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. (MO,
USM); Lampa, 4000 m, Rauh & Hirsch P650 (NY). TAC-
NA. Tacna: Lomas cerca al Morro Sama, 500-600 m,
Ferreyra 12551 (MO, USM).
IX. Gnaphalium
Gnaphalium L., Sp. PI. 850. 1753. LECTOTYPE:
G. uliginosum L., designated by Hitchcock &
Green (1 929). '
Annual or perennial herbs, rarely suffrutescent,
usually lanate or tomentose. Leaves alternate, sim-
ple, sessile or petiolate, at times decurrent upon
the stem; blade lanate or tomentose, occasionally
glandular-puberulent, the margins entire or cren-
ulate. Capitulescences corymbose to paniculate or
cymose, open or dense, terminal and axillary, rare-
ly of solitary heads, bracteolate. Capitula disci-
form, heterogamous; involucres ovoid to campan-
ulate; phyllaries 3 4-seriate, imbricate, scarious,
white, stramineous, brown, or sometimes reddish,
the margins usually hyaline; receptacles flat, gla-
brous or sometimes with rudimentary caducous
paleae; pistillate florets, (25-)40-130, the corolla
filiform, the base slightly expanded, apex contract-
ed, minutely 3 4-lobed, the style branches slender;
hermaphroditic florets functionally staminate, 5-
10(-25), the corolla tubular, yellow, white, or pur-
plish, 3-5-lobed, the lobes usually pubescent with
multicellular, stalked, capitate-glandular trich-
omes, the anthers sagittate, caudate, the terminal
appendages obtuse, the style branches truncate,
penicillate, the nectary conspicuous. Achenes ob-
long, subterete, glabrous, or papillose, rarely
sparsely pubescent with multicellular, biseriate
trichomes; pappus of smooth to scabrid, barbellate
bristles, uniseriate, free, usually caducous, white.
Chromosome numbers: n = 7, 14, 21, 28.
Gnaphalium contains over 1 50 species and has
a cosmopolitan distribution, with centers of di-
versity in Africa, Mexico, and upland South
1 For a discussion of the choice of lectotype, see Jeffrey
(1979).
32
FIELDIANA: BOTANY
America. The genus is represented by seven spe-
cies in Peru occurring in a wide variety of habitats.
The generic limits between Gnaphaliwn and its
close relatives, Achyrocline and Helichrysum, are
ill-defined and often arbitrary. Gnaphalium is here
considered to include those species with campan-
ulate involucres, truncate style branches (her-
maphroditic florets), free pappus bristles, and a
large ratio of pistillate (25-120) to hermaphroditic
(5-25) florets).
Milliard and Burtt (1981) have provided evi-
dence to support the acceptance of the segregate
genus Pseudognaphalium Kirpichnikov. This ge-
nus was based upon the Mexican species, Gna-
phalium oxyphyllum DC, and they expanded it
to include 40 to 50 taxa from Africa, Asia, and
several from South America. Furthermore, they
maintain that Pseudognaphalium has little affinity
with Gnaphalium (sensu stricto) (Milliard & Burtt,
1981, p. 202) and suggest greatest affinity with
Helichrysum. Only one South American species,
Gnaphalium cheiranthifolium Lam., has been
transferred, and numerous other American species
need to be reclassified and transferred to Pseu-
dognaphalium. While we recognize this genus, it
is beyond the scope of this study to provide the
transfers necessary to accept the genus in practice.
Species boundaries in Gnaphalium are often ill
defined. McVaugh (Rora Novo-Galiciana 1 2: 447.
1 984) has discussed the problem of specific bound-
aries in Gnaphalium and his remarks are appli-
cable to the genus in Peru:
Most of the species, even the ones that are pretty
surely native plants with well-defined natural ranges,
behave as weeds when their surroundings are dis-
turbed. Under these circumstances they exhibit a great
deal of vegetative variability, making identification
difficult without prolonged study. The treatment of-
fered below is a provisional one, as the diagnostic
characters employed are quite empirical, and the ob-
served variation (e.g., in number of phyllaries, relative
numbers of perfect and pistillate flowers, etc.) should
be interpreted anew in the light of field study and
experimentation.
With these words of caution, the treatment pre-
sented here should be regarded as provisional, for
many taxonomic and nomenclature problems re-
main to be solved within South American Gna-
phalium.
References
CABRERA, A. L. 1961. Observaciones sobre las Inuleae-
Gnaphalineae (Compositae) de America del sur. Bol.
Soc. Argent. Bot., 9: 359-386.
. 1978. Compositae. Flora de la Provincia de
Jujuy. Colecc. Cient., Inst. Natl. Teen. Agropec., 13:
275-288.
MILLIARD, O. M., AND B. L. BURTT. 1981. Some generic
concepts in Compositae Gnaphaliinae. J. Linn. Soc.
Bot., 82: 181-232.
JANSEN, R. K., T. STUESSY, S. DIAZ-PIEDRAHITA, AND V.
FUNK. 1984. Recuentos cromosomicos en Compos-
itae de Colombia. Caldasia, 14: 7-20.
JEFFREY, C. 1979. Note on the lectotypification of the
names Cacalia L., Matricaria L. and Gnaphalium L.
Taxon, 28: 349-351.
KOCKX-VAN ROON, M., AND J. H. WlEFFERING. 1982.
IOPB Chromosome Number Reports LXXV. Taxon,
31: 367.
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.
Key to Species of Gnaphalium
1 . Plants prostrate, decumbent, or suberect, often cushion-shaped; stems usually less than 1 2 cm long
2
2. Phyllaries milk-white, leaves subtending capitulescences linear-spatulate, apex rounded to obtuse
G. lacteum
2. Phyllaries yellowish or brown, never milk-white; leaves subtending capitulescences spatulate to
linear-lanceolate, apex acute to obtuse or rounded 3
3. Perennials; leaves densely and compactly white-tomentose, linear to narrowly spatulate, the
margins often revolute and the leaves appearing cylindrical or thickened G. polium
3. Annuals; leaves laxly arachnoid-lanate and gray-tomentose, lanceolate to spatulate, the margins
not revolute, the leaves appearing plane G. badium
1 . Plants erect or rarely decumbent; stems usually 1 2 cm long or longer 4
4. Stoloniferous, single-stemmed, rosulate herbs; phyllaries with white, petaloid lamina at apex . .
. . G. antennarioides
DILLON & SAGASTEGUI: FLORA OF PERU. V.
33
4. Multistemmed herbs, rarely with persistent, rosulate basal leaves; phyllaries lacking petaloid
lamina at apex 5
5. Plants annual; capitula 2.5-3 mm high; achenes minutely pubescent with multicellular, biseriate
trichomes G. luteo-album
5. Plants perennial; capitula 4-7 mm high; achenes imbricate-papillose, but lacking multicellular
trichomes 6
6. Plants robust often to 1 m tall or greater; cauline leaves lanceolate to elliptic, 3.5-7 cm
long, 5-20 mm wide, strongly discolorous; capitula 5-7 mm high; inner phyllaries usually
acute or apiculate at apex; achenes ca. 0.6 mm long, 5-ribbed G. elegans
6. Plants usually 0.5 m tall or shorter; cauline leaves linear to lanceolate, (l-)2-5(-7) cm long,
2-5 mm wide, not strongly discolorous; capitula ca. 4 mm high; inner phyllaries usually
obtuse at apex; achenes ca. 0.5 mm long, not obviously ribbed G. dombeyanum
1. Gnaphalium antennarioides DC., Prodr. 6: 224.
1838.
HelichrysumgnaphaloidesKunlh, Nov. Gen. PI. (fol.)
4: 68. 1820. TYPE: Peru, Dept. Piura, Prov. Ay-
abaca, "crescit prope pagum Ayavacae Peruvi-
anorum," A. Humboldt & A. Bonpland s.n. (ho-
lotype, P, not seen, IDC Microfiche 6209. 97: I.
7, F neg. 37977). Non H. gnaphaloides Steud.
Antennaria monica Wedd., Chlor. And. 1: 150. 1856.
TYPE: Based upon Kunth's Helichrysum gna-
phaloides. Weddell also cited the following col-
lections: Colombia, Dept. Tolima, Quebrada del
Azufral, J. Goudot (P, not seen); Ecuador, Prov.
Pichincha, Quito, A. Humboldt & A. Bonpland
s.n. (P, not seen, F neg. 37700).
Rosulate herbs, (9-) 12-30 cm tall, stolonifer-
ous; stems erect, foliaceous, lanate. Basal leaves
sessile; blade oblanceolate to spatulate, 2-5 cm
long, 3-5 mm wide, densely lanate-tomentose;
cauline leaves linear to narrowly spatulate, 14 cm
long, 1.5-2.5 mm wide, sessile, discolorous, often
falcate, densely lanate-tomentose. Capitulescences
densely glomerulate, terminal, peduncles to 2 mm
long, subtended by densely lanate-tomentose fo-
liaceous bracts. Capitula 6-7 mm high, 3-4 mm
wide; involucres narrowly campanulate; phyllaries
ca. 3-seriate, the outer ovate to lanceolate, 3-3.5
mm long, 1-1.5 mm wide, the inner linear-lan-
ceolate, 5.5-6 mm long, ca. 1 mm wide, all with
apices prolonged into a petaloid lamina, white,
rounded to acute; pistillate florets 50-75, the co-
rollas 2.6-3 mm long, apex pink; hermaphroditic
florets 7-15, the corollas 2.5-3.3 mm long, pink.
Achenes 0.6-0.8 mm long; pappus bristles 3.5-4
mm long.
Gnaphalium antennarioides is a relatively un-
common species distributed from Colombia to
Bolivia. In Peru, it is known from scattered col-
lections in high-elevation sites from the Depart-
ments of Piura to Cuzco. It is characterized by its
erect, white petaloid phyllaries, stoloniferous hab-
it, and terminal congested capitulescences. It has
been confused with Antennaria linearifolia, an-
other species with white petaloid phyllaries, but
G. antennarioides may be readily distinguished by
its strictly heterogamous capitula.
Gnaphalium antennarioides appears most close-
ly related to G. paramorum S. F. Blake, a species
with a rosulate habit and white, petaloid phyllaries
from Colombia and Venezuela.
AMAZON AS. Bagua: Cordillera Colin NE of La Peca,
10,800 ft, Barbour 3428 (F, MO). CUZCO. Paucartambo:
Tres Cruces, upper edge of Parque Nacional de Manu,
1-13 km NW of Parcartambo-Pilcopata Road, 3330-
3500 m, Gentry et al. 23439 (F, MO); Cerro de Cusilluyoc,
3800-3900 m, Pennell 13869 (F p.p., GH, NY p.p.); Cori-
huayrachina, Callanga, 3800 m, Woytkowski 565 (MO,
USM). Urubamba: Machu Picchu, on the Palcay side of
the Salcantay-Palcay pass, 3500 m, Peyton & Peyton 711
(F, MO).
2. Gnaphalium badium Wedd., Chlor. And. 1 : 145.
1856. TYPE: Bolivia, Dept. Carangas, A. C.
V. d'Orbigny 1366 (lectotype, P, designated by
Cabrera (1978), not seen, F neg. 37581; lec-
totype fragment, F).
Prostrate to decumbent, annual herbs; stems 5-
1 5 cm long, much-branched. Leaves sessile; blade
oblong-spatulate, 0.5-1.5 cm long, 3-5 mm wide,
base attenuate, apex obtuse to rounded, both sur-
faces laxly arachnoid-lanate. Capitulescences cor-
ymbose or glomerulate with 2-many heads. Ca-
pitula 4-5 mm high, 4-5 mm wide, subtended by
leaves; phyllaries ca. 3-5 -seriate, the outer ovate,
the inner oblong, apex brown; pistillate florets ca.
100, the corollas 2.5-2.9 mm long; hermaphro-
ditic florets 12-25, the corollas 2.5-2.9 mm long.
34
FIELDIANA: BOTANY
Achenes ca. 0.5 mm long; pappus bristles ca. 3
mm long.
Gnaphalium badium is distributed from north-
ern Argentina to central Peru (3200-4800 m). It
is distinguished by its small, annual habit and spat-
ulate leaves with lax, gray, arachnoid-lanate to-
mentum. Its capitulescences are often surrounded
and obscured by the upper leaves. The majority
of collections have been from moist areas near
high-elevation lakes.
This species may be confused with G. polium,
another high-elevation species with a reduced hab-
it, but is distinguished from the latter by its annual
habit and more arachnoid-tomentose leaves that
engulf and obscure the capitulescences.
ANCASH. Carhuaz: Huascaran National Park, Que-
brada Ishinca, 4250-4100 m, Smith et al. 9555 (F, MO).
AYACUCHO. No Exact Locality: Valley Puquio, 4000
m, Rauh & Hirsch P452 (NY). JUNIN. Yauli: near Atoc-
sayco, ca. 4200 m, Pearsall 897 (F); Marcapomacocha,
4300 m, Velarde 6593a (us). LIMA. Huarochiri: Rio
Blanco, Macbride & Featherstone 782, 3026 (F, us); Ti-
clio, 4700 m, Rauh & Hirsch P255 (NY). PUNO. Huan-
cane: Cerro Calvario, Conima, 3900 m, Aguilars.n. (USM),
443 (MO). Melgar: Orurillo, Soukup 554 (F). Puno:
Amantani, 3900 m, Aguilar s.n. (USM).
3. Gnaphalium dombeyanum DC., Prodr. 6: 225.
1838. TYPE: Peru, Dombey 38 (holotype, G-
DC, not seen, IDC Microfiche 800. 1066: II.
4, F neg. 8223; isotype, F).
Erect or ascending herbs to 50 cm tall; stems
densely lanate. Lower cauline leaves sessile to sub-
amplexicaul, not decurrent on stems; blade linear-
oblong to spatulate, (l-)2-5(-7) cm long, 2-5 mm
wide, apex obtuse, lower surfaces white-tomen-
tose, upper surfaces glandular-puberulent, the
margins entire, often undulate; upper cauline leaves
gradually smaller, base auriculate, amplexicaul,
apex long acuminate. Capitulescences corymbose
in semiglobose clusters of 4-12 heads. Capitula ca.
4 mm high, ca. 4 mm wide; involucres ovoid;
phyllaries pale yellow to brown, glabrous, the out-
er broadly ovate, 34 mm long, ca. 1.5 mm wide,
apex acute, the inner oblong, 3.6-4 mm long, 1.5-
1.6 mm wide, apex obtuse, rarely mucronate; re-
ceptacles ca. 2 mm wide; pistillate florets 40-120
(-130), the corollas 2-3 mm long, 3-lobed, often
reddish, glandular; hermaphroditic florets (4-)7-
15(-20), the corollas 2-3 mm long, 5-lobed.
Achenes ca. 0.5 mm long; pappus bristles 2-2.5
mm long. Chromosome number: n = 1 (as G.
chieranthifoliwri).
Gnaphalium dombeyanum is a ubiquitous
member of the genus in Peru and found in a wide
range of habitats at various elevations. The con-
cept adopted here recognizes a "species complex"
that includes several previously recognized spe-
cies; however, the pattern of morphological vari-
ability exhibited by these Andean elements makes
application of names difficult. Populations are of-
ten morphologically heterogeneous with individ-
uals having quite different character combinations
found within a particular geographic area or en-
vironment. Characters that tend to exhibit the
greatest variability include overall size of individ-
uals, size and shape of the leaves, the density of
pubescence, degree of expansion of the capitu-
lescences, and color of phyllaries. Until a detailed
biosystematic study of the genus is undertaken,
this broad species delineation must be applied.
Label data indicate that Gnaphalium dombey-
anum is occasionally used in folk medicine. It has
the following common names: hot on de oro, huira-
huira, qheto qheto, vira-vira, wila wila.
Gnaphalium dombeyanum Complex
Several species originally described from Pe-
ruvian material are here considered to belong to
the Gnaphalium dombeyanum complex. Because
these taxa, upon further study, may prove worthy
of taxonomic recognition, they are not here placed
into formal synonymy. Rather, their distribution
and/or salient characters are discussed; their final
disposition must await further study.
Gnaphalium peruvianum Sprengel, Syst. veg. 3:
473. 1826 and G. dysodes Sprengel, Syst. veg.
4: 776. 1827
These two species represent the oldest names
for Peruvian material here placed under G. dom-
beyanum. Types of these species have not been
located. Gnaphalium dysodes has occasionally been
applied to Peruvian collections; no material an-
notated as G. peruviana was seen in this study.
Gnaphalium jelskii Hieron., Bot. Jahr. 36: 483.
1905. SYNTYPES: Peru, Dept. Cajamarca,
DILLON & SAGASTEGUI: FLORA OF PERU. V.
35
Cutervo, K. Jelski 707, 710, 711 (B, probably
destroyed, not seen, syntype 707 = F neg.
15104).
This species appears to fall into the range of
morphological variability of G. dombeyanum;
however, no type material has been studied.
Gnaphalium helichrysoides Wedd., Chlor. And. 1 :
146. 1856. SYNTYPES: Peru, Dept. Cuzco,
Oct 1839-Feb 1840, C. Gays.n. (P, not seen,
F neg. 37598; fragment, F 971889; Bolivia,
Prov. Caranges, A. C. V. d'Orbigny 1377 (P,
not seen).
This species appears to differ from G. dombey-
anum in possessing white phyllaries.
Gnaphalium melanosphaeroides Schultz-Bip. ex
Wedd., Chlor. And. 1: 148. 1856. TYPE: Peru,
Dept. Puno, Macusani, W. Lechler 1838 (ho-
lotype, P, not seen; isotype, w, not seen, F neg.
33150).
This species differs from G. dombeyanum in
possessing dark brown phyllaries.
Gnaphalium nanum Kunth, Nov. Gen. Sp. (fol.)
4: 62. 1820. TYPE: Peru, Dept. Piura, Prov.
Ayabaca, "juxta pagum Ayavaca, alt. 1410
hex.," A. Humboldt & A. Bonpland s.n. (ho-
lotype, P, not seen, IDC Microfiche 6209. 97:
I. 1, F neg. 15111; isotype, B, probably de-
stroyed).
The type consists of a single small stem and
quite possibly represents a depauperate specimen
of G. dombeyanum.
Gnaphalium lanuginosum Kunth, Nov. Gen. Sp.
(fol.) 4: 65. 1820. TYPE: Peru, Dept. Piura,
Prov. Ayabaca, "juxta pagum Ayavaca, alt.
1410 hex.," A. Humboldt & A. Bonpland s.n.
(holotype, P, not seen, IDC Microfiche 6209.
96: III. 7, F neg. 37602).
This species was collected at the same locality
as G. nanum and consists of a single stem (33 cm
long) with past bloom heads. No material resem
bling the specimen in the type photograph ha;
been encountered in this study.
Material here placed under Gnaphalium dom
beyanum has been previously annotated under ad
ditional names, including G. cheiranthifolium Lam
and G. graveolens Kunth. A discussion of thes(
two taxa is provided under Doubtful and Exclude*
Names.
AMAZONAS. Chachapoyas: ca. 43 km NE of Balsas
ca. 2900 m, Dillon & Turner 1721 (F.USM); Chachapoyas
2700 m, Williams 7546 (F, us). ANCASH. Bolognesi
abajo de Raquia, 1900 m, Sagdstegui et al. 12296 (F
HUT, MO). Huaraz: 4 km N of Monterrey, ca. 8400 ft
King& Collins 9021 (F,US); Cachabamba, 2870 m, Smiti
& Buddensiek 10934 (F). Huaylas: Parque Naciona
Huascaran, 4000 m, Bernardi et al. 16492 (us); Laguna
Llanganuco, 3500-3800 m, Gentry et al. 37405 (F, MO)
Zardini 1540 (MO). APURIMAC. Abancay: Sayhuite
Km 45 E of Abancay, 3600 m, Hutchison 1738 (F, NY
us, USM); 6-7 km NNW of Abancay, ca. 3100 m, Iltis d
Ugent 700 (MO, USM); Curahuasi, 2800 m, Vargas 126.
(F). AREQUIPA. Arequipa: ca. 40 km NNE of Arequip;
on road to Nevado Chachani, 3500-4000 m, Dillon e
al. 3277 (F); near Arequipa, 8000 ft, Harrison s.n. (GH)
southern slopes of Chachani, 2700 m, Hinkley & Hinkle_
45 (GH.US); Nevado de Chachani, 4100-4200 m, Fennel
13290 (F), 13294 (F, GH); Arequipa, campo de aviation
2700 m, Ponce 109 (USM); Km 18 on road between Ar
equipa and Yura, 3300 m, Solomon 2847 (MO); Chilen;
Valley, 7500 ft, Stafford 5 63 (F). Camana: 14 km on roa<
S of Chala, 350-400 m, Worth & Morrison 15688 (GH)
Caraveli: Lomas of Atiquipa, 150-200 m, Dillon & Dil
Ion 3790 (F, USM); Lomas de Pongo, cerca a Acari, 700-
800 m, Ferrerya 13426 (MO); Atico, 100 m, Tovar 266!
(USM). Islay: Lomas of Mollendo, ca. 8 km NW of Islay
ca. 510 m, Dillon & Dillon 3940 (F, USM); 10 km E o
Islay, 300 m, Worth & Morrison 15722 (F.GH.MO). AY A
CUCHO. La Mar: 40-46 km NNE of Tambo, 2710-
3050 m, Luteyn & Lebron-Luteyn 6359 (NY, us). Lucan
as: Aucara, Chinchay 3665 (us); entre Nazca y Puquio
3200-3300 m, Ferrerya 5483 (MO); Marcahuasi, arrib*
de Puquio, 3400-3450 m, Ferrerya 7213 (MO, us). CA
JAMARCA. Cajamarca: ca. 27 km NNW of Cajamara
on road to Hualgayoc, Pampa de Cerro Negro, 3600 m
Dillon & Molau 3017 (F); ca. 6.5 km SW of Cajamarca
2980 m, Dillon et al. 2901 (F, USM); ca. 22.5 km S o
Cajamarca on road to Chilete, 2740 m, Dillon et al
3036A (F). Celendin: 31 km NE of Cajamarca, 2950 m
Dillon & Turner 1588, 1589 (F); Jalca de Kumulca, 335(
m, Sagdstegui et al. 12038 (F, HUT, MO); Usnio, 3 120 m
Sanchez et al. 2776 (MO). Contumaza: La Herilla (Guz^
mango), 3050 m, Sagdstegui et al. 6456 (MO), 9696 (F
MO); Guzmango, 2600 m, Sagdstegui 7726 (MO). CUZ
CO. Paucartambo: Llulluchayoc, 3100-3700 m, Varga
4327 (F). Quispicanchis: 95 km above Quincemil on th
Rio Marcapata, 3480 m, Madison 1031 (GH). Urubamba
Chincheros, quebrada above Pojpoj waterfall, 3450-355C
m, Davis et al. 1474 (F, us); Chincheros, Taucca, 405C
m, Davis et al. 1575 (F); Chincheros, 3750 m, King e t
al. 120 (F,US); camino a Puyupatamarca, 3290 m, Drey-
36
FIELDIANA: BOTANY
fus 12831 (MO); Machupicchu, 2500-2600 m, Iltis et al.
1045 (us); Ollantaytambo, 3650 m, Peyton & Peyton 52
(F, MO p.p.); Machupicchu, 2675 m, Peyton & Peyton 201
(MO); Machupicchu, 2400 m, Vargas 819 (F). HUAN-
CAVELICA. Huancavelica: Orccon, arriba de Conaica,
3550-3580 m, Tovar 83 (us); Caniorccona, 3 km SE de
Conaica, 3560-3590 m, Tovar 98 (us p.p.). Tayacaja: 3
km N of Salcabamba, 3500 m, Stork & Morton 10328
(F). HUANUCO. Huanuco: Carpish Pass, ca. 2750 m,
Allard 20994 (F, us); Cerro Carpish, ca. 48.5 km N of
Huanuco, 8600 ft, Davidson & Jones 9188 (us); Carpish,
2700-2900 m, Ferrerya 2343 (USM); Mitotambo, arriba
de Mito, 3000-3100 m, Ferreyra 6678 (us, USM); Car-
pish, cumbre entre Huanuco y Tingo Maria, 2700-2900
m, Ferreyra 8154 (F, MO, us); Yanano, ca. 6000 ft, Mac-
bride 4944 (F, GH, us); Yanahuanca, Macbride & Feath-
erstone 1192 (F, us); Mito, ca. 9000 ft, Macbride & Feath-
erstone 1531 (F, us), 1801 (F, GH, us); N of Carpish Pass,
48.6 km NE of Huanuco, 2490 m, Plowman & Rury
11133 (F,US); Acomayo, Ridoutt 11499 (MO); Tumanga,
2400 m, Woytkowski 7958 (MO); Pillao, 2700 m, Woyt-
kowski 34149 (F). JUNIN. Conception: Chicchi, 3500-
3700 m, Ochoa 1989 (us). Huancayo: San Jeronimo,
3340 m, Proano 13 (MO, USM); Huancayo, cerca del Ce-
menterio, Ridoutt 10318 (MO p.p.); Huancayo, 3317 m,
Soukup 2955 (F.GH); Huancayo, 3800 m, Velarde 2342
(us). Tarma: 28-32 km NE of Tarma, ca. 2480 m, Dillon
& Turner 1348 A (F, USM); ca. 35 km NE of Tarma along
road to San Ramon, ca. 2300 m, Dillon & Turner 1408
(F, USM); Chuquishunca, 2 km arriba de Huacapistana,
2000-2400 m, Ferreyra 458 (MO, USM), 77242 (MO);
Chanchamayo valley, 1000 m, Schunke 412 (F); Aco-
bamba, 3500 m, Woytkowski 71 (F). Yauli: Oroya, Ka-
lenborn 42 (GH, MO, us), Kalenborn & Kalenborn 83 (NY);
La Oroya, ca. 12,000 ft, Macbride & Featherstone 979
(F, us); Banos, Capt. Wilkes s.n. (NY p.p.). LA LIBER-
TAD. Huamachuco: Rio Colorado, 3215 m, La Rosa &
Riccio 3519, 3520 (us); Marca Huamachuco, 3300-3400
m, La Rosa & Riccio 3601 (us). Otuzco: Cerro Ragache
(Salpo), 3200 m, Sagdstegui et al. 11595 (F, HUT, MO);
Shorey, Trujillo-Huamachuco, 3300 m, Smith & Vas-
quez 3271 (F, MO). Sanchez Carrion: Jalca de Quespuen-
da-Huamachuco, 3800 m, Sagdstegui 9432 (F, HUT, MO).
LAMBAYEQUE. Chiclayo: Cerro Reque, 540 m, Llatas
313 (F, HUT, MO). LIMA. Canete, Lomas de Quilmana,
entre Mala y Canete, 250-320 m, Ferreyra 6317 (MO).
Canta: La Florida, 2400-2500 m, Ferreyra 18423 (USM);
Ijadero, Canta-Lachoqui, 3700 m, Meza 108 (MO), 752
(USM); Llullum, 12 km arriba de Canta, 4100 m, Meza
210 (MO); near Antaicocha, Cerro Colorado, E of Canta,
3400-3800 m, Pennell 14631 (F); Huaros, 3400-3600
m, Pennell 14725 (F, GH, NY), 74726 (F); Purumarca,
Sanchez 48 (USM). Chancay: Lomas de Lachay, entre
Chancay y Huacho, 400-450 m, Ferreya 8774 (MO);
Chancay, Ridoutt 12208 (MO). Huarochiri: Surco, 3000-
3200 m, Ferreyra 676 (USM); Chicla, 3700 m, Ferreyra
6518 (us, USM); Matucana, ca. 8000 ft, Macbride &
Featherstone 298 (us), 360 (F, us p.p.); Rio Blanco, ca.
12,000 ft, Macbride & Featherstone 740 (F, us); Matu-
cana-San Mateo, 3000 m, Rauh & Hirsch P76 (NY p.p.);
Chicla, 3733m, Riccio 3716 (us). Yauyos: Huacracocha,
17 km N de Tupe, 4400 m, Cerrate 1225 (USM);
Cruzpampa, 2950 m, Cerrate & Tovar 1105 (F, MO, us).
MOQUEGUA. Mariscal Nieto: Quebrada de Torata,
Nunez 38 (USM). PIURA. Huancabamba: Hacienda Por-
culla, 1800-2000 m, Ferreyra 14175 (USM); Mitopampa,
Huancabamba-Cuello del Indio, 2650 m, Sagdstegui et
al. 8242 (MO, NY). PUNO. Azangaro: Amantani, 3900
m, Aguilar 438 (MO); Isla Arapa, Aguilar s. n. (MO). Huan-
cane: Miajachi, Isla Chirone, s.d. (MO). Melgar: Orurillo,
Soukup 503 (F, GH); Araranca, 4100-4300 m, Pennell
13472 (F, GH, NY, us, USM). Prov. Unknown: Tahuaco,
3850-3900 m, Tovar 5196 (USM).
4. Gnaphalium elegans Kunth, Nov. Gen. Sp. (fol.)
4: 63. 1820. TYPE: Ecuador, Prov. Pichincha,
"prope Chillo et Cachambamba, 1340 hex,
Jun" A. Humboldt & A. Bonpland 3001 (ho-
lotype, P, not seen, IDC Microfiche 6209. 96:
III. 1, F neg. 37591, fragment, F). Figure 11.
Gnaphalium poeppigianum DC., Prodr. 6: 227. 1838.
TYPE: Peru, Dept. Huanuco, Cuchero, 1829, E.
Poeppig 1368 (holotype, G-DC, not seen; IDC Mi-
crofiche 800. 1067: I. 1.; isotypes, B, probably
destroyed, F neg. 1 5 1 38, F, GH, w, not seen, Fneg.
33151).
Perennial herbs to 1 m tall; stems densely lanate
or tomentose. Leaves sessile, the base subamplex-
icaul; blade lanceolate to elliptic, 3.5-7 cm long,
5-20 mm wide, apex acute to attenuate, discolo-
rous, lower surface densely lanate, upper surface
glandular-puberulent, the margins entire to cren-
ulate. Capitulescences corymbose, terminal and
axillary. Capitula 5-7 mm high, 3-5 mm wide;
involucres ovoid to campanulate; phyllaries 3-4-
seriate, white or straminous, the outer ovate, ca.
2.5 mm long, ca. 1.5 mm long, apex rounded to
obtuse, the inner oblong to lanceolate, 4-7 mm
long, apex acute to apiculate; receptacles 2-2.5
mm wide; pistillate florets 80-100, the corolla ca.
3 mm long, base slightly expanded, apex minutely
3-4-cleft; hermaphroditic florets 5-18, the corolla
2.5-3 mm long, yellowish, 3-5-lobed. Achenes ca.
0.6 mm long, 5-ribbed; pappus bristles 34 mm
long. Chromosome number: n = 14 (Jansen et al.,
1984).
Gnaphalium elegans is distributed from Mexico
to Panama and throughout the Andes of northern
South America from Venezuela to southern Peru
(1400-3000 m). It is readily recognized by its ro-
bust habit, strongly discolorous cauline leaves, and
large capitula with white or rarely stramineous
phyllaries.
Several collections from central and northern
Peru are tentatively placed under this species (Fer-
reyra 5730, p.p.; Killip & Smith 21844; Sagdstegui
et al. 11516; Macbride & Featherstone 271). Their
leaves and bracts subtending the capitulescences
DILLON & SAGASTEGUI: FLORA OF PERU. V.
37
FIG. 1 1 . Gnaphalium elegans. A, habit; B, capitulum; C, outer phyllary; D, inner phyllary; E, pistillate floret; F,
hermaphroditic floret; G, achene. (Drawn from Lopez et al. 9031, HUT.)
are more similar to material placed in Gnaphalium
dombeyanum, but they have apiculate inner phyl-
laries and strongly 5 -ribbed achenes of typical G.
elegans. Further sampling in these regions may
help to define the nature of this variation.
AMAZONAS. Chachapoyas: 2 kms along road W of
Chachapoyas, ca. 7200 ft, King & Bishop 9177 (us). AN-
CASH. Bolognesi: Chiquian, 3500-3600 m, Ferrerya
5730 (us p.p.). CAJAMARCA. Cajamarca: ca. 22.5 km
S of Cajamarca on road to Chilete, 2740 m, Dillon et al.
3036 (F); Asuncion-San Juan, 2500 m, Sagdstegui et al.
38
FIELDIANA: BOTANY
10153 (F, HUT, MO). Contumaza: Tambo La Lima (Cas-
cas-Contumaza), 1 650 m, Lopez et al. 9031 (F, HUT, MO).
San Ignacio: Maisgua (San Ignacio-Nueva Esperanza),
Lopez et al. 8996 (F, HUT, K, MO, NY). CUZCO. Calca:
Lares-Manto, 2400 m, Marin 2289 (F). La Convencion:
Amaylamba, 1800 m, Mann 1514 (F, us). Urubamba:
Rio Urubamba, 2000 m, Angulo 1769 (HUT); Hacienda
Santa Rita, Dreyfus s.n. (USM 12815); Machupicchu, 2400
m, Ferreyra 2731 (us, USM), Machupicchu, 2400 m, Var-
gas 811 (F). HUANUCO. Huanuco: Carpish, 2650 m,
Asphund 13 119 (us); between Huanuco and Tingo Mar-
ia, 6 km N of Acomayo, 2450 m, Croat 57870 (F, MO).
Pachitea: Piedra Grande, estacion near Rio Santo Do-
mingo, ca. 5000 ft, Macbride 3688 (F, us). JUNIN. Tar-
ma: Huacapistana, 1800-1900 m, Ferreyra 11306 (MO,
USM); Tarma, 3000-3200 m, Killip & Smith 21844 (F,
NY, us). Prov. Unknown: Agua Dulce, 1600 m, Woyt-
kowski 7427 (GH, MO); Yucapata, 1400 m, Woytkowski
6622 (MO). LA LIBERT AD. Otuzco: Casmiche (Samne-
Otuzco), 1700 m, Sagdstegui et al. 11516 (F, HUT, K, MO,
NY). LIMA. Huarochiri: Matucana, 8000 ft (2424 m),
Macbride & Featherstone 271 (F, us). PIURA. Huanca-
bamba: Cerro Porculla, 3-11 km W of Continental Di-
vide down W slope, Edwin & Schunke 3764 (COL, F, NY,
us); 37 km E of Olmos on road to Pucara, 1 500 m, Gentry
et al. 22626 (F, MO). PUNO. Puno: region of Puno, Sou-
kup 518 (F). NO EXACT LOCALITY. "Peruvia," 1862,
Matthews s.n. (NY p.p.).
5. Gnaphaliutn lacteum Meyen & Walpers, Nov.
Actorum Acad. Caes. Leop.-Carol. Nat. Cur.
19: 276. 1843. TYPE: Chile, Prov. Tarapaca,
Tacora, F. J. F. Meyen s.n. (holotype, B, prob-
ably destroyed, F neg. 15105, fragment, GH).
Small, perennial herbs; stems prostrate,
branched, densely white- to gray-tomentose, leafy
to the apex, 5-10(-20) cm long. Leaves sessile;
blade linear to spatulate, 3-15(-40) mm long, 2-
3 mm wide, rarely decurrent, apex rounded, both
surfaces densely white-tomentose, the margins en-
tire. Capitulescences corymbose to corymbose-pa-
niculate in semiglobose clusters of 3-8 capitula,
terminal and axillary. Capitula 4-5 mm high, 3-
4 mm wide; involucres campanulate; phyllaries
white, the outer ovate, 2-3 mm long, 1.5-2 mm
wide, apex obtuse to rounded, the inner oblong,
ca. 4 mm long, 0.5-1.5 mm wide, apex rounded;
receptacles ca. 2 mm wide; pistillate florets 75-
100, the corollas 2.5-2.7 mm long; hermaphro-
ditic florets ca. 12, the corollas ca. 3 mm long.
Achenes ca. 0.6 mm long; pappus bristles ca. 3
mm long.
Gnaphalium lacteum is distributed from ex-
treme northern Chile and adjacent Argentina and
Bolivia to central Peru. It is distinguished by its
small, predominately prostrate habit and pure
white phyllaries with obtuse to rounded apices.
The overall size of individuals is variable, with
some plants having quite long, multi-branched
stems (e.g., Hinkley & Hinkley 3, GH). It superfi-
cially resembles other small prostrate species, e.g.,
G. badium and G. polium, and apparently occurs
sympatrically with G. polium in one locality. A
gathering by Pennell from Puno appears to be
mixed; two duplicates of Pennell 13444 (GH, NY)
are clearly G. lacteum, but the F sheet is clearly
different, and here assigned to G. polium. Another
collection from the same locality (Pennell 13470a,
F) is also assigned to G. polium and indicates that
these two species occur in the same geographic
area.
Cabrera (1978) placed Gnaphalium argyrolepis
Philippi under the synonymy of G. lacteum, and
based on the type photograph and description, it
appears that another species, G. frigidum Wedd.,
described from Tacora, Chile, may also be con-
specific with G. lacteum.
ANCASH. Bolognesi: Chiquian, 4000 m, Ferreyra
5825 (us, USM). AREQUIPA. Arequipa: southern slopes
of Chachani, north of Arequipa, 3600 m, Hinkley &
Hinkley 3 (GH); above Arequipa, 3000-3300 m, Pennell
13219 (F, GH, NY); Volcan Chachani, 3700 m, Rauh &
Hirsch P530 (NY). HUANCAVELICA. Castrovirreina:
near Cordova, 3050-3300 m, Metcalf 30266 (GH, F, MO).
LIMA. Canta: carretera Canta-Lachaqui, 3600 m, Meza
211 (MO). PUNO. Melgar: Araranca, 4100-4300 m,
Pennell 13444 p.p. (GH, NY). NO EXACT LOCALITY.
1838-42, Wilkess.n. (NY).
6. Gnaphalium luteo-album L., Sp. PI. 851. 1753.
TYPE: Europe, no exact locality, (holotype,
LINN, 900. 286-294, not seen).
Annual herbs to 30-50 cm tall; stems white-
lanate pubescent, erect to decumbent. Basal leaves
sessile; blade oblong-spatulate, ca. 6 cm long, ca.
6 mm wide, apex obtuse; cauline leaves sessile,
subamplexicaul to subdecurrent; blade lanceolate,
1-5 cm long, 2-5 mm wide, apex acute, all arach-
noid-tomentose. Capitulescences corymbose or
subcymose, congested. Capitula 2.5-3(-4) mm
high, 2.5-4 mm wide; involucres campanulate;
phyllaries 2.5-3(-4) mm long, ca. 1 mm wide, ob-
long, apex obtuse; pistillate florets 100-120, the
corollas ca. 2.5 mm long; hermaphroditic florets
ca. 30, the corollas ca. 2.5 mm long. Achenes ca.
0.5 mm long, minutely pubescent with multicel-
lular, biseriate trichomes; pappus bristles ca. 2.5
mm long. Chromosome number: n = 1 (Kockx-
van Roon, M. & J. H. Wieftering, 1982).
DILLON & SAGASTEGUI: FLORA OF PERU. V.
39
Gnaphalium luteo-album is native to central and
southern Europe and has been introduced infre-
quently in South America. In Peru, it is repre-
sented only by a few collections from the vicinity
of Arequipa. It is readily distinguished from native
members of the genus by its annual habit, small
capitula, and achenes with multicellular, biseriate
trichomes.
AREQUIPA. Arequipa: Cerrillo Savandia (Lavan-
cha?) near Laspinas, 14 km N of Arequipa, 2200 m,
Eyerdam & Beetle 22147 (MO); vicinity of Arequipa,
Rose & Rose 18828 (GH, NY, us), 19011 (NY).
7. Gnaphalium polium Wedd., Chlor. And. 1: 147.
1856. TYPE: Peru, Dept. Cuzco, no exact lo-
cality, C. Gay 1709 (holotype, P, not seen, F
neg. 37605, fragment, F).
Prostrate to decumbent, perennial herbs; stems
suffrutescent, much-branched at the base, 5-1 0(-
20) cm long. Leaves sessile or slightly decurrent
on stem; blade linear to spatulate, 5- 15 (-20) mm
long, 2-2.5(-3.5) mm wide, apex rounded to acute,
both surfaces densely tomentose. Capitulescences
corymbose in globose clusters, terminal. Capitula
ca. 4 mm high, ca. 4 mm wide; phyllaries 3-5-
seriate, the outer ovate, 2.5-3 mm long, ca. 1.5
mm wide, apex acute, the inner oblong, 3.54 mm
long, 1-1.5 mm wide, apex obtuse to rounded;
pistillate florets ca. 100, the corollas ca. 2.2 mm
long, 3-4-lobed; hermaphroditic florets 8-12, the
corollas ca. 2.2 mm long, 5-lobed. Achenes 0.5-
0.7 mm long; pappus bristles ca. 3 mm long.
Gnaphalium polium is a highly variable and
possibly polyphyletic assemblage of small, high-
elevation perennials distributed from northern Peru
to Bolivia. Its thickened lignescent stems and dense,
white-tomentose pubescence that gives the leaves
a thickened circular or rounded form are its most
readily observable characteristics. It is readily dis-
tinguished from the annual, G. badium, in not
having its capitulescences immersed in the upper
leaves as in that species.
Urubamba: Machupicchu, 3350 m, Peyton & Peyton 709
(F, MO); Salcantay, 4500 m, Rauh & Hirsch P1424 (NY).
JUNIN. Janja: Hacienda San Juan, entre Janja y Ma-
tahuasi, 3200-3300 m, Ferreyra 12901 (MO). LA LIB-
ERTAD. Santiago de Chuco: Chota (Motil-Shorey), 3200
m, Sagdstegui et al. 11700 (F, HUT, MO). LIMA. Huar-
ochiri: Casapalta [Casaplaca], Ball s.n. (GH); 20 km NE
of San Mateo, 4050 m, Gentry et al. 19197 (MO). PUNO.
Azangaro: Tequena, 20 km N de Arapa, Aguilar 417
(MO, USM). Melgar: Araranca, 4100-^4300 m, Pennell
13444 (F), 13470 (F), 13470a (F).
Doubtful and Excluded Species of
Gnaphalium in Peru
Gnaphalium cheiranthifolia Lam., Encyl. Method.
2: 752. 1786. TYPE: Uruguay, Montevideo,
May 1767, P. Commerson s.n. (holotype, P-
LAM, not seen, F neg. 37587, fragment, F).
Pseudognaphalium cheiranthifolia (Lam.) Milliard &
Hurt, J. Linn. Soc. Hot. 82(3): 205. 1981.
Cabrera (1978) considered this species to be a
southern South American element extending only
into southern Bolivia, Brazil, Argentina, and Chile.
It is a commonly used name throughout northern
South America; the large basal rosette in the pho-
tograph of the holotype has not been observed in
any Peruvian plants.
Gnaphalium frigidum Wedd., Chlor. And. 1: 147.
1856. TYPE: Bolivia, Dept. Botosi: A. C. V.
d'Orbigny 1372 (lectotype, P, designated by
Cabrera (1978), not seen, F neg. 37594).
From an examination of the type photograph
and the description, this species appears to be quite
close to G. lacteum, and the two may prove to be
conspecific. Material assigned to G. frigidum agrees
with G. lacteum in most respects but lacks the pure
white phyllaries associated with the latter species.
It is not placed into formal synonymy pending
examination of type material.
ANCASH. Huaraz: Cerro San Cristobal, 3800 m,
Evangelista s.n. (F, MO); Huascaran National Park, Que-
brada Shallap, 3690-4100 m, Smith et al. 9646 (F, MO);
Quebrada Llaca, 4400-4600 m, Smith et al. 10824 (F
P.P., MO). Recuay: Km 125, Pativilca to Recuay, 3870 m,
Dillon et al. 31 10 (F, HUT, MO, NY, TEX, us, USM). CA-
JAMARCA. Contumaza: Pozo Kuan, 3600-3800 m, Sa-
gdstegui et al. 10053 (F, HUT, MO). CUZCO. Quispican-
chis: Auzangate, 4700 m, Rauh & Hirsch PI 149 (NY).
Gnaphalium glandulosum Klatt, Linnaea 42: 129.
1878. TYPE: Chile ["Peru"], Tacora, F. J. F.
Meyen s.n. (holotype, B, probably destroyed,
F neg. 15099, fragment, GH).
No material referable to this Chilean species has
been examined in this study.
40
FIELDIANA: BOTANY
Gnaphalium graveolens Kunth, Nov. Gen. Sp. (fol.)
4: 64. 1820. TYPE: Ecuador, Prov. Chim-
borazo, Riobamba, A. Humboldt & A. Bonp-
land 3193 (holotype, P, not seen, IDC Micro-
fiche 6209. 96: III. 2, F neg. 37586), non G.
graveolens M. Bieb. ( 1 808, 1 8 1 9), nee G. grav-
eolens Fenzl ex Schultz-Bip. (1835- 1 850), nee
G. graveolens Henning (1823).
This is one of the most commonly applied names
for Andean Gnaphalium', however, it is invalid
due to an earlier homonym. The type photograph
appears to show some characters of G. elegans,
including its large discolorous leaves and large
spreading capitulescences. Most of the material
previously annotated under this name is here placed
with G. dombeyanum.
X. Helichrysum
Helichrysum Miller, Gard. Diet., ed. 4, 1754.
TYPE: Gnaphalium orientale L. = Helichry-
sum orientate (L.) Gaertn.
Perennial herbs or shrubs, cespitose or suffru-
tescent, usually lanate, arachnoid, or glandular-
pubescent. Leaves alternate, simple, sessile or sub-
petiolate, the margins entire to subentire. Capi-
tulescences of solitary heads or compound cor-
ymbs or panicles. Capitula disciform or discoid,
heterogamous, female florets fewer than hermaph-
roditic; involucres cylindrical-campanulate to
hemispherical; phyllaries multiseriate, imbricate,
persistent, scarious, white or colored distally; re-
ceptacles flat, glabrous; pistillate florets (2-)45-
55(-850), the corollas narrowly tubular, rarely fi-
liform; hermaphroditic florets 20-200(-700), the
corollas tubular, yellow, 5-lobed, the anther bases
sagittate, caudate, the terminal appendages oblong
to lanceolate, obtuse to rounded, the styles bifid,
the branches spatulate, flattened, apex deltoid, the
nectary often prominent. Achenes cylindrical,
glandular-papillose; pappus bristles scabrid, bar-
bellate, uniseriate, free or united at the base, ca-
ducous, stramineous to white. Chromosome num-
bers: n = 7, 8, 10, 1 1, 14, 15, 28, 38.
Helichrysum is a genus of perhaps 500 species
widely distributed in Eurasia, Africa, Madagascar,
Australia, and New Zealand. It is closely allied to
Gnaphalium and the latter is weakly distinguished
from it by the number of pistillate florets outnum-
bering the hermaphroditic florets. Several mem-
bers are cultivated as ornamentals in the New
World, the following throughout Latin America.
1 . Helichrysum bracteatum (Vent.) Andrews, Bot.
Rip. Sup. 1, t. 428. 1805.
Xeranthemum bracteatum Vent., Jard. Malm,
t. 2. 1803. TYPE: not seen.
Perennial herbs to 1 m tall, glabrate or puber-
ulent with short glandular trichomes and sparse,
arachnoid hairs; stems striate. Leaves alternate,
sessile or the base narrowed to an indistinct winged
petiole; blade oblanceolate, to 12 cm long, 1.5-2
cm wide, the margins entire and undulate, both
surfaces sparsely arachnoid-pubescent and glan-
dular. Capitulescences of solitary heads, terminal
and axillary, braceolate. Capitula ca. 2 cm high,
2.5-5 cm wide; involucres hemispheric; phyllaries
multiseriate, subequal or the innermost shorter,
oblong to lanceolate, 10-15 mm long, 3-5 mm
wide, apex rounded to acute, variously red, orange,
yellow, purple, or white, the innermost usually
pale; receptacles flat, 15-25 mm in diameter; pis-
tillate florets few or rarely absent; hermaphroditic
florets usually over 500, the corollas tubular, 8-
10 mm long, the limb slightly expanded, 5-lobed,
the base expanded at the nectary. Achenes fusi-
form to cylindrical, ca. 2 mm long, striate, dark
brown, apex dilated; pappus bristles ca. 8 mm
long, stramineous to white. Chromosome num-
bers: n = 12, 14.
This species is widely cultivated as an orna-
mental in flower gardens and for sale in larger
cities. It is known as "immortal." No herbarium
specimens collected in Peru have been examined
in the present study.
XI. Jalcophila
Jalcophila Dillon & Sagast., Brittonia 38: 162.
1986. TYPE: Jalcophila peruviana Dillon &
Sagast.
Cespitose, perennial herbs; stems branched,
densely compact. Leaves alternate, rosulate, ses-
sile, marcescent in age, coriaceous; blades ovate-
lanceolate to lanceolate or obdeltate, 3.5-9 mm
long, 0.5-2 mm wide, lower surface tomentose-
DILLON & SAGASTEGUI: FLORA OF PERU. V.
41
arachnose, 3-costate, upper surface glabrous, 2-
canaliculate. the margins entire. Capitulescences
of solitary heads, sessile at an thesis, becoming pe-
dunculate at maturity, the peduncles 1-5 mm long,
sparsely arachnose. Capitula discoid, heteroga-
mous; involucres ovoid to campanulate; phyllaries
2-seriate, equal to subequal, scarious, strami-
neous, ovate-lanceolate to lanceolate; receptacle
plane; pistillate florets 4-6, the corollas filiform,
the style branches linear, exserted; hermaphroditic
florets 2-6, the corollas tubular, the limb slightly
expanded, 4-5-lobed, the anther bases sagittate,
the terminal appendage lanceolate, the style
branches truncate to clavate, papillose, exserted,
the stigmatic lines short and parallel. Achenes
ovoid, 4-costate, appearing glabrous, scattered
biseriate, multicelluar, glandular trichomes; pap-
pus bristles uniseriate, the bases fused.
Jalcophila contains two species and is known
from high-elevation localities in the paramo of
Volcan Galeras (Dept. Narino) in southwestern
Colombia, Paramo El Angel (Prov. Carchi) in
northern Ecuador, the paramos of the Cordillera
de los Llangantes (Prov. Napo) in central Ecuador,
and Pampas de la Julia (Dept. La Libertad) in
north-central Peru. This genus appears to lack close
relatives among Central and South American In-
uleae (Dillon & Sagastegui, 1986).
Reference
DILLON, M. O., AND A. SAGASTEGUI A. 1986. Jalco-
phila, a new genus of Andean Inuleae (Asteraceae).
Brittonia, 38: 162-167.
1 . Jalcophila peruviana Dillon & Sagast. . Britton-
ia 38: 163. 1986. TYPE: Peru, Dept. La Lib-
ertad, Prov. Santiago de Chuco, arriba de Qui-
ruvilca, Pampas de la Julia, 3800 m, 15 Nov
1983, A. Sagastegui A., J. Mostacero L., S.
Lopez M. & S. Leiva G. 11131 (holotype, HUT ;
isotype, F). Figure 12.
Cespitose herbs; stems 1-2 cm long. Leaves
ovate-lanceolate to obdeltate, 3.5-5 mm long, 0.5-
1.5 mm wide, apex rounded, often with tomentose
tuft. Capitulescences of solitary heads, peduncu-
late, the peduncles 1-3 mm long. Capitula 2-2.5
mm high, 1-1.4 mm wide; involucre campanulate;
phyllaries 8-13, subequal, the outer ovate-lance-
olate, 2-2.3 mm long, 0.5-0.6 mm wide, apex acute
to obtuse, lacerate, the inner lanceolate, 1.8-2.1
mm long, 0.4-0.5 mm wide, apex acute; pistillate
florets ca. 4, the corollas ca 1.25 mm long, the
style branches ca. 0.2 mm long; hermaphrodite
florets ca. 2, the corollas ca. 1.75 mm long, the
limb 5-lobed, the lobes 0.25 mm long, the anthers
ca. 1 mm long, the style branches truncate, ca. 0. 1
mm long. Achenes ovoid, ca. 0.75 mm long, ca.
0.4 mm wide; pappus bristles 1.2-1.5 mm long.
Jalcophila peruviana is known only from the
type and one additional locality in northern Peru.
Its small, cespitose habit makes it an inconspic-
uous element within the jalca formation. It su-
perficially resembles Cuatrecasasiella isernii in
general habit; however, it is easily separable from
the latter by having alternate leaves and heterog-
amous capitula. The achenes of this genus were
originally described as gabrous; however, they do
possess scattered biseriate, multicellular, glandu-
lar trichomes common in the tribe.
LA LIBERTAD. Santiago de Chuco: Chota-Shorey,
3250 m, Sagastegui et al. 11104 (F, HUT, MO, NY).
XII. Loricaria
Loricaria 2 Wedd., Chlor. And. 1: 165. t. 27. 1856.
TYPE: Conyza thuyoides Lam. = Loricaria
thuyoides (Lam.) Schultz-Bip.
Shrubs or subshrubs. dioecious, stems erect,
branched, densely foliaceous, laterally com-
pressed, narrowly rhomboid to subterete in sec-
tion. Leaves alternate, distichous, sessile; blade
laterally compressed, coriaceous, often dorsally
carinate, strongly imbricate, appressed to stems,
the margins entire, involute. Capitulescences of
solitary heads, axillary or terminal. Capitula dis-
coid, homogamous, sessile; involucre campanu-
late to cylindrical; phyllaries 2-5 -seriate, imbri-
cate, subequal; receptacles paleaceous or naked;
masculine florets functionally staminate, the co-
rollas tubular, 5 -dentate, the anther bases sagit-
tate, caudate, the styles undivided or bifid, the
ovary sterile, the pappus bristles uniseriate, the
bases fused, the apices clavellate; feminine florets
with corollas filiform, 5-fid, the style branches lin-
ear, acute, exserted, the ovary fertile, the pappus
bristles uniseriate, the bases fused, the apices slen-
der and acute. Achenes cylindrical, 4-5-costate,
For additional synonymy, see Cuatrecasas (1954).
42
FIELDIANA: BOTANY
FIG. 1 2. Jalcophila peruviana. A, habit; B, leaf (lower surface); C, capitulum; D, phyllary; E, hermaphroditic floret;
F, anthers; G, style and achene of pistillate floret; H, achene. (Drawn from Sagdstegui et al. 11131, HUT.)
the surface reticulate, usually glabrous, rarely pu-
bescent with bicellular, capitate-glandular trich-
omes (e.g., L. graveolens).
A genus with approximately 1 9 species distrib-
uted in high-elevation Andean habitats from Co-
lombia to Bolivia. Nine species and one variety
are currently recognized from Peru.
Loricaria is distinctive among South American
Inuleae in possessing a shrubby habit with laterally
compressed, foliaceous stems and densely imbri-
cate, involute, coriaceous leaves. The frequent oc-
currence of paleaceous receptacles is a character
DILLON & SAGASTEGUI: FLORA OF PERU. V.
43
not found in other Peruvian Inuleae. Cuatrecasas
(1954) recognized three sections based primarily
upon position of capitula, either terminal or ax-
illary, the presence or absence of receptacular pa-
leae, and achene pubescence.
DILLON, M. O., AND A. SAGASTEGUI A. 1986. Ne\
species and status changes in Andean Inuleae. Phy
tologia, 59: 227-233.
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.
References
CUATRECASAS, J . 1954. Synopsis der Gattung Loricaria
Wedd. Feddes Repert. Spec. Nov. Regni Veg., 56: 149-
172.
Key to Species of Loricaria
1 . Capitulescences of solitary, terminal heads ',
2. Foliaceous stems slightly flattened, 2-2.5 mm wide (including leaves); leaves ovate in lateral view
2-2.5 mm long, ca. 1 mm wide, apex obtuse L. leptothamnc
2. Foliaceous stems strongly flattened, 4-9 mm wide (including leaves); leaves triangular or linea
in lateral view, 4-8 mm long, 1.5-2.5 mm wide, apex acute or subacute '.
3. Leaves triangular or triangular-oblong in lateral view, apices acute <
4. Foliaceous stems 7-9 mm wide (including leaves); leaves 7-8 mm long, ca. 2 mm wide
vernicose-ferruginous L. ferruginet
4. Foliaceous stems 45 mm wide (including leaves); leaves 5-8 mm long, 2-2.5 mm wide
olivaceous green L. lycopodinei
3. Leaves linear-oblong in lateral view, apices subacute '.
5. Foliaceous stems 5-7 mm wide (including leaves); leaves 5-8 mm long, 2-2.5 mm wide
lustrous, green to yellowish white L. macbride
5. Foliaceous stems 4-5 mm wide (including leaves); leaves 4-7 mm long, ca. 1.5 mm widi
brownish roseate L. lucidc
1 . Capitulescences of solitary, axillary heads, or thyrsiform (
6. Capitulescences thyrsiform L. thyrsoidei
6. Capitulescences of solitary heads, axillary
7. Foliaceous stems subterete; leaves with dorsal surfaces tomentose; achenes pubescent will
bicellular, capitate-glandular trichomes L. graveolen.
1. Foliaceous stems flattened; leaves with dorsal surfaces glabrous; achenes glabrous
L. thuyoide.
1. Loricaria ferruginea (Ruiz & Pavon) Wedd.,
Chlor. And. 1: 166. 1856. Figure 13.
Molina ferruginea Ruiz & Pavon, Syst. Veget. 211.
1 798. TYPE: Peru, "Cordilleras et Punas per Tar-
mae Cantae et Huarocheri Provincias," H. Ruiz
& J. Pavon 28/10 (holotype, MA, not seen, F neg.
49225; isotype, F).
Shrubs to 80 cm tall, much-branched; stems 7-
9 mm wide. Leaves triangular in lateral view, 7-
8 mm long, ca. 2 mm wide, dorsally carinate, gla-
brous, vernicose-ferruginous, inner surface lanate.
Capitulescences of solitary, terminal heads. Mas-
culine capitula 6-7 mm high, 4-5 mm wide; in-
volucres campanulate; phyllaries 15-22, ovate-
oblong to oblong-linear, 3.5-5 mm long, 1-2 mm
wide, apex obtuse to rounded, glabrous; florets 1 7
20, the corollas ca. 4 mm long; pappus bristle
clavellate, ca. 4 mm long; paleae ca. 6, linear, ca
5 mm long, scarious, apex obtuse. Feminine ca
pitula 6-7 mm high, 2.5-3 mm wide; involucre
cylindrical; phyllaries 15-18, ovate-oblong to ob
long-linear, 4-6.5 mm long, 1-2 mm wide, ape:
obtuse, lacerate; florets 12-22, the corollas 44.
mm long; paleae ca. 12, lanceolate-linear, ca. i
mm long, apex acute, lacerate. Achenes 1-1.5 mn
long, glabrous; pappus bristles acute, ca. 5 mn
long.
Loricaria ferruginea is distributed from centra
Ecuador to central Peru (3300-4800 m). It is readi
ly distinguished by its vernicose, ferruginous leaves
terminal capitula, and paleaceous receptacles.
44
FIELDIANA: BOTANY
K
B
FIG. 13. Loricaria lucida. A, habit. (Drawn from Macbride & Featherstone 2171, F). Loricaria leptothamna. B,
portion of stem with capitulum; C, lateral view of leaf; D, outer phyllary; E, inner phyllary; F, pistillate floret; G,
masculine floret. (Drawn from Lopez & Sagdstegui 3341, HUT). Loricaria ferruginea. H, portion of stem with capit-
ulum; I, lateral view of leaf; J, outer phyllary; K, inner phyllary; L, pistillate floret; M, masculine floret. (Drawn from
Lopez 8076, HUT.)
DILLON & SAGASTEGUI: FLORA OF PERU. V.
45
This species is purported to have medicinal
properties and is often sold in herb markets. Label
data (Sagdstegui & Skillman 12782) indicate that
the ". . . entire plant is boiled in water and the
liquid is then drunk. Used to change a person's
character, to make him stronger." In Peru, it is
known by the following common names: chin-
chango, huallpapa-chaquin, palmilla blanca, and
pata de gallo.
ANCASH. Bolognesi: Cordillera Raura, 4300 m, Rauh
& Hirsch P1813 (NY). Carhuaz: Huascaran National Park,
Quebrada Ulta, 4700 m, Smith 11324 (F,MO); Quebrada
Ishinca, 4400 m, Smith et al. 9534 (F, MO); Vinoyapam-
pa, Quebrada Honda, 4200 m, Smith et al. 11600 (F,
MO). Huaraz: Laguna de Shurup, 3800 m, Evangelista
s.n. (F, HUT, MO); Cerro San Cristobal, 3800 m, Evan-
gelista s.n. (F, HUT, MO); Huascaran National Park, Que-
brada Llaca, 4500 m, Smith et al. 8994 (F, MO). Huari:
Huascaran National Park, 3-4 km below Cahuish Tun-
nel, 4500-4550 m, Smith & Buddensiek 11120 (F, MO);
2 km from Cahuish Tunnel, 4400 m, Smith & Goodwin
8746 (F, MO); Quebrada Pucaraju, 3900-4200 m, Smith
et al. 12673 (F, MO); Quebrada Rima Rima, 4200-4440
m, Smith et al. 12302 (F, MO). Huaylas: Huascaran Na-
tional Park, Paron Valley, E of lake, 4220-4600 m, Smith
11518 (F.MO); Quebrada Alpamayo, 4600-4750 m, Smith
et al. 9755 (F, MO). Recuay: Huascaran National Park,
Rio Pachacoto drainage, Ojo de Puma spring, 4200 m,
Smith 11433 (F, MO); Quebrada Queshque, 4550-4600
m, Smith et al. 11893 (F). Yungay: Laguna de Llac, 4200
m, Antenuez de Mayolo 321 (F, uc, us, USM); Parque Na-
tional Huascaran, Porta Chiello, Bernardi et al. 16539
(F); Laguna Llanganuco, 4200-4800 m, Gentry et al.
37430 (F, HUT, MO); Huascaran National Park, between
Lake Llanganuco and Portachuelo, 4420-4620 m, Smith
8257 (F,MO); Quebrada Ranincuray, 4000-4300 m, Smith
et al. 9143 (F, MO); Quebrada Demanda, 4800 m, Smith
& Cautivo 10271 (F, MO); Laguna Llanganuco-Porta-
chuelo, 4000-5000 m, Zardini 1553 (LP, MO). CAJA-
MARCA. Cajamarca: Km 45 Cajamarca-Bambamarca,
3640 m, Antunez de Mayolo 364 (uc, us); lagunas de la
Choclla, 3800 m, Llatas & Vdsquez 870 (F, PRO). San
Miguel: La Quinua-Calquis (jalca), 3300 m, Sagdstegui
et al. 8851 (F, HUT, MO). LA LIBERT AD. Pataz: Paso de
Alaska, carretera a Tayabamba, 3900 m, Lopez & Sa-
gdstegui 8178 (GH, HUT, MO). Santiago de Chuco: Jalca
de Quiruvilca, 4000 m, Lopez 8076 (F, GH, HUT, MO, NY);
Pampas de la Julia (Quiruvilca-Huamachuco), 4000 m,
Sagdstegui & Fabris 7572 (HUT, MO, NY), Sagdstegui et
al. 11116 (HUT, MO). PASCO. Pasco: cerca de Huallay
(Junin), ca. 4500 m, Koepcke s.n. (F, USM). LAMBA-
YEQUE. Ferrenafe: Incahuasi, Sagdstegui & Skillman
12782 (F, HUT).
nio," 17,000ft, 1854, W. Lechler 7S75(holotype,
B, probably destroyed; isotypes, F.G-DEL, not seen,
F neg. 28509, NY, p, not seen).
Shrubs to 30-50 cm tall, much-branched; stems
subterete, ca. 1 .5 cm wide (including leaves). Leaves
broadly ovate, 5-6 mm long, 34 mm wide, dor-
sally rounded, dorsal surface tomentose, glandu-
lar, inner surface lanate. Capitulescences of soli-
tary, axillary heads. Masculine capitula 4.5-5.5
mm high, 2-3 mm wide; involucres campanulate;
phyllaries 12-14, obovate-oblong, 4-4.5 mm long,
1-1.5 mm wide, apex expanded and rounded, lac-
erate; receptacles epaleaceous; florets 6-14, the co-
rollas 3-4 mm long; pappus bristles clavellate, 4-
4.5 mm long. Feminine capitula 6-7 mm high,
1.5-2.5 mm wide; involucres cylindrical; phyllar-
ies 13-15, linear, 5-7 mm long, 0.8-1.2 mm wide,
apex obtuse or acute, lacerate; receptacles epalea-
ceous; florets 6-12, the corollas 3.54 mm long;
pappus bristles acute, ca. 5 mm long. Achenes ca.
1 mm long, pubescent with bicellular, capitate-
glandular trichomes.
Loricaria graveolens is endemic to Peru, occur-
ring in jalca or puna formations from La Libertad
to Puno (3900-4400 m). It is distinguished by its
densely tomentose leaves, strictly axillary capitula,
and achenes pubescent with bicellular, capitate-
glandular trichomes.
This species is purported to have medicinal
properties and is sold in herb markets, as indicated
by label data (Cerrate 2920). In Peru, it is known
by the following common names: pata de gallo,
palmita, hualcayo, tola, and quera.
AREQUIPA. Arequipa: Charchani [Chachani], 16,400
ft, Douglass 5 (F, GH, NY); Rauh & Hirsch P544 (NY);
Pichupichu, 13,500 ft, Strafford689 (F). Cailloma: Cail-
loma, Schmidt s.n. (F); Sumbay, Schmidt s.n. (F). CUZ-
CO. Canchis: La Raya, 4400-4500 m, Pennell 13517 (F,
GH, NY, us). Espinar: Yauri, 3900 m, Vargas 4380 (F).
LA LIBERTAD. Santiago de Chuco, Quiruvilca, 4100
m, Lopez 1140 (F, USM). LIMA. Yauyos: Huacracocha,
a 17 kms de Yauyos, 43504400 m, Cerrate & Tovar
1227 (F, MO, USM). PUNO. Azangaro: Checayani [Che-
cayane], 4000 m, Ellenberg 548 (us). Melgar: Orurillo,
Soukup 578 (F, us); Lago Titicaca, Monheim M45 (NY).
2. Loricaria graveolens (Schultz-Bip.) Wedd.,
Chlor. And. 1: 167. 1856.
3. Loricaria leptothamna (Mattf.) Cuatr., Feddes
Repert. Spec. Nov. Regni Veg. 56: 163. 1954.
Figure 13.
Baccharis graveolens Schultz-Bip., Bonplandia 4: 51.
1856. TYPE: Peru, Dept. Cuzco, San Antonio,
"rupium fissuris Cordiller. summ. pr. S. Anto-
Tafalla leptothamna Mattf., Notizbl. Bot. Gart. Ber-
lin-Dahlem 10: 775. 1929. TYPE: Peru, Dept. La
Libertad, Prov. Pataz, entre Tayabamba y Valle
46
FIELDIANA: BOTANY
de Mixiollo [Mishollo], afluente izquierda del
Huallaga, 3600-3700 m, 2 Aug 1914, A. Weber-
bauer 7027 (holotype, B, probably destroyed; iso-
types, F, F neg. 49220, OH, F neg. 40754).
Shrubs to 80 cm tall, much-branched; primary
stems ca. 1 cm in diameter, the foliaceous stems
flattened, 2-2.5 mm wide (including leaves).
Leaves ovate in lateral view, ca. 2.5 mm long, ca.
1 mm wide, dorsally rounded, dorsal surface gla-
brous, brownish green, inner surface lanate. Ca-
pitulescences of solitary, terminal heads. Mascu-
line capitula ca. 4 mm high, ca. 3 mm wide;
involucres campanulate; phyllaries 12-13, obo-
vate-oblong to oblong, 34 mm long, 0.5-1.5 mm
wide, apex expanded and rounded, lacerate; re-
ceptacles epaleaceous; florets ca. 10, the corollas
2.5-3 mm long; pappus bristles clavellate, 3-3.5
mm long. Feminine capitula 5-6 mm high, ca. 3
mm wide; involucres cylindrical; phyllaries ca. 1 2,
elliptic-oblong to oblong-linear, 3.5-5.5 mm long,
1-1.5 mm wide, apex acute, lacerate; receptacles
epaleaceous; florets 3-9, the corollas 3.54 mm
long; pappus bristles acute, ca. 4 mm long. Achenes
1-1.5 mm long, glabrous.
Loricaria leptothamna is endemic to Peru and
known only from a few collections in Departments
of Amazonas, Cuzco, and La Libertad. It has the
shortest leaves of any Peruvian species, ap-
proached only by those of L. thuyiodes var. mi-
crophylla (Wedd.) Cuatr. and L. ollgaardii Dillon
& Sagast. of Colombia and Ecuador, respectively.
This character, combined with terminal capitula
and epaleaceous receptacles, makes L. leptotham-
na a readily distinguishable member of the genus.
A series of collections from between Cajamarca
and Hualgayoc superficially match L. leptotham-
na; however, they differ in several characteristics
including: (1) leaves more ovate-oblong in lateral
view and dorsally convex, and (2) masculine ca-
pitula with fewer florets (5 versus 12). Further
study may prove these collections worthy of tax-
onomic recognition at subspecific rank in L. lep-
tothamna.
AMAZONAS. Chachapoyas: ca. 30 km SW of Lei-
mebamba on road to Balsas, 3540 m, Dillon & Turner
1738 (F, HUT, MO, USM); Jalca de Calla-Calla, 3650 m,
Sagdstegui 7480 (HUT, us); Sanchez & Ruiz 548 (CPUN,
F). CAJAMARCA. Cajamarca: La Pajuela (carretera Ca-
jamarca-Hualgayoc), 3500 m, Lopez et al. 7832 (HUT,
MO-2); ruta Cajamarca-Hualgayoc, 3500 m, Sdnchez 776
(CPUN, F) . Celendin: cumbre Komulca, entre Cajamarca
y Celendin, 3800-3850 m, Ferrerya 13250 (USM). LA
LIBERTAD. Bolivar: Travesia Laplap, 3600 m, Lopez
& Sagdstegui 3341 (GH, HUT, MO, us). CUZCO. La Con-
vencion: Cordillera Vilcabamba, 3400 m, Dudley 11185
(F, MO, USM).
4. Loricaria lucida Cuatr., Feddes Repert. Spec.
Nov. Regni Veg. 56: 163. 1954. TYPE: Peru,
Dept. Huanuco, Prov. Huanuco, 15 mi NE
of Huanuco, about 15,000ft, 12-22 Jun 1922,
J. F. Macbride & W. Feather stone 2171 (ho-
lotype, F, F neg. 40751; isotypes, GH, us). Figure
13.
Shrubs 50-80 cm tall, much-branched; stems
flattened, 4-5 mm wide (including leaves). Leaves
linear-oblong in lateral view, 4-7 mm long, ca. 1 .5
mm wide, dorsally carinate, apex subacute to ob-
tuse, dorsal surface glabrous, vernicose, reddish
brown, inner surface lanate. Capitulescences of
solitary, terminal heads. Masculine capitula 4.5-
5 mm high, ca. 3 mm wide; involucres campan-
ulate; phyllaries ca. 13, ovate to ovate-oblong, 3-
4.5 mm long, 1.5-2 mm wide, apex obtuse, lac-
erate; florets 1 1-13, sterile, the corollas 3.5-4 mm
long; pappus bristles clavellate, 3-3.5 mm long;
paleae oblong, ca. 4 mm long, scarious, apex ob-
tuse. (Feminine individuals not observed.)
Loricaria lucida is a rare Peruvian endemic
known only from the type and one other collection
from central Peru. It most closely resembles L.
ferruginea but differs from that species in having
much narrower stems (4-5 mm versus 7-9 mm)
and linear-oblong leaves. Feminine individuals
were not encountered by Cuatrecasas (1954) or in
this study.
JUNIN. Huancayo: Santo Domingo de Acobamba,
4300 m, Ochoa 166 (F, USM).
5. Loricaria lycopodinea Cuatr., Feddes Repert.
Spec. Nov. Regni Veg. 56: 164. 1954. TYPE:
Peru, Dept. Huanuco, Prov. Dos de Mayo,
San Carlos Mines, 6 mi W of Huallanca, ca.
9000 ft, J. F. Macbride 2471 (holotype, F, F
neg. 40750; isotypes, GH, us).
Shrubs to 80 cm tall, much-branched; stems
flattened, 4-6 mm wide (including leaves). Leaves
triangular-oblong in lateral view, 5-8 mm long,
2-2.5 mm wide, dorsally carinate, apex subacute,
dorsal surface glabrous, olive-green, inner surface
lanate. Capitulescences of solitary, terminal heads.
DILLON & SAGASTEGUI: FLORA OF PERU. V.
47
Masculine capitula 4-5 mm high, 3-3.5 mm wide;
involucres campanulate; phyllaries 15-20, ovate
to elliptic or oblong, 3-3.5 mm long, 1.2-2 mm
wide, apex obtuse, lacerate; florets 16-22, the co-
rollas 4 mm long; pappus bristles clavellate, ca. 4
mm long; paleae ca. 6, oblong-linear, 4-5 mm long,
scarious, apex obtuse. Feminine capitula 67.5 mm
high, ca. 4 mm wide; involucres cylindrical; phyl-
laries ca. 12, the outer oval to ovate, ca. 4 mm
long, ca. 2 mm wide, the inner lanceolate, 6-6.5
mm long, 1-1.5 mm wide, apex acute; receptacles
epaleaceous; florets 1 1-13, the corollas ca. 5 mm
long; pappus bristles acute, ca. 5 mm long. Achenes
ca. 1.3 mm long, glabrous.
Loricaria lycopodinea is a Peruvian endemic
known from a few collections from north-central
Peru. It most closely resembles L. ferruginea but
differs in possessing narrower stems.
ANCASH. Yungay: Cordillera Blanca, Llanganuco,
Bathgate 16 (USM). LA LIBERT AD. Huamachuco: road
to Huamachuco, 8 km E of Quiruvilca, 4100 m, Hutch-
ison et al. 6139 (F, GH, MO, NY, uc); El Portachuelo Pass,
S of Cerro Huaylillas, 4000 m, West 8136 (GH.MO.UC).
JUNIN. Yauli: Oroya, Maisch s.n. (USM).
6. Loricaria macbridei Cuatr., Feddes Repert.
Spec. Nov. Regni Veg. 56: 164. 1954. TYPE:
Peru, Dept. Junin, Prov. Junin, San Jose, ca.
13,000 ft, /. F. Macbride & W. Featherstone
1093 (holotype, F, F neg. 40752; isotypes, GH,
us).
Shrubs to 60 cm tall, much-branched; stems
flattened, 5-7 mm wide (including leaves). Leaves
linear-oblong in lateral view, 5-8 mm long, 2-2.5
mm wide, dorsally carinate, apex subacute, curved
inward, dorsal surface glabrous, opaque, olive to
ochraceous, inner surface densely lanate. Capitu-
lescences of solitary, terminal heads. Masculine
capitula 6-7 mm high, 3-4 mm wide; involucres
campanulate; phyllaries 12-14, elliptic to oblong,
(4-)5-6 mm long, 1-2 mm wide, apex obtuse, lac-
erate; florets 10-24, sterile, the corollas ca. 4 mm
long; pappus bristles clavellate, ca. 4 mm long.
Feminine capitula 5-6 mm high, 1 .5-2.5 mm wide;
involucres cylindrical; phyllaries 1 3-22, elliptic to
oblong-linear, (3-)5-6 mm long, 1-2 mm wide,
apex subacute; florets 20-26, the corollas 4-4.5
mm long; pappus bristles acute, ca. 4 mm long;
paleae ca. 12, linear, ca. 6 mm long, 0.6-1 mm
wide, apex acute, lacerate. Achenes ca. 1.5 mm
long, glabrous.
Loricaria macbridei is a Peruvian endemic
known from three collections. This is the only spe-
cies that possesses paleas within the feminine ca-
pitula, but lacks them in the masculine capitula.
CUZCO. Paucartambo: Dist. Marcachea, Escalerayoc,
3700-4200 m, Vargas 11181 (F, uc). NO EXACT LO-
CALITY. Soukup s.n. (F).
7. Loricaria thuyoides (Lam.) Schultz-Bip., Bon-
plandia 8: 258. 1860.
Conyza thuyoides Lam., Encycl. 2: 90. 1786. TYPE:
Peru, no exact locality, /. deJussieu 42 (holotype,
p, not seen, F neg. 37750).
Molina incana Ruiz & Pavon, Syst. Veget. 211. 1794.
TYPE: Peru, Dept. Huanuco, Prov. Pachitea, Po-
zuzo, H. Ruiz & J. Pavon 28/18 (holotype, MA,
not seen; isotype, F).
Shrubs to 1 m tall, much-branched; stems slightly
to strongly flattened, 4-10 mm wide (including
leaves). Leaves linear in lateral view, 5-10 mm
long, 1-1.5 mm wide, dorsally carinate, apex sub-
acute to obtuse, dorsal surface glabrous, green to
gray-green, inner surface lanate. Capitulescences
of solitary, axillary heads. Masculine capitula 4-
5 mm high, 2-2.5 mm wide; involucres campan-
ulate; phyllaries 7-10, oblong to oblanceolate, 3-
4 mm long, 1-1.2 mm wide, apex obtuse, lacerate;
receptacles epaleaceous; florets 5 or 12-14, the
corollas ca. 3 mm long; pappus bristles clavellate,
ca. 3 mm long. Feminine capitula 4-5.5 mm high,
1-1.5 mm wide; involucres cylindrical; phyllaries
1 1-13, oblong, 4-5 mm long, ca. 1 mm wide, apex
obtuse, lacerate; receptacles epaleaceous; florets 5
or 1 0-1 2, the corollas 4-4.5 mm long; pappus bris-
tles acute, ca. 5 mm long. Achenes 1-1.5 mm long,
glabrous.
Loricaria thuyoides is widely distributed from
Colombia to Bolivia and in Peru ranges from
Amazonas to Cuzco, 3000-4500 m. This species
possesses considerable morphological variability,
especially in the width of the foliaceous stems and
the number of florets per head. Several varieties
have been recognized, including L. thuyoides var.
thyrsoidea Cuatr., a taxon here treated as a distinct
species (Dillon & Sagastegui, 1986). Two mor-
phological varieties are recognized.
48
FIELDIANA: BOTANY
Key to Varieties of Loricaria thuyoides
1 . Foliaceous stems strongly flattened, narrowly rhomboid in section; masculine and feminine capitula
with 10-14 florets var. thuyoides
1 . Foliaceous stems slightly compressed, subterete in section; masculine and feminine capitula with ca.
5 florets . . var. stuebelii
7a. Loricaria thuyoides var. thuyoides
Foliaceous stems strongly flattened, narrowly
rhomboid in section, 4-10 mm wide (including
leaves). Leaves 5-10 mm long, 1-1.5 mm wide,
apex obtuse to subobtuse. Masculine capitula 4
5 mm high, 2-2.5 mm wide; florets 12-14. Fem-
inine capitula 5-5.5 mm high, 1-1.5 mm wide;
florets 10-12. Chromosome number: n = ca. 14
(Turner et al., 1967).
Variety thuyoides has a wide distribution
throughout the Andean Cordillera and is distinc-
tive with its strongly flattened stems and axillary
capitula with 10-14 florets.
Masculine capitula ca. 4 mm high, ca. 2 mm wide;
florets ca. 5. Feminine capitula ca. 4 mm high, ca.
1.5 mm wide; florets ca. 5.
Variety stuebelii is readily distinguished from
the typical variety by possessing heads with only
5 florets. This variety may, upon further study,
prove to be worthy of specific status. It most close-
ly resembles L. pauciflora Cuatr. of Ecuador, a
species with only four (4) female florets per head.
LA LIBERT AD. Bolivar: Las Quinuas, 3350 m, Lopez
6 Sagdstegui 3345 (HUT, MO). HUANUCO. Huanuco:
Pachitea, Tambo de Vaca, 13,000 ft, Macbride 4385 (F,
GH, US).
AMAZON AS. Bagua: Cordillera Colan NE of La Peca,
10,400 ft, Barbour 3546 (F, MO). Chachapoyas: Cerros
de Calla-Calla, between Leimebamba-Balsas road pass
and the "camino de herradura," 3500-3750 m, Wurdack
1189 (F, GH, NY, uc, us); ca. 30 km SW of Leimebamba
on road to Balsas, ca. 3540 m, Dillon & Turner 1738A
(F). AYACHUCO. Huamanga: above Sotorabamba,
Weberbauer 5460 (F, GH). CUZCO. Quispicanchis: Am-
patuni, 4400-4500 m, Vargas 12157 (us). Urubamba:
Pumawanca, 3900-4350 m, Vargas 13701 (us). LA LIB-
ERT AD. Bolivar: Nevado de Cajmarquilla, Ferreyra 1291
(F, MO, USM); Ferreyra 1293 (MO, USM). HUANUCO.
Huanuco: Sariapampa, 3600 m, Woytkowski 34305 (F,
HUT, MO, uc, us, USM). Prov. Unknown: Shishmay, ca.
3000 m, Woytkowski s.n. (F). JUNIN. Yauli: cerca Ti-
clio, ferrocarril Lima- La Oroyo, Esposto s.n. (F, USM).
PASCO. Oxapampa: Cordillera Yanachaga, 3 1 50-3300
m, Smith 7701 (F, MO); between Santa Barbara and Lan-
turachi, 3000 m, Smith 8196 (F, MO). SAN MARTIN.
Mariscal Caceres: Puerto del Monte, 3500 m, Young
1799(p).
7b. Loricaria thuyoides var. stuebelii (Hieron.)
Cuatr.
Loricaria stuebelii Hieron., Bot. Jahrb. Syst. 21: 346.
1896. TYPE: Peru, "prope Pishenununa, 3200
m, et prope Calle-Calle, 3400-3600 m, inter Pas-
casmayo et Moyobamba," A. Stuebel 53 (holo-
type, B, probably destroyed, F neg. 15139).
Foliaceous stems slightly flattened, subterete in
section, 5-8 mm wide (including leaves). Leaves
7-10 mm long, ca. 1 mm wide, apex subacute.
8. Loricaria thyrsoidea (Cuatr.) Dillon & Sagast.,
Phytologia 59: 230. 1986.
Loricaria thuyoides var. thyrsoidea Cuatr., Feddes Re-
pert. Spec. Nov. Regni Veg. 56: 170. 1954. TYPE:
Peru, Dept. Lima, Prov. Canta, near Antaicocha,
Cerro Colorado, E of Canta, 4000-4100 m, 20
Jun 1925, F. W. Pennell 14685 (holotype, F; iso-
types, GH, NY, us).
Shrubs to 1 m tall, much-branched; primary
stems 7-8 cm in diameter, the foliaceous stems
flattened, 5-7 mm wide (including leaves). Leaves
triangular-oblong in lateral view, 5-7 mm long,
1.5-2 mm wide, dorsally carinate, apex obtuse,
dorsal surface glabrous, vernicose, greenish, inner
surface lanate. Capitulescences thyrsiform, ter-
minal and axillary. Masculine capitula 5-7 mm
high, 3-4 mm wide; involucres campanulate; phyl-
laries 20-22, obovate-oblong to oblanceolate, 4.5-
5.5 mm long, 1-2 mm wide, apex rounded, lac-
erate; receptacles epaleaceous; florets 16-20, the
corollas 4-^4.5 mm long; pappus bristles clavellate,
4.5 mm long. Feminine capitula 7-8 mm high, 2-
3 mm wide; involucres cylindrical; phyllaries 1 7-
21, oblong to oblong-linear, 4-5.5 mm long, 1-1.5
mm wide, apex obtuse, lacerate; receptacles epa-
leaceous; florets 12-15, the corollas ca. 4 mm long;
pappus bristles acute, ca. 5 mm long. Achenes 1-
1.5 mm long, glabrous.
DILLON & SAGASTEGUI: FLORA OF PERU. V.
49
Loricaria thyrsoidea is distributed from central
to southern Peru. The thyrsoid capitulescences of
this species are unique for the genus and quite
unlike those of all other members. In the Depart-
ment of Lima, it is known by the common name
of matara.
CUZCO. Quispicanchis: Marcapata, 10,000 ft, Staf-
ford 988 (F). JUNIN. Yauli: Tucto, cerca a Morococha,
4800-4900 m, Weberling 5907 (USM). LIMA. Canta:
cerca Canta, Sanchez 043 (F, USM). Huarochiri: Dist. San
Mateo, Rio Blanco, ca. 1 5,000 ft, Macbride 2985 (F, GH);
Sounders 388 (GH, NY). Prov. Unknown: Auquimarca,
5000 m, Geraldo 3274 (F).
XIII. Lucilia
Lucilia Cass., Bull. Soc. Philom. Paris 1817: 32.
1817. TYPE: Serratula acutifolia Poir. = Lu-
cilia acutifolia (Poir.) Cass.
Perennial herbs, erect or ascending, less often
cespitose, generally lanate or tomentose. Leaves
alternate and imbricate or rosulate, the margins
entire. Capitulescences glomerulate, few-headed
or of solitary, terminal heads. Capitula disciform;
involucres cylindrical to ovoid; phyllaries 4-6-se-
riate, imbricate, scarious, hyaline at margins, the
inner gradually longer; receptacle plane, naked;
pistillate florets 10-155, multiseriate, the corollas
filiform; hermaphroditic florets 3-23, the corollas
narrowly tubular, 5-lobed, the anther bases sag-
ittate, caudate, the terminal appendages ovate, the
style branches rounded, dorsal surfaces puberu-
lent. Achenes obovoid to turbinate, sericeo-pu-
bescent; pappus bristles scabrid, barbellate, the
bases fused, deciduous together as a unit. Chro-
mosome number: n = 14 (Freire, 1986b).
A genus of perhaps 12 species distributed
throughout South America, but with a center of
diversity in Argentina, southeast Brazil, Uruguay
and Paraguay; two are known from Peru.
There has been considerable difference of op-
pinion as to the generic limits between Lucilia and
Belloa (see discussion under Belloa). The gross
appearance of some members of each genus is quite
similar and especially those which occupy high-
elevation Andean habitats. Cassini's original de-
scription of Lucilia described sericeous achenes;
however, Freire (1986, 1987) expanded the genus
and transferred several species possessing achenes
with biseriate, multicellular, glandular trichomes
to the genus. Sagastegui and Dillon (1985) re-
moved from Lucilia all members with glandular
achenes (e.g., L. longifolid) and placed them in
Belloa. In the present treatment, only those taxa
with consistently sericeous achenes are included
in Lucilia.
The monotypic genus Novenia was established
by Freire (1986a) to accommodate one species
previously placed in Lucilia as L. tunariense
(Kuntze) K. Schumann (= N. acaulis).
References
CABRERA, A. L. 1978. Compositae. Flora de la Prov-
incia de Jujuy. Colecc. Cient., Inst. Natl. Teen. Agro-
pec., 13: 288-293.
FREIRE, S. E. 1 986a. Novenia: Nuevo genero de Inuleae
(Compositae). Bol. Soc. Argent. Bot., 24: 295-304.
. 1986b. Numeros cromosomicos en el genero
Lucilia (Compositae, Inuleae). Bol. Soc. Argent. Bot.,
24:411-413.
1986c. Revision del genero Lucilia (Compos-
itae, Inuleae). Darwiniana, 27: 431-490.
SAGASTEGUI-ALVA, A., AND M. O. DILLON. 1985. New
species and combinations in Belloa (Inuleae-Astera-
ceae). Phytologia, 58: 392-400.
ZARDINI, E. 1987. A new combination in Lucilia (Com-
positae-Inuleae). Ann. Missouri Bot. Gard., 74: 431.
Key to Species of Lucilia
1. Leaves spatulate, plane to falcate, 10-20 mm long, 1-2 mm wide L. kunthiana
1. Leaves linear, pleated or canaliculate, 5-15 mm long, 0.8-1 mm wide L. conoidea
1. Lucilia conoidea Wedd., Chlor. And. 1: 154.
1856. TYPE: Peru, Dept. Cuzco, "cordilleres
du departement de Cuzco," C. Gay s.n. (ho-
lotype, P, not seen, fragment, F). Figure 14, A-
G.
Cespitose, perennial herbs; stems much-
branched, decumbent. Leaves imbricate, sessile;
blade linear to spatulate, 5-1 5 mm long, 0.8-1 mm
wide, marcescent, the base slightly expanded and
clasping stem, pleated or canaliculate, apex obtuse,
mucronate, lower surface arachnoid-tomentose,
glabrescent, upper surface densely silvery-tomen-
tose. Capitulescences of solitary heads, terminal.
50
FIELDIANA: BOTANY
FIG. 14. Lucilia conoidea. A, habit; B, leaf, upper surface; C, leaf enlargement, lower surface; D, capitulum; E,
outer phyllary; F, inner phyllary; G, hermaphroditic floret. (Drawn from Dillon et al. 1082, HUT). Lucilia kunthiana.
H, leaf, lower surface. (Drawn from Dillon & Turner 1392, HUT.)
Capitula 7-8 mm high, ca. 4 mm wide; phyllaries
3540, 5-seriate, the outer ovate, 3-4 mm long,
1.5-2 mm wide, concave, dorsally lanuginous, apex
obtuse, the inner linear, ca. 6 mm wide, 0.8-1 mm
wide; pistillate florets 40-50, the corolla 4-5 mm
long; hermaphroditic florets 5-6, the corollas 4-5
mm long. Achenes obovoid, ca. 1 mm long, pilose,
brown; pappus bristles ca. 6 mm long.
Lucilia conoidea is known from three collections
in southern Peru and northern Bolivia. It ap-
proaches L. kunthiana in general appearance but
is distinctive in having leaves that are pubescent
on the upper surface and glabrescent below, the
reverse of the usual condition in Lucilia. Although
Lucilia conoidea was reduced to synonymy of L.
kunthiana (as L. pusilla) by Freire (1986b), it is
here accepted as distinct and readily recognizable.
PUNO. Carabaya: ca. 20 km SE of Macusani,
ca. 4500 m, Dillon et al. 1082 (F, HUT, LP, MO, NY,
USM); Lago Titicaca, 4000 m, Monheim Ml 57 (NY).
2. Lucilia kunthiana (DC.) Zardini, Ann. Mis-
souri Hot. Gard. 74: 431. 1987. Figure 14, H.
Conyza kunthiana DC., Prodr. 5: 379. 1836. (nomen
novum for Conyza pusilla Kunth).
DILLON & SAGASTEGUI: FLORA OF PERU. V.
51
Gnaphalium kunthianum (DC.) Kuntze, Revis. Gen.
PL, 3: 152. 1898.
Lucilia pusilla (Kunth) Hieron., Bot. Jahrb. Syst., 29.
29. 1900.
Conyza pusilla Kunth, Nov. Gen. Sp. (fol.) 4: 54.
1820. TYPE: Ecuador, Prov. Pichincha, "crescit
regione alpina Regni Quitensis," A. Humboldt &
A. Bonpland s.n. (holotype, p, not seen, IDC Mi-
crofiche 6209. 95: II. 3, F neg. 37649). Non Hout-
tuyn(1779).
Lucilia lehmannii Hieron., Bot. Jahrb. Syst. 19: 51.
1895. TYPE: Ecuador, Prov. Azuay, Paramo del
Cajas, 3600-4200 m, F. C. Lehmann 4649 (ho-
lotype, B, probably destroyed, F neg. 15126).
Cespitose perennial herbs; stems much-
branched, decumbent, densely foliaceous. Leaves
imbricate, sessile; blade linear-spatulate to spat-
ulate, 10-20 mm long, 1-2 mm wide, plane to
falcate, marcescent, the base attenuate and clasp-
ing stem, apex rounded, both surfaces tomentose.
Capitulescences of solitary heads, terminal. Ca-
pitula 7-8 mm high, ca. 5 mm wide; phyllaries ca.
35, 5-6-seriate, brownish black to stramineous,
the outer ovate, ca. 3 mm long, ca. 2 mm wide,
concave, dorsally glabrescent, apex obtuse, the in-
ner linear, 7-8 mm wide, ca. 1 mm wide, glabrous,
apex acute; pistillate florets ca. 50, the corolla fi-
liform, 5-6 mm long; hermaphroditic florets ca.
8, the corolla narrowly tubular, 4-5 mm long.
Achenes obovoid, ca. 1 mm long, pilose, brown;
pappus bristles 6-7 mm long.
Lucilia kunthiana is distributed throughout high-
elevation Andean habitats from Ecuador to Bo-
livia, 3000-4800 m. In Peru, it has a wide range
and is a relatively common element in jalca and
puna formations, 3000-4600 m.
ANCASH. Bolognesi: Quebrada Cancha-cancha, cer-
ca a Colca, 14,300 ft, Roberts 69 (USM). Carhuaz: Huas-
caran National Park, Quebrada Honda, between Vinoy-
apampa and Portachuelo Honda, 4200-4400 m, Smith
et al. 11669 (F, MO). Huari: Huascaran National Park,
Quebrada Pucaraju, 3900-4200 m, Smith et al. 12681.
Huaylas: Huascaran National Park, Paron Valley, E of
lake, 4200 m, Smith 11483 (F, MO). Recuay: Huascaran
National Park, Rio Pachacoto drainage, moraine below
Nevado Pasto Ruri, 4700 m, Smith 11426 (F.MO). APU-
RIMAC. No Exact Locality: Rauh & Hirsch P1426 (NY
p.p.). AYACUCHO. Parinacochas: 57 km W of Cotaruse,
ca. 1 1 1 km E of Puquio, 4100 m, Gentry et al. 23300
(F, MO, USM). CAJAMARCA. Cajabamba: Cajabamba-
Luchubamba, 3800 m, Sagdstegui et al. 11163 (F, HUT,
MO). CUZCO. Cuzco: Cordillera Veronica (Vilcanota),
4100 m, Rauh & Hirsch P999 (NY). Quispicanchis: La-
bramarca, 3800 m, Rauh & Hirsch PI 222 (NY); Auzan-
gate, 4600 m, Rauh & Hirsch P7 259 (NY p.p.). Urubamba:
Machupicchu, 4430 m, Peyton & Peyton 102 (MO); Ol-
lantaytambo, 4285 m, Peyton & Peyton 1136 (F, MO).
HUANCAVELICA. Huancavelica: San Jose de Aco-
bambilla, 4000 m, Cerrate & Marshall 3811 (USM).
HUANUCO. Huanuco: 6 mi S of Mito, 10,500 ft, Mac-
bride & Featherstone 1861 (F). JUNIN. Jauja: Mt. La
Juntay, near Huancayo, ca. 4700 m, Killip & Smith 22109
(F, NY). Junin: ca. 6 km N of La Cima, ca. 56 km N of
La Oroya, 10 km S of Junin, 4100 m, Dillon & Turner
1392 (F, HUT, MO, TEX, USM); Atocsayco, 4166 m, Johns
81-85 (F, MICH, USM). Yauli: Km 125 E of Casapalca,
14,250 ft, Davidson & Jones 9005 (us); ca. 17 km E of
La Oroya on road to Tarma, 3900-4000 m, Dillon &
Turner 1466 (F, MO, USM); 1 5 km SSW of La Oroyo, 3850
m, Iltis & Ugent 9 (uc, us); Laguna Pomacocha, 4200
m, Ochoa 606 (GH); near Atocsayco, ca. 4200 m, Pearsall
888 (F). PASCO. Pasco: Cerro de Pasco, ca. 14,000 ft,
Macbride & Featherstone 2400 (F). PUNO. Melgar: be-
tween Santa Rosa and Ayaviri, ca. 3500 m, Dillon et al.
1069 (F, LP, USM); Chuquibambilla, 3900-4000 m, Pen-
nell 13371 (F). Puno: Cerros de Puno, 3000-3900 m,
Tovar s.n. (USM). NO EXACT LOCALITY. Ellenberg
626 (USM), Soukup 3779 (F).
XIV. Mniodes
Mniodes A. Gray ex Benth. & Hook., Gen. PI. 2:
301. 1876. TYPE: Mniodes andina (A. Gray)
A. Gray ex J. D. Hook. & A. B. Jackson.
AntennariaGaertn. sect. Mniodes A. Gray, Proc. Amer.
Acad. Arts 5: 138. 1861.
Subshrubs or perennial herbs, suffrutescent, ces-
pitose, cushion-shaped, dioecious; stems much-
branched, often tightly compacted. Leaves densely
imbricate, appressed to stems, sessile; blade 2-5
mm long, both surfaces tomentose to villose, often
glabrescent, the margins entire. Capitulescences of
solitary heads, terminal or subterminal. Capitula
discoid, homogamous, sessile; involucres cylin-
drical; phyllaries multiseriate, imbricate, sub-
equal, scarious, stramineous, rarely with white
petaloid apex; receptacles naked; masculine florets
with the corollas tubular, apically dilated, 5 -fid,
the anthers sagittate, caudate, the style branches
obtuse, the ovary sterile, the pappus bristles uni-
seriate, clavellate, rarely slender and acute; femi-
nine florets with the corollas filiform, 5-fid or trun-
cate, the style branches linear-oblong, dorsally
pubescent, apex subobtuse, the ovary fertile, the
pappus bristles uniseriate, apex slender and acute.
Achenes fusiform to ovoid, 4-angled, brown, usu-
ally pubescent with biseriate, multicellular, capi-
tate-glandular trichomes or glabrous.
52
FIELDIANA: BOTANY
Mniodes is a distinctive genus of 4 species known
only from the Peruvian and adjacent Chilean An-
des (2730-4900 m).
Reference
CUATRECASAS, J. 1954.
Andina, 1: 1-7.
El genero Mniodes. Folia Biol.
Key to Species of Mniodes
1. Leaves obtrullate or fan-shaped, 4-5 mm long, 3-5 mm wide, 3-nerved; capitulescences subterminal,
axillary M. pulvinulata
1. Leaves oblong to obovate or obovoid, 3-4 mm long, (1-) 1.5-3 mm wide, 1 -nerved; capitulescences
terminal 2
2. Leaves oblong to subobovate, 1-1.2(-1.7) mm wide, apex truncate to rounded; achenes glabrous
M. andina
2. Leaves obovate to obovoid, 1.2-2 mm wide, apex subacute to rounded; achenes glabrous or
glandular 3
3. Leaves obovate-oblong, apex subacute, often briefly mucronate; achenes glandular
M. aretioides
3. Leaves ovoid to elliptic, apex obtuse to rounded; achenes glabrous M. coarctata
1. Mniodes andina (A. Gray) A. Gray ex J. D.
Hook. & A. B. Jackson, Ind. Kew. 2: 250.
1893. Figure 15.
Antennaria andina A. Gray, Proc. Amer. Acad. Arts
5: 138. 1861. TYPE: Peru, Dept. Junin, Prov.
Yauli, Alpamarca, Wilkes s.n. (holotype, GH; iso-
types, MO, NY, us).
Suffrutescent subshrub, pulvinate-cespitose;
stems foliaceous, cylindrical, 3-5 mm in diameter
(including leaves). Leaves with blades oblong to
subobovate, 34 mm long, 1 .0-1 .2(-l .7) mm wide,
1 -nerved, apex rounded or rarely truncate, both
surfaces incano-villous. Capitulescences of soli-
tary heads, terminal. Masculine capitula 3-3.5 mm
high, ca. 1.5 mm wide; phyllaries 7-8, oblong-
linear, 3.2-3.5 mm long, 0.6-0.8 mm wide, sub-
equal, apex obtuse, lacerate; florets ca. 7, the co-
rollas ca. 2.5 mm long; pappus bristles ca. 3 mm
long, clavellate. Feminine capitula ca. 4 mm high,
ca. 1.5 mm wide; phyllaries 8-10, oblong-linear,
3-3.5 mm long, 0.8-1 mm wide, subequal, apex
obtuse, lacerate; florets 10-14, the corollas 1.8-
2.3 mm long; pappus bristles 2.5-3 mm long, acute.
Achenes ca. 1.6 mm long, glabrous.
Mniodes andina is distributed from central to
southern Peru. It most closely resembles M. coarc-
tata, but differs from that species in having leaves
widest at the apex. This species forms large round-
ed cushions of stems that are easily separated.
ANCASH. Bolognesi: Dist. Ticllos, paso de Chonta,
4400 m, Cerrate 2658 (USM); Cushish, punta entre Chi-
quian y la Pampa de Lampas, 4300 m, Cerrate 2713
(USM). Carhuaz: Huascaran National Park, Quebrada
Ishinca, 4700-4930 m, Smith & Buddensiek 11207 (F,
MO). Recuay: Huascaran National Park, Quebrada
Quenua Ragra, 4600-4700, Smith et al. 10659 (F, MO).
Yungay: Cordillera Blanca, Llanganuco Valley, 16,000-
16,500 ft, Bathgate 11 (F.USM); Huascaran National Park,
Llanganuco sector, Quebrada Demanda, 4750-4900 m,
Smith & Cautivo 10296 (F, MO, USM); Pisco Creek, Llan-
ganuco Valley, Tothil 162 (uc). AREQUIPA. Cailloma:
San Ignacio above Cailloma, 4500 m, Schmidt s.n. (F).
JUNIN. Huancayo: Cordillera Huaytapallana, 4800 m,
Rauh & Hirsch PI 781 (NY). Yauli: Anticona Pass, 4800-
4900 m, Dillon & Turner 1479 (F, USM); Morococha,
Isern 2201 (F). NO EXACT LOCALITY. Ellenberg 2 1 8 1
(USM).
2. Mniodes aretioides (Schultz-Bip.) Cuatr., Folia
Biol. Andina, 1: 3. 1954. Figure 15.
Baccharis aretioides Schultz-Bip., Bonplandia 4: 51.
1856. TYPE: Peru, Dept. Puno, Prov. Carabaya,
Agapata [Ayapata], W. Lechler 1823 (holotype,
B, probably destroyed; isotypes, F, GH, G-DEL, not
seen, F neg. 28502, fragment, F).
Merope aretioides (Schultz-Bip.) Wedd., Chlor. And.
1: 164. 1856.
Antennaria aretioides (Schultz-Bip.) A. Gray, Proc.
Amer. Acad. Arts 5:139. 1861.
Suffrutescent subshrubs, pulvinate-cespitose;
stems foliaceous, cylindrical, 2-3 cm long, 3-4 mm
in diameter (including leaves). Leaves with blades
obovate to oblong, ca. 3 mm long, 1 .2-2 mm wide,
1 -nerved, base cuneate, apex truncate to subacute,
both surfaces incano-villous, the abaxial surface
DILLON & SAGASTEGUI: FLORA OF PERU. V.
53
FIG. 1 5. Mniodes andina. A, habit; B, leaf with pubescence removed; C, masculine floret; D, enlargment of pappus
apex, (drawn from Ellenberg 2181, USM); E, leaf of M. aretioides with pubescence removed; F, leaf of M. coarctata
with pubescence removed; G, leaf of M. pulvinulata.
rapidly glabrescent. Capitulescences of solitary
heads, terminal. Masculine capitula 3-3.5 mm
high, 1-1.2 mm wide; phyllaries 8-12, oblong-
linear, 3-3.5 mm long, 0.5-0.8 mm wide, sub-
equal, apex obtuse, lacerate; florets ca. 3, the co-
rollas ca. 2.5 mm long; pappus bristles ca. 2.5 mm
long, apex slightly dilated, acute. Feminine capit-
ula ca. 3.5 mm high, ca. 1.5 mm wide; phyllaries
8-12, oblong-linear, 2.5-3 mm long, 0.5-1 mm
wide, subequal, apex obtuse, lacerate; florets 3-5,
the corollas 1.5 mm long; pappus bristles ca. 2
mm long, acute. Achenes ca. 1 mm long, glandular.
54
FIELDIANA: BOTANY
Mniodes aretioides appears to have a narrow
distribution at very high elevations (ca. 4500 m)
in southern Peru. It is characterized by its narrow,
readily separable, cylindrical stems that are leafy
for their entire length. Mniodes aretioides most
closely resembles M. andina but it is readily dis-
tinguished by its glandular achenes. It is also dis-
tinguished from both M. andina and M. coarctata
by its acute, slightly dilated pappus tips of the male
florets.
CUZCO. Quispicanchis: Auzangate, 4500 m, Rauh &
Hirsch PI 142 (NY). PUNO. Carabaya: ca. 20 km SE of
Macusani, ca. 4500 m, Dillon et al. 1083 (F, HUT, K, MO,
NY, USM).
3. Mniodes coarctata Cuatr., Folia Biol. Andina
1: 4. 1954. TYPE: Peru, Dept. Arequipa, Prov.
Arequipa, 52 mi ENE of Arequipa, 15,000 ft,
18 Apr 1952, C. B. Koford P. 25 (holotype,
UQ isotype, F). Figure 15.
Suffruticose perennial herbs, pulvinate-cespi-
tose; stems foliaceous, cylindrically compressed,
2-3 cm long, 3-5 mm in diameter (including
leaves). Leaves with blades ovate to elliptic, 3-3.5
mm long, ca. 2 mm wide, 1 -nerved, apex obtuse
to rounded, both surfaces densely villous. Capit-
ulescences of solitary heads, terminal. Masculine
ca pi tula 3-3.5 mm high, 1-1.5 mm wide; phyl-
laries 7-8, oblong, 3-3.2 mm long, 0.3-1 mm wide,
apex obtuse, lacerate; florets ca. 8, the corollas ca.
2.8 mm long; pappus bristles ca. 3 mm long, clav-
ellate. Feminine capitula ca. 3 mm high, ca. 1.5
mm wide, sessile; phyllaries 7-8, oblong, ca. 2.5
mm long, ca. 0.5-0.9 mm wide, apex obtuse to
rounded; florets ca. 9, the corollas ca. 1.8 mm long;
pappus bristles ca. 2 mm long, acute. Achenes ca.
1 mm long, glabrous.
Mniodes coarctata is only known from a few
collections from central and southern Peru. While
it has been reported from northern Chile, no ma-
terial from that region has been seen. It most close-
ly resembles M. aretioides in general habit but is
distinguished from that species by its ovate to el-
liptic leaves with rounded to obtuse apices.
AREQUIPA. Arequipa: Paso del Condor, 4693 m,
lit is & Ugent I486 (GH, MO, NY, uc, USM); Volcan Pichu-
pichu, 4500 m, Rauh & Hirsch P622 (NY). La Union:
Cordillerana Rio Armas, 4300-4500 m, Vargas 19547
(uc). AYACHUCO. Parinachocas: Negro Mayo, Km 75-
80, Iltis & Ugent 504 (GH, MO). LIMA. Huarochiri: Ca-
prichosa, above Casapalca, 4750 m, Amstutz 114 (MO).
4. Mniodes pulvinulata Cuatr., Folia Biol. Andina
1: 5. 1954. TYPE: Peru, Dept. Huanuco,
Rumichaca, estacion 8 millas SW de Chasqui,
1 1,500 ft, 28 Sep 1922, /. F. Macbride & W.
Featherstone 2460 (holotype, F, F neg. 49244;
isotype, us). Figure 15.
Mniodes ferreyrae Cuatr., Folia Biol. Andina 1: 6.
1954. TYPE: Peru, Dept. Ancash, Prov. Bolog-
nesi, Conococha, 4000 m, 8 Aug 1949, R. Fer-
reyra 6231 (holotype, us; isotype, USM).
Suffrutescent herbs, pulvinate-cespitose; stems
4-8 cm long, 4-6 mm in diameter (including
leaves). Leaves with blades widely obtrullate or
fan-shaped, 3.54 mm long, (3-)3.5-4 mm wide,
3-nerved, base encircling the stems ca. half the
circumference, apex rounded to truncate, both sur-
faces densely villous distally. Capitulescences of
solitary heads, subterminal. Masculine capitula 5-
6 mm high, 2-2.5 mm wide; phyllaries ca. 14,
oblong, 3-3.5 mm long, 1-2 mm wide, subequal,
scarious, apex obtuse, revolute, petaloid, white;
florets 10-12, the corollas 3-3.5 mm long; pappus
bristles ca. 4 mm long, clavellate. Feminine ca-
pitula 6-7 mm high, 1.5-2.5 mm wide; phyllaries
14-20, oblong, 4-6 mm long, 1-2 mm wide, sub-
equal, apex obtuse, revolute, petaloid, white; flo-
rets 12-14, the corollas 4 mm long; pappus bristles
ca. 5 mm long, acute. Achenes ca. 1.5 mm long,
densely glandular.
Mniodes pulvinulata is widely distributed
throughout puna habitats from northern to south-
ern Peru. It is characterized by its 3-nerved, wide
obtrullate or fan-shaped leaves encircling the stems
for nearly half the circumference.
Cuatrecasas (1954) described M. ferreyrae as
distinct from M. pulvinulata and stated that it had
slightly narrower leaves that were less densely im-
bricate. After intensive study of considerable ad-
ditional material, the present authors can find no
basis to maintain this species. It appears that slight
variations in leaf size and degree of compactness
are under environmental control and a full range
of leaf sizes and orientations is present in an in-
dividual gathering (Sagdstegui & Dillon 12342)
collected from the type locality.
ANCASH. Bolognesi: Cordillera Raura, 4900 m, Rauh
& Hirsch PI 899 (NY). Carhuaz: Cordillera Blanca, above
Vicos on trail to Llacshacocha, at Carhua Catac, 4000
m, Hutchison & Wright 4401 (F, NY, uc); Huascaran Na-
tional Park, Quebrada Ishinca, 4730-4930 m, Smith et
al. 9456 (F, MO, USM), Smith & Buddensiek 11210 (F.MO).
Huaylas: Huascaran National Park, Quebrada Santa Cruz,
DILLON & SAGASTEGUI: FLORA OF PERU. V.
55
4300-4800 m, Smith et al. 9313 (F, MO, USM). Recuay:
Conococha, 4 1 00 m, Sagdstegui & Dillon 12342 (F, HUT);
Huascaran National Park, Quebrada Queshque, 4550-
4600 m, Smith et al. 11884 (F, MO, USM), 11858 (F, MO,
USM). CAJAMARCA. Cajabamba: Cajabamba-Luchu-
bamba, 3800 m, Sagdstegui et al. 11154 (F, HUT, K, MO,
us). Cajamarca: desvio al Arboretum Negritos, Km 35
de la carretera Cajamarca-Hualgayoc, 4050 m, Sanchez
el al. 2558 (CPUN, F). Celendin: Pumarrume, 2730 m,
Mostacero et al. 1098 (HUT). Otusco: Julcan, 3000 m,
sin coll. (HUT #16935). PUNO. Huancane: Miajachi, 3900
m, Aguilars.n. (F.USM). LA LIBERTAD. Huamachuco:
ca. 25 km E of Quirivilca, 13,600 ft, Duncan et al. 2643
(F, MO, uc); Jalca de Huaguil, 3950 m, Lopez 2731 (HUT),
1424 (USM); Pallar-Huaguil, carretera a Tayabamba, 3300
m, Lopez & Sagdstegui 8139 (GH, HUT, MO). Santiago de
Chuco: Huillilas N of Cachicadan, 4000 m, Stork & Hor-
ton 10002 (F). LIMA. Canta: La Viuda, arriba de Cul-
Ihuay, 4250 m, Meza 200 (F, MO, USM).
Brazilian genus Chionolaena for possible generi
relationships.
References
ANDERBERG, A. A. 1989. Phylogeny and reclassifica
tion of the tribe Inuleae (Asteraceae). Can. J. Bot., 61
2277-2296.
FREIRE, S. E. 1 986a. Novenia: Nuevo genero de Inulea
(Compositae). Bol. Soc. Argent. Bot., 24: 295-304.
. 1986b. Numeros cromosomicos en el gener
Lucilia (Compositae, Inuleae). Bol. Soc. Argent. Bot
24: 411-413.
AND F. HELLWIG. 1990. A new combination i:
Novenia (Compositae: Inuleae). Taxon, 39: 124-12f
XV. Novenia
Novenia Freire, Bol. Soc. Argent. Bot. 24: 296.
1986. TYPE: Gnaphalium tunariense Kuntze
= Novenia acaulis (Benth.) Freire & Hellwig.
Acaulescent, cespitose, perennial herbs. Leaves
densely rosulate, sessile; blade narrowly linear, co-
riaceous, canaliculate, the base slightly expanded
and densely villous, apex attenuate, mucronate,
both surfaces glabrous distally. Capitulescences of
1-4 heads, glomerulate and sessile in the central
rosette, rarely of a solitary head. Capitula disci-
form, heterogamous; phyllaries subequal, ca. 4-
seriate, stramineous, scarious, glabrous, indurate;
pistillate florets 3-9, the corolla filiform, briefly
ligulate; hermaphroditic florets 3-6, the corollas
narrowly tubular, 5-lobed. Achenes obovoid to
oblong, sericeo-pubescent, brownish; pappus bris-
tles uniseriate, the bases fused.
A monotypic genus distributed in high-eleva-
tion habitats of Peru, Bolivia, and extreme north-
western Argentina. Freire (1986a) established
Novenia as distinct from Lucilia on the basis of
differences in habit, foliar trichomes, and chro-
mosome number. In fact, the characteristic n = 9
chromosome number was considered to be of such
significance that she named the genus for the num-
ber nine and placed the genus within Merxmuller's
broad "Lucilia group." The relationships of this
genus are not clear but it is noteworthy that An-
derberg (1989) has questioned its position even
within his Gnaphalieae. We accept this genus
within the Gnaphaliinae and would look to the
1. Novenia acaulis (Wedd. ex Benth.) Freire &
Hellwig, Taxon, 39: 125. 1990. Figure 16.
Dolichogyne acaulis Wedd. ex Benth., Gen. PI. 2: 258
1873. TYPE: Bolivia, "Prov. a Larecaja, vicinni
Sorata," Mandon 221 (BM, lectotype, not seen).
Lepidophyllum acaule (Wedd. ex Benth.) Benth. i
Hook, ex J. D. Hook. & A. B. Jackson, Ind. Kev,
3: 58. 1894.
Gnaphalium tunariense Kuntze, Revis. Gen. PI. 2
155. 1898. TYPE: Bolivia, Dept. Cochabambz
Tunarigebirge, 4000-5000 m, Apr-May 189^
"besonders auf Torfmorren haufig," O. Kuntz
s.n. (NY, holotype, not seen).
Lucilia tunariensis (Kuntze) K. Schumann, Just. Boi
Jahresber. 28: 378. 1898.
Mniodes tunariensis (Kuntze) Hieron. ex Weberbauei
El Mundo Vegetal de los Andes Peruanos 731
1945.
Novenia tunariensis (Kuntze) Freire, Bol. Soc. Argen
Bot. 24: 295. 1986.
Acaulescent herbs, 1 .5-4 cm in diameter, form
ing cushions. Leaves with blades 1.5-2.5 cm long
0.8-1 mm wide. Capitula 9-10 mm high, 4-5 mr
wide; phyllaries ca. 18, the outer ovate to lance
ovate, 5-6 mm long, 1.5-2 mm wide, apex atten
uate, the inner linear, 6-7(-10) mm long, 0.8-1.
mm wide, apex acute; marginal florets (3-)7-8(
9), the corolla ca. 3.5 mm long; hermaphroditi
florets 3-4(-6), the corollas ca. 4 mm long. Achene
ca. 3 mm long; pappus bristles 4-5 mm long. Chro
mosome number: n = 9 (Freire, 1986b).
Novenia acaulis is distributed from northern Per
to Bolivia and into extreme northwestern Argen
tina, 3100-4600 m. Its acaulescent habit an
densely rosulate, apiculate, coriaceous leaves ar
distinctive within the Peruvian Inuleae.
56
FIELDIANA: BOTAISH
FIG. 16. Novenia acaulis. A, habit; B, leaf; C, capitulum; D, outer phyllary; E, center phyllary; F, inner phyllary;
G, pistillate floret; H, hermaphroditic floret; I, enlargement of corolla lobes and style branches of hermaphroditic
floret; J, achene. (Drawn from Lopez & Sagdstegui 8243, HUT.)
ANCASH. Carhuaz: Cordillera Blanca, above Viscos
[Viscas] on trail to Llaschacocha, at Carhua Catac, 4000
m, Hutchison & Wright 4398 (uc). Huari: Huascaran
National Park, 1 km below Manto Mina, 4300 m, Smith
& Buddensiek 11012 (F, MO). Recuay: Quebrada Huanca,
3950 m, Smith & Buddensiek 10971 (F, MO); Quebrada
Queshque, 4550-4600 m, Smith et al. 11887 (F). CA-
JAMARCA. Contumaza: Jalca de las Quinuas (Contu-
DILLON & SAGASTEGUI: FLORA OF PERU. V.
maza-Asuncion), 3900 m, Sagdstegui et al. 10089 (F,
HUT, MO). San Miguel: Taulis Alto, 3100 m, Sagdstegui
et al. 9545 (F, HUT). CUZCO. Canchis: La Raya, 4400-
4500 m, Pennell 13508 (F). HUANUCO. Dos de Mayo:
Lauricocha, Cardich s.n. (F, MO, USM). JUNIN. Junin:
Junin, Aguilar s.n. (F, MO, USM); Yaoli: near Atocsayco,
ca. 4200 m, Pearsall 825 (F). LA LIBERTAD. Pataz:
carretera Huancaspata-Tayabamba, 3700 m, Lopez &
Sagdstegui 8243 (HUT, MO). LIMA. Huarochiri: laguna
de Chumpicocha, 4600 m, Cerrate2001 (USM). PASCO.
Pasco: Huaron, ca. 14,000 ft, Macbride & Featherstone
1144 (F). PUNO. Carabaya: Macusani, 4300 m, Marin
2092 (F). NO EXACT LOCALITY. Ellenberg654 (USM),
679 (USM); Haenke 1928 (F); Weberbauer 5452 (F).
XVI. Pluchea
Pluchea Cass., Bull. Sci. Soc. Philom. Paris 1817:
31. 1817. TYPE: Conyza marilandica A. Mi-
chaux = Pluchea marylandica (A. Michaux)
Cass. 3
Perennial herbs or shrubs, often aromatic; stems
glabrous, pubescent, or glandular, sometimes
winged. Leaves alternate, simple, sessile, auricu-
late-clasping, or occasionally petiolate, often de-
current on stem; blade ovate to linear-lanceolate,
glabrous, pubescent, or glandular, the margins en-
tire or toothed. Capitulescences corymbose-panic-
ulate or elongate to flat-topped, cymes. Capitula
disciform, heterogamous; involucres ovoid or
campanulate to hemispheric; phyllaries 2-5-seri-
ate, imbricate, herbaceous to chartaceous, indu-
rate, or scarious; receptacles flat, glabrous; pistil-
late florets multiseriate, the corollas filiform,
slightly dilated distally, 3(-4)-lobed, purplish to
cream-white, the styles greatly divided, the
branches slender, exserted; disc florets hermaph-
roditic, the corollas tubular, slender, distally di-
lated, cylindrical or slightly campanulate, 5-lobed,
rose to rose-purple, the anther bases sagittate, cau-
date, the terminal appendages deltoid, rounded,
the styles bifid, the branches slender, separate or
united, apex rounded. Achenes cylindrical, prom-
inently 3-6-ribbed, strigose, glandular, or gla-
brous; pappus bristles barbellate, uniseriate, the
bases fused. Chromosome numbers: n = 10, 15.
Pluchea includes about 40 species occurring in
the warmer regions of both hemispheres, with four
species known from Peru. The generic limits have
been emended by Robinson and Cuatrecasas (1973)
to include several species previously placed in Tes-
saria (e.g., P. absinthioides and P. zamalloae).
References
ARIZA E., L. 1969. Notas sobre Compositae Argenti-
nas, I. La distribution geografica de Pluchea micro-
cephala Godfrey, con una clave de las especies con-
genericas. Kurtziana, 5: 297-302.
. 1979. Contribution al conocimiento del genero
Tessaria (Compositae), I. Considerations sobre los
generos Tessaria y Pluchea. Kurtziana, 12-13: 47-49.
GODFREY, R. K. 1962. Pluchea, section Stylimnus, in
North America. J. Elisha Mitchell Sci. Soc., 68: 238-
271, pis. 20-23.
ROBINSON, H., AND J. CUATRECASAS. 1 973. The generic
limits of Pluchea and Tessaria (Inuleae, Asteraceae).
Phytologia, 27: 277-285.
SUNDBERG, S., AND M. DILLON. 1986. Chromosome
number reports XCI. Taxon, 35: 404-410.
Key to Species of Pluchea
1 . Perennial herbs usually 1 m or less tall; stems winged; leaves linear-lanceolate to obovate, generally
greater than 6 cm long P. microcephala
1. Shrubs usually over 1 m tall; stems unwinged; leaves oblanceolate, elliptic, or ovate, l-5(-8) cm long
2
2. Leaves 5-8 cm long, usually oblanceolate, densely cano-tomentulose P. absinthioides
2. Leaves 1-3.7 cm long, ovate to oval or elliptic (rarely oblanceolate), tomentose to strigillose . 3
3. Spreading shrubs to 3.5 m tall; leaves oval to ovate, 8-25 mm wide; distributed in arid coast
habitats from northern Peru to northern Chile (usually below 1000 m) P. chingoyo
3. Erect shrubs, usually less than 2 m tall; leaves elliptic or rarely oblanceolate, 7-15 mm wide;
distributed in interior southern Peru, above 1000 m P. zamalloae
1 . Pluchea absinthioides (Hook. & Arn.) H. Rob-
inson & Cuatr., Phytologia 27: 284. 1973.
3 For additional generic synonymy, see Godfrey ( 1 962).
Only the above name has been used for Peruvian ma-
terial.
Baccharis absinthioides Hook. & Arn., Bot. Beechey
Voy. p. 57. 1830. TYPE: Chile, Valparaiso, T.
Bridges 55 (holotype, GL, not seen).
Tessaria absinthioides (Hook. & Arn.) DC., Prodr. 5:
457. 1836.
58
FIELDIANA: BOTANY
Shrubs or subshrubs to 2 m tall; stems erect,
densely tomentulose. Leaves oblanceolate, 5-8 cm
long, 5-12 mm wide, base attenuate to a pseudo-
petiole, apex acute to subobtuse, both surfaces
densely cano-tomentose, the margins dentate or
rarely entire. Capitulescences cymose-corymbose,
terminal. Capitula 6-8 mm high, 4-5 mm wide;
involucres campanulate; phyllaries 3-4-seriate,
scarious, ciliate, the outer ovate, 2.5-3 mm long,
1.2-1.5 mm wide, apex obtuse, the inner lanceo-
late to linear, 5-6 mm long, 0.6-1 mm wide, apex
obtuse to attenuate; pistillate florets ca. 100, the
corollas 5-6 mm long, lilac; hermaphroditic florets
6-14, the corollas ca. 6 mm long, the lobes 1-1.5
mm long. Achenes ca. 0.6 mm long, glabrous; pap-
pus bristles 4.5-5 mm long.
Pluchea absinthioides occurs frequently in sandy
soils from southern Peru, Bolivia, Chile, Uruguay,
and Argentina. In Peru, it is known from two col-
lections in the Department of Arequipa (2 1 00-
2300 m) where it is known by the following com-
mon names: brea, pdjaro bobo, suncho negro.
AREQUIPA. Arequipa: southern slopes of Chachani,
2287 m, Hinkley 33 (F, GH, us); Tiabaya, 2100-2200 m,
Pennell 13084 (GH).
2. Pluchea chingoyo (Kunth) DC., Prodr. 5: 452.
1836. Figure 17.
Conyza chingoyo Kunth, Nov. Gen. Sp. (fol.) 4: 59.
1820. TYPE: Peru, Dept. Ancash, Prov. Santa,
A. Bonpland 3758 (holotype, P, not seen, IDC
Microfiche 6209. 96: 1. 5, Fneg. 37652, fragment,
Shrubs to 3.5 m tall; stems densely branched,
spreading, glandular-pubescent. Leaves sessile or
with petiolates to 1 cm long; blade ovate to oval,
l-3(-3.5) cm long, 8-25 mm wide, upper surfaces
glabrous to sparsely tomentose, lower surfaces gla-
brous to densely tomentose, both surfaces glan-
dular-punctate, base cuneate to truncate or cor-
date, apex rounded to obtuse, the margins entire
to serrate. Capitulescences densely corymbose-pa-
niculate, terminal and axillary; peduncles 1-5 mm
long, puberulent. Capitula 4-5 mm high, 2.5-3
mm wide; involucres campanulate; phyllaries 4-
5-seriate, the outer ovate to lanceolate, 1-1.5 mm
long, 0.5-0.7 mm wide, glandular, resinous, apex
acute, ciliolate, the inner linear to lanceolate, 3.5-
4.5 mm long, 0.3-0.5 mm wide, apex attenuate,
glandular, resinous; pistillate florets ca. 100, the
FIG. 17. Flowering branch of Pluchea chingoyo.
(Drawn from Ferrerya 6298, F.)
corollas 2.5-3 mm long, 3 -fid, the style branches
linear, ca. 0.5 mm long; hermaphroditic florets ca.
1 5, the corollas 4.2-4.7 mm long, the limb slightly
expanded, 5-lobed, the lobes ca. 0.5 mm long,
deltate, the style branches ca. 0.5 mm long, stri-
gillose below the point of bifurcation. Achenes ca.
0.6 mm long, hirtellous; pappus bristles 1 .5-3 mm
long. Chromosome number: n = 1 (Sundberg &
Dillon, 1986).
Pluchea chingoyo occurs in xeric, coastal habi-
tats (10-800 m) from northern Peru to northern
Chile; occasional individuals are found at higher
elevations further inland. This species is quite dis-
tinctive in having densely branched, shrubby habit
and small ovate leaves. In Peru, populations in
the north (Department of Piura) have considerably
more pubescence than those in the south (De-
partments of lea and Arequipa), possibly due to
environmental effects.
It is known by the following common names:
parrano, tonuz.
ANCASH. Bolognesi: abajo de Raquis, ruta Pativilca-
Huaraz, 1900 m, Sagdstegui et al. 12298 (F, HUT). Cas-
ma: ca. 48 km N of Pativilca on Pan American Hwy.,
ca. 10 m, Dillon & Whalen 4006 (F.USM); Yautan, 2000
ft, Macbride & Featherstone 2562 (GH, us). Bolognesi:
DILLON & SAGASTEGUI: FLORA OF PERU. V.
59
abajo de Raquia, ruta a Pativilca-Huaraz, 1900 m, Sa-
gdstegui el al. 12298 (F, HUT). Santa: Samanco, cerca a
Chimbote, 10-15 m, Ferreyra 18529 (F, MO, USM). AR-
EQUIPA. Camana: 25 km NW of Camana, I It is et al.
1557 (MO, us). Caraveli: Lomas de Jahuay, entre Chala
y Nazca, 300-400 m, Ferreyra 14010 (MO, USM). Islay:
14 km S of Mollendo, Hutchison & Wright 7138 (F, us,
USM); 7 km de Matarani, 500 m, Lopez 64 (MO); Mol-
lendo, 600 ft, Stafford 902 (F). ICA. lea: laguna de La
Victoria, 500 m, Ferreyra 565 (F, MO, USM); Hda. Ocu-
caje, 300-400 m, Ferreyra 1381 (uc, us, USM); Laguna
de Orovilca, a 6-7 km de lea, 400 m, Cerrate 897 (MO,
USM), Ferrerya 2496 (MO, NY, us), 6277 (MO, us, USM);
Huaga, 300 m, Ferreyra 8205 (MO, USM); lea, Ridoutt
s.n. (MO, USM mos); Ica-Ocucaje, 300 m, Weberbauer
7194 (GH, us). Nazca: ca. 85 km S of Nazca, 90 m,
Ellenberg 4143 (us). Pisco: Chincha, valley of Rio San
Pisco, 900-1000 m, Weberbauer 5369 (F, GH, us). JUN-
IN. Tarma: cerca Fundo Puguio, 800-900 m, Ferreyra
17702 (MO, USM). LA LIBERT AD. Pascasmayo: 7 km E
of San Pedro de Lloc, ca. 120 m, Dillon & Whalen 4020
(F, MO, NY, TEX, USM). Trujillo: between Viru and Chim-
bote, 485 km N of Lima on Pan-Amer Hwy, Rahn 323
(MO). LAMBAYEQUE. Chiclayo: ca. 2 km E of Zana,
100-1 50 m, Dillon et al. 4095 (F, MO, TEX, USM); 28 km
S of Lambayeque, 30 m, Ellenberg 3749 (us); desvio a
Oyotun, cerca a Bebedero, 200 m, Sagdstegui et al. 12352
(F, HUT, K, MO); Chiclayo-Patapo, 50 m, West 3580 (GH,
uc). Ferreflafe: Pitipo (tres Tomas), 80 m, Llatas & Laos
822 (F). LIMA. Canete: Chilca, Asplund 13849 (us).
Huarochiri: Cocachacra, Km 60 entre Lima y La Oroyo,
Ridoutt s.n. (MO, USM 126399). Lima: Chosica, 40 km de
Lima, 800 m, Ferreyra 546 (MO, USM); La Cantuta, cerca
Chosica, Lima-Huancayo, 700-800 m, Ferreyra 6298 (F,
us, USM); ruinas de Cajamarquilla, 300-400 m, Ferreyra
2847 (MO, USM), 16886 (MO, USM); Ricardo Palma, 850
m, Riccio 4677 (us). Hda. Zarate, Soukup 3133 (us).
PIURA. Huancabamba: arriba de Palambla, 1 500-1 600
m, Cerrate & Tovar 10809 (MO, USM). Morropon: road
to Huancabamba, 3 km W of Serran, 1 2 km E of Ma-
lacasi, 200 m, Hutchison & Wright 6548 (F, GH, MO, NY,
uc, us). Talara: 25 mi E of Cabo Blanco, Cana Dulce,
Haught 200 (F, GH, NY). TACNA. Tacna: Boca del Rio,
50 m, Zegarra 3344 (us).
3. Pluchea microcephala Godfrey, Jour. Elisha
Mitchell Soc. 68: 270. 1952. TYPE: Argen-
tina, Prov. Formosa, Dept. Pirane, al N.O. a
6 km Pirane, 5 Dec 1945, T. Morel 518 (ho-
lotype, F, isotype, LIL, not seen). Figure 18.
Perennial herbs to 1.2 m tall; stems erect,
branched, glabrous to glandular-pubescent, leafy,
alate, the wings 1.5-5 mm wide. Leaves sessile;
blade narrowly obovate, elliptic, or linear-lance-
olate, 6-9(-l 1) cm long, l-3(-4) cm wide, upper
and lower surfaces glandular pubescent to gla-
brous, base attenuate, decurrent on stems, apex
acute to obtuse, the margins serrate. Capitules-
cences densely cymose-paniculate, 5 0-7 5 -headed;
peduncles glandular pubescent. Capitula (3-)4-5(-
6) mm high, 2-3.5(-4) mm wide; involucres nar-
rowly campanulate; phyllaries 4-5 -seriate, gla-
brous, the outer ovate, 1.5-2 mm long, 0.9-1.2
mm wide, submembranous, apex obtuse, the inner
oblong to linear-lanceolate, 3.5-4.5 mm long, 0.4-
0.7 mm wide, membranous, apex acute to atten-
uate; pistillate florets 25-50, the corollas ca. 44.3
mm long, 3-dentate, rose-purple; hermaphroditic
florets 3-5, the corollas 4.5-5.5 mm long, apex
dilated, 5-lobed, the lobes lanceolate, ca. 0.5 mm
long. Achenes 0.8-1 mm long, papillose to gla-
brous, 4-6-ribbed; pappus bristles, 44.5 mm long.
Chromosome number: n = 10 (Sundberg & Dillon,
1986).
Pluchea microcephala is known from eastern
Bolivia, northern Argentina, and northeastern Peru
(0-200 m). It is perhaps most closely related to P.
suaveolens of Argentina, which shares its winged
stems and leaves, but P. microcephala is distin-
guished by its smaller capitula with narrowly cam-
panulate involucres.
LA LIBERTAD. Pacasmayo: Rio Jequetepeque, Sa-
gdstegui et al. 8530 (F, HUT, MO); Chepen-Pacanguilla,
180 m, Sagdstegui & Diestra 11486 (F, HUT, K). LAM-
BAYEQUE. Chiclayo: ca. 2 km E of Zana, 100-150 m,
Dillon et al. 4094 (CPUN, F, GH, HUT, K, MO, NY, TEX, us,
USM); ca. 2.5 km E of Chiclayo, Hudson 965 (F, MO);
Chiclayo-Lambayeque, Km 764, Lopez 8108 (F), Sa-
gdstegui 8353 (F, HUT, MO); common in plains around
Chiclayo, Stork 11429 (F, uc) .
4. Pluchea zamalloae (Cabr.) H. Robinson &
Cuatr., Phytologia 27: 284. 1973.
Tessaria zamalloae Cabr., Notas Mus. La Plata, Bot.
19: 201. 1959. TYPE: Peru, Dept. Cuzco, Prov.
Urubamba, 3870 m, H. Zamalloa 1007 (holo-
type, LP, not seen).
Shrubs to 3 m; stems densely villous to gla-
brescent, striate. Leaves sessile or with petioles 1-
2 mm long; blade elliptic to oblanceolate, 1.5-3.7
cm long, 7-15 mm wide, upper surfaces sparsely
strigillose, glandular-punctate, resinous, lower
surfaces strigillose, glandular-puncate, resinous,
base attenuate, apex obtuse, the margins entire or
dentate with 6-8 pairs of teeth. Capitulescences
dense corymbose cymes, terminal; peduncles 3-5
mm long, puberulent. Capitula 4-5 mm high, 2.5-3
mm wide; involucres campanulate; phyllaries 4-
5-seriate, the outer ovate to oblong, 1-2 mm long,
ca. 0.6 mm wide, apex obtuse, the inner lanceolate
to oblong-lanceolate, 3.5^4 mm long, apex acute
60
FIELDIANA: BOTANY
FIG. 18. Pluchea microcephala. A, habit; B, capitulum; C, pistillate floret; D, hermaphroditic floret; E, achene
with pappus. (Drawn from Sagdstegui et al. 8530, HUT.)
to attenuate; pistillate florets ca. 20, the corollas
ca. 3 mm long, 3-4-fid, the style branches ca. 0.5
mm long; hermaphrodite florets ca. 4, the corollas
4-5 mm long, the limb slightly expanded, 5-lobed,
the teeth triangular, ca. 0.5 mm long, the style
branches ca. 0.5 mm long, strigillose below the
point of bifurcation. Achenes 0.6-0.8 mm long,
sericeous; pappus bristles 2.5-3.5 mm long.
DILLON & SAGASTEGUI: FLORA OF PERU. V.
61
Pluchea zamalloae is apparently a rare species
that occurs in dry sites along river valleys in south-
ern Peru (1850-3300 m). This species is most
closely related to P. chingoyo but is readily distin-
guished from the latter by its leaf-shape and eco-
logical preference.
APURIMAC. Abancay: cerca de Cunyac, 2300 m,
Ferreyra 9829 (F, MO, us, USM); 33 km SW of Abancay,
Hac. Casinchihua, 2300-2500 m, Iltis & Ugent 814 (wis,
us), 814a (wis); Quebrada Honda, Vargas 465 (us); E
side of Rio Pacachacas, 1850 m, West 3795 (MO, uc).
AYACUCHO. Huanta: Hda. Bado Grande, 2300 m,
Vargas 15670 (us). CUZCO. Ante: cerca de Yucay, 3300
m, Ferreyra 9875 (MO, USM).
XVII. Pterocaulon
Pterocaulon Elliott, Sketch Bot. S. Carol. 2: 323.
1823. TYPE: Conyza pycnostachya A. Mi-
chaux = Pterocaulon pycnostachyum (A. Mi-
chaux) Elliott.
Perennial herbs, often suffruticose, glabrous or
tomentose. Leaves alternate, sessile, decurrent on
alate stems, the margins entire or dentate. Capit-
ulescences glomerate or spicate. Capitula disci-
form, heterogamous, sessile; involucres campan-
ulate; phyllaries imbricate, linear-lanceolate,
scarious, caducous; receptacles hirsute or glabrous;
pistillate florets multiseriate, the corollas filiform,
truncate or 2-3-dentate, the styles of pistillate flo-
rets bifid, filiform, glabrous; disc florets hermaph-
roditic or functionally staminate, the corollas nar-
rowly tubular, the limb 5-lobed, the anther bases
sagittate, the terminal appendages ovate-lanceo-
late, rounded, the styles of hermaphroditic florets
bifid, pubescent distally. Achenes cylindrical to
fusiform, angled or compressed, costate; pappus
bristles 1-2-seriate. Chromosome number: n = 10.
Pterocaulon contains 25-30 species in North and
South America, Madagascar, Mauritius, Asia, and
Australia. Some 12 species occur in South Amer-
ica, with only the following species represented in
Peru.
References
CABRERA, A. L., AND A. M. RAGONESE. 1978. Revision
del genero Pterocaulon (Compositae). Darwiniana, 21:
185-257.
COLEMAN, R. 1968. Chromosome numbers in some
Brazilian Compositae. Rhodora, 70: 95-102.
1 . Pterocaulon alopecuroides (Lam.) DC. 4 , Prodr.
5: 454. 1836. Figure 19.
Conyza alopecuroides Lam., Encycl. 2: 93. 1786.
TYPE: Martinique, "on trouve cette plante a la
Martinique, dans les pres qu'on nomme Sa-
vannes. M. Commerson 1'a observee au Brasil,"
collector unknown (holotype, p, not seen).
Perennial herbs to 1.5 m tall, suflruticose; stems
erect, branched above, tomentose, alate, the wings
entire, discolorous, 2-3 mm wide. Leaves sessile,
decurrent; blade obovate-lanceolate to elliptic, 5-
7 cm long, 2-3 cm wide, discolorous, upper sur-
faces glabrescent, lower surfaces lanuginous, the
margins denticulate. Capitulescences spicate, ter-
minal, 3-17 cm long, compact, frequently inter-
rupted at the base. Capitula 5-8 mm high, ca. 3
mm wide; involucres campanulate; outer phyllar-
ies oblong-lanceolate, 2-3 mm long, ca. 0.5 mm
wide, lanulose, apiculate, the inner phyllaries nar-
rowly lanceolate, 4-6 mm long, 0.3-0.5 mm wide,
ciliolate, acuminate; pistillate florets 25-50, the
corollas 4-5 mm long; hermaphroditic florets 1-
2, the corollas ca. 4 mm long. Achenes ca. 3 mm
long, pilose, glandular; pappus bristles ca. 5 mm
long, white. Chromosome number: n = 1 (Cole-
man, 1968).
Pterocaulon alopecuroides is a widely distrib-
uted species ranging from the Caribbean Antilles
to southern South America, including Brazil, Peru,
Bolivia, Paraguay, Uruguay, and northern Argen-
tina. In Peru, it is found on the eastern slope of
the Andes at elevations below 1 200 m.
JUNIN. Tarma: Chanchamayo Valley, 1200 m,
Schunke 1533 (F). SAN MARTIN. San Martin: Tara-
poto, 1855, Spruce 4488 (F, GH, NY); Tarapoto, 840 m,
Woytkowski 35101 (F, MO).
XVIII. Stuckertiella
Stuckertiella Beauverd, Bull. Soc. Bot. Geneve,
ser. 2, 5: 205. 1913. TYPE: Gamochaeta cap-
itata Wedd. = Stuckertiella capitata (Wedd.)
Beauverd.
4 For complete synonymy, see Cabrera and Ragonese
(1978).
62
FIELDIANA: BOTANY
Annual to perennial herbs, branched, prostrate
to ascending. Leaves alternate, the margins entire.
Capitulescences glomerate, terminal, subtended by
upper leaves. Capitula disciform, sessile; involu-
cres hemispherical to campanulate; phyllaries 3-
4-seriate, imbricate, scarious; receptacles plane,
alveolate; pistillate florets multiseriate, the corol-
las filiform, irregularly lobed; disc florets her-
maphroditic, 4-5, functionally staminate, the co-
rollas tubular, 4-lobed, the anthers 4, base sagittate,
caudate, 3 with short, obtuse apical appendages,
1 longer, lanceolate, the style branches truncate.
Achenes fusiform to obovoid, surface with sinu-
ate-reticulate sculpturing, glandular with bicelled,
sessile papillae or trichomes, mucilaginous when
wet; pappus bristles scabrid, barbellate, apex blunt,
base fused, deciduous together as a unit, white.
A genus of two species confined to South Amer-
ica: Stuckertiella capitata (Wedd.) Beauverd from
Peru, Bolivia, and northern Argentina and S. per-
egrina Beauverd from central Argentina.
Reference
BEAUVERD, G. 1913. Le genre Stuckertiella Beauverd,
gen. nov. Bull. Soc. Bot. Geneve, ser. 2, 5: 205-209.
1. Stuckertiella capitata (Wedd.) Beauverd, Bull.
Soc. Bot. Geneve, ser. 2, 5: 206. 1913. Figure
20.
Gamochaeta capitata Wedd., Chlor. And. 1: 153. 1855.
TYPE: Peru, Dept. Cuzco, C. Gay s.n. (holotype,
p. not seen, F neg. 37584, fragment, F).
Gnaphalium capitatum (Wedd.) Griseb., Abh. Konigl.
Ges. Wiss. Gottingen, 24: 186. 1879. Non La-
mark (1786), nee Thunberg (1799).
Gnaphalium weddellianum Rusby, Mem. Torrey Bot.
Club 3: 57. 1893 (nomen novum for Gnaphalium
capitatum (Wedd.) Griseb.).
Annual to perennial herbs, to 25 cm tall; stems
usually prostrate, arachnoid-lanate, branched.
Leaves linear-lanceolate, 5-30 mm long, 1-2 mm
wide, sessile, apex acute, both surfaces arachnoid-
lanate, or glabrescent below. Capitulescences
glomerate at tips of branchlets, overtopped by ter-
minal leaves. Capitula 4-5 mm high, 4-5 mm wide;
involucres campanulate; outer phyllaries ovate, ca.
2.5 mm long, ca. 1.5 mm wide, base woolly, apex
obtuse to rounded, the margins scarious, the inner
lanceolate to oblong-lanceolate, ca. 3 mm long, ca.
FIG. 19. Flowering stem of Pterocaulon alopecur-
oides. (Drawn from Woytkowski 35101, MO.)
1 mm wide, apex rounded, scarious; marginal flo-
rets pistillate, 50-75, the corollas ca. 2 mm long;
hermaphroditic florets 4-5, the corollas 2-2.5 mm
long, 4-merous. Achenes obovoid, 0.4-0.5 mm
long; pappus bristles ca. 2.5 mm long.
Stuckertiella capitata is distributed from north-
ern Peru to northwestern Argentina, 2000-4000
m. In Peru, it occupies moist habitats between
30004000 m. This species is readily distinguished
by its capitula, which are subtended by upper
leaves, and by its 4-merous hermaphroditic flo-
rets.
ANCASH. Carhuaz: Cordillera Blanca, valley of Rio
Marcara, Viscos [Viscas], 2960 m, Hutchison & Wright
4291 (us). Huaylas: Huascaran National Park, Quebrada
DILLON & SAGASTEGUI: FLORA OF PERU. V.
63
B
FIG. 20. Stuckertiella capitata. A, habit; B, capitulum; C, outer phyllary; D, inner phyllary; E, pistillate floret; F,
hermaphroditic floret; G, anthers. (Drawn from Sagdstegui et al. 8993, HUT.)
Alpamayo, 4020-^4250 m, Smith et al. 9828. Yungay:
ca. 25 km NE of Yungay, slopes below Laguna de Llan-
ganuco, 3380 m, Dillon et al. 31 15 (F, MO, USM); Yungay-
Llanganuco, 3950 m, Lopez et al. 7442 (HUT). APU-
RIMAC. Abancay: Sayhuite, Km 45 E of Abancay, 3600
m; Hutchison 1737 (F, us); Ampay, 3500 m, Vargas &
SantanderSlO (F, MO), 57 3 (F). AYACUCHO. Prov. Un-
known: near Mataral, 3600 m, West 3670 (GH, MO). CA-
JAMARCA. Contumaza: Portachuelo de Nanra (Ven-
tarron-Totorillas), 3350 m, Sagdstegui et al. 8993 (F,
64
FIELDIANA: BOTANY
HUT, MO). CUZCO. Canas: Langui, 4000 m, Chavez 2324
(MO). Paucartambo: Hacienda Churu, 3500 m, Herrera
1049 (F, GH), 1399 (us). HUANUCO. Huanuco: Pam-
payacu, Sawada 95 (F). JUNIN. Huancayo: Quebrada
Occopilla, Soukup 3631 (F); Yauli: Laguna de Poma-
cocha, 4100 m, Ochoa 1991 (us). LIMA. Huarochiri:
Chicla, 3700 m, Asplund 11318 (us); Ticllo-Pap, 4800
m, Rauh & Hirsch P84 (NY). PASCO. Pasco: Km 5 Cerro
de Pasco-Huanuco, 4125 m, Ellenberg 4 1 30 (us). PUNO.
Huancane: Miajachi, Isla Chirone, 3900 m, Ortiz s.n.
(USM); Lampa: 9 km SW of Santa Lucia, 4000 m, Iltis
& Ugent 1426 (us); Melgar: Orurillo, Soukup 504 (F),
103 17 (MO).
XIX. Tessaria
Tessaria Ruiz & Pavon, Fl. Peruv. Prodr. 1 12, t.
24. 1794. TYPE: T. integrifolia Ruiz & Pa-
von.
Trees or large shrubs; roots giving rise to ad-
ventitious stems. Leaves alternate, the margins en-
tire or rarely dentate. Capitulescences cymose-pa-
niculate. Capitula disciform, short pedunculate;
involucres turbinate; phyllaries 3-5-seriate, im-
bricate, scarious, glabrous; receptacles conical, al-
veolate, pubescent; marginal florets pistillate, the
corollas filiform, 2-3-fid; disc floret hermaphro-
ditic, functionally staminate, solitary, the corolla
tubular, the limb 5-lobed, the anther bases sagit-
tate, caudate, the terminal appendages narrowly
ovate, the style unbranched, scabrous. Achenes
cylindrical, glabrous; pappus bristles scabrid, bar-
bellate, uniseriate, base fused, deciduous together
as a unit, white.
A monotypic genus as circumscribed by Rob-
inson and Cuatrecasas (1973). They transferred
several species traditionally placed in Tessaria to
Pluchea (see discussion under that genus).
References
ARIZA E., L. 1979. Contribution al conocimiento del
genera Tessaria (Compositae), I. Considerations sobre
los generos Tessaria y Pluchea. Kurtziana, 12-13: 47-
49.
CABRERA, A. L. 1939. Las especies Argentinas del ge-
nero Tessaria. Lilloa, 4: 181-189.
ROBINSON, H., AND J. CUATRECASAS. 1 973. The generic
limits of Pluchea and Tessaria (Inuleae, Asteraceae).
Phytologia, 27: 277-285.
WAISMAN, C. E., E. ROZENBLUM, AND J. H. HUNZIKER.
1984. Estudios cariologicos en Compositae. I. Dar-
winiana, 25: 217-226.
1 . Tessaria integrifolia Ruiz & Pavon, Syst. Veg.
Fl. Peruv. Chil. 213. 1798. TYPE: Peru [exact
locality unknown], H. Ruiz & J. Pavon 30/26
(holotype, MA, not seen; isotype, F.Fneg. 49304).
Figure 21.
Tessaria dentata Ruiz & Pavon, Syst. Veg. Fl. Peruv.
Chil. 214. 1798. TYPE: Peru, Dept. Lima, Prov.
Chancay, Chancay, H. Ruiz & J. Pavon 30/25
(holotype, MA, not seen, F neg. 29309; isotype, F).
Trees or large shrubs, 3-10(-15) m tall; stems
green to brownish green. Leaves oblanceolate to
lanceolate or elliptic, 3-9 cm long, 0.8-3.5 cm
wide, base attenuate, apex acute, mucronulate, both
surfaces minutely puberulent, glandular, grayish.
Capitula 4-5 mm high, 3-3.5 mm wide; outer
phyllaries ovate, 1 .5-2.5 mm long, 1-1 .2 mm wide,
apex acute, the inner linear to lanceolate, 4-5 mm
long, 0.5-1 mm wide, apex reflexed; marginal flo-
rets 40-80, corollas 3-3.5 mm long, styles exsert-
ed; hermaphroditic floret 1 , the corolla ca. 4 mm
long. Achenes 0.5-0.8 mm long; pappus bristles
ca. 20, ca. 5 mm long. Chromosome number: n =
ca. 10 (Waisman et al. 1984).
Tessaria integrifolia is widely distributed
throughout southern Central and South America.
It generally prefers sandy soils along the margins
of rivers, and at times forms pure stands from
adventitious root sprouts.
Leaves of this species are used medicinally for
hepatitis and as a diuretic, its wood is often used
in the construction of corrals, and its bark is said
to be used in the tanning of leather. In Argentina,
its wood is utilized in the production of paper (A.
L. Cabrera, pers. comm.), and around Iquitos, Peru,
it is an important forage source for cattle (F. Ayala,
pers. comm.).
In Peru, it known by the common names bobo,
chamchi, and mayu-chamchi, in Argentina as pd-
jaro bobo or aliso, and in Colombia as aliso, cen-
izo, sauce de playa, olivo, and sauce playero.
AMAZONAS. Bagua: between Bagua and Nazareth,
430 m, Croat 58344 (F, MO); trail from La Peca into
Serrania de Bagua, 1000-1400 m, Gentry et al. 23081
(F, HUT, MO, USM); 1 km de bajo La Poza, 180 m, Hu-
ashikat 44 (F, MO); rain forest along Rio Santiago 3-5
km above mouth, 250-300 m, Wurdack 2227 (F, GH, NY,
us). ANCASH. Carhuaz: Yungay-Marcara, 2400 m, Sa-
gdstegui et al. 12327 (F, HUT, TEX). Santa: Rio Santa
floodplain, 2 km N of Santa, 5 m, Stork & Horton 9153
(GH). Yungay: Mancos-Yungay, Smith & Bas 4939 (F,
us, USM). APURIMAC. Abancay: Abancay, 2400 m,
DILLON & SAGASTEGUI: FLORA OF PERU. V.
65
FIG. 21. Tessaria integrifolia. A, flowering branch; B, capitulum; C, pistillate floret; D, hermaphroditic floret.
(Drawn from Ridoutt 44a, HUT.)
Vargas 468 (F); Hacienda Matara, edge of Rio Matara,
2200 m, Vargas 9806 (F). AREQUIPA. Arequipa: Rio
Yarabamba, 2800-2900 m, Arenas 53 (USM); Tingo, ca.
7500 ft, Munz 15491 (F). Camana: Rio Ocona at Ocona,
4 m, Iltis et al. 1553 (MO, us). CAJAMARCA. Chota:
Carhuaquero, entre Chongayape y Llama, 500 m, Fer-
reyra 3165 (MO, USM). Jaen: Jaen, 500 m, Woytkowski
5585 (GH, MO, us). CUZCO. La Convention: ca. 5 km N
of Aguas Calientes, 2000 m, Solomon 3178 (MO); Machu
Picchu, 2000 m, Stork et al. 10490 (F), Vargas 823 (F,
MO). Urubamba: San Miguel, ca. 1 800 m, Cook & Gilbert
970 (GH, us). HUANCAVELICA. Angaraes: 4 km W of
66
FIELDIANA: BOTANY
Huanta, 2000 m, Stork & Morton 10806 (F.GH). HUAN-
UCO. Huanuco: downriver from Tingo Maria, 7000 ft,
Belshaw 3094 (F, GH, MO, NY, us); Huanuco, ca. 7000 ft,
Macbride 3231 (F, GH, us). Leoncio Prado: La Divisoria,
ca. 1 8 km NNE of Tingo Maria on road to Pucallpa,
1500-1600 m, Dillon 2647 (F, MO, USM), Gentry et al.
18845 (MO, USM), Plowman & Schunke 11705 (F, USM).
JUNIN. Tarma: La Merced, Acosta 19 (MO, USM), La
Merced, 700 m, Killip & Smith 23466 (GH.NY.US), 23690
(NY, us); Rio Perene, 2300 ft, Townsend 765 (F). LA
LIBERT AD: Pascasmayo: Pascasmayo, Fosberg 27995
(F, NY, us), Rose & Rose 18524 (NY), Williams 2910 (NY).
Trujillo: Trujillo, near sea level, Killip & Smith 21503
(GH, NY, us). LAMBAYEQUE: Lambayeque: E side of
Chiclayo, Hudson 938 (MO); road to Jaen, Km 17 E of
Olmos, at the Bridge of Silence, 570 m, Hutchison &
Wright 4423 (F, GH, MO, NY, us). LIMA. Callao: San Lor-
enzo, Andersson s.n. (F, GH, us). Chancay: Chancay, Ri-
doutts.n. (MO, USM 12195); Pativilca, Ridoutt s.n. (MO, USM
13946); Estanque Lampay, Hda. Paramonga, Ridoutt s.n.
(MO, USM 13999). Lima: Lima, Ball 176 (GH); between
Lima and Callao, Cook & Gilbert 2095 (us); Hacienda
Villa, ca. 20 km S of Lima, 100-120 m, Ferreyra 6133
(MO, USM); Chosica, ca. 3000 ft, Macbride & Featherstone
518 (F, GH), Holway & Holway 778 (GH); ca. 12 km SW
of Matucana, ca. 5800 ft, King & Collins 9002 (F, us);
Lima, Bajada de Miraflores, along Malecon, 100 m,
Plowman 11015 (F.USM); Santarrositas, Ridoutt s.n. (MO,
USM 10276); Lima, Soukup 1921 (us), 2054 (F); Barranco,
50 m, Weberbauer 7224 (F.GH.US). LORETO: Alto Ama-
zonas: Quebrada Shanuce above Yurimaguas, Croat
19775 (F, MO); entre Andoas y Capahuari Sur, 210m,
Diaz & Jaramillo 1371 (F, MO, USM); Andoas, Rio Pastaza
near Ecuador border, 210m, Gentry et al. 29725 (F, MO,
USM); Balsapuerto, 220 m, King 2999 (F, GH, MO, NY, us);
Maucallacta, Rio Paranapura, 200 m, Klug 3940 (F, GH,
MO, NY, us). Maynas: Lupuna, 100 m, Ayala 528 (MO);
Dtto. Iquitos, Isla de Escabino near Santa Maria de Na-
nay, McDaniel & Rimachi 17324 (F, NY); Padre Isla,
McDaniel & Rimachi 22297 (F); Rio Amazonas, Isla
Rondina, opposite Leticia, Plowman et al. 6384 (ECON,
F); Quebrada del caserio de Bombonaje Yanayacu, Ri-
machi 2567 (F), 2652 (F); lower Rio Nanay, Williams
494 (F, GH, us); Pebas, Williams 1863 (F, GH, us); Iquitos,
120 m, Williams 8024 (F). MADRE DE DIGS. Tam-
bopata: ca. 3 air km or 70-80 river km SSW Puerto
Maldonado, ca. 260 m, Barbour 5174 (F, MO, USM). PAS-
CO. Oxapampa: Huancabamba, 1830 m, Smith et al.
1665 (F, MO). PIURA. Piura: Piura, Townsend 1399 (us).
Talara: Parins Valley, Haught 146 (NY, us). SAN MAR-
TIN. Mariscal Caceres: Quebrada de Cachiyacu, 500-
600 m, Schunke 8494 (F, MO). San Martin: Vaca Pozo,
Rio Chumbazo al SE de Tarapoto, 830 m, Woytkowski
35178 (F.MO); Alto Rio Huallaga, 360-900 m, Williams
6891 (F.GH.US). UCAYALI. Coronel Portillo: cerca Div-
isoria, 1400 m, Ferreyra & Jones 19020 (USM). TACNA.
Tacna: near Tacna on road to Calientes, 500 m, Metcalf
30356 (GH, MO). TUMBES. Prov. Unknown: Ferreyra
10744 (MO, USM). NO EXACT LOCALITY. Matthews
s.n. (us); Wilkes s.n. (GH, MO, NY).
Acknowledgments
Illustrations were prepared by Segundo Leiva
Gonzalez, Universidad Nacional de Trujillo, and
Marlene Werner, Field Museum of Natural His-
tory. We wish to thank Timothy Plowman, Scott
Sundberg, and John Strother for helpful criticisms
and comments during the review process. Field-
work and manuscript preparation were assisted,
in part, by National Science Foundation grant
DEB-79-05078 (Alwyn H. Gentry, Principal In-
vestigator), NSF-Science in Developing Countries
Program Grant INT-8512104 (M.O.D., Principal
Investigator), and the National Geographic Soci-
ety. We thank Alicia Lourteig (p), Guy Nesom
(TEX), and Dan Nicolson (us) for help with no-
menclatural questions. The following herbaria have
lent or otherwise made their specimens available
for study, and their help is acknowledged: CPUN,
COL, GH, HUT, MO, NY, PRO, uc, us, USM. The acro-
nyms follow Holmgren et al. (1981).
DILLON & SAGASTEGUI: FLORA OF PERU. V.
67
1. Tumbes
2. Piura
3. Lambayeque
4. Cajamarca
5. Amazonas
6. La Libertad
7. San Martin
8. Loreto
9. Ancash
10. HuaViuco
11. Lima
12. Pasco
13. Junlh
14. Ucayali
15. lea
16. Huancavelica
1 7. Ayacucho
1 8. Apurfmac
1 9. Cuzco
20. Madre de Dios
2 1 . Arequipa
22. Puno
23. Moquegua
24. Tacna
DEPARTMENTS OF PERU
FIELDIANA: BOTANY
Index to Latin Names
Names for new species or combinations are in boldface, accepted names are in Roman type, and
synonyms or doubtful species are in italics. Page numbers in boldface refer to descriptions and numbers
in italics refer to illustrations.
Achyrocline 3
alata 4
madioides 4
peruviana 6, 7
ramosissima 6
satureioides 6
Antennaria 9
andina 53
aretioides 53
dioica 9
linearifolia 9, 10, 34
monica 34
sect. Mniodes 52
Baccharis
absinthioides 58
aretioides 53
graveolens 46
Belloa 11
chilensis 1 1
longifolia 12, 13
lopezmirandae 14
pickeringii 14
piptolepis 14
plicatifolia 13, 15
punae 1 1
schultzii 16
spathulifolia 16, 17
subspicata 16
turner! 18, 19
Blumea 18
balsamifera 18
lyrata 20
viscosa 20, 21
Chevreulia 22
acuminata 22, 23
filiformis 22
longipes 22
sarmentosa 22
Chionolaena 56
Conyza
alopecuroides 62
balsamifera 18
chingoyo 59
kunthiana 5 1
lyrata 20
marilandica 58
pusilla 51, 52
pycnostachya 62
thuyoides 42, 48
viscosa 20
Cuatrecasasiella 22
isernii 22, 24, 24, 42
Dolichogyne
acaulis 56
Ernstia
lyrata 20
Eschenbachia
lyrata 20
Facelis 24
lasiocarpha 25, 26
plumosa 23, 25, 26
retusa 24
schultziana 25
weddelliana 25
Filago
lasiocarpha 25
Gamochaeta 27
capitata 62, 63
americana 27, 28
humilis 29
monticola 29, 30
oreophila 29, 30
purpurea 31
spicata 28
Gnaphalieae 1, 56
Gnaphaliinae 56
Gnaphalium 1 , 32
alatum 4
americanum 27
antennarioides 34
badium 34
capitatum 63
celosioides 5
cheiranthifolium 33, 40
dioicum 9
dombeyanum 35
dysodes 35
elegans 37, 38
frigidum 39, 40
glandulosum 40
graveolens 4 1
helichrysoides 36
incanum 4
jelskii 35
kunthianum 52
lacteum 39
lanuginosum 36
linearifolium 9
luteo-album 39
melanosphaeroides 36
nanum 36
orientate 4 1
oxyphyllum 33
paramorum 34
pellitum 4
peruvianum 35
piptolepis 14
polium 40
purpureum 3 1
ramosissimum 6
retusum 24
rufescens 4
satureioides 8
sedoides 9
spicatum 28
subgenus Achyrocline 3
tunariense 56
uliginosum 32
weddellianum 63
Helichrysum 41
bracteatum 41
gnaphaloides 34
orientate 41
Inula 1
Inuleae 1
Jalcophila 41
peruviana 41, 42,
Leontopodium
linearifolium 9
Lepidophyllum
acaule 56
Loricaria 42
ferruginea 44, 45
graveolens 46
leptothamna 45, 46
lucida 45, 47
lycopodinea 47
macbridei 48
ollgaardii 47
DILLON & SAGASTEGUI: FLORA OF PERU. V.
69
pauciflora 49
stuebelii 49
thuyoides 42, 48
var. microphylla 47
var. stuebelii 49
var. thuyoides 49
var. thyrsoidea 48, 49
thyrsoidea 49
Lucilia 50
acutifolia 50
chilensis 1 1
conoidea 50, 51
kunthiana 51, 51
lehmannii 52
longifolia 12
lopezmirandae 14
pickeringii 14
var. /3 minor 14
piptolepis 14
plicatifolia 15
plumosa 25
pusilla 52
santanica 16
sect. Gamochaeta 27
sect. Lucilioides 1 1
tunariense 50, 56
Luciliopsis 23
isernii 22
perpusilla 23
Merope
aretioides 53
piptolepis 53
schultzii 16
Mniodes 52, 53
andina 52, 53, 54
aretioides 53, 54
cerratei 16
coarctata 54, 55
ferreyrae 55
pulvinulata 54, 55
tunariensis 56
Molina
ferruginea 44
incana 48
Novenia 56
acaulis 56, 57
tunariensis 56
Pluchea 58
absinthioides 58
chingoyo 59, 59
marylandica 58
microcephala 60, 67
suaveolens 60
zamalloae 60
Pseudoconyza
lyrata 20
viscosa 20
Pseudognaphalium 33
cheiranthifolia 40
Pterocaulon 62
alopecuroides 62, 65
pycnostachyum 62
virgatum 28
Serratula
acutifolia 50
Stuckertiella 62
capitata 63, 64
peregrina 63
Taffala
leptothamna 46
Tessaria 65
absinthioides 58
dent at a 65
integrifolia 65, 66
zamalloae 60
Tussilago
sarmentosa 22
Xeranthemum
bracteatum 4 1
70
FIELDIANA: BOTANY
Other Fieldiana: Botany Titles Available
1.ORA OF PERU. By J. Francis Macbride and collaborators (Alwyn H. Gentry, Michael O. Dillon,
and Samuel B. Jones). Conspectus and Index to Families. Family Compositae: Part I. Tribe Vemonieae.
Fieldiana: Botany, n.s. no. 5, 1980. 73 pages, 9 illus.
Publication 1314, $10.00
LORA OF PERU. By J. Francis Macbride and collaborators. Family Compositae: Part II. The Tribe
Anthemideae. By Michael O. Dillon. Fieldiana: Botany, n.s. no. 7, 1981. 21 pages, 3 illus.
Publication 1319, $3.00
LORA OF PERU. By J. Francis Macbride and collaborators. Family Compositae: Part HI. Genus
Mikania Tribe Eupatorieae. By Walter C. Holmes and Sidney McDaniel. Fieldiana: Botany, n.s. no.
9, 1982. 56 pages, 11 illus.
Publication 1333, $7.00
LORA OF PERU. By J. Francis Macbride and collaborators. Family Compositae: Part IV. Tribe
Cardueae. By Michael O. Dillon. Fieldiana: Botany, n.s. no. 10, 1982. 8 pages, 3 illus.
Publication 1335, $3.00
LORA OF PERU. By J. Francis Macbride and collaborators. Additions to Tribe Vernonieae (Com-
positae). I. By Michael O. Dillon. Fieldiana: Botany, n.s. no. 11, 1982. 7 pages, 3 illus.
Publication 1336, $3.00
Systematic Study of Flourensia (Asteraceae, Heliantheae). By Michael O. Dillon. Fieldiana: Botany,
n.s. no. 16, 1984. 63 pages, 41 illus.
Publication 1357, $10.00
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