ILL { 



AT URBA ..AMPAI 

 BIOLOGY 



JUL 1 9 JQ9? 






5 



FIELDIANA 

 Botany 



Published by Field Museum of Natural History 



New Series, No. 5 



OCT22-/9C 



FLORA OF PERU 



J. FRANCIS MACBRIDE 

 AND COLLABORATORS 



CONSPECTUS AND INDEX TO FAMILIES 



ALWYN H. GENTRY 



FAMILY COMPOSITAE: PART I 



INTRODUCTION TO FAMILY 

 MICHAEL O. DILLON 



TRIBE VERNONIEAE 

 SAMUEL B. JONES 



December 31, 1980 

 Publication 1314 





FLORA OF PERU 



CONSPECTUS AND INDEX TO FAMILIES 



FAMILY COMPOSITAE: PART I 



INTRODUCTION TO FAMILY 



TRIBE VERNONIEAE 





FIELDIANA 

 Botany 



Published by Field Museum of Natural History 



New Series, No. 5 



FLORA OF PERU 



J. FRANCIS MACBRIDE 

 AND COLLABORATORS 



CONSPECTUS AND INDEX TO FAMILIES 



ALWYN H. GENTRY 



Missouri Botanical Garden 

 St. Louis, Missouri 



FAMILY COMPOSITAE: PART I 



INTRODUCTION TO FAMILY 

 MICHAEL O. DILLON 



Department of Botany 



Field Museum of Natural History 



TRIBE VERNONIEAE 

 SAMUEL B. JONES 



University of Georgia 

 Athens, Georgia 



December 31, 1980 



Publication 1314 Accepted for publication July 12, 1979. 




Library of Congress Catalog Card No.: 80-66384 



ISSN 0015-0746 

 PRINTED IN THE UNITED STATES OF AMERICA 




CONTENTS 



List of Illustrations v 



List of Tables v 



Conspectus by Alwyn H. Gentry 1 



Index to Published Families 6 



Compositae by Michael O. Dillon 12 



Morphology 14 



Key to Tribes of Peruvian Compositae 20 



Tribe Vernonieae by Samuel B. Jones 22 



Key to Genera of Vernonieae 23 



I. Vernonia 24 



Key to Species of Vernonia 25 



1 . Vernonia pycnantha 28 



2. Vernonia lambayequensis 28 



3. Vernonia jalcana 29 



4. Vernonia woytkowskii 29 



5. Vernonia peruviana 30 



6. Vernonia jelskii 30 



7. Vernonia libertadensis 31 



8. Vernonia gracilis 31 



9. Vernonia laurifolia 32 



10. Vernonia sordidopapposa 32 



1 1 . Vernonia mapirensis 33 



12. Vernonia ferruginea 33 



13. Vernonia costata 34 



14. Vernonia stuebelii 34 



15. Vernonia sambrayana 34 



16. Vernonia patens 35 



17. Vernonia fulta 36 



18. Vernonia apurimacensis 38 



19. Vernonia scorpioides 38 



20. Vernonia brachiata 41 



21 . Vernonia cainarachiensis 41 



22. Vernonia yurimaguasensis 43 



23. Vernonia myriocephala 43 



24. Vernonia canescens 44 



25. Vernonia fieldiana 45 



26. Vernonia salzmanii 46 



27. Vernonia herbacea . . 46 




IV 



II. Piptocarpha 47 



Key to Species of Piptocarpha 48 



1 . Piptocarpha poeppigiana 48 



2. Piptocarpha asterotrichia 49 



3. Piptocarpha opaca 51 



4. Piptocarpha canescens 52 



5. Piptocarpha sprucei 52 



6. Piptocarpha lechleri 53 



7. Piptocarpha gutierrezii 54 



III. Pollalesta 54 



1 . Pollalesta discolor 55 



IV. Centratherum 57 



1 . Centratherum punctatum 59 



V. Struchium 60 



1 . Struchium sparganophorum 60 



VI. Elephantopus 62 



Key to Species of Elephantopus 63 



1 . Elephantopus mollis 63 



2. Elephantopus angustifolius 65 



VII. Pseudelephantopus 65 



Key to Species of Pseudelephantopus 66 



1 . Pseudelephantopus spicatus 66 



2. Pseudelephantopus spiralis 68 



Acknowledgments 68 



Index . . 70 




LIST OF ILLUSTRATIONS 



1 . Vernonia patens 37 



2. Vernonia scorpioides 40 



3. Vernonia brachiata 42 



4. Piptocarpha asterotrichia 50 



5 . Pollalesta discolor 56 



6. Centratherum punctatum 58 



7. Struchium sparganophorum 61 



8. Elephantopus mollis 64 



9. Pseudelephantopus spiralis 67 



LIST OF TABLES 



1 . Largest families in Flora of Peru 4 



2. Comparison of species numbers in Flora of Peru and Peruvian species 



in Flora Neotropica monographs 4 



3. Estimates of the number of genera and species for the tribes represented 



in the Compositae of Peru 13 





THE FLORA OF PERU: A CONSPECTUS 



ALWYN H. GENTRY 



Missouri Botanical Garden 



2345 Tower Grove Ave. 



St. Louis, Mo. 63110 



The Flora of Peru is the only floristic treatment of Andean or upper 

 Amazonian plants and is one of the most significant of all floristic 

 works. Except for the monumental 19th century Flora Brasiliensis, the 

 Flora of Peru is today the closest thing to a complete Flora enjoyed by 

 any South American country. The present volume marks the re- 

 inauguration of this project after a hiatus in publication of almost a 

 decade. We anticipate that the Flora of Peru will be completed in 1986. 



The Flora of Peru was begun in 1936 under the direction of J. Francis 

 Macbride who had been hired by Field Museum in 1922 specifically to 

 study Peruvian plants. Macbride and associates made several plant- 

 collecting expeditions to Peru in the 1920's, and other collections of 

 Peruvian plants were also accumulated by Field Museum during this 

 time. By 1936 when the first volume of the Flora of Peru was pub- 

 lished, Dahlgren (1936) estimated that Field Museum collections in- 

 cluded over 33,000 sheets of Peruvian plants which were "undoubtedly 

 the most complete representation of the flora of that country in exis- 

 tence." Although this data base seems quite comparable to that on 

 which initial publications of other Latin American Floras now nearing 

 completion the largely concurrent Floras of Panama and 

 Guatemala were undertaken, it was hopelessly inadequate for 

 Amazonian Peru and far from complete for large areas of the Peruvian 

 uplands and many of the coastal lomas as well. As a result the Flora of 

 Peru shares with these other attempts to document the incredibly rich 

 neotropical flora many faults attributable to an inadequate collection 

 base (Gentry, 1978). 



Another problem with which Macbride was forced to cope, in an era 

 when transoceanic loans of specimens were not so readily available as 

 today, were many species described from Peru but inadequately known 




FIELDIANA: BOTANY 



to him. His solution was generally to accept essentially all species 

 which had been proposed: "It soon became evident that an attempt to 

 express an opinion on the merit or lack of merit of every species pro- 

 posed was impractical if the whole work was to be completed within a 

 reasonable period" (Macbride, 1936). Though perhaps inevitable in the 

 context of 1936, this lack of a thoroughgoing attempt to critically eval- 

 uate the species accepted in the Flora poses problems for its users 

 today. 



Despite the criticisms to which the perspective of half a century can 

 lead, Macbride's compilation of the Flora of Peru is universally rec- 

 ognized as having been a truly herculean task. Macbride' s own con- 

 tributions to the Flora spanned a quarter century from 1936 until 1962, 

 and several subsequent specialists' contributions swelled the number 

 of species published in the Flora to 11,789 (plus an additional 246 

 varieties) at the cessation of its active publication in 1971. 



Although contributing specialists were used when available, the 

 great majority of the compilation of Peruvian plants was by Macbride 

 himself. Paul Standley contributed 10 familial treatments, including the 

 large ones for Gramineae and Rubiaceae. Ellsworth Killip contributed 

 the treatments of Passifloraceae, Caprifoliaceae, Valerianaceae, Ur- 

 ticaceae, and the genus Bomarea. Charles Baehni contributed three 

 familial treatments: Lacistemaceae, Violaceae, and Sapotaceae, the 

 latter two co-authored with associates. Lyman Smith contributed 

 treatments of Bromeliaceae and Begoniaceae, the latter jointly au- 

 thored with B. Schubert, and J. Steyermark treated Fumariaceae and 

 Connaraceae. Fifteen other taxonomists contributed treatments of 

 families or important genera to the Flora, including Schweinfurth's 

 monumental Orchidaceae work and treatments of Piperaceae by Tre- 

 lease, Rum ex by Rechinger, Annonaceae by R. Fries, Myristicaceae by 

 A. C. Smith, Krameria by Hartmann, Monnina by R. Ferreyra, 

 Callitrichaceae by N. Fassett, Myrtaceae by R. McVaugh, Umbel- 

 liferae by M. Mathias and L. Constance, Hydrophyllaceae and 

 Polemoniaceae by D. Gibson, Solatium by D. Correll, Scrophu- 

 lariaceae by G. Edwin, Plantaginaceae by R. Pilger, and Cam- 

 panulaceae by F. Wimmer. Some of these are still considered among 

 the definitive taxonomic works on major plant groups. Altogether 32 

 families and an additional five genera were treated by specialists and 84 

 families were treated by Macbride. 



In 1975 the Flora of Peru was revitalized as a joint project of Field 

 Museum and the Missouri Botanical Garden under this author's direc- 

 tion and supported by the National Science Foundation. An additional 

 15 spermatophyte families remain to be treated, including the Com- 




MACBRIDE: FLORA OF PERU 3 



positae, the largest family of the Peruvian flora, which has been sub- 

 divided into tribes to be published individually. Specialists' treatments 

 of all the remaining families and the tribes of Compositae have been 

 arranged, with the last promised by 1986. 



Pteridophytes were not included in the original Flora of Peru but 

 Rolla Tryon (1964) has separately published an account of an estimated 

 quarter (187 species) of the Peruvian ferns using a somewhat more 

 elaborate format than Macbride's. The remainder of the Peruvian 

 pteridophytes will also be treated under the reactivated Flora of Peru, 

 although arrangements for specialist contributions of only part of the 

 pteridophytes have been completed to date. 



At this point a preliminary analysis of the Peruvian flora and the 

 completeness of its coverage by the published Flora seems appropri- 

 ate. To the 11,789 species of spermatophytes treated to date can be 

 added 2,148 additional species, the sum of the estimates of numbers of 

 species in their groups by the various contributors, to give a total 

 number of 13,937 species expected to have been treated in the Flora of 

 Peru when it is completed. Similarly 1,654 genera of spermatophytes 

 have already been treated and 298 remain to be covered, for a total of 

 1,952 genera to be included in the Flora. Table 1 lists the largest 

 families and genera in Peru, as treated in the Flora. It is noteworthy 

 that the number of species included is nearly double that included in 

 the Flora of Guatemala and approaching triple the number of species 

 included in the Flora of Panama (see Gentry, 1978), a striking indica- 

 tion of the immensity of the task undertaken by Macbride. Only the 

 19th century Flora Bras Hie nsis covers a larger portion of the neotropi- 

 cal flora. 



It is obvious that a neotropical Flora whose publication was begun in 

 1936 is likely to omit many species which actually occur in the country. 

 For example the Flora of Panama will treat only 5,000 of the 8,000- 

 9,000 species estimated to actually occur in that country (Gentry, 

 1978). In Peru coverage of the floristically rich Amazonian region is 

 especially incomplete, suggesting that many more than the 14,000 

 species treated in the Flora may actually occur in Peru. 



On the other hand, many of the published treatments in the Flora of 

 Peru were prone to excessive taxonomic splitting. For example, the 

 genus Peperomia was treated as including 342 species and varieties in 

 Peru; the fact that 78% of the accepted taxa were based on single 

 collections and 90% on two or fewer collections is highly suggestive of 

 unwarranted splitting. An average of 1.59 collections per accepted 

 taxon could hardly be adequate for understanding intraspecific varia- 




TABLE 1. Largest families in Flora of Peru. 



Family No. of species 



Compositae 1 ,432* 



Orchidaceae 1,290 (+38 var.) 



Leguminosae 751 (+7 var.) 



Piperaceae 726 (+64 var.) 



Melastomataceae 509 



Rubiaceae 480 



Gramineae 408 



Solanaceae 401 ( + 29 var.) 



Euphorbiaceae 269 



Scrophulariaceae 229 (+12 var.) 



Malvaceae 218 



Campanulaceae 192 (+42 var.) 



Myrtaceae 178 (+6 var.) 



Bromeliaceae 175 



Verbenaceae 174 ( + 2 var.) 



Labiatae 173 



Araceae 165 



Cyperaceae 156 (+3 var.) 



Gesneriaceae 155* 



Gentianaceae 1 50 



Guttiferae 150* 



Cactaceae 150* 



*Estimated. 



TABLE 2. Comparison of species numbers in Flora of Peru and Peruvian species in 

 Flora Neotropica monographs. 



No. of genera No. of species (+var.) 



Taxon Fl. of Peru FI. Neotr. Fl. of Peru Fl. Neotr. 



Swartzia 1 1 11 13(+1) 



Brunelliaceae 1 1 88 

 Moraceae (Olmedieae and 



Brosimeae) 9 9 29 30(+2) 



Zingiberaceae 4 4 37 29(+3) 



Chrysobalanaceae 4 4 29 36(+l) 



Dichapetalaceae 4 3 15 14(+1) 



Caryocaraceae 22 86 



Manihot 1 1 65 

 Bromeliaceae (Pitcairnioideae 



and Tillandsioideae) 8 8 148 270(+13) 



Memecyleae 1 1 911 



Trigoniaceae 1 1 76 



Bignoniaceae 44 41 106 125 



Totals 80 76 413 553 




MACBRIDE: FLORA OF PERU 5 



tion in Peperomia. Uncritical treatments of other groups similarly led 

 to inclusion of some variable species under several different names, 

 inflating the number of Peruvian species. 



Is it possible to reconcile these two opposing trends and arrive at a 

 meaningful estimate of the actual number of species in the Peruvian 

 flora without critically reworking the entire Flora? One feasible ap- 

 proach is to compare the number of species "lost" and "gained" from 

 the Flora when compared with that in recent monographs of plant 

 groups occurring in Peru. I have used the Flora Neotropica monograph 

 series to arrive at such an estimate. My own specialty group Big- 

 noniaceae is also included, since I have the data readily available, even 

 though only part of the family has been monographed to date for Flora 

 Neotropica. Table 2 compares the number of species and genera re- 

 corded from Peru in each of the relevant Flora Neotropica treatments 

 with the number treated in the Flora. The total of 413 species treated in 

 the Flora of Peru for these 12 groups increased to 553 species in the 

 Flora Neotropica monographs, a 34% average increase. Although 

 acknowledging that 413 species is a perilously small (3%) sample of the 

 total Flora of Peru species, we may tentatively extrapolate from these 

 figures an average increase of 34% in number of species now known 

 from Peru as compared with the number included in the Flora. Applied 

 to the 13,937 treated species, this would project to 18,676 Peruvian 

 plant species. 



Actually the Peruvian flora might be expected to be significantly 

 richer than this since very few of the 15,000 collections generated by 

 the current Flora of Peru project have been included in the Flora 

 Neotropica treatments on which these estimates were based. Many 

 new and new-to-Peru species have already been discovered in some of 

 these groups subsequent to the Flora Neotropica monographs. 

 Moreover the rate of discovery of new species in these groups shows 

 no signs of leveling off. Thus it seems likely that a definitive tabulation 

 of Peruvian plants would eventually be expected to include well over 

 20,000 species, approximately as many plant species as are included in 

 such very much larger areas as North America or tropical continental 

 Africa (cf. Raven, 1976). Thanks to its Flora, Peru is the only tropical 

 South American country (except tiny Surinam) for which such a figure, 

 useful however tentative, can somewhat meaningfully be extrapolated. 



REFERENCES 



DAHLGREN, B. E. 1936. Preface to Flora of Peru. Field Mus. Nat. Hist., Bot. Ser., 13, 




6 FIELDIANA: BOTANY 



GENTRY, A. H. 1978. Floristic knowledge and needs in Pacific Tropical America. Brit- 



tonia30, pp. 134-153. 

 MACBRIDE, J. F. 1936. Introduction to Flora of Peru. Field Mus. Nat. Hist., Bot. Ser., 



13, pp. 9-13. 

 RAVEN, P. 1976. Ethics and attitudes. In Simmons, J. et al., eds., Conservation of 



Threatened Plants. Plenum, New York and London, pp. 155-179. 

 TRYON, R. 1964. The ferns of Peru: Polypodiaceae (Dennstaedtieae to Oleandreae). 



Contr. Gray Herb. 194, pp. 1-253. 



INDEX TO PUBLISHED FAMILIES 



Family Publication Data 



Acanthaceae (Wasshausen) 



Actinidiaceae 3A(2): 677-686. 1956. 



Aizoaceae 2(2): 558-562. 1937. 



Alismataceae 1(1): 91-94. 1936. 



Amaranthaceae (Standley) 2(2): 478-518. 1937. 



Supplement 2(3): 1134-1136. 1938. 

 Amaryllidaceae (Bomarea 



by Killip) 1(3): 631-690. 1936. 



Anacardiaceae 3A(1): 238-258. 1951. 



Annonaceae (Fries) 2(3): 700-766. 1938. 



Apocynaceae 5(1): 363-455. 1959. 



Aquifoliaceae 3A(1): 270-288. 1951. 



Araceae 1(3): 428-486. 1936. 



Araliaceae 5(1): 8-44. 1959. 



Aristolochiaceae 2(2): 431-443. 1937. 

 Asclepiadaceae (Spellman & 



Morillo) 



Balanophoraceae 2(2): 427-431. 1937. 

 Balsaminaceae (Gentry) 



Basellaceae 2(2): 573-578. 1937. 



Bataceae 2(2): 546. 1937. 

 Begoniaceae (L. Smith & 



B. Schubert) 4(1): 181-202. 1941. 



Berberidaceae 2(3): 665-680. 1938. 




MACBRIDE: FLORA OF PERU 



Family 



Betulaceae 



Bignoniaceae 



Bixaceae 



Bombacaceae 



Boraginaceae 



Bromeliaceae (L. Smith) 



Brunelliaceae 



Burmanniaceae 



Burseraceae 



Butomaceae 



Buxaceae 



Cactacea (Solomon) 



Callitrichaceae (Fassett) 



Calyceraceae 



Campanulaceae (Wimmer) 



Cannaceae 



Capparaceae 



Caprifoliaceae (Killip) 



Caricaceae 



Caryocaraceae 



Caryophyllaceae 



Celastraceae 



Ceratophyllaceae (Gentry) 



Chenopodiaceae (Standley) 



Chloranthaceae 



Clethraceae 



Cochlospermaceae 



Columelliaceae 



Combretaceae 



Commelinaceae 



Compositae (Dillon, Jones, 



King, Holmes, McDaniel, 



Turner, Robinson, 



Sagastegui, Keil, Barkley, 



Ferreyra) 



Connaraceae (Steyermark) 

 Convolvulaceae 

 Coriariaceae 

 Cornaceae 

 Crassulaceae 



Publication Data 



2(2): 267-268. 1937. 

 5C(1): 3-101. 1961. 

 (as Flacourtiaceae) 

 3A(2): 593-622. 1956. 

 5(2): 539-609. 1960. 

 1(3): 495-592. 1936. 

 (as Cunoniaceae) 

 1(3): 767-768. 1936. 

 3(2): 703-717. 1949. 

 1(1): 94-95. 1936. 

 3A(1): 220-221. 1951. 



3A(1): 235-237. 1951. 

 6(2): 489-491. 1937. 

 6(2): 383-489. 1937. 

 1(3): 738-741. 1936. 

 2(3): 984-1006. 1938. 

 6(2): 281-287. 1937. 

 4(1): 132-143. 1941. 

 3A(2): 697-703. 1956. 

 2(2): 578-638. 1937. 

 3A(1): 259-270. 1951. 



2(2): 469-478. 1937. 

 2(2): 257-260. 1937. 

 5(1): 45-50, 1959. 

 (as Flacourtiaceae) 

 5C(1): 101-103. 1961 

 4(1): 221-229. 1941. 

 1(3): 592-608. 1936. 



2(3): 1119-1125. 1938. 

 5(1): 455-536. 1959. 

 3A(1): 237-238. 1951. 

 5(1): 44-45. 1959. 

 2(3): 1007-1015. 1938. 




8 



FIELDIANA: BOTANY 



Family 



Cruciferae 



Cucurbitaceae 



Cunoniaceae 



Cycadaceae 



Cyclanthaceae (Standley) 



Cyperaceae 



Dichapetalaceae 



Dilleniaceae 



Dioscoreaceae 



Dipsacaceae 



Ebenaceae 



Elaeocarpaceae 



Elatinaceae 



Ephedraceae 



Ericaceae 



Eriocaulaceae 



Erythroxylaceae 



Euphorbiaceae 



Flacourtiaceae 



Frankeniaceae 



Fumariaceae (Steyermark) 



Gentianaceae 



Geraniaceae 



Gesneriaceae (Skog) 



Gnetaceae 



Gramineae (Standley) 



Guttiferae (Maguire) 



Haemodoraceae 



Haloragaceae 



Hernandiaceae 



Hippocrateaceae 



Humiriaceae 



Hydrocharitaceae 



Hydrophyllaceae (Gibson) 



Hypericaceae (Robson) 



Icacinaceae 



Iridaceae 



Juglandaceae 



Julianaceae 



Juncaceae 



Labiatae 



Publication Data 



2(3): 937-983. 1938. 

 6(2): 321-383. 1937. 

 2(3): 1038-1063. 1938. 

 1(1): 81-82. 1936. 

 1(3): 421-428. 1936. 

 1(1): 261-320. 1936. 

 3(3): 954-964. 1950. 

 3A(2): 667-677. 1956. 

 1(3): 690-707. 1936. 



5(1): 205-214. 1959. 

 (as Tiliaceae) 

 1(1): 84-86. 1936. 

 1(1): 84-86. 1936. 

 5(1): 50-149. 1959. 

 1(3): 489-494. 1936. 

 3(2): 632-647. 1949. 

 3A(1): 3-200. 1951. 

 4(1): 5-52. 1941. 

 4(1): 4-5. 1941. 

 2(3): 936-937. 1938. 

 5(1): 270-363. 1959. 

 3(2): 511-544. 1949. 



1(1): 86. 1936. 

 1(1): 96-261. 1936. 



1(3): 630-631. 1936. 

 5(1): 3-8. 1959. 

 2(3): 931-933. 1938. 

 3A(1): 200-220. 1951. 

 (as Linaceae) 

 1(1): 95-%. 1936. 

 5A(2): 101-112. 1967. 



3A(1): 221-233. 1951 

 1(3): 707-717. 1936. 

 2(2): 263-266. 1937. 

 2(2): 266-267. 1937. 

 1(3): 609-617. 1936. 

 5(2): 721-829. 1960. 




MACBRIDE: FLORA OF PERU 



Family 



Lacistemataceae (Baehni) 

 Lauraceae 

 Lecythidaceae 

 Leguminosae (Krameria 



by Hartmann) 

 Lemnaceae 



Lentibulariaceae (Taylor) 

 Liliaceae 

 Linaceae 

 Loasaceae 

 Loganiaceae 

 Loranthaceae 

 Lythraceae 



Magnoliaceae (Lozano) 

 Malesherbiaceae 

 Malpighiaceae 

 Malvaceae 



Addendum 

 Marantaceae 

 Marcgraviaceae 

 Martyniaceae (Gentry) 

 Mayacaceae 

 Melastomataceae 

 Meliaceae 

 Menispermaceae 

 Monimiaceae 

 Moraceae 



Supplement 

 Musaceae 

 Myricaceae 



Myristicaceae (A. C. Smith) 

 Myrsinaceae 

 Myrtaceae (McVaugh) 

 (Najadaceae) 

 Nolanaceae 



Nyctaginaceae (Standley) 

 Nymphaeaceae (Standley) 

 Ochnaceae 

 Olacaceae (Standley) 



Supplement 

 Oleaceae 



Publication Data 



4(1): 52-56. 1941. 

 2(3): 819-931. 1938. 

 4(1): 229-249. 1941. 



3(1): 3-507. 1943. 

 1(3): 486-487. 1936. 



1(3): 617-630. 1936. 

 3(2): 621-632. 1949. 

 4(1): 143-181. 1941. 

 5(1): 239-269. 1959. 

 2(2): 375-416. 1937. 

 4(1): 206-219. 1941. 



4(1): 85-90. 1941. 

 3(3): 781-871. 1950. 

 3A(2): 442-593. 1956. 



3A(2): 742-744. 1956. 

 1(3): 741-767. 1936. 

 3A(2): 703-717. 1956. 



1(3): 487. 1936. 

 4(1): 249-521. 1941. 

 3(2): 717-777. 1949. 

 2(3): 680-699. 1938. 

 2(3): 784-819. 1938. 

 2(2): 274-331. 1937. 



2(3): 1126-1127. 1938. 

 1(3): 717-726. 1936. 

 2(2): 261-263. 1937. 

 2(3): 766-784. 1938. 

 5(1): 163-203. 1959. 

 4(2): 567-818. 1958. 

 1(1): 89. 1936. 

 5(2): 829-854. 1960. 

 2(2): 518-546. 1937. 

 2(2): 638-639. 1937. 

 3A(2): 686-697. 1956. 

 2(2): 421-427. 1937. 



2(3): 1127-1132. 1938. 

 5(1): 235-239. 1959. 




10 



FIELDIANA: BOTANY 



Family 



Onagraceae 

 Opiliaceae (Standley) 

 Orchidaceae (Schweinfurth) 



Supplement 

 Orobanchaceae 

 Oxalidaceae 

 Palmae 

 Papaveraceae 

 Passifloraceae (Killip) 

 Pedaliaceae (Gentry) 

 Phytolaccaceae 

 Piperaceae (Trelease) 

 Plantaginaceae (Pilger) 

 Plumbaginaceae 

 Podocarpaceae 

 Podostemaceae 

 Polemoniaceae (Gibson) 

 Polygalaceae (Monnina by 



Ferreyra) 

 Polygonaceae (Standley) 



(Rumex by Rechinger) 

 Pontederiaceae 

 Portulacaceae 

 Potamogetonaceae 

 Primulaceae 

 Proteaceae 

 Quiinaceae 

 Raffle siaceae 

 Ranunculaceae 

 Rapateaceae 

 Rhamnaceae 

 Rhizophoraceae 

 Rosaceae 



Rubiaceae (Standley) 

 Rutaceae 



Sabiaceae (Gentry) 

 Salicaceae 

 Santalaceae 



Publication Data 



4(1): 521-566. 1941. 

 2(2): 420-421. 1937. 

 30(1): 1-260. 1958. 

 30(2): 261-531. 1959. 

 30(3): 533-786. 1960. 

 30(4): 787-1005. 1961. 



33: 1-80. 1970. 

 5C(1): 103-104. 1961. 

 3(2): 544-608. 1949. 

 1(2): 321-418. 1960. 

 2(3): 933-936. 1938. 

 4(1): 90-132. 1941. 



2(2): 546-558. 1937. 

 2(1): 3-253. 1936. 

 6(2): 265-281. 1937. 

 5(1): 203-205. 1959. 

 (as Taxaceae) 

 2(3): 1007. 1938. 

 5A(2): 112-131. 1967. 



3(3): 891-950. 1950. 



2(2): 444-468. 1937. 

 1(3): 608-609. 1936. 

 2(2): 562-573. 1937. 

 1(1): 87-89. 1936. 

 5(1): 149-152. 1959. 

 2(2): 367-375. 1937. 

 3A(2): 717-726. 1956. 

 2(2): 443-444. 1937. 

 2(2): 639-661. 1937. 

 1(3): 494-495. 1936. 

 3A(2): 391-408. 1956. 

 4(1): 219-221. 1941. 

 2(3): 1063-1119. 1938. 

 6(1): 3-261. 1936. 

 3(2): 655-689. 1949. 



2(2): 260-261. 1937. 

 2(2): 416-420. 1937. 




MACBRIDE: FLORA OF PERU 



11 



Family 



Sapindaceae 

 Sapotaceae (Baehni & 



Bernard!) 

 Saxifragaceae 

 Scheuchzeriaceae 

 Scrophulariaceae (Edwin) 

 Simaroubaceae 

 Solanaceae (excluding 



Solarium) 



Solanum (Correll) 

 Staphyleaceae 

 Sterculiaceae 

 Styracaceae 

 Symplocaceae 

 (Taccaceae) 

 Taxaceae 

 Theaceae 

 Theophrastaceae 

 (Thurniaceae) 

 Thymelaeaceae 

 Tiliaceae 

 Tovariaceae 

 Trigoniaceae 

 Triuridaceae 

 Tropaeolaceae 

 Turneraceae 

 Typhaceae 

 Ulmaceae 

 Umbelliferae (Mathias & 



Constance) 

 Urticaceae (Killip) 

 Valerianaceae (Killip) 

 Velloziaceae (Gentry) 

 Verbenaceae 



Violaceae (Baehni & Weibel) 

 Vitaceae 

 Vochysiaceae 

 Winteraceae 

 Xyridaceae 

 Zingiberaceae 

 Zygophyllaceae 



Publication Data 



3A(2): 291-391. 1956. 



5A(3): 135-177. 1970. 

 2(3): 1015-1038. 1938. 

 1(1): 90-91. 1936. 

 5B(3): 459-717. 1971. 

 3(2): 689-703. 1949. 



5B(1): 3-267. 1962. 



5B(2): 271-458. 1967. 

 3A(1): 233-235. 1951. 

 3A(2): 622-667. 1956. 

 5(1): 225-235. 1959. 

 5(1): 214-225. 1959. 

 1(3): 690. 1936. 

 1(1): 82-84. 1936. 

 3A(2): 726-741. 1956. 

 5(1): 153-163. 1959. 

 1(3): 494. 1936. 

 4(1): 203-206. 1941. 

 3A(2): 413-442. 1956. 

 2(3): 1006-1007. 1938. 

 3(3): 950-954. 1950. 

 1(1): 96. 1936. 

 3(2): 608-620. 1949. 

 4(1): 82-85. 1941. 

 1(1): 87. 1936. 

 2(2): 268-274. 1937. 



5A(1): 1-97. 1962. 

 2(2): 331-367. 1937. 

 6(2): 287-321. 1937. 



5(2): 609-721. 1960. 

 4(1): 56-82. 1941. 

 3A(2): 408-413. 1956. 

 3(3): 872-891. 1950. 

 2(3): 699-700. 1938. 

 1(3): 487-489. 1936. 

 1(3): 726-738. 1936. 

 3(2): 647-654. 1949. 




12 FIELDIANA: BOTANY 



COMPOSITAE Giseke 1 2 



Annual, biennial or perennial herbs or sometimes shrubs, trees, or vines, variously 

 pubescent or glandular, sometimes glabrous, lactiferous or not; stems terete, sometimes 

 winged or flattened into cladodes. Leaves alternate, verticillate, or opposite, sometimes 

 basal, rarely reduced to scales, spines, or wanting, simple or 2- to many-foliolate, entire, 

 or variously toothed, lobed or dissected; petioles present or wanting; the leaf bases 

 sometimes decurrent or clasping; exstipulate, but pseudostipules sometimes present. 

 Inflorescence cymose, racemose, paniculate, umbellate, or of solitary capitula, some- 

 times in indefinite aggregates; usually pedunculate, rarely with the capitula in glomerules 

 (pseudocephalium); often bracteate; usually pedicellate, sometimes bracteolate. Capitula 

 with 1-many florets inserted on a receptacle; heterogamous, radiate or disciform, or 

 homogamous, discoid or ligulate; basally enclosed in an involucre; phyllaries (involucral 

 bracts) few to many in 1 to several similar, differentiated, or evenly graded series, free or 

 connate, valvate or imbricate; receptacle convex, concave, flat, or conical; paleae flat or 

 keeled and enfolding the florets, or reduced to hairs or short scales, or wanting; florets 

 epigynous, either all hermaphrodite and protandrous, or female, male, or neuter (sterile); 

 corollas gamopetalous, tubular, filiform, ligulate or bilabiate, usually 3- to 5-toothed, 

 rarely absent, the stamens 5, rarely 3 or 4, epipetalous, filaments usually free, the anthers 

 mostly oblong, marginally connate, introrse with sterile appendages, basally truncate to 

 tailed, the style branches 2, pubescent, glabrate or glandular, the ovary terete or com- 

 pound, often with apical nectary. Fruit usually an achene (cypsela), rarely baccate or 

 drupaceous, or a utricle formed by fusion of the achene with paleae, bracts or other 

 parts, the pericarp mostly hard; pappus usually present, of bristles, awns, or scales; 

 sometimes with a distinct carpopodium. 



The Compositae is one of the largest flowering plant families in the 

 world, represented by over 1,400 genera and estimates of between 

 20,000 and 30,000 species. Only the Orchidaceae is comparable in 

 number with about 750 genera and some 18,000 species. The Com- 

 positae is cosmopolitan in distribution, occurring on all continents, 

 except Antarctica. The family is well developed in the New World, 

 with Peru being a center of diversity for several tribes. In Peru, there 

 are over 1 ,400 species of Compositae representing approximately 10% 

 of the total Peruvian flora (see Gentry, A Conspectus). 



Presently, 13 tribes are recognized as occurring in Peru. The 

 generic composition of the tribes reflects the results of the Reading 

 Symposium on the Biology and Chemistry of the Compositae (1977). 



'The treatment for the family Compositae is being coordinated by Michael O. Dillon, 

 Field Museum of Natural History, who wrote introductory material including the family 

 description and key to tribes. Tribes are being published separately as they are com- 

 pleted, with authorship indicated at the beginning of the taxon. Each author is solely 

 responsible for his treatment. 



2 Assisted by National Science Foundation Grant DEB-79-05078 (Alwyn H. Gentry, 

 principal investigator). 




MACBRIDE: FLORA OF PERU 13 



TABLE 3. Estimates of the number of genera and species for the tribes represented 

 in the Compositae of Peru. 



Tribe No. of genera No. of species 



VERNONIEAE 7 39 



LIABEAE 12 50 



EUPATORIEAE 39 290 



ASTEREAE 15 200 



INULEAE 10 67 



HELIANTHEAE 70 289 



TAGETEAE 5 23 



ANTHEMIDEAE 6 11 



SENECIONEAE 9 231 



CALENDULEAE 1 1 



CARDUEAE 2 2 



MUTISIEAE 21 200 



LACTUCEAE 6 29 



TOTAL 203 1 ,432 



Estimates of the number of genera and species within each tribe are 

 given in Table 3. The tribes Eupatorieae and Heliantheae are the 

 largest, with each containing some 20% of the total, followed by the 

 Senecionieae (ca. 16%), the Astereae (ca. 14%), and the Mutisieae (ca. 

 14%). The tribe Liabeae is here recognized and considered most 

 closely aligned with the tribe Vernonieae. The polyphyletic tribe 

 Helenieae (sensu Bentham) is not maintained, with constituent genera 

 being realigned with the tribes Heliantheae, Senecioneae, and 

 Tageteae. The tribe Calenduleae is represented by the introduced or- 

 namental Calendula officinalis L. Only the African tribe Arctoteae is 

 unrepresented in the flora. 



In Peru, the family has radiated into a wide variety of habitats, 

 including the puna, inter-montane valleys, the lomas of the coastal 

 desert, and the ceja de la montana; however, few are present in the 

 tropical and subtropical rain forests. Nearly every type of habit is to be 

 found, with perennial herbs and shrubs predominating. 



Despite their abundance in the flora, few members of the family have 

 any economic importance in Peru. Several introduced ornamentals are 

 cultivated and sold in the markets (e.g., Calendula, Chrysanthemum), 

 and some native species are used in folk medicine (e.g., Spilanthes, 

 Tagetes). At least some members of the genus Clibadium and possibly 

 Ichthyothere (Heliantheae) are used as fish poison in the lower Amazon 

 basin. 




14 FIELDIANA: BOTANY 



MORPHOLOGY 3 



Plants of the Compositae display a range of specialized morphology 

 not found in other families, and terminology is often particular to the 

 family. A hand lens or dissecting microscope is useful in examining 

 these plants and some features must be studied with a compound mi- 

 croscope. Literature citations in the following survey of terminology 

 refer mainly to good illustrations of Compositae structures. 



Pubescence and glands. Characteristic hair (trichome) types are 

 found in several groups of Compositae (cf. D'Arcy, 1975; fig. 1). In the 

 Vernonieae hairs are sometimes sturdy, elongate, and single-celled. In 

 the Eupatorieae and Astereae hairs are usually many-celled and uni- 

 seriate or moniliform, with the basal or apical cell sometimes slightly 

 differentiated. Arachnoid hairs, too fine to be seen in cellular detail 

 under magnifications less than x45, occur and may form tomentum in 

 the Inuleae, Liabeae, Senecioneae, and Cardueae. A specialized "ver- 

 rucose hair" occurs in many genera of the Heliantheae. This hair con- 

 sists of a multicellular basal rosette, one or two sturdy, distinctly ver- 

 rucose, erect cells, and an apex of one or two smooth, acicular cells. 

 The basal rosette of cells is sometimes calcified giving the leaf a 

 punctate appearance, and the sometimes calcified rugose and apical 

 cells may result in a scabrous leaf surface. Large multiseriate hairs 

 occurring in Trixis (Mutisieae), Hieracium (Lactuceae), and Pectis 

 (Tageteae), and others may be termed bristles. Branched hairs occur 

 on Hieracium and some species of Senecio. For a discussion of the 

 double hairs (Zwillingshaare) found on the ovaries of many genera and 

 especially of some primitive elements, see Hess (1938). 



Paleae (chaff) and receptacle (torus). Convention refers to bracts 

 external to the outermost whorl of florets as involucral bracts and those 

 internal to it as paleae. Although artificial, this distinction causes little 

 difficulty. The two structures are homologous with leaves but the 

 paleae are usually considerably more modified. Paleae are best devel- 

 oped in the Heliantheae and Mutisieae but isolated species or genera of 

 the Eupatorieae, Astereae, Liabeae, and Lactuceae and perhaps other 

 tribes also have paleae. In the Heliantheae the paleae frequently enfold 

 the ovary and may be bent over the corolla in bud or occasionally are 

 apically modified into awns or cusps. The paleae of Eclipta, Cirsium 

 and some Liabeae are narrowed into bristles or awns. In many genera 

 paleae are reduced to hairs or low scales which may persist on the 

 receptacle. In some genera, low hairs or spicules on the receptacle are 

 referred to as paleae although they may consist of enations of the 



3 Adapted largely from D'Arcy, 1975; pp. 837-843. 




MACBRIDE: FLORA OF PERU 15 



receptacle, or remains of carpopodia and are not homologous with the 

 bracts noted above. Aged receptacles may be fimbrillate (fringed), 

 pilose, foveate (pitted), verrucose (warty or knobby), alveolate (hon- 

 eycombed), spiculiferous, muricate (spiny), or naked (lacking paleae). 

 The receptacle tissue may be completely sclerified or include paren- 

 chyma. 



Corollas. Corollas (Hoffman 1894: 99, 101; Solbrig 1963: 451; 

 Bentham 1873: tab. 8; D'Arcy 1975: figs. 34E, 104, 106, 34B, 48B, 81A, 

 93B, 98B, 57C, 58C) are considered to be either ligulate (rays) or tubu- 

 lar (disc), although the tubular form includes modifications to cam- 

 panulate, funnelform, etc., and ligulate corollas usually consist of a 

 tube and a straplike ligule. When extremely narrow, corollas are 

 termed filiform or capillary. The outline made by the top of the corollas 

 and paleae is referred to as the disc. In the Lactuceae all corollas have 

 a 5-lobed ligule. In other groups, ligulate corollas are confined to the 

 outer whorls of florets on the head or are lacking. In the Mutisieae, 

 ligulate corollas have a 3- or 4-lobed ligule and short, opposing lobes at 

 the top of the tube (bilabiate). In the Astereae, Inuleae, Heliantheae, 

 Tageteae, Senecioneae, and Anthemideae, ligules are 2- to 3-lobed or 

 entire, and an opposing lobe is seldom present. In Zinnia and Heliopsis 

 (Heliantheae) the corolla consists of a ligule persistent on the achene 

 and a tube is lacking, and in Melampodium also the tube may be obso- 

 lete. Ligulate corollas are lacking in all Peruvian taxa of Vernonieae, 

 Eupatorieae, and Cardueae and only tubular corollas are present. 

 Tubular corollas consist of a basal tube, an expanded limb, and 4-5 

 apical lobes. They are mostly actinomorphic but sometimes one suture 

 of the limb is deeper than the others (e.g., Elephantopus and Pseudel- 

 ephantopus), and in other cases two sutures are deeper, producing 

 slightly bilabiate corollas. In Cotula mexicana (Anthemideae) the disc 

 corollas are regularly 3-lobed, a rarity in the family. 



Sexual condition. Sexual condition of the florets is of great system- 

 atic utility. In the Vernonieae, Eupatorieae, and Cardueae (Peru) and 

 in a few genera in other groups, all florets are alike, perfect, and have 

 tubular corollas. Such heads are termed discoid. All florets of the Lac- 

 tuceae are also perfect and have only ligulate corollas. These heads are 

 termed ligulate. In the above mentioned groups all florets are fertile, 

 producing mostly viable achenes. In most other groups, the outer 

 florets are pistillate, lack stamens, and only rarely produce staminodes. 

 The outer florets may have tubular or ligulate corollas and the heads 

 are termed radiate or disciform depending on whether the ligules are 

 elongate (exceeding the stigmas and pappus) or short and inconspicu- 




16 FIELDIANA: BOTANY 



ous. The ovaries may be fertile or sterile. Variations in the above 

 conditions occur in a few groups. Some Mutisieae have two peripheral 

 whorls of pistillate florets, the outer with ligulate corollas and the inner 

 with tubular corollas. Whorls internal to these have perfect florets with 

 tubular corollas. In a few cultivated plants (e.g., some strains of Den- 

 dranthema and Tagetes) proliferation of pistillate, often abortive, 

 florets with ligulate corollas may supplant normal florets with tubular 

 corollas. 



Conspicuous, often pellucid, oil glands of various shapes are ar- 

 ranged characteristically on leaves and involucres in the Tageteae. In 

 Siegesbeckia (Heliantheae), Hieracium, and Sonchus (both Lac- 

 tuceae), large globose glands are displayed on bristles. In Baccharis 

 (Astereae), and Flourensia (Heliantheae), a coating of glandular mate- 

 rial may make the leaf shiny. With the aid of a lens, punctate glands in 

 the leaf surface or globose glandular materials on the surface may be 

 observed in many species. In the Lactuceae a network of laticifers 

 invisible without special techniques yields copious milky sap. 



Leaf arrangement. In Peru leaves are opposite or rarely verticillate 

 in most Eupatorieae, Tageteae, many Heliantheae, and Liabeae, but 

 are alternate in all other groups. Plants with leaves in basal rosettes 

 belong to groups with usually alternate leaves, but can occur in oppo- 

 site leaved members (e.g., Paranephelius, Liabeae). In plants with 

 opposite leaves, it is not unusual for some leaves and branches in the 

 region of the inflorescence to be alternate. 



Inflorescence (Capitulescence). Capitula (heads) are often grouped 

 into recognizable general inflorescences (capitulescences), i.e., cymes, 

 corymbs, racemes, panicles. A capitulum occurring singly is described 

 as solitary. When capitula are aggregated into a secondary capitulum, 

 it is termed a glomerule (pseudocephalium) or synflorescence (e.g., 

 Elephantopus). 



Involucral bracts (phyllaries). These are mostly numerous and in 

 most groups are overlapping in several graded series. Except in the 

 Eupatorieae this is referred to as imbricate, but in the Eupatorieae the 

 terms eximbricate, subimbricate, and imbricate are used to refer to 

 degrees of overlapping. In some species of Tageteae, Senecioneae, 

 Mutisieae, and Lactuceae, the bracts do not overlap but are valvate, 

 touching only at the margins, or they may sometimes be marginally 

 connate for part of their length. A whorl of short bracts at the base of 

 the involucre may be referred to as either subinvolucral bracts or as 

 calyculate bracts. Commonly one or more subinvolucral bracts may be 

 found on the pedicel, sometimes in a different phyllotaxy from the rest 




MACBRIDE: FLORA OF PERU 17 



of the plant. In Elephantopus and Pseudelephantopus (both Ver- 

 nonieae), the involucral bracts are decussate, and in these genera with 

 their heads fused into a common receptacle, a series of subinvolucral 

 bracts forms a pseudoreceptacle around the glomerule. 



Stamens. Stamens (Fig. 1, Hoffmann, 1894: 104; Bentham, 1873: 

 tab. 9; Cabrera, 1974: fig. 52, 53; D'Arcy 1975: fig. 1) are usually of the 

 same number as the corolla lobes. Filaments are usually compressed 

 and the anthers are connate or coherent into a narrow tube. The anther 

 apex is usually sterile and differentiated into a distinct, hyaline appen- 

 dage. In Ophryosporus andAdenostemma (Eupatorieae) and inEclipta 

 and Eleutheranthera (both Heliantheae), the appendage is much re- 

 duced or wanting. In the Mutisieae the anther apex is sterile but not de- 

 marcated on the dorsal (outer) side, appearing as a homogeneous con- 

 tinuation of the thecae. Anther bases may be blunt, auriculate, sagittate, 

 or with variously elaborated tails. The auricles of adjacent anthers are 

 sometimes united. In some cases short auricles appear to be derived 

 from longer but crumpled tails. Tails are present in most taxa of 

 Inuleae and Mutisieae. 



A ring or region of specialized cells near the top to the filaments, the 

 anther collar, acts as a hinge to permit straightening of the filaments at 

 anthesis when the style pushes through the anther tube with much of 

 the pollen. Characteristics of the anther collar have been used system- 

 atically in the Eupatorieae and Senecioneae. Endothecial cells of the 

 anthers, visible under a compound microscope after special prepara- 

 tion, have also been of systematic use in the Eupatorieae (King & 

 Robinson, 1970). 



Styles. The style (Hoffmann, 1894: 107, 109; Bentham, 1873: tab. 

 10; Solbrig, 1963: 443; Cabrera, 1974: 54-56; D'Arcy, 1975: fig. 1) is 

 typically a 2-branched shaft which may have an expansion (node) near 

 the base. The basal expansion sometimes is stipitate above the ovary 

 by a slender pedicel. The base of the shaft is frequently immersed in a 

 cupular nectary on the ovary apex. In some species the branches do 

 not separate and the shaft is entire. In most cases the dorsal (abaxial) 

 surface is pubescent and the ventral (adaxial) surface is more or less 

 flat. The stigmatic region is on the edge or ventral surface in a con- 

 figuration characteristic of the tribe. Not always correlated with stig- 

 matic position, several shapes of style branch are common: 



Lactucoid: Branches slender, longitudinally uniform, and sparingly 

 pubescent. The apex is acute or obtuse. This type occurs in the 

 Lactuceae and in pistillate florets of other tribes. 




18 FIELDIANA: BOTANY 



Vernonioid: Branches elongate, longitudinally uniform, and often 

 copiously pubescent. This type occurs in the Vernonieae and 

 Liabeae. 



Eupatorioid: Branches elongate, gradually expanded near the apex, 

 minutely pubescent, papillose, or smooth. It is stigmatic at the mar- 

 gins near the base, and distal portions of the branches may be re- 

 ferred to as appendages. This type occurs in only the Eupatorieae. 



Senecioid: Branches often short, truncate, the apex with a fringe of 

 papillae or hairs (penicillate). This type occurs in some species of 

 Senecioneae, Anthemideae, and Inuleae. 



Helianthoid: Branches are short, pilose near the apex, and sometimes 

 with a triangular or filiform appendage at the tip. This type occurs in 

 several genera of Astereae, Inuleae, and Heliantheae, and inter- 

 grades with the Senecioid type. 



Carduoid: Branches short and smooth, the shaft has an annulus of hairs 

 or thickening near the apex. This type occurs in the Cardueae. 



Ovaries. Taxonomic characters of the ovary are usually expressed 

 in terms of the achene, and younger stages may be misleading. Wings 

 in some Verbesina (Heliantheae) species do not develop until after 

 anthesis, while in Wulffia (Heliantheae) the awn (pappus) is deciduous 

 soon after anthesis. In many groups a copular nectary is present at the 

 apex of the ovary and in some genera, e.g., Ayapana (Eupatorieae), it 

 is conspicuous. This is distinct from the expanded style base which 

 resembles a nectary in some groups. The nectary may be stipitate. It 

 may envelop the basal enlargement of the style shaft or end below it, in 

 which case the stylar expansion appears stipitate. The nectary and 

 style shaft are adnate only at the base. In several tribes Vernonieae, 

 Eupatorieae, Inuleae, Tageteae, Liabeae, and Anthemideae the 

 ovaries are characteristically terete, often ribbed, while in the Astereae 

 and Heliantheae they are often compressed laterally (radially) or dorsi- 

 ventrally (tangentially). 



Fruits (achene s). The usual dispersal unit in the Compositae is the 

 achene, which consists of pericarp, endosperm and embryo, and 

 sometimes includes a pappus, persistent nectary, and carpopodium. 

 The pericarp (rind) is usually hard but is soft and fleshy in Wulffia. The 

 exocarp is sometimes transparent. The achene may be apically nar- 

 rowed into a beak which subtends the pappus, and the top of the beak 

 may be expanded in a flange. All structures surmounting the achene 

 except the nectary are referred to as pappus. This may consist of hairs, 

 bristles, scales (squamellae), awns or rarely glands, and sometimes 

 these elements are fused in a corona or annulus. Bristles or hairs are 




MACBRIDE: FLORA OF PERU 19 



usually strigulose (barbellate, scabrid) and are especially fine and 

 numerous in the Senecioneae and Lactuceae. Stout bristles are some- 

 times basally flattened or expanded. Scales may be lacerate. In the 

 Heliantheae awns are common. While the pappus is of great utility in 

 identifying Compositae, it is not unusual to find epappose (calvous) 

 achenes in individuals or species of normally pappose groups (e.g., 

 Galinsoga}. The carpopodium (hypophysis) is sometimes conspicuous, 

 and the cellular arrangement has been given taxonomic weight in the 

 Eupatorieae. A stipe arising above the carpopodium occurs in some 

 species of Verbesina. 



Frequently, the achene is united with enveloping bracts or paleae or 

 with adjacent florets, and the compound structure falls together. This 

 compound fruit may be termed a utricle in the same sense as the term is 

 used in the Chenopodiaceae and Urticaceae. It has also been known as 

 an involucral fruit or fruiting involucre. The utricle may be flat and 

 winglike or samaroid as in Delila, covered with hooks or spines and 

 burlike as \nAcanthospermum, or the bract may be tightly fused to and 

 hardly distinguishable from the achene as in Melampodium (all 

 Heliantheae). Several achenes (or heads) may be held in glomerules 

 with associated bracts to form a burlike utricle in members of the 

 Vernonieae. 



In a number of Peruvian Compositae the fruit is fleshy and bird- 

 dispersed. The inulin-rich pericarp of Wulffia is soft and fleshy and this 

 baccate fruit is technically a drupe. In Clibadium and Milleria (both 

 Heliantheae) parts of the involucre are fleshy or even juicy and form a 

 baccate structure. The baccate condition is best noted in fresh material 

 and may pass unnoticed when dry. 



Achene shape is sometimes indicative of tribe; thus, the Heliantheae 

 and Cardueae have generally larger achenes than those of other tribes, 

 and in the Lactuceae, Tageteae, and Mutisieae, fruits are often long 

 and thin. Achenes are often compressed in the Astereae and Helian- 

 theae and sometimes in the Lactuceae, but are mostly oblong and 

 cylindrical in the Vernonieae, Liabeae, Eupatorieae, Inuleae, and 

 Senecioneae. Winged achenes occur in the Heliantheae and An- 

 themideae. 



REFERENCES 



BENTHAM, G. 1873. Compositae. In G. Bentham & J. D. Hooker, Genera Plantarum. 

 Vol. 2, pp. 163-533. Lovell Reeve, London. 



CABRERA, A. L. 1974. Compuestas. In A. Burkart, Flora llustradade Entre Rios (Argen- 

 tina). Vol. 6, pp. 106-554. Coleccion Cientifica del I.N.T.A., Buenos Aires. 




20 FIELDIANA: BOTANY 



D'ARCY, W. G. 1975 [1976]. Compositae. //; R. E. Woodson, Jr., et al.. Flora of Panama. 



Ann. Missouri Bot. Gard. 62, pp. 835-1322. 

 HESS, R. 1938. Vergleichende Untersuchungen uber die Zwillingshaare der Compositen. 



Bot. Jahrb. 68, pp. 435-4%. 

 HEYWOOD, V. H., J. B. HARBORNE, & B. L. TURNER. 1977. The Biology and Chemistry 



of the Compositae. Vol. 1 & 2. Academic Press, London. 

 HOFFMANN, O. 1894. Compositae. //; "Die naturlichen Pflanzenfamilien" (Engler and 



Prantl, eds.), 4 (5), pp. 87-387. 



KING, R. M. & H. ROBINSON. 1970. The new synantherology. Taxon 19, pp. 6-11. 

 SOLBRIG, O. T. 1963. The tribes of Compositae in the Southeastern United States. Jour. 



Arnold Arbor. 44, pp. 436-461. 



KEY TO TRIBES OF PERUVIAN CoMPosiTAE 4 



Heads with staminate or perfect florets towards the middle, the corollas tubular or 

 bilabiate; sometimes with pistillate florets towards the outside: usually sap not milky. 

 2. Anther tips with sterile, tonguelike, often hyaline appendages. 



3. Florets all alike, perfect, corollas tubular, not yellow: anthers not tailed: 

 receptacle usually naked. 



4. Leaves alternate; style branches slender, terete, hairy all over, the style 

 shaft apically hairy; anthers auricled (tailed in Piptocarpha); hairs often 

 1-celled Tribe VERNONIEAE 



4'. Leaves mostly opposite (except sometimes in the region of inflorescence): 

 style branches gradually expanded near the tips, papillose or short-hairy, 

 the shaft often glabrous; anthers obtuse or rounded: hairs multicellular. 



often moniliform Tribe EUPATORIEAE 



3'. Florets often not all alike, corollas often yellow; anthers sometimes tailed; 

 receptacle naked or with paleae. 



5. Leaves mostly not spiny; involucral bracts not spiny; anthers tailed or 

 not: style shaft without an apical ring. 



6. Leaves alternate: style branches flattened-fusiform. sometimes api- 

 cally appendaged or rounded: anthers tailed or not: receptacle mostly 

 naked: pappus mostly bristles. 

 7. Anthers obtuse: style branches often appendaged; achene often 



compressed; hairs multicellular Tribe ASTEREAE 



7'. Anthers tailed: style branches rounded: achene plump: hairs 

 arachnoid Tribe INULEAE 



6'. Leaves alternate or opposite; style branches flattened-fusiform, 

 sometimes apically appendaged: anthers not tailed: receptacle with 

 paleae or naked: pappus of bristles, awns or scales. 



"Adapted in part from D'Arcy (1975). 




MACBRIDE: FLORA OF PERU 21 



8. Pappus of awns, bristles or scales; style branches often appen- 

 daged. 



9. Involucre without transparent margins: leaves mostly oppo- 

 site, often 3-nerved from base or trifoliolate. 

 10. Receptacle naked: involucral bracts equal, mostly val- 

 vate (biseriate in Schizothrichia), with pronounced pel- 

 lucid glands: leaves glabrous to puberulent, typically 

 bearing conspicuous pellucid secretory cavities or glands 



filled with strongly scented essential oils 



Tribe TAGETEAE 



10'. Receptacle with paleae, squamellae, bristles or merely 

 deeply alveolate (rarely truely naked): involucral bracts 

 unequal, overlapping, 2- to many-seriate, lacking pel- 

 lucid glands; leaves variously pubescent or glabrous, 

 pellucid glands absent. 



11. Receptacle with costate paleae, enfolding the 

 achenes; achenes usually compressed; pappus of 

 scales, awns, or rarely of numerous, strigose bris- 

 tles; leaves opposite or alternate, mostly eglandular; 



hairs often verrucose Tribe HELIANTHEAE 



11 '. Receptacle deeply alveolate, with the margins of the 

 alveolae prolonged into stiff mostly subulate awns, 

 squamellae or bristles, rarely with true paleae (i.e., 

 Chionopappus) or naked (i.e., Cacosmia, Philo- 

 glossa); achenes usually cylindric to turbinate, (2-) 

 5- to 10-angled; pappus generally biseriate, the inner 

 series of bristles and the outer of bristles or 

 squamellae, rarely absent (i.e., Cacosmia); leaves 

 opposite or whorled in a basal rosette, usually to- 



mentose below Tribe LIABEAE 



9'. Involucre with hyaline, transparent margins: leaves alternate, 

 with strong midrib. 



12. Leaves usually dissected, often aromatic; style branches 

 in disc and ray florets truncate, penicillate; pappus 



paleaceous, coroniform, or absent 



Tribe ANTHEMIDEAE 



12'. Leaves entire, not aromatic; style branches of ray florets 

 filiform, glabrous, and of the disc florets, undivided; 



pappus lacking Tribe CALENDULEAE 



8'. Pappus of soft, silky, hairlike bristles; style branches not appen- 



daged Tribe SENECIONEAE 



5'. Leaves and involucral bracts spiny: anthers tailed; style shaft with an 



apical ring Tribe CARDUEAE 



2'. Anther tips sterile, but not differentiated into hyaline, tonguelike appendages: 



anthers mostly tailed Tribe MUTISIEAE 



1'. Heads with only perfect florets, the corollas ligulate, 5-denticulate; sap milky 



Tribe LACTUCEAE 




FIELDIANA: BOTANY 



Tribe VERNONIEAE 



SAMUEL B. JONES 



Professor of Botany 



University of Georgia 



Athens, Georgia 



Vernonieae Cass., J. Phys. Chim. Hist. Nat. Arts 88: 203. 1819. 

 TYPE: Vernonia Schreb. 



Vernoniaceae Bessey, Ann. Missouri Bot. Card. 2: 163. 1915. TYPE: Vernonia 

 Schreb. 



Perennial or rarely annual herbs, shrubs, trees, or scandent vines. Leaves alternate, 

 rarely opposite or whorled, sometimes in a basal rosette, sessile or petiolate, entire or 

 remotely toothed, rarely lobed, usually revolute. Inflorescences various, heads separate 

 or united in glomerules. Heads discoid, homogamous, 1-many flowered, sometimes re- 

 duced and syncephalous, florets normally bisexual and fertile; involucre usually cam- 

 panulate, ovoid, or globular; phyllaries many, closely or loosely imbricated in several 

 series, or rarely few in one series; receptacle flat or subconvex, either smooth or pitted, 

 rarely alveolate, sometimes with palea-like bracts. Corollas tubular, usually regular (sub- 

 ligulate in Stokesia), tube elongate, with five narrow lobes to the limb, rarely 3-4 lobed, 

 or somewhat bilabiate (e.g., Elephantopus), deep purplish-red to white or blue (rarely 

 yellow-orange in a few Old World species), often glandular; anthers with terminal 

 appendages, basally sagittate, the auricles obtuse, acute or rarely tailed, pollen grains 

 echinate to echinolophate, filaments inserted high above the base; style branches semi- 

 cylindrical, long and slender, narrowed to the acute tips, usually short-hirsute outside, 

 rarely glabrate, stigmatic papillae on the inner surface. Pappus usually elongate and 

 setose, sometimes of scales or coroniform, often in two series, the outer reduced or 

 rarely absent. Achenes variable, terete to slightly flattened, often 10-ribbed or 4- or 

 5-angled, occasionally smooth, rarely dimorphic. 



Vernonieae may be recognized by their usually alternate leaves, 

 their slender, pubescent style branches tapering to slender tips, their 

 involucre of similar imbricate phyllaries in graded series, and (in Peru) 

 by their reddish-purple, or pink to whitish corollas. Vernonieae are 

 most likely to be confused with Eupatorieae since the heads of both are 

 homogamous and their corollas are similarly colored. The leaves of 

 most Vernonieae, however, are alternate as opposed to those of 

 Eupatorieae, which are mostly opposite. In Vernonieae, the stigmatic 

 papillae of the style branches are on the inner surface, but in 

 Eupatorieae, they are restricted to the lower half of the lateral margins. 



The tribe (worldwide) has ca. 1,456 species and over 70 genera. 

 There is little doubt that this tribe originated in the tropics, since that is 

 its center of diversity, the area where its primitive species occur, and 

 the region where the majority of its genera are located. The tribe Ver- 

 nonieae seemingly has two centers of distribution, one in southern 

 Brazil and the second in tropical Africa. Vernonieae are also com- 

 monly found in Southeast Asia and associated archipelagos and in the 




MACBRIDE: FLORA OF PERU 23 



West Indies, Central America, and North America. Carlquist (1976) 

 argues that the tribe originated in the New World. 



Chromosome numbers are known from 16 of the 70 genera of Ver- 

 nonieae. On a worldwide basis, genera with* = 10 predominate, with 

 the second greatest number having x = 9. The Old World Vernonias 

 are dibasic with* = 9, or 10, and have polyploids derived from either 

 base number. Vernonia in the New World has a base number of x = 17 

 which is assumed to represent ancient polyploids derived by aneu- 

 ploidy from a base of x = 9. Cytologically, this tribe has less known 

 about it than any of the other Compositae tribes. 



REFERENCES 



CARLQUIST, S. 1976. Tribal interrelationships and phylogeny of the Asteraceae. Aliso 8, 



pp. 465-492. 

 JONES, S. B. 1977. Vernonieae Systematic review. In Heywood, V. H., J. B. Har- 



borne, and B. L. Turner, The Biology and Chemistry of the Compositae. Vol. I, pp. 



503-521. Academic Press, London. 

 WAGENITZ, G. 1976. Systematics and phylogeny of the Compositae (Asteraceae). PI. 



Syst. Evol. 125, pp. 29-46. 



KEY TO GENERA OF VERNONIEAE 

 a. Heads united in glomerules, syncephalous. 



b. Pappus of straight bristles which are all alike VI. Elephantopus. 



bb. Pappus of bristles, at least two of which are spirally twisted or doubly bent . . . 



VII. Pseudelephantopus. 

 aa. Heads separate from each other, not syncephalous. 



c. Pappus a ring or corona shorter than the achene V. Struchium. 



cc. Pappus of strigose bristles or of scales longer than achene, often biseriate, the 



outer shorter. 



d. Outer phyllaries leaflike, wide-spreading; pappus easily deciduous; inner 



phyllaries usually distinctly awn-tipped IV. Cenlratherum. 



dd. Outer phyllaries scalelike, mostly appressed; pappus persistent; inner phyl- 

 laries acute to acuminate or mucronate. 



e. Heads with 2 (rarely 1 or 3) florets HI. Pollalesta. 



ee. Heads with more than 3 florets. 



f. Inflorescences terminal, composed of scorpioid cymes or becoming 

 paniculate or corymbiform; anthers saggitate at base; pubescence not 



stellate-tomentose I. Vernonia. 



ff. Inflorescences aggregated in rounded axillary corymbs or sessile in 

 rounded axillary clusters. Anthers caudate at base; pubescence often 

 stellate-tomentose II. Piptocarpha. 




24 FIELDIANA: BOTANY 



I. VERNONIA 



Vernonia Schreb., Gen. PI. 2: 541. 1791. nom. cons. TYPE: V. 

 noveboracensis (L.) Willd. 



Serratula noveboracensis L., Sp. PI. 818. 1753. TYPE: S. noveboracensis L. typ. 



cons. 



Behen Hill, Veg. Syst. 4: 41. 1762. TYPE: B. noveboracensis (L.) Hill. 

 Suprago Gaertn., Fruct. 2: 402. 1791. TYPE: 5. glauca Gaertn. 

 Baccaroides Moench, Meth. 578. 1794. TYPE: B. anthelmintica (L.) Moench. 

 Hololepis DC., Ann. Mus. Natl. Hist. Nat. 16: 190. 1810. TYPE: H. pedunculata DC. 

 Teichostemma R. Br. ex Salt, Abyss. App. 65. 1814. TYPE: T.fruticosum R. Br. 

 Bracheilema R. Br. ex Salt, Abyss. App. 65. 1814. TYPE: B. paniculatum R. Br. 

 Ascaricida Cass., Diet. Sc. Nat. 3: Suppl. 38. 1816. TYPE: A. indica Cass. 

 Centrapalus Cass., Diet. Sc, Nat. 7: 382. 1817. TYPE: C. galamensis Cass. 

 Isonema Cass., Bull. Soc. Philom. Paris 1817: 152. 1817. TYPE: /. ovata Cass. 

 Distephanus Cass., Bull. Soc. Philom. Paris 1817: 151. 1817. TYPE: Conyza 



populifolia Lam. 

 Lepidaploa Cass., Bull. Soc. Philom. Paris. 1817: 66. 1817. TYPE: V. glauca (L.) 



Willd. 

 Gymnanthemum Cass., Bull. Soc. Philom. Paris 1817: 10. 1817. TYPE: G. congestum 



Cass. 

 Turpinia Lex. ex LaLlave & Lex., Nov. Veg. Desc. fasc. 1: 22. 1824. TYPE: T. 



tomentosa Lex. ex LaLlave & Lex. 



Acilepsis D. Don, Prod. Fl. Nep. 169. 1825. TYPE: A. squarrosa D. Don. 

 Cyanthillium Bl., Bijdr. 889. 1826. TYPE: C. moluccense Bl. 

 Achyrocoma Cass., Diet. Sc. Nat. 5: 57. 1828. TYPE: A. tomentosa Cass. 

 Cyanopis Bl. ex DC., Prodr. 5: 69. 1836. TYPE: C. villosa (Bl.) DC. 

 Plectreca Raf., Fl. Tellur. 4: 119. 1836. TYPE: P. corymbosa (Schwein.) Raf. 

 Webbia DC., Prodr. 5: 72. 1836. TYPE: W. pinifolia (Less.) DC. 

 Monosis DC., Prodr. 5: 77. 1836. TYPE: M. wightiana DC. ex Wight. 

 Keringa Raf., Sylva Tellur. 144. 1838. TYPE: K. amygdalina (Delile) Raf. 

 Flustula Raf., Sylva Tellur. 116. 1838. TYPE: F. tomentosa Raf. 

 Candidea Ten., Atti Accad. Sci. Fis. 4: 104. 1839. TYPE: C. senegalensis Ten. 

 Cyanopsis Endl., Ench. 232. 1841. 

 Trianthaea Spach, Hist. Veg. Phan. 10: 39. 1841. 



Linzia Sch. Bip., Flora 24. I. Intell. 26. 1841. TYPE: L. glabra (Steetz) Sch. Bip. 

 Cheliusia Sch. Bip., Flora 24. I. Intell. 26. 1841. TYPE: C. abyssinica Sch. Bip. 

 Stengelia Sch. Bip., Flora 24. I. Intell. 26. 1841. TYPE: S. adoensis Sch. Bip. 

 Polydora Fenzl, Flora 27: 312. 1844. TYPE: P. stoechadifolia Fenzl. 

 Claotrachelus Zoll., Natuur- Geneesk. Arch. Ned. -Indie. 2: 565. 1845. TYPE: C. 



rupestris Zoll. & Mor. 



Leiboldia Schlecht., Linnaea 19: 742. 1847. TYPE: L. leiboldiana Schlecht. 

 Vernonella Sond., Linnaea 23: 62. 1850. TYPE: V. africana Sond. 

 Llerasia Triana., Ann. Sci. Nat. Ser. 4: 10. 1858. TYPE: L. lindeni Triana. 




MACBRIDE: FLORA OF PERU 25 



Strobocalyx Sch. Bip., Pollichia 28/29: 170. 1861. TYPE: 5. arborea (Buch.-Ham.) 



Sch. Bip. 

 Crystallopollen Steetz ex Peters., Reise Mossamb. Bot. part 6: 363. 1862-1864. TYPE: 



C. angustifolium Steetz. 

 Ambassa Steetz ex Peters., Reise Mossamb. Bot. part 6: 346. 1862-1864. TYPE: A. 



hochstetteri (Sch. Bip. ex Hochst.) Steetz ex Peters. 

 Xipholepis Steetz ex Peters., Reise Mossamb. Bot. part 6: 344. 1862-1864. TYPE: X. 



silhetensis (DC.) Steetz. 

 Punduana Steetz ex Peters., Reise Mossamb. Bot. part 6: 345. 1862-1864. TYPE: P. 



volkameriaefolia (DC.) Steetz ex Peters. 

 Lysistemma Steetz ex Peters., Reise Mossamb. Bot. part 6: 340. 1862-1864. TYPE: L. 



indica (Wall, ex Clarke) Steetz ex Peters. 

 Stenocephalum Sch. Bip., Pollichia 20/21: 385. 1863. TYPE: 5. monticolum (DC.) Sch. 



Bip. 

 Tephrothamnus Sch. Bip., Pollichia 20/21: 431. 1863. TYPE: T. pycnanthus (Benth.) 



Sch. Bip. 



Critoniopsis Sch. Bip., Pollichia 20/21: 430. 1863. TYPE: C. lindenii Sch. Bip. 

 Senecioides Post & O. Ktze., Lex. Gen. Phan. 2: 515. 1903. TYPE: S. cinereum (L.) 



Post & O. Ktze. 

 Eremosis (DC.) Gleason, Bull. New York Bot. Card. 4: 227. 1906. TYPE: E. salicifolia 



(DC.) Gleason. 



Perennial herbs, shrubs, or small trees, scandent lianas, or rarely annuals. Leaves 

 alternate, simple, pinnately veined, usually cauline, or sometimes basal in herbaceous 

 perennials: blades various, lanceolate to ovate or elliptic. Inflorescences terminal or upper 

 axillary or scorpioid cymes, panicles, corymbs, of combinations thereof, or reduced to 

 solitary terminal or axillary heads. Heads discoid, homogamous, with 1-many florets; 

 involucre cylindric to broadly hemispheric or campanulate; phyllaries loosely or closely 

 imbricate in several series, the inner phyllaries progressively longer; receptacle flat to 

 subconvex. Corollas tubular, regular, 5-lobed, deep reddish purple to whitish or pinkish 

 (blue and yellow in the Old World); often slightly glandular; anthers sagittate at the base; 

 style branches elongate, filiform-subulate, outer surface hispid throughout, with 

 stigmatic pappillae on inner surfaces. Pappus usually in 2 series, the inner pappus of 

 capillary, terete, or slightly flattened, purple to white bristles; the outer series short, of 

 bristles or scales, or pappus bristles subequal and not in distinct series. Achenes ribbed or 

 sometimes ribless, commonly resinous-dotted between the ribs. Chromosome number: 

 x = 17 in New World. 



KEY TO SPECIES OF Vernonia* 

 a. Heads with 7 or fewer florets. 



b. Inner pappus bristles ca. 3.5 mm long: corollas ca. 3 mm long . 



1. V. pycnantha. 



5 As the present manuscript went to press, two additional Vernonia species were 

 described from Peru; see Robinson, H., 1980. Phytologia 45(2): 158-165. M.O.D. 




26 FIELDIANA: BOTANY 



bb. Inner pappus bristles 4 mm or more long; corollas 5 mm or more long. 



c. Leaves glabrate or with scattered small trichomes beneath: inflorescences 



large (2-3 dm broad and tall) with scorpioid-cymose branches 



21. V. cainarachiensis. 



cc. Leaves tomentose, softly pubescent, or with tomentum beneath; in- 

 florescences smaller (less than 2 dm broad), branches not scorpioid. 

 d. Inner pappus bristles ca. 9 mm long; corollas ca. 8 mm long; achenes 



strigose 2. V. lambayequensis. 



dd. Inner pappus bristles ca. 6.5 mm or less long; corollas 7 mm or less 

 long; achenes glandular to sparsely pilose. 



e. Leaf blades 3.5-6 cm long, 1 .5-2.4 cm wide, coriaceous 



3. V. jalcana. 

 ee. Leaf blades 7-20 cm long, 2.5-5.5 cm wide, not coriaceous. 



f. Achenes sparsely pilose; inner phyllary tips obtuse; leaf blades 



elliptic to ovate 4. V. woytkowskii. 



ff. Achenes glabrous to glandular; inner phyllary tips acute; leaf 

 blades lanceolate to long-elliptic. 



g. Inner pappus bristles 6.5 mm long; heads with 4-5 florets; 

 pappus white; leaf blades tomentose beneath, with scat- 

 tered longer dark-brown villous trichomes arising above the 



tomentum 5. V. peruviana. 



gg. Inner pappus bristles 5 mm long; heads with 5-7 florets; 

 pappus straw-colored; leaf blades tomentose beneath with 



no long villous trichomes 6. V. jelskii. 



aa. Heads with 8 or more florets. 



h. Heads with more than 50 florets. 



i. Heads with 80-90 florets; corollas ca. 5.5 mm long; leaf blades rigid or coria- 

 ceous 7. V. libertadensis. 



ii. Heads with ca. 50 florets; corollas ca. 2.5 mm long; leaf blades thin 



8. V. gracilis. 

 hh. Heads with 36 or less florets. 



j. Pappus straw-colored, brown or pinkish. 



k. Inner pappus bristles ca. 10-11 mm long, corollas 12-13 mm long. 

 1. Heads with ca. 20 florets; corolla throats glandular; phyllary tips 



acute; inflorescences of axillary, leafy cymes 9. V. laurifolia. 



11. Heads with ca. 12 florets; corolla throats glandular; phyllary tips 



acuminate; inflorescences paniculate-corymbose 



10. V. sordidopapposa. 



kk. Inner pappus bristles ca. 7 mm or less long; corollas 10 mm or less long, 

 m. Corollas ca. 10mm long; heads with 7-13 florets; inner phyllary tips 



obtuse: pappus pinkish 21. V. cainarachiensis. 



mm. Corollas ca. 8 mm long; heads with 14-26 florets; inner phyllary tips 

 acute to long-acuminate; pappus straw-colored to brown, 

 n. Outer pappus of fimbriate scales ca. 1.2 mm long: pappus light 

 brown; corollas ca. 8 mm long; inner phyllary tips long- 

 acuminate 1 1 . V. mapirensis. 




MACBRIDE: FLORA OF PERU 27 



nn. Outer pappus of bristles 0.8 mm or less long; pappus straw- 

 colored; corollas ca. 6.5 mm or less long; inner phyllary tips 

 acute to slightly acuminate, 

 o. Leaf blades densely tomentose beneath, oblong-elliptic; 



achenes faintly strigose 12. V. ferruginea. 



oo. Leaf blades glabrate to hispid or downy beneath, elliptic 

 to broadly elliptic or ovate-lanceolate; achenes 



glandular-hispid 16. V. patens. 



Pappus white, 

 p. Inflorescences paniculate-corymbose or cymose. 



q. Inner bristles of pappus ca. 6-7 mm long 17. V. fulta. 



qq. Inner bristles of pappus ca. 4.5 mm or less long, 

 r. Leaf blades 2-6 cm long, 1-2.7 cm wide. 



s. Corollas ca. 9 mm long; leaf blades cordate to ovate or 

 ovate-elliptic, densely white tomentose beneath; inner 



phyllary tips long-acuminate 18. V. apurimacensis. 



ss. Corollas ca. 4.5-5 mm long; leaf blades lanceolate, glabrate 

 beneath; inner phyllary tips acute to obtuse or mucronate . 



14. V. stuebellii. 

 rr. Leaf blades ca. 12-26 cm long, ca. 5-15 cm wide. 



t. Heads with ca. 36 florets; leaf blades elliptic to elliptic- 

 oblong, villous beneath 13. V. costata. 



tt. Heads with ca. 20 florets; leaf blades ovate to ovate- 

 lanceolate, tomentose beneath 15. V. sambrayana. 



pp. Inflorescences scorpioid-cymose or somewhat scorpioid-paniculate. 

 u. Leaf blades 3.5 cm or less long. 



v. Inner phyllary tips slightly recurved; heads with 14-24 florets . 



19. V. scorpioides. 



vv. Inner phyllary tips flat or straight; heads with 11-13 florets, 

 w. Corollas ca. 5 mm long; leaf blades ca. 1.9 cm wide, 

 closely pubescent with minute slender hairs, ovate-oblong 



to elliptic-ovate 25. V. fieldiana. 



ww. Corollas ca. 8 mm long; leaf blades ca. 2.5 cm wide, vil- 

 lous to hirsute with straw-colored trichomes; obovate to 



obovate-lanceolate 27. V. herbacea. 



uu. Leaf blades (4)6-70 cm long. 



x. Inner phyllary tips slightly recurved 19. V. scorpioides. 



xx. Inner phyllary tips flat or straight. 



y. Achenes brownish, with round glandular trichomes 



22. V. yurimaguasensis. 

 yy. Achenes not brownish, with hairlike trichomes. 



z. Leaf blades minutely or sparsely pubescent beneath; 

 inner phyllary tips acute to acuminate or fimbriate. 

 a' Achenes strigose; leaf blades 10-17 cm long, 3.5-7 

 cm wide 23. V. myriocephala. 




28 FIELDIANA: BOTANY 



aa' Achenes sparsely pubescent; leaf blades 20-70 cm 



long, 8-19 cm wide 20. V. brachiata. 



zz. Leaf blades densely or sparsely strigose or strigose- 

 hirsute beneath; inner phyllary tips acute, subulate or 

 spinose. 



b' Inner pappus bristles 4 mm long; corollas pinkish 

 to whitish; leaf blades 4-7 cm wide; inflorescences 

 of scorpioid cymes arranged in spreading panicles 



or corymbs 24. V. canescens. 



bb' Inner pappus 6-8 mm long; corollas reddish- 

 purple; leaf blades 1.5-3 cm wide; inflorescences 



divaricately spreading scorpioid cymes 



26. V. salzmannii. 



1. Vernonia pycnantha Benth., PI. Hartw. 134. 1844. TYPE: in mon- 

 tibus Paccha (K, not seen). 



Critoniopsis lindenii Sch. Bip., Pollichia 20/21: 431. 1863. TYPE: Colombia: Quindiu, 



Los Volcancitos, Linden 1054 (Holotype P, as photo F!). 

 Vernonia lindenii (Sch. Bip.) Cuatr., Bot. Jahrb. Syst. 77: 72. 1956. 



Shrub with long scandent branches, sometimes forming a tree, young stems 

 brownish-tomentose to almost glabrate. Leaves cauline, petiolate; petiole ca. 0.8-1.5 cm 

 long; blades ovate-elliptic, elliptic, or elliptic-lanceolate, acuminate to acute at the apex, 

 cuneate to cuneate-rounded at the base, ca. 8-15 cm long, ca. 3.5-7 cm wide, margins 

 re volute, and sometimes remotely toothed, largely glabrous but remotely glandular 

 above, glabrate and glandular to tomentose beneath. Inflorescence of terminal, corym- 

 bose cymes with reduced bracteal leaves along main axis. Heads with ca. 6 florets, sessile 

 in dense pedunculate clusters; involucres campanulate, ca. 4 mm long, loosely im- 

 bricated; phyllaries soon deciduous, glabrous to slightly pubescent, green, tipped with 

 purple; inner phyllaries oblong, tips rounded; outer phyllaries ovate. Corollas ca. 3 mm 

 long. Pappus white; inner bristles 3.5 mm long, outer bristles ca. 1 mm long. Achenes ca. 

 2.2 mm long, ribbed, lightly strigose. 



This species is distributed from Ecuador south to Peru. In Peru, it 

 has been collected at 1,750 m elevation within a forest border. Flower- 

 ing and fruiting occur from July to September. 



HUANUCO: Churubamba, Mexia 8229 (F, MO, NY, UC). 



2. Vernonia lambayequensis S. B. Jones, sp. nov. TYPE: Peru: Lam- 

 bayeque: km 28 E of Olmos, Hutchison and Wright 3473 (Holotype 

 UC! Isotypes F! MO! USM!). 



Frutex 2.5 m altus. Foliorum laminae ellipticae ad elliptico-obovatae, ca. 8-12 cm 

 longae, ca. 4-5 cm latae. Inflorescentia terminalis, paniculato-corymbiformis, 

 capitulis in fasciculos compactos, rotundatos, conspicue aggregatis. Capitula 5 flos- 

 culos habentia. Achenia strigosa. 



Erect shrub, up to 2.5 m tall, young stems canescent. Leaves cauline; petioles ca. 0.7-1 

 cm long; blades elliptic to elliptic-obovate, acute to rounded or mucronate at the apex, 

 cuneate at the base, ca. 8-12 cm long, 4-5 cm wide, margins revolute, very faintly 




MACBRIDE: FLORA OF PERU 29 



toothed, glabrate to slightly canescent above, veins canescent above, softly pubescent 

 beneath. Inflorescences terminal, paniculate-corymbiform, heads grouped in compact, 

 rounded clusters within the inflorescence, branches canescent. Heads with 5 florets, 

 sessile; involucres cylindric, ca. 6 mm long, 4- to 5-seriate; phyllaries canescent and dark 

 at tips, yellowish; inner phyllaries oblong-lanceolate, tips obtuse to acute; outer phyllaries 

 ovate, arachnoid. Corollas ca. 8 mm long, pale purple to almost white, glandular on tube. 

 Pappus white; inner bristles ca. 9 mm long, outer bristles ca. 1 mm long. Achenes ca. 3.5 

 mm long, strigose, ribbed. 



This species is known only from the type location in Depto. Lam- 

 bay eque, where it was collected at 1,150 to 1,200 m elevation. Habitat 

 information was not available on the label; however, it was described 

 as being rare. 



3. Vernonia (alcana Cuatrec., Ann. Missouri Bot. Gard. 52: 312. 

 1965. TYPE: Peru: Amazonas: Prov. Chachapoyas, Molinopampa. 

 Wurdack 1359 (Holotype US, Isotype UC!). 



Shrub, 1.5-2 m tall; stems grayish to brownish-tomentose to almost black, with scat- 

 tered long purple trichomes. Leaves crowded, coriaceous; petiolate, petioles ca. 7 mm 

 long; blades ovate-elliptic, acute at the apex, cuneate to slightly rounded at the base, 

 1.5-2. 4 cm long, 3. 5-6 cm wide, margins entire, upper surface reticulate and tomentose on 

 lower part of midvein, gray tomentose, with scattered long purple trichomes beneath. 

 Inflorescences densely corymbose-paniculate. Heads with 3 florets, compact and almost 

 sessile; involucres campanulate-cylindric, 7-8.5 mm long, 5- to 6-seriate; phyllaries 

 arachnoid, glandular near tips, tightly appressed, purplish; inner phyllaries oblong, tips 

 acute; outer phyllaries lanceolate. Corollas ca. 6-7 mm long, reddish-purple, with scat- 

 tered glands. Pappus white; inner bristles ca. 6.5 mm long, outer bristles ca. 1-1.5 mm 

 long. Achenes 3 mm long, glandular, very faintly ribbed. 



This species occurs in Depto. Amazonas in the jalca zone (north 

 Peruvian paramo) at 2,000-3,000 m elevation. Flowering and fruiting 

 occur in June. 



AMAZONAS: Chachapoyas, Cerros Calla Calla, 19 km above 

 Leimebama on road to Balsas, Hutchison and Wright 5515 (F, MO, 

 NY, USM); Bongara, 3 km S of Pomacocha, Wurdack 971 (F, USM). 

 CAJAMARCA: Cutervo: Cerros de Cutervo, 2,500-2,600 m, Ferreyra 

 0810 (USM). 



4. Vernonia woytkowskii S. B. Jones, sp. nov. TYPE: Peru: Lam- 

 bayeque: Porcullaad Olmos, Woytkowski 6770 (Holotype MO! Isotype 

 GA!). 



Frutex scandens, ca. 7 m altus, caulibus dense canescentibus. Foliorum laminae 

 ellipticae vel oblongo-ellipticae vel ovatae, ca. 7-12 cm longae, ca. 4-5 cm latae. 

 Inflorescentia terminalis, compacta, capitulis dense conglomeratis. Capitula 5-6 

 flosculos habentia. Achenia sparsim pilosa. 



Liana, ca. 7 m tall, young stems densely canescent. Leaves cauline; petioles canescent, 

 ca. 1 cm long; blades elliptic, oblong-elliptic, or ovate, acute to obtuse at the apex, 




30 FIELDIANA: BOTANY 



cuneate at the base, 7-12 cm long, 4-5 cm wide, margins mostly entire, slightly revolute, 

 very remotely fine-toothed, finely and remotely canescent above, softly tomentose and 

 with raised veins beneath. Inflorescences terminal, very compact (actually forming a 

 dense mass of heads) corymbose-paniculate, bracts only at very base of inflorescence. 

 Heads with 5-6 florets, sessile; involucres cylindric-campanulate, 5.5 mm long, 4- to 

 5-seriate; phyllaries pubescent at tips, wide spreading when mature and deciduous along 

 with achenes; inner phyllaries oblong, tips obtuse, dark brown; outer phyllaries obtuse. 

 Corollas ca. 5.5. mm long, white (from label), sparsely glandular. Pappus whitish; inner 

 bristles ca. 4.5 mm long, outer bristles ca. 0.5 mm long. Achenes ca. 2.4 mm long, very 

 sparsely pilose, faintly ribbed. 



This species is known only from the type location in Depto. Lam- 

 bayeque. It was collected on the barren slope of a hill, sprawling upon 

 Cereus at an elevation of 2,100 m. It apparently flowers and fruits in 

 August and September. 



5. Vernonia peruviana Cuatrec., Bot. Jahrb. Syst. 77: 75. 1956. 

 TYPE: Peru: Villcabamba, Hacienda on Rio Chinchao, Macbride 5150 

 (Holotype F! as photo F! Isotype NY). 



Shrub 3-4 m tall, with spreading branches, younger stems pubescent. Leaves cauline, 

 coriaceous; petioles pubescent, 1-2.5 cm long; blades oblong-lanceolate to oblong- 

 elliptic, acute to slightly acuminate at the apex, rounded or obtuse at the base, 10-20 cm 

 long, 3-5.5 cm wide, margins mostly entire, but sometimes remotely toothed, slightly 

 revolute, mostly glabrate except pubescent along midvein above, densely tomentose 

 beneath, also having scattered dark brown, villous trichomes beneath. Inflorescences 

 terminal, paniculate-corymbose. Heads with (4)5 florets, mostly sessile or subsessile; 

 involucre broadly campanulate, ca. 6 mm long, 5- to 6-seriate; phyllaries arachnoid to 

 ciliate, mostly deciduous when achenes mature; inner phyllaries oblong, tips acute; outer 

 phyllaries ovate. Corollas ca. 7 mm long. Pappus white; inner bristles ca. 6.5 mm long, 

 outer bristles ca. 2-4 mm long. Achenes ca. 3 mm long, glabrous or sparsely glandular, 

 ribbed. 



This species is known only from the type location where it was 

 collected on a mountain slope at 2,000 m elevation. Flowering and 

 fruiting occur in July and August. 



6. Vernonia jelskii Hieron., Bot. Jahrb. Syst. 36: 459. 1905. TYPE: 

 Tambillo, Jelski 602 (Holotype B, as photo F! Isotype MO!). 



V. jelskii Hieron. var. virescens Hieron., Bot. Jahrb. Syst. 36: 459. 1905. TYPE: Peru: 

 Tambillo, Jelski 623 (Holotype B, not seen). 



Shrub, stems slightly brownish-tomentose. Leaves cauline, prominately pinnately 

 nerved; petiolate, petioles ca. 1 cm long with brownish tomentum; blades narrowly, 

 long-elliptic, acuminate at the apex, cuneate at the base, 12-18 cm long, 2.5-4 cm wide, 

 margins revolute, glabrous above, reticulate veined, finely glandular, and with tomentum 

 beneath. Inflorescences paniculate-corymbose, leafy. Heads with 5-7 florets, sessile; in- 

 volucres campanulate, ca. 6 mm long, loosely imbricate, 6- to 9-seriate; phyllaries 

 arachnoid with tomentum at base, loosely appressed, brownish-straw colored; inner 




MACBRIDE: FLORA OF PERU 31 



phy Maries oblong, deciduous, tips acute to slightly fimbriate; outer phyllaries lanceolate, 

 tips acute. Corollas ca. 5 mm long, light reddish-purple, glandular. Pappus straw-colored; 

 inner bristles 5 mm long, outer bristles 0.5-0.7 mm long. Achenes ca. 2.5 mm long, 

 glandular, slightly ribbed. 



This species is known only from the type location of Tambillo. Flow- 

 ering and fruiting occur in August. 



7. Vernonia libertadensis S. B. Jones, sp. nov. TYPE: Peru: La 

 Libertad: Otuzco: Cerro Sango (Motil-Shorey), Lopez 1947 (Holotype 

 GA!). 



Frutex caule glanduloso. Folia rigida, laminis ca. 2.5 cm longis, ca. 0.8-1 cm latis, 

 resinoso-glandulo-punctatis. Inflorescentia parva, terminalis, corymbosa. Capitula 

 80-90 flosculos habentia. Involucrum 10-11 mm longum. Setae pappi subaequales, 

 ca. 7 mm longae. 



Shrub, stems glandular. Leaves rigid, cauline, crowded, sessile; blades oblong- 

 lanceolate, obtuse to acute at the apex, cuneate at the base, ca. 2.5 cm long, ca. 0.8-1 cm 

 wide, margins entire, resinous, glandular-punctate both above and beneath. In- 

 florescences relatively small, terminal, corymbose-cymose, the few heads terminal on 

 short branches, the heads subtended by bracteal leaves which are only slightly reduced 

 from the cauline leaves. Heads with 80-90 florets; involucres campanulate, 10-11 mm 

 long, ca. 6-seriate; phyllaries slightly fimbriate, resinous, tightly appressed; inner phyl- 

 laries oblong, tips obtuse to rounded or cuspidate; outer phyllaries oblong-lanceolate to 

 ovate-lanceolate. Corollas ca. 5.5 mm long, reddish-purple, tube slender. Pappus straw- 

 colored; bristles in one series ca. 7.5 mm long. Achenes ca. 2.4 mm long, ribbed, remotely 

 strigose. 



This species is known only from the type location where it was 

 collected at the border of a field at an elevation of 3,300 to 3,400 m. 

 Flowering and fruiting occur in June and July. 



8. Vernonia gracilis H.B.K., Nov. Gen. & Sp. 4: 34. 1820. TYPE: 

 Colombia: Turbaco, Humboldt and Bonpland 1439 (Holotype P, as 

 IDC microfiche!). 



V. moritziana Sch. Bip., Linnaea 20: 511. 1847. TYPE: Venezuela (not seen). 

 Cacalia gracilis (H.B.K.) O. Ktze., Rev. Gen. PI. 970. 1891. 

 C. moritziana (Sch. Bip.) O. Ktze., Rev. Gen. PI. 970. 1891. 



Annual herbs, 2-3 dm tall, stems reddish-purple, sparsely strigose. Leaves cauline, 

 thin; petioles 0-5 mm long; blades lanceolate, elliptic to elliptic-lanceolate, rounded to 

 acute at the apex, cuneate at the base, 4-5 cm long, 1-1.6 cm wide, margins remotely 

 toothed, minutely scabrous above, glandular punctate and remotely pubescent beneath. 

 Inflorescences cymose, weakly branching, bracteal leaves present and similar to the stem 

 leaves. Heads with ca. 50 florets, sessile; involucres broadly campanulate, ca. 5 mm long, 

 3- to 4-seriate; phyllaries minutely glandular, ciliate, arachnoid, greenish; inner phyl- 

 laries oblong-lanceolate, tips acuminate; outer phyllaries ovate-lanceolate. Corollas 

 pinkish, ca. 2.5 mm long. Pappus straw-colored, of indurate, thick bristles; inner 2.5 mm 

 long, outer 0.3 mm long. Achenes ca. 2 mm long, slightly pubescent, ribbed. 




FIELDIANA: BOTANY 



This species is distributed from northern South America south into 

 Peru. Only one collection has been seen. It was flowering in Septem- 

 ber. 



LORETO: Rio Mamon near Rio Nanay, Croat 19916 (MO, NY). 



9. Vernonia laurifolia DC., Prodr. 5: 30. 1836. TYPE: (G-DC, as IDC 

 microfiche!). 



Cacalia laurifolia (DC.) O. Ktze., Rev. Gen. PI. 2: 970. 1891. 



Herb 1 m tall, stems brownish-tomentose. Leaves coriaceous, cauline; petiolate. 

 petiole 0.3-0.8 cm long; blades elliptic to lanceolate, acute at the apex, cuneate to 

 rounded at the base, ca. 5-7.5 cm long, 2-3.5 cm wide, margins revolute, glabrous except 

 along midvein above, glandular and prominently veined beneath. Inflorescences of axil- 

 lary leafy cymes, heads usually arising in the internodes of the bracteal leaves. Heads 

 with ca. 20 florets, long peduncled; involucres narrowly campanulate, ca. 14 mm long, 

 tightly imbricate, 7-seriate; phyllaries slightly arachnoid at base, reddish-purple; inner 

 series of phyllaries linear-lanceolate and much longer than the other series, tips acute; 

 outer phyllaries lanceolate. Corollas ca. 13 mm long, reddish-purple, glandular on outer 

 throat. Pappus light brown; inner bristles ca. 10 mm long, outer bristles ca. 2 mm long. 

 Achenes (immature) brownish-pubescent. 



This species has been collected in Depto. Puno at elevations of 1,900 

 m, growing in a moist, shady place in rocky soil. Flowering and fruiting 

 occur from May to June. 



PUNO: Carabaya: trail Santo Domingo to Chabucamine, Metcalf 

 30660 (MO, UC, US). 



10. Vernonia sordidopapposa Hieron., Bot. Jahrb. Syst. 22: 697. 

 1897. TYPE: Peru: Sandia, Weberbauer 759 (Holotype B, as photo F! 



NY!). 

 Cacalia sordidopapposa (Hieron.) O. Ktze., Rev. Gen. PI. 2: 971. 1891. 



Shrub 1-2 m tall, stems strigose to long strigose-pilose. Leaves cauline; petioles 3-4 mm 

 long; blades elliptic-lanceolate, acuminate at the apex, broadly cuneate at the base, 3-8 

 cm long, 1.5-2.5 cm wide, margins distinctly revolute, pilose-hispid and reticulate veined 

 above, glandular and pilose-hispid to pilose beneath. Inflorescences paniculate- 

 corymbose, with bracteal leaves. Heads with ca. 12 florets, subsessile; involucres nar- 

 rowly campanulate, ca. 7 mm long, imbricate, 3- to 4-seriate; phyllaries ciliate, arachnoid 

 to pilose-hispid, appressed, greenish-purple; inner phyllaries oblong-lanceolate, tips 

 acuminate; outer phyllaries lanceolate. Corollas ca. 12 mm long, reddish-purple, gla- 

 brous. Pappus brown; inner bristles ca. 11 mm long, outer scales fimbriate ca. 1.5 mm 

 long. Achenes ca. 2 mm long, pilose, faintly ribbed. 



This species is distributed from Depto. Amazonas south to Depto. 

 Puno at elevations of 2,400 to 3,400 m. It grows in the jalca zone and 

 puna in moist soil. Flowering and fruiting occur from May to June. 




MACBRIDE: FLORA OF PERU 33 



AMAZON AS: Chachapoyas, west of Molinopampa, Wurdack 1371 

 (NY, US). PUNO: Sandia, near Limbani, Metcalf 30513 (MO). 



11. Vernonia mapirensis Gleason, Amer. J. Bot. 10: 307. 1923. 

 TYPE: Bolivia: Mapiri, Buchtien 1533 (Holotype NY!). 



V. trichoclada Gleason, Bull. Torrey Bot. Club 52: 184. 1925. TYPE: Peru: La 

 Merced, Hacienda Schunke, Macbride 5775 (Holotype F! as photo F! Isotype NY). 



Perennial herb, erect, ca. 3.5 m tall, stems long hirsute-villous. Leaves cauline: petioles 

 ca. 1 cm long; blades elliptic-ovate, acuminate at the apex, rounded at the base, 10-14 cm 

 long, 5-6 cm wide, margins revolute, slightly crenate and remotely callus toothed, 

 rugose, slightly pubescent above, hirsute-villous on midvein above, rugose and hirsute- 

 villous beneath. Inflorescences paniculate to cymose. Heads with (10)14-20(23) florets, 

 sessile: involucres campanulate, 8-9 mm long, imbricate, ca. 4-seriate; phyllaries ciliate, 

 loosely appressed, greenish to reddish-purple; phyllaries long-lanceolate, tips long- 

 acuminate. Corollas ca. 8 mm long, reddish-purple, glabrous. Pappus light brown; inner 

 bristles ca. 7 mm long, outer scales fimbriate, ca. 1.2 mm long. Achenes 3 mm long, 

 densely pilose. 



This species occurs in Peru from Depto. Junin south to Depto. Puno 

 at elevations of 1,300 to 2,600 m in open areas in the mountains. Flow- 

 ering and fruiting occur from June to September. 



JUNIN: La Merced, Macbride 5775 (F). CUZCO: Tambopata, 

 Machupijcho, Vargas 13539 (US). PUNO: Sandia, 2-6 km Oconeque, 

 Metcalf 30603 (UC). 



12. Vernonia ferruginea Less., Linnaea 4: 271. 1829. TYPE: Brasil: 

 Sellow s.n. (not seen). 



Cacalia ferruginea (Less.) O. Ktze., Rev. Gen. PI. 2: 970. 1891. 



A small tree or shrub, 2-4 m tall, crown bushy, stems tomentose. Leaves cauline; 

 petiolate, petioles 0.5-1 cm long; blades oblong-elliptic, obtuse at the apex, truncate to 

 slightly rounded at the base, 8-16 cm long, 3-5 cm wide, margins remotely callus toothed, 

 revolute, undulate to crenate, arachnoid to glabrate, tomentose on large veins above, 

 tomentose beneath. Inflorescences paniculate-cymose with slightly scorpioid branches. 

 Heads with 20 to 26 florets, sessile; involucres campanulate, 5-6.5 mm long, ca. 5-seriate; 

 phyllaries arachnoid-tomentose , appressed, greenish with lighter green margins; inner 

 phyllaries ovate-lanceolate, tips acute to slightly acuminate; outer phyllaries oblong- 

 lanceolate. Corollas 4.5-5 mm long, reddish-purple, sometimes slightly glandular. Pappus 

 straw-colored; inner bristles 3.5-4 mm long, outer bristles ca. 0.7 mm long. Achenes ca. 

 1.8 mm long, faintly strigose, weakly ribbed. 



This species is distributed from Depto. Junin south to Depto. Cuzco 

 into Brazil at elevations of 800 to 1 ,000 m on open hillsides and grassy 

 slopes. Flowering and fruiting occur from June to August. 



JUNIN: San Ramon, Killip and Smith 24780 (F, NY, US). CUZCO: 

 Convencion, Chahuares, Vargas 21674 (US). 




34 FIELDIANA: BOTANY 



13. Vernonia costata Rusby, Mem. Torrey Bot. Club 6: 53. 1896. 

 TYPE; Bolivia: Mapiri, Rusby 1472 (not seen). 



Slender, erect shrub, 1-2 m tall, stems brownish-tomentose to villous. Leaves 

 cauline; petioles brownish-villous, 1-1.5 cm long; blades elliptic to elliptic-oblong, acute 

 to acuminate at the apex, cuneate to slightly rounded at the base, 12-26 cm long, 5-15 cm 

 broad, margins re volute, sometimes with callous teeth, villous wide, densely brownish- 

 villous and prominently veined beneath. Inflorescences cymose-paniculate. Heads with 

 ca. 36 florets, sessile; involucres campanulate, 7-8 mm long, 6- to 7-seriate; phyllaries 

 slightly arachnoid, tightly appressed, greenish to reddish-purple; inner phyllaries long- 

 lanceolate, tips subacute; outer phyllaries lanceolate. Corollas ca. 5 mm long; reddish- 

 purple, glandular and hairy on outside of lobes. Pappus white; inner bristles ca. 4.5 mm 

 long, outer bristles ca. 0.8 mm long. Achenes 2-3 mm long, strigose. 



This species is distributed from Depto. Junin to Depto. Cuzco 

 south into Bolivia at elevations of 600 to 1 ,300 m, growing in thickets 

 and thin woods. Flowering and fruiting occur from June to August. 



JUNIN: Colonia Perene, Killip and Smith 25012 (F, NY, US). 

 CUZCO: Convencion, Cuesta de Ichiquiato, Vargas 14495 (US). 



14. Vernonia stuebelii Hieron., Bot. Jahrb. Syst. 21: 327. 1895. 

 TYPE: Peru: San Martin: Cerro de la Campana between Moyobamba 

 and Rio Huallaga, St'ubel 58b (Holotype B, as photo F! USM). 



Perennial herb or suffrute scent, stems striate, puberulent to glabrate. Leaves cauline; 

 petioles short to indistinct; blades lanceolate, acute or short acuminate at the apex, 

 cuneate at the base, ca. 5-6 cm long, ca. 1.4-1.6 cm wide, margins remotely toothed, 

 slightly scabrous and subrugose above, glabrate beneath. Inflorescences corymbose- 

 paniculate, heads numerous. Heads with 11-16 florets; involucres campanulate, 5- to 

 6-seriate; phyllaries slightly pubescent to glabrate, minutely ciliate at tips, purplish; inner 

 phyllaries lanceolate, tips acute to obtuse or mucronate; outer phyllaries ovate. Corollas 

 4.5-5 mm long. Pappus white; inner bristles ca. 4 mm long, outer pappus almost scalelike, 

 ca. 0.3 mm long. Achenes (immature) pubescent, turbinate. 



Vernonia stuebelii is known only from the type collection from Cerro 

 de la Campana, a remote area of Peru. Its habitat is not known. Flow- 

 ering and fruiting occur in July and August. 



15. Vernonia sambrayana S. B. Jones, sp. nov. TYPE: Peru: Cuzco: 

 La Convencion: upper valley of Rio Sambray; western affluent of Vil- 

 canota, open woods along trail, 1,600 m elevation, Mexia 8055a 

 (Holotype UC!). 



Arbor ca. 7 m alta. Foliorum laminae ovatae vel ovato-lanceolatae, longo- 

 acuminatae, rotundatae vel rotundato-cuneatae versus basim, 12-15 cm longae, 5-6 

 cm latae. Inflorescentia terminalis, obovata. Capitula ca. 20 flosculos habentia. 

 Achenia remote strigosa. 



Small tree ca. 7 m tall, young stems brownish-tomentose, older stems becoming gla- 

 brate. Leaves cauline; petioles canescent, ca. 1.5 cm long; blades ovate to ovate- 




MACBRIDE: FLORA OF PERU 35 



lanceolate, long acuminate at the apex, rounded to rounded-cuneate at the base, 12-15 cm 

 long, 5-6 cm wide, margins entire, revolute, faintly glandular-punctate and lightly pubes- 

 cent above, softly tomentose and with brownish, elevated veins beneath. Inflorescences 

 terminal, obovate, paniculate-corymbiform with branching of a scorpioid-cymose nature, 

 a few foliaceous bracts are present in the inflorescence. Heads with ca. 20 florets, sessile 

 to nearly sessile; involucres broadly campanulate, ca. 3.2 mm long, 4-seriate; phyllaries 

 slightly arachnoid; inner phyllaries ovate to ovate-oblong, tips obtuse to acute; outer 

 phyllaries ovate. Corollas ca. 3 mm long, reddish-purple, faintly glandular. Pappus 

 whitish; inner bristles ca. 2.2 mm long, outer bristles ca. 0.1 mm long. Achenes ca. 1.7 

 mm long, faintly glandular, especially at base, very remotely strigose. 



This species is known only from the type location in Depto. Cuzco. 

 It apparently flowers and fruits in May and June. 



16. Vernonia patens H.B.K., Nov. Gen. & Sp. 4: 41. 1820. TYPE: 

 Humboldt and Bonpland s.n. (Holotype P, as IDC microfiche!). 



V. baccharoides H.B.K., Nov. Gen. & Sp. 4: 40. 1820. TYPE: Colombia: Andium 



Novo-Granatensium juxta Gonzanama et Salto del Fraile, Humboldt and Bonpland 



3438 (Holotype P, as IDC microfiche!). 



V. suaveolens H.B.K., Nov. Gen. & Sp. 4: 38. 1820. TYPE: Novo-Granatensi, Hum- 

 boldt s.n. (Holotype P, as photo F! Isotype B, as photo F!). 

 V. floribunda H.B.K., Nov. Gen & Sp. 4: 38. 1820. TYPE: Peru: Humboldt and 



Bonpland s.n. (Holotype P, as photo F! as IDC microfiche!). 

 V. micradenia DC., Prodr. 5: 38. 1836. TYPE: Poeppig 1215 (Holotype G-DC, as 



photo NY!). 

 V. lanceolaris DC., Prodr. 5: 37. 1836. TYPE: Mexico: Haenke s.n. (Holotype G-DC, 



as microfiche!). 

 V. haenkeana DC., Prodr. 5: 37. 1836. TYPE: Peru: Haenke 8122. (Holotype G-DC, as 



microfiche! as photo F! NY!). 

 V. pacchensis Benth., PI. Hartw. 134. 1844. TYPE: montibus Paccha, Hartweg s.n. 



(Holotype K). 



V. aschenborniana Schauer, Linnaea 19: 714. 1847. 

 Cacalia lanceolaris (DC.) O. Ktze., Rev. Gen. PI. 2: 970. 1891. 

 C. patens (H.B.K.) O. Ktze., Rev. Gen. PI. 2: 970. 1891. 

 C. baccharoides (H.B.K.) O. Ktze., Rev. Gen. PI. 2: 969. 1891. 

 C. suaveolens (H.B.K.) O. Ktze., Rev. Gen. PI. 2: 970. 1891. 

 C. aschenborniana (Schauer) O. Ktze., Rev. Gen. PI. 2: 969. 1891. 

 C. haenkeana (DC.) O. Ktze., Rev. Gen. PI. 2: 970. 1891. 

 Vernonia bangii Rusby, Mem. Torrey Bot. Club 6: 52. 18%. TYPE: Bolivia: between 



Mapiri and Tipuani, Bang 1483 (Holotype NY). 

 V. pacchensis Benth. var. tambillensis Hieron., Bot. Jahrb. Syst. 36: 460. 1905. 



TYPE: Peru: Tambillo, Jelski 699 (Holotype B, as photo F! NY!). 

 V. monsonensis Hieron., Bot. Jahrb. Syst. 40: 335. 1908. TYPE: Peru: Weberbauer 



3489 (Holotype B, as photo F!). 

 V. weberbaueri Hieron., Bot Jahrb. Syst. 40: 354. 1908. TYPE: Peru: Weberbauer 



5023 (Holotype B, as photo F!). 




36 FIELDIANA: BOTANY 



V. salamana Gleason, Bull. Torrey Bot. Club 46: 242. 1919. TYPE: Guatemala: 

 Salama, Maxon and Hay 3385 (Holotype NY). 



Large shrubs or small branched trees, 1.5-7 m tall, stems glabrate to lanate or tomen- 

 tose, younger stems sometimes brownish-lanate. Leaves cauline, slightly coriaceous; 

 petiole 0.7-3 cm long; blades elliptic to broadly elliptic or ovate-lanceolate, acuminate to 

 acute at the apex, attentuate or rounded or truncate at the base, 12-22 cm long (2)3-6(10) 

 cm wide, margins revolute, remotely callus-toothed to serrate, shiny when fresh, surface 

 variable, glabrate to glandular-scabrous above, almost glabrate to hispid or downy, 

 rarely brownish-tomentose beneath. Inflorescences in terminal, much branched panicles 

 or corymbs, the branches sometimes slightly scorpioid. Heads with 14-24 florets, sessile; 

 involucres campanulate, 3.5-5.5 mm long, loosely imbricate, 3- to 6-seriate; phyllaries 

 arachnoid to ciliate, glandular, greenish to reddish-purple; inner phyllaries oblong-ovate 

 to ovate-lanceolate, tips acute; outer phyllaries ovate, tips acute to apiculate. Corollas 

 ca. 5-6.5 mm long, whitish to pinkish, glabrous, sweet-scented. Pappus straw-colored; 

 inner bristles 4-4.8(6) mm long, outer bristles 0.3-0.8 mm long. Achenes 1.5-2 mm long, 

 glandular, hispid, ribbed. Chromosome number: n = 17. 



This species is distributed from Mexico into South America at 

 altitudes of 100 to 2,300 m. In Peru it occurs from Deptos. Amazonas 

 and Loreto south to Cuzco. It is very common in old clearings, along 

 roadsides, and various open places in forests where it is an important 

 part of secondary tropical communities. Flowering and fruiting occur 

 from May to October. Minor variations are common within this wide 

 ranging species; however, it is not possible to separate morphologically 

 the Central and South American material into V. patens and V. bac- 

 charoides. 



LORETO: Boqueron Padre Abad, Woytkowski 34350 (F, MO, UC). 

 AMAZONAS: Chachapoyas: Rio Utcubamba, Hutchison and Wright 

 5854 (F, MO, UC, US, USM). CAJAMARCA: Celendin: Canyon of 

 Rio Marahon above Balsas, Hutchison and Wright 5399 (F, MO, UC, 

 USM). PIURA: Ayabaca: road to Ayabaca, 18 km above Puente Tan- 

 dopa, Hutchison and Wright 6690 (F, UC, US, USM). SAN MARTIN: 

 San Martin: 1-4 km NE Tarapoto, Belshaw 3252 (F, UC, US). 

 HUANUCO: Tingo Maria, Ferreyra 879 (F, UC, US). PASCO: be- 

 tween Oxapampa and LaMerced, R.P.s.n. (USM). JUNIN: Chan- 

 chamayo Valley, Schunke 1586 (F). AYACUCHO: LaMar: between 

 Ayna and Hacienda Luisiana, Dudley 11764 (USM). CUZCO: 

 Machupicchu, Vargas 4557 (F). MADRE DE DIGS: Iberia, Seibert 

 2126 (F). 



17. Vernonia fulta Griseb., Goett. Abh. 24: 164. 1879. TYPE: not 

 seen. 



V. trixioides Rusby, Mem. Torrey Bot. Club 6: 54. 18%. TYPE: Bolivia: Mapiri, 

 Rusby 1484 (Holotype NY, Isotype MO!). 




MACBRIDE: FLORA OF PERU 



37 



4mm 



2cm 

 FIG. 1. Vernonia patens. A, habit; B, head. (From Belshaw 3284, F.) 



V. cotaniensis Hieron., Bot. Jahrb. Syst. 40: 352. 1908. TYPE: Peru: PUNO: Cotani, 

 Weberbauer 1290 (Holotype B, as photo F! NY! USM!). 



Liana 2-4 m, sprawling over other vegetation, stems tomentose to glabrate. Leaves 

 cauline; petioles glabrate to tomentose, 1-2 cm long; blades elliptic, acute to acuminate at 

 the apex, cuneate at the base, 7-18 cm long, 3.5-7 cm wide, margins very remotely callus 

 toothed and slightly revolute, glabrous to glabrate or scabrous above, punctate, and 

 sometimes pilose-hispid beneath. Inflorescences paniculate-cymose. Heads with 22-36 

 florets, stalked; involucres campanulate, 8-11 mm long, 5-seriate; phyllaries arachnoid, 

 loosely appressed, brownish-green with a lighter margin; inner phyllaries oblong- 

 lanceolate, tips acute to slightly apiculate; outer phyllaries lanceolate. Corollas 10 mm 

 long, light reddish-purple, glandular on the lobes. Pappus whitish; inner bristles 7 mm 

 long, outer bristles 0.8 mm long. Achenes 1.8 mm long, strigose, faintly ribbed. Chromo- 

 some number: n = 17. 




38 FIELDIANA: BOTANY 



This species is distributed from Depto. Amazonas in Peru south to 

 Bolivia at elevations of 1,450 to 1,800 m. Flowering and fruiting occur 

 from July to September. In the field, it is a very attractive and striking 

 plant. 



AMAZONAS: Cascadas de Mayasi, Bagua, Wurdack 1830 (US). 

 LORETO: Coronel Portillo: Boqueron del Padre Abad, entre Tingo 

 Maria y Pucallpa, 400-500 m, Ridoutt s.n. (USM). SAN MARTIN: 

 Jepelacio near Moyobamba, Klug 3734 (MO, NY, US). PASCO: Villa 

 Rica, Soukup 4378 (US). JUNIN: Chanchamayo Valley, Schunke 1786 

 (F). CUZCO: Amaytamta, Convencion, Mann 1602 (F). 



18. Vernonia apurimacensis S. B. Jones, sp. nov. TYPE: Peru: 

 Apurimac: 84 miles E of Abancay, Hutchison 1748 (Holotype UC! 

 Isotypes F! NY!). 



Frutex 1 m altus, caulibus albido-canescentibus. Foliorum laminae ca. 2-3.5 cm 

 longae, ca. 1-2.7 cm latae, subtus dense albotomentosae. Inflorescentia composita 

 ex cymis compactis, reductis. Capitula 18 flosculos habentia. Phyllariorum interi- 

 orum apices longoacuminati. Achenia pubescentia. 



Shrub up to 1 m tall, stems whitish, canescent. Leaves relatively small, cauline, some- 

 times crowded; petiolate to almost sessile, petioles to 0.5 cm long; blades cordate to 

 ovate or ovate-elliptic, acute to mucronate at the apex, cordate to rounded or cuneate at 

 the base, 2-3.5 cm long, 1-2.7 cm wide, margins revolute, remotely toothed, rugose and 

 scabrous above and pubescent on large veins above, densely white tomentose beneath. 

 Inflorescences usually of compact reduced cymes, but sometimes with elongated cymes, 

 small bracteal leaves present at base of cymes. Heads with ca. 18 florets, sessile or short 

 peduncled; involucres campanulate, 8-9 mm long, imbricated, 5-seriate: phyllaries wide- 

 spreading when achenes are mature, arachnoid; inner phyllaries oblong-lanceolate, tips 

 long-acuminate; outer phyllaries ovate-lanceolate. Corollas ca. 9 mm long, reddish- 

 purple. Pappus whitish; inner bristles ca. 4.5 mm long, outer bristles ca. 1 mm long. 

 Achenes ca. 1.5 mm long, pubescent. 



This species occurs in Depto. Apurimac and Depto. Cuzco at eleva- 

 tions of 2,200 to 2,700 m in open shrubland. Flowering and fruiting 

 occur from November to February. 



APURIMAC: Andahuaylas: Pincos, Stork and Norton 10668 (F, 

 UC); Rio Pinkos, Weberbauer 5859 (F). CUZCO: Anta: quebrada de 

 Sisal, hasta el puente de Cunyac, hoya del Apurimac, hacia Cuzco, 

 Vargas 412 (F). Puente Cunyac, Ferreyra 2744 (USM). 



19. Vernonia scorpioides (Lam.) Pers., Syn. PI. 2: 404. 1807. 



Conyza scorpioides Lam., Encycl. Meth. 2: 88. 1783-1817. TYPE: Brasil: Commerson 



s.n. (Holotype: P-JU, as IDC microfiche P-JU!). 

 Vernonia subrepanda Pers., Syn. PI. 2: 404. 1807. TYPE: based upon C. scorpioides 



Lam. 




MACBRIDE: FLORA OF PERU 39 



V. lournefortioides H.B.K., Nov. Gen. & Sp. 4: 34-35. 1818. TYPE: Venezuela: 



Caracas, Humboldt s.n. (Holotype: B, as photo B!). 

 Lepidaploa scorpioides (Lam.) Cass., Diet. Sc. Nat. 2: 16. 1823. 

 Staehelina solidaginoides Willd. ex Less., Linnaea 4: 281-282. 1829. TYPE: based 



upon V. tournefortioides H.B.K. 

 Vernonia flavescens Less., Linnaea 6: 657. 1831. TYPE: based upon C. scorpioides 



Lam. 

 V. scorpioides (Lam.) Pers. acentriflora DC., Prodr. 5: 42. 1836. TYPE: Brasil: Bahia, 



April 1831, <f //os/A v s.n. (Holotype: G-DC, as IDC microfiche G-DC!). 

 V. scorpioides (Lam.) Pers. fisubrepanda DC., Prodr. 5: 42. 1836. TYPE: P, not seen. 

 V. scorpioides (Lam.) Pers. y subtomentosa DC., Prodr. 5: 42. 1836. TYPE: Brasil: 



1834, Lund 479 (Holotype: G-DC, as IDC microfiche, G-DC!). 

 V. scorpioides (Lam.) Pers. 8 longifolia DC., Prodr. 5: 42. 1836. TYPE: Brasil: Bahia 



1830, Salzmann s.n. (Holotype: G-DC, as IDC microfiche G-DC!). 

 V. scorpioides (Lam.) Pers. longeracemosa DC., Prodr. 5: 42. 1836. TYPE: Brasil: 



1832, Poeppig 33 (1203) (Holotype: G-DC, as IDC microfiche G-DC!). 

 V. longeracemosa Mart, ex DC., Prodr. 5: 42. 1836. TYPE: not seen. 

 V. languinosa Gardn., J. Bot. 5: 219. 1846. TYPE: Brasil: Minas Geras Prov.: In 



fruticetis propre Formigas in Sertao, Gardner 4764 (Holotype: BM! Isotype: NY!). 

 Cacalia tournefortioides (H.B.K.) O. Ktze., Rev. Gen. PI. 2: 971. 1891. 

 C. scorpioides (Lam.) O. Ktze., Rev. Gen. PI. 2: 971. 1891. 

 V. saepium Ekman., Ark. Bot. 17: 63. 1929. TYPE: Haiti: Depart du Sud: Morne de la 



Hotte ad Ma Blanche, prope Dayette, 7 Aug. 1917, Ekman 463 (Holotype: S! 



Isotypes: F! GH! NY! S! US!). 



Perennial herb to almost shrublike, upright or often scandent and sprawling over other 

 vegetation, stems densely pubescent, strigose to villous, ribbed when dry. Leaves 

 numerous, but not crowded along stem; petioles densely pubescent, 4-7(10) mm long: 

 blades ovate to elliptic, obtuse to acute or acuminate at the apex, cuneate and tapering at 

 the base, (1.8)3.3-15 cm long, (1.2)2.2-8 cm wide, margins entire to rarely denticulate, 

 strigose to pilose-hispid above, hispid to hirsute below. Inflorescences of scorpioid 

 cymes, heads very close together. Heads with 14 to 22 florets, sessile to 1 mm peduncles; 

 involucres campanulate, 3.5-4.5(6) mm long; phyllaries ciliate, pubescent, firmly appress- 

 ed, often purple tinged; inner phyllaries linear-lanceolate with curled tips, 3.8-4.5 mm 

 long; outer phyllaries lanceolate, 2 mm long. Corollas 6-7(8) mm long, reddish-purple, 

 lobes ciliate. Pappus white; inner bristles 4. 5-6 mm long, outer scales fimbriate, (0.8)1-1.6 

 mm long. Achenes 1.5 mm long, with sparse, stiff hairs between ribs. Chromosome 

 number: n = 17. 



This species is distributed from Central America and the West Indies 

 southward into Argentina. In Peru it occurs in the selva east of the 

 Andes. Vernonia scorpioides is a widely distributed species, rather 

 weedy, and commonly occurs on disturbed sites. Flowering and fruit- 

 ing occur throughout the year. 



LORETO: Prov. Coronel Portillo, Ferreyra 18029 (MO). LA 

 LIBERTAD: Otuzco: Huaranchal, Otuzco, Miranda 1334 (USM). 

 AMAZONAS: Prov. Bongara, Hutchison and Wright 6829 (F, MO, 




FIG. 2. Vernonia scorpioides. A, habit; B, head. (From Hutchison & Wright 3848, F.) 



40 




MACBRIDE: FLORA OF PERU 41 



UC, US). CAJAMARCA: Prov. Hualgayoc, Monte Seco, Soukup 3821 

 (US). SAN MARTIN: Jepelacio, near Moyobamba, Klug 3297 (F, MO, 

 NY, US). HUANUCO: Prov. Huanuco, Tingo Maria, Asplund 12962 

 (US). PASCO: Tarma: San Luis de Shuaro, Ferreyra 18608 (USM). 

 AYACUCHO: La Mar: between Ayna and Hacienda Luisiana, Dudley 

 11661 (USM). JUNIN: Pichis Trail, Enehas, Killip and Smith 25782 (F, 

 US). CUZCO: Macchu-Picchu, Ferreyra 2700 (MO, US). 



20. Vernonia brachiata Benth. ex Oerst., Vidensk. Meddel. Dansk 

 Naturhist. Foren. Kjobenhavn 1852: 67. 1852. TYPE: Oersted s.n. (K). 



Cacalia brachiata (Benth.) O. Ktze., Rev. Gen. PI. 2: 969. 1891. 



Vernonia megaphylla Hieron., Verh. Bot. Vereins Prov. Brandenburg 48: 195. 1906. 



TYPE: Peru: Loreto: Pongo de Cainarachi, Ule 6386 (B, as photo F! NY!). 

 V. digitata Rusby, Bull. New York Bot. Gard. 8: 125. 1912. TYPE: Bolivia: Mapiri, 



Williams 713 (NY, not seen). 



Coarse suffrute scent, perennial herb, 1.5-6 m tall, stems glabrate to lanate. Leaves 

 cauline on current season's growth; petiole ca. 1 cm long; blades elliptic, coriaceous, 

 acuminate at the apex, cuneate to auriculate at the base, 20-70 cm long, 8-19 cm wide, 

 margins entire to remotely toothed, glabrous above, remotely glandular to sparsely 

 pubescent beneath. Inflorescences terminal, large, branches of scorpioid cymes with 

 numerous, relatively small heads. Heads with 24-34 florets, sessile in 1 or 2 rows along 

 branches of inflorescence; involucres campanulate. 3-6 mm long, loosely imbricate, 4- to 

 5-seriate; phyllaries lanate-glandular, loosely appressed, reddish-purple; inner phyllaries 

 long-lanceolate, tips acuminate; outer phyllaries ovate-lanceolate. Corollas ca. 6.5 mm 

 long, reddish-purple, lobes glandular. Pappus white; inner bristles ca. 5 mm long, outer 

 bristles ca. 0.7 mm long. Achenes ca. 2.8 mm long, sparsely pubescent, ribbed. Chromo- 

 some number: n = ca. 17. 



It is found from Costa Rica and Panama southward into Peru. This 

 species is distributed in Peru from Depto. Amazonas south to Depto. 

 Cuzco in the selva. It occurs at the edge of woods, along streams, and 

 in tropical woodlands at elevations of 135 to 1,000 m. Flowering and 

 fruiting occur from June to November. 



AMAZONAS: Bagua, Aramango, Sagastegui s.n. (GA). SAN 

 MARTIN: Lamas, Belshaw 3423 (F, MO). HUANUCO: Tingo Maria, 

 Ferreyra 10293 (MO). LORETO: Previsto, Woytkowski 7585 (US). 

 JUNIN: Tarma: Puente Perene, Ferreyra 11349 (USM). CUZCO: In- 

 ambari, Vargas 16514 (US). 



21. Vernonia cainarachiensis Hieron., Verh. Bot. Vereins Prov. 

 Brandenburg 48: 196. 1906. TYPE: Peru: Loreto: Pongo de Cainarachi, 

 Ule 6387 (Holotype B, as photo F! Isotype F!). 



Herbaceous to suffrutescent shrub, 2.5-5 m tall, often sprawling over other vegetation, 

 stems brownish, pilose-hispid to glabrate. Leaves cauline; petioles (0)1.5-3.5(4) cm long; 

 blades elliptic -obovate, long-acuminate at the apex, cuneate at the base, 14-40 cm long, 




1mm 



2mm 



FIG. 3. Vernonia brachiata. A, habit; B, head; C, achene. (From Liesner 221, MO.) 



42 




MACBRIDE: FLORA OF PERU 43 



5-14 cm wide, margins slightly re volute, very remotely callus toothed, glabrous above, 

 almost glabrate or with small closely appressed hairs and strigose on veins beneath. 

 Inflorescences terminal, paniculate-scorpioid-cymose. Heads with 7-13 florets, sessile; 

 involucres campanulate, 5.5-6.9 mm long, ca. 4-seriate; phyllaries ciliate to arachnoid, 

 and sometimes finely strigose, appressed, reddish-purple; inner phyllaries oblong, tips 

 obtuse; outer phyllaries oblong and obtuse. Corollas 10 mm long, reddish-purple, glan- 

 dular. Pappus whitish to pinkish; inner bristles ca. 6 mm long, outer bristles ca. 1 mm 

 long. Achenes ca. 3 mm long, short pilose, ribbed. Chromosome number: n = 17. 



This species is distributed in Peru from Depto. Lore to south to 

 Depto. Cuzco at 400 to 1,600 m elevation in open forest or brushland. 

 Flowering and fruiting occur from July to October. 



AMAZONAS: Huampami, Kayap 1413 (MO). LORETO: Coronel 

 Portillo, Ferreyra 18048 (MO). SAN MARTIN: Puerto Pizana, Rio 

 Huallaga, Schunke 6453 (MO). HUANUCO: Tingo Maria, Ferreyra 

 2281 (US). JUNIN: San Ramon, Schunke A-l (F, NY, US). CUZCO: 

 Paucartambo, Mann 1716 (F). 



22. Vernonia yurimaguasensis Hieron., Verh. Bot. Vereins Prov. 

 Brandenburg 48: 195. 1907. TYPE: Peru: Loreto: Yurimaguas, Vie 

 6270 (Holotype B, as photo F! Isotype F!). 



V. vargasii Cuatrec., Bot. Jahrb. Syst. 77: 83. 1956. TYPE: Peru: Cuzco: Urubamba, 

 Machupicchu, Vargas 6236 (Holotype F!). 



Scandent shrub, stems grayish-brown, velutinous. Leaves cauline; petiole up to 1 cm 

 long; blades broadly elliptic, acute to acuminate at the apex, cuneate at the base, 11-16 

 cm long, 4-8 cm wide, margins slightly revolute, sparsely pubescent to glabrate above, 

 sparsely pubescent and glandular-punctate beneath. Inflorescences terminal, scorpioid 

 cymes, branches divaricate. Heads with ca. 20 florets; involucres campanulate, ca. 5 mm 

 long; phyllaries pubescent, loosely imbricated, grayish; inner phyllaries oblong, tips 

 acute; outer phyllaries ovate. Corollas ca. 4 mm long, violet. Pappus whitish; inner 

 bristles ca. 4 mm long, outer bristles ca. 0.6 mm long. Achenes ca. 2.2 mm long, glandu- 

 lar, brownish. 



This species occurs in Deptos. Loreto, Amazonas and Cuzco in the 

 tropical selva. Flowering and fruiting occur from May to August. 



AMAZONAS: Bagua, Sagastegui, Lopez and Collantes 4248 (GA). 

 LORETO: as type. CUZCO: Urubamba: Machupicchu, Vargas 6236 



(F). 



23. Vernonia myriocephala DC., Prodr. 5: 40. 1836. TYPE: Peru: 

 Haenke s.n. (Holotype G-DC, as IDC microfiche! Isotype F! NY!). 



Cacalia myriocephala (DC.) O. Ktze, Rev. Gen. PI. 2: 970. 1891. 



Shrub, 1.5-6 m tall, branches erect, stems strigose to glabrate. Leaves cauline, firm; 

 petioles 0.2-1.3 cm long; blades elliptic to ovate or lanceolate, acuminate at the apex, 

 cuneate and slightly decurrent at the base, 10-17 cm long, 3.5-7 cm wide, margins rev- 

 olute (sometimes only slightly so) and remotely toothed, sometimes sparsely glandular or 




44 FIELDIANA: BOTANY 



faintly scabrous, glabrate, pubescent on large veins above, glandular and minutely 

 pubescent (best viewed by turning leaf at oblique angle) beneath. Inflorescences scorpioid 

 with a few very small bracteal leaves scattered in the inflorescence. Heads with 17-22 

 florets, sessile to short-stalked; involucres campanulate, (6)6.5-6.8(8) mm long, im- 

 bricate, ca. 5-seriate; phyllaries arachnoid, and sometimes minutely strigose, somewhat 

 loosely arranged, reddish-purple to greenish; inner phyllaries oblong-lanceolate, tips 

 acute rarely acute-acuminate or fimbriate; outer phyllaries tips acuminate to long- 

 acuminate. Corollas ca. 7 mm long, reddish-purple fading to whitish, glabrous. Pappus 

 white; inner bristles ca. 6 mm long, outer scales ca 1.1 mm long. Achenes ca. 2.5 mm 

 long, strigose, ribbed. 



This species is distributed from Depto. San Martin south to Depto. 

 Cuzco at an elevation of 200 to 1,000 m. It occurs in tropical forests, 

 sunny clearings, and brushlands. Vernonia myriocephala appears 

 closely related to V. canescens, differing in amount of pubescence on 

 the lower surfaces of the leaves. The former has leaf blades minutely or 

 sparsely pubescent beneath, whereas V. canescens is densely to 

 sparsely strigose to strigose hirsute beneath. Flowering and fruiting 

 occur from June to August. 



LORETO: Alto Amazonas: Yurimaguas, Ferreyra 4979 (USM). 

 SAN MARTIN: Mishquiyacu, Ferreyra 4622 (MO, USM). 

 HUANUCO: Tulumayo near Tingo Maria, Ferreyra 2168 (US, USM). 

 JUNIN: Satipo, Ridoutt 11718 (MO, USM). CUZCO: Quispicanchis, 

 Vargas 16495 (F). 



24. Vernonia canescens H.B.K., Nov. Gen. & Sp. 4: 35, tab. 317. 

 1820. TYPE: Peru: Guancabamba, Bonpland 3529 (Holotype P, as IDC 

 microfiche! as photo F!). 



V. mollis H.B.K., Nov. Gen. & Sp. 4: 36. 1820. TYPE: (Holotype P, as IDC 



microfiche!). 



Lepidaploa canescens (H.B.K.) Cass., Diet. Sc. Nat. 26: 18. 1823. 

 Vernonia bullata Benth. ex. Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. 



Kjobenhavn 1852: 67. 1852. TYPE: Costa Rica: Cartago, Bjergene s.n. (K). 

 V. arborescens Sw. var. cuneifolia Britt., Bull. Torrey Bot. Club 18: 311. 1891. TYPE: 



Bolivia: Reis, Rusby 2148 (Holotype NY!). 



Cacalia canescens (H.B.K.) O. Ktze., Rev. Gen. PI. 2: 969. 1891. 

 C. mollis (H.B.K.) O. Ktze., Rev. Gen. PI. 2: 970. 1891. 

 C. bullata (Benth. ex Oerst) O. Ktze., Rev. Gen. PI. 2: 969. 1891. 

 Vernonia volubilis Hieron., Bot. Jahrb. Syst. 36: 460. 1905. TYPE: Peru, Tambillo, 



Jelskii 775 (Holotype B, as photo NY!). 

 V. patuliflora Rusby, Bull. New York Bot. Gard. 4: 376. 1907. TYPE: Bolivia: 



Coroico, Yungas, Bang 2396 (Holotype NY! Isotype NY!). 

 V. cuneifolia (Britt.) Gleason, Amer. J. Bot. 10: 301. 1923. (non V. cuneifolia Gardn., 



Hooker's J. Bot. Kew Gard. Misc. 5: 215. 1846). 




MACBRIDE: FLORA OF PERU 45 



V. rusbyi Gleason, Amer. J. Bot. 2: 753. 1932. (based upon V. arborescens Sw. var. 



cuneifolia Britt.). 

 V. pseudomollis Gleason, Amer. J. Bot. 10: 307. 1932. TYPE: Bolivia: Yungas, Rusby 



1658 (Holotype NY! Isotype NY!). 



Semi-woody perennial herbs, to sprawling shrubs, to 3 m tall, often much branched, 

 stems densely pubescent above especially near the inflorescence. Leaves cauline, usually 

 not crowded; petioles 4-14 mm long; blades broadly lanceolate to ovate-elliptic, acumi- 

 nate to acute at the apex, rounded to cuneate at the base, 9-20 cm long, 4-7 cm wide, 

 margins almost entire to remotely toothed, sometimes revolute, scabrous to sparsely 

 strigillose above, densely to sparsely strigillose, and sometimes glandular beneath. In- 

 florescences of terminal, scorpioid-cymes arranged into spreading panicles or corymbs. 

 Heads with 18-24 florets, sessile; involucres campanulate, 5-6 mm long, loosely im- 

 bricated; phyllaries tomentose; inner phyllaries linear-lanceolate, tips acute to slightly 

 spinulose tipped; middle phyllaries spinulose tipped, outer phyllaries lanceolate 

 spinulose tipped. Corollas ca. 5 mm long, pinkish fading to white. Pappus white; inner 

 bristles ca. 4 mm long, outer bristles ca. 0.8 mm long, sometimes slightly flattened. 

 Achenes ca. 2.5 mm long, densely strigillose. faintly ribbed. Chromosome number: 

 n = 17. 



This species is distributed from Mexico and Central America south- 

 ward into Bolivia. It occurs in tropical vegetation often in secondary 

 scrub. Flowering and fruiting occur from July to December. 



LORETO: Ucayali, Contamana, McDaniel 14091 (F, MO, MISSA). 

 PIURA: Huancabamba, Palambla, Sagastegui, Cabanillas, Dios 8139 

 (MO) AMAZONAS: Bongara, Rio Utcubamba, Hutchison and Wright 

 5865 (MO, UC, US). SAN MARTIN: Lamas, Ferreyra 17285 (MO). 

 HUANUCO: Cayumba entre Huanuco y Tingo Maria, Ferreyra 4196 

 (MO). JUNIN: Colonia Perene, Killip and Smith 24974 (F, NY). 

 CUZCO: Machupicchu, Vargas 19902 (US). 



25. Vernonia fieldiana Gleason, Bull. Torrey Bot. Club 59: 374. 1932. 

 TYPE: Peru: San Martin: San Roque, Williams 7663 (Holotype US, 

 Isotype F!). 



Shrub, ca. 1 m tall, upper stems slender, densely and closely cinereous-tomentose. 

 Leaves firm, dull green; petioles stout, ca. 1 mm long; blades ovate-oblong to elliptic- 

 ovate, sharply acute or subacuminate at the apex, rounded-cuneate at the base, ca. 3.5 

 cm long, ca. 1.9 cm wide, margins mostly entire, but slightly revolute and very remotely 

 toothed, both sides inconspicuously pubescent with minute slender hairs; lateral veins 

 curved, ascending, and parallel, strongly elevated beneath. Inflorescences somewhat 

 crowded, many-flowered, compound scorpioid cymose-paniculate, its branches densely 

 and softly cinereous-tomentose. Heads with ca. 11 florets, sessile; involucres broadly 

 campanulate, 4-5 mm long, ca. 4-seriate; phyllaries densely subtomentose, loosely im- 

 bricate; inner phyllaries oblong-lanceolate, tips sharply acute; outer phyllaries 

 triangular-lanceolate. Corollas ca. 5 mm long, reddish-purple. Pappus whitish; inner bris- 

 tles ca. 4.5 mm long, outer scales ca. 1 mm long. Achenes ca. 1 mm long, densely 

 sericeous. 




46 FIELDIANA: BOTANY 



This species is known only from Depto. San Martin in Peru. It has 

 been collected in mountain forests at elevations of 1,200 to 1,600 m. 

 Flowering and fruiting occur from December to February. 



SAN MARTIN: Jepelacio near Moyobamba, Klug 3423 (F, MO, 

 NY, US). 



26. Vernonia salzmannii DC., Prodr. 5: 55. 1836. TYPE: Brasil: 

 Salzmann 1830 (Holotype G-DC, as IDC microfiche!). 



V. poeppigiana DC., Prodr. 5: 55. 1836. TYPE: Peru: Poeppig 1204 (Holotype G-DC. 



as IDC microfiche!) non. V. poeppigiana DC., Prodr. 5: 20. 1836. 

 V. argyropappa Buck. Index Prodr. I:IX Tom. V. (based upon V. poeppigiana DC.. 



Prodr. 5: 55. 1836). 

 V. geminiflora Poepp., Poepp. & Endl. Nov. Gen. & Sp. 3: 42. 1845. (based upon V. 



poeppigiana DC., Prodr. 5: 55. 1836). 



Cacalia argyropappa (Buck) O. Ktze., Rev. Gen. PI. 2: 969. 1891. 

 C. salzmannii (DC.) O. Ktze., Rev. Gen. PI. 2: 971. 1891. 



Herb 1-2 m tall, branched, stems sparsely hirsute-pubescent with brownish hairs. 

 Leaves thin but firm: almost sessile; blades ovate-lanceolate to oblong-lanceolate, acute 

 or acuminate at the apex, gradually narrowed or obtuse at the base, 6-12 cm long, 1.5-3 

 cm wide, margins entire or minutely serrulate, rugose and papillate-pilose above, softly 

 strigose-hirsute and resinous beneath. Inflorescences sparingly branched, of several di- 

 varicately spreading scorpioid cymes each bearing 4-10 heads, bracteal leaves oblong to 

 oblong-lanceolate. Heads with 21-34 florets, 1-3 cm apart; involucres campanulate or 

 nearly hemispheric, 8-10 mm long; phyllaries sparsely pilose, erect, linear, tips narrowed 

 to a subulate, spinose tip. Corollas 5-6 mm long, reddish-purple. Pappus white; inner 

 bristles 6-8 mm long, outer scales ca. 1 mm long. Achenes ca. 3 mm long, hirsute. 



This species is distributed from southern Mexico south through 

 Central America and northern South America into Brazil. In Peru, it 

 occurs in the tropical selva in old clearings or secondary growth from 

 Depto. San Martin to Depto. Madre de Dios. Flowering and fruiting 

 occur from May to August. 



SAN MARTIN: San Martin, Ferreyra 17401 (MO, USM). 

 HUANUCO: Huanuco: Concordia, cerca a Puente Durand, Ferreyra 

 9327 (USM). JUNIN: Tarma: arriba de San Ramon, Ferreyra 16321 

 (USM). CUZCO: Convencion: Vargas 13170 (US). MADRE DE 

 DIOS: Iberia, vie. Rio Thuamanu, Seibert 2125 (MO, F, US). 



27. Vernonia herbacea (Veil.) Rusby, Mem. Torrey Bot. Club 4: 209. 

 1895. 



Chrysocoma herbacea Veil., Fl. Flum. 330. 1825. TYPE: as illustration, Atlas Tab. 29. 



T 8. 1835. 



Vernonia obovata Less., Linnaea4: 279. 1829. TYPE: Brasil: Sellow s.n. (not seen). 

 V. paucifolia Rusby, Mem. Torrey Bot. Club 3: 50. 1893. TYPE: Bolivia: Yungas, 



Bang 247 (NY). 




MACBRIDE: FLORA OF PERU 47 



Perennial herb, stems villous to hirsute, pubescence straw-colored. Leaves cauline; 

 sessile; blades obovate to obovate-lanceolate, obtuse at the apex, cuneate at the base, 6-7 

 cm long, 2.5-3.5 cm wide, margins slightly revolute, pubescent, remotely hirsute above, 

 upper surface of leaf dark brown when dry, villous to hirsute with straw-colored pubes- 

 cence beneath. Inflorescences terminal, condensed scorpioid-cymose to corymbose. 

 Heads with 12-13 florets, short pedunculate; involucres broadly campanulate, ca. 7 mm 

 long, 3- to 4-seriate; phyllaries strigose, loosely appressed; inner phyllaries lanceolate- 

 oblong, tips acute to acuminate or slightly aristate; outer phyllaries tips lanceolate. 

 Corollas ca. 8 mm long, dark reddish-purple, sparsely glandular. Pappus white; inner 

 bristles ca. 7 mm long, outer bristles ca. 1 mm long. Achenes ca. 2 mm long, strigose. 



This species is distributed from southern Brazil northward into Peru. 

 It is apparently uncommon in Peru since only one collection has been 

 seen. Flowering and fruiting occur from May to June. 



JUNIN: Chanchamayo Valley, Schunke 1527 (F). 



II. PIPTOCARPHA 6 



Piptocarpha R. Br., Trans. Linn. Soc. London 12: 121. 1817. (1818). 

 TYPE: P. brasiliana Cass., Diet. Sc. Nat. 41: 109. 1826. 



Carphobolus Schott, Spreng. Syst. iv. Cur. Post. 409. 1827. TYPE: C. sessiliflorus 



Schott. 

 Monanthemum Griseb., Fl. Brit. W. Ind. 354. 1861. TYPE: M. cruegerii Griseb. 



Shrubs, usually scandent or infrequently trees, especially in some Brazilian species; 

 branches pubescent, stellate-tomentulose or lepidote. Leaves alternate, petiolate, blades 

 large, ovate to lanceolate, entire to subentire, pinnately veined with prominent ascending 

 lateral veins, arching and anastomosing near the margins, glabrous above, often to- 

 mentose with stellate trichomes or lepidote beneath, bases usually oblique. Inflorescences 

 aggregated at base of leaves (often reduced toward apex of stem and on secondary 

 branches) in axillary corymbs, umbels or sessile to subsessile in axillary clusters or in 

 axillary and terminal panicles. Heads with 1-35 florets; involucre campanulate, 

 cylindrical-campanulate to turbinate; phyllaries imbricated in several series, the 

 outer bracts persistent, small, triangular-ovate, apex obtuse, tomentulose, upper margin 

 ciliate to fimbriate; inner bracts narrowly ovate to oblong to lanceolate, apex tomen- 

 tulose to glabrous, acute, often with a dark tip, curling and usually deciduous with 

 achenes; receptacle convex, flowers in species with turbinate or broadly campanulate 

 involucres subtended by distinct, linear-lanceolate, scarious paleas with acuminate tips, 

 deciduous with achenes; in species with cylindrical to narrowly campanulate involucres 

 (usually with 6 or less florets), paleas absent. Corollas regular, narrowly tubular, 5-lobed; 

 stamens 5, anthers apically acute, bases sagittate with auricles acute to caudate; style 

 branches slender, acute, stigma bifid, the stigmatic surface hispid. Pappus biseriate, the 

 inner series of long, filiform, equal bristles, the outer series of shorter, filiform, unequal 

 bristles or paleaceous scales, often inconspicuous or absent in some species. Achenes 

 glabrous or infrequently pilose, cylindrical or angled, often 10-costate, apex truncate. 



6 Mr. Gerald L. Smith, a pre-doctoral student, is presently revising Piptocarpha and 

 generously contributed to this treatment. His assistance is gratefully acknowledged. 




48 FIELDIANA: BOTANY 



Piptocarpha is a small neotropical genus of ca. 40 species, extending 

 southward from the West Indies and Central America into northern 

 and central South America. 



REFERENCE 



ELIAS, THOMAS S. 1975. Fl. of Panama. Part IX. Compositae. Ann. Missouri Bot. Gard. 

 62(4): 860. 



KEY TO SPECIES OF Piptocarpha 

 a. Inflorescences of 20-60 heads. 



b. Inflorescences of terminal and axillary panicles of heads clustered at the ends of 



branchlets 7. P. gutierrezii. 



bb. Inflorescences of axillary, branching corymbose clusters of heads. 



c. Heads with 3 florets; involucre campanulate; leaves basally cuneate 



5. P. sprucei. 



cc. Heads with 6 florets; involucre ovoid-campanulate; leaves basally oblique, 

 d. Lower leaf surface densely appressed, cinereous-tomentose with stellate 



trichomes; phyllaries sparsely tomentose at apex \. P. poeppigiana. 



dd. Lower leaf surface thinly gray-pubescent with stellate trichomes; phyl- 

 laries densely tomentose at apex 4. P. canescens. 



aa. Inflorescences of 4-16 heads. 



e. Inflorescences of axillary, stoutly pedunculate clusters of heads; involucre cam- 

 panulate 6. P. lechleri. 



ee. Inflorescences of axillary clusters of subsessile heads; involucre turbinate. 



f. Lower leaf surface yellow-gray tomentose with stellate trichomes; branches 



cinnamon-tomentose with stellate trichomes 2. P. asterotrichia. 



ff. Lower leaf surface cinereous to yellow-brown tomentose with stellate 

 trichomes; branches cinereous-tomentulose or lepidote 3. P. opaca. 



1 . Piptocarpha poeppigiana (DC.) Baker in Mart. , Fl. Bras. 6(2): 131. 

 1873. 



Vernonia poeppigiana DC., Prodr. 5:20. 1836. TYPE: Peru; Poeppig 1425 (Holotype: 



G-DC, as microfiche G-DC!, photo NY! Isotypes: G! P! BM! B as photos GH! NY!). 

 Vernonia tereticaulis DC., Prodr. 5:20. 1836. TYPE: Peru: Haenke s.n. (Holotype: 



PR! Isotypes: G-DC, as microfiche G-DC! P! F!). 

 Carphobolus poeppigiana (DC.) Sch. Bip., Pollichia 20/21:422. 1863. 

 Carphobolus tereticaulis (DC.) Sch. Bip., Pollichia 20/21: 422. 1863. 

 Piptocarpha tereticaulis (DC.) Baker in Mart., Fl. Bras 6(2): 131. 1873. 

 Piptocarpha chontalensis Baker in Mart., Fl. Bras. 6(2): 132. 1873. TYPE: Nicaragua: 



Tate 163 (Lectotype: K! [selected from among syntypes] Isolectotype: BM! Syn- 



types: BM!). 

 Piptocarpha costaricensis Klatt, Bull. Soc. Bot. Belg. 31(1): 184. 1892. TYPE: Costa 



Rica: Pittier 4927 (Lectotype: GH! [selected from among syntypes] Isolectotypes: 



GH! M! BR!). 




MACBRIDE: FLORA OF PERU 49 



Piptocarpha laxa Rusby, Bull. New York Bot. Card. 8(28): 123. 1912. TYPE: Bolivia: 



Charopampa, Williams 703 (Holotype: NY! Isotypes: K! BM! US!). 

 Piptocarpha foliosa Cuatrec., Brittonia 8: 161. 1955. TYPE: Colombia: Amazonas, 



Tacana, Castanal igapo, Schultes & Black 46-82 (Holotype: US!). 

 Piptocarpha paraensis Cabrera, Arquiv. Jard. Bot. Rio de Janeiro 15: 73. 1957. TYPE: 



Brazil: Para: Rio Tapajoz, Pimentel, Ducke 21-VIII-1923 (Holotype: LP! Isotypes: 



RB!). 



Subscandent shrub to drooping liana with branches 3-30 m long; stems cinereous to 

 yellow-brown tomentulose-lepidote. Leaves cauline, not crowded; petioles sulcate, 1-2.5 

 cm long; blades coriaceous, elliptic to oblong-ovate to broadly ovate, acute to acuminate 

 at the apex, oblique at the base, 7-20 cm long, 4-10 cm broad, margin revolute, sometimes 

 faintly toothed; glabrous above except tomentulose on midvein, densely cinereous to 

 yellow-brown stellate-tomentose, occasionally glandular beneath, lateral veins 6-8 pairs. 

 Inflorescences in dense axillary corymbose clusters of 20-60 heads. Heads usually with 6 

 florets, sessile or shortly pedicellate in groups of 2 or 3 at the ends of tomentose pedun- 

 cles; involucre ovoid to narrowly campanulate, 5-7 mm long, 3-4 mm broad, 4-seriate; 

 phyllaries tomentulose at tips, margins ciliate to fimbriate; outer phyllaries triangular- 

 ovate, acute, persistent; inner phyllaries oblong to lanceolate, acute, deciduous. Corollas 

 4-5 mm long, glabrous, the lobes 1.5-3 mm long, revolute, white, fragrant; anthers 3 mm 

 long, basal auricles caudate, 0.4 mm long. Pappus white, biseriate, the inner series of 

 equal, filiform bristles 5-6.5 mm long, the outer series of short, inconspicuous bristles 

 0.5-1 mm long. Achenes 2.5-3 mm long, 10-costate, glabrous or sparsely pilose. 



This species is distributed throughout tropical regions from southern 

 Mexico to central Bolivia. It is most frequently found along rivers in 

 tropical forests at elevations of 250-1,000 m. Flowering and fruiting 

 occur mainly from July to November but occasionally the year round. 



LORETO: Previsto, Woytkowski 7566 (MO, UC, US). 

 AMAZONAS: Bagua, Rio Santiago, Wurdack 2501 (F, UC, USM). 

 SAN MARTIN: Lamas, Belshaw 3445 (F, MO, UC). HUANUCO: 

 Tingo Maria, Ferreyra 891 (US, USM). JUNIN: Mazamar., Woyt- 

 kowski 5966 (MO, US). CUZCO; Paucartambo, Marin 1697 (F). 



2. Piptocarpha asterotrichia (Poepp. & Endl.) Baker in Mart., Fl. 

 Bras. 6(2): 127. 1873. 



Vernonia asterotrichia Poepp. & Endl., Nov. Gen. & Sp. 3:41-42 t. 247. 1843. TYPE: 



Peru: Poeppig 1887 (Holotype: W! Isotypes: P! NY! GH!). 



Carphobolus asterotrichus (Poepp. & Endl.) Sch. Bip., Pollichia 20/21: 426. 1863. 

 Piptocarpha insignis Gleason, Bull. Torrey Bot. Club 59: 371-372. 1932. TYPE: Peru: 



Junin, San Nicolas, Pichis Trail, Killip & Smith 26083 ( Holotype: NY! Isotype: US!). 



A high-climbing, wide-spreading, much-branched liana with branches reaching 3-15 m, 

 often in trees, showy; stems densely cinnamon-stellate tomentose. Leaves cauline, not 

 crowded; petioles sulcate, densely yellow-gray stellate-tomentose, 0.8-1.7(2.5) cm long; 

 blades ovate-oblong to lanceolate, acuminate at the apex, oblique or rounded to sub- 

 cordate at the base, 6-19 cm long, 3-9 cm wide, margins slightly revolute and very faintly 

 remotely toothed, glabrous, opaque, glandular above except yellow-gray stellate- 

 tomentose on major veins, prominently reticulately veined when dry, 5-8 pairs of lateral 




2mm 



FIG. 4. Piptocarpha asterotrichia. A, habit; B, achene; C, anther. (From Schnnke 

 263. F.) 



50 




MACBRIDE: FLORA OF PERU 51 



veins, densely yellow-gray stellate-tomentose beneath. Inflorescences in dense, rounded, 

 axillary, clusters of 4-10 subsessile heads, number of heads in glomerules reduced toward 

 apex of stem (seemingly forming flat "sprays" in the fresh condition). Heads with 11-35 

 florets, subsessile; involucre broadly turbinate, 10-17 mm long, 4-10 mm wide, closely 

 imbricate in 5-6 series, when fresh the involucre is brilliant yellow-green; phyllaries 

 ciliate on the margins, rarely pubescent; outer phyllaries persistent, ovate, apex acute 

 with a dark tip; inner phyllaries deciduous, lanceolate, apex acute; flowers subtended by 

 linear-lanceolate paleas, tips acuminate, deciduous with achenes. Corollas 6-7.5 mm 

 long, creamy white, fragrant, sometimes glandular, lobes revolute, 2-3 mm long; anthers 

 ca. 4 mm long, basal auricles sharply acute, 0.2 mm long. Pappus white, predominately 

 uniseriate, inner bristles equal, 6-8 mm long, outer bristles less than 1 mm long, totally 

 absent in some specimens. Achenes 3.5-4.5 mm long, angled, indistinctly costate, gla- 

 brous. 



This species is distributed from Colombia south to the Cordillera 

 Real in eastern Bolivia. It occurs at elevations of 425-1,500 m in tropi- 

 cal forests, montane rain forests, at lower edges of cloud forests, and in 

 secondary growth. Flowering and fruiting occur mainly from June to 

 December but occasionally throughout the year. 



Piptocarpha insignis Gleason is considered to be a very robust form 

 of P. asterotrichia. It is known only from the type collection and differs 

 only from typical P. asterotrichia specimens by its larger heads. 



AMAZONAS: Quebrada Aintami, Kayap 690 (GA, MO). SAN 

 MARTIN: Lamas, Belshaw 3434 (F, MO, UC). JUNIN: San Ramon, 

 Killip & Smith 24747 (F, US). CUZCO: Convencion, Dudley 10282 

 (MO). LORETO: Maynas, Tocache, Poeppig 1887 (W, P, NY, GH). 



3. Piptocarpha opaca (Benth.) Baker in Mart., Fl. Bras. 6(2): 124. 

 1873. 



Vernonia opaca Benth., in Hooker Lond. J. Hot. 2: 39. 1840. TYPE: Guyana: Serra 



Mey, Schomburgk 1016 (Holotype: K! Isotypes: K! BM!). 

 Carphobolus latifolius Sch. Bip., Pollichia 20/21: 426. 1863. TYPE: Brasil: Para: in 



vicinity of Obidos, Spruce Dec. 1849 (Holotype: K! Isotypes: BM! M! NY! GH!). 

 Piptocarpha opaca (Benth.) Baker var. latifolia (Sch. Bip.) Baker in Mart., Fl. Bras. 



6(2): 124. 1873. 



A highly scandent, much-branched liana with branches 3-12 m long, often in trees, 

 stems cinereous, appressed tomentulose-lepidote. Leaves cauline, not crowded; petioles 

 sulcate, cinereous tomentose-lepidote, 1-2 cm long; blades coriaceous, elliptic to ovate- 

 oblong to broadly ovate, acuminate at the apex, oblique at the base, 7-13 cm long, 3-5 cm 

 wide, margins slightly revolute and remotely toothed, glabrous and somewhat lustrous 

 above, 6-8 pairs of lateral veins, densely stellate-tomentose and yellow-brown lepidote 

 beneath. Inflorescences in dense axillary, hemispheric to rounded clusters of 6-15 heads. 

 Heads with 9-12 florets, subsessile; involucre turbinate, 7.5-11 mm long, 3.5-5 mm 

 wide, imbricate in 5-6 series; phyllaries darkened and tomentulose at tips, especially in 

 outer phyllaries, margins ciliate to fimbriate; outer phyllaries persistent, ovate, apex 




52 FIELDIANA: BOTANY 



acute to obtuse; inner phyllaries deciduous, narrowly ovate to oblong, apex acute: 

 flowers subtended by linear-lanceolate paleas, tips acuminate, deciduous with achenes. 

 Corollas 4-5 mm long, purple-white, very fragrant, glabrate, lobes revolute, 1.5-2.5 mm 

 long: anthers 3-4 mm long, basal auricles caudate, 0.4 mm long. Pappus white, biseriate, 

 the inner series of bristles equal, filiform, 5-6 mm long, the outer series poorly developed, 

 bristles short, unequal, 0.5-1 mm long. Achenes 2.5-3 mm long, cylindrical to 3-angled, 

 10-costate, glabrous or sparsely pilose. 



This species is distributed throughout the range of the Amazon 

 River. It occurs most frequently in secondary woods with sandy soil 

 along the margins and upland regions of the Amazon River at eleva- 

 tions of 25-700 m. 



LORETO: tributary of Rio Nanay near Iquitos, McDaniel 10749 

 (GA). LORETO: Iquitos, Vandemann 2260 (K). AMAZONAS: Rio 

 Santiago-Rio Pongo de Manseriche, Tessmann 3694 (NY, G, S). 



4. Piptocarpha canescens Gleason, Bull. Torrey Bot. Club 59: 373. 

 1932. TYPE: Peru: Junin: San Nicholas, Killip & Smith 26084 

 (Holotype: NY! Isotype: US!). 



Subscandent shrub, ca. 3-4 m tall, stems densely cinereous-pubescent. Leaves cauline, 

 petioles stout, densely tomentose, 1-2 cm long; blades thin, elliptic-oblong to elliptic- 

 ovate, acute to acuminate at apex, oblique at base, ca. 15-21 cm long, 6-10 cm wide, mar- 

 gins revolute, primary and secondary veins elevated beneath, glabrous, opaque above ex- 

 cept densely stellate-tomentose on the midvein, 8-10 pairs of lateral veins, stellate- 

 pubescent beneath. Inflorescences in axillary, branching, corymbose clusters, ca. 2.5 cm in 

 diameter (when pressed) with ca. 40 heads. Heads with 6 florets, shortly pedicellate 

 terminating peduncles, pedicels and peduncles densely stellate-tomentose, heads imma- 

 ture on the specimen examined; phyllaries (when dry) brown with dense grayish brown 

 pubescence at the tips; inner phyllaries ovate, tips acute: outer phyllaries ovate. 



According to Gleason, this species is distributed in Depto. Junin. It 

 occurs in dense forest at ca. 1,100 m. It either is not abundant or has 

 been poorly collected. Flowering and fruiting occur from July to Sep- 

 tember. The only collection examined is the type. 



5. Piptocarpha spruce! Baker in Mart., Fl. Bras 6(2): 129. 1873. 

 TYPE: Peru: Tarapoto, Spruce 4362 (Holotype: K! Isotypes: BR! P! 

 F! NY! BM! G! W! GH! E!). 



Liana, stems densely and finely canescent, strongly 4-angled. Leaves cauline: petioles 

 slender, ca. 0.5 cm long; blades elliptic, somewhat rigid and coriaceous when dry, 

 acuminate at the apex, cuneate at the base, 8-13 cm long, 2.5-6 cm wide, margins slightly 

 revolute, remotely toothed, glabrous above but canescent along midvein, veins elevated 

 (in dry specimens), glabrate to cinereous lepidote-black glandular beneath. Inflorescences 

 in dense axillary corymbose clusters of ca. 35 heads, ca. 1.2 cm wide. Heads with 3 

 florets, sessile in groups of 2 at the ends of stout, tomentose peduncles; involucre cam- 

 panulate, 4-5 mm long; phyllaries yellow-brown with a dark tip when dry, tomentulose at 




MACBRIDE: FLORA OF PERU 53 



apex, upper margin finely ciliate; outer phyllaries persistent, ovate, apex obtuse; inner 

 phyllaries deciduous with achenes, ovate-oblong, apex acute to obtuse. Corollas ca. 3 

 mm long, white, glabrate but with occasional glands, lobes revolute 1.5-3 mm long; 

 anthers ca. 3 mm long, basal auricles acute, less than 0.1 mm. Pappus white biseriate, inner 

 bristles equal, filiform, ca. 6 mm long, outer bristles inconspicuous, short, unequal, 0.5- 

 1 mm long. Achenes ca. 3 mm long, 10-costate, glabrous or sparsely pilose. 



This species is found in forests in Loreto where it is rare or poorly 

 collected. Flowering and fruiting occur from August to September. It 

 appears to be closely allied with the Brazilian species Piptocarpha 

 leprosa (Less.) Baker, and further study may show P. sprucei to be the 

 northern range of P. leprosa. 



LORETO: Pumayacu, King 3167 (F, MO). LORETO: Tarapoto, 

 Spruce 4362 (K, BR, P, F, NY). 



6. Piptocarpha lechleri (Sch. Bip.) Baker in Mart., Fl. Bras. 6(2): 

 127. 1873. 



Carphobolus lechleri Sch. Bip., Pollichia 20/21: 428. 1863. TYPE: Peru: prope St. 



Gavan, Lechler 2479 (Holotype: B as photos F! NY! GH! Isotypes: K! G!). 

 Piptocarpha vismiaefolia Gleason, Bull. Torrey Bot. Club 59: 372. 1932. TYPE: Peru: 



Junin: La Merced, Killip & Smith 23848 (Holotype: NY! Isotypes: K! F! US!). 

 Piptocarpha longifolia Gleason, Bull. Torrey Bot. Club 59: 372-373. 1932. TYPE: 



Peru: Junin, Pichis Trail, Yapas, Killip & Smith 25459 (Holotype: NY! Isotype: 



US!). 



Shrubs to slender trees, 3-6 m high, branches slender, long, widely spreading, stems 

 cinereous, appressed tomentulose-lepidote. Leaves cauline, not crowded; petioles stout, 

 sulcate, densely tomentose, 1-2 cm long; blades large, coriaceous, oblong to lanceolate, 

 acuminate at the apex, oblique at the base, 10-22 cm long, 3-10 cm wide, margins slightly 

 revolute and remotely toothed, glabrous and lustrous above, 6-10 pairs of prominent 

 lateral veins, densely cinereous stellate-tomentose and yellow-brown lepidote beneath. 

 Inflorescences in dense, axillary, rounded, umbellate clusters of 9-16 heads. Heads with 

 10-20 florets, stoutly pedicellate, singly or in groups of 2 or 3, at the ends of stout 

 peduncles of a uniform length giving an umbellate appearance to inflorescences; in- 

 volucres broadly campanulate at maturity, 8-10 mm long, 4.5-5.5 mm wide, imbricate in 

 5-6 series; phyllaries uniformly brown when dried, nearly glabrous; outer phyllaries 

 persistent, ovate, margin minutely ciliate, apex obtuse; inner phyllaries deciduous with 

 achenes, oblong-lanceolate, apex acute; flowers subtended by linear, acuminate paleas, 

 deciduous with inner involucral phyllaries. Corollas 4-5 mm long, white, glabrate, lobes 

 revolute, 1.5-2.5 mm long; anthers 3.5-4 mm long, basal auricles caudate, 0.3 mm long. 

 Pappus white, predominately uniseriate, inner bristles equal, 5-6 mm long, outer bristles 

 inconspicuous, less than 1 mm long, totally absent in some specimens. Achenes 3-3.5 mm 

 long. 10-costate, glabrous or sparsely pilose. 



This species is distributed in the Peruvian Andes to eastern Bolivia. 

 It occurs at elevations of 360-1,600 m in dense montane rain forests. 

 Flowering and fruiting occur from June to November. 




54 FIELDIANA: BOTANY 



PERU: near St. Gavan, Lechler2479 (B, K, G). JUNIN: La Merced, 

 Killip & Smith 23848 (NY, K, F, US). JUNIN: Pichis Trail, Yapas, 

 Killip & Smith 25459 (NY, US). SAN MARTIN: Alto Rio Huallaga, 

 Williams 6675 (US). CUZCO: San Lorenzo, C. Vargas C. 11749 (US). 



7. Piptocarpha gutierrezii Cuatrec., Brittonia 8(2): 161-162. 1955. 

 TYPE: Colombia: Antioquia, Municipio Sonson: region de Rioverde, 

 Orilla de Rio Verde de los montes, Gutierrez 35633 (Holotype: F! 

 Isotype: MO!). 



Piptocarpha umbricola Cuatrec.. Brittonia 8(2): 163. 1955. Type: Colombia: Comisaria 

 de Putumayo: Umbria, Klug 1863 (Holotype: F! Isotypes: GH! NY! S! US!). 



Scandent shrubs, 3-6 m high, branches slender, spreading, stems glabrate to minutely 

 silvery lepidote. Leaves cauline, not crowded; petioles slender, sulcate, appressed 

 yellow-brown tomentulose-lepidote, 0.5-2 cm long; blades large, papery, elliptic to obo- 

 vate to ovate-lanceolate, acute to abruptly acuminate at apex, oblique or cuneate to 

 slightly rounded at the base, 9-16 cm long, 4-8 cm wide, margins slightly revolute, entire 

 to very faintly and remotely toothed, glabrous, opaque above, 6-8 pairs of prominent 

 lateral veins, densely and closely silvery cinereous lepidote-tomentose beneath. In- 

 florescences in axillary and terminal panicles. Heads with 6 florets, clustered in groups of 

 6-12 on stout peduncles at the ends of lateral branchlets; involucres cylindrical- 

 campanulate, 6-7 mm long, ca. 3 mm wide, closely imbricate in 3-4 series; phyllaries 

 straw-colored with brown-purple tips, nearly glabrous; outer phyllaries persistent, ovate, 

 margins ciliate, apex obtuse, inner phyllaries deciduous with achenes, ovate-oblong, 

 apex obtuse to acute. Corollas 3.5-4 mm long, white, glabrate, lobes revolute 1.5-2 mm 

 long; anthers ca. 3 mm long, basal auricles caudate, 0.3 mm long. Pappus biseriate, the 

 inner bristles equal, filiform, 5-5.5 mm long, the outer bristles irregular, short, unequal, 

 0.5-1 mm long. Achenes immature, ca. 2.5 mm long, costate, sparsely pilose. 



This species is distributed in the northern Andes ranging from NW 

 Venezuela to N Peru. It occurs in dense montane rain forest and along 

 rivers at elevations of 300-700 m. Flowering and fruiting occur mainly 

 from July to November but occasionally the year round. 



AMAZON AS: Lugar Aintami, Kayap 356 (NY). 



III. POLLALESTA 



Pollalesta H.B.K., Nov. Gen. & Sp. 4: 46. 1820. TYPE: P. ver- 

 nonioides H.B.K.. 



Oliganthes Cass., Bull. Soc. Philom. Paris 10. 1817. TYPE: O. triflora Cass. 

 Odontoloma H.B.K., Nov. Gen. & Sp. 4: 43. 1820. TYPE: O. acuminata H.B.K. 

 Dialesta H.B.K. , Nov. Gen. & Sp. 4: 45. 1820. TYPE: D. discolor H.B.K. 

 Adenocyclus Less., Linnaea 4: 337. 1829. TYPE: A. condensatus Less. 



Shrubs to trees, usually diffusely branched, branches often tomentose. Leaves 

 alternate, petiolate: blades lanceolate to ovate, usually elliptic, cuneate at the base, 

 occasionally oblique, apex acute to long acuminate, margins entire to subserrate, be- 




MACBRIDE: FLORA OF PERU 55 



coming glabrate above, densely stellate below, punctate-glandular both above and 

 below. Inflorescences terminal, corymbose-paniculate. Heads with 1-5 florets; involucre 

 cylindric to narrowly campanulate to strongly compressed; phyllaries 5-18, imbricate, 

 membranous to scarious, receptacle subconvex to flat, naked. Corolla tubular, 5-lobed; 

 stamens 5, anthers basally sagittate: style branches slender. Pappus variable, usually of 2 

 series, outer series of short scales, usually separate but occasionally coroniform, some- 

 times absent; inner pappus of 0-15 aristate bristles. Achenes obconic to slightly truncate, 

 8- 10 ribbed. 



Pollalesta is a small neotropical genus ranging from Central America 

 south into Peru and northern Brasil. One species occurs in Peru. 



REFERENCE 



ARISTEGUIETA, L. 1963. El genero Oliganthes de Madagascar y su equivalente Ameri- 

 cano Pollale sta. Bol. Soc. Venez. Ci. Nat. 23(103): 255-288. 



1. Pollalesta discolor (H.B.K.) Aristeg., Bol. Soc. Venez. Ci. Nat. 

 23(103): 275. 1963. 



Dialesta discolor H.B.K., Nov. Gen. & Sp. 4: 45. 1820. TYPE: Colombia: Honda, 

 Bonpland s.n. (Holotype P, as photo GH! Isotype B, as photo GH!). 



Eupatorium cuspidatum Willd. ex Less., Linnaea 4: 315. 1829. TYPE: 15156 

 (Holotype B, as microfiche Willd. Herb.!). 



Oliganthes discolor (H.B.K.) Sch. Bip., Linnaea 20: 502. 1847. 



O. karstenii Sch. Bip., Linnaea 30: 116. 1859-1860. TYPE: Colombia: Guaduas, 

 Karsten s.n. (Isotype F!). 



O.ferruginea Gleason, N. Amer. Fl. 33: 102. 1922. TYPE: Costa Rica: Forests of Alto 

 de Mano Tigre, Diquis Valley, Pittier 12138 (Holotype US!). 



O. corei Cuatrec., Brittonia 8: 185. 1956. TYPE: Colombia: Dept. Antioquia, El Radio, 

 Core 720 (Holotype WVA! as photo GH! NY! Isotype US!). 



Pollalesta argentea Aristeg., Bol. Soc. Venez. Ci. Nat. 23(103): 275. 1963. TYPE: 

 Peru: Dept. Cajamarca: Valle del Rio Tabaconas, Weberbauer6162 (Holotype F!). 



P. brasiliana Aristeg., Bol. Soc. Venez. Ci. Nat. 23(103): 280. 1963. TYPE: Brasil: 

 Amazonas: Sao Paulo de Olivenca, Ducke 298 (Holotype NY! Isotype MO!). 



P. colombiana Aristeg., Bol. Soc. Venez. Ci. Nat. 23(103): 274. 1963. TYPE: Colom- 

 bia: Villavincencio, Pennell 1406 (Holotype NY! Isotypes GH! US!). 



P. corei (Cuatrec.) Aristeg., Bol. Soc. Venez. Ci. Nat. 23(103): 276. 1963. 



P. ecuatoriana Aristeg., Bol. Soc. Venez. Ci. Nat. 23(103): 277. 1963. TYPE: 



Ecuador: Prov. Napo-Pastaza: cerca de Puyo, Skutch 4428 (Holotype NY, Isotype 



MO!). 



P.ferruginea (Gleason) Aristeg., Bol. Soc. Venez. Ci. Nat. 23(103): 273. 1963. 

 P. karstenii (Sch. Bip.) Aristeg., Bol. Soc. Venez. Ci. Nat. 23(103): 273. 1963. 

 P. klugii Aristeg., Bol. Soc. Venez. Ci. Nat. 23(103): 278. 1963. TYPE: Peru: Dept. 



Loreto: Fortaleza, cerca de Yurimaguas, King 2819 (Holotype GH! Isotype MO!). 

 P. peruviana Aristeg., Bol. Soc. Venez. Ci. Nat. 23(103): 277. 1963. TYPE: Peru: 



Dept. Loreto: Mishuyaca, cerca de Iquitos, King 1242 (Holotype F!). 




56 



FIELDIANA: BOTANY 



FIG. 5. Pollalesta discolor. A, habit; B, head. (From Klug 2220, F.) 



Tree, 10 to 30 m tall, of a single trunk branched in the crown, young stems stellate 

 pubescent, brown to grayish. Leaves somewhat crowded at tips of stems; petiole 1-3.5 cm 

 long; blades elliptic to lanceolate or ovate, acute to long acuminate at the apex, cuneate 

 to oblique at the base, 5-20 cm long, 1.5-9 cm wide, margins mostly entire, sometimes 

 remotely serrate, becoming glandular punctate and glabrate except pubescent on midvein 

 and at base above, densely stellate-pubescent beneath. Inflorescences terminal 

 corymbose-paniculate. Heads with (1)2(3) florets, pedunculate; involucre narrowly cam- 




MACBRIDE: FLORA OF PERU 57 



panulate, 4.5-9 mm long; phyllaries often ciliate, glabrous to slightly pubescent, glandular 

 punctuate near the tips, yellowish-brown, sometimes with dark tips; inner phyllaries 

 oblanceolate; tips acute; outer phyllaries elliptic-ovate. Corollas 5.5-7.5 mm long, whitish 

 to light purple with glandular dots, fragrant. Pappus straw-colored; inner bristles ca. 3-4 

 mm long, outer scales minute to 1.2 mm long. Achenes 1.8-2.4 mm long, obconic, gland- 

 dotted, sometimes thinly pubescent, 8-10 ribbed. 



This species is distributed from Costa Rica into Peru in tropical 

 forest or secondary vegetation from 100-1,600 m elevation. Flowering 

 and fruiting throughout the year. 



LORETO: Florida, Rio Putumayo at mouth of Rio Zubineta, Klug 

 2220 (BM, F, GH, MO, NY, S, US). AMAZONAS: Rio Cenepa, An- 

 cuash 302 (MO). CAJAMARCA: Valle del Rio Tabaconas, Weber- 

 bauer 6162 (F, GH, US). SAN MARTIN: Moyobamba, Klug 3578 (F, 

 GH, K, MO, NY, S). 



IV. CENTRATHERUM 



Centratherum Cass., Diet. Sci. Nat. 7: 384. 1817. TYPE: C. 

 punctatum Cass. 



Spixia Schrank, PI. Rar. Hort. Monac. tab. 80. 1819. TYPE: 5. violacea Schrank. 

 Ampherephis H.B.K., Nov. Gen. & Sp. 4: 31. 1820. TYPE: A. mutica H.B.K. 

 Amphibecis Shrank, Syll. PI. Nov. 1: 86. 1824. TYPE: A. violacea Schrank. 

 Crantzia Veil., Fl. Flum. viii. tab. 153. 1835. TYPE: C. ovata Veil. 



Herbs to subshrubs, often branched, stems glabrescent to villous. Leaves alternate, 

 petiolate to sessile; petioles often indistinct, blades ovate, linear, or oblanceolate, ob- 

 tuse, or subacute to blunt at the apex, cuneate to attenuate at the base, margins serrate, 

 lobed, glabrous, punctate, or pubescent above and beneath. Inflorescences with heads 

 terminal on axillary branches, occasionally 2 or 3 heads clustered together. Heads with 

 numerous florets, sessile; involucre cylindric-campanulate, 8-25 mm wide; phyllaries in 

 several series, outer series foliaceous, intergrading to firm scales, tips variable, rounded 

 to long awned. Corollas tubular, 5-lobed, reddish-purple, glandular, tube sometimes 

 pubescent. Pappus straw-colored, deciduous, of bristles, infrequently absent. Achenes 

 8-10 ribbed, obconic. Chromosome number: n = 16, 32. 



Centratherum is a small tropical genus of two species found in the 

 New World, in Australia, and the Philippines. Formerly, the genus 

 Centratherum included species from India and Java. Based on chromo- 

 some numbers, pollen morphology, and trichome morphology corre- 

 lated with geographical distribution, these Old World species are pres- 

 ently recognized as the genus Phyllocephalum. 



REFERENCE 



KIRKMAN, L. K. Revision of Centratherum and Phyllocephalum (Compositae: Ver- 

 nonieae), Rhodora (in press). 




2cm 



B 



FIG. 6. Centratherum punctatum. A, habit; B, head. (From Woytkowski 7643. F.) 



58 




MACBRIDE: FLORA OF PERU 59 



1 . Centratherum punctatum Cass. Diet. Sc. Nat. 7: 384. 1817. TYPE: 

 Panama: Jussieu s.n. (Holotype: P-JU, as IDC microfiche cat. number 

 8420- JU!). 



Spixia violacea Schrank, PI. Rar. Hort. Monac. Tab. 80. 1819. TYPE: as illustration 

 GH!. 



Ampherephis aristata H.B.K., Nov. Gen. & Sp. 4: 31. 1820. TYPE: Bonpland s.n. 



(Holotype: P, as photo TEX!; Isotype P!). 

 A. mutica H.B.K., Nov. Gen. & Sp. 4: 31. 1820. TYPE: Humboldt & Bonpland 



(Holotype: P, as photo GH!). 



Amphibecis violacea (Schrank) Schrank, Syll. PI. Nov. 1: 86. 1824. 

 Ampherephis pulchella Cass., Diet. Sc. Nat. 57: 346. 1828. TYPE: d'Urville and 



Lesson s.n. not seen. 



A. pilosa Cass., Diet. Sc. Nat. 57: 346. 1828. TYPE: based upon A. mutica Kunth. 

 Centratherum brevispinum Cass., Diet. Sc. Nat. 57: 346. 1828. TYPE: same as A. 



aristata H.B.K. 

 C. longispinum Cass., Diet. Sc. Nat. 57: 346. 1828. TYPE: based upon C. punctatum 



Cass. 



Ampherephis intermedia Link, Abbild. 5 tab. 29. 1828. TYPE: not seen. 

 Centratherum muticum (H.B.K.) Less., Linnaea 4: 320. 1829. 

 C. intermedium (Link) Less., Linnaea 4: 320. 1829. 



Crantzia ovata Veil., Fl. Flum. viii. tab. 153. 1835. TYPE: as illustration!. 

 Centratherum pulchellum (Cass.) Steud., Norn. ed. II. 324. 1840. 

 C. punctatum Cass. var. parviflorum Baker in Mart., Fl. Bras. 6(2): 12. 1873. TYPE: 



Blanche! 3689 (BRAZIL: Bahia: Holotype: K! Isotypes: BR! F! G! LE! MO! P!). 

 C. holotoni Baker in Mart., Fl. Bras 6(2): 12. 1873. TYPE: BRAZIL: Ibague, Holton 



301 (Holotype: K!). 

 C. brachylepis Sch. Bip. ex Baker in Mart., Fl. Bras. 6(2): 12. 1973. TYPE: BRAZIL: 



Martius 461 (Holotype: M! as photo GH! NY! TEX!). 

 Baccarodes holtonii (Baker) O. Ktze., Rev. Gen. PI. 1: 320. 1891. 



B. brachylepis (Sch. Bip. ex Baker) O. Ktze., Rev. Gen. PI. 1: 320. 1891. 

 B. violaceum (Schrank) O. Ktze., Rev. Gen. PI. 1: 320. 1891. 



B. punctatum (Cass.) O. Ktze., Rev. Gen. PI. 1: 320. 1891. 



B. muticum (H.B.K.) O. Ktze., Rev. Gen. PI. 1: 320. 1891. 

 Centratherum aristatum non Cass. Index Kew. 1: 478. 1895. 



C. punctatum Cass. var.foliosa Chod., Bull. Herb. Boissier 2(2): 298. 1902. TYPE: 

 PARAGUAY: Capibuy, Hassler4378. (Holotype: G! Isotype: BM! K! NY! P!). 



C. punctatum Cass. ssp. camporum Hassl. var. viscosissimum Hassl. /. foliosum 

 (Chod.) Hassl., Feddes Repert. Spec. Nov. Regni Veg. 12: 369. 1913. 



C. punctatum Cass. ssp. camporum Hassl. var. viscosissimum Hassl. f. brachyphyl- 

 lum Hassl. Feddes Repert. Spec. Nov. Regni Veg. 12: 369. 1913. TYPE: 

 PARAGUAY: In regione vicine Igatimi, Hassler4768 (Holotype: G! Isotypes: GH! 

 MO! MPU! NY! P! S!). 



C. punctatum Cass. ssp. camporum Hassl. var. longipes Hassl.. Feddes Repert. Spec. 




60 FIELDIANA: BOTANY 



Nov. Regni Veg. 12: 369. 1913. TYPE: PARAGUAY, Fiebrig4532 (Holotype: B, as 



photo GH! TEX!). 



C. violaceum (Schrank) Gleason, N. Amer. Fl. 33: 49. 1922. 

 C. camporum (Hassl.) Malme var. longipes (Hassl.) Malme, Ark. Bot. 24A 6: 15. 1931. 



Sprawling to erect herb becoming suffrutescent with age, stems strigose, often ridged. 

 Leaves cauline, often crowded; short petiolate to sessile; blades ovate to elliptic to 

 spatulate, obtuse at the apex, cuneate to attenuate at the base, (1)2-7 cm long, (0.5)0.8-3 

 cm wide, margins serrate, often ciliate, glandular-punctate, often pubescent (especially 

 on veins) above and beneath. Inflorescences of terminal heads, or occasionally 2-3 headed 

 clusters, peduncles 2-7 cm long. Heads with numerous florets; involucre cylindric- 

 campanulate, imbricate hi several series; phyllaries glandular, membranaceous, outer 

 foliaceous, greenish; inner phyllaries purplish, rounded to aristate (when awned, awns to 

 3 mm). Corollas 5-8(10) mm long, glandular. Pappus straw-colored; bristles numerous, 

 deciduous 1.5-2.8(3.5) mm long, rarely absent. Achenes (1.2)1.6-2.6 mm long, 8-10 

 ribbed. Chromosome number: n = 16, 32. 



This subspecies occurs in South and Central America and the West 

 Indies. It grows in pastures and waste places, flowering the year 

 around. It is sometimes cultivated as an ornamental. 



The one specimen seen was: PERU: no locality cited, Woytkowski 

 7643 (MO). The material from Peru represents C. punctatum Cass. ssp. 

 punctatum. 



V. STRUCHIUM 



Struchium P. Br., Civ. Nat. Hist. Jam. 312, tab. 34, fig. 2. 1756. 

 TYPE: S. herbaceum St.-Hil. 



Athenaea Adans. Fam. 2: 121. 1763, non Sendtn. (Solanaceae), nom. cons. 

 Sparganophorus Vaill. ex Crantz, Inst. 1: 261. 1766. 



Erect, weedy, annual herbs, stems simple or branched, somewhat succulent. Leaves 

 alternate, simple, petiolate, blades subentire to serrate, pinnately veined. Inflorescences 

 axillary, of single or glomerate heads. Heads discoid; involucre hemispheric; phyllaries 

 numerous, imbricated in several series. Corollas tubular, 3-lobed, white; style branches 

 reddish-purple. Pappus a cartilaginous, whitish corona, ca. one-half the length of the 

 achene. Achenes 3-4 angled. 



A monotypic genus of the New World tropics that is reportedly 

 weedy in Africa. 



1. Struchium sparganophorum (L.) O. Ktze., Rev. Gen. PI. 366. 

 1891. 



Ethulia sparganophora L., Sp. PI., ed. 2. 1171. 1763. TYPE: not seen. 

 Struchium herbaceum P. Br. ex St.-Hil., Expos. Fam. 1: 406. 1805. 

 Sparganophorus Struchium Poir. in Lam., Encycl. Meth. 7: 302. 1806. (July). TYPE: 

 P. Br., Civ. Nat. Hist. Jam. 312, tab. 34, fig. 2. 1756. 




f 



FIG. 7. Struchium sparganophorum. A, habit, x Vi\ B, detail showing axillary in- 

 florescences, natural size; C, mature head showing achenes and receptacle, x 5; D, 

 mature head with fallen corollas showing top achenes, x 3Vi; E, flower at anthesis, x 16; 

 F, corolla showing position of lobes and stamens, x 18, stigma, x 25. (From Steyermark 

 46308, F.) 



61 




62 FIELDIANA: BOTANY 



S.fasciatus Poir. in Lam., Encycl. Meth. 7: 302. 1806. TYPE: Lam. 111. Genres, tab. 



670. 1823. 

 Struchium americannm Poir. in Lam., Encycl. Meth. 7: 475. 1806. TYPE: P. Br., Civ. 



Nat. Hist. Jam. 312, tab. 34, fig. 2. 1756. 

 Ethulia struchium Sw., Prod. Veg. Ind. Occ. 3: 1297. 1806. TYPE: P. Br., Civ. Nat. 



Hist. Jam. 312, tab. 34, fig. 2. 1756. 



Annual herbs, 0.4-1 m tall, stems simple or branched, somewhat succulent, stout, 

 sparsely short-strigose. Leaves cauline: petioles ca. 1 cm long; blades elliptic, acuminate 

 at the apex, cuneate at the base, 9-12 cm long, 3-7 cm wide, margins subentire to serrate, 

 inconspicuously strigose to glabrate on both surfaces. Inflorescences of solitary or glom- 

 erate heads in the leaf axils. Heads with 60-70 florets, sessile or on short branches; 

 involucre hemispheric, 3-5 mm long; phyllaries arachnoid, appressed, greenish with 

 white margins, tips acuminate. Corollas 2 mm long, white, 3-lobed, glabrous; style 

 branches reddish purple. Pappus a cartilaginous corona. Achenes 2 mm long, 3- to 4- 

 angled, glandular. 



This species is distributed throughout tropical America and is re- 

 portedly adventive in Africa. It grows in moist alluvial or sandy soil 

 along streams or in flood plains. Flowering and fruiting occur through- 

 out the year. 



LORETO: Pongo de Manseriche, Mexia 6350 (MO, UC, US). SAN 

 MARTIN: Tocache Nuevo, Vigo 7144 (MO). 



VI. ELEPHANTOPUS 

 Elephantopus L., Sp. PI. 814. 1753. TYPE: Elephantopus scaber L. 



Orthopappus Gleason, Bull. New York Bot. Card. 4: 237. 1906. TYPE: Elephantopus 

 angustifolius Sw. 



Erect perennial herbs, simple or sparsely branched; stems usually solitary and pubes- 

 cent. Leaves cauline or chiefly basal; petioles usually indistinct; blades elliptic to lanceo- 

 late or ovate, acute at the apex, attenuate at the base, margins entire to crenate or 

 dentate. Inflorescences spicate, corymbose, or slightly paniculate; bracteate; the heads in 

 ovoid or globose glomerules, the glomerules dense with many heads. Heads with (1)2-4(5) 

 florets; involucre of 8 phyllaries, in 4 decussate pairs. Corollas blue to white, tubes 

 slender, the limb unequally 5-cleft with a deeper fissure on the inner side. Chromosome 

 number: n = 11, 22. 



Elephantopus is a largely tropical genus of ca. 30 species centered in 

 the New World but also occurring in the Old World tropics. 



REFERENCES 



BAKER, C.F. 1902. A revision of the Elephantopeae. Trans. Acad. Sci. St. Louis. 12, pp. 



43-56. 

 CLONTS, J. A. 1972. A revision of the genus Elephantopus, including Orthopappus and 



Pseudelephantopus (Compositae). Unpublished Ph.D. dissertation. Miss. State Univ. 

 CLONTS, J. A. AND S. MCDANIEL. 1978. Elephantopus. N. Amer. Fl. Ser. II, pt. 10, pp. 



196-202. 




MACBRIDE: FLORA OF PERU 63 



KEY TO SPECIES OF Elephantopus 



a. Inflorescence paniculate, glomerules pedunculate; pappus bristles 5-8 



1. E. mollis. 



aa. Inflorescence appearing spicate. glomerules sessile, pappus bristles numerous 

 (ca. 20-30) 2. E. angustifolius. 



1. Elephantopus mollis H.B.K., Nov. Gen. & Sp. 4: 26. 1820. TYPE: 

 Caracas, Caracas, Humboldt and Bonpland 627 (Holotype P, as IDC 

 microfiche!). 



E. martii Gran., Edinburgh New Philos. J. Jan.-Mar. 378. 1830. TYPE: Brazil: Rio 



Janeiro, Harris s.n. (not seen). 

 E. sericeus Grab., Edinburgh New Philos. J. Jan.-Mar. 373. 1831. TYPE: Dominica: 



Krous s.n. (not seen). 



E. serratus Blanco, Fl. Filip. ed 1. 635. 1837. 



E. carolinanus var. mollis (H.B.K.) Beurl., Bidr. Portobellensis F. 134. 1854. 

 E. hypomalacus Blake, Contr. Gray Herb. 52: 20. 1917. TYPE: Costa Rica, Holway 



314 (Holotype GH). 

 E. pilosus Philipson, J. Bot. 77: 314. 1939. TYPE: Dutch Guiana: Hostmann 875 



(Holotype BM, Isotype K). 



Erect perennial herbs, 3-10(20) cm tall, from a creeping rootstock, stems pilose or 

 hirsute. Leaves cauline or basal, greatly reduced upward: petioles short and clasping the 

 stem; blades ovate, obovate, or oblanceolate, acute to short acuminate at the apex, 

 attenuate to the base, 8-20 cm long, 5-10 cm wide, margins crenate. serrate to subentire. 

 thinly pilose to slightly scabrous above, softly pilose beneath. Inflorescences 

 corymbose-paniculate, glomerules terminal, to 2.3 cm wide: bracts 3, cordate to deltoid. 

 0.7-1 cm wide, 0.7-1.3 cm long, occasionally longer or shorter than the glomerules, acute 

 to short acuminate, pilose. Heads with 4 florets; phyllaries lanceolate, 1 .5-2 mm wide, 6-8 

 mm long, sharply acuminate, membranous along the margin, at least below, sparsely 

 pubescent, at least above the middle. Corollas 5-6 mm long, white to bluish pink, deeply 

 divided on the adaxial side. Pappus of 5(8) bristles unseriate, ca. 4 mm long dilated into a 

 narrow to broad triangular base. Achenes 2.5-3.5 mm long, ribbed minutely pubescent. 

 Chromosome number: n = 11. 



This species is widely distributed in the American tropics and has 

 been introduced into tropical Africa and southeast Asia. Flowering and 

 fruiting occur throughout the year. 



AMAZONAS: Mendoza, Woytkowski 81 1 1 (MO). LORETO: Lago 

 Llanchama near Rio Nanay, Croat 18756 (MISS A, MO). SAN MAR- 

 TIN: Jepelacio near Moyobamba, King 3464 (F, MO, US). 

 HUANUCO: Tingo Maria, Asplund 12155 (US). JUNIN: near La 

 Merced, Killip and Smith 23972 (US). AYACUCHO: La Mar: Between 

 Ayna and Hacienda Luisiana, Dudley 11686 (USM). CUZCO: 

 Machupicchu, Vargas 806 (F, USM). MADRE DE DIGS: ca. 20 km W 

 of Puerto Maldonado, Gentry, Revilla, Alfaro, Daly 19677 (MO). 




FIG. 8. Elephantopus mollis. A, habit, x Vi; B, inflorescence, x 2; C, flowering head 

 with detail, x 3; D, corollas, one dissected to show detail, x 13; E, anther, x 25. (From 

 Standley 76193, F.) 



64 




MACBRIDE: FLORA OF PERU 65 



2. Elephantopus angustifolius Sw., Prod. Veg. Ind. Occ. Prodr. 115. 

 1788. Based on Sloane, Voy. Isl. Madera. 1: 256, pi. 148, fig. 4. 1707. 



E. nudiflorus Willd. Sp. PI. 3: 2390. 1804. TYPE: St. Domingo, Poiteau (not seen). 

 Elephantosis quadriflora Less., Linnaea4: 323. 1829. TYPE: Brazil, Beyrich s.n. (not 



seen). 



Elephantopus quadriflorus (Less.) D. Dietr., Syn. PI. 4: 1372. 1847. 

 Orthopappus angustifolius (Sw.) Gleason. Bull. New York Bot. Gard. 4:237. 1906. 



Perennial erect herbs arising from a short caudex, usually 3-12(15) dm tall, stems 

 pilose. Leaves cauline and basal, crowded near the base; petioles short and broad; blades 

 narrowly oblanceolate to oblong-lanceolate or linear to oblong, acute to obtuse at the 

 apex, long attenuate at the base, (5)10-35(50) cm long, 1.3-4(5.5) cm wide, margins 

 shallowly and irregularly crenate, thinly and softly strigose on both surfaces, the 

 trichomes somewhat silvery beneath. Inflorescences spicate or racemose-spicate below, 

 glomerules lateral and terminal, to 2 cm wide, bracts 1 or 2, lanceolate, to 1 cm long. 

 Heads with 4 florets; phyllaries 8, in 2 descussate series, acute to acuminate, membra- 

 nous along the margin, sparsely strigose at least above the middle. Corolla tubes to 6 mm 

 long, the limb 2 mm long, white to lavender, 5 parted, deeply divided on the adaxial side. 

 Pappus of 20-40 bristles, uniseriate, 6-7 mm long, gradually dilated near the base. Achenes 

 1.5-2.5 mm long, ribbed, pubescent. Chromosome number: n = 11. 



This species is distributed from southern Mexico south to northern 

 Argentina and Chile and into the West Indies. Flowering and fruiting 

 occur throughout the year. 



AMAZONAS: 4 km from Campamento Ingenio, Hutchison and 

 Wright 3980 (F, MO, US). LORETO: Yurimaguas, McDaniel and 

 Rimachi Y 16554 (MISSA, F, MO). SAN MARTIN: Tarapoto, 

 McDaniel 13724 (F, MO, MISSA). HUANUCO: Pachitea, Camino a 

 Shahuinto a 5 km del campamento de Iparia, Schunke 1667 (F). 

 JUNIN: San Ramon, Woytkowski 7487 (MO, US). CUZCO: Herrera 

 3232 (F). 



VII. PSEUDELEPHANTOPUS 



Pseudelephantopus Rohr, Skr. Naturhist.-Selsk. 2: 213. 1792. 

 "PseudoElephantopus," TYPE: Elephantopus spicatus Juss. ex Aubl. 



Distreptus Cass., Bull. Soc. Philom. Paris 1817: 66. 1817. TYPE: Elephantopus 



spicatus Juss. 

 Matamoria La Llave & Lex., Nov. Veg. Desc. fasc. 1: 8. 1824. TYPE: Elephantopus 



spicata Juss. ex Aubl. 

 Spirochaeta Turcz., Bull. Soc. Imp. Naturalistes Moscou 24: 166. 1851. TYPE: 5. 



funckii Turcz. 

 Chaetospira Blake, J. Wash. Acad. Sci. 25: 311. 1935. TYPE: Spirochaeta funckii 



Turcz. 




66 FIELDIANA: BOTANY 



Erect herbs, stems solitary, branched. Leaves cauline, clasping, alternate, pinnately 

 veined; petioles indistinct. Inflorescences terminal, slender, racemose-spicate, in 

 glomerules of 1-5 heads usually subtended by 2 foliaceous bracts. Heads with 4 florets: 

 involucre of 8 phyllaries. Corollas tubular-funnelform, the tube slender, the limb 5-cleft, 

 deeply divided on one side; anthers sagittate at the base; style slender. Pappus of 5-15 

 unequal or subequal bristles, uniseriate, plicate or spiraled at the tip, some straight 

 bristles in 1 species. Achenes 10-ribbed. 



A neotropical genus of two species, both known from Peru; in- 

 troduced pantropically. 



Cronquist (1971) discusses the spelling of Pseudelephantopus and 

 maintains it as a genus. It was recognized as a genus by Busey (1975); 

 however, Clonts (1972) submerged it in Elephantopus. The two genera 

 Pseudelephantopus and Elephantopus differ in gross morphology of 

 the inflorescence and in other features. Pseudelephantopus differs 

 cytologically having a chromosome number of A? = 13; Elephantopus 

 has = 11, 22. Because of the cytological differences and its spe- 

 cialized morphology, it clearly merits generic status. 



REFERENCES 



BUSEY, P. 1975. Elephantopodinae. Flora of Panama, part IX, Ann. Missouri Bot. Card. 



62, pp. 873-883. 

 CLONTS, J. A. 1972. A revision of the genus Elephantopus including Orthopappus and 



Pseudelephantopus (Compositae). Ph.D. dissertation. Miss. State Univ., Starkville. 

 CRONQUIST, A. 1971. Composite. In I. L. Wiggins and D. M. Porter, Flora of the 



Galapagos, pp. 350-353. Stanford Univ. Press, Stanford. 

 GLEASON, H. A. 1922. Pseudelephantopus, N. Amer. Fl. 33, p. 109. 



KEY TO SPECIES OF Pseudelephantopus 



a. Pappus consisting of 2 bristles bent and curled and several short straight bristles . . 



1. P. spicatus. 

 aa. Pappus bristles curled or twisted 2. P . spiralis. 



1. Pseudelephantopus spicatus (Juss. ex Aubl.) C. F. Baker, Trans. 

 Acad. Sci. St. Louis 12: 55. 1902. Based on Sloane, Voy. Isl. Madera 1: 

 256, pi. 150, fig. 3-4. 1707. 



Elephantopus spicatus Juss. ex Aubl. PI. Gui. 2: 808. 1775. 

 Distreptus spicatus (Juss. ex Aubl.) Cass., Diet. Sc. Nat. 13: 667. 1819. 

 Matamoria spicata (Juss. ex Aubl.) La Llave & Lex., Nov. Veg. Desc. fasc. 1: 8. 

 1824. 



Erect perennial herb up to 1 m tall, stems pilose or hirsute, striate. Leaves cauline; 

 petioles indistinct; blades lanceolate, oblanceolate, obovate, reduced above, acute at the 

 apex, attenuate, clasping at the base, (3)4-10(17) cm long, 1-5 cm wide, margins remotely 

 serrate to sinuate, hispid above, pilose to hirsute and punctate beneath. Inflorescences 

 racemose-spicate; clusters of heads subsessile. lateral and terminal, 3-5 headed. Heads 

 with 4 florets; phyllaries 8, similar, in 4 pairs, lanceolate, keeled, 9-12 mm long, 1.5-2(3) 




1mm 



B 



2cm 



FIG. 9. Pseudelephantopus spiralis. A, habit; B, head; C, achene. (From Dillon & 

 Turner 1421, F.) 



67 




68 FIELDIANA: BOTANY 



mm wide, sparsely pubescent at apex, tips sharply acuminate. Corollas 6-10 mm long, 

 white to blue-purple. Pappus of 4-10 bristles, 2-seriate, 2-plicate, 5-7 mm long, the re- 

 mainder straight and shorter, gradually dilated at the base. Achenes 5-7 mm long, ribbed, 

 pubescent. Chromosome number: n = 13. 



This species is distributed from Central Mexico and the West Indies 

 south to Chile, and has become a pantropical weed. Flowering and 

 fruiting occur throughout the year. 



SAN MARTIN: San Martin: Tarapoto Hills, Plowman 6026 (USM). 

 LORETO: Coronel Portillo: cerca a Neshuya, Ferreyra 17193 (USM). 

 JUNIN: Tarma: La Merced, Chanchamayo, Ferreyra 0490 (USM). 



2. Pseudelephantopus spiralis (Less.) Cronq., Madrono 20: 255. 1970. 



Distreptus spiralis Less. Linnaea6: 690. 1831. TYPE: Jamaica, Herb. Thunberg 20920 



(UPS). 

 Spirochaetafunckii Turcz., Bull. Soc. Imp. Naturalistes Moscou 24: 167. 1851. TYPE: 



Venezuela, LaGuayra, Funck 358, Galeotii Herb. 380 (G-Delessert Herb. 28530). 

 Chaetospira funckii (Turcz.) Balke, J. Wash. Acad. Sci. 25: 311. 1935. 

 Pseudelephantopus funckii (Turcz.) Philipson, J. Bot. 76: 301. 1938. 

 Chaetospira spiralis (Less.) Aspl. & Blake, Svensk Bot. Tidskr., 52: 50. 1958. 



Erect, perennial, stoloniferous herb, stems pilose to hirsute. Leaves cauline; petioles 

 indistinct; blades oblanceolate to obovate, acute to obtuse at the apex, attenuate at the 

 base, 3-7(15) cm long, 1.2-3(5) cm wide, margins crenate, hispid above, punctate and 

 hispid beneath. Inflorescences racemose-spicate, bracteate, clusters of heads subsessile, 

 5-10 headed. Heads with 4 florets; phyllaries 8, similar, in 4 pairs, oblong-lanceolate, 

 keeled, 7-8 mm long, 1-2 mm wide, pubescent above middle, tips acuminate. Corollas 6-7 

 mm long, whitish to blue-purple. Pappus of 4-6 bristles, uniseriate, 4-6 mm long, strongly 

 twisted above the middle, dilated at the base. Achenes 2.5-3 mm long, ribbed, pubescent. 



This species is distributed from Costa Rica and the Lesser Antilles 

 throughout northern South America south to northern Argentina and 

 occurs as a weed in pastures and waste places. Flowering and fruiting 

 occur throughout the year. 



SAN MARTIN: San Martin: cerca a Tarapoto, Ferreyra 17863 

 (USM). LORETO: Mishuyacu, King 1331 (F). MADRE DE DIGS: 

 Iberia, Seibert 1928 (US, USM). MAYNAS: Bellavista, McDaniel 

 16052 (MISSA, MO). JUNIN: Rio Pinedo, N of La Merced, Killip and 

 Smith 23596 (F). 



ACKNOWLEDGMENTS 



The assistance of Ms. Kay Kirkman and Mr. John Stutts with the 

 treatments ofCentratherum and Pollalesta is gratefully acknowledged. 

 My wife, Carleen A. Jones, assisted with the field work in Peru. Ap- 

 preciation is extended to Dr. Ramon Ferreyra for his hospitality during 




MACBRIDE: FLORA OF PERU 69 



our visit to Peru. Mrs. Nancy C. Coile, Mr. Greg Jones, and Mr. Oliver 

 Ware assisted with the preparation of the manuscript, literature re- 

 view, and herbarium tasks. The work was supported by the University 

 of Georgia and by a grant from the National Science Foundation. Dr. 

 Al Gentry supplied the Latin diagnoses for the new species. Drs. 

 Michael Dillon and Ramon Ferreyra provided helpful suggestions re- 

 garding the manuscript. 



Figures 7 and 8 were drawn by Marion Pahl and are used with the 

 permission of the Editor of Fieldiana. The remaining figures were 

 drawn by Marlene Werner, Department of Exhibition, Field Museum 

 of Natural History. 




INDEX OF LATIN NAMES 



Names in boldface refer to new species; names in Roman type refer 

 to valid species; names in italics refer to synonyms. Numbers in 

 boldface refer to descriptions; numbers in italics refer to illustrations. 



Achyrocoma 24 



tomentosa 24 

 Ac Heps is 24 



squarrosa 24 

 Ambassa 25 



hochstetteri 25 

 Ampherephis 57 



arista ta 59 



intermedia 59 



mutica 57, 59 



pilosa 59 



pulchella 59 

 Amphibecis 57 



violacea 57, 59 

 Ascaricida 24 



indica 24 

 Athenaea 60 

 Baccarodes 24 



anthelmintica 24 



brachylepis 59 



holtonii 59 



muticum 59 



punctatum 59 



violaceum 59 

 fle/7? 24 



noveboracensis 24 

 Bracheilema 24 



paniculatum 24 

 Cacalia argyropappa 46 



aschenbarniana 35 



baccharoides 35 



brachiata 41 



bullata 44 



canescens 44 



ferruginea 33 

 gracilis 3 1 

 haenkeana 35 

 lanceolaris 35 

 laurifolia 32 

 mollis 44 

 moritziana 3 1 

 myriocephala 43 

 patens 35 

 salzamannii 46 

 scorpioides 39 

 sordidopapposa 32 

 suave olens 35 

 tournefortioides 39 



Candidea 24 

 senegalensis 24 



Carphobolus 47 

 asterotrichus 49 

 latifolius 5 1 

 lechleri 53 

 poeppigiana 48 

 sessUjflorus 47 

 tereticaulis 48 



Centrapalus 24 

 galamensis 24 



Centratherum 57 

 aristatum 59 

 brachylepis 59 

 brevispinum 59 

 camporum 60 

 holotoni 59 

 intermedium 59 

 longispinum 59 

 muticum 59 



70 




MACBRIDE: FLORA OF PERU 



71 



pulchellum 59 



var. pan'iflorum 59 



punctatum 57, 58, 59 

 var. foliosa 59 



ssp. camporum var. longipes 59 

 var. viscosissimum f. brachyphyl- 

 lum 59 



f . foliosum 59 

 ssp. punctatum 60 



violaceum 60 

 Chaetospira 65 



funckii 68 



spiralis 68 

 Cheliusia 24 



abyssinica 24 

 Chrysocoma herbaceae 46 

 Claotrichelus 24 



rupestris 24 

 Conyza scorpioides 38 



populifolia 24 

 Crantzia 57 



ova/a 57, 59 

 Critoniopsis 25,28 



/mrff/i/V 25,28 

 Crystallopollen 25 



angustifolium 25 

 Cyanopis 24 



villosa 24 

 Cyanopsis 24 

 Cyanthillium 24 



moluccense 24 

 Dialesta 54 



discolor 54, 55 

 Distephanus 24 

 Distreptus 65 



spicatus 66 



spiralis 68 

 Elephantopus 22, 62, 66 



angustifolius 62, 65 



carolinanus var. mollis 63 



hypomalacus 63 



martii 63 



mollis 63, 64 



nudiflorus 65 



pilosus 63 



quadriflorus 65 



scaber 62 



sericeus 63 



serratus 63 



spicatus 65, 66 



Elephantosis quadriflora 65 

 Eremosis 25 



salicifolia 25 

 Ethulia sparganophora 60 



struchium 62 

 Eupatorieae 22 

 Eupatorium cuspidatum 55 

 Flustula 24 



tomentosa 24 

 Gymnanthemum 24 



congestum 24 

 Hololepis 24 



pedunculata 24 

 Isonema 24 



ovata 24 

 Leiboldia 24 



leiboldiana 24 

 Lepidaploa 24 



canescens 44 



scorpioides 39 

 Linzia 24 



glabra 24 

 Llerasia 24 



lindeni 24 

 Lysistemma 25 



indica 25 

 Matamoria 65 



spicata 66 

 Monanthemum 47 



cruegerii 47 

 Monosis 24 



wightiana 24 

 Odontoloma 54 



acuminata 54 

 Oliganthes 54 



corf/ 55 



discolor 55 



ferruginea 55 



karstenii 55 



triflora 54 

 Orthopappus 62 



angustifolius 65 

 Phyllocephalum 57 

 Piptocarpha 47, 48 



asterotrichia 49,50,51 



brasiliana 47 



canescens 52 



chontalensis 48 



costaricensis 48 



foliosa 49 




72 



FIELDIANA: BOTANY 



gutierrezii 54 



insiginis 49, 51 



laxa 49 



lechleri 53 



leprosa 53 



longifolia 53 



opaca 51 



var. latifolia 51 



paraensis 49 



poeppigiana 48 



sprucei 52, 53 



tereticaulis 48 



umbricola 54 



vismiaefolia 53 

 Plectreca 24 



corymbosa 24 

 Pollalesta 54, 55 



argentea 55 



brasiliana 55 



colombiana 55 



corf/ 55 



discolor 55, 56 



ecuatoriana 55 



ferruginea 55 



karstenii 55 



klugii 55 



peruviana 55 



vernonioides 54 

 Polydora 24 



stoechadifolia 24 

 Pseudelephantopus 65, 66 



funckii 68 



spicatus 66 



spiralis 67, 68 

 Punduana 25 



volkameriaefolia 25 

 Senecioides 25 



cinereum 25 



Serratula noveboracensis 24 

 Sparganophorus 60 



fasciatus 62 



struchium 60 

 Spirochaeta 65 



/u/icM 65, 68 

 Spixia 57 



violacea 57, 59 

 Staehelina solidaginoides 39 

 Stengelia 24 



adoensis 24 

 Stenocephallum 25 



monticolum 25 



Stoke sia 22 

 Strobocalyx 25 



arborea 25 

 Struchium 60 



americanum 62 



herbaceum 60 



sparganophorum 60,67 

 Suprago 24 



glauca 24 

 Teichostemma 24 



fruticosum 24 

 Tephrothamnus 25 



pycnanthus 25 

 Trianthaea 24 

 Turpina 24 



tomentosa 24 

 Vernonia 22, 23 



africana 24 



apurimacensis 38 



arborescens var. cuneifolia 44. 45 



argyropappa 46 



aschenborniana 35 



asterotrichia 49 



baccharoides 35, 36 



bangii 35 



brachiata 41,42 



bullata 44 



cainarachiensis 41 



canescens 44 



costata 34 



cotaniensis 37 



cuneifolia 44 



digit at a 41 



ferruginea 33 



fieldiana 45 



flavescens 39 



floribunda 35 



fulta 36 



geminiflora 46 



glauca 24 



gracilis 31 



haenkeana 35 



herbaceae 46 



jalcana 29 



jelskii 30 



var. virescens 30 



lambayequensis 28 



lanceolaris 25 



languinosa 39 



laurifolia 32 



Hbertadensis 31 




MACBRIDE: FLORA OF PERU 



73 



Undent! 28 

 longeracemosa 39 

 mapirensis 33 

 megaphylla 41 

 micradenia 35 

 mollis 44 

 monsonensis 35 

 moritziana 31 

 myriocephala 43, 44 

 noveboracensis 24 

 obovata 46 

 opaca 51 

 pacchensis 35 



var. tambillensis 35 

 patens 35, 36, 57 

 patulifolia 44 

 paucifolia 46 

 peruviana 30 

 poeppigiana 46, 48 

 pseudomollis 45 

 pycnantha 28 

 rusbyi 45 

 saepium 39 

 salamana 36 

 salzmanii 46 

 sambravana 34 



scorpioides 38, 39, 40 

 a centriflora 39 

 longeracemosa 39 

 8 longifolia 39 

 subrepanda 39 

 y subtomentosa 39 



sordidopapposa 32 



stuebelii 34 



suaveolens 35 



subrepanda 38 



tereticaulis 48 



tournefortioides 39 



trichoclada 33 



trixioides 36 



vargasii 43 



volubilis 44 



weberbaueri 35 



woytkowskii 29 



yurimaguasensis 43 

 Vernoniaceae 22 

 Vernonieae 22, 23 

 We>6/a 24 



pinifolia 24 

 Yernonella 24 

 Xipholepis 25 



silhetensis 25 





EQUADOR 



^ v^/*\ 



8 ,.. *V_ 



COLOMBIA 



BRAZIL 



1 TUMBES 



2 PIURA 



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10 HUANUCO 



11 LIMA 



12 PASCO 



13 JUNIN 



14 HUANCAVELICA 



15 ICA 



16 AYACUCHO 



17 APURIMAC 



18 CUZCO 



19 MADRE DE DIGS 



20 AREQUIPA 



21 PUNO 



22 MOQUEGUA 



23 TACNA 



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K 



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PERU 

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