UNIVERSITY OF 

ILLINOIS LIBRARY 

AT URBANA-CHAMPAIGN 

BIOLOGY 

< 2001 




Botany 

NEW SERIES, NO. 35 




Ramon Ferreyra 



May 31, 1995 
Publication 1466 



PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY 



rs to Fieldiana 












ATION IS PRINTED ON ACID-FREE PAPER. 



FIELDIANA 



Botany 

NEW SERIES, NO. 35 



FLORA OF PERU 

J. Francis Macbride and Collaborators 

Family Asteraceae: Part VI 

Ramon Ferreyra 

Museo de Historia Natural "Javier Prado" 
Lima, Peru 



Accepted February 17, 1993 

Published May 31, 1995 B|QLOGY L|BRARY 

Publication 1466 1Q1 BURR|L| _ HALl 

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY 






1995 Field Museum of Natural History 

ISSN 00 15-0746 
PRINTED IN THE UNITED STATES OF AMERICA 



Table of Contents 



ABSTRACT 1 

TRIBE MUTISIEAE 1 

Key to Genera of Mutisieae in Peru 2 

I. Arnaldoa 4 

Key to Species of Arnaldoa 4 

1. Arnaldoa coccinosantha 4 

2. Arnaldoa macbrideana 5 

3. Arnaldoa weberbaueri 5 

II. Barnadesia 7 

Key to Peruvian Species of 

Barnadesia 7 

1 . Barnadesia berberoides 8 

2. Barnadesia blakeana 10 

3. Barnadesia caryophylla 10 

4. Barnadesia corymbosa 11 

5. Barnadesia dombeyana 11 

6. Barnadesia horrida 12 

7. Barnadesia hutchisoniana 13 

8. Barnadesia jelskii 13 

9. Barnadesia lehmannii 14 

1 0. Barnadesia macbridei 14 

1 1 . Barnadesia polyacantha 14 

12. Barnadesia reticulata 15 

1 3. Barnadesia wurdackii 15 

III. Chaetanthera 16 

Key to Peruvian Species of 

Chaetanthera 16 

1 . Chaetanthera chiquian- 

ensis 17 

2. Chaetanthera cochleari- 

folia 17 

3. Chaetanthera peruviana 19 

4. Chaetanthera stuebelii 19 

IV. Chaptalia 19 

Key to Peruvian Species of 

Chaptalia 20 

1 . Chaptalia callacallensis 20 

2. Chaptalia cordata 21 

3. Chaptalia exscapa 21 

3a. Chaptalia exscapa var. 

microcephala 21 

4. Chaptalia integerrima 22 

5. Chaptalia isernina 22 

6. Chaptalia malcabalensis 22 

7. Chaptalia nutans 23 

8. Chaptalia oblonga 24 

9. Chaptalia rotundifolia 24 

10. Chaptalia similis 25 

V. Chucoa 25 

1. Chucoa ilicifolia 25 

VI. Chuquiraga 27 



Key to Peruvian Species of 

Chuquiraga 27 

1. Chuquiraga johnstonii 28 

2. Chuquiraga jussieui 28 

3. Chuquiraga rotundifolia 29 

4. Chuquiraga spinosa 29 

5. Chuquiraga weberbaueri 31 

VII. Dasyphyllum 31 

Key to Peruvian Species of 

Dasyphyllum 32 

1. Dasyphyllum brasiliense 32 

la. Dasyphyllum brasiliense 

var. barnadesioides 32 

2. Dasyphyllum brevispinum .... 34 

3. Dasyphyllum cabrerae 34 

4. Dasyphyllum ferox 34 

5. Dasyphyllum horridum 35 

6. Dasyphyllum hystrix 35 

Key to Peruvian Varieties of Dasy- 
phyllum hystrix 35 

6a. Dasyphyllum hystrix var. 

hystrix 35 

6b. Dasyphyllum hystrix var. 

peruvianum 36 

7. Dasyphyllum leiocephalum ... 36 

8. Dasyphyllum weberbaueri .... 36 
VIII. Fulcaldea 36 

1. Fulcaldea laurifolia 37 

IX. Gerbera 37 

1. Gerbera jamesonii 39 

X. Gochnatia 39 

Key to Peruvian Species of 

Gochnatia 40 

1 . Gochnatia arequipensis 40 

2. Gochnatia patazina 40 

3. Gochnatia vargasii 41 

4. Gochnatia vernonioides 41 

XI. Jungia 41 

Key to Peruvian Species of 

Jungia 43 

1. Jungia amplistipula 43 

2. Jungia axillaris 44 

3. Jungia discolor 44 

4. Jungia floribunda 45 

5. Jungia paniculata 45 

6. Jungia rugosa 46 

7. Jungia spectabilis 47 

8. Jungia stuebelii 47 

9. Jungia vitocensis 48 

10. Jungia weberbaueri 48 

XII. Leucheria 50 

1 . Leucheria daucifolia 50 

XIII. Lycoseris 51 

Key to Peruvian Species of 



in 



Lycoseris 51 

1 . Lycoseris peruviana 51 

2. Lycoseris trinervis 52 

XIV. Mutisia 54 

Key to Peruvian Species of 

Mutisia 54 

1. Mutisia acuminata 55 

Key to Peruvian Varieties of 

Mutisia acuminata 55 

la. Mutisia acuminata var. 

acuminata 56 

Ib. Mutisia acuminata var. 

bicolor 56 

Ic. Mutisia acuminata var. 

hirsuta 56 

2. Mutisia alata 57 

3. Mutisia andersonii 57 

4. Mutisia arequipensis 57 

5. Mutisia cochabambensis 58 

6. Mutisia hastata 58 

7. Mutisia lanata 59 

8. Mutisia lanigera 59 

9. Mutisia ledifolia 59 

10. Mutisia mathewsii 60 

1 1 . Mutisia orbignyana 60 

12. Mutisia pulcherrima 61 

13. Mutisia rauhii 61 

14. Mutisia venusta 61 

1 5. Mutisia wurdackii 63 

XV. Onoseris 63 

Key to Peruvian Species of 

Onoseris 64 

1. Onoseris acerifolia 65 

2. Onoseris albicans 65 

3. Onoseris amplexicaulis 66 

4. Onoseris annua 67 

5. Onoseris cabrerae 67 

6. Onoseris castelnaeana 68 

7. Onoseris chrysactinioides 68 

8. Onoseris cumingii 68 

9. Onoseris gnaphalioides 69 

10. Onoseris humboldtiana 69 

1 1 . Onoseris linearifolia 70 

12. Onoseris lopezii 70 

13. Onoseris macbridei 71 

14. Onoseris minima 72 

15. Onoseris odorata 72 

16. Onoseris peruviana 73 

1 7. Onoseris speciosa 74 

18. Onoseris weberbaueri 74 

XVI. Perezia 75 

Key to Peruvian Species of 

Perezia 76 

1. Perezia ciliosa . .76 



2. Perezia coerulescens 76 

3. Perezia multijlora 77 

4. Perezia pinnatifida 78 

5. Perezia pungens 79 

6. Perezia pygmaea 81 

7. Perezia sublyrata 81 

XVII. Plazia 82 

Key to Peruvian Species of 

Plazia 82 

1. Plazia conferta 83 

2. Plazia daphnoides 83 

XVIII. Polyachyrus 83 

Key to Peruvian Species of 

Polyachyrus 85 

1. Polyachyrus annuus 85 

2. Polyachyrus fuscus 85 

3. Polyachyrus sphaero- 

cephalus 87 

XIX. Proustia 87 

Key to Peruvian Species of 

Proustia 88 

1. Proustia berberidifolia 88 

2. Proustia cuneata 88 

3. Proustia cuneifolia 89 

4. Proustia foliosa 90 

5. Proustia peruviana 90 

XX. Trichocline 91 

Key to Peruvian Species of 

Trichocline 91 

1 . Trichocline caulescens 91 

2. Trichocline peruviana 92 

XXI. Trixis 93 

Key to Peruvian Species of 

Trixis 94 

1 . Trixis cacalioides 94 

2. Trixis churinensis 95 

3. Trixis divaricata 95 

4. Trixis neaeana 96 

5. Trixis sagasteguii 96 

6. Trixis subparadoxa 96 

ACKNOWLEDGMENTS 98 

MAP OF PERU 99 

INDEX TO LATIN NAMES . .100 



List of Illustrations 



1. Arnaldoa weberbaueri 6 

2. Barnadesia berberoides 9 

3. Chaetanthera cochlearifolia 18 

4. Chaptalia malcabalensis 23 

5. Chucoa ilicifolia 26 



IV 



6. Chuquiraga rotundifolia 30 14. Onoseris lopezii 71 

7. Dasyphyllum brasiliense 33 15. Perezia pungens 80 

8. Fulcaldea laurifolia 38 16. Plazia daphnoides 84 

9. Gochnatia vernonioides 42 17. Polyachyrus sphaerocephalus 86 

10. Jungia weberbaueri 49 18. Proustia cuneifolia 89 

1 1. Leucheria daucifolia 50 19. Trichocline caulescens 92 

12. Lycoseris trinervis 53 20. Trixis sagasteguii 97 

13. Mutisia venusta . . 62 



Back cover: Leucheria daucifolia 



FLORA OF PERU 
Family Asteraceae: Part VI 



Abstract 

The tribe Mutisieae (Asteraceae) is treated for 
Peru and includes 21 genera, 121 species, and 7 
varieties. Keys to genera, species, and varieties, 
descriptions, and illustrations are provided for Ar- 
naldoa (3 spp.), Barnadesia (\3 spp.), Chaetanthe- 
ra (4 spp.), Chaptalia (10 spp.), Chucoa (1 sp.), 
Chuquiraga (5 spp.), Dasyphyllum (8 spp.), Ful- 
caldea (1 sp.), Gerbera (1 sp.), Gochnatia (4 spp.), 
Jungia (10 spp.), Leucheria (1 sp.), Lycoseris (2 
spp.), Mutisia ( 1 5 spp.), Onoseris (18 spp.), Perezia 
(7 spp.), Plazia (2 spp.), Polyachyrus (3 spp.), 
Proustia (5 spp.), Trichocline (2 spp.), and Trixis 
(6 spp.). The generic limits of Proustia are here 
expanded to include species previously accepted 
in Lophopappus and require the following new 
combinations: P. berberidifolia (Cabr.) Ferreyra, 
P. foliosa (Rusby) Ferreyra, and P. peruviana 
(Cabr.) Ferreyra. Lectotypification is provided for 
Dasyphyllum ferox (Wedd.) Cabr. and Plazia 
daphnoides Wedd. 



Tribe MUTISIEAE 

Mutisieae Cass., J. Phys. Chem. Hist. Nat. Arts 
88: 199. 1819. TYPE: Mutisia L. fil. 

Annual or perennial herbs, lianas, shrubs, or 
small trees. Leaves alternate and cauline, or con- 
fined to a basal rosette, rarely opposite. Capitu- 
lescences of solitary heads, sessile, pedunculate, or 
scapose, or several-headed panicles or cymes, oc- 
casionally glomerulate in pseudocephalia or syn- 
florescences. Capitula homogamous, discoid, or 
heterogamous, radiate to disciform; involucres 
globose to cylindric; receptacles flat or convex, 
naked (paleate in Jungia, Arnaldod), foveolate, or 
hispid-fimbrillate or glandular pubescent; phylla- 
ries (l-)2-12(-14)-seriate, usually unequal, usu- 
ally imbricate; florets iso-, di-, or trimorphic, the 
corollas bilabiate, the outer lip ligulate, 3-4-lobed 
to 3 4-dentate, the interior lip single or of 2 fused 
or unfused lobes; or actinomorphic, tubular, the 
limb 5-lobed, the lobes equal to subequal; anthers 



with truncate to pointed terminal appendage, the 
bases usually caudate with long tails (ecaudate in 
Barnadesia), the tails sometimes fringed at the tips; 
style branches short, truncate or obtuse to rounded 
or acute, penicillate, papillose, or glabrous. Achenes 
columnar, turbinate, or fusiform, ribbed to costate 
or smooth, sericeous, glandular, or glabrous; pap- 
pus of bristles, usually 1-2-seriate, scabrid, bar- 
bellate, or plumose, or occasionally scalelike, flat- 
tened, or setaceous, glabrous to hirtellous. 

Mutisieae (sensu latd) contains 89 genera with 
over 975 recorded species in tropical and sub- 
tropical America, Africa, Asia, Australia, and Ha- 
waii (Cabrera, 1977). It is predominantly a Neo- 
tropical tribe with greatest generic representation 
in South America, where no fewer than 65 of its 
genera occur. The tribe contains endemic genera 
in several other areas for example, North Amer- 
ica (5 genera), Asia (5 genera), Australia ( 1 genus), 
Africa (10 genera), and several Pacific Islands (1 
genus). Western South America is a center of high 
generic diversity; the Andean Cordillera contains 
over 30 genera with distributions generally above 
2000 m. Peru is particularly rich with 21 genera 
and over 120 species distributed in various eco- 
systems of the Coast, Sierra, and Selva. Species 
diversity and distribution patterns are quite vari- 
able; intermontane valleys and the ceja de la mon- 
tana are richest in taxa. Fewer than six species are 
found in the more humid tropics of the Amazon 
basin of eastern Peru (e.g., Barnadesia caryophyl- 
la, Chaptalia nutans, Lycoseris spp., Trixis divar- 
icatd). Gerbera is predominantly an Asian and Af- 
rican genus, with only one species cultivated in 
Peru. Diversity within genera is also variable; the 
monotypic genus Chucoa is endemic to a single 
valley in northern Peru, whereas Onoseris and 
Trixis each occur in both hemispheres from the 
United States to the Patagonian region of Argen- 
tina and Chile. Some pairs of genera and species 
exhibit overall morphological convergence. Ful- 
caldea and Barnadesia, Chuquiraga and Dasy- 
phyllum, and even Chuquiraga spinosa and Chu- 
quiraga johnstonii are quite similar as herbarium 
specimens but easily distinguishable in the field. 
The color of the corolla is an important character, 



FIELDIANA: BOTANY, N.S., NO. 35, MAY 31, 1995, PP. 1-101 



particularly within species with close resem- 
blancefor example, both Mutisia pulcherrima 
and M. wurdackii are quite similar lianas, but the 
former has orange corollas and the latter red. 

References 

CABRERA, A. L. 1977. Mutisieae Systematic re- 
view, pp. 141-248. In Heywood, V. H., et al., 
eds., The Biology and Chemistry of the Com- 
positae. Academic Press, London. 

CRISCI, J. V. 1974. A numerical-taxonomic study 
of the subtribe Nassau viinae (Compositae, Mu- 
tisieae). J. Arnold Arb, 55: 568-610. 



GRAU, J. 1980. Die testa der Mutisieae und ihre 
systematische bedeutung. Mitt. Bot. Munchen, 
16: 269-332. 

HANSEN, H. V. 1990. Phylogenetic studies in 
Compositae tribe Mutisieae. Opera Bot., 109: 
1-50. 

. 1991. SEM-studies and general com- 
ments on pollen in tribe Mutisieae (Compositae) 
sensu Cabrera. Nord. J. Bot., 10: 607-623. 

HARLING, G. 1991 190(10). Compositae Mu- 
tisieae, pp. 1-106. In Harling, G., and Anders- 
son, L., eds., Flora of Ecuador, No. 42. Swedish 
Natural Science Research Council, Stockholm. 



Key to Genera of Mutisieae in Peru 

The following artificial generic key is adapted from that presented by Cabrera (1977) and will allow 
determination of representatives treated here but may not function for taxa outside of Peru. 

1. Disc florets more or less actinomorphic, deeply 5-lobed or 5 -cleft, with equal or unequal segments, 
occasionally weakly zygomorphic with single lobe opposite a 4-lobed segment, never truly bilabiate 

(i.e., 3 lobes opposite 2 lobes); ray florets, when present, bilabiate or subligulate 2 

2. Disc florets with corollas usually villous; pappus of plumose bristles (setaceous to barbellate in 
Barnadesia); axillary or nodal spines frequently present (rarely reduced or absent, e.g., Chuquiraga 

or Dasyphyllum spp.) [Barnadesiinae] 3 

3. Capitula with 1 disc floret and lacking ray florets Fulcaldea 

3. Capitula with 2 or more disc florets; if fewer than 2 disc florets then accompanied by 8 or 

more ray florets (cf. Barnadesia macbridei) 4 

4. Capitula heterogamous; anthers ecaudate and obtuse to rounded basally .... Barnadesia 

4. Capitula heterogamous; anthers caudate to sagittate at base 5 

5. Corollas zygomorphic, the limb deeply divided (4 lobes/ 1 lobe) or subligulate; involucres 

25-60 mm high; phyllaries 10- 13 -seriate; florets 50-150; corollas orange to red 

Arnaldoa 

5. Corollas actinomorphic, the limb equally to subequally 5-lobed; involucres 25-35(-50) 
mm high; phyllaries 5-8(-12)-seriate; florets 10-50; corollas yellow to orange or white 

to violet 6 

6. Leaves 1 -nerved; corollas yellow to orange; anthers long-tailed, apical appendages 

entire, linear- lanceolate, apically acute to obtuse Chuquiraga 

6. Leaves 3-5-nerved; corollas white to violet; anthers short-tailed, apical appendages 

usually bifid, occasionally emarginate or truncate Dasyphyllum 

2. Disc florets with corollas usually glabrous or at most puberulent; pappus of bristles, scabrous to 

barbellate, never truly plumose; branches lacking axillary or nodal spines [Gochnatiinae] 7 

7. Plants dioecious; i.e., capitula functionally unisexual, radiate Lycoseris 

7. Plants monoecious; i.e., capitula functionally bisexual, discoid or radiate 8 

8. Disc florets actinomorphic or slightly zygomorphic, the corollas 5-lobed or 5-dentate, the 

lobes or teeth less than '/3 the length of the entire corolla 9 

9. Shrubs; leaf blades oblanceolate, the margins strongly dentate, the 4-6 pairs of teeth 
spinulose; capitula homogamous, discoid; florets with yellow corollas; achenes glabrous 

or papillose Chucoa 

9. Annual or perennial herbs, subshrubs, shrubs, lianas; leaf blades linear to cordiform, 
the margins entire to incised or lyrate-lobed, never with 4-6 pairs of spinulose teeth; 
capitula heterogamous, radiate; disc florets with yellow corollas; ray florets with purple 
to violet corollas; achenes pubescent, glabresenct, or rarely glabrous Onoseris 



FIELDIANA: BOTANY 



8. Disc florets actinomorphic, the corollas deeply 5-lobed, the lobes linear and greater than 

Vs the length of the corolla 10 

10. Leaves in dense whorls at branch apices; involucres 18-22 mm high; achenes glabrous 

or glandular pubescent Plazia 

10. Leaves evenly spaced on stems, not densely clustered at apices; involucres 8-12 mm; 

achenes densely sericeous or glandular Gochnatia 

1 . Disc florets zygomorphic, the corollas clearly bilabiate with the outer lip 3-dentate and the interior 
lip simple or bifid (for exceptions see Proust ia), or ligulate with 5 -dentate ligule; ray florets bilabiate, 

ligulate 11 

1 1 . Style branches apically acute to rounded, dorsally glabrous or papillose but lacking conspicuous 

ring of collecting trichomes [Mutisiinae] 12 

1 2. Plants perennial herbs; leaves oval to lanceolate or lyrate-pinnatifid in basal rosette; capi- 

tulescences scapose; capitula borne singly (rarely 2-3) on elongate peduncles 13 

1 3. Capitula nutant or rarely erect; florets typically trimorphic; ray florets equalling or rarely 

exceeding the involucre, staminodia absent Chaptalia 

13. Capitula erect; florets dimorphic; ray florets clearly exceeding involucre, staminodia 

present 14 

14. Peduncles bracteate, tomentose; achenial pubescence of 2 types: biseriate trichomes 
(Zwillingshaare), the hair shaft typically oblong to spathulate, often appearing in- 
flated, the apices rounded to obtuse, and biseriate, multicellular glandular 

trichomes Trichocline 

14. Peduncles ebracteate, villous; achenial pubescence of only 1 type: biseriate tri- 
chomes (Zwillingshaare), the hair shaft typically linear-elongate and tapering, the 
apices attenuate to acute, lacking biseriate, multicellular glandular trichomes .... 

Gerbera 

1 2. Plants subshrubs, lianas, or perennial or annual herbs; leaves not disposed in basal rosettes; 

capitulescences axillary and terminal, never scapose 15 

15. Small annual or perennial herbs (rarely subshrubs); leaves simple, linear to spathulate, 

never with tendril at apex Chaetanthera 

1 5. Shrubs or lianas; leaves generally pinnately compound, if simple then usually possessing 

apical tendril Mutisia 

1 1 . Style branches apically truncate (rounded in Proustia), usually with a crown of collecting tri- 
chomes (penicillate) [Nassauviinae] ' 16 

16. Leaf blades oblong-lanceolate to orbicular-cordate; receptacles paleate; corollas usually white, 

yellow, or occasionally pink, lilac, or violaceous Jungia 

16. Leaf blades variously linear, lanceolate, oblong, ovate, or lyrate-pinnatifid, never truly or- 
bicular-cordate in outline; receptacles fimbriate or naked; corollas yellow or orange, blue, 

purple or violet, or rarely white to pink 17 

17. Capitulescences glomerulate in pseudocephalia or synflorescences of 9 or more heads 

Polyachyrus 

17. Capitulescences of solitary heads, or cymes to panicles, never glomerulate in synflo- 
rescences 18 

18. Shrubs or vines; corollas yellow, whitish yellow, or yellow-orange Trixis 

18. Shrubs without yellow florets, or perennial or annual herbs with corollas white, 

blue, lavender, or violet 19 

19. Shrubs or subshrubs, occasionally with infrapetiolar spines; leaves simple, the 
margins entire to denticulate; capitulescences solitary or 2-4 heads at apex of 
branchlets, or panicles or racemes of spikes; capitula with 1 5 or fewer florets; 
corollas white to lavender Proustia (including Lophopappus) 



1 Proustia (including Lophopappus) is included within the Nassauviinae on the basis of pollen type and is a notable 
exception in possessing actinomorphic disc florets and rounded, dorsally pubescent style branches (Cabrera, 1977). 



FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI. 



19. Perennial herbs, caulescent or scapose; leaves typically in a basal rosette, the 
margins entire, lobed, or pinnatisect; capitulescences cymose-corymbose or, 

rarely, solitary; capitula with 15-150 florets; corollas white to blue 20 

20. Corollas white; pappus of plumose bristles Leucheria 

20. Corollas violet to blue; pappus of scabrid or barbellate bristles 

. Perezia 



I. ARNALDOA 

Arnaldoa Cabr., Bol. Soc. Argent. Hot. 10: 39. 1962. 
TYPE: Arnaldoa magnifica Cabr. = Arnaldoa 
weberbaueri (Muschl.) Ferreyra 

Shrubs to 5 m tall, much-branched, the branch- 
es usually spiny at least below, the nodal spines 
geminate, divergent, fuscous. Leaves alternate, 
petiolate; blades oblong- to ovate-elliptic, coria- 
ceous, the abaxial surfaces with midrib prominent, 
penninerved with 2-3 pairs of secondary veins, 
the margins entire. Capitulescences of solitary, ter- 
minal heads. Capitula homogamous, discoid, ses- 
sile; involucres broadly campanulate; receptacles 
plane, paleate, the pales yellowish; phyllaries 1 0- 
13-seriate, acuminate; florets isomorphic, her- 
maphroditic, the corollas orange or red, bilabiate, 
the outer lip liguliform, 5-nerved, apically 4-lobed, 
the inner lip filiform; anthers sagittate; styles cy- 
lindric, 30-32 mm long, the branches small, api- 
cally acute. Achenes turbinate or cylindric; pappus 
of plumose bristles, yellowish. 



Arnaldoa consists of three species, all confined 
to central and northern Peru. The genus is most 
easily recognized by its large, homogamous capit- 
ula with zygomorphic corollas. It shares relation- 
ships with the other members of the subtribe Bar- 
nadesiinae, including Barnadesia, Chuquiraga, and 
Dasyphyllum. 



References 

CABRERA, A. L. 1962. Compuestas Andinas 
Nuevas. Bol. Soc. Argent. Bot, 10: 39. 

FERREYRA, R. 1965. Las especies de Arnaldoa 
del Peru (Compositae). Publ. Mus. Hist. Nat. 
"Javier Prado," Ser. B, Bot., 19: 1-8. 

. 1 980. Notas sobre especies Peruanas de 

los generos Arnaldoa, Chucoa (Compositae) y 
Monnina (Polygalaceae). Bol. Soc. Peruana Bot., 
8: 106-109. 



Key to Species of Arnaldoa 

1. Branches hirsute or tomentose; leaves pubescent; corollas orange A. weberbaueri 

1 . Branches glabrous; leaves glabrescent; corollas red 2 

2. Upper branches spiny, spines ca. 38 mm long; apex of leaves spinulose; capitula to 60 mm . . . 

A. coccinosantha 

2. Upper branches usually without spines, rarely with few spines to 22 mm long; apex of leaves 
obtuse mucronate or emarginate; capitula ca. 30 mm A. macbrideana 



1. Arnaldoa coccinosantha (Muschl.) Ferreyra, 
Publ. Mus. Hist. Nat. "Javier Prado," Ser. B, 
Bot. 19:4. 1965. 



Barnadesia coccinosantha Muschl., Bot. Jahrb. Syst. 
50, Beibl. Ill: 98. 1913. TYPE: Peru, Ancash, 
Huari, Chavin de Huantar, A. Weberbauer 3295 
(holotype, B, presumably destroyed; isotype, USM; 
isotype fragment ex w, F, photograph ex B, FM 
neg. 15908; photograph ex w, FM neg. 52100). 

Chuquiraga coccinosantha (Muschl.) Chung, Rev. 
Barnadesia 7. 1965. 



Arnaldoa peruviana Lopez & Sagast.. Revista Fac. Ci. 
Biol. Univ. Nac. Trujillo 1(1): 12. 1964. TYPE: 
Peru, Dept. La Libertad, Pataz, entre Huaylillas 
y Tayabamba, A. Lopez M. & A. Sagdstegui A. 
3410 (holotype, HUT). 

Shrubs 1 5-20 dm high, the branches striate, gla- 
brescent, spiny, the spines to 35 mm long. Leaves 
with petioles 2-5 mm long, sericeous; blades ob- 
long to ovate-elliptic, 30-50 mm long, 12-32 mm 
wide, obtuse, mucronate. Capitula with involucres 
50-60 mm high, 30-38 mm wide, campanulate; 



FIELDIANA: BOTANY 



phyllaries 1 1-12-seriate, the inner linear, 45-50 
mm long, 1.8-3 mm wide, attenuate at apex, se- 
riceous outside, the outer ovate triangular, 8-10 
mm long, 3.54 mm wide, acute, lanuginous out- 
side; florets ca. 145, the corollas red, 45-50 mm 
long, the tube 6-8 mm long, the outer lip 3542 
mm long, 3.8-5 mm wide, pubescent outside, the 
inner lip 35-40 mm long; anthers 9-9.5 mm long. 
Achenes cylindric, 9-11 mm long, 1.8-2.2 mm 
wide, pubescent, costate; pappus ca. 30 mm long. 

DISTRIBUTION Native to the intermontane re- 
gion of northern Peru (2350-2700 m). 

Arnaldoa coccinosantha is apparently quite rare 
and weakly distinguished from A. weberbaueri by 
its more oblong leaves and deep red corollas. It is 
quite distinct from A. macbrideana, with much 
larger capitula and leaves than that species, but it 
does share that species' reddish corollas. 

LA LIBERTAD. Pataz: Chagual, Huaquil-Tayabam- 
ba, Lopez & Sagdstegui 8159 (GH, MO); Huancaspata- 
Mamahuaje, Lopez & Sagdstegui 8218 (GH, MO). 

2. Arnaldoa macbrideana Ferreyra, Publ. Mus. 
Hist. Nat. "Javier Prado," Ser. B, Bot. 19: 6. 
1965. TYPE: Peru, Piura, Huancabamba, near 
Abra Porculla, R. Ferreyra & J. J. Wurdack 
14415 (holotype, USM). 

Dasyphyllum barbatum Chung, Phytologia 14: 321. 
1967. TYPE: Peru, Dept. Lima, Botanical Gar- 
den, cultivated, 7 Aug 1945, R. J. Seibert 2096 
(holotype, us 1 90 1 49 1 ; photograph ex us, FM neg. 
52103). 

Shrubs 2-5 m high, the branches sericeous, spiny, 
the spines to 25 mm long. Leaves with petioles 2- 
6 mm long, pubescent; blades oblong-elliptic, 22- 
55 mm long, 1842 mm wide, obtuse, mucronate, 
rarely emarginate, sericeous. Capitula with invo- 
lucres 25-30 mm high, 14-18 mm wide, campan- 
ulate; phyllaries 10-1 1 -seriate, the inner linear, 
22-25 mm long, 1.2-1.8 mm wide, acute, gla- 
brescent, the margins pubescent, the outer trian- 
gular, 6-6.5 mm long, 4.54.8 mm wide, acute, 
mucronate; florets ca. 68, the corollas red-purple, 
1 9-24 mm long, the tube 6-9 mm long, the outer 
lip 12-14.5 mm long, 3-3.2 mm wide, acuminate, 
the inner lip 1 0- 1 5 mm long; anthers 1 0- 1 0. 5 mm 
long. Achenes turbinate, 6.5-7 mm long, pubes- 
cent; pappus ca. 1 6 mm long. 

DISTRIBUTION Endemic to the intermontane 
region of northern Peru (1000-2200 m). 



Arnaldoa macbrideana is distinguished by its 
smaller capitula with generally fewer florets (usu- 
ally fewer than 70 per head) and virtually glabrous 
leaves. 



LAMBAYEQUE. Lambayeque: Olmos-Porculla, El- 
lenberg 3723 (us), 5725 (us); W side Abra Porculla Pass 
km 35 E of Olmos, Hutchison 1379 (F, uc, us, USM); 
between Olmos and Abra Porculla, Plowman 5516 (us); 
road from Olmos to Pucara, km 32 E of Olmos, Plowman 
et al. 14202 (F); Abra de Porculla, km 45 E of Olmos, 
Plowman et al. 14266 (F); Sagdstegui et al. 7140 (GH). 
PIURA. Huancabamba: near Abra Porculla, Ferreyra 
13628 (USM), 13731 (USM). NO EXACT LOCALITY. 
Peru, "inter Bagua et Olmos," Maekawa 602307 (F 
1615406). 

3. Arnaldoa weberbaueri (Muschl.) Ferreyra, Publ. 
Mus. Hist. Nat. "Javier Prado," Ser. B, Bot. 
19: 2. 1965. Figure 1. 

Barnadesia weberbaueri Muschl., Bot. Jahrb. Syst. 50, 
Beibl. Ill: 97. 1913. TYPE: Peru, Amazonas, 
Chachapoyas, prope Balsas, 25 Jun 1904, 2000- 
2100 m, A. Weberbauer 4278 (holotype, B, pre- 
sumably destroyed; isotype, GH; photograph ex B, 
FM neg. 15920). 

Chuquiraga muschleri Chung, Rev. Barnadesia 8. 1 965. 
NEOTYPE (designated by Chung, 1965): Peru, 
Amazonas, "Balza" [Balsas], May 1877, Vidal- 
Seneges.n. (neotype, P, not seen; isotype fragment 
ex p, F, photograph ex p, FM neg. 052101), illeg. 
nom. nov. for B. weberbaueri Muschl. 

Arnaldoa magnifica Cabr., Bol. Soc. Argent. Bot. 10: 
39. 1962. TYPE: Peru, Cajamarca, Celendin, La 
Lina, Celendin-Maranon, A. Lopez M. & A. Sa- 
gdstegui A. 3372 (holotype, LP; isotype, HUT). 



Shrubs 1 5-20 dm high, the branches striate, hir- 
sute-tomentose, glabrescent, spiny, the spines to 
25 mm long. Leaves with petioles 3-6 mm long, 
lanuginous; blades ovate-elliptic, 32-62 mm long, 
2040 mm wide, apex obtuse, mucronate, seri- 
ceous or lanuginous. Capitula with involucres 45- 
55 mm high, 24-30 mm wide, campanulate; phyl- 
laries 12-13-seriate, the inner linear, 41-43 mm 
long, 1.5-2 mm wide, acuminate, recurved, pu- 
bescent outside, trinerved, the outer triangular, 8- 
9 mm long, 34 mm wide, acuminate, mucronate 
or spinulose, densely pubescent outside; florets ca. 
91, the corollas orange, 32-33 mm long, the tube 
3.54 mm long, 1.8-2 mm wide, glabrous, the 
outer lip of the limb 27-30 mm long, 4-5 mm 
wide, acute, pubescent outside, the inner lip 27- 
28 mm long, 1-1.2 mm wide; anthers 14-18 mm 
long. Achenes cylindric, 8-1 1 mm long, 2-2.2 mm 
wide, pubescent; pappus ca. 22 mm long. 



FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI. 




5mm 



FIG. 1. Arnaldoa weberbaueri. 1, habit; 2 floret; 3, achene with pappus; 4, anther; 5, style apex. (Drawn from D. 
N. Smith & I. Sanchez V. 4323, F.) 



FIELDIANA: BOTANY 



DISTRIBUTION Endemic to both sides of the 
valley of the Upper Rio Maranon (1900-2400 m), 
where it grows on the steep slopes with xerophytic 
vegetation. Arnaldoa weberbaueri is distinguished 
by its densely sericeous or lanuginous leaves, large 
capitula, recurved phyllaries, and conspicuously 
orange corollas. 

AMAZONAS. Chachapoyas: Balsas-Leimebamba, 
Duncan 2602 (F, MO); western base of the Cerro Calla 
Calla, 22 km E of Balsas on the road to Leimebamba, 
Hutchison 5454 (F, us, USM). CAJAMARCA. Celendin: 
above Hacienda El Limon, Ferreyra 13322 (USM), 15068 
(us, USM); Hacienda Limon, 10 mi W of Balsas, Osgood 
& Anderson 62 (F); Hda. El Limon, Sagdstegui 7410 (F, 
us); Huasho, entre Gelig-Balsas, Sanchez 339 (CPUN, F). 



II. BARNADESIA 

Barnadesia Mutis ex L. fil., Suppl. pi. 55, 348. 
1782. TYPE: B. spinosa L. fil. 

Shrubs or small trees, the branches usually with 
geminate or fasciculate nodes. Leaves alternate, 
often fasciculate in groups of 2-8, subsessile to 
petiolate; blades lanceolate to oblong-elliptic, pen- 
ninerved, glabrous to sericeous, the margins en- 
tire. Capitulescences of solitary heads, terminal or 
axillary, rarely cymose or corymbose, sessile or 
pedunculate. Captitula heterogamous, radiate; in- 
volucres campanulate to cylindric; receptacles 
plane, epaleate, fimbriate to villous; phyllaries 7- 
1 4-seriate, imbricate, linear or lanceolate to ovate, 
plane to concave, apically acute, acuminate, or 
long-attenuate; ray florets pistillate, 8-13, 1 -seriate, 
the corollas red to purple, occasionally pink or 
white, bilabiate with outer lip liguliform, 5-nerved, 
apically 4-toothed, the inner lip filiform, entire, 
glabrous to pilose, the tube usually pubescent to 
villous; stames 5, filaments free, or connate and 
forming a fused membranous tube, the anthers 
ecaudate, basally obtuse or rounded, the apical 
appendages linear-lanceolate, apically obtuse; styles 



claviform, the branches short, rounded, papillate; 
disc florets hermaphroditic, 1-3, the corollas tu- 
bular, the lim 4-5-toothed, the teeth equal to un- 
equal, rarely bilabiate, the outer lip ligulate, 
4-toothed, the inner lip simple; stamens 4-5, the 
anthers ecaudate, basally obtuse or rounded, the 
apical appendages linear-lanceolate, apically ob- 
tuse; styles claviform, the branches short, round- 
ed, papillate. Achenes [ray] cylindric to turbinate, 
subterete to flattened, villous; pappus of bristles, 
plumose to barbellate, persistent, yellow to brown- 
ish-yellow; [disc] turbinate, densely villous; pap- 
pus of setae, yellowish, rigid, coiled to recurved, 
subulate, glabrous to hirtellous, rarely of barbellate 
bristles. Chromosome number: n = 25. 

Barnadesia consists of approximately 20 species 
confined to the Andean Cordillera from Colombia 
to northern Argentina and southern Brazil. The 
genus records it greatest diversity in Peru with 1 4 
species distributed from the Cordillera Occidental 
to the ceja de la montana of the Cordillera Oriental 
(6004100 m). Only one species, Barnadesia car- 
yophylla, has a distribution at lower elevation on 
the eastern escarpment and into Brazil. 

References 

CABRERA, A. L. 1 960. Notas sobre tipos de Com- 
puestas Sudamericanas en herbarios Europeos. 
HI, Los tipos de Ruiz y Pavon. Bol. Soc. Argent. 
Bot., 8: 195-215. 

CHUNG, I. 1965. Revision of Barnadesia (Com- 
positae Mutisieae), pp. 1-86. Chicago, 111. (pri- 
vate edition). 

FERREYRA, R. 1964. Revision de las especies 
Peruanas del genero Barnadesia (Compositae). 
Publ. Mus. Hist. Nat. "Javier Prado," Ser. B, 
Bot., 18: 1-35. 

ROBINSON, H. 1 975. A new species of Barnadesia 
from Ecuador. Phytologia, 32: 414-417. 



Key to Peruvian Species of Barnadesia 

1 . Stamen filaments of the ray florets free and distinct, 1-5 mm long 2 

2. Ray florets 8; disc floret 1 3 

3. Capitula 40-55 mm high, 10-12 mm wide B. berberoides 

3. Capitula 25-30 mm high, 5-8 mm wide 4 

4. Leaf blades linear to lanceolate, 15-25 mm long, 3-5 mm wide B. horrida 

4. Leaf blades oblong-elliptic, 15-60 mm long, 1 1-21 mm wide B. macbridei 

2. Ray florets 1 3; disc florets 3 5 

5. Phyllaries 6-8-seriate B. lehmannii 



FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI. 



5. Phyllaries 10-14-seriate 6 

6. Phyllaries 1 3-14-seriate; inner phyllaries oblong- lanceolate, ca. 30 mm long, ca. 3 mm wide 

B. hutchisoniana 

6. Phyllaries 10-13-seriate; inner phyllaries linear, 30-40 mm long, 1.2-2 mm wide 7 

7. Leaf blades elliptic to obovate-lanceolate, 30-80 mm long, 1 1-34 mm wide; phyllaries 

1 1-1 3-seriate B. caryophylla 

1. Leaf blades oblong-elliptic, 12-35 mm long, 5-14 mm wide; phyllaries 10- 11 -seriate 

B. polyacantha 

1. Stamen filaments of the ray florets connate and forming a fused membranous tube, 3-13 mm long 

8 

8. Ray florets 8; disc floret 1 9 

9. Trees; leaves clearly alternate; petioles 5-30 mm long; blades (35-)60-140 mm long; capitu- 
lescences corymbose, 10-50-headed B. corymbosa 

9. Shrubs; leaves fasciculate; petioles 5-7 mm long or lacking; blades 7-28 mm long; capitules- 
cences solitary or in 3-5-headed clusters B. jelskii 

8. Ray florets 1 3; disc florets 3 10 

10. Plants glabrous 11 

1 1 . Upper branches densely spiny, the spines ca. 30 mm long; leaves elliptic . B. blakeana 

11. Upper branches without spines, or rarely with few spines, 4-10 mm long; leaves oblan- 
ceolate B. wurdackii 

1 0. Plants tomentose or hirsute 12 

12. Spines 25-50 mm long; leaves with inconspicuous lateral nerves; phyllaries 5-9-seriate, 
recurved B. dombeyana 

12. Spines 10-18 mm long; leaves with conspicuous lateral nerves; phyllaries 9-12-seriate, 
erect . . B. reticulata 



1. Barnadesia berberoides Schultz-Bip. in Lechl., 
Berberid. Amer. Austral. 57. 1857. TYPE: 
Peru, Puno, Agapata, 1844, W. Lechler s.n. 
(holotype, B, presumably destroyed; photo- 
graph ex B, FM neg. 1 5907; isotype, K, not seen; 
photograph ex K, FM neg. 52068). Figure 2. 

Barnadesia macrocephala Kuntze, Revis. gen. pi. 3(2): 
135. 1898. TYPE: Bolivia, "Sierra Colomi," 1- 
4 Apr 1892, 3000 m, O. Kuntze s.n. (holotype, 
NY; photograph ex NY, FM neg. 52070). 

Barnadesia pycnophylla Muschl., Bot. Jahrb. Syst. 50, 
Beibl. Ill: 101. 1903. TYPE: Peru, Apurimac, 
valley of Pieni, A. Weberbauer 5558 (holotype, B, 
presumably destroyed; photograph ex B, FM neg. 
15917). 

Shrubs 2-6 m high, branched, the branches fus- 
cous, hirsute, glabrescent, spiny, the spines divar- 
icate or fasciculate, to 28 mm long. Leaves fascic- 
ulate, 3-5 per node, subsessile; blades lanceolate, 
25-42 mm long, 8-14 mm wide, apically acute, 
mucronate, both surfaces sericeous. Capitules- 
cences of solitary heads, terminal or axillary, ses- 
sile, occasionally weakly cymose with 2-3 heads. 
Capitula with involucres cylindric, 40-55 mm high, 
10-12 mm wide; phyllaries 9-1 1 -seriate, the outer 
ovate-triangular, apex strongly spinulose, re- 



curved, the inner linear, 2649 mm long, 1.8-2.2 
mm wide, acuminate, the abaxial surfaces seri- 
ceous; ray florets 8, the corollas pink-purple, 32- 
61 mm long, the tube 15-33 mm long, pubescent, 
the outer lip 1 5-24 mm long, 3-7 mm wide, the 
inner lip 10-1 1.5 mm long, filiform, flexuose; fil- 
aments free, the anthers 10-13 mm long; disc floret 
1, the corollas 1 1-17 mm long, the tube 3.5-4.2 
mm long, 3.84.5 mm wide, pubescent. Achenes 
[ray] turbinate, 3.5-5.5 mm long, 3-5.2 mm wide; 
pappus ca. 21 mm long; [disc] 3.2-5.5 mm long, 
1.4-2.2 mm wide; pappus ca. 25 mm long. 

DISTRIBUTION Native to intermontane valleys 
from central Peru to northern Bolivia (3000-3700 
m). 

Barnadesia berberoides is distinguished by its 
large heads, strongly spinose and recurved phyl- 
laries, and pink-purple corollas of the ray florets. 
Chung (1965) reduced Barnadesia berberoides to 
synonymy under B. pycnophylla; the former spe- 
cies was validly published, however, and is here 
accepted. 

APURIMAC. Abancay: Ampay, Stork 10615 (F). An- 
dahuaylas: between Pajonal and Andahuaylas, Hjerting 
& Petersen 1432 (USM). CUSCO. Anta: Sillapuyu, Brunei 



FIELDIANA: BOTANY 




FIG. 2. Barnadesia berberoides. 1, habit; 2, ray floret; 3, achene with pappus; 4, disc floret. (Drawn from J.F. 
Macbride & W. Featherstone 2230, F.) 



242 (MO). Claca: vicinity of Lares, Vargas 3607 (cuz, F); 
Pilco River, Woytkowski 344 (USM). Paucartambo: 
Acjanaco, Parque Nacional de Manu, Cano 3255 (F); 9- 
1 2 km NE of Paucartambo, Gentry et al. 23420 (F, MO); 
Paucartambo-Pilcopata road, 4 km below Tres Cruces, 



Gentry et al. 23493 (F); Cachupata, Vargas 2861 (F). 
Quispicanchis: Ocongate to Marcapata, Aronson & Berry 
509 (F, MO). Urubamba: Urubamba, Peyton 1541, 1563 
(MO). HUANCAVELICA. Tayacaja: Tranca-Pampa, 
between Huari and Acobamba, Tovar 4544 (USM); above 



FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI. 



Ampurco, Tovar 3795 (USM); 3 km N of Salcabamba, 
Stork & Morton 10319 (F). HUANUCO. Huanuco: 6 mi 

5 of Mito, Macbride & Featherstone 1830 (F, us); 15 mi 
SE of Huanuco, Macbride & Featherstone 2230 (F, us). 
JUNIN. Prov. unknown: "Tarhuish," Soukup 2883 (F, 
us). PUNO. Sandia: 2-6 km S of Limbani, Metcalf 30446 
(F). NO EXACT LOCALITY. Soukup 1225 (F). 

2. Barnadesia blakeana Ferreyra, Publ. Mus. Hist. 

Nat. "Javier Prado," Ser. B, Bot. 18: 14. 1964. 
TYPE: Peru, Lima, Huarochiri, Monte de 
Zarate, above San Bartolome, R. Ferreyra 
10427 (holotype, USM 15730; isotype, MO). 

Shrubs 15-20 dm high, branched, spiny, the 
spines ca. 30 mm long, divaricate, fuscous, diver- 
gent. Leaves fasciculate, 34 per node; petioles 1.5- 
2.5 mm long; blades lanceolate to elliptic, 12-32 
mm long, 7-13 mm wide, apically acute, strongly 
mucronate, glabrous. Capitulescences of solitary 
heads, terminal or axillary, sessile. Capitula with 
involucres 4045 mm high, 10-12 mm wide, cy- 
lindrical to campanulate; phyllaries 1 1-12-seriate, 
the outer ovate-lanceolate, 9.5-10 mm long, 5-5.5 
mm wide, the inner linear, 38-40 mm long, 1.6- 
2.2 mm wide, acute, the abaxial surfaces sericeous; 
ray florets 1 3, the corollas purple, 52-55 mm long, 
the tube 32-33 mm long, pubescent, the outer lip 
20-21 mm long, 3.84.2 mm wide, the abaxial 
surface pubescent, the inner lip 15-16 mm long; 
filaments connate, the anthers 10-11 mm long; 
disc florets 3, the corollas 1 8-2 1 mm long, the tube 
7.5-8 mm long, 3-3.2 mm wide, pubescent, api- 
cally 5-lobed, usually 4 equal, the inner lip fili- 
form; anthers 9-9.5 mm long. Achenes [ray] 5.5- 

6 mm long, 1.2-1.5 mm wide; pappus 21-23 mm 
long; [disc] 4.5-5 mm long, 3.5-4 mm wide, tur- 
binate; pappus 13-16 mm long. 

DISTRIBUTION Endemic to the type locality of 
Monte de Zarate, between Lima and Matucana 
(2900 m) in central Peru. 

Barnadesia blakeana most closely resembles B. 
dombeyana but differs from that species with its 
glabrous habit, leaves with strongly mucronate 
apices, and longer involucres. 

LIMA. Huarochiri: Monte de Zarate, Diers 1147 (USM); 
Hutchison & Wright 7091 (USM). 

3. Barnadesia caryophylla (Veil.) S. F. Blake, Proc. 

Biol. Soc. Wash. 38: 86. 1925. 

Xenophontia caryophylla Veil., Fl. Flum. 347, t. 85. 
(1825) x 1829. NEOTYPE (designated by Chung, 
1965): Brazil, Goyaz, 1841, G. Gardner 4268 



(neotype, K, not seen; photograph ex K, FM neg. 
52085; isoneotypes, F 1 1 3 1 97 ex G, F 974790 ex 
P). 

Shrubs 1 840 dm high, branched, hirsute, gla- 
brescent, spiny, the spines divaricate to 30 mm 
long, caducous on distal branchlets. Leaves fas- 
ciculate, 3-4 per node; petioles 3-8 mm long; blades 
elliptic to obovate-lanceolate, 30-80 mm long, 1 1- 
34 mm wide, apically acute, mucronate, the ad- 
axial surfaces glabrous, the abaxial surfaces seri- 
ceous. Capitulescences of usually solitary heads, 
terminal or axillary, sessile, or, occasionally weak- 
ly cymose with 2-3 heads. Capitula with involu- 
cres campanulate, 3442 mm high, 8-10 mm wide; 
phyllaries 1 1-13-seriate, the outer ovate-triangu- 
lar, apically acute, mucronate, 5-5.5 mm long, 2.2- 
2.5 mm wide, the inner linear, 3435 mm long, 
1.2-2 mm wide, apically acute; ray florets 13, the 
corolla pink, 4648 mm long, the tube 33-35 mm 
long, the outer lip 12-16 mm long, 34 mm wide, 
the inner lip filiform, 1 1-12 mm long, pubescent; 
filaments free, the anthers 9.5-10 mm long; disc 
florets 3, the corollas 18-18.5 mm long, the tube 
7-7.5 mm long, pubescent, the limb 5-toothed, 
the teeth acuminate; anthers 8-8.5 mm long. 
Achenes [ray] 3-5 mm long, 1.4-1.6 mm wide; 
pappus 18-19 mm long; [disc] 4-4.2 mm long, 3- 
3.2 mm wide; pappus 18-19 mm long. 

DISTRIBUTION Known from northern Peru to 
Bolivia and the only member of the genus to reach 
Brazil; in Peru it grows within the Andean Cor- 
dillera and adjacent lowlands from Lambayeque 
to Cusco (600-1500 m). 

Barnadesia caryophylla has the widest distri- 
bution at the lowest elevations of any member of 
the genus. Its attractive rose-colored corollas and 
absence of spines make it a popular ornamental 
in subtropical regions. 



AMAZONAS. Chachapoyas: Mendoza, Woytkowski 
8121 (MO). CAJAMARCA. Jaen: Colasay, Woytkowski 
6880 (GH, MO). CUSCO. Convention: Alto Urubamba, 
Diehl 2496 (F). JUNIN. Tarma: Chanchamayo, La 
Merced-Quimiri, Ferreyra 3696 (MO, USM); Montayaco, 
W of San Ramon Gentry & Prance 16429 (F); La Merced, 
Macbride & Featherstone 5252 (F); Hacienda Genova, 
Woytkowski 7398 (GH, MO); San Luis de Shuaro, Ridoutt 
s.n. (USM 14564); Satipo, Soukup 2818 (USM); Yucapata, 
Woytkowski 6637 (F, MO); Vitoc, Soukup 4415 (us); La 
Merced, Killip & Smith 23774 (F); Colonia Perene, Killip 
& Smith 25043 (F, us); along Rio Perene, Killip & Smith 
25142 (F, NY, us); entre San Ramon y Palca, Hjerting & 
Petersen 1214 (USM); Valle Chanchamayo, Schunke 334 
(F), 7605 (F); La Merced, Soukup 3391 (F, GH, us); San 
Ramon, Constance & Tovar 2227 (uc). LIMA. Lima: 



10 



FIELDIANA: BOTANY 



Jardin Botanico de la Universidad de San Marcos, Fer- 
reyra 12946 (MO, USM); Plowman 11037 (F); Soukup 2918 
(F, us, USM). SAN MARTIN. Lamas: Roque, Williams 
7241 (F, GH, us). Moyobamba: Moyobamba, Woyt- 
kowski 35300 (F), 15482 (MO, USM). Rioja: Zepelacio 
[Jepelacio], Klug 3318 (F, GH, NY, us). San Martin: Tar- 
apoto, Spruce s.n. (GH). NO EXACT LOCALITY: Peru, 
Diehl 2538 (F). 



4. Barnadesia corymbosa (Ruiz Lopez & Pa von) 
D. Don, Trans. Linn. Soc. London 16: 278. 
1833. 

Bacasia corymbosa Ruiz Lopez & Pavon, Syst. veg. 
fl. peruv. chil. 189. 1798. TYPE: Peru, Huanuco, 
Huanuco, Ruiz Lopez & Pavon s. n. (holotype, MA; 
fragment, F 84551 1; isotype BM, not seen). 

Barnadesia venosa Rusby, Mem. Torrey Bot. Club 
6(1): 69. 1896. TYPE: Bolivia, "Bolivia Plateau," 
M. Bang 1205 (holotype, GH; iso types, F, MO, uc, 
us). 



Trees to 8 m high; branched, glabrous, usually 
without spines. Leaves alternate; petioles 5-30 mm 
long; blades ovate-lanceolate, (35-)60-140 mm 
long, (16-)25-60 mm wide, coriaceous, apically 
acute-acuminate, the adaxial surfaces glabrous, the 
abaxial surfaces glabrescent. Capitulescences cor- 
ymbose, 10-50-headed, the axis 50-75 mm long, 
bracteolate, the bracts lanceolate, 9-28 mm long, 
312 mm wide, pubescent. Capitula with invo- 
lucres campanulate, 1 5-24 mm high, 7-9 mm wide; 
phyllaries 9-10-seriate, the inner linear, 1 7-20 mm 
long, 1.8-2.4 mm wide, apically acute to acumi- 
nate, mucronate, pubescent; ray florets ca. 8, the 
corollas 28-38(43) mm long, 2.8-3 mm wide, the 
tube 20-27 mm long, the outer lip 1 5-1 7 mm long, 
3-3.5 mm wide, the inner lip filiform, 1 1-15 mm 
long, pilose; filaments connate, the anthers 4.5-5 
mm long; disc floret 1 , the corolla 15-18 mm long, 
the tube ca. 16 mm long, pubescent, the limb 
5-toothed, the teeth acute, pubescent; anthers 3.5- 
4 mm long. Achenes [ray] 2.5 mm long, 1.2 mm 
wide; pappus ca. 12 mm long; [disc] turbinate, 2.5 
mm long, 2.2 mm wide; pappus ca. 8 mm long. 

DISTRIBUTION Known from the eastern es- 
carpment of the Andean Cordillera of north-cen- 
tral Peru to central Bolivia (1000-2600 m). 

Barnadesia corymbosa is distinguished by its ar- 
borescent habit, predominately spineless branch- 
es, petiolate leaves, and multiheaded corymbose 
capitulescences. This species superficially resem- 
bles B. parviflora of Ecuador and Colombia but is 
easily distinguished by its fused staminal fila- 
ments. Cabrera (1960, p. 209) determined that 



Barnadesia venosa was conspecific with B. cor- 
ymbosa, and I agree with that placement. 

CAJAMARCA. Jaen: Colasay, W oytkowski 6955 (MO). 
CUSCO. Paucartambo: Santa Isabel, Vargas 5170 (cuz, 
USM); Yanamayo, valle Cosnipata, Vargas 14710 (cuz, 
USM). HUANUCO. Huanuco: Mufia, Macbride 3953 (F, 
GH); Huanuco-Tingo Maria, N of Carpish Pass, Plow- 
man & Rury 11166 (F, 2 sheets); Huallaga, Muna, We- 
berbauer 6717 (F, USM). 

5. Barnadesia dombeyana Less., Linnaea 5: 246. 
1830. LECTOTYPE (designated by Chung, 
1965): Peru, without locality, /. Dombey 448 
(lectotype, p, not seen; photograph ex p, FM 
neg. 52090; isolectotype ex p, F 974660). 

Bacasia spinosa Ruiz Lopez & Pavon, Syst. veg. fl. 
peruv. chil. 188. 1798. TYPE: Peru, no exact lo- 
cality, Ruiz Lopez & Pavon s.n. (holotype, MA, 
not seen; photograph ex MA, USM), non Barna- 
desia spinosa L. fil., 1781. 

Bacasia lanceolata Ruiz Lopez & Pavon ex DC., Prodr. 
7: 3. 1838. TYPE: Peru, no exact locality, Ruiz 
Lopez & Pavon s.n. (not seen). 

Barnadesia lanceolata D. Don, Trans. Linn. Soc. Lon- 
don 16: 277. 1830. 

Shrubs 1 2-30 dm high, branched, branches hir- 
sute to sericeous, spiny, spines divaricate, fuscous, 
2548 mm long. Leaves in fascicles of 3-8; petioles 
1-3 mm long; blades ovate-oblong, 13-60 mm 
long, 4-17 mm wide, acute, mucronate, sericeous. 
Capitulescences of solitary heads, terminal or ax- 
illary, sessile. Capitula with involucres campan- 
ulate, 26-28 mm high, 7-9 mm wide; phyllaries 
(5-)8-9-seriate, the outer ovate 4.2-9 mm long, 
1.8-3.6 mm wide, acuminate, long-apiculate, the 
inner linear, 21-30 mm long, 1-1.8 mm wide, 
acute, the abaxial surfaces sericeous; ray florets 1 3, 
the corollas pink of red-purple, 2740 mm long, 
the tube 1 2-2 1 mm long, 1 .2-1 .8 mm wide, pilose 
to villous, the outer lip 1 5-2 1 mm long, 3.2-5 mm 
wide, pubescent, the teeth acuminate, the inner lip 
12-17 mm long, filiform; filaments connate, the 
anthers 7.5-9 mm long; disc florets 3, the corollas 
12.5-19 mm long, the tube 4-7 mm long, 1.4-2.6 
mm wide, pubescent, the limb 45-toothed, the 
teeth acuminate, the inner lip 7-12.5 mm long, 
filiform; anthers 6.8-7 mm long. Achenes [ray] 3- 
4 mm long, 0.8-2 mm wide, pubescent; pappus 
ca. 18 mm long; [disc] turbinate, 3-4. 2 mm long, 
1.8-3 mm wide; pappus 7-1 1 mm long. 

DISTRIBUTION Known from the Andean Cor- 
dillera of northern to central Peru (2300-4100 m). 
Barnadesia dombeyana is easily distinguished 



FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI. 



11 



dth its densely pubescent habit, long spines, and 
>ng-apiculate outer phyllaries. 
Cabrera (1960, p. 210) examined the holotype 
fBacasia spinosa Ruiz Lopez & Pa von at MA and 
etermined it to be identical to Barnadesia dom- 
eyana. While type material of D. Don's Barna- 
esia lanceolata has not been consulted, his de- 
sription agrees with the concept of B. dombeyana 
dopted here. 

ANCASH. Bolognesi: Ocros, Weberbauer 2755 (USM); 
ear Laguna Conococha, Ferreyra 14453 (USM); Chi- 
uian, Cerrate 523 (MO, USM), 419, 1318, 1407 (USM), 
'erreyra 6216 (USM), 7325 (MO, USM); Matarragra, Fer- 
ryra & Cerrate 5595 (MO, USM); Chiquian, Ferreyra 12136 
fO, USM); Huacar, Cerrate 27 6 (MO, USM), 77 (MO, USM). 
asma: El Tambo, Mostacero 532 (F). Recuay: Pampa 
e Lampa, Edwin & Schunke 3846 (F); Rio Santa, Gentry 
' al. 37463 (F, MO); Catac, near Conococha, Ferreyra 
4292 (MO, USM), Ames 29 (USM), between Chavin and 
.ecuay, Ferreyra 14550 (USM). Santa: El Tambo, Boeke 
32 (MO). CAJAMARCA. Cajamarca: Sunchubamba, 
'llenberg 1701 (USM); 3 km E of Encanada, Hutchison 
: Wright 5120 (F, uc, us, USM); Michiquillo, Pennell & 
eichlin 15041 (USM); Hacienda Polloquito, Sanchez 312 
JSM); Cajamarca-Pacasmayo, cerca El Gavilan, Sdn- 
hez 3340 (F). Celendin: between Balsas and Celendin, 
^utchison & Wright 5312 (F, GH, uc, us, USM). Chota: 
lama, Sandeman 4087 (F). Contumaza: Jalca El Chuno 
*ozo Chuno), Sagastegui 9385 (F, HUT, MO); Cascabam- 
a, Sagastegui et al. 9013 (F, MO), 70007 (MO). HUAN- 
CO. Ambo: Chasqui, Macbride & Featherstone 1759 
; , GH, us). Huamalies: Llata, Macbride & Featherstone 
250 (F, GH, us). LA LIBERTAD. Bolivar: Nevado Ca- 
imarquilla, Ferreyra 1335 (USM); Quishuar, Lopez & 
agdstegui 3280 (HUT); Huamachuco, Lopez 1019 (USM). 
tusco: Agallpampa, Lopez 1059 (HUT, USM). Santiago 
B Chuco: near Shorey, Hutchison et al. 6289 (USM); 
erro la Botica, Lopez 1010 (USM); Chulite, Sagastegui 
' al. 11814 (F). LIMA. Canta: Huaros, Pennell 14720 
0; Buenaventura, Nee s.n. (F); Pachacama, Meza 86 
;SM); Pucachaca, near Buenaventura, Diaz s.n. (USM); 
brajillo, near Canta, Pennell 14397 (F, GH); Capt. Wilkes 
xped. s.n. (us); Churcac, near Canta, Acleto 603 (USM); 
.guallin, near Canta, Sanchez 15 (USM). PASCO. Daniel 
arrion: Yanahuanca, Macbride & Featherstone 1222 (F, 
s). Pasco: between Salcachupan and Cerro de Pasco, 
erreyra 6630 (USM), 8062 (USM); La Quinua-Salcachu- 
an, Ochoa 350 (GH, USM). 

. Barnadesia horrida Muschl., Bot. Jahrb. Syst. 
50, Beibl. Ill: 100. 1913. TYPE: Peru, Cus- 
co, Sacsahuaman, A. Weberbauer 4859 (ho- 
lotype, B, presumably destroyed; photograph 
ex B, FM neg. 1591 1, USM). 

Chuquiraga seleriana Muschl., Bot. Jahrb. Syst. 50, 
Beibl. Ill: 100. 1913. TYPE: Peru, Cusco, Sac- 
sahuaman, E. G. Seler 161 (holotype, B, presum- 
ably destroyed; isotype, GH; photograph ex GH, 
FMneg. 52107). 



Shrubs 1-3 m high, erect, branched, spiny, the 
spines divaricate, ca. 22 mm long. Leaves fascic- 
ulate, 34 per node, subsessile; blades linear to 
lanceolate, 15-25 mm long, 3-5 mm wide, apical - 
ly acuminate-spinulose, glabrescent. Capitules- 
cences of solitary heads, terminal or axillary, sub- 
sessile. Capitula with involucres narrowly com- 
panulate, 28-30 mm high, 5-6 mm wide; phyl- 
laries 7-8-seriate, the inner linear, 26-27 mm long, 
1.41.6 mm wide, apically acuminate, mucronate, 
the abaxial surfaces sericeous; ray florets 8, the 
corollas purple, 36-38 mm long, the tube 18-19 
mm long, the outer lip 17-18 mm long, 3.4-3.8 
mm wide, pubescent, the inner lip 10-1 3 mm long, 
filiform; filaments free, the anthers 8.8-9 mm long; 
disc floret 1, the corollas 17-18 mm long, the tube 
5-6 mm long, pubescent, the limb 5-lobed, the 
teeth acuminate; anthers 8.5-9 mm long. Achenes 
[ray] cylindric, 4.5-5 mm long, 1-1.2 mm wide; 
pappus 1416 mm long; [disc] pyriform, 34 mm 
long, 2.8-3.8 mm wide; pappus 14-17 mm long. 



DISTRIBUTION Endemic to intermontane val- 
leys in the Andean Cordillera of southern Peru 
(3000-3800 m). 

Barnadesia horrida closely resembles B. jelskii 
of northern Peru and southern Ecuador. The for- 
mer species differs in having linear-lanceolate 
leaves, more narrow involucres, and, generally, 7- 
8-seriate phyllaries. 

Chung (1965) established a neotype (Herrera 
1544, us) for Barnadesia horrida; until an ex- 
haustive search for additional Weberbauer 4859 
is conducted, however, the neotype will not be 
recognized. 



AYACUCHO. Andahuaylas: Chincheros, Hutchison 
1718 (F, us). CUSCO. Anta: between Cusco and Lim- 
atambo, Hjerting & Petersen 1513 (USM). Chumbivilcas: 
Bosque de Miraflores, Nunez & Delgado 7974 (F); Velille, 
Vargas 6520 (cuz). Cusco: Cerro Picchu, Ferreyra 9843 
(USM); Cusco, Cardenas 4 182 (us); Ferreyra 20847 (USM); 
Tankarpata, Gentry 43223 (F); Saxaihuaman, Herrera 
1544 (F, us), Herrera s.n. (USM); Lopez 615 (HUT); Sac- 
sahuaman, Pennell 13556 (F, GH); Tambomachay, Ugent 
4372 (us), Vargas 1896 (cuz, GH, NY), Rose 19036 (NY, 
us), Gay s.n. (GH); Socorro, Vargas 3627 (cuz). Paucar- 
tambo: Paucartambo, Balls 6683 (us), Vuilleumier 247 
(GH), Herrera 2956 (us); Quebrada Llulluchayocc, Var- 
gas 4360 (cuz). Quispicanchis: Hacienda Ccapanco, 
Herrera 1055 (us); Hacienda Chiraura, Herrera 2611 
(us); Marcapata, Vargas 14559 (cuz, USM). Urubamba: 
Chupani, Vargas 11119 (us, USM). HUANCAVELICA. 
Tayacaja: Hacienda Tocas, between Colcabamba and 
Paucartambo, Tovar 1931 (USM). 



FIELDIANA: BOTANY 



7. Barnadesia hutchisoniana Ferreyra, Publ. Mus. 

Hist. Nat. "Javier Prado," Ser. B, Bot. 18: 30. 
1 964. TYPE: Peru, Amazonas, Bongara, frente 
al pueblo de Pomacochas, 17 Aug 1963, R. 
Ferreyra & C. Acleto 15201 (holotype, USM 
15732; isotype, us). 

Shrubs 4-6 m high, branched, hirsute, spiny, 
the spines divaricate, fuscous, ca. 25 mm long. 
Leaves fasciculate, 2-5 per node, subsessile; blades 
oblong-elliptic, 10-56 mm long, 6-20 mm wide, 
apically obtuse, mucronate, both surfaces seri- 
ceous. Capitulescences of solitary heads, terminal 
or axillary, the peduncles 5-7 mm long. Capitula 
with involucres campanulate, 30-33 mm high, 10- 
12 mm wide; phyllaries 13-14-seriate, the outer 
ovate, 6-6.5 mm long, 4-5 mm wide, the inner 
oblong-lanceolate ca. 30 mm long, ca. 3 mm wide, 
acute, mucronate, sericeous; ray florets 1 3, the co- 
rollas pink, 40-42 mm long, the tube 27-28 mm 
long, 1.2-1.5 mm wide, pubescent, the outer lip 
15-16 mm long, 44.2 m wide, pubescent, the 
teeth acute, the inner lip 10-1 1 mm long, filiform; 
filaments free, the anthers 9-9.5 mm long; disc 
florets 3, the corollas 14-15 mm long, the tube 5- 
6 mm long, 2.8-3 mm wide, pubescent, the limb 
4 5-lobed, the inner lip filiform. Achenes [ray] 4- 
4.5 mm long, 1.8-2 mm wide; pappus ca. 18 mm 
long; [disc] conical, 5-5.5 mm long, 2.5-3 mm 
wide; pappus 10-15 mm long. 

DISTRIBUTION Endemic to the intermontane 
valleys in the Andean Cordillera of northern Peru 
(1300-3 100m). 

Barnadesia hutchisoniana most closely resem- 
bles B. lehmannii and is distinguished by its small- 
er, sericeous leaves and shorter involucres with 
1 3- 1 4-seriate phyllaries. 

AMAZONAS. Rodriguez de Mendoza: no exact lo- 
cality, Soukup 5032 (USM). CAJAMARCA. Cajamarca: 
San Pablo, Munoz 1015 (USM). Chota: Llama-Chongo- 
yape, Ferreyra 862 (USM); near Llama, Ferreyra 8394 
(USM). Contumaza: arriba de Trinidad, Sagdstegui et al. 
8945 (F); Rupe-Contumaza, Sagdstegui et al. 9819 (F). 
Hualgayoc: Hacienda Lives, Diaz s.n. (USM). LAMBA- 
YEQUE. Lambayeque: W side Abra Porculla Pass km 
35 E of Olmos, Hutchison 1378 (F, GH, uc). PIURA. 
Huancabamba: arriba de Palambla, Lopez & Sagdstegui 
8802 (F, MO). 

8. Barnadesia jelskii Hieron., Bot. Jahrb. Syst. 36: 

511. 1905. TYPE: Peru, Cajamarca, Chota, 
entre Chota y Cutervo, Jun 1879, Jelski 748 
(holotype, B, presumably destroyed; photo- 



graph ex B, FM neg. 15912, USM). NEOTYPE 
(designated by Chung, 1965): Peru, Amazo- 
nas, Chachapoyas, Mat hews s.n. (K, not seen; 
photograph ex K, FM neg. 52088). 

Shrubs 2-6 m high, erect, branched, the branch- 
es tomentose, spiny, the spines divaricate, fuscous, 
ca. 38 mm long. Leaves alternate, fasciculate; pet- 
ioles 1-2 mm long, sericeous; blades oblong-ellip- 
tic, rarely spatulate, 7-28 mm long, 5-10 mm wide, 
apically obtuse, mucronate to spinulose, the ad- 
axial surfaces glabrous, the abaxial surfaces seri- 
ceous. Capitulescences of solitary heads, terminal 
and axillary. Capitula with involucres campanu- 
late, 19-25 mm high, 7-9 mm wide; phyllaries 
10-12-seriate, the inner linear 18-24 mm long, 
1.4-1.8 mm wide, acute, pubescent, fuscous; ray 
florets 8, the corolla pink-purple, 26-33 mm long, 
the tube 13-17 mm long, pubescent, the outer lip 
ca. 16 mm long, ca. 4 mm wide, the teeth acute, 
the inner lip 9-1 1 mm long, filiform, pubescent; 
filaments connate, the anthers 6.4-7.5 mm long; 
disc floret 1, the corollas 1 1-15 mm long, the tube 
3.64.2 mm long, 2-2.4 mm wide, pubescent, the 
limb 5-lobed, the teeth acute; anthers 4.5-5.5 mm 
long. Achenes [ray] 2.2-3.2 mm long, 1-1.4 mm 
wide; pappus ca. 1 1 .5 mm long; [disc] conical, 2.5- 
3.2 mm long, pubescent; pappus ca. 6.8 mm long. 

DISTRIBUTION Known from the Andean Cor- 
dillera from northern Ecuador to northwestern Peru 
(2500-3700 m). 

Barnadesia jelskii appears to be confined to small 
relict forest patches in interandean valleys in 
northern Peru and is known locally as cara-cash- 
ua. 

AMAZONAS. Bongara: Pomacocha, Ldpez et al. 4406 
(HUT). CAJAMARCA. Cajabamba: entre Wacraruco y 
Succhubamba, Diaz 2154 (F); near Cajabamba, Ferreyra 
3039 (USM). Cajamarca: Cumbre El Gavilan, Ferreyra 
3331 (USM); below Cumbe Mayo, Muller & Gutte s.n. 
(USM); San Miguel, El Tingo, Sagdstegui et al. 8798 (MO); 
Cumbre El Gavilan, Sdnchez 582 (MO); Cajamarca- 
Bam bam area. Smith & Vasquez 3445 (F). Celendin: Ce- 
lendin, Dillon & Turner 1637 (F, MO); near Celendin, 
Ferreyra 15032 (USM); Cruz Conga, between Cumullca 
and Celendin, Ferreyra 13821 (USM); Sendamal, Sagdste- 
gui et al. 12235 (F). Chota: 12 km SE of Chota, Gentry 
et al. 61608 (F); above Chota, Ferreyra 8493 (USM). Cu- 
tervo: between Cutervo and Cochabamba, Ferreyra 838 
(USM). Hualgayoc: above San Miguel, Weberbauer 3948 
(USM); Tahona, between Bambamarca and Hualgayoc, 
Ferreyra 8533 (USM). HUANUCO. Huanuco: Quichqui, 
Apac 4115 (USM). LA LIBERTAD. Bolivar: near Nevado 
Cajamarquilla, Infantes s.n. (USM). Huamachuco: Motil, 



FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI. 



13 



Ferreyra 3003 (USM), 3008 (USM); Yanazara-Huaquil, 
Lopez & Sagdstegui 2755 (F). Otusco: Agallpampa, Lopez 
311 (USM). PIURA. Huancabamba: above Huancabam- 
ba, Hutchison 1611 (USM); Mitopampa, Sagdstegui 8239 
(F, MO). 

9. Barnadesia lehmannii Hieron., Bot. Jahrb. Syst. 
29: 77. 1901. LECTOTYPE (designated by 
Harling, 1991): Ecuador, Chimborazo, Pan- 
za-Cajabamba, F. C. Lehmann 5238 (lecto- 
type, K, not seen; isolectotype, NY). 

Barnadesia lehmannii Hieron. var. angustifolia Chung, 
Rev. Barnadesia 50. 1965. TYPE: Peru, Lima, 
Canta, Sandeman 216 (holotype, BM, not seen; 
photograph ex BM, FM neg. 52084; isotype, K, not 
seen). 

Shrubs 1-4 m high, branched, upper branches 
hirsute, spiny, spines divaricate, fuscous, pilose, 
ca. 42 mm long. Leaves alternate, fasciculate, 2- 
4 per node; petioles 3-7 mm long; blades ovate- 
elliptic, 26-70 mm long, 1 5-36 mm wide, apically 
acute or rarely obtuse, mucronate, the adaxial sur- 
faces hirtellous, glabrescent, the abaxial surfaces 
sericeous. Capitulescences of solitary heads, ses- 
sile, terminal and axillary, or weakly cymose with 
2-3 heads. Capitula with involucres cylindric, 30- 
46 mm high, 9-11 mm wide; phyllaries 6-8 -se- 
riate, the outer ovate, acute, 5-7 mm long, 4.2-5 
mm wide, the inner oblong-lanceolate, 2642 mm 
long, 2-4 mm wide, acute, sericeous; ray florets 
13, the corollas 32-54 mm long, the tube 17-25 
mm long, 1.5-1.8 mm wide, the outer lip 15-26 
mm long, 4.5-5 mm wide, pubescent, the teeth 
acuminate, the inner lip 1 1-12 mm long, filiform, 
glabrous; filaments free, the anthers 9-10 mm long; 
disc florets 3, the corollas 1 7-24 mm long, the tube 
6-7 mm long, 2.2-3 mm wide, pubescent, the limb 
4-toothed, the teeth acuminate, the inner lip 11- 
12 mm long, filiform, glabrous; anthers 8-8.5 mm 
long. Achenes [ray] 4-4.2 mm long, 1.2-1.4 mm 
wide; pappus ca. 1 8 mm long; [disc] turbinate, 4 
4.2 mm long, 2.5-3.2 mm wide; pappus ca. 18 
mm long. 

DISTRIBUTION Known from the Andean Cor- 
dillera from central Ecuador to northern Peru 
(1 500-2200 m). 

Barnadesia lehmannii closely resembles B. spi- 
nosa of Colombia but differs in its larger leaves, 
larger involucres, and 6-8-seriate phyllaries. 
Chung's (1965) variety angustifolia is not consid- 
ered sufficiently distinct to allow recognition in 
this treatment. 



PIURA. Ayavaca: above Tandopa, Hutchison & Wright 
6684 (uc, us, USM). Morropon: entre Chalaco y San Mi- 
guel, Diaz & Vdsquez 3031 (F). TUMBES. Zarumilla: 
near Caucho, Maekawa s.n. (USM). 

10. Barnadesia macbridei Ferreyra, Publ. Mus. 
Hist. Nat. "Javier Prado," Ser. B, Bot. 18: 23. 
1 964. TYPE: Peru, Huanuco, Huanuco, Hua- 
cachi, estacion near Muna, J. F. Macbride 3878 
(holotype, F 534941). 

Barnadesia glomerata Kuntze var. distincta Chung, 
Rev. Barnadesia 36. 1965. TYPE: Peru, Junin, 
Jauja, Comas, C. Ochoa 553 (holotype, us 
1952591; isotype, F 1626953). 

Shrubs 12-25 dm high, branched, the upper 
branches hirsute, spiny, the spines divaricate, yel- 
lowish-fuscous, ca. 2 1 mm long. Leaves alternate, 
fasciculate, 3 per node; petioles 1.5-2 mm long; 
blades oblong-elliptic, 1 5-60 mm long, 11-21 mm 
wide, apically obtuse, mucronate, both surfaces 
pubescent. Capitulescences of solitary heads, ter- 
minal or axillary, rarely weakly cymose with 2-3 
heads. Capitula with involucres cylindric, 25-28 
mm high, 6-8 mm wide; phyllaries 9-1 1 -seriate, 
the outer ovate-lanceolate, 5-7 mm long, 1.52 
mm wide, the inner linear, 22-26 mm long, 1 .6- 
1 .8 mm wide, acuminate; ray florets 8, the corollas 
pink or purple, 22-30 mm long, the tube 15-22 
mm long, 0.6-1 mm wide, pubescent, the outer lip 
8-10 mm long, 3.54 mm wide, pubescent, the 
inner lip 6.5-9 mm long, linear, glabrous; fila- 
ments free, the anthers 6.5-7 mm long; disc floret 
1, the corollas 9.5-1 1 mm long, the tube 3.54.5 
mm long, 1.5-2.8 mm wide, pubescent, the limb 
5-toothed; anthers 6-6.5 mm long. Achenes [ray] 
2.53 mm long, 1-1.3 mm wide; pappus ca. 12 
mm long; [disc] conical, 2.8-3.8 mm long, 2.2-3 
mm wide; pappus 8-9 mm long. 

DISTRIBUTION Endemic to the intermontane 
region in the Andean Cordillera of central Peru 
(2200-3350 m). 

Barnadesia macbridei closely resembles B. re- 
ticulata, but, unlike that species, it has conspicu- 
ously hirsute branches and capitula with 8 ray flo- 
rets and only 1 disc floret. 

HUANUCO. Huanuco: near Acomayo, Woytkowski 
34335 (F, USM). JUNIN. Jauja: Comas, Ochoa 553 (F, 
us, USM). LIMA. Lima: Jardin Botanico, Ferreyra 12 946 A 
(MO, USM). 

1 1. Barnadesia polyacantha Wedd., Chlor. And. 
1: 13,t. l.A. 1855. LECTOTYPE (designated 



14 



FIELDIANA: BOTANY 



by Chung, 1965): Bolivia, Larecaja, "Envir. 
de Sorata," 1848, H. A. Weddell 4468 (lec- 
totype, P, not seen; photograph ex p, FM neg. 
52077; isolectotype, F 974857). 

Shrubs 25-40 dm high, branched, the branches 
hirsute, spiny, the spines divaricate, fuscous, ca. 
25 mm long. Leaves alternate, fasciculate, 3-5 per 
node; petioles 1.54 mm long; blades oblong-el- 
liptic, 12-35 mm long, 5-14 mm wide, apically 
acute, spinulose, both surfaces sericeous. Capitu- 
lescences of solitary heads, terminal or axillary, 
sessile. Ca pi tula with involucres campanulate, 25- 
30 mm high, 8-10 mm wide; phyllaries 10-11- 
seriate, the outer ovate, 6-7 mm long, 2.8-4 mm 
wide, acute, mucronate, the inner linear, 23-38 
mm long, 1-2 mm wide, acuminate, sericeous; ray 
florets 13, the corollas 3846 mm long, the tube 
25-29 mm long, 0.8-1 mm wide, pubescent, the 
outer lip 12-16 mm long, 3.84.8 mm wide, pu- 
bescent, the teeth acute, the inner lip 8-10 mm 
long, filiform, flexuose, glabrous; filaments free, 
the anthers 6-7 mm long; disc florets 3, the corollas 
12-17 mm long, the tube 4.5-6.5 mm long, 2-2.2 
mm wide, pubescent, the limb 5-lobed. Achenes 
[ray] cylindric, 2.8-3.2 mm long, 1 .2-1 .4 mm wide; 
pappus 12-16 mm long; [disc] turbinate, 2.5-3.5 
mm long, 2.6-3.2 mm wide; pappus ca. 10 mm 
long. 

DISTRIBUTION Native to the Andean Cordil- 
lera from central Peru to Bolivia (2300-3500 m). 

Barnadesia polyacantha is distinguished by its 
hirsute branches, oblong-elliptic leaves with spi- 
nulose-mucronate apices, and fuscous phyllaries. 
Chung (1 965) described several varieties under this 
species that have not been recognized in this treat- 
ment, including B. polyacantha var. attenuata 
Chung, which Harling (1991) placed under the 
Ecuadorian B. kingii H. Robinson. 

CUSCO. Ante: Cuesta Limatambo, Varbas 2960 (cuz). 
Calca: Vilcabamba, Vargas 3966 (cuz, USM). Paruro: 
Araypallpa, Vargas 440 (cuz, F, USM). Urubamba: Penas, 
Vargas 8120 (us, USM); Ollataytambo, Cook & Gilbert 
657 (us); Janccac, Marin 1614 (us); km 88-95, Ferro- 
carril Santa Ana, Vargas 3395 (cuz, USM). JUNIN. Tar- 
ma: Tiambra, km 14 desde Huasahuasi, Diaz 2919 (F); 
Carpapata, above Huacapistana, Killip & Smith 24391 
(F); near Tarma, Palmer 64 (USM). PUNO. Sandia: entre 
Sandia y Cuyo-Cuyo, Weberbauer 838 (USM); Ferreyra 
16742 (MO, USM). 

12. Barnadesia reticulata D. Don, Trans. Linn. 
Soc. London 16: 277. 1830. TYPE: Peru, Ju- 
nin, "Tarmac Peruvianorum" [Tarma], Ruiz 



Lopez & Pavon s.n. (holotype, MA, not seen; 
isotype fragment, BM). 

Shrubs 2-3 m high, branched, upper branches 
sericeous, spiny, spines divaricate, fuscous, 10-18 
mm long. Leaves alternate, fasciculate, 3-6 per 
node; petioles 1.5-3 mm long; blades oblong-el- 
liptic, 20-^48 mm long, 9-20 mm wide, obtuse, 
mucronate, the adaxial surfaces glabrous, the ab- 
axial surfaces sericeous. Capirulescences of soli- 
tary heads, terminal or axillary, subsessile. Capit- 
ula with involucres campanulate, 43-45 mm high, 
10-11 mm wide; phyllaries 9-10(-12)-seriate, the 
outer ovate-lanceolate 7-7.5 mm long, 4.6-4.8 mm 
wide, brown, the inner linear, 3840 mm long, 
2.3-2.5 mm wide, membranaceous, acute, mucro- 
nate, sericeous; ray florets 13, the corollas pink, 
5051 mm long, the tube 31-32 mm long, 11.2 
mm wide, pubescent, the outer lip 20-2 1 mm long, 
3.84 mm wide, pubescent, the teeth acuminate, 
the inner lip 9-10 mm long, filiform; filaments 
connate, the anthers 11-12 mm long; disc florets 
3, the corollas 18.5-19 mm long, the tube 6-7 mm 
long, 2.8-3 mm wide, pubescent, the limb 
4-toothed, the inner lip 8-9 mm long, filiform, 
glabrous; anthers 10-11 mm long. Achenes [ray] 
4.5-4.8 mm long, 1.2-1.4 mm wide; pappus ca. 
22 mm long; [disc] turbinate, 4.5-4.8 mm long, 
3.8-4 mm wide; pappus ca. 13 mm long. 

DISTRIBUTION Endemic to the Andean Cor- 
dillera of central Peru (2800-3000 m). 

ICA. Pisco: above Pisco, Rauh 386 (USM). LIMA. Can- 
ta: vicinity of Canta, Ferreyra 6914 (USM); Pachacama, 
near Canta, Meza 86A (USM). 

13. Barnadesia wurdackii Ferreyra, Publ. Mus. 
Hist. Nat. "Javier Prado," Ser. B, Bot. 18: 16. 
1864. TYPE: Peru, Amazonas, Chachapoyas, 
upper slopes of Puma-urcu ESE of Chacha- 
poyas, 1 Jun 1962, 2700-3000 m , J. J. Wur- 
dack 681 (holotype, USM 15731; isotypes, F 
1635434, MO, us). 

Barnadesia chachapoyasensis Chung, Rev. Barnade- 
sia 64. 1965. TYPE: Peru, Amazonas, Chacha- 
poyas, Cerro Chunchula, 2500 m, C. Ochoa 1665 
(holotype, us 2059 1 45; photograph ex us, FM neg. 
52096; isotype, USM). 

Shrubs 2-6 m high, spiny, the spines ca. 8 mm 
long, divaricate, caducous on distal portions. 
Leaves alternate, fasciculate, 3-6 per node; peti- 
oles 1-2.5 mm long; blades oblanceolate, 20-50 



FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI. 



15 



im long, 9-16 mm wide, apically acute, mucro- 
ate, both surfaces glabrous. Capitulescences of 
)litary heads, terminal or axillary, weakly cymose 
ith 1-4 heads, the peduncles 30-80 mm long or 
:ssile, subtended by reduced leaf. Capitula with 
ivolucres campanulate, 23-30 mm high, 10-12 
im wide; phyllaries 1 0-1 2-seriate, the outer ovate, 
-9 mm long, 4-5 mm wide, the inner linear, 23- 
2 mm long, 1.2-2 mm wide, acute, recurved, 
:riceous, the margins pilose; ray florets 13, the 
>rollas pink, 32-48 mm long, the tube 1 5-23 mm 
ing, 1-1.2 mm wide, pubescent, the outer lip ca. 
2 mm long, ca. 4 mm wide, the teeth acuminate, 
ic inner lip 13-14 mm long, filiform, flexuose; 
laments connate, the anthers 8.5 9 mm long; 
isc florets 3, the corollas 1418 mm long, the tube 
-6.5 mm long, 2-4.5 mm wide, pubescent, the 
mb 4-toothed, the teeth acuminate, the inner lip 
3-1 1 mm long, filiform; anthers 7-8 mm long. 
chenes [ray] 4-5.2 mm long, 1.2-1.8 mm wide; 
ippus 15-17 mm long; [disc] 3-5 mm long, 2.8- 
mm wide; pappus 8-9.5 mm long. 

DISTRIBUTION Endemic to the interandean re- 
on of northern Peru (2300-3500 m). 
Barnadesia wurdackii resembles B. lehmannii 
it differs in its smaller, glabrous, oblanceolate 
aves and shorter, recurved phyllaries. Robinson 
975) suggested that B. wurdackii was synony- 
ious with Chung's (1965) B. chachapoyasensis, 
id I agree with this placement, even though Har- 
ig (1991, p. 17) questioned the placement of the 
tter species. 

AMAZONAS. Bongara: WNW of Pomacocha, Wur- 
ick 930 (F, us, USM). Chachapoyas: Cerro Calla-Calla, 
twin & Schunke3651 (F, us, USM); above Leimebamba, 
?rreyra 12325 (USM); 15-21 km SW of Leimebamba, 
entry et al. 23188 (F, MO); Cerros Calla Calla, E side, 
> km above Leimebamba, Hutchison & Wright 5596 
, uc, USM); 25 km above Leimebamba on road to Bal- 
s, Hutchison & Wright 6972 (F, uc, USM); SE of Cha- 
lapoyas, King & Bishop 9295 (us); Chachacomo-Lei- 
ebamba, Ochoa 266 9 (USM); Pumaurco, Pennell 15553 
H, USM); Cerro Puma Urco, Soukup 4047 (F). 

[. CHAETANTHERA 

haetanthera Ruiz Lopez & Pavon, Fl. peruv. 
prodr. 106. 1794. TYPE: Chaetanthera ciliata 
Ruiz Lopez & Pavon. 

L'y to Peruvian Species of Chaetanthera 

Plants procumbent, the stems 3^4 cm long; leaves oblong-spathulate, lanate-tomentose, the margins 
entire, apically obtuse; capitula sessile; involucres hemispheric; ray florets ca. 25; disc florets ca. 27 



Erect to prostrate, annual or perennial herbs, 
rarely subshrubs, glabrous, pilose, or lanate; mon- 
oecious (extra-Peruvian species occasionally di- 
oecious). Leaves alternate or opposite; blades lin- 
ear-apiculate to lanceolate, oblanceolate, oblong, 
spathulate, or flabellate, the margins entire to den- 
tate, denticulate, or spinulose. Capitulescences of 
solitary, terminal heads, sessile. Capitula heter- 
ogamous, radiate; involucres campanulate to cy- 
lindric; receptacles plane, epaleate, pubescent to 
naked; phyllaries 3 4-seriate; ray florets 1 -seriate, 
functionally pistillate or neuter, the corollas bila- 
biate, yellow, orange, white, rarely reddish, the 
outer lip ligulate, 3-toothed, the inner lip filiform 
or absent; anthers sterile; styles slender, cylindric, 
bifid; disc florets perfect or functionally staminate, 
the corollas tubular, bilabiate, the outer lip tri- 
dentate, the inner lip bifid; anthers basally caudate, 
the tails plumose, the terminal appendages lan- 
ceolate; styles deeply bifid, the branches apically 
obtuse, papillose distally. Achenes [ray] turbinate 
or lanceolate, glabrous or densely papillose; pap- 
pus of ciliolate bristles, 1 -2-seriate; [disc] cylindric 
or turbinate, glabrous or densely papillose; pappus 
of ciliolate bristles, 1 -2-seriate, white. 

Chaetanthera consists of nearly 40 species, all 
confined to southern South America from central 
Peru to 42S latitude in Chile and Argentina. Only 
four species are recorded from Peru; all tend to 
occur at high elevations (27004900 m) within the 
Andean Cordillera. 



References 

CABRERA, A. L. 1937. Revision del genero Chae- 
tanthera (Compositae). Revista Mus. La Plata, 
Secc. Bot., 1: 1-210. 

FERREYRA, R. 1953. Las especies peruanas del 
genero Chaetanthera. Publ. Mus. Hist. Nat. 
"Javier Prado," Ser. B, Bot., 6: 1-8. 



FIELDIANA: BOTANY 



C. cochlearifolia 

Plants erect, the stems (6-)9-32 cm long; leaves lanceolate to spathulate, villous to glabrous or 
glandular-pubescent, apically 3-7 -dentate or entire; capitula pedunculate or subtended by upper 

leaves; involucres campanulate; ray florets 5-16; disc florets 5-20 2 

2. Stems to 6 cm long; leaves opposite, decussate, the blades oblong-lanceolate, the bases connate, 

the margins strictly entire, strongly involute and leaves appearing cylindric C. stuebelii 

2. Stems 9-32 cm long; leaves alternate to subopposite, never decussate, the blades linear-lanceolate 

to spathulate, the bases never connate, the margins dentate 3 

3. Stems to 9 cm long, or rarely to 20 cm; leaves lanceolate 5-10(-14) mm long, villous to 

glabrescent, apically 3-5-dentate; capitula containing ca. 6 ray florets, 5-7 disc florets 

C. peruviana 

3. Stems to 32 cm long; leaves spathulate, 10-22 mm long, glandular-pubescent, apically 7-dentate; 
capitula containing ca. 1 6 ray florets, ca. 20 disc florets C. chiquianensis 



1. Chaetanthera chiquianensis Ferreyra, Publ. 

Mus. Hist. Nat. "Javier Prado," Ser. B, Bot., 
6: 5. 1953. TYPE: Peru, Ancash, Bolognesi, 
Chiquian, E. Cerrate 1323 (holotype, USM 
15710; isotype, MO). 

Annual herbs, the stems 14-32 cm high, erect, 
pubescent. Leaves alternate, rarely opposite, ses- 
sile; blades spathulate, 10-22 mm long, 3-8 mm 
wide, attenuate at base, glandular-pubescent, api- 
cally ca. 7-dentate, the teeth acute, mucronate, the 
margins plane. Capitulescences of solitary heads, 
axillary or terminal, pedunculate, subtended by 
apical leaves, spathulate, 5 -dentate, 5-8 mm long, 
2-2.8 mm wide, pubescent. Capitula with invo- 
lucres 7-10 mm high, 5-7 mm wide, campanulate; 
phyllaries 3-seriate, the outer lanceolate, 6.6-7.5 
mm long, 1.8-2 mm wide, the inner lanceolate, 
7-8 mm long, 1.8-2 mm wide, membranaceous, 
obtuse, mucronate, glabrous; ray florets ca. 1 6, the 
corollas 5.8-6.2 mm long, the tube 44.2 mm long, 
glabrous, the outer lip ligulate, 1.8-2 mm long, 
0.8-1 mm wide, tridentate, the inner lip bifid, the 
lobes acute, 0.4-0.5 mm long; disc florets ca. 20, 
the corollas 4.5-5 mm long, the tube 3.84.2 mm 
long, glabrous. Achenes [ray] turbinate, 2-2.2 mm 
long, papillose; pappus ca. 4.5 mm long; [disc] 
turbinate, 1.6-2 mm long, 1-1.2 mm wide, pa- 
pillose; pappus 44.5 mm long. 

DISTRIBUTION Endemic to the Andean Cor- 
dillera of central Peru (2700-3500 m). 

Chaetanthera chiquianensis most closely resem- 
bles C. peruviana but it differs in having 7-nerved 
leaves and larger heads with more florets. 

ANCASH. Recuay: Marca, Gomez 345 (USM). 

2. Chaetanthera cochlearifolia (A. Gray) Robin- 



son, Proc. Amer. Acad. Arts 49: 514. 1913. 
Figure 3. 

Oreastrwn cochlearifolium A. Gray, Proc. Amer. Acad. 
Arts 5: 144. 1861. TYPE: Peru, Junin, Yauli, Al- 
pamarca, Capt. WilkesExped. s.n. (holotype, GH). 

Annual herbs, or short-lived perennials, 34 cm 
high, glabrescent. Leaves alternate, sessile; blades 
oblong-spathulate, ca. 10 mm long, ca. 3 mm wide, 
basally attenuate, apically obtuse, both surfaces 
dense lanuginous, the margins entire, revolute. 
Capitulescences of solitary , terminal heads, sessile. 
Capitula with involucres hemispheric, 10-12(-16) 
mm high, ca. 1 5 mm wide; phyllaries 3 4-seriate, 
the outer elliptic, 1 1-12 mm long, 4.5-5 mm wide, 
acute, foliaceous, lanose basally, slightly coria- 
ceous at the apex, the inner lanceolate, 14-16 mm 
long, 2.2-2.8 mm wide, membranaceous, acumi- 
nate, glabrescent, 1-3-nerved, transparent at the 
base; ray florets ca. 25, the corollas 7-7.5 mm long, 
bilabiate, the tube 3-3.2 mm long, glabrous, the 
outer lip 4-4.5 mm long, 0.7-0.8 mm wide, gla- 
brous; disc florets ca. 27, the corollas 5.8-7 mm 
long, the tube 4.2-5.2 mm long, glabrous. Achenes 
[ray] cylindric, 2.2-2.6 mm long, 0.2-0.6 mm wide, 
glabrous; pappus ca. 8.5 mm long; [disc] cylindric, 
2.4-2.8 mm long, 0.4-0.5 mm wide, glabrous; 
pappus 7.5-8 mm long. 

DISTRIBUTION Endemic to the Andean Cor- 
dillera of central Peru (4800-5000 m). 

Chaetanthera cochlearifolia most closely resem- 
bles C. pulvinata (Phil.) Hauman from the Andean 
Cordillera of Chile and Argentina [27-34S lati- 
tude]; however, it differs in its larger leaves, phyl- 
laries, and ray florets. 

JUNIN. Yauli: near Anticona Pass, Dillon & Turner 
1315 (F), 1477 (F); Ticlio, Tovar 7192 (USM); above Ha- 



FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI. 



17 







5mm 



1cm 



FIG. 3. Chaetanthera cochlearifolia. 1, habit; 2, leaf. (Drawn from J.F. Macbride & W. Featherstone 845, F.) 



18 



FIELDIANA: BOTANY 



cienda Acopalca, Weberbauer 6524 (USM). LIMA. Can- 
ta: La Viuda, Meza 225 (USM). Huarochiri: Casapalca, 
Macbride & Featherstone 845 (F). 

3. Chaetanthera peruviana A. Gray, Proc. Amer. 

Acad. Arts 5: 144. 1861. TYPE: Peru, Lima, 
Canta, Banos, Capt. Wilkes Exped. s.n. (ho- 
lotype, GH). 

Annual herbs, 4.5-9 cm high, erect, slightly pu- 
bescent. Leaves alternate, sessile; blades lanceolate 
to spathulate, 6-14 mm long, 24 mm wide, the 
adaxial surfaces glabrous, the abaxial surfaces vil- 
lous to glabrescent, basally attenuate, apically 
3-dentate (rarely 5-dentate), the teeth acute, mu- 
cronate. Capitulescences of solitary, terminal heads, 
peduncles with lanceolate-linear bracts, tridentate, 
5-6 mm long, 1-1.6 mm wide. Capitula with in- 
volucres campanulate, 5-8 mm high, 3-5 mm wide; 
phyllaries 3-^4-seriate, the outer ovate, 4.5-5.8 mm 
long, 1.6-1.8 mm wide, foliaceous, mucronate, 
glabrous, the inner elliptic, 4.2-6.2 mm long, 1.5- 
1.8 mm wide, membranaceous, obtuse, glabrous; 
ray florets 6, the corollas 4.8-6 mm long, the tube 
2.5-3 mm long, glabrous, the outer lip 2.3-3 mm 
long, 0.8-1 mm wide, pubescent; disc florets 5- 
7(-13), hermaphroditic, the corollas 2.5-3 mm 
long, the tube 2-2.5 mm long, glabrous. Achenes 
[ray] turbinate, papillose; pappus ca. 4 mm long; 
[disc] turbinate, 1-1.2 mm long, 0.5-0.6 mm wide, 
papillose, costate; pappus ca. 4 mm long. 

DISTRIBUTION Native to the Andean Cordil- 
lera from central and southern Peru (3000-4000 
m). 

Chaetanthera peruviana most closely resembles 
C. tenella Less, from the Andean Cordillera of 
northern to central Chile; however, it differs in its 
shorter, branched habit and smaller leaves, in- 
volucres, and florets. 

AREQUIPA. La Union: above Alca, Cotahuasi, We- 
berbauer 6876 (F, GH, USM). LIMA. Huarochiri: Matu- 
cana, Macbride & Featherstone 197 (F, USM). NO EX- 
ACT LOCALITY. Peru, 1904-1914, Weberbauer 5419 
(F). 

4. Chaetanthera stuebelii Hieron., Bot. Jahrb. Syst. 

21: 368. 1896. TYPE: Bolivia, La Paz, "cres- 
cit prope Sisasica inter Tomarape et La Paz," 
3800 m, A. Sttibel 15a (holotype, B, presum- 
ably destroyed). 

Annual herbs, ca. 6 cm high. Leaves opposite, 
decussate; blades oblong-lanceolate, 3.5-9 mm 



long, ca. 1.5 mm wide, the adaxial surfaces densely 
tomentose, the abaxial surfaces glabrous, apically 
obtuse to acute, mucronulate, the bases connate, 
the margins strictly entire, strongly involute and 
appearing cylindric. Capitulescences of solitary, 
terminal, sessile heads. Capitula with involucres 
narrowly campanulate, 5-6 mm high, 4-5 mm 
wide; phyllaries 3-4-seriate, the outer oblanceo- 
late, ca. 5 mm long, ca. 1.5 mm wide, glabrous, 
the margins broadly hyaline, obtuse, brownish, the 
inner elliptic, ca. 6.5 mm long, ca. 1.8 mm wide, 
broadly hyaline, obtuse, brownish; ray florets ca. 
1 1, the corolla ca. 6 mm long, the tube ca. 3 mm 
long, glabrous, the outer lip ca. 3 mm long, 0.5 
mm wide, the inner lip ca. 1 mm long; disc florets 
ca. 1 3, the corollas ca. 5 mm long, the tube ca. 4.5 
mm long, glabrous. Achenes columnar, ca. 1 mm 
long, isomorphic, glabrous; pappus ca. 5 mm long. 

PUNO. Puno: San Antonio de Esquilache, Stafford 
873 (F). 



IV. CHAPTALIA 

Chaptalia Venten., Descr. pi. nouv. 61. 1802. 
TYPE: Chaptalia tomentosa Venten. 

Perennial herbs, scapose, acaulescent, rhizo- 
matous. Leaves in basal rosettes, the blades ob- 
lanceolate, lyrate, or lanceolate-ovate, rarely cor- 
diform or elliptic-ovate, lanuginous or tomentose, 
the margins entire, runcinate, crenate, denticulate, 
lobed. Capitulescences of solitary, terminal, nu- 
tant heads; peduncles 2-60 cm long, lanate or 
arachnoid. Capitula disciform, heterogamous; in- 
volucres campanulate or turbinate; receptacles 
plane, epaleate, foveolate; phyllaries 3-7-seriate, 
graduated, glabrous or lanuginous; florets 20-300, 
trimorphic, occasionally dimorphic, the outer- 
most 1 -several rows of pistillate florets, the co- 
rollas ligulate, tridentate, the tube shorter than the 
style, staminodia absent; inner pistillate florets 1- 
several-seriate, the corollas filiform, tubular, styles 
slender, exceeding the tube; central disc florets her- 
maphroditic or more rarely pseudohermaphrodit- 
ic with sterile ovaries, the corollas bilabiate (the 
outer lip 3-parted, the inner lip 2-parted) or reg- 
ularly 5 -lobed; anthers sagittate; styles thickened, 
the branches short, papillose. Achenes columnar 
to fusiform, (4-)5(-l l)-costate, glabrous or pu- 
bescent; pappus of scabrous bristles, white or pink. 

Chaptalia consists of about 50 species distrib- 



FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI. 



19 



uted from the United States to southern South 
America, with 1 here treated for Peru. There has 
been considerable taxonomic disagreement as to 
the generic limits among Chaptalia, Gerbera, and 
Trichocline (Burkart, 1944; Zardini, 1974; Han- 
sen, 1 990). The three are separated with some dif- 
ficulty and various species have been moved among 
these genera. Chaptalia is characterized by tri- 
morphic florets (rarely strictly dimorphic), ray flo- 
rets with corollas merely equalling or slightly sur- 
passing the phyllaries, staminodia absent, and ros- 
trate achenes with inflated trichomes or, more 
commonly, glabrous. Gerbera possesses stamino- 
dia in the ray florets and achenes with slender, 
acute trichomes; Trichocline also possesses ray flo- 
rets with staminodia and achenes with slightly in- 
flated, myxogenic trichomes mixed with capitate- 
glandular types. Burkart (1944) provided a dis- 

Key to Peruvian Species of Chaptalia 



cussion of the characters separating Chaptalia from 
its close relatives and provided a key to putative 
sections of the genus. The treatment presented here 
should be considered provisional; the Andean spe- 
cies of the genus are in need of monographic treat- 
ment. 



References 

BURKART, A. 1944. Estudio del genero de Com- 
puestas Chaptalia con especial referencia a las 
especies argentinas. Darwiniana, 6: 505-594. 

HIERONYMUS, G. 1896. Plantae Stuebelianae no- 
vae. Bot. Jahrb. Syst. 21: 325-377. 

ZARDINI, E. M. 1975. Revision del genero Tri- 
chocline (Compositae). Darwiniana, 19: 618- 
733. 



1. Heads sessile in leaf bases or borne on peduncles 1-2 cm long C. exscapa var. microcephala 

1 . Heads borne on peduncles 5-60 cm long 2 

2. Leaves sessile or with winged petioles to 20 mm long; blades oblanceolate to spathulate with entire 

margins, or lyrate-pinnatifid 3 

3. Leaf blades oblanceolate to spathulate, the margins entire C. integerrima 

3. Leaf blades lyrate-pinnatifid 4 

4. Leaf blades 20-1 10 mm long, 7-30 mm wide; achenes ca. 5 mm long C. similis 

4. Leaf blades 90-350 mm long, 40-100 mm wide; achenes 10-15 mm long . . . . C. nutans 
2. Leaves with unwinged petioles, 20-320 mm long; blades generally ovate, oval, or oblong, the 

margins entire to remotely crenate-denticulate 5 

5. Leaf blades with the abaxial and adaxial surfaces densely tomentose 6 

6. Adaxial and abaxial leaf surfaces densely cinereo-tomentose; peduncles stout, ebracteate, 
densely pubescent flavo-tomentose, to 20 cm long; involucres densely flavo-tomentose; ray 

florets 15-22, the corollas white, the outer lip 13-14 mm long, ca. 1.5 mm wide 

C. malcabalensis 

6. Abaxial leaf surfaces densely flavo-tomentose, the adaxial surfaces arachnoid-tomentose; 
ray florets ca. 25-27, the corollas yellow to orange, the outer lip 6-7 mm long, ca. 1 mm 
wide C. isernina 

5. Leaf blades with abaxial surfaces densely tomentose, the adaxial surfaces glabrous or rarely 
simply glabrescent 7 

7. Leaf blades ovate to oval, (20-)30-50 mm long, (15-)20-35 mm wide, basally cordate to 

truncate 8 

8. Involucres 10-12 mm high C. cordata 

8. Involcures 18-20 mm high C. rotundifolia 

7. Leaf blades oblong-elliptic, 60-110 mm long, 10-25(-^40) mm wide, basally cuneate to 

attenuate 9 

9. Outer phyllaries densely ferrugineo-tomentose C. oblonga 

9. Outer phyllaries lanuginous C. callacallensis 



1. Chaptalia callacallensis Cuatr., Fontqueria 9: 
5. 1985. TYPE: Peru, Amazonas, Chacha- 
poyas, Cerros Calla-Calla, 3100 m, 18 km 



above Leimebamba on road to Balsas, 7 Jun 
1964, P. C. Hutchison & J. K. Wright 5589 
(holotype, us). 



20 



FIELDIANA: BOTANY 



Perennial rosulate herbs to 30 cm tall, rhizo- 
matous. Leaves in basal rosette, petiolate; petioles 
23-32 cm long; blades oblong-elliptic, 80-1 10 mm 
long, 20-23 mm wide, apically attenuate, obtuse 
or subobtuse, basally cuneate, the adaxial surfaces 
glabrescent, pale green, the abaxial surfaces dense- 
ly albo-lanate, the margins remotely sinuate-den- 
ticulate, revolute. Capitulescences of solitary, sea- 
pose heads, 5-6 per rosette, the scapes 30-40 cm 
long, bracteolate. Capitula cernuous or nutant; in- 
volucres narrowly cylindric-campanulate, 15-16 
mm high, ca. 8 mm wide; phyllaries ca. 6-seriate, 
the outer lanceolate, 4-4.5 mm long, 1-1.2 mm 
wide, lanuginous, the inner linear to linear-lan- 
ceolate, 14-15 mm long, 1.5-2 mm wide, scarious; 
florets ca. 55 per head; ray florets ca. 28, the co- 
rollas ligulate, pinkish, 1 1-13 mm long, the tube 
3-4 mm long, the ligules 8-9 mm long, 1-1.2 mm 
wide, the ovary fertile; disc florets ca. 27, the co- 
rollas tubular, bilabiate, yellowish, ca. 8.3 mm long, 
the tube 3.2-3.5 mm long, the abaxial lip ca. 4 
mm long; anthers ca. 4.5 mm long; ovary 3-3.5 
mm long, sterile. Achenes [ray] oblong-elliptic, ca. 
5 mm long, pilose; pappus 5.5-6.7 mm long. 

DISTRIBUTION Only known from the type lo- 
cality. 

2. Chaptalia cordata Hieron., Bot. Jahrb. Syst. 2 1 : 
370. 1 896. LECTOTYPE (designated by Har- 
ling, 1991): Ecuador, Carchi, Paramo del An- 
gel, near El Voladero, 3500 m, 23 Feb 1974, 
G. Marling & L. Andersson 12104 (lectotype, 
GH, not seen). 

Perennial herbs, rhizomatous. Leaves in basal 
rosette; petioles 2-12 cm long; blades cordiform, 
20-40 mm long, 1 8-38 mm wide, basally cordate, 
apically obtuse, the adaxial surfaces arachnoid- 
lanate, glabrescent, the abaxial surfaces white-to- 
mentose, the margins revolute, entire to denticu- 
late. Capitulescences with scapes 9-24 cm long, 
lanate, bracteate, the bracts subulate. Capitula 
usually nutant; involucres campanulate, 10-12 mm 
high, 5-6 mm wide; phyllaries 4-5 -seriate, imbri- 
cate, acute, glabrous, the margins rose to pink, the 
outer linear, 5-6 mm long, ca. 0.7 mm wide, the 
inner linear-lanceolate, 10-12 mm long, 1.2-1.5 
mm wide; ray florets reddish to whitish; disc florets 
white or yellow. Achenes fusiform, 5-costate, gla- 
brous, 4.5-5.5 mm long, 1-1.2 mm wide; pappus 
5-6 mm long, yellowish. 

DISTRIBUTION Known from the Andean Cor- 



dillera from Colombia to northern Peru (2900- 
3700 m). 

Chaptalia cordata is very similar to C. rotun- 
difolia, from which it differs in having cordate 
leaves and smaller, campanulate involucres. These 
two species are easily confused, and detailed pop- 
ulational studies might suggest that they are con- 
specific. The collections cited here are considered 
as one end of a great deal of morphological vari- 
ability in leaf shape and involucre size. 

AMAZONAS. Bongara: Yambarasbamba-Pomaco- 
cha trail between Yanayacu and Pomacocha, Wurdack 
1062 (us). Chachapoyas: between Balsas to Chachapo- 
yas, Dillon & Turner 1735 (F, USM); middle eastern Calla- 
Calla slopes, near km 416-419 of Leimebamba-Balsas 
road, Wurdack 1276 (us, USM); 22 km along road from 
Leimebamba SW toward Celendin, King & Bishop 9249 
(MO, us); Cerros Calla-Calla, Hutchison & Wright 5566 
(uc, us, USM). CAJAMARCA. Cajamarca: Cumbe Mayo 
(Frailones), Becker & Terrones 817 (PPEA, us); Negritos, 
Becker & Terrones 1715 (PPEA, us). PIURA: Huanca- 
bamba: La Cruz, Acleto 399 (USM). 

3. Chaptalia exscapa (Pers.) Baker, Mart. Fl. Br. 
6(3): 379. 1884. 

Tussilago (Chaptalia) exscapa Pers., Syn. pi. 2: 456. 
1807. TYPE: P-JU, not seen. 

Chaptalia exscapa is a highly variable species 
of austral South America, and Burkart (1944) rec- 
ognized three varieties based largely on corolla size 
and achenial pubescence. The following variety 
was described from Peru and accepted by Burkart 
(1944; p. 539) with some doubts. 

3a. Chaptalia exscapa (Pers.) Baker var. micro- 
cephala Domke, Notizbl. Bot. Gart. Berlin- 
Dahlem 13: 246. 1936. TYPE: Peru, Puno, 
Santa Rosa, Nov 1933, D. Stafford 243 (ho- 
lotype, K). 

Acaulescent, perennial herbs, 10-25 mm tall. 
Leaves in basal rosette; sessile or petioles to 8 mm 
long; blades oval to oblanceolate, 10-20 mm long, 
5_8(-14) mm wide, the abaxial surfaces densely 
albo-tomentose, the adaxial surfaces glabrescent, 
the margins retrorse-dentate to crenate. Capitu- 
lescences of solitary capitula, sessile or on pedun- 
cles to 20 mm long. Capitula erect; involucres cy- 
lindrical, 6-10 mm high, ca. 6 mm wide; phyllaries 
2-3 -seriate, the outer subulate, ca. 4 mm long, the 
inner lanceolate, 7-8 mm long, apically attenuate, 
all glabrous; ray florets ca. 2 1 , the corollas bilabiate, 
the tube ca. 3 mm long, the outer lip ca. 4.2 mm 



FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI. 



21 



3ng, ca. 0.8 mm wide, the adaxial surfaces white, 
lie abaxial surfaces pink, 3-toothed; disc florets 
ermaphroditic, the corollas white, tubular, acti- 
omorphic, the limb 5-lobed, the lobes equal or 
nequal. Achenes fusiform, 4-7 mm long, 7-8- 
ostate, glabrous; pappus 3.2-3.5 mm long. 

Chaptalia exscapa var. microcephala is appar- 
ntly a rare variety, known only from a few col- 
;ctions. It is easily distinguished by its essentially 
essile heads on nonelongating peduncles con- 
ealed in the leaf bases; it varies from the typical 
ariety in possessing smaller heads and glabrous 
chenes. The collection cited here from Cajamarca 
Becker & Terrones 2479, us) varies from the type 
ollection of C. exscapa var. microcephala in pos- 
essing long-attenuate inner phyllaries. Further 
ollections must be examined before the extent of 
ariation of this and other characters can be eval- 
lated. 



CAJAMARCA. Cajamarca: Llano Cruz, Becker & 
^errones 2479 (us). 

. Chaptalia integerrima (Veil.) Burkart, Darwin- 
ian 6: 576. 1944. 

Tussilago integerrima Veil., Fl. flumin. Icon. 8, t. 140. 
(1827) 1831. TYPE: not seen. 

Perennial herbs to 60 cm. Leaves in basal ro- 
ette; sessile; blades oblanceolate to spathulate, 80- 
:50 mm long, 15-35 mm wide, basally attenuate, 
pically acute to obtuse, the abaxial surfaces densely 
Ibo-tomentose, the adaxial surfaces arachnoid- 
omentose, glabrescent, the margins entire. Capi- 
ulescences of solitary heads, the scapes 14-60 cm 
Dng, lanuginous. Capitula erect; involucres cam- 
>anulate, 15-18 mm high, 12-15 mm wide; phyl- 
aries 3-4-seriate, albo-tomentose, the outer lin- 
ar, 4-5 mm long, apiculate, the inner lanceolate, 
4-15 mm long, apiculate; ray florets white or 
eddish, the corollas ligulate, the outer lip linear, 
lot exceeding involucre; disc florets white. Achenes 
usiform, ca. 8 mm long, 5-costate, papillose with 
>iseriate trichomes; pappus 10-12 mm long, red- 
lish. 

DISTRIBUTION Known from the Andean Cor- 
lillera from Venezuela to Bolivia. In Peru, it is 
ipparently a rare species known only from two 
ocalities in northern Peru (1800-2500 m). 

Chaptalia integerrima is distinguished by its en- 
ire, oblanceolate to spathulate leaves with long- 



attenuate bases and scapes with erect, densely 
white-tomentose heads. 

AMAZONAS. Chachapoyas: mountains behind Tin- 
go, King & Bishop 9282 (us). JUNIN. Tarma: Chan- 
chamayo valley, Schunke 1392 (F). SAN MARTIN. San 
Martin: Tarapoto, Spruce 4181 (GH). 

5. Chaptalia isernina Cuatr., Anales Univ. Ma- 

drid 4(2): 212. 1935. TYPE: Peru, Junin, 
Chanchamayo, "in sylvis," 12 Oct 1863, J. 
Isern 42 (holotype, F). 

Perennial rosulate herbs, 10-45 cm tall. Leaves 
in basal rosette, petiolate; petioles 1 5-25 mm long; 
blades oblong-elliptic, 50-90 mm long, 1 7-30 mm 
wide, subcoriaceous, apically obtuse to rounded, 
basally attenuate, the adaxial surfaces arachnoid- 
tomentose, glabrescent, the abaxial surfaces dense- 
ly stramineous-tomentose, the margins remotely 
sinuate-denticulate. Capitulescences of solitary, 
scapose heads, usually 1 per rosette, the scapes 
10-^45 cm long, bracteolate, densely stramineous- 
tomentose. Capitula erect; involucres cylindric, ca. 
15 mm high, 3.54 mm wide; phyllaries 3-5-se- 
riate, the outer lanceolate, ca. 5 mm long, 0.5-1 
mm wide, apically attenuate, the inner lanceolate, 
11-13 mm long, 1-1.5 mm wide, apically atten- 
uate, the margins scarious; florets heteromorphic; 
pistillate florets 2-seriate, the corollas ligulate, yel- 
lowish-orange, 13-14 mm long, the tube ca. 5 mm 
long, the ligules 3-5 mm long, the ovary fertile; 
disc florets hermaphroditic, the corollas tubular, 
bilabiate, yellowish, ca. 10 mm long. Achenes [ray] 
fusiform, 2.5-3 mm long, glabrous; pappus ca. 7 
mm long. 

Chaptalia isernina is readily recognized by its 
short petiolate leaves with densely yellowish-to- 
mentose abaxial surfaces. 

HUANUCO. Huanuco: Pillao, 2700 m, 19 Feb 1946, 
Woytkowski 34110 (F). 

6. Chaptalia malcabalensis Cuatr., Ciencia (Mex- 

ico) 24: 123. 1965. TYPE: Peru, Amazonas, 
Chachapoyas, summit of Cerro Malcabal 
(Cerro Tumbe) 3-6 km SW of Molinopampa, 
2850-2900 m, /. J. Wurdack 1414 (holotype, 
us 2373662; isotype, USM 32682). Figure 4. 

Perennial herbs, 20-30 cm high. Leaves in basal 
rosette; petioles ca. 2040 mm long, ca. 3 mm 
wide, densely tomentose; blades ovate-elliptic, 35- 
40(-55) mm long, 1 5-26 mm wide, basally cuneate 



FIELDIANA: BOTANY 



to rounded, apically obtuse, both adaxial and ab- 
axial surfaces densely cinereo-tomentose, the mar- 
gins entire or slightly crenate-undulate. Capitu- 
lescences with scapes to 30 cm long, densely flavo- 
tomentose, ebracteate. Capitula nutant; involucres 
campanulate, ca. 20 mm high, ca. 25 mm wide; 
phyllaries 3-4-seriate, all densely ochraceous-to- 
mentose, the outer linear, 5-8 mm long, ca. 0.8 
mm wide, the inner linear-lanceolate, 12-15 mm 
long, 1.5-1.8 mm wide, all apically acute; ray flo- 
rets ca. 13, white; disc florets ca. 20, cream-col- 
ored. Achenes cylindric, 2.6-2.8 mm long, 
5-costate, glabrous; pappus 1 2-14 mm long, white. 

DISTRIBUTION Known only from the type lo- 
cality. 

Chaptalia malcabalensis is distinguished by its 
ovate-elliptic, short petiolate leaves and densely 
lanate scapes without bracts. 

7. Chaptalia nutans (L.) Polak., Linnaea 41: 582. 
1878. 

Tussilago nutans L., Syst. Nat. 10(2): 1214. 1795. 
TYPE: America, collector unknown, Herb. Linn. 
995.5 (holotype, LINN, not seen). 

Perennial herbs, 40-90 cm high. Leaves in basal 
rosette; blades lyrate-pinnatifid, 90-350 mm long, 
40-100 mm wide, the adaxial surfaces glabrous or 
slightly pubescent, the abaxial surfaces densely la- 
nuginous. Capitulescences of 1 -several scapes. Ca- 
pitula nutant; involucres campanulate to turbi- 
nate, 11-15 mm high, 8-10 mm wide; phyllaries 
3-4-seriate, linear-lanceolate, acute; florets tri- 
morphic, white; outer series pistillate, the corollas 
ligulate, the ligules narrow, ca. 1 2 mm long; second 
series pistillate, the corolla tubular, shorter than 
the style; innermost disc florets hermaphroditic, 
the corolla bilabiate. Achenes fusiform or linear, 
10-15 mm long, reddish-brown to green, 5-8-cos- 
tate, papillose with minute, biseriate trichomes; 
pappus 12-15 mm long, yellowish to pink. Chro- 
mosome number: 2n = 48. 

DISTRIBUTION Known from North and Central 
America to southern South America (600-1200 
m). In Peru, it is usually found in the ceja de la 
montana or the eastern escarpment of the Cor- 
dillera Oriental. 

Chaptalia nutans is easily recognized by its large 
spathulate to lyrate basal leaves and nutant heads. 

AMAZONAS. Chachapoyas: Mendoza, Woytkowski 
8144 (MO). CUSCO. La Convention: Rosario Mayo, Var- 
gas 21180 (us); Maranura, Chaullay, Nunez 8143 (F). 




FIG. 4. Flowering individual of Chaptalia malca- 
balensis. (Drawn from /./. Wurdack 1414, us.) 



Urubamba: between Molino and Maras, Vargas 12474 
(us). HUANUCO. Huanuco: Puente Durand, Stork & 
Norton 9584 (F, GH, MO, uc). Leoncio Prado: near Tingo 
Maria, Allard 21654 (F, uc, us); Tingo Maria, Ferreyra 
12769 (MO, USM); Cayumba, Ferreyra 6743 (USM). JU- 
NIN. Jauja: Satipo, Ridoutt s.n. (USM); Satipo, Woyt- 
kowski 5887 (us). Tarma: Chanchamayo, Esposto s.n. 
(USM); near San Ramon, Ferreyra 379 (USM); Tulumayo, 
Ferreyra 1618 (GH, MO, USM); San Luis de Shuaro, Fer- 
reyra 18619 (USM); San Ramon, Hermann 11327 (us); 
La Merced, Killip & Smith 23427 (us); Perene, Kunkel 
826 (us); Chanchamayo valley, Schunke 388 (F), 390 (F); 
La Merced, Soukup 3394 (F). MADRE DE DIGS. Tam- 
bopata: 0-4 km from Puerto Maldonado, Gentry & Re- 
villa 16343 (F). PASCO. Oxypampa: Pozuzo, Macbride 
4704 (F). SAN MARTIN. San Martin: Tarapoto, Fer- 



FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI. 



23 



eyra 5066 (MO, USM); Pucaloma, Ferreyra 7795 (us, 
JSM); Alto Puca Yacu, Monies 55 (F); Tarapoto, Wil- 
iams 5500 (F). Lamas: Cumbaza River, Belshaw 3526 
uc); near Lamas, Ferreyra 4661 (USM); Zepelacio [Je- 
>elacio], near Moyobamba, Klug 3476 (F, GH, MO). Ma- 
iscal Caceres: Tocache Nuevo, Schunke 4945 (F, us), 
1204 (F), 72575 (F, us). UCAYALI. Coronel Portillo: 
:ordillera Azul, Young & Sullivan 710 (F). 

\. Chaptalia oblonga D. Don, Trans. Linn. Soc. 
London 16: 240. 1830. 

Perdicium oblongum Ruiz Lopez & Pavon ex D. Don, 
Trans. Linn. Soc. London 16: 240. 1830. TYPE: 
Peru, "in Peruviae Andibus prope Panao," Ruiz 
Lopez & Pavon s.n. (holotype, MA, not seen). 

Trichodine oblonga (D. Don) Burkart, Darwiniana 6: 
592. 1944. 

Trichodine oblonga Hieron., Bot. Jahrb. Syst. 2 1 : 370. 
1 896. TYPE: Peru, "crescit prope Fraileyacu inter 
Pacasmayo et Moyobamba, alt. 3300 m," A. Stu- 
bel 24 (holotype, B, presumably destroyed). Non 
T. oblonga (D. Don) Burkart, 1944. 

Rosulate, perennial herbs. Leaves petiolate; pet- 
oles 20-50 mm long, tomentose; blades oblong 
o elliptic, 30-55 mm long, 10-30 mm wide, api- 
:ally obtuse to rounded, basally rounded to ob- 
use, the adaxial surfaces densely arachnoid-to- 
nentose, the abaxial surfaces densely tomentose, 
>chraceous to ferruginous, the margins remotely 
lenticulate. Capitulescences of solitary heads, the 
icapes bracteolate, tomentose. Capitula nutant; 
nvolucres campanulate, ca. 1 5 mm high, ca. 1 5 
nm wide; phyllaries 3-4-seriate, narrowly lanceo- 
ate, apically acute, densely ferrugineo-tomentose, 
he outer ca. 9 mm long, the inner ca. 1 6 mm long; 
>uter florets bilabiate, the corolla 12-15 mm long, 
he tube 34 mm long, the outer lip ligulate, 7-1 1 
nm long, apically 3-toothed, the inner lip 1-5 mm 
ong, bifid; disc florets 8-9 mm long, the corollas 
ailabate, the outer lip 3-dentate, the inner lip bifid, 
^chenes fusiform, 2.5-6 mm long, distally con- 
tricted, 5-6-costate, glabrous; pappus 7-9 mm 
ong. 

DISTRIBUTION Known from high-elevation 
labitats within the Andean Cordillera from south- 
ern Ecuador to north-central Peru (3000-3500 m). 

Chaptalia oblonga is distinguished by its long- 
jetiolate, oblong leaves and the densely ferrugi- 
leo-tomentose phyllaries. Hieronymous (1896) 
vas apparently unsure of D. Don's species and 
iescribed what he took as a new species under 
Trichodine and cited a northern Peruvian speci- 
nen (A. Stubel 24, B) under the description. Zar- 
lini (1975) argued that it was best accommodated 



in Chaptalia since it lacks staminodia and achenial 
pubescence. A complete specimen of C. callacal- 
lensis has not been examined; however, the de- 
scription and fragments (ex us) suggest close sim- 
ilarity to C. oblonga. 

AMAZON AS. Bagua: Cordillera Colan, E of La Peca, 
Barbour3157 (F), 3243 (F). HUANUCO. Huanuco: Mito, 
Macbride & Featherstone 1793 (F). 

9. Chaptalia rotundifolia D. Don, Trans. Linn. 
Soc. London 16: 242. 1830. TYPE: Peru, 
Huanuco, Pillao, Ruiz Lopez & Pavon s.n. 
(holotype, MA, not seen). 

Chaptalia cordata \ar.ferrugineo-tomentosa Hieron., 
Bot. Jahrb. Syst. 36: 512. 1905. TYPE: Peru, Ca- 
jamarca, Jelski 726 (holotype, B, presumably de- 
stroyed; photograph ex B, FM neg. 1 6000). 

Perdicium ovale Ruiz Lopez & Pavon ex D. Don, 
Trans. Linn. Soc. London 16: 241. 1830. TYPE: 
Peru, Junin, Tarma, "in Peruvia ad Huassahuassi 
et Churapallanam" (holotype, MA, not seen). 

Chaptalia ovalis D. Don, Trans. Linn. Soc. London 
16: 240. 1830. 

Trichodine ovalis (D. Don) Hieron., Bot. Jahrb. Syst. 
21: 370. 1896. 

Perennial herbs; rhizomatous. Leaves in basal 
rosette; petioles 25-60 mm long, conspicuously 
lanose; blades oval to round, 25-45 mm long, 15- 
35 mm wide, lanuginous, base amplexicaul, api- 
cally obtuse to acute, the adaxial surfaces arach- 
noid, the abaxial sufaces white-tomentose, the 
margins denticulate. Capitulescences with scapes 
1 2-27 cm long, lanose, bracteate, bracts subulate, 
glabrous. Capitula nutant; involucres turbinate, 1 8- 
20 mm high, 10-12 mm wide; phyllaries 5-6-se- 
riate, the outer linear, 5-10 mm long, 0.5-1 mm 
wide, acuminate, the inner lanceolate, 10-18 mm 
long, 1 .8-2 mm wide, more or less reddish at apex, 
acuminate; ray florets ligulate, the ligule linear, 
reddish, female; disc florets linear, white, bilabi- 
ate. Achenes fusiform, costate, glabrous, 7-9 mm 
long, 4-5 mm wide; pappus 9-10 mm, yellowish. 



DISTRIBUTION Known from the Andean Cor- 
dillera in northern to southern Peru (3300-4200 
m). 

Chaptalia rotundifolia most closely resembles 
C. cordata and is rather arbitrarily distinguished 
from that species by its rotund to ovate, noncor- 
date leaf blades and larger involucres. Detailed 
populational studies are needed, especially in 
northern Peru, where the two species are appar- 
ently sympatric. 



FIELDIANA: BOTANY 



Hieronymus (1905) described C. cordata var. 
ferrugineo-tomentosa, calling attention to its 
densely ferrugineo-tomentose phyllaries. The type 
material from northern Peru is no longer available 
for study; however, a photograph of the type (FM 
neg. 1 6000 ex B) shows that Burkart examined the 
holotype collection in 1930 and annotated it as 
equaling C. rotundifolia. The photograph of C. 
ovalis (FM neg. 3807 1 ex p) and D. Don's descrip- 
tion both conform to material here placed under 
C. rotundifolia. 

ANCASH. Carhuaz: Huascaran National Park, Que- 
brada Ishinca, Smith & Buddensiek 1 1272 (F). Huaras: 
Quebrada Churup, Smith et al. 9622 (F); Laguna Quero- 
cocha, Tovar & Rivas 9549 (USM). Huaylas: Quebrada 
Alpamayo, Smith et al. 9847 (us). Yungay: Quebrada 
Ranincuray, Smith et al. 10373 (F). CAJAMARCA. Ca- 
jamarca: Cumbe Mayo, Sanchez 3443 (F, MO). CUSCO. 
Paucartambo: Acjanaco, Parque Nacional del Manu, 
Cano 3142 (F, USM), 3204 (F, USM); Pinasniocj, Pantia- 
calla Pass, Cook & Gilbert 1881 (us); Aconaco, Woyt- 
kowski 056 (USM). HUANUCO. Dos de Mayo: Huallan- 
ca Valley, Tovar & Rivas 9895 (USM). Huanuco: 1 5 mi 
NE of Huanuco, Macbride & Featherstone 2189 (F, us). 
JUNIN. Huancayo: Huancayo, Soukup 2735 (F). LIMA. 
Huarochiri: Viso, Macbride & Featherstone 585 (F); Rio 
Blanco, Macbride 3014 (F, us). SAN MARTIN. Maris- 
cal Caceres: Chochos, Young & Leon 4829 (F). 

10. Chaptalia similis R. E. Fries, Nova. Acta Re- 
giae Soc. Sci. Upsal., ser. 4, 1: 95. 1905. CO- 
TYPES: Argentina, Jujuy, Puna, Sta. Catalina 
Kurtz 14460 (BAF, not seen); Bolivia, La Paz, 
Potopoto, Mandon 13 (K, not seen). 

Perennial herbs to 22 cm tall; rhizomatous. 
Leaves in basal rosette; petioles 1 5-20 mm long; 
blades lyrate-pinnatifid, 20-110 mm long, 7-30 
mm wide, basally attenuate, apically acute to ob- 
tuse, mucronate, the adaxial surfaces lanuginous- 
tomentose, the abaxial surfaces densely albo-to- 
mentose, the margins sinuate-crenate. Capitules- 
cences with scapes 3-20 cm high, tomentose, brac- 
teate, the bracts subulate. Capitula nutant; invo- 
lucre campanulate, 10-15 mm high, 7-8 mm wide; 
phyllaries 3-4-seriate, the outer linear, 4-5 mm 
long, apiculate, the inner linear, ca. 1 8 mm long, 
acute, tomentose to glabrous; florets 25-40, tri- 
morphic; outer ray florets ligulate, the corollas 
white, the outer lip ligulate, inconspicuous, not 
surpassing involucre, the inner lip bifid; inner ray 
florets hermaphroditic, ca. 1 0, the corollas tubular, 
the limb bilabiate; disc florets hermaphroditic, 7- 

1 1 , the corollas tubular, the limb bilabiate. Achenes 
fusiform, ca. 5 mm long, papillose with biseriate 
trichomes; pappus 8-10 mm long, white. 



DISTRIBUTION Known from extreme southern 
Peru, through Bolivia, and into northern Argen- 
tina (1500-4500 m). 

Chaptalia similis is distinguished by its lyrate- 
pinnatifid leaf blades with sinuate-crenate margins 
and lower surfaces lanate tomentose. Burkart 
( 1 944) cited this taxon from Lago Titicaca to west- 
central Argentina and the Peruvian collection cit- 
ed here (Vargas 12474, us) is the northernmost 
recorded occurrence of the species. 

CUSCO. Urubamba: entre Molino Waycko y Maras, 
Vargas 12474 (us). 



V. CHUCOA 

Chucoa Cabr., Bol. Soc. Argent. Bot. 6: 40. 1955. 
TYPE: C. ilicifolia Cabr. 

Shrubs much-branched. Leaves alternate, sub- 
sessile; blades lanceolate to oblanceolate, the mar- 
gins unequally dentate with 46 pairs of spinulose 
teeth. Capitulescences of solitary, terminal heads, 
pedunculate. Capitula homogamous, discoid; in- 
volucres turbinate; receptacles plane, alveolate, 
epaleate, densely pilose-fimbrillate; phyllaries 4- 
5-seriate; florets isomorphic, or occasionally, di- 
morphic with abortion of anthers in marginal flo- 
rets, hermaphroditic, the corollas yellow, the limb 
5-lobed, the lobes linear; stamens 5, the anther 
appendages linear-lanceolate, the bases caudate, 
glabrous; styles bilobed, the branches apically ob- 
tuse, the abaxial surfaces papillose. Achenes cy- 
lindric, glabrous or papillose; pappus of scabrid 
bristles, 2-3-seriate, isomorphic. 

Chucoa is a monotypic genus with only the fol- 
lowing northern Peruvian species recorded. 

References 

CABRERA, A. L. 1955. Un nuevo genero de Mu- 
tisieas del Peru. Biol. Soc. Argent. Bot., 6: 40- 
44. 

FERREYRA, R. 1980. Notas sobre especies Per- 
uanas de los generos Arnaldoa, Chucoa (Com- 
positae) y Monnina (Polygalaceae). Bol. Soc. 
Peruana Bot., 8: 106-109. 

1. Chucoa ilicifolia Cabr., Bol. Soc. Argent. Bot. 
6: 42. 1955. TYPE: Peru, La Libertad, San- 
tiago de Chuco, Angasmarca-Tulpo, 2930 m, 
1 9 Jun 1 954 A. Lopez M. 1090 (holotype, LP). 
Figure 5. 



FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI. 



25 




FIG. 5. Chucoa ilicifolia. 1, habit; 2, capitulum; 3, floret with achene. (Drawn from A. Weberbauer 7017, F.) 



FIELDIANA: BOTANY 



Weberbaueriellajohnstoniana Ferreyra, Bol. Soc. Pe- 
ruana Bot. 5: 2. 1955. TYPE: Peru, La Libertad, 
Santiago de Chuco, below Mollepata, A. Weber- 
bauer 7017 (holotype, GH; isotype, F, USM). 

Suffrutescent shrubs, 40-50 cm high. Leaves with 
petioles 2-6 mm long, tomentose; blades oblan- 
ceolate or lanceolate, 22-70 mm long, 10-28 mm 
wide, the adaxial surfaces glabrous, the abaxial 
surfaces white-tomentose, apex acute, spinulose, 
the margins unequally dentate, the teeth 1-3 mm 
long, the spines 2-3 mm long. Capitulescences of 
solitary, terminal or axillary heads or weakly cy- 
mose, 1-3-headed; peduncles 32-130 mm long, 
lanuginous-glandulose, bracts 3-8 mm long, subu- 
late. Capitula with involucres 14-17 mm high, 8- 
1 5 mm wide; phyllaries 4-5 -seriate, the outer su- 
bulate, ca. 5 mm long, ca. 1 mm wide, the inner 
lanceolate, 8-16 mm long, 1-2 mm wide, atten- 
uate; florets 15-20, the corollas 14-18 mm long, 
the tube 11-15 mm long, 5-lobed, the lobes linear, 
2-3 mm long, coiled; anthers 8-10 mm long, the 
apical appendage ca. 3 mm long, the tails ca. 2.5 
mm long. Achenes cylindric, ca. 3 mm long; pap- 
pus 12-15 mm long, yellowish-white. 

DISTRIBUTION Endemic to the intermontane 
region of northern Peru (2200-3000 m). 

Chucoa ilicifolia is apparently a rare species 
known only from the type and one other collec- 
tion. 



sessile. Capitula homogamous, discoid; involucre 
cylindric, campanulate, or turbinate; receptacle al- 
veolate, epaleate, pilose; phyllaries 5-10(-12)-se- 
riate, scarious to coriaceous, imbricate, pubescent, 
the outer ovate-lanceolate, sericeous, the inner lin- 
ear-lanceolate, mucronate; florets hermaphroditic, 
the corollas yellow to orange, densely villous, tu- 
bular, the limb 5-lobed; stamens 5, the anthers 
basally caudate, the terminal appendages acute to 
obtuse; styles filiform, bifid, the branches rounded, 
papillose-glandular distally. Achenes conical, 
densely sericeo-villous; pappus of plumose bris- 
tles, 1 -seriate, yellowish to purple, united at base. 

Chuquiraga consists of approximately 20 spe- 
cies largely confined to the Andean Cordillera from 
Colombia to Argentina and Chile. In Peru the ge- 
nus is represented by five species distributed in 
the jalca and puna formations at elevations above 
3000 m. Chuquiraga is most similar to Arnaldoa 
and distinguished from that genus by possessing 
actinomorphic corollas with five lobes and sta- 
minal filaments inserted at the base of the corollas 
rather than above the base as in Arnaldoa. Chu- 
quiraga is readily separated from Dasyphyllum with 
the former possessing coriaceous leaves with only 
one central nerve, yellow to orange corollas, and 
anthers with linear-lanceolate, apical appendages. 
Various species of Chuquiraga are used in folk 
medicine and are commonly found in herb mar- 
kets. 



VI. CHUQUIRAGA 

Chuquiraga Juss., Gen. pi. 178. 1789. TYPE: Chu- 
quiraga jussieui J. F. Gmelin 

Intricately branched shrubs, frequently with ax- 
illary spines, erect or rounded to flattened cush- 
ions. Leaves opposite or alternate; sessile or short- 
ly petiolate; blades coriaceous, uninerved, often 
apically spinulose, the margins entire, plane, in- 
crassate, or revolute. Capitulescences of solitary 
heads or loose aggregates, terminal and/or axillary, 



References 

ESCURRA, C. 1985. Revision del genero Chu- 
quiraga (Compositae Mutisieae). Darwiniana, 
26(1-4): 219-284. 

ESCURRA, C., AND J. V. CRISCI. 1 987. Relaciones 
feneticas entre las especies del genero Chuquira- 
ga (Compositae Mutisieae): Un analisis nu- 
merico. Darwiniana, 28(1^4): 219-229. 

TOVAR, O. 1952. Revision de las especies peru- 
anas del genero Chuquiraga, Publ. Mus. Hist. 
Nat. "Javier Prado," Ser. B, Bot., 5: 1-29. 



Key to Peruvian Species of Chuquiraga 

1 . Plants with spines 2 

2. Leaf blades rounded, sessile; involucres turbinate, 34-40 mm high; florets 10-16 

C. rotundifolia 

2. Leaf blades ovate-elliptic or oblanceolate, subsessile; involucre campanulate, 25-30 mm high; 
florets 15-20 . . 3 



FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI. 



27 



3. Leaves ovate-lanceolate, pubescent, then glabrescent; phyllaries 8-10-seriate, orange; corollas 

19-22 mm long C. spinosa 

3. Leaves oblanceolate, tomentose; phyllaries 7-9-seriate, greenish to brunneous; corollas 16-17 

mm long C. johnstonii 

Plants without spines 4 

4. Leaves alternate, subsessile, ovate-lanceolate, 5-15 mm long, 3-7 mm wide; florets 21-40 

C. jussieui 

4. Leaves opposite, sessile, elliptic-lanceolate, 6-28 mm long, 3-8.5 mm wide; florets 2045 

. C. weberbaueri 



. Chuquiraga johnstonii Tovar, Publ. Mus. Hist. 
Nat. "Javier Prado," Ser. B, Dot. 5: 8. 1952. 
TYPE: Pasco, Daniel Carrion, Yanahuanca, 
J. F. Macbride & W. Feather stone 1209 (ho- 
lotype, OH; isotypes, F, us). 

Shrubs 9-12 dm high, much-branched, the 
ranches pubescent, spiny, the spines axillary, 3.5- 
mm long, curved at the base. Leaves opposite, 
ubsessile; blades oblanceolate, rarely lanceolate, 
-20 mm long, 3-6.5 mm wide, coriaceous, the 
daxial surfaces glabrous, lustrous, the abaxial sur- 
ices tomentose, usually 1 -nerved, apically mu- 
ronate, the margins incrassate. Capitulescences 
f solitary heads, terminal and axillary. Capitula 
dth involucres campanulate, 28-30 mm high, 20- 
5 mm wide; phyllaries 7-9-seriate, the outer tri- 
ngular-ovate, ca. 4 mm long, ca. 4 mm wide, the 
iner linear-lanceolate, 25-29 mm long, 1.4-1.8 
im wide, apex spinulose, pubescent; florets 18- 
0, the corollas yellow, 16-17 mm long, the tube 
.5-7 mm long, 2-2.2 mm wide, pubescent, the 
sbes linear-lanceolate, 4 lobes 6-7 mm long, equal, 
lobe 10-12 mm long, 0.8-1 mm wide at the base, 
cute, pubescent; anthers 1 1-12 mm long, the ter- 
linal appendage membranous, obtuse; styles 25- 
8 mm long. Achenes 3.3-3.5 mm long, 1 .8-2 mm 
/ide, pubescent, the hairs rigid, whitish then brun- 
ieus; pappus of 1 7-1 9 bristles, yellow-orange, 10- 
4 mm long. 

DISTRIBUTION Native of the interandean re- 
ion of central Peru (3600-3900 m). 

Chuquiraga johnstonii most closely resembles 
7. spinosa but differs in possessing stems with 
nfrequent, small spines to only 1 mm long, leaves 
onspicuously tomentose beneath, yellow capitula 
nd restricted distribution in the south-central An- 
lean Cordillera. Chuquiraga weberbaueri also has 
ericeous leaves, but it is readily distinguished from 
7. johnstonii by its much larger leaves and capit- 
ila. 



HUANCAVELICA. Huancavelica: near Vichi, Proano 
1 9 (USM). Tayacaja: 3 km N of Salcabamba Village, Stork 
& Morton 10329 (F). HUANUCO. Ambo: km 324 Hua- 
nuco to Cerro de Pasco, Duncan et al. 2702 (F, MO); 
Quinna, Sawada 84 (F). PASCO. Pasco: cerca a Salca- 
chupan, entre Cerro de Pasco y San Rafael, Ferreyra 
6598 (F, MO, USM); La Quinua and Chicrin, Ellenberg 
3769 (us), 3773 (us); Gentry et al. 19222 (MO); 95 km S 
from Huanuco, on road to Pasco, Gentry et al. 37501 
(F, MO). 

2. Chuquiraga jussieui J. F. Gmelin, Syst. Nat. 
1205. 1796. TYPE: Peru, without exact lo- 
cality, /. de Jussieu, herb. A. L. de Jussieu 
8320 (holotype, P-JU, not seen). 

C. peruviana]. St. Hil., Fam. Nat. 1: 394. 1805. TYPE: 
not seen. 

C. pseudoruscifolia Muschl., Hot. Jahrb. Syst. 50, Beibl. 
1 1 1: 93. 1913. SYNTYPES: Peru, Puno, Sandia, 
Cuyocuyo, A. Weberbauer 936 (syntype, B, pre- 
sumably destroyed; isosyntype, MOL; photograph 
ex B, FM neg. 15847); Puno, A. Weberbauer 500 
(B, presumably destroyed); Ancash, Cajatambo, 
Ocros, A. Weberbauer 2805 (B, presumably de- 
stroyed); La Oroya, A. Weberbauer s.n. (B, pre- 
sumably destroyed). 

[For additional synonymy, consult Escurra (1985).] 

Shrubs 6-12 dm high, much-branched, the 
branches pubescent, hairs flexuose. Leaves alter- 
nate, subsessile; blades ovate-lanceolate, 5-1 5 mm 
long, 3-7 mm wide, glabrous, coriaceous, lustrous, 
mucronate, the margins incrassate, uninerved, the 
midrib prominent. Capitulescences of solitary, 
sessile, terminal heads. Capitula with involucres 
campanulate, 3450 mm high, 15-22 mm wide; 
phyllaries 8-12-seriate, orange, the outer elliptic- 
lanceolate, 8-18 mm long, ca. 3 mm wide, the 
inner linear-lanceolate, 22-32 mm long, 1.5-1.8 
mm wide, yellowish acute, spinulose, pubescent; 
florets 2 1-40, the corollas yellow, 1 7-22 mm long, 
the tube 7-9 mm long, 1.4-1.5 mm wide, pubes- 
cent, the lobes acute, pubescent; anthers 11-12 
mm long; styles 21-22 mm long, brown-orange, 
sericeous, glandular. Achenes 3-5 mm long, 1 .2- 



FIELDIANA: BOTANY 



1.5 mm wide, dense pubescent; pappus of 20-22 
bristles, yellowish, 14-15 mm long. 



DISTRIBUTION Known from the paramos of 
Colombia and Ecuador, jalca formations in north- 
ern Peru, and the drier puna formations of central 
and southern Peru and Bolivia (3100-4500 m). 

Escurra (1985) did not lectotypify C. pseudo- 
ruscifolia, but the only likely extant element is 
Weberbauer 936 (MOL), since all others were pre- 
sumably destroyed at B. 



AREQUIPA. Condesuyos: Chuquiribamba, Hartweg 
18 (NY). CUSCO. Quispicanchi: Marcapata-Cusco, 1 1 
km from peak, Aronson & Berry 540 (F); Marcapata, 
Stafford 991 (F). LAMBAYEQUE. Ferreflafe: 7 km NW 
of Incahuasi, Dillon & Skillman 4139 (F); Huasicaj, In- 
cahuasi, Llatas 1333 (F). LA LIBERTAD. Pataz: Tira- 
gra, Young 3059 (F). PIURA. Huancabamba: San An- 
tonio-Talaneo, Sagdstegui & Cabanillas 8631 (F). PUNO. 
Azangaro: Muiiani, Macedo s.n. (USM). Chuchuito: Juli, 
Soukup231 (F). Huancane: Miajachi-Huancane, Aguilar 
s.n. (MO); near Huancane, Shepard 85 (NY). Puno: Cerro 
Putina, Soukup 420 (F, GH). Sandia: Limbani, Stafford 
1111 (F). 



3. Chuquiraga rotundifoliaWedd., Chlor. And. 1: 
4. t. 4A. 1855. LECTOTYPE (designated by 
Escurra, 1985): Peru, "sommet des Andes Ta- 
cora," d'Orbigny s.n. (lectotype, P, not seen; 
photograph ex P, FM neg. 52102). Figure 6. 

Chuquiraga spinosa subsp. rotundifolia (Wedd.) Es- 
curra, Darwiniana 26: 242. 1985. 

Shrubs 4-1 2 dm high, branched, upper branches 
pubescent, spiny, spines divaricate, 4-15 mm long, 
yellowish. Leaves opposite, sessile; blades round 
to oval, 4-14 mm long, 3.5-10 mm wide, coria- 
ceous, entire, uninerved, acuminate, the adaxial 
surfaces slightly pubescent, the abaxial surfaces 
densely pubescent. Capitulescences of solitary 
heads or aggregates of 2-3, sessile heads. Capitula 
with involucres turbinate, 34-40 mm high, 7.5- 
9.5 mm wide; phyllaries 5-8-seriate, the outer 
ovate-lanceolate, 5-9 mm long, ca. 4 mm wide, 
brunneous, mucronate, pubescent outside, the in- 
ner linear-lanceolate, 26-31 mm long, 2-3 mm 
wide, apex spinulose; florets 10-16, the corollas 
yellow, 19-22 mm long, the tube 7-9 mm long, 
1.4-1.8 mm wide, pubescent, the lobes linear-lan- 
ceolate; anthers 12-14 mm long, apex obtuse; styles 
28-35 mm long. Achenes 3-4 mm long, 1.4-1.5 
mm wide, densely pubescent; pappus of 17-21 
bristles, yellowish, 15-18 mm long. 



DISTRIBUTION Known from the puna of south- 
ern Peru and adjacent northern Chile and Bolivia 
(3000-4000 m). 

Chuquiraga rotundifolia is characterized by its 
round or oval leaves, turbinate involucres, and 
brown, mucronate, 5-6-seriate phyllaries. 

AREQUIPA. Arequipa: entre Chihuata y la Cumbre, 
Ferreyra 2592 (F, MO, USM); S slopes of Nevado Chacha- 
ni, on road to Sumbay, Hutchison & Wright 7234 (F, 
MO); 26 mi out of Arequipa on road to Puno, Maguire 
& Maguire 44451 (F, MO, NY); above Arequipa, Pennell 
13214 (F, GH, NY); Nevado de Chachani, Pennell 13271 
(F, GH, NY); Lopez 131 (us); Rauh 40534 (USM); West 
7135 (MO). Condesuyos: above Salamanca, Weberbauer 
6858 (F, MOL, us, USM). AYACUCHO. Lucanas: 83 km 
W of Puquio, 76 km E of Nazca, Gentry et at. 23271 (F); 
Puquio, Rauh & Hirsch P427 (NY), P428 (NY, USM). MO- 
QUEGUA. Mariscal Nieto: cordillera above Torata, 
Weberbauer 4778 (F). TACNA. Tarata: 16 km on trail 
from Candarave to Puno, Metcalf 30394 (MO, us); 4 km 
NE of Tarata, Pearson 31 (F). Without locality data, 
Rauh 526 (USM). 

4. Chuquiraga spinosa Less., Linnaea 5: 259. 1 830. 
LECTOTYPE (designated by Escurra, 1985): 
Peru, without exact locality, J. Dombey 447 
(lectotype, P, not seen). Non Chuquiraga spi- 
nosa (Ruiz Lopez & Pa von) D. Don, 1830. 

Chuquiraga spinosa subsp. huamanpinta Escurra, 
Darwiniana 26: 243. 1985. TYPE: Peru, Lima, 
"bei Ticlio Bajo, ostl. v. Lima," L. Diers 946 
(holotype, LP, not seen). 



Shrubs 4-1 5 dm high, branched, upper branches 
finely pubescent, spiny, the spines axillary, yellow- 
ish, divaricate, 12-18 mm long. Leaves opposite, 
sessile; blades ovate-lanceolate, 4.5-20 mm long, 
2.6-5 mm wide, pubescent, then glabrescent, lus- 
trous, uninerved, midrib prominent, attenuate at 
base, apex spinulose, entire, the margins incras- 
sate. Capitulescences of solitary heads, terminal 
and axillary. Capitula with involucres 32-45 mm 
high, 1 2-24 mm wide, campanulate; phyllaries 8- 
10-seriate, deep red-orange, the outer lanceolate, 
apically rounded-mucronate to acute-spinulose, 
densely pubescent, the inner linear-lanceolate, 25- 
30 mm long, 1.8-2.8 mm wide, pubescent; florets 
9-14, the corollas yellowish-orange, 19-22 mm 
long, the tube 6.5-8 mm long, 1.7-2.2 mm wide, 
pubescent, the lobes 12-14 mm long, acute, pu- 
bescent; anthers 12.5-15 mm long, obtuse, sagit- 
tate; styles 30-34 mm long, deep red-orange. 
Achenes 3-4 mm long, 1.9-2.2 mm wide, densely 
pubescent; pappus of 1 5-2 1 bristles, yellowish, 1 3- 
20 mm long. 



FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI. 



29 




FIG. 6. Chuquiraga rotundifolia. 1, habit; 2, floret; 3, achene with pappus. (Drawn from P.C. Hutchison & J.K. 
right 7234, F.) 

DISTRIBUTION Known from the Andean Cor- glabrescent leaves, and yellowish-orange florets 

Hera of central Peru (3500-4800 m). distinguish it from the other related species. 
Chuquiraga spinosa is one of the most common 

id characteristic species in the Central Puna of ANCASH. Bolognesi: near Carpa, Antunez 332 (USM); 

:ru. The dense spiny habit, conspicuous rigid and Pampa de Lampas, Cerrate 1511 (USM); between Tal- 



FIELDIANA: BOTANY 



lenga and Pachapaque, Ferreyra 7473 (USM); 10 km E 
of Catac, Gentry et al. 37460 (F, MO, USM); Pachacoto, 
Tovar & Rivas 9618 (USM); between Ocros and Paso de 
Chonta, Weberbauer 2805 (USM). Huaras: near Huaras, 
Bernardi et al. 16605 (F, us). Recuay: arriba de Carpa, 
Lopez et al. 8327 (F); Huascaran National Park, Todzia 
et al. 2600 (F). Yungay: Parque Nacional Huascaran, 
Zardini 1561 (MO). APURIMAC. Andahuaylas: Cora- 
cora-Andahuaylas, Weberbauer 5834 (f). AYACUCHO. 
Huanta: N of Huanta, near Huaillay, Weberbauer 7593 
(F). CUSCO. Urubamba: Chinchero, Sallo & Franque- 
mont 276 (F). HUANCAVELICA. Huancavelica: near 
Conaica, Tovar 740 (USM); Chassca, above Conaica, Fer- 
reyra & Tovar 8008 (USM). JUNIN. Huancayo: near 
Huancayo, Killip & Smith 22135 (F); Hda. Acopalca, 
Ochoa 637 (F, GH); Acopalca, Soukup 3683 (F, us), 3691 
(F, us); between Acopalca and Pariahuanca, Stork 10926 
(F, uc). Jauja: above Chicchi, Ochoa 29 (USM). Junin: 
entre Carhuamayo y Junin, Ferreyra 5237 (F, MO, USM); 
Ondores, Johns 81-3 (F); Lago Junin, Landrum 4629 
(NY); 3 mi N of Junin on road to Cerro de Pasco, Maguire 
& Maguire 44401 (F, NY, USM). Tarma: between Tarma 
and Huancayo, Anderson 1045 (us); San Pedro de Cajas, 
Antunez 26 (F, USM); 1 1 km W of Tarma, Dillon & Turner 
1387 (F); between Tarma and the Cumbre, Ferreyra 3780 
(USM); Pampa de Junin, Ferreyra 3945 (USM); Logo Ju- 
nin, Hutchison et al. 5886 (F, K, MICH, MO, NY, us, USM); 
between Tarma and Jauja, Iltis & Ugent 153 (us); above 
Tarma, Infantes 637 (USM); between Oroya and Tarma, 
Palmer 94 (USM); Huacapa, Smith et al. 1395 (F); Laguna 
Parinacochas, Tovar 423 (MO); Tarmatambo, Tovar 2334 
(USM); Incatacuna, Tovar 1081 (MO, USM); Acobamba, 
Woytkowski 53 (F). Yauli: above Oroya, Cerrate 91 4 (MO, 
USM); Lima-Oroya, Davidson 9010 (MO); 1 km W of 
Anticona Pass, Dillon & Turner 1482 (F); 3 km from 
Yauli, Duncan et al. 2651 (F); near Morococha, Grant 
7567 (F); near Oroya, Kalenborn 34 (GH, us, MO); near 
Yauli, Soukup 3999 (F, USM); between Tarma and Huan- 
cayo, Sullivan et al. 1050 (USM); Towsend 1523 (us); 
Tucto, near Morococha, Weberling 5909 (USM). LIMA. 
Canta: near La Viuda, Meza 1 97 (MO, USM); above Canta, 
Sanchez 41 (USM); Mishquipuquio, Sanchez 40 (MO, USM); 
Chuchun, Cordillera La Viuda, Meza s.n. (USM). Huaro- 
chiri: km 94, near San Mateo, Ferreyra 5322 (USM); above 
Chicla, Ferreyra 6904 (USM); Casapalca, Ferreyra 8009 
(USM); 20 km NE of San Mateo, Gentry et al. 19184 (F, 
MO); 10 km NE of Suchi, Gentry 21678 (F, MO, USM); 
NW of Ticlio, Gentry & Smith 44837 (F); Simpson 8558 
(USM); Maisch s.n. (USM); between San Mateo and Ticlio, 
Hjerting & Petersen 1283 (MO, USM); near Huarochiri, 
Hrdlica s.n. (us); Viso, Macbride & Featherstone 632 (F, 
GH); near Escomarca, Ferreyra 19090 (USM). Yauyos: Co- 
chapunco, above Tupe, Cerrate 1215 (MO, USM); 17 km 
de Tupe, Cerrate & Tovar 1234 (F, USM); Omas, Cerro 
Quilca, entre Tamara y Ayaviri, Ochoa & Solas 14666 
(F, us); Huacracocha, near Tupe, Tovar 650 (USM); above 
Tupinacha, Tovar 632 (USM). PASCO. Pasco: La Viuda, 
Ellenberg 4064 (us); Bosque de Piedras, Urquizo 27 (USM). 
Without locality data, Rauh PI 7 58 (USM); Peruvian An- 
des, Capt. Wilkes Exped. s.n. (GH). 

5. Chuquiraga weberbaueri Tovar, Publ. Mus. 
Hist. Nat. "Javier Prado," Ser. B, Bot. 5: 9. 
1952. TYPE: Peru, La Libertad, Bolivar, cer- 
ca Nevado Cajamarquilla, Sep 1946, 4200- 



4600 m, R. Ferreyra 1304 (holotype, F 
1488740; iso types, MO, USM). 

Shrubs 8-15 dm high, branched, the branches 
pubescent. Leaves opposite, sessile; blades elliptic- 
lanceolate, 6-28 mm long, 3-8.5 mm wide, cori- 
aceous, apically mucronate, the adaxial surfaces 
lustrous, the abaxial surfaces sericeous, uninerved, 
the margins entire, incrassate. Capitulescences of 
solitary, terminal heads. Capitula with involucres 
campanulate, 28-45 mm high, 15-30 mm wide; 
phyllaries 6-7 -seriate, orange-yellowish, the outer 
triangular-ovate, 4-6 mm long, ca. 4 mm wide, 
the inner linear-lanceolate, 27-33 mm long, 2-2.5 
mm wide, sericeous, spinulose, abaxial surfaces 
pubescent; florets 2045, the corollas yellow, 17- 
23 mm long, the tube 5-7 mm long, 1.6-2 mm 
wide, pubescent, the lobes 1 1.5-16 mm long, acute, 
abaxial surfaces pubescent; anthers 10.5-15 mm 
long; styles 25-33 mm long, red-orange. Achenes 
3-4 mm long, 1.8-2.2 mm wide, densely pubes- 
cent, brunneous; pappus of 17-18 bristles, brun- 
neous, 14-18 mm long. 

DISTRIBUTION Endemic to the jalca forma- 
tions of northern Peru (3200-4600 m). 

Chuquiraga weberbaueri is characterized by its 
dense, nodose branches; sessile, sericeous, tomen- 
tose leaves; and campanulate involucres with 
phyllaries 6-7-seriate. 

CAJAMARCA. Cajamarca: cerca Cajamarca, San- 
chez 1146 (CPUN). LA LIBERTAD. Huamachuco: Mon- 
tana de Huaylillas, Weberbauer 7009 (F, GH, MOL); West 
8128 (GH). 



VII. DASYPHYLLUM 

Dasyphyllum Kunth, Nov. Gen. Sp. 4: 13. (fol.) 
1 8 1 6, 1 7 (quarto). 1 820. TYPE: D. argenteum 
Kunth 

Flotovia Spreng. Syst. Veg. 3: 359. 1826. TYPE: F. 
glabra Spreng. 

Trees or shrubs, erect to scandent, with or without 
axillary spines. Leaves alternate, shortly petiolate; 
blades lanceolate to oblong or oval, coriaceous, 3- 
5 -nerved, the margins entire. Capitulescences of 
solitary heads or racemose or glomerulate, ter- 
minal or axillary. Capitula homogamous; invo- 
lucres campanulate or turbinate; receptacles plane, 
pubescent with bristles or scales, epaleate; phyl- 
laries 5-8-seriate, imbricate, the outer lanceolate, 



FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI. 



31 



nucronate, the inner lanceolate, sericeous; florets 
5-many, hermaphroditic, the corollas tubular, 
.vhite to violet or rarely yellow, usually isomorphic 
Dr slightly dimorphic, usually villous, the limb 
5-lobed; anthers 5, basally sagittate, terminal ap- 
pendage bifid to emarginate, rarely truncate; styles 
iliform, bifid. Achenes cylindric, densely villous 
:o glabrous; pappus of plumose bristles, 1 -seriate. 

Dasyphyllum consists of approximately 37 spe- 
:ies of shrubs or small trees restricted to South 
\merica from the Andean Cordillera of Colombia 
md Venezuela to eastern Brazil and central Chile. 
Die habit of Dasyphyllum is very close to Chu- 
juiraga; it can be distinguished from that genus, 
lowever, in having three- or five-nerved leaves, 
:he stamens united at the lower part of the tube 
)f the corolla, the anthers slightly sagittate with 
:he terminal appendage bifid or truncate, the flo- 
ets with white, pale violet, or yellowish corollas, 
md tricolpate pollen grains. It is readily distin- 



guished from Barnadesia, which possesses gener- 
ally reddish corollas, acute anther appendages, and 
lophate pollen. In addition, Dasyphyllum finds its 
greatest diversity in the subtropical region east of 
the Andean Cordillera, while Chuquiraga and Bar- 
nadesia find their greatest diversity within the Cor- 
dillera. 

Although Tovar (1953) recognized Flotovia as 
a distinct genus, Cabrera (1959) combined it with 
Dasyphyllum, and it is not accepted in the present 
treatment. 



References 

CABRERA, A. L. 1959. Revision del genero Dasyphyl- 
lum (Compositae). Revista Mus. La Plata, Secc. Bot. 
9: 21-200. 

TOVAR, O. 1953. Las especies peruanas del genero Flo- 
tovia. Publ. Mus. Hist. Nat. "Javier Prado," Ser. B, 
Bot., 7: 1-12. 



tey to Peruvian Species of Dasyphyllum 

. Corolla apices velutinous 2 

2. Stems without spines 3 

3. Leaves glabrescent, the blades obovate-elliptic, 20-35 mm long, 10-18 mm wide; capitula with 
(1 1-)12(-13) florets D. cabrerae 

3. Leaves pubescent, the blades ovate-oblong, ca. 33 mm long, ca. 14 mm wide; capitula with 
1 3-1 5 florets D. weberbaueri 

2. Stems with paired, divergent, straight or curved spines 4 

4. Spines longer than leaves, 25-50 mm long D. ferox 

4. Spines greatly reduced, 3-5 mm long 5 

5. Stems velutinous, spines straight, 3-5 mm long; capitulescences racemose 

D. brevispinum 

5. Stems glabrous, spines strongly curved proximally, ca. 5 mm long; capitulescences cymose- 

paniculate D. brasiliense 

. Corolla apices glabrous 6 

6. Capitula with (1 1-)1 5-20(-30) florets; achenes glabrous D. hystrix 

6. Capitula with 6-10 florets; achenes pubescent 7 

7. Leaf blades elliptic lanceolate, generally 15-30 mm long, 4-12 mm wide; involucres turbinate, 

1 1-1 3 mm long; florets 6 D. leiocephalum 

7. Leaf blades ovate-elliptic, generally 5-15 mm long, 3.5-5 mm wide; involucres campanulate, 
8-1 1 mm long; florets 7-10 . . D. horrida 



. Dasyphyllum brasiliense (Spreng.) Cabr., Re- 
vista Mus. La Plata, Secc. Bot. 9: 74. 1954. 
Figure 7. 

Joannea brasiliensis Spreng., Neu. Entd. 2: 132. 1821. 
TYPE: Brazil, "Brasilia aequinoctialis," Sello s.n. 
(holotype, p, not seen; photograph ex p, FM neg. 
15838). 



Dasyphyllum brasiliense is composed of some 
five varieties differing primarily in their leaf shape 
and size, pubescence type, and number of florets 
per capitula (Cabrera, 1959). Peruvian material is 
referable to following variety. 

la. Dasyphyllum brasiliense var. barnadesioides 



FIELDIANA: BOTANY 




smm 



FIG. 7. Dasyphyllum brasiliense. 1, habit; 2, capitulum; 3, achene with pappus. (Drawn from Marin 1567B, F.) 



(Tovar) Cabr., Revista Mus. La Plata, Secc. 
Bot. 9: 76. 1954. 

Flotovia barnadesioides Tovar, Publ. Mus. Hist. Nat. 
"Javier Prado," Ser. B, Bot. 7: 10. 1953. TYPE: 
Peru, Junin, Tarma, Colonia Perene, 9 Sep 1920, 



800-900 m, A. Weberbauer s.n. (holotype, USM 
10322). 

Shrubs slightly voluble, 1-1 .4 m high, branched, 
the branches tomentose, axillary spines curved 
proximally, ca. 5 mm long. Leaves lanceolate-el- 



FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI. 



33 



ptic, 20-92 mm long, 7-32 mm wide, mucronate, 
inerved, pubescent, the margins revolute. Capit- 
lescences cymose-paniculate, axillary. Capitula 
1-12 mm high, 6-7 mm wide; involucres cam- 
anulate; phyllaries 7-8-seriate, the outer ovate, 
.5-2.5 mm long, 1-1.5 mm wide, acuminate, 
ensely pubescent, the inner lanceolate-linear, 9.5- 

mm long, 1.2-1.4 mm wide, mucronate, pu- 
escent; florets 10-12, the corollas 7.5-7.8 mm 
>ng, the tube 3-3.2 mm long, yellowish; anthers 
.3-2.4 mm long; styles 9-9.5 mm long, papillose 
t apex. Achenes 2-2.4 mm long, campanulate, 
ubescent; pappus of 16-17 bristles, 7-7.5 mm 
>ng. 

DISTRIBUTION Known from the subtropical re- 
ion of the eastern slope of the Andean Cordillera 
f central Peru to Bolivia (800-1800 m). 

Dasyphyllum brasiliense var. barnadesioides is 
istinguishable from the typical variety by pubes- 
jnt leaves and is distinctive among Peruvian spe- 
cs in possessing small downwardly curved spines, 
he other varieties are restricted to lower eleva- 
ons in Argentina, Brazil, and Paraguay (Cabrera, 
959). 

USCO.