CHROMOSOME COUXTSCOUNTS OF COMPOSITAECOi\lPOSITAE FR01.1FROM THE UNITEDUXITED STATES.STATES, MEXICO,MEXICO. AND GUATEMALA'GUATEMALA1
D,\\,II)DAVID .r.J. KEILKEII, AND TOllTOD F. STl'Ei'SYSTUESSY
Chromosome numbersnumber.'> can be exb'emelyextremely u,;efuluseful inill .'>y,;temsystem atic studie>;,studies, particularly for hl'1pinl{helping to reveal e\'olutionaryevolutionary relationship".relationships. For the past1)1\st fifteen yean;years numerous chromo-chromo ,;omesome reports from plant"plants have been published,pllblishe,l. especially in thethe CompositHe.Compositae, and thesethelie counts have been compiled illin several major >:ouree>:sources (Darlin~toll(Darlington & Wylie,'Vylie, 1955;1!);')!i: Cave,Col\'e, 1!).,H-r,5;1958-65; OrndulT.Ornduff, 1967-69;1967-(i!); Fedoro\",Fedorov, 1969;1%9; Moore,Moore. 1970-72).1970-7:!), Howew!!'.However, a rapid A"lanceglance through these references indicatesindicate;; not only that many I\pecielispecies never have been counted, but ullioalso that many taxa are known only from a sillA"lesingle plant inin oneone population,population. In view of the common occurrence of euploid and aneuploid racesrace,; in plants asail illustrated by-by ,;evemlseveral (ldaileddetailed investigationsillve~ti~ations (e.g'"(e.g., Lewis,Lewis. 1962,1%2. 1970:1970; Stueslif.Stuessy, 1971a), it is desirable to haveha\'e several toto many counts from eHcheach specie,;species before accurate judgmentsjudg-Illents cancnn be made regardinl{regarding evolutionary relatiollshipsrelationships (Stuessy,(Stuessy. 1!"l71b;1971b; Kovanda.Kovanda, 1972:1972; Strothel',Strother, 1972).1972), The present paperpapel' helps tIlto rerne,lyremedy these deficiencies in the Compositae by: (1) re porting"porting fir>:tfirst chromosome counts for several genera,g"cnera, species, and varieties; aMIand (2) reporting additionala(lditionnl populational Chl'OlllOSUmechromosome counts for taxa documented previously.
MATERIALSMATERIAL:': AND I'R()('EDURESPROCEDURES
The meiotic chrolllosumnlchromosomal material for this ,~tudystudy was collected during the past '1everalseveral ycar,~years by thethe senior and junior author:;authors on various field excursions.cxcursioll~, Immature capit ula were killed and fixed in modified Carnoy's fluid (4(I chloroform: ;\3 ab'loluteabsolute alcohol: 1 glacialg"lacial accticacetic acid) and refrigerated in the laboratory until later preparedprepare(! by con-Cflll
1 'I'ublie"tion Publication :\"0,No. Rl:>843 fromft'om th,'the Departmentn~l'"rtment of Botany,Botany. TheThp Ohio St"t.'State University,\'ni"p".ity, ('Columbus.... Iumbus.
171 172172 RhodoraHhodora [Vol. 777i ventional acetocarmine squa.'lhsquash techniques. Voucher speci-speci+ mens collected by KeilReil and assistants are on deposit inin the herbarium of The Ohio State University (OS)(OS); ; vouchers\·ouchers collected by Stuessy are in the herbarium of the University of Texas at Austin (TEX).
RESULTS The chromosome counts obtained in the present study are listed in Table 1. First counts are reported for two genera, 16 additional sllecies,species, and one variety; 112 addiaddi. tional counts are for taxa counted previously, seven of which are new numbers. The first counts for genera are from Epaltes CassoCass. (n ==10) 10) and TricarphaT'ricarplta Long-preLon~pre (/I.(n = 8),S), and first counts for species are in BidensRidcnl/ L., GaleaCalea L., Guardiola Cerv. ex H. & B., MachaerantheraMac1wcranlhera. Nees, MelampodiumMeklmpodilon L., OtOPflJiPIISOtopappus Benth., SclerocarpusSclerOC(ll'pU8 Jacq., Srnrcio Senecio L., Sif/csbeckia.Sigesbeckia L., Simsia Pers.,Pel's., SpilanthesSpilallfhel/ Jaeq.,Jacq., T-rida:tTridux L., and Z(lmzamaZaluzania Pers.Pel's.
DISCUSSIONIlISCUSSIOl\' Because many of the counts counts presented herehere corroboratecOIToborate previous chromosomal reports,reports. the discussions are restricted either to first counts or to new reports for genera, species,specie.;;;, or varieties. The order of cornment.:\rycommentary will follow the se-se (illeneequence of tribes tl'ilJes in the classification of Hoffmann (1890.94),(1890-94), which is the same as that used in Table 1. References for statements regarding the range of chromosomal variation within genera will wilJ not be given; docnmentationdocumentation for thesethef\e counts comes from the several major sources cited in thethe introduction to thisthi~ paper.pape!'.
IWPATORIEAE.EUPATORIEAE. Counts for three herbaceous species of SteviaStevia Cav. representrepresent new reports.reporL.,. SteriaStevia elatior H.B.K.H.RK. is cited here as n = 121112n & 12i121 (Fig. 1), whereas thethe twotWll previously recorded numbers have been n = 34i3<11 (Powell & Tlirner,Turner, 1963) and nIt =• :l3133i (Grashoff, Bierner, & NorthNorth+ illgton,ington, 1972). Our count {orfor StcviaStevia ongrwoidesoriganoides H.B.K.,H.R.K., 1975]1!.17ft1 ChromoMmeChromosome Counts -• Keil & Stue~;;yStuessy 173
reported here as nII = 11J1 (Fig. 2), isi" the first for thisthi,; taxon at what appears to be the diploid level; the previouspl'eviolls cOllntscounts were n = 34134t and n/I ::-:-= ca. 4:143 .....± li1] (Gr.1shoff(Grashoff 1'/et al., 1\)72).1972). Stm'iaStevia plumml'rflPplummerae A. GmyGray var.var, rhU"ulIYCllsisdurangensis Robins,Robins. has been reported before only once by GrashoffGra,;holf etrt 1/1.al. (1972) 11Sas nII = ca,ca. 17, but our count isi;>. n1/ =.J= 44iII (Fig".(Fig. :q,3). AsA" pointed out b~'by Gra:-lhoffGrashoff ptet tr/.al. (1972), it isi,; common to findlind "al'ringvarying meiotic chromosomal associations and numbers in specie;
Asn:REAKASTEREAE. Sl'\'eralSeveral previous count;>.counts have beenbcen reportedrel>orteil fOl"for EriflerollErigeron knl"lI'iI,s1."itmIlSkarwinskianus DC.'DC: 2/12n =:{2= 32 (Carano.(Carano, 1924;1921; Rattag-lia,Battaglia, 1950)l!).-;O) ; 2/1 2n = 36:1(; (Fag-el'lillil,(Fagerlind, 1947;1917; Larsen, 1953,19,,:1. 19.-;·1;1954; Kliphui;;Kliphuis & Wieffering,WiefTering, 1972); nIt = 9 and n/I = 27i27, (Turne;·.(Turner, Elli"on,Ellison, & Kill~,King, 1961)19(1) ; andanrl n = ca,ca. 27 (Turner.(Turner, Powell, & King, 1962). Considel'in~Considering thcthe variation in chrochro- mosome number that has beenuccn documented previously]lrcviowdy in this spcde.",species, as well as our new reportI'eport of nII = ,,"5n & 17i17[ (Fig.(Fi~, 4),,I), it is likely that B,E. karwinskianus"(lrWill.~kiaIlIlS i;;is apomictic throughthroug-h at leailtleast part of its ]'anf{erange from Mexicoi\Iexico to northernnOl'thern South America (S()lbri~,(Solbrig, 1962).19fi2). It is interesting that our count come"comes from a pormlatinnpopulation veryvcry nearneal' the locality cited by TunlCrTurner ctet al. (19GI)(1961 ) forfOl' their counts of n1/ = 9 and n1/ = 27i.27[. MarJlflPlulIfhcl"/lMachaeranthera ('ou/t(Ticoulteri (A. Gray) TurncrTurner & HomeHOl'ne (a"(as P.~ilactisPsilactis I'u/dlericoulteri A. Gra,\')Gray) was reportedreporterl as nn:-: = i)5 by 801Sol- brig-,brig, Anderson,Andcrson, Kyho'i,Kyhos, Raven,Ra\'en, and RüdenbergRi.idcnbcrg" (19(;(1964).I). Jhl\reveJ',However, based on the recentreccnt revision of sect,sect. PsilactisP,~i/(/cti.~ of Mflcllfll'mJlfhenlMachaeranthera by TUl"nerTurner and Home (196.1),(1964), the geo-geo gnqlhicgraphic location of the voucher\'ollchcr for the count seems more aplll'olH'ialeappropriate for M. adduarida Turner & Horne than for M. cIJ/tU!'l'i,coulteri. The lattcrlatter "pecics,species, as recentlyrcccntly interpreter!,interpreted, is known only from the \'icinityvicinity of Guaymas, Sonora, where our voucher \\'aswas collccted.collected. OllrOur first count of n/I ="= 5 (Fig.(Fig. ii)5) for Macllll(!1"((1/thl'/'aMachaeranthera CO/t!tl'ricoulteri is particularly inter-inter. esting becau"ebecause in thcthe previouslypre\'iou;;ly mentioned revisionrevi,'l,inn of sect. P.,il(II·li,~Psilactis of the ~enusgenus (Turner & Horne, 1964;1961; ef,cf. their \7·1174 Rhodora {Vol.[Vol. 77 .. 0 •o er* •*•• 00-&0 q0 o ...%| - "... ."" ~ •• reo 0 •• • 'to 0 IIQ °o° •••• 1 2 03 04o
~ ~. \0 '4..* ^ ~«**... «for« 6 5b Iff o >6 • ' .." ,• - ~'<£.. 7 *•..«~ . " • ...~ . •« 8 8 9 ,^ - o -, "0*o 0 • J -, o~ 0 00 , •% 1 o/.r; , • \ , .. .. o JII.,o""41 k \Jo 10 00° o 11
•tr••... : ... •••• •Va .-.i>•.: . ':""',.,.f".':1 r?yv•• • .:%er..;~ + 12 13 '. !-'ib'1<.Figs. 1-22.1.22. Camera lucida drawings of meiotic ehromo30m"schromosomes of Sl~ci"sspecies of Coml'0.itae.Compositae. llil'loI"n",Diplotene, Fig.Fig". 18; diakin"sis,diakinesis, Figs.Fij{s. ;;.5, 7-10,7_10, 12,15,19-21;12, 15, 19-21; metaphasemet"phast' I, Fi"$.Figs. 1,I, 3, 4, G,6, 11,Il, 14,H, 16,IG, 17; metaphasem"laphase IT,IT, Fi"s.Figs. 2 (on,>(one half of 1',,11cell shown), 13.13, All filo."'l'esfigures sa1lWsame scale.S('( ~ -~.. ,-:.-.,. .. ., 4, ";."=I~ JJ .. -' _ F3LT» 17? 16 ~~{I ~1~t''°0 ^ O0 18"«W ~ , () CC* & ,• *<) 20 ., '-0w x o0 o~ - V J .... 5p 21 *4 ^ & if\22 ril'lIrillriparia \',:IJ',var. orr/nu:I",refracta, KCKG 8710, mIl = 12;12: Fig.Fil:'. 8, (i''''''r/;'II"Guardiola platyphyUa,1,/"1111'''1111,,, 1 KMK,l1S33S, 8558, 11n = 12; Fi~,!1,Fig. 9, .1[,I"ml'"I/i"",Melampodium appendiculatum,up/ln"li""I"t,,,,,. KCKG S701;A,8706A, ••n = 10; Fig.fo'i~, 10, OI"I"'I;""~Otopappus imbricatus,i",/",;",,,,,.,, KCKG !!I12,,,9112, w = 16;Ie: Fig.F"i~, 11,II. Partheniumr"rlh,,,inm "''''",wm,incanum, KMK,ll 77r;'-,A,7765A, '"'n = 18nlS11 & 18i;lS[; Fig.Fig:, 12, ScleroSrl,'ro· c"rp!l-~carpus : Fi/,;.Fig. 3), M. coulteri on morphological grounds was placed in the x = 5 cytophyletic group along with M. aridaarid(l and M,M. crispa (Brandg.) Turner & Home,Horne, both known chromo somally as n11=5. = 5. More recently M. ariz011ieaarizonica Jackson & ItR. R. Johnson and M. yawparvijlora ¡flora A. Gray haveha\'e been added to this group and both species have been counted as n = 5 (Jackson & Johnson, 1967). This fir.'itfirst chromosomal report (orfor M. coulteri substantiate.'!substantiates its phyletic association with these other species. All other taxa in sect. PsilactisPsi/netis are known chromosomally as either n1/ = 4 or n = 9 (Turner &&Horne, Horne, 1964). INULEAE. The fir;ltfirst reportI'eport for Epaltes (E. mexicana),me;ric(wa) , n = 10 (Fig.(Fig. 6).6), is in keeping with its present subtribal di.'ipo;litiondisposition in the Plucheinae. Of the related genera of the samesame subtribe (Hoffmann.(Hoffmann, 1890-94) that areure known chromo-chromo .'iomallysomally (BIl/mea(Blumea DC,DC., PlucheaPlllc}lPa Cass., PterigeronPtrrigcron (DC.) BenthBenth.,.. PtrroenlliollPterocaulon Ell., Spl1neTanUllUJSphaeranthus L., and TessariaTc.~saria Ruiz & Pav.), all are based on .tx =• 10 except BlumeaBlumra which appears multibusicmultibasic with .tx = 9, 10, and 11. On morphologi cal and geQgraphicalgeographical grounds, in OUI'our opinion, EpaltesEpnltes mexime.ri can~tcana Less, is quite similar to ;lOmesome species of Pluchea,Plllchea. the former differing mainly in its .'imaJlersmaller heads and flowers and in itsits epappose achenes. As emphasized by Bentham (I8n)(1873) and Godfrey (1952).(1952), the generic boundariesboundarie.'i in thethe Plucheinae are not well defined and perhaps should be re-re eV:l.llI:l.ted.evaluated. lIELIANTIiEAE.HELIANTHEAE. BidensBidrlls ripal'iariparia is reported forfor the first time as n = 12 (Fig. 7) in a genus that has most frequently counted number~numbers of n = 12, 24, lindand ~686 (clearly based on :l'x = 12). The first count for Calea zacatecl1ichl~zacatechichi Schlecht., n = ca. 19, is in keeping with previous reports forfol' other species of the genus (n=(n = 9, 16.16, IS, 18, 19, 24, ~2).32). According to the mostmO.'it recent revision of the Mexican and Central American taxa (Robinson & Greenman, 1896), C. zacatechichizacatcchichi is most closely related to C. ne/soniinelsonii Robins. & Greenm. which has lJeenbeen counted as n = ell.ca. 18 (Turner et aI.,al., 1962). The 1975] Chromosome Counts -• KeilRei! & Stuessy 177 mOI'pholoJ,:"icalmorphological and chromosomal hetcrog-cneityheterogeneity within Ca[roCalea and thethc absence of a recent l'edsionrevision of the entire genus suggcstsuggest that a thorough modernmodcrn .~tudystudy is much needed. (;uunliola,Guardiola, a genu~genus of about ten species.species, has been placed traditionnl1ytraditionally in the subtribe Melampodiinae (Hoffmann.(Hoffmann, 1~90-9l).1890-94). Howevcr.However, recent studie!'.studies byuy the junior author sUg'l{estsuggest thntthat on morphologicalmorpholog'ical and cyto!og-icalcytological evidence itit belong'Sbelongs morcmore properly in the CorCDpsidinaeCoreopsidinae (Stuessy, 1973). Our first COlllltcount of 11n = 12 (Fig. 8)R) for G. platyphyllaP[lltlfphyl/a A. Gray is consistent with the recent reportsrepOlis of n = 12 for both G. tlllocarpw~tulocarpus A. Gray (Grashoff etct afal,.• 1972) and D.G. mexicanamNictll1fl IT.H. & B. (SolbriJ..:",(Solbrig, Kyhos, Powell, & Raven,Ra\·en. 1972),-lfl72)," and with the base nlllnbernumber of x = 12 forfor se\'eralseveral other members of this subtribe.' The count of nII = 10 (Fig. 9) isis a first report forfOl' !U('/llmpodillntMelampodium aplJClldiclilaflill/.appendiculatum Robins. In a recent revision ofof the J..:"cnu."genus (StuCR."y,(Stuessy, 1972) this species is placed inin seriesserics CllpuhltaCupulata of scct.sect. Mcflllllpodillm;Melampodium; three other related species (M.(M. cN}nllotllmcupulatum A. Gray.Gray, M. roseiro.~, i Robins., and M. f'lIdllOlttenellum Hook. & Arn.) also are known chromosomally asa.~ n=/I = 10 (StllesS~',(Stuessy, 1971b).l!.l71L). The addition of this new count increasesincreaseR thcthe number of ;;pecicl'species "un'cycdsurveyed within thethc genusg-enus to 27 Ol:ltout of:nof 37. The first gcnericgeneric reportrejlOl"t for OIQjJflPPusOtopappus (0(O... scaber~("ab('r S. F. Blake) haRhas been publi! 'The-The COllntcount pllulj"h~~lpublished brby ~"lh]"j"Solbrig ,Iet "I.al. was li~tl~1listed fo,-for G.(;. IlIrip/;('"iatriplici /Ohflfolia A. (;ray.Gray, but in theIh<, m,'~tmost r,'C<,ntrecent publishedpubli~h<,d revision"""ision of 1h<,the g-enus,L...... nu9 (n"bjn~on.(Robinson, 1899)18~)9) Ihi_this <,pithetepithet is ""~"I"ftregarded...d allas "rnonrmoussynonymous with G.C. 'nemexicana..rieallll. 3 ((e.g.,.... 1':"•• Bidens1!idl'lI,~ 1..,L., C(J"'''I', Parthe»illmParthenium L., and particularly P. ar,qenlat1lmargentatum A. Gray, has been studied extensi\'elyextensively for many years (cf.(d. HammondHanunond & Polhamns,Polhamus, 1965), including a comprehensive revision by Rollins (1950). Parfl!('11i11mParthenium illCallllmincanum H.B.K. has been re-re ported previously as having a polyploid series of n1~ = 18, 27, 36, and 45, but our new count is n = 181118n & 18118i (Fig,(Fig. 11).11). 'T'hi.~This interploid number could represent the product of hybridization between P. incanumincmmm and other species of the genus that J,::'rowgrow in the vicinity,\'icinity, such as P. argfltlafum,argentatum, but our voucher specimens :'Ihowshow no morphological indicaindica~ tion of such intergradation. Alternatively, the meiotic conCOIl~ figurntionfiguration couldcould indicate a hybrid between n = 18 and 11n =:16= 36 cytotypes of the !>arnesame species. The plants under comiderationconsideration alfwalso could be apomictic, a condition that isis known to occur in populations in the northern range of P. ii/callUmincanum (Rollins, 19(0)1950) where our material was collected. The COllntcount of tln = 11 (Fig. 12) for SclerocarpusSclcl'Qcarpu.~ spatu.~paf1t latllslatus Rose is consistent with previously reported numbers of n = 11, 12, 14, and 18 in the genus as recently defined by Feddema (1971).(1971). The closely related genus,~el1US, Aldama LaLlave & Lex., is known chromosomally as n11 = 17]7 (Turner f'let 01.,al, 1962; Powell & Cuatrecasas, 1970; Feddema, 1971). Sige.~ueckhlSigesbeckia L. of the subtribe Helianthinae is a small genus of less thanthim ten species. It is worthwhile to mention that a cloReclose morpholoR"icalmorphological resemblance]'esemblance exists withwith Tri !Jono.~/lermllmgonospermum Less.LeM. (i\1cVallgh(McVaugh & Anderson, 1972; Stuessy,Stue;;.sy, 197:1)1973) and perhaps also with RumfordiaRlImfordin DC,DC., the former of the subtribe Melampodiinae and the latter of the Helian thinae. OU]'Our COUlltcount of t/.n = 15 is a first report for S. agrestisayresfis Pocpp.Poepp. & Endl. All but two other reports in thethe genusj{enus [n[n = 12 (Subramanyam & Kamble, 1967) and 2n = 20 (H.'lu,(Hsu, 1967) for S. orinlfaTisorientalis L.] have been either n=n = 15 or n = 30. Infraspecific euploidy is known to occur in S. ol'icllhlTiSorientalis (Mehra, Gill, Mehta,Meht.'\, & Sidhu, 1965) and it is now documented for S. jorullensisjOl'UlletlSis H.B.K. by our countscounls of n =]5= 15 and 30 (Fig. 13).J3). Only one count of 1/n = 15113 (Sol(301 hrigbrig et al.,ah, 1972) hashaff been recorded previously forfor thisthis species. In;,) 1975] Chromosome Counts -• Keil & Stuessy 179 Of the approximately 353.'i species of Simsia Silllsiu recognizedl'ecognized by yariousvarious authors (Blake, 1913,19I~, 1!H7,1917, 1928; Cuatrecasas,Cuatrecasa,~, 1%1;1954; Robinson & Brettell, 1972), ten havehaye been counted from morpholos;ricallymorphologically diverse parts of the gl'nus,genus, and :Illall count.~counts ha\'ehave been n = 17. Our first counts of n = 17 (Figs.(FiJ.:"S. 1414 & I.j)15) for S. rl/l·y/l'pi.'feurylepis S. F. Blake and S. grayigm!li SchSch.-Bip..•llip. exex S. F. Blake emphasize the chl'omosomalchromosomal uniformity within the genu:'!,genus. SpiltlllfJu.sSpilanthes with upproxim:ltel.\"approximately GO60 species (Moore,(:Moore, 1907) i:>is a t:lxonomicallytaxonomically complex genu;;genus much in need of revision-revision [lryary attention. It appears to belong in the :'!ubtribesubtribe GalinGalin• .'!oginaesoginae rather than in the Helianthinae [ISas traditionallytraditiOlmlly placed (Jloffmann.(Hoffmann, 1890-9·1).1890-94). Chromo;;omallyChromosomally the situation:-iluution al;;oalso is complex. Even thoughthous;rh only ,;ixsix species haveha\'e been counted, fourfOlll' base Humbers,numbers, .1'x = 7.7, 12.12, 13,1:1, and 16,JG, arearc present. Our firstflr.'!t report of 11n = ca. ".j45 for £.S. o('ymijoliflocymifolia (Lam,)(Lam.) A.A. H. :\IooreMoore add.'!adds anotherarlOther c1lromo'lomallen:!1chromosomal level toto the already chromosomally dh'ersediverse sect. "Sali\'uria""Salivaría" (==(= sect.;;ect. Spj/ulIl!lrx)Spilanthes) known with 1/n = 7.12,7, 12, Hi.16, and 26. AllAIlI)reviolis previous re[)orti\reports for .'lectsect.. .Armella4/'I11(/ll/ (Rich.) DC. have been clearly based on .1'x = 131:1 (only nII = 131:1 and n = :W26 counts reported).rl'ported). Our :lCWnew report of 11=n = 4111 (Fij..(".(Fig. 16)1Ii) forfOl' S. phaneractis}lhIOl('l"II('li.~ (Greenm.) A. H. Moorej\loore increll.~(>Sincreases the chromos()ma!llivelchromosomal diversity..~it~· of thi~this section as well. TricillpimTricarpha is ,Ia J.:"enus genus of two spL'Cie,;species recently described by !.ong-prl'Longpre (1!l70).(1970). Our firsttll'st count forfol' the genus (from T.7'. d!lI·allfJ('lIsi.~durangensis Longpre)LonJ.:"pre) of n11=8 = 8 (Fig".(Fig. 17)17) substantiatessubstantiate;! itsit.'i presllmpth'epresumptive close relationshiprelntionshi,) tnto SHlm:;il/Sabazia Cn,;;;.Cass. (n(II = 4,I, ~, 8, and Hi)16) and SI'/ll/(!Selloa Kunth (II(n = 8) as mentioned by Long-},onj..(" pre (J!J70).(1970). The problem of ,genericgeneric delimitatiolldelimitation in the sllbtribesubtribe Galinsogillae.Galinsoginae, ill\'o!dng"involving TJ'il'llrphu,Tricarpha, Sabazia811/mzi!1 andanrl SI'//OllSelloa a;:as well as (;lIlhlNOflllGalinsoga RuizRui;>; & Pavon,Pm'on, Stenocarpha81/,1/('('111'])1/11 S,S. 1".F. Rinke,Blake, TridrO"Tridax and Ja"{fI'rl(/Jaegeria Kunth,Kllllth, is much in neednCI'd ofof further study, de"pitedespite the appeHl'anceappearance in recent yearsyeaI'" of se\'cral several excellent revisions (Powell, 196::);1965; TUl'Iwr,Turner, 1965;l~Hl.-); Torres, 1968;1!l6~; Longpre,Longpl'c, 1970). Part of the diflicultydifficulty in .'iodsort- ing out the properpropel' aflinitiesaffinities of all the taxa in the Galinsllg-iGalinsogi nae is that [lre\'iou"previous workers workers ha\'ehave been working from thethe 180 RhodoraRhodQra [Vol.l Vol. 77 perspective primarily of a single genus and not fromfrom a perspicacious overview of many of thethe genera within thethe subtrihe.subtribe. An added difficulty is the absence of a recentrecent revision of GlifinsogaGalinsoga (most recent treatment that of Robin-Rohin son,son, IR94),1894), the understanding of which clearly isiR central toto sortingsorting out these generic relationships. QlIrOur first report for Tr;tlm:Tridax {f'"lliloihltenuifolia Rose, n = 9 (Fig. 18), fits well with the established base number of x = 9 for !lect.sect. T,-idlIXTridax to which T. tCllui[oliatenuifolia belongs (Powell,(Powel\, 196!i).1965). Of the 14 specie,.,species of Zall/z(wiflZaluzania recognized by Sharp (!9:lrl),(1935), six have been counted with definite numbersllum1Jers of nn=l&= 16 and 18. QurOur first report, n11= = 17 (Fig. 19), for Z. (JmyalUlgrayana Robins. & Greenm. firmly e,<;tablishesestablishes this as a new chromosomal level for the genus (a previous count of n = 17 ± 1 for Z. montagnaefoliamontomwc/olia Sch.-Bip. was reportedreported uyby Powell and Turner, 1£163).1963). HELENIEAE.HELF.NIEAE. ScllkllhriaSchkuhria pinnatapil/nala (Lam.) Cabrera has been counted before as 2n = 20 (Co\'as(Covas and Schnack, 1946), and var.val" virgalavirgata (LaLlave) Heiser of the same species has been reported a:o.as nw = ca. 20 (Turner 1'1et al., 1962). Recently fo.IcVaughMcVaugh (1972) transferred S. anlhcmoidcaanthemoidea (DC.) Coult.COlill. val".var. flual1'1IlalCtlBisguatemalensis (Rydb.) Heiser to S. lJillnala,pinnata, and our counts of n =)0= 10 (Fig. 20) forfOi' this taxon are the firstfit'st reports. A count of nII = 11 (Table 1) was obtainedobtailled fromfrom material tentatively identified asliS S. (wlhl'moidru.anthemoidea. However, our voucher specimen differs from thethe chal'aeterscharacters of thisthis speciesspecies as delimited by Heiser (19.lfi)(1945) in having more numerous disc florets and large raymy florets, as in S. schkuhschkllh rioidesrioidN'. (Link & Otto) Thellung in Fedde.Fedele. In pappus strucstJ'uc ture, though, our plants are much more similar to S. antheanthl' moidrnmoidea than to S. sch!whl'l·oidrs.schkuhrioides. This collection may reprerepre sentsent a previously undescribed taxon. RENECIONEAE.SENECIONEAE. Our first reports of nII = en.ca. 30 for Senecio hal'fll)cr,iihartwegii Benth. :lIldand 11n = 22 (Fig. 21) for S. i-iincinatus1'1l11citwtuJ; Less,Less. Meare consistent with counts reported previously for nthel'other taxa of the genus. Thirty-three specie!!species of SenecioSCIlI'r;rJ 1975] Chromosome Counts •- KeilReil & Stuessy 181181 have been reported a"as Itn = 30 and six arcare known with nIt = 22. Although the Kenusgenus is based either on x.1' = 5 (Barkley, 1962)191;~) or .rx = 10 (Ornduff,(Ol'ndufT. Raven, Kyhos, & Kruckeberg,KruckeberK, 19(;3;1963; Orndllff,Ornduff, Mosquin,r.to.~quin, Kyhos,Kyho.'1, & Raven,Ra"en, 1967),19(7), the diver sity of haploid numbers isi.~ great, representing 30 different chromosomal levels from nII -= fi5 to n = ca. 92. 92. CICHOR1F;AE.CICHORIEAE. The small genus~enus PinnfopnpplIsPinaropappus Less,Lei's. has been counted from only one species, P. 1'().'{I'II.~roseus Less.,Less.. and the reported counti!counts arcare nIt = 9 andaml 18 (Darlington & Wylie, 19551955: ; Turner ctet Ill.,al, 1961;1961 ; Powell & Turner, 1963196:~; ; Powell & Sikes, 1!)70).1970). Our present count of n = 20n2011 & 1rli (Fig. 22) is a new report for thisthii! taxon. The meiotic configura-confi~ura. tions of celli!cells in our preparation were irregular with bridges,bridges. lagging'lagging chromosomes.chromosomes, and varying\'arying' numbers of univalents.uniyalent.~. ACACKNOWLEDGMENTS K NO\\'LEOG"IENTS Field work for this investigationinvestiJ,:'ation was completed largely with support from NSF~SF grant GB-30240;GB-30~.IO; publicationpublic~\tion costs werewerc defrayed partially lWby funds from NSF grant GR.GB ~7678.37678. Thanks ai'eare extended to Judith M. Canne and Lyle A. ?lIcGlllMcGill for assistance 011on collecting trips.tril)s. Table 1. Chromol'tlmeChromosome counbcounts ufof Composit:l.eCompositae from the United States,Stales, Mexico,)!exko, and Guatemala.Cntemala. QC ChromosomeChro~l' Taxon Locality and voucher"oucher number (n)(n)"a EUPATORIEAE ..Iutrnl,.",Ageratum <"Ol'l'lIIOO",.",corymbosurn ZUClClIJ{.Zuccag. :'oIEXlCO:MEXICO: Sinaloa: 16.4 mi. NE. :'\E. of Santa Lucia,Lucia, 10 KC 88~9.b8859> BritkdliflBrickellia <"ON/lrricoulteri A. GrllyGray )tEXICO:MEXICO: Durango:nuran~o: 8.8 mi. S. of Nazareno,~aureno, KMK:\l 9 802;;." 8025.C Sln'WIStevia flu'I./nloln',ltlJanadenotricha (Robins.) MEXICO: !)urant:o:Durango: 12.7 mi. SW. of La Ciudad,Ciudlld. 1212 GrosholTGrashofF KGKC 8883. tSI,r;lJfStcvia "'"lio'elatior U.n.H.B.K.... GUATE:lIGUATEMALA:.... LA: Gualrmal.:Guatemala: 12.8 km. E. of Cd. 12n12[[ &&: 12i12[(Fig.1) (Fig. 1) GU:l.tPmala,Guatemala, K 9396.<19396.d ao tHIn';"fStevia oriY"'loi&I.8884. pi tSln'iflfStevia I,IlHIt"U'rfleplunimerae A. Gra)'Gray )IEXICO:MEXICO: Durango:f)uranlto: 9.-:9.7 mi. NE.I'E. of La Ciud3d.Ciudad, KC 44i44[ (Fig.(Fig.3) 3) '·ar.var. dllm"I1""ai.odurangensis Robins.nobin5. 8927-1.8927-1. SI.-";"Stevia salicifolialirifolifl Ca'·.Cav. :'oIEXICO:MEXICO: Chihuahua: 9,59.5 mi. W. of GeneralGe"tltral Trias, 12 '·:Irvar. salicifolia/it"ifolia K.llKM 8288. ASTEREAE Aph""'Uff'J'lIlUlAplianostephiis flri.o"inuarizoniciis "f:XICO:MEXICO: Chihuahua: 10.4 mi. S. of Cd. Chihu3'Chihua 4 .....A. GrnyGray hila,hua, K:\lKM 82l'3A.8263A. Apltflno.I'lmuApílanoste pit us M_~iui"'JfIlram-osissimus nCoDC. )IEXICO:MEXICO: Chihuahola:Chihuahua: 2-124 n,i.mi. 1'.N. otof Cd. Chihuahua, 4 S 1097.~1097.e < c< Aster exilis Ell. MEXICO: Sinaloa: 11 mi. XE.NE. of SanlaSanta Lucia, KC , ~ 88508850A...... AslerAster alli~osllsspinosus Benth.Ilellih. l\1f';XICO:MEXICO: Coahuila: 5.5 mi.mt E. 01of Nazareno, KMK,11 9, 8010. Baccharis glutinosa Pers. )IEXICO:MEXICO: Sonora: ArroyoArro~'o ClIchujaqlliCuchujaqui on ÁlamosAlamo!! 9, C;"i"",obaGuirocoba rd.,nl., KC 8675,8675. Conyza sophiaefolia H.B.K. MEXICO:)lEXICO: Sinatoa:Sinaloa: 11II mi. NE.:\K 1)1of Santa Lucia, KC 9, 8848. =r~ >•<, t:riyrr""E r ig ero n tli,"di i 'erg ry,·"" ens MEXICO:)tJ::XICO: Chihuallua:Chihuahua: 39 mi.mi, W. of Cd. Chihuahua, , 9 '" o Torr. &:& A. GnI}'Gray S 1025. 3 3 o •J. tf:l·iy"ro".fErigeron l"'f,.'ill"l;i""u,karwinskianus DC. IIC. MEXICO:)IJ::XICO: Veracruz:,'darru"t: 11.1 mi.mt E. of Puebla-Vera-J'uehla_Vera_ 5n & 17i (Fig. 4) o" Cru1cruz boundaryboundar)" Onon Rte.Hte. 150-D,150-1l, KCKG 9178.!H711. 3 o Grindelia sp. nov.f MEXICO:)IEXICO: Chihllallua:Chihuahua: 2 mi. S. of Cuauhtemoc,CU:luhtcmoc, , 6 O S 1041.IOH. g ".II"dltlrr'I"II,rrn**Machaeranthera ~""lIf'ri(A.coulteri(A. Gra}')Gray) MEXICO:lIEXICO: $onora:Sonora: ca. 15 mi. E.Jo:. of Guaymas,(;u:l)'ma", KCl,e 5 (Fig. 5) = Turn,'rTurner &:& HörneIlorn(' 8r.:l7.8637. • .IIMachaeranthera" .."m·rn"'h..ro ,,;1111/11pinnatifida ifi4" .'IEXICO:MEXICO: Chihuahua: ca. 2 mi. W.W. of Parral,I'arml. S 4 • (Ilook.)(Hook.) ShinnenShinners "'1.993. .IIMachaeranthera"dltlrrml'h.. m M'"brrl/n.scabrella MEXICO:)IEXICO: Co.lluiECoahuila:.. : 110 mi. S.~. of Piedras1'i(' INULEAEI:":ULEAE "''':1'/11/''1/***Epaltes mexicana,"'11"11"11 Less.Le~ MEXICO:lIEXICO: Veracruz:"eranUl: 33.5aa.;; mi. E.};. of PozaI'on Rica,Wea, KCKG 10 (Fig.(Fig. 6) 9211. HELJANTHEAEHELIANTHEAE Iffc Ambrosia artemisiifolia L. UNITED STATES: Illinois:IlIino;.: LAKEuxJ: CO.:ro,: LakeLak" For-For 18 est,,"t CoU"J::eCollege c.mpu§,campus, S 1132. Ambrosia psilostachya DC. :\IEXICO:MEXICO: Tamaulipas:Tamaulipu: ca.~a. 2 mi. SE.SF.. ofQf Reynosa,llernOlla. 1818 S 774 (ca. 18). UNITEDU:-IITEU STATJo;S:STATES: Texas:Tun: KINKIS NEYSEY CO.: 42 mi. NW.~w. orof Eagle Pass,'>au, S 877a,871a, b.b. Ambrosia psilostachya DC. :\fEXIOO:MEXICO: O\ihuahua:Chihuahua: 7 mi. :-IW.NW. ofor Cuauht.em(K',Cuauhtemoc, ca.~:a. 3636 S 10C2a.. 1062a. Ambrosia psilostachya DC. UNITEDU:-:ITED STATES;STATES: Texas:TUail; BRAZORIABUZORI" ro.:CO.: 1212 mi. 54 XE.NE. of Freeport,Fre<"purt, Til",,,,,Turner 5,475747 (ca.(ea. 1)4);54); GALVESTONIHLH:s'l"OS co.;CO.: 60 S. Thereu,Theresa, Galveston,Gah'eclon, Miller.Hill" s.n.,.n. BIIltiBaltimore...orr ""cllIrecta IL... GUATF.:\IALA:GUATEMALA: Sa"'aSanta Rosa:ROlla: 3.5 km.11m. NW.XW. of Culi-Culi. 15 apa, K 9403.94.03. Q Bid"Bidens... pil_pilosa L. varvar..... minori,u,r MEXICO::\lEXICO; Guerrero:Gunrf'ro: 10 mi. S. of Chilpancingo, KC 12 a o f. ",i_ I' (Blume)(Blume) ShertrSherff f. minor 9126. 2 BitlnuBidens piloJnpilosa 1...L. Vllr,var. radiataf'fIdiolfl MEXICO:MEXICO; DillinioDistrito Federal:.-~f'ral: 17.7 mi. W. of D.F.-D.F. 1212 SchAllp.Sch.-Bip. in Webb"Webb & Berth.Berth, Puebla boundaryboundar')' Onon Rte. 190-0,190-D, KC 9164. f. dOlldindo/i"dondiaefolia (1--"5.)(Less.) Sh"rtrSherff Bitl,"~Bidens pil_pilosa 1..L. ":at,var. rotlilJlnradiata ;\1F.XICO:MEXICO: Chihuahua;Chihuahua: 15.5Hi..5 mi. W. of Santa Lucia,Luda, 1212 S~h.-Bip,Sch.-Bip. inin Webb &:& B"rth.Berth. KMK.lf 83W.8304. ff...... ,intradiata.. ••**BidensBidnlll ripDr;"riparia JI.H.K.H.B.K. "ar,var. "I'Mfn.refracta MEXICO;MEXICO: ~,,_:Sonora: 5.5 mi. E.E, of AI'TO)'oArroyo Cuchujaqui 12 (Fig. 7)i) (Brundll'.)(Brandg.) O.E.O.E. &hub Schulz on ,Uamo,·:\liJpilluÁlamos-Milpillas rd., KGKC 8710. l.'ftlmCalea IIrl~{oliourticifolia (!o!iIl,)(Mill.) DC. GUATEMALA: Guatelllala;Guatemala: 1;.86.8 11m,km. NE.~K of Ama-Ama. 19 tilla.titla. K 94.30.9430. < " o **Calea zacatechichi Schlecht. MEXICO: Jali:>to:Jalisco: 1.2 mi. ~W.NW. of :\1:aKdalpnaMagdalena :airair- ('a.ca. 19 port,po'Irt, KC 9008. CJir/lMIllh,IIIlUlChrysanthellum mexicanumm~rir;l""m )IEXICO:MEXICO: Nayarit:Xa}'aril: ca.ra. 18I!' mi. SE. of Tepic,Tellk, KCliC 8 • -,0 (;""<"lIm,Greenm. 8965. .=; C"R",Cosmos..,t lillruriJoIiuRlinearifolius (Sch,·Rip.)(Sch.-Bip.) :.n;xICO;MEXICO: Durango:Huranll:n: 10.510,:' mi.ml. SW.S\\'. ofnf LaI"'" Ciudad,Ciudall, 23 Hemsl.lIem~1. var.\·lIr. linrllriJ"/i,,.linearifolius KCliC 8892.811ftZ. ('''R''I/.RCosmos fl',n·ijl"rrtparvifiortls.• (JaC<'].)(Jacq.) Pers.Pen. MEXICO:)IEXICO, ....Sinaloa:i"alo.: 16.416,4 mi. NE.XK of SantaSanla Lucia,"uriI', 12 KC 8858. " a v~ Dicranocarpus parviflorus A. Gray MEXICO:}IEXICO, Coahuila:C03hui1:l: jctn.}ctn. Rte.Hlf'. 40 &&: id.nl. to10 San 1010 »i, c" 0 Pedro,I'l."dro, KMKJf 7991. 3 Ecliptaf:dip'" alba (I..)(L.) Hassk.Ha~sk. o<5= ,,11'(1 MEXICO:).IEXICO: Sin.loa,Sinaloa: 14.5U..5 mi. S,S. of Sinaloa·SonnraSinaloa-Sonora 11IJ M boundary on Rte.HI .... 15, KCliC 8736.8.36. C•0 1 = FI""cri'lFlaveria tri""",'i"trinervia eSI,n-ng.)(Spreng.) ).IEXICO:MEXICO: Jalisco:J.liliiW: O'Ocotlán,...lll1n, KCliC !I('3fi.9036. 1818 re C.C. }lohr.Mohr. G "0 G~I,..Ii"/nGuardiola "I.mexicana H. &:& B. Gu~ntro: j T;l..~~o, ~'IO', c ri,.""n H. R. :\n;xICO:MEXICO: Guerrero: 7 mi. NE.:";'t:. ofnf Taxco, KC 9107. 121211n &&. frag.fmlt. " 0 ··GlUlmi**Giiardiola..l" 1,1"'Nlm/llfnplatyphylla A. Gra}'Gray UNITEDL:XITI;n STATt:S,STATES: Arizona:A';>.On.: COCHISEPl('IIISt: "u.:Co.: ca. 10.510.5 12I:! (Fig.n'ij{.8) 8) =~ mi.....i. W. of Coronado Nati.Xall. Mem.:\I('m. headqtrs.,h",adqln<.. KMK.\' • 8558.~. I Helianthus laciniatus A. Gray MEXICO: Durango: 2 mi. NW. of Bermejillo, S 17 ;lH:XICO: Iluran2Q: 2 mL XW, of Il<"mlPjillo, S 17 1.~, 940. '" Heterosperma pinnatum Cav. ).IEXICO:MEXICO: Sin.lo.:Sinaloa: 3.8 mi. S\\',SW. ofnf !oilln!.Santa Lucia, 252;:; KGKC 88.13.8833. r!'"' Jaegeria hirta (Lag.) Less. ~ J'UI/uiu hirt" (Lag" 1.t:"M. Gl'ATB;lIAl.A:GUATEMALA: AltaAha Verapaz:\·tr.paz: 4 mi.ml. NE.Xt:. of San 18 s. Pedro Car-h:i,Carcha, S r.!l6.596. MEXICO:)IEXICO, Durango:nuralll'" 8.38;: • ~• mi. SW. nfof LaL:J, Ciudad, KClie 8903C,89u3C, 8fflf.);8909; 4.8".Il mi. S\\,SW.. .,fof I..aLa Ciud:Ciudad,...I, KCKe 8916; 15.61:1.6 mi. NE.Xt;. of La Ciudad. KC l\92!t.8929. :\licht)llonn:Michoacán: :;454 mi. E.K .,rof I'.amora,Zamora, J,:CKC !l(lo(;U.9060. Puebla:l'u..bl.: Tételes,TN..I S ":.Ill.498. Sina-:,in:l_ 10:1:loa: :1.83.8 mi. ::W.SW. of Santa Lucia,I.uda., KClie 1'o~'2A.8832A. X Ja,ycnaJaegeria macrOCCl'hala•macrocephala Less. MEXICO: Michoacán:~Iiehoac:in' 262G mi. S. of Pátzcuaro,f'atzeuaro, S 18 G86.686. Veracruz:Verauu7" 8 mi. N. of Jalapa, S 491, 492. *.* *MMelampodium~blml",dil.'m aP1'~lldicul«lu"lappendicidatum ~IEXICO:MEXICO: Sonora: 0.3 mi. N. of Arroyo Cuchu-Cuchu 10 (Fig.(~·ig. 9) Robins,Robins. jaqui on Alamos-GuirocoooÁlamos-Guirocoba rd., KC 8672A;8G72A; 4.7 mi. E. of ArroroArroyo Cuchujaqui on Álamos-MilpillasAlamos·~lilpillas rd., KGKC 8706A. Mewml'odiumMelampodium CIlpuwlumcupulatum A. Gray ~IEXIGO:MEXICO: Sinaloa: 27.327.:1 mi. S. of Sinaloa-Sonora 10 boundary on Rte. 15, KC 8742A. ,11cu/mJ>QdiumMelampodium P3 HF.LEmEAEHELENIEAE :l.Q"'" Bfl/tiaBahia ul.sil1/hi/l)/iaabsinthifolia Benth. var. :'IIEXICO:MEXICO: Coahuila: 0,20.2 mi.mi, E. of El Numbre,Kumbre, 24 O0 •1 dt'alllllilldealhata {A.(A. GranGray) A. Gray KMK.ll 7963 (ca. 24); 8.98.~\ mi. N.No of RanchoRancha Acatita, ~ KM BOnA,8073A. Bahia pedata A. Gray MEXICO:~IEXICO: Chihuahua: 29.9 mi. S. ofaf Cd. Jiménez,Jimenez, 12 KM 8221A.8221 A. UNITED STATES: Texa~:Texas: CULBEReULBEIl 80:<;SON co.:CO.: 3.7 mi.m;, W. of Van Horn, KMKJf 7688.7GBB, PRF.g/l)1OPRESIDIO co,:CO.: 13.113,1 mi. SE. of Presidio, KM 7776.7776, OysRooia.Dyssodia uureaaurea (A.(A. Gra}')Gray) Neb.Neis. UNITEDUXITED STATES: Texas: BREWSTERIIRF:\\"STF.R CO.:co,: BigHil:" BendBeud 8 var. poll/chattapolychaeta (A.(A. Gra}')Gray) Nati.Kat!. PkPk.,.• 8.7 mi. N.:-:. of headqtrs.,headqtrs,. KM 7821A. M. C. Johnst.Johnst. DyRRodi"Dyssodia pa/'PORIIpapposa (Vent.)(Vent.) Hitchc.Hitchc. MEXICO: Plwhla:Puebla: 8 mi.m!. E. afof Puebla.Puel!la, KCKG 9172. 1913 HI/RRodinDyssodia pMltud",ctupentachaeta (DC.)(DC.) .MEXICO:\lEXICO: Coahuila: 2,12.1 mi.mi, N.X, of Ranchon"ucho Acatita,Acatita. 16 Itobin~.Robins, subsp.subsp. ptl1tuc1Ultt'lpentachaeta var.":lr. KMK.\f SG66,8066. beleniFlorestina tripteris DC. MEXICO:)IEX1CO: Coahuila:("""huila: 63Ii:! mi. S.:-. of Piedrasl'iedrll~ Negras,X'·lIn'~. 10'0 KMK.I/ 7847A.78~7A. Durango:DUralll:O: 16]l'i mi. S.S, of Nazareno,Nluareno, -,• KM1\.\1 8030A. 803M, Gaillardia pulchella Foug'. UNITEDeXrrl·;n STATES:~l'A1'ES: NewSf'''' Mexico:)I.. ~i(n: GRANTGIlMIT CO.:n,.: 0.31).:1 17 " mi. !'.S. of jctn.jrtn. Rtes.H,c~. USl:~ 180181),!" & NM~:'l 61,GI. KMJo:.II 8460A.8lGOA, " t:X1T~'n STATES: Xc..' :'Il"~ieo: HIDALGOltlll\U"oO t:",: 14 Gaillardia pinnatifida Torr. UNITED STATES: New Mexico: CO.: 14 1717 ~ mi. NE. of Lordsburg,Lordshul'lI, KMK.II 8491A.1l,1!!)A. -s", :0:. o0 (;"iI/,,,.,/illGaillardia pinnatifida Ton.Torr. var.'·"r. MEXICO::\IEX1CO: Chihuahua:('hihuahua, 2 mi.",I. S. of Cllauhtcrno.-,Cuauhtemoc, S 17 3 /,;''''''Ii/i'''' o0 I;I/o,,"'~linearis (1:)<110.)(Rydb.) Bidd.fll;d SENECIOl\"EAESENECIONEAE OdontolricloumOdontotrichum Hi1!ualltlllsinuatwm Mf;XICO:MEXICO: Jalisco: 1.2 mi. NW. of MagdalenaMall;daJena air-air_ 30 (Cen·.)(Cerv.) Rfdb.Rydb. port, KGKC 9{l11.9011. Schistoc"r"lwSchistocarpha 0oppositifolia p1ffisit;/()} ia MEXICO:)lI!;XICO: Veracruz:"eracruz: 11.8 mi. W. of Tuxpán,Tuxpan, KC 8 (Kuntze)(Kuntze) Rydb.h Rydb.h 9222. **Senecio hartwegii Benth. MEXICO:~lEXICO: Durango:Durl\n~o: 8.3 mi. SW. of LaI.a Ciudad,Ciudtld. ca. 30 KC ~!lOf"8906. **Senccio runcinatus Less. MEXICO:'IEXICO: Veracruz:\·cn,HU1.: cn.ca. r,6 mi. E. orof LasJ.:u Vigas,\'iJ{U, KC 22 (Fig.O"i~. 21) 91!12.9192. MUTTMEAl-:MUTISIEAE ~T.-\TE~: Triri!lTrixis ..califomica"lif"r>!;"" KKellogg...lIol!:l!: UXITEOUNITED STATES: Texas:Tcu!': BREWSTERIlRt:W! ,I,HOP-IF:ACICHOPvIEAE E tl>i""rr,,..,,'p,,.fPinaropappus roseusrau,,, Less.IA."D. MEXICO:lIEXICO: Veracruz:\'cr,I('ru7.: 11.1 mi. E.E, "rof Puebla-Vera-ru..bla'\'l'ra· 202I1rr&IIIFil!;,22)n & U (Fig. 22) 1~>I1·n. cruz boundary onnn Rte.Htp. 150-D, KCKG !11'j'j,9177. no G= -t:nlc.!<»Unless indicat",1indicated ntb"otherwise,.....·i*, theth" reportedTPpnrtprl meiotic",pioli<' ('btorunrom"chromosome numbersnU1I11 ... r.o representrl'l'n'~l'nl bivalents.h;'·al.. nt~. • "KC»•KC = KrilKeil ,c..& C"ltlt~:Canne; ~'OlcRM = "".il.(·Keil & .l1rr.ilI:McGill; "f.:<'K = Keil;K"il; P:-;<\S =~ Stuessy.,"11"«'" I ,\'ou('h"p^Voucher d"'"rmlneidetermined bybr P.II. L.I.. Turner,Tun eA.J.. M.:\1. Powell,PO"'dl, hR.hr:. :\I,·\'au<:"h.McVaugh. ''-il'1'l*First report....""rt for \aril'l)',variety, ...**species,1'.'0;; ***genus.• ..~....IU •. 2.O. ;'1\_-{•New r~I"'rtl'(lreported nun,h"'rnumber fn.for t:non.taxon. '" to 192192 Rhodora [Vol.(Vol. 777j LITERATURE CITED 1l,IRKr,l-:\·. BARKLEY, T. M. 1962.19/i2. A revisionr"v;s;on of SenecioSc"uio aureua,Wr'·II.' Linn,l.;nn. and allied ~I\(",;es.species. 1'nons.Trans. Kansasi\.1n~as Acad.Acm!. Sci.Sei. 65:6;': 318-408.318-408. IlATTAGLIA,BATTAGLIA, E.F.. 1950.19.>0. L'alterazioneL'alt('razionc della meiosimejosi nella riproduzionel'iproduzion" apomitt;apomittica...a di ErigeronJ-:ri!I' fOIl KarwinskianusI\"r,,·i». IIF.I;lF:R,HEISER, C.G. 11B.,.. JI:.JR. 1945.I~.l.j. A revision,"(',-[s;on of lh"the genusI::enu~ Schk'l"r''''Schkilhria. Ann. )Ijsrolll'iMissouri not.Bot. (;:ml.Gard. 32: ~".j·278.265-278. HOFnlA;'>';o;",HOFFMANN, O. 1890·!J~,1890-94. Compo",t"",Compositae, 1'1'.87-:101.pp. 87-391. In/11 A. ENGLER~;;O;"GU:R & A.K. PIl,\;,>,Tl.,PRANTL, Die natürlichennaW,.l,.. I"'n Pflanzenfamilien,I'Ilnoz,'nf"milien, vol.\'01. H·;).4(5). Leipzig.J... ipzil::. Hn:,Hsu, C.,C,C.-C. 1067.1967. l'rdimin"ryPreliminary chromo. POWELL,1'0\\'1':1.1•• A. :\1.M. 1965.l!1(l:,. T"xollomyTaxonomy of TridaxT,.ill", (Compositae).(Conll'0silac). Brit-Brit· Ioniatonia 17: 47_!lG,47-96. -- ," &". J. CUATBECASAS.CU.\TR'X",ISMI. 1970.197\). ChromosomeChl"QrnoMrne numbersIlUl11bel'~ inin C<;>m)",sitnf':Compositae: CillomhianColombian :mrland VenezuelanY,·"<,zueln,, species.speci,',. Ann.A,m. Missouri:\lis!'()ul'; Hot.Rot. r:al'dGard... 57:i7: .17-\-379.374-379. -----.• • , & S.R. !'i1l":S.SIKHS. 1970.\!l'jO. ChromosomeChl'"mo~om(' numbersnUI"I",,". of somei'(lme ('hi.Chi- huahuan <1"""1"1desert Compositae.romp".it"c. South\\'.Southw. Naturalist:';:ntur:tlisl 15:I:;, 175-186.17.'·18r.. -----, , & R.n. L. TURNER.Tl'R:st:ll. 1963.19(1:1. ChromosomeChromownl(' numbersnumbers inin thethe Cornpos;t:Compositae...:. VTI.VII. Additional .,,,·ri,-sspecies fromf,"(lrn the south",,,.!southwestern...,.,, United St"tCgStates """and Mexico. MadroñoM:lflr()]io 17:11: 128-140.128.140. ROBINSON,P',.IITX.«(lX, n.R. L. 1894.1891. NotesK()I"~ up""upon the genusll:cnus ",,1;"'011'"Galinsoga, Proc.p,"f>(', Am. Arm!.Acad. ArtsArt.. 29: :12.;·:127.325-327. . 1899. RevisionP."l'ig;on of thetl,... genusl.'"tJnug Guardiola.r,,,,,,·l1i,,lo. Rull.Rull. TorreyToney Tl,..(.Rot. Club 26: 2:'12.23232-235..•• -----, •, k& JJ. . .\1.M. GI11:t::-:M,\X.GREENMAN. 1896.18!1r.. Revision1:'-"';.;011 of ~'·H1(.\.\U;"T,\)I,SUBRAMANYAM, K.,K.. & N.X. 1'.P. RAMBLE.K \\lBU:. 1967.I~)i;'. hiI,,: : IOPBIOI'Il ...chromosomeh,.,.,mo"ol11<: numLHnumber n'I'OI'lsreports XII 'faxonTaxon 16: :~"O.350. T'>1mE~,TORRES, A,A. )1.M. 1~1r.8.1968. RevisionJ:c\"ision of JaegeriaIn,'yui,, (Coml'o."ita(Compositae-Helian...... llclian ththeae)... uc). B,·;tloniuBrittonia 20: .-,2-,,1.52-73. TUR:-:rn,TURNER, B. L. 1[11;."1965. Taxonomy of Stn""",)"!,""Stenocarpha (Coll1l'o(Compositae-Heli-.• it:,,--lIcli_ allth<:'l('_"nlin,,<)~jna<'),antheae-Galinsoginae). ~"ulll\\·.Southw. NaturalistXnlul':lli,,( 10: 238-240.2~8-24U. -----" , W.\\'. L. ELLISON,1':1.1.''''':-:, & R.It. M. KING.K,:-:,:. 1961.191;1. ChromosomeChrol1losonw num11UI1I I~-l'"bers in the COIllI'O";I,,,,.Compositae. IV.1\', NorthXorth American,\m"r;... an "11<'species,... i",,_ withw;lh phyI'h~' Idicletic ;nte"l"","t,,tioll".interpretations. Am. Jou,',Jour. Bot.Ilot. 48:l~: 211;·22:1,216-223. -----" , & D.I). HORNE.H"lt: IlErARnIE;>.:TDEPARTMENT OF lJOTA:-iYBOTANY TilETHE 01110OHIO :'T'\T~:STATE U:\ln:ltSlTYUNIVERSITY (,OLl1:'1lJU~,COLUMBUS, 01110OHIO ·1;~2\()43210