OF
ILLINOIS
AT URBANA
BIOLOGY
'S
FIELDIA
Botany
MEW SERIES, NO. 26
FLORA OF PERU
J. Francis Macbride and Collaborators
Family Asteraceae: Part V
Michael O. Dillon
Abundio Sagastegui Alva
January 31, 1991
Publication 1422
PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY
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Croat, T. B. 1978. Flora of Barro Colorado Island. Stanford University Press, Stanford, Calif., 943 pp.
ib, P. J., J. R. Lloyd, and T. D. Pennington. 1963. A comparison of montane and lowland rain forest in Ecuador.
I. The forest structure, physiognomy, and floristics. Journal of Ecology, 51: 567-601.
Langdon, E. J. M. 1979. Yage among the Siona: Cultural patterns in visions, pp. 63-80. In Browman, D. L.,and R. A.
Schwarz, eds., Spirits, Shamans, and Stars. Mouton Publishers, The Hague, Netherlands.
1946. The historic tribes of Ecuador, pp. 785-821. In Steward, J. H., ed., Handbook of South American
Indians. Vol. 2, The Andean Civilizations. Bulletin 143, Bureau of American Ethnology, Smithsonian
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/e, R. G. 1981. Ferns and fern allies of Guatemala. Part II. Polypodiaceae. Fieldiana: Botany, n.s., 6: 1-522,
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FIELDIANA
Botany
NEW SERIES, NO. 26
FLORA OF PERU
J. Francis Macbride and Collaborators
Family Asteraceae: Part V
Michael O. Dillon
Department of Botany
Field Museum of Natural History
Chicago, Illinois 60605-2496
Abundio Sagastegui Alva
Universidad Antenor
Orrego de Trujillo
Trujillo, Peru
Accepted September 12, 1988
Published January 31, 1991
Publication 1422
PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY
1991 Field Museum of Natural History
Library of Congress Catalog Card Number: 90-85756
ISSN 00 15-0746
PRINTED IN THE UNITED STATES OF AMERICA
Table of Contents
ABSTRACT 1
TRIBE INULEAE 1
Key to Genera of Inuleae in Peru 2
I. Achyrocline 3
Key to Species of Achyrocline 3
1 . Achyrocline alata 4
2. Achyrocline celosioides 5
3. Achyrocline peruviana 6
4. Achyrocline ramosissima 6
5. Achyrocline satureioides 8
II. Antennaria 9
1. Antennaria linear ifolia 9
III. Belloa 11
Key to Species of Belloa 12
1. Belloa longifolia 12
2. Belloa lopezmirandae 14
3. Belloa pickeringii 14
4. Belloa piptolepis 14
5. Belloa plicatifolia 15
6. Belloa schultzii 16
7. Belloa spathulifolia 16
8. Belloa subspicata 16
9. Belloa turneri 18
IV. Blumea 18
1 . Blumea viscosa 20
V. Chevreulia 22
1 . Chevreulia acuminata 22
VI. Cuatrecasasiella 22
1 . Cuatrecasasiella isernii 24
VII. Facelis 24
Key to Species of Facelis 25
1. Facelis lasiocarpha 25
2. Facelis plumosa 25
VIII. Gamochaeta 27
Key to Species of Gamochaeta 28
1 . Gamochaeta americana 28
2. Gamochaeta humilis 29
3. Gamochaeta oreophila 29
4. Gamochaeta purpurea 31
IX. Gnaphalium 32
Key to Species of Gnaphalium 33
1 . Gnaphalium antennarioides 34
2. Gnaphalium badium 34
3. Gnaphalium dombeyanum 35
4. Gnaphalium elegans 37
5. Gnaphalium lacteum 39
6. Gnaphalium luteo-album 39
7. Gnaphalium polium 40
X. Helichrysum 41
1. Helichrysum bracteatum 41
XI. Jalcophila 41
1 . Jalcophila peruviana 42
XII. Loricaria 42
Key to Species of Loricaria 44
1 . Loricaria ferruginea 44
2. Loricaria graveolens 46
3. Loricaria leptothamna 46
4. Loricaria lucida 47
5. Loricaria lycopodinea 47
6. Loricaria macbridei 48
7. Loricaria thuyoides 48
Key to Varieties of
Loricaria thuyoides 49
7a. Loricaria thuyoides var.
thuyoides 49
7b. Loricaria thuyoides var.
stuebelii 49
8. Loricaria thyrsoidea 49
XIII. Lucilia 50
Key to Species of Lucilia 50
1 . Lucilia conoidea 50
2. Lucilia kunthiana 51
XIV. Mniodes 52
Key to Species of Mniodes 53
1. Mniodes andina 53
2. Mniodes aretioides 53
3. Mniodes coarctata 55
4. Mniodes pulvinulata 55
XV. Novenia 56
1 . Novenia acaulis 56
XVI. Pluchea 58
Key to Species of Pluchea 58
1. Pluchea absinthioides 58
2. Pluchea chingoyo 59
3. Pluchea microcephala 60
4. Pluchea zamalloae 60
XVII. Pterocaulon 62
1. Pterocaulon alopecuroides 62
XVIII. Stuckertiella 62
1. Stuckertiella capitata 63
XIX. Tessaria 65
1. Tessaria integrifolia 65
ACKNOWLEDGMENTS 67
MAP OF PERU 68
INDEX TO LATIN NAMES . . 69
List of Illustrations
1 . Achyrocline peruviana 7
2. Antennaria linearifolia 10
3. Belloa longifolia, B. plicatifolia 13
111
4. Belloa spathulifolia 17
5. Belloa turneri 19
6. Blumea viscosa 21
7. Chevreulia acuminata 23
8. Cuatrecasasiella isernii 24
9. Facelis lasiocarpha, F. plumosa 26
1 0. Gamochaeta oreophila 30
1 1 . Gnaphalium elegans 38
12. Jalcophila peruviana 43
13. Loricaria lucida, L. leptothamna, L.fer-
ruginea 45
14. Lucilia conoidea, L. kunthiana 51
15. Mniodes andina, M. aretioides, M.
coarctata, M. pulvinulata 54
16. Novenia acaulis 57
1 7. Pluchea chingoyo 59
1 8. Pluchea microcephala 61
19. Pterocaulon alopecuroides 63
20. Stuckertiella capitata 64
2 1 . Tessaria integrifolia 66
IV
FLORA OF PERU
Family Asteraceae: Part V
Abstract
The tribe Inuleae (Asteraceae) is treated for Peru
and includes 19 genera, 55 species, and 2 varieties.
Keys to genera and species, descriptions, and il-
lustrations are provided for Achyrocline (5 spp.),
Antennaria (1 sp.), Belloa (9 spp.), Blumea (1 sp.),
Chevreulia (1 sp.), Cuatrecasasiella (1 sp.), Facelis
(2 spp.), Gamochaeta (4 spp.), Gnaphalium (7 spp.),
Helichrysum (1 sp.), Jalcophila (1 sp.), Loricaria
(7 spp., 2 vars.), Lucilia (2 spp.), Mniodes (4 spp.),
Novenia (1 sp.), Pluchea (4 spp.), Pterocaulon (1
sp.), Stuckertiella (1 sp.), and Tessaria (1 sp.).
Gamochaeta oreophila Dillon & Sagast. (nomen
novum) is provided to replace G. monticola Dillon
& Sagast., an invalid name due to an earlier hom-
onym.
Tribe INULEAE
Inuleae Cass., Jour. Phys. 88: 193. 1819. TYPE:
Inula L.
GnaphalieaeRydb., Fl. Rocky Mts. 833. 1917. TYPE:
Gnaphalium L.
Dioecious or bisexual, annual or perennial herbs,
shrubs, rarely trees; stems usually arachnoid or
glandular. Leaves alternate or opposite, cauline or
rosulate, the margins mostly entire. Capitulesc-
ences paniculate, spiciform, cymose-corymbi-
form, glomerulate, or of solitary heads. Capitula
disciform, discoid, or rarely radiate; involucres cy-
lindrical to campanulate or hemispherical; phyl-
laries numerous, usually imbricate, indurate or hy-
aline, scarious, usually stramineous, sometimes
white, or variously colored; receptacles plane or
convex, usually naked; marginal florets pistillate,
the corollas filiform or ligulate (rarely tubular), the
styles divided into linear branches; disc florets
usually hermaphroditic, functionally staminate or
occasionally gynoecia fertile, the corollas tubular,
the anthers connate, appendaged, bases obtuse to
sagittate, usually caudate, the styles divided, bi-
dentate, or undivided, the branches slender, su-
bulate, and attenuate, or oblong-linear and round-
ed, obtuse, or truncate, unappendaged; ovaries
mostly terete. Achenes mostly terete, small, often
ribbed, variously pubescent, glandular, or gla-
brous; pappus bristles usually 1-2-seriate, smooth
to scabrid, barbellate, or plumose, often fused at
the base, deciduous together as a unit or persistent
on the achene.
Inuleae is well defined in Peru; members may
be recognized by alternate or at times rosulate
leaves (opposite in Chevreulia and Cuatrecasa-
siella), usually with arachnoid pubescence and
caudate anthers. In Peru, it includes 55 species
and two varieties distributed in 1 9 genera. Several
genera have centers of diversity in the Peruvian
Andes, including Belloa, Loricaria, and Mniodes.
A few members are used medicinally (e.g., Achy-
rocline, Gnaphalium, Loricaria, Tessaria) and one,
Helichrysum, is cultivated for ornamental value.
References
MILLIARD, O. M., AND B. L. BURTT. 1981. Some generic
concepts in Compositae Gnaphaliinae. J. Linn. Soc.
Bot., 82: 181-232.
MERXMULLER, H., P. LEINS, AND H. ROESSLER. 1977.
Inuleae Systematic review, pp. 577-602. In Hey-
wood, V. H., et al. (eds.), The Biology and Chemistry
of the Compositae, Academic Press, London.
TURNER, B. L., A. M. POWELL, AND J. CUATRECASAS.
1967. Chromosome numbers in Compositae. XI. Pe-
ruvian species. Ann. Missouri Bot. Gard., 54: 172-
177.
DILLON & SAGASTEGUI: FLORA OF PERU. V.
Key to Genera of Inuleae in Peru
1. Plants dioecious; individual capitula exclusively homogamous (polygamo-dioecious in Antennarid)
2
2. Erect shrubs or subshrubs, 20 cm tall or greater; stems laterally flattened; leaves laterally
compressed XII. Loricaria
2. Cespitose, perennial herbs or suffruticose perennials, less than 20 cm tall;
stems cylindrical; leaves not laterally compressed 3
3. Stems clearly visible, internodes evident; basal leaves rosulate and cauline leaves alternate
or basal leaves lacking and cauline leaves opposite, decussate, mostly linear to oblong . . 4
4. Basal leaves rosulate; capitulescences scapose, cymose-corymbose . . II. Antennaria
4. All leaves opposite, cauline; capitula solitary, terminal VI. Cuatrecasasiella
3. Stems shortened into a dense cushion, internodes not evident; leaves spiral and tightly
appressed to the stem, imbricate, usually ovate to obovate XIV. Mniodes
1 . Plants bisexual; individual capitula heterogamous 5
5. Styles of hermaphroditic (or functionally staminate) florets briefly divided at the apex, the
branches pilose along the dorsal side and the distal portion of the shaft, apex rounded; or greatly
divided with two linear branches, apex acute to rounded, pilose 6
6. Stems winged, the wings discolorous; leaves discolorous; capitulescences spiciform-glo-
merulate; capitula sessile XVII. Pterocaulon
6. Stems un winged, or if winged, not discolorous; leaves not discolorous; capitulescences of
corymbiform cymes; capitula pedunculate 7
7. Large shrubs or more often trees; capitula with only one hermaphroditic floret
XIX. Tessaria
7. Shrubs or herbs; capitula with (3-)5-15 hermaphroditic florets XVI. Pluchea
5. Styles of hermaphroditic florets divided, the branches linear to oblong, apex truncate to acute
or rounded, dorsally papillose or with an apical ring of papillae 8
8. Achenes densely sericeo-pubescent 9
9. Plants caulescent; stems sparsely branched and erect or much-branched and cespitose
or decumbent 10
10. Erect annual herbs; pappus bristles plumose VII. Facelis
10. Cespitose perennial herbs; pappus bristles scabrous XIII. Lucilia
9. Plants acaulescent, pulvinate XV. Novenia
8. Achenes glabrous, papillose, or pubescent with scattered glandular or duplex trichomes .
11
1 1 . Achenes fusiform, terminally rostulate V. Chevreulia
1 1 . Achenes ovoid to oblong-cylindrical, not rostulate 12
12. Pappus bristles free at the base (except Helichryswri), individually deciduous . .
13
13. Capitula solitary, pedicels evident; leaves glabrous or glandular on both sur-
faces 14
14. Capitula 20-50 mm wide; phyllaries variously colored, greater than 2 mm
wide (cultivated) X. Helichrysum
14. Capitula 4-5 mm wide; phyllaries green, less than 1 mm wide (native
species) IV. Blumea
1 3. Capitula glomerulate, pedicels obscure; leaves tomentose (at least lower sur-
faces) 14
15. Involucres cylindrical to narrowly oblong or narrowly campanulate; pis-
tillate florets 1-8 1. Achyrocline
15. Involucres campanulate; pistillate florets 25-150 IX. Gnaphalium
12. Pappus bristles fused at the base, deciduous together as a unit 16
16. Stems 1-2 cm long; leaves oblong-lanceolate, 3.5-5 mm long, 0.5-1.5 mm
wide; pistillate florets usually 4 per capitulum XI. Jalcophila
FIELDIANA: BOTANY
16. Stems greater than 2 cm long; leaves linear-lanceolate to ovate, usually greater
than 5 mm long, greater than 2 mm wide; pistillate florets ususally 10 or more
per capitulum 17
17. Hermaphroditic florets 4-merous XVIII. Stuckertiella
17. Hermaphroditic florets 5-merous 18
18. Style branches of hermaphroditic florets acute to rounded, evenly
papillose externally and lacking an apical ring of collecting trichomes
III. Belloa
1 8 . Style branches of hermaphroditic florets truncate, not evenly papillose
externally, but with an apical ring of collecting trichomes
. . VIII. Gamochaeta
I. Achyrocline
Achyrocline (Less.) DC., Prodr. 6: 219. 1838.
TYPE: Gnaphalium satureioides Lam. =
Achyrocline satureioides (Lam.) DC.
Gnaphalium subgenus Achyrocline Less., Syn. Comp.
332. 1832.
Perennial herbs, suffrutescent or shrubs, to-
mentose or rarely glabrate; stems leafy, erect to
decumbent or rarely scandent. Leaves alternate;
petiolate or sessile; blades linear to ovate, arach-
noid to lanate, the margins entire to crenulate.
Capitulescences glomerulate, cymose-paniculate
to corymbose, open or dense, rarely of solitary
heads. Capitula 3-5.5 mm high, ca. 1.5-2 mm
wide, disciform; involucres cylindric to narrowly
oblong or narrowly campanulate; phyllaries im-
bricate, 2-3-seriate, scarious, brown to white or
rarely reddish, the outer gradually shorter, base
lanate, the inner glandular on abaxial surface; re-
ceptacles flat, naked or rarely with rudimentary
paleae; pistillate florets 1-1 1(-23), the corollas fi-
liform, dentate or shallowly 2-4-cleft; hermaph-
roditic florets l-4(-6), fertile, the corollas tubular,
base dilated, 4-5-lobed, the lobes usually pubes-
cent with multicellular, stalked, capitate-glandular
trichomes, the anther bases sagittate, caudate, the
terminal appendages lanceolate, obtuse, the style
branches slender, unappendaged, apex truncate,
penicillate, the nectary conspicuous. Achenes ob-
long to ovoid, subcompressed, ca. 4-5 -ribbed,
papillose with imbricate cells; pappus bristles sca-
brid, barbellate, uniseriate, bases cohering by pa-
tent cilia, readily deciduous, apical cell rounded.
Chromosome number: n = 14.
A genus of 1 5-20 species distributed primarily
in tropical and subtropical South America, with
five in Mexico and Central America and a few in
tropical Africa, Madagascar, and Comores. In Peru,
it is represented by five species distributed in a
wide variety of habitats and elevations throughout
the Andean Cordillera. This genus most closely
resembles Gnaphalium in habit and floral struc-
ture; however, Achyrocline is readily distinguished
by its narrow capitula with fewer than 25 florets
and glabrous achenes.
This genus is in need of experimental and field
study throughout its range. Several of its species
exhibit considerable morphological and ecological
variability that presents problems for determining
species boundaries.
References
DILLON, M. O., AND A. SAGASTEGUI A. 1986. Anew
species of Achyrocline (Inuleae- Asteraceae) from Peru.
Phytologia, 59: 107-110.
GIANGUALANI, R. N. 1 976. Las especies argentinas del
genero Achyrocline (Compositae). Darwiniana, 20: 549-
576.
MILLIARD, O.M., AND B. L.BURTT. 1981. Some generic
concepts in Compositae Gnaphaliinae. J. Linn. Soc.
Bot., 82: 181-232.
JANSEN, R. K., AND T. F. STUESSY. 1980. Chromosome
counts of Compositae from Latin America. Amer. J.
Bot., 67: 585-594.
JANSEN, R. K.., T. F. STUESSY, S. DiAZ-PiEDRAHfrA, AND
V. FUNK. 1984. Recuentos cromosomicos en Com-
positae de Colombia. Caldasia, 14: 7-20.
NESOM, G. L. 1990. Taxonomy of Achyrocline (Aster-
aceae: Inuleae) in Mexico and Central America. Phy-
tologia, 68: 181-185.
Key to Species of Achyrocline
1 . Leaves decurrent; stems winged A. alata
1 . Leaves not decurrent; stems not winged 2
2. Leaves linear to linear-lanceolate; outer phyllaries ovate, apex obtuse or rounded 3
DILLON & SAGASTEGUI: FLORA OF PERU. V.
3. Plants with reduced habits, usually less than 50 cm tall; leaves 2-3(-5) cm long, 1-3 mm wide
A. ramosissima
3. Plants not reduced in habit, usually greater than 1 m tall; leaves 4-10 cm long, 2-9 mm wide
A. satureioides
2. Leaves lanceolate to elliptic or obovate; outer phyllaries lanceolate, apex acute 4
4. Leaves elliptic to obovate, (0.5-)1.3-3.5 cm long, 5-16 mm wide A. celosioides
4. Leaves lanceolate to elliptic, 1.5-5 cm long, 2-8 wide A. peruviana
1. Achyrocline alata (Kunth) DC., Prodr. 6: 221.
1838.
Gnaphalium alatum Kunth, Nov. Gen. Sp. (fol.) 4:
62. 1820. TYPE: Ecuador, Prov. Chimborazo,
"circa Alausi Quitensium," A. Bonpland 3243
(holotype, P, not seen, IDC Microfiche 6209. 96:
II. 4, F neg. 37578, fragment, F).
Gnaphalium incanum Kunth, Nov. Gen. Sp. (fol.) 4:
63. 1820. TYPE: Peru, Dept. Cajamarca, Prov.
Hualgayoc, "juxta urbem Micuipampa," 1 830 m,
A. Bonpland 3680 (holotype, P, not seen, IDC
Microfiche 6209. 96: II. 5, F neg. 37601).
Gnaphalium pellitum Kunth, Nov. Gen. Sp. (fol.) 4:
63. 1820. TYPE: Peru, Dept. Cajamarca, Prov.
Hualgayoc, "juxta urbem Micuipampa," 1 830 m,
A. Bonpland s.n. (holotype, P, not seen, IDC Mi-
crofiche 6209. 96: II. 6, F neg. 37604, fragment,
F). Non G. pellitum Hort. Getting, ex DC.
Gnaphalium rufescens Kunth, Nov. Gen. Sp. (fol.) 4:
61. 1820. TYPE: Colombia, Prov. Cauca, "inter
fluvium Putes et villam San Miguel," A. Bonp-
land s.n. (holotype, P, not seen, IDC Microfiche
6209. 96: I. 7, F neg. 37571, fragment, F).
Achyrocline madioides Meyen & Walpers, Nov. Ac-
torum Acad. Caes. Leop.- Carol. German. Nat.
Cur. (suppl.) 19: 275. 1843. TYPE: Peru, Dept.
Arequipa, W. G. Walpers s.n. (holotype, B, prob-
ably destroyed, F neg. 1 5 1 34).
Perennial herbs to 1.5 m tall; stems ascending
to decumbent or scandent, glabrous to tomentose,
alate. Leaves sessile; blade linear-lanceolate to ob-
lanceolate, decurrent on stems, 2-12 cm long, 2-
15 mm wide, trinerved from near the base, gla-
brous and glandular to densely tomentose beneath
and arachnoid above, base attenuate, apex acute,
the margins entire to crenulate. Capitulescences
glomerulate, cymose-paniculate, dense or open.
Capitula 3.0-5.5 mm high, ca. 1.5 mm wide, ses-
sile; involucres cylindrical to narrowly campanu-
late, yellow to cream-white; outer phyllaries ovate
to lanceolate, 2-3 mm long, 1-1.5 mm wide, apex
acute, the inner phyllaries linear-spatulate, 4-5 mm
long, 0.5-1.5 mm wide, apex acute to obtuse; pis-
tillate florets 3-1 1(-23), the corollas 2.5^4.0 mm
long; hermaphroditic florets l-3(-6), the corollas
2.54.0 mm long. Achenes narrowly ellipsoid to
ovoid, 0.5-0.7 mm long; pappus ca. 2.5 mm long.
Chromosome number: n = 14 (Jansen & Stuessy,
1980).
Achyrocline alata has a wide distribution
throughout the Andean Cordillera from Colombia
to Argentina, Brazil, and Paraguay. It inhabits a
variety of environments within its range, but in
Peru it is usually found in secondary vegetation
or disturbed roadsides (2400-4100 m). In Peru it
is known by the common names: ajenjo, Foster-
sacha, huira-huira, ishpingo amarillo, vira-vira,
yatama blanca, and yshpingo.
Achyrocline alata is readily distinguished from
other Peruvian species by its decurrent leaf bases
and alate stems. Pubescence is quite variable; in-
dividuals range from predominately glabrous to
densely arachnoid-tomentose. No ecological or
geographical patterns are obvious for these pu-
bescence morphs, and the capitular and floral mor-
phology is relatively constant. Gnaphalium incan-
um, G. pellitum, and G. rufescens are all readily
referable to Achyrocline. They are all quite pu-
bescent, and their decurrent leaf bases forming
winged stems and few-flowered capitula agree well
with the present broad concept of A. alata. Achy-
rocline madioides Meyen & Walpers is provision-
ally placed here also. Further detailed populational
investigations may lead to the taxonomic recog-
nition of these pubescent morphs.
Individuals with smaller leaves, greatly reduced
stem wings, and smaller, few-flowered capitula ap-
proach A satureioides, a species with linear leaves,
unwinged stems, and narrowly cylindrical capit-
ula. While A. alata and A. satureioides are closely
related, the latter species is typically found at el-
evations below 1 500 m. We have not found sym-
patric populations to date, but two collections
(Vargas 456, F, and Weberbauer 5638, GH) have
stems both with and without wings.
AMAZONAS. Chachapoyas: 28-31 km SW of Lei-
mebamba on road to Balsas, 3350-3450 m, Gentry et
al. 23158 (F, MO); 41 km SW of Leimebamba on road
to Balsas, 3110m, Gentry et al. 23147 (F, MO); Cerros
Calla Calla, W side, 45 km above Balsas, 3 1 00 m, Hutch-
ison & Wright 5774 (F, GH, MO, NY); Cerro Chimchula,
cerca a Chachapoyas, 2500 m, Ochoa 1628, 1637 (F);
rocky slopes of Cano Santa Lucia just E of Chachapoyas,
2200-2300 m, Wurdack 726 (GH, NY, us). Rodriguez de
Mendoza: Mendoza, 1600 m, Woytkowski 8273 (GH, MO).
ANCASH. Bolognesi: Valle de Rio Fortaleza, 3600 m,
FIELDIANA: BOTANY
Cerrate & Gayoso 7819 (MO); Chiquian, 3350 m, Fer-
reyra 7455 (MO, us). Casma: abra de la Cordillera Negra,
4050 m, Mostacero et al. 536 (F, HUT, MO). Huaraz: Cerro
San Cristobal, 3800 m, Evangelista s.n. (F, HUT, MO);
Laguna de Shurup, 3800 m, Evangelista s.n. (F, HUT, MO).
Huaylas: Laguna de Paron, 4100 m, Mostacero et al.
540, 553 (F, HUT, MO). Yungay: Llanganuco, Ferreyra
14342, 16826, 16863 (MO, USM); slopes below Laguna
de Llanganuco in Quebrada de Llanganuco, ca. 25 km
above Yungay, 4000 m, Edwin & Schunke 3818 (COL, F,
us); Yungay- Yauya road, 3480 m, Gentry et al. 37366
(F,MO). APURIMAC. Abancay: arriba de Abancay, 3350-
3450 m, Ferreyra 9812 (MO, USM). Aymaraes: 4 km SW
of Cotaruse, 26 km SW of Chalhuanca, 3220 m, Gentry
et al. 23302 (F, MO, USM). AYACUCHO. Huanta: Ccar-
rapa, between Huanta and Rio Apurimac, 1 200 m, Killip
& Smith 22430 (F, us). La Mar: above Ayna (Aina), 1 600
m, Weberbauer 5638 (F, GH, us). CAJAMARCA. Caja-
marca: ca. 6.5 km SW of Cajamarca, 2980 m, Dillon et
al. 2904 (F, MO, TEX, USM); San Juan a Cajamarca, 2800
m, Ochoa 1475 (GH, F); La Encanada- Jalca de Kumulca,
3300 m, Sagdstegui et al. 8100 (F, HUT, MO, NY), 8105A
(HUT, MO); NE of Porcon, Sanchez 2858 (CPUN, F). Ce-
lendin: 4 km NNE of Celendin on road to Balsas, Edwin
& Schunke 3582 (COL, F, NY, us); entre Celendin y Jalca
Cumullca, 3000-3 100 m, Ferreyra 15087 (MO, us, USM);
Celendin, 2700 m, Hutchison & Wright 5127 (F,MO,NY);
Canyon of the Rio Maranon above Balsas, 2930 m,
Hutchison & Wright 5374 (F.MO.NY). Chota: Cochabam-
ba, 1 800-2200 m, Ferreyra 783 (USM). Contumaza: Guz-
mango, Sagdstegui 7726 (F, HUT); Cascabamba, 3050 m,
Sagdstegui et al. 9038 (F, HUT, K, MO); Jalca El Chuno
(Pozo Chuno), 4500 m, Sagdstegui et al. 9373 (F, HUT,
MO); Cruz del Hueco, 2800 m, Sagdstegui et al. 9918 (F,
HUT, MO). Cutervo: 4 km E of Huambos on Cutervo trail,
2800 m, Stork & Horton 19199 (F). CUZCO. Anta: Lim-
atambo, 2400 m, Vargas 456 (MO). Calca: Amparacs,
3300 m, Chavez 3283 (MO). Cuzco: San Sebastian, 3300-
3400 m, Pennell 13626 (F, GH, NY, us). Paruro: Aray-
pallpa, 3200 m, Vargas 456 (F). Paucartambo: 9-12 km
NE of Paucartambo, 2980-3050 m, Gentry et al. 23413,
23423 (F, MO, USM); 22 km NE of Paucartambo, 3350 m,
Gentry et al. 23430 (F, MO, USM); Cerro de Cusilluyoc,
Pillahuata, 2700-2900 m, Pennell 14089 (GH). Urubam-
ba: Machupicchu, 2400 m, Herrera 1974 (F); Ollantay-
tambo, 2800 m, Herrera 3447 (F); Machu Picchu, above
Paucarcancha, 3225 m, Peyton & Peyton 133 (F).
HUANUCO. Huamalies: Llata, 7000 ft, Macbride &
Featherstone 2242 (F, GH, us). Huanuco: cerca a Mito-
cucho, 2600 m, Cerrate et al. 6887 (MO, USM); between
Huanuco and Chinchao, Dwyer6115 (MO); Sheiricancha,
cerca a Acomayo, 2500-2600 m, Ferreyra 9333 (MO,
USM); Acomayo, Distrito de Chinchao, 2500 m, Ridoutt
11560 (MO, USM). Pachitea: Mitotambo, arriba de Mito,
3200-3300 m, Ferreyra 10380, 10380A (USM). ICA.
Nazca: Puquio Valley, 3400 m, Rauh & Hirsch P435
(NY). Pisco: Valle Rio Pisco, 3700 m, Rauh & Hirsch
P354 (NY). JUNIN. Conception: Mito, 9000 ft, Macbride
& Featherstone 1474, 1475 (F), 1644 (F.US), 1818 (F.GH,
us). Huancayo: Km 7 Huancayo-Pariahuanca, 3300 m,
Antunez de Mayolo 38 (F); Huancayo, 3317 m, Soukup
2955 (COL), 3003 (F), 3151 (COL, us, USM). Jauja: Comas,
Ochoa 549 (F). Tarma: cumbre a Tarma, 3500 m, Cerrate
922 (MO, USM); 28-32 km NE of Tarma on road to San
Ramon, 2480 m, Dillon & Turner 1348 (F, MO, TEX, USM);
entre Tarma y San Ramon, 1 800-2 1 00 m, Ferreyra 3606
(MO, USM); Tarma, 3000-3200 m, Killip & Smith 21914
(F, NY, us); Huacapistana, 1 800-2400 m, Killip & Smith
24206 (NY, us); Tarma, Macbride & Featherstone 988
(F); Sacsamarca, Riccio 1541 (us); entre Tarma y Oroya,
10 km de Tarma, 3600 m, Tovar 2380 (USM). Yauli: San
Pedro de Cajas, S of Lake Junin, 3800-3900 m, Pearsall
924 (F). LA LIBERTAD. Otuzco: Agallpampa, 3050 m,
Lopez 1873 (F, HUT). Pataz: Huancaspata-Tayabamba,
3900 m, Lopez & Sagdstegui 8266 (HUT, MO). Santiago
de Chuco: Laguna La Victoria, camino a Conzuzo, 4000
m, Sagdstegui et al. 6190 (HUT, MO). LIMA. Canta: 3 100-
3200 m, Pennell 14612 (F, NY, us). Huarochiri: Puente
de Infiernillo, Cerrate et al. 4278 (MO, USM); Chicla, entre
San Mateo y Casapalca, 3700 m, Ferreyra 6505 (MO, us,
USM); Infiernillo, 3300 m, Ferreyra 8318 (MO, USM); 4
km E of Chicla, 3800 m, Gentry & Prance 16448 (F, MO);
Rio Blanco, 3000-3500 m, Killip & Smith 21637 (F.NY,
us), Macbride & Featherstone 672 (F, us). PASCO. Dan-
iel Carrion: Yanahuanca, ca. 10,000 ft, Macbride &
Feather stone 11 93 (F,US). Pasco: ca. 23 km NNE of Cerro
de Pasco on road to Huanuco, 3810 m, Dillon 2552 (F,
MO, USM); La Quinua, ca. 12,000 ft, Macbride & Feath-
erstone 2021 (F, us). PIURA. Huancabamba: Huanca-
bamba, 2950 m, Hutchison 1603 (F, NY); Mitopampa
(Huancabamba-Cuello del Indio), 2650 m, Sagdstegui
et al. 8227 (MO, NY). PUNO. Carabaya: Macusani to
Ollachea, 2000-3000 m, Dillon et al. 1099, 1265 (F, MO,
USM); Ollachea to San Gabon, 1000-2000 m, Dillon et
al. 1103 (F, MO, USM). SAN MARTIN. Mariscal Caceres:
Rio Abiseo National Park, 3425 m, Young & Watson
3519(p).
2. Achyrocline celosioides (Kunth) DC., Prodr. 6:
221. 1838.
Gnaphalium celosioides Kunth, Nov. Gen. Sp. (fol.)
4: 61. 1820. TYPE: Ecuador, Prov. Loja, "crescit
juxta Loxam Peruvianorum," A. Humboldt & A.
Bonplands.n. (holotype, P, not seen, IDC Micro-
fiche 6209. 96: II. 2, F neg. 37585).
Perennial herbs or shrubs to 0.5 m tall; stems
much-branched, ascending to procumbent. Leaves
sessile; blade elliptic to obovate, (0. 5-) 1.5-3. 5 cm
long, (0.3-)0.5-1.6 cm wide, trinerved from near
the base, discolorous, both surfaces tomentose,
apex obtuse to rounded, base obtuse to cuneate.
Capitulescences glomerulate, cymose-paniculate,
dense or open. Capitula 45 mm high, ca. 1.5 mm
wide, sessile; involucres cylindrical, white; outer
phyllaries ovate, ca. 2.5 mm long, 0.8-1 mm wide,
apex acute, the inner phyllaries lanceolate to ob-
lanceolate, 3.54.0 mm long, 0.40.8 mm wide,
apex acute to apiculate; pistillate florets 2, the co-
rollas 2.0-2.5 mm long; hermaphroditic florets 3,
the corollas 2.2-2.5 mm long. Achenes ovoid, 0.7-
0.8 mm long; pappus ca. 2.5 mm long.
Achyrocline celosioides ranges from southern
Ecuador to south-central Peru (2000-3400 m) and
DILLON & SAGASTEGUI: FLORA OF PERU. V.
normally occupies relatively moist sites in inter-
Andean valleys. It is readily distinguished from A.
peruviana, its nearest relative, by the former's
shorter, elliptic leaves.
A few populations display considerable varia-
tion in leaf size and shape. Two unusual collections
from the Cumbre El Gavilan (above Cajamarca)
display the following variability: ( 1 ) Ferreyra 3277
(us), has very densely lanate stems with short in-
ternodes; ovate, sessile leaves; and sessile, glo-
merulate capitulescences in the upper leaf axils;
(2) Ferreyra 8538 (MO) has very small leaves (ca.
5 mm long, ca. 3 mm wide) and capitulescences
on short peduncles.
Two collections, Sagdstegui & Dillon 12312
(Dept. Ancash) and Chavez 3372 (Dept. Apuri-
mac) have more narrowly elliptic, sessile leaves
and compact, nearly sessile glomerulate capitu-
lescences. Last, a collection from a dry site above
the Rio Maranon (Evinger & Hodge 514, US) has
smaller, more narrowly lanceolate leaves, but with
capitulescences typical for the species. All these
specimens essentially agree with A. celosioides in
capitular and floral characteristics, but further study
may prove them to be worthy of specific rank.
AMAZONAS. Bongara: between Rio Utcubamba and
Shipasbamba, 1520 m, Hutchison & Wright 3994 (uc).
Chachapoyas: 10 km E of Balzas [Balsas], Evinger &
Hodge 514 (us); rocky slopes of Cano Santa Lucia just
E of Chachapoyas, 2000-2400 m, Wurdack 532, 579
(GH, NY, uc, us). ANCASH. Recuay: rocky river bluffs,
ca. 9000 ft, Macbride & Featherstone 2517 (F). APU-
RIMAC. Abancay: Curahuasi, 2550 m, Chavez 3327
(MO). CAJAMARCA. Cajabamba: camino a Langla, 2600
m, Mostacero & Guerra 0008 (HUT). Cajamarca: Cumbre
El Gavilan, entre Cajamarca y San Juan, 3200 m, Fer-
reyra 3277 (us), 8583 (MO); La Encanada-Jalca de Ku-
mulca, 3300 m, Sagdstegui et al. 8106 (HUT); La Encan-
ada, 2750 m, Sagdstegui et al. 12013 (HUT). Celendin:
ca. 31 km NE of Cajamarca along road to Celendin, E
slopes, ca. 2950 m, Dillon & Turner 1587 (F); hills above
and to N of town, ca. 2700 m, Hutchison & Wright 5136
(uc). Hualgayoc: Bambamarca. 2600 m, Soukup & Car-
mona 4935 (us). JUNIN. Conception: Mito, ca. 9000
ft, Macbride & Featherstone 1489 (F). Prov. Unknown:
Uspachaca, ca. 8500 ft, Macbride & Featherstone 1295
(F). LA LIBERT AD. Santiago de Chuco: Hacienda An-
gasmarca, 2850 m, Weberbauer 7205 (F, GH). PIURA.
Huancabamba: on road to Canchaque, 18.5 km above
and W of Huancabamba, 2900 m, Hutchison & Wright
6635 (F, MO, NY, uc, USM).
3. Achyrocline peruviana Dillon & Sagast., Phy-
tologia 60: 107. 1986. TYPE: Peru, Dept. Ca-
jamarca, Contumaza, El Granero, laderas
abiertas, 2800 m, 14 Jun 1983, A. Sagdstegui
A. & S. Lopez M. 10661 (holotype, HUT). Fig-
ure 1.
Suffrutescent perennials to 75 cm tall, much-
branched at the base; stems arachnoid-tomentose.
Leaves sessile; blade narrowly lanceolate to ellip-
tic, (1.5-)4-5 cm long, 2-8 mm wide, discolorous,
base attenuate, apex acute, mucronulate, lower
surface densely arachnoid-tomentose, upper sur-
face sparsely arachnoid-tomentose, glabrescent,
obscurely trinervate, the margins entire. Capitu-
lescences glomerulate, cymose-paniculate, open,
terminal and axillary. Capitula ca. 4 mm high, 1 .2-
1.5 mm wide, sessile; involucres narrowly cam-
panulate, stramineous; outer phyllaries lanceolate,
2-3 mm long, ca. 1 mm wide, apex acute, the inner
phyllaries lanceolate, 3-4 mm long, 1-1.2 mm
wide, apex acute; pistillate florets 1-2, the corollas
ca. 2 mm long; hermaphroditic florets 2-3, the
corollas ca. 2.5 mm long. Achenes ovoid, ca. 1
mm long; pappus ca. 2.5 mm long.
Achyrocline peruviana is a narrow endemic
known only from dry intermontane localities in
the Departments of Cajamarca and La Libertad
(2600-3 100 m) and is primarily restricted to river
valley systems with Pacific drainage. It most close-
ly resembles A. celosioides, but differs from that
species by possessing more narrowly elliptic or
lanceolate leaves, usually at least 4 cm long, with
acute, mucronulate apices.
Some individuals of A. satureioides approach A.
peruviana in general appearance, but the former
species has strictly cylindrical capitula, with a
greater number of pistillate florets and a distri-
bution usually below 1500 m on the eastern es-
carpment of the Andean Cordillera.
CAJAMARCA. Contumaza: 1-2 km S of Contumaza,
2620 m, Dillon et al. 4455 (F, HUT, MO, TEX, us, USM);
Contumaza, 2600 m, Sagdstegui et al. 10685 (F, HUT,
MO). LA LIBERTAD. Huamachuco: Hda. Cochabamba,
2750 m, Lopez & Sagdstegui 2781 (HUT). Otuzco: Agall-
pampa, 3100 m, Lopez 895 (us), 1867 (F, HUT).
4. Achyrocline ramosissima (Schultz-Bip.) Brit-
ton, Bull. Torrey Bot. Club 19: 148. 1892.
Gnaphalium ramosissimum Schultz-Bip., Bonplandia
4: 42. 1856. TYPE: Peru, Dept. Puno, Azangaro,
cordillera prope Azangaro, 1854, W. Lechler 1780
(holotype, B, probably destroyed; isotypes, F, NY,
G-DEL, not seen, F neg. 28706, w, not seen, F neg.
33153).
FIELDIANA: BOTANY
FIG. 1. Achyrodine peruviana. A, habit; B; capitulum; C, inner phyllary; D, outer phyllary; E, hermaphroditic
floret; F, pistillate floret; G, stamen; H, style branches of hermaphroditic floret. (Drawn from Sagdstegui & Lopez
19661, HUT.)
Perennial herbs, 15-50(-70) cm tall, suffruti-
cose; stems much-branched, decumbent to as-
cending, densely lanate. Leaves sessile; blade lin-
ear to linear-lanceolate, 2-3(-5) cm long, 1-3 mm
wide, discolorous, the lower surfaces densely la-
nate, the upper surfaces tomentose, the margins
entire, revolute. Capitulescences corymbiform
cymes of glomerules, dense or open. Capitula ca.
4 mm high, ca. 1.5 mm wide, stramineous; in-
volucres cylindrical; outer phyllaries ovate, 1.5-2
mm long, 1-1.2 mm wide, 2-3-fid, base lanate,
the inner phyllaries lanceolate to oblanceolate, 3-
DILLON & SAGASTEGUI: FLORA OF PERU. V.
3.5 mm long, 0.5-0.8 mm wide; pistillate florets
generally 4, the corollas ca. 2.5 mm long; her-
maphroditic floret 1 , the corollas ca. 2.8 mm long.
Achenes ellipsoid, ca. 0.7 mm long; pappus 2.5-
3 mm long.
Achyrocline ramosissima is distributed from
central Peru to northwestern Argentina. In Peru,
it occurs in puna formations from Junin to Puno
(2800-4300 m). It most closely resembles A. sa-
tureioides but is distinguished from that species
by its reduced habit, smaller leaves and capitula,
and high-elevation distribution.
Several collections appear to approach A. sa-
tureioides in habit, but these collections have the
small capitula characteristic of A. ramosissima:
Lopez M. 618 (HUT), Cook & Gilbert 523, 702 (us),
Macbride & Featherstone 239 (F), 1020 (F, OH, us),
Killip & Smith 21828 (F, NY), Iltis et al. 66 (F).
ANCASH. Huaylas: Huascaran National Park, Au-
quispuquio, 3900-4000 m, Smith et al. 12104 (USM).
AREQUIPA. Arequipa: Km 18 on road between Are-
quipa and Yura, 3300 m, Solomon 2848 (MO). CUZCO.
Anta: Huanacori, El Chaccan, 3690 m, Brunei 901 (MO).
Calca: Hda. Urco, 2800 m, Vargas 1768 (GH,NY). Cuzco:
Saqsaywaman, 3600 m, Lopez 0618 (HUT); ca. 2 km S
of San Geronimo, 10 km SE of Cuzco, 3200 m, Solomon
3000 (MO). Espinar: Puerto Rosario, Rio Apurimac, 3900
m, Vargas 10103 (F). Urubamba: Ollantaytambo, 2800
m, Herrera 3399 (F), Cook & Gilbert 523, 702 (us).
HUANCAVELICA. Huancavelica: Caniorccona, 3 km
SE of Conaica, Tovar 98 (us p.p.); Peccsuca, a 5 km E
de Conaica, 3250-3280 m, Tovar 276 (us); Pana, cerca
a Conaica, 3320 m, Tovar 760 (us, USM). JUNIN. Huan-
cayo: Huancayo, 3317m, Soukup 1877 (us, USM). Tarma:
ca. 1 8 km SSE of Tarma, Rio Quishuarcaucha, Iltis et
al. 66 (F, us); Tarma, 3000-3200 m, Killip & Smith 21828
(F, NY, us); Tarma, Macbride & Featherstone 1020 (F, us).
Yauli: Banos, Capt. Wilkes s.n. (NY p.p.). LIMA. Huar-
ochiri: Matucana, 8000 ft, Macbride & Featherstone 239
(F). PUNO. Azangaro: Cuturi, entre Aropa y Azangaro,
4200 m, Aguilar 411 (MO, USM); Isla Arapa, Aguilar s.n.
(USM). Huancane: Moho, 3900 m, Aguilar 43 5 (MO, USM).
Melgar: Araranca, 4100-4300 m, Pennell 13484 (F, GH,
NY, us, USM). Puno: near Puno, Soukup 93 (F). NO EX-
ACT LOCALITY. "Perou austral," 3500 m, Weddell
4506 (F).
5. Achyrocline satureioides (Lam.) DC., Prodr. 6:
220. 1838.
Gnaphalium satureioides Lam., Encycl. 2: 747. 1788.
TYPE: Uruguay, Montevideo, 1767, P. Com-
merson s.n. (holotype, P, not seen, F neg. 37572;
isotype, G-DEL, not seen, F neg. 28707).
Perennial herbs to 2 m tall, suffruticose; stems
much-branched, ascending to decumbent, densely
lanate. Leaves sessile; blade linear to linear-lan-
ceolate, 4-10 cm long, 2-9 mm wide, apex acute.
Capitulescences glomerate, cymose-paniculate,
terminal and axillary. Capitula 4.5-5.5 mm high,
ca. 1.5 mm wide; involucres narrowly cylindrical;
outer phyllaries narrowly ovate to lanceolate, 2.5-
3.5 mm long, ca. 1 mm wide, the inner phyllaries
lanceolate to linear, 4-4.5 mm long, 0.5-1 mm
wide; pistillate florets 4(-8), the corollas 3.2-3.6
mm long; hermaphroditic florets 1-2, the corollas
3.4-3.6 mm long. Achenes ovoid, 0.6-0.7 mm long;
pappus ca. 4 mm long. Chromosome number: n
= 14 (Jansen et al., 1984).
Achyrocline satureioides is widely distributed
throughout South America in sandy soils, usually
at elevations below 2000 m. In Peru it is most
commonly found in forest clearings and disturbed
sites along the eastern slope of the Andean Cor-
dillera (500-1500 m).
It appears most closely related to A. ramosis-
sima, a smaller, alpine species, which most likely
is a high-elevation derivative of A. satureioides.
Label data from King 3257 (F) state that the
plant is used medicinally: "A preparation from
this plant is taken as a drink for kidney troubles.
The natives are acquainted with 'Foster's Pills,'
used for the same disorders, whence the name Wild
Foster." The gathering and transport of this spe-
cies for sale in local markets may explain the oc-
casional plant found outside its usual distribu-
tional range at lower elevations. In Peru, it is known
by the following common names: coronilla-sacha,
huira-huira, huira huayo, Foster-sacha.
AMAZONAS. Bongara: between Rio Utcubamba and
Shipasbamba, 4 km from Campamento Ingenio, 1520
m, Hutchison & Wright 3954 (F, MO, NY, uc). Rodriguez
de Mendoza: Mendoza, 1500 m, Woytkowski 8115 (GH,
MO). CUZCO. La Convencion: mountains between Po-
tocusi and Sahuayacu, 1000-1 100 m, Weberbauer 7946
(F,GH,MO,US). HUANUCO. Huanuco: Cotirarda to Mer-
cedes, 1875 m, Mexia 8194 (F, GH, MO, NY, us). Leoncio
Prado: Pampayacu, hacienda at mouth of Rio Chinchao,
3500 ft, Macbride 5039 (F, GH, us), Kanehira 287 (GH).
JUNIN. Tarma: Chuyquishunca, 2 km arriba de Hu-
acapistana, entre Tarma y San Ramon, 2000-2400 m,
Ferreyra 466 (USM); Chanchamayo Valley, 1200-1500
m, Schunke 365, 1074, 1536, 1622 (F). PUNO. Sandia:
along Rio Tambopata in vicinity of San Juan del Oro,
1900 m, Wasshausen & Solas 1231 (us); Sandia, 2250
m, Vargas 14800 (us). SAN MARTIN. Huallaga: cerca
a Bellavista, 250-350 m, Ferreyra 10080 (MO, USM). La-
mas: Lamas, below English Evangelical Mission, Bel-
shaw 3447 (F, GH, MO, NY, us); San Roque, 1350-1500
m, Williams 7282 (F). Moyobamba: Zepelacio, near
Moyobamba, 1200-1600 m, Klug 3257 (F, MO, NY, us).
San Martin: Tarapoto, 400 m, Schunke 6420 (F); prope
FIELDIANA: BOTANY
Tarapoto, Spruce 4174 (GH, NY). Rioja: Rioja, Woyt-
kowski6169(GH,MO,us). NO EXACT LOCALITY. 1862,
Matthews s.n. (NY p.p.).
II. Antennaria
Antennaria Gaertn., Fruct. Sem. PI. 2: 410. 1791.
TYPE: Gnaphalium dioicum L. = Antennaria
dioica (L.) Gaertn.
Dioecious or polygamo-dioecious, perennial
herbs; stems simple, ascending or erect, tomentose
or lanate. Basal leaves rosulate, oblanceolate to
spatulate, entire; cauline leaves alternate, smaller.
Capitulescences cymose-corymbose, racemose, or
glomerulate, terminal, occasionally of a solitary
head. Capitula discoid or rarely disciform; invo-
lucres ovoid or campanulate; phyllaries imbricate,
scarious, the outer gradually narrowing, the inner
prolonged into a petaloid lamina; receptacles con-
vex to plane, epaleaceous; pistillate florets with
corollas filiform, white or lilac, truncate or sub-
dentate; staminate florets with corollas tubular, 5-
lobed or 5-dentate, the anther bases sagittate, cau-
date, the terminal appendages ovate, the styles un-
divided or briefly bifid, the branches truncate.
Achenes cylindrical or ellipsoidal, rounded or sub-
compressed; pappus bristles uniseriate, scabrid,
barbellate, fused at base or free, apices clavellate
(staminate) or acute (pistillate). Chromosome
numbers: x = 14.
A genus of some 50 species distributed princi-
pally in arctic to temperate regions of the Northern
Hemisphere. Three species are known from South
America and only one from Peru. In many species,
masculine individuals are rare or completely ab-
sent, suggesting apomictic seed production.
Reference
CABRERA, A. L. 1957. Una nueva especie del genero
Antennaria (Compositae). Notas Mus. La Plata, Bot.,
19: 73-79.
1. Antennaria linearifolia Wedd., Chlor. And. 1:
150. 1856. TYPE: Peru, no exact locality, J.
Dombey or J. Pavon (syntypes probably P, not
seen). (Two syntype collections were cited by
Weddell, but no lectotype will be chosen until
the type material can be studied.) Figure 2.
Leontopodium linearifolium (Wedd.) Benth. & Hook.,
Gen. PI. 2: 303. 1873.
Gnaphalium linearifolium (Wedd.) Franchet, Bull. Soc.
Bot. France 39: 135. 1892.
Gnaphalium sedoides F. W. Klatt, Linnaea 42: 135.
1 878-79. TYPE: Peru, J. Dombey 252 (holotype,
p, not seen, F neg. 37609; isotype, GH).
Dioecious or rarely polygamo-dioecious, peren-
nial herbs, rhizomatous; stems unbranched, 3-2 1
cm tall. Leaves basal, rosulate, sessile; blade ob-
lanceolate-linear, 2-8 cm long, 1.5-3 mm wide,
the lower surface densely tomentose, midrib
prominent, the upper surface glabrescent, the mar-
gins entire. Capitulescences glomerulate, subtend-
ed by foliaceous bracts. Capitula 5-7 mm high, 3-
5 mm wide, discoid, homogamous, rarely disci-
form and heterogamous; involucres campanulate;
phyllaries 4-5-seriate, the outer ovate, 34 mm
long, ca. 2.5 mm wide, apex obtuse, the inner ob-
ovate to oblong-lanceolate, 4-5 mm long, 1-2 mm
wide, the apex prolonged into a white, petaloid
lamina; masculine capitula with 40-50 function-
ally staminate florets, the corollas narrowly tu-
bular, 3-3.5 mm long; feminine capitula with 50-
60 pistillate florets, the corollas filiform, 2.5-3 mm
long; heterogamous capitula disciform with 50-60
pistillate florets, the corollas filiform, 2.5-3 mm
long, functionally staminate florets 6-8, the co-
rollas narrowly tubular, 3.5-4 mm long. Achenes
cylindrical, 0.5-0.7 mm long, pubescent with ses-
sile, biseriate, capitate-glandular trichomes; pap-
pus bristles ca. 3.5 mm long, apices clavellate (sta-
minate) or acute (pistillate).
Antennaria linearifolia is distributed through-
out the Andean Cordillera from southern Ecuador
to Bolivia. In Peru it occurs from Amazonas to
Cuzco at elevations of 2700-4000 m.
Collections of this species are commonly an-
notated as Gnaphalium; however, the dioecious
or polygamo-dioecious reproductive system and
dimorphic pappus bristles clearly place these taxa
in Antennaria. Masculine, feminine, and occa-
sionally individuals with heterogamous capitula
are encountered within Peruvian populations.
In Peru A. linearifolia is known as champito, a
name applied to many rosulate or cespitose species
of Asteraceae.
AMAZONAS. Chachapoyas: Cerros Calla Calla, E
side, 19 km above Leimebamba on road to Balsas, 3100
m, Hutchison & Wright 5560 (F, MO, NY, us); Puma-urcu,
SE of Chachapoyas, 3100-3200 m, Wurdack 1135 (us);
Cerro Campanario NNE of Diosan, 3200-3500 m, Wur-
dack 1603 (us). ANCASH. Bolognesi: Chiquian, 3840-
DILLON & SAGASTEGUI: FLORA OF PERU. V.
FIG. 2. Antennaria linearifolia. A, habit; B, masculine capitulum; C, outer phyllary; D, inner phyllary; E, pistillate
floret; F, masculine floret; G, stamen; H, achene. (Drawn from Sagdstegui et al. 11588, HUT.)
3860 m, Ferreyra 5766 (us, USM). Huaraz: Cerro San
Cristobal, NE of Huaraz, 3800 m, Evangelista s.n. (HUT).
Huari: Huascaran National Park, 1 km below Manto
Mina, 4300 m, Smith & Buddensiek 11010 (F P . P .), 14.8
km NE of Tunel Cahuish along road between Catac and
Chavin de Huantar, 3000-4000 m, Stevens 21965 (MO).
CAJAMARCA. Cajamarca: ca. 27 km NNW of Caja-
marca on road to Hualgayoc, Pampa de Cerro Negro,
3600 m, Dillon & Molau 302 3 A (F). Celendin: ca. 57 km
NE of Cajamarca along road to Celendin, 3650 m., Dil-
10
FIELDIANA: BOTANY
Ion & Turner 1611 (F). Contumaza: above Contumaza,
2700 m, Sagdstegui & Fukushima 6098 (HUT, us); Pozo
Kuan, 3600-3800 m, Sagdstegui et al. 10086 (HUT), 12451
(HUT, MO). San Miguel: Taulis Alto (jalca), 3100 m, Sa-
gdstegui et al. 9543 (F.HUT). CUZCO. Paucartambo: Tres
Cruces, 3330-3500 m, Gentry et al. 23439 (F, MO); Co-
rihuayrachina, Callanga, 3800 m, Woytkowski 565 (MO).
Urubamba: Machu Picchu, on the Palcay side of the
Salcantay-Palcay pass, 3500 m, Peyton & Peyton 711
(MO). HUANUCO. Dos de Mayo: Probresco, 12 mi E
of Huallanca, ca. 10,500 ft, Macbride & Featherstone
2463 (F.US). Huanuco: Pillao, 2700 m, Woytkowski 34041
(F). JUNIN. Conception: 6 mi S of Mito, ca. 1 1,000 ft,
Macbride & Featherstone 1819(f, us). Huancayo: Huan-
cayo, 4000 m, Soukup 1870 (us); Laguna Huacracocha,
5000 m, Soukup 3612 (F). LA LIBERT AD. Huamachu-
co: ca. 20 km W of Huamachuco, ca. 3400 m, Dillon et
al. 2809 (F, USM); Cacana, 3300-3400 m, Ricchio & La
Rosa 3564 (us). Otuzco: Cerro Sango, Motil-Shorey,
3300-3400 m, Lopez 968 (us), 7950 (HUT); Sagdstegui
et al. 11588 (HUT, MO). Pataz: Quebrada Rangia, Taya-
bamba-Huancaspata, 3600 m, Lopez & Sagdstegui 8208
(HUT, MO, NY). Sanchez Carrion: ca. 10 km N of Laguna
Saucacocha, ca. 20 km NE of Huamachuco, ca. 3370 m,
Dillon et al. 2845 (F, MO, TEX, USM). LIMA. Chancay:
Auquimarca, 4000 m, Abdon 3271 (F). PASCO. Pasco:
Cerro de Pasco, 4135 m, Ellenberg 4123 (us). NO EX-
ACT LOCALITY. Weberbauer 7004 (F).
III. Holloa
Belloa Remy in Gay, Fl. Chil. 3: 336. 1 848. TYPE:
Lucilia chilensis Hook. & Arn. = Belloa chi-
lensis (Hook. & Arn.) Remy.
Perennial herbs, pulvinate or erect, generally
lanate or tomentose. Leaves alternate or rosulate,
rarely distichous, the margins entire. Capitulesc-
ences pseudospicate, glomerulate, or of solitary
heads, terminal and axillary. Capitula disciform;
involucres ovoid, campanulate, or narrowly cylin-
drical; phyllaries 3-5 -seriate, imbricate, scarious,
hyaline at margin, concave, stramineous, the inner
gradually longer; receptacles plane, naked, alve-
olate; pistillate florets 1-6-seriate, the corollas fi-
liform, lacerate, the styles exserted; hermaphro-
ditic florets (1-)10-15, the corollas narrowly
tubular, 5-lobed, the anther bases caudate, the ter-
minal appendages ovate to oblong, the style branch
apices obtuse or rounded, dorsal surfaces papil-
lose. Achenes obovoid or ellipsoid, pubescent with
biseriate, multicellular, capitate-glandular trich-
omes; pappus bristles 40-80, scabrid, white to lu-
tescent, the bases fused, deciduous together as a
unit, apical cells acute or rounded. Chromosome
number: n = 12 (Fernandez Casas & Fernandez
Piqueras, 1981).
Belloa consists of 1 8 species distributed in high-
elevation habitats throughout the Andes from
Venezuela to Chile and Argentina. Peru appears
to be a center of diversity for the genus, with eight
species represented.
Freire (1986) expanded the generic limits of Lu-
cilia to include all taxa previously accorded to
Belloa (Cabrera, 1958; Sagdstegui & Dillon, 1985)
with the statement that the presence or absence of
trichomes on the achenes was not sufficient for
recognition of the genus. She did recognize the
monophyletic nature of Belloa and chose to place
its component species into section Lucilioides DC.
She subsequently (Freire, 1987) continued her as-
sault on the genus by placing two recently de-
scribed species (Sagastegui & Dillon, 1985) into
synonymy (see discussion under individual spe-
cies). We have chosen to maintain Belloa and re-
main in agreement with Cabrera (1958) and Ariza
(1989), who both recognized the genus as a dis-
tinct, easily distinguishable, entity. In addition to
the strikingly different pubescence of the achenes,
several other important characteristics separate
these two genera, including the apices of the style
branches of hermaphroditic florets, pappus char-
acteristics, and overall distribution and ecology.
Initial cytological evidence also points to funda-
mental differences between these two genera: n =
1 2 for Belloa punae (Cabr.) Cabr., the only Belloa
counted thus far, and n= 14 for all Lucilia species
counted (Freire, 1986b).
Belloa is recognized by the following suite of
characteristics: heterogamous capitula, pappus
bristles fused at the base, style branches of her-
maphroditic florets rounded or obtuse, achenes
glandular with biseriate, multicellular, capitate-
glandular trichomes, and probable chromosome
base of n = 12. Suites of similar characters have
been used to establish the monotypic genus Nov-
enia (Freire, 1 986a). In our view, Belloa is perhaps
more closely related to Gamochaeta than it is to
Lucilia. A few highly reduced Belloa species ap-
proach Mniodes in general habit; however, that
genus is strictly dioecious.
A more detailed analysis and discussion of these
and other differences will be published in a future
work dealing with generic relationships among
South American Inuleae.
References
ARIZA-ESPINAR, L. 1 989. Las especies centroargentinas
de Belloa (Asteraceae). Kurtziana, 20: 173-179.
CABRERA, A. L. 1958. El genero Belloa Remy. Bol. Soc.
Argent. Bot., 7: 79-85.
DILLON & SAGASTEGUI: FLORA OF PERU. V.
11
. 1978. Compositae. Flora de la Provincia de
Jujuy. Colecc. Cient., Inst. Natl. Teen. Agropec., 13:
294-301.
FERNANDEZ CASAS, J., AND J. FERNANDEZ PIQUERAS.
1981. Estudio cariologico de algunas plantas bolivi-
anas. Anales Jard. Hot. Madrid, 38: 149-152.
FREIRE, S. E. 1 986a. Novenia: Nuevo genero de Inuleae
(Compositae). Bol. Soc. Argent. Bot., 24: 295-304.
. 1986b. Revision del genero Lucilia (Compos-
itae, Inuleae). Darwiniana, 27: 431-490.
1987. Nuevos sinonimos y una nueva com-
bination en el genero Lucilia (Compositae, Inuleae).
Darwiniana, 28: 409-41 1.
SAGASTEGUI-ALVA, A., AND M. O. DILLON. 1985. New
species and combinations in Belloa (Inuleae-Astera-
ceae). Phytologia, 58: 392-400.
Key to Species of Belloa
1 . Plants with stems erect or ascending, not cespitose, pulvinate, or acaulescent 2
2. Foliaceous stems compressed; leaves distichous, orbicular to suborbicular, folded upward, subam-
plexicaul B. plicatifolia
2. Foliaceous stems terete; leaves alternate or rosulate, oblanceolate to linear-lanceolate, plane, at-
tenuate, and sessile 3
3. Capitulescences glomerulate with 2-5 capitula sessile in upper leaf axils . . B. lopezmirandae
3. Capitulescences pseudospicate, bracteate B. turned
1 . Plants with stems prostrate to decumbent, generally cespitose or pulvinate, or rarely acaulescent with
rosulate basal leaves 4
4. Acaulescent perennial herbs; leaves all basal, rosulate; capitula
solitary, sessile or shortly pedunculate 5
5. Leaves oblanceolate, 2-5 cm long, 6-10 mm wide, upper surfaces usually glabrescent
B. longifolia
5. Leaves spatulate, 2-3.5 cm long, 24 mm wide, both surfaces densely silvery-tomentose . . .
B. spathulifolia
4. Caulescent, prostrate to decumbent, cespitose or pulvinate perennial herbs; leaves basal, rosulate
and/or caulescent; capitula in glomerules or solitary, usually sessile 6
6. Compact, cespitose habit; leaves 4-10 mm long 7
7. Capitula solitary; leaves 44.5 mm long B. schultzii
1 . Capitula in glomerules; leaves 6-10 mm long B. piptolepis
6. Branched, loosely cespitose, prostrate habit; leaves 10-50 mm long 8
8. Leaves alternate, congested, obovate to spatulate, 10-20 mm long; capitula solitary
B. pickeringii
8. Leaves rosulate, oblong-linear to spatulate, (10-)20-50 mm long; Capitulescences pseudo-
spicate, capitula sessile in upper leaf axils B. subspicata
1 . Belloa longifolia (Cuatr. & Aristeg.) Sagast. &
Dillon, Phytologia 58: 396. 1985. Figure 3.
Lucilia longifolia Cuatr. & Aristeg., Fl Venezuela 10:
367. 1964. TYPE: Venezuela, Edo. Merida, cam-
ino a Pico Bolivar, 1 5 km al sudeste de Merida,
4100-4300 m, Oct 1953, E. L. Little 15725 (ho-
lotype, VEN, not seen).
Cespitose, perennial herbs; rhizomes oblique to
horizontal; roots filiform. Leaves rosulate, sessile;
blade oblanceolate, 2-5 cm long, 6-10 mm wide,
marcescent, base attenuate to a winged petiole,
apex obtuse, mucronate, the margins entire, lower
surface silvery-tomentose, upper surface densely
lanate. Capitulescences of solitary heads, terminal,
sessile or pedunculate, the peduncles to 5 mm long.
Capitula 7.5-10 mm high, 4.5-8 mm wide; in-
volucres cylindrical to narrowly campanulate;
phyllaries 4-5-seriate, the outer ovate, 4-5.5 mm
long, 2.5-3 mm wide, apex obtuse, dorsally la-
nuginous, the inner linear, 7.5-10 mm long, 2.5-
3 mm wide, apex obtuse, glabrous; pistillate florets
ca. 80, the corollas 6-7 mm long; hermaphroditic
florets ca. 1 2, the corollas ca. 6 mm long. Achenes
obovate to oblong, 1-1.2 mm long, glandular; pap-
pus bristles 6-8 mm long, lutescent.
Belloa longifolia is distributed from Venezuela
to southern Peru. In Peru, it is known from jalca
and puna formations (3600-4000 m) in the De-
partments of Ancash, Cajamarca, Cuzco, and La
Libertad.
ANCASH. Huaraz: Parque Nacional de Huascaran,
Quebrada Alpamayo, 4600-4750 m, Smith et al. 9733
12
F1ELDIANA: BOTANY
H
FIG. 3. Belloa longifolia. A, habit; B, capitulum; C, pistillate floret; D, hermaphroditic floret; E, achene. (Drawn
from Sagdstegui et al. 10060, HUT). Belloa plicatifolia. F, habit; G, capitulum; H, pistillate floret; I, hermaphroditic
floret; J, style branches of pistillate floret; K, achene. (Drawn from Sagdstegui et al. 101 17, HUT.)
(HUT, MO, USM), Smith 10376 (HUT, MO). Huari: Huas-
caran National Park, Quebrada Pucaraju, a lateral valley
to Quebrada Rurichinchay, 3900-4200 m, Smith et al.
12708 (F, MO). Santa: Laguna Capao, 4350 m, Mostacero
et al. 1932 (F, HUT). AYACUCHO. Prov. Unknown: Au-
casime 10 (USM). CAJAMARCA. Contumaza: alrede-
dores del Pozo Kuan, 3600-3800 m, Sagdstegui et al.
10060 (F, HUT); Jalca del Pozo Kuan, 4000 m, Sagdstegui
DILLON & SAGASTEGUI: FLORA OF PERU. V.
13
et al. 10763 (F, HUT). San Miguel: Taulis Alto (jalca),
3100 m, Sagdstegui et al. 9546 (F, HUT). CUZCO. Uru-
bamba: Machupicchu, along Inca Trail in the pass of
Huarmihuanusca, 4205 m, Peyton & Peyton 279 (MO).
JUNIN. Yauli: near Atocsayco, ca. 4200 m, Pearsall837
(F). Prov. Unknown: Patarcocha, 4300 m, Aguilar s.n.
(MO, USM). LA LIBERT AD. Pataz: Paso de Alaska (Re-
tamas-La Paccha), 4000 m, Lopez & Sagdstegui 3560
(HUT, us). Santiago de Chuco: alrededores de Laguna El
Toro (Jalca de Quiruvilca), 4000 m, Sagdstegui & Bernal
3019 (HUT, us). Chota: Motil-Shorey (jalca), 3200 m,
Sagdstegui et al. 11697 (F, HUT); Jalca de Coipin, 4000
m, Sagdstegui et al. 11981 (F, HUT). LAMBAYEQUE.
Ferrenafe: Sinchigual-Laguna Tembladera, 3 100 m, Sa-
gdstegui et al. 12853 (F, HUT, MO, K).
2. Belloa lopezmirandae Cabr., Bol. Soc. Argent.
Dot. 7: 83. 1958. TYPE: Peru, Dept. La Lib-
ertad, Prov. Otuzco, Agallpampa, 3100 m, 22
May 1952, A. Lopez M. 858 (holotype, LP;
isotype, HUT).
Lucilia lopezmirandae (Cabr.) Freire, Darwiniana, 28:
409. 1987.
Perennial herbs to 20 cm tall; stems un-
branched, ascending, cylindrical, leafy to the api-
ces. Leaves alternate, sessile; blade oblanceolate,
1-3 cm long, 3-5 mm wide, base attenuate, apex
acute, mucronate, lower surface tomentose, upper
surface lanuginous or glabrescent. Capitulescences
glomerulate with 2-5 heads sessile in upper leaf
axils. Ca pi tula 7-7.5 mm high, 1-1.5 mm wide;
involucres cylindrical; phyllaries 12-15, ca. 5 -se-
riate, the outer ovate, 34 mm long, 1-1.5 mm
wide, dorsally lanuginous, apex acute, the inner
linear, ca. 7 mm long, ca. 1 mm wide, glabrous,
apex acute to subacute; pistillate florets ca. 20, the
corollas 6.5-7 mm long; hermaphroditic florets 1-
2, the corollas 5-6 mm long. Achenes obovoid,
ca. 1 mm long, brown, glandular; pappus bristles
5.5-6 mm long, white.
Belloa lopezmirandae is known from two col-
lections from open grassy sites in the region of the
type locality (3100 m). Freire (1987) placed B.
turneri Sagast. & Dillon into the synonymy of this
species and chose to dismiss the numerous qual-
itative and quantitative characteristics that clearly
separate these two entities.
LA LIBERT AD. Otuzco: Agallpampa, 3 100 m, Lopez
1864 (F, HUT).
3. Belloa pickeringii (A. Gray) Sagast. & Dillon,
Phytologia 58: 396. 1985.
Lucilia pickeringii A. Gray, Proc. Amer. Acad. Arts
5: 138.1862. TYPE: Peru, Dept. Junin, Prov.
Yauli, Banos-Alpamarca, Capt. Wilkes s.n. (ho-
lotype, GH; isotype, us).
Lucilia pickeringii var. #.? minor A.. Gray, Proc. Amer.
Acad. Arts 5: 138. 1862. TYPE: Peru, Dept. and
Prov. Unknown, Casa Cancha, Capt. Wilkes s.n.
(holotype, GH p.p.).
Prostrate perennial herbs to 5 cm tall; stems 2-
8 cm long, branched, ascending to decumbent.
Leaves alternate, sessile, congested; blade obovate
to spatulate, 1-2 cm long, 3-6 mm wide, base
attenuate, apex rounded, both surfaces densely la-
nate, incanous. Capitulescences usually of solitary
heads, terminal, sessile, rarely with 2-4 heads in
terminal glomerules. Capitula 8-9 mm high, 3-4
mm wide; involucres cylindrical; phyllaries ca. 24,
4-5 -seriate, the outer ovate, 4-5 mm long, 2-3
mm wide, dorsally lanuginous, acute, the inner
linear, 7-9 mm long, 1-2 mm wide, glabrous, sub-
acute to obtuse; pistillate florets 40-60, the corol-
las 4-5 mm long; hermaphroditic florets ca. 10,
the corollas 34 mm long. Achenes (immature)
oblong, 0.8-1 mm long, brownish, glandular; pap-
pus bristles ca. 5 mm long, white.
Belloa pickeringii is found infrequently in high-
elevation puna formations (3600 m) from central
to southern Peru.
CUZCO. Cuzco: Cuzco, 3600 m, Herrera 2397 (F).
JUNIN. Tarma: 25 km W of Tarma, 4100 m, Hutchison
659 (uc). LIMA. Huarochiri: Rio Blanco, 12,000 ft,
Macbride & Featherstone 705 (F, us).
4. Belloa piptolepis (Wedd.) Cabr., Bol. Soc. Ar-
gent. Dot. 7: 81. 1958.
Merope piptolepis Wedd., Chlor. And. 1: 162. 1856.
SYNTYPES: Peru, Dept. Puno, Maravillas, H. A.
Weddell 4514 (lectotype, P, designated by Ca-
brera, 1978). Bolivia, Dept. Potosi: "aux environs
des lagunas," A. C. V. d'Orbigny 1371 (paralec-
totype, P, not seen, fragment, F; isoparalectotype,
G, not seen, F neg. 2861 1).
Lucilia piptolepis Wedd., Chlor. And. 1, t. 26 B, 1855.
Gnaphalium piptolepis (Wedd.) Griseb., Abh. Konigl.
Ges. Wiss. Gottingen. 24: 186. 1879.
Compact cespitose, perennial herbs to 5 cm tall;
rhizomes oblique, roots filiform; stems 540 cm
long, decumbent, tomentose to glabrate. Leaves
rosulate, sessile; blade obovate-spatulate to obo-
vate-oblong, 610 mm long, 34 mm wide, mar-
cescent, base attenuate, apex obtuse, mucronulate,
both surfaces tomentose, incanous. Capitules-
14
FIELDIANA: BOTANY
cences glomerulate, sessile, terminal, occasionally
of solitary heads. Capitula ca. 6 mm high, ca. 4
mm wide; involucres narrowly campanulate; phyl-
laries ca. 18, ca. 3-seriate, the outer ovate-oblong,
34 mm long, 1-1.5 mm wide, lanuginous, obtuse,
the inner linear, ca. 6 mm long, ca. 1 mm wide,
glabrous, obtuse; pistillate florets 18-20, the co-
rollas ca. 4 mm long; hermaphroditic florets 2-5,
the corollas 3.5-4 mm long. Achenes ellipsoid,
0.7-0.8 mm long, brown, glandular; pappus bris-
tles ca. 5 mm long, white.
Belloa piptolepis is distributed from Venezuela
to northwestern Argentina. In Peru it occupies a
wide range of habitats within puna and jalca for-
mations and is normally found above 3500 m.
The overall habit varies considerably with the
smallest individuals found in southern Peru, and
the larger, to the north. Belloa piptolepis most
closely resembles B. subspicata, but the former
generally has a more compact habit, smaller leaves,
slightly broader capitula, and smaller achenes. In
addition, the capitulescence of B. piptolepis is
composed of terminal capitula subtended by fo-
liaceous bracts, whereas that of B. subspicata is
pseudospicate, with axillary capitula subtended by
the upper leaves. Individuals from northern Peru
(e.g., Dillon & Skillman 4127) often have a more
open habit that approaches B. subspicata, but con-
form in all other characters to the species delin-
eation presented here. Further sampling in this
region will be necessary to assess variability in
habit.
ANCASH. Bolognesi: Chiquian, 3840-3860 m, Fer-
reyra 5768 (us, USM). Carhuaz: Huascaran National Park,
Quebrada Ishinca, 4730-4930 m, Smith et al. 9459 (MO,
USM); Quebrada Ulta, 4400-4600 m, Smith 11354 (F,
MO). Huaraz: Huascaran National Park, Quebrada Lla-
ca, 4400-4600 m, Smith et al. 10806 (F, MO). Huari:
Tunel Kahuish, 4250 m, Lopez et al. 7569 (HUT, MO);
Huascaran National Park, Quebrada Rurichinchay be-
tween Mina Esparta at Quebrada Pachachaca, 3700-4060
m, Smith et al. 12658 (F, MO). AYACUCHO. Huanta:
Pampalca, between Huanta and Rio Apurimac, ca. 3200
m, Killip & Smith 23265 (NY). Santa: Laguna Capao,
4400 m, Mostacero et al. 1942 (F, HUT, MO). CAJA-
MARCA. Chota: Laguna Yahuarococha, arriba de In-
cahuasi, 3600 m, Sagdstegui et al. 12906 (F, HUT, K, LP,
MO). Contumaza: Jalca del Pozo Kuan, 4000 m, Sagds-
tegui et al. 10764, 10791 (F, HUT, MO). CUZCO. Cuzco:
5 km N of Cuzco, 3700 m, West 7165 (GH.UC). JUNIN.
Junin: between Tarma and Jauja, ca. 4500 m, Killip &
Smith 21962 (NY). Yauli: Yauli, ca. 13,500 ft, Macbride
6 Featherstone 931 (F,GH). Prov. Unknown: Patarcocha,
4300 m, Aguilar s.n. (USM). LA LIBERTAD. Pataz:
Huancaspata-Tayabamba, 3900 m, Lopez & Sagdstegui
8267 (HUT, MO). Santiago de Chuco: Jalca de Corpm,
4000 m, Sagdstegui et al. 11980 (F, HUT, MO, NY, us).
LAMBAYEQUE. Ferrenafe: ca. 7 km NW of Incahuasi;
near Cerro Punamachay on trail to Laguna Hualtaco,
3300-3550 m, Dillon & Skillman 4127 (F). MOQUE-
GUA. Mariscal Nieto: ca. 77 km NE of Moquegua on
road to Carumus (Km 80), ca. 4050 m, Dillon et al. 4812
(F.USM). PUNO. Azangaro: Arapa, 3900 m, Aguilar s.n.
(USM p.p.); Arapa, Aguilar 130 (MO p.p.); Tequena, 20
km N de Arapa, Aguilar 418 (USM p.p.). Huancane: Moho,
4000 m, Aguilar s.n. (USM); Granja Salcedo, Canon Vis-
cachani, 4500 m, Mexia 4262 (GH.MO.UC); Melgar: Chu-
quibambilla, 3900-4000 m, Pennell 13368 (GH, F, NY);
Araranca, 4100-4300 m, Pennell 13437 (F, NY). Puno:
Amantani, 3900 m, Aguilar 427 (MO p.p., USM).
5. Belloa plicatifolia Sagast. & Dillon, Phytologia
58: 394. 1985. TYPE: Peru, Dept. Cajamarca,
Prov. Contumaza, Cascabamba, arriba de
Contumaza, ladera, 3200 m, 14 Jun 1981, A.
Sagdstegui A., E. Garcia A., S. Lopez M. &
J. Mostacero L. 70777(holotype, HUT; isotype
F, HUT, K, MO). Figure 3.
Lucilia plicatifolia (Sagast. & Dillon) Freire, Darwin-
iana, 28:411. 1987.
Perennial herbs to 5-20 cm tall; stems branched,
erect to ascending, leafy to the apex. Leaves alter-
nate, distichous, sessile; blade orbicular to subor-
bicular, 5-14 mm long, 3-7 mm wide, marcescent,
folded, base attenuate, subamplexicaul, apex
rounded, both surfaces densely lanate, the margins
entire. Capitulescences of solitary heads in upper
leaf axils, subsessile. Capitula (6-)7-8(-9) mm high,
3-4 mm wide; involucres cylindrical; phyllaries
15-20, 4-5-seriate, the outer ovate, 4-6 mm long,
34 mm wide, apex subacute, dorsally lanuginous,
the inner linear to lanceolate, 7-8 mm long, 1-1.5
mm wide, apex acute, glabrous; pistillate florets
12-14, the corollas 4-5 mm long; hermaphroditic
florets ca. 1 0, the corollas 4-5 mm long. Achenes
obovate to oblong, 1-1.5 mm long, brown, glan-
dular; pappus bristles 5-6 mm long, white.
Belloa plicatifolia is found frequently among
sheltered rocky sites in jalca formations of north-
ern Peru (Departments of Cajamarca and La Lib-
ertad, 3000-3500 m).
This species is morphologically distinct and has
no apparent close relatives. Its erect lanate branch-
es with folded, distichous leaves give the stems a
flattened appearance unknown elsewhere in the
genus.
CAJAMARCA. Cajabamba: Cajabamba-Luchubam-
ba, 3800 m, Sagdstegui et al. 11199 (F, HUT, K, MO, NY,
DILLON & SAGASTEGUI: FLORA OF PERU. V.
15
USM). Cajamarca: Cumbemayo, 3200 m, Sagdstegui &
Tellez 12686 (F, HUT, MO). Contumaza: Cascabamba, 3050
m, Sagdstegui et al. 9022 (F, HUT, MO); Cascabamba,
3100 m, Sagdstegui et al. 10010 (F, HUT, K, MO); Casca-
bamba, 3000 m, Sagdstegui et al. 10719 (F, HUT, K, MO).
LA LIBERTAD. Otuzco: Salpo, Cerro Ragache, 3500
m, Sagdstegui et al. 11631 (F, HUT, K, MO).
palca-Marcapomacocha, ca. 1 5,000 ft, Sounders 807 (NY).
MOQUEGUA. Mariscal Nieto: between Torata and
Carumas, Km 75-76 from Moquegua to Puno, ca. 3600
m, Dillon et al. 3 346 A (F, HUT, K, MO, NY, us, USM). PUNO.
Puno: San Antonio de Esquilache, 15,500 ft, Stafford
746 (F). Prov. Unknown: Casa Cancha, Wilkes s.n. (GH
p.p., NY).
6. Holloa schultzii (Wedd.) Cabr., Re vista Invest.
Agric. 11:404. 1957.
Merope schultzii Wedd., Chlor. And. 1: 163. 1856.
TYPE: Peru, Dept. Puno, Prov. Carabaya, Aya-
pata, W. Lechler 1984 (holotype, P, not seen, F
neg. 37608).
Compact, cespitose, perennial herbs; stems
much-branched, 2-3 cm long, compacted. Leaves
sessile, densely imbricate; blade obovate, 44.5
mm long, 2-2.5 mm wide, marcescent, base at-
tenuate, subamplexicaul, apex rounded, both sur-
faces densely lanate, gray. Capitulescences of sol-
itary heads, terminal, sessile. Capitula 4-^.5 mm
high, 1.5-2 mm wide; involucres cylindrical; phyl-
laries ca. 18, 4-seriate, the outer ovate, 2-3 mm
long, 1-1.5 mm wide, dorsally lanuginous, acute,
the inner linear, 4-4.5 mm long, 1-1.5 mm wide,
glabrous, apex subacute, purplish; pistillate florets
9-1 1, the corollas 3-3.5 mm long; hermaphroditic
florets 1-3, the corollas 2.5-3 mm long. Achenes
(immature) oblong, 0.6-0.8 mm long, brown,
sparsely glandular; pappus bristles ca. 4 mm long,
white.
Belloa schultzii is distributed from southern Peru
to northern Chile and adjacent northwestern Ar-
gentina in dry puna habitats usually above 4000
m. It is distinctive among the Peruvian members
of the genus in possessing a tightly compact habit,
densely imbricate leaves, capitula only 44.5 mm
high, and sparasely glandular achenes. Superfi-
cially, its overall habit approaches that ofMniodes,
but it is readily distinguished by its heterogamous
capitula.
Freire (1987) placed Mniodes cerratei Ferreyra
under the synonymy of B. schultizii (as Lucilid).
Sagastegui and Dillon (1985) recognized the type
as a species of Belloa and will follow Freire's sug-
gestion until further material can be examined.
7. Belloa spathulifolia Sagast. & Dillon, Phyto-
logia 58: 394. 1985. TYPE: Peru, Dept. La
Libertad, Prov. Santiago de Chuco, entre
Chota Motil y Shorey, jalca, 3200 m, 6 Dec
1984, A. Sagdstegui A., J. Mostacero L. & M.
Diestra Q. 11695 (holotype, HUT; isotypes, F,
MO, NY). Figure 4.
Cespitose, perennial herbs to 3 cm tall; rhizomes
oblique, roots filiform. Leaves rosulate, sessile;
blade spatulate to oblanceolate-spatulate, 2-3.5 cm
long, 2-4 mm wide, coriaceous, marcescent, the
base expanded and partially sheathing the stem,
both surfaces densely silvery-tomentose, the mar-
gins entire. Capitulescences 2-3 -headed, sessile
glomerules or, rarely, of solitary heads. Capitula
7-8 mm high, ca. 5 mm wide; involucres narrowly
campanulate; phyllaries ca. 24, 4-seriate, the outer
ovate, ca. 4 mm long, ca. 2 mm wide, lanuginous,
apex obtuse, the inner linear-oblong, 7-8 mm long,
ca. 2 mm wide, glabrous, apex obtuse; pistillate
florets 15-16, the corollas ca. 4 mm long, the style
branches exserted; hermaphroditic florets ca. 10,
the corollas ca. 5 mm long. Achenes obovate to
oblong, ca. 1 mm long, brown, glandular; pappus
bristles ca. 4 mm long, white.
Belloa spathulifolia is known only from the type
locality in open spaces between clumps of Stipa
ichu within the jalca formation (ca. 3200 m).
Among the Peruvian species, it most closely re-
sembles B. longifolia with its cespitose, rosulate
habit; however, the latter has much wider leaves
(6-10 mm) and larger capitula with more pistillate
florets (ca. 80). Freire (1987) placed B. spathuli-
folia under the synonymy ofLucilia (Belloa) san-
tanica (Cabr.) Freire, choosing to ignore the dif-
ferences in habit, capitulescence, floral number,
and ecological and geographic preferences (Sagas-
tegui & Dillon, 1985).
AREQUIPA. Arequipa: Nevado de Chachani, 4300-
4600, Pennell 13310, 13311 (F.NY); Pichupichu, 13,500
ft, Stafford 697 (F). CUZCO. Quispicanchis: Auzangate,
4300-4600, Rauh & Hirsch PI 190 (NY). JUNIN. Yauli:
Tucto, cerca a Morococha, 4800-4900 m, Weberling
5915 (USM). LIMA. Huarochiri: ca. 1 km from Casa-
Belloa subspicata Wedd., Chlor. And. 1: 159.
1855. TYPE: Bolivia, Dept. Potosi, "aux en-
virons de Potosi," A. C. V. d'Orbigny 1373,
1374 (holotype, P, not seen, F neg. 38022).
16
FIELDIANA: BOTANY
FIG. 4. Belloa spathulifolia. A, habit; B, leaf (underside); C, capitulum; D, outer phyllary; E, inner phyllary; F,
pistillate floret; G, hermaphroditic floret; H, anther; I, style branches of pistillate floret; J, achene. (Drawn from
Sagdstegui et al. 1 1695, HUT.)
Prostrate, perennial herbs to 5 cm tall; roots
fibrous; stems numerous from caudex, 2-7 cm long,
prostrate, decumbent, or ascending, lanulose. Bas-
al leaves sessile, rosulate, oblong-linear to spatu-
late-oblong, (l-)2-5 cm long, 2-7 mm wide, base
attenuate, apex acute to obtuse, lower surface
densely tomentose, upper surface lanuginous, the
cauline leaves gradually smaller. Capitulescences
DILLON & SAGASTEGUI: FLORA OF PERU. V.
17
pseudospicate, heads sessile in upper leaf axils or,
rarely, of terminal, solitary heads. Capitula 6-8
mm high, 2-3 mm wide; involucres cylindrical;
phyllaries ca. 14, 4-seriate, the outer ovate, 34
mm long, 1-1.5 mm wide, lanuginous, acute to
subacute, the inner oblong-linear, 7-8 mm long,
1-1.5 mm wide, glabrous, acute to obtuse, brown-
ish; pistillate florets 14-20, the corollas 4-5 mm
long; hermaphroditic florets 3-6, the corollas ca.
5 mm long. Achenes oblong, ca. 1 mm long,
brownish, glandular, pappus bristles ca. 6 mm long,
white.
Belloa subspicata is distributed from northern
Peru to northwestern Argentina. Within its range
this species is polymorphic for leaf shape: collec-
tions from the north usually have spatulate to ob-
long leaves with obtuse to rounded apices (rarely
acute), and those from the south have more ob-
long-linear leaves and acute apices. These morphs
may prove to be worthy of taxonomic recognition;
however, further collections and field observations
are needed.
This species most closely resembles Belloa pip-
tolepis; however, it may be separated by the form-
er's larger leaves, cylindrical involucres, and more
pseudospicate capitulescences. The overall size of
the achenes varies considerably, but those in B.
piptolepis tend to be smaller than those of B. sub-
spicata.
CAJAMARCA. Contumaza: El Granero, 2800 m, Sa-
gdstegui & Lopez 10640 (F, HUT, MO). CUZCO. Canas:
San Andres de Checca, 3870 m, Vargas 11002 (F, uc),
Vargas 1479 (GH p.p.). HUANUCO. Huanuco: San Car-
los Mines, estacion 6 mi W of Huallanca, Macbride &
Featherstone 2467 (F). JUNIN. Jauja: Muquiyauyo, Rio
Mantaro, 3500 m, s.d. (F.USM). Tarma: road from Huan-
cayo to Tarma, 3000-3300 m, Sullivan, et al. 1043 (F,
MO). LA LIBERTAD. Santiago de Chuco: Laguna El
Toro, 4 100 m, Sagdstegui 3020 (HUT, us). LIMA. Huar-
ochiri: Viso, ca. 9000 ft, Macbride & Featherstone 614,
615 (F); Rio Blanco, ca. 12,000 ft, Macbride & Feath-
erstone 704, 741 (F), 742 (F, us). PUNO. Carabaya: be-
tween Santa Rosa and Ayaviri; ca. 3500 m, Dillon et al.
1070 (F, MO, USM). Melgar: Chuquibambilla, 3900-4000
m, Pennell 13369 (F); Araranca, 4100-4300 m, Pennell
13471 (F). NO EXACT UOCALIT.Ellenberg613(usM).
Perennial herbs to 20 cm tall; stems erect, un-
branched, cylindrical, densely sericeous-lanose.
Basal leaves rosulate, sessile; blade oblong-lance-
olate to linear-lanceolate, 2-4 cm long, 3-5 mm
wide, coriaceous, marcescent, discolorous, base at-
tenuate, apex acute, mucronulate, parallel-nerved,
lower surface silvery-tomentose, upper surface la-
nuginous; cauline leaves alternate, sessile; blade
linear to lanceolate, 1-3 cm long, 1.5-3 mm wide.
Capitulescences spiciform, bracteate. Capitula 5-
6 mm high, 5 mm wide; involucres campanulate;
phyllaries ca. 25, ca. 4-seriate, the outer ovate,
3.5-4.5 mm long, 2.5-3.5 mm wide, lanuginous,
apex acute, the inner linear to oblong-linear, 5-6
mm long, 1-2 mm wide, apex acute, glabrous;
pistillate florets 2540, the corollas ca. 3.5 mm
long; hermaphroditic florets (7-) 10-12, the corolla
3-3.5 mm long. Achenes oblong, ca. 1 mm long,
brownish, glandular; pappus bristles ca. 4 mm long,
white.
Belloa turneri occurs frequently in open areas
within jalca formations in northern Peru (De-
partments of Ancash and Cajamarca, 3100-3800
m) and southern Ecuador (Province of Loja, 2400-
2600 m).
This species is distinctive within the genus,
bearing rosulate basal leaves and robust, erect, spi-
ciform capitulescences. It most closely resembles
B. lopezmirandae Cabr. of Department La Lib-
ertad; however, the latter has smaller capitula with
cylindrical involucres and fewer florets (ca. 20 pis-
tillate, 1-2 hermaphroditic). It is known by the
local name of champito in Ancash.
ANCASH. Huaraz: Cerro San Cristobal, 3800 m,
Evangelista s. n. (F, HUT, MO); Huari: Huascaran National
Park, passing Manto Mina, 4.8 km from Catac-Chavin
road, 4410 m, Smith & Buddensiek 10988 (MO); 1 km
below Manto Mina, 4300 m, Smith & Buddensiek 11011
p.p. (F). CAJAMARCA. Contumaza: Pampa de la Sal,
3500 m, Sagdstegui et al. 10731 (F, HUT, MO); San Mi-
guel: Taulis Alto (jalca), 3100 m, Sagdstegui et al. 9547
(F, HUT, MO). LA LIBERTAD. Santiago de Chuco: Chota
(Motil-Shorey), 3200 m, Sagdstegui et al. 11693 (F.HUT,
MO).
9. Belloa turneri Sagast. & Dillon, Phytologia 58:
392. 1985. TYPE: Peru. Dept. Cajamarca.
Prov. Contumaza: alrededores del Pozo Kuan,
ladera, 3600-3800 m, 13 Jun 1981, A. Sa-
gdstegui A., E. Garcia A., S. Lopez M. & J.
MostaceroL. 10087 (holotype, HUT; isotypes,
F, HUT, MO, TEX). Figure 5.
IV. Blumea
Blumea DC, Arch. Bot. (Paris) 2: 514. 1833,nom.
cons. TYPE: Conyza balsamifera L. = Blu-
mea balsamifera (L.) DC.
Annual or perennial or rarely biennial herbs or
shrubs; stems erect, ascending or occasionally pro-
18
FIELDIANA: BOTANY
B
FIG. 5. Belloa turneri. A, habit; B, leaf (underside); C, capitulum; D, outer phyllary; E, inner phyllary; F, pistillate
floret; G, hermaphroditic floret; H, style branches of hermaphroditic floret; I, achene. (Drawn from Sagdstegui et al.
10087, HUT.)
cumbent or climbing. Leaves alternate, simple;
petiole wanting or short, sometimes decurrent on
the stem; blade base rounded, amplexicaulous, or
attenuate, often glandular-pubescent, the margins
entire, toothed, or pinnately or runcinately lobed.
Capitulescences paniculate or of solitary heads
(rarely glomerulate), terminal or axillary, sessile
or pedunculate, bracteolate or not. Capitula dis-
ciform, 3-18 mm in diameter; involucres hemi-
spherical to cylindrical; phyllaries imbricate, mul-
DILLON & SAGASTEGUI: FLORA OF PERU. V.
19
tiseriate, dorsally pubescent, the margins scarious;
receptacles flat or convex, epaleaceous, puberulent
to pilose; pistillate florets numerous, the corollas
filiform, mostly yellowish white, apex 2-4-lobed
or bilabiate, the style sometimes conspicuously
exserted; hermaphroditic florets fertile or func-
tionally staminate, the corollas tubular, the limb
slightly expanded, (4-)5-lobed, the anther bases
sagittate, caudate, the terminal appendages vari-
ously shaped, the styles bifid, the branches slender,
strigose to papillose. Achenes small, brown, ob-
long, 5-10-ribbed, plump, terete or obscurely 4-
angled; pappus bristles scabrid, barbellate, uni-
seriate. Chromosome numbers: n = 9, 10, 11, 18,
22, 27.
Blumea is a tropical and subtropical genus con-
taining some 75 species, from Africa, Asia, Aus-
tralia, and the Pacific Islands. Only the following
species is known from the New World. Blumea is
placed within the Plucheinae, which is character-
ized by alternate leaves, heterogamous capitula
(either ligulate or disciform) with predominately
yellow corollas, and sagittate or caudate anthers
(Randeria, 1960). Its nearest Peruvian relative is
Pluchea, a genus possessing a more shrubby habit,
corymbose capitulescences, more coriaceous phyl-
laries, and functionally staminate central florets
with undivided styles.
References
BADILLO, V. M. 1974. Blumea viscosa y Piptocarpha
cuatrecasiana, dos nuevas combinaciones en Com-
positae. Revista Fac. Agron. (Maracay), 7: 9-16.
MCVAUGH, R. 1972. Nomenclatural and taxonomic
notes on Mexican Compositae. Rhodora, 54: 495-5 1 6.
RANDERIA, A. J. 1960. The composite genus Blumea,
a taxonomic revision. Blumea, 10: 176-317.
1. Blumea viscosa (Miller) Badillo, Revista Fac.
Agron. (Maracay) 7(3): 9. 1974. Figure 6.
Conyza viscosa Miller, Gard. Diet., ed. 8. 1768. TYPE:
Mexico, Veracruz, W. Houston s.n. (holotype, BM,
not seen).
Conyza lyrata Kunth, Nov. Gen. Sp. (fol.) 4: 55. 1 820.
TYPE: Ecuador, Guayaquil, A. Humboldt & A.
Bonpland 3812 (holotype P, not seen, IDC Mi-
crofiche 6209. 95: II. 5; isotype, B, probably de-
stroyed, F neg. 14934).
Eschenbachia lyrata (Kunth) Britton & Millsp., Fl.
Baham. 444. 1920.
Blumea lyrata (Kunth) Badillo, Bol. Soc. Venez. Ci.
Nat. 10: 257. 1946.
Ernstia lyrata (Kunth) Badillo, Cat. Fl. Venez. 505.
1947, genus ined.
Pseudoconyza lyrata (Kunth) Cuatr., Ciencia (Mexi-
co). 21: 31. 1961.
Pseudoconyza viscosa (Miller) D'Arcy, Phytologia 25:
281. 1973.
Blumea viscosa (Miller) D'Arcy, Phytologia 30: 5. 1 975,
redundant comb.
Herbs to 8 dm tall; stems erect, terete, striate,
sparsely pilose and glandular-puberulent, viscid.
Leaves sessile, simple or divided into stipuleform
auricles; blade obovate to spatulate, l-3(-7) cm
long, 0.5-l(-3.5) cm wide, base attenuate, apex
acute, mucronate, the margins dentate, the teeth
mucronate, both sides sparsely pilose and glan-
dular-puberulent. Capitulescences cymose-panic-
ulate, subtended by a foliaceous bracts; peduncles
1-4 cm long, glandular-puberulent, usually brac-
teolate. Capitula 5-7 mm high, 4-5 mm wide;
involucres narrowly campanulate; phyllaries 3-4-
seriate, the outer lanceolate, 1.5-3 mm long, ca.
0.5 mm wide, pilose and glandular-puberulent,
apex acute, the inner to 6 mm long, ca. 0.7 mm
wide, glandular-puberulent, apex acuminate, stri-
gulose; receptacle flat to depressed-hemispheric,
3-4 mm in diameter, puberulent to glabrescent;
pistillate florets 50-300, the corollas ca. 3.5-4 mm
long, yellowish-white, slightly expanded at the base,
apex 3 -notched, the style 44.5 mm long, the
branches filiform, 0.4-0.5 mm long; hermaphro-
ditic florets 2-12, fertile, the corollas white or
pinkish, tubular, 5-5.5 mm long, the limb slightly
expanded, 5-lobed, the lobes ca. 0.4 mm long,
narrowly triangular, the anthers ca. 1.2 mm long,
caudate, the terminal appendages oblong, apex
rounded, the style 5-5.8 mm long, the branches
cylindrical, 0.2-0.3 mm long, strigillose. Achenes
fusiform, 0.8-1 mm long, ribbed, strigillose, dark
brown; pappus of ca. 10 bristles, ca. 4.5 mm long,
white, persistent, not deciduous together as a unit.
Blumea viscosa is distributed in litoral habitats
throughout the Caribbean region and along the
western coast of South America from Colombia
to northern Peru. It is apparently rare in Peru and
represented by only two recent collections.
This species superficially resembles Conyza (As-
tereae), but is readily distinguished from all Pe-
ruvian members of that genus by its dentate leaves,
tailed anthers, and coastal habitat preference.
20
FIELDIANA: BOTANY
B
FIG. 6. Blumea viscosa. A, flowering branch; B, capitulum; C, outer phyllary; D, inner phyllary; E, pistillate floret;
F, hermaphroditic floret; G, style branches of hermaphroditic floret; H, stamen; I, achene. (Drawn from Sagdstegui
& Mostacero 1 1361, HUT.)
McVaugh (1972) recognized two varieties ofBlu- on Ecuadorian material (Guayaquil, Humboldt &
mea viscosa, a pale-pilose form, based on Mexican Bonpland, P). Badillo ( 1 974) considered this taxon
material (Veracruz, Houston, BM), and a much less to be one highly variable species, and this concept
hairy form with more conspicuous glands, based is followed in the present treatment.
DILLON & SAGASTEGUI: FLORA OF PERU. V.
21
CAJAMARCA. Contumaza: Ascope-San Benito, 500
m, Sagdstegui & Mostacero 11361 (HUT, MO). LAM-
BAYEQUE. Lambayeque: Motupe (Anchoviera), 50 m,
Llatas 1012 (F, PRO).
V. Chevreulia
Chevreulia Cass., Bull. Soc. Philom. Paris 1817:
69. 1817. TYPE: Tussilago sarmentosa Pers.
= Chevreulia sarmentosa (Pers.) Blake.
itary heads at tips of branches, sessile during an-
thesis, the peduncles elongating with maturity of
fruit, to 6(-12) cm long. Capitula 8-10 mm high,
ca. 3 mm wide; involucre cylindrical; phyllaries
ca. 3-seriate, the outer lanceolate 3-5 mm long,
ca. 1 mm wide, dorsally lanuginous, apex acute,
the inner linear-oblong to linear, 7-8 mm long,
0.4-0.6 mm wide, glabrous, apex obtuse; pistillate
florets 10-12, the corollas ca. 3.5 mm long; her-
maphroditic florets ca. 4, the corollas ca. 4 mm
long. Achenes ca. 2 mm long; pappus bristles ca.
3.5 mm long.
Perennial herbs; stems rhizomatous. Leaves ro-
sulate or opposite, the margins entire. Capitulesc-
ences of solitary heads, long-pedunculate at ma-
turity. Capitula disciform; involucres cylindrical
to campanulate; phyllaries 3-5 -seriate, imbricate,
scarious, hyaline, the outer gradually smaller; re-
ceptacles plane, naked; pistillate florets 1-2-seri-
ate, the corollas filiform, white or violet, apex trun-
cate or dentate; disc florets hermaphroditic, the
corollas narrowly tubular, 5-lobed, the anthers
sagittate, caudate, the terminal appendages ovate,
the style branches truncate, dorsally papillose.
Achenes fusiform, contracted into a filiform ros-
trum, asperous; pappus bristles capillary, scabrid,
barbellate, 1-2-seriate, the bases fused. Chromo-
some number: n = 14.
A genus of some six species distributed through-
out South America. Only the following species is
recorded for Peru.
1. Chevreulia acuminate Less., Linnaea 5: 261.
1 830. TYPE: Brazil, Rio de Janeiro, Serra dos
Orgaos, Jan 1828, H. K. Beyrich s.n. (p, ho-
lotype, not seen). Figure 7.
Chevreulia JUiformis Hook. & Arn., Companion Bot.
Mag. 1: 102. 1836. TYPE: Brazil, St. Catherine's,
J. Tweedie s.n. (holotype, GL, not seen).
C. longipes Wedd., Chlor. And. 1: 157. 1856. TYPE:
Peru, Dept. Cuzco, 1839-40, C. Gay s.n. (holo-
type, P, not seen).
Creeping, perennial herbs; stems to 8 cm long,
much-branched, densely foliaceous, procumbent,
laxly lanate. Leaves decussate, sessile; blade ob-
long-lanceolate, 5-15 mm long, 1-4 mm wide, apex
acuminate, mucronate, the lower surface densely
lanate, the upper surface lanuginous to glabres-
cent, the margins revolute. Capitulescences of sol-
Chevreulia acuminata is distributed from Ec-
uador to northern Argentina and Brazil. In Peru,
it is represented by only three collections. It is a
distinctive species with greatly elongating pedun-
cles and rostrate achenes, features unknown in any
other Peruvian Inuleae.
AMAZONAS. Rodriguez de Mendoza: Mendoza, 1 700
m, Woytkowski 8082 (MO). HUANUCO. Huanuco: Mito,
ca. 9000 ft, Macbride& Featherstone 1723 (F,GH); Chas-
qui, ca. 10,500 ft, Macbride & Featherstone 2457 (F).
VI. Cuatrecasasiella
Cuatrecasasiella H. Robinson, Flora Neotropica
39: 14. 1986. TYPE: LuciliopsisiserniiCualT.
= Cuatrecasasiella isernii (Cuatr.) H. Rob-
inson.
Annual or perennial herbs, dioecious; stems ces-
pitose or prostrate. Leaves opposite, decussate,
sessile to subsessile; blade oblong, apex rounded
or subtruncate, the margins entire. Capitulesc-
ences of solitary heads, terminal on branchlets.
Capitula discoid; involucres ovoid or campanulate
to cylindrical; phyllaries 2-3 -seriate, imbricate,
hyaline; receptacles plane, naked; florets in mas-
culine capitula 5-14, the corolla tubular, 5-lobed,
the anther bases sagittate, caudate, the terminal
appendages ovate, the ovary sterile, the style
branches rounded; florets in feminine capitula 8-
15, the corolla filiform, 2 4-lobed. Achenes cylin-
drical, brown, glabrous; pappus bristles scabrid,
barbellate, uniseriate, the bases fused, deciduous
together as a unit, white to reddish brown.
A genus of four species distributed from Ecua-
dor to Argentina in high-elevation habitats. Only
one species is known from Peru.
22
FIELDIANA: BOTANY
Fio. 7. Chevreulia acuminata. A, habit; B, enlargement of stem section; C, capitulum; D, outer phyllary; E, inner
phyllary; F, pistillate floret; G, hermaphroditic floret. (Drawn from Macbride & Featherstone 1723, F.)
Robinson (1986) established Cuatrecasasiella ciliopsis is not dioecious and lacks opposite leaves
for species previously placed in Luciliopsis Wed- throughout, and suggested that Luciliopsis per-
dell (Chloris Andina 1: 159-160. 1856; pi. 26A. pusilla Wedd. is based on a small individual of
1855). He observed that the type species of Lu- Facelis plumosa (Wedd.) Schultz-Bip.
DILLON & SAGASTEGUI: FLORA OF PERU. V.
23
FIG. 8. Cuatrecasasiella isernii. A, habit; B, enlargement of leaves; C, feminine capitulum; D, pistillate floret.
(Drawn from Dillon 2517, HUT.)
References
ROBINSON, H. 1986. Cuatrecasas Festschrift In Hon-
or of the Botanical Career of Jose Cuatrecasas. Flora
Neotropica, 39: 13-16.
1. Cuatrecasasiella isernii (Cuatr.) H. Robinson,
Flora Neotropica, 39: 15. 1986. Figure 8.
Luciliopsis isernii Cuatr., Anal. Univ. Madrid 4: 28.
1935. TYPE: Ecuador, Prov. Napo, paramo Ja-
cupampa, Cerro Antisana, Jan 1865, J. Isern 70
(probably MA, not seen).
Perennial herbs; stems much-branched, 3-5 cm
long. Leaves sessile; blade oval to ovate or oblong,
2-3 mm long, 0.6-2 mm wide, marcescent, apex
obtuse to rounded, often with a tuft of tomentum
persisting, lower surface arachnoid-tomentose to
glabrescent, upper surface glabrous. Capitula 3-5
mm high, 11.5 mm wide; involucres cylindrical;
phyllaries 3-seriate, stramineous, hyaline, the out-
er ovate, ca. 2 mm long, 0.5-0.7 mm wide, arach-
noid-tomentose dorsally, apex obtuse, the inner
oblong to linear, ca. 4 mm long, ca. 1 mm wide,
glabrous, apex acute to obtuse; florets in masculine
capitula ca. 8, the corolla ca. 3 mm long, the an-
thers ca. 1 mm long; florets in feminine capitula
ca. 15, the corolla 3.2-3.5 mm long. Achenes (im-
mature), 0.5 mm long, brown, glabrous; pappus
bristles ca. 4 mm long.
This species is known from high-elevation lo-
calities in Ecuador and Peru, 4100-^4700 m. The
collection from Junin (Dillon 2517) was found
growing within moist hummocks of a Distichia
muscoides moor at the south end of Lago Junin.
No male individuals were found in this popula-
tion. Conversely, the two collections from Ancash
(Bernardi et al. 16591, Smith 11431) appear to
contain only male individuals. Additional field ob-
servations are necessary to ascertain if entire pop-
ulations are actually unisexual or what sex ratios
exist.
ANCASH. Huaraz: Carpa, 4100-^4300 m, Bernardi et
al. 16591 (us). Recuay: Huascaran National Park, mo-
raine below Nevado Pasto Ruri, 4700 m, Smith 11431
(F, MO). JUNIN. Junin: ca. 9 km N of Junin on road to
Cerro de Pasco, 4160 m, Dillon 2517 (F, HUT, MO, us,
USM); Mount La Juntay, near Huancayo, ca. 4700 m,
Killip & Smith 22105 (us).
VII. Facelis
Facelis Cass., Bull. Soc. Philom. Paris 1819: 94.
1819. TYPE: Gnaphalium retusum Lam. =
Facelis retusa (Lam.) Schultz-Bip.
Annual herbs; stems simple or branched, erect
to decumbent. Leaves alternate, the margins en-
24
FIELDIANA: BOTANY
tire. Capitulescences of solitary heads, axillary or
terminal, occasionally glomerulate in axils of the
upper leaves. Capitula disciform; involucres cy-
lindrical to ovoid or campanulate; phyllaries 3-5-
seriate, imbricate, the outer foliaceous, the inner
membranous, hyaline; receptacle plane, naked;
pistillate florets 1-2-seriate, the corollas filiform;
disc florets hermaphroditic, the corollas narrowly
tubular, 5-lobed, the anther bases sagittate, cau-
date, the terminal appendages ovate, the style
branches linear, apex acute, dorsal surfaces pu-
bescent. Achenes turbinate, densely sericeo-velu-
tinous; pappus bristles plumose, uniseriate, the
bases fused, deciduous together as a unit.
Facelis consists of four species distributed
throughout South America. This genus is readily
distinguished by its densely sericeo-pubescent
achenes and pappus of plumose bristles, both char-
acteristics not found elsewhere in the Peruvian
Inuleae. Two species are known from Peru.
References
BEAUVERD, G. 1913. Le Genre Facelis Cassini (emend.
Beauverd). Bull. Soc. Bot. Geneve, ser. 2, 5: 212-220.
CABRERA, A. 1978. Compositae. Flora de la Provincia
de Jujuy. Colecc. Cient., Inst. Natl. Teen. Agropec.,
13: 259-260.
Key to Species of Facelis
1. Herbs with stems generally simple, erect, 5-35 cm high; leaves 15-25 mm long; capitula 5-7 mm
high, 2-3 mm wide F. lasiocarpha
1. Herbs with stems generally much-branched from the base, ascending or decumbent, 2-7 cm high;
leaves 5-10 mm long; capitula 4-5 mm high, ca. 1.5 mm wide F. plumosa
1. Facelis lasiocarpha (Griseb.) Cabr., Physis 10:
280. 1931. Figure 9.
Filago lasiocarpha Griseb., Abh. Konigl. Ges. Wiss.
Gottingen. 19: 180. 1874. TYPE: Argentina, Prov.
Tucuman, Cienaga, Sierra de Tucuman, 13/31
Mar 1872, P. G. Lorentz 123 (holotype, GOET,
not seen; isotype, CORD, not seen).
Facelis schultziana Beauverd, Bull. Soc. Bot. Geneve,
ser. 2, 5: 219. 1913. TYPE: Bolivia, Dept. La Paz,
Prov. Larecaja, "viciniis Sorata, clivosis collis
Catarguata, in dumosis, 2700 m," G. Mandon
177 (holotype, G-BOIS, not seen).
Annual herbs, 5-35 cm tall; stems simple or
sometimes branched from the base, slender, la-
nate, foliaceous to the apex. Leaves sessile; blade
narrowly linear, 1 5-25 cm long, ca. 1 mm wide,
apex attenuate, mucronate, lower surface densely
lanate, upper surface laxly lanate, the margins rev-
olute. Capitulescences glomerulate or of solitary
heads in the axils of the upper leaves, sessile. Ca-
pitula 5-7 mm high, 2-3 mm wide; involucres
ovoid; phyllaries 3-4-seriate, the outer lance-ovate
to lanceolate, 2.54 mm long, 0.5-1.5 mm wide,
foliaceous, lanate, apex acute, mucronate, the in-
ner ovate to oblong, 5-6 mm long, 0.8-1 mm wide,
scarious, apex obtuse to rounded, glabrous; pis-
tillate florets 4-8, the corollas ca. 2.5 mm long;
hermaphroditic florets 2-5, the corollas ca. 3 mm
long. Achenes 1-1.5 mm long; pappus bristles 5-
6 mm long.
Facelis lasiocarpha is distributed throughout the
Andean Cordillera from Ecuador to northern Ar-
gentina, 1800-3800 m. It is very similar to F.
plumosa, differing only in its non-branching habit
and larger capitula and with a greater number of
pistillate florets. Further study may show it to be
conspecific with F. plumosa.
ANCASH. Bolognesi: Cerros al E de Chiquian, 3500-
3600 m, Cerrate 542 (F, MO, USM). CAJAMARCA. Con-
tumaza: Ventarron (Trinidad-Totorillas, 3000 m, Sa-
gdstegui et al. 8988 (F, HUT, MO); Las Campanulas (Guz-
mango), 2700 m, Sagdstegui & Mostacero 9159 (F, HUT,
MO); El Mojon (Las Quinuas-Huatum), 3000 m, Sagds-
tegui et al. 10125 (F, HUT, MO); Yeton-Guzmango, 1800
m, Sagdstegui et al. 10326 (F, HUT, MO), Sagdstegui et al.
10559 (F, HUT). LA LIBERT AD. Otuzco: desvio a Otuz-
co-Agallpampa, 2800 m, Sagdstegui et al. 11534 (F, HUT,
MO). LIMA. Huarochiri: Matucana, ca. 8000 ft, Mac-
bride & Featherstone 446 (F, us).
2. Facelis plumosa (Wedd.) Schultz-Bip., Isinnaea
34: 532. 1866. Figure 9.
Lucilia plumosa Wedd., Chlor. And. 1: 155. 1856.
TYPE: Peru, "Cordilleres du Perou, entre Puno
et Arequipa, 4000 m," H. A. Weddell s.n. (ho-
lotype, P, not seen, F neg. 37819).
Facelis weddelliana Beauverd, Bull. Soc. Bot. Geneve,
ser. 2, 5: 217. 1913. (nomen novum based upon
Lucilia plumosa Wedd.).
DILLON & SAGASTEGUI: FLORA OF PERU. V.
25
FIG. 9. Left side: Facelis lasiocarpha. A, habit; B, capitulum, C, outer phyllary; D, inner phyllary, E, pistillate
floret; F, hermaphroditic floret; G, stamen; H, style branches of hermaphroditic florets. (Drawn from Sagdstegui
3802, HUT.) Right side: Facelis plumosa. A, habit; B, capitulum; C, outer phyllary; D, inner phyllary, E, pistillate
floret; F, hermaphriditic floret; G, stamen; H, style branches of hermaphroditic florets. (Drawn from Sagdstegui 4508,
HUT.)
26
FIELDIANA: BOTANY
Annual herbs, 2-7 cm tall; stems usually
branched from the base, slender, lanate, foliaceous
to the apex. Leaves sessile; blade narrowly linear,
5-10 mm long, ca. 1 mm wide, apex attenuate,
mucronate, lower surface densely lanate, upper
surface laxly lanate, the margins revolute. Capi-
tulescences of 1-3 heads glomerulate in the axils
of the upper leaves, sessile. Capitula 4-5 mm high,
ca. 1.5 mm wide; involucres ovoid-cylindrical;
phyllaries 3-4-seriate, the outer lanceolate to lin-
ear, 2.5-3 mm long, 0.5-1 mm wide, foliaceous,
lanate, apex acute, mucronate, the inner ovate to
oblong, 4-5 mm long, 0.5-1 mm wide, scarious,
apex obtuse to rounded, glabrous; pistillate florets
3-6, the corollas ca. 2.5 mm long; hermaphroditic
florets 2-4, the corollas ca. 2.5 mm long. Achenes
1-1.5 mm long; pappus bristles 34 mm long.
Facelis plumosa is distributed from central Peru
to northern Argentina, 28004300 m. It is similar
to F. lasiocarpha but is distinguished from the
latter by its more branched habit and smaller ca-
pitula with fewer pistillate florets.
AREQUIPA. Arequipa: above Arequipa, 2800 m,
Pennell 13251 (F, GH, us); Pampa de Arrieros, 3750 m,
Pennell 13326 (F). AYACUCHO. Huanta: Pampalca,
between Huanta and Rio Apurimac, ca. 3200 m, Killip
& Smith 23291 (us). LIMA. Lima: Rio Blanco, 12,000
ft, Macbride & Featherstone 738 (F, us). JUNIN. Yauli:
Yauli, ca. 13,500 ft, Macbride & Featherstone 915 (F).
PUNO. Melgar: Araranca, 4 100-4300 m, Pennell 13459
(F, GH, us).
slender; hermaphroditic florets 1-5, the corollas
tubular, yellowish, 5-lobed, the anthers tailed, the
terminal appendages obtuse, the style branches
truncate, penicillate. Achenes obovoid to ellip-
soid, surface with sinuate-reticulate sculpturing,
glandular with bicelled, sessile papillae or trich-
omes, mucilaginous when wet, stramineous to
brown; pappus bristles scabrid, barbellate, uni-
seriate, the bases fused into a ring, lacking basal
cilia, deciduous together as a unit, white. Chro-
mosome number: n = 14.
Gamochaeta is a genus of some 80 species dis-
tributed primarily in the warmer regions of the
New World, but with several species adventive in
the Old World. The taxonomy of Gamochaeta and
its close relatives is extremely difficult because eas-
ily observable or quantifiable characters are lack-
ing. Gamochaeta is here accepted on the strength
of the suite of characteristics that allows for the
recognition of its constituent elements: obovate to
spathulate leaf form, usually spiciform capitules-
cence, involucre cylindrical to conical, number and
ratio of masculine to feminine florets per capitu-
lum, pappus bristles fused at the base, and achenal
surface sculpturing and pubescence.
The discrimination of species within Gamo-
chaeta poses additional problems. Many species
exhibit considerable morphological plasticity, pre-
sumably in response to environmental and edaph-
ic influences. This treatment should be considered
provisional; extensive biosystematic study is be-
yond the scope of the present work.
VIII. Gamochaeta
Gamochaeta Wedd., Chlor. And. 1: 151. 1856.
TYPE: Gnaphalium americanum Miller =
Gamochaeta americana (Miller) Wedd.
Gnaphalium section Gamochaeta (Wedd.) O. Hoffm.
Annual or perennial herbs, usually tomentose
or lanate. Leaves alternate, simple, often with a
basal rosette; blade oblanceolate to spatulate, the
margins entire or crenulate, lanate or tomentose,
occasionally discolorous. Capitulescences glomer-
ulate, spiciform, or paniculate. Capitula disciform,
heterogamous; involucres cylindrical to conical,
rarely campanulate; phyllaries 3-4-seriate, imbri-
cate, scarious, stramineous to dark brown or oc-
casionally reddish, the margins usually hyaline;
receptacles flat, glabrous; pistillate florets 35-100,
the corollas filiform, 5-dentate, the style branches
References
CABRERA, A. L. 1961. Observaciones sobre las Inuleae-
Gnaphalineae (Compositae) de America del Sur. Bol.
Soc. Argent. Hot., 9: 359-386.
DILLON, M. O., AND A. SAGASTEGUI A. 1986. New
species and status changes in Andean Inuleae (Aster-
aceae). Phytologia, 59: 227-233.
DRURY, D. G. 1970. A fresh approach to the classifi-
cation of the genus Gnaphalium with particular ref-
erence to the species present in New Zealand (Inuleae-
Compositae). New Zealand J. Hot., 8: 222-248.
. 1971. The American spicate cudweeds adven-
tive to New Zealand. New Zealand J. Bot., 9: 157-
185.
MILLIARD, O. M., AND B. L. BURTT. 198 la. Some ge-
neric concepts in Compositae Gnaphaliinae. J. Linn.
Soc. Bot., 82: 181-232.
. 1981b. Names in Gnaphalium, Xeranthemum
and Helichrysum published between 1753 and 1800.
J. Linn. Soc. Bot., 82: 233-265.
HOLUB, J. 1976. Gamochaeta. In Tutin, T. G., et al.
(eds.), Flora Europaea, 4: 127.
DILLON & SAGASTEGUI: FLORA OF PERU. V.
27
JANSEN, R. K., T. STUESSY, S. DfAZ-PiEDRAHiTA, AND V.
FUNK. 1984. Recuentos cromosomicos en Compos-
itae de Colombia. Caldasia, 14: 7-20.
MERXMULLER, H., P. LEINS, AND H. ROESSLER. 1977.
Inuleae-Systematic review, pp. 577-602. In Heywood,
V. H., et al. (eds.), The Biology and Chemistry of the
Compositae, Academic Press, London.
NESOM, G. 1990. The taxomonic status of Gamochaeta
(Asteraceae: Inuleae) and the species of the United
States. In press.
TURNER, B. L., A. M. POWELL, AND J. CUATRECASAS.
1967. Chromosome numbers in Compositae. XI. Pe-
ruvian species. Ann. Missouri Bot. Gard., 54: 172-
177.
Key to Species of Gamochaeta
1 . Leaf surfaces discolorous, lower surfaces white or silvery tomentose, upper surfaces glabrescent, gray
or green G. americana
1 . Leaf surfaces similar in color and pubescence, lower and upper surfaces tomentose or lanuginose
2
2. Erect annual or biennial herbs, stems never cespitose, generally greater than 12 cm tall
G. purpurea
2. Cespitose perennial herbs, generally less than 12 cm tall 3
3. Leaves generally obovate, 4-22 mm long, 1.54 mm wide; capitula ca. 4.5 mm high, ca. 4 mm
wide G. humilis
3. Leaves generally oblanceolate, 10-40 mm long, 4-7 mm wide; capitula ca. 4 mm high, ca. 2.5
mm wide G. oreophila
1 . Gamochaeta americana (Miller) Wedd., Chlor.
And. 1: 151. 1856.
Gnaphalium americanum Miller, Gard. Diet.,
ed. 8. 1768. TYPE: Jamaica, 1731, W. Hous-
ton s.n. (holotype, BM, not seen).
Gamochaeta spicata (Lam.) Cabr., Bol. Soc. Ar-
gent. Bot. 9: 380. 1961.
Gnaphalium spicatum Lam., Encycl. 2: 757. 1788.
TYPE: Uruguay, Montevideo, (? "Des envi-
rons de Buenos Ayres") P. Commerson s.n.
(lectotype, p, designated by Cabrera (1961),
not seen, F neg. 37573, fragment, F). Non
Miller (1768).
Annual or biennial herbs to 60 cm tall; stems
branched, decumbent or erect. Basal leaves rosu-
late, sessile; blade linear- to oblong-spatulate, 3-
10 cm long, 8-20 mm wide, marcescent, base trun-
cate and clasping the stem, apex obtuse to round-
ed, rarely acute, apiculate, strongly discolorous,
lower surface tomentose, upper surface glabrescent
or more commonly glabrous and bright green, the
margins entire, sometimes undulate distally; cau-
line leaves linear, apex acute, gradually smaller
upwards, pubescence similar to basal leaves. Ca-
pitulescences spiciform, terminal and axillary. Ca-
pitula 4-4.5 mm high, 2.5-3 mm wide; phyllaries
ca. 24, 3 4-seriate, the outer ovate, 2-3 mm long,
1-1.2 mm wide, apex obtuse to acute, the inner
liner-oblong, 3.5-4 mm long, 0.8-1 mm wide, gla-
brous, apex acuminate, apiculate; pistillate florets
ca. 80, the corollas ca. 2 mm long; hermaphroditic
florets 34, the corollas ca. 2.2 mm long. Achenes
0.5-0.6 mm long; pappus bristles ca. 2.5 mm long.
Chromosome number: n = 14 (Jansen et al., 1984;
Turner et al., 1967, as G. spicata).
Gamochaeta americana is distributed through-
out Central and South America and is adventive
in the United States and New Zealand. In Peru, it
is distributed in a wide range of habits and is quite
variable in overall size. Its common names include
lengua de perro.
There has been considerable disagreement as to
the specific boundaries of Gamochaeta americana.
Adams (Flowering Plants of Jamaica, 1972), and
followed by D'Arcy (Flora of Panama, 1975),
placed G. spicata into synonymy under G. amer-
icana. Cabrera ( 1 96 1 ), Godfrey (1958), and Drury
( 1 97 1 ) all maintained G. americana and G. spicata
and distinguished them by differences in the upper
leaf pubescence and phyllary shape. Both Hillard
and Burtt ( 1 98 1 b) and Nesom ( 1 990) have pointed
out that Lamark's Gnaphalium spicatum is ille-
gitimate as a later homonym of Miller's species
which possibly represents Pterocaulon virgatum
(L.) DC. We have adopted a broad concept for G.
americana and inculde taxa with acute to obtuse
phyllaries and grabrous to glabrescent upper leaf
surfaces.
AMAZONAS. Bagua: 48 km NW of Chachapoyas on
road to Bagua, 1 280 m, Gentry et al. 23210 (F, MO, USM).
Mendoza de Rodriguez: Mendoza, 1 700 m, Woytkowski
8086 (MO). ANCASH. Yungay: Quebrada de Llangan-
uco, 3900 m, Lopez et al. 8347 (HUT, MO); Huascaran
National Park, Llanganuco sector, 3580-3850 m, Smith
28
FIELDIANA: BOTANY
& Goodwin 8838 (F). APURIMAC. Abancay: Sayhuite,
Km 45 E of Abancay, 3600 m, Hutchison 1736 (F, NY
us). CAJAMARCA. Cajamarca: ca. 43 km E of Caja-
marca, 27 km N of San Marcos, 2765 m, Dillon et al.
2861 (CPUN, F, HUT, MO, NY, TEX, US, USM). (VU> 11(1 ill: Pu-
marrume, 2800 m, Mostacero et al. 0999 (F.HUT). CUZ-
CO. Paucartambo: Km 1 4 1 of Kosnipata road, 1 6 km
below Pillahuata, 2050 m, Stein 2525 (F, MO, USM).
HUANCAVELICA. Huancavelica: entre Huancavelica
y Lachoj, 3800 m, Tovar 3080 (USM). HUANUCO.
Huanuco: between Huanuco and Tingo Maria, ca. Km
443.5, 2010 m, Croat 57825 (MO); ca. 47 km NNE of
Huanuco on road to Tingo Maria, Carpish Pass, Dillon
2599 (F); Carpish, entre Huanuco y Tingo Maria, 2700-
2900 m, Ferreyra 2344 (USM). JUNIN. 1 anna: 28-32
km NE of Tarma on road to San Ramon, ca. 2480 m,
Dillon & Turner 1350 (F, MO, USM); ca. 35 km NE of
Tarma along road to San Ramon, ca. 2300 m, Dillon &
Turner 1407 (F, MO, USM); Huacapistana, entre Tarma y
San Ramon, 1800-1900 m, Ferreyra 287 a (USM). Prov.
Unknown: Yaupi, 1 580 m, Woytkowski 6426 (MO), 6428
(MO). LA LIBERTAD. Otuzco: Chota (Yamobamba-
Shorey), 2900 m, Sagdstegui et al. 11081 (F, HUT). San-
chez Carrion: Laguna Sausacocha, 3000 m, Sagdstegui
9444 (F, HUT, MO). LAMBAYEQUE. Lambayeque: Km
28 W of Olmos, 1 1 50-1 200 m, Hutchison & Wright 3474
(F, MO, us). LIMA. Canta: Canta, 2942 m, Lopez 3297
(us). MOQUEGUA. Mariscal Nieto: between Torata
and Carumas, km 75-76 from Moquegua to Puno, 3600
m, Dillon et al. 3357 (F). PASCO. Oxapampa: Oxapam-
pa, Esposto s.n. (USM #19319). PIURA. Huancabamba:
Cuello del Indio (ruta Huancabamba), 2800 m, Lopez
et al. 8889 (F, HUT, MO). PUNO. Carabaya: across Rio
San Gaban from Ollachea, Boeke & Boeke 3053 (us).
SAN MARTIN. Lamas: Alonso de Alvarado, Cerro
Blanco (carretera a Moyobamba), 1000 m, Schunke 6192
(NY, us); San Roque, 1350-1500 m, Williams 7445 (F,
GH p.p, US).
2. GamochaetahumilisWedd.,Chlor. And. 1: 153.
1856. TYPE: Bolivia, Dept. La Paz, Chiquia-
guillo, 4500 m, H. A. Weddell s.n. (holotype,
p, not seen, F neg. 37600).
Perennial herbs to 1 2 cm tall; stems branched,
cespitose to decumbent or erect. Basal leaves ro-
sulate, sessile; blade obovate to oblanceolate or
spatulate, 4-22 mm long, 1.5-4 mm wide, mar-
cescent, base attenuate to a pseudopetiole, apex
obtuse to rounded, rarely acute, slightly discol-
orous, lower surface canescent-tomentose, upper
surface incano-lanate, the margins entire; cauline
leaves oblanceolate to spatulate, often subfalcate,
5-20 mm long, 2-5 mm wide, base attenuate, apex
acute, pubescence similar to basal leaves. Capitu-
lescences laxly spiciform or terminally glomer-
ulate. Capitula ca. 4.5 mm high, ca. 4 mm wide;
involucres campanulate; phyllaries ca. 24, 4-se-
riate, stramineous, the outer ovate, 1.5-2.2 mm
long, 0.5-1 mm wide, dorsally lanate, apex acu-
minate, the inner oblong-linear, 2.8-3 mm long,
0.4-0.8 mm wide, lanate to glabrous, apex acute;
pistillate florets 3540, the corollas ca. 2 mm long;
hermaphroditic florets ca. 3, the corollas ca. 2 mm
long. Achenes 0.5-0.7 mm long, sparsely glandular
to glabrous; pappus bristles ca. 2 mm long.
Gamochaeta humilis is distributed from the De-
partment of Ancash in central Peru to northern
Bolivia. In Peru, it occurs in open areas of puna
vegetation, 3560-4600 m.
This species is characterized by its short, ces-
pitose habit (usually less than 1 2 cm tall), mostly
falcate leaves, and more glomerulate capitules-
cences. In general habit, Gamochaeta humilis ap-
proaches the form of various Belloa species, but
G. humilis may be distinguished by the truncate
style branches of the hermaphroditic florets. Gam-
ochaeta humilis appears most closely related to G.
oreophila, which occurs in central and northern
Peru, and it appears that the distributional ranges
of the two overlap to some degree in Ancash and
La Libertad. Further sampling is necessary to de-
termined if these two species occur sympatrically
or occupy different microhabitats.
ANCASH. Huaraz: Huascaran National Park, Que-
brada Llaca, 4400-4600 m, Smith et al. 10807 (F). AYA-
CUCHO. Huanta: between Huanta and Hacienda Par-
gora, 4150 m, Killip & Smith 22192 (F, NY). CUZCO.
Quispicanchis: Auzangate, 4600 m, Rauh & Hirsch PI 244
(NY). Paucartambo: Tres Cruces, 1-13 km NW of Pau-
cartambo-Pilcopata road, 3330-3500 m, Gentry et al.
23470 (F p.p., MO, USM); paso de Tres Cruces, Cerro de
Cusilluyoc, 3800-3900 m, Pennell 13869 (F p.p., NY p.p.).
Paruro: Cerro Sullccan, Hda. Ayusbamba, 4020 m, Var-
gas 845 (F). Urubamba: Pampaccahuana, 4100 m, Pey-
ton & Peyton 77 (MO). JUNIN. Tarma: Oroya-Tarma,
4200 m, Rauh & Hirsch P269 (NY). Yauli: Yauli, ca.
13,500 ft, Macbride & Featherstone 930 (F). LA LIB-
ERTAD. Santiago de Chuco: Jalca de Coipin, 4000 m,
Sagdstegui et al. 11982 (F, HUT). MOQUEGUA. Mar-
iscal Nieto: between Torata and Carumas, Km 75-76
from Moquegua to Puno, ca. 3600 m, Dillon et al. 3356
(F). PUNO. Azangaro: Arapa, 3900 m, Aguilars.n. (USM
P.P.). Carabaya: Macusani-Condorquina, 4400 m, Var-
gas 17630 (us). Huancane: Majachi, 3950 m, Aguilar
s.n. (F.USM); Moho, 3950 m, Aguilar 4 14 (USM). Melgar:
La Raya-Sicuani, 4300 m, Rauh & Hirsch P678 (NY).
Prov. Unknown: Lago Titicaca, 4000 m, Monheim Ml 02
(NY).
3. Gamochaeta oreophila Dillon & Sagast., no-
men novum. Figure 10.
Gamochaeta monticola Dillon & Sagast., Phytologia
59: 227. 1986. TYPE: Peru, Dept. La Libertad,
Prov. Santiago de Chuco, entre Chota y Shorey,
DILLON & SAGASTEGUI: FLORA OF PERU. V.
29
H
FIG. 10. Gamochaeta oreophila. A, habit; B, leaf (underside); C, capitulum, D, outer phyllary; E, inner phyllary;
F, pistillate floret; G, hermpahroditic floret; H, style branches of hermaphroditic floret. (Drawn from Sagdstegui et
al. 11 100, HUT.)
ladera, 3250 m, 15 Nov 1983, A. Sagdstegui A.
et al. 11100 (holotype, HUT; isotypes, F, MO). A
new specific epithet is required because of the
discovery of the earlier homonym Gamochaeta
monticola (Phil.) Cabrera, Bol. Soc. Argent. Bot.
9: 374. 1961.
Cespitose, perennial herbs; stems simple or
30
FIELDIANA: BOTANY
branched, 1-6 cm tall. Basal leaves densely ro-
sulate, sessile; blade oblanceolate, 1-4 cm long, 4
7 mm wide, base attenuate to a winged petiole,
apex acute to rounded, mucronate, discolorous,
lower surface tomentose, upper surface lanate, the
margins entire, the cauline leaves gradually small-
er. Capitulescences spiciform. Capitula 3.54 mm
high, 2-2.5 mm wide; involucres campanulate;
phyllaries ca. 22, 3 4-seriate, stramineous, the
outer ovate, ca. 2.2 mm long, ca. 1.2 mm wide,
dorsally lanate, apex acuminate, the inner oblong-
linear, 2.9-3.2 mm long, 0.4-0.7 mm wide, gla-
brous, apex acute; pistillate florets 42-45, the co-
rollas ca. 2 mm long; hermaphroditic florets 2-3,
the corollas ca. 2 mm long. Achenes 0.5-0.7 mm
long; pappus bristles ca. 2.5 mm long.
Gamochaeta oreophila is infrequent in open ar-
eas within ya/ca formations of northern and central
Peru (Departments of Ancash, Cajamarca, and La
Libertad, 31004870 m). It is a distinctive mem-
ber of the genus, possessing densely rosulate, ca-
nescent, basal leaves and erect, compressed spi-
ciform capitulescences. It most closely resembles
G. humilis of central and southern Peru and north-
ern Bolivia; however, the latter species normally
has a more branched, decumbent habit, smaller
spatulate and falcate leaves, and an open capitu-
lescence.
ANCASH. Carhuaz: Huascaran National Park, Que-
brada Ishinca, 4380-4500 m, Smith et al. 9486 (F, MO,
USM). Huaraz: Huascaran National Park, Quebrada
Shallap, 3690-4100 m, Smith et al. 9650 (F,MO). Huay-
las: Huascaran National Park, Quebrada Alpamayo,
4350-4500 m, Smith et al. 9801 (USM), 9849 (F). Yungay:
Huascaran National Park, Quebrada Ancosh, Smith &
Goodwin 8886 (MO, USM). CAJAMARCA. Celendin: ca.
57 km NE of Cajamarca along road to Celendin, ca. 3650
m, Dillon & Turner 1614 (F); Sendamal, 3100 m, Sa-
gdstegui et al. 12228 (F, HUT, K, MO). San Miguel: Taulis
Alto Galca), ladera, 3100 m, Sagdstegui et al. 9548 (F,
HUT, MO). LA LIBERTAD. Otuzco: Motil-Shorey, 3500
m, Sagdstegui & Fernandez 4914 (HUT, us), Sagdstegui
et al. 11698 (F, HUT, MO); Cerro Ragache, Salpo, 3500
m, Sagdstegui et al. 11627 (HUT, MO). LAMBAYEQUE.
Ferreflafe: ca. 7 km NW of I ncahuasi , near Cerro Pun-
amachay on trail to Laguna Hualtaco, 3300-3550 m,
Dillon & Skillman 4128 (F), Sagdstegui et al. 12842 (F,
HUT, MO), Sagdstegui et al. 12865 (F, HUT), Sagdstegui et
al. 12914 (F, HUT, MO, K).
4. Gamochaeta purpurea (L.) Cabr., Bol. Soc. Ar-
gent. Bot. 9: 377. 1961.
Gnaphaliumpurpureum L., Sp. PI. 2: 854. 1 753. TYPE:
United States, "habitat in Carolina, Virginia,
Pensylvania [Pennsylvania]," P. Kalm s.n. (ho-
lotype, LINN, not seen, IDC Microfiche 117. 572.
Ill: 2.).
Annual or biennial herbs to 40 cm tall; stems
branched, decumbent or erect. Basal leaves rosu-
late, sessile; blade spatulate to oblanceolate, 1.5-
3 cm long, 5- 1 5 mm wide, marcescent, base trun-
cate and clasping the stem, apex obtuse to round-
ed, rarely acute, apiculate, discolorous, lower sur-
face tomentose, upper surface glabrescent or more
commonly glabrous and bright green, the margins
entire, sometimes undulate distally; cauline leaves
linear, apex acute, gradually smaller upwards, pu-
bescence similar to basal leaves. Capitulescences
spiciform, terminal. Capitula ca. 3.5 mm high, ca.
2.5 mm wide; phyllaries ca. 24, 3 4-seriate, stra-
mineous, the outer ovate, 2-2.5 mm long, 1-1.5
mm wide, apex obtuse to acute, the inner linear-
oblong, 3-3.5 mm long, 0.5-0.8 mm wide, gla-
brous, apex acute; pistillate florets 70-80, the co-
rollas ca. 2.2 mm long; hermaphroditic florets 3-
4, the corollas ca. 2 mm long. Achenes ca. 0.5 mm
long; pappus bristles ca. 2 mm long. Chromosome
number: n = 14.
Gamochaeta purpurea has a wide distribution,
including North and South America and Europe,
and is adventive in New Zealand. In Peru, it is a
common, weedy species and occupies a wide range
of habitats, from coastal lomas formations to high-
elevation Andean sites, 54300 m.
Three collections from southern Peru (Vargas
844, 845, & 9812) possess more narrowly linear
leaves with denser, lanate-tomentose pubescence
and congested, glomerulate capitulescences. While
these collections are quite distinct, we are hesitant
to give them formal taxonomic status until this
genus is better understood.
Common names include qheto qheto, lengua de
perro, allucupa callun, and lling-llicha del cerro.
ANCASH. Bolognesi: cerca a Chiquian, 3240-3300
m, Cerrate 785 (MO, USM); Ferreyra 7586 (MO, USM). Hu-
ari: Huascaran National Park, Quebrada Pucaraju, 3900-
4200 m, Smith et al. 12705 (USM); 14.8 km NE of Tunel
Cahuish, 3000-4000 m, Stevens 21964 (MO). Huaylas:
carretera a Laguna Paron, ca. 24 km de Caraz, 3100 m,
Ldpez 1865 (us); Huascaran National Park, Quebrada
Santa Cruz, 4300 m, Smith et al. 9321 (F, MO, USM).
APURIMAC. Abancay: between Saywite and Chachora,
4000 m, Vargas 9812 (F). AREQUIPA. Arequipa: south-
ern slopes of Chachani, N of Arequipa, Hinkley 6 (F);
Nevado de Chachani, 4 100-4200 m, Pennell 13290 (GH,
NY); Tiabaya, 2100-2200 m, Pennell 13094 (F, NY). Ca-
mana: Lomas de Camana, ca. 32 km SE of Camana, ca.
1000 m, Dillon & Dillon 3878 (F, HUT, K, MO, NY, USM);
DILLON & SAGASTEGUI: FLORA OF PERU. V.
31
Lomas de Camana, 600 m, Ferreyra 11688 (USM). Islay:
Lomas de Mejia, ca. 8 km N of Mejia, 500-600 m, Dillon
& Dillon 3725 (F, HUT, MO, NY, USM); Lomas de Mollendo,
ca. 8 km NW of Islay, ca. 510 m, Dillon & Dillon 3941
(F, HUT, MO, NY, USM); bajando la Jolla a Matarani, 500
m, Ferreyra 6381 (USM); Mollendo, Hitchcock 22411
(us); 7 km NE de Matarani, 450 m, Lopez 046 (MO).
CAJAMARCA. Cajamarca: ca. 24 km SW of Cajamarca
on road to Cumbemayo, ca. 3600 m, Dillon et al. 2918
(F, USM) Namora, 2600 m, Sagdstegui 7738 (HUT, MO);
Santa Apolonia-Cumbe Mayo, 3000 m, Sagdstegui et al.
11276 (F, HUT). Celendin: Pumarrume, 28850 m, Mos-
tacero et al. 1002 (F, HUT). Chota: Laguna Yahuarcocha,
arriba de Incahuasi, 3600 m, Sagdstegui 12897 (F, HUT).
Contumaza: Yeton, 2200 m, Sagdstegui et al. 9742 (F,
HUT, MO); Sagdstegui & Lopez 10551 (F, HUT, MO); El
Granero, 2800 m, Sagdstegui & Lopez 10651 (F, HUT),
70652 (F, HUT). CUZCO. Cuzco: Kaira, 3200 m, Vargas
659 (F). Paucartambo: Tres Cruces, 3600 m, Marin 1245
(F); Acanacu, 3500 m, West 7040 (GH). Paruro: cerro
Sullccan, Hda. Ayusbamba, 4020 m, Vargas 844 (F), 845
(MO). Urubamba: Ollantaytambo, 3000 m, Cook & Gil-
bert 707 (us); summit of Antakillqua, 4500 m, Davis et
al. 1712 (F); Vilcanota Valley, 3000 m, Herrera 1068
(us). HUANCAVELICA. Huancavelica: Caniorccona, a
3 km SE de Conaica, 3560-3590 m, Tovar 96 (us).
HUANUCO. Huanuco: Mito, 9000 ft, Macbride &
Featherstone 1750 (F, us), 1814 (F, us); Huanuco, Mac-
bride & Featherstone 2446 (F, us). JUNIN. Tarma: Aco-
bamba, hern 2029 (F). Yauli: Oroya, Kalenborn & Ka-
lenborn 65 (NY); near Atocsayco, ca. 4200 m, Pearsall
1149 (F). LA LIBERTAD. Otuzco: Plazapampa, 1800
m, Cevascos.n. (MO); Chota (Yamobamba-Shorey), 2900
m, Sagdstegui et al. 11081 (F, HUT); Cerro Ragache (Sal-
po), 3200 m, Sagdstegui et al. 11572 (F, HUT, MO). Pataz:
Huancaspata-Tayabamba, 3800 m, Lopez & Sagdstegui
8260 (HUT, MO). Santiago de Chuco: Santiago de Chuco,
2800 m, Sagdstegui et al. 11725 (F, HUT, MO). Trujillo:
between Trujillo and Salaverry, 5 m, Eyerdam 9052 (GH);
Cerro Cabezon, 600 m, Sagdstegui & Lopez 11328 (F,
HUT); Cerro Campana, 500 m, Sagdstegui 4041 (HUT,
us), Sagdstegui & Lopez 10416 (F, HUT, K, MO); Barraza,
60 m, Sagdstegui 7802 (F, HUT, MO, NY). LAMBA-
YEQUE. Chiclayo: Cerro Reque, 540 m, Llatas 358 (F,
HUT). Ferreflafe: Laguna Tembladera, Distrito Incahu-
asi, 3150 m, Sagdstegui et al. 12800 (F, HUT), 72522 (F,
HUT). Lambayeque: Abra de Porculla, km 45 E of Olmos,
1920 m, Plowman et al. 14267 (F, HUT). LIMA. Canete:
Lomas de Quilmana, entre Mala y Canete, 300-480 m,
Ferreyra 4006 (us, USM); Lomas de Grandos, 1 2 km N
of Huaral, 600 m, Stork & Vargas 9330 (GH); hills of
Asia, 300 m, Vargas 9302 (GH). Canta: ca. 1.5 km NE
of Canta, ca. 3000 m, Dillon et al. 3195 (F). Chancay:
Lomas de Lachay, entre Chancay y Huacho, 560 m,
Cerrate 3826 (USM); Lomas de Lachay, ca. 105 km N of
Lima, 300-500 m, Dillon et al. 3618 (F, HUT, MO, USM);
Lomas de Granados, cerca a Huaral, 250-300 m, Fer-
reyra 19417 (us, USM). Huarochiri: Matucana, 8000 ft,
Macbride & Featherstone 184 (F,US); Rio Blanco, 12,000
ft, Macbride & Featherstone 739 (F, us). Lima: Lomas
de Amancaes, 400 m, Cerrate 2081 (MO, USM); Lomas
de Pachacamac, 360 m, Cerrate 3552 (F, USM); Lomas
de Amancaes, 300-400 m, Ferreyra 4042 (us, USM); Lo-
mas de Lurin, 400-500 m, Ferreyra 9546 (MO, USM).
MOQUEGUA. Mariscal Nieto: Lomas de Ilo, ca. 20 km
E of Ilo, ca. 620 m, Dillon et al. 3326 (F, HUT, K, MO, NY,
us, USM), Dillon & Dillon 3649 (F, MO, USM), Ferreyra
12568 (MO, USM). PIURA. Ayabaca: Ayabaca, 2742 m,
Soukup 4341 (us). Huancabamba: 40-43 km E of Olmos
on road to Pucara, 1600-1850 m, Gentry et al. 22661
(F, HUT, MO, USM). PUNO. Azangaro: Tequena, 20 km N
de Arapa, Aguilar 418 (USM p.p.). Melgar: Nanoa, 3900-
4000 m, Vargas 16254 (us). Puno: Amantani, 3900 m,
Aguilar 439 (MO, USM); Puno, 3900 m, Aguilar s.n. (
