Flora of Peru: Family Asteraceae: Part V. Tribe Inulae

73 Pages·1991·29.6 MB·

by  Michael Owen Dillon

FIELDIANA
Botany
NEW SERIES, NO. 26
FLORA OF PERU
J. Francis Macbride and Collaborators
V
Family Asteraceae: Part
Michael O. Dillon
DepartmentofBotany
FieldMuseumofNaturalHistory
Chicago, Illinois60605-2496
Abundio Alva
Sagastegui
UniversidadAntenor
Orregode Trujillo
Trujillo, Peru
Accepted September 12, 1988
Published January 31, 1991
Publication 1422
PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY
Table of Contents XI. Jalcophila 41
1. Jalcophilaperuviana 42
XII. Loricaria 42
Key to Species ofLoricaria 44
ABSTRACT 1 1. Loricariaferruginea 44
TRIBE INULEAE 1 2. Loricariagraveolens 46
Key to Genera ofInuleae in Peru 2 3. Loricaria leptothamna 46
I. Achyrocline 3 4. Loricaria lucida 47
Key to Species ofAchyrocline 3 5. Loricaria lycopodinea 47
1. Achyroclinealata 4 6. Loricaria macbridei 48
2. Achyroclinecelosioides 5 7. Loricaria thuyoides 48
3. Achyroclineperuviana 6 Key to Varieties of
4. Achyroclineramosissima 6 Loricaria thuyoides 49
5. Achyroclinesatureioides 8 7a. Loricaria thuyoides var.
II. Antennaria 9 thuyoides 49
1. Antennaria linearifolia 9 7b. Loricaria thuyoidesvar.
III. Belloa 11 stuebelii 49
Key to Species ofBelloa 12 8. Loricaria thyrsoidea 49
1. Belloa longifolia 12 XIII. Lucilia 50
2. Belloa lopezmirandae 14 Key to Species ofLucilia 50
3. Belloapickeringii 14 1. Lucilia conoidea 50
4. Belloapiptolepis 14 2. Lucilia kunthiana 51
5. Belloaplicatifolia 15 XIV. Mniodes 52
6. Belloa schultzii 16 Key to Species ofMniodes 53
7. Belloa spathulifolia 16 1. Mniodes andina 53
8. Belloa subspicata 16 2. Mniodes aretioides 53
9. Belloa turneri 18 3. Mniodes coarctata 55
IV. Blumea 18 4. Mniodespulvinulata 55
1. Blumea viscosa 20 XV. Novenia 56
V. Chevreulia 22 1. Novenia acaulis 56
1. Chevreulia acuminata 22 XVI. Pluchea 58
VI. Cuatrecasasiella 22 Key to Species ofPluchea 58
1. Cuatrecasasiella isernii 24 1. Pluchea absinthioides 58
VII. Facelis 24 2. Pluchea chingoyo 59
Key to Species ofFacelis 25 3. Pluchea microcephala 60
1. Facelis lasiocarpha 25 4. Pluchea zamalloae 60
2. Facelisplumosa 25 XVII. Pterocaulon 62
VIII. Gamochaeta 27 1. Pterocaulon alopecuroides 62
Key to Species ofGamochaeta 28 XVIII. Stuckertiella 62
1. Gamochaeta americana 28 1. Stuckertiella capitata 63
2. Gamochaeta humilis 29 XIX. Tessaria 65
3. Gamochaeta oreophila 29 1. Tessaria integrifolia 65
4. Gamochaetapurpurea 31 ACKNOWLEDGMENTS 67
IX. Gnaphalium 32 MAPOF PERU 68
Key to Species ofGnaphalium 33 INDEX TO LATIN NAMES . . 69
1. Gnaphalium antennarioides 34
2. Gnaphalium badium 34
3. Gnaphalium dombeyanum 35
4. Gnaphalium elegans 37 List of Illustrations
5. Gnaphalium lacteum 39
6. Gnaphalium luteo-album 39
7. Gnaphaliumpolium 40 1. Achyroclineperuviana 7
X. Helichrysum 41 2. Antennaria linearifolia 10
1. Helichrysum bracteatum 41 3. Belloa longifolia, B. plicatifolia 13
111
4. Belloa spathulifolia 17 14. Lucilia conoidea, L. kunthiana 51
5. Belloa turneri 19 15. Mniodes andina, M. aretioides, M.
6. Blumea viscosa 21 coarctata, M. pulvinulata 54
7. Chevreulia acuminata 23 16. Novenia acaulis 57
8. Cuatrecasasiella isernii 24 17. Pluchea chingoyo 59
9. Facelis lasiocarpha, F. plumosa 26 18. Pluchea microcephala 61
10. Gamochaeta oreophila 30 19. Pterocaulon alopecuroides 63
11. Gnaphalium elegans 38 20. Stuckertiella capitata 64
12. Jalcophilaperuviana 43 21. Tessaria integrifolia 66
13. Loricaria lucida, L. leptothamna, L.fer-
ruginea 45
IV
FLORA OF PERU
V
Family Asteraceae: Part
Abstract thecorollasfiliformorligulate(rarelytubular),the
styles divided into linear branches; disc florets
ThetribeInuleae(Asteraceae)istreatedforPeru usuallyhermaphroditic,functionallystaminateor
andincludes 19genera, 55 species,and2varieties. occasionallygynoecia fertile, the corollastubular,
Keys to genera and species, descriptions, and il- the anthersconnate, appendaged, bases obtuse to
lustrations are provided forAchyrocline (5 spp.), sagittate, usually caudate, the styles divided, bi-
Antennaria(1 sp.),Belloa(9 spp.),Blumea(1 sp.), dentate, or undivided, the branches slender, su-
Chevreulia(1 sp.), Cuatrecasasiella(1 sp.),Facelis bulate,andattenuate,oroblong-linearandround-
(2spp.),Gamochaeta(4spp.),Gnaphalium(7spp.), ed, obtuse, or truncate, unappendaged; ovaries
Helichrysum (1 sp.), Jalcophila (1 sp.), Loricaria mostlyterete. Achenes mostlyterete, small, often
(7 spp., 2vars.),Lucilia(2spp.),Mniodes(4spp.), ribbed, variously pubescent, glandular, or gla-
Novenia (1 sp.), Pluchea (4 spp.), Pterocaulon (1 brous;pappusbristlesusually 1-2-seriate,smooth
sp.), Stuckertiella (1 sp.), and Tessaria (1 sp.). to scabrid, barbellate, or plumose, often fused at
Gamochaeta oreophila Dillon & Sagast. (nomen thebase,deciduoustogetherasaunitorpersistent
novum)isprovidedtoreplaceG. monticolaDillon on the achene.
&Sagast.,an invalidnameduetoanearlierhom-
onym. 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
Tribe INULEAE caudate anthers. In Peru, it includes 55 species
andtwovarietiesdistributedin 19genera. Several
Inuleae Cass., Jour. Phys. 88: 193. 1819. TYPE: gAenndeersa,hiancvleudcienngteBreslloofa,diLvoerrisciatryiai,natnhde MPneiroudveisa.n
Inula L. A few members are used medicinally (e.g., Achy-
GnaphalieaeRydb.,Fl.RockyMts.833. 1917.TYPE: rocline, Gnaphalium,Loricaria, Tessaria)andone,
GnaphaliumL. Helichrysum, is cultivated for ornamental value.
Dioeciousorbisexual,annualorperennialherbs,
References
shrubs, rarely trees; stems usually arachnoid or
glandular. Leavesalternateoropposite,caulineor MILLIARD,O.M.,ANDB.L.BURTT. 1981. Somegeneric
rosulate, the margins mostly entire. Capitulesc- conceptsinCompositae Gnaphaliinae.J. Linn.Soc.
ences paniculate, spiciform, cymose-corymbi- Bot., 82: 181-232.
form, glomerulate, or ofsolitary heads. Capitula MERXMULLER, H., P. LEINS, AND H. ROESSLER. 1977.
disciform,discoid,orrarelyradiate;involucrescy- Inuleae Systematic review, pp. 577-602. In Hey-
lindrical to campanulate or hemispherical; phyl- wood, V. H.,etal. (eds.),TheBiologyandChemistry
lariesnumerous,usuallyimbricate,indurateorhy- oftheCompositae, Academic Press, London.
aline, scarious, usually stramineous, sometimes TUR19N6E7R.,CBh.rLo.,moAs.omM.enPuOmWbEeLrLs,iAnNCDomJp.osCiUtAaTeR.ECXIA.SAPSe.-
white, or variously colored; receptacles plane or ruvian species. Ann. Missouri Bot. Gard., 54: 172-
convex, usually naked; marginal florets pistillate, 177.
DILLON & SAGASTEGUI: FLORA OF PERU. V.
Key to Genera of Inuleae in Peru
1. Plantsdioecious; individualcapitulaexclusivelyhomogamous(polygamo-dioeciousinAntennarid)
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. Stemsclearlyvisible, internodesevident; basalleavesrosulateandcaulineleavesalternate
orbasal leaves lackingandcauline leaves opposite, decussate, mostly linearto 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. Plantsbisexual; individual capitula heterogamous 5
5. Styles of hermaphroditic (or functionally staminate) florets briefly divided at the apex, the
branchespilosealongthedorsalsideandthedistalportionoftheshaft,apexrounded;orgreatly
divided with two linearbranches, apex acute to rounded, pilose 6
6. Stems winged, the wings discolorous; leaves discolorous; capitulescences spiciform-glo-
merulate; capitula sessile XVII. Pterocaulon
6. Stems unwinged, or ifwinged, 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 ofhermaphroditic florets divided, the branches linearto oblong, apex truncate to acute
or rounded, dorsally papillose orwith an apical ring ofpapillae 8
8. Achenes densely sericeo-pubescent 9
9. Plantscaulescent; stems sparselybranchedanderector much-branchedand cespitose
ordecumbent 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 scatteredglandularor duplex trichomes .
11
11. Achenes fusiform, terminally rostulate V. Chevreulia
11. Achenes ovoid to oblong-cylindrical, not rostulate 12
12. Pappus bristles free at the base (except Helichryswri), individuallydeciduous . .
13
13. Capitula solitary, pedicels evident; leaves glabrous orglandularon both sur-
faces 14
mm mm
14. Capitula20-50 wide;phyllariesvariouslycolored,greaterthan2
wide (cultivated) X. Helichrysum
mm mm
14. Capitula 4-5 wide; phyllaries green, less than 1 wide (native
species) IV. Blumea
13. Capitula glomerulate, pedicels obscure; leaves tomentose (at least lower sur-
faces) 14
15. Involucrescylindrical to narrowly oblongornarrowlycampanulate; 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 percapitulum XI. Jalcophila
FIELDIANA: BOTANY
16. Stemsgreaterthan2cmlong;leaveslinear-lanceolatetoovate,usuallygreater
mm mm
than 5 long,greaterthan 2 wide;pistillatefloretsususally 10ormore
percapitulum 17
17. Hermaphroditic florets 4-merous XVIII. Stuckertiella
17. Hermaphroditic florets 5-merous 18
18. Style branches of hermaphroditic florets acute to rounded, evenly
papilloseexternallyandlackinganapicalringofcollectingtrichomes
III. Belloa
18. Stylebranchesofhermaphroditicfloretstruncate,notevenlypapillose
externally, but with an apical ring ofcollecting trichomes
VIII. Gamochaeta
. .
I. Achyrocline A genus of 15-20 species distributed primarily
in tropical and subtropical South America, with
Achyrocline (Less.) DC., Prodr. 6: 219. 1838. five in Mexico and Central America and a few in
TYPE: Gnaphalium satureioides Lam. = tropicalAfrica,Madagascar,andComores.InPeru,
Achyroclinesatureioides (Lam.) DC. it is represented by five species distributed in a
widevarietyofhabitatsandelevationsthroughout
GnaphaliumsubgenusAchyroclineLess.,Syn. Comp. the Andean Cordillera. This genus most closely
332. 1832. resembles Gnaphalium in habit and floral struc-
Perennial herbs, suffrutescent or shrubs, to- ture;however,Achyroclineisreadilydistinguished
mentose or rarely glabrate; stems leafy, erect to by its narrow capitula with fewer than 25 florets
decumbent or rarely scandent. Leaves alternate; and glabrous achenes.
petiolate or sessile; blades linear to ovate, arach- Thisgenus is in need ofexperimental and field
noid to lanate, the margins entire to crenulate. study throughout its range. Several ofits species
exhibitconsiderablemorphologicalandecological
Capitulescences glomerulate, cymose-paniculate
to corymbose, open or dense, rarely of solitary variabilitythatpresentsproblemsfordetermining
heads. Capitula 3-5.5 mm high, ca. 1.5-2 mm species boundaries.
wide, disciform; involucres cylindric to narrowly
oblong or narrowly campanulate; phyllaries im- References
bricate, 2-3-seriate, scarious, brown to white or
rarely reddish, the outer gradually shorter, base DILLON, M. O., AND A. SAGASTEGUI A. 1986. Anew
lanate, the innerglandular on abaxial surface; re- speciesofAchyrocline(Inuleae-Asteraceae)fromPeru.
ceptacles flat, naked or rarely with rudimentary Phytologia, 59: 107-110.
paleae; pistillate florets 1-11(-23), the corollas fi- GIgAeNnGeUrAoLAAcNhIyr,oRc.liNn.e(C1o9m7p6o.sitLaaes)e.sDpaercwieisniaarngae,nt2i0n:a5s4d9e-l
liform, dentate or shallowly 2-4-cleft; hermaph- 576.
roditicflorets l-4(-6), fertile, thecorollastubular, MILLIARD,O.M.,ANDB.L.BURTT. 1981. Somegeneric
base dilated, 4-5-lobed, the lobes usually pubes- conceptsinCompositae Gnaphaliinae.J. Linn.Soc.
centwithmulticellular,stalked,capitate-glandular Bot.,82: 181-232.
trichomes,theantherbases sagittate,caudate, the JANSEN,R.K.,ANDT.F.STUESSY. 1980. Chromosome
terminal appendages lanceolate, obtuse, the style counts ofCompositae from Latin America. Amer. J.
branches slender, unappendaged, apex truncate, Bot.,67: 585-594.
penicillate, the nectary conspicuous. Achenes ob- JANVS.EFN,UNRK..K..1,9T8.4.F.RSeTcUuEeSSnYt,osS.cDrioAmZo-sPoimEiDcRoAsHfernA,CoAmN-D
long to ovoid, subcompressed, ca. 4-5-ribbed, positaede Colombia. Caldasia, 14: 7-20.
papillosewith imbricatecells;pappusbristlessca- NESOM,G. L. 1990. TaxonomyofAchyrocline(Aster-
brid, barbellate, uniseriate, bases cohering by pa- aceae: Inuleae)in MexicoandCentralAmerica. Phy-
tent cilia, readily deciduous, apical cell rounded. tologia,68: 181-185.
Chromosome number: n = 14.
Key to Species ofAchyrocline
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. Plantswith reducedhabits, usually lessthan 50 cm tall; leaves2-3(-5)cm long, 1-3 mm wide
A. ramosissima
3. Plants not reduced in habit, usuallygreaterthan 1 m tall; leaves 4-10 cm long, 2-9 mm wide
A. satureioides
2. Leaves lanceolate to elliptic or obovate; outer phyllaries lanmcemolate, apex acute 4
4. Leaves elliptic to obovate, (0.5-)1.3-3.5 cm long, 5-16 wide A. celosioides
4. Leaves lanceolate to elliptic, 1.5-5 cm long, 2-8 wide A. peruviana
1. Achyroclinealata(Kunth) DC., Prodr. 6: 221. Achyrocline alata has a wide distribution
1838. throughouttheAndeanCordillerafromColombia
to Argentina, Brazil, and Paraguay. It inhabits a
Gnaphalium alatum Kunth, Nov. Gen. Sp. (fol.) 4: variety ofenvironments within its range, but in
62. 1820. TYPE: Ecuador, Prov. Chimborazo, Peru it is usually found in secondary vegetation
"circa Alausi Quitensium," A. Bonpland 3243 ordisturbed roadsides(2400-4100 m). In Peru it
(holotype,P,notseen,IDCMicrofiche6209. 96: is known by the common names: ajenjo, Foster-
II. 4, Fneg. 37578, fragment, F).
Gnaphalium incanumKunth, Nov. Gen. Sp. (fol.)4: sacha, huira-huira, ishpingo amarillo, vira-vira,
63. 1820. TYPE: Peru, Dept. Cajamarca, Prov. yatama blanca, andyshpingo.
Hualgayoc,"juxtaurbemMicuipampa," 1830m, Achyroclinealata is readily distinguished from
A. Bonpland 3680 (holotype, P, not seen, IDC otherPeruvian species by its decurrentleafbases
GnapMhiaclriofuimchpeel6l2i0t9u.m K96u:ntIhI.,5N,oFvn.egG.en3.76S0p1.).(fol.) 4: and alate stems. Pubescence is quite variable; in-
63. 1820. TYPE: Peru, Dept. Cajamarca, Prov. dividuals range from predominately glabrous to
Hualgayoc,"juxtaurbemMicuipampa," 1830m, densely arachnoid-tomentose. No ecological or
A. Bonplands.n. (holotype,P,notseen,IDCMi- geographical patterns are obvious for these pu-
cF)r.ofNiochneG6.20p9e.ll9i6t:umII.Ho6r,t.FnGeegt.ti3n7g,60e4x,DfCra.gment, bescencemorphs,andthecapitularandfloralmor-
GnaphaliumrufescensKunth,Nov. Gen. Sp. (fol.)4: phologyisrelativelyconstant. Gnaphaliumincan-
61. 1820. TYPE: Colombia, Prov. Cauca, "inter um, G. pellitum, and G. rufescens are all readily
fluvium Putes et villam San Miguel," A. Bonp- referable to Achyrocline. They are all quite pu-
lands.n. (holotype, P, not seen, IDC Microfiche bescent, and their decurrent leaf bases forming
Achyr6o2c0l9i.ne96m:aId.io7,idFensegM.ey37e5n71&, fWraalgpmeernst,,NFo).v. Ac- wingedstemsandfew-floweredcapitulaagreewell
torum Acad. Caes. Leop.- Carol. German. Nat. withthe presentbroadconceptofA. alata.Achy-
Cur. (suppl.) 19: 275. 1843. TYPE: Peru, Dept. roclinemadioidesMeyen &Walpersisprovision-
Arequipa, W. G. Walperss.n. (holotype,B,prob- allyplacedherealso.Furtherdetailedpopulational
ablydestroyed, Fneg. 15134). investigations may lead to the taxonomic recog-
m nition ofthese pubescent morphs.
toPdeerceunnmibaelntheorrbssctaond1e.n5t,gltaalblr;ousstetmostoamsecnetnodsien,g stIenmdwiivnigdsu,alasnwdistmhalslmearl,lefrewl-efavleosw,ergeredactalpyirteuldaucaepd-
bla1lar5anotcueme.somlLaaentawdevi,egddlseea,scnedutsurrsilrilaenerne;rttbvoloeaddndefenssrtleeoilmmnsye,antreo2-am-lrea1nn2tctheoceosmlbeaatlbseoeent,ngoe,gaol2tba-h-- puulrnaow.aiWcnhhgiAeldessaAtt.eurmaesli,aotiaadneasdn,dnaaAsr.presocawitleusyrewciiyotlihidneldsirniaecraaerllcelcaoavspeiestl,-y
and arachnoidabove, base attenuate, apex acute, erevlaattieodn,stbheelloawtte1r50sp0ecmi.esWies thyapvicealnloytffoouunnddastyeml--
the margins entire to crenulate. Capitulescences
glomerulate, cymose-paniculate, dense or open. patric populations to date, but two collections
Capitula 3.0-5.5 mm high, ca. 1.5 mmwide, ses- (Vargas 456, F, and Weberbauer 5638, GH) have
stems both with and without wings.
sile; involucres cylindrical to narrowly campanu-
late,yellowtocream-white;outerphyllariesovate
tolanceolate, 2-3 mmlong, 1-1.5 mmwide,apex AMAZONAS. Chachapoyas: 28-31 km SW ofLei-
talicolunltgae,t,e0t.fhl5eo-r1ie.nt5sne3mr-pm1hy1lw(l-ia2dr3ei),e,satlphieenxecaaorcr-uospltaletautsloa2ot.eb5,t^u44s.-e05;mmpimms- atmCloae.lbBl2aaa3lCms1aab5ls8la,a,(3oFW,n1M1rOs0oi)mda;e,d,44t1Go5eknkBtammrlsyaSabseW,tova3ole.3fB5L2a0l3e-s1ia34sm4,7e53b0(aF1,0mmM0,bOam)G,;eonHnCtuertrrycorhaoe-dst
long; hermaphroditic florets l-3(-6), the corollas ison & Wright 5774 (F,GH, MO,NY); Cerro Chimchula,
Co2v.ho5ri4od.,m0o0.sm5o-mm0.e7lonmnugm.mbAelcroh:nge;nnep=sap1np4aur(srJocaawn.lsye2n.e5l&lmiSpmtsuoelisodsnygt.,o crM2eoe2rcn0ckd0ayo-azs2la3Co:0ph0eMascemhon,fadpoCWozauyanra,ods,a1Sc6a20nk5t070am20L,6umcW(,iGoaHyO,jtcuNkshYoto,wEasukso)1if.68C2Rh28oa7,dc3rh1i(a6gGp3Huo,7eyzMa(OFsd)),e.;
1980). ANCASH. Bolognesi: Valle de Rio Fortaleza, 3600 m,
FIELDIANA: BOTANY
Cerrate & Gayoso 7819 (MO); Chiquian, 3350 m, Fer- (MO,USM);Tarma, 3000-3200m,Killip&Smith21914
reyra7455(MO,us).Casma:abradelaCordilleraNegra, (F,NY,us); Huacapistana, 1800-2400m,Killip&Smith
4050m,Mostaceroetal.536(F,HUT,MO).Huaraz:Cerro 24206 (NY, us); Tarma, Macbride & Featherstone 988
San Cristobal, 3800 m, Evangelista s.n. (F, HUT, MO); (F);Sacsamarca,Riccio1541(us);entreTarmayOroya,
LagunadeShurup,3800m,Evangelistas.n.(F,HUT,MO). 10kmdeTarma,3600m, Tovar2380(USM).Yauli:San
Huaylas: Laguna de Paron, 4100 m, Mostacero et al. PedrodeCajas,SofLakeJunin,3800-3900m,Pearsall
540, 553 (F, HUT, MO). Yungay: Llanganuco, Ferreyra 924(F).LALIBERTAD.Otuzco:Agallpampa, 3050m,
14342, 16826, 16863 (MO, USM); slopes below Laguna Lopez 1873 (F, HUT). Pataz: Huancaspata-Tayabamba,
de Llanganuco in Quebrada de Llanganuco, ca. 25 km 3900 m, Lopez& Sagdstegui8266 (HUT,MO). Santiago
aboveYungay,4000m,Edwin&Schunke3818(COL,F, deChuco:LagunaLaVictoria,caminoaConzuzo,4000
us); Yungay-Yauya road, 3480 m, Gentryet al. 37366 m,Sagdsteguietal.6190(HUT,MO).LIMA.Canta:3100-
(3F4,5M0O).m,APFeUrRreIyMrAaC9.81A2ba(nMcO,ayU:SMa)rr.ibAaymdearAabeasn:ca4y,km335S0W- d3e20I0nfime,rniPlelno,neClelrr1a4t6e1e2ta(lF.,4N2Y7,8us()M.O,HUuSaMr)o;chCihriic:laP,ueennttree
ofCotaruse,26 kmSWofChalhuanca, 3220m, Gentry SanMateoyCasapalca, 3700m,Ferreyra6505(MO,us,
etal. 23302(F,MO,USM). AYACUCHO. Huanta: Ccar- USM); Infiernillo, 3300 m, Ferreyra 8318 (MO, USM); 4
rapa,betweenHuantaandRioApurimac, 1200m,Killip kmEofChicla,3800m,Gentry&Prance16448(F,MO);
&Smith22430(F,us).LaMar:aboveAyna(Aina), 1600 RioBlanco, 3000-3500m,Killip&Smith21637(F.NY,
mmala,.r2cWa9e:0b4ecar(.bF,a6M.u5Oe,rkT5mE6X3S,8WUS(MFo,)fG;CHSa,ajuansm)Ja.urCacnaA,Ja2AC9aM8jA0aRmmaC,rAcDa.i,llC2oa8nj0ae0-t Fuiesel)a,tCMhaearrcrsibtoronin:deeY1&1an9Fa3eh(auFt,ahUneSc)r.as,tPoansceca.o6:71c2a0.,(F02,03u0sk).mft,PNAMNSaEcCbOor.fiCdDeearnr&-o
m, Ochoa 1475(GH,F);LaEncanada-JalcadeKumulca, de Pasco on road to Huanuco, 3810 m, Dillon 2552(F,
(l3eH3nU0dT0i,n:mMO,4);SkamNgdENstNoeEfguPioofreCtceoalnl.e,n8Sd1ai0nn0coh(neF,zrHo2UaT8d,5t8MoOB(,aClPNsUYaN)s,,,F8E)1.d0wC5ieAn- ebMrOas,mtboUanSe,M)2;209L25a10Q(mFu,,iunsuH)au.,tcPchIai.UsRo1nA2,.0106H00u3aftn,(cFM,aabNcYab)mr;biadMe:it&HouFpaeanamctpaha--
& Schunke3582(COL,F,NY,us); entre Celendin yJalca (Huancabamba-Cuello del Indio), 2650 m, Sagdstegui
Cumullca, 3000-3100m,Ferreyra 15087(MO,us,USM); et al. 8227 (MO, NY). PUNO. Carabaya: Macusani to
Celendin,2700m,Hutchison& Wright5127(F,MO,NY); Ollachea, 2000-3000m,Dillonetal. 1099, 1265(F,MO,
Canyon of the Rio Maranon above Balsas, 2930 m, USM); Ollachea to San Gabon, 1000-2000 m, Dillon et
Hutchison& Wright5374(F.MO.NY).Chota:Cochabam- al. 1103(F,MO,USM).SANMARTIN.MariscalCaceres:
ba, 1800-2200m,Ferreyra783(USM).Contumaza:Guz- Rio Abiseo National Park, 3425 m, Young & Watson
mango,Sagdstegui7726(F,HUT);Cascabamba,3050m, 3519(p).
Sagdstegui et al. 9038 (F, HUT, K, MO); Jalca El Chuno
(Pozo Chuno), 4500 m, Sagdsteguiet al. 9373 (F,HUT,
HM2OU8)T0;,0MCmOr,)u.zStCdouetrlekrH&vuoeH:co4or,tko2mn8E0109o1fm9,H9uS(aaF)gm.dbCsotUseZgouCniOCe.tutAalne.tra9v:9o1Lt8riami(lF-,, 2. Ac2h2y1r.oc1l8i3n8e.celosioides(Kunth)DC., Prodr. 6:
aNp33at34Ela00lm0p0obafm,omP,,,3a2Cu2P0ch4e0aa0nrv0nmtee,azlmml3,Vb2ao18r,V3g3a62ar2(9sgM68aO4s0)5(.-F64,3C50G(u6FH5z),.0c(oPMNm:OaY,)u,S.caGuanesCrn)aSt.tlearcbPmyaaab:serottuiA:aramlon9.p:,-a21r33A2a34rc01aks03ym,,-- GnapjB4hu:oaxn6tlp1ai.luaL1mno8d2xsc0ae..lnmo.TsPYi(oePhiroEdule:vostiEyKacpunuenoa,trdhPuo,,mrn,,No"PotrAvs.o.eveH.nGu,eLmonIbj.Dao,lCSpd"M.ctir(ec&fsrocloAi.-.t)
23423(F,MO,USM);22kmNEofPaucartambo,3350m, fiche 6209. 96: II. 2, Fneg. 37585).
Gentry et al. 23430 (F, MO, USM); Cerro de Cusilluyoc, m
Pillahuata,2700-2900m,Pennell14089(GH).Urubam- Perennial herbs or shrubs to 0.5 tall; stems
ba: Machupicchu, 2400 m, Herrera 1974(F); Ollantay- much-branched,ascendingtoprocumbent.Leaves
tHPaaUumcAbaoNr,cU2aCn8cO0h0.a,m,Hu3Ha2em2ra5rleiremas,:34PL4el7aytt(aFo,)n;7M0&a0c0hPuefty,PtioMcnacchub1,r3i3adbeo(vF)&e. sleosnsgi,le(;0b.l3a-d)0e.e5l-l1i.pt6icctmoowibdoev,atter,in(0e.r5v-e)d1f.5r-o3m.5necamr
Featherstone 2242 (F,GH, us). Huanuco: cerca a Mito- the base, discolorous, both surfaces tomentose,
cucho, 2600 m, Cerrateet al. 6887 (MO,USM); between apex obtuse to rounded, base obtuse to cuneate.
HuanucoandChinchao,Dwyer6115(MO);Sheiricancha, Capitulescences glomerulate, cymose-paniculate,
cerca a Acomayo, 2500-2600 m, Ferreyra 9333 (MO, 45 mm mm
USM);Acomayo,DistritodeChinchao, 2500m,Ridoutt denseoropen.Capitula high,ca. 1.5
11560(MO,USM). Pachitea: Mitotambo,arribadeMito, wide, sessile; involucresmcmylindrical, whimtem; outer
3200-3300 m, Ferreyra 10380, 10380A (USM). ICA. phyllariesovate,ca. 2.5 long,0.8-1 wide,
P(&uNNsa3)YzF5.)ec.4aaHt:(uPNhiaYesPn)rcuc.osqa:tJuyoUiVonNao:elIl1KVNe4am.7lRl4Cei,7oyo,n1Hc4Pu3e7iap4s5ntc0coi(0a,Fo)yn,mo3:,1-7M6P0iRa40tar4oium,(a,hF9h.0uUR&Sa0a)n0,uHcfiha1tr,,8sM&1c3a8h3cH(0biPFr0r.4isG3mdcH5,he, araloapplneelcxxaesoaal2cca.uut0tte-ee,,2t.3to5.ha5emp4iim.cnu0nlleamomrtnemgmp;;hyphllieslotrnaigrml,ialeaps0the.rl4foal0dnoc.iret8eotilscma2tf,melottrhwoeetisodcbeo3,--,
AntunezdeMayolo38(F); Huancayo, 3317 m, Soukup thecorollas2.2-2.5 long.Achenesovoid,0.7-
2O9c5h5oa(C5O4L)9,(F3)0.0T3ar(Fm)a,:3c1u51mb(rCeOLa,Tusa,rUmSaM,).35J0au0jma:,CCoermraast,e 0.8 mm long; pappus ca. 2.5 mm long.
922(MO,USM); 28-32 km NEofTarma on roadto San
Ramon,2480m,Dillon& Turner1348(F,MO,TEX,USM); Achyrocline celosioides ranges from southern
entreTarmaySanRamon, 1800-2100m,Ferreyra3606 Ecuadortosouth-centralPeru(2000-3400m)and
DILLON & SAGASTEGUI: FLORA OF PERU. V.
normally occupies relatively moist sites in inter- A. &S. LopezM. 10661 (holotype,HUT). Fig-
Andeanvalleys. ItisreadilydistinguishedfromA. ure 1.
peruviana, its nearest relative, by the former's
shorter, elliptic leaves. Suffrutescent perennials to 75 cm tall, much-
A few populations display considerable varia- branchedatthebase; stemsarachnoid-tomentose.
tioninleafsizeandshape.Twounusualcollections Leaves sessile; blade narrowly lanceolate to ellip-
from the Cumbre El Gavilan (above Cajamarca) tic,(1.5-)4-5cmlong, 2-8 mmwide,discolorous,
displaythefollowingvariability:(1)Ferreyra3277 base attenuate, apex acute, mucronulate, lower
(us), has very densely lanate stems with short in- surface densely arachnoid-tomentose, upper sur-
ternodes; ovate, sessile leaves; and sessile, glo- face sparsely arachnoid-tomentose, glabrescent,
merulate capitulescences in the upper leafaxils; obscurely trinervate, the margins entire. Capitu-
(2) Ferreyra 8538 (MO) has very small leaves (ca. lescences glomerulate, cymose-paniculate, open,
mm mm mm
5 long, ca. 3 wide) and capitulescences terminalandaxillary.Capitulaca.4 high, 1.2-
mm
on short peduncles. 1.5 wide, sessile; involucres narrowly cam-
Two collections, Sagdstegui & Dillon 12312 panulate,stramineous;outerphyllarieslanceolate,
mm mm
(Dept. Ancash) and Chavez 3372 (Dept. Apuri- 2-3 long,ca. 1 wide,apexacute,theinner
mm mm
mac) have more narrowly elliptic, sessile leaves phyllaries lanceolate, 3-4 long, 1-1.2
and compact, nearly sessile glomerulate capitu- wide,apexacute;pistillateflorets 1-2,thecorollas
mm
lescences. Last, acollection from a dry site above ca. 2 long; hermaphroditic florets 2-3, the
the Rio Maranon(Evinger& Hodge514, US) has corollas ca. 2.5 mm long. Achenes ovoid, ca. 1
mm mm
smaller,morenarrowlylanceolateleaves,butwith long; pappus ca. 2.5 long.
capitulescences typical for the species. All these
specimens essentially agree with A. celosioides in Achyrocline peruviana is a narrow endemic
capitularandfloralcharacteristics,butfurtherstudy known only from dry intermontane localities in
may prove them to be worthy ofspecific rank. the Departments ofCajamarca and La Libertad
(2600-3100m)andisprimarilyrestrictedtoriver
CShhAaicpMhaAaspbZoaOymaNbsaA:,S11.502B0konmmg,aErHau:otfbcehBtaiwlsezoeansn&R[iBWaolrsUiagtsh]c,tub3Eav9im9nb4gae(uracn)&d. vlyalrleeysesmybslteesmsA.wictehloPsaicoiifdiecs,drbauintagdief.feItrsmofsrtomcltohsea-t
Hodge514 (us); rocky slopes ofCano Santa Luciajust species by possessing more narrowly elliptic or
E ofChachapoyas, 2000-2400 m, Wurdack 532, 579 lanceolate leaves, usually at least 4 cm long, with
c(RaGI.H,M9AN0YC0,.0ucfAt,,bauMsna)c.cabyAr:NidCCeuAr&SahHuF.aesaRite,hceur2as5yt5:o0nremo2c,k5y1C7hrai(vvFe)er.zbAl3Pu3fUf2s-7, acuStoe,memuicnrdoinvuidluaatlesaopfiAc.ess.atureioidesapproachA.
(MO).CAJAMARCA.Cajabamba:caminoaLangla,2600 peruviana in general appearance, but the former
m,Mostacero&Guerra0008(HUT).Cajamarca:Cumbre species has strictly cylindrical capitula, with a
El Gavilan, entreCajamarcay San Juan, 3200 m, Fer- greater number of pistillate florets and a distri-
reyra 3277 (us), 8583 (MO); La Encanada-Jalcade Ku- bution usually below 1500 m on the eastern es-
amdual,ca2,735300m0,m,SaSgadgsdtsetgeugiuietetala.l.182100163((HHUUTT));.LCaelEenncdainn-: carpment ofthe Andean Cordillera.
ca. 31 km NE ofCajamarca along road to Celendin, E
aslnodpetso,Nca.of29to5w0nm,,caD.il2l7o0n0&m,TuHruntecrhi1s5o8n7&(F)W;rhiilglhsta5b1o3v6e 26C20AJmA,MDAilRlConA.etCaol.nt4u4m5a5za(:F,1H-U2T,kmMOS,oTfECX,onutsu,mUaSzMa),;
(uc).Hualgayoc:Bambamarca. 2600m,Soukup&Car- Contumaza, 2600 m, Sagdstegui et al. 10685 (F, HUT,
mona 4935 (us). JUNIN. Conception: Mito, ca. 9000 MO).LALIBERTAD.Huamachuco:Hda.Cochabamba,
ft, Macbride& Featherstone 1489 (F). Prov. Unknown: 2750m,Lopez&Sagdstegui2781(HUT).Otuzco:Agall-
(UFs).paLcAhaLcaI,BcEaR.T8A5D0.0Sfta,nMtaicabgoriddeeC&huFceoa:thHearcsiteonndea1A2n9-5 pampa, 3100 m, Lopez895 (us), 1867(F,HUT).
gasmarca, 2850 m, Weberbauer 7205 (F,GH). PIURA.
aHnudanWcaobfamHbuaa:ncoanbarmobada,to29C0a0ncmh,aqHuuet,ch1i8s.o5nk&mWarbiogvhet 4. Achyrocline ramosissima (Schultz-Bip.) Brit-
6635(F,MO,NY,uc,USM). ton, Bull. Torrey Bot. Club 19: 148. 1892.
GnaphaliumramosissimumSchultz-Bip.,Bonplandia
3. Achyrocline peruviana Dillon & Sagast., Phy- 4:42. 1856.TYPE:Peru,Dept. Puno,Azangaro,
tologia60: 107. 1986.TYPE: Peru,Dept. Ca- cordillerapropeAzangaro, 1854, W.Lechler1780
(holotype, B, probablydestroyed; isotypes,F,NY,
jamarca, Contumaza, El Granero, laderas G-DEL,notseen,Fneg. 28706,w,notseen, Fneg.
abiertas, 2800 m, 14Jun 1983,A. Sagdstegui 33153).
FIELDIANA: BOTANY
FIG. 1. Achyrodineperuviana. A, habit; B; capitulum; C, inner phyllary; D, outer phyllary; E, hermaphroditic
floret; F, pistillate floret; G, stamen; H, style branches ofhermaphroditic floret. (Drawn from Sagdstegui & Lopez
19661, HUT.)
Perennial herbs, 15-50(-70) cm tall, suffruti- entire, revolute. Capitulescences corymbiform
cose; stems much-branched, decumbent to as- cymes ofglomerules, dense or open. Capitula ca.
mm mm
cending, densely lanate. Leaves sessile; blade lin- 4 high, ca. 1.5 wide, stramineous; in-
eartolinear-lanceolate, 2-3(-5)cmlong, 1-3 mm volucrescylindrical; outerphyllaries ovate, 1.5-2
mm mm
wide, discolorous, the lower surfaces densely la- long, 1-1.2 wide, 2-3-fid, base lanate,
nate, the upper surfaces tomentose, the margins the innerphyllarieslanceolate to oblanceolate, 3-
DILLON & SAGASTEGUI: FLORA OF PERU. V.