1977]1977) ChromosomeChromosome Studies•Studies-KeilKeil 7979

CHROMOSOME STUDIESSTUDIES INI~ NORTHNORTII ANDAND CENTRALCENTRAL AMERICANAMERICAN SPECIESSPECIES OFOF PECTISPEenS L.L (COMPOSITAE:(COMPOSITAE: TAGETEAE)TAGETEAE)

DAVIDDAVID J.J. KEILKEll

Chromosomal modificationmodification isis oftenoflen aa majormajor factorfactor inin thelhl: evo­e\'o­ lutionaryIUlioltllry diversificationdl\'ersiflCalion of aa large,large. rapidlyrapidly evolvinge\ohmg genus.genus. ,PUIU. thethe largestlargesl and most widespread genusgenus inin thelhl: NewNe~ WorldWorld tribetri~ TageteaeTagelC'ae (Compositae),(Composit3e). hashas radiatedradiated intointo a variety\'ariet) ofof habitatshabilats during itsiu evolutionarye\'Olutionarv history.historv. UntilUntil relativelyrclati\'ely recently,recently. how-hollo­ ever,e\"Cr. littlelillie has beenbeoen knownk~own of the thl: rolerole ofof chromosomal change in thethl: evolutione\olution of the genus. PectisPrctis wasIloas cytologically cy'lologicall) unknownunknown untilunlillhe the studiessludles of RavenRa\"Cn and KyhosK)hOS (1961) andand of Turner,Turner. Elli­flli­ SO'l,son, and King (1961). Subsequently, additional counts haveha\'e beenbttn reported for several taxataxa (Table 1).I), To date, however,ho"e\'er, only a few(~, of thethe species have been studied cytologically, and thesethese only from a few populations.popUlations, Of thelhe fourrour largelargc genera of thelhe Tagcteae,'ageteae, PectisPro;s has been the most poorly representedrepresented inin cyto­cyto­ logical studies. This investigation is one of a seriesserics of studies of thethe evolution ~ndand of Pectis.Pecfis. The genus reachesreaches itsits greatest diversity mIn the warmer areas of North America,America. and itit isis fromfrom thislhis region that most of thethc collectionscollcctions upon which thisthis study is based were Illade.made. Through an examination ofor the chromosomes of as many species~P«ies as possible,possible. and from numerous populations of wide-"ide­ ranging ~pccies.species, information regarding thelhe variability\'ariabililY of chromo-chromo­ SOmessomes inh PectisPur;s can be betterbeller correlated with\loith thelhe: evolutionarye\o]utionary h· historyISlory and syslematlcssystematics of thelhe genus.

MATERIALS\lATEKIA15 "''l:DAND METHODS\IEfHODS Immature capilUbcapitula were killed and fixed in thelhe field in modi-modi· r~led U1moy'sCarnoy's fixalhefixative (4 chloroform:chloroform : 3 elhanolethanol : 1 I glacial acetica~lic aci aetdl.d), and were kept under refrigerationrefrigeralion prior to examinalion.examination. In a modification of lhethe methods of BeeksReeks (1955).(1955), wholeIlohole florclfloret buds --ere^!re slainedGained (usually(usually withoulwithout heal)heat) for 2-48 hourshOUfS in standard Itt0 acClocarmineacet°carmine or iron acelocarmincacetocarmine solution and were\I"Cre squ3!ihcdsquashed ~~OUt[Vltnout Prio~Prior ma~rationmaceration in a drop of Hoyer'sHO)'~r's sol~tlOn.solution. Camera ucidaida draWingsdrawings \\ocrewere prepared from meIoticmeiotic mlcrosporcx)"tesmicrosporocytes to0 documentdocument eacheach reported counl.count. Table I. ~ Table I. Previously'·...iou'l) reportedropor••d chromosomedlOomoo.o.... countsoounh forroo Pectis.1'",... o0 Reported1<,p<>rtN OOunlcount I,,)(n) TaxonTa.Of1 ReferenceRef,rence P. angustifoliadnf:l'tltfolHl Torr.rorr. var.''ilr. dn1:angustifolia....'ijol'" 12 (1"(as P.r .fcf. ,'texana..." .... Co'}lCory) PowellI'o""n &II. TurnerTum•• (1963)(1%3) "12 la.(as P.r. angustifoliaIHlKO'm/oI", Torr.)run-.) PowellF'o".,,11 &II. TurnerTu...... (1963)Il96)) " P. ""1:"-"if''''''angustifolia Torr.Torr.• var.ar. ,~,...II"tenella {DC.)(DC.) KeilI:.il 12 laJ(as P. Im~/'"tenella LX")DC.) Powellpo..,,11 &II. TurnerTum... (1963);(l96ll: StrotherSI,odM" (1972);(1971~ " UrbatchUrhlidl in'10 LoveI ii•• (1974)I 197.) 12 PP..• coulter00"I~n i /faHarv.... &&I GrayGra) Keil(1975B)K,;11I975K) "12 1'.P. ~",n.udepressa IFern. efn. PowellI'o..en &&: TurnerTtl'..... (1963)tl96ll " PP. filipesfibf't'. Ifal'\'Harv. &.t GrayG",) ''iI.var. so... (1974)(l97.) "12 l'P. 1""pur"purpurea KtandeBrandeg... ,".var. sonorae___ ".ilKeil 1(edKeil 1197.)(1974) "12 fl.(as 1'.P. ".1-.-1palmeri 'iS. Wats.)WIIS) TTurner*u,...., .t Flyr(1966)f I)" 119661 " ,P. ""I>"rusbyi GGreene...... OKex A. GB)Gray 12 las(as PP. ,..b¥npalmeri SS, ""liSWats.) I TurnerTUlnel &:& Flyr(1966)n), 1I1l66) "12 (as 1',P. ".,,,,,,,,,/11puberula Glttnm.lGreenm.) Turner*h,,,,,, II Flyr(1966)1"1>, 1196b) "12 ,P. ..saturejaoides,,,",.,....I\ldlI (Mill.) 'ithSch. R,pBip. Turner,Tu...... Ellison[Iii""" &A: King1('101 (1961):(1961); PowellI'o..,n &II. TurnerTu.....r 11963)(1963) "12 rP. '.-10sinaloensis ...... 1. I"ernFern. Turner*Turne,&; Flyr11)'11061» (1966) < 36 "" I'I', ""'\If''''"''''subsquarrosa lll",,~(Hook, {f.) I 'khSch. Bip.lI,p. KyhosK>t- '"~in Wiggins""lU"" &III PorterI'on", 119111(1971) • 1977]1977] Chromosome Studies-KeilStudies•Keil 81

RESULTS AND DISCUSSION Chromosome counts were obtained from 28 taxa and from fivefive natural interspecificinterspecific hybridshyhrids Cfable(Table 2; Figures 1-34).1~34), Of thethe taxataxa studied.studied, 22 arcare diploids with n = 12,12. three arcare tetraploids with 11:=n = 24,24. ontone has both a diploid and a tetraploid race, and twotv.o arearc hexaploids with n = 36. Two of the hybrids are triploids with 8-J28-12 bivaJentsbivalents and correspondingly 12-20 univalentsunivaJcnts at meta­ phase 1I and with very irregular meioticmciotic behavior. The remaining h}'bridshybrids areare functional diploids with only sporadic incidence of ~eioticmeiotic irregularities. Supernumerary chromosomes were noted Inin three speciesspecies and in some of the interspecific hybrids.hyhrids, Of the counts reported, 18 of Ihethe taxa and all of the interspecific hybrids are first repons,reports. SeveralSeveral evolutionary lines arcare represented in the newly reportedreponed COUIllS.counts. Some of these Jineslines correspond to named sections of the genus. However, since sectional limits have never been applied t~to the entire genus, and in some cases, previouslyprc\'iously publishedpuhlished sec­ tional limitslimits have been found to10 hebe in error (Keil.(Keil, 1977),1977). not all taxataxa arcare presentpresently I)' assignable to sections. Consequently, in the followingfollowing discussion, taxa will be referred to section only whenv.hen thethe affinities are wellv.ell established. Section Pt(,/OIhrixPectothrix AA. Gray is easily Ihethe best-knownbest-knov.n portion of the genus cytologically. Counts are knownkoo\\" for all except twot\\O s~cies.species, PeNi.iPectis vol/muivollmeri Wiggins and the newly described P. peru­pl'rll­ \'lanaviana Keil (Keil.(Keil, 1977),1977). although varieties remain uncounted in 1"'0two otherother species. First reports for this section are two western Mexican taxa, P. jifipesfilipes vaT.var. jilipesfilipes and P. stenophyllasll'1Iophyl/a var.vaT. biari-Mari· S/ala,stata, a Chihuahuan Desert endemic, P.f'. illdsijofia.incisifolia, and a ~f1d~mic,endemic, P. all~u.f/ijofiaangustifolia var. faSlif{ialII.fastigiata. Numerous additional COuntscounts are reported for P. allgustijoliaangustifolia vars. anf{lIstijoliaangustifolia and It'wl/a.tenella, P. ji/ipe.rßlipes varvaT. slIbnllda,subnuda, P.P, papposa(lapposa vars. papposa and g'andis.grandis, P. prinK/d.pringlei, P. rusb)·i.rusbyi, and P. sinaloensis..iilla/o('mis. Section l'e('/otltrixPectothrix is interesting from the standpoint of chro­chro- IJlOs~malmosomal uniformity and the relative lack of sterility which"hich char­ ~Ct~TIleacterize the known natural hybrids within the group. Of the \anousvarious hybrid combinations which haveha\'e been studied cytolo­cylo[o­ logically,°gically, Onlyonly one.one, PectisP('Nis anXlIs/ijoliaangustifolia var.\1Ir. angustifoliaall~lIs/irolia X P.P./>O/I­ pap­ PosaP°sa var. Kfandis.grandis, was found to possess individuals displayingdIsplaying ~ioticmeiotic irrelllularitics,irregularities. Inversion bridge-fragment formations and 82 RhodoraRhodora [Vol.[Vol 7979

TableTabl. 2.2. Newiii.... · chromosomed1'0m0\Om~ countscoun" forfor taxa"" ofof Pectis.I'm;'.

ChromosomeChrom(KoO"'" numbernumbrr (n)rn) TaxonTuon with",nil locationIonlt'OOlnd and voucher'OIJ<'hcr'11

12 PectisPIOdi. angustifolia.n'....ilolb Torr.Torr. var.'At. angustifolian,,,,,ik>ti. Mexico,\lhieo. CHIHUAHUAOlJH'A."'''' 13.613" mi"" SS ofof Cd.Cd Jiménez,Ji",",",. K&MK ", .1.1 8387A;J81A; 25.6!S.6 mimi N, of Janos,JllIO>. K,,&. & MIf 8432A,lUJ}A. B.B. :Pu ... -.oo 1.8I mi E.. of Nazareno,Nu... fftlG. KA. 8c" M\I 8014A.M)UA Unitedt"'tt'Cf Co.: Big8ig BendHmd Nati"'all Park,Par~. K&MA .. If 7795A;7NJA; 15.11'.1 mi"', N" ofof Cas-ca,.. tolon,lolon. K&MK & If 7798A;liWJA; Culberson('.. IMuon Co.:Co. 1.4 mi SESE of HudspethItudoipcln Co. lineIi"" on 1-10,I 10. K&MK &. 1/ 7687; HudspethlIud'~lh Co.: SESf of FtH Han­It~n· cock,rod. K & MIf 7665A,7665A. B;11: PresidioPr~id,o Co.: 18.5Ig.S mimi S ofof Marfa,Morf•. K &&. M.II 7141,-7741; 13.113.' min,i SESF of Presidio,"r••ld.o, K&MK '" .11 7774A,7714A, B­8.

12°12* PectisPmi$ angustifolia.n~u,.i'o~1 Torr.Tnrr. var.\I. fastigiataf.,.I~;II' (A. Gray)Gflyl KeilK.il UnitedI·nil.d States.SI.In. TEXAS:rn~s SuttonSulTon Co.: ,Sono.a. KeilK.d &d RobertsRI,/vrlJ 1001'10074 pop. (FigureIFigure 2).

12 PectisPm;" angustifoliaIn~"'liro~. Torr.To,. var.\If tenella.....11. IDe.)(DC.) KeilKril México.\I.,ioo. rn~Htll-~: 107 m,mi S of PiedrasPlOdru Negras,....gr... K&MK &. 1/ 18'9A.7849A, B,8. CC;.. 114 min,i S of PiedrasPlOd... :-'.graNegras,.. K&MK d If 7860A;7MO.· 1Inbetween.....n Nueva,_-. RositaROIJLlI andlnet Monclova,Mon<:kn-.. KA <&I: M\t]86J 7865A; -t. 1L4It 4 mi S orof Mondcr.Monclova,... A"K & M\11 7875; ". 27 m,mi S of Monclova,Mondo'... K &&. MIf ]7879 1'9A,A, B/FiruroB (Figure 3);J): 60 miIll' S of Monclova,\fondou. K &&. M1I1'9t1 7911 A;A ll-S22.5 mi E orof P.,g.Paila, KA 6.Si M,\1 79441'9'4-", A; 16.7 mim, E of Paila,P.ila, K&MA 01 1/ 1'9jlJA.7950A, 8.B; 5.3U ....mi F.E of Paila.P••Ia. KA 01& M\t "'95'A7954A'; .. 13IJ mi W" of Paila­1',,110. AK d& M\I 7960A,"I'lWJA. B;B. 86m,8.6 mi E of Emli~noEmiliano Zapata,7..apall. K&MA &. If 7996;'9\11I; 44.2 2miSof mi S of AcaIIQ.Acatita, A"K&M II Mil!A.8052A; 2.1 mifl1I'ofAcalill." N of Acatita, K &4" M 8OISJA.8065 A; 7.8 mi ,N of Anllla,Acatita, K&MA 6. If Ibn8072 ...A; 1.1 mi S of RanchoR.ndtO El Cinco.Cinco, AK&M & It II0!A,8102A; 10J10.3 Ill'mi S orof RanchoR.nd>o los\ol; Charcos,ChlfC'Cl'. AK&M &. 1/,I6JA. 8165A, 8B. ,DURANGO..... ~ ..c.o E of ""''''11I1.Mapimí, K&M" '" II 8186A­IMA lnht1lUnited S'.'"".States, TE.XAS:TH"" IIldll",Hidalgo CoCo.: 010.8 miIll' £E 0'-of jctn.;e.n." w/Texas T..u' F\lFM rdlS60nrd 886 on US 83,J. AK & C9141 C 9247..A,, 8.B; S1."Co,:Starr Co.: 1ImiWd­ mi W of joe.n."jctn. w/TexasTu,..t"" FM rd 1.'01430 on US 83. AK & ("91'1/6,1L6C 9248B; 11.6 n".... SF.SE orof jeln.jctn. '4'w/TexasTu.. f\lFM rdrd 2098 on usUS 83.83, KA &.& C 9149,49249 A­ 1977]1971] ChromosomeChromosome Studies•Studies Keil Keil 8383

TableTab'- 22-t<>~tinuW • continued

Chromosome number (n) Taxon with location and voucher

12* PcctisI"IOCI;' berlandieribtrl:ondini DC.IX' Mexico.:'>Ih>to. TAMAULIPAS.T~\t~nll'\~ justj ....t EE ofol Magiscatzan,\1ap.ealUn. KK'" & CC 92359!JJ A,A. B,B. C;C: 3J mi EE of Mante,MIDlt. KA &'" C 9238B;91J81J: 8.9 mi"" WW ofof Magi-Mac>­ scatzan,lQlUn. KK'" & C 9240A9140A (FigureIf"ipt 4),4). B.II.

24* Pectis"l'diJ bonplandianabonpl.ndian. H.B.K.II RIC. Guatemala,G ..., ...... Ia. SUCHITEPEQUEZ:~~("Kll[I'FOUI PuentePuente Siguacan,Si,...can. KK 9439A~J9A (Figure(Fillu.e 5);S); 5.4SA kmIfm E£ of jctn.jeln. w/rd'" ,d to10 ChicanoaCIuclanoa onOll. CA-2,CA 2, KA 9442A;'#4}A; 11.5II.S km E ofol PuentePuente SiguacanS>,~n on00 CA-2,CA 2. K9448AA"U#.A·,d.

12 ".* PectisI"IOCI;" capillipescapilli~ (Benth.)(!knill.) Hemsl.llem,l. Honduras,llondu..... CHOLUTECA:UIOLUT[CA justj""t S'i ofor San AntonioAnto~i<.> ded. Flores,F1ocfl. K 9505A9JQJA (Figure(Fi,u,. 6),6). B,B. pop.;pop,; 8.4 kmkm WYo' ofo( CholutecaCboluttc:o KK 95179JI1 pop.""

".2^* PectisPte"lo qlind,itacylindrica (Fern.)(Fttn.) Rydb.R)·db. Mexico.:'olhito. COAHUILA:roMRtt ~ 9.4 mi S of Rancho los"" Charcos, C~UC05. KK & MII 8161816/ pop. (Fi,ure(Figure 7).1). SONORA:'.,N.'.A 28.5285 mim; WYo' of A'lamos,Alamos. KK & C 8663861H pop.

". Pectis"I'dil fiNp<'Sfilipes Harv.Ibn". &.& GrayGn.y var.\lr. filipesfilipn México,'lhito. SO'OfI\,SONORA. 282lI mi S ofllmnosillo.of Hermosillo, KA &&: C 8617MiI7A.A, B,B. C; J,lI3.8 m,mi E ofol jctn.jetn. w/Mexico"" \I.ut<> 15IJ on rdId to'0 Alamos,^Alamoo.. K'" & CC &lJ9A.8659A, B;B. 11.111.8 mi E of jctn.jan. '"w/Mexico" ••it<:! 15IS on rd,d to1O Alamos, Ab..-.. KA

12 PtnftPectis I'iliPftfilipes II.",Harv. &.& Gn.)Gray 'atvar. ,\1bft1Olbsubnuda "an.Fern. " "México,..ito. (""'"lCHIHUAHUA:\Ill \ 242a IlUmi N, of Nueva,""".. CCasas...,u Gta.ndn.Grandes, AK '"& M\f MOllA.8406A, B. l,.;United.... NaIl".States. ARIZONA:\."0'\ Coc"",,,Cochise Co.:Co. O.S0.5 mrmi S of PI..ad"".Paradise, AK '"Sí .IfM BJIJA;8512A; Ou.oll""Chiricahua ",.,1Nati Monu-Mon... _nt.ment, K '"& MIf BJ18A8528A lfi,lurc(Figure 9). TEXAS:IfH' P'~IOPresidio Co.,Co.: 18.5IllS mimi SS of Marfa,\!a.fa. AK

Table 22 • continuedeol'lunUotd

Chromosome('h'OlnO'Omc numbernumbu (n)1,,1 Taxonrunn with",th locationIocauon anda.... voucher\oltChe.

12*12" PectisPtnio haenkeanahamL.."" (DC.)(DC,) Sch.ow,h Bip.DIp México,"hiro. GUERRERO:(.l HUM) 3.22 mirn' S ofof Iguala.I,w.l:a. K&CA JI {' 9144A,9/-1"", B;.. 1.414 mimi:'> N of SabinaSo.b"u Grande.OnllCk. K&CA.I C'IIISA 9115A (FigureIM"'ll' 10),10\.11::2 B; 22 mi.... S of Iguala,lpqb. K&CA .( C 9117A;9111A. 22:: mim, N" ofof ZumpangoZunl~n", del dd Rio.R'" K&CA & C 9119o/IOA. A, B;B. 4.54 j mim, NN of011"...1:0 Iguala on MéxicoMhlC'O 95-D,95 D. K&CA .I C 9137A,91J1A. B. MORELOS:_riO' 25.7!S.l mim, N..; of Morelos-GuerreroMortlol-G""rm.. boundarybound1f) onOn MéxicoMhic'll 95-D,9S D. K&CA ... C 9140A.9/lfJA

12*12' PectisPM;" incisifoliaiotci..tolia I.I M.111 JohnstonJohMlon México.:\Ihiro. COAHI'H.A:co,II. "" S<; of"f Laguna1.a,ul\lI delIkl Rey,Rey. KA.( & M118101 8101AA (Figure(h,o.( 11).Ill. B. D. E;£.810114. 8102A; 8105A,a/OJA. B;R. 8109A,ll/()QA. B.Il. C.

12*12' PectisPff';" leavenworthiiIUH""'o"hii Standl.S,andl. México.'1"'''0. MICHOACÁN:\l1("11O"C"~' 5j mi Er of Apatzingan,Apall1npn. StuessyS"""O' &J RobertsHoj"rtl 3744J7U pop.pop. (Figure(fi8u,e 12).

12*12° Pectis1'«1i< longipesIon~ilJ'f' A. GrayGra>' Unitedlnited SlateStates.•. ARIZONA:UI/"'" SantaSan,a CruzC'UI Co.: 2.5 mi"]I N... of So-So­ noita,nona. KK.I & MII 8566A8jf)tJA (Figure(Fi,ure 13).13)

36*36" PectisPf<'" multiflosculosamultin.,.,....l0", (DC.)(1)(')!Kh Sch. Bip.H,p. México.'Ihito. :"'MO. Mazatlán,Ma/a,!jn. KeilAtil & RobertsR,,fo,m 10I60B;IOI6IJB. Mazatlán,\1a/aILin. StuessyS"...... ,·.4 & GardnerG""ht" 3040J{)40 pop. (Figure(f'I!UK 14).141.

12*12· PectisPut;" ~ncrdiaoerstediana... Rydb.R>db COGuatemala,...,nn."'. JIJUTIAPA: n~~~ 7.77.1 km\m E of jctn.Jetn." w/rd rd,o to Quesada~ on CACA-1, 1. KA. IUIJBIF"9413B (Figure...." 15).lSI

12 POd;"Pectis P"~papposa IhI'"Harv.

TableTable 22-eo01;nued • continued

Chromosome nu.nbe,number ('I)(n) Taxon ...withith 'o<'ationlocation and voucher\'Oueher

12 1',<1Pectis.. papJlO'apapposa Harv. &:& Gray var."a" lr.ndi.grandis Keil " ~Ihie".Mexico, CIlIIIIIM'IACHIHUAHUA: 15,415.4 mi 1'1N of Cd Canmrgo.Camargo. KA" & M 1I!51-f/4;8252-1 A; 20.22(1,2 mi N" of Janos.Jano$. K & MIf 84.'9:8429; 25.62H. mi N" of JanoJanos,•. K ..& M 84)),4.8433A. Unitedllnilrd SIalStates,..... ~f"NEW "n'n'MEXICO: Doñao.:.~a AnaAna Co,:Co.: 2~28 mi N" of la,Las Cruces,Cru~~. A'K 7654/4.7654A, ('C (figu",(Figure 17);17): I1 mimi I'N of jeln.jctn. I1-10 10 & I 25. KA' 16J.5/4.7655A. B. TEXAS:'OXA' BrewsterBre",I,r Co.: BiglIig BendBond NatiNall Park,Park. K & MIf 78/8A';7818Á: ElEI PasoPa.o Co.: llor;7onHorizon Cil)'City eXilexit on 1t 10.10, K ..& M 165911.7659A. B;8; 2.62.b mi NW:-. W ofor Tornillo uilexit on I-10,I 10. KK" & M\f 16,'j,fA;7664A; PresidioPrC)idio Co.: 2.7 mi01' Sf.SE orof Redford.Redford, K ..& ,IIM 7118A.7778A.

'2 P«li,Pectis prinll.ipringlei Fern. " Mexico,Mh"o. nnWCHIHUAHUA:A"I'A 0.6 miOIl I'N of jeln.jctn. w/rd" rd 10to Escalónf...:al~n onon MhicoMéxico 49.49, KK" & MIf 810fJ-1A.8206-2A. B,8. C:C; 29.9 mi S of Jiménez,Jim~l\Cj. AK .t& M If 8l18A8218A (F;gur,(Figure 18). COAlIl",ACOAHUILA. 36.9Jt>.9 mi W ofor Paila.Pait... K & .ItM 7916A.7976A, S.B, C.C, D;I); jctn.jeln.... w/rdrd to10 San PedroPCilro on MéxicoMhieo 40.40, K ..& M 79//IIA.7988A, B;II: 10.3 01;mi S of Rancho los10. Charcos,Charco•• K ..& M Sf".,A.8I64A. "lRA'

". 1'«*Pectis pprostrata.....".la Ca>'.Cav. [/El Salvador,S.'~.dor. lA'LA LIBERTAD: '''!.RIM' I km E of SantaSan~ Tecla,'tecla. K9400~:9JOO pop,pop. GUll.mala.Guatemala, ELII p"ooussoPROGRESSO. 43 km W of ElFI Progresso,ProgrC)'o. I(K 9417A;NIlA; 23 km W of El£1 Progresso,P,OgreiSO. K NI8A9418A (Figure(Fi~ur. 19). 0'GUATEMALA:... HMAl.... 30 km S[SE of Cd,Cd. GUII'mala.Guatemala, K1(9398A. 9398A. JUTIAPA. Jl.""'P'" 7.3 km W of jctn.jeln.... w/dirtd,n Idrd 10to JalpalaguaJalpatagua on CA-J,CA I. K 94099~09A.A. S"'''TASANTA ftosROSA:... · 3.5 km NW of jctn.jeln, w/rd" rd to10 ColiapaCuliapa on CA-1, CA-1. K 94ooA:9400A; II11 km E of PuentePoen(e ColiapaCuliapa Onon CACA-1, l, K 94059~05A:A; 2.1 km E of ,KIn,jctn. w/rd" rd '0to Jalpa'aguaJalpatagua on CA-1.CA-1, K 9406A.9~OfJA 7AC"'PA,ZACAPA: 3.1 km E of 1'1ICnl~Puente ".lmillaPalmilla on CA-J.CA-1, KA 942494:4A.A. Mhieo.México, nmWMtllACHIHUAHUA: 4.5 nlimi N" of Galeana,Gakana. K & MIf 8388. 83//8. COAHUILA.COAHUILA: 47 mi S of MOllC'lo\·,.Monclova, K & M~f 7901A.7901A, C. MEXICO. MH'CO 20 m;mi SSE of Zitacuaro,Zitaeuaro. S/lU'H.~Stuessy <& GafJ"",Gardner 3126A.3/MA. NAYARIT:~AYA~'l 44 mimi SE of Acaponeta,Aeaponela. K & C S1I59A.8959A, pop. SINALOA:''''''LOA 8.7mi8.7 mi SE of Ro"",no.Rosario, K & C 8948A8948A.. VERACRUZ:• , RACRll I mi01' Sf.SE of Tan-Tan• 10)o<:a.toyuca, KA·" & C 11117S,9227B. Unitedl'ni'e GrantGram Co.,Co.: 6'h6'/2 mi EF of SanllSanta Rnl.Rita, K & M 8~1!8472 pop.

2.% 1'«1Pectis.. f'jlt'Mrepens Brandeg.IIrandeg. ". .\I"'i. 6.16.' ntimi E of Nayarit-Jalisco"a}'ari.-Jali...:o boundaryboondary onon M'xiroMexico 15.15, K & ('C 899)8993 pop. (Figor~(Figure 20). 12* Mhieo.México. "rx,coMÉXICO: 0.5 n,;mi N" of Goe"ero·MhieoGuerrero-México boundary Onon FederalFed~ral Rl'Rte 55.55, K"K & CIl(}97C 9097 pop.fN'Ir (Figure(F;gu~ 21).11). 86 RhodoraRhodora [Vol.{Vol 7979

Table 2 • continued

Chromosome number («) Taxon with location and voucher

12 PectisP~I;" rusbyi.....b)i GreeneCrft_ exu A.1\ GrayGr.) " Mexico.\1hico. siNALOA:~1'~10~ 20.6206 mim, SS ofof Sonora-SinaloaSonon-Si....10.1 boundary...... 1IliI~ onon Mexico\lhico 15,IS. KA &cl CC"'J,fA, 8738 A, B;B. 1.8II milII' EEofs"n of San Bias,1l1a•• K&CK" C 8744A,6UU, B,B. C.C SONORA:\&\0... justju" SS of Magdalena,Mapl.lena, K&CA & C 8594A,Ij94A. B,II, C\C". D;D. 71 minil S of01 Magdalena,\hpkl\.&. K&CA

12 Pectispmi'. saturejaoides...'''uj.old.... (Mill.)(M,It) Sch.5ch Dip.flIp. ~'o ~m " Guatemala,r."atcmll.. EL[J PROGRESSO.rROGR' 5.65.6 km E of turnoffturnoff to10 ElEI RanchoRanch" Onon CA-9,ell 9. KI( 9423A941),01 (Figure(Ilwur< 23),23). B,B. C;C: 0.5 km W of turnoffcurnofflO to ElEI Rancho on CA-9,ell 9, KA 9427A,941714, B.Ii

12 PectisPC' ";n.]O<'",I,sinaloensis Fern.Fe.n " México..\lhie<>. siNALOA:~I"'OA Altata,Ahall. KA &" C('67866; 8786G; Mazatlán,MI,.llin. K&CI." ( 88/0A.8810A, B,Il. C,C Df) (Figure(Fi,un 24),24). E.F:.

12·12* Pectis,"«I;" "ftlophjll.stenophylla A. GGray..) var. , .., biaristatahl..i'lIl. (Rydb.)CR)db.) Keil Kril Mexico.\Ihito. siNALOA:~''',IO, 16.816.' mim' W.... of ElEI Fuerte,f-lItn~. KA. &JI CC 8752A,·5....1· Il.B, C (Figure(FillU~ 25).~)

36 1-'«,0.Pectis .ukqoq.rro..subsquarrosa (llook(Hook, nf.) Sch. Bip.HlP Gua,.to\Galapagos 1".IICk·Islands: Hood1l00d IWand.Island, C.C H.If R¥INRacine s.n.UI· (Fig-Ifill'" "tture 26).261.

24·24* PectisP«I" ,swartziana...,riafta 1"'1\Less. 11.-.Honduras. ,VALLE: ..'Il I'1.9 In'km S 01of jctn.jelO. "'.w/CA-5 CA ~ 0'1on CA-ell I.• K 9510A (F(Figure ,~ 17~27), poppop.

12·12* Pn1Pectis... uniaristata"",'''.,. DC. '~rvar. IKlIo\lrm.holostema AA. Gr.)Gray e UEl Slhador.Salvador, ,LA.. 11111LIBERTAD:....0 10.1 k",km 'IN of Qur:1r.~<'P"'l'"Quetzaltepequ u;,exit on EIEl $at,Salvador..dor I.I, AK 'U51A.9452A, B. 1I0ndi>,Honduras..... ("HOI'CHOLUTEC* rr("~ 24 km W 01of Chol"'~ClI.Choluteca, AK 9~/lA9512A IFi,ur.(Figure 2S);28); :020 km 1\W o CholultCll.Choluteca, AK 95/5..19515A. ,977] ChromosomeChromosome Studies•Studies Keil"cil 8787

Table 22 • continuedCOfIllnued

Chromosome number (n) TaxonTuon with."th locationloallon andand voucher HI''''',,",

12*12" PectisPuriollniariuata uniaristata DC.DC. var.nr jangadensisjan~ad~tS, (S. Moore)\~) KeilKnl México,\lhiN. GUERRERO:GLfllllUO 1212mi mi SS ofof Chilpancingo,Chdpa..... n.... KK& & CC9/!1 9127 pop.pop. JALISCO:Htr-a;> 2.12.1 minll NW'''' of MagdalenaMap""" airport,airport KA & ".'2* Pectisf"ftth angustifoliaal\J:USlifo(ia Torr.To.. var._ar angustifoliaanfU\fifolia Xx P.f". papposapap""", Harv.lIaO', &&I: GrayGra)" var.,-ar grandisuand" KeilKfd UnitedI niled States,~Ia.~ TEXAS:If'''' Brewster!ltn>',tft Co.:Co.; Big11<1 Bendflfnd Nati'atl Park,Pa,~. KII &.. M" 7797;1191: Presidio" ....0<1'0 Co.:Co., 1010 mim, S"of of CíbolaObol.a Creek,Cr..L. KII & MIf 7756ny, hII'A (Figure(hflure 30),30). VlO (Figure(h,urf 31).JI~

1211·J2I'.12 II + 12I*; p1'«1;"ect¡s bonplandiana_plandiana H.II B.9. K.K X)( P. prostrataprmlr". Cav.eav 8 + 8Jj'21)I' " 2° I* Guatemala.(;lIat"",ala. SUCHITEPEQUEZ:~"CIllH"-Ql U PuenteI'u.n'. SiguacanSi.""un K&II & C9438A,(,'NJ/IA. BB(FiEu (Figure•• 32).J2).

".•2* PectisI'fClh filipesfltiJlf> Harv.Ih,,'. &&: Gray var._If, filipesIlll~ X P.1'. papposap'Pf'D'-ll Harv.Ib,,'. &&: Gray var."ar. papposapapp"sa México,.\Ihioo. SONORA.W'O." 72 mi S of BenjaminflfnJam,n HillIJili on Mexico,\lox;';" b 15.,~. KK& & C8611DC8611r1Wigurt'JJl (Figure 33). 12 ".* PectisPf.l;" papposapappo>a Harv.lIa". &" GrayGral var.'If, papposapa"posa X P. sinaloensissin.IofoI>" Fern.r.m (Pff,i,(Pectis X salinakll"", Brandeg.)Brandfl.) Mexico.\Ih;'o. siNALOA:SI'AlOA Altata,Allata. KJo. & ('C 11668.8786B, C,C. Dn (Figure )4).34), FF.

Cll.ca. 9 II +• PurPectis.. surlrn.naswartziana Less.I.ns. X P.P..... uniaristatarMala DC.DC. var.•at. holostemah.....oma II18 I'!* AA. Gu.yGray Honduras,lion...... OK.lll'l£C"CHOLUTECA: 20 kmk'" '"W of Choluteca,C't>oIut«-a.. KA 9514119514B.

.~ Previously...... sI) n'fpOflaiunreported tawntaxon or hybridh}b"d CombonillOflcombination. \I AIlem.Abbreviations...hollS 01of «lImon.collectors' ""names:..... AK • UaudDavid Keil;"nl; KA .II& M=101· Keil"f,J.t & Lylel)k A.A C ';;GI1I:.Gill; AK "& Cc _= KfllKeil &.& JlldnhJudith 101M. CanCanne.... Voucher\'OUfhf, 'r«'IlW'"specimens a",are depositeddqIowled atal ..e^Ornoot.t. StaffState I'n"fn'l)University Ikmnu"'los~Herbarium (OS). ,.;II.• 12 II+-2 w:qdsmall ...supernumeraries.prmUmn3f1n ~Anaphase... I lM'1d,.bridge obwr\a1observed in """'"'some ""lb.cells. h,^ome ocn~cells fOn''',ncontain Ontwo anaanaphase...... lof I1 bridgesbndl6 and a>O'_ ~ ' .. V .. --,::.. ..,p' ,. ~~ ~ ....'\ .14 .. l' "'~2 • '01 1 -3 ~ ..... • ,-" A ?*• .....".'.: 8 "i~":1 ...... I" . • I'\1\"" .., • ~•• .­ 1\. 7 ...... -...... 5 .,­'-,·,'-6 ", Visv­ '-6 "7 -*r•

1 I,J'" .~/~ 1, I • I '-1 r "*£~ "1010 **#•.~:t "12 :'''12 .,. /~ ¿V 11 .~ f~~ ~9 I,: ^9 ¿fit~r" . 0'0) ',­' .. ;$''':15,-/ "'13 1'","-- ,_ .(. ~ I ... ~ J\ -;,.. .. ~ ''l<,-:}14 ~ A/tIi /'' lOa10u - .=:16^216 Q4~

FilUm.Figures IU16. 16 Ca-...Camera lucidalwid. ddrawings...... t!' ofoIllW'iotic meiotic ."'_chromosomes of01 Pectisrod_ U'"a ..and h)briob..hybrids. Voucher\our:hrr ~cirncl5specimens areaR lOll.angustifolia, diakinesis,dIlL,-. In2If =.. 12 II.II 2,2. P. angusttfo4~1l" Utvar. f"l'~""ll.fastigiata, d",LIllCSOs,diakinesis, 2»2Jl ..= 12 II.II J.3, .P.I' II~Jlifolangustifolia.. var.•:a'. "..ntenella,•. metaphaseI••:t.~ I. 2It2« ..= 12 II.II ...4, ,.P. ~'W"'/wr~berlandieri, metaphaselnfIS.sis, 211-2/7 = 24 II. 6,1'.6, P. ,tqJfl5pn.capillipes, ...apllaKmetaphase I,I. 2»2If •.. 12 II.II 7,7." P. nilltd,,,,cylindrica... meta­.-.. ph.aKphase I.I, 2rr2/7 ..= 24 IIII. 8."8, P. flipsßlipes 'Ifwar. ßlipes,fl1'P'J, anaphase.~phu< I.""I, n - 12. 9,1'.9, P.filipe*** filJrti'~ SIIM_subnuda, mnaphametaphase I.I, 2nIn ~- 12 IIII. 10."10, P. ~,.,_.haenkeana, metaphase_taplla... I,I. In2n -.. 1^11 '2 1 II.11, P. INtii/O""incisifolia, anaphasephase I.I, "n ..= 12. '2.12, I'P. IN,leavenworthii,..." .._,"". telophaseulophaK I,I."" n ­ ll.13, PP. ,,~,(1'#'J.longipes, d",L.nel.1l.diakinesis, 2n"2/7 = 1212 IIII. 14, PP. ""'/,,/1oJfulomultiflosculosa,.... ddiakinesis,...~inw.. 2«l .. =-)6 36 t1 I~.15, 1'.P. MWrd_.oerstediana, "",.aph3Sl:metaphase I.I, 1n"2/7 = 12 IIII. 16.1'.16, P. papposaJ>QPPOSIl var.'"ill.~' papposa, me,...eJ'" pha~phase I.I, 2Ir2« _= 1212 II. 1977]1977] Chromosome Studies•Studies~KeilKeil 8989

~ C' ... / *-* t ~.> ~ y .... y 20 o~ * *18 19 20 17

· ' x ,) .. I 7 * • • "25 • «• ^23~23 24 21 22

.... II • ..t: 5 +i -",,"- ::. " .~,.. fr .. .. --~,.. ,::" ....' ,.. , d'Aï »'26 -27"27 w.. po " 30 28 29 .. ,~ *"•• ...*• ...­ ^ v^ . -­ « °-, !I »* *» .. • ..•..... • V... .' I ^33 ^*34",34 0 ^¿^ I • • • lOa 31 3232 '~ *>" h'lI'.,Hgures 17-3417-34. C._r.Camera lucid.lucida drawingsd, i"C'I ofoK meiotic....iolit dlt_chromosomes ofor Pectis"KI.. 1:Ioutaxa .ndand h}brid...hybrids. 17.P.17,P. f1GN>OfIIpapposa var.' , ~n.,.d".grandis, diakinesis.d.aIt'IWi.... 2«br" = 12 II.II 18. P. 1"'"",~~Pringlei, dll~In<'\fS.diakinesis. br"In = 12 II.II 19.19, P.P prostrata,proW.'., metaphasemt1.ptaa,.. I,l. In2rt =.. 12 II. 20.:!O, P.P "Prmrepens tntI.Iph.,~metaphase I (poI;Ir(polar 'Yview),.. l, 2n"In = 24 II.II 21,21. P.P "pm',repens, ddiakinesis...It",n... In2n" = 121211 II. 2 ;-- PP. ,..$/>rusbyi,...... ,.pNStmetaphase I, 2n"In - 12 IIII. 23, PP. saturejaoides,"'U~,J". metaphaseme1.pM~ I,I.:!II In = 12 II.II 4 •- PP- '_klm",.sinahensis, d"'~I1"'''s.diakinesis, 2n"2n = 12 II. ~.25, PP. stenophyUa."",-",rt•• var.,-aT. """1Jlbiaristata, meta-mtU­ Pba'CPhase l.I, z"2n ..= 1211i2 ¡j 26, PP. suhsquarrosa.""'~qud"ottI mrup/LImetaphase... I,r. 2nIn =~ 36 II.II 27. P. ,swartziana, ,,:..,_ ~1tlM1".diakinesis, 2"2n ..= 24 IIII. .28,28, PP. u"",ml~'11uniaristata var.••• 1rtoH,,_.holostema. doakinnis.diakinesis, 2n:2n" = 12I~ II.II •. PP- --"'","uniaristata ...... var. jangadensis,/II1'¢'I. f'Uf'f"OJQpapposa Ut.var. f{rtI",hgrandis,...... anaphaserha

- - ..... , ...... adpne- e^ ' Braz,lian and Mexican collections of Pec/is uniaristata van jang­ Malme 31 "' '0 (GH) and Feddema 2190 (MICH).

T}"picalypical PectisPtrlis uniaristata,lI1/ioriJ/olo, alsoal.so a Mexican\1c~ican taxon,laxon. hashas notnot yet)CI *en~n stStudicdudied cytologically.c)tological1v. tio, Stablc table .polypIoids polyploids naveha~e arisen inin atal leastleaSI threelhree differentdifferent evolu­e\'olu­ lIOnaryry uneslines inin Pectis.Peetis. Section LorenteaLoren/eo A.A. Gray,Gray, perhapsperhaps thethe most Plodrn~t Pnrnitprimiti\e've °fof tnelhe genus,genus. contains bothboth diploids andand poly­pol}"­ worth-ploids_ ThreeTee speciesspec.:ies fromfrom thethe section,stttion. P.P. haenkeana,haml..l'tIna, P.P. leaven­leall'n­ "'of/h" a o pIf allid P-P. saturejaoides,sorurejDoideJ. areare knownkno.... " onlyonly asas diploids,dIploids. andand oncPl' . Pectip, '. • OllSquan1a.squama> nashas been~n reportedreporled onlyonly atal thethe tetraploidtetraplOId level,level. both::'" H'T* reptns,7" aalso'S° aa memmemberber ofof sect.secl, Lorentea,Lo,entt'O. apparentlyapparently hashas Populh diploid1P and and tetra tetraploidP'°'d races.races. FurtherFurther studystudy ofof additionaladdilional 0ns w frPU!ahO0r th ns will^l bebe necessarynecessary forfor anan understandingunderstanding ofof thelhe basisbasis t0Or r theS(' differentdifferent races.races. TheThe twoIwO levelslc\'cls maymay eventuallyeventually provePTO\'C a~o widr~preS('nl reSent cryCl)'plicallyPtical,y differingdiffering species.spec-its. PectisPtrlis multiflosculosa,mll"if1o~uIOJQ. °each~lde:spread a(* P!'<'rennl31erennial speciesspecies whichwhich occursoccurs onon PacificPacific Ocean<:>cc:an he:s fromrom SinaloaSinaloa to10 Peru,Peru. isis oneone ofoflhe the two tv.o knownlnollin hexaploidshexaploids Vo1 79 9292 RhodoraRhodora i[Vol.­ 79

inin thethe genus.genus. This unusualunusual species,species. uniqueunique inin itsits long-creepinglong-ereeplO& habit,habit. mayrna)' bebe aa membermember of sect.sect. Lorentea.l..or('lIIt'u. SectionSeclion Pectis~('/is isis representedrtpresented atat bothboth thethe: diploiddiploid andand thethe tetra­tetra­ ploid levels.k\"c:ls. The discoverydiscovcr)' that lhal Pectis 1'('('1'$ prostrataprostrolD isis aa diploiddiploid whereas.... hcr(as thethe very\1"1')' similar P.P. cylindrica{-.rUne/rl('o isis aa tetraploidtetraploid addsadds con-con• siderable weight....~i8ht to10 thethe argumentargument that[hal thethe two' .... 0 shouldshould bebe treatedtruled as distinct speciesspecies (Keil,(Kcil. 1975A). PectisPUlis bonplandianabar/plane/iaM andand P.P swartziana,SMQrt:/ona, twoI\loO closelyclosel)' relatedrelated andand very\1"1') similarSImilar speciesspecies (Keil,(Knl 1974),1914). are bothbOth tetraploids.tttraploid<. FurtherFunher studystud) mayrna)' indicateindicate thatlhal thesethot two(\000 taxata~a are actually conspecific.conspecirlC. TwoT",o triploids,lnploids. both thethe resultmiU11 of diploid-tetraploiddiploId-tetraploid hybrids,hybrid!. represent thethe firstfirst reportsreports of thisthis polyploidpol) ploid levelIc\(1 inin Pectis.PU-IU. InIn both hybrids,h)bnds. P. bonplandianabOllplQIIl!iana X P.P. prostratapmuroto and P.P. swartzianau"QrI:iantl Xx P. uniaristata,IlI/iuri.flOIa. partial pairingpairin!'l occurred, occurred. indicatingindicating atal leastleast somesome homology or homoeology of the parentpall:nt chromosome comple-comp~ ments. The evidence is insufficient,insufficient, however,ho\\ever, toto indicateindicate that(hat thethe diploidsdiploid~ involvedin\"ohed inin these hybrids are directly involvedin\olved inill thethe ancestry of the respectiverespective tetraploids.telraploids. As was indicatedindicated above,abo\'e, evolution in some partspans of PectisPuris hasha~ notnOl been accompanied byby chromosomal isolation,isolation, and thethe chromosomeschrom~omes of quite different species may be capable of full pairing.pairing, TheThc ability of P. uniaristatauniarislalO var.\ar. holostemaJIOJosltma to hybridize with a member of sect.'ecl. PectisPeC/is sug-sug' gests thatthaI there may be a fairlyfairly' close relationship betweenbety,een thelhe P.p elongatatJo,,~ola complucomplex and sect. Pectis.PeC/is. The only species knownkno.....n cytologically fromfrom the South Ameri-Amen­ can region is PectisPenis subsquarrosa,slIhsquorrnMJ, a hexaploid endemic of the(he Galapagos Islands. The hexaploid levelle\'el suggests thatIhat this speciesspee~ has had a complex e\OIUlional')'evolutionary history.hislol'). CurrenlCurrent researchusearch (Ktll(Ken- unpubl.) suggesusuggests thatthai a patternpallern of adaplh'eadaptive radiationradialion and sub­ sequentsequent migration may haveha\e played a partrart in theIhe e\'0Iull01'1evolution oof both P,P. subsquarroMJsubsquarrosa and P. Ittllli(oliotenuifolia (DC.) Sch. Bip-,Bip.. alsoakO en(lt-micendemic 10to IhI:the Galapagos Islands. Further samples of theseItldC species from addilionaladditional islands may pro\K1eprovide more informationmformauall regarding lhethe n'olUlionevolution of IheSoethese species. BothBolh Galapagos Islandslsland~ species are members of an unnamed SoulhSouth American sectionseclion 01o PUliS.Pectis. Diploid counlscounts are reported here for theIhe firstfint timelime for 1,,0two perennial speciesspecies of uncerlainuncertain affinilles.affinities. Pectis~rlis lonf(iP'slongipes appearSappeal 10to be m~tmost closely Il:latedrelated 10to secl.sect. Pl'rlUlhri:rPectothrix but also posstSst">possesses features in commoncommon with somesome members of secl.sect. Lorentea.Lo""/I'o. PPtrleC lJl 1977]1977] Chromosome SlUdics-KcilStudies•Keil 939J capillil"'Scapillipes (I'.(P. creCfaerecta Fern.)fern.) appears to be mostmOSI closely related to members of the P. elollgalaelongata complex.complex, perhaps through the peren­ nial P.p. diffllSadiffusa Hook. & Arn. Thirty-four taxa in PectisPeelis are now known cytologically,cytologicillIy. somesomc from milnymany populations.populations, some from only one. Thus far,far. only one base number, x = 12, hasha, been reponedreported for the genus. This situa­ tiontion isis inin strongstrong contrast to that found in the other large genera of the Tageteae, which are all multibasic:muhibasic: Dyssodia,D.1"Ssodia. xx:: = 7,8,(12?),7.8.(12·!). 13 (Johnston(Johnston & Turner.Turner, 1962:1962; Strother.Strother, 1969):1969); Porophyllum,POfUphdhllll. x = IL11, 12.12, 15 (Johnson.(Johnson, 1969):1969); i1ndand Tageles.Tagetes, x=x = 11.12 (Strother. 1969). Inin light of thethe vilriabilityvariability of the chromosome numbers of thesethcse related genera, it is surprising that such a widespread and diversediven.e gellusgenus as PectisPeelis has remained monobasic.monoba,ic. Certainly notnOt all of thethe taxa of PectisPeois have been examinl'dexamined c}lOlogically.cytologically, and thethe po,sibilitypossibility still exists that additional baseba\c numbersnumber, may yet be diSCOvered.discovered. The probabilityprohability of such an event is lessened,lessened. how­how· ever.ever, by the broad spectrum of thethc genus which has nowno"' been sampled. The largest gaps in the present knowledge of the cytology of PectisPeetis lie in the WeslWest Indian and South American rangerange of thethe genus. Of particular valuev'alue will be counts from additional mem­ befl;bers of sect.sect. Peetis.Pectis, particularly from thethc Caribbean region.

ACKNOWLEDGMENTSACKl'o:QWl.FD(JM£STS

Field sludiesstudies wcrewere supportcdsupported by National Science Foundation Grant GB-30240. I wish to thank Dr. TodToo F. Stuessy and Dr. DonaldRonald J. Pinkava for critically reviewing the manuscript. I am tePly.gratc.fuldeeply grateful to Miss Judith M.~. Canne and Mr. Lyle A. McGill1vlcGiJl foror thclrtheir assistance in the field.

LITERATURE CITED Rl:us.EE*s, R. M. 1955.1955. tmp.O\~menl'Improvements in thelhe .qua,hsquash techniquelechnique for plantptanl chromo­ 'omes.somes. AI;'o.1,Aliso 3: LJI131 1.\4134. )oH,\O, R " C °"NSO\,. R. . ".R 1969,]969 Monograph of theIh. plantplanl genus~cnul PorophyllumPomplwlllll" (Composi-l ompol'­ tae: Helenie Joj0 lar, H'lenicael.ae). Univ. KansasKansa' Sci.Sei. Rull.Bull. 48,48: 225-267.225 267 NS H"ro,TON, . .\t.M. C.C, & B. L.L TLR'I".TURNER. 1962. C'hromQSomcChromosome numbersnumber. of DyssodiaDrHodllJ K ,(Composilac:(Compositae: Ta~linae)Tagetinae) and ph)'lcricphyletic lnlerprela,lion,.interpretations. Rhodora 64:64, 2-15.2 15 Ell,, D. J.J. 1974.1974 NewI'.'e'" la.utaxa injn pectisPff,i, (Compo"lae:(Cornposjtae: Pectidinae)PCClid,nac) from Mexico\1exico and lhethe SOUlh",esl.rnsouthwestern UnitedUnlle

1975A.1975A Pectis1'....,;. cylindrica r, I"";,; ..(Compositae),, (C'ompMit ••established, o:lot.l>h.IKd as Ifa member• """mhn of d the,hot Texash.u floraflora andand confirmedconfirmed" as a• distinctdl~IIM:1 species..penn.. Southw.South.... NaturalistS.turali,' 20::.: 286-287.!86 2117. 1975B.IUSII Revision It."""" of ofPectisPUll' sect...n. Heteropectis IIr't'f'<¥'"fu (Compositae: Koml'O'iLac; Tageteae). T.~"'I, Madrono.\lMI,oIo 23:U,I81 181-191. I'll. 1977.1977. A revisionA '''''''001 of clPectis/W,n section1C<11On PeclothrixI'ulm"ri.r (Compositae: (Com~~. Tageteae). TaVlcw1 RhodoraRIM;Idu. 79:n: 32l! 78.78 __.& AT. I F.1-. STUESSY.S"""'Y. 1975.1975. ChromosomeChr_ numbersn..mlxn in ,nCompositaeCompo<>lK from f.­ thethe UnitedLl....., States,Sui.., Mexico\lultO.nd and Guatemala.C....lmALi. RhodoraRhodo... 77:71: 171-195.171 19S. LOVE,Lt»t. A.A 1974.1914 lOPBIOPIl.h'.....,..""., chromosome numbernumber reports",pore, XLV.XL\'. Taxon"la_on 23:U: 619-624.619 614 POWELL,Pol'"lili. A.A M.,\1 .• &.t S. SIKES.Sins, 1970. ChromosomeOr~ numbersn.. mlx~ of01 someoomr ChihuahuanO"h....hoIu DesertOt.nt Compositae.Compo&luc. Southw.SouIhw. Naturalist"'l..n~ ... 15:I~ 175-186.17S 180 __ •& ,,,",«" UnitedUnl1cd StatesSt.lr-s and.nd Mexico.MesIC" Madrono\hdfoM 17:17: 128 140. 11,"'.'.RAVEN, P. H.,H.. &&. D.D W.III KYHOS."rMOS 1961.1961 ChromosomeChromO'SO"", numbersnllmb!:crs inin Compositae.Comp of Dyssodia/)..." ..1", Cavanilles(".,.",11"" (Compositae:lCum/lO>llu; Tageteae).1.~...1 Univ.L"" C.hfCalif. Publ.l'uhL Bot.11<:0:. 48:411: 1I 88.&ll 1972.1972 ChromosomeOr_orne studies u...t1tS in ,nwestern ...... ,nNorth North American AltKnc.u Compositae. Compo;ilU Am. Juur.Jour. Bot.Rol 59:5.; 242-247.242 247. TURNER.I,"". IIB. tL, .. W.W L.I, EIIISON,E",,,,,. &"" R.11., M.M KINGK",. 1961.IW.I, ChromosomeCh,,,,no,,,nl~ numbersnun,l'>.ilU fromfrom the1M south-SOllI~' western.. 01t~rn UnitedUnm:d States.SUOI.... Southw.Soolll...' Naturalist"alur.li~t 18: 117-124.117 124 WIGGINS,WI(.c;.I\\. II. L., I .. &:& D.D M.\-1 PORTER.1'oIl1£0. 1971.1971 FloraFlo.. of thelilt GalapagosGalapap Islands.Island.. Stan-Sill>­ ford I'n"Univ. Press.!'rca Sunford.Stanford, xxu. + 99899fl pp.pp

I>EPARl\IF,,.DEPARTMENT OF BOT"BOTANY.... y 'IHFTHE 01110OHIO !>TATESTATE UNIVERSITYL"I\~RSITY I1]S1735 NEIL\[11 A\(\I'EAVENUE COLt\18t'S.COLUMBUS, OHIO 4J21043210

Pre-sen(Present Address: 810LOGICALBIOLOGICAL SnE\USSCIENCES DEI'ARHIE',.DEPARTMENT CALlFrnt"ACALIFORNIA POIPOLYTECHNIC YlFnl\IC SlATESTATE U'I\fRSITYUNIVERSITY SA'SAN lUISLUIS ORISPO.OBISPO, CAllFQR'IACALIFORNIA 4J40J93407