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.

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1946. The historic tribes of Ecuador, pp. 785-821. In Steward, J. H., ed., Handbook of South American

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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

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

2.5—4.0 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 4—5 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, 3.5—4.0 mm long, 0.4—0.8 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, 3—4 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 (FP.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, 2—4 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 4—4.5 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, 3—4 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 3—4 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 5—40 cm

long, decumbent, tomentose to glabrate. Leaves

rosulate, sessile; blade obovate-spatulate to obo-

vate-oblong, 6—10 mm long, 3—4 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,

3—4 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,

3—4 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, 4—4.5

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 4—4.5 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, 3—4

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 25—40, 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 4—4.5 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, 1—1.5 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, 2.5—4 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 3—4 mm long.

Facelis plumosa is distributed from central Peru

to northern Argentina, 2800—4300 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, 1.5—4 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 3—4, 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 35—40, 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 3.5—4 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, 3100—4870 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, 5—4300 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, 1—4 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, 3—4 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 et

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 3—4 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, 3.5—4 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

Loricaria2 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 4—5 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 4—4.

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, 3—4 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 3.5—4 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, 4350—4400 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, 3—4 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 3.5—4 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 6—7.5 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

35—40, 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, 3—4 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, 3.5—4 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 0 (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. 4—4.3

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, 4—4.5 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 0 (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

3000—4000 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

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Other Fieldiana: Botany Titles Available

1.ORA OF PERU. By J. Francis Macbride and collaborators (Alwyn H. Gentry, Michael O. Dillon,

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Fieldiana: Botany, n.s. no. 5, 1980. 73 pages, 9 illus.

Publication 1314, $10.00

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Publication 1319, $3.00

LORA OF PERU. By J. Francis Macbride and collaborators. Family Compositae: Part HI. Genus

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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-

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Publication 1336, $3.00

Systematic Study of Flourensia (Asteraceae, Heliantheae). By Michael O. Dillon. Fieldiana: Botany,

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Publication 1357, $10.00

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