Number Seven De cember 1978

Ro al Bot anic Charles Je ffrey , Edi t or , Herbarium , y Gardens , Kew , Ri chmond ,

Surrey TW9 BAB , U . K .

Financial support o f the NEWSLETTER is generously provi ded by Ot to Koel t z - P . O . 1 60 624 . . . Ant i quariat , Box 3 , Koenigst ein Taunus , B R D

BOOK REVIEW

THE BIOLOGY AND CHEMISTRY OF THE COMPOS ITAE , edit ed by V . H . Heywood ,

J . B . Harborne and B . L . Turner , Academi c Press , LO n don , New York and £ 118 in m 1 San Franc i sc o , 55 . Pp . 9 ( two volu es ) . 977 .

A brie f ac count o f ail the genesi s o f thi s work was given in t he last n In the openi g chapt er , the edi t ors give a bri e f hist ory o f

t he t axonomy o f the family , c onsider i t s vari at ion in t erms o f adaptat i on t o habit at s and pollinat ors and review it s chemistry in relat ion t o it s ec on omi c import anc e and the role o f me tabolit es as chemi cal de fence mechani sms . Turner gives an ac count o f i t s meagre fossil hist ory and it s phyt ogeography , c ons idering the family t o be po ssibly mid-Cret aceous in origin and explaining i t s present dist ribut i on in t erms o f c ont inental dri ft and rapid adapt ive radiat i on from Mi oc ene t imes onwards . Burt t relat es much o f the diversit y observed in involucre , fl ore t and frui t types t o selec t ive pressures involving m poll inat i on e chanisms , bree ding syst ems , ovary prot ec t i on , and seed dispersal . Leppik shows t hat t he pat tern o f evolut ionary modi ficat ion o f t he capitulum in relat ion t o t he sensory percept ion o f pollen ve ct ors parallels t hat exhib it ed by sin gle flowers . Je ffrey speculat es on t he possible evolut i onary impli cat i ons o f t he oc currence o f t he bilabiat e c orolla type . St ebbins evaluat es t he evolut ionary signi ficanc e o f studi e s in comparat ive development and func t i onal anat omy . I t i s un fortunat e t hat he has c hosen a taxonomi cally unsat i s fac t ory Senec io sensu lat o on whi ch t o base in part hi s argument in favour o f a shrubby ancestry for the Composi tae , for as a result , t he stat ement s he makes wit h respect t o t he so c ircumscribed are wi thout signi fi canc e for the argument e it her way . No clear a ffini ty wit h any other family i s c oncluded from t he admi t t edly equivocal evidenc e , and i t i s c onsi dered all t he spe c iali zat ions o f t he Composi t ae except for t he gamope talous c orolla evolved independent ly o f Baa o e similar spe c iali zat ions in other famil ies . g breaks int erest ing new ground in considering microcharac t ers o f t he ligule epidermi s in relat ion t o pollinat i on llum heliant hoid enec ionoid b iology and ; Eri o h and Arni ca are , not s , 2 .

Pollen morphology in the Composit ae and some postulat ed relat ed families is

revi ewed in a long paper by Skvar l a e t al . Of t he types o f exine ultrast ructure anth moid exhib it ed by t he Composi t ae , only t hat designat ed e is approached by

any o f t hese other famili es , it s c ommon possession doubtless a ple s iomorphy H nni ian A th moid an themoid i n t in t he e g sense . n e or modi fie d pollen s ' e wit in

Gor t eriinae Tarc honan thinae Er ioc e halus Osmit o s is Er iachaenium , , p , p , , Aden ocaul on G arrhena , ymn and t he Ant hemi deae i t sel f , while the apparently progressively more derived s enec ionoid and helian thoid types charact eri ze t he Heliant heae Hel enieae Eu tor ieae Cal en dul eae As t er eae In ul eae , , pa , , , and S ene c ion eae Elachant hus Cerat o e I s o e t o s is , as well as Cent ipeda , , gyn , p , l n Abr o tan e la , Ursinia and , i n modi fied form , the Ac t ot idi ae and Cat ananche . S olbr ig shows t hat t he chromosome cyt ology o f abou t if3 o f the family is known ;

even so , t he sample i s far from uni formly represent at ive o f all t ribes and

genera and other aspect s o f cyt ology are much more poorly known .

The abundance o f secondary metaboli t es in Composit ae i s emphasi zed by He nauer g , who discusses t he distribut ion in t he family o f members o f

charact eri st i c ubiquit ous and ot her import ant classes o f c ompounds . The

sesqui terpene lact ones are di scussed in more detail by Herz , who emphasi zes the signi fi cance for chemot axonomy o f knowledge o f t he biosynt het i c relat i onships Eu at or ieae He liantheae He len ie ae o f compounds . The af finity o f t he p , and i s h emphasi zed by t he ir possessi on o f eliangiol ide s , also recorded from Peuc e 11 h lian olide h while t he possess ion o f e s links Arni ca , Balduina , Ps il ot rophe wit h Helenium and Gaillardia ; Part henium and

Part henic e show obvious af finity wit h t he Ambr os iinae . Flavonoid variat ion ,

revi ewed by Harborne , appears to have great er t axonomi c s igni fi cance at generic - and lower t han at t ribal and subtribal levels . Re s c orcin ol based chalc ones and aurone s are c haract eri st i c o f C or eogs idinae and a few ot her members o f the H lian th eae He len i a e and e e . The flavonoid syndrome o f t he Composit ae is unl ike

t hat o f any o f the famil ies postulat ed as closely relat ed t o it . Polyac e tylene s (revi ewe d by Sorensen ) occur in all t ribes o f t he family and t he ab ility t o i synthesi ze t hem is probably anc est ral . While cr e penyn c ac id derived poly

ac etylenes are also developed in Pit t osporaceae , Araliaceae , Umbelli ferae and Campanulac eae , in the Compositae they are charact eri zed by the presenc e o f cycli c , aromat ic or het ero cycli c end groups . Two dist inc t groups o f tribes H l anth a are t he e i e e Hel enieae In ul eae on the one hand , and the Ant hemideae t ( except Cent i eda sand As e reae on t he other , although polyac e tyl enes are absent m from some austral genera , e . g . Vit t adina , Minur ia , Olearia , C el is ia , east ern

Elachanthus Is oe t o s is Abr ot an ell a . Australian Brachycome , and from , p and

Dit hiin Eu t or ieae Helian theae Hel enieae . c ompounds are restri ct ed t o pa , and Pharmaceut i cal and ec onomi c uses o f t he Compositae are considere d by Wagner in t he final general chapt er ; i t s ec onomic pot ent ial is far from being fully

exploite d .

There follow t ribal revi ews , st art ing with an almost complet e reworking o f Eu t orieae t he t axonomy o f the pg by King and Robinson , very welcome in vi ew o f t he very unsat is fact ory generi c t reatment o f Bent ham and o f t he important weedy er atum Aust r oeu at or ium Chr omolaena spec ies in Agerat ina , ég , p , Ayapana , ,

Fl e is chmann ia Heb clinum . Is ocar rmum idesmia , e and Mikania #ha , Mi cros , Le and Ilt is ia Ad no t Sene c ioneae c r it ot hamnus are ac cept ed int o t he tribe , e s les ( s ,

Mar s hall ia and Pala foxia (all Hel iant heae are not . The chemi st ry o f the t ribe ,

reviewed by D om1 ngue z , has obvious t axonomic pot ential but is at present t oo

li tt le known . The t axonomy o f the Vernonieae , reviewed by Jones , is st ill in an unrevi sed st at e and our chemical knowledge o f i t , as shown by Harborne and Williams , is also very inc omplet e . There i s some indi cat ion that most , i f not all , o f the Old-World spec i es included now in such genera as Vernonia and C ent rat he rum will prove eventually t o be generically di st inct from t hose o f t he New . The Vernonieae i s now t he least well known o f all t he tribes .

As t e reae . The , reviewed by Grau , are in lit t le bet ter taxonomi c order Ge is s ol e pis (perhaps ) and Is oe t ops is (perhaps Inul eae ) are anomalous in t he t ribe in fruit structure , and Ab r otan ella is excluded . Rham is A odoc e hala Nanno l ot t is Pla ioche ilus Ix1 ochla is placed near p p , g near g and s is segregat ed from Podo coma . The chemi stry o f t he t ribe , reviewed by Herz , is lit t le known ; sesquit erpene lact ones are unc ommon and the polyac e tylen e s resemble those o f .

‘ A t entat ive arrangement o f t he Inul eae int o three subtribes is propos ed by n ha t Merxmfille r , Leins and Roessler . A is oc e a and Fedde s are ac c ept ed but

S m h ll oc ar us Ant hemi deae ) , G arrhena Cardueae Dimer es ia (Hel iantheae E Osmit o s Tar c honanthinae s, , p t he Ar c t ot ideae Adenocaul on E r iachaen ium (probably Anthemideae ) , Het erolepis s, and

(b ot h Mut is ieae sare excluded , mainly on palynol ogi cal grounds . The generi c chaos

H lic h sum . o f the Gnaphalium e gy complex i s duly acknowledged The chemistry , reviewed by Harb orne , shows promi se but lit t le is known in det ail and even why some Anaphali s and Heli chrysum spe c ies smell o f curry i s le ft chemi cally

explained .

The revi ew o f t he He lianthe ae by St eus sy shows signs o f hasty preparat i on ; H en os t e hium Verb s inae Helian t hinae 1 i s re ferred bot h t o e group 3 and group , 4 h n 5 and (correct ly ) 45 are bot h quot ed for Fit c ia , and Melanthera is not rec orde d as present in t he Old World as well as the New (alt hough t he Old-World Li ha ta n - Melanthera spec ies are perhaps nearer t o po c e t ha t o typi cal New World ) .

Nevertheless , t he new syst ematy i s welc ome , like all such hypothe se s a st imulus Trichos ira Ph il o lo a t o furt her work . p is removed t o Vernoni eae , as i s g ss (prope rly

Liab eae . Mic r ol e cane and ) , Coreopsi s Cosmos are maintained dist inct from Bi dens wi t hout c omment . Ot her point s t o be queri ed are t he re ferenc e o f Mar shallia t o Eu at orieae G i s ol e i A t er eae I s ocar ha t he p , o f e s p s t o t he s , t he ac cept anc e o f p int o Galin so He tant hus Dile is the inae , and t he plac ing o f a group o f genera ( p , p Coult erella Te t ranthu Se ne c ion eae Flaveria i, Flaveria ) and g int o t he , where t hey appear dist inct ly out o f place . Swain and Williams revi ew t he chemistry .

Se squi t erpene lac t one s are confined , as far as is known , t am Bahiinae and Ambr os iinae ; chal cones and aurone s are known V1 and C or o idina Last henia as well as the e ps e . Gui zot ia appears out o f place in the

by t he presence o f thioether s , it would b e bet t er re ferred t o E c l1 t i nae W d lia Sh ar er ia m ear Eclipt a and e e . The wholly enigmat i c e i s ad it t ed

r rvat ion .

Turner and Powell review and dismember t he Hel en ieae . Some o f t heir prO posal s Ol ivaea Ri io us can be hardly disput ed ; such are t he re ference o f , g gapp and Thurovia As t ereae Il t i ia t orieae t o , Mi crospermum and s t o Eu , and o f various H i N urola ninae groups to t he el ant heae Neur olaen inae , Var illinae (included in e e S u i la St u by t e s sy ) , Ga l r dinae (ac c ept ed by St eus sy s, Bahiinae (ac cept ed by e s sy , Flore t ina P il t o hinae en O inae Gaillar diinae including s s, s o r and gym EEEE ( included in Bahiinae St eus s Pala foxia Mars hallia and re spect ively by y S. The plac ing o f (with s Eu t or ie ae Th o s is in t he pg i s more doub t ful , while zm p is acc ept ed int o the Bahiinae S t u Jaum in by e s sy . The e ae o f these aut hors is het erogenous and i t s plac ement in Cor e ops idinae (newly elevat ed t o tribal rank by t hem ) is likewise dubious . The ma j or controversy lies , however , in t he re ference o f a number o f S n ion a Bl nno rmat inae groups t o the e e c e e int o which , with the exc ept ion o f t he e spe

wi t h some re servat ion , they are not ac cepted by No r den s tam . The chemi cal revi ew 1 l o- - eude sman ol ide s by Bohm provides some guidance . , sec o are known in Ven e as ia e rmac ran o Eriophyllum , whic h , like Chaenact is , Bahia and g , also makes g An t h o hlor s S n t r ic h0 u li des . c are recorded in Last henia and y papp s . Chaenact i s disul hate d ol ac e t l e n e s Pala foxia S chkur ia and Eriophyllum have p p y y , as do and , u - - while Schk r ia and Last henia have polyac e tylen e s o f the ene t e t rayn e ene syst em , C ore o s idinae like most o f t he p . Flaveria and Lasthenia have t hiophenes similar

H r ic o h llum . t o those in Taget es , but t hiophenes have not been found in ype p y

Sulphat ed flavonoi ds are found in Last henia , Flaveria and Sartwe ll ia . Flaveria resembles Pect is and a group o f closely relat ed genera in t he C o re ops idinae in C t elaborat ing a Kranz syndrome ( h) phot osynt he t i c pa hway . In short , what is known o f t he c hemist ry o f t hose genera t hat Turner and Powell woul d re fer t o the Sen e c ione ae shows t hat t hey have a typi cally he liantho id chemi cal fac i es ; there i s no t rac e o f the s enec ion o id er emophilan e s and alkaloi ds so charact erist ic o f Arni ina P u h um t hat tribe . c e ( including e c e p yll ) are closely relat ed to the Gail lardiin ae Chae nac t idin ae E r i ll ina ; , t y e (perhaps het erogenous ) and possibly Cr o c idium and Bl enn o s perma appear t o fit between t he Bahiinae and Coreo s idinae ' St eu s s s Per it l inae Flave r iinae Jaume inae in y syst em , and the y , and (in part ; between t he Gaillar diinae and Coreo s idinae ; the affinity o f Welwit s c hiella

ve rn on io id ) is unknown . Cadis c us (like Doroni cum ) i s truly s e ne c iono id . The Ta et inae Ta e t e ae g are reviewed as a t ribe ( g 5 by Strother , who provi des a use ful key t o the genera . Morphologically and chemi cally (as shown by Rodriguez and

Mabry ) t hey fall within the helian th oid ambit , near t o Fl ave r iinae , Jaume inae Cor e o s idinae and E .

o n m a n n N rde s ta reviews the Li b e ae and S e e c io eae , t he former rece iving t he

tribal charact erizat ion and recognit ion i t has long warrant ed . From the lat t er

he c orrec tly excludes Arni ca and vari ous relat ed h e liant hoid genera . In t he Bl e nn os ermat in ae Isc hn ea Abr otanella Cr oc idium p , he includes and as well as and Bl enn os perma ; the lat t er two appear t o have li t tle affinity wit h t he former two . I s chne a Ab r o tan ella i s probably asteroid , near Brachycome , is perhaps a reduced H ianth a inuloid ; i t has kaurene dit er peno ids , which are more typi cal o f el e e , ' Eu t or ieae Be dfordia S ene c ione ae Rob ins s although also recorded in pa and in o f the . review o f t he chemist ry emphasi zes the taxonomi c i solat i on o f t he true S ene c ioneae .

Heywood and Humphrie s show in t he Ant hemideae a t ribe in whi ch progress

t owards a sat is fact ory taxonomy is st eadily being made , b oth morphology and

chemi st ry (reviewed by Greger ) making use ful contribut i ons . Excluded from the Pol ne He lian t he ae Inul eae ' but tribe are ygz ( ) , Cerat o e ( re ferred t o the perhaps As t er eae Pla io c he il us As t er ea b et t er in , near Brachycome s, g ( ear La eno hora Ab o tanella I hn ea an Cen ti e da In ul e ae ne r s c . and Soleno s, , d p probably , near ' Car esium s. Elachan thus and Dimor hoc oma are ac c e t ed withou t comment , t hough t hey p p p _ Minur ia Os mit o s is may be ast eroid , o f affini ty . p and Ursinia are ac cept ed wit h i enta nene reservat ion . Cent peda contains t he c ommon p y , whic h has ot herwise never t been found in As er eae or Anthemideae . Formal subtribes are not establi shed , but

a number o f informal groups is rec ogni zed . The southern hemisphe re genera appear h t o fall int o two groups , t he c o tul oid wit h c haract erist ic ant emoid acetylenes , e Er io c e halus S haer ocl in ium Adenant hemu m C mb o a us . g . Cotula , Soliva , Hippia , p , p , , y p pp , Leuc o t er a Aden o lo s sa Pe rous ia Lidbe ckia Thamni h llum p , g , y , , p y , Ine zia and Pent zia u and an r s inioid , devoid o f acetylenes but with charact eri st i c benzo furans , Las ios er mum Ph rmas ermum As aemia Eumor hia including Ursinia , p , y p , , p , Athanasia ,

St il n o h t um G o ent z ia . Tar ch onant hinae p p y and ymn g , al l con fined t o Afri ca The are E ioc e halus c otul o id not considered but may be relat ed t hrough r p t o the group , as Os mit o i S h ll o ar us G arrhena may be also ps s . ymp y c p and ymn should also be considered

as possibly anthemoid . The syst emat ic s and c hemistry o f the two unrelat ed t rib es Cal endul eae and

Ar c t ot ideae are revi ewed by Norlindh and Valadon respe c t ively . Our knowledge o f bot h tribes i s inadequat e in bot h respec t s . The Ar c t ot ideae are t aken t o - include four well marked subt ribes , including Er emo thamn inae whi ch fit s easily int o t hi s geographically c ircumscribe d but morphologically diverse assemblage . Plat car ha Mut is ieae y p (probably ) i s exc luded , Het erolepi s is not c onsidered .

The Cal en dul eae are l ikewise ge ographi cally nat ural but unlike t he Ar c t o t ideae are (except for Dipt er oc ome and Garul eum ) also morphologically uni form . Pre sent generi c concept s are based almost ent irely on achene dimor h oc o lic charac t ers and ne ed revision . Chemi cal (presence o f p e ac id ) and morphological (ligul e mic r ocharac t er s ) evidence favours the uni fi cat ion o f Cas t alis Os t eo s er mum Blaxium Dimorphot heca , and p sect . int o a di st inct taxon , for whi ch generi c rank woul d seem appropriat e .

Dit t ri c h divides t he tradit i onal Cardueae int o three dist inc t tribes , E chin o eae Carl ineae Car dueae Car duinae Ce ntaure inae p , and sensu stri ct o (wi th and ) , ' but Wagner s revi ew o f t he chemistry , while support ing thi s t axonomic division , reveals no di fferenc es emphat i c enough t o warrant more t han subtribal ranking for t hese four taxa . As a single t rib e , t hey are certainly more homogeneous t han , e . g . , Hel ian th eae , Ar c t ot ideae and Mut is ieae . Excluded are Cavea (re ferred t o t he Inul eae but surely near t o Saussurea and War iona ( t o Mu t is ieae ) as well as ar c t ot oid Plat c ar ha Mu t is ieae Er t hr oc e halum Gundelia ( ) and y p (probably , o f y p a ffinity ) ; Be rar dia i s acc ept ed from Mut is ieae .

Mut The is ieae themselves are revi ewed by Cabrera , being , l ike the Liabeae , t oo lit t le known chemi cally t o meri t a chemi cal review . Of t he four subtribes , Ba na iina uv n - t he r de s e and Nas s a ii ae are well de fined and homogeneous , the

Gochnat iinae and Mu t is iinae het erogenous and needing taxonomic elucidat ion . The - key to t he subt rib es on p . 1041 has suf fered disast rously in the type set t ing ; t he first line o f t he second lead in couplet B shoul d read " Corollas mostly Chiono us glabrous . Pappus o f simple bri stles ( or villous paleae in papp

( Liab eae ) i s excluded ) .

Closing t he tribal reviews are t hose by Tomb ( syst emat ic s ) and G onzalez

( chemistry ) o f the Lactu c eae . The trib e has been li t t le studied chemi cally and generi c boundari es in t he Lac tuca Prenanthe s c omplex are st ill unc ertain . The P D " important paper by . . Sell , Taxonomi c and nomenclatural not e s on t he Composit ae ub fam Ci hor ioid a 1 : 2 6- 266 1 6 s . c e e in Bot . J . Linn . Soc . 7 3 ( 97 ) should be added t o t he syst emat i c re ferenc es .

Mabry and Bohlmann summari ze t he chemist ry , Turner t he b iology , in the Chem1 c al l os it ae closing chapt ers . y , the Co are plac ed in dire ct relat ionship t o the Umbell i ferae and nulac b e i ng l inked t o both by t he sesquit erpene m lac t one and acetylene chemi st ry , t o t he former by the cou arins and the lat t er by inulin ac cumulat i on . The Calycerac eae have no acetylenes and contain sec o iridoids and are t here fore placed remot e ; but perhaps the presence o f t hi s - lat t er class o f compounds has been over we ight ed taxonomi cally , and they do - o ccur in t he other inulin c ontaining group , the Goodeniaceae Brunoniaceae alliance . Chemi cally , t he tribe s relate more t o one anot her than t o any other

Lac tuc ea S n ionea . family , even t he e and e e c e whi ch are most di st inct chemi cally A very use ful alphabet i cal list o f ac cept ed genera ( and some synonyms ) , wit h t he t ribal allo cat i on o f each , followed by sub j ect , organi sm and chemi cal compound indexes , complet es the work . In view o f the vast amount o f new in format ion t hat has ac crued during the

past 20 years , it is remarkable t hat the t ribal c oncept s o f Cronqui st are nearly

wholly support ed by t he data here present ed . Of ot her groups acc orded tribal b rank , both be fore and since , only t he Lia eae remain support ed as a dist inct

t axon at t hi s level . Of the groupings o f t he t ribes , t hat o f Wagenit z receives

most support . The sub family Cich or io ide ae includes t he t ribes in which the plesiomorphous anth emoid pollen type preponderat es and in which the sesquit erpene ma an o m lact ones are confined t o the ge r c r lide , el e anolide , guaniol ide and u man e de s olide st ructural classes ; t hese are the Lac tuc eae , Mut is ieae , Vernonieae , Liab eae Cardueae E c hin 0 eae Carlineae Ar c t o t ide a , ( including p and ) and e (including Er e mot hamn eae t r o id a The sub family As e e e , on t he ot her hand , commonly exhib it s n ion h h t he more derived s e e c oid and e lian t oid pollen types and , i n t he more

advanced tribes , development o f b iogenet i cally more advance d sesquit erpene lactone

Ur ini ae n h m - - s e ) have mostly a t e oid pollen , he liangiolide s and ene t e t rayne ene type ace tylenes ; As t ere ae share the lat t er but are depauperat e in sesquit erpenes and ' have helian t hoid pollen ; the Cal en dul eae resemble the As t er e ae in the lat t er respe ct but have pen taynenes ; advanced sesquit erpene lact one classes charact eri ze Inul eae Hel ian theae Hel en ieae C or eo s ide ae some members o f t he , ( including , p and Ta e t eae Eu t o r ieae g ) and , t o a l esser ext ent , pa ; the last two t ribes are also h linked closely by the development o f p yt omelan in s , helianth o id type pollen and

similar floral epidermis features ; finally , the Sen e c ion e ae have dist inct

e r emophilan e type sesqui t erpene lact ones and pyrroli zidine alkaloids , t he lat ter Eu t or ie ae known ot herwise (albe it wi t h di fferent necio aci d moit ies ) only in the pg .

C . JEFFREY

ARTICLE

NOTES ON THE EVOLUTION OF COMPOSITAE

B . L . Burt t

(Royal Bot ani c Garden , Edinburgh )

Sel f- incompat ibility i s widespread in Compositae and may well have been a

basic feature o f the family sinc e i t s incept i on . There t hus seems t o be li ttle reason for the di fferent iat ion o f peripheral flowers that have only a female

funct i on . This problem puzzled me when I spoke at t he Reading Symposium in

July 1975 (Burt t , Female rays , by O pening first , render the capi tulum - - prot ogynous and may contribut e t o cross pollinat ion in sel f compat ible s ,

for the st igmata are re cept ive be fore t he pollen o f t he disc flowers is shed . But this i s clearly a secondary advantage i f Compositae are primarily sel f

incompat ible .

Got t sb er er In Sept ember 1976 , at t he IAPT Symposium at Hamburg , Dr . G . g

gave a st imulat ing paper on pollinat ion by beetle s , whi ch i s wi dely beli eve d

t o be t he primit ive form o f zoophily i n angiosperms . He point ed out that the beetles are an ol d and numerous group t hat have always been present in the

hist ory o f angi osperms . The primit ive angiosperms may well have been beetle

pollinat ed , alt hough the canthar ophily now seen in many plant s undoubt edly G ott sb er er represent s a secondary re currence o f t his qe o f pollinat ion . g made 7 .

one observat ion t hat i s o f great importanc e in the c ont ext o f the evolut ion o f an thar o hil Compositae . He point ed out t he close correlat ion between c p y and - prot ogyny . The reason , he beli eve s , i s t hat unl ike ot her pollen ve ct ors beet les are long-t erm visi t ors t o flowers ; i f t hey are t o be e ffect ive agent s o f cross pollinat ion t hey need t o wipe t he fore ign pollen from their bodies on t o a receptive st igma on arrival at a flower ; ot herwise t he foreign pollen will soon

be swampe d by t hat o f t he flower ( or capit ulum ) i t sel f (Got t sbe rger ,

h As already not ed , female ray flowers c onfer e ffec t ive prot ogyny on t e

capitulum , and amongst t he basi c t ribes o f Composit ae they are part i cularly - c harac teri st i c o f t he yellow flowered Helian t heae and relat ed t ribes , whic h - are o ft en bee t le poll inat ed . It i s thus a reasonable hypot hesi s t hat female

rays evolved in relat ion t o cantharO phily . The fact t hat t he ray flowers have - b ec ome male st erile is not , o f i t sel f , o f any funct ional import ance ; it follows from t he fac t t hat t he floral me chanism o f Composit ae does not easily permit

prot ogyny in an indivi dual flower .

The above hypothesis gives an answer t o t he quest i on ' Why are t he capi tula o f some Composi tae The answer t o the quest ion ' Why are t he rays o f some sel f- inc ompat ible Composit ae female " ‘ is simply ' because it con fers ' func t i onal prot ogyny on t he capitulum . It is not t he primary quest ion .

A c orollary o f t hi s hypot hesis should be t hat s e1f— incompat ible members o f _ e non c antharO phil ous genera do not develop femal e rays . Wit hout c antharo phily t he peripheral di fferent iat ion o f a s emaphyll should retain the bisexu al nature o f t he flower or lead t o a neut er fl ower . This i s what we find : hermaphrodi te Mu peri pheral flowers in , for exampl e , St okesia (Vernonieae ) and Mut i sia ( tise ae ) , st erile radiant flowers in Centaurea Car dueae

The ot her st imulus t o t hought on Compositae , s inc e I wrot e the paper for t he Reading Symposium , has been t hat delivered at t he same meet ing by C . Je ffrey on the evolut i on o f c orolla type s i n t he family ( Je f frey , Je ffrey put forward t he unort hodox vi ew t hat t he b ilabiat e corolla o f Mut is e ae i s t he basi c type in Composit ae , and t hat i t has been derived from an anc est or bearing zygomorphi c flowers in an elongat e raceme . Al though I have come t o disagree wi t h t hi s view , it has made me t hink more thoroughly about relat ive primit iveness among t he t r ibe s o f Composit ae , a mat t er I had previously t ended t o dismi ss as unpro fi t able spe cul at ion .

The t hought s t hat now c ome t o my mind are these . In the first place zygo morphi c fl owers in an elongat e raceme would almost certainly have reached the stage o f having numerous ovul es and a more or less spe ci ali zed pollinat ion

1 61 . 228 mechani sm . They woul d have embarked on what I have calle d ( Burt t 9 , p ) t he individualist path , where a flower produces a large amount o f see d from a single succ ess ful act o f pollinati on , o ft en by a spe ciali zed pollinat or . : Composit ae belong t o t he ot her main t rend , t he collec t ivi st s there is an - agglomerat ion o f small one se eded flowers , so that each seed requires a separat e act o f pollinat ion , but pollinat ors ar e generalised . Now I do not deny for a moment t hat there are o ft en deviant s from a main evolut i onary trend , or t hat t here are such deviant s from the indivi dualist t o t he c ollect ivist pat h . One has only t o re call the conge st ed b i or uni -ovul at e flowers o f Selagineae and Gl obular ieae i n Scrophulariaceae . Nevertheless , t hese are rest ri ct ed groups o f l imit ed scope . In seeking t he or igins o f Composi tae we are seeking a group that could have advanc ed t owards t he capi tulum on a broad front . I find i t di ffi cult t o c onc e ive a select ion pressure that would have induc ed the sharp diversion o f a large group o f indivi duali st s on t o a collect ivist pat h . The pat t ern o f c hanges shown in Je ffrey ' s diagram o f corolla- forms is very much more plausible t han t hat involved in a scheme o f Composit e phylogeny that

start s wit h He l ian t heae alone . Nevert heless it assigns a derived st atus to t he most widely spread type : the 5-lobed t ubular corolla o f many hermaphrodit e

flowers .

- Since an elongat e raceme will t end to have mult i ovulat e zygomorphic flowers ,

i t i s , in my view , simpler t o look for Composi te origins in a corymbose raceme .

In such an in florescence the flowers remain erect and regular , but wi t h a t endency ,

‘ enhance d as the in floresc enc e bec ome s more compac t , t o peripheral di fferent iat ion

( c f . Iberis in Cruci ferae , for example ) .

1 At t he Reading Symposium , Leppik ( 977 ) restat ed his t hesis that , in t he

organi zat ion o f the capitulum , t he Compositae have passed t hrough t he same seri es o f ant he cologi cal stages t hat he had previously re cogni zed in t he ' evolut ion o f t he form o f individual flowers . However , Leppik s sequence in regard t o the flowers o f early angiosperms had as it s central t hesis t he idea t hat t he floral forms evolved in correlat ion wit h the sensory powers o f the available - poll inat ors that were themselves evolving alongsi de . It was a c o evolut ionary

sequence . Composit ae are a highly evolved family and i t must be almost c ert ain t hat t he whole range o f pot ent ial pollen-vect ors was already in exist ence when - t he first rec ognizable Composi tae appeare d . There was thus no c o evolut ionary sequence and no obvious reason why capitular forms should evolve in sequenc e rat her t han as an explosion o f direc t adapt ive radiat i on t owards t he whole range

o f pollinat ors .

There are four fundamental pat t erns in Com os itae : (a ) a head o f anthocyanic , t he relat ively long , tubular corollas (Vernonieae et c . (b ) liguli florous head a - - ( L c t uc e ae ) ; ( c ) the radiat e head , wit h 3 t oothed rays and 5 lobed disc H ian th a ( el e e , S ene c ion eae o ft en re duced to a di scoi d head by loss o f rays ;

( d ) t he head o f bilabiat e flowers (Mut is eae ) .

It i s one o f the st riking t axonomi c fact s about Compositae that , despit e the uni formit y o f the family , the divi sions be tween the maj or tribes are deep and t he " " number o f int ermediat e genera remarkably few . Thi s i s just t he situat ion t hat would be expec t ed i f there had been an early explosive diversi ficat i on conne ct ed with the evolut ion o f t he di fferent capitular types when this v irtually new floral organizat ion o f high pot ent ial int erac t ed wi t h t he wi de range o f potent ial

pollinators .

Re ferences

Burt t , B . L . , 1961 . Composit ae and t he study o f func ti onal evolut ion . Trans .

1977 . Aspe c t s o f diversi fi cat ion in the capitulum . In Biology and - Harb ourn e 1 : 41 . Chemistry o f Composit ae : ed . Heywood , Turner 59

t t s ber er G . 1 . g , , 977 Some aspec t s o f beetle poll inat ion in t he evolut ion o f

s . In Flowering Plant s : evolut ion and classi ficat ion o f — 1 : 211 226 . higher cat egories : ed . K . Kubit ski . Plant Syst . Evol . Suppl . C 1 . Je ffrey , . , 977 Corolla forms in Compositae some evolut i onary and taxonomi c : speculat i ons . In Biology and Chemist ry o f Compositae ed . Heywood , - Harbourne Turner 1 : 111 118 .

Leppik , E . E . , 1977 . The evolut ion o f capitulum types o f t he Compositae in t he

light o f insect fl ower int erac t i on . In Biology and Chemi st ry o f n 1 : 61- 8 Compos it ae : ed . Heywood , Harb our e Turner 9 .

CURTIS CLARK , Department o f Botany , Universit y o f Cali fornia , Davis ,

Cali fornia 95616 , U . S . A . RESEARCH : Experiment al syst emat i c st udy o f Enc elia wit h D . W . Kyhos , experiment al syst emat i c study o f Eschscholzia ( Papaveraceae ) , invest igat i on o f UV floral pat t erns in Composit ae and ot her famili es . TRAVEL

Desert s o f Cali fornia , Ari zona , Sonora , Baj a Cali fornia , Baj a Cali fornia Sur ,

1 8 1 . Spring 97 and 979 , collect ing Enc elia and relat ives

KYHOS DONALD W . and CURTIS CLARK , Department o f Bot any , University o f 616 Cal i fornia , Davi s , Cali fornia 95 , would be grat e ful for viable achenes G raea s ea o f Encelia , Enc eli opsis and e and especially o f Encelia osa ( Texas , N ew Mexic o ) , Enc elia linearis (Nayarit ) , Encelia hi s i da (Galapagos ) , Encelia _ p halimi fol ia ( Sonora , Ba j a Cali fornia Sur ) , Verbesina an us t is s ima (Nayarit ) , Verbesina vir at a (Cent ral Mexi co lia raven ii (Ba j a Cali fornia ) and

Ge rae a vi sc i da (Cali fornia , Ba j a Cal An quant ity great er t han two i s adequat e .

DR . W . DENNIS CLARK , Department o f Bot any and Mi crobi ology , Ari zona St at e

Univ i T m 8 f ~ ers ty, e pe , Ar i zona 852 1 , i s working on t he syst emat ics o

Haplopappus sect ion Haplopappus o f Sout h Ameri ca . Denni s R . Bi scho ff , a student m o f Dr . Clark , i s currently doing syst emat i c research on I s oc o a Nut t . v " Gregory K . Brown , a student o f Dr . D . J . Pinka a and Dr . Clark , has complet e d a Plat hkuhr ia bi osyst emat i c study o f ys c Rydb . and i s current ly working on Sout h Pinkava D J Ameri can Haplopappus . Dr . Clark , Dr . , Dr . . . Keil (Department o f

Bi ologi cal Sc i ences , Cal i fornia Polyt e chni c Ins t itut e , San Lui s Obispo ,

Cali fornia 93407 , and Dr . R . A . Mayes (Department o f Biological S cienc es ,

Virginia Polyt ec hni c Inst it ut e , Blacksburg , Virginia 20550 , have recently H n o thr ix l oomi ii complet ed a st udy on t he ori gin o f yme s Blake .

DAVID R . GIVEN , Bot any Divisi on , DSIR , Privat e Bag , Christ church , New Zealand . RESEARCH : Taxonomy o f introduced Composit ae in NZ botani cal region ; taxonomy As t ereae Celmis ia o f Composi tae t ribe in Aust ralasia e spec ially , Olearia and derivat ive genera ; relat i onships be tween Sout h Ameri can and Aust ralasian E r i o i ger d genera ; floret t richomes and classi fi cat ion . N . Z . Journal Bot any 11 : 86- 6 - 7 9 loc . c it . 13 : 547 56 loc . c it . 17 in press RE"UESTS FOR MATERIAL : Aut hent i c fl oral mat erial o f Composit ae t ribe t As er eae (worldwide ) .

DR . WALTER A . KELLY , School o f Natural Sc iences , Mesa College , Grand Junct i on , 81 01 Colorado 5 , is current ly start ing a research programme on Agoseri s .

DR . JOHN C . MITCHELL , Dermat ology Laborat ory , Shaughnessy Hospit al , M 00 V6H N1 5 Oak St reet , Vancouver , Brit ish Columbia 3 , Canada , announc es the

J A Green ras . publi cat i on o f Mi t chell , . , Rook , . , Bot ani cal Dermat ology , g s Lt d , 6 1 8t h 9 W . 2 Avenue , Vancouver , Bri t i sh Columbia V5H 2H4 , Canada ; 800 pp . approx . , 10 .

8 publ . January 1979 . Pri ce $2 (US ) unt il £31 . 95p . a ft er

Has mat erial o f c hemotaxonomi c int erest c oncerning Compositae , plant product s and c ompounds irritant , toxi c and allergeni c for the human skin , eye and respirat ory trac t .

PRITH PALS INGH D hban dhu I , e s College , University o f Delhi , Kalkaj i , New Delhi

110019 , India , report s recent publi cat ions

DAKSHINI , K . M . M . and PRITHIPALS INGH , 1977 . Numerical Taxonomy o f t he

genus Blumea DC . ( Ast eraceae Inul eae ) in India . Phyt omorphology

DAKSHINI , K . M . M . and PRITHIPALS INGH , 1978 . Sec t ion Hier aciifoliae

Rand . o f t he genus Blumea DC . ( Ast eraceae Inul e ae ) . Taxon - 27 : 21 25 .

H PRIT IPALSINGH and DAKSHINI , 1977 . St ructural variat ions in t he style and pollen present at ion me chanisms in the genus Blumea - in India . Bot . Mag . (Tokyo ) 90 : 291 299 .

PRITHIPALSINGH and DAKSHINI , 1977 . Chemotaxonomic studies on

t he genus Blumea DC . ( Ast eraceae Inul eae ) in India . Acta Bot . - Indica 5 : 143 159 .

PULLA T . IAH , Department o f Botany , Andhra Universi ty , Waltair 530003 , A . P . , : odi India . RESEARCH Embryology and syst emat ic s o f Composi tae , lam t humu Car s , As t er eae , Vernonieae , Sene c ion eae . Phyt omorphol o 2 - Bot anique (Nagpur ) 7 : 63 70 Acta Bot . Indica 5 Mature viable achenes o f any spe c ies o f He lenieae request ed .

T M 6 001 HANIKAI ONI , inst it ut Francais , B . P . 33 , Pondicherry 05 , India , the recent work o f Miss Vasant hy Ge orge o f t he same Inst itut e

- u VASANTHY , G . Structure e t t erminologie de la paroi sporo polliniqe ; Ast eraceae : Vernonia and Cen t rat herum APLF Document s pour la r eun ion du groupe de travail morphologi e s por O pollin ique e t numé ri 44-h - c latur e , p . 8 , 6 7 Oc t obre Paris . See also Pro f . St raka ' s comment on it and Vasanthy ' s reply in Bull et in de la S oc ié t é Botani que de France 122 : 39 56

VASANTHY , G . Pollen o f t he Sout h Indian hills . Trav . Sect . Sc i . Te ch .

Inst . Fr . Pondichery , "V , 7h p . 36 planches phot o , wi th English

version (1976 ) Morphologi cal descript ion , key and phot omi crographs Cent r at h er m c our tall o f pollen o f u Vernonia conyzoides , - 7 20 , 1 , 50 51 and pl . VI I

I".

VASANTHY , G . Complexit ies o f aperture , columella and t ectum . Pro ceedings - o f IV IFC (1976 77 ) in press ; see also IV IPC Abstrac t s , 1976 . - pp . 189 190 . Aperture complexit ies in Pavet ta , Moonia and Sene ci o C olumellar t e c t al and complexit i es in Teucrium , Dipsacus , Vernonia

(13 spp . ) Cent ratherum ( 6

N 104 0 AN ETTE WIKLUND , Bo taniska Inst itut ionen , Lilla Frescat i , 5 St ockholm

5 0 , Sweden , i s undert aking a t axonomi c revisi on o f the Medit erranean group o f t um genera Ast eri scus s . l . , Pall enis and Rhan eri . RECENT LITERATURE

The se re ferenc es have b een abstrac t ed from t he Current Awareness List s o f the Library , Royal Botani c Gardens , Kew . A c omprehensive bibliography o f n Composi tae , publi shed an ually , i s cont ained in the Kew Re c ord o f Taxonomic

Lit erature .

For explanat i on o f t he S i gni fi canc e (geographical ) o f t he numbers at t he lower le ft o f each c it at ion , see Newslet t er No . 6 .

Abba, G . Mears , J . A . La di ffusione dei generi Types o f Ast erac eae at t he " " " " Ambrosia e Galinsoga Philadelphia Academy o f Natural

(Ast erac eae ) in Pi emont e e in Sci ences part 5 . Segre ' Valle d Aos t a . (The spreading gat ed types and sele ct ed aut hent ic " " o f t he genera Ambrosia and mat erial o f Ast e r eae and Inul eae . " " - os it ae 1 . 6 : 2 6 Gal insoga ( Ast eraceae ) in Com News , no . Pi edmont and in t he Aost a 1- 7

Valley ) . In form . Bot . I ta1 . ,

Morris , D . I . Miscellaneous not es on endemic

Anon . Tasmani an plant s in t he genera

Re cent lit erat ure . Olearia , Ixodia . Rec . ueen

os it ae 1 . . 6 . " . . Com News , no Vic t oria Mus (Tasmania no 55 - 12 30 Re ferences on 7 5p . (1977 ) Chrom . no . Composi tae t aken from t he

Current Awareness List , VERNONIEAE - G 1 7 Library , RE , Kew .

Rob inson , H .

Badillo , V . M . See under GENERAL .

Tre s Compuest as de Venezuela . E Act a Bot . Vene zue1 . , EUPATORIEA - 109 115 (1978 ) Illus .

Mont an oa Lour t e i ia F . n P . K . angulat a , g Bohlmann , , Maha ta , ,

A A R M . fimbriat a , Elephant opus Natu , . . , King , . and

4 hir t ifl oru s . Robinson , H . New germacranolide s from o ar Ph t ochemis t Hilli ard , 0 . M . I s c ha spe c ies . The geographi cal dis t ribu 471-h74 t ion o f Compos it ae nat i ve t o o Natal . Not es R Bot . Gard . Bohlmann , F . , Suwit a , A . , and

Edinb 5 Mabry , T . J . n Map . New labda e derivat ives and furt her c onst i tuent s o f Bri ckellia

. Ph t oc hemis t 1 (E) : Rob inson , H . speci es ry , 7 An analysis o f t he c haract ers fi and relat ionships o f t he t rib es Eupat or ieae and Vernonieae

(Ast eraceae ) . Syst . Bot . , - 199 208 (1977 Publ o 1978 )

111118 0 (Cont inued )

R M King , R . M . and Robinson , H . King , . . and Robinson , H . Guayan ia davids e i and Studi es in the Eupat or ieae H be clin ium e nt r i : 182 e g y , new ( Ast eraceae ) . A new Bi hov a h spe cie s from nort hern Sout h genus , s i . P tolo ia , a 4 - Ameri ca (Eupat or ie e 339 342 (1978; Illus .

Ast eraceae ) . Ann . Missouri - Ma z n a h Bot . Gard . , 336 370 t e b c er , N . I .

(1977 publ . 1978 ) Illus . Uma nova c ombin acao no

género Mikania Willd .

(Composit ae Bre dea , - King , R . M . and Rob inson , H . 258 259 (1978 ) En Studies in the Eupa tor ieae : 168 ( Ast eraceae ) . Addi t i ons Robinson , H .

t o t he genus A erat ina . See under GENERAL . 4 - Ph t olo ia , 3 323 355

(1978 ) Illus . , chrom . nos .

key .

Anderson , L . C .

R M . Bi el owia King , . . and Rob inson , H Studies on g u t o a Studi es in t he E pa r ie e ( Ast erac eae ) : 3 . Cyt otaxonomy

(Ast eraceae ) : 169 . Two new and biogeography . Bot spec i es o f Fl e is ch mann ia from 209- 218 ( 197 Ph Guat emala . t olo ia , Maps , chrom . nos . - 417 423 (1978 ) Illus .

Anderson , L . C . o h mnu King , R . M . and Robinson , H . New t axa in Chr s t a s ,

St udies in t he Eupat orieae se cti on Nauseosi "Ast eraceae ) . - : 1 0 . Ph t olo ia 0 20 1 8 ) ( Ast eraceae ) 7 Addi t ions y g , 3 9 3 ( 97 N o ab r i t o t he genus e c r e a . Illus . , map , chrom . nos .

J Clark , W . D . and Mabry , T . . Syst emat ic impl icat ions o f M King , R . . and Robinson , H . flavonoids in Hazardia . S Studi es in the Eupat orieae Biochem . st . Ecol . ,

: - 8 . ( Ast erac eae ) 171 . A new 19 20 (197 ) Chrom . nos

genus , Core t hamn ium . - P l : 4 C . R . h t olo ia , 39 ( ) 5 57 Cart er ,

8 : 8 . (197 ) Illus . The cyt ology o f Brachycome

The inheritance , frequency and

King , R . M . and Robinson , H . distribut ion o f B chromosomes h m m t i a Studi es in t he Eupat orieae in B . dic r o o s o a c (n

( ) : 172 . A new n i h omo oma genus , Cas t e e d a . loba . C r s , 8-60 - Ph t olo ia , 5 109 121 (1973) Illus . , map ,

(1978) 111118 0 chrom . nos .

n Ki ng , R . M . and Rob inso , H . Hobhouse , P .

Studies in t he Eupat or ieae Olearias . Garden (London ) , 22 - 2 5 1 8 (As t eraceae ) : 181 . Two new 9 3 ( 97 ) (En ) ;

spec ies o f Tri cho onia . col . illus . 4- Ph t olo ia , 39 5 33 338

197 Illus . Jones , A . G . Observat i ons on reproduct i on and phenology in some perennial

Ast ers . Amer . M idl . Nat . 184- 197 7 ( Cont inued )

Speirs , D . C . Koyama H . The genus Townsendia in On a new name o f Gnaphalium

west ern Canada . art . Bull . from Falkland Islands . Acta 16 -16 8 Al Gard . Soc . , 7

P M Semple , J . C . Mat hew , A . and Mathew , . . The cyt oge ography o f Ast er Studi es on South Indian pilosus (Composi tae ) : Ontari o Compositae : cytology o f the

and t he adj acent Uni t ed St at es . genus Blumea DC . C t olo ia 12 4 - Canad . J . Bot . , 7 365 370 (1975 5 Illus . ,

1 1 8 Fr . 279 ( 97 ) ( ) ; Illus , chrom . nos .

map , chrom . nos . V la u Namur , C . de , and er qe , R . ' . a l é tu de Stucky , J Cont ribut ion bio é du enre 1 e Hybri di zat ion between Ast er g ographique , g l hr sum

and Machaerant hera and it s Miller . Biol . Ecol . i t - t axonomic signi fi canc e . Amer . ' 17 22 (1 E map , chrom . 12 -1 1 8 - J . Bot . , 5 33 ( 97 ) 5 7 nos .

Illus . , chrom . nos .

Nogueira , I . The genus St o eb e ( Composit ae ) in t he Flora Zamb e s iaca area and

R . K . : Brummit t , Angola . BO 1 . Soc . Bro t . , 51 127 ( Report o f t he Commi t t ee for 5 133 1977 ) Illus . 4 Spermat ophyta , 20 : Proposal 03 . hania L ' H r it ier 1 8 k 9050 Re l é ( 7 9 ) Ovdienk o , 0 . A . , Litvinen o , V . I . mit (Composi tae ) vs Os e s Linnaeus and Shr e t er , A . I . - nn - i t m t ich ko l76 ) . Taxon , 285 289 Kol iche s tve o s s e a es e 2197g izuc henie Kr ymsk o -Kavkaz skikh vi dov

roda Hel ichrysum . (A quant it at ive ' Capin er i, R . , D Amat o , G . and t axonomi c study o f the Crimean

Marchi , P . Caucasian spe ci es o f the genus h h Nuclear and reproduct ive H B ull . Mosk . Ob s c .

charact ers in t he It alian t axa Is t . Pr ir . Bi ol . , 7

o f Ast eri scus Mill . ( incl . (1977 ) (En ).

Pall enis Cass . ) (Composit ae ) .

n h D k shini K . M . M . In OPTIMA Second Meet ing , Prithipal si g and a , - Firenze ( It aly ) 23 29 May 1977 . Chemotaxonomi c studies on the

Abst rac t s [1 ] p . genus Blumea DC . (Ast eraceae

Ab st rac t only . Inul eae in India . Acta Bot . Indica , 6 143- 159

Ge orgiadou , E . E . Three new speci es o f Anaphali s HELIANTHEAE (Composi t ae ) from "Fl ora Irani ca " e s é c e s area . (Troi s nouvelle s p Anas hchenko , A . V . d ' Anaphalis ( Compositae ) dans S ovr emennye voprosy pr ikladn oi ' " ki l kt s ii l aire de la Flora b o taniki , gen e t i i s e e - 1 1 8 1 8 ods oln e chnika . B ll . Vs e s . or d . Candollea , 35 3 ( 97 ) p yu

Fr . . N . I . ( i. Lenina Inst Rast Vavilova , - 69 : 47 51 Heliant hus .

Kitamura , S .

La c orre spondance du Musé um Burt on , R . M . ' nat ional d his t o ir e naturelle , connata . News , o t Paris . Act a t ot ax G e ob . , 8 ) (En ) ; Illus .

0 Recorded in Bri t ain . m Gnaphaliu . 14 . HELIANTHEAE (Cont inued )

G R R Butler , M . . and Weedon , . . Robinson , H . Chromosome analysi s o f Bidens Studi es in t he Hel ian the ae

coronata (Compositae ) . Canad . ( Ast eraceae ) : 9 . Rest orat ion o f

J . Genet . C tol . , t he genus Alloi s rmum . - - 14 14 . . Ph t ol o ia 11 412 7 9 Illus , chrom y g , 3 5

nos .

Robinson , H .

Cm me , J M . St udi es in the Hel ian t heae

C ircumscript ion and generic ( Ast eraceae ) : 10 . The relat ion

relat ionships o f Galins o a ship o f Calea sku t chii . - (Composit ae : Hel iant heae ). Ph t olo ia ; 413 414 - Madr ofio , 81 88 (1978 )

Chrom . nos . , keys .

Robinson , H .

Fay , J . J . Studi es in t he Hel ianth eae Per en ium : 11 Revi sion o f ym ( Asteraceae ) . A new genus (Ast eraceae He l ian th eae ) in

Mexico and Cent ral America . 3 - All er t on ia , 235 296

(1978) Illus . , chrom . nos . , Robinson , H .

keys . Studies in t he He l ian th eae : 12 Re - ( Ast eraceae ) . establi sh

Hart , C . R . ment o f t he genus Smallant hus . A new spec ie s o f Bidens

(Compositae ) from Mexic o . Illus . , chrom . nos . - Brit t onia , 289 292

(1978 ) Illus . , c hrom . no . Rock , B . N . Wood anat omy o f members o f the

Horner , H . T . , j r . and subtribe Es el e t iinae

C . B . ). Bu11 . . Pearson , (Compos itae IAWA , no Pollen wall and aperture de 2- 3 : 49 vel opmen t in Helianthus annuus

Tarasov , V . V . - ov n zn akhi k a adv nt vn kh Amer . J . Bo 293 309 P t or a d e y y i karan tynnykh roslyn Salvia

re flexa Horn em . ta Ambrosia URSR Je ffrey , C . tri fi da L . v . . (Repeat ed ' Helianthus annuus . Curt is s finding o f advent i t i ous and quar

1 tab . 755 ant ine Salvia re flexa

197 n ) ; Illus . , col . Horn em . and Ambrosia t ri fi da L . Ukr illus . in t he Ukrainian SSR ) . . Bot . - Zhurn o , 188 189 (1978 )

Port er , D . M . Nomenclatural changes in B L Spilanthes for the Galapagos Turner , . . 8- Axini llum Islands . Madr ofio , 5 59 Taxonomy o f h

(Ast eraceae He liant h eae ) . - Madr ofio , 46 52 (1978 )

u v t V . . P s ta oi , G . . and Illus , map , keys ' Slyu sar , E . L . ' B L Is p ol zovanie dikikh vidov Turner , . .

l kt s ii B ull . Hel iant hus v se e . y New spec ies and comb inat i ons in b kia Vse s . or d . Lenina Inst . Rast . t he genera Si e s e c and - 1 e rmum N . I . Vavilova , 69 : 11 19 ( 977 ) Tri onos (Compositae m iina Br itt onia Chrom . nos . Mela p od e ) . , - 64 68 (1978 ) Chrom . nos . An dr os hchuk A . F Ko t i . , . , s nenko , Greger , H L D ' . . and Khmel , N . V . Comparat ive phyt ochemistry and V vch nn y e ya kar iotypiv u syst emat i c s o f Anac clus . Biochem . - r e ds t avn kiv rodu . . 6 (l 11 1 p y _ Ac hillea L . Syst Ecol , 7 URSR flory . (Study o f caryo 0 types in representat ives o f Illuminat i , . and Marchi , P .

t he genus Ac hillea L . o f the Contribut i on t o the biosyst emat i c

flora o f t he Ukrainian SSR ) . knowledge o f t he genus Leucanthemum

Ukr . Zhu Bot . r n . , (Composit ae ) . In OPTIMA Second 2 -2 8 1 8 2 -2 73 7 ( 97 5 (Rus , En ) Meet ing , Firenze ( Italy ) , 3 9 May

Illus . , maps , c hrom . nos . 1977 . Abst ract s [1] p .

Abstract only .

Arr igoni , P . V . R K Le genre Sant olina en Itali e . I ssar , . .

. In OPTIMA Sec ond Meet ing , Achillea mille folium Linn ( 2 -2 Firenze Italy ) , 3 9 May 1977 . florist ic variabil ity o f morpho

Abst ract s Ll jp . logi cal charact ers . ( Achillea

Abstrac t only . mille fol ium Linn . Blumene Verfimderlichke it der morpho Bi ok K j , . , Pawlak , T . and logischen Charakt eren ) . Indian Kr n - 8 e ska B . Forest er 90 93 (197 )

Embryologi cal st udie s on Fr ) ; Illus .

Anthemis t inct oria L . (Studia embr iol o i zn Y -R g c e nad Anthemi s Ling , Y . and Ling , . . Elachanthemum t inc t oria Ac ta Soc . Bot . [ , genus novum

o - m itarum a Pol . 547 557 (1977 pub familiae Co p os ] . Act 1 8 Ph t ot ax 61-6 1 8 97 ) ( Pol ) ; Illus . , chr om . 5 ( 97 ) nos . key .

Bist e , C . Shih , c . Z t ot ax m A anio s is y on o is ch e Unt ersuchungen ( j p Shih , genus novum des Formenkrei ses Achillea familiae C ompos it arum s inens ium ) . 86-8 mille fol ium ( Ast eraceae ) in der Ac ta P t otax . Sin . , 9

. Re r t 1 8 . DDR Feddes . , ( 97 ) (Ch ) ; Illus 533-61 1 3 97 En ) ; Illus . , maps , chrom . nos . , key . Thompson , J .

Y arrow , herb o f Achille s . m 1 Bohlmann , F . and Zder o , C . Vasculu , 3 Dit erpenes wi th a new carb on skelet on from Print zia laxa . 9 SENECIONEAE

Barkley T . M .

Bohlmann , F . and Zder o , C . Three new spe c i es o f Seneci o fu New rano s es quiterpene s from (Ast eraceae ) from Mexi co . n ia 6 - 1 8 pec ie s . o c hem1 s t Br itt o , 9 75 ( 97 ) - 1159 (1 G Illus . , chrom . nos .

Bohlmann , F . , Ehlers , D . and

Bremer , K . ZdEro , C . Proposal ( 44 5 ) t o change the Einige neue Furan oer emO philan e Ph t ochemis t r type spec ies o f Pe ous ea DC . aus Seneci o Art en . , - . 56 A O 1 8 (EH) . . 7 ( 9 ) nom c ons (Composi tae ) . Taxon , 7 7 305-306 SENECIONEAE (Cont inued)

Bohlmann , F . and Le Van , N . Koyama , H . Neue Sesqui und Dit erpene Not e s on some spe cie s o f Be dfordia Ph aus sali cina . t o Chinese Cacalia : 1 . Act a - b t - 7 1173 1178 t ax Ge o o . , 65 84

Ja Illus .

Zder o Bohlmann , F . and , C . Nor dens tam , B . Neue Furanoe remophilane und Taxonomic st udies in the tribe

andere Sesquit erpene aus Sene c ioneae (Compositae ) .

Eur o s . 44 8 1 8 Vert ret ern der Gattung y p Bot . , no . . 3p . ( 97 ) - 1 7 Illus . , map .

Rei chert , H . Zdero Bohlmann , F . and , C . Senec io jac obaea L . und Senec io u n o h New f ra oer em p il an es from eruc i folius L . e rganzen de

Sout h Afri can Sene c io spe c ies . Hinweise zur Unt erscheidung der

Ph chemist 1161 bei den Art en . Gat t in er Flor . - Run db r . , 102 109 (1977

Illus . , key .

Bohlmann , F . and Zder o , C .

Hilliar din ol e s t er und andere Robinson , H . Furan oeremophilane aus Senec i o St udi es in t he Sen ec ioneae

maur ic e i . Ph t oche mis t r (Ast eraceae ) : 8 . A new spec ies - l7 (8 ) : 1333 1335 197 (En ) . o f Ps acalium from Mexi co . Ph t ol o ia 6- 8 1 8 y g , 35 35 ( 97 )

Correa , S . and Pippen , R . W . Illus . The biology o f Pi enalia

de l hin ifol ia . p (Compositae ) Robinson , H . No 131 (l ) : 27 -30 Studies in the Liabeae

(Ast eraceae ) : 12 . A new spec ies

o f Munn oz ia from Costa Ri ca . - Dick or é W . B . Adol hi . Ph t ol o ia 1 1 8 ) , and p , K y g , 33 333 ( 97

Senec io inae nidens DC . im Illus .

MTE 4908 Burschei d . G6 t t in er 4 - Flor . Run db r . , ll ( ) z 9 99 CALENDULEAE

(1977 ) Illus . K P Pandey , A . . and Singh , R . . - Douglas , G . W . and Ruyle Douglas , Development o f seed and frui t

G . in Dimor hot he c a sinuata DC . - 64 Nomenclatural changes in t he Flora (Germany ) , 57 Ast erac eae o f Brit ish Columbia : 5

1 . Se n e c ion eae . Canad . J . - 1 8 Bot . , 56 1710 1711 ( 97 ) CARDUEAE

(Fr ) . Arni ca , Luina and

Sene c i o . Brullo , S . Taxonomic and nomenclatural

Jas on ia . Gar dinal e , R . and Sc iandra , A . not es on t he genera Cass " Nuova stazi one di Senec io and Chil iadenu s Cass . (Compositae ) . " ' inae u idens g nell Italia In OPTIMA Second Meet ing , Firenze - 1 . Set t entrionale . ( A new stat i on ( Italy ) . 23 29 May 977 " " o f Sene cio inae uidens in Abstrac t s [1 ] p .

Nort hern Italy ) . In form . Bot . 1 Abstrac t only .

Ita1 . . 290

18 .

LACTUCEAE

K L Bachmann , K . and Chambers , . . Harborne , J . B . Pappus part number in annual The rare flavone is o et in as spec ies o f Microseris a yellow flower pigment in H die lla oli oc hala (Compositae , Cichoriaceae ) eywo o g ep and - Pl . 3 st . Evol . , 2 ) in ot her Cich or ieae . Ph t o - - 11 1 4 (1 8 ) . chemis t 1 1 9 3 97 Illus zz, 9 5 9 7 Ameri can spec ies only .

Kh arkevic h , and J R Barber , S . C . and Est es , . . Tsvel ev , N . N . Comparat ive pollinat ion ec ology Novye vidy oduvan chika i r i iz v rnoi o f Pyrrhopappus ge s e and (Taraxacum Wigg . ) s e e

ho u . ha t K m ha k Pyrr pgp s carolinianus . Amer c s i a c t s oi oblast i . g 6 2- 6 1 J . Bot . , 5 5 5 5 ( New spe ci es o f dandelion

Illus . , map . (Taraxacum Wigg . ) from the nort hern part o f Kamchatka di s E Bjfirndalen , J . . t ri e t ) . Bot . Zhurn . , o -8 4 8 Aconitum sept entri onale g 837 3 (197 ) Illus . Lac tuca alpina som bar skogs ar t er

i Ski en kommune , Telemark . Lack , H . W . H o diella (Aconi tum septentrionale and Die Gat tung eyw o Svent .

s Bramw Lac tuc eae . Lac tuca alpina in t he c oni ferou . ( Ast eraceae , )

H diella . forest s o f Skien , Telemark , SE (The genus eywoo Svent

Norway ) . Bl t t ia , Bramw . (Ast eraceae , - 4 8 - 125 13 (197 ) (En ) ; Illus . , Willdenowia , 329 339 (1978 )

maps . En , Fr Illus . , map , chrom .

nos .

Doll , R . Zur H W Taraxacum Flora der Lack , . .

Kasachischen SSR . Feddes Die Abbildungsvorlagen (Ski zzen , - zu Z Re ert . , 615 627 He rbarb el ege ) ahns Hierac ium

197 (En ) ; Illus . , chrom . Monographie . (The original draw Z no . ings and spe c imens used by ahn t o illust rat e his Hieracium mono

Doll , R . graph ) Willde n owia , Zur Taraxacum Flores des 439- 442 (De Herbario - Rila Geb irges . Feddes Re ert . , Be r olin en s i Not ulae no . 6 ) (En ) ; - 629 643 (1978) Illus .

(En ) ; Illus . , chrom . nos .

Ret z , B . de a Duwen s e e , H . A . Contribut ions la connai ssanc e Zu einem Fund con Lactuca de la flore hié rac iol ogique de la ' virosa im Oberharz . i n r France e t de l E s pagn e : 5 . Taxons un l Flor . R dbr . , l2 (2 ) nouveaux pour e genre Hierac ium n n a t (1978 ) Illus . dans les Pyr é é e s fra c is es e en

Espagne . Bull . Soc . Bot . France ,

Got t lieb , L . D . St e hanomer ia malheur en s is

(Composit ae ) , a new spec ies Schnaeppi , H . Madr ofio U from Oregon . , ber den Habi tus von Urospermum 44- 46 1 8 ( 97 ) Illus . , chrom . dale champii (Compositae ) . Ber .

no . Schwei z . Bot . Ges . , ' ‘ Ifl72153 11977 56611 1978 ) (Fr ) :

Illus . 5 18 5 0028 8 3799

LACTUCEAE (Cont inued ) 19 .

Schn e dl er , W .

Pflanzen , von denen in der mit t e l europais ch en Lit eratur s et t en oder gar keine Abbil dungen zu finden sind : Folge

4. 3 : Cic horium endivia L .

Singh , D . and Kaul , V . Cyt oe mbryological st udies

on Indian Tra o 0 on L . -44 C t olo ia , 1 35

1975 Illus .

Voss , E . G . and Bbhlke , M . W . The status o f c ert ain hawkweeds (Hierac ium subgenus il os e lla )

in Mi chigan . Mi ch . Bot . , 17 2 -4 35 7 (1978 ) Maps , chrom . no . ,

3 key .