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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 Giseke1 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.

2Assisted 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

MORPHOLOGY3

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

3Adapted 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 CoMPosiTAE4

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.

5As 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. PIPTOCARPHA6

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.

6Mr. 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

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