PHYTOLOGIA

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. L PA ONew m and infrafarnilial in A . J . NER , co binations taxa the steraceae

d mb B B Mu fer J. R . an ln . S , ; PENCE , New genera co inations ryaceae ( ryales sci)

North America .

B WA D K flor F 1 2 R ubus R D . . a R ; eys to the of lorida , ( osaceae)

‘ L T A S fneria u f m a c B . e . URNER, new species of y (Scroph lariaceae) ro O xa a,

L( T URNE new A eratum e Eu atorieae from B . R , A species of g (Ast raceae; p ) f

B L T N Bidens melchertii A a h r . . UR ER, ( steraceae) , new species o n Jalisco ,

B L s mes f Trzxzs : Mutisieae m . T e o . URNER, A new p (Asteraceae ) fro Oaxaca , Mexico

’ his idula : i mmonii v ar. L. Stev a le B . T NE UR R, Elevation of p (Asteraceae

Botanical t B acalid moramz Aster ce e: Ta eteae - cemmentsre rdin its Anecdo es: aj ( a a g ) , ga g discovery

Botan c l Humor i a .

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PHYTOLOGIA

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m R infraf lial x . L . ami J PANE O , New co binations and ta a in the Asteraceae \ R . m in B B Mu e 1 J. SPENCE , New genera and co binations ryaceae ( ryales , s i) for Am / North erica .

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B L T Bidens melchertii a f m . . URNER, (Asteraceae) , new species ro Jalisco , Mexico 47

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' dula ; E m mi v ar. his i B L. T R Stevia lem o . U NER, levation of p (Asteraceae

Botan c l Anecdotes: Ba acalia moranii Aster ceae; Ta eteae comments re rd n its i a j ( a g ), ga i g di scovery

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NEW COM B INATIONS AND INF R AFAM ILIAL TAXA IN THE ASTER ACEAE

J ose L . P an ero

B u TX 787 1 2 Section of Integrative iology , University of Texas , A stin , USA

AB STR AC T

Molecu lar studies of Asteraceae support the recognition of tribe D oroniceae u m Asteroideae u r Aniso a in ae of s bfa ily , s bt ibes p pp , Ath roismin ae C enti edin ae Athroismeae , and p of tribe , Ch m‘mole idin ae D u esiin ae En celiin ae S ilanth in ae p , g , , and p of tribe Helianth eae D scritoth amnin ae J ae eriin ae , and y and g of tribe Mill ri a m E u h r n a r a E u h r n e e e. os e ce os os e The co binations p y , p y

dealbata E u h ros n e n evaden sis . , and p y are also proposed

KEY WO D S: Athroismeae Asteroideae Classification R , Asteraceae , , ,

Eu hros ne Heliantheae Millerieae . p y , ,

Comparative studies of D NA sequ ence data for several coding regions of the ' chloroplast D NA of the Asteraceae have revealed several lineages that requ ire names to maintain a classification that refl ects m tu u onophyletic groups . These s dies expand on the res lts of Panero and Funk (2002) and are aimed at elu cidating the major lineages of the

Asteraceae recogniz ed at the tribal level .

" L D oronicea trib u n ov : D oronicum . e . Panero , s Type

Asteroideae characteribus con uncte: u A ceteris , differt j habit herbaceo foliis alterniis etiolatis laminis inte ris ca itu lis li u latis ; , p , g ; p g ; phyllariis 2 - 3 seriatis; receptaculis epaleatis; fl osculis radiorum istillatis fl oscu lis discorum bisexualibu s corollis actinomor his lobis p ; , p , 5 ; antheris ecaudatis; ramis stylorum truncatis; cypselae teretibus 1 0 an ularibu s radiorum resentibu s absentibus g ; pappi p vel , pappi 2 Phytologia (Mar 2 005)

discorum resentibus 1 - 3 seriatis setis a orum p , pappi , p pp albis vel albo

fl avis .

z m u . Perennial rhi o ato s herbs Leaves alternate , petiolate , Ca itulescence m m . s so eti es clasping , blades ovate , elliptic or obovate p i lif rm m m an cu o . u . ter inal , solitary or open p cy es Capit la radiate

u m 2 - 3 u Invol cres he ispherical , phyllaries in series , herbaceo s to R a fl orets bifid trifid m u . so ewhat chartaceo s y fertile , or , corollas D fl orets u yellow . isc bisex al , fertile , corollas yellow , essentially glabrou s or sparsely to moderately pubescent at base . Cypselas 1 0 cylindrical to obovate with ribs , brown to greenish or blackish , u u m m essentially glabro s to p bescent, pappose , so eti es absent on ray

fl orets u 1 - 3 , papp s of capillary bristles arranged in series , white to m m m x 0 B u 3 . yellowish . ase chro oso e n ber,

M D oron icum 26 u m onotypic , contains species fo nd ostly in Asia ,

’ u Afn ca z - Femandez 2003 E rope and northern (Alvare , )

u D oron icum u m The gen s , beca se of its distinctive che istry and

mu u Arnica L. ltiseriate invol cres , has traditionally been allied to and relatives of the Madieae (Norden stam Nordenstam (1 994)

' believed D oromcum to be one of the most plesiomorphic members of the Senecioneae and consider the genu s as an outgroup in his cladistic l n l m D NA u u studies of the Ca e du eae . Co parative st dies s pport the distinctive nature of D oronicum as an independent lineage of the F r I Asteroideae affinities Calendu leae . o with to the clade this reason , D r ni a propose the recognition of tribe o o ce e .

New subtribes of the Athroismeae

- min ri A r i ma D C . Ath rois ae su b t b . n ov . : th o s Panero , Type

Athroismeae ch aracteribus con uncte: u A ceteris , differt j habit h erbaceo foliis alterniis fascicu latis ca itulis discoiformis ; vel , p ;

h llariis absentibu s l - seriatis rece taculis aleatis fl oscu lorum p y vel , p p ; eri herialum istillatis corollis c lindraceis fl oscu li discorum p p , p , y , Phytologia (Mar 2 005) 8 3

bisexu alibu s fun enter staminatis lobis 4 5 anth eris caudatis vel g , ( ; , m r i ami brev ioribu s co ess s s u s. ; cypselae p , pappi vel q

m u . Perennial herbs , shr bs or s all trees Leaves alternate or u brach blasts m m fascic late in y , petiolate or sessile , petioles so eti es with a basal spine ; blades linear, lanceolate to ovate , obovate , venation r Ca itu lescence t iplinerved or with a single vein . p a congested

m ru - m O m glo e le like solitary cy e or arranged in pen to co pact , i lif rm m m m m an cu o . u u p cy es Capit la discifor or discoid , ho oga o s or

m u r m - 2 u u 0 . heteroga o s , invol cre na rowly ca pan late , phyllaries R F m m fl orets eceptacle paleate . lorets all actino orphic , fe ale present or

2 tu u u fl orets 2 - 25 absent , to several , corollas b lar, bisex al , corollas

m u - u - 4 ca pan late , white , greenish white , p rple or yellow white , lobes ( 5 4 5 u u ; anthers ( , ecalcarate , ca date , thecae white to pale p rple , appendages sometimes with apical gland; styles divided or filiform and u filiform u m fl orets ndivided , styles associated with f nctionally sta inate , m confl uent stig atic area divided and at style branch apices , sweeping hairs present at style apex , or slightly below , rarely present below the fl att ned u . e style branch bif rcation Cypselae tangentially , ovate to oval , u u u obovate to obcordate , elliptic to s borbic lar in o tline , lateral ridges u fl oret ciliate , black, papp s of fertile cypselas either a crown of twin r m m m u t icho es , so eti es with rec rved apices or of deeply lacerated and u u m m m f sed sq a ellae , so eti es with two stronger awns at lateral ridge of u u m fl oret cypselae , papp s of f nctionally sta inate ovaries with a B u . vario sly lacerated crown or of a few , free , lacerated scales ase r m m um x 1 0 ch o oso e n ber,

The subtribe contains three genera including Athroisma (1 2

B r r L r Am 1 1 5 . eucoble ha is . lepha ispe mum Wight ex D C . ( spp ) and p (

s . m r p ) with ost species in easte n Africa, and a few species in western

M u . Africa , India , adagascar, so theast Asia , and Indonesia

- C en ti edin ae sub trib n ov . : Cen ti eda u . p Panero , Type p Lo r

Athroismeae characteribu s con uncte: tu A ceteris , differt j habi h erbaceo foliis alterniis ca itulis discoiformis radiatis h llariis ; , p vel ; p y

1 - 2 seriatis rece tacu lis e aleatis fl oscu lorum eri herialum , p p ; p p 4 Phytologia (Mar 2 005) 8 7(1)

istillatis corollis z omor his c lindraceis li u lis brev ioribus lobis p , yg p , y , g ,

2 - 3 absentibu s fl osculi discorum bisexualibu s corollis vel ; , actinomor his lobis 4 anth eris caudatis brev ioribu s m st lorum p , ; , ; ra is y i r i ri ubt r ti a nti subtruncat s b ev o bu s s e e bu s bse bus. , ; cypselae , pappi

u . Ann al or perennial herbs Leaves alternate , sessile , blades i l n u . Ca tu esce ce obovate , vario sly toothed , rarely entire p sessile to m u u . u shortly ped nc late , axillary , rarely ter inal , solitary Capit la m m m . u u discifor or radiate Invol cres ca pan late to he ispherical ,

1 - 2 u R u u . phyllaries in series , s beq al , herbaceo s eceptacles shallowly l Peri h eral/radiate fl orets e a eate . convex , p p in several series , pistillate , m m m u m fertile , so eti es lobes extre ely red ced and corolla see ingly m u D tu u . b lar, corollas crea y white , green , light yellow or p rplish isc fl orets u m u m bisex al , fertile , corollas tetra ero s , crea y white , green , light u e u u m yellow or p rplish , ssentially glabro s with a few gland lar tricho es , u z m 4 lobes of corollas not vasc lari ed; sta ens , anthers hyaline , tailed , m u t ecalcarate , appendages wanting or if present , in te , oval; s yles with u t m u two vasc lar strands , s yle branches with divided stig atic s rfaces ,

u u u . concave , rceolate in o tline , apices ac te , papillose Cypselas u s bterete , deeply ridged , glabrescent on basal end , cells of distal end u m with raised apical tips , event ally tips elongating and for ing biseriate m m t m tricho es , tricho es increasing in densi y at apical end for ing a cu u tu m shallow p aro nd corolla be , tricho e cells shallowly ridged; u papp s absent .

M t Cen ti eda fiv e u u ono ypic , p contains species fo nd in A stralia , m u r u u . so theast Asia, Af ica and so thern So th A erica

The genu s Centipeda has been placed in several tribes including Anthemideae and Astereae but Bremer in his excellent account m u difficult but m of the fa ily , considered the gen s to place aintained it Asteroideae u m 1 994 u in , nassigned to a tribe . Neso ( ) believed the gen s m m Astereae m tu to be a e ber of the . Co parative s dies of the ITS region of the nu clear ribosomal D NA by Wagstaff and Breitwieser (2002) u u m A t r a fo nd strong s pport for the exclu sion of the genu s fro the s e e e . Their studies revealed Centipeda to be an isolated lineage sister to Phytologia (Mar 2 005) 8 5

Athr i mea H lianth a M o s e and the e e e alliance . ore recent studies (Panero A et al . . u u Centi eda thr i m a o s e e . , in prep ) s pport the incl sion of p in the

An iso a in ae su b trib n v - A iso a o . : n us . p pp Panero , Type p pp Hook Am .

Athroismeae characteribu s con uncte: tu A ceteris , differt j habi herbaceo; foliis altemiis; capitu lis radiates plerumqu e discoideis; phyllariis 1 - 2 seriatis; receptacu lis paleatis plerumque epaleatis; fl osculi radiorum istillatis fl oscu li discorum bisexu alibu s p ; , corollas actinomorhis lerum ue z omor his lobis 5 anth eris caudatis p q yg p , ; , brev ioribu s s bteretibu s ab sentibu s mu s ami u u s. ; cypselae , pappi vel lti q

u r Ann al or perennial herbs . Leaves alte nate , petiolate to u m m u m s bsessile , blades linear to ovate so eti es s bcordate , si ple to innatifid m m Ca itulescences m p , triplinerved , so eti es pinnate . p ter inal , ani lif rm m bumbillif rm m cu o su o . solitary , open p cy es , rarely cy es tu u m u m Capi la radiate , rarely discoid . Invol cres ca pan late to ostly m R he ispherical . eceptacles convex , rarely shallowly conical , paleate , l at R a fl r e a e e . o ets m m m rarely p y pistillate , so eti es with sta inodes ,

D fl r ts u - . o e 5 fertile , corollas yellow isc bisex al , fertile , corollas lobed ,

- m 5 golden yellow; sta ens , anthers yellowish to brown , shallowly t calcarate or ecalcarate , tailed , appendages ovate ; s yle branches with m u tu . u divided stig atic s rfaces , with ob se papillae Cypselae s bterete , rm u u m edges wanting , epide is witho t crystals , papp s of s all scales or absent .

M Aniso a us 40 u m onotypic , p pp contains species fo nd ostly in M u Africa and adagascar , with one r deral species extending into u so theast Asia and China .

New subtribes of the Heliantheae

h r - hromole is C omole idin ae su b trib u s n ov . : C p Panero , Type p B enth . 6 Phytologia (Ma r 2 005) 8

Helianth eae ch aracteribu s con uncte: tu A ceteris , differt j habi herbaceo u , pal stris f lii altemiis etiolatis laminis inte ris lob atis/run ina i o s , , c t s; p I g vel ca itulis li ulatis h llariis 2 - 3 seriatis dimor his rece taculis p g ; p y , p ; p aleatis fl osculis radiorum istillatis fl osculis discorum bisexualib us p ; p ; , corollis actinomor his lobis 5 anth eris ecau datis m st lorum p , ; ; ra is y sublinearis deltatis radiorum com ressis discorum , ; cypselae p ; cypselae udratis radiorum absentibu s discorum coronniformibu s q ; pappi , pappi ,

brev iorum.

R - m u . osette like , perennial , se iaq atic herbs Leaves alternate

m - u u petiolate , se i s cc lent, ovate to narrowly lanceolate , entire or inn ifi a i l n at d. C tu esce ces u p p axillary , solitary . Capit la radiate .

u m r m r 2 - 3 u u Invol cres ca panulate , phylla ies di o phic in series , s beq al , m m h m R u . e branaceo s wit black arkings eceptacles convex , paleate . R a fl r t D fl oret u o e s . s y pistillate , fertile , corollas white isc bisex al , m u m 5 fertile , corollas penta ero s , yellow; sta ens , anthers yellow m m n a hyaline so eti es with dark con ectives , ppendages papillose ; styles u m u with two vasc lar strands , style branch with divided stig atic s rfaces , u lt t R a apices broadly ac te to shallowly de a e . y cypselas tangentially fl ttened u u a u u . , triq etro s to weakly q adrate in cross section , glabro s D m u u isc cypselas obpyra idal , q adrate , sparsely p bescent , u u m u m u conspic o sly s aller than ray cypselas , papp s a crown of in te u m awns and sq a ellae , absent in ray cypselas .

M t Chromole is u ono ypic , p contains one species fo nd in vernal mid M pools in elevation grasslands of central and western exico .

The Chromolepidinae occupy a derived position within the Helianth eae D u esiinae Zaluz aniinae , sister to g and collectively sister to t al R 1 8 1 hr m le s (Panero e . obinson ( 9 ) placed C o o pi in the Zaluz aniinae because of its fertile ray fl orets and shallowly quadrartic u un m M nfl r cypselas . The gen s is usual a ong exican su owe s becau se of

u - u u its aq atic , rosette like habit, and conspic o s blackish colorations on r m phyllaries and abaxial side of ray corollas . The heteroch o atic capitula and dimorphic involu cre of Chromolepis are characteristics not Zaluz aniina seen elsewhere in e . Phytologia (Mar 2005) 7

tri n - D v ia A . G D u esu n ae su b b u s o . : u es g Panero , Type g ray

Helianth eae ch aracteribus con u ncte: u A ceteris , differt j habit h erbaceo foliis altemiis etiolatis laminis runcinatis lobatis ca itulis ; , p , , ; p

ligu latis; phyllariis 2 - 3 seriatis; receptacu lis paleatis; fl oscu lis radiorum istillatis fl osculis discorum fun enter staminatis corollis p ; g , actinomor his lobis 5 anth eris ecaudatis m st lorum sublinearis p , ; ; ra is y ; com ressis radiorum absentibu s lerum ue cypselae p ; pappi , p q i i r i m coron form bu s b ev oru .

r u n Prost ate , stolonifero s peren ial herbs . Leaves alternate , innatifi u u d. petiolate , blades obovate to oval in o tline , r ncinate to p i l n m ani lif rm m Ca tu esce ces cu o . p axillary , solitary , or s all p cy es

2 - u . u m 3 Capit la radiate Invol cres he ispherical , phyllaries in series , m R fl at u u u u . s beq al , o ter ost foliaceo s eceptacles to slightly convex , R a fl orets two m paleate . y pistillate , in series , fertile , corollas le on to

- golden yellow with greenish to black veins on abaxial side , corolla

- D fl oret u m 2 3 . s apices deeply rarely lobed isc f nctionally sta inate , m u m 5 corollas penta ero s , yellow; sta ens , anthers black, appendages oval to trullate with a visible constriction between appendage and fl orets thecae , shallowly carinate ; style branches of disc narrowly t fl oret tapered and papillose , s yle branches or ray spreading with broad , m R a fl attened u . divided stig atic s rfaces y cypselas tangentially , u m biconvex , essentially glabro s with a few tricho es on the neck , m u m u argins with shallowly lacerate wings , papp s absent or of a in te

crown .

M t D u esia u tu ono ypic , g contains one species fo nd in dis rbed ,

- eastem M . gravelly or sandy soils , in the dry highlands of central exico

The D ugesiinae are sister to P arthenium with which they share bifid fl attened strongly ray corollas , tangentially ray cypselas , and M m functionally staminate disc fl orets. ore extensive sa pling of the Helianth eae in the future may reveal subtribe Ambrosiinae to be G u ara h i r Parthenice A . p p ylet c and that P a thenium L. and ray sho ld be D included in the ugesiinae . 8 Phytologia (Mar 2 005) 8 7(1)

tri n - E lia . En cell mae su b b u s ov . : nce Panero , Type Adans

Heliantheae ch aracteribu s con uncte: u A ceteris , differt j habit herbaceo u foliis altemiis etiolatis laminis inte ris vel fr ticoso , , p , g plerumque lobatis/dissectis; capitulis ligulatis plerumque discoideis;

h llariis 2 - 5 seriatis rece tacu lis aleatis deciduis p y ; p p , paleae ; fl oscu lis radiorum plerumqu e pistillatis; fl oscu lis discorum bisexu alibu s corollis actinomor his lobis 5 antheris ecaudatis , p , ; ; ramis stylorum sublinearis; cypselae compressis; pappi absentibu s

1 - 2 ami vel squ s.

u u . Ann al or perennial herbs , shr bs , rarely trees Leaves alternate , m m u rarely opposite , petiolate , rarely sessile , so eti es with resino s " : u u m m ex dates , blades linear to ovate or tr llate , so eti es laciniate , n Ca itule cenc enta erv ed. s es pinnately veined or triplinerved , rarely p p m aniculiform m m ter inal , solitary and scapose , or in p or cory bifor m m u . u ur u cy es . Capit la radiate or discoid Invol cres t binate , ca pan late

m 2 - 5 u u u or he ispherical , phyllaries in series , s beq al , rarely grad ated ,

u u m m W u u . herbaceo s , rarely chartaceo s so eti es ith resino s ex dates R a fl r t R fl at u u . o e s u eceptacles to convex , pales decid o s y ne ter or

- m m rarely pistillate and sterile , corollas golden yellow so eti es with m u m D fl orets u biseriate tricho es on t be , rarely on li b . isc bisex al ,

5 - - u m u fibers fertile , corollas lobed , golden yellow or p rple , ostly witho t m u m m e bedding the vasc lar strands , lobes so eti es with thickened cells or with glandu lar or mu lticelu lar trichomes on abaxial surfaces of the m 5 lobes ; sta ens , anthers yellow , brown or black, appendages ovate to m m u m um linear, so eti es with gland lar tricho es , endotheci cells

u rm u 1 - 3 f sifo , rarely q adrate with polar bridges ; style branches with u m u m m u f sed stig atic s rfaces , so e species with two stig atic s rfaces that u ur f se slightly above style bif cation point, densely papillose below ,

u u . fl attened apices ac te to broadly ac te Cypselae radially , rarely u thickened or terete , obovate to oblong , densely sericeo s , rarely u m m u u W glabro s , so eti es with conspic o s ings or corky edges on sides u u and neck, papp s of two slender awns (rarely absent) with or witho t u m m u sq a ellae in between the , rarely of awns f sed to a crown u u s rro nding the neck . Phytologia (Mar 2 005) 8 9

The subtribe contains fiv e genera including Encelia (1 5 E A A 4 G . F lour n A D ncelio sis . ia s C . 3 3 p ( ray) Nelson ( e . ( r A G 2 H l a t e l A Ge aea . h i n l a . . G . e 8 Torr ray ( Torr ray ( spp . ) u m u m fo nd ostly in western North and So th A erica .

The Enceliinae are an isolated lineage of the Heliantheae nestled En elmanniinae Ambrosiinae between the g and the clade containing the , Chromole idinae D u esiinae Helianthinae S ilanthinae p , g , , p , Zal z aniina Zinniina t al u e e e . u , and (Panero The s btribe shares H li nthinae fl r u e a o ets. with s btribe sterile ray In addition , the strongly fl attened u u u and sericeo s cypselas , along with the cad co s pales of the Enceliinae are only seen elsewhere in the Helianth eae in a few m m H lianthin a r F B e e e S ncretoca us S. . e bers of the y p lake , and m K so e species of Viguiem unth) .

S ilan th in ae tri n - b b u s ov . la he su : S i nt s . p Panero , Type p Jacq

Heliantheae ch aracteribu s con un cte: tu A ceteris , differt j habi h erbaceo u foliis o ositiis etiolatis lerum u e vel fr ticoso ; pp , p p q sessilis , lamin is integris plerumqu e lobatis; capitulis ligulatis prlerumque discoideis; phyllariis 1 - 5 seriatis; receptaculis paleatis plerumque e aleatis conoideis fl oscu lis radiorum istillatis fl oscu lis p , ; p ; bisexualibu s corollis actinomor his lobis 5 lerum u e 4 antheris , p , p q ; ecaudatis; ramis stylorum sublinearis; cypselae compressis; pappi

r ntibu s 1 - 3 ami ese s u s. p , q

um u n m m Erect or dec bent ann al or peren ial herbs , so eti es rooting

u . at the nodes , rarely scandent shr bs Leaves opposite , petiolate or u m m m s bsessile , blades linear to ovate , so eti es renifor , entire , Ca itule n m m . sce ces triplinerved p axillary or ter inal , solitary, si ple to m m l un m fistu ose . u congested cy es , ped cles so eti es Capit la discoid or

m m 1 - 5 . u u radiate Invol cres ca pan late to he ispherical , phyllaries in u u m m u m m series , s beq al , rarely di orphic , ostly herbaceo s , so eti es bases ur u R ind ate and coriaceo s . eceptacles convex to conical , especially m e aleate u with age , ostly paleate , rarely p , pales chartaceo s , rarely R a fl r t u u . o e s coriaceo s y pistillate , fertile , corollas p rplish , yellow D fl r t m u . o e s u orange or white isc , bisex al , fertile , corollas penta ero s , 1 0 Phytologia (Mar 2 005)

m u u m 5 4 rarely tetra ero s , p rplish , white , or yellow; sta ens , rarely , deltate u anthers brown or black, appendages ovate or with or witho t t u m u u glands ; s yle branches with f sed stig atic s rfaces , apices ac te and R a fl att n u . e ed papillose , witho t appendages y cypselae tangentially , r u u m t iq etro s , obovoid , sparsely to densely ciliate , tricho es with D fl attened m m u . rec rved tips with age isc cypselae radially , so eti es

' tn u etrou s m m u peripheral ones q , terete , so eti es q adrate and narrowly m u u m rho bic , rarely sq are in o tline , ostly shallowly winged or m m u u k u so eti es conspic o sly winged , cor y , ciliate , glabro s to sparsely u u m u u p bescent, papp s of a in te crown , a single awn f sed to a broad

u m mm 2 - 3 ring aro nd the neck of the cypsela , or ore co only of slender u m m u m awns as a contin ation of the wings , so eti es with sq a ellae in t be ween .

fi A R The subtribe contains v e genera including cmella ich . ex Pers . r F B 1 Salmea D C . 1 0 . (3 0 Oxyca pha S. . lake ( ( spp )

2 - 4 M Spilan thes (6 and Tetranthus SW . ( ost species of the u u W but s btribe are fo nd in the New orld tropics , a few species of Acmella and Spilanthes are endemic to the tropical and subtropical

regions of the Old World .

The Spilanthinae are sister to the Zinn iinae (Panero et al 200 1 ) and m share with the opposite leaves , the tendency for solitary, long a i l ilanthinae u u c tu escences . S ped nc late p , and conical receptacles The p m Zinniinae m differ fro the in lacking arcescent ray corollas .

New subtribes of tribe Millerieae

D itoth amnin n scr ae su b trib u s ov . y Panero , : D scrit t nu B L R ob . Type y o ham s . .

Millerieae ch aracteribu s con uncte: tu A ceteris , differt j habi h erbaceo u foliis o ositiis lerum ue altemiis etiolatis vel fr ticoso pp p q , p laminis inte ris lerum ue lobatis/dissectis ca itulis vel sessilis , g p q ; p

ligulatis plerumque discoideis; phyllariis 1 - 4 - seriatis; receptacu lis paleatis plerumque epaleatis; fl osculis radiorum pistillatis; fl oscu lis discorum bisexualibu s corollis actinomor his lobis 5 anth eris , p , ; Phytologia (Mar 2005) 8 1 1

ecaudatis; ramis stylorum sublinearis; cypselae subteretibu s; pappi

resentibu s setis a orum lumosis . p , p pp p

u u . m Ann al or perennial herbs or shr bs Leaves ostly opposite , m m m m so eti es alternate , petiolate or sessile , so eti es clasping or m m perfoliate , blades linear to broadly ovate , so eti es trilobed to deeply deltate u u . lobed , rarely or acic lar, triplinerved , rarely ninerved Ca itulescences m anicu liform p ter inal , solitary and scapose or open p m m . u . cy es , rarely congested cy es Capit la radiate , rarely discoid

u m u m 1 - 4 Invol cres cylindrical , ca pan late or he ispherical , phyllaries in u u m m r u m n series , s beq al , so eti es g ad ated and di orphic with in er series u mu u u chartaceo s and ch longer than the o ter herbaceo s series , m m u 4 fl esh u so eti es red ced to only y, herbaceo s phyllaries enclosing

fl oret m - R s r bud. in a py a id like , valvate eceptacles convex to conical , leate m m R a e a u . paleate , rarely p , pales so eti es chartaceo s , hyaline y

fl r t - nk m D o e s . fertile , corollas golden yellow , white , pi or agenta isc

fl orets u 5 - - bisex al , fertile , corollas lobed , golden yellow , green , white , u m 5 pink or p rplish; sta ens , anthers yellow or brown , appendages m m ovate ; style branches with divided stig atic surfaces . Cypselae ostly fl attened u u terete to shallowly radially or q adrate , glabro s to densely u u mu u sericeo s , papp s of ltiple bristles or scales , rarely a very red ced crown .

The subtribe contains fiv e genera including Bebbia Greene (2 m h r B L R D r t hamnus 2 ob . 5 sc i ot Cy op o a . . ( y ( T etra onotheca L 4 Tridax L. 3 0 M g . ( and ( ost species are u m m distrib ted in ontane , tropical North A erica with a few species in An m the north and central des of South A erica .

The D yscritothamn inae are sister to the Melampodnnae (Panero et al m D scritothamn inae u . The onophyly of the y and their s btribal relationships are well supported .

J ae eru n r n " Jae eria Ku ae su b t ib u s ov . : g Panero , Type g nth

Millerieae ch aracteribu s con uncte: tu A ceteris , differt j habi herbaceo u u u foliis o ositiis etiolatis , terrestris , pal stris vel aq atic s ; pp , p 1 2 Phytologia (Mar 2 005) 8

laminis inte ris ca itulis li u latis h llariis l seriatis vel sessilis , g ; p g ; p y receptacu lis paleatis; fl oscu lis radiorum pistillatis; fl oscu lis discorum bisexualibu s corollis actinomor his lobis 5 anth eris ecaudatis m , p , ; ; ra is stylorum sublinearis; cypselae teretibu s; pappi absentibu s plerumque r n if rmi br v i rum co o o bu s e o .

u . Ann al or perennial herbs , rooting at the nodes Leaves opposite , Ca itulescen ces sessile to petiolate , blades linear to ovate . p axillary or m anicu liform m u ter inal , solitary or of open p cy es . Capit la radiate . u m u m 1 Invol cres ca pan late to he ispherical , phyllaries in series , u u u W u s beq al , herbaceo s with hyaline , ciliate ings wrapping aro nd the u u u ray cypselae , cad co s and shed along with cypsela as a nit i eri n a . R R a (p gy ) eceptacles strongly convex to conical , paleate . y

' fl orets - - - u pistillate , fertile , corollas white , white pink , white p rple , m D fl r t tu . o e s u yellow , arcescent , be short or absent isc bisex al , fertile ,

- m m 5 corollas yellow or yellow green , so eti es whitish gray , lobes rarely

' 4 u u th e throat m 5 , t be abr ptly narrowed into ; sta ens , anthers yellow , t u un u . appendages ovate; s yle branches rec rved , apices ro d to obt se u u Cypselae terete to obsc rely angled , clavate , bases tapered , glabro s , shiny black , ray cypselas slightly larger than disc cypselas otherwise u u m u B r m m eq ivalent; papp s absent or a in te crown . ase ch o oso e um x 9 n ber,

M t Jae eria u ono ypic , g contains nine species fo nd in the New World tropics with most species concentrated in the highlands of M exico .

Jae eriinae th e Milleriinae The g are sister to clade containing , D esmanth diinae Es el tiina alin in t al o e e G so ae e . , p , and g (Panero

m u Am r iina New Co binations in s btribe b os e .

tu Helianth eae u Am r iinae M t al 1 995 S dies of s btribe b os by iao e . ( ) u E r D u fiv e s pport the expansion of the genu s uph osyne C . to incl de E r species . This new concept of uph osyne requ ires the creation of the following combinations :

1 4 Phytologia (Mar 2 005)

m B 1 4 al n l a In B C e du e e . : m K rden sta . 99 . ed No , Tribe re er . ( ) classification m Asteraceae cladistics and , Ti ber Press . Portland ,

- 3 65 3 76 . Oregon , pp.

P n ero J . L . B . G. B aldwin E . E . ch illin J . A . le in r a , , , S g, and C v ge . 200 1 M u tu m m . olec lar phylogenetic s dies of e bers of tribes H l ni a H lianth a Eu at ri a e e e e e e e o e e 3 . G , , and p (Asteraceae) eneral systematics and proposed taxonomic changes in cu rrent la ifi ati n t AIB S 200 1 m u u c ss c o . u Abs racts of the eeting , Alb q erq e ,

M . 1 32 . New exico , p

P n ro L. V A . F u nk . 2002 a e , J . and . . Toward a phylogenetic subfamilial classification m for the Co positae (Asteraceae) . Proc .

B a 1 1 0 - 22 W s 5 : 9 9 9 . iol . Soc . hington

R ob in son H. 1 1 u r m , 98 . A revision of the tribal and s bt ibal li its of the

- m B t . 1 1 1 2 Heliantheae . 0 o 5 : . (Asteraceae) S ithsonian Contr .

t ff I B r W a s a S. J . . eitwieser. 2002 . g , and Phylogenetic relationships

m u . Pl . of New Zealand Asteraceae inferred fro ITS seq ences Syst .

E l 2 1 2 - 224 v o . 3 : 03 . Phytologia (Mar 2005) 8 7(1) 1 5

NEW GENER A AND C OM B INATIONS IN BR YACEAE BR M F OR R H M R ( YALES , USC I) NO T A E I CA

en ce J oh n R . Sp

G R P O . National Park Service , len Canyon National ecreation Area ,

B x 1 AZ 4 - 1 m o 507 860 0 507 US A . : , Page , , E ail

John_Spence@nps. gov

AB STR ACT

P h stomum H rn h u The genu s tyc o o sc . is res rrected for species of Br um Ambl o h llum Caes itibr um 32 y in sections y p y and p y , with well m established species found in North A erica transferred . The section Le tostomo sis Brach menium k p p of y is raised to generic ran , with the m u one species found in North A erican transferred . The gen s Plagiobryoides is newly described for the neotropical Bryum Mielichho er a m incrassa tolimba tum. i Two species of f in North A erica ,

M. macrocar a M. tehamensis p and , are transferred to the neotropical H genu s aplodontium.

K e words: m m B Ha lodontium y osses , North A erica , ryaceae , p , Pla iobr oides Pt chostomum g y , y

I ntrodu ction

A considerable amount of new research has been published B u u Br um recently on the ryaceae , and in partic lar the gen s y (Cox H d r n 1 2 2 2002 e de so 998 003 et al. 000 , ; Cox ; Pedersen ; Pedersen Hedena 2 02 2 1 87 1 996 s 0 ai al. 003 9 ; Pedersen ; Spence , ; Spence R m 1 996 1 999 Br um m a say , , y is an extre ely large and diverse u u m m t u gen s , nited pri arily by plesio orphic sporophy ic characters fo nd

- f et throughout the Bryalean alternate diplolepidou s taxa (o . Pedersen m al. Although there is considerable orphological variation u t m m Br um within the gen s , no at e pts have been ade to divide y into m u um m tu u . s aller and pres ably ore na ral gro ps To date , the vario s phylogenetic studies u sing chloroplast D NA sequ ences and 1 6 Phytologia (Mar 2005) 8

m u m u orphology have not converged on a partic lar taxono ic sol tion , other than indicating that Bryum and the related genu s Brachymenium

are polyphyletic .

Although trying to understand the relationships among variou s B m r um s . l. m proposed genera , species and sections of y see s at ti es to I define tu u be hopeless , believe that it is possible to na ral gro ps within m the genu s based on a combination of characters fro both generations . D r R m 1 m Br um F . Helen a say and , while onographing y for the lora of Au stralia (Spence R amsay have identified groups among the We u m gametophytic diversity in the genu s . have foc sed pri arily on m t u characters of the ga etophy e generation , as all work to date s ggests u m tu u that the caps le orientation and peristo e fea res , traditionally sed to m in m u deli it genera the fa ily , are not indicative of evol tionary We relationships above the species or perhaps sectional levels . feel that restricting analysis to primarily gametophyte characters provides a valu able and relatively stable basis for the delimitation of natural u B c 1 993 u gro ps (genera) in the ryaceae ( f. Zander for an analogo s m our situation in the Pottiaceae) . There are several i plications of work

u D NA u . and other st dies based on characters , which are disc ssed below

Our work clearly supports the presence of W m major lineages B m i um u . within y , in addition to several s aller gro ps Interestingly , these two major groups receive at least partial support from the other

D NA m c et al. and orphological studies cited above ( f. Pedersen One lineage inclu des the many traditional Bry um species first grouped “ ” as the informal Cernu ibryum by one of u s (Spence along with Acidodontium Brach menium Brach menium R hodobr um , y section y , y R r u um u m and osulab yum. The second lineage incl des the n ero s s all mm u Br um Br um A alodict on ge ifero s species of y in sections y , p y , Al ini ormia D oliolidium Anomobr um p f , and , along with y and Brach m m D r r u u ic anob um. We out y eni sect . y have pointed that tr e Bryum Bry um argenteum) is closely related morphologically to Anomobr um u accomodated u y , and co ld easily be in that gen s (Spence R m G B lecto ified u E. t a say However, . ritton yp the gen s Br um r m u u B. a enteu y with g , which we regard as an nfort nate choice R m Anomobr um Br um (Spence a say jy , and the type of y are Phytologia (Mar 2 005) 8 1 7

su fficiently distinct from other members of the genu s to warrant two m m generic status . There are potential taxono ic i plications of this fir t B l t t ifi ation m . ec o yp c if revisions are atte pted The s is that . argenteum and its allies cou ld be treated as a genu s closely related to B Anomobr um r um m . y , with all other y species given a new na e The m m second is that the two taxa ight be co bined , in which case a new Br For name wou ld still have to be found for all other yum species . u t R m We A stralia , we chose the lat er of the two (Spence a say u m Br um t also arg ed for the conservation of the na e y with a new ype , fir t B es iti um s . ca ci for which we selected the species in Hedwig , p R m B t mm (Spence a say However, the ryophy e Co ittee on Nomenclature rejected our proposal (Zijl stra based in part on a use Br um caes iticium concern for the of y p as the conserved type , and Br m also becau se further studies on yum were pending . So e of these m l presu ed studies have now been either published Pedersen et a . 2003) or are cited in Pedersen and as noted above they do not u Br um agree with each . This leaves two choices , contin e to treat y in a u very broad sense , knowing f ll well that it is polyphyletic , as Ochi 1 992 m u classification u ( ) chose to do , or co e p with a new of the gen s m tu based on the evidence to date fro all s dies , both genetic and m r first u o phological . The approach is nacceptable , so this paper as well as the treatment in Spence R amsay (2005) represents the first attempt since Fleischer and Broth erus (1 925) to generically re- classify Bry um

th u 1 9 u u Perhaps not s rprisingly , the cent ry also witnessed conf sion m Br um 1 80 1 with the exact deli itation and nature of y . Hedwig ( ) u t u m incl ded a varie y of species in his concept of the gen s , any of F Homschuch which now reside elsewhere . airly soon thereafter both (1 822) and Bridel (1 826) described new genera long since ’ m z m h h u Br um. Ho sc uc s synony i ed nder y In case , it was P t chostomum Br um y , described for several species now residing in y , B u l ovicu B . uli nosum. B including . a g m and gi ridel p blished = Cladodium B rr B inclinatum B . ma atii . for the species . and ( M u amblyodon) . ost of the other species they incl ded in these new

Pohlia Mnium h . genera have since been transferred to , and ot er genera 1 8 Phytologia (Mar 2 005) 8

Validly P u b lish ed Name C u rrent Name Bri Br um calo h llum Br. d. calo h llum R . C. p y ( ) y p y Nees Pla iobr um demissum demissum Hook . C. ( ) g y * B ambl odon Brid. r um inclinatum SW . ex C. ( ) y y

rham hoste ium Ham e Acidodontium rham hoste ium C. p g p p g socorrense Ham e Br um limbatum C. p y Bri Br um uli i nosum uli inosum d. C. g y g

1 umm u Cladodium B Table . S ary of validly p blished species of ridel , m u u . year of p blication , and c rrently accepted na e The lectotype species is indicated by an asterisk .

F or m m u so e reason , these generic na es never ca ght on with later u u Br um bryologists , and have lang ished nder y ever since , probably B iticium u first Br um . caes beca se the species of y in Hedwig p ,

mbl odon . al ovicum B . a is morphologically similar to both B . g and y All those species still remaining in Bryum that were placed by these two in Cladodium and P lychos tomum are in the informal group “ ” ” ar enteum u B um Cemu ibryum while B . g is in the gro p ry of Spence (1 987)

B u I re- eca se of the above considerations , propose here to instate the genu s P tychostomum Homschu ch for those species of Bryum in the ” informal group Cernu ibryum of Spence Since all three first B Homschu ch species cited by both ridel and are related , either u u u but P l chostomum t Cladodium gen s co ld be sed , y has priori y over m u u . by fo r years Interestingly , y work also s ggests two closely related B u P t chostomum . gro ps within y , one corresponding to the species B um li inos . al ovicum . u g , and the second to those species related to g For I two u u u u now prefer to keep these gro ps together, altho gh f t re work may indicate that they represent distinct but closely related

u Hedderson 200 1 et al. genera . The st dies of Cox and ( ) and Pedersen (2003) both show a well supported clade that consists of species in

P t chostomum u u Br um donianum y , altho gh incl ding the species y , which r based on its gamteophyte may be a R osulab yum. Phytologia (Mar 2005) 1 9

Validly P u b lish ed Name C u rren t Name

’ um Bri Br um al ovicum caes itzci d. P . p y g * H m ch . Br um uli in su cernuum Hedw. o s o m P . ( ) y g Br m al ovicum com actum Homsch . u P . p y g Br m a l ovicum endulum Homsch . u P . p y g r B um a l vi P uchellum R . B . r o cum . p y g Br um uli inosum radiculosum Brid. P . y g

2 umm u P t ch ostomum Table . S ary of validly p blished species of y m Homsch . u u . , year of p blication , and c rrently accepted na e The lectotype species is indicated by an asterisk .

Tables 1 and 2 list all validly published species in Cladodium and P t mum 1 873 lectot ified Cladodium tychos o . Pfeiffer ( ) yp with the in linatum but P chostomum C. c t species , y has not yet been ’ l t t ifi Homschuch s u u ec o yp ed. nderstanding of the gen s was based on m t m . certain characters of the sporophy e , pri arily the peristo e His “ essential characters inclu ded P eristomium duplex: exterius dentibus sedecim erectis interius membrana h alia licata eristomio externo , y p p adhaerens et denta ejusdem inter se conjungens The adherence of the endostome to the exostome is a critical character in the Homschuch u circumscription of the genu s . The two species incl ded in

P tychostomum are cu rrently kn own as Bry um algovicum and B . B z u m uliginosum. oth these species are characteri ed by red ced peristo es P m m . with the endosto e adherent to the exosto e . I have selected = P =B dulum . cernu m B uli inosum u . en u ( . g ) as the lectotype beca se p ( m inclinatu . al ovicum t Cladodium C. g ) is closely related to the ype of , This selection was made in the interests of preserving both generic

m Cladodium z na es , in case is also recogni ed at the generic level at m P t chostomum cernuum so e point in the future . y is in a different

inclina tum it . section than C. and is not closely related to

2 P t m h 2 2 1 8 2 . chostomum Ho sc . F 5 : . 6 y , lora , syll , Lectotype 4 22 2 6 1 8 . F 5 : . : P t chostomum cernuum Homsch . species y , lora , syll , Homschu ch u u In addition , incl ded the species c rrently known as Br P t chostomum um al ovicum B B S. G. u y g ( rid . ) . in the gen s as y

dulum Hom h F 2 62 1 822 . en sc . 5 : p , lora , syll , 20 Phytologia (Mar 2005) 8

The following 27 species found in North America are also transferred to the genu s .

m n R b . ov . P t chostomum archan elicum . . co y g J Spence , r r um Br ol E r 4 1 5 B m: B um a chan elic . u . : 3 asiony y g y , 1 4 8 6 .

R m n Br. I . . co b . ov . P t chostomum arcticum R . y ( ) Spence ,

P r R Br. M l ill . 3 8 . 1 823 B m: ohlia a ctica . . e v . asiony , Chlor

B m n R . co b . ov . P t ch ostomum badium . . y ( rid ) J Spence ,

B m B a um B . B l 1 di r . : r um caes iticium . b o . : asiony y p var rid , y Univ 1 2 850 . 8 7 .

P t R m n chostomum bimum r . . . co b . ov . y (Sch eb ) J Spence ,

’

B m M r B t R l . : mum bimam . o . . u : 79 asiony Sch eb , Zeit ( egensb rg) 1 802 .

mb n ov P t chost mum calo m R B r R co . o h llu . . . . y p y ( ) J Spence , 2 B m: Br um calo h llum R Br Melv ill. 3 8 . 1 8 3 . asiony y p y . . , Chlor . 1 m Cladodium calo h llum R Br B . B ol . . : : . . r Synony p y ( ) rid , y Univ 2 2 6 0 . 1 8 6 .

P t m n chostomum cr o hilum Mat . R . co b . ov . y y p ( r ) J Spence , B m B Afl l Na k 1 : r um cr O hilum Mat K . V A . . turs . 5 : asiony y y p r , . 1 83 1 N m in r F rb 956 . o . . B . obtusi olium L db . Ofv e s. o . , nov f ,

Kon h nk a t n " 2 44 1 B c Sv e V e e sk . . 3 : 5 . 866 . s . Akad ; not btu i B 2 1 o s olium Mu scol. R 5 . 803 . f rid . , ecent

Pt ch tomum curvatum K Am R comb . n ov . os u . . I . . y ( a r ) Spence , B m: B um curv K Am B t . 6 1 8 97 . r atum ur. . o . 7 . asiony y a , Not

P t hostomum o h r mb . n ov . c c cl h llum Sc wae . . R . co y y p y ( g ) J Spence , B m M 2 Mu u . : n ium c cl h llum chwa r S . o S e . . asiony y p y g p sc S ppl , 1 60 l 1 4 1 2 9 . 8 7 , p . .

22 Phytologia (Mar 2 005) 8

R m n r r R Br . b . ov . chostomum u u ascens . . . co s p p ( ) J Spence , R Br l r M l ill m P ohlia ur urascens . . C o . e v . 3 9. 1 823 B : . asiony p p ,

R R . comb . n ov P l chostomum reedit s . . y ( obin ) J Spence , R B l 1 1 B m: Br um reedii s r o . 69: 07 966 . asiony y obin , y ,

B R mb . n v P t chostomum rutilans . . . co o . y ( rid ) J Spence , B 4 B m Br um rutilans B . r ol. . 1 : 68 1 826 : . . asiony y rid , y Univ

im L r. . R . comb . n v P t chostomum salinum . ex o . y (Hag p ) J Spence , im r L r L . aubm. D eutschl 2 B m: B um salinum . ex : asiony y Hag p . 4 1 2 3 3 . 89 .

m n P r hwa r R b . ov . t chostomum schieiche i Sc e . . . co y ( g ) J Spence , B m Br r hwa M F : um schleiche i Sc e r. S . u . . u . asiony y g , p sc rond S ppl 1 1 1 2 1 1 l . 6 : 3 . 73 . 8 . , , p p p

P r m n I . R . co b . ov . t chostomum tu bina tum Hedw. y ( ) Spence ,

B m M r H M 1 : n ium tu binatum edw. S . u . F . 1 91 . 1 80 asiony , p sc rond .

P ch s o u w rn m R R m n v t o t m m a eu . . . co b . o . y ( ohl ) J Spence , B m: Mn um e i w rneum R D Pl i ca s ticium . a o l u . . asiony p var h , e tsch Kr d. to . Gew. 3 : 5 1 8 1 3 e 9 . ( yp g .

P t h m m w R m n c osto u ei elii . . . co b . ov . y g (Spreng ) J Spence , B m: Br w M m Fl um ei elii . . . . . 55 . asiony y g Spreng , ant Pri Hals 1 807 .

P t chos tomum wr htii L R m n i u . es . I . . co b . ov . y g (S ll q ) Spence , B m: B um wri h L m Ar r tii u . es . . . . asiony y g S ll q , Proc A er Acad ts 4 2 : 78 . 1 860 . Sci .

Several additional species of Bryum that may belong in P t chostomum m m M y have been reported fro North A erica . ost are k m m poorly nown Arctic species , and ore work is needed to deter ine

- tu . B i i B ti orm Lim r B ax l bl ttii Ph l b . . : . acu e . e their sta s These are f p . , y , Phytologia (Mar 2005) 8 23

Kin B R Br An t B . br ch neuron db . . ha en ii Lim r br oides . . s r a . y ( ) g , y , g p , B Lin B d n e Kindb . . ter s Lin bneo ame s e db . B nitidulum db . . su . , , and

Br um u u The following three species of y also occ r in A stralia , and are transferred for the Flora of Au stralia (Spence R amsay

B R H P R m P t chostomum an usti olium . . . . . y g f ( rid ) J Spence a say , M m no B m Br um an sti olium B . u . R . co b . v . : u asiony y g f rid , sc ecent

’ 1 1 1 m B m ae cium Hedw u . 3 : 3 8 7 . : r u c s itz . S . S ppl , Synony y p , p B B l M F 1 80 1 P hostomum caes iticium . r o . u . . l c sc rond , , not y p rid , y 2 1 : 83 7 1 8 7 . Univ . ,

Since the combination P tychostomum caespiticium has priority as a syn onym of P tychostomum pendulum a new name is needed for the fir m c es iticium Hedw. st species known as B . a p The na e that still B B an sti olium . remains valid within that species is . gu f ridel

P r rr R H. P R m t chostomum c ebe imum I . . . y (Taylor) Spence a say , r B m n B m B um r rr m m . ot . co b . ov . : c ebe i u asiony y Taylor, Lond J 4 5 : 54 1 8 6 . .

m P H w R H. P . R t chostomum seudotri uetrum ed . . . y p q ( ) J Spence a say ,

om n ov B m: Mn u eudotri ue rum Hedw. c b . . asiony i mps q t ( ) 1 2 Schwae r. . M . u : 1 1 0 . 1 8 1 6 u . . g , Spec sc S ppl ,

L M R H P R m tat . n ov . e tostomo sis C . u . . . . . p p ( ell ) J Spence a say , s P R m R H. . t L M . lium C . u . . : . s sl Lecto ype y y ( ell ) J Spence a say , M m n B m B s Mu . . u . co b . ov . : r um st lium C . asiony y y y ell , Syn sc F 1 2 4 : 3 0 . 1 8 8 . rond .

The species in section Leptostomopsis of Brachymenium comprise m a distinctive group that superfi cially resembles Leptostomu . They are z m u characteri ed by densely co pact c shions on bark or rock, dense m hair oint u la ina areolation , a long hyaline spinose p , and erect caps les m 1 996 first with redu ced peristo es . In an earlier paper (Spence ) I

su ggested that the section may represent a distinct genu s . The work of 2003 u r Hedderson 2003 ci al. Cox ( ) and Pedersen ( ) s ppo t this , 24 Phytologia (Ma r 2 005) 8

u u m B s ggesting that the gro p is basal to the re ainder of the ryaceae , and Brach menium Bi only distantly related to the rest of y as well as fyum. This section is sufficiently distinct from other members of the Bryaceae

in both genetic and morphological characters to warrant generic rank .

u 8 - 1 0 u m There are abo t species , pantropical and so thern he ispheric in

u u . distrib tion in drier tropical and s btropical regions One species , the

n m u US A . type , is k own fro so thern areas of the

R n n P la iobr oides I . . e . ov . g y Spence , g

r f liati F u sicco u uniformite o . Ca les olia ovata , in stat contorta , areolatione laminali subh etero enea cellulis distalibus brev ibu s g , irre ulariter sub u adratis - rhombicis m u g q vel brevi , in parte ediana atq e

‘ roximali cellulis- u - rhombicis mar inibu s limbatis p pa lo longi , g distincte

bi- V el mu ltistratosis sectione transv ersali cellu lis du cum valde , costa in in strato unico adaxialiter supra stratum stereidarum bene effectum sitis

ra dita G mm u nullae dioicae . u p e . e ae asex ales . Plantae Caps lae subz omor h ae u inclinatae subrectae m um yg p , ore obliq o , vel ; peristo i di lole idum se mentis endostomialibu s u u p p , g longit dine aeq is

i i i - m exostomio cili s m erfect s mu . S orae 1 0 1 5 u . , p vel llis p parvae ,

tem 30 mm m S s elongate , to , si ple , rarely branched , in transverse

- section with weakly developed central strand , with large thin walled

m m - cells exterior, and so ewhat s aller and thicker walled peripheral

- z u . ea e cells ; evenly foliate; rhi oids ab ndant , red brown , papillose L v s dr pink , red to brown , ovate , keeled , contorted and twisted when y, mm u tu spreading when wet , concave , long , apex ac te to ob se , m u oderately to strongly and longly dec rrent; costa broad , strong , not

ur - reaching apex to rarely perc rent, in cross section with enlarged

u - - g ide cell like layer, one to two layered , ventral to a well developed

m m u - m stereid band , so eti es g ide like cells lacking; la ina areolation

m u - u so ewhat heterogeno s , thin walled , with short irreg larly

subquadrate to short- rhomboidal cells distally and somewhat longer u m m m 60 irreg larly rho boidal cells in edian and proxi al part, typically ,

1 00 m 1 2 - 3 0 m m u x u wide , argins with strongly differentiated border, bi mu ltistratose - ecializ ed a exu al to , of thick walled cells . Sp s mm l iti n fl x e e ae exu a con d o u . eta e u os g lacking . S dioico s S slender, to Phytologia (Mar 2 005) 8 25

a u le u m z m bent near base . C ps s s berect , so ewhat ygo orphic and mm ri u m u asy et c with obliq e o th , apophysis slender, well differentiated , u um u u u m u m operc l conic , witho t apic l s , peristo e do ble , exosto e teeth m u m m lanceolate , so ewhat nited at base , yellowish , basal e brane m hyaline , high , seg ents slightly keeled , with narrow perforations or m m u m r . o es slits , the sa e length as exosto e , cilia r di entary or lacking Sp m 1 1 - 1 4 um t m z s all , (description of sporophy e fro Ochi and Sala ar Allen

P r : la iobr oides inc assa tolimbata . I R Type species g y (Card ) . .

mb . nov . B m B r co : r um inc assatolimbatum . Spence , asiony y Card , R B l 1 1 4 1 ev . r o . 3 6 : 909. y ,

Plagiobryoides incrassatolimbata is a distinctive species that n m m B can ot be confused with any other e ber of the ryaceae . The extremely lax lamina areolation and often mu ltistratose limbidium are R m m m diagnostic . e arkable for the fa ily , la ina cells near the leaf tip are m u m often nearly isodia etric to subqu adrate . The caps le is si ilar to that u Pla iobr um u Br um fo nd in g jy and its allies in the broad sense , incl ding y cellulare and related species . This distinctive plant is found on wet u u r m calcareo s rock and is a Neotropical species , distrib ted f o the

u M m . southwestern U . S . A . thro gh exico and Central A erica

Haplodontium Hampe

Haplodontium consists of species with lateral sporophytes and an r m m 1 985 u t . ext e ely red ced peristo e , ypically of one layer Shaw ( ) transferred all species in the genu s to Mielichhoferia Homsch . u D NA Hedderson However, the recent work sing analysis (Cox and 2 2 2 2 Hedenas 2002 003 ; Cox et al . 000; Pedersen 00 ; Pedersen and ;

20 Mielichho eria M. et al . 03 Pedersen ) indicates that the type of f , mielichho eriana m P ahlia m f , is ore closely related to , while any other species originally in Haplodontium are deeply nested with the B M m . ryaceae . These include the two North A erican species macrocar M tehamen is ameto h ticall r a . s p and , which are g p y y ve y close Ha lodon tium Pla iobr um m to other species of p and g y , and very si ilar to 26 Phytologia (Mar 2005)

H ar u . me aloc um . two the type of the gen s, g p Arnott These species are thu s transferred to a resu rrected Haplodon tium as :

R m r r m D umm. Ha lodontium ac oca u . ex J. . comb . p p (Hook r ) Spence , D r r ex umm . M n ov . Basionym: Weissia mac oca pa Hook . r u sci 2 m B N m . 74 1 8 8 . A er . , rit . . A er .

H i m ehame e R comb . n a lodont u t ns . . ov . p (Showers) J Spence , B m: Mielichho eria tehamens is Br ol 83 : 3 6 . 5 asiony f Showers , y 1 0 98 .

Ack n owledgments

This work was supported in part by a grant from the Flora of North m M ur B G ffi A erica Association . The isso i otanical ardens provided o ce due R M B u space . Thanks are to ichard Zander, arshall Crosby and r ce m am u Allen for their support during y stay . I gratef l to Patricia Eckel M R for providing the Latin diagnoses . arshall Crosby and ichard

Zander kindly reviewed an early draft of this work .

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h i H l z r" ll n r c . N. a a a e . 1 990 . 3 1 mm inc assatolimba tum O , and S A 3 new m fir t 2 t . B ol 3 86 s r . 9 : to Pana a , with description of its sporophy e y 28 7 .

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d r n L e e se N. . Heden as 2002 . P , and . Phylogenetic relationships Br 4 2 B ol . 2 : 77 between yum and supposedly related genera . J ry 28 9.

f iffer L 1 8 N m ni sum tibu s e . 73 . o enclatur Bota cus. . P , I(I) Cassellis , p F 808 Theodori ischeri , vi pp.

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B t 1 1 - 1 1 r J. o . G . 7 : 07 8 . B yum from Au stralia . Adelaide ard

H P R m 1 n ce JR . and . . a sa . 999. Spe , y Proposal to conserve m Br um Mu B the na e y ( sci , ryaceae) with a conserved

4 2 - 2 8 : 8 7 8 8 . type . Taxon

msa 2002 . u Anomo r ence JR . H.P . R a . b um Sp , and y The gen s y

B B - Schim . u . 9: 777 792 . p ( ryopsida, ryaceae) in A stralia Telopea

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1 m in . 5 . G Vol . overn ent Printing Service , Canberra ( press)

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Z I I tl ‘ R mm t : 7 . S a . 2002 . t J , G eport of the co it ee for bryophy es

5 1 793 - 794 Taxon : .

3 0 Phytologia (Mar 2005) 8 7(1) genera in terms of the ease by which a specimen or even an entire plant r mu ifi may be assigned to its co rect species . The ltiplicity of spec c u fl oristic u epithets fo nd in works , together with the s btle m u orphological differences by which the vario s entities are separated , has presented the u ser with an excessive number of alternatives and R little prospect that his choice will be correct . eproducibility the bedrock of scientific enqu iry in this case the probability that independent observers of competent background will assign the same m R ubus u na e to a given clone , has been replaced by a q icksand of individu al interpretation .

This imprecision of identification has produ ced in the minds of many workers distaste for the genu s R ubus and a disinclination to " i n u spend time and attent o in the resolution of its problems . The hiat s has been filled by a few dedicated and exceedingly patient stu dents whose very expertise has resu lted in the recognition of ever more taxa r and a further widening of the ba rier to conventional enqu iry .

A casual observer whose interests lie primarily in the identification of plants of a limited area may not fully appreciate the explosive proliferation in the names assigned to R ubus in eastern North America . Until the last years of the 1 9th century less than a doz en names were of mm u u fl owerin co on sage , and incl ded the wholly distinct raspberries , g

- u . m raspberries , and clo d berries The blackberries , where ost of the m i l H d fficu ties u m u W . . taxono ic lie , received no serio s exa ination ntil Blanchard studied them in the field from Newfoundland to Florida and B 1 1 m u . 91 u west to the geographic li it of the gro p y , altho gh Blanchard had concluded that eight species include the great bu lk of " our t m blackberries , perhaps nine y percent of the , he felt obligated to z m 3 6 s ecific B G recogni e and na e additional p entities ( ailey , entes Herbarum

u m M L. Other a thors then saw need to na e still other species . . ' F u m B m 24 ernald , altho gh he accepted any of lanchard s na es , gave additional specific epithets to North American blackberries (Gray m T P A R 24 H. A . . u . . u Herbari index) . ydberg described a f rther , and D 9 m m avis described ore , while still others were described and na ed Phytologia (Mar 2005) 8 3 1

N L B K m T C . . . W . W . J. . L H by . Porter, ritton , Ashe , S all , and . .

Shinners .

But u m field u the a thor of greatest i portance , whose intensive st dies um u u R ubus m tu and vol ino s p blications on span ore than half a cen ry ,

L. H. B . R ubus was ailey His interest in grew slowly , with a single ifi u 1 1 2 B 1 934 spec c epithet p blished in 898 and a second in 90 . y

Bailey had published a total of 52 specific names . His perceptions by u z u m : then were ac tely sensiti ed , and a prof sion of new na es appeared 42 1 94 1 9 1 943 40 1 944 74 1 45 8 1 1 4 W 7 9 9 7 . in , in , in , in , and in ith a m B few additional na es in following years , ailey was responsible for the m u description , the na ing , and in nearly all cases the ill stration of a total R of 3 83 species of ubus .

Yet Bailey did not feel that the reservoir of undescribed species m u B ato m was near depletion. His co pendio s Species ru (Gentes

H r a 1 - 2 1 94 1 - 45 3 90 e b rum 5 : 93 . ) provided coverage of the species of R u u n z him m two b s then recog i ed by in North A erica . In the next years umu u 76 u he acc lated a f rther species , as described and ill strated in his

u m R ubu Her a m - 4 s pple entary Studies in s (Gentes b ru 3 9. ' But 466 u z m B the species he th s recogni ed were no ore than , in ailey s " u - um opinion , nearly or q ite one half the n ber of species native on the " continent .

' u m B Altho gh North A erica is , in ailey s words , " m R ubus u ost fertile area on the globe for , E rope has fared no less well m m ifi in ter s of the nu ber of spec c epithets applied to the genu s . Perhaps refl ecting the greater density of botanists more than the ' u u m u fl ora E ropean s bcontinent s relatively i poverished vasc lar , the basic texts of Europe treat large numbers of minutely distingu ished R ubus um z m m species . The vol e of recogni ed na es has co pelled a m stratage for their handling , by the segregation of species into m categories , or levels of i portance , with only the principal species or " " - H u in m u . u . circle species given f ll treat ent In central E rope , H ber (

F M 2 4 - 4 1 1 1 964 - 66 z 3 3 u IV/2A z 7 . Hegi , lora von ittele ropa ) recogni ed 252 u basic species and s bordinate species ; these latter were described , but in different type siz e and without inclu sion in the main key to 3 2 Phytologia (Mar 2005) 8

m m m u fl oristic species . In the ost co prehensive odern E ropean

- F u - 2 5 . 1 68 m Y . 9 treat ent , Heslop Harrison ( lora E ropaea ) " " n 75 R ubus 66 - ack owledged species of , of which were circle species representative of groups which contained an additional 374 i undescribed and unkeyed related spec fic names . Since there is no r m m provision in the Inte national Code , nor accepted syste atic dog a , for the existence of species of more than one level of biological m u u m i portance , the E ropean stylistic practice , tho gh aintaining the m u tu u m se blance of a workable str c re , is as yet nexplored in its ore fundamental implications .

u s ecific R ubus u The red ndancy of p epithets in , both in E rope and m u A erica , has not failed to bring forth skeptics who q estioned the fic m biological signi ance of the na ed entities , even their very existence

m u . m E . P B n and the otivation of their a thors The A erican . ick ell , m u 70 u defined Sis rinchium hi self the a thor of obsc rely species in y , ' expressed his unease at Blanchard s many R ubus names with an article “ ” plaintively entitled Have we enou gh New England blackberries"

- Bu B t . 4 3 D o u 0 . t ( ll . Torrey Cl b efense of the reali y of L H B the new entities has rested largely with . . ailey who provided a At m rationale with each of his major treatments of the genu s . no ti e was Bailey unaware of the unconventional image that his many species u m m He m 1 923 prod ced in the inds of classical syste atists . re arked in “ (Gentes Herbarum In is not possible always to apply the formal species concept of ante - evolutionary days with either ” 4 “ 1 9 1 G. H precision or satisfaction . In ( . he said , The reader ma u fi l y s ppose that I have split the species ne y. The opposite is the u u um m u u tr th . I co ld have described any n ber ore if I had cared to p rs e 1 " u . 943 G. H. mm W a separatist co rse In ( he co ented , ith so m ma any kinds now separated , the reader y wonder whether every tu He colony is not a distinct species . This is a na ral u 1 44 H R ubus ca tioned in 9 (G. . not critical in will u m m mu have little s ccess with either pictures or speci ens . The ind st " first m m be free of notions and then the eye u st be able to discri inate . m 1 947 G H “ u . And he rested his arg ents in ( . with the defense , I “m z m have never ade a species ; I have only recogni ed , na ed and m described the . Phytologia (Mar 2005) 8 3 3

Bailey gave repeated attention to the supposition that the many closely defined species recogniz ed by him were the product of prolific z m m m B n hybridi ation a ong a li ited nu ber of true species . ick ell had been insistent that the entities named by Blanchard could be accounted t m 1 1 for as hybrids be ween no ore than species . Unwillingness to accept su ch an explanation was a theme that persisted through all of B ' ailey s works . His hostility to what he considered a glib and fi ial r H. supe c interpretation was scarcely concealed (G. “ Early in the present century began the singu lar hybridity postulate in R ubus r um m um work , whereby hybrids were f eely ass ed fro herbari specimens so fragmentary that not even the species themselves can be determined; thereby was the fear of making new species escaped and diffic lt a He the u y of underst nding the plants was assuaged . did not t tu t u but deny the possibili y of na ral hybridi y in the gen s , challenged that its existence had yet to be proved and insisted that the entities described by him were readily recogniz ed by a person of observant eye ffi i n and su c e t field experience .

Yet even as stu dents of R ubus determinedly pursu ed the self immolating course of recogniz ing and naming a seemingly endless u u B u series of d bio s new species , and as ailey shr gged off the shallow proposals that nothing more was involved than a massive hybrid m m fields m swar , conte poraries in the of e bryology and cytogenetics were disclosing a framework of understanding on which a workable m taxono ic structure cou ld be hung . This was the discovery that reprodu ction by seed was a less than universally sexu al process . In

u infl uenced u A . E rope , partic larly by the work and writings of Gu m m u staffson , the concept ca e to be held that any genera of vasc lar m m m u plants display the pheno enon of aga osper y , or reprod ction by but u z seeds witho t fertili ation , as a special case within the general R ubus m process of asexu al reproduction or apomixis . was a ong the numerou s genera in which agamospermic reproduction was su spected identified or .

m u m m In North A erica , and partic larly a ong classical syste atists , B such a concept was not quickly applied or perhaps understood . ailey 34 Phytologia (Mar 2 005) 8 seems never to have given written expression of awareness that the u u m m u individ als constit ting any of his species ight be of asex al origin , althou gh his activities were indeed largely prior to widespread He acceptance of the agamospermou s pathway . referred to the process “ H am u G. . only once , by indirect incl sion ( I asked for proof or at least for opinion that my novelties are not hybrids or apomicts or other course no man has such

But B u ailey was nonetheless in search of s ch proofs , whichever

‘ m In 1 944 u u way they ight point . he had beg n the s pport and u m Einset u enco rage ent of John in New York State , in his st dy of the R B 1 m 1 95 . cytology and embryology of variou s ubus clones . y (A er J

B - 772 Einset u first otany . ) was able to p blish the satisfactory evidence that in North America R ubus there operated certain apomictic phenomena that had previou sly been demonstrated only in Old World 1 spe0 es .

Einset worked with 24 wild selections of R ubus which he brou ght u B i ntifi He u into c ltivation and which ailey de ed. fo nd the chromosome numbers to form a regu lar series of multiples of the basic 7 r m 1 4 hr u 63 u W , ranging f o t o gh in the species st died , ith only a single

u um u . clone having an ane ploid n ber o tside this series Triploids , with m um 2 1 m m mm u the so atic n ber of , for ed the ost co on gro ping , m u representing a third of the clones exa ined . As had E ropean workers u u previo sly , he fo nd that pollination was necessary for seed B m m um produ ction . y crossing clones with different chro oso e n bers u m m s u and by co nting the chro oso e of the res ltant seedlings , if the seedling count matched that of the seed parent and differed from that of m m m the pollen parent he could assu e that aga osper y was present .

' Einset s work strongly supports the assumption of apomixis in the Am r m m um erican blackberries . When the ch o oso e n bers of the seed parent and the pollen parent were different , a high proportion of the 80 96 seedlings ( per cent in the case of tetraploid seed parents , per cent with triploid seed parents) gave the same chromosome count as that of m m um the seed parent . Had there been redu ction of chro oso e n ber with Phytologia (Mar 2005) 8 3 5

m m u m u a chro oso al contrib tion fro each parent , as in sex al u m u u mu reprod ction , inter ediate co nts wo ld have been obtained with ch Ein set greater frequ ency . cou ld only conclude that a high percentage of his R ubus progeny resu lted from the parthenogenetic development of r un educed eggs .

m u R ubus u m The echanics of reprod ction in th s beca e clearer . B m u lackberries spread by vegetative eans , with r nners and rooting

- m shoot tips increasing the extent of the colony . Seeds are also for ed , and serve as a means of dissemination across natural barriers and over At m distance . ti es these seeds are produ ced by familiar sexu al u m u processes , and generate individ als that show the in te differences m characteristic of genetic reco bination . Perhaps also disparate u m m m individ als co bine at ti es , to yield hybrids which differ fro the offspring of conventional sexu al reprodu ction only in the magnitude of t their variabili y .

But R ubus m m in a less fa iliar generative ode is also present , and ma m m u t y well for the do inant reprod ctive pa tern within the genu s . a amos ermou s u This is the g p pathway , in partic lar the pollination u n m seudo a . u u req iring variant k own as p g y The progeny th s prod ced , mu u u while si lating the offspring of sex al reprod ction , are identical in genetic composition and essentially identical in morphology to their m B aternal parent . y the agamospermou s replication of these u tu um ma individ als in their rn , large n bers of clones y be generated .

W - u m ith seed eating birds as vectors , their distrib tion will be li ited only by factors of habitat availability and physiologic adaptability .

A student inexperienced in R ubus who detects one of these agamospermic mu lti - clonal series is compelled to V iew it as he would u He find an ndescribed species . s the plants to have a coherent u u u distrib tion , occ r in predictable habitats , req ire pollination and u m reprod ce by seed , and vary orphologically within a narrow range . z him m His novelty is recogni able to , and in al ost every regard possesses the criteria that he associates with conventional species . It is perhaps inevitable that he should wish to bring legitimacy to his 3 6 Phytologia (Mar 2 005) 8

discovery by publishing its description and by coining a new name to serve as its label .

R On And thus confl ict arises in the study of ubus . the one hand is u u m m the insistence of close st dents of the gen s , ost of the of u u m m field nq estioned taxono ic co petence and extensive experience , z u m m m that they can recogni e entities that are nifor , at ti es in any m On colonies distributed over hundreds of iles . the other hand is the practical inability of less practiced workers to distingu ish reliably among entities so very numerou s that only the finite energies and i h ma i ntifi r m lifetimes of the r u n de e s appear to restrict their nu bers .

m m This confl ict has no elegant solution . The organis s the selves create the hierarchy to be described , and it is neither linear nor u u u consistent in its structure . The series of eq ivalent nits of a sex al m a am rmi m syste does not have a parallel in an g ospe c co plex . No u seful purpose is achieved by insisting that apomictic microspecies can z sufficient u be recogni ed with st dy , for they can neither be keyed by i ntifi m de ed . the specialist nor by the workaday taxono ist A coarser, less precise classificatory structure seems to be the only feasible approach .

Thu s one is led to a system that recogniz es as species only major u u R ubus gro pings of the gen s . In the blackberries only one species

u u z . m B appears sef lly recogni ed in each section In North A erica , ailey

G H r a m - 4 1 94 1 n ( entes e b ru 6 . ) ack owledged ten sections in the ru ri u u t e blackber es ; the present scenario , if rigoro sly followed , wo ld u red ce the North American blackberries to this number of species .

m r fl owerin u - A ong the raspbe ries , g raspberries , and clo d berries the m u u sa e degree of red ction is perhaps nwarranted , for it is not clear that a am mi m B t os er c . u the g p process is as do inant there the blackberries , m at least , appear ore easily handled as a few aggregations of related m for s than as constellations of numerou s related microspecies .

Such an approach implies the assumption that each aggregation a sin le m consists of , g basic species and its derived apo icts . Extensive R ubu G s u mu Gu V . work on in E rope , ch of it by stafsson (see rant ,

3 8 Phytologia (Mar 2005) 8

four sections and shou ld be u sed to denote the major aggregations of R ubus u F : R ubus ar u tus n R ubus cunei olius as fo nd in lorida g Li k, f r R bus tr vialis M R b a ella is W . u i u u us . P rsh , fl g illd , and ichx

L B 1 R ubus . lackberries

m u 2 Ste s trailing or s pported by low vegetation , elongate (to u um u densely set with both sto t prickles and n ero s stiff bristles , u u m r u u s ally ahogany red; leaves persistent th o gho t winter, u u leafl ets u fl owers u u s bcoriaceo s , glabro s , dark glossy; s ally

u . solitary , on erect pedicels ; petals white . Prickly trailing shr b ’ fi l r u e ds. u u mm Thickets , open woodlands , br shy Th o gho t; co on . R B R B a ilis . continen talis R . lucid Spring . [ . g ailey; ailey; us R R r B R B . mi us . Okeechobeus ydb ; . ailey; ailey" R D WB R R R t ri i li M ub u s a s . SOUTHE N E E Y . v ichx

1 m . Ste s erect, arching at tips , or declining and rooting , with prickles but r u u leafl ets with few or no slender b istles; leaves decid o s , thin ,

glabrou s or densely pubescent below; fl owers mostly in 3 - several fl owered panicles .

m u but t Ste s erect when yo ng soon arching and declining , of en m rooting at tips ; prickles weak , sparse , often al ost lacking;

u m. M petals white . Prickly arching shr b , to esic fi l M e ds. woodlands , clearings , old Panhandle (east to adison u t R t S . ubus enslenii Co n y); infrequent . pring [ Tra t" R R D r R u b u fl ella is W . NO THE N EWBER R Y . s ag illd

2 m m . u Ste s re aining erect , altho gh often arching toward tip , not m u u u u . rooting; prickles sto t, n ero s , rec rved

asic mor o o ica framework of a conv entiona ic otomou s ke as we as ata on b ph l g l l d h y, ll d a itat ran an T i a r i ntin ati n f a ri in the 1 970s e d fre uenc . s e s a co u o o se es e un h b , g , q y h p p b g

V i e Ph tolo ia - 4 1 I w t nk i n K nt D P rkin is o t a D av W . Ha a d e . e s ( d y g 3 . h h d ll for constructiv e rev iewin th e man scri t ly g u p . Phytologia (Mar 2005) 3 9

Leafl ets u so m ovate to oblong , glabro s or nearly ; ste s mm 2 m m 8 . u co only to . (to , when s pported by

surrounding vegetation); panicle often several - fl owered;

u . M petals white ; fr its sweet Prickly arching shrub . oist m m r u u to dry thickets , pond argins , swa ps . Th o gho t

nh F u mid- u pa andle and north lorida , so th to penins la R u mm . . (Highlands , Okeechobee co nties); co on . Spring [

rid R . r r B R . betulifolius Small; R . fl o us Tratt; ka pe i ailey; B R r R m R ran hodo h llus . . penet s ailey; . p y ydb in S all ; B u e a u B R . z oae tallahasseanus ailey; R . c t n s ailey; Bailey" G B B KB R R R u b u s ar u tu s HI H USH LAC E Y . g Link

Leafl ets - u m obovate , densely gray p bescent beneath; ste s

u u 1 - 3 - fl owered u su ally less than 1 m. ; panicle s ally ; u D r petals white; fruits bland . Prickly arching shr b . y fi ld i tu r u u e s . sands , old , d s rbed areas Nearly th o gho t R u u mm . . . audax (excl . so th penins la); co on Spring [ R r B R B . cha manii . o idensis Bailey; R . p ailey; fl ailey; B R in erior B humei ailey; . f ailey" D B KB R R R u b u cu n eifoliu s u SAN LAC E Y . s P rsh 40 Phytologia (Mar 2005)

A NEW SPEC IES OF SE YMER IA (SCR OPHULAR IACEAE) F M M RO OAXACA, EXIC O

B L rn . . T u er

R u u 787 1 2 047 1 Plant eso rces Center, University of Texas , A stin USA

AB STR AC T

m ria maz a ecana B . L. u s n Se e t . ov . m y T rner, p , is described fro

M . de r S. curva but northeaste n Oaxaca , exico It is closely related to is readily distingu ished from that taxon by vestitu re and shape of its m u . at re capsu les A photograph of the holotype is provided .

KEY W R D m r M : u Se e ia . O S Scroph lariaceae , y , exico , Oaxaca

R outine identification of Mexican plants has revealed the following novelty :

m r m Se e ia a atecan a B . L s n o Fi 1 u v . . y z . T rner , p. g

m r Se me iae decurvae B . D h r a C . e b e Si ilis y enth ex , sed plantae com actae u V et u ov oideae p , ca les dense illosi , caps lae late (vice ca su larum lli i a m p e pso de ru ) sunt .

m - 1 5 30 cm . m Co pact perennial herbs high , drying black Ste s densely villou s with multicellular spreading or refl exed hairs mm mi - m m - 1 2 mm . d 6 long Leaves pinnately lobed , those at ste ostly

5 - 8 mm u u u long , wide , p bescent with both short hispid lo s hairs and

- u m m capitate gland lar hairs , interspersed a ong these a s attering of u m Infl orescence longer crinkly hairs s ch as occur on the ste s . a

m m 1 2 m - 2 fl w r ter inal leafy race e up to c long bearing 1 0 0 o e s. Pedicels

8 - 1 1 mm - mm - 2 - 4 long . Calices 6 7 long having 5 8 irregu lar lobes

mm - . u u u u long Corollas yellow , arc ate rceolate , the t be glabro s

u 4 mm mm ca. 8 u ca. 3 witho t, p bescent within , long , wide , the throats

mm 6 mm - u long , wide , capitate gland lar , their lobes broadly cordate , Phytologia (Mar 2005) 8 7(1) 4 1

Fi . 1 . Se meria maz a tecana g y , holotype . 42 Phytologia (Mar 2 005) 8

4 - m 4 mm 5 mm m . 4 ca. long , wide , having ciliate argins Sta ens ;

4 - 5 mm filaments mm anthers long; long , densely pilose to the mm so . u ca. 8 6 mm apex or nearly Caps les broadly ovoid , high ,

- u t 6 - 8 mm across , capitate gland lar, the s yle persistent , long . Seeds

1 u ca. 2 mm mm ca. 5 a m ur c . mm (so ewhat i at e) per loc le , long ,

m u u . wide , their argins with narrow nd late wings

M M : . : . m TYPE EXIC O OAXACA pio San Jeroni o Tecoatl , ” Vereda (antiguo camino real) del Plan de Guadalupe al Cerro Pelon a 2640 m 6 Jul 200 1 Xochitl Munn 1 8 08 1 0 x 96 58 20 W c . ( N ) , , l 1 1 MEX 3 5 : U : . Estrada et a . (HOLOTYPE ; ISOTYPE TEX)

This species clearly belongs to the V irgatae group of Seymeria as n 1 2 treated by Pe nell ( 9 5) where it will key to the widespread S.

' m r m Se merza decurva . It will also key to the latter in ytwo t eat ents of y M 1 82 t m u 9 S. for exico (T rner, ; The present novel y differs fro decurva m m m u in being a s aller, ore co pact plant having a villo s vestitu re on its stems and broadly ovoid capsules (as opposed to ovate

u u . dec a ellipsoid caps les as consistently fo nd in S urv ) .

m decurva u The only speci en of S. previo sly collected in the state F ar as 3264 V of Oaxaca has been that of p ( icinity of Cerro Verde , W m m tu this hou sed at UC) . hile I exa ined the latter in y initial s dy of the group (Turner I can not recall if it was in fruit or not; at least u m I did not record the caps le as broadly ovoid , nor the ste s as being u m z villo s , characters e phasi ed here . Nevertheless , it is possible that the

m z m m S. ma atecana speci en concerned belongs to , e ories being fallible .

m The species is na ed for the sierra where it was first collected . I m but intended to na e the taxon for its principal collector, when told of ” m um u y intentions she h bly replied , Please no , I deplore s ch honors , besides it wou ld be more meaningfully named for the sierra in which it ” But m u was collected , or words to that effect . left to yself I wo ld have m preferred this attractive herb bear the na e of its collector, Xochitl

Mu n - a u m n Estr da . She is c rrently enrolled in a graduate progra in Phytologia (Mar 2005) 8 43

t u fl oristic u botany at The Universi y of Texas , A stin working on a st dy of the mountains concerned .

ACKNOW LED GMENTS

I am grateful to my wife Gayle Tu rner for the Latin diagnosis and M M nn to her and s u Estrada for reviewing the paper .

LITER ATUR E CITED

1 2 A elia : m P n n ell F .W . 9 5 . u z e , The gen s f a taxono ic stu dy in

- 77 : 3 3 5 3 3 . evolution . Proc . Acad . Nat . Sci . Phila . 7

u rn er EL . 1 982 . R m M T , evisional treat ent of the exican species of

r Ph t l i 1 4 - 4 o o a 5 03 1 9. Seyme ia (Scrophu lariaceae) . y g

1 Se meria u r m 995 . A new species of y (Scroph lariaceae) f o

- M Ph tolo ia 78 : 28 3 5 . exico . y g 44 Phytologia (Mar 2005) 8

A NEW SPECIES OF AGER ATUM (ASTER ACEAE ' E PAT R IEAE F M R I M U O ) RO NAYA T , EXIC O

B L rn r . . T u e

R ur u 787 1 2 - 047 1 Plant eso ces Center, University of Texas , A stin , USA

AB STR ACT

A eratum solisti s . n ov IS m M A new species , g p described fro pio . i M m Huacor a . de , N yarit, exico It rese bles the widespread , highly A cor mbosum but u m variable , . y is readily disting ished by a co bination u m u m of characters incl ding deltoid , arkedly retic late leaves , and ore m m m t nu erou s uch s aller heads . A photograph of the holo ype is provided .

KEY W R D : A era tum Eu atorieae O S g , Asteraceae , p , Nayarit , M exico .

In The Comps of Mexico (Turner 1 997) Agera tum is recogniz ed as 1 1 1 2 m having species; the present novelty brings this total to . In y m A solisii treat ent, the new species described below ( . ) will key to or A toment sum B m . o . u u u near ( enth . ) He sl beca se of its s ffr ticose habit

confined - m and thick deltoid leaves . The latter taxon is to the tri co er uz u region of the states of Veracr , P eblo , and Oaxaca and can be

mm m A solisii m i ediately distingu ished fro . by its arkedly bicolored

- i A. so isii ca tu lescence . t l leaves and few headed p In reali y, is probably m r mbosu u . . A. co m ore closely related to the widespread y Z cc ex Pers , but u m m is readily disting ished fro this by a co bination of characters , as noted in the above abstract

A r tum soli B L e a su . . u s . n ov Fi 1 . g T rner, p . g

m A r r laminis foliorum Si ilis ge ato co ymboso Zucc . ex Pers . sed deltatis nerv atis u et ca itulis arvioribus , valde retic late infra , p p luribus ue a i l p q per c p tu escentiam.

46 Phytologia (Mar 2 005) 8

u 1 m m Su ffruticose herbs or shrublets p to high . Ste s densely mm u tur ca. . u hirs te , the vesti e high Leaves opposite , p bescent like

S m cm 2 - 3 cm e cm the te s , long , wide; petiol s long;

u m u - blades deltoid , r gose above , arkedly retic late veined beneath , m m u u . u u s bcordate at the base; argins irreg larly crenate Heads n ero s ,

m m m fl at- ca itulescences arranged in ter inal cory bose , so ewhat topped p

1 2 - 1 5 cm u u m u m 1 - 5 across , and abo t as high , the lti ate ped ncles ostly mm m m u u a 4 c . mm long . Heads relatively s all , ca pan late , the invol cres m a 2 1 2 R c . . high , co posed of linear scales arranged in series eceptacle F 2 u u ca 1 mm mm . 0 s bconical , naked , glabro s , across , high lorets m mm 3 0 m ca. per head; corollas white , long; throats long ,

defined m u - u grading into a weakly throat , ato ifero s gland lar 1 mm u u u . ca. u thro gho t, the lobes p bescent Achenes long , glabro s ;

‘ mallt a r- u s s uce u ca. mm papp s a shaped crown of nited scales high .

M R M H a ri R : . : . u co TYPE EXIC O NAYA IT pio de j , ancho de Los “ u m u B u m ch a orros Sa ces , o th of canyon in osq e Te plado (encinos p y

28 1 8 I. olis 5 1 7 . 9 5 S : . otros arboles) Sep , (HOLOTYPE TEX)

D D MI D : M A ITIONAL COLLECTION EXA NE EXI CO . R m Sol s 52 7 . : t 3 0 1 985 I. i NAYA IT sa e locality as ype , Sep , (TEX)

The species is named for Ign acio Solis Cumplido D on Nacho who is a Tepahu an Indian and collected for CIID R " D urango for several m u r D u years in extre e so thweste n rango and nearby Nayarit, in the canyon country .

ACKNOWLED GEM ENTS

I am grateful to Gayle Turner for the Latin diagnosis and to Tom W a m endt for obt ining infor ation as to the credentials of the collector . M D M H B . S. . ias de oraes and . erg provided the digital photos

LITER ATUR E CITED

u rn er EL . 1 A m M 1 Ph olo ia 997 . eratum t T , g , in Co ps of exico , y g

M m 1 1 - e oirs : 5 1 5 5 . Phytologia (Mar 2 005) 8 47

BID ENS MELCHER TII R E (ASTE ACEAE) , A N W SPEC IES F R OM J M ALI SC O , EXICO

B L . . T u rn er

R u t u 787 1 2 047 1 Plant eso rces Center Universi y of Texas , A stin , USA

AB STR ACT

Bidens melchertii s . nov IS m A new species , p described fro the

- M . u mm state of Jalisco , exico It is closely related to the sq are ste ed ,

- B rin lei but m . u white rayed , perennial , p g , is readily disting ished fro the latter by characters of the achene .

KEY W R D : Bidens M . O S , Asteraceae exico , Jalisco

Preparation of a taxonomic treatment of the genu s Bidens for m M Mexico by Prof. Tho as elchert of the University of Iowa has necessitated description of the following novelty :

Bid n m h r B L s . n ov . Fi . 1 i . e s elc e ti . . u T rner, p g

h nii W . ac e s Similis Bidens pringlei S. ats sed sine pappis (non ti biaristatis et brev ioribus cor oribu s tetra onis et recurv a s. ) , p g non

m u u z m u 80 cm . Erect rhi o ato s herbs to high Ste s s ffr ticose , leafy u u multinerv ed u thro gho t, the lower portions , the pper essentially

1 - 2 u u so . sq are in cross section , glabro s or nearly Leaves deeply ( )

cm 1 - 2 cm u m pinnatisect , long; petioles long; lti ate divisions

- mm m u . linear to lance linear, wide , their apices so ewhat p ngent

- R a fl orets 8 u 2 3 cm . Heads showy , across the expanded rays y ; lig les

- m D fl orets um u m 1 2 mm ca 5 m . 6 . crea y white , long , wide isk n ero s , u ca 1 0 u yellow . O ter phyllaries , linear with sharply ac te apices ,

- R m . u recu rving with age . Inner phyllaries yellow argined eceptac lar

- bracts linear lanceolate , their apices extending beyond the 48 Phytologia (Mar 2 005) 8

1 Bidens melchertii . Fi . g. , holotype (TEX) Phytologia (Mar 2 005) 8 49

fl orets. u ca disk Achenes epappose , black, glabro s , . as long as the u chaff, linear, tetragonal in cross section , the o ter series notably u m m mm inc rved and so ewhat obco pressed , long (the very

u m - n m o ter ost nearly linear clavate , the in er ost nearly straight , to mm long) .

M D M . J Km 1 : : . e 5 m TYPE EXI C O ALI SCO pio Talpa , . Ca ino E1 u M m 1 650 m 1 4 1 989 T ito a ina de Zi apan , pine forests , , Oct , Lomal 2 r D az Luna os A. i 0968 : Ca los i J e S. (HOLOTYPE TEX;

ISOTYPE : MEXU) .

D D M M D M A ITIONAL SPECI EN EXA INE : EXICO . JALI SC O

M D e r M - 2080 pio . Talpa , Ce ros de Las inas de Oro , pine oak forests , 2 M m 0 1 1 R . nz e 6 3 97 Go al s T. 5 . , Oct , ( ICH)

Biden tal na rfi iall m B rin s a su e c . lei p p y rese bles p g , both being squ are - stemmed perennials with deeply pinnatisect leaves having

filiform u m m - nearly lti ate seg ents , and possessing showy white rayed

B r u . in lei heads . Their achenes , however, are q ite distinct , those of p g

fl - u u biaristate having atter, long atten ate achenes with slightly rec rved

B m B . molli olia u . idens ta ana apices (v s. papp s absent) lp also rese bles f

Sherff . , the two taxa having short, clavate to linear clavate achenes

m m m m The species is na ed for y sixth acade ic son , Tho as m M m . u . . elchert, c rrently Prof E er at the Univ of Iowa , A es , and H m Bidens Cosmos Theles erma . e longti e scholar of the genera , , and p called this novelty to my attention in his submitted manu script for the f ur m u m m Bidens M c . pco ing treat ent of for exico ( T ner, Co ps of 1 1 W u M 1 ri Ph t l ia M m . Eu ato eae. o o exico , Vol . . p y g e oirs itho t

u m . his knowledge , I pro dly provided the epony concerned

ACKNOWLED GEMENTS

I am grateful to Gayle Tu rner for the Latin diagnosis ; Marta D ias

M B . de oraes and Heidi S. erg prepared the digital photos 50 Phytologia (Mar 2 005)

A NEW SPECIES OF TR D HS (ASTER ACEAE : MUTISIEAE) F M M RO OAXACA, EXICO

B L u . . T rner

R u t u 787 1 2 047 1 Plant eso rces Center, Universi y of Texas , A stin USA

AB STR A CT

L n m Trixis hinton iorum B . . u s . ov . T rner, p , is described fro

M . So u Oaxaca, exico far as known , it is a prostrate shr b with winged M m confined u . ste s to the rocky beaches of so thern Oaxaca , pio

Pachutla. A photograph of the holotype is provided .

i i Tr M KEY W R D : A Mut s eae ixis . O S steraceae , , , exico , Oaxaca

R outine identification of Mexican Asteraceae has revealed the following novelty :

' r Fi 1 L. u s . n ov . . . Trix is h in tomo u m B . T rner, p g

m i f lii ra i ri s caulibus T. s lvaticae o s c ss o bu Si ilis sed , sine alis ,

fl li m imilari la ii x ri ri 1 - 2 mm oscu s u s bu s b s e te o bus . o nib s , longis

u m mm m Shr b , reportedly (fro label data) co on and for ing “ m ” u . m u u u W . proc bent colonies Ste s p ber lo s , ingless Larger leaves

1 0 m i l n - m m ca. c ca tu esce ce 6 8 c c ( below p ) long , wide , sessile so n m or nearly , relatively thick , pi nately venose , so ewhat bicolored , m - m r u . oderately appressed pilose beneath , the argins se r late Heads

2 - 4 m u ca. u s btended by ovate bracts as long as the invol cres , ostly arranged 3 - 5 in terminal congested cymes scarcely exceeding the

- - . u 1 1 u leaves Invol cral bracts , linear lanceolate , appressed p bescent , u um R u the apices grad ally ac inate . eceptacles densely p bescent , the mm F 2 m ca. 3 hairs long . lorets , the corollas yellow , all ore or

- u 1 2 mm . less alike , those at the periphery with o ter lips long Achenes mm a 5 mm r u u u tu c . u (i a re) long , densely p bescent th o gho t; papp s of

um u - 1 mm n ero s tawny bristles 9 0 long . Phytologia (Mar 2 005) 8 7(1) 5 1

nmmas wm we 37 m m s ama s W . 3 9 43 4 xz xm xxw a m v m s x - a an gz n gggmgssmgaw m z amz m 8 1 1 935 <9: 33 3 3 . 3 9936 59 3 3 3 3 ;

Fi . 1 . Trixis hintoniorum t g (Holo ype TEX) . 52 Phytologia (Mar 2005) 8

M . u : M . : TYPE EXICO OAXACA pio Poch tla , Playa La Tijera , 27 1 9 5 Hinton l 2 9 et a . 6474 sea level along rocky beaches , Oct ,

(Holotype TEX) .

’ 1 972 t m Trixis u W In Anderson s ( ) reat ent of , beca se of its ingless l h r nm m T. me a o lla G ee . / T. ste s , this taxon will key to or near g p y and or

R Greenm. b t m m silvatica u . obinson , it is arkedly different fro both

m T. According to label data and appearance of pressed aterial , r m u u hintonio u is a prostrate shr b occ rring along the beach . In habit m T. silvatica but and leaf shape it rese bles , possesses thicker leaves

m um u - fl owered 1 1 u and larger, ore n ero s heads , having invol cral bracts u Tr hintoniorum arvi l r ixis T. o a (vers s also has characteristics of p f , Gu but a coastal species of northern Oaxaca and errero , the latter n 8 u possesses narrower thin er leaves , the heads with only invol cral

1 0 - 1 fl r t bracts and 3 o e s.

u u T. silvatica It sho ld be noted that Anderson , in her disc ssion of , “ ” mentioned a single popu lation of the latter to have 1 1 - 1 3 involucral

27 - 29 fl oret m bracts and s per head (speci en not cited) . This

fit r s T. hinton io um but t description the heads of , corollas of the lat er

m m 1 - 2 mm v s mm have arkedly s all lips ( long ) and , as already m m . noted , has arkedly winged ste s

ACKNOWLED GEM ENTS

I am grateful to Gayle Turner for the Latin diagnosis and to her

M " s. M and Xochitl unn Estrada for reviewing the paper .

LITER ATUR E CITED

n d r n e so C . 1 72 . m M A , 9 A onograph of the exican and Central m m t r Me . N. Y . Bo . G . A erican species of T ixis (Compositae) . ard

22 1 - : 68 .

54 Phytologia (Mar 2 005)

’

Fi 1 D t vi mo S. g . istribution of S e a lem mi (closed circles) and

(open circles) .

L ITER ATUR E C ITED

m t r Grash off J . 1 972 c u , A syste ati st dy of the Nor h and Cent al m tevia D D S . A erican species of octoral issertation , the University

u . of Texas , A stin

i a 1 4 Stevia m : Eu ator e e . 97 . Novelties in (Co positae p )

Britt nia 26 : 347 - 3 84 o .

u rner B L 1 m M 1 Ph t lo ia M m 1 1 : . 997 . o T , . in Co ps of exico , y g e oirs

1 70 - 1 97 . Phytologia (Mar 2 005) 8 55

om. 2000 . . an d G . Nes Use of variety and subspecies and

’ m St rax latam olius new varietal co binations for y p f (Styracaceae) . ' 2 Sida l 9: 25 7 62 . 56 Phytologia (Mar 2005)

B OTANICAL ANECD OTES : Bajacalia moran u (Asteraceae : Ta eteae mm m R g ) , co ents of Ja es eveal regarding its discovery

B n r . L . Tu r e

R u u 787 1 2 - 047 1 Plant eso rces Center, University of Texas , A stin , USA

2003 R Loockermann et al . ecent description ( ) of the new species , r B L u m r m m Bajacalia mo anii . . T rner elicited the following e ail f o Ja es L R t m . eveal , who participated in its discovery, ype aterial having been collected with R eid Moran on a joint collecting trip to Baja California

during February of 1 973 .

Morning Billie :

A tale about the discovery of Bajacalia moranii .

We drove down Arroyo de Portez u elo looking for Eriogomum in encelioides u 1 947 G . find , a plant fo nd in by Scott entry Upon g a few u out m tu shr bs , we stopped the jeep and I got y s ff to write a description fi l based on what I cou ld see in the e d.

R eid said he had to V isit a bu sh .

I said I wou ld look further down the arroyo to see what the m H e u d . variation ight be . went p the raw; I went down the draw

Ju st around the comer I found what proved to be Eriogonum preclarum and yelled back to R eid that we wou ld be a while as I found

a new bu ckwheat .

He He um yelled back that it was okay . was taking a d p and a new P oro h llum p y was by his side .

one finds Ahhh , how new Phytologia (Mar 2 005) 57

P oro h llum R The new species of p y , declared to be so by eid and confirmed m tu B in the afore entioned s dy with the description of .

' maranti u u but , was never p blished of co rse , had I known the details of its discovery I might have provided a very different Latin name "

I am grateful to my fri ed Jim who permitted my u se of his email r communication dated 1 4 Ma 2003 .

L ITER ATUR E CITED

k rm n D B L u rn r R K n n 2 Looc e a . J . . . e . . J a se . 003 . , , T , and Phylogenetic relationships within the Tageteae (Asteraceae) based r m on nu clear iboso al ITS and chloroplast 71 t gene sequences . B 2 1 1 2 Systematic otany 8 : 9 07 . 5 8 Phytologia (Mar 2 005) 8

B OTANICAL HUM OR

In June of 1 994 my family and I traveled through S cotland for two weeks. It was the second summer after I h ad discovered Botany and I was determined to learn every plant in S cotland by feverishly collecting every species and k in th m n t l r n L wa de ribed e e out i ce One nt Ci cea lutetia a . s sc y g S a p a , , “ ” as r r in l n Not uite underst ndin if rare meant a e Northern S cot a d. q a g endangered or simply not common I fretted that I had extirpated the last o ulation of in tl n M th r r l t thi t r to a ri end lutetiana co d. e e ed s s o p p C. S a y fa a y f of th M ll a teacher of E L at Amarillo Colle e e amil Moll adler. o S f y, y S y, g , illu strates as a hobby and has had works published in th e Wall Street Journal ’ n ith m th r h m lli t as in iration he dre a d Forbes. W y fa e s (muc e be shed) s ory sp s w th The ill tr ti n i r dl i la ed in m office and it is e above picture . us a o s p ou y d sp y y ith much o th I i tin William Editor at resent t in Ph olo ia. J u s s w j y p yt g ,

t New F ra th ti i r i f am ri Pr ss. P 1 226 . S ace C . 1 991 . I t e o of e Bri s s es. Un ve s o C e , l h l y b dg p Phytologia (Mar 2 005) 8 7(1) 59

IND EX TO NEW NAMES AND COMB INATIONS IN PHYTOLOGIA

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P ch ostomu m n W n comb . nov . t ulle s S . . R . e ce y p ( ) J Sp , 60 Phytologia (Mar 2005)

m n hl i h . n m b . nov Pt ch ostomu allesce s c e c . . e ce co . y p (S ) I R Sp ,

m r m H dw R . n P m seu dotri u et u e . J. S e ce H. . R amsa c m n Pt ch osto u o b . ov y p q ( ) p y,

mu m u r u rascen R . Br. . R . ence c m n Pt ch osto s o b. ov . y p p ( ) J Sp ,

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mum tu rbin atu m Hedw. . . ence omb . n Pt ch osto c ov . y ( ) I R Sp , h m m rn m hl n P l c osto u wa eu o . . . e ce comb . nov . y (R ) I R Sp ,

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