Boise State University ScholarWorks Biology Faculty Publications and Presentations Department of Biological Sciences 1-1-1993 Phylogenetic Analysis of the Zingiberales Based on rbcL Sequences James F. Smith Boise State University W. John Kress Smithsonian Institution Elizabeth A. Zimmer Smithsonian Institution This document was originally published by Biodiversity Heritage Library in Annals of the Missouri Botanical Garden. Copyright restrictions may apply. Image courtesy of Biodiversity Heritage Library. http://biodiversitylibrary.org/page/553107 PHYLOGENETIC ANALYSIS OF J(lfI/(>S F. Srnitli / ·H IV l ohn Krc:}s,3 and Z 3 THE ZING IBERALES BASED I:'/izabeth A. Zimmer • ON rbeL SEQUENCES' AnSTRAcr Morphological data have bet:n used previously to construct phylogenies of the eight families of the Zingiberales, one of the most wi dely accepted monophyletic groups of nowcring plants. To provide additional suppon for phylogenetic relationships within Ihe order, and placement of the order among monacots. we present a parsimony analysis of DNA sequences from the chloroplast-encoded gene, rbe L. for 21 species of Zingiberales and proposed relatives. Five analyses with equal, and differential weights were performed. All analyses resulted in the salllC most parsimonious tree for taxa within the Zingiberales and the irnmediate outgroup. The closest sister group to the Zingiberales based on these data is a clade containing Commelinaceae/ Haemodoraceae/ Pontederiaceae. The tree topology within the order based on r/)('l sequence data is different from previous morphological analyses. The order can be divided into two sister groups, one containing the Costaceae and Marantaceae, and the other, the remaining six families . All recognized families are monophyletic with the exception of the Musaceae, which is paraphyletic with the Cannaceae. With trees one and two steps longer than the most parsimonious trees, phylogenetic resolution is rapidly lost. suggesting that the phylogenetic utility of rbe l sequence data for the Zingiberales is limited to interordinal and intrafamilial relationships. T he Zi ngi berales. a morphologicall y di stinctive classifica tion, including delim it ation and rank of oreler of monocots. arc one of the most widely ac­ these families, has been subject to many changes cepted monophyletic groups of pl ants (Bentham & (reviewed in Kress. 1990). Cladistic analyses of Il ooker. 1883: Petersen. 1889: Schumann. 1900, morphological charal:1ers ha ve greatly improved 1902. 1904: Hutchinson, 1934, 1959, 1973: Na­ the understanding of ph ylogenetit: relat ionships of kai, 1941: Tomlinson, 1962, 1969: Stebbin :-s, 197 k the families (Dahlgren & Rasmussen, 1983: Cronquist, 1978, 1981: Dahlgren & HasllIu ssen, Kress, 1990). Dahlgren & Rasmussen (1983) per­ 1983; Dahlgren et aI. , 198.'): Kress, 1990). Dahl­ fon ned the fir st cladisti c analysis of the Zi ngiberales gren el aL (1985) li sted six apomorphies for the using the eight families listed above and polarized Zi ngiberales: root hair cells short er than other epi­ <.: hara <.: ters using their Commeliniflorae. This anal­ dermal cells. sieve tube plastids containing starch, ysis resulted in a single tree (Fig. I) composed of presence of silica bodies, epigynous fl owers, lack three main clades that included the gi nger group of distincti ve apert ures on the pollen grains. ami (Z ingiberaceae/ Costaceae and rVl arantaceae/ Can­ the occurrence of arill ate seeds. In addition_ the naceac). the banana group (Musaceae/ Heliconi­ herbaceous arborescent stem, distichous phyllo ­ aceae). and the bird -of- paradi se group (Strclit zia­ taxy. large pet iolate leaves wi th blades possessing ceae i Lowiaecae). The relationships among the three transverse ve nation, conspicuolls colorfu l hral:1eate groups remained equi vocal. in fl orescences. and the substit ut ion of olle to five Kress (1990) re-a nalyzed th e data of Dahlgren staminodia for the fer til e stamens a re characters & Ra smussen ( 1983) and performed a separate easil y lI sed to ident ify members of the Zi ngiberales analys is that included ot her characters which we re (Kress, \990). rooted wit h th e Brorne li ales, T his second analysis As currently classified, the order consists of eight resulted in a different cladogram (Fig. 2) from that fami lies (K ress. 1990): Musaceae. Low iaceae. I-Iel­ of Dahlgren & Hastnlt ssen ( 1983). Although the iconi aceae. Strelitziaceac, Zi ngibcraceae, Costa­ ginger group relationship wa s retai ned in bot h. the ceae, Marantaceae, and Cannaceae. SuLordi nal fam il ies of the banana and bird-o f-paradise groups , We thank YOllngbae Sli h for expert advice on cloning and sequencing. We acknowledge the Scholarly Studies Program of the Smithsonian Institution fo r providing fundi ng and a grant to WJK, the staff of the US Botall ical Carden for cure of li villg coll ectiuns, and G. Zurawski (DNAX) for providing primer sequences, , l aooratory of Molecular Systematics, Museum Support Center, Smithsonian Insti tution, Washington, D.C. 20560, U.S.A. ' Department of Bota ny N HB· 166, Smithsollian Illst itution, \Va shington, D. C. 20560, USA. , Current address: Department of Biology, Boise State University. Boise. Idaho 83725, U.s.A. A NN. MISSO UIl' BOT. GAllI). 80: 620- 630. 1993. Volume 80. Number 3 Smith et al. 621 1993 Phylogenetic Analysis of Zingiberales • • • • • • •v • •v v < • V • • • • <• c • • ,- • v • v • •w ~ •w - • < • •0 •, • w •v < -~ v• • < •c - - < • - • •c u• • 0 - , v 0• U ,• • , v • -- ~ " " "• " • f lt :1 liE [. Cladogram of the Zill~ibl'ralcs frollt Dahlgren & Ra"lIIu!'o"clI ([ 983. their figu re 9 ). we re show n to he paraphylc tit' (K n·:-:--. 19 90). T hese I-I a rbo rne . 1977 ). T hol"ll(' ( 1992). !J(I\\t. \ t·r. 1'1','­ C1 na]vscs• have heen highv l•\' ill forlna tiq' ill t(,rIl lS of ognized thrce .'- eparalt" o rder" ill hi ... "'lIl" 'rord('r f.. tuilia l relationships. vel the ut il i!\' of m a ll \' o f the Cornrnclin<ln<le : Brorneiia l(':-. COIl llllt'lirlait· .... and • • • luorphological c ha racter;; at th i:- It'vel is q u('sli on+ Z ingihera Ie- !-t. alIII..'. The ('ig ht fami li es of tht· Zingi hcralcs a rc Alt c rnali \,,·I\". \\ a lk e r ( 1987) t1 criH'd hi ... I.in- • highl y d i~ ' i ll (' l i\' ( ' wi th man )' uniq ue a nd highl y gi beridae (excluding the Ihorllt: lial(':-) ;lIltl It i:- POll ­ modified morphological ~ tr uctll rt ':-. Ill l e f p r cta l ion tcde rii dae ( Il ueillodoracc<lc. I)OIll t·t!t·riir,·t·irt'. alltl of homology between famjli(' ~ ha ... posed di nicult Phil yd ra t:cae) d irectly frolll a liliid l i rlt'a ~ ( '. ~l']JaI problc ilis without deta iled de\'doprllL'lI tal a nalysi" nectaries. ve... :-cl ... prirnaril~ in 11)(' root .... and "'('\ n al for ma n)' of the structures (Ki rdlofL 1991 ). c he mical ciwra(·ter:' (e.g ._ I"Ill·lidonit' iwid) fOlllld The Il Ulllcro us aU lapo rnor phi t· ...; u r th e Z ingi bc r ­ ill the I.ingilwralcs s u pport tllt 'ir pla t"t"IIH"ll t \\ ilh a ales ha ve a l ~o Iliade diffie nll lilt' dt'l nminalio ll o f lili a lcall l i n ca~(' (T a khlajan. 1980). Ihe evolu lionary relalio nship o f tilt' o rder 10 o lhe r T he li se of molccular c har<H'tcr" ill diUli ... li\" <lnal­ lIIo nocols_ The Zing iberalc!> 1...1\ t' gCllerally been yscs has !tn'n hig hl y Stu·c(' ...... ful in plall t... (1'':IIIIU'r allied wilh the Bromeliales and or COlllmelinalcs e t a l. 1988: Crawford. 19(0). pa r t il"lilarl ~ illla \ ­ ( Il u ic hinson . 19i3: Dahlgrell ('I al .. 1985: T horne. ollolllica ll y diflinllt group" ill Idlich it i ... hard 10 1992). T he p re:o;.e ll ce of s tarc h y (·nduspe nn. cpi. intc rp ret morphological Ir o illolog~ (c./-! .. Cha ... t· & I"lltie ular wax or the Sln'lil:.ifl type. L· \'-fluores· Palmer. 1989: Smilh. 1991). H l' t'l'll ll ~ . ..; t·q lll·rw(' ("(! lIt o rganic M'ids in th(' ("{,II \\all--. and Iwo or fou r da ta derived frolll the- chloropia ... t gt'lH' ,.Iw l .. "hich :-i ub:,id ia r y cell::. in the S\Ollia lal ("oili plex a re a ll ('nc()( I (· ~ the large- s ubu nit o r ri bulose "i ~ l' ll u :-o pll<l l (' s pet:ia lized c ha rac te rs Iha l ullitt' I Ill' Zi llg iberiflo rae carhoxyla s('. has provid NI I"\'solu tio ll a l highn lax ­ wilh the IJ roll lc1i ifl orae and COll ltl lt·lillifl orae ( Da hl­ onomic k" cI ~ in planls (e.g., DOl' hlt,y , '1 a l. 1990: gren e l a l.. 1985). Mos t m Od C1'I1 t·la ...;si fi l"alioll .'; ha ve Donoghue l't al. 199 2: Olrtls tead t'I ;d .. 1992: placed Ihe Zingiberales Iwar tilt" Bromcliales (COII ­ o lhe r pape rs this issue). Thl' I"\ 'ialivt'ly :-o l!)\\' ra t(' of taill ing the single ramily Brollldiaecae) (l iulc hi ll' 1l11llation o f th e- riu'L gell (' ha :-o lIIadt' 1111 :-0 gl'lle :'011. 1973: Siebbi n :> . 197 k T akhlajan_ 1980: usc rul for taxonolllit: iln t'"ligatiun:-o al alld abo\"{' C ronquist. 19i5. 1981) based on the sim ilarit ), o f Ihe gell eri(' 1('\"t·1. 1I 0\\e\l'r. Ihl' li mi t:- ,IIHI taxo­ in fl orescencc a nd flowe r sl n H·turt·" (primaril y the nomic range ro r wh ic h rlw l .