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A Revision of Pectis Section Pectothrix (Compositae: Tageteae)

A Revision of Pectis Section Pectothrix (Compositae: Tageteae)

A REVISIONREVISION OF PECTISPEens SECTIONSECTlO~ PECTOTHRIXPECTOTHRIX (COMPOSITAE.(COMPOSITAE: )TAGETEAE)

DAVIDDA\ID J.J. KEILKEIl

PectisP~('lis L.l. sect. PectothrixPUlOlhri.'C A. GrayGray (Compositae:(Compositae: Tageteae)Tageteat) comprises a group of showy,showy. yellow-headedyellow-headed species which.... hich occursoccurs mainlymainl} inin thethe arid and semi-arid portionsportions ofof thethe southwesternsouthwestern UnitedUnIted States .and northernnorthern Mexico.Meltico. AlthoughAllhough severalse\-eral regionalregional floras have includedincluded one or moremore of thethe species inin thisIhis group,group. nono workersworl.:el'$ since Fernald (1897) and RydbergR}dbcrg (1916) have attemptedattempted to10 treatlreal this portion of thethe genus inin itsits entirety.emirely. SinceSince 1916,1916. severalsc\'eral workers have described new taxataxa (Cory, 1937; Johnston,Johnston. 1940;1940: Wiggins,Wiggins. 1951;1951: Keil,Keil. 1974),1974). and manymany new collections haveha\'C beenbttn made. Recent studiessiudies (Keil,(Keil. 1974) haveha\'e suggested thatthat thethe species relationships proposed by earlier workers.... orkers (Fernald,(Fernald, 1897;1897: Gray,Gray. 1884;1884: Rydberg,Rydberg. 1916) should be re-examined.rc-uamined.

TAXONOMICTAXO-':O\1IC HISTORYIIISTOR Y The species included in thethe present reportreport belong toto a complexl;omp1ex treatedtrealed in the past as two subgenera, PectidopsisPI'C/i(IQPsis and PectothrixPeclOtllri.r (Fcrnald,(Fernald, 1897). Pectidopsis,Pectitlopsis. originallyoriginall}' described as a mono-mono­ typict}pic genus (DeCandolle.(DeCandolle, 1836),1836). was reduced in rankrank to a section of PectisPI'Ct;s byb}' Gray (1852), and was subsequentlysul>sequenlly mergedmcrged withwilh sect.sec\. £llpt'ClisEupectis by Gray (1884). Fernald (1897) resurrected PectidopsisPu/idopsis as a SUbgenussubgenus and included "'itbinwithin it twelvetwelve species with thethe pappus coromformcoroniform and few-bristledfe .... -bristled or few-awned.few-a'" ned. Pectothrix,putothri:r. dcscnbeddescribed by Gra}Gray (1&49)(1849) as a subgenus of ,PI'C/is, was",,-as reduced to sectional rank three:three years}ears later (Gray.(Gray, 1852).1852), but was""as returned to10 subgcncnesubgeneric 1e\,<:1level in FernakTsFernald's (1897) revision. Throughout its hIstory.history, P~(,'OIh,ixPectothrix has been used for species with""llh a multisetosemulliselOSC pappus. Although RydbergR)"dbcrg (1916).(1916), the last worker to10 revisercvise this part of I'utis,Pectis, did not accept any infragenericIOfragcncric categories, inIn most cases he retained the:the arrangement of sp«iesspecies thatthai FernaldFe:rnald had proposed.

RF-F\I\IRE-EVALUATION tAllO' Of- OF 1'\1INFRAGENERIC IlA(,1 '[RtC IlOlBOUNDARIES 'OARIF'

Previous imcstigalorsinvestigators have:have e:mphasindemphasized in varying deg~esdegrees thethe' features of thethe pappus as characters for subdividing PeClis,Pectis, firstfirs!

J232 1977] Pectis•KeilPectis-Keil 333J into segrcgatesegregate genera (I..es.~ing.(Lessing, 1830, 1831;1831: DeCandolle, 1836), and later into sections or subgem:rasubgenera (Gnly,(Gray, 1849,1849. 1852, 1884:1884; Fernald.Fernald, 1897). Although pappus characters can be used in some cascscases in conjunction with other features in delineating infrageneric categories in PectisPee/is (e.g., .~ect.sect. HeteropectisHe/eropeClis A. Gray), the use of thesethese features alnnc.alone, like any other one-eharaeterone-character , isis SUbjectsubject to error. Fernald's a priori reliancerclianee upon pappus char­ acters inin his treatment of subgenera PectidopsisPeClidopsis and PectothrixPeclOfhrix illustrates the problems of such an approach. After field and herbarium studies of a number of the taxa included by Fernald in subg. PeClidop~·i.~Pectidopsis and subg. Pectothrix,PeClofhrix, I began to doubt the naturalness of thesethe.~e subgl'ncra.subgenera. SC\cr.l1Several lines of evidence led me to this viewpoint. First,First. the taxa included in subg.subg. PectidopsisPee/it/opsis sensu Fernald fall into t"Otwo morphologi­ cally dissimilar groups which differ in .severalseveral features (Table 1).I).

TablcTable I.1. \lorphologiealMorphological ~rollpsgroups of PectisPeelis subg. PectidopsisJ'ectidopsis sensu Fernald

CH>\R>\CTERCHARACTER GROUP I1 GROUP II11

Positionposition of foliar inframarginal or oil glands marginal scatteredscallered apex of immature rounded or narrowly conical, capitUlacapitula truncate acuteacule 'lumbernumber of ray florets 5or85 or 8 ,5 nUmbernumber of disc florets 10-40 5-10 PUbeSCl'nCCPubescence of glandular-glandular­ Corollascorollas puberulcntpuberulent glabrous

TheseThes two groups of taxa have different geographical and eco­ceo­ IO~ical•ogical ranges as well, with the taxa of Group I1 occurring in thethe andarid and semi-arid areas of the southwestern United States and nonheron°rthern Mexico.Mexico, and the species of Group II occupying a more SOuthernsouthern distribution, extending well into the tropicstropiOl of Mexico [Vol.[Vol. 7919 34 J4 RhodoraRhodora

SUBGENUSSUBGEtWS PECTIDOPSISPECTI OOPS IS SUBGENUS PECTOTHRIXPECTOTHR\X (~: ~;~~.~~.; P. ..ANGUSTíFOLI'A"*" Foi..l,,············· -'. '" _ ...... ~ P,P. FILIPES FILl PES .•.... . P.P. PALMERIPAUtERI : P. PRINGLEIPRINGLEl GROUP I P,P PAPPOSA PAPPOS~ : P.p, ROSEI ROSEl • P, STENOPHYLLA STEtIOPHYL.1.A .: : P. RUSSYI P. P. RUSBYI ...... ,.... :.. P. TENELLA TEKELLA •.•.•.•••••.••.•••.•.•••••••• ", . " " . ..-...­ .... . ~ p,P. BERLANDIERÎ" BERLANDIEiii··· .. •••·••••· .••••••••••.•• ...... ~ P. CAPILLARISCAPILLARIS P ..•.•.•• P,• DIFFUSADIFFUSA : P,P. DICHOTOMAOJ CHDTOM GROUP II P P.• ELONGATAEL011GATA : P. FASCICULIFLORAFASCICULlFLORA •••...••••. - ...... \ p,P. UNIARI~~~~~UNIARISTATA ...... •. ······· ......

P. FASCICULAT" FASCICULATA P.p, CANESCENS P. HAENKEANAHAElIKEA.NA P. LIEBMANNLlEBKANNII I I

P. LONGIPES

P.P, AQUATICAAQUATIC"

fiFigure....e I.1. Rflall"Relationships....bJpo llmOllIamong ~species includedindudcd by FernaldHfn:llid in PectisP,'WJ sub-lUb­ amel1lgenera ~'idopJ"Pectidopsis and hnOl_i-

andand CentralCentral Amc:rica.America. Second.Second, ifif J-ernald'sFernald's taxonomyluonomy is followed,foUo\Ooeil. each of these groups of species has a counterpancounterpart in his subg. PI'('IO/h"x.Pectothrix. Thus somesome species of subg. PI'('lidoP5isPectidopsis appear to10 be more c10stlyclosely related to some species of subg. PectothrixPUIOIh,/x than they dodo toto other speciesspecies of theirtheir o\Oo'nown subgenus (Figure 1).I). 1917)1977] PeclisPectis•Keil Keil 35J5

The morphological similarity between taxa of different sub-sub­ genera reflects lhcthe biological proximity of these species. Among lhethe species of Group I (Figure 1),I), several cases of interspecific hybridi7l11ionhybridization involving taxa from bothbOlh subgenera havehaw been ob­olr served (e.g(e.g.,.. PectisPeni.'! f'Uf'f'O$Opapposa X P. ol/gwr!(olilJ.angustifolia. P. papposaPUI'flWIJ X P.p, filifWS).filipes). The hybrids are generally r:llherrather vigorous and fertileferlile and evidenC1:evidence of backerossingbackcrossing to thelhe parentparei'll species has been found.found. This hybridi7alionhybridization is an indicationindicalion of the close relation­rclalion­ ship of the parent taxa. In one case,ea~e. taxa of Fernald's different subgenera have even proved to beIloe conspecific (P. palmeripurJl/ai of subg. 1'eflol/tr;.'(;Pectothrix is the same as P.I'. rusbyirushri of subg. Pectidopsis).Pef/idvpl;S). A final line of ev"idencrevidence is the variability"'ariability of the pappus itself.ilself. In Somesome species (e.g(e.g.,.• PectisPeCl;S IlI/KIISr!ro!ia.angustifolia, P. papposa,PllppOSU. P.1'. rusbyi),rusbp1. the pappus is sometimes so variable at the population level that the reliability of even infraspecific classifications,classifications. which relyrely on pappus structure.structure, are doubtfuLdoubtful. Based upon this evidence.evidence, I have redrawn thelhe infrageneric boundaries to reflect more aet;uratel)accurately thelhe species relationships.relationships, [I have chosen to designate the newly defined infragcnericinfrageneric taxon for which the species of Group I (Fi,2ure(Figure 1)/) form a nucleus, as a ~ctionsection rather lhanthan as a subgenus. In rejectingreJccting the subgenus inin favor of the ~ction.section, I am following the lead of Gray (1884) who.... ho wrOlewrote concerning Pet:I;S:Pectis: 'The"The genus is very natural; and, althoughallhough two or lhrrethree genera have been made from it,il. itit seems incapableincapable of°f division even into well-marked subgenera."subgenera.~ In my opinion, the degree of differentialiondifferentiation among the species of PectisPet:lis isis notnOI sufficientsufficient to justify lhethe recognition of subgenera. The name.name, Pct:lidopsis,Pectidopsis, cannOIcannot be used for the newly defined section, because the provisions of Article 21 of thelhe InternationalImernOlional Code of Botanical NomenclatureNomendOlUre (Slafleu.(Stafleu, 1972) prohibitprohibil itsits use alat eilhereither the sectional or subgeneric level within Pectis.P('("/is. However,Howewr, oneone of the twolwo original species of sect. PectothrixPet:lolhr;x (for which no ~ypetype species was ever designated).designated), P. papposa,papposo. is now included Inm lhethe section as I have defined it. I have designated P. papposaPOppOSIl as the lectotype species of sect.~ct. Pectothrix,Pectothr;x, which is thus adopted as thethe name for the newly defined section. To the species of Group I (Figure I),1), I hav'ehave added se\'eralseveral taxa described since F~rnaldFernald (1897) published his revision. This group of taxa con­ Stitutesstitutes ~el.sect. PectothrixPecto/hrix as I am treating it. 3636 RhodoraRhodora [Vol.[\,,~ 79N

TheThe taxataxa which... hieh arearc includedincluded inin GroupGroup IIII (Figure(Figure 1)I) constituteconstitute aa distinct sectionsection notnOl atat allall closely relatedrelated to10 theI~ speciesspecies ofof thethe redefinedrcddined sect.set!. Pectothrix.Putothrix. ThisThis section,section. forfor whichI'ohich nono namenarm isis knownkno.....n toto bebe available,3\"ailablc:. will bebe thethe subjectsubject ofof aa future(uture report.report. TheThe remainingremaining speciessptties whichwhich werewere includedincluded inin subg.subg. PectothrixP~IOfhri:c byby FernaldFernald are excluded fromfrom thetbe presentprescnt treatment.!rcatmcm. OneOne ofof these,these. PectisPutis aquatica,aquotic-a. isis betterbettcr treatedtreated asas aa membermember of aa differentdifferent genus,genus. Hydropectis.Hl"d'o~C'lis. The remainingremaining species will.... ill bebe thethe subjectsubj~t of furtherfurther investigations.;n~·estigations. PectisP«tlS JOKiwlota.fasciculata, thethe remainingremaining speciesspecies inin subg.subg. Pectidopsis,Ptctidopsjs. is a doubtful species of uncertain affinityamnii)' forfor whichIlohich II haveha.. /: beenbeen unableunable: toto locatelocale a type.type. UnlessUnless or until aa typet)pe isis locatedlocatC'd forfor thisthis species,sp«ies. IJ will",ill excludee:o.:clude itit from furtherfurther consideration.

TAXONOM1CTA.l(O,"O\ltC TREATMENTTREAl'.l['.1

PectisPcclis L. sect. PectothrixPcctothrh: (A. Gray) A. Gray, Smithsonian Contr.Conlr. Know. 3:J: 83. 1852.18S2. LECTOTYPE LF(""rorYPE SPECIES (here designated): PectisP~('liJ papposapappQJa Harv. lIarv. & Gray inin A. Gray.

PectisP",·,i, .ubg.subg. PectothrixP....""h'i.T A.,\. Gray,Gray. Mem.\tern. Am. Acad."~ad. 4: 62.62, 1849.11\49. Helioreos1/~'HJr;n .• Raf.,ltar.. Atlantic Jour.Jnur. 1:I: 145.143. 1832. TYPET.~, SPECIES:'~"'II~: Pectisr(rti, angustifolial~"l:lIm(nt,a forrey. orrey. PectidopsisPtrt"Jop"" DC.DC.. Prodr. 5:S: 98. 1836.1&.16. nom.nom illeg.illel. (superfluoushupc:rfluou. when",ben published).pubh~l>c:dl. PectisPH'''' sect.~. PectidopsisP«,"I"f'SiS (DC.) A. Gray,Gray. SmithsonianSm'lhloub,.subg. 1'r'

pubescent. pubescent, black. Pappus coroniform and/or setose or ariSlate.aristate, thethe ra)'ray pappus often differing from the disc pappus. Chromosome base number: xJV: = 12.

DtSTRIHllllo,:DISTRIBUTION: California and Nebraska south to10 thelhe Mexican statesstates ofof Sinaloa.Sinaloa, Aguascalientes.Aguascalientes, and San Luis Potosí;POlosi; one species disjunct in in Peru. Locally abundant throughout muchmilch of thelhe arid and semi-arid porlionportion of North America from sea~ea levelIc\cllO to 2000 m.

Kr.YKEY TO TIlETHE TAXA OfOF PEcnsPECTIS SECnOSECTION..... I'ECTOTHRIXPECTOTHRIX

I. 1 • PltyllariesPhyllaries and rays 8 (rart'ly(rarely 7.7, 9 or to).10) 2. 2. 2. l,caYe,Leaves pinnalifid.pinnatifid . . 7,7. P. incisifolia.,nmifoiJo. 2. 2. lea,csLeaves entir.entire or ncarlynearly so.so ...... 3.3, 3. lolJoreSl:.l>Ce Inflorescence rat~crrather C(m8esled;congested; peduncles mostlymoslly less thanl/.an 15 mm long.lool!...... 4.4. 4. Pappus brislle.bristles (if(if prescnl)present) .ub·plum05C;sub-plumose; trichomestrichom•• on achenesachencs with",ilh curled bUlbousbulbous lips;tips; leaf 1>.0"",bases ""Inot flaredflart'd, 5. 5. 5. PhyllariesPhyllari." 3-5 mm 1001;long; disc floretsfIor.ls 66-14; 14: disc pappuspappu, 1.5-2.5 mm long.long ...... 4a4a. P. papposapappOlO var.,·ar. papposa.pappow 5. Phyllarics Phyllaries 5 8 mm long; disc floretsflow, 12-24 (-34); disc pappus 2.52.5-4 4 mm loog.long .. 4b. P. papposapappOJo var.'..r. grandis.Il:'ondis, 4. Pappu.Pappus bristles (If(if pres.nllpresent) ShoMshort scabrid; Irichome.trichomes on achenesachen.s with....ith .....straight,igltl. bifurcal.,;p5.bifurcate tips ...... • 6.6. 6. 6. Ila"",Bases of upp.rupper I.a,'.sleaves conspicuou.lyconspicuously flared,f{aICd. on.noften partiallyparllall)' COlIQal_conceal­ mging I~ethe peduod.,,;peduncles; pappus usually only coroniformcoromform (weakly aris­ari,_ lal.tate Oror f."'....,,"'"few-setose in ""mesome T.xanTexan or ChihuahuanChillualluan populations); plantsplanl, usually l.mon_sccnle.1'i. 38 38 Rhodora Rhodora [Vol.[VoI,79 79

8. 8. Ph~lIaficsPhyllaries ~il~reither nOlnot broadly obtuobtuse•• 0'or not longitudinallylongiludinall) striate,trialo...... 9. 9. 9. Pappus o(of tsachenes ,hofler.shorter 10. 10. Ray pal'pulcoroniformpappus coroniform or multisetose. muhisclosC. 11.II. 11. 11. RayKay pappu,pappus coton,fo,m.coroniform 12. 12. 12. DiscDi", pappuspappu, usuallyusuaU} setose;lale. obtuse,Oblll"':. mostlymoslly 66-1212 mm broad;blood: enodem;cendemic to10 coaslalcoastal areasarras of southernsOlllllern Sinaloa.Sinaloa ...... 9. P.1'. sinaloensis.,inalUf'n.'i•. 13, 13. Leaves linear 10to narrowlynam)"ly ellipticelliplic or ohlanceolale.oblanceolate, mostlymosll)' aeuleacute 10to acutish,acllli.lI. usually less thanlha n I>6 mm wide"ide. 14­14. 14. Ray corollas 77-12 12 mm long;10nB: pappu,pappus biseriatehi,.riale. •... 15.IS. 15.IS. OUlerOuter pappu.pappus of disc achene.achenes of slender.lender bristles;hri,ll..: lea'·leaves... Wilhwith 11-3 3 pairs of marginal cilia;ciha; foliar oil glands~!and. in submarginal rowsro"'., ..... "•...... • Sb.8b. P. purpureap"'p",~a'ar, var. sonorae.Wtk"IN. 15. IS. OUlerOuter pappuspappu. of flattened.flattened, fimbriatefimhriale pales;pales: leaveslea,'es usually "'lhwith 44-6 (, pairs of marginal cilia; foliarfohar oil glandsg\;loos submarginalsuhmarginal and ..::aHeredscattered on thelhe under-unde,­ .unace.surface 8c. P.1'. purpureapU'f""{,(} 'M.var. land,ifolia.lann/piia. lao 14. II.I)'corollas2Ray corollas 2 6 mm longlong. '6.16, 16. 16. Planuperrnnialwilhawff"ood)caude~. perennial with a stiff woody caudex. ... 17­11. 17.11erbaceglahrou,or""arlyso17. Herbage glabrous or nearly so , ...... • la. P.Pj/hwf'iidla,ar.5/en0l'iidIQ. stenophylla var. stenophvlla. 17.17. llerhagcHerbage densel)'densely hirtelloushillelloll•. lb. P.P.. stenophylla'/mopii.-/la var.,ca •. gentryi.gmlrl", 16. PlantsPlom, annual ..without ilhoUI a woody"ood}' caudexcaudex. '$•IS. IS,18. Heads moMlymostly b'''''dl}broadly eampanulale.campanulate, 17-2617 26 flowered . .. , ... , .....le. Ie. P. P. stenophylla

I.1. Ray 1l0rclSflorets 5 (rarcly4(rarely 4 oT6). or 6) 20. 20. I'h}1LancsPhyllaries J.S3.5-6 6 mm loog;long; dIscdisc corollas 2.5-4 mm long; pappuspappu. 00-3 3 aristate.anSlale...... •...... •• ..21.21. 21. Fa,ciclesFascicles of .mallsmall leavesIea ...,s usually presentptc~nl injn thethe upper leaf axils;a~ib: pedurn:lc.peduncles mo.llymostly 1010-25 25 mm long;lonll: disc floretsOo",u 22-8; 8: plantsplanlS of c<:nlralcentral and southemsouthern Sonora.Sonora 10a.lOa. P.P.filipt filipes.• "aT.filiptvar. filipes.•. 21. Fascicles of ka,""sleaves usuallyusuall)' ababsent;..nt; pedunclespedunc..... mostlymoMI} 20-50 mm long;loog: dISCdisc floretsnoret. 77-22; 22: plantsplann ..-ilhwith morCmore northernnurlhern or eastern dislribUliondistribution...... ,.... 10b.lOb. P. filipes(i/ipt., ".r,var. subnuda.J"hm"'D. 20. I'h}lIariesPhyllaries 6 8 mm 101lg:long; dISCdisc (orollascorollas 44-55 mm long;lung: pappuspappu, 2-62 6 aristate;.ri.tale; plamsplants of BajaRaja CalifornIa.California ,. 11. II. P.I'. '·011",,,,;.vollmeri.

I.1. PectisPutis stenoph}·llastenophylla A. Gray, Proc.Proe. Am. Acad. 21: 393. 1886. TYPE: Mnieo.Mexico. CIlIHUAllUA:CHIHUAHUA: near Batopilas,Balopilas. PalmerParmer 81 (11010­(Holo- type. type, Gil!:GH!; isotypes, K!,K'. I'll!).PH!). Figure 2. Annual or perennial herbs from a slender taproot or woody caudell.caudex. Stems JO10-30 30 emcm long, one to many from the base, erect or decumbent.decumbent, glabrous 10to densely hirtellous, ciliate with 1-3I 3 pairs of basal bristles 1-3 mm long, punctate marginally withwilh golden-brown oil glands or.or, in some forms.forms, punctatepunctale submar­ ginally on the undersurface with tiny black glands 0.1-0.2 mm diameter. Peduncles filiform.filiform, 2.5-72.5 7 erncm long,tong. glabrous or hir­ tellous:tellous; bracteolcsbracteoles 3-8, linear to lanceolate,lanceolatc, glabrous or hirtel­ lous.lous, sometimes punctate. Heads terminal or axillary;axillary: involucres campanulate. Phyllaries linear 10to oblong, 3.5-5.5 mm long, 0.5-20-5-2 mm broad,broad. obtuse to acutish,aeutish. basally rounded and gibbous,gibbous. slender-keeled nead)'nearly to the apex,apex. glabrous to hirtellous,hirtellous. punctate with I1 or 2 Subterminalsubterminal and 2-3 pairs of slender submarginal glands, these either brown and swollen or tiny and black. Ray Il.orets florets 8; corollas 2-6 mm long, yellow or becoming tinged red­ dish.dish, with ligules glabrous and tubes glandular-pubcrulcnt.glandular-puberulent. Disc 110retsflorets 8-26.8-26, bilabiate.bilabiate, 1.7-3 mm 1011g.long, glandular-puberulent.glandular-puberulcnt. A.chenesAchenes 1-2.5 mm long, strigillose. Pappus ofo( the ray achenes btarislale,biaristate, 1-2 mm long:long; pappus of the disc achenesachcnes variable, setose, aristate, and/or eoroniform.coroniform, 1-2I 2 mm long. Chromosome number (from vaLvar. biari.flarabiarisiata onty):only); n/I =- 12.

DISTRIRUTlO-'::DISTRIBUTION: Eastern Sonora and sombwesternsouthwestern Chihuahua to Centralcentral Sinaloa and western Durango,Durango. from 300 to 1500 meters elevation. Very local in thorn forest and oak forest zones lones (Fig­ ure 3). 4040 RhodoraRhodora [Vol[Vol. 7979 ~ I\ { I

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Ii ,i I ~? ~'S IT I , . ',I a ~dD k %m

• i ( 'Its "e

,:~I\ j I fill." ,I • • \. ." , !; t 'A Ct H \~I ".'.1 r;N o w•',P

Figurel'~rc 2.2. A-P,A-I'. PeáishUll stenophyila;.,....,oph'ff"" A-C..... -e. var.'ar biahstatato"'''III'. (KeilrAnI && CanneC...... 8752,~" .. OS);0-): D-F,D-F, var.,-a' gentryi~Ir" (Gentry((;....,.. 7323,"J.'J. MICH);"IO;'H); G-J,G-J. var.>:,,, puberula"..lvrul" (GentrylOrn,.. 5495.'4~J. NY);"k K.-M,It.\!. var.'ar. roseirov' (Palmerf f'"tmn 730.'JO. GH);G"I: N-P,\'.1', var.u, stenophyila,,",,,,,,,"1<1 (Kimnachf """.....-It &di BrandtB,,,,tJr 950)\9~ Q-T,Q-T. Pectis1'rr1lJ rusbyi.""10,, A,A. D,D. H,II. K,K. N,\.. R,k. phyllaries;ph)lLlIrlt'; B,8. E.E. I,I, L.l. O,O. S,S. rayfa) achenes:uti","; C. F,F. J.J. M,\1. P,p. T,T. discd"'" achenes;.chcnn; G,G. Q,Q. habit."""". 19771I9?7] Pectis•Pectis-KeilKeil 41

PenisPectis s/el/opnl'ffastenophylla is an unusually vilTiablevariable species.spel:ies. Local races,races, isOlatedisolated from other populations,populations. often by considerable distances,dbtanl:cs, h.avehave undergone marked divergence. In /9161916 Rydberg recog­recog· nized a total of five species from this complex,complex. a logical treat­treat­ ment based upon the limited samples then available. Each taxon,taxon, h.owever.however, was known from only one or two collections.collections, In the intervening period since Rydberg's treatment, a num­ ber of wllectionscollections from additional populations have been made. Members of several of thesethesc populations have character com-l:om­ binations intermediateintermediate to those of specicsspecies accepted by Rydberg.Rydbcrg. Because of this obServedobserved intermediacy.intermediacy, I1 havehav~ chosen to regard all members of this complex as one polymorphic species and to recognilerecognize the variant forms atOIl the varietal level.levcl. The species thus defined is comparable 10to olherother variable specicsspecies within sect.sect, PeClo/h,ix.Pectothrix. Some Objectionobjection might be raised to the inclusion of the peren­peren· nial PeNisPectis slt'//ophylla,stenophylla, sensu stricto, in the samcsame species with severalseveral annual forms. There is, however, precedentprecedcnt and ample justification for such a treatment. Both!loth perennial and annual forms exist in another species of the section, P. anglls/ijoJia,angustifolia, and a tcndcncytendency toward the perennial habit is found in some annual species of other sections (e.g. some populations of P. prostratapmSlralO of sectsect. Penis).Pectis). In one population of P. s/enophylfastenophylla var.\ar. puberula (Gemry(Gentry 5282), annuals and apparently perennial forms occurred together, thus indicating theIhe plasticity of the habit in thesethese plants. 011eOne feature of particular interest in PectisPertis s/<'IlOphyflastenophylla is thethe \'esligialvestigial nature of the oil glands in some local races. Collectors' notes 011on herbarium herbarium labels indicate that the oils of plantsplanlS with s\>,ollenswollen brown glands are are quite aromatic. In[n some populations,populations, however, the glands are small.small, dark brown or black,black. and shrunk-shrunk· e.n.en, and apparently lack any det~ctabledetectable odor (personal observa­ tion).ron). The odorless glands are apparently displaced from thethc leafleaf margin during ontogcny.ontogeny. This relationshiprclationship is quite apparent fr fro~°m an examination of theIhe type collection of P.f'. scabra,scahra. inin \>,hlehwhich plants wilhwith either one or the othcrother typetr~ of oil glands co­ e.xistcdexisted in the same population. Those foliar oil glands thatthat are hnytlny and black arcare inframarginal on the undersurface of thetbe leaves.'eaves. Because this variation can occur within a population, I[ co.nsiderconsider Ihethe taxonomic significance of this charactercharncter toto be of mmmi . noror Importanceimportance in P.P, slel/oplz.rlla.stenophylla. 4242 RhodoraRhodora [Vol.[Vol. 7919

la.1a. PectisI'eclh stenophyllasltnoph}lIa A.A. Gray,Ora)'. Proc.Proc. Am.Am. Acad.Acad. 21:21: 393.393. 1886.1886. var.'oar. stenophylla.slcnoph}lla. FigureFigure 2.2. PlantsPlants perennial.perennial. Stems 15-2515-25 cmem long, long. woody woody andand rigidrigid atal thethe base,bast. glabrousglabrous or nearlynearly so.so. Leavesua\'es linear,linear. 1-3I 3 cmem long,long. 1-21-2 mm wide,""ide. glabrous,glabrous. marginallymarginally punctatepunClale with.... ilh golden-browngolden-brown oiloil glands. Peduncles 2.5-52.s~S cmem long, long. glabrous. PhyllariesPhyllanes 4-4.54-4.5 mmmm long,long. 1I mmmm wide, punctatepunclatc withwith aa conspicuously enlargedenlarged sub- sub­ terminalterminal oil gland and smaller submarginalsubmargmal glands. RayRay corollascorollu 44-6 6 mm long. Pappus of thethe rays 1.5-2 mmmm long;long; pappuspappus ofof thelhe disc 1010-20 20 SC'IOse.setose, 1-2 mm long.long. Chromosome numbernumbe'r unknown.unknown. FloweringAO'Acring AugustAugusl to10 April. DISTRIBUTION:DISTRI8UTIO,\ EasternEaslern Sonora andand southwesternsouthwestern Chihuahua atal 600 to10 1500ISQO meters elevationdc\'alion (Figure 3).3). InIn some areas thethe local residents boilboll thethe herbageherbage of PectisPl!rtis -ffefmphyflastenophylla and useusc thethe decoction as aa medicine (Gentry,(Gentry. 1942). SineeSince the plants of var. stenophyllaJ/e/lophyllu are notnot very common (as(as evidenced by thethc paucity of collections),eollei:tions). such usage,usage. ifif veryvcry common,common. must put a considerable strainSlrain on thethe populationspopulations of this taxon. InIn species with large population systems,systems. locallocal har­har­ vesting is probably of little overall significance. ForFor a taxonlaxon as uncommon as var. stenophylla,J/enophrlla, however,howe\'cr, man's influenceinnuence may have profound effects.effects. 2 REPRESENTATIVERr~u.sr\UT1n SPECIMENS'~f(l,,[\~l,: México, "I"ico. CHIHUAHUA:("IlIM ...."\ .. Rio Batopilas,BloIOI"Lo~ KimnachKimnt')bo.Guagueybo, PenningtonP",n/J1X"'" 424! (TEX),IIl\) SONORA:'O'

lb. Pectis Ptclk slmoph}lIastenophylla A. Gray var. gentryiJtfllf)i Keil,Kcil. var. nov.no\'. Figure 2.1. TVPETYPE: :\le.\Ko.Mexico. 'i1:"AlOA.SINALOA: VaromenaVaromcna and vicinity,\kinity. GentryGm"r 7)157325 (Holol)pc.(Holotype, Gil';GH!; isol)pes.isotypes, F!.f!. MICH!,'liCit'. NY!).'V'). Plantae perennes. Caules 15 15-30 30 cm longi.longi, basi lignosi\ignOS! rigides-rigidcs< que.que, dense hinelli.hirtelli. Folia Iinearialinearia vel\~I anguste elliptica.elliptica, 1-2 cm

2 'V.In,Unless •a 1',0"taxon Itis knoknown..n fromfrom ''nJvery rfew.... collcctlOftl,collections, peamespecimens... p",paJ"Odprepared dun""during lhethe present lUx!)study: KA -~ Keil,Ant AK ..& MI( z- AtliKeil '"& MeGill,IkGiII, and K '"& C ~- A~i1Keil '"& CD"""Canne. BecauseIkcaU1t duplOIe1duplicates of >Ornesome ofof thethe col,",t"''''collections ma.xmade durin,during 1Mthe prepar.nonpreparation of thisIhlS "'udy:lrestudy are ...still,11 beingbe,n, dislnb\llcd.distributed, onlyonly lhethe Iw'banumherbarium ofof Ihethe OlnoOhio 'i1l10State Vn,~nilyUniversity (OiS)(os) i.is listed for Iho~those coltoel'O",collections. 1977] Pectis•Keil 43

'gure 3. Geographical distribution of Pectis stenophyUa and P. rusbyi. Half­ arred hne indicates Sonoran Desert boundary (redrawn from Shreve, 1942).

0n ga, 1-3 mm jata^ dense hirtella, margine punctata. Pedun­ p!|! 3­ 5 Cm longi' mrtem' vel glabrescentes, 3-5 bracteolati. yllaria 4-4.5 mm longa, dense hirtella. Corollae flosculorum radiorum 4 mm longae. Flosculi disci 8-10, corollis 2.5-2.7 mm longis. Achenia 2.5 mm longa. Pappus radiorum biarista­ s> 1.5 mm longus; pappus discorum circa 20-setosus, 1-2 mm ngus. Chromosomatum numerus ignotus. , ISTR!BUTiON: Known only from the type locality in nor- ern Sinaloa at 300 meters elevation (Figure 3). entry's collection is from the Sierra Surotato, a mountain SS 1SoI Q^ ated from the main cordillera of the Sierra Madre cddental by a distance of about 40 miles (Gentry, 1946). This d,ffers de from the more northern var. stenophyUa in being In %tfU- . Scaberulous rather than almost completely glabrous. a dltion, the leaves appear to be mostly shorter and thicker r- gentryi than in var. stenophyUa. 4444 RhodoraRhodora [Vol.['101.19 79

1c.Ie. Pectis1'«115 stenophyllasl~nophllla A.A. GrayGray var.var. roseirDSti (Fern.)(Fern) Keil,Keil, comb,comb. nov.no\'. BASIONYM:RASIO'YM: PectisP«tis roseiroyi Fern.,Fern., Proc.Proc. Am.Am. Acad.Acad. 33:]]: 77. 1897. TYPE:TYPE: Mexico.l\Iuico. SONORA:sm.ORA: Alamos,Alamos. PalmerPalmer 730730 (Holotype,(llo1otype. GH!;Gil'; isotype,isol)'J}t. US!;US': photophoto ofof USus isotype,isOI)'pt. NY!).."Y!). FigureFigure 22 Plants annual. Stems 20-302Q-30 cmem long,long. glabrous. Leaves~a\"eS linear,linear. 2-3 cmem long,long. 1I mmmm wide,""ide. stronglystrongly revolute,re\olute. glabrous,glabrous, punctatepunctate on thethe undersurface withwilh tinytiny blackblack glands.glands. PedunclesPeduncles 3-5]-5 cmem long,long. glabrous.g\.abrous. HeadsHeads cylindric.C)lindric. PhyllariesPhyllanes very,"cry narrowlynarrowly linear,linear. 3.5-43.5 4 mm long,long. ca 0.5 mm broad,broad. punctatepunctate: withwilh linearlinear red-blackred-blat" glands. Ray corollas 3 mm long.long. DiscDisc floretsOOrtl.5 11-12;11-12; corollascorollas 2 mm long. AchenesAchcnes 2 mm long.long. Pappus of thethe raysrays 1-1.51-1.5 mmmm long;long: pappus of thethe: disc 2-32 3 aristate,arist:lIt. caea 1.5 mmmm long.long. Chromo­Chromo­ somesome: number unknown. DISTRIBUTION:DI<;TRlllUTIO';: Known only from thethe: typeI)'PC localitylocality inin southernsouthern Sonora at ca. 1500 meters elevation (Figure(Figure 3). PectisPee/is stenophyllas/enopJryl/a var. rosei isis perhaps most closely relatedrclat('(! to10 var.\ar. stenophylla,slenophrlfa. which itit resembles inin being glabrous andand inin having very\'ery narrow leaves. The twotwo differ inin habit,habit, foliarfoliar oiloil glands, and pappus structure. Since var. roseirose; isis stillstill knownknown only from itsits type collection, the fullfull range of variation remainsremains unknown.

Id,Id. PectisPeclis slenophyllastenophylla A. Gray var.\ar. puberulapubtrula (Greenm.)(Grttnm.) Keil,Ktil. comb. nov.no\". BASIONYM.BASIOSYM: PectisPeNis puberula Greenm., Proc. Am.Am Acad. 40: 49. 1904. TlTYPE:PE Mexico,Muteo, SINALOA?SISALOA~ Lodiego,Lodiego. PalmerPalmu 1605J6()j (Synt}"J't$,(Syntypes, Gil',GH!, US!;US'; isosyntypes,isosyntypes. F!,"', GH!.Gil', NY!;...,!; lectotypelectotype: (here chosen).chosen), syntype at Gil).GH). Figure 2.

,,",i>Pectis """'Qscabra 8nndcJ.Brandeg.. Zoe S:5: 2..'6.226. 111061906. -h",TYPE: Mexico,\luiro. SINALOA:~r"...to" Cofradía.Cofradia. ,.~Brandegee J.II.s.n. (HoIotH~.(Holotype, te'.UC!; isotypes,~H".·" GN'GH!, O'(N'POM!. 1'S').US!). PlanaPlants annual.annual, or sometirnt$sometimes persistingpersistlflg more than one grow-go'" inging season. SiemsStems 10-30 cm long.long, sparsely to densely hirtellous.hirtelloll'i· Lt:a\'C'SLeaves hnearlinear to narrowlynarro~ly elliptic.elliptic, 1-4 cm long, 1-5 mm wide,~ide, denselydensely hirtellous.hirtellous, punctate tithereither marginally with golden-browngolden-bro,,-n glands oror on the ahaxialabaxial surfacesurface ~with ith tiny black glands. PedunclesPeduncleS )3-6 6 cmcm long.long, glabrous to pubctulenl,puberulent, )3-6 6 bracteolate. Heads cylindriccylindric to narrowly campanulalt,campanulate. Phyllaries 3-4.5 mm long,long. glabrousglabrous to denselydensely hirtellous, punctate with either brown or blackblae!. 1977]1977] Pectis•KeilPectis-Keil 45

glands. Ray corollas narrow, 2-3.7 mm long. Disc florets 8-17; corollas 1.7-3 mm long. ÀchenesAchencs 1-2.5 mm long. Pappus of the raysrays 1-1.5 mm long; pappus of thelhe disc coroniform, 2-4 aristatearislate or 10-20 setose, highly variable,variable. 1-2 mm long. Chromosome number unknown. Flowering October to March.

DISTRllIUTlOS:DISTRIBUTION: Central SinaloaSinaioa and western Durango atal 200 10to 900 meters elevation (Figure 3). The pappus of var. puberulapuberu/a is highly variable. Within popu­ lations from bOlhboth Ihethe typelype locality of PectisPutis puberula {Palmer(Palmer 1605)¡605) and from the type locality of P. scabra (Gentr)'(Gentry 5495),5495). a full rangerange from coroniform to short arislatearistate 10to setose pappus formsforms OCCurs.occurs. PeaisPectis scabra and P. puberula puberu/a were described from dif­ fcrelltferent pappus forms. Although the exaclexact localitylocalily of Lodiego,Lodiego, the type localilylocality of P. puberula,pl/bem/a, has been 10Sllost (McVaugh, 1956),1956). and it is notnOl certain whether the silesite is in eaSlerneastern Sinaloa or westernweslern Durango, lherethere is no doubldoubt that itil is only a few miles fromfrom Co­ fradia.fradía, Sinaloa, the type locality of P. scabra.

R'PRESr-q"'T!vfREPRESENTATIVE ~P[C1"1l"'SSPECIMENS: Mhiro.México, DtR""GODURANGO: Sierra TTC'Tres Picos,Pic,,", Grm,]Gentry 5282518J ('.'l.(ARiz. rx.~.Ds, F. GilGH, '-l0.MO. 'Y,NY, I'C).uc). "SINALOA: ...... ,0... LasLa. Mllpas,Milpas, Gmlfl'Gentry 5J6J5361 (DS.(lY<, (;II,GH. MO):MO); Colradia.Cofradía, Gr,myGentry 5~Qj5495 ("'klf.(ARIZ. u~,DS. J.F. Gil.GH, MICH,"IICH. MO,"10, NY,,yo t'e.VC, ts):US); Badiraguato,BadirallualO, GentryGrnlrl' 57815781 ("'.'l.(ARIZ, os.DS, IlII,GH. MICII.MICH, MO. NY).H). Ie.]e. PectisP~lis stenophylla A. Gray var. biaristatabiuislata (Rydb.) Keil,Keil. comb. nov,nov. BASIONYM: Pectisfpctis hiaris/(llabiaristata Rydb., N. Am. F1.34: 211.2/ r. 1916. TypF.:TYPE: "Juleo,Mexico, SINALOA: vicinity of Fuerte,Fucrte. Rose,Rose. S/andlel'Standley & Russell 13537JJ5J7(Holotype, (Hololypc, NY!;~y!; isotype,iSOl}·pe. trd).uc!). Figure 2. Plants annual. Stems 5-25 emcm long, sparsely to densely hirtel-hirtel­ lous.0US- Leaves linearlinear to narrowly elliptic.elliptic, J-31-3 emcm long, 1-3 mm wide, nearly glabrous to densely hirtellous, punctatepunclale marginally with\lith brown oil glands or SUbmarginally submarginally on thelhe abaxial surface with blackpack glands. Peduncles 3-5 emcm long, glabrous to densely hirtel­hirtel­ lous.0us. Heads broadly campanulale.campanulate. Phyllaries linear to oblance­ olate.olate, 4~54-5 mm long, 1-21~2 mm broad,broad. glabrous or hirtellous,hirtellous. punctate ~IlhJWh Smallsmall dark glands. Ray corollas 4.5-54.5 5 mm long. Disc corol­ *s 17-26.,7~26> ca 3 mm long. Achenes 2.3-2.5 mm long. Pappus of ~~ne rays 1.7-1.8 mm long; pappus of the disedisc 1-3 aristatearistale or 10-2010.20 S*etos ose,e, 1-2 mm long. Chromosome number: nII == 12. FloweringFIO\lertng Septembercplember 10to March.

DI DISn,IRUTlO,:STRIBUTION: SoulhernSouthern Sonora (1)(?) 10to central Sinaloa (Figure 3).3), 4646 RhodoraRhodora [(\'01Vo1­79 7'J

A collectionrolkclion fromfrom SonoraSonora (Sikes(Sikn && BabcockBahcock 178)178) isis referredreferred with..... Ith some doubt to10 var.\1Ir. biaristata.biariSIOla. ItIt appearsappears toto bebe aa mixedmilled collectioncollection includingincluding some putativeputative hybrids hybrlds withwith anotheranother undeterminedundetermined spe­spe­ cies. The plantsplants resembleresemble var.\'IlT. biaristata,bioris/%, butbut becausebceause ofof thedK de-de­ pauperatepauperate naturenature of thethe specimens,specimens. thethe determinationdetermination isis mademad~ withII> ilh some reservations.rnen:alions.

REPRESENTATIVER'!1l'''''.'''' SPECIMENS:""<1",,,,; México,Mhko. SINALOA:"~"lJ..' WW ofof Pericos,~~ GentryGnfU" 5751J7jJ (GH,{<--, MICH,..Ie" MO);\10); 16.816.1 mi.i SWSilo' of ElII Fuerte,Fuone. K &'" C 8752IiSlIO'<) (os).

2.2. PectisPtttis rusbyirusbli GreeneGreen!" exell A. Gray,Gra)'. Syn. Fl.FI. N.No Am.Am. 1(2):1(2): 361.36J. 1884. TYPE: Unitedlnittd States.SIIIU. ARIZONA:.1\.11.170''': YavapaiYavapai Co.,Co., BeaverBea,"cr Creek.Creek, RusbyRushy 317 (= s.n.HI. or 655) (Holotype,(HololypC. GH!;GH!: isotypes,isolYpes. F!,fl. K!,It!, MICH!,\lICU!. PH!,PH!. RSA!,RS.... !. UC!,uc!. US!).L's!). FigureFigur"t 2.2.

PectisI't'rliJ palmeriptJlmr.; S. Wats.,Wals.., Proc.P'O(. Am. Acad.And. 24:U, 58.sa. 1889. TYPE:T,"" Mexico,""ui, TYPE:Tn'" UnitedI nlltd States.SI.'ts. ARIZONA:UI/O" YavapaiYa\llpai CoCo.,.. FortFori Verde,\'trdt. Mearns,lftarttJ 184IIU (Holotype,(Ilolol)"~. NY!).'y!l. Strong-scented tap-rooted annuals,annuals. erect or branching fromfroll1 thethe base. StemsSIems 5-50 cm long, erect or decumbent,decumbent. often dichot-dichot­ omously branched,branched. glabrous or sparsely puberulent on thethe angles.angles. Leaves linear to narrowly elliptic,elliptic. 1-5 cm long,long. 1-5 mmmm broad,brood. often revolute,revolute. ciliateciliale with 1-3 pairs of basal bristlesbristks 121-2 mm long,long. otherwise entire.entire, glabrous or glabrescent,g13breslXnt. marginally punctatepunClate with rounded oil glands O.2..().70.2-0.7 mm diameter. Pedunclesredunc1es 2-8 cm long.long, glabrous. glabrous, "'ithwith 3-6 lance-attenuatelanlX-allenuate or setiform bracteolesbraeteoles 1-3 mm long. Heads in forks of stems and axillary,altillary. becomingbecOmHl' cro",dedcrowded in ag<'".age. PhyllariesPh}'llaries 8.8, oblong or narrowly obovate,obovate. 4-7 mmmlfl long.long, I1-2 2 mm broad,broad. "cryvery obtuse to10 subacute,subacule. longitudinally10ngilUdinally striate wilhwith 3-53 5 usually conspicuous nerves on each side of thethe conspicu­conspicU­ ous.ous, corky_corky, basallyba~lIy gibbous midrib,midrib. hyaline margined,margined. apicallyapJcaIl) olio13te.ciliolate, otherwise glabrous, punctate "'ithwith 0-2 subterminalsublerminal oil glands and 2-4 pairs of inconspicuous.inconspicuous, rounded to narrow-elon­narro",-clolt" gated submarginalsubmarginal glands. Ray floretsnorets 8; corollas 5-115-J I mm long,Ion,. with brOadl}broadly ovate glabrous ligutesligules and slender.slender, glandular-pu­g1andular-pu' berolentberulent lUbes.tubes. Disc florets (1-)(7-) 2020-55;55; corollas 3.5-5 mmrom long,Ion" bilabialebilabiate with the narrownarro'" lobe ca. twicel"'ice the length length of the teethleeth on the broad lobe.lobe, glandular-puberulent on the tubelube below.belo"'· Achenes 3-4.5 mm long.long, slrigillosestrigillose or pilosulose. Pappus of the ray achenesachenes 0-4 anstatearistate andand/oror coroniform; awns slender.slender, 1-4 mmmil! 1977)1977] Pectis•KeilPectis-Keil 47 long.long, antrorsely antrorsely barbed towards the tips. Pappus of thethe disc aehenesachenes eoroniformcoroniform or 15-30 setose; bristles 2.5-5 mmmOl long,long. antrorsely barbed. Chromosome number: n = 12. FloweringFlowering August to February. February.

DISTRIKUTlOS;DISTRIBUTION: Central Arizona to sOUlhernsouthern Baja California and northern Sinaloa at elevations of 0-1200 meters (Figure 3). Pec/isPectis rwbyirusbyi has a mostly Sonoran Desert distribution.distribution, barely extending beyond the deserldesert into the grassland in central Arizona and occurring sporadically in the thorn forest of southern Sonora and northern Sinaloa. In central Arizona,Ari7ona, PectisPec/is rusbyi exhibits a striking dimorphism of pappus structure. The type of P.1'. rusb.rirusbyi is essentially epappose,epappose. Whereaswhereas the type specimen of P.P. mearnsii, collected only a fewfew miles from that of P. rushyi,rusbyi, has a setose pappus. However, Hown-er. based upon personal field observations, both pappus forms may coexist inin the same population.popUlation. For example, in both Maricopa and Ya\'apaiYavapai Counties of Arizona.Arizona, some individuals were found with a setose pappus and numerous others with only a coroniform pappusPappus (Keil{Keil 8569, 8575). As a result of these observations.observations, II regard both P. rusbyirusbl'i and P. mearmiimearnsii as eonspecilicconspecific.. .Kearney and Peebles (1942) placed PectisPee/is mearnsii inm synonymy wlIllwith P. palmeri,palmeri. a decision with which I heartily concur. The two taxa are essentially indistinguishable. However,However. with the present e\'idenceevidence of the conspecificity of P. mearnsii and P.1'. rusbyi,rusbyi. P. Palmeripalmeri must necessarily also be treated as a synonym of P.1'. rusbyi.rush)'i. In lowland areas of sOUlhernsouthern Sonora,Sonora. a weedy, large-headed race OCcursoccurs commonly along roadsides and in cultivated areas (Keil{Keil & Canne 8631.8(51).8632, 8652). On some sites.sites, this form is so common that it becomes an aspect dominant,dominant. coloring large areas with itsits Showyshowy yellow heads. heads. In less disturbed sites, this form grades intointo the smaller-headed forms moremorc typical of the species.specics. Large-Large­ headed forms Occuroccur throughout much of the range of the species aBdand are not limited to the weedy race. On the basis of the evidence presentlyPresently available.available, taxonomic recognition of these forms does notn°t seem justified,justified.

P~(,lisPectis rush)'irusbyi usually can be distinguished from other relatedrcl~ted speclcssPecies by its broad, apically flattened,flattened. longitudinally striate phYllaries.Phyllaries. A few populations from Baja California and Sinaloa 4848 RhodoraRhodora [Vol.[Vol. 79W

(e.g. FlyrFlrf 68,68. Wiggins 15533A)/5533.4) haveha\'c smaller,smaller, moremore convex,convex. lessless conspicuouslyconspicuousl)' striate phyllaries.phyllaries. However,Ho....-ever. inin otherolher respectsrcspe<:IS thesethese plantsplants areare soso similar toto other populationspopulations ofof P.P. rusbyirusb.ri thatthai tax­13lt­ onomic anomie separationseparation eveneven atat thethe varietalvatltla] levellevel doesdoes notnot seemsttm war-war­ ranted.ranted. PectisPUlis rusbyirusbfi sometimes~mc:times hybridizesh}bridizes with.... ith P.P. papposapapposQ var.\'aT. papposapapposa and with P. filipes/ili/Hs var.var. filipesfiliJHs inin Sonora where.... here complexcomple}[ hybridhybrid swarms mayrna)' developdc\'dop locally.locally. InIn Arizona,Ari10na. however,ho.....c\·cr. wherewhere: P.P. pap­po~ posaposQ var.\'ar. papposapapposa and P.P. rusbyirusb); haveh,n'c beenbeen observedobscned toto grow together, together. no hybrids have beenb«n found.found.

REPRESENTATIVERE"'lq'I~"'l SPECIMENS:'"O'U'S: México,.\Ihiro. BAJA1>\.1.. CALIFORNIA('..l"....' ... SUR:~Il: Purissima,PllrmilTl&. Brandegee8T~ s.n..l.n. (COl.'(GH, US); ~l; WYo' of LosI.", Dolores,00I0rtS. WigginsII iain. 15533ÄHJJJA (MICH,TEX).1..1Ot. TH). SINALOA:'"",0.\: nearTopolo­Mar Topokt­ bampo,Nmpo, FlyrFI,. 696\1(,l'. (TEX, WIS);ons); 1.8 mi E of SanSa" Blas,BIaI. K&CA "C 8744'JUI (os),.....), SONORA:",...,...: Guaymas,GU}mu. GentryGrit") 46821M! (F,(r.GH. GH, MICH,"K;H MO,"0.'" PH, SD,'-incraUllasuberoso-incrassata fere u!iqueusque ad apicem,apiecm. tenuimarginatis,lenuimarginalis. ad apices ciliolatis..ciliolatis, alileraliter glabris.glabris, glandlbu$glandibus oleosis anguste aUIaut late elliplicisellipticis ad apiecmapicem elet utroque coslaecostae dense punctatis.punclalis. FlosculiF1()Sl;\11l r3diiradii 8.8, corollis 66-8 8 mm longis.longis, cilriOlScitrinis aut tub~ntibus.rubrescentibus, glabris,glabris. ligUlalígula ]~53 5 mm lonplonga 1-1.51 1.5 mm lata apiceápice minute Iridenliculala.tridenticulata. FloscuhFlosculi disci 1010-12, 12. coroUiscorollis 5 mm longis,longis. cilrinis.citrinis, glabris.glabris, paene regul:ltibus.regularibus, 5-denlalis.5-dentatis, denlibusdentibus 1.5 mm 'ongis.longis, antheris 1.5 mm longis.longis, basalilerbasaliter sagittatis.sagittatis, ad apicem appcndiceappendice minuleminute emargi­ nala.nata. Acheni3Achenia C)lindrica.cylindrica, discorum 3.5-4 mm longa.longa, radiorum 4.5 mm 10n1l:3.longa, brunnea aut nigra.nigra, uichomatibustrichomatibus brevibusbrevi bus badiis 1977]1977) Pectis•KeilPtttis-Keil 4949

, . ~:~ .,l;:j".­ d;; ,, ;;..,.. " .'j '10''.

\JJ.J!F al -,W 1¿A

I B H G " L

FFigure...... e 44. A-D.A-D, IWmPectis p"peruviana....itmD {fNlrl,,~(Hutchison & 14''lt'hlWright J4/i5414, ~1:F); f-l,F-L, Pectisp..mJ ~U.Papposa; E-H.E-H, <:arvar. pGpf"Wilpapposa (1.",1(Keil d& CMfnrIJ1J8,Canne 8758, 0'>1;os); 1.1_I-L, >lrvar. 8.lJnIiagrandis (A"I(Keil A& dMcCillrG.N 1l~~.7786, (K)os). A.A, E.E, I.I. ""1),\;habit; BB, F.F, LL. p/I)lIanphyllaries;...; C.C. G, J.J, "')ray a<:",",":achenes; D.D, II. H, K.K, disc1IoC -.:hent1achenes. 50 '0 RhodoraRhodora [Vol.[Vol 7979 clavatisc1avalis bicellularibus puberulenta.puberuknl3. PappusPappus idemidem inin flosculisf10sculis radiiradii acae disci,di$(';, 12-20 setosus,5C'lOSUS. setisSt:lis 44 mmrom longis,longis. antrorseanlro~ scabridis.~abridis. Chro­Chro­ mosomatumffiosomatum numerusnumerus ignotus.ignolus, DISTRIBUTION:DISTJl.ItIl no'· PresentlyPrt"scntl} knownkno.... n onlyonl} fromfrom thethe typetype collectioncollection inin northernnonhern PeruPeru at an elevation e!t:'"ation of 1100 meters.meters. PectisP~(,I;S peruvianapt'fUl'/onIJ is thethe onlyonl} representativen:pn:sentali,c of sect.sect. PectothrixPt'clfIIlm,' knownlna",n toto occuroccur outside of North1'l'onh America.Amcrka. The occurrenceocrurrence of taxalalla inin thethe NorthNorth American desertsdesuts and alsoalso inin PeruPeru isis notnot limitedlimited inin Pectis,PeClis. however,howc,'er. to sect. Pectothrix.P('C/mllr;T, PectisP«tif IIillifolio in i/o lia L.L of sect.$C'C1. PectidiumP,'aidillm hashas racesraCC" in the Sonoran DesertDeSt:r1 and also inin thelhe deserts of Peru.Peru. Indeed.lodc-ro. HutchisonlIuu:hison &&. WrightWrighl collected P.P. linifolia'mijofio near~ar thethe typetype localitylocalit) of P. peruviana.pt'fU\'i(mo. PectisPUlIS peruvianapeflll-fana bears an overallo\'crall resemblancen:semblaT'lCe toto P.P. stenophyllaJlenoph."'/a var. stenophylla.s/enophyfla. Both80th are31l: perennial taxataxa with narrowlynarrowly linearlinear leaves,lea~'es. elongate peduncles and a multisetosemullisetosc disc pappus. How-Hoy,­ e\'er.ever, these taxa differ inin several respects.respects. PectisPtc/is stenophyllaJII'"ophylla gen­ erally has smaller flowers, flowers. glandular puberulent ratherrather thanthan glabrousglabroUS corol1as.corollas, a biaristate rather thanthan multisetose rayray pappus andand slender rather than clávate clavate trichomes on the achenes.achcnes, ThcThe evolutionary relationship between PectisPen;s peruvianapt'IlH'iaIlU and thcthe remaining species of sect. PectothrixPecl/lI/my: isi~ notnot certain. The ffiultisetosemultisetose pappus is apparently a primitive feature in Pectisfrnis andand the aristate types haveha\e beenb«n derived from it.il. The perennial habithabit and muhisetoscmultisetose pappus of 1'.P. peruviana per/ll'iUllu suggest that this is a rela­rela­ tilel)'tively primitive species in the section. If these assumptions are true,true. then P. peruviana perm·jalla is probably an early offshootofbhoot from thetM.sec­ sec­ tion Pectothrix P'clofJr,i.Y: progenitor and has been !>eparatwseparated from thethe restn:st of of the section for a considerable period of time.lime. Whatever thethe length of time that P. ,,"u\·iatlaperuviana has remained in\f1 isolation fromfrolP the rest of the !iC'ction.section, it apparently has not enjoyed the evolution-evolution­ ..ary ry successsuccess that the North American species ha~~have displayed. 4. 4. PectisP«lis P'ppo!'papposa Harv. &.& Gray in A. Gray.Gray, Mem. Am. Acad.Acad. ..4: : 62. 1849. TYPE: Mn:ieo.Mexico, -CAUFO"'I"CALIFORNIA,"....- (probably collected near HermO!iilloHermosillo or Guaymas.Guaymas, Sonora).Sonora), CoullerCoulter 331JJI (Holot)PC'(Holotype, TeD!:TCD!; isotypes.isotypes, GM!.GH!, K.K., 1two.... 0 sheets!). Figure 44.

Prr,;,Pectis f'IlPpoJ"papposa lIan',Harv. Ii:& Gray ,var....r. 'fJIIPPOJiIepapposa AA. Gray.Gray, G~oI,Geol. SUr\'Surv. CalifCalif. Bot. 1:617­t: 611 18&0.1880. T'-,rTYPE: lniltdUnited SUln.States, r"llfOll"~CALIFORNIA: San DoeaoDiego CoCo.,.. t.JlflnLarkens,.. P"I_,Palmer 200!OOp.rp-P­ (IlolOlypc,(Holotype, GH').GHI). 1977J1977] PectisPc<:tis~Keil Keil 51

Bushy strong-scented annuals. Stems 1-30 cmem long, much branched, glabrous or minutelyminutel)' puberulent.pubewlenl. Leaves linear,linear. 1-6I 6 cm long, 1-2 mm wide, basally ciliate with 1-3I 3 pairs of bristles 1-2 mm long.long, otherwise entire, glabrous.glabrous, marginally punctate with rounded to oval oil glands 0.3-0.50.3 0.5 mm10m diameter. Peduncles 3--403-40 mm long, glabrous, with 2-6 hyaline,hyaline. lanceolatelanccolate bracteolesbrdcteoles ca. 1I mm long. Heads clustered at tips of branches. InvolucreInvolucre cylindric to eampanulate.campanulate. Phyllaries 8,8. rarely 7. 7, 9 or 10,10. linear, 3-8 mm long.long, 0.5-1.7 10mmm wide,wide. apically aeutish,acutish, basally truncate,truncate. gibbous.gibbous, strongly convex-keeled almost to the tips.tips, narrowly hya­ line-margined.line-margined, apically ciliolale,ciliolate, otherwise glabrous.glabrous, punctate with 1-5 swollen subapical glands lindand 2-5 pairs of smaller.smaller, inconspicuous SUbmarginalsubmarginal glands, occasionally with additional dorsal glands. Ray florets 8, rarely 7.7, 9, or 10; corollas 3-8 mm10m long, with narrowly to broadly ovate glabrous ligu[esligules and slender glabrous or glandu­g[andu­ lar+puberu[entlar-puberulent tubes. Disc florets 6-34; corollas 2-5.5 mm10m long,long. bilabiate.bilabiate, with the narrow lobe ca. twice the length of the teeth on the broad lobe.lobe, glabrous or glandular-puberulent below. AehenesAchenes 2-5.5 mmrom long, strigillose or pilosulose with slender, bulbous-tipped unicellular triehomes.trichomes. Pappus of ray florets co­ roniform.roniform, ca. 0.2 10mmm long,long. rarclyrarely I-several1-several aristate or setose. Disc pappus 16-24[6-24 setose.setose, 1.5-4 10mmm long.long, or occasionally only coroni­ form; bristles slender.slender, sub-plumose.sub-plumose, spreading. Chromosome number;number: n::n - 12.

DI~nRI8uno~:DISTRIBUTION: Southern California and southwestern Utah to westernWcstern Texas.Texas, south 10to Baja California, Sinaloa.Sinaloa, Chihuahua and Coahuila (Figure 5). PectisPe("/is papposapupposu occurs,occurs. often inin great abundance.abundance, in all four of North America's deserts, occupying a variety\'arielY of habitats. UnderUndcr favorable conditions.conditions, this species some-some­ times becomes an aspect dominant.dominant, coloring wide areas with its bright yellow heads. PectisPer/is papposafJUPpoS(l is divided into two well-we[1­ marked geographic varieties. The typical one occurs in the Mojave and SOnoranSonoran Deserts,Deserts. whereas var. grandisgrundis is restricted mostly to the Chihuahuan Desert. 4a.4a. Harv.lIan'. & GrayGra)' in A. Gray, Mem. Am. Acad. 4:4:62. 62. 1849. var. pallpnsa.papposa. Figure 4. Stems 1-20 emcm long. Leaves 1-313 cm long,long. 1-2 mm broad. Pe-Pe­ duncles 3-[03-10 (-25) mm long.long, 3-6 bracteolate. Heads densely 52" RhodoraRhodora [Vol.[Vol 7979 clustered at tipstips ofof thethe branches,branches. oror inin somesome MexicanMexican formsforms moremore diffuse. PhyllariesPhyllaries 3-53-5 mmmm long,long. ca.ca. 0.50.5 mmmm broad,broad. punctatepunctate withwith 1-3 subapical glandsglands and 2-3 pairspairs of submarginalsubmarginal glands.glands. RayRa}' corollas 3-6 mmmm long,long. ca. 0.50.5 mmmm broad,broad. glabrousglabrous oror glandular-gJandular­ puberulent. DiscDisc floretsOorets 6-14 (-18);(-18); corollascorollas 2-3.52-3.5 (-4)(-4) mmmm long,long. glabrous or glandular-puberulent.g1andular-puberuknl. AchenesAchenes 2-4.52-4.5 mmmm long.long. Pap­Pap­ pus of disc achenes setoseselOse or sometimessometimes coroniform;cOTaniform; bristlesbristles 1-2.51-2.5 mm long. Chromosome number:number; n"", - 12. FloweringFlollocring mostlymostly JuneJune to10 December.

DISTRIBUTION-.Dl,rRleCTIO', SouthernSoUlhcrn California and southwesternsouthwestern UtahUtah toto southwestern NewNell> Mexico,Mexico. south toto BajaBaja California and centralcentral Sinaloa at elevationsele\3tions rangingranging fromfrom minus 60 toto 1500 metersmeters (Figure 5).

PeclisPl'(f;S papposaPOfJPOJO var.vaT. papposapapposo often colors largelarge areas ofof thet~ Sonoran Desert during late summer.summer, ItIt isis particularlyparticularly abundantabundant in centmlcentral ArizonaArilOna onOn the broad inter-mountaininter-mountain alluvial plainsplains and also occurs on thethe lowlow desert mountain ranges.ranges. TheThe plantsplants of var. papposapopposa arearc morphologically variable,variable. and a number ofof intergrading locallocal races occur within itsits range.range. PedunclePeduncle length,length, branching habit,habit. and pappus structure are among thethe featuresfeatures that vary, but none of these populational differences seems worthyworthy of formal taxonomicta",onomic recognition.recognition. Bradley and Haagen-SmitlIaagen-Smit (1949) reported thatthat a largelarge portionportion of the essential oils of PectisPeetis papposaPQ{JpO.fQ (\ar.(var. papposa)ptJppoSQ) isis composed of cumin, and theythe)' sugg~tedsuggested that P. papposapaplWSQ might be grown as a commercial crop for this oil. To my knowledge,kno"'ledge. thisthis suggestion apparenllyapparently has not been pursued.

JhREPRESENTATIVE ....Sl"AlIH vt:Cl"£'~:SPECIMENS: ;\lhico.México, aAJ~BAJA ("'11_\''''CALIFORNIA: S of LacunaLaguna SecaSta Chápala.Oarab. C"nn.Carter, Aln"""'.1Alexander & KelloggK,'~ 18\101890 ("".(DS, Gil,GH, _.MO, SD);'0): LOSLoo. "n&donAngeles Bay,&y. PalmerP,,/-, 657dJ1 1<011(GH, '[l-4;.ND-G, ,y,NY, l~,UC, lS).us), BAJAa.v~c CALIFORNIA... lltoa'IA:;t1l SUR: 6 numi S ofor La Paz,Pu. HammerlylI"mnwrlr 235~)5 (CAS.(eA' I>\,.DS. so).SD), '\.1'.010"SINALOA. 1I mim. EF. orof Altata,Ah"Ul. FIyrnlr II)113 In\(TEX. "'1\).WIS). 'O-.p;I. Pringle,1't,~Ir, s.n...... {I(F. lonCharleston Mts.,\\ls,. C/oi.",Clokey 8198197 1<" OS.I,DS. F. (,11,GH. ..KSC...c, ...MICH,Klt. "'''.MIL, "0,'"MO. PH. '>P, SD, ~"l.SMU. TEX,,n.... wis);,,): t.,na>InLincoln Co"Co.: Timpa"nllcTimpahute Range.R,,"'" R,w"l8:Reveal 8L II",J",~,Holmgren... JIl(U1904 (D'>(DS. ~"'

4b. 4b. Pectis papposa Harv. & Gray in A. Gray var. grandis Keil, Brittonia 26: 32. 1974. TYPE: United States.States, TEXAS: Presidio Co., 3 mi E ofjunctof junct US RteRle 67 on Texas RteRtc 170, E of Presidio, McGill & Keil 7768 (Holotypc,(Holotype, us!; isotypes, ,l.Rl7!,ARIZ!, ,l.su!,ASU!, ENCB!,Er>CII!. f!,F!, Gil!,GH!, MICH!,MICU!. MO!, NY!, os!,OS!, RSA!, SMU!,S1>HI!. nx!,TEX!, UC!).IX!). Figure 4. Stems 5-30 emcm long. Leaves 2-6 emcm long,long. 1-2I 2 mm broad. Pe­ duncles 1-4 emcm long, 2-6 bracteolate. Heads usually several at tips of branches. Phyllaries 5-8 mm long, 1-1.7 mm broad,broad. punc-punc­ tate with 1-5 subterminal glands and 2-52 5 pairs of submarginal glands. Ray corollas 5-8 mm long, glabrous or sparsely glandu-glandu­ lar-puberulen\.lar-puberulent. Disc florets 12-24 (-34); corollas 3-5 mm long,long. glabrous or glandular-puberulent. Achenes 3-5.5 mm long. Pap­ pus of disc achenes 2.5-42.5--4 mm long,long. setose or rarely short-coroni­short-coroni- form. Chromosome number: IIn = \2.12. Flowering June to October.

DISTRIIIUTIOr>:DISTRIBUTION: SOuthernSouthern New:"lew Mexico and southwestern Texas to Chihuahua and northern Coahuila; disjunct in ArizonaAri70na and nOrtheasternnortheastern New Mexico (Figure 5). The distribution of this variety is primarily in thcthe Chihuahuan Desert at elevations of 1000 ~oto 2000 meters. The plants of var. xrandiJgrandis are particularly common Inin the Rio Grande valley in Texas.Texas, New Mexico and northern Mex­ ICO.ico. This taxon is undoubtedlyundoubtcdly more common in northern Coahuila than is indicated by presently available collections. The plants ~owgrow almost to water's edge along the Rio Grande on the Texas Sideside of the border,border. but I have han seen no collections from thethe oppo-oppo­ site bank.

RfrR[~"TA[I\[REPRESENTATIVE ~rln"r'sSPECIMENS: S« See Keil (1974) (orfor specimen$JlC'cimen eilalion,citations for PectisPerti, ""ppa'oPapposa Var.var. troll"i,.granáis.

5.5- PettisPectis anguslifoliaangustifolia Torrey.Torrey, Ann. Lyceum Nat. Hist.His\. Newl"ew York 2: 214. 1828. TYPE: UnitedUniled Statrs,States, (COLOR,l.I)()(COLORADO or NEW'EW MEXICO): ~on"on the Rocky Mountains," 1820.1820, James HI.s.n. (Hulolype,(Holotype, NY!).w1). Figure 6. ;~~rrQjHehoreos ,onguJrif"/'uJangustifolius (Tor",)')(Torrey) Raf.,Raf.. AtlanticAllanl;c Jour. 1:I: 145.145, 1832. 1'.KtidopsislidopSl.< al/g,wifutioangustifolia (Torrey) DC,DC.. Prodr.Prod,. 5: 98. 1836. nom. illeg.illeg, 6C S an r~:"" 0"llriJIifolio8ustifolia Torrey "ar.var. JUbOr;j/0I0subaristata A. Gra)".Gray, SmithsomanSmithsonian Contr.Con". Knowl. : 82 L: n - 1852,1852. TvrrTYPE: UnitedL'ni'~ 51.lh.States. TEXAS:Tn~~ "''IIIIc:}'"valley bel",..,nbetween the Pecos"""OS andand thelhe lm la 1'.p "t'P! Socorro CoCo.,.. SO<:O.1)OO' ' • var. grandis J' • .- var. grandis '::0<,.0,,0D0ttz "",,: ' f=,"' . - "I ' •• intermediatesintermediates o ~ ~.O~. ~ .f • ~" __' o PectisPedis pringleipringlei ,. '-~"i"':: PectisPeelis incisifoliaincisifolia 1 • L i :r"!:~-.,, " . ..J, •. .,I· . . , • ,'1. '-. ,\ ,", "/~-~~...... , 1 ? ' \ r 0 i"" 0:-'-' 1!1'·\:::r'-·~·' \ ...... -. , , " ;' /" ,-' , :",., '.-' , , l',F'Ul _-..~ r^rn, "/, .

FigureFilur~ 5.5 GeographicalG~&raphl<.l distributiondi$uib1ltion of PectisPrrm papposa,paP(HM4. P. pringlei",ing~j andand ".P. ""·"ifol'

Annual or perennial herbs from a slender tap-rootlap-rool or ligneousligneouS caudc.K.caudex. SIemsStems 1-20 emcm long,long., usually branched fromfrom thethe base,base. glabrous or minutely puberulent.puberulcnl. LeavesLe:avC5 linear, 1-4.5 cmem long,long, 1-3 mm broad.broad, 2-52~S cilialcciliate alat the base with bristlesbrislks 1-2 mm long,long, mucronale.mucronate, oflcnoften slronglystrongly ~\OIUIC.revolute, glabrous, marginally punctate ~ilhwith glands 0.2-0.7 mm diameter. Peduncles 1-20 mm long,long. with\lolh 2-5 braeteolesbracteoles 1-3 mm long. HeadsH~ads congest~dcongested at ~ndsends orof branches,branches. narro\\,lynarrowly campanulalecampanulate or cylindrical. PhyllariesPhyllarin linearlin~ar or nar-nar­ ro\\,lyrowly oblanceolal~,oblanceolate, 2.5-5.5 mm long.long, obtuse,ObluM:. basally truncatetruncate and strongly gibbous.gibbous, vcryvery strongly con\ex-kecledconvex-keeled nearly to thelbe apex,apex, glabrous, punctate sublermlnallysubterminally with 1I or 2 swollen oil glands:.glands, and submarginallysubmarginally \\"ithwith 2-5 painpairs orof smaller glands. RayRa} floretsflorets 8:8; corollascorollas 3-5 (-7)(-7) mm long, glandular-puberulcnlglandular-puberulent or nearlyn~rl) glabrous,glabrous. Disc florets (7-)(7-) 10-20:10-20; corollas 2.52.5-3.5 3.5 mm long, regular oror bilabiatc,bilabiate, glandular-puberulent. Achenes 2,5-42.5-4 mm long,long. stn­stri- 1977])9771 Pectis•KeilPectis~ Keil 55" gillosepilose withwith bifurcatebifurcate trichomes.trichomcs, PappusPappus variable,variable, coroniformcoroniform and/orand or 1-7 aristatearistatc oror setose,sc:tose, 1-21~2 mmmm long.long. ChromosomeChromosome num-num­ ber:ber: nII = 12.

DISTRIBUTION:DISTlllllllO': WyomingWyoming and NebraskaNebraska southsouth toto Durango,Durango, SanSan LuisLuis PotosíPotosi andInd VeracruzVeracruz (Figure(Figure 7).7).

InIn thethe PectisP'etiJ angustifoliaangwlijolia complex previouspre\'ious workersworken (Gray,(Gra), 1884;1&84; Fernald,Fernald. 1897;1897; Rydberg,Rydberg, 1916) havehave reliedrelied primarilyprimaril)' onon thethe: featuresfUtures of theIhe pappus as taxonomiclaxonomic characters.characlers, ThreeThree taxataJIa tra­Ira· ditionallyd~lionally havehne been recognizedrttogniztd inin thethe: complex on thetnc basisbasis ofof differencesdlffercnc:e:s inin pappus structure:struClure: PectisP,('/u angustifolia,allgus/i/olta, P.P, angusti-angus/i­ folia var.\ar. subaristatasl/haris/a/a and P. tenella./f'nf'lfa, A fourthfourlh group of plantsplants hashas twicelwice been named as a species {P.(P,jastigia/Q fastigiata Gray,Gray. 1849; P.P. texanaff'xana Cory, 1937) and twicetwice re-unitedre.united withwilh P.P. angustifoliaa"gustijolia (Gray, 1852;1852: Correll &do:. Johnston, 1970). The value\alue of the pappus as a charactereharaeler in thisIhis complex has beenbeen greatly Overemphasized.overemphasized. Although easily distinguishable formsforms exist~Xist in someSOme regions, in other areas there is so much variation mI~ pappus SlnlCIUrestructure that as many as threethrec "taxa"~taxa~ could beDc recog-recog­ mUdnized from thethe members of an otherwise homogeneous population.population. A re-examination of taxonomic boundaries,boundaries. necessitatedneces.silated byby thethe breakdown of thethc traditionally accepted scheme, has ledled me to10 twot.... o eonclusions.conclusions. First, thetnc PectisPUfis angus/i/o/iaangustifolia complex should be:e .rccogni~edrecognized as a single species with threethree varieties: P.P, angusti­angus/i­ foilo*°ha var.\'ar, ol/gus/l{Qlia,angustifolia, var.\'ar. ff'tNd/atenella and var.var, laJ/iKlofO.fastigiata. Second. Gra}'sray's var.\ar. sllharis/atosubaristata has ofoftenlen beenbttn misapplied,misaprJit'd. and in itsIts strict Interpretalion,lr>terpretation, cannOIcannot be distinguished from var.\ar. angustifolia.a/lK/lSfi(oJia. Sa.a PectisPettis tnluslifolitangustifolia TolTt)'.Torrey, Ann. LyceumL)'~um Nat.:'\11. Hist.His!' New~ew York 2;2: 214,214. 1828,1828. var,var. InCuslifolil.angustifolia. Figure 6. bj BuSh*!usn)y lemon·scenledlemon-scented annuals. SiemsStems 1-20 cmem long,long. densel)densely at the tips Le fyY i1t the tips.- Leavesaves 1-4.5 emcm long,long. I1-3 3 mm wide,wide. conspicu­ :usl)° y finedflared at the bases. Peduncles mostly 1-10I 10 mm long, 1-4 r1cteolilte,racteolate, ofoftenlen "'holl)'wholly or n"r1iall)'partially con~aledconcealed byb\' lhethe bases of theb' ...- , e susubtendingtendmg leavcs.leaves. Ph\"lIariesPhyllaries 2.5-5.5 mm long, linear,Iincar, strongly •eeledee ed t0 th . lert£ . to thee lips.tips, abrupllyabruptly truncale,truncate, punctate wilhwith 1I or 2 sub­ rminal R llllnal glands 0.2-0.50.2-0.5 mm long.long, and smallersmaller submarginal glands. a l~~2 y Corollascorollas 3-5 mm long. Disc florets (7-) 10-20:10-20; corollas 3.S35 mm long, slightly bilabiate. Achenes 2.5-4 mm long. 56" RhodoraRhodora [Vol.[\'01 79N

...., , /'1, /.~z .( / '.' I ,~ . ", (1': , J mm em

l ~ .. , A , I ( 'i • ~, ,-', ,. ,, \ I ., ' , UGjH ",F I B ,C ','0

FiIUftFigure ll.6. PectisPrr'lJ il1fXIIJIl/oli«angustifolia;.. A_D,A-D, var.'"v angustifolia~J(ifQ/•• ("t>l1(Keil 7634,os);76,U.OS): E[-II. H, ,aIv JfIJl'8"'liIfastigiata (1JlIJ'1'(Tharp "& W.,POOdWarnock 45-30.45-30, uu):ARIZ); I-L,[.L 'var..., ,rnr/JtItenella (Kn/"(Keil & McGill"di.N 784°t": • (5).os). A.A, E.E, I,I. habol;habit; B.B, F.F, J.J, plI)Ib.~;phyllaries; C.C, H,H.... K, ,a)'ray achenes;.dW:Tla; n,D, G.G, L,t. discd~ achenesitChdO'

A Pappus cOToniform.coroniform, 0.1-0.3 mm long,long. or in !Ornesome TuasTexas anoand Chihuahua populations also 1-7 aristate or sC'tosc.setose. ChromosomChrolTloSonll' numbernumber, 11n =- I~.12. Flowering July to October. D'SrII.ISUTIO':DISTRIBUTION: Western Nebraska to eastern Ari7.ona.Arizona, western\lester' Texas.Texas, Chihuahua andand northern Durango,Durango. grading into var. l,nttenelilU inin easlerneastern Chihuahua and Durango (Figure(Figure 7). 1977]1977) Pectis•Pectis-KeilKeil 5757

••':J • •

Pectis angustifolja • var. angustifolia A var. fastigiata ° var. tenella * « intermediates

Figure 7. Geographical distribution of . 5858 RhodoraRhodora [Vol.[\'ol.1'9 79

The presentpresent interpretationinterpretation of morpho-geographicalmorpho-geographical variationvariation inin thethe PectisI'et'/ls angustifoliaanKll$/ifo/io complexcomplex hashas necessitatednecessitated aa re-interpre­re-inlcrpre­ tationla1ion of var."ar. subaristata.suhD,isfa(u. This namename hashas beenbeen appliedapplied widely.....idely overO\Tr theIhe: years)'t:ar§ to10 any individualmdividual withwith aa few-bristledfc:",-bristled pappus.pappus. AnAn exam­exam­ inationination of thethe: typet)'J)e materialmaterial of thisthis taxontaxon revealedrevealed that,that. withwith theIhe exception of thethe slightly bristly pappus,pappus, thesettlne: plantsplants areare notnot dis­dis­ tinguishablelingl,llShablc from(rom individualsindi\'iduals of var."ar. angustifolia.anRlISllfu/ia. TheThe: "subaristate"~subaristatc:~ forms(onns toto which.... hich thisthis namename hashas beenbeen appliedapplied arearc: of several51:\c1al differentdirren:nl origins. SpontaneousSpontaneous "subaristate"hsubaristatc~ vari­\'an­ ants occur sporadically throughthrough muchmuch of thethe rangerange of PectisP«tiJ angustifolia(lajlus/i{ol'Q var."ar. angustifoliaanJ(uslifofia and are particularlypanicularly commoncommon inin western""estern TexasTc~as and northernnOflhcrn Mexico.Mc}[ico. WithinWithin populationspopulations ofof var.\'ar. tenella,1t'n~If(J. individuals may be "subaristate"Msubarislatc~ or evencvcn completelycomplc:tc:ly epap­cpap­ pose.pose:. A/I. similar condition sometimessomclimc:s occurs inin var.var. fastigiata./QJ/if{IQIQ. InIn somesomc areasarc:as of northern Mexico,Meltico. individualsindl\'iduals intermediateintermediatc be­be­ tweent"een var. angustifoliaan1{lUli{QIiQ and var. tenellaleneJfa may havehave aa subaristatesubaristatc: pappus. Finally,Finally. inin eastern Chihuahua and western Texas,Teltas, "sub­"sub­ aristate"aristatc" races have developed apparently as a resultresult of hybridiza­hybridiza­ tion and introgression betweenbetwecn var. angustifoliaQfl1{lIsli/oIiQ andand P.P. papposaPQPposo var. grandis.Krandis.

REPRESENTATIVERIPH~I "Amf SPECIMENS:'PIO"'" Mexico,:\1;,1<0. CHIHUAHUA:C'IIIHI ""I A nearno" Chihuahua.Ch,huahua, Pringle/'Ti,,~~ 648648 (""(Ds. IF...... GH. MICH."I("II MO."0 NY.'v PH.~". RSA.R~A us);l"): 43 minli N of VillaVItia Ahumada,AhullUldl. StuessyStu<"JJI ¡114/JU (DS,(1"­ ENCB.NY.TEX).",I.,y ,rY) DURANGO:III R"'(;() 1.8 mim, E of Nazareno,:'\uartno. KK.I & M8014(os).\f 8(lU (0'1 SONORA:""'ORA Guade-Guade­ lupelup< Can)on.Canyon. MearnsIf"",,,. &A Mertonlit""" .'(IJJ2034 ((DS..., NY.,y US).~~l. UnitedI niltd SialStates..... ARIZONA....uO".. CoconinoComllll\O Co.:Co. 12 mi N" or Cameron,Camt.on. Cutlere",I" 3122Jill (DS,(pS -E of La Junta,Junia RollinsRoll... 1~1J¡873 I'''',(DS. \00.MO. 't>.ND. ".NY, l'),US). KANSAS:~""A~ EllisEll .. Co.:Co., 12 miml SW of Hays,III) RunyonRWIl"" A& Bondy&>I1J. :M288 I"(MO... rllPH. ,,,,):SMU); Tregor~ Co.Co.: "'"houlwithout locality,~luy. HitchcockU"tIrnx4' 294."91 (GH,1<''' KSC~<,(" "'0.MO. '\0(NMC. ").NY). "NEBRASKA:....,~.. ~H)Cherry CO.Co.: Ft.Fl. Niobrara,"Obrl'l. BateslIoJu. s.n.,.... (CAS.I,A.... GH); Deuel(kud Co.: 8<1Big Spnnf5Springs RdRd. IIat SS. PlantPlatte R"n.River, KA 161J7633 Irui Ana Co.:Co.' OrpnOrgan \ttMts..... II.,."Wooton.... 4!f>426 (I>'\.(DS, G"GH. ~<;CKsc, MO.\to. ND-G,'1)0(, ,,,..NMC. NY.,y "'"POM, 'c.UC, US);l~); GrantG..... CoCo.: BnlBear Mts..\h~. MetcalfeIrn••lft 699 4uLI.(ARIZ, (.HGH. MO.1.lO. NMC.'''C' NY.".PO" POM. UC.IC.,~): US); Lincolnllncola Co.:Co G",Gray,... .5'tlaoSkehan.. ~J45 II(F. (';"GH. K\C.KSC. '000.MO, ''OCNMC. "NY, "'"POM. \C.UC. \US); ,); "~.raSierra Co.:Co Berendo""tndo Creek,Crcd Bl«kBlack RIRange,...... MetcalfeI/t,...lft IJ1387 1Ic"" (CAS. IF. (,,,GH. 1.lO.MO. NMC.''"' "NY, PO".lCPOM. UC. US).l-.oj. Ol"OKLAHOMA. " ..0.... CimvNnCimarrón Co:Co.: 2 m,mi S of K:tolKenton,..... ~Demaree....'" /J)IU13381 (';"(GH, MO,"'0 ".NY, rllPH. 1'0"POM, SMU)-~"'L). TEXAS:rn", OulDeaf SmllhSmith CoCo.:.. I1 mi S of Gknno.Glenrio, 14Waller ..11" 1526u.'¥, ((ASU...'it "cENCB,•. ",,'SMU. TTC);nc); JeffJeff ~,~Davis Co.Co.: FIFt. 01Davis, Ptt~,Palmer J]IJII"'O32151 (MO. '''1.PH). 'HIIUTAH: San Juan Co.Co.: CopfltfCopper e"Mon.Canyon, CutinCutler J14~3146 1(;(GH. '<0MO. '"NY. ..SMU).", t, ....WYOMING:()..."t. (jos~"Goshen Co.Co.: 1112 mim' bolo'"below flFt. la"m~Laramie on"on N. Plane.Platte, SimpJOor"sSimpson's L~JHJExped. s.".I\OO1s.n. (MO) Sb.5b. PtclisPectis .ngusliroli.angustifolia Torr~Torrey 'ar.var. tenellaItndl. (DC.) Keil,Kc:il, comb.comb, nov. BAS10,"Y\I:BASIONYM: P"crisPectis I"n,,/fatenella DC,nc., Prodr. S:5:99.99. 1836. TypfTYPE: 1977)I977J Peclis-KcilPectis•Keil 59

Mexico. TA.\lAUIIPAS:TAMAULIPAS: "prope~prope Laraedo,"Laracdo," Berlandierlkrfandier 2009 ( = 599) (Holotype, G-De;G-DC; photo of holotype,hololype. US!; microfiche, JDCIDC 800. 787: III. 5!;51; isolypes.isotypes, (l009)(2009) Gil!.GH!, MO!,MOi. NY!,NYi. US!.US!, (599) Gil!.GH!, K!,K'. MOi.MO!, I'll!).PH!). Figure 6. Bushy slrongstrong scented annuals. SiemsStems 2-15 cm long.long. Leaves 1-4 cm long, not or scarcelyscarcely flared at the base,base. evenly distributed on the stems. Peduncles 5-15 mm long, 1-4 bracteolate,bracleolate. not or s<:arcelyscarcely concealed by the bases of the sublendingsubtending leaves, thethe heads evidently slender-peduncled. PhyllariesPhyllarics 3-5 mm long, linear, widest ncarnear the middle, punctate with a solitary sub-sub­ terminal gland 0.2-0.5 rommm long, and smaller submarginal glands. Ray corollas 3-5 mm long. Disc florets 10-18; corollas 2.5-3.5 mm IOllg,long, regular or slightly bilabiale.bilabiate. Achenes 2.5-4 mm long.long. Pappus highly variable, coroniform and/or 1-7 SClosesetose or aristate,aristale, 1-3 mm long. Chromosome number: n -= 12. Flowering June 10to January.

DISTRJRUTION:DISTRIBUTION: Eastern Chihuahua and southernsoulhern Texas to Agua~calicnlcs.Aguascalientes, San Luis PotosiPotosí and Veracruz. Also one station in nOrthernnorthern New Mexico where probably adventivcadventive (Figure 7).7). . In most areas, PectisPee/is anxus/ifoliaangustifolia var. tenella/ene/fa is easily dis­dis­ llnguishedtinguished from var. angusfifolia.angustifolia. Some individuals from Chi­Chi­ huahua.huahua, Durango.Durango, Aguascalientes and Zacatecas,Zacalecas. however,howC\"er. pos-pos­ sess thethe flared leaf bases or lemon scent of var. angustifolia,angus/ijofia. butout inin most other respeclsrespects Iheythey resemble var. fenefla.tenella. Because thesethese individuals have the open branching pattern and bristly pappusPaPpus of var.var, fenella.tenella, they are referred to thislhis variety in thelhe presentPresent treatment. treatment.

RrPREsnrAl"rREPRESENTATIVE 'PH""'ssSPECIMENS: Mhko.México, A"lA'>CAllrsAGUASCALIENTES: If' nearneu Aguascalientes,Agul>Clllienlc>. RouRose &:& }fa)Hay 775J7753 (GIl.(GH, SV,NY. l'~),us), OHHlAIlUACHIHUAHUA. 27 mi E of Jim';nc/Jiménez,.. ShreveSh't,',8867 8867 (~Rll.¡ARiz, GH. "ICHMICH, us), COAlWllACOAHUILA: 27 rnlmi S of Monclova,Moncion, KK" & M~f 7879 (0'\):(os); 252S mi SWW of Sa ' Y of Sab,nal,binas, II.,,,,;Wynd &.& .l!ulk,Müller 215]J' JAR(ARIZ. II. "",GH. .\to.MO, ND,'0. NY,\>. US),l'i). DURANGO:m R~"l0 betweenbci"'ecn e ~rblnilrbanis &:& C,,~ncame.Cuencamé, (;M1r\Gentry 69-1/6941 (GH. MICH,'IIC". us);I'S): 252S mirni N'" of Bermejillo,RerrrtCl,1I0. KA &J. MIf :/,9}192 /0'<1.(os), sn,,,NUEVO LHhLEóN: 2S25 km ....NE E of MoolMonterrey,.....y, Dieterlem<'luk 3678J678 (MICH),("ICH) SANs~s LUIS'11' r Tos Tos!í: San Luis Potosí,Potosi. ShaffM'rShaffner J]5325 f:]J5,[=235, 75JI751] II(F. Gil.GH, MICH,'UCH. NY.S\ 1(",VC. us).lS). ;~.\tAlIIPA'AMAULIPAS: Micr.Mier, G,eu,Gregg ,.".s.n. (Gil,(GH, MO. PH);PHI: vicinityviconily of Victoria,Victoria. PalmerPalmer 43101 F ;.< - "0Mo, sv,NY, U',UC, L').us). vrRAcaUlVERACRUZ: Chinameca, PMiche]7Péniche 27 (F,(F. us),V'i). ZACATECAS:I~CAlfC"S near an a~ ~aphacl,Raphael, .'lI,,,wShreve 9J9Q9390 (A"ll),(ARIZ). nllodnited Stale..States, SEWNEW "'H'COMEXICO: Rio Arriba Co.: Capulin Volcano, ClarkClmk 4608-1608 ~;"').^NM). TEXAS.Tf~~s McMullen Co.: 11V.l¡% Ill;mi S of Tilden,Tildcn. Co,,"Cory 5JJJ955339 (SMU.('i"~' "I;US); WebbWd.b Wlth °", "'llhou!°ut localily,locality, 17IarpTharp "ns.n. (Gil.(GH. MO.\10. TEX.!IX. "'1·VC). 6060 RhodoraRhodora [Vol.IVol.19 79

5c. 5c Pectis PKlis angustifolia TorreyTorrey var.\'ar. fastigiatafllJlieial. (A.(A. Gray)Gray) Keil,Keil. comb. nov.no.... BASIONYM:BASIO"1VM: PectisPre,;s fastigiatajtuliRio/Q A.A. Gray,Gray. Mem.Mem. Am.Am. Acad. 4: 62.62 1849.1849. TYPE:TYPE: UnitedUnited States.Sfalrs. TEXAS:rEXAS: TravisTravis Co.,Co.. Austin,Auslin, WrightII'riRhr s.n.S.tI. (Holotype,(Holotype, GH!;OH!: isotypes,isotypes. GH!,(iH! ...NY!,y~. US!).US'). FigureFigure 6.6.

Pectis(W.n texanaIH_ Cory,COf). RhodoraIl.hodon 39:39: 421.421. 1937.1937. TYPE:rtP1: UnitedI nitll'd States.~.IOS. TEXAS:TfX,\S SuttonSUlIOll Co.,Co.. RanchRallC'h Experimentupl'mnnt Station,5tallon. CoryCo...· 15382ISM] (Holotype,(llolol)pt'. GH!;GH'; isotype,iKlfH"'. TEXI).Tt'<'). Fibrous-rootedFibtous-rOOled strong-scentedslrong-scented perennials or sometimessometimes annuals.annuals. Stems 5-15 cmem long,long. sometimes woody"'oody atat thethe: base.base. LeavesLca,'cs 1-41--4 cmtill long,long. 1-212 mm wide,",ide. not or scarcely flaredflared atalthe the base,base. notnot crowdedcroll,dtd at thethe tipslips of thethe branches. PedunclesPeduncles 3-303-30 mmmm long,long, 3-6 bracte-br.lctr­ olate. PhyllariesPhyllarit's 2.5-4.5 mm long,long. narrowly oblanceolate,oblanceolatc:. punc­punc­ tatetate with a conspicuous subterminal oil gland 0.5-1 mmmm longlong andand 1 I or 2 pairs of smaller submarginal glands. RayRay corollas 4-64--6 mmrom long. Disc florets 8-21;8-21: corollas regular,regular. 2.7-42.1-4 mmrom long.long. AchenesAdK::nes 25-3.52.5-3.5 mm long. Pappus 0-40 4 aristate,aristate. 0.5-10.5 I (-2) mmmm long,long. also more-or-lessmore-or-Iess coroniform. Chromosome number: n -= 12. Flower-Flower­ ing September toto November.

DISTRIBUTION:DISTRIHUTION: Endemic to thethe Edwards PlateauPlateau area of cen-cen­ tral Te;\as.Texas, mostly on limestone soils (Figure 7).1). Until now,now. the perennial nature of some plants of PectisPee/is an­on' 1!./lstifoJiogustifolia var.vat. !osrigio/(lfastigiata has not been noted. Some individualsindi\'iduals of this taxon develop strongly lignified rootroot crowns.cro..... ns. Although both P. fastigiatajosrigiulo and P. texanau'xono were\liere both described as annuals,annuals. ilit is apparent from examination of additional specimens of thesethese planuplants that the perennial condition prevails in severalse\'eral populations. Correll and Johnston (1910)(1970) evidentlye\;dently were\liere referring primarily to populations of var.jasfiglotovar. fastigiata when they indicated that,that. in Texas,Texas. PUlisPectis onKUJ/ifoliaangustifolia occ.:UTSoccurs primarily on limestone soils.~ils. Both var.\'ar. angzlJlifolioangustifolia and var.\"1U. un~f{otenella occur commonly on a variety of substnltessubstrates and are Commoncommon on sandy soils. Only var. fastigiatajaniy,io10 isis largel)'largely restricted to limestone soils. In \l;eslo«ntralwest-central Texas.Texas, on the \l;csternwestern edge of the EdwardsEdl\"ards Plateau.Plateau, se\'eralseveral populations occur ....which hich are some\lihatsomewhat similar morphologically'morphologically 10to PeclisPectis on1!.uSlijoliaangustifolia var.\'3r. fen~1I0tenella (Troq(Tracy 1852;185].­ Egg~"Eggert j,n,s.n., 13 Jun 19(0).1900). These sites, hO\l;e\·er.however, are well\l;ell outside the general rangerange of this variel)'.variety. The populations are situated within",ilhln the rangerange ofof \'aT.var. anguslifoliaangustifolia hO\l;ever.however, and not far outside the tangerange 1977] Pectis•Keil PeClis-Keil 61 of var. fastigiata.jasligiara. Although none of these plants appear to haveha\'e the perennial habit of var. fastigiata,jastigiala. some individuals possess the elongated subterminal oil glands of the phyllariesphylJaries and the shonshort aristate pappus of this variety as well as the flared(Jared leaf bases of var. var. ongllJtifolio.angustifolia. Because of their geographical location and thethe evidenccevidence of morphological intermediacy, it scemsseems best to treattrcat these populations as intermediates between var. angustifoliaanguSlifo/ia and var·lastig/alavar. fastigiata rather than as disjunct populations of var. tenella.

RCPHSE'iIATIV[REPRESENTATIVE SP~\'I'lE"SPECIMENS: t:lll1NlUnited SialStates,... 'IEXA

6. 6. PettisPectis pringlei Fern., Fern.. Proc. Am. Acad. 33: 76. 1897. TYPE:TYPE; Me;o:ico.Mexico. COAHUILA: Jimulco,JimuJco, Pringle 125/15 (Holotype,(Holotype. GH!).GH~). Figure 8. Strong-scented, tap-rooledtap-rooted annuals.annuals, branching from the base. Stems 2-25 em cm long,long. erect or decumbent,decumbent. pseudodichotomously brart<:hed,branched, glabrous or puberulent. Leaves linear, 1-5 emcm long,long. 1-2 mm broad, often revolute, basally ciliate with 2-4 pairs of b~is!lesbristles 1-2 mm long, glabrous or abaxiatlyabaxially puberulent,puberu!ent, mar­ &Iginallyna lly punctate with conspicuous elliptic to round oil glands 0.2-0.7 0.2-0.7 mm diameter. PedunclesPedunctes 8-50 mm long,long. glabrous, with 1-4 scalteredscattered lanceolate bracteolesbracleoJes 1-2 mm long. Heads inin forks 1 of°f .Stemsstems and aJlitlaryaxillary, • becoming somewhat crowded in age. Phyl-Phyl­ lariesanes 8, linear to narrowly oblong, 4-6.5 mm long,long. 1-1.5 mm broad, broad, obtuse.obtuse, conspicuously hyaline-margined,hyaline-margined. with slender,slender. c~rkr,c°rky, basally gibbous midribs.midribs, apically villous-ciliolate,villous-eiliolate. other­other- WIse wise glabrous.glabrous, each punClatepunctate with a solitary subterminal oil gland and 1-4 papairsirs of smaller, elongated submarginal glands. Ray floretsfl°rets 8; corollas 5.5-6 mm long,long. with narrowly to broadly obo­ ~.te^te glabrous ligules and sparsely glandular-puberu!entglandular-puberulent tubes.tubes. lsc IS( floretsforets 1O~21;10-21; corollas 3-4 mm long,long. regular.regular, glandular-gtandular­ PU.berulent.Puberulent. AehenesAchenes 3-4.5 mm long, strigillose.strigiltose. Pappus 11-4 4 arJstate;estate; awns slender, antrorsely barbed,barbc

DISTRIBUTtO:';,DISTRIBUTION.- Southern Chihuahua.Chihuahua, southern Coahuila and llOrthernn°rthern Durango at 1100 10to 1800 meters elevation (Figure 5). [Vol.[Vol. 7979 6262 RhodoraRhodora

.~ (.

B

",preFigure I8. A_D.A-D, Pur;'Pectis pn1t(/"ipringlei (Mil(Keil & McGill\f,G,N 7976,jJ"&. O'i)'os); [.11.E-H, PeclisfWm incisifol*_Ill"" 110.".1"(Keil & McGill\/.GI118/fll.0">1 8105, os). A. E.E, h..Ibll;habit; e.B. F.F, ph)lbphyllaries,..".. O.D. G. raynI) ¥ht~achenes; C,C. H-H d",disc achoachenes....

PtellsPectis pringleipringle; is locally quite common on dry stony sites in thtthe southernsouthern pansparts of the Chihuahuan Dutn.Desert. Unlike the more wide­wid~ spreadspread P. utlKlJsrifolia.angustifolia, P. pringle;pringlei usuall)'usually does not OCl;UP)'occupy sites on 1977J1977] Pectis•Pecti~ KeilKcil 63" tilethe broad alluvial plains of this region. The latter species occurs on a variety of substrates including both igneous and sedimentarysedimcutary rocks.rocks. Extreme forms of PectisPee/is pringlei approach P. papposa var. grandis in habit and general aspect. The two taxa are easily dis-dis­ tinguishable by their very different pappus structure and by thethe morphology of their phyllaries (Figures 4, 8). Some small, rather depauperate individualsindividuals of P. pringlei, on the other hand, resemble P. angustifo/iaangustifolia var. fenella.tenella. The overall silesize and less congested appearanceappearance of the heads on P. pringlei, however,howevcr. make distin­ gUishingguishing the two taxa rather easy. I haveha,'e observed P. pringlei growing with both P. papposa var. grandis and with P. angusti-angus/i­ folia vaevar. /enel/atenella without an~'any evidence of hybridization.hybridi7.ation. The similarities observed with these taxa have probably resulted from localizedlocalized selection rather than from hybridization and intro­intro· gression.

RfPR~S£~T~TIHREPRESENTATIVE SPfCl"[~S.SPECIMENS: ,"hiM.Mexico, CllllllAlltACHIHUAHUA: 48 km~m S of Jiménez,Jim~nel. Cron­e,o,,· quillquist & Fa.'Fay 107JII10759 ('IIC1!.(MICH. 'YoNY. 0<»:OS); 19.5tll.5 mi S of Cd,Cd. Camargo,Camargo. K & MIf 8251tllJI (os).to'». OO~Ht'ILACOAHUILA: .;clll.jctn. ofof rd. to SailSan PedroP~dH> &.& Rte.Rlt. 40.40, K & M.If 79887988(O';f: (os); 10.3 miml S of"r RanchoRanch" loslos Ch.arc",.Charcos, ~K & M 8164 (os):(os); Jim"lco.Jimulco, PringlePril1J;:k l009f10091 (ARll.A"·.(ARIZ. ASU. CAS.CA'. F.F, GH.GII.Ll. LL. MICH,'IiCU \to.MO.NY.PH.SMU.UC. W PH. S\.Il'.' C l').US). IRIlA'!',ODURANGO: 11.5-2217.5-22 mi SW of Nazareno,Natarcno. K & MII 8033,~(}}). 80}8, 8038. 80l/8041 (0');(os); lluar;chi~.Huarichic, Pem'i'llPennell /860}18603 (\.lICit.(MICH. NY."Y. PH).PilI. 7.7- reedsPectis incisifolia I. M. JohnstJohnst.,.. Jour. Arnold Arb. 21: 75. 1940. TYPE: Mexico. COAHUILA: just S of Laguna del Rey, Rey. Johnston 7824 (Holotypc,(Holotype, GH!:GH!; isotypes.isotypes, CAS!.CAS!, LL!,LLI. US!). Figure 8. Strong-scented taprootedtapromed annuals. Stems decumbent to pros-pros· !rate,trate, dichotomously branched,branched. 5-40 cnlcm long.long, glabrous,glabrous. strami­ neous toto red-purple. Leaves linear to elliptic in outline, 1-4 cmem long.long, 2-15 mm wide.wide, irregularly pinnatifid with 2-4 pairs of re-re­ mote,m°te, linear to triangular.triangular, acute, mucronate or bristle-tipped lobeslo°es 1-5 mm long and 1-2 mm wide,Il.·ide. basally incised-ciliate with 2-42-4 pairs of slt:nderslender bristle-tippedbristle-lipped teeth 1-2 mm long,long. glabrous on0n both surfaces.surfaces, marginally punctate with rounded oil glands 0.2-0.4 mm diameter. Peduncles 5-10 mm long, glabrous,glabrous. with 1-3J-3 linear-acute bractco1csbracteoles 10to 2.5 mm long. Heads solitary inin thethe forksforks of the stem.stem, cylindric or campanulatc.campanulate. Phyllaries 8, nar­ ro",l)'rowly linear.linear, 5-6 mm long.long, 0.5-0.7 mm wide,wide. strongl}'strongly convex­cOO\'ex­ keeledKeeled,. b·basallyasa II y gIbbous.gibbous,. very narrowly scartouS-marglflcscarious-margined,.' d , api­ .apl' . callycally narrowed and sllbacutish.subacutish, tipped with a tuft of minute hairs, 64 RhodoraRhoc.lora [Vol.[Vol. 79?9 otherwise glabrous,glabrous. eacheach punctatepunctate withI!dlh aa solitarysolitary roundedroundl"d sub­suI). terminalterminal oil gland caca 0.50.5 mmmm diameterdiameter andand 1-41--4 pairspairs ofof slender,slender. inconspicuousinconspicuous submarginal glands.glands. RayRay floretsnorets 8;8; corollascorollas 4-64-4:1 mm long,long. with\!;ith narrowly narro\!;ly toto broadlybroadly obovateobovate ligulesligules 2-42-4 mmmm long,lon!. glandular-puberulent g[andular'pu~rulent on thethe tubes.tubes. DiscDisc floretsnorets 12-15;12-15; corollascorollas 3-4 mm long,long. regular,regular. withwith teethteeth ca.ca. II mmmm long,long. glandular­glanduLar­ puberulent on thethe tubetube and throat.throat. AchenesAchenes 3.5-53.5 5 mmmm long,long. glandular puberulent.pubC'ruknt. Pappus absent.absent. Chromosome number:number. ntI -,. 12. FloweringFlo\!;l:Ting August toto October.

DISTRIBUTION:OISIRIBL'TIO'· Very locallocal inin southeastern M1uthe:lS!ern Chihuahua andand west­west­ centralccntral CoahuilaCoahUila on sand dunes. Chihuahuan DesertDesc:rt endemicendemic (Figure 5). PectisP~tis incisifoliamciszjoliu appears toto haveha\'e aa relictrelict distribution. This species occursoccun only on the sand dunes which borderborder certain playasplayu or lagunaslaguna~ in thethe interior of thethe Chihuahuan Desert.Desert. Pectisptnis incisifolia il/fiji/olia may have occupied a widerwider rangerange beforebefore post-Pleisto­ cene drought brought about thethe formationformation of thethe largelarge playas.pla)·u. As noted by Johnston (1940),([940). thesethese areas are remarkableremarkable forfor thethe endemism of their floras. PectisPef/is incisifolia incisi/olia is a very distinctive species,species. well differentiated from all other species in sect. PectothrixPt'flOl/'rlJr byb)' itsiu prostrateprostrate habit,habit. pinnatirKlpinnatifid 1C'aves.leaves, and glandular,glandular. completelyeompletely epappose achenes.achents. This taxonIII",on does not appear toto be closely related to any of thethe other species. John~tonJohnston (1940) suggested that P. incisifoliaim:isijoliu might bebe most closclyclosely relatedrelaled to P. ullgustijoliu.angustifolia. AlthoughAllhough II cannotcannol rulerule OUllhatout that possibility.possibility, I fedfeel that a closer relative might be~ P.papposaP.puppoJIJ because of similarities in the phyllary morphology. However,)iowe\"C'f. any relationship of P. incui/oliuincisifolia to 10 either of these species is distant.distant

RIREPRESENTATIVE....r.. I'r~"'1 ""C'I"'['~SPECIMENS: México,\thico. ntIH~~ln,CHIHUAHUA: 9 km NE'E of.... Carillo toward1"",.,.nI GlnmNlne.Guimbalete, C"-'ttt_Chiang, MmJlWendt & JoJm~""Johnston 9OSl111);9052 (LL); 36J6 mi1111 ENE[>.;e of EscalónEKak\Ol alongaloe 1""1trail 10 to ~.Esmerelda, 1I"""d.JO"Henrickson 77661111;7766 (u.); j5 mi11\1 NE'E of Lagunal.apIna Palomas,PIIo-. JohnstonJoIuul•• ~&l~7827 ((,H):(GH); j5 mi E of Canllo.Carillo, Muller",,11;.)) 3317 IllCiIl (GH, 1LL. r. MICH."l1e1l.1 UC). C)_ COAHUILA:("{)~1l1 I~'" .,.lIIifttïà E d.of s..Salinas..nal onon rd.rd. CfCM'''1crossing L>ktlake bed lead'''Il'oleading to La Cllcma.Chemica, 1J«-"1<11Leverich 1221I.'.': In(TEX); \):.s S of !.IfUlIIlLaguna dddel R~.Rey, AK & M116101. 8/01, III()lA.8102A, 6105.8105, 61()9.8109, 6117.8117, 6L'O(0'ir.8120 (os); 6"6 n" ...N ofof 'oOaNoria Mnear ..... lll"Laguna.... d'"del R.Rey,..... Sh,n'f'Shreve 8844II (~~Il.(ARIZ. IUS). ~J_ 8. 8. Pecti)Pectis purpurnpurpurea BrandegBrandeg.,.. Zoe 5: 225. 1905. TYPE: Mexico.;\1uico. SI:,\AlOA·SINALOA: Cofradia.Cofradía, BrUnlWXtt'Brandegee $.n.s.n. (llolot)'pt.(Holotype, UC!.UC!; isotype.isot)pC'· Gll'). GH!). Figure 9­9 19771 19?7] Pectis•Pectis-KeitKeil 65

Plants annual. Stems 15-50 cm tall.tall, one to several from thethe base, erect or ascending, branching above. Leaves 1.5-6 cmem tong.long, 1-9 mm wide, linear to narrowly elliptic or lanceolate,laoceolatc. sometimc.~sometimes strongly revolute, punctate on the undersurface with circular or elliptic oil glands. Peduncles 4-13 cmem long,long. with 4-10 bracteoles, these 1.5-3 mm long, scare-like.scale-like, linear to lanceo­lanceo­ late.late, hyaline except for the darkened midribs, marginally cilio­ late,late, often punctate with 1-2 glands. Heads solitary, terminalterminal or ax.illary,axillary, campanulate. Phyllaries 8 or 9, 4-6.5 mm long,long. nar­ rowlyrowly to broadly angular-obovate, obtuse to acute, with thin,thin. hyaline margins and rounded backs, obtusely keeled belowbclow and basally gibbous, punctate with scattered linear to ovate oil glands, marginally ci[io[ate,ciliolate, glabrous to puberulent on the back. Ray Ooretsflorets 8 or 9:9; corollas 17-11.5 11.5 mm long.long, yellow,yellow. oftenoflen becoming Whitishwhitish wilhwith purple veins when dry, glandular-puberulentglan

DISTRtBUTlON:DISTRIBUTION: Southern Sonora to northern Nayarit. Local-Local­ [Yly common in thorn forcstforest and semi-desertscmi-desert sites (Figure 10). PI'ClisPectis J)urpl/rl'apurpurea is apparently most closely related to P. sinalo­sina/o­ I'nsis.ensis. Unlike most spl'ciesspecies in 5('ct.sect. Pectothrix,Pl'ctolhrix, both of these have glands scalleredscattered on the undersurface of the leaves or inin SUbmarginalsubmarginal rows on the undersurface. In addition,addition, thesethese srt:ciesspecies both ha\'ehave many-floweredmany-Oowerl.'d heads, angular-obovate phyl­phyl­ laneslaries and a biserialCbiseriate disc pappus (except in one variety of P. PI/rpl/rea).Purpurea). Although some forms of P. purpureapurpureo resemble p.p- rushyirusbyi (Keil.(Keil, 1974).1974), theIhe relationship betweenbelween thesethe~c twotwo specie~sPecies is not as doseclose as thatthaI between P. purpureapurpureo and P.P. sinalo-sin%­ fnsis.ensis.

8a.8a PcetisPectis purpurupurpurea Brandeg.,BrarKleg., Zoe 5: 225. 1905.1905. var.var. purpurea.purpurt'a. Figure 9. 6666 RhodoraRhodora IVoi.[Vol. "79

o '" ."\ I

i I AB mm • , , , ~ , , ( , • ~ .. \ / em ( -', '( • .c:G ;;,~":...- ., E F I ,J FilUr~ Figure 9.9 A-F,A-F. PectisPH,i. purpurea;pIN_N, A-B,A·B. var.Waf purpureaP"'PUutI (Gentryj(i

Stems to 50 em.cm, minutely puberulentpuberulcnt on the angles or gla-gla­ brate. Leaves 1.5-4.51.5 4.5 emcm long.long, 1-4 mm broad,broad. acute to10 acumi­ nate.nate, mucronate, ciliate near the base with 1-3I-J pairs of bristles 1-31-3 mm long.long, glabrous.glabrous, punctate with glands 0.4-0.7OA...().7 mm inin diameter scalleredscattered on the undersurface or in submarginal rows.rows. Peduncles 6-11 emcm long.long, g.labrous.glabrous, with 55-7 7 bracteolesbractcoles 1-1.5 mm long.long. Phyllaries 4-4.8 mm long,long. 1-1.3 mm broad,broad. glabrous,glabrous. punctate wilhwith 5-10 glands in two vertical rows or scattered.~allered. Ray corollas 7-8 mm long.long, 1-1.3 mm broad. Disc corollas 20-20 40.40, 3-3.3 mm long. Ray achenes 2.3 3 mm long, strigillose;strigillose: disc achenes 3.3-3.7 mm long.long, strigillose. Pappus of both rayray andand disc eoroniform.coroniform, 0.3 mm long. Chromosome number un­ known. Flowering October to February.

DISTRtIlUTlON:DISTRIBUTION: Known only from the vicinity of the type locality in central Sinaloa al at ca. 200 m elevation (Figure 10).10). . Pectis Pectis purpureopurpurea var. purpureapurpureo has been collected several timestimes in and around Cofradia.Cofradia, Sinaloa since its initial discovery b)'by T.T. S. Brandegee. There is no evidence of intergradation of°f pappus structure between this variety and the other twotwo varie­ ties of P. purpurea,f'urpurel1. such as are found in some other species of°f sect. sect. Peclofhrix.Pectothrix. The very\"Cfy local and isolated range of var. purpurel1Purpurea has apparently allowed this race toto maintain its identity.

REn[s£q~llvFREPRESENTATIVE SPECIMENS.SP'('I\t~'S "'t~;

8b.8D - PtctisPectis purpurellpurpurea Brandeg. var. sonorasonorae.. Keil,Keit Brittonia 26: 35. 1974,1974. TYPE: l\1rxico.Mexico. SO:-;ORA:SONORA: 1.9 mi NW of Vicam on Mexico Rte 15, ca. 66 mi SE of Guaymas,Guaymas. Canne & KeilKeil 8644 (Holotype.(Holotype, US!; isotypes,isotypcs. ARIZ!, ASU!,ASU!. OFS!.DES!, DS!,OS!, ENCB!,E....CB!. p F',!. GH!, MICII'.MICH!, :'>10!.MO!, NY'.NY!, OS!, RSA!, SO!.SD!. SMU!. TEX!.TEX!, UC!).ue'). Figure 9. Stems to 50 em,cm, glabrous to densely puberulent.puberulenl. Leaves 1.5­ 6 emcm long.long, 2.5-9 mm mm broad, obtuse to acute,acute. mucronate,mucronate. ciliate f\Carnear thethe base with 1-3 pairs of bristles 1-2.4 mm long,long. glabrous toJ° densely puberulent on the upper surfaces, glabrous or puberu­ ~ent. nt Onon the midribs below,below. punctate with glands 0.4-0.70.4...(),7 mm In 'n diameter in SUbmarginalsubmarginal rows on the undersurface. Peduncles 4-1313 cm long, glabrous to densely puberulent,puberulent. with 4-10 brae­brac­ 6868 RhodoraRhodora [Vol.[Vol. 7979 teolesteoles ca.C3. 22 mmrom long.long. PhyllariesPhyllaries 5-6.55-6.5 mmmm long,long. 1.8-31.8-3 mmmm wide,....'ide. glabrousglabrous oror puberulent,puberulent. punctatepunctate withwith 4-124-12 scatteredscattered glands.glands. RayRay corollas 9-11.5 mmmm long,long, 22-3 3 mmmm wide...... ide. DiscDisc floretsnorelS 40-50,40-50. theIhe corollascoroll,l5 44-5 5 mmmm long.long. RayRay and disc achenesacllenes bothboth 3-43-4 mmmm long,long. pilose or strigillose.strigillaSC'. PappusPappus of thethe raysrays biaristate,bianstate. 2-32-3 mm long,long, with.... ith 1-41-4 shortshort scalessca'es betweenbet.....een thethe awnsawns onon thethe sideside adjacent to thethe: subtending5ubtending phyllary;phyllary; pappuspappus ofof thethe discdisc biseri­biseri· ate,ale. thethe innerinner series of 5-10 stiff bristlesbristles 44 5S mmmm long,long. thethe outerouter series _of 8-208 2{l shorter, more slenderslender bristles.bristles. Chromosome number,number. n,. -= 12. FloweringFlo.....ering August andand September.

DISTRIBUTION:D1SlIURUT10S. Southern Sonora andand northernnorthern Sinaloa alongalong the coastal plain (Figure 10).

PectisPUlis purpureaplI'pur~" var.\"3T. sonoraewnoroe isis thethe most widespread\\idespread and most variable of thethe threethree varieties of P.P. purpurea.pmpurea. The pubes-pubeS­ cence of the plants of var. sonoraesonoroe rangesranges fromfrom subglabrous toto densely puberulentpubcrulent within individual populations.populations. The plants of this variety"ariety appear to be somewhat weedyweedy andand are often locally abundant. Some of thethe areas where thisIhis variety isis foundfound are heavily browsed,browsed. but P. purpureapurpurl!(J does notnor appear toto bebe touched. Perhaps the strongly scentedseemed foliagefoliage serves as a de-de­ terrent to10 browsing as the oils havehave a very\'ery disagreeable odor resembling that of stinkbugs (Pentatomidae).

REPRESENTATIVER,,.,,,,,... ,,,, SPECIMENS:~",,,,..t" 5Cesee KeilKei] (19"')(1974) forfo< specimen~pccill"Ol:n citationscltallon. forfo< Pectisf't

hPectis..... $I_lorsinaloemis..... remFern. ,ar,var. ItlnrrfolllJlancifolia G<< R)-.lbRydb.,..... N. Am. FlFl. 34:U: 203. 1916.1916

SiemsStems to 40 em.cm, hinellou$hirtellous on thethe' angles. angles. LeavesLea\'l:$ 2-3 cm long,long. 3-63-6 mm broad.broad, acute.acute, mucronate.mucronate, ciliate along thethe margins wilhwith 44-6 6 pairJpairs ofof bristles 2-3 mm long.long, densel)'densely shonshort puberu­ lentlent on lhethe upper surfaCC'$.surfaces, hinelloushirtellous along the midribs beneath,beneath. punctate wilhwith glands ca.ca. 0.25 mm diameter both submarginal andand scalleredscattered on thethe undersurface. Peduncles 77-1212 emcm long,long. 1971J1977] Pectis•Pcctis-KeilKeil 69 short pubcrulenl.puberulent, with 4-84 8 bnlcteo1csbracteoles ca. 3 mm long. Phyl­ brieslaries 5-5.5 mm long, 1.6-2 mm broad, glabrous or minutely pubcrulem,puberulent, punctatcpunctate with 4-6 scattcrcdscattered glands. Ray corollas 9-10 rnmmm long.long, 2-3 mm wide. Disc[Jisc florets!lorets ca. 40, thethc corollas 4.5-4.8 mm long. Ray and disc aehcnesachenes both ca. 3.5 mm long, strigillosc.strigillose. Pappus of the rays biaristatc,biaristate, 3-4 mm long, with",ith ca,ca. 8 shortshort scales betweenbetwcen the awns on the side adjacent toto thethe subtending phyllary; pappus of thethc disc biseriate,biscriatc, the innerinncr series of 4-6 stiff bristlcsbristles ca. 6 mm long, the outer scricsseries of 8­ 16 16 Ilallened,flattened, fimbriate pales to 2 mm long. Chromosome num-num­ b<'rber unknown.

DISTI(IRUTIO:";DISTRIBUTION: Known only from the type collection in nonhcrnnorthern Nayarit (Figure 10).

Pem'sPectis purpurea var.vaL lanci/olialancifolia is somewhat of an enigma. Known only from its Iypetype collection.collection, this taxon is clearly a mem-mcm­ b-erber of the P. purpureapllfpUrl'u complex. It is far removedremo\'ed geographically,gcognlphically, however, from the nearest population of var. sonorae,.!Onorat', which it mostmost closely resembles. PeclisPectis purpureapurpureu var. lancifolia!unci/olia may represent a relict population of a once more "idcspreadwidespread P. pur-pur­ purea, or conversely, may be a pioneer population. In either caSt,ca se, irsits present status is rather uncertain, because extensiveextensi\'c modification of thcthe habitat has occurred in the region where the t}petype collection was made:made; in fact, thistbis variety may wellwcll be extinc!.extinct.

9. 9- PeetisPectis sinaloensis Fern.,Fern .. Proc.Proe. Am. Acad. 33: 69. 1897. TYPE:TVPF: Mnieo,Mexico. SI:"AlOAoSINALOA: Mazatlan,Mazatlán, WriKhlWright 1204 (Lectotype (here Chosen),chosen), Gil!:GH!; isolectotypes,iso!ectotypes, F!, MO!,\10', MSC!.MSC!, US!).US'). Figure 9. b Unscented, tap-rooted annuals.annuals, usually branching from the a.l.e.ase. Stems 10-30 cm long.long, erect, or more commonly decumbent,decumbent. branchingranching above, short villous\'it!ous in lineslincs or glabrate. LcavesLeaves linear-linear­ lanceolate,anceolate, elliptic or or oblanceolale,oblanceolate, 1-5 emcm long, 5-12 mm wide,widc, ~b.tuse? . tUse to acutish, mucronatc.mucronate, basally ciliate with 2-6 pairs of nst1esnstles 1-3U3 mm long,jong glabrousgiabrous or pupubcrulentberulent at the base, punctate On"thethe Undersurfaceundersurface with numerous scattered round oil glands. PedunCles^uncles 2-6 cm long,long. glabrous, with 4-6 scale-like bracteolesbracteolc.~ ,1~2 mm long. Heads terminal and axillary.axillary, few. Phyllaries 8 (rarelyrarely 9 Or°r 10), oblan'eolate.oblanceolate, S5 mm long,long. 1.5 mm wide, rounded 7070 RhodoraRhodora [Vol.(Vol. 7919 to10 subacute,subacmc. broadlybroadly hyaline-margined,hyaline-margined, dorsallydorsally rounded,rounded. basallybasally gibbous,gibbous. apically short-ciliolate,short-ciliolatc, otherwiseotherwise glabrous,glabrous, punctatepunctate with\lith 6-12 scatteredscallerro ellipticaldliplical oiloil glands.glands. RayRay floretsflorelS 8-10;8-10: corollascorollas 55-7 7 mm long,long, withwith narrowlynarrowly toto broadlybroadly ovateo\'ate: glabrousglabrous ligulesliguks and sparsely glandular-puberulentglandular-puberuknl tubes.lUbeS. DiscDisc flowersnOIlo"cr5 25-60;25-60: corollas 4-5 mmmm long,long, bilabiate,bilabial!:, withwith thethe narrownarrow lobelobe ca.ca. 22 mmrom long,long. two1.... 0 or threethree timeslimes thethe lengthlength of thethe teethteeth on thethe broadbroad lobe,lobt'. sparsely glandular-puberulent. Achenes 2-32-3 mmmm long,long, thosethose ofof the disc sparselysparsel) strigillose allall over,o\'cr, thosethose of theIhe raysrays glabrousglabr()lrt on thethe side adjacent toto thethe subtending phyllary,phyllary. otherwiseotherwrsc denselydenstl~ strigillose. Pappus of theIhe raysra)"s biaristate,biaTislale, 2-32 3 mmmm long,long. withwith aa single fimbriatefimbriate: scale betweenbct\\-ecn thethe bristlesbristles on thethe side adjacent toto the subtending phyllary. Pappus of thethe disc biseriate,biseriate. of 5 stiff,sliff, antrorsely scabrid~cabrid bristlesbristks 3-3.53 3.5 mmmOl long,long, alternatingallcmating withwith 55 fimbriate acute toto bristle-tippedbristle-lipped scales 1-2 mmmOl long.long. Chromo­ some number: nII -= 12.12, Flowering September to10 January.

DISTRIBUTION:D1STR1DUTIO:-l; Known only fromfrom thethe coast of Sinaloa atat a few metersmelers elevationelcvation (Figure 10).(0). This speciesspccies isis particularly common on thethe sea-cliffssca-eliffs at Mazatlán,Malatlan, thethc typetypc locality.localily.

At Altata,Altala, Sinaloa,Sinaloa. PectisPeOiJ sinaloensis has formed an extensiveextenSI\'e hybrid swarm with PectisP{'ois papposapappOJQ var.vaT. papposa.pappusa, T.T, S.S, BrandegeeBrandes« described PectisPutis solillOsalina from thisIhis population.population, The typestypes of ?•P. salina are intermediate between P.P, sinaloensissina/~nJis and P. papposa.papplJso The hybridsh)brids in this population are apparentlyapparenlly fertile,fertile. and no detec1abledetectable chromosomal irregularitiesirregUlarities were detecteddele<:led inin any olof the membenmembers of the populationpopulalion that II examined cytologically-C)'101ogicallY· PlantsPlan15 ....with;th \'aT)'ingvarying degrC"C'sdegrees of intermediate characters are found in the populalionpopulation with"'ith C'Xtremeextreme forms approaching both parents.parent1­ Because of its origin, therefore, thisthis hybrid population is to bebr lno""nknown as PectisP('('tis X salina Brandegee,Braooegec. pro sp. The plants in the P('('tisPectis sinaloensissillalot'flsis population at Altata pos-pos­ sesssess some""h.atsomewhat largerlarger heads than those atal Mantlan.Mazatlán. This featurefeatUrt ma)may ""ellwell be a result of instability introduced intoinlO the P.P, sina/«n­sinaloen­ sissis population b)'by the 10ng-l((mlong-term hybridizationh)'bridiTation with P.P, papposa.poppaso If thisthis ISis thethe case.case, Iherethere rna)may nOInot be any ~pure~"pure" P. sinaltWnJissinaloensis alat Altata.Altata, onl)only extremesextremes of the hybrid swarm.

Rr'.''''I~!IHREPRESENTATIVE ~P.cr".,~SPECIMENS: .\1"I«t,México, '1"1("SINALOA: Mazatlán,\Ia,allon. AK &-& C UJ108810 (()'o);(os): ULa r..H'.a,"Nevería, N ofof \IualJ3n,Mazatlán, MexiaI/ula /()9.(¡094 (~,(CAS. >.1,11F. GH. "1(11MICH. ..MO./0 ,~_NY. 1US).'I 1977]1977/ Pectis•Pectis-KCtlKeil 7171

• r­ : I, , i ,· i •

'<.• ( ,

, ~ f'­ ~ f.. \ PeetisPectis ppurpureaurea \~ T •var.var. purpurea Peelis fi pes o var. filipes • var. lancifolia x\ •var. lancifolia • var. subnuda • var. sonorae . Pectis vollmeri o PeetisPectis sinaloensis

Fi,u.~ 10 G h' f iii igure 10.. Geographical(o,.ap ..al distributiondi,l"bulion of0 . Pectisp,,,,, purpurea,1""1""Nl. P.1'. sinaloensis,JI""I"',,'i,'. P.P. Jupes/ 'Pn .ndand P.P ''OIlmni.vollmeri. Half-barredIblf.ba.ted line indicates",,;hC.lCl Sonoran Desert~n boundary (redrawn'edl1l"'n from Sh.ew.Shreve, 1942).

10. 10- Pectisr«tis filiJn5filipes Harv. &:& Gray in A. Gray, Mem. Am.Am.Acad.~: Acad. 4: 62.62­ 1849. TYPE: Mexico.l\Iu:ico. '"CALlFOR'IA.-"CALIFORNIA," (probably collectedeolleaed inin Sonora near Guaymas or Hermosillo),lIermosillo). CoulterCouflt'r 129329 (Halo­(Holo- type.type, Teo,:TCD!; isOlypes,isotypes, GH (fragment)'.(fragment)!, K!).Ie!). Figure II. S Sirong-scenied.Strong-scented, diffusely branched tap-rootedlap-rool«l annuals. Stems c -4() emm long.long, slender.slender, pscudodichotomouslypseudodichotomously branched,branehro. glabrous ~s~uberulemPuberulent... Leaves linear 10to narrO'ol.lynarrowly elliptic.elliptic, 1-6 .emcm long,lo~g. I. . .S."5 mm wIde.wide, often Stronglystrongly fC\oIUle.revolute, basally ("lhaleciliate with... lth -:eP-aPairs11'5 of briStlesbristles 1-2I 2 mm long.long, othcr'ol.iseotherwise entire, glabrous or ~ erulent,ndent, marginally punctate 'ol.'fthwith rounded oil glands ca. o'^ mm diameter. Peduncles filiform.filiform, I1-6.5 6.S emcm long.long, glabrousglabrous puberu t:te tllberulenl,lent, with 1-3.1-3, usually subterminal, lanceolate brae­brac­ e ca ey~i:r ^ ~.- I* mm long.long. Heads solitarysolitary in the forks of the stem.stem, lndricI'1C 10to narrowly Campanulate.campanulate. PhyllariesPhyllarits S,5, narrolOolynarrowly oblong 7272 RhodoraRhodora [Vol.IVo1. 7979 toto oblong-obovateoblong-ooov3le 3.5-63.5-6 mmmm high,high. ca.ca. 1I mmmm broad,broad, basallybasally trun­trun­ cate and gibbous,gibbous. apicallyapically ciliolate,cil10lale, obtuseOblUSC to10 acute,acute. hyalinehyaline mar­mar­ gined,gined. convex and slender-keeled,sknder-keekd, glabrousglabrous oror puberulent, pubcrulcnl. punctatepunctate with\lilh 0-2 smallsmall to10 largelarge subterminal glandsglands andand 2-32 3 pairspairs ofofsmallcT, smaller, rounded to10 elongatedelongated submarginalsubmarginal glands. RayRay floretsnouts 5;5; corollasoorolbs 4-9 mmmm long,long. with",ith narrowlynarro.....ly to10 broadlybroadly obovateObo\'aIC glabrousglabrous ligulesliguks andand glandular-puberulentgbodular-pubcrulc:nl tubes.lUbes. DiscDisc floretsOOTets 2-22;2-22; corollascorollas 2.5-42.5-.4 mmmm long,long. bilabiate,bilabl3lc. with thethe narrownarrow lobelobe ca.ca. twicetwice thethe lengthlength ofof thethe teethI~h on thethe broad lobe,lobe. sparsely to10 densely glandular-puberu­glandular+pubcru­ lent.lent. Achenes 2.5-4 mm long,long. shortershor1er thanthan thethe phyllaries,ph)ILaries. strigil­ loselose or shortshan pilosepilose withwith bifurcatebifurcate hairs.hairs. PappusPappus 0-3 aristate,ari5tatc:. antrorselyantrorsel) barbedbarbed to nearlyncarly smooth,smooth. usually with",-jlh an additional low10.... crown ca. 0.2 mm high. Chromosome number:number: nII ­= 12.12.

DISTRIBUTION:D1STklHUlIO",' Southeastern Arizona and westernweSlern Texas to10 southern Sonora and southern Chihuahua at elevationsdevations of 0-2000 meters (Figure 10). Within thethe broad altitudinal tolerancestolerances of PectisPeclis jilipt!5.filipes, two varieties may behe distinguished. The typicaltypical va-va­ riety has a primarily Sonoran DesertDesert distribution,distribution. occurring mostlymostly below 500 meters elevation. The variety subnudaJllhflllda grows mostly at higher elevations and barely reachesreaches thethe fringes of thethe Sonoran Desert. This latter variety occurs mostly inin the grassland andand pine-oak-juniper zones70nes and extendsextertc/is filipesjifip'J isis relathelyrelatively common, there is lntlelittle e"idC'nceevidence to suggeslsuggest thatIhat the effeclseffects of the crossing are more than locallocal. ThroughoulThroughout most of ilsits range.range, P. fi/ifNsfilipes remains well differentiated from other specieSspecies inin the section.section. AIAt Ihethe local populational level,level. some bizarre re­~ combination types have been obsenedobserved which combine the fea­ tures of two or more species,species. 1977]1917] Pectis•KeilPectis-Keil 737J

, E

em

K L -­mm

lgU ll A H ,:'::~.var r "." A·H." ' Premfilipn**ctis filipts; A·D.A-D, 'a.var.ßüpes filll"'J fA,J(Keil ..& ell"'"Canne 8659.8659, 00;);os); E-H, Uda {KeÍl & McGiU 85U os l L Pec,is vollmeri C~;)CAS) AA ItUdIl (1."/,, I/eG"I.' 8J1!.- (5);); 1-1..' ' ,,"liS .'011",", fC",,"(Carter ..& MoranVO'iUI 5]50,5250, 1 A . >,F. E, I,I, habll;habit; 8.B, F, J.J, jlllyllana:phyllaries; C.C, G.G, K, d,s<:disc acMI\C';achenes; D.D, II,H, l.L, rayrayachona. achenes. 7474 RhodoraRhodonl [Vol.[Vol. 7919

10a.lOa. PectisI'celis filipesfilipes Harv.lIarv. && GrayGray inin A.A. Gray,Gray. Mem.Mem. Am.Am. Acad.Acad. 4:62...: 62. 1849. var.\"aT. filipes.filjp". FigureFigun: 11.11. HerbageHerbage densely puberulent.puberuknt. LeavesLeavcs linear,linear. 1-31-3 cmem long,Ion!. 0.5-2.5 mmmm wide. FasciclesFascicles ofof severalseveral shortsnon narrownarrow leavesIea\'cs usuallyusuall} developeddc\'eloped inin upperuppl:r axils.axils. PedunclesPeduncles 1-2.5I 2.5 (-3.5)(-3.5) cmem long.long. In-In­ volucre cylindric;C}lindri~ phyllariesphyllarics acute,acute. lackinglacking subterminalsubterminal oiloil glands.glands. Disc floretsnorets 2-8; corollascorolla~ 2.5-3.52.5 3.5 mmmm long.lon!_ OilsOils of plantsplants fromfrom southern Sonora lemon-scented,kmon-M:l:nled. from(rom other populationspopulations spicy-SPIC)­ scented. Chromosome number: «=12.n= 12. FloweringFlowering AugustAugus! toto October. DISTRIBUTION:DISTRIRUTlO' Central and southern Sonora,Sonora. mostlymostly belowbelo\\ 500 meters elevationclcutian (Figure 10).

Coulter's typetype localitylocality was listedlisted as "California,""California." a statestatc where.....hert PectisPeelis filipesjifipt'J isis not known to occur. RydbergRrdbcrg (1916)(1916) suggestedsuggested thatthaI this species was collected by Coulter inin Arizona.Arilona. CoultersCoulter's itinerary in ArizonaAri70na (Coville,(Coville. 1895), however,however. did notnOI includeinclude any areas where P. filipesjilipl'$ grows.grows. Upon examination of thethe typelYpe of P. filipes,jilipt>s. II found that itit very closely resembles rcsembles thethe Sonoran race of the species. Coulter is known toto have collected ata\ both GuaymasGuayma~ and Hermosillo,Hermosillo. Sonora (Coville,(Coville. 1895; McVaugh,McVaugh. 1943).1943), and therefore it is likely lik.ely that thethe type collection of P.p.filipn filipes came from this region.

RIH.n.'~r~",.REPRESENTATIVE ~"'-{1"f"SPECIMENS: "'I"iro.México, SONORA:"""011' 282 mi S of Hermosillo.Iltnnooillo. K/<.. &&. C MJI18617 10'-1:(os); Il.K11.8 m,mi E of jctn.jcr..... w/Mexico \Iuiro 15 on rd.rd to10 Alamos,"I.on..... K &<4 C 1I!l658665 (os);to'l~ 14 ....mi 'iS of Divisaderos,O',~de,~ U'ui",Wiggins U~97469 f(ARIZ,.. ~". 1>"DS...... MO, TEX,'" US).l~)

lOb. 10b. PectisPteti~ fili~filipes Harv. &.& Gray in A. Gray var.\'ar. subnud.subnuda Fern.,Fern.. Proc. Am. Acad. 33: 76. 1897. nTYPE: Pf. Mexico.Muieo. CHIHUAHUA:ClllHUAllUA Janos.Janos, A. SchouSchott s.n.s.n. (l..ci.:Iotype(h¢re(Lectotype(herechosen), chosen),GIl!).GH'.). Figure 11.I', Herbage glabrous 10to moderately puberulent.puberulenl. LeavesLea\'cs linearliflC1lr to narronarrowly.... ly elliptic, 1-6 cm long.long, 1-5.5 mm wide. PedunclesPedufK"k:! (1-)(1-) 2-6,52-6.5 emcm long. Involucres cylindrkcylindric 10to campanulate; phyl­ph)1­ larieslaries obtuse 10to acute, ....withilh or \loilhoUIwithout subterminal oil glands. Disc Ooretsflorets 7-22; corollas 3-4 mm long. Oils spicy-scented butbUI nolnot lemon-scented,lemon-scented. Chromosome number:number, n :- 12. Flo\!."(:ringFlowering July 10to No\ember.November. 197711977] Peelis-KeilPectis• Keil 75" DIS1RIRUTlO:-.':DISTRIBUTION: SoutheaslernSoutheastern ArilonaArizona and weslernwestern Texas to10 northeastern SOnoraSonora and southern Chihuahua, mostly atal 1000­ 3000 meters elevation (Figure 10). The presentpresenl treatment of PectisPeflis filipesfilipes var. Sllhlludasubnuda as a wide-wide­ spll:adspread eeo,geographicaleco-geographical race differs markedly from thelhe much more restricted concept of Fernald (1897),(1897). who considered var. sllbnudasubnuda 10to be merely a minor morphological variant in thelhe much more widespread species, P. filipes. As the epithet.epithet, sUbllutla.subnuda, is available in the varietal rank and my concept of thethc upper elcva,eleva­ liontion race of P. filipesjilipes includes thelhe types of var. subnudaslIhlluda I have necessarily used lhisthis epithet for the redefined variety.''ariety.

RHKfSf'1.nl\lfREPRESENTATIVE ~Pt("l\lE"SPECIMENS: México,Mhie... CltII",AIlUACHIHUAHUA: 10-1410 14 mi W of Hidalgo/lld.lgo del Parral.Parral, ut"'",Gentry 8:578257 ("11.(GH, MICH,\lKIl. lfS):US); S.45.4 mi N/'.. of Janos.Janos, KA & MIf UN8414 (os); Mts.M,s. NW/'.·w of Cllihu.hua,Chihuahua, LeSueuruS"tu' 10/1}1019 (CAS. F. GH. \10.MO, TEX,1F~ lC).UC). SONORA:MJ'ORA GuadelupeGuadelulX Can'Can­ yon.yon, MearnslIt"",. & .lIntorr!048Merton 2048 (1)5.(DS, '<.States. ARI70~AARIZONA. Coch,S(Cochise Co.: ChiriealunChiricahua Mts.,MIS.. BlumerBlumtr 1708 (ARI/.'K.(ARIZ. DS. r.F. Gil,GH. KSC.KSC, "10,MO, '. c.lt.GH. MICH,",CIt. MO,MO. NY.'<1. PH.p\l 'S).us). 'EwNEW MlMEXICO: 'KO G",n,Grant Co.;Co.: 18 mim; NW olSil,·c.of Silver Cily.City, MetcalfeII"lfolf"68S 685 (ARIZ.(·'Rlf MO.ND-G,YO. ,,}-,; NAH'''C, 'YoNY. PO'!.POM, lC);uc); Burro MtsMts.,.. RusbyHu.,") 18$185 (0.(F. "ICII.MICH. MO."O. ,<\'NY, PIIPH. UC.l'C.lS); US); Corptrmin.Coppermine C'eek.Creek, WriglrtWright 11151125 (HI.(GH. 1::.K. MO.MU. 'YoNY, rHoPH, TCD.'0>.' UC. C.l'); US); SienaSierra Co.,Co.: Black Range,RanFt. Met-.Ift'· c CQlftalfe Ij!J71357 «·AS.(CAS. >,GlT.F. GH. MO,'1O. ,,,cNMC... NY.'Y. POM.PO", UC).l'('j, TEXAS.,nA' BrewsterSr<:\O."c' Co.: Bj'Big BendSend Nat'l."aT'l p Parl.ark, Co",,11Correll &;&. Co""11Correll 35379j5j71} (''.SD. TEX.,n. ~s);US); 15.1IS.I mim;" N of Castolon.Cam,lon. }(K &:& .IIM 7797 (os). It.U. Pectis \'ollmtrivollmeri Wiggins, Contr. Dudley Herb. 4: 27. 1950.19SO. TYPE: Muico,Mexico. BAJA CAUFOIlSIACALIFORNIA SUR: foot of Coyote Grade 20 mi S of Mulcge,Mulegé, WigginsWi~gi11S 11407 (Holotype, DS!;os'; iSotypes,isotypes, CAS!, OS!.DS!, Gil'.GH!, Ut'!.UC!, IJS!).US!). Figure 11.J I. Slrong-scenled.Strong-scented, tap-rootedlap-rooted annuals. Stems 5-SO5-50 cmem long,long. ereClerect to decumbent.decumbent, pseudodichotomously branched,branched. glabrous, pUrplish. Purplish. Leaves linear.linear, 1-6 cmem long, 1-2 mmmOl broad,broad. often revo­ l~le,ke, basally ciliate Wilhwith 2-3 pairs of bristles 1-2I 2 mmmOl long,long. rarelyrarely WIthWlth 1-3 short basal lobes, Olherwiseotherwise cmire.entire, glabrous,glabrous. marginally PUnctalePunctate with round to elliplicelliptic oil glands ca. 0.3OJ mmmOl diameter. PedunclesP~duncles 3-8 emcm long.long, glabrous, withwilh 2-3 scattered, linear acu­ minate glabrous bracteolesbraeteoles 1-2 mmmOl long. Heads solitary inin the forks°rks of the stem. Phyllaries 5, narrowlynanowly obovate, 6-8 mmmOl long,long. l.5-2.5•5-2.5 mOlmm broad, apically obtuse.obtuse, ciliolate,eiliolate, basally truncate and 76 RhodoraRhodora [Vol.[V'" 791'9 gibbous,gibbous. hyaline-margined,hyaline-margined, convexcon\'ex onon thethe back,back. round-keeledround-keded below,below, glabrous,glabrous. punctatepunctate withwith 0-30-3 small,small, roundedrounded subterminalsubterminal oil glands,glands. and 3-73-7 pairspairs of narrownarrow scatteredscattered submarginalsubmarginal glands.glands. RayROIl' floretsOoren 5;s: corollas 6-10 mmmm long,long. withwith broadlybroadly ovateovate glabrousglabrous ligulesligules and glandular-puberulent tubes.tubes. DiscDisc floretsflorets 9-19;9-19: corol-corol­ laslas 4-5 mmmm long,long. weaklyweakly bilabiatebilabiate withwith thethe narrownarrow lobelobe ca.ca. twicetwi« as longIons as thethe teethteeth on thethe broadbroad lobe,lobe:. glandular-puberulentglandular·pubc:rulent onon thethe tubetube: and throat.throat. Achenes 4-6 mmmm long,long.. cylindric,cylindric. strigilloseslrigil1()S( to short pilose. PappusPappus 2-62-6 aristate;anstate; awnsalA ns stiff,stiff, spreading,spreading. 3-43 4 mmmm long,long. purplish,purplish. antrorselyanlTorsely scabrid towardtoward thethe tips.tips. ChromosomeChromosome number unknown.unknov.n. Flowering September to\0 NovemberNovember andand againagain from FebruaryFebruary toto April.

DISTRIBUTION:DIS·IRJ8UTIO~. Endemic toto a narrowIlarrow strip along thethe easternea.m·rlJ coast of Baja California at elevationsc1e\'ations of 0-500 meters (Figure(Figure 10).10). PecPt'Cli$ iis vollmermllml'ri i appears to be most closely relatedrelated toto P.P, filipes.fl1iptJ. Both speciesspecIes share thethe characteristics of slender,slender, diffuse habit,habit, five-rayed heads, and aristatearistale pappus. PectisI7cliJ vollmeri,I·ollmui. however, is somewhat stouter and has larger headsheads and flowers.l1owers. Wiggins (1950) suggested thatthat P. vollmeriI'vllmeri waswas closely relatedrelated to10 P.P. linifoliaIinijoJio L. of sect. Pectidium.l'ecridiulIl. Although both species do havehave similar pappus structure,StrUCture. the two taxa differ greatly inin size of thethe florets,norris. number of disc florets,l1orets. placement and sizesile of foliarfoliar oil glands, glandularilalionglandularization of the discdi~c corollas and inin overall0\era1l habit.habil. InIn mym} opinion.opinion, the relationship betweenbet"ccn P. vollmeri\·oJ/m,.,i and P. linifolia lilli/olia isis distant.distant, and the similarities in pappus structurestructure: are thethe resultmull of parallel evolution rather than of patristic relationship.

I/:REPRESENTATIVEFI'-.r~, H,1I\ [ sna"r's.cSPECIMENS: Mhito.México, BAJAaVA CALIFORNIA(,"'II~(HI;"1< SUR:~I ~ 40.6 mi S of Mulegé.\l"IrIi, Cllrlrr.Carter, AItAlexander..",,*, & Al'lloaKellogg J'MI1988 (O!..(DS, GilGH. us);I'); SWS ..... of SapiaSanta Rosalia,Rooalia. C,,""Carter &&. Arlon./(\9:Kellogg 3082 leA,(CAS. GilG H. \toMO, <;D.SD. IUS);"I; RanchoRallCho Aguajito,""..alii.:>. CarterC""rr,ll & MoranIf",." 5250j!.VlI<" (CAS. ~/)SD. ,('uc. •us); "I; CO)O<.Coyote &Y.Bay, GmtGentry... WM4068 haIL(ARIZ. DS.D\ ""GH. MO."0.\<'. vc. us);I ,j: Arroyo"fl'O)O de SanS,n JoséJ_ d~de \badalt-na.Magdalena, "~Wiggins.... 11J.J611386 ID'\...(DS, GilGH...... uc. Ius). "I.

OO1.8TFUlDOUBTFUL "'\0AND [XClLOEOEXCLUDED TAXA PKtkPectis aqu~licaaquatica S. WatsWats.,.• PrIX.Proa Am. Aead.Acad. 23:B: 279. 1888. T'lPETYPE: Mt~ico.Mexico. ClllIIUAlIU"CHIHUAHUA: at base of Sierra Madre (S of Cd. Guerrero).Guerrero), PringleP'",gl,. 1296 (llolot)pe.(Holotype, GIl';GH!; isotypes, Ft.F!. MO!).\10'). =Hydropectis Iquatic:aaquatica (5.(S. Wats.) Rydb. PtctisPectis fuciculalafasciculata Poir. inin Lam. &:& Poir., Encycl.Eneye!. Method. Bot. 5:~: 120. 1804.J804. TYPE;TYPE: grown from seed of unknown origin (nO(no typetype specimensspecimens loeattd).located). ThISThis species is indeterminable. 1977)•977] Pectis-Pectis•KeilKeil 777J

Pect~Pectis mulliradilltamultiradiata Sesse & Mocino.Mocino, Fl.H Mex. cd.ed. 2. 190. 1894. TYPf:TYPE: Mexico. SI"AIOA:SINALOA: prope Oppidum (no type speci-speci­ mens located). This species is indeterminable.

ACKSOWLfDGMENTSACKNOWLEDGMENTS This study isis based in part upon a dissertation presented in partial fulfillment of the requirements for the Ph.D. degree at the Ohio State University. Field studies were supported by National Science Foundation Grant GB-J0240.GB-30240. II wish to thank LV.Dr. TooTod F. Stuessy and Dr. Donald J. Pinkavapinka\'a for critically re­rc­ viNingviewing the manuscript. Thanks are also extended to Miss Judith M. Canne and Mr. Lyle A. McGill for their assistance during the field studies. LITERATURE CITEDClrFO

III,..o,Kf.BLAKE. S. 1'.F. 1945. A'lera<;eaeAsteraceae deS\:r;beddescribed from MexicoMe~ico and thelhe southwesternSOuth"nlern UniledUnited STalesStates by M. E. Jones.Jones, 19081908-1935. 1935. COnlr,Contr. U.1I. S.S.l"all.lIerb. Nati. Herb. 29:117-137.29: 117 137. B~"'OlH,BRADLEY, C. EE.,.. &:& A. J. 11""<;FHAAGEN-SMIT...·$'''r. 1949. The e,,,,,nlialessential oils0;1. of"f Pectisp~(/i,·pQJ>­ pap- f'OJQ.posa. Ecrn>,Econ. Bot. J:3: 4il7-412.407-412. G.'DOllf.CANDOLLE, A. P. DE.nE 1836. Prodromus S)'OlemaTi.systematis naturisnaluris regniregn; vegetabilis,,egelabili•• •siive.. "nUmeral;Oenumeratio conlraClacontracta ordmun,ordinum genegenerum.rum. spcc;erumquespecierumque plantarum.planlarum. vol.'01 S.5- TreulldTreuttel & Wiirtz,WUrlz. Paris. CO.UlICORRELL,.. D. SS.,.. 8.:& M,M. C. JOH"'ro".JOHNSTON. 1970. Manual "fof theIhe vascular,'asrolar plants of TnaTexas.•. Te~hTexas RC!;earchResearch Foundation,Foundali"n. Renner. eoOY.CORY, V. L. 1937.1937. Some M"new plantsplanls fromfrom TexaTexas.•. Rhodora 39:39; 417-123.417--423 eo\ill~,COVILLE, f.F. V. 1895. The botanical exploraTionsexplorations of,,{Thoma. Thomas CoulterCouller inin Mexico and California. Bot. Gar.Gaz. (Crawfords,;lle)(Crawfordsville) 20: 519-531.519-S31. fuFERNALD,..... ln. .'.1,M. l.L. 1897.1897. A 'y>l1.Bot. Club n:73: 451 462. 462. AV G~~>.. A. 1849.1849 piPlanTaeantae FendkrianacFendlerianae ;"o,;.Mu;canac,Novi-Mexicanae. Mem.\lenL Am.'\nT. Acad.,kad 4:,(, 11-116. 116, _." 1852. PlantaePiantae Wrightianae TTexano-Neo-Mexicanae...ano."~o.Me~icanae, I.l. Sn"lhsonianSmithsonian COnic.c°ntr. KnowLKnowl. 3(5): 1-146.I 146. -~"~ 1884. Conlribution,Contributions 10to North American botany.bG.0*0, C. F. 1830. De synanThe,c;,synanthereis herbarii regnireg", Berolinensis.Ke.oli.... nsis. DissertatioDi.sertalio secunda.secunda. Nassau'~e.Nassauvieae. Linnaea 5: I1-236. 236. _.- 1831.'831. D~De S)synanthereis. nanthcreis. l)'ssertal;oDissertatio qua"quarta.•. Linnaealinnaea 6: 624624-721. 721 78 RhodoraRhodora [Vol.[\'01.79 79

MCVAUGH,MeV",

DEPARTMENTDEPART\lF' r OFOf BOTANY8OTA:

Present address: BIOLOGICAL8101 OGleAt. SCIENCESSCIE~CES DEPARTMENTDEPARTM"....T CALIFORNIA POLYTECHNICI'OLYTFCIINIC STATE UNIVERSITYUN1V\'RSITY SAN LUIS 08151'0,OBISPO, CALIFORNIACAUFORNIA 93407