UNIVERSITY OF
ILLINOIS LIBRARY
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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-
blance—for 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, 3.5—4 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 35—42
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, 18—42 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, 4.5—4.8 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,
20—40 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, 3—4 mm wide, acuminate, mucronate
or spinulose, densely pubescent outside; florets ca.
91, the corollas orange, 32-33 mm long, the tube
3.5—4 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
(600—4100 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, 26—49 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, 3.8—4.5 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, 3—4 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 40—45 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, 3.8—4.2 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 0 1 3 1 97 ex G, F 974790 ex
P).
Shrubs 1 8—40 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, 34—42 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, 34—35 mm long,
1.2-2 mm wide, apically acute; ray florets 13, the
corolla pink, 46—48 mm long, the tube 33-35 mm
long, the outer lip 12-16 mm long, 3—4 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,
3—12 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,
25—48 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, 27—40 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 4—5-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, 3—4 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,
1.4—1.6 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 14—16 mm long; [disc] pyriform, 3—4 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, 4—4.2 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
3.6—4.2 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, 26—42 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, 1.5—2
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, 3.5—4 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 3.5—4.5
mm long, 1.5-2.8 mm wide, pubescent, the limb
5-toothed; anthers 6-6.5 mm long. Achenes [ray]
2.5—3 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 1.5—4 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 38—46 mm long, the tube
25-29 mm long, 0.8-1 mm wide, pubescent, the
outer lip 12-16 mm long, 3.8—4.8 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, 38—40 mm long,
2.3-2.5 mm wide, membranaceous, acute, mucro-
nate, sericeous; ray florets 13, the corollas pink,
50—51 mm long, the tube 31-32 mm long, 1—1.2
mm wide, pubescent, the outer lip 20-2 1 mm long,
3.8—4 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 14—18 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 42°S latitude in Chile and Argentina. Only
four species are recorded from Peru; all tend to
occur at high elevations (2700—4900 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 4—4.2 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 3.8—4.2 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 4—4.5 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, 3—4 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°-34°S 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, 2—4 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 0 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, 3.5—4 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. 20—40 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 3—4 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 4—6 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 20—45
. 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-
0 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 0 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, 34—50 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 20—45, 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-
0 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. Convention: Hacienda Choquellohuanca, Mar-
1 7567 (F).
. Dasyphyllum brevispinum Sagast. & Dillon,
Brittonia 37: 6. 1985. TYPE: Peru, Cajamar-
ca, desvio Bosque de Cachil (Cascas-Contu-
maza), 25 Jun 1982, 2200-2300 m, A. Lopez
M.,A. SagdsteguiA., J. Mostacero, & S. Lopez
9136 (holotype, HUT; isotypes, F 1964136, G,
MO, NY, us).
Shrubs to 3 m tall, much-branched; stems erect,
glutinous, axillary spines paired, 3-5 mm long,
ivergent, straight. Leaves with petioles 2-3 mm
mg, velutinous; blades elliptic, 30—45 mm long,
5-20 mm wide, coriaceous, both surfaces densely
glutinous, basally acute, apically obtuse to round-
I. Capitulescences racemose on axillary bran-
ilets, 3-8 -headed, often glomerulate distally. Ca-
itula sessile to subsessile, 5-7 mm high, 5-6 mm
ide; involucres cylindric to narrowly campanu-
ite; phyllaries ca. 7-seriate, the outer narrowly
vate, 3-4 mm long, 2-3 mm wide, coriaceous,
elutinous, ciliolate, obtuse, spinulose, the inner
near to lanceolate, 7-8 mm long, 1-1.5 mm wide,
jriaceous, velutinous, acute to obtuse, spinulose;
orets 13-16, the corollas whitish, ca. 5 mm long,
ic lobes ca. 1.5 mm long, narrowly lanceolate,
apically velutinous; anthers ca. 1.7 mm long.
Achenes cylindric, ca. 2 mm long, sericeous to
velutinous; pappus ca. 5.5 mm long, yellowish-
brown.
DISTRIBUTION— Known only from the type lo-
cality.
The description presented here was taken from
the original description.
3. Dasyphyllum cabrerae Sagast., Bol. Soc. Ar-
gent. Bot. 19: 61. 1980. TYPE: Peru, Ama-
zonas, Bongara, distrito Pomacocha, El In-
genio (km 12), 1500 m, 28 Jul 1970, A. Diaz,
L. Vdsquez, & L. Cerna 1105 (holotype, HUT;
isotype, PRO).
Shrubs to 2 m; stems sericeous, lacking spines.
Leaves petiolate; petioles 1-2 mm long; blades ob-
ovate to elliptic, 20-35 mm long, 10-18 mm wide,
coriaceous, trinervate, the abaxial surfaces laxly
pubescent, the adaxial surfaces glabrous, basally
attenuate, apically rounded to obtuse, mucronu-
late. Capitulescences racemose, terminal and ax-
illary. Capitula sessile to subsessile, ca. 9 mm high,
ca. 5 mm wide; involucres narrowly campanulate;
phyllaries 5-6-seriate, the outer ovate, 3-5 mm
long, coriaceous, acute, mucronate, tomentose, the
inner linear-lanceolate, 7-9 mm long; florets (11-
12(-13), the corollas ca. 5 mm long, the limb gla-
brous. Achenes (immature) cylindric, ca. 2 mm
long, velutinous; pappus of 15-18 bristles, ca. 5
mm long.
DISTRIBUTION— Known only from the type lo-
cality.
The description presented here was taken from
the original description.
4. Dasyphyllum ferox (Wedd.) Cabr., Revista Mus.
La Plata, Secc. Bot. 9: 54. 1959.
Flotovia ferox Wedd. Chlor. And. 1: 5. 1855. LEC-
TOTYPE (here designated): Bolivia, La Paz,
D'Orbigny 1533 (lectotype, p, not seen; photo-
graph of isolectotype ex w, FM neg. 33314).
Chuquiraga ferox (Wedd.) Britton, Bull. Torrey Bot.
Club 19: 266. 1892.
Shrubs to 3 m high, branched, branches pubes-
cent, axillary spines 10-40 mm long, divaricate.
Leaves oblong-lanceolate, 7-32 mm long, 6-24
mm wide, glabrescent, trinerved, subsessile. Cap-
itulescences glomerulate, terminal. Capitula sub-
sessile, 11-13 mm high, 4-5 mm wide; involucre
FIELDIANA: BOTANY
turbinate-campanulate; phyllaries 6-7-seriate, the
outer ovate-lanceolate, 2-4 mm long, 1.5-2 mm
wide, apiculate, the inner lanceolate, 10-11 mm
long, 1 .2-1 .4 mm wide, mucronate, pubescent; flo-
rets 10-19, the corollas white, 8-10 mm long, the
tube 3.5-5 mm long, the lobes 4.5-5 mm long,
linear; anthers 3-4 mm long, the terminal ap-
pendage apically bifid; styles 9-13 mm long, pa-
pillose. Achenes 4-5 mm long, cylindric; pappus
of 1 8-20 bristles, 7-9 mm long, whitish-yellow.
DISTRIBUTION— Known from throughout the
Andean Cordillera of Peru to Bolivia (3000-3700
m).
Dasyphyllum ferox most closely resembles D.
armatum (Koster) Cabr., a native of southern Bo-
livia and northern Argentina, but D. ferox differs
in possessing oblong-elliptic leaves, densely pu-
bescent phyllaries, and pubescent corolla apices.
Cabrera (1959) cited the two collections from
WeddelFs protolog as "cotypus" but did not choose
a lectotype collection. Here, the lectotype speci-
men of d'Orbigny was chosen over Weddell's un-
numbered collection (F 970337 ex p).
ANCASH. Recuay: Recuay, Macbride 2513 (F, GH,
us); Ames 38 (USM); Lopez et al. 7488 (F, MO). Huaras:
Puente Bedoya, Proano 32 (USM). Bolognesi: Chiquian,
Ferreyra & Cerrate 7335 (USM). CUSCO. Canchis: Si-
cuani, Vargas 930 (F, MO, USM), 9834 (F, GH, uc). Uru-
bamba: near Urubamba, Solomon 3091 (MO); Vilcanota
Valley, Cook & Gilbert 101 (us). JUNIN. Tarma: Cerro
San Sebastian, Ferreyra 510 (USM); Macbride & Feath-
erstone 1028 (GH, us); 1 km W of Tarma, Stork 10944
(F, uc). LA LIBERT AD. Otusco: Puente Chiquin, Lopez
& Sagdstegui 8065 (F, MO). Santiago de Chuco: Angas-
marca, Lopez 1121 (us); 70 km N of Agallpampa, Lopez
1128 (USM). LIMA. Canta: San Buenaventura, Pennell
14509 (F); near Canta, Sanchez 019 (USM); Obrajillo,
Capt. Wilkes Exped. s.n. (GH, us).
5. Dasyphyllum horridum (Muschl.) Cabr., Re-
vista Mus. La Plata, Secc. Bot. 9: 49. 1959.
Chuquiraga horrida Muschl., Bot. Jahrb. Syst. 50,
Beibl. 111:92. 19 13. TYPE: Peru, Apurimac, An-
Key to Peruvian Varieties of Dasyphyllum hystrix
1. Leaf blades elliptic-lanceolate, 10-25 mm long, 2-6 mm wide D. hystrix var. hystrix
1. Leaf blades narrowly lanceolate, 25-30 mm long, 9-12 mm wide D. hystrix var. peruvianum
dahuaylas, Hacienda Cotahuacho,^. Weberbauer
5847 (holotype, B, presumably destroyed; iso-
types, F, GH, us).
Flotovia horrida (Muschl.) Tovar, Publ. Mus. Hist.
Nat. "Javier Prado," Ser. B, Bot. 7: 7. 1953.
Shrubs 1-3 m high, branched, branches some-
times ferruginous, spines axillary, divaricate, 4-
1 7 mm long. Leaves with blades ovate-elliptic, 5-
15 mm long, 3.6-5 mm wide, slightly pubescent,
trinerved. Capitulescences of solitary heads or small
aggregates, subsessile, terminal or axillary. Capit-
ula with involucres 11-12 mm high, 4-^4.5 mm
wide, campanulate or turbinate; phyllaries 5-6-
seriate, the outer ovate, 1-2 mm long, 1.5-2 mm
wide, the inner lanceolate, 8-9 mm long, mucro-
nate, pubescent; florets 7-10, the corollas 7-8 mm
long, the tube 3-5 mm long, the lobes 4 mm long,
linear-lanceolate, slightly coiled; anthers 3-3.2 mm
long, the terminal appendage bifid; style 9.5-10.5
mm long, papillose. Achenes 3—4 mm long, cylin-
dric, densely pubescent; pappus of 22-24 bristles,
ca. 6.2 mm long.
DISTRIBUTION— Endemic to the Andean Cor-
dillera of southern Peru (2600-3200 m).
Dasyphyllum horridum is strongly spiny and
most closely resembles Dasyphyllum ferox, but it
differs in its smaller, ovate-elliptic leaves, capitula
with fewer florets (7-10), and glabrous corolla lobes.
CUSCO. Calca: near Calca, Vargas 708 (MO).
6. Dasyphyllum hystrix (Wedd.) Cabr., Revista
Mus. La Plata, Secc. Bot. 9: 51. 1959.
Flotovia hystrix Wedd., Chlor. And. 1 : 6. pi. 3B. 1 855.
TYPE: Bolivia, La Paz, Lagunillas, D'Orbigny
1381 (holotype, P, not seen).
Dasyphyllum hystrix contains two varieties based
on leaf shape and size.
6a. Dasyphyllum hystrix var. hystrix.
Shrubs to 2 m high, branched, branches pubes-
cent, spines to 25 mm long, divaricate. Leaves
elliptic-lanceolate, 10-25 mm long, 2-6 mm wide,
trinerved. Capitulescences of solitary heads, ter-
minal. Capitula sessile; involucres globose; florets
ca. 20, the corollas white, glabrous; anthers with
terminal appendages bifid. Achenes glabrous.
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
35
DISTRIBUTION— Known from the Andean Cor-
lillera of southern Peru to central Bolivia (3400-
J700 m).
CUSCO. Convention: Rosariomayo, Vargas 20693
us).
>b. Dasyphyllum hystrix var. peruvianum Cabr.,
Revista Mus. La Plata, Secc. Bot. 9: 52. 1959.
TYPE: Peru, La Libertad, Santiago de Chuco,
alrededores de Santiago de Chuco, A. Lopez
Miranda 980 (holotype, LP; isotype, USM).
DISTRIBUTION— Known from intermontane
egions of northern Peru (3000-3100 m).
This variety is quite similar to variety hystrix,
>ut it is distinguished by possessing narrowly lan-
;eolate, 5-nerved leaf blades, 25-30 cm long, 9-
12 mm wide.
LA LIBERTAD. Santiago de Chuco: Agua de Loro,
sdpez Miranda 1961 (HUT, USM).
. Dasyphyllum leiocephalum (Wedd.) Cabr., Re-
vista Mus. La Plata, Secc. Bot. 9: 50. 1959.
Flotovia leiocephala Wedd., Chlor. And. 1: 6. 1855.
TYPE: Peru, Cusco, Gay 1220 (holotype, p, not
seen; isotype fragment ex p, F 971917).
Shrubs 15-25 dm high, branched, spiny, spines
-16 mm long, axillary, divaricate. Leaves alter-
late, elliptic-lanceolate, (5-) 1 5-30 mm long, 4-12
nm wide, mucronate, trinerved, more or less pu-
•escent. Capitulescences of solitary heads, sessile,
xillary. Capitula with involucres 11-13 mm high,
L5-5 mm wide, turbinate-fusiform; phyllaries
(-seriate, brunneous, lustrous, the outer ovate-
anceolate 2-3 mm long, ca. 2 mm wide, the inner
anceolate, 1 1-1 1.5 mm long, acuminate, mucro-
late; florets 6, the corollas yellow, 6-8 mm long,
he tube 4.5-5 mm long, the lobes coiled; anthers
>.2-3.6 mm long, bifid; style filiform, 12-14 mm
ong, papillose. Achenes 4-5 mm long, cylindric,
lensely pubescent; pappus of 15-19 bristles, 6-7
nm long.
DISTRIBUTION— Endemic to the Andean Cor-
lillera of southern Peru (2800-3000 m).
Dasyphyllum leiocephalum suggests D. horri-
'um but differs in the sessile and solitary capitula
rith 6 florets, the turbinate involucre, and the
ustrous brunneous phyllaries.
APURIMAC. Andahuaylas: Chincheros, Ricci 5730
(USM); Stork & Morton 10737 (GH, uc). CUSCO. Calca:
San Salvador, Mann 2053 (USM). Quispicanchis: Hua-
carpay, Chavez 2018 (MO). Urubamba: Pumamanca,
Vargas 7602 (USM); Urubamba, Zamalloa 56 (USM); Var-
gas 11060 (uc). NO EXACT LOCALITY. Ellenberg 1004
(USM).
8. Dasyphyllum weberbaueri (To var) Cabr., Re-
vista Mus. La Plata, Secc. Bot. 9: 66. 1959.
Flotovia weberbaueri Tovar, Publ. Mus. Hist. Nat.
"Javier Prado," Ser. B, Bot. 7: 1 1. 1953. TYPE:
Peru, Piura, Ayavaca, Quiros, A Weberbauer6345
(holotype, GH; isotype, F, 2 sheets).
Shrubs to 3 m high, branched, branches striate,
pubescent. Leaves with blades ovate-oblong, 12-
33 mm long, 7-14 mm wide, rounded at the apex,
more or less pubescent. Capitulescences of aggre-
gate racemes. Capitula with involucres 9-10 mm
high, 5-6 mm wide, campanulate; receptacle plane,
pubescent; phyllaries 6-7-seriate, yellowish, the
outer ovate, 2-2.5 mm long, ca. 1.5 mm wide,
mucronate, pubescent, the inner lanceolate, 6.8-
7 mm long, 1-1.5 mm wide, spinulose, pubescent;
florets 13-15, the corollas 6.8-7.2 mm long, the
tube 3.4-3.6 mm long; anthers 3-3.5 mm long,
the terminal appendage bifid; styles 7-7.8 mm long,
papillose near apex. Achenes cylindric, 3-4 mm
long, pubescent; pappus of 16-19 bristles, ca. 5.2
mm long.
DISTRIBUTION— Endemic to the intermontane
region of northern Peru (2000-2200 m).
Dasyphyllum weberbaueri resembles D. cabrer-
ae in overall habit and Capitulescences, but it dif-
fers by having more pubescent ovate-oblong leaves
and capitula with 13-15 florets.
This species is rare and currently only known
from the type.
VIII. FULCALDEA
Fulcaldea Poiret, Encycl. Meth., Bot. Suppl. 5: 375.
1827. TYPE: Turpinia laurifolia Humb. &
Bonp. = Fulcaldea laurifolia (Humb. & Bonpl.)
Poiret
Shrubs or small trees to 8 m in height; trunks
to 25 cm in diameter, paired axillary spines usually
present. Leaves alternate, subsessile to petiolate;
blades ovate, triplinerved, the margins entire.
Capitulescences of corymbose or paniculate cymes,
FIELDIANA: BOTANY
terminal and axillary. Capitula homogamous with
only 1 floret; involucres cylindric; phyllaries 5-7-
seriate; floret hermaphroditic, actinomorphic,
5-merous, the corolla tubular, densely villous,
5-lobed; anthers ecaudate; styles exserted, apically
bifid, the distal portion swollen and papillose, the
branches rounded, glabrous. Achenes cylindric;
pappus of plumose bristles, 1 -seriate.
Fulcaldea consists of a single distinctive species
distributed from southern Ecuador to northwest-
ern Peru (650-2800 m). Although Marling (1991)
stated that the genus was endemic to Ecuador,
ample collections exist from northern Peru.
1 . Fulcaldea laurifolia (Humb. & Bonpl.) Poiret,
Encycl. Meth., Bot. Suppl. 5: 375. 1827. Fig-
ure 8.
Turpinia laurifolia Humb. & Bonpl., PL Aequin. 1:
1 13. pi. 33. 1808. TYPE: Ecuador, Loja, "crescit
in calidis Andium Peruvianorum inter vicum
Lucarque [Lucarqui] et amnem Macara [Macara].
Floret Novembri," Bonpland s.n. (holotype, p,
not seen; isotypes, F ex p, FM 974434; o, not seen,
photograph ex o, FM neg. 52108).
Barnadesia laurifolia (Humb. & Bonpl.) Kuntze, Rev.
Gen. PL 3: 135. 1898.
B. laurifolia (Humb. & Bonpl.) Hieron., Bot. Jahrb.
Syst. 28: 651. 1900.
Shrubs 3—4 m high, much-branched; stems 4-
8 mm in diameter, glabrous, axillary spines 10-
20 mm long, the lower branches nodulose. Leaves
petiolate, the petioles 5-10 mm long, glabrous;
blades oblong-lanceolate to ovate, 55-90 mm long,
20-34 mm wide, occasionally falcate, apically acute
or acuminate, both surfaces glabrous, 3-nerved
from near the base. Capitulescences corymbose
cymes, terminal or axillary. Capitula 10-15 mm
high, 2-3.5 mm wide; involucre 9-12 mm high,
2-3 mm wide; outer phyllaries ovate, 1.5-4 mm
long, 1.5-2 mm wide, densely pilose to villous,
ciliolate the inner lanceolate, 11-12 mm long, 1 .2-
1.5 mm wide, acuminate, the faces glabrous,
densely ciliate; the single floret with the corollas
white or violaceous, 8-10 mm long, 1-1.2 mm
wide, the lobes 1.8-2.5 mm long, pubescent; an-
thers 2.8-3.2 mm long. Achenes 6-7 mm long,
1.2-1.4 mm wide, pubescent; pappus ca. 10 mm
long, spreading, whitish to purplish.
DISTRIBUTION— Known from the intermontane
region of southern Ecuador to northwestern Peru
(1000- 1200m).
Fulcaldea laurifolia possesses a habit quite sim-
ilar to some Barnadesia species; however, it differs
in having a solitary floret per head, strongly plu-
mose pappus bristles, a swollen style apex, and
unique pollen grains. Other than Peruvian collec-
tions having slightly smaller and more falcate
leaves, there appear to be few differences between
the Ecuadorian and Peruvian representatives.
The isotype collection housed at F (neg. 974434)
is probably from the type gathering of Bonpland
originally held by Paris. The specimen contains a
hand-written label with the protolog, the stamp
"Ex Herbario Musei Parisiensis," and a small label
indicating the origin of the specimen as "Herb.
Schultz Bip." Since this species is still extremely
rare in herbaria, it is probable that the specimen,
sent to F on exchange from Paris, was a portion
of the collection made by Bonpland.
PIURA. Ayavaca: Puente Tandopa, Hutchison &
Wright 6685 (F, MO, us); Olleros (Pacae-La Pampa), Sa-
gastegui & Cabanillas 8675 (F, MO); Ayavaca, Soukup
4324 (F, us); Olleros, Weberbauer 6357 (F, GH, us).
Huancabamba: ca. 1 1 km S of Canchaque, Dillon & San-
chez 6213 (F, CPUN); below Canchaque, Ferreyra et al.
10932 (MO. USM); La Afiladera, carretera a Canchaque,
Lopez & Sagstegui 8774 (F, MO).
IX. GERBERA
Gerbera L., Opera Varia 247. 1758 (nom. cons.),
non Boehmer, 1760, nee. J. F. Gmelin, 1791.
TYPE: Gerbera linnaeiCass. = Arnica gerbera
L. (typ. cons.)
Acaulescent, scapose, perennial herbs, often with
a woolly crown. Leaves basal, usually obviously
petiolate; blades oval, lanceolate, or lyrate in out-
line, the margins entire, serrulate, dentate, or sin-
uate-pinnatifid, the abaxial surfaces usually to-
mentose, woolly, or sometimes glabrescent, the
adaxial surfaces usually glabrous. Capitulescences
of solitary, monocephalous scapes, the peduncles
bracteate or ebracteate, the heads usually erect.
Capitula heterogamous, radiate; involucres tur-
binate to broadly campanulate; receptacles flat,
epaleate; phyllaries 2-6-seriate, the outer shorter
than the inner; ray florets 1-2-seriate, usually pis-
tillate, staminodia often present, the corollas white,
yellow, pink, or red, bilabiate, the outer lip pro-
duced into a generally elongate ligula, the inner lip
short, usually bifid; disc florets hermaphroditic and
fertile, or the inner functionally staminate, the co-
rollas bilabiate, both lips short; anther bases sag-
ittate, caudate, the tails entire or ciliate; style
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
37
FIG. 8. Fulcaldea laurifolia. 1, habit; 2, capitulum; 3, floret with style; achene with pappus. (Drawn from A.
Sagdstegui & J. Cabanillas 8675, F.)
38
FIELDIANA: BOTANY
branches of the disc florets broadly lanceolate, the
tips rounded or subacute with short pollen-sweep-
ing trichomes outside. Achenes cylindrical, ribbed,
villous to pilose, rarely glabrous or nearly so, at-
tenuated at the apex; pappus of scabrid bristles,
1 -seriate.
Gerbera is an Old World genus of approximately
35 species, distributed in the Sino-Himalayan re-
gion, Madagascar, and tropical and southern Af-
rica; only one South African species is cultivated
in Peru. The limits between Gerbera and Tricho-
cline are established with some difficulty (Hansen,
1 990); the former genus is here denned with achenes
lacking multicellular, glandular trichomes. Zardini
(1974) transfered one native South American spe-
cies to Gerbera; however, that species is here treat-
ed under Trichocline (Hansen, 1991).
References
HANSEN, H. V. 1990. Phylogenetic studies in the
Gerbera-complex (Compositae, tribe Mutisieae,
subtribe Mutisiinae). Nord. J. Hot., 9: 469-485.
. 1991. Phylogenetic studies in Compos-
itae tribe Mutisieae. Opera Botanica, 109: 5-50.
JEFFREY, C. 1967. Notes on Compositae: II. The
Mutisieae in east tropical Africa. Kew Bull., 21:
177-223.
ZARDINI, E. M. 1974. Sobre la presencia del ge-
nero Gerbera en America. Bol. Soc. Argent. Bot.,
16: 105.
1 . Gerbera jamesonii Bolus ex Hook. f.
Scapose, perennial herbs to 5 dm. Leaves peti-
olate; petioles 15-18 cm long; blades oblong-
spathulate in outline, 15-18 cm long, 4-7 cm wide,
the abaxial surfaces villous, the adaxial surfaces
strigillous, the margins deeply lobed to lyrate-pin-
natifid. Capitulescences of solitary heads, the pe-
duncles 30-50 cm long, 3-5 mm in diameter,
ebracteate, striate, villous. Capitula with hemi-
spheric involucre, ca. 20 mm long, ca. 20 mm
wide; phyllaries 3 -seriate, the outer linear 7-8 mm
long, the inner lanceolate 15-16 mm long, 2-2.5
mm wide, all arachnoid- villous; ray florets 25-33,
the corollas scarlet to orange-red or salmon, the
outer lip 30-35 mm long, 3.5-4 mm wide, apically
3-toothed; disc florets 75-100, the corollas yellow,
8-9 mm long, the outer lip 3-3.5 mm long, ca. 1.5
mm wide, 3-toothed, the inner lip ca. 3 mm long,
ca. 1 mm wide, bipartite. Achenes isomorphic,
cylindrical, 4-5 mm long, densely villous, the tri-
chomes slender, acute; pappus ca. 7-9 mm long,
stramineous.
Gerberia jamesonii is a native of South Africa
(Transvaal) and only found in cultivation in Peru.
It is easily recognized by its scapose habit with
long peduncles and large heads with showy rays.
LIMA. Lima: Jesus Maria, cultivated, Museo de His-
toria Natural, Plowman 10989 (F).
X. GOCHNATIA
Gochnatia Kunth, Nov. Gen. Sp. 4: (fol.) 15,1818,
(qu.) 19. 1820. TYPE: G. vernonioides Kunih
Trees or shrubs, frequently tomentose (extra-
Peruvian species occasionally dioecious or gyno-
dioecious); stems lacking spines. Leaves alternate,
sessile or petiolate; blades circular or oval to lin-
ear-lanceolate, coriaceous to subcoriaceous, the
margins entire or dentate. Capitulescences cy-
mose-corymbose or racemose, or of solitary heads,
terminal and axillary. Capitula homogamous, dis-
coid, sessile or pedicellate; involucres campanu-
late to turbinate; receptacles plane or alveolate or,
more rarely, pubescent with glands or hairs; phyl-
laries imbricate, multiseriate, ovate to lanceolate;
florets isomorphic, hermaphroditic, the corollas
actinomorphic, yellow, tubular, deeply 5 -lobed,
the lobes linear, acute, recurved, equal to or short-
er than the tube; anthers linear, the terminal ap-
pendages lanceolate, the bases caudate, the tails
entire or laciniate; styles bilobed, the branches api-
cally rounded, glabrous. Achenes cylindric to tur-
binate, sericeous or glandular; pappus of scabrid
bristles, 2-many-seriate, yellowish.
Gochnatia consists of some 66 species with a
disjunct distribution between the paleotropics of
Southeastern Asia, China, India, Thailand, Laos,
Vietnam (2 spp.), and the Neotropics (64 spp).
Approximately 6 species are recorded from Mex-
ico and adjacent Texas, 26 species are endemic to
the Antilles, Cuba, Bahamas, Haiti, Dominican
Republic, and Puerto Rico, and the remaining 34
species are confined to South America. The South
American species are found in Peru, Chile, Ar-
gentina, Bolivia, Brazil, and Paraguay and occupy
a wide variety of habitats from intermontane val-
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
39
leys of the western Andean Cordillera to campo
and sierra habitats in Argentina, Paraguay, and
Brazil. In Peru, all Gochnatia are small shrubs
confined to mid-elevation (1500-3100 m), sea-
sonally dry sites within the Andean Cordillera.
Reference
CABRERA, A. L. 1971. Revision del genero Goch-
natia (Compositae). Revista Mus. La Plata, Secc.
Bot., 12: 1-160.
Key to Peruvian Species of Gochnatia
1. Leaves oblong, elliptic, or obovate; involucres cylindric; total florets 9-12 G. arequipensis
1 . Leaves ovate; involucres campanulate; total florets 1 5-30 2
2. Leaf margins dentate to serrate G. vargasii
2. Leaf margins entire 3
3. Leaves to 80 mm long; involucres 10-1 1 mm high; phyllaries acute, mucronate
G. vernonioides
3. Leaves to 25 mm long; involucres 14-17 mm high; phyllaries attenuate, aristate-mucronate
G. patazina
1. Gochnatia arequipensis Sandwith, Kew Bull.
1934: 100. 1934. TYPE: Peru, Arequipa,
"Volcan de Arequipa, in collibus aridis," 1 854,
W. Lechler 2751 (holotype, K).
G. iserniana Cuatr., Anal. Univ. Madrid 4: 22. 1935.
TYPE: Peru, Arequipa, Quenquena, 1 1 Aug 1863,
/. hern 448 (holotype, MA, not seen; isotype, LP).
Shrubs to 2 m high, much-branched, the
branches tomentose. Leaves subsessile; blades ob-
long, elliptic or obovate, 10-25 mm long, 6-8 mm
wide, apically obtuse or acute, the adaxial surfaces
slightly tomentose and glandular, the abaxial sur-
faces densely tomentose, the margins entire or re-
motely denticulate. Capitulescences of solitary
heads or in groups of 2-3. Capitula sessile; invo-
lucres cylindric, 10-12 mm high, 5-6 mm wide;
phyllaries imbricate, 6-7-seriate, the outer ovate-
lanceolate, 2-2.5 mm long, 1-1.5 mm wide, acute,
mucronate, lanuginose, the inner linear-lanceo-
late, 8-9 mm long, 1-1.2 mm wide, acute, mu-
cronate, glabrous; florets 9-12, the corollas with
tubes 8-9 mm long, the lobes linear, 4-5 mm long.
Achenes turbinate, densely sericeous, 4-5 mm long;
pappus ca. 1 2 mm long.
DISTRIBUTION— Endemic to the intermontane
region of southern Peru (2400-3100 m).
Gochnatia arequipensis most closely resembles
G. cardenasii with its more cylindric involucres
and reduced number of florets. It is easily distin-
guished from its nearest geographic neighbor, G.
vargasii, in possessing smaller, strictly entire leaves
and much smaller capitula.
AREQUIPA. Arequipa: slopes of Misti Volcano, Nu-
nez 92 (USM); slopes of Chachani, Rauh & Hirsch P579
(NY); Hutchison & Wright 7220 (MO, us, USM), Vargas
8281 (us), 79920 (us); Eyerdam & Beetle 22120 (GH);
Chiguata, Tovar 5130 (MO, USM). La Union: Cotahuasi,
Weberbauer 6866 (GH, MO, us, USM); without locality,
Rauh s. n. (USM), 7 969 (USM). TACNA. Tacna: near Tara-
ta, Beck 2024 (us).
2. Gochnatia patazina Cabr., Revista Mus. La
Plata, Secc. Bot. 12: 28. 1971. TYPE: Peru,
La Libertad, Pataz, entre Huaylillas y Taya-
bamba, A. Lopez M. & A. SagdsteguiA. 3409
(holotype, LP; isotype, HUT).
Shrubs to 1.5 m high, branches more or less
tomentose. Leaves subsessile; blades ovate, 10-25
mm long, 8-20 mm wide, obtuse, the adaxial sur-
faces slightly tomentose, the abaxial surfaces to-
mentose, the margins entire. Capitulescences of
solitary heads at branch apices, terminal or axil-
lary. Capitula sessile; involucres campanulate, 10-
14 mm high, 8-10 mm wide; phyllaries imbricate,
5-seriate, all aristate-mucronate, the outer ovate,
4-6 mm long, 1.5-2 mm wide, attenuate, slightly
lanuginose, the inner linear-lanceolate, 8-9 mm
long, 1-1.2 mm wide, lanuginose, attenuate; flo-
rets 30-40, the corollas with tubes 5-8 mm long,
the lobes linear, 3—4 mm long; anthers 5-6 mm
long. Achenes turbinate, 4-5 mm long, dense se-
riceous; pappus ca. 9 mm long.
DISTRIBUTION— Native to the intermontane re-
gion within the Departments of Ancash and La
Libertad (2300-3000 m).
Gochnatia patazina most closely resembles G.
40
FIELDIANA: BOTANY
vernonioides but differs in possessing smaller leaves
(< 25 mm long) and larger capitula with 40 florets.
ANCASH. Huaras: near Huaraz, Macbride & Feath-
erstone 2520 (us); Velarde Nunez 3178 (LP), 3179 (LP).
Yungay: between Rio Santa and Mancos, Smith & Bias
4901 (USM); Keusho, La Torre 425 (USM).
3. Gochnatia vargasii Cabr., Revista Mus. La Pla-
ta, Secc. Bot. 12: 19. 1971. TYPE: Peru, Apu-
rimac, Abancay, "bajando a C'onocc," 2150
m, C. Vargas 16317 (holotype, LP; isotype,
USM).
Shrubs to 2 m high, much-branched, the
branches tomentose. Leaves petiolate; petioles 2-
3 mm long, tomentose; blades ovate, 20-25 mm
long, 10-15 mm wide, obtuse, the adaxial surfaces
glabrous, the abaxial surfaces tomentose, the mar-
gins dentate. Capitulescences of solitary heads, ter-
minal. Capitula sessile; involucres campanulate,
10-12 mm high, 8-10 mm wide; phyllaries
5 -seriate, coriaceous, imbricate, the outer ovate,
mucronate, tomentose, glabrescent, the inner lan-
ceolate, 8-10 mm long, 1.5-2 mm wide, acute,
mucronate, tomentose; florets 30-35, the corollas
with tubes 7.5-8 mm long, the lobes 3.5-4 mm
long. Achenes turbinate, ca. 5 mm long; pappus
ca. 8 mm long.
DISTRIBUTION— Known only from the type lo-
cality.
Gochnatia vargasii appears to share relation-
ships with G. boliviana S. F. Blake and G. ver-
nonioides and G. patazina but is distinguished by
its dentate leaves and the ovate outer phyllaries.
APURIMAC. without exact locality, Rauh 1556 (USM).
4. Gochnatia vernonioides Kunth, Nov. Gen. Sp.
4: 20, t. 309. 1820. TYPE: Peru, "crescit in
regione calida Provinciae Bracamorensis in
ripa fluvii Amazonum, alt. 195 hex," Hum-
boldt & Bonpland s.n. (holotype, p, not seen).
Figure 9.
Shrubs 1-2 m high much-branched, the branch-
es dense pubescent, white-tomentose. Leaves sub-
sessile; blades elliptic-lanceolate to ovate, apically
acute, 25-60 mm long, 12-25 mm wide, the ad-
axial surfaces slightly glandulose, the abaxial sur-
faces tomentose, the margins entire. Capitules-
cences of solitary or weakly cymose heads, 2-3-
headed, terminal and axillary. Capitula sessile; in-
volucres campanulate, 8-12 mm high, 6-8 mm
wide; phyllaries 4-5 -seriate, imbricate, the outer
ovate, 4-6 mm long, 2-2.5 mm wide, mucronate,
lanuginose, the inner linear-lanceolate, 8-9 mm
long, 1.2-1.6 mm wide, mucronate, densely pu-
bescent; florets 26-29, the corollas with tubes 6-
7 mm long, the lobes 4-5 mm long, glabrous; an-
thers 5-6 mm long. Achenes turbinate, 3.8-5 mm
long, 0.8-1 mm wide, densely sericeous; pappus
8-10 mm long.
DISTRIBUTION— Known from the intermontane
region of northern Peru (1600-2600 m).
Gochnatia vernonioides is abundant in season-
ally dry habitats on both sides of the valley of the
Rio Maranon at Balsas and is represented by more
collections than any other Peruvian species. It un-
doubtedly shares relationships with G. patazina,
its nearest geographic neighbor; however, G. ver-
nonioides is readily distinguished by its smaller
involucres and generally larger leaves.
AMAZONAS. Chachapoyas: Tingo, Ferreyra 7097
(MO, us, USM); near Chachapoyas, Ferreyra 14428 (MO,
USM); E of Chachapoyas, King & Bishop 9169 (MO, us)
9756 (us), 9227 (MO, us); Chachapoyas-Leimebamba,
Lopez et al. 4364 (F, MO, NY); Balsas-Leimebamba, Sa-
gastegui 7487 (F, us), Hutchison & Wright 5856 (F, OH,
MO, NY, uc, us, USM); Chachapoyas, 1834-1840, Ma-
thews s.n. (GH). CAJAMARCA. Celendin: Limon, San-
chez 2804 (CPUN, F).
XI. JUNGIA
Jungia L. fil., Suppl. pi. 58: 1781. TYPE: Jungia
ferruginea L. fil.
Subshrubs, annual or perennial herbs, or lianes;
stems lacking spines. Leaves alternate, cauline or
rarely strictly basal; petiolate, petioles with or
without basal stipuliform bracts; blades oblong-
lanceolate to cordiform or round, pinnate- or pal-
mate-veined, the margins lobed, serrate, dentate,
or incised. Capitulescences paniculate or corym-
biform, terminal or axillary. Capitula homoga-
mous; involucres campanulate, cylindric, or tur-
binate; receptacles plane, paleate, the pales scari-
ous or similar to phyllaries; phyllaries 2-seriate,
the inner series embracing florets, equal to un-
equal, imbricate, lanceolate, oblong, or spathulate,
apically acute to acuminate, mucronate, glabrous
to densely pubescent and/or glandular; florets iso-
morphic, hermaphroditic, the corollas white, yel-
low, pink, reddish, or violaceous, bilabiate, the
outer lip lanceolate or elliptic, 4-nerved, triden-
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
41
FIG. 9. Gochnatia vernonioides. 1, habit; 2 capitulum; 3, floret with achene. (Drawn from P.C. Hutchison & J.K.
Wright 5856, F.)
plumose bristles, 1 -seriate, white to gray, reddish,
tate, the inner lip bind to subentire, laciniate, coiled; tawny, or stramineous,
anthers caudate; styles branches claviform, trun-
cate, penicillate. Achenes cylindric, rarely 4-costate, Jungia contains approximately 30 species rang-
glabrous to pubescent and/or glandular; pappus of ing from Mexico through Central America and
42
FIELDIANA: BOTANY
along the Andean Cordillera from Colombia to
Argentina, Brazil, and Chile. No fewer than 10
species are recorded from Peru and these generally
occupy moist, montane environments at eleva-
tions above 2000 m. Of the Peruvian species, only
J. axillaris occupies drier sites below 2000 m on
the western escarpment.
References
CERRATE-VALENZUELA, E. 1951. Revision de las
especies peruanas del genero Jungia. Publ. Mus.
Hist. Nat. "Javier Prado," Ser. B, Bot., 4: 1-24.
HARLING, G. 1992. Two new species of Jungia
(Compositae) from Peru. Novon, 2: 16-18.
Key to Peruvian Species of Jungia
1 . Annual herbs or short-lived perennial herbs 2
2. Leaves essentially basal, the blades oblanceolate to oblong-elliptic, penni-nerved; petioles without
stipuliform bracts; florets 70-80 J. stuebelii
2. Leaves cauline, the blades orbicular to cordiform, palmate-nerved; petioles generally with basal,
stipuliform bracts; florets 14-1 8 3
3. Capitulescences densely corymbose to paniculate; phyllaries densely villous; pappus strami-
neous /. vitocensis
3. Capitulescences open paniculate; phyllaries glandular-puberulent to pilose; pappus whitish to
blackish-brown 4
4. Stems succulent; pappus blackish-brown; achenes 4.5-6 mm long J. amplistipula
4. Stems not succulent; pappus whitish; achenes 3-5 mm long J. floribunda
1 . Erect, ascending, scandent, or vining, perennial herbs, or shrubs 5
5. Cauline leaves generally less than 4 cm long and 4 cm wide J. axillaris
5. Cauline leaves generally equal to or greater than 4 cm long and 4 cm wide 6
6. Abaxial leaf surfaces densely black puncticulate J. spectabilis
6. Abaxial leaf surfaces lacking black punctations 7
7. Leaf blades triangular-cordate to ovate-cordate, concolorous, the abaxial surfaces densely
sericeous-tomentose, the adaxial surfaces densely strigose J. weberbaueri
1 . Leaf blades orbicular, rotund-cordate, or cordiform, discolorous, the abaxial surfaces to-
mentose to strigillose, the adaxial surfaces bullate, strigillose, or glabrous 8
8. Petioles possessing stipuliform bases; leaf blades with adaxial surfaces bullate: phyllaries
6-7.5 mm long, densely glandular, apically acuminate J. rugosa
8. Petioles lacking stipuliform bases; leaf blades with adaxial surfaces smooth; phyllaries
3.5-6 mm long, ferruginous-pilose to flavo-tomentose, apically acute to obtuse or round-
ed 9
9. Phyllaries ferruginous-pilose, apically obtuse or rounded; corollas white to purple or
lavender /. discolor
9. Phyllaries flavo-tomentose, apically acute; corollas yellow J. paniculata
1. Jungia amplistipula Cerrate, Publ. Mus. Hist.
Nat. "Javier Prado," Ser. B, Bot. 4: 10. 1951.
TYPE: Peru, Ayacucho, Lucancas, near Pu-
quio, R. Ferreyra 7202 (holotype, USM; iso-
types, MO).
Annual herbs, 12-14 dm high, erect, stems
semisucculent, pubescent. Leaves cauline; petioles
9-120 mm long, stipuliform bracts 25-40 mm long,
25-45 mm wide, round, the margins lobed; blades
cordiform, 50-120 mm long, 50-155 mm wide,
membranaceous, the adaxial surfaces pubescent-
glabrescent, the abaxial surfaces arachnoid, palmi-
nerved 5-7-veined, the margins 9-12-lobed, the
lobes unequal, dentate, mucronate. Capitules-
cences panicles to 30 cm long, terminal or axillary.
Capitula pedunculate; involucres 6.5-7 mm high,
5-6 mm wide; phyllaries 2-seriate, the inner 6.5-
7 mm long, 1.8-2 mm wide, acute; florets 17-18,
the corollas white, 8-8.5 mm long, the tube 4—4.5
mm long, the outer lip 4.5-5 mm long, 4-nerved,
tridentate, the inner lip 2.5-3 mm long; anthers
3.8-4 mm long, the tails to 1 mm long. Achenes
(4.5-)5.5-6 mm long, glandular-puberulent; pap-
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
43
pus of 30-32 bristles, ca. 5 mm long, blackish-
brown.
DISTRIBUTION— Known from the Andean Cor-
dillera of northern to southern Peru (2700-3400
m).
Jungia amplistipula most closely resembles J.
floribunda but differs in its succulent stem, leaves
with 9-12 lobes and paniculate capitulescences.
CUSCO. Calca: Pisac, Maren 308 (F); Pissac, Vargas
5064 (USM); Ccachca, Vargas 13217 (USM). Convencion:
Lucumayo, Vargas 4237 (us). Cusco: Cusco, Soukup 79
(F), 252 (F), 929 (USM); Saxaihuman, Herrera 2365 (F);
Sacsahuman, Pennell 13580 (F), Vargas 4165 (USM). Pa-
ruro: Ayusbamba a Mayhus, Vargas 888 (F). Paucartam-
bo: Llulluchayoc, Vargas 4356 (USM); Pilco, Woytkowski
280 (USM). I rubamba: Ollataytambo, Cook & Gilbert
498 (us); Ellenberg 1159 (us); Huayoccari to Yanacocha,
Nunez et al. 7006 (F); Machupicchu y el Camino Inca,
Nunez & Luna 8855 (F); Huinaihuaina, Vargas 4138
(USM); Santa Rita, Vargas 2662 (USM). HUANCAVE-
LICA. Castrovirreyna: near Cordova, Metcalf 30275 (GH,
uc, us). LIMA. Huarochiri: above San Bartolome, Mon-
te Zarate, Ferreyra 9719 (MO, USM). PIURA. Huanca-
bamba: Abra de Porculla, Ferreyra 13745 (USM).
2. Jungia axillaris (DC.) Spreng., Syst. Veg. 4.
Park 1: 301. 1827.
Dumerilla axillaris DC., Ann. Mus. Hist. Nat. Par.
19: 72. t. 15, pi. 6. 1812. TYPE: "in Peruvia,
Chili et Panamaide," Lagasca s.n. (holotype, P,
not seen).
Jungia seleriana Muschl., Hot. Jahrb. Syst. 50, Beibl.
Ill: 107. 1913. SYNTYPES: Peru, "Agua ver-
rugas ad viam ferream Lima-Oroya," Seler 227
(B, presumably destroyed, photograph ex B, FM
neg. 16127); Peru, "inter Narquimam et Matu-
canam, ad viam ferream Lima-Oroya," A. We-
berbauer 69 (B, presumably destroyed).
Perennial herbs, 12-15 dm high, branched, the
branches 5-6 cm long, pubescent. Leaves petio-
late; petioles 7-23 mm long, 2 stipuliform bracts
at the base, amplexicaul, 2-8 mm long, the mar-
gins dentate-mucronate; blades cordiform, 14-37
mm long, 13-63 mm wide, palmi-nerved, 5 veins
prominent, the adaxial surfaces glabrous, the ab-
axial surfaces glandulose, the margins 7-lobed,
acute, unequal, dentate-mucronate. Capitules-
cences paniculate, terminal or axillary. Capitula
pedunculate; involucre 9-9.8 mm high, 5-6 mm
wide, campanulate; phyllaries 2-seriate, the inner
8-9 mm long, 1.8-2 mm wide, acuminate, ciliate,
red-purple, lanceolate, pubescent outside, hairs
white at apex, brown at base; florets 20-25, the
corollas violaceous, 10-12 mm long, the tube 6-
6.5 mm long, the outer lip 4-4.5 mm long,
4-nerved, tridentate, pubescent, the inner lip 3.5-
4.2 mm long; anthers 6-6.5 mm. Achenes 2-7 mm
long, glabrescent; pappus of 45-5 1 bristles, ca. 7
mm long, white.
DISTRIBUTION— Known from the western es-
carpment of the Andean Cordillera in northern to
southern Peru (1000-3400 m).
Jungia axillaris is distinguished by its numer-
ous, cordiform leaves, large paniculate capitules-
cences, and violet corollas. This species occupies
drier sites usually at elevations below 2000 m but
occasionally can be found higher.
ANCASH. Bolognesi: abajo de Raquia, Pativilca-
Huaras, Sagdstegui et al. 12292 (F). Huaras: Tambo de
Pariocoto [Pariacota], Macbride & Featherstone 2551
(GH). Huarmey: Lomas de Lupin, Cerrate 803 (MO, USM),
812 (MO, USM); Ferreyra 764A (MO, USM), 3922 (MO, USM),
8061 (MO, USM), 8372 (MO, USM), 73790 (MO, USM). AYA-
CUCHO. Lucanas: 43 km E of Nazca on road to Puquio,
Gentry et al. 23253 (F, MO). CUSCO. Cusco: near Cusco,
Gay s.n. (GH). LA LIBERTAD. Trujillo: Pate, Hda. Cal-
ambay, Ridoutt & Vargas 2248 (F). LIMA. Cajatambo:
Churin, Ferreyra 6160 (MO, USM). Canta: Obrajillo, Ma-
thews s.n. (GH, us). Huarochiri: Rio Rimac, Angulo 2298
(F); between Lima and San Mateo, Duncan et al. 2711
(F, MO); between Lima and Oroya, Ferreyra 3922 (USM),
5327 (USM); Santa Eulalia Valley, Hutchison & Saravia
7096 (F); Chosica, Rose 18540 (us); between Lima and
Oroya, Stevens 22101 (F). NO EXACT LOCALITY.
Capt. Wilkes Exped. s.n. (us), Haenke 1831 (F).
3. Jungia discolor Muschl., Bot. Jahrb. Syst. 50,
Beibl. Ill: 106. 1913. TYPE: Peru, Caja-
marca, Hualgayoc, Chugur, 2700-2900 m, 21
May 1904, A. Weberbauer 4083 (holotype, B,
presumably destroyed; photograph ex B, FM
neg. 664115).
Scandent shrubs or climbing vines; stems 5-8
m long, striate, puberulent, glabrescent. Leaves
petiolate; petioles 20-50(-70) mm long, strigillose-
puberulent, lacking stipuliform bases; blades ro-
tund-cordate, 35-90 mm long, 55-100 mm wide,
generally strongly discolorous, the adaxial surfaces
sparsely strigillose, prominent-reticulate, ferrugi-
nous, the abaxial surfaces sparsely to densely stri-
gillose, tawny, the margins shallowly 6-8-lobed,
crenate-undulate, mucronulate. Capitulescences
cymose-paniculate, terminal and axillary, the pe-
duncles 50-100 mm long, the pedicles 10-20 mm
long, ferruginous-tomentose. Capitula sessile to
pedicellate; involucres 4-6 mm long, 4-6 mm wide,
campanulate; phyllaries 2-seriate, all apically ob-
tuse to rounded, mucronulate, the outer oblong to
oblong-linear, 4-5 mm long, 1-1.5 mm wide,
44
FIELDIANA: BOTANY
densely ferruginous-pilose, the inner 5.5-6 mm
long, oblong to obovate, the keel ferruginous-pi-
lose; florets 12-15(-20), the corollas white to pur-
ple or lavender, 6-7 mm long, the tube ca. 4.5 mm
long, the outer lip 3-4 mm long, ca. 1.5 mm wide,
3-fid, the inner lip ca. 2.5 mm long, deeply 2-fid;
anthers 4.5-5 mm long. Achenes cylindric, ca. 2
mm long, glabrous; pappus of 30— 40 bristles, white,
5-6 mm long.
DISTRIBUTION— Known from the Andean Cor-
dillera in north-central Peru (2700-3400 m).
Jungia discolor is distinctive among the Peru-
vian species in possessing exstipulate, discolorous,
shallowly lobed leaves and oblong to obovate
phyllaries with obtuse to subtruncate apices. Jun-
gia discolor most closely resembles J. coarctata
Hieron., an Ecuadorian species, which shares the
characteristic exstipulate leaves with abundant,
mucronate teeth on the shallow lobes, but that
species has longer, acute phyllaries and nearly con-
colorous leaves.
AMAZONAS. Chachapoyas: Puma-urcu, SSE of
Chachapoyas, Wurdack 690 (F). CAJAMARCA. Hual-
gayoc: Hacienda Taulis, Hutchison & Bismarck 6510 (F,
MO). HUANUCO. Huanuco: Villcabamba, Hacienda on
Rio Chinchao, Macbride 5180 (F). Pachitea: Mima, trail
to Tambo de Vaca, Macbride 4331 (F). SAN MARTIN.
Mariscal Caceres: Rio Abiseo National Park, Young 37 2 3
(F).
4. Jungia jloribunda Less., Linnaea 5: 38. 1830.
TYPE: Eastern Brazil, F. Sellow s.n. (holo-
type, B, presumably destroyed; photograph ex
B, FM neg. 16119).
Annual herbs, 60-70 cm high, pubescent. Leaves
petiolate; petioles 2.2-6.2 mm long, pubescent,
upper leaves with paired stipuliform bracts, un-
equal, 7-20 mm long, 10-25 mm wide; blades
cordiform, 30—40 mm long, 40-70 mm wide, the
adaxial surfaces glabrous, the abaxial surfaces
arachnoid, palmi-nerved, 5-7-veined, the margins
7-8-lobed, crenate, rounded, unequal. Capitules-
cences cymose, 10-15-headed. Capitula pedun-
culate; involucres 6-9 mm high, 3.5-5 mm wide,
turbinate; phyllaries 2-seriate, the inner 7-8 mm
long, 1.8-2.5 mm wide, acute; florets 14-15, the
corollas yellowish-white, 7.5-8 mm long, the tube
4-4.6 mm long, the outer lip ligulate, 4.2-4.6 mm
long, 1-2 mm wide, 4-nerved, tridentate, the inner
lip 2-4 mm long, 0.8-1 .2 mm wide, bifid. Achenes
cylindric, 3-5 mm long, 0.8-1 mm wide, pubes-
cent; pappus of 23-24 bristles, ca. 5 mm long, gray
to white.
DISTRIBUTION— Known from the Andean Cor-
dillera from northern to central Peru (2000-3400
m).
Jungia jloribunda is a herbaceous annual char-
acterized by upper leaves with large stipules and
the capitula in dichotomous cymes. Potential is-
otypes may be housed at P and a fragment from
Sellow 3643 (F 971243 ex P) matches the species.
Harling (1992) recently described a new species,
Jungia gracilis, based on material from the Cha-
chapoyas region (King & Bishop 9278) that he
compares to J. spectabilis. No decision is here made
as to the validity of this new species, and the type
material is cited under J. floribunda.
AMAZONAS. Chachapoyas: Utcubamba Valley, King
& Bishop 9165 (MO, us), 9213 (MO), 9275 (MO); Ubilon,
Ferreyra 15622 (F, us, USM). CAJAMARCA. Cajamarca:
Chetilla, ruta a Llullapuquio, Sanchez & Cabanillas 4104
(F). Chota: between Chota and Cajamarca, Cerrate 1625
(USM). Contumaza: Tambo de Luria, Lopez & Sagdstegui
9032 (MO); above Cascas, Lopez & Sagdstegui 9003 (MO);
Guzmango, Sagdstegui 2901 (F); Quebrada Honda, Sa-
gdstegui 3766 (F); Trinidad, Sagdstegui 8950 (MO); Cruz
del Hueco, Sagdstegui 9945 (MO); Cerro Chungarran,
Guzmango, Sagdstegui 9210a (F, MO); Pampa de la Sal,
Sagdstegui 10751 (MO), 10032 (MO); Cruz Grande, Sa-
gdstegui 8996 (MO); Herilla, Sagdstegui & Samame 2945
(F). HUANCAVELICA. Castrovirreyna: near Cordova,
Metcalf 30275 (MO). HUANUCO. Huanuco: near Hua-
nuco, Macbride & Featherstone 2134 (F, us); Mito, Mac-
bride & Featherstone 1586 (F, us); Huacachi, estacion
near Muna, Macbride 4067 (F). LA LIBERTAD. Otuzco:
Agallpampa, Ldpez 1866 (F). Pataz: Tayabamba, Lopez
& Sagdstegui 8163 (MO).
5. Jungia paniculata (DC.) A. Gray, Proc. Am.
Acad. Sci. 5: 145. 1861.
Dumerilia paniculata DC., Ann. Mus. Hist. Nat. Paris
1 9: 72, 1. 1 6, pi. 7. 1 8 1 2. TYPE: Peru, J. deJussieu
s.n. (holotype, P-JU, not seen; photograph ex p,
FM neg. 1037999).
Shrubs 1 5-20 dm high, branched, the branches
glandular pubescent. Leaves petiolate; petioles 5-
25 mm long, lacking stipuliform bracts; blades cor-
diform, 40-90 mm long, 40-90 mm wide, the ad-
axial surfaces puberulent, the abaxial surfaces white
tomentose, palmi-nerved, 5 veins prominent, the
margins 5-7-lobed, acute, unequal, mucronate-
dentatc. Capitulescences paniculate; bracts fili-
form. Capitula peduncles 15-80 mm long; invo-
lucre 4.2-6 mm high, 2.5-4.5 mm wide, turbinate;
phyllaries 2-seriate, the inner 3.5-5.5 mm long,
1.2-1.8 mm wide, flavo-tomentose, acute; florets
17-27, the corollas yellow, 6-8 mm long, the tube
3.5-5.2 mm long, the outer lip liguliform 2-3.2
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
45
mm long, 4-nerved, tridentate, the inner lip 2-3.2
mm long; anthers 2.8-5 mm long. Achenes tur-
binate, 2-2.8 mm long, sparsely pubescent; pappus
of 25-31 bristles, ca. 5.8 mm long, white.
DISTRIBUTION— Known from the Andean Cor-
dillera of northern and central Peru (2500-3500
m).
Jungia paniculata has been traditionally ac-
cepted as the valid name for a species distributed
from central and north-central Peru. Harling (1 992)
argued that the "true" J. paniculata is known from
Colombia and Ecuador and is rare in northern
Peru (only recorded in Cajamarca) and has de-
scribed a new species J. schuerae, based on ma-
terial gathered near Matucana, Lima. The illus-
tration in DeCandolle's publication depicts a plant
with much less congested capitulescences and the
photograph from J. de Jussieu's herbarium in Paris
also shows this characteristic. Since the exact lo-
cality for J. de Jussieu's collection has not been
established and since type material of Harling' s
new species has not been studied yet, some of the
material treated here may potentially represent his
new taxon.
ANCASH. Bolognesi: Usgor, Ferreyra & Cerrate 7563
(USM), Cerrate 459 (USM); Chiquian, Mostacero et al.
1344 (F). Carhuas: Vicos, Hutchison & Wright 4299 (F,
MO, us). Huaras: Marcara, Proaho 149 (USM); Puente
Bedoya, Proano 39 (USM); Banos de Chancos, Sandeman
4593 (F); above Cochabamba, Smith & Buddensiek 10958
(F). Huari: Chavin, Rauh 40472 (USM). Huaylas: Santo
Toribio, Mostacero et al. 1975 (F). Yungay: Lagunas
Llanganuco, Gentry et al. 37370 (F, MO, USM); Huascaran
National Park, Smith 11295 (F); Llanganuco, Zardini
7576 (MO). CAJAMARCA. Cajamarca: El Gavilan, San-
chez 237 (CPUN, F, USM); Llacanora, Sanchez 2246 (F,
MO, us). Celendin: above Balsas, Hutchison & Wright
5232 (F, MO, uc, USM); Llanguat, Mostacero et al. 947
(F). HUANUCO. Dos de Mayo: below Chavanillo, Mac-
bride & Featherstone 1987 (F, GH, us). LA LIBERT AD.
Pataz: Huancaspata, Lopez & Sagdstegui 8236 (GH, MO).
Santiago de Chuco: Santiago de Chuco, Lopez 977 (USM),
7957 (F), Sagdstegui et al. 11712 (F). LIMA. Canta: road
to Canta, Cerrate 8680 (USM); ca. 1.5 km NE of Canta,
Dillon & Molau 3180 (CPUN, F, HUT, MO, NY, TEX, USM);
Chiuchin, Ferreyra 19167 (USM); San Juan, Ferreyra
18358 (USM); Ingenio, Meza 73 (USM); along Rio Chillon,
below Obrajillo, Pennell 14430 (F, GH, us, USM); Huaripa,
Sanchez 56 (USM); Canta, Soukup 2815 (F). Huarochiri:
San Mateo Cerrate 5328 (USM); Santa Eulalia, Hutchison
& Saravia 7097 (F, USM); Viso, Macbride & Featherstone
757 (F, GH, us). Lima: near Lima, Cerrate 2924 (USM);
Cerrate 2393 (USM); market in Lima, Cerrate 7709 (MO);
Infiernillo, Ferreyre 6246 (USM); Monte Zarate, Gutte &
Muller 9509 (USM); Matucana, Infantes 1987 (USM), Ri-
doutt s.n. (USM 15371), Vargas 4750 (USM). NO EXACT
LOCALITY. Ellenberg 1724 (USM), Ames s.n. (USM).
6. Jungia rugosa Less., Linnaea 5: 36. 1 830. TYPE:
Ecuador, Prov. Chimborazo, "locis frigidis in
radicibus montis nobilissimi Chimborazo, alt.
1860 hex. Regno Quitensi," Humboldt 3143
(holotype, B, presumably destroyed; photo-
graph ex B, FM neg. 664249; isotype fragment,
F 10 12092 ex P).
Jungia jelskii Hieron., Bot. Jahrb. Syst. 36: 512. 1905.
TYPE: Peru, Cajamarca, Chota, entre Chota y
Cutervo, Jelski 639 (holotype, B, presumably de-
stroyed; photograph ex B, FM neg. 664243).
Jungia malvaefolia Muschl., Bot. Jahrb. Syst. 50, Beibl.
1 1 1: 105. 1913. TYPE: Peru, Ancash, Huari, Co-
nin, A. Weberbauer 2912 (holotype, B, presum-
ably destroyed; photograph ex B, FM neg. 16122).
Shrubs 2-3 m high; stems glandular pubescent,
ferruginous. Leaves petiolate; petioles 10-65 mm
long, hirsute to glandular pubescent, the bases with
stipuliform bracts, occasionally reduced or absent;
blades cordiform, 13-70(-100) mm long, (9-) 12-
78 mm wide, the adaxial surfaces conspicuously
rugose-bullate, strigillose or villosulous to gla-
brescent, the abaxial surfaces prominently retic-
ulate-nerved, tomentose, the margins 5-lobed, un-
equal, obtuse, crenate. Capitulescences paniculate,
terminal or axillary. Capitula peduncles (17-)22-
65 mm long, pubescent, the bracteoles densely cil-
iolate, glandular; involucres 6-9(-10) mm high,
4-6(-7) mm wide, campanulate; phyllaries
2-seriate, the outer lanceolate, 6-7 mm long, ca.
1.5 mm wide, glandular, apically acute to atten-
uate, the inner oblong-lanceolate, 6.8-7.5(-11.5)
mm long, 1.8-2.6(-3) mm wide, apically acute to
attenuate; florets 20-35, the corollas yellowish-
white to white, 9-9.5(-14) mm long, the tube 5-
6.5 mm long, the outer lip 2.5-4.5(-6.5) mm long,
4-5-nerved, tridentate, the inner lip 2.5-4(-6) mm
long; anthers 4-6 mm long. Achenes cylindric, 2.5-
5 mm long, 0.4-1 mm wide, cylindric, subglabrous
to puberulent with 1 -seriate, multicellular, glan-
dular trichomes; pappus of 35-37 bristles, 8-9 mm
long, white.
DISTRIBUTION— Known from the Andean Cor-
dillera of Ecuador to southern Peru (2900-3800
m).
Jungia rugosa is readily distinguished by its
characteristic bullate upper leaf surfaces and glan-
dular phyllaries. The apex of the phyllaries is vari-
able from acute to attenuate, but the densely glan-
dular pubescence and ciliolate margins is essen-
tially invariable. The foliar characteristics of both
J. jelski and J. malvaefolia agree well with those
46
FIELDIANA: BOTANY
of J. rugosa, and their distributional ranges also
coincide.
AMAZONAS. Luya: 41 km SW of Leimebamba,
Gentry et al. 23148 (F). ANCASH. Carhuaz: Quebrada
Ulta, Smith 11408 (F). Huari: Quebrada Rurichinchay,
Smith et al. 12472 (F). Huaylas: Caraz-Laguna Paron,
Lopez et al. 8361 (F); Quebrada Paron, Smith 10572 (F);
Caras, Weberbauer 3224 (MOL). Yungay: Laguna Llan-
ganuco, Mostacero et al. 1398 (F); Quebrada Yanapac-
cha, Smith et al. 10450 (F). APURIMAC. Abancay: Say-
roite, Vargas 1032 (GH); near Socclaccasa, West 3925
(uc). Grau: Lambrana-Chuquibambilla, Hoogte &
Roersch 1451 (F). CAJAMARCA. Contumaza: Pampa
de la Sal, Sagdstegui et al. 10751 (F); Salcot, Sanchez &
Sanchez 3138 (F). CUSCO. Anta: Limatambo, Vargas
8198 (MO, USM). Calca: Amparaes, Nunez 6763 (MO);
near Lares, Vargas 3581 (MO, USM). Paucartambo: Pau-
cartambo Valley, H err era 2975 (us, USM); Paucartambo-
Pilcopata, Molau 764 (USM); Tres Cruces, Pennell 14149
(F, GH, NY); Pillahuata, Vargas 1926 (USM); Llulluchayoc,
Vargas 4309 (USM). Urubamba: Ollantaytambo, Cook &
Gilbert 802 (us), 493 (us), 7567 (us); Pisac, Ellenberg
4833 (us); Tanccac, Marin 1579 (us); Penas, road to
Panticalla, Vargas 4398 (USM). LA LIBERT AD. Bolivar:
Unamen, Lopez & Sagdstegui 3321 (F); Laguna de Srur-
huas, Young 3555 (F). Pataz: Yalen to los Alisos, Young
2929 (F). LAMBAYEQUE. Ferrenafe: near Incahuasi,
Ferreyra & Vreeland 20905 (USM). PIURA. Huancabam-
ba: Cienego largo, Sagdstegui et al. 8261 (F).
7. Jungia spectabilis D. Don, Trans. Linn. Soc.
London 16: 227. 1833. TYPE: Ecuador, Prov.
Guayas, Guayaquil, Tafalla s.n. (holotype, MA,
not seen).
Jungia floribunda Spruce ex Benth. & Hook., Gen. pi.
2: 502. 1873, non Less. 1830. TYPE: Ecuador,
"in Andibus Ecuadorensibus," R. Spruce 5966
(holotype, K, not seen; photograph ex c, FM neg.
22568).
Shrubs 8-15 dm high, branched, branches more
or less tomentose. Leaves petiolate; petioles 3-6.5
mm long, puberulent-tomentose; blades cordi-
form 150-200 mm long, 200-300 mm wide, the
adaxial surfaces sparsely strigillose to glabrescent,
the abaxial surfaces finely strigillose, the veins
prominent-reticulate, densely black puncticulate,
the margins 7-9-lobed, crenate. Capitulescences
paniculate. Capitula pedunculate, the pedicles to
5 cm long; involucres campanulate, 6-9 mm long,
3-7 mm wide; phyllaries 1 -seriate, 10-12, lanceo-
late, glandular-puberulent, basally callose, apically
attenuate; florets 2 1 , the corollas white, bilabiate,
the outer lip elliptic, tridentate, the inner lip bifid,
coiled, linear. Achenes cylindric-fusiform, 5-6 mm
long; pappus of ca. 24 bristles, 5-6 mm long, white.
DISTRIBUTION— Known from the intermontane
region of Ecuador and northern Peru (1400-3200
m).
Jungia spectabilis is readily distinguished by its
large leaves with numerous black punctations on
the abaxial surfaces. Marling (1991) attributed type
fragments (Tafalla s.n.) to F; however, no such
collection has been located at F.
CAJAMARCA. Contumaza: Cascas, Zubiate 1050
(USM); Contumaza, Pena s.n. (USM). Hualgayoc: Monte
Seco, Soukup 3824 (F, us). San Miguel: entre el Naranjo
y La Alhaja, Niepos, Llatas 1242 (F). San Pablo: Sangal,
Sanchez 2849 (F). LA LIBERT AD. Otusco: Agallpampa,
Lopez 868 (USM). PIURA. Huancabamba: El Tambo,
Acleto 250 (USM); Canchaque, Sagdstegui 8131 (MO), 8545
(F, MO).
8. Jungia stuebelii (Hieron.) Crisci, Bol. Soc. Ar-
gent. Bot. 13: 341. 1971.
Leuceria stuebelii Hieron., Bot. Jahrb. Syst. 2 1 : 372.
1896. TYPE: Peru, Cajamarca, Celendin, "crescit
supra Celendin inter Pascasmayo et Moyobamba,
3200 m," A. Stubel 35g (holotype, B, presumably
destroyed; photograph ex B, FM neg. 16059).
Jungia longifolia Cerrate, Publ. Mus. Hist. Nat. "Ja-
vier Prado," Ser. B, Bot. 4: 7. 195 1. TYPE: Peru,
Ancash, Chiquian, Bolognesi, Huancar, Cerrate
193 (holotype, USM).
Annual herbs, 4.5-1 2 dm high, erect, pubescent.
Leaves petiolate or sessile; petioles 30-85 mm long;
blades oblanceolate to oblong-elliptic 95-205 mm
long, 40-80 mm wide, penninerved, 7-8-veined,
pubescent. Capitulescences paniculate. Capitula
peduncles 3-8 mm long; involucres 10-13 mm
high, 8-12 mm wide, campanulate; phyllaries
2-seriate, 10-12 mm long, 2-2.5 mm wide, pu-
bescent, acute, reddish-brown; florets 70-80, the
corollas white, 10.5-12.5 mm long, the tube 5-5.5
mm long, the outer lip 6-7.2 mm long, 1-2.8 mm
wide, 4-nerved, tridentate, the inner lip bifid, 3.8-
4 mm long. Achenes cylindric, 3.2-3.5 mm long,
0.8-1 mm wide, pubescent; pappus of 28-32 bris-
tles, 5-6 mm long, tawny.
DiSTRiBUTiON-Known from the intermontane
region of north and central Peru (2600-3800 m).
Jungia stuebelii closely resembles J. floribunda
from which it differs in having basal, lanceolate,
penninerved leaves and capitula with 70-80 flo-
rets.
ANCASH. Bolognesi: Chiquian, Cerrate 461 (MO, USM);
Pasamarca, near Chiquian, Cerrate 1614 (USM); near
Chiquian Ferreyra 5744 (MO, USM, us). Carhuas: Cor-
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
47
dillera Blanca, valley of the Rio Marcara, road to Vicos,
Hutchison & Wright 4385 (F, uc, USM); near Carhuas,
Infantes 1131 (USM). Huaras: Olleros, King & Collins
9048 (us). Huari: above Chavin, Ferreyra 14555 (USM).
Huaylas: Tocanca-Punta, Mostacero et al. 1961 (F);
Huascaran National Park, Alpamayo-Cashapampa,
Smith & Valencia 10040 (F); Quebrada Paron, Smith
10609 (F); Auquispuquio, Smith et al. 11954 (F); 72075
(F). Yungay: Quebrada de Llanganuco, Edwin & Schunke
3838 (F); Llanganuco, Ferreyra 14355 (MO, USM), Lopez
et al. 8342 (F, MO), Smith & Cautivo 10313 (F), Smith
et al. 10338 (F), Perez 22 (USM). CAJAMARCA. Caja-
marca: El Gavilan, Ferreyra 3267 A (USM); Kumulca baja,
Sanchez 195 (CPUN, F); Quinuamayo, entre Kumulca y
Encanada, Sanchez & Ruiz 466 (F, USM); entre Cerro
Gavilan-Cajamarca, Sanchez 578 (F, USM), Sanchez &
Molau 2938 (F); Encanada, Sagdstegui 7385 (us); Jalca
Kumulca, Sagdstegui et al. 81 16a (F, MO); Banos del Inca,
Soukup 4636 (F, us). Celendin: Challuayaco, Sanchez &
Cabanillas 3477 (MO). Contumaza: Pampa de la Sal, Sa-
gdstegui et al. 10033 (F, MO); arriba de Lleden, Sagdstegui
et al. 10809 (F, MO); Cascabamba, Sagdstegui 9035 (MO);
Guzmango, Sagdstegui & Lopez 10560 (MO). San Mi-
guel: Cerro Quillon, Sagdstegui 9603a (F, MO), 6 36 3 (MO).
LA LIBERTAD. Huamachuco: Nevado Huaylillas, Ric-
cio & La Rosa 3626 (us). Otusco: Agallpampa, Lopez
341 (USM). Santiago de Chuco: Cerro La Botica, Lopez
1016 (F, USM); Sagdstegui et al. 11885 (F). NO EXACT
LOCALITY. Ellenberg 1688 (USM).
9. Jungia vitocensis Cuatr., Anales Ci. Univ. Ma-
drid, Fasc. 2. 230. 1935. TYPE: Peru, Junin,
Tarma, camino a Vitoc, en las alturas de Pal-
ca, 4 Nov 1863, /. Isern 462 (holotype, MA,
not seen; isotype fragment F 843173 ex MA).
Annual or perennial herbs, erect, branched, the
stems tomentose. Leaves petiolate; petioles 10-
40(-170) mm long, tomentose; blades orbicular,
70-1 10(-180) mm long, 70-100(-180) mm wide,
the adaxial surfaces pilose to strigillose, the abaxial
surfaces densely to sparsely tomentose, the mar-
gins ca. 9-lobed, the lobes rounded, mucronate-
dentate. Capitulescences corymbose to paniculate,
bracts to 25 mm long, linear. Capitula peduncles
20-25 mm long, tomentose; involucres campan-
ulate; phyllaries 12-14, lanceolate, densely villous,
acuminate, yellowish, the inner 6-7 mm long, 1-
1.5 mm wide; florets 14-18, the corollas bilabiate,
6-7 mm long, the tube 2.5-3.5 mm long, the outer
lip ovate, tridentate, the inner lip lanceolate, bifid.
Achenes cylindric, ca. 3.5 mm long, 5-costate, pi-
lose; pappus of ca. 30 bristles, 5-6 mm long, stra-
mineous.
DISTRIBUTION— Known from the eastern slopes
of the Andean Cordillera of central Peru.
Jungia vitocensis is readily distinguished by its
densely villous phyllaries and long-pedunculate
capitulescences. This is apparently a rare species
and is potentially related to /. rugosa, from which
it differs in possessing more orbicular leaves and
pilose achenes.
HUANUCO. Huanuco: Huacachi, estacion near Mufia,
Macbride 4138 (F, us).
10. Jungia weberbaueri Cerrate, Publ. Mus. Hist.
Nat. "Javier Prado," Ser. B, Bot. 4: 22. 1951.
TYPE: Peru, Huanuco, Pachitea, [Rio] Hua-
llaga, Chaglla, A Weberbauer6701 (holotype,
MOL; isotype, F 62861 1; photograph ex F, FM
neg. 1 504494). Figure 10.
Shrubs 1 5-20 dm high, scandent, branched, the
branches terete, pubescent, ochraceous. Leaves
petiolate; petioles 20-55 mm long, pubescent,
paired stipuliform bracts; blades triangular-cor-
date to ovate-cordate, 35-140 mm long, 25-92
mm wide, ochraceous, palmi-nerved, the bases
cordiform, the apices obtuse, the adaxial surfaces
densely strigose, the abaxial surfaces densely to-
mentose, the veins prominent, the margins sub-
entire to remotely lobed. Capitulescences panic-
ulate, terminal or axillary. Capitula with involu-
cres 3.5-5 mm high, 4.5-5.5 mm wide, campan-
ulate; phyllaries 2-seriate, the inner 4.2-4.5 mm
long, 2.2-2.4 mm wide, apically obtuse to trun-
cate, mucronate, strigulose pubescent, ochraceous,
coriaceous; florets 1 1-12, the corollas violaceous,
to 8 mm long, the tube 3-5 mm long, the outer
lip 2.8-3 mm long, 4-nerved, tridentate, the inner
lip 2.8-3 mm long, bifid; anthers 4-4.2 mm long.
Achenes cylindric, 1-2 mm long, glabrous; pappus
of 58-61 bristles, ca. 7 mm long, yellowish-brown.
DISTRIBUTION— Known from the eastern slope
of the Andean Cordillera of central Peru (2800-
2900 m).
Jungia weberbaueri is distinctive among the Pe-
ruvian species in possessing ochraceous, triangu-
lar-cordate to ovate-cordate leaf blades. The in-
volucre of J. weberbaueri most closely resembles
that of J. discolor in possessing phyllaries with
obtuse to truncate apices, but the latter species has
rotund-cordate, strongly discolorous leaves.
HUANUCO: Huanuco: Carpish, between Huanuco and
Tingo Maria, Ferreyra 2098 (MO, us, USM), 2294 (MO,
us, USM).
48
FIELDIANA: BOTANY
FIG. 10. Jungia weberbaueri. 1, habit; 2, capitulum; 3, floret with achene. (Drawn from /. Sanchez et al. 6068, F.)
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
49
3cm
FIG. 1 1. Flowering individual Leucheria daucifolia.
(Drawn from D. Stafford 686, F.)
XII. LEUCHERIA
Leucheria Lag., Amer. nat. Espafi. 1: 32. 1811.
TYPE: Leucheria hieracioides Cass.
Chabraea DC, Ann. Mus. Nat. Hist. 19: 65, 71. t.
13. 1812. TYPE: Perdicium purpureum Vahl. =
Chabraea purpurea (Vahl) DC. (non Chabraea
Adanson, 1763).
Annual or perennial herbs, lanuginose to glan-
dulose; stems lacking spines. Leaves in basal ro-
sette, rarely alternate; blades linear, lanceolate,
spathulate, oblong or ovate, the margins entire,
dentate, or pinnate- to bipinnate-lobed. Capitu-
lescences of solitary heads or terminal and/or ax-
illary cymose-corymbs or cymose-paniculate. Ca-
pitula homogamous, conspicuously pedunculate;
involucres campanulate to hemispheric; recepta-
cle slightly convex to plane, glabrous or paleate;
phyllaries 2-3-seriate, the outer phyllaries lanceo-
late, acute, lanuginose or glabrescent, the inner
smaller; florets isomorphic, hermaphroditic, the
corollas white, the limb bilabiate, the outer lip
ligulate, 3-dentate, the inner lip bifid, coiled; an-
thers linear, sagittate; styles cylindric, the branches
truncate. Achenes ovate or oblong, papillose or
villose; pappus of plumose bristles, isomorphic,
united at base.
Leucheria is a genus of some 46 species and
exclusively distributed in South America from
central Peru to Tierra del Fuego (0-5000 m). The
genus is easily recognized by its basal rosette of
bipinnatifid leaves, white corollas, and plumose
pappus bristles. Leucheria is readily distinguished
from Perezia, in possessing 1-2-seriate involucres
rather than the 3-6-seriate involucres in the latter
genus. Only the following species is known from
Peru.
Reference
CRISCI, J. G. 1976. Revision del genero Leu-
cheria (Compositae: Mutisieae). Darwiniana, 20:
9-126.
1 . Leucheria daucifolia (D. Don) Crisci, Darwin-
iana 20: 52. 1976. Figure 11.
Ptilurus daucifolius D. Don, Trans. Linn. Soc. London
16:219.1830. TYPE: Peru, "summis alpibus cor-
dilleras de los Andes Hispanice dictis," Ruiz Lo-
pez & Pavon s.n. (holotype, P, not seen).
Chabraea daucifolia (D. Don) Wedd., Chi. And. 1 : 35.
1855.
Chabraea laciniata Wedd., Chlor. And. 1: 34, t. 10.
1855. TYPE: Peru, no exact locality, Gay s.n.
(holotype, P, not seen). Non Leuceria laciniata
Hook. & Arn., 1835.
Perennial, cespitose herbs. Leaves petiolate, pet-
ioles 20-40 mm long; blades oblong in outline,
30-50 mm long, 10-15 mm wide, membrana-
ceous, the margins bipinnatifid. Capitulescences
cymose-corymbiform. Capitula 4-12, peduncu-
late, densely pubescent at the base; involucre cam-
panulate to hemispherical; receptacle slightly con-
vex, glabrous; phyllaries 2-3-seriate, lanceolate,
12-15 mm long, 3-5 mm wide lanuginose-fusces-
cent, acute; florets ca. 150, the corollas white, bi-
lobed, the lobes coiled; anthers ca. 3 mm long,
bases sagittate; styles filiform, the branches ca. 1
mm long. Achenes 2-3 mm long, oblong.
DISTRIBUTION— Known from puna within the
Andean Cordillera of central Peru to central Bo-
livia (3800-4800 m).
Leucheria daucifolia is separated from all other
members of the tribe as perennial herbs with a
basal rosette of bipinnatifid leaves, pedunculate
capitulescences, and florets with white corollas,
truncate style branches, and plumose pappus bris-
tles. While no authentic type material has been
examined in the current study, there can be no
50
FIELDIANA: BOTANY
confusion as to the application of the name when
Crisci (1976) is followed.
ANCASH. Bolognesi: above Chiquian, Weberbauer
224 (F). Recuay: Huascaran National Park, Smith & Es-
calona 10206 (F). AREQUIPA. Arequipa: Shihuata-La
Cumbre, Ferreyra 2599 (us, USM), C. & B. Maguire 44453
(us); Nevado Chachani, Pennell 13309 (F, GH); Nevado
Chachani, Rauh & Hirsch P552 (NY); Arequipa market,
Rose 18817 (us); Pichu Pichu, Stafford 686 (F); between
Arequipa to Puno, Straw 2307 (us, USM). Nevado Cha-
chani, Straw 2333 (USM). CUSCO. Chumbivilcas: Sto
Tomas, Hoogte & Roersch 1580 (F). Espinar: Yauri, Car-
Her 195 (USM); Wara-Wara, Tocroyoc, Hoogte & Roersch
2364 (F). JUNIN. Huancayo: Huaytapayana, Rauh &
Hirsch PI 776 (NY). Junin: Huampucocha, Cerrate s.n.
(USM 32738). Yauli: above Yauli, Weberbauer 222 (F).
LIMA. Canta: Culluay, Capt. Wilkes Exped. s.n. (GH).
Huarochiri: above Tucto, Lima-Oroya, Duncan 2675
(F, MO); Casapalca, Macbride & Featherstone 865 (F, us);
above Casapalca, Caprichosa, Amstuts 137 (us). PAS-
CO. Daniel Carrion: Cordillera Huayhuash, Adcook s.n.
(us). PUNO. Carabaya: Macusani, Soukup 474 (F); Bal-
canpata-Apacheta, Vargas 17618 (us). NO EXACT LO-
CALITY. Rauh 552 (USM), 7776 (USM).
XIII. LYCOSERIS
Lycoseris Cass., Diet. Sci. Nat. 33: 474. 1824.
TYPE: Atractylis mexicana L. fil. = Lycoseris
mexicana (L. fil.) Cass.
DiazeuxisD. Don, Trans. Linn. Soc. London 16: 251.
1830. TYPE: Diazeuxis mutisiana D. Don = Ly-
coseris mexicana (L. fil.) Cass.
Dioecious, perennial herbs, lianas, or sub-
shrubs; stems scandent, lacking spines. Leaves al-
ternate, simple, shortly petiolate to subsessile;
blades ovate to lanceolate or elliptic, 3-5-penni-
nerved (camptodromous) or strongly 3 -nerved
from near the base (perfect to imperfect acrod-
romous), the adaxial surfaces glabrescent, the ab-
axial surfaces arachnoid to lanuginose, white, the
margins entire to remotely denticulate. Capitu-
lescences of solitary, terminal heads, rarely mul-
tiheaded, corymbose or racemose. Capitula func-
tionally unisexual: involucres campanulate to glo-
bose; receptacle convex, pubescent with needlelike
bristles between florets; phyllaries 6-8-seriate, im-
bricate, the outer lanceolate, the inner linear to
oblong, apically acuminate to obtuse, appressed
to reflexed; staminate heads radiate; ray florets
pistillate, 1 -seriate, the ovaries rudimentary, the
corollas orange to orange-red, occasionally yellow
or violet, bilabiate, the outer lip ligulate, (l-)3(-
5)-dentate; disc florets tubular, the corollas 5-lobed,
orange; anthers basally sagittate, the apical ap-
pendage lanceolate; styles cylindric, the style
branches connate; pistillate heads radiate or dis-
coid; ray florets with bilabiate corollas or absent;
disc florets narrowly tubular, the limb 5-lobed,
actinomorphic, the stamens rudimentary, ovary
well developed, style exserted, branches short,
rounded, papillose. Achenes columnar, glabrous,
5-costate; pappus of 150-200 bristles, 1 -seriate,
yellowish-white.
Lycoseris consists of 1 1 species of dioecious sub-
shrubs distributed from Guatemala to northwest-
ern and western South America. The genus is dis-
tinctive among the Peruvian members of the tribe
in possessing scandent or vining habits, rather large
capitula with orange-yellow to orange-red corollas,
and a distribution below 1 000 m along the eastern
escarpment of the Andean Cordillera.
Reference
EGEROD, K., AND B. STAHL. 1991. Revision of
Lycoseris (Compositae— Mutisieae). Nord. J.
Hot., 11: 549-574.
Key to Peruvian Species of Lycoseris
1 . Leaf blades narrowly ovate to lanceolate; phyllaries reflexed L. peruviana
1 . Leaf blades elliptic to narrowly lanceolate; phyllaries appressed L. trinervis
Lycoseris peruviana Egerod, Nord. J. Bot. 1 1:
564. 1991. TYPE: Peru, Dept. San Martin,
Pasaraya to Saposoa, 700 m, 6 Aug 1958,
[male], F. Woytkowski 5080 (holotype, F; iso-
types, NY, us).
Procumbent shrubs, or lianas, branched, the
branches lanuginose, striate. Leaves subsessile or
petioles 5-8 mm long; blades narrowly ovate to
lanceolate, 55-150 mm long, 25-55 mm wide, ba-
sally obtuse to rounded, apically acute, venation
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
51
perfect suprabasal acrodromous. Capitulescences
of solitary, terminal heads, subsessile and usually
subtended by reduced leaf. Staminate capitula ca.
1 5 mm high, 1 2-20 mm wide; phyllaries 5-6-se-
riate, strongly reflexed, the outer linear-lanceolate
to spathulate, ( 1 0-) 1 5-25 mm long, 2-3 mm wide,
the inner oblanceolate, 1 5-20 mm long, ca. 5 mm
wide, reflexed; disc florets 87-90, the corollas 1 1-
12 mm long, the lobes 3-3.5 mm long, the anthers
6-6.5 mm long. Pistillate capitula 20-25 mm high,
20-30 mm wide; phyllaries 6-8-seriate, reflexed,
the outer lanceolate, 12-16 mm long, 2.3-2.8 mm
wide, acute, the inner gradually longer, 25-28 mm
long, ca. 3 mm wide; ray florets 1 9-20, the corollas
1 8-20 mm long, the tube 8-9 mm long, the ligule
10-11 mm long, 2.2-3 mm wide. Achenes 4-6 mm
long; pappus 12-15 mm long.
DISTRIBUTION— Endemic to quebradas and low
forests along the valley of the Rio Huallaga in the
Department of San Martin (200-900 m).
Lycoseris peruviana is weakly distinguished from
L. trinervis by the former's narrowly ovate to lan-
ceolate leaf blades, short petioles, smaller female
capitula, and longer, strongly reflexed phyllaries.
SAN MARTIN. Lamas. San Jose de Sisa, Ferreyra
7942 (MO, us, USM); 20 km SE of Moyobamba, Woyt-
kowski 35256 (F). San Martin: W of Shapaja 2-8 km on
trail to Tarapoto, Belshaw 3200 (F, MO, NY), 3201 (F);
Pucayacu, 1 1 km S of Tarapoto, Ferreyra 7743 (F); near
Tarapoto, Ferreyra 7749 (MO, USM), 5005 (MO, USM);
Puente Colombia, Ferreyra 17521 (USM); 31 km S of
Tarapoto, Gentry et al. 37734 (F); Tarapoto, Williams
5488 (F, us), 5744 (F, us), 5557 (F, us).
2. Lycoseris trinervis (D. Don) S. F. Blake, Contr.
U.S. Natl. Herb. 22: 653. 1924. Figure 12.
Diazeuxis trinervis Ruiz Lopez & Pavon ex D. Don.
Trans. Linn. Soc. London 16: 253. 1830. TYPE:
based upon Aster trinervis Ruiz Lopez & Pavon,
ined. LECTOTYPE (designated by Egerod and
Stahl, 1991): Ecuador, Prov. Guayas, "Aster de
Huayaquil, Sp. nova," J. Tafalla s.n. (lectotype,
BM, not seen).
Centroclinium altissimum Poepp. & Endl., Nov. gen.
sp. pi. 3: 52, t. 259. 1843. TYPE: Peru, "Tocache
Huahaga, 1 830," [female], E. F. Poeppig 163 (ho-
lotype, w, not seen; isotype, NY).
Onoseris altissima (Poepp.) Kuntze, Revis gen. pi. 354.
1891.
Lycoseris trinervis (D. Don) S. F. Blake subsp. altis-
sima (Poepp.) Egerod, Nord. J. Bot. 1 1 : 56 1 . 1 99 1 .
Shrubs scandent or subscandent, branched, the
stems lanuginose, glabrescent. Leaves subsessile,
or petioles 5-13 mm long; blades oblong-lanceo-
late to elliptic, 90-180 mm long, 20-50 mm wide,
basally attenuate to truncate, apically acuminate,
venation perfect suprabasal acrodromous. Capitu-
lescences of solitary, terminal heads, the peduncles
15-25 mm long. Staminate capitula 11-16 mm
high, 1 2-24 mm wide; phyllaries 5-6-seriate, ap-
pressed, the outer lanceolate, 11-15 mm long, ca.
2.5 mm wide, the inner linear-lanceolate to ob-
long, 1 5-20 mm long, ca. 1.5 mm wide; disc florets
ca. 100, the corollas ca. 20 mm long, unequally
5-lobed, the lobes 1.5-2 mm long, the anthers ca.
6 mm long. Pistillate capitula 22-40 mm high,
15-30 mm wide; phyllaries 8-9-seriate, the outer
lanceolate, ca. 15 mm long, ca. 3.2 mm wide, the
inner linear-lanceolate, 20-22 mm long, 2.8-3 mm
wide, acuminate, glabrous; ray florets ca. 21, the
corollas subliguate, ca. 20 mm long, the tube ca.
15 mm long, the ligule ca. 5 mm long, 1-2 mm
wide; disc florets 260-270, the corollas 1 8-20 mm
long, the lobes ca. 1.5 mm long. Achenes 3.5-4
mm long; pappus 16-18 mm long.
DISTRIBUTION— Known from coastal Ecuador
and disjunct to the eastern escarpment of the An-
dean Cordillera of north-central Peru (350-900
m).
Lycoseris trinervis is distinguished by its elliptic,
strongly trinerved leaves, large pistillate capitula
with up to 270 florets, and appressed phyllaries.
Material annotated by Egerod and Stahl (1991)
has been examined, and the application of this
name appears unequivocal. Egerod and Stahl
(1991) treated the Ecuadorian and Peruvian col-
lections as a subspecies trinervis and altissima, re-
spectively; the morphological distinctions appear
so weak, however, that the subspecific taxa are not
recognized here. Furthermore, with some hesita-
tion, Egerod and Stahl (1991) placed two central
Peruvian collections (Soukup 2422, Sandeman
4992) under L. retrqflexa Koster, a central Boli-
vian species. Because these collections are stated
to lack certain of the denning characteristics of L.
retrojlexa, that species is not recognized from Peru
in this treatment and is here placed under L. tri-
nervis.
HUANUCO. Leoncio Prado: 69 km NE of Tingo Ma-
ria on road to Tocache, Gentry et al. 37630 (F). Pachitea:
Pozuzo, Macbride 4593 (F, GH, us). JUNIN. Satipo: Sa-
tipo, Rio Chiquireni, Madison 10455-70 (us, USM). Tar-
ma: between La Merced to Quimiri, Ferreyra 3689 (us,
USM); 10 km S of San Ramon, Gentry et al. 41529 (F,
MO, USM); Chanchamayo, hern 37 (F 157054, F 1570155);
Quimiri-La Merced, Killip & Smith 23933 (us); San Ra-
mon, Killip & Smith 24741 (F, us); Colonia Perene, Killip
& Smith 25099 (us); Chanchamayo Valley, Schunke 1607
52
FIELDIANA: BOTANY
FIG. 12. Lycoseris trinervis. 1, flowering branch of female individual. (Drawn from 5. Knapp 8160, F.) 2, flowering
branch of male individual. (Drawn from /. Schunke 4880, F.)
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
53
(F); San Ramon, Soukup 1835 (GH, us); San Ramon,
Tovar 2288 (USM); Woytkowski 7402 (GH, us). LORE-
TO. Alto Amazonas: Santa Rosa, lower Rio Huallaga
below Yurimaguas, Killip & Smith 28721 (us). SAN
MARTIN. Mariscal Caceres: 4 km de Puerto Pizana,
Schunke 4480 (F, us). San Martin: Tarapoto, Ferreyra
7877 (us, USM), 17413 (MO, USM); Chazuta, Klug 4148
(F, GH, MO, NY); Chazuta, Knapp 8160 (F); Huaquisha,
Schunke 7084 (MO, NY, USM), 72577 (F); Tocache Nuevo,
Schunke 12428 (USM). UCAYALI. Coronel Portillo: Bos-
que Nacional Alexander von Humboldt, Gentry et al.
41415 (F); road to Tournavista, Gentry & Diaz 58357
(F); Pucallpa, Soukup 3063 (f). NO EXACT LOCALITY.
"Peru," Cuming 1160 (F), 7767 (F).
XIV. MUTISIA
MutisiaL. fil., Suppl. pi. 57. 1781. TYPE: Mutisia
clematis L. fil.
Perennial herbs, shrubs, or vines; stems with or
without wings, lacking spines. Leaves alternate,
sessile or shortly petiolate; blades simple or pin-
nately compound, apically with or without a ten-
dril, the margins entire to dentate. Capirulescences
of solitary heads, terminal or axillary, or occa-
sionally weakly corymbose; sessile or long-pedun-
culate. Capitula heterogamous, radiate, or ho-
mogamous, discoid; involucres cylindric to cam-
panulate; receptacle plane to convex, foveolate,
glabrous; phyllaries (3-)4-8(-10)-seriate, imbri-
cate; ray florets pistillate, the corollas bilabiate with
the outer three lobes fused into a tridentate ligule
and the two inner lobes free; disc florets her-
maphroditic, the corollas bilabiate, the outer lip
ligulate, apically 3-fid, the inner lip divided deeply,
the lobes filiform; anthers long-exserted, 15-30
mm long, the terminal appendages lanceolate, the
bases sagittate or more typically auriculate, the
tails sometimes fringed; style branches short, trun-
cate to rounded. Achenes columnar or fusiform,
ribbed or smooth, glabrous or pubescent; pappus
of plumose bristles, 1 -seriate, white to yellow or
brown.
Mutisia contains more than 60 species, the ma-
jority of which occur in Chile, Argentina, Peru,
and Bolivia. No fewer than 1 5 species are currently
recorded from Peru. Although type material has
not been studied for some species, material an-
notated by Cabrera (1965) was available for com-
parison.
References
CABRERA, A. L. 1965. Revision del genero Mu-
tisia (Compositae). Opera Lilloana, 13: 1-227.
FERREYRA, R. 1980. Especies nuevas de Com-
puestas Peruanas. Bol. Soc. Peruana Bot. Nos.
1-2, 8: 75-82.
Key to Peruvian species of Mutisia
1 . Lianas or shrubs; leaves pinnate, the leaflets lanceolate, ovate or elliptic 2
2. Median and inner phyllaries with recurved apices; pappus of ligulate florets longer than the tube
3
3. Leaflets lanate; capitula with involucres to 40 mm wide; disc florets ca. 100 M. lanata
3. Leaflets glabrous or glabrescent; capitula with involucres to 30 mm or less wide; disc florets
no more than ca. 60 M. wurdackii
2. Median and inner phyllaries appressed, only outer phyllaries with recurved apices; pappus of
ligulate florets less than half the length of the tube 4
4. Leaves with 9-14 pairs of leaflets M. acuminata
4. Leaves with 3-9 pairs of leaflets 5
5. Leaves with 3-5 pairs of leaflets; disc florets 30-60; inner phyllaries apically obtuse ....
M. pulcherrima
5. Leaves with 8-9 pairs of leaflets; disc florets 10-25; inner phyllaries apically acute
M. venusta
1 . Shrubs; leaves simple, the blades linear to oblong or lanceolate 6
6. Leaf blades apically acute, mucronate, without terminal tendrils 7
7. Leaf blades with abaxial surfaces densely white-tomentose M. ledifolia
1. Leaf blades with abaxial surfaces glabrous M. orbignyana
6. Leaf blades apically attenuate, with terminal tendrils 8
54
FIELDIANA: BOTANY
8. Stems lacking wings 9
9. Leaves to 16 cm long, to 6 mm wide; involucres to 45 mm high, ca. 30 mm wide
M. rauhii
9. Leaves to 9 cm long, to 4 mm wide; involucres to 35 mm high, ca. 10 mm wide
M. mathewsii
8. Stems winged, the wings entire to dentate, rarely reduced or absent (cf. cochabambensis,
lanigerd) 10
10. Leaf margins denticulate M. lanigera
10. Leaf margins essentially entire 11
1 1 . Stems with wings entire; leaves glabrous, bases rounded or cordate M. alata
1 1 . Stems with wings dentate; leaves glabrous or tomentose, bases sagittate to hastate . . .
12
1 2. Leaves shortly petiolate M. cochabambensis
1 2. Leaves sessile 13
13. Leaves with abaxial surfaces glabrous M. arequipensis
1 3. Leaves with abaxial surfaces tomentose 14
14. Phyllaries glabrous, the outer series not recurved M. andersonii
14. Phyllaries lanate, the outer series with strongly recurved apices
. . M. hastata
1 . Mutisia acuminata Ruiz Lopez & Pavon, Syst.
veg. fl. peruv. chil. 192. 1798. TYPE: Peru,
"Ceuchin [Chiuchin] 1 779 mayo," J. Dombey
s.n. (holotype MA, not seen).
Mutisia peduncularisCav., Icon. PI. 5: 62. 1799. TYPE:
Peru, San Buenaventura, Nee s.n. (holotype, MA,
not seen).
Mutisia viciaefolia Cav., Icon. PI. 5: 62. 1799. TYPE:
Chile, Valparaiso, Nee s.n. (holotype, MA, not seen;
isotype fragment ex MA, F 844679).
Shrubs 10-25 dm high, branched, branches stri-
ate, glabrous to lanuginose, erect to procumbent,
sometimes semivoluble, rarely forming rounded
cushions. Leaves sessile, pinnate; rachis with ten-
drils trifid, 7-1 5 cm long, the leaflets in 9-14 pairs,
opposite to alternate, lanceolate or elliptic-lanceo-
late, 10-35 mm long, 5-10 mm long, apically acu-
minate with terminal tendril, gradually attenuate
at base, entire. Capitulescences of solitary, ter-
minal or axillary heads; pedunculate, peduncles
3-15 cm long, tomentose to glabrescent. Capitula
radiate; involucres 45-60 mm high, 7-1 5 mm wide,
cylindric; phyllaries 18-22, 6-7-seriate, oblong-
lanceolate, 6-16 mm long, 4-12 mm wide, apically
obtuse, glabrous to tomentose; ray florets 5-8, the
corollas red, orange, or yellow, the tube 35-50 mm
long, 0.8-1.2 mm wide, the ligule 10-18 mm long,
8-10 mm wide, obtuse, sometimes 2-dentate; disc
florets 10-20, the corollas yellowish or reddish,
bilabiate, the tube 20-30 mm long, the outer lip
tridentate, the inner lip bifid, 4—4.5 mm long; styles
4-5 mm long; anthers ca. 17 mm long. Achenes
fusiform, 8-18 mm long, 1-2 mm wide, glabrous;
pappus 1 8-20 mm long, yellowish brown.
DISTRIBUTION— Known throughout the Andean
Cordillera from northern Peru to Bolivia and ad-
jacent northern Argentina and Chile (1200-3800
m).
Cabrera (1966) recognized five varieties, and
Peruvian material is referable to the following three.
Key to Peruvian Varieties of Mutisia acuminata
1 . Leaflets with abaxial surfaces whitish- to yellowish-tomentose; phyllaries apically white-tomentose;
corollas orange-yellow M. a. var. bicolor
1 . Leaflets glabrous; phyllaries glabrous or yellowish-brown-tomentose; corollas yellow or reddish-yellow
2
2. Involucre cylindric, 45-60 mm high; corollas red to reddish-yellow M. a. var. acuminata
2. Involucre campanulate, 55-60 mm high; corollas yellow M. a. var. hirsuta
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
55
la. Mutisia acuminata var. acuminata.
DISTRIBUTION— Known from the western and
central portions of the Andean Cordillera ( 1 200-
3800 m). The protolog states "habitat in Prov.
Tarmac, Huarocheri et Caxatambo ad Cheuchin
vicum" and the label indicates "Ceuchin," which
is undoubtedly a misspelling of Cheuchin, De-
partment Lima.
Variety acuminata is distinguished by its gla-
brous leaflets and red to reddish-orange florets. It
is used in folk medicine, especially in Cusco, and
has the common name of "chinchircuma." Chro-
mosome number: n = 21.
ANCASH. Bolognesi: Chiquian, Cerrate 1319 (USM);
entre Chachash y Chiquian, Ferreyra 6186 (us, USM),
Mostacero et al. 1337 (F); Cajacay, Ferreyra 6232 (us,
USM); below Puente Santa Rosa, Gentry & Zardini 37316
(F, MO); near Conococha, Lopez et al. 7062 (NY); Pativil-
ca-Huaras, km 99, Smith 8312 (F); 15 km SE of Chi-
quian, Young & Eisenberg 905 (F); Pativilca-Huaras,
Zardini 1492 (MO). Casma: arriba de Chacchan, Mos-
tacero et al. 525 (F). Huaras: Pampas Grande, entre San
Juan y Huinapajatum, Diaz 1978 (F); Huaras-Recuay,
Lopez & Sagdstegui 7485 (M), 7602 (MO); between Casma
and Huaras, Maekawa s. n. (USM). Recuay: Marca, Gomez
120 (USM); Shucta, Sanchez 252 (F). Santa: arriba de
Lamparin, Mostacero et al. 1829 (F). HUANUCO. Hua-
malies: Llata, Macbride & Featherstone 2255 (GH, us).
JUNIN. Huancayo: Huancayo, Soukup 1198 (MO). Tar-
ma: between Tarma and Oroya, Ferreyra 3821 (MO, us,
USM). LIMA. Cajatambo: Churin, Ferreyra 3542 (us,
USM). Canta: below San Juan, Chancay Valley, Ferreyra
18357 (MO, USM); Guamantanga, Mathews 457 (GH); Ay-
asura Meza 04 (USM); near Viscas, Pennell 14318 (GH,
NY, us); Obrajillo, Velarde 951 (USM). Huarochiri: Cha-
cahuaro, Cerrate s.n. (USM); San Lorenzo, Cerrate 1976
(MO, USM), 1045 (MO, USM); 7775 (MO, USM); Rio Blanco,
Cerrate 4285 (USM); San Mateo, Cerrate 5325 (USM); 4
km E of San Mateo, Dillon 2505 (F, MO); near Surco,
Ferreyra 3418 (USM); between Huaquicha and Palacala,
Ferreyra 3455 (MO, USM); San Mateo, Ferreyra 5314 (us,
USM); Infiernillo, Ferreyra 6254 (MO, us, USM); San Ma-
teo, Ferreyra 8295 (MO, USM); between Surco and Ma-
tucana Ferreyra 5414 (USM); Huariquina, Ferreyra 9189
(MO, USM); Matucana, Ferreyra & Tovar 20132 (USM);
Suchi, NE of Chosica, Gentry 21681 (MO), Goodspeed et
al. 11550 (F, uc), Hutchison 604 (uc); Zarate, above San
Bartolome, Hutchison & Wright 7075 (uc); Rio Blanco,
Killip & Smith 21655 (F, us, NY); Matucana, Macbride
& Featherstone 192 (GH, us), Rose 18640 (NY); Matu-
cana, Stork & Morton 9133 (GH, uc); Rio Blanco, Vargas
s.n. (USM); Vargas s.n. (USM); San Pedro de Huancaure,
Soler 28 (USM); between Chosica and Matucana, Tovar
1749 (USM). Lima: Lomas de Asia, Grant 7487 (GH, NY,
us). Yauyos: near Tupe, Tovar 466 (USM).
lb. Mutisia acuminata var. bicolor Cabr., Opera
Lilloana 13: 63. 1966. TYPE: Peru, Dept.
Arequipa, Rio Chulucanas, N. Angulo 1794
(holotype, LP, not seen).
DISTRIBUTION— Endemic to the intermontane
regions of southwestern Peru (2400-3200 m).
Variety bicolor is distinguished by its white-to-
mentose leaflets, densely ferrugino-tomentose
phyllaries, and orange-yellow florets.
APURIMAC. Aimaraes: Chalhuanca, Gentry et al.
23306 (F p.p.). AREQUIPA. Arequipa: near Arequipa,
Arenas 27 (USM), 69 (USM), 767 (USM), Rauh 541 (USM),
Simpson 8567 (USM), Straw 2279 (USM); above Yura,
Straw 2348 (USM). AYACUCHO. La Mar: Tambo Cusi-
cancha, Madison 10398-70 (USM). Lucanas: near Puquio,
Gentry et al. 2 3 28 3 (F, MO). TACNA. Tarata: near Tarata,
Muller 3697 (USM), Rauh 690 (USM).
lc. Mutisia acuminata var. hirsuta (Meyen) Cabr.,
Opera Lilloana 13: 59. 1966.
Mutisia hirsuta Meyen, Reise um die Erde 1: 451.
1834. TYPE: Peru, Meyen s.n. (holotype, B, pre-
sumably destroyed; photograph ex B, FM neg.
15934).
DISTRIBUTION— Common along the Andean
Cordillera from north-central Peru to northern
Bolivia and adjacent northern Chile (2000-4000
m).
Variety hirsuta has glabrous leaves, yellowish-
orange florets, and conspicuous yellowish-brown-
tomentose phyllaries.
APURIMAC. Aimaraes: Chalhuanca, Gentry et al.
23306 (F p.p., MO). Andahuaylas: near Andahuaylas,
West 3728 (uc). AREQUIPA. Arequipa: slopes of
Chachani, Hutchison & Wright 7222 (F, MO, NY, uc, us,
USM); 1 2 mi NE of Arequipa, Maguire & Maguire 44447
(F, NY, USM). Cailloma: Valle del Colca, Treacy 656 (F);
Qayra, Treacy 791 (F). AYACUCHO. Lucanas: San An-
tonio to Malaya, Metcalf 30310 (uc). CAJAMARCA.
Contumaza: Cascas, Lopez & Sagdstegui 9051 (F, MO);
arriba de Trinidad, Sagdstegui et al. 8955 (F); Guzman-
go, Sagdstegui et al. 9211 (F); La Montana, Sagdstegui
et al. 9313 (F); Contumaza-Cascabamba, Sagdstegui et
al. 9996 (F, MO). CUSCO. Calca: Pissac, Marin 199 (USM),
Ugent 3874 (USM). Canchis: Sicuani, Cook & Gilbert 126
(GH); Sicuani to Marangani, Dillon et al. 1055 (F, NY);
Marangani-Sicuani, Rauh & Hirsch 690 (NY, USM). Cus-
co: Sacsayhuaman, Ferreyra 17099 (USM); Huancar,
Muller & Gutte 9420a (USM); San Sebastian, Pennell 13601
(F, GH, NY); San Geronimo, Solomon 3002 (MO, USM).
Urubamba: Tarapata, above Ollataytambo, Plowman &
Davis 4743 (GH); near Cusco, West 8054 (GH, MO, uc).
HUANCAVELICA: Huancavelica: Yauli, Stork & Nor-
ton 10876 (GH, uc). JUNIN. Jauja: between Araura and
Llocllapampa, Sounders 482 (F); km 61 between Jauja
and La Oroya, Smith 1859 (F, USM). Tarma: Tarma,
Constance & Tovar 2211 (uc); 19 km NE of Tarma,
Dillon & Turner 1328 (F); 1 1 km W of Tarma, Edwin &
Schunke 3884 (F, USM); entre Tarma and Oroya, Ferreyra
3781 (USM); Tarma, Hutchison 627 (uc); Oroya, Kalen-
born 153 (GH); near Tarma, Killip & Smith 21846 (us);
56
FIELDIANA: BOTANY
hills above Tarma, Landrum 4636 (F, NY), Macbride &
Featherstone 1014 (GH, us); Tarmatambo, Tovar 1087
(USM). LA LIBERTAD. Santiago de Chuco: Santiago de
Chuco, Sagdstegui et al. 11765 (F). LIMA. Huarochiri:
5 km W of Matucana, Edwin & Schunke 3792 (F, NY);
1 1 km E of San Mateo, Edwin & Schunke 3800 (F, NY);
between San Mateo and Matucana, Ferreyra 7697 (us,
USM); 2 km W of Matucana, Hutchison 1044 (F, GH, MO,
uc); Matucana, Soukup 2081 (us); Rio Blanco, Vargas
s.n. (USM). MOQUEGUA. Mariscal Nieto: between To-
rata and Carumas, Dillon et al. 3330 (F), 4805 (F).
2. Mutisia alata Hieron., Bot. Jahrb. Syst. 19: 73.
1894. TYPE: Ecuador, Prov. Azuay, "inter
Huasi-huaico et Contrayerba," 2900-3400 m,
F. C. Lehmann 4571 (holotype, B, presum-
ably destroyed; photograph ex B, FM neg.
15921; isotypes, F 6851 13, GH, us).
Scandent shrubs, to 5 m high; stems 3-winged,
the wings entire glabrous, 3-6 mm wide. Leaves
simple, sessile; blades linear- lanceolate, 80-120
mm long, 10-20 mm wide, apically attenuate, ten-
dril elongate, curved, basally amplexicaul, gla-
brous, the margins entire, revolute. Capitules-
cences of solitary heads, erect. Capitula radiate;
involucres campanulate, 30-40 mm high, 12-20
mm wide; phyllaries 5-6-seriate, the outer ovate,
5-10 mm long, 10-12 mm wide, the apical ap-
pendage attenuate, 7-10 mm long, squarrose, the
inner ovate-lanceolate, 35-40 mm long, 10-12 mm
wide, apically obtuse, purplish, mucronate; ray flo-
rets 8-10, the corollas orange, bilabiate, the tube
20-24 mm long, ligulate, the ligule lanceolate, 35-
40 mm long, 7-10 mm wide, tridentate, the inner
lip bifid; disc florets 30-35, the corollas bilabiate,
35-40 mm long, yellow. Achenes ca. 5 mm long,
cylindric, costate: pappus 20-25 mm long, tawny.
DISTRIBUTION— Known from the intermontane
region of northern Peru and southern Ecuador
(2000-4000 m).
Mutisia alata most closely resembles M. an-
dersonii but is distinguished from that species in
possessing leaf blades with glabrous abaxial sur-
faces and rounded basal auricles.
CAJAMARCA. Celendin: between Cajamarca and
Celendin, Gibbs G74-12 (F); entre Celendin-Gelig y Bal-
sas, Sanchez 478 (F). Chota: Huambos, Ochoa 1594 (us);
without locality, Hutchison Wright 5242 (uc, USM).
3. Mutisia andersonii Sodiro ex Hieron., Bot.
Jahrb. Syst. 29: 80. 1900. TYPE; Ecuador,
Pichincha, Pangor, Sodiro 63/1 (holotype, B,
presumably destroyed, photograph ex B, FM
neg. 15923).
Suffrutescent vines, climbing, branched, branch-
es lanuginose, glabrescent, winged, the wings nar-
row, dentate. Leaves simple, sessile; blades linear-
lanceolate, 70-1 10 mm long, 8-10 mm wide, api-
cally attenuate, tendril simple, curved, basally am-
plexicaul, the adaxial surfaces reticulate- veined,
the abaxial surfaces densely lanuginose, the mar-
gins entire, revolute. Capitulescences of solitary,
terminal heads; peduncles 10-30 mm long. Ca-
pitula radiate; involucres campanulate, 35-45 mm
high, 1 5-28 mm wide; phyllaries 6-7-seriate, red-
dish, glabrous, the outer ovate 10-12 mm long,
ca. 1 0 mm wide, apically lanceolate, the inner ob-
long, ca. 30 mm long, 6-7 mm wide, glabrous,
apically obtuse, mucronate; florets dimorphic; ray
florets 10-15; corolla yellow, rarely orange, bila-
biate, ligulate, the tube 20-22 mm long, ligule lan-
ceolate, tridentate, 30-40 mm long, 5-8 mm wide,
inner lip bifid; disc florets with corollas bilabiate,
the tube 20-22 mm long, outer lip 12-14 mm long,
tridentate, the inner lip bifid. Achenes cylindric,
ca. 5 mm long; pappus 25-30 mm long.
DISTRIBUTION— Known from the Andean Cor-
dillera of central Ecuador and disjunct to south-
central Peru (2000-4000 m).
HUANCAVELICA. Tayacaja: Hacienda Huari, Ve-
larde 2100 (USM). LIMA. Canta: "Chipraca,'Mcfe/0 733
(USM); Purumarca, near Canta, Sanchez 01 (USM).
4. Mutisia arequipensis Cabr., Opera Lilloana 1 3:
171. 1966. TYPE: Peru, Arequipa, Conde-
suyos, encima de Salamanca, 3900—4000 m,
A. Weberbauer 6857 (holotype, GH; isotypes,
us 1473434, USM).
Suffrutescent vines, climbing or procumbent, the
branches glabrous, winged, the wings dentate.
Leaves simple, sessile; blades linear-lanceolate, SO-
SO mm long, 3-4 mm wide, inconspicuously has-
tate, apically attenuate, terminal tendril simple,
basally sagittate, the margins entire or with scat-
tered denticulations. Capitulescences of solitary,
terminal heads; peduncles 5-10 mm long. Capit-
ula radiate; involucres campanulate, 20-30 mm
long, 8-10 mm wide; phyllaries 4-5-seriate, api-
cally lanuginose, the outer ovate, 2—4 mm long,
ca. 5 mm wide, the apical appendage coiled, 3-4
mm long, the inner oblong-ovate, 1 5-20 mm long,
7-8 mm wide, apically obtuse; florets dimorphic;
ray florets 9-10, the corollas red, the ligules lan-
ceolate, tridentate, 15-17 mm long, 3-3.5 mm
wide, inner lip bisect; disc florets 1 5-20, the co-
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
57
rollas bilabiate, the tube 20-22 mm long, the outer
lip tridentate, the interior bifid. Achenes cylindric,
ca. 3 mm long, glabrous; pappus ca. 1 8 mm long.
DISTRIBUTION— Endemic to the puna of central
Peru (3400-4000 m).
Mutisia arequipensis closely resembles M. an-
dersonii and M. hastata but differs in possessing
a glabrous habit, narrower leaves, and reddish flo-
rets. It is also close to M. mathewsii but distin-
guished from that species by its dentate stem wings
and leaves with plane margins.
AREQUIPA. Cailloma: Valle del Colca, ca. 100 km
N de Arequipa, Treacy 829 (F), 840 (F).
5. Mutisia cochabambensis Hieron., Bot. Jahrb.
Syst. 19: 74. 1894. TYPE: Bolivia, Cocha-
bamba, N. H. Bang 881 (holotype, B, presum-
ably destroyed; isotypes, GH, MO, NY, us).
Suffrutescent herbs, climbing, branched, the
branches narrowly winged or unwinged, densely
leafy, glabrous. Leaves simple, subsessile; petioles
1-2 mm long; blades linear to lanceolate, 60-100
mm long, 5-1 5 mm wide, the adaxial surfaces gla-
brous or tomentose, apically attenuate with curved,
solitary tendril, bases sagittate, the lobes acute, the
margins entire. Capitulescences of solitary, ter-
minal heads; peduncles 15-25 mm long. Cap! tula
radiate; involucre campanulate, 25-35 mm high,
10-15 mm wide; phyllaries 16-20, 3-4-seriate,
reddish, the outer narrowly ovate, 5-10 mm long,
7-8 mm wide, apical appendage apiculate, the in-
ner oblong, 25-30 mm long, 8-10 mm wide, api-
cally obtuse, lanuginose; florets dimorphic; ray flo-
rets 9-10, orange, the ligule lanceolate, tridentate,
ca. 25 mm long, 4-6 mm wide, inner lip bisect;
disc florets bilabiate, the tube 20-24 mm long, the
outer lip tridentate, the inner lip bisect. Achenes
cylindric, 4-5 mm long, striate; pappus ca. 25 mm
long, white.
DISTRIBUTION— Known from the Andean Cor-
dillera of southeastern Peru to central Bolivia
(3200-3700 m).
Mutisia cochabambensis most closely resembles
M. mandoniana Wedd. ex Schultz-Bip. but differs
in possessing shortly petiolate, strictly glabrous
leaves and much smaller heads.
APURIMAC. Aimaraes: Lucre, Abra Ratcay, Vargas
1616 (GH). CUSCO. Calca: Totora, Mann 2130 (USM).
Espinar: Yauri, Virginniyoc, Nunez et al. 7891 (F). Pa-
ruro: Pucaccasa, Vargas 1195 (MO, uc). Quispicanchis:
Hacienda Ccapana, Herrera 1128 (us), 1051 (GH); Oro-
pesa, Balls 6833 (uc, us). Urubamba: Gutierrezchayoj,
King et al. 288 (F); trail up Antakillqu hillside, Davis et
al. 1688 (F). TACNA. Tacna: NE of Tarata, Pearson &
Pearson 1 155 (uc).
6. Mutisia hastata Cav., Icon. PI. 5: 64. 1799.
TYPE: No exact locality, "ex Cordillera del
Planchon," L. Nees.n. (holotype, MA, not seen,
isotype ex MA, F 845291).
Suflruticose vines, procumbent; stems 4-winged,
conspicuously dentate, the teeth triangular, acute,
retrorse, 3-5 mm long, lanuginose to glabrescent.
Leaves simple, sessile; blades linear-lanceolate, 50-
100 mm long, 5-10 mm wide, apically attenuate
with curved solitary tendril, the bases sagittate with
acute lobes, the adaxial surfaces glabrous, the ab-
axial surfaces densely tomentose, the margins en-
tire to distally dentate, revolute. Capitulescences
of solitary, terminal heads; peduncles 1 0-40 mm
long. Capitula radiate; involucres campanulate, 35-
40 mm high, 15-20 mm wide; phyllaries 8-10-
seriate, reddish, the outer ovate, ca. 10 mm long,
ca. 5 mm wide, apically lanceolate or linear, re-
curved, lanuginose, mucronate, the inner ovate-
oblong, 25-30 mm long, 10-12 mm wide obtuse,
mucronate, apices lanuginose; florets dimorphic;
ray florets ca. 10, the corollas orange to red, the
tube ca. 12 mm long, the ligule lanceolate, 25-35
mm long, 4-5 mm wide, apically tridentate, the
inner lobe bisect; disc florets with corollas yellow,
bilabiate, the tube ca. 20 mm long, the lobes ca.
1 0 mm long, outer lobe tridentate, the inner lobe
bifid. Achenes cylindric, glabrous; pappus 22-25
mm long, grayish.
DISTRIBUTION— Endemic to the Andean Cor-
dillera of central Peru (3400-3600 m).
Mutisia hastata most closely resembles M. an-
dersonii but differs in possessing outer phyllaries
with strongly recurved and lanose apices. The type
locality was cited as from Chile, "Cordillera del
Planchon"; however, it is likely an error in labeling
and actually collected by Nee in Peru. The type
material at F was sent from MA and has a separate
label indicating the origin of the material as "Ca-
vanilles."
ANCASH. Bolognesi: entre Tallenga y Pachapaque,
Ferreyra & Cerrate 7475 (us, USM). Recuay: Marca, Go-
mez 163 (USM). HUANCAVELICA. Tayacaja: Tran-
capampa, entre Huari y Acobamba, Tovar 4550 (USM).
HUANUCO. Huanuco: Mito, Macbride & Featherstone
1857 (us). LIMA. Canta: Obrajillo, Capt. WilkesExped.
58
FIELDIANA: BOTANY
s.n. (GH, us). Huarochiri: 10 km NE of Suchi, Gentry
21639 (F).
7. Mutisia lanata Ruiz Lopez & Pavon, Syst. veg.
fl. peruv. chil. 1: 192. 1798. TYPE: Peru,
Huanuco, "Habitat in Peruviae nemoribus
prope Muna vicum in Pozuzo via," Ruiz Lo-
pez & Pavon s.n. (holotype, MA, not seen; iso-
type, BM, not seen).
Suffruticose herbs, scandent, to 10 m high, the
stems narrowly winged, tomentose. Leaves sessile,
pinnate; rachis 8-20 cm long, apex with trifid ten-
dril, the leaflets 4-5 pairs, ovate to lanceolate, 45-
1 00 mm long, 1 7-35 mm wide, alternate or subop-
posite, apically acute to obtuse, basally rounded,
the adaxial surfaces albo-arachnoid to glabrescent,
the abaxial surfaces ferrugino-tomentose, the mar-
gins entire. Capitulescences of solitary heads, more
or less nutant, the peduncles 6-20 cm long, to-
mentose. Capitula radiate: involucres campanu-
late, 35-40 mm high, 30-50 mm wide; phyllaries
6-7-seriate, the outer ovate, the apical appendage
linear-lanceolate densely ferrugino-tomentose,
strongly recurved, the inner oblong, the apical ap-
pendage attenuate, densely ferrugino-tomentose;
florets dimorphic; ray florets 20-25, the ligules
lanceolate, red, tridentate, ca. 40 mm long, ca. 10
mm wide; disc florets 90-100, the corollas yellow,
bilabiate, 12-15 mm long, the outer lip tridentate,
the inner lip bifid. Achenes cylindric, glabrous;
pappus 25-30 mm long.
DISTRIBUTION— Known from the intermontane
region from central Peru to Bolivia (1800-3400
m).
Mutisia lanata is easily distinguished by its con-
spicuous tomentose habit, red ray florets and yel-
low disc florets. While no authentic type material
has been seen in this study, Cabrera (1960) con-
sulted the type of this species in both Madrid (MA)
and London (BM), and comparison with material
annotated by him allows for the application of the
name.
CUSCO. Calca: San Miguel, Cook & Gilbert 1145 (us);
San Miguel Valley, Herrera 2020 (GH); Monte Lares,
Marin 2139 (USM); Lares, Vargas 3593 (MO). Paucartam-
bo: near Paucartambo, Balls 6713 (uc, us); 9-12 km NE
ofPaucartambo, Gentry etal. 23419(p);A.canaco,Molau
774 (USM), Plowman & Davis 4923 (GH, USM); Tres Cru-
ces, Pennell 14142 (GH, NY, us); Sunchubamba, Vargas
2267 (NY); Hacienda Paccho, Vargas 9469 (MO); Pilla-
huata, Velarde 1 165 (USM), Woytkowski 21 (USM); Tam-
bomayo River, West 7096 (uc); Pavayoc, Woytkowski
623 (USM). Quispicanchis: Ocongate to Marcapata, Aron-
son & Berry 523 (F); near Marcapata, Metcalf '30728 (GH,
MO, uc, us); entre Abra Walla Walla y Marcapata, Nunez
et al. 9102 (F); above Marcapata, Wasshausen & En-
carnadon 822 (us); Marcapata, Weberbauer 7880 (GH,
NY, us). Urubamba: Torontoy-Ollantaytambo, Cook &
Gilbert 834 (us); near Urubamba, Herrera 2020 (GH,
USM). HUANCAVELICA.Tayacaja: Huari-Acobamba,
Tovar 4563 (USM). HUANUCO. Huanuco: Huacachi,
near Muna, Macbride 4060 (GH, us). Pachitea: Panao,
Macbride & Featherstone 2223 (GH, us). JUNIN. Huan-
cayo: "Ceja de Montana," Esposto s.n. (USM). Tarma:
between Pinto and Andamarca, Raimondi s.n. (USM).
PASCO. Oxapampa: Los Chacos, near Oxapampa, Smith
& Pretel 1495 (F). PUNO. Carabaya: Ollachea to San
Gaban, Boeke 3048 (us); Ollachea, Soukup 457 (GH, USM).
8. Mutisia lanigera Wedd., Chlor. And. 1: 16.
1855. TYPE: Bolivia, Carangas, d'Orbigny
1341 (holotype, P, not seen; photograph ex p,
FM neg. 38028).
Suffrutescent, scandent to vining herbs, pros-
trate, branched, the branches glabrous, winged, the
wing conspicuously dentate. Leaves simple, ses-
sile; blades lanceolate, 35-60 mm long, 5-12 mm
wide, the margins of basal portion dentate, with
6-8 pairs of teeth, glabrous, apically rounded, ten-
dril simple. Capitulescences of solitary, terminal
heads; peduncles 5-15 mm long, lanuginose. Ca-
pitula radiate; involucres campanulate, 25-40 mm
high, 10-15 mm wide; phyllaries 5-7-seriate, the
outer ovate, 3-6 mm long, 5-6 mm wide, apically
recurved, lanuginose, the inner lanceolate to ob-
long, 25-28 mm long, 7-8 mm wide, lanuginose;
florets dimorphic; ray florets 9-10, the corollas
orange, pistillate, the corollas ligulate, the ligule
elliptic, tridentate, 12-15 mm long, 4-5 mm wide,
the inner lip bisect; disc florets bilabiate, the outer
lip tridentate, the inner lip bifid. Achenes cylin-
dric, glabrous; pappus ca. 20 mm long.
DISTRIBUTION— Endemic to the altiplano of
southern Peru, northern Bolivia, and adjacent Chile
(3 700-4 100m).
Mutisia lanigera is readily recognized by its
prostrate habit, winged branches, and dentate
leaves.
AREQUIPA. Arequipa: Above Chihuata, Straw 2316
(us, USM); between Sumbay and Canahuas, Weberbauer
1388 (USM). Condesuyos: near Chichas, Weibel 07 (USM).
PUNO. Puno: Santa Rosa, near Have, Pearson & Pearson
22 (us).
9. Mutisia ledifolia Dec. ex Wedd., Chlor. And.
1 : 20. 1 855. LECTOTYPE (designated by Ca-
brera, 1965): Bolivia, "cumbre de la cuesta
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
59
de Lagunillas departamento Potosi," d'Orbig-
ny 1272 (lectotype, P, not seen).
Shrubs 5-20 dm high, branched, the branches
glabrous, costate, the branchlets brown, laxly white-
tomentose. Leaves simple, sessile; blades linear-
oblong, 14-40 mm long, 2-5 mm wide, apically
acute, mucronate-spinulose, basally attenuate, the
adaxial surfaces glabrous, the abaxial surfaces
densely white-tomentose, the margins entire, rev-
olute. Capitulescences of solitary, sessile, terminal
heads. Capitula radiate; involucres campanulate,
1 5-20 mm high, 5-7 mm wide; phyllaries 5-seriate,
oblong, 10-14 mm long, 2-3 mm wide, glabrous,
apically slightly lanuginose, obtuse, rarely acute,
mucronate; florets dimorphic; ray florets ca. 6, lig-
ulate, the ligules white, the abaxial surfaces red-
dish, 10-25 mm long, 2-3 mm wide, tridentate,
the tube 10-12 mm long, the inner lip bisect; disc
florets 10-15, yellow, the tube 9-1 1 mm long, bi-
labiate. Achenes cylindric, 9-10 mm long; pappus
10-12 mm long, white to violaceous.
DISTRIBUTION— Known from the altiplano of
southern Peru, central Bolivia, and northern Ar-
gentina (3000-4200 m).
PUNO. Puno: vicinity of Lake Titicaca, Monheim 159
(NY).
1 0. Mutisia mathewsii Hook. & Arn., Companion
Hot. Mag. 1: 107. 1835. TYPE: Peru, Aya-
cucho, Vilcanota, Mathews 1119 (holotype,
BM, not seen; isotypes, GH, P; photograph ex
GH, USM).
Mutisia weberbaueri Muschl., Bot. Jahrb. Syst. 50,
Beibl. Ill: 104. 1913. TYPE: Peru, Ayacucho,
Huamanga, Totorabamba, 3600-3700 m, A. We-
berbauer 5490 (holotype, B, presumably de-
stroyed; isotype F).
Mutisia mathewsii var. anomala Cabr., Opera Lil-
loana 13: 117. TYPE: Peru, Ayacucho, Lucanas,
between Nasca and Puquio, Hutchison 1244 (ho-
lotype, uc; istoypes, F 1569717, us, USM).
Suffrutescent vines, scandent, branched, the
branches striate, glabrous. Leaves simple, sessile;
blades linear, 30-90 mm long, 1.5-4 mm wide,
the adaxial surfaces glabrous, the abaxial surfaces
lanuginose, acuminate with terminal, simple ten-
dril, basally sagittate, the margins entire, revolute.
Capitulescences of solitary, terminal heads, the pe-
duncle 6-20 mm long, glabrous. Capitula radiate;
involucres campanulate, 25-35 mm high, 8-10
mm wide; phyllaries 5-6-seriate, glabrous, api-
cally lanuginose, the outer ovate, apical appendage
attenuate, more or less recurved, albo-tomentose,
the inner oblong, 28-30 mm long, 5-6 mm wide,
apically obtuse, mucronate; florets dimorphic; ray
florets 9-10, orange, the tube 16 mm long, the
ligule lanceolate, 20-25 mm long, 3-5 mm wide,
inner lip absent; disc florets 20-25, bilabiate, the
tube 1 5-20 mm long, the outer lip tridentate, the
inner lip bifid. Achenes cylindric or fusiform, cos-
tate, glabrous; pappus ca. 25 mm long.
DISTRIBUTION— Known from the puna forma-
tions from central to southern Peru (3500-4300
m).
Mutisia mathewsii is easily recognized by its
linear leaf blades and arachnoid-tomentose phyl-
laries. Cabrera (1966) established the variety
anomala for Peruvian material with the characters
of retrorsely divided stem wings and leaf blades
with distal 1-2-denticulations; however, capitular
characteristics are uniform within the species.
ANCASH." Huaras: Parque Huascaran, Quebrada
Shallap, Smith et al. 10770 (F). Recuay: near Tunel Ca-
huish, between Recuay and Chavin, Ferreyra 14527
(USM). Yungay: Huascaran National Park, Quebrada De-
manda, Smith & Cautivo 10303 (F). AYACUCHO. No
exact locality: Palomino s. n. (USM). LIMA. Yauyos: Hua-
cracocha, Cerrate 1270 (USM). JUNIN. Yauli: Macbride
& Featherstone 907 (us). LIMA. Huarochiri: above Sur-
co, Ferreyra 6338a (USM).
1 1 . Mutisia orbignyana Wedd., Chlor. And. 1 : 22.
1855. LECTOTYPE (designated by Cabrera,
1965): Bolivia, Potosi, St. Bartolo, d'Orbigny
1387 (lectotype, P, not seen; isotype fragment
ex p, F 972437; isotype fragment ex G, F
1023190; photograph ex G, FM neg. 28876).
Shrubs erect, 1-2 m high, branched, branches
glabrous. Leaves simple, sessile; blades linear-lan-
ceolate, 35-70 mm long, 4-8 mm wide, acute,
attenuate at base, glabrous, the margins entire.
Capitulescences of solitary, terminal heads, sub-
sessile. Capitula discoid; involucres turbinate, 25-
30 mm high, 7-10 mm wide; phyllaries 4-5-se-
riate, reddish, glabrous, the outer ovate, 5-10 mm
long, apically linear, the inner oblong-lanceolate,
30-35 mm long, 3.5-4.5 mm wide, mucronate;
florets 5-8, isomorphic, orange, the corolla bila-
biate, the tube 10-12 mm long, the outer lip
4-dentate, apically lanuginose, the inner lip linear.
Achenes cylindric, costate, glabrous, 4-5 mm long;
pappus 17-18 mm long.
60
FIELDIANA: BOTANY
DISTRIBUTION— Known from southern Peru to
Bolivia and northern Argentina (3400-4000 m).
AREQUIPA. Cailloma: Chivay, Lopez Guillen 305
(us); Valle del Colca, Treacy 841 (F). Condesuyos: above
Chuquibamba, Weberbauer 6847 (F, GH, us, USM).
12. Mutisia pulcherrima Muschl., Bot. Jahrb. Syst.
50, Biebl. Ill: 102. 1913. TYPE: Peru, Cha-
chapoyas, inter Tambos Bajazan et Almiran-
te, A. Weberbauer 4444 (holotype, B, presum-
ably destroyed; photograph ex B, FM neg.
15945).
M. macrantha Muschl., Bot. Jahrb. Syst. 50, Biebl.
Ill: 103. 1913. TYPE: Peru, Amazonas, Cha-
chapoyas, Molinopampa, 2400 m, A. Weberbauer
4383 (holotype, B, presumably destroyed; pho-
tograph ex B, FM neg. 15941).
Vines, to 8 m high, branched, branches hanging,
glabrous. Leaves pinnate, sessile; rachis 12-22 cm
long, apex with trifid tendril, leaflets 3-5 pairs,
alternate to subopposite, elliptic-lanceolate, 50-80
mm long, 20-27 mm wide, petiolules 1-2 mm
long, apically acute, mucronate, basally attenuate
to rounded, glabrous. Capitulescences of solitary,
terminal heads, pendulous, the peduncles 16-30
cm long. Capitula radiate; involucres cylindric, 60-
70 mm high, 25-30 mm wide; phyllaries 5-6-se-
riate, the outer ovate, 10-20 mm long, ca. 5 mm
wide, the inner oblong, 55-60 mm long, 10-12
mm wide, apically rounded, mucronate; florets di-
morphic; ray florets 14-15, pistillate, the corollas
orange, ligulate, the ligules lanceolate, 50-60 mm
long, 12-15 mm wide; disc florets bilabiate, the
tube 12-15 mm long, the outer lip tridentate, the
inner lip bifid; anthers 25-30 mm long. Achenes
cylindric, glabrous; pappus 20-25 mm long.
DISTRIBUTION— Native to the intermontane re-
gion of northern Peru (2300-2800 m).
Mutisia pulcherrima is distinguished by its elon-
gate pendent branches and large capitula with con-
spicuous orange florets.
AMAZONAS. Chachapoyas: Leimebamba, Ferreyra
& Acleto 15276 (MO, USM); Calla-Calla, Hutchison &
Wright 5613 (NY, uc, USM); "Peruvia," Mathews 18 (GH);
between Jumbilla and Pomacocha, Tillet 673 (us); Calla-
Calla, Wurdack 1162 (us, USM). CAJAMARCA. Hual-
gayoc: Taulis, Hutchison & Wright 6504 (uc, USM).
1 3. Mutisia rauhii Ferreyra, Bol. Soc. Peruana Bot.
Nos. 1-2, 8: 75. 1980. TYPE: Peru, Cusco,
Anta, E of Cordillera Salcantay, 3600 m, W.
Rauh 1451 (holotype, USM 29417).
Scandent, shrubs, 2-3 m high, branched,
branches ascendent, 3-4 mm in diameter, striate,
glabrous. Leaves simple, sessile; blades linear, 80-
1 60 mm long, 3-6 mm wide, the adaxial surfaces
glabrous, the abaxial surfaces lanuginose, apically
with simple tendril, basally amplexicaul, the mar-
gins entire, revolute. Capitulescences of solitary,
terminal heads, the peduncles 45-70 mm long,
glabrous, bracteate, the bracts 30-45 mm long, 5-
6 mm wide, basally sagittate, glabrous. Capitula
radiate; involucres campanulate, 40-45 mm high,
25-30 mm wide; phyllaries 6-7-seriate, the outer
ovate, 7-16 mm long, subulate, rigid, the inner
elliptic, 40-45 mm long, 14-16 mm wide, obtuse,
mucronate, glabrous, the margins more or less
transparent; ray florets 12-13, the corollas 75-80
mm long, the tube 28-30 mm long, glabrous, the
outer lip 5.2-5.5 mm long, 6-11 mm wide,
6-nerved, glabrous, acute, the inner lip bifid, 2.5-
3 mm long, glabrous; styles 48-50 mm long; disc
florets 38-40, the corollas 48-52 mm long, the
tube 35-37 mm long, 3-4 mm wide, the limb
5-lobed, the outer lip ca. 14 mm long, the inner
lip subequal, ca. 8 mm long; anthers 2.2-2.4 mm
long; style 5.2-5.5 mm long. Achenes cylindric,
3-4 mm long, glabrous; pappus 25-30 mm long,
yellowish.
DISTRIBUTION— Known only from the type lo-
cality.
Mutisia rauhii most closely resembles M. cocha-
bambensis, from which it differs in having larger
leaves and capitula, phyllaries 6-7-seriate and lig-
ules to 55 mm long.
14. Mutisia venusta S. F. Blake, Proc. Biol. Soc.
Wash. 46: 191. 1933. TYPE: Peru, Cusco,
Urubamba, Pinasniocj, Panticalla Pass, O. F.
Cook & G. B. Gilbert 1235 (holotype, us
604430; isotype, GH). Figure 13.
Scandent, vining shrubs, branched, branches
more or less lanuginose. Leaves sessile, pinnate;
rachis 6-15 cm long, trifid at apex, the leaflets
oblong in 6-10 pairs, opposite to subopposite, ses-
sile, glabrous, 20-45 mm long, 8-15 mm wide.
Capitulescences of solitary, terminal heads, the pe-
duncles 5-18 cm long, glabrescent. Capitula ra-
diate; involucres cylindric, 60-90 mm high, 15-
20 mm wide; phyllaries 7-8 -seriate, glabrous, the
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
61
FIG. 13. Mutisia venusta. I, habit; 2, detail of leaflet; 3, achene with pappus. (Drawn from O.F. Cook & G.B.
Gilbert 1235, us.)
62
FIELDIANA: BOTANY
inner oblong-lanceolate, acute, 65-80 mm long,
5-6 mm wide; florets dimorphic; ray florets 7-8,
the tube 6-7 mm long ligulate, ligules lanceolate,
tridentate, 30-40 mm long, 8-10 mm wide; disc
florets 18-20, the corollas bilabiate, the tube 10-
1 2 mm long, the outer lip tridentate, the inner lip
bipartite. Achenes cylindric, glabrous; pappus ca.
24 mm long.
DISTRIBUTION— Known from the Andean Cor-
dillera of southern Peru (3400-4000 m).
Mutisia venusta most closely resembles M. acu-
minata, from which it differs in having longer in-
volucres, lanuginose leaf rachis with 8-9 pairs of
obtuse leaflets, and gradually attenuate phyllaries.
APURIMAC. Abancay: Ampai, without collector,
2416 (USM). CUSCO. Calca: Lares, Mann 2141 (USM).
Urubamba: Pinasniocj, Panticalla Pass, Cook & Gilbert
1879 (us); Penas, Marin 1634 (USM), Vargas 4420 (MO);
Machupicchu, Nunez & Arque 8342 (F).
15. Mutisia wurdackii Cabr., Opera Lilloana 13:
43. 1966. TYPE: Peru, Amazonas, Bongara,
Pomacocha, J. J. Wurdack 840 (holotype, LP;
isotype, us 2373593).
Vines, to 9 m high, branched, branch striate,
hirsute, glabrescent. Leaves sessile, pinnate; rachis
12-22.5 cm long, apex with trifid tendril, twisted,
the leaflets ovate-lanceolate in 6-8 pairs, 5-6 cm
long, 2-4 cm wide, acute, the adaxial surfaces gla-
brous, the abaxial surfaces lanuginose. Capitu-
lescences of solitary, terminal heads, the peduncles
8-15 cm long, lanuginose at apex. Capitula radi-
ate; involucre cylindric, 35-45 mm high, 20-25
mm wide; phyllaries 6-7-seriate, the outer ovate-
deltoid, 6-12 mm long, 6-7 mm wide, apically
attenuate, densely tomentose; the inner oblong,
30-35 mm long, 8-12 mm wide, apically rounded
or truncate, densely tomentose; florets dimorphic;
ray florets 8-12, the corollas red, ligulate, the tube
25-35 mm long, the ligules lanceolate, tridentate,
25-30 mm long, 4-5 mm wide, the inner lip bifid;
disc florets 40-60 mm long, yellow, bilabiate, the
tube 2.8-3.4 mm long, the outer lip tridentate, the
inner lip bifid. Achenes cylindric, 1 5-25 mm long,
1.5-2 mm wide, glabrous; pappus ca. 40 mm long.
Chromosome number: n = ca. 25.
DISTRIBUTION— Native to the intermontane re-
gion of northern Peru (1700-2400 m).
Mutisia wurdackii most closely resembles M.
microcephala Sodiro of Ecuador, but it differs in
having six to eight pairs of leaflets, red corollas,
and a pappus to 40 mm long.
AMAZONAS. Bongara: between Pomacocha and
Jumbilla, Ferreyra 20611 (USM); Pomacocha to Jazan,
Ferreyra & Acleto 15200 (MO, USM), 75255 (USM); be-
tween Utcubamba and Pomacocha, Hutchison & Wright
3881 (F, MO, NY, uc, us, USM); without locality, Hutchison
4886 (USM). Chachapoyas: Leimebamba, Boeke 1784 (MO,
NY); 16 km SW of Pomacocha, Gentry et al. 61354 (F);
Cerros Calla Calla, Hutchison & Wright 5613 (F); entre
Calla-Calla y Leimebamba, Sanchez 971 (F).
XV. ONOSERIS
Onoseris Willd., Sp. PI. 3: 1702. 1804. TYPE:
Atractylis purpureaL. fil. = Onoseris purpurea
(L. fil.) S. F. Blake2
Cursonia Nutt, Trans. Amer. Philos. Soc. 7: 422. 1 84 1 .
TYPE: Cursonia peruviana Nutt. = Onoseris
odorata (D. Don) Hook. & Arn.
Shrubs, lianas, or perennial or annual herbs;
stems lacking spines. Leaves alternate or entirely
basal; sessile or petiolate; blades linear to cordi-
form, usually lanceolate in outline, hastate, sag-
ittate, or lyrate, usually discolorous, the adaxial
surfaces glabrescent, the abaxial surfaces arach-
noid to lanate, the margins crenate to denticulate.
Capitulescences of solitary, terminal heads or pa-
niculate in groups of 2-8 (-3 00). Capitula usually
heterogamous, radiate, or occasionally homoga-
mous, discoid; involucres campanulate, turbinate,
hemispheric; receptacles plane to convex, fim-
brillate, pubescent to glabrous; phyllaries 4-12-
seriate, imbricate; florets isomorphic or hetero-
morphic; ray florets pistillate, the corollas bilabi-
ate, violet or purple, the outer lip ligulate, 6-nerved,
3 -dentate, the inner lip bifid or entire; disc florets
hermaphroditic, tubular, yellow, the limb cylin-
dric, 5-lobed, the lobes equal or unequal, less than
'/3 the length of the corolla, erect to recurved.
Achenes cylindric to subterete, (4-)5(-6)-costate,
glabrous to pubescent; pappus of barbellate bris-
tles, 2-many-seriate, isomorphic or rarely with in-
ner bristles longer, yellowish.
Onoseris contains approximately 30 species dis-
tributed from Mexico and Central America south
throughout the Andean Cordillera from Colombia
and Venezuela to northern Argentina. In Peru, no
fewer than 1 8 species are currently recognized.
For additional synonymy, consult Ferreyra (1944).
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
63
References . 1959. Dos especies nuevas de Onoseris
(Compositae) para la Rora Peruana. Publ. Mus.
FERREYRA, R. 1944. Revision del genero Ono- Hist. Nat. "Javier Prado," Ser. B, Bot., 11: 1-6.
seris. J. Arnold Arb., 25: 349-395. . 1980. Una especies nueva de Onoseris
. 1949. Una nueva especie de Onoseris (Compositae) del Peru. Bol. Soc. Arg. Bot., 19:
(Compositae) procedente del Peru. Publ. Mus. 19-23.
Hist. Nat. "Javier Prado," Ser. B, Bot., 1: 1-4.
Key to Peruvian Species of Onoseris
1 . Leaf blades orbicular or cordiform, palmate- veined from base 2
2. Involucres hemispherical, ca. 30 mm high, ca. 30 mm wide; phyllaries densely ferrugino-glan-
dulose, the inner ca. 25 mm long, strongly flexuose O. acerifolia
2. Involucres campanulate, ca. 20 mm high, ca. 15 mm wide; phyllaries densely albo-lanuginose,
sparsely glandulose, the inner ca. 1 5 mm long, erect O. cabrerae
1 . Leaf blades linear, lanceolate, or oblanceolate to spathulate or lyrate, pinnate- veined 3
3. Plants annual herbs 4
4. Leaf bases auriculate, amplexicaul O. amplexicaulis
4. Leaf bases acute to attenuate or cuneate 5
5. Plants to 50 cm tall; leaf blades typically > 5 cm long, > 2 cm wide; inner phyllaries
filiform, long-attenuate, strongly reflexed or twisted O. odorata
5. Plants to 20 cm tall; leaf blades typically < 5 cm long, < 2 cm wide; inner phyllaries
acuminate, erect 6
6. Involucres narrowly campanulate, 4-6 mm wide; ray florets 7-9; disc florets 8-10 . . .
O. minima
6. Involucres campanulate, (5-)6-9 mm wide; ray florets 8-13; disc florets 20 or more
7
7. Stems erect; leaf blades entire to subentire; involucres ca. 1 2 mm high; inner phyllaries
0.7-0.8 mm wide O. cumingii
7. Stems prostrate to ascending; leaf blades dentate; involucres ca. 8 mm high; inner
phyllaries 1 .5-2 mm wide O. annua
3. Plants perennial herbs 8
8. Leaf blades linear to linear-lanceolate or narrowly spathulate, typically 1.5-10 mm wide ... 9
9. Phyllaries 9-1 1 -seriate, the inner linear, ca. 1 mm wide, apically long-attenuate; ray florets
20-24 O. lopezii
9. Phyllaries 4-7 -seriate, the inner lanceolate to oblong, 1.5-2.5 mm wide, apically acute to
acuminate; ray florets 5-15 10
10. Leaf blades linear, the margins strictly entire; phyllaries 4-5-seriate, the inner oblong,
ca. 2.5 mm wide, apically rounded to obtuse; ray florets 9-1 1 O. linearifolia
10. Leaf blades narrowly lanceolate to elliptic, the margins remotely denticulate to sinuose-
dentate; phyllaries 6-7-seriate, the inner lanceolate, 1.2-1.5 mm wide, apically acute
to acuminate; ray florets 5-8 or 1 5 11
1 1 . Phyllaries nigrescent, ferrugino-tomentose; ray florets ca. 15 . 0. chrysactinioides
1 1 . Phyllaries yellowish-green, albo-tomentose; ray florets 5-8 . . . O. gnaphalioides
8. Leaf blades lanceolate, oblanceolate, spathulate, or lyrate, typically 10-30(-120) mm wide
12
1 2. Plants caulescent; leaves cauline, the blades narrowly lanceolate or hastate 13
13. Leaf blades hastate to sagittate; phyllaries 10-12-seriate, the inner long-attenuate,
yellowish-green O. castelnaeana
13. Leaf blades lanceolate; phyllaries 5-6-seriate, the inner acuminate, nigrescent, stra-
mineous-tomentose O. albicans
12. Plants acaulescent; leaves forming a basal rosette, the blades lanceolate, oblanceolate,
spathulate, or lyrate 14
64 FIELDIANA: BOTANY
14. Capitulescences paniculate with 7-10 heads; peduncles 40-60 cm long; leaves (in-
cluding petioles) 30-40 cm long, the distal portion hastate, apically attenuate
O. peruviana
1 4. Capitulescences of solitary heads or rarely of 2-3 heads on a single peduncle; peduncles
10-40 cm long; leaves (including petioles) 5-30 cm long, the distal portion sagittate
to cordate or rhomboid to spathulate, apically subacute to obtuse or rounded ..15
15. Plants procumbent; leaves with distal portion rhomboid to spathulate; Capitu-
lescences procumbent or laxly ascending (southern Peru, coastal habitats below
500 m) O. humboldtiana
15. Plants erect; leaves with distal portion sagittate to cordate; Capitulescences erect
(northern Peru, montane habits above 1000 m) 16
16. Leaves lanceolate; sessile; outer phyllaries ovate, erect, the inner lanceolate,
ca. 1.5 mm wide, apically acuminate, erect; achenes densely pilose
O. \veberbaueri
16. Leaves oblanceolate or lyrate; petiolate; outer phyllaries apiculate, strongly
recurved, the inner long-attenuate, ca. 1 mm wide, apically twisted; achenes
sparsely puberulent to glabrous 17
17. Leaf blades lyrate; petioles 4-15 cm long, winged and lobed; achenes
glabrous O. speciosa
1 7. Leaf blades oblanceolate; petioles 3—4 cm long, winged; achenes sparsely
puberulent O. macbridei
1. Onoseris acerifolia Kunth, Nov. Gen. Sp. 4: 8.
1820. TYPE: Peru, "in prov. Bracamorensis
in devexis Parami de Yamoca inter pagos Co-
lazey et Chontali," Humboldt & Bonplands.n.
(holotype, p, not seen; photograph ex p, FM
neg. 38101).
Suffruticose, perennial herbs, 50-80 cm high,
erect, stems terete, densely glandular-pubescent.
Leaves alternate; petioles to 75 mm long, glan-
dular-pubescent; blades orbicular-cordiform, 10-
90 mm long, 11-120 mm wide, the adaxial sur-
faces arachnoid and glandular-pubescent, the ab-
axial surfaces white-lanuginose and glandular-pu-
bescent, the margins lobed. Capitulescences of ter-
minal, solitary heads; peduncles 2-8 cm long, glan-
dular, bracteate, the bracts 5-10 mm long, 1.5-2
mm wide, glandular. Capitula radiate; involucres
7-25 mm high, 5-20 mm wide, hemispheric; phyl-
laries 6-7-seriate, the inner 22-29 mm long, 1.5-
3 mm wide, attenuate at apex, the abaxial surfaces
arachnoid and glandular; ray florets 13-21, the
corolla 33-44 mm long, the tube 6-9 mm long,
glandular, the outer lip 27-35 mm long, viola-
ceous, the abaxial surfaces arachnoid, glandular,
tridentate, the inner lip 5-17 mm long, attenuate,
twisted; disc florets with corollas 14-16.5 mm long,
the tube 1 2.5-1 4 mm long, 5-lobed, the lobes acute,
slightly coiled; anthers 5.6-7 mm long. Achenes
[ray] 2.4-4.5 mm long, 1 mm wide, pubescent;
pappus ca. 12 mm long, yellow; [disc] 2.5-5 mm
long, 1-1.5 mm wide, pubescent; pappus ca. 13
mm long, isomorphic, yellow.
DISTRIBUTION— Endemic to the intermontane
region of northern Peru (380-1800 m).
Onoseris acerifolia is characterized by its arach-
noid-tomentose pubescence mixed with glandular
trichomes, orbicular-cordate leaves, large capitu-
la, and elongate, attenuate, flexuose, densely glan-
dular phyllaries. It most closely resembles O. ca-
brerae; however, that species has smaller leaves
and capitula and more rigid phyllaries.
AMAZON AS. Bagua: km 248, between Pongo de
Rentema and Amojao, Ferreyra 13676 (MO, USM); Pongo
de Rentema, Ferreyra 19440 (USM); Bagua, Rauh 40314
(USM); 22 km N of Bagua Chica on road to Chiriaco,
Gentry et al. 23108 (F). Luya: arriba de Balsas, Sanchez
348 (F). CAJAMARCA. Jaen: cerca de la carretera a
Jaen, Angulo 2096 (F, NY); between Porculla and Jaen,
Ferreyra 13636 (USM); Porculla-Pucara, Ferreyra 14417
(MO, USM); road to Las Pirias, Ferreyra & Sanchez 19569
(USM); above Shumbe Alto, Ferreyra & Sanchez 19611
(USM); 6 km E of Pucara, Gentry et al. 22704 (F); Rio
Huancabamba, 90 km E of Olmos, Hutchison 1395 (F,
NY, USM); between Acapulco and Pucara, Hutchison &
Wright 3514 (F, uc, USM); Pucara to Chamaya, Plowman
et al. 14249 (F); km 81, Chamaya, Velarde 5880 (USM);
Jaen, Weberbauer 6203 (F). San Ignacio: Chuchuhuasi-
Huaquilla, Lopez et al. 8970 (F). LA LIBERT AD. Pataz:
Chagual-Pataz, Alayo 74 (F).
2. Onoseris albicans (D. Don) Ferreyra, J. Arnold
Arb. 25: 389. 1994.
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
65
Centroclinium albicans D. Don, Trans. Linn. Soc.
London 16: 256. 1829. TYPE: Peru, Ruiz Lopez
& Pavon s.n. (holotype, p, not seen).
Onoseris integrifolia Less., Linnaea 5: 343. 1830.
TYPE: Peru, /. Dombey 25 (holotype, B, presum-
ably destroyed; isotypes, MA, P, not seen; isotype
ex P, F 972679; photograph ex p, FM neg. 8266).
Perennial herbs, 30-80 cm high, white-lanugi-
nose. Leaves alternate, subsessile to petiolate; pet-
ioles 5-10 mm long; blades lanceolate, 20-95 mm
long, 6-35 mm wide, the adaxial surfaces arach-
noid, the abaxial surfaces lanuginose, apically and
basally acuminate to acute, the margins sinuose-
dentate, revolute. Capitulescences of solitary, ter-
minal capitula; peduncle 70-245 mm long, white-
lanuginose, bracts subulate, 4-9 mm long. Capit-
ula radiate; involucres 12-20 mm high, 7-13 mm
wide, campanulate; phyllaries 5-6-seriate, the in-
ner 10-12 mm long, 2-2.4 mm wide, acuminate;
ray florets ( 1 3-)2 1-34, the corollas violaceous, 26-
40 mm long, the tube 5.5-7 mm long, ca. 0.8 mm
wide, the outer lip 20-33 mm long, 3.6-4.8 mm
wide, 6-nerved, the abaxial surfaces arachnoid, tri-
dentate, the inner lip bipartite, 2.2-3.5 mm long,
ca. 0.9 mm wide; disc florets with corollas, 12-16
mm long, the tube 10-13 mm long, the lobes yel-
low; anthers 5-7.5 mm long. Achenes [ray] 2.5-5
mm long, 0.8-1 mm wide, glabrescent; pappus ca.
13 mm long, heteromorphic, yellow; [disc] 2.5-
6.5 mm long, 1-1.3 mm wide, glabrescent; pappus
ca. 14 mm long, heteromorphic, yellow.
DISTRIBUTION— Known from the Andean Cor-
dillera of northern Peru to central Bolivia ( 1 400-
3200 m).
Onoseris albicans closely resembles O. salici-
folia, an Ecuadorian species, but differs in pos-
sessing leaves with white-arachnoid pubescence,
revolute margins, campanulate involucres, and
glabrescent achenes.
APURIMAC. Abancay: near Abancay, Ferreyra 9818
(USM). Grau: Kairancka, Vargas 5816 (MO, USM). CA-
JAMARCA. Cajamarca: below San Pablo, Hutchison &
Wright 5072 (F, MO, NY, USM); Vista Alegre, Sagdstegui
et al. 10142 (F); Asuncion-San Juan, Sagdstegui et al.
10150 (F), 11994 (F); Yumagual, Sanchez 233 (F); Ha-
cienda Huacaruco, Ute s.n. (MO, USM). Chota: Llama,
Ferreyra 8402 (MO, USM); between Cochabamba and La-
jas, Ferreyra 8470 (MO, USM); Lajas, Mostacero et al.
1533 (F). Contumaza: Tambo La Lima, Lopez et al. 9025
(F); Contumaza, Sagdstegui et al 9842 (F). Cutervo: near
Cutervo, Ferreyra & Acleto 15390 (USM). Hualgayoc: San
Miguel, s.d. (USM). CUSCO. Anta: Challatambo, Cuzco
to Abancay, Balls 6835 (F). Cusco: Oropeza, Beck 8368
(F). Paruro: near Paruro, Vargas 132 (F, GH); Hacienda
Ayusbamba, Vargas 981 (MO). Urubamba: Chinchero to
Urquillos, King et al. 301 (F); Limatambo, Rauh & Hirsch
P1352 (F, NY); Urquillos, Vargas 11097 (F). No exact
locality: Herrera 2586 (F). HUANCAVELICA. Huan-
cavelica: 5 km E of Mejorada, Stork & Norton 10903
(F). HUANUCO. Ambo: near Ambo, Ferreyra 6627
(USM). Huanuco: Mitotambo, Ferreyra 6695 (F, MO, USM);
Acomayo, Ridoutt s.n. (USM); Dunkafael, Sawada P77
(F); Acomayo, Woytkowski 34277 (F, MO). Pachitea: near
Panao, Ferreyra 1806 (GH, MO, USM). JUNIN. Huan-
cayo: near Huancayo, Hjerting & Petersen 1386 (USM).
Yauli: near Oroya, Kalenborn 71 (MO). LA LIBERTAD.
Otusco: 65 km E of Trujillo on road to Otusco, Dillon
& Turner 1495 (F); near Otusco, Ferreyra 2970 (MO,
USM); Agallpampa-Samne, Ferreyra 3084 (MO, USM);
Samne-Otusco, Ferreyra 7615 (MO, USM); Sinsicap, Lopez
1078 (USM); Casmiche, Sagdstegui et al. 11685 (F).
LAMBAYEQUE. Lambayeque: Olmos-Porculla, Fer-
reyra 13619 (USM), 75767 (MO, USM); Abra de Porculla,
Plowman et al. 14268 (F). LIMA. Canta: near Canta,
Soukup 2814 (GH). Huarochiri: 3 km W of San Mateo,
Dillon 2501 (F); between Lima and San Mateo, Duncan
et al. 2707 (F); near Matucana, Ferreyra 718 (USM), 2003
(USM), 7695 (F), 8293 (MO, USM), 76596 (MO, USM); near
Surco, Ferreyra 14870 (MO, USM); San Mateo, Ferreyra
5269 (MO, USM), 5572 (MO, USM), 7695 (MO, USM); Ob-
rajillo, Ferreyra 18336 (MO, USM); Viso, Goodspeedet al.
11524 (F, GH); km 90 Lima-Oroya, Goodspeed & We-
berbauer 33062 (GH, MO), Macbride & Featherstone 240
(F); Rose 18641 (F, NY), Ugent 5295 (USM), Hutchison
690 (USM), Soukup 2076 (GH, MO, USM); Puruchuco, Ma-
thews s.n. (GH); Collca, Meza 153 (USM); below Obrajillo,
Pennell 14496 (F, GH); Chosica-Matucana, Vargas 4781
(USM). PASCO. Daniel Carrion: 13 km NE of Yana-
huanca, Bird 1265 (F). PIURA. Huancabamba: Cerro
Porculla, Edwin & Schunke 3740 (F, USM); Abra de Por-
culla, Gentry et al. 22670 (F). WITHOUT EXACT LO-
CALITY. Ochoa 1559 (USM), 365 (USM), Rauh 1352
(USM).
3. Onoseris amplexicaulis Ferreyra, J. Ajnold Arb.
25: 364. 1944. TYPE: Peru, Lima, Huaro-
chiri, Matucana, J. F. Macbride & W. Feath-
erstone 310 (holotype, F 5 16844; isotype, GH).
Annual herbs, 11-25 cm high, erect, lanuginose.
Leaves sessile; blades oblanceolate to lanceolate,
14-75 mm long, 3-20 mm wide, the adaxial sur-
faces arachnoid, the abaxial surfaces lanuginose,
apically acuminate to acute, basally auriculate,
amplexicaul, the margins sinuose-dentate, revo-
lute. Capitulescences of solitary, terminal heads;
peduncles 1 5-48 mm long, lanuginose, bracteate,
the bracts 3-5 mm long, subulate, the abaxial sur-
faces arachnoid. Capitula radiate; involucres 7-1 1
mm high, 5-6 mm wide, campanulate; phyllaries
4-5-seriate, the inner 1 1-12 mm long, 2-2.2 mm
wide; ray florets 8-13, the corollas 1 8-2 1 mm long,
the tube 3-3.5 mm long, the outer lobe 15-18 mm
long, 3.5-4 mm wide, 6-nerved, the abaxial sur-
faces arachnoid, tridentate, the inner lobe bipar-
tite, 2-2.5 mm long, filiform; disc florets with co-
66
FIELDIANA: BOTANY
rollas 7.8-8.5 mm long, the tube 6.6-7 mm long,
the lobes acute; anthers 3-3.5 mm long. Achenes
[ray] 2-4 mm long, 1 mm wide, pubescent; pappus
ca. 6 mm long, isomorphic, yellow; [disc] 2.5-4.2
mm long, 1 mm wide, pubescent; pappus ca. 8
mm long, isomorphic, yellow.
DISTRIBUTION— Endemic to Andean Cordillera
of central Peru (1800-2400 m). Onoseris amplex-
icaulis is very similar to O. annua, but it differs
in possessing amplexicaul leaf bases.
LIMA. Huarochiri: near Matucana, Ferreyra 1375
(USM), 7600 (F, MO, USM); Huariquina, Ferreyra 9172
(USM), 9755 (MO, USM); near Surco, Ferreyra 656 (MO,
USM), 2025 (USM), 3420 (MO, USM), 16894 (MO, USM);
Matucana, Macbride & Featherstone 131 (F); near Ma-
tucana, Rose 18668 (NY).
4. Onoseris annua Less., Linnaea 5: 341. 1830.
TYPE: Peru, "Chinchin" [Chiuchin], J. Dom-
bey s.n. (holotype, B, presumably destroyed;
isotype, F ex p; photograph ex B, FM neg.
15887).
Annual herbs, 4-16 cm high, erect, lanuginose.
Leaves alternate, petiolate; petioles 1.5-2 mm long,
lanuginose; blades lanceolate, 10-25 mm long, 5-
10 mm wide, bases attenuate, the adaxial surfaces
arachnoid, the abaxial surfaces lanuginose, the
margins dentate. Capitulescences of solitary, ter-
minal heads; peduncles 1-7 cm long, lanuginose.
Capitula radiate; involucres 9-10 mm high, 5-7
mm wide, campanulate; phyllaries 5-6-seriate, the
inner 9-10 mm long, 1.4-2 mm wide, acute, the
abaxial surfaces arachnoid; ray florets ca. 13, the
corollas 18-20 mm long, the tube 3.5-4 mm long,
the outer lip 10.5-16 mm long, 2.2-2.5 mm wide,
6-nerved, tridentate, the inner lip bipartite, 2-2.3
mm long, filiform. Achenes [ray] 2-2.2 mm long,
pubescent; pappus ca. 5 mm long, heteromorphic,
yellow; disc florets yellow, the corollas 7-7.5 mm
long, the tube 5.6-6 mm long, the lobes acute;
[disc] 2.2-3.5 mm long, pubescent; pappus ca. 6
mm long, heteromorphic, yellow.
DISTRIBUTION— Known from the Andean Cor-
dillera of northern and central Peru (2200-3400
m).
Onoseris annua most closely resembles O. am-
plexicaulis but lacks auriculate leaf bases.
ANCASH. Bolognesi: near Chiquian, Pennell 15408
(MO, USM), Ferreyra 5670 (F, MO, USM), 7376 (MO, USM).
Huaylas: between Caras and Huallanca, Canon del Pato,
Ferreyra & Rojas 14578 (USM); "Incapamanam," Car-
rillo 647 (USM). LIMA. Cajatambo: near Churin, Fer-
reyra 3518 (USM), 3520 (USM), 3548 (MO, USM). Chancay:
Chiuchin, Ferreyra 19164 (USM). Huarochiri: Monte
Zarate, above San Bartolome, Ferreyra 97 11 A (USM).
5. Onoseris cabrerae Ferreyra, Bol. Soc. Argent.
Bot. 19: 19. 1980. TYPE: Peru, Amazonas,
Chachapoyas, encima de Balsas, hacia Cor-
dillera Calla-Calla, 1300-1400 m, 21 Aug
1963, R. Ferreyra & C. Acleto 15339 (holo-
type, USM 15746).
Perennial herbs, suffrutescent, 5-10 dm high,
erect, stem pubescent, glandulose. Leaves alter-
nate, petiolate; petioles 40-80 mm long; blades
cordiform, 50-110 mm long, 60-130 mm wide,
palmatilobed with 8-10 segments, acute, mucro-
nate, the margins dentate. Capitulescences ter-
minal cymes, 2-4-headed; peduncles hirsute-glan-
dular, bracteate, bracts 3-7 mm long, 1-1.5 mm
wide. Capitula radiate; involucres campanulate,
18-20 mm high, 12-16 mm wide; phyllaries 5-6-
seriate, lanceolate, the inner 16-17 mm long, 1-
1.2 mm wide, attenuate at apex, the abaxial sur-
faces lanuginose, hirsute-glandular; ray florets 1 7-
18, the corollas purple, 30-32 mm long, the tube
5-6 mm long, hirsute-glandular, the outer lip 26-
32 mm long, 2.5-3 mm wide, 6-nerved, the ab-
axial surfaces lanuginose, hirsute-glandular, tri-
dentate, the 3 lobes acute, coiled, the inner lip 3-
6 mm long, filiform, twisted; disc florets 36-43,
the corollas yellow, 12-13 mm long, the tube 1 1 .5-
12.5 mm long, the lobes acute, coiled; anthers 8.5-
9 mm long. Achenes [ray] cylindric, ca. 3 mm long,
pubescent; pappus ca. 10 mm long, isomorphic,
yellowish; [disc] cylindric, 3-5 mm long, 1.2 mm
wide, hirsute; pappus ca. 1 1 mm long, isomorphic,
yellowish.
DISTRIBUTION— Known from the intermontane
region of north-central to southern Peru (1200-
2600 m).
Onoseris cabrerae most closely resembles O. ac-
erifolia, but, unlike that species, it possesses a more
sparse hirsute-glandular pubescence, peduncles
with bracts conspicuously rigid, and campanulate
involucres with ascending phyllaries.
AMAZONAS. Chachapoyas: above Balsas, Hutchi-
son & Wright 5448 (uc, us, USM). APURIMAC. An-
dahuaylas: Pincos, Stork & Horton 10655 (F, GH, MO,
uc, USM). CUSCO. Anta: between Limatambo and Apu-
rimac, Rauh 1528 (USM); Cuesta Mandor, Vargas 5874
(USM). HUANCAVELICA. Tayacaja: above Quinta-
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
67
bamba, Tovar 4587 (USM). LA LIBERTAD. Pataz: 5 km
above Chagual, Hutchison et al. 6233 (F, uc, us, USM).
6. Onoseris castelnaeana Wedd., Chlor. And. 1:
10. 1855. TYPE: Peru, [Cusco], Jun 1847, M.
F. Castelnau 35 (holotype, P, not seen; isotype
fragment ex P, F; photograph ex P, FM neg.
38105).
Seris castelnaeana (Wedd.) Kuntze, Revis. gen. pi. 1 :
364. 1891.
Perennial herbs, decumbent, lanuginose. Leaves
alternate; petioles 8-30 mm long; blades sagittate
to hastate, 26-60 mm long, 7-29 mm wide, the
adaxial surfaces arachnoid, the abaxial surfaces
lanuginose, apically acute, the margins slightly
dentate. Capitulescences weakly cymose, 2-3-
headed; peduncles 5-16 cm long, lanuginose, brac-
teate, the bracts 4-6 mm long, subulate, arach-
noid-tomentose. Capitula radiate; involucres 1 8-
28 mm high, 8-12 mm wide, campanulate; phyl-
laries 10-12-seriate, the inner 22-24 mm long,
1 .8-2 mm wide, acuminate, arachnoid-tomentose;
ray florets 13-21, the corollas violaceous, 46-48
mm long, the tube 11-12 mm long, the outer lip
35-36 mm long, 3-3.5 mm wide, 6-nerved, the
abaxial surfaces arachnoid, tridentate, the inner
lip bipartite, 6-7 mm long, glabrous, attenuate,
filiform; disc florets yellow, the corollas 1 8-22 mm
long, the tube 15-19 mm long, the lobes acute, the
abaxial surfaces pubescent; anthers 7-8 mm long.
Achenes [ray] 4-4.2 mm long, 1-1.2 mm wide,
pubescent; pappus ca. 18 mm long, isomorphic,
yellow; [disc] 3-3.5 mm long, 1-1.2 mm wide,
pubescent; pappus ca. 19 mm long, isomorphic,
yellow.
DISTRIBUTION— Known from the intermontane
region of southern Peru (2400-2900 m).
Onoseris castelnaeana is characterized by its
sagittate to hastate leaves, 10-12-seriate involu-
cre, and phyllaries with apices conspicuously re-
flexed.
APURIMAC. Abancay: near Abancay, Vargas 404
(GH). Aimaracs: 1 5 km NW of Chalhuanca, below Pakay-
ca, Gentry et al. 23341 (F). CUSCO. Anta: Limatambo,
Rauh & Hirsch 1527 (NY, USM). Paruro: Chaupimoco,
Vargas 403 (GH).
7. Onoseria chrysactinioides Sagast. & Dillon,
Brittonia 37: 8. 1985. TYPE: Peru, Cajamar-
ca, Contumaza, Lleden, Sagdstegui et al.
10839 (holotype, HUT; isotypes, F, HUT, MO).
Perennial herbs to 4 dm tall, branched; stems
ascending, lanulose. Leaves clustered at apices of
branchlets; petioles 2-12 mm long; blade linear,
30-65 mm long, 2-7 mm wide, the abaxial sur-
faces densely albo-tomentose, the adaxial surfaces
lanate, grey, basally acuminate, apically acute, the
margins shallowly repand, remotely denticulate,
revolute. Capitulescences of solitary heads, sea-
pose, the peduncles to 20 cm long, minutely brac-
teolate distally. Capitula 12-15 mm high, ca. 12
mm wide; involucres campanulate; receptacle with
prominent laciniate crests, planar; phyllaries 6-7-
seriate, linear-lanceolate, the outer (2-)4-6 mm
long, 0.5-1.5 mm wide, the abaxial surfaces lanate,
apically attenuate, the inner (10-)13-17(-20) mm
long, l-2(-3) mm wide, the abaxial surfaces fer-
rugino-tomentose, the margins scarious, apically
acuminate; ray florets ca. 15, 1 -seriate, pistillate,
the corollas lilac, bilabiate, the outer lip ligulate,
20-25 mm long, 4-5 mm wide, 6-nerved, apically
tridentate, the abaxial surfaces lanate, the inner
lip reduced to 2 linear lobes, 3-5 mm long, apically
clavate, glabrous; disc florets 25-40, the corollas
lilac, tubular, 5-6 mm long, glabrous, the 5 lobes
1.5-2.5 mm long. Achenes cylindric, (3)4-5(6) mm
long, ribbed, glabrous or sparsely pilose apically;
pappus 8-12 mm long, isomorphic, stramineous.
DISTRIBUTION— Known only from the type lo-
cality.
Onoseris chrysactinioides resembles both O. li-
nearifolia and O. lopezii but is readily distin-
guished from both by its petiolate, shallowly re-
pand, dentate leaves clustered at the branch apices,
and ferrugino-tomentose inner phyllaries.
CAJAMARCA. Contumaza: arriba de Lleden, Sa-
gdstegui et al. 14018 (F); San Martin, Sagdstegui et al.
14025 (F).
8. Onoseris cumingii Hook. & Arn., Companion
Bot.Mag. 1: 103. 1835. TYPE: "North Peru,"
without exact locality, H. Cuming 995 (ho-
lotype, K; photographs ex P, FM neg. 38106).
Annual herbs, erect, 12-30 mm high, lanugi-
nose. Leaves sessile to petiolate; petioles 2-4 mm
long, lanuginose; blades oblanceolate, 8-65 mm
long, 3-20 mm wide, the adaxial surfaces arach-
noid, the abaxial surfaces lanuginose, acute, the
margins entire to remotely denticulate, revolute.
Capitulescences of solitary, terminal heads; pe-
duncles 22-120 mm long, lanuginose, bracteate,
bracts 2.5-5 mm long, subulate, the abaxial sur-
68
FIELDIANA: BOTANY
faces arachnoid. Capitula radiate; involucres 8-12
mm high, 7-9 mm wide, campanulate; phyllaries
5-6-seriate, the inner 8-10 mm long, 0.8-1.4 mm
wide, acuminate, the abaxial surfaces pubescent
arachnoid-tomentose; ray florets 8-13, the corol-
las 15-25 mm long, the tube 2.5-4 mm long, the
outer lip 12.5-21 mm long, 2.5-3.5 mm wide,
6 -nerved, the abaxial surfaces arachnoid, triden-
tate, the inner lip bipartite, 0.5-3 mm long, fili-
form; disc florets with corollas 8-1 1 mm long, the
tube 6-8.5 mm long, geniculate, the lobes acute;
anthers 3.8-4.5 mm long. Achenes [ray] 1 .5-3 mm
long, glabrescent; pappus ca. 5 mm long, hetero-
morphic, yellow; [disc] ca. 3.5 mm long, ca. 1 mm
wide, glabrescent; pappus ca. 6 mm long, heter-
omorphic, yellow.
DISTRIBUTION— Endemic to coast region of
northern Peru.
Onoseris cumingii very closely resembles O.
odorata but differs in possessing oblanceolate leaves
and tomentose, acuminate phyllaries that lack pi-
lose or hirsute pubescence common to the invo-
lucre of the latter species.
PIURA. Talara: Cerro Pan de Azucar, Naught 30 (F,
NY, US).
9. Onoseris gnaphalioides Muschl., Bot. Jahrb.
Syst. 50, Beibl. Ill: 94. 1913. TYPE: Peru,
Ancash, Caraz, A. Weberbauer 3012 (holo-
type, B, presumably destroyed; photograph ex
B, FM neg. 15891; isotype, USM 33393; isotype
fragment ex B, F 1013035).
O. hyssopifolia Kunth & teretifolia Wedd., Chlor. And.
1: 10. 1855. SYNTYPES: Peru, "Cuzco," Gay
s.n. (P, not seen); Bolivia, "d'Ayopaya," Weddell
4140 (P, not seen).
Perennial herbs, 1 1-70 cm high, decumbent,
lanuginose. Leaves subsessile; blades lanceolate,
12-55 mm long, 3-8 mm wide, the adaxial sur-
faces arachnoid, the abaxial surfaces tomentose,
apically acuminate to acute, the margins revolute,
sinuose-dentate. Capitulescences of solitary, ter-
minal heads; peduncle 22-165 mm long, bracts 2-
7 mm long, subulate. Capitula radiate; involucres
8-12 mm high, 4-7 mm wide, campanulate; phyl-
laries 6-7-seriate, the inner 1 1-12 mm long, 1.8-
2.2 mm wide, acute, the abaxial surfaces arach-
noid; ray florets 5-8, the corollas 17-19 mm long,
violaceous, the tube 5-6 mm long, the outer lip
12-13 mm long, 2.8-3.4 mm wide, 6-nerved, the
abaxial surfaces arachnoid, tridentate, the inner
lip bipartite, the lobes filiform, attenuate; disc flo-
rets ca. 15, the corollas 7.5-10 mm long, the tube
6.5-8 mm long the lobes acute; anthers 3-3.5 mm
long. Achenes [ray] 2.5-5 mm long, 1 mm wide,
glabrescent; pappus ca. 7 mm long, isomorphic,
yellow; [disc] 2-4 mm long, 1 mm wide, glabres-
cent; pappus ca. 8 mm long, isomorphic, yellow.
DISTRIBUTION— Known from the Andean Cor-
dillera of northern Peru to Bolivia (1 700-3200 m).
Onoseris gnaphalioides most closely resembles
O. hyssopifolia Kunth, a Colombian and Ecua-
dorian species, but differs in having more dentate
leaves and more conspicuous arachnoid pubes-
cence on the adaxial surfaces.
AMAZONAS. Bagua: between Bagua and Bellavista,
Diaz s.n. (USM); near Bagua Grande, Hutchison 1495 (F,
NY, USM). Chachapoyas: Cerro Calla-Calla, Edwin &
Schunke 3621 (F); above Balsas, Pennell 15182 (USM).
ANCASH. Huaylas: Caraz, Ute s.n. (USM). Yungay:
Mancos, Cordillera Negra, Smith & Bias 4869 (USM).
APURIMAC. Abancay: near Abancay, Hutchison 1749
(USM); Curahuasi, Marin 2066 (F); Puente Cunyac, Fer-
reyra 2747 (MO, USM), 9835 (MO, USM); Saunders 746 (F,
NY), Chavez 3068 (MO); Rio Pachachaca, 10 km SW of
Abancay, Iltis & Ugent 796 (F, NY); 20 km N of Abancay,
Stork et al. 10542 (F, GH, MO); Cconoc, Chavez 3246
(MO); Abancay, Vargas 405 (F). Grau: Kairancka, Vargas
5860 (USM). AYACUCHO. Andahuaylas: Pinkos, Her-
rera 1491 (GH). Cangallo: Ocros-Andahuaylas, Hjerting
& Petersen 1415 (USM). Huanta: near Huanta, Hjerting
& Petersen 1394 (USM). CAJAMARCA. Cajamarca: near
Cajamarca, Coronado s.n. (USM); Cumbre El Gavilan,
Ferreyra 8596 (USM); Chaquil, Ferreyra 13233 (USM);
near Cajamarca, Pennell 15071 (USM); Fundo Colpa,
Sanchez 324 (USM). Celendin: 10 km SW of Balsas, Dil-
lon & Turner 1701 (F). Jaen: between Porculla and Jaen,
Ferreyra 13639 (MO, USM); Pucara, Ferreyra 14416 (MO,
USM), 75650 (USM); 6 km E of Pucara, Gentry et al. 22713
(F); 90 km E of Olmos, Hutchison 1394 (GH, uc, USM);
15 km E of Acapulco, between Olmos and Jaen, Hutch-
ison & Wright 3512 (F, GH, K, MO, us, USM); 1 km W of
Pucara, Hutchison & Wright 3608 (F, MO, NY, us, USM);
97 km E of Olmos, Plowman et al. 14228 (F); 1 km W
of Chamaya, Stein & Todzia 2227 (F). CUSCO. Anta:
Sisal, Limatambo, Vargas 3693 (MO). HUANCAVELI-
CA. Tayacaja: between Quichicapota and Mantaro, Stork
& Horton 10399 (F); Surcubamba, Tovar 3709 (USM);
Valle del Mantaro, bajo Colcabamba, Weberbauer 6470
(F, GH). LA LIBERTAD. Pataz: Tayabamba, Lopez &
Sagdstegui 3405 (USM). PIURA. Huancabamba: near
Huancabamba, Friedberg s.n. (USM); no exact locality,
Ellenberg 1116 (USM), Edwin 3621 (USM).
10. Onoseris humboldtiana Ferreyra, Publ. Mus.
Hist. Nat. "Javier Prado," Ser. B, Bot. 11.2.
1959. TYPE: Peru, lea, Nasca, Lomas de San
Nicolas, 500 m, Ferreyra 13400 (holotype,
USM 33422; isotype, USM 16000, MO).
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
69
Perennial herbs, procumbent, acaulescent, cau-
dex woody, lanuginose. Leaves alternate; petioles
5-10 mm long, lanuginose; blades spathulate, 15-
40 mm long, 1 1-20 mm wide, attenuate at base,
lanuginose, acute, the margins more or less den-
tate. Capitulescences of terminal, solitary heads,
peduncles 38-75 mm long, lanuginose. Capitula
radiate; involucres 1 8-2 1 mm high, 7-9 mm wide,
campanulate; phyllaries 7-8-seriate, lanceolate, the
inner 12-15 mm long, 1.6-2 mm wide, attenuate
at apex, the abaxial surfaces arachnoid; ray florets
21-28, the corollas 18-22 mm long, the tube 5-
5.5 mm long, pubescent, the outer lip 13-16.5 mm
long, 3-3.8 mm wide, violaceous, 6-nerved, the
abaxial surfaces arachnoid, tridentate, the inner
lip bipartite, 2-2.5 mm long, filiform; disc florets
14-16, the corollas 10-12 mm long, yellow, the
tube 9-10.5 mm long, the lobes acute; anthers 5-
5.6 mm long. Achenes [ray] 2-2.6 mm long, 1 mm
wide, pubescent; pappus ca. 8 mm long, isomor-
phic, yellow; [disc] 1.8-2.2 mm long, pubescent;
pappus ca. 9 mm long, isomorphic, yellow.
DISTRIBUTION— Known only from the type lo-
cality.
Onoseris humboldtiana closely resembles O.
amplexicaulis, from which it differs in having a
woody, basal caudex, spathulate leaves, and 7-8-
seriate involucres.
11. Onoseris linearifolia Sagast., Phytologia 57:
415. 1985. TYPE: Peru, Cajamarca, Caja-
bamba, alrededores de Cajabamba, 2600 m,
18 Nov 1983, Sagdstegui et al. 11264 (ho-
lotype, HUT; isotypes, F, HUT).
Perennial herbs, stems to 30 cm long, erect to
ascending, terete, lanate. Leaves sessile; blades
narrowly linear, 10-25 mm long, 1.5-2.5 mm wide,
basally attenuate, apically obtuse, mucronate, the
adaxial surfaces lanuginose, the abaxial surfaces
densely tomentose, the margins entire, revolute.
Capitulescences of solitary, terminal heads; pe-
duncles ca. 1 5 cm long, lanuginose, minutely brac-
teate, the bracts ca. 1.5 mm long. Capitula radiate;
involucres campanulate, 10-12 mm high, 8-10
mm wide; phyllaries 4-5 -seriate, the outer ovate
to ovate-lanceolate, 1.5-2.5 mm long, 0.8-1 .2 mm
wide, apically acute, the abaxial surfaces lanose,
the inner oblanceolate, 8-10 mm long, 1.8-2.2
mm wide, apically acute, the abaxial surfaces lan-
ose; ray florets 9-1 1, the corollas lilac, ca. 25 mm
long, the tube ca. 6 mm long, the abaxial surfaces
glabrous, the outer lip violaceous, ca. 1 2-20 mm
long, ca. 3 mm wide, 6-nerved, the abaxial surfaces
arachnoid, tridentate, the inner lip bifid, the lobes
filiform, ca. 1 mm long; disc florets 15, the corolla
8-10 mm long, the tube ca. 7-8 mm long, the lobes
acute, yellow, glabrous; anthers 3 mm long.
Achenes [ray] cylindric, ca. 6 mm long, sparsely
pilose; pappus ca. 8 mm long, isomorphic, stra-
mineous; [disc] cylindric, ca. 6 mm long, sparsely
pilose; pappus 8-10 mm long, isomorphic, stra-
mineous.
DISTRIBUTION— Known only from region of the
type locality.
Onoseris linearifolia most closely resembles O.
chrysactinioides but is easily distinguished by its
narrowly linear leaves with obtuse apices, smaller
capitula, and 4-5-seriate involucre.
CAJAMARCA. Cajabamba: ca. 8 km NW of Caja-
bamba, Dillon et al. 2857 (F); entre Chancay y Valle
Condebamba, Sanchez et al. 956 (F, USM).
12. Onoseris lopezii Ferreyra, Publ. Mus. Hist.
Nat. "Javier Prado," Ser. B, Bot. 11:4. 1959.
TYPE: Peru, La Libertad, Santiago de Chuco,
Los Quengos, A. Lopez M. 439 (holotype, USM
33500, originally cited as USM 1 600 1 ; isotype,
HUT). Figure 14.
Perennial herbs, to 40 cm high; stems procum-
bent, lanuginose. Leaves sessile; blades linear, 1 7-
65 mm long, 2.5-8 mm wide, apically acuminate,
bases semiamplexicaul, the adaxial surfaces arach-
noid, the abaxial surfaces lanuginose, the margins
entire. Capitulescences of solitary, terminal heads;
peduncles 17-24 cm long, lanuginose, bracteate,
the bracts 5-25 mm long, 1-2 mm wide, subulate,
lanuginose. Capitula radiate; involucres 16-19 mm
high, 14—17 mm wide, campanulate; phyllaries 9-
11 -seriate, lanceolate, the inner 11-16 mm long,
1.8-2.2 mm wide, attenuate at apex, the abaxial
surfaces arachnoid; ray florets 20-24, the corollas
35—44 mm long, the tube 4.8-6 mm long, 1 mm
wide, glabrous, the outer lip violaceous, 33-38
mm long, 4-5 mm wide, 6-nerved, the abaxial
surfaces arachnoid, tridentate, the inner lip bipar-
tite, 1.5-3 mm long, attenuate at apex, twisted;
disc florets 50-55, the corolla 13-16 mm long, the
tube 10.5-13 mm long, the lobes acute, yellow,
glabrous; anthers 7-8 mm long. Achenes [ray] ca.
3.5 mm long, 1 mm wide, pubescent; pappus ca.
1 1 mm long, isomorphic, yellow; [disc] 2.5—4.2
mm long, 1 mm wide, pubescent; pappus ca. 14
mm long, isomorphic, yellow.
70
FIELDIANA: BOTANY
3
1cm
FIG. 14. Onoseris lopezii. 1, habit; 2, ray floret with achene; 3, disc floret with achene. (Drawn from M.O. Dillon
& M. Whalen 4053, F.)
DISTRIBUTION— Known from the Andean Cor-
dillera of northern Peru (2500-3500 m).
Onoseris lopezii is related to O. gnaphalioides
but differs in its basal amplexicaul leaves and larg-
er involucre and phyllaries.
ANCASH. Huaras: above Huaras, Ferreyra 14305
(USM), Munoz 1004 (USM); Tacllan, Proano 85 (USM); SW
of Huaras, Pennell 15275 (USM), 75279 (USM). CAJA-
MARCA. Cajamarca: Encanada, Ferreyra 15001 (F, MO,
NY, USM); near Cajamarca, Fosberg 28144 (MO, USM); 8
km E of Cajamarca, King & Bishop 9124 (F); El Cumbe,
Lopez 2618 (USM), 1 302 (USM); arriba de San Juan, Lopez
8736 (F), 8744 (F); SW of Cajamarca, Pennell 15014 (MO,
USM); Pullucana, Sagdstegui et al. 8056 (F, MO); arriba
de Santa Apolonia, Sagdstegui 9965 (F); Quebrada de la
Esperanza, Sanchez & Tonal 3281 (F); ca. km 25 be-
tween Cajamarca and Celendin, Smith & Sanchez 4233
(F). Celendin: ca. 3 1 km NE of Cajamarca, Dillon &
Turner 1579 (F); ca. 24 km E of Celendin on road to
Balsas, Dillon & Whalen 4053 (F); above Balsas, Hutch-
ison & Wright 5210 (F, MO, NY, USM); Llanguat, Mos-
tacero et al 963 (F); near Celendin, Rauh 40295 (USM).
LA LIBERTAD. Huamachuco: near Huamachuco, Lopez
1005 (USM); Hacienda Cochabamba, Lopez & Sagdstegui
2783 (USM); Cachibamba, Infantes s.n. (MO, USM). San-
tiago de Chuco: Cachicadan, Sagdstegui et al. 1 1821 (F).
1 3. Onoseris macbridei Ferreyra, Publ. Mus. Hist.
Nat. "Javier Prado," Ser. B, Bot. 2: 2. 1949.
TYPE: Peru, Cajamarca, Chota, Yamaluc, R.
Ferreyra 846 (holotype, USM 33511, USM
15629 in original citation; isotype, F).
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
71
Perrenial herbs, 6-14 cm high, acaulescent, cau-
dex tomentose. Leaves in basal rosettes, 12-14,
petiolate; petioles 8-32 mm long, winged, the lobes
triangular, acute; blades oblanceolate in outline,
the distal portion ovate-hastate, 28-100 mm long,
1 0—40 mm wide, acute, the adaxial surfaces arach-
noid, the abaxial surfaces lanuginose, the margins
sinuose-dentate. Capitulescences of solitary, ter-
minal heads; peduncles 55-100 mm high, lanu-
ginose, bracteate, bracts 3.5-6 mm long, 1-1.5 mm
wide, subulate, the abaxial surfaces lanuginose.
Capitula radiate; involucres 20-27 mm high, 10-
12 mm wide, turbinate; phyllaries 1 1-12-seriate,
3-nerved, the inner 21-24 mm long, 1.4-2 mm
wide, attenuate at apex, the abaxial surfaces arach-
noid-glandulose; ray florets 1 8-23, the corollas 2.9-
3.3 mm long, the tube 9.5-10 mm long, glandular-
pubescent, the outer lip 19-23 mm long, 3-3.5
mm wide, 6-nerved, the abaxial surfaces arach-
noid-glandulose, tridentate, the inner lip bipartite,
filiform; disc florets with corollas 13.6-15.2 mm
long, the tube 11.6-12.8 mm long, the lobes acute,
coiled, yellow, the abaxial surfaces pubescent; an-
thers 5.8-6 mm long. Achenes [ray] 3-6 mm long,
1 mm wide, pubescent; pappus ca. 1 1 mm long,
isomorphic, yellow; [disc] 3.6-4 mm long, pubes-
cent; pappus ca. 1 3 mm long, isomorphic, yellow.
DISTRIBUTION— Native to the intermontane re-
gion of northern Peru (1800-2900 m).
Onoseris macbridei most closely resembles O.
speciosa, from which it differs in having smaller
habit, solitary capitula, turbinate involucres, and
1 1-12-seriate phyllaries with recurved apices.
ANCASH. Yungay: Mancos, Smith & Bias 4914 (F,
USM). CAJAMARCA. Chota: Ferreyra 8458 (MO, USM);
Lajas, Sagdstegui 14044 (F). Cutervo: near Cutervo, Fer-
reyra & Acleto 15379 (MO, USM). Jaen: Pucara to Cha-
maya, Plowman et al. 14257 A (F). Santa Cruz: Hacienda
El Molino, Munoz 1012 (USM).
14. Onoseris minima Domke, Notizbl. Bot. Gart.
Berlin-Dahlem 13: 247. 1936. TYPE: Peru,
Arequipa, Socosani, NW of Arequipa, 1933-
1934, D. B. Stafford 295a (holotype, K).
Annual herbs, 2-10(-14) cm high, erect, lanu-
ginose. Leaves sessile; blades lanceolate, 1 8-45 mm
long, 3-12 mm wide, basally attenuate, apically
acuminate, the adaxial surfaces arachnoid, the ab-
axial surfaces lanuginose, the margins slightly den-
tate. Capitulescences of solitary, terminal heads;
peduncles 5-35 mm long, lanuginose. Capitula ra-
diate; involucre 9-12 mm high, 4-6 mm, narrowly
campanulate; phyllaries 4-5 -seriate, the inner 10-
12 mm long, 2-2.2 mm wide lanceolate, attenuate
at apex, the abaxial surfaces arachnoid; ray florets
7-9, the corollas 10-11 mm long, the tube 3.2-3.5
mm long, glabrous, the outer lip 6.8-7.5 mm long,
1.6-1.8 mm wide, 6-nerved, the abaxial surfaces
arachnoid, tridentate, the inner lip filiform; disc
florets 8-10, the corollas 6-6.5 mm long, the tube
5-5.2 mm long, 5-lobed, the lobes acute; anthers
2.2-2.4 mm long. Achenes [ray] ca. 2.7 mm long,
pubescent; pappus ca. 4.5 mm long, heteromor-
phic, yellow; [disc] 2.6-3 mm long, pubescent;
pappus ca. 6 mm long, heteromorphic, yellow.
DISTRIBUTION— Known only from the type lo-
cality in southwestern Peru (1200-1900 m).
Onoseris minima closely resembles O. annua
but differs in its slightly sinuous-dentate leaf mar-
gins and attenuate, glabrous phyllaries.
MOQUEGUA. Mariscal Nieto: Monte Estuquina, NW
of Moquegua, Weberbauer 7440 (F, GH, NY, us); between
Rio Las Trancas and "Llaxwa," Hrdlicka s.n. (us).
15. Onoseris odorata (D. Don) Hook. & Arn.,
Companion Bot. Mag. 1: 103. 1835.
Chaetachlaena odorata D. Don, Trans. Linn. Soc.
London 16: 257. 1832. TYPE: "in Guayaquila
Peruvianorum," /. Tafalla s.n. (holotype, P, not
seen).
O. longipedicellata Muschl., Bot. Jahrb. Syst. 50, Beibl.
Ill: 96. 1913. TYPE: Peru, Lima, San Barto-
lome, estacion via ferrea entre Lima y Oroya,
1700-1800 m, A. Weberbauer 1698 (holotype, B,
presumably destroyed; photograph ex B, FM neg.
15894).
O. parva Muschl., Bot. Jahrb. Syst. 50, Beibl. 1 1 1: 95.
1913. TYPE: Peru, Arequipa, Mollendo, A. We-
berbauer 1492 (holotype, B, presumably de-
stroyed; photograph ex B, FM neg. 15895).
Centroclinium reflexum Hook., Bot. Mag. 58. pi. 3114.
1831. TYPE: Peru, cultivated at GL from seed
supplied by Cruickshanks, no voucher seen.
O. reflexa (Hook.) Less., Syn. Comp. 1 19. 1832.
[For additional synonymy, see Ferreyra ( 1 944).]
Annual herbs, 4-70 cm high, lanuginose. Leaves
subsessile to petiolate; petioles 5-10 mm long;
blades lanceolate, 1 2-90 mm long, 4-25 mm wide,
the adaxial surfaces arachnoid, the abaxial sur-
faces lanuginose, acute at each end, the margins
dentate. Capitulescences of solitary, terminal heads;
peduncles 25-230 mm long, bracteate. Capitula
radiate; involucres 8-15 mm high, 5-8 mm wide,
campanulate; phyllaries 5-6-seriate, the inner 1 2-
18 mm long, 1.5-2 mm wide, apically long-atten-
uate to acuminate, recurved; ray florets 1 3-2 1 , the
72
FIELDIANA: BOTANY
corollas violaceous or purple, 1 4-30 mm long, the
tube glabrous, 2.5-4 mm long, the outer lip 11-
26 mm long, 2.5-5 mm wide, glabrous, 6-nerved,
tridentate, the outer lip bipartite; disc florets with
corollas 7-1 1 mm long, the tube geniculate; an-
thers 3-5 mm long. Achenes [ray] 1.8-4 mm long,
slightly pubescent; pappus ca. 6 mm long, hetero-
morphic, yellow; [disc] 2.2—4 mm long, pubescent;
pappus ca. 8 mm long, heteromorphic, yellow.
DISTRIBUTION— Known from along the Peru-
vian coast and western slopes of the Andean Cor-
dillera (100-3700 m).
Onoseris odorata has a remarkable distribution
extending from the coastal desert to high-elevation
habitats within the Andean Cordillera. It is easily
recognized by its long-attenuate to acuminate, re-
curved phyllaries and membraneous, lanceolate
leaves.
No type material of O. longipedicellata has been
located; however, a photograph of the holotype
clearly shows the reflexed, long-attenuate phyllar-
ies common to O. odorata, and the leaves are with-
in the size range for that species (e.g., Sagdstegui
& Dillon 12291).
While no type material of O. reflexa has been
examined in this study, the excellent illustration
in the original publication shows the reflexed, long-
attenuate phyllaries characteristic of O. odorata.
ANCASH. Bolognesi: between Chasquitambo and
Cajacay, Ferreyra 14289 (USM); between Conococha and
Chasquitambo, Ferreyra 14470 (USM); 70 km E of Para-
monga, Richardson 2084 (MO, USM); abajo de Raquia,
Sagdstegui et al. 12291 (F). Huaras: Chancas, below Pun-
ta Caillan, Ferreyra 14631 (USM), 14395 (MO, USM).
Huaylas: bajada de Huaylas a Huallanca, Mostacero et
al 1977 (F). Recuay: Marca, Gomez 121 (USM). ARE-
QUIPA. Camana: Lomas de Ocona, Ferreyra 1 1712 (MO,
USM); Lomas de Camana, Ferreyra 8850 (MO, USM), Rich-
ardson 2152 (USM), Tovar 3449 (USM). Caraveli: between
Nasca and Chala, Ferreyra 1384 (USM); Lomas de Ati-
quipa, Ferreyra 6488 (USM), 8820 (USM), 727/2 (MO,
USM), 13492 (MO, USM), 19834 (USM); Lomas de San
Nicolas, Ferreyra 13417 (MO, USM); Lomas de Jahuay,
Ferreyra 19853 (USM); Lomas de Chaparra, Ferreyra 1409
(MO, USM); between Chala and Atico, Ferreyra 6353 (MO,
USM); Atico, Ferreyra 2513 (MO, USM), 8884 (MO, USM),
77555 (MO, USM), 72075 (MO, USM), 75957 (USM), 75625
(MO, USM); Lomas de Capacc, Hutchison 1279 (F, NY);
near Chala, Raimondi s.n. (USM); Puerto Chala Rauh 62
(MO); Lomas de Capac, near Chala, Tovar 2694 (USM),
2705 (MO, USM), 2667 (MO, USM). Islay: Mollendo, Cha-
vez et al. 43 (USM); Lomas de Mollendo, Ferreyra 12594
(MO, USM); between Mollendo and Matarani, Ferreyra
11568 (MO, USM); Lomas de Mejia, Ferreyra 6411 (USM);
Mollendo, West 8240 (MO). AYACUCHO. Lucanas: be-
tween Nasca and Puquio, Ferreyra 5454 (F, MO, USM),
5481 (USM). CAJAMARCA. Cajamarca: La Yunga, Ca-
banillas & Dioses 14 (MO); Chilete-San Pablo, Sagdstegui
et al. 7915 (MO); between Magdalena and San Juan, Ugent
5392 (USM). Chota: between Huambos and Llama, Fer-
reyra 871 (MO, USM). Contumaza: between Llallen and
Tembladera, Ferreyra 8598 (MO, USM); El Portachuelo
de San Antonio, Lopez et al. 9208 (F); abajo de Rupe,
Lopez et al. 9218 (F); El Portachuelo, Sagdstegui et al.
9227 (F). Jaen: near Jaen, Angulo 2115 (F, NY). San Mi-
guel: Casa Blanca, Sagdstegui et al. 8775 (F); Quinden,
Sanchez et al. 3038 (F). San Pablo: Tayas, Sanchez 2813
(F); La Moyuna, Sanchez et al. 2997 (F). ICA. lea: near
lea, Ferreyra 2542 (USM). Nasca: Lomas de Pongo, Rauh
& Hirsch P476 (NY). LA LIBERTAD. Otusco: between
Samne and Otusco, Ferreyra 7659 (USM); above Samne,
Ferreyra 14087 (MO, USM); Hacienda Membrillar, Var-
gas 370 (USM). Trujillo: Laredo to Simbal, Angulo &
Lopez 459 (USM); Lomas de Viru, Angulo & Lopez 460
(USM); Cerro Chiputur, Lopez & Lopez 9150 (F); Lomas
de Viru, Mostacero et al. 1444 (F); Cerro Cabezon, Sa-
gdstegui & Cabanillas 8744 (F), Sagdstegui & Mostacero
10444 (F). LAMBAYEQUE. Chiclayo: Tinajones, Cer-
rate 5254 (MO, USM); Hacienda Espinal, Ferreyra 12377
(MO, USM). Jaen: between Olmos and Motupe, Cerrate
4996 (USM); km 713, Cerrate 5201 (USM); Portachuelo a
Olmos, Cerrate 5220 (MO, USM), Ferreyra 6043 (MO, USM),
9757 (MO, USM); Naupe, Lopez et al. 7680 (MO); Olmos,
Ferreyra et al. 16304 (USM). LIMA. Canta: near Quives,
Ferreyra 19870 (USM); near Canta, Ferreyra 7249 (MO,
USM); Canta-Yaso, Ferreyra 18354 (USM); near Yaso,
Tovar 1025 (MO, USM), Ferreyra 7249 (MO, USM); near
Viscas, Ferreyra 14454 (F), 14468 (F, GH, USM), 74479
(F, USM); Quive, Pennell 14302 (F); Obrajillo, Capt. Wilkes
Exped. s.n. (F, GH); cerros N de Chosica, Weberbauer
170 (USM). Huarochiri: near Chosica, Weberbauer 5314
(F), Ferreyra 6110 (USM); between Oscolla and Surco,
Ferreyra 9179 (MO, USM); Surco, Ferreyra 2026 (MO, USM);
between Chosica and Matucana, Ugent 5286 (USM); Tor-
namesa, Ferreyra 7031 (USM); km 49, Lima-Oroya, Cer-
rate 4295 (USM); km 56, Lima-Oroya, Ferreyra 8240
(MO, USM), 70455 (USM), 77077 (MO, USM), 75556 (MO,
USM), Cerrate 5330 (MO, USM); km 61-62, Lima-Oroya,
Ferreyra 9163 (MO, USM), 9777 (USM); km 70, Lima-
Oroya, Cerrate 5090 (MO, USM), Ferreyra 3477 (MO, USM),
5927 (MO, USM), 10545 (USM); below Surco, Ferreyra
16893 (MO, USM); Monte Zarate, Ferreyra 9746 (USM);
near Matucana, Mexia 4088 (MO). Lima: Lomas de Man-
chay, Biegman s.n. (USM); 7 km E of Trapiche, Hutchison
1014 (F, USM), 7027 (F, USM). TUMBES. near Cancas,
Ferreyra 12205 (MO, USM).
16. Onoseris peruviana Ferreyra, J. Arnold Arb.
25: 373. pi. 4. 1944. TYPE: Peru, Junin, Tar-
ma, La Merced, J. F. Macbride 5425 (holo-
type, us 1191547; isotype, F 536457).
Perennial herbs, 1-2 m high, erect, canescent-
tomentose. Leaves petiolate, rosulate at base; pet-
ioles 85-195 mm long, winged with ovate-lanceo-
late lobes, 3-8 pairs, opposite or alternate, 48-58
mm long, 1 8-28 mm wide; blades oblanceolate in
outline, the distal portion hastate, 100-190 cm
long, 86-205 mm wide, the adaxial surfaces arach-
noid, glabrescent, the abaxial surfaces lanuginose,
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
73
apically acuminate, the margins sinuose-dentate.
Capitulescences paniculate with 7-9 heads; pe-
duncles 40-46 cm long, lanuginose, bracteate, the
bracts 3-6 mm long, subulate, arachnoid. Capitula
radiate; involucres 14-20 mm high, 7-10 mm wide,
campanulate; phyllaries 6-7-seriate, the inner 1 8-
20 mm long, 2.4-2.5 mm wide, acuminate, arach-
noid; ray florets ca. 13, the corollas 19-20 mm
long, the tube 9.5-10 mm long, the outer lip vi-
olaceous, 9.5-10 mm long, 1.8-2 mm wide,
6-nerved, the abaxial surfaces arachnoid, triden-
tate, the inner lip 6-7 mm long, attenuate, twisted;
disc florets with corollas 16-17 mm long, the tube
14.8-15.6 mm long, the lobes acute, yellow; an-
thers 4.5-4.8 mm long. Achenes [ray] 4.2-5 mm
long, 1 mm wide, pubescent; pappus ca. 14 mm
long, isomorphic, yellow; [disc] 4.5-5 mm long,
1-1.2 mm wide, pubescent; pappus ca. 15 mm
long, isomorphic, yellow.
DISTRIBUTION— Known from lower elevations
along the eastern escarpment of the Andean Cor-
dillera of central to southern Peru (600-900 m).
Onoseris peruviana most closely resembles O.
silvatica Greenman from Costa Rica but differs in
possessing a tomentose habit and hastate leaves
with dentate margins.
CUSCO. Convencion: Hacienda Luisiana, Dudley
11487 (F, USM). JUNIN. Tarma: Valley of Chancha-
mayo, Ferreyra 20294 (USM); Colonia Perene, Killip &
Smith 24937 (us); Rio Seco, Chanchamayo, Soukup 2501
(USM); Satipo, Woytkowski 5882 (GH, uc).
17. Onoseris speciosa Kunth, Nov. Gen. Sp. 4: 7.
t. 305. 1820. TYPE: Ecuador, Chimborazo,
"Alausi," Bonpland 3235 (holotype, B, pre-
sumably destroyed; isotype, F; photograph ex
B, FM neg. 15897).
Onoseris stuebelii Hieron., Bot. Jahrb. Syst. 21: 366.
1895. TYPE: Peru, "Tambo de Carizal, in valle
fluminis Utcubamba," A. Stiibel 35a (holotype,
B, presumably destroyed; isotype, GH; photograph
CXB, FM neg. 15898).
Perennial herbs, 30-65 cm high, acaulescent,
tomentose. Leaves in basal rosette; petioles 18-
160 mm long, winged, lobed, the lobes ovate 1-4
pairs opposite or alternate; blades lyrate in outline,
35-100(-300) mm long, 20-90(-150) mm wide,
apically acute, rarely obtuse, the adaxial surfaces
arachnoid, the abaxial surfaces lanuginose, the
margins sinuose-dentate. Capitulescences corym-
bose, 1-3-headed; peduncles to 60 cm long, lanugi-
nose-pubescent, bracteate, bracts 3-5 mm long,
subulate. Capitula radiate; involucre 18-24 mm
high, 1 1-16 mm wide, campanulate; phyllaries 6-
7-seriate, the inner 20-24 mm long, 1.2-2 mm
wide, lanceolate, attenuate at apex, arachnoid-
glandulose; ray florets ca. 21, the corollas 29-39
mm long, the tube 8-9.5 mm long, pubescent, the
outer lip 21-30 mm long, 2.8-3.8 mm wide,
6-nerved, arachnoid-glandulose, tridentate, the in-
ner lip bipartite, 8-14 mm long, attenuate, twisted
at apex; disc florets with corollas 14-16 mm long,
the tube 12.2-15 mm long, the lobes 1.8-2.8 mm
long, acute; anthers 6-7 mm long. Achenes [ray]
4-6 mm long, 1.2 mm wide, pubescent; pappus
ca. 12 mm long, isomorphic, yellow; [disc] 3.8-6
mm long, 1.2 mm wide, pubescent; pappus ca. 14
mm long, isomorphic, yellow.
DISTRIBUTION— Known from the Andean Cor-
dillera of southern Ecuador to northern Peru ( 1 1 00-
2600 m).
Onoseris speciosa is easily recognized by its
acaulescent habit and Capitulescences with 1-3 large
heads on peduncles to 40 cm long.
AMAZON AS. Bagua: Bagua Grande, Ferreyra 14433
(MO, USM). Bongara: Pomacocha-Shipasbamba, Wur-
dack 1097 (MO, USM). Chachapoyas: between Chacha-
poyas and Tingo, Ferreyra 7132 (F, MO, NY, USM); near
Chachapoyas, Ferreyra 14442 (MO, USM); Pumachaca,
Friedberg s.n. (USM); Cerros Calla-Calla, Hutchison &
Wright 5453 (F, USM); Taquia, SW of Chachapoyas, Pen-
nell 15175 (USM), 75572 (USM); Sonche to Chachapoyas,
Wurdack 1533 (GH, NY, USM); without locality, Maekawa
3053 (NY). Luya: Yunquilla Cevasco s.n. (USM). CAJA-
MARCA. Cajabamba: Condebamba valley, Smith &
Vasquez 3393 (F). Cajamarca: Magdalena, Guzman s.n.
(USM); Quebrada San Vicente, Pennell & Reichlin 15015
(USM); E of Cajamarca, Sanchez 762 (F, USM); between
Chancay and Valley of Condebamba, Sanchez 949 (F,
USM); Vista Alegre, Sagdstegui et al. 10141 (F). Celendin:
Hacienda El Limon, Ferreyra 15069 (USM). Chota: Mon-
tan, near Cochabamba, Ochoa 1544 (USM). Cutervo: above
Socota, Stork & Morton 10147 (F, uc). Jaen: Chontali,
Chimoy 33 (USM); between Pucara and Porculla, Ferreyra
13712A (USM); 13712 (USM); 13632 (USM); Colasay,
Woytkowski 7046 (uc). LAMBAYEQUE. Lambayeque:
near Abra Porculla, Ferreyra 13624 (USM), 13751 (MO,
USM). PIURA. Ayavaca: below Ayavaca, Ochoa 1799
(USM). Huancabamba: El Tambo, Acleto 238 (USM), 255
(USM); near Huancabamba, Acleto 467 (USM); Quebrada
Shumaya, Fosberg 27874b (MO, USM); Porculla, Llatas
2449. (F).
18. Onoseris weberbaueri Ferreyra, Journ. Arn.
Arb. 25: 386. 1944. TYPE: Peru, Cajamarca,
Jaen, between Rios Tabaconas and Maranon,
1100-1300 m, A. Weberbauer 6163 (holo-
74
FIELDIANA: BOTANY
type, F 628892; photograph ex F, FM neg.
49246).
Perennial herbs, to 65 cm high, erect, lanugi-
nose. Leaves in basal rosettes; blades lanceolate,
70-120 mm long, 15-25 mm wide, sessile, the
adaxial surfaces arachnoid, the abaxial surfaces
lanuginose, apically acuminate, the margins sin-
uose-dentate. Capitulescences of solitary, terminal
heads; peduncles 35-42 cm long, lanuginose, bracts
2-2.5 mm long, subulate. Capitula radiate; invo-
lucres 18-20 mm high, 8-10 mm wide, campan-
ulate; phyllaries 6-7-seriate, the inner 1 7-20 mm
long, 1.2-2 mm wide, acute, arachnoid; ray florets
ca. 13, the corollas violaceous, 36-38 mm long,
the tube 1 1-12 mm long, the outer lip 25-26 mm
long, 3.6-4 mm wide, 6-nerved, the abaxial sur-
faces arachnoid-pubescent, tridentate, the inner lip
bipartite, 13-14 mm long, attenuate at apex, twist-
ed; disc florets with corollas 17-18 mm long, the
tube 15-16 mm long, the lobes 2.5-2.8 mm long,
violaceous; anthers 5-6 mm long. Achenes [ray]
4-5 mm long, to 1 mm wide, pubescent; pappus
ca. 14 mm long, isomorphic, yellow; [disc] 3-4.5
mm long, ca. 1 mm wide; pappus ca. 1 5 mm long,
isomorphic, yellow.
DISTRIBUTION— Native to the savannas of in-
termontane regions in northern Peru (380-1300
m).
Onoseris voeberbaueri most closely resembles O.
speciosa but is distinguished by the long peduncles
and lanceolate leaves with dentate margins.
AMAZONAS. Bagua: near Pongo de Rentema, Rio
Maranon, Ferreyra 13671 (MO, USM); Bagua Chica, Fer-
reyra 14186 (MO, USM), 13643 (MO, USM); 22 km N of
Bagua Chica, Gentry et al. 23110A (F); Bagua Chica,
Hutchison & Wright 3632 (F, K, MO, NY, us, USM). CA-
J AM ARC A. Jaen: above Shume Alto, Ferreyra 19630
(USM); Quebrada San Agustin, Ferreyra & Sanchez 1 9657
(USM); near Jaen, Friedberg 579 (MO, USM); Pucara, Rio
Huancabamba, Hutchison & Wright 3522 (F, MO, NY);
Pucara to Chamaya, Plowman et al. 14257 (F); Santa
Rosa, Rio Maranon, Woytkowski 5735 (USM).
XVI. PEREZIA
Perezia Lag., Amer. nat. Espan. 1:31.1811. TYPE:
Perdicium magellanicum L. fil. = Perezia ma-
gellanica (L. fil.) Lag.3
For additional synonymy, consult Vuilleumier ( 1 969).
Perennial herbs, rhizomatose, or tap rooted,
caulescent or scapose; caudex glabrous; stems erect
or procumbent, pubescent, glabrous or glandular,
lacking spines. Leaves basal or caulescent, the
margins entire, lobed, or pinnatilobed, the lobes
rounded, ciliate, spinulose. Capitulescences many-
branched panicles or cymes or of solitary capitula.
Capitula homogamous, radiate; involucres cam-
panulate to cylindric; receptacles plane, epaleate;
phyllaries 3-7-seriate, imbricate, lanceolate to ob-
long, soft or rigid; florets hermaphroditic, iso-
morphic, the corollas blue, pink, or whitish, bi-
labiate, the outer lip tridentate, the inner lip bifid;
anthers sagittate; styles cylindric, the branches api-
cally truncate, papillose. Achenes cylindric or ob-
long, glandulose, papillose-pubescent; pappus of
scabrid to barbellate bristles, 1-2-seriate, white,
tawny, or ochraceous.
Perezia has recently been redefined to include
only the South American section Perezia; North
American members of Perezia section Acourtia
have been elevated to generic status (Reveal and
King, 1973; Crisci 1974). Vuilleumier (1969)
monographed section Perezia and treated 30 spe-
cies distributed in high-elevation habitats of the
Andean Cordillera (Colombia, Ecuador, Peru,
Chile, Bolivia, and Argentina) to lowland steppe
and grassland habitats (Argentina, Brazil, Para-
guay, and Uruguay). In Peru, seven species are
recorded from high-elevation environments
throughout the Andean Cordillera.
The species concepts and synonymy presented
here largely reflect the detailed monographic work
of Vuilleumier (1969), and more detailed discus-
sions of variation and distribution can be found
in that publication.
References
CRISCI, J. V. 1974. A numerical-taxonomic study
of the subtribe Nassau viinae (Compositae, Mu-
tisieae). J. Arnold Arb., 55: 568-610.
REVEAL, J. L., AND R. M. KING. 1973. Re-es-
tablishment of Acourtia D. Don. Phytologia, 15:
228-232.
TOVAR, O. 1955. Revision de las especies perua-
nas del genero Perezia (Compositae). Publ. Mus.
Hist. Nat. "Javier Prado," Ser. B, Bot., 8: 1-35.
VUILLEUMIER, B. 1969 (1970). The systematics
of Perezia sect. Perezia (Compositae). Contr.
Gray Herb., 199: 1-163.
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
75
Key to Peruvian Species of Perezia
1 . Plants essentially acaulescent with heads sessile in leafy rosette, or with flowering scapes equal to or
less than 10 cm high; cauline leaves less than 2 cm long, bracteiform 2
2. Plants generally shorter than 3 cm tall; basal rosette of 5 or fewer, lyrate leaves 1-3 cm long, 4-
8 mm wide P. pygmaea
2. Plants generally taller than 3 cm; basal rosette of 5 or more, pinnatifid to entire leaves (2-)4-10
cm long, (5-)10-20 mm wide 3
3. Leaf blades with entire, densely and evenly ciliolate to spinulose margins P. ciliosa
3. Leaf blades pinnatilobed with 6-15 pairs of lobes, the margins with long, contorted cilia . 4
4. Leaves with lanceolate lobes, apically acute to attenuate, the margins serrate to dentate;
the outer phyllaries ovate, apically acute, entire or ciliate-spinulose P. coerulescens
4. Leaves with lance-ovate lobes, apically obtuse to rounded, the margins conduplicate; the
outer phyllaries oblong to pandurate, apically rounded, ciliate P. pinnatifida
1 . Plants caulescent with heads borne on leafy scapes or stems ( 1 0-)20-80 cm high; cauline leaves greater
than 2 cm long 5
5. Basal leaf blades lanceolate to oblanceolate, the margins irregularly dentate; capitulescences of
dense panicles or corymbs of 10-50 heads; involucres 10-12 mm high, 8-9 mm wide
P. multiflora
5. Basal leaf blades oblanceolate to spathulate, the margins entire to lobed-dentate; capitulescences
of solitary, terminal heads or weakly paniculate with 2-9 heads; involucres 15-30 mm high, 10-
20(-35) mm wide 6
6. Flowering scapes decumbent; outer phyllaries entire P. sublyrata
6. Flowering scapes erect; outer phyllaries serrate or spiny P. pungens
1 . Perezia ciliosa (Phil.) Reiche, Anal. Univ. Chile
116:426. 1905.
Clarionea ciliosa Phil., Anal. Mus. Nac. Chile 8: 35.
1891. TYPE: Chile, Tarapaca, Cerro de Copa-
coya, 3500 m, 18 Feb 1885, F. Philippi 2250
(holotype, SGO, not seen; isotype, LP, not seen;
photograph ex B, FM neg. 1 6066).
Small resetted, perennial herbs, 3-10 cm tall;
underground rhizome thick. Basal leaves elliptical
or lanceolate, 20-100 mm long, 5-20 mm wide,
glabrous, apically acute, the margins entire, dense-
ly and evenly ciliate; petioles winged and mem-
branous; stem leaves 2-11, lanceolate, 5-10 mm
long, 2-3 mm wide, scalelike apically acute, ciliate,
glabrous. Capitulescences of solitary heads, ter-
minal; peduncles 3-10 cm long, 1-7 per rosette.
Capitula with involucres campanulate, 8-2 1 mm
high, 9-20 mm wide; phyllaries 4-6-seriate, the
outer ovate, 3-9 mm long, 1-3 mm wide, apically
acute, entire or ciliate, the margins scarious, the
inner lanceolate, 7-20 mm long, 1-3 mm wide,
apically acute, entire, the margins scarious, red-
dish; florets blue or whitish, 14-43, the outer flo-
rets 8-25 mm long, the ligules 2-7 mm long.
Achenes 4-5 mm long, villous; pappus 6-13 mm
long.
DISTRIBUTION— Known from the Andean Cor-
dillera of southern Peru (Arequipa) to Bolivia and
Argentina (3500-4500 m).
Perezia ciliosa was reported from Arequipa
(Volcan El Misti, Stafford 578, K) by Vuilleumier
(1969, p. 104); however, no material referable to
this species has been encountered in this study and
the description is taken from the previously men-
tioned work.
2. Perezia coerulescens Wedd., Chlor. And. 1: 39,
10 A. 1855. TYPE: Peru, Cusco, C. Gay s.n.
(holotype, p, not seen).
Perezia nivalis Wedd., Chlor. And. 1 : 39. 1855. TYPE:
Peru, Carabaya, 1847, Weddell 1848 (holotype,
p, not seen; photograph ex p, FM neg. 38 1 26), non
Homoianthus nivalis Phil.
Perezia coerulescens Wedd. var. amplibracteata To-
var, Publ. Mus. Hist. Nat. "Javier Prado," Ser.
B, Bot. 8: 16. 1955. TYPE: Peru, Huancavelica,
Huantanayoc-Tansiri, O. Tovar 1129 (holotype,
USM 15714).
Perezia cirsiifolia Weddell, Chlor. And. 1:41. 1855.
TYPE: Bolivia, Larecaja, Cordillera de Sorata,
5100 m, H. Weddell s.n. (holotype, P, not seen;
isotype fragment ex P, F 971659; photograph ex
p, FMneg. 38122).
Perennial herbs, acaulescent, rhizomatose. Bas-
76
FIELDIANA: BOTANY
al leaves in rosettes, petiolate; petioles 6-27 mm
long, 4-6 mm wide, winged, membranaceous;
blades oblanceolate to spathulate, 1 5-38 mm long,
6-12 mm wide, apically ovate, the margins deep
wavy, pinnatilobed, 6-1 1 pairs of lobes, or sinu-
ate-denticulate, rarely more or less entire, ciliate,
slightly pubescent; cauline blades lanceolate, 10-
20 mm long, 3-5 mm wide, glabrescent, the mar-
gins dentate-spinulose. Capitulescences of soli-
tary, terminal heads; scape to 30 mm long, 1-4 in
each rosette. Capitula with involucres campanu-
late 12-22 mm high, 7-12 mm wide; phyllaries
4-6-seriate, the outer oblong to ovate, dentate to
spiny or rarely entire, the inner 1 3-20 mm long,
2-5 mm wide, elliptic-lanceolate to linear, apex
spinose, pubescent; florets 7-25, the corollas blue,
violet or whitish, 20-25 mm long, the tube 12-16
mm long, the outer lip 8-9 mm long, 2-4 mm
wide, glabrous, the inner lip bipartite, coiled; an-
thers ca. 7 mm long; styles 14-16 mm long. Achenes
cylindric, 2.5-3.5 mm long, 1-1.5 mm wide, 5-6-
costate, pubescent; pappus ca. 1 5 mm long.
DISTRIBUTION— Known from the Andean Cor-
dillera of Ecuador, Peru, Bolivia, and northwest-
ern Argentina (3800-5000 m).
Perezia coerulescens is a highly variable species
most closely resembling P. pinnatifida but is dis-
tinguished from that species by its lanceolate lobes
and ovate outer phyllaries with dentate, ciliate-
spinose margins. Vuilleumier (1969, p. 138) re-
ported hybridization between the two aforemen-
tioned species and some collections possess char-
acteristics of both. In P. coerulescens, the leaf mor-
phology is highly variable with a trend from dis-
sected to entire leaves visible from mid-Peru
through Bolivia. The name P. virens (D. Don)
Hook. & Arn. has been applied to material here
referred to as P. coerulescens; however, P. virens
is a synonym of P. poeppigii Less., a central Chil-
ean species (Vuilleumier, 1969, p. 144).
ANCASH. Carhuaz: Quebrada Ishinca, Smith & Bud-
densiek 11209 (F). Recuay: ca. 3 km below Cahuish Tun-
nel, Smith & Goodwin 8757 (F). Santa: Conocrangra,
Mostacero et al. 1929 (F). Yungay: above Yungay, We-
berbauer 3277 (USM). AREQUIPA. Arequipa: above
Chiguata, Straw 2303 (USM). Cailloma: Viscachani-
Pampa de Tocra, Perea 10 (USM); Cailloma, Schmidt s.n.
(F 1222045); Pichu Pichu, Stafford 808 (F). AYACU-
CHO. Lucanas: 93 km E of Puquio, Gentry et al. 23293
(F). CUSCO. Cusco: Pico Seneca, Vargas 2171 (F). Quis-
picanchis: Hacienda de Urco, Schmidt s.n. (F 1222644).
Urubamba: Chincheros, Taucca, Kinget al. 187 (F); Mac-
chu-Picchu, Peyton 99 (MO), 724 (MO); Yucay, Soukup
721 (F); Santa Rosa, Stafford 556 (F). HUANCAVELI-
CA. Castrovirreyna: Chiclococha, Tovar 2888 (GH, USM).
Huancavelica: above Kircay, Amstutz 115 (MO); Huay-
tanayocc-Tansiri, Tovar 1149 (USM); Tansiri, near Man-
ta, Tovar 1152 (USM), 7756 (GH), 7756 (USM). HUANU-
CO. Dos de Mayo: Valle Huallanca, Tovar & Rivas 9922
(USM); Yanashallas, Macbride & Featherstone 2481 (F,
GH, us). Huanuco: Mito, Macbride & Featherstone 1936
(F, GH, us). JUNIN. Huancayo: Huancayo, Soukup 3235
(F). Jauja: Ricran, Perez 37 (USM). Yauli: Laguna Punrun,
Aguilar s.n. (USM); Yauli-San Cristobal, Duncan et al.
2664 (F, MO); Cazapata, Encarnacion 193 (USM); near
Morococha, Grant 7562 (F); Oroya, Kalenborn & Kalen-
born 178 (MO, us); Pampa de Junin, Tovar 2384 (USM);
near Capillacocha, Carhuamayo, Tovar 2414 (USM).
LIMA. Canta: La Viuda, Diaz, s.n. (USM); Mishquipu-
quio, Huamantanga, Sanchez 30 (USM), 31 (MO, USM);
Carhuapampa, carretera Canta-Lachaqui, Meza 160
(USM); above Huascoy, Waechter s.n. (USM). Huarochiri:
Ticlio, between Casapalca and Oroya, Diers 926 (USM);
above Casapalca, Mina Caprichosa, Hutchison & Tovar
4250 (F, GH, MO, uc, us, USM); Casapalca, Macbride &
Featherstone 843 (F, GH); Antaicocha, Pennell 14661 (F,
us). MOQUEGUA. Mariscal Nieto: between Torata &
Carumas, Dillon et al. 3349 (F). PUNO. Azangaro: Sal-
cedo, Mt. Putina, Vargas 9626 (F). Carabaya: ca. 20 km
SEofMacusani, Dillon etal. 7059(F);Fauchinta, Vargas
7169 (USM); without locality, Soukup 364 (F, USM); Rauh
s.n. (USM). Huancane: Moho, Aguilar s.n. (USM). Puno:
San Antonio de Esquilache, Stafford 706 (F), 738 (F), 741
(F), 577 (F).
3. Perezia multiflora (Humb. & Bonpl.) Less.,
Linnaea 5: 19. 1830.
Chaetanthera multiflora Humb. & Bonpl., PI. Aequin.
2: 168, pi. 135. 1809. TYPE: Ecuador, Pichincha,
Volcan Antisana, Humboldt & Bonplands.n. (ho-
lotype, P, not seen; photograph ex p, FM neg.
38125).
Perezia bidentata Meyen, Reise urn die Erde 1: 470.
1 834. TYPE: Peru, Puno, Talaram, 1 4000-1 5000
ft, Apr 1831, Meyen s.n. (presumably destroyed,
B, photograph ex B, GH).
Perennial herbs, erect, 10-70 cm high, glandu-
lar-pubescent. Basal leaves in rosette, petiolate or
sessile; petioles 10-20 mm long, membranaceous;
blades usually lanceolate to oblanceolate in out-
line, 60-250 mm long, 10-30 mm wide, coria-
ceous, glabrescent, the margins irregular dentate,
spinulose, mucronate; cauline leaves linear-lan-
ceolate, 20-75 mm long, 8-26 mm wide, basally
amplexicaul, glandular-pubescent. Capitules-
cences of panicles or corymbs; scapes 1 0—40 cm
high, glandulose. Capitula with involucres cam-
panulate, 10-12 mm high, 8-9 mm wide; phyl-
laries 3-4-seriate, the outer ovate to triangular,
spiny toward the bases, the margins irregular den-
tate, the inner ovate-lanceolate, 9-10 mm long, 3-
3.2 mm wide, acuminate, spinulose, 3-nerved,
glandular-pubescent; florets 34-44, the corollas
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
77
blue, whitish or violaceous, 6-10 mm long, the
tube 5-6 mm long, glabrous, the outer lip 3.5-4
mm long, 1.8-2 mm wide, 4-nerved, glabrous, tri-
dentate, the inner lip bipartite, ca. 3 mm long,
linear-lanceolate, slightly coiled; anthers 2.8-3 mm
long; styles filiform, 6-7 mm long. Achenes 2.5-
4 mm long, 1-1.5 mm wide, cylindric, pubescent;
pappus ca. 8.6 mm long.
DISTRIBUTION— Known from the Andean Cor-
dillera of northern Peru, Bolivia, and northwestern
Argentina (3500^800 m).
Perezia multiflora is readily distinguished by its
basal rosette of lanceolate, spinulose and irregu-
larly dentate leaves, robust paniculate capitules-
cences with 9 or more heads, and blue corollas. It
is abundant along the Andean Cordillera where it
is used in folk medicine and called escorzonera.
ANCASH. Huaras: ca. 35 km W of Huaras, Dillon et
al. 3140 (F); Punta Cayllan, Ferreyra 14326 (USM). Huari:
near Tunel Cahuish, Ames s.n. (USM), Smith & Budden-
siek 1 1086 (MO, USM). Huaylas: Quebrada Los Cedros,
Alpamayo, Smith et al. 9879 (F). Recuay: Carpa, Smith
& Escalona 10220 (F). APURIMAC. Andahuaylas: near
Andahuaylas, Ferreyra 2806 (MO, USM). AREQUIPA.
Arequipa: no exact locality, Guenther & Buchtien 911
(HBG). AYACUCHO. Lucanas: near Puquio, Ferreyra
5496 (MO, USM). CAJAMARCA. Cajamarca: Porcon,
Sanchez 123 (F). Hualgayoc: near Coymolache, Ferreyra
8570 (MO, USM); vicinity of Bambamaraca, King & Bish-
op 9116 (MO). CUSCO. Calca: Amparaes, Chavez 3308
(MO); Pisac, Marfn 807 (USM), 1327 (F). Canas: Checca,
Vargas 1485 (MO); slopes of San Andres de Checca, Var-
gas 11021 (F). Espinar: Yauri, Nunez et al. 7855 (F).
Paruro: cerca a Occopata, Vargas 889 (F). Quispicanchis:
Ausangate, Humbert 30787 (us). Urubamba: Taucca, King
et al. 180 (F). LA LIBERTAD. Jalca de Huaguil, Lopez
1416 (MO, USM). HUANCAVELICA. Huancavelica:
Quebradas W from Huancavelica, Stork & Horton 10846
(F, GH); Machacchuay, Tovar 867 (GH); Tansiri, Huay-
tanayoc, cerca a Manta, Tovar 1174 (GH). HUANUCO.
Huanuco: above Huanuco, Ridoutt s.n. (USM); Huanuco,
Soukup 2182 (F). JUNIN. Huancayo: between Acopalca
and Pariahuanca, Stork 10937 (F); near Huancayo, We-
berbauers.n. (USM). Jauja: between Conception and Sa-
tipo, Ochoa 252 (USM). Junin: Oroya, Kalenborn 112
(MO, us); Hondores, Tovar 2395 (USM); Nevado Aco-
palca, Soukup 3684 (F, USM). Tarma: puna between Tar-
ma and Oroya, Ferreyra 3826 (USM), 70965 (USM); La
Oroya, Grant 7522 (F). Yauli: ca. 1 km W of Anticona
Pass, Dillon & Turner 1481 (F, MO, USM); Hacienda Pu-
cara, Hutchison & Tovar 4217 (F); near Atocsayco, Pear-
sail 845 (F), 7 182 (F). LIMA. Canta: Carhuapampa, Meza
165 (MO, USM); Auquimarca, Peraldo 3263 (F). Huaro-
chiri: Casapalca, Asplund 1 1440 (us); above Chicla, Fer-
reyra 6512 (MO, us, USM); Rio Blanco, Macbride & Feath-
erstone 654 (F, GH, us); Rio Blanco, Killip & Smith 21705
(F, us); Cerro Colorado near Antaicocha, E of Canta,
Pennell 14652 (F, GH, us). PASCO. Cerro de Pasco: Bos-
que de Piedras, Urquizo 3 (USM). PUNO. Carabaya: cerca
de abra de Antapampa, Vargas 7004 (F); between Ma-
cusani and Nunoa, Vargas 7143 (USM). Huancane: Moho,
Shepard 109 (MO, USM), Tincopalca, Lopez 7361 (MO);
near Huancane, Aguilar 149 (MO, USM). Puno: Cerros de
Puno, Zuniga 33 (USM); Araranca, Pennell 13429 (F, GH,
us). TACNA. Tacna: Cordillera de Volcan Tacora, Chis-
lluma, Werdermann 1 142 (F, GH, us).
4. Perezia pinnatifida (Humb. & Bonpl.) Wedd.,
Chlor. And. 1:40. 1855.
Chaetanthera pinnatifida Humb. & Bonpl., PI. Ae-
quin. 2: 170, pi. 136. 1809. LECTOTYPE (des-
ignated by Vuilleumier, 1969): Ecuador, Coto-
paxi, Humboldt & Bonpland s.n. (lectotype, pi.
136 in Plantae Aequinoctialis).
Perennial herbs to 10 cm tall, rhizomatose, the
rhizome incrassate. Basal leaves rosette, usually
longer than the stem, petiolate; petioles 27-70 mm
long, 5-8.5 mm wide, membranaceous, transpar-
ent; blades lanceolate to lyrate in outline, 35-60
mm long, 10-15 mm wide, the margins pinnatifid
with 10-15 pairs of lobes, slightly imbricate,
rounded, obtuse, glandular-pubescent, entire, cil-
iate; cauline leaves lanceolate, 23-45 mm long,
3.5-5.5 mm wide, sessile, the margins dentate,
pubescent, gradually smaller to the apex. Capitu-
lescences of solitary, terminal heads, the scapes 4-
9 cm long, pubescent. Capitula with involucres
campanulate, 10-25 mm high, 10-13 mm wide;
phyllaries 4-5-seriate, the outer oblong, rounded,
the inner linear-lanceolate, 1 5-24 mm long, 3-5
mm wide, acuminate, membranaceous, 3-nerved,
glandulose-pubescent, glabrescent distally; florets
20-25, the corollas blue-whitish, 15-24 mm long,
the tube 9-14 mm long, the outer lip 6-9 mm long,
2-3 mm wide, 4-nerved, apex tridentate, glabrous,
the inner lip bipartite, coiled; anthers 5-6 mm
long, sagittate; style filiform, 11-13 mm long,
branches truncate, papillose. Achenes 3-4 mm
long, 1-1.2 mm wide, cylindric, pubescent, 6-8
costate; pappus ca. 1 8 mm long.
DISTRIBUTION— Known from the Andean Cor-
dillera of Colombia, Ecuador, Peru, Bolivia, and
northwestern Argentina (3000-4600 m).
Perezia pinnatifida closely resembles P. coeru-
lescens, from which it differs in its incrassate rhi-
zome, elongated petioles, strongly pedunculate ca-
pitula, and pandurate outer phyllaries.
ANCASH. Bolognesi: Pampa de Lampas, Cerrate 1450
(MO, USM); Ocros, Weberbauer 2780 (USM); Rio Puma-
pampa, Tovar & Rivas 9640 (USM). Huaras: near Huaras,
Cerrate 7707 (MO, USM); between Huaras and La Union,
Tovar & Rivas 9852 (USM). APURIMAC. Andahuaylas:
near Andahuaylas, Hjerting & Petersen 1441 (USM).
78
FIELDIANA: BOTANY
AREQUIPA. Caraveli: near Quicacha, Guevara s.n.
(USM). CUSCO. Urubamba: Antakillqa, Davis et al. 1703
(F, USM); Chinchero, Franquemont & Franquemont 369
(F). HUANCAVELICA. Huancavelica: Paso de Chonta,
Tovar 2947 (GH, USM). JUNIN. Yauli: Yauli, Macbride
& Featherstone 929 (F); Nevado Acopalca, Soukup 3698
(MO, USM). LIMA. Canta: Auquimarca, Peraldo 3280 (F).
Huarochiri: Langa, Cerrate et al. 4725 (MO, USM); Lake
Avascocha, Casapalca, Saunders 1 150 (F). Yauyos: Hua-
cracocha, Cerrate & Tovar 639 (USM); Rio Blanco, Mac-
bride 3031 (F, GH). PASCO. Pasco: Huaron, Macbride
& Featherstone 1136 (F). PUNO. Puno: San Antonio de
Esquilache, Stafford 1280 (F).
5. Perezia pungens (Humb. & Bonpl.) Less., Lin-
naea 5: 20. 1830. Figure 15.
Chaetanthera pungens Humb. & Bonpl., PL Aequin.
2: 146, pi. 127. 1809. TYPE: Ecuador, Pichincha,
Humboldt & Bonpland s.n. (holotype, B, presum-
ably destroyed; photographs ex B, FM neg. 1 6084;
isotype, P, not seen).
Perezia ciliaris D. Don ex Hook. & Arn., Companion
Bot. Mag. 1: 34. 1835. LECTOTYPE (designated
by Vuilleumier, 1969): Chile,/. Dombeys.n. (lec-
totype, P; isolectotype, G, photograph ex G, FM
neg. 8278; isolectotype fragment ex P, F 972652).
Clarionea macrocephala Schultz-Bip. in Lechler, Berb.
Amer. Austr. 57. 1857. Vuilleumier (1969) lists
this taxon as a nomen nudum and cites Tovar's
(1955) transfer to Perezia as invalid.
Perezia stuebelii Hieron., Bot. Jahrb. Syst. 21: 372.
1895. TYPE: Peru, Cajamarca, between Pacas-
mayo and Moyobamba, A. Stubel 34 (holotype,
B, presumably destroyed, photographs ex B, FM
neg. 16088).
Perezia weberbaueri Hieron. ex Domke, Notizbl. Bot.
Gart. Berlin-Dahlem 13: 249. 1936. TYPE: Peru,
Puno, Sandia, Cuyocuyo, A. Weberbauer 933 (ho-
lotype, B, presumably destroyed; photograph ex
B, FM neg. 16091; isotype fragment, F 1012351).
Perezia conaicaensis Tovar, Publ. Mus. Hist. Nat.
"Javier Prado," Ser. B, Bot. 8: 31. 1955. TYPE:
Peru, Huancavelica, Huancavelica, Conaica, Lar-
ia, 8 km SW of Conaica, 3900-4000 m, 30 Mar
1952, O. Tovar 903 (holotype, USM; isotye, GH).
Perezia coriacea Tovar, Publ. Mus. Hist. Nat. "Javier
Prado," Ser. B, Bot. 8: 21. 1955. TYPE: Peru,
Huanuco, Huanuco, Mitotambo, above Mito, R.
Ferreyra 9431 (holotype, USM 31602; isotypes,
GH, MO).
Perezia fosbergii Tovar, Publ. Mus. Hist. Nat. "Javier
Prado," Ser. B, Bot. 8: 22. 1955. Peru, Cajamarca,
Celendin, Las Lajas, F. R. Fosberg 28123 (ho-
lotype, USM).
Caulescent, perennial herbs, 10-80 cm tall, rhi-
zomatose. Basal leaves in a loose rosette, or rarely
marcescent at flowering, sessile to petiolate; peti-
oles 10-120 mm long; blades spathulate to oblan-
ceolate, 50-200 mm long, 1 5-50 mm wide, grad-
ually tapering to an attenuate base, apically obtuse
to rounded, glabrous to strigillose or glandular-
puberulent, the margins entire to lobed-dentate,
slightly revolute, ciliate to spinulose; cauline leaves
lanceolate to elliptic or oblong, 1 5-50 mm long,
3-18 mm wide, sessile, the bases truncate or au-
riculate and clasping, apically attenuate to obtuse,
the margins entire to denticulate, ciliate to spi-
nulose. Capitulescences of solitary, terminal heads
or weakly paniculate with 2-9 heads, the primary
peduncles 10-80 cm long, bracteolate. Capitula
with involucres campanulate to turbinate, 1 5-30
mm high, 10-20 mm wide; phyllaries 4-6-seriate,
the outer lanceolate, narrowly ovate, or spathulate,
5-20 mm long, 3-7 mm wide, ciliolate, serrate or
spiny, coriaceous to scarious or membranous, the
inner lanceolate to oblong, 14-18 mm long, 3-5
mm wide, apically acuminate, entire to denticu-
late, glandular-puberulent, 3-nerved; florets 10-
68, the corollas white, blue, violet, or pinkish, 1 6-
21 mm long, the tube 8-10 mm long, the outer lip
7-10 mm long, 2.8-3.2 mm wide, 4-nerved, tri-
dentate, pubescent, the inner lip bifid, 6-7 mm
long, coiled; anthers 5-6 mm long; styles 10-12
mm long. Achenes cylindric, 2.2-4 mm long, 1-
1.5 mm wide, glandular-papillose to strigillose, 5-
8-costate; pappus 12-13 mm long.
DISTRIBUTION— Known from high-elevation
habitats within the Andean Cordillera of southern
Ecuador, Peru, Bolivia, and adjacent northern Ar-
getina (3000-4600 m).
Perezia pungens is readily recognized among Pe-
ruvian species by its basal rosette of spathulate
leaves and erect, foliose capitulescences with one
to nine heads. It most closely resembles P. sub-
lyrata but is distinguished from that species by its
generally longer stems, larger capitula, entire to
weakly dentate basal leaf blades, and ciliolate to
spinulose outer phyllaries. Perezia pungens is here
considered a highly variable species with several
different ecotypes recorded from within its range.
Vuilleumier (1969) recognized P. ciliaris as a dis-
tinct species but detected putative hybrids be-
tween it and P. pungens in the region of northern
Peru and southern Ecuador. She speculated (p. 95)
that these two species might merely be ecotypes,
and, after examining a large suite of specimens
from throughout the range, it appears likely that
P. ciliaris merely occupies drier sites and only dif-
fers quantitatively from typical P. pungens. Har-
ling (1991, p. 102) was also unable to distinguish
these two species in Ecuador and placed all within
the range of P. pungens.
AMAZONAS. Chachapoyas: Calla-Calla, Raimondi
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
79
FIG. 15. Perezia pungens. 1, habit; 2, floret with achene. (Drawn from A. Weberbauer 6973, F.)
80
FIELDIANA: BOTANY
s.n. (USM). ANCASH. Bolognesi: Laguna Huamanhue-
que, Cerrate 1556 (USM); Pampa de Lampas, Cerrate
1463 (USM); Cerros de Chiquian, Cerrate 1391 (USM).
Huaras: between Punta Cayllan and Cajamarquilla, Fer-
reyra 14310 (USM); arriba Huaras, Tovar & Rivas 9527
(USM); near Huaras, Weberbauer s.n. (USM). Huari: Que-
brada Rima Rima, Smith et al. 12206 (F); Quebrada de
Yuraccocha, Smith et al. 12725 (F). Huaylas: Quebrada
Alpamayo, Smith et al. 9797 (F); Auquispuquio, Smith
et al. 12096 (F). Recuay: between Recuay and Chavin,
Ferreyra 14532 (USM). Santa: arriba de Lamparin, Mos-
tacero et al. 1843 (F). Yungay: 25 km E of Yungay, Edwin
& Schunke 3808 (F); Llanganuco, Ferreyra 14357 (GH,
USM), 7 6545 (USM); Yungay to Yauya, Gentry et al. 37395
(F, MO, USM). AREQUIPA. Cailloma: Valle de Colca, ca.
100 km N de Arequipa, Treaty 770 (F). AYACUCHO.
Huanta: Hacienda Pargora, Killip & Smith 23303 (us).
CAJAMARCA. Cajamarca: Cumbe Mayo, Sanchez 3431
(F, CPUN). Celendin: Cruz Conga, Cumullca, Ferreyra
13277 (USM); Cumullca, jalca near Celendin, Ferreyra
15128 (USM); above Balsas, Hutchison & Wright 5214
(F, MO, uc); near Celendin, Ochoa 1515 (USM); Quinu-
amayo, Pennell & Reichlin 15055 (USM); Jalca de Ku-
mulca, Sagdstegui et al. 12055 (F). Contumaza: Casca-
bamba, Sagdstegui et al. 9030 (F); Las Campanulas, Sa-
gdstegui et al. 9153 (F); Pozo Kuan, Sagdstegui et al. 10076
(F), 70770 (F); Pozo Chuno, Sagdstegui et al. 14005 (F).
Hualgayoc: near Coymolache, Ferreyra 8573 (USM); Koi-
molache, Sanchez et al. 795 (F); Hualgayoc, Weberbauer
3957 (USM). San Miguel: arriba de El Tingo, Mostacero
et al. 1209 (F); Cerro Quillon, Sagdstegui et al. 9590 (F).
Santiago de Chuco: Cerro La Botica, Sagdstegui et al.
11880 (F). CUSCO. Calca: Pisac, Mann 1898 (F, us).
Convencion: Lucumayo Valley, Cook & Gilbert 1261 (us).
Cusco: Sacsayhuaman, Vargas 4091 (F); Ayanacu, Var-
gas 16268 (us). Paucartambo: Acjanaco, Cano 3404 (F);
Valle del Paucartambo, hacienda Churu, Herrera 2328
(F); Pillahuata Valley, Herrera 3335 (us); Tres Cruces,
Pennell 13814 (F, GH, us); Escalerayoc, Vargas 11180
(F); Acjanacu Pass, Vuilleumier 250 (F); Corihuayra-
china, Woytkowski 624 (USM). Quispicanchis: Quispi-
canchis, Herrera 1085 (us); Urubamba: Chincheros, Da-
vis et al. 1648 (F, USM), 7695 (F); Gutierrezchayoj, King
et al. 285 (F, USM); Lomas de Puyupatamarca, Vargas
2730 (F, USM). HUANCAVELICA. Huancavelica: Con-
aica-Laria, Tovar 792 (USM); Huando, Tovar 1281 (USM).
Tayacaja: Hacienda Alalay, between Mariscal Caceres
and Pampas, Tovar 1289 (USM); Motcca, Tovar 252 (us,
USM); Sachahuaccta, Tovar 934 (USM). HUANUCO. Dos
de Mayo: Huallanca, Tovar & Rivas 9906 (USM). Huan-
uco: Chinche, Macbride & Featherstone 1258 (F); Mito,
Macbride & Featherstone 1664 (F); 15 km SE of Huan-
uco, Macbride & Featherstone 2116 (F, GH). JUNIN.
Huancayo: Huancayo, Soukup 2971 (F). Junin: Laguna
Punrun, Aguilar s.n. (USM); Cordillera Raura, Rauh &
Hirsch P1831 (NY). Tarma: Acobamba, Woytkowski 69
(F). Yauli: Yauli, Macbride & Featherstone 925 (F). LA
LIBERTAD. Ottusco: Hacienda Llaguen, Lopez 605a
(USM). Pataz: between Tayabamba and Huancaspata,
Lopez & Sagdstegui 8204 (GH). LIMA. Cajatambo: Oyon,
Ferreyra 3552 (USM). Canta: Paella, above Canta, Meza
196 (USM). Huarochiri: Chaccha, Cerrate & Tovar 1600
(USM); Manzanacha, Tovar & Cerrate 1737 (USM); In-
fiernillo, Ferreyra 6243 (USM). PIURA. Huancabamba:
Huancabamba, TownsendA215 (F). PUNO. Huancane:
Moho, Aguillar 177 (USM); Amantani, Aguilar 446 (USM).
Sandia: above Cuyocuyo, Diaz s.n. (USM). No exact lo-
cality: Weberbauer 6973 (USM).
6. Perezia pygmaea Wedd., Chlor. And. 1: 40.
1855. TYPE: Bolivia, La Paz, ravine of Chu-
quiaguillo, H. Weddell s.n. (holotype, p, not
seen; photograph ex p, FM neg. 38128).
Perennial herbs to 3 cm high, rhizomatose.
Leaves in rosette, petiolate; petioles 3-10 mm long;
blades oblanceolate or elliptic-lanceolate, 10-30
mm long, 5-6 mm wide, basally attenuate, apically
obtuse, the margins pinnatilobed, rarely entire, 3-
5 pairs of lobes, ciliate, glabrescent. Capitules-
cences of solitary, terminal heads; subsessile, the
scapes glandulose-pubescent, bracteate, bracts lin-
ear. Capitula with involucres cylindric, 10-15 mm
high, 5-6 mm wide; phyllaries 4-5-seriate, the out-
er ovate, 7-15 mm long, 2-5 mm wide, broadly
scarious, the inner elliptic-lanceolate, 10-12 mm
long, 2-3 mm wide, greenish, mucronate, the api-
ces entire, slightly pubescent, 3-nerved; florets 7-
15, the corollas blue, 13-15 mm long, the tube 7-
8 mm long, the outer lip 6-7 mm long, 3-3.5 mm
wide, glabrous, 4-nerved, tridentate, the inner lip
linear-lanceolate, bipartite, 4-5 mm long; styles
9-1 1 mm long. Achenes 2-3 mm long, 1-1.5 mm
wide, 5-7-costate, glandular-papillose; pappus ca.
1 1 mm long, white.
DISTRIBUTION— Known from high-elevation
habitats within the Andean Cordillera from cen-
tral Peru to northwestern Argentina (4000-5100
m).
Perezia pygmaea closely resembles P. coerules-
cens but differs in possessing smaller leaves, cy-
lindric involucres, and phyllaries and pappus to
only 12 mm long.
CUSCO. Quispicanchis: Ausangate, Rauh 1140 (USM).
HUANCAVELICA. Castrovirreyna: Laguna Choclo-
cocha, Tovar 2920 (GH, USM). Huancavelica: cerca Man-
ta, Tovar 1115 (USM); JUNIN. Junin: Cazapato, Lago
Junin, Aguilar 1089 (USM). Yauli: near Morococha, Grant
7576 (GH). LIMA. Huarochiri: Mina Caprichosa, above
Casapalca, Hutchison & Tovar 4259 (F, NY, uc, us, USM);
Casapalca, Macbride & Featherstone 846 (F, GH); Laguna
Tuctococha, Tovar & Cerrate 1624 (USM). PUNO. Azan-
garo: Putina, Monheim 55 (USM).
7. Perezia sublyrata Domke, Notizbl. Bot. Gart.
Berlin-Dahlem 13: 248. 1936. TYPE: Peru,
Puno, Santa Rosa, 14,500 ft, Apr-May 1934,
D. Stafford 345 (holotype, K; isotype, BM, not
seen).
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
81
Perezia sublyrata var. glabrescens Tovar, Publ. Mus.
Hist. Nat. "Javier Prado," Ser. B, Bot. 8: 11.
1955. TYPE: Peru, Cusco, Paucartambo, near Rio
Quencomayo, below Colquepata, F. W. Fennel I
13789 (holotype; GH; isotypes, F 558098, NY).
Perennial herbs to 25 cm tall, rhizomatose. Bas-
al leaves in rosette, petiolate; petioles 20-40 mm
long, 3-3.5 mm wide at the base, membranaceous;
blades oblanceolate to spathulate in outline, con-
spicuously lyrate, 20-100 mm long, 11-20 mm
wide, basally attenuate, apically obtuse, glabres-
cent, the margins ciliate-spinulose; cauline leaves
linear-lanceolate to spathulate, 7-26 mm long, 2-
5.4 mm wide, sessile, the margins entire to re-
motely sinuate-dentate, mucronate. Capitules-
cences of solitary, terminal heads; scapes 35-1 10
mm high, decumbent, glandulose-pubescent. Ca-
pitula solitary, terminal; involucres campanulate,
16-20 mm high, 9-12 mm wide; phyllaries 4-6-
seriate, the outer oblong-lanceolate, 7-13 mm long,
2-6 mm wide, entire or slightly dentate at the base,
apically acute, mucronate, the inner 14-18 mm
long, 3-5 mm wide, ovate-lanceolate to lanceolate,
membranaceous, glabrescent, rarely glandulose
outside, 3-nerved, apically acute, the margins scar-
ious, entire; florets 20-24, the corollas violaceous,
21-24 mm long, the tubes 11-14 mm long, the
outer lip 8-10 mm long, 2-3 mm wide, glabrous,
4-nerved, tridentate, the inner lip bipartite, coiled;
anthers 5-6 mm long; styles 16-18 mm long.
Achenes 2.5-3 mm long, 1-1.5 mm wide, ovoid,
glandulose, 5-6-costate; pappus ca. 14 mm long.
DISTRIBUTION— Known from high-elevation
habitats from central Peru to Bolivia (3300-4400
m).
Perezia sublyrata is characterized by the lyrate
leaves with laminar petioles and decumbent flow-
ering scapes.
JUNIN. Yauli: near Oroya, Kalenborn 115 (GH, us).
PUNO. Puno: Araranca, Pennell 13442 (F, GH, us).
XVII. PLAZIA
Plazia Ruiz Lopez & Pavon, Fl. peruv. prodr. 104.
1794. TYPE: Plazia conferta Ruiz Lopez &
Pavon
Harthamnus Yi. Robinson, Phytologia 45: 451. 1980.
TYPE: Harthamnus boliviensis H. Robinson =
Plazia daphnoides Wedd.
Shrubs to 1 m, the branches erect; stems lacking
spines. Leaves simple, sessile, in whorls at branch
tips; blades oblanceolate, glabrous to glandular,
usually appressed, the margins entire. Capitules-
cences of solitary, terminal heads, sessile. Capitula
heterogamous, radiate; involucres campanulate;
receptacles plane, glabrous; phyllaries 5-7-seriate,
lanceolate; ray florets 7-20, the corollas ligulate-
bilabiate, the outer lip 4-nerved, tridentate, the
inner lip bifid; styles cylindric, glabrous, bifid, the
branches short, inconspicuous; disc florets 10—42,
the corollas tubular, actinomorphic, glabrous, the
limb deeply 5-lobed, the lobes greater than '/? the
length of the corolla, coiled; anthers linear, the
terminal appendages lanceolate, fused into a col-
umn, truncate, the bases caudate; styles claviform,
the branches short, rounded. Achenes dimorphic,
glabrous or glandular-pubescent; pappus of sca-
brid bristles, isomorphic, yellow.
Plazia consists of perhaps three species confined
to the Andean Cordillera of Peru, Bolivia, Chile,
and Argentina. The genus is easily recognized by
its woody habit, sessile capitula, and purple to
pink, connate anther appendages exserted from the
disc florets. In Peru, the following two species are
recorded from central to southern departments,
usually in high-elevation, dry sites.
References
CABRERA, A. L. 1951. Notas sobre Compuestas
de la America Austral. Darwiniana, 9: 363-386.
. 1 960. Notas sobre tipos de Compuestas
Sudamericanas en herbarios Europeos. Ill, Los
tipos de Ruiz y Pavon. Bol. Soc. Argent. Bot.,
8: 195-215.
FERREYRA, R. 1980. El genero Plazia en el Peru.
Bol. Soc. Peruana Bot., 8: 101-105.
ROBINSON, H. 1980. Harthamnus, a new genus
of Mutisieae from Bolivia. Phytologia, 45: 45 1-
455.
Key to Peruvian Species
1 . Leaves 22-42 mm long, 5-6 mm wide, glabrous; capitula with ca. 42 florets; ray florets 1 8-20; achenes
glabrous P. conferta
82
FIELDIANA: BOTANY
1. Leaves 10-20 mm long, 3-4 mm wide, glandular-pubescent; capitula with 17-19 florets; ray florets
7-8; achenes glandular-pubescent P. daphnoides
1 . Plazia conferta Ruiz Lopez & Pavon, Syst. veg.
fl. peruv. chil. 187. 1798. TYPE: Peru, Junin,
Acobamba [near Tarma], Ruiz Lopez & Pa-
von s.n. (holotype, MA, not seen).
Shrubs, branched, branches glabrous. Leaves
oblanceolate, 22-42 mm long, 5-6 mm wide, gla-
brous, sessile, acute-mucronate, margin entire.
Capitula with involucres 18-22 mm high, 12-16
mm wide; phyllaries 6-7-seriate, glabrous, lanceo-
late, the inner 22-25 mm long, 3-3.5 mm wide,
acute, the outer gradually smaller; ray florets 1 8-
20, the corollas 26-28 mm long, the tube 10-1 1
mm long, glabrous, the outer lip 14-15 mm long,
5-6 mm wide, 4-nerved, tridentate, the inner lip
bipartite; disc florets 40-42, the corollas 15-18
mm long, the tube glabrous, the lobes 9-10 mm
long, 1-1.2 mm wide, coiled; anthers 6-7 mm long.
Achenes [ray] 4-5 mm long, 1.5-1.6 mm wide,
glabrous; pappus ca. 12 mm long; [disc] 4.5-5 mm
long, 1-1.2 mm wide; pappus ca. 14 mm long.
Plazia conferta is a rare species, and no new
material has been collected until the gathering made
by Felix Woytkowski at the type locality more than
100 years after its original description. Cabrera
(1960) was unsuccessful in locating Ruiz and Pa-
von's type material in the major European her-
baria, including Madrid.
DISTRIBUTION— Endemic to the Andean Cordi-
llera of central Peru.
JUNIN. Tarma: Acobamba, betweenTarma and San
Ramon, Woytkowski 52 (USM).
2. Plazia daphnoides Wedd. Chlor. And. 1: 13,
pi. 2B. 1 855. LECTOTYPE (here designated):
Peru, "Cordillera de Tacora, entre Tacna et
La Paz," Weddell s.n. (lectotype, P, not seen;
isolectotype ex P, F 971331). Figure 16.
Harthamnus boliviensis H. Robinson, Phytologia 45:
451. 1980. TYPE: Bolivia, Cochabamba, SE of
Cochabamba, vicinity of Rodeo, 3500 m, 5 Mar
1979, .7. A. Hart 1 73 9 (holotype, us 2854177, not
seen; photograph ex us, FM neg. 1944785).
Shrubs to 2 m, resinose. Leaves subsessile; blades
oblanceolate to lanceolate, 10-20 mm long, 3^4
mm wide, apically acute, basally cuneate, glan-
dular-pubescent, the margins ciliate. Capitula with
involucres 18-20 mm high, 7-8 mm wide; phyl-
laries 5-6-seriate, pubescent, lanceolate, the inner
20-21 mm long, 2.4-3 mm wide, acuminate, the
outer smaller; ray florets 7-8, the corollas 20-21
mm long, geniculate, the tube 1 1-1 1.5 mm long,
the outer lip 9-9.5 mm long, 2-3 mm wide, tri-
dentate, the inner lip bipartite, 6-7 mm long, coiled;
disc florets 10-11, the corollas white, 13-14 mm
long, the tube geniculate, 10-11 mm long, the lobes
7-8 mm long, coiled; anthers 5-6 mm long.
Achenes [ray] 4.5-5 mm long, 1.2-1.4 mm wide,
glandular; pappus ca. 13 mm long; [disc] 5-5.5
mm long, 1 mm wide, glandular; pappus ca. 12.5
mm long.
DISTRIBUTION— Known from the Andean Cor-
dillera of southern Peru, Bolivia, Chile, and Ar-
gentina (3500-4000 m).
Plazia daphnoides displays the greatest distri-
bution and frequency within the genus. The lec-
totype was selected from the available syntypes in
the original protolog, which also included a Bo-
livian collection (d'Orbigny 1386, P) designated as
(8. The lectotype was collected by Weddell in the
Cordillera Tacora near Tacora, a region now part
of northern Chile, and there can be little doubt
that Weddell derived his description from mate-
rial he had collected in the area that was Peru until
the 1870s.
Robinson's (1980) new genus and species, Har-
thamnus boliviensis, was discovered to be a syn-
onym of Plazia daphnoides by Robinson shortly
after its publication.
AYACUCHO. Ayacucho: 83 km W of Puquio, Gentry
et al. 23273 (F); Coracora, Keel 698 (USM); near Puquio,
Rauh P423 (USM).
XVIII. POLYACHYRUS
Polyachyrus Lag., Amer. nat. Espan. 1: 37. 1811.
TYPE: Cephaloseris poeppigii Kunze ex Less.
= Polyachyrus poeppigii (Kunze ex Less.) Less.
Annual or perennial herbs, erect or decumbent,
branched, branches terete, pubescent or glabrous.
Leaves alternate; blades lanceolate to oblanceolate
in outline, the bases amplexicaul to decurrent, the
apices more or less mucronate, membranaceous
or succulent, glabrous to glandular, the adaxial
surfaces lanose, the abaxial surfaces albo-lanugi-
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
83
FIG. 16. Plazia daphnoides. 1, habit; 2, capitulum, 3, disc floret with achene; 4, ray floret with achene. (Drawn
from A. Gentry et al. 23273, F.)
nose, the margins entire to pinnatilobed or pin-
natisect, the lobes or teeth mucronate or spiny.
Capitulescences of glomerulate clusters of 9 or more
heads, solitary or pseudocorymbose. Capitula with
2 (or rarely 3) florets, sessile; involucre cochlei-
form with 5 bracts; receptacles plane, glabrous;
phyllaries 4-5-seriate, oblong, glabrous; florets
homogamous, the corollas reddish or violaceous,
84
FIELDIANA: BOTANY
bilabiate; anthers 5, exserted, sagittate; style
branches apically obtuse. Achenes oblong, glan-
dular-puberulent to pilose; pappus of plumose
bristles, 1 -seriate.
Polyachyrus contains seven species distributed
throughout the Andean Cordillera of Peru and
Chile, with the greatest diversity in the coastal
lomas formations. In Peru, one species is recorded
from Andean Cordillera and two from adjacent
coastal localities.
References
RICHARDI, M., AND E. WELDT. 1974. Revision
genero Polyachyrus (Compositae). Gayana, 26:
3-34, pis. 1-7.
Key to Peruvian Species of Polyachyrus
1 . Plant annual, erect P. annuus
1 . Plant perennial, decumbent 2
2. Leaves succulent, the blades oblong, the adaxial surface glandular, the abaxial surfaces arachnoid
P. fuscus
2. Leaves membranaceous, the blades lanceolate, the adaxial surfaces arachnoid, the abaxial surfaces
white-tomentose P. sphaerocephalus
1. Polyachyrus annuus Johnston, Contr. Gray
Herb. 8: 134. 1929. TYPE: Chile, Antofagas-
ta, Tocopilla, Caleta Duendes, I.M. Johnston
3538 (holotype, GH).
P. mollendoensis Johnston, Contr. Gray Herb. 8: 134.
1929. TYPE: Peru, Arequipa, Islay-Mollendo, /.
M. Johnston 3539 (holotype, GH).
Annual herbs, 18-35 cm high, erect, stem pu-
bescent. Leaves amplexicaul, membranaceous;
blades oblong to lanceolate, 35-1 80 mm long, 10-
50 mm wide, the adaxial surfaces arachnoid, the
abaxial surfaces lanuginose, the bases auriculate,
the auricle orbicular, 1-3 mm wide, the margins
pinna tilobed. Capitulescences of 1-3 glomerules,
weakly cymose. Capitula biflorous, terminal or ax-
illary; peduncles 2-10 cm long, lanuginose; florets
homogamous in pairs: 1 outer lip floret with 1
bract and 1 interior floret with 4 bracts, 4-5 mm
long, 2.5-3 mm wide, the corollas reddish to vi-
olaceous, fragrant, the tube 2.5-3 mm long, the
outer lip 3.5-3.8 mm long, 1.8-2 mm wide, the
interior bilobed, 3.5-3.8 mm long, the lobes coiled,
glabrous. Achenes oblong, ca. 1 mm long, pilose;
pappus (outer florets) 0.7-1 mm long, (inner flo-
rets) 3-4 mm long.
DISTRIBUTION— Known from coastal habitats in
southern Peru to northern Chile (50-700 m).
AREQUIPA. Camana: near Camana, Ferreyra 6437
(MO, USM), 8833 (MO, USM), 77654 (MO, USM). Caraveli:
Atico, Ferreyra 2518 (us, USM), 2675 (MO, USM), 11538
(MO, USM), 72027 (USM), 75625 (MO, USM). Islay: ca. 10
km NW of Islay, Dillon & Dillon 3957 (F); Mollendo,
Ferreyra 12598 (MO, USM), 75650 (MO, USM); Mejia, Fer-
reyra 6407 (MO, USM); Mollendo, Stafford 901 (F); Mol-
lendo, Vargas 8591 (GH); E of Mollendo, Worth & Mor-
rison 15758 (GH, MO). MOQUEGUA. Mariscal Nieto:
near Ilo, Ferreyra 11611 (MO, USM), 72577 (MO, USM);
Mostacilla, Vargas 8591 (USM). TACNA. Tacna: Morro
Sama, Ferreyra 12556 (MO, USM).
2. Polyachyrus fuscus (Meyen) Walpers in Meyen,
Nov. Actorum Acad. Caes. Leop.-Carol. Nat.
Cur. 19, Suppl. 1: 288. 1843.
Diaphoranthus fuscus Meyen, Reise um die Erde 1:
406. 1864. TYPE: Chile, Cordillera de Copiapo,
4000 ft, F. J. F. Meyen s.n. (holotype, B, presum-
ably destroyed; photograph ex B, FM neg. 1 5992).
P. nesites Johnston, Contr. Gray Herb. 95: 85. pi. 7.
fig. 4.1931. TYPE: Peru, lea, Pisco, Isla San Gal-
Ian, R. Murphy 3467 (holotype, GH).
Suffrutescent herbs, decumbent, branched,
branches terete, glandular, 30-80 cm tall. Leaves
succulent, amplexicaul, sessile; blades oblong, 40-
70 mm long, 12-21 mm wide, the adaxial surfaces
densely glandular, the abaxial surfaces lanuginose
arachnoid, the margins pinnatilobed, the segments
6-10 pairs, opposite or alternate, the lobes mostly
triangular, revolute. Capitulescences of 1-3 glom-
erules, 15-17 mm in diameter. Capitula with in-
volucres of several phyllaries, linear, glabrous; flo-
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
85
FIG. 17. Polyachyrussphaerocephalus. 1, habit; 2, capitulum; 3, achene with pappus. (Drawn from P.C. Hutchison
1256, F.)
86
FIELDIANA: BOTANY
rets reddish. Achenes ca. 1 mm long, densely glan-
dular-puberulent; pappus ca. 5 mm long.
DISTRIBUTION— Known from the coastal desert
of northern Chile and the Peruvian island of San
Gallan, near Pisco, Department of lea.
Polyachyrus fuscus is apparently a rare species
in Peru, with only one collection recorded from
Isla San Gallan, but it is more common and widely
distributed in northern Chile.
3. Polyachyrus sphaerocephalus D. Don, Trans.
Linn. Soc. London 16: 230. 1830. TYPE: Peru,
Ruiz Lopez & Pavon s.n. (holotype, P, not
seen). Figure 17.
P. glandulosusNutt., Trans. Amer. Philos. Soc.7 : 423.
1 84 1 . TYPE: Peru, Curson s.n. (holotype, PH, not
seen).
P. villosus Wedd., Chlor. And. 1: 56. 1855. TYPE:
Peru, Tacna, Cordillera de Tacora, d'Orbigny290
(holotype, P, not seen; isotype ex P, F 971771).
Suffrutescent herbs, decumbent, branched,
branches to 80 cm long, terete, glandular-arach-
noid, white-lanuginose. Leaves amplexicaul, the
upper sessile; blades oblong-lanceolate, 50-150
mm long, 12-60 mm wide, the adaxial surfaces
arachnoid, the abaxial surfaces white lanuginose,
the margins pinnatifid, 6-8 pairs of segments op-
posite. Capitulescences of 1-2 glomerules, termi-
nal, 18-23 mm in diameter. Capitula with corolla
reddish, fragrant, the tube 4-5 mm long, the outer
lip 3-3.5 mm long, 4-nerved, tridentate, the inner
lip bisect, coiled, 3-3.3 mm long; anthers 1.8-2
mm long. Achenes ca. 1 mm long, densely pilose;
pappus 5-6 mm long.
DISTRIBUTION— Known from the Andean Cor-
dillera of central to southern Peru (800-3800 m).
Polyachyrus sphaerocephalus is distinguished by
its white habit, reddish florets,and fragrant smell,
and it is the only member of the genus in Peru
occupying habitats above 1000 m.
ANCASH. Huaras: Punta Caillan, between Casma and
Huaras, Ferreyra 14394 (USM). Recuay: km 104 on road
from Pativilca to Recuay, Dillon et al. 3093 (F); Pativil-
ca-Huaras, Gentry et al. 37333 (F, MO, USM). AREQUI-
PA. Arequipa: Chachani, Arenas 168 (USM); Arequipa,
Douglass s.n. (GH); Chachani, Hinkley & Hinkley 49
(GH,US); Yura, Straw 2346 (us, USM). AYACUCHO. Lu-
canas: Puquio, Ferreyra 7200 (us, USM); Pachan, Hutch-
ison 1256 (F, GH, MO, uc, us, USM). Parinacochoas: Cor-
acora, Weberbauer 5795 (GH). LA LIBERTAD. Otuzco:
Cerro de los Enamorados, Leiva & Leiva 328 (F). LIMA.
Cajatambo: near Oyon, Ferreyra 3533 (MO, us, USM).
Cant a: Culluay, Capt. Wilkes Exped. s.n. (GH); Obrajillo,
Capt. Wilkes Exped. s.n. (us), Meza 217 (USM). Huar-
ochiri: Rio Blanco, Cerrate 909 (USM); Innernillo, Cerrate
& Jibaja 4872 (MO, USM); Tambo de Viso, Ferreyra 1241
(USM); Innernillo, Ferreyra 8812 (USM), Ferreyra et al.
14909 (USM); Innernillo, Goodspeed et al. 11606 (GH,
uc), Killip & Smith 21574 (F, us), Ferreyra 6989 (F, us,
USM), 8946 (USM), 5579 (us, USM); Lopez 487 (us), Fer-
reyra 13600 (USM); Chicla, Ferreyra & Tovar 20135 (USM),
Ball s.n. (GH); Chicla, Seler & Seler 238 (GH, us), Ma-
thews 641 (GH); Matucana, Macbride & Featherstone 333
(GH, us); Huaros, Pennell 14711 (GH, us); Santa Eulalia,
Goodspeed 33096 (GH, MO, us); Langa, Barclay 2341 (F,
us). MOQUEGUA. Mariscal Nieto: Mina Cuajone, Dil-
lon & Matekaitis 3401 (F); Minas Toquepala, Vargas
13067 (us); near Carumas, Weberbauer 7336 (us, USM).
XIX. PROUSTIA
Proustia Lag. Amenidades Naturales de las Es-
panas 1: 33. 1811. TYPE: Proustia pyrifolia
DC.
Lophopappus Rusby, Bull. Torrey Bot. Club 21: 487,
pi. 225. 1894. TYPE: Lophopappus foliosus Rus-
by.
Shrubs branched, unarmed, rarely with few nodal
spines or with branchlets apically spinescent.
Leaves alternate; blades ovate-lanceolate, lanceo-
late, oblong, or, rarely, linear, the margins entire
or denticulate. Capitulescences of solitary heads,
terminal and axillary or racemose. Capitula ho-
mogamous, discoid; involucres cylindric to cam-
panulate; receptacles naked; phyllaries 4-6-seri-
ate, ovate, oblong, or lanceolate, subcoriaceous,
glabrous or finely tomentose, puberulent, or res-
inous, apically acute to acuminate, often mucro-
nate; florets 5-15, the corollas bilabiate, white to
lavender, the lobes linear, coiled, the outer lip tri-
dentate, the inner lip bidentate; anthers sagittate,
apex acute; style cylindric, apically bifid, the
branches papillate. Achenes cylindric or turbinate,
4-5 costate, pubescent; pappus of barbellate to
plumose bristles, 1 -seriate.
Proustia is a Neotropical genus with eight spe-
cies, five of which occur within the Andean Cor-
dillera. After examining several species of Lopho-
pappus, I believe it is congeneric with Proustia. In
addition, the structure of the pollen grains of Loph-
opappus and Proustia are similar with tricolpate
and spinulose exine in both.
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
87
References
CABRERA, A. L. 1953. Compuestas peruanas
nuevas o criticas. Bol. Soc. Argent. Bot., 5: 45-
50.
FABRIS, H. A. 1968. Revision genero Proustia
(Compositae). Revista Mus. La Plata, Secc. Bot.,
11: 23-49.
Key to Peruvian Species of Proustia
1 . Leaves linear, the apices long-apiculate P. foliosa
1 . Leaves ovate or lanceolate, oblong, or spathulate 2
2. Upper branchets flexuose, apically acicular or spinulose P. cuneifolia
2. Upper branchets not apically acicular or spinulose 3
3. Leaves densely tomentose on abaxial surfaces; capitula with 5-6 florets per head P. cuneata
3. Leaves glabrous on abaxial surfaces; capitula with 7-1 5 florets per head 4
4. Leaf blades with only midvein prominent, the margins essentially entire or remotely den-
ticulate; capitula with 1 8 florets P. peruviana
4. Leaf blades with midvein and 2 lateral veins prominent, the margins repand, spinulose-
dentate; capitula with 7-9 florets P. berberidifolia
1 . Proustia berberidifolia (Cabr.) Ferreyra, comb,
nov.
Lophopappus berberidifolius Cabr., Bol. Soc. Argent.
Bot. 5: 48. pi. 8. 1953. TYPE: Peru, Huancave-
lica, Sachahuaceta, Conaica, O. TovarS. 961 (ho-
lotype, LP; isotypes, MO, USM).
Shrubs, 1-1.6 m high, branched, the branches
striate, glabrous, lacking spines. Leaves subsessile;
petioles 0.5-1 mm long; blades ovate-spathulate,
10-25 mm long, 5-10 mm wide, both surfaces
glabrous, resinous, basally cuneate, trinerved from
the base, apically rounded, the abaxial surfaces
prominent-reticulate, the margins repand, spinu-
lose-denticulate. Capitulescences of 3-4 heads,
more or less glomerate. Capitula sessile; involu-
cres campanulate, 10-12 mm high, 4-5 mm wide;
phyllaries 14-18, 3 -seriate, the outer ovate, 3-4
mm long, 2-2.5 mm wide, the inner oblong-ellip-
tic, (6-)l 1-12 mm long, 2-2.5 mm wide, apically
obtuse-mucronate, the margins pilose; florets 6-
9, the corollas white, 12-13 mm long, bilabiate,
outer lip tridentate, inner lip bipartite, coiled, 6-
7 mm long; tube 5-6 mm long, cylindric; anthers
5-6 mm long. Achenes turbinate, 3-5 mm long,
0.8-1 mm wide, pubescent, 5-costate; pappus of
plumose bristles, ochraceous, 7-8.5(-12) mm long.
DISTRIBUTION— Known from throughout the
Andean Cordillera of central to southern Peru
(2700-3600 m).
Proustia berberidifolia most closely resembles P.
peruviana but is readily distinguished from that
species by its resinous leaves with repand, spi-
nulose-dentate margins and typically solitary ca-
pitula. It is known by the common names aucu-
pichana and pucka-pucka.
AREQUIPA. Arequipa: Morro Verde, above Yura,
Straw 2356 (USM), Eyerdam & Beetle 22135 (GH, MO,
uc). Cailloma: Valle del Colca, Treacy 833 (F). AYA-
CUCHO. Lucanas: Puquio, Ferreyra 9972 (MO, USM),
9775 (MO, USM). Parinacochas: Chaipi, Leon 10 (F).
HUANCAVELICA. Huancavelica: Sachahuaceta, Alau-
ma, near Conaica, Tovar 921 (USM); between Mejorada
and Anco, Tovar 3330 (USM). JUNIN. Yauli: near Yauli,
Tovar 3016 (USM); Paccha, Tovar 3287 (USM). LIMA.
Huarochiri: Cuchilla-Blanca, Cerrate et al. 4675 (USM);
above San Mateo, Guttes.n. (USM). MOQUEGUA. Ma-
riscal Nieto: 50-57 km NE of Moquegua, Dillon et al.
4797 (F). TACNA. Tarata: above Tarata 1 5 km, Hutch-
ison & Wright 7204 (F); near Tarata, Rauh 40756 (USM).
2. Proustia cuneata S. F. Blake, Contr. U.S. Natl.
Herb. 22: 653. t. 63. 1924. TYPE: Peru, Ol-
lantaytambo, Cook & Gilbert 538 (holotype,
us). Non Lophopappus cuneata R.E. Fries,
1906.
L. blakei Cabr., Bol. Soc. Argent. Bot. 5: 48. 1953.
TYPE: (nom. nov. for P. cuneata S. F. Blake).
Much-branched, spineless shrubs; branchlets
albo-tomentose, glabrescent. Leaves subpetiolate;
petioles 1-2 mm long; blades oblanceolate, 12-25
mm long, 4-8 mm wide, basally cuneate, apically
obtuse or rounded, spinose-mucronate, repand-
denticulate distally, coriaceous, the adaxial sur-
faces light green, inconspicuously tomentose, gla-
brescent, the abaxial surfaces conspicuously to-
mentose, the veins concealed by tomentum,
88
FIELDIANA: BOTANY
densely prominulose-reticulate. Capitulescences
of glomerules of 2-5 heads, terminal on branch-
lets. Capitula sessile; involucres campanulate, 8-
10 mm long, 4-10 mm wide; phyllaries ca.
4-seriate, ca. 14, the outer ovate, 2-2.5 mm long,
1.5-2.5 mm wide, apically acute, stiffly mucro-
nate, tomentose, the inner oblong-elliptic, 7-7.5
mm long, mucronulate, glandular-puberulent,
densely ciliolate, brownish or purplish; florets 5-
6, the corollas bilabiate, white, glabrous, ca. 7.5
mm long; anthers ca. 5 mm long. Achenes oblong,
ca. 3 mm long, 5 -angled, stipitate -glandular,
densely hispidulous; pappus of subplumose to bar-
bellate bristles, ca. 7 mm long.
Proustia cuneata most closely resembles P. ber-
berdifolius but is distinguished from that species
by its generally larger, abaxially tomentose leaves
and glomerulate clusters of two to five heads.
The new name provided by Cabrera (1953),
Lophopappus blakei, was necessary only when
Blake's Proustia cuneata was placed in Lophopap-
pus, where the name was preoccupied by L. cu-
neatus R. E. Fries ( 1 960), a Bolivian and Argentine
species. No nomen novum is here provided for
the latter species.
CUSCO. Urubamba: Ollantaitambo, Pennell
13664 (F, us).
3. Proustia cuneifolia D. Don, Trans. Linn. Soc.
London 16: 202. 1830. TYPE: Chile, Co-
quimbo, Calddeugh s.n. (holotype, P, not
seen). Figure 18.
Proustia pungens Poepp. ex Less., Syn. Comp. 110.
1832. TYPE: Chile, Aconcagua, Poeppigs.n. [884]
(holotype, B, presumably destroyed; photograph
ex B, FM neg. 20730).
Intricately branched shrubs, 1-2 m tall, the
branches flexuose, the branchlets 1-8 cm long, the
apex acicular or spinulose, striate, nodose, lacking
nodal spines, finely pubescent, glabrescent. Leaves
subsessile or petiolate; petioles 2-5 mm long; blades
lanceolate to oblong-lanceolate or spathulate,
(12-)40-90 mm long, 4-22 mm wide, coriaceous,
the abaxial surfaces prominulose-reticulate, the
margins entire or slightly repand-denticulate api-
cally. Capitulescences of racemose, glomerulate
spikes. Capitula sessile; involucres campanulate,
7-8 mm high, 4-5 mm wide; phyllaries 14, 5-6-
seriate, the outer ovate, ca. 1 mm long, ca. 1 mm
wide, coriaceous, apically obtuse, the middle ovate-
lanceolate, 4-6 mm long, 2-2.5 mm wide, apically
1cm
i | O.Scm
FIG. 18. Proustia cuneifolia. 1, habit; 2, capitulum;
3, floret with achene. (Drawn from C. Vargas C. 740,
F.)
obtuse, the inner oblong to lanceolate, 6-9 mm
long, 2-2.2 mm wide, apically rounded, ciliolate;
florets 5-6, the corollas violaceous to purple, bi-
labiate, 10-15 mm long, the outer lip tridentate,
5-6 mm long, the inner lip bipartite, 5-5.5 mm
long, coiled; anthers 4.5-6 mm long. Achenes tur-
binate, 3-4 mm long, 5-costate, glandular-pubes-
cent; pappus of barbellate bristles, 9-12 mm long,
tawny to violaceous.
DISTRIBUTION— Known from the intermontane
region of southern Peru (3100-3600 m) and dis-
junct populations in the lomas formations along
the southern coast in the Department of Arequipa
(200-700 m).
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
89
Proustia cuneifolia most closely resembles P. pe-
ruviana but is unique among the members of the
genus in Peru in that it possesses spinose branch-
lets. Fabris (1968) proposed several formas based
on leaf form and pubescence, and Peruvian ma-
terial was referred to forma cuneifolia and forma
oblongifolia. An examination of an isotype of D.
Don's P. oblongifolia (Ruiz Lopez & Pavon s.n.,F
639998) shows it to be conspecific with Proustia
pyrifolia Lag. ex DC., a Chilean species. No sub-
specific categories are recognized in the current
study. It is known by the common names chamo,
tantar, huanchinco, and huacata.
APURIMAC. Abancay: entre Abancay y Curahuasi,
Ferreyra 9819 (MO, USM); Cusahuasi, Vargas 740 (F).
Aimaraes: 1 5 km NW of Chalhuanca, Gentry et al. 23340
(F, MO). No exact locality: Weberbauer 5762 (F). ARE-
QUIPA. Cailloma: Valle del Colca, Treacy 723 (F). Car-
aveli: Lomas de Atiquipa, near Chala, Ferreyra 8894
(USM), 72727 (USM), 73525 (MO, USM), 73995 (MO, USM),
Ono 1 10104 (USM); Lomas de Okopa, near Chala, Fer-
reyra 11472 (USM); Lomas de Taimara, near Atiquipa,
Ferreyra 13939 (MO, USM). AYACUCHO. La Mar: Tam-
bo Cuchicancha, Madison 10399-70 (USM). Lucanas: be-
tween Lucanas and Puquio, Ferreyra 9773 (MO, USM).
CUSCO. Calca: near Calca, Ferreyra 9981 (USM). Uru-
bamba: Caicay, Herrera 2580 (F); Kaira, Vargas 6747
(F); Salapuncu, Far#asS779(us);Tarapata, Vargas 11076
(F).
4. Proustia foliosa (Rusby) Ferreyra, comb. nov.
Lophopappus foliosus Rusby, Bull. Torrey Bot. Club
21: 487, pi. 225. 1894. SYNTYPES: Bolivia, La
Paz, M. Bang 66 (NY); Bolivia, Chuguiaguilla, M.
Bang 791 (NY).
Much-branched shrubs, the branches glabrous,
erect, striate. Leaves sessile; blades linear, rarely
narrow oblanceolate, 7-16 mm long, 0.8-1 mm
wide, aristate, the adaxial surfaces canescent, the
abaxial surfaces glandular-muricate, apically at-
tenuate-apiculate. Capitulescences of solitary, ter-
minal heads. Capitula sessile; involucres cylindric
to campanulate, 1 1-14 mm long, 4-5 mm wide,
subtended by linear, glandular bracts; phyllaries
12, 4-5-seriate, the outer lance-ovate, 5-7 mm
long, 2-2.5 mm wide, apically attenuate, the inner
linear-lanceolate, 9-10 mm long, 1.6-1.8 mm wide,
apically apiculate; florets 4-5(-8), the corollas 9-
10 mm long, bilabiate, the outer lobe tridentate,
the inner lip bipartite, coiled, 3-3.5 mm long, the
tube 5-6 mm long; anthers 4—4.5 mm long. Achenes
cylindric, 3-4 mm long, 0.6-0.8 mm wide, his-
pidulose-pubescent, 5-costate; pappus of plumose
bristles, 7-9 mm long.
DISTRIBUTION— Known from Andean Cordil-
lera of southern Peru to Bolivia (3300-3750 m).
Proustia foliosa is distinguished by its linear,
aristate leaves and linear, apiculate phyllaries.
AREQUIPA. Arequipa: Chivay, Weberbauer 6888 (F,
LP, us, USM). Cailloma: Valle del Colca, Treacy 654 (F).
5. Proustia peruviana (Cabr.) Ferreyra, comb. nov.
Lophopappus peruvianusCabr., Bol. Soc. Argent. Bot.
5: 50. 1953. TYPE: Peru, Ancash, Bolognesi, cer-
ca de Aquia, E. Cerrate 1552 (holotype, LP; iso-
type, USM).
Much-branched shrubs, 1.5-2 m high, the upper
branches striate, glabrescent. Leaves sessile or sub-
sessile; blades ovate-elliptic, 10-20 mm long, 7-
9 mm wide, coriaceous, both surfaces glabrous,
basally cuneate, apically acute to obtuse, mucro-
nate, the abaxial surfaces prominulose-reticulate,
the margins entire to finely denticulate. Capitu-
lescences of solitary heads, occasionally glomer-
ulate at branch tips with 2-3 heads. Capitula ses-
sile; involucres cylindric, 9-11 mm high, 5-6 mm
wide; phyllaries 18-20, 4-5-seriate, the inner lan-
ceolate, 8-9.5 mm long, 2-2.2 mm wide, apex
brown, mucronate, the outer ovate, apex deep
brown, shortly bipartite, 5-7 mm long, 2.6-3 mm
wide; florets 14-18, the corollas pink to whitish,
10-12 mm long, bilabiate, the outer lip tridentate,
coiled, 6-6.5 mm long, the inner lip bifid, 5-5.5
mm long, coiled, the tube 4.5-5 mm long; anthers
4-5 mm long. Achenes cylindric, 3.8-4 mm long,
0.5-0.8 mm wide, glandular- puberulent, glabres-
cent, 5-costate; pappus of barbellate bristles, 7-9
mm long.
DISTRIBUTION— Known from the Andean Cor-
dillera of northern to central Peru (3200-3500 m).
Proustia peruviana most closely resembles P.
berberidifolius from which it differs in having typ-
ically solitary capitula with more florets and larger
leaves with entire margins.
ANCASH. Bolognesi: near Chiquian, Cerrate 2380
(USM); S of Huaraz, Gentry et al. 37470 (F, MO). AYA-
CUCHO. Lucanas: between Puquio and Abancay, El-
lenberg 4956 (us). CAJAMARCA. Contumaza: Guz-
mango, Sagdstegui et al. 9690 (F, MO). HUANUCO.
Huamalies: near Llata, Macbride & Featherstone 2296
(F, us). LA LIBERTAD. Santiago de Chuco: Santiago
de Chuco, Sagdstegui et al. 1 1760 (F); Hacienda Angas-
marca, Weberbauer 7206 (F, us). PIURA. Huancabam-
ba: Rumitana, Lopez et al. 8849 (F, MO).
90
FIELDIANA: BOTANY
XX. TRICHOCLINE
Trichocline Cass., Bull. Sci. Soc. Philom. Paris
1817. 13. 1817. TYPE: Doronicum incanum
Lam. = Trichocline incana (Lam.) Cass.
Acaulescent or caulescent, rhizomatous, peren-
nial herbs. Leaves sessile or petiolate, generally in
a basal rosette, or if shortly cauline, alternate; blades
simple, the adaxial surfaces tomentose or glabres-
cent, the abaxial surfaces usually tomentose or la-
nate, the margins entire, dentate, crenate, lobed,
or lyrate-pinnatifid. Capitulescences of solitary,
terminal heads; peduncles erect or ascending,
bracteate or ebracteate, hollow or solid. Capitula
heterogamous; involucres hemispherical or cam-
panulate; receptacles plane, naked or alveolate;
phyllaries imbricate, 2-6-seriate, subequal or the
outer gradually shorter, often tomentose or lanate;
ray florets pistillate, the corollas yellow or orange
to red, violet or rarely white, bilabiate, the outer
lip ligulate, tridentate, the inner lip bifid, the lobes
filiform; staminodia present; styles lobed, the
branches glabrous or pilose; disc florets hermaph-
roditic, the corollas bilabiate, the outer lip triden-
tate, the inner lip bifid; anthers basally sagittate;
style filiform, the branches apically obtuse or sub-
acute, papillose to pilose. Achenes cylindric, tur-
binate, ovoid, obovoid, or elliptic, apically trun-
cate or with a short neck, pubescent with both
swollen or inflated, biseriate trichomes, and mul-
ticellular, glandular trichomes; pappus of barbel-
late bristles, white.
Trichocline consists of approximately 25 species
primarily restricted to austral South America and
one native to Australia. In Peru, two species are
currently recognized. The genus is distinguished
from Chaptalia in possessing larger ray florets with
staminodia and nonrostrate achenes, pubescent
with both swollen or inflated hairs and multicel-
lular, glandular trichomes.
The placement of several species has been con-
troversial. For example, Hieronymus (1896), and
later Burkart (1944), suggested that two species,
originally placed in Chaptalia (C. ovalis and C.
oblonga), be moved to Trichocline. Zardini (1975)
argued that these species were misplaced in Trich-
ocline and returned to Chaptalia; that placement
is adopted here. Alternatively, Zardini (1974)
moved Trichocline hieracioides to Gerbera; how-
ever, Hansen (1990, 1991) returned it to Tricho-
cline; that placement is adopted here.
References
BURKART, A. 1 944. Estudio del genero de Com-
puestas Chaptalia con especial referencia a la
especies argentinas. Darwiniana, 6: 505-594.
HANSEN, H. V. 1 990. Phylogenetic studies in the
Gerbera-complex (Compositae, tribe Mutisieae,
subtribe Mutisiinae). Nord. J. Bot, 9: 469-485.
. 1991. Phylogenetic studies in Compos-
itae tribe Mutisieae. Opera Botanica, 109: 5-50.
HIERONYMUS, G. 1896. Plantae Stuebelianae no-
vae. Bot. Jahrb. Syst., 21: 325-377.
ROBINSON, H. 1988. A new species of Trichocline
from northern Peru. Phytologia, 65: 47-49.
ZARDINI, E. M. 1974. Sobre la presencia del ge-
nero Gerbera en America. Bol. Soc. Argent. Bot.,
16: 105.
. 1975. Revision del genero Trichocline
(Compositae). Darwiniana, 19: 618-733.
Key to Peruvian Species of Trichocline
1 . Leaf blades 1 5-50 mm long, 5- 1 4 mm wide, the abaxial and adaxial surfaces densely white-tomentose,
the margins irregularly crenate, serrate, or dentate; peduncles 2-6 cm long; involucres 5-10 mm high;
ray corollas yellow, the outer lip 5-7 mm long (southern Peru) T. caulescens
1. Leaf blades 50-120 mm long, 15-45 mm wide, the abaxial surface sparsely to densely tomentose,
the margins crenate to lyrate-pinnatifid; peduncles 8-30(-48) cm long; involucres 1 7-30 mm high;
ray corollas pink, violet, or purple, the outer lip 1 5-30 mm long (northern Peru) T. peruviana
Trichocline caulescens Phil., Anal. Mus. Nac.
Chile, Bot. 8:31.1891. TYPE: Chile, Iquique,
"Medanos de Pica," Mar 1885, F. Philippi
2069 (holotype, SGO, not seen; photograph ex
B, FM neg. 16013). Figure 19.
Trichocline stuebelii Hieron., Bot. Jahrb. Syst. 2 1 : 369.
1896. TYPE: Chile, Parinacota, "crescit inter Ta-
cora et Tomarape," 4200-4400 m, Oct, A. Stiibel
Wla (holotype, B, presumably destroyed; pho-
tograph ex B, FM neg. 16025).
Perennial herbs, essentially acaulescent, 6-12
cm tall, rhizomes vertical, 1-10 mm in diameter,
lignescent. Leaves basally rosulate; blades oblan-
ceolate to spathulate in outline, 1 5-50 mm long,
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
91
FIG. 19. Trichodine caulescens. 1, habit; 2, ray floret; 3, disc floret. (Drawn from A. Weberbauer 7383, F.)
5-14 mm wide, the abaxial and adaxial surfaces
densely white-tomentose, the margins irregularly
crenate, serrate, or dentate. Capitulescences of sol-
itary, terminal heads; peduncles erect to procum-
bent, 2-6 cm long, densely albo-tomentose, brac-
teate, the bracts linear, 3-5 mm long. Capitula
with involucres hemispherical, 5-10 mm high, 10-
25 mm wide; phyllaries 2-3 seriate, densely albo-
tomentose, the outer linear, ca. 3 mm long, 0.5
mm wide, the inner lanceolate, 6-7 mm long, 1.5-
2 mm wide; ray florets 12-14, the corollas yellow,
the outer lip 5-7 mm long, 1.5-2 mm wide; disc
florets 15-25, the corollas yellow, 6-7 mm long,
the outer lip ca. 1 mm long, 3-toothed, the inner
lip ca. 1 mm long, bifid. Achenes ellipsoid, ca. 4
mm long, ca. 1.5 mm wide; pappus 4-6 mm long,
white.
Trichodine caulescens is rare in southern Peru
but more common in northern Chile (1850-3500
m). It is distinctive among the Peruvian species
with densely white-tomentose leaves and yellow-
ish ray corollas.
PERU. Tacna: Candarave, Weberbauer 7383 (F, OH,
us).
2. Trichodine peruviana Hieron., Bot. Jahrb. Syst.
21: 368. 1896. TYPE: Peru, "crescit infra La
Cruz de Celendin et prope Tambo Mayo inter
Pacasmayo et Moyobamba," A. Stiibel 481
(holotype, B, presumably destroyed; photo-
graph ex B, FM neg. 16023).
Onoseris hieracioides Kunth, Nov. Gen. Sp. 4: 7. t.
304. 1820. TYPE: Ecuador, Chimborazo, Hum-
boldt & Bonplands.n. (holotype, p, not seen; pho-
tograph ex P, FM neg. 38102).
Trichodine hieracioides (Kunth) Ferreyra, Journ. Arb.
25: 344. 1944. Non Baker, 1884.
Gerbera hieracioides (Kunth) Zardini, Bol. Soc. Ar-
gent. Bot. 16: 105. 1974.
Trichodine beckeri H. Robinson, Phytologia 65: 47.
1988. TYPE: Peru, Cajamarca, surroundings of
Cajamarca, 2700-3700 m, 30 May 1 986, B. Beck-
er & F. M. Terr ones H. 1305 (holotype, us
3092896; isotype, PPEA).
Perennial herbs, acaulescent; rhizomes erect, to
ca. 4 mm in diameter. Leaves basally rosulate;
92
FIELDIANA: BOTANY
petioles 10-120 mm long; blades oblanceolate to
spathulate or lyrate, 50-120 mm long, 15-70 mm
wide, acute to obtuse, the adaxial surfaces gla-
brous, the abaxial surfaces white-tomentose, the
margins entire, weakly serrate, or lyrate-pinnati-
fid. Capitulescences of solitary, terminal heads;
peduncles erect, tomentose, 1 4-70 cm long, brac-
teate, the bracts linear, 5-20 mm long, 1-2 mm
wide. Capitula with involucres campanulate, 1 7-
30 mm high, 1 5-25 mm wide; phyllaries 4-5-se-
riate, the outer linear, 5-12 mm long, ca. 1 mm
wide, acuminate, lanuginose, the margins purple,
the inner 16-30 mm long, 1.5-2.5 mm wide; ray
florets 14-24, pistillate, bilobed, the tube 5-12 mm
long, 0.5-1 mm wide, the outer lip ligulate, tri-
dentate, 15-30 mm long, 2-3 mm wide, the ad-
axial surfaces pink, violet, or purple, the abaxial
surfaces dark pink, the inner lip bifid, the lobes
filiform, ca. 4 mm long, coiled; disc florets 1 5-26,
the corollas white, violet, or pink, 13-15 mm long,
the tube 4-5 mm long, the limb 8-9 mm long;
anthers 6-8 mm long; style branches linear, ca. 1
mm long, papillose. Achenes cylindrical to fusi-
form, (3.5-)4-7 mm long, 1.5-1.8 mm wide; pap-
pus 12-20 mm long.
DISTRIBUTION— Known from the Andean Cor-
dillera of southern Ecuador to north-central Peru
(1500-3 800m).
Trichocline peruviana is a common element in
the intermontane, subtropical region of the Pe-
ruvian Andes. Representative collections show
much variability in the leaf size, amount of pu-
bescence, and ray floret coloration. Examples with
strongly lyrate leaf blades superficially resemble
Onoseris speciosa, but they are readily distin-
guished by the species' blue ray florets and often
multiheaded, branched peduncles.
Trichocline peruviana possesses characteristics
of both Gerbera and Trichocine and, until recently,
the collections cited here were treated in Gerbera
(Zardini, 1974) under the name G. hieracioides
(Kunth) Ferreyra. Hansen (1990, 1991) suggested
that this species be removed from Gerbera and
tentatively placed in Trichocline. Because the name
Trichocline hieracioides is preoccupied by Baker's
Brazilian species, T. peruviana was adopted as the
next available name.
After examining a suite of specimens from
throughout its range, including the type locality of
T. peruviana and T. beckeri, it is obvious that these
two species are conspecific. The smaller, strongly
lyrate leaves found in collections from Cajamarca
to Ancash and Huanuco are but the end of a range
of variation in leaf size and degree of dentation.
Additional collections (e.g., Dillon et al. 6202) from
the type locality of T. beckeri exhibit a wide range
of leaf size and lobing while maintaining rather
constant capitular size and floral morphology.
AMAZONAS. Chachapoyas: Cerros Calla-Calla, 8 km
above Leimebamba on road to Balsas, Hutchison &
Wright 5653 (F, uc, us, USM); 1 km SW of Chachapoyas,
Wurdack 470 (us). ANCASH. Bolognesi: Chiquian, Fer-
reyra 5794 (us). Corongo: Nueva Victoria, Mostacero et
al. 1999 (F). Huari: between Pichiu and Recuay, We-
berbauer 2943 (USM); near Recuay, Ferreyra 14556 (USM).
Yungay: Llanganuco, above Yungay, Ferreyra 14341
(USM). CAJAMARCA. Cajamarca: Rio Manzana, Beck-
er & Terrenes 1916 (PPEA, us); 15 km NW of Cajamarca,
road to Chamis, Dillon et al. 6202 (CPUN, F); San Pablo,
Hutchison & Wright 5041 (uc, us, USM); Chiquiden to
Cajamarca, Ochoa 1488 (USM); La Encanada-Jalca de
Kumulca, Sagdstegui et al. 8081 (F); Hacienda Pollo-
quito, Sanchez 298 (CPUN, F). Celendin: 4 km NNE of
Celendin on road to Balsas, Edwin & Schunke 3588 (F);
arriba de Celendin, entre Celendin y Balsas, Ferreyra
15055 (us, USM); on road to Celendin, 31 km E of Ca-
jamarca, Hutchison & Wright 5109 (F, us), 5132 (uc,
USM); 2 km E of Celendin on road to Balsas, Hutchison
& Wright 5153 (F, uc, us, USM); Canyon of Rio Maranon
above Balsas, 3-4 km below summit of the road to Ce-
lendin, Hutchison & Wright 5218 (uc, us, USM); vicinity
of Celendin, Woytkowski 233 (F). Chota: between Chota
and Bambamarca, Ferreyra 8496 (USM), Hutchison &
Wrigth 5229 (uc); Llama to Chongoyape, Ferreyra 0869
(USM); Cerro El Leonero, arriba de Chuyubamba, Sa-
gdstegui 14062 (F). Contumaza: Contumaza-Cascabam-
ba, Sagdstegui et al. 9974 (F); Guzmango, Sagdstegui &
Lopez 10573 (F); Bosque Cachil, Sanchez 3155 (F). Cu-
tervo: near Cutervo, Ferreyra & Acleto 15362 (USM).
HUANUCO. Huamalies: Llata, Macbride & Feather-
stone 2233 (F). LA LIBERT AD. Otusco: Agallpampa,
Lopez 479 (us, USM); above Huaranchal, Lopez 1349
(USM). PIURA. Huancabamba: camino Suruguna, Acleto
495 (USM); carretera entre Canchaque y Huancabamba,
Diaz & Baldeon 2391 (F); ca. 15 km NE of Canchaque,
Dillon & Sanchez 6244 (CPUN, F); Abra de Porculla, etre
Olmos y Jaen, Ferreyra 13756 (USM), 75666 (us, USM);
Mitopampa, Sagdstegui et al. 8230 (F); Ayabaca, Soukup
4352 (us); El Tambo, Soukup 4644 (USM); between
Huancabamba and Ayavaca, Weberbauer 6336 (F, us).
XXI. TRIXIS
Trixis P. Br., Civ. nat. hist. Jamaica 312. 1756.
TYPE: Trixis inula Crantz
Perennial herbs, shrubs, or, occasionally, scan-
dent, climbing. Leaves alternate, simple, sessile to
petiolate; blades linear to lanceolate, elliptic, ovate
or obovate, the margins entire to denticulate. Ca-
pitulescences paniculate or corymbose, terminal,
rarely solitary. Capitula radiate; involucres cam-
panulate to cylindric, usually subtended by (l-)3-
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
93
5 (-12) accessory bracts; receptacles plane, alveo-
late, pilose; phyllaries 5-13(-16), 1-3-seriate; flo-
rets 4-30(-60), isomorphic, hermaphroditic, the
corollas yellow, yellow-orange, or whitish-yellow,
often maturing white, bilabiate, the outer lip el-
liptic or oblong, tridentate, the inner lip bifid; sta-
mens 5, the anthers with oblong apical append-
ages, basally caudate; style branches apically trun-
cate, penicillate, the adaxial surfaces papillose, the
abaxial surfaces glabrous. Achenes cylindric to fu-
siform, 5-ribbed, usually pubescent with short,
biseriate, myxogenic trichomes, and/or uniseriate,
glandular trichomes; pappus of 50-150, hispidu-
lous bristles, (2-)3-4-seriate, isomorphic, tawny to
white.
Trixis contains 50-60 species ranging from
southern United States to southern South Amer-
ica. In Peru, the genus is distributed in a wide
variety of habitats from coastal desert to montane
forests.
References
ANDERSON, C. 1972. A monograph of the Mex-
ican and Central American species of Trixis
(Compositae). Mem. New York Bot. Garden,
22: 1-68.
CABRERA, A.L. 1936. Las especies argentinas y
uruguayas del genero Trixis. Rev. Mus. La Pla-
ta, Secc. Bot., 1: 31-86.
DlAZ-PlEDRAHITA, S., AND C. VEZEZ-NAUER. 1991.
Nueva especie de Trixis (Asteraceae) de Colom-
bia. Rev. Acad. Colomb., 18: 149-151.
HARLING, G. 1991.190(10). Compositate-Mu-
tisieae, pp. 76-80. In Harling, G., & L. An-
dersson, eds., Flora of Ecuador, No. 42, Swedish
Natural Science Research Council, Stockholm.
LOJA, B. H. A. 1969. Revision de las especies
Peruanas del genero Trixis (Compositae). Publ.
Mus. Hist. Nat. "Javier Prado," Ser. B, Bot.,
25: 1-36.
Key to Peruvian Species of Trixis
1. Phyllaries 1 -seriate 2
2. Leaves tomentose T. subparadoxa
2. Leaves hirsute 3
3. Capitula subtended by foliose bracts T. neaeana
3. Capitula not subtended by foliose bracts T cacalioides
1 . Phyllaries 2- or more-seriate 4
4. Leaves acuminate; capitula with 10-12 florets, white T. divaricata
4. Leaves acute; capitula with 6 florets, yellow 5
5. Blades oblanceolate, to 15 cm long; phyllaries 14-16, 3—4 seriate T. sagasteguii
5. Blades oblong-lanceolate, to 6 cm long; phyllaries 7-8, 3-seriate T. churinensis
1 . Trixis cacalioides (Kunth) D. Don., Trans. Linn.
Soc. London 16: 187. 1830.
Perdicium cacalioides Kunth, Nov. Gen. Sp. 4: (fol.)
121. 1818; (quatro) 154. 1820. TYPE: Peru, Ca-
jamarca, Jaen, "in declivitate orientalis Andium
Peruvianorum ad ripam fluvii Matara, 430 hex,"
Humboldt & Bonplands.n. (holotype, p, not seen).
Trixis paradoxa Cass, Opusc. phytol. 2: 160. 1826.
TYPE: Peru, /. Dombeys.n. (holotype, p, not seen).
Trixis hexantha S. Moore, J. Bot. 52: 264. 1914. TYPE:
Peru, "valley between Pasasmayo and rail-head,
7000 ft," H. O. Forbes s.n. (holotype, BM, not
seen).
Shrubs 8-12 dm high, branched, branches stri-
ate, pubescent. Leaves subsessile; blades lanceo-
late to oblanceolate, 10-80 mm long, 4-30 mm
wide, bases attenuate to cuneate, apically acute,
conspicuously pubescent, hairs strigose, glandular,
the margins subentire to denticulate, mucronate.
Capitulescences corymbose, the peduncles short
with 3 bracteoles. Capitula with involucres nar-
rowly campanulate, phyllaries 5, 1 -seriate, obtuse,
rarely bidentate; florets 6-7(-10), the corollas yel-
low, 9-12 mm long, the outer lip 3-5 mm long,
pubescent, tridentate, the inner lip bifid, laciniate,
coiled. Achenes 2-6 mm long, 0.4-1 mm wide,
pubescent with glandular trichomes; pappus 7-12
mm long, tawny.
DISTRIBUTION— Known from coastal and adja-
cent montane habitats of southern Ecuador, Peru,
and Chile. In Peru, it is well represented in the
coastal lomas formation, with occasional individ-
uals occurring at higher elevations along the west-
94
FIELDIANA: BOTANY
ern slope of the Cordillera Occidental (100-2500
m).
Trixis cacalioides is characterized by its highly
variable, strigose and glandular leaves with entire
to denticulate margins and the absence of enlarged
subtending bracts at the base of the involucre.
There has been some disagreement as to the
correct name for this taxon, and both T. caca-
lioides and T. paradoxa Cass. have been applied
to Peruvian material. The type material of T. ca-
calioides was gathered near Matara (Prov. Jaen,
Dept. Cajamarca). The type material of T. para-
doxa was a Peruvian collection by Dombey, and
the name has been applied to both Peruvian and
Ecuadorian collections (Harling, 1991) with more
denticulate leaf blades with strigose pubescence.
After examining a large suite of collections from
varied ecological situations throughout Peru and
Ecuador, it appears that the presence of teeth and
pubescence of the leaves are variable. Therefore,
the two are considered conspecific, with Kunth's
T. cacalioides having priority.
ANCASH. Santa: Lomas de Lupin, Cerrate 814 (USM);
between Barranca and Huarmey, Ferreyra 13789 (USM).
AREQUIPA: Caraveli: Lomas de Capacc, Ferreyra 7230
(USM); Lomas de Okopa, near Atiquipa, Ridoutt 14522
(USM). Arequipa: slopes of Misti, Ferreyra 14272 (USM),
Pennell 13157 (USM). CAJAMARCA. Chota: Carhua-
quero, Ferreyra 3168 (USM). Contumaza: near Contu-
maza, Ferreyra 8597 (USM). ICA. lea: Santa Cruz, Fer-
reyra 13460 (USM); Los Molinos, Angulo 2421 (HUT).
LIMA. Cajatambo: Churin, Ferreyra 5351 (USM), 3522,
(USM), 6757 (USM). Canta: Quives, Araguay, Pennell 14297
(NY), Dreyfus 12754 (USM), Ferreyra 11781 (USM). Chan-
cay: Lachay, Ferreyra 9631 (USM), 9694 (USM), Macbride
& Featherstone 550 (GH, us); Magdalena, Ridoutt 10290
(USM); San Bartolo, Ferreyra 8214 (USM); near Trapiche,
Ferreyra 9668 (USM). Lima: Atocongo, Maisch 13815
(USM), Pennell 14766 (NY), Ferreyra 2058 (USM), 3471
(USM), 12451 (USM), Cerrate 3020 (USM); Cajamarquilla,
Ferreyra 2851 (USM), 8347 (USM), 9755 (USM), Ridoutt
12219 (USM); near Chosica, Ferreyra 522 (USM), 6720
(USM). Huarochiri: above Surco, Ferreyra 6055 (USM);
km 57 Lima to Oroya, Ferreyra 3484 (USM); Cocachacra,
Ridoutt 12484 (USM); Santa Eulalia, Ridoutt 13037 (USM);
near Surco, Ferreyra 5330 (USM). PIURA. Piura: near
Piura, Haught s.n. (GH).
2. Trixis churinensis Herrera, Publ. Mus. Hist.
Nat. "Javier Prado," Ser. B, Bot. 25: 14. 1969.
TYPE: Peru, Lima, Cajatambo, Churin, 2100-
2200 m, R. Ferreyra 6151 (holotype, USM
15728).
Shrubs 1.5-2 m high, branched, the branches
striate, pubescent. Leaves subsessile; blades lan-
ceolate to oblong-lanceolate, 20-60 mm long, 8-
22 mm wide, the adaxial surfaces glabrous, the
abaxial surfaces pubescent, the margins entire.
Capitulescences cymose. Capitula subsessile; in-
volucres cylindric; phyllaries 7-8, 3 -seriate, 2 out-
er, 3 middle, 2-3 inner; florets 5, the corollas yel-
lowish, the tube 5.8-6.2 mm long, the outer lip
tridentate, 4-nerved, the inner lip bifid, laciniate,
coiled. Achenes 3.5—4.5 mm long, densely papil-
lose; pappus ca. 8.5 mm long, yellowish.
DISTRIBUTION— Known only from the type lo-
cality.
Trixis churinensis most closely resembles T. sa-
gasteguii, from which it differs in having a gla-
brescent habit and 7-8 phyllaries.
3. Trixis divaricata (Kunth) Spreng., Syst. Veg. 3:
501. 1826.
Perdicium divaricatum Kunth, Nov. Gen. Sp. [quarto]
4: 155. 1820, [fol.] 4: 121. 1818. TYPE: Peru,
Cajamarca, "prope Cavico ad ripam fluvii Cha-
maya, Jaen de Bracamoros," Humboldt & Bon-
pland s.n. (holotype, p, not seen; photograph ex
p, FM neg. 37859).
T. antimenorrhoea (Schrank) Mart, ex Baker, Mart.
Fl. Br. 6(3): 385. 1884.
Prionanthes antimenorrhoea Schrank, PI. Rar. Hort.
Acad. Monac. 2(6): 51. 1820. LECTOTYPE (des-
ignated by Harling, 1991): Brazil, no specimen
known, 5 1 chosen as lectotype, not seen.
Shrubs 2-2.5 m high, erect to procumbent; stems
cylindric, striate, scarcely pubescent to tomentose.
Leaves subsessile; blades lanceolate to elliptic-lan-
ceolate, 40-170 mm long. 10-50 mm wide, pu-
bescent, acuminate, the margins entire to slightly
dentate. Capitulescences paniculate-corymbose;
peduncles 6-10 mm long, pubescent, 1-3 bracte-
ate, the bracts lanceolate, pubescent. Capitula with
involucres campanulate to turbinate; phyllaries
2-seriate, the outer 4—5, unequal, the inner ca. 8,
equal, acute; florets 10-12, the corollas white or
yellowish, the tube 5-9 mm long, the outer lip
4-nerved, tridentate, the inner lip bifid, laciniate,
coiled. Achenes 2.8-6 mm long, 0.5-1 mm wide,
pubescent with uniseriate glandular trichomes;
pappus ca. 1 1 mm long, tawny.
DISTRIBUTION— Known from the Neotropics
from Costa Rica and Hispaniola to Argentina. In
Peru, it is largely restricted to the eastern Andean
Cordillera (1800-3000 m).
Trixis divaricata is easily distinguished by its
auriculate, clasping leaf bases, abaxial arachnoid-
tomentum, and flexuose, divaricately branched
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
95
capitulescences, Cabrera (1936) treated Peruvian
material under T. antimenorrhoea var. auriculata
(Hook.) Cabr., and Marling (1991) and Diaz-Pied-
rahita and Velez-Nauer (1991) followed by ac-
cepting T. antimenorrhoea as the valid name.
However, Kunth's name T. divaricata, should have
priority by several months and, pending detailed
studies, T. antimenorrhoea will be restricted to
austral South America.
APURIMAC. Apurimac: Rio Chalhuanca, Angulo
1741 (HUT). CUSCO. Convencion: Santa Ana Valley,
Herrera 3617 (cuz). Paucartambo: Patria, Mann 1708
(USM). HUANCAVELICA. Tayacaja: below Surobam-
ba, Tovar 3744 (USM); below Andamarca, Tovar 1851
(USM). HUANUCO. Estacion Pozuzo, Cueva Grande,
Macbride 4784 (us); Yamano, Chacalla, Macbride 3793
(us). JUNIN. Tarma: Palca, Soukup 3492 (USM). PIU-
RA. Huancabamba: Canchaque, Calle 1003 (USM). SAN
MARTIN. Lamas: near Lamas, Ferreyra 7807 (USM).
Mariscal Caceres: Chambira, near Juanjui, Ferreyra 4574
(USM). San Martin: Tarapoto, Ferreyra 7852 (USM); Pu-
cayacu, Ferreyra 7738 (USM). Saposoa: near Saposoa,
Ferreyra 4620 (USM); between Bellavista and Tingo de
Saposa, Ferreyra 4766 (USM); near Banos, Ferreyra 4729
(USM).
4. Trixis neaeana DC., Prodr. 7: 67. 1838. TYPE:
Peru, Nee s.n. (holotype, p, not seen).
Shrubs to 1 m high, the stems striate. Leaves
subsessile; blades oblong-lanceolate, 35-60 mm
long, 15-25 mm wide, the adaxial surfaces gla-
brescent, the abaxial surfaces densely pubescent,
apically acute, the margins entire. Capitulescences
of panicles or corymbose with 2-3 leaves basal.
Capitula with involucres cylindric; phyllaries 5,
1 -seriate, oblong, obtuse or acute; florets 5-6, the
corollas yellow, bilabiate, the tube 7-8 mm long,
the outer lip 2-nerved, tridentate, the inner lip
bifid. Achenes 4-5 mm long, 1-1.2 mm wide, pu-
bescent with uniseriate glandular trichomes; pap-
pus 9-10 mm long, yellowish.
DISTRIBUTION— Known only from intermon-
tane valleys of the north-central Peruvian Andes
(2000-2400 m).
Trixis neaeana resembles T. cacalioides, but it
differs in its two to three foliose bracts subtending
the involucres.
ANCASH. Huaylas: near Canon del Pato, Ferreyra
14585 (USM); road to Huallanca, Luna 85 (USM).
5. Trixis sagasteguii Cabr., Bol. Soc. Argent. Bot.
10: 42. 1962. TYPE: Peru, La Libertad, Bo-
livar, Catal, route Maranon-Longotea, 200 m,
A. Lopez M. & A. Sagdstegui A. 3163 (holo-
type, LP, not seen; isotype, HUT). Figure 20.
Shrubs, branched, the stems pubescent. Leaves
subsessile, blades oblanceolate to lanceolate, 90-
150 mm long, 25-40 mm wide, pubescent, acute,
the margins entire or denticulate. Capitulescences
of multiheaded, terminal panicles, shortly pedun-
culate. Capitula with involucres cylindric; phyl-
laries 14-16, 3—4 seriate, lanceolate, acute, ciliate;
florets 5, the corollas yellow, the tube 4-4.5 mm
long, the outer lip tridentate, the inner lip bifid,
laciniate, to 3 mm long. Achenes ca. 5 mm long,
pubescent with biseriate trichomes; pappus 6-7
mm long, yellowish-white.
DISTRIBUTION— Known from extreme southern
Ecuador and northern Peru. Trixis sagasteguii is
distinguished by its capitulescences of large, tri-
angular panicles, large sericeous leaves, and three-
to four-seriate, densely pubescent involucres.
AMAZONAS. Bongara: Rio Utcubamba, 18-19 km
below Gaelic, Hutchison & Wright 5864 (F).
6. Trixis subparadoxa Herrera, Publ. Mus. Hist.
Nat. "Javier Prado," Ser. B, Bot. 25: 6. 1969.
TYPE: Peru, Cajamarca, Celendin: entre Bal-
sas y Hacienda El Limon, R. Ferreyra 15059
(holotype, USM 15729).
Shrubs branched, branches pubescent. Leaves
subsessile; blades elliptic-lanceolate, 50-105 mm
long, 10-30 mm wide, the abaxial surfaces densely
strigose, the margins entire. Capitulescences pa-
niculate-corymbose, terminal, pedunculate, the
peduncle with 2 bracts linear, 6-9 mm long. Ca-
pitula with involucres cylindric; phyllaries
2-seriate, 2 linear to filiform and 5 lanceolate; flo-
rets 7-8, the corollas yellow, bilabiate, the outer
lip tridentate, the inner lip bifid. Achenes 4-5.5.
mm long, pubescent with glandular trichomes;
pappus 8-10 mm long, yellowish.
DISTRIBUTION— Endemic to the intermontane
regions of northern Peru (900-2000 m).
Trixis subparadoxa most closely resembles T.
cacalioides but differs in having densely strigose,
strictly entire leaves and strongly differentiated,
biseriate phyllaries.
AMAZONAS. Chachapoyas: above Balsas, Ferreyra
20761 (USM), 13335 (USM). CAJAMARCA. Jaen: be-
tween Porculla and Jaen, Ferreyra 13635 (USM).
96
FIELDIANA: BOTANY
FIG. 20. Trixis sagasteguii. 1, habit; 2, capitulum; 3, floret with achene. (Drawn from P.C. Hutchison & J.K.
Wright 5864, F.)
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
97
Acknowledgments
I dedicate this treatment to the memory of J.
Francis Macbride, a personal friend and the first
Field Museum of Natural History botanist to work
on the Flora of Peru.
Study for this treatment was conducted in the
following institutions: Gray Herbarium of Har-
vard University (GH); Field Museum of Natural
History (F); The New York Botanical Garden (NY);
United States National Herbarium, Smithsonian
Institution (us); Missouri Botanical Garden (MO);
University of California, Berkeley (uc); and Her-
bario San Marcos, Museo de Historia Natural "Ja-
vier Prado," Universidad San Marcos, Lima (USM).
I wish to thank the directors, curators, and col-
leagues of these institutions, who have extended
facilities and hospitality, and particularly Dr. Mar-
cia Koth, Director of the Fulbright Commission,
Lima; Laurence Skog, Dieter Wasshausen, John
Wurdack, and Jose Cuatrecasas of the Smithson-
ian Institution; the late Timothy Plowman, Field
Museum of Natural History; Peter Raven and Al-
wyn Gentry, Missouri Botanical Garden; Ghillean
Prance, Kew Botanical Garden; and Lincoln Con-
stance, University of California at Berkeley.
I am most grateful to Michael Dillon, Nancy
Hensold, and Elizabeth Moore for their help with
translating and editing earlier versions. John
Strother and an anonymous reviewer made many
corrections and constructive criticisms that greatly
improved the manuscript.
I also extend a special thanks to Field Museum
scientific illustrator, Zorica Dabich, for providing
the excellent drawings that appear here.
98
FIELDIANA: BOTANY
Colombia
1. Tumbes
2. Piura
3. Lambayeque
4. Cajamarca
5. Amazonas
6. La Libertad
7. San Martin
8. Loreto
9. Ancash
1 0. Huanuco
11. Lima
1 2. Pa sco
1 3. JunTn
14. Ucayali
15. lea
1 6. Huancavelica
1 7. Ayacucho
1 8. Apurfmac
1 9. Cuzco
20. Madre de Dios
2 1 . Arequipa
22. Puno
23. Moquegua
24. Tacna
Chile
DEPARTMENTS OF PERU
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
99
Index to Latin Names
Names for new combinations are in boldface, accepted names are in Roman type, and synonyms or
doubtful species are in italics. Page numbers in boldface refer to descriptions and numbers in italics refer
to illustrations.
Arnaldoa 1 , 4
coccinosantha 4
macbrideana 5
magnified 5
peruviana 4
weberbaueri 5, 6
Arnica gerbera 37
Aster trinervis 52
Atractylis mexicana 5 1
purpurea 63
Bacasia corymbcsa 1 1
lanceolata 1 1
spinosa 1 1
Barnadesia 1, 7
berberoides 8, 9
blakeana 10
caryophylla 1, 10
chachapoyasensis 15
coccinosantha 4
corymbosa 11
dombeyana 11
glomerata var. distincta 14
horrida 12
hutchisoniana 13
jelskii 13
kingii 15
lanceolata 1 1
laurifolia 37
lehmanni 14
var. angustifolia 14
macbridei 14
macrocephala 8
polyacantha 14
var. attenuata 1 5
pycnophylla 8
reticulata 15
spinosa 1 1
venosa 1 1
weberbaueri 5
wurdackii 15
Centroclinium altissimum 52
albicans 66
reflexum 72
Cephaloseris poeppigii 83
Chabraea 50
daucifolia 50
laciniata 50
purpurea 50
Chaetachlaena odorata 72
Chaetanthera 16
chiquianensis 17
ciliata 16
cochlearifolia 11, 18
multiflora 77
peruviana 17
pinnatifida 78
pulvinata 17
pungens 79
stuebelii 19
tenella 19
Chaptalia 20
callacallensis 20
cordata 21
var. ferrugineo-tomentosa 24
exscapa 21
var. microcephala 21
integerrima 22
isernina 22
malcabalensis 23, 23
nut an s 1, 23
oblonga 24
ovalis 24
rotundifolia 24
similis 25
tomentosa 19
Chucoa 1, 25
ilicifolia 25, 26
Chuquiraga 1, 27
coccinosantha 4
ferox 34
horrida 35
johnstonii 1, 28
jussieui 28
muschleri 5
peruviana 28
pseudoruscifolia 28
rotundifolia 29, 30
seleriana 12
spinosa 1, 29
subsp. huamanpinta 29
subsp. rotundifolia 29
weberbaueri 31
Clarionea cilosa 76
macrocephala 79
Cursonia 63
peruviana 63
Dasyphyllum 31
barbatum 5
brasiliense 32, 33
brasiliense var. barnadesioides 32
brevispinum 34
cabrerae 34
ferox 34
horridum 34
hystrix 34
var. hystrix 35
var. peruvianum 36
leiocephala 36
weberbaueri 36
Diaphoranthus fuscus 85
Diazeuxis 5 1
mutisiana 5 1
trinervis 52
Doronicum incanum 9 1
Dumerilla axillaris 44
mutisiana 5 1
paniculata 45
Flotovia barnadesioides 32
ferox 34
horrida 35
hystrix 35
leiocephala 36
weberbaueri 36
Fulcaldea 36
laurifolia 36, 37
Gerbera 37
hieracioides 92
jamesonii 37
linnaei 37
Gochnatia 39
arequipensis 40
boliviana 41
cardenasii 40
iserniana 40
patazina 40
vargasii 41
vernonioides 39, 41, 42
Harthamnus 82
boliviensis 82, 83
Joannea brasiliensis 32
Jungia 41
amplistipula 44
axillaris 44
discolor 44
floribunda 45
floribunda 47
ferruginea 41
jelskii 46
longifolia 47
malvaefolia 46
paniculata 45
rugosa 46
schuerae 46
seleriana 44
spectabilis 47
stuebelii 47
vitocensis 48
weberbaueri 48, 49
Leucheria 50
daucifolia 50, 50
100
FIELDIANA: BOTANY
hieracioides 50
stuebelii 47
Lophopappus 1, 87
berberidifolius 88
blakei 88
cuneata 88
cuneatus 89
foliosus 87, 90
peruvianus 90
Lycoseris 51
mexicana 51
peruviana 51
retroflexa 52
trinervis 52, 53
var. altissima 52
Mutisia 1, 54
acuminata 55
var. acuminata 56
var. bicolor 56
var. hirsuta 56
alata 57
andersonii 57
arequipensis 57
clematis 54
cochabambensis 58
hastata 58
lanata 59
lanigera 59
ledifolia 59
macrantha 6 1
mandoniana 58
mathewsii 60
var. anomala 60
microcephala 63
orbignyana 60
peduncularis 55
pulcherrima 2, 61
rauhii 61
venusta 61, 62
viciaefolia 55
weberbaueri 60
wurdackii 2, 63
Mutisieae 1
Onoseris 63
acerifolia 65
albicans 65
altissima 52
amplexicaulis 66
annua 67
cabrerae 67
castelnaeana 68
chrysactinioides 68
cumingii 68
gnaphalioides 69
hieracioides 92
humboldtiana 69
hyssopifolia 69
var. teretifolia 69
integrifolia 66
linearifolia 70
longipedicellata 72
lopezii 70, 71
macbridei 71
minima 72
odorata 63, 72
parva 72
peruviana 73
purpurea 63
reflexa 72
salicifolia 66
silvatica 74
speciosa 74, 93
stuebelii 74
weberbaueri 74
Oreastrum cochlearifolium 1 7
Perdicium cacalioides 94
divaricatum 95
magellanicum 75
oblongum 24
ovale 24
purpureum 50
Perezia 75
section Acourtia 75
bidentata 77
ciliaris 79
ciliosa 76
cirsiifolia 76
coerulescens 76
var. amplibracteata 76
conaicaensis 79
coriacea 79
fosbergii 79
magellanica 75
multiflora 77
nivalis 76
pinnatifida 78
poeppigii 77
pungens 79, 80
pygmaea 81
stuebelii 79
sublyrata 81
var. glabrescens 82
virens 77
weberbaueri 79
Plazia 82
conferta 82, 83
daphnoides 83, 84
Polyachyrus 83
annuus 85
fuscus 85
glandulosus 87
mollendoensis 85
nesites 85
poeppigii 83
sphaerocephalus 86, 87
villosus 87
Prionanthes antimenorrhoea 95
Proustia 87
berberidifolia 1, 88
cuneata 88
cuneifolia 89, 89
foliosa 1, 90
oblongifolia 90
peruviana 1, 90
pungens 89
pyrifolia 87
Ptilurus daucifolius 50
Seris castelnaeana 68
Trichocline 20, 91
beckeri 92
caulescens 91, 92
hieracioides 92
incana 9 1
oblonga 24
ovalis 24
peruviana 92
stuebelii 9 1
Trixis 93
antimenorrhoea 95
var. auriculata 96
cacalioides 94
churinensis 95
divaricata 95
hexantha 94
inula 93
neaeana 96
oblonga 24
paradoxa 94
sagasteguii 96, 97
subparadoxa 96
Turpinia laurifolia 37
Tussilago exscapa 2 1
integerrima 22
nutans 23
Weberbaueriella johnstoniana 27
Xenophontia caryophylla 10
FERREYRA: FLORA OF PERU. FAMILY ASTERACEAE. VI.
101
A Selected Listing of Other Fieldiana: Botany Titles Available
FLORA OF PERU. By j
and Samuel B. Jones). Conspectus and Ind
Publication 1314, $10.00
FLORA OF PERU. By J. Francis Macbrid<
Anthemideac\ ael O. Dillon
1 page
Publication 1319, $3.00
FLORA OF PERU. By J. Francis Macbridc and ositae: Part III. Genus
Mikania-Tribe Eupatorieae. By W« Sidney McD;
9, 1982. 56 pages, 11 illus.
Publication 1333. $7.00
FLORA OF PERU. By J. Francis Macbride and collaborators. Family
Cardueae By Michael O Dillo • 10> 1982. 8
Publication 1335, $3.00
T.ORA OF PERU. By J. Francis Macbride and <?ollabora
tae). By Michael O. Dillon.
ilus.
Publication 1336, $3.00
A Systematic Study of Flourensia (As;
n.s., no. 16. 1984. 63 pages, 41 illus.
Publication 1357. $10.00
FLORA OF PERU. By J. Francis Macbride and coli
O. Dillon and Abundio Sagastegui Alva.
Publication 1422, $15.00
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