The ^ • A-"..* the BOTANICAL REVIEW
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THE BOTANICAL REVIEW Founded in 1935 HENRY ALLAN GLEASON AND EDMUND H. FULLING Published by THE NEW YORK BOTANICAL GARDEN Editor THE BOTANICAL REVIEW DENNIS WM. STEVENSON VOL. 64 JANUARY-MARCH 1998 Project Editor No. l JOY E. RUNYON Articles arc obtained primarily by invitation, but unsolicited manuscripts are also consid- ered. All manuscripts are subjectto critical review before acceptance, but it is to be expected Phylogenetics and Character Evolution in the Grass Family that most solicited manuscripts will, in fact, be acceptable. In all cases, the Editors reserve (Poaceae): Simultaneous Analysis of Morphological and the right to accept, reject, or suggest revisions in submitted material. Instructions to contribu- Chloroplast DNA Restriction Site Character Sets tors appear in issue 1 of the current volume. Copies may be obtained from the Editor. Articles submitted for publication as well as any editorial questions should be addressed ROBERT J. SORENG1 to: The Editor, The Botanical Review, The New York Botanical Garden, Bronx N. Y. 10458 USA. " ' AND The. function of The Botanical Review is to present syntheses of the state of knowledge and understanding of individual segments of botany. It is to be hoped that these syntheses can JERROLD I. DAVIS be read and understood by other botanists less well informed than the authors. Photographs electron micrographs, and drawings may be included as necessary. Enough papers should be L. H. Bailey Hortorium cited to document the information and interpretation presented, to help the reader learn how 462 Mann Library we arrived at the present state of understanding, and to demonstrate that the author is thor- Cornell University oughly familiar with the literature in his or her field. New information can and sometimes Ithaca, NY 14853 should be incorporated into the articles, but The Botanical Review is not the place for articles primarily concerned with presenting a limited set of new data. I. Abstract II. Introduction " ' III. Methods 3 S»b>eripdon price (1998 rate.), $82.00 U.S.; 589.00 elsewhere. Subscriptions entered on a cal- A. Taxon Sample endar year basis (Jan-Dec) for the current volume unless otherwise specified. Cancellations B. Chloroplast DNA Restriction Site Characters' I 1 (U-S currcncy drawn on a u s bank should be mad BOTANI- C. Structural Characters SfmSTr - - - ) = <° .D. Data Analysis ' U IV. Results I2 Claims for missing issues must be made within one year of date of issue. Non-U.S. deliveries at A. Colinearity of the Chloroplast Genomes.' j^ subScnbersrisk.Non-U.S.subscribersmaypurchaserePlacementcopieSata20%discount B. Characteristics of the Data Matrix-. .].[ ., ^TlttZlSend~?pl"60Idandnew«ldress«'°^Sei tif,cPublieaiionsD pan- C. Cladistic Analyses ment. Allow eight weeks for change to become effective en e 1. Chloroplast DNA Restriction Site Data.'....". ]l Back issues and volumes are available from the Scientific Publications Department. 2. Structural Character Data 3. Simultaneous Analysis ofRestrictionSi'te'and Structural Characters 21 4. Topological Similarities among Unweighted Analyses ->< 5. Incongruence among Unweighted Analyses ,, BOTANICAL.REVIEW (ISSN: 0006-8101) is published quarterly forS82.00per year forUS 6. Implied Weighting .' ,'' V Discussion ""' : • ^2 TEHW?••; rSUge P cd.St Br0nXl NY lnd ldditional mailin« oflfce,. POSTMAS- SS SSEISKTpublioations Depanment-The ^ • a-"..* A. Scoring of Structural Characters andUse of Exemplar' Taxa! \\\\'.'.'.'.'.'.'.'.'.\'. 34 DCPW Na£i nal MUSeUm fNatUraI Hist0 © 1998, by The New York Botanical Garden. • 5SSSl ' ° ° * a"*"""-" >^tion, Composition by Sarah MacLennan Kerr, Ph.D., Editorial and Production Services Printed by Allen Press, Inc., Lawrence, Kansas 66044, U.S.A. Library of Congress Catalog Card Number Agr 36-330 rev 4 Copies of this issue [64(1)] may be purchased from the Scientific Pub- (ISSN 0006-8101) The New Y rk Botanical Garden Bro 1U45S-5125Mss°7i« ?mUSA. PleasePT' inquire as° to prices. - -- ^ THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER. The Botanical Review M(l): 1-45, Juiaiy-Much 1998 O 1998 Tot New York Botuicil Guden THE BOTANICAL, REVIEW PHYLOGENETICS AND CHARACTER EVOLUTION IN POACEAE 7 B. Analyses of Molecular Data Sets I C. Phylogenetic Structure of the Grasses.' 35 Nardus, Olyra, Pharus, and ftrep/ocAaeto) also were included with the <•i of *•, • • 1. Support for Groups Resolved by fte Present'Analysis' It the,r phylogenetic affinities. The two character sets wer T^LTsZ^LT, 7" 2. Companion of Results of Simultaneous Analysis with Previous Classifications and Hypotheses ofPhylogcny of the Grasses neous analysis of the combined matrices also was conductedcombTdda a < , D. Structural Character Evolution in the Grasses 3S analyzed using homoplasy-implied weights. Among mJr^S? the com iS T 1. Spikelets ; 43 weighted analyse wcreresolution of a sister-grotm relationship between ^indPoa" 2. Lodicules 43 3. Stamen Number " '• ' 46 4. Stigmatic Branching ,'' 47 5. Fruit Characteristics : 47 6. First Seedling Leaf 4S 7. Cross-Venation ' 49 8. Microhairs SO 9. Arm Cells 50 10. Fusoid Cells 51 11. Photosynthetic Pathway 51 12. Chromosome Numbers. 5' E. Synapomorphies of Various Groupings of'Ehrhanoid'eac'. Poo'ideae " Brachyelytrum, and the PACC Clade . late mcludiST f^"ftWea<^ Agrostideae [i.e., Aveneae], and Poeae, the F. Placement of Five Problematic Taxa within Poo'ideae <? latter including Seslena) are resolved as a monophyletic group that is nested amon* th, ~ 53 CJ. Taxonomic Conclusions main.ng elements of the subfamily (Brachypodieae, MelLe" Stip aTnloZ* Dar 54 H. Breeding Systems in Grasses .... •^W •f'rf^AS£COndPrinCipdS°a!°f*e-alysiswastoidZfy^cmr; 1. Evolution of Wind Pollination in Grasses 55 ynapomorphies of clades. Among the synapomorphies identified for some Xe So 2. Evolution of Self-Incompatibility in Grasses " I. Biogeography So VI. Acknowledgments S7 VII. Literature Cited 59 VIII. Appendix 1: Restriction Site Character Data 59 IX. Appendix 2: Structural Characters and States 67 X. Appendix 3: Structural Character Data 74 W»«-«B.»d»^ •«1, ^i• ta fc, d.de tot comprises Ceo" toSdSe I. Abstract veT,•^ A phylogenetic analysis of the grass family (Poaceae) was conducted ••• * i. sets, one representing variation in 364 mapped and clad slal v inforSlv S ^^ from all regions of the chloroplast genome L •J~Z' mf0miatlVe reStnctlon sites "structural characters " The strucmral ,>,'",!,> ' P f"tmg Vanatlon m'42 ^formative chrotnosotnal, and S^S^^^^S^^^ •'-I, II. Introduction The man simple comprises 7S e*e•i«;•vY • TT t, the ehlor°Plas' genome. In recent years agrostologists have generated several explicit phylogenetic hypotheses vita);n*«a*.M^^J^Z^T.-}**•*"* '»d">'- oncemmg major d.verstfication patterns in the grass family (Poaceae). fn c«wta 5 haVC bCen baS6d n 1987,7° ttr^XT^rKellogg & Campbell, 1987); on chloroplast DNA °(cpDNA) -^logical restriction chaTac sites e, (Davis 5a m& X£: HaLov * 7•hltB ^ a,!o i"3): °n nUCle°tide Se^=nC£S of nuclear nSm^t f^A2 Hamby & Z«•«. 1988,B 1992), genes of the chloroplast genome (Doebley eVal 1,1994; Nad0tcta1 ' & MonT^ZV T '. IS^Barkeretal., 1995; LkJS^tln &. Morton, 1996; Hongpmg L.ang & Hilu, 1996), and the nuclear gene Alcohol-dehvdT S; ^TJLt "}%* and,°n T°US C°mbinati0nS °f *«* ^-acter set iSS •ithhto..ffenpeudothers. Sever.lgeder,ofeonw^SllSL^S?1"',?' "••" Under, 1995) Most of ftese analyses have yielded results consistent with the recognition of two major Cades m the family, one of them corresponding to subfamily PooideaeZ second compnsmg subfamihes Panicoideae, Arundinoideae, Chloridoidcae, and CentotheE coJIecnvely referred to as the "PACC clade" (Davis & Soreng, 1993). However, there have THE BOTANICAL REVIEW I'HYLOGENETICS AND CHARACTER EVOLUTION IN POACEAE j subfamilies of their own (e.g., Anomochlooideae Potztal StreDtochaetoirf«, rr u « uu , Butzin, Pharoideae L. G. Clark & Judz. Parianoideae rc E Sim , n [ V?1 Ubb'] 1 0ryZ ldeae hartoideae Link if »*,*, is iactadcflfSSSSL^ ^ ° ^ ' potheses of r^^X^SS^- ^^SS^SfT^ ^ * ! dci 1957 I962 Another complicating factor encountered by grass systematise ;.fh,t••M • _• , . Tateoka, 1957; Prat, 1960; Stebbins & C^nJ iwi n • ,« " "' ' i fications of the grasses are intuitive and idiosyncm cto! nf lsthatl"°^t existing classi- gree f an then Be0ause the ford eta,., 1969; Sharma, i979 ;cl mT£'&SSB ^ "^ '^ "* cs family is so large groups once reL^S * ° ° ton & Renvoize, 1986; Soderstrom & Ellis lMrlsTi ,•i,: atS°n et aL' 1985= Clay- £ be cohere & objective analyses of date to d^onstraX" ' t'?• 'T^ ° "^ithout 1992), some of which f^S^^^i^^^ **« *«*£ neticanalysesofgrassdatareSZt3tSZ^^Z^""^?* as later detected by cladistic analyses AlmTehXr * and *= PACC dade' family (Hilu & Wright 1982- Watson et .1 108< f ? haVe Provided Programs of the these treatments, Ldflc^oa ^J^StSS^T ^T^,,a^ f two ma or clades, aside from the disposition of tax" assSed to R K "j " ° ^ J ^-aandgenerio•^^ y m cult t0 lace pelodesmos Link, Anisopogon R Br Danthn»i•*, £ , V. i • P are ^m- A* M^Conert, NaJsl, P^danttS^Tc^£T'^ ^ <" Stipeae. These so-called orphan genera and trib ° , i, , ' S,ePhanachn° Keng, and either in Pooidea terminals of analysis (e.g., Hilu SlS^ ST^^^S^ T Arundinoideae (compare thTckssSon of S• S" " ^ = " reviewed therein; Kellogg & Campbell, 1987; Kellogg & Wateon 1 w A ' " approach is that all putative groups of interes can bf rfn^v ? }" advantaSe in ^ H te^*^ approach involves theoretical .5 SSSSfe r^ °T'" phyly of groups, creation of nonexistent characte «tei7r• v • ^^Pt1011 of mon°- based studies