendleri Rusby in Bull. Torr. Bot. — 26: 197 (1899). B. minutiflora Rusby in Mem. N. Y. Bot. Gard. 7: 3438 (1927). B. platyneura H. Winkl. in Fedde, —. Spec. Nov. 7: 248 (1909). B. solanacea (Miers) Benth. & Hook. f. Gen. Pl. 2: 891 (1876), combination implied but not actually made. Brachistus hebephyllus Miers in Ann. & Mag. Nat. Hist. (ser. 2) 3: 266 (1 : B. lasiophyllus (Humb. & Bonpl.) Rusby in Bull. Torr. Bot. Club, 26: 198 (1899). B. virgatus H. Winkl. in Fedde, Repert. Spec. Nov. 7: 245 (1909) — Lycianthes actinocalyx. Browallia viscosa HBK. Nov. Gen. & Spec. 2: 273 (1818). Brugmansia arborea (L.) Steud. Nomencl. (ed. 2) 1: 230 (1840). B. bicolor (R. & P.) Pers. Synops. Pl. 1: 216 (1805). B. sanguinea (R. & P.) D. Don in Sweet, Brit. Fl. Gard. (ser 2) 3: t. 272 (1835). B. suaveolens (Humb. & Bonpl.) Bercht. & Presl, Rostl. 1. Solanac. 45 (1823). Brunfelsia Mire Monachino in Phytologia, 4:342 (1953). B. latifolia (Pohl) Benth. in DC. Prodr. 10: 199 (1846). Cacabus parviflorus Rusby in Mem. Torr. Bot. Club, 4(3): 233 (1895). Capsicum baccatum L. Mant. 47 (1767). C. frutescens L. Sp. Pl. 189 (17538). C. glomeruliflorum (Sendt.) O. Ktze. Rev. Gen. 3(2): 218 (1898). C. grandiflorum O. Ktze. Rev. Gen. 3(2): 218 (1898) — Dunalia fasciculata. C. punctatum (R. & P.) O. Ktze. Rev. Gen. 3(2): 219 (1898). C. solanaceum (L’Hérit.) O. Ktze. Rev. Gen. 3(2): 219 (1898). Cestrum albotomentosum U. Damm. ex Francey in Candollea, 6: 152 (1936). C. atroxanthum O. Ktze. Rev. Gen. 3(2): 219 (1898). C. Baenitzii Lingelsh. in Fedde, Repert. Spec. Nov. 7: 248 (1909). Baenitzii var. angustifolium Francey in aa 6: 312 (1936). bolivianum Francey in Candollea, 6: 230 ih Buchtienii Francey in Candollea, 6: 300 (19 bor calycinum Willd. ex Roem. & Schult. Syst. shin 4: 808 (1819). ellipticum Francey in Candollea, 6: 216 (1986). floribundum Britton in Mem. Torr. Bot. Saco 6: 92 (1896) = C. reflexum. foliosum Francey in Candollea, 6: 260 (193 Herzogii E. Damm. in Meded. Rijks Herbar. voi 28 (1916). impressum Rusby in Bull. N. Y. Bot. Gard. 4: 425 (1907) —= C. ~— Mandonii Rusby in Bull. N. Y. Bot. Gard. 4: 425 (1907) = C. Parqui Mathewsii Dunal in DC. Prodr. 13(1): 637 (1852). papyraceum Rusby, Descr. S. Am. Pls. 120 (1920). Parqui L’Hérit. Stirp. Nov. 4: 73, t. 36 (1788) . Parqui var. macrocalyx Francey in Candollea, 7(preprint): 45 (1936). . Parqui var. oranense Scolnik in Revist. Argent. Agron. 21: 30 (1954). PAAgaaanagananas 175 Parqui var. tomentistipes O. Ktze. Rev. Gen. 3(2): 219 (1898). plicatum Francey in Candollea, 7: 52 (1936). racemosum var. bolivianum Francey in Candollea, 6: 274 (1936). reflexum Sendt. in Mart. Fl. Bras. 10: 218 (1846). reflexum var. densiflorum Francey in Candollea, 6: 267 (1936). rigidum Rusby in Mem. Torr. Bot. Club, 4(3): 234 (1895). sparsiflorum Britton ex Rusby in Bull. Torr. Bot. Club, 27: 22 (1900). strigillatum R. & P. Fl. Peruv. 2: 29, t. 156 (1799) [as C. strigilatum]. strigillatum var. calycinum (HBK.)O. Ktze. Rev. Gen. 3(2): 220 (1898). tenuissimum Francey in Candollea, 6: 248 (1936). Chamaesaracha boliviensis U. Damm. in Engler, Bot. Jahrb. 49: 215 (1913) = Solanum chamaesarachidium Cyphomandra acuminata Rusby in Bull. Torr. Bot. Club, 26: 196 (1899). arborea H. Winkl. in Fedde, Repert. Spec. Nov. 7: 246 (1909). artocarpophylla H. Winkl. in Fedde, Repert. Spec. Nov. 7: 246 (1909). benensis Britton ex Rusby in Bull. Torr. Bot. Club, 26: 196 (1899). crassifolia (Ort.) O. Ktze. Rev. Gen. 3(2): 220 (1898). Fraxinella Sendt. in Mart. Fl. Bras. 10: 122 (1846). subcordata Rusby in Bull. N. Y. Bot. Gard. 8: 119 (1912). uniloba Rusby in Mem. Torr. Bot. Club, 6: 90 (1896). yungasensis Rusby in Bull. Torr. Bot. Club, 26: 195 (1899). angaaaagaaa aaaaaaaa Dunalia brachyantha Miers in Hook. Lond. Journ. Bot. 7: 339 (1848). D. breviflora var. glabrata (Sendt.) Sleum. in Lilloa, 23: 181 (1950). D. fasciculata (Miers) Sleum. in Lilloa, 23: 135 (1950). D. Hauthalii U. Damm. in Engler, Bot. Jahrb. 50, Beibl. 111: 55 (1918). D. Herzogii U. Damm. in Meded. Rijks Herbar. 29: 25 (19 16) = D. senticosa. D. lycioides Miers in Hook. Lond. Journ. Bot. 4: 334 (1845). D. Pflanzii U. Damm. in Engler, Bot. Jahrb. 50, Beibl. 111: 57 (1918) = D. lycioides. D. senticosa Miers in Hook. Lond. Journ. Bot. 4: 340 (1848). D. Steinbachii Sleum. in Lilloa, 23: 136 (1950). Fabiana densa Remy in Ann. Sci. Nat. (ser. 3) 8: 225 (1847). Fregirardia Dunaliana Wedd. Chlor. And. 2: 100 (1859). Grabowskia boerhaaviaefolia var. obtusifolia O. Ktze. Rev. Gen. 8(2): 221 G. schizocalyx U. Damm. in Meded. Rijks Herbar. 29: 22 (1916). Iochroma lyciifolium U. Damm. in Engler, Bot. Jahrb. 37: 637 (1906). Jaborosa floccosa U. Damm. in Engler, Bot. Jahrb. 37: 641 (1906). J. leiocalyx U. Damm. in Meded. Rijks Herbar. 29: 27 (1916). Juanulloa Hookeriana Miers in Ann. & Mag. Nat. Hist. (ser. 2) 4: 189 (1849). J. membranacea Rusby in Mem. Torr. Bot. Club, 4(3): 233 (1895). J. mexicana (Schlechtd.) Miers in Ann. & Mag. Nat. Hist. (ser. 2) 4: 188 849). J. pedunculata Rusby in Mem. Torr. Bot. Club, 4(3): 234 (1895). Lonchestigma crispum (Miers) Dunal in DC. Prodr. 13(1): 477 (1852). Lycianthes actinocalyx (H. Winkl.) Bitter in Abh. Mateo Ver. Bremen, 24: (1920 L. apiculata Bitter in Abh. Naturw. Ver. Bremen, 24: 452 (1920). L. Buchtienii Bitter in Abh. Naturw. Ver. Bremen, 24: 337 (1920) = L. acti- nocalyx. L. candicans (Dun.) Hass]. in Ann. Conserv. & Jard. Bot. Genéve, 20: 181 (1917). L. coccinea (Rusby) Rusby in Bull. Torr. Bot. Club, 53: 210 (1926). L. fasciculata (Rusby) Bitter in Abh. Naturw. Ver. Bremen, 24: 334 (1920). 176 Fendleri (Rusby) Rusby in Bull. Torr. Bot. Club, 53: 210 (1926). heterodoxa Bitter in Abh. Naturw. Ver. Bremen, 24: 333 (1920). hispida (Rusby) Rusby in Bull. Torr. Bot. Club, 53: 210 (1926). hylophila Bitter in Abh. Naturw. Ver. Bremen, 24: 336 (1920). inaequilatera (Rusby) Bitter in Abh. Naturw. Ver. Bremen, 24: 439 (1920). leptocaulis (Rusby) Rusby in Bull. Torr. Bot. Club, 53: 218 (1926). lycioides (L.) Hassl. in Ann. Conserv. & Jard. Bot. Genéve, 20: 181 (1917). lycioides var. parvifolia (Wedd.) Bitter in Abh. Naturw. Ver. Bremen, 24: 828 (1920). medians Bitter in Fedde, Repert. Spec. Nov. 18: 317 (1922). polycarpa Rusby in Bull. Torr. Bot. Club, 53: 212 (1926). pseudolycioides (Chod. & Hassl.) Bitter in Abh. Naturw. Ver. Bremen, 24: 352 ? pyrifolia Rusby in Bull. Torr. Bot. Club, 53: 212 (1926). reflexa Rusby in Bull. Torr. Bot. Club, 53: 211 (1926). sancti-caroli (H. Winkl.) Bitter in Abh. Naturw. Ver. Bremen, 24: 354 (1920 oy “al aye ait ok al a ool aot ad x strigosa (Rusby) Bitter in Abh. Naturw. Ver. Bremen, 24: 434. (1920). subfalcata (Rusby) Rusby in Mem. N. Y. Bot. Gard. 7: 349 (1927). tomentella Rusby in Bull. Torr. Bot. Club, 53: 211 (1926). L. viridis Rusby in Mem. N. Y. Bot. Gard. 7: 350 (1927). Lycium cestroides Schlechtd. in Linnaea, 7: 70 (1832). ree ee L. ciliatum Schlechtd. in Linnaea, 7: 69 (1832). L. ciliatum var. latifolium O. Ktze. Rev. Gen. 3(2): 221 (1898). L. colorans U. Damm. in Meded, Rijks Herbar. 29: 22 (1916) = L. cuneatum. L. cuneatum U. Damm. in Engler, Bot. Jahrb. 37: 169, t. 14, fig. 17-19 (1905). L. cyathiformum C. L. Hitche. in Ann. Missouri Bot. Gard. 19: 218, t. 14, fig. 32-34, t. 20 (1982). L. divaricatum Rusby in Bull. N. Y. Bot. Gard. 8: 117 (1912). L. glomeratum Sendt. in Mart. Fl. Bras. 10: 154 (1846). L. graciliflorum U. Damm. in Meded. Rijks Herbar. 29: 23 (1916). L. Herzogii U. Damm. in Meded. Rijks Herbar. 29: 24 (1916). L. nodosum Miers in Ann. & Mag. Nat. Hist. (ser. 2) 14: 189 (1854). L. paucifolium Rusby in Mem. Torr. Bot. Club, 6: 91 (1896) —= L. Tweedianum. L. subtridentatum U. Damm. in Meded. Rijks Herbar. 29: 25 (1916) = L. Tweedianum. L. Tweedianum Griseb. in Goett. Abh. 19: Pe (1874). Markea verrucosa Rusby in Mem. Torr. Bot. Club, 6: 92 (1896). Nicandra physaloides Gaertn. Fruct. 2: 237, t. 181, fig. 2 (1791). Nicotiana Arentsii Goodsp. in Proc. Calif. Acad. Sci. (ser. 4) 25: 297 (1944). Friesii U. Damm. in Fries in Ark. Bot. 1: 409 (1903). glauca R. Grah. in Edinb. New Phil. Journ. 1828: 175. Herzogii U. Damm. in Meded. Rijks Herbar. 29: 30 (1916) = N. wigandio- ides Leguia ana Macbr. in Field Mus. Publ. Bot. 8: 105 (1930). otophora Griseb. in Goett. Abh. 24: 243 (1879). pandurata Dunal in DC. Prodr. 18(1): 569 (1852). plumbaginifolia var. chlorantha Dunal in DC. Prodr. 13(1): 569 (1852). Rusbyi Britton ex Rusby in Bull. Torr. Bot. Club, 27: 23 (1900) = N. wigandioides. sylvestris Speg. in Gartenfl. 47: 131 (1898). tomentosa R. & P. Fl. Peruv. 2: 16, t. 129 (1799). tomentosiformis Goodsp, in Ostenia, 309 (1933). undulata R. & P. Fl. Peruv. 2: 16, t. 180 (1799). wigandioides Koch & Fint. in Wochenschr. 1: 65 (1858). ierembergia boliviana Millan in Darwiniana, 5: 537 (1941). pls mh: BARE 177 N. pulchella Gill. ex Miers in Hook. Lond. Journ. Bot. 5: 173 (1846). Petunia axillaris (Lam.) BSP. Prelim. Cat. N. Y. Pls. 38 (1888). P. occidentalis R. E. Fries in Kgl. Sv. Vetensk. ae Handl. 46(5): 37 (1911). Physalis barbadensis Jacq. Misc. Austr. 2: 359 (1781). P. margaranthoides Rusby in Bull. Torr. Bot. Club, 26: 197 (1899). P. peruviana L. Sp. Pl. (ed. 2) 2: 1670 (1763). P. Rydbergii Rusby in Bull. N. Y. Bot. Gard. 4: 423 (1907). P. surinamensis G. Miq. in Linnaea, 18: 741 (1844). P. viscosa L. Sp. Pl. 183 (1753). Poecilochroma brevifolium Rusby in Bull. N. Y. Bot. Gard. 4: 424 (1907). P. Lobbianum Miers in Hook. Lond. Journ. Bot. 7: 367 (1848). P. macrophyllum por in Bull. N. Y. Bot. Gard. 4: 423 (1907). P. punctatum (R. & P.) Miers in Hook. Lond. Journ. < 7: 354 (1848). P. venosum Rusby in Bull. N. Y. Bot. Gard. 4: 424 (19 Salpichroa amoena Benoist in Bull. Soc. Bot. France, 5: 409 (1938). S. diffusa Miers in Hook. Lond. Journ. Bot. 7: 335 (184 S. glandulosa Miers in Hook. Lond. Journ. Bot. 4: 325 ne S. origanifolia (Lam.) Thell. Fl. Advent. Montpell. 452 (1912). S. sarmentosa Benoist in Bull. Soc. Bot. France, 85: 410 (1938). S. Weddellii Benoist in Bull. Soc. Bot. France, 85: 408 (1938). Salpichroma alatum U. Damm. in Meded. Rijks Herbar. 29: 26 (1916). S. foetidum U. Damm. in Engler, Bot. Jahrb. 37: 639 (1906). S. hirsutum Miers in Hook. Lond. Journ. Bot. 7: 334 (1848). S. Mandonianum Wedd. Chlor. And. 2: 98 (1859). S. rhomboideum var. molle U. Damm. in Engler, Bot. Jahrb. 37: 640 (1906). S. scandens U. Damm. in Engler, Bot. Jahrb. 37: 641 ee S. triste Miers in Hook. Lond. Journ. Bot. 7: 335 (184 Saracha diffusa Miers in Ann. & Mag. Nat. Hist. (ser. ea 3: 447 (1849). Schwenkia americana HBK. Nov. Gen. & Spec. 2: 375, t. 180 (1818). S. hirta Klotzsch in Linnaea, 14: 289 (1840). S. Mandonii Rusby in Bull. N. Y. Bot. Gard. 4: 426 (1907). Schwenkiopsis Herzogii U. Damm. in Meded. Rijks Herbar. 29: 31 (1916). Sessea dependens R. & P. Fl. Peruv. 2: 9, t. 116 (1799). Herzogii U. Damm. in Meded. Rijks Rates 29: 29 (1916). hypotephrodes Bitter in Fedde, Repert. Spec. Nov. 18: 209 (1922). rugosa Rusby in Bull. N. Y. Bot. Gard. 8: 119 (1912) = Cestrum strigillat- um, . stipulata R. & P. Fl. Peruv. 2: 9 (1789). . suaveolens (Rusby) Francey in Notizbl. 11: 984 (1934). . vestita (Hook.) Miers in Hook. Lond. Journ. Bot. 5: 155 (1846). Solandra boliviana Britton ex Rusby in Bull. Torr. Bot. Club, 26: 199 (1899). Solanocharis albescens (Britton) Bitter in Fedde, Repert. Spec. Nov. 15: 153 NQnn (1918). Solanum abutilifolium Rusby in Mem. Torr. Bot. Club, 4(3): 229 (1895). S. acaule Bitter in Fedde, Repert. Spec. Nov. 11: 391 (1912). S. acaule var, subexinterruptum Bitter in Fedde, Repert. Spec. Nov. 11: 393 (1912). actaeabotrys Rusby in Bull. Torr. Bot. Club, 26: 192 (1899). Ahanhuri Juz. & Buk. in Trudy Vsecouz. Sezda Genetike, 3: 605 (1929). aloysiaefolium Dunal in DC. Prodr. 13(1): 73 (1852). alticola Bitter in Fedde, Repert. Spec. Nov. 12: 5 (1913). amaranthifolium Gill. ex Rusby in Bull. Torr. Bot. Club, 26: 152 (1899). andigenum Juz. & Buk. in Trudy Vsecouz. Sezda Genetike, 3: 609 (1929). andigenum var. aymaranum Buk. & Lechn. in Bull. Appl. Bot. Genet. & PI.- Breed. Leningr. Suppl. 58: 68 (1933). min nn nn in 178 mM Nn Ann NANANDAANIAN[n . andigenum var. bolivianum Buk. & Lechn. in Bull. Appl. Bot. Genet. & PIl.- Breed. Leningr. Suppl. 58: 68 (1933). . andigenum var. brevicalyx Buk. & Lechn. in Bull. Appl. Bot. Genet. & Pl.- Breed. Leningr. Suppl. 58: 75 (1933). . andigenum var. Imilla Buk. & Lechn. in Bull. Appl. Bot. Genet. & Pl.-Breed. Leningr. Suppl. 58: 66 (1933). . andigenum var. longibaccatum Buk. & Lechn. in Bull. Appl. Bot. Genet. & Pl.-Breed. Leningr. Suppl. 58: 73 (1933). . andigenum var. stenophyllum Buk. & Lechn. in Bull. Appl. Bot. Genet. & Pl.-Breed. Leningr. Suppl. 58: 74 (1933) . anomalocalyx Hawkes, Potato Coll. Exped. 2: 126 (1944). . anomalocalyx var. brachystylum Card. & Hawkes in Journ. Linn. Soc. 53: 105 (1946). . anomalocalyx var. llallaguanianum Card. & Hawkes in Journ. Linn. Soc. 53: 104 (1946). . anomalocalyx var. murale Card. & Hawkes in Journ. Linn. Soc. 53: 106 946). arachnidanthum Rusby in Mem. N. Y. Bot. Gard. 7: 345 (1927). argenteum Dunal in Poir. Encycl. Suppl. 3: 755 (1818). argentinum Bitter & Lillo in Fedde, Repert. Spec. Nov. 12: 547 (1913). argillicola Dunal in DC. Prodr. 18(1): 236 (1852). asarifolium Kunth & Bouché, Ind. Sem. Hort. Berol. 1845: 10 (1845). asperum Vahl, Symb. 2: 17 (1798). asterosetosum O. Ktze. Rev. Gen. 3(2): 224 (1898). atricoeruleum Bitter in Fedde, Repert. Spec. Nov. 10: 563 (1912). atriplicifolium Gill. ex Nees in Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 386 aureifolium Rusby in Mem. Torr. Bot. Club, 6: 87 (1896). auriculatum Ait. Hort. Kew. 1: 246 (1789). Bangii Bitter in Fedde, Repert. Spec. Nov. 10: 552 (1912). bassoviicarpum Rusby in Bull. N. Y. Bot. Gard. 4: 419 (1907). bermejense Bitter in Fedde, Repert. Spec. Nov. 12: 87 (1913). Berthaultii Hawkes, Potato Coll. Exped. 2: 122 (1944). biceps Dunal in DC. Prodr. 13(1): 341 (1852). bijugum Bitter in Fedde, Repert. Spec. Nov. 10: 583 (1912). bolivianum Britton ex Rusby in Bull. N. Y. Bot. Gard. 4: 416 (1907). boliviense Dunal in DC. Prodr. 18(1): 43 (1852). boliviense subsp. virgultorum Bitter in Fedde, Repert. Spec. Nov. 12: 153 (1913). brevicaule Bitter in Fedde, Repert. Spec. Nov. 11: 390 (1912). brevimucronatum Hawkes, Potato Coll. Exped. 2: 127 (1944). brevipedunculatum Rusby. in Bull. N. Y. Bot. Gard, 4: 421 (1907). aS ita Bitter in Fedde, Repert. Spec. Nov. 10: 558 (1912) = S. lilacin- ae | Kunth & Bouché, Sec. Nov. & Emend. Hort. Bot. Berol. 10 (1853). campylocladum var. magdalense sha in DC. 13(1): 178 (1852). canaminense Rusby in Mem. N. Y. Bot. Gard. 7: 347 (1927). capsibaccatum Cardenas in Rev. oe Univ. Cochabamba, 2(2): 35 (1944). caricaefolium Rusby in Bull. N. Y. Bot. Gard. 8: 118 (1912) Cardenasii Hawkes, Potato Coll. Exped. 2: 120 (1944). carnosipes Rusby in Bull. N. Y. Bot. Gard. 4: 420 (1907). Cervantesii Lag. Gen. & Sp. Nov. 10 (1816). chamaesarachidium Bitter in Fedde, Repert. Spec. Nov. 15: 94 (1917). Chaucha Juz. & Buk. in Trudy Vsecouz. Sezda Genetike, 3: 609 (1929). circaeifolium Bitter in Fedde, Repert. Spec. Nov. 11: 385 (1912). ANNAN mM Minin tn . ANANDNNAANNANNNANNAN Mmnnininh Per errr 179 . clavatum Rusby in Mem. Torr. Bot. Club, 6: 87 (1896). cochabambense Bitter in Fedde, Repert. Spec. Nov. 10: 553 (1912). coerulescens Bitter in Fedde, Repert. Spec. Nov. 10: 554 (1912). coerulescens var. manophyes Bitter in Fedde, Repert. Spec. Nov. 10: 554 (1912). coerulescens var. pycnophyes Bitter in Fedde, Repert. Spec. Nov. 10: 554 (1912). confusum Morton in Contrib. U. S. Nat. Herb. 29: 70 (1944). crinitum Lam, Tabl. Encycl. 2: 20 (1793). crotalobasis Bitter in Fedde, Repert. Spec. Nov. 13: 96 (1914). curtilobum Juz. & Buk. in Trudy Vsecouz. Sezda Genetike, 3: 609 (1929). cymosum R. & P. Fl. Peruv. 2: 31, t. 160 (1799). daphnophyllum Bitter in Fedde, Repert. Spec. Nov. 18: 67 (1922). decachondrum Bitter in Fedde, Repert. Spec. Nov. 11: 228 (1912). decachondrum var. latiusculum Bitter in Fedde, Repert. Spec. Nov. 11: 229 (1912). decachondrum var. longiusculum Bitter in Fedde, Repert. Spec. Nov. 11: 229 decompositiflorum Sendt. in Mart. Fl. Bras. 10: 88 (1846). decorum Sendt. in Mart. Fl. Bras. 10: 83 (1846). decurrentilibum Card. & Hawkes in Journ. Linn. Soc. 53: 97 (1946). deltoideum Rusby, Descr. S. Am. Pls. 115 (1920). devernicascens Bitter in Fedde, Repert. Spec. Nov. 11: 483 (1913) = S. psidiifolium. dianthum Rusby in Bull. N. Y. Bot. Gard. 4: 420 (1907). dibrachiatum v. Heurck & Muell. Arg. in v. Heurck, Obs. Bot. 59 (1870). edinense Berthaut in Ann. Sci. Agron. France (ser. 3) 6(2): 194 (1911). ellipsifolium Card. & Hawkes in Journ. Linn. Soc. 53: 100 (1946). extuspellitum Bitter in Fedde, Repert. Spec. Nov. 10: 555 (1912). extuspellitum var. subcoeruleum Bitter in Fedde, Repert. Spec. Nov. 10: 556 (1912). Fiebrigii Bitter in Fedde, Repert. Spec. Nov. 10: 556 (1912). filiforme R. & P. Fl. Peruv. 2: 31, t. 159 (1799). fragile Wedd. Chlor. And. 2: 105 (1859). Gayanum (Remy) Phil. in Anal. Univ. Chile, 59: 272 (1881). gilioides Rusby in Mem. Torr. Bot. Club, 4(3): 228 (1895). glandulosum R. & P. Fl. Peruv. 2: 35, t. 167 (1799) glandulosum forma acuminatissimum O. Ktze. Rev. Gen. 3(2)+ 226 (1898). glandulosum forma stellatopubescens O. Ktze. Rev. Gen. 3(2): 226 (1898). gonocladum Dunal in DC. Prodr. 13(1): 93 (1852). gonatotrichum Bitter in Fedde, Repert. Spec. Nov. 11: 230 (1912). grandifiorum R, & P. Fl. Peruv. 2: 35, t. 168 (1799). granuloso-leprosum Dunal in DC. Prodr. 13(1): pe (1852). haematocladum Dunal in DC. Prodr. 13(1): 230 (1852). Hauthalii Bitter in Engler, Bot. Jahrb. 50, Beibl. as 61 (1913). hibiscifolium Rusby in Mem. Torr. Bot. Club, 6: 88 (1896). hylobium Bitter in Fedde, Repert. Spec. Nov. 11: = (1912). hyoscyamifolium Rusby in Bull. Torr. Bot. Club, 26: 190 (1899). hyoscyamoides Bitter in Fedde, Repert. Spec. Nov. 11: 236 (1912). hypomalacothrix Bitter in Fedde, Repert. Spec. Nov. 18: 70 (1922). incarceratum R. & P. Fl. Peruv. 2: 40, t. 176, fig. a (1799). inelegans Rusby in Mem. Torr. Bot. Club, 4(3): 229 (1895). insulae-solis Bitter in Fedde, Repert. Spec. Nov. 10: 563 (1912). iodotrichum v. Heurck & Muell. Arg. in v. Heurck, Obs. Bot. 81 (1870). irenaeum Bitter in Fedde, Repert. Spec. sea 10: 551 (1912). jasminoides Paxt. in Mag. Bot. 8: 5 (1841). 180 nnn mn open ae n tn mn th 1 th on . Nnnnnn mM ool akg aaa eae juglandifolium Humb. & Bonpl. ex Dunal, Solan. Syn. 6 (1816). Juzepezukii Buk. in Trudy Vsecouz. Sezda Genetike, 3: 603 (1929). lanceaefolium Jacq. Collect. 2: 286 (1788). lapazense Hawkes, Potato Coll. Exped. 2: 127 (1944). Lauterbachii (H. Winkl.) Bitter in Fedde, Repert. Spec. Nov. 15: 155 (1918). Lechleri Rusby in Bull. Torr. Bot. Club, 26: 193 (1899). leptocaulon v. Heurck & Muell. Arg. in v. Heurck, Obs. Bot. 40 (1870). leptophyes Bitter in Fedde, Repert. Spec. Nov. 12: 448 (1913). lilacinum Rusby in Bull. Torr. Bot. Club, 26: 192 (1899). Lindenii Rusby in Mem. Torr. Bot. Club, 6: 88 (1896). liriunianum Card. & Hawkes in Journ. Linn. Soc. 53: 106 (1946). lycopersicoides Dunal in DC. Prodr. 13(1): 38 (1852). maglia Schlechtd. Hort. Hal. 6 (1841). mammosum L. Sp. Pl. 187 (1753). Mandonii A. DC. in Bibl. Univ.: Arch. Sci. Phys. & Nat. (ser. 3) 15: 4388 (1886). Mandonis v. Heurck & Muell. Arg. in v. Heurck, Obs. Bot. 78 (1870). Mandonis var. dryophyllum Bitter in Fedde, Repert. Spec. Nov. 11: 487 (1913). . Mandonis var. tardecalvescens Bitter in Fedde, Repert. Spec. Nov. 11: 487 (1913). maptriense Bitter in Fedde, Repert. Spec. Nov. 11: 16 (1913) = S. phytolac- coides. medianiviolaceum Bitter in Fedde, Repert. Spec. Nov. 10: 562 (1912 yes megalochiton var. villosotomentosum Dunal in DC. Prodr. 13(1): 124 (1852). megistacrolobium Bitter in Fedde, Repert. Spec. Nov. 10: microdontum Bitter in Fedde, Repert. Spec. Nov. 10: 535 (1912). minutibaccatum Bitter in Fedde, Repert. Spec. Nov. 10: 549 (1912). minutibaccatum var. curtipedunculatum Bitter in Fedde, Repert. Spec. Nov. 11: 205 (1912). mite subsp. hexazygum Bitter in Fedde, Repert. Spec. Nov. 11: 10 (1912). mollepujroénse Card. & Hawkes in Journ. Linn. Soc. 53: 108 (1946). montanum L. Sp. Pl, 186 (1753). muricatum var. papillosistylum Bitter in Fedde, Repert. Spec. Nov. 11: 359 (1912). muricatum var. teleutogenum Bitter in Fedde, Repert. Spec. Nov. 12: 443 (1913). myrianthum Britton ex Rusby in Bull. Torr. Bot. Club, 26: 191 (1899). nanum Bitter in Fedde, Repert. Spec. Nov. 10: 564 (1912). narcoticum Bitter in Fedde, Repert. Spec. Nov. 13: 97 (1914). nigrum L. Sp. Pl. 186 (1753). nitidibaccatum var. robusticalyx Bitter in Fedde, Repert. Spec. Nov. 11: 209 (1912). . nitidum R. & P. Fl. Peruv. 2: 33, t. 163, fig. a (1799 ). nudum Humb. & Bonpl. ex Dunal, Solan. Syn. (ed. 2) 20 (1816). nutans R. & P. Fl. Peruv. 2: 34, t. 166 (1799). ochrophyllum v. Heurck & Muell. Arg. in v. Heurck, Obs. Bot. 50 (1870). oligodontum Bitter in Fedde, Repert. Spec. Nov. 11: 215 (1912). oplocense Hawkes, Potato Coll. Exped. 2: 119 (1944). pachyantherum Bitter in Fedde, Repert. Spec. Nov. 11: 206 (1912). pachytrichum Hawkes, Potato Coll. Exped. 2: 121 (1944). pallidum Rusby in Mem. Torr. Bot. Club, 4(3): 228 (1895). Pearcei Britton ex Rusby in Mem. Torr. Bot. Club, 4(3): 227 (1895). Pentlandii Dunal in DC. Prodr. 13(1): 51 (1852). Phureja Juz. & Buk. in Trudy Vsecouz. Sezda Genetike, 3: 604 (1929). tn os beaaalm ANNARANDANANDNANNANANDNANNNNNANANDANNNANANDANDMNMM Nin Phureja var. Pujeri Hawkes, Potato Coll. Exped. 2: 128 (1944). physalifolium Rusby in Mem. Torr. Bot. Club, 6: 88 (1896). phytolaccoides (Rusby) Bitter in Fedde, Repert. Spec. Nov. 13: 172 (1914). pinnatifidum Cardenas in Rev. Agric. Univ. Cochabamba, 2(2): 33 (1944). platypterum Hawkes, Potato Coll. Exped. 2: 118 (1944). poecilochromifolium Rusby in Bull. N. Y. Bot. Gard. 4: 419 (1907). poinsettiaefolium Rusby in Mem. N. Y. Bot. Gard. 7: 344 (1927). polytrichostylum Bitter in Fedde, Repert. Spec. Nov. 10: 550 (1912). polytrichum Moric. P!. Nouv. Amér. 32, t. 22 (1837). pongoénse Rusby in Mem. N. Y. Bot. Gard. 7: 348 (1927) = S. leptocaulon. psidiifolium Rusby in Bull. Torr. Bot. Club, 26: 194 (1899). pterocladum v. Heurck & Muell. Arg. in v. Heurck, Obs. Bot. 44 (1870). pulverulentum Pers. Synops. Pl. 1: 223 (1805). Pycnanthemum Mart. in Flora, 20, II Beibl.: 120 (1837). radicans L. f. Dec. 1: 19, t. 10 (1762). rheithrocharis Bitter in Fedde, Repert. Spec. Nov. 13: 91 (1914). rogaguense Rusby in Mem. N. Y. Bot. Gard. 7: 346 (1927). rosulatum Rusby in Bull. N. Y. Bot. Gard. 4: 418 (1907). rugosum Dunal in DC. Prodr. 13(1): 108 (1852). Rusbyi Britton ex Rusby in Bull. Torr. Bot. Club, 26: 191 (1899). Saccianum Carr. & André in Rev. Hort. 64: 50 (1892 Salzmannii Dunal in DC. Prodr. 13(1): 206 (1852). sanctae-crucis Dunal in DC. Prodr. 13(1): 274 (1852). sarachioides Rusby in Bull. N. Y. Bot. Gard. 4: 420 (1907). sassafrideum Rusby in Mem. N. Y. Bot. Gard. 7: 348 (1927). Schlechtendalianum Walp. Repert. 3: 61 (1844). scotinectarium Bitter in Fedde, Repert. Spec. Nov. 10: 560 (1912). sericeum R. & P. Fl. Peruv. 2: 33, t. 161 (1799). setosicalyx Rusby in Mem. N. Y. Bot. Gard. 7: 344 (1927). simplicifolium Bitter in Fedde, Repert. Spec. Nov. 11: 369 (1912). sinuatirecurvum Bitter in Fedde, Repert. Spec. Nov. 11: 241 (1912). sinuatiexcisum Bitter in Fedde, Repert. Spec. Nov. 10: 558 (1912). sisymbrifolium Lam. Tabl. Encycl. 2: 25 (1793). sisymbrifolium forma lilacinum O. Ktze. Rev. Gen. 3(2): 227 (1898). sordidum Sendt. in Mart. Fl. Bras. 10: 53, t. 4 (1846). Sprucei v. Heurck & Muell. Arg. in v. Heurck, Obs. Bot. 67 (1870). steironematophyllum Rusby in Mem. N. Y. Bot. Gard. 7: 346 (1927). stellativelutinum Bitter in Fedde, Repert. Spec. Nov. 16: 397 (1920). stenotomum Juz. & Buk. in Trudy Vsecouz. Sezda Genetike, 3: 604 (1929). stenotomum var. cyaneum Hawkes, Potato Coll. Exped. 2: 57 (1944). stenotomum var. megalocalyx Hawkes, Potato Coll. Exped. 2: 59 (1944). stenotomum var. Pitiquilla Hawkes, Potato Coll. Exped. 2: 54 (1944). stenotomum var. Puca-lunca Hawkes, Potato Coll. Exped. 2: 61 (1944). stipuloideum Rusby in Bull. N. Y. Bot. Gard. 4: 419 (1907). styracioides Rusby in Mem. Torr. Bot. Club, 4(3): 230 (1895). suaveolens var. heterotrichostylum Bitter in Fedde, Repert. Spec. Nov. 11: 856 (1912) suaveolens subsp. microphyllidium Bitter in Fedde, Repert. Spec. Nov. 11: 357 (1912). subandigenum Hawkes, Potato Coll. Exped. 2: 128 (1944). subauriferum Bitter in Fedde, Repert. Spec. Nov. 10: 559 (1912). subtusviolaceum Bitter in Fedde, Repert. Spec. Nov. 11: 207 (1912) sucrense Hawkes, Potato Coll., Exped. 2: 126 (1944). sucrense var. brevifolium Biwkes: Potato Coll. Exped. 2: 51 (1944). symmetricum Rusby in Mem. Torr. Bot. Club, 6: 89 (1896). spaicanatéiiein Rusby in Bull. N. Y. Bot. Gard. 4: 418 (1907). 182 tabacifolium Salzm. ex Dunal in DC. Prodr. 13(1): 261 (1852). tarijense Hawkes, Potato Coll. Exped. 2: 114 (1944). tenuifilamentum Juz. & Buk. in Trudy Vsecouz. Sezda Genetike, 3: 603 mn ta ta tenuispinum Rusby in Phytologia, 1: 75 (1934) mR. & Pi Fl. Perevi 2: 88; t:: 172 (1799). Theresiae A. Zahlbr. in Beih. Bot. Centralbl. 13: 83 (1902). toralopanum Card. & Hawkes in Journ. Linn. Soc. 53: 98 (1946) toralopanum var. subintegrifolium Card. & Hawkes in Journ. Linn. Soc. 53: 99 (1946). trifurcum Dunal in DC. Prodr. 13(1): 272 (1852). tripartitum Dunal in DC. Prodr. — 72 (1852). tuberosum L. Sp. Pl. 185 (1753 . tuberosum subsp. sparsipilum Lf in Fedde, Repert. Spec. Nov. 12: 152 (1913). tunariense O. Ktze. Rev. Gen. 3(2): 228 (1898). turneroides Chod. in Bull. Herb. Boiss. (ser. 2) 2: 814) (1902). ursinum Rusby in Mem. Torr. Bot. Club, 4(3): 227 (1895). urticans Dunal in DC. Prodr. 13(1): 324 (1852). utile Klotzsch in Allg. Gartenz. 17: 315 (1839). validum Rusby in Mem. Torr. Bot. Club, 4(3): 230 (1895). velutissimum Rusby in Mem. Torr. Bot. Club, 6: 89 (1896). verbascifolium L. Sp. Pl. 184 (1753). verbascifolium var. caducum O. Ktze. Rev. Gen. 3(2): 228 (1898). verniciflorum Bitter in Fedde, Repert. Spec. Nov. 18: 66 (1922). violaceimarmoratum Bitter in Fedde, Repert. Spec. Nov. 11: 389 (1912). violaceimarmoratum var. papillosum Hawkes, Potato Coll. Exped. 2: 113 (1944). violaceistriatum Bitter in Fedde, Repert. Spec. Nov. 9: 550 (1912). violifolium Schott ex Spreng. Syst. Veg. 4: Cur. Post. 408, no. 5 (1827). volubilis Rusby in Bull. Torr. Bot. Club, 26: 194 (1899). vulpinum Rusby in Bull. N. Y. Bot. Gard. 4: 418 (1907). Williamsii Rusby in Bull. N. Y. Bot. Gard. 8: 118 (1912). Wrightii Benth. Fl. Hongk. 248 (1861). yapacaniense O. Ktze. Rev. Gen. 3(2): 228 (1898). yungasense Hawkes in Ann. & Mag. Nat. Hist. 7 (81): 697 (1954). Tunaria albida O. Ktze. Rev. Gen. 3(2): 228 (1898). Vassobia atropoides Rusby in Bull. N. Y. Bot. Gard. 4: 422 (1907). V. dichotoma (Rusby) Bitter in Fedde, Repert. Spec. Nov. 15: 150 (1918). mnt nn ANNAnMnNMnAM nin in Nnnnnnn yn SCROPHULARIACEAE Alonsoa acutifolia R. & P. Syst. 153 (1798). A. incisaefolia R. & P. Syst. 154 (1798). Angelonia acuminatissima Herzog in Meded. Rijks Herbar. 29: 33 (1916). A. chiquitensis Herzog in Meded. Rijks Herbar. 29: 33 (1916). Bacopa Monniera (L.) Wettst. in Engl. & Prantl, Nat. Pflanzenfam. 4(3b): 77 (1891). Bartsia altissima Rusby in Phytologia, 1: 75 (1934). . breviflora Benth. in DC. Prodr. 10: 545 (1846). ciliolata Wedd. Chlor. And. 2: 123 (1860). crenoloba Wedd. Chlor. And. 2: 124 (1860). . diffusa Benth. in DC. Prodr. 10: 546 bo bd bo od bo bo be ( Guggenheimiana Rusby in Phytologia, 1: 76 (1934). bo td bd be be be bd bo 183 hispida Benth. in DC. Prodr. 10: 547 (1846). inaequalis Benth. in DC. Prodr. 10: 547 (1846). laxiflora Benth. in DC. Prodr. 10: 547 (1846). Meyeniana Benth. in DC. Prodr. 10: 546 (1846). mutica (HBK). Benth. in DC. Prodr. 10: 548 (1846). patens Benth. in DC. Prodr. 10: 546 (1846). peruviana Walp. in Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 400 (1843). pumila Benth. in DC. Prodr. 10: 546 (1846). sanguinea Diels in Engler, Bot. Jahrb. 37: 433 (1906). Basistemon Rusbyi Moldenke in Bull. Torr. Bot. Club, 63: 348 (1936). B. B. silvaticum (Herzog) Baehni & Macbr. in Candollea, 5: 345 (1934). spinosum (Chod.) Moldenke in Bull. Torr. Bot. Club, 63: 347 (1936). Buchnera elongata Sw. Prodr. 92 (1788), nomen illegitimum B. elongata forma rigidior O. Ktze. Rev. Gen. 3(2): 230 (1898). Calceolaria aquatica A. Br. & Bouché, Ind. Sem. Hort. Berol. 2 (1852). AH aaAAaAsaAanaAaAaaaaA ANRAA ANAAHAAA eat VN ae a Atahualpae Kranzl. in Engler, Pflanzenr. IV, 257C: 107 (1907). bartsiaefolia Wedd. Chlor. And. 2: 134 (1860). boliviana (Britton) Pennell in Proc. Acad. Nat. Sci. Philad. 97: 170 (1945). Buchtieniana Kranzl. in Fedde, Repert. Spec. Nov. 5: 369 (1908). camptoclada Krinzl. in Engler, Pflanzenr. IV, 257C: 74 (1907). canescens Willd. ex Roem, & Schult. Syst. Mant. + 165 (1822). chelidonioides HBK. Nov. Gen. & Spec. 2: 378 (1818). conocarpa Pennell in Proc. Acad. Nat. Sci. Philad. 97: 172 (1945). cryptantha Rusby in Phytologia, 1: 77 (1934). C. cuneiformis R. & P. Fl. Peruv. 1: 19, t. 27, fig. 6 (1798). defiexa R. & P. Fl. Peruv. 1: 18, t. 30b (1798). dichotoma Lam. Encycl. 1: 555 (1783). elatior Griseb. in Goett. Abh. 24: 237 (1879). elliptica Wedd. Chlor. And. 2: 137 (1860). endotrachys Kranzl. in Fedde, Repert. Spec. Nov. 27: 22 (1929) = C, En- gleriana. . Engleriana Krinzl. in Fedde, Repert. Spec. Nov. 1: 106 (1905). oneoraceee Kranzl. in Ann. Naturh. Hofm. Wien, 22: 194 (1907) = C. En- leri Renae ‘Kranal. in Fedde, Repert. Spec. Nov. 27: 24 (1929). extensa Benth. in DC. Prodr. 10: 214 (1846). Fiebrigiana Kranz]. in Fedde, Repert. Spec. Nov. 1: 84 (1905). glacialis Wedd. Chlor. And. 2: 143, t. 59A (1860). Guentheri Kranzl. in Fedde, Repert. Spec. Nov. 27: 17 (1929). Halliana Kranzl. in Fedde, Repert. Spec. Nov. 1: 85 (1905). Herzogiana Krianzl. in Engler, Bot. Jahrb. 50, Beibl. — 69 (1913). hypericina Poepp. ex Benth. in DC. Prodr. 10: 222 (184 hypoleuca Krianzl. in Fedde, Repert. Spec. Nov. 27: = (1929). incachacensis Kranz]. in Fedde, Repert. Spec. Nov. 27: 20 (1929) = C. En- gleriana. inflexa R. & P. Fl. Peruv. 1: 16, t. 25 (1798). larecajensis Kranzl. in Fedde, Repert. Spec. Nov. 16: 450 (1920). lechioides Rusby in Phytologia, 1: 77 (1934) = C. parvifolia. leiophylla Kranzl. in Engler, Bot. Jahrb. 50, Beibl. 111: 70 (1913). lobata Cav. Ic. 5: 26, t. 443 (1799). hylla Kranzl. in Engler, Bot. Jahrb. 50, Beibl. 111: 67 (19138) = C. larecajensis. melissaefolia Benth. in DC. Prodr. 10: 214 (1846). monantha Kranzl. in Fedde, Repert. Spec. Nov. 27: 20 (1929). palustris Sodiro & Kranzl. in Engler, Pflanzenr. IV, 257C: 27 (1907). parvifolia Wedd. Chlor. And. 2: 134 (1860). 184 Pavonii Benth. in DC. Prodr. 10: 211 (1846). Pearceana Kranzl. in Engler, Pflanzenr. IV, 257C: 107 (1907). Pflanzii Perkins in Engler, Bot. Jahrb. 49: 218 (1913). pinnata L, Mant. 171 (1771). poikilantha Kranzl. in Fedde, Repert. Spec. Nov. 27: 12 (1929). polyclada Krianzl. in Fedde, Repert. Spec. Nov. 1: 105 (1905). ribesiaefolia Rusby in Phytologia, 1: 78 (1934). rivularis Kranzl. in Fedde, Repert. Spec. Nov. 1: 83 (1905). C. saxatilis HBK. Nov. Gen. & Spec. 2: 382 (18 vi scabra R. & P. Fl. Peruv. 1: 19, t. 29, fig. a (1798). soratensis Kranzl. in Engler, Pflanzenr. IV, eae 105 (1907). teucrioides Griseb. in Goett. Abh. 19: 212 (18 74). trilobata Hemsl. in Biol. Centr. Am. Bot. 2: 439 (1882). tripartita Mandoniana (Krianzl.) Pennell in Proc. Acad. Nat. Sci. Philad. 97: 175 (1945). umbellata Wedd. Chlor. And. 2: 142 (1860). . vaccinioides Kranzl. in Fedde, Repert. Spec. Nov. 27: 17 (1929). . virgata R. & P. Fl. Peruv. 1: 20, t. 31 (1798). Capraria biflora L. Sp. Pl. 628 (1753). Castilleja communis Benth. in DC. Prodr. 10: 529 (1846). C. fissifolia L. f. Suppl. 293 (1781). C. fissifolia subsp. fruticosa Wedd. Chlor. And. 2: 118 (1860). C. fissifolia subsp. pumila (Benth.) Wedd. Chlor. And. 2: 119 (1860). Escobedia brevipes Pennell in Proc. Acad. Nat. Sci. Philad. 83: 418 (1931). E. obtusifolia Pennell in Proc. Acad. Nat. Sci. Philad. 83: 419 (1981). E. scabrifolia R. & P. Syst. 159 (1798). Esterhazya andina Herzog in Meded. Rijks Herbar. 29: 36 (1916). Fagelia Bangii Rusby in Mem. Torr. Bot. Club, 4(3) : 286 (1895) — Calceolaria bartsiifolia ? F. boliviana Britton ex Rusby in Bull. Torr. Bot. Club, 27: 24 (1900) — Cal- ceolaria sp. Gerardia Bangii O. Ktze. Rev. Gen. 3(2): 233 (1898). brevifolia Rusby in Bull. Torr. Bot. Club, 27: 25 (1900). Fiebrigii Diels in Engler, Bot. Jahrb. 37: 428 (1906). Fiebrigii var. brevidens Herzog in Meded. Rijks Herbar. 29: 38 (1916). genistifolia Cham. & Schlechtd. in Linnaea, 3: 15 (1828). humilis Diels in Engler, Bot. Jahrb. 37: 429 (1906). lanceolata (R. & P.) Benth. in Hook. Compan. Bot. Mag. 1: 207 (18386). lanceolata var. revoluta (R. & P.) O. Ktze. Rev. Gen. 3(2): 233 (1898). linarioides Cham. & Schlechtd. in Linnaea, 3: 18 ( : macrodonta var. latifolia O. Ktze. Rev. Gen. 3(2): 234 (1898). ovatifolia Rusby in Bull. N. Y. Bot. Gard. 4: 427 (1907). reflexidens Herzog in Meded. Rijks Herbar. 29: 37 (1916). rigida Gill. ex Benth. in Hook: Compan. Bot. Mag. 1: 206 (1836). scarlatina Herzog in Meded. Rijks Herbar. 29: 38 (1916). splendida (Mikan) O. Ktze. Rev. 3(2): 234 (1898). tarijensis R. E. Fries in Ark. Bot. 6(11): 19 (1906). Ilysanthes gratioloides (L.) Benth. in DC. Prodr. 10: 419 (1846)). Lendneria mone: (Soland.) Minod in Bull. Soc. Bot. Genéve, 10: 241 (1918) == Capraria ? Leucocarpus slab G Don ex Sweet, Brit. Fl. Gard. 2: t. 124 (1831). Limosella americana Gliick in Notizbl. 12: 75 (1984). L. aquatica L. Sp. Pl. 631 (1753). Linaria canadensis var. texana (Scheele) Pennell in Proc. Acad. Nat. Sci Philad. 73: 502 (1922). Mimulus glabratus HBK. Nov. Gen. & Spec. 2: 370 (1818). ana as a aleahla ye AA AAARAARARAAAA 185 M. parviflorus var. nanus Wedd. Chlor. And. 2: 132 (1860). Ourisia biflora Wedd. Chlor. And. 2: 117 (1860). chamaedrifolia Benth. in DC. Prodr. oe Ae (184 . chamaedrifolia var. elegans Wedd. Chlor. And. 114 (1860). - muscosa Benth. in DC. Prodr. 10: 493 (1848). nana Benth. in DC. Prodr. 10: 493 (1846). pulchella Wedd. Chlor. And. 2: 116 (1860). rupicola Wedd. Chlor. And. 2: 115 (1860). Saccanthus violaceus Herzog in Meded. Rijks Herbar. 29: 47 (1916) = Basi- stemon spinosum. Scoparia annua Cham. & Schlechtd. in Linnaea, 6: 375 (1831). S. dulcis L. Sp. Pl. 116 (1758). S. nudicaulis subsp. praedensa R. E. Fries in Ark. Bot. 6(9): 12 (1906). Sibthorpia conspicua Diels in Engler, Bot. Jahrb. 37: 428 (1906 S. nectarifera Wedd. Chlor. And. 2: 211, t. 60B (1859). S. pichinchensis HBK. Nov. Gen. & Spec. 2: 390, t. 176 (1818) . S. retusa HBK. Nov. Gen. & Spec. 2: 391, t. 177 (1818) Stemodia lanceolata Benth. in DC. Prodr. 10: 384 (1846). S. parviflora Ait. Hort. Kew. (ed. 2) 4: 52 (1812). S. stricta Cham. & Schlechtd. in Linnaea, 3: 10 (1828). Vandellia diffusa L. Mant. 89 (1767). Verbascum virgatum Stokes in With. Bot. Arr. Brit. Pls. (ed. 2) 1: 227 (1781). Veronica peregrina L. Sp. Pl. 14 (1758). Virgularia ochrophylla Rusby in Mem. N. Y. Bot. Gard. 7: 350 (1927) = Gerardia sp. 29990900" BIGNONIACEAE Adenocalymma bracteatum (Cham.) DC. Prodr. 9: 200 (1845). A. densiflorum Rusby in Mem. N. Y. Bot. Gard. 7: 355 (1927). A. impressum (Rusby) Sandw. in Rec. Trav. Bot. Néerl. 34: 212 (1937). A. latifolium Rusby, Descr. S. Am, Pls. 121 (1920). Amphilophium molle Cham. & Schlechtd. in Linnaea, 5: 120 (1830). A. paniculatum (L.) HBK. Nov. Gen. & Spec. 3: 149 (1818). A. Vauthieri DC. Prodr. 9: 193 (1845). Anemopaegma huachianum Rusby in Mem. N. Y. Bot. Gard. 7: 352 (1927). A. leptosiphon Rusby in Mem. N. Y. Bot. Gard. 7: 354 (1927). A. symmetricum Rusby in Mem. N. Y. Bot. Gard. 7: 353 (1927). Arrabidaea arthrerion (DC.) Bureau ex K. Schum. in Engl. & Prantl, Nat. Pflanzenfam. 4(3b): 213 (1894). . Bangii Sprague in Bull. Herb. Boiss. (ser. 2) 6: 371 (1906). candicans (Rich.) DC. Prodr. 9: 185 (1845). . Cardenasii Rusby in Mem. N. Y. Bot. Gard. 7: 351 (1927). . floribunda (HBK.) Loes. in Fedde, Repert. Spec. Nov. 16: 209 (1919). . florida DC. Prodr. 9: 184 (1845 macrocarpa O. Ktze. Rev. Gen. 3(2): 242 (1898). obovata DC. Prodr. 9: 185 (1845). . Orbignyana DC. Prodr. 9: 184 (1845). . pachycalyx Sprague in Bull. Herb. Boiss. (ser. 2) 6: 373 (1906). . pentstemonoides Krinzl. in Notizbl. 6: 370 (1915). . platyphylla (Cham.) Bur. & K. Schum. in Mart. Fl. Bras. 8(2): 38 (1896). Hignanin benensis Britton ex Rusby in Bull. Torr. Bot. Club, 27: 70 (1900). B. boliviana Rusby in Bull. Torr. Bot. Club, 27: 70 (1900). B. brachypoda DC. Prodr. 9: 145 (1845). B. brevipes Rusby in Bull. Torr. Bot. Club, 27: 71 (1900). B. capreolata L. Sp. Pl. 624 (1753). >>> >> >>> >>> 186 glutinosa DC. Prodr. 9: 162 (1845). nivea Kranzl. in Engler, Bot. Jahrb. 54, Beibl. 119: 26 (1916). Pearcei Rusby in Mem. Torr. Bot. Club, 6: 100 (1896) pyramidata Rich. in Act. Soc. Hist. Nat. Paris, 1: 110 (1792). Rusbyi Britton ex Rusby in Bull. Torr. Bot. Club, 27: 71 (1900). tecomaeflora Rusby in Mem. oe Bot. Club, 6: 101 (18 o6)- unguis-cati L. Sp. Pl. 623 (175 venusta Ker in Bot. Reg. 3: t. on (1818). Callichlamys riparia Miq. in Linnaea, 18: 254 (1844). Cremastus pulcher (Cham.) Bureau in Vidensk. Meddel. Kjoebenh. 1893: 101 4). bo td bd te be bo bd be C. rufo-villosus Herzog in Meded. Rijks Herbar. 29: 40 (1916). Crescentia Cujete L. Sp. Pl. 626 (1753). Cuspidaria ovalis Rusby, Descr. S. Am. Pls. 120 (1920). C. Schumanniana O. Ktze. Rev. Gen, 3(2): 243 (1898). Cybistax antisiphilitica Mart. ex DC. Prodr. 9: 199 (1845). Gelseminum amoenum var. tomentosum O. Ktze. Rev. Gen. 3(2): 245 (1898). . amoenum var. pubescens O. Ktze. Rev. Gen. 3(2): 245 (1898). G. Avellanedae (Griseb.) O. Ktze. Rev. Gen. 3(2): 245 (1898). G. Garrocha var. bicolor O. Ktze. Rev. Gen. 3(2): 245 (1898). G. Garrocha var. rubrum O. Ktze. Rev. Gen. 3(2): 245 (1898). G. molle (HBK.) O. Ktze. Rev. Gen. 3(2): 245 (1898). Glaziova sp. Godmania aesculifolia ap Standl. in Standl. & Calderén, List. Prelim. PI. El Salvador, 200 (1925). Jacaranda acutifolia Humb. & Bonpl. Pl. Aequin. 1: 59, t. 17 (1805). J. atropurpurea Rusby in Mem. N. Y. Bot. Gard. 7: 357 (1927). J. Copaia var. spectabilis (Mart. ex DC.) Bureau in Mart. Fl. Bras. 8(2): 387 (1897). J. cuspidifolia Mart. in Flora, 24, II Beibl.: 51 (1841). J. longiflora Britton ex Rusby in Bull. Torr. Bot. Club, 27: 73 (1900). Lundia densiflora DC. Prodr. 9: 181 (1845). L, phaseolifolia Rusby in Mem. Torr. Bot. Club, 6: 99 (1896). L. Spruceana Bureau in Adansonia, 8: 279 (1867-68). L. truncata Rusby in Mem. N. Y. Bot. Gard. 7: 356 (1927). Macfadyena Bangii Rusby in Bull. Torr. Bot. Club, 27: 72 (1900). M. uncata (Andr.) Sprague & Sandw. in Meded. Bot. Mus. Utrecht, 40: 215 (1937). M. undulata K. Schum. in Engl. & Prantl, Nat. Pflanzenfam. 4(3b): 227 (1894), nomen nudum. M. violacea Rusby in Mem. N. Y. Bot. Gard. 7: 356 (1927). Martinella obovata (Spreng.) Bur. & K. Schum. in Mart. Fl. Bras. 8(2): 161 (1896). Melloa populifolia (DC.) Britton in Ann. N. Y. Acad. Sci. 7: 188 (1893). Mussatia hyacinthina (Standl.) Sandw. in Rec. Trav. Bot. Néerl. 34: 218 (1937). Paragonia pyramidata (Rich.) Bureau in Vidensk. Medd. Naturh. Foren. 1893: 104 (1894). Petastoma laurifolium Kranzl. in Fedde, Repert. Spec. Nov. 17: 57 (1921). . multiglandulosum (Benth.) Krianzl. in Fedde, Repert. Spec. Nov. 17: 58 (1921). patelliferum (Schlechtd.) Miers in Proc. Roy. Hort. Soc. 3: 195 (1863). samydoides Miers in Proc. Roy. Hort. Soc. 3: 195 (1863). truncatum (Sprague) Hassl. in Fedde, Repert. Spec. Nov. 9: 53 (1910). Whitei (Rusby) Sandw. in Kew Bull. 1953: 462. Pithecoctenium cynanchoides DC. Prodr. 9: 195 (1845). ao) ty td 187 P. echinatum K. Schum. in Mart. Fl. Bras. 8(2): 168, t. 86 (1896). P. glaucum Rusby in Mem. N. Y. Bot. Gard. 7: 354 (1927) P. Lundii P. DC. Prodr. 9: 196 (1845). Pleonotoma jasminifolium (HBK.) Miers in Proc. Roy. Hort. Soc. 3: 184 (1863) Pyrostegia venusta Miers in Proc. Roy. Hort. Soc. 3: 188 (1863). Saldanhaea confertiflora Bureau in Adansonia, 8: 356 (1867-68). ? S. mollis Kranzl. in Engler, Bot. Jahrb. 54, Beibl. 119: 26 (1916). Scobinaria s Stenolobium Cac tocbk (Hieron.) R. E. Fries in Ark. Bot. 6(11): 16 (1904). Tabebuia nodosa Griseb. in Goett. Abh. 24: 258 (1879 T. Caraiba (Mart.) Bureau in Vidensk. Meddel. 1893: 113 (1894). T. roseo-alba (Ridley) Sandw. in Kew Bull. 1954: 597 (1955). T. serratifolia (Vahl) Nicholson, Dict. Gard. 4: 1 (1888). T. suberosa Rusby in Mem. N. Y. Bot. Gard. 7: 358 (1927). Tecoma capensis Lindl. in Bot. Reg. 18: t. 1117 (1828). T. cochabambensis (Herzog) Sandw. in Kew Bull. 1953: 455. T. Gaudichaudii DC. Prodr. 9: 223 (1845). T. mollis HBK. Nov. Gen. & Spec. 3: 144 (1819). T. sambucifolia HBK. Nov. Gen. & Spec. 3: 143 (1819). Tynnanthus micranthus Mello in Arg. Mus. Paranaénse, 9: 63 (1952). T. myrianthus (Poepp.) Bur. & K. Schum. in Mart. Fl. Bras. 8(2): 197 (1896). T. polyanthus (Bur.) Sandw. in Kew Bull. 1953: 465. Zeyhera Kuntzei K. Schum. in O. Ktze. Rev. Gen. 3(2): 246 (1898). MARTYNIACEAE Craniolaria integrifolia Cham. in Linnaea, 7: 725 (1832). GESNERIACEAE Achimenes flaccida Rusby in Mem. Torr. Bot. Club, 6: 95 (1896). A. gracilis Britton ex Rusby in Bull. Torr. Bot. Club, 27: 27 (1900). A. heppielloides Fritsch in Bull. Torr. Bot. Club, 23: 151 (1896). A. Rusbyi Britton ex Rusby in Mem. Torr. Bot. Club, 6: 95 (1896). Alloplectus dichrous DC. Prodr. 7: 546 (1839). A. grandifolius Britton ex Rusby in Bull. Torr. Bot. Club, 27: 30 (1900). A. Patrisii DC. Prodr. 7: 545 (1839). A. solitarius Rusby in Mem. Torr. Bot. Club, 4(3): 238 (1895). Besleria aurantiaca Fritsch in Meded. Rijks Herbar. 29: 51 (1916) — B. longipedunculata. boliviana Morton in Contrib. U. S. Nat. Herb. 26: 432 (1939). elegans HBK. Nov. Gen. & Spec. 2: 397 (1817). foliacea Rusby in Mem. Torr. Bot. Club, 4(3): 240 (1895). longipedunculata Britton ex Rusby in Bull. Torr. Bot. Club, 27: 69 (1900). montana Britton ex Rusby in Mem. Torr. Bot. Club, 4(3): 240 (1895). ovalifolia Rusby in Mem. Torr. Bot. Club, 4(3): 240 (1895). pauciflora Rusby in Bull. Torr. Bot. Club, 27: 31 (1900). Rhytidophyllum Hanst. in Linnaea, 34: 332 (1865-66)). rotundifolia Rusby in Mem. Torr. Bot. Club, 6: 98 (1896). Sprucei Britton ex Rusby in Bull. Torr. Bot. Club, 27: 31 (1900). Uleana Fritsch in Engler, Bot. Jahrb. 37: 483 (1906) — B. Sprucei. Columnea ascendens Rusby in Mem. Torr. Bot. Club, 4(3): 239 (1895). C. boliviana Britton ex Rusby in Mem. Torr. Bot. Club, 4(3): 238 (1895). C. grandifolia Rusby, Descr. S. Am. Pls. 126 Sony C. inaequilatera Poepp. & Endl. Nov. Gen. 3: 1 (1845). C. latisepala Rusby in Mem. Torr. Bot. Club, 4(3): 239 (1895). DH W WNW 188 C. oblongifolia Rusby in Mem. Torr. Bot. Club, 6: 98 (1896). C. pallida Rusby, Descr. S. Am. Pls. 125 (1920). C. stricta Rusby, Descr. S. Am. Pls. 125 (1920). C. Trollii Mansf. in Fedde, Repert. Spec. Nov. 36: 122 (1934). Corytholoma paludosum Rusby in Mem. N. Y. Bot. Gard. 7: 360 (1927). Diastema galeopsis Fritsch in Engler, Bot. Jahrb. 50: — (1913). D. latiflorum Rusby in Mem. Torr. Bot. Club, 6: 96 (189 D. parviflorum (Rusby) Fritsch in Engler, Bot. Jahrb. wf 406 (1913). D. purpurascens Rusby in Bull. Torr. Bot. Club, 27: 28 (1900). D. Williamsii Rusby, Descr. S. Am. Pls. 124 (1920). Diplolegnon Riceanum Rusby in Bull. Torr. Bot. Club, 27: 30 (1900). Drymonia Campbellii Rusby in Mem. N. Y. Bot. Gard. 7: 359 (1927). | D. serrata (Jacq.) Mart. Nov. Gen. & Spec. 3: 59 (1829), combination implied but not actually made. Episcia Buchtienii Mansf. in Fedde, Repert. Spec. Nov. 38: 25 (1935). Fiebrigia digitaliflora Fritsch in Engler, Bot. Jahrb. 50: 397 (1913). Fritschiantha nematanthoides O. Ktze. Rev. Gen. 3(2): 241 (1898). Gesneria stachydifolia Benth. Pl. Hartw. 230 (1846). Gloxinia perennis (L.) Fritsch in Engl. & Prantl, Nat. Pflanzenfam. 4(3b): 174 (1894). G. reflexa Rusby in Mem. Torr. Bot. Club, 6: 94 (1896). Isoloma flexuosa Rusby in Bull. Torr. Bot. Club, 27: 28 (1900). I. urticifolia Rusby in Bull. Torr. Bot. Club, 27: 28 (1900). Koellikeria argyrostigma (Hook.) Regel in Flora, 31: 250 (1848), combination implied but not actually made. K. major Fritsch in Engler, Bot. Jahrb. 50: 398 (1913). Kohleria patentipilosa (O. Ktze.) K. Schum. in Just, Bot. Jahresber. 26(1): 386 (1900). K. Sprucei (Britton) Fritsch in Engler, Bot. Jahrb. 50: 429 (1913). Kohlerianthus Fritschii (Rusby) Fritsch in Engl. & Prantl, Nat. Pflanzen- fam. Nachtr. 300 (1897). Monopyle divaricata Rusby in Bull. N. Y. Bot. Gard. 8: 119 (1912). Napeanthus andinus Rusby in Mem. Torr. Bot. Club, 6: 98 (1896). N. rigidus Rusby in Mem. Torr. Bot. Club, 6: 99 (1896). Nautilocalyx Whitei Rusby in Mem. N. Y. Bot. Gard. 7: 358 (1927). Rechsteinera ignea (Mart.) Fritsch in Engler, Bot. Jahrb. 50: 486 (1913). R. microphylla Fritsch in Meded. Rijks Herbar. 29: 53 (1916). R. suleata (Rusby) Fritsch in Engler, Bot. Jahrb. 50: 436 (1913). Seemannia albescens (Rusby) Fritsch in Engler, yes Jahrb. 50: 403 (1913). S. cuneata Rusby in Mem. Torr. Bot. Club, 6: 96 (1896). S. dioica Rusby in Mem. N. Y. Bot. Gard. 7: 360 (1927). S. latifolia Fritsch in Meded. Rijks Herbar. 29: 52 (1916). major Baill. in Bull. Mens. Soc. Linn. Paris, 1: 710 (1887). purpurascens Rusby in Mem. Torr. Bot. Club, a 237 (1895). . Silvatica (HBK.) Hanst. in Linnaea, 29: 540 (1857-58). ternifolia Regel in Gartenfl. 4: 183, t. 126 (1855). uniflora Baill. in Bull. Mens. Soc. Linn. Paris, 1: 710 (1887). nnn tn ta COLUMELLIACEAE pel sthecre excelsa Rusby in Mem. Torr. Bot. Club, 6: 17 (1896) — Columellia rata. Coluiiatite serrata Rusby in Mem. Torr. Bot. Club, 6: 93 (1896). C. subsessilis Schltr. in Notizbl. 7: 12 [856]. (1920). LENTIBULARIACEAE Pinguicula antarctica Vahl, Enum. 1: 192 (1804) Utricularia alpina Jacq. Enum. Pl. Carib. 11 (1760). cornuta Michx. Fl. Bor.-Am. 1: 12 (1803). globulariaefolia Mart. ex Benj. in Mart. Fl. Bras. 10: 241 (1847). Herzogii Liitzelberg in Fedde, Repert. Spec. Nov. 7: 356 (1909). obtusa Sw. Prodr. 14 (1788). pusilla Vahl, Enum. 1: 202 (1804). subulata L. Sp. Pl. 18 (1753). unifolia R. & P. Fl. Peruv. 1: 20, t. 31 (1798). velascoénsis O. Ktze. Rev. Gen. 3(2): 240 (1898). qaaqceaagce ACANTHACEAE Aphelandra acutifolia Nees in DC. Prodr. 11: 299 (1847). A. albadenia Rusby in Mem. N. Y. Bot. Gard. 7: 364 (1927). castanaefolia Britton ex Rusby in Bull. Torr. Bot. Club, 27: 76 (1900). cryptantha Rusby in Mem. N. Y. Bot. Gard. 7: 364 (1927). Hieronymi Griseb. in Goett. Abh. 24: 260 (1879). inaequalis Lindau in Bull. Herb. Boiss. (ser. 1) 3: 368 (1895). kolobantha Lindau in Ann. Conserv. & Jard. Bot. Genéve, 2: 39 (1898). longibracteolata Lindau in Bull. Herb. Boiss. (ser. 1) 3: 367 (1895). macrosiphon Lindau in Bull. Herb. Boiss. (ser. 1) 3: 367 (1895). Rusbyi Britton ex Rusby in Bull. Torr. Bot. Club, 27: 77 (1900). simplex Lindau in Bull. Herb. Boiss. (ser. 1) 3: 366 (1895). plied (Vahl) Nees in DC. Prodr. 11: 295 (1847). eloperone Amherstiae Nees ex Wall. Pl. As. Rar. 3: 102 (1832). Besnitell H. Winkl. in Fedde, Repert. Spec. Nov. 7: 113 (1909). Bangii Rusby in Mem. Torr. Bot. Club, 6: 104 (1896). cochabambensis Rusby in Mem. Torr. Bot. Club, 6: 103 es consanguinea Lindau in Bull. Herb. Boiss. (ser. 1) 5: 676 (18 cg en ae var. pubescens Lindau in Bull. Herb. Boiss. rts 1) 5: 676 (189 sack Nees in DC. Prodr. 11: 423 (1847). Mandonii Lindau in Bull. Herb. Boiss. (ser. 1) 5: 675 (1897). Matthewsii Lindau in Bull. Herb. Boiss. 6, App. 1: 30 (1898). nuda Rusby in Mem. Torr. Bot. Club, 6: 103 (1896). pseudociliata Mildbr. in Notizbl. 9: 1159 (1927). ramulosa Morong in Ann. N. Y. Acad. Sci. 7: 194 (1893). tetramerioides Lindau in Bull. Herb. Boiss. (ser. 1) 3: 488 (1895). velascana Lindau in Bull. Herb. Boiss. (ser. 1) 3: 489 (1895). viridissima Rusby in Mem. N. Y. Bot. Gard. 7: 367 (1927). Chaetochlamys Lindavii Rusby in Bull. N. Y. Bot. Gard. 4: 429 (1907). C. macrosiphon Lindau in Bull. Herb. Boiss. (ser. 1) 3: 490 (1895). C. Rusbyi Lindau in Bull. Herb. Boiss. (ser. 1) 3: 491 (1895). Chaetothylax boliviensis Lindau in Bull. Herb. Boiss. (ser. 1) 3: 492 (1895). C. tocantinus Nees in Mart. Fl. Bras. 9: 153 (1847). Dianthera graminifolium Rusby in Mem. N. Y. Bot. Gard. 7: 366 (1927). D. multicaule Rusby in Mem. Torr. Bot. Club, 6: 105 (1896). . resupinatum (Juss.) O. Ktze. Rev. Gen. 3(2): 248 (1898). scutellatum var. coeruleum O. Ktze. Rev. Gen. 3(2): 248 (1898). scutellatum var. flavum O. Ktze. Rev. Gen. 3(2): 248 (1898). Dicliptera cochabambensis Lindau in Bull. Herb. Boiss. (ser. 1) 3: 479 (1895). D. Niederleiniana Lindau in Engler, Bot. Jahrb. 19, Beibl. 48: 15 (1894). Ecbolium Lindavianum O. Ktze. Rev. Gen. 3(2): 248 (1898). E. minimiflorum O. Ktze. Rev. Gen. 3(2): 248 (1898). FeevEreense . DROW WWW Pah h DOD 190 E. oreadum (S. Moore) O. Ktze. Rev. Gen. 3(2): 248 (1898). Elytraria imbricata (Vahl) Pers. Synops. Pl. 1: 23 (1805). Eranthemum cordatum Nees ex Benth. Bot. Voy. Sulphur, 147 (1846). Geissomeria cincinnata Nees in Mart. Fl. Bras. 9: 81 (1847). Habracanthus pyramidalis Lindau in Bull. Herb. Boiss. (ser. 1) 3: 482 (1895). H. sanguinalis Nees in DC. Prodr. 11: 312 (1847). Hansteinia crenulata Britton ex Rusby in Mem. Torr. Bot. Club, 4(3): 242 1895). Jacobinia glabribracteata Lindau in Bull. Herb. Boiss. (ser. 1) 3: 486 (1895). J. Rusbyi Britton ex Rusby in Bull. Torr. Bot. Club, 27: 79 (1900) J. tenuistachys Rusby in Mem. Torr. Bot. Club, 6: 105 (1896). Justicia boliviana Rusby in Mem. Torr. Bot. Club, 6: 104 (1896). comata (L.) Sw. Obs. 14 (1791). Kuntzei Lindau in Bull. Herb. Boiss. (ser. 1) 3: 483 (1895). laeta (Nees) Lindau in Engl. & Prantl, Nat. Pflanzenfam. 4(3b): 350 (1895). longiacuminata Rusby in Bull. Torr. Bot. Club, 27: 78 (1900). Lorentziana Lindau in Engler, Bot. Jahrb. 19, Beibl. 48: 20 (1894). obtusifolia (Nees) Lindau in Engler, Bot. Jahrb. 19, Beibl. 48: 20 (1894). parviflora (Nees) Lindau in Engl. & Prantl, Nat. Pflanzenfam. 4(3b): 350 5). pg ts reisensis Rusby in Bull. Torr. Bot. Club, 27: 79 (1900). riojana Lindau in Engler, Bot. Jahrb. 19, Beibl. 48: 19 (1894). robusta Rusby in Bull. N. Y. Bot. Gard. 4: 480 (1907). Rusbyana Lindau in Mem. Torr. Bot. Club, 4(3): 248 (1895). subintegrifolia Rusby in Bull. N. Y. Bot. Gard. 4: 430 (1907). velascana Lindau in Bull. Herb. Boiss. (ser. 1) 3: 484 (1895). Lepidagathis justicioides Britton ex Rusby in Bull. Torr. Bot. Club, OT: 76 (1900). Sette tars Lophostachys conferta Rusby in Mem. Torr. Bot. Club, 6: 103 (1896). ? Mendoncia glabra Nees in DC. Prodr. 11: 52 (1847 M. hirsuta Poepp. & Endl. Nov. Gen. 3: 10 (1845). M. Lindavii Rusby in Mem. Torr. Bot. Club, 4(3): 241 (1895). M. puberula var. micropus (Mart.) Nees in DC. Prodr. 11: 53 (1847). M. robusta Rusby in Mem. N, Y. Bot. Gard. 7: 361 (1927). M. robusta var. alba Rusby in Mem. N. Y. Bot. Gard. 7: 362 (1927). M. Schomburgkiana Nees in DC. Prodr. 11: 50 (1847). M. Velloziana Nees in DC. Prodr. 11: 52 (1847). Pachystachys Riedeliana Nees in Mart. Fl. Bras. 9: 99 (1847). enoortre ry bolivianum Britton ex Rusby in Bull. Torr. Bot. Club, 27: 77 Ruellia amoena Nees ex Rusby in Mem. Torr. Bot. Club, 6: 102 (1896). bahiensis (Nees) Morong in Ann. N. Y. Acad. Sci. 7: 192 (1893). Bangii Rusby in Mem. Torr. Bot. Club, 6: 102 (1896). chiquitensis Baill. in Bull. Mens. Soc. Linn. Paris, 2 853 (1890). elliptica Rusby in Bull. Torr. Bot. Club, 27: 74 (19 euantha Lindau in Bull. Herb. Boiss. (ser. 1) 3: ate (1895). filicalyx Lindau in Bull. Herb. Boiss. (Ser. 1) 3: 362 (1895). geminiflora HBK. Nov. Gen. & Spec. 2: 240 (1817). gracilis Rusby in Mem. Torr. Bot. Club, 6: 102 (1896). Herzogii Lindau in Fedde, Repert. Spec. Nov. 7: 67 (1909). Humboldtiana (Nees) Lindau in Bull. Herb. Boiss. (ser. 1) 3: 366 (1895). hypericifolia Rusby in Mem. N. Y. Bot. Gard. 7: 363 (1927). Kuntzei Lindau in Bull. Herb. Boiss. (ser. 1) 3: 365 (1895). Lechleri Britton ex Rusby in Bull. Torr. Bot. Club, 27: 75 (1900). Lechleri var. grandifolia Britton ex Rusby in Bull. Torr. Bot. Club, 27: 75 (1900). mi PO Py Pd Pd md Pd po ad bd oo po Ba 191 longipedunculata Lindau in Bull. Herb. Boiss. (ser.1) 3: 365 (1895). Lorentziana Griseb. in Goett. Abh. 24: 259 (1879). Morongii Britton in Ann. N. Y. Acad. Sci. 7: 192 (1893). multisetosa Rusby in Mem. N. Y. Bot. Gard. 7: 362 (1927). paniculata L. Sp. Pl. 685 (1753). Pearcei Rusby in Bull. N. Y. Bot. Gard. 4: 429 (1907). pedunculosa (Nees) Lindau in Engl. & Prantl, Nat. Pflanzenfam. 4(3b): 311 (1895). proxima Lindau in Bull. Herb. Boiss. (ser. 1) 3: 365 (1895). Puri Mart. ex Nees in Mart. Fl. Bras. 9: 35 (1847), in synon. Ruiziana (Nees) Lindau in Engl. & Prantl, Nat. Pflanzenfam. 4(3b): 311 95). sanguinea Griseb. in Goett. Abh. 24: 260 (1879). serratitheca Rusby in Mem. N. Y. Bot. Gard. 7: 362 (1927). tuberosa L. Sp. Pl. 635 (1753). velascana Lindau in Bull. Herb. Boiss. (ser. 1) 3: 363 (1895). Willdenowiana (Nees) Lindau ex Rusby in Bull. N. Y. Bot. Gard. 4: 429 (1907). Sanchezia peruviana (DC.) Rusby in Mem. Torr. Bot. Club, 6: 103 (1896). Schaueria azaleaeflora Rusby in Mem. N. Y. Bot. Gard. 7: 365 (1927). Staurogyne diantheroides Lindau in Bull. Herb. Boiss. (ser. 1) 5: 645 (1897). Stenandrium diphyllum Nees in Mart. Fl. Bras. 9: 75 (1847). S. dulce (Cav.) Nees in DC. Prodr. 11: 282 (1847). S. mandioccanum Nees in Mart. Fl. Bras. 9: 76 (1847). S. trinerve Nees in Mart. Fl. Bras. 9: 75 (1847). Stenostephanus bolivianus Rusby in Mem. N. Y. Bot. Gard. 7: 366 (1927). Streblacanthus boliviensis Lindau in Fedde, Repert. Spec. Nov. 7: 67 (1909). Suessenguthia trochilophila Merxm. in Mitt. Bot. Staatssamml. Miinch. 6: 178 1953 FARRAR AR AR BB Tacoanthus Pearcei Baill. in Bull. Mens. Soc. Linn. Paris, 2: 832 (1890). PLANTAGINACEAE Bougueria nubicola Decne. in Ann. Sci. Nat. (ser. 2) 5: 133 (1836). Plantago Asplundii Pilger in Engler, Bot. Jahrb. 62: 27 (1928). Buchtienii Pilger in Engler, Bot. Jahrb. 50: 251 (1913). Durvillei subsp. mollior (Pilg.) Pilger in Engler, Pflanzenr. IV, 269: 234 (1937). Durvillei subsp. Pflanzii (Pilg.) Pilger in Engler, Pflanzenr. IV, 269: 232 (1937). Durvillei subsp. Pflanzii var. chamaeclina (Pilg.) Pilger in Engler, Pflan- zenr. IV, 269: 2382 (1937). Durvillei subsp. Pflanzii var. grandidens (Pilg.) Pilger in Engler, Pflan- zenr. IV, 269: 233 (1937). Durvillei subsp. Pflanzii var. Hauthalii (Pilg.) Pilger in Engler, Pflanzenr. IV, 269: 233 (19387). Durvillei subsp. Pflanzii var. latifolia (Pilg.) Pilger in Engler, Pflanzenr. IV, 269: 233 (1937). Fiebrigii Pilger in Fedde, Repert. Spec. Nov. 15: 420 (1919). Hillii Pilger in Notizbl. 10: 833- (1929). hirtella HBK. Nov. Gen. & Spec. 2: 229, t. 127 (1818). hirtella forma minor Pilger in Engler, Bot. Jahrb. 50: 276 (1913). humilior Pilger in Field Mus. Publ. Bot. 13(6): 271 (1937). lanceolata L. Sp. Pl. 113 (1753). litorea Phil. Fl. Atac. 46 (1860). major L. Sp. Pl. 112 (1753). Po re ae oe Oem om me we 192 major forma sinuata (Lam.) Pilger in Fedde, Repert. Spec. Nov. 18: 271 (1922). monticola Decne. in DC. Prodr. 13(1): 711 (1852). monticola subsp. angusta Pilger in Engler, Bot. Jahrb. 62: 80 (1928). monticola subsp. crispula Pilger in Engler, Bot. Jahrb. 62: 81 (1928). monticola subsp. eumonticola var. humillima Pilger in Engler, Bot. Jahrb. 62: 80 (1928). monticola subsp. sericans Pilger in Engler, Bot. Jahrb. 62: 80 (1928). Orbignyana Steinh. ex Decne. in DC. Prodr. 13 (1): 704 (1852). paralias subsp. affinis (Decne.) Pilger in Notizbl. 11: 328 (1932). paralias subsp. Grisebachii (Hieron.) Pilger in Notizbl. 11: 328 (1932). Psyllium L. Sp. Pl. 115 (1753). rigida HBK. Nov. Gen. & Spec. 2: 227, t. 126, fig. 2 (1818). sericea R. & P. Fl. Peruv. 1: 51, t. 79, fig. b (1798). tarijensis Pilger in Fedde, Repert. pe Nov. 15: 420 (1919). truncata Barn. Monog. Plantag. 14 (1845). tubulosa Decne. in DC. Prodr. 13(1): ie (1852). tubulosa forma maxima Pilger in Engler, Bot. Jahrb. 62: 92 (1928). virginica L. Sp. Pl. 118 (1753). Weddelliana Decne. in DC. Prodr. 13(1): 712 (1852). SUN DeV I ee V Y RUBIACEAE Alibertia benensis Standl. in Ficld Mus. Publ. Bot. 7: 291 (1931). A. Steinbachii Standl. in Field Mus. Publ. Bot. 4: 287 (1929). A. Tutumilla Rusby in Mem. N. Y. Bot. Gard. 7: 375 (1927). Amaioua corymbosa HBK. Nov. Gen. & Spec. 3: 419, t. 294 (1820). Anisomeris albicaulis (Rusby) Standl. in Field ‘Mus. Publ. Bot. 4: 293 (1929). A. apodantha Standl. in Field Mus. Publ. Bot. 4: 293 (1929). A. boliviana (Standl.) Rusby in Bull. Torr. Bot. Club, 52: 142 (1925). A. rauwolfioides Standl. in Field Mus. Publ. Bot. 4: 292 (1929). Arcytophyllum filiforme (R. & P.) Standl. in Field Mus. Publ. Bot. 11: 183 (1936). A. nodosum Rusby in Phytologia, 1: 78 (1934). A. setosum (R. & P.) Schlechtd. in Linnaea, 28: 492 (1856). Basanacantha erythropoda Rusby in Mem, N. Y. Bot. Gard. 7: 873 (1927) = Randia spinosa. carpa Rusby in Mem. N. Y. Bot. Gard. 7: 374 (1927) = Randia formosa. B. mollis Rusby in Mem. N. Y. Bot. Gard. 7: 375 (1927) = Randia formosa. B. mucronata Rusby in Mem. N. Y. Bot. Gard. 7: 373 (1927) —= Randia spinosa. Bathysa obovata Schum. ex Standl. in Field Mus. Publ. Bot. 7: 280 (1931). Bertiera guianensis Aubl. Pl. Guian. Fr. 1: 180, t. 69 (1775). Borreria Brownii (Rusby) Standl. in Field Mus. Publ. Bot. 7: 333 (1931). capitata (R. & P.) DC. Prodr. 4: 545 (1830). B. corymbosa (R. & P.) DC. Prodr. 4: 550 (1830). densiflora var. perennis Standl. in Field Mus. Publ. Bot. 7: 334 (1931). exigua S. Moore in Meded. Rijks Herbar. 46: 28 (1922). Herzogii S. Moore in Meded. Rijks Herbar. 46: 26 (1922). laevis (Lam.) Griseb. in Goett. Abh. 7: 231 (1857). latifolia (Aubl.) K. Schum. in Mart. Fl. Bras. 6(6): 61 (1888). mectarifera Rusby in Mem, N. Y. Bot. Gard. 7: 381 (1927) = B. capitata. ocimoides (Burm.) DC. Prodr. 4: 544 (1830). staurochlamys R, E. Fries in Ark. Bot. 6(11): 9, t. 2 (1907) — B. Brownii. B. suaveolens Meyer, Prim. FI. Esseq. 81, t. 1 (1818). Calycophyllum multifiorum Griseb. in Goett. Abh. 24: 155 (1879). So oy Bd by bd bY be te by 193 C. Spruceanum (Benth.) Hook.f. ex K. Schum. in Mart. Fl. Bras. 6(6): 191, t. 106 (1889). Cephaélis conephoroides Rusby in Bull. N. Y. Bot. Gard. 4: 372 (1907); == C. umbellata. C. tomentosa (Aubl.) Vahl, Eclog. Amer. 1: 19 (1796). C. umbellata (R. & P.) Standl. in Field Mus. Publ. Bot. 7: 301 (1931). Chiococea alba (L.) A. S. Hitche. in Rept. Missouri Bot. Gard. 4: 94 (1893). Chomelia brevicornu Rusby in Bull. Torr. Bot. Club, 52: 140 (1925). C. dimorpha Rusby in Bull. Torr. Bot. Club, 52: 140 (1925). C. multiflora Rusby in Bull. Torr. Bot. Club, 52: 141 (1925). C. unguis-cati Standl. in Field Mus. Publ. Bot. 8: 172 (1930). masa ey ee Wedd. Hist. Nat. Quing. 100 (1849) == Pogonopus tubulos Cinchona snaatola Wedd. in Ann. Sci. Nat. (ser. 7 ay 7 (1848). C. calisaya Wedd. in Ann. Sci. Nat. (ser. 3) 10: 6 (18 C. Humboldtiana Lamb. Ill. Cinch. 7 (1821). C. Josephiana Wedd. in Ann. Sci. Nat. (ser. 5) 12: 58 (1869). C. micrantha R. & P. Fl. Peruv. 2: 52, t. 194 (1799). C. officinalis L. Sp. Pl. 172 (1753). C. ovata R. & P. Fl. Peruv. 2: 52, t. 195 (17 C. pubescens Vahl in Skrivt. Naturh. Selsk. - 19 (1790). Coccocypselum Brittonii Rusby in Bull. N. Y. Bot. Gard. 4: 368 (1907) = C. Condalia. C. Condalia Pers. Synops. Pl. 1: 132 (1805). C. cordatum Krause in Anex. Mem. Inst. Butantan, 1(3): 13, t. 1 (1922). C. glabrum Britton in Bull. Torr. Bot. wien 18: 108 (1891) — C. Condalia. C. hirsutum Bartl. ex DC. Prodr. 4: 396 (1830). C. lanceolatum (R. & P.) Pers. Synops. Pl. 1: 132 (1805). Coffea arabica L. Sp. Pl. 172 (1753). Condaminea angustifolia ie in Mem. Torr. Bot. Club, 6: 45 (1896) = C. corymbosa var. pubescen C. corymbosa (R. & P.) DC. mai 4; 402 (1830). C. corymbosa var. pubescens Spruce ex K. Schum, in Mart. Fl. Bras. 6(6): 258 (1889). Corynula pilosa (Benth.) Hook. f. in Hook. Ic. Pl. 18:22, t. _ (1872). Cosmibuena grandifiora (R. & P.) Rusby in Bull. N. . a . Gard. 4: 368 (1907). Coussarea auriculata Standl. in Field Mus. Publ. Bot. 4: 330 (1929). benensis Britton ex Standl. in Field Mus. Publ. Bot. 7: 296 (1931) = C. brevicaulis. brevicaulis Krause in Verh. Bot. Ver. Brandenb. 50: 117 (1908). . hydrangeaefolia (Benth.) Benth. & Hook. f. ex Muell. Arg. in Mart. FI. Bras. 6(5): 94 (1881). rudgeoides Rusby in Mem. Torr. Bot. Club, 6: 48 — urophylla Standl. in Field Mus. Publ. Bot. 7: 298 (1981 Diodia hyssopifolia (Willd.) Cham. & Schlechtd. in Linsaee: 8: 350 (1828), combination implied but not actually made. D. Kuntzei K. Schum. in Mart. Fl. Bras. 6(6): 15 (1888). D. rogaguana Rusby in Mem. N. Y. Bot. Gard. 7: 380 (1927). Duroia Steinbachii Standl. in Field Mus. Publ. Bot. 7: 290 (1931). Elaeagia glomiflora Standl. in Field Mus. Publ. Bot. 7: 281 (1931). E. grandis (Rusby) Rusby in Mem. Torr. Bot. Club, 4(3): 208 (1895). E. Mariae Wedd. Hist. Nat. Quing. 94 (1849). E. mollis Rusby, Descr. S. Am. Pls. 130 (1920). E. obovata Rusby, Descr. S. Am. Pls. 130 (1920). E. utilis (Goud.) Wedd. Hist. Nat. Quing. 94 (1849). Jam aa A 194 Emmeorrhiza umbellata (Spreng.) K. Schum. in Mart. Fl. Bras. 6(6): 408 (1889). Evea radiata Rusby in Phytologia, 1: 79 (1934). Faramea anisocalyx Poepp. & Endl. Nov. Gen. 3: 28 (1845). Bangii Rusby in Mem. Torr. Bot. Club, 3(3): 45 (1893). enensis Rusby in Mem. N., Y. Bot. Gard, 7: 379 (1927) — F. maynensis. candelabrum Standl. in Field Mus. Publ. Bot. 7: 299 (1931). glandulosa Poepp. & Endl. Nov. Gen. 3: 29 (1845). maynensis Spruce ex Rusby in Bull. N. Y. Bot. Gard. 4: 370 (1907). tenuifolia Rusby in Mem. N. Y. Bot. Gard. 7: 380 (1927 Ferdinandusa Paxii H. Winkl. in Fedde, Repert. Spec. Nov. 7: 249 (1909). Galium Aparine L. Sp. Pl. 108 (1753). canescens HBK. Nov. Gen. & Spec. 3: 336 (1819). charoides Rusby in Phytologia, 1: 80 (1934). cochabambense Rusby in Mem. Torr. Bot. Club, 6: 52 (1896) = G. plu- mosum. larecajense Wernh. in Journ. Bot. 50: 244 (1912). Mandonii Britton in Bull. Torr. Bot. Club, 18: 263 (1891). obovatum HBK. Nov. Gen. & Spec. 2: 336, t. 227 (1819). plumosum Rusby in Mem. Torr. Bot. Ciub, 3(3): 49 (1893). Genipa americana L. Syst. (ed. 10) 931 (1759). G. Caruto HBK. Nov. Gen. & Spec. 3: 407 (1820) . Geophila herbacea (L.) Morong in Ann. N. Y. Acad. Sci. 7: 129 (1893). Gomozia granadensis L. f. Suppl. 129 (1781). Gonzalagunia Whitei (Rusby) Standl. in Field Mus. Publ. Bot. 4: 280 (1929). Guettarda boliviana Standl. in Field Mus. Publ. Bot. 4: 289 (1929). G. Spruceana Muell. Arg. in Flora, 58: 449 (1875). G. Tournefortiopsis Standl. in Field Mus. Publ. Bot. 7: 293 (1931). Hamelia lutea Rohr ex Smith in Rees, Cycl. 17: no. 4 (1819). H. patens Jacq. Enum. Pl. Carib. 16 (1760). H. pedicellata Wernh. in Journ. Bot. 49: 212 (1911). Hemidiodia ocimifolia (Willd.) K. Schum. in Mart. Fl. Bras. 6(6): 29 (1888). pasar age Fiebrigii (Krause) Standl. in Field Mus. Publ. Bot. 11: 214 PROP DOE H. yeas (Rusby) Sandw. in Kew Bull. 1949: 256. ta Hook. f. in Hook. Ic. Pl. 12: 31, t. 1184 (1878). Hillia rare Britton in Mem. Torr. Bot. Club, 3(3): 43 (1893). H. parasitica Jacq. Enum. Pl. Carib. 18 (1760), nomen nudum. H. Ulei Krause in Verh. Bot. Ver. Brandenb. 50: 97 (1908). Hoffmannia brachycarpa Britton in Bull. Torr. Bot. Club, 18: 108 (1891) = H. latifolia. H. latifolia (Bartl.) O. Ktze. Rev. Gen. 1: 284 (1891). H. pallida Rusby in Mem. Torr. Bot. Club, 3(3): 44 (1898). H. Pearcei Rusby in Mem. Torr. Bot. Club, 3(3): 44 (1898). Isertia reticulata Britton in Mem. Torr. Bot. Club, 6: 46 (1896). Ixora Killipii Stand]. in Field Mus. Publ. Bot. 8: 174 (1930). I. peruviana (Spruce) Standl. in Field Mus. Publ. Bot. 7: 296 (1931). Ladenburgia Carua (Wedd.) Standl. in Field Mus. Publ. Bot. 7: 274 (1931). L, magnifolia (R. & P.) Klotzsch in Hayne, Arzneigew. 14: sub t. 15 (1856). L. magnifolia var. rostrata (Wedd.) Standl. in Field Mus. Publ. Bot. 7: 275 (1981.) L. Riveroana (Wedd.) Standl. in Field Mus. Publ. Bot. 7: 201 (1931). L. sericea Standl. in Field Mus. Publ. Bot. 7: 275 (1931). Limnosipanea Kuntzei Standl. in Field Mus. Publ. Bot. 7: 280 (19381). M cnemum hirsutum Rusby in Mem. N. Y. Bot. Gard. 7: 370 (1927) = M. roseum. 195 M. roseum (R. & P.) Wedd. in Ann. Sci. Nat. (ser. 4) 1: 76 (1854). M. Sprucei Rusby in Mem. Torr. Bot. Club, 6: 44 (1896) = M. roseum. M. tortuosum Herzog in Fedde, Repert. Spec. Nov. 7: 68 (1909). Malanea boliviana Standl. in Field Mus. Publ. Bot. 7: 295 (1931). Manettia asperifolia Standl. in Field Mus. Publ. Bot. 4: 270 (1929). Bangii (Rusby) Standl. in Field Mus. Publ. Bot. 4: 271 (1929). boliviana Wernh. in Journ. Bot. 57, Suppl.: 20 (1919) = M. cordifolia var. glabra. coccinea (Aubl.) Willd. Sp. Pl. 1(2): 625 (1798). cordifolia Mart. Spec. Mat. Med. Bras. 19, t. 7 (1824). cordifolia var. glabra (Cham. & Schlechtd.) Standl. in Field Mus. Publ. Bot. 7: 263 (1931). divaricata Wernh. in Journ. Bot. 57, Suppl.: 41 (1919). Fiebrigii Stand]. in Field Mus. Publ. Bot. 7: 264 (1931). hispida Poepp. & Endl. Nov. Gen. 3: 24, t. 228 (1845). Pearcei Wernh. in Journ. Bot. 57, Suppl.: 36 (1919). Tatei Standl. in Field Mus. Publ. Bot, 7: 265 (1981). tenuis (Britton) Wernh. in Journ. Bot. 57, Suppl.: 33 (1919). tomentosa (Rusby) Standl. in Field Mus. Publ. Bot. 7: 265 (1931). Mapouria rigida Rusby in Mem. N. Y. Bot. Gard. 7: 376 (1927) = Psychotria a S 5 SSSS555 SES Iba. Mitracarpus frigidus (Willd.) K. Schum. in Mart. Fl. Bras. 6(6): 81, t. 85 ). M. hirtus (L.) DC. Prodr. 4: 572 (1830). M. megapotamicus (Spreng.) Standl. in Field Mus. Publ. Bot. 7: 331 (1931). M. simplex Rusby in Mem. N. Y. Bot. Gard. 7: 381 (1927) = M. hirtus. Paederia diffusa (Britton) Standl. in Field Mus, Publ. Bot. 7: 330 (1931). Palicourea amethystina (R. & P.) DC. Prodr. 4: 527 (1830). attenuata Rusby in Bull. N. Y. Bot. Gard. 4: 371 (1907). Buchtienii Standl. in Field Mus. Publ. Bot. 7: 317 (1981). Cardenasii Standl. in Field Mus. Publ. Bot. 22: 119 (1940). cornifolia Standl. in Field Mus. Publ. Bot. 7: 318 (19381). corymbifera (Muell. Arg.) Standl. in Field Mus. Publ. Bot. 7: 127 (1930). crocea (Sw.) Roem. & Schult. Syst. Veg. 5: 193 (1819). exiguiflora Standl. in Field Mus. Publ. Bot. 7: 819 (1931). fastigiata HBK. Nov. Gen. & Spec. 3: 368 (1819). flavifolia (Rusby) Standl. in Field Mus. Publ. Bot. 7: 320 (1931). glabrata H. Winkl. in Fedde, Repert. Spec. Nov. 8: 3 (1910). Herzogii Standl. in Field Mus. Publ. Bot. 11: 226 (1936). lasiantha Krause in Engler, Bot. Jahrb. 40: 341 (1908). longipes Rusby in Bull. N. Y. Bot. Gard. 8: 121 (1912) = P. triphylla. macrobotrys (R. & P.) DC. Prodr. 4: 527 ( 1830). macrophylla (HBK.) Standl. in Field Mus. Publ. Bot. 7: 321 (1931). malacophylla Standl. in Field Mus. Publ. Bot. 7: 322 (1931). mapirensis Standl. in Field Mus. Publ. Bot. 7: 822 (1931). mollis H. Winkl. in Fedde, Repert. Spec. Nov. 8: 4 (1910) = P triphylla. obliqua H. Winkl. in Fedde, Repert. Spec. Nov. 8: 4 (1910) = P. ovalifolia. ovalifolia (Rusby) Standl. in Field Mus. Publ. Bot. 7: 323 (1931). papyracea Rusby in Bull. N. Y. Bot. Gard. 4: 370 (1907). Pearcei Standl. in Field Mus. Publ. Bot. 7: 324 (1931). punicea (R. & P.) DC. Prodr. 4: 526 (1830). radicans Standl. in Field Mus. Publ. Bot. 7: 325 (1931). rigida HBK. Nov. Gen. & Spec. 3: 370 (1819). stipularis Benth. Pl. Hartw. 133 (1844). . tenuis Stand]. in Field Mus. Publ. Bot. 7: 326 (1931). . triphylla DC. Prodr. 4: 526 (1830). . we we oe ee ee Meee 0 oY OD NY eee ee 196 P. verrucifera Standl. in Field Mus. Publ. Bot. 7: 327 (1931). P. Williamsii Rusby, Descr. S. Am. Pls. 142 (1920) = P. macrobotrys. Pogonopus febrifugus var. macrosema Hutchinson in Kew Bull. 1910: 200 = P. tubulosus. P. tubulosus (DC.) K. Schum. in Mart. Fl. Bras. 6(6): 265 (1889). Posoqueria latifolia (Rudge) Roem. & Schult. Syst. Veg. 5: 227 (1819). Psychotria alba R. & P. Fl. Peruv. 2: 58, t. 205, fig. a (1799). axillaris Willd. Sp. Pl. 1(2): 962 (1798). Bangii Rusby in Mem. Torr. Bot. Club, 3(3): 47 (1893). Bangii Romero in Caldasia, 7: 50 (1955) = P. salicifolia Rusby — Pali- courea attenuata Rusby. boliviana Standl. in Field nie Publ. Bot. 7: 308 (1981). brachiata Sw. Prodr. 45 (1788). Buchtienii (H. Winkl.) Standl. in Field Mus. Publ. Bot. 7: 303 (1931). capitata R. & P. Fl. Peruv. 2: 59 (1799). Cardenasii Standl. in Field Mus. Publ. Bot. 7: 303 (1931). carthagenensis Jacq. Enum. Pl. Carib. 16 (1760). chlorotica Muell. Arg. in Flora, 59: 542 (1876) = P. capitata. costata (Rusby) Standl. in Field Mus. Publ. Bot. 7: 305 (1931). cuspidata Bredem. ex Roem. & Schult. Syst. Veg. 5: 192 (1819). emetica L. f. Suppl. 144 (1781). falcata Rusby in Mem. Torr. Bot. Club, 3(3): 47 (1893). Hartwegiana Standl. in Field Mus. Publ. Bot. 7: 95 (1930). Herzogii S. Moore in Meded. Rijks Herbar. 46: 25 (1922). involucrata (Aubl.) Sw. Prodr. 45 (1788). lassula Standl. in Field Mus. Publ. Bot. 7: 100 (1930). lupulina Benth. in Hook. Journ. Bot. 3: 230 (1841). luxurians Rusby in Mem. Torr. Bot. Club, 6: 50 (1896). macrophylla R. & P. Fl. Peruv. 2: 56, t. 202, fig. a (1799). mapirensis Standl. in Field Mus. Publ. Bot. 7: 308 (1931). marginata Sw. Prodr. 43 (1788). microbotrys Ruiz ex Standl. in Field Mus. Publ. Bot. 8: 204 (1930). nana Krause in Verh. Bot. Ver. Brandenb. 50: 109 (1908). niveo-barbata (Muell. Arg.) Britton in Bull. Torr. Bot. Club, 18: 110 (1891). olyphylla Rusby, Deser. S. Am. Pls. 188 (1920) = ae triphylla. Ottonis Standl. in Field Mus. Publ. Bot. 7: 310 (193 pallescens (Rusby) ssa in Field Mus. Publ. Bot. ny "310 (1931). patens Sw. Prodr. 45 (17 Pearcei Standl. in Field ie. Publ. Bot. 7: 311 (1931). pervicax Standl. in Field Mus. Publ. Bot. 7: 311 (1931). pilosa R. & P. Fl. Peruv. 2: 50, t. 208, fig. a (1799). racemosa (Aubl.) Willd. Sp. Pl. 1: 966 (1797). ramiflora Rusby in Mem. N. Y. Bot. Gard. 7: 377 (1927). reticulata R. & P. Fl. Peruv. 2: 56, t. 202, fig. b (1799). rhodothamna Standl. in Field Mus. Publ. Bot. 8: 210 (1930). igida var. brevipes Rusby in Mem. Torr. Bot. Club, 6: 50 (1896) = Pali- courea papyracea Ruizii Stand. in Field Mus. Publ. Bot. 7: 313 (1931). salicifolia Rusby, Deser. S. Am. Pls. 140 (1920) = Palicourea attenuata. scabrifolia Rusby, Descr. S. Am. Pls. 137 (1920). Steinbachii Standl. in Field Mus. Publ. Bot. 7: 314 (1931). tipuanensis Standl. in Field Mus. Publ. Bot. 7: 315 (1931). tristis H. Winkl. in Fedde, Repert. Spec. Nov. 7: 249 (1909). trivialis Rusby in Mem. N. Y. Bot. Club, 6: 50 (1896). viburnifolia Rusby in Mem, N. Y. Bot. Gard. 7: 378 (1927) —= Rudgea to- mentosa. ~ B90 Te FO PO BP 0 tr 0 Pe Oe ari to Po or Te MUM NN ~ 197 P. viridis R. & P. Fl. Peruv. 2: 61, t. 210, fig. b (1799). P. yungasensis Rusby in Mem. Torr. Bot. Club, 3(3): 46 (1893). Randia boliviana Rusby in Bull. N. Y. Bot. Gard. 4: 368 (1907). R. calycina Cham. in Linnaea, 9: 246 (1835). R. formosa (Jacq.) K. Schum. in Mart. Fl. Bras. 6(6): 342 (1889). R. oblanceolata Rusby in Bull. N. Y. Bot. Gard. 8: 120 (1912) = R. spinosa. R. spinosa (Jacq.) Karst. Fl. Colomb. 2: 128 (1869). Relbunium alpicola K. Schum. & R. E. Fries in Nov. Act. Soc. Sci. Upsal. (ser. 4) 1(1): 101 (1905). Bangii Rusby in Bull. N. Y. Bot. Gard. 4: 374 (1907) = R. ovale. ciliatum (R. & P.) Hemsl. in Biol. Centr. Am. Bot. 2: 62 (1881). compactum Rusby in Mem. Torr. Bot. Club, 3(3): 49 (1893) = R. ciliatum. croceum (R. & P.) K. Schum. in Mart. Fl. Bras. 6(6): 116 (1888). hirsutum (R. & P.) K. Schum. in Mart. Fl. Bras. 6(6): 116 (1888). hypocarpium (L.) Hemsl. in Biol. Centr. Am. Bot. 2: 63 (1881) . nitidum (HBK.) K. Schum. in Mart, Fl. Bras. 6(6): 114 (1888). ovale (R. & P.) K. Schum. in Mart. Fl. Bras. 6(6): 115 (1888). tenuissimum Krause in Engler, Bot. Jahrb. 40: 350 (1908). vile (Cham. & Schlechtd.) K. Schum. in Mart. Fl. Bras. 6(6):.116 (1888). Richardia brasiliensis Gomez, Mem. Ipecac. 31, t. 2 (1801). R. coldenioides Rusby in Mem. Torr. Bot. Club, an at (1895). R. cruciata Rusby in Mem. Torr. Bot. Club, 6: 51 R. grandiflora (Cham. & Schlechtd.) Steud. vt cathe ted. 2) 2: 459 (1841). Rudgea acuminata (R. & P.) Standl. in Field Mus. Publ. Bot. 7: 327 (1931). R. amazonica Muell. Arg. in Flora, 59: 449 (1876). IPP AAD AANA R. Buchtienii Stand]. in Field Mus. Publ. Bot. 7: 328 (1931). R. ciliata (R. & P.) Spreng. Syst. Veg. 1: 755 (1825). R. fimbriata (Benth.) Standl. in Standl. & Calderén, List. Pl. Salvador, 274 (1925). R. tomentosa Rusby in Mem. Torr. Bot. Club, 3(3): 48 (1893). RB. Mrreg Rusby in Mem. Torr. Bot. Club, 6: 51 (1896) = Psychotria reticu- lat R. vtpaehotads (Cham. & Schlechtd.) Benth. in Linnaea, 23: 458 (1850). Sabicea acutissima Rusby in Mem. N. Y. Bot. Gard. 7: 371 (1927) = S. erecta. boliviensis Wernh. Monog. Sabicea, 37 (1914). brasiliensis Wernh. Monog. Sabicea, 51 (1914). cuneata Rusby in Mem. Torr. Bot. Club, 6: 47 (1896). erecta Rusby ex Wernh. Monog. Sabicea, 36, t. 12, fig. 23 (1914). Pearcei Wernh. Monog. Sabicea, 38, t. 3, fig. 1 (1914). setiloba Wernh. Monog. Sabicea, 37 (1914). villosa Roem. & Schult. Syst. Veg. 5: 265 (1819). villosa var. adpressa (Wernh.) Standl. in Field Mus. Publ. Bot. = 52 (1930). Sickingia catappifolia Standl. in Field Mus. Publ. Bot. 7: 279 (1931). S. fragrans (Rusby) Standl. in Field Mus. Publ. Bot. 7: 279 (1931). Sipanea hispida Benth. ex Wernh. in Journ. Bot. 55: 173 (1917 Spermacoce cephalophora Rusby in Bull. N. Y. Bot. Gard. 4: 374 oe Borreria latifolia. S. tenuior L. Sp. Pl. 102 (1753). Staelia filifolia Rusby in Mem. Torr. Bot. Club, 4(3): 209 (1895). Tocoyena formosa (Cham. & Schlechtd.) K. Schum. in Mart. Fl. Bras. 6(6): 347 (1889). Tournefortiopsis reticulata Rusby in Bull. N. Y. Bot. Gard. 4: 369 (1907) = Guettarda Tournefortiopsis. Uncaria guianensis (Aubl.) Gmel. Syst. Veg. 370 (1791). Warscewiezia coccinea (Vahl) Klotzsch in Monatsber. Akad. Berlin, 1853: 497 853) 1NAAMAN IN 198 W. splendens Wedd. in Ann. Sci. Nat. (ser. 4) 1: 72 (1854) = W. coccinea. Watsonamra sordidiflora Rusby in Mem. N. Y. Bot. Gard. 7: 372 (1927) = Palicourea lasiantha. Wernhamia boliviensis S. Moore in Meded. Rijks Herbar. 46: 23 (1922). CAPRIFOLIACEAE Sambucus peruviana HBK. Nov. Gen. & Spec. 3: 429 (1820). Viburnum ayavacense HBK. Nov. Gen. & Spec. 3: 428 (1820). ? V. glabratum HBK. Nov. Gen. & Spec. 3: 428 (1820). lasiophyllum Benth. Pl. Hartw. 189 (1845). pichinchense Benth. Pl. Hartw. 188 (1845). Seemenii Graebn. in Engler, Bot. Jahrb. 37: 435 (1906). Seemenii forma bolivianum (Gandog.) Killip & Smith in Bull. Torr. Bot. Club. 57: 258 (1930). Seemenii forma minus Killip & Smith in Bull Torr. Bot. Club, 57: 258 (1930). Spruceanum Rusby in Bull. N. Y. Bot. Gard. 8: 121 (1912). ? V. tinoides L. f. Suppl. 184 (1781). = Sta VALERIANACEAE Belonanthus angustifolius Schmale in Notizbl. 13: 25 (1936). B. crassipes (Wedd.) Graebn. in Engler, Bot. Jahrb. 37: 447 (1906). B. hispidus (Wedd.) Graebn. in Engler, Bot. Jahrb. 37: 447 (1906). Phyllactis corymbulosa Wedd. Chlor. And. 2: 34 (1858). P. densa Wedd. Chlor. And. 2: 31 (1858). P. inconspicua Wedd. Chlor. And. 2: 31 (1858). P. Mandoniana Wedd. Chlor. And. 2: 34 (1858). P. mapirensis Britton in Bull. Torr. Bot. Club, 18: 263 (1891). Valeriana andina Britton in Bull. Torr. Bot. Club, 18: 264 (1891). Bangiana Graebn. in Engler, Bot. Jahrb. 37: 437 (1906). boliviana Britton in Bull Torr. Bot. Club, 18: 263 (1891). bulbosa Wedd. Chlor. And. 2: 24 (1858). calvescens Briq. in Ann. Conserv. & Jard. Bot. Genéve, 17: 334 (1914). decussata R. & P. Fl. Peruv. 1: 42, t. 70 (1798). effusa Griseb. in Goett. Abh. 19: 162 (1874). effusa var. Fiebrigii Brig. in Ann. Conserv. & Jard. Bot. Genéve, 20: 434 (1919). glauca Poepp. ex DC. Prodr. 4: 639 (1830). hyalinorrhiza R. & P. Fl. Peruv. 1: 41, t. 67 (1798). jasminoides Briq. in Ann. Conserv. & Jard. Bot. Genéve, 17: 336 (1914). leptothyrsos Graebn. in Engler, Bot. Jahrb. 26: 425 (1899). macrorhiza Poepp. ex DC. Prodr. 4: 635 (1830). Mandonii Britton in Bull. Torr. Bot. Club, 18: 264 (1891). micropterina Wedd. Chlor. And. 2: 26, t. 49B (1858). miphobia Brig. in Ann. Conserv. & Jard. Bot. Genéve, 17: 347 (1914) = Belonanthus hispidus. nivalis Wedd. Chlor. And. 2: 238, t. 48A (1858). paniculata R. & P. Fl. Peruv. 1: 41, t. 70 (1798). Pavonii Poepp. & Endl. Nov. Gen. 3: 16, t. 215 (1845). Pavonii var. yungasensis Briq. in Ann. Conserv. & Jard. Bot. Genéve, 17: 337 (1914). polemonioides HBK. Nov. Gen. & Spec. 3: 332 (1819). polyclada Briq. in Ann. Conserv. & Jard. Bot. Genéve, 17: 343 (1914). poterioides Graebn. in Engler, Bot. Jahrb. 37: 441 (1906). potopensis Brig. in An. Conserv. & Jard. Bot. Genéve, 17: 335 (1914). psychrophila Brig. in Ann. Conserv. & Jard. Bot. Genéve, 17: 331 (1914). SSSBSMSR SSeS ao SASS 199 pygmaea Graebn. in Engler, Bot. Jahrb. 37: 446 (1906). Rusbyi Britton in Bull. Torr. Bot. Club, 18: 264 (1891). scandens L. Sp. Pl. (ed. 2) 1: 47 (1762) soratensis Brig. in Ann. Conserv. & Jard. Bot. Genéve, 17: 346 (1914). tuberifera Graebn. in Engler, Bot. Jahrb. 26: 428 (1899). <<<<< DIPSACACEAE Dipsacus fullonum L. Sp. Pl. 97 (1753). CUCURBITACEAE Anguria bignoniacea Poepp. & Endl. Nov. Gen. 2: 53 (1838). A. grandiflora Cogn. in Mém. Cour. Acad. Belg. 28: 22 (1877). A. longipedunculata Cogn. in Mém. Cour. Acad. Belg. 28: 21 (1877). A. spinulosa Poepp. & Endl. Nov. Gen. 2: 52, t. 170 (1838). A. Warszewiczii Hook. in Bot. Mag. 88: t. 5304 (1862). Apodanthera hirtella Cogn. in Engler, Pflanzenr. IV, 275(1): 61 (1916). A. Mandonii Cogn. in Mém. Cour. Acad. Belg. 28: 41 (1877). Calycophysum pedunculatum var. villosum Cogn. in Bull. Torr. Bot. Club, 23: 17(1896). Cayaponia boliviensis Cogn. in Mem. Torr. Bot. Club, 3(3): 39 (1893). C. citrullifolia var. breviloba iriti, ex Cogn. in DC. Monog. Phan. 3: 749 (1881). C. pentaphylla Cogn. in DC. Monog. Phan. 3: 760 (1881). C. Tayuya (Vell.) Cogn. in DC. Monog. Phan. 3: 772 (1881). Citrullus vulgaris Schrad. in Linnaea, 12: 412 (1838). Cucurbita urkupifiana Cardenas in Revist. Agric. Univ. Cochabamba, 2(3): 76 (1946). Cucurbitella Duriaei (Naud.) Cogn. in Mart. Fl. Bras. 6(4): 70, t. 19 (1878). Cyclanthera boliviensis Crovetto in Bol. Soc. Argent. Bot. 2: 174 (1948). C. brachybotrys (Poepp. & Endl.) Cogn. in Mém. Cour. Acad. Belg. 28: 73 (1877). C. explodens var. intermedia Cogn. in O. Ktze. Rev. Gen. 3(2): 103 (1898). C. explodens var. trifida Cogn. in O. Ktze. Rev. Gen. 3(2): 103 (1898). C. Hystrix (Gill.) Arn. in Hook. Journ. Bot. 3: 280 (1841). C. Matthewsii Arn. in Hook. Journ. Bot. 3: 280 (1841), nomen nudum. C. microcarpa Cogn in Mém. Cour. Acad. Belg. 28: 75 (1877). C. montana Cogn. in Meded. Rijks Herbar. 19: 69 (1913). C. pedata Schrad. in Ind. Sem. Hort. Gotting. (1831 C. quinquelobata (Vell.) Cogn. in Mém. Cour. Acad. Belg. 28: 64 (1877). C. Rusbyi Britton in Bull. Torr. Bot. Club, 17: 284 (1890). C. tomentosa Cogn. in Mém. Cour. Acad. Belg. 28: 77 (1877). Echinopepon araneosus (Griseb.) Cogn. in Fedde, Repert. Spec. Nov. 19: 171 (1923). Elaterium amazonicum Mart. in Cogn. in Mém. Cour. Acad. Belg. 28: 55 (1877). Fevillea cordifolia L. Sp. Pl. 1013 (1753). F. Harmsii (0. Ktze.) K. Schum. in O. Ktze. Rev. Gen. 3(2): 103 (1898). F. pergamentacea Cogn. ex O. Ktze. Rev. Gen. 3(2): 104 (1898). Gurania annulata Rusby in Mem. N. Y. Bot. Gard. 7: 382 (1927). G. boliviana Rusby in Mem. Torr. Bot. Club, 3(3): 38 (1893). G. costaricensis var. subtrilobata Cogn. in DC. Monog. Phan. 3: 708 (1881). G. lanata Cogn. in Mém. Cour. Acad. Belg. 27: 26 (1876). G. latifolia Rusby in Mem. N. Y. Bot. Grad. 7: 383 (1927). G. neogranatensis Cogn. in Mém. Cour. Acad. Belg. 27: 34 (1876). G. plumosa Rusby in Mem. N. Y. Bot. Gard. 7: 382 (1927). 200 repandodentata Herzog in Fedde, Repert. Spec. Nov. 7: 69 (1909). spinulosa (Poepp. & Endl.) Cogn. Mém. Cour. Acad. Belg. 27: 17 (1876). Steinbachii Harms in Notizbl. 9: 1042 (1926). variabilis Cogn. in Engler, Pflanzenr. IV, 275 (1): 217 (1916). Weberbaueri Harms in Notizbl. 7: 502 (1921). Lagenaria siceraria (Mol.) Standl. in Field Mus. Publ. Bot. 3: 435 (19380). Luffa cylindrica (L.) Roem. Syn. 2: 63 (1846). Marah Rusbyi (Greene) Greene in Leafl. Bot. Obs. 2: 36 (1910). Melothria Cucumis Vell. Fl. Flum. 29, t. 70 (1825). M. fluminensis Gardn. in Hook. Journ. Bot. 1: 173 (1842). M. Hookeri Cogn. in DC. Monog. Phan. 3: 588 (1881). Momordica Charantia L. Sp. Pl. 1009 (1753). M. Charantia var. abbreviata Ser. in DC. Prodr. 3: 311 (1828). M. balsamina L. Sp. Pl. 1009 (1753). Sicydium tamnifolium (HBK.) Cong. in DC. Monog. Phan. 3: 905 (1881). S. tamnifolium var. grandifolium Cogn. in Engler, Pflanzenr. IV, 275(1): 259 ARAAR Sicyos aculeatus R. E. Fries in Ark. Bot. 6(11): 5, t. 2, fig. 6-7 (1906). S. australis Endl. Prodr. Fl. Norf. 67 (1833). S. debilis Cogn. in O. Ktze. Rev. Gen. 3(2): 104 (1898). S. Kuntzei Cogn. in O. Ktze. Rev. Gen. 3(2): 104 (1898). S. malvifolius Griseb. in Goett. Abh. 19: 147 (1874). S. montanus Poepp. & Endl. Nov. Gen. 2: 53 (1838). S. subcorymbosus Cogn. in DC. Monog. Phan. 3: 887 (1881). S. trigeminus Rusby in Mem. Torr. Bot. Club, 3(3): 38 (1893). Siolmatra brasiliensis (Cogn.) Baill. in Bull. Mens. Soc. Linn. Paris, 1: 458 (1885 CAMPANULACEAE Centropogon aggregatus (Rusby) Gleason in Bull. Torr. Bot. Club, 48: 199 1}. amplifolius Vatke in Linnaea, 38: 716 (1874). aquilinus E. Wimm. in Engler, Pflanzenr. IV, 276b: 259 (1948). aquilinus var. campestris E. Wimm. in Engler, Pflanzenr. IV, 276b: 260 (1943). aquilinus var. integer E. Wimm. in Engler, Pflanzenr. IV, 276b: 260 (1948). Bangii A. Zahlbr. in Bull. Torr. Bot. Club, 24: 372 (1897). Brittonianus A. Zahlbr. in Bull. Torr. Bot. Club, 24: 373 (1897). Brittonianus var. brevidentatus A. Zahlbr. & Rech. in Meded. Rijks Herbar. 19: 51 (1918). cardinalis A. Zahlbr. & Rech. in Meded. Rijks Herbar. 19: 51 (1918). cornutus (L.) Druce in Rept. Bot. Exch. Club Brit. Isl. 3: 416 (1914). fulvus Gleason in Buli. Torr. Bot. Club, 52: 7, t. 1, fig. 10 (1925). dubius (A. Zahlbr.) E. Wimm. in Engler, Pflanzenr. IV, 276b: 196 (1943). gloriosus (Britton) A. Zahlbr. in Bull. Torr. Bot. Club, 24: 873 (1897). Herzogii A. Zahlbr. & Rech. in Meded. Rijks Herbar. 19: 49 (1918). incanus (Britton) A. Zahlbr. in Bull. Torr. Bot. Club, 24: 374 (1897). macrophyllus (G. Don) E. Wimm. in Notizbl. 10: 733 (1929). magnificus A. Zahlbr. & Rech. in Meded. Rijks Herbar. 19: 50 (1913). Mandonis A. Zahlbr. in Ann. Naturh. Hofmus. Wien, 6: 438 (1891). roseus Rusby in Bull. N. Y. Bot. Gard. 8: 123 (1912). unduavensis (Britton) A. Zahlbr. in Bull. Torr. Bot. Club, 24: 374 (1897). unduavensis var. aduanus E. Wimm. in Engler, Pflanzenr. IV, 276b: 230 (1943). yungasensis Britton in Bull. Torr. Bot. Club, 19: 371 (1892). SP NAAKHAHAAHRAARHAAHGH AAAH AAH 201 Diastatea micrantha (HBK.) McVaugh in Bull. Torr. Bot. Club, oi 143 (1940). Hypsela reniformis (HBK.) Presl, Prodr. Monog. Lobel. 45 (183 Lobelia Gardneriana Kanitz in Mart. Fl. Bras. 6(4): 138 agian L. nana HBK. Nov. Gen. & Spec. 8: 317, t. 272 (1819). L, nana var. cymbalarioides (A. Zahlbr.) E. Wimm. in Macbr. in Field Mus. Publ. Bot. 18(6): 479 (1987). . nana var. flagelliformis Wedd. Chlor. And. 2: 13 (1857). . ruderalis Willd. ex Roem. & Schult. Syst. Veg. 6: 56 (1819). . xalapensis HBK. Nov. Gen. & Spec. 3: 315 (1819). Lysipomia glandulifera Schlechtd. in Lechl. Berb. Am. Aust. 58 (1857), nomen ud ec nudum. L. laciniata A. DC. Prodr. 7(2): 349 (1839). L. laciniata var. vulgaris (Wedd.) E. Wimm. in Macbr. in Field Mus. Publ. Bot. 13(6): 486 (1987). L. linearifolia E. Wimm. in Macbr. in Field Mus. Publ. Bot. 18(6): 487 (1937). L. pumila (Wedd.) E. Wimm. in Macbr. in Feld Mus. Publ. Bot. 13(6): 488 ). Pratia boliviensis A, DC. Prodr. 7(2): 340 (1839) — Lobelia nana var. eym- balarioides. P. hederacea Cham. in Linnaea, 8: 212 (1833). Siphocampylus altiscandens Gleason in Bull. Torr. Bot. Club, 48: 198 (1921) = S. flagelliformis. andinus Britton in Bull. Torr. Bot. Club, 19: 373 (1892). andinus var. elegantissimus E. Wimm. in Fedde, Repert, Spec. Nov. 38: 22 (1935 andbins: var. solemnis E. Wimm. in Fedde, Repert. Spec. Nov. 38: 23 (1935). angustiflorus Schlechtd. ex A. Zahlbr. in Bull. Torr. Bot. Club, 24: 379 (1897). argutus A. Zahlbr. in Bull. Torr. Bot. Club, 24: 383 (1897). aureus Rusby in Mem. Torr. Bot. Club, 6: 72 (1896). aureus var. latior A. Zahlbr. in Bull. Torr. Bot. Club, 24: 378 (1897). bilabiatus A. Zahlbr. in Bull. Torr. Bot. Club, 24: 382 (1897). . bilabiatus var. dives (E. Wimm.) E. Wimm. in Engler, Pflanzenr. IV, 276b: 343 (1953). bilabiatus var. gisbrates Lauterb. in Buchtien, Contrib. Fl. Boliv. 1: 187 (1910). boliviensis A. Zahlbr. in Ann. Naturh. Hofmus. Wien, 6: 443 (1891). correoides A. Zahlbr. in Bull. Torr. Pe Club, a 382 (1897). corymbiferus Pohl, Pl. Bras. 2: 112, t. 175 (183 corymbiferus var. gracilis (Britton) A. a in . Bull. Torr. Bot. Club, 24: 384 (1897). crenatus (E. Wimm.) E. Wimm. in Ann. Naturh. Mus. Wien, 56: 323 (1948). elegans var. boliviensis A. Zahlbr. in Bull. Torr. Bot, coke 24: 381 (1897) = S. soraticus in part and S. andinus var. solemnis in elegans var. cordatus A. Zahlbr. in Bull. Torr. Bot. Club, Ae 381 (1897) = S. reflexu Fiebrigii ‘Wimm: in Fedde, Repert, Spec. ‘Nov. 22: 209 (1926). Fiebrigii var. intermedius E. Wimm. in Engler, Pflanzenr. IV, 276b: 310 (1953). flagellformis A. Zahlbr. in Bull. Torr. Bot. Club, 24: 380 (1897). flagelliformis var. glaber E. Wimm. in Ann. Naturh. Mus. Wien, 56: 324 (1948). flavoruber Gleason in Torreya, 25: 94 (1925). foliosus var. =— E. Wimm. in Macbr. in Field Mus. Publ. Bot. 13(6): 453 (1937). S. Kuntzeanus A. Zahlbr.-in Bull. Torr. Bot. Club, 24: 378 (1897). wa ta DnnnAn nin ”R le Pm mn th 202 Lorentzii E. Wimm. in Fedde, Repert. Spec. Nov. 29: 85 membranaceus Britton in Bull. Torr. Bot. Club, 19: 372 (1892 nemoralis var. tarijanus E. Wimm. in Revist. Sudam. Bot. 2: "92 (1935). neurotrichus E. Wimm. in Fedde, Repert. Spec. Nov. 38: 24 (1935). nummularius E. Wimm. in Fedde, Repert. Spec. Nov. 38: 18 (1935). oblongifolius Rusby in Mem. Torr. Bot. Club, 6: 73 (1896). Orbignyanus DC. Prodr. 7(2): 405 (1839). Orbignyanus var. discurrens E. Wimm. in Ann. Naturh. Mus. Wien, 56: 323 1948). puberulus E. Wimm. in Fedde, Repert. Spec. Nov. a 13 (1929). radiatus Rusby in Mem. Torr. Bot. Club, 6: 73 (18 radiatus var. brevidentatus (A Zahlbr. & Rech.) fe ee in Ann. Naturh. Mus. Wien, 56: 331 (1948). radiatus var. minor A. Zahlbr. in Bull. Torr. Bot. Club. 24: 376 (1897). reflexus Rusby in Bull. N. Y. Bot. Gard. 4: 403 (1907). Rusbyanus Britton in Bull. Torr. Bot. Club, 19: 372 (1892). soraticus E. Wimm. in Fedde, Repert. Spec. Nov. 38: 23 (1935). soraticus var. angustatus E. Wimm. in Fedde, Repert. Spec. Nov. 38: 23 (1935). sparsipilus E. Wimm. in Fedde, Repert. Spec. Nov. 19: 388 (1924). subcordatus Rusby in Bull. N. Y. Bot. Gard. 8: 121 (1912). tunarensis A. Zahlbr. in Bull. Torr. Bot. Club, 24: 376 (1896). tunicatus A. Zahlbr. ex O. Ktze. Rev. Gen. 3(2): 189 (1898). tupaeformis A. Zahlbr. in Ann. Naturh. Hofmus. Wien, 6: 440 (1891). tupaeformis var. diversifolius Hicken in Darwiniana, 1: 142 (1924). Vatkeanus A. Zahlbr. in Bull. Torr. Bot. Club, 24: 377 (1897). virgatus A. DC. Prodr. 7(2): 398 (1839). Werdermannii E. Wimm. in Fedde, Repert. Spec. Nov. 38: 76 (19385). Williamsii Rusby in Bull. N. Y. Bot. Gard. 8: 122 (1912). Sphenoclea zeylanica Gaertn. Fruct. 1: 118, t. 24 (1788). Wahlenbergia arida (HBK.) Griseb. in Goett. Abh. 19: 200 (1874). W. linarioides (Lam.) A. DC. Monog. Campan. 158 (1830). W. peruviana A. Gray in Proc. Amer. Acad. 5: 152 (1861). . . . DAnnnAnAnAAIn Minninin Win tn CALYCERACEAE Acicarpha laxa R. E. Fries in Ark. Bot. 6(11): 2, t. 1, fig. 1-5 (1907). A. tribuloides Juss. in Ann. Mus. Paris, 2: 348, t. 58, fig. 1 (1803). Calycera pulvinata Remy in Ann. Sci. Nat. (ser. 3) 6: 352 (1846). COMPOSITAE Acanthospermum australe (L.) O. Ktze. Rev. Gen. 3(2): 128 (1898). A. hispidum DC. Prodr. 5: 522 (1836). Achyrocline alata DC. Prodr. 6: 221 (1838). A. candicans (HBK.) DC. Prodr. 6: 221 (1838). A. celosioides (HBK.) DC. Prodr. 6: 221 (1838). hyperchlora Blake in Bot. Gaz. 74: 415 (1922). latifolia Wedd. Chlor. And. 1: 148 (1856). polycephala Rusby in Bull. N. Y. Bot. Gard. 4: 388 (1907). ramosissima Britton ex Rusby in Mem. Torr. Bot. Club, 3(3): 57 (1893). rufescens (HBK.) DC. Prodr. 6: 220 (1838). satureioides (Lam.) DC. Prodr. 6: 220 (1838). tomentosa Rusby in Bull. N. Y. Bot. Gard. 4: 388 (1907). Vauthieriana DC. Prodr. 6: 220 (1838). venosa Rusby in Mem. Torr. Bot. Club, 3(3): 57 (1893). Addisonia boliviana Rusby, Descr. S. Am. Pls. 147 (1920). . PPP Pp > bbb 203 Adenostemma triangulare DC. Prodr. 5: 118 (1836). A. viscosum var. brasilianum (Cass.) Baker in Mart. Fl. Bras. 6(2): 185 (1876). Ageratum conyzoides L. Sp. Pl. 839 (1753). Ambrosia elatior L. Sp. Pl. 987 (1753). Anthemis Cotula L. Sp. Pl. 894 (1753). Artemisia Absinthium L. Sp. Pl. 848 (1753). Aspilia lucidula Blake in Proc. Biol. Soc. Wash. 36: 52 (1923). Aster acaulis Wedd. Chlor. And. 1: 189, t. 833A (1857). Bangii Rusby in Mem. Torr. Bot. Club, 4(3): 213 (1895). divaricatus var. graminifolius (Spreng.) Baker in Mart. Fl. Bras. 6(3): 22 exilis Ell, Sketch, 2: 344 (1828). exilis forma subalpinus R. E. Fries in Ark. Bot. 5(18): 12 (1906). limophilus (Sch. Bip.) Hemsl. & Pearson in Journ. Linn. Soc. 35: 86 (1901). marginatus HBK. Nov. Gen. & Spec. 4: 91 (1820). Vahlii (Gaud.) Hook. & Arn. in Hook. Compan. Bot. Mag. 2: 49 (1836). Baccharis alpina HBK. Nov. Gen. & Spec. 4: 48 (1820). alpina var. nummularioides Heering in Jahrb. Hamb. Wiss. Anst. 21, Beih. 3: 35 (1904). alpina var. serpyllifolia Wedd. Chlor. And. 1: 168 (1856). aphylla var. boliviensis Sch. Bip. in Bull. Soc. Bot. France, 12: 81 (1865), Pre r re. ee nomen nudum. articulata (Lam.) Pers. Synops. Pl. 2: 425 (1807). caespitosa (R. & P.) Pers. Synops. Pl. 2: 425 (1807). capitalensis Heering in Jahrb. Hamb. Wiss. Anst. 31, Beih. 3: 102 (1915). caprariaefolia DC. Prodr. 5: 416 (1836). cassinoides DC. Prodr. 5: 412 (1836). condensata Rusby, Descr. S. Am. Pls. 148 (1920). Conwayi Rusby in Bull. N. Y. Bot. Gard. 8: 130 (1912). coridifolia DC, Prodr. 5: 422 (1836). cylindrica (Less.) DC. Prodr. 5: 426 (1836). debilis Rusby in Mem. Torr. Bot. Club, 6: 60 (1896). densiflora Wedd. Chlor. And. 1: 175 (1856). dracunculifolia DC. Prodr. 5: 421 (1836). dracunculifolia var. integerrima O. Ktze. Rev. Gen. 3(2): 132 (1898). dracunculifolia var. integerrima forma subviscosa O. Ktze. Rev. Gen. 3(2): 132 (1898). effusa Griseb. in Goett. Abh. 19: 177 (1874). fallax O. Ktze. Rev. Gen. 3(2): 1382 (1898). floribunda HBK. Nov. Gen. & Spec. 4: 64, t. 325 (1820). genistelloides (Lam.) Pers. Synops. Pl. 2: 425 (1807). Gilliesii A. Gray in Proc. Am. Acad. 5: 123 (1861). glutinosa Pers. Synops. Pl. 2: 425 (1807). grandicapitulata Hieron. in Engler, Bot. Jahrb. 36: 481 (1905). grindeliaefolia Wedd. Chlor. And. 1: 176 (1856). Grisebachii Hieron. in Bol. Acad. Nac. Cérdova, 4: 36 (1881). hemiprionoides Buek, Ind. 1: x (1842) = B. Sternbergiana. heterothalamoides Britton in Bull. Torr. Bot. Club, 19: 4 (1892). juncea Desf. Cat. Hort. Paris (ed. 3), 163 (1829). Kuntzeana Teodoro in Contrib. Inst. Geobiol. La Salle, Canoas, no. 2: 46 (1952 lanceolata HBK. Nov. Gen. & Spec. 4: 63 (1820). latifolia Pers. Synops. Pl. 2: 424 (1807 laxiflora Rusby in Bull. N. Y. Bot. Gard. 8: 129 (1912). magellanica var. subviscosa O. Ktze. Rev. Gen. 3(2): 183 (1898). Dem Pe hw BO bo De Sd BS Oo be be Oe bo do bb OO bb bY ae 204 VETERE Eee eee e Ww Wm wl DWWWHDHDND DOWD Mandonii Sch. Bip. in Leopoldina, 25: 108 (1889). mapirensis Rusby in Mem. Torr. Bot. Club, 6: 61 (1896). marginalis var. coerulescens (DC.) Heering ex Reiche, Fl. Chil. 4: 11 (1905). marginalis var. viminea Heering ex Reiche, Fl. Chil. 4: 10 (1905). medullosa DC. Prodr. 5: 405 (1836). microphylla HBK. Nov. Gen. & Spec. 4: 55 (1820). microphylla var. incarum Wedd. Chlor. And. 1: 170 (1856). microphylla var. linearifolia Wedd. ex Sch. Bip. in Bull. Soc. Bot. France, 12: 81 (1865), nomen nudum. microphylla var. pulverulenta Rusby in Mem. Torr. Bot. Club, 3(3): (1893). myriocephala Baker in Mart. Fl. Bras. 6(3): 93 (1882). nitida (R. & P.) Pers. Synops. Pl. 2: 425 (1807). oblanceolata Rusby in Mem. Torr. Bot. Club, 6: 61 BhGP OD: obtusifolia HBK. Nov. Gen. & Spec. 4: 51 (1820 odorata HBK. Nov. Gen. & Spec. 4: 52 (1820). oppositifolia O. Ktze. Rev. Gen. 3(2): 1383 (1898). Orbignyana Klatt in Abh. Naturf. Ges. Halle, 15: 327 (1881). oxyodonta var. punctulata (DC.) Baker in Mart. Fl. Bras. 6(3): 77 (1882). . papillosa Rusby in Bull. N. Y. Bot. Gard. 8: 129 (1912). paucidentata DC. Prodr. 5: 420 (1836). Pentlandii DC. Prodr. 5: 416 (1836). perulata O. Ktze. Rev. Gen. 3(2): 133 (1898). Pflanzii Perkins in Engler, Bot. Jahrb. a 224 (1913). polycephala Wedd. Chlor. And. 1: 178 (1856). prostrata (R. & P.) Pers. Synops. Pl. 2: 425 (1807). pulchella Sch. Bip. in Bull. Soc. Bot. France, 12: 81 (1865), nomen nacdune, pulverulenta Klatt in Abh. Naturf. Ges. ele 15: 327 (1881). quitensis HBK. Nov. Gen. & Spec. 4: 57 (18 resinosa var. truncatifolia Perkins in i Bot. Jahrb. 49: 224 (1913). retusa DC, Prodr. 5: 412 (1836). rhexioides HBK. Nov. Gen. & Spec. 4: 66 (1820). riparia HBK. Nov. Gen. & Spec. 4: 65 (1820). rubricaulis Rusby in Bull N. Y. Bot. Gard. 8: 129 (1912). salicifolia (R. & P.) Pers. Synops. Pl. 2: 425 (1807). saliens Rusby in Bull. N. Y. Bot. Gard. 4: 387 (1907). scandens (R. & P.) Pers. Synops. Pl. 2: 424 (1807). semiserrata DC. Prodr. 5: 404 (1836). serrulata Pers. Synops. Pl. 2: 423 (1807). spartea Benth. Bot. Voy. Sulphur, 114 (1845). Sternbergiana Steud. Nomencl. (ed. 2) 1: 179 (1840). Sternbergiana var. pubescens Perkins in Engler, Bot. Jahrb. 49: 225 (1913). subalata Wedd. Chlor. And. 1: 174 (1856). subpenninervis Sch. Bip. in Linnaea, 34: 532 (1865-66), nomen nudum. syncephala Sch. Bip. ex Rusby in Bull. N. Y. Bot. Gard. 4: 386 (1907). tomentosa Pers. Synops. Pl. 2: 424 (1807). trichoclada DC. Prodr. 5: 400 (1836). tridentata DC. Prodr. 5: 409 (1836). tridentata var. pluridentata DC. Prodr. 5: 409 (1836). trinervis (Lam.) Pers. Synops. Pl. 2: 423 (1807). tucumanensis Hook. & Arn. in Hook. Journ. Bot. 3: 34 (1841). ulicina var. multifida (Griseb.) O. Ktze. Rev. Gen. 3(2): 135 (1898). viscosa var. nigricans O, Ktze. Rev. Gen. 1: 320 (1891). Vitis-Idaea O. Ktze. Rev. Gen. 3(2): 185 (1898) = B. Kuntzeana. Barnadesia glomerata O. Ktze. Rev. Gen. 3(2): 135 (1898). WD ROW Wh Ol lkh Oh PeEreryye 205 inermis Rusby in Bull. N. Y. Bot. Gard. 4: 399 (1907). polyacantha Wedd. Chlor. And. 1: 13, t. 1A (1855). . Seleriana Muschl. in Engler, Bot. Jahrb. 50, Beibl. 111: 99 (1913). spinosa L. f. Suppl. 348 (1781). venosa Rusby in Mem. Torr. Bot. Club, 6: 69 (1896). idens andicola HBK. Nov. Gen. & Spec. 4: 237 (1820). andicola var. Cosmantha (Griseb.) Sherff in Bot. Gaz. 85: 2 (1928). andicola var. Cosmantha forma Buchtienii (Sherff) Sherff in Field Mus. Publ. Bot. 16: 502 (1937). andicola var. decomposita O. Ktze. Rev. Gen. 3(2): 136 (1898). andicola var. Mandonii Sherff in Bot. Gaz. 80: 380 (1925). andicola var. tarijensis Sherff in Bot. Gaz. 85: 14 (1928). andicola var. tarijensis forma dissecta Sherff in Bot. Gaz. 85: 14 (1928). . grandiflora var. breviloba O, Ktze. Rev. Gen. 3(2): 136 (1898) = B. an- dicola var. decomposita. . grandiflora var. longiloba O. Ktze. Rev. Gen. 3(2): 136 (1898). . humilis HBK. Nov. Gen. & Spec. 4: 234 (1820). . humilis var. tenuifolia Sch. Bip. ex Griseb. in Goett. Abh. 24: 198 (1879) = B. triplinervia var. macrantha. . longipetiolata Rusby in Bull. N. Y. Bot. Gard. 8: 131 (1912) = B. segetum var. patula. ; macrantha Griseb. in Goett. Abh. 19: 186 (1874). . pallida Rusby in Bull. N. Y. Bot. Gard. 4: 389 (1907) = B. se . peucedanifolia var. soratensis O. Ktze. Rev. Gen. 3(2): 137 Py Cosmos peucedanifolius. . pilosa L. Sp. Pl. 832 (1753). . pilosa var. alausensis (HBK.) Sherff in Bot. Gaz. 81: 35 (1926). . pilosa var. alausensis forma scandicina (HBK.) Sherff in Bot. Gaz. 81: 36 (1926). . pilosa var. minor (Blume) Sherff in Bot. Gaz. 80: 387 (1925). . pilosa var. radiata (Sch. Bip.) J. A. Schmidt, Beitr. Fl. Cap Verdischer Ins. 197 (1852). . pseudocosmos Sherff in Bot. Gaz. 76: 151 (1923). rubifolia HBK. Nov. Gen. & Spec. 4: 237, t. 881 (1820). segetum Mart. ex Colla, Herb. Pedem. 3: 307 (1835). segetum var. patula (Gardn.) Sherff in Field Mus. Publ. Bot. 16: 197 (1937). squarrosa HBK. Nov. Gen. & Spec. 4: 238 (1820). _ tenera var. tetracera Sherff in Bot. Gaz. 88: 293 (1929). . triplinervia var. macrantha (Wedd.) Sherff in Bot. Gaz. 80: 383 (1925). . triplinervia var. macrantha forma octoradiata Sherff in Bot. Gaz. 92: 203 31). ee hes mollis (Poepp. & Endl.) Sherff in Bot. Gaz. 80: 384, t. 22, fig. a-i (19 25). Brickellia cay (Vahl) A. Gray, Pl. Wright. 1: 86 (1852). B. paucidentata Klatt in Abh. Nat turf. Ges. Halle, 15: 326 (1881) = Cacalia ysis O. Ktze. Rev. Gen. 3(2): 138 (1898) = Vernonia je oer Calea anomala Hassl. in Fedde, Repert. Spec. Nov. 7: 856 (1909). ANQNAAN Cep. brevifolia Rusby in Bull. N. Y. Bot. Gard. 8: 132 (1912). coriacea DC. Prodr. 5: 675 (1836). cymosa Less. in Linnaea, 5: 158 (1880). rhombifolia Blake in Proc. Biol. Soc. Wash. 36: 53 (1923). robusta Britton in Bull. Torr. Bot. Club, 19: 151 (1892) = C. coriacea. solidaginea HBK. Nov. Gen. & Spec. 4: 295 (1820). halophora robusta Rusby in Mem. Torr. Bot. Club, 3(3): 63 (1893) = Helenium sp. ? 206 Chaetanthera boliviensis Koster in Blumea, 5: 673 (1945). C. Stuebelii Hieron. in Engler, Bot. Jahrb. 21: 368 (1895). Chaptalia ebracteata (O. Ktze.) K. Schum. in Just, Bot. Jahresber. 26(1): 376 (1900). integrifolia (Cass.) Baker in Mart. Fl. Bras. ee 377 (1884). majuscula Greene, Leafi. Bot. Obs. 1: 196 (19 Mandonii Burkart in Darwiniana, 6: 551 SHY microdonta Greene, Leafi. Bot. Obs. 1: 196 (1905). nutans (L.) Hemsl. in Biol. Centr. Am. Bot. 2: 255 (1881). piloselloides (Vahl) Baker in Mart. Fl. Bras. 6(3): 378 (1884). rotundifolia D. Don in Trans. Linn. Soc. 16: 242 (1830). Chersodoma Antennaria (Wedd.) Cabrera in Rev. Mus. La Plata, 6: 352 (1946). C. candida Phil. in Anal. Mus. Nac. Chile, Sec. 2, Bot. 8: 33 (1891). C. diclina (Wedd.) Cabrera in Rev. Mus.- La Plata, 6: 353 (1946). C. iodopappa (Sch. Bip.) Cabrera in Rev. Mus. La Plata 6: 350 (1946). Chevreulia elegans Rusby in Bull. N. Y. Bot. Gard. 4: 389 (1907). C. sarmentosa (Pers.) Blake in Proc. Biol. Soc. Wash. 38: 85 (1925). Chiliotrichiopsis Keidelii Cabrera in Not. Mus. La Plata, 2: 172, fig. 1 (1937). Chrysanthellum americanum (L.) Vatke in Bremen Abh. 9: 122 (1885). Chrysanthemum Parthenium Bernh. Syst. Verz. Erf. 145 (1800). Chuquiraga acanthophylla Wedd. Chlor. And. 1: 5 (1855). armata Koster in Blumea, 5: 662 (1945). brasiliensis (Spreng.) O. Ktze. Rev. Gen. 3(2): 141 (1898). brasiliensis var. divaricata (Griseb.) O. Ktze. Rev. Gen. 3(2): 141 (1898). ferox (Wedd.) Britton in Bull. Torr. Bot. Club, 19: 266 (1892). insignis var. armata Wedd. Chlor. And. 1: 3 (1855). Jussieui var. lanceifolia (Humb. & Bonpl.) Koster in Blumea, 5:662 (1945). longiflora (Griseb.) Hieron. Plantae Diaphoricae, 165 (1882 oppositifolia Gill. & Don in Phil. Mag. 11: 392 (1832). oppositifolia var. macrocephala Wedd. Chlor. And. 1: 3 (1855). oppositifolia var. microcephala Wedd. Chlor. And. 1: 3 (1855). parviflora (Griseb.) Hieron. in Engler, Bot. Jahrb. 49: 231 (1913). rotundifolia Wedd. Chlor. And. 1: 4. t. 4A (1855). varians (Gardn.) Rusby in Bull. N. Y. Gard. 4: 399 (1907). Clibadium asperum (Aubl.) DC. Prodr. 5: 506 (1836). C. heterotrichium Blake in Contrib. Gray Herb. me 8 (1917). C. peruvianum Poepp. in DC. Prodr. 5: 505 (18 C. remotiflorum O. E. Schulz in ea Bot. ake 46: 621 (1912). C. surinamense L. Mant. 2: 294 (177 Cnicothamnus Azafran (Cabr.) faece in Not. Mus. La Plata, 9: 256 (1944). C. Lorentzii Griseb. in Goett. Abh. 19: 197 (1874). Conyza andicola Phil. in Anal. Mus. Nac. Chile, 8: 38 (18 C. artemisioides Meyen & Walp. in Nov. Act. Acad. se ae 19, Suppl. 1: 262 (1848). C. chilensis Spreng. Novi Proventus, 14 (1819). evacioides Rusby in Bull. N. Y. Bot. Gard. 4: 385 (1907) — C. gnaphalioides. .- gnaphaliciaea HBK. Nov. Gen. & Spec. 4: 73, t. 327 (1820). lignescens Rusby in Bull. N. Y. Bot. Gard. 4: 385 (1907). lyrata HBK. Nov. Gen. & Spec. 4: 70 (1820) obtusa HBK. Nov. Gen. & Spec. 4: 71 (1820). suffruticosa Phil. in Linnaea, 28: 735 (1858). . yungasensis Rusby in Mem. Torr. Bot. Club, 3(3): 55 (189 pre boliviana Blake in Contrib. U. S. Nat. Herb. 22: ne (1924) == C. Pickeringii. C. fasciculata Wedd. Chlor. And. 1: 71 (1856). PNANAAAHRAHAAAAA Ree hee gEEGED? 207 C. Pickeringii A. Gray in Proc. Am. Acad. 5: 124 (1861). C. spectabilis A. Gray in Proc. Am. Acai 5: 125 (1861). . diversifolius Otto in Knowles & Westc. Fl. Cab. 2: 6, ‘ 47 (1838). Herzogii Sherff in Bot. Gaz. 96: 148 (1934). integrifolius Wedd. Chlor. And. 1: 70 (1856) — C. peucedanifolius var. peucedanifolius Wedd. Chlor. And, 1: 70 (1856). peucedanifolius var. cochabambensis (O. Ktze.) Sherff in Bot. Gaz. 88: 307 (1929 ‘ peucedanifolius var. tiraquensis (O. Ktze.) Sherff in Bot. Gaz. 88: 307 1929 Qa S29 ao Cotula oyemaca (HBK.) Hemsl. in Biol. Centr. Am. Bot. 2: 230 (1881). Crepis patorephaiiie Klatt in Ann. Naturh. Hofmus. Wien, 9: 368 (1894) = Hieracium sp. Culcitium Neaei pen Sch. Bip. ex Wedd. Chlor. And 1: 140 (1856). C. nivale HBK. Nov. Gen. & Spec. 4: 171, t. 363 (1820). Dimerostemma asperatum Blake in Contrib. Gray Herb. 52: 12 (1917). Diplostephium atropurpureum Rusby in Bull. N. Y. Bot. Gard. 4: 384 (1907) = D. Haenkei. - Haenkei (DC.) Wedd. Chlor. And. 1: 203 (1857). liaboides Rusby in Bull. N. Y. Bot. Gard. 4: 384 (1907) = D. Haenkei. Mandonii Rusby in Bull. N. Y. Bot. Gard. 4: 383 (1907) = D. Haenkei. sejaénse (O. Ktze.) Blake in Contrib. U. S. Nat. Herb. 24: 80 (1929) = D. Haenkei. Dyssodia fastigiata DC. Prodr. 5: 640 (1836). Eclipta alba (L.) Hassk. Pl. Rar. Jav. 528 (1848). Egletes viscosa (L.) Less. Syn. 252 (1832). Elephantopus angustifolius Sw. Prodr. 115 (1788). E. scaber L. Sp, Pl. 814 (1753). E. spicatus B. Juss. ex Aubl. Pl. Guian. Fr. 2: 808 (1775). E. tomentosus L. Sp. Pl. 814 (1753). Eleutheranthera ruderalis Sch. Bip. in Bot. Zeit. 24: 165 (1866). Elvira biflora (L.) DC. Prodr. 5: 503 (1836) Encelia hirsuta O. Ktze. Rev. Gen. 3(2): 145 (1898). E. soratensis Rusby, Descr. S. Am. Pls. 154 (1920). Erechtites hieracifolia (L.) Raf. ex DC. Prodr. 6: 294 (1838). E. valerianaefolia (Wolf) DC. Prodr. 6: 295 (1838). Erigeron bonariensis L. Sp. Pl. 863 (1753 rittonianus Rusby in Mem. Torr. Bot. ‘Club, 3(3): 54 (1893) = E. rosu- latus. canadensis L. Sp. Pl. 863 (1753). canescens Sch. Bip. in Linnaea, 34: 534 (1866), nomen nudum. cinerascens Sch. Bip. in Bonplandia, 4: 54 (1856), nomen nudum. ferrugineus Wedd. Chlor. And. 1: 195 (1857). floribundus (HBK.) Sch. Bip. in Bull. Soc. Bot. France, 12: 81 (1865). frigidus Wedd. Chlor. And. 1: 231 (1857). hieracioides Wedd. Chlor. And. 1: 194, t. 34B (1857). Hillii Domke in Notizbl. 13: 244 (1936). lanceolatus Wedd. Chlor. And. 1: 193 (1857). lanceolatus var. subacaulis Wedd. Chlor. And. 1: 193 (1857). laxiflorus Baker in Mart. Fl. Bras 6(3): 31 (1882). pazensis Sch. Bip. ex Rusby in Mem. Torr. Bot. Club, 3(3): 54 (1893). pulvinatus Wedd. Chlor. And. 1: 194, t. 833B (1857). . rosulatus Wedd. Chlor. And. 1: 193, t. 33C (1857). bebe he to S oo emiamplexicaulis Meyen, Reise, 1: 311 (1834). sekaaibaraii Blake in Biol. Soc. Wash. 8 51 (1923). senecioides Wedd. Chlor. And. 1: 198 (185 tunariensis O. Ktze. Rev. Gen. 3(2): 146 pee 8). upatorium alternifolium var. genuinum forma nitidum Koster in Blumea, 5: 651 (1945). amygdalinum Lam. Encycl. 2: 408 (1786). Arnottianum Griseb. in Goett. Abh. 19: 167 (1874). austerum Robinson in Contrib. Gray Herb. 68: 9 (1923). azarangoénse Sch. Bip. ex Wedd. Chlor. And. 1: 217 (1857). Bangii Rusby in Mem. Torr. Bot. Club, 6: 56 (1896). betonicaeforme (DC.) Baker in Mart. Fl. Bras. ane 362, t. 96 (1876). Bridgesii Robinson in Proc. Am. Acad. 55: 7 (19 buniifolium Hook. & Arn. in Compan. Bot. Mag. 4: 240 (1836). bupleurifolium DC. Prodr. 5: 149 (1836). calderillense Hieron. in Engler, Bot. Jahrb. 40: 381 (1908). camachense Hieron. in Engler, Bot. Jahrb. 40: 386 (1908). camataquiense Hieron. in Engler, Bot. Jahrb. 40: 377 (1908). chaparense Robinson in Contrib. Gray Herb. 90: 24 (1930). chiquitense Robinson in Contrib. Gray Herb. 68: 11 (1923). clematideum Griseb. in Goett. Abh. 24: 172 (1879). cochabambense Hieron. in Engler, Bot. Jahrb. 22: 745 (1897) = E. con- nivens. confluentis Robinson in Contrib. Gray Herb. 77: 11 (1926). connivens Rusby in Mem. Torr. Bot. Club, 6: 57 (1896). conoclinanthium Hieron. in Engler, Bot. Jahrb. 40: 388 (1908). conyzoides var. ciliatum (Hook. & Arn.) Hieron. in Engler, Bot. Jahrb. 22: 741 (1897). crenulatum Spreng. ex Hieron. in Engler, Bot. Jahrb. 22: 776 (1897). dejectum Robinson in Contrib. Gray Herb. 77: 12 (1926). dentatum Gardn. in Hook. Journ. Bot. 6: 443 (1847). desmocephalum Robinson in Contrib. Gray Herb. 68: 14 (1923). didymum Klatt in Ann. Naturh. Hofmus. Wien, 9: 356 (1894). endytum Robinson in Proc. Am. Acad. 55: 18 (1919). eucosmum Robinson in Contrib. Gray Herb. 61: 6 (1920) — E. saltense. euphyes Robinson in Contrib. Gray Herb. 68: 16 (1923). extensum Gardn. in Hook. Journ. Bot. 6: 440 (1847). Fiebrigii Hieron. in Engler, Bot. Jahrb. 40: 371 (1908). gloeocladum Robinson in Proc. Am. Acad. 55: 17 (1919). grossidentatum Hieron. in Engler, Bot. Jahrb. 40: 377 (1908). naiense Britton in Bull. Torr. Bot. Club, 18: 333 (1891) = E. ivaefolium. gynoxioides Rusby in Bull. N. Y. Bot. Gard. 4: 380 (1907) = E. gynoxo- morphum. gynox xomorphurh Rusby in Robinson in Contrib. Gray Herb. oe 7 (1920). hecatanthum (DC.) Baker in Mart. Fl. Bras. 6(2): 365 (1876). Herzogii Robinson in Contrib. Gray Herb. 68: 19 (1923). hirsutum Hook. & Arn. in Compan. Bot. Mag. 1: 239 (1836). hosanense Robinson in Contrib. Gray Herb. 100: 14 (19382). ignoratum Hieron. in Engler, Bot. Jahrb. 40: 379 (1908). inulaefolium HBK. Nov. Gen. & Spec. 4: 109 (1820). inulaefolium forma suaveolens (HBK.) Hieron. in Engler, Bot. Jahrb. 29: 11 (1900). iresinoides HBK. Nov. Gen. & Spec. 4: 106, t. 340 (1820). ivaefolium L. Syst. (ed. 10) 1205 (1759). ivaefolium var. extrorsum Baker in Mart, Fl. Bras. 6(2): 290 (1876). . jugipaniculatum Rusby in Bull. N. Y. Bot. Gard. 4: 379 (1907). ahah alata atte 5: ous bak les hel Rakai SER EE Ef sedan bet is Bi id et ttt be . E. BOP iS td yd Bt bt oy w 209 kleinioides HBK. Nov. Gen. & Spec. 4: 120 (1820). Kuntzei Hieron. in Engler, Bot. Jahrb. 22: 766 (1897) = Ophryosporus macrodon. laevigatum Lam. Encycl. 2: 408 (1786). laevigatum forma albiflorum O, Ktze. Rev. Gen. 3(2): 147 (1898). laevigatum forma flavidum O. Ktze. Rev. Gen. 3(2): 147 (1898). laevigatum forma lilacinum O. Ktze. Rev. Gen. 3(2): 147 (1898). lanigerum var. longicuneatum Robinson in Ostenia, 354 (1933). lasiophthalmum Griseb. in Goett. Abh. 19: 167 (1874). latipaniculatum Rusby in Bull. N. Y. Bot. Gard. 4: 380 (1907). leptocephalum DC. Prodr. 5: 148 (1836). ge pon arse var. hypomalacum Robinson in Contrib. Gray Herb. 80: 24 8). lobatum Robinson in Proc. Am. Acad. 55: 21 (1919). Lobbii Klatt in Ann. Naturh. Hofmus. Wien, 9: 356 (1894). longipetiolatum Sch. Bip. ex Rusby in Mem. Torr. Bot. Club, 3(3): 52 (1893). macrocephalum Less. in Linnaea, 5: 1386 (1830). macrophyllum L. Sp. Pl. (ed. 2) 2: 1175 (1763). mallotum Robinson in Proc. Am. Acad. 55: 22 (1919). mallotum var. aporum Robinson in Proc. Am. Acad. 55: 23 (1919). Mandonii Sch. Bip. in Linnaea, 34: 533 (1865-66) = E, bupleurifolium. mapiriense Hieron. in Engler, Bot. Jahrb. 40: 374 (1908). marginatum Poepp. & Endl. Nov. Gen. 3: 54 (1845). microstemon Cass. in Dict. Sci. Nat. 25: 4382 (1822). morifolium Mill. Gard. Dict. (ed. 8) no. 10 (1768). patens var. rhodolaenum Griseb. in Goett. Abh. 24: 170 (1879). patens var. tomentosum Hieron. in Engler, Bot. Jahrb. 22: 773 (1897). Pentlandianum DC. Prodr. 5: 157 (1836). phyllocephalum Klatt in Ann. Naturh. Hofmus. Wien, 9: 358 (1894). polopolense Robinson in Contrib. Gray Herb. 61: 10 (1920) = E. pyeno- cephalum. porophylloides Robinson in Contrib. Gray Herb. 68: 29 (1923). pteropodum Hieron. in Engler, Bot. Jahrb. 29: 15 (1900) — E. nemorosum. pycnocephalum Less. in Linnaea, 6: 404 (1831). pyramidale Klatt in Abh. Naturf. Ges. Halle, 15: 323 (1881). pyramidale forma angustifolium (Hieron.) Robinson in Contrib. Gray Herb. 61: 56 (1920). rufescens var. glabratum Hieron. ex Robinson in Contrib. Gray Herb. 61: 67 (1920). Rusbyi Britton in Bull, Torr. Bot. Club, 18: 334 (1891) = E. nemorosum. santacruzense Hieron. in Engler, Bot. Jahrb. 22: 762 (1897). scopulorum Wedd. Chlor. And. 1: 216, t. 40B (1857) = E. azangaroénse. simillimum Robinson in Contrib. Gray Herb. 77: 38 (1926). soratae Sch. Bip. ex Robinson in Contrib. Gray Herb. 61: 51 (1920). solidaginoides HBK. Nov. Gen. & Spec. 4: 126 (1820). ordescens var. bolivianum Rusby in Mem. Torr. Bot. Club, 6: 56 (1896) = E. endytum. squalidum var. Rusbyanum Robinson in Proc. Am. Acad. 55: 34 (1919). squarroso-ramosum Hieron. in Engler, Bot. Jahrb. 22: 7 53 (1897). stachyophyllum Spreng. Syst. Veg. 3: 420 (1826). Sternbergianum DC. Prodr. 5: 167 (1836). steviaefolium DC. Prodr. 5: 158 ( 1836). stipuliferwm Rusby in Mem. Torr. Bot. Club, 4(3): 210 (1895) = E. soli- daginoides. subseandens Hieron. in Engler, Bot. Jahrb. 22: 742 (1897). tamboénse Hieron. in Engler, Bot. Jahrb. 22: 770 (1897). 210 thymifolium Britton in Bull. Torr. Bot. Club, 19: 1 (1892). toldense Hieron. in Engler, Bot. Jahrb. 40: 378 (1908). triosteifolium Rusby in Bull. N. Y. Bot. Gard. 4: 379 (1907). tucumanense Lillo & Robinson in Contrib. Gray Herb. 90: 32 (1930). tunariense (Hieron.) Robinson in Contrib. Gray Herb. 61: 39 (1920). vitalbae DC. Prodr. 5: 163 (1836). yungasense Robinson in Contrib. Gray Herb. 104: 30 (1934). Facelis capillaris Rusby in Mem. Torr. Bot. Club, 6: 62 (1896). F. Schultziana Beauverd in Bull. Soc. Bot. Genéve (ser. 2), 5: 219 (1913). F. Weddelliana Beauverd in Bull. Soc. Bot. Genéve (ser. 2), 5: 217 (1913). Filago lasiocarpa Griseb. in Goett. Abh. 19: 180 (1874). Flaveria chilensis J. F. Gmel. Syst. 1269 (1791). F. contrayerba (Cav.) Pers. Synops. Pl. 2: 489 (1807). Flourensia Fiebrigii Blake in Engler, Bot. Jahrb. 54, ae 119: 47 (1916). F. heterolepis Blake in Contrib. Gray Herb. 54: 186 (19 Franseria artemisioides Willd. Sp. Pl. 4(1): 378 se F. Conwayi Rusby in Bull. N. Y. Bot. Gard. 8: 130 RG F. recurva Rusby in Bull. N. Y. Bot. Gard. 8: 131 (1912). Galinsoga calva Rusby in Mem. Torr. Bot. Club, 3(3): 61 (1893). G. parviflora Cav. Icon. 3: 41, t. 281 (1795). G. parviflora var. hispida DC. Prodr. 5: 677 (1836). bs bed et bt bt badium Wedd. Chlor. And. 1: 145 (1856). cheiranthifolium Lam. Encycl. 2: 752 (1786). cheiranthifolium var. multiflorum Koster in Blumea, 5: 655 (1945). cymatoides Kze. ex DC. Prodr. 6: 225 (1838). ecuadorense Hieron. in Engler, Bot. Jahrb. 21: 347 (1895). ecuadorense var. boliviense Cuatr. in Pl. Isernianae, 1: 223 (1935). frigidum Wedd. Chlor. And. 1: 147, t. 24A (1856). Gaudichaudiana DC. Prodr. 6: 226 (1838). helichrysoides Wedd. Chlor. And. 1: 146 (1856). Kunthianum (Wedd.) O. Ktze. Rev. Gen. 3(2): 152 (1898). lacteum Meyen & Walp. in Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 276 (1843). melanosphaeroides Sch. Bip. ex Wedd. Chlor. And. 1: 148 (1856). monticola Phil. in Anal. Univ. Chile, 112: 117 (1903). purpureum L. Sp. Pl. 854 (1753). satureioides var. candicans (HBK.) O. Ktze. Rev. Gen. 3(2): 153 (1898). sphacelatum HBK. Nov. Gen. & Spec. 4: 86 (1820). spicatum Lam. Encycl. 2: 757 (1786). tunariense O. Ktze. Rev. Gen. 3(2): 155 (1898 versatile Rusby in Mem. Torr. Bot. Club, 6: 62 (1896). Weddellianum var. nanum Cuatr. in PI. Isernianae, 1: 225 (1935). Gochnatia boliviana Blake in Contrib. U. S. Nat. Herb. 22: a (1924). G. Cardenasii Blake in Journ. Wash. Acad. Sci. 25: 324 (192 G. curviflora (Griseb.) O. Hoffm. in Engl. & Prantl, Nat. Pte 4(5): 337 (1893). G. macrocephala (Rusby) Cabrera in Not. Mus. La Plata, 15, Bot. 74: 41 950). AANAAHANA ARARAARARAD G. Rusbyana Cabrera in Not. Mus. La Plata, 15, Bot. 74: 41 (1950). Grindelia boliviana Rusby in Mem. Torr. Bot. Club, 6: 60 (1896). Gutierrezia Gilliesii var. scabriuscula Griseb. in Goett. Abh. 19: 173 (1874). Gynoxys baccharoides Cass. in Dict. Sci. Nat. 48: 455 (1827). G. boliviana (Klatt) Blake in Contrib. Gray Herb. 53: 28 (1918). 211 cochabambensis Cabrera in Not. Mus. La Plata, 14: 194 (1949). cruzensis Cuatr. in Collect. Bot. [Barcelona] 3(3): 295 (1953). discolor Rusby in Bull. N. Y. Bot. Gard. 4: 398 (1907). folfosa (Rusby) Blake in Contrib. U. S. Nat. Herb. 24: 86 (1922). glabriuscula Rusby in Mem. Torr. Bot. Club, 6: 68 (1896). Hallii Hieron. in Engler, Bot. Jahrb. 19: 64 (1894). Hoffmannii O. Ktze. Rev. Gen, 3(2): 156 (1898). hypomalaca Blake in Bot. Gaz. 74: 427 (1922). laurifolia (HBK.) Cass. in Dict. Sci. Nat. 48: 435 (1827), combination im- plied but not actually made. Mandonii Sch. Bip. ex Rusby in Mem. Torr. Bot. Club, 6: 67 (1896). megacephala Rusby in Bull. N. Y. Bot. Gard. 4: 398 (1907). neovelutina Cuatr. in Fieldiana, Bot. 27(2): 11 (1951). psilophylla Klatt in Ann. Naturh. Hofmus. Wien, 9: 367 (1894). repanda Wedd. Chlor. And. 1: 77 (1856). Rusbyi Cuatr. in Fieldiana, Bot. 27(2): 10 (1951 sorataénsis Cuatr. in Fieldiana, Bot. At 12 sieiis . tablaénsis Cabrera in Blumea, 7: 197 52). Haplopappus boliviensis Cabrera in tot ot a 193 (1952). Heliopsis buphthalmoides (Jacq.) Dunal in Mém. Mus. Paris, 5: 57 (1819). Helogyne Fiebrigii Hieron. in Engler, Bot. Jahrb. 40: 368 (1908). H. virgata (Rusby) Robinson in Proc. Am. Acad. 42: 31 (1906). Heterosperma diversifolium HBK. Nov. Gen. & Spec. 4: 246 (1820). H. maritimum HBK. Nov. Gen. & Spec, 4: 245, t. 383 (1820). H. pinnatum Cav. Icon. 3: 34, t. 267 (1795). H. pinnatum var. biternatum O. Ktze. Rev. Gen. 3(2): 158 (1898). H. rhombifolium Griseb. in Goett. Abh. 19: 187 (1874). Heterotheca deltoidea Klatt in Ann, Naturh. Hofmus. Wien, 9: 358 (1894) = Liabum hastifolium Hieracium ailenocephaluin (Sch. Bip.) Arvet-Touvet, Spicil. 8 (1881). apoloénse Rusby in Bull. N. Y. Bot. Gard. 8: 135 (1912). Bangii Rusby in Mem. Torr. Bot. Club, 3(3): 66 (1898). Bangii subsp. austroboliviense Zahn in Engler, Pflanzenr. IV, 280: 1099 QAANANAAA AnAAgADAD boliviense (Wedd.) Sch. Bip. in Bonplandia, 9: 173 (1861). eriocephalum Wedd. Chlor. And. 1: 226 (1857 Fiebrigianum Zahn in Engler, Pflanzenr. IV, 280: 1098 (1922). fimbriatum Arvet-Touvet, Spicil. 6 (1881). Hauthalianum Zahn in Engler, Pflanzenr. IV, 280: 1090 (1922). lagopus D. Don in Trans. Linn. Soc. 16(2): 176 (1830). leptocephalum var. microcephalum (Sch. Bip.) Zahn in Engler, Pflanzenr. IV, 280: 1096 (1922). Mandonii (Sch. Bip.) Arvet-Touvet, Spicil. 16 (1881). mapirense Britton in Bull. Torr. Bot. Club, 19: 371 (1892). megalochaetum Zahn in Engler, Pflanzenr. IV, 280: 1144 (1922). pazense Blake in Bot. Gaz. 74: 429 (1922). stachyoideum ahi Touvet, Spicil. 21 (1881). strigosum D. Don in Trans. Linn, Soc. 16: 175 (1830). tacense Hieron. in Engler, Bot. Jahes. 21: 375 (1896). trichodontum (Sch. Bip.) Arvet-Touvet, Spicil. 16 (1881). trichodontum Rusby in Bull. N. Y. Bot. Gard. 4: 402 (1907). Trollii Sleumer in Fedde, Repert. Spec. Nov. 41: 119 (1936). Hyalis lancifolia Baker in Mart. Fl. Bras. 6(3): 368 (1884). Hyaloseris boliviensis Koster in Blumea, 5: 668 (1945). H. camataquiensis Hieron. ex Fiebrig in Engler, Bot. Jahrb. 45: 43 (1910). H. quadrifiora Koster in Blumea, 5: 667 (1945). bapa as a 212 Hypochoeris acaulis (Remy) Britton in Bull. Torr. Bot. Club, 19: 871 (1892). brasiliensis var. sulfurea O. Ktze. Rev. Gen. 3(2): 159 (1898). chilensis Britton in Bull. Torr. Bot. Club, 19: 371 (1892). elata (Wedd.) Griseb. in Goett. Abh. 24: 218 (1879). Meyeniana (Walp.) Griseb. in Goett. Abh. 19: 199 (1874). Meyeniana var. ciliata (Wedd.) Perkins in Engler, Bot. Jahrb. 49: 232 (1913 ornata Sante in Blumea, 5: 660 (1945). parvifolia Koster in Blumea, 5: 661 (1945). sessiliflora HBK. Nov. Gen. & Spec. 4: 2 (1820). setosa (Wedd.) Rusby in Bull. N. Y. Bot. Gard. 4: 402 (1907). stenocephala (A. Gray) O. Ktze. Rev. Gen. 3(2): 160 (1898). stenocephala var. integrifolia O. Ktze. Rev. Gen. 3(2): 160 (1898). . taraxacoides (Meyen & Walp.) Ball in Journ. Linn. Soc. 22: 48 (1885). Hysterionica nidorelloides (DC.) Baker in Mart. Fl. Bras. 6(3): 12 (1882). Isostigma Herzogii Hass]. in Fedde, Repert. Spec. Nov. 7: 358 (1909). I. Hoffmannii O. Ktze. Rev. Gen. 3(2): 160 (1898). Jaegeria hirta (Lag.) Less. Syn. Comp. 223 (1832). Jungia affinis Gardn. in Hook. Journ. Bot. 6: 460 (1847). divaricata Rusby in Mem. Torr. Bot. Club, 6: 71 (1896). . ferruginea L. f. Suppl. 390 (1781). floribunda Less. in Linnaea, 5: 38 (1830). grossulariaefolia Rusby, Descr. S. Am. Pls. 164 (1920). Herzogiana Beauverd ex Koster in Blumea, 5: 683 (1945). orbicularis Rusby in Bull. N. Y. Bot. Gard. 4: 401 (1907). pauciflora Rusby in Bull. N. Y. Bot. Gard. 4: 401 (1907). polita var. tomentosa O. Ktze. Rev. Gen. 3(2): 161 (1898). pubescens var. boliviensis O. Ktze. Rev. Gen, 3(2): 161 (1898). sordida Koster in Blumea, 5: 683 (1945). Laestadia Lechleri Wedd. Chlor. And. 1: 184 (1856). Lagascea mollis Cav. in Anal. Cienc. Nat. 6: 332, t. 44 (1803). Leontopodium linearifolium (Wedd.) Britton in Bull. Torr. Bot. Club, 19: 148 (1892). Lepidophyllum lucidum (Meyen) Cabrera in Bol. Soc. Argent. Bot. 1: 51 (1945). L. phylicaeforme (Meyen) Hieron. ex Fries in Nov. Act. Soc. Sci. Upsal. (ser. 4) 1(1): 77 (1905). L. quadrangulare (Meyen) Benth. & Hook. f. Gen. Pl. 2(1): 258 (1873). L. teretiusculum O. Ktze. Rev. Gen. 3(2): 162 (1898 L. Tola Cabrera in Bol. Soc. Argent. Bot. 1: 56 (1945). i acuminatum Rusby, Descr. S. Am. Pls. 161 (1920). L. asperifolium Muschl. in Engler, Bot. Jahrb. 50, Beibl. 111: 78 (1918). L. Cardenasii Cabrera in Not. Mus. La Plata, 14: 191 (1949). L. corymbosum Sch. Bip. ex Klatt in Ann. Naturh. Hofmus. Wien, 9: 363 (1894). L. foliosum (Rusby) Cabrera in Not. Mus. La Plata, 14: 193 (1949). L. fulvotomentosum O, Ktze. Rev. Gen. 3(2): 163 (1898). L. giganteum Rusby in Bull. N. Y. Bot. Gard. 4: 391 (1907). L. glandulosum O. Ktze. Rev. Gen. 3(2): 163 (1898). L. hastatum (Wedd.) Britton in Bull. Torr. Bot. Club, 19: 263 (1892). L. hastifolium Poepp. & Endl. Nov. Gen. 3: 43 (1845). L. hexagonum Blake in Journ. Wash. Acad. Sci. 25: 322 (1935) = L. longifoli- teen el ae i att St Se Se um. L. hirtum O. Ktze. Rev. Gen. 3(2): 163 (1898). L. isodontum Blake in Journ. Wash. Acad. Sci. 17: pats (1927). : Jelskii Hieron. in Engler, Bot. Jahrb. 36: 499 (1905). L. longifolium (Rusby) Blake in Journ. Wash. fe Sci. 25: 322 (1935). 2138 L. mee Muschl. in Engler, Bot. Jahrb. 50, Beibl. 111: 85 (1913). L. ovatum (A. Gray) Ball in Journ. Linn. Soc. 22: 46 (1885). L. oan var. hirtum Perkins in Engler, Bot. Jahrb. 49: 229 (1913). L. pinnulosum O. Ktze. Rev. Gen. 3(2): 163 (1898). L. Rusbyi Britton in Bull. Torr. Bot. Club, 19: 263 (1892). L. subviride Blake in Journ. Wash. Acad. Sci. 17: 294 (1927). L. uniflorum (Poepp. & Endl.) Sch. Bip. in Flora, 36: 34 (1853). Lophopappus cuneatus R. E. Fries in Ark. Bot. 5(13): 29, t. 1 (1906). L. foliosus Rusby in Bull. Torr. Bot. Club, 21: 487 (1894). Loricaria graveolens Wedd. Chlor. And. 1: 167, t. 27C (1856). L. thuyoides (Lam.) Sch. Bip. in Bonplandia, 8: 260 (1860). L. unduaviensis Cuatr. in Fedde, Repert. Spec. Nov. 56: 170 (1954). Lucilia affinis Wedd. Chlor. And. 1: 230 (1857). L. flagelliformis Wedd. Chlor. And. 1: 157, t. 26D (1856). L. Jamesonii Baker in Mart. Fl. Bras. 6(3): 113 (1882). L. recurva Wedd. Chlor. And. 1: 156, t. 25B (1856). L. Schultzii (Wedd.) A. Gray in Proc. Am. Acad. 5: 138 (1862). L. squarrosa Baker in Mart. Fl. Bras. 6(3): 114 (1882). L. subspicata (Wedd.) Hieron. in Engler, Bot. Jahrb. 29: 29 (1900). L. tomentosa Wedd. Chlor. And 1: 157 (1856). L. tunariensis (O. Ktze.) K. Schum. in Just, Bot. Jahresber. 26(1): 378 (1900). L. violacea Wedd. Chlor. And. 1: 155 (1856). Luciliopsis perpusilla Wedd. Chlor. And. 1: 160, t. 26A (1856). Lycoseris boliviana Britton in Bull. Torr. Bot. Club, 19: 266 (1892). L. retroflexa Koster in Blumea, 5: 664 (1945). Mikania baccharoidea Rusby in Bull. N. Y. Bot. Gard. 8: 127 (1912). M. Buchtienii Robinson in Contrib. Gray Herb. 64: 7 (1922). M. cinnamomifolia Lingelsh. in Fedde, Repert. Spec. Nov. 7: 251 (1909). M. cochabambana Robinson in Contrib. Gray Herb. 90: 33 (1930). M. comarapana Robinson in Contrib. Gray Herb. 90: 34 (1930). M. cordifolia (L. f.) Willd. Sp. Pl. 3(1): 1746 (1804). M. decora Poepp. in Poepp. & Endl. Nov. Gen. 3: 53 (1845). M. desmocephala Robinson in Contrib. Gray Herb. 64: 7 (1922). M. dictyota Robinson in Contrib. Gray Herb. 68: 37 (1923). M. dioscoreoides (Rusby) Robinson in Contrib. Gray Herb. 64: 97 (1922). M. eucosma Robinson in Contrib. Gray Herb. 64: 9 (1922). M. ferruginea (Rusby) Rusby ex Robinson in Contrib. Gray Herb. 64: 101 (1922). M. ferruginea var. subglabra Robinson in Contrib. Gray Herb. 104: 36 (1934). M. Fiebrigii Hieron. in Engler, Bot. Jahrb. 40: 390 (1908). M. flaccida Robinson in Contrib. Gray Herb. 64: 9 (1922). M. Guaco Humb. & Bonpl. Pl. Aequin. 2: 84, t. 105 (1809). M. Haenkeana DC. Prodr. 5: 196 (1836). M. lanuginosa DC. Prodr. 5: 201 (1836). M. leucophylla (Rusby) Robinson in Proc. Am. Acad. 47: 196 (1911). M. longiacuminata (Rusby) Rusby ex Robinson in Contrib. Gray Herb. 64: 103 1922). M. longiflora (Rusby) Robinson in Proc. Am. Acad. 47: 196 (1911). M. Mandonii Sch. Bip. ex Baker in Mart. Fl. Bras. 6(2): 188 (1876) = Ophryosporus piquerioides. ’ M. micrantha HBK. Nov. Gen. & Spec. 4: 134 (1820). M. micrantha forma congesta (DC). Robinson in Contrib. Gray Herb. 64: 43 (1922). M. microptera DC. Prodr. 5: 196 (1836). M. officinalis Mart. Reise, 1: 283 (1823). M. oreimeles Robinson in Contrib. Gray Herb. 68: 39 (1923). bo ee ~ = 5 i gS Pennellii Robinson in Contrib. Gray Herb. 61: 19 (1920). periplocifolia Hook. & Arn. in Hook. Compan. Bot. Mag. 1: 243 (1836). phyllopoda Griseb. in Goett. Abh. 19: 170 (1874). platyphylla DC. Prodr. 5: 195 (1836). psilostachya DC. Prodr. 5: 190 (1836). rubella Lingelsh. in Fedde, Repert. Spec. Nov. 7: 250 (1909). Rusbyi Robinson in Contrib. Gray Herb. 64: 18 (1922) . Schultzii Robinson in Contrib. Gray Herb. 64: 19 (1922). sinuata Rusby in Bull. N. Y. Bot. Gard. 8: 127 (1912) = M. micrantha. speciosa DC. Prodr. 5: 196 (1836). Steinbachii Robinson in Contrib. Gray Herb. 80: 40 (1928). stygia Robinson in Contrib. Gray Herb. 90: 36 (1930). trifolia (Rusby) Robinson in Contrib. Gray Herb. 64: 107 (1922). vitifolia forma boliviensis (Lingelsh.) Robinson in Contrib. Gray Herb. 104: 54 (1934). Werdermannii Robinson in Contrib. Gray Herb. 104: ve (1934). . Williamsii Robinson in Contrib. Gray Herb. 64: 19 (19 Montanoa Orbignyana Klatt in Abh. Naturf. Ges. Halle, ~ 328 (1881). Moquinia boliviana Rusby in Bull. N. Y. Bot. Gard. 4: 399 (1907) = Gochnatia us usbyana. ies Bipontina Sch. Bip. ex Rusby in Mem. Torr. Bot. Club, 6: 68 (1896). SS BSSSSSSS 55555555 mptosorifolia Rusby in Mem. Torr. Bot. Club, 4(8): 213 (1895) = M. comhabeatenaii Clematis L. f. Suppl. 373 (1781). cochabambensis Hieron. in Engler, Bot. Jahrb. 19: 74 (18 comptoniaefolia Rusby in Mem. Torr. Bot. Club, 3(3): 65 meseo8y Flagellaria Koster in Blumea, 5: 672 (1945 hastata Cav. Icon. 5: 64, t. 494 (1799). homoeantha Wedd. Chlor. And. 1: 21, t. 2A (1855). Isernii Phil. in Anal. Univ. Chile, 85: 823 (1894). lanigera Wedd. Chlor. And. 1: 16 (1855). ledifolia Decne. ex Wedd. Chlor. And. 1: 20 (1855). ledifolia forma glabrata Cuatr. Pl. Isernianae, 1: 236 (1935). ledifolia forma longifiora Koster in Blumea, 5: 673 (1945). Orbignyana Wedd. Chlor. And. 1: 22 (1855). subulata R. & P. Syst. 193 (1798). Vicia Koster in Blumea, 5: 670 (1945). viciaefolia Cav. Icon. 5: 62, t. 490 (1799). viciaefolia var. Candolleana (Gardn. & Field) Wedd. Chlor. And. 1: 15 (1855). viciaefolia var. hirsuta (Meyen ex Walp.) Wedd. Chlor. And. 15 (1855). viciaefolia forma intermedia Cuatr. Pl. Isernianae, 1: 287 (193 Nardophyllum armatum (Wedd.) Reiche in Anal. Univ. Chile, arte "29 (1901). Onoseris acerifolia HBK. Nov. Gen. & Spec. 4: 8 (1820). Sfoo9999 alata Rusby, Descr. S. Am. Pls. 163 (1920). albicans (D. Don) Ferreyra in Journ. Arnold Arb. 25: 389 (1944). fraterna Blake in Journ. Wash. Acad. Sci. 33: 368 (194 3). gnaphalioides Muschl. in Engler, Bot. Jahrb. 50, Beibl. 111: 94 (1913). hastata Wedd. Chlor. And. 1: 9, t. 7 (1855). hyssopifolia HBK. Nov. Gen. & Spec. 4: 9, t. 306 (1820). sagittata (Rusby) Rusby, Deser. S. Am. Pls. 164 (1920). Ophryosporus angustifolius Robinson in Contrib. Gray Herb. 90: 3 (1930). Oo. Cumingii (Sch. Bip.) Benth. ex Baker in Mart. Fl. Bras. 6(2): 188 (1876). O. eleutheranthus (Rusby) Robinson in Contrib. Gray Herb. 61: 27 (1920). O. Kuntzei Hieron. in Engler, Bot. Jahrb. 22: 707 (1897). 215 origanoides var. microcephalus Hieron. in Engler, Bot. Jahrb. 22: 708 (1897). piquerioides (DC.) Benth. ex Baker in Mart. Fl. Bras. 6(2): 188 (1876). . Steinbachii Robinson in Contrib. Gray Herb. 77: 5 (1926). - venosissimus (Rusby) Robinson in Proc. Am. Acad. 42: 24 (1906). Oyedaea boliviana Britton in Bull. Torr. Bot. Club, 19: 149 (1892). O. bullata Koster in Blumea, 6: 269 (1948). O. lanceolata (Rusby) Blake in Contrib. U. S. Nat. Herb. 20: 416 (1921). O. Pearcei Rusby in Mem. Torr. Bot. Club, 3(3): 59 (1893) = O. boliviana. O. Rusbyi Blake in Contrib. U. S. Nat. Herb. 20: 416 (1921). Pappobolus macranthus Blake in Hook. Ic. Pl. 31: t. 83057 (1916). Parthenium cineraceum Rollins in Contrib. Gray Herb. 172: 32 (1950). P. glomeratum Rollins in Contrib. Gray Herb. 172: 59 (1950). P. Hysterophorus L. Sp. Pl. 988 (1758). Pectis linifolia L. Sp. Pl. (ed. 2) 2: 1150 (1768). P. odorata Griseb. in Goett. Abh. 24: 200 (1879). P. sessiliflora Sch. Bip. ex Rusby in Mem. Torr. Bot. Club, 3(3): 62 (1893). P. substriata Rusby in Bull. N. Y. Bot. Gard. 4: 390 (1907). P. Swartziana Less. in Linnaea, 6: 711 (1831). Perezia aracensis Koster in Blumea, 5: 678 (1945). P. coerulescens Wedd. Chlor. And. 1: 39, t. 10A (1855). P. cirsiifolia Wedd. Chlor. And. 1: 41 (1855). P. elongata O. Ktze. Rev. Gen. 3(2): 166 (1898). P. foliosa Rusby in Mem. Torr. Bot. Club, 6: 71 (1896). P. glomerata Rusby in Mem. Torr. Bot. Club, 4(3): 214 (1895). P. integrifolia Wedd. Chlor. And. 1: 40 (1855). P. laurifolia O, Ktze. Rev. Gen. 3(2): 166 (1898). P. Mandonii Rusby in Mem. Torr. Bot. Club, ge sho (1893). P. multiflora (HBK.) Less. in Linnaea, 5: 19 (1830). P. nitidifolia Koster in Blumea, 5: 677 (1945). P. obtusisquama Koster in Blumea, 5: 680 (1945). P. pungens (HBK.) Less. in Linnaea, 5: 20 (1830). P. pungens var. cernua Rusby in Mem. Torr. Bot. Club, 6: 70 (1896). P. purpurata Wedd. Chlor. And. 1: 43 (1855). P. pygmaea Wedd. Chlor. And. 1: 40 (1855). P. scalpellifolia Koster in Blumea, 5: 680 (1945). P. scalpellifolia var. parvifolia Koster in Blumea, 5: 680 (1945). P. violacea Wedd. Chlor. And. 1: 42 (1855). Picrosia longifolia D. Don in Trans. Linn. Soc. 16: 184 (1830). Piptocarpha laxa Rusby in Bull. N. Y. Bot. Gard. 8: 123 (1912). P. Lechleri (Sch. Bip.) Baker in Mart. Fl. Bras. oe 127 (1873). Plagiochilus ciliaris Wedd. Chlor. And. 1: 227 (1857). r. phy Rusby in Mem. Torr. Bot. Club, 4(3): 212 (1895) — Chrysanthel- lum ? P; slleker ante DC. Prodr. 6: 142 (183 Plazia acaciifolia Koster in Blumea, 5: ed (1945). P. argentea (D. Don) O. Ktze. Rev. Gen. 3(2): 167 (1898). P. daphnoides var. glabrescens Wedd. Chlor. And. 1: 13, t. 2B (1855). P. spartioides (Wedd.) O. Ktze. Rev. Gen. 3(2): 167 (1898). Pluchea fastigiata Griseb. in Goett. Abh. 24: 184 (1879). P. glabra Griseb. in Goett. Abh. 24: 183 (1879). P. odorata var. ferruginea Rusby in Mem. Torr. Bot. Club, 4(3): 211 (1895). P. suaveolens (Vell.) O. Ktze. Rev. Gen. 3(2): 168 (1898). Podocoma hieracifolia (Poir.) Cass. in Dict. Sci. Nat. 42: 60 (1826). Polyachyrus oblongifiorus Koster in Blumea, 5: 675 (1945). Polymnia edulis Wedd. in Ann. Sci. Nat. (ser. 4) 7: 114 (1857). 299 © 216 P. glabrata DC. Prodr. 5: 515 (1836). P. Siegesbeckia DC. Prodr. 5: 515 (1836). Porophyllum lanceolatum DC. Prodr. 5: 649 (1836). P. oblanceolatum Rusby in Mem. Torr. Bot. Club, 6: 64 (1896). P. platyphyllum Chod. in Bull. Herb. Bois. (ser. 2) 2: 397 (1902). P. ruderale (Jacq.) Cass. in Dict. Sci. Nat. 43: 56 (1826). P. ruderale var. angustifolium Hassl. in Trab. Mus. Farm. Med. Buenos Aires, 21: 181 (1909). P. ruderale var. at aphae (Cass.) Gray ex Robinson & Greenman in Proc. Am. Acad. 49: 509 (1913). Proustia angustifolia Wedd. Chlor. And. 1: 24 (1855). P. angustifolia var. mollis O. Ktze. Rev. Gen. wc 168 (1898). P. pungens Poepp. ex Less. Syn. Comp. 110 (18 P. pungens var. cuneifolia (Don) Wedd. Chlor. sa 1: 23 (1855). P. pungens var. oblongifolia Wedd. Chlor. And. 1: 23 (1855). Pseudobaccharis acaulis (Wedd.) Cabrera in Not. Mus. La Plata, 9: 248 (1944). P. boliviensis (Wedd.) Cabrera in Not. Mus. La Plata, 9: 249 (1944). Pseudogynoxys Benthamii Cabrera in Brittonia, 7: 56 (1950). Pterocaulon alopecuroideum (Lam.) DC. Prodr. 5: 454 (1836). . lanatum O. Ktze. Rev. Gen. 3(2): 169 (1898). P. latifolium O. Ktze. Rev. Gen. 3(2): 169 (1898). P. Lorentzii Malme in Kgl. Sv. Vet. Akad. Handl. 27: 22 (1901). P. purpurascens Malme in Kgl. Sv. Vet. Akad. Handl. 32(5): 55 (1899), no- men a") P. virgatum DC. Prodr. 5: 454 (1836). Salmea scandens (L.) DC. Prodr. 5: 493 (1836). Sanvitalia versicolor Griseb. in Goett. Abh. 24: 189 (1879). Schistocarpha Hoffmannii O. Ktze. Rev. Gen. 3(2): 170 ae S. paniculata Klatt in Bull. Soc. Bot. Belg. 31: 210 (189 S. triangularis Rusby in Bull. N. Y. Bot. Gard. 4: 392 las 7). Schkuhria degenerica (O. Ktze.) R. E. Fries in Ark. Bot. 5(18): 22 (1906). . multiflora Hook. & Arn. in Journ. Bot. 3: 322 (1841). mutiflora var. pusilla (Wedd.) Cabrera in Anal. Soc. Cient. Argent. 114: 192 (1932). pinnata (Lam.) O. Ktze. Rev. Gen. 3(2): 170 (1898), in synon. pinnata ee abrotanoides (Roth) Cabrera in Anal. Soc. Cient. Argent. 114: 189 (19 S. pinnata var. otemeliaie (DC.) Cabrera in Anal. Soc. Cient. Argent. 114: 190 (1982). Senecio adamantinus Bong. in Bull. Sc. Acad. Pétersb. 5: 97 (1838). adenophyllus Meyen & Walp. in Nov. Act. Acad. Caes. Leop. 19, Suppl. 1; 282 (1843). agapatensis Sch. Bip. in Bonplandia, 4: 55 (1856), nomen nudum. algens Wedd. Chlor. And. 1: 104 (1856). alternifolius (Rusby) Greenm. in Ann. Missouri Bot. Gard. 10: 76 (1923). amphibolus Wedd. Chlor. And. 1: 109 (1856). attenuatus Sch. Bip. ex Rusby in Mem. Torr. Bot. Club, 3(3): 63 (1893). attenuatus var. microphyllus Britton in Bull. Torr. Bot. Club, 19: 264 (1892) = S. clavifolius. ayopayensis Cuatr. in Fieldiana, Bot. 27 (2): 50 (1951). baccharidifolius Rusby in Bull. N. Y. Bot. Gard. 4: 397 (1907). Bangii Rusby in Mem. Torr. Bot. Club, 3(3): 64 (1893). biacuminatus Rusby in Bull. N. Y. Bot. Gard. 4: 394 (1907). biserrifolius O. Ktze. Rev. Gen. 3(2): 171 (1898). boliviensis Sch. Bip. ex Klatt in Leopoldina, 24: 127 (1888). . Brittonianus Hieron. in Engler, Bot. Jahrb. 29: 72 (1900). min min in ERC EC ECE nnn nia tn in nnn In tr 1D th NNANANANANANAMININnN A . pee ty enor PP RERE rer heme ren hes 217 Buchtienii Greenm. in Ann. Missouri Bot. Gard. 10: 78 (1923). Cabrerae Cuatr. in Fieldiana, Bot. 27(2): 61 (1951). Cabrerianus Greenm. & Cuatr. in Fedde, Repert. Spec. Nov. 55: 134 (1953). campanulatus Sch. Bip. ex Klatt in Leopoldina, 24: 126 (1888 campanulatus var. glabrescens Cabrera in Blumea, 7: 198 (1952). canescens (Humb. & Bonpl.) Cuatr. in Fieldiana, Bot. 27: 43 (1950). canescens var. monocephalus (Wedd.) Cuatr. in Fieldiana, Bot. 27(2): 37 (1951). Cardenasii Cuatr. in Fieldiana, Bot. 27(2): 48 (1951). Chaenocephalus Cabrera in Not. Mus. La Plata, 9: 192 (1944). charaguensis Cuatr. in Fieldiana, Bot. 27(2): 59 (1951). Chodatianus Cuatr. in Collect. Bot. [Barcelona] 3(3): 278 (1953). clavifolius Rusby in Mem. Torr. Bot. Club, 3(3): 64 (1893). clivicola Wedd. Chlor. And. 1: 130 (1856). cochabambensis Cabrera in Blumea, 7: 201 (1952). colaminus Cuatr. in Fieldiana, Bot. 27(2): 58 (1951). comarapensis Cabrera in Darwiniana, 10: 599 (1954). coroicensis Rusby in Bull. N. Y. Bot. Gard. 4: 395 wre culcitioides Sch. Bip. ex Wedd. Chlor. And. 1: 103 (185 dictyophlebius Greenm. in Ann. Missouri Bot. Gard. ste 801 (1938). epiphyticus O. Ktze. Rev. Gen. 3(2): 173 (1898). expansus Wedd. Chlor. And. 1: 107 (1856). evacoides Sch. Bip. in Bonplandia, 4: 52 (1856). floscosus Britton in Bull. Torr. Bot. Club, 19: 264 (1892). formosus HBK. Nov. Gen. & Spec. 4: 177 (1820). glacialis Wedd. Chlor. And. 1: 113, t. 18A (1856). glacialis var. adenophylloides (Sch. Bip.) Perkins in Engler, Bot. Jahrb. 49: 230 (1913). glacialis (Meyen & Walp.) Cuatr. in Fieldiana, Bot. 27: 44 (1950). graveolens Wedd. Chlor. And. 1: 111 (1856). helianthemoides Wedd. Chlor. And. 1: 115 (1856). Herzogii Cabrera in Blumea, 7: 202 (1952). heterotrichius DC. Prodr. 6: 419 (1838). Hohenackeri Sch. Bip. ex Wedd. Chlor. And. 1: 131 (1856). Hualtata Bert. ex DC. Prodr. 6: 417 (1838 humillimus var. vegetus Wedd. Chlor. And. "44 104 (1856). icoglossus var. araneosus DC. Prodr. 6: 420 (1838). innovans Klatt in Ann. Naturh. Hofmus. Wien, 9: 365 (1894). Jarae Phil. in Anal. Mus. Nac. Chile, Bot. 8: 44 (1891). Klattii Greenm. in Ann. Missouri Bot. Gard. 1: 281 (1914). Kosterae Cabrera in Blumea, 7: 201 (1952). Krukoffii Cuatr. in Fieldiana, Bot. 27(2): 60 (1951). leuceria Cabrera in Blumea, 7: 202 (1952 liabifolius Rusby in Bull. N. Y. Bot. Gard. 4: 396 (1907). longilinguae Cuatr. in Fieldiana, Bot. 27(2): 51 (1951). Mandonianus Wedd. Chlor. And. 1: 228 (1857). medullosus Sch. Bip. ex Greenm. in Ann. Missouri Bot. Gard. 10: 85 (1923). melanolepis DC. Prodr. 6: 424 (1838). Miguelii Cuatr. in Fieldiana, Bot. 27(2): 54 sn modestus Wedd. Chlor. And. 1: 105, t. 18B (1856). multinervis Sch. Bip. ex Klatt in Leopoldina, 24: 127 (1888). myrianthus Klatt in Leopoldina, 24: 127 (1888). oblanceolatus Rusby in Bull. N. Y. Bot. Gard. 4: 394 (1907). octophyllus Sch. Bip. ex Rusby in Bull. N. Y. Bot. Gard. 4: 393 (1907). oronocensis DC. Prodr. 6: 423 (1837). pampae Lingelsh. in Fedde, Repert. Spec. Nov. 8: 6 (1910). 218 pampae var. penicillatus Lingelsh. in Fedde, Repert. Spec. Nov. 8: 6 (1910). pectioides Rusby in Bull. IN. Y. Bot. Gard. 4: 395 ( 1907). pensilis Greenm. in Ann. Missouri Bot. Gard. 25: 813 (1938). pentamerus Cuatr. in Fieldiana, Bot. 27(2): 57 (1951). Pentlandianus DC. Prodr. 6: 421 (1838). Pflanzii (Perkins) Cuatr. in Fieldiana, Bot. 27: 44 (1950). pongoénsis Cuatr. in Fieldiana, Bot. 27(2): 61 (1951). nearaae Klatt in Abh. Naturf. Ges. Halle, 15: 331 (1881). praeru m Sch. Bip. ex Klatt in Leopoldina, 24: 127 (1888). prunifolius = Wedd. Chlor. And. 1: 102 (1856). prunioides Rusby in Bull. N. Y. Bot. Gard. 4: 396 (1907). psidiifolius Rusby in Mem. Torr. Bot. Club, 6: 66 (1896). psychrophilus Wedd. Chlor. And. 1: 112 (1856). pulviniformis Hieron. in Engler, Bot. Jahrb. 21: 359 (1895). Reicheanus Cabrera in Lilloa, 15: 403 (1949). repens DC. Prodr. 6: 423 (1838). rhizomatus Rusby in Mem. Torr. Bot. Club. 6: 66 (1896). . rudbeckiaefolius Meyen & Walp. in Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 283 (18438). rufescens (Humb. & Bonpl.) Cuatr. in Fieldiana, Bot. 27: 45 (1950). sailapatensis Cuatr. in Fieldiana, Bot. 27(2): 52 (1951). senecioides (HBK.) O. Ktze. Rev. Gen. 3(2): 177 (1898). Sepium Sch. Bip. ex Rusby in Bull. N. Y. Bot. Gard. 4: 394 (1907). serratifolius (Meyen & Walp.) Cuatr. in Fieldiana, Bot. 27: 45 (1950). sinapoides Rusby in Mem. Torr. Bot. Club, 6: 65 (1896) Smithii DC. Prodr. 6: 412 (1838). i Britton in Bull. Torr. Bot. Club, 19: 265 (1892) = S. Brittonianus. Steinbachianus Cuatr. in Fieldiana, Bot. 27(2): 55 (1951). stylotrichus Cabrera in Not. Mus. La Plata, 15: 107 ports subdecurrens Sch. Bip. ex Wedd. Chlor. And. 1: 109 (1856). subglomerosus Greenm. in Ann. Missouri Bot. Gard. 10: 93 (1923). subvulgaris O. Ktze. Rev. Gen. 3(2): 178 (1898). tabacifolius Rusby in Bull. N. Y. Bot. Gard. 4: 396 (1907). tablensis Cabrera in Blumea, 7: 203 (1952). tenuicaulis Sch. Bip. ex Klatt in Leopoldina, 24: 128 (1888). tephrosioides Turcz. in Bull. Soc. Nat. Mose. 24(2): 92 (1851). unduavianus Cuatr. in Fieldiana, Bot. 27(2): 49 (1951). viridilacus Cabrera in Blumea, 7: 199 (1952). vulgaris L. Sp. Pl. 867 (1753). Weddellii Cabrera in Not. Prelim. Mus. La Plata, 3: 122 (19384). Wuliamsti Rusby in Bull. N. Y. Bot. Gard. 8: 184 (1912) = S. Cabrerianus. yungasensis Britton in Bull. Torr. Bot. Club, 19: 264 (1892). . yurensis Rusby in Bull. N. Y. Bot. Gard. 8: 133 (1912). Siegesbeckia flosculosa L’Hérit. Stirp. Nov. 37, t. 19 (1786). S. orientalis L. Sp. Pl. 900 (1753). Solidago microglossa DC. Prodr. 5: 332 (1836). S. polyglossa DC. Prodr. 5: 332 (1836). Sonchus asper (L.) Vill. Hist. Pl. Dauph. 3: 158 (1789). S. oleraceus L. Sp. Pl. 794 (1753). Sphaereupatorium Hoffmannii O. Ktze. Rev. Gen. 3(2): 147 (1898). Spilanthes Acmella L. Mant. 2: 475 (1771). S. ocymifolia forma radiifera A. H. Moore in Proc. Am. Acad. 42: 533 (1907). Stemodontia elongata Rusby in Mem. Torr. Bot. Club, 3(3): 58 (1893) = Heliopsis buphthalmoides. Stevia Bangii Rusby in Mem. Torr. Bot. Club, 4(3): 209 (1895). S. Bangii var. dyscrita Robinson in Contrib. Gray Herb. 100: 3 (1932). DNANNANRNANNAMAIN NA In ANRNANANAAANANNANADN InN tA In RANNNANANANAANNANANNARNAR ANNANNNANANRNNANANNDNANNANN 219 Benderi Perkins in Engler, Bot. Jahrb. 49: 221 (1913). Benderi var. Cardenasii Robinson in Contrib. Gray Herb. 104: 7 (1934). bermejensis Hieron. in Engler, Bot. Jahrb. 40: 361 (1908). boliviensis Sch. Bip. ex Griseb. in Goett. Abh. 24: 166 (1879). brevipapposa Hieron. in Engler, Bot. Jahrb. 22: 718 (1897) = S. boliviensis. Bridgesii Rusby in Bull. N. Y. Bot. Gard. 4: 377 (1907). calderillensis Hieron. in Engler, Bot. Jahrb. 40: 356 (1908). camachensis Hieron. in Engler, Bot. Jahrb. 40: 359 (1908). cardiatica Perkins in Engler, Bot. Jahrb. 49: 222 (1913). chacoénsis R. E. Fries in Ark. Bot. 5(13): 7, t. 2 (1906). Chamaedrys Griseb. in Goett. Abh. 24: 167 (1879). clivicola Robinson in Contrib. Gray Herb. 100: 4 (19382). cochabambensis Hieron. in Engler, Bot. Jahrb. 22: 726 (1897). copiosa Koster in Blumea, 5: 645 (1945). discolor Robinson in Contrib. Gray Herb. 100: 5 (1982). eclipes Robinson in Contrib. Gray Herb. 96: 5 (1931). elatior var. austrina Robinson in Contrib. Gray are 96: 7 (1931). Fiebrigii Hieron. in Engler, Bot. Jahrb. 40: 365 (1908). filipes Rusby in Bull. N. Y. Bot. Gard. 8: 126 (1912). fruticosa Griseb. in Goett. Abh. 24: 167 (1879). galeopsidifolia Hieron. in Engler, Bot. Jahrb. 22: 719 (1897). glanduloso-pubescens Hieron. in Engler, Bot. Jahrb. st 860 (1908). glomerata Hieron. in Engler, Bot. Jahrb. 40: 357 (190 grandidentata Sch. Bip. ex Rusby in Mem. Torr. Bot. eis 6: 55 (1896) = S. soratensis. humilis Hieron. in Engler, Bot. Jahrb. 22: 730 (1897) = S. obovata. kuhnioides Rusby ex Robinson in Contrib. Gray Herb. 96: 7 (19381). Kuntzei Hieron. in Engler, Bot. Jahrb. 22: 733 (1897). Mandonii Sch. Bip. ex Robinson in Contrib. Gray Herb. 77: 6 (1926). melancolica Robinson in Contrib. Gray Herb. 90: 15 (1930). mercedensis Hieron. in Engler, Bot. Jahrb. 22: 735 (1897). mercedensis var. glanditecta Robinson in Contrib. Gray Herb. 90: 16 (1930). neglecta Rusby in Mem. Torr. Bot. Club. 4(3): 209 (1895). obovata Rusby in Mem. Torr. Bot. Club, 6: 55 (1896). obovata var. aristifera Robinson in Contrib. Gray Herb. 90: 17 (1930). pauciflora Koster in Blumea, 6: 266 (1948). reclinata Rusby in Bull. N. Y. Bot. Gard. 8: 127 (1912). samaipatensis Robinson in Contrib. Gray Herb. 96: 15 (1931). santacruzensis Hieron. in Engler, Bot. Jahrb. 22: 731 (1897). sarensis Robinson in Contrib. Gray Herb. 100: 9 (1932). sarensis var. dissiticeps Robinson in Contrib. Gray Herb. 100: 10 (1932). Schultzii Hieron. in Engler, Bot. Jahrb. 22: 721 (1897) = S. boliviensis. setifera Rusby ex Robinson in Contrib. Gray Herb. 100: 10 (1932). soratensis Hieron. in Engler, Bot. Jahrb. 40: 356 (1908) soratensis var. mecoyensis Robinson in Contrib. Gray Herb. 100: 11 (1932). gprs ee subeglandulosa (Hieron.) Hieron. in Engler, Bot. Jahrb. 40: 359 (19 Stuebelii ts in Engler, Bot. Jahrb. 21: 328 (1895). tapacariensis Hieron. in Engler, Bot. Jahrb. 22: 734 (1897) = S. Bangii. tarijensis Hieron. in Engler, Bot. Jahrb. 40: 362 (1908). totorensis Robinson in Contrib. Gray Herb. 96: 16 (1931). triaristata Hieron. in Engler, Bot. Jahrb. 40: 358 (1908). tunariensis Hieron. in Engler, Bot. Jahrb. 22: 713 (1897). urticaefolia var. boliviensis (Hieron.) Robinson in Contrib. Gray Herb. 100: 44 (1932 220 S. urticaefolia var. pailidiflora Robinson in Contrib. Gray Herb. 96: 16 (1931). S. vaccinioides Koster in Blumea, 5: 646 (1945). S. yaconensis var. subeglandulosa Hieron. in Engler, Bot. Jahrb. 40: 366 (190 8). Stuckertiella capitata (Wedd.) Beauverd in Bull. Soc. Bot. Genéve (ser. 2), (1913). 5: 206 Synedrellopsis Grisebachii Hieron. ex O. Ktze. Rev. Gen. one 180 (1898). Tagetes daucoides Schrad. Ind. Sem. Hort. Govt 5 (1833). SHAS erythrocephala Rusby in Bull. N. Y. Bot. Gard. 8: 133 (1912). gigantea Carr. in Rev. Hort. 58: 107 (1886). graveolens L’Hérit. in DC. Prodr. 5: 644 (1836). Mandonii Sch. Bip. ex Klatt in Leopoldina, on 109 (1889). maxima O. Ktze. Rev. Gen. 3(2): 181 (1898 micrantha Cav. Icon, 4: 31, t. 352 (1797). multiflora HBK. Nov. Gen. & Spec. 4: 197 (1820). pusilla HBK. Nov. Gen. & Spec. 4: 194 (1820). silenoides Meyen & Walp. in Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 272 (1843). Tessaria absinthoides (Hook. & Arn.) DC. Prodr. 5: 457 (1836). <3 integrifolia R. & P. Syst. 213 (1798). Trichocline incana (Lam.) Cass. in Dict. Sci. Nat. 55: 216 < ye reptans (Wedd.) Robinson in Proc. Am. Acad. 49: 515 (1913 Trichogonia capitata (Rusby) Robinson in Proc. Am. Acad. 47: "193 (1911). Tridax boliviensis (Wedd.) R. E. Fries in Ark. Bot. 5(13): 21 (1906). - procumbens L. Sp. Pl. 900 (1753). Trixis aggregata Rusby in Mem. Torr. Bot. Club, 6: 71 (189 a 3 i zs 7 T. 6). antimenorrhoea var. heterophylla O. Ktze. Rev. Gen. 3(2): 183 (1898). antimenorrhoea var. petiolata O. Ktze. Rev. Gen. 3(2): 183 (1898). diffusa Rusby in Bull. N. Y. Bot. Gard. 8: 184 (1912). divaricata (HBK.) Spreng. Syst. Veg. 3: 501 (1826). rigida Koster in Blumea, 5: 682 (1945) Verbesina australis Baker in Mart. Fl. Bras. 6(3): 215 (1884). 2. V. y. . Ndds <444 444462; ss © Benderi Perkins in Engler, Bot. Jahrb. 49: 227 (1913). boliviana Klatt in Ann. Naturh. Hofmus. Wien, 9: 361 (1894). Bridgesii Rusby in Mem. Torr. Bot. Club, 4(3): 212 (1895) = V. boliviana. . cinerea Rusby in Mem. Torr. Bot. Club, 6: 63 (1896). . Cumingii Sch. Bip. ex Blake in Amer. Journ. — 12: 634 (1925). . elegans HBK. Nov. Gen. & Spec. 4: 204 (182 flavovirens R. E. Fries in Ark. Bot. 5(13): ‘ 3 (1906). leucactinota Robinson in Proc. Am. Acad. 47: 213 (1911). macrophylla (Cass.) Blake in Bull. Torr. Bot. Club, 51: 430 (1924). Mandonii Sch. Bip. ex Robinson & Greenm. in Proc. Am. Acad. 34: 547 (1899). Pflanzii Perkins in Engler, Bot. Jahrb. 49: 227 (1913): rhomboidea Koster in Blumea, 6: 270 (1948). semidecurrens O. Ktze. Rev. Gen. 3(2): 183 (1898). soratae Sch. Bip. ex Robinson & Greenm. in Proc. Am. Acad. 34: 551 (1899 suboordate DC. Prodr. 5: 614 (1836). rnonia apiculata Mart. ex DC. Prodr. 5: 51 (1836). arborescens Sw. FI. Ind. Occ. 3: 1320 (1806). arborescens var. cuneifolia Britton in Bull. Torr. Bot. Club, 18: 331 (1891) a V, Rue argyropappa Buck, Index Prodr. 2: v (1840). aristosquamea Britton in Bull. Torr. Bot. Club, 18: 332 (1891) = V. ar- gyropappa ? V. auriculata Griseb. in Goett. Abh. 24: 163 saci 08 V. baccharoides HBK. Nov. Gen. & Spec. 4: 40 (182 V. Bakerana Britton in Bull. Torr. Bot. Club, 18: “Pe (1891). Bangii Rusby in Mem. Torr. Bot. Club, 6: 52 (1896) = V. baccharoides. boliviana Britton in Bull. Torr. Bot. Club, 18: 332 (1891). brachylepis Griseb. in Goett. Abh. 24: 164 (1879). brasiliana (L.) Druce in Rept. Bot. Exch. Club Brit. Isl. 3: 426 (1914). brevipetiolata Sch. Bip. ex Baker in Mart. Fl. Bras. 6(2): 85 (1873). breviramosa Rusby in Bull. N. Y. Bot. Gard. 8: 123 (1912) = V. scorpioides. Buchtienii Gleason in Amer. Journ. Bot. 10: 302 (1923). canaminana Gleason in Amer. Journ. Bot. 10: 309 (1923). centauropsidea Hieron. in Engler, Bot. Jahrb. 40: 353 (1908). Conwayi Rusby in Bull. N. Y. Bot. Gard. 8: 125 (1912). cordiaefolia HBK. Nov. Gen. & Spec. 4: 38 (1820). coriacea Less. in Linnaea, 6: 661 (1831). costata Rusby in Mem. Torr. Bot. Club, 6: 53 (1896). crassifolia Rusby in Bull. N. Y. Bot. Gard. 8: 124 Cation deflexa Rusby in Bull. N. Y. Bot. Gard. 4: 376 (1907). densipaniculata Rusby in Bull. N. Y. Bot. Gard. 8: 126 (1912). digitata Rusby in Bull. N. Y. Bot. Gard. 8: 125 (1912) = V. megaphylla. echitifolia Mart. ex DC. Prodr. 5: 60 (1836). ferruginea Less. in Linnaea, 4: 271 (1829). fulta Griseb. in Goett. Abh. 24: 164 (1879). fulta forma tomentosa Koster in Blumea, 5: 643 (1945). grandiflora Less. in Linnaea, 6: 660 (1831 herbacea (Vell.) Rusby in Mem. Torr. Bot. ‘Club, 4(3): 209 (1895). ixiamensis Rusby in Bull. N. Y. Bot. Gard. 8: 125 (1912). jubifera Rusby in Mem. Torr. Bot. Club, 6: 53 (1896). Kuntzei Hieron. in Engler, Bot. Jahrb. 22: 678 (1897). laurifolia DC. Prodr. 5: 30 (1836). Lehmannii Hieron. in Engler, Bot. Jahrb. 19: 44 (1894). ligulaefolia Mart. ex DC. Prodr. 5: 45 (1836). Mandonii Sch. Bip. ex Gleason in Amer. Journ. Bot. 10: 301 (1923). mapirensis Gleason in Amer. Journ. Bot. 10: 307 (1923). mattogrossensis Hieron. in Engler, Bot. Jahrb. 22: 696 (1897). megaphylla Hieron. in Verh. Bot. Ver. Brandenb. 48: 195 (1907). membranacea Gardn. in Hook. Journ. Bot. 5: 217 (1846). mollis HBK. Nov. Gen. & Spec. 4: 36 (1820). myriocephala DC. Prodr. 5: 40 (1836). obtusata Less. in Linnaea, 6: 662 (1831). patuliflora Rusby in Bull. N. Y. Bot. Gard. 4: 376 (1907) = V. —"* — Rusby in Mem. Torr. Bot. Club, 3(3): 50 (1893) = V. mem branac auceenamats Rusby in Bull. N. Y. Bot. Gard. 4: 376 (1907). pinguis Griseb. in Goett. Abh. 24: 165 (1879). polycephala DC. Prodr. 5: 39 (1836). prenanthoides Gleason in Amer. Journ. Bot. 10: 308 (1923). pseudomollis Gleason in Amer. Journ. Bot. 10: 307 (1923). pycnantha Benth. Pl. Hartw. 134 (1844). remotiflora L. C. Rich. in Act. Soc. Hist. Nat. es 1: 112 (1792). robusta Rusby in Mem. Torr. Bot. Club, 6: 54 (189 rubricaulis HBK. Nov. Gen. & Spec. 4: 33 (1820). rufo-papposa var. latifolia Hieron. in Engler, Bot. Jahrb. 22: 700 (1897) = V. membranacea. Rusbyi Gleason in Amer. Journ. Bot. 19: 753 (19382). saltensis Hieron. in Engler, Bot. Jahrb. 22: 691 (1897). s NRE AI oa ncaa c cs cdeecdeere, s bo i bo santacruzensis Hieron. in Engler, Bot. Jahrb. 22: 699 (1897). scorpioides (Lam.) Pers. Synops. Pl. 2: 404 (1807). scorpioides var. sororia (DC.) Baker in Mart. Fl. Bras. 6(2): 101 (1873). senecionaefolia Britton in Bull. Torr. Bot. Club, 18: 331 (1891) — V. fulta. setososquamosa Hieron. in Engler, Bot. Jahrb. 22: 684 (1897). simplex Less. in Linnaea, 4: 280 (1829). sordido-papposa Hieron. in Engler, Bot. Jahrb. 22: 697 (1897). squamipes Rusby in Bull. N. Y. Bot. Gard, 8: 124 (1912) = V. mem- branacea. squamulosa Hook. & Arn. in Hook. Compan. Bot. Mag. 2: 44 (1836). subacuminata Hieron. in Engler, Bot. Jahrb. 22: 691 (1897). tarijensis (Griseb.) Hieron. in Engler, Bot. Jahrb. 22: 682 (1897). tricholepis DC, Prodr. 5: 54 (1836). tristis Hieron. in Engler, Bot. Jahrb. 22: 683 (1897). trixioides Rusby in Mem. Torr. Bot. Club, 6: 54 (1896) — V. fulta. varroniaefolia DC. Prodr. 5: 56 (1836 . yungasensis Britton in Bull. Torr. Bot. Club, 18: 332 (1891). Viguiora anchusaefolia (DC.) Baker in Mart. Fl. Bras. 6(3): 222 (1884). V. australis Blake in Contrib. Gray Herb. 54: 148 (1918). V. fusiformis Blake in Contrib. Gray Herb. 54: 145 (1918). V. glutinosa Rusby in Mem. Torr. Bot. Club, 4(3): 211 (1895) — Flourensia heterolepi V. lanceolata Britton in Bull. Torr. Bot. Club, 19: 149 ay V. macrorhiza Baker in Mart. Fl. Bras. 6(3): 225 (188 V. Mandonii Sch. Bip. ex Rusby in Mem. Torr. Bot. eee ‘gay 60 (18938) = V. lanceolata. : V. pazensis Rusby in Mem. Torr. Bot. Club, 3(3): 59 (1893). V. Pflanzii Perkins in Engler, Bot. Jahrb. 49: 226 (1918). V. retroflexa Blake in Contrib. Gray Herb. 54: 146 (1918). Villanova oppositifolia Lag. Nov. Gen. & Spec. 31 (1815). Wedelia brachycarpa Baker in Mart. Fl. Bras. 6(3): 181 (1884). W. Holwayi Blake in Bot. Gaz. 74: 420 (1922). W. isolepis Blake in Bot. Gaz. ig 421 Adve: oe a ee boraginifolia O. Ktze. Rev. Gen. a 184 (1898) = W. strigosissima. caespitosa var. Haenkei Wedd. Chlor. And. 1: 83 (1856). seme se (Wedd.) Rusby in Bull. N. Y. Bot. Gard. 4: 398 (1907). ta Wedd. ex Sch. Bip. in Linnaea, 34: 530 (1866), in synon. = W. pecti- diliciete A. Gray in Proc. Am. Acad. 5: 140 (1861). dactylophylla Sch. Bip. in Bonplandia, 4: 53 (1858). dactylophylla forma glabriuscula Rockhausen in Engler, Bot. Jahrb. 70: 286 (1939 digitata Wedd. Chlor. And. 1: 86, t. 17D (1856). heteroloba Wedd. Chlor. And. 1: 88, t. 16A (1856). Knocheae Perkins in Engler, Bot. Jahrb. 49: 230 (1913) — W. pectinata. Mandoniana Wedd. ex Klatt in Ann. Naturh. Hofmus. Wien, 9: 367 (1894) = W. Orbignyana. melanandra Wedd. Chlor. And. 1: 88 (1856). nubigena HBK. Nov. Gen. & Spec. 4: 198 (1820). Orbignyana Wedd. Chlor. And. 1: 85 (1856). pectinata Lingelsh. in Fedde, Repert. Spec. Nov. 8: 6 (1910). plantaginifolia Wedd. ex Klatt in Ann. Naturh. Hofmus. Wien, 9: 367 (1894). bpbb S344 442 sia 223 pygmaea Gill. ex Hook. & Arn. in Journ. Bot. 3: 348 (1841). setosa Wedd. ex Sch. Bip. in Linnaea, 34: 530 (1865-66), in synon. = W. strigosissima. solivaefolia Sch. Bip. in Bonplandia, 4: 53 (1856). spathulata Wedd. Chlor. And. 1: 85, t. 17A (1856). staticaefolia Sch. Bip. in Bonplandia, 4: 53 (1856). strigosissima A. Gray in Proc. Am. Acad. 5: 140 (1861). villosa A. Gray in Proc. Am. Acad. 5: 139 (1861). Wulffia baccata (L. f.) O. Ktze. Rev. Gen. 3(2): 184 (1898). Xanthium orientale L. Sp. Pl. (ed. 2) 2: 1400 (1763). X. spinosum L. Sp. Pl. 987 (1753). Zexmenia foliosa Rusby ex W. W. Jones in Proc. Am, Acad. 41: 162 (1905). Z. Herzogii Hassl. in Fedde, Repert. Spec. Nov. 7: 357 (1909). Z. mikanioides (Britton) Blake in Journ. Bot. 53: 200 (1915). Zinnia paucifiora L. Sp. Pl. (ed. 2) 2: 1269 (1763). 42822 33