ANNALS OF THE MISSOURI BOTANICAL GARDEN VOLUME 81 1994 Colophon APS Bodoni. The text is set in 9 point typ g 10# i! Vintage Gloss. This is an acid-free paper sig to have a d shelf: life of over 100 Gloss is manufactured by the Potlatch Paper Comp lft үе oe used in 8 р, ү оер T } а s а : лл The ANNALS is printed and distributed by Allen Press, Inc. of Lawrence, Kansas 66044, U.S.A 1 Garden 1994 © Mi ISSN 0026-6493 Annals | of the — Missouri Botanical Garden umber . ‚ їп the back of the last issue of each volume. Volume 84, Number 4 Annals of the Winter 1994 Missouri Botanical Garden E The Annals, published quarterly, contains papers, primarily in systematic botany, con- / tributed from the Missouri Botanical Garden, St. Louis. Papers originating outside the Garden will also be accepted. Authors should write the Managing Editor for information ` concerning arrangements for publishing in the ANNALS. Instructions to Authors are printed | РР: М”. 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The journal Novon is included paid at St. Louis, MO and additional mailing offices: in the дейсе price of the ANNALS. POSTMASTER: Send address coe to eer, on ARDEN, Department Eleven, P.O. Box 299, St. Louis, Hio: 63166- on. (O Missouri Botanical Garden 1994 | © Thi te th n 4, f ANCI/NICA 720 AR_4009 ғр ч €t D, r) кы ч Volume 81 Number 1 1994 Annals of the Missouri Botanical Garden КД MI, [| "31 1994 DEN LIBR SYSTEMATICS OF THE EUPHORBIACEAE: INTRODUCTION! Grady L. Webster e first International Conference on the Sys- tematics of the үөс organized by Mi- chael Huft and Grady Webster, was held at the Missouri Botanical Garden in St. Louis on 14-16 August 1989. Papers were presented by 21 people ain: of these with one or more collaborators); f these, 15 authors submitted papers that were oen for publication and are presented in this issue and the one following. Mo the St. Louis conference was techni- ca e first international symposium on the Eu- EDU it ha d been pr ЧУН by ше {ушр Royal Botanic E Kew, in as can be s that symposium (Jury et al., 1987, Bot. J. Linn. Soc. 94:1-326), the scope of the Kew m was different; there was much more emphasis che шы апа medical applications, and. most ч 1986. However, f eke of the Euphorbiaceae with other an- Я famili e St. Loui sium, in contrast, was “‘in- ner-directed,” vh De primarily on е» of diversity within the Euphorbiaceae. How considerable attention was given to uc ph i E а with other families in the papers by Vogel et al n эы matics of storage ‘proteins, agar о and by Sel gle er on comparative biochemistry. Ta bet at the s symposium was the für ther elaborati ion on, and perhaps progress toward resolution of, ques- tions about the **membership" of certai bind Mithin. ше Euppertiacess. Pa exam studies by ), Kap nd oo apnd di ESER and Levin phology) indicated that controversial genera ‘a such as Antidesma, Bischofia, an nested within Euphorbiaceae, ar recognition of separate families poen eae, Bischofiaceae, and ymenocardiaceae would render the Euphorbi- aceae paraphyletic. А significant part of the symposium contribu- at the level of sub- n, Levin pso e, pollen m phology), Acalyphoideae tribe Plukenetieae (Gil- Жыр pollen morphology), Euphorbioideae (Men- ne wood anatomy, not published here), Енот (Gilbert; and Koutnik, not published). the generic level, there were four studies, on ' The International Conference on the ane of the Euphorbiaceae was supported this issue of the Annals and the one sep, 15 mcs {то eviews of Michael 4 qae Geoffrey A. Levin, and Grady S.A Foundation Grant BSR- 061799 m The pape ? Section of Plant Biology, Beaty of California by National Soc vis, California 95616, U. ANN. Missouri Bor. Garp. 81- 1-9 199A Annals of the Missouri Botanical Garden Phyllanthus с, = КОК Jatropha (Dehgan & Schutzman), d a (Carter), and Dale- фо ia (A p ie collectivelv. indicate of cladistic the bd х, intra- or intergeneric re- lationships. The p y Armbruster—with its елдери ion of pollination relationships, cladistic af- that Wee is a 51 шаса t interest in the Euphor- with symp positae, Legumino: 0 and Orch х8) the Spy eens sul unexplored terri шогу. It is be that there was no compar- udy of DNA or R DE it is apparent c seed ае proteins. pu а эеле 15 жен клр е жор a model for the kind of f studies that lê be done wis of the taxa in the 1 ге regret ча їп publication of tm iu m, we believe family. obsolescence by suggesting pi ios the profitable directions for research in the CLASSIFICATION OF THE EUPHORBIACEAE! Grady L. Webster? ABSTRACT ү f. T he family Euphorbiaceae The display a great variety of growth forms, aa at least 17 “models” of Hallé. Anatomical pean particula rly useful for classification includ pran a m pse паш thia Ж, evolved several tim crit haract tribes ^ e for a e wood structure, laticifer type, trichomes, and stomata es. Pollen nw dr Sub nu. erede of the seed c h f the fam Sd E ly a а dispersal oe ants is importa t in many taxa. primitive taxa in Africa an Madagascar, ораза 2 DETTA nuclear y. Pollination i is t prevaligy entomophilous, and 5 on. Benth ted. гей е orbus uter appear to reflec tha id ntham’s Types of an Old World origin of the у (Cretaceous and Paleogene) f long-distance depo r but rather trivial i inst 4 so r bar and s in the late Tertiary tog ال ا‎ i high: a س‎ via the Bering land bridge appear to have been relatively insignifica es Euphorbiaceae, Vinum one hs the ANIN ics, have been relatively neglected by systematists in the 20th centu oth = n E Б g : 8. 8 family and its Constituent infrafamilial taxa is the major focus of attention e history of classification systems for the Eu- абы at the subfamilial and tribal level has been reviewed in de Ke e the ru series of genera сая 7 muah jator in э, ш: Миейег ( 1866), who provided. the fust tive about relationships, and the later (1880) and Pax (1890), who uninforma ones of Bentham . La & Ho man no (1291) ona ihe Nie by visas u^ E ы. 0 ueller. i Alt g my recent classification (Webster, 1975) a appears ery different from that of Mueller at first glance, stru ith se apt 5 the tribes divided into is con- s ies, wi su eu The classification presented at th ference shows sear a limitea number of aluages, арш апа subtribes. The originality ы Mueller is is is show that substantial fomes іван - ve be made ear Particularly System of Baillon (1858), sil was cisci кош of taxa in a classification is an im- ' I wish to thank Robert Rhode were provided through the ore of Daniel Axe т his assistance in Y ge gebe of the manuscript and illustrations. Outline maps * Section of Plant Biology, University of California, Davis, California 95616, U.S.A. ANN. MissounRi Bor. GARD. 81: 3-32. 1994. 4 Annals of the Missouri Botanical Garden perfect way of indicating рона оь Ixonanthaceae are accepted as possible “ош shi ips. Cladistic analysis of the family hypotheses that in the classification cheme. Unfortunately, the information available for the Euphorbiac imperf: d this, alon with ips size oi the gtoupi- make it аш йо proceed congress in 1987 SNC I | provided some mily t nd s intended to set forth this monea in greaten detail, in de hope that it will In doing so, 1 emphasise that, while our classification system Mu eller i in t an of the a ne is indebted to iis s dem ма "io Gon ly + + prove ibs ‘almost sivit adig in its е of later pend on the biogeography of major angio- sperm RELATIONSHIPS BETWEEN THE SUBFAMILIES ince the p мү discussion of Bentham (1878), it has чач accepted s later нева (е. Рах, 1924), that the subfamily Ph Пано dis is ilie are within the aai Wielandieae, nme eywoodia and Savia, appear t orig- inal N E complex that arose probably in the = ee: Those primitive Eupho fbi: forest habitats in tropical latitudes. They have, as expected, dei d HMM пении wit vs 1 Ж г 1 staminate flower m iss pisos 2x has a 3-locular ovary, with two anatropous ovul a large emb ce the ete dons. ain far as the evidence goes, the chrome sve edes 3 in th archaic" Phyllanthoideae. An- atomically, rus sy pd ideae have such unepecialisid features as vessel slemen ts with sca- nd e @ Ф чі nonglandular trichomes. Lotes ap- to be absent. I f Geranialean families such as groups," as speculated earlier (Webster, 1967), then it is feasible to compile a list of characters in the р for which plesiomorphic (prim- зех ee Mt eres A=. А | (Table Bos An important (ош to Мона eigo llan tł Yoidea has пва made by ren (19860) i in his cladistic Pollen eod. on ка subfamily, cx still un pu wena (Pun & Sim promise o ate din structing phyletic m in our knowledge, for T is in n biochemical dab Although освен г (1962- f f. ү | X) ү IT I this information is frustrating to the systematist, because it derno ons Бп atoe a dis biochemical di- versity in the t at the same time reveals a nu bet erratic level of sampling. As a орехи і ical dat till of limited use fulnes £3 z МОРЕ "= and ipit leve ls. 1987 review of the classification of the eagues. Hymenocardiaceae are accepted as a family by Léonard & Mosango (1985) and by Radcliffe-Smith (1987). Furthermore, Radcliffe-Smith accepts Pandacea d retains three genera ( desma, ischofia, and Uapaca) only provisionally within ien he Euphorbiaceae, Jensen (1994) s sts that gical data there appear to two main groups of Euphorbiaceae: Phyllanthoi- deae + Oldfieldioideae; and Ac ho | Е bioi roposed е Mahlberg (1987) on the basis of laticifer morphology. On a phenetic asis, this basic dichotomy чаан сая rea- sonable, but it remains to be seen whether it be substantiated by critical pibadi studies. se (199 0) went muc he rt nine families wi ing restricted mainly to the т 5 ilies a RET Ri and Euphor- bioideae. There is not sufficient space here to analyze rec — Volume 81, Number 1 Webster 5 1994 Classification of the i Euphorbiaceae TABLE l. List of primitive (plesi phic) and derived (aj phic) ch in the Euphorbiaceae. Character Primitive state Derived state l. Habit Trees/shrubs Herbs/vines 2. Branching Monopodial Sympodial 3. Phyllotaxy Alternate Opposite 4. Leaf shape Simple Lobed; compound 5. Leaf venation Pinnate Palmate 6. Stip Absent de Vessel perforation Scalariform Simple 8. Vascular rays ultiseriate Uniseriate 9. Internal phloem t resent 10. Latici sent Present 11. Trichomes Simple Stellate; lepidote 12. Foliar glands Absent resent 13. Inflorescence Axillary Terminal 14. Calyx aestivation Imbricate Valvate; reduplicate 15. Petals Present; free Absent; connate 16 Present Absen: 17. Stamen number 0 1-4; over 10 18. Filaments e onna 19. Anther dehiscence Longitudinal (vertical) Horizontal 20. Pollen nucle 2-nucle nu 21. Pollen exin Semitectate Tectate; intectate 22. Pollen apertures Colpi ores; inaperturate 3. Apert mber 3 4 or more 24. Pistill Present ^as 25. Carpel number 3-5 26 or m 26. Style branches Bifid Une шша 27. Style union Free 28. Ovule numbe 2/locule me 29. Ovule жарга Anatropous — ороп 30. 2 Monosporic E EE 31. Dehiscent Indehisc du Hid Ecarunculate arunculate 33. See y Fles 34. Present Scanty or absent 35. Cotyledo = han 2 EESE Less than 2 36. Cotyledon/ d width 2+ is provocative arguments, but I seram that he ual tribes or subfamilies, particular morphological has overemphasized the seed-coat data of Corner characters have proven to especially useful. So (1976) that the case for ponent fthe far only p orph s provided micro- emains unpersuasive. Clearly, though, the morphological characters that are useful in deter- monophyly e Euphorbiaceae as a whole, an mining systematic affinitie all fi bfamilies, f hes nd subfamilies, needs be as orks of Erdtman (1952 shown by w Punt (1962, 1987). However, the studies of Levin demonstrated in the course of any ; ровде; Ь, с) demonstrate. "mw foliar venation pro- vU ingly дем ш of the infrafamilial tax ily Phyllanthoideae, this may o be true for other char s well. The REVIEW OF SYSTEMATIC CRITERIA HS nt, the overwhelming preponderance of тоѕоте кеи Bs "we asa) i nft alan At pre һе the data iiti ble for d the patterns of these may well уте clues to de ines of sys- phylogeny within the — ae comes from tematic affinity, but at present on „йн data gross vegetative and flora idu charac- аге frastratingly seas it Biochemical data from ters that have been used as анасон taxonomic parati dies of dary pound h features. In evaluating relationships within individ- Ikaloids and tery hardly fail to produce Annals oe Pu E Garden results of great interest, especially at the tribal and generic levels, resent we have only tan- à ар а эжең scarcely more than a In f the Euphorbiaceae proposed рге, PRO: the magn criteria were obser уаше Eob e т oJ wath th ked d + ] J © 1 characters by Erdman a (1952) the systems s Hu- 3 : on classification of f the family, m seems nidis уе : ting evolutionary trends and lines of “аши арен a are not cussed, exce dispersal a le significant effect on em evolution of reproductive structure À. GROWTH FORM e Eu diete display an extraordinary range ge growth f which show a wide adaptations to relatively mesic and xeric habitats, Êr In ars the nee of =з oe їгеез at has been. pie on a compar e basis sis, or the first tim bewild diis in tropical taxa can be comprehended beni em scheme of about two Pos Pasie m odels proposed & Oldeman, duy 1978 Determination o y Hallé, et sla E kh nz Ol 1 X | r cL. system is difficult бой examination of herbarium specimens; nent lings or ы may be required. Despite the can- studied, th li f tribes is sun incomplete. However, further studies may p g phylogeny of suprageneric- taxa t the present time, no less than 17 models have been reported for the woody Euphorbiaceae; and a significant number of herbaceous, scandent, and 1 Е. ЕРЕ 77) } 33: 1 dol: All of the кошш өлер me abe Ош: llé that cannot be easily assigned to any of the Hallé and there appear to be diaaa betwee of the маснав that тау Бе о signific In bani carni branchin terns tend to be monopodial and axillary; the ERE ip Te patter those shown by the mode 15 of At Hone, Aube il ih. A Mm podial plagiotropic branching (Roux's model) as in ueg, Phyllanthu pecialized р йош m, ching " by Webster (1956), me penal (1. e 1), Roux (19 ad Lom & бышы (19 subfamily SEP ао Phyllanthoideae i in a having relati e roa th the mode i of Attims "te ‘Ra uh. Нус Stachys- a few other taxa, appear to fit the del of a in which extension of the main axis is interrupted by a ter els of Koriba and beurre in which the main axis is terminated b d branch- ore or less dichasial in nature. In the е d model of Koriba seems to be the most common, and that of Rauh is abo alent as that of Attims. Finally, in the Euphor- riis of Koriba а owth patte The vegetative and reproductive morphology oa Mairie iic noit de ing pat- inflorescences rns are le, dM ERE es Volume 81, Number 1 1994 Webst Cassifeation the Euphorbiac pad nee has received a i dien "d of болани апа а Шш сап е ‘traved и до — Euphorbieae. It is particularly of Anthostema is referred to the model BENZ by Hallé & Oldeman (1970); the ае іп that genus recapitulates the origin of the cyathium { + Uer Paar r ТЕРИ? 1 1 with the abortive axis surmounted by a pistillate flower. Nozeran (1953) had i EHRE applied the term *pre-cyathium" to the ual spikes o nea such as Sapium i voali чуге) К on a e nt growing ac 1 f Noz zeran n could i indeed liie g I i al though there are still pro oblems—in the absence of any "missing link"—in specifying how the ictu ا‎ basal flower in the Hippomanean thyrse becomes the terminal flower in the cyathium (see Gilbert, 1994). One argue that the ultimate point in veg- etative керн in the Euphorbiaceae is attained in Cham jeg Kati ee "t ‘seedling axis is Mil н — "n is virtually all sympodial. In the — of Nozeran, but the subsequent dichasial pattern t gs E Б © gressing through subgenus Agaloma by retardation of the mai i lati he lateral anthocladia J "1 J (1ORR)\ch * | that the lateral branclis-i in Chamaesyce arise не the cotyledonary nodes, and that th mology with the terminal pleiochasial s in Eu- Phorbia is doubtful. The adaptive significance of the growth forms in the Euphorbiaceae is not as clear as their чав as systematic characters (а common situatio many angiosperm taxa). Part of the diffic za is e ^e бнкы ез in some тшш párteme m are model. Cremers (1977) seesaw 10 ee M orbia, and there ap- Baillon and E. abyssinica Gmel., which conform to the models of Atti and Rauh, have surely be ived from ancestral taxa with the mperfect Min i sie каз тариад — eap interact with omplicated fashion. Cert tain models, — as ie itm and Nozeran's, clearly predominate i in m те ҳач oth- ers, such as Attims’s, are f ests and deserts. Evidently the Euphorbiaceae өнен tered such a wide variety of habitats during ie radiation eli no та — can be ex ected. Neve онай ере герг ченее ctive structures in some taxa of Euphorbiaceae is so striking that m more the architectural models. B. ANATOMY rene ont adve rud —_ Ered as since the essay of Pax (1884), which first estab- ed the h or de g natural g (art meri vs. nonarticulated), ph and trichom a as the major ака char- t his redivision of the major su- studi extended in the following decade by Radlkofer and s students. Radlkofer (1870) emphasized ana tomical characters in a paper establishing the ne uphorbiac genus Pausandra; and a series of is students surveyed most of the family anatom- ically (Rittershausen, 1892; е 1896; dauscher, 1896; Herbert, 1897). The first really critical anatomical survey «t the family was provided by OES Ey who reviewed the activity, so that reversals of trends and conver- 0 wn by of Troll's ate ries the Ba poricee in the Phyl- lanthoideae, it поа t peus and негњфеббоц Рат wh rived from Attims’s model; bet in the Бабына. deae, it seems to ien evolved in Pedilanthus via Attims's model and in Actinostemon via Prévost's observations of his own. Gaucher (1902) published пото d his obse ns. Howeve ve the merit of providing эч eae — arranged by tr w y Pax id diri; Annals of the Missouri Botanical Garden up to the turn of the century, seem mainly to hav honnen ma айо) оп tig majai al. ‘The first important wood fr om h 1 РТ же! 90 1930), bead аем оп Indonesian taxa. Jansson- Da m as an indi relationship to the Euphorbiaceae. The most in- teresting of his findings was his designation o group of era of subfamily Phyllanthoideae (Aporosa, Baccaurea, Drypetes, and Putranjiva) as characterized by thick-walled nonseptate fibers, ala the work on fiber пор ology by showing th thick-w alled 1 fibers (“уре IT”) occur in genera of subfamily Онбеш (шина апа Ре- aime vanes and showed the presence of type II fibers in a ticifers i in the single genus Jatropha: single-celled respects contradic findings o & ya (1964). viet (1975, Mahlberg et 7), in a series м ара showed the sys- Eu phorbia Mu | Ж раг- ticularly with respect to the type of starch grains produced. Оша ге R uda Conty however, han types within the family and a ply demonstrated the importance of laticifers i in ыы relation- 1987) interpreted this e data radically i própneing.a subdi- das (1980) reviewed the wood anatom amily Pic. as part of a BHO И ар Де surv Metcalfe & Chalk (1950), in ee most recent companhona anatomical 1 review of there famil y J correspond to accepted: subfamilial and tribal con- — Hayden à ntire 1 Their “Group A” of the Phyllanthoidese i incl cters first vessel silvia. It appears that “Group A" taxa h as tribes Wiela: ane eae and Antiderntae iat the mos 1 1 ыкы Bois cal (and Janss = with the thoideae; but this disposition is contradicted чыш 5 ch other evi dence th semblances of Acalypha to the Phyllanthoideae must be due entirely to conver- gence. The studies of Mennega (1987) on w natomy in the Phyllanthoidea general sup- hgan & Craig (1978) reported a variety of la- on the groups Phyllanthoideae, Acalyphineae, and Hippomanoineae gi Бал (1 884); Clearly there Е still unio ovulate пане ог mor- нант d anatomical dat far ilies. ilie subsequent HEEE of hos char- y the fa ct that the last detailed systematic review is that of more compre- ningful. A hensive sampleja 250 species by Rao Raju (1985) э He Mz ail af E R | M. Баг AR e JE ан f ТЕ: E a "m fam ilies Acalyphoideae and Crotonoideae, where var- ious kinds о malpighiaceous, stellate, and ue types sconto in the other three subfamilies simple unic siete multicellular hairs overwhelmingly predomin. The то T trichomes ned in the ne phorbiaceae are stinging hairs, ean rain in the нгө але ae and Crotonoi- deae. The hairs in Cnidoscolus (Crotonoid Urti Volume 81, Number 1 199 Webst ышын of the Euphorbiaceae ican €: of COS ems dis кй hoidea 1 + auterent type are em own vn from the tribe Plukenetieae. These ). The possession of this unique ty ype “i trichome (along with other characters such compou "€ iini неу а powerful argument for Sepe lechampia, a separate "s o the Plukenetieae. The Buphorbiaceae are also impressively rich in foliar glands, which appear to have arisen de novo within t ® @ Ф Ро o B > t B d and а Idioideae, foliar glands are restricte occurrence, but they are very common in the uni- lat hf. n rs n Le | 1 А | ve been reported in the unrelated genera Clutia үн a Suregada Cna are wi cu pheae, Alchorneae, Bernar rse; glands on leaf ften appear to Nt with petiolar glands. Bernh niy et бири fallen ard (1966), in a survey of the pee ands, has teeth h in шапу siofia a separate origin from petiola nds. Belin-Depoux & Clair-Maczulajtys (1974, 1975) saraaa the an- tomi of Aleurites ‘Delis: орах of Alchornea Euphorbiaceae may b th dominance of paracytic stomata in the woody taxa не most of the 17 tribes sampled, with а oyi Plana tui indicated рч for Phyl lan- y Webster, 1956). The berrant sit- Mies oar by Raju & us is the high per- centage of anomocytic stomata in Chamae Not unexpectedly, Kakkar es) Е Е * ge ez m fo e two subgenera of Jatropha can be distinguished by stomatal type and also surface of hairs (sm s verruculose). ooth versu: Foliar venation, a neglected character, has only mone ко used by Levin (1986a b, o1 fo шо Phylanthoideae. Similar studies on the other sub: ni ould no do ubt as illuminating So far, done on tribe Eu hee bites. (Sehgal & Paliwal, 1974). The characteristic jacketed venation in Cham aesyce ome attracted interest because of its correlation with C, photosynthesis (Webster et al., 75). Although the vast majority of Phyllanthoideae and Euphorbioideae have simple entire leaves, the aaa ae and Crotonoideae have predomi- itely: ini e or shed pm оу & Wolle in “Dilleniid” ‘leaf type on the basis of actinod- romous venation and *'Violoid" marginal teeth, in ich the medial vein expands into a glandula termination without a Euphorbiaceae a are variable; for example, decidu- setae occur in the Hippoman eae, which would nite them “‘Theoid” instead of **Violoi venation and tooth t of Euphorbiaceae betray a "'Dilleniid" instead of “Rosid d" affinity. Vascular барт of petioles has been studied y Dehay (1935), who showed — vari шой in stelar H rate u p (1982) found that different degrees of petiolar ste- ; > d ructure in many Euphor- pules ы at the base of the biace: aide (Ublarz, 1978). In many of the Phyllan- Annals ambe A IMS Garden thoideae and occasional genera (e. g^ Manihot) else аке ee — are s (ca- ducous) .g., Ну stipules, whereas they are absent in related species. Many "aig of Croton lack ае сике they re large a Most notably, Eu ШО. or absence of ee is a orbia diagnostic character f sitis role in protecting ш ор pis lea di primor la en for their presence or то, in closely related tax, C. INFLORESCENCES AND FLOWERS Gis, F. Be 1 = 1g 3 are often re educed, grouping of flowers into inflo- rescences is often im character ny instances. Monoecious and di- oecious flower production are widespread, with onoecious perhap ewhat more com t judging from sexual dispositions in *basal" taxa (e.g., Wielandieae) in the subfamily Phyllanthoi- deae. However, this seems to have been readil t een since monoecious taxa appear to have Croton, Euphorbia, nera. Inflorescences as h Pinus e е a whole are pro mop phily nt): Hyme r ge nera n re- ioar see (Crotonoideae). Wind- polinated | taxa the flowers in axillary clusters are produc pce The гк өг ойеп — pene as pike, ommon in all five s ubfamilies. In diu карнай there i duction series from the diffuse pseudanthial structure, with eine e athial genera apparently evolved from a ancestor. Floral form in the Euphorbiaceae shows tre- "— БЫ ЫГЫ indie asa плен псе = in- ers Pad partly as the result of adaptation to a variety of vectors cialists on t & е s work there has been only one other comprehensive survey, kata Rao that of Michaelis : (1924) However, Ven tly (1971, 1972) studied vascular anat- nsiderable num of a co ingle inst. Tae nen нен the fevers of tein ause vitri are often her- y the ii ammo ed maphroditic; his i | 1 a Rao pointed ой ats of trends in most obvious being that of reduction in size an n r of parts. Reduction is sometimes correlated with shift to wind jon, as in е рһеае; phorbieae) it clearly is Rao, there are also tre s to increase the number of cuius nim here аши! the ere is no conan usually entomophilous It seems apparent that inflorescence SUES in the h ta have elaborate compound dichasia. H the aS, it can usually be seen that mens, for ex cur in i GFE (entomophi- lous) and pin (inopia A particularly evident seduction клу is ji 1095 of the Mm кка . placed in “apet talous” groups. This affects V — ңе» is most о for нн т flowers; pet talous pistil- at y = РА Volume 81, Number 1 1994 late flowers occur in most species of Croton and in other Crotonoideae, as well as many Acaly- phoideae (e.g., Agrostistach ydeae, Chrozopho- eae). Two а. Oldfieldioideae (except Croizatia) and E orbioideae, are apet both sexes. In the presumably pollin Wines — Androstachys and nali flow inate fl in these genera are modified inflo. rescences. The calyx in DR AP d varies greatly in rt configuration and number of parts. Where the co- rolla is absent, pero calyces may occur, as in Cellos and al ipu (1943) sug: that the hix had been lost; however, the aac from vascular traces in Manihot c cate or ractice initiate (1866). меме Baillon (1873) дану eee: to the excessive weight accorded the c by Mueller, it was still extensively used by Pax (1890) and Pax & 1931). Co aren, of the evidence from other characters (wood anatomy осе and йч strongly vith en euphorbiaceous аш where present is hi te variab flow r mainly in the кайган ai чаў g., in hse ites, dicen. The petals are usually dis- ti b i ct, but m e coherent or te in so Crotonoideae such tropha ( Web- ster, 1979) and some Aleuritideae (Léonard, 1962) Ve amalakshmi (1968) reported that vascularization of petals (with three bundles) is usua r to that of the sim although it may 1 е | 1 1 1 TAE gm با‎ and n. The floral disk (nectary) is a His feature of many euphorbiaceous flow n ( x & Hoffmann, 1931). On the other hand, кы, (1924) proposed cone ма disk is piet staminodial origin, partly bees of he anna as гче and р artly ‘ebuild of the stami- nodelike EA of disk-segments in various such as Chiropetalum, Clutia, and Por- nther а. However, Venkata Rao & Ramalakshmi Webst ni à the Euphorbiacea (1968) have shown that the vascularization of the disk is is highly variable; and Webster (1 ct ЕЛ — EU = ith mass; small disk-se ents are usually nonvascularized. The prepon- erance of evidence against the view lis and in fav the classical concept tacular nature and not an independent whorl of the flower The eup абнои androecium displays а pro- tribes, Michaelis nad Вси that the sia condition i in > family was ап androecium of a indefinite f stamens in many whorls. lilio kato Rao & € 1(1968), in contrast, sug- gested that the primitive condition is 10 stamens in two whorls, as in Jatropha. yin r, on the basis of the prevalent condition: in the Wielandiese I think that an Se a of a single whorl of five or six sta likely to be ancestral within the fa hod Моне ш. in defining subtribes, placed reat е on the insertion of the stam practice, cin usually depended on whether they were inserted around a NOM or d disk; Hoyer in some groups such as t yP to distinguish a from a eres central disk; and in Crotonoideae such as Cnidoscolus or Jatropha, it is not easy to tell scr ^e slender projections at the и. of the — column rep- resent staminodes or a pistillod Mueller was ou ia by Baillon (1873) for placing undue emphasis on anther form and de ens as E characters. Because of trans mens; in es difficult to ver- ty There are all degre “of hif me the рев of anther dehiscence, not only between extrorse and intro orse en bet pre pince sitse d lon- l. The мане: dee Lasi ion are found in the A sio- — seri es am ان‎ of the stamens are ed into b ing fascicles and the number of anther peer. —— the giis „e degree labora a fi lat; к I © 1 1 ; гр F n. he role of telomes in the dicc flower, as pointed out by van der Pijl (1952). Venkata Rao & Ramalakshmi (1968) made the fantastic sug- ion that the branched lex of Rici- о Е 3 nus is primitive and that conventional dithecous Annals of the vinti Botanical Garden anthers are derived by reduction; abnormal numerarily lobe super- 111p ICI! t кый тог olga cia is needed, it apie ars that Airy Shaw may have b monoia us were derived independently ibus those in Rici Pollen grains Bii the Euphorbiaceae are produced in tetrads by simultaneous cytoki naw te trinucleate pollen occur. In a study of the tri Euphorbieae, Webster, Rupert & Kout (1982) howed th cleate pollen originated several times and t he distribution of poll ucle important tium in defining pul i Pagi ce suppor the *'Schürhoff- "ыен biet ndi qo that the shift from binucleate to Меге е pollen is йен The morphology of is grains in Eupho aceae is so diverse that only the f ien diram of Brazilian phology fining tax E at any ve Pa x (1884) 1: d systematic utility lin Radlkof ао th pollen n despite йер рарег fer (1870) тананы the pollen char- bie Pausan me We owe to Erdtman (1952) hology offers decisive clues to s systematic rela- hi within the Euphorbiaceae. Erdtm est in Hoffmann (1931) has been adipe. by Punt (1962) and later workers. Köh- r (1965 ed ight microscopic studies and by unpublished SEM observations (Webster et al., un- published). The importance of palynological char- acters is further highlighted by the studies in this issue on Oldfieldioideae, Crotonoideae, and tribe — great — were ma in Euphorbiaceae makes it difficult presumed to partial exception; although t tation is reduced, th colpi. Within all of the subfamilies except the Eu- phertéoiioéo, there isa tengenty си порно grains; in the Crotonoideae, the hS we obsolete. Crotonoid sexinous ornamentation is а — e synapomorphy n subfamily Crotono- ae. Tr киш his BEEBE wi ERE within the subfamilies uphorbiaceae i is е igh " r is three i in га; эру e taxa, it is i eiii to two in a number enera and to ne in several, includin. irp (in s Cro- tonopsis, Drypete es үм "хайыр к Jatrop . On other han phorbi- ac "e since wit un ists Wielandieae add are ue five indi М Маггагйагїа pies number i uctuates between two and six, and varies from € to six in the single species M. nobilis L.f. iren 1979). Stylar variation is marked a often for specific i 0 — the two style branches may be further subdivid $0 that each style m may have four or eight stigmas, n in bizarre shapes. Finally, number of genera (e.g., Endospermum, Glo = ч د ن‎ ud Fes SpA ا ایو‎ ure rm y UA if Volume 81, Number 1 1994 Webster Classification of the Euphorbiaceae = ышы ы л g RERO а chidion, Tetrorchidium) the styles are reduced to sessile, often petaloid, stigmas. Rather une 2 : шор have either highly dissected ч натен ог undivided entire бе (Al Ovular ch н ауе ee e as ден nificant within тр м Sealy енын the distinction de- as There i is no eins of reversion гош esumed d , uniovulate, to the home edis deer it remaing unclear whet 1 ith DlIOVULaT ia with шош locules oc- curred only once. In other words, we cannot yet i uniovulate taxa, treated by Pax (1890) as чыту Crotonoideae, represent а monophyletic ж ut by E Baillon (1858), ovules of = & a Iporusa. sive oliin lans ers (e. B^ ма Phy llanthus) hav turators. Schweiger (1905) aca the variation ovular it m in thie Биро and fout phorbia, and tetrasporic types in Acalypha, Eu- phorbia, and Mallotus. A characteristic of most EHR female is the early di ft ld а їп репега mixis is rare їп enema ap азы even though the berto report o of a apogamy is is fro in tem bi 1 cis L. (Carano, 1926; Cesca, 1961; Kapil, 1961). D. FRUIT AND SEEDS ruit in most Euphorbiaceae is an explo- sie dehisc ent schizocarp, but indehiscent sed have armen Picton in all of the subfamilies h state, both in the ‚л аз ш within individual genera. In Pianta (Webster, 19507 1958) there is a pan larly pholog ical series: typical c 1 t sections) to bac- cate fruits (sect. Ani nema); кеш but de- а uence does use a transition within a single phyletic superg: (1975 b has provided the most one ana tomy 0 n some тл especially t tribe Phyllan- theae, the ovules are dist ation in 77 ovule is hecha dines to m» parenta - — a distinct fun via illustrated si йна речовин " d b families Crotonoideae and Acal ахы respec- tively; in the Crotonoideae the ovule s elongated and has a nucellar beak projecting be- I the micropyle, whereas in the Ac Ferien ^ ovule is less elongated with a thickened inner integument and without a projecting nucellus. Ovules in the Euphorbioideae conform to those of the Acalyphoideae Considerable ied _— work has been done on the mie copus (Davis, 1966; Rao, pies showin о sac develop ent is of t Кит most of та fam x t has feta only in the Агайды, яз e disporic (Allium) types occur in Chrozophora and Eu- ogy of - — euphorbiaceous fruit as r sented in ralian genus Micrantheum. Berg related ma аи we roam features of the eu- Новасовь pistillate flower, such as the vascular in the colu vv to ada dum dehiscence an its tip." However, as shown in the че asi and illustrations of Venkata Rao & malakshmi (1968), the three basal central ена іп the axis each one © to о paired аган, that sii ai the 1 lies |! Б ump. 1 з + ph h E is typically axile, and Berg’s ingenious hypothesis is unnecessary. Another provocative suggestion of Berg is that the primary function of the obturator is not to facilitate passage of the pollen tube but rather to displace the seed lower into the coccus for more Annals of the Missouri Botanical Garden effective seed dispersal. Against this view may be - ned great — і in obturator dimensions although it is pos- that size omis could be correlated wit Iruitea the biovulate taxa, there are typically two seeds ule the number i EN. in Pm Meineckia, and Savia). In orie taxa with ia tien fruits, ы опе ог tw в б б Aporus и с: varies greatly, HE less than 1 Chamaesyce or a a to ov d Omp Ма (926), Wunderlich (1968), and Cor- ne t th e anatomica struc- + faeh А | "A £ in relating the орносо is eu families, as well ships. The testa ев the outer integument) рег- es +h j * | $ а |I 3 e.g., in Baccaurea, Mallotus, Mar- ium, and ‘Tetror © +h subfamilies differ from the darin in sometimes having tegmic vascular bundles, this character i is inconsistent: hod ner's EOS nd 3 families, and group 2 contains both Acalyphoideae rotono Although “ч ee is pe no "— a ubiq- Raillon рашоп (1873) — Mueller а «ыр it as a gone ch 1aracter d in terest. Bresinsky (1963) and Berg 5a, b ehe 0 the caruncle always arises from the region (exostome) of the outer integume - horbia, Micrantheum, and doubtless the other gener: h carunculate seeds, the carunc - tions аз an elaiosome to attract ants for dispersal Berg noted that in Micrantheum it may also play the build-up of tension within the devel- oping capsule, leading up to the explosive dehis- cence. E 3 z - 8 Ф б Е 3 . B 5 3 c = from the ouen gaira of ihe: чын. oer integumen t), uniovulate and biovulste taxa. In the Phyl- aoe, the mechanical cells consist of cuboid only slightly dece quic саар cells or e tangentially elongated fibers ntrast, in the uni iovulate tax de mec Mica “layer is rend 1 f£ at 2) t frankly admitted that the picture is not crystal-clear. n my opinion, the primitive Bises in th o the аи ough de шай lat d pete ч ү pae е А. е Phyllant in that subfamily er are sometimes miero- аня outgrowths that could be regarded as ient caruncles). However, their di i riage A is ee ا‎ related gen while in various ‚ genera (e.g. Euphorbia, sa y no means all) оГ peres eed character that appears particularly im- ortant in the Phyllanthoideae is the presence or renis of endosperm, which develops as the Nu- clear Pid (Rao, 1970). Itis copious ir in most genera, but sc of the spottiness in repo uch as an Amanoa, Spondianthus, and Wielandi exalbuminous seeds, the cotyledons are often distinctly folded. Out the Phyllan- osperm, but there are a few exceptions (e.g ateriospermum in the Crotonoideae Cotyledon characters ng tional weight in the classification of Euphorbiaceae since t hey were strongly emphasized by Mueller т — ` Volume 81, Number 1 Webst 15 1994 EE of the Euphorbiaceae (1866), = ee his € pane M number o сш Lad 3. 23:78. 10, with other numbers originating in a isang aneuploid aoe: ‘Jones & Smith to this о overemphasis on the “ү "e аан and his rejection of Mueller’s scheme has been ы e 2 the pa нет studies of Erdt- 19 Punt (1962 куе ae with intain t relation between cotyledon shape, ericoid habit, and Australasian distribution. As a matter of fact, it is still n ar why the character of narrow cotyled b ost con us tralia, except for the anomalous North American enus Reverchonia (Webster & Miller, 1963). E. CYTOLOGICAL CHARACTERS Perry isi was the first to provide a surv of Sification. Hans c d there a f damental basic numbers amily: x — eu gen ‚ of which the latter is A sone 5 desse B da D © = Dn ya 8 m’s (1880) tribal arrangement, and unfor- ely the worst mistake in B treatment S his inclusion of the Buxaceae (with x = 7) a опе of the tri Hans ano is probably correct in ii ioi that x = 11 is the original bas mber in the Aca alyphoideae and Fo need The same con- clusion c $ö Opinion, the evidence clearly suggests an original g y of basic num bers i in Mona denium: x = 12716; the lowest number of x = 12 occurs in species. M «омак Stapf) rank eur as E ) s LE | OA PS PEPS PE | n are of polyploid origin from an ancestor with x 9, followed by a mostly decreasing aneuploid series. Gaede all of the oytologically hub n po- tential succ the case remains not proven SUMMARY OF TAXONOMIC CHARACTERS IN THE EUPHORBIACEAE At = m о опе ен po êê and cladistic methóds has ред made. lovin yP (1986b, c) classified th hyl- lanthoideae pri ийре basis of leaf anatom- ical data ladistic i of Oldfeldioideae based = 8 0 T e 1 atomical, ate b гечи cal (пої to mention т ular) work needs to be done before the didis set к adequate. In Table 1 are listed a the eh situation хавь е that панок ted by Ehrenfeld 1 (1976) collected 175 species T ies, and wasps. us HBK a xican d C. suberos n the role of floral nectaries in Croton 22 Doct 1 > 2: Ls "^ z and his collaborators have not yet demon e benefit from visits of oO © S а E] as 2. © =] л = E I] n peded ICE in floral ua Логове received even —— ورا‎ is жоо number of instances of m for pollination rons rag нка ted, апа по oubt many remain to be detected. Pollination by i ally unspeciali re- a Picks хе onde as Hauman suggested, a similar syn- ome may occur in other taxa ч tribe йы maneae, but there are few In contrast to this, however, Warmke с dem- onstrated nat a z ification of this is desirable, it appears that Hevea mily for is o be the c tilobus (Pax gni umbers of у by taxa of Heliconidae, Pieridae, and Hee zil situation. in Cnidoscolus is similar, ar Je a visits to Cnidosco / РР P Aiscct pollination. In my opinion, th ts for wind lh Ч ЕР e^ : Z 1 n © reve MC: 1 г m: = morphology, are not entirely persuasive; Domin- lus niger da were seen by hummingbirds (Webster, летна ве, urn-shaped corolla and the RAA o E Volume 81, Number 1 Webster 17 1994 Classification of the Euphorbiaceae listi lelphous) to have bracts are usually glandless. In contrast, the c ved in this one speci es concomitan tly with a sm from lepidopteran to ornithophilous pollina- ion. Pollination by small Hymenoptera is doubtless very widespread in the Euphorbiaceae. In Califor- nia, wasps were f be the most important pollinators for Croton and Euphorbia by Moldenke (1976). In ey pollination of Tragia. uhi tiana EARN ona (Webster & — ae tion by small wasps and bees may Дер то elucidate кошо of the variations in eu- h EF rer I mysterious. s presumably are major pollinators most species produces resin for a floral re (Webster & Webster, 1972). Armbruster & Web- one species, Dalech of Webster & Armbr ampia magnistipulata ected by female o er cham visits b = чаны, and euglossine ees The cons h ts. Armbruster (1992) showed a fascinating correlation betes Rhylogeny of taxa in Dale- € convincingly demonstrated that че i by fra grance- Bios om ing euglossine bees has evolved ently in three хаа species lineages of pseudan thial i in- Dalechampia may m the thyrse of Acalypheae, in which the floral been reviewed by Dressler (1957) and Webster (1967). Other pseudanthia, of i БУ. m of Euphorbia and Cham syce is к. Piae by small generalist id ga as a og nis e стен р P ааа: ше Ашы ш а "m su аен buriningbird pollination. з. meder (1957) clearly illustrated how the more o dially symmetrical cyathium of Euphorbia has been high ified into the bilaterally symmet- rical cyathium of Pedilanthus that is pollinated mainly by hummingbirds. At least fragmentary accounts of most major s lination of a occidentalis Benth. by bats (Gloss порае СагоШа) and the red woolly pos- sum (Caluro гуз), шоп visit the inilorescnnnes, fo 1 obtain nectar . Ап even more remarkable instance of bat minate and pistillate me inflorescence, in Hura the fleshy staminate 1 41 k satil a me 7 е 12а lination ie Steine segete that the а ““сопез” аге 1а istaking them for fleshy аа са ase of , a ae. examples indicate that the great diversity hl a | : +] + +] These £f 1 related toa iong гу of shifting adaptation to gre flow ers However, becaus ег ч ini r асеае are less than ЄТ? di- ameter, the effective unit of pollination has ба been the inflorescence ог s florescence. Func- scenc tional studies of evolution of reproductive struc- es must therefore take into account the in Annals of the Missouri Botanical Garden TABLE 2. Taxonomic distribution of disseminule types in the Euphorbiaceae А mper of genera have not been scored due эрт oldfieldioid dpa non: is men- which dispersal of the этине оп attraction ч ants to = ree Num- Fruit ber Seeds Seeds inde of carun- - Testa s- Subfamily genera culate culate fleshy cent Phyllanthoideae 58 0 38 5 12 Oldfieldioideae 28 14 0 e Acalyphoideae 118 — 48 XU [ssi Ws 5 Crotonoide: 29 3 8 Euphorbioideae 42 16 18 1 4 Тота 3071 БА 170 24 31 rescence as a whole, as well as its constituent parts. Furthermore, as discussed below, adaptati dispersal may also have a “ in proje 8 developmental restraints back о the flow 2. DISPERSAL The prevalent fruit type in € кие seni E: Es (Di n 1 LP \ + > J is t Sera schizo- carp that dehisces year ii into three segments (cocci) each wit a dry in iis sub- r fle e spicuous elaiosome (caruncle) fou ilies; elty (in- d seed mor- f fruit an phology, Berg's (1975b) died in on the Aus- TaBLE 3. Relation of dispersal type to e онар distribution of genera: carunculate seeds аге treated as myrmecochorous; fleshy ар we indehiscent fruits as ornithochorous. Many g ble, are probably Cred s Myrmeco- Ornitho- chorous chorous Neo 18 10 үүчү, Madagascar 8 21 Tropical А: 6 28 ARM 21 6 Holarctic 4 0 Totals 57 65 Ber g э stud 1 f Mic — seeds with a careful anatomical study of fruit and seed comparative data M у " era of M ao Ld кер Since a very rudimentary elaiosome occurs in e Phyllan- thoideae (e.g., Phyllan duped it is not used ruit—in- cluding the reduced ovule ышты: et e dthe RÊ т ialize derive from an ancestral state of dee osive capsular fruits, and that morphological changes M | m А 14 1h ] d from the fruit back into the flower. Berg also sug- ested, somewhat less iilii ig that this char- смеа tic euphorbiaceous gyno ium/ fruit/seed character syndrome ar opical monsoon climate; this could be so, but there are many taxa with typ = explosive fruits in "si i rain- forest enera arê badly needed in ет to test cie d s hypotheses me of t n the adap- ive aspec gar dispersal ar are sit ee in Tables 2-4. It is dis from Table 2 that ‘“‘unspecialized” sete (i.e., ancien ита dry seed coat) pre- minate in all subfamilies. However, in the Olin ind d Варнаве there are near- as ma ecarunculate genera. The Kishi аа s dii БЫН pe of the four subfamilies with Yon seg idis the largest number of we with fleshy Geo opti (Table 3), there are some inter- esting aei nces in n" dispersal type, since +h ї пу е путева genera ‘the: n ornithocho- st of the кн агеа us, whereas in the r the situation is reve si n view of the interest tropical ecologists have taken i in = Dess of dioecy asa ере пуе Me of tn and seeds with the monoecious verdi D Y e Givnish ie and Steiner (1988) that wis taxa should have a higher percentage of ornitho- Volume 81, Number 1 Webst 19 1994 Овенот т {һе Euphorbiac li n do monoecious taxa. In TABLE 4. Relation of sexual expression to nt contrast, Prae "Pian to be no significant differ- are presumabl Although Berg is suggesting that the Sutpehorous Piccopine fruits r KPIS primitiv ve state l developed su. D- sequently, further evolution to ignei more or less indehiscent fruits has all of the sub- fa milis: es. hn the Phyllanthoidea [иза аге qe js birds: examples UNGE Б їп pakos and es in Drypetes and Uapaca. In Phyllanthus (Webster, 4956), thexe is a particularly marked Isocladus) to berries (subgenus Kirganelia) and finally to y, drupaceou sed by specialist frugivores, but we have ver little data on Euphorbiaceae. A stri elop- nt in lyphoideae and Crotonoideae is th evolution of large oily seeds in ge 5 s Caryodendron, Hevea, Joannesia, Aleurites, and Omphalea. 'The seeds of Hevea are ipse أ‎ dispersed to gu dintapees Мир to 45m gcording to van der Pijl, 1 rivers in the Amazon basin. Hevea ang these other Е seeded taxa аге mostly polis is a liana), but another group of gene uch as Homonoia, rheophytes that are adapted to extensive peri s of su ably depen preyed upon. Wind dispersal is uphorbi- aceae and occurs ma in taxa with samaroi fruits adapted for open tropical woodland (e.g.. Hymenocardia); there are no good examples of winged seeds in t e Euphorbiac eae Bresinsky (1963), on the ben. of studies of cm iem s that seed =“; in temperate x more complex. Primitively, seeds in the Euphor- уре monoec /D, monoecious or dioecious different eames of a genus); D, consistently dioecious. Seeds Seeds/ arun- fruits Totals Seeds culate flesh for dry (myrme-(ornith- each (auto- cocho- ocho- sub- Subfamily chorous) rous) rous) family Phyllanthoideae M 191 0 8 14 M/D 5 0 1 6 D 22 0 9 SI Oldfieldioideae M 1 3 0 4 M/D 1 1 0 2 р 12 7 3 22 Acalyphoideae 44 4 6 54 M/D 3 0 T 4 D 30 14 11 55 Crotonoideae 15 13 4 32 M/D 0 2 0 2 D 14 4 9 27 Euphorbioideae M 14 13 0 27 M/D 2 2 1 р 2 1 4 7 Totals M 85 33 13 131 M/D Lr 5 19 р 80 26 36 142 176 64 52 292 biaceae appear to haye been E pa ld. lanthoideae. Ht is учтен» that carunculate seeds ос аха of the Oldfieldioideae (Tetra- co ideae (Clutia), and Crotonoideae (Mier re M Seeds with sarcotesta are relati e a lar- пс a) in Phy atively "basal genera in ig iie eae (Cheilosa) nt hu Klainea hus) n appears that diaspore evolution has followed a n many еа between dry/ tortuous path, w езһу ап y isa general tendency lor pes with Sarooteste ы testa. There occur u caruncules to occur in open dry habitats evident, dr d is there is no unilinear direction 20 Annals of the | selene Botanical Garden I . Summary of diversity in subfamilies and tribes of Euphorbiaceae as measured by numbers of native and endemic genera ee emic genera indicated in parentheses). Tropical Arabia is counted as part of Africa, Pacific islands with Australasia Subfamily America Africa Madagascar Asia Australasia Phyllanthoideae Wielandieae 4 (3) 4 (3) 3 (2) 1 (0) 1 (0) Amanoeae 1 (0) 2(1) Bridelieae 2 (0) 2 (0) 2 (0) 2 (0) Phyllantheae 8 (3) 14 (4) 6 (1) 10 (1) 7 (0) Drypeteae 1 (0) 3 (2) 1 (0) 2(1) 1 (0) Antidesmeae 6 (5) 7 (4) 4 (1) 4 (1) 5 (2) Hymenocardieae GY) 1 (0) 1 (0) Віѕсһобеае 1 (0) 1 (0) Totals 21 (12) 33 (14) 16 (4) 21 (3) 17 (2) Oldfieldioideae Croizateae TC) Podocalyceae 3 (3) Caletieae 1 (1) 2(1) 13 (12) Picrodendreae 4 (4) 4 (3) 3 (2) 1 (1) Totals 8 (8) 5 (4) 3 (2) 3 (2) 13 (12) Acalyphoideae Clutieae 1(1) Pogonophoreae 1 (0) 1 (0) Chaetocarpeae 1 (0) 1(0) 1 (0) 2(1) 1(1) Сһе 2(1) 1(0) улашы. 22) 1 (0) Dicoelieae 1(1) Galearieae 2(1) 2 (0) 1 (0) ae 2 (2) d Agrostistachydeae 2(2 2(1) 1 (0) } phor 5 (4) 2(0 6 (4) 1 (0) i Caryodendreae 2(2) 1(1 Bernardieae 2 (2) 242 1 (1) Pycnocomeae 2(1 2(1) 4 (2) 2 (0) Epiprineae 2(1 2 (0) 8 (6) elieae 4 (4) 1(1 Alchorneae 5 (4) 1 (0) 3 (2) (0 2 Acalypheae 3(1) 11 (6 10 (5) 14 (4) 11(3) Plukenetieae 11 (8) 3 3(1 5 (3) leae (0) 1(0 1 (0) 1 (0) 1 (0) Totals 36 (26) 33 (17) 22 (9) 52 (26) 23 (7) Crotonoideae icrandreae 4 (4) Manihoteae 2 (2) Adenoclineae 3 (2) 3 (2) 1(0 onieae 1 (0) 1 (0) 1 (0 1 (0) Elateriospermeae 1(1 Jatropheae 3 (2) 2(1) 3 (2 Codiaeae 4 (4) 1 (0) 8 (5 5 (3) Trigonostemoneae 1(0 1 (0) cino ae 1(1 6 (6) Crotoneae 4 (3) 20) 1(0) 2(1) 1 (0) Ricinodendreae 2(1) 1 (0) 1 (0) Aleuritideae 2 (2) 7 (6) 3 (2) 4 (3) 0) Totals 22 (19) 18 (11) 6 (2) 23 (13) 15 (9) Volume 81, Number 1 Webster 21 1994 Classification of the Euphorbiaceae TABLE 5. Continued. Subfamily America Africa Madagascar Asia Australasia Euphorbioideae Stomatocalyceae (1) 2 (2) 1 (0) 1 (0) Hippomaneae 14 (10) 4 (2) 1 (0) 5 (1) 4 (0) Pachystromateae (1) Hureae 4 (4) Euphorbieae 4 (2) 7 (4) 3 (0) 2 (0) 4 (2) otals 24 (18) 13 (8) 4 (0) 8(1) 9 (2) Grand totals 111 (83) 102 (54) 51 (17) 107 (45) 77 (32) Їз of the subfamilies some of the most striking disjunct or relict distri- except 8 the уйе which lack eit butions. There do not appe be any critically mecochory, and the Oldfieldioide on erate documented maps of suprageneric taxa in the lit А The a s and 3) is oc cal deer Australian regions mai k enn ro ame chem or- J with the pect opit nithochorous. P. and the Айз dos redominane ce “of myrme- ean he dispersal in Australia, confirming the statements of Berg (1975a, b, 1981). ittle has ias studied so far on the effects of seod predation on dispersal systems in Euphorbi- e He ЕНЕ со туча: e such genera e p. ic- F but th y in Africa, qium Дк) fenem udi t be indicate lower levels of pre- = ыы PERS T is Bde: cent at ай іп vid have been nalyses besi distribution У, ома ае by Рах (1910), а tially) of = Euphorbieae by Croizat (1972) and Leach (1976). A review of the geographic distribution of the tribes and genera of Siti sae соме brings out the striking promin f Africa and deber d ри 5). The highly disjunct der 1); including the related genus Heywoodia, the diversity in Africa/Madagascar is clearly greater an the New Work he occurrence of чё relict genus Wielandia r and t in Madagasca e Sey- chelles is pha тн with the x fn Pei 22. and Petalodiscus, it indicates a ma- jor c in Malagasia (Fig. 2). Discocarpus and Lachnostylis crane ота а уісагіапі per in So d Afr: са кү ofS vica South rican/ African dis junétià n is ies wit ilies, as noted in Table 5. It informati ve to look “ the Бэрш of the more It is notable that in nthe s Oldfeldioideae the Dasal tribes Croiza- 1 (Fig inus, patterns. 3. GEOGRAPHIC DISTRIBUTION A ii ным of the e geographic niam patte bey бы the scope of this essay. TEn it seem appropriate here to outline some of the ог geo- бары: patterns of di five Loblanifis Vid indica te 1eotropical a е the Caleti tieae are айыу Old World, nly the Picrodendreae (Fig. 4) are represented c еу : may represent the taxon in which the important Annals of th Missouri ied Garden PHYLLANTHOIDEAE Wielandieae . Distribution of Phyllanthoideae, tribe Wielandieae: 1, Heywoodia (dotted line, Africa); 2, Savia Wobei line); 6, Actephila (dashed line); 9, Chonocentrum (dotted line, America). apomorphy of carunculate seeds first appeared in the family. In Te "enlyphoidese; there is one relict neo- onophora, one of east anl southern prog Со, опе neotropical and Asiatic, Chaetocarpus, and three Asiatic, Chei- losa, Neoscortechinia, and Trigonopleura (Figs fri о be the epicenter z узе hier that it vulat pouer basal йа, sch as | Cheiloseae and Chae The Crotonoideae (Figs. 7, 8) differ from all ‘the other subfamilies in a clear ndron, He- vea, Micrandra, Micrandronps. and Tetrorchi- dium. Ноев ma qe i Africa, and A nd К. Te E The. ре therefore show con- n pn 0 occurs in , except that the с center of gravity is кше to the west of the Atlantic rather than y, th primitive Euphorbioideae К ya Stomatocalyceae (Fig. 9) give an ambiguous picture, with one neo- ` Distributions are based on references cited in Webster (1994) bet examination of herbariu um spec imens, E marily at DAV. G ster (1994). Distributions are rece Rea лу ind eralized and do not show minor gaps. tropical genus, Nealchornea, two African, Ham- ilc tr ibe Euphorbieae (Fig. 10) has a distributional with tribe Anthosteminae їп in Africa/ Madagascar, the Neoguillauminii led c by Raven & Axelr - (1974 ), deployment of a significant number of tribes o ау ауе taken place before Graco plate tecto onic a of the family into till limited, but the faci that fossils ot he re A atively “advanced” аа, from the Eocene (oed & Daglian, 1982, Dilch t „Мадоне, хара) suggests that the ori gin a go Danks into ci Cretaceous, or the габий 13 tiary. For e the pollen records from Aus- tra hus discus ed T Martin (1974, 1 978) үн се that relatively “advanced” genera of Oldfieldioi- Wein 1а plac e in Australia in the Paleocene, er. ЧЗ Tx hfamily well PR into the корон. It asy to summ. the overall distri- ова ү ано of Me Ennis, put I am str uck Volume 81, Number 1 Webster 1994 Classification of the Euphorbiaceae PHYLLANTHOIDEAE Wielandieae FIGURE 2. Distribution of Phyllanthoideae, tribe Wielandieae ( in d) 3, Gonatogyne, (dotted line, Brazil); 4, 5, Blotia, Petalodiscus (dotted line, adagascar); 7, Di ‘pus (d America); 8. line, Africa); 10, Wielandia (solid line). by Bentham (1878), who presented the first model Order was in the Old World, but that several of for the geographic patterns involved in the origin the principal forms differentiated end widely h l1 ry L *, p J + and radiation of horbiaceae. Bentham’s sp p p words deserve to be quoted: *... we may be led impassible barriers opposed by the Atlantic and ; AE ЛАД eU d : to conjecture that the most ancient home of the Pacific did ‚ or in some OLDFIELDIOIDEAE Croizateae Podocalyceae ананда : — EBT io =з ЖЕЛЕУ. Baer E. Idioid Tribe Croizateae: 61, Croizatia. Tribe Podocalyceae: 62, Podocalyx; FIGURE 35° йн f Olds 63, Tetracoccus; 64, Paradrypetes. 24 Annals of the Missouri Botanical Garden OLDFIELDIOIDEAE Picrodendreae _ FIGURE 4.° Distributions of Oldfieldioideae, tribe Picrodendreae: 79, Cel dend (dot, Piranhea (dashed line); 81, Picrodendron (solid line, North America); 82, Parodiodendron (dot, South America); 83, Oldfieldia (solid line, Africa). North America); 80, T ы ү manner oi hick plausi pl i і of the new American forms." Later in this essay suggested. It would also appear that interchange Bentham clearly stated “that the most ancient home of forms between the principal Old-World centres of the order was in the Old World, whence it spread idi е à I kee of differentiation . . . continued long after the in- іп very remote times to America. terposition of the obstacles preventing the spread This scenario proposed by Bentham over a cen- ACALYPHOIDEAE Clutieae Pogonophoreae | Ж. c 1 —— um ta" a Cheiloseae _„— ЗИЙА ATY GE OTOP Ig p с Дл 4 FIGURE 5.° Distribution of Acalyphoideae. Tribe Clutieae: 89, Clutia (solid line, Africa). Tribe Pogonophoreae - 90, Pogonophora (dashed line). Tribe Cheil (solid line, Asi ges of the t bined); 94, Cheilosa; 95, Neoscortechinia. Volume 81, Number 1 1994 Classification of the Euphorbiaceae ACALYPHOIDEAE Chaetocarpeae —— ——— Тийе ин ата лыш ст. FIGURE 6.3 (solid line). tury ago now appears as a prescient anticipation of models of intercontinental relationships based on plate tectonic models. Pax (1924), although not mentioning Bentham’s essay, accepted much the rall explanation of intercontinental “апа same ove bridge" distributions in Euphorbiaceae, and—un- CROTONOIDEAE Micrandreae eject “continental drift" as an explanation. The very d paleontological record still d ot provide critical evidence for Euphorbiaceae, but th utions and relationships of the taxa are now better understood, so that Bentham’s model Manihoteae FIGURE 7.3 istrib crandra [and 206, Mi ДУ c Distribution of Crotonoideae. Tribe Micrandreae (dotted line, generic ranges combined); 205, Mi- icrandropsis; 207, Cunuria; 208, Hevea]. Tribe Manihoteae (northern and southern limits): 209, Manihot; 210, Cnidoscolus. Tribe Gelonieae: 217, Suregada 26 Annals of the Missouri Botanical Garden CROTONOIDEAE Adenoclineae FIGURE 8. Distribution of Crotonoideae, tribe Adenoclineae: 211, Glycydendron (solid line, America); 212, Klaineanthus (solid line, Africa); 213, Tetrorchidium (dashed line); 214, Adenocline (dotted line, Africa); 215, Ditta (dotted line, North America); 216, Endos; (solid line, Asia & Australasia). can be refined. It now appears that Africa/Mad- the Acalyphoideae and Euphorbioideae, the trail g ins the larg ber of primitive taxa of clues leads back to a joint South American/ of Phyllanthoideae, and is the most likely center African center, but a lesignation of orig of origin for the family, but South America would inal locality must await further studies. the cockpit of initial differentiation for With regard to the role of long-distance dispersal both the Oldfieldioideae and C. id Forboth in possibly accounting for these geographic pat- © EUPHORBIOIDEAE Stomatocalyceae ae 29 2 «щу S E FIGURE 9.’ Distribution of Euphorbioideae, tribe Stomatocalyceae: 272, Plagiostyles (solid line, Africa); 219 — Pimelodendron (solid line, Asia); 274, Hamil. (d 1 line); 275, Nealch (solid line, America). i Volume 81, Number 1 1994 Webster Classification of the Euphorbiaceae EUPHORBIOIDEAE Euphorbieae S FIGURE 107 à „ Distribution of ero eris tribe сона Subtribe Anth E PS line, Afri a). Subtrihe N. a 1, Dichostemma (solid | th 302, Neoguillauminia 6 (solid line, ny Caledonia); 303, увя пэт КЎ Australia). eee which: Thorne (1979) and more recently Ive for БШ. tropical intercontinental disjanetions, it is notable that the “basal” taxa in the Phyllanthoideae, ОМ. fieldioideae, and Acalyphoideae ia eae fruits with dry seed-c coats, which certainly do not suggest a high degree of B ios: ded qe ig pecori ic dispersal Only in and the Санс де. of the Eu- phorbioideae (Fig. 9) do aliy id or Bondi реш Bos 4 +h A : Агы PE g African disjuncts such as Mayaca, Pitcairnia, and Sac- coglottis represent recent long-distance dispersal, but he goes pra in iconotuding that this expla- nation ma f the taxa that link Africa. The transoceanic vicariant » America with patterns of genera and tribes in the Eup ceae agree with those of many other tropical families mapped and discussed by Axelrod (1970, 1972, 1975); these кш. EE that the Bereh ation of genera that seem more likel : » birds. The n nature e many Euphorbiacea would ind ed appea їп establish t afte LH patterns d above Lio that the Euphorbiaceae, as in many other ade des families, the spectac be widely sep- arated. " Suggested by: Sohuner 1929), the pro- gressiy, po ore of the Tertiary leads pem pido їаегу кас апа id >the dramatic ones of hag? аер -distance transoce anic e literatu tell us a grea t deal а the MORD recent Stocking of oceanic islands but uti little about the RE of less h evolved taxa that‏ ا Persing in the late Сы ceous or earliest‏ * ertiary. Because of this paradox, Thorne (1973)‏ by ocean-floor s ing. The ere isa дейын alternative explanation for trans- latitudes during the proposed e early Tertiary; Wolfe (1972) that оше Ызы groups. f ко into the Berir 15 land bridge. It is notable, however, that the genera he cited— Meliosma, Sagaretia, and Saurauia— m n offer Bucks “ше xi ше кшп alternative ,8 pos- sible example i is H lueggea (Webster, 1984); | how- 1987). There are also possible trans- cific or trans- Atlantic links in the subtribe A дынын, "The Oldfieldioideae show no indication of possible use Annals of the Missouri Botanical Garden of the Alaskan corridor. In the Acalyphoideae and Crotonoideae, an intrusion from Asia into America is men n у p iuc" pn of Aleurites & Leopold, 1967). There an is dure sign of СЯ іп а Бурк к) тус рея ini. - North American genus Acidocroton ap- obe a vicariant of the Mn оси Ла th erefore a from Eurasia seems to have bien an xplaining transoceanic cing model w С: о Americ. v have to involve plate tectonics as a major deter- minant. Although the overall ecology of the Euphorbi- aceae has not been reviewed in detail, it is worth- wi noting that ро: of ысы Bike oes Heywoodia, CMS occur as heale or small К with many b ge in the ot occur in rainforest or at least more mesi itats; thus the spectacular rescence of hytes such as the African succulent Eup ieae, Jatro- ih rs to a devel- has e ү deu deny Nm sach subfa Relatively - 1 ке ES OM CREE | + i | | opportunistic life-styl at fragmented, рее or extreme ps E ra habita In de osing this essay, I feel obliged to ne ран the large amount of conjecture that has been nec in discussing x major w ted phy- asa Pas ae alkene di Pes M wipes contrived, and evanescent. There may be J f 4l : 141 1 £ 1 ro pe peer states), and their гянт ‘the onward rush in e the > turbulent; convoluta d, and ultimately party "s taxa. LITERATURE CITED The к ph bá 363. 975. The Euphorbiaceae of Borneo. Kew Bull. ла Ser. 4: 1-245. 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Tropical Trees een For ts: Ап Architectural va ysis. Springer eril Heidelberg 30 Annals Missouri жай Garden Hans, A. S. 1973. Chromosomal conspectus of the relation to angiosperm taxonomy. Pp. 272-295 in uphorbiaceae. Taxon 22: 591-636. P. K. K. Nair tar EM in Plant Research, HAUMAN- d K,L. 1912. — d'éthologie flo- 5. Vikas Publ. House, New Delhi. rale ar quelqu espéces ae ntines et chiliennes. princes A 1905. aie der Euphorbiaceen- е t. Bot. Léo Errera ungen Dalecham € ia und Tragia. Sitzungsber. HAYDEN, uw. 3 7. Com mparative anatom my and sys- serl. Akad. Wiss. Math.-Nat. Klasse 114: 29- завуч of Ap ec imo enus Incertae sedis. J. 48. nold Arb. E -— 219. KónLER, E. 1965. Die MT der Biovu 1980. ана ои of aan laten Euphorbiaceae und ihre ee g fiir die Tax- (Euphorb ia aca) d ral Dissertation Title, onomie. Grana Palynol. 6: 26-1 EACH, L. C. 1976. Drivel an morphological versity of сои 1 Һе e identity e the genus /Veowawraea 9: 268-277. eny ie is e cotyledonary region nho Amer ыкы Britto 1988. Ontog of Chamaesyce кезү а 1701-1 J. Bot. а= 3 GILLIS, D. E. of Рісг e vih 2 ilius ан J. Arn 1962- 13, Chemo a ا‎ Übersicht über die Verbre und o реа. er de conde f 6 n. Birkhauser, Basel. Hon H. Blat Or номина, В. : Ein Anatomische Unt = von und Axe der Hippomaneen. Inaug.-Diss., Mu- ines ша . J. & J. А. Wor 975. The уч of M “phylogeny: Vegetative ee nn. Misso t. Gard. 62: 538-589. Durchsicht pen teren Sinne. c. Sci. Univ. Tokyo, Bot. 6: 2 09- HUTCHINSON, J. 1969. Tribalism in the жез Euphor- biaceae. Amer. J 56: 738-758 INAMDAR, J. A. ANGADHARA Studies on the power of vw Euphorbiaceae. Feddes Re- pert. 88: JANSSONIU s, H. AX classification of de Codes "Trop. Woods 19: 1990. Mikrographie des Holzes der auf Java n Baumarten [Euphorbiaceae, pp. 442- Weis "Brill, parkais JENSEN, U., ITSCH 4. Legu- minlike proteins and the beim of хе Euphor- Ann. Missouri Bot. Gard. 81: 160-1 J. B. Ѕмітн. 1 VocrL-BAUER & 4 N 1989, Strüctihe, ontogeny and biology of nectaries in Croton ens ee dianus oe Beitr. Biol. de n 64: 157- S. L., T. REYNO ns, D. F. Corm E F. J. Evans (editors). 1987. , tax- mic & economic iti af Bot. J. Linn. Soc. 94: Jury, l1 326 JussiEU, А. 1824. кре generibus med- icisque earumdem viribus ten n. Paris. Kakkan, L. & С. S. PALIWAL. 197 . Studies on the leaf anatomy of Eu ate L. Epidermis. Proc. In- mac bryological aspects of Eu phorbia dulcis L. Phytomorphology 11: 24-36. ———— & А. К. BHATNAGAR. Embryology in Dist studies of the tribe Euphorbieae берки нн чи their relevance to its classification and possibl lution. Excelsa _ „е 969. a oer Fl. Congo 8(1): 1- ANGO. Hymenocardiaceae. коа 147-200 i and Paleofloristics eii bk and Eastern Noi erica. heir, Amster Lenor Я. -F. 1 несе тунш ‘sur em nature et origin la fleur spem ie Por we oupe singulier hen te chdi He bd. Séances Acad. Sci 41-250. P d Euphoriaa ‚ Sér: D. и °C. re 986a. Systematic foliar MAS of Ph а qui (Euphorbiaceae), I. Con tus. Ann. Misso dimus Tío 29- 85. matic foliar morphology © of Phyl-‏ پا note (кросс ае). б. oe‏ r HETE III. Cladistic analysis. Syst. Bot. 11: 515- & М. С. Simpson. 1994a. plications of pollen ultrastructure in th e Oldfieldioi- dea oe nr onte Ann. Missouri Bot. Gard. 81: 1994b. Phylogea re- Fy. жык к ЭШ, MPSON. laos a ee and By nen сй a рһог ae). Ann. Missouri Bot. Gar KA MAHESHWARI, P. 1942. f Euphorbia heterophylla L.—A reinvestigation. Proc. Indian Acad. Sci. 15B: 158-166. MARIS, P.G Evolution ie the айса а in m starch grain m . MANNERS: 1987. айай and the classification of Euphor ue tax Bot. J’ The chemo of Euphorbia esula L. Linn. Soc. 0. Martin, Н. А. 1974. The identification of some Te tiary pollen be ves to the family oec Austral. J. Bot. 22: 271-291. i Brain x Australian flora oug y: Evidence 1 81 202.2 . TIFFN Frui rela and Palace Бойдо e) from Eoc eine inia and Mar 4: 3 MEEUSE, A D. T^ . The dies ied auct. ріш“ Ап unnatural а Eburon, Delft etic analysis. | 6c. pete tamatic foliar vite of Phyl Ph ee im- | zE Volume 81, Number 1 1994 ebster ron E sia of Euphorbi metophyte binucleate; pistillode present or absen 2-5(-20) as (1-) tyles mostly free hircos at base, usually bifid; ovu ch locule, anat- hemitropous, nucellar beak sometimes is so diverse that it into six separate families. T are grouped into 10 tribon a Pee fo om Web- ster (1975), where 13 tribes This large “basal” subfamily of Euphorbiaceae Meeuse yit pa ipee jimi 0 genera included d. The work of км (9000 b, e) has shown that some m aruncle perm present or ab- I sent; cotyledons i-em or r folded. Base chromosome _ within ће Antidesmeae. 2 or numbers mostly 1 KEY TO THE TRIBES AND SUBTRIBES OF SUBFAMILY PHYLLANTHOIDEAE la. кэм simple е, mostly entire; styles usually bifid. less th n lem long except in some Phyllantheae); seeds a. F lowers in ал Шагу юп pi е withont endosperm; leaves mostly ith F y 1 : о £A 1 £ 1 y 24-9 capsular or rp less commonly drupaceous. 4 Petals present fl ls imbri lp hy nsp 8 Rs | 71. 3 E. £ : = | b 3, 1 | аһзе 5a. Pollen exine finely ee not coarsely spinose; stipules deciduous РОЦЕ АМАМОЕАЕ Sb. 3 llen exine coarsely and deeply reticulate 5 spinose; s LL persisten . Petals absent, or ifi Lesen then sepals valvate, or wood and ovule сян SW en desinis usually present. 6a. Sepals valvate; ape present, us rter than sepals BRIEDELIEAE 6b. Sepals imbricate; petals absent. nt then often as long as "os YLLANTHEAE 3b. nd dink intrastaminal; leaves ola denk wood fibers thick- umm da xn fruit dru ovary 1-2-loc cular DRYPETEAE н> E -— 2b. Fl in Uapes ©; oi akent a if present, inflorescence КЕ 1еауез with stomata Gene ic or anomocytic; seeds with end osperm. мр Маа люс жаруу with t sent or absent; pollen = orate ient vay a bifi 7b. Pm vinged, Pn aro vies ANTIDESMEAE germ tanniniferous epidermal cells; inflo- styles unlobed Minden lb. Leaves trifoliolate; decis ст flowers mien инг disk absent; styles Gie fruits Басса! 8. BISCHOFIEAE nniniferous риши cells, not granulose-glandular; floral disk 6. rescences Tribe 1. WIELANDIEAE Baillon ex Hurusawa, t. 6: 339. ш eat gic cru with a deu pap ae J. Fac. Sci. Univ. Tokyo 1 or 2 ег E prese Pe or abe ent; embryo with шк ера adicle. 1954. Wielandiidées Baillon, e oh ч phorb. 18 picea subtribe or folded, much longer than the r e ax > zenr with the greatest numbers of unspe- Monoecious or dioecious trees or ames ж mentum sim mple; lea ves alternate, entire, nation ў с). M eglandular; papules deciduo us. pone phyletic in cladistic terms, which is essentially in- ipd hepistillate evitable for the basal in a large family such coker bracts inconspicuous. Sepals m честе 4-6, as the Euphorbiaceae. Relationships ат the imbricate; NM ually 5; disk usually annular; nine genera appear геп i stam -10, free or ғ flame ants connate at Dese; feasible to group them inia hisci that all of the genera occur in South America, larged; pe grains 3. b ee dorus e, re- Indian Ocean islands, and only Savia extends into the Northern Hemisphere. ticulate: s bifid; hi le SE styles DL KEY TO THE GENERA OF TRIBE WIELANDIEAE la. Seeds with end osperm. y 2a. Stamens 8-10: pal tal tly 4, very similar; ovary 4-5docular с — 1. Heywoodia Annals of the Missouri Botanical Garden 2b. Stamens mostly 5; sepals and petals mostly 5; ovary mostly 3-locular. 3a. Pistillate disk entire, not Dau d M 2. Savia ЗЬ. Pistillate disk glandularlobed. — — 3. Gonatogyne lb. ete with little or no endosperm. Staminate disk dissected 4. Petalodiscus es Staminate disk annular, sometimes lobed 5a. age 3- or Sethe чы ppc ves coriaceous; а longer po A CMM ‘Stamens Ree dne at Баве: — 7 SS Tb. Leaves chartaceous; petals shortei column __ 6. Actephila 6b. "реше leaves coriaceous. un als p resent; staminate "en шы Styles ted; stante k glabrous; from pistillod 7. Discocarpus à Styles slender; st. adnate to pistill biocg d cd b.. Tae hnostylis 8b. Petals absent il nocentrum Sb. Ovary 5-locular 1. Heywoodia Sim, For. Fl. Cape Col. 326, pl. 140. & Hoffm., Pflanzenr. 147. 1907; Pax XV. (Heft 81): 280. 1922; Hutchins Bull. 1922: 114. 1922; Milne-Redhead, Bull. Jard. Bot. Brux. 27: 327, pl. X. 1957; Rad- cliffe- -— Fl. Trop. E. Afr., Euphorb. 1: 85, fig. 12. 1987. TYPE: аыр lucens Sim (үне: лаг and Kenya south to Natal ibe Ph iu rs to me as perhaps nearest in morphological e to the hypo- thetical ancestor of the preclude an affinit may connect the tribes Wielandieae and Phyllantheae. 2. Savia Willdenow, Sp. Pl. 4: 771. 1805; Muell. Arg. in DC Trop. E. Afr., orb. 1: 83, fig. Li. 1987. TYPE: Savia purum (Sw. ice seid Smith & Downs, Sellowia 16: 177. т, : Kleinodendron piar Smith & ns 5 а via dictyocarpa Muell. Агр.]. About 25 species of America, d and Ma id agascar. he sections have been r Savia, 2 spp., the type and S. dieryocarpa x Brazil; sect. Не Urb., ca. 20 spp. of the on, Kew ce шш; ir Wielandia West Indies and Madagascar; and the recently described sec and other workers. 3. ве Muell. Arg. 1873; Pax & Hoffm., dud , Fl. Sgen 11(2) 13. t. Sp. Nov. 31: (“Klotzsch *) Pax & = (Hos 81): 187. 1922. Pu Gonatogill is (Baillon) Muell. Arg. [Amanoa ? ира Badlanl Monotypic; Brazil. 4. — Md Pax, Natürl. Pflanzen- ar Eod Savia — Petalodis- cus s Bis n. Euphorb. 571. 1858. Meses ay techs a Pax гаанс selected sii this was the species illustrated by Pax, loc. cit.). s of five or six species endemic to Mad- agascar; Басы with Savia by Pax & Hoffmann (1922, 1931) and Leandri (1958), but differing rom Savia in bein: g monoecious and having ех- albuminous seeds. n? yg ы Mém. Inst. Sci. Madag. 19 uell. Arg. Prodr. 15(2): 229. 1866. Betólodisces sect. Charid- ia (Baillon) Pax, ү РВапгепїат. 1. 3(5): 15. 1890. TYPE: EORNM (Baillon) Leandri [Savia longifolia Baill. ]. This genus of five or six Madagascan жо is мен NAE to Sa ea dete and ma dist Volume 81, Number 1 1994 Webste Syn opas = ^ of Euphorbi 6. Actephila Blume, Bijdr. Fl. Ned. Ind. 581. 1825; Muell. Arg., DC. Prodr. 15(2): 221. 1922; Airy Shaw, Kew Bull. 25: 496. 1971; Kew Bull. Add. Ser. 8: 21. 1980. TYPE: Ac- tephila javanica Miq. Anomospermum Dalz k. J. Bot. Tor Pi d. 3: 228. 1851 е Miers 1851). T mum excelsum z. [= жерш aee (Dalz. ) Muell. Arg.]. s of ca. e- scribed species, ey eur эм a taxonomi- cally. The position of ie genus has been variously interpreted. Pa ffmann (1922) associated it with Amanoa, which has similar flowers bu h Жа pm Kóhler oe pointed out a pal- drachne, and Levin (1986 ound a тетен іп leaf venation to present, it seems best to retain Actephila in the tribe Wielandieae. 7i ENS Klotzsch, Arch. Naturgesch. 7 , DC. Prodr. Pflanzenr. 147. XV (Hef 81): 202. 1922: Jablonski, Mem. New York Bot. Gard. 17(1): 84. 1967. TYPE: алас та essequiboensis Klotzsch. ording to the recent treatment of Jablonski, RT p | speci es of Discoca ари t are restricted to merica The ‘pene position di на bà uncertain. Pax & Hoffmann (1922 eiliad epar btribe (including o 0 965) euggeótedi it was close to Amanoeae and Bridelieae; and Mennega (1987) regarded it 35 out of place in the Wielandieae. 8. Lachnostylis Turczaninov, Bull. Soc. Imp. "on alistes Moscou 19: 503. 1846; Muell. E ^ut en 224. 1866; Bentham, hei. . Pl. 13: 61, t. 1279. 1879; Gen. PI. 3: “Dyer Cet 8. Afr. Fl. PL 1: 309. 1975. TYPE: Lachn urcz. [7 Lach- "m hirta 4 ) Muell. nii This genus, endemic to South Africa, includes iab Disc (1922), it differs in a number of characters. 9. Chonocentrum Pierre ex Hoffm Pflanzenr. 147. XV (Heft 81): 205. 1922; Jablonski, Mem. New хар Pt Sen, dd Bole er YPE: Cho rum (Muell. Arg.) Pax & Hoffm. nue cyathophora Muell. Arg. ]. notypic genus of Amazonian South Amer ica, regarded by Pax & Hoffmann as related to г Lo qs ү ا‎ t o 10. Wielandia Baillon, Etude Сеп. Euphorb 568. 1858; Bentham, Gen. Pl. 3: 270. 1880: Hemsley, Hook. Ic. Pl. 29: t. 2813. 1906; Pax & Hoffm., Pflanzenr. 147. XV (Heft 81): 181. Mns Leandri, Notul. Syst. (Paris) 7: 190. 1 ; Fl. Madag. ma 135. 1958. TYPE: НА elegans ВаШоп Monotypic; restricted to Madagascar апа the Seychelles. Although combined with Savia b Mueller, Wielandia appears closer to Petalodiscus hd flower production. Tribe 2. AMANOEAE (Pax & Hoffmann) Web- ster, Taxon 24: 594. 1975. eg era sub- Pflan tribe Amanoinae offm., 147. XV (Heft 81): 190. 1922. TYPE: nr noa, Aublet Moncecious mee» ип m ooo ly en penninerved, 1 dul 1 1 t some E. © Т 19 dnate t tiol l these sometimes forming d bracts conspicuous an tals 5, imbricate; sta- БЕ ў: оушез а ро ч styles рсе ‘bifid or iid l per docile endosperm pr resent, ы ог absent; соїу massive but not plicate, much longer than ыйы here interpreted, the tribe Amanoeae in- ME the genera Amanoa and Pentabrachion. In qued circumscription of the tribe (Webster, Annals eens P AEA Garden 1975), Actephila was also included. However, ev- idence from pollen morphology (Kóhler, 1965), leaf venation (Levin, 1986a, b, c), and wood anat- omy (Mennega, 1987) argues against a close as- sociation of Amanoa and Actephila. KEY TO THE GENERA OF TRIBE AMANOEAE la. Stipules not ге aad floral spicuous, not indurate; styles 5 ; frui 52 acts incon- ; floral bra cts conspic- urate; styles diated, ,emarginate; fruit ees walled, seeds 1 p e teens Sie tae ыы MM UE “12. Amanoa 11. Pentabrachion Muell. Arg., Flora 4 864; DC. Prodr. Mod 223. 1866 4 Pen оома: P chion биа MA Monotypic; West Africa (Cameroon, Gabon). Levin (1986a, b, c) suggested a possible e on- ship to r on ш basis of. f oliar nation characters acters x E баг Kahler pe cee еа е Атапоеае and Bridelieae becau di its pen reticulate pollen It лау be a better с to affinity, so » Pentabrachion i is here tribe with Amanoa, ee the obvious differences Tem the gene 12. rique Duc Fl. Guiane Fr. 256, pl. 101. ; Mue Arg., DC. Prodr. 15(2): 219. idee M ue Gen. PI. . 1880; Bail- lon, in Grandidier, Atlas Hist ipe d PL 210, 2 ax offm 147. XV (Heft 81): 195. 1922; pus Mem. New York Bot. Gard. 17: 967; 0 0. TYPE: Amano EINEN Au A mainly neotropical genus of 16 od the three African pene have been referred t endemic sectio TE tee vee oe Muell. Arg., Bot. Zeit. 1864 (as Bridelieae). Tribe Biovu- е sect. Brid delieae Thwaites, Enum. Pl. Zeyl. 861. Antidesmataceae subfam. Bridel- ioideae Hurusawa, J. Fac. Sci. Uni v. Tokyo Sect. 3, Bot. 6: 321. 1954. TYPE: акла Willd Monoecious (rarely dioecious) trees or shrubs; leaves alternate, entire, pinnately veined, without ds; siduous. Flowers drupaceous, 1-3-celled; seeds w e cotyledons much longer and broader than the rad- icle. е tribe has traditionally been considered to ime furthee anay ан, пе regarde Бий aw ED ب‎ hit ss of A К ie African species of Cleistanthus; follow latter's suggestion, it would be possible to -— the Amanoeae and Bridelieae as subtribes o ertheless, neither the floral nor of the Bridelieae appear su to Amanoa to associate them in їй single tribe. Nev ollen characters ciently close same tribe KEY TO THE GENERA OF TRIBE BRIEDELIEAE la. Ovary mainly of 3 кай [ux 2 or 4); fruits capsular, lobed, mainly 3 with г eeds per locule; leaf ventation orn reticulate ...... 13 Cleistanthus lb. Ovary mainly of 2 pd Rape Mee capsular, unlobe drupaceous or rarely ca 2-locular with 1 же рег bake leaf ui mainly percur 14. Briedelia 13. Cleistanthus € f. ex Planchon, Hook. Ic. Pl. 8: t e 848; Muell. Arg., e Sit Bentham, Gen. 3(1): 268. qe две, Pflanzenr. ud Р rop. E. Afr TYPE: Cleistanthus polystachyus Nanopetalum Hassk., Versl. Meded. Afd. Natuurk. Коп. coti Volume 81, Number 1 1994 Aka "n С Amsterdam s 140. 1856. T talum myrianthum Hassk. [— Cadi d (Hassk.) Kurz]. Leiopys Miq uel, Fl. Ned. Ind., Erste Bijv. 445. st cee sumatrana Miq. [= саар М g.]. Lebidieropsis "Mu ell. A а 32: 863. TYPE: ebidier e ойлы ve м Mael im [7 Cleis- tanthus coi ont eee ) Be Schistostigma ye m utsches уре, ds v с 190 = атаан pua. ut. [= Cice ibe papuanus (Laut.) Jabl. 1 bolts. us Gagnepain, Bull. Soc. Bot. France 70: 499. 1923 E: Paracleistus "тае Cangen: [= Cleis ssi an saichikii Merr. & Chun; lectotype, desi озн ted by Wheeler, 1975 Al сойо Gagnepain, Bull. Bot. Fran defroya p (Tabl. ) n cia MAS rotundatus Jabl.]. А large and тае а genus of over 100 species, of which t 30 occur in Africa/Mad- agascar and the rest in in кара Asia, from India esia. Both Koh s s of pollen an characters, respectively; clearly, generic limits in the tribe need to be critically reevaluated. 14. uM Willdenow, Sp. Pl. 4(2): 978. eandri, Fl. Madag. 58; rd Fl. Congo a1) 21. 1962; ые e th, Fl. E. Trop. Afr., Euphorb. 120. 1987; Chapman, Austral. PI. Name Index jg 1991. ТҮРЕ: Briedelia oxb.) Willd. [Clutyia sc Roxb.; н lectotype, selected here; of the з ѕре- ed іп the pisei description of laced in subg. illdenow saw herbarium material Candelabria еек Flora 26: 1843. ТҮРЕ: tha Hochst. i Briedelia mi- Candelabria micran x crantha (Hochst.) Cs ¢ogoetzea Pax, Bot. . 28: 419. 1900. TYPE: Neo goetzea brideliifolia | Pax [= J жун brideliifolia (Pax) Fedde]. Gentilia Веће, Compte Rend. Acad. Sci. Paris 114: 1294. т ТҮРЕ: боа аат ВеШе p» Brie- " а ndellensis Вее]. Penta Chiovenda, Ann. Bot. Roma 9: To TYPE: Tzellemtinia nervosa Chiov. [= scleroneura Muell. Arg.]. Webster Synopsis of Taxa of Euphorbiaceae An Old World genus of more than 60 species, the ma to match the Bridel) is contrary to the current rules of nomen- cl те 4. PHYLLANTHEAE Dumortier, Anal. 45. 1829. Euphorbiaceae sect. ied ae NEM Bijdr. 578. 1826 — Akad. = Berlin 1859: 1859. TYPE: Phyllanth SaL; Monoecious or dioecious trees, shrubs, o fedi leaves буо entire, pinnately veined, withou с п pistillate sepals mostly 5, imbricate, persistent or deci ae disk cupular to lobed pd commonly dissected or absent); ovary mostly 3-5-locular; vules ropous or hemitropous; styles bifid or entire ni БЫНА. Fruit usually capsular (rare- ly baccate or drupaceous); seeds 1 or 2 per locule; 1 + h ж laa } 1 1 J 1 igi, 1 hy зет WIth the radicie. € м here interpreted, tribe к, is con- si "deo narrower in scope than the circumscrip- of Pax & Hoffmann (1922), т p than inii of Webster (1975). T осазіа, Тро) point to- era to pollen (Punt, 1962; Kohler, 1965) and some o the leaf venation characters pointed out by (1986a ): Subdivision of this Beterogeneque ўров пер med 1 with six subtribes must be considered provisional. The Ды осы between Шоро: and hemitro- pous ovules, pointed out b systematically пирса, еуеп T not all gen- and there may pea rs ета һау prove to abe transitional stages. Annals of the Missouri Botanical Garden KEY TO THE SUBTRIBES OF TRIBE PHYLLANTHEAE la. Petals present; ovules an: ous; seeds smooth. 2a. yes tillate sepals бои staminal col- dnate to peltate pistillode 4a. Astrocasiinae 2b. Pistillate sepals Proms stamens free or connate b. Leptopinae lb. Petals absent (except i in some An drachninae ih ovules anatropous or hemitropous; seeds sm т dr tured. "Ovals anatropous; pistillode prese . Staminate disk annular; га" ms reticulate; seed e dull or sculptured c. Pseudola achnostylidinae ч уюн ed disk мее pollen grains uculose; seed coat blackish, smooth curineginae 3b. Ovules hemitropous; staminate disk an- nular or dissected; pistillode present or ab- sent. P. Petals present; stamens 5, free; sta- minate disk dissected ... 4e. An 5b. Med absent; stamens 2-10 or more © e G* free iid or dissected __ 4f. Flueggeinae patron: a rale Webster, Syst. . 1992. TYPE: Astrocasia Rob. Кы Dioecious (or subdioecious) trees or shrubs; sti T ules MAU poda in ax vuv y glo cus ts column; pollen grains 3- E reticulate pistllode peltate; рае sepals 5, de- oe styles bifid; fruits сара seeds 1 or т locule, smooth; cotyledons much longer du му ا‎ This monogeneric tribe includes only the type ocasia, which was i ibe Wie- A show to the tribe Тый, particularly to genera of subtribes Leptopinae and Pseudolachnostylidinae. . Astrocasia Robinson & Greenman, Bot. ebster, Syst Astrocasia phyllanthoides Rob. & Greenm [= Astrocasia tremula (Griseb. ) Webster] A neotropical genus of five species distributed from Mexico and Cuba south to Bolivia and eastern Brazil Subtribe 4b. LEPTOPINAE Webster, subtribe nov. TYPE: Leptopus Dcne tipulae persistentae; petala conspicua; stamina b f. lamenta libera tillata persistent ovarium 3: и ovulis anatropis; fructus capsularis; semina lae Dioecious shrubs or herbs; stipules persistent; азе їп аа агу кре, Roe Ke 59; , filaments free r connate below: pollen grains 3- рони retic- aii pistillode 3-fid; pistillate sepals 5, persistent; wage г ог dissected nid hers от veni testa smooth; waa et pai d than radicle. This e SI only the single genus Lep- top s submerged in Andrachne by Mueller ( (1806), AG (1880), and Pax & Hoff- mann (1922). However, Leptopus differs from An- on (sensu stricto) in its айар ороп оү а апа ae Packie: It ебе appears best to assign it to a separate subtribe 16. чү Decaisne, in Jacquemont, Voy. Inde 155. 1836; Pojarkova, p Syst. Herb. теч Bot. Acad. Sci. USSR es 8 cordifolius es Nuttall, . Phil. So 7 (no n Lepdan wd fue А phyllanthoides Nuit: (loc. da [= Lep- topus phy une ip (Nutt.) Webster, coal Thelypetalum te LE, м 5ос. Bot. France Ly 876. E: Thelype = Lep- ea E ) Pojark i peager an oi Airy Shaw, Kew Bull. 23: 40. ке E: Choi tria ive diplosperma Airy Sh енер us diplospermus (Airy Shaw) Webster, comb. поу.]. Ag 10 species of apes sc санай istribution w Old World (India to China, In distribu donesia, and tropical Australia) and North pe (Mexico, southern U.S., and Greater Antilles). засы $ масо AE Pax & Нов Pflanzenr. 147. ү Коя 81): 206. 192974 Pd Pseudolachnos- tylis Pax Volume 81, Number 1 Webst 41 1994 inane of Taxa of Euphorbiaceae Моше ог dioecious nett баг or un- ee nn ps "ww ionem мем | ог M US | 58 2p in axillary gl lomérules or cymes, bracts inconspic- men erar than radicle. sepals an- ous; staminate se 5, petals absent; disk a и азада а 5 as = ранено aii This subtribe of six genera is heterogeneous and pollen grains 3-colporate, reticulate; pistillode pres- yrs puts, Ce ie QUIS ровные tat Fett eiit йн Se Dal Bi pen ў мата dolachnostylis апі Keayodendron may not b P petals absent; disk annular; ovary 3-locular; ovules шер ташына oaar ponera KEY TO THE GENERA OF SUBTRIBE PSEUDOLACHNOSTYLIDINAE la. E Apre 3-locular, with 99 б seeds; staminate disk annular sil aie walled; cymes pedunculate; odi with sculptured testa; pistillate sepals ba. “Ch x za d eed; i fl bsessil 17. Chascotheca 3b. f gr^ ventr. in dicellat . Filaments За sms pisti valle de. . Stipules foliaceous, persistent; sepals broad and imbricate mt. 18. Zimmermannia 5b. Stipules narrow, deciduo у, зер als me apa not peus mua {ме iopsis 4b. Filaments connate to pistillode; карта 20. Meineckia 2b. C lled; pis allan cymes edite sell with smooth testa; pistillate nes deciduous; fen Pseudolachnostylis lb. Fruit Th ET with one seed; staminate disk dissected 22. Keayodendron Lo сее Urban, Symb. Аш. 5: 14. t. 3698. 1971; Brunel 5 Roux, Bull. Mus in, Fl. Cuba 3: 44. 1953. Plut Hist. Nat. Paris, IV. 4: 79. 1982; Radcliffe- e Un Symb. Ant. 3: 284. 1902 (non Smith, Fl. Trop. E. Afr. Bd 16. 1987. T. M. s. 1860). TYPE: быы пео- TYPE: Meineckia PERNA Baillon. lt RE ey (Grech, ) Urb ees Muell ТВО: 1864. ТҮРЕ Pow species have been described in this Nis Cluytia adra Trichopoda on : хь {= Meineckia ene а (Muell. Arg.) Webster o Men 4 Exe. (Сара: Шо Peltandra Wight, Icon. Pl. Ind. Or. 5(2): 24. 1852 (non ascotheca domingensis К, ) Urb. is до Үз Peltandra Raf., 18 9). Ne opeltandra Gamble, Fl a synonym of the type specie: NUR 1285. 1925. Peltandra parvifolia olia ht [7 Meineckia parvifolia (Wight) Webster; 18. Zimmermannia Pax, Bot. Jahrb. 45: 235. суро] 1910; Hutchinson, FI. Trop. Afr. 6(1): 739. A genus of 20 species with a dures distribution 1912; Verdcourt, Kew Bull. 9: 38. 1954; in the ES World an d Old нлр от Мехісо Poole, Kew Bull. 36: 129. 1981; Radel to Colombia and Bra n tral Africa to Smith, H. E. Trop. Afr. Euphorb. 1987 EE southern India, Sri Due and As- TYPE: Zimmermannia capillipes ti = eat һу Bodo nih (1987), Zim- st African species, plus transitional to Mei- 21. e er унум Pax, Bot. Jahrb. Hutchinson, Hook. Ic. Pl. 31: m neck ы nie Pax , Pflanzenr 147. XV (Heft 81): 206. 1922; Phillips, Gen. S. 19. V pie me mora Radcliffe-Smith, Kew Afr. Pl. ed. 2, 457. 1951; uei eg Bull. 45: 152. 1990. TYPE: Zimmermanniop- H. E. тнр. М. putes 80. 1987. sis хара Radcl.-Sm. Pse kindtii Pax A monotypic genus from Tanzania (Iringa Distr.), An African genus of six described keid, re- m = to Zimmermannia, and questionably duced by Radcliffe-Smith (1987) to four varieties o maprouneifolia Pax phylogenetic po- sition of Pseudolachnostylis remains uncertain, 20. Meineckia Ba illon, Etude Gén. Euphorb. although the arse of Pax & Hoffmann (1922) 586. 1858; Webster, Acta Bot. Neerl. 14: іп locating it adjacent to Meineckia is followed 323. 1965; Radcliffe- Smith, Hook. Ic. Pl. 37: бА However, ће pedunculate inflorescence and Annals of the Missouri Botanical Garden massive fruits of Pseudolachnostylis are quite ferent and it is no dM in pollen ornamentation represents con- rgence rather than close affinity. 22; мрн. лот, Soc. Bot. Franc 1959. : Keayodendron bri OR. Кең ех x Hutch & Dalz.) Leandri. notypic genus from tropical west Africa, es flowers are rapetaleus, with imbricate sepals. referred to this m УА lack of a better ик a Subtribe ES SECURINEGINAE Muell. Arg., DC. ipods 46. 1866. TYPE: Securi- a m Dioecious shrubs or small trees; usd decid- uous; flowers in дарии ня а bracts incon- са 5; 5 absents disk 1 ies primm Meca 1 or 2 per E + as Alt бей Mueller’ s name is EA to this sub- ircumscription is very different, since Mueller (1866) included Hymenoca and genera of subtribe Scepinae. The distinctive pollen and seeds of SE oe set i rt from other 0 ibe ntheae and suggest a possible affinity with do Oldfieldioideae. issieu, Gen. Pl. 23. Securinega C 388. 1789 (nom. ini Lea и Fl. Madag. 111: 107. 1958. TYPE: Санаан durissima ns.). J. F. Gmel. (typ. co A Malagasian genus of five closely related s 0 others from Madagascar. The majo have confounded the genus with Flue eggea, and only Leandri among modern writers has presented the genus as accepted here. Subtribe 4e. ANDRACHNINAE Muell. Arg., Linnaea 34: 64. 1865 (as Pax, & Hoffm., Pfla mein XV (Heft 81 ); 160 1922. TYPE: gemi ed Monoecious herbs or subshrubs; UE persis- incon us; staminate sepals mens 5, filaments m or ae pollen grains 3-colp striate; pis- AIE Pra pistillate epale 2: pore petii 5; locular; ovules айр pow bifid; fran TS seeds 2 er locule, testa sculptured; cotyledons much lon- ger than г е orate, The ing Al Uoc un "ys ere Andro ers included Leptopus within Andrachne. " f aa point toward affinity with tribe Antidesmeae. 24. Andrachne L., Spec. Pl. аа 1753; Gen. РІ. ed. 5, 4 4; Mue ., DC. Prodr. 15(2) 232: 1866; rm M Pl. 3: 270. A ax n Pflanzenr. 147. XV (Heft 81): 169. 1922; roizat, J Acad. Sci. 33: 42; Vindt, Trav. Inst. Sci. Chérifien 6: 4. 1953; Pojarkova, Not. Syst. Herb. Inst. Bot. Acad. Sci. USSR 20: 256. 1960; Webster, J. Arnold rbor 27. 1967; Radcliffe-Smith, Fl- Trop. E. Afr., Euphorb. 7. 1987; Gilbert & Thulin, Nordic J. Bot. 8: 159. 1988. TYPE Andrachne telephioides L. [lectotype, 45 ignated by Small in Brit 0 H N. U.S. ed. 2, 2: 453 Eraclissa Forsskál, Fl. Aegypt.-Arab. 208. 1775. Eracliss a hexagyna Forssk. [= Andrachne “de phioides 4 пар р ez Ortega, Tabulae Bot. 15. 1773. TÊ Ra procumbens Moench [= 4 Andrachne _ telephioides L.]. Volume 81, Number 1 Webster 43 1994 Synopsis of Taxa of Euphorbiaceae pe fo sk Didrichsen, Vidensk. Medd. Naturh. For- pun aida ener in some Ses OWES i in ax- n. Kjøbenhavn 1857: 450. 1857. TYPE: Phyllan- : У thidea microphylla (Lam.) Didr. [= Andrachne mi- И ОШО (Lenk) Ballon. s 4 6, free or connate: stamens 2— 15, connate; ins ex ке кш and ну dissected the restricted circumscription adopted here, oy co In Andrachne includes abou t species with a pri- гт! pistillode present or absent; pistillate marily Tethyan distribution from Persia through im RISUS 5 or б, регинеп ити; ak usu Нау the Mediterranean to the West Indies, with one cu pular, oft I species (4. microphylla) disjunct from Baja Cali- mostly 8 (riy 2 or 4-10); styles free or ЖОНЕ; fornia to Pacific South America. The majority of bifid o entire; ovules Шр fruit My species listed under Andrachne by most authors ah are better classified in a separate genus Leptopus. ^ usually 2 per locule; testa smooth. or sculptured; endosperm copious; cotyledons broader than and dicle. Subtribe 4f. FLUEGGEINAE Muell. Arg., Lin- 1-2 times longer than the radic PE т id a ee oru A variable subtribe of nine genera with over 1000 species, evidently monophyletic (possibly ex- Tribe ees ord Euphyllantheae Muell. Arg., cluding Aerisilvaea, however). The vast majority in a 34: 64. 1865 (nom. illeg.). TYPE: Phyllan- м the species in genera lacking pistillodes (except Tribe. Phyllantheae E Beh ee Muell. Arg., or Margaritaria and some species of Phyllan- a 34: 64. E: Sauropus Bl. тит share a distinctive е оа Tribe Phylantheae Као Phyllanthinac Pax, Natürl. leaves are redu x to s us m and are ed. 1, 3(5): 17. 1890. TYEE Phyl- deck only This has been оази аз jw big branch- ing" ri 1956). Monoecious or dioecious trees, shrubs, or herbs; KEY To THE GENERA OF SUBTRIBE FLUEGGEINAE la. tale present in staminate flower (sometimes minute). - Dioecious; stamens 3-7; pill E not ке Flower ill clus: e flow z. Hat, Я » i ly 2 T locule i 25. Flueggea 3b. Flowers in axillary or lif or panicles; inate flowers subsessile; seeds mostly T 26. Richeri ly 27. Aerisilvaea ent 2b. Monoecious; stamens 17-21; 1b. Pistilode а j rio or dioe - Seed coat with bony endotesta (not ventrally invaginated) and fleshy exotesta; i» dry, irregularly e а e dU м зш 28. Margaritaria 4b. Seed coat seas bony "otc and fleshy exotesta. (or jr endotesta a aey ventrally i invaginate ted); fruits iem NI usually regularly аы cidal; monoeciou: trees, a S, 0 erbs. 5a. Floral disk а present; seeds Ж, not ventrally invaginated - Staminate dis x xtrastaminal; stamens various, but rarely 2 and introrse; vulneris much dicle Phyllanthus broader than 6b. Staminate disk lobed, i taminal; st. 2. anthers introrse; cotyledons =? ална radicle eur ei. a d. ord disk absent or completely adnate to calyx; pollen h rate; styles bifid o ,an Lá real invagina ated. ular л a ga € 5 = а entire; seeds with thickened dry or fleshy exotesta 7a. Styles bid or r emarginate;, anthers not apiculate; ovary 3-loc ME + discoid, not turbinate 31. Загоры 2:01:92 Brey! Tb. ads гай up entire; anthers capt seed-coat usually fleshy; ovary 3-8- DE AB Glochidion 2 caly vx turbinate зс э al Browne, Civ. Nat. Hist. н 385. 17 25. Flue W 1.1806; 56 Be pair ty ee oe parlor: rej.; non Ac idoton Sw., 1788). TYPE: Adelia am ‚ 1880; Weheter, = i dido Fh L. [= Flueggea acidoton Ae ) Webster ae + 273. 1984: D yden, Brittonia Бейш Sprengel, Pl. Minus Cogn. Pugillus 2: 90. 1815 268. 1987. TYPE: Е lueggea leucopyrus TYPE: Bessera inermis onam en an = Flueggea virosa (Roxb. ex Willd.) Voi Annals epis MR Garden н Fischer & Meyer, Index Sem оро : 28. 1835. TYPE: Geblera e Mr allas) Fac & Meyer [7 Flueggea e RR (Pallas) Baillon]. Colmeiroa Reuter, Bibliot. Univers. e 3827-215: 1842. TYPE: Colmeiroa buxifolia Reuter [= Flueg- gea tinctoria (L. in Loefl.) Was ; Pleiostemon Sonder, Linnaea 23: PE: Pleiostemon verrucosum (Thunb. ) ves © Buse? gea verrucosa (Thunb.) Webster]. — Rock, Indig. Tr 913. E: Neowawraea phyllanthoides Rock vo Flug. d neowawraea Hayden]. А dine of 14 specie and temperate eastern Asia [un with a disjunct relict КЕЙТ, The genus was merged with Se- curine, ga by бав ох (1866), who was hi ea by owever, Baillon (1858), Bentham (1880), and J. ч Hooker (1887) m tained Flueggea as distinct; and indeed it is not ees related to Securi e as indicated by both en and seed character 26. eee Pax & Hoffmann, Pflanzenr ions. Р i 26. 1922; "Natür b am. : 48. 1931; Hore Garden ^ o vom 0927 ЗР 933: Airy Shaw, Kew Bull. 25: 489. 1971. ТҮРЕ: Richeriella gracilis бе Рах & Hoffm [Baccaurea gracilis Merr. |. nus of two species native to southern China, з; Pine N Malaya, and Borneo. It is ver close to Flueg, 27; е mst iris Kew Bull. 4 49. 1990; 4 7.1992. TYPE: нр pnt enfe enus of two species, one from Tanzania and the dis from Malawi. It is anomalous within this goes because ot its mee stamen ШШ ап nd the pollen can be examined. Levin (pers. comm.) Ея } E ы. TY. 55 7 JP 28. gS ge L. 9s e 66. 17 ‚ Kew ew aede 63. 1987. TYPE: Маш ш, по- bilis Prosorus Dalz., J. Bo en Misc. 4: 345. 1852. T Р, z. [= Margaritaria india x Baillon, Etude Gén. Euphorb. e np дерет ит zeylanicum Thw. ex Baill bs Margaritaria cyanosperma (Gaert- er) À ec de Wartsia Ba illon, A Adansonia I. 1: 186. 1861. TYPE: Wurt- sage rand Baillon [= edere tetracocca п) Webster]. on a, pro ex Таја. ud 21 Nat. Tijdschr. Ned. B 8. 1864. ». Calocaceus sundaicus & s ism. & Binn. Е Margaritaria indica Uh у Airy Shaw]. genus of 14 species occurring in tropical America, Africa, Asia, and Australia 29. hitman a x Р: 981. 1753; беш ед. oni Brown, 2, 2: 453 Niruri Adanson, Fam. Pl. 2: 356. 1763. TYPE: Phyllan- ap ex Cicca L., Syst. Nat. spam Cicca disticha L. [= Ph hats p [ ) Se els]. Xylophylla L., Mant. E: Xy- lophylla latifolia L м яа и. куша thus L; lectotype]. Conami eng Hist. Pl. Guiane Fr. 926. i brasiliensis и T Ae das ‘rast is (Aubl. ) mn u^ Genesiphylla L'Heri Е: бе nesiphylla ИШ г Wis 1- Pilan la- tolius Sw w.) Ca pw Fl. Cochinch. 607. 1790. TYPE: Cath- us fasciculata Lour. [= Phyllanthus cochi inchi- nensis Nymphanthus patr eiro, Fl. Cochinch. 543. 1790. TY Nymphanthus ruber Lour. [= Phyllanthus rule (Lour.) Sra ng.]. — л Volume 81, Number 1 1994 Webster Synopsis of Taxa of Euphorbiaceae Emblica psu. Fruct. Sem. Pl. 2: 122. 1790. T s a officinalis Gaertner = рге eta ет- Lh А ar Gen. Pl. 387. 1789. TYPE: Kirgane a . F. Gmelin [= Phyllanthus virgineu (Cm ДО) Pers. Tricarium оне Fl. Cochinch. 557. 1790. TYPE: Tri- carium cochinchinense Lour. [= Phyllanthus aci- dus eels]. ie vel Swart, Fl. Ind. Occ. 2: 1095. 1800. Ty к» m azillare (Sw.) Sw. [= К ax ax- “ape OM l. Arg.; lectotype]. ao Ка ме ы Minerva 42. 1821. TYPE: Geminaria ERR Raf. [= Phyllanthus caroli- Walt ab A. Jussieu, Бүр, Tent. 23, 4. TYP nk И: Nada cryptophila Comm. ex s. [= OUR APRS (Comm. ex Anisonema Ti u, Euphorb. Tent. 4 (nom E: Anisonema р n (Poir 3 А. Juss. i УП а pens ix uas Blume, Bijdr. 582. се- кечен BI. [= Pilani URE ا‎ ‚ Neogenyton 2. 1825. TYPE exemia caroliniana Raf. [= niensis Walt.; tot d Hex ylva : Syn- Phyllanthus caroli- ll. 92. 1838. TYPE: Hex- а angustifolia | (Sw. E Raf. [7 Phyllanthus an- gusti us (Sw.) Sw.]. Moeroris Rafine үн, Sylva Tell. 91. 1838. Е: Мо- a Raf. [= Phyllanthus A atus Sylva Tell. 92. 1838. TYPE: deni s (L.) Raf. меер = Phyllanthu $ maderaspaten Asterandra Klo . Na seca, 7(1): 200. 1841. Pe Kin andra ‘cornifolia (H BK) КІ. [= Phyl- есще НВК]. лоста hrs ж» Tijdschr. Natuurl. Gesch. Physiol. : Eriococcus gracilis Hassk. P va see gracilipes Muell. Árg.]. Ceramanthus Ha лит їз. д m Hort. Bogor. Alt. 240. 44 us gracilis Hassk. [= Phyl- lanthus М (као) ay Shaw]. Macraea dv , Icon. РІ. OP OR AEE мы 1852 (поп Мала Lindley, 1828). TYP ne Wight [= Phyllanthus sim- 5(2): 27, t. 1903, 1904. ҮРЕ: Reidia fortuna teen Hr SE эш C thus u Arg.; lectotype c ere] ra Wa Ico n. Pl. Ind. Or. 6 Үч d i oa: 3: 375. 1852. iche; lactis Modis Hance [= Phyllan- thus noises т .) Skee бый саш ах Griffith, N вы Pi. Аче, 4: 476. 1854 = Phyllanthus aci- tude Gén. Euphorb. 645. E: Hemicicca japonica Baillon A ны» ы, (Sieb. & Zucc.) Muell. Arg.]. Williamia Baillon, Etude Gén. Euphorb. 559. 1858. TYPE: Williamia pruinosa Baill. [= Phyllanthus warp oepp. г; ice emn on, ‘Etude Gén. Euphorb. 616. 1858. eee ruote phyllanthoides Baillon [nom. il- Ye nthus orbicularis HEN Phyilanihodendron gruth y, Hook. Ic. t. 256 8. TYPE: Phylianthodsadron гаа мы Аг n) ) Hemsl. [= Phyllanthus mirabilis Muell. Apres uud Bull. Herb. Boiss. II. 5: 488. 1905. orosella Moet al Chod. [= Phyllan- ik us hacen * Men pex totem (Muell. Arg) К . Schuma nn, Fl. Deut Schutzgeb. Nach. 289. 1905. TYPE: Phyllanthus ичин uell. A Nymania K. ба FI. 2а sche е Nachtr. p. 1905. TYPE: Nymania insig TE {м hum. [= hyllanthus schumannia LS ы ы Pax & Hoff: Palin hg T p 47): 95. 1911. TYPE: ашаа siamensis Pax offm. [= chs saa ^ roseus (Craib & Hutch.) Beille]. Torrey Bot. Club 16 adium formosum (Urb.) и e нөк у Bot. Clu ib 16: 12. : Кат. = Phyllanthus seen Ur M Кра Briton, чамі у Bot. Club 192 тй oigia comosa ia (Urb. vest PE. us Ur Dendrophyllanthus pera Moore, J. Linn. Soc. Bot. : 395. 1921. onii Sp. Moore [- S з © N m u TYPE: зону итеге {асты Gamble [= Phyllan r oon: аң (Gamble) Web- ег, со Hexaspermum Domin , Bibl t. 89: 315. 1927. TYPE: Hexaspermum paniuatum Domin [= Phyllanthus E =й, uell.) Diels]. митра Gagnepain, Notul. Spt اا‎ 14: 32. 1950; J Shaw, Kew ew Bul 14 praedo (Gagnep. ) Webster, comb. п A large and very diverse genus of 750-800 species, about 200 of which are American, 100 African, 70 Madagascan, and Ec rem iude Asian and Australasian. Many attempts have been made to subdivide the genus, which pire аул is un- natural. Here, the | d f Muell (1866) A 1 end} n C "Гы ЖЕ AGE B 10 and Margaritaria as distinct. 30. ica A. Gray, Proc. Amer. Acad. Arts Sci. 16: 107. 1880; Webster & Miller, Rhodora а 200. 1963. ТҮРЕ: Reverchonia агепагіа А. Ста A monotypic genus confined to sand dunes the ыл a United States iad кеб em Annals of the Missouri Botanical Garden ico. Although it is very close to Phyllanthus, it would appear excessively foreign if placed in that nus. 31. hanc n quor ge N 1826; Muell. Рт 15(2): 239. 1866; Bentham, p ) me [= Sauropus о T ) Merr.; A posue selected here ]. Cores Wight, Icon. РІ. Ог. 5(2): 26. 1852. TYPE: = Sauropus 1854. :B 2 £z. з ® ` o & . Phyt. Austral. 1: 32. mon ramosissimus F. Muell. s (F. Muell.) vid Shaw; уре, designated by Wheeler, 1975 Breyniopsis Beile, E Soc. psis pierrei Beill peers: pierrei (Beile) Webster, comb. p Heterocalym a Domin, Bibl. Bot. 1 927. TYPE: Hae calymnantha dre Domin [= Sauropus аи (Е. Muell.) Airy Shaw]. A genus of about 50 species im from India and Sri Lanka to southern Chin: Phi i as problematical as its separation from Synoste- mon. 32. бурек. R. & С. Forster, Char. Gen. Pl. 1 & С. Forste AC a Bijdr. 590. 1826. түре: Melanthesa a Blume мїм ye here; = Brey- nia racemosa а ) Мое Melanthesopsis Muell. , Linnae y m Melanthesopsis к. (Poir.) ye jh T nia fruticosa (L.) Hook. f.; lenny Pies b Wheeler, 1975]. A difficult genus of 10-15 highly variable spe- cies, f 1 Asia, Indonesia trop- ical Asia, barely separable from Sauropus on the one hand, and Glochidion on the other ЭЗ. Mie no eee R. & G. Forster, Char. Gen. adeniya 1 8; & Hoffm., Natürl Pflanzenfam. ed. 2, 19c: 56. 1931; Airy Shaw, Kew Add. Ser. 4: 115. 1975; Kew Bull. Add. Ser. 8: 92. 1980; Kew Bull ::6831 1980; 36: 298. 1981; A. C. Sm 1. Vi tiensis Nova 2: kt CHE TYPE: Glochidion ramiflorum J. R . For Agyneia L., Mantissa Pl. 2: 1771. TYPE: Agyneja pubera L. [= Glo. d$ d. йа ся, (L.) Muell. Arg.; lec КҮР, г һеге dips E^ x Gae ermer, Fruct. Sem et adleja sinica Ciera & Goa ion sinicum i Caeci H.& A Gynoon A. Juss., Mém. Mus. Hist Ма. ума 10: 335. di YPE: Gynoon rigidum A. Juss. [= Glochid- gidum (A. Juss.) Mu SN Glochidionopsis Blume, Bijdr. 588. 1826. E: Glo- S ionopsis sericea Blume [— Glochidion seri- m (Blume) Hook. f.]. Glochisandra Wight, Icon. Pl. Ind. Or. E: Glo chi sandra. acuminata Want 2 "n z.]. ee Ponte Bot. Zeit 1857. TYPE: Zarcoa Sati ppica Llanos А hue album (Blanco) chum., Nachtr. Fl. Schutzgeb. 292. Coccogloehidion ert ococcus [= Glo- - Hemiglochi m y Mueller} Teroglochdion K. Schum., Nachtr. Fl. Schutzgeb. 291. t: Tetraglochidion gimi K. Schum. [= Glochidion gimi (K. Schum.) Pax & Hoffm.]. An Asian and Australasian genus ога more than w extending to Polynesia. E ( workers have treated it as distinct, even though ! = py SE et £ tral taxê within Phyllanthus. Tribe = ee pepe ae J. Fac. Sci. Univ. Tokyo, Sect. 3, В 6: 334. 1954 lea tribe in Макылы: E and the Pacific islands. The genus FF | Volume 81, Number 1 1994 Webster Synopsis of Taxa of Euphorbiaceae ee eee Drypeteae Griseb., Fl. Br. W. Ind. 859. ТҮРЕ: Drypetes Vahl. donimo: Putranjiveae Endl., on Pk 28741837 feudis Беи, Sgn Nat. 837. Putranjivaceae En dl. ex ,E офісне nd Plur. 30. 1990. TYPE: PRAG , Wal lich. ill ide Сеп Euphorb 561.1858 CHEER sie DEP aea ad Baillon ex Muell. Arg., Linnaea 34: 64. 1865. TYPE: Cyclos- temon Blume [= нурї Vahl]. ecious trees or shrubs; leaves entire or den- te absent; staminate disk intrastaminal; stamens (2—) 3-20(-50), free, anthers introrse or extrorse; pollen grains 3-colporate, êre pistillode small uous фран ovules апаї hope us, with ma capsular ме per locule or fruit; endosperm copious; ганка plane, broader and somewhat longer than radicle. 8 b tribe Drypeteae includes four genera, with th ed in the pua b Webster (1975), is really a species of Flueg, КЕҮ ТО THE GENERA OF TRIBE DRYPETEAE la. Ovary 3-locular; fruit capsular; styles bifid, style- Ке slender .. ‚ 34. Lingelsheimia lb. Ovary mostly 1- or 2. Jocular; fruit drupaceous; Im t bifid. Disk present; stamens mostly 4 or more; styles stigmatiform 3a. Pistillate beak imbricate in 2 de- ciduous in fruit ... . Drypetes 3b. ГИ sepals open i in bud, S dt 36. Sibangea - Dis Е. Eie stamens се 2 i `g styles petaloid-dilated _ 7. Putranjiva 34. Lingelsheimia Pax, Bot. Jahrb. 43: 317. 9 db t. Belg. 84: 49. 1951; Brenan, Kew B ne 1952: 445. 1953; Léonard, Bull rux. 32: 513. 1962. TYPE: Ишет ышна ein ion Pax. Danguyodrypetes Leandri, Bull. Soc. Bot. France 85: 524. 1938. TYPE E: Danguyodrypetes E vensis Leandri [= Lingelsheimia manongari- vensis (Leandri) Webster, comb. nov.]. As interpreted by Léonard (1951, 1962b), Lin- gelsheimia is an Afri ou g ppe no n the Dry- Botas "t ^ is aei Mut its correct yis is Levin ( рег uggests 7 55 a possible affinity to pt ume 35. ا ی‎ "ein Amer. 3: 49. 18 Muell. Fl. Vitiensis 1 Nova 2: 455. 1981. TYPE: Dry- petes glau Koelera Willd., Sp. Pl. ed. 4, 750. 1806, nom. illeg. (non Koeleria ari 1805). Limacia Dietr., Nachr. Vollst. Lex. Gar 383. 1818, nom. illeg. peg ue Lour., 179 X TYPE: Koelera laurifolia d. [= Drypetes lateriflora (Sw.) Kr. & Urb.]. Liparena Poit. ex Leman, Dict. Sci. Nat. 27: 6. 1823. : not designated. Cyclostemon Blume, Bijdr. 597. 1826. TYPE: Cycloste- macrophyllum Blume [lectotype, selected here; tes dert: (Blume) Pax & Hoffm.]. , Edinb J. 14: 297. = Drype = Wight 833 (as Henieyelia). 1 TYPE: Hemicyclia Ке Wight & Arn. [= Drypetes sepiaria (Wight & Arn.) Pax & H ae — ыл pun l. Ind. Orient. 6: t. 1992. 1853. tylis aer Wight [= Drypetes venusta & Hoffm t. Kew Misc. 1: Т; . Akad. Wet. 4: 141. 1856. TYPE: Dodecas- emon teysmanii Hassk. [= Drypetes teysmanii Hassk.) Bakh. f. & van oe Pycnosandra Blume, Mus. Bot. 2: 191. 1856. TYPE: PEC serrata en) ا‎ h Dry- Blume) Pax : offm., nom. illeg. b.) = Drypetes teys- g&B з ЧА Pl T ب‎ @ @ horb. 6 [= Dr ne онык (Baill п) Апаиа nm ed. I Eerste Bijv 410. 1861. TY naua ‘sumatrana iq. [= Dr rypetes suma- di (Miq.) Pax & Hoffm.]. Annals of the Missouri Botanical Garden nee м» оше, Madras J. Lit. Sci., se . 1861. Е: Laneasa gum oblongifolia Beid. = “р me longifolia T ) Pax Stelechanteria Thouars ex Ba 1864. TYPE: Stelechan wn y 147. eria а thouarsiana Baillon [= Drypetes etin iana (lin) Capuro iiber Muell. Arg., DC. 15) 245. 1866. EAR Abs teme sessili AK ш Allem.) Mue g. [= bah tes sessiliflora m.]. Hunbloria Bails, B ull. Mens. Soc. lie. hi is 593. E: Humblotia comorensis Baillon [= Dry- petes ) Pax offm.]. Guya] Frappier ex Cordemoy . Ile Réunion 350. 1895. YPE: Guya caustica "Te. ex Cordem. [= Dry. s caustica е. ех Cordemoy) Airy Shaw]. е лея Hemsley 7. TYPE: ыа griffith Л (Hoo f) Hue. [= Drypetes indica (М uell. Arg.) D & Hoffm.]. Calyptosepalum Spen еназа J: Me YPE: Calypt palum in Drypet s ulyptosepala a vent pss Brexiopsis l Perrier, Norul Sys ier 10: 192. 1942. E: Brexiopsis aquifolia т. [= Drypetes dd E bes 3 bathi iei Capuro TE Таа r A large and variable genus of ca. 200 species; New World, with ca. 10 mos & Hoffman 22) in combining . Th i er study. I am indebted Levin (pers. T for critical sug- to Geoffrey gestions on the generic synonym 36. Mee Oliver, Hook. Ic. Pl. 15: 9, t. 1411. 1883; Radcliffe-Smith, Kew Bull. 32: i 1978; Fl. Trop. E. Afr., Euphorb. 101. 1987. TYPE: Sibangea arborescens Oliver. As interpreted by Radcliffe-Smith, Sibangea cludes three African species. The genus is very close to Drvpetes d it 1 } } 1 wnetin- Ж er it can be maintained as distinct. 37. piss caede Wallich, Tent. Fl. Nepal 61. ; Muell. Arg., DC. Prodr. 1502). 443 prey Bab: Gen. Pl. 3: 277 "ud Pax & Hoffm., Natürl. Pflanzenfam. ed. 2, 19c: 1931. 9. TYPE: Putranjiva МА а Wal- lich [Nageia putranjiva Roxburgh]. Palenga Preis ee 27 “ 1856. ТҮРЕ: Pale 2 Putranjiva i (Thwaites) Muell. аср Liodendron Keng, J. Washington Acad. Sci. TYPE: Liodnidiés on matsumurae E Keng = Putranjiva matsumurae Koidz.]. . Misc. 8: не genus of three or four Asian species н гі Lanka, Taiwan, and the Ryukyu Is- lands), vaina separable from Drypetes. Tribe 6. ANTIDESMEAE (Sweet) Hurusawa, J. Fac. Sci. Univ. T . 6:822] 1954 [as tribe of Antidesmataceae ]. Ordo Sti lagineae Agardh, Aphor. 199, Class. Pl. 9. rd ort. Brit tidesmeae Tho Enum. Pl. Zeyl. 289. 1861. E: Antidesm Dioecious tarot monoecious) trees or shrubs 5р 3. colporate, maios pistillode pre 2 or 3 (-5); ovules anatropous; styl bifid to шө (rarely entire fruit ی‎ eed or Hp seeds mostly | per locule (or 1 per fruit); seed-coat dry or fleshy; endosperm p to copious; cotyledons mostly broader than radicle. This mainly Роек. tribe of 18 e in 5 subtribes is well characterized by its leaves wi Berens epidermis oou. 1986a) and | roel pollen grains with 3 narrow colpi (Köhler, 1965). The tribe is кые given a broader circumscription than disi ef oe re i acinae, or Porantherinae KEY TO THE SUBTRIBES OF TRIBE ANTIDESMEAE la. Stems with resinous secretion; inflorescence pa- Ero. ог d 2a. Pe tals present; inflorescence terminal, pa- niculate, pera small; pollen sexine not spinulose; fruit capsular; «кан aracytic E xc) p p i а 205P absent; inflorescen: LE ai gen large and оор, ро len sexine spin inulose; fruit drupaceous; stomata ic b. Uapacinae lb. Stems lacking resin; inflorescence we panicu- d or capitate; p india abse I dle trees or e; anthe — es epa) 4b. Pollen ica distinctly ea anther locules + pendulous; leaves not glan- dular; stomata абуйс. cs o m 6d. ‘Antidesminae | " Volume 81, Number 1 1994 Webster Synopsis of Taxa of Euphorbiaceae 3b. Inflorescences terminal, mene herbs e. Porantherinae Subtribe 6a. SPONDIANTHINAE (Webster) Webster, stat. nov. it Spondiantheae Web ster, Taxon 24: . 1975. TYPE: гніт dia Engler Dioecious trees; leaves entire, penninerved, stip- east in filament ant e; pi ode e present; ik disk vais styles free; fruit sistent; er locule, testa de жүүн scanty; embryo um cotyledons much broader than radicle. otypic tribe, including only the African nthus. Pax & Hoffmann (1922) i group of the other woody tribes e. The Nee of Köhler (1965) on pollen mail a similar pos 38. Spondian 1905; ni Jahrb. 36: 215 1958; Aubreville, Fl. Forest. Cóte d'Ivoire, 2, 2: 62. 1959; Radcliffe-Smith, Fl. E. Trop. Afr. Euphorb. 1 1987; Léonard & is db Bull. Bot. Nat. Belg. 59: Jard. 1989. TYPE: Spondianthus preussii En- Megabaria De wia. d. Fl. Bas & Moy. Congo 2: 284. 1908. T Ho minia trillesii De Wild. [= Spondi Ба preussii Engler]. otypic genus of a a single variable pd аы. іп et Africa from Liberia to An- gola а and Tan Subtribe б, MAPACINAL Muell. Arg., Lin naea . 1865 (as Phyllantheae er ribe paceae (Muell. Ar, . TYPE: Uapaca Baillon Dioe us trees o nately ve Com stipule S; уа apitate, А crate, pedun Petalous and without floral disk; pua pamo: ; leaves entire, pin- phyllous; stamens 5, filaments free, anthers in- trorse; pistillode apically dilated: pistillate calyx gamophyllous; ovary mostly 3-locular; styles free, it drupa 1 buminous; cotyledons much longer than radicle. his subtribe includes only the single gen i ii o which ha as a always PIT isolated zg Airy Shaw (1965) and Meeuse oe) have as- signed it to a Bessa tie mily, Uapa aceae. How- v and Pax (1890) АЕ а it to his subtri an affinity vith учн ie Sce Дт un the other bend, the are diver rgent characters, and it appears bidet re- ferred to a separate subtribe. 39. Uapaca Baillon, Etude Gén. Euphorb. 595. 1858; Muell. Arg., DC. Prodr. 15(2): 489. 1866; Pax & Hoff (Heft 81): 298. 1922: Leandri, Fl. Madag. ШЧ). 163. d пакіне Зан ELE, Trop. Afr., Euphorb. 567. 1988. TYPE: Uapa- ca thouarsii E. Test, ed by Airy-Shaw, Kew Bull. 18: 1965]. А genus of ca. 60 species in Africa and Mad- agascar. Subtribe 6c. SCEPINAE (Lindley) SAL stat. nov. Ordo Scepaceae Lindl., Nat. Bot. ed. 2, 171. 1836. TYPE: Scepa Lindl. ^ni Aporusa Bl.]. 24: 594. 1975. TYPE: E: Áporu. sa onoecious trees or shrubs, stems n with lam- ubdrup 8 per fruit, seed-coat dry or fleshy; ا‎ thin 50 Annals Ка не Garden to copious 1 ly distinctly broad d Airy Shaw (1974) at the tribal rank. Since Lindley longer than radicle. published the Scepaceae over 20 years before the o Aporuseae was proposed, it seems preferable һ y Lindley to preserve his name. The seven genera of Scepinae (1836) as ап “ого, ” was adelaide Mueller are scattered through the tropics in America, Af- (1866) and later authors with the Antidesminae, ica, and Asia, with only Baccaurea attaining the and was first recognized in the 20th century by Pacific islands. KEY TO THE GENERA OF SUBTRIBE SCEPINAE la. Pistillate disk ba styles dilated, entire to slightly lobed 40. Protomegabaria lb. Pistillate disk cupular or absent. 2a. рык. inate disk k present. . Pistillate disk prema pistillate oe persistent in fruit. 4a. РЕ ious; flowe: аттана ules deciduous. a. Capsule lo Seal minate flowe aes one d ррасб на. ~. 41. Maesobotrya 2 ee ae насы us ral pam Draci. d гше 0 n 42. Richeria 4b. us; flow merules; picis nian capsule septicidal _ 43. Jablonskia 3b. Pil ate е disk absent райе Ын contra us. ous; 44. Baccaurea eris us; stole eine 45. Apodiscus 2b. Bago disk abse istillode massive, е, pelate stamens 5 or 6; carpels mostly 3 or 4; styles stigmatif __ 46. Ashtonia гв Pi tly 2 (rarely 3-5); carpels mostly 2 (rarely 3 or 4); wis BEA RTE 47. Aporusa 40. nk acer ee Hutchinson, Hook. I 866; Fl. Bras. "ig 13: Su: Bent TIBI Prop: Afr. 6). a Plantarum 3: 286. 18 ax & de pA Fa = Hoffm., Pflanzenr. 147. Hoffm., Pflanzenr. vci XV Ln 81: 26. XV (Heft 81): 43. 1922; Keay, Fl. W. Trop. 1922; Jablonski, Mem. New York Bot. Gard. Afr. ed. 2, 1: 373. 1958; Aubreville, Fl. For 17: 124. 1967; Philcox, Fl. Trin. Tob. 2(9): Cóte, d'Ivoire 2: 68, t. 147. 1959. TYPE: 638. 1979; WEBE Ann. Missouri Bot. Gard. Protomega itx ia үтел (Beille) Hutch. 75: 1093. 1989; Secco & Webster, Bol. Миз. (lectotype, selected her Para. Emilio Goeldi, Bot. 6 ОЛ? 158. 1990. Tuo species of West ХО. TYPE: Richeria grandis Vahl. Guarania wien ex Baillon, Etude Gén. Euphorb. 598. 41. Maesobotrya Bentham, Hook. Icon. Pl. 13: IA. re Girani ares o г 75, t. 1296. 1879; Gen. Pl. 3: 284. 1880; Rickert” Алыке (Baillon) Baillon; mee Hutchinson, Fl. Trop. Afr. 6(1): 663. 1912; a Pax & Hoffm., Pflanzenr. 147. XV (Heft 81): A genus of several ке defined taxa, treat 17. 1922; Léonard, Bull. Jard. Bot. Brux. 17: ав two species by Secco & Webster (1990); °x 256. 1945; Keay р. Afr. ed. 2, tending in lowland рам montane tropical sin Fl. W. Trop. Afr. 1: 373. 1958; Radcliffe-Smith, Fl. Trop. E. from Costa Rica and the Lesser Antilles o Peru Afr. Euphorb. 112. 1987. TYPE: Maesobotrya and Brazil. floribunda Bentham. Staphysora a Bull. So din дее 1233; 1896; 43. hens rary int Bot. Pax, Bot. Jahrb. 23: 521. 1897. Staphysora PE: Jablonskia conges га (Benth dusenii Pax [= г ка he (Pax) Hutch. ]. мык Arg.) Webster indes s Benth. ex Muell. Arg.]. can genus of ca. 20 species, all in West Africa ed only 1 species extending east to Ugan nda. Monotypic, the single species confined to The кн is extremely close to the neotropical A azonian South America. The genus is We in Rich subtribe Scepinae by its distinctive pollen t paracytic stomata and shows some similarities w 42. отта Vahl, Eclog. Amer. 1: 3 Celianella Jabl. in subtribe Antidesminae- Its ; Muell. Arg., DC. Prodr. ae dc placement requires further evaluation. — e Volume 81, Number 1 1994 44. Baccaurea Loureiro, Fl. Cochinch. 661. = 1880; Pax & m. V (Heft 81): 45. 1922; Airy Shaw, Kew bel 14: 353. 1960; acker & Bakh., Fl. Java 1: 453. 1963; Whitmore, Tree Fl. Malaya 2: 63. 1973; C. Smith, Fl. Vit. Nov. 2: 450. 1981; Thin, Cóng Trinh Nghién Cá'u Khoa Hoc 10: 78 1 РЕ: Ва 98 геа ramiflora Lour. Пес- totype, selected by Merrill, 1935]. Firari Roxb. ex Jack, Trans. Linn. Soc. London 14: u сое TYPE: Pierardia dulcis Jack [= Bac- a dulcis (Jack) Muell. T Bn Adererpis M Bijdr. 579. 1825. ТҮРЕ: Adenocre- a Bl. [= Ba T eau ed Jag (ВІ.) Muell. Calyptroon el Fl. Ned. Ind., Erste is 411. 1861. 7 Bac tro q ccaurea sumatrana ый? ) м Arg.]. d. Ind., Erste Bijv 444. 1861. diis wt Miq. I^ Baccaurea x Ac gl De (Bl. у ты J. Sci. Bot. 4: 279. 1909. E Bron digesta Merr. [= in. accaurea philippinensis [ue aie Gatnaia Gagnepain, Bull. Soc. gere 71: 870. 1924. TYPE: Gatnaia annamica Gagnep. [7 Bac- caurea oxycarpa Gagnep.]. tsified genus of ca. 75 paleotropical spe- cies, ы, from India to the Philippines, New Guinea, and de Pacific islands. 45. Apodiscus dern e Bull. Soc. Bot France 58, Mem 1912; Fl. Trop. Afr. 6(1): 1045. is Lab Icon. Pl. 31: t. 3032 15; Pax & Hoffm., Pflanz 147. XV (Heft al): 45. 1922. ТҮРЕ: lode cus chevalieri Hut poorly ssn monotypic genus from the байр. region of west Africa; said by Pax and Hoffmann to be d to Maesobotrya. 46. p Airy ge Kew pus 357. 1968; Hook. Icon. Pl. 38: t. 74; Анн Тгее Fl. "Mal laya 2: it edd y Shaw, Kew Bull. Add. Ser. 4: 42. 96 ptit Ash- tonia excelsa Airy Shaw orld genus of two species in Borneo and "n E peninsula. 47. Aporusa Blume, Bijdr. 514. 1826 e Apo- DC И» Jav. 1: уш. 1828; Mue Prodr. 15(2): 469. 1866; Bentham, ri РІ. Webster 51 Synopsis of Taxa of Euphorbiaceae 3: 282. 1880; Hook. f., Hook. Icon. Pl. 16 45. ТҮРЕ: Aporusa frutescens Blume Leiocarpus Blume, Bijdr. ge 1826. TYPE: Leiocarpus foe ae fines e [= orusa fruticosa (Blume) ani pia Lapis Wate ex 836. D ay Bi وو‎ dioica (Roxb.) Muell. rg.]. Scepa Lindley, Nat. Syst. ed. 2, 441. 1836. TYPE: Scepa stipulacea Lindley [= Aporusa dioica (Roxb.) Muell. Arg.]. A genus of 80 Old World species, from India Islands). Subtribe 6d. ANTIDESMINAE Muell. Arg., @ zh ЧЁ e ب‎ Ф Pax & Hoffm., Pflanzen. 147. XV (Heft 81): 3. 1922. TYPE: Antidesma L. Ordo Stilagineae C. A. Agardh, Aphor. Bot. 14: 199. 1824. TYPE: аса i Т. В ^ Antidesma Lj Vigo ong ies ponen 34: 64. 1865. E: Hyeronima Fr r. ioecious (rarely monoecious) trees or shrubs r nearly so, anthe nl connectives, anther-sacs usually ulous; pollen rai rolate, 3 orate, reticulate; pistillode b seeds ostiy beet r locule or init: sord: poat dry; ah thal but — exceeding radicle. ubtribe Antidesminae corresponds to the family Stilaginaceae as recognized by Ai qu tonem are not well understood, and their inclusion here must be regarded as provisional. Annals of the Missouri Botanical Garden KEY TO THE GENERA OF SUBTRIBE ANTIDESMINAE la. dupl or of = or stellate hairs. . Carpels 3- е "frui dehisc 3a. "Dis k pre esent; ovary 3. ocular. 4a. I I З, f. s I 4 in A £ s+} P 1 and stamin inodes. 5a. Leaves entire; pistillate flowers not bracteolate; fruits pedicellate __ 48. Thecacoris 5b. Leaves dentate; pistillate flowers bracteolate; fruits subsessile 49. Phyllanoa 4b. Pistillode absent; pistillate sepals accrescent; pistillate flowers lacking petals or staminodes _ ies 50. Celia cre ovary 4— жан Шаг 51. р fruit 52. Antidesma lb. ы sit See fruit dhien 53. Hyeronima . Thecacoris A. Jussieu, Euphorb. Tent. 12. 1824; Baillon, Etude Се orb. 605. ntham n PE: Thecacoris madagascariensis А. Juss Cometia Du Petit Thouars ex Baillon, Etude, Gén. Eu- phorb. 642. 1858. TYPE: Cometia ЙАЛТ Baillon = Thecacoris cometia Leandri]. TERON Mue Flora А 536. 1864. TYPE: Суа в. [= Thecacoris vir- Baccaureopsis Pax, Bot. 319. 1909. accaureopsis lucida Pax ах E Thecacoris ludica (Pax) Hutch.]. As here delimited, Thecacoris includes about 20 genus Mi is ea Muelle (1866) = де & Hoffmann (1922, dcliffe Smith 1987) epe usambarensis ande. bridges the gap between the two generic concepts. 49. peut Croizat, Caldasia 2: < 1943. Phyllanoa colombiana Croiza notypic genus from the Colo e tho чыкт resem- blance to Richeria, in subtribe 5сер 50. e ано, Mem. New York Bot. Gard. 1 1965. TYPE: Celianella mon- tana ts otypic genus of the lan G highlands The plant i is ditintive i in the semisue- Le (1986c) a it dies Hyeronima on the ied p foliar chara 512 — e A. Jussieu, d Tent. 19. 275. 1880; Pax 2; 198 M 8B: 212. 1957; Fl. Madag. 111(1): 1 Wn TYPE: Lepto- nema venosum (Poir. у A. poorly known genus of two species endemic to ils L., Sp. Pl. 1027. GAS ben iun Burman ex 988. Bestram Adanson, Fam 354. 1763. TYPE: Antidesma Pie La Stilago L., Mant. Pl. 16. 1767. TYPE: un. bunius L. [^ Antidesma bunius (L.) Sp er Rhytis шы, Pe Cochinchin. 660. . TYPE: Rhytis frutic Lour. aa ntidesma giantess (Lour.) Muell. J IE A large Old World genus of ca. 200 species species are Asian ex a few species in Actis and the Pacific islands. 53. dediti Mentes , Pl. Novas Bras. 1. ; Muell. Arg., DC. Prodr. 15(2): 268: "9v Spe riage Bentham, Gen. pl. E 284. ax & Hoffm., Pflanzenr. 14 . Jah E: Hyeronima WBE Allemao. Stlaginete Tul., 15: 240. eg PE: Stilaginella lesa du. ic Hyeronima ‘fe а (Tul.) Mue! g.l. | | Volume 81, Number 1 1994 A neotropical genus of more than 20 described ica, and th re a few additional species in the West Indies Subtribe 6e. ll Arg ) Kohler, Grana E жен 99. 1965. Tribe EER ue ‚ Bot. ае 22 . 1864. TYPE: Pistes Rudge Monoecious or dioecious herbs or subshrubs; ; Ь а ule; endosperm copious; cotyledons about as broad as radicle. wo closely mes dettes genera. Their Systematic position r controversial. Pax & Hoffmann (1 931) followed. Mue ller (1866) in plac- ing tribe Porantheae in the division “stele,” because of the narrow cotyledons. In 1975, I sociated амга we Andrachne, partly т nt (1962). However, pe her Sol ied e pollen of rond 4 was more compatible with an affinit ty to piso. complex, and he demoted etin to a subtribe of Antidesminae. Because he did not cite the basionym, his combination was not asd published, and it presumably is validated here Cause > о KEY To THE GENERA OF SUBTRIBE PORANTHERINAE 1а. Monoecious; petals and pistillode present .......... E EN ший 54. Poranthera LEUR petals and pistillode absent Buc н ыс oral 55. Oreoporanther a 94. к" Rudge, Trans. Linn. Soc. Lon- e 10: 302. 1811; Ben =ч Fl. Austral. 6: £ 1873; бы Pl. 3: 262. 1880; Griining, Pda s4 n 58): M 1913. TYPE: Как Теп Species from Australia, эз. Oreoporanthera Hutchinson, Amer. J. Bot. 56: 747. 1969. Po ranthera sect. Oreopor- anthera Grüning, Pflanzenr. 147. (Heft 58): 21. 1913 : CNPR ан alpina а боне К Monotypic genus endemic to New Zealand. tb t d Pointed ou Webster 53 d sd 28 of Euphorb Latin description), even trous he was obviously basing it on Grüning's sectio. Tribe 7. SEAM ПАР ү» An) Hutchin: Amer. J. Bot. 56: a aceae io ies Shaw, Kew мач 18: Léonard & Mosango, Fl. Afr. om menocardiaceae 1. 1985. TYPE: id j ley Dioecious trees or shrubs; indumentum of sim ple hairs nai sessil ала; aie bie alternate, aen 5 penninerved, sti axillary, spi cate, racemose, or paniculate; staminate sepals = 8, connate; petals absent; stam 4—8, filaments free or basally opem i Кы. de- hiscent; pollen суша сыш; pue fre capsular or winged, samaroid; seeds 1 per locule; endosperm scanty; cotyledons broader and longer than radicle. À distinctive group originally including a single Old World genus, considered to be a distinct family f moder a number of m n workers (e. d Léonard & Mosango, 1985; Meeuse, 1990). However, most oral and anatomical details , in Levin, 5 luded | altho u provisionally tinct from Hymenocardia in man perhaps be placed in a separate tribe or subtribe. KEY TO THE GENERA OF TRIBE HYMENOCARDIEAE la. Fruit capsular; inflorescence pgs styles lacerate 6. Di a 1b. et Fe inflorescence racemose; styles 251 ттен 56. раро Kuhlmann, An. Prim Sud-Amer. Bot. 3: 82. 1940. TyPE: sent mocistus esis Kuhlm nepi, genus of Amazonian Brazil. Kuhl- t Di Didymocistus may eda represent а a link r2 — eoe ex Lindley, Nat. Syst. ed. 2, ; Muell. Arg., DC. Prodr. 15(2): UA ipd Oliver, Hook. Ic. Pt 12:20 1481: ; Bentham, Gen. Annals of the Missouri Botanical Garden Pl. 3: 284. 1880; Pax & Hoffm., Pflanzenr. 147. XV (Heft 81): 72. I 2; Léonard & e, do gne Afr., Euphorbiaceae 57 menocardia punctata Wallich ex x Lindl. Samaropyxis Miquel, Fl. Ned. Ind., Erste Bijv. y» 1861. TYPE: Чоло ty elliptica Miq. [= menocardia wallichii Tul.]. An Old World genus of eight species, of which seven are African and fro m Madagascar. j one southeast Asian; absent Tribe 8. BISCHOFIEAE oe at) Huru- sawa, J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 6: 339. 1954. Piplisiese side Bis- choffieae Muell. Arg., Linnaea 34: 64. 1865. v Pu E (Muell. n. ) Airy dus Kew Bull. 2. 1965. ТҮРЕ: Bischofia pic tribe of a single Asian species, very 6a, b, c) supports retention in the Euphorbiaceae 58. nem Blume, Bijdr. 1168. 1826; Muell. Arg., DC. Prodr. 15(2): 478. 1866; Bentham, n. Pi 3: 280. 1880; Pa 1981. TYPE: Васи javanica Blume. oe Wight & Arnot, Edinb. New Pate sA Cd 298 O > 3 س‎ со دب‎ = Bischofia жаша Blume] 33. 1840 PE: iscus elias iA ) Be nnett i Nh trifoliata Roxb. = Bischofia javanica Blume]. A genus of a single species distributed from India and i Lanka to China, Indonesia, and the Pacific islands. INCERTAE SEDIS 59. ет, € Bull. Mens. Soc. uem s II. 1: 9; Gilg, Bae Jahr a 6. 1908; Dmitri FL pcr Afr. а 629. 1912; Рах & seins » fam. ed. 2, 19c: 46. 19 cus glaucinus Pierre. atürl. Pflanzen- . TYPE: Centropla- This genus is based on a single poorly known species from ч that was ns e a fla courtiaceous y by P e Eu- phorbiseese by tur (1908) ым e ou Pax & Hoffmann (1931) placed it in the Antides- minae next to 5 ondianthus, but its place within the Euphorbiaceae still appears uncertain. 60. Meborea Aublet, Hist. Pl. Guiane e 1775; Meu жыды b. : Meb Suriname 9, t. 1. 1931. rea guianensis Aublet This monotypic genus from French Guiana was referred by Mueller (1866) to Phyllanthus. Lan- gard antheae, but it cannot be identified with certainty Subfamily II. OLDFIELDIOIDEAE P th bster, J. Arnold Arbor. 48: 308. d Mes 595. 1975. TYPE: "ов fieldia В Auct. Plur x Benth. [= ài Paivaeusaceae Meeuse, Борик 1990. ТҮРЕ: Paivaeusa Welw. fieldia Benth. & Hook.]. Monoecious or dioecious trees, pa or sub- shrubs; leaves alternate ог more commonly 6| posite or setters, simple or palmately а сош- pot in or neo sent; inflorescences axillary, ae r etalous spines, шд е tam Us rarely dr ирас ceous); seeds , testa usually smooth à and ly obsolete ). hi: a 1! ( ys y copious (rare This most ien ien ed subfamily ™ cludes 28 genera with only own 0 species, whelmingly Cia in the a Непизрһеге. KEY TO THE TRIBES OF SUBFAMILY OLDFIELDIOIDEAE ic ee pre: disk extrastaminal, staminate nnular; Pte twice bifid; ovules ops EAE i Volume 81, Number 1 1994 lb. Petals absent; staminate disk dissected, intra- staminal or absent; ig entire or rarely once bifid; ovules anatro 2a. ds ws apertures idi: r; vessel elements at with scalariform pain rates (except in Wok илсе и. amin. ple; monoecious or dioe- cious trees, shrubs, or herbs; seeds mostly carunculate ....... CALETIEAE 3b. Leaves compound, or i ple then lamina stipellate or stipules adnate to petiole; dioecious trees or shrubs; seed: carunculate or e pülate 5 3,43 0^ iet: cp LE 2. PICRODENDREAE Tits, td CROIZATIEAE Webster, trib. nov. ark. E: Croizatia Steyerm Folia alterna integra; discus annularis; petalae 5, im- bricatae; stamina 5, antheris extror orsis; pollinis 0 mutropis; semina ecarunculata, exalbuminata loecious trees or shrubs; leaves alternate, en- introrse; pollen g r Spines conspicuous; pistill resent; pistillate se- pals and petals 5, imbricate; disk annular, glabrous; b m дема E broader than and equaling radicle. The tribe is E eue by the single gen Croizatia. This on shows clear affinities w ith the tribe Amanoeae in subfamily Phyllanthoideae, and it is more or less arbitrary dis. it is placed. 61. Croizatia Steyermark, Fieldiana 28: 308, fig. 57. 19 52; vi ods a & Steyer- mark, Syst. Bot 1987 TYPE: Croizatia neotropica iiie he. a genus of four кү дус m Panama о Venezuela and Еси T . ribe 10. PODOCALYC Hegt Webster, trib. nov. TYPE: Podocalyx Klotzs ud alterna vel opposita, integra vel serrata; rari us ê nullus; petalae nullae; stamina 5- ме. a €xtrorsis nae introrsis; Rima grana Webster Synopsis of Taxa of Euphorbiaceae 4-brevicolporata, echinata; ovarium 3- юеш, stylis in- tegris, ovulis anatropis; semina albumina Dioecious ше, у: сае or opposite, entire or orescences in thyrses or ebracteate cymes. Flowers apetalous, disk absent; staminate sepals (3)4 ae free or р, imbricate; pamens 9- 3. е; anthers 4 brevicolpor ate nt or ap echinate, tectum smooth; pistillode pres sent; pistillate sepals 4 or 5, imbricate, persistent; disk present or absent; ovary 3-locular; styles di- lated, stigmatiform; чэнне апа! торек Fruit ar or ous; ate; endos seien much broader t cap- r ecarun- osperm present; Merci plane or than radic This tribe includes three American genera that re different in many s re classified gether on the basis llen characters (polle eh with four shortened colpi) differe tween the genera are so striking that it seems necessary to assign them to different subtribes. KEY TO THE SUBTRIBES OF TRIBE PODOCALYCEAE la. аи ен free from petiole; fruit capsu ane "esit ign ecarunculate; leaves 10a. ; йш d iscrta; pistillode maa flowers solitary or clustered; seeds e; leaves alternate, opposite, or E ire or dentat N c pas lb. Peduncle adnate to the petiole; staminate se discrete; pistillode absent; fruit e ceous ... une ar RN Segun e 10a. PODOCALYCINAE Webster, . TYPE: Podocalyx Kl. Folia alterna, i integra; flores sinters entiers. extrorsis; LAM, ты gee planis. Trees with alternate leaves; stipules caducous; flowers in a con- ry L columella slender, kis ni оа та This subtribe includes only the single genus Po- ocalyx 62. haa erty Klotzsch, Arch. Naturgesch. 7 1841. Richeria sect. Podocalyx (Kl. Annals of the Missouri Botanical Garden Muell. Arg., DC. Prod eon nA 1866. TYPE: Podocalyx ARAS A monotypic genus restricted to Amazonian South America. Although P docalyx as a ES E аена its peer is completely 4 f the subfamily eee ioideae. Subtribe 10b. TETRACOCCINAE С. Levin, subtr. nov. TYPE: Tetracoccus Engelm. ex Par- utices dioicae; folia simplicia, alterna, opposita, vel vert cita dentata vel integra; рше 0; discus ê in M : trasta. lis, lobatus; st ‚ antheris Ts pollinis grana 4-porata, echinata; pistillodio 0; discu: glabrus, lobatus; styl if lari 1 © 23 d carunculata, albuminata, cotyledonibus plicatis. hrubs; leaves alternate, opposite, or whorled, кек toothe r entire; sti ipule es absent; stami- e one in ахау racemes ог panicl h f н ог asciculate, axillary; staminate sepals 4— б crete disk in trastaminal, lobed; stamen: 0, anthers ex- trorse; pollen grains е ectum verr ose tween spines; pistillode absent; pistillat 5-13; disk eo ovary (2-)3-4 e sepals – Pj ond styles elongated; fruit capsular; seeds carunculate; en dosperm a FERA cotyledons plicate. E E subtribe is very difficult to 14 2 Erunt. Ф toh Vegeta Térac coccus is intermediate betwee Podocalyx and Paradrypetes in phyllotaxy "e i e characters, such as the in- exine, inb isa а Соза; resemblance between Tet- tribes of Oldfieldioideae. Levin & Simpson ie odd refer the Tetracoccinae to the tribe Picrodendrea 63. Tetracoccus Engelmann e ni itin Scientist 1: 13. 1885; Croizat, Bul dias Bot. Club 69: 456. 1942; DM msi ra 56: 49. 1954. TYPE: Tetracoccus diia Par- ry. Halliophytum 1. M. Johnston, Mound Сга Mercer 68: 88. T TYPE: Halliophytum к АРА т (5. Wats. ) I. M. Johnston [= [ais occus pueris (S. Wats) Croat lectotype]. Аз шш һу Dressler hE a D of fo ica "possc California and adjacent Toma, southern Baja California, and northern Mexico) cina тер AEE Levin, Sys 1992. TYPE: Paradrypetes Mpeg. aves opposite, margin + spinose; stipules en- = n E E © ы” Ф © = Ф + Ф Б n - Ф 5 an ar; fruit drupaceous, 1-seeded; em- bryo with plicate cotyledons A monogeneric subtribe, eri Levin to be moy to subtribe Podocal 64. Paradrypetes Klis Ar 14. Ins - "m ral Jan. 2: 84. 1935 . 1992. TYPE: Parody Er "hm А genus of two South American species, one in the Amazonian rainforests, the other in the Bra zilian Mastic rainforest Tribe 11. CALETIEAE Muell. e Bot. 2: 3 190. а баг. е] A Б N o “Шш ч - > - = "VH e т л сот М t2 © — as © = دب‎ TYPE: OF Baillon [= Micrantheum Dest. Dioecious or monoecious trees ог EA M alte a s opposite, or verticellate, simple, or dentate, sti ipules obscure or suet flowers E se- al - Р mens 4-30, id cS ad Bui seeds dial per coccus, carunc ciliis d unculate; endosper ually copious; cotyledon i ёл. broader ог л broader than the radic This variable tribe of 4 subtribes with 15 gent? is entirely estoy and Hyaenanche, the gen are nearly со Indon KEY TO THE SUBTRIBES OF TRIBE CALETIEAE Bre whorled pw least in part); filam pus dics anthers; pollen tectum gan апсы - us iced lar D. ves кы uk or opposite; Е pe than anthers; pollen tectum smooth. 6 Dude e PE КАЛ lb. indeed except fot nfined ® — у | ш Australasia (with a Bes навела of Austrobuxus esia). і { | Volume 81, Number 1 1994 Webster Synopsis of Taxa of Euphorbiaceae ciduous or reduced; leaves opposite or if apa ipen capsule fleshy. 3a. Leaves opposite; stamens fre: thers н а capsule a bi . Leaves alternate; stamens connate; th 1 lat le flesh 5 i T А г КО tillate sepals deciduous .. { Pollen grains pantoporate; stipules dec or persistent; leaves oe nate; е Tu KETA отити pese lla. HYAENANCHINAE Bailon ex неч Arg., Linnaea 34: 64. 1865 (as Hyae- ae); DC. Prodr. 15(2): 479. 1866. Tribe ws ir (Baillon ex Muell. Arg.) Hutch., Amer. J. Bot. 56: 746. 1969. TYPE: кдн che Lambert Toxicodendrinae Pax, Natiirl. remm. ed. 1; 3(5) 3 1. 1890; Pax = Ho atm.» Pflanzen WS белде Lambert]. us trees or shru vericilate, Î ne ns ary clus minate sepals 4-8(- "pun stamens 8-30, fila poi anthers introrse or extrorse je: grains or 7-zonoporat ose 11b. Dissiliariinae etalostigmatinae id- Cap- 114. Pseudanthinae bs; leaves opposite or obscure о or in frui absent; ovary 3—4-lo €; endosperm spa ubtribe is here circumscribed to include Pholo clear whether the two genera should be Sie together 65. n che Lambert, Descr. Cinchona 52, САТ нан 68, m Thunberg, Kongl. M Acad. Handl. Bills. b (non Toxicodendron P. Miller, 1754); P Gen. Pl. 3: 280 dee Pax & Hoffm E 47. XV (Heft 81): 284. 1922. TYPE: tilode absent; ай nidi 3-8, free; deciduous t; disk PP sericeous; ks xtti + dilated; fruit capsular; seeds runculat rse. Toxicodendrum capense Thunb. [= Hyaenanche &lobosa (Gaertner) Lambert]. £c Tes TUN (d pm rica. The des bof Hyae базавага мару: with- in the Oldfieldioideae in “he scan uy endos speci (Hyaenanchin). iN IIb. a E Pax & Hoff- nn, Pflanzenr. 147. XV виб 81): 288. 1922.1 YPE: dete F. Mue = Monoecious or dioecious Hoda or Y shrubs; leaves opposite, simp e, absent; Abos 1 in ers panicles ог PT staminate sepals 4-6 v ricate, free; disk a Bn stamens —50 iii + tectum smooth; pistillode absent; ite sepals 4-6, free; disk cupulate or abse (rarely 4- Jocular, glabrous; rg inl fale cap- sular; seed culate or ecarunculat This Australasian subtribe includes 6 genera with a. 25 species. Generic limits require further crit- idi tudy. КЕҮ TO THE GENERA OF SUBTRIBE DISSILARIINAE la. Pollen exine with elongated spines; pistillode mostly absent. 2a. Dioecious; stamens 8-25; pistillate disk cu- ular o facra b 3 h fimbriate caruncle; ovary 3(= diera, styles ovate, erect .......... 66. Austrobuxus 3b. Seeds ecarunculate 4a. Styles elongated recurved; ovary 3(-4)-locular ................ 67. Dissuliaria 4b. Styles loc ovate, erect; ы 8. Сапаса 2Ь. Monoecious; stamens ¢ ca. i. 50; pon disk 9. Whya "pon lb. Pollen exine with reduced spines; pistillode pre 5a. Stamens 4- 6; styles Край at base, г cca D ud 5b. Stamens 8-17; styles Ee not re- urved; seeds carunculate ........... 71. Longetia 66. — (€ Miquel, Fl. Ned. Ind. Suppl. 1980; A. C. Smith, Fl. Maven, E 495. 1981; McPherson & Tirel, Fl. Nouv.-Caléd. dad 193. 1987. TYPE: Pd eps nitidus Miqu пее А Baillon, Adansonia 11: 83. 1873. ТҮРЕ: Bur- avia carunculata (bon) Baillon [= Austrobuxus Choriophyllum Bentham, Hock. Icon. Pl. 13: 62, t. Annals of th RE س‎ Garden 1280. 1879. TYPE: Choriophyllum malayanum Bentham [= Austrobuxus nitidus Miq.]. the circumscription of Airy Shaw (1974) and Mn & B uel (1 pesi Us Austrobuxus is the dfieldioideae, with ca. 20: species. ШЫ distribution is disjunct ‚ with areas Queensland New Coa. and Fiji, but absent Vin New Guinea. 67. € F. Mueller ex Baillon, Adansonia 66. 1867; Bentham, Fl. Austral. 6: 90. Je Pax & Hoffm., Pflanzenr. 147. X (Heft 81): 291. 1922; Airy iuh Ma 1980. a baloghioides т ell. ex Baillon Que a selected here; the generic name was associated with that species in Bail- lon's original description] E to Airy Shaw (1980), Dissiliaria in- to Australia (Queens- land). 68. Canaca Guillaumin, Arch. Bot. Caen 1: 74. 1927; Airy Shaw, Kew Bull. 25: 508. 1971. TYPE: Canaca vieillardii Guill. ted by Airy Shaw (1971), Canaca was Guil- th laumin but recognized as related to trobu. by McKee. McPherson & Tirel (1987) followed Air aw in relegating Canara to syn Austrobuxu owev i as distinct ге Ышш, it maar that Can- is eq lly g possi ible P Mete ee Lin US lak the g name Austrobuxus. 69. Whyanbeelia ae Shaw & Tay Kev Bull. 31: 375, 1976; Kew Bull. 35 691. 1980. TYPE: за terra-reginae i Shaw & Hyland. monotypic genus of tropical Australia eee 70. eet ee Baillon, Adansonia I. 11: 119. 3; Ai B eae Р e A ceras australiana Baillon [= C Кога» tricorne (Bentham) Airy Shaw]. f two species restricted to tropical ym aes Airy Shaw мы, кагу out its Sou. uishing charact T1. EREIN Baillon ex Muell. Arg., DC. Prodr. (2k 1866; Baillon, Adansonia 1. rel, Fl. Nouv. -Caléd. 14(1): 1 Longetia buxoides Baillon ex Modi As interpreted by McPherson S Tirel, Ln is a monotypic genus endemic t w Caled Pax & Hoffmann р, н ene ша broad sense, includin cies now placed in Aus trobuxus. Airy Shaw “1971, 1974) held a similar broad view, Mn that Longetia species were now transferred to Austrobuxus. In view of the distinct difference in pollen, McPherson's return t o the classical concept of Baillon and Mueller seems m justified. Subtribe I lc. PETALOSTICMA S Pax V (Heft 81): m.) Webster, Taxon 24: 595. 1975. TYPE: hey F. Muell Dioecious trees or shrubs; leaves alternate, sim- eee entire; eee WE RUN flowers in axillary clus 4:6, imbricate; di 7 sant = 20-40, ‘fi к grains with short spines; pist tillode o ob sent; ieee sepals 4-8, imbricate; disk e | ovary 3-4-locular, pubescent; — dilated; fruit a x capsule; seeds carunculat nogeneric subtribe restricted to Australis — sd Bow Guinea (Papua | 1922; Ау Shaw, Kew Bull. 3 : Kew Bull. 35: 661. deum TYPE: Peal quadriloculare F. Mue As treated by Airy Shaw (1980), Peal | is a genus of six variable Australian id есте І of which extends to New Guinea (Pap Subiribe 114. PSEUDANTHINAE "m | MEN Linnaea 34: 55. 1865; DC. Prodr. 15( . 1866. TYPE: рү Sprengel- Monoecious (rarely dioecious) trees shrubs, re 4 herbs; leaves mostly alternate, entire; stipules 1 Volume 81, Number 1 1994 Webst Syn ок of Таха of Euphorbiaceae ciduous or persistent; flowers i in axillary mia epals 3 i ula styles unlobed, iat or йерей: fruit capsular; 0 carunculat The subtribe Pseudanthinae ence s here circum- m ppe eros with the anatomical си palynological eviden KEY TO THE GENERA OF SUBTRIBE PSEUDANTHINAE la. eire broader than radicle; trees R a . Staminate e 4; pou NUTS 3. Kairorhamnus 2S За. Gres І e e грешне present staminat nce racemose ШЕЛ: ae conspicuous but deciduous . . Scagea iden 2 per locus ias absent minate flow етим stipules ШЫ ог roe w = hi “Sip foliose; E pollen grains һ lon ng е xino spi . 76. Micrantheum 4b. Sti seeded: pollen grains with reduc ed's spin 5a. Stamens free; pistillode present ыст 11. Pseudanthus 5b. Stamens connate; PN absent 8. St tachystemon 73. Coon Airy Shaw, Kew Bull. 34: rn 0; Kew Bull. Add. Ser. 8: 121, t. 1 1980. т YPE: Еи phyllan- т, E ipee Airy Sh w Guinea (Mo- iuda te-acuminate an e sug- E. N è = ct). The Ded cau еауез of this plant indeed g dle of Phyllan the thus, as p b y Shaw tits pollen is typical of the Онаа ана §enus appears very close to Austrob 14. Scagea McPherson, Bull. Mus. Nat. Hist. at. Paris, ТУВ. | Бы 1985; Fl. Nou Caléd. 14(1): 90. 7. ТҮРЕ: e Pauperata ы, Жор Peri 9f two species endemic to New Cale- ` “Me genus was originally referred to sub- family Crotonoideae by McPherson because of the uniovulate carpels, but the cle й о ment in the 'Oldfeldioideae. The very simil Choriceras, but pesa in By. ica r to vule number ed pollen morpholog 5. Neoroepera Muell. Arg. & F. Mueller, DC. Prodr. 15(2): 488. 1866; n Fl. Aus- je 6: 116. 1873; — bg 276. 1880; y Sha w Bull. 3 . 1980. TYPE: Peces buxifolia bd т; & Е. Muell. A genus to two species endemic to Australia (Queensland). 76. мич Desf., Мет. Mus. Hist. Nat. ‚ 14. 1818; Muell. Arg., DC. EA iso $n 1866; Bentham, Fl. Aus- tral. 6: ЫР, 95:202. 1890; Grüning, read 147. (Heft 58): 23. 1913; & Hoffm., Natürl. Pflanzenfam. ed. 2, TYPE: Micrantheum eri- chides Desf. Caletia ape Etude Gén. Euphorb. 553. 1858. alet pati ides Baillon, nom. Шер. [= ook. f.]. use oe Proc Ros . Vict. II. 22: 7. 1909. E: Allenia blackiana T Шер. [= Mi- ntheum dem ll A genus of three Australian species. TYPE: Mi- cra Mee kerea Sieber ex Sprengel, Syst. Veg. b DC. Prodr. Pflanzenr. 147. (Hef Pseudanthus pimeleoides Seber ex Scares Chrysostemon Klotzsch, Pl. Preiss. 2: 232. 18 YPE: Chrysostemon virgatus Kl. [= Ps nik to vir- us (К1.) Muell. A horizotheca Mue Arg., nnaea 32: 76. 1863. TYPE: e izotheca micra ieoidos Muell vus [- Pseu- anthus virgatus (Kl.) Muell. Arg An Australian genus of seven or eight species. 78. CLINT 1 Hook. London J. I5: ell. Arg., DC am. ed. 1. TYPE: Wc КУЛИН» ver- süculre Planch. A genus of three species in western Australia. Annals deni P RS Garden ha 12. ее Dre Web- , Taxon 24: 1975. Family Picr беба Smal " "New Yor k Bot pese 18: 184 . TYPE: oec catum Plan- chon. Monoecious or dioecious trees or shrubs; leave alternate or opposite, 3—9-parted (rarely куы late), entire; stipules caducous, fused to petiole, or sent; fl i i s ers in axillary cymes, racemes, - rules; staminate sepals 4—8 (or olete), free; disk int inal or absent; stamens 4—50, 3 anthers extrorse introrse; pollen grains sphe- roidal, brevicolporate or porate, with ines, ectum granulose; pistill te or is- tillate sepals 4—8, portii or deciduous; disk present or absent; ovary 2—3-locular; dn elon- ceous; seeds carunculate or ecarunculate sperm usually copious; cotyledons каг ап a longer than radicle. is tribe of ten genera in three see lk con- trasts Ке with on Саза. in its gu rican Madagascan = one ee (Mischod m in Sri Lank well characterized by its compound ts е the leaf blade ap- pears unifoliolat te in Mischodon, some species of Androstachys, Rn Parodiodendro KEY TO THE SUBTRIBES OF TRIBE PICRODENDREAE la. ee caducous ог obsolete, free from petiole. caps = posed crodendrinae 2b. Staminate sepals connate; apanle loculi- b. Paivaeusinae lb. Stipules рене. + adnate " petiole; seeds ecarunculate we c. Mischodontinae mages et donee e (Small) yiip nov. Family iori am зу id ork Bot. s 18:1 1917 ТҮРЕ: А Planchon. Dioecious trees or shrubs; leaves alternate; leaves 3-foliolate or 1-foliolate, margins entire; sti ecarunculate e four American genera of tribe Picroden- dreae are all referable to this subtribe. KEY TO THE GENERA OF SUBTRIBE PICRODENDRINAE la. Leaves они 3-5-foliolate. 2a. Fruits dehiscent; staminate sepals not re- "i P4 ИРС upto EEE тат: ОАЕ oe 3b. Stamens 8-15; ae vel race- jose 80. de 2b. due eh die drupaceous; staminate 81. ГЭ 1Ь. bid iy diis) жыйуу yes pals foliose, persistent. — orodiodendill 79. Celaenodendron Standley, Contr. Dudley | Herb. 1: 76. 1927. TYPE: Celaenodendron mex icanum Standl notypic genus of western Mexico (Tres Ma- rias faa Jalisco 80. Piranhea Baillon, Adansonia I. 6: 235. 1866; M Pflanze ‚147. ху (ней 81) 295. 1922; Ja blonski, Matic w York Bot. Gard. 17: 121 1967. TYPE: 6 Sei муы, иш Baillon. A genus of two species endemic to South America. | 81. ede Planchon, Hook. Londo i cons.): Picrodendron baccatum (L.) K Urban onotypic genus endemic to the Wes t Indies (ioi Antilles, Bahamas, Cayman Is Islands, Swan Islands). 82. Parodiodendron Hunziker, Kurtziana 5: 33l. fig. 1. 1969. TYPE: : Parodiodendron margin | villosum (Speg.) H | A monotypic genus of northern Argentina. Subtribe 12b. PAIVAEUSINAE P шаш Pa 14%: ху (кар “204. X TYPE: Paivaeusa 1 am [= I arti Benth. & Hoo! | Dioe trees; leaves alternate or opposite, | 8- а stipules dede staminate infloresc rescence _ nrn E — cymes, the pistillate ш y ally reduced to 1-3 flowers; staminate e sepals 5 ‚А basally ма yen intrastaminal, lobed; E Volume 81, Number 1 bster 61 1994 Synopsis of Taxa of Euphorbiaceae 4-12, free, абад ехігогѕе; pistillode minute or 1Ь. Leaves opposite; staminate disk ог pistillode (or — pistillate se —8, persistent; disk annular; both) absent. — 2-3-loc учы сн dilated, lobed; capsule tar- 29: weit filaments elongated, free; leaves е dily loculicidally dehiscent; seeds carunculate. Ja. жесен disk absent; pistillode present; ree В ; Д tillat — 85. Mischodon i yc gegen i in agreement Es the stud 3b. S е disk pr esent; pistillode aheent; D aivaeusinae includes only the biu sepals accrescent in fruit ___ single African genus Oldfield x & Hoffmann 86. Voatomalo к= со which appears better placed | in Mischo- don 83. Oldfieldia Bentham & Hooker f., Hook. J mith, Fl. Trop. E. Afr., Euphorb. 114. 1987. TYPE: Oldfieldia ndm Benth. & Hoo Paivaeusa Welw. ex Benth., Gen. Pl. 1 #5 ТҮРЕ: TYPE: Cecc на ена Milne-R by Léonard (1956) and Radcliffe- si ew Oldfieldia includes four African spe- * The distinction in phyllota asy (Не уз. a posite leaves) used by Pax & Н Paivaeusa is clearly unworkable, because Eines and opposite leaves occur in both Paivaeusa d Oldfieldia Webster, ee 24: 595. Mischodon Thwa Androstac hyda ei iit Kew Bull. 18: 250. 1965. TYPE: E 1975. TYPE: shrubs; 1 or 3- бенен е, entire; xilla pistillate sepals 5 or 6, ; disk annular or omen ovary 3-4(-5)- NR glabrous or pubescent; styles slightly to dis- tinci] Y connate below: fruit capsular; seeds ecarun- Culate, Key TO 3 ада GENERA ОЕ SUBTRIBE MISCHODONTINAE Ves alternate; pistillode presentan iu ee BG, Aristogeitonia 2b. Stamens with filaments united into * long vue anthers subsessile; leaves 1- 7-fo- iolate. Да. Leaves unifoliolate; fruit esr pressed же рий 4b. Leaves 3-7- foliolate; fruit А 88 NOTRE 84. Aristogeitonia Prain, Kew Bull. Misc. Inf. Radcliffe-Smith, Kew TYPE: Aristogeitonia уон doe Pris бимо Leandri, Bull. Soc. Bot. France 85: 231 1939. TY РЕ: Paragelonium perrieri i Leandri [= Aristo- geitonia perrieri (Leandri) Ra m.]. Ап African genus of three species (Angola, Kenya, Tanzania, and Madagascar). Radcliffe-Smith (1988) has recently demonstrated that Paragelonium from Madagascar cannot be separated from the mainland African Aristogeitonia. 85. Mischodon Thwaites, Hook. J. Bot. Kew Gard. Misc. 6: 299. 1854; Muell. Arg., DC. Prodr. 15(2): 1124. 1866; Bentham, Gen. PI. 3: 280 Fl. Br. Ind. 5: 344. 1887; Pax E: Mischodon муйнын Thwaites. monotypic genus of Sri moe perhaps native (but very rare) in southern Ind 86. Voatomalo Capuron ex Bosser, Adansonia II. 33. 1976. TYPE: Voatomalo eugenioidea Capuron ex Bosser This of two species from Madagascar was Scat hy M dri (1958) but not formally pub- lished until 1 87. Androstachys Prain, Kew Bull. Misc. Inf. 1958; Airy Shaw Adansonia II. 10- 519. 1970; Dyer, Gen. 62 Annals Missouri айй Garden Afr. Fl. Pl. ed. 3, 312. 1975; Leroy, С. R. leaves alternate (rarely opposite), usually stipulate; Acad б Paris 283D: 147. 1976; Alvin, Ann. blade simple or palmately lobed (rarely compound) Bot. 59: 579. 1987. TYPE: Androstachys john- often with f glands. Inflor ces axilla sonii i Prain [= Androstachys subpeltata (Sim) inal, mose or DOM or еш e H- erules or ary fl А : ics Staminate ая мег: ѕера ls imbricate o or often valvate If Stachyandra is recognized as a distinct genus, ( and then often е d rupturin at айй бреу А аз becomes a monotypic genus confined 2100, to eastern Afric 88. Stachyandra Leroy ex Radcliffe-Smith, Kew Bull. 45: 562. 1990. TYPE: a чы mera- na (Airy Shaw) Leroy ex Radcl. enus of four Madagascan species has re- el ght propriate to treat Stachyandra as a subgenus o Androstachys. поран Ш. ACALYPHOIDEAE Ascherson, FL Pr тапдепЬ. 1: 58. 1864. Family Aca- lyphaceae e Kio tzsch & Garcke, Monatsber. Kön igl. Preuss. Akad. Wiss. Berlin 1859: 246. 1859. TYPE: E rees, shrubs, or herbs; milky latex absent wowed present, or a by colored exudate); in tum simple or hairs often branched or KEY TO THE TRIBES OF SUBFAMILY ACALYPHOIDEAE free or содае: pollen grains Bi 3-4- саа exine semitec tate (very rarely echinate); male ga- ee s ucleate; pistillode present or 16 Pistillate ove: ве pal. (2-)3-6(-12), а open at an p. or pens ey шу 2; 4- ocular; a (except їп Dicoelia), anatropous. Fruit ca apsular or rarely baccate or drupaceous; e саги а or ecarunculate rm ust- ally copious; cotyledons longer and ‘broader than rad hi rs mostly 9, 10, or 1l. icle. Base chromosome п 116 genera in 20 tribes, subfamily Acaly- rd is the largest and most ec of the s of Through Clutia it ap- roaches the Phyllanthoideae, m rough alea both the Crotonoideae an En Euphorbioideae multiple connections s that the e suggest phoideae may not be a monophyletic group. 1 f 1, indumentum la. ] sm or ago d 2a. t adaxially barbate. linens connate; disk segments glandular-lobed; seeds carunculate; r llucid-punctate, i tum simple 25 stamens. F © leaves ` CLUTIEAE © indum р 2Ь. Petals adaxially barbate; vw free; disk segments not glandular-lobed; seeds шш. leaves not pellu ucid- -punctate 4. POCONOPHOREM | lb. S itl bri pals and p d 5 st (or if so, then pollen not perforate-tectaté f or fruit not capsular). 3a. EET Yu in | 1: р 1 ` 1 a petalous; docu trees or shrubs. g 1 eine 5a. Fil Я pi ‘ad т 1 3 ulate, testa dry. i 6a. Flowers in axillary glomerules, not involucrate; stamens 8-15; styles pene bifid; capsule echinate; leaves stipulate 15. CHAE 6b. Flowers enclosed in ed; bibracteolate involucres; stamens 2-6; styles дка E og capsule not echinate; leaves exstipulate 5b. Pene free; pistillode small or absent; pollen tectum spinulose; seeds IE ПОРА | 4b. Leaves oj te; poll i ticulat d 18. dac 3b. Staminate se о ог е beg tals present or slightly i _ pollen grains various, not micropunctate; indumentum various. EC T Ovules 2 per das ee capsular; plants monoecious; styles unlobed . Ovule 1 1 es 1 per осш ruit drupaceous; plants dioecious; s tyles bifid ______. 2 per 9a. Cotyledons cary broader than NON Minas бш dioecious) ae е lai Se as TAMPERE k gut C tulad, E ҒО 1 d : y у E 1 dicle; h t Volume 81, Number 1 Webst 63 1994 Syn x of Taxa of Euphorbiaceae 10а. Petals present, at least i lla. Dioeci ind i anther sacs pendulous; pollen sexine coarsely еа пої eraat РЯ AGROSTISTACHYEAE llb. Monoecious (rarely edes yd indum tum (at least i in par art) bris, мс» not 23. CHROZOPHOREAE 1 ап! еп sexine often plebs rare her iube f h (£1 E Lan AT 10b. Petals absent. 12a. pr Jui. inci scandent; stinging hairs absent e disk present; pollen exine coarsely reticulate, colpi mostly pte Leaves usually with laminar glands; ind impl stel- Я ill erminal. 154 Pollen pu not marinate staminate disk massive, pu- bescent 24. CARYODENDREAE 15b. Ройеп colpi marginate; staminate disk tenuous, glabrous; 25. BERNARDIEAE 14b. Leaves eglandular; indumentum simple; stamens many; inflo- 26 PYCNOCOMEAE 13b. Staminate ier b ( if prese 16a. Pollen grains coarsely rece or prot -tectate; indumen- tum stellate; stamens inflexed in bud (ex чайкасан monoecious; pistillate calyx Ду М. carunculate т, 7 BPIPRINEAE 16Ь. е ко cim ето отав to rugulose; іп stellate; ot inflexed in bud. ps E ollen kar Tex prelate colpi; staminate disk prese т absent: mostly ecarunculate, testa not foam stipules deciduous or d te. Leaves fse dular; styles subentire to — HS sta: minate disk (if present) extr. astaminal oi inal; pollen sexine finely perforate- T jii ds cape M or ecarunculate, testa or flesh isexual; seeds p 19a. Staminate sepals valvate, splitting apart at anthesis; latex nd in- florescence not paniculate; bracts 4 31. PLUKENETIEAE 19b. Staminate sepals imbricate; latex clear or reddish; inflorescence pa pes bracts foliose and biglandular; stamens 2 or 3, rad е; pollen grains colporate, exine rugulose _ 32. OMPHALEAE Tribe 13, bes (Muell. Arg.) Pax, Natürl. imbricate; in 5, free; disk distccted, €— 5, Pflanzenfam, ed 3(5): 81. 1890 (as Cluy- + palmat filaments ы, nan m luytieae Muell. d conl 34:2 1865. Cluytieae subtribe late, 3- irme, gem ор ont slightly n í “onthe Ld 47. III (Heft 47): distinctly margina 1911. apes Clutia L. present; ovary 3- candy + pa glabrous or pu- Dioecious besoa nt; styles ae so, bifid; fruit араз shrubs; ind о" monoecious) perennial herbs or : eS А ur punctate), testa nate, simple, entin imple or absent; leaves alter- dry; endosperm copious; cotyledons broader. and » enlre, penninerved, often pellucid-punc- ilice: longer than the radicle. tate, egland RE ular; stipules small or obsolete; flowers in glomerul rules (pistillate often solitary); sepals 5, As here interpreted, the tribe Clutieae is restricted Annals of the Missouri Botanical Garden to the type genus Clutia, in accordance with the original "xpo of Mueller. None of the genera as- pia be шш ve ү (1 BR Ја аге Clutia in tibe tribe sp еза of its ne жа! їп habit Baillon (1858), pee clea rly demor ed atu pels, aside the ee ee to Briedelia and ice eto to be ocior and suggested an af- finity with C ever. е nan struc- ture ue not suppor Вай 5 view, and i in fact is phi ginal suggestion of a тешип vit the Phyllanthoideae. 89. Clutia 8 Sp. Pl. 1042. 1753; Gen. Pl. ed 5, 464. 1754; Muell. Arg., DC. Prodr. 15(2): 1043. e не, Сеп. Pl. 3; 302. 1880; ах, Pflanzenr. 147. Ш (Heft 47): 50. 1911; Hutchinson, FI. ri Afr. 6(1): 801. 1912; Gen. S. ith, Fl. E. Trop. Afr., Euphorb. 1: 331. 1987. TYPE: Clutia pulchella L. A large Afri o 4. AC 75 species, with tw Y Arabia; the greatest Pisae ds is in South Afia 14. Tribe POGONOPHOREAE (Muell. Arg.) Webster, Taxon 24: 595. 1975. Hippomaneae subtribe Pogonophoreae Muell. Arg., Linnaea 34: 202. 1865; DC. Prodr. 152): 1040. 1866. TYPE: Pogonophora Miers cious trees; indumentum simple or (in inflo- e lenses aceous; ps alternate, simple, ms rved, egland He very = s; staminate sepals 5, coriaceous, br oadly nd (st: uibs petals 5, imbricate, MGR barbate; disk intrastaminal, urceolate, 5-lobed; mens 5, filaments |, capsular, ilum large; endosperm copious; piene much dua and broader than radicle. The эхе ена is unigeneric, with the ription as that of Mueller (1866). The n Mi anen Microdesmis, and it has even been claimed that the membership of the taxon in the w Euphorbiaceae is suspect. Airy Shaw (1966) sug- gested that it may mo ne allied to the Icacinaceae. Howe to niga Icacinaceae are probably superficial. 90. الیو‎ ex Bentha: . Kew Gar er. Sul, TYPE: Pogonophora pe | "s Miers ex Bentham A genus of two species, one neotropical (Атал? nian and coastal Brazil), the other west African. Tribe 15. CHAETOCARPEAE (Muell. Arg) Webster, Taxon 24: 595. 1975. Hippomaneat subtribe Chaetocarpeae Muell. Arg., Linnaea ea 34: 202. 1865. TYPE: t Chaeocarpus Thwaites. h imple oF Dioecious trees or sh rubs; i absent; leaves alternate, ims се) Eve б ipules deciduo А xm Hi co MR stamens grains 1 i si exine ose and micropunctate; pistillode minent, hirsute; pistillate sepals 4- ‚ imbrical® disk urceolate; ovary 3- БОЙЫ, dB bipartite P К | pillose; fruit capsular; seeds sm th, blackish, car , | culate; endosperm copious; broader longer than radicle. taxon, a cluding only the type us, was enlarged by Webster (1975) ire i chien of UNT The close | the roe ш "e two genera see indi The blackish carunculate are reminiscent of those of Clutia. There may de heilosa, a8 80 independently sugges Chaetocarpe2' Tri formally published) м9 Chaetocarpus and E gonopleura, Run saw a resemblance to Caseari? the Flacourtiacea | КЕҮ To THE GENERA OF TRIBE ARRA | 1 - tals present; ne ly sessile on staminal a. d йазы нона, ап! nearly 91. Trigono noia | |] Volume 81, Number 1 Webster 65 1994 Synopsis of Taxa of Euphorbiaceae 1b. нк absent; filaments well PNE ovary ec ers apetalous, disk absent; staminate c. 2-4-fid echi- 2. Chaetocarpus 91. Trigonopleura Hook. f., Fl. Brit. Ind. 5: 399 87 Bull. 36: 350. 1981. TYPE: 5 Таор та- бена Hook. f. otypic genus of southeast Asia (Malaya to Ф Philippines and Borneo). 92. Chaetocarpus Thwaites, Hook. J. Bot. Kew : 76. 1973 aeto pungens Thwiiités [= Chaetocarpus castan carpus (Roxb.) Thw.]. Mettenia Grisebach, Fl. Brit. W. Ind. 1859. TYPE: Mettenia globosa Griseb. bim а дод globosus Griseb.) hg с. & Re 1 b. асам . 1861. TY FA naldia ну eom js кебу castan carpus (Roxb.) Thw. [secre Beille, Compt. Rend. Hebd. Acad. Sci. cvs 1907. TvPE: Neocheva ims ae Beille [= Chaetocarpus peo Pax]. À genus of 12 species with a ct distribution West Indies, South America, west id ike Mos саг, and tropical Asia. Tribe 16. PEREAE (Kl. & Gcke.) Pax & Hoff- с 147. ХШ (Ней 68): 1. 1919. Регасеае КІ. reuss. A am. ed. 1, 3(5): 69. 1890 TYPE: Pera Mu lineae Klotzsch, Arch. à TYPE: pae sch, Arch. Naturgesch. 7: 176. 1841. a or shrubs; indumentum simple, © te, ог d leaves alternate (dis opposite), = pinnately Flowers in ax- veined, exstipulate. ilary i dal valvate сег us glomerules; involucre usually of 2 tirely enclosing flowers in bud. Flow- 3-locular; rosea stigmatiform. Fruit capsular, valves woody; seeds smooth, dark, биш: endosperm copious; es much longer and broader than adicle. is tribe, by the unanimous consensus of workers since x ha includes only the single genus Pera. The relationships of Pera have been in doubt, how- ever, because of the reduced nature of the Pup and the peculiar inflorescences S have n oun- жй 1 Pera at the "id. of the Acalypheae (Acalyphoideae), Pax & Homa pigi pde iine Airy Shaw (Dict. Fl. Pl. ed. 7, x ее mia Peraceae as a distinct family, oed of its affinities. I believe that the T matin indicated J . Li $1 1 1 | 7 E. 1 1 J Te J F with smooth carunculate seeds. 93. Pera Mutis, Kongl. Vetensk. Akad. Nya Handl. { 15(2): blonski, Mem. New York Bo 1967. Perula Schreber, Gen. Pl. 703. 1791. TYPE: Pera arborea Mutis Spixia Leandro de Sacrameto, Denkschr. Kó n < Wiss. München, Math. Arch. Naturgesch. 7: 178. 184 41. ismatopera martiana Kl. [= Pera disticho- phylla (Mart.) Baillon]. А neotropical € of ca. . 30 Mee wei from Cuba and Cent Brazi the um concentration of taxa in the ue basin. [= Pe Schismatope era TYPE: Schis E zd CHEILOSEAE (Muell. UE ) Airy Shaw Webster, Taxon 24: 595. 1 ippoma- subtribe Cheiloseae Muell. ay Linnaea 3 "202. 1865. ree subtribe Chaetocar- pinae series Cheilos es Pax & Hoffm. Pflanzenr. 147. XIV а 68): 50. 1919. ТҮРЕ Cheilosa Blume. Dioecious trees; indumentum simple, stellate, or Annals eis н. Garden dé асан нз ог ec iode. caducous; inflorescences ary or sometimes ter- inal, paniculate, bracts eglandular; flowers apetal- us; sepals 4 or 5, imbricate; staminat cted t introrse; pollen grains globose, 3- соро ойы терини ср ес рүш pis- or absent; ovary 2-3- lular; styles bifid; ы gadi seeds eca- 1 runcul at than EU ribe includes two бене but has а different circumscription from "series" Cheilosiform: appears to have been the first to mcn e Neos- adi should be placed in the tribe as ilosa KEY TO THE GENERA OF TRIBE CHEILOSEAE a. m NS rupe inge zv 3 ; Stam ar b. Frui a r Жы, stamens 5 or m seeds; indu- — 94. Cheilosa reine indumentum si 95. А соо сыа T 94. Cheilosa Blume, Bijdr. 613. 1826; Muell. Arg., A Prodr. 15(2): 1122. 1866; Bentham, Gen. 975; Ke 1981. TYPE: Cheilosa montana Blume of two species (perhaps conspecific) in smth yin: (Malaya to the Philippines and Bor- eo). 95. Neoscortechinia Pax, Natürl. Pflanzenfam. Nachtr. 1: 213. 1897; Pax & Hoffm., Pflan- zenr. 147. XIV (Heft 2 52. 1919; ior Shaw, Kew Bull. 16: 368. 3; Whi e, Tree Е. c PL I8: 7 (non b dk m 1885). ТҮРЕ: is techinia kingii 'eoscortechinia. kingii (Hook. "n m.]. H P die Hoff Ai NE rig Phil. J. Sci. Bot. 7: 379. 1912. TYPE: Alcina philippinensis Merr. [= Neoscor- са pike 9 {.) Рах ех $. Moore 1. A tropical Asian genus of four species, distributed from Burma to the Philippines and Solomon Islands. Tribe wn ERISMANTHEAE Webster, E 24: 595. 1975. ТҮРЕ: Erismanthus Wall. e Muell. Arg. onoecious trees or shrubs; indumentum simple; leaves opposite, d simple, entire or obscurel blique, penninerved, stipulate; inflo- dis muticous, uds pollen reme 3-colporate, angu- erturate, sexine not marginat e. lap зле d istillode present; pistillate sepals 5-6, im bricate; petals present or absent; disk absent; ovary 3-locular; ovules anatropous; styles free to connate, entire or bifid: fruit capsular; seeds smooth, ecarun- culate, testa dry This tribe includes three tropical Asian genera that share some common features with tribe P and Moultonianthus have pe KEY TO THE GENERA OF TRIBE ERISMANTHEAE la. Pistillode in staminate flower gus elongated, clavate; уо vlde staminate inflorescence covered with а эу bracts; pistlate flowers apet- petals shorter than calyx; stamens 96. gehe lb. Peake not T elongated; young s inflorescences a Eae with densely att stamens | е с tal nger фан calyx; pis istillate E туеш stip- ules persistent, foliaceous, an ges 2b. Styles entire or nearly so shorter than sepals; реше green apie ous; stipules small, deciduous —_______—— 98. Syndyophyllum 96. Erismanthus Wallich ex Muell. , DC. Prodr. 15(2): 1138. 1866; в. je pl. 3: 325. 1880; Hook. f., . 5: 405. Shaw, уш Bull d. Ser. . 1975; Kew Bull. 36: 294. nae TYPE: fue obli- quus Muell. Arg An Asian genus of two species, the type species —— Volume 81, Number 1 1994 Webster Synopsis of Taxa of Euphorbiaceae occurring from Thailand to Sumatra and Borneo, the other species (E. sinensis Oliver) in Indochina and Hainan. 97. ^ai grip Merrill, Phil. J. Sci. Bot. 11: 0. 1916; Pax & Hoffm., Pflanzenr. 147. XIV 1; Steenis, t. d. Вакво; Ш. 17: Aves Mene "es Shaw, Kew Bull. Add. Ser. . 1975; Kew Bull. 36: 332 1981. TYPE: vid anthus borneensis [= Moulinionhus leembruggianus eh & Koord.) Steenis]. A monotypic genus known only from Sumatra and rneo. 98. RN gear Lauterbach & K. Schu utsche gv 403. 1901; Pax 47. III (Heft 47): 104. illum excelsum Lauterb. & K. Sch. totype } monotypic genus, the single variable tie ا دا‎ tra, Borneo, and New Guin Tribe д9. PCO КАЕ Hurusawa, J. Fac. S Soc. 94: 6. 1987. TYPE: Monoecious trees or s haia indumentum simple; leaves sim imple, entire, pinnately veined, eglandular; stipules caducous. sient os xual, racemose. x te sepals 5, valvate or nearly so; petals 5 valvate, with 2 Teas opposite eis disk S2 nn ula: &natropous; styles free or nearly so. jue bed, jism eds ecarunculat Fruit capsular; seeds s original suggestion of a rela- бы to the Galearieae seems же эле tive and best (y hus by the available d Thy last review of ds à f Dicoeli s MR 0 гсоепа NE E bu tribe ped was кеа he, bes, the being t ы тт If the а меге ES VAL CERE SE es : , this arrange- ment still ир [^ However, i in bi re synopsis, it appears necessary to include the Diccelieae і in the Acalyphoideae adjacent to the Galearieae. It might be күр рер a separate subfamily for the tribes Di » ba at this s еш» pre TREE. family le s are much better understood. 99. Dicoelia Bentham, Hook. Ic. Pl. 13: 70, t. 1289. 1879; Gen. Pl. 8: A: а LF reine wed Hee Bot. Buitenz. III. 1: 392. 1920; ., Pflanzenr. 147. es mh 81) " Airy Shaw, Kew Bull. 27: 3. 197 1981. ТҮРЕ: Dicoelia beccariana Benth. À monotypic a of Malaya, Sumatra, and Bor- neo according to Airy Shaw pian who is not accept D. affinis J. i Sm. as distinct Tribe 20. GALEARIEAE Bentham, Gen. Pl. 3: 241, 287. 1880. err ribe Galeariina Pax, Natürl. Pflanzenfam. ed. 1, 3(5): 81. 1890. TYPE: Galearia Zoll. & els Bennettiaceae R. Brown, РІ. uot дт 250. 1850 à Саа ae & М тыш Pierre, Bull. ie nn. Paris 1255. 1896; Eng er & Gil че бшшш, ед. 7, 223. 1913; Жош: w Bul . 1966; Webster, Bot. Lini agr "94: 6 т es Panda Pierre. Dioecious trees or leaves alternate, simple, entir nately veined, eglandular, indue Inflorescences terminal t thyrse es or axillar y glomerules; bracts ti tent. S sepals free or con- bets slightly imbricate or valvate; petals valvate or somewhat imbricate, with venir pal р aired de- 5 indumentum simple; ire or dentate, pin- pes free; ‘ushers introrse; pistillode arie t som times peltate; pollen grains subglobose, angulap- erturate, a: colpors se sexine finely reticulate. Pis- onnate; petals valvate; ovary ovules 1 per locule, anat- mult pe к ROÊ 8 mainly اک و‎ ropous or orthotropous; styles bifid or Fruit drupaceous; exocarp fleshy or d bony and e or ERE d ecarunculate, endosperm со А tribe Galearieae as treated here corresponds exactly, except for rank, with the family Pandaceae Annals of the d Botanical Garden as circumscribed by Forman (1966). The group has had a rather complicated taxonomic history, i fa n th eta oe operi ae (no denti because of ite бше carpel s). I ostiis asa dide ride group between Ph yllan theae and Crotoneae, included Galearia and Mi- crodesmis buts he Pandaceae м generally been accepted in the 20th century a ат а (e.g.. 1987). I appears that Dicoelieae and Gal defined) form a coherent and apparently mono- phyletic group. The key to the three genera follows that of Forman KEY TO THE GENERA OF TRIBE GALEARIEAE la. Inflorescences axillary, flowers solitary or in glomerules; staminate pet tals imbricate; leaves usually pellucid p edis crodesmis ce: us, thyr- cauliflor T: аан ог ate во; ; leaves not pellucid-pun: 2a. элү: thin. walled; ovules yeay Бера ropous; petals valvate; infloresc ost- lyt terminal e nire 2b. Endocarp thick-walled; ovules onono pous; petals + imbricate; inflore: es cauliflorous 102 Mida 100. ae арво Hook. Ic. Pl. 8: t. 758 8; Mue УрСрг в 15(2): 1041. озн apris се Pl. 3: 287. 1880; Hoffm., Plfanzenr. 147. III (Heft 47): TYPE: Micradesmis puberuka Hook: f ex Planch. асте = of ten species, eight Af- rican and two 101. тунш Zollinger & Moritzi, Syst. Verz. 846 (nom. cons. prop.); Muell. Arg., DC. Head 15(2): 1036. 1866; pi Gen. PI. g^ n ert Hook. f., Fl. Brit. Ind. 5: 377. & Hoffm., Pflanzenr. 147. Ш (iet yei 97. 1911; Gagnepain, Fl. pe Bull linger & Moritzi, G. pedice confused with Bennettia pedicellata R. Br. |. AL Tulasne, Ann. Sci. Nat. III. 15: 259. 1851. E: Cr emostachys filiformis (BL ) Tul. [= Cue . Jav. ien 9. 1852 ong F. YPE: pi javanica r. [= cinis pA (ВІ.) Boerl.]. (1071 11 eatea D А F l'a | bes 9 in X J> six species of tropical southeast Asia (Burma to ds). Solomon Islands 1966. TYPE: 2 oleosa Pierre. Syst. 26: 367. 189 dignes. bou Bot. Jahrb. be Porphyranthus zenkeri Engler [= deba Pi hails typic genus of west tropical Africa. Al- nearly all other Eu- within the Galearieae.‏ ر ا AMPEREAE Muell. Arg., Bot. Zeit. 4. 1864; DC. Prodr. 15(2): 211. 1866. TYPE: ат А. Juss. Ampereae subtribe Monotaxideae Muell. Arg., Linnaea 34: 63. 1865. TYPE: Monotaxis Brongn. Monoecious (rarely dioecious) h shrubs; indumentum simple; leave coid, stipulate; fa flowers in terminal or axillary 819 rul minate calyx 3-5- lobed, lobes хати erbs or s alternate, еп- slightly Cine petals present or absent; ire or dissected; stamens 6-10, biseriate, fila ents free, anthers t or pendulous loc = ^ 5 Prd bifid to multifid; fruit uon: seeds smooth, imam Volume 81, Number 1 1994 Webster Synopsis of Taxa of Euphorbiaceae runculate; endosperm copious; embryo cylindric, cotyledons на: broader than radicle. n the circumscription of Pax о) and Grü- ning (1913), which is followed here, tr mper- era confined to Australia Al- ways, they should be associated in the same tribe. KEY TO THE GENERA OF TRIBE AMPEREAE A sepals imbricate; an- cnm lo iui i 5 жаз grains 3-angle er sexine reticulate; style-branches fimbriate; cym rmina 1 onotaxis lb. Samina petals absent; sepals valvate; ‘anther ocules pendulous from glandular connective; Pel en bte 3-lobed, sexine perforate-foveo- late; style-branches entire; flowers i LER 4. Amperea 103. Monotaxis Brongniart, Voy. Coquille Bot. 223. 182 ll. А rg., DC. Prodr. 15(2) 212. 1866; Bentham, Fl. Austral. 6: 78. 1873; en: РЗ 1880; Grüning, Pflanzenr y Shaw. ы. men Biong ee mics тры Muell. ‚арма 34: б Е: Нїрросг SIL acilis Muell. Mo onotaxis lurida (Muell. P ic) Benth.; designated by Wheeler, 1975]. wl Чел. ыб An Australian genus of about ten species. Brong- iart, in his origina enus of BM 104. Amperea y Mas Euphorb. Tent. 35. 1824; Muell. s DC. PF 6; Bot. (in press). mperea ericoides Accordin g to pyes 1993 , Amperea is a Benus of eight Aus , к, Confined to West tern ack . On the ba pollen Characters, Punt (1962) related Атрегеа further inquiry. Tribe 22. ru ide Na pepe (Muell. i Aca- lypheae subtribe "e Muell. Arg., Linnaea 34: 143. 1865. TYPE: Agros- tistachys Dalz Dioecious (rarely monoecious) trees or shrubs inay simple or a imple, entire or denti eglandular: stipules kaa rescences xillaı ne racemose 0 ок sometimes with sca mbricate bracts; staminate calyx closed in ing: йы pera 5 valvate lobes; petals 3-8; disk nd dis- — or extrastaminal a тан stamen 0, a ше free (at aiti in nthers introrse n pen ж sacs often pendu lous; pollen grains subglo- bose, 3-colpora ‚ pitted disse often tomentose; styles bifid; fruit capsular; seeds h or granulose, ecarunculate, testa some- times fles ibe is here —— to include four This tri genera, two African and two Asia KEY TO THE GENERA OF TRIBE AGROSTISTACHYDEAE Т8; deg present; flowers i in bows or racemes. petals present in N Во flow 3a. Staminate dis ر‎ F stamens 8- 10; stipules free; leaves e glandula 105. Agrostista hys 3b. Staminate disk re rece cepacular; st stan a. 55; stipules connate, lea an nular scar; leaves glandula ular at base _ 106. Pseudagrostistac hys 2b. Monoecious; stamine ate petals not mbricate in bu n peut flowers ШЕТ alous leav Jan „С yttaranthu gla se lb. Petals i rim in T monoe leaves glandular at base . 108. С Жен 105. se TT err Hook. J. Bot. Kew Gard 0; Muell. Arg Рой. 152) сд D Bentham, uM Pl. 26: 210. 1972; Whitmore, Tree Fl. Malaya 9: 52. 1973; Airy Shaw, Kew Bull. Add. Ser. 26. 1975; Kew Bull. 36: 248. 1981. TYPE: pedites ns indica Dalz pepe ne Gagnepain, Notul. Syst. (Paris) 14: 33. 1950. перса laoticus Gagnep. [= Agrostis- vets indica Dalz.]. Asiatic genus of ten species, some difficult to distinguish, distributed from India and Sri Lanka to New Guinea. 106. Pseudagrostistachys Pax & Hoffmann, Pflanzenr. vas VI bcm 57): 96. 1912; Le- brun, Bull. Soc y. Belge 67: 97. 1934; Léonard, Fl. с PN Hd 1962; Rad- cliffe-Smith, Fl. E. Trop. Afr. Euphorb. 1: 166. 1987. TYPE: deg tee af- ricana (Muell. Arg.) Pax & Hoffm > 9 A f. + е Дафе аавал fi Sao Time and Nigeria to 7а aire and Uganda. 107. C толстое, Léonard, Bull. Jard. Bot. Brux. 25: 286 955; Jl. = SIE де 1962. ТҮРЕ п- ard. fined t Africa (Con f ыр чы | 1 on fh 21. ES is go and ) y cl and perhaps not separable. 108. Chondrostylis Boerlage, Ic. Bogor. 1 23. 1897; Koorders, vu т . Bot. tli 9: 45. 1904; Pax & H ., Pflanzenr. 147. VII (Heft 63): 15. TU Airy Shaw, Kew Bull. 14: 358. 1960; Whitmore, Tree Fl. Ma- laya 2: ve е Airy Show: ps Bull. Add. Ser. 4: 6 5; Kew Bull. 36: 276. 1981. TYPE: ranean bancana rl. Kot Ridley, Fl. Malay Penin. 3: 283. p E: Ku uns stlerodendron ts naa up ileg, i (King un t Ho 3 A monotypic genus of southeast Asia (Thailand to Sumatra and Borneo). Tribe 23. CHROZOPHOREAE (Muell. Arg.) Pax & Hoffmann, Pfanne. de = Lag y 5. 1919. Ch lares (Pax & Hoffm.) je & Hoffm., iut ed. 2, 19c: 89. 1931. TYPE: Chrozophora Necker ex A. Juss Trees, shrubs, or he cious; indumentum va s stellate, lepidote, or rt; rarely entirely malpighiaceous (at least in part simple); leaves alternate, su veas or lobed, entire Annals of the Missouri Botanical Garden or E pinnately or e veined, soi stamens 5—many, Мате colporate, colpi sometimes operculate ly marginate, sexine reticulate, often distinctly het- ubica di: pistillode present or absent. Ріѕ Ша flower: sepals 5 ог 6, imbricate ог valvate; ре 5 (в ometi mes reduced. or «решу, ras an or , smooth or ната аилы styles i bed ip ud ог twiodl ы sometimes lacerate. Fruit capsular; seeds dry fleshy, endotesta smooth or roughened to inal ecarunculate. As delimited by Webster (1975) and here, tribe Chroz ophoreae corresponds; inest exactly to subtribe ax & Но (1931); their other subtribe ا‎ in genera belonging to subfamily Crotonoideae. Pullen charac rs in gen- era such as Doryxylon suggest a on i relation- — ship between Chrozophoreae and tribe Epip rineae. — KEY TO THE SUBTRIBES OF TRIBE C HOREAE ains not eines heterobrochate; in- malpighiaceous; leaves not y^55 simple; st ^ es nces ax indumentum ighiaceous or dd (at n in part); не, amens, if more than 6, connate; pistillate dis слей or obsolete... 23b. Ditaxinae - lb. Seni n ач veces Мекеме in — stellate with basal la A. ires За. Shrul rees; pollen grains 3 ate; Ў stamens ys eek fleshy __ 23c. Doryxyli 3b. Herbs; be; pollen ДУ stephanocolporate; stamens conna not fleshy —— . РОЗУ: "934, Chrosophori i Subtribe 23a. SPERANSKIINAE Webs Taxon 24: 596. 1975. TYPE: Speranski И lon. herbs; indumentum simple; lea alternate, mda dentate or lobulate, with gla? T dular teeth, stipula rescenc terminal, = : cate, bisexual, pistillate flowers proximal; 5 5 | and petas 5 disk dissected; stamens 10-1 ffmann — aS tribe TERNS and Al i | i a EI f Volume 81, Number 1 1994 Webster Synopsis of Taxa of Euphorbiaceae filaments free; pollen grains 3-colporate, evenly homes pistillode absent; pistillate gu 5; pet- als 5, small or absent; disk annular; ovary 3-loc ular, hifid } 1 which resembles the Chinese genus in its muricate ovary and laciniate style-branches, but differs in йз axillary racemes and stephanocolporate pollen, a: athe Beak ie hie ос иена capsular; seeds dry, ecarunculate, testa foveolate- sperate. monogeneric subtribe is morphologically Foo to subtribe Ditaxinae, v — Mueller (1866) included Speranskia as tion of Ar- gythamnia. However, the инд diverge from the Ditaxinae in a number of characters, particularly the androecium of free stamens and the terminal infloresc 109, Speranskia veris Etude Gén. Euphorb. 388. 1858; Ben . Univ. Tokyo m. Bot. 6: 310. 1954. TYPE: Speranskia (Bunge) Baillon A genus restricted to China; Ps are three species according to Pax & Hoffmann (1912) or two according to Hurusawa (19 54). Subtribe 23b. — Grisebach, FI. Brit. W. Ind. 43. 1859; Abh. Gesell. Wiss. Goet- Fa 15. 1861. TYPE: pena Vahl ex A. Acalypheae subtribe Caperonieae Muell. Arg., Linnaea 34: 143. 1865. TYPE: Caperonia St. Hil. Monoecious (rarely dioecious) trees, shrubs, or herbs: in. sometimes also st tellate, simple, or glandular; les alternate, entire or serrate, without laminar D es axillary, racemose or glomerulate, usually oye AGAM. sepals mostly 4 or he petals 4 or 15, Абай connate (rarely ‘canes кр һегв introrse; p^ grains age to dide о stephanocolpo icula sexine retic ® ornamented (foveolate, saporita, or aR erarunculate, " In the circumscription of ет 1 а pr Continued here, subtribe Dita subtribe ap via Caperonia, troversial, but pollen characters such as pointed mte delimitation of Mueller (1866) and атат (1979). Key ТО THE GENERA OF SUBTRIBE DITAXINAE la. Pistillode present; "e ed; indumentum simple or glandular (по! eis eous); leaves finely serrate with bacis — lateral M -colpora pollen gr ate 110 - ота . Pis оде, года s i sy 112. Ditaxis 4b. -— уен ү “anthers in | whorl; pollen grains stephanocol- porate 113. Argythamnia 3b. Petals toothed or lacerate; styles once bifid; stellate hairs sometimes present; pollen gr 110. Caperonia St. Hilaire, PI 244. 1826; у Arg., DC. Р 15(2): 151. 1866; Fl. Bras. 11(2): 315. 1874; Ben- tham, Gen. Pl. 3: 304. 1880; Pax & — nold Arbor. Afr. Fl. Pl. 1: 314. 1975; Radcliffe-Smith, Fl. E. Trop. Afr. Euphorb. 1: 163. 1987. TY [lectot уре, — x Britton Bot. 925]. subtribe Caperonieae. On the basis Hans, 1973), Caperonia appears M ت‎ x = 11) iem the м ет Klotzsch, Arch. Naturgesch. 7010: 1841; Baillon, Etude Gén. Euphorb. 297. Annals of the Sachi Botanical Garden x & Hoffmann, Pflanzenr. 147. VI мар ә 49.1912; Чөн, рен 2, 19c: 92. 1931; O’Donell & Lourteig, Lox 8: 60. 1942. TYPE: Philyra е КІ. A monotypic genus of Paraguay and southern Brazil TS vee Vahl ex A. Jussieu, аран oT 4; Pax, Natürl. am. ed. ste y 1890; Pax & , Pflanzenr. 147. T (Heft 57): 51. 1912; Natiirl. Pflan- zenfam. ed. 2, 19c: 93. 1931. TYPE: Ditaxis pons da Vahl ex A. Jus Pflanzenf. Hoffm Ee ayr Trans. Amer. Phil. Soc. H. 5: 174. 1837. ог mercurialina Nutt. [= Ditaxis mer- Serophyton Bentham, Bot. Voy. Sulphur 52. 1844. TYPE ophyton lanceolatum Benth. [= Ditaxis is. offm.]. Stenonia Didrichse ` . Da Foren dre 1857f: 146. 1857 А Endl er, 1947 pel, E: Stenonia Sun idensis Didr. [^ Ditaxis ) Рах]. ee Herter, Rev mer. Bot. 6: 92. 1941. E: Pa r: И (Herter ех Arech.) Herter da " Ditaxis acaulis Herter]. AGE 1 f AOSA Ly io 4 TU L ited States south to о in drier areas from the Un Argentina. Ditaxis я very close ie: Argythamnia, Br - 1 1 1 2 1 14 combined with that genus, as many workers, including Mueller (1866), Va pns (1880), áng Ingram (1979), have proposed. However, evidence from pollen morphology (Punt, 1962; Webster, med Y 3a E $, ph ا‎ + rr 113. Argythamnia P. wn, Civ. Nat. Hist. Jamaica 338. 1756; sit Fl. Ind. Occid. Warnoc 962; "rr Brittonia 16: 271. 426. cd 5 Correll: Fl. Ba ama th. Ant. 3: 250. 1984. TYPE: Aresthemuia can- dicans Swartz. the revisional study by Ingram (1967), 18 species of Argyt M mnia (sensu ни) аге гес- ognized. It is a with over half the species in the West Indies, but a number also occurring in Mexico and Central America 114. Chiropetalum A. Jussieu, Ann. Sci. Nat. I. 25: 21. 1832; Pax & Hoffm., Pflanzenr. 147. VI (Heft 57): 86. 1912; ehe Gentes h et а H. E: Chi- ropetalum сеен за ) fe pe ir Mai Vellozo, Fl. Flum. 95. 1825. Type: Desfon- ricocca Vell. z Chiropetalum tricoccum Cho . PI. Suppl. 43): 89. EE TAE, schiedeanum (Muell Arg.) P x& Hoffm Aonikena н Anal. Ме . Nac. Hist. Nat. Buenos es П. 7: 162. 1902. TYPE: Aonikena ра! йй nica к Chiropetalum dik ia (Speg.) О” M rt]. s of ca. 20 species, almost entirely in a single Mex- zn J ts lobed petals and айе pollen type (Punt, 1962). Арине, 2502 DORYXYLINAE Me Tax- . 1975. ТҮРЕ: Doryxylo noecious shrubs or trees; indumentum stel- n dt 3- БЕШИ stellate-pubescent; fid; seeds with fleshy exotesta, рыны endotest4 This subtribe of four Asian men sta p from other Chrozophoreae by th aeos oody habit and stellate ead ce КЕҮ ТО THE GENERA ОЕ SuBTRIBE DORYXYLINAE la. Petals present; ovary 3-locular; stamens Са. 30-100. 2a. Inflorescences E € unlobed, ba- ; в free _ 115. Dory 2b. Inflorescences ter sina st rth bifid, + free. 3a. Stamens free; staminate disk absent; x 11 Sumbaviops 3b. Stamens connate; staminate ; disk pr ent; pistillate sepals v —€— 5 - here 17. Thyr Petals sess ovary usually 2- xr lar; stamens ed or more; inflorescences rid soles 118 lb. 115. Doryxylon Zollinger, Tijdschr. Ned.- -Ind _ 14: 172. lanolepis i 1857; Linnaea 29: 469. 1859; Be : Volume 81, Number 1 1994 Webster Synopsis of Taxa of Euphorbiaceae lakrishnan, Bull. Bot. Surv. India 9: 56. 1967. TYPE: Doryxylon spinosum Zoll зеи ВаШоп, Etude Сеп. Euphorb. 390. 1858; Muell. bie ped E 727. 1866; Pax bti ffm a VI (Heft 57): 11. 1912 bavia жзнд Baillon [= Ооба йезен Zoll.]. Tue nino genus, Ania to Mele ды ‚ but he treated Sumbaviopsis ira m ot Dor yxylon. He may be correct dE E until the relationships can be further investigated, abet aking Thyr Pereas Earlier doles from Mueller (1866) to Pax & 31) ү san ра into ac- Doryxylon was first published in Linnaea in 1859, and hence did not have priority 116. Ped bt geen J. Smith, Add. Fl. Arb. av ; Air ry Shaw, Кеў Bull. 14: 357. 1960: Фе, Tree Fl. рея 2:79 oer н Airy Shaw, Kew Bull. A 4: 197. 1975. TYPE: Sumbaviopsis POL a ) J. J. Sm. A monotypic ee Asian genus (Assam to Borneo and Palawan). 117. Thyrsant Soc. Bot. [ndi nce e 71: 878. 1925; Deum FI. Indochine 5: 299. 972. TYPE: E suborbicularis Pierre ex Gagne onotypic genus known from Thailand and Canin. Airy Shaw (1972, in clavi) suggested tit may be more DK related to Chrozophora than to Sumbav viopsis 118. Кеннер Reichb. f. & Zoll., Verh. Ма- tuur ا‎ Ind. 1: 22. 1856; Linnaea 28: Ep 1 Hoffm., Pflanz 47. VII (Heh 63): 142. 1914; тор. Fi. nd Fl. 0. TYPE: Melànolepts SUL (Reinw. ) Reichb. f. & Zoll. MR genus of two species extending from nd and Cambodia to Taiwan, Marianas, and Bismarck Archipelago. poer mé — боер нед Muell. ea 34: 14 Prodr. 15(2): 726. 1866; Pax a oe yt anzenr. 147. VI (Heft 57): 1. pone TYPE: Bleck ora tinctoria (L.) A. J Monoecious herbs or subshrubs; indumentum stellate; leaves alternate, entire or lobed, biglan- dular at base, stipulate; inflorescences axillary, ra- cemose, bisexual; staminate sepals and petals 5; ee є A + + + s 1 1 mens 4-15, filaments connate into a ae 1; an- thers introrse; a. grains oblate, stephanocol- colpi s and broad, sexine dicitis Матко, dd illode absent; pistillate sepals petals 5, small or absent; disk lobed or dissected; ova hes 3-locular, stellate or lepidote, sometimes mu- ricate; Mies bifid; fruit capsular; seeds caruncu late, somewhat fleshy, endotesta smooth or t меан d ge In the present circumscription, subtribe Chro- very close to sub- apiid bridges the apparent gap between the wo taxa. 119: Reread ipte Necker ex A. Jussieu 746. 1866; heared is Pi 3: 305. 1880; Pax & Hoffi j rra 147. VI (Heft sa 17. 1912; Praia, K w Bull. Misc. In 49. dew Puja Fl. (он 14: m. — nica 6:5. t, Fl. x i: k 307. 1975; pg Smith, Fl. E. Trop. Afr. Eu- phorb. 1: 160. 128K.. TY TYPE: Chrozophora с (L.) A. Juss An Old World genus of ca. ten species distrib- uted from southern Europe to east Africa, central and southern Asia. Tribe 24. ee с. egenis T on 24: 596. 1975. TYPE sten. oecious trees or shrubs; indumentum simple; nis dep alternate, penninerved or triplinerved, with basal glands; oe ears reduced, or absent; axillary, spicate. Flowers apetalous; аю saab 3-5, valvate; disk in- trasta! ubescent; stamens 4-15, ents М "erue introrse, connective nett or glan Bere en grains oblate, 3-colporat colpi not arginate, sexine coarsely Mei pistilode Annals of th Missouri Воны Сагаеп present or absent. Pistillate sepals 4—6, imbricate vate, deciduous; disk annular or dissecte m esta dry or fles This tribe of three genera is represented by two be the most closely wi qe nao however, they оу differ from more stamens, a glabrous floral disk, and bifid styles. (^ KEY TO THE GENERA ОЕ Т Va: дыы s absent; inflorescence terminal. . Staminate disk massi k- ninerved, stipules py 130. Caryodendron 2b. Staminate disk dissected; px thin- walled, seed coat fleshy; leaves tripli- nerved, exstipulate .......... iscogly premna lb trot: з nt; Lione ae E ; sta e disk nular udi ч аа арн Ten iphened е stipulat ес 120. Сакистон Karsten, Fl. Colombiae 1: 901. t. 1860; Muell. Arg., Fl. Peas 11(2): 706. fe Bentham, Gen. Pl. . 1880; Pax & Hoffm., Pflanzenr. 147. VII us 63): 77. 1989. nocense Karst Саг SS TYPE: КА огі- A neotropical genus of three 5 ir recorded from Panama to Colombia and Brazil. Huft (1989) has recently shown that Centrodiscus M Arg., cited by Pax & Hoffmann and others as a synony m of Caryodendron, was not validly published; hence it is not listed as a synonym. 121. Discoglypremna Prain, Kew Bull. 1911: 3 191 1958; Radcliffe-Smith, Fl. E. Trop. Afr. Б. . 1987. TYPE: Боса caloneura (Pax) Pra A mo otypic African genus, distributed from Guinea sad o Cabind São Tome t a and Uganda. Discoglypremna is lekeli the African vicariant of Alchorneopsis; the two taxa could as easily be regarded as subgenera of a single genus. 122, a Muell. 65; DC. P Arg., Linnaea 34: Missouri Bot. Gard. 54: Alchorneopsis floribunda (Benth. ) Muell. fe A neotropical genus of three described species that are probably conspecific, recorded from West Indies к уйан. Rico), Panama, Brazil, and the Gui a fisci ne Webster, Taxon ubtribe Mer- Hoffm., Pflanzenr 1914. ТҮРЕ: Bernardia ‘Hostê ex M Monoecious or dioecious shrubs or herbs; m piculate; po 3- HM ы inoperculate, conspicuously шаг ginate, sexine par рше. pide e; pistillode es tary or absent. Р! —6, imbricate; td, annular or dissected; ovary 3-locular; styles b anches sometimes lacerate. Fruit capsular; gras smooth, roundish, sometimes cari Б m > е ра sexine; tribe Adelieae do not appear to be deed relat inson (1969) referred Be joe but its pollen is very different from wo апра; KEY TO THE GENERA OF TRIBE BERNARDIEAE la. Se i bifid; stamens 4—50, anther connective or enlarg ам Leaf blades ш ‘stipellate. іна | seeds диннге таныу др ipae tuae Volume 81, Number 1 1994 Webste Syn int ا‎ ы of Euphorbiac 3a. Stamens 30 or fewer, d emar- ginate; seeds carinate 3. Bernardia 3b. Stamens аа Дап зо, ante apic- —— 124. Necepsia 2b. бе ie cated Lea penninerved, short-petioled; КЕШЕ ers jm culate ___ 5. Paranecepsia Ab. e palmatinerved, long-petioled; nthers muticous . Discocleidion 1b. uM unlobed, apia еты А сар over the of the anther con- halt айу. dil a ый st ies Adenophaedra 123. 55 Houstoun ех Miller, Gard. Dict. abr. ed. ч & H a Pflanzenr. ee VII (Heft 63): 21. 1914; Fawc. & Rend., Fl. Jam. 4: 290. 1920; Pax & , Natürl. Pflanzenfam ed. 2, 19c: 105. 1931; Standley & Steyerm Fieldiana Bot. 24(6): 52. 1949; Buchheim, Willdenowia 2: 291. 1960; 3: 217. 1962; Allem, Rev. Brasil. Biol. 39: 529. 1979; Lio- gier, Fl. Espanola 4: 85. 1986. TYPE: Ber- nardia carpinifolia Griseb. td. Pflanzenk. 2: 116. 1820. Bivonia RM Neu Bernardia ax- e En TYPE: Bivonia axillaris ae eng. [= illaris Бр: )M rg.]. tinci Klotzsch, in ek AR 7(1): 188. 1841. : Tra gents و‎ an Kl. [= Bernardia si- es (КЇ) ) Mue А Klotzsch ex LIS Gen. Pl. Suppl. 4(3): 88. - TYPE: Bernardia jacquiniana Muell. Arg. le ecto nd e: here HE К а ех Endlicher, Gen. РІ. Suppl. ps pain Acalypha corensis Jacq. [= dia corensis (Jacq. M uell. A Tria Кыз h ex Endlicher, Gen, Pl E A. x TYPE: Ве = ernardia mexicana (Hoo Muell. Arg. Lectotype, iced felis Alevia Baillon , Et ude b. 508. 1858. TYPE: mmi Im. [= Báriardin spha rupia бешм, ) Muell ate чей 507. 1858. T п [= Bernardia ЕЕ За Bailo assa mente a tioides (Ball ) Muell. Arg A diverse Kee ican genus of ca. 50 species, т into seven sections by Pax Hoffmann Т, 14, 1931); so f these, especially section А amc may din e generic status on further y. 124, Necepsia Prain, Kew Bull. Misc. Inf. 1910: 343. 1910; Hook. Ic. Pl. 30: t. 2987. 1913; Pax & Hoffm., Pflanzenr. 147. ie ва 63): 16. 1914; fisucks t & Léona ull. Jard. Bot. Brux. 56: e 1986; eben Smith, Fl. E. Trop. Afr. rien di 1:5238:51087. TYPE: Necepsia afzelii Pra Neopalissya Pax, in Pax & Hoffm., Pflanzenr. 147. VII (Heft 63): 16. 1914. Palissya Baillon, Ede Cén. ea b. 502. 1858( lli : Neopalissya castaneifolia (Baillon) Pax [= A алаа (ВаШоп) Bouch. & Léon.]. As interpreted by Bouchat & Léonard (1986), Necepsia is an African genus of three species. 125. Paranecepsia Radcliffe-Smith, pus Bull. 30: e 1976; Fl. E. Trop. Afr. Euphorb. TOB TS TYPE: АНЯ epsia alchor- таваа Radcl.-Sm monotypic east African genus (Tanzania and cen e). 126. nd 2 Arg.) Pax & a mann, Pfla . VII (Heft 63): 1914. TYPE: т ulmifolium i get Arg.) Pax & Hoffm A genus of two species found in southeast China and the Ryukyus; reduced to Alchornea by Hu- 4). rusawa (195 127. Adenophaedra (Muell. Arg.) Muell. Arg., Fl. Bras. 11(2): 385. 1874; Bentham, Gen. Pl. 3: 314. 1880; Pax & Hoffmann, Pflan- zenr. 147. VII (Heft 63): 261. 1914; Croizat, Trop. Wood . Miss TYPE: s ыра grandes (К1.) Mue 1. ipi [lectotype, designated by Jablonski, 67]. A neotropical genus of three species extendin from Costa Rica south to Brazil. The r lationship of the genus require further stud ril next to Caryodendron, as did Pax me resemblance, but е т the intuition of Mueller that the closer relationship Adenophaedra is with Berna rdia. Tribe 26. T Hutchinson, Amer. J. Bot. 56: 753. 1969; Webster, Taxon 24: 596. "Er ТҮРЕ: Pycnocoma Bentham um simple, often glan- Dioecious trees; indument posite, or ver- dular, or absent; leaves alternate, op ticillate, rset penninerved, eglandular, some- caducous or absent; or шеш з ы Ганы 15-many, m anthers some- or donc co 3. едри, colpi short ies narrow, ho ulate, ate or nM pistillode sigi ‘Pistillate e sepals 3-7, i a de ; disk glabro (2-)3- hick-walled and subdrupaceous; seeds e late, testa dry or fleshy. As defined by Webster (1975) and here, only us Pycnocoma, of the genera included in the tribe em- bership of the taxa h uded in the Pycno- comeae is janl a ри of doubt. The m ш andi in Soe distribution, so that they pne have to be separated on further study. However, ded. is some resemblance in pollen characters ween Pycnocoma and Blumeodendron, so the two subtribes are kept together provisionally. KEY TO THE SUBTRIBES OF TRIBE PYCNOCOMEAE — a. Monoecious; seed-coat not Ms € sexine gemmate a. Pycnocominae Dioecious; seed-coat fleshy; ig sexine punc- tate-tectate or coarsely reticulate .. 26 b. Вил a 1 = Subtribe 26а. PYCNOCOMINAE eigo subtr. nov. TYPE: Pycnocoma Benth Monoicae; stamina numerosa; antherarum connecti- vum incrassatum; pollinis n 3-colpoidorata, sexino ge emmato; semina non сагпо: Monoecious shrubs mall trees; leaves alter- nate: or peeude: verticilate; men 50 or more; ES sent; styles ongated, connate below, barely lobed at tip; fruit ie -walled, horned or not; seeds with dry testa. This subtribe eins three genera, two African and one Malaga КЕҮ TO THE GENERA OF SUBTRIBE PYCNOCOMINAE - a. Pistillate flower — the inflorescence; pile Pet abse; y 6- тед pubescent; staminate Annals of the Missouri Botanical Garden flowers vri pedicellate; filaments in flexéd in budsizse ed yon . Ovary unappendaged, glabrous; stamina’ M wers subsessile, filaments по Fen “din 9. Droceloncia I BE iini flower not — М ers pistillate disk annular 30. Argomuellera N -a е, airy Bentham, Niger Fl. 508. uell. od phorb. 1: 228. rophylla Bent M TYPE: Руойоб ОШ тас- A tropical African genus of ca. 15 species. 129. Droceloncia Léonard, Bull. Soc. Roy. Bot. Belge 91: 279. 1959. TYPE: Droceloncia rig- idifolia (Baillon) Léonard. A monotypic genus of the Comores and Mad- agascar. 130. Argomuellera Pax, Bo hrb. 19: 90. 1894; Prain, Fl. Trop. Afr. En ie P eke $ Notul Syst. (Paris) 9: 161. 1941; Be Léonard, В с. Ro t. Belge 91: 274 1959; Radcliffe-Smith, Fl. E. Trop. Afr. Eu phorb. 1 87. Pycnocoma sect. We sate Muell. DG. :P. Ee 952 866. петата (Sel Arg. es: Pfla тод VII (Heft 63): 49. 1914. ТҮРЕ: p muellera Ми: эЧ з Рах An Afro-Malagasian g f ca. ten species, % delimited by Léonard (1959). Subtribe 26b. дин INAE Web ster, Taxon 24: 596. TYPE: Blume dendron (Muell. Arg.) E Di us trees; leaves long- geo, entire, with xS orm veinlets; in e огра" niculate; stamens 15—many; anther px pem grains perforate peat of r co: orescences еце @ large, thick- vidi: ск with 71 testa. t This subtribe includes three Asian genera thal differ in aspect from the Pycnocominae by the Jong petiolate leaves and more or less panic culate i rescence permer eem Volume 81, Number 1 1994 Webster 77 Synopsis of Taxa of Euphorbiaceae KEY TO THE GENERA OF SUBTRIBE BLUMEODENDRINAE la. Anther connective than moderately Fn ed, not ніна бнт pistillate disk ten- ; styles elongated. 2a. A her parta 2; fruit pena spines or ridg- a. tillate disk тшшш, эбзе Witholit glandular hai - Blumeodendron А ruin er-cells aan лн ponis disk Meri flowers with stal гч ое, - Podadenia 2b. ibo pé 4; fruit with sins or ridges 133. Ptychopyxis Ww =m lb. Anther connective umbraculiform, with 4 pen- dent locules; inus disk nn pulviniform; styles stigmatiform ........................... 4. Botryophora 131. Blumeodendron (Muell. Arg.) Kurz, J. Asiatic Soc. Bengal 42: 245. Pup J. J. Smith, Meded. Depart. Landbou 458. 1910; Pax & Hoffm -» Pflanzenr. pi" VII (Heft 63): S . Ad . 8: 37. 1980; Ke . 1981. TYPE: Blumeodendron tokbrai jum Kurz [lectotype, designated by Wheeler, 1975]. : Àn Asian genus of six species distributed from the And Tal 1 1D 1 L3 | D to the Bismarck Archipelago. pus орнау ae Enum. Pl. Zey 1861; Muell. ; DG. Prodr: uu c foco ese KA Pl. 3: 318. 1880; Hook. f., Fl. Brit. Ind. 5: 422. 1887; Trimen, Handb. Fl. Ceylon 4: 62. 1898; Pax & Hoffm ^a 147. VII (Heft 3 19.1914. TYPE: adenia sapida Thw otypic genus endemic to Sri Lanka. Al- b is ы, ed by Croizat domi with polt pyxis, it is a very aimi group, " 9 hy - Shaw (19 963). Un uated, it seems best to retain it as a distinct us. 133. jo o tog Miquel, Fl. Ned. Ind. ч, Hooker ie: Pi f., Hook. tcl 1887; Croizat, J. Arnold кы. ed 47. 1942; Air : Bull. 63 1960; 16: 347. 1963; Whitmore, Tree rive Ser. 4: 188. 1975; Add. Ser. bos 980; Kew Bull. 36: 340. 1981. TYPE: Pty- chopyxis costata Miq Clarorivinia Pax & Hoffi ‚ Pflanzenr. 147. VII "e ӨЗА (К. БАБ ) 2 Shaw]. A tropical Asian genus of 10-12 species dis- шы pos Thailand through Indonesia to the Philippines and New Guinea. 134. "Arb узо Hooker f., Fl. Brit. Ind. 5: 476 Hoffm., Na- = © = 5 © B wag d= an = mu = »* 261. 1981. TYPE: Во калыйк kingii Hook. f. [= Botryophora geniculata (Miq.) Beumée ex Airy Shaw]. otypic genus of tropical Asia (Burma to бошан P. Borneo). Tribe 27. EPIPRINEAE (Muell. =н Ниги- Tokyo, Sec Sci. Univ. neae Muell DC. Prodr. me 1024. 1866. TYPE: Ері. prinus Griffith. noecious trees or shrubs; indumentum stel- late; ims alternate, simple, pinnately veined, eglandular or with petiolar r glands, stipulate; inflo- termin iculate 5- 39, osea uve ог шш usually meet hore to subprolate, : 3- арна angulaperturate, cely inate, ne per- 7 argın: lorate. tectate or “rather coarsely reticalate Pistil. lodi) styles 0 Fruit je costed "ars ope semana seeds subglo bose, smooth, dry, ecarunculat This tribe includes nine paleotropical genera that have ne ll ssociated in previous treat- ments, although a number were brought together unde tri phalocroto: ueller 31) in resemblance between Agrostistachydeae, and in some respects with the heae. rismantheae 78 Annals of the s Botanical Garden KEY TO THE SUBTRIBES OF TRIBE EPIPRINEAE Leaves entire or dentate, plui or p la. Pee calyx splitting into distinct segments; veined; staminate flowers in clusters on spica ne not sp xen pistillate sepals often racemose axes, or in e oh Я He k жыган d ac nt Re Epiprininae ^ same nodes as staminate, or lb. Staminate са es каң z 5-lobed, lobes ver- 10, filaments free; pollen e nd globose, will E. rucose; р s; pistilate sepals d tillat сайры i St a EE b. Cephalomappinae реа large colpi, sexine not spinulose; pistillate sepals 4—6, + persistent; capsule smooth. не 27a. EPIPRININAE Muell. Arg., The eight genera of this subtribe occur in Afric Lin aea 34: 144. 1865. TYPE: Epiprinus Grif- Madagascar, and tropical Asia. Although Pax 8 o 919 31) а Acalypheae subtribe Cephalocrotoneae Muell. Arg., Lin- maintained Ep ipr inus in a шыу tribe or sub- naea 34: 143. 1865. TYPE: Cephalocroton Hochst. tribe, its relationship to the other genera of Epi- i m Acalypheae subtribe Mercurialinae series Cladogy: prine a elow seems cl re | a ffm., tns rd VII (Heft semblances in pollen and floral structures (e.g« : TYPE: ас Ipp. ех à 5 ‚ . the pe oes ите е асан Thin, Tap Chi Sinh St@munate flowers with filaments inflexed in 10(2) 32. 1988. TYPE: Cleidocarpon Airy ud). КЕҮ TO THE GENERA OF SUBTRIBE EPIPRINAE la. Staminate flow cemes or spikes; stipules (if present) glandular 2a. lae agli inse PERRA dag ay ate with decidu uous biglandular bracts; styles наш into а 1, distally or multifid; stamens 5-15; leaves with petiolar glands .. 135. Epiprinus 2b. Plate calyx n c т accrescent nor AUS ml styles + free, distally multifid; stamens yi e ments се in bud. . Fruits S ра leases „glandular. heads 136. Symphyllia istillate e sep P dec staminate flowers in heads — ..... 137. Adenochlaena b. Fruit inde эн рт biglandular 138. Cleidiocarpo" 3b. Fila Ts not inflexed in bud 139. Koilodepas lb. Saree? ч ers in pedunculate capitula; сони пої Lge m ences axillary, inconspicuous; stamens 3-5 l t, unlobed; pec white- ose beneath 0. Cladogynos 5b. [rusas ds terminal; leaves b 6a. Pis tillat ate are entre, connate styles free; leaves coriaceous pa ae WA psis 6b. ; styles connate; leaves thinner ephalocroton 135. Epiprinus Griffith, Notul. Pl. Asiat. 4: 487. 763. 1866; Gagnepain, Fl. Indochine 5: 477. 1854; Muell. Arg., DC. Prodr. 15(2): 1024. 1926. TYPE: Symphyllia siletiana Baillon. 1866; Bentham, Gen. Pl. 3: 325. 1880; Hook. f., Fl. Brit. Ind. 5: 463. 1887; Pax & Hoff- A genus of three species distributed from India ann Gagnepain, Bull. Soc. Bot. France 72: 465. Symphyllia with Epiprinus on the basis that Ept 1925; Fl. Indochine 5: 474. 1926; Croizat, aig balansae (Pax а пиво, ) Савра No J. Arnold Arbor. 23: 52. 1942; Airy Shaw, a transitional link Kew Bull. 16: 356. 1963; Kew Bull. 26: 259. Shaw (1963, 1972) and Thin (1988) нен him 1972; Whitmore, Tree Fl. Malaya 2: 95. in this, but neither discussed the morphology # 73; э Tap Chi Sinh Hoc 10(2): 30. the pistillate flower or demonstrate t the "m z 1988. : Epiprinus malayanus Griff. volucre” described by Mueller is without systemalie A rather variable genus of ca. six species in tropica ia, fro (1866), followed by Pax & Hoffmann (1919), placed 5€*"* preferable to maintain Symphy; Epiprinus by itself in subtribe E d опе "M basis that the pistillate flower is involuer: 137. Adenochlaena Baillon, Erie Gén. E | pue уер Baillon, Etude Gén. Euphorb. phorb. 472. 1858; Pax, Pflanz пг. 147. uid. 858; Muell. Arg., DC. Prodr. 15(2): (Heft 44): 12. 1910; Pax & Hot i | Volume 81, Number 1 1994 Webste Syn sen of Taxa of Euphorbiaceae Pflanzenfam. ed. 2, 19c: Adenochlaena leucocephala Baillon. бенен, Pelle, Etude Gén. Euphorb. 469. 1858. ostylis ea Baillon [= Adenoch- (Ba Thw He ie оп) Niedenzua Pax, Bot. ieee: 19: 10 ; E: Nie- denzua tarditas Pax [7 AURORE ета mies ala Baillon]. of two disjunct species (Madagascar/ aliae si Lanka); treated by Mueller (1866) as a section of Cephalocroton. 138. errem Air d Kew Bull. 31 Bg Sinh Hoc 10(2): d 1988. TYPE: iPad laurinum Airy Shaw aa Tsiang, Acta Bot. Eee 15: 132. : Sinopimelodendro angsiense Tsang is ure kien (Lew) Airy enus of two closely related species of Burma and China; Thin (1988) placed it in a separ subtribe, Cleidiocarpinae ate 139. а» мла joa Versl. Med. Afd. tuurk. Kon. Akad. Wetensch. 4: 139. 1856: Flora 40: 531. 1857 (as C us Muell. Arg., DC. Prodr. 15(2): 759. 1866 k. I 137. 1975; Airy Shaw, Kew Bull. 36: 310. 1981. TYPE: Koilode epas bantamense Hassk "шы Blume, Ann. Mus. Lugd.-Batav. 2: id . Soc. Bot. France 72: 467. ephrostylus poilanei Gagnep. [— B longifolium Hook. f.]. À genus of t Asia as TYPI Vries we epa in, Ne en species of India and southeastern ar as Hainan and Borneo 140 ogynos es ex балаа Lin- naea 15: 349. 1841; Muel Arg., DC. Prodr. 15(2): 895. 1866; Backer Gen. PI. 3: 323. 1880; J. А Smith Bakhuizen, Fl. Java 1: . 1963; Airy Shaw, Kew Bull. 26: 232. 124. 1931. TYPE: 1972. TYPE: Cladogynos orientalis Zipp. ex Span. a AES, Берас & кш. Acta Soc. Regia Sci. Indo-Neerl. - TYPE: Adenogynum discolor Reich & Zoli. E С ladogynos orientalis ipp- ex Span.]. otypic genus of tropical pera ^w Sees to the Philippines, Java, and Tin 141. Cep halocrotonopsis Pax, Pflanzenr. 147. . 1973. TYPE: Cephaloc ice opsis socotrana (Balf. f.) Pax o monona е of Socotra; reduced b y Rad- to a section of Cephalocroton. 142. Cephalocroton Hochstetter, Flora 24 370. 1841; Muell. Arg., DC. Prodr Trop. Afr. Eu Cephalocroton mn Hochs An Old World genus of five species recorded from east ака Madagascar and the Comores, and Sri L Subtribe 27b. CEPHALOMAPPINAE Web- ie Taxon 24: 597. 1975. TYPE: Cephalo- appa Baillon Leaves entire, pinnately veined; staminate flow- stillate flowers of inflorescence; stamens Afta aments aue: pollen dein globose, coarsely reticulat pistillate sea 4- 6, аса сарѕше ‘verrucose. This subtribe includes only the type genus. 143. Cephalomappa Baillon, Adansonia I. 130. 1874; Bentham, Gen. Pl. 3: 323 Kew B 975; Kew Bull. 36: 274. 1981. TYPE: Cephalomappa beccariana Baillon Tried Chun Acta r Am uy 5:14 1956. E: Muricococcum sinense & How r- a otir sinensis (Chun & gom Kos- m.]. enus of five species in southeast Asia (Malaya to Sumatra and Borneo, southern China). Annals of the Missouri Botanical Garden ае" I АРКЫБАВ ояи Тахоп ени 597. гіеѕ оина ormes Pax & Н , Pflanzenr. 147. VII (Heft 63): 59. ums TYPE: Adelia I oecious (rarely monoecious) trees or shrubs; ‚ + connate at base; anthers versatile; pollen grains 3-(4-)colporate, col- pi rculate, exine tectate-perforate with polyg- hdi: often reflexed; disk annular, glabrous or pube t; seeds roundish, smooth, ecarunculate ү carunculate) An American tribe of five genera. KEY TO THE GENERA OF TRIBE ADELIEAE la. Indumentum simple; stipules deciduous; m grains with finely perforate tectum; styles lac erate 2a. Ovary T е staminate disk present; eeds penc te. 3a. "Fille peek staminate disk an- or segmented; sae | in ү 4. Adelia Pace 3b. Pistillode rum staminate M fu in terstaminal pro ; flowers in €—— сы roionogynopsis 2b. ре 2-locular; йиш. disk d mens more than 20; seeds carun juu E ESE 146. epee Ê lb. Wig ias stellate or stellate-lepidote; stip- m “т pollen grains with coarsely perfo- ra um. de halisi disk entire; styles bifid or sub- entire; filaments free; рда. 3-colporate; leaves triplinerved, indumen’ stellate Lasiocroton 4b. Pistillate disk lobed; н шаш fila- ; lpo penninerved, ind t tells ecc ctu у нр aD 148. T rion 2: 1298. 175 9 (nom. E) e P3 312. nzenr. Ga 272. 1968; Sneep & De Roon, Fl. N 3: 254. 1984. TYPE: Adelia rouco, ii». cons.). Ricinella Muell. Arg., Linnaea 34: 153. 1865. TYPE: Ricinella pedunculosa (A. Rich.) Muell. Arg. [= Adelia ricinella L.]. А neotropical genus of ca. 10—12 species, Mex- ico to Paraguay and Brazil but best represented in the West Indies. 145. eee аи. Pax, Bot. Jahrb. 2 ste cap неч H ee P ey 14. 1914; Keay, Fl. W. Trop. Afr. ‚ 2, 1: 404. 1958; bui Smith, Fl. Trop. Afr. Euphorb. 1: 213 7. TYPE: Cro- tonogynopsis usambarica я monotypic African genus (Zaire to Uganda i Tanzania). 146. дне” ape immi Rzedowski, B 55-1970 da, Bol 7 (non жакры Pansies 1882). TYPE: ‘Enriquebel trania crenatifolia (Miranda) Rzedow ski [ Beltrania asia. Miranda ]. otypic genus of the Am Yucatan penins very ta to Adelia and ا ا‎ 147. T perii Grisebach, Fl. Brit. Ww. Б: 46. ; Abh. Ko 9: ^ Tur Bentham, Gen. P. Pax, Pflanzenr 4 7. VII ae i 60. 1914; Fawc. & Rend., Fl. Jam. 4: 293. 1920 0; Alain, Fl. Cuba 3: 87. 1953. et Lasiocroton mae rophyllus pie ) Gris A West Indian genus of five species (Bahamas, Cuba, Jamaica, and Hispaniola). Kus 1 h, Wiss. Gottingen 9: 20. DC. Prodr. 1 89. 195% Borhidi, Acta Bot. Acad. gar. 1975. TYPE: e d ae A genus of 20 species of Cuba and Hispaniob- | Tribe 29. ALCHORNEAE (Hurusawa) Hutt inson, Amer. J. e ate 752. 1969. a pheae subtribe A inae Hurus Sci. Univ. Tokyo УП. 6: 302. 1954 Ж | Alchornea Sw. | H imes stipe! Volume 81, Number 1 Webster 81 1994 Synopsis of Taxa of Euphorbiaceae nflorescences terminal or ax- 3b. Styles Vanno or dilated; indumentum illary, spicate or paniculate (compounded spikes); simple; y 3-locular bracts glandular or eglandular; flowers apetalous fa; Stamens uly ее; stigmas Staminate calyx splitting in valvate seg- ا‎ pilae тоз ments; disk aminal or absent; stamens (2—) Ab. Stamens 2 y 4, , basally conn inii I oe 4—60, free; anthers introrse, muticous or a 5 . Sta nitens i styles elongated, di- pollen grains 3-colporate, colpi operculate, sexine rugulose to vermiculate; pistillode rud tary or 5b. — dune 3. M р aristhmium bud дага нери als дг 8; itobricete; gd. per- form; seeds аму, crunch en-us elie a NR Меш 4. Bocquillonia tary or absent; pvary 2- or 3- (rarely 4- рет Du entire to bifid or multifid. TM capsular; es Wes owe Etude Gén. ctim 452. seeds smooth or issue testa not fleshy E: Orfilea coriacea Bail Ln ielimited earlier а 1975) and here, tri сте the circumscription of ть pow since Conceveiba and its allies— referred to a гї o Mallotus and related genera, but are iinet i in the operculate colpi. KEY TO THE SuBTRIBES OF TRIBE ALCHORNEAE la. T entire or if distinctly bifid, then indu- um simple; stamens 4—9; staminate inflo- i rescence axilla о. 29a. ALC eon stellate; stamens 60; staminate inflorescences Ae or eL 29b. CONCEVEIBINAE Ste 29 29a. ee зе 302. Indumentum simple or stellate; staminate inflo- " ces axillary; stamens 2—9; pistillate sepals hs andular; me mostly entire, often m 9r dilated; ovary 2. or 3. (rarely 4-)locul : ғ ubtribe of six genera wi cies Bor гайн. їп bes the Neotropics and Pa- Topics. Most o genera are only weakly e parated from a and the number ma uced on further st tudy. КЕ Y TO THE GENERA OF SUBTRIBE ALCHORNEINAE la, d bifid; indumentum s Dioe ; ovary edens: ык Ее seeds ecarun neulat HE ee end аз ds caru ger SU Np M" 0. Bo sera lb. Sole кыа ог bif reis at tip. = а. Styles elo n: and "qna indumentum le or stellate; ovary 2- or 3-locular __ is 151. Alchornea Diderotia Baillon, Adansonia I. 1: 274. 1861. Laurem- ergi i 1 rgi- 1767). TvPE: Diderotia ына (ВаШоп) ВаШоп [= Orfilea multispicata (ВаШоп) Webster, с А Ма ov.]. lagasian genus of four species, closely re ong to this genus: Orfilea neraudiana ib Webster (comb. nov.; based on Claoxylon neraudianum Baillon, Adansonia I. 1: 280. 1861). 150. Bossera Leandri, Adansonia II. TYPE: 1962. This E: Bossera cristatocarpa е monotypic genus described from Mada- 1 gascar is very close to Alchornea and is weakly ae $c Ж ИА J : £ 4 distinguished Py the distinctively cristate ovary. 1515 рыта Swartz, Prodr. 6, 98. 1788; Oc Fl. Ind. Occ. 2 53. 1800; Muell. Arg., DC. Prodr. 15(2): 899. 1866; ЕІ. Bras. 11(2) 374. 1874; Bentham, Gen. 1 80; Pax, Pflanzenr. 147. "үп (Heft 63): 220. 1914; Webster, Ann. Missouri Bot. Gard. 54: 279. 1968; 75: 1100. 1989; Whitmore, Tree ЕІ. nh Hoc Арии 26. 1984. ТҮРЕ: Alchornea lati- eae Fl. Cochinch. 574. 1790. TYPE: Clad- 1. 1 odes rust Lour. [= Alchornea rugosa (Lour.) Muell. Arg.]. Hermesia Humb. & Bonpl. ex Willd., Sp. Pl. 4: 809. 1805. TYPE: Hermesia castanifolia ja Humb. & Bonpl. Willd. [= Alchornea castanifolia (Humb. & . Ju ) ТҮ aes a rt (Schum.) Muell. Annals of the Missouri Botanical Garden Зеби Bentham, J. Bot. Kew Gard. Misc. 6: 2. 1854. E: Stipellaria trewioides Benth. [= Alchornea ипшде, е Muell. Arg.; lectotype, desig- 84]. nO ber: ex 'Bailon, pute "беп Euphorb. 4 48. 858. ТҮРЕ: о). ех ВаШоп [= Alc чаары anata (Bailen) Pax & Hoffm.]. Bleekeria Miquel, Fl 1(2): 407. 1859 (non m 2:1855y.3 Meses zollingeri rea ) q. [= а mh (Benth.) Muell. A A variable genus of ca. 50 species distributed orld and О the tropics of both the New W ld World. Pax (1914) drew the generic ани 5 more narrowly than Mueller (1866) by recognizing such segregates as Aparisthmium and Coelebo ryne. However, the Paxian genus still includes ; 8 three sections, and the relationships betwe and the segregate taxa badly need clarification. Ee Beh. ics J. Smith, Proc. Linn. Soc. 839; Baillon, Etude Gén. Eu- ee gua ASA Pax, Pflanzenr. 147. VII (Heft 63): hE vigi TYPE: Coelebogyne il- ipee J.S otypic genus from tropical Australia, Men we a distinguished from Alchornea and p better treated as a section of that genus 153. apir rt Gen. Pl. 1112. 1840 r. 147. . Jussieu, Euphorb. Tent. 42. 1824. TYPE: Concevelinis orda tum A. Juss. [= Aparisthmium ва (А. Juss.) Baillon). the concept of Pax (1914), e by clearly seems to fit the plant that Baillon and later writers have accepted as Aparisthmium. If End- licher’s tion is regarded as a misprint, the t abundance of the plant in neotropical vegetation. 154. асе ВаШоп, Адапѕопіа І. 2: 225. 861; М Р 1; Muell. Arg., DC. r. 15(2): 894. 1866; Bertani: Gen. Pl 13. 1880; Pax, Pflanzenr. 14 II (Heft 63): 260. 1914; Airy Shaw, Kew Bull. 29: 321. 1974; Me- Sh aries & Tirel, Fl. Nouv.-Calédonie 14): . TYPE: Bocquillonia sessiliflora чеде (кедей by McPherson & Tirel, 1987]. Ramelia Baillon, Adansonia I. 11: 132. 1874. Ra melia codonost tylis Baillon [= Bocquillonia a co- ) 1. ) Airy Shaw As treated * McPherson & Tirel M E CA is a genus of Caled зарате 29b. CONCEVEIBINAE Webster, n 24: 597. 1975. TYPE: Conceveiba Aubl. or subentire, not t lea st in part); stillate mr es long-petiolate, entire нт s indumentum stellate y» oral disk absen subtribe of three genera and species was thought to be entirely American until the recent discov f Conceveiba in west Africa. Pax & Hoffmann (191 ы) included Conceveiba and Ga i es Tre mes (Mallo- arretia in their ser teae), an entirely ا‎ grou (^. JAF KEY TO THE GENERA OF SUBT la. Inflorescences terminal. eg E: illate ра aii. often glandular cous; y 3- ics gi branches blun ae c: gl d N c iere sp ponte TS ie pee 2. locular; style-branches acu “ x кн 56. Gavarretia 1b. Inflorescences axillary; anthers mico ME 7. Polyandra = pe еа one Hist. We Fr. ; Muell. "poet г. 1502 i bos Jose Fas РІ. 22 pii Pax, Pflanzen. 147. VII (Heft бз, a 12 0 New York . Gard. 1 P135 ri Bot. call 77: 856. 1990. TYPE: ESAE ; guianensi Aubl. — — (Muell. Arg.) Pax & Hoffm ., Pfla айы I (Heft 63): 217. 1914. n: Conc fasi m martianum рет Pax & Hoffm. [= -UR ba martia 147. Veconcibea spes red ) nme & Hoffm., Pflanze re II (Heft 63): 218. 1914. TYPE: pem \ $ Volume 81, Number 1 1994 folia (Benth.) Pax & Hoffm. [= Conceveiba latifolia Benth.]. A eg die seven or eight Ae tor: species Ri n Peru and Brazil, mu one rae discovered erate species from Gabon (Thomas, 1990). ica to Amazoni YT үт не “api Baillon, Adansonia I. 1: 186 SPS SIG: ew York 130. 1967. TYPE: Clue RE lon. A monotypic South American genus; the single отап Venezuel е ап о be the best iit disposition 157. اا‎ Leal, Arch. Janeiro 11 19 teosa Le a Jard. Bot. Rio de 51. TYPE: ы brac- A poorly lei seria Des known ШШ, n the ype S ers жы fruits are still unknown. KEY TO THE SUBTRIBES OF TRIBE ACALYPHEAE la. р к canünculate; inflorescences terminal; indumentum stellate оа sd е bifid. corn Webster Synopsis of Taxa of Euphorbiaceae Is 905 {АСМАРИРАВ rau Anal. . 1829. TYPE: Аса a L. Monoecious or dioecious trees, shrub, or herbs; P glan = сга inflorescences ев or axil- хөй, һгас imeem اا‎ dular: ud, valvately ming into 2- 5 з гел ents; disk real (intrastam 5 or absent; stam E pha maly aito: Pistillate sepals mostly 3— or absent; ovary 2— 4-locular; pus "ion ог basally connate, unlobed to lacerate. Fruit capsular or drupaceous; seeds carunculate or ae testa dry or fle ie de ig tribe of Acalyphoideae, here а subtribes with 30 genera and over 1000 ies s; filaments fre 30a. Ricininae 30b. Adrianinae 2 a оесіоц lb. ipe ccarunciate Stamens g © © 5 5 @ E 8 E zi © P Ф s Ф Ф ЖЕ {те е, or if carun еше then inflorescences cc axis or гала unlobed. stipu ules d pendulous; pollen colporate; seeds usually ecarunculate. mentum usually imple (except Lobaniliinae); leaves alternate, eglandular or with em- ded laminar glands, not gra alee -glandu 6a. р rbs, often with opposite казы indumentum simple; Чөп undiv =: St 3 ens free; ovary 2-loc sepals free; stam tam. minate thi anthers not No oo lar. ided. r; pollen gu 3-colporate: broader than не 9 e Mecania Tb. о: xd n. als aments connate; ovary 3- locular; pollen atii ina a skakis: cotyledons scarcely broader than radicle сс SOU. е 6b. Shrubs or heen orm herbs with bifid styles). 8a. р "Sab a |, ni with кеш. Жыр 9a. 1 9] | " 1 30e. c ilie 9b. "ir "nie anthers 3- fles usually © >" styles unlobed. 1 4-locular; pollen grains spinulose-rugulose; seed-coat 30f. Mereenie . Anther-sacs ê to the connective, erect or pendulous; interstaminal disk present — -y ухсан а. Indumentum simple LL 10b. Indumentum stellate ШИВА oe a at UE, 5b. Ind t fi te, ofi n with embedded — vira an S lly di d > D esent or absent; ollen grains + Cees -rugulose ne 0 ت‎ 4b. Ant ermiform and ا‎ vn disk sem t; pollen porate; styles Tere ghe + сагип 3b. „ады та simple or le; epi idot ate; pistillate bracts us aie [ie and foliaceo а Ri connate into fascicles; shrubs or trees with pi энш е ager le leaves, жеч ите 30). —— rere ИҢ Annals of the sided: Botanical Garden эс. we аот Grisebach, Fl. Brit. . Konigl. Ges. ed. Taxon 24: 597. 1975. TYPE: Ricinu SI ecious trees or shrubs (herbs in temperate ; indumentum abse n grains 3 colporate, cop narrow and шору late; peti, ; Sty ; capsule echinate; seeds carunculaté. As here defined, subtribe Ricininae is restricted to the single genus Ricinus. Pax & Hoffmann (1919) E Pone Homonoia and Lasiococca пеше, ашу because yi he rather However, Ricinus differs from those genera (here subtribe Lasiococcinae) in е distinctly its terminal carunculate seeds, pal- вач oe with ш peile, ш ae st кешл / that the character of | ei ae omo plasious, and the ninae do not appear а related to the La On the other han Adriana is much more similar to Ricinus e appears to be the most closely related genus. m pie me Pl. 1007. 1753; Gen. PI. , 437. АВ: ене 1:322 TYPE: Ricinus commun A monotypic genus native to northeast а а » катон, now wide ly c worldwide. — pore ADRIANINAE m Gen. 597. pisa ee Adrianeae (Benth.) im Pflan. zenr. 147. Б (Heft 44): 1. 1910. TYPE: Ad- riana Gau Dioecious; indumentum stellate or absent; lea alternate or opposite, unlobed or palmately lobed, stipules glandular; inflorescences spicate, terminal glandular; stamens 5 dnate to 3-locular, smooth; styles bifid; fruit Sie ا‎ carunculate. In contrast to the Wu of Bentham (1880) and Pax (1910), su e Adrianinae is here ге stricted to the at genus Aduana Fag (19 © 1 Manihot rather sla Ind in his and Pachystroma. In his later work (Pax & Hof mann, 1931), M аши ha trike Adi and included adjacent to Conceveiba and Gavarretia, which is a better and not implausible suggestion of affinity. 159. Ann. Sci. Nat. Paris 5: 223. 1825; Muell. زا‎ Prodr. 1 889. 1866; Bentham, Ps Aus 1873; Gen. PI. 3: 306. 0; Pax 147. II (Heft 44): 17. ib be Shai Kew Bull. 35: 589. 1980. TYPE: Adriana tomen- tosa Gaud. pa ype LEE y geographical disjunction. How ever, Adriana such as Cephalocroton and Cephalom appa d in the = may be necessary to remove the rianinae and Ricininae from the accumulate further knowledge venia e TDN Pax, Ne curialiiformes Pax & Hoffm., Pss Wi. drianeae by Pax (Ie | Acalypheae as "° 890 Aer es Mer i уп ыны 6З): 270. 1914. ТҮРЕ: Mercurial | onoecious or dioecious herbs; indum styles it ca meret smooth dus scis сые cotyledons much broader icle. entum - Volume 81, Number 1 1994 This subtribe includes three vem one Eur- fric asian and the other two South Afric KEY TO THE GENERA OF SUBTRIBE MERCURIALINAE la. Dioecious; stamens AS 20; pistillate flower with disk- ve eni seeds carunculate DAN Poca FO PUTER 7 0. Me rcurialis lb. N 2-7; pistill тке or very small; seeds ecarunc 2a. Capsule smooth; pistillate crab P ; leaves entire or denticulate |... . Seidelia 2b. Capsule setose-muricate; e TT CRT nearly obsolete; leaves crenulat 160. Mercurialis L., Sp. Ро T1035: T 153: Gen. Pl. ed. 5 VII (Heft 63): 271. ; Zimmermann e al., in Hegi, Ш. ЕІ. “peti 5(1): 126. 1925; Siig 1. Arnold Arbor. 48: 3 erennis urasian genus of eight species, seven in Europe and з ае Africa, and опе іп east temperate A b тад uen Etude n Anse 465. m, Gen. PI. 3 1880; Prain, ,6d-3, 316. 1975. T TYPE: Seidelia Mer E Baillon (nom. Шер.) [= Seidelia triandra (E. Mey.) Pax; pin ignated by Pfeiffer, Nomencl. Bot. 2: 1128. A iun African genus of two species. 162. Leidesia Muell. Arg., DC. Prodr. 15(2): 792. 1866; Bentham, Hook. Ic. bn. Panzer 147. VII (Heft 63): 284. ^: Prain, Ann. Bot. 27: 399. 1913; T Capensis 5(2: 1920; рун tn S.A 2 * ed. 3, 316. 1975. ge cs (Thunb) Mon Arg. Toe. cit. 793; type, chosen here]. African genus of two or three species, 277 ав to Seidelia; the two genera ш) je be combined without obscurin g phy- ка Рах & Hoffmann (1914) used A South very closely fi Webster Synopsis of Taxa of Euphorbiaceae the name Leidesia procumbens (L.) Prain for the fwo ponies of Mueller андин this vecti be Arg., which unfortunately i is based on a confusion choose Mueller’s thie species as lectotype. Subtribe 30d. DYSOPSIDINAE apap 1 Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 6: : 1954. ТҮРЕ: Dysopsis Baillon. Monoecious herbs; leaves alternate, simple, cre- a nate, eglandular; stipules caducous; flowers axil lary, mostly solitary; staminate calyx gamophyl us; stamens 3-6, interio ; anthers e to connective; pollen grains ovary 3-locular, ate; fruit capsular; apiculate, sacs adna globose, eap iila DUE 3; о pubescent; styles seeds with obsolete UM А monogeneric tribe of South America; closely ide 15(2) 63): 286. 1914. Molina Gay, Hist. Chile ier 5: 345. 1851 (non Molina Cav., 1790). T olina chilensis — Dysopsis dibus Baillon, nom. illeg. T DUD glechomoides (A. Rich.) Muell. Arg.]. otypic genus of the South American An- des (аша to Chile), Juan ee and Costa Ric — ная CLEIDIINAE hag ooi Taxon 1975. ТҮРЕ: Cleidion Blu Monoecious or dioecious trees or shrubs; indu mentum simple; leaves alternate, simple, pin reel » veined, with laminar glands, stipulate; inflores cences axillary, spicate or racts ssh dular; flowers apetalous, without dink; staminate sepals 3 or 4; stamens 25-80, free or basally 3-colporate, colpi not operculate; sexine ru pee te, spinules vestigial; pistillod istillate vm 3- i imbricate; ovary (2- et locular aia bifid I with dry or r fleshy testa. - here construed, subtribe Cleidiinae includes three genera, of which two are paleotropical and e absent Annals of the Missouri Botanical Garden the third pantropical. The characteristic globose pollen grains with vermiculate sexi rnamenta- tion indicate an y between Weitria and Clei dion not clearly expressed in previous classifica- tion & m is stellate and the pollen grains oper- КЕҮ ТО THE GENERA OF SUBTRIBE CLEIDIINAE la. Staminate flowers пе usually 2 ог more per bract; stamens 25-8 a. An ther: s muticous, pied stamens 25- 30; Бейне pelier lessthan"] сш лоп 9 Wetria 2b. Anthers кагыш connective еп ыз stamens mostly 30-80 or more; pistillat резе Ww over 1 cm longs сс; 165. Cleidion Ib. St per bract; stamens 15-25, кау apiculate, ет ї larged Hif abba 164. ерта ВаШоп, Etude Сеп. Кр, 409. КД Smith; ГУ e Dept. Landb. 10 Fl. Java 1: 485. 1963; Airy Shaw, Kew Bull 26: 350. 1972; Whitmore, Tree Fl Malaya 2:136; d Airy Shaw, Kew m 58. o : Wetria trewioides g. [= Wein insignis (Steud. Е A Shan] А Шш Burma to ae and New Gunea; Airy o Alchornea, but it di ffers ir in its simple Шш кек bifid styles, and larger stamen number. o i eu oss Bijdr. Fl. Ned. Ind. 612. Muell. Arg. 6; ., DC. Prodr. 15(2): 983 eni ras Gen. Pl. 3: 320. 1880; Pax & ‚ Pflanzenr. 147. VII (Heft 63): 288 javanicum Blum Redia Casaretto, Nov. Stirp. Bras. n 1843. TYPE: Re- dia tricocca Casar. [= Cleidio. ) Baillon]. ss. Turcz., Bull. So 58. 1843. TYPE: Pench a timete Turcz. a ей ion t Lasiostyles Presl, Abh. Kónigl. Bohm. Ges. Wiss. V. 579. 1845. TYPE: Meets salicifolia Presl К leidion javani um Tetraglossa Pelion Madea ET Lit. Sci, Il; 1861. TYPE: ETE E See "di dion tied Blume]. A well-marked genus of ca. 25 species, of which 5 are neotropical, 1 west African, 12 New Cale donian, and the remainder tropical Asian. um fum cipue Airy Shaw, Kew Bull. 26: 2; Hook. Ic. Pl. 38: t. 3717. 1974. ТҮРЕ: Sampanaca amentiflora (Airy Shaw) m Shaw typic pene restricted to Thailand and Ca ut It appears closely related to Wetria and may prove congeneric. Subtribe 30f. MACA (Hutchin Web E Bot. 56: 755. 1969. TYPE: Macaranga Du Рей Dioecious trees or shrubs; indumentum simple | leaves alternate, unlobed or arian be d, | ped to pa Imately ve чырыш дн а xillary. racemose or pani Re RER $ glandular; E king disk; staminate calyx als 3-5, free or c EC ocular; styles unlobed; fruit capsular; | ecarunculate, testa fleshy. As defined by Webster (1975) and bee subirle | seven other genera fro cta ce ae and Crotonoideae only the &* s Масагапда is included h 167. e Du Petit Thouars, беп. N 2 | Madag. 26. 1806; Muell. Arg., DC. Pr . Ser. 123. 1980; A ШЕ NS 2: 500. 1981; s Тий i Volume 81, Number 1 1994 25: 184. 1976; McPherson & Tirel, Fl. Nouv.- Radcliffe-Smith, Fl. E. 39. 1987. TYPE: jected the proposed selection of M. roxburgii Wight by Wheeler, 1975]. pee Tu pai Tent. 44. 1824. TYPE: Map- ) s. [7 Macaranga glabra (A. Juss. Es Pachystemon Blume, Bijdr. Fl. Ned. Ind. 626. 1826 TYPE: Pac +hystemon trilobum (Reinw. ex Bl.) В Macaranga triloba (Reinw Bl.) Muell. Arg.] Mecostylis Kurz ex Teysmann & nendijk, Natuurk. Tijdschr. edm ik 44 zs hoides 4. TYPE: ешн lyp Kurz ex Tey: Binn. [= Maca anga involucrata ш (Roxb) Ballon pi Seemann, J. . 1870. TYPE: Phocea andersonii i Seem. ар М, ified yis & Hoffm., 1931, or McPherson & Tirel, А very large and diverse paleotropical genus of bout 50 spec ca. 300 s s; there ar les in Africa/Madagascar, over 200 in tropical Asia from India to New Guinea, and a few in Fiji and o KEY TO THE GENERA ОЕ S la. те v ae pistillate disk unlobed to 5-lobed. erula о Ф = < p- о Webster Synopsis of Taxa of Euphorbiaceae Subtribe ee оа INAE урлык i ac. Sci. Univ. Tokyo, Sect. 3, Bot. 6: 301. Tribe Acalypha subiribe аз B Cla Pax 1969. ТҮРЕ: Claoxylon A. Juss Tribe Mareyeae Hutchinson, Amer. J. Bot. 56: 751. 1969. ТҮРЕ: Mareya Baillon cious or dioecious ures shrubs, or herbs; altern x eglandulas; Bowers apetalous; staminate sepals d D: Tu (rarely absent stamens 5- -40(- 200), free; anther sacs separa o the connective, 4 erect; dom gra Meses spiro pistillate pepals 2-4, о or ly 2- or 3-locular; styles + Sank bnt but lacerate; fruit capsular; seeds ecarunculate, testa isi fleshy paleotropical subtribe o enera, some hanh distinct, and others not je belonging her 168. Erythrococca uds ate; stipules m ud te; ther sac Staminate disk of intersta 5a. re 6a. Stamens mos ii: ister deciduo niformly floriferous; m D coriaceous; leaves not stipellate. minal segments. s erect; mostly dioecious. 20 or more; styles recurved, papillose but hardly duh ^s "T rrupted; capsules crustaceous; pini stipella Cla xylon 170. Соор ; monoecious . . Mar tyles A edi di ОА м те . Micrococca 1 b. e мш, ola dii 8-10-lobed; staminate disk of interstaminal segments; stamens up k :25 174. EER A 168. Erythrococca Bentham, ie» Fl. 506. 1849; Muell, Аг 15(2): 790. А 400. 1954; Ral Sith, FI E Afr. uphorb. 1 E вне а пот. Шер. [= thrococca а mala (Juss. ex Pak ) Prain]. Po sBrophyton Pax, Bot. Jahrb. 19: 88. 1894, TYPE Жы: a лын aculeatum Pax [= Erythrococca нне e Pra et Chl b. 26: 383. 1899. TYPE: Chlo oropat ane айкала (Bailon) Tnd [= Ery- thrococca africa арч оп) Prain]. Athroandra (Hook. DE loft. Pflanzenr 147. II 63): 76 1914. E: Claoxylon vpn Prain = Erythrococca mannii (Hook. f. А diverse African iege of ca. 50 species, whose delimitation apparent analysis. Although Pra pou Athroandra as an of Repth boats Airy Annals of th Sedul inna Garden (1966) suggested that Erythrococca is close to Claoxylon. е, Claoxylon A. Jussieu, Euphorb. Tent. 43. 1914; Merrill, ai hil Pl. 2: 4 3; Whitmore, Tree Fl. Malaya 2: 78. 1973; A . Smith, Fl. Vit. Nova 2: 516. 1981. TYPE: Claoxylon parviflorum A. Jus er уок € x Blume, Bijdr. Fl. Ned. pee 614. m trois i pg Reinw ВІ. [= Clay m (Rein x Bl.) Hk desig- ted by ан 1975]. Quadrasia pow Leafl. Phil. Bot. T 2656. 1915. TYPE: iadrasi — Claoxylon eu- phone (Elmer) Merr.]. А paleotropical n of c from Africa but e Melanesia and Haw ca. 75 species, absent enge: from Madagascar to id uc apis Leandri, Bull. Soc. Bot 85: 526. 1938; Radcliffe-Smith, Kew Bs. 43: 642. hk, TYPE: Claoxylopsis per- rieri Leandri. A genus of three species to Madagas- car. es apparently very close to Claox nee it is still poorly known and its = sti and status ust be regarded as tentati К. ае ВаШоп, Adansonia І. 1: 73. 1860; ‚ DC. Prodr. 15(2): 792. 1866; bu Hook. 16. Pi ES: 63, t. 128. 1879; 3e, XIV oo bay 11. S. Léonard, Bull. Jar ..29:.291. ; Radcliffe- ue Fl. E. "Tro Аб: LR 1: 216. 987. TYPE: Men spicata Baillon. d Pax & Hoffm., Pflanzenr. 147. XIV (Heft . 1919. TYPE: "Mare reyopsis longifolia (Pax) h Hoff m. [= Mareya longifolia Pax]. An African genus of three species. 172. Discoclaoxylon (Muell. Arg.) Pax & Hoffmann, Wiss. E 912; Pflanzenr. 147. x Жз 63): 137. 1914; Radcliffe-Smith, E. Trop. Afr. Euphorb. 1: 279. e Discoclaoxylon Muell. A Flora 47: 13 4. TYPE: SORE гин, hexandrum ub ell. Arg.) Pax & Hoffm. [lec- e totype, chosen here; ms larger staminate disk in the lectotype species appropriately reflects the taxon name]. mainly west African genus of three species, extending from Sierra Leone and Fernando Póo to ganda. 173. e Niger Fi. 503. 1849; Gen. Pl. 3: 309. 1880; Prain, Ann. Bot. 25: 628. -— Fl. Trop. Afr. 61) 816. Dyer, Gen. S. Afr. Fl. PL, ed. 1975; Radcliffe-Smith, Fl. E. Trop. Afr. Eu- 7. TYPE: Micrococca mer. curialis (L.) Benth A paleotropical genus of 12 species, repa from tropical Africa and Madagascar to Malay та А тугеа Leandri, Notul. Syst. (Paris) 9: 68. 1940. TYPE: Amyrea sambiranen nsis ta нин selected here ic to Madagascar. | Amyrea diverges "i om all of the к: genera of subtribe Claoxylinae in its bifid styles, and its sition requires evaluatio: Subtribe 30h. BONON Radcliffe- Smith, Kew Bull. 9. 1989. TYPE: Lo banilia Radcl.-Sm. : е | oecious trees or shrubs; indumentum stella | deciduous; inflore: ary, г cemose; flo ax er- ers apetalous; staminate sepals 3: disk of int culate. | This monotypic subtribe, recently proposed | by Ragelide iib. appear s very similar to dumentum Е t 2i. he бепан is Балуч какыр һеге until detailed we parisons with taxa of Clao can be made {| determine whether Lobanilia should be kept in separate subtribe. 175. Lebanilia bae iun Kew Bull. Eel 334. 1989. TYPE: Lobanilia luteobrum | (Baker) ы Сы, | A genus of seven species confined to Мада | car. Originally these were treated as а section 0 " Volume 81, Number 1 1994 Webster Synopsis of Taxa of Euphorbiaceae Claoxylon, but Radcliffe-Smith presented persua- : SA A Ё E = 3 lg ко МАЕ бике pi : 339. 1841; Webster, Taxon а а элыш ТҮРЕ: Rottlera Roxb, W Mal- T Family Trewiaceae Lindley, Intr. Nat. Syst. ed. 2, 174. 1835. Acalypheae subtribe Mercurialinae series Tre- willormos Pax & Hoffm Mon 147. VII (Heft . 1914. TYPE: Tribe Acdphese s subtribe Сагды ае » Muel 1. Arg., Lin- 1865. TYPE: Doslodiveus Baillon [= Ma de art Tribe Maleae diss. Amer. J. Bot. 56: 752. 1969. E: Mallotus Lour nate or opposite, err or lobed, pinnately or palmately veine imes with embedded lam- inar Bland: “stipules presen or obsolete; inflores; flowers apetalous; staminate sepals 3— 5; disk trastaminal, of interstaminal segments c nate; ais unlobed, + нш ог каше, {гий capsular or runculate), testa often fleshy. \ This Old Y 14 Hesh f eight | ly related ioecious (ra arely monoecious) trees or shrubs; g gly rep li 1 Asi indumentum stellate (rarely simple); leaves alter- KEY TO THE GENERA ОЕ SUBTRIBE ROTTLERINAE la. Carpels 2-4 (rarely 5); fruit capsular or Tense. 2a. Pi ы unlobed; leaves alee lan dul ruit capsular, dehiscent; pistillate sep connate. 4a. Un smooth t inate, not winged axillary as well); bracts small; stamens 20-200 „рн ө sexine tectate- peto to rl 5a. Styles elongate; leaves usually ыйы, -glandu lar 176. Mallotus 5b. Styles contracted; leaves eglandu es EAT : ~- 177. Deuteromallotus 4b. Ovary + winged; M. UH 6a. Stamens many __ 178. Cordemoya 6b. Stamens 15-20; иы caruncul: 179. Coccoceras 3b. E indehiseen ghey calyx gam s. А pan e disk absent; ovary 2- Allo cular 80. Trewia scd disk наан ovary 1(2)-locular ..... ndul: T 2b. Styles bifid; xis egla Жур trewia lb. Ovary 8-locular; ; fruit joris: leaves alternate, glandular; stamens 15-20 ч e Loureiro, Fl. Coch e ; Schum. & P. Afr. Eup 987. PE: Cog EEE Lour Echi j tnus Loureiro, F]. Cochinch. 633. 1790. TYPE: Echi- 181. 182. Rockinghamia 183. Octospermum nus trisulcus Lour. [= Mallotus cochinchensis кошта олш n Coromandel 2: 1798 (non Willd., 1797). TYPE: Rottlera obs Roxb. [= Mallotus philippensis Sg ) Muell. Adisca Blume, Bijdr. . Ind. 609. 1826. TYPE: a se (RL) M ell. Аг et е, ед һег е}. : Rei "D m ger, Verh. Natuu rk. Ver. ` Ned 1(4) . TYPE: Plagianthera op- posi (ВІ. ) паа Y & Zoll. [= Mallotus blu- DI. j Voy. Herald 409. 1857. Mallotus чанае Hana Seemann Hancea hookeriam a Seem. [= u —— ell. Arg.]. Axenfe anus pisa tude Gén. Euphorb. 419. 1858. е rE y rini Baillon [= Mallotus muricatus (Wight) Muell. Ar Солана ‘Baillon, Etude Gén. брой 293. 1858 ТҮРЕ Е: Ricinus көрт T ill. [7 Mallotus arpus (Th Aconceveibum Miquel, Fl : Aconceve cibum: trinerve Miq. [= uell. Arg.]. Fragm. Poeni: Austral. ges Bat. 1(2): 389. 1859. Mallotus phili ippensis (Lam. )M Gens F. Mueller, Annals of the Missouri Botanical Garden 1: 31. 1859. TYPE: Echinocroton claoxyloides КЁ. Posi [= Mallotus claoxyloides (F. Muell.) Muell. rg.]. Bod Muell. Arg., Flora 47: 539. 1864. TYPE: ee chlamys griffithianus Muell. Arg. [= Mallo- us griffithianus (Muell. Arg.) Hook. f.]. This large and complex paleotropical genus cludes ca. 150 species, of which there are ^ two in Africa and Madagascar, the rest in Asia and Australasia (to Fiji). 177. Deuteromallotus Pax & Hoffman Pflanzenr. 147. VII (Heft 63): 212. tis i aud Bull. Soc. Bot. France 103: 605 956. TYPE: Boum allotus acuminatus ا‎ Pax & Hoffm A genus of two speci demic to Madagascar. 178. Cordemoya Baillon, Adansonia 1. 1: 255. 1861; Pax а Hoffm., фы 147. ҮП (Heft 63): 208. 1914. TYPE: Cordemoya їп- tegrifolia v. ex Bail, о A monotypic genus endemic to Malagasia. 179. Coccoceras Miquel, Fl. Ind. Batav. Suppl. 455. 1860; Muell. Arg., DC. does 15(2): 949. 1866; Bentham, oe PS Pax & Hoffm., Pflanzenr. 147. Үй dE 63): 209. 1914; Airy Shaw, Kew Bull. 16: 349 1963. TYPE: Coccoceras sumatranus Miq. A tropical Asian genus of three or four species, distributed from Burma to Borneo. Airy Shaw (1963) reduced Coccoceras to a section of Mal- lotus, perhaps correctly so. 180. Trewia L., Sp. Pl. 1193. 1753; Klotzsch, A Add. хач . 4: 200. 1975. TYPE: Trewia nu- notypic genus, the single species widely nd distributed from India to southern China and In- don 181. Neotrewia Pax & Hoff ffmann, Pflanzenr. 147. beide 211. 1914; Merrill, Enum . 2: 437. 1923. TYPE: Neotrewia cum- ingii i (Muell. Arg.) Pax & Hoffm otypic genus of th арт and Cele- Am bes, дажы, separable from Trewia were йын Airy Shaw, Kew Bull. 20: . 1966; 35: 667. 1980. TYPE: o" Ls е (Benth. ) Airy Sha A tralia MU ic to tropical Aus- 183. Octospermum Airy Shaw, Kew Bull. 19: BUB M dde Hook. Ic. Pl. 38: t. 3716. 1974; Kew Bull. Add. Ser. 8: 176. 1980. TYPE Airy А single species of New Guinea. Subtribe aga POTERE a Fl. Br. 45. 1859; P еы pleiogynum (Pax & Hota Shaw ., Pflan- zenr. нй XVI (Heft 85): ds: 1924. TYPE: pha L. Acaly pieno or dioecious trees, shrubs, o r herbs; indumentum simple or stellate, often glandular; leaves "eae m peer or pina lat tillate (in a [ge pecies) wen te, ter Ew 4 axillary, наар or bisexual; pistillate bracts ust- ally enlarged in fruit; staminate sepals 4; dis sent; ovary (2-)3-locular; "a mostly free, ^ | iate (rarely TA fruit capsular; seeds runcle minute or absent. k | | is here defined, subtribe Acalyphinae inc M | only the genus Acalypha; it appears closely relat to dui m Claoxylinae, but differs in its unique anthers, enlarged pistillate bracts, distinctive PO ate pollen, and dissected styles. Ин, Agrl Sp. Pl. 1003. 1753; GF | . 5, 436. 1754; Muell. Arg., DC. Ргой | aay 199. (us Bentham, Gem Pl. 3: 311. | 1880; Hutchinson, Fl. Trop. Afr. 6(1): pe | 1912; Pax & Hoffm., Pflanzenr. 147. ХҮ | (Ней ж 12. 1924; ; Natürl. Pllanzenfam. 9t ed. 34. 193 ignated = Small, in Вг - & Brown №. 0.5. ed. 2, 2: 457. 3]. Caturus L., Iyn Nat. ed. 12, 2: 650. 19, 127. 1767. TYPE: Caturus Morte Acalypha iida Burm. f.]. — "LE | Volume 81, Number 1 1994 ронку" Klotzsch ex Schlect., Linnaea 19: 235. 1846. E: Linostachys padifolia Schlecht. [7 Acalypha sari ала Muell. A Odonteil Turc sak Bull. So p. Natur. Moscou E: Odo. suce. claussenii Tur urcz. nd calypha VASA (Turez.) M es kl: onda Grisebach, Bo: onplandia 6: 2. m . TYPE: Gymnaly, . Wat cad 451. 1887. TYPE: Corythea fpes pon P VA м filipes (S. Watson) McVaugh & Acalyphopsis Pax & Hoffm., РАС do VI (Heft 85): 178. 1924. TYPE: Асау hopsis celebica Рах Hoffm. [= Acalypha КАЛМАК Hurus.]. With ca. 450 species, Acalypha is the fourth largest genus of Euphorbiaceae; in addition to the small number of lemperate species, species are wid А +} 1 1 z н and a few Pacific archipelagos. MENS: ng Pe ig Ninh del Webster, 1975. ТҮРЕ: Lasiococca noecious or dioecious trees or shrubs; indu- nise the ti Loue E bracts dS sta- nate 8 Коше) pom on filaments of ERES M to papillose ta usually fles This sie includes three genera that w treated by Pax & Hoffmann (1919) as part ot sioe Ricininae, doubtlessly because of the an- ый many stamens with anthers borne ranches о ndritic column d - Airy Shaw (1974) has suggested a baie relationship of subtribe Lasiococcinae to Mal- s. Key | TO THE GENERA OF SUBTRIBE LASIOCOCCINAE à. Mo noecious; ages sepals uec indu- lb. D; mentum sim muricate — — vary ; istilla t 2 ttn Le he vw sepals dade indünsen 2а. idote Lococca dd. absent ovary tuberculate; 186. paeme 2b. lero se seals present ovary pemi ; pol- 87. EN pha jacquinii Griseb., nom. illeg. fe Aca- . a aen] Webster 91 Synopsis of Taxa of Euphorbiaceae ая: пие Hooker f., Hook. Ic. Pl. 16: 5817 Fl. Brit rie 5: Fok 1887; 104. 1973; ' Thin, J. Biol. Sinh Hos 8(3): 36. 1986. TYPE: Lasiococca symphyllifolia (Kurz) Hook. f. A tropical Asian genus of three species, with a e Vy M in India, Malaya and Indo- china, and Hai 186. amr AN OLE Blume, Bijdr. Fl. Ned. Ind. 62 у 196. 1975; 8: 202. 1980. ТҮРЕ: Epa Инрйетоп javense Bl. Polydragma Hook. f., Hook. Ic. Pl. 18: t. 1701. 1887; Fl. Brit. Ind. 5: 456. 1887. TYPE: Polydragma mallotiforme Hook. f. [= A eel javense BI.]. Ca Sp. Moo YPE: Сонун ўя 5р. Мооге [= Врв forbesii (Sp. Moore) Airy Sha w]. . Bot. 63 Шр 101. 1925. Spathios- A [iim] of two specie bodie "et Thailand igh united with Menor noia by Mueller (18 visis Bentham (1880), and Pax & Hoffmann (1919), it may be distinct, as indicated by Airy Shaw 187. Homonoia Loureiro, si Cochinch. 636. noia riparia Loi 1837. TYPE: Lumanaja cepi riparia Lour.]. Lumanaja Blanco, Fl. Filip. 8 fluviatilis Blanco [= of two Asian species (India to Taiwan, the ү mare and New Guinea). Tribe 31. oe ا‎ Hutchinson, Amer. J. Bot. i бен аад n. Pl. 3: 252. 1880. TYPE: Plukenetia a Monoecious (rarely е trees, shrubs, о her ning; = mentum бене, often glandular, ее urticant; leaves alternate, unlobed to palmately sat pinnately to palmately veined, sometimes glandular or sti- | 92 Annals of the | Missouri Botanical Garden | [ pellate at base; stipules present. Inflorescences ter- mann (1919, 1931). In contrast to subtribe Tra: * minal or axillary, usually bisexual, mostly racemose — giinae, the distribution of the Pluk ieae is al or spicate; bracts eglandular; flowers cpg entirely neotropical (a few species of Plukenetia Sta [ш dn splitting into 3-5 valvate seg- occur in Africa). extrastaminal, or di stamens (2)3-100, usually free; anthers introrse KEY TO THE GENERA OF SUBTRIBE PLUKENETIINAE extrorse, muticous or apiculate; pollen in la. Ovary алар МИМ, sepals 5 ог 6; trees, e соросае, porate, or Aper с. sexine varying shrubs i Pistillate 2a. 3 staminate disk segments 4 (or als 3-6, imbricate; disk absent; ovary 3- if absent аа by 4-lobed ر‎ i leaves рї dria DU trees or shrubs 4-locular; styles unlobed, slender or dilated, some- EU Gass of Hub dilated! mes р ose or lacerate, usually connate into a formi: E an intrastaminal pseudodisk; distinct column. Fruit capsular; columella Ata style urceolate ......... 188. Haematostemon tent; seeds поа: їезїа mee endo- 3b. Disk наат: style ovoid ...... 9. e | | * iid broad- 2b. Stamens 10 or more; staminate disk se Өг than radicle. ments absent or minate. numer bis de Stam te; Пом | This distinctive tribe includes 16 genera with е REEL pn in oa basis c: pin- | more than 250 species in the tropics of both the nately veined; tree or shrub О Old ма, and New World: the majority are neo- 190 Angostyles trop Ab. Sta 10 1, clavate, | ар 8 lobed; flowers in huc ra- | emes; leaves palmately veined; twin- | KEY TO THE SUBTRIBES OF TRIBE PLUKENETIEAE 198 Do; | la. Flowers in racemes or ipe s, not bibracteate lb. Ovary 4- bodie pistillate sepals 4; twining vin | pseudanthia; pollen grain s subglobose. or lianas. i 2a. Stinging hairs absent; fruit ined crested, 5a. Je mostly free, connate for ү сети, or indehiscent; De grains inel their d b m bifid; m | colp. Bib a NAT lukenetiinae 40-5 д 2. Eleusherosis 2b. risu hairs present; fruit Men dom nap- 5b. Styles entir ely < t iiy ed; pollen grains 3-colpate, 3-porate, mn; stamens 8 40). | s inaperturate, rugulose ...... 3lb. Тайне ба. Fr Seneca ek indehiscent, but | 1b. Flowers in bibracteate pseudanthia; pollen grain then staminate receptacle n not d E | proie coarsely reticulate; stinging hairs pai 93. Plukene! ento eus o ACC E 31c. Dalechampiinae 6b. shit indehiscent; Vara Er Í globus kii son ы шш 4. Vigia | Subtribe е soe ea em 188. ores ts (Muell. Arg.) : i Gen. Р 3. 1880; Pax, Natiirl. Pflan Hoffmann, Pflanzenr. 147. IX (Heft 68): 3 | zenfa ed ‚ 3(5): 62. 1890; Pax & Hoffm., 1919; Sandwith, Kew Bull. 1950: 133 1951; Pflanzenr. 147. IX (Heft 68): 1 ш Na- Jablonski, Mem. New York Bot. G türl. Pflanzenfam. ed. 2, 19с: 1931; 143. 1967. Astrococcus sect. Haematosl? Weise Tad 24: 598. 9 TYPE: Plu- mon Muell. Arg., Linnaea : Р Б kenetia L. TYPE: Haematostemon coriaceus us (Вай) " & Hoffm shrubs, lianas, or ; indu- mentum ا‎ of simple se leaf blades simple and unlobed, entire n ЕД E В 8 S pose ns nthers subsessile; pollen g grains pan to sane, tricolpate, t 0 di buit e ê sty "M es massive, partly to e connate; fruit capsular or rarely indehiscent. As delimited by kira (1975), subtribe Plu- kenetiinae is here treated in a narrower sense than the concepts of Baath (1880) or Pax’ & Hoff- genus of two species from rainforests in south- ern Venezuela and Guiana. The gynoe oecium is Ve d similar to that of жи га the staminale — flowers are quite differen m. 30. 1919. TYPE: Astrococcus cornutus A monotypic genus from Amazonian Brazil and nezuela. Volume 81, Number 1 Webster 93 1994 Synopsis of Taxa of Euphorbiaceae 190. Angostyles Bentham, Hook. J. Bot. Kew (Muell. Arg.) Pax & Hoffm. [= Plukeneti ph Gard. Misc. 6: 328. 1854; Muell. Arg., DC 3. "M jd wm ^ Ta podandra Pax ve 147. IX (Heft oe Prodr. 15(2): 767. 1866; Fl. Bras. 11(2) 20. 1919. TYPE: yee hei tne (Ule) P 331, t. 50. 1874; Bentham, Gen. Pl. 3: 32 & Hoffm. [7 Plukenetia bes 1880; Pax & flanze 147. qi rens Ducke Ann. Jard. Bot. Done 4: ff an IX (Heft m ЕЗ 1919. ТҮРЕ: reai lon- gifolia B A monotypic genus from the Brazilian Amazon. 191. Romanoa Trevisan, Saggio Monogr. Algh Cocc. 99. 1848 (nom. cons.); Radcliffe- Өй : Anabaena tamnoides A. Jus E иші, tamnoides (A. Juss.) Radel. m.]. otypic genus of Brazil, e Cia close to Paget and questionably distin 192. Eleutherostigma Pax & Hoffm Pflanzenr. 147. 1X (Heft xs 11. 1919. TYPE: Eleutherostigma lehmannianum Pax & Hoffm. A poorly known monotypic genus of Colombia and Ecuador, apparently closely related to Plu- kenetia, and combined with it ud Gillespie (1993). 193. mete L., Sp. PI:1192...1753; Gen; Pl. ed. 5, 438. 1754; Muell. Arg., DC. Prodr. 15(2): 768. Eo sane ad Gen. Pl. 3: 327. Hoff 1880; nzenr. 147. IX (Heft 68): 12. 1919; alo Mem. New ork Bot. Gard. 17 967; Webster, Ann н Bot. С к 4: 293 8; Huft, Ann. Missouri Bot. Gard. 75: 1105 1989; Cik espie, Fu Bot. e 575. 1993. TYPE: Plukenetia volubilis P, po Hasskarl, Flora 2009) Beibl. 41. 1842 (nom. “Bull. Airy d узуны ‚ J. Arnold Arbor. 22: 423. 1941; ЖЫ? Ке ети d. Ser. 4: 187. 1975. Sajori occus Meissner, Pl. Vasc. Gen. 2: 369. 1843 A Endl n. Pl. Suppl. 3: 98. 1843. TYPE Voies "occus glaberrimus Hassk., nom. Шер. [= Pl Hedrai orniculata J. E. Smit im. xs Ne Tijdschr. Natuurl. Gesch. TYPE: Hedra rimus Hai ы. da Ж aiostylus pias th]. LE Plukenetia corniculata Te = idium Pax, Bot. Jahrb, 26: 329. 1899. TYPE Age rpidium staudtii Pax [— Plukenetia co- га Muell. Ar. xr. Il. Herb. Boiss. II. 8: 635. 1908. corniculata ги seeds Pax [= Plukenetia 17. m. Hoffm., Pflanzenr. 147. IX (Heft 9. TYPE: Angostylidium conophorum 1925. ТҮРЕ: Elaeophor WES ie Ducke [= Plu kenetia "abutifolia (Ducke) Pax & Hoffm.]. А Jim of about 15 E mostly шогон а, M frica and one in southeast Asia. The genus is “rather variable, and a number of workers have accepted either Pterococcus or Tetracarpidium as distinct OW genera. How may no bend à im since some of the other genera of вае Fitikenetiinae listed — (e. £^ Vigia and ^ win i fiis Plukenetia. v Bus Vellozo, Fl. Flum. 9: t. 127. 1832. E: Vigia serrata Ve Fragariopsis St. Hilaire, Теса Morph. Veg. 426. 1840. St. Hil. [= Vigia ѕег- apes Шр Veg. 499. 1840. TYPE: Vigia serrata Vell. ]. Boryone пастет Кат ОЬ 7(1): 190. 1841. Botryanthe discolor К. [= Vigia serrata Ve "n uth American genus, eres monotypic, 31) rec a гё IL} Ac cia St dum sally used for this vide from Рисан si since 858), the name Vigia ru valid as ee "vind — of Fragariopsis by St There seems no p reason ropose the name Fragariopsis for rva- tion, irum dn since it is only ddr eu from Pluken oe edes TRAGIINAE Hr Taxon 1975. TYPE: Trag оде. ог herbs, often twining; iumentum тн oe ntire or зле; without ا‎ о ш seer not сеча inflorescences axillary ог ter- and о e leaves; stamens (2)3-50, fila- ments jar or spp N pollen grains subglobose to suboblate, 3-colpate, bs a is HM. or р DE E e. s 3 f 3-locular; че nearly ча lar to io détails connate; fruit capsula 94 Annals of the Missouri Botanical Garden here delimited, tl btrib ludes 8 g Nair, Gardener’s ыг? 31: 49. 1978. TYPE with ca. 150 species; besides the widespread Tra- Cnesmone javanic gia, there are 5 Old World and only 2 neotropical genera. KEY TO THE GENERA OF SUBTRIBE TRAGIINAE la. Staminate — d inflexed to form a seudodisk; stamens 3 or 4, anthers introrse; e sped 2a. Pistillate woe entire or ee pistillode algal minate flow 1 a. соппес- tive е triangular, sometimes audate; stamens free. 5s Styles free or nearly so, long-pap- i on adaxial surface; inflores- cence terminal and leaf-oppo leaves with stinging hairs . 5. Cne nesmone 4b. Styles connate into a a massive glo- ар1 il- p osed; > lose; inflorescence и jis subglabrous ..... 196. Megistostigma 3b. Pistillate sepals toothed; anther nective: nok е nlar arged or caudate; n column a inflorescence ter ermi nal and leaf- dice емш subgla ves 198. Tragiella pete ally ed; stamens 0 + deck Msn or extrorse; styles usually hen er. 5a Anthers 3 or more, if 2 then not Е staminate calyx concave; styles free хари connate. Anther connective bean tuft of stinging hairs; stam subs, eed be 29 usually т Ta. Styles puse papi ir bi eptacle con — s (2)3- many; OF ueri vines, O e SLAE N SR 7b. у ma not а Had sta- nal r acle plane; stamens 50). ERE 200. Tragia 6b. Kui connective ending in tuft of stinging hairs; erect shrubs, leaves gla- brescent, di 201. Acidoton Sb. Anthers 2, subsessile; staminate calyx flat with gis pe. style: елат into thick с Pd E РЕЩ os Peers н > Bijdr. Fl. Ned. Ind. 630. Fl. Javae vi. 1828; Muell. ., DC. Prodr. 15(2): 926. 1866; Bentham, 7. 1941; Airy Shaw, w Bull. 26: 240. 1972; Balakrisnan & hi Gagnep., Bull. Soc. Bot. France 71: 866. 192% 1. Gen. ا‎ 5: 386. Ke ТҮРЕ: Cents tonkinense nep. [= Cnesmone tonkinense o ai ) Croizat lectotype, designated Wheeler, n 24: 534. 1975]. This Asian genus of ca. ten species is here de limited Mem Croizat (1941). is retia Hooker f., Hook. Ic. Р. | 1592. 1887 | Croizat, Arnold Arbor. 22: 425. 1941; | haw, Bull. 23: 119. 1969. Merson malaccense Hook. f. е кор Мед. Landb. 10: E f 1910. TYPE: Cats pelts J J. Sm. [= gistostigma peltatu roizat]. tropical southeast Asian genus of five specl® | distributed from Burma to China, the P ips | and Borneo; combined with рин by Pax | & Hoffmann (1919) but maintained as distin ct by Croizat (1941). | 207 Rd sai pastes pois Etude Gén. I 1858; Mue E IX (Heft 68) 106. 1919; Croizat, J. А т . 1941. TYPE: Sphaerel ls Baill. A genus of t lemic to Мей | E: pice P adr pin Pax & Hoffmann, | IX (Heft 68): 104. 1919; in | Smith Kew Bull. 35: 777. 1981; Fl. E. p fr. Euphorb. 1: 318. 1987. TYPE: Wes natalensis (Sond.) Pax & Hoffm. [le ectotyP^ | ] chosen African genus of five species, extremely dec ш Seen and doubtfully separable. — — ME Aare doe Bull. Bot. "E 1830; ll. Arg., DC. Prodr. 1 | nee 1866 ш Platygynes Pens PE 3:328; x&H 147. IX mee "i En (919: Ati "ros | Yor t Ме. 21: 132. 1971: шы Ann. ны, "€ ! Nat. Hung. 6 1972. түрЕ: Plat (jact) urens ye ч е pue Muell. Arg.]. Volume 81, Number 1 1994 Webste 95 Syno al аа of Euphorbiac Acanthocaulon odi ex v Std им еп. у erd Kn 88. 1850. Endl. [7 Жа па я Уе ы ) Muell ju. T n the most recent revision of Borhidi ае ic t onym of Tragia, a disposition that merits further study 200. vade Plumier dns Sp. Pl. 980. 1753; Gen. Pl. ed. 5, 421. 1754; Muell. Arg., DC. Prodr. cm 927. es Bentham, Gen. PI. 880; eai Fl. Trop. Afr. 6(1): 964. 1913 f ах & Н ., Pflanzenr. 147. (Heft 68): 82) yw Pee ain, Fl. Capensis 5(2): 5 192 ourteig & O'Donell, Lilloa 6 347. 1941; Johnston, Rhodora 64: 137. 1962 Miller & Webster, Rhodora 69: 241. 1967; Webster, J. Arnold Arbor. 4 1967; eandri, Adansonia II. 11: 437. 1971; Rad- cliffe-Smith, E. Trop. Afr. Euphorb. 1 01 Ann. Missouri Bot. Gar 1106. 1989. T e | ү ley gaiga by Sm in Britto Brown, Ill. Fl. N. U.S., ed. 2 2: 458. m Bia Klotzsch, Arch. Naturgesch. CL) a! ME TYPE: ч Io gi Na Aor 7(1): 189. © Leptorhachis hastata Kl. [= Tragia L a (Kl. ) neue Piers Klotzsch, Arch. Ne aturgesch. 7(1): 188. 1841. E: Leucandra b o viden KI. [= Tragia leu- y, Hoo Кой) Bot. 1: 29. 1842. woe | omeria cordaia Harv. [= iris capen- un| 2 Baillon, Etude Gén. Euphorb. 464. 1858. ТҮР с Жарыш Baill. [= Tragia scandens (Baill.) candra Pax CEE Harve pax Z coriis de Gén. Euphorb. 495. 185 58. TY aig v ie Baill. [= Tragia bailloniana A large and diverse aos of ca. 125 species, a he gne is m Zuckert rtia ma 5 "ец жы. Y prove to merit segregation on fur 20 1. cop Swartz, Prodr. 6, = бы (пот. DR - Ind. Occ. 2: 952, . 1800; ek Arg Arg., DC. Prodr. you чуч 1866; S ntham, Gen. Pl. 3: 328. 1880; Urban, Ymb. Ant. 7: 513. 1913; Pax & Hoffm., Pflanzenr. 147. IX que =ч 24. 1919; Fawc. & Rend., Fl. Jam Ann. Missouri Bot. Cad 289. 1968; Liogier, Fl. Batole 4r 71. 1986. ТҮРЕ: Acidoton urens Sw. paris Pax & Hoffm., Pflanzenr. 147. bb Soh ay 87. died TYPE: Gitara venezolar je Acidoto otropical genus of six or seven species, in the Greater Antilles (Jamaica and Hispaniola), Central America, and northern South 202. sis wage Pax & Hoffmann, Pflan- X (Heft 68): 108. 1919; Backer & B Es Java 1 : 491. 1963; Airy Shaw, Kew. nr 23: 115. 1969. TYPE: Pachystylid: ium aout (Bl) Pax & Hoffm. [Tragia hirsuta. Bl. typic genus of tropical Asia (India to a Java, and the Philippines; not reported from Sumatra or Born valise 31c. DALECHAMPIINAE (Muell. rg.) We alecham- 4. 1864; s eus 1232. 1866. TYPE: Dale- mune Plum. ex Monoecious шин Шш ог more спі о bering or ing hairs {бешен almost abenn; leaves Ше le, lobed or partite, usually nate, blades simple, or partite, йер р stipules три infloresc cenc danthial, subtended by a pair of usually showy involucral bracts; eiit lowers 3 per pse eudan- t ium, in a basal c i es pseu- with mostl -1 wers; staminate ir, mostly resiniferous; stamens 8-10 re, filaments connate; pollen grains prolate, mo a = ex imi pias with ышкы ed re eticu- асе оуагу 3- locular; stylar column caen stigmatic bad extending down upper / to %; capsular. т This subtribe contains only the pud genus Dal- rank b SE as links t ing hairs) and ERN (elongate stylar col- mn). 203. Dalechampia Plumier ex L., Sp. Pl. 1054. 1753; Gen. Pl. ed. 5, 473. 1754; sis y. Annals of th Missouri итна Garden es Prodr. 15(2): 1232. 1866; Fl. Bras. 11(2): 633. ede Bentham, Gen. PI. З 0. 1880; flanzenr. 147. XII ч 68): rd. 54: 308. 1968; Webster & Arm Pres Brittonia 31: 352. 1979; sieh Bot. 7: 1982; Armbruster, Syst. Bot. 9: 272. ae Rae Smith, Fl. E. Т ме Euphorb. 1: 285 tonia 4] 1989; Webster, Brit 1. 1989; Webster & Armbruste t inn Soc. 10 37. 1991. TYPE: Dalechampia Cremophyllum eres Bull. Acad. Roy. Sci. Brux (1): 23. 1842. TYPE: Cremophyllum spathulatum ale Pini: spathulata (Scheidw.) ' Bail Megalostyli Spencer Moore, J. 8. egalostylis Борн = as ie »^ T ns micrantha Poepp.]. А genus of slightly over 100 OY about 9 merican, with a fe and фо ster xd Armbruster 31) рахе recog- leise ut т. Mies & Hoffmann) mid Dn ae ns & Hoff Pflanzenr. 147. IV (Heft 52): 14. 1912. TYPE: Omphalea L. once bas shrubs, or lianas; "m with Ра ч simple or lobed, pinnately or r palmately ined, 2 sessile basal glands; ENT. decid inal о or axillary aments coded anther con- re n шош oblate, 3-colporate, sexine finel tate-sp жыз 1 гея imbricate; connate into Fruit thick-walled, caps large, globose copious. кл. г ‚ ecarunculate, testa dry; endosperm This tribe contains the single genus Omphalea, whose affinities have been controversial for тап h ussil р maneae. Mueller (1866) kept it in the tribe Hip- pomaneae, in subtribe Gelonieae. Pax & Hoffmann 1912) created a special subtribe of Hi aneae tr! Croi zat (1942), in contrast, pointed out affini with the tribe Plukenetieae, кые Pent (1962) ough Omphalea shows striking resem: a number of char- its affinities appears to be critical in the phylogeny of the uniovulate Euphorbiaceae. m MAn зир L., Syst. Nat. ed. 10, D V (Heft 52) 14. 19 | . 1920; Coat Bull. D аа Bot. к. Onghdica leandra L. (ty cons.) 9. Ronnowia Buchoz, Pl. Nouv. Decouv. 6, t. 4 pu PE: Ronnowia че Buchoz (пот. | [= Omphalea triandra 1,05 | Hecatea (алкым rM d ce — A er Om 1804. TYP a oppositifolia оу} | phalea зрнаца (Willd.) rd к гацен. by illespie Hebecocca Beur! Me Vetera Akad. Handl occa XT ‚ 147. XIV ( P ax 1919. TYPE: Neomphalea. papan б | 54 & Hoffm.) Pax & Hoffm. [= Omphalea ари? Pax & Hoffm.] | A genus of са. 15 species in both the Old Worl and New World, but with the greatest specia centrations in the Greater Antilles and Madag Volume 81, Number 1 1994 Webster Synopsis of Taxa of Euphorbiaceae — ow lente ne Pax ahrb 3. 1884; Natürl. Pansat ed. T won " 1890. ТҮРЕ: Cro es, shrubs, or herbs; latex colored or absent; umentum simple, stellate, or lepidote; leaves alternate, Opposite. or rely wheres stille or pound, often ids foliar glands. Inflorescences ax- ary or ter spica te. REE FP Е" vate; pet- als and disk Козер! ог eats stamens (3-)5—- many, filame e; pollen grains bi- nucleate or trinucleate, indes to more - only p or ina rate, sexine mostly wi ornamentation of and disk p or absent; ovary mostly 2-4- | PCT e d | i \ у ); solitary ё each locule, anatropous. Fruit capsular (rarely indehiscent); seeds carunculate or ecarun- culate, testa sometimes fleshy; endosperm usually copious, often oily. As defined here, following Webster (1975), sub- family Eu includes 12 tribes. This is a KEY TO THE TRIBES OF SUBFAMILY CROTONOIDEAE 1а. d a grains colporate or porate, sexine reticulate to clavate; EI icifers articulated; pollen grains 3-nucleate; plants mostly monoecious; = 9. 3a. Pollen grains colporate; styles bifid; seeds carunculate or ecarunculate; dep much narrower circumscription than that of Pax mily Cr H (954), н include d only the tribe Crotoneae. f Mueller (1866), genera of the presen accepted Crotonoideae were dis- iip e lyphe ther subtribes had a mix- ture et Aeplyphoideae and Rratonosinse:. pinten: la 20, 38, ‘and 39; but 5 of those tribes also incl juded un" of ide Acalyphoideae or Eu- phorbioid As e eed by Erdtman (1952), pollen mor- xn has provided decisive ed in estab- es of phylogenetic affinity in the Cro- TRAT Punt (1 ou foll qi: Erdman bod oral grouped most ы the gen is “С 2 ver, “crotonoid” g oweve ful for PEE me xn and ex cept in the tr с does not help to clarify ides of tribal delimitation. petals absent; indumentum mostly simple. basic chromosome number — oly- s 3. MICRANDREAE tate, stipules free; inflorescences bier mostly AD 4а. Ройеп ртаїпз colporate; | Р spicate or panicu ate . = ENOCLINEAE flowers i in ater opposite the 4b. uni grains periporate; leave pelle ici ves . lb. Heel rains i " i ‚ 36. GELONIEAE attert n; ; petals aot, present; “indumentum simple, aper th *'crotonoi "p malpi eous, or ЫЫЫ laticifers ae ae (excep ї in some Jat ge ds without endosperm, cotyledons massive; inflorescences ; dichasial og absent; styles un- divided, dilated; latex mi 5b. otyle in Neobnutonae inflorescences various; е bifid to mul 6a. tyledons thin; petals present at : ELATERIOGPERMEAE p cept a ава, less commonly u 5 free or if co а caruneulate or vae ul A ndum imple. grim ens md more than 5, free or connat ym 9a. pacar S te pound; fruits capsular or drupaceo ` unlobed, u œ c , g . Stame ыба connate; pollen se cences terina or axillary, racemose or thyr nae сагип or ecarunculate _ ne . JATROPHEAE ose or spicate to panicula leaves .. 39. CODIAEAE — inflores- soid; кы ecaruneula 0 TRICONOSTEMONEAE 7b. Ind (at least in | Annals of th Missouri s sl Garden — о S Pall, в un e seeds ecarunculate; petals free or absent. Гопеп sex pinose or often КОКА, ч: o i in part; cotyledons broad or narrow . 41. PEAE 11b. Pollen sexinous, 10b. 6b. ignis ane Т їп үе жез plitt or drupaceous; seeds ecarunculat cenis t spinose or reduced; leaves Me eni Pus aments free; cotyledons broad rupaceous, seeds oru cres £ coherent о ОМЕАЕ 43. RICINODENDREAE 8 fruit capsular Tribe 33. a aeneus ge Arg.) Web- ster, Taxon 24: 598. . Crotoneae sub- 1 Ми an dk , DC. Prodr. 866. Gelonieae siib Chae- s Micrandriformes & . 147. XIV (Heft Mie 49. 1919. TYPE: модо; Benth Monoecious or dioecious trees; stems with ar ticulate laticifers and u: pollen таш, 3- -nucleate, ‚ colpi иеге i tillode present or r absent. ENE spe 5, im- bricate, fre 3-locular; ES free, bifid. Fruit cala nu mella persistent; seeds large, carunculate a- unculate, endosperm oily. As treated - шч и the tribe includ three or four he Mi a link etait p families jahren d "Clio e KEY TO THE SUBTRIBES OF TRIBE MICRANDREAE ta; in bud; srl rin lage imbricate or valvate - Micrandrinae ately m stamens ae 33b. bg connate, valvat Bevilüne Subtribe 33a. MICRANDRINAE Muell. Arg., odr. 15(2): 709. 1866. TYPE: Micran- dra Benth eaves simple, unlobed. pisei with кн als i aments free, + inflexed in bud; pist stillode pre pistillate disk annular. A subtribe of three South American genera. 44. ALEURITIDEAE KEY TO THE GENERA OF SUBTRIBE MICRANDRINAE la. — free; Bi eem disk present; stamens 5- 8; floral bracts small. 2a. ben elliptical indumentum simple 205. Моча 2b. Anthers linear; en stella б. Micrandrpi lb. "Ps connate; staminate disk AA stam : 8-10; floral bracts large .. 1120721 pe 205. iae Bentham, Hook. Kew J. Bot. 6:3 4 (nom. cons.); , 9 NS 709. 1866; Fl. Bras. 11(2): Bentham aie, 15: 2 : Micrandra siphonioides Benth. (typ- ih y Mil; Medd. Dansk Naturh. sip dA 144. 1857. TYPE: Po r. [7 Micrandra elata Говор iria бе п. Kj aM ке (Оа ) Muell. MN understood genus of about seven Am- azonian speci 206. роки Rodrigues, Acta Аша? ica 3(2): 5. 1973. TYPE: Ми ropsis E roxylon (W. Rodr.) W. К A monotypic genus of Amazonian Brazil. & pee sop: Mus 325 Univ. 12: ceana Baillo oem ches E Arg., dais 47: det pese pe E: Clusiophyllum sprucei Muell. Arg. [= uria ке» Muell. T rg.]. Baldwin & Schultes s det re five E cies of Cunuria, all from Amazonian South Amer a ge Schulte (1952) « on киирии =: redu f Micrandra, and because of his intimate knowledge of these South America? | Volume 81, Number 1 1994 Webster Synopsis of Taxa of Euphorbiaceae trees his opinion must be weighted heavily. The listin, ing id (шша аза separate сеня is ич һеге aly t its generic distinctness. kom 33b. HEVEINAE Muell. Arg., Lin 34: 202. 1865; DC. Prodr. 15(2): 716. 1866. TYPE: Hevea А Leaves palmately compound. Panicles with in- conspicuous bracts. Staminate sepals c 0 s valvate; disk lobed or ене sta- ments connate into a column, an- thers pores pistillode present; и disk dis- ted or obsolete; seeds ecaruncu M. А g lly Tk] 1 by ( 865, 1866), subtribe Heveinae is here interpreted to include the single genus Hevea. Pax (1910) placed Hevea in subtribe Jat стак, а рое that is соп- tr adicted by МАДЫ, oJ ber, and s Ee 208. Hevea Aublet, Hist. Pl. Guiane Fr. 2: 871, D 33 T uell. Arg., DC. Prodr. 15(2) 716. 6; Fl. Brasil. 11(2): 297. 1874; Pax, Pflanzenr m 1 191 5 217. 1935; Sh AN Mas Lea 1:21945:.13 PE: Bev guianensis Aubl. Walle N ides Richard, in Schreber, Gen. Pl. 2: 656. = Hevea gui- * anensis meta unie i i F. Gmelin qox. Nat. 2: 677. 1791. TYPE: utchoua adea Gmel. [7 Hevea guianensis bl.]. ; According to ge (1990), Hevea includes Ha species with several varieties, all found within mazonian So атф America. Because есо- по пис fee of Me genus, there is extensive be liter. i pa on syst заана. references ma 990). found in "ind (1970, 1 = 34. MANIHOTEAE (Muell. Arg.) Pax, n. e DC. pe 180 1056. 1866. ТҮРЕ: Ma. nihot Mill cious (rarely dioecious) trees, a ada cifers articulated, latex white; indumen- Mon herbs; y дошу Inflorescences terminal or sen e de y, ione late or die hasial- n St taminate fl r неч гыл intrastamin: As Оше etr ey — Ó includes two World. The close genera, b affinity between Manihot and Cnidoscolus was in- dependently confirmed on paly ynological grounds by Punt (1962) and Miller & Webster (1962). KEv TO THE GENERA OF TRIBE MANIHOTEAE . Stinging hairs absent; stamens free, stami disk intrastaminal; stam indes биси А изи: ue nish; leaf P stipellate (not ; infloresce e glandular) se o racemose- Ded 9. Manihot lb. — hairs pre esent; amens connate (very rely free); stamina te disk extrastaminal, an miles staminate perianth ee leaf moa glandular (not welt) at at base; E ^s escen ulat colis dichasial-panicul 209. Manihot Miller veri Dict. ed. 4. 1754; P Pl. 3: 306. 1880; Pax ee 44): 21. 1910; Croizat, J. Arnold Arbor. 16. 1942; Rogers & Appan, Fl. Neotrop. 26. 1989. Mandioca Link, Handb. 2: 436. 1831. TYPE: Manihot esculenta Crantz [Ja- tropha manihot L.] — unn Nov .:2:: 106, t. 81 TYPE: Janipha a HBK [= tittle aes- culifolia a (HBK ) Po! Ы Ма Eg ng. ers & Appan fi 1973 E: Manihotoides ГИС ble ) Rog- ers & ka n [= Manihot pauciflo ora Bdge.]. A neotropical genus of с . 60 species, mer native to Brazil. Rogers & rab (1973) created a new genus, Manihotoides, for an rir spe- 100 Аппа! | heys Ae Garden s in Mexico characterized by reduced inflores- ifoliolate leaves clustered on short shoots. ent discovery of another Mexican species (ined.) with unlobed leaves on 1 1 1 1 1 31.24 2 as Most of the genera of this tribe were referred to the Geloniae by Pax & p Ee bui Li 2:28 0 1 © o 5 кн” ааа Pohl, e Bras. 1: 56. 1827; . 1979 m otype, uS Small, in Britton & Brown, Ill. Fl. N. U.S. s 2, 2: 462. 1913]. ê Пои Reliq. Houst. 6. 1781 (non see L TYPE: Jatro Pur I L. [= Cnidos pus Johnston]. ecchi . 1814 pos rej.). TY Biv ivonea !stimulosa (Mich. Raf. [= idis qus T Victorinia León, Me ; b. ‘Hist . Na t Felipe Poe 15:242. 194 1. TYPE: Fora regina e pac кл regina ix. — Cnidoscolus regina (León) ugh]. otropical genus of ca. 50 eed with the reatest concentration in Mex he as reviewed by McVaugh (1944), id his jm f Victorinia to synonymy appears justified. Tribe 35. ADENOCLINEAE TM Arg. W ebster, Taxon 24: 59 u 5; DC. Prodr. X 1139. 1866. ТҮРЕ: тазы кат Тигс onoecious or dioecious trees, shrubs, or herbs; latex clear (+ colored); indum minal, ax- illary, or opposite the guis ages e to oe or glomerular; flowe 3-5, imbri E dele or TTR disk a or ; ovary 2-6-locular; paceous; see езпу; endosperm Srt Roll g th t уез the six ae are grouped into two subtribes. KEY TO THE SUBTRIBES OF TRIBE ADENOCLINEAE la. Indumentum simple; stamens бер fruit dehis- cent or indehiscent, 2~3-locular ............................... EOW Ade noclininae lb. Indumentum stellate; stamens connate; fruit baccate, 1-7-locular 35. Endosperminae las hos ile piter Muell. Arg. 34: . Prodr. 15(2): A rus Paw TYPE: а. Turcz. Gelonieae subtribe Tetrorchidiinae Pax, Pflanzenr. 147. IV (Heft 52): 29. 1912. TYPE: Tetrorchidium Poepp. Trees, shrubs, or herbs; t йай: aes diverse; stamens ns 3- 2 , free; ud or 3-locular; stigmas distinct; | id E (oy or drupaceo | subtribe inclu des 5 wine with ca. 30 | species у the Neotropics and Afric KEY TO THE GENERA OF SUBTRIBE ADENOCLININAE la. eani ue present, at least as staminodia. | Жз 30; frui uits s drupaceous ovary | ith laminar В 211 Glycydendron | 2b. poe less than 20; fruits cl ova- ry 2-3-locular; leaves + dentate, with or without pei gla nds. 3a. es zt t 13 j or | itángls disk a of edes к | сеззез. 4a. Stamens 8-10; anthers 2 : 2-locular, caducous; indumentum c smooth.. a — . Stamens 3; а са гв 4-loc | peltate; stamina ate disk absent | eA c 3b. Herbs; seed-coat dry; staminate ae‏ سس pis Moria MB pene? "214. Adenodlit lb. Pistillate disk inii foggy Шашы pn without laminar glands _....................... | 211. Glycydendron Ducke, Arq. Jard. Bot E Janeiro 3: 199. 1922; 4: 107, pl. me : Volume 81, Number 1 1994 Webster Synopsis of Taxa of Euphorbiaceae i. 1925. TYPE: Glycydendron amazonicum Ducke. f Amazonian Brazil. Pax notypic genus of Nealcho the Pate (Ducke, 1925), m pno contains both g P: e niani ws Pierre ex Prain, Kew Bull. 105; d ae e 6(1): 963. 1913; TYPE: PLC E gaboniae Pierre ex Prain. А monotypic genus of West Africa (Nigeria to Gabon). 213. Tetrorchidium dece in dpi: & En- j . Gen. Sp. 1989, TYPE: Terarken A PETE Hernia JUS Adansonia I. 1: 843). Tetrorchidiopsis fine scher 559. 0 TYPE: Hasskarlia E Вай. [= Errore hidium didymostemon (Baillon) Pax & pa (асв; Мон iw = i А genus of са. 20 species, mostly in tropical Е: тегіса but са. 5 species і іп Attica. i nen ts dr Pa ax & ofan (1912), are бең ae ax & Hoffmann, 1931) reduced it to a section of Tetrorchidium. 214. Adenocline р, ws уна Imp. at. Moscou 16: 59. ; Mue , DC. dps ers s iet mes freut бес PI. 310 27: 404. 1913: Pax & "Hoff П (Heft 63): 409. 1914: Prain, Fl. Cap. 5(2): 855 Y BE: FE > В; 488. 1920; Milne- ome Kew Bull. 5: 349. 1951; Dyer, Gen. S. Afr. Fl. Pl. ed. 3 315. 1975. Mercurialis sect. Adenocline FUP) Baillon, Adansonia I. 3: 159. 1864. TYPE: not designated [lectotypification not de- sirable at this time due to confusion in Turcza- ninov’s application of names to specimens]. Diplostylis Sonder, Linnaea 23: 113. 1850. TYPE: not designated. ее Muell. Arg., DC. Prodr. 15(2): 1141. TYPE: Paradenocline procumbens Muell. Arg. [— riesen violifolia (Kunze) Prain]. reated by Prain (1920), Adenocline includes eight species of the Cape region of South Africa. 215. Ditta Grisebach, Mem. Amer. Acad. Arts. Sci. II. 8: 160. 1861; Muell. bc DC. Prodr. 15(2): Mas. 1866,4 Рах гіо Rico 2: 383. 1988. TYPE: Ditta а Griseb. us of one or two species of the Greater Айй (Cuba, Hispaniola, Puerto Rico). MAN wem ENDOSPERMINAE Pax & nn, Pflanzenr. 147. XIV (Heft 68): s "PM (comen) Natürl. Pflanzenfam. ed. 19c: 42. 1931 (in clavi). TYPE: vette бн Bentha DERE ee or aud pie eo stellate; lous; stamens 6-10, filaments coniate pis small or absent; ovary 2-6-locular; styles stig- етай confluent into а cap; fruit capsular. The — includes only the type genus En- dosperm 216. Endospermum Bentham, Fl. vq a: 861 (nom. cons.); Muell. Arg., DC. vein 15(2): 1131. 1866; Bentham, Gen. Pl. 1884; n Schaeffer, Blumea 19: 171. шы му Shaw, ull. Add. Ser. 8: 78. dospermum chinense Bon. n & Binn pd E Tijdscr. е! ӨГ ee Ned.-I ia hn . 29: 238. uccana а Teijsm. m (Teijsm. & Bin. ) Kur, A йин genus of ten species, occurring 102 Annals of th Мо س‎ Garden from China to Malaya, New Guinea, and tropical “ie ш чыр Мыз чыр. Webster, Australia emis nd). Tribe 36. GELONIEAE Mts Arg.) Pax, N türl. Pflanzenfam. ed. 1, 3(5): 88. 1890. Hip. pomaneae "open Gelonieae Muell. Arg., Lin- naea 3 dg Prodr. E 124, 866. TYPE: arse m Roxb. e illd. [= Suregada Roxb. ex Roul «di Шш (уегу теу шшш trees or 1 usually very scant or absent; leaves alternate, simple erved, mig usually pellucid- JR cH nine caducous. ow- ers мсн in glomer ded. Реза E usu- 5 or 6, imbri len grains 2- TERNS subglo! with C pistillod pals moana, Sa- 8. imbricate, eee s glandula ar roton- e 21 istillate ovary 3 (rarely 2- or 4- Nin e bifid fearely multifid). Fruit capsular ometimes indehiscent and drupaceous; seeds pee py it coat + fleshy; endosperm present А monogeneric poison tribe, as treated here. Pax & Ho Hs nn (1 931), in their final treat- ment, a lar tribe with 18 dits in 7 асаан а à considerable ене of 217. eoe Roxburgh ex Rottler, Ges. Na- 1951; на Вий. Јаг 443. 1958; Ai Cong e ы гор. uphorb. . TYPE: Sel ebi Gelonium и Sp. Pl. 4: 831. 1806 it pe tn., 1791). T v: Celonigim lanceolatum Bie ii lanceolata (Willd.) Croiz.]. оо Blume, Bijdr. Fl. Jav. Pe 1825. TY pte TA eene E [= cs erulata (Bl.) pts ашуре} C «орот Sonder, Linnaea pes Rt E us cana (бод) 17.]. 20. 1850. ТҮРЕ: Sond. 2 с afri- A paleotropical genus of ca. 40 species, the majority in tropical Asia. n 24: 5 1975. TYPE: Elateriosper- 1 тит a B ume oecious ا‎ stems with whitish latex; in in- a us. нер епсеѕ (ещ _ ог r oa cy- ate зер escent; stamens 5, free, imbricate; disk dd pub filam s ЧЕ 10-18, ents free; anthers Rae grains glo obose, MIU SON with Cro or absen p fleshy; seeds ecarun- culate endosperm scanty, gres cs massive. A monotypic tribe i ing only Elateriosper- mum. The genus was i. between Suregada and ospe by Mueller (18 e Ben- tham (1880) and Pax (1910) positioned it with Airy y Sha aw (1975) included it in his subtribe Ja- with Aleurites, Annesijoa, Ta- us appears to be is a distant relative of Micrandra and Man мы мүн ер) Blume, Bijdr. Fl. Jav. 18 C. rodr. 15(2): 1294. 88. 1880; Pax Pflanzenr. 147. I (Heft 42): 17. 1910; J- E Smith, M ept. Landb. 10: 571 1910; YPE: Elaterispt the other 5 220 IT mum tapos Bl. ome ae p rai, was transferre typic genus of SEE southeast Asi: ао to Java and Вог bes 34. ДАТИОРНКАБ E Pax, in М БУУ, Д н Й or care le 0. TYPE: Jatropha b pese (Muell. Arg.) Pax, Bot. Jahrb. "m 142.1 alypheae subtribe Joannesieae (‘Johann [^ Moll Arg., Linnaea 34: 201. 1865. ТҮКЕ: ей. nesia V tA. naQ ÀÀ — Volume 81, Number 1 1994 Webster Synopsis of Taxa of Euphorbiaceae Monoecious = commonly dioecious) trees, shru ems with articulated and/or nonartioulated ина latex clear, white, or red- distinct, аары реа. 5, bat or s Meer asi S CO- 6)8- 30, filaments at least partly connate; pollen grains gl , binucleate, inaperturate, se with roton-pattern; pistillode present or a Pistil- late sepal , free, imbricate, + p ent in ruit; disk annular, lobed, or dissected, sometime: with staminodia; vary mostly 2—3-locular; styles unlobed or bifid, Soutiorimre Заме: Fruit imine ular or dn unculat е, bee. bo ur As ci l tribe Jatropheae includes of the genera d A and Joannesia) (1910). None of the genera enu- included two genera, Givotia and n, n, that are now т alea ga he Ja tr ropheae, based on peni and Joannesia знах but Deutzianthus and Oligoceras are interme: diate in m many respects, and it now seems better not to recognize any subtribes within th Jatropheae KEY TO THE GENERA OF TRIBE JATROPHEAE la. eş simple (lobed or unlobed); monoecious 0i cious; E gera free or connate; АЖА usually reddish. - Styles bifid or cns sepals distinct; mon- oecious (rarely dioecious). a. Fruit c. liie idi not horned on ack. 4a. Seeds carunculate; leaves often lobed or dentate, without distinct ре рен аї is of petiole; not in- ack _ 1. Oligoceras 2b. Styles twice big t erect connate below; podes, 222. Deutzianthus lb. Leaves palmately compound; monoecious; seeds ecarunculate. 5a. ze pilates subentire to lacinate; ovary cular; anthers apiculate; foliage with malpighacen s hairs. ses minate calyx cu cupular, not 69 tokio bods stamen: s 7-10, і ong Aa РЕЯ "а ан dies 223. Joannesia 6b. Stam te calyx fused in bud, com- ll covering the petals; stamens 18-33, inner filaments connate; sta- minate disk annular E igh Pah oso fruit agri See не uwenbergia 5b. Styles slender cular; an- thers muticous; i ange saws s staminate caly. stamens 15- r fila- ments connate staminate dui Masts d; Irübc4psulr e с NUR 225. Annesijoa p ы L., Sp. Pl. 1006. 1753; Gen. d. 5, 437. 1754; Muell. Arg., DC. Prod 150) 1076. 1866; Bentham, Gen. РІ. 1880; Pax, Pflanzenr. 147. I (Heft oS 2j 1910; Hutchinson, d nU E pn 775. 1912; Fawc. & Rend., Fl. Jam H ürl. Mun. or 1979; Radcliffe-Smith, Fl. Trop. E. Ай phorb. 1: 343. epe enemi Pohl, РІ. as. Ic. D 12. TYPE: Jatropha Br. nios i L peat eg к, ор Fam. Рі. 2: 356. 1763. TYPE: Jatropha sL. iz & Pavón, Fl. Peruv. 139. 1794. TYPE: Coton 290 Ruiz & Pavón, пот. Шев. [= Ја ropha curcas L.]. C a escr. Pl. 5: 17. 1799. TYPE: Loureira glandulifera Cav. [= Jatropha cordata (Ort.) Muell. Arg} Mozinna отча N v. Rar. Pl. Hort. Bot. Mat seti YPE: Ил уйш Ort. [= М лк йо iion нее Engler & Pax, Natürl. Pflanzenfam. ed. dy 2 D 1891. ан арар schiedeana En ах [= Jatr dioica Sessé]. T e I da, FL ie 1: 177. 1929. TYPE: Collen a paradoxa a Chiov. [7 Jatropha para- doxa (Chiov .) Chiov.]. A diverse genus of ca. 175 species of tropics and subtropics i in mo: Old World and New Mp rld; 33 inde nds. "The assignment of species into n and sections is given by Dehgan & Webster (1979). 220. Vau "pne Me Bot. Mus fl. Har- vard Univ. 1955. TYPE: слад cataractarum Кыны 104 Annals rimi са Сагаеп кше genus of the Rio Negro region, kable Beme (pers. comm.) regards it as better placed n the Micrandre er e V mE Bull. Soc. Bot. ce 11: 872. гу ҮРЕ: еи Т. COE: otypic genus of Indochina; Du related to PAE but distinguished b onoe- cious flower production and curious сс вера. 222. Vee eae Gagnepain, Bull. Soc. Bot. 346. 1963. TYPE: инс, аме Gagnep Loerzingia Airy Shaw Kew Bull. fae 365. е 25: 54 43. r9 : Loer eutzianthus ov.]. r to the Indochinese ee шы m their differences seem of specific rather than generic di 223. Joannesia Muir Alogr. imer 199. 1798; Muell ; Prodr. Т Catar., FE str. EUFO 165. 1988. TYPE: рон ргіпсерѕ Vell Anda A. Juss., t. Euphorb. 3. 1824. : Anda em ü A. Vau nom. illeg. [— Pina, as Andic us Vellozo . Flum. 80, t. E: An- 1825. T s : pense ee Vell. [= удава phe Vell. А neotropical genus of two South American spe- ies. менеен», Letouzey & Halle, sonia IT. 14 . 1974. T YPE: e gia letestui SM Cu & Hall A tropical west African genus of two species in Cameroon, Congo Gabon. The enus is is in a many ways Libr modal E Joann E = m. б; nate flowers of the second species, Leen | africana Let. & Hallé, are still unknown. 25. Annesijoa Pax & H 147. ffmann, Pflanzenr. 1980. TYPE: Annesijoa BARS. Pax & Hoffm A monotypic genus endemic to New Guinea. — Tribe ue di ips (Pax) Hutchinson, Amer. | ibe Codi- PE: Codiaeum Rumph. ex A Monoecious or dioecious trees or shrubs; 5 | h nonarticulated laticifers, p clear “= nontoxic; in s E illary, racemose or ge ulate. Staminate pti ree or c imbri rica ate M Mar reg "n Ph free, p ; ra (5 ) se 100 e or Me nnate; кан grains binucleate, it, e sexine with Croton-patte pistillode Ф دم‎ я M s colume sually persistent; nculate or oe кы testa sometimes fleshy: ondotan present This чк of 15 genera with over 100 spec has the greatest generic diversity in the e ideae, аа нена nships between the genera are clear; it seems RU: to PS to rec formal subtribes at this | с=т=н M Volume 81, Number 1 1994 Webster 105 Synopsis of Taxa of Euphorbiaceae KEY TO THE GENERA OF TRIBE CODIAEAE la. Ate present, at least in staminate flowers. Petals present in pistillate flower: t e calyx lobed; fruit capsular. without beadlike glands beneath anthers and petals not gland-tipped; inflorescences al or axillary, not ЕЯ е leaves. 5 е terminal; seeds carunculate; staminate disk and ptacle glab 6a. Ovary 3-lo nid petals cni outside 226. Baloghia 6b. Ova id 2-locular; Киев атар -sericeous outside xillar ar 5b. Inflorescences a min 227. Hylandia receptacle pilose. 228. Ostodes 7a. Indumentum anes ach ifid inflo tice malpighiaceous; a. Styles ne. staminate disk i eds carunculate glabrous; stamens Ша, ri f Petals KR 9. Pa ini mdi staminate ast pubescent; stamens 14-16; petals es gare 230. jp NR adax ally; . Leaves with beadlike к beneath; nud and petals with apical glands; AE DR opposite кт; lea . Pantadenia ri truncate. ien accr 2b. road eer in €— ate Infi @ © c Ф Кя lowers in racemes; stipules caducous ог absent; anthers muticous. escent; petals glabrous; fruit capsular . ea cds s at accrescent; petals tomentose; fruit drupaceous . - 232. regi moj E . Fontainea odiaeum a. Styles entire, slender tyles twice iphyranthera 12b. St 5. 5 11b. Reaves in axillary clusters; stipules persistent, + spinose; anthers aia styles bifid = 236. r dilated Acidocroton 10b. Inflorescences ter Pistilla: te se Баг ас crescent, not recurved; floral disk present y stilla te sepals re 226. Baloghia Endlicher, dash his Nor 1833; Bentham s . Pl. 3: 300. 1880; ht Pflanzenr. 147. III (Heft 15 12. ДОТ Maiden, For а .W TOSE 28 1904; White, Proc y. Queensl. 53 226. 1942; Airy Sha > Kew Bull. 35: 1980; McPherson & Tirel, Fl. Nouv.-Caléd 14(1): 43. 1987; James & Harden, Fl. N. S. W. 1: 410, 1990. түре: Baloghia lucida ndl. Steigeria Muell. Arg., Linnaea 34: 215. 1865; DC. Prodr rus ): 1121. 1866. TYPE: Stei, igeria montana Muell. 8. [= Baloghia montana (Muell. Arg.) Pax]. Bree SEM s of 15 iiw 12in N Caledonia d the others in Norfolk Island, Lord Howe bu and Ais alia 227. Hylandia Ay Shaw, Kew Bull. 29: 329. 74; 35: 643, fig. 4. 1980. TYPE: Hylandia =н, Airy кы Pon 9typic genus of tropical Australia E icem i Shaw (1974) suggested a r tionship with Dimorphocalyx, Ostodes, isa ncul 13b. curved, not accrescent; floral disk : absent . lb. Petals absent; ess bifid; seeds eae eves glandular 240. Baliospermum Loerzingia (Deutzianthus), especially the latter. Th ese ни are gene erally pip e wit ghia Re prove to be the closest genus. 2% Ostodes Blume, Bijdr. Fl. Ned. Ind. 619. 35: 334. 1981. TYPE: Ostodes püntcilità ВІ. Дд ch (1966) Ostodes A is a genus of four species of tropical Asia, from A num- of species included by Pax & Hoffmann (1911) were removed by Airy Shaw (1966) to Fahren- heitia, a "арон that is accepted here pending further study. 229. Pausandra Radlkofer, ee 53: 92, t. 2. Sig 1870; Baillon, Adansonia 92. 1873; Muell. Arg., Fl. Brasil. nos 503, t. 99. 106 Annals of the Missouri Botanical Garden Pax & Hoffm., Pflanzenr. 147. HI jt 15 со, Bol. Mu ar. Emilio Goeldi, Bot. 3: 59. 1987; Huft, Ann. Missouri Bot. Gard. 75: 1115. 1989; Secco, Rev. Gen. Anomalocalyx . Amer. Sul 58. uh TYPE: Plus d morisoniana (Casar.) R As revised by Secco (1990), Pausandra is a h neotropical genus of ca. eight species, extendin, Honduras to Brazi enus is very dis tinctive and isolated from other gene he tribe inate disk, but very different in a number of other characters. 230. Dodecastigma Ducke, Notizbl. Bot. Gart. Berlin 11: эө 1932 Janeiro 6: E S Hm Sar TYPE: Dedena amazonicum Ducke of two species of the Brazilian Amazon and ri ipie 231. Pantadenia Gagnepain, Bull. Soc France 71: 873. 19 PE: Pantad. d ptg m Capuron, Adansonia II. 12: 206. pa E: Parapantadenia chauvetiae Leandri [= taden ia chauvetiae (Leandri) Webster, AE ov.]. A paleotropical genus of two widely disjunct e endemic to h puntata enia wee distinguished Sy me 2- шап 8: гу th 1- ; Kew Bull. 35: 624. 1980. TYPE: Di. pr Pc glabellus Thw 5 species, mainly in tropical Asia (India and Sr Lanka to Hainan and Borneo) but extending to Australia. 233. Fontainea Heckel, soe sur ig pancheri. 1870; Baillon, : 19% 1874; Pax, Pflanzenr. 147. ee па E 30. 1911; Airy Shaw, Kew Bull. 35: P Jessup & Guymer, Austrobaileya 19653 McPherson & Tirel, Fl. d E 14(1) 74. 1987. TYPE: Fontainea pancheri (Baillon) Heckel. Ап Augiralasian „Бере of six Ашу Ж extending uinea w Caledonia, and елй, 234. Codiaeum Rumphius ex A. Jussieu, Eu- phorb. Tent. 33. 1824 (nom. cons.); Muel. Arg., DC. Prod McPherson & "e 5 BET: А. Juss 1790. TYPE Phyllaurea Loureiro, Fl. Cochinch. ы: 4 Phyl . illeg., = Codi- Кайни urea codiaeum Lour. [no riegatum (L.) A p pa Synapisma "Endliche т, Gen. Р pi ra peltata ‘Labill. i anode gu . Gr Fl. Ind. Batav. 1(2): 412 n Corda, 1842). TYPE: A glabra = Codiaeum variegatum (L.) A. Juss.]. ca. 15 Ma P^ n Australasian genus of c -seeded Howeve the | species are otherwise so similar in the реши indumentum and inflorescence n" it seems best to regard them as congeneric. 232. Dimorphocalyx Thwaites, e Pl. Zey ‚ 186 ED Handb. Fl. Ceylon 4: 54, 1898; Pax, Pflanzenr. 147. III (Heft 47): 31. 1911; Airy Ke ull. 23: 123. 1969; 26: 2 1972; Whitmore, Tree Fl. Malaya 2: 86. 1973; Airy Shaw, Kew Bull. Add. Ser. 4: 9 95. s u narrower poi niti 235. Sphyranthera Hooker 18: t. 1702. 1887; Chakrabarty & Vasudev 6: Rao, J. Econ. Tax. Bot. 1984; 129. 1985.1 тш Урат aa ellai illeg. [m^ nthera lutes- cens a a os & Hoff endemic genus of the Tr T: n Beretta ago; there are two species, the & the recently described S. airyshawii Cha krab. | эче Volume 81, Number 1 1994 Webster Synopsis of Taxa of Euphorbiaceae of кан amans are still | un- Vasu p кар ү 1 Th lati hi 1 belong: to the ing. fees sie е Grisebach, Fl. Br. Ind. 9 (no s. prop.; dry hitch, ipa iis n DG. гу 15(2): 1042. 1866; Bentham, Gen. Pl. 3 d., , Symb. Ant. 9: 208. 1924; d. Bot 20А(15): 62. 1926; Alain, Fl. Cuba 3 1. Espanola 4: 69. 1986. TYPE: i atin pi tans Gri- seb. анса sae ад J. Wash. Acad. Sci. 14: 97. еза : Ophellantha spinosa Standl., sod '98 8 [2 yit эла аи spinosus (Standl.) Web. ster, comb. nov.]. mainly Caribbean genus of 12 species, 2 from Central America (Chiapas/Guatemala) and the re- mainder from the has been subject to both nomenclatural and tax onomic problems since its de Grisebach cited Acid. naeus represents Flueggea acidoton (L.) We (Allertonia 3: ЕЯ 2984); ptem Je Grisebach's desc тїрїї accepted by later authors as Acidocroton, by typ- ification his genus is a synonym of Flueggea. Con- servation of this genus that is common and wide- Spread in the Greater Antilles would seem to be the best solution d Nene Орла described from Central meri Ereater spinosity of the T in зн they TY Met similar in floral r the * accrescent ca the Central pin lyx in ime a It vci e: that Ophe E ellantha is reated Aci t. Ophellantha (Standl.) агае ov nly other s ies in the section trt the уре іѕ ege dnos tryermarkii (Stan bster, comb. phellantha steyermarkii ri Field Мо affinities of Acidic io. have been vari- ously interpreted. Mueller (1866) created for it a subtribe placed between Pogonophora and Cluti eat bat оса ne in ишә Micrandrinae still needs more study, the totality of floral characters support a placement in tribe Codiae 237. Blachia Baillon, Etude Сеп. gsm 385. 1 YPE: umbellata (Willd.) Baill. Toto bell Willd. J. us of ca. ten species, in то: Asia from жам ‘India to Hainan, the Philippines, and per- haps Borneo. The genus appears related to Aci- docroton E the New World, especially the section Ophellant 238. Pr puente Boerlage, Handl. Fl. Ned. 11(3): 15. 1989. TYPE: Strophioblachia fim- bricalyx Boerl. A genus of two species in southeast t Asia: In- bes. dochina to Hainan, the Philippines, and Cele 239. rat Baillon, Adansonia I. 1: 53. 1860 (no ns.); Muell. Arg., DC. Prodr. 15(2): ас 6; Fl. Bras. 11(2): 504. 1874; ntham 2218: . 1880; Pax, n- Be ‚ Ge zenr. 147. Ш е 47): 39. 1911; Jablonski, ard. 17: 151. 1967; Secco, Acta Amazonica 15(1-2, P 81. calyx 1985; Rev. Anomalo: Sul 99. е ТҮРЕ: онн гасетоѕа pa lon. According to Secco (1985), gon includes ; Levin f Amazonian South anama. Th enus is rather doubtful; it shows m to Ostodes but also to Croton ый se a. gener 108 Annals of the Missouri Botanical Garden 240. Baliospermum Blume, Bijdr. Fl. Ned. Ind. Ap zenr. 147. IV (Heft 52): 24. 1912; бейе пе- pain, Fl. Indochine 5: 429. 1926; Airy ote Kew Bull re 22219723336: 2674 ls Lus Tap Chi Sinh Hoc 11(3): 16. jen YPE: Rond axillare Bl. n Asian genus of ca. 12 species, distributed from India to China (Yunnan) and Java. Mueller (1866) associated Baliospermum with Суйп, hich it l bl in habit, and has been followed in this by Pax & Hoffmann (1912) and h workers. H section, ru. based on бечет balansae Сарп Tribe 40. TRIGONOSTEMONEAE Webster, Taxon 24: 599. 1975. ТҮРЕ: Trigonostemon Bl. ious trees or shrubs; indumentum sim- Monoec A ple; тее шшш шоро. eu oe veined, t apex of petiole; кй persistent « ог ОО. l. Infi nces terminal or axillary, ra- S sepals 5, free, im- ] zisal 1 exceed J › x or dissected; ашып: 3(- 5) fil t ther connective e rged, often ps ge М inaperturate е, with e grains globose pattern, 1 I Ws tillate 5, free, Шш еа disk cupular; ovary 3. түйү styles free or nearly so, unlobed or bifid. Fruit capsular; seeds écaruneulets: testa dry; en- dosperm present. е tribe contains the single paleotropical genus оаа, иод 241. рее. Blume, Bijdr. Fl. Ned. 825 (as fissi ); Fl. Javae ll 395. 7 IH Tee 47): de 1911; фен epain, FL In- dochine 5: 309. 1925; Quisumbing, Phil. J. Sci. 41: 329. 1930; Jablonski, Brittonia 15: ew Bull. 23: 126. 1981. TYPE: iode Rer serratus Bl. Er user n Malayan Misc. 2(7): 89. 1822 (nom. ej.). TYPE: Enchidium verticillatum Jack [= Tri койы» verticillatus (Jac k) P ax]. Silvaea Hook. & Arn., Bot. Beechey’ des 211. 1837. TYPE: Silvaea а semperflorens (R Roxb.) Hook. P [= Trigonostemon semperflorens (Roch) il Athroisma Griffith, Not. Pl. As. 4: 477. 4 (non A. PD andolle, 1833. TYPE: a берч ү Ара wo sp emn neither p Hoffm: Telogyne Bailon, "Eie Gén. Euphorb. 327. 1858. TYPE Telog Baill. [= жейин) vi latu M ЭР ах]. Tritaxis Baillon E tude Gén. Euphorb. 3 1858. TY Trit axis gaudichaudii Baillon [= duel ) Muel . Arg.]. ihe Tylosepalum Kurz ex . & Binn., Natuurk. Tijdschr. Ned.-Indië 27: 50. 1864. түре: Tylosepalum alr i inn. [= Trigonos re с. Bot. "Frane 71: 815. = Tri 80 Poilaniella Gag . Bot. i aoe niella la fragilis Cap gonostemon fags (Gagnep.) Airy Shaw]. Каа vhetes offm., Notizbl. Bot. e Berlin 10: 3: ү ТҮРЕ: Neotrigonostem Trigonostemon viri versifolius jk & Hoffm. [= ] ) Airy Shaw]. & n Balakrishnan, Bull. Bot. Sur v. India Ж TYPE: Kurziodendron were Balakr. [= Trigonostemon viridissimus s (Kurz) Air? Shaw]. sim Kurziodendro 68. 1966. rather diverse genus of tropical Asia, extend- ea. The Ostodinae еы Ж it, ein it is possible that the tribe Trigonostemonet ot be maintained as distinct from the Codiaea® || Volume 81, Number 1 1994 Webster Synopsis of Taxa of Euphorbiaceae Tribe 41. oe Muell. Arg., Bot. Zeit 4; DC. Prodr. 15(2): 199. onini анато Pax, Natür 1, 3(5): 113. =. — ci. Univ. Tokyo, 5 TYPE: ficticias 1954. Monoecious (rarely dioecious) shrubs or herbs; or absent; disk dissected or nt; sta-‏ ,5 اا mens н 15 ог тоге, flame onnate into a‏ olum rt and apparently free; anthers ex-‏ short a a io pollen Tias. aed inaperturate, sexine reduced); pistillode Eug Pistillate sepals (4)5. imbricate; petals 5, sometimes reduced or absent; disk dissected or absent; ovary 3-locu is styles unlo 3 bifid, or multifid. Fruit capsular, 1-3- d 1 1 + locula B . B ^ x 3 bryo cylindrical, with narrow cotyledons. As indicated by y pollen characters, tribe Ricin- rubias Pax ud Hoffmatin (1931). The tribe bees work may how that the two groups gren j^ combined. K EY TO THE SUBTRIBES OF TRIBE RICINOCARPEAE 1 е a. Pollen grains with lar rge clavate клн, ре. эй. petals and disk usually present; ovary E alternate ма Ricin AUS inocarpinae lb. Pollen gr. ith 4 preis та rer nt; ovary 1-3- locular; leaves ernate, кайыны or whorled 41b. Bertyinae Subtribe МА RICINOCARPINAE Webster, Тахо 1975. TYPE: Ricinocarpos Desf. x Trees Ка ees, shrubs, ог undershrubs; leaves alternate, metimes resinous; fl in te mor е ро у 3-locular; styles bifid or ; fruit сары ыр 3 seeds. An Australasian group of three genera, two en- demic to Australia KEY TO THE GENERA OF SUBTRIBE RICINOCARPINAE la. Styles bifid to multifid; stamens at least partially connate. 2a. Staminate calyx lobed halfway or М filaments all connate; petioles shor ig ie TT Be Gee er E 42. Ric inocarpos 2b. oan calyx lobed less than halfway r filaments free; petioles lon Ao RI UPC A ede Mr: 243. “Alphandia lb. ion unlobed, рны forming а cap over the ary; stamens all fre 244. Beyeria 242. т зедов Мет. аг © 47. 1826. TYPE: Roe peria pinta Шит аня [= Кин рое pinifolius Spr g]. fes Australian genus a ca. 2 spain pronped NES pec it does not oe r there; ше нр d Ne d x Palatio by McPherson & Tirel (1987). 243. Alphandia i dd Adansonia I. 1873: Pax & Hoffmann, Pflanzenr. B ге Fl. Nouv.-Caléd. 14(1): 86. 1987. TYPE: AL phandia furfuracea Baillon. A Melanesian genus of three set in New Кыа, New Caledonia, and Vanu a. Miquel, Ann. Sci. Nat. III. 1: 350, 4; Muell. Arg., DC. Prodr. 15(2): 6: 63. 1873; Bentham, Gen. Grüning, Pflanzenr. 147. (Heft 58): 63. 1913; m flanzenfam. ed. 2, Annals of the Missouri Botanical Garden Снори Klotzsch іп Lehm., Pl. Preiss 1845. т E: Calyptros stigma viscosum (laa ki [^ Beyeria үке (Labill.) Miq. Lo Beyer Mue ll. in я es бта d Bey inay brea Muell. Arg. = brev- ifolia (Mue g-) Benth.; ыо, эана һу Wheeler, es s А genus of ca. 15 speci d to Australia. incon md ese Muell. Arg., Lin- :1865; DC. ake 15(2): 208. oe TYPE: н Pla Trees r shrubs; leaves alternate, opposite, or verticillate; flowers in axillary Тасев br clusters or absent; st stamens numerous, filaments connate into a colum with sexinous processes ге duced; ova E 2- 3 loc- © ену s bifid to multifid; fruit 1—3-seeded. is Australasian subtribe includes one genus from Australia, two from New Caledonia, and one from Borneo. KEY TO THE GENERA OF SUBTRIBE BERTYINAE la. St ; petals present or ab- sent; ovary 3. loc ular 2a. Leaves alternate; anthers glabrous; fruit bord e aN 2b. ee es Menus eed Pit 3 dul 3a. Leaves е site ; inflorescences race- mose, terminal; styles m Бу fid PES tuin Ыр RESA _ Myricanthe ЗЬ. Leaves whorled; inloresene 1-flowered, axillary; "m -fi Ба ы 7. Cocconerion Ib. Staminate sepals absent; petals absent; ovar 2-locular; staminate flowers in terminal эре teate catkins, pistillate prei axillary; ant pubescent; styles bifid .. .. 248 В dori гоп 245. Bertya Planchon, Hook. Lond. J. Bot. 72. 1845; Muell. Arg., ~ f oon 19c: 226. 1931; Gu ymer, Austrobaileya 2: 427. 1988; James & Harden, Fl. N. S. W. 1990. ТҮРЕ: Bertya rosmarinifolia Ри [lectotype, chosen here]. A genus of ca. 20 species endemic to Australia. 246. Myricanthe Airy Shaw, Kew Bull. 35: 390. 1980; McPherson 2 zt 14(1): 72. 1987. T Airy Shaw el, Fl. v.-Calé i Мун ЙЫ, discolor onotypic genus restricted to northwestern = Caledonia san. Goteenérion Baillon, Adansonia 11: 87. 1. Syn. Nouv.-Caléd. 18 1948; Airy Shaw, Kew Bul. 25: t 1971; 32: 382. 1978; McPherson & Tire Caléd. 14(1): 38. 1987. TYPE: Cocco balansae Baillon. A genus of two species endemic to New Cale- оша. 248. Borneodendron Airy Shaw, Kew Bull 16: 359. 1963; Hook. Ic. Pl. 7(2): t. t. 3633 1967; Kew Bull. Add. Ser. 4: 60. 1975. TYPE: Borneodendron aenigmaticum Airy Shaw. Am c genus endemic to Born TEN RAND Airy Shaw (1963) E enus that it some: Shaw, 1975), he regarded it as most c with Cocconerion, a disposition that is followed ere. Tribe 42. CROTONEAE Dumort., Anal. a Pl. 45. 1829 (**Crotonieae"). TYPE: Croton inous Mii 4- The "eai t Leda tribe panded from my previous treatment (W 1975), in ы, only Croton апа uation, but at least has the merit of not | mma = Volume 81, Number 1 1994 Webster 111 Synopsis of Taxa of Euphorbiaceae roton in a taxonomic “black box" without any apparent уыш КЕҮ TO THE GENERA OF TRIBE CROTONEAE la. uc. not aene inflexed in bud; pistillate peta ium taminate petals thinner in dri aie xir pistillate flowers pet- ers styles bi tamens free; БИШ petals S 'inflotescerices axillary; seed coat d 249. Mildbraedia Stamens with fi s connate; sta- minate petals arenen n terminal; seed coat fles w p: p^. Fahrenheitia 2b. Stamens 3-6; staminate petals similar in MM ure to n pistillate flowers apetal- ous; styles unlobed or emargina e = - Moacroton lb. Filaments + distinctly inflexed in bud; pistillate | petals mostly reduced to absent ............. 252. Cro pau ц ia Рах, Bot. Jahrb 9; Pflanzenr. 147. III (Heft AT) = 1911; "pe di Fl. Trop. Afr. 6(1): 798. 1912; Léonard, Fl. Congo Belge 8(1): 85. 1962; Radcliffe-Smith, Fl. E. Trop. Afr., Euphorb. 340. 1987. TYPE: Mildbraedia paniculata Neojatropha Pax, Pflanzenr. 147. I (Heft 42): 114. 1910. TYPE: /Veojat et carpinifolia (Pax) Pax [= Mildbraedia ранда (Pax) Hutch.]. A tropical African genus of four species. 250. күн Reichb. f. & Zoll., Linnaea М bot. & Zoll. [= Fahrenheitia pendula. (Hassk.) aw]. А tropical Asian genus of four species, found em d i India and Sri Lanka to the Philip- es an Porn . Pa - uui. Dom по ax & Hoffmann MH ) : Ai i Shaw 5 66) slowed thet Fahrenheitia br te distinct, although he later (Airy Shaw 11979) eal to the same informal ‘‘subtribe”’ 5 2 Bh; Meee roton Croizat, J. Ап Arbor. 26: 9. 1945; Alain, Fl. Cuba 3: 85. 1953. TYPE: Moacroton leonis Croiz. As — by Croizat ем Her laisse i is . It appears deb related to Croton, and its status requires evaluation (Webster, 1992). 252. Croton L., Sp. Pl. 2: 1004. 1753; Gen. Pl., ed. 5, 436. PRO: Klotzsch, Hook. Lon- don J. Bot. 2: 48. 1843; Muell. Arg., DC. Prodr.15(2): 512. о. Fl. wet л 81. Hook. со = = = دو‎ o = 1943) in ш, 5: di: Sodio Ni dou 42: 793. 1993. TYPE: Croton aromaticus L. M saps by Webster, J. Arnold 1967; this de es the ear- 1 ү ape selection of Croton tiglium L. by Small, in Britton & Brown, N. U.S. ed. 2, 2: 254. 1913] Cieca та Fam, Pl. 2: 355. 1763 (nom. rej.). TYPE: Cro n LS us ЕІ. Bor.-Amer. 2: АМ 803. Lep- gro Raf., А Repos. П. 5: 353. 1808 (пот. superfl.). TY є: Crotonopsi linearis andis [= Croto Pa ii ster]. кони tas bec Kavi Gewächse, ed. 2, 2(2): : Friesia argentea Sprengel [= Cr d poste "Webster isa Raf., Neogenyton 1. т glandulosum (L.) Raf. [ 1825. ТҮРЕ: Decari- (C J lul 5 Drepadenium Raf., Neogenyton 2. 1825. TYPE: Dre " vm ium maritimum (Walt.) Raf. a Croton mar- ненай. Raf., Neagenyto n l. 1825. TYPE: — ды An Croton capitatus Michx totype Julocroton п Marius Flora ra 20(2 ) Beibl.: 119. 1837. TYPE: Ju мц phagedaenicus Mart. [= Croton tri- que qp i dure entium yi a Hartweg. 14. 1839. dac a trogyne crot noides Benth. Astraea ша, Te Nd 7:1 4.1841. TYPE: straea lobata Kl. [= Croton tats L.]. Cleodora Klotzsch, v ч. Naturgesch. 7: 1 841. E: Cleodora sellowiana Kl. [= С теда sphaero- e s Baill.]. Eutropia Klotzsch, Arch. Naturgesch. 7: 196. 1841. Annals of the Missouri Botanical Garden TYPE: Eutropia arasiliensis KL, nom. illeg. [= Cro- Medea Klotzsch, Arch. el di zB 41. TYPE: a hirta Klotzsch [= Croton КЕК ы (Didr.) йт Arg.]. Ocalia Kinase, Arch. i-a ead 7: 195. . TYPE: Cr : t ed ‚ Arch. Naturgesch. 7: 255. 1841. еме nophytum capitatum (Michx.) Kl. [= Cro- atus Michx. Podostachys Кыш, Arch. pelea ovi 7: 1841. Е: Podostachys subfloccosa Didr. [= ao len fü n- did (Didr -) Muell. Arg - var. subfloccosa (Didr.) Wheeler Muell. Arg. lectotype, designated by Ф, (1975 рерна Klotzsch, Arch. Naturgesch. 7: 197. 1841 E: Croton serratus Muell. Arg. Гаравы des- ted by Wheeler (1975)]. Horsham Ta urcz., Bull. Soc. Imp. Nat. Moscou 16: : [уке Mam s еы кач Tur z. [= Croton argent ja Е Lasiogyne Klos h, Nov. Act d. Caes. Leop.-Carol. j J1: 1843. ТҮРЕ: шуге brasiliensis KI. [= Croton compressus Lam.]. Tiglium Klotzs Act. Acad. Caes. Leop.-Carol Nat. Gir. it = p . U.S. Nat . Herb. ТҮРЕ: (Hook.) Benth. [= oton setigerus Hook. P Aaa Endlicher, Gen. PI. Up pl. 5: 91. TYPE i mC -ell itia Сај. seler [= оп m dh Barhamia Kich, in in Seem um Me y. Herald 104. 853. TYPE: Barham mia pan nsis Kl. [= Croton Meroe Wd lectotype, purse by Wheeler (1975)]. Cyclostigma iy: ch, in Seem., Bot. Voy. Herald 104. 1653. T РЕ: Cyclostigma | panamensis Kl. [= Cr ssp. panamensis (Kl.) Webster; designated by REI de (1975)]. hee Mane ‚ Re ep. Mar TY : С 1857.7 E: Cen- s Croton ү. Myriogonphos Didrichsen, Vidensk. Medd. Dansk Na- irh. For n. Kjoeben ha avn 1857: 142. 1857. ТҮРЕ ау Moore, Trans. Linn. Soc. 5. NE Heteroc Shire п тено 5р. Ул [= Croton mentiens (Sp. Moore) Pax]. d ell. / E Heteroc ате oo 461.189 A large and highly diverse but definitely mono- phyletic genus of at least 800 sp: ric ta mplexity of е genus is indicated by the 40 sections recent tly леке (Webster. 1993). The relationships. af: Croton within, the шщ h h R. been referred to a separate tribe associated only obea likely калыд ie E e since has in mens and seeds, but it diverges i in its axillary, eate po n suggest a pos- sible Teatri with the tribe Ricinocarpeae. Tribe hae De Ecc Ает (Pax) Hutch- in . Bot 969. Cluytieae n, Amer kind i rd ut Ew P zenfam. ed. 1, 3(5): 87. 1890. TYPE: Rici- ndron Muell. Ar loecious trees or shrubs; vam es leaves alternate, disc to lobed o ntire to lo n c. as) E c a a yis: ps styles bifid. Frui eous, die че eede d; seeds ecarunculate; E sperm copious This Soi litum tribe of two genera is entirely in African or Ma can except fo for one spe) he tr e uncertain. ^t cence; neverthe SS, ПІСТ ed t о Jatropheae KEY TO THE GENERA OF TRIBE RICINODENDREAE la. Leaves unlobed to 5-lobed; ete small ЮБИ Т ee a ee lb. naL palmately parted. jus s > flabellate- lobed, Аарне? p tendo € 253. p wib 2^ uote ROSE ret E К Stipules small, unlobed, d eid or а tt ا‎ 1-locular; тее Г уол N gl Rc ae oat Calcutta J. hus 2 P rg., nw 1866; ad Gen. Pl. И ү; 1880: $e M, — o — EL o S Ed Volume 81, Number 1 1994 Webster 113 Synopsis s bos of Euphorbiac Pax & Hoffm., Pflanzenr. 147. III (Heft iili 44. 1911; Radcliffe-Smith, Kew Bull. 22: 1968; Fl. Trop. E. Afr. Euphorb. 1 en 1987. TYPE: Givotia PUDE Griffith ex À genus of four species, one from Africa, two from Madagascar, and one from India. — etsi чар Muell. E ., Flora Fl. Congo 8(1): 116. 1962; i: Smith, Fl. E. Trop. Afr. Euphorb. 1: 1987. TYPE: oque EE ns Arg. Ag о African species, d related to Givotia and apparently differing only in vege- tative characte 255. Schinziophyton Hutchinson ex Radcliffe- Smith, Kew Bull. 45: 157. 1990. TYPE: shee ziophyton rautanenii (Schinz) Radel A monotypic African т discriminated by Hutchinson in an unpub i by Radeliffe-Smi th a ci ® a Р Б 5 T л [=] КА, © ee & t ruly compound leaves, small s arge fruits with du ا‎ peht Tribe 44. ALEURITIDEAE tre: ety Fac. Sci. Univ. Tokyo, x Y Bot. 954. TYPE: Aleurites G. For Mon ad us trees or shrubs; stems with nonar- тузду, hing ua scanty or absent; indu- m pos à Mone о stellate; leaves alternate, pin ae Sadia mately veined or lobed, eglandular or eso glands; stipules present or abse zd nces axillary, dichasial-pa- ate or red to cymose clu St calyx cl isse st ? Ра mg Fruit drupaceous or capsular and tar- hos: iscent; seeds ecarunculate, testa dry; en- rm Copious, Tribe Aleuritideae is circumscribed as previously ( ebster, TOT At kala & Ме: аА total of +e genera and ооо ресе. It is overwhelm- tribe as — constituted is still not sat- fin tham (1880) included vnd & ER a 931 s sirable to resurrect tribe Joannesieae (Pa 1924 in order to arrive at a more natural Eher KEY TO THE SUBTRIBES OF TRIBE ALEURITIDEAE la. Мопоесіош 2a. Penals 5, glabrous; stamens 8-20, fila- ents connate; inflorescence paula Pes palmately veined or lo arr а. Aleuritinae Pet из; ns ner filaments free: Sina uli a cymose clu ter; pon pinnately veined ..... 44b. ici lb. Dioeciou: 3a. Penals Душ at least in staminate flower. s entum simple or absent; bey lorescence ly t 2Ь. "Gre 4b. Indumentum ‘stellate or ерде; ре!- als often coherent ог соп inflo- rescence terminal or axillar os БЕ pret Cr ne 3b. Petals absent: indumentum — aves with beadlike glands ....... e. Neoboutoninae Subtribe 44a. ALEURITINAE (Hurusawa) Webster, Taxon 24: 599. 1975. TYPE: Aleu- rites J. R. & G. Forster. shrubs; indumentum simple or stellate leaf blades оон! veined or lobed, ith glands at apex рео e; пеене E 5, re glabrous; disk- „segments 5; stam Ku 20, laments connate; sen 2- or 3 Jocular fruit daas or са r; seeds ecarunculat The subtribe Aleuritinae is here equivalent to the tribe Aleuritideae of Hurusawa (1954), except into three genera, following bis ere is considerable doubt whether these segre- ates need to be recognized, as I indicated earlier Annals of th Missouri вни Garden bue 1967); however, they are peen cepted here, pending further critical s KEY TO THE GENERA OF SUBTRIBE ALEURITINAE la. Ovary 2-locular; fruit drupaceous; stamens 15- 20, anthers introrse; indumentum distinctly stellate 256. Aleurites lb. Ovary 3-5-locular; fruit dehiscent; stamens 7— 12 2a. Тл + Pallas ly pubescent, bracts c 257 ай, : indes) of bifid or simple hairs; an- thers introrse; pina not densely роь bracts inconspicuo: N c 8. Vernicia — pies J- & G. она ا‎ met: Arg., D TES Smith, Md . Trop. 1987. TYPE: Aleurites & с. гад T Aleurites moluccana (L.) ld. ]. the Pete ыр sense adopted by Airy Shaw e Aleurites is a genus of two species ex- tending from India to the Pacific islands. 251. Siegen Airy Shaw, Kew Bull. 20: 394. 19 E: Reutealis trisperma (Blanco) Airy notypic genus endemic to the Philippines; more or ‘ike Pueri between Aleurites (sensu stricto) and Vernic IT Baere is Loureiro, Fl. E 586. ae | t. 2801, рыр b: 157078721087: TYPE: “Кенге montan n Asian genus of three species rst from Burma to Indochina, China, and Japan; found in more temperate latitudes or ditti than Aleu- rites and Reutealis Subtribe 44b. GARCIINAE Muell. Arg., Lin- naea 34: 143. 1865; DC. Prodr. 15(2): 719. 1866; Pax & Hoffm., Pflanzenr. 1 € 68); 9. 1919. TYPE: Garcia Vahl in noecious trees or shrubs; indumentum sim styles bifid; fruit behavior: seeds ecarunculate. | Although Mueller (1866) iE three other genera (Aleu rites, Crotonogyne, and Manniop yton) in his subtribe Garciinae, Pas ч Но pise (191 3 ENNA them. In ihe treatment d Me ( 1975 and it is still accepted here. 259. Garcia s in вене € Naturh.-Selsk. jobenh. 2: ; Muell. Arg. 1. 1945; Webster, Ann 8. 1968. TYPE: pies nutans Vahl ex Rohr. I А neotropical genus of two species distributed from Mexico to Colombia ссе ets GROSSERINAE Me Tax . 1975. TYPE: Grossera Dioecious trees or shrubs; P ES or absent; leaves pinnately veined or tripline a glandular or eglandular; stipules decime o or | | zi onnate; p “ 5, free; ovary 3-5-locular; styles bifid; fruit sular seeds ecarunculate | s subtribe are n genera of thi дей | ee for Tapoides Ч | he Асап i їп Н tribution, Indone KEY TO THE GENERA OF SUBTRIBE GROSSERINAE . Sexinous processes of ШЕ rounded to obtuse 2 cien ve ine leaves pellucid- pun 3a. “Brats large, imbrica ie е before anthesis; forming a cone sr, Oo entire; eri scars УШАШ 1 kened a Cava Å 3b. Bracts small, persistent ы pe emu е; Зен scars very small; pistil- late sepals medially e ө. ome ost Volume 81, Number 1 1994 Webster Synopsis of Taxa of Euphorbiaceae 2b. Staminate petals pubescent; leaves not pel- lucid-punctate. 4a In TI axillary; ie ecarun- culate; stam TUN pe nular i EET 6 t 2. Tapoides 5b. Stamens over 20; ke gla- товба 2:3 263. An ا‎ 4b. Inflorescences terminal; seeds culate; staminate disk E peu mens over 20; capsule tomentose ... . Sandwithia . Sexinous processes of pollen echinate: leaves ES caedi staminate disk dissected. 6a. кане sepals 4 or 5, imbricate: stami- ptacle glabrous ............... . Tannodia 6b. Pilate Дн 2 or 3; Ig rece acle 266 DoDD 6 Сауасоа Léonard, Bull. Bot. Brux. 1955; quintasii (Pax & Hoffm.) тда [Grossera quintasii Pax & Hoffm.]. А genus of three species of central and east Africa 261. Grossera Pax, Bot. Jahrb. 33: 281. 1903; Hoffm., Pflanzenr. 147. VI (Heft 57): 958; F1. Солго, 8(1): 18 ins. a paniculata Pax ia totype, Fade by Léonard, 1955]. Ап African genus of seven species. 262. Tapoides Airy Shaw, Kew Bull. 14: 473. 1960; 20: 412. 1966; Hook. Ic. Pl. 37: t. 3632. 1967; Kew Bull. Add. Ser. 4: 200. 1975. ТҮРЕ: ) Airy Shaw [Ostodes villamili Merr. е A monotypic genus of Borneo. 263. Anomalocal ke, Notizbl. Bot. Gart. Berlin 11: 344. 1932; A hid Bot. Rio Janeiro 6: 60. Mes 3; Secco, Rev. Gen. Anom- alocalyx . er. Sul 39. 1990. TYPE: Ал- burm M (Pax) Ducke. ph > otypic genus of Amazonian Brazil. The у Stel position "d Anomalocalyx is саай. and it possibly belongs close to Dodecastigma and Sagotia. ге E Lanjouw, Kew Bull. 1932: 933; Secco, Bull. Mus. fond di Bot. 3: 157. TYPE: Sandwithia guyanensis Lanj. of two species of the Guyana region and Amazonian Brazil and Venezuela. Secco (1988) А Tal 8 an opinion that А withia is here provisionally retained Я p ибне Grosserinae because of its resemblance alocalyx, but it is quite possible that ab к genera may prove to belong to the same tribe о subtribe Tm Tannodia Baillon, Adansonia I. 1: 251. odr. 15(2): 728. m, & Hoffm., Pflanzenr. ur. VI (Heft o4): 110. 1912; Prain, J. Bot. 5 25. 1912; Léonard, 1955; FI. Bull. Jard. Bot. Brux. os Congo 8(1): 186. 1962; dovesse Smith, Fl. : al 1987. Tan- Te am cordifolia Vir e a cor- difolia Baill.] Hen dads b. 43: 220. p Tannodia sect. ix (Pax) Trois, J. Bot. 50: 127. 1912. Neo- ана aes 1982 (nom. Tax jf `5 : stia звя Рах [= Tannodia млн E (Рах) Prain; lectotyp A genus of three or four species in east Afr and the Comoro Islands. Although Radcliffe- Smith per rianth character, and Prain's reduction of it to a section of Tannodia appears justified. 66 ohinea Leandri, Bull. Soc. Bot. афт ‚81: 285. 1940. ТҮРЕ: роо рег- гїегї Leandri. A monotypic genus from Madagascar. In de- scribing it, Leandri noted its affinity to Grossera and ТЫЛЫШ. and this disposition appears correct. be 44d. CORET Web- ster, acis 24: 600. 1975. TYPE: Crotono- gyne Muell. ms metimes scandent; indumen- Trees or shru tum Min or dE leaves pinnately or pal- Subt Annals of the Missouri Botanical Garden mately veined, biglandular at juncture with Deng Be icate or Seni disk cupular; ovary 3-locular; styles multifid; fruit capsular; seeds e cuim Ап noa tribe of three genera: К» апа ibe Garciinae = е » (1866); however, they difer from Сап апд fewer petals that are more coheren The Crot er seem most tew decus to y dio Grosserin (^ KEv TO THE GENERA ОЕ S INAE la. Trees or shrubs; leaves pinay veined; peti- oles s lacking a ed tricho a. Styles bifid; кыыстан. rminal, panic- ы i staminate petals free; s aminte жел nee acular and extrastaminal, of mo than segments "261. Селде 2Ь. Sila: multifid: inflorescences a axillary, ra- cemose or spicate; staminate petals mostly rent or co inate disk ех trastaminal, of 5 d zs en rotonogyne Lia pa alm ately veined; maius with иймей trichomes; styles bifid; staminate bone als D VERIS 269. Manniophyton 267. ities gece dps Kew Bull. 1911: 231. 1911 : ойи argentea (Pax) Pat [C rotonogyne argentea Рах]. A monotypic genus of west Africa (Nigeria to Cameroon). 8 ара, да Muell. А; -, Flora 47: 535. : 720. 1866; Ben- "i m, x & Hoffm Pflanzenr. 147. VI (Heft 57) ^u 1912: ke Hook. Ic. Pl. 31: t. 3019 9. 1915; Léon- Bas Congo 8(1): 174. 1962; Adam, F i. mba 468, t. 165. 1971. TYPE: Croto- nogyne manniana Muell. Arg Neomanniophyton Pax & Hoffm., Pflanzenr. ,147. VI (Heft 57): 115. dn TYPE: Ne peditum (Prain) Pax [— Crotonogyne ls Prain; lectotype, chon here]. 15 species in west Africa (Sierra Leone to cado and Angola). 270. Neoboutonia Muell. Arg., J. Bot. pd уто Muell. "e e 4T: 1864; 3 Fl. det 8(1): 171. 1962; Adam, Меш Mus. Hist. Nat. Paris 20: 495, t. 183. 1971. TYPE: n africanum ac АЕ to a and E) f west Africa Leone эмы 44e. AOS E 24: e i: ser. d. u oniiforme: Hoffm., Pflanzenr. 147 VII (Heft 63): 71. 1914. Tribe Neoboutonieae .. Amer. 56 Neoboutonia Muell. Ar Tribe M MS Radcliffe-Smith, Kew Bull. 43: 632. 1988 E: Benoistia Perrier & Leandri. Dioecious trees or shrubs; indumentum stellate or Dacis leaves unlobed, p yo r palmately ined, glandular-dotted, stipulate; — _ Mini or axilla e segments 2 or 3; 15-40, filaments free, shorte thers with glandular connective; ovary 3 styles iia ae capsular; seeds carunculate 0 ecaruncu ry, racemose to рап! Pe 0 E ey X 15 А Pn of two ud one Re and one Madag . The era toj and Radcliffe. (1988) has prs to p т taxa, SO provisionally kept in the same subtribe KEY TO THE GENERA OF SUBTRIBE NEOBOUTONINAE la. Leaves m ———— —— M — —— — palmately veined; m. persistent; ота seeds carunculate 1. 0. Neobout ` 1Ь. meus. pinnately veined; ips decd P- | seeds ecarunculate 271. Ben 1864; DC. Prodr. 15(2): 892. 1860 ger Fl. Ibe ae кы Smith, ЕІ. Е. Trop. Afr. T3 lano нь gn Volume 81, Number 1 1994 Webster Synopsis of Taxa of Euphorbiaceae phorb. 1: 231. 1987. ТҮРЕ: Neoboutonia af- ricana Muell. Arg. A genus of three African species. 271. sa wage т» & Leandri, Bull. Bot. Franc 528 Kew Bull. js 632. 1988. Ty perrieri Leandri [lectotype, chosen her A Madagascan genus of three species. Subfamily V. EUPHORBIOIDEAE. TYPE: dips 1 bf usawa, J. Fac. Sci. Univ. Tokyo, 3, oy 6: 310. 1954. TYPE: iris Browne. Monoecious or less monly dioecious trees shrubs, or herbs an зари Heus non- sponte dates ually. whitish, sometimes scanty ene in Veiis leaves нра ог ант simple апа unlobed, entire о or dentate, usually pin- nately veined, often with glands at base of blade: ии arih sma ae da redi ced or absent ace- niculate, or Рен и (orbium хелай Pu EUN base; flowers apetalous. Sta- inate sepals 3-6, CAT e to valvate, mostly n mmonly reduced or Оргого; disk E соп- е to pati 3-colporate, educed or obsolete; disk absent; ovary 2-3(-20) fi om- y connate int mn. Fruit capsular (rarely rupaceous); seeds carunculate or eca t testa dry or fleshy ; endosperm copious circ рет of s subfamily Euphorbioi- five (1931) ( us » except that the tribe Stomatocalyceae (in- the Gelonieae by Pax & Hoffmann) is Key TO THE TRIBES OF SUBFAMILY EUPHORBIOIDEAE la. Р, olle : ere reticulate; inflorescences race- or ўме] bracts eglandular; stamens staminate coe 3-8- rice € ы ийй ог һап 5. STOMATOCALYCEAE lb. Pollen sexine perforate-tectate; inflorescenc ces n 3 E о دم‎ E 5 "a E 5 B a Е am aS gB с si ш Гы D or adnate to r е а; КЕРДЕ or herbs, not scandent 2a. Infi ences mostly se or spicat (if c , then not pseudanthial); styles А rae staminate calyx usually develo 3a. Fal bracts usually biglandular at ta ES = base, not pel r adnate to the rachis; stam or connate 4a. Staminat ор ud; leaves mostly not spinulose-den- tate; stamen: ee laments > n E 8. = [1 = ® о m = о w > = 5 ч = gm ч 2 < т л @ 9. .m 3 E Ex 5 3 E. E = nate to Фи rach ally with a single terminal и fonet zu 4 or 5 M ar aminate monochasia r dichasia; perianth reduced absent; sles mos bifid; seeds ge or culate Je. E 9. EUPHORBIEAE Tg., от Muell. Arg. t о йтгоп Hassk. ]. perui trees, shrubs, or kanaa; [у pue ish, alternate e, unlobed, pinnately veined, without glands mall and cad c 30, fi reticulate or reticu 5 or 6, free or pointe a ree, filaments. mon pem grains coarsely Pis at carunculate; endosperm ӨК ойу. This tribe includes one neotropical and three paleotropical genera фы ted into two subtribes. me characteris |р» within the ‘megan such as the liana habit, colored latex, and oily endosperm, suggest possible affinity with Ompha- lea, in the Acalyphoideae. Annals of th Missouri êê Garden KEY TO THE SUBTRIBES OF TRIBE STOMATOCALYCEAE 1а. Pollen grains finely re indehiscent; е Trevi. hee I RLS Кыйгы daba а lb. Pollen grains coarsely reticulate; Fen lar; styles + elongated; ovary 2-3-locular ...... 45b. Hamilcoinae culate- оила fruits 0-loc- omatocalycinae ca 452a. Sul АЕ Muell Arg. 1. a 34: 202. 1865; des Prodr. 15(2): 139: 1866. TYPE: Stomatocalyx Muell. Arg. [= Pimelodendron Hassk. ]. Trees or shrubs; leaves entire or dentate; ra- cemes simple or slightly branched; sepals Seid 10-20; pollen grains finely ocular; styles cime du кн tribe includes two genera, one of In- donesia, wes other of Afric КЕҮ TO THE GENERA OF SUBTRIBE STOMATOCALYCINAE la. Sepals 6-8, free; ovary l-locular _ 72. Plagiostyles lb. Sepals connate, staminate Miner 2-lipped; ovary 2-10-locular .. 273. Pimelodendron 272. Вон Pierre, Bull. Men: Pans 2: 1326. s. Soc. Linn 897; pe Kew Bul ull. 1912: ): 1001. 1913; S Ога Fl. : Plagiosty iuc PRASE (Muell СОО 8(1): 131. 1962. es Bu E. [= Pla- ) Prain]. typic genus of west REA (southern Nia to o Са and Congo). 273. Pimelodendron Hasskarl, Versl. bon; n. Ak ТАЕ k овес om д, , DC. Prodr. 15(2): пе. 1866. ҮРЕ: Sto А Агр. [= Pimelodendron griffithianum (Muell. уча Benth. x Hook. f.]. An Australasian genus of six to eight speci dictated f om Malaya to New Guinea, oben Australia ENE and ii Solomon Islands. 45b. Subtribe HAMILCOINAE Pax & Hoff mann, d anzenr. 147. VII (Heft 63): 419, 1914. TYPE: Hamilcoa Prain Trees or lianas; leaves long-petiolate, entire or 1 ranched; fr vaio pistillate sepals 4—6, 2- or 3-locular; styles connate e erect or bien. fruit capsular. f two genera, one American and one tively coarse-reticulate pollen grain riously similar in ornamentation to those of Dal- echampia KEY TO THE GENERA OF SUBTRIBE HAMILCOINAE la. Ovary 3-locular; styles erect; stam 18-20; pistillate sepals not glandular; stems E € t 4. Hamileoa 1G UN tamens b. Ova: 2-loc cul a^ pile e with 1 basal al es stem anden ITS. ТЕ leki * 107. р. Afr., ed. 2, 1 "t : Ham iiv onse (Pax) Pu ? кй zenkeri Рах]. A monotypic genus of west African rainforests (Cameroon). А ера Huber, В Bol. Mus. Goeldi 1: 3; Pax & Hoff d n try 68): SE d аце Au Ja t Bot. Rio de Henri «t Field Mus. Nat. Hist., A ЗША, 1): 1 1951. ТҮРЕ: Nealchornea yapurensis Huber. A monotypic genus of the Amazonian rainforest (eastern Colombia and Peru to Brazil). Tribe 46. HIPPOMANEAE A. Jusi pee Hist. Nat. Veg. 2: 522. 1834- eL. oecious (less commonly dioecious) tree a or nares ; latex usually milky, re ja acrid o «| dritic in фти leaves alternate e oppo le or ——— — س ج س Ооз‏ n nn ннан Volume 81, Number 1 1994 Webster Synopsis of Taxa of Euphorbiaceae simple and unlobed, esata veined (rarely pal- mo mately veined), com 1 ar or petiol glands; stipules conspicuous to reduced or absent Inflorescences terminal or axillary cemose or spicate е paticulote) usualy Autem Ws ndular. Staminate calyx open in bud (exposing WO. sometimes v кее we co M a colpi usu- R^ mergit, sexine тад tectate. Pistillate s 3-6, ate, sometimes reduced or ab- E - Fruit capsular or drupaceous; seeds carunculate or ecarunculate, testa tig or fleshy; endosperm copious, not ma rkedly This divided by , tribes «(А Page ЕТИ ФИТ diverse ре of ое "500 FEES was Pax & H su 7 — to maintain "oss С ic delimitati pi e uncertain and controversial, and may change ee when a monographic revision is com- eted. KEY TO THE SUBTRIBES OF TRIBE HIPPOMANEAE la. 6o receptacle minate onvex or conical; sta- pals 3-6, imbricate stamens mostly 070, P seeds са! ulate; inflorescences inal, paniculate; Tares sti gen V d s. pet iolar алав LE EASE y go . Mabeinae aminate Mor pue масаите. mak ostiy 2 or 3 һ2 c i ly metim only 1 o abse sent; ee 1-62 ). В ts fr r united; in- din ences ter idm or axil ery, rm cemose or picate ne хез not compound); eH often with Peut ands. "roa calyx laterally compressed; sta- mens 5-20 b. ree; leaves wit th petiolar glands stipule atex tery; Carumbiinae BUE 1 28 c. Hippomaninae $ "ee 46a. MABEINAE Pax & Hoffmann NEM 147. V (Heft 52): 22. 1912. TYPE: Mabea Aubl. Monoecious trees or shrubs; latex mil le 5; indumentum of den dri i : эм : *rnate, pinnately ve ы, "iun cn cem » PR in тезс ky, innoc- r ences termi- racts biglan- dular; staminate sepals 3—6, imbricate; stamens 2— 70, free from an into a column la р or connate ; pistillate sepals 3—6, sometimes gla ndular; ovary 3-locular, eve styles connate into a column, tips slender and unlobed; ul fruit capsular; seeds carunculate This neotropical videi includes three genera as conventionally defin КЕҮ ТО THE GENERA ОЕ SUBTRIBE MABEINAE la. Stylar column elongate; style tips slender, un- lobed; inflorescence гасыз + flexuous; stami- nate eve long-pedicellate; stamens (3- )10- 70; leaves and stems with dendrit dy" EE 276. Mabea lb. Stylar column ао e Jue. Men thick; ; сеу -12. 2a. Latex very scant y o камый ae mentum absent; stamens 5-12; leaves eee нее 2b. Latex copays milky; карис of dritic hairs present on inflorescence axes; stamens 2 or 3; leaves rigid .... 278. ‘Senefelderopsis 276. Mabea Aublet, Hist. Pl. Guyane Fr. 2: 867, t. 334. 1775; DC. Prodr. Arg. Ў fen 515. 1874; Bentham, Gen 3: 363. 1880; Pax & Hoffm., Pflanzenr. 147. V (Heft 52): 26. 1912; Jablonski, Mem. New York Bot. 17: 164. 1967; den Hollander & Berg, Proc. Kon Ned. Akad. Wet. 89C: 147. 1986; Huft, ы: tologia 62: 339. 1987; Missouri rd. 75: 1125. 1989. TYPE: Mabea piri үү Haiti s T by Pfeiffer, No- 2(1): 191. 1874]. A diverse and difficult genus of ca. 50 neotrop- ical specie p ee Martius, Flora С (Beibl. 2): uell. A DC. Missouri Bot 0 species, mainly in А f 8- Amazonian ‘South America but reaching north to Panama. 278. Senefelderopsis Steyermark, Bot. Leafl. Harvard Univ. 15: 45. 1951; рада 120 Annals of the Missouri Botanical Garden Mem. New York Bot. Сага. 12: 174. 1965; — Wartmannia Muell. Arg., Linnaea 34: 2 Gillespie, Brittonia 45: 92. 1993. TYPE: Se- Wartmannia stillingi ifolia (F. Mel b мый pe AS Орай еен [= Omalanthus Mia fa F. Muell.]. оре у RP Алый à. Index Nes t. Bot. Peu ү. i i E 866. hion peltatum Regel [= A genus of four species endemic to South Amer Г үт" зии (Linden) Villar] ica (tepui areas of Colombia, Venezuela, northern razil). — utm CARUMBIINAE Muell. enia ni a 34: 203. 1865; DC. Prodr. 15(2): 1142. 1866. TYPE: Санти Reinw. [= Omalanthus A. Jus Нерон. subtribe Homalanthinae Pax & Н PMN ‚ 147. ын 52): 42. 1912. TYPE: E x Jus Monoecious or dioecious trees or r shrubs; latex o unlobed to. bifa, usually glandular near tip; fruit dehiscent or indehiscent; seeds with large caruncle. As interpreted earlier ed e ж a vw iub ur is s ere um y the x& Hoffman o EU шаса нарты 6 in the sub ever, Pimelodendron differs in its pollen sexine ornamentation, eglandular wid and ecaruncu- late seeds. — edere A. Jussieu, eu Euphorb. Pl. 3: 33451880 offm., i Pill 147. V (Heft 52): 42. 1912; J. J. Smith, Nova Gui lanthus populneus (Geisel.) Pax]. Сахір Reinwardt, Syll. Pl. Nov. 2(1): 6. arumbium C" einw. [= О Fos a populneus (Geisel.) P. KEY TO THE GENERA OF SUBTRIBE HIPPOMANINAE la. mise 2-3-locular; fruits capsular or drupaceo 2a. Staminate spikes not capitular ps if so, Lm not foveolate). I Ап Old World genus of ca. 30 prona e | from Malaya through Indonesia to the Pacific | lands (as far as Tahiti and Ht ye ve | Subtribe 46c. HIPPOMANINAE Grisebach, Í Fl. Brit. W. Ind. 49. 1859; Pax, Natürl. Pflan- | zenfam. ed. 1, 3(5): 91. 1890. TYPE: Hip | pomane L. | —— subtribe Adenopeltinae Pax & Hoffm., . 147. V. (Heft 52): 57. 1912. TYPE: Ai. €: Мемед | Hippomaneae subude be Excoecariinae Pax & Hoffm, TEN enr. E. V (Heft 52): 153. 1912. ТҮРЕ | mnanthinae Pax & Ho! fim., Hippomaneae ме RE 2. TYPE: бут: Pflanzenr. 147. А 527: 57-191 nthes — subici Stillingiinae Pax & Hoffm., Pflan- | 47. X e t 52): 174. 1912. TYPE: Sulling | oecious or di shrubs, or wa latex milky, often caustic; indum entum simple absent; leaves alternate rt оро us innately veined, often with lamina pencil sands: Mirr ve small, e obsolete or absent; inflorescences terminal or 2* illary, m vara spicate (sometimes racemose), bracts usually biglandular; staminate calyx open in sepals 0—3; stamens 1-20 [=] subd at present, & vision seems premature Volume 81, Number 1 Webster 121 1994 Synopsis of Taxa of Euphorbiaceae 3a. Staminat il pi y bracteate; tami 1 ly 3-5; w oT . Seeds 5а. Spas tis rally carun иеЫ | culate : Stamina ae usually not ia seared bracteate; staminate sepals 2 or 3 (or obsolete). act 280. yen aves; staminate calyx 2-3-lobed; pistillate flowers ually e lea sessile or vorrei (iting pedicels less than cm long). e; pistillate flowers oken pedicellate ба. Columella not 3-horned at base; pistillate flowers often pedicellate ........ 281. Nossa 6b. Columella 3-horned at base; neos flowers sessi 282. Stillin 5b. Spikes axillary; staminate sepals 1 or 2, often баши or obsolete; pistillate flowers distinctly pedicella cm more in fruit) 283. Gymnanthes 4b. Seeds generally Seige testa eshy. Fruit sular (or if drupaceous eal bracts glandular). 8a ecious (rarely monoecious); petioles usually eglandular; inflorescences axillary; staminate sepals distinct; ovary not appendiculate; caruncle deciduous from seed and persistent lumella 4. Excoecaria 8b n n glandular; inflorescences terminal or axillary. ndular; M iie mostly w coat dry; calyx su| pre та petiolar glands. in ks is s not 3-hor . Stamens 2-4; pis pinnately veined, unlobed 285. Colliguaja b. lOb. Seed 9b. са eglan n 1; leaves mella 3. de rned; coat ү" һу; staminate calyx рг es palmately veined, + lobed ... ee Dalembertia leaves анын veined ... 287. ree 288. Sap . Spikes аш arent 3, filaments exserted beyond h gamophyllous calyx 13b. mr elongat styles 289. Glyphostylus inb Spikes terminal; filaments connate; leaves with glands at apex жга pikes axill 14b. $ uit drupaceous; inflorescences Tb. 2b. Staminate a ازا‎ нен EC bracts glandular; seeds foveolate, with. large crane 292. Duvigneaudia y; filamen xillary; bracts per ce Dux EE ERA 293. Maprounea 280. Spirostachys Sonder, Linnaea 23: 106. 1850; Pax & Hoffm ., Pflanzenr. 147. MR 52): 153. 1912; Prain, ЕІ. ч 1005. 1913 camb., Pu " A18: 1. 1964 че 29; Afr. Fl. Pl. ed. 3, 322. 1975; adele, . E. Trop. Afr., Euphorb. 1: 385. TYPE: Spirostachys р Sond An Ex Africa n genus of one species (or two, if Coecaria venenifera Pax is include d). 281. ia р diee Entdeck. 2: 118 KVA : Klein, Fl. Ilustr. eem Webster & Huft, ina M issouri ee 75: 1127. 1989. = ИЧ}, йа ЩЫ Spreng. ee Sprenge 1, ru Entdeck. Pflanzenk. 2: a 35: иѕѕопіа АЗУ, Spreng. [= Бера а discolor (Spreng.) Muell. Arg.]. __ 294. Hippomane % > 8 € а S 8 з 8 E ix] су > ELI. 3 Б = a A P 29 M uell. Tragiopsis ро Wochenschr. Саег tnerei Pane 2: 5. 1859. TYPE Eg brem Kars Sebastiani uell Dendrocousinsia Mills us. . Bot. 2: 374. 1913. TYPE: De ndrocousinsia spicata Millsp. [= Sebastiania spicata (Millsp.) Pax & Hoffm.]. A genus of ca. 100 species, mainly neotropical, } AE 1; : te North America. Asia. and Australasia. The herbaceous species of section 122 Annals of the ү шы Botanical Garden M; NF, f, З. ооду опеѕ апі тау poss ie represent : a фено genus. such as Se- ma aintain the — — saga unnatural) ge- x & Hoffmann (1931). — bees m Garden ex L., Syst. Nat. ed. 12, 767; Mant. Pl. 19, 126. 1767; FI. m 565. к ТҮРЕ: Stillingia = L So Muell. nnaea 32: E: Gymnost тоа ены Banh, ) Pe = Stillingia M ole (Benth.) Hemsl.; totype, designated by Wheeler, 1975]. 863. Mel ainly American genus of ca. 25 species, with a ee Piin taxa in the Mascarene Islands, eastern Malaysia, and Fiji. e pee Swartz, Prodr. Veg. Ind. Oe . 1788; Grise rit. W. Ind Pil. 3: 337. 1880; гач t. 309. 1895; Рах & Я и. V. (Heft 52): 81. 1912; Fawc. & Rend., Fl. Jam. 4: 329. 1920; Webster, J. Arnold Arbor. 48: 387. 1967; Taxon 32: 304. 1983; Ann. Mioun Bot. Gard. 75: 1129. 198 9. ТҮРЕ: Gymnanthes lucida Sw. [lectotype, chosen by Grisebach, 1859]. Actinostemon Martius ex Seige. Arch. Naturgesch. 7: 184. 1841; Pax flanzenr. 147, УШ 52): 57. Pues Jablonski, I Fhyologi TYPE: Acti ) jM (S eng.) [Gussonia piste Spren = Сунь d Il: agros rts КЕ, а Ьу і, Mem. New York Bot. Gard. 17: 177. Yu incorrectly cited Actinos- n grandifolius I KI., an invalid name]. Dactylostemon Klotzsch, Arch. Naturgesch. 7: 181. 1841. YPE: Dac ble stemon les mbur, gui en Si Lon- d J. Bot. 2 43 [7 Gym schom ман н SS xu nov.; a n. mscribed here following Webster dh бло is an American genus of са. 4 . A ost 20th-century workers h followed Pax & Нена (1912) in neni m деш аз e: ama. gie eg: spp to previous century, Baillor on (1858) К аш. оп oie gen- s р com суо with Sehen lem in this com iq of neotropical Hippomaninae with reduced staminate flow T name Ateramnus a species et Sapium (Webster, 1983 284. Excoecaria L., Syst. Nat. ed. 10, 1288. Ss & бави. 147. у deeds e e 1912; Cag: 26; Airy Shaw, nepa in, oe ah 26: 268. ue url Tree | Add. Ser. 4: 112. 1975; 8 980; Kev Bull. 35: 630. 1980; 36 1981; Rad cliffe-Smith, Fl. E. Tro fr Бор 1: . 1987. ТҮРЕ: Excoecaria agallocha al. chinchin. Her Commia Loureiro, Fl. Co 1790. THE ss [se Ee Com pis а Lou agalloc улуы ion Hurusawa, J. Tokyo Bot. 6: 311. 1954. TYPE: M | Ba Eie Ab gie lastellii (Baillon n) Muell. Arg: A paleotropical genus of ca. 40 species, тойу cluding н but this has not been 1o recent author: 285. си Moli y Sa gio Chile ! ; Cacia eec e ег 830; м | 3: 338. 1880; Ра У (Heft 52): 265. 1912; І. Smi aed mre EUFO 326. ا‎ TYPE: ligu orifera Mol A South American genus of five species. th et al fl Col T Dalembertia Baillon, Etude Ge horb. 545. 1858; Muell. Arg.» hee 1225. 1866; Bentham, pc. " Gen. Р. 3 | halfway e; petioles mostly 1 cm PA Or eh о a long c EL за Оуагу возе stylar column араш 3-lobed; stamen 1; staminate calyx rup- "s into 2 or 3 segments; ta КАСЫ ог 1еауез ENER v^ cuneat cma 298. Opht thalmoblapton 5-20-loc оси ular; stylar column api- caly dilated pele bed disk; stamens 10- 5 connate; staminate calyx cu- А oars terminal (or neg ^ flowers mer leaves ovate, unded to cordate a .. 299. Hura 3b. DU eure Baillon, Ann. Sci. Nat. 200 зто, 94. 1981 . TYPE: rne lean- genus of seven m mostly in » but also recorded from the A fi nian region as T comm.). ar west as ius (Huft, pers. A neotr tropical eastern Вг; razil 20) Alge €rnonia Baillon, Ann. Sci. Nat. IV. 9: aro Etude Gén. Euphorb. 546. 1858; Carb 5х du Safe 1230. 1866; Bentham, шр шы ; ; Pax & fla ЁЁ А. Es 52): 276. 1912; Pu 1981 q Nac. Rio de Janeiro 56: us ; Bradea 3: 148. 1981. TYPE: male 1. попіа Кайы, Bail A neotropical genus of five species, endemic to eastern Brazil. The genera Algernonia and Tetra- ndra appear very closely related, and it might 1.2 } h = $, |, к" genus. 006, SR E EA Fr. Allemao, PI. ds 849; Ann. Sci. Nat. Bot. III. 1 us. dep? L. Smith et al., Fl. Illustr. Catar. EUFO 325. 1988; ard, Fl. Lesser How: е ps TYPE: Ophthalmoblapton Fr. Alm. A ponia genus of four species endemic to eastern Вга 299. Hura L., Sp. Pl. 1008. 1753; Gen. PI. ed 5, 439. 1754; Muell. Arg., DC. Prod 1228. 1866; grow Gen. PI. 3 Pax & Hoffm POOR Arch. 821. 1982. TYPE: Hive cre- om epos Ecol ii "nr арен ie. type ipic polyandra B Baill ) d M ico to Costa ‘Rica; widely introduced n th tropics. bius 49. EUPHORBIEAE [Blume, Bijdr. 631. Natürl. Pflanzen- Midl. Nat. 30: “477. 1943. TYPE: Euphor- bia L. onoecious (or less commonly dioecious) trees, shrubs, or sh latex milky, innocuous or to oxic; indumentum simple or absent; leaves alt erate: opposite, or whorled, r veined, without era or райы glands; € seudanthial, 1; pollen -colporate, colpi marginat ше ре бешене. Pistillate flower a ded 126 Annals of the Missouri Botanical Garden -lobed or absent; ovary (2-)3-(4-)locular; styles 0 Baie dtu кч ha ecarunculate; testa dry, bolso or ornamented. tribe of more than 1500 species, here ii ЫР Webster, 1975) into ; subtribes e han 200 Dad names in tr ae T4 recently been published by Oudejans (1990). KEY TO THE SUBTRIBES OF TRIBE EUPHORBIEAE dens ate calyx present; involucre of 4 bracts, arti ally fused; cyathi al glands a p ial tube 49a Анален absent; involucral bracts ух ally 5, connate into а 2a. Petaloid a ppendagos bracteal (dilated in- volucral lobes) opposite the staminate 1 рна cyathial glande commissural; e, im- bat, түрүлө. staminate flowers; pistil- late sepals 5 or 6, large, ee те E ML uad irs oguillauminiinae . Petaloid appendages NEE (i.e posite cyathial glands), alternate with. M N c staminate monochasia small "e obsolete IE c M аЗ c. Euphorbiinae Subtribe 49a. ae ee уызы Webster, Taxon 24: 600. ostem- i ill nn. Sci. Nat. "e ^ pes 58 Anthostemeae Kl. & Gcke., Monatsber. Кбп igl. Preuss. Akad. Wiss. Berlin Kos 24 1859. TYPE: Anthostema A. Jus onoecious or е trees; leaves alternate, s inconspicuous; cyathia in axillary di ien ; cyathial involucre of dio. a isti minal (or apparently lateral); staminate dichasia + enclosed by 4 secondary (inner) bracts; cyathial glands projecting inward from infolded he ry + h + 2 A Л M B uter > м tillate flowers with gamophyllous calyx; ovary 3- Al 1 PERS Ч (ЧТ. Mus 1 eo E seeds carunculate or ecarunculate. This paleotropical subtribe includes two genera of Africa and ghe una The cyathia in thé An- thoct H 1 КЕ hat bilat t n at Diiaterall KEY TO THE GENERA OF SUBTRIBE ANTHOSTEMINAE Таг yathia bisexual; involucral bracts laterally con- nat pen 4-lobed involucre; cyathial glands at margins of internal bracts; inflores- cences axillary; ovary 3-locular. 300. Anthostema b. Cyathia mostly unisexual; outer involucral bracts | connate into a closed tube; cyathial glands fused by pairs into 4 lobes alternating with involucral lobes; inflorescences termi 4-locular . Dichostemma egal omen bane A. Jussieu, Euphorb. Tent. | 824; Baillon, Ann. Sc. Nat. 9: 193. POM Boissier, DC. Prodr. 15(2): 188. 1862; entham, Gen. 3: 261. 1880; Hutchinson, Fl. Trop. Afr. 6(1): 607. 1912; Pax & Hoffm., — atürl. Pflanzenfam. ed 19c: 207. 193]; | eay, . Trop. Afr. ed. 2, 1: 416. 1958; | Berhaut, Fl. Ill. Sénégal 3: 379. 1975. ТҮРЕ Anthostema senegalense A. Juss. two in A paleotropical genus of three species, west Africa and one in Madagascar 301. Dichostemma Pierre, Bull. Mens. Soc. | Y 59. 1896; Hutchinson. Keay, Fl. W. Trop. Afr. TYPE: Dichostemma puis Pierre. А genus of three species in tropical west Africa. | Subtribe 49b. Nr to vate seg a» p Sci. 64: 408. 1938; Webs : 1975. TYPE: Neoguillaum' oe ae | rees or shrubs; indumentum absent; E ternate or tpe entire; stipules — athia in axillary pedunculate cymes; !nY es bracts 4(5), diated or petaloid, connate pé glands 4, 8, o Lope атое “a the staminate чна ріѕ Ша! Mee | imbricate; ovary i locular; styles connate 2 bed, moo "1 unlo| dilated; fruit capsular; seeds 5 runc As defined by Webster (1975), this a | mande avy PME alasian genera. In their "m d cts, J Carre’ over to some exin in the Neo кыа е, suggesting that the pentamerous с i ig in the Euphorbiinae may be a derived conditi rom the involucral bra éyat thia of the Neoguillauminiinae are dis p different from those іп Euphorbia, with а ages developed from the interbracteal £ lands. Volume 81, Number 1 1994 ei Due of Taxa of Euphorbiaceae КЕҮ TO THE GENERA OF SUBTRIBE NEOGUILLAUMINIIAE la. Cyathial glands 8 or 10, €— Ate реи не al bracts large (са. 1 с пр); Іеа ral 302 Neo, bern lb. Cyathi hial glands 4, vem involucral bracts, Ji petaloid, less than 1 cm long; leaves oppos FOLE um aes nena 2, 303. у җон 302. Neoguillauminia Croizat, Phil. J. Sci. 64: 398. 1938; Bull. Jard. Bot. Buit. III. 17: 206. 1941; Guillaumin, Fl. Anal. Syn ouv.- Caléd. 18 McPherson & Tirel, Fl n -Caled. 14(1): 22, t. TYPE: : B Neoguillauminia cleopatra (Baill.)- Croiz. [Euphorbia cleopatra Baill.]. typic New Caledonian genus, мея dif- сан іп O from its Australian sister genu . Cal ем. ара Bull. Soc. Bot. кт d E Euphorb. t. 120. ^ aia 6: 52.1873; Gen ш “4 261. vue pia Bir Natürl. Pflanzenfam. des hà 19c: 221. 1931; Airy Shaw, Kew : 603. 1980. TYPE: Ca- HIM е Plane E of three species endemic to Australia; F: ma xeromorphic in vegetative morphology, ae have sah bel T those of the An- temina — 49c. EUPHORBIINAE. TYPE: Eu- Phorbia L, family Pelilanthaceae Kl. & Gcke., Monatsber. Preuss. ad. Wiss, Berlin 1859: 247, asa: 1859. Eu- Key | ТО THE GENERA OF SUBTRIBE EUPHORBIINAE ы: Cyathia (+) radial] сое р; utes e S Gcke.) yi a, J. Fac. Sci. . 3, Bot. 226. 1954 ҮРЕ: Pe iim a us A Euphorbieae ei иа КІ. & Gcke., Mon- atsber. P: ad. Wiss. Berlin 1859: 247. 1859. TYPE: Anisophyllum bon . [= Chamaesyce S. F Gray]. кышын subtribe Tithymalinae Kl. & Gcke., Mon- r. Preuss. Akad. Wiss. Berlin 1859: 247. 1859. TYPE: Tithymalus Scopoli [= Euphorbia L.]. onoecious s (rarely dioecious) trees, shrubs, or Neon uer ulate or exstipulate pac es € lor axi llary, bei cya NOM distinctly cupulate with x or 5 тиан glands on the rim (rarely glands only or 2); glands usually discrete, with or without pet in oe pistillate flower central, sta- flow ro a minate monochasia, monochasial bracts nee staminate calyx absent; pistillate cal —6-l , or often vestigial or absent; ovary entire); fruit capsular (rarely drupaceous); seeds carunculate or ecarunculate. The subtribe is here considered to include seven genera, pris h some authors have subdivided Euphorbi ia into many am ag There eere th y for fu Leeds such as the Pe dila xm ir odia of the taxa is reviewed by Wheeler (1943 y symmetrical, the glands not hidden within a nectar spur; styles mostly free or near- ly 50. 2а. involucra] а $ —) or 5, free, alternating with the lobes of the cyathium along the . Leaves alterna e. opposite, or whorled, but if opposite then not inaequilateral at P. vd veins of ed act clrenchya sheathed; stipules present or absent; main (embryo nic) axis not aborting; see tarunculate or ecarunculate SERES . 304. Eu phorbia 3b. Leaves Sopa stipulate, inaequilateral at base, veins pera: 'chlorenchyma s sheaths; = axis a . Chamaesyce seeds сша 2b. 1 ceasing growth above the cotyledons; "ей usua ally ecaruncul ula te ; "Ae wg t 4 or 5 alternating wit nate po a cup. v Hte illat * flower calyculate; involucral bids fused and the 2 eR pi^ sto a ie ORM late: Sterne DOT аыл Н ЖИК. UN EO CEL 4b. Реа fens N: : C RSEN тг i ж Ca succulent. " а. Cyathium hat bi l. the gland with a gap on the lower side ~- : somewhat bilaterally symmetrical, the gland wi gap doce oce SERE b. C ppsa without paka crests __ 308. ated below the crenulate rim; ovary ; with Permis double Synadenium crests at 4 ens . Endadenium 128 Annals of th Missouri س‎ Garden \ | 1Ь. = ир ани bilaterally symmetrical, the glands hidden within the nectar spur; styles connate into а jong сой 310. Pedilanths | 304. ee Ti SPB 450.1753; Gen. Galarhoeus helioscopius (L.) Haw. [= Euphorbia \ ed. 5, 208. 1754; Roeper, Enum. Eu- и i BIDS) би 5 позз | 3 edusea Haworth, n. uccu Е | ps Germ. 9. 1824; Boissier, DC. Prodr Medüsod major , Ait.) Haw. [= Euphorbia M | 15(2): 7. 1862; Icones Euphorbiarum, 1. L. var. major rit lectotype]. 1866; Muell. Arg., Fl. Bras. 11(2): 666. 1874; sr ber аж ped Succul. 131. 1812. TYPE: Te Bentham, Gen. Pl. 3: 258. 1880; Norton, а nez) Haw. [7 Euphorhi A m Ann. Rep. Missouri Bot. Gard. 11: 85. 1899; за , Edinb. New Phil | > h = | Pax, Bot. Jahrb. 34: 61. 1905: Berger, Suk- ҮРЕ: РЕ ЕШШ pirma (Willd.) Сга am [= | Euphorbia чеди ку a Wil | kulente Euphorbien 1. 1907; М. E. Brown, ми Rafinesque, Fl. Tell. 2: E La | Fl. Trop. Afr. 6(1): 470. 1911; Fl. Capensis nthis splen ndens (Boj.) Raf. i “Euphorbia ple 5(2): 222. 1925; Pax & Hoffm., Natürl. de bia e «b | Pflanzenfam. ed. 2, 19c: 208. 1931; Prok der Rafinesque, FL Tell, O ME h C Ё : d orbia Л; L. Mo ae ype anov, Consp. Syst. Tithymal. Asiae Mediae. E ud tuU num Schlecht., Linnaea 19: 252. 1933; Croizat, Bull. Jard. Bot. Buitenzorg III. Alectoroctonum uo Schlecht. [7 pn 16: 351. 1940; Hurusawa, J. Jap. Bot. 16: scotana Schlecht. 330. 1940; White, A., ў er & B. L Anthacantha ее Ш.Н ort. 4: Misc. ae 1857. TYPE ; Euphorbia heptagona a [lectotype]. à Sloane, Succul Euphorbieae. 1941; ler, À denoperalum | Klo tzsch & Garcke, Monatsber- Konig. Amer. Midl. Nat. 30: 478. 1943; Leandri, Preuss. Akad. Wiss. Berlin 1859: 250. 1859 (no Not. Syst. Paris 12: 64. 1946; Vindt, Trav. Turcz., ru TYPE: Euphorbia graminea Јас, Inst. Sci. Chérifien 6: 23. 1953; Ursch & [lectotype]. 1 Pre Leandri, Mém. Inst. Sci dag. 5B: 109. Bu ser = ORI 1050 нам | 1954; Dyer, Bull. Jard. Bot. Etat 27: 487. Akad Wiss. Berlin 1859: 251 calli (ШКЕ | А 1848). ТҮРЕ: Arthrothamnus tiruc li (L pn Dressler, Ann. Missouri Ж Gard. 48: Gcke. [= Euphorbia tirucalli L.]. Konig. 1961; been Webbia 20: 573. 1965; Борыны наср & Garcke, Monatsber Es gs ү d Arbor. Pa 395. 1967; Жү ss. Berlin 1859: 249. A au Radcliffe. d & T. ‚ Fl. Europaea 2: 213. Dill margen (ШЕ К | 2 -— hori ia mar, 1968; Allem & Irgang, Fl. Ilustr. Rio Grande f, phorbiastrum Klo ae h & qu Monatsber. Ee do Sul 11: 15. 1975; M. C. Johnston, Wright- Preuss. Akad. Yes Berlin 1859: 252. 1859. ie 2 ia 5: 120. 1975; Leach, Dinteria 12: 1. 1976; Euphorbiastrum hoffmannianum КІ. E Hassall, Austral. J. Bot. 25: 430. 1977; Su. Bs hoffmanniana (Kl. & Секе) Boisi bils, Kurtziana 10: 83. 1977; Carter, Hook ? , тез бу the Klot 5 & Garcke, Monatsbe Ic. Pl. 39: 5. 1982; Huft, Ann. Missouri Bot. ue сыл 24 1 Wi Beg omg та 252; 1859. m | Gard. 71: 1021. 1984; McPherson & Тї horbia diee ns Во}. [= Euphorbia . Nouv.-Caléd. 14(1) 10. 1987; Gilbert, Desm.; lectotype]. г. König Kew Bull. 42: 231. 1987; Carter, Kew Bull. Тиһу malop qM Pan reke, Мо АШ TYPE ‘USS. а 1 Tin 42: 673. 1 1. Е Trop. Afr., Е phorb Pires corollata L. [lectotype] Kónig- 2: 409. 1988; Deil & Müller-Hohenstein, Eu- Шырын у tzsch & Garcke, Мола nm phorbia J. 5: 1 988; Lebrun & Stork, Pre . Wiss. Berli u 48. 1859. Enum. Pl. Afr. Trop. 1: 213. 1991; Mayfield, Euphorbia fulgens Ms оз, YPC) aet. Lyciopsis р sier) Sch a a 7 (non Spach, 1835 ). TY TYPE: Eu a rum L. | (lectotype, ре by Mill augh, Publ. Оле Stapf, in Johnston, Liberia 2: 6 646. 1 34 Field Columbian Mus., Bot. 11 y 1909]. TYPE: Elaeophorbia drupacea (Sc chum.) ЗР Euphorbia ‘drupa ea um.]. 921} шуш Gaertner, Fruct. nom. cons. Diplocyathium Н chmidt, Beih. Bot. Centr (Ка) E: Tithymalus pow s ^ tow (7 Eupho 1907 „ТҮРЕ: Diplocyathium capitulatum | bia eplus L.]. Sch midt [= Euphorbia Ре е БДУ —€— (Ait.) Haw. [= Euphorbia an- — Euphorbiodendron Millspa iag” iodendtt f , ,acantha Ait; lectotype]. Mus. Bot. 2: 305. 1909. TYPE: Eupho ^ Esula (Persoon) Hay mid Syn. Pl. Succul. 153. 1812 latazii pd Min. Es n: latazit it gt (non ү andi, 1761). TYPE: Esula dalec champii Haw. lectot chosen h = Eu uphor phorbia esula ifolia [= № Саш Haworth, Syn. РІ. Succul. 143. 1812. TYPE: — Ctenadena r Consp. Syst. Tithymal. Mt | Volume 81, Number 1 994 Webster 129 Synopsis of Taxa of Euphorbiaceae 8. TYPE: Ctenadena lanata (Sieb.) h. [= Euphorbia lanata Sieb.]. mum Prokhanov, Consp. Syst. Tithymal. Asiae Mediae 25. 1933. TYPE: Cystidospermum cheiro- ida s (Fisch. & Mey.) Prokh. [= Euphorbia chei- rolepis Fisch. & Mey.]. C Rm lap e Mesa Syst. Tithymal. Asiae E . 1933. : Sclerocy: EE радо vii ОГА, se [Чу sc ус, Когоу Po] с А vast genus of over 1000 species, subdivided ny subgener mber of ОЛО cited here (lectotypifi- units by Wheeler, a frac- tion of the published names that are referable to y Wheeler (1943), Vindt (1960), Re осу. Carter & Radcliffe-Smith ) — No] eo re ere is great need for a ыл. ду be ИН gu guide to the зори, Еир jans (1990). Some segregates, ай 0 Poinset- жени һауе been ado opted by a a nu umber of workers, Е f future Conservative generic circumscription adopted у Boissier and his followers. 305. i EER F. Gray, Nat. Arrang. Brit. Pl. 2: 260. & ina. = tnik, S. Afr. J. Bot. 3: 262. са ; Allertonia E 331. 1987. TYPE: amaesyce maritima S. F. Gray [= Cha- maesyce peplis (L. ) Prokh. ]. "орут Haworth, Syn. Pl. Succ. 159. 1812 (non , 1763). TYPE: oru d peplis E ) Haw. ie Chamaesyce peplis (L.) Prokh.]. А . тч uk Eum genus of ca. 250 species, m in tropical America and Afric ca; often ti asa Ра t Ei Кезшде the iost de uphorbia. Koutnik (1987) tailed analysis of the char- PE g t f Chamaesyce. However, it t be admitted that Eu pho. rbia sub maesyce and Euphorbia subg. т та. Ву pris шшс ы, ко conservative ера of Euphorbia, because its recognition makes the An 10 cladistic gee off me hook, tailed phyloge кл ‚ we practical alternative to con- ing the use of the present artificial (but rea- PAM usable) system. 306. arrow es gut teat ii eet Mus. Nat. Hi 913; Alai ifolius (сть) Millsp. EEE linear- s ius Griseb.]. of three species ipiis to i West bes (Cuba and | Hispaniola чн г to spe- cies of Euphorb . Adenorima, hail perhaps о be included in ibit group "e SIN Pax, Bot. Jahrb. 19: 126. 903. 1987; Fl. E. Trop. Afr., Eupho 540. 1988. TYPE: Monadenium coccineum Pax hrb. 30: 343. 1901. Экей Рах, [= PAPAE hare ei 19, Stenadenium spinescens Pax nescens (Pax) Bally]. An African genus of ca. 50 species, distributed from Somalia to the Transvaal. 308. Synadenium Boissier, DC. Prodr. 15(2): 187. 187. 1862; N. E. Brown, Fl. Trop. Afr. ee ERA arborescens Boissier [7 Syna- denium cupulare (Boiss s.) L. C. Wheeler]. 130 Annals of the Missouri Botanical Garden A genus of 10—15 species of eastern and south- ern Africa, closely related to Monadeniu atte? d Leach, Garcia de Ort TYPE poe enium weht diee E d BT А monotypic genus of Meum PoE not dis- tinct Eds the genus Monade MR raras X cker Poi Ann. Mus 1812; Boissier, DC uphorbia J. 985. Pedilanthus pede à ) Pad. Am : zi E T5 P E w Tithymalus | Miller, Gard. Dict. Abr. Ed. 4. 1754. abl. Bot. 9. "Us Tithy- i н ej.). Crepidaria Ha awo rth n. PI. S Hi E: Tithymalus myriflns Mil] о нй tithymaloides (L.) Р. М bun A Сеп, Г! 86. 1802 Gee S qp 1798 at. [= ninis b (1 ae — Klotzsch & Gar iss Мовай ber. Kónigl. s. Akad. Wiss. Berlin m 254. 1859. TYPE: Ba pavonis Kl. & Gcke. [= Pedilanthus \ Н, Л, } ЗЕРЕ KGnigl. Preuss. kad. Wiss. ыз 1859: 253. 1859. ТҮРЕ: Нех- d macrocarpa eren Kl. & Gcke. [= Pe- th.]. dilanthus macrocarpus Ве А neotropical genus of ca. 15 species, all except ^. д M. х 1 z: C PES рете 2 іса. INCERTAE SEDIS 311. т Pax & Hoffmann, nze T ME m 125. 1912; EE C дый e ed 2167 193] TYPE: Chlamydojatropha podes d Pax & Hoffm This mysterious plant, known only from рер ns fi x the staminate flowers are described, it cannot be definitely placed. ee CN Beille, Compt. Rend. Hebd. в Acad. Sci. (Paris) 145: 1294. 1907; Bul us . France 55 (Mém. 2) 8: 1508 Pax pa Чакы, Natürl. Ёш сщш 19c: 52. 1931; Léonard, Bull. Jard. m t. tie Belg. 59: 319. 1989. TYPE: Mar- tretia quadricornis Вее. n west African genus of two wes was placed next to Aporusa by Pax & Hoffm an (1931), but á “Léonard Scien уор that it Be б ina tribe Mar e Köhler ex Léo say Phyllanthodeae Meeuse (1990) vega: ea lara (шоло е) ^ 313. Tacarcuna Huft, Ann. Missouri Bot. Gard. 76: 1080. 1989. TYPE: Tacarcuna gentryi Huft. This recently described genus of three species from Panama and South America was proposed by the author without any иша affinity, al- аи the description о les of the ary, together with the g gest arm aged Acalyphoideae. eneral m would sug- wever, exami- an isotype of 8 type species JE has (Gen : 6560. А, ше thas me over) a that Tacarcuna belongs | to subfamily Ph yllant ae and лү s to E beu: em it y be close à БАЙ ), scanning micrographs of gue fena of Joan Non icke; Webster, ol) show a striate exine suggestive of pollen in other tribes such as Antidesmeae or Briedelieae. 314. پو‎ A. Philippi, Linnaea 33: pr 1864. Е: Avellanita bustillosii R. A- lippi in patente. position of this genus has rê to the present. In his orig: ша) е Philippi assigned Ач ош to the n the basé lowing these earlier dispositions, located within tribe Jatrophea e following С us flowers and e be mw! an plant wo quite discordant with that Volume 81, Number 1 1994 Webster 131 Synopsis of Taxa of Euphorbiaceae Examination of scanning micrographs fr specimen in the Museo Nacional (SGO 051523) clearly contradicts an ass RÎ of emr to subfamil a iena -pattern. The 3. colporate grains with rath- Crotono coarse muri to nanospinules ar e- Из t similar HANS, in tribe Adelieae subfa Acalyphoideae, by tt in tribe oo vem ‚ Trewia. Although the precise affinity of sinite remains ambiguous, the evidence from pollen and floral characters clearly Io to its placement within subfamily Acalyphoidea a A ponis Alain, Candollea 17: 116. E: Cubacroton maestrense Alain. not been ues f thi aliti genus, which is known САН from the type allen in d Sierra de Maestra, Cuba. Judging from the rae ne the plant probably = ong s in vire Crotonea appears similar to the Cuba oacroton, and possibly may prove to ЕИ ап aberrant species of that genus. ADDENDUM 316. er hao Esser, Novon 3: 245. 1993. TYPE: Dendrothrix yutajensis (Jabl.) ats This 5 genus, referred b Мањ s author to subtribe abemae, wa $ published too Ay to be accom- ated in the u of this synopsis. It ps provides several keys to riae uen Es the other genera of Mabe 317. y pem wi sri Bull. Soc France 85: : 1939 РЕ: E eei ah vto 4 Te onotypic genus from Madagascar remains ii "Ed Judging from the description and Eig it does ear to be similar to the oes app а Spread paleotropical genus Su mesg so it У Prove to belong to tribe Gelonie EXCLUDED TAXA l. Burse ranthe Ri izzini, denm A/5: 5. 1974. РЕ: comes бы "m ا‎ genus, based on a single species any in the family (except for the dubious case of Bischofia). Examination of authentic material a RR < = 5 5 Ё @ = c a т < 25 N D. B м n = © = ى‎ e = ЕЧ m ct = owing th of Pennington (1981), clearly nonoa Tri- chilia lepidota Mart. subsp. lepidot 1980; 2. E dU Phytologia 47: 175. 92. TYPE: Casabitoa Fl. Espa 1986. perfae A This inadequately characterized genus, based on pistillate specimens of a single species from His- paniola, is not euphorbiaceous. According to Tom e noni ii (in press), I Sd the Pn] Ili, th the iyoneura (Urb. ) Urb.). 3. priae Ducke, Notizbl. Bot. Gar 1 1932. ТҮРЕ: ыйы amazonicus Andi The genus is not d although its systematic position has controversial. Al- bero Pu uni S ше (1039) немча Poly- ). pla ce- in the Anisophylleaceae by Pires and ens (1971) probably has the most support (e.g., in Cronquist, 1981). 4. Pseudocroton Muell. Arg., Flora 55: А MAE Huft, каш Missouri Bot. Сага 9. 1982. T e tinctorius et pies (L.) Fawc. & Rend. ]. 5. pape Pax & Hoffmann, J. Wash. Acad. Sci 06. 995 HIE Felina me 40: 2 12. 1977. T & Hoffm. [= Trophis mexicana vien Bur. ]. c ие Baillon, Adansonia 11: ; Bauman-Bodenheim, Bull. us тоң мы. gine II. 25: 420. 1953; Steenis, Adan- sonia II. 11: 615. 1971. TYPE: not ўт? ated (Baillon published 2 species: ee رو‎ bal- ansae and 7. codon andra; but either р man-Bodenheim NES The N ryne was re- duced Pr v van Steenis (1971) to dpt ah sub- sect. Bipartita LITERATURE CITED Notes on Malaysian Eu- AIRY S H. K. facies Kew Boll 14: 353-397. 132 Annals of the Missouri Botanical Garden پھچ‎ A synopsis of the genus Ptychopyxis la гайр amazonienne. Ш. Arq. Jard. Bot. Rio Ја. gai Ma Kew [n 14: 363-374. neiro 4: 1-208. Dia, рар of Dew families, new names, SUDAN C. 1952. Pollen Morphology and Plant Tax- etc. "iei Willis’s “Dictionary.” Be a a v bi ells, Uppsa Kew Bull. 18: m Farr, ый, т^ J. A. LEUSSINK & F. A A. STAFLEU ie Wilis Fe of the Flowering уте Index Nominum Genericorum (Plantarum Plants m Pons. ey ind [pp. xxi, xxii]. Cambridge 3 vols. Bohn, ie ыр & Holkema, Utrecht. Uni ire s, Cambrid| Forman, L. L. 1966. The reinstatement of Galearia 961. Notes Malaysian xen 2 Asiatic Zoll. et Mor. and Microd k the P. кама Kew Bull = Acai tien = S Salle e те Haee ee of Siam. synopsis of Galearia Zoll. and Mor. Kew Bull. 26: (Pand vds за Ви ii 26: 153-1 1974. Noteworthy у An from trop- САСМЕРАІМ, F. & L. BEILLE. 1926. Euphorbiacées. Е. ical dius (Bu c. Pl. 38(1): Generale AU ва 5: 313-516; is 3701- ar a EHRMANN. arbeiten zu einer Monographie The Биргана of Borneo. Kew der nae Be ide lia mit besonderer Ваа Bull. m Ser 4: z gung der e Arten. Bot. Jahrb. 41(Bei Ар ad synopsis of the Euphorbi- : 1-42. хеее. Platylobeze of Australia foc Phan Сис, E. 190 Пасбипирие Africanae. In: A. Engler thus, Eupho 39: (editor), Beiträge zur Flora von Afriken 32. Bot. E Jahrb. 40: 444 51 в. 81. The Euphorbiaceae of Sumatra. Kew mun L. synopsis of neotropical Pluke- ES. 36. um 374. p (Euphorbiaceae) including two new species. ез. Syst. BAEHNI, C. & P. AU. 1939. Polygonanthus, E re de зорнен: Bull. Soc. Bot. France 86: байан. k un it iphorbiaceae- Porantheroideae & 83-186. icarpoideae. In: A. Eng A- ditor), Das Pflan es H. 1858. Etude Générale du Groupe des E zenreich IV. 147 (Heft 58) 1 | phorbia Mx ve Masson, Par 973. акаю, conspectus of the 865. Remarques Я ba les Stillingi- Euphorbiaceae Taxon 22: 591-636. idées rs Brésil. Adansonia, I. 5: 336-344 HENDERSON, F Studies in Euphorbia ا‎ БОЛ Es Кв ox Eüphoricie: Bé ACTI ns. lat revision of Amp rea Ай. Adansonia, I. 6: Juss. Vip sie Бас r cat Muell. BALDWIN, J. & R. an diei 1947. А conspectus of ). Aus ts st. Bot. 10 (in press). the Без Сипигіа. Bot. Mus. Leafl. Harvard Univ. Hooker, J. D. ie of British "India, Vol. 5. L. 12: 3 51. Reeve, pee BENTHAM, С. 1878. Mess on Борооснаочав, J. Linn. Hurr, M. 1989. Nis and critical taxa of Кирһо Soc. London, Bot. 185-2 fro xs th America. Ann. Missouri Bot. Gard 7 0. dd aceae. ©: ГС. Bentham & J. 1077-1086. D. Hooker, Genera Plantarum 3: 239- 340. Seit > 1954. Eine nochmalige Durchsicht des mE o = К. & R. N. КАРП. Bischofia he et 3mmlichen Systems de Euphorbiaceen im wer javanica —Its enia with Vidi dip Phy- n Sinne. J. Fac. Sci. Univ. Tokyo, Sect. 3, Во. tomorphuley 23: 264-267. 6: a 342. BORHIDI, A. 2. La taxonomia del género Platygyne HUTCHINSON, J. -1912. Euphorbiaceae [in p Me A Hist. Nat. Mus g. 64: 89-94 In: W. T. Taisen -Dyer Де tor), Flora of Trop Carter, S A. ADCLIFFE-SMITH 1988. Euphorbi Africa, Vol. sy 41-1059. aceae Part 2. Fl. Trop. E. Africa. Mes HUTCHINSON, J. 1969. “Tribals in on family nC CHAPMAN, А. 91. Australian Plant Name Index. biaceae. Amer Australian Cove nma Publishing Service, СОЙ INGRAM, J. 1967. А reridend st ay tes ra. e et nete сене Croizat, L. The Tribe Plukenetiinae ig the Herb. lo Euphorbiaceae і in eastern tropical Asia. J. Arnold he generic limits of Argytha % Arbor. 22: 417-431. Е) defined. Gentes Herb. 11: 426- 1 Notes on the Euphorbiaceae П. 1. JABLONSKI, E. Euphorbiaceae. In: B. VII. The systematic position т ү, кы еен . Bull. Jard. (editor), "Sos of Gua am e es nd — Part Bot. Buitenzorg. ser. 3, Mem. New York Bot. Gard. med- 2. On visis. Eup orbiaceae from the Jussieu, А. 1824. Басен gener tropical Far East. J. Arnold Arbor. 29-5. icisque earumdem tent. . Didot, Paris. | C тый уле п Integrated анко of бы: KÖHLER, E. 1965. Die Pollenmorphologie de fication of Flowering Plants Columbia Univ. Press, laten Euphorbiaceae und ihre utung fü a cw оп Grana Palynol. 6: 26-120 the DEHGAN, В. & С. L. Wess Morphology and — KourNik, D. 1987 taxonomic re (pr fragen relationships E ud ke. Sees d ios Hawaiian species of the genus Сһатаеѕусё phorbiaceae). U Lo lifornia eis bl. Bot. phorbiaceae) crees : 331-388. сот DRESSLER, R. Eos pe raat (Eu. Ккорт, В. C. & 1989. Proponi prow phorbiacea). Воо 45-61. serve 4483 S d beds 1760 against P. Ducke, А. 1925. Panis ou peu connues de 1756 (Euphorbiaceae). Taxon 38: 320- -325. Volume 81, Number 1 1994 Webst Syno ses of Taxa of Euphorbiaceae 133 pera J. 1958. Euphorbiacées. Fl. Madagascar 111: eae J.-P Enumeération des Plantes a d^ linh d Afrique Tropicale, Vol. I. Général- ités et Annonaceae à Pan . Conservatoire et i i néve, Geneva. Jar LÊONARD, J. 195 spêces congolaises pilaster: ou . Be intéressantes. Bull. Soc. Roy. Bot. Belg. 84: 47-60. ==, 1956. = ulae Systematicae DE. ins sur les genres Oldfieldia, P horbiaceae су Bull. Jard. Bot. Brux. “26: 343. 1959 servations sur les genres Pycnocoma et E Pu (ШОШ Айрис). Bull. Soc. Roy. Bot. Belg 1962a "Euphesliancae! Fl. Congo 8(1): 1- . 1962b. а — 33: Sur е lim- ites entre les rypetes Vahl et Li Рах (Euphorbiaceae) Bull. Tard. Bot. Brux. 32: 513- 989. Revision du genre africain Martretia Beille (Eu; s orig et la nouvelle tribu des Mar- tretieae. Bull. Jard. Bot. Nat. Belg. 59: 319-332. az & ANGO. Hymenocardiaceae. Flore d frique Centrale: Serin erm atophytes Jardin Bo- É tanique National de Belgique, Bruxelles EVIN, С. А. 1986a. Systematic foliar dons of Meta or Car athe sine I. Cons; Teer. | I986b ctus. Ann Й е утаж ЖОООК of Phyl- sen ( уте 4 II. Phenetic analysis. 1 ї. Саг . 1986c. чаш tok hol oe l- ме III. Cladistic anal ar mo p B Бы | 1 5 E MPSON. 1994. Phylogenetic maps cts ultrastructure i in the Oldfie ie idea: биос), Ann. Missouri Bot. Gard. ian d 18 A Natural System of Botany, 2nd o Rees, Orme, Brown, Green, and Long- Novitates antillanae. Mem. New 107-157. 1987. Euphorbiacées. Fl. 226. и N v. Lal ting R. 1944. The e genus Cnidoscolus: Generic Dur andi оне groups. Bull. Torrey Bot. Club MEEU ст 4 1990. The Euphorbiaceae auct. plur.: аы п Unnatural Taxon. Eburon, Delft. ча ss s . W. ood a of the " iaceae, in particular of ш» subfamily Phyl- Mer toe e. Bot. a Linn. Soc. 26. LL, E. D. Ac а оп nas Д. е анн. " Trans. Amer. Philos. Soc. MILLER «с. L. Wessr 1962. Systematic po- o Cnidoscolus а ees Jatropha. Brittonia 14: $4 5. Euphorbiaceae. Linnaea 34: 1- Te. 1866. 15(2 189. Pr maneno In: DC., Prodromus ——. 1874 rene [part 2]. /n: Martius, Fl. Db L2y.2 OupEJANs, К. С. H. M. e d Catalogue of Spec "i iis н uphorbia In: Engler Prantl Die Natiirlichen emet A ed. 1, ds Euphorbiaceae-Jatropheae. In: gler (elor), Das Pflanzenreich IV. 147. d A 42): 1 Pee Die Phylogenie der Euphorbiaceae. Bot. Jahrb. 59: 12 e &K. пине Euphorbiaceae-Cluy- tieae. /n: sein (elitr Das Pflanzenreich IV. 147. Ш Eres 47): 1- 1912 Euphorb iaceae—Gelonieae, Hippomaneae. In: A Engler — m Pflanzen- reich IV. 147. V. (eft ven дый чен Acaly- pheae-Merc urialinae. ers Ene (editor), Das РВ еш ОН! Iv. Vai VII gc 63): 1-473. wie ی‎ sta Plukene s Pflanzenreich IV. пае. In: А. Еда (editor), D 147. > (Heft md Euphorbiaceae- Phyllan- Pe In: А. A (edion) Das Pflanzenreich IV. 147. XV. (Heft — & horbiaceae. /n: A. E gler & К. ranil [ Die Natiirlichen Phan. zenfamilien un 2, 19c PENNINGTON, T. D. 1981. tropical Meliaceae. Fl. Neotrop. 28: 1- 410. 1874. —À не Т. Fischer, PFEIFFER, L. Cassell. Pires, J. M. & W. A. RODRIGUES. 1971. Notas sobre os géner os Polygonanthus e Anisophyllea. Acta riesco 1(2 = RAIN, D. 1 Euphorbia ceae [in part]. / 99 sip Dyer (editor), Flora Capensis, Vol. ‘5, sect. Par T. eere ollen morphology of the — aceae with special reference to taxonomy . Wen 7: 1-116. RADCLIFFE-SMITH, An account of the genu Ce, n aoa à ariy (Euphorbiaceae). Kew Bu i 28: 2 eee Euphorbiaceae, Part 1. Fl. Trop. E. Afr. 1-407 1987b. S gre Масев. Bot. J. ver 1988. ilies from the Euphor- Not ioe scan n Euphorbiaceae 1. On the identity of Priel and the affinities of ns MX oxylopsis (Euphorbiaceae). Kew c ud is Roc e 1973. Manihot, Ma- айоо aper eae). Fl. ERA 13: em T8 d € ita Nov. 53: 1-37. 1952 Studies i in the genus Micrandra ‘The — of taxonomic c studies in ————. 1970. _ Э Вог. Веу. 3 6. 1990. A e = view of the genus Hevea. Malaysian perg at x & Development Board pec ograph 14: 985. Notas ie 9 novo conceito de Sa- cta ee biete ela. "Bol, Mus. Par. Emilio Rew o dos géneros Anomalocalyx uck iig а Ducke, Pausa ndra Radlk., Pogoni ophora “Mi iers ex Ben teriais tei a ora amazonica IX. Елый ше. a sistematica de o Richeria. Bol. Mus. Para. Emilio Goeldi, Bot. 41-158. Sree, = С.С.].у к Caledonii Adansonia " bn bird -624. М днн A. L. 1980 f flowering plants (Mag li ). Bot. Rev. 46: 225- THIN, N. N. 984. Tribus Alchornieae (Euphorbiaceae) of ر‎ Flora. Tap Chi Sinh Hoc 6(3): 26- —. Tribe Epiprineae (Muell. Arg.) Huru- sawa (Euphorbiaceae) in Vietnam. Tap Chi Sinh Hoc 10(2 к ribus Codiaeae (Pax) Hutch. in Vi- etnam. Tes Chi Sinh Hoc 11(3): 14-17. Linie D. W. - Conceveiba Aubl emen ae) new to Africa. Ann. Missouri Bot. Gar 856 6-858. Vinpt, J. 19 Monographie des Pho M du Maroc. зен a ique. Trav. Inst. Sci. Chér if., Bot. 19: 4 WEBSTER, Gals efe monographic study of the West Indian oe of Pylian har J. Arnold Arbor. 37: 91-122, 217-268 ior: А п оаа of the West Indian species of ОШ n prm Arbor. 38: 51- n 170-198, 295- 958. Ат ae study of the West Pss ا‎ мер thus. J. Arnold Arbor. 39: 49-100, 11 1967. The genera of Euphorbiaceae in the southeastern United States. J. Arnold Arbor. 48: 03 roo nspectus of a new classification of the Fuporicza Taxon 24: 593-60 ———. A botanical emen knot: The case of m and G pl ). Tax laceae). n 32: 304-305 1987. The f the spurges: A review of classification and relational in the Euphorbiales. Bot. J. Linn. Soc. 94: 3-4 46. ——. 1992. Realignment A Ci (Eu _ < Novon 2: 269-2 273. A аа ибо synopsis of the sections of die genus Croton a Gar serio Taxon 42: 3-823. 793 W. BRUSTER. 1991. A synopsis of Ре ана конагы: of pyar Aam ig aceae). Bot. J. Linn. Soc. eg. wv. 1968. deo -— of Panamanian Euphorbiaceae. Ann. Missouri Bot. Gard. 75: 1087-1144. Annals of the Missouri Botanical Garden WHEELER, L. C. 1943. The genera of heel Euphor- bieae. Amer. Midl. Naturalist 30: 456 -503. Miroi 24: 534-5 APPENDIX 1. Outline of taxa as treated in the text. E сои hae ings be WIEL $ ser ia 2. Savia 3. Gonatogyn 4. Petalodiscus E otia Actephila Discocarpus Lachnostylis . Chonocentrum 10. Wielandi 2. Tribe AMANOEAE 11. Pentabrachion 12. Amanoa 3. Tribe retro a Cleistanthus . Brie س‎ $ 4. Tribe eem 4a. Subtribe е 15. Astrocasia 4b. Subtribe LEPTOPINAE 16. Leptopus 4с. ctr PSEUDOLACHNOSTYLIDI- y T; Chascotheca 21. Pse затон Keayodendron 4d. sisi: SECURINECINAE (Sie ANDRACHNINAE Andrachne 4f. see FLUEGGEINAE 5 єл 33. - Tribe DRYPETEAE 34. Lingelsheimia 35. utranjiva 6. Tribe ANTIDESME EAE 6a. Sui s ibe SPONDIANTHINAE ndianthus эзин ire gage 39. Subtribe | SCEPINAE 40. Protomegabaria esobotrya 6b. 6c. Pee genera lectotypified. а У Volume 81, Number 1 1994 Webster Synopsis of Taxa of Euphorbiaceae 45. Apodiscus 46. Ashtonia | 47. Aporusa | 6d. Subtribe ANTIDESMINAE | 48 acoris 53. Hyeronima : Yo ea E ERAN a @ = донн 7. Tribe HYMENOCARDIESE 6. Didymocistus Hymenocardia 8. Tribe PISCHOTTEAE ischofi Bis Incertae Sedis itn Phyllanthoideae) Centroplacus 50. Meborea Il. ну К УККУС И Дайы Tribe CROIZA b roiz 10. > POD ALY CEA AE А, Hu ibe TERES оаа Родос 10b. эмне TETRACOCCINAE us 10c. Site PARADRYPETINAE Par И zw et TRAE a. Su peti БАЕ ONGTANAR nche llb. SAN DISSILARIINAE 71. Longetia lle. Fut PETALOSTIGMATINAE Petalostigma па. Sie ر‎ NDE нө 12. I e PICROD DEND EAE "ur e. PICRODENDRINAE 12b. му PA IRUSINÁE Oldfieldia 12c. Subtribe MISCHODONTINAE Aristogei eitonia . Misc 86. Voa дн 87. Ап Ker ЗА 1 88. Stachyandra Il. Subfamily AC 13. Tribe C AE p 14. Tribe POGONOPHOREAE 15. Tribe CHAETOCARPEAE Tri - Chaetocarpus 16. Tribe PEREA um 17. Tribe C EILOSEAE Cheilosa : -Nececbttedliiá 18. Tribe E SMANTHEAE . Erismanthus . Moultonianthus 19. Tribe DICOELIEAE Dicoelia 20. Tribe € LEARIEAE 1 23. Tribe CHROZOPHOREAE 23 ra SPERANSKIINAE Б otc 9. 23b. S iie DITAXINAE 5. Doryxylon эб че n= ЕБ “hag, > ا ا‎ c E E. $ © 3 В. 3 Melanolepis 25. Tribe 26. Tribe PYCNOCOMEAE 26a. Subtribe ee ee . Pycnoco: 29. Droerloncia 30. ине BLUMEODENDRINAE ventes ع تت تن بن د ا ع دب ب . . را ا . ч‏ tri‏ oe Stier‏ Е 2‏ 5 E‏ > © = 4 27. Tribe EPIPRINEAE 27a. Subtribe EPIPRININAE 5 6. Symphyllia 136 Annals of the Missouri Botanical Garden 2 -1 c . Tribe . Tribe A 904: دب © = 30). зок. 137. Ade nochlaena эз Cephaloerotonopsis Cephaloc Si CEPHALOMAPEINAE 43. Cephalomap ADELA AE 144. Adelia 145. Cr rotonogynopsis ЕЕЕ 5, > disi @ mog orze E z t — z > ti VE ubtribe CONCEVEIBINAE 55. Conceveiba 56. Gavarretia 7. Polyandra ALYPHEAE a. — DISTR us wal. "ADRIANINAE Adriana 59. . Subtribe аша Ашейин. 60. Mercurialis 61. Seidelia 62. Leidesia . Subtribe atc pai 8 ubtribe CLEIDIINAE o = @ Hd S 66. pantaea ^ bie MACARANGINAE $ te “CLAOXYLINAE 68. Er у һгососса obanilia aie ACALYPHINAE Acalypha £ ai LASIOCOCCINAE siococca A P ауд 7. Hom — 31. Tribe PLUKENETIEAE 31a. Subtribe PLUKENETIINAE 38. Haematostemon 39. Astrococcus si Angostyles SR 4 ченее суз E - sey 315 eine TRAGIINAE . Che мнений . Sphaerostylis Aci Pa sre pdt СЕ Бйр DALECHAMPIINAE Dalechampia 32. Tribe OMPHA LEAE 3 - 4. Omphalea . Subfamily CRONE 33: boi MICRANDREAE . Subtribe MICRANDRINAE 205. Micrandra 206. Micrandropsis 207. Cunuria 33b. ney HEVEINAE Hevea 34. Tribe MANIBOTEAE 215. 35b. Subtribe ENDOSPERMINAE 216. End 36. Tribe CELONIEA E . Suregada 37. Tribe ни um 38. Tribe JATROPHEA E : iaeum 235. FN coment 236. Acidoc . ВІ 238. Strophioblachia — А ———— — — Volume 81, Number 1 1994 Webster Synopsis of Taxa of Euphorbiaceae TO ; . Ad 1 + созы ОШ ы ~ 40. Tribe TRIGONOSTEMONEAE ү Glyphostylus Trig 41. Tribe a. RP DEAL ^ E 41а. Subtr = RICINOCARPINAE . Duvigneaudia 24 . Maprou ion mu on . Hippomane 47. Tribe PACHYSTROMATEAE 41b. BERTYINAE s 295. Pachystroma rtya 48. Tribe REA Myricanthe Р евз) occonerion Alge Borneodendron 42. Tribe CROTONEAE d aem í Mildbraedia 49. Tribe UPHORB . Fahrenheitia 49 Lewes ANTHOSTEMINAE : je rot . Anthos Cro : or ruinam 43. Tribe "INODENDREAE 49b. Subtribe NEOGUILLAUMINIINAE Givotia 02. Neoguillaumi . Ricinodendron 8: "рз aka: А . Schinziophyton 49c. Subtribe EUPHORBIINAE 4. Tribe ALEURIT uphorbia 44 чире ЕЛЕЕ 05. Chamaesyce 56. 306. Cubanthus MR Reus 307. Monadenium 308. Synadenium 44b. е Ан CARCIINAE 309. E dei 59. Garcia 310. Pedilanthus 44c tribe GROSSERINAE ava 1 Sedis (within Euphorbiaceae) Е "ipei : ?. Tapoi 3IL Chlamydojatropha Кешш н 312. Магїтепа L San АЫ 313. Tacarcuna nnodia 314. Avellanit 315. Cubacroton 444. Subtribe CROTONOGYNINAE Fio ee 67. Cyrtogonone 317. Cladogelonium 6 8. Crotono aye APPENDIX 2. List of new taxa. 44e. 69. Man ЖО NEOHOUTONINAE - Neobout Bard М; ww EUPHORBIGIDEA m E STOMATOCALYCEA = bt 2 Равон STOMATOCALYCINAE e war 45b. Satie HAMILCOINAE заан 46. ie HIPPOM MANEAE > e c > e ё ES €— MABEINAE . Mabe 279 Pes Ider ن ا‎ 278. : Sie CARUMBIINAE Omalanthus : Sate HIPPOMANINAE cen l; 282. 283. 284. Excoecaria 285. iguaia 286. Dalembertia CROIZATIEAE Webster, trib. nov e TA ACOCCINAE Levin, subtr. n TURAE (Muell. Arg.) AW bec stat. nov. PICRODENDRINAE (Small) Webster, stat. nov oe pesi Arg Е) Ns bster, st. nov. un à ә as] = 2 > т E > = е SPONDIANTHINAE Wie bier ig Webst ter, Acidocroton sect. Ophellantha (Standl.) Webster, pot nov 1 Webster, comb. ni Acidocroton оета (Stan ) Кене» ‘em nov. Webster, com! Desens thyrsiflorus (Airy Shaw) Webster, comb. i ) неч, сотЬ. поу e eres us (Ai haw) Webs Leptopus — (Nin. ) Webster, с . nov. Lee "yt manongarivensis (Leandri) Webster, comb. Orfilea — (Baillon) Webster, comb. no audiana (Baillon) Webster, с comb. no’ алаг chauvetiae (Leandri) Webster, Е поу. 138 Annals of the Missouri Botanical Garden Phyllanthus — = Webster, comb. nov. Amanoa ا‎ Pile thus hm + ut Д Webster, uk nov AMANOEAE uropus d ei yov comb. nov Amanoinae Amperea 1 DIX 3. t a. This AMPEREAE T mage includes all the gener: ee taxa існа myrea 1 > paper. pte М taxa аге = in ith Anabaena 1 ale ageneric taxa all capitalized. S: nonymised taxa аге — Anabaenella 1 italicized, and those merel: ti Anaua d text. Taxa are Bagues rete m in the text, dida 2 those in Roman numerals referring to subfamilies, _4ndicus 2 on preceded by “т” referring to re dta or subtribes, Andrachne those preceded by “E” watts to excluded taxa, and ANDRACHNINAE те others referring to Androstachys 7 Acalypha 184 Angelandra " : Acalyphaceae Ill] Angosty ud ACALYPH EAE T30 р Rk — 9 ACALYPHINAE T30j T ACALYPHOIDEAE ш pe sees 2 Acalyphopsis 184 Anisophyllum э Асын онин 199 Annesijoa » Acantholom 29 6 ccia 194 ospermu 2 roton 236 Anomostachys * Acidoton P. Browne 25 Anthacantha 3 idoton Sw. 201 nth ma Tis Aconceveibum 176 — Anthostem Т49 Actephila Ahoutmideió Ti Actinostemon 283 ANTHOSTEMINAE 144 са tidesma ADELIEA T28 ANTIDESMEAE T Adenochlaena 137 bir — À | - see 214 1 35 Fes coo иа | ADENOCLININAE T35a Aphora 1 Adenocrepis 4 Apodandra Adenogyne 281 iscus num 140 — Aporosella Adenopeltinae T46c porusa T pelti 287 Арогиѕеае Adenopetalum 304 rachn 1 Adenophaedra 1277 Argomuellera 1 pete ene 219 Argythamnia Adis De А eitonia 3 Adriana 159 rthrothamnus Adrianeae T30b Ashtonia ADRIANINAE T30b Asterandra 2 Аедоргїсит 293 Astraea Aerisilvaea 27 Astrocasia T Agaloma 304 ASTROCASIINAE 1 AGROSTISTACHYDEAE T22 Astrococcus 2 Agrostistachys 105 Astrogy Agyneia 33 Astylis 1 Alchornea 151 Athroandra 2 Alchorneiformes T29a Athrois ALCHORNEAE » Austrobuxus 3 ALCHORNEINAE T29a Avellanita 1 psis 122 Axenfeldia Alcinaeanthus 95 Baccaurea lcoceria 286 Вассаигеорѕіѕ 240 — Alectoroctonum 304 Baliospermum 226 u 256 Baloghia 952 ALEURITIDEAE T44 Barhamia 146 ALEURITINAE T44a — Beltrania 10 Alevia 123 Bennettia T20 Algernonia 297 Bennettieaceae 10 Allenia 76 Bennettieae xb Alphandia 243 Benoistia r3 Volume 81, Number 1 1994 Webster Synopsis of Taxa of Euphorbiaceae 139 Benoistieae — DIEAE ERN Pere Bertya BLUMEODENDRINAE Blum vo Bocquillon Bonania Borneodendron Botryanthe LIEAE D B Ti iedelioideae Bureavi t$ t$ м - e ч — N س‎ N = = دب ای تا چ ي و دب دب دپ D ECC — cc Centrost ylis Cephalocroton Cephalocrotoneae Cephalocrotonopsis Cephaloma CEPH ALOMAPPINAE C rant dta Chlamydoja dd Chlor: v Chlor. ane асау Chonocentrum Choriophyllum 6G — Á Cnesm ن‎ Сос Fone - N‏ wy‏ بن تح قي > к кеке U سے‎ е N -N H‏ يمس اس к о 9 Annals of the Missouri Botanical Garden Coelodisceae CROIZATIEAE Croton леш Croto CROTONOGYNINAE Crotonogynopsis CROTONOIDEAE Dalecham DALECHAMPIINAE Dalem Danguyodrypetes Decar rinium Dendrocous Dendrophylantus Dendrothri Dimorphocaly Dimorphocladium Diploc Dli - w N kel ONAN “ae PNP لد‎ 2 л (л у d юн م م“ خم‎ Ww 316 4 Di iplomorpha Discoglypremna Dispermae Dissiliaria DISSILIARIINAE eyri XINAE xis Ditrysinia Ditt Bocas Dodecastigma omohinea DORYXYLINAE Du DYSOPSIDINAE Dyso Eehinocroton ноен Elaeo, Elae drin rbia ELATERIOSPERMEAE cer Endaden ENDOSPERMINAE Endos rne hire Уј Е Еѕша EUPHORBIOIDEAE Euphyllantheae Eutropia Everettiodendron FLUEGGEINAE (or PHYLLANTHINAE) Flueggeopsis = 7 ) х ә 4 ү И 252 264 A и 46 250 28 % 4 9 - Volume 81, Number 1 1994 Webster Synopsis of Taxa of Euphorbiaceae = ae | Foersteria Fontainea Fragariopsis Fı reireodendr от CALEARIEAE CARCIINAE Gonatogyne ren GROSSERINAE HAMILCOINAE Hancea ermesia Bo. Heteroc. alyx HIPPOMANE AE HIPPOMANINAE Г. — o Ix {— x t ee] чале Ф d + Holstia Homalanthinae Homalanthus Homonoia Humblotia enanche Hyaenancheae HYAENANCHINAE enoca rdi 1a йүрек Kleinodendron Koelera й س ی یا ج‎ = o о Ше] T30f AN ea -2 = i سس‎ toU E EE NN о лон сс оо СС ® Wwunod6 wo س یں 9 سے re N ь Myricanthe Myriogomphos Nageia Nanopetalum ealchornea cepsia 7: сандан NEOGUILLAUMINIINAE Neoholstia Neojatropha етар eom Mopalissyn Neoroeper: Neoscortechinia Neot Neotrigonostemon eo Nepe. int s кону Nied м e as Осака Отр Ophellan 0 PA RADRYPETINAE Paragelonium Paran ia Parapantadenia Parodiodendro we Beaks Volume 81, Number 1 1994 Webster Synopsis of Taxa of Euphorbiaceae assaea 123 Pseudagrostistachys 106 ausandr: 229 PSEUDANTHINAE Tlld шеи 112 Pseudanthus 7 edilanthaceae T49c Pseudocroton 4 edilanthinae T49c Pseudoglochidion 9 edilanthus 310 PSEUDOLACHNOSTYLIDINAE T4c eltandra 0 Pseudolachnostylis entabrachion 1 udotragia 1 ега 3 Psilostachys 1 Peraceae T16 erococcus 1 РЕВЕАЕ T16 tychopyxis 1 erula J3 Putranjiva Petalodiscus 4 Putranjiveae Petalostigma 72 yenocom 1 Petalostigmateae Tlld PYCNOCOM T PETALOSTIGMATINAE Т11 PYCNOCOMINAE T26a Phaedra 123 Pycnosandra Philyra 111 Quadrasi 169 Phocea 167 Ramelia 1 Phyllano: 49 Ramsdenia ) Phyllanthaceae I edia 1 Phyllanthaceae T4 egnaldi PHYLLANTHEAE T4 Regulares T Phyllanthidea 24 eidi Phyllanthinae Тағ Reuteali 257 Phyllanthodendron 29 Reverchonia PHYLLANTHOIDEAE I Rhy Phyllanthus 29 Richeria 4 Р hyllaure 234 Richeriella 5 Picrodendrace T12 Ricinella PICRODENDREAE T12 RICININAE T30a PICRODENDRINAE Tl2a Ricinocarpaceae T41 Picrodendron RICINOCARPEA T41 Pierardia 4 ICINOCARPINAE T4la Pierardieae П Ricinocarpoideae 3 Pilinophytum 2 Prisa га Pimelodendron 273 RICINODENDREAE 1 Piranhea Ricinodendrinae 1 Plagianthera 1 Ricinodendron piagiostyles 2 icinus roa ma 199 Riseleya etostemon Rockinghamia 1 Plukenetia 1 Roeperia 2 PLUKENETIEAE T31 Roigia PLUKENETIINAE T3la Romanoa 1 podadenia 132 Ronnowia 204 PODOCALYCEAE T10 Rottlera ws Pod CALYCINAE T10a ROTTLERINAE ү" ; E р odostachys 2 Sagotia 1 Poggeoph $ 252 Sajorium yton 168 Samaropyxis порһог: ampantae 166 poCONOPHOREAE T14 Sandwithia e р. опорћуШт 2 Sapioideae : oilaniella PI: 288 Poinsettia ры h 281 Polyan, 304 Sarothrostachys т Polyboea 1 Sauropodeae 3] Polydragm S TI 2 rs = s oranthe S 0 47 ршатһеге Є Тбе оен и Р, RAN RINAE T6e 5сераѕта m orphyranthy EPINA Тбс Prosartema 102 E 255 Prosopidocli 241 Schinziophyton 93 Pros ttneae T16 cassie r 3 8 chistostigma - 9megabaria 40 . Schousboea - n Annals of the Missouri Botanical Garden Sclerocyathium Pea ea ae SPONDIANTHINAE ondianthus STOMATOCALYCEAE MATOCALYCINAE Tapo таре, Telog Tusc т. TETRACOCCINAE Tetracoccus Tetraglochidion Volume 81, Number 1, PP. = hs of the ANNALS ОЕ gr ан BOTANICAL GARDEN ublished on March 21, w etraglossa N Ыы N б О awe op E du l5 Led S 3 ^ =: br У‏ س bh b2‏ س t» о -1‏ 66 Sz = Ё‏ E ч‏ НЕЕ ЕЕЕЕ 5 аё б Е. > - & т 2 3 м ry = & 3 d ut ده‎ СО دب نھ‎ С ي ي‎ 1 ® © 3 aR рч © "S 2 a "Ins me^ “aR $955 2555 е з Ej ЕЕ B3 Pi Tri TRIGONOSTEMONEAE Trisyngyne Tritaxis Tylosepalum Tyria Tzellemtinia Uapaca Uapaceae UAPACINAE e ج سي‎ СИ СКИ ДБ Victorinia Vigi Voata mal 7 Wa "та пта TEETH moyo 5 е 3 Р. 5 һм м м دب‎ u N 58 > © N gospermum Recent Issues of the Annals of the Missouri Botanical Garden Phylogeny of Asteridae Annals Vd the Missouri Botanical Garden 79(2) (Spring iro T nii sister group (or sponsor was held at their annual meeting, 7 Aug. postage. nd praon of relationships of the Asterida by L ord he Bo in Richmond, Virginia. 238 pages. $ INT Brokering: The Mechanics of Synthesis Annal. s of t Systematics Symposium of the Missouri B discusses the val m “Phylogen ny of 27.00, plus Missouri Botanical Garden ЗЕР: Spring: Seu Vs symposium, the 38th Annual 1991, in St. Louis, Missouri), alue and future necessity of сакате Рмн Эг NEE into new and broader кыы: taking advantage of computer-based data-management systems. 41 pages (entire issue 230 pages). $35.00, plus postage. rbcL Sequence Data and Phylogenetic Reconstruction in Seed Plants Annals of the Missouri дени ИГ ER 80(3) (Summer ЭРЕ a. 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Box 299, St. Louis, MO 63166 0: | Thi ' is 1 i РУУ Злез $46 am Чал 72 of paper): | Volume 81 Number 2 1994 SYSTEMATICS OF THE EUPHORBIACEAE 5 ^ me аараан оосар оп фе Systematics of t ERÎ Annals of the Missouri Botanical Garden КД | hi er Support for the confer p roceedir ided by National Science Foundation den in St. v on 14-16 August 1989. Papers were nted by 2 people expel of Pe "m one or mo icto $ eek hat ыш for pienin and are d in this and the one prece edin Grant BSR- 881 17991. The denen papers benefitted from the editorial assistance and scientific reviews of Mi chael J. Huft, Geoffrey A. Levin, and ps us L. Webste MISSOURI BOTANICAL MAY 1 0 1994 GARDEN LIBRARY THE CONTRIBUTION OF R. №. Kapil? and А. K. Bhatnagar? EMBYROLOGY TO THE SYSTEMATICS OF THE EUPHORBIACEAE! ABSTRACT ы ffiniti Although their vegetative 3e d s ^ E - з { ‹ 1 ix € PERS ү the embryological wd da edm are markedly р labes nships. Tu "rcd sisti uod Diac of the fami чи аге: (1) is pue anther, 4 S-la Mu ed anther wall, fibrous endothecium, ephemeral SM la cae sy rar tapetum with m camer i simultaneous с cytokinesis, SHS nay decussate and e" bila "mi nic rads, and 2- or 3-nucleate pollen; (2) Дре; syncarpous, and trilocular ovary with o axile sq entation, sno е or h lo ос r hemian nat ropous, bite egmic, crassinucellate ovule, placental pes rand presence of hypostase; (3) em embryo sac both integuments forming seed coat, and outer epidermis of inner integument developing into a fibrous «n s P^ aie fi fre Моң ot 3 gnifica: nt Я the D uphor асое pops It UA more appropriate that the E ph biales should inclu e rm f the Euphorbiac e sub Cn ^e t the eis diverged ater ay di the Pla "Olen, and later the tri попой and ће Eu ideae. Since only 16 of the 50 ve so far been embryologically me he tribal probes into their reproductive biology. 1980). Wagenitz wn rightly em- av e Euphorbiaceae represent a large group of крш h plants distributed in almost all parts of ized shat “for elucidation of the systematic the world, except the arctic pat e family p pem es 300 genera and 8000 species (Webster, an Mee arie characters are especially valu- ма №, predominantly ا‎ in the tropical ا‎ as classical characters of flower may World. tropical areas of both the New and Old seriously altered by aie ” The embry ological cause of the great АЦА іп vege- characters have ak ig useful in: (i) d E f floral characters, the gro s been e th axon, especially at the level (Webst as het "CO PR and highly. adem pi ler; (ii) It D. ЩА Согпег, 1976). э ыл ies аге suspected; and (iii) or phyi ing inc де А ealized that a natural di ар a scheme of classifica Phylogenetic irn ation, be it of the angio- In the Eu ene e, the BRE UAR charac- нац ла а whole, or of a class, order, ily, ters exhibit t seit i diversity. They help in based ely on morphological charac- —recognizi atural groups down to the level of 1 Оп г hand, the therefore z "ox » seo an rata ы ae ynology the ge ric level and above. They are t only i circumscribing the family, but es SHER as cy os anatomy, det vinci ing n рың ependable tool also in j bios its а вА ematics. Embryo Sich бзи аге es- into subfamiie es anı x m in classifying it Н pholo, Y significant in situations where exomor- Cons may а either leads to inconclusive correlations or ины! ds *n delude due to convergence (Kapil & i about 30 genera), out y rge € the appa tribes nown. Only hm Michael J. Huft, Geoffrey A. Levin, and two anonymous reviewers for comments and suggestions. ent of Botany, University of Delhi, Delhi-110 007, India. ANN. Missouri Вот. Garp. 81: 145-159. 1994. 146 Annals of the Missouri Botanical Garden by Webster не 1989, privately muda have been investigated. Some genera of econ importance б б, Б, Ri icinus L. and Hevea Aublet, „© g Acalyp a Eb: н L., and Euphoria L., are well ом ied uis has dep iE d o ility of research material és een: on pes atic considerations. In the Phyllanthoideae, Putranjiva Wall., Phyl- J.R. & 34 lanthus L., Breynia . Forster, and Bis- chofia Blume are embryologically. wel: known, whereas eL al uropus E. Blume, and Secu nega Juss. are La under- 6 The Oldfeldioideae are almost unk cept for information on the ovule and seed of Micrantheum Desf. and Androstachys Prain, and the pollen and ovule of Picrodendron Planch. ашар" Acalyphoideae, ie = tribes, i is familiar belonging to three tribes, ins Chrozophora Me Juss. of Chrozophoreae, Mallotus Lour., е b Ricinus, Micrococca Benth., Trew ., and Аса lypha of Acalphyeae, and Tragia d and Data: champia L. of Plukenet In the Crotonoideae, which have been subdivi to 13 vibes by Web- ster (1987a; 1989, privately circulated), only sev- en genera belonging t ibes ha een ex- plored. These are Hevea, Manihot Mill. ‚ Jatropha, CHAR Juss., Croton L., B Blume, and Aleurites J. R. & G i b RT large bod data are able PEN rbia е ЧУ other genera that pay ve in TE Sapi E um P. Browne , Excoecar ia L., and Hura L. The paucity ‘oe embryological data can be part — to the difficulty i in n collecting a close series f and fruits Th e F 5 е laborious and time-consuming procedures. In Р nagar, 1980). More and more taxonomists аге a Dx ү, ОЛИ" £ : : 1 suggesting new schemes classification and tax- onomic i lS {ый Dahlgren, 1975, 1980; Takhtajan, 1980; ай 1981; Thorne, 1983; Goldberg, 1986). Нат main wd. of this contribution is to document the ways in which embr pu ae in ‘the Euphorbiales, asin affinities ‚ and in deter: g how best this guess can be classified into зайн ies. EMBRYOLOGICAL CHARACTERS OF THE EUPHORBIACEAE The following set of correlated embryological (see Wunderlich, 1967; Kapil & Bhat- nagar, 1972; Webster & Rupert, 1973) charac- terizes the Euphorbiaceae: 1. Tetrasporangiate anther, 4- or 5-layered anther wall, fibrous thickenings in endothecium, ephemeral middle layer(s), single layered ta- of t i i uclei, simultan cytokinesis, nee or ыштан microspore tetrads, 2- or 3-nucleate Б, grains. 2. Tricarpellary, syncarpous, trilocular ova sinucellate ovules, ventral raphe, funicula 1 inating in chal entering ments, placental © the base of nucellus or inte: a Е б. о Я 8: 3 ga 5. 8 к 3 leafy cotyledons S ; 4. Seed with copious endosperm conta aining oil oil and starch, эеле of hypostase in the di formation of seed coat by both mes uter epid of inner integument d ? 4 tiating into mechanical palisade or fibrous me and proliferative tip of outer integumen producing the caruncle. DISTRIBUTION OF SOME EMBRYOLOGICAL H ERS IN THE EUPHORBIACEAE EE nuclear n ere is an pU -nuc esting assortment of distrib tion of 2- and 3 ge is pollen grains in horbiaceae. Euphor at е one of the five exceptional angiosperm gen 16 species of Euphorbia investigated, some had 2-nucleate and others 3-nucleate Le grains а He concluded that the distributio Volume 81, Number 2 Kapil & Bhatnagar 147 1994 Embryology of the Euphorbiaceae TABLE 1. Embryological studies in the Euphorbiaceae. +: presence; —: absence. An- o Bud do- Em Genus (tribe) Author(s) ther SES s sperm ics Seed Phyllanthoideae (11 tribes) Putranjiva FN еи, Dutt eng yee s Dut crate ot + + - + (1944); R (1 d a) Neope E eei (7 Meineckia Bail- P. N. Rao oes RR + + - =з = = lon) (Ph ae) d et al. (19 veia ram " (Phyllantheae) TONN ar & Ka ^ AM Merc humo em icm lished); Corner (1976) Breynia J. R. & G. Forster (Phyllan- Thathacha we Venka- + + + + + + theae) teswarlu et al. (1973); Cor- ner Ki Melanthesa Blume (— Breynia J. R. & R. P. Singh (1968) : :E m Т лж $ G. Forster) Serer mre a L. (= Glo hidion 1. n 6. Mukherjee (1962); Corner + + + - - t Pal (Phy n 1 197 Phyllanthus L. (Phyllantheae) R. P. Singh (1956, 1972); + + + + Ж T Dang- Van- Liem n 2); Mukher (1964); estate уч Singh 1969) Sauropus Blume (Phyllantheae) P. N. Rao (1970) HES x NEM Bischofia Blume (Bischofieae) Bhatnagar & Kapil (1973, graue Dope See UR 1979); Corner (1976) Oldfieldioideae (3 tribes) ps Prain (Picrodendreae) Dahlgren & van Wyk (1988) = + - - - + Picrodendr Hain RM о - - - - 4 (1985) Micrantheum Desf. (Caletieae) Berg (1975) ол о OT Acalyphoideae (18 tribe Chrozophora A. Prid сане Кард. (1956a, b); Dang-V fae и MES тп (1962); Вог & Kapil Ricinus L a от a үр: " - (Acalypheae) R. P. Singh (1954); Dang- pu Jo. Г Van-Liem (1962) Місгососса Bentham (Acalypheae) PON Rad ПОК Мк (o ot To t t. or am (1963) Acalypha L, (Acalypheae) Tateishi (1927); Landes mug oom rt T y Thathacha ( ); Johri & (1953); Kapil (1960); Mu- M д kherjee (1958, 1 Mercurialis L, (Acalypheae) aan Lení(1962) GET |. илышы boh Mallo, А 229) + - tus Loureiro (Acalypheae) йн (1934); ponow +з о IA (1953a); Raju & К (1953) Se pe L. (Acalypheae) Banerji & Dutt (1944) WOW To da ES, = авіа L. (Plukenetieae) Mukherjee (1962, 1968); BE E То, Nair & Abraham (1980) Palechampia L. (Plukenetieae) R. P. Singh & Pal (1968); G. — + Ec y m L. Webster & B. D. Web- ster (1972) 148 Annals of the Missouri Botanical Garden TABLE 1. Continued. Em- ryo Endo- Em An- bry - Em- ther Ovule sac sperm bryo Seed Genus (tribe) Author(s) Crotonoideae (13 tribes) а Aublet (Mi lreae) Muzik (1954); Bouharmont UM Ee DE (1962); A. N. Rao (1964) Manihot Miller (Manihoteae) P. N. Rao & Sarveswara Rao — е 5 a nido (1976); Dang-Van-Liem (1962); Corner (1976) Jatropha L. (Jatropheae) Mukherjee (1962); R. T T = E Singh (1970b) Baliospermum Blume (Codiaeae) P. N. Rao (1970) XD E ins = Codiaeum A. Juss. (Codiaeae) я (1963); К. = T T ет Croton L. (Crotoneae) P Sgh Aleurites J. R. & С. Forster (Aleuritidae) Wiehr (19 h (1965); Bor & Bou- 1974 a ) таваг (1 953b); Venka- + JU ЫЫ u & Rao (1963); R “а (1970) Сапп of E (1945); au (1955) Euphorbioideae (5 tribes) Exe xcoecaria L. (Шр) PN. ~ 1970) sh sb m 95 TOR Sapium P. B ) Ch 970); Corner 0940) то Pa tA Sprengel Ho) Thathachar (1953b); Nair F He i + ШР: Maitreyi (1962) Нига L. (Нигеае) Р Као ) + zm Vo rr Euphorbia L. (Euphorbieae) Kapil (1961); R. P. Singh & + + 4o Cope с Jain ; R. P. Singh (1 apil (1975); Battaglia (1986) 3-nucleate pollen within the Euphorbiaceae was random ане & Rupert (1973) аа 66 Ж cies belongin da e he 2-nucleate condition. In the Crotonoideae; 9 1 1 » f a: o h гр a 1 e, C Ric- о Се] odiaeae. inocarpeae, Aleurtidene, Neoboutonieac) and te nih һе! 198 Ча а) both the large tribes ippomineas ' and h 9 na 2 ster et al., 1982). In angi osperms, the 2- nucleate condition of pol- red pri a e ad- condition is regarded as irreversible (Webst Rupert, 1973; Goldberg, 1986). It не й seems » that in the 2 nucleate condition has evolved pede ярык, Mene: more than once in the Crotonoideae and Euphorbioideae. пенен initiation іп the ovule. nsidered to be ip Gron aquist, 1 1900h Hor e рм e layer and the outer chiefly from subderm of f the ovular primordium (Bouman, 1974). Bor 4 Bouman (1974) and Bor & Kapil (1975. sai the wipe occurrence ot subde rmal ini p of both inn i ents In pu ilii Moulins, E. geniculata M (Euphorbia oi ma сы m variegatum Bi (Crotonoideae), and Crozopho ora о obliqua (Aca vee oideae). In Bischofia le (Kapil & Bhatnagar, 1980) and Securinega we copyrus (Willd.) Muell.-Arg. (unpublish lished obs.) | EDEN o — |A—e | BEES IS2c—————————— MM T. Volume 81, Number 2 Kapil & BMC. 149 1994 Embryology of th integuments are initiated by divisions in Webster & B. D. Webster, 1972) ањ Tragia of the dermal cells of the ovular primordium. Thus, the zm 2 в: о e. 8 БЕ £ 2 == © Гј E £g e sig anthoideae, with duis initiation of ws ушы, the үч се елү too, con- Рр 78 y + ened base of the intepumients, particularly of the outer one. Similarly, in the dermal initiation of the integuments, the epidermal lav, T dv Mod f. P CLAN | т . B ж % portions (rim) of the integuments. It is therefore мө SANT that dit dermal иесше ath e e ааа Crotonoideae, and Eupho rbioideae uted to the development of the integuments 3. er peg eet nucellar cap, and obtura- A maj herje & т © 28 Am -1 a $ hed obs.) of the РЕЙДЫ һауе vel developed Am beak that projects beyond e i rds t er of the apical of nucell V par us. ieu et al. (1 973) pont ovules with promine; ent nuce iia cell. The vil ermal x the Prominent nucellar cap, which d @ ^a micropyle. Putranjiva roxburghii Wall. : Р. Singh, 1970 ever, - ntheum and Picrodendron ephemeral. Mic ind Е (Bere, 1975; fig. 29 in Hakki, tends fa and subdermal cells. The beak I Dalecha : rl the micropyle in Chrozophora. а (В. P. Singh & Pal, 1968; С. L. Mere, 1962) it grows up to the exostome as in Ricinus (R. 3 Singh, 1954), at (Tathachar, 1953, EA and Aca alypha Jatropha, манн 1962; сда Вог & Као, 1970), река Nair & мия 1962) posses a vai developed nucellar cap that does not extend outside the mi- cropyle. In Euphorbia, some species (e.g., E. hy- Lira pino 85 чне и 1957; 69) һауе а on Gellê beak, E. milii, Bor Roxb., Make i 1965) kave’ a short beak or a nucellar cap that does not extend beyond the mi- cropyle. Presence of a nucellar beak in the Euphorbi- aceae is more widespread and seems to represent the primitive state of the character. The beak ex- tends Toni "f Be micropyle and comes in contact (so that it is in the form of a extend beyond the micropyle) is correlated with of t bturator. In maller in extent, the obturator its trichomelike —— cell project into the micropyle and come in contact with the nucellar tip. Both болое and the nucellar Baky ар! ee РЕТ tube as well as іп guiding it to the embryo в за Their correlated development is, therefore, com- r plemen 4. Ontogeny of the — gametophyte. The monosporic, 8-nucleate female iem metophyte of the Polygonum type asians er 80 per cent of the investigated angios pow rns (Maheshwari, 1950) de Eu phorbiacene, the Pinte (Phyllan- a, Bischofia) have the Polygonum type о o ак phytes is lacking in the Oldfieldioideae. In the; Acalyphoidese, | —Ó Micrococca, and Poly apex of » male etophyte, м 1; Mukherjee, bern "Johri & ne een st 150 Annals of the Missouri Botanical Garden and Mallotus nne reprae, 1934; o 1953a) hav of Penaea, Acalypha indica, Peperomia hispidu. la, or Drusa types. decns boh too, are reported to have xen the Po- sac. In the а 0 ine onum type of embryois P» However, е genus Euphorbia is unique (e.g., E. geniculat, E. тій, Е. heterophylla Б hen Е. а L.) have the Pol type of embryo sac, dine show рон ie " Fritillaria/ Euphorbia dulcis type (E. dulcis; Kapil, 1961; 86) or Penaea type (E. palustris L.; 23. It thus seems obvious het a „Вобан тре Euphorbiaceae. From this, pears to have evolved S илы есин in some = дев: еве б апа Euphorbioideae. 5. Development of endosperm. Irrespective of ihe number of nuclei, participating in enc in J t GE 142 is remarkably nits orm in the — (see Kapil & Bhatnagar, 1972). The primary aperi puce bte earlier pe the zygote. Free 32-256 nuclei are produced. Wall formation is beds ined at the globular or heart-shaped stages o bryo. It usually begins at the micropylar cit e later киш along the, per peu of the embryo n Ricinus, Aleurites Hevea, Chroz еби: Micrococca Sebastiania, Euphorbia) or the w: y be laid down sim v taneously e: ка уау of the embryo sa in Acaly, ‚ 1953). Wall "a il the entire en- f £g o nuclear three Bis of Croton—C. sparsiflorus fed = клар ымы Ваш ws C. klotzschianus (Venkate eswarlu sa Rao 3), an t ; A 1 x al, маа it аа develops into an dinde ate struc- e. Simultaneously, free uclear divisions contin- e к, micropylar part the embryo sac, еш the erae portion forms a | free nuclear caecumlike н. which some- and behaves as an aggressive chalazal des riu h с eyes erm Т is a spe- o have evolved only in the advanced Haein o (Cea pe 6. Caruncle. In several Euphorbiaceae the es 75). The caru seed germination by absorption of water, deserves boe attention o Руано many of which are tree the seeds are mostly without with puse qs fruit, L 1 N a a (Р. М. Rao, 1970 Hippomaneae. In Барон, some species : well-developed caruncle (i.e , E. dulcis, Kapil 961), whereas others have a structure on os ventral micropylar side b 1974; E. geniculata, Bor 1975). In the Ph llanthoideae the outer integu™ xi 2-4 ce eer ks the outer integument seed is ecarunculate. In Micranthe Volume 81, Number 2 1994 Kapil & Bhatnagar 151 9 Embryology of the Euphorbiaceae Oldfieldioideae the caruncle is produced primarily isi f the inner dermal layer at the apical = ca- К рч ba Pp ru and dermal apical bel of the outer integument. Interestingly, in Eupho lata (Bor & Kapil, 1 975) : an inconspicuous ca around the micropyle, on the ii side of i young seed, by divisions of the dermal cells. эке а cpat of Euphorbia i d a ca- runc "nr т f n (F h ds ЧАН ` т & Bouma 74). , It seems, Er that formation of a caruncle is a derived feature in the Euphorbiaceae, and its absence in some Euphorbioideae represents the result of Belo loss of this feature. CIRCUMSCRIPTION OF THE EUPHORBIALES According to Dein (19872), “Over 30 ilies hav ve been included fam- es in eac aptati t of convergence on account o mates a similar noce ox Vaid and see ist) ` mbryol gether with ev- ous lines B. lynolo: (P 1p such as palynology unt, 1962; Köhler, 1965), floral ARDI (V. Asha Singh, 1975), and w nat пору idence from uphorbi- $E asam оге “i natural group, are considered here. cause of their simple flowers, Гу ovary, axile placentation, presence of mi each locule, and seeds that are some- times carunculate, were, and still are, often re- ferred close to the Euphorbiaceae i ins, 1974; Bens 1979; ООШ 1981; ДОШ аше example, in Buxus L. an obturator is present and the ovule has a nucellar cap formed by periclinal divisions i i i the Buxaceae integument (see fig. 3D in antipodal Legis videt ern ttp. and ia sion polars пи псев lie in the de sien of endos 2н, which is A nd seeds, Tte are exo-mesotestal (Wiger, 1935; Corner, 1976). А report of nuclear endosperm a Lindl. d n. There is thus little дикан even for the grouping together of the Bux and the Euphorbiaceae in the same order 2. DAPHNIPHYLLACEAE ost taxonomists now agree that Daph- should be berg (1986) consider it closer to the Hamameli- dales. palynological and =. investiga- iphyllum e Muell. Arg tions of рар imalayense (Bhatnagar & Garg, To: Bhatnagar & Kapil, 1982) assign it a distinctive position from the Eu- m an КЕ is ab- QR all ent, the en ular sm the seeds BAR perisperm pel are pene gii and ecarunculate Daphni phyllum resembles the Hamamelidales aul, ; Ka pil, 1974) bilocular ovary with incomplete p , two o in each locule, parietal placentation, anatropous, eit crassin te ovul ermal outer and dermal inner integument, bsence of an чнай persistent antipodals, cellular en dosperm and polyembryony. Ne vertheless, it is different from the Hamamelidaceae in having 3-traced stamens Annals of the Missouri Botanical Garden and an endotegmic seed coat. Thus, its inclusion in an independent йу; Эң енн, within the order Hamamelidales or in an independent or "à Daphniphyllales s. to it, appears more sat- isfactory. 3. SIMMONDSIACEAE Although there is consensus that Simmondsia t Nu d form an indepe mily, the Sim- mondsiaceae, its phylogenetic relationship is st debated. Cronquist (1981 ), Thorne (1983), aceae (Ember- ger, 1960), DR un уннан 1983), ог the Fagales (Goldbe immondsia io: ares a es Ша traits with the Euphorbiaceae—both have на bitegmic, disci ella te ovules and nuclea dosperm vin obturator and e ase Andi in its is pontepsing, exo- mesotestal that s Simmondsiaceae are not related to the Euphorbiales, and are perhaps quite out of place in the ales or the Hamamelidales as well. Gold- her s (1986) suggestion that Sim lated to the F mondsia is re- agales spes consideration. The a3- o 5-lay a us, e EA des bsc nr nn Ps ke colon (Corner elat tionship 4. BISCHOFIA Bischofia javanica, a tall deciduous tree, has been placed she mi tho tite Fhylaptteae o or in x уйш инек (лм ча 1969; m Airy Shaw (1965, 1966) as- rp nus to an independent family, СЯ which he thought was related to the —— hn елй study of Bischofia javanica eec tnagar & Kapil 1 ephemeral middle layers, a secretory tapetum with ultinucleate ce utn a tricar rpellary ovary. twò anat- b egm ип a ven tral raphe siti on axile placenta in each locule, dermal integuments (as in Securinega), formation of a nucellar cap and placental obturator, differ entiation of Rope a Polygonu ryo sac, раа seeds. Н m type of em- embryogeny, and exotegmic ce, the removal of Bischofia from the Eu я 18 ME for 5. PICRODENDRON This genus has been variously classified in the Bombacaceae, Burse i Ev phorbiacene, 4 glandaceae, Sapindac im- e (see Hakki, 1985). Wee яп Ка ig as a tribe Picrodendreae in the s Oldfieldioideae of the Euphorbiacea Hakki 5) iie n some Bis. ol floral mor- the West Indies species ) Krug & Urban. It e Euphorbiaceae in having late s bo: n axile place of a nuc cap, obturator, and hypostase. orphological and palynological features, Hay den et al. (1984) strongly supported an alliance ° Picrodendron with th fieldioideae je is distinct in havi bicarpellary, : ovary and embryo with folded cotyledons. Anm саи of the anther, female gamete A m, and seed of Picroden dron and the elie would be of considerable interest from poi xonomic view 6. CALLITRICHACEAE їп euphorbiacean connection of үе. d uite eril (We cell ferentiation of endot ре welll erminal haus mentous proembryos ith membranous ar the 1080; Lamiales (Stebbins, 1 akhtajan. ^ Thorne, 1983; apes: 1983). 7. AEXTOXICACEAE onotypic Put endemic to Chile. € toxicaceae ер to the Eup rbiales signed 914) Тайна (09 (1983), 85), 1019 87a) prefer it to be in th taxon is embryologically not well underst EE e Volume 81, Number 2 1994 Kapil & Bhatnagar Embryology of the Euphorbiaceae anatropous, bitegmic, crassinucellate ovules with massive nucellar beaks (Mauritzon, 19 ever, its En unilocular ovar ubapic o extending only up to half th di the cado pert, and degens apes the leng to the Euphorbiaceae. 8. DICHAPETALACEAE e Dic фар ийселе, a small family of four е rica, have been included in Milo anv hy p Pe PARTE (Cronquist, 1 ilarity with the Malpighiaceae. From what is known about its embryology, it se that it has a bi- or tricarpellary ovary, two ovules in each lo o axile placentation, and carunculate seeds like those of the Euphorbiaceae. However, s unitegmic ovules (Corner, 1976) and exalbuminous seeds do not accord well with the acinaceae (see он 1966) апа оїһег оа (Goldberg, 9. THYMELAEACEAE melaeaceae have been pes in the m elis, 1 974; Bens ; Cro oe? 198 Bl) ana i the Euphorbiales ( Chore, 1983) t zm independent order, the Thymelaeales, which Mii. to the Euphorbiales in the superorder tman (1952) €— the resemblance in pat " pape Pu Diae Mineli Co; панн (1976) Sues out that both thes taxa are racterized b ans id otegmic seeds. Other c on ogica dino are: tetr: asporan, em an ther, fibro d Ў tend; e 1976). ^ long the see (see Davis, 1966; Corner, melaeacea sd E bees phorbiaceae, the Thy- and multi е hav tylar obturator and persistent бшу: карда cells, and the endo- sperm is usually scanty or absent. It is за better to assign the Thymelaeaceae to а s order, the Thymela iiid next to the Euphorbiales in ше вир кеш эм lviflorae. The manae үе tales is much less convincing. ;. The Myrtaceae have п an obturator requ Ney, done. Corner, 1976 )a are ecarunculate, with a testal d coat. 10. PANDACEAE Pandaceae, another small family with four gen- a, is di tributed o in western Africa and ee D fruits ducis by valves (Goldberg, 86), th deserve ve to be segregated endent family, Em ical data become ше гы оп Шш Paniaccae. AFFINITIES OF THE EUPHORBIALES e Euphorbiales have been considered by dif- affiliated with one of the two ges with regar Faria have been mostly confined to these four order: 1. Celastrales (Stebbins, 1974; Cronquist, 1981, Ei Maps 1964; Webster, 1987a); . Violales (Hickey & Wo 1975); і Malvales (Croat. 1973; Takhtajan, 1980, 1987; Dahlgren, 1983; nio 1983; Gold- berg, 1986). The Celastrales (which may inc clude Aquifoli- ha aceae, Stackhausiaceae, Icacinaceae, Siphonodon- e taceae, Sphenostemonacea and Goupiaceae, for purposes о the present compa ison) are mostl trees and s t two to five carpels, one to fiv to ten locules (Siphonodontaceae), mostly one or two ovules per locule (two to six in Siphono- 5 154 Annals of the Missouri Botanical Garden dontaceae and up to 18 per locule in Celastraceae), axile El anatropous, edi egmic or uniteg- d 5a or- d raphe in ren мн чну Seng micro- pyle or y directed n ссн of ойнен а Polygonum ad or т Caryophyllad сре of embryogeny, minc opin ы capsular fruit жн, Euphorbiales in 5, unitegmic, tenuinucellat ovul es (in EE a dorsal raphe in ovules ith + iri. bisexual fl upward, single-celled arc sporium, 8- 1 9 antipodal: cells (in ау, cellular endos d Icacin Gana- layer. T structure of pida er, o enings in the outer epidermis of the inner integu- ment. The M however, far outweigh the similarit PCR (including Geraniaceae, Oxali- nt olaceae, Viviani- к tapetum, ve three to five pels, on ‚ or several ovul per locule, axile placentation, anatr pous, cam- pylotropous or amphitrop bitegmic, crassinu- cellate (Geraniaceae), u , tenuinucella (Limnanthaceae) or bi ui сно) ovules. Endothelium, eee and hy- I . The female e Po olygonu um type, the helobial, copious or metophyte is of ا‎ is nuclear or absent, the embryo has haus- шы атда SUD (Tropaeolaceae), а and Pini.) s arillate, and the exotegmen eso a me- onsi ме very among themselves. They differ from the Euphor in many features, e.g., in the presence of oeboid tapetum, several ovules in each locule, wid and tenuinucellate ovules, endothelium, lL : FLA RO 1 \ 4 pensor haustorium (in Tropaeolaceae), Solanad t of embryogeny, and artillate seeds. Although there is some resemblance in the fibrous or palisadelike exotegmen of the lidaceae and Ge — with the Euphorbiales, the behavior of the out integument is quite differqu кыша Сеа аге, therefore, unlikely to be e Eu phorbiales. he Violales are a pantropical and subtropical as the Flacour- es superior-inferior, usually monoloc ostly numerous ovules in each locu rarely axile or apical placent hemianatropous or orthotropous, br i ш ез viih саса He D урон н соп- sisting о (podium), t LET halazal part. Polygonum type of embryo sac, nuclear g sperm (often scanty or absent in see e acd um pied of e кучы straight е "s o co vl ed rom raphe, fibrou s exotegmen, and frui mere berry AD 1966; D. Singh, vorat Каш, 1970; Согпег, 1976). a ушЫ dice ni the Euphorbiales in 9 structure of a seed. How certain cha such as their monolocular ovary parietal placentation, absence of obturator: pe tent ve en tubes, ryogeny g from the raphe, and reductio a dhi Sion give the impression that they rsis- y, aril es, postase, multicellular захарни ода! с type of embryo sac, anii ly multiplicative), nuclear endosperm, with chalazal haustorium (Bombac сасеае), | | " Volume 81, Number 2 | 1994 Kapil & Bhatnagar Embryology of the Euphorbiaceae 155 or Onagrad type of embryogeny, straight, curved or folded embryo, extending all along the length of the endosperm, winged, hairy or arillate seed. with exotegm a palisadelike layer of sclerotic nec с, and capsule or schizo- ic fruit (Davi 966; 6). Thu major groups of dicotyledons is difficult to decide without some element of ambiguity ресама н» Various taxonomists, such аз Dahlgren reg Fon ap EUA 1988), Thorne (1983), Takh an (1987), and Goldberg (1986) have sabe their Cela: Sod Geraniales, Violales, and "a in different wa E and have placed them a beo from t ot Ф е Euphorbiales апа from у f the а, At the present level of under- standing, the E Bee are A closer to the Malvales than to the Be re. In view з among the Dilleniidae is justified о ds e grounds. CLASSIFICATION OF THE EUPHORBIACEAE The Subfamilies of the Euphorbiaceae as cir- kis by а 1025, 1987а) can be racterized embryologically as follows: cumse SUBF. ied l. PHYLLANTHOIDEAE: 2-nucleate pollen = D exine, two ovu! : ermal атт оѓ both integu eveloped nucellar beak, Pol bryo sac, a fibrous ex, onum type of e "n Wi doe LENE KEY m SUBF A 2 bonae 2-nucleate pollen usually spinulos sium inner epi- outer integument) with fibrous exoteg- SUBFAM] ILY 3. ACALYPHOIDEAE: e setis qu» i чац and noncrotonoi e, E € subdermal etas x oe inte; = ins ell-developed nucellar beak, monosporic i well as subdermal cells of outer inte 8- 0 ^ niic A exotegmen SUBFAMILY 4. CROTONOIDEAE: 2- or 3-nucleate pol- exine, one ovule y lr mostly carunculate seeds with palisadelike exotegmen SUBFAMILY 5. EUPHORBI OIDEAE: 2- or 3-nucleate р ps presence of nucellar beak or cap, monospo- E man sac saaie se аа е basis of ы distribution of the above- menti me feat no doubt that the they differ, usu № tion is саны, able. . As indice ted by Кечер & of 3-nucleate ‘pollen from a 2-nucleate condition within vd Gicupanidese and Euphorbio ad id A nuc r g is a trend of ер in the Ph е Oldfieldioideae, and Euphorbioideae. In Euphorbia ecie a nucella reas others — embry en remarka bly xinifomn, wit th initi; m fn » nuclear Croton spp. are exceptional in a having endosperm ll. The seeds are exotegmic and with the о some extent, indicating a secondary loss. thus 3 apparent that behind the facade ot т, that could have evolved on a a monophyletic basis from an as yet unknown group within the Dillen- " is likely that the evolutionary line that gave to the Acalyphoideae, Crotonoideae, and Eu- phorbioideae split rather early fra om ье A ied This is in pred because the biovulate condition, the she initiation of bot int eguments, а and the fieldioideae ?) are fundamental» different from the uniovulate condition, the s bdermal initiation of Old 156 Annals of the Missouri Botanical Garden both integuments, and the palisadelike exotegmen Euphor- goer both der . The өн otonoideae Euphorbioideae seem to hav ved as in- Puis lines from the اا‎ aa (cf., Web- ster, 1 b). Thus, the pattern of е of embryolog- ical desta rs in different subfamilies gives an Euph br ipsas that the evihuion depre the Euphor- ly ancestral condition one phyletic line gave rise the Phyllanthoideae and the Oldfieldioideae; it a second line gave rise to the Acalyphoideae from which the Crotonoideae and the Euphorbioideae evolved as separate lines. Corner (1976) proposed a cleavage in the ph on the basis of the fact that the Phyl- lanthoideae have fibrous exotegmen, whereas the otonoideae possess exotegmen with palisade (Malphighian) cells. According to him the pali- sadelike exotegmen of the Cr — "bow them ombacaceae- Malvaceae-Stercu RE On the other hand, “the fibrous exotegmen of Phyll аба їо the Celastraceae- Flacourtiaceae- Viola- ceae. Їп our opi he differences ous in —— family example, in the chalazal ne iD тайш is ma de palisadelike ne whereas in the me has fibers w ribbonlike thickenings. казао, se support the ана of the family Euphorbiaceae into five subfamilies, as pos- tastes by Webster (1975, 19872). Since only 16 are explored r premature to out of 50 tribes of the Euphorbiaceae a ve Ge it would be rather nion on tribal classification. Such a ps must await further of reproduction of quite a йе number of tax CONCLUSIONS А general survey of the Ea gis on the Euphorbiaceae gives a that i is an extensively eske ات‎ era a up of dian regions it work on the bi ology critical appraisal leads us to conclude that o information is confined to only abou o tribes recognized by Webste preposterous to list out the embry- ifi characterize this cuna, ìt is not ological features that uniformly family. чанат: display a peere homogeneity in e and development of anther. paiay ate ci meia d kie en- dosperm, embryo, and seed. Variation in ollen nucle umbe tegument initiation, extent of nucellar growth, development of the fe ga metop ncle ems to represent an uph a mo ong its инел allies, the Diaeta ctr and Ta пине аге get ee к: a >> inclusion i in the Euphorbiales along wit ith th Callitri We e tte: m the basis of distri es рнк Webster’s басан jon “of Eupho! orbi- subfamilies nm the "indt Aca Iyphoid early from the biovulate Phyl КО) the most pre nd the Eup ments on the tribal classification of this inte uphorbi- e embryology of жыз = de m [v] oO © oO = = Ф о E BEES g Ф 4 w ES @ © 5 В. > d = zc g ibes, for mp th and interpretation of the data, together ™ e formation from other disciplines, would go û way toward iren the saga o LITERATURE CITED f. new families of wilis otes on the genus Bischo аса Kew Bull. 21: 327-329. лашы, — m АЙЕ <> na Volume 81, Number 2 1994 Kapil & Bhatnagar 157 Embryology of the Euphorbiaceae ВАХЕВЛ, I. 1949. A contribution to the life a ie рт fallax ا‎ Arg. Bull. Bot. Soc. 3: 29-3 —— 2 M. K. I es) nA Аат of female gametophyte of the Euphorbiaceae. Pan bold heii eang Sec. B 20: BATTAGLIA, E 1986. Embryological У 8. Eu- dulcis type Mies. Fritillaria type. Annali nica (Roma) 44 -136. 25419692: Fécondation de l'ovule et dé- velop pement de la graine après croisement et auto- dla n chez Hevea [He майы Cellule 62: 119- BEER, Т. & & M. D. Sw 1977. On the theory of Н id к New Phytol. 78: 681- 4 Benson, L. D. t Cl 2nd ed. Heath Lexington, Маны Berc, R. Y. 197 Кош, | seed, and myrmecochorus dispersal i 5 i m TRAN Norwe, 3- J. Bot. 17 BHATNAGAR, е: к. 19 Morphology and Embryology of аха of rcd Affinities. Ph.D. The- sis. Univ. of A lhi, Indi. 95 MEENAKSH 1 GAR 1977. Affinities of Dap hniphylun — Palynological approach. Phyto- mor Lig logy Жыл ; етее N. Кур 19 Bischofia javanica— Its m with Rad eres Phytomorphol- Те 1979. Ontogen d y and taxonomic significance of anther in aiios javanica. Phy- __ morphology 29: 298-306 — Re deve tbe nt in with a discussion on aphniphyllaceae. Plrticio Bor, J. E ^4 Bor 1974. 1982. See Daphniphyllum himalayens taxonomic posi о of 32: 66-8 Development of ovule teguments in Euphorbia milii and Codiaeum _tariegatum Piytonerphology 2 24: 280-296. От REN: КАР 1975. Eup pee genicula- 5T 97 9, Апаїг xdi ete ontogeny of MN bit . L. Jussieu s тості Acta Bot. eerl. 2 5: 385 -400. с Е 1974, el de Studies of the Ore on p KD. байы University of Amsterdam, The | rattan te "у J. L. 1967. The тойы = phylo- c significance of binucleate and trinucleate pol- 5 s in the angiosperms. Amer. J. Bot 1970. EL sebi ife Рена eni : Car. Sei gametophytes GH. 39: 17-18. ure and devel- eee in уды urinaria 1.. J. Indian ч 1 ЕР 19 "je uem cm L от Сонце t (Шу. Press, Cambridge. Les Euphor' biacées vues en elles чөн leurs 20 envers Pingieapernilc am 206. бока System of Classifi- o of of Flowering Plants. Columbia Univ. Press ay 098 : 1963. The Evolution and Classification of Flowering Plants, 2nd ed. The New York Botanical den, Bronx, New York. 75. A Lone of caon of the to demonstrate the distri- bution x charactors В T xs 128: 119-14 stem of classification of the ; Жеш pu a es Soc. 80: 91-124. 3. ral aspects of ^ oriens evolu- tion and ral epe e Nordic J. Bot. 3: 119- 149. @ 6 5 © ore A: 1988. Structures and Pa eae of le E ил to or centered in uthern Africa. Pp. 1-94 in P. Goldblatt & P. P. Us y H (edito но) эшн Systematic Studies їп African cn dte . Syst. Bot. Missouri Bot. De БЫ 1962. Recherches sur l'Embryogéne йе s icoques. Ph.D. Thése. Université de Paris, Davis, A eli: 1966. Systemic Embryology of the ngiosperms John Wiley & Sons, New ive 2. Developmen t of the microspores and the a behaviour in the pel e E н Wall. Sci. & Cult J 310. Sed RR L. 1960. Traité de Botanique (Systéma- ue), du T Les Vegétaux Vasculaires. Masson, Pi Exon, g 1952. € Morphology and Plant Tax- y: Angiosperms. 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E a 2: 197-201. 1953 аин ds cal Mises e p Lag чий Ire ae. П. Мн ay g. Pro Natl. FS Sci. India, 469 T4, 53b. Mor a ee studies in the Euphor- ae. S Mysore Univ. 13B: 363-388. 3. Proposed new realignments in 117. the * angiosperms Nordic ‚ Bot. А сув VENKATESWARLU, J. & Р ‚ Rao. Endosperm in Euphorbia асеае ку» occurrence of sat agg haus- ies of Croton Linn. Curr. Sci. 32: toria in two spec 514-516. —— — & D. S. Rao. 1973. Pvt un x чушг‹ obturator in two Euphorbiaceae. Curr, Sci. 128- ENTURA, M. 1934. Sulla poliembrionia di Mallotus japonicus Muell. Arg. Ann. Bot. (Roma) 20: 568- Flower reduction as baci prob. b. 96: 448- 578. WaGENITZ, G. 19 lem of a eats taxonomy. Bot. Jah 470. WEBSTER, G. L. 1967. The genera of Euphorbiaceae rA southeastern United States. J. Arnold Arbor. 48: 3 T Conspectus of a new — of the Euphoric. -— 24: 593-6 ہے‎ The saga of the spu ү review of lationshi ips in the Euphorbiales. асе dd ни Bot. J. Linn. Soc. 9 . 1987b. etes dd distributional history of the Euphorbiaceae. P. 309 in XIV Int. Bot. Con- gress, Berlin. [Abstract. p к. A. Rupert. 1973. ir Aap ge signif- ican pollen nuclear num! the Euphorbi- aceae. ee 27: 524- тта & р. Kournik. 1982. Systematic signific cance of pollen nuclear number in ر‎ aceae, tribe Euphorbieae. Amer. J. Bot. 69: . WEBSTER. The exire of Wieur, E. 1930. Beitr rüge zur r Kenntnis der Anatomie der wichtigsten Euphorbiaceen Samen unter beson- derer cn уй ihrer Erkennungsmerkma in Futter . Landw. Versuchsstationen 110: 39 98. WIGER, м ан — ета Studies оп the сее Meliacea , Simarubacea a Bur: PhD. Thesis. ni Lund. фе. Wowpenuics, R 1967. в on cer of the seed coat. ыу 17:3 ——————————————M——r 007 ш нн со 2. 1 LEGUMINLIKE PROTEINS Uwe Jensen,’ Ina Vogel Bauer, AND THE SYSTEMATICS OF %4 Marei Nitschke THE EUPHORBIACEAE! ABSTRACT Serological methods were used to elucidate legumin | ни нш ге, similarities between the major seed storage і 1 f l шап ите 1 1 subfamilies i bi d 1 witl from Violiflorae, Malvifiorae (both s: ie e gue and ni mn x {ай-е maintaining these t bel t categories is not supported by the data reported her оао сорри. of approximately аатай eiaa ij the major seed proteins dto speci range in gymnosperms (Jensen & Berthold, 1989, for of morpho ese erra such as broad- brad UR. and unpublished ш for the с as trees from the tropical rainforest, succulents from arid ecosystems, екн weeds, and even a Sal. 1981; Jensen & Cringe, 1983; ae Greven, vinia-like water plant و‎ fluitans Benth.) 1984; cem ‚ 1984; Bergner & Jensen, 1989; ке of this great div: d I ischer & Jensen, 1992). ese proteins have stantial family characteristics (latex, number of proven to be important characters for the eluci- аи: carpels, basic chromosome vo the dation of phylogenetic relationships in similar in- taxonomic position б the family continues to be — Thus, a priori ЕРУ have been ind 987). Are the Euphorbiaceae i oteins ical and a i n seed او‎ УН their relatively hh € ncluded in the Dilleniidae (e.g., Dahlgren, 1983; i evolution. A pos teriori Имеш s hav койы & Jensen, 1992), the rae (e.g., Eh- supported "p I i i rendorfer, 1991) or both (e g- Baillon, 1858; Pax, legu ave provided in a v temang y of syste lthough most authors consider the Eu- йы) ы 1980; Fa Mus. et T 1975; Min ы-у to m arisen monophyletically, the Fairbrothers, 1977, 1983; Jensen & Fairbrothers, ro indi wo (Dahlgren, 1983) am- lies give gives rise to ant n of di- or polyphyletic METHODS AND. MiTERDUS origin (Schultes, 1987). bl The classification dilemma illustrates the need Immunogenic legumins were isolated from t for additional taxonomic and phylogenetic criteria seeds of Ei j us Labill. (Ph Ha to improve our understanding of the systematics — thoideae), Ricinus communis L. (Acalyphoideae of this family. Molecular data ob ained from the Hevea brasiliensis ( An investigation of nucleic acids and proteins con- — Reutealis trisperma (Blanco) A. Shaw th lá ducted for other taxonomic g € proven to tonoideae), and a crepitans L (Eupho "i le for such taxonomic research. In thi deae). Sources of the are given in the , investigation we u comparative data from pendix. T proteins we cted in legumin molecules, proteins that have been suc- rabbits to elicit specific antisera (“reference 5Y* cessfully included in t systematic research for tems”). These antisera and the antigens from е Я similar projects (e.g., Jensen & Greven, 1984; 30 Euphorbiaceae species and 100 not PME Bergner & Jensen, 1989). biaceae species were used in the subsequent the ' We gratefully acknowledge the assistance of Jórg Herbst for the statistic analysis, and the revision “ manuscript Ьу D. Е. Fairbrothers. This investigation was partially funded by grants from the Deutsche Forschung gemeinschaft. * Lehrstuhl für Pflanzenókologie und Systematik, Universitit Bayreuth, D-95440 Bayreuth, Germany. ANN. MISSOURI Bor. Garp. 81: 160- 179. 1994. — Volume 81, Number 2 1994 Jensen et al. 161 Leguminlike Proteins dimensional immunodiffusion experiments on agar elec see e (10 they separate qualitative units; quantitative mea- беш» о the acterization of systematic useless whe tigating То? ү ems used аге re represented by the тысе е ап nti- h of antihadi determinants of the bec molecule of one spe- Ne These different kinds of antibodies а оп ospecific antiserum аге the “reference seg in our systematic research RESULTS INTRAFAMILIAR LEGUMIN AFFINITIES five reference systems, i. or the nthus calycinus, Rici inus со e., Phylla r е dat rding to the teas sida similarity in relation to the reference stem For the systematic evaluation of the taxa in- n cluded in riteria are es- relative ode of se- llarity in iiis to the re "ord taxon. y the iones anking gro l. (ii) Identical serological reac- are ant if gumi reese Pondence to the reference system only rep- ts a very small ah of к potential antigenic ^ар n abd (Tab * data are e ae on bh basis of a dis- crimination of the five subfamilies: Голое deae, ia поген od Acalyphoideae, Crot ideae, and Euphorbioi ideae, ТЫ owin ster's classification OTH 1 ). ing We d Hn Pie umm В СЕ чаа ae ien е а Fig. Іа), the def Phyllathoideae з ^ чай чан Вгеу- та, produ nd—some им er ME similar taxa (i.e., had the edd. similarity indices). ntidesma bunius Spreng., another member of the iiri gn is distantly connected. The reactions of all Crotonoideae and Acalyphoideae a did not oed th ter E! tivities; onl Ls | Н, ит апа Hura, have sie cm usd ud been obse gis УИ Chrozophora and Mer а райе ef tie is pene Acalyphoideae, re re- indices with the Ricinus communis reference system (Table 1b, and indice Fig. 1b). The similarit ices between Ricinus and p: Hen two me s of the Acalyphoideae, i.e., ham and especially Mallotus, are onoide: and Ho (о; "e orbioideae) a are serologically эй to react in a range of к. outgro rotonoideae. Tee sm di and pe highest tonoideae)r reference system (Table 1с and Fig. 1с): Mani or (Сое and Ri icinus. (Acalyphoi- 7 © ith Hevea than the other Euphorbiaceae ge ested. The Phyllanthoideae and Oldfieldioideae go ra үр е юн reactivi he serological re- es, Phyllanthus, Andrachne, and le “all Phyllanthoideac) with the pue of Antidesma, another member the Ph жеен thoideae, *°зригз”” the legumin precipitation o of the four genera indicating а a somewhat higher degree of similarity with Hevea other reference system from the subfamily Ee Reutealis trisperma (tribe Aleuri- TABLE 1. Data matrices for the serological results Р two l-dimensional gel diffusion experiments using m о systems, i.e., Phyllanthus eoe Labill. (a), Ricinus communis L. (b), Hevea brasiliensis (Willd. Juss.) Müll. Arg. (c), Reutealis trisperma (Blanco) A. Shaw (d), Hura crepitans L. (e). | legumin of the left species spurs the upper species; — legumin of the upper ma ы the left species; + double spurring; О identical pad @ blank. All generic names are райы! out in the Appendix. The data h trically. Boxes indicate taxa groups with identical or similar cross-reactivity. The interrupted line marks the ximum reactivity of the o оц Right column: Similarity ince ا‎ according to the formula 5-2 х 100 (9%) S = Z| + 42+ + V4ZO for each species, using the row data, where R = reference taxon and С = cross-reacting taxon. a 1 2 > АҢ 5 6 7 ЗО ШОЕ 432033. 14^ 15 16717 8 190. 20921] "29 э3 94 S Phyllanthus M 1 e | | | | | | | | | | | | | | | | | | | | | | | 100 Reverchonia arenaria 2 — € o | | | | | | | | | | | | | | | | | | | | | 93 Breynia fruticosa fin Ofer Ооо | | | | | | | | | (81) Andrachne colchica ao aoe Що! | | | | | | | | | | | | | | | | | | | 87 Securinega durissima 5 - — |O О e| | | | | | | | | | | | | | | | | | | | 87 ‚ populifolius orar — ee ee у | = |, ът Sapium sebiferum 7 DE CUM: 109 тәң E 9) | | | | | | | | | | | | | | 72 ura crepitans 6 = — poo o— TO Сс к x | | | | | | | | | | | | | | | 72 Margaritaria scandens 9 — — = ee nn XU ++ Lc EE | | | | | | 50 Om phalea trichotoma 10 — — жы у к=з a qme duo | zs | | | | | | | | 52 Adelia ricinella ЖЕ з = E ы а Бус а | + | | | | | | | | 50 Ricinus communis 12 = - m d xk Ee pln CE NP now 0 L5. | | | | | | | 48 Manihot laziovii 13 - m E - = = x ES t ё + o | > | | | | | | | 50 а brasiliensis EE eee кы сш SEE + e o + + | | | | | | | 48 Mercuri annua Ei. — Ec Se 2-5 0E о о ө + | | | | | | | | 46 Antidesma bunius е та .— ae Ser ee Е | | | | | | | 46 есе tinctoria 17 — — EC eM uc оо Е E E ome ЗООГ o | | | | 35 S. fimbricalyx = 8 Eum xc 1 8-409 RE m Ее е. OS OO Ox к | 22 Joannesia princeps 1 л E3272 1x ER meee TCT IGS 0-970 420 t] | 22 Mallotus tenuifolius cues EE E c4 Ec E ue o кыл NE. E Ou ON Os. 0.20. гот 17 Euphorbia characias 21 — — = = =o ee m Ше meg pce Bs iE = = е эе бузау | o І 15 urites montana = = = = = - = = = - ^ = = o o о = e o о 11 Croton tiglium Ек т 2 = = am E де me m э. = x хе mj e- oO. O сш o ej pha curcas 24 - Xu = не) = em 5 "c n zm s vu = E T E zi o o e 4 ay} JO sjeuuy uapsed jeoiuejog HNOSSIN TABLE 1. Continued. 7661 b ТОРЕ lo 14 15 16° 17 DE E E ES SE E 8 "9 18 19 20 21 22 23 24 25 26 27 28 29 30 оме ѕ comm ӨЛ ШБ S 35 УР eg vul E GR. E DPI АЙС EIS d4 ozophora tinctoria E € 1 uz vb ИЕА M pP emper qp] ЖӨН boc deos d god ecd а. асе Eom кашу, Жыкы perenni. 3 c" pono wp €] Wbepe Ар pcc OM c TETTE WE d o» Joannesi 5 Дс Е ОЕ Е E eq ВАЕ р [o o4 зр з ыы rM БЕШ Hevea brasi E oc cde — w-poo XS c] рсы a Fw pores ар. „ый ШЕЛ al ми ү ига crepitans К з o O | А a cp зн ыр, Ic р ЕДЕ S 31 EE EL E S H. populifolius Nc = o Ue M D MM cr *Lornempr oro gp 9 | alechampia scandens 8 — — — - — + + Ф | + | Ол CE = c ЛО гр Ер at Тү RS д Manihot esculenta Ио = че Е Р Ф ЕЕК А эү Е es ee fe es үнү үз I Eremocarpus setigerus 10 — + | | + + + + 4 "ccc it Hf eH a RE Town Sapium sebiferum BR v o pu coc cc TIMES Uu qo еро CD TL USD MEUS EIMEE E atropha curcas Ee m = pacco ee lee eam x 31 uuo tC E E E. aria bicolor pol e «tuc T r.a qui d ds rua IUE ol Euphorbia lathyris IR | c со 5 — 2 beu t e 1 1. SEE КАЛЫ ТЕТЕ а 2| ernicia montana ae РЕ E cb ОК x "LI dug ucut uus i T Croton tiglium Ко ане o o Luc. [i ыру n ED eov SONT ACH Lo ENS ты д Reutealis trisperma К occ nor з -.—uv c uM. «4| hoop xp E p | i ontana ER — — m e o oo c Ce c Haro | C d Lo EVITER ASIE v] Baccaurea tetrandra ga o с c шшс 0 o == (ЕР comet nt SO Шу 4 4 A hb p ыр SH p DÎ colchica NR —- o Se — coge E Eh UNE TD COE АА Xp oo eae peg А чан imum suffruticosa NO o ы 0 o0 — e d E pop ы i cg uw e Jb SE A КУА ES — — ri e чысы O X ы к иш уш — ne cue е йл суыкты еы | SEM Aa petes eustralasica MO oco d e NES — du ш fq E еы ue чы ш Tee iin c РЕ ЕПРУВЕТИ] one nee M — — -—-5- a TC TEE voe е D oa a Tet bres 0 ue ҮТМИ рыш 25 — COE — — оа ои = т ЕЕЕ кел АГ Fee | Bridelia tomentos е. Ено сена Ed = quee I m mr ы э Set eU cop ОО | | | Glochidion glaucum oim dp Tee qu e c ME c miu cu куер SL BR RUP Cie bales Mul philippinensis 2d з — — — — — — 4 — 4$ 1. 1 Wo косе i Rede c cp. X Antidesma bunius mop qM NU ico amm uocum n DIM ар x M mx m d e c eu ee READ уа! P. quadriloculare BER v == c woe e ЕТ ЕЕЕ SEC TECTA wore A кши TEE Loc. с JOQUINN ‘18 eunjoA че јә иәѕиәг $шәзолд әзцшшпбәл E91 TABLE 1. Continued. с үй 6 10-11-12 13 14 15 16 17 18 19 20 21 22.23.94 25 26 21 928 29° 30 S Hevea brasiliensis 1 I d qq qu | gc espe p 100 Manihot esculenta 2 pc ge qe $4 Eoi qe spe] 95 s communis 3 vro Merete ere це [eg] ode ЕА 91 H. populifolius 4 LU dmg r3 Exp trap px 2-06 Hura crepitans 5 IE px du cepe qo hele Т П 08 Croton tigliu 6 porc reus SY pm e disi Joannesia prince 7 poc uec deo noel cal do obtu 76 hrozophora tinctoria 8 p EA | | | | | 11 72 Sapium sebiferum 9 [p PESE TEST 4. d ада ра оа 7 Reutealis trisperma 10 E Iq re A AE И а pw peor Euphorbia lathyris 11 ut. dq мүш ушен Г | ЕССЕ E у ӨН Excoecaria bicolo 2 o. . 1 I. дызы ls reco Wr: 262 Jatropha curc 13 09. -1. 4 uo Г үт | SRE УЕЛ Л Ma 99 Dalechampia scandens 14 Eo 7 9. ua ee коор у + Ip cho deas a2 Mercurialis peren = E EE 9 5 — lh l. 1 ud m2 Aleuritis montana 0 = — = Elo oc MA +. ! ! Dese v 52 Mallotus philippinensis 17 — — — — = 1 Ф. 1 | | | | | | | 50 Putranjiva roxburghii 18 — — = DON LUE Ф r + Ser Е | Vernicia montana 9 - - = WP cuc ТЕ -— 4. | OE Ie mle A cae 90 Eremocarpus spec. 20 - - - ux re к сы o ы o. | | | | | 38 Antidesma bunius 219 — — - - - - - = - | - - | gps E] 34 Drypetes austra. 22 — — + “ыр — Sr uae к=к ie MEE MES eee be Н 28 Phyllanthus grandifolius 23 — — - RI ae س اب‎ — ++ = (КАО Bah TEA 24 Andrachne colchica 24 = - - - = — - = = = - -— ISOLE TS C Nod Securinega suffrutic. до = — E LUE LR ct ci ыз р с; ТО qx opes 19 Baccaurea tetrandra 26 = — = eee Gh Cee or er 315 b x sd 14 Glochidion glaucu. 25 — 3 = Rp эку рышы i m ш. Окас 10 Bridelia tomentosa 8 — - E mE m. yb de i Lb ш xni olm ESAMI 9 Micrantheum ericoides 29 — — - d M a c ee аата А = {=e i 7 Petalostigma quadriloc. 30 — — - DIL 2 ee ee SCSI ELS 0 ee (oot — 9 € —— gà Уйный ni du a 1; ati — ÁÀÀ——M—— Ql т pbb. aie чі әсл o cle oda uapıe5 [eoiuejog unossiIA v9L au} JO sjeuuy TABLE 1. Continued. d BTR N12 AS MOS d6 17:18 19. 20 2E 27252904 25 20 75 S Reutealis trisperma 1 | | | ا‎ b AS a | | | | | | | | | | | | 1557100 Гегпісіа montana 2 | | | | | | | | | | | | | | | | | | | | | 94 Aleurites montana 3 | | | | | | | | | | | E e | | | | | | | 92 Jatropha curcas 4 | | | | | | | | | l | | | | | | | | | | | 88 D. scan 5 | | | | | | | | | | | | | | | | | | | | | 85 Joannesia princeps 6 | | | | PAGE | | | | | | | | | | | | | | | 81 Croton tiglium 7 © I- E E | | RR a d | | | | | | | | | | 69 Euphorbia lathyris 8 —— 51 | | | | | | | | | | | | | | | | | 71 Hura crepitans 9 Au „есы зук „чар eee ged | | | | | | | | | | | | | | 65 Manihot esculenta 10 dE owed NEN | | | | Зи ыз | | | | | | | | | | 65 Excoecaria bicolor 11 ye me 4E o D AM | | | | | | | | | | | | | | І 65 hilippinensis 12 m rd c E e | | | | | | І І | | | | | | 62 Mercurialis perennis 13 у атуны кщн ыла о то | | | | Ea | | | | | | 50 Baccaurea tetrandra 14 = mu lu е. сш E) | | | | | | І | | | | | 50 Antides ius 5 Bo om ey рш Е ао aE | | | | | | | т | | | 48 E. setigerus 16 pin эш I укр Мыз жы IW Жо ШИГ | | | | | | | | 40 Hevea brasiliensis T - = -—- + Bare een se = а | | | | | | | | 37 Andrachne colchica 18 LoT ee уке c Пс ОсОО, me { КЕ е, | | | | | 38 Sapium sebiferum 19 = = ЕИ Е "c a | Es bp І | 29 Н. populifolius 20 = cy em e m у= pu xc cx E Qc | ав І | 27 Р. quadriloculare 21 = Teel Co ы M wes жа кен ЕМ ей uno 9:3 | | | | | 29 icinus communis 22 addc. сме r ы ше E e Жытын Жо гр ird um Qa | | | 17 ridelia toi 3 ex aem у ws c M C T MD СТЕ Ro ow CURT 1l — x 8 Glochidion glaucum 24 Col E хен Um. da E c To ocho. od | І 17 М. ericoi 5 тт а ge к с ie шш. чылы ы е O 12 Р. grandi 26 ЖЕСЕ МЫШ ме йү ч-сы. Шкок коры сч el Та cuc vr J 6 S. suffruticosa 21 a tI eM mE eA cmo m ep ы и е ызы д x. 4 v661 | с Jequinw ‘L8 eunjoA ѕшәјола әзүшшпбәл е Je uesuer 991 E TABLE 1. Continued. 991 _ on — œ 24 27 28 w © n Hura crepitans Ricinus communis Excoecaria bicolor H. populifolius Sapium sebiferum Joannesia princeps ernicia montana Dalechampia scandens us philippinensis ——-— t2! o00-1O0 0 р ом – —— ль w - – ч е 'e IU 10112. 13 | | | | | | | | | | | | | | | | | | | | | | | | ME sod | | | | | | | | Oe + us. NT оске M 1 + - - ө ТИЕ - = - + OO = + = SS a „ы pé eL es usu a. a to P. quadriloculare Micrantheum ericoides : à: Phyllant randifolius Andrachne c ridelia tomentosa Antidesma Glochidion glaucum БЕ 84} jo sjeuuy иәрлесу [E9Iuejog unossilN Volume 81, Number 2 1994 Jensen et al. 167 Leguminlike Proteins tideae), has been tested (Table ld and Fig. ld). Clearly the two other members of the tribe Aleu- didi. A opha- Caton Hura- Мо апї ne -Eremocarpus Again however, additional taxa were include d in this group of low cre RF OY (Euphorbioi- deae), E аны (дому акей. зекий Moe ine n ШЕ чу indices in both graphs (based on different, nonrelated sta- tistical procedures) арш аге ш Mey adequately axa. For the des reference taxa the relative po- r tan case for Mallo tus within Be deae (e.g cimi sod Reutealis) are e distantly locat- melos Sapium) within the Euphorbioi- with t A (Table le rand 752 Ме Ь df ° PCA has also been used for an ordination of a ; из со which is placed in the Aca- higher f taxa, t are included yphoideae. "he e identical genera Ex- in at least four reference systems ). Although со ria, Homalanthus, and Sapi e separation of th yllanthoideae ldfiel- phorbioideae) follow horbia reveals affinities dioideae from the Acalyphoidea Crotonoideae i are lower than most members of the A caly- + Euphorbioideae becomes less distinct, a high p ика and Emtonnideae. phe similarity indices Decimi t can be о d. The re- сок pattern g the cus en se- ical discrimination capacity y between Ricinus t nee and do not exceed the outgroup reac- ivities, м d таш aphic representation of the data patterns М tionship pet tween taxa, aratra dion tech- c oe U ee M rds ike UPGMA or similar hierarchical meth- of their т e ^i ix pen For the five reference e taxa, the similarity values ith little de- experimentally obtained data (Fig. is ers clearly demonstrates close serolog- CM ~~ between Ricinus, Hura, and He- ы istant position for Phyllanthus is re- hs for‏ اھ ры иа axa plus the 13 genera that have‏ 3a, b). Sin е reference experiments (Fig.‏ ce relative positions for the taxa included‏ Ac alyphoidese), Hura (Eh and He- vea (Crotonoideae) reference s n the con- trary, the taxa connected to ine rund Кеша (both Crotonoideae) are clearly separa EXTRAFAMILIAR CONNECTIONS For th idati f tematic con- or ms between uae rbiaceae pe other an- ctio giospermous families, we investigated approxi- ately 100 species from a large number of тез ае ek ilies. Th test ie ed for the rol res reaction of their leguminlike proteins wit ntiserum against Ricinus com- munis. One tot vn species were used to represent 0 окы serological reactions) were Asteraceae, Lauraceae, Pittosporaceae, and Ra- nunculaceae; families with low legumin similarity indices serological reactions) are show in Tabl Only the species of Table 3 revealed high legumin similarit indices, exceeding those of Euphorbiaceae species (rankin ed r h че Ricinus communis antisystem, wi ies as the t similar species; se lah. 9 in Vog el, 1986). All taxa of high legumin similarity (Table 3) belon 4 superorders lviflorae, Rutiflorae, and Кылы They are of interest in attempting 168 Annals Sehen "© Garden a» LJ LJ A a A A я a a п | ommuemus — | o O BREYNIA O ANDRACHNE O SECURINEGA 1.€., f SECURINEGA FiGURE i Phat? HEVEA @ MERCURIALIS О ANTIDESMA @ CHROZOPHORA Ш MANIHOT m HEVEA А НОНА “hy H @ RICINUS سے م نے سے — —— © tem. sys Niere; interpretation of the relative serological similarities. in relation to each reference (d), Hura € crepitans Volume 81, Number 2 94 Jensen et al. 169 Leguminlike Proteins © PHYLLANTHUS SECURINEGA с The interrupted line marks the maximum reactivity о е à y insignificant or undetectable serolo, Yphoideae; Ш Crotonoideae; А E e тесин t 0 interpret the ato oii of the Euphorbiaceae to the other cue ledon Discussion nsidered in this investigation m the other sub- les, i.e., Ac calyphoideae, Crotonoideae, and of the outgroups. Only generic names are used because of gical Ponss differences. O Phyllanthoideae; O Oldfeldioideae; LJ Euphorbioideae: the biovular locule (in contrast to the uniovular locule, characteristic for the other Ir iê Û ig phe ura (Euphorbioideae), Ric inus “hey em of th ae), Р Cte ОРУЙ do not —— gine Е affin- 170 Annals of the Missouri Botanical Garden PHYLLANTHUS ө Whe HEVEA E 2. des (S. D. 4 fo ities of the outgroups. However REUTEALIS Graphic Meri i of serological dis- erence taxa Phyllanthus, ‚ Ricinus, Ниг eutealis, ا‎ ж from a. Similarity pé of Table 1 (S.D. =S). cases our outgroup families: In the аа. (Сго- стат antisystem, the serological similarities of the —— taxa Baccaurea, Antides- ma, and к ne with Reutealis somewhat ех- ceed the serological e of the outgroups 1). S | with Reutealis (Fig. 1d). Similarly, 1 lanthus antisystem, the Euphorbioideae genera Ho us, Sapium, and Hura pl а communis апа Phyllanthus y s (Table 4), support proposals that suggest two dif- nin ee roots (Huber, 1985) for the Euphorbia FIVE OR TWO SUBFAMILIES? The serological data (Figs. 3 and 4) show that Ric a e the legumins from inus communis (Acalyphoi- deae), Hevea brasiliensis аиа апі Нига crepitans (Euphorbioideae), i TO of different subfamilies, are о Нига (Euphorbioideae) reference system, Ricinus commu р T taxa from o e genera Excoecaria, Homala 2. Families, taxa of which show low legumin similarity indices with a Ricinus communis (Euphorbiaceae) antiserum. бы 1 is according to Dahlgren (1980). Family Order Superorder Araliacea Araliale: Araliiflor: Berberidaceae Ranunculales ваи. Buxaceae ales Rosiflorae Casuarinaceae Casuarinales Rosiflorae Celastraceae Celastral Santaliflorae Combretaceae Myrt: Myrtiflorae Daphniphyllaceae Buxales Rosiflorae ce Fabales Fabiflorae gaceae Fagales Rosiflor: Gentianaceae Gentianales Gentianiflorae Juglandacea: Juglandale Rosiflorae ythraceae Myrtales Myrtiflor: Magnoliace: Magnoliales Magnoliiflorae Melastomataceae Myrtales Myrtiflorae Myristicace Annonales Magnoliiflorae Myrtaceae tales Myrtiflorae Onagraceae tale: Myrtiflorae Papaveraceae Papaverales Ranunculiflorae Primulaceae Primulales Primuliflorae Proteaceae Proteal Proteiflorae Rosace: Rosales osiflorae Saxifrag. e Saxifragales Rosiflorae Sc наба Lamiiflorae Solar Эа ы ЕЛ Solanales ЧЧ — BEEN Volume 81, Number 2 Jensen et al. 171 1994 Leguminlike Proteins 07 9) wis @ RICINUS 05 А Sapium HURA А Mil HEVEA @ Mercurialis a А Joannesia Mj $ 03 - О PHYLLANTHUS O Securinega O Andrachne Euphorbia Д Ng 3 04 3 7 a REUTEALIS @ Mallotus O Antidesma -0.1 T T T -0.3 -0.1 0.1 0.3 0.5 PCA | MDS | logical similarities between five CURE А & relerence oe тн апа * nonreference taxa ex Plaining 91% of variance. — b. s. For кле sie rk 5; ne S on pairwise average taxonomic PCA, two factors اا‎ (Sneath & Sokal, 1978 ) clue] ed from five variables. “Str i: Tr by the d epic Picted ihtecpoint distances, i.e., very obs fit to observed distances. the j ium, and Eupho d я of сае Meer from the thre ai e Systems mentioned is high; the г about y = x and almost identical for all Corr "c т combinations (Vogel, 1986). This is not ), а or the Reutealis reference system (Fig * another reference in Corre: ik spondence obtained among the three sub- Serological data do not support Thus the зер, aration of f the s е subfamilies, but clearly indicate м © 5 © E б e rad а & = 3 2 =» 2 © E о agreement wit sur оно who designate the two stares ا‎ Gi yuan 1980; Ehrendorfer, 1991). EROLOGICAL RELATIONSHIPS WITHIN THE AMILIES use the Oldfieldioideae have been m ally Scusa only data obtained for the Phyllan- 172 Annals of th Missouri SN Garden PCA | g | | к г TE JN XI MER rS 1.2.0] eimilarities between five " f, Ж experiments. I | FiGURE 4. di i | еше сня кайшы) апа rok reference exp к РСА, | 91% of variance. For ыл; end ie see cad 1. ч Species with high legumin similarity indices with a Ricinus communis (Euphorbiaceae) antiser ABLE 3. Classification is according to Dahlgren (1980). Species Family Superorder | Adenia pechuelii Passifloraceae Violiflorae | Passiflora caerulea — | ugonia orientalis Linaceae Rutiflorae Linum usitatissimum ux Cneorum tricoccon aceae Rutiflorae Tropaeolum majus Tropaeolace Rutiflorae Daphne gnidi Thymelaeaceae Malviflorae Antiaris africana Moraceae Malviflorae Morus rubra E Celtis occidentalis Ulmaceae Malviflorae Dictamnus albus Rutaceae Rutiflorae Casearia gladiiformis Flacourtiaceae Violiflorae усагр == Laportea moroides Urticaceae Malviflorae Parietaria officinalis um Urtica d oica E Anacardium occidentale Anacardiaceae Rutiflorae Toe verniciflua же Spondias mombin ج‎ porien eard MM а matrix for the serological results from and Lee dim этан" (b). For further explanation see Table 1. The ps iex ‘aceae”’ has been using three outgroup tax pu h. f to allow for fi Rici alignment of the table. a ТОМИ 14-15-10 37518 N © 21 22 25 28 communis огиз Celtis е Dictamnus albus Fla courtia inermis dio: ») ox 8 bd Е; 5 E 3 e х ота ац R hamnus catharticus wh © о © ЕДА s us li sages c o IET Wm I ce Me vcio I qc ote t ld pe pcs p NM шш Лат cst uu ОЕ О UL ИЕ А р ee Toa der del 3l e dg 2 ESI жы s DANCE е O4« ut О IL tub dm 4 cl EI c oap Ө rt 3 = 4 c T Ost L + — — = = + ©0 + + + - + + - + ө + - + - +++ + ө س‎ -++--++ HESTE E — = — + - а аа 5 -= = - + —-—- + + - =- — - - س‎ + + - = = - = = — + = - - -+ — —- د‎ -+ ppp ee e = aaa ————— КТ ТӨ зч = нее үш ee пешае eu M NC шш тасса ше Euphorbiaceae Passifloraceae Linaceae Cneoraceae Tropaeolaceae v661 z JOQUINN ‘Lg әшпүод че зә uəsuəf $шәўолд әзүшшпбәл [741 TABLE 4. Continued. b 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Ti 18 19 Phyllanthus calycinus 1 & | | | | | | | | | | | | | | | | | | Euphorbiaceae Linum usitatissi 2 = E + + + | | + | | | | + | | | | | | Linaceae Pimelea ligi 3 + 0 + + + Hs | | | | | | ar 25 | | | | e Pteroceltis tatarinowii 4 - + + ө O | ar | + | | | | | | | | | | Ulmaceae eltis occide 5 - + + [9] @ Е + | + E | [9] | | | | | | | Sos Passiflora caerulea 6 E E= + E f ө + ax + | | | | | | | | | | iflor F. cataphracta EL -— = + E + To. E + + z + T F | | | | | Flacourtiaceae Ficus carica 8 = + - = — t t e T T [9] [9] | | | | | | | огасеае пеогит tricoccon 9 — = a + + + + S e O ae + [9] | | | | | Спеогасеае С. pulverulentum 10 = Ed = + ES + + O ө + t zm | | | | | | Cneoraceae Daphne mezereum 11 т = = m = = + O zr 25 e o E a еј | | | | hymelaeaceae Parietaria officinalis 12 E ЕЕ э з; O + O T 6 о ө O кт UR | | O | Urticaceae tica dioica 13 - + - - - - + - + + + O ө + + + + | | Urticaceae Ptelea — 14 -= — + = — $ Q = + + + e + | | + | tace Zanthoxylum alatum 15 — + - - - - -— - - о + + + ө O | | | Rutaceae Citrus limon 16 - — -= — — - - - =- — + = о & + | | їасеае Hydnocarpus alpina l7 - — - - - — — Е — + - — Е © + | Flacourtiaceae Laportea €: 18 — > = - -— - - — - - + — -— + © + гїїсасеае Toxicodendro Md a a = E = = E = E = = = + ө nacardiaceae ciflua au} Jo sjeuuy ZL uapsed jeoiuejog HNOSSIWN Volume 81, Number 2 1994 Jensen et al. 175 Leguminlike Proteins thoideae, nies reine Crotonoideae, and Eu- phorbioideae are disc Phyllanthoideae. ا ر‎ genus: Phyllan- thus Te sted genera: E ا‎ Bridelieae, ac- 4) thus, no honia, piis (all tribe зеге theae). Only seven genera included in the cross-reaction experiments arith he eti q^ pd iunior "e tem (Fig. 1a) Bre nia, Andrachne, and S. the most similar genera to Phyllanthus pie the tested taxa, in agreement with the inclusion of these в found to be taxonomically and serologically Seit from the genera included in the Phyllanthea M iine Reference genus: Ric Tested ge Chrozophora (tribe inibi cd nera: А ). Mallotus, Mercuri s, Ricinus (all tribe calypheae), Dalechampia (tribe Plukenetieae), tribe Ade- um alea (tribe Omphaleae), Adelia (t e жеўи, {зш їп the sequence Ricinus -Chrozophora net — 5€ the three genera should be phyloge- connected to non-Acalyphoideae subfam- ал rough Ricinus. ш ER although ed in the tribe Aca- Sri emonstrates only a distant Y rhon ip я A Dale ae ire Pluken eae) has nt legumin, whic with the Reutealis (Cro- Fig. 4). ) reference ee m (Fig. ideae. Reference genera: Hevea, Reu- Crotono tealis, ested genera: Hevea (tribe Micrandreae), Tes M Sy Meme f anihot (tribe Manihoteae), Jatropha, Joannesia Passiflor ас Violiflorae logical similarities between the Eu- Metus (E) pum utgroup families or orders of the Rutiflorae, Malviflorae, i Vio liflo rae. ари Да index fro г (both tribe a Strophioblachia (tribe Codieae), A leurites, Reutealis, Vernicia (all tribe Aleuritideae), Croton, Eremocarpus (tribe Croto- neae) e relationships of taxa included in the Cro- tonoidene were este with wo Herne v— 1 пеу for this subfamily. From the comparison with the c, Fig. 1c), the high serological correspondence es € (tribe Manihoteae) and—although of less d —Croton (tribe т сап Бе interpreted as supporting close relatio оир. Bow- ‘da en » interpreting the high serological correspon- xa from other subfamilies, i.e., Rici- calyphoi Te sm and dh ra (both Euphorbioideae) The close relationship of Jatro- pha and er d was not confirmed; Jatropha is loosely connected t e и онн. whereas Joannesia is con vea, Rici- nus, са Hura (Table 1c; Figs. lc, е Strophiob- h Codieae b) was emp! ploy ed in t the expor- nus (À iments with the Phy y is close to dominera, Aleurites, Reutealis, and } П те bers of the tribe Aleuritideae subtribe Aleuritinae. They produce a similar legumin, which establishes ret el relationship. The significance of this 176 Annals of the Missouri Botanical Garden atement is supported by the prominent cross- sin е m (Fig. her ref- eronen «уе; Be distant spe of Hevea in of considerable phyldiinctia differences iecur the Aleuritideae and the Hev groups Euphorbioideae. Reference hart Hura. Tested genera: E ea—Hura-Ricinus о ‚ Homalanthus, Sapium (all tribe у Нига (tribe Hu- i rbie ally in e of ona lanthus, to i Cr ploneideas * aea mod to either th e Pare ues reference e er refe em ея ге- , which is s солы with morphological characteristics. strate ee d position of "Eüphas bia RELATIONSHIPS BETWEEN EUPHORBIACEAE AND OTHER ANGIOSPERMS onomic and phylog: ions on reli iotaipo of the Euphorbiaceae within t osperm s (see Webster, 1 Corner, 1976; Croizat, 1973; Dahlgren, 1983; Huber, 1985; Hutchinson, 1926; Lindl E836, 1853; Mueller, 1866; P. 1924; Takh- per E TON serological an ne ae aie char- Violiflorae, or Rutiforae, this does not apply to all г? ха v the orders equally. alviflorae. “Dahi hlgren (1980, 1983), Dahlgren et : (1981), Takhtajan (1980), Thorne (1976, 1981), and many other authors regarded Euphor- Ф es = eo un — c = E. 5 # = © = През кейт the Malvales nor the Выс have been found to be krea related to the Euphorbiace as between Euphorbiaceae and Rhamnales, are un- ike In contrast to the orders previously discussed, the Urticales (Moraceae, Urticaceae, Ulmaceae) to hypothesize close relationships between these two taxonomic groups. 2 The шше similarities betwee Euph between Euphorbiaceae and Thymelaeaceae: Tos phorbolic esters are produced in these tw o fa (Frohne & Jensen, 1992). A crotonoid pollen exine ; with a palisade layer ) oa гер, Phyllanthus and ly Pe Howev mbers of the Violales, ene y t this context the Flacourtiaceae have p ed e scil of interest because this family is regar Volume 81, Number 2 1994 Jensen et al. Leguminlike Proteins pecially closely connected to the phylogenetic or- Vi igin of the Violales. Passifl ). A valvate calyx, A ES E ре. Ыт ite and an ex- pry cr ties of the Passifloraceae, and these characters in ad- dition to the seed protein data reinforce the theory of a close EC between Violales and Eu- phorbiace Rutiflorae. Many Rutiflorae species included in our serological experiments revealed important same range as the cross- re Re many ырший ыа. is merge in contrast to the ic. (Geraniales), failed significenfly t ш cra Jes). The following three Rutiflorae families vealed especially high legumin correspondence with the Euphorbiaceae: Tropaeolaceae, Cneora- ceae, Linacea varies of the part into as do t upon mandi a three Басе They also ion it is ilte pr n Ке пеогасеае and Euphorbiaceae; however, тм Characters have not been utilized in form classifications. Na min si Phyllanthus) nacen ы аз M E йыш, cond placement (reference: Ri- ч the families tested, an ve ized when the relationships о Euphor- characters ттеу ane ae "йен Lr. er c aracters indicate nificant ween the ee and Euphorbi- ac eae that cannot be exp lained by convergence alone. THE Linaceae seeds ж жаа ЧОРУ апу 1 a hypo gynous с one or two bitegmic sro for each carpel, an obturator, intra- or extrastami- nal floral disks, and stipulate leaves ARE THE НИВ AND ROSIDAE SUBCLASSES NATURAL ur discussion of the relationships of the Eu- y асошцасеве ant iaceae are к"; ае), which а (e.g., Cronquis fore, the separation of the Dilleniidae ae Boadas subclasses hes d be toned and a taxonomic is con E sidered necessary. Within th is large collection of orders in the “Mittelbau” (Ehrendor dent i ) of the angiosperms, there is a group s that show sine аг char acters, Len ipie Tares re- 1 al disks ith cases per с eai bit tegm c and c аа А ea with obturator, нов, and cyanogenic and glucosinolate compoun this taxonomic complex, the a aa are included. LITERATURE CITED Etude Générale du sm des Eu- : 185- 1 ‘Euphorbiaceae. In: G. Bentham & J. р. Hoo ker (e og Ge 3: 239-340. BERG, C. C. А Urticales, their differentiation and наза) position. Pl. Syst. Evol., Suppl. 1: 349- BERCN ER, I. & U. JENSEN. 1989. Phytoserological con- tribution to the rapis. adus of the Typhales. ordic J. Bot. 8: 447-4 Corner, Е. J. H. 1916, The dub of Dicotyledons. 2 idge Univ. Press 0. Interpretation and analysis of sero nee data. Pp. 269-288 in F. A. isby, J. С. Vau n& D. . Wr ight (editors) отоу с Press, London Кг New э Croizat, L. 1973. Les Euphorbiacées vues en elles- 178 Annals of the Missouri Botanical Garden mémes, et dans leurs rapports envers l'angiospermie énéral. Mem . Soc. Brot. 23: 5-206. ае. А. 198 п Integrated System of Classi- =ч tion of пяе Plants. Columbia Univ. Press, ork. реа Re ah A revised syst f cl of the angiosperms. Bot. J. Linn. Soc. 80: 91 1983. General aspects of angiosperm evolu- di and macrosystematics. Nordic J. Bot. 3: 119- 149. ‚ S. К. JENSEN & В. J. NIELSEN. 1981. A revised classification of ie сестее with c ents on "wel T between chemical and other characters. - Young & D. S. Seigler е 199 ; УВ AA pies and Angiosperm Phylogen Praeger, New York. E т 1991. Samenpflanzen. Pp. 699-8 itte, H. Ziegler, F. Ehrendorfer & A. чай Ой ЫЫ buch der Botanik fiir Hochschulen. С. Fischer Verlag, e New York. FAIRBROTHERS, D. E. 1977. Chemosystematics. Bull. Serol. Mus 52: 3. Ev iden nce from nucleic acid and protein chemistry, i in Tear serology, in angiosperm clas- c J. E 3: 35-41. sification. Nor D ju BEL SG B. L. Tur The bases of angiosperm оше: ае . Missouri Bot. Gard. 62: motaxonomy. Ann 800. ‚ Н. & U. JENSEN. 1992. Utilization of i un to е و‎ in plants: Serology. A dis- cussion based on eraceae— Cichorioideae. Belg. ih um 125: 243-2 Froune, D. & U. JENSEN. 1992. Systematik des Pflan vibe: 4. Aufl. G. Fischer Verlag, Stuttgart, Né York. HUBER, un ae Angiospermen. Leitfaden durch die d Familien der Bedecktsamer. Gustav Fischer Verlag, Stuttgart, New York. HurcHIN isi ; ae و ا‎ of Flowering Plants 8 Dico die Macm d 1969. Grobe us Phylogeny of Flowering Plan nts. Academic Press, London. JENsEN, U. 1984. Legumin- like and vicilin-like pea proteins in Nigella damasce. ena (Ranunculaceae) an six other J. Pl. Physiol. AUS Steps toward the natural Miri of E dicotyledons: Serological characters. Aliso 13: 18 ———— & Н. BER Legumin-like proteins in E. "Phyocheniaty 28: 1389-1394, —— . BÜTTN distribution of storage proteins in Ме olio ophytina reg pe) and their serological similarities. Taxon 3 9: E FAIRBROTHERS (editors). 1983. Pro- Nucleic 1 Spring. er- T Berlin, СЯ p & B. GREVEN. 1 1 a phylogenetic relati р Magnoliidae. Taxon B - 1983. Seed storage proteins. Pp. 238-254 in U. Jensen & D prit (editors), P. ucleic Acide in Plan tematics. rep Co in Berlin, Heide no KoLBE, К.Р. 1978. Serologischer "Beitrag zur Syste- morik der Capparales. Bot. Jahrb. Syst. 99: 468- 489. ———— & J. Jonn. 1979. Serologische Untersuchun- gen zur e re der Violales. Bot. Jahrb. Syst. Lem R RE. A. К & CL r cation ion techniques. Pp. 275-300 in U. Jensen & р. Е! Fairbrothers (editors), ant Systematics. LINDLEY, J. 1836. A Natural cui of Botan y. London. . 1853. The ee table Kingdom, 3rd ed. Brad- Evans, Londo : d Euphorbiaceae In: A. р: del ae itor), Prodromus Sy 3 — Paris Pax, F. orbia In: A. Engler & ran Га Die Narirlichen Pte ok te Auflage, III. 3: 1-119. W. Engelmann Fae Tee bee кү der Euphorbiaceae. Bot. Jahrb. i "59: dei 5% Rizk, A.-F. М. 1987. The al constituents anf economic Cu of Es рни, m 293-32 in S. E; . F. Cutler & F. J. ken (edi: npa Bat ичиле лы Chemistry, Hed y and Economic Botany. Academic Press, R ;] 13 : ‚1989. „МТЗҮЅ- с; Numerical Taxonomy "e " i i S t Exeter Publishing, Жы New t di ScHULTEs, R. E. 1987. A new generic concept in . Leafl. 17: 27-36. О Я .& Е. Soka. 1973 Numerical Taxonom A aN Du iem . Flow rig Plast is Evolution le species Level. —— ap Press, Harv: 9 Cambridge, Massachus TAKHTAJAN, A. 1969. ы Гара Origin and ip 4A ral Oliver & Boyd, Edinbur; ae 1980. Outline of the Чаш of w plants (Magnoliophyta). Bot. Rev. Lancaster 46: M Magnoliophytorum. Nauka, 1968. "Syn opsis of a be р net : аа абса of the flowering plants. A phylogenetic Mines of the ks pior е Angiospermae. Evol. Biol. 9: 1981. Ph dopnas i and a ngospo in D logeny Ks summary s nt. Pp. 233- ud A. Young & D. S. Seigler (oir) Phytoc val k. and Angiosperm Phylogeny. Praeger, New hea oposed наф realignments m giosperms. Nordic LE Bot. 3: 85-117. VocEL, CH. 1986 Aes Untersuchung e. Dissertationes zur Syste matik der Euphor À tanicae, 98. J. Cramer, , Stuttgart. й WEBSTER, С. 1. 1967. The кеч of Euphorbia in the south-eastern United Sta . Arnold 48: 303 of nspectus of a new classification the E Taxon 24: 593- Mr review 9f 1987. The saga of the е pue horbiales- classification and relationships in the uP’ Volume 81, Number 2 1994 Jense al. Gao Proteins 179 Pp. 3-46 in S. L. Jury, T. Reynolds, D. F. Cutler & F. J. Evans iiis "The Euphorbiales: Chemistry, Taxonomy and Economic Botany. Academic Press, London. 1994. Synopsis of the genera and suprage- neric ic taxa a EE Ann. Missouri Bot. Gard. 81: 33-1 т, R. dee Handbuch der Systematischen Ж nik. Vierte umgearbeitete Auflage. Deuticke, s ig. WUNDERLICH, R. 1968. Some remark: nomic significance of the seed coat. Debe cs 17: 310-311. APPENDIX. List of species used, with their sources. Euphorbiaceae ane ricinella L., seed stock Lehrstuhl Jensen Aleurites montana (Lour.) Wils., Bot. Gard. Sri Lanka 1985 Andrachne hiss Fisch. et Mey. ex. Boiss., Bot. Gard. n 19 ne ed (L.) Spreng., Bot. Gard. Salisburg Baccaurea tetrandra Muell. d. Basel Br Оша fruticosa (Muell. Arg. ) Hook., seed stock Lehr- ensen Bes tomentosa Bl., Bot. Gard. Wien eis zophora Ns (L. ) А. Juss., Bot. Gard. Gót- Cron tiglium L., Bot. Gard. Gießen 1 lechampia s candens , Bot. Gard. e Drypetes pira asien Pax & Hoffm., seed stock о И Jens Ere (Hook.) Benth., Bot. Gard. Ho- nol Euphor lathyris L., Bot. Ga ни йоны, Lo bicolor Hassk., rew d. Base » ochidion glaucum Muell. A oed Basel evea Biens (Willd. ex s nu Muell. Arg., xt Gard. Sri Lanka 1985 omalanthus populifolius Grah., Bot. Gard. Wien Hura repitans L., atropha Ja си Bot. Gard. Sri Lanka 1985 rcas ER Bot. Gard. Bogor 1984 AN 4 princeps Vell., Cutt. Lyon Arboretum, Ho- Mallotus phdippinensis (Lam.) Muell. Arg., Bot. Gard. Moir ese culenta Crantz, Bot. Gard. Düsseldorf Tgaritaria discoidea (Baill.) Webster, Bot. Gard. E je ercurialis perennis L., Bot. Gard. Basel tie ericoides Desf., ا‎ stock Lehrstuhl Omphalea trichotoma Muell. Arg., seed stock Lehr- р stuhl Jense aine quas quadriloculare F. Muell., seed stock Mos c yee. Labill., 1 Reut ае roxburghii Wall., Bot. Gard. Basel 1 (RI \ w. Bot Gar d. Bo- gor 198 Rev - none arenaria A. Gray, coll. Webster Texas ac communis L., Bot. Gard. Sri Lanka 1985 Pium sebiferum (L.) Roxb., Bot. Gard. Siena 1983 Securinega durissima J. F. Gmel., Bot. Gard. Stutt- gart-Hohenheim Strophioblachia fimbricalyx Boerl., seed stock Lehr- stuhl Jensen Vernicia montana Lour., seed stock Lehrstuhl Jensen Anacardiaceae Anacardium occidentale L., Bot. Gard. Basel Spondias mombin L., Bot. Gard. Wien 1984 Toxic ie € verniciflua (Stokes) Barkl., Bot. Gard. Got Cneoraceae Cneorum tricoccon L., Bot. Gard. GieBen Flacourtiaceae Casearia gladiiformis Mast., seed stock Lehrstuhl Jen- sen Flacourtia inermis Roxb., Bot. Gard. Basel 1978 Idesia polycarpa Maxim., Bot. Gard. Wien 1988 Linaceae Hugonia orientalis Engl., seed stock Lehrstuhl Jensen Linum usitatissimum L., Reformhaus Sattran, Bay- reuth Moraceae a Engl., seed stock Lehrstuhl чесен Antiaris africana Lavall., Cudrania iare qi (Carr.) Bur. ex ard. Wie OF 2 rubra ка -— Gard. Wien 1988 Passifloraceae Adenia pechuelii (Engl.) Harms, seed stock Lehrstuhl ensen Passiflora caerulea L., Terra, Rosdorf Rutaceae Dictamnus albus L., seed гей — — Phellodendron japonicum Max t. Gard. Basel Ruta graveolens L., seed Беа Lehrsuhl Jensen AN simulans Hance, seed stock Lehrstuhl Jen: Thymelaeaceae Daphne gnidium seed stock Lehrstuhl Jensen Da phne mezereum ты pa st tock Lehrstuhl Jensen Tropaeolaceae Tropaeolum majus L., priv. Garden 1989 Ulmace Celtis occidentalis stock Lehrstuhl Jensen Ulmus parvifolia Mis “hot Gard. Siena 1984 элө ча са —€— (Pall.) К. Koch, seed stock Lehr- hl Jen Urticacea Laportea moroides Wedd., Bot. Gard. Gottingen 1989 Parietaria officinalis L., Bot. Gar d. Wien 1988 Urtica dioica L., Bot. Card. ar ‘Dahlem 1984 SYSTEMATIC ANATOMY OF W. John Hayden? EUPHORBIACEA BF UBFAMILY OLDFIELDIOIDEAE. . OVERVIEW! ABSTRACT characterized ur spiny polle therwise apparent livers pees or of ГА зат Hemisphere trees and shrubs that traditiona ally tive been allied with genera x and Hoffmann. Altho: A "fairly ibi anatomically, the following structures characterize the subfamily with o nud few exceptions: pinnate brochidodromous venation with generally randomly organized tertiary and pie her ped venation; eme and petiolar glands absent; unice e ie uniseriate trichomes; latex absent; mucilaginous epidermis or hypodermis; brachyparacytic с stomata; 5208 d i with s simple perforation plates iai alternate, often very s ‘small, intervascular pits; thic k-walled nonseptate ИШ "t banded patterns and Ше heating prismatic crystals, Anatomically, the sl shrubby Australian ericoid er [T : kia еше рр ий Aus tralasian genera, "sine cd кш илт up somewhat more isolated and seem to represent E in various cases, elements that are primi erh ani Mee i Mint or xvii derived from the group bearing compound leaves. Presence of хобе a and р compound leaves in Oldfieldioideae are features consistent with Dilleniid origin for Euphorbiacea: d | xonomic entity the euphorbiaceous sub- dendron was shown to be е А de Nu Oldfieldioideae Kahle er & A 55 ter _ Sates deae (Hayden, 1977; K к onceptually and Hakki, 1985), an айа SU ney o (1962) and Kohler (1965) whe poted the spiny entire subfamily was initiated d 1 a i m sess relationships from data pende x recogniz ed by pollen structure was subsequently productive (including pollen) morphology. Ee formalized pesce ally as a new subfamily part on these anatomical studies, йаз ШШ (Webster, 1967). In essence, pollen characters Oldfieldioideae has been bare fro e dii absque: a group of genera thai previously (1975) original proposal; two broadly similar © ilies Phyl- — sifications, one by G. L. Webster (the prec lanthoideae dnd Porantheroideae in the system of issue) and another by С. A. Levin and © Pax & Hoffmann (1931). Webster's first (1975) Simpson (this issue), are proposed elsewhere. classification of oldfieldioid genera contained sev- ovel ta associations at variance wi earlier classifications, notably those of Pax & Hoff- nonu the first mann (1931) and Hutchinson (1 ). rane of Euphorbiaceae кишш pa Interest in the anatomy of Oldfieldioideae half of the 19th century (e.g., Ju ussieu, 1f to reli- stemmed from the need for comparative data to licher, Rt offer little insight for- aid po S Me problematic genus Picroden- eei among bi eee genera. А! үт zi dro ‚ 1977). Despite a number ative time in the ition of Euphor known of AR dealing a ith the anatomy of old- (Webster, TI үй is Puts genera more- PEE к ra (Appendix 2), detailed oer often s attered widely the family: ' Genera t of the mnis is lacking. Once Picro- over, son ref ther families k. I ' This study is based largely on a Ph.D. dissertation submitted to the University of pigri: College Jie thank W. L. Stern for assistance and Sali H. К. Airy Shaw, L. J. Hickey, С. A. Levin A le provided and G. L. Webster were particularly specimens. * Department of Biology, University of Richmond, Richmond, Virginia 23173, U.S.A. ANN. Missouni Bor. GARD. 81: 180-202. 1994. ime ———— —— ӘӘ ——— — ө ёӨ ө ЧЁҤ[Ҥ[Ёү-ҥҥ-}.И Ю Ү ?? 9 ўўттЕ — ————— Ne ЧЫРЫНАН Volume 81, Number 2 1994 Hayden Euphorbiaceae Subfamily Oldfieldioideae of ини a Caletieae in the traditional/ h classified to- D which prb 1859) added йеп Plan- s followed in all subeeqeebs ngu until the а first, of Neoroepera Mue by Kóh- ler (1965) and, now, the inclusion E al Austral- asian oldfieldioids as proposed by Le 1 o ар = E N [7] Dn = i) LL e [= [8] Е" Ф e c = ~ = $5 Ke) A = He =| ч Б ge c > о г e primary criterion for ivision of the ا‎ (Mueller, 1866; 890; Hofmann, 1935 iie the genera SEE EM 1 mall cot- group of аап ene obe with narrow vedo ns. Their somewhat isolated position in Sten- inimi 1 # i гр, 1 74 Ко ith other oldfieldioid genera Other ol fie ldioid id similarly suffered early at у casside s relationships with tl Bail- on (1858) submerged Podocalyx Klotzsch within i "bs MM genus Richeria Vahl, a disposition lfieldia Ben h. ries of misa ents at both generic and ep levels (Hayden et al., 1984) al e sensu stricto, the genera m Caletiea ^ oom cun were scattered widely in Muel- ег' (1866) treatment of the family for the Prod- ch h included with ч чч : Choric 1 as in- within Wee —— Miq. (as Bur- Pru Baill.), Z getia Baillon ex Muell. Arg Stigma Е. Mel „апа Hyaenanche Lambert on and issiliaria F. Muell. ex байар е повна (which included the old- d оаоса lyx). (See Webster (1987) for com- n's peculiar *serial" system esed accreted to this loo d nucleus bon pe. dioid genera. e (1880), i indus neat: Euphorbiaceae in the Genera Plantarum, included, using current nomenclature, Dissiliaria, Longetia, Austrobuxus, Hyaenanche, Mischo- don, Oldfield, Piranhea Baillon, and only one Bischofi a Blume. within gen- era 46-54. Although these plants constituted b Bentham (1878) Bionic some doubt about their relatedness and he with other phylla M мавод genera much as h Tetracoccus Engelm; є x C. Кашу, pages: was included ta- pi dade Prain, and e a Prain to various subtribes consisting — of oldfieldioid genera. * Per haps the first clear indica- relat edness. d oldfieldioid ا‎ can =: Although Fat of the detail in this be fieldioid genera (as usual, minus lade distinct from other biovulate ) and bend каше ken of Pa ax & Hoffmann 69 ега. Ae light tof Pax’s (1925) phylogeny and because of its disregard for the palynological data av ailable the time at (Punt, 1962; Köhler, 1965), Hutch- inso 69) classification was a retrograde de- velopment; oldfieldioid genera were widely distrib- uted in five of 12 biovulate tribes, with three tribes er ldfieldioid and g^ iain E em ts inson was the first, how Hut Pardee Kuhlm. with Mer oldfeldioid £ gen- е i (1965) entertained serious doubts about inclusion of several oldfieldioid genera in drin enm Accordingly, he proposed Andro- stachydaceae, of uncert rtain relationships, to accom- modate Androstachys; he r recognize we Pe "ee А and he viewed Aristogeitonia, Calaenodendron Sey. “Mischodon, Qidfieldia a, and, Pitgahes s (Airy Shaw, 1966, 1972, манаа ни; Shaw dn continued to accept an iso onomic position for Stenolobeae in pala anthinae), but he did, consistently, group the re- mainin E Australasian aure of Oldfieldioideae 1 in classification systems (e.g., Airy Shaw, 1975, 1980b, 1 83). Annals of the Missouri Botanical Garden WIELAN ANDRACHNINAE ANTIDESMINAE AMANOINAE DISCOCARPINAE UAPACINAE BISCHOFFIINAE PHYLLANTHINAE GLOCHIDIINAE DIINAE PSEUDOL ACHNOSTYLIDINAE PETALOSTIGMATINAE Petalostigma DRYPETINAE oldfieldioid taxa in bold italics. Other th desminae]) is ine an element of Ol indic an Pseudant r^ Podocalys (treated an a mapa eee Drypetes Heywoodi Lingelsheimia Neoroepera Tetracoccus PAIVAEUSINAE SAUROPODINAE Aristogeitonia Oldfie dia TOXICO DENDRINAE Androstachys Hyaenan DISSILIARIINAE FICHA: Longe (ле. "Austrobuxus) Mischodon f Phyllantheae, after m Aus Right-hand clade is the earliest p PPAR ae Olli e Gener ic composition of subtribes, ublished & Hoffmann (1931} ated, follows Pax at Riche Anti- DESCRIPTION, CIRCUMSCRIPTION, AND DISTRIBUTION Plants of Oldfieldioideae are Ират ie low depressed shrubs to tall tree d tri — Leaves are atoma e, opposite, or "nene d лаў ъс эп. пр le Ul „р апа маб зні ave кн stipules may be present or absent. Leave: st 5 n ч دم‎ 3 ga @ m © В e ree to many. Pollen is bi- — with {ш to — brevicolporate to po- 1 be ars yer, a discontinuous interstitium and thick perforate tectum with prominent е spines (Levin & Simpson, 1994, this volume). Gynoecia consist styles are gene erally p a pair of pen ue of two to five carpels; vided; each locule houses a ropous ovules. Seeds are Pur AX bg s copiou osp - r Pseudan e bens 1976). Ae bi We Pieradendron (Fritsch, d 73), and 8 Mischodon a We. ры н, = 24, 2п = тн Misokodon did 1980) cci suggest a base number 12 for cu sub sive a of en ded 21 £ The HORAN eme (Webster, 1975) in malo Capuron ex Bosser (19 Airy Shaw & B. Hyland ii he —— ———————"" Volume 81, Number 2 1994 Hayden Euphorbiaceae Subfamily Oldfieldioideae been referred to the subfamily. ations listed in Appendix 2 has been pee era, Croizatia e "(Weber et ue 1987 ) an » Paradrypetes (Levin, 1992), have been as- gned to Oldfieldioideae following ш of di- pi flowering material and, especially, е " udy of pollen from these plants. Kairotham QI X 1Q MaD} (1085) (1090 ` © E rn f 4 $ 1 as has Сапаса Guillaumin, if only пан (Webster, 1994). In recent years Radcliffe-Smith has redefined generic limits for several Oldfieldioi- dea e. He has included Para nium Léandri with- in Aristogeitonia ате, 1988), he seg- ih préchyandra A Sei rom онаси ing compound pu (d desine 1990), and he expanded Pseudanthus to include Sta (уне mon (Radcliffe- Smith. 1993). mber of gen era now ш at = ы because of me re- treated as a distinct genu — a of Oldfieldicidene are mostly mono- or sii Austrobuxus, with perhaps 20 species, risus (sev- pecies dica E is largely a southern евге u m feet the breakup of Gondwanaland; "е, are Ce- й enodendron, from western Mexico, roden- ri ren fom fhe Bahamas aod | Greater niles, эй i ce genera and 16 s pecies occur in New s are Australasia genus of UTE Aut a bicontinental yia ution, gh gh Androstachys and s PN occur in Madagascar and Afric MATERIALS of a | port is based on first-hand examination eaf and 87 wood specimens of 61 species f hà hes dfieldioideae. Specimens examined are listed ü ndix 1. The specimens available represent of the subfamily except nus, З pe and Stachystemon are lacking in this Fais ous anatomical literature concerning a of Oldfieldioideae provided by the publi- into the following account as appropriat LEAF ARCHITECTURE st part, leaves are > either simple or nsi es occur in 4 Parodiodendron Hunz. аге 1mifolioste, as dg de (H unziker, dandi and by ды frequent disarticu- min: e point (Fig. 4). eaf which, by loss of their common p sess зве | on the stem (Вайоп, 1858); alternatively, x 1 (Gri üning g, 1913; ; Webster & Miler, I a he few he ана Eon Е з = = leri (Baill.) ex Kuhlm. (Levin, 1986), P. subintegrifolia G. Levi (Levin, 1992), Tetracoccus dioicus Parry зш some — ps T. ilicifolius Coville a on er ул eeth (Levin, ‘1986, 1992). Teeth i of 1 Тағасоссиз media paced lv: ilicifolius veins of these teeth are derived from d» E secondary veins; there are prominent brochidod- PEL G d di romouslike | Р veins extend nearly to the apices, which in the | cans specimens examin possess at least in young leaves. Teeth of the ‘remaining species are much smaller, consisting of a vein end- a small protrusion n of leaf tissue. зе, ing in however deciduous apical cape are also v visible (Figs 7, 8). Tooth morphology in риает | hus theoid type (Hi ckey & W always p tooned brochidodromous (Figs. 2, to brochidodromy, however, leaves of Longetia 184 Annals of the Missouri Botanical Garden | \ i i A E | \ E] i um Fic 2-8. Leaf architectural энше of cene, —2. Austrobuxus rubiginosus (Baumann: Boden“ T 15010), gon ared | аг = 1 Celaenodendron mexicanum (Ortega 6367), cleared leaf, bar = 1 © . 4. е таи ран 469), ve leaf, note disarticulation of petiole from bas = 25 . Picrodendron baccatum (Gillis 6963), note lack of vein orders beyond tertiaries, bar ba Volume 81, Number 2 1994 Hayden 185 Euphorbiaceae Subfamily Oldfieldioideae ibit a tenden 1 exhi ey wh Aereas those of Dissiliaria, BRENT рмен end toward reticulodromy. Leaves Picrodendron (Fig. 5), and Piranhea (all New d gene Wor pice this c WAY state (T J H k us of the Australian genera of Penis к in the dicots a ( pers tieae sensu strict ibit a is tl ially rol ро! š х ГА г кА. ing disorganization, pre ly a consequence of ing these gene era. Areole ll developed in the tk іп opera leaves are weakly fes- — genera with com des ives (Figs. 5, 12) plus t brochidrodromous; in Micrantheum one can — Dissiliaria, Mischo! n, Parodiodendron (Fig. 6), find Focus of oer Smit узын аз ne p dens as seconda ana there, areoles are coa ually arranged rand web и. ki Intersecondary veins are usually absent, al- though one or two simple inte ee. per in- tercostal area ш: сазда ustrobuxus and Mischodon 1986), Old E ee amely, Amanoa ubl., па Leandri, odia Sim, Petalodiscus Baill 5 ia Willd., a L ); their presence о be primitive for Oldfieldio iae ally random reticu- n- ep ve- à massive intram Pse marginal vein a (Fig. ае А ронан furcate -fla- nis n Hyae- orm of mar he. goa wae > venation varies considerably within currence д h е. Perhaps most notable is the oc- 2 ultimate reticula in which vein order tinguishable, a feature of Celaenodendron, a 6. Parodiodendron ing tenes (Hueck 469), num. v T leaf, bar = 250 um cleared Po. bar andit à пиц, tri, icorne pula ап s.n .), toot! Paradermal section thr ough tooth re quar ds note Au) cmt E in Picrodendron, Piranhea, and Dissilia- vanlar вз usually present and mostly pple еШ ing to be most highly branched i in leaves with low ades of Elit т yas т ane veins. g. 12), but they may appear to be present u upon superfici cial examination M eus of the em sclereids lo- cated in each areole (see belo LEAF ANATOMY Trichomes have been observed in about two- thirds of the genera of Oldfieldioideae. Many spe- cies are glabrescent with аре, а although mature leaves of Androstachys, P ы. гоп, апа aaan of petalo: stigma, fot: се аге реу Hula scent. Trichom or ы ыы: cub consisting el cmd € six ithin Pseu a nthinae, nap emer- ce ell, 1 h ed by Alvin (1987) to func- spheric moisture (mists y arid en- tion in of atmos and drizzles) in the otherwise extr ете vironment of southern Africa trichomes similar to t those of Androstachys, ar- a. Viaj i o genera despite = manifest — in . Dependin the spe- en trich or bifurcate с unique within the ubfamily in pos 69 strongly ciliate leaf margin: s (Fig. . Rao & Raju (1985) ported unise seriate, stellate, d glandular hairs in айан characteristics reminiscent of uniovu- — 7. Choriceras dee lin ies tricorne (Forman s.n.), ve d - um. 186 Annals of the Missouri Botanical Garden FIGURES 9-12. Leaf archit f Oldfieldioid ) кыд venation, massive primary and intramarginal veins, bar = 250 um.—10. Celaenodendron mexican us hi 6367), fimbrial vein (far right), bar = 250 um.— 11. Petalostigma banksii (Perry 1981), branched vr ory e Bee PU 1 2220 TEL 17 (^L : I Researc y аваны 9. Pseudanthus orientalis (Clemens 44092 , bar = 100 um.—12 J s.n.), well-developed areoles with columnar sclereids, veinlets absent, bar = 1 mm. late euphorbs tł ld be uniq Oldfieldioid cell or chamber thus formed contains m ae; however, neither trichomes nor their bases were which is common in other genera a well. pow detected in the leaves available to me. of the ilagi in Australian Figs The epidermis is fundamentally uniseriate. How- ^ thinae greatly distends the adaxial epidermis ( ous ever, a single | hypodermis has been re- 18, 19); e the grossly expanded mu s it is 0 r) ayer may well be an artifact of preparation or all epidermal cells of the Australasian genera derived solely from the lower portions of s ee à have a horizontal partition resulting in a poorly to epidermal cells and is not thickly multicell кн well-defined hypodermis (Figs. 17-19). The inner depicted by Gaucher (1902), whose nr Volume 81, Number 2 1994 Hayden Euphorbiaceae Subfamily Oldfieldioideae Fı URES mate from 6. Leaf араш or oa —13 Parade cross section, ре 4. Scage 10 ш зра ^ ; a note c mulate Pic e fro; a 4850), s ихиз s huerlimannii (McKee 20 ш m paradermal section, bar = . Lo рене: b а oligostem n (Mc Kee —16. Podocalyx loranthoides d denheim 5605), sto- { stomate from bar — = = & : ubdivided crenulate subsidiary cells e base (Krukoff 811), vache idioblasts nem macérated mesophyll, bar = 20 um. СС. = guard cell, sc — subsidiary cell, TB — trichom ter, : ане ауа has been reiterated by others A "it е & Chalk, 1950; Raju & Rao, 1977). S lonia, ане е has been du ted in Aristogei- Podo calyx aenodendron, Mischodon, Piranhea, and several CÓ M HK mis is k BL, An bac. “agents for exa Caur, ma ea Lour. ‚ Bridelia Willd., mo М AL p Phyllanthus L., Richeria, and Savia (Met- e& Chalk, 1950). In surface view, едм па! 14, 15 >), t the only exceptions being Aristgeitoni, ee че Piranhea, (all members of Pi- endreae), and Dissiliaria, in w walls wich these , par Jermal se sec- ^B = B. du 9 di ауе a 2; tions of . Ari cui and C 'elaenode seni reveal 188 Annals ыл „ш Garden Je 5 dead, ] soe E we х ш FIGURES Ay EAS Bauman" reais anatomy and wood parenchyma of Oldfieldioideae. me I жой buxoides ( үи” odenheim 5 eaf 100, —18. Pseu e and am н ode orientalis (W is e 9. Mif o cross section, note grossly expanded mucilaginous тшне marginal veins, bar = 100 um.—19. Micrantheum uper ego MA ina vray 3196), cross sectio 5 й i7 л note grossly е apenas vcio Шуро rmis, bar = um.— pnr, globosa (Go " © ке mbered cross section of leaf, bar 100 um.— 21. Pir anhea ‘pli Duete s 5. p a 1485), norma! a сту gnis axial xylem parenchyma cells, bar — m. PF — EEE ————————X— Sa Volume 81, Number 2 1994 Hayden Euphorbiaceae Subfamily Oldfieldioideae inner portions of epidermal cells to possess ingly hypostomatic, al- nditi Мару, с pa For mites intrud end to form an anomocytic pattern in E. as viewed оа the surface; pad is peotions, howe ever, reveal typical e most frequent modification ё stomat tal t type is the tendency for f the subsidiary cells to become sub- of subsidiary cells ma some stomata or onl bsidiary cell of a given pair. Subdivi орат does occur in Androst eh where the patte s bee h n de- scribed as ponds biles — 1 987), il E some species of А 15), Hyaenanche, Misc Pseudanthus. The P рагаруйе stomata of Por adr ry- 1986) The distribution of subdivided «пне, ы ithin the асалу. Within Phyllanthoideae, rachyparacytic stomata are characteristic of Bri- delieae, Drypeteae, Ph certain genera of Wielandieae; r, subdi- vided b, achyparacytic stomata o Lachn tylis Turcz. and Bridel (Levin, 1986). Raju & most common type among woody genera nera subsidiary cells extend partially periclinal walls of adjacent guard : ai EIN fashion fe. 13 Sk t the it hie extension of the бс pore. Within ( n tial hylan ideae, Levin (1986) noted a similar spa- relationship of guard and subsidiary cells only rype es. The inner anticlinal walls of “pig ok subsidiary cells may have ан out- ne 14, 15), a feature first observed hus eod (1908) in Micrantheum joris hides w pro characteristic of the Aus- genera (Misch d cells mily. Crenul Mis аге only weakly developed in the single pe n of Whyanbeelia avaiable for study. In con trast to most genera, subsidiary and guard cells are coplak in ала ia snd e ron: Mesop hyll i is bilateral Petalostigma with ا‎ leaves; ү of LI. L 2 a ‚Ж: 1 3545 (Е; 20). Spongy mesophyll with well-developed inter- теп ds cells occurs in a als Oldfieldia, м апа Piranhea, all members of Pi- j yll usually contains scattered druses; prismatic crystals are also present in Ar- these genera usually co see with ramified tips ae run parallel with unique mesophyll feature is found in Podocalyx, which bears unbran €: sinuous iraebeoid a blasts with spirally th 16). The ai و‎ "occurring most nent are re sufficiently abundant to obscure the | селе le ; they are not, however, ow ye curvature (Figs. 18, 19). P the ary v in Petalostigma and due Australian ьн harp contrast to ssociate: parenchymatous bun (Fig. 12. ‘Bunde sheath extensions that are merely cur in Longetia (Fig. 17); crys- рагепсг һу 190 Annals of the een Botanical Garden ct р fæ sheath extensions of Aristogeitonia, series ndron, nanche, Mischodon, Oldfieldia, Parodiodendron, Peta- из ste Picrodendron, and Pin anhea; and at в are present in bundle of Androstachys, Mischodon and nea h veloped in the genera bearing compound Bee but also шоп sporadically wiped in the sub- milv. S УСИШС э OCCUT some Austrobuxus, Choriceras, Hyaenanche, Longe к. (Fig. 11), and Terie Woop ANATOMY wth л аге often absent or only faintly vis ible, Gro е the upper Ата Orinoco » these are plants of dry weet with st aie seasonal нац of moisture. Pore ., diffuse porous in species with grow vth rings), excep in the ring ро of Tetracoccus, which extends into desert regions of western North America. Pore outlines are mostly circular, generally small, and usually less than 100 um diam.; pore diameters are often less than 50 um in das its mentioned ш fom: dry ha ib- itats 9 but wh plates are found in Paradrpeies sue во according to Mennega (1987), mix: with s паре perforations {есы to Araujo т Matt (1 ) „ѕса alariform (Fig. 30), and reticulate plates characterize Podo- calyx. Intervascular pits are transitional in Par- adrypetes and Podocalyx; т ise they are al- ternat small, ca. 5 um or less. Very small intervascular pits (2-3 m) are found in Aristogeitoni eoroepera, and Pseudanthus; somewhat | than usual pits (6-8 um) ur їп Austrobuxus, Celaeno endron, Parodioden- dron, Pi endron, Piranhea, and Podocal s r some на shorter i in Picrode E - 300 in Dissiliaria (c m), one trobuxus (cai: and Pa парче (са. 1200 um), and lon ogen | in Podocalyx ( 12 um). Sculpture on ckenings are Wh yanbeelia (spiral only) and moet (spiral to reticulate). Tyloses and/or tannins are comm in lumina of heartwood vessels; silica deposits occur li in lumina of Whyanbeelia. Imperforate penne ае bear AM sim- ple or bordered pits ple pits, i.e., librifor rm wood fibers, charact th ith compound leaves (Aristogeitonia, Cel oden- dron, Oldfieldia, Piranhea, and Picrodendron) plus potenti Hyaen anche, Eny Pet talos der ; The r rem, icing genera possess only el- п of ment, hardly seems appropria ells. ah e other hand, the tracheids M Pon be а diameters ир to 13 шт ngl to primitive word and both ges inch кА tracheids are present in Austrobux tinguish- able by both wall ен (Fig 22) and pit size. us fibers occur in woods of Aus w 59 a S g [<] аге septate only in Раг odiodendron TN are numerous, r, there are e more did 20 pe e tha n 15 per r mm in е but up to d or 5-seriat racoccus “Fig. 3 olymerous (verti r Fas: riceras, Parodiodendr idi n pi udanthinae. Aggr uideo rays W E . Ray: usually het Visite | (Figs sidered the БЕЙ condition ent procumbent rays are shared by “Androstach73 Volume 81, Number 2 Hayden 191 1994 Euphorbiaceae Subfamily Oldfieldioideae CM v ШЕТУ n 21 lle MADw 104 49), ^ el. ood anatomy of Oldfieldioideae.— 22. Austrobuxus swainii (deBeuzeville s.n., 1 kd thin- a thick- wa d fibers. D Celae nodendron mexicanum (Ortega 35, USw 3886), note long 1 radia tiples of а М 3536) daries nd bott anih ук+ bands of ents 2 due nhea — ata 1 (Ducke e ѕ.п., L Sw 31 " 5), nchyma.— E x = > 3 = a 3T Id 8 3 а x 28 of а ef м гоб 88, USw ne x All vires - 192 Annals of th d чен Я Garden TABLE 1. Wood structure of biovulate euphorbs, after Metcalfe & Chalk (1950). Feature Aporusa-type “Other genera” Glochidion-type Perforations scalariform, simple, or both simple simple i nonseptate, thick-walled nonseptate, thick-walled septate, thin to moderately thick-walled Parenchyma abundant, diffuse or nar- abundant, diffuse to wide absent or scanty row bands some species of MAU which serves to ent Axial xylem parenchyma is mostly diffuse and onstrate that these woods are not a cally чы л Fig. 27), «e E also distinct as i Chalk (950) sugges id present in Aristogeitonia, Dissiliaria, Parodio- Но omocellul mare to erect rays are one of sev- dendron к з initial bands), РЕЦ ndron, Pi- wis alian ge ranhea (Fig. 25), and Voatamalo; on the othe era of Pseudanthinae as a distinct Statice hand, parenchyma is agit: and restricted to group. Vessel to ray pits are of two forms that a few abaxial scanty paratracheal cells in Andro- appesi to hold great the ura Spare ier The stachys, Stachyandra, and the Australian genera 1 z 1 = se Fi ghly irregular of Pseudanthinae (Fig. 24). Parenchyma strands pits that range from circular to elongate with the usually contain a ordinary cells and short scler- elongate pits at various ori ations: vertic 1, hor- ified cells single prismatic crystal (Fig izontal, or diagonal (Fig. 28). This irregular pattern 21); Са parenchyma is absent, how. is interpreted as primitive by virtue of its resem- ever, in Paradr rypetes, deer alyx, e c blance to primitive transitional intervascular pit- Whyanbeelia, the Australian genera of Pseudan g, occurrence aradrypetes and t х of Tetra coccus. Since Podocalyx (see below); irregular vesse el-ray pits sclerified crystalliferous axial xylem parenchy have been retaine ll arborescent Australasian cells al phyllanthoid «t gener nera possess uniform vessel- (е.в. Amanoa and es), presence of p ray pits that are circular and alternate. If circular, cells may be considered primi Oldfieldios lternate vessel-ray pits are indeed the derived deae; their loss may well be apomorphous A character state, this feature evolved twice, since — Petalostigma, Wh lia, and prs ate, t gma, Whyanbee е ааа Дт Pseudanthinae GE e., qante sensu genera of Pseudanthinae; los i in others = ет ў p es a rype tes; Podocal VX. the ре етее of натен ê below), racoccus) may represent convergent pe rather than the lineage of African and Am meri me genera. Ray cells generally bear Uy tan RA several exceptions occur: prismatic crystals are ORIGIN OF OLDFIELDIOIDEAE sporadic in Podocalyx, abundant in Paradrypetes (Araujo & Mattos Filho, 1984 ; silica Ma are FYVIDENCE FROM WOOD АНАТОМ common in Petalostigma; and sclerified cells bear- Pu pomo ht ev structure of biovul eu a prismatic ү are abundant in Aristogei- calfe т Chalk (950; : ischodon Tetracoccus fasciculat vides ient starti int for discussio? these зунан, E are only rarely ec а between Руане апі Oldfield- o. Perforated ray cells are sporadic in ioideae. In this work e groups of gener? A: Аф cime (Milanez, 1 > ede a d distinguished, as indic € in Table 1. At the mos Hyaenanche, but have no superficial of wc = of Oita wo Ободо Pire ed ray cells are fairly дв. ly included in Меїс e Chalk's M а ead orbiaceae according to Giraud та y be char: ase combina паге (1983) A RY this feature in the following e Glock oT in ok y the fiber ee UM genera: Арек Baccaurea, Bri- and | ures of Apor orusa-type v thus Hook. f. ex Planch., Drypetes, es latter tw реноме consisting prim rimaril 9 » yeronima, and Richeria. Perfor ated ray cells phyllanthoid genera. In some insta ces these are also present in Putranjiva (Nazma et al., 1981) coarsely defined categories provi au and Amanoa (unpublished obs Ж spective for placing genera; for onan ше Volume 81, Number 2 1994 Hayden Euphorbiaceae Subfamily Oldfieldioideae moval of Neowawraea Rock, which has Glochidi- ear a (1984) found type woods. [io ЧА ш the reverse of the combi- nation net acterizes most o i ] additio an match the typical oldfeldioid уаш п de above. Further, within Oldfieldioideae, Podoca lyx 0 A n onary p perspective. sot DERN char- rs seen ps to clarify ai oods of b bes ^u idi orbs. In accord with M widely accepted hypotheses of w volution, ennega (1987) has Н ae woods "un long vascular pits, long thick- = ate fibers with bordered pits on both radial n tangential walls, and di z sonally drier ied by pus olution of more song hy E e каш ‚ опе тау геа- Una exploitation of Ripe and/ or of “om nical and storage тешне of the i changes would be manifest in both fiber and Бон ood structure may thus be in- in terms of divergence from primitive Aporusa-type structure, an ecologically restrictive bones: of xylem features. E Red, Sort А sa- and Glochid- ent ed from ample, tracking both perforation plates and de- ve aparon of septate pete yields Ше following between Aporusa- and Pon. -type structure: bd esc A урн Urb., and Pentabrachi- m Mue with exclusively scalariform per- orans is some septate e fibers ; Сеналеца РЫ, Didymo И Kakka. „апа Ji simple а са 2 and some sep- a-type woods, there is a гож ms from exclusively scalariform in Apo- He onima, Maesob S ER ES E yer Benth., and RIMIS to mixed simple and iE fumer ne Кепе mi idend to cmd a. Of co such sequences of erini genera к not ‘be оп желе Phyllanthoideae. In terms of the gross vessel, fiber, and paren- eh ma features of Table tb ithe eygünonary ene HI nad for it sa-type woods. In com- nthoideae experienced a greater ific ation with the additional muli ple perforation plates, an intervascular pits; presence of perforated ray »» eatures. . Overall, their woods are eium to structure. 1 and Podocalyx excepted, Oldfieldioideae aly їау los риту гау cells, but ot therwise retain the features of Aporusa- уре structure oe comments h above). M on Lachnosty: Annals of the 194 Missouri Botanical Garden my of Oldfieldioideae.—28. Austr. y and irregular slitlike vessel d simple perforation plates. 11) dial rley IND USw wide % = Шей ESE ES St SbF А ч > : So Ss 43-353 T EEE S BSR EES Бер ES ко S38 AED DE SIREN — EG ~~ WS BR "ә ч б о gogo. SSS в 8-5-5 5. оок gi Xess БЕЛДЕРДЕ IN INY] рарын хс сеш 2X 8ai- €. 80 Sal ¢ =m Же, 5158 Wood anato 33; radial section, heterocellular ra section, heterocellular ra 42 t JRES 28- Fict ys an 22429), radial 1991), tangential section ential section. — 33. Tetra £ E ‚ Short and narrow ra coccus fasciculatus var section. scalarif« 95, MADw ,Uw2 97 "7 1 3886), tang rays. All bars — 100 um. Volume 81, Number 2 1994 Hayden Euphorbiaceae Subfamily Oldfieldioideae siderable diversity of oldfieldioid woods; base f oldfieldioid genera with woods from other subfam- esence of somewhat primitive woods (Par. gests that the div: Phylluthoideae. осте quite pag in the evo- relatively primitive A poraga: type woods. Ts sym- plesi lutionary | transition between the subfamilies pre- cludes the possibility of specifying one or another phyllanthoid group as closest to the origin of Old- fieldioideae based on this tissue alone. ptate fibers, common in phyllanthoids with Glochidion n-type structure, occur in only one old- fieldioid, Parodiodendron. Опе prominent differ- ene; ow , between thi od and the Glo- chidion-type phyllanthoids is its fairly abundant parenchyma present in tracheal an nt nfigurations. Typic l expressions of Gloch type structure include v The occu 2 Parodiodendron may us represent convergence EVIDENCE FROM STOMATAL STRUCTURE ration of the stomatal pern is one of Old- genera is ошадымы atterns in which the subsidiary cells substantially nd from ог brachyparacytic stomata are аы їп *maining tribes und ideration, i.e., oeae, Antidesmeae, Drypeteae, and Wieland- , A brachyparacytic i in onl ad ypeteae and like M ieae. Subsidiary cell n Dd we he дё of most ae анар peri all Jacent guard cells, The Oldfieldioideae. Unfort nate] у, йы сайы ы of stomatal adi i in Phyllanthoideae and Oldfieldioideae precludes any simple qais eme i abo gut t the Kon t of di iverder nce of important characters quss e ia a lanar- k groups. This situatio of Cronquist's (1988) eie Be seas ten- onabl dencies are reasonably g ships; a rigorous cladistic ae of Phyllanthoi- deae and Oldfieldioideae should distinguish the napomorphic stomatal characters from the par- isms. ADDITIONAL EVIDENCE + ical ch umber a such as th occurrence of theoid teeth, intersecondary veins, and mucilaginous epidermis in Oldfeldioideae and prünitive MEE ae are consistent with the Floral structure ormer’ of Wielandieae i is stint ane ba to be con- ncestral status. Hens wever, plesiomor- e do not present by themselves an overly aen case for consan- guinity. On the other hand, the often reduced carpel d drupa 1967) are difficult to reso ith the c sug gesting x — 12 dfieldioideae (see above) oe E are mes unsuitable as progeni- ains € diit these taxa may have shared a common cestor. Levin & Simpson е aus that the spinules oot and discontinuous ex n pollen of окалы mple та jeny the w DE sim PE type Securinega саш мы 1 as plea ii by shakers (1987)). Securinega aracytic stomata Кан, 02). useful. In our present state of ignorance sem nega stan as another possible near rditive to Oldfeldioideae. SUMMARY Taken together, stomatal and wood characters 1 nuu org 22 1; 1 f£ th. | Желе „ 5 IT * . differentiation of the primitive biovulate tribe Wie- 1 1; | 1 г. h E^ ач Ж ) d Р 196 Annals of th уңа ES Garden ly led to tł ] Drypetes and Tribe Pierii. (Small) Webster. The gen- or ; "Sed nega. In a la piper: context, prosaic < of ега with palmately compound leaves form the con- re vad teeth 1 panne ly comp un = серія m. ar pss Fics odie Bentham Rosid ee of "perti (cf. ва іп Геуш (1986)). RELATIONSHIPS WITHIN OLDFIELDIOIDEAE Classification of subfamily Oldfieldioideae has been in a stat fA th P MER i 5 3 rn а a i3 Ty PEN ү puri o 1 Ра ч р рр ра es че the role of mc in modification ud ear my supports r Ee e TRE & jd 1994; Webster, Tribe Croizatieae We bster. Croizatia, the uen Picrodendreae plus, again, Tetracoccus. As cussed кы а 5 tur: of Picrode сс tne er bearing si or compound leaves, form a teeny. well pier clade and are thus better cla here. Of the two classifications pre- volume, both link Podocalyx with adrypetes and ойе (see above) suggest that at least thes е the Жанасу stem of Oldfi leae; as such, Hy may not be expected to share many synapomor- phies with sach ber. Fach genus of очу еае the Artes of idee the tracheoid pem blasts of sii and the ring porous wood of Tetrac morphic a - irae Шы, je à as easily ob r late di- Mig cig Кош ancestral volte UR Tetra- sel- Eri pitting with ould be used Ii rr and this AK to argue for its inclusion in that vibe. as classified by Levin & Simpson (1994, this volume). {ior 78) the biovulate а with ошо bad associating Old- fieldia and Piranhea XN the по! Bs r a nus Bischofia Pax & Ho ШИП 1922, 1931) апа б + this om ater spiny pm (КОШЕ 1965) апа Mee s -type ood 1987) shows Bischofia to be better pagal in aire Webster (1975) maintained Picrodendron in a monogeneric tribe, te an. {бис eviden (Hayden, 1977, eal Mer Bec Presently. Pibrodendienc includes a ts cu po a), plus eci ide leaves (Androstachys, Mischodon, and Voata- alo) (Levin & Simpson, 4: Webster, 1994). Picrodendreae is characterized Bre ically by al- ternate vessel-ray pits, bundle s bearing RATE well-developed Wu brial veins at leaf margins. Anato: nica data su dendreae that has been adopted by Levin & Simp son (1994) ad Webster (1994). The first group is African-Madagascan-Sri Lankan and consists ischodon, ort a subdivision of Tir Stachyandra, and Voatamalo; these genera ma be defined by stipules adnate to is wer be Shaw, 1970, 1972; Bosser, 1976; e-Smit 19 nd a strong Me tonal u foliolate or wank leaves. Further, Ari t atamalo desc sclerified ray mong the genera o see also blance to any genus of tribe Placement in Pi clusion co by Airy Shaw av base pipi i| Et ш St та The second subgroup of Рой js neo p а а consists of dee Volume 81, Number 2 1994 Hayden Euphorbiaceae Subfamily Oldfieldioideae ndron, and Piranhea; these genera are uni- formly trifoliolate, e alternate phyllotaxy, and so w read ra ae e vein зетін, of A other members 9t the iiba. rth u family by virtue of its septate sd fibers. Nev- ertheless, Parodiodendron seem m dated among the other саң Picrodendreae, ен if its oc ee features are viewed as utapomorphie Oldfieldia is not included in either of the above resemble Ll n in i e oth different ре gen ldfieldia ribe Caletieae Po b opinion on record, Micr ant. us, and ЕЗ bie] jede recognized tog "y tives for well over a century (Agardh, 1918, rege 1866; Bentham, 1880; Griining, iiie р ‚ 1931; Hutchinson, 1969) у hytes of Australia constitute Nooroeper е systematics of Euphorbiaceae. кетон ا‎ added to what was a wholly pressed, sm xd катар (1975) who was im- ertures (um ubt, by the numerous scattered ap- four gener t, 19 bea Kahler, 1965) shared by al iral mal palynological and anatom- efined concepts of re л E ce of crenulate “piggyback” subsid- i a particularly striking synapomorphy йыл. Australasian genera of Oldfieldioideae, monotypic Hyaenanche, endemic to the western Cape of South Africa. Hyaenanche has no other obvious close relatives within Oldfieldioi- deae. Anatomically, its margin ion i markable, but, ae autapomorp perhaps amu vie pn a ant tuf Me diti: separated since the sexes of Gondwanaland. Placement of Hyaenanche in a separate mono- generic nde seems ri Hyaenanche excluded, all Tri ко inally с on deposits in in pines lower hal yes: Te Г; n to- mis provide a robust definition of the Aus- Ње Australasian genera also tend ures buccina epidermal features lato are found in Scagea, which Е strongly for its placement here and notin Acalypho Че һе uniovulate carpels of eti thu: homoplasious with Acalyphoideae, кэз tits and Euphorbioideae oth: М (1994). and рее "i шр (1 н Вазе Petalostigmatinae, and Pseudanthin Aside from he epidermal E e че ihe cahe genera of subtribe Dissiliariinae share few pd genetica ally BARAT anatomical structures. Woods f Dissili vessel-ray pits (Fig. 28), but this is probably. a plesiomorphic feature L ins tend to be ) in the subfamily. Leaf entire Oldfieldioideae, but, as noted above, ккан t aves occur in ies of Austrobuxus, obably ple span ad Austrobuxus, wit y pores; uod ves: is кей ments (often mm), forations, and decina ES seems to be ЫЗА least ното genus of the subt ribe in terms of ood a . Anatom ically, Dissiliaria stands ibe by virtue of high ra oriented areoles, narro rial veins, wavy an lets, ins, ticlina р. extensions, and libri- epidermis cells, bundle form wood fibers Whyanbeelia lacks crystal- жолы axial xylem parenchyma; in this respect it mbles wood of Dalai and сну ip if its retention in 198 Annals of the Missouri Botanical Garden Dissiliariinae may be challenged on this character. Lack of wood material pre saa ermining whether Scagea а nd K in- cluded in Di ( it em Mi dari pn or either sj ысышына gests relationship with Pseudanthinae (Levin & pson, 1 ubtribe D tr be a para ahy assemblage at the Base: of en Australasian clade. Petalostigma is a distinctive genus of Australia and southern New Guinea characterized, as it epidermal cells with mucilage sho s- sociation of Petalostigma with Australasian oar Idfieldioideae to be umet to its e sociation with Androstachys (Webster, 1915; Hayden, 1982); this Missed is also supported biogeographically Both Lev diy Жерен (1994) and Webster (1994) Т Petalostigma in its own monogeneric tribe, e ntly because = ollen bears at leas so rtures out of the equatorial pla unt, 1962; Köhler, 1965), thus differing from the pollen of udant vertheless, Petalostigma res some an al features with Pseudan- thinae: lack of crystalliferous axial xylem paren- chyma the presence of nonlignified phloem fibers in foliar s. In Petalostigma the phlo fibers are thin-walled, where P thinae th pee occluded, so the similarity is only partial in is regard rom the numerous comments above in the an- sees descriptions, н is cis thet the fou Aus- , tribe Caletione in нй traditional or narrow seo} are ted by e of characters involving leaf ar- sr ure, phe anatom vies and wood structure (see also Hayden, 1981). Many of the anatomical char- acters = diced genera suggest a чп е of in- Pan size, increasing disorganization of the онар, matous marginal vein. /Veoroepera is particular! ell. 5 аг marginal ultimate venation alis: lign ified e in f vei arginal vein and LN nified p hoes iat die Si (19 within Pseuda iia ieee the diversity of ve- nation patterns in the latter, but otherwise these taxa are quite similar Meum and evi s evi othamnus, Neoroepera, Petalostigma, and Sca- | gea with the other genera of Pseudanthinae. LITERATURE CITED AGARDH, G. 1858 — Systematis Plantarum... 6 Gleerup, AIRY cii IL EK. Ў Wo ee of new families, S, new names, ete., for the seventh edition of Willis’s “Dictionary.” Kew Bull. 18: 249-2 1966. A Dictionary be the Flowering Plants and isa ын 8 s, Са: mbr idge. enu rica. Ea ser. 21 uu 524. 1972. A seco аан of the genus Aris togeitonia chan (Euphorbiaceae), from East Africa. Kew Bull. 2 197 ^n i е нз of the Flowering Plants Ese eret Ed. 8. Univ. Press, Cont 1975. The опо of ео. Kew Bull. Additional Series IV. Her mes s Stationery Office, Lon 197 i ew or noteworthy Australian Eu- (0 phorbiaceue Kev Bull. 31: 341-398. 80a. New Сои from New бшп- ea. Kew m pd 598. ЖЕП? of New Bull Aina Series VIII. Her beh зке Office, ^ 19 Euphorbiaceae Pp. 129-135 in м Моге & H. R. Tolken (editors), Flowering Plan | ohnsonii Prain and its functional significance 1987. On the unusual pani sais pattern of leaf sclereids in in Androstachys s ii Prain (Euphorbiaceae). Bot. J. Linn. 55- ‘60. pics frt 1909. Makruss or Inform. 1909: 201-204. [Probably ees D. Pra in & L. A. Boodle. 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Wood anatomy of the Euphorbiaceae, in panier p пе түгү "3 Phyllanthoideae. Bot. -126 METCALFE, C. R wd des 71950. Anatomy of the Dicotyledons. Clarendon Press, Oxfor MILANEZ, F. R tomie de Paradrypetes il- icifolia. t. Biol. Veg. 2: 133- 1866. Euphorbiaceae. MUE Am J. abend ARG.). е € (editor), Prodromus systematis getabilis. 15(2): 189-1286. Fortin 1981. «йым Е asson NAZMA, B. Soup & К. VUENDRARAO. 200 Annals of th Missouri Botanical Garden Occurrence of perforated ray cells in the wood of | SOLEREDER, Н. 1908. Systematic es of the Di- A dnos DN (Wall) Hurusawa. I.A.W.A. cotyledons. лекс by L. e & F. Е. Bull., 201- Fritsch; revised by D. H. p ) сй Press, Pax, F. 1884 abide der Euphorbiaceen. Bot. Jahrb. Oxfor Syst. 5: 384- i ont STERN, W. L. 1988. Index xylariorum. 3. Institutional 1890. Euphorbiaceae. In: А. Engler & К. ood collections of the world. IAWA Prantl (editors), Die Natiirlichen ра aA Ed. 1. Teil III. Abteilung 5: 1-119. Wilhelm En- STONE, H. 1904. 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Notes on Australian каси ef II: Pewee and Stachystemon. Kew Bull. 42: 165- Raju, ү, S; 1984. Notes on Mischodon zeylanicus Thwaites: A little-known euphorbiaceous sity from India. J. Econ. Taxon. Bot. ah Lanka and southern 165- & P.N.Rao, 1977. Variation in the structure and development of foliar stomata in the Euphorbi- Bot . Soc. 75: 69-97. 1985. Foliar a i a in sang M. vil & V t Re- search. Bishen Singh & Mahendra Pal Singh, ‘Dehra Dun. RECORD, A 1928. The as ie * of west central Mexico. Trop. Woods 1 The зин each of the family Euphorbiaceae. Trop. Woods 54: 7-4 . W. Hess. pn Timbers of the New World. Yale Univ. Pre s, New Haven. Struct i patterns Котн, I. 1981. s of tropical barks. In: Encyclopedia of Plant Anatomy, Volume 9, Part 3: traeger, Ber m ROTHDAUSCHER, H. 6. halt von Blat Axe ег ычча айныр, (mi Ausschluss der Esphylanthe “| m Centralbl. 6 65-79, 97-108, 129-136, -169, 193- 203, 248-253, 280- as, ps 315, pte eis 385- iiim SARKAR, А. he & N. ytological asse wee Í the famil ly Euphorbiaceae. II. Tribe P Phyl. УШ, Proc. Indian Sci. Congr. Assoc. (III, С) 67: abi thot Tod States. J. Arnold Arbor. 48: 303-4 em Conspectus of a new classification of the гачы us Taxon 24: 593-601. 1987. The saga of the spurges: A review of classification and КУГ ЧОР УЕ. in the Euphorbiales. t. J. Linn. Soc. 1994. Syn x ahs пы genera and suprage- HAE cuna 9 cl Ann. Missouri Bot. Gard. 81: TES та 1. MILLER. 1963. et Rever- ) Ганча) 193-207. — mb: Сш Е & J 1987. Sys tematics of ения нн. Syst. Bot. 12: 89. Growth rings, increment and age forests, savannas and a cx 5 Үл. М. of trees in inundation tain forest in the Neotropics. IAWA Bull. 109-122. APPENDIX 1. Specimens of Oldfieldioideae examined. Xy- larium acronyms follow Stern (1988). Anea s johns ain. Leaf: Gomes & Sous 202 (Ky Wellome Ch Chemical ыы pec ond: Capuron do ARA 4, USw 27434 (= CTFw 13817) (TAN D 5; Pardy S-N- z 5 (= Uw 1 à giu "ЛАО 2127 1991; P jt осе ازا‎ Sei REA Airy Shaw. Leaf: Bal P К) Smith В aa (K). Eo twig: Tanner 3386 TAN Aristogeitonia sp. Begue 727.R. 1, CTFw HP a buss brevipes Air a 51 (К). W руа аа or t -) Air n ar es TET Шун уай 7 (05). К у шой 2 ов, aill.) Airy Shaw. Leaf: ee 2 US); Webster 14972 (DAV). Woody twig: ster 14972 (DAV). Lost Austrobuxus cuneatus (Airy Shaw) Airy Shaw. McKee 12184 4 (K). w. Leaf: quU (Guill S Guillaumin é Baumann. a #77 (09 Нигітапп 1331 (US). Sha w. Leaf: Austr yan horneanus (A. C: iw Mm бан. 1 6872 LO. m h 6669 (US). Wood: А. б, Ам 2; og USw 30522), e 4850 ine huerlimannii Airy Sha Leaf: McKee 713 Austrobuxus lugubris Airy Shaw. Leaf: McKee 25 & s обихиз monianus (Ridl.) Airy Shaw. Leaf: Wray Robinson 5424 (K). NB wie Ms. hent hew VAM 07 (US); С Py FPA« DFP 30 vu ae 21393 es ч (SAR); Si Воесса 7983, USw 29100 (М ‚213 Р СА Sy —————Ó Ы БН Volume 81, Number 2 1994 Hayden Euphorbiaceae Subfamily Oldfieldioideae mington s.n., Uw 21411 (= KEPw 3602) (KEP); USw 1120796, Uw So Es mut. Heu (SAR). f: McKee 26894 (K). Minas Fon yu roi eaf: Baumann- Bodenheim 14113 (K); McPherson 3935 (MO). Wood: McPherson 3923 (MO); McPherson 3935 (MO); Mc- qe? 4603 Austrobuxus rubiginosus (Guillaum.) Airy Shaw. Leaf: “ш n-Bodenheim 15010 (US); Hürlimann 1486 ( pu swainii (de Beuzev. & C. T af: E S White) se P ا‎ vieillardii (Guillaum.) Airy Shaw ‘Leaf: Schmid 2514 (K). Wood: McKee 25222, FPAw DFP sp. AH E ier of California, Davis use B73 Wood: Hyland 6918 (DAV); er & Hyland Ae (DAV ). endron тех dley. Leaf: Ortega 4962 us Onega 6367 (US). Wood: Ortega 35, USw 3886 E cn 29901, MAD-SJRw 1200); Or- FC 67 (05); Uw Hen jd Aw 30657). oriceras tricorne (Benth.) А si Shaw. Leaf: Forman s.n. (LAE, US). W n 7135, Mans 29097, ayliss 6 (US); Cod Sit 1257 ^ билд PRÉ E od: Boh Ag у Wordeman AT ВЕ); d Pretoria s.n. dieck 528 (9 з), (Airy Shaw) Airy Shaw. hns NGF 47324 (K); os с NGF 45108 у ВЕ); pine an & Oberdi shan ЕА Longetia iein (Baill.) Airy Shaw. Leaf: Bauma БСО un 5605 (US). Wood: Baumann- CAL Al ер Mi QUUM dem F. v. Muell. Leaf: Hunt 2802 Moni oody inc pou 2802 (US nt eum ericoides Desf. Leaf: Hahk 428 (US); Johns on & Constab 909. W le 19095 ba e 495 SN € (US). Woody twig: Cam- Micrantheu andru : Boo Us), SM lur 3196 (DAV); Schodde 1 17 77 (US). 3536 NS oorman s.n. (US); Whaite & Whaite odon zeylanicus Th Wheel cus ep Leaf: Ripley 05 (US); > Whe, d 12079 (US) W ood: Jayasuriya 2434 (PDA); Use а banksii Benth. L Leaf: pees " о s.n. Eie die id & Ta e n. (NSW). as Cooper 29. ge 15207) (MA 5, SH 4882, (= Aw 16932, Old, lfieldia No (Welw. ex Oliv.) Léonard. Wood: А. Tanganyika Territory 254, Uw 10951 Rte 495). Old, died. pin Léonar echamps (Comité Nat. du Kivu) s.n., TERV» 1 160 5 (BR). ошеша MM (Chiov.) Milne-Redh af: Рег ead. Le e & Kibuw dicis (NA). Wood: Schlieben 6371 ў 8), U 15635 (= MAD- Te 34030). Oldfieldia sp. Wood: үг 330, TERVw 7522; Dechamps 335, TERVw 7527; Perak 655, TERVw 8499 (BR). Parodiodendron marginivillosum (Speg. JF Hunziker. Leaf: S). Wood: } Hueck 469 (0 7.610C (LIL). Petalostigma banksii Britten & S. Moore. Leaf: Perry 1981 (US); Perry 3531 (US); Perry 3562 (US). Wood: oherty s. п., Usw 21280. (probably ae to Р Petalostigma “glabrescens” ^ е). Leaf: Clem vi uit pubescens or P. trilocular: (US); Wilson 679 D ood: SFCw R 5 lostig ma pubescens Leaf: White pen. (US). 190 К 594-3; Wed ri 977-256; White s.n., re F. v. Muell. Leaf: Clemens 5). ен PRFw лы 10913, (= U Раното qua. 4256 10 US); е pon Е 24259, (= Uw 10949); PRFw 948); Webster & Hyland 18879 (DAV) кыы longepedunculata Jablonski. Leaf: Breteler 4969 (US). Wood: Breteler 4970, MAD-SJRw 55650, (= Uw 12254) (MER, US); Breteler 5096, USw 35682, (= Uw 12306, MAD-SJRw 55702) (MER, NY, U, АС). Piranhea ү Baill. Leaf: UE 5924 (US); Stey- 15 (US). Wood: Capucho 493, USw 22377 CA 6, MADw 30166, 6, MAD- SJRw 23457) (F, AN); viste s.n., USw 31485 (MAD); Krukoff 6163, US 7524 Podocalyx loranth hoides Kl h. Leaf: Bernardi 1675 Dee чер és 21543 fa "rof at NA Williams 0 (US). Wood: Wurdack & Adderle 42795, ) Pseudanthus divaricatissimus PON Arg.) Benth. Leaf: Constable 53354 (US); Ingram s.n. (NSW). a nsta ble рЫ a Ingram s.n. (NSW). Ps lp ori у. af: Cleme 44092 ү vhi 0 (US). Woody twig: Faas 44092 (U Pseudanthus purs Pay ll. Leaf: Beauglehole & Orchards 30452 (N SW); pre 906 (US). Woody twig: Beauglehole & Orchards 30452 (NSW); Muir 906 (US). iaioe u ы Sieb. ex das e ed ii US); Clemens 42750 (US ). W ; Clemens 42750 (US); ns NSW 55972 (NSW 7). Scagea depauperata (Baill.) McPherson. Leaf: Franc 1642a (US). Sca, agea (Guill ry Shaw uil- laumin & norant 11808 се ғоя 2352 (05); eke 2651 is ME merana (Air nA Radcliffe-Smith Leaf: Bar n 6431 ЧК}; [Asa 23335- 5F (K). Stachyanra viii (Airy Shaw) A ЮМО, Smith. Cap 20975 y (К). Wood: Capuron 1914 4.R. "à CT 9 9069 (TAN). Stachyandra sp. Wood: Belin 218.R.6, CTFw 13786 (TAN). Annals of the Missouri Botanical Garden TE d George. Leaf: Blackwell & Grif- fin 3132 (PERTH). Stachystemon үчн T Muell. Arg. Leaf: George 12928 (PERTH). Stachystemon polyandrus (F. Muell.) Benth. Leaf: Hna- tiuk 761262 (PERTH). tachystemon vermiculare Planch. Leaf: Pritzel s.n. (US); Royce 5207 (PERTH). Eve dioicus Parry. Leaf: Moran 13170 (US); Terrell & Gordon 4004 (US); Webster & Hildreth A Campbell 214 Tetracoccus fasciculatu ats.) Croi latus. Leaf: Johnston 7783 (GH, US); Webster, 21221 (DAV). Wood: mats die А S Tetracoccus fasciculatus (Wats.) С hallii (Brandeg d ing pe Webster. & Hildreth 7460 (DAV); Wi s 6617 (US). Wood: Webster & Hil- dreth 7 460 {DA V). Ec c ilicifolius Coville & Gilman. Leaf: Gilma US); Gilman 2181 (US). Wood: Gilman 2181 (09. Voatamalo eugenioides Capuron ex Bosser. Wood: SF 2 R 259, (= CTFT 13816, Uw 23104); SF 5327 R 4 (= CTFT 15079, Uw 23105). Whyanbeelia t & Hyland. Leaf: Irvine 1399 (К). Wood: ana 7945 (К). APPEN ee Oldfelain: se bet works abbreviated as follows: G = Gaucher dpa S = Solereder (1908); MC = Metcalfe & Chalk (1950). puce Prain. Anonymous (1909); MC; Bolza Pp i prs Rao & Raju (1 ras 987); Dahlgren & van Wyk (1988). psc uel (often reported as *Longetia," see below). Rothdeascher (1896), G; Dehay (1935); Math- ou (1 940); Heimsch (1942); MC; Bamber (1974); Rao & Raju (1985). Prin sie Standley. Qoo Record (1938); Record & Hess Bah Hayden (1977). Croizatia Steyerm. Levin (1986 Dissiliaria $^ in ent МС; Dehay (1935); Bamber (1974); Као &A 85). Hyaenan bert & Vahl. Pax (1884); G; S; Dehay (1935), il (1954 x Longe: “Lon PN ains to Аиии ys (= E nget nine Maia there appears to be no previous онаа study of Longetia pens (Baill.) Airy Shaw MM Desfontaines. Pax (1884); С; 5; MC; Rao & Raju pane К е ү 96). Neoroepera Mueller Arg Oldfieldia Bentham. Stone (1904); M C; Assailly (1954); ebacq & Decham iUm s (1964); No & Keating (1972); Hayden (1977); Rao ef М a (19 Ms Araujo & Mattos 198 77) (q.v. for earlier a Raju (1985). Ok (1938); MC: еп (1977); a Roth (1981); Worbes d ie Ps nur Sieber ex Sprengel. Pax (1884); С; S; Rao & Raj Stach E Bl E Pax (1884); G; MC. Tb Rl Engelman ex Parry. Heimsch (1942); MC. — ——— Nen PHYLOGENETIC Geoffrey A. Levin? and Michael G. Simpson? IMPLICATIONS OF POLLEN TRASTRUCTURE IN THE (EUPHORBIACEAE)! ABSTRACT oc pollen ined sla members FR o" uphorbiaceae subfamilies е = apg was studied "p pe sion elec microscopy in order to assess t relationships. We identified 10 rue and tra palynological characters ees Appear t “have Hr sii е. We al a so i i 37 characters of vegetative Hayden of reproductive morphology, 1 D ho Phullanthoid on datai in the literature. Cladistic analysis of the “Odea f the Eu uphor orbiaceae as ge provided g insight into the phylog Л Mle Sy echinate : А егор pu present). э t M from the foot-l ih t ue exception of Ку e basal member of the seri Oldfieldioideae also share the orphies, an exinous foot- s ‘yhapomorphies $i pur or more е pollen apertures and no petals; two homoplastic уы ls o ize these gen roa r to absent and a caruncle on the seeds, also c eats hese era. A clade consisting of a xoi ind Podocaly is is weder! by the synapomorphies o of a ly interstitium fa "mi ystallifero xylem m parenc chym ng Oldfieldioideae < share the synapomorphies We ts. some of which аге co E ontinuous wit with. the columellae and/or wins uniformly зен perforation plates; and alternate intervascular ag КҮ. diagnos sed by the synapomorphy of alternate, circular vessel-ray pitting, and except к ; cus, the mber of the clade, also share four additional synapomorphies, all foliar: compound or eoles, Withi elop dle ns, venation, and well-developed Indian dad he Picrodendreae, synapomorphi ез distinguish а South American clade and an African/ Мае ап/ ИЕ eres Ra ut до пої resolv ve the position of Oldfie ldia within the crode ndreae. The Calet etie e share the ЕТИ. Australas: tribe, the African genus ; Hyaenanche i is the member, ith th all of ends are asian, being united by the estan. a of chambered fair epidermal cells. The Australasian genera es ча E E car: ut the vant the Ph: mee the Oldfieldioideae, these results (1) support the transfer of rte tia and Paradrypetes from ^ oideae to the Oldfieldioidea SE support pe of Scagea, despite its single ovule /locule, in the mbers of the Oldfieldioideae letic. Oldfieldioi me (3) demonstrate that Androstachys and Stachyandra are bona de me pr n belonging in their own family, бег (4) indicate die Neoroepera, as currently circ umscribed, is diphyletic Propose a revised classification of n subfamily in which all suprageneric taxa are Secolo: according to this Phylogeny W быы bine our warmest thanks to John Hayden, ind Largus shared his data and thoughts ı Our study. Without his generous cooperation, o ork w ould hav ve been far jess оле о M SE full ay differ from arbe We tl tors at DAV, GH, K, MO, P, QRS, llen from collections in thei care. Jon Blevitt, ee ost of the Hun _ 7» lor removing or a send us to remove pollen. fr i пере, Nicki Watson embedded an sectio. ned much о. the pol ollen n for | EM and p rinted mos at the I HAVE Mas кнн Mich wo thank G valuable а) 9 Webster and Mike Huft for inviting us to participate Mike er and an anonymous Pena made Invertebra on. on a aes version of this paper. Levin thanks Rick Brusca and Regina Wetzer of Marine Mark Тута. partment San Diego Natural History Museum for ip ieee him use of their Apple computers, and illustrations e Exhibits Dep tiic, San Diego Natural History Department, for geli erage the с — анн of pollen expertise ап : t this research тм по! ефе been possible. researc supported in needs by a San Diego State Car Partme uc st addres: nt of Botany n Diego N ll Histor ry Museum, P.O. Box 1390, San Diego, California 92112, U.S.A. quer Center f for Biodiversity, "llinois Natural такан Ѕигуеу, 607 East Peabody Drive, Champaign, Illinois a of Biology, San Diego State University, San Diego, California 92182, U.S.A. ANN. Missouni Bor. GARD. 81: 203-238. 1994. Annals of the Missouri Botanical Garden Light microscopy (LM) of re morphology in the Тарни асеае (Erdtma 165-175; 1962; Kohler, Ma. i e er ed ecent classifi- y (SEM) to study pollen of КО ае d th veys have yet been published several papers have treated a variety of genera throughout the family, пеач Phyllanthoideae and E (Bonnell & Riollet, 1980; De- Per El-Ghazaly & Raj, 1986; sare ia & Tirel, = (Hayden et al., 1984; microscopy (TEM exine rchitecture, and these papers treated only five genera. We initiated a study of pollen ultrastructure in е, which is gen- oe regarded ; as a: eq cans PUTES in ing three Жл ез. . Of the two biovulate берчи; Phyllanthoideae is by far the larger and pre bly is paraphyletic and basal to the Oldfieldioideae uniovulate subfamilies Meme 1967, 1975, 1994; Webster et al., 19 g tails of our pollen study will be published ченге (Simpson & Levin, in press). Here we present an overview and ا‎ ROW ie of the Ii. concentrating on the Yee To evaluate pollen characters and better u phological and anatomical data compiled by Hay- den (1980, 1994, this issue) and of selected re- productive characters. MATERIALS AND METHODS TAXA We examined 40 species in 34 genera with LM many of these genera we also examined d TEM; m with SEM (Table 1). Nine genera are members of by prior researchers (e.g., Kohler, 65; Webster, 1975, 1994; Levin, 1986c). We included A is and Securinega dur. |! issima becaus cent of pollen “of Ollie (Webster, 1984; Webster et al., 1987) mined ‚ placed D Our selection of Oldfieldioideae inclu 5 We chose not to se and Picroden pollen ultrastructure information 7 o r (1994) inc (dod: in the P rient Webste dioideae. Like Webster бом we accept Par gelonium as a s of Aristogeitonia (Rad- cliffe-Smith, 1987b) e Stachyandra as distinct from Androstachys (Radcliffe-Smith, 1990). We have been unable to get pollen of Canaca, à segregate from юй i (Webster, 1994). SPECIMEN PREPARATION Flowers from dried erben pe (cited in Table 1) were rehydrated at For Hs studies pollen dium mess et al., rv dehydrated to 100% ethanol, then рга aduall T to S Au methylal (dimethoxyme Fre . An aced in Т or - T zi = о 4 = @ ч [:] [nd = о crosco e also examined ае Іоапеа Бу С. ерѕіег М observations, nthers in 0-1 pollen we in c % emm M اا‎ s phosphate buffer fo hey appear in "po h f investigated Euphorbiaceae. Taxa wh one o of our preferred cladograms (Figs. 31-334A). Data are from Simpson & І are d following the voucher citation. d; techniques used t in Phyllanthoideae are sessi Dep Rai Taxa in pulido are efti in the order in observe each taxon йо var. alchorneo [Croat 51307, MO] LM, H thin (0.21 um) Ektexine Pollen Peri eriaper- Size tural X: RE, Exine thick- Shape um) Aperture type sculpturing Foot-layer Interstitium Tectum enings PHYLLANTHOIDEAE manoa guianensis Aublet Oblate 36 x 42 3-colporate Echinate, spines 4.1 | Homogeneous, ase (homolo- Absent (?) Absent [Tillett et al. 45256 um long thick (1.0 um) with echinae?) A , SEM, TE Amanoa strobilacea Muell. Oblate 38 x 54 3-colporate Reticulate Homogeneous, Columellate Perforate/homoge- Absent Arg. [Thoma. E thick (1.4 um) eous (2.7 um) M, T Antidesma membranaceum _ Prolate 23 x 12 3-colporate Reticulate Homogeneous, Columellate Perforate/homoge- Absent Muell. е ov 4973, thin (0.11 um) neous (0.30 um) MO] LM, T Antidesma — Tul. Prolate 20 x 12 3-colporate Reticulate Homogeneous, Columellate Perforate/homoge- Absent [Kemp 525, MO] LM, thin (0.15 um) eous (0.31 um) TEM Aporusa falcifera Hook. f. — Globose to 16 x 15 3-colporate Reticulate Homogeneous, Columellate Perforate/homoge- Absent 1, MO] LM, sub-pro- thin (0.17 um) neous (0.29 um) SEM, TEM ate Did ymocistus xen cau Globose 12 3-colporate Rugulate, scabrate, Homogeneous, Columellate Perforate/homoge- Absent . [Dodso M Tor scabrae 0.08 um thin (0.12 um) neous (0.20 um) 2961, MO] LM, SEM, Drypetes gps al Prolate 30 x 20 3-colporate Reticulate Homogeneous, Columellate Perforate/homoge- Absent Krug & Urban [Chambers thin (0.18 um) neous (0.37 um) & Stern 266, US] ye SEM Hyeronima alchorneoides Al- Prolate 33 x 19 3-colporate Reticulate omogeneous, Columellate Absent lem ides Perforate/homoge- neous (0.40 um) | 766: z JOQUINN ‘1g eunioA eeepioipjeljpio ш әлцопдѕедіп ueliod uosduulS * ui^91 TaBLE 1. Continued. Ektexine Pollen Peri eriaper- Size ural x E, Exine thick Taxon Shape ит) Aperture type sculpturing Foot-layer Interstitium Tectum enings enocardia acid. Hive A Oblate 25 x 28 3-porate Rugulate, scabrate, omogeneous, Columellate Perforate/homoge- Present We Wilde 4044, MO] L scabrae 0.07 thin (0.18 um) nous (0.26 um) (Жык. ulmoides Oli- Oblate 19 x 21 3-porate Rugulate, scabrate, Homogeneous, Columellate Perforate/homoge- Present ver [Lebrun 2119, MO] scabrae 0.07 um thin (0.16 um) neous (0.19 um) SEM, TEM long ju PIA discoide Prolate 27 x 22 3-colporate Reticulate gcc uo Columellate Perforate/homoge- Absent (Baillon) Webster ssp. niti n (0.17 ш neous (0.43 um) ^. (Pax) Webster [Chase 150, МО] IM, T R spp." Prolate to 13 x 13 3-colporate to Reticulate Details unknown — Columellate Perforate/homoge- ? globose to 4 60-pantopor- neous x 33 ate Securinega durissima J. Oblate 22 x 25 3-colporate Echinate, spines 1.0 Homogeneous, Columellate Perforate/homoge- Absent melin [Lorence 1397, um long thin (0.28 um) us (0.32 um) MO] LM, SEM, TEM OLDFIELDIOIDEAE roizatia naiguatensis Stey- Globose H2 3-brevicolporate Foveolate, echinate, Homogeneous, Irr xim discontinu- Microperforate/ Present n. [Berry et al. 4124, spines 3.3 um long thick (0.63 um) w/ foot-lay granular (0.24 DAV] LM, SEM, TEM um Paradrypetes subintegrifolia Oblate 22 x 27 4-brevicolporate, Psilate-wrinkled, echi- Irregular (0.12 Absent Imperforate (0.35 Absent i off 4999, zoni-apertur- nate, spines 1.4 um дш) um) U] LM at Я rerit Oblate 22 x 25 4-porate, zoni- Psilate-wrinkled, echi- Irregular (0.11 Absent ape Absent Men eta al. aperturate ate, spines 2.8 um um) baculate (0.33 tea LM, SEM, p TEM Tetr C. Parry Globose 46 x 47 4-pororate, zoni- Verrucate, echinate, Irregular (0.35 Tregear, gonn Microperforate/ 2 (Henry i Dice s.n. (8 aperturate spines 4.7 um long um) w/foot-lay: baculate (0.63 um) April 1980), SD] LM, SEM, TEM иәрео jeolue}Og unossiIN 902 әш JO sjeuuy TABLE 1. Continued. Ektexine Pollen ; Регїарег- Size tural (P x E, Exine thick Taxon Shape um) Aperture type sculpturing Foot-layer Interstitium Tectum enings Parodiodendron marginivil- Oblate 24 x 27 ca. 7- “porate, Verrucate, echinate, ^ Absent n discontinu- — Microperforate/ Absent losum (Speg.) Hunz. [Ver- oni-apertur- spines 3.1 um lon; ous w/foot-layer baculate (1.1 veerst & Cuezzo 7610. ate um) US] LM, SEM, TEM Picrodendron baccatum (L.) Oblate 26 x 29 5-8-brevicolpor- Verrucate, ea Irregular, thin Irregular, discontinu- — Microperforate/ Absent Urban’ LM, SEM ate, zoni-aper- spines 3 um lon ous w/foot-layer baculate (0.6— TE urate 0.8 um Piranhea ipie Baillon Oblate 32 x 35 6-porate, zoni- ^ Verrucate, echinate, ^ Absent Irregular Microperforate/ ? [Berg l. P19789, MO] aperturate spines 2.7 um long "ien (0.39 LM, SEM. le Celaenodendron mexicanum Sub-oblate 39 x 40 7(8)-porate, Echinate, spines 2.9 Absent Irregular Метей) Absent ce mais [Perez & оды ае шт lon, baculate (0.59 6, MO] LM i nee africana a & Oblate 28 × 30 5- 6. -рог Verrucate, echinate, ^ Absent Irregular Microperforate/ Absent Hook. f. [Fox 157, К] zoni- ae pines 3.5 um long baculate (0.44 LM, > аїе ит Androstachys s johnson Globose 50 5-7-pantoporate voi iip Homogeneous, Columellate Microperforate/ho- Absent ain Hh ean 3549, spin 1 um long thin (0.07 um) mogeneous (0.36 ee SEM, TEM um оу. merana (Airy — Globose 44 4—6-porate, Echinate, spines 1.4 Absent + Irregular Microperforate/ho- Absent haw) R.-Sm. [Capuron zoni-apertur- um long geneous (0.52 23335-SF, P] LM, SEM, ate TEM Stachyandra deci ET Globose 46 5-7-por Echinate, spines 0.9 Absent Irregular b coda Absent Em w) R.-Sm. [Сариг zoni- eid um long us (0.50 ii PI IM “TEM ate а Bos- Globose 44 5-6-por Echinate, spines 3.3 а to absent ie pe Microperforate/ ? ser т (apa 22327 -SF, zoni- ape um long (0.14 um) w/foot-laye baculate (0.38 K]L um) 766} z JOQUINN ‘18 әшпүол eeeproipjeupiO U! әлпуәпд$едп uerog uosduulS § ui^e1 10€ TABLE 1. Continued. Ektexine Pollen 2 Регіарег- Size tural {Рх Е; Exine thick- Taxon Shape ит) — type sculpturing Foot-layer Interstitium Tectum enings Mischodon zeylanicus Globose 38 5- T-por Echinate, spines 3.8 Irregular (0.25 Irregular, cba Microperforate/ ? Thwaites е 336, МО] zoni- ee um long um) ous w/foot-lay baculate (0.44 LM, SEM, TEM ate um) Axisiogétionia st ded Oblate 30 x 34 5-7- egit Echinate, spines 4.1 ^ Absent Irregular Microperforate/ Absent Airy Sha ta ate, zoni-aper- um long culate (0.24 45256, K] L ТЕ turate um gemere DO (Gaert- Suboblate 40 x 42 6-7-brevicolpor- те, echinate, Irregular (0.34 Irregular, discontinu- become Absent er) Lambert aes & ate, zoni-aper- spines 2.1 um long um) ous w/foot-layer baculate (0.47 Bleck 20843A, MO] L turate е ЅЕМ, ТЕ ШО spp.'^^* LM, Oblate 25 x 28 5-T-porate, Foveolate, echinate, Irregular Irregular, discontinu- — Microperforate/ ? SEM, ТЕМ to 29 pM spines 3-5.5 um us w/foot-layer baculate (0.9— x 33 ong 4 um Whyanbeelia terrae-reginae Oblate 21 x 30 5- а -рог Verrucate, echinate, Absent Irregular Microperforate/ Present Airy Shaw & Hyland [Hy- zoni- a spines 4.1 um long culate (0.48 land 3041, DAV] LM ate SEM, TEM Longetia buxoides Baillon Oblate 27 х 30 6-7-colporate, Psilate-wrinkled, sca- Absent Irregular X Imperforate Present [McPherson 3789, MO] oni-apertur- br: dinum cabrae 0.4 (0.84 um) LM, SEM, TE um Choriceras majus Airy Shaw Oblate 26 x 28 O-porate, zoni- Psilate-wrinkled, sca- Absent Irregular Microperforate/ho- Present [Hyland 10633, K] LM, aperturate brate, scabrae 0.2 mogeneous (0.43 EM, TEM um long um Dissiliaria baloghioides Е. ^ Subglobose 36 x 38 5-6-porate, Echinate, spines 4.1 ^ Absent Irregular nde aca Present Muell. [MacPherson s.n. zoni-apertur- um long baculate (0.75 (23 Jan 1980), QRS] LM, ate тоот pubescens Oblate 32 x 34 5-porate, zoni- ^ Echinate, spines 0.9 Absent Irregular Microperforate/ Absent in [Speck 1818, K) aperturate um long baculate (0.38 : ‚ TEM um) uepJer) [eoiuejog unossiIA 80c ay} jo sjeuuy TABLE 1. Continued. Ektexine Pollen P Periaper- Size tural (px: E; Exine thick- Taxon Shape um) Aperture type sculpturing Foot-layer Interstitium Tectum enings кг pei vein oe Oblate 32 x 34 5-6-porate, Echinate, spines 1.0 Absent Irregular Microperforate/ Absent F. Muell. я zoni-apertur- um long ba up e (0.51 1993, K] L TEM ate Neoroepera buxifolia Muell. Globose 33 16-20-pantopor- тене, penes Absent Irregular ТРИЕ. + Arg. el ate Л um lon baculate (0.63 LM, SEM, TE um) Kairotha phyll Globose 26 10-1 2-pantopor- е echinate, Homogeneous, Irregular, cot аср берейн Absent thoides (Airy Shaw) Airy ate spines 1.8 um long thick (0.61 um) ous w/foo ar (0.21 [Streiman pm) 24462, K] LM, SEM, TEM Scagea oligostemon (Guillau- Globose 30 16-20-pantopor- к» echinate, Homogeneous, Irregular, iy a Microperforate/ Absent in) McPh [Webst ate spines 1.9 шт long thick (0.70 um) ous w/foot-lay granular (0.19 19181, DAV] LM, SEM, TEM Neoroepera banksii Benth. Сіороѕе 36 ca. 25-pantopor- Echinate, spines 3.7 | Homogeneous, Irr iique оаа T ate/ Absent [Hyland 6945, K] LM, ate um long thick (0.98 um) ous w/foot-lay: granular (0.26 емее hexandrum Globose 34 30-40-pantopor- и. echinate, Homogeneous, Irregular, discontinu- са др Absent ook. f. [Briggs & John- ate spines 3.0 um long thick (1.5 um) ous w/foot-layer granular (0.33 son 4419, DA AV] IM, Pseudanthus Lea Globose 23 10-14-pantopor- cipe pucr Homogeneo rregular, discontinu- — Microperforate/ Absent mus (Muell. Arg.) Be B ate s 1.2 um lon thick (0. 65 и m) ous w/foot-layer granular (0.22 Det ^ DAV] LM, um) SEM, T Stachystemon polyandrus ^ Globose 22 l2-pantoporate — Echinate, spines 1.8 | Homogeneous, Irregular, discontinu- Mi a Absent (F. Muell.) Benth. [Young um long thin (0.34 um) ous w/foot-layer 5 LM, TE granular (0.14 кж ' Punt, 1962, 1980, 1987; ? Kohler, 1965; * Martin, 1974; * Hayden et al., 1984. 7661 с 1әдшпм '{8 ӘШПоЛ әвәріотрјәүріо U! әлтдопдѕедіп uerod иоѕашіѕ § ui^e1 602 210 Annals of the Missouri Botanical Garden 2% tetroxide for 2 hours. The material was then quickly rinsed twice in buffer sively dehydrated to 100% e iamond kni Reichert Ultracut-E ultramicrotome, mounted on nw о ретш 00 Ew ‘copper рза, mi in in 50% ало, 15 minutes) and lead citrate (0.2% aq., 7 minutes). To inhibit stain precipitation, grids © J | d o of filtered, distilled water between and after post- staining changes. sevi ors and photographs were made on a electron microscope s EM 410 E scant PHYLOGENETIC ANALYSIS We analyzed our data using Wagner parsimony as d аана by PAUP version 3.1 (Swofford, 1993) апа HENNIG86 version 1.5 (Farris, 1988). енш b both квга and random 209 repli- reconnection (TBR) branch-swa ир! ing. With poy N we used the options whennig* and а ENPE са реадер. ыша g. In a T ш Mus aud bb to с the effect of Farris’ в (1969) ий. O weighting of characters, i in which th ach character ees using MacClade ver- sion 3.0 (Maddison & Maddison, 1992). RESULTS AND DISCUSSION CHARACTER DEFINITION dd. ind. are listed i in Table 2. Of the 52 odere. 45 are binary characters. We treate all but one of the seven multistate characters as unordered. The Phe sae is снае ter 24, lea we coded this character. The f following i isa бесов of the rationale for our decisions of haracte state definition for the bela selected d etative morphological and дык and герго- ductive morphological characters. Palynological characters. Table 1 summa- rizes hog pollen morplitdogy of ke Lise we ex- rr E M PM унт. L & Levin (in prep.). We recognized 10 WES characters Е бузы shape. This character distinguishes t equatorial axes) those such eoroe pera (Figs. 3 Parodiodendron (Figs. 5 6), Androstachys (Figs. 7, 8), Micrantheum character state because = © e ay oblate shapes occ & Doyle, 1975). pL those taxa with we than three aperture ges from four (e.g., Par rad UN Levin, 1 1992: fig. 1) Neoroepera, Figs. 3, k Micrantheum, Figs. 9, 10; see also Table 1). ecause we c quie see no clear discontin inuity thes among e latter taxê, we treated the presence of 4-20 or more p ingle a А 3. Aperture position. This character distingus s between zoni-aperturate gr ins (apertures present only along the equatorial plane; €^ Hyeronima, Fig. 1) and pan-aperturate rains Pon hic pear: oo" surface; ё." Neoroepera, Figs. 4; Pseudanthus, Figs 1 2) Report aperture number and distri? tion of yen ait differ — Erdtman (1952: 167) described the po PN polyforate, i.e., pantoporate with Ë 12@ 12 apertures (not oligoforate, i.e., wit & on fewer p as misquoted by Dahlgren ler (1965), studying the sam e sample, и scribed the pollen as ame pantopor? | & E E f Volume 81, Number 2 1994 evin & Simpso 211 Pil пани in Oldfieldioideae ТАВ Characters and character states used in cladistic Bos of the Euphorbiaceae. 1. Pollen ик е: 0 = БЫШ tea 1 = prolate. 2. Apert ber: 3. iere Ls 0 = zoni- йи 1 = pan- apertur. 4. ie =e 0 = colporate; 1 = brevicolporate- l): О = absent; 1 = minute («0.5 mk s - um (>0. 9 u m). 6. S reticulat eolate; = гара ulat sis — verrucate; 4 — miele tie Fs ne " — homogeneous/ ы = hom -abse ntinuous ш ee HN 1 = к vith foot d er; 2 = c0 = fi E ;2- Мав 5 л ме 3= | о granular; 4 = imperforate; = absent. 10. Bicis periapertural thickenings: 0 = absent; 1 d = diffuse; 1 = © Parlin plates: 0 = 8-4 а and simple; = all simple. 13. ^" ervascular pits: 0 = transitional; 1 = alternate. t < pits: 0 = small, 1 = larger. 5: 0 = по septate; = septate. 16. es 0 = heterocellular; 1 = homocellular erect. gregate; 1 = aggregate. rm 5: 0 = irregular shape and pattern; 1 is rds and alternate. d = tay cells: 0 = unsclerified; 1 = scleri- эр АХ parenchyma: = present; | = absent. ystalliferous бај 1 en оа 0 = scleri- ; 1 = unscleri 22. Secondary phloem a. 0 = present; 1 = 23. Кы 0 = wider than radicle; 1 = narrower Pri than radicle. эзы lade: 0 = simple; 1 = compound; 2 = uni- 5. % EUM Y: 0 = alternate; 1 = opposite. 27, рій 1 muci ве: O = present; 1 = absent. 28, Epidermal cells: 0 — undivi 29. Epidermal anticlinal walls: 0 = straight; und я om 1 п: 0 = ai es or нарта 30. § = anomocytic; 2 = vog Sy А niic val боо stomatal pore): 3l ~ Straight; 1 = crenulate. * Phloem fibe rs of primary vein nified: thick-walled and unlignified; 2 = aude did "p un- 33, Ciystals i in mesophyll: 0 = present; 1 = absent. 34, Be ем, extension: 0 = absent; 1 = present. аа, е sl extension crystals: 0 = absent; 1 = TABLE 2. Continued. 35. Venation: 0 = brochidodromous; 1 = irregularly mou: irregularly brochidodromous; 1 = an ET S9 © о ll 37. Venation: 0 = all others; 1 = reticulodromous or hy ous. 8. Intram: кыны п: 0 = absent; 1 = present. 39. Marginal eden venation: o = looped or incom- plete; 1 = fimbriate. 40. diee venation: 0 = irregular; 1 = ortho 41. учо ак р 0= e 1 42 — 0 = ed or ا‎ 1 = well de- velo 43. id idioblas' = abse M 44. Columnar foliar gy ids: 0 = ds rue anti 45, Stipules resent; 1 = shea: 46. Stipules: 0 = on stem; 1 = epipetiolar 47. Raphides: 0 = absent; 1 = жыл 3 :0=а 48. Sexuality: 0 = dioecious; 49 ў : 5: 50. Оушез/1ос опе. 51. Caruncle on seeds: 0 = absent; 1 = presen 52. Endosperm: 0 = abundant; Г = scanty or nr or possibly зше E with 14—18 ine ped apertures най P П “pore” with a granular mem- examined pollen o у both Androstachys and Stachyandra (= Androstachys subg. drehandratachys vem which : — but clos rela ROR leaf morphology and deus deor, К, 1988; Radcliffe- ia easily seen wi edem ed were zoni-ape P 6 pores and S. eb v with 5- E pores Erdtman (1952: 170), studying what be 212 Annals of the Missouri Botanical Garden Fic Scanning aoa macrogre hs of ollen of Eu orbitae subfamil Phyllanthoidea айу сма (3-6 2 à р es ш Pein dron marginivillosum. Scale DEB іп 1,3,5 = 5 ит; іп 2, 4, 6 = a um. called Petalostigma pubescens, and Punt bescens we found that some of the ape (1962) and Kohler (1965), studying кс they indeed lay outside the equatorial plane, " "i called P. quadriloculare, reported that the P. quadriloculare all the apertures Mr aperture s of Petalostigma do not a exactly strictly equatorial. Prior to Airy Shaw > р? 1 the equatorial plane. Our observations are vision of the genus in 1976, many сме not entirely consistent with theirs. In Р. ри- were misidentified (Airy Shaw, 1976). Ane Volume 81, Number 2 994 evin & Sim 213 mpso € Areas in Oldfieldioideae Fic {nd 9/712. $c са rostac Iba johns Nonii ur sz » Rd amples came from spec imens at Kew nnotated by Airy Shaw Bicis he y the Or Sir 'at * have J ons. У there > tentati 5 refore tentatively coded the pollen of Pet, 1 t y ае а e А ‚ Stigma as zoni-aperturate. It is possible, nning electron micrographs of pollen of iwi y Dem — Oldfieldioideae. Figure 11 i : 10. Mic rantheum hexandrum. indicates an aperture. Scale bars in 7, 9, 11 = 5 125 разре из divaricatissimus. Arrow in pm; 1 Uu 10 2 um. however, that the Mem distribution i in Pet- alostigma may zoni-aperturat в: рап- spertanto, particu larly if the species чабыла аге =ч эмс dn nong the species we sample ^d, except for Androstachys, taxa with 7 apertures are zoni-aperturate or essentially so, aed those 214 Annals of the Missouri Botanical Garden 10 or more apertures are all pan-aper- taxa were coded **?" because we were unable turate (see Table 1). A literature review, sum- to determine from the micrographs whether marized in Simpson evin (in prep.), shows the pollen was verrucate or f this pattern to be generally true, the sole ex- sculpturing of Picrodendron might be termed ception we are aw being some species *vermifo: at the tectal processes are of аот which аге pan-aperturate with somewhat elongated and twisted, but we coded 6-8 apertu it as verrucate because of its basic similarity 4. Aperture d is Apertures were either elon- to that pattern and because the sculp of gate and colporate (e.g., ima, Figs. 1, Parodiodendron (Fig. 6) is somewhat t 2) or considerably shorter, intergrading from sitional. Psilate-wrinkled pollen has a relatively brevicolporate 7 кк. to porate (e.g., Neo- шоо ес tum Wohin y devoid of шо i- roepera, Figs. 3, 4; Pseudanthus, Figs. 11 Androstachys, Fig. 8 12); т may appear кошсун: ї Mud , per "e M ti We divided exine wall sculpturing into the fol- fonction of specimen Pcp aigi а. two characters: We treated the structure of the exine wall as ee pene iria! tied: үн fr apparato debes p M The frst he ыер эр rie ا‎ he three pud regions DESEE ES £s "H nima, керене ea Figs. 1, 2). However, a number of taxa possess TA "ыр ed Several taxa have a typical con- ines, the length of which ranges fr el- s, homogeneous ektexinous foot-layer atively long (e.g., Parodiodendron, Figs. 5, Np tene Fig. 13; Amanoa, Figs 14, ) to short (e.g., Androstachys, Figs. 7, 8) 15; Croizatia, Fig. 16). Analysis of the vari und a clear discontinuity be- ation in foot-layer thickness within and among tween short te ines um long, these taxa (Fig. 17 ws four distinct size called *scabrae" by Reitsma, 1970) and spines lasses that could a recognized as of greater length um, called “ ae" character states (Stevens, 1991). In prelimi by Reitsma), we coded these as separate states nary cladistic analyses we tried dividing (see Table 1 characte ter mio four fium: MEC the two 6. Sculpturing ا‎ This character de- g ngle state, or com notes the sculpturing of the outer exine sur- чыга all but the smallest pe E into à face between spines (if present) or the general single state. Because the resulting trees were rface sculpturing if spines are not present he same Il three cases so few t Reticul en has a perforate tectum in fell into the two largest size classes, We decided ch a network of muri c delimited, to recognize only P regardless of the relative size of the perfora- > 0.5 um) for the homogeneous foot-layer. tions (e.g., nima, Figs ). Foveolate I ining , the ektexino fa pollen also has a perforate tectum but with layer was very irregular, consistin| f on) no clear indication of tectal muri (see Croiza- tinuous, almost gran Jements (Figs: 18- ‚ Webster et al., 1 figs. 2, 3). Inr 3 extreme (e.g tracoccus, Ёё ognizing both states we differentiated between 19), this irregular foot-layer was present micropores, tiny perfi ns in the tectum around the entire grain; at the othe “a present in most pollen, and the considerably (e.g., Oldfieldia, Fig. 18), the ektexinous pd larger perforations found in reticulate and fo- жт was essentially а ет: герге - 7 4 veolate pollen. Rugulate pollen has a tectum by ta and scattered granules. Muss that appears conspicuously wrinkled or folded; ^ ^ term edia g., Longetia, z a regular pattern of perforations is lacking 23), preve E us from sine the tw (see Hymenocardia and Didymocistus, figs. extremes as oe е inn эк 1-6 in Levin & Simpson, 1994, this issue). layer of ekte og Policy (Fig- oe The remaining states lack large perforations, з vete (Fig. 21) w interpret һе though they may be microperforate. Verru- an irregular foot-layer (see uisi #8, cate pollen has the tectal surface covered with low | small rounded processes, or verrucae (see Tet- En dexine was found as a thin to noi racoccus, Webster et al., 1987: fig. 5). Some thick ied layer pies thickening а! Volume 81, Number 2 1994 Levin & Simpso Pollen fec AS in Oldfieldioideae 215 stri obilac. cea. MA. € ЦРО ННЦ ge © 1 Yo hf. MI . —16. Cro oizatia naiguatensis. Arrow ius periphery) in almost all taxa (se tidesma, Fig. 13; Oldfieldia, Fig. 19) id E sep relatively thick outer layer discon- ru rom m a thin, granular inner layer, we Е preted the inner layer as an ake eee r and the interstitium as absent. tia E variables affect tectum struc- toss n of perforation and its orga- е tectum of some taxa (e.g., An- 15. Amanoa Scale bars — 0.5 um ES 13- Тт. ам. Кыш, ou modd — 13. yp MR ыс — 1 4. A in Figure i tidesma, Fig. 13; Amanoa strobilacea, Fig. 15) is homogeneous and penetrated by Bie perforations rather than gee cu. a state we called “perfora z Because of the large з size i the perforations, е some TEM sectio ~ = m wit Hiree states Were vices The tectum may ., An- — drostachys, F n mirer) bac- ul i итего ар- e © 5 2 [7] E d 5°98 aa © к, © both perforation rate, perforations (e.g., Paradrypetes, Fig. 21; 216 Annals of the "rire Botanical Garden Foot-layer thickness (um) е р ERA ы FIGURE 17. Distribution of pollen exine foot-layer thickness among sampled species of Euphorbiaceae. a (solid f sp eg ‹ the mean thickness, a vertical bar the range, and a rectangle dard deviati side of the = = minimum of 10 meas surements from one to three grains). a ies are arranged by in my thickness WI subfamily, with the Phy th Oldfieldioideae on the right ek arate size-c! poi that we considered pot Hoge character states. Mirpo for species names: А = Amanoa пос guiant ШЕ = Amanoa strobilacea, ANjo = Androstac chys joh ii, ANme = Antidesma dens aceum, ANve = Anti venosa, APfa = Aporusa falcifera, ARmo = Aris togeitonia monophylla, CEme = Celaen odendron pine CHma = Choriceras majus, CRna = Croizatia а ы Diba = Dissiliaria baloghioides, Dich = be aii chrysadenius, DRla — Drypetes niii а Гутепосагііа acida, HYal = Hyeronima а ir Ygl = enanche globosa, HYul = Hym онаа ulmoides, К. = Kairothamnus phyllanthoides аб getia buxoides, MAdi = Marsan taria discoidea, M rantheum grt m, т РАша = zeylanicus, NEba = Neoroepera "i NEbu = Neo uxifolia, OLaf = Oldfieldia africana, P. PEqu ee marginivillosum, PAsu = Parad. arypetes шев PEpu ; = . Petalostig ma pub ХО des ostigma — ulare, Phir = Pi ranhea ae ers POlo s Р5й = Р» STpo divaricatissimus, inega inane ssima, STme tende тегапа, = Stachyste e amalo mon polyandras een: = 5 achyandra rate TEdi = Tetracoccus dioicus, VOeu Voata eugenioides, WHte = Whyanbeelia terrae-regin Longetia, Fig. 23). Finally, we interpreted the only supportable breaks associated = the exine of Amanoa guianensis (Fig. 14) as single taxa and bone leading to ol acking a tectum; the echinae of this taxon netically uninformative autapomo e ae we believe to be homologous to columellae. coded as айыны states. We the ro * Evidence for this interpretation is the diff use tec ickness as a c The structure of these echinae (distally г "eri 10. Ektexinous periapertural sickening had ike е ана Ear pes poss nay pollen of most of the taxa we examin! sharply point ) bable ho much thicker exine around the apertures кг mology of the echi сене in the nonapertural regions. In most ri with the columellae of няк тейден Ао these thickenings were formed by the were ee (Fig. 15). ine, but in a few genera the thicken a ài Tec thickne ss varied considerably formed by the ektexine; in no са thick mong t чы ха we examined (Fig. 26). The observe both endexinous and ektexinous thic variation was са continuous, however, with Volume 81, Number 2 1994 Levin & Simpson 217 Pollen Ultrastructure in Oldfieldioideae Еск! eae subfamily Oldfieldioideae. — 18. e though in several of these the endexine ele fe d eu. а enings had little Hien peers a ce, we have included only ektexinous ickenings in gic analys Alternative codings. we have coded ет eed in which Though the pollen SENE y ap to permit а Чол them to vary Sapa i die pattern coding аш within the taxa combi with the e E and жам 8 is problem, demonst to address it, easi у r umber ated with the vien. for aperture charact "m perm A ave coded these cter. NN a group potentially supported by the а urate йет дек is nested within a group ga y defined by aperture number greater than er, . 9 OUP with more than three zonate ures is n teeters E t paraphyletic based on these 5 alone. seg with pan- te pollen, the derived state of one char- RES 18-21. T 5 Oldfieldia afric. апа. — 19. zer == Arrows indicate endexi it Таласы dius 20. Кы ркан gen 21. Paradrypetes subintegrifolia. acter, have more than three ее» the derived these two n three оше, zs pan -apertur- apertures to be bón venia einn GEO based on this single pollen characte e tried recoding these быы and found des p codings s us same cladogr Similar apere (уре, poten rini and aperture. number. perhaps others. As with the example we дор above, combining a pai characters into a one unordered multistate cha r did not affect the results our 1 tic analysis. We have a as we have, therefore chosen to presen which we consider simpler and more informative. Vegetative morphology and anatomy. Most of the wood and leaf нн (11-47) that we included in our analysis are discussed by Hayden Annals of the Missouri Botanical Garden FiGURES 22-25. Transmission electron micrographs of pollen о in Fig ale bars = 0.5 (1994, this issue). We obtained data for the Old- fieldioideae from his paper, and data for genera in Phyllanthoideae and the uniovulate subfamilies of аа Matos ae & Mattos Filho (1984), 1 ed b most cases, we could c е directly from Hayden (1994). Following is a dis- cussion of those characters he did not include or f ich our interpretation may differ from Hay- > 12. Perforation plates. Rather than distin- guishing exclusively scalariform from — simple qus "A we com- a we treated them as separate states, po- larizing the character would have been difficult because both conditions are n within some of the genera of Phy thoideae we included as outgroup $ соз ray cell h r =] © cells containing one or сеа utgro 20. Axial vien hegi Axial xyle mly Oldfieldioideae.— 22. EE loeum Bid Longetia buxoides. — 24. Fea ЛК 7-20. pueden polyandrus. ITOW . Se £g. 1935; Matos А aujo & Mattos dea: "a XM In Aristogeitonia, Mischocon, l4he nlternate-lea r cies Ф ТОБО occus, howe 2 Ha n talliferous ray cells are ceri the literature g thoideae prevent us е of ges about km conditi of som m pa- present in the Old- nchym п varies мышы, though ees distribution eu . Only limited data Ой 50 e we were able o to record 3ta Pe or . Despite this ext л E e cation, we include! harac among cause it helps resolve relationships | Volume 81, Number 2 1994 Levin Pollen & Sim pson 219 Ultrastructure in Oldfieldioideae 244 1994), as equivocal (“?”) for this char- acter. hi opposite or whorled leaves. cause the phylogeny of species within THHIDTEPEESShRPh-AB,!áP—O»NTEPEPEPEDUEESATIT- EPSP?LEZIZDNII280G20EROSSZQGOLESNTIEGQEEQBÍS «TorqZtXoozr«coorü.Xozüordan2^5orryortoz-el i Taxa PORE 6 — Distribution of nal 1 h TEN the same as in Figure 17. а the taxa we included ps for the these genera is not well understood, we Ж vol Srey : coded these taxa as polymorphic for this . Leaf blade. We tried treating this char- character. acter both as an dered ch ter ani 29. Stomatal pattern. Because Phyllanthoi- аз an ordered character arranged simple deae have both paracytic and brachypar- to compound to unifoliolate. In our ex- cytic stomata, s es on the same Perience with other groups, unifoliolate leaf (Levin 1986a), we treated these as leaves appear to be reductions from com- he same state. Though the ound leaves rather than evolved stomatal pattern in Oldfieldioideae is an- directly from simple leaves. With this data cytic, Aporusa, one of outgroup Set, we found that it is more parsimonious genera, has anisocytic stomata. We di to derive unifoliolate leaves from com- ot differentia tween undivided and Pound leaves, regardless of whether the subdivided subsidiary cells because this character is or unor may within a single leaf (Hayden, ed Mier. eum, interpreted as hav- 994). ing either sessile c und leaves or sim- 31. Phloem fibers of primary vein. In most fibe ayde s difference in wall thickness sure a priori that the unlignified condition was homologous in these five taxa. We 220 Annals of the Missouri Botanical Garden 000 of our proposed leaf venation transfor- BROCHIDODROMOUS ation series and i (1) (1) 45—46. Stipules es МР ЫН & Tirel (1987) stat- that the stipules of Austrobuxus and 100 1 ?01 ; ý IRREGULAR <-> RETICULODROMOUS- Longetia are extremely reduced to ah BROCHIDODROMOUS HYPHODROMOUS sent. The same may be true of other Dis siliariinae sensu Webster (1994), but we "m 2 са ific reference to the stip- 110 ше г genera in the subtribe RETICULODROMOUS- KLADODROMOUS and have therefore coded these genera as FIGURE 27. Ch ling for leaf venation, char- equivo i) acters 35-37. w & 35—37. . Bundle erefore recognized three unordered states for the phloem fibers: lignified, thick- walled and unlignified, and thin-walled and unli nified. gar d crystals. Som enera have no bundle sheath extensions (character 33, state 0). We coded these genera as equivocal (‘‘?’’) for character Vena on. Four basic leaf venation pat- ur а brochidodromous pattern, which e coded “100.” More irregular still is M nei but it prob- dod allow the two li code odromous vena- tion *?01." See Figure 27 for a diagram iry Shaw & H Airy Shaw, 1976) stated |^ ipiis certe visae, die obsoleto con not T pi 5 of us (L Pu ) sol k terrae-reginae bland 3041, "DAY) and unable to find evidence of stipules. We have therefore Е содей Whyanbeelia as la ipules (see fur ther discussio section on taxonomic implication: oded Micrantheum as equivoca a (?") for this character; see the discussion of character 24 a > -1 biaceae with ‘crystals have druses а prismatic crystals completen ness Re du tive morpholo, For eproductiv phology: s of repr oductive some characte: cult to interp those characters w (48- reasonable confidence, only five оһагаб ug 52) remained. We expect that a care eful 5 eee n _ M —— —— M — Volume 81, Number 2 1994 Levin & Sim 221 pson Pollen Ultrastructure in Oldfieldioideae E reproductive morphology will yield sig- ant characters beyond those in our meager list. owing is a discu rs a each e the Pico ^ : W f our data on re productive фы ору from Рах а Hoffmann (1922, 1931), Bosser (1976), А Shaw ый 1974, = 1980), and McPherson & Tirel ither monoecious 48. Sexuality. Plants may be e regarding the character state in each genus except Aristogeitonia. Pax Нов (1931) described this genus as monoecious, but both Airy Shaw (1972 Radcliffe- more recent authors who with far more specimens. 49. i Though petals a are — — in ре ne ldioideae, ia Sog relationships among the out- 50. Oves осше. Since Jussieu (1824) first rec- ogniz FEREN has bee regarded as one of ‘dl rincipal taxonomic pin Phyllanthoideae ar tw ain: cule. Scagea, however, is uniovu- late (McPherson, 1985; McPherson & Tirel, к, also the cladistic analysis below). Caruncle on m seeds. Most Peete ш have is 8 Apparent exceptions are ndrostachys (Dahlgren & v yk, ) Chi erial a ffmann, 1922; R cliffe-Smi dedii ше элү ley, 1927 1980), áp pocos (Hayden, 1977 oe et al., 1984). dat on Neoroepera Росса Voatamalo, 52 dei Whyanbe des x the exception of Hyaen e (Pax & & Hofmann , 1922) and Picadas п (Hayden, 1977; Hayden et al., 1984), ошын dese have seeds w k osperm ere unable to yim data on Kairothamnus, Voatamalo, and Whyanbee- сн ARACTER POLARIZATION binge logeneti analysis of the primitive pue © 18 complicated by the absence of a clea (see review by ا رو‎ fog and therefore taxa sho characters. Taxa that have been proposed i in recent & Wolfe, be chosen In the absence of a well-a ted outg pe c & Cantino (1984; see also Maddison et al., 1984) this method, the potential outgroups are used alone and in plausibly related groups in separate mat resulting cladograms regions of consensus. In our analyses, we peri each tt riii Elaeo- plus m p p aene carpaceae pl lacour- tiaceae plus Elae aeocarpaceae ee Sterculiaceae Dilleniidae of Cronquist, 1981), Linales plus Fla- courtiaceae (Chase et у 1991; hase, pers. comm. 1l e either without any ogy. гейде topol CLADISTIC ANALYSIS Table 3 contains the character X taxon data m r systematic implications a the pollen data we first ош ed ШЕН analyses on — data е. n the com plete o dat; set. In both cases both RUM HENNIG86 yielded numerous equally par ы. t ous trees, differing mainly in local een f taxa. We therefore produced a strict consensus tree for each d t. Choice id not ns remaining taxa enoug t a consensus among the various trees has limited lution. use of th ur limited of non-oldfieldioid tax estrict most of our discuss e will re relatore among Oldfieldioidea Pollen. Initial analysis showed that nonspinal sculpturing (character 6) was ve D Hove of this and the larg e number e a for t ring we 16 dropped this character from further т Тһе — tree based on the data al wn in Figure 28. We show only the sre alee кк Croizatia, дй petes, and Scagea); the tree is rooted the 222 Annals hires ен Garden | га . Character X taxon matrix for cladi ti Euphorbiace eae u red. ^ ex IEA an ا‎ or энен штар state. Taxa hia at cha are dca a an “&” utgroup = Baal first, followed by taxa in the Euphorbiaceae. Taxa in the Phy llanthoideae are listed древа, iis taxa in the Oldfieldioideae are in the order shown іп one of our preferred cladograms (Figs. 31-334). Characters 000000000111111111122222222223333333333444444444455 Таха pv tbrebtpepbhtie eps pui O SS Outgroups Flacourtiaceae Celastraceae 000000770701 inales UYT i iiU 1 Elaeocarpaceae Sterculiaceae 0272292722270] Euphorbiaceae Acalyphoideae Crotonoideae 0 Euphorbioideae бере. Pilate strobilacea 0001 ر‎ поа guianensis 0001 Antide. S ARA Aporusa 01000 Didymocistus n100? Drypetes 71000 Hyeronim n1000 Hymenocardia 000112000101 01001 Margaritaria 001000 Phyllanthus Securi nnnon1009 Oldfieldioideac саби roizatia Parddryperes 0101000 Pod 6 nnnin Tetracoccus onnn1010 T 0 Parodiodendron 000010?0 Picrodendron 0001001 Piranhea 42001010 6 EEE min 10001000 Oldfield: оппо 1 Androstachys 01001010 Stachya Boi 01000000 ۲ oatamalo 10010?? Mischodon 41001010 Aristogeitonia Hyaenanche z 4510001011 Austrobuxus nni0001010 Whyanbeelia 001 00] wi Longetia 01011421 оой) 1010 Сһо icera 0010001010 Dissiliaria 10 Petalosti д ^ Neoroepera buxifolia 00000115 t Kairothamnus 0007001100 соз ае Volume 81, Number 2 Levin & Simpso 1994 Pollen dis so Н in Oldfieldioideae 223 TABLE 3. Continued. Characters 0000000001111111111222222222233333333334444444444555 Таха 1234567890123456789012345678901234567890123456789012 Sca agea 111159 Neoroepera banksii 0i | 1271 Micrantheum 011 Pseudanthu. Stachystemon ntire tree nate (5: 2; character number: state number from ). The e (with all the RA їп i А “bia ceae we included) Table 2) and has apertures that are brevicolporate has a length of 34 steps and a consistency index to pororate to porate (4: 1) and greater than three (C.I) of 0.47. The portion of the tree i in number (2: 1). It was these characteristics that just the Oldfieldioideae, as sh i „is initially pro d Köhler (1965) and Webster eps 1975) to recognize the subfamily. When Kuhl- Though large portions of the tree are unresolved mann (19 cribed Paradrypete allied it (as would be expected with only eight characters), with Drypetes Vahl llanthoideae. Hutchin- our pollen data show several significant syna son (1969) was the first to place Pa маре morphies. As previously noted ee be: (1962, with genera now placed in Oldfiel ioideae, i um- 1987), Köhler (1965), Webs ter (1975, 1994), and ably because of its opposite leaves; ie gas Hayden et al. (1984), A Kog isechi- data offer clear support for ea й $ 2 o б б o 3 n E c E oo 7 o: ES E EES СЕТЕ ТЕ TREET T T siti Bee ee S rer E rT ж PSPS SSC ae 01:214 SSE 28 3.28245 2525 FRSC FES EE Ss у PAPE sc te SA ор SE AESES EES x Le OPO UP RN E QR ee ee e z с Ф *91 :4 Y9: 0 7:2, 8:1, 9:2 2:1, 4:1, 5:2 га Strict consensus cladogram of the Oldfieldioideae based on palynological characters (1-5, the character number, 4 n alo, M In heim and the following cladograms c oe state changes are shown by number ot the derived state (see ‘fable 2). 224 Annals of the я Botanical Garden too, shares these synapomorphies. Note that though Croizatia i is well ет чаша th « Oldfeldoideae, s it has only th We echinate pollen, but Nas — = 0, тоу it б m wer d ро: would appear then that aire sculpture Ы convergent in Securinega re in the Phyllantho 26 to ‘ace shape is apomorphic or plesiomorphic Based on узе alone, Androstachys would ap- pear t diede th kde ded of the rest of the Old- fieldi o be closely related to Stach- pee Three vitium we ures appear to columellae are where и is aps хаш 8: K 8 recognized by amined Austrobuxus and Picrodendron. The pol- len Шаку) v used are теш rahe шиш ly chotomy at this level in the tree. Palynolo ogical characters allow г recognition of st кф Meter (Fig. AER Another clade, consisting of Mns (Fig. 16), (Fig. roepe remaining t share pantoporate To (3: 1). On this BPE Мех epera, Mi- antheu. um, Pseudanthus. an group by Punt (1962), 975), who united mnus and previously recognized а: Köhler (1965), and Webster (1 them in the Pseudanthinae TN Scagea had not yet been described). As previously noted by Kohler (1965), Neoroepera бүг: (Figs. 3, 4), шш panes por hs ine е fore lacks the synapomorphies that place N. и sii їп this clade. In all the е nious trees, pa Androstachys is convergent with the pantoporate Na ж Ж е МА ee ys E | AN, | buxifo- lia (cf. Fig. 28). Combined character set. As might be € perte iam the ospiti — vs hart beue 1 on f relati nshir We again, dropp £dd the Шеш reduced or absent (8: 2). Because we a ted the озн character as unorder ed. i P. ч фы ү 1 1 In that case, havin mellae discontinuous with the foot- fis peed be зарини and thus opener for these two Back. Eo is more parsimonious to interpret the төй of the interstitium as apomorphic. e nd clade, consisting of Choriceras, Dis- siliaria, Longetia (Fig. писмо. do « Within this clade, Choriceras and L clade, sharing reduced ines les ss than 1 um long (5: 1); it is easy to ee late ici and e queries: tectum (9: 4) of Lon analysis because of its relatively high бошору of many taxa (but see hE discussion d m character below). Several hundred equally ра arsi- monious trees resulted. We did not search the sets We did compare the c fieldioideae, however, and found 24 equally "d hese taxa. Figure 2 r imonious arrangem ft di illustrates the strict consensus tree for t n tionships among the of Euphorbiaceae lon i5 = 0.45) if the uen jes are ех ng C e 24 trees includ the oe deae are 104 steps long = 0.54) if all i acters that vary among thes taxa are incl in or 97 steps long (C.I. — if auti un are excluded ee analysis of the data vegetative шы and anatomy data less (taken from челе 1094) yielded similar but resolved tre Volume 81, Number 2 1994 Levin & Simpson 225 Pollen Ultrastructure in Oldfieldioideae Amanoa strobil. Amanoa guian. Drypetes Hieronyma Aporusa Antidesma ACALYPHOIDEAE CROTONOIDEAE EUPHORBIOIDEAE Hymenocardia Phyllanthus Margaritaria Securinega Didymocistus { OLDFIELDIOIDEAE FIGURE 29, vegetative, and Str: f +} repr em tive порина (1-5,.7-52).N ames in all apial le letters are of subfamilies. the ехе. f of the Oldfieldioideae, represented here by a triangle, is shown in Figur ymorphism hyllot dus ldia; d ridi й ипге- s for ile 30B-D). All asic e- eng Figure 30B require two reversals nsider unlikely: derivation of scalariform Oration р 1 аз (ch, rvascular pitting water 13). E. bi aig of beth genera of Old- fiel т ed known perforation nd transitional ма pitting, Раг- Ypetes and Pod are about 1.0-1.4 mm "^ Я ° s E E esed =~ 0.3-0. d m 1984). whi 0; Matos aujo & Mattos Filho, ch i iS consistent iura the козе SINE that these character states are plesiomorphic. These 1 Sio 4:6, ] = } 1 E 123 of Austrobuxus, Dissiliaria, Hyaenanche, Peta- lostigma, Whyanbeelia, and the clade consisting of Choriceras plus Longetia within clade B of беш иге is ave of these three giclee are ag i о Банй uin low homoplasy greater n those with high ho- moplasy. These two cladograms, which we prefer because they do not require um hate reversals and are оңо supported by what w we consider He y (es ht (1 O&O) whic are shown in Figures (The third cladogram, not shown, is similar h but nanche the sister эш ey th Austrobuxus, Choriceras, Disa, с^ ^ nbeelia. 2 In contrast to the polychotomy a (Fi igs. 304 and 20, the. poly- in chotomies чыл in Figur omorphies fici — lengths) rather therefore cannot be further — with our data. d" es sisi ed using pollen data alone (e.g., Fig 28) a e almost entirely consistent with our bed i trees produced using all the data (Figs. 226 Annals of th Missouri Botanica Garden Clade A Clade B Parodiodendron Picrodendron Celaenodendron Stachyandra Aristogeitonia Hyaenanche Whyanbeelia Kairothamnus Neoroepera banks. Pseudanthus Stachystemon Croizatia GAP po ee D poA dm В C P ATE B C D еше берет. 15.7% 30. а of the ООба, ary on edi ied vegetative, and reproduc equally а -5, 7-59). — Clades A and В а major clad f ene cladograms.— B- D. Cladograms toa the iP imi) most parsimonious per men d Goin atia (“С”), Tetracoccus (“Т”), th g p Podocalyx (“P”), and clades A two exceptions are the positions of (Airy Shaw, 1970; Dahlgren & van : A: 90). In -33). The Androstachys and Croizatia. Instead of being bas Radcliffe-Smith, 1990). turn al, the я based cladogram (Fig. 2 8), morphological synapomorphies (notably с! АШКАНЫ hys forms a clade with Stachyandra, alternate vessel-ray pits (18: 1), compound wir which it аншы is closely related (Radcliffe- —foliolate leaves 1 well-developed be ul- Smith, - This relationship is supported by extensions ge 1), riate mà — three synapomorphies: microperforate/homoge- timate venation (39: 1), well- developed areo ele ( » neous tectum (9: 1), foliar veins beyond the ter- 1), and etes stipules (4 46: 1)) caus se Andros: tiaries indistinguishable in a reticulum (41: 1), and — chys and es - : An- the presence of columnar foliar sclereids (44: 1); the Bier тараана (Figs. 31, 32). Placing e in addition, these genera share rather unusual floral drostachys ase of subfamily, a5 sho morphology, notably the staminate inflorescences — in Fis ure 28,. increases the length of the com in£ which consist of a triad of flowers, each with nu- data set tree by at least nine steps and FU r merous stamens borne on an elongate androphore Androstachys and Stachyandra togethe Volume 81, Number 2 1994 Levin & Simpson 227 Pollen Ultrastructure in Oldfieldioideae Croizatia Paradrypetes Podocalyx PICRODENDREAE CALETIEAE 2:3, 7:2, 49:1, 51:1 абе; vegetative, and Ficure 31. P reproductive an ters (1—5, 7e 52) ЕС эзи гергевеп 32 and 33. Only synapomorphies are shown, so some ch Tetracoccus Parodiodendron Picrodendron Piranhea Celaenodendron t tribes whose s mede eia m are ciini in Figures aracters are more homoplastic than indicated. Oldfieldia Androstachys Stachyandra Voatamalo Mischodon Aristogeitonia 24:1, 33:1, 39:1, 42:1 d ` Fia = Pref, : А ae | сма 14:54 “1 4 1 1 A m hai (1-5 1-52). Only synapomorphies are shown, so som тор! е bo lastic a aie 228 Annals of the Missouri Botanical Garden 5 © t. а. 3 i; 6 lr cE Ss 5 5 © 3 с o o E © © = £ S 2 2 E: 5 Е = a E с 2 = Е 2 = 9 5 © 2 K o o z o o = 5 2 Ф + > о = = = Les o o о S 9 s x 5 E © 8 A Е © = = © 5 2 8 cm ч z EI o a a = x $ 2 5 e 6 5:1 36:1 22:1, gan 23:1, 35:1 31:1, 38:1 31:2 7:1, 9:3 3:1, 16:1, 18:1, 48:1 10:1, 45:1 21:1 27:1 5 5 © є 2 8 a " б SE а ee ur o 2 2 3 @ ° @ Ф © Е o E t E С y 8 g = © 2 9 2 © © z Е > 5 = S = o 2 o о 5 o © E 5 £ Б] 42 о геа > hed o > о © 5 $ b u a. ч a 21 о 2 2 8 8 2 > c [7] re 48:0 5:1 25 A э 7:1, 9:3 31:2 10:1, 45:1 3:1, 16:1, 18:1 B 48:1 REZE Fic The two preferred most parsimonious ayo a hes B) of керегине: tribe Caletiea d p SS vegetative, тайы reproductive characters (1-5, 7-52). With е. tion of character synapomorphies are shown, o some characters are more mills than indica Volume 81, Number 2 1994 Levin & Simpson 229 Pollen Ultrastructure in Oldfieldioideae аи с Є A 5 5 Е v S Eus? 5 2 ze E Коз с = o = > 9 vigi S Ө © л Ост a Bore HORNS ow cow. Ox SOG o © ос $$ s D o MeO. 5c t s E BID m хоо 8 One. 0 9. Dr gm o Bons o - б Ww p DADA жш Peet GOON сыр даза 8 в Ми ш а аса соо бф 2 SE F пш шг oS си на x Ё 5 c a o о S в 2 5 a Е = o S$ E o E BE в § § 2 в 3 OOK c ШАО ОР o of б E n dmg Rr c DE QE M 2 Вто dom OLE o 0 B gw 9 muto Qu OU AS D. p een e o 8 2 fo 8 д БО DIS URSUS ime oO E .9.9 р ЮЕ 0 md + <3 л о балш з бе я саїе ЖЫШ pellet wrinkled ЕЗ eq cal haracter states fo са А Ficu E 34. Cladogram of the Old In the d ldi h 6, are s shown by boxes below the Bison names; ; where there i is no ‘box we do not кы the state e foun pl 7 that taxon. Note wide би 1 base oo | at least seven additional steps. ea ast, placing Androstachys with St ft р Sadia dc эш length of the pollen-based tree istic, а namely reversals to a thin, ho- tinuous ded cot-layer (7: 0) and columellae co Phic states f, the foot-layer (8: 0). The plesiomor- char or the exine foot-layer and SM "ters 7 and 8) found in Andro achys a Pt ore ie better regarded as reversals sched ncestral conditions. Characters of res UD ab- =~ -d s | 1) a and pre. f a caruncle ix ; combine with poll g more t pertur. ) to support the monophyly беа, ы. ae tachystemon, as 5 ggested by pollen мер 28), increases the length of the com- Set tree by five steps, whereas placing б ata го и за at the base of the pollen-based У опе additional step. The similarities in exine structure these taxa share, i.e., thick studied, such data may support or contradict this p em is. The results of this analysis clarify the synapo- Sorte of the Oldfieldioideae. As suggested b dat ne echinate pollen apertures e iiis Ted having more » than three apertures (2: 1) is not a synapo- morphy of the entire subfamily be the basal genus, Croizatia, retains the plesiomorphic state of three КОС, The phylogenies illustrated in igures 3 w that having с a llae dis- s a further continuous from A foot-layer (8: morphy of the x waar in Mares bad Ж уар Б LÀ ae те d data аге missing ee Б our several taxa, several ра atterns сап be 4). Notably, no Е ЕЕЕ ar аге Кпо heise sculpturing (6: 0), the сое condi- n to have TABLE 4. Proposed classification of the Oldfieldioi- deae. Зирор tap are arrang ged such Her each I it in sequence. чту” indicates each type genus. Tribe I. Croizatieae Webste Genus: Croizatia med (T) ribe I eben e Webster Subtribe Podocalycinae Webster Genus: Podocalyx Klotzsch (T) Levin ос 5 Subtribe Paradrypetinae С. Genus: Pa корее рш (Т) Tribe II. Pi Webster Subtribe Tetracoccinae С. Lev Genus: Mirac Engl € ex к p (T) Subtribe Р. Сысы! Cel: dend Standl gya dron Hunz., Picrodendron Кош a pem hea Baillon ubtribe don eusinae Pax & K. Hoffma Genus: Oldfieldia Benth. & Hook. f. incl. Pai- a Welw. ex Benth., T) sine Mischodontinae Müll. Ar nera: Androstachys Prain, Aristogeitonia Prain ee ari (rada Léandri), Mı diia s (Т), Stachyandra sj А P gos o Cap Tribe ш. rares Subtri be anc Baillon ex Müll. Arg. Genus: Hyaenanche Lambert (T) Subtribe Беда шын a K. Hoffi Genera: Austrobux (incl. C illau- min?), Choriceras Baillon, Dissiliaria F. Muell ex Baillo: ), Longetia Baillon ex l. Arg., hy ia Airy Shaw Hylan Subtribe Pseudanthinae enera: Kairothamnus Airy Shas. Micrantheum esf., Neoroepera Müll. Arg., Pseudanthus Sieber ex Sprengel (T), Е McPherson, Stachystemon Plancho: tion in the Phyllanthoideae. Therefore nonreticu- late Sa bi may be said to characterize the Oldfieldioi Further pass of pollen character evolution are discussed by Simpson & Levin (in press), and evolutionary panon of mos a я: the кеше Hayden (1 1994). Here we will concentrate on the taxonomic impli- cations of the synapomorphies shown in Figures 31-33. TAXONOMIC IMPLICATIONS Since Kóhler (1965) first recognized the Old- fieldioideae as a distinct subfamily and subdivided Annals of the Missouri Botanical Garden it into trikes, three further теак of the sub- O60) 4:4 far nil not recognize any s subfamilies, and his ‘treatment d Kohler (1965) a genera among five tribes, of also taining phyllanthoid genera. He was the first, ho ver, t ce Paradrypetes with other genera of the Oldfieldioideae. Webster (1975, 1994), in con tras n palynology and presented nts е either of the d 4 erred we genies е оп ош set (Figs Lu d reluc combined d d. it use findings in the form of a s propose the аео: shown in e which all the suprageneric tax monophyletic Dis to our preferred ini igs. 31-33). W enies (Fig ) have not given € taxa equal rank, but for suprageneric taxà мг боме the seque nona pe tion prop We divide the Oldfieldioideae into е ee Cr yd t i “consist? Te first pn oiza ieae S Oel Шеп with more than ese aperture: oh poet opposite or the Oldfie Moin and leaves alternate. Ar the other commo: way that it is impossible, using P to determine the state at most le Volume 81, Number 2 1994 Levin & S 231 трѕо Pollen ieee ae in Oldfieldioideae <, &, bl d N^ uU S S 9 PI D <, Ў б 9 g 8 p z © © Q & A é Ў Ы i Ir а —— o o 8 S o S 9 = © A Jum C0 € = c t © 5 5 $ = 8 3 ‘= E: Ё E Qu» 0 2 £ © 5 c E £ 8 = 8 .E a 2 m 2 = o [7] i є 59 69 so 3 о c @ E] © s wees © o & с = о © @ 9 5 e E: o Ф = — 5 б S Ed a © o 8 2 EIN à g = f 5 5 а а à = Е & d d rrt Ir AT ME i ee i c c : 5 " ола 2 = = 5 E Eos: Е в Е ов а eee AES хоу 9 o o Dou o t Q. m a £ © € m page Peete oS иба SE GS ee Ss S mco 'E ES BBS EAE CG HEHE TE EEES Te T e eaa LCE GOES ы a cee Tr а?» Bese ге MESES ete ce Re ЕЕЕ Ade 5 > Bd = © n 5:123 LEE: D B NH - Б зге, л тле, к еек 2 Sû BÛ Û ù Ó Ont SB MTT CO AS A г 200 OOS HE ЕЗЕП ES C3 C3 NH NAE ERO UNO NOUO UO GE S | Оду CS Gaf ар», 9a | (7 - 5 7 | 7 Distribution: ег, 42 New World LZ Africa E ric Cladogram of Oldfieldioideae showing the geographic distribution of the taxa and o „н 35. Classification of the Oldfieldioideae. bfa ig (Fig. 37). In ne the first pair di r es above the cotyle i e 1984) p. leaves are ves e (Hayden et al., Oldfeldicid ation of the other alternate-leaved Va eae could clarify vers er this is devel- чан M that could aid in t termi- the заны, of this character. of sy Wily пар речну shared by (Fig ive 9 REE lacking in Croizatia ould argued th shits genus shoul Реа ы. г data, how- E. s reta ever, arm v е little doubt that Croizatia is the sister EE India E Sepa E equivocal ur proposed group of the remaining Oldfieldioideae. Further- mbi if i reat Croizatia as the basal membe t fieldioideae. The second tribe, the Podocalyceae, is supported duction or loss ine (8: 2). In у u apomorphy, r of the interst elements in the exine ( addition, unsclerified crystalliferous axial м а геп 1), сопуег m elsewhere in tree, supports this clade (Fig. 31) Psilate- saute’ 232 Annals of the Missouri Botanical Garden E z x e 9 c Lm É 9 o E: 0 o "Es ә > с E & B8B S g 2 LE e de = = o коо с в Bs E Qi X Dno D oo $ p.029 50 S с о © 9 m ua o xum mu race m cn тш. даш. е Шаш m e r1 чел боо жр с БЕ Әәә Л ос & 24224 2 =й. © o 9 ug E == оо SBeseEcSesesSFSugses Ss F275 BE S 9 = qq o0 0 Ма. co Pic a 5 92 9 o cnc $ СОО БОА UO «Ою >“ ч D €. €) B» R Ез шш E шшш шшг шш M M EH M EM M BM B NB BN Nd Character 7 FicunE 36. nonspinal sculpturing (6: 4) may also be a synapo- mo: rphy of Шш pue: but hne 5g the, an- is is ambiguous oe 34) Because Pani. gue and Pod. 1992), we place each in its own subtr The rest © ee die ai dde two ique E4 equivocal Cladogram of Oldfieldioideae showing character 7, pollen ektexir sing АШ and Podocalyx, has fonr МЕШ Ре еѕ tracoccus pigs the Podo! u we >ч 33) by ы. ste AN 'The rest of (3 иеа share Me ` le ti pau Ne (Fig. 32): compo mop astic еудар P N Cb2: 1) and lt te int l йлы, (18: 1), 2), 9: 1), апа опе, ті forate/ba ulat tec nm o. 2^ Ons (ЭЗ сас гіаїе ieta venation (39 : Fig. 38), that ther d ] and well-developed areoles (42: . We гео ШИ within the clade. It is tempting to interpret + ver- three subtribes in t a job the rucate اا‎ sculpturing (6: 3) as a synapo- the three clades of this trichotom Idfielat shies orphy of this clade, but the ancestral state is sole genus of Paivaeusinae, has no at: E equivocal gis 4). We divide this clade into two that clearly unite it with either clade. Inde 0 g jor split within tl bf. ata provide no support for the monophyl d e ily. eldia, which has four species; given th ks e The first of th , the Picrodendreae e (Fig. n phyllotaxy within the genus (Fig. d ud 32), is defined by eed alterate vessel-ray pitting sibi that it i not monophyletic pi Picro- (18: 1), a synap hy that is convergent else- examined. Genera in the American n subtr pee where. Tetracoccus, the basal member of this clade, ^ dendrinae (Fig. эй share one unique Re ani has several autapomorphies and we therefore pro- phys loss of bati in шеи esophyll (32: ^^ pose placing it in its own su eae, emphasiz- ing that the pollen of теда. like that of both two ical iio pitting (14: :1( an tals in the bundle sheath extensions (34: Picr oden- are uniformly alternate (25: 0) in the | | ут 81, Number 2 Levin & Simpson 233 Pollen Ultrastructure in Oldfieldioideae 5 5 x i " B x g ра m 3 S ? 5 E ә ‚© a.” E ESI is Ei es? 281] CT5ERBSEEBIILILDL $ s об Е $255 т > $35593292955E2255:22255235 Е? ЖЕ КЕ ЕБ ЕКЕ ERE ТК Т ЕВ: КО ре ос б Sse ss gpeseesSatzsFgtrscaers ss КЕЛ Б 56828 o ® @ E] 2 co n eo cO WW ax о t 3 6 odirigiBoiosziriríoóblii2l22215 O O rums BH p^ NNNM IS EN BEN L Character 25 a Ф 27 Phyllotaxy VE у=? CÛ alternate A ZF ШШ opposite/whorled Il ЕЗ polymorphic ЕЗ equivocal Fides жт, АМА fCYWJCALE. 1 : 1 95 nhvll withi SRM g phyllotaxy. Note that the polarity of this cl the Oldfieldioideae cannot be determined in most of the tree. dri ron. but whether this is a synapomorphy or à Plesiomorphy is equivocal (Fig. 37). Further i : y the hauügblistie s р упаротогрһу о opi eres ao lacking (26: 1). Reversal to tamalo (Fig. 32). 0) unites Mischodon and Voa- e tribe, Caletieae (Fig. 33), is defined walls que synapomorphy of crenulate anticlinal South African genus Hyaenanche, the basal mem- ber of the clade, also is the sole member foun outside Australasia (Fig. 35). We propose main- c A apud EUH ч © © © 4 distribution and gross morphology. We are unable 1 } Зе IN y } 1 £ for the remaining genera in the tribe (Fig. 33) because our data support both equally well. All IN h 1 l a 24. 1 We (97 1), a unique synapomorphy, and depending on the phylogeny chosen, the clade may also be defined by sclerified crystalliferous axial xylem parench ma (21: 1). These genera also share nonverrucate sculpturing, either foveolate (6: 1) or psilate-wrin- kled (6: 4), but whether either of these states can ren | 4 1 г 4 ocal (Fig. 34). In either case, we recognize three additional subtribes within the Caletieae. Reduced or obsolete stipules (45: 1), a unique synapomorphy lel i 1 t l thick phy репар enings (10: 1), convergent in Croizatia, unite the genera we assign to the subtribe Dissiliariinae (Fig. 33). I t h ters are Though our data for tl mplete for these taxa (see Table 3 and the 9n the subsidi incomplet sidiary cells (30: 1, Fig. 31). The discussion above of character 45), for each genus 234 Аппа occa чә Garden E E = x ع‎ 9 e © Б 9 8 c отти EE e238 сәг wEB? ox д © © ee рс ® we ee 7 5 е Do Е $955; о > 9 D е m 5.9154 w.3^7s.E i à L iB e Ix с £ С 2 > © О оф Осу 9 £ o o 9 9 P 5, 9 s у OE а о еж 5 Se 5 Ot С % OF г oF Rs о о = о о 6 OCs SESESSESSSESR ES EPP SES SESE EE >= ¬ ONES ЖА - = 4 з КД P4 s GgebofbgsiktkOodsbp2barfzuzbhioncs22B ES NE E3 E3 E3 EJ Ез EJ EJ EJ EJ Ed EJ EJ Ed ГЛ Ed ШШ OSHA EH E EE а ES о Character 9 Tectum C perf./homog. [3] microperf./homog. EJ microperf./baculate БЕШ microperf./granular EN imperforate FIGURE 38. Cladogram of Oldfieldioideae showing character 9, the tectum. we have definite data for at least one of the char- d DIU ПЛ, h 2: ,and атро; "including a рае ora ore, ria m afio m Austro busus) this is is the paiva demonstration ч the DETTE ari the polarity of this ти at this hylo level i y chosen —" е" кеди AES ng a close relationship. from рт as advocated by imei a Tirel 5 сбой n of its distinctive эма teres a 33), In and its Ke placement (Fig. we maintain Petalostigma in its own subt ribe, the ри matinae. Whether this E or the de o the final Mes the have poll hich the apertures do on the equator. If this is transitional to the eu heec gn dora tei ues Pseudanthus, and Stachys 18 first placed Жо in 1858 (Agardh, em. vir rtually every KE (1965) rec since Kai their pollen, too, is pantoporate. These Volume 81, Number 2 1994 Levin & Simpso Pollen et 0 in Oldfieldioideae [] Celaenodendron [] Oldfieldia [] Aristogeitonia ll Androstachys [] Parodiodendron [] Picrodendron О Croizatia i FIGURE 39. as evolved twice, once in reinfor : а ce our earlier id rae that —€— e Pertures evolved іп, ntly in Androstachys the pe palyn ological data, ipm ies es » kis aie е abes кона "us. of the e Jam n 7e Decl ч see Fig. 33), notebly. а Е oot-layer (7: 1; Fig. 36) and а mi- granular бой (9: 3; Fig. 38). So ibin x 2 E o 9 S a5 2 ع‎ S 2 275 с С c 2 о x - E E SEES 20 E с 9 5 چ‎ 2 2 a c a £ © со Е v RR. q 9 m os 8 D-39950 > [ge eS D 9 SE $ 5 5 & 9.358 ог б El т Ж cent А оф [ке ие И шг | 8 EH E B | ES] Character 3 Aperture position С zoni-aperturate ШШ pan-aperturate Cladogram ( of | Oldfieldioideae da character 3, pon aperture position. Pan-aperturate pollen he Pseudanthinae wood has not been examined), far as is known (its e Eu uphorbiaceae (Hayden, 1994). Also unique than the radicle (23: b a e shared nthus Stachyste- nus) should be e its embryo. Like other Pseudanthinae, Scagea is monoecio On the bebe of our data Neoroepera appears banksii Los ih o be diphyletic, in that N. ese Pseudan ynapom: lacking in N. buxifolia. With Kairothamnus, Sca- k Micran theum, pede iud and Sta сө из foo то ы d: Fig. sounds mer e ar tectum (9: 3; 38), both unique within the 236 Annals of the Missouri Botanical Garden EF and with the last three genera shares ified ph imary leaf simple with foliaceous stipules BIOGEOGRAPHY e phylogenies shown in Figures 31-33 fit rypetes, and Podocalyx, the rema rican endemic Hyaenanche, are all Australasian. ining genera RELATIONSHIPS WITH OTHER EUPHORBIACEAE It i is clear that Oldfieldioideae are derived from en, 1994; Web- ed the pollen unn: cture of e enough ENN particularly the rela a did study tween these paa aod the Oldfieldioideac. К this inii relatives of the Oldfieldioideae are incid ong the relatively basal Phyllanthoideae. Аз pointed out ur Webster et a mblance betwee foot- ах (7: О) like other Мий йа Fur. thermore, pollen of A. guianensis is intectate: the verrucae and round-ended spines—or baculae— f not Pp a oped A m between this Жз ies nd . Instead, pears to be a derived s otherwise has tectate- d te pollen Levin, in prep.; 1965; Punt, seen, for ا ا‎ i in Pi strobilacea (Fig. 15). RECOMMENDATIONS FOR FUTURE RESEARCH This s tudy confirms that pollen studies and 5 data of systematic Kart for the Euphorbi eae. Nowicke's иду of Crotonoideae provides additional evidence for the u of ata. Given our ignorance of pollen ultrastructure t Re most of the family, we ommend with general surveys of the r ining three startin, sub families, paying аар attention to E basal genera in rious lineages i com sahen r po ein en data xiii the excellent data on the Bis morphology and anatomy ыы by Hayden (1980, oe we have been hesis for the phy- ortunate Ei © Et. n @ A @ 5 ص‎ i= | > Ф ч MA Ф a - m 3 с. ч li © [nd ds aston we have f in taxa also Жеб rer s 0 vegeta anatomy beca ese d unav vailable i Table. 3. Critical E x re poo wn. quiring драча study are Kairotharm lid tia, Scagea, and Stachystemon, for all hurt we lack data on wood anatomy. d Voat pe for which we lack data on foliar morphology anatomy Ch ro! known for only 100 me numbers тон in pic Oldfieldio ese s Mischodon 6) cus and Tetracoccus fasc tus (5. т: Pert залы Both have x = 2 а > Le 1943; Sarkar & Datta, 1980), w herea! 3 (io number for most Phyllanthoi ge is ‘2 id 1973). Chromosome counts 0 deae are needed to determine if chromosome ж xa] Volume 81, Number 2 1994 Levin & Simpson Pollen Ultrastructure in Oldfieldioideae ber provides an additional synapomorphy for the subfamily, further clarifying its limits. LITERATURE CITED 1858. Theoria Systematis Plantarum. Lund. AIRY SHAW, H. K. 1970. The genus AM Prain in Madagascar. Adansonia. Sér 277085 1972. A second species of the genus Aris- togeitonia Prin (Euphorbiaceae) from East Africa. We > s: 26: 495-498. es on Malesian and other Asiatic те M CLXXV. V. New species of Austrobux: A ДЕ еу їо the whole genus. Kew Bu " 7— —. 1976. New or noteworthy Australian Eu- УН Kew Bull. 31: 341-398. . 1980. New Euphorbiaceae from New Guinea. li w Bull. 34: 591-598. ыы , R. & G. RIOLLET. 1980. Pollens des Sa- vanes d'Afrique Oriental: бшк National de la Re- Nr" Scient ntifique, Paris. s Jd an nouveau genre e d'Eu- 5 cesa de Мез Adansonia, ѕёг. 2, 15: W. W. R. ANDERSON > H. С. Hırs. 1991 olecular ori. p "middle rosid” fa milies: 8(6, vorne dis C «i idu А. 1981. EE ONE and Sr pres ў ing Plants, гга n Мі іп, Boston, Mas sach DAHLGREN, R. ola da 83. General aspects of angiosperm à ; i io: s macrosystematics. Nordic J. Bot. 3: —— m E x 1988. i ont and dm ps E енн доно сора dn En Monogr. Syst. Bot. Missouri Bot. D eae ATT, ni 1967. | А. Lóve € IOPB аг Бей umber и ХП. Taxon 16: 341 ee » M. Mosanco EN d Afri ystematiques sur les Hymenocardiaceae di ie La morphologie du pollen et l'an Didi s ВЕ Etat 55: 5 E J. & P. D. Cantino. 1984. The logie limitations of the ou "i tgrouj чанына roach to cladistic , дота Syst. Bot. т 2-20 02. à В. Ras. 1986. ntribution to t & r of Andrachne re Алу 297- Pollen & Spo ERDTMAN $e Pede 1952. Pollen "Moa and Plant Tax- У. Angiosperms. Almqvist & Wiksell, Stock- = 1.5 А successive approximatio ions ap- sch зак, weighting. Syst. Zool. 18: 374- . 1988. Hennig86, gram nnig86, ver. 1.5. Computer pro- арду by James S. Farris, Port Jefferson Hans, A ا‎ 175. Chroni somal conspectus of the Havnen, W. ceae. Taxon са Ts 636. J. 1977. Com y and sys- tematics of Picrodendron, genus incertae sedis. J. Arnold Arbor. wm 251-219. matic anatomy of Oldfieldioideae Кееш ы ‘np e җе * Ph.D. Dissertation. ——. 1994. WEM ашу of е ars Olden. I. Overview. Ann. Misso ї. Саг D E. S pn. G.L LW im E ауный» nd palynolog of Picrode n: F ence for relationship vih ~~ [re (Euphorbiaceae). J. Arn Arb 1i HERR, J. M, E dh new clearing- squash aa for the ud of ovule аер жар nt in angiosper Bot. 58: wid classification bà the Dicotyledons, 2nd ed., Vol. 1. Clarendon Press, Ox- ford. ————— & J. А. WOLFE. 1975. The bases of angio- bad phylogeny: Vegetative morphology. Ann. Mis- uri Bot. Gard. 62: 538-589. EEN J. 1969. ines in the family Euphor- biaceae. Amer. J. Bot. 56: 738- Jussieu, A. H. L. DE. 1824. М Euphorbiacearum Ge- eribus Me dicisque Earumdem Viribus Tentamen. пег Didot, is OHLER, E. 1965. Die анон ишы der Biovu- laten Euphorbiaceae und ihre Bedeutung fiir die Tax: onomie. g ol. : 26- 120. KUHLMANN, J. G. P Hylea. Arq Ins ge iol. i 2: 83- 89. 40. toriais. Anais Re- ul.-Amer. Bot. 3: 78- = mai? d + 1976. Recherches sur la natur ur angiospermienne: [шеггп pe struc- tures fe un grou| in yin ci i rbiaceae. ompt. Rend. Hebd , Sér. D, 28 -150. Levin, С. А. 1986a. Systematic foliar = of Phyllanthoideae ати e I. Conspectus. Missouri Bot. С: 6b. matic liar morphology о of Phyl- Sys‏ 19 ,ہس lanthoideae ‘porns ae), ч. fog tic analysis. Ann d ar Systematic foliar энең КИ a of Phyl- C ыйойче “Euphoria rbiaceae). Ш. Cladistic analysis. Sys' ace 5-530. Systematics D oo (Eu- = ре Во!. 17 Pogot rela- PSON. ү d" and Нут теп. rd. tionships 0 ocardia (Eu- phorbiaceae). Ann. Missouri Bot. Gard. 81: 239- 244. Mappison, W. P. & D. R. Mappison. 1992. MacCla — is of idein and d Character Evol тезә ae ue КМ J. зви sims & D. R. Мар! POM. Outgroup analysis and parsimony. Syst. Soak ot MARTIN, H. А. 1974. The identification of some Ter i longing to the ins mily Eupborhiscese. ATOS Anat, Р. A. DE & A. DE iron Fino. 1984. rutura das madeiras brasileiras de Dicotiledó- 238 Annals of th Missouri "ls Garden didi Euphorbiaceae. Rodriguésia 36: 25- MEP ERSON ‚ С. 1985. Scagea, а new genus of Eu- phorbiaceae from New Caledonia. Bull. Mus. a t. (Paris), sér. 4, sect. B, 3 S. Mackee (editors), Flore а Dépndane 14: 1-226. u abe ea Nat MENNEGA, A. М. W - blonskia has emen Syst. Bot. 9: 236-239. 1 87. Wood anatomy of the Euphorbiaceae, in particular of the subfa 2 Phyllanthoideae. J. Linn. Soc., Bot. 94: 111- 126 ME С. R. & L. CHALK 1950. Anatomy of the Dicotyledons, lst ed., Val. 2. Oxford Univ. Press, Oxford. MILANEZ, F Anat me de Paradrypetes il- ede pus vana ‘Biol. V : 133-156. Nuvi: J. 1994. А smi d study of Croton- oidea e En Dedi Ann. Missouri Bot. Gard. 1s 2: Pax, F. & K. Hor. 922. Euphorbiaceae-Phyl- ‘lanthoideae- “Phjlntheae, y А. Engler PETAS Das Pflanzenreich, € V (Heft 81): 1-34 ce horbiaceae. In: A. En- gler & K. Peat ‘ction Die Natürlichen Pflan- темава, e PERRY, B. A. EER ر‎ and phylo- genetic ае н зү in the Pu Meer Amer. J. Bot. e 521-543 POOLE, 981 Pollen diversity in Zimmermannia (ізбен) Kew Bull. 36: 129-138. Punt, W. Pollen n morphology of the ауан асеае pats wi. reference to taxonom 7: 1-11 980. Pollen pi sa d E the visti species кне е) о іп New Guin Rev eobot. Pain 31: L 155-177 e survey ll in Eu- phorbiaceae with special reference to К, , Bot. pet 127-142. n. Soc. Rapcuree Surra, À. Buphor biaceae (Part 1). In: dris (etter), Жозе of Tropical East а. i А. В, а, Rotterdam. . 1987b Not on i Madagascan gei I. On the identity ч aragelonium and on the af- finities of Benoistia and PADOMEI (Euphorbi- Mere m Bul E 625-647 Notes on Madagascan Euphorbiaceae. ПІ “ siachyandra. Kew 2 ll. 45: 561-568. RADFORD, A. E N, J. R. Massey & C. ELL, 1974. feo Plant Systematics. Harper & Row, New York. REITSMA, Т. 1970. Suggestions towards unification of descriptive terminology of angiosperm pollen i 60. са KS A. 1980. Cytological assess- ment of the family Euphorbiaceae. П. Tribe Phyl- lantheae. Proc. Indian Sci. Congr. Assoc. (III, C) 67: 48- a SIMPSO! С. & С. A. Levin. Pollen ultrastructure of те E Euphorbiaceae. Int. J. Pl. Sci. (in p ress). PURR, shee ік electron microscopy. J. Ultrastruct. Res. 26: 2b: te P. C. 927. Celaenodendron. Pp. 16-11 n R. S. Ferris E, Preliminary Report on the Flora of the Tres Marias Islands. Contr. Dudley Herb. 1: 63-88. STEVENS, Р. К. 1991. Character states, morpho ological variation, and phylogenetic analysis: A review. Syst. ot. -5 SWOFFORD, D. D 1993. AEG Per Analysis Using P. n omputer program dis tributed by the bei Eus History Survey, hampaign, Illinois. d TAKHTAJAN, А. ine of the classification 0 flo is 25- owering plants Re л. Bot. Rev. 46:2 359. THORNE, В. T Proposed new realignments in shes Bot. 3: 85-117. the ап im erms. Nordic J. Bot. АЈА 1975. The bean Ga WEBSTER, С. Ts The genera of Euphorbia d ne in the SES United States. J. Arnol 48: ae 430. 975. Conspectus of a new classification of Бас Euphorbiaceae. Taxon 24: 593-601. e Jablon ipis a new genus pc biaceae боп South America. Syst. Bot. 9: ii 987. The = a the s rs e seni за and rotis nships in e Eul J. Linn. Soc., Bot. е 1994. Ѕуп du or the suprageneric er Euphorbiaceae. Ann. Missouri Bot. Gard. 81: 00 ————., L. GILLESPIE & J. STEYERMARK. 4 981-4 tematics of dn (Euphorbiaceae). ae 1-8. An sunt Linnean an Бене WILEY, E. О. ш me We 1979. with comments o o tems. TUE Zool. 2 and practice Р 1981. Pigeon Te ME Sons, Ner PHYLOGENETIC RELATIONSHIPS OF DYMOCISTUS AND HYMENOCARDIA (EUPHORBIACEAE)! Geoffrey A. Levin? and Michael С. Simpson? Жы | Fe | TENERE | ABSTRACT The g us Hymenocardia has been placed either in E and t und ith the lacks the microchanneled t ctum, gr. 1 ma subfamily. Didymocistus, which has H са Ymenocardia, should be transferred [т the Phyllanthoideae-Aporuseae to a position near чеин, Urticales, artists the ee However, the ollen kenin nocardia instead п exine wall o nd ехїпе sculpturing an ucture similar to cardia. | T relationships of Hymenocardia Wallich e Shaw (i ey have been controversial ever since p aw (1965) segregated the genus as its own fam- rdiace жш (1985) ie also ac- 40) made w ribe Le that Webster (1991) nere re- d the Aporuseae баваць" Antidesmeae: for convenience we evil О refer to this group as the Aporuseae.) ree | MET ҮТИ, us of sori ae a на DIEN ge ae an sits beat stomata, all s ies of pomorp. genus and Hym menoeardia = relatively sie leaf gap which in n linked th M. велеа with som lantheae. We have undertaken a review of the morp ine literature esie р р енеш propsebve Ec упа: g We have also obtained new data on the with е s ill show, these results lead us to оле that Didymoc d enocardia are closely re- lated members of the Phyllanthoideae. MATERIALS AND METHODS mined pollen of Г: рш denius Kuhlm aet & Torre ‚ MO), Hymen a Tul. (de Wilde 20 MO), H. ulmoides Oliver (Lebrun 2119, МО)! ( (TEM). Details of specimen тиске сап photogr in a POM к к Blevitt "es теч and Nicki Watson helped with specimen preparation, sectioning, p P. О, ho 1390, San Diego, California 92112, U.S.A. ‚‚ 607 East Peabody Drive, Champaign, Illinois "De Gud mu nt of Botany, San ecd rede History Museum, 61820 USA. Center for Biodivers Departmen of Biology, San Diego State University, San Diego, California 92182, U.S.A. ANN. Missouri Bor. GARD. 81: 239-244. 1994. Annals of the т, Botanical Garden found i in Levin & Simpson (1994, this issue) and evin (in press). The Hymenocardia ed to represent the two subgen- 7). sad (Enti R.755, RSA) using LM and SEM. Leaf frag- ments from dried herbarium specimens were firs rehydrated at 60°C in 10% Aerosol OT for two days, then stored in F.A.A. (formalin/ acetic acid/ зн a For LM иер rehy- drated leaf fragments wer е paraffin crophot-FX photomicroscope equipped with a camera lucida. servations, the rehy- йе ME fragments were prepare ed and photo- ше pollen (Levin & SE 1994). RESULTS AND DISCUSSION Those authors who have segregated Hymeno- male Гоа, which lack both ped and a disk, “de cidedly *Urticaceous' ous' in appearance а adele, Smith (9870) re- like some Ulmaceae. The flowers of Pec also lack both petals and a disk, as do flowers o many орык апа Ulmaceae. Given the striking reductio: et the 9 ONST, it is diffi etermine whe ds: mon ancestry or convergence. The winged fru its of Hymenocardia also r ble those of some Ulmaceae, notably Нараа. Plancho: on (Airy Shaw, 1965). This resemblance is no d at right angles to the ern, and dehiscent, whereas тына of Hol- gl BEE а, and almost ‘tans uni thermore, carpels of H ys uniloc gs ur- Hymenocardia are ge ulate uet seg of Didymocistus and other Phylla an- ves in of Didymocistus are bilocular and de: hiscent. and ivabl 14 rie b 1 һа ee | ] CP 6 Ec AUN Phyllanthoideae and the unusual fruits of Hymen- ocardia. Pollen morphology has also been cited as evi- dence for a relationship between Hymenocardia c and Kohler other Phyllan- thoideae they examined (neither studied pollen of Didymocistus). gums one (1967) was the first to epit his ше Oplata шрот e pollens is e| (Ulmaceae) at least with LM. Dechamps et al. (1985) oa Hymenocardia and Ulma Our TEM studies n E this ыш ity though а & Tiles (1986) showed that the exine s has a micro- strik ing foot-layer. It is very unlikely og moderately ies would be found in a close these character states relative of a Ulm macea Exine scul к structure of Didymoc Figs. c n is quite similar to that " зуп EOIN ie Like o reflect increased adaptation for wind etaining menocardia i in the Phy llanthoideae mr ма have well Volume 81, Number 2 Levin & Simpson 1994 | i Didymocistus and Hymenocardia FIGURES e (Figs. 4-6; Dodson £ pape acida rn 1- аз de Wilde 4044, MO. congue € sadenius — micrographs. Arr " orre MO.— 4, 5. Scanning electron eb —3, 6. Transmission elec ow in Figure 6 ао "Bde Scale bars in 1, 2, 4, 5 = 1 um; in 3 a nd 6 = 0.2 2 ит. Меппе ga, 1987). sere wood is found in genera fibers and abundant t axial parenchyma, and gen of P уре wond (Меш wit манне: Мын PM rm perforation plates ple esiomorphic condition it zer > th not рукн ta I n Contras t, wood of the Aporuseae has non-septate 242 Annals of the Missouri Botanical Garden ‘menocardia acida leaf trichomes (Fig. 7A, B): Enti R.755, SACAR ion througl FIGURE 7. Hy leaf trichomes (Fig. 7C, D): Davidson 5356, RSA © relationship leaf archite he gen- ood, particularly gen- placed in the tribe У ба, с). е also found that both Hymenocardia and 75, 1994) er : Phyllantheae (Levin, 198 i denius A. Didymocistus chrysacen ве Longitudinal section d г axial leaf surface (Fig. 7). These trichomes diffe in size and struc ils. 1 ees (Fig. 7A, B) the trichomes ha and a head 77-102 um i a central region of more or less isod is and an outer region of radially-oriented сез, Volume 81, Number 2 1994 Levin & Simpson Didymocistus and Hymenocardia outermost of which have thickened walls. In Didy- mocistus E, 7C, D) the trichomes have a uni- rate stalk and a head 40-51 in diameter m g of about eight cells. ` Though somewhat different in BE perha a consequence lelike йен are otherwise smapomorphy linking Didymocistus and Hyme- nocardia CD additional lines of evidence argue that Hy- 1 the Phyllanthoideae. Fi irst, its ovules s are Кү рй and epitropous, bite sg crassinucellate, and inserted below a placen tu "Rd (Baillon, 1858), a structure that is bes B (Webster 1967) bu ent pos h \ y | 3 9 the "ciui number is n — 13 as in most Phyllanthoideae (Hans, 1973), a number unknown in the Ulmaceae (Cronquist, 1981). These data are unknown for Didym Chemistry offers some additional evidence re- rur ud relationships of Didymocistus. Runde М Levin (un unpublished ee found that pande Yperaccumulation is a synapomorphy of the Paini Didymocistus, е all other PRS ба. ае Outside de Aporuseae, does пої hyperaccumulate aluminum CONCLUSIONS imilarities with ы в also clear that Didymocistus B E a from each other in ih en natomy, foliar mo y i y that paint rm them is highly unlikely. au included both Didymocistus and Hyme bis a in a cladistic analysis of selec a a Rocistus and e did find that af а monoph a ocardia consistently form Phyletic ¢ de эы pes was the sister group of Tan L. f. and verge a two of the thre nthea t study (see 29). т relation- L. Juss., the 0 ineg KCN, hird member d Phylantheae we studied, are am- 4; Webster, E ) Sn паро meses shared by Didymocistus, Hy- menocardia, and the Phyllantheae vnde. wood with simple perforation plates, septate fibers, and lacking axial paren j leaves with per morphies shared by D. id y. n and утоа d inc guas rugu- highly organised higher-order leaf venation, scalelike fo- liar trichomes, and bilocular ovaries (the last two characteris tics were not included in our cladistic we have reviewed here and the Waster . The ась in turn, hould be placed near the Phyllanthea LITERATURE CITED AIRY SHAW, H. К Diagnoses of new families, 1965. new names, е sig y TA = of Willis's “Dicti ср ё Kew ll. 18: 249 BAILLON, Н. Etude Gé phor a eit ak Masson, Par 1874. Eup iE Histoire des Plantes 5: 105-256. arn aris. BENTHAM, С. 1880. Euphorbiaceae. In: С. Bentham & JD. ker (editors), Genera Plantaru К: berii lower o of owering eoi Houghton Mifflin, Boston, Mas- sachusetts Decuamps, R., M. M & E. ROBBRECHT. 1985. Etudes systém tine ues sur les Hymenocardiaceae d'Afrique: la morphologie du pollen et l'anatomie du is. B . Jard. Bot. Eta 473-485. Hans, 73 d conspectus of the Euphorbiaceae. Taxon 22: 5 I- HUTCHINSON, J. 196 — ^ 9 family Euphor- biaceae. Amer. J. Bot. 56: 7 Конік, E. 1965. ¢ Pulenmorphelogie der laten асарч gee ihre varia g für die Tax- omie. Gra vie lynol. 6: 26-1 Spe J. С. эе novas жин. Anais Reuniao ЗДА т. Bot. 3: 78-86 LEONARD, J. 1957. бин. = iques et cécidolo- giques sur les espéces du nre Hymenocardia Wall. (Enphorbiacées) Bull. ie gs Bot. Belgique 90: 1985. H Iymenocardiaceae. ety M.M In: P. Bamps (editor or), Fl тая азиа . Jardin Botanique cese de Belgique, Brusse 244 Annals of the Missouri Botanical Garden Levin, G. A. 1986a. Systematic foliar aay i of Planes Бараа) І. Сопзресїиз. Апп Miss peat ——. ж) foliar morphology of Phyl- бойне (курошсо da II. Phenetic analysis. t. Gar -98. ematic fla morphology Phyl- ie wa Черте Syst. Bot. of ae). III. Cladistic analysis. а IMPSON. 1994. Phylogenetic impli- of pollen ultrastructure i in the et mier cac Ann. Missouri Bot. Gard. 8 238. Чень, 4 А. 1967. Postglacial vegetation of the i Mountains in equatorial Africa. Ecol Me 3n 25- gr MENNEGA, A. Wood structure Pu Ja- blonskia congesta (ышы Syst. Bot. 9: 236-239. 1987. Wood anatomy of the Euphorbiaceae, in particular wi bs Y Phyllanthoideae. J. nn. Soc., кала C. R. Pus Ln HALK. Mos Anatomy of the Dicotyledons, 1st ed., Vol. 2. Oxford Univ. Pr ress, Oxford. MUELLER, J. 1866. сро, In: A dolle (editor), P ao Vegetabilis Maio 18 Pax, F. & К.Н ы е. In: gler & К. Pra Ty De ا‎ [^ zenfamilien, ed. "2, 19c: bw m 1962. Pollen morphology of the Re e with special reference to taxonomy. T 116: gonads SMITH, A. 1973. A new variety of Hymeno- < acida Ee, Kew Bull. 28: Euphorbiaceae (Part ce ee 1). In: К. M. Polhill (eie. Ie of Tropical East Africa. A. A. Balkem - С ви from the Euphor- ceae. J. Lin dons MPH ae M.G.& m Pollen Вебе ad biovulate Euphorbiaceae. Int. J. Pl. Sci. (in press.) WEBSTER, G. L. e genera of Euphorbiaceae in t in southeastern United States. J. Arnold Arbor. 48: 303 pe Conspectus of a new classification of йе. Euphorbiaceae a 24: SAATANA a of the spurges w of Lag ae ad ‘relationships i A the HE inn. Soc., Bot. 94: Synopsis ч po genera and suprage- neric taxa of Euphorbiaceae. Ann. Missouri Bot. Gard. 144. 81: 33- AVADA, M. S. & W. L. Сак. Es Investigations of оао from t e Eocene of No e America: flowers of ay deco Amer. J. Bo! 68: “933, & D. L. Dic 1986. Comparative " mor; boe and its үзе ыис to phylogeny ої pol len È n the Ната melidae да МЕ i Bot. Gard. T3: 348- 381. A PALYNOLOGICAL STUDY OF CROTONOIDEAE (EUPHORBIACEAE)! Joan W. Nowicke? ABSTRACT of 69 species Shenton 34 genera from 12 of the 13 tribes of Crotonoideae sensu Webster was examined in ШЫ, microscopy (LM), s canning electron microsc opy (SEM), and Wansiniasion electron microscopy Аун Тһе pollen of the vast majority of age subfamily is inaperturate, hitecture: Us ar supratectal elements attached to a network “ muri having vm or поез columellae. These three ponents, triangular su sentem elements, muri, and modified columellae constitute the Croton structure. The phe ar ele: ee designate d subunits, can be psilate, striate, furrowed and d or pitted, and with echinate, nte n ong-attenuate apices; the subunits can be closely spaced or arrayed in more open c ion: bia dai refers to the presence of triangular sculpture elements, whic! T not have muri an ilu co rg cn Inaperturate grain footl d exines. a colpate pollen has зад wa = exine well deve eloped; їп porate grains, the foo d and endexine appear intermediate betw va and colpate. Tribes Micrandreae (Hevea, Micrand. ; Micrandropsis examined and Adenoclinene e, Ditta, Glycydendron, Endospermum, Klain ennt, Tarore im- itive, haying retained not only aperture s but (prominent) footlay nidos; ues endexines as well; ‚ Manihoteae (Manihot, The thick y be ا ا‎ retention of apertures. The remaining | 22 genera examined, including the two largest, and a tectum grains, Croton ай Jatrop ilm is found in some Acalyphoideae. In both morphology and exine structure, the pollen of ngular Gotoniden closely resembles the (porate) pollen of Thymelaeaceae. In a famil y as scd and diverse as Euphorbi- aceae (300 gen cies), digi: in expected pollen mo a ollen the tax- oe llowed the treatment x & Hoffmann (19 i^ rec- subfa Rim Crotonoideae and 987 ре E e t elev ads three Cro- 'onoideae Aa Xy ily status, Oldfieldioi- and RA RNR in a бш о Phyllan i ide eae and the remaining eae, to represent dis best concept of the family to date. In part, auch data support Web- ster's classification (see Disc Ву s egr egating the above three tribes pom Cro- dea tonor form a much more natural alliance that may or may ne ii monem en ic. u Webster, comprise 1 s and «ротах rs E genera, of which doe ара culate, baculoidte, or spongy la layer." He noted turate апа spheroidal. Punt (1962) examined at l thank Carol Annable, Christine Begle, and Lura Williamson for technical assistance, Susann Braden and Walter t. G rady воч с ur with s the lat Pan ad T 1, n ан provided Background i iyi ig | joe kv can = ransmission e a arc thenaultii, 0, C. texensis, and Joa aphs of enia adenanthera, and Ricinocarpos stylos Skvai t electron etn ip of D.C. 20560, ANN. Missouni Вот. GARD. 81: 245-269. 1994. Annals of the Missouri Botanical Garden least 50 species that are referable to the reduced ype that is Sonde on was delimited = (957 ps “Cro- Glass slides for ud were made with glycerin jelly and sealed with p For SE M racetlyrd poll ps ранк са J t Afi : 15 р Вч м LI nes tud are usually clavae but can also be echinae, baculae etc. S ti the st t l tsa H А | Mani- hot saxicola). Tecate.” Punt r reserved “Croton e" for the mended. рее, ро: poten ека pictormed thin from li na кадир а which was all that was avail- dtman (195 Crotonoideae provides a rare opportunity to com- SOM PEN : E + + Ж: perturate g ted Sixty nine species were examined in light PM AT (LM) and scanning electron microscopy (SEM), a 69 were embedded and exa PATER in 1 ру (TEM). In аа- dition, representative кысу» Acalyphoideae, Euphorbioideae, оде апа Oldfieldioi- were examin ed and i are briefly discus pea A ег CO. s ырым were examined and photographed i in сгоѕсоре. s incorporated «id ти юе stained Jw ккан Acrylic Resin. After sectioning Me disi citrate, ud be examined ud photographed with a ick EX ained ү ааа ig and glass slides are dps icum at ils Palynological Laboratory, National Museum of Natural History, Smithsonian Institu- tion RESULTS e Results are organized as follows: a Crotonoi- deae pollen description; d— of exine structure for i and с х te pol- = the ear Picea etn ji in tecta, s s, muri, eee footlayers, and endexines; ix a brief discus ion of pollen in each of the 12 t ribes ex- CROTONOIDEAE, FIGURES 1-88 n mostly spher ik occasionally 5 mu es panto operis t — р ct e; inaperturate, some rarely 3- -colpate, on ds з i ai e ia an Тһуп melaeaceae Чома Skvarla & Pat unpublished data ds on pollen E oe is also discussed. For ease ue reference, examples of bfamili hvmelae. illustrat d. The thrust of this effort is to pare the ра data set for Crotonoideae and not to provide а ка: search of the Шеги! is Punt, 1987, for references to that date). All species examin ed, with voucher data and figure number(s), are given in Table 1 MATERIALS AND METHODS Anthers ere roared from herbarium speci- mens. lyzed as outlined Т Erdtman (1966), but heating time was re m two minutes to one in a attempt to alle viate oils This Medo had var Б and poss ying degrees of succe e lumina usually with shallow free ihi the muri with triangular o E nits, ubunits frequently inuous Pu nl агг: ays, pu surface pes: pitted, striate, o ed, their apice echinate, roun dot flat, or EINEN di E densely spaced or arrayed in open confi і tio In M, the endex ine mostly very thin, irregula: ar and discontinuous, in or som metimes lamellae, oF r ( but more consis- continuous; the supporting colum Volume 81, Number 2 1994 Nowicke 247 Palynological Study of Crotonoideae R dos 1 2. ometimes n grains) Jm but we aring to be in two , ог (in porate grains) the subunits (a tangential Mun) triangular to rounded-triangu- lar, with or without grooves сай {азо їп ше ратар етуи amined here (Table 1), inaperturate and apertur- all umellae, constitute the Croton structure. This definition n incorporates informati adial thin s ns show that the subunits are always attached t muri, e.g., Figures 11, 14, 17, У 26, 31, 36, 39, 41 inset, 47 inset, 63, 69, CN 75, e And 88; and Magenta] sections ү is tri ae or rounded- so eg., roe 6, 33, 8, 13, 18, 21, 27, Е ‚ 79, апа 85. en Croton sculpture r efers t to the presence Meu supratectal elements. INAPERTURATE POLLEN, FIGURES 34—88 erturate «ӘНЕ, сап be further bid tlayer r/ e.g., 82, 83); oa a thick, texine in the lumina, Rum more or less solid = bed g he ricted muri et tricted beca in Species the mu 4 e E.$ demonstrate the thin footlayer/ orant charac- teristic of most inaperturate Crotonoidea PORATE POLLEN, FIGURES 25-33 One of the a tribes, Ише rud 25- 30 ). hasi ге gu mellae, n uneven yet Me s footlayer, э a thin endexine. However, only one species of Manihot (Figs. 26, 27) wA one pI (ыо: (Fig. 29 шах characterization і is a preliminary The of the second porate she, SR (Figs. 31-33) hasa КОНТИ footlayer/endexine and is similar to the inaperturate grains (see dis- cussion of tribe). COLPATE POLLEN, FIGURES 1—24 The exines of the colpate tribes, Micrandreae (Figs. à -9) s Alain hd 10 -24), : are distinguish 1l gs > x db 13, UA a recogniza able i deus (e.g., Fi al 14 23), at least under the abi ee Fig. 17), and ed granular columellae (e.g., Figs. 6, 18, 23; unpublished data). 14, 17, w POLLEN MORPHOLOGY/EXINE STRUCTURE At most, hu сш һауе ап incomplete of the outer muri, i.e., the columellae. The tectum y the nág see margins of attached subunits are considered sculpture. Subunit refers to the tr iangular ed el- ements attached to the muri. Their fundamental shape in cross section is triangular, e these el- ements form or can form a continuous triangular .g., Figures 25, 28, 62, and 67. Although uni i 18, 21, 33, modified by striations, e.g., Figures s 24, de 66, idges, e.g. Figu s 43, urrows and r "p шшш изе In some (rotos dene the muri are well devel- oped, e.g., Figures 8, 18, 38, and 39; in others дә аге a e.g., Figures 41, 78, 79, and 82. 248 Annals of the ау Botanical Garden E 1. Species examined, voucher data, and figure number if illustrated. All collections US unless indicated otherwise. Euphorbiaceae Subfamily Crotonoideae ri . Micrandreae Micrandra elata Müll. Arg. Haught 1 Colombia M. kubeorum R. E. Schult. Schultes "4 Cabrera 13922 Colombia 4 M. lopezii R. E. Schult. Tillet et al. 4 British Guiana 5, 6 M. minor Benth. Schultes & eae 13820 Colombia M. siphonioides Benth. Schultes & Lopez 8700 Brazil 1-3 i mia scleroxylon W. A. Rodrigues et al. 6961, NY Brazil 7 Rodrig Rodrigues & Loureiro Brazil Hevea guianensis Aubl. Asplund 13299 Peru 8,9 Tribe 2. Adenoclineae Adenoline bupl ides (Meisn.) Rogers 24550 Natal 10-12 Prain Ditta myricoides Griseb. Ekman 12802 Hispaniola 13-15 Glycydendron amazonicum Ducke Black 148 Brazil 18 Klaineanthus gaboniae Pierre Zenker 583 Cameroon 16, 17 ex Prain Tetrorchidium rubrivenium Patino 556 Colombia 19-21 Endosperinum diadenum Krukoff 4109, MO Sumatra (Miq.) Airy Shaw E. formicarium Becc. Floyd 3452 New Britain 24 E. moluccanum Becc. Havel е МО New Guinea E. ovatum Merr. Rojo 2 Philippines E. peltatum Merr. jon Pi sinh Philippines 22, 23 Philippines us deo MO Philippines ibe 3. Manihoteae Cnidoscolus angustidens Torr. Gentry Mexico latus (Brandegee) Pax Carter 5 y 4569 Mexico & K. Hoffm. C. rotundifolius (Müll. Arg.) J. & M. Breckon 1295 Mexico 29 McVaugh C. sinaloensis Breckon Breckon & Webster 947 Mexico nit 30 Manihot epee bors d Pohl McVaugh 15355 Mexico 25-21 M. brachyloba Miil Haught 1 Col M. esculenta 1 Fosb Carolina Islands . grahami envoize et al. 3177 Argent M. irwinii D. J. Rogers & Appan Irwin et al. 34338 Bra ‚ marajoara Н uber s.n., BAT Brazil M. Rn Pohl Maxwell & qe ideni Brazil 28 19774 М. pentaphylla Pohl Irwin et al. Te 9 Brazil M. pringlei S. Watson Pringle Mexico . tweediana Müll. Arg. ојах s.n., Ni 1933 Brazil M. tripartita (Spreng.) Müll. Arg. Irwin et al. 18650 Brazil emend. D. J. Rogers & Appan M. violacea Pohl Heringer 18246 Brazil Tribe 4. Gelonieae Suregada glomerulata (Blume) Ahern s.n., 1901 Philippines 31-33 Baill. Volume 81, Number 2 Nowicke 249 1994 Palynological Study of Crotonoideae TABLE 1. Continued. Tribe 5. Jatro ide Deutzianthus tonkinensis Gagnep. Petelot 6388 Vietnam 34 Jatropha cinerea (Ortega) Müll. Wiggins 16117 Mexico Arg. J. glandulifera Roxb. Nowicke & Jayasuriya Sri Lanka J. hastata Jacq. Lundell 1577 Yucatan 35-37 J. podagrica Hook. Blackbern E10 Caroline Island Tribe 6. Elateriospermeae Elateriospermum tapos Blume Suvarnakoses 1872 Thailand 38, 39 Tribe 7. Cod Noii um axillare Blume Nicolson 2870 Nepal Codiaeum variegatum (L.) Blume yas 1339 E] Salvador 40, 41 Dimorphocalyx аа Elmer s.n., 18 Mar. Philippines 45 Wee Ostodes zeylanica Müll. Arg. edu > чыз на Sri Lanka 46, 47 nayake Pantadenia adenanthera agam Poilane diee: А а 42 Pausandra densiflora ер cke Brazil 48 Sagotia racemosa Bail Prance et al. 22670 Brazil 43, 44 Mexia 6051, A Brazil Tribe 8. Trigonostemoneae 7 9; panes Iphandia fu Pur 809 New Hebrides roms leschenaui 3 ) Ваш. r 1130 tralia 52-54 Ricinocarpos bowmannii F. Muell. Borman I Australia Bo ustra! 51 R. glaucus Endl. Australia 50 R. stylosus Diels теше 18714, DAV Australia 5 Bertya gummifera Planch. Johnson & Constable Australia 56, 57 Tribe 10. Crotoneae Croton argyratus Blume Toroe. tra C. balsamifer rubi m West Indies 60 C. californicus Müll. Arg. Wheeler s.n., 16 Nov. California 1 C. draco Schltr. Salazar s.n., 11 Mar. Mexico 62, 63 1913 C. floribundus Spreng. Mexia 5292 Brazil 61 С. lachnocarpus Bent Chun 6261 China С. matourensis Aubl Arch 820 Brazil 58, 59 С. mutisianus HBK Uribe Colombia C. pyr LE Donn. Sm. Dwyer 11073 i 64 C. texensis Müll. Arg. White s.n., 26 Aug. 1946 Nebraska 65, 66 Tribe 11. Joannesieae Annesijoa novoguineensis Pax Brass 32333 New Guinea 70-72 . Hoff Joannesia princeps Vell. Macedo 5335 Brazil 07769 Tribe 2, E 19 Rici п rautanenii Schinz Gibson 19 Angola 8 o. po dii Hemsl. Peterson 2805 Argentina Аппа! of the eset Botanical Garden TABLE 1. Continued. Tribe 13. Neoboutonieae Anomalocalyx uleanus (Pax & K. Hoffm.) Ducke Grossera major Pax Crotonogyne argentea Pax Subfamily кааш Drypetes laterif g & Urb Phyllanthus acuminatus Vahl P. Melee و‎ А Р. simplex Flueggea test Mill. Arg. Subfamily Oldfieldi Hyaenanche p (Gaertn.) amb. Longetia buxoides Baill. enr eis Da mpia gens iege Arg. D. icon olia 8 р. & Endl. Galearia celebi a G. filiformis (Бил) Рах С. fulva (Т G. ЕТУ чы ү (Е. Вг.) Miq. Масагапда са Müll. Arg. M. griffithiana Mü = М. tanarius @. )M d erri Planch, rlotiana J. L Panda oleosa Pierre Subfamily Euphorbioideae Euphorbia pulcherrima Willd. Hippomane mancinella L Mabea montana Müll. A Thymelaeaceae Gnidia glauca (Fresen.) Gilg Gyrinopsis cumingiana Decne. orst.) Wil Schoenobiblus grandifolia Urb. Ducke 64 Zenker 469 Zenker 3078 Contreras 1788 Standley 23316 Godfrey SH-1257 Baumann 14418 end » Rizzini 17544 Allen hmat "n Sog 4892 Fou Mie Sinclair 354 Rabil 338 Lei 5 Trou upin 3549 ladik s.n., 12 Nov. 1971 Hall & Amponsah 46547 Wheeler 12310 Killip 41829 Standley 25156 Hoogland 11563 R Anderson 186 Cowan 1548 Brazil 80-82 Cameroon 83, 84 Cameroon 85-88 Guatemala El i 96 Seats Philippines South Africa New Caledonia 92, 93 89-91 94, 95 Panama Ceylon Philippines 97, 98 New South Wales 102 New Zealand 100 Tobago 99 In the Ricinocarpeae (Figs. 49-57), the muri ap- pear to be reduced (see discussion of tribe). тж epore e columellte Ho variable, but in r эпо 1 апа any appear to be in tw d 5 Für s 37, 41 inset, АТ, "34, 57, 78, pee 83. All ү ынады and most Micrandreae hav о e columellae ea 6;. 165 23: айра ДЫ dat n the exines of inaperturate Hag EIE TENE the footlayer and endexine are pe so thin ge be distin, from each otir TE cannot guished proximal dary iei the vini e there are sma granules abo d below a "thread of uet e.g., Figures 44, 59, 78, 82, and 83. Som. all the granules have the same electron et! (Fig. 44). In a few taxa, a eti 1 endexine is present (Figs ; 54. 91). -24 The footlayer of фору (Figs. 10 24) Volume 81, Number 2 1994 Nowicke Palynological Study of Crotonoideae Elateriospermeae (Fig. 39), and some Micrandreae (Figs. 3, 6), is uniformly well-developed, whereas in the single species of the porate Manihot ex- min thin section (Fig. 26), it is irregular. ae and Micrandreae have a dex- a one in Manihot aesculifolia (Fig. f + Pun procedure (see на gia Methods s some inaperturate pollen retained part, if not all, of the Kis oram contents m intine (Figs. 47 inset, 72, and 88). erturate Crotonoideae are known to et al., 1984: f pere oT her : i: : figs. 14, 15) may provide some me- nical/structural support, partially compensat- ing or t the extreme reduction of footlayer and епаехіпе. TRIBAL DESCRIPTIONS Micrandreae, Figures 1—9 үз even copa (Table 1) examined in the ndreae are 3-col i (Figs. 5, б), М. ere examined in thin ps has a a recognizable, у соп- granu a prom- In radial sections at " lopezit, Species is on lack a C he very few in Crotonoideae that чы ea t s sinis pture. In M. kubeorum (Fig. 4) $ " ir d while m SEM. v of eh E od Figure 4, they are in other data), . Webster, unpublished A fourth species, M. elata, examined only SEM, a tectum with triangular subunits, much like ан illustrated in Figures 25, 28, and . A more extensive examination of Micrandra is warranted. Micrandropsis (Fig. 7) also has a modified sculpture—t are finely divided or very reduced, and the network of muri is irregular. ре (Figs. 8, 9) has very small feet Лы subun a aes netw ork of muri with smaller lumina Presta Fig. 6 ig. 8; the latter is illustrated at a magnification five times spec ntinuous ende келде. icem а esc columellae. Adenoclineae, Figures 10—24 Ten species representing six genera were amined. The 3-colpate grains of Adenoclineae on a distinctive E ыла uniform exine: a well- developed. endete e ie frequently detached footlayer (Figs. iy ^ a 17, 18, 23), gran Ei p um, de. 1 hus, Tetrorchidium have striate subunits with echinate pices. Tetrorchid rubri re- ium 3-colporate (Punt, t ere found in the amined here (Table 1). Klaineanthus however, is 3-colporate as demonstrated in Figure 16 and con- firmed in LM. A few tetrads а in Endosper- mum peltatum. Yn thin sect ion, E. formacarium is very similar to the exine portrayed in Figure 23 of E. peltatum I , the Arosa can be easily distin- guished from all other Crotonoideae by ed colpi and thick ех elongate or pou iN nits. The deep subunits are аа notic eable in Ditta and Tetrorchidium These thick exines may be correlated with the es or retention of apertures since such wall pon prove difficult to p e in the ger- i sed pressure might be lethal to the sperm idi Manihoteae, Figures 25-30 Cnidoscolus m: Manihot are characterized by larg tweediana may be excepted) po ollen i Only two species, one in Manihot and one in Cnidos revi Med examined in TEM and any sco Struc ctural characterization is premature. Manihot i e that are tin ges pha yet collectively form a significant co tial muri 252 Annals о Missouri — Garden 1 } JEN S BS S 1 ponent a dincüntinüods. endexine (Figs. 26, 27). The he structure of Cnidoscolus (Fig. 29 inset) is similar to Manih i the presence/reten- of apertures appears correlated with thicker xines In Manihot, the subunits are conspicuously tri- ie., their diana is t some grains PME 15- а аж (5-5-5) нау 15-pantoporate. Gelonieae, Figures 31-33 regada, the only genus in this tribe, com- s 40 species found in the warmer World. The pollen is of interest because it is porate, yet has an exine structure more eser ofi шрек га еп. пер огеѕ of Su sets pia retain a poorly defined aperture cov sma is e- - Аз a result, the b are not conspicuous in — (с; Fig. 981 кш in section, colu- frequently there is empty space between the muri and the thread- like TORTO нетов (Fig. id or iri is are ine. In tangential section, the free Шш ар 2 A ТОШ tribes have inaperturate pollen. Jatropheae, Figures 34-37 In this tribe, only one species, рө hastata (Figs. 35-37), was examined in thin section, but of other э Сго- e subunits аге roun ded i w, uri ver and irregular. Both footlayer and endexin € constitute only a thread with Кинза. =” anules. n & Webster (1979), in an infrageneric ы pr Jatropha, illustrated the pollen of 12 se in SEM (plates XXVII-XXIX), E ing the thin muri as well a al sub se striations; smooth subunit in Jatropha subg. Jatropha sect. Peltat Elateriospermeae, Figures 38-39 e exine of Elateriospermum can be distin- other inaperturate Crotonoideae by veloped, contin layer and thick muri, beth. eas ily к іп LM. In some sections, f th i RS there are some grains with one or two penings, pns abl << e Е о 8 © 5 = mall о olay The ra mination difficult, the presence of apertures 1s not unexpected Codiaeae, Figures 40-48 This tribe reflects it khu. size, some 20 genera, 40, 42, 43, 46, 48). (Fig. ll remaining taxa examined have козы furrowed, or ridged subunits. Co gun (Fig s. bd ud a large lumina obules. Pantade- i n diae- um variega with fr nia adenanthera (Fig. 42) uA Ra large lumina, but me ss) muri are obscure. Sagotia poe subunits үн longs cate tips (Figs. (Fig. "penal tlay Fig 44 Be appears to consist of small gra above his below at кед of exine. Аро election oft atourensis (Fig. collections was d Ostodes 46, faintly grooved subunits ; À ins the widest part above the middle; some gra s 47 inset) including intine. Pausandra (Fig. 48) h lender subunits wt ations. Ricinocarpeae, Figures 49—57 In this small dos of five doer ue ndi, у Вегіуа, Веуег ое the ро ie of some hoo sc be Russ ad derived—both muri and Тыын modified а (Fig or difficult to лаш. Beyeria loma coneri Volume 81, Number 2 1994 Nowicke 253 Palynological Study of Crotonoideae 54) appears to have lost definable muri—the sub- units rest on a trace of footlayer over a thin la- xiii Bes: ефе по Crotonoideae pollen ined in TEM has hollow supratectal subunits, axi if the areas ange by small arrowheads (Fig. Lus are An ed Ai XR muri of a sort, ick pair ‚ would be pres eee. (Fig. =n e shallow muri with paired (үөт In ix append онц (Fig. 49) and mmifera ( 56), some adjacent нв ме fused iis ачы In Ricinocarpos bow mannii lim- subunits ited to e junctions of the widely spaced muri, and most of th re c ed in thin section, Beyer- si Кешш: (Fig. 54) and Bertya gummifera ; ig. 57), show a very жыз sparsely granular foot- ayer and a thin lamellate endexine. n B and in TEM, he exine of Bertya has s s in the Ше, endexine where the upper У texine appears to “unravel” outward (Fig. 57, right), forming a Star protrusion, giving the appearance of por Crotoneae, Figures 58-66 m the 13 species of Croton examined, C. ma- rensis (Figs. 58, 59) is the most ше aelike supratectal elements occur r onl se rows, Clearly, tributio of s n, Mid striation reflects the large number Species in Cro Joannesieae, Figures 67—72 © Species examined in this small tribe, ё- prominent furrows/ridges; t uate tips foo dne (barely visible in Fig. 72) appears to con- ist of a thin] y f yg ] and is subtended thin but recognizable endexine in both tan- evil (Fig. 71) and radial sections s (Fig. 72). The nl in other thin sections is present as a sparsely and loosely granular zone. Aleuritideae, Figures 73-79 n another small шы comprising dry a dozen species, two 5 and TEM. Alerts fordi (Figs. 73-75) has vs units with roun regular, шше us nodendron rauta exine. ndis ASA d within a collection 1), bu the . open улсын (Fig. 7 more common than the d the Жаматы of the subunits in robably from a a more е like that in Figure T than in Figure Neoboutonieae, oe 80-88 In this large tribe of 13 genera, the three species examined, Ano mala uleanus (Figs. 80-82), Gains major (Figs. 83, 84), and Crotonogyne argentea (Figs. 85—88), have striate subunits, well- d i ts e, an Although Шш иш ео of fG. i m ajor гү 83) FM Fig are vr 84) and in other sections. The deep subunit stri- ations in C. argentea are prominent even in TEM (Figs. 85, he pollen morphology /exine structure of the Tt four subfamilies is briefly described. deae, a Webster (1987) pone Анарай аз Асс din E т сн Acaly phoi- nd Eupho to to Punt (1962, 1987), this subfamily has two i lin orate, tinct, yet related, pollen lines: one is 3-colp rolate, a os ticulate without m r- ате the о line is distinctly, often ore or less obl Cer- coa arsely, r reticulate, and the vast majority of Phyllanthoideae pollen M be m M, docume cies of Zimmermannia. The pollen of ha 254 Annals of the hea Botanical Garden Р ч ke, unpublished data), is very лый in TEM to караг rbia Дш £5 ig ul ЭРЕ which in turn со ral Bhadlantkus; however, кеша diverse Mer types as well as a wide array of tecta. Punt (1987: are more endoapertures [th biculatu. and they are usually ed at om." How ever, the collection examined here (Table 1) has is pollen (Fig. 96) ber “> three (ecto-) ap- prominently bo ا‎ ‘colpus” that is joined 1 mid PT bain the rounded ends ese “bowtielike” apertures are placed equidistant from each and, if m the range of variation of those remaining identified as Р. acuminatus. st Oldfieldioideae appear to be multiapertur- ate, brevicolporate or porat ularly thick- ened near ‘the вао In terms of g а 10), if not species (1500) kal 1987). узе hoideae are the largest mil е у and exine structure are unremark thus have limited axonomic value. The same conclusion and inter- ertain m Acalyphoide , Aci doton nicaraguensis ies 1994, "Ыз ау: and certain taxa in 1 tr ibe Adelieae, Adelia triloba, t id tectum-triangular supratectal el- ements, but the elements are small and not always e tectum in 4. triloba does form a continuous triangular array. However, none of the above species have been examined in TEM Euphorbioideae have never been the subject of a i detailed ышчу in SEM and TEM, but Punt hie acterized the pollen as being consistent in shape, mentation, and size. Based o above con- Pics by Punt (1962, 1987), vd Ghobary (1985), and El-Ghazaly (1989), the pollen of Eu- phorbia pulcherrima (Figs. 94, 95) is represen- ta Mise 9 the uu and cult be representa lp Е Sus pus um, an en dex: bd t P ckened fuera the apertures, ber devel oped, even elongate, columellae, and a in foot- layer. In a largely unpublished study of Thymelae- aceae pollen (N ть Patel es Skv arla), 70 spe- cies from 39 era were examined. These рап- poete Logs illustrated here en 97-102) in SEM in Nowicke et al. (1985) in SEM and M. are nete de Sed in exine structure to Crotonoideae: triangular supratectal elements are attached to prominent thick muri with short or narrow columellae, and both footlayer and endex- ine are thin and/or irregular. u B DISCUSSION llen data from electron microscopy support ate, ha n e gular supr dic Mesi calle hec to a network of muri having n or irregular be furthe hreadlike urate exines can mellae. Inapert characterized by a very thin or even t [y adde are ni the уч large assemblage in the cto to have a preponderance of nape turate n. Of the 2000 dias that. Webster (1987) ыгы rae this subfamily, at most only 300 have apertures. д ticular ехше Colpate apertures and a par wet бойун, d well-developed endexine—reintorce the ili regation of Adenocline, Ditta. em Шит Glycydendron, Klaineanthus, idi Tetrorcht as the tribe Adenoclineae and emphasize their close r. cture It is difficult to characterize the exine = pe of the 3-colpate Micrandreae until more tà | Volume 81, Number 2 1994 Nowicke 255 Palynological Study of Crotonoideae investigated, especially in Улым, Of all spe- cies examined here (Table 1), only M. siphonioides (Figs. 1-3) clearly lacks the died subunits, although they appear modified in M. kubeo (Fig. 4). ithin the subfamily, the pollen data suggest that Micrandreae and Adenoc itive, having retaine ith well-developed Ie d endexines anihoteae large genera 8 (Manihot (170 a Cnidose olus (75)) ac- count cl most of the 300 реш Cangi Wes have p umellae, and : a thinner (than Adenoclineae) foot- layer xd endexine. This base on TEM data from only two species, REE and thus sh v be considered as Pee. р f Suregada, the in Ge- lonieae, is poor ate but ma ж ‘mead foot- ids ndexine characteristic of the scri oes 2M Pun Pun ba 962) cla ы the two species he xamine ging to the Manihot type " ri- in 5,10 seemingly absent suppor a d eis further е its HE with other Crotonoideae а Micandrens Adenoclineae, Manihot- С, С elonieae have apertures, they have the ructu; bas Groton re Mac are clearly allied to inaper- ining qe have inaperpurase c S 7), the Crotonoideae exine is Consi iderably iskor. Clearly, the pollen wall of the ТЕМ» illustrated here would sugge Inaperturate ро еп is a reotricted colton i in the dicotyledons subfamily and rarity in ‘the remaining Euphorbi- aceae, would support the monophyly of inapertur- c о tri It piece be difficult to characterize any one iversity of mod- ifications to the subunits might mds оаа but that diversity eae: in рап ш larg mber of ү: kei nde to Crot p ; "Willis, 1966). Some Ricinocarpeae s tov mo: i- fication, a size reduction and loss d deii of subunits. The exine structure of Crotonoideae is more те Һау ins T ЯРА 1989) while Aoknontedging | v тї 1 1 4 late | Ше. assigned uM to the Myrtales and uphorbiales. But dn Punt (198 "m E recognized b “mo the subfamily Аса Iyphoidea š is an с n to this it ‚ at t to me, that eption onsidering all Euphor Eid i. be pollen є Oldfeldioideae, С Crotono = Pod pollen & 4 eralizi in the thi rd; Plots and ctii oideae have diverse pollen . In Oldfieldioideae, the miel is СТЕЙ spinulos with a thick tec- tum colur n footlayer, and well- dev ram und endexin e. The pollen of Cro- Croton structure and is mostly tonoideae has the i llen of Euphorbioideae, un- phology certainly doe: rgue agains ent concept of гад Pollen siet aa 256 Annals of th Missouri iwi Garden ns к these end iege он are natural ages. How o judge the taxonomic e Ф @ = 8 е o дай esi in Phyllanthoideae and Acalyphoideae? Probably on a generic and/or tri as much pollen diversity, e.g., Polygonum (No- ke & Skvarla, 1977). LITERATURE CITED ся . An Integrated System of Classi- fication of е Plants. Columbia Univ. Press, N "Хос a 9. е Evolution and Classification of Flowering Plants, 2nd ed. New York Botanical Gar- den, Bronx, New , G. TER. 1979. bei ea and infr; rageneric relationships = the genus er "^ и phorbiaceae). Uni Publ. Bot. sane pons G. 1989. p» E and узы cin эсш me Euphorbia species. Grana 28: 243-259. 1952. Pollen Mo чу 24 ology and Plant Тах- my. Almqvist & Wiksell, Stockholm. нэа Pollen Morphology and Plant Taxon- Angiosperm: "Wir i Wiksell, Stockholm. Gust L. J. 1994. orphology and ke eny of the tribe Pete (Euphorbiaceae) Ann. Missouri Bot. Gar KGHLER, E. 1965. Die ” Pollen morphologie der Biovu- pet psg ue und ihre Bedeutung für die Tax Gra 1 6:2 nomie. a Palynol. 6: 26-120. Kass, W. J. & D. E. TO 82. Nature of the ки їп monocotyledons, with A E ую and Heliconia. Grana 21: NowICKE, rg. 1984. А маар "e of the Pandaceae. Pollen 5 uie c Io Ev lle en morphology and the idis, ^t the Plumbaginaceae, Poly- 9 5. cation of athe Борне Tax ل‎ The of the PE and — М pes npr: Centrosper- Mur Ter Contr. 1-64. L& ла ORAE P the relationships of Aetoxylon, Amyxa and ey iat us to the Thymelaeaceae. Amer. J. Bot. 72: Few .&K E rer NN. 1931 D: ae. Pp. -233.i Engler & К. tl (editors), Die nan lichen i dus A. M Vol. 19c. Wil- helm Engelmann, Leipzig. OOL. . M. 1981. Е bui M eges in Zimmerman- Pollen MALE of Euphorbiaceae with special reference to taxon пошу. Wentia 7: l- 987. олаи É edid reference to gw . Soc., Bot. 94: 127- ROWLEY, 128 & J. р SKVARLA. 1986. Devi of the pollen grain wall in Canna. Nordic J. Bo STONE, D. E. onv 1987. Developmental evidence for the ma of S ) d Heliconia Zingiberales) pollen. Grana 26: 179-191. сана G., С. CanariNI, S. NILSSON & E. G 5ТЕ Omniaperturate Euphorbiaceae pol- ies vit diritto spines. Bull. Jard. Bot. Belg. 54: = о аны n Снов ү, М. О. 1985. Pollen morphology of four succulent Lao of Euphorbia AE aceae). Anales oa Мп а Ез WEBSTER, sp фы A revi oki and relationships in the га J. Linn. Soc., Bot. 94: 1 . Revised EER of the Euphor! Euphorbiaceae Newsle ў А Dictionary ‘of the Flowering m Н. К. Airy Shaw), Tth . Univ. Press, Cambridge | Volume 81, Number 2 | 1994 | Nowicke 257 Palynological Study of Crotonoideae IGURES 1-93 İS a very r d 9^ SEM a EM of pollen in tribe Micrandreae. 1-3. Micrandra siphonioides . Tectum. This е example bus a Meneses speci cies in which the muri lack ipe най subunits; this en. ‘could ut the absence of any clear vestiges of the Te "present а lo ss of the subunits stage wie to their дейри, ol. subunits wo 1 rm Radi al section. Noe prominent footlayer and Well-dey e Pport the later interpretation —2. i 8; f Brain. Althou arrow).—4. M. — K i 5, 6. M. lopezii. —5. Whole of subunit d not apparent on this grain cis species is 3-colpate —6. Tan eig section. Note: triangular shape Shaped Co ete section; muri (arrow); Ё nular di (arrowhead); a fet cut ut obliquely ( ак mow Table 1) has solid area). — 7. Marque is oxylon s are finely ; a sec аге Prominent tectum more similar to that n Figt 29. 8, 9. evea en sis. — 8. Tan ngential section. The Subunits and the very small lumina appear —9, Tectum with very small маном Ма are p 'ated by a straight arrow; supporting columellae by an arrowhead; free columellae/globules of *ktexine by Y а wavy arrow, All scale bars equal 1 micron unless otherwise indicated. 258 Annals of the Missouri Botanical Garden ата from -— ron also Figs. 14, d 12: Te ctum. 13 EE Ditta кенне; 13. сш structures, in order from the outside, are Pur эн subunits, , then solid | im Ж muri, dd columellae, did T yer; rue seeming ly . Ra dial чаи er Mert F poilayet, i Fi 7 i with ndexine.— 15. Tectum. 16, 17. Kb бона, —16. Тыв was the only rues canine тб olporate аже rtu —17. Radial Tange ential had a fragm зё чиде аг endexine similar to that in Figure 23.—18. Glyeydendron amazonicum. section. Structural components are same as in Figure 13. en = endexine; fl = Volume 81, Number 2 Nowicke 259 1994 Palynological Study of Crotonoideae Ficures 19- 245 SEM and TEM of pollen in se Adenoclineae. 19-21. Tetrorchidium rubrivenium. —19. = um at high m magnification. i are prominent, subunits are deep/long, and most lumina are covered "m in int glob les. —20. Radial section. Althou muri are poet Че ейпей, шу аге sah present in Figure 19 an the tangential section in Figure 21. hi Tangential s nclude gran nular columellae ^ payer. 22, 23. ndospermum peltatum. ni^ 22. ho ain, polar view. de minor portion this sa ро Radi Table 1) appears to ab been shed as t gat but the caret mbers ret arying degrees of cohesion E Оп near aperture. Note: endexine layer separating from ektexine; thick irregular cds n l secti со] "Poser еч od — 24. E. formicarium. Tectum with variously у striate subunit: 260 Annals of the Missouri Botanical Garden FIGURES 25-33. SEM and TEM f pollen in tribes Manil (25-30) and Gelonieae (31-33). 2 27. Moni aesc ulifolia. —25.T 2 А ап 5) a have det sparse shallow на = ektexine. ‘Of all genera examined Dre Manihot has the most ie an tr es gp subunits. — 26. Radial section. Arrows connect (fro top downward) кик columellae, muri, and {сии this radial section bee. Чы counterparts in the tangential meat ion in Figure 27; "e порае columellae are irregular, ev ar; note discontinuous but recognizable, . Tangential sec as See ү” gend of Fem ure yo 28. Manihot ping Note pit aem surface of о а char: teristic vici ye Ман: species examined here.— Pd Cridosc olus rotundifolius. Fracture illustrating relative thick foot? ting endexine. Inset, C. sinaloensis. Note — columellae and irr al footlayer; in other sec ctions part ada columellae appear similar to those in ga e 26.—30. C. sinaloe Tectum with ad ore. — 31-35. — ri; in leveloped m" glomerulata. —31. Radial sections of two o grains. There is a paucity a аар below the well-d Volume 81, Number 2 Nowicke 261 199 Palynological Study of Crotonoideae n in Bis dor at Uum 37) and ягана (38, 39). — ur -37. Jat abe hus a.—35. Shoe The large size and den nae p = en Muri ar е muri; some subunit appear t o be connecte thin ra section. the Б dim Mir the ( SLT "Coria ^ h a few g anul i are fi iln 37. Ta angential section. Small Weare 11 With conspic olumellae (s (see Fi ig. 35); note ра виррогї шш ellae. 38, is teriospermum tapos A . Tec ^ Suggests Radial section. The gradual jets: (left) of the rip foot D з а preformed aperture Ра Орго es a break). However, apertures could not be verified in LM. ~ both Brains the zT dor А reduc tion of sins н proximity to an aperture, but even here the scart is s thread 'overed with ылы, free llae/globules of ektexine 109, чи section. upper ike. — 262 Annals of the Missouri Botanical Garden Ficunes 40-482 SEM and T in tribe Codiaeae. 40, 41. Codi gatum, —40. Теша open vor irregularly ei subunits itl ss 2015, ‚44. sie. Inset, radial section. —42, Pant Pc adenanthera. 'Tectum with irre сштш muri F and ire lum th е are Sagotia racemosa. —43. Tectum. This is а of the more e н їп е! su gun т wr areas where the subunits fuse together. A simi us ar tectum is present in Annesij os. жег tri. po of Oblique section. Mis section emphasi zes е unusual f granules ve and below 62 densely exine.—45. Dimorphocalyx luzoniensis. 7. Ostodes z Bis fie ca. —46. um. Although subunits il section. spaced here, E grains had a € onfi uration; note faint furrows in subunits. — 47. Ten d g Inset: radial section of portion of i incom n күге, жн area between open arrowheads is Pausandra densiflora. Supratectal subunits are deeply st Volume 81, Number 2 Nowicke 263 1994 Palynological Study of Crotonoideae К Е. в а 0 P. SEM and TEM o eia in tribe Ricinocarpeae. —49. Aiphandie fury Ficures 57 urac ). Ricinocar- pos ~ a Whole grain, N collapsed.—51. К. bow mannii. Tectum. This species and ү; oR б hena i 39) have the mos tal configuration of all Crotonoideae са Бес -54. s dde d "Mos 74.52. Whole са о) ectum. See discussion of еп 4.—54. Radial section. Altho apes the кы. Fest оп a trace of footlayer ond thin lamellate endexine, no subunit examined i in TEM i is йы thus be y 5. R; i i enizable. - 55. Riein P portion is ihe subunit. If the gaps ‘onal arrowheads) were ке ed up, the mur p cvm shows o "ocarpos stylosus. This species has a typical tectum. SO 94, а gummifera. —5 ils y й: д specie ne of the more modified Croton structures — many subunits have fused ote Moe collection a ida muri; granu loot similar to the . Radial section. Note: p layer: and thin, but well-delimited pele aa 264 Annals of the Missouri Botanical Garden FIGURES 58-66.3 SEM TEM of pollen in tribe Crotoneae, Croton. 58, 59. C. pe от = 58. Whole grain. One of the most distinct рө, morphologies in Crotonoideae, but a similar опе 0002 Шш papa bowmannii (Fig. 51).— apes үүтүн fine granules above and des qeu in, irregular footlay ine; t lid ts at mde ‘and pat muri cut at right angles to the ir Neves axes; at right is a la arge subunit cu ] section; although 2 61 C apparent at this magnification, the thick muri rest on granules or E. "Lotus — 60. C. balsamifer. ~ s ol тавоне Hus lum a have scattered free columellae. at 63. C. draco. —62. Tectum.—63. Radial ен гаїпз . С. pyramidalis. The rounded apices and d oft ча ебе ex are similar to Figure 61. E 66. rae Vise — 65. Tangential section. The thin mur! $ е5 evidence of paired columellae.— 66. Tectur Volume 81, Number 2 Nowicke 265 1994 Palynological Study of Crotonoideae Ficures 67- ida _ SEM а nd TEM of selle | in tribe Joannesieae. 67-69. Joannesia princeps. —61. Whole res pie з 9. Radial section. Columellae here are = delimited; other sections shov ar s the d or below a а qme of footlayer. 70- 72. Annesijoa novoguineensis. —70. Tectum. Even though obscur у sha Т? ee surf , at least some аса (those іп frontal чеч) show the basic triangular alm Fu 7l. Tangential aine Even i int angential section, which enhances the thickness of et the в footlayer is secti Onexistent, but the xine is ide ntifiable as the aic асаан less electron-dense layer.— 72. Radial 90 of portion of e r acet olyzed grain; area betw open à owheads is as — E be intine; note thin, but co, "sistent, lamellate endexine; muri are much better deo m than in Figur 266 Annals of the Missouri Botanical Garden ain. FIGURES 73-79. SEM and TEM of m in tribe uiui) 73-75. Aleurites РЕ Dm T5 m 74. Tectum.— 75. Radial section. The foo ait ndexine are T thin, but Mw. ut pollen ge and a trace of footlayer. 16-79. "Ric nodendron rautanen 76, 77. fecal s Tie 7 than Figure ample. —18. Radial section. The spacing о оГ ts sugg hat thi grain is more like Fig зе" ^ ntial section. Mun are very thin; footlayer/ endexine are highly irregular. Volume 81, Number 2 Nowicke 267 1994 Palynological Study of Crotonoideae Fia RES 80- 88.2 &rain.— 8] T, SEM and TEM of poleni in tribe Neoboutonieae. 80- va Anom [ 1 =< оче ection. eee Radial : sectio 83 4: Grossera та ira е the lumina h , Spacing of the subun idis the pattern in Figure 84; the muri x сме p oring «== — reflect ui much йай sible x eee, 85-88. Crotonogyne argentea. —85. Tangential section. Sub e section ; ер striae illustrate d in Fig e 87. Бозй c el near center are free columellae i n cross section; the lke Fı n not deep enou 86. Whole grain. —87. Tectum.— "98. adial section. oth бге 47 and 72, this grain is only sig er and the layer between the open arrowheads includes *ndexine and intin 268 Annals Иол eae Garden humilis. doar erture; و“‎ Ficures 89-96. SEM and TEM of pollen of Euphorbiaceae. 89-91. rak get Dalechampia 89. Equatorial view.—90. Slightly o ae bd gases section. Grain is 3-colporate and the zonorate Lt is delimited by very prominent costae ight, n has pas E AMOR center y сор апа endoaper prominent costae s angles band. “л. Тес ctum re colpus. 92, 93. Oldfieldioideae. Longetia т агаа view ids oblat ‚ 9-b "brevicolporate grain.— = ee Radial section through a pert ure ioe a i (9 Чепзїїу аз p es lundi ond tuc tum) and partially endexine. 94, 95. Eu phorbioideae ks horbia Bose: 94. Tectum near po 95. Radial section of whole grain. Note elongate “= thin but с in mesocolpus, wih cat accumulation under apertures, and pe wich tect чш кык à acuminatus. This unusual grain has three of these bordered * er at are а midlength and а D in which the pila at one pole are noticeably larger than at the other ate tee Volume 81, Number 2 Nowicke 1994 Palynological Study of Crotonoideae 0 Е + € SEM of pollen of Thymelaeaceae, 97, Gyrinopsis cumingiana. —97. um. Arrow ure. Arrow marks broken ту cete Tectum Зани g роге. — close spacing of БЫН рее тигі. — 10 nidia glauca. Fracture illustrating —98. T. Pim песа Prostrata. The Trow) of vertically elongated muri.— 102. Pimelea M "fectum broken edges (ar LATICIFERS IN Paula Rudal? CROTONOIDEAE (EUPHORBIACEAE): HOMOLOGY AND EVOLUTION! ABSTRACT Laticifer distribution is foie и several genera of Euphorbiaceae, particularly in subfamily Crotonoidese, previously neglected in this respect. Branched, nonarticulated laticifers are каш in both Crotonoideae | е Ец делі io анн = in ee ste (occurring in, e.g., Omphalea), and absent in the other two subfamili However, era without latex ips elongated, highly branched sclereids i in n ‘mesophyll, which may in n sone ткан nd һе homologous with € ifer ll i (ane although several anomalo us and transitional forms are а reported Crotonoideae differs from Euphorbioideae i in that nonarticulated laticifers originate in the e prot ophloem, ux i ri in general more abundant in the phloem, in common with articulated laticifers. Laticifer to the phylogeny of Euphorbiaceae. = > я Ф @ og @ @ a m g The objectives of this investigation are to assess — tropha curcas, but without bae a the phylogenetic significance of laticifers in Eu- (1884a, b, 1886) examined articulated e ers phorbiaceae with particular referen the in din of Manihot and Hoven, both Crot subfamily Crotonoideae and to т extent А Чеа lyphoideae, both less well know n Euphor Webster (1975) used latex рш and pres- upho а. in this Yee (Rudall, 1987). Ontogenetic ence or absence of laticifers as characte > e in e are limited by the availability of sufficient оба | lies; however, information on both is frequen The role of ontogenetic data in systematics has lacking in tha literature, Repani i Hes "e seen a recent resurgence of interest, especially with fers ack of sve respect to the establishment of homologies of fea- records may s ge mean that the he if presen e tures present at maturity. This aspect has partic- is not conspicuous. Latex is of very wid ü 4 ular relevance to a study of laticifers, where de- occurrence among Crotonoideae and Pop velopment is important in the classification of t eae, but varies in color and abundance. rA different types. Most recent investigations of latic- — whitis sap is particularly characte: 9 m ifer development in сеае have concen- phorbioideae; in Euphorbia phosph e М rated on Euphorbia and closely related genera comes phosphorescent on exposure (2p* » Euphorbioideae (e.g., lberg & Sabharwal, tius, 1828). ong Crotonoideae, айо = d elopmental studies on other genera are genera have white | Maniho many scant gely owing to the difficulty of obtaining Hevea, genera with ated кшй Neo- suitable material. Th at do exist (e.g., Chau- others, such as Aleurites, Codiaeum, ап ‘Red veaud, 1891; Gaucher, 1902) need updating in — boutonia, have a clear, often viscous, pid the light of modern methods and taxonomic con- latex occurs in occasional — of sever ral ре Cro cepts. Chauveaud (1891) examined several бе such as Alphandia, Baloghia, Cocconi a of Euphorbioideae and three species of Croton ton, ка rmum, and Pausandra (Me 1971) ideae, Aleurites triloba, Croton pungens, and Ja. & Tirel, 1987; Roosmalen, 1985; Schaeffer, l РА the feline f, H 11.3 D pap iens | nasa: бесш, Чу Botanic Gardens, Zimbabw ee Mount Coot-tha Botanic Gardens, Queensland, hnsonii}: p. Botánico National, La Habana Cuba; K. Alvin, Imperial College, London (material of плата john "m P. Lees (material from Panama and Cameroon). I am very grateful to T. Lawr T x п, ©? ns (Thames Polytechnic), and : Hewis (Hatfield Polytechnic) for ич assistance and here al Botanic Humphries (British Museum, Natural History) for comments on the manuscrip d В. M. Harley (Roy Gardens, Kew) and A. M. Giulietti Universi z São Paulo) for help in the fie Fa. * Jodrell Laboratory, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3DS, U.K. ANN. Missouni Bor. Garp. 81: 270-282. 1994. | Volume 81, Number 2 1994 | Rudall Laticifers in Crotonoideae There are also records of latex in Acalyphoideae: species of Omphalea exude a clear or slightly milky latex, ^em BE to red; Macaranga has latex rangin, clear to Dn orange, or red (Mc- md А та. кө Apart from а АТА records і in Acalyphoi- ad laticifers are iris in subfamilies 1987 nly, e.g., Hevea) and non- articula "s poen vidual cells with intrusive growth, eg, in Ё other genera). Some authors (e e.g. Lado 67) mh BRNS this as evidence of p чуруу. i МЕЧЕ еї а]. (1987) proposed that garded as three distinct {а milies, one aking latic- A one En articulated laticifers, and o dues , although most г "cm (e.g., We ler а) would bie ү оп the basis of other charac E AND METHODS erial was obtained from various sources (Ta- s (S), and in some cases, гов, бхе malin acetic Bor Алу were ed from м author’ n Bra- zil, living collections at Kew (HK), Limbe Botanic K), "iie » Cameroon, and the Smithsonian Ps y Ep иеше (STRI), Panis. 2% ds i E xaminati n (E ) Y A. : rit (кишш (5С), Ке» vi collections (HK), ег botanic gardens, and the author's own col- y instances apparently k embryos, or the ; 9S were not sufficiently well preserved for inclusion in this S surv ge Eur was ENS in FAA for at ем 48 5, then stor in 70% alcohol. Herbariu i erial was г Ed by boilin Ф = water. "pee and Ste "e Were sectioned using a үк sliding mi- ; ‘à me Embr s and apices were embedded in raplast using standard met SE d ed in safranin and Alci ‘ion el , “ask conta arts of cial acetic acid and 20-volume ied. per- about бш g the mixture on a water bath for cell pa dp shaking it vigorously to achieve Separa OBSERVATIONS EMBRYOS The distribution of nonarticulated laticifer ini- tials in the = росе епн S ех каш here adie a А 1877), and is а consistent with records of develo pe S: od E SE = 5 ©. articulated laticifer Dae are prese ni in t iut P of th node, number Fs points уйй the axis. sg pe be permeate throughout other tis- es by rapid intrusive growth, as Mahlberg Sabharwal 1968) domon Шара h UE of i varies Бе 4 or 8 to o 12 or more (Chau veaud, 1891), a variation ылу relating to diff yo size and number of juna ] h 1| the diff; { I H In matu b b d here of anl ca. 8 initials in E grantii (Eu. enerally more in о = Е] — Dn ( this stage. Furthermore, in Crotonoideae the initials occur toward the о dge of the e tis (1.е., within the protophloem ) con- trary to the situation in Eu dese (e.g., S. itials elongate and ош ah intrusively, forming a complex laticif In Baloghia feta ell Figs. 3, 4), indicating ‘possible polyploidy, is no evidence for this. Branches of thus forming two separate Bary (1884) and Chauveaud (1891) also noted in Eupho orbia. Two керки concen rings з g. 7) +h copay (Euphorbia lathyris a synade roton spp.). Further branches proliferate around the cotyledonary node, close to th and ex- te pwards into the coty ons. ure embryos of iram confusa ad a very Ma short hypocotyledonary stem and e broad cotyledons (Fig. 9). Typical ана laticifers TABLE 1. Taxa and material examined, with observations оп laticifers and sclereids. Taxon Source & data* Plant part” Laticifers: Sclereids in leaves: Acalyphoideae Omphalea diandra L. Omphalea triandra L. Omphalea oleifera Hem Mallotus subulatus Mk Arg. Crotonoideae Adenocline acuta B Aleurites bes J. е; T & G. Forster Baloghia luc Bertya mitche и (Sond. ) Muell. Arg. Beyeria leschenaultii (DC.) Baill. C i tum (L.) A. Jus Croton antisyphiliticus Mart. ex Muell. Arg. G roton comosus Muell. A Croton co s licatus Kunth. Croton occidentalis Muell. Arg Croton sagreanus Muell. Arg Croton sylvaticus Hochst. Соо argentea Prain cult. STRI, Panama Limbe B.G., Cameroon K: Muller & roe 4T HK: Hyland 2 Adelaide B.G., oe K: Sharrad 779 K: уыз. 408 HK (no data) Brazil: bas et al. 25010 (K) Brazil: Harley et al. 25109 (K) Brazil: Harley et al. 25156 (K) dá Z: Muller Brazil: Harkey et al. 24827 (K) cult. Cuba B.G. : Ritchken s.n. Brazil: Harley et al. 24832 (K) Brazil: Harley et al. 25091 (K) Brazil: Harley et al. 25226 (K) Brazil: Harley et al. 24986 (K) K: Thomas 4738 + nonarticulated + nonarticulated + nonarticulated + nonarticulated + (older leaves) uapsed jeoiuejog unossilN ole au} JO sjeuuy TABLE 1. Continued. Taxon Source & data* Plant рагі» Laticifers* Sclereids in leaves: Micrandra elata M uell. A. or Bent Micrandra siphonoides Bent Micrandra prucei (Muell. ‘a ) Schultes Neoboutonia macrocalyx Pax Pausandra alien dul. Ducke инак o ledifolius F. Muell. Ricinocarpus pinifolius Desf. Ricinodendron heudolotii (Baill. T Pierre ex Hecke ax А E rigonostemon BP is Mer s L. Synadenium grantii Hook. STRI = Smithsonian Tropical Research Institute; В.С. = of the Institut für is matische Botanik, соны 5С = a * E = seeds for Sn int raction; L = leaves; S = s ‘+= present; — = absent. К: Gillis 15070 : Bamps 2363 SC: cult. Castleton Gdns. HK (no d Brazil: Harley et al. 25098 (K) : Lindeman et al. 256 K: Ducke 7148 K: Coveney 6842 se K: Thow K: s.n. er Eo HK: Aronson s.n. HK: Brandham et al. 2549 co cn mandi eic чы уе ве eie t per ue вд. жр ча Botanic Garden; K = herbarium of the Royal Botanic Gardens, Kew; HK w n collection. t nonarticulated + articulated + + nonarticulated + nonarticulated + nonarticulated + nonarticulated = living collections at Kew; Z = herbarium #661 z Jequunw ‘18 euinjoA әрәрюицо}олг) ul SJejrone] перпн ele y of vascular Scale bars - ег icifer (L) branching h perip with laticifers (L) toward -— 4. Cross section of cotyledons. L = laticifer. Missouri Botanical Garden Annals of the уен 3 ius, +9 эз ade v LU HP 2*4. Ida v | ê yos 5059959, Joe Jc) os Mh BTS a E Aa ate, abe 4% SUN 1? БАТР ghia lucida. —1. Longi re d х Wey T Embryos of Balo, oucher or source information is in Table 1, unless given in the caption. thee ? ge qr titt FIGURES 1-4.? 50 um. Volume 81, Number 2 Rudall 1994 Laticifers in Crotonoideae Is ied бире рз? tyls.—5, 6. S nader nium grantii. Laticifer initia ш, КЕ mw e wipe натаў wig m in initial i sent on e each side of each o Usi ming ca. eight initials, which then rami mify. —1. А nd inner ramifying laticifers (L).—8. a oba. Laticifer initia 276 Annals of the Missouri Botanical Garden Wa iT тысе : 7 ledons. 100 ит. Ficures 9-11. Longitudinal ; tH. jfi b 9. Short hypocotyl and broad cotyl 10. Articulated laticifers (L).— 11. Region near apex (A), showing laticifers (L) in cotyledon. Scale bars = (Fig. 10) are present in the outer part of the pro- also found in Hevea brasiliensis. Intrusive gro ath vascular tissue (ie., the protophloem region, as is not pronounced, although some 5 ort branches with nonarticulated laticifers of other genera) of аге present. Later articulated laticifers form e "i the cotyledonary node and in the base of the cot- gated intrusive branches between surroun ing с yledons (Fig. 11). The cotyledonary initials are the the same way as nonarticulated laticifers most well developed at this stage, as Scott (1886) pecially in the leaf mesophyll (Rudall, 1987). Volume 81, Number 2 1994 Rudall Laticifers in Crotonoideae LEAVES AND STEMS hoideae e lea, which exude es a Mp um stem. La Е were also uie: bee leaf surface. In Mallotus subulatus, although la- ticifers are absent in mature leaves, many narrow intrusive thick-walled sclereid branches are pres- Crotonoideae. Thin walled laticifers were ер l): Adenocline acuta, Baloghia lucida, par mitchellii, Beyeria leschenaultii, Codiaeum v legatum (Figs. 15, 16), Croton spp. (Figs. 13, LE 19, Cyrtogonone arge (stem only), Garcia nutans (Fig. 12), opha. curcas (Fig. 20), Mi- crandra spp utonia calyx, san- ra mac i leaves of Grossera veginei and Trigonostemon chi- nensis (Fig. 22) (in the latter sometimes protrudin between epidermal cells or to hair bases). Pax (1884) н dae hiis in Trigonostemon; йир аге Present in the stem КБИ у ап a pith, eaves, indi- rved in the petiole, clere ids i in Pu mesophyll. In s Rao d iaeum var- ude t a tie as aloghia lucida (Ru- P 87), the fact that Moni are continuousl Towing is demons t mach cells (г. er ones. In the literature laticife with intrusive branches) are terme Ts if E эн thin-walled, and sclereids if in old Bary, 1884, in Euphorbia lathyris; Dehgan & Craig, 1978, in Jatropha ѕрр.). ае Fond ibis is ны similar in MER Hev iio: Manihot, and жнг the main epee being in size and qua BOY. 18 potonar stem, pons sud midrib bundles, ad: may be large and inguishable in cross section (Fi the plant sur rface (e.g., in 15; 16) o or xs the bases of trichom . In 12974 гіа К Ricinoc car- pu contact E small glandular haire on the Pon surface (Fig. 14). In many cases (e.g., Codiaeum garn sgam) bos шше laticifors, immediately A wide variatio on of nonarticulated laticifers is displayed hr various speret: of Viu “n spe- oug gh sometimes less Pi pE in older stems (e.g., in C. omosus). Distribution ranges from extremely cpi: and highly branched in all aerial tissues ү 8» SG anm dd 1 és to isp ely) ap- Q & pet ht т, hve ror latex apparently absent in s of nie n, and this variation may well ies di ic significance within the genus. However, species pa lack laticifers may tal in is is cycle, and a the function of latex in different species would be necessary to establish this. = = investigation, typical articulated laticifers ме in mee и Е t sp. aff. caeru- cous va 21) an ). In Manihot, prs and Оман, which шо present fpr in am aiti «(full 1967, ied Cnidoscolus: d Pen are ra produced in the developing primary phloem, ха also і ue the ates e cambia m. However, in Croto: b conaup served | of ray initials in the —€— cambium in- truding yet adjacent phloem ray cells in the manner of young latic ier initials (Rudall, 1989), ie. new ess initials forming in the vascula cambium. Since no evidence of cell fusion was 278 Annals of the Missouri Botanical Garden 4 К DW. . IGU 12-16. Laticifers (L) in leaves.— 12. Garcia nutans, — 13. Croton sp. (Harley 5 24986), laticifers branching = base of stellate pu with ү region ш 14. Borena leschenaultit, cos section. Laticifers extending to bases of glandular hairs (G). 1 . Codiaeum variegatum. — Leaf surfa showing point of intrusion L t hii between epidermal cells ante ps ross section, with laticifers intruding epidermal cells. Scale bar Rudall 279 Volume 81, Number 2 1994 Laticifers in Crotonoideae e a et tA °ч 16. „6:690: Х IR (cdd — 18. Stem, cross section.— 21. Manihot leaf cross section, s T 20. Jatropha curcas, stem cross "hinensis, . Midrib, cross section. ey et al. 24843).—1 stem cross section. — showing laticifer.— 22. Trigonostemon c phloem region, , 17, 18. Croton sp. (Harl -Toton conduplicatus, gitudinal s s (S). Scale bars — 50 um. L - laticifer. section of stem 7-293 <2. Ficungs 1 section, — 19 ( SP., lon, Wing sclereids howi E 280 nnals of the Missouri Botanical Garden found, these are suce tee laticifers. This spe- cies is unusual in the abundance of its laticifers efor and Shot snares » Un and it үш ply more SPATS in que material because new laticifers produced in the vascular mbium begin to branch и га Сом dan elongating for. a period. A * соргу ОШ enon is simply in the region close to the vascular cambium vould be virtually indistinguishable from a “primar үне (шщ їп primary oe) Mer extended "o mt sp е- cies have АШ. laticifers in the most aE formed secondary phl ficult to A д oem, and their origin is dif- DISCUSSION LATICIFER EVOLUTION The tra т difference between the articu- lated and nonarticul t (Fig. 21). N ss a tes as de Bary. (18 2 and other early ies , ther are several similarities and transitional fonds p te ш to narrow ше gap between the extre of mes о laticifers жулт in the pue of the em- o the condition of Eu- E V4 * 1 1 laticifers in leaves of, e.g., He evea grow intrusively be ophyll cells, in the same manner as nonarticulated laticifers (Rudall, 1987). piens (1 TE срести these intrusive -amoeboid form: |" of Hevea. Other anomalous сей types have also reat length. The “‘articulat laticifers" that jat бп & Craig (1978) Niere in Jatro & n anihot, but may represent an- other fo Nonarticulated (i.e., intrusive, uni- unc Inter, althoug rmally primary in о f о been spe erved i in Стою n to ies cambium е ll 1989), a type of deve ius! usually associated only with articulated laticifers. Scott (1886) believed, following Pax (1884), separately, both from secretory sacs; in fact, row of lotus Шш Dono um M of several species, ‘such as Micrandra elata. How ever, an alter bably more КУЛ п the two laticifer types, is that а la ticifera of Hevea, Cnidoscolus, and МОЛЕ and the чапаи “forms of other gener тараны type (although further data are essential, par ularly on pat aoe ontogeny). The rena type is by far the most common type among bu- phorbiaceae, Ex indeed other dicot inrer per therefore probably the plesiomorphic type in this family. LATICIFERS AND SCLEREIDS Elongated "fiberlike," occasionally brani ched sclereids occur in mesophyll of leaves of many euphorbiaceous genera, generally (though not ex- eden whore laticifere are ke ene E iew of tvpes JE (see Pelon and Орке: ratione. above NE his exclusiv € is consistent with a probable wins between them, ae this hypothesis г equires ee testing. Metcalfe & Chalk (1950) and Rao & Das ( ) ere sclereids i in several genera, as Acaly- highly bra таан leaves, similar to laticifers in many T E s sociated with the epidermis and veins, à also with the bases of probable water- vin, 1987). Am shable in sections, € Nea pet. Минен r, thick-wal pri of Euph n (e.g., in stems ) ыны Rudall, 1987). Rao & Tewari (1960 | Volume 81, Number 2 1994 | Rudall 281 Laticifers in Crotonoideae чө that in Codiaeum variegatum, laticifers ^ Acalyphoide i i h m bw tur conor M ds i in о viter lyphoideae nd a few Croronsideae) i indiennes araphyly, altho ir it is also possible ids were present in older leaves, aod absent in versal (i.e., loss of latex) may yet be j ar on the basis of congruence 5 ae characters. LITERATURE CITED pce es A 1987. Leaf anatomy of е» аге Кпо o produce latex, but ii Prain and its functional significan itself war 59: 5 79-591. BARY, A. DE. 1884. Comparative Anatomy of the Veg- in e Organs of the Phanerogams and Ferns. [En- SYSTEMATIC AND PHYLOGENETIC CONSIDERATIONS iadaceae, Euco : ard р mmiaceae, Sapotaceae, and Mo- БОШО, wW. 23. Anatomy of Hevea brasiliensis. raceae, are widely distributed throughout various uric dicotyledon orders. Nevertheless, as ma any author CHAUVEAUD, M. L. G. 1891. Recherches embryogé- (e.g., Mahlberg & eat 1968) have also niques sur l'appareil laticifére des ap observed, there is remarkable simi arity in laticifer rusos Apoyo a рше AI. oe BL ‘ind: devel b at., Botani 14(1-2): 1-160. tivel m etween compara- Dgncaw, B. & M. E. CRAIG. Types of cepa ely unrelated dicotyledons, especially consider- e crystals i in Jatropha and diet taxonomic impli- J. Bot. 65: 435-352. ing the anomalous nature xs the cell types con- ations. Amer cerned. The apparent relationshi ме oss I st Recherches anatomiques sur p between laticifers d Е “ | branched scler EIS aedes ids) i Hose het Euphorbiacées. Ann. Sci. Nat., Botanique 15(1): n the two MAHLBERC, Р. С. & P. S. SABHARWAL. 1968. Origin non-articulated asd in r. J. Bot. ilie: cll types a e often mutually nell sive in related and early development о of imp ako indicative of a possible homology be- ила of Euphorbia marginata. Am ween them. A . fae М -3 ix om, "s survey of dicotyledon families in D. С. Davis, D. S. Gaurz & С. D. MANNERS. furth » though inappropriate here, might ims Lard the E of Euphorbia: шы h he subject an o indicate The chemotaxonomy of Euphorbia esula L. Bot. J. shi hes of Eup aceae. Sister-group relation inn. Soc. : 165-1 80. E Nd but ү Büphorbiacaae have been тане р ? uc AG nee pé vod e d ТМ 3 sd probably exist among the Malvales, Geraniales, n T P E inales (We bster, 1987a, b). Although ses Meu CR. 1967. enews а latex іп the plant f cur widely among these orders, there a m. Econ. Bot. 21: ew latex-producing families; therefore, in еа; ind . CHALK. 1950. ду көше. of the Dicoty- Sence of a latex-produci ; difE ledons. Clarendon Press, Oxfor phorbiaceae is ; 1 -— sister group, апан Eu- px к, 1884. Die Anatomie der Euphorbiaceae in ihrer inf eae is indeed monophyletic, the most likely ziehung zum System Derselben. Bot. Jahrb. 5: 18 p latex be ON evolved de novo -42]. E mily. A0, А. В. & J. P. TEW 1990; oe делал thin == and ontogeny of ы dier sclereids of Codiaeum tulated th ABER SEO d rud variegatum Blume. Proc. Nat. Inst. Sci. India. 26B: i at Crotonoideae and Euphorbioideae sn 6: de = ee tors usually present) evolved indepen- 0; ArT.: . Das. 1979. Leaf sclereids— Occur- ds м fro alyphoideae (latex and laticifers rence no distribution in the angiosperms. Bot. Not. ey present). ili oi 132: 319-324. Seema — E pean mio - ROOSMALEN, M G.M. v one 1985. Fruits of the Guianan closely link atex and laticifers absent) are Flora Inst. Syst. Bo " Direcht University. main divi Ed to each other. Thus there are three ке Р. J. 1987. Laticifers езун Mino: н ithin Euphorbiaceae on the basi nspectus. Bot. J. Li M e 143- nic) š and other characters: Oldfieldioideae/Phyl- =. 1989. Гай а an lantho; г в; eldioideae/ Phy ге TAW A Bull. re Acalyphoideae, and Crotonoideae/Eu- de of 579-383; TP: . Euphor aceae). ав поіеае. However, Acalyphoideae divides into сен, uM 197 revision of Endospermum Benth. late Broups: without latex (most genera) and wit (Euphorbiaceae, Blues 19: 171-192. рш e.g., caranga, Omphalea). If (as seems | SCHMALHAUSEN, Beitráge E zi a im. ä flanzen. mi E latex production evolved within Eu- ^y» mee dh 7 24: mom P . | Màceae (in the Euphorbioi ni e iupra 5сотт, D. H. 1884 Or if hot ET he Ceara rubber). Qai. J. det $c. 2 Annals of the thesis Botanical Garden ——. 1884Ь. Not he laticif: i f He- vea spruceana. Quart. J. Microsp. Sci. 24: 204- . 1886. th f articulated lati- ciferous vessels in Hevea. J. Linn. Soc. (Bot.) 21: 566-573. SPIX, т B. & C. F. P. Martius. 1828. Pp. 612, 726 n Reise in Brasilien; M. Lindauer, Miinchen WEBSTER, С. L. 1975. Conspectus of a new classifi- cation of the Euphorbiaceae. o i es vie he saga of the classification. and relationships 4 in de puru. Bot. J. Linn. Soc. 1987b. Evo п and db UN tional history of the = Euphorbiaceae: Abstracts, XIV International Bo- tanical Congres THE RELATIONSHIPS OF М. С. Gilbert’ THE EUPHORBIEAE (EUPHORBIACEAE)! ABSTRACT „Пе Euphorbieae and Hippomaneae, though both Ar placed in subfamily Euphorbioideae, differ in many 3 ew While ше > Hippomaneae have inflorescences very similar to Mis of many other members of | the Euphor- ky e E. through a hypothetical ancestor ӨШ a synflorescence of axillary bisexual cymes more primitive than most extant taxa possibly other than Jatropha. rimary aim of this symposium is to review between the Old World and New World species of ety Webster’s (1994) dr bnc of the Eu- Роне з How de fie dpi Euphorbia and phorbiac e. This paper is an attempt to share a ups elate to the rest йы arising from misgivings over the sup- of the goni? mia an an vh my view, could н or ipd of the sub amily Euphorbioi- = much to clarify these асры» and thus the . This is not a presentation of the results of taxonomy of the tribe as a whole. It would dem че and detailed research but rather spec- much greater knowle ie of the эн. origins tions arising while producing a routine Flora ч the Euphorbise — is available. In a group hologically isolated, speculation is needed. © рһ UE а Нір 1 ii dijs 5 pled with The ong-standing interest in Euphorbia (the largest within the subfamily Euphorbioideae along with t iopi her tribe thin the Ethiopian flora), particularly th ree other s (Stomatocalyceae, Pachystro- i ent spe А Flora wr hould delve mateae, and ч ) that have usually been as- id scale taxonomy of those families that sociated with the peines ae (Webster, 1994), ^ covered, but is rarely all arily on t caustic milky latex ‘ry out anything more than superficial investi- ti "onaricte hiicifers PREA glandular К Таха пої actually included in the Flora. bra reduced flowers that alw. Ny following ideas must not be regarded as lack ponen This juxtaposition ing cate not always 8 more than simple-minded speculations. been the case: Mueller Argoviensis (1866) and — the relationships of a group as spe- Bentham (1878) laced these groups at opposite "— Б the ieae used to be of little ends in their sequences. ther deed ma a writer. The matter became a — differences in inflorescence morphology between ifferenc de two groups such that 1 felt forced ! i ми poth y demands a workin = We е ty tl esis of probable sister groups and character я со Кун ‚ One ош m роне tization. Sooner or later such methods shoul very arch grains found ms d the Euphorbiaceae in general and in the Jide; isa bos perd for a synapo- there are nd Euphorbieae in particular, where et which — fae ё. do have а сот- itati inconsistencies in с — pM -— n origin. This in turn led to an attempt to re- ом, for instanc seni a vultis ancestral inflorescence ty : e re Я elate Chamaesyce, very ers cena as a bin which the modern plants could have evolved cUm to the various sections within Eu- most parsimoniously. bia? galoma and t est of € The rod-shaped odi grains have not been wai * ‘here are many other interesting problems reported in other me embers of the Euphorbiaceae, Ung to be t ckled: What is the йене although many taxa appear never to have been dim. cd was employ Р t. funded by the Swedish Ag for В h Cooperation ing C * Flora h M eie ЗАКЕ. Я the bd «de ete рег тй e dence: Department of Botany, Natural f China P; , Hist O! China Project, Missouri Botanical Garden; fies for correspon ory Museum, Cromwell Road, London SW? 5BD, U.K. ANN. Missouni Вот. GARD. 81: 283-288. 1994. Annals of the Missouri Botanical Garden prs a ore spate ме their е else- wher rious challen en me character but the major compa ede ed di ite онг Crotonoideae (Ве wide e et al., 1989). The pollen ше 18 rather upiform (Pu unt, 1987), but siomor- phic; thus the similarity sh ould be ed with caution. Morphologic cal ev idence is even less clear- similarities in t Q ; а о 4 rs many species o Stllingio viti the Hippomaneae, ad such ev- nce can onl g yn НЧЕ 1: 1 ER 1 tia tionshi The a ret toward ers кайы flowers, rare in the family as a whole, has also been invoked to support a Panik ore pinê: It i is a theo- retically, t he flowers egt e: within the Hippomaneae from genera ses as Ma- flowers with many stamens, through to Dalember- tia, in which the male flower is reduced to a single sepal and a s en and the female flower s temptation to suggest that iis nm gives a true guide to N but this must be ques- tioned—could the E y have given rise ај to the Eu uphorbieae? jor difference between the two groups is the dc structure, whic et he a hav in ects, the exception being a group of "ies cialized in New Wo rid taxa, most obviously the genus Pedi- lanthus but also various red-flowered Euphorbia d irds. I embers of d tion, perha insects (Bawa al, 1985), but in at least one case, Mabea occidentalis Benth. (Steiner, 1983), р in ts. Thus the significance of the cy of succule withi Euphorbieae where i has clearly evolved several times indepe у, һегеаз succulence appears 1 t e Hippomaneae, recorded from a few Br 1951). ein еч of Stillingia (Rogers, he nte ast in infloresc ence organiza i hax one must consider whether the sim- so great ilarities between them со result of con- i a 2 опе of the other MAR otably the Crotonor- deae, rather than еи pho orbi oideae, the Eu- bis regarded as a gra ade tribe Euphoric 1ants s even a whorl о below each lobe and a to a naked ovary. There is quite a within this theme with regard to e number of parts within the involu Volume 81, Number 2 1994 Gilbert 285 Relationships of the Euphorbieae amount of Buon aite men ид ш оооп г flowers, but по gener a can be regarded as ae rescence mur [ешеш between y- a the fiil: ‘There has been much debate on the weight of evidence (e.g., Schoute, 1937) seems to ыг the scheme nd by Eichler (1878), in which each invo- | а. } 1 a 2 Ж. 1 The cyathium lexus de very. ee in orga- nizat vast major ity ah +h of other ат, RO tribes ер within samy канот ойе һу Webster. There is a fundamental div within the е between dis ees tribes, with o vule per ovary locule, and the lorte wes ih two оле per locule. The so many features of natomy and morphology that hes will not erm in detail. [ ki ing the observation that ill lowers characteristic of st majority biovulate taxa mus considered rather special- wed red st other uniovulate the stem due many nutrients for so long, to be sia е: n support rel- simi “at fruits, and X as Hav; € male flower LS С their arty E чы said this, шш п obvious exception to logic— Acal ing a ы. жанча, кар a bewilde tray of inflorescence types, не with regard to the relative positions of male and female fl and must be жч а c diim mon evolutionary cess. However, at the same time one must also pt 1 1 Я } 1 . ое of the male and female flowers or the loss of the proximal female flowers plus the de novo produc tion of a terminal female не The latter process seems пе difficult to account for unless one invokes minal allom ternative sch wou d rom a group of thyrses. Such a scenario is very similar to that proposed below iv sideration. The one fact against it is the cymos rrangeme ps of male ers within cyathium Iti is eig to imagine a racemelike uch а groupin nario is to postulate e Euphorbieae ea sexual cymes (Fig. 1). T monochasial or, perhaps quite р monochasial ra The Hippomaneae inflorescence could (a ie Жа, did) evolve Br a simple process ш опоо р боне апі the distal cymes reduced to vale е would have to be deriv se nary process s centered marily on a great condena e» of шеш Hor / the central = perhaps in response to the p reduced to a tua fe- E =. = B ^ @ became all male bracts and dmm glands fused to form an in- volucre. » ptam ДЫ snp is that the cyathium ha has me with a prim ary female ed de bel ef that 4-lobed inv voluc type for the Eupho orbieae, d deed амы бен the initial two levels of branching of a dichasium. Volume 81, Number 2 1994 Gilbert 287 Relationships of the Euphorbieae | An immediate p is the frequency of 5-lobed involucres, which wo ifficult to derive from i ul à regular dichasial cyme but which could be ex- pected as the norm if the involucre w product of the condensation of a Fibonacci spiral derived from a thyrse. Two further bits of e argu | against а lopment from a in the ae very much more А. = imple Iu are of rather rare PE, а, a genus in idi the rently primitive Mage vits n well- угз d corolla and female flower: $ with staminodes occasionall Rudall jue : су f most species of Jatro- Us Te primarily dichasial, but in any 5 5 picata Pa a good example) the ultimat d a chasial, which would su E ih Eichler” s (1878) i interpretation of the cy- Dehgan & С rai sie a rand pha. like pla ( besten the possibility olde: EA С id g ancestral to th iif t seems a барны > possibility that thorn EM of the Hippomaneae and the Oa ight also have had Jatropha-like nie up the question of subfamily relati ioi ips. The cymes of e Jatropha species 22 d tendency tow are strictly terminal and show no ard the production of a synflorescence. Perhaps the ps terminal position of the inflorescence in Jatropha is a factor in their having retained 1 L „л +h = suc i 1 cymes have evolved the thyrse fend cs How- h + у with axillary cya udi ) now so ыыы їп үш fa ever, nclusion, it is ipa e that it is not pos- sible s the cyathium to have evolved from any infin ше и. within modera der Hippama. ber jme common ancestor belonging neither group i but rather to ше Crotonoideae. if to extend the Euphorbioideae to include the Crotonoideae, “ = sal in е E by Мацнее. еї à (1987 ) ery primitive са of the family be roup to i r placin 8 єл due pisi. the Hippo and allies es in a subfamil own and pru. the Euphorbioideae to the phorbieae. LITERATURE CITED oe In: Hist. Pl. 2: 105- Bawa, К: S., S. re D. R. PERRY, R. E. Covite & M. H. GRA ум. _ 1985 Pollination systems. Amer. 346-3 356. otes on Euphorbiaceae. J. Lin J. Bot. 72: BENTHAM, С. 1878. N ., Bot. 17: 185- Se La 201-204, vid 245). BEUTLER, T Jen A. B. A ро, T. G G. Mc Сію p & С. М. Ска 198 bio- pina in 1 the Euphorbiaceae case Re- search 3: 188-1 — L. 1938. си on the Euphorbiaceae, with w genus of the Euphorbieae. Philipp. J. Sci. 64: 397-4 412. DEHGAN, B. & M. E. Craic. 1978. Types of lacticifers and crystals in Jatropha ae "e taxonomic impli- tions. Amer. J. Bot. 65: 352. 2M ч ге шш hi etica] ama sy of ах еки, е» D ore likely : the rem в. On the Jmes, a c centra Ed termi aneae as most ا‎ an: perit ual c . For simplicity the cymes o be рату "dichasial On pos left thers is a progressive song right there is a shor ning of the mi the | echa an id on Fi ng d the pesca pany a monochasia, but in ion sof the main = and orm a dense er of an roue d a whorl of all male є "ymes, while th M PEE a Бей бейе quie to form an enclosing involucr © Annals of the Missouri Botanical Garden х В. ScHUTZMAN. 1994. Contributions toward nograph of потона Jatropha: Phen " "m warme analyses. Ann. Missouri Bot. Gar 349-367. EICHLER, А. W. 1878 erläutert, 2. ps Eng GILBERT, M. C. 1987 Euphorbi о construit und elmann, Lei "Two new “geophytic species of © ings of the Afri b f th Б 5 Kew Bull. 42: 231- Lamarck, J. B. A. P. M. ре. 1788. Euphorbe ou Ti- Шү Euphorbia; Caractére Générique. Encycl. Манцшио, P. С. D. С. Davis, D. S. Gar MAN esu c. MUELLER ARGOVIENSIS, 1. `1866. Жаай 1п: А. Р. de Candolle, Prodr. 15(2): 1-1286. Punt, W. 1987. A survey of pollen morphology in PARADA with special reference to Phyllan- thus. Bot. J. Linn. Soc. 94: m pe J. 1951. A revision of Stillingia à in the orld. Ann. Missouri Bot. Gard. 38: 207-259. mere ч ] . 1987 egere in Euphorbiaceae—A conspectus. Bot inn 4: 14-163 ScHovurE, J. C. n hê aestivation in the cy atheum of ith s remar! me CE A Sy suprageneric taxa of Болон е Апп. Misso Bot. Gard. 81: 33-144. PATTERNS OF DIVERSIFICATION AND ION IN ADENIUS (EUPHORBIACEAE)! R. Haicour,? L. Rossignol,? M. Rossignol,’ and C. Gaisne? ABSTRACT 1 GUT Experimental crossings within Phyllanthus subse = gem taxa of varyin vitro culture. Cytological giras Hatin about the processes of d Phyllan A dei complex pee us. г g pes diu. Some of the reproductive barriers ological as of the proge в may be ydus we ny of experimental crosses has yielded a within subsection Odont 1 i . The wi of this ay suppor the recognition | of taxa earlier Y uisa in these А two соп P fees of x — 10. ‘ r to understand the genetic ocesses involved in Siam of populations ànd species, | In presenting the findings of this study, we will ш. provide a ches of our results with the Previous experimental studies on herbaceous in 1 ylla 980; B t he d edit hus urinaria complex (Haicour, 1983, а, b; Rossignol et al., 7). Two comple differ eie Sis of closely ee айни that numbe TS, ribs ogical characteristics, and morphi ie inated | i orig- Базан m Africa. This study considers it аз а ч containing eight species with three basic umbers = = = t ۷ | re жар, morpholo; d cal in acteristics (2n = 19 981): P. braunii (Pax) Hutch ), Р. mieschii Brunel & Roux (2n = ^18. P. odontadenius Muell. Arg. (2n — 24), P. gag- nioevae Brunel & Roux (2n — 28), P. magnificens un QAM l aunii tos leaves). бү braunii (typical iom and its two variants were each crossed wi nis ie evae, P. bancilhonae, E F. P marne de; The ‘Phyllanthus urinaria com- © { ber hod ek been described ке = Lud ey ~~ ished within seed ornamentation гш г, pos al., 1987). Each group contains d and puapii taxa (2n = 100). Our studies are focu: sexual incomparisiiy phenomena seen in experi- sings и ны from West га char )А е Р. urinaria cones using Rossignol et loid (2n = 50) sed on three areas: (1) the mental с e 1 1 f, nox t po crossings, either cidem conditions or after culturing extracted embryos in vitro (differences ` We thank L. Webster for critically reading the manuscript and for suggestions about changes in w ording. We Rn acknowl edge P. Kupfer for БЕД агіда counts ard J. Defoug and D. Froger for illustration dhi Cedex, Sisty de Morphogentse Végétale Expériment а ntale, Bátiment t 360, Université Paris-Sud, F 91405 Orsay rance. Author of correspondence: L. Rossign ANN. Missouni Bor. GARD. 81: 289-301. 1994. 290 Annals of the си Botanical Garden in fertility of hybrids are noted). Finally, we include 3) a preliminary morphological study of the var- ious parental groups and of some of the hybrids MATERIALS AND METHODS PLANT MATERIAL Plant material used in this study came from taxa of Phyllanthus maintained i инкар. in section Phyllanthus. The taxa included are Phyllanthus braunii (Pax) d 2n — 12. ree morphotypes were found in a single popu- lation near Abidjan, Côte d'Ivoire: i» typical form (12b) and the two variant forms (121 with long branchlets, 12m with ave Phyllanthus ie me Muell. Arg. 2n — m Bouafle, Có voire. Our material ap- E closest to the typical e of Mueller’s ecies. "Phyllanthus Li hte lc prung € Roux an == t Cote d^ Пуд: most of the seeds from this d cality were very close to the next s speci Phyllanthus bancilhonae Brunel E kai 2n = 56, from Bouaké. Phyllanthus e a й Roux, col- n a 1 as a taxon runel & ows Bin vs Len ibe: es it as a tax a ч that Р. bancilhonae and ; nioevae were grown from the same popu- lation of seeds. hould bee noted F: т the species теп- Ire 1980). extended this work by = ln degree of merges ta eg agnificens. pices рор пош: of Р. braunii (3 from Céte d’Ivoire, 2 from Cam- Togo) All these species of Ры аге мов with stems more ог less ligneous at the base and pluriannual (annua or more additional years). The plagiotropic ia ribs (characteristic of s ovary; seeds е oot are dorsally батбай swith жит: ribs. METHODS Crossing and fertility testing of hybrids The crossing and fertility testing of hybrids could in our greenhouses 0 ing the warm ri wi axes, ie crossing experiments were conduc weeks later. or is ed program, four clonal нең yer bra бенен an ds), and е ortho buds po Tig s were cut off to eliminate any possibility of sta- i р after the first marking leaves corresponding to flowers, the (os eration was repeated three жп consecutively newly n е ers). Using the носа, а ens plant was self- l. All th NE GRE e i e plants treated were to isolated with transparent paper bags in ОГ rder avoid alien pollen. eeds produced by — pene n" pa to soil with peat as soon green leaves appeared. For fertility tests, we carri tions and backcrosses with parent then e first ied out self-fertiliza- al plants. In vitro embryo culture ome crosses that produced plants with Ee ns harv dropping before complete ripeness, we nd sterilized ie iae. 13 days after d а xdi mi in » du toa medium with "eap osmotic press T sac cha rose 120 Se the testa of the seed, we ma Наїсоиг РДАН, HRS | uw 81, Number 2 | | | | | Ld Фу Flowers z Q Solitary flower Ficure 1 4 К leaves n le. fa = gree above the ons. sc = scars of c otyledo: P этер in daylight on єс arose (20 ач respon any amino oe у oung bee ts then can be o rt time and btained in a sho rown out on soil compost Cytologi Ytologica] observation The T niques used in this study h vin described by ае “нк ЧОТ “signol (1976). Architectural scheme of a ое from the tyledon: af; fe — cataphylk bos — secondary phi end bud; fl — solitary Pistillate flower Le E 0 СОЗ Р. odontadenius jer Original characteristic: s; AO = orthotropic ax s; RP = deciduous branchlet; je cells were studied using young root api- ces. Roo e immersed in a-chloronaphthalene, fixed i in acetic i. alcol (34—1A), washed, hydrolized ed with Schiff reagent, ss cover slips in a Merck aceto-carmine solution. For meiotic ге Митин staminate floral buds solution, dissected to were fixed in + and squashed in separate the рва обе cells Merck aceto-carmine solution. 292 Annals of th Missouri Botanical Garden Pollen grains were obtained for morphological to be a relation between the seed condition and Putent by "qe di mature anthers in 276 Merck aceto-carmine solut CULTURE CONDITIONS Plant culture conditions in the tropical green- houses at 0 7 80%, temperature 25-40°С, daylight with com- plementary o light. Plants were vegetatively propagated on a stable soil compost. RESULTS CROSSINGS Crossings in greenhouse conditions Two series of experimental e were made. First, 36 crosses were made betwee: a from Cote d'Ivoire: three morp of P braunii (2n = 2 d single forms of P. odontadenius (2n = Ep: gagnioevae m = 28) and P а (2п = 56). А f was as between these taxa and P. magnificens (2n — 40). The results were the same when using different sia ау of the same pope tion. The results of the total of 49 crossings accom- و‎ are буе іп Table l. big: ba include taxon аю with pollen from the same taxon), 2 cross-pollinations. All experimental polli- d i varies min Owers were о P timal, nearly 100% n ue elling of ovary) could be ob- tained from self-pollination Amon ug t the new cross- polinatons, ue hy ar ones are the crosses between the 121 апа 12m {оган e e depends on genotypes р ч the poisson of the cross. The success ra any event, is always smaller than Ма, (I, ) Only 20 42 cross-pollinations yielded viable F, hybrids. 2 crossi ut of i le ven though well devel- 8 days gos Pollination Ea Fable with either abby teguments and DECIES tary em- bryo, or well-developed teguments with unviable embr € degree of delay in fruit abortion does not appear well correlated with the state of develop- t of sh | Tha th } 1 ther e appears ET. Б | © ^" EI = Ф Ф 5 S e ` g tadenius (2n — ا‎ we never nic p iss f h is one of the ie taxa with 2 12, wher ereas there is always an embryo developed hs any x ihe 25 — sed Similar results were found in the pue of P. ws Sa (forms 12] and 12m) with P. Да ые the embryo is viable or not depending о tà directionality of the cross ral features of Кы crossing program stand tible ined taxa are 1 tween tetraploid and octoploid (P. gagnioevae X . bancilhonae). In contrast, a strong genetic relationship exists t f P. braunii, especially between nd 12m; the results of a cross between those ilar to those a a a sd e effect") may. be Bie. between 12b in relation to 12l and 12m. Pg ne have ale so been dem- n P. bra 12) and the gagnioevae (2n "à or P. odontadenius (2n — an magnificens (2n — 40) and P. odontadenius га = Ол) (Table 2). a z = E In vitro ena culture in case of non-viable progen have In ee yielding no viable progeny» s 2d observed a kind of embryo rejection prior t uration vis = Е -— isolation of the 13 days a E mitted i development. Rapi transforma the seedling into a matu hybrid р = 28 crossing, and them rep ings (тене 2). Hybrid plants were obtain! instan Results from о een vario netic entities of Phyllanthus odontadenius in greenhouse conditions. First generation I, and F, descendants from auto- es); SE = seed without embryo; Ep = seed with minute embryo; Ed = seed with ict embryo; M = meiosis or cuc sings. T — fruit abortion се mip boxes); (MN, normal meiosis; MA, i. rrant meiosis; MTA, ery aberrant meiosis). P. bancilhonae P. magnificens P. gagnioevae P. odontadenius P. braunii BUS сзади“ S typical long branchlets brown leaves С 12 chr. Q 56 chr. 40 chr. 28 chr. 24 chr. 12b 121 12m 3 Fyclose to 56 | ^ T (circ. 13 - 14d) Fı T(circ. 10 - 11d) T(cir. 10d) — 7] T(circ. 10d) S(| 56 Ц 48 chr. N 40chr. E sterile br sterile PAM MTA | Ep MTA | SE SE SE S Fı close to 56 Fy Fy T(circ. 16d) 7 T(cire. 16d) T(circ. 16d) *| 4 48 chr. ц 34 - 32 chr. 32 chr. 2 m DM Ў sterile terile sterile x: MTA MTA | SE SE SE За T(circ. 15 - 16d) Fi F1 F1 F1 F] $ 28 Ei 34 - 32 chr. ц 26 с. 20 - 24 - 26 20 - 24 - 26 с. 20 chr. 50 sterile sterile sterile sterile sterile ct Ed A A MT MT MTA 3 F1 F1 F1 T(circ. 114) T(circ. 144) Т(сїгс. 15d) 3 40 chr 32 chr. 26 chr. 11 ed IM $ 24 ile sterile sterile n MTA MTA MTA SE SE SE T(circ. 10 - 12d) T(circ. 10 - 15d)... | F1 T(circ. 16d) F1 Fi 12b 20 - 24 - 26 chr. 11 12 chr. 12 chr. sterile Ep SE MA MA Е T(circ. 12 - 14d) T(circ. 10 - 15d) Т(сис. 15 - 18d). | Т(сис.12-144 Fj F1 8 121 = 12 chr. n 12 chr. © به‎ Ep SE Ed Ed MA MN 7 (circ. 14d) =] ~~ T(cire. 10 - 15d) T(circ. 15 - 18d) T(circ. 15d) —. | F1 F1 12m D PA ee) 12 chr. 12 chr. 11 Ep SE Ed Ed MA MN p66! с JOQUINN ‘18 eumjoA sniuapejuopo snujue[|/Kud че зә поен €6c 294 Annals of the Missouri Botanical Garden TABLE 2. ph Hann Кош opone progeny present or not depending on cross. SE = seed without embryo; Ed — Crossings Results in greenhouse After in vitro culture ruit abortion and no progeny except after in vitro culture (22 cases) N 7) — gagnioevae X à P. bancilhonae M a eder X & P. braunii typical (12b) 9 F е В ar mu x 6 P. gagnioevae 7) (28 = 4 x “ү 12=2 x 6) P uds typical (12b) x ё P. bancilhonae ce iis en x âP. үч hg gs (121) (56 = 8 x 7 ) 9 P. long E (12) x г P. baron QR: эк ges x à P. — mp Pod х 7) (12 MK Р, WEM d (12m) x 6 P. DRE VASE oP: | жул. x à P. jose ee (12b) 10 6) QP. орен a XR: pissi < $ P. magnificens x à P. — d ge (121) (40 = 4 x 10) -2 x 6) 9 P. long branchlets (121) x ; p magnificens 9 P. pr oo X 6 P. brown ие (12т) х 10 ) BP; pw paak (12m) x à Ê.) magni ens 9 iy are x 6 P. braunii typical (12 = 2 x 6 ~ 12b) x 6) SP ranni typical (12b) x â P. odontadenius P: ntadenius x 8 P. long y s (12b) x 6) 12 = ow. pau ni (121) x 6 P. Noc 9 P. odontadenius x 8 P. brown leaves (12m) (24 = 4 x 6) 12= 2 х 9 P. brown leaves (12m) x 8 P. odontadenius 9 P. long branchlets (121) x 8 P. gagnioevae 9 P. brown leaves (12m) x 6 P. gagnioevae 9. P. PEE x à P. porci sinn =8 x EF gs es x 6 P. EEN (28 x 7) 24=4 x Ct ae EE уан x à P. gagn 28 = 4 x 7) $ P. gagnioevae x à Р. Mari ува (12b) 12 2 P. braunii typical (12b) x 8 P. gagnioevae Sterile hybrid F, (14 cases) F о descent (Ep) F, 42 chr. No descent (Ed) F, 42 chr. No descent (SE) No descent (Ep) F, 34 chr.? No descent (SE) No descent (Ep) F, 34 chr.? No descent (SE) No descent (Ep) F, 34 chr? No descent (SE) No descent (SE) No descent (SE) No descent (SE) No descent (SE) No descent (SE) No descent (SE) No descent (Ed) F, 18 chr.? No descent (SE) No descent (Ed) No descent (SE) F, 18 chr.? No descent (Ed) F, 18 chr.? No descent (Ed) unknown — No descent (Ed) unknown res ‚ 48 chr. F, 48 chr. F, 40 chr. F, 40 chr. F, 34-32 chr. F, 34-32 chr. F, 32 chr. F, 32 chr. F, 26 chr. F, 26 chr. F, 20-24-26 chr. | Volume 81, Number 2 1994 | | | | | —— — Haicour e 295 Pene ese TaBLE 2. Continued. Crossings Results in greenhouse After in vitro culture 2 P. gagnioevae х 8 P. long branchlets (121) ? P. gagnioevae X 8 P. brown leaves (12m F, 20-24-26 chr. r ба fertile hybrid Е, (4 cases) | ê P. br es (12m) 9 : Em leaves (12m) : x е Е тн Ere (12b) f Е Нр" 1 (12b) x 6 P. long branchlets (121) | E ( ê P. braunii typical (12b) (191 o fertile hybrid F, (2 cases) ё P. br es (12m) 9 : m leaves (12m). x pA P. long ымын (121) — — © F, 12 chr. F, 12 chr. CYTOLOGICAL ANALYSIS OF PROGENY Chromosome counts It was first necessary to confirm the chromosome abl morpho нена al 4 and amis весе distribution: own by the related spec ies n odontadenius (n = 24 Жыш nd Р. braunii (2n = 12). ver, in examining P. magnificens, we al- Ways Hind the chromosome Ee to 2n = 40. To confirm this ples to P. Kupfer (Neuchatel, who ba found a count of ШОШ. y) as 48 (Fig. 2C, С), 42, 40, 34, 32 (Fig. 2B, de ‚ 20, s 12 represent exactly half of the 2060) unt of the respective parents, а h and - to ү confirm the hybrid nature of the plants Studies of meiosis ain Three categories of F, hybrids may be recog- ET osis winds risen nas 1 (MTA in Ба ): little p ing, with n ous univalents : е I and weed in воено I. This In 14 cases from the following crosses: 228 x à 121 928 x à 12b 928 x 812m |? 12b x 6 28 Berk 9 40 x à 24 224x656 |? 24 x 840 2928x624 |? 56 x 640 2924x828 |? 40 x 656 240 x à 28 228 x 640 (2) Meiosis with some anomalies in Table 1), especially an emis umber of bivalents in metaphase I (i.e., 4 or 5 bivalents nu of 6). 912b x 8 121 jJ? 12b x 612m 9121 x 8 12b |? 12m x 8 12b (3) Meiosis perfectly normal, with all chromo- somes paired (MN in Table 1). 9 121 x 612m $ 12m x ê 121 PRELIMINARY MORPHOLOGICAL ANALYSIS OF HYBRIDS AND PARENTAL POPULAT е found a PN homogeneity of F, hybrids f g. It is уегу ‹ difficult (нон trom u tween evel i hybrid of a given ipee о, very ro from any hybrids obtained bon dod between other species. ybrid nature can be visually determined js able 0 and 56 easil is different, e.g., in brids are closer to 56 parent); or b d'Ivoire gave similar characters between plants with 2n = 12 (121 and 12m) and 2n = 4 on th one hand, and bet an = nd on the & Bancilhon-Rossignol, 1976). the basis of this, we concluded that Р. odontadeni- 296 Annals of the Missouri Botanical Garden Es ái "T te ne Popule quid i mplements.—A, А'. Ph ge dene F, ‚ A’, line et s EET 2 тиен iio satellites indicated by markers). — B, B'. Ny pt bat hybrid i: P. odontadenius (24) x & P. INR (40)).—C, C'. Chromosomes of F, hybri ( (40) x 8 P. ls apa (56)). us represents tetraploid form related to ds, diploid P. braunii, and P. banci bicis ae the ploid form RAE to P. gagnioe EXAMINATION OF POLLEN FROM F, HYBRIDS— RTILITY F, hybrids from pollen gr grain examination agrees „ik the preceding one estab- lished from studies of meiosis. Thus e observed Rai "МТА that hybrids with very abnormal m Ame e pollen is very heterogeneous, with g a fering in nd length, and so (hetero - or deformed. " In ў with abnormal meiosis on less he depen in the pollen, and in a crossing 12] with 12m, the pollen ee à quite normal. a is iet surprising, Volume 81, Number 2 Haicour et al. 297 1994 Phyllanthus odontadenius Morphological E: nates characters from parents and hybrids. Degree of pedicel length th. |o 5 symbols +, ++, ‚ +++ +. ІЛ = length/wid Genotype Р. odontadenius Р. magnificens Hybrids Parameters 2n = 24 2n = 40 24 x 40 Deciduous branchlet umber of green leaves 38 (16-50) 14 (6-20) Leaf blade form subquadrangular void with acut pi ovoid lightly acute ul Number of secondary veins 5 pairs 5-6 pairs 6 pairs LA 1.4-1.6 1.7-2 1.7-2 wer Flower and pedicel length H ++ Calyx-lobe ch teristi acute and thin with bod rounded at the tip acute and thin with rosy- white-edged v with red-edged vein dged vein Style characteristics bifid pas with к bifid style with red bifid aa with i bra branches apart branches ap: ; oge Disk characteristics Seren, crenulate dissected into 6 segments dissected into 6 segments Genotype P. bancilhonae Р. magnificens Hybrids Parameters 2n = 56 2n = 40 56 x 40 T branchlet of green leaves 36 (20-49) 16 (6-20) 28 (18-30) ve blade form broadly ovate ovoid with acute red broadly ovate apiculum Number of secondary veins 4 pairs 6 7 pairs ki 1.8-2 6-1. 1.8-2 Terminal 9 flower present missing present 9 flower Flower and pedicel length ++++ Calyx-lobe characteristics ке апа UN with bod rounded at the tip — color and S reen-edge red-edged vein tyle characteristics sloping style ay pnt more ed style with jen p intermediate branches red branc = ope and rosy branch- Disk characteristics continuous, crenulated, dissected into 6 segments scene into 6 segments edge slightly 6-lobed id fertile, while the hybrids from 12] x 12m are “ы trad and aberrant pollen grains. totally fertile (Table 3). mo- Some : numbers (Table 2) are sterile. After manual DISCUSSION AND CONCLUSIONS invari- h ay М m 10 The seeds have esults of our от appear to be of so Ccid teguments containing a che agis an aes esp they are compar with pe hh OF Жеш an abnormally pale embr ble studies of d t ху ИРЕЙ urinaria n Bac ings of F, hybrids with parents are a complex of eastern Asia (Haicou ur, 1 ды; са ад as did crossing between two гесір- In the Phyllanthus odontadenius complex et С "аі hybrids. The F, generation is ME com- sect. Odontadenii) we i tified five taxa with a А Tt atn ile. different karyotype with diploid complements of X hybrids yia b ossing tax 12, 24, 28, 40, and 56 chromosomes. taxon, Sa 12 а es (Р. Ь d the fou qud identified as P. braunii, E three subgroups: b rids from 12b x 12 and 12b > x 12m are dir 12b, 121, and 12m (Fig 298 Annals Фра а Garden Deciduous йе ыгы epp Penis flowers ot Phyllanthus асра i = 56), 15. 3) of FIGURE 3. (2n = 40, and the two hybric P. mag gnificens and 9 flower of hybrid 56 x 40.- branchlet and 9 flower of parent i e In our earlier studies of the i Moers’ uri- e ur STE jos, SOR. "IOOR: > SOR group is polymorphic with two subgroups: soni and 50R2 (Fig ныз sult of our crossing рг ograms using ta Ey bie ене на ры r апа Р. uri- пагїа complexes, it а appears that there are repro- ductive barriers between taxa not previously rec- ognized by earlier Nene In some instances, th а barriers to inte almost Eu /. Deciduous and 9 flower eds 40 x 5 B'. Deciduous branchlet 6.—D, D’, Deciduous А : „е has 0 and indicate that considerable divergen¢ curred. Lb onsequently, we consider Phyllanthus К па! 1 1 ; broad, pc tadenius anc sely relate species. In the P. urinaria complex it 15 inc make so inferences about the en most ution if we consider the 50S group to Фр itv (50S p lates were found only ш ay then sed original dd The 50R group п a d considered as a mutant ri with Mer characters; this pues beyond up appear and an 1). B gro stage of diversification a ie original center (50R Volume 81, Number 2 Haicour et al. 299 o mus "5 4. Deciduous sraa and | pistillate flower. of Phyllanthus odontadenius (2n — 24), P. magnificens = ) and the two hy ybrids. E. of parent 24. F, F'. Dec Ди БЕ. hlet Ts of hybri 0.- —G, G'. Deciduous branchlet and 9 flower of hybrid 40 x 24.- -H, H'. Deciduous чэй, hlet and 9 flower of deem 40. analogy with what we observed in the P. numbers: diploid and tetraploid eon ies with x — ные, complex, and by comparing the two den- , tetraploid апа o oc ja qi with x 7, and a M (Fig. 5), i ppears that the taxa with 2n Briss with 10 (P. magnificens), newly I 12 are of t most recent netic origi rted in is pape I w basic num mb r his assumes an aneuploid reduction from taxa reported for the first time ir Ph lanthus rein- E 10 (P. magnificens) to those with x = forces the hypothesis suggested by Webster (1967: "sadi 377) that the basic numbers of herbaceous Phyl- the ^a : er altho ugh there are some E in "ai might form a desc wu eries fr з х = h versification within the two complexes, ther to x = 12, (...), , 8, 7, 6. We can = eg differences. In the P. urinaria com- now ow almost complete ibis series, giving x — 12 » dts z à : e e a a and 2n У z Ek сш ge 5 р. i ds nally based crossing barriers " as diploids and tetraploids on chromosor : sed on a common basic number (x = 25). This observed by us in two complexes of herbaceous biis. Их = 25 is most probably the result of Phyllanthus (P. odontadenius and P. urinaria) га amphiploidization between species with x = 12 between morphologically similar plants were not ure unique to this genus. This pattern is common in Sidi mart, in the P. odontadenius complex annual quem SH ы, 1) made Te point € are several groups of taxa with different base numer 1F This is the case, for example 13+12 (amphiploidy) о X= 25 0 G (evouurıon (decreasing any) | FIGURE 5. evouurion D D D (2) (em) (ш 10| | 40 х= 10 А ODONTADENIUS COMPLEX В URINARIA COMPLEX Dendrograms showing proposed relationships in the Phyllanthus odontadenius complex and P. urinaria complex. 00€ ay} Jo sjeuuy uepJer) jeoiuejog ипо$5!у Volume 81, Number 2 1994 Haicour et al. Phyllanthus odontadenius 301 everal LE of Compositae [Madia, Layia ) Caen, 1 эе Microseris (Chambers, 1955), and Hi hus (Heiser et al., 1962)], in the genus Gili алмашы) tie pi and in Clark- ( Lewis, 1955). In th геге we find the ом exemplified sica (Yarnel ollinsi Graminaceae it is edd Eu alia by Bromus sect. Ceratochloa 55). (Hall, 1 LITERATURE CITED =» J.& J.B 1980. Etude du róle des feuilles иш n phénomène d'iapnnapatibilbé entre Р їах- Ми ell. A deep ES sora France, 127, Lettres I va (25:9 Brunel, J. Е. & J. Roux. 1981. hyllanthus نما‎ Odon tadenii (E uphorbiacea: e) 5 va Ag ar de үүн TM 11: 69-89. -HAMBERS, К. L. ystematic od of the trib. Dudley Her- А . Stage Species. Cornell Univ. Prees, Ithaca, New , E. D. . The genus Collinsia TX: nas T сч chromosome repatterning. Cytologia 25: GOODSPEED, T. H. 19 Chroni- ES ca 09:4 vanis E rana usetts. "RANT, V. 1954. G dc and taxonomic studies in Gilia. VI Е EE in the leafy- stemmed Gilias. Aliso ln ds Y t Plant اا‎ пекан Univ. Press, or Hatcour, В. bilit x taxons de Ph rhb eae urinaria L. (Euphor- -— Mise : enters t perspectives ouvertes ч e. Bull. Soc. 0, Lettr Ee (3); 2 "E Таш 198 тка d'analyse de la structure ее et de l'évolution d'une espéce rud- Bot. France érale pe Phyllanthus urinaria 1. (Eu- phorbiace ae). 1. Et d P. urinaria et premier es sai de classification. Bull. Nn Paris, IV. 6 (sect. B, Adansonia Mus. ^ Hist 1): 63- Eléments d'analyse de la structure me des s pop pulations et de Feldes. dix Heiser, C. B., C. МАК N&D M. 5м! 1962. Spe- cies crosses l Вані 1. Diploid s Ses Brittonia $e E H. & M. wis. 1955. The genus Clarkia. iv. Calif. Publ Bot. 20: 241-392. Croissance et développement des ма М. 1 embryons globulaires de Capsella Bursa-pastoris cult in vitro dans un milieu à base d'une nou m solution minérale. Bull. Soc. Bot. France, Mém., Morphologique, Aat, ie (Euphorbiaceae). Amer. TRAN Ha, N. M. & 1. BANCILHON- ROSSIGNOL. — 1976, Phyllanthus r découlant т ипе m arée de e P.o " g. à proie ана io 56). Rev. Cyt. Biol. —, . BELLIARD. 1976. Analyse m phologique et biométrique comparée ies six taxons n Phyllanthus odontadenius Muell. Arg. à garnitures аан miques différentes (2n = 12, 24, 28, 56) et de leur descendance. Rev. Gen. Bot. 83: 269- WEBSTER, G. L. 1967. The genera of Euphorbiaceae in the Рат TE United States. J. Arnold Arbor. Cytogenetics of — 81- 48: Mapas on 1986: И, Crucifers. Bot. Rev. 22: EARLY EVOLUTION OF DALECHAMPIA (EUPHORBIACEAE): INSIGHTS FROM PHYLOGENY, ВІОСЕОСКА COMPARATIVE ECOLOGY! W. Scott Armbruster? ABSTRACT сае ampia contains over 120 spec species in both .M the Old and t orld secrete КЫЫ: in the уе ica раи oe obi S and Usin en 1 construction. Ориг mica ascertai n the on volu per bi ies nx v throughout the lowland tropics of Asia, Africa, and the New tare resin from large glan вар aggr egations e pollinated by bees that ое п for use іп nest m > la and “ecological data and phylogenetic aa : с al asi na or Sout a in Aus mid Cretaceou: a the rafting of. land n masses, or by may have involved pollination л I voll g ted in wes ous or early pra ah vin Md. and spread E the tropics run across higher poe n they had d nts between the Old a ew Worlds. a po ollinatior I I ЕХ ۱ e earliest FE ч апа геѕіп лабаата аѕ ser the role y herbi vores par) or mi le of micis reward. The inuitus libto ii E wiih resin- a bees may have originated independently in the Old and New Worlds. тацы боеашо шшш of similar plant spe- i: botanical explorer began to report them. We have lo nta | Оа An rS RE challenge is to the еу e: din that s occurred since the isolation of plant populati R earch и on ps ре hes rarely proctedad r of фо sister taxa and their Ош la - ord. The development of powerful oon Raise methods of phylogeny y reconstruction, ever, now makes it realistic to attempt retrospective diss of ancient biogeographic and evolutionary processes in the absence of a fossil record. » Donoghue, 1989). The field of vicariance E oe represents one such roach in its tempt t anal f phylogenetic relationships of several independent taxa as d to à d dist g the processes that affected the z those taxa (see review in Humphries & Yu 19 In the Ba itis paper I consider p ec about the early evolutionary and b re events is affected the members of t he pantrop tionships. This is because research on historical ? (2) W at е enomena, especially those as ancient as the events led to the present geographical distribu : ems? (3) creatin ns-oceanic disjunctions, is extre ly cies, character states, and p ion sy$ in the difficul i What was the orig inal pollination $ system L M. Huft and С. L. Web: thi HS Lees and р. Ode ior „ generously providing me pe snubs ad and M. E. "Eva, С. Levin, P. M. Rie ts BSR- an anonym or comments on earlie rch was 5 supported b dir from the 8509031, ISR. 900 06607. BSR-9020265 ج‎ the National Science ensi (USA) and а & National | Geographie So ? De nt of Biology d Wildlife and Institute of Arctic Biology, University of Alaska- Fair Alaska 99775. 0180, U.S.A. ANN. Missouni Bor. GARD. 81: 302-316. 1994. banks, Fairbark* F Volume 81, Number 2 1994 Ar 303 mbruster Early Evolution of Dalechampia genus, and how did the early diversification of pm id tid gerun (4 ) H ow, when, an } 1 DNE ti bees arise? Using the cladistic method of олие reconstruction, I develop explicit hypotheses "i dressing me yur Ratones! шоо 1 ой th hy the наьа between the plants and their her- bivores and pollin NATURAL History AND DISTRIBUTION OF DALECHAMPIA Dalechampia is distinctive within the Eupho biaceae: the ca. 120 species have rangement is probably homologous to of the eo of Tragia and pee е їп thes prse viny species and Dalechampia comprises mostly viny € dodi and uode are also similar g ing, rystaliferous tri- орм of virtually identical (W & Webster, 1972 r these Roi Webst ed has recently ойна» that Dalechampia uded in tribe Plu каса Ava n and W /ebster ster Бак onfidence, Tragia an 1а as candidate outgroups in the phylogenetic oe Кеч below í the relatively uniform bloss phology à їп Dalec. mpia, there is parie à gp in the MEE biology. Most aer th the e Paleotropics and Neotropics offer sin as a pollin female or tii за that collect rini from a variety of in nest construction (Cam- grance-collecting male euglossine bees (Apidae: Euglossini) ics & Webster, 1979; Arm- ш! ster ree al. с ИЕ et al., 1992). in the гелин of sex pheromones (Dressler, 1982; Whitten et al., 1989). Several species in the NESE its and in Madagascar appear to pro- duge no reward ion pollinators other than pollen; llecti h and Безден (Armbruster et a 1993). Species of Dalechampia are found throughout New the ud tro bye of AN Africa, and the rid d 2 k Atere ca (Fi ig: I): ee resin n (end pollinated by resin- iE bees) are euglossin cies rud ri resin pe and лш by he: seeking пиеси аге t di- South ERES and southern Central America s b Magin, 1987), while Webster & Armbruster (1991) ge | A E ч 1 Al < г } © + America (sect. Riel to form probably the most basal group. It is possible that the Mad- nd Soe th ayant poe species, jf арча ME an orm a sin- v был] о ог рагарһу cluster of s the possibility in Dalec ham- species. This г pia has a c side biogeographic history, with multiple migrations among the continents and in- dependent evolution of the mutualism with resin collecting bees i ld and New World tropics (Armbruster & Mziray, 1987). Alternatively, if t semblance bet Madagascan species and ween members of section ГА рай is the sed of homoplasy (character convergence and/or rever- sals), there may be no reason to ste a very =н tbi history involving an origin in New E 0 followed by Fas to ы other ‘mh Armbruster * Mziray (1987) proposed four ypotheses dat could = nt з 1 Armbruster ebster, 1979; ternative a sheds rmbruster, ө, О: & Maray 1999. epr — distribution of Paler бнри pee resin ic “1 amon st (the “resin deni ") associated with the sta : mate flowers (Webster & Wb 1972). About Ozen species in the New World offer —— à pollinator reward and are pollinated by fra соіа, (2) газа еза mid- to high- к. tude land bridges during the Тегі y, when these latitudes had subtropical climates, (Э) а пеаг у © on- tinuous we: FIGURE l1. App Хх, ۹ À 6—2 Spp. 96 J 7 5 spp. N Je _ PH F /2 Spp. and number of species of Dalechampia in the Paleo- and Neotropics. eu} Jo sjeuuy РОЄ иәрео jeoiuejog unossiIA Volume 81, Number 2 1994 Armb 305 Early bum of Dalechampia o late Cretaceous disrupted by subsequent conti- nental drift, and (4) Are of primitive Dal- champia in Sout h Africa, and Mad و‎ n. continental drin, LL о extinction n Afr ааш a second mi gra £ fp m tion (eit er by dist IY 1igration via mm [atitudes) of more dd species be- tween South America, Africa, and Asia. Circum- е E po bh d supported the sec- ond, t ut not the firs E They P not attempt 9 ‘identify t the site of origin of the genus nor the Hinection dispersal суеш. The Di f 4h analyses of the New d Old Wo rld mm to refine thes e biogeographic hypotheses and specify possible sites of origin of the genus, directions of dispersal, alton eei of the earliest species, and the s) of origin of the novel mutualis with resin- Вора bs ees. Р HYLOGENETIC INFERENCE METHODS To reconstruct ا‎ relationships amon Old an New World species of Dalechampia, I polariz he ancestral n Ves global par- ric taxa to и volution, assess tree congru- nce with alternative biogeographic hypotheses, and Produce graphics. After initial run with the full set of taxa including the sister genera), I eliminated all but member of eac sed AUP run time Sod computer. To further reduce run times the ts of the initial cladistic analysis were used to tees ү 1 s. 1 E: г Ј states, and the outgroups were eliminated. These теа е. етн the relative positions of Old and М t екы мн» е conducted to de- termine the sensitivity of cladistic reni m initial varied the assumption of character seversibility one vs. irreversible between plesiomorphic [primitive] and apomorphic [derived] states) for | PL 1 e 2 1 il combinations of the three characters. hen examined each of the numerous maxi- 4 x d + * РЕЧ 4 all е 1 + MY ы А | ех zin ain d distribution of Hu character states, consistent wi = = аїа mption (see Maddis son & Меда, 199 RESULTS When all characters were assumed reversible, cladistic analysis of the set of 34 taxa yielded 436 equally e trees өнүү 0 ste eps, consistency index [CI] = 0.47). The large number of equally parsimonious trees reflects instability i in the a. ace- ы! а б so Don (ШЕ 2). The gland ве and geograph- characteristics. e 3 25 эй. Ф f +} tree) were inferred by parsimony from the аныи © llowed f the history of these sud шир [ш pope he ewe ecological biogeographic implicatio nerally suppor Hype 1 1 a cr Dal. ia ori n the Neo есп. 5 the dew spe or pollen- -collecting insects mu s (Fi 3). volved a и (olin e Neo ized Madaga Cladistic а of the reduced set of taxa 306 Annals of the Missouri Botanical Garden d in “o? TABLE l. Characters and character states use: phylogenetic analyses. The character state is the ancestral conditi pie as калдан by global- NUN outgroup analysis. * indica the phylogenetic bot sis. ates character was not u: . Nature of pome ERA bract: O — stipuli- form; 1 = involuc 2. Nature of distal ed bract: 0 = stipuliform; 1 = involucral. 3. Pos ition of proximal involucral bract: 0 = separated from ee cymule; 1 = adjacent to pistillate YI 4. рд оп proximal involucral bract: 0 = present; = absent. " nies ip Hn veins of involucral bracts: 0 — 3; e 6. ( T of involucral Би a = white to pale Bree = pink; 2 = yellow wn; 4 = deep green. 7. Shape of boss of um КАСЫ 0 = cuneate; 1 = cordate. 8. Shape of tip of involucral bracts: 0 = entire, acute; 1 = lobe 9. Persistence of involucral bracts: 0 = deciduo shortly after anthesis; 1 = persistent into ring Ў Mumbar of pistillate sepals: 0 = 5-6; 1 = >7; > s: 0 = obed. : pene "i icd реше Ө a ee. 0 d Size of distal pistilate dnvolitedllar bractlets: 0 = ulifor: small, stip — large, enveloping 14. өре of реше sepals (excluding lobes) 0 = е; 1 = lin 9: Syl 2* ape: 0“ = саи to PO hers at tip; 1 — clavate; 2 — ngly dilat ; 3 = ae ie 4 = de cn oed 16. Number of stamens: 0 — Lv L7. rs of staminate eel eae p PE 0 — free; — connate. 18. Symmetry of staminate pleiochasial arms: 0 = de- ; 1 = bilateral. 19 : Nun са of staminate pleiochasial arms: 0 = 4; 1 BAY, inate bractlets: 0 = symmet- really ebie 1 = tue et "gland." УА acute; 1 = truncate. 22. Margins of staminate bracei О = entire; 1 = ate. 23. e. secretion by staminate bractlets: 0 = absent; sent. 24. Resin color: » - clear, whitish; 1 — yellow, orange; = maroo 1 25. Закін a piros 0 — green to whitish; 1 — 26. Stigma color: 0 — pale to deep green; 1 — pink to maroon Zt. :0= rub. 28. Relative м. Жы. ч = > limb length; 1 = ^4» limb length. 29. ра shape: 0 = simple, entire; 1 = compound; 2 = simple, 3—5-lobed. 30. Leaf venation: 0 — palm nate. 31. Leaf stipules: 0 = stipuli ulifor = : fo lio: - Involucral hi stipules: 0 - - aire l = fo- liose. TABLE l. Continued. 33. Pollen узди 0 = үн Mises 34. "à d s :0 = oth; 1 = rugulose; 2 = lt 35. Seal shape: 0 = 02 = 3-angled; 2 = lentic- шаг; 3- slightly po vid 36 : ens le of da roximal m bract: 0 = present; юе : 0 = reticulate; 1 = гамія аїе. 3 ed size e of реше sepals in in frui capsule len ximately Rr cae 2 = fully dud d e. 39. Stinging удава trichomes: 0 = absent; 1 = resent. ^t Number of carpels: 0 — 3; 1 — 4. «t De x ofi purs arms in staminate pleiochasium: 1; 2 = 2. ollen-collecting = (Trigona e beetles; 1 = resin-collecting bees ( raices females, Trigona when Euglossa, Eulaema a, and Жз. females), 2 "42. Panton: 5 sa ‚ Eu lae a, Eufries 43. ue usion ы gode ido bractlets: 0 = free; 1 44. Pre resence i ut pof gue. 0 = absent; 1 = present. 45. Leaf nectar nt; 1 = present. iti 46. Dine onde condensa n: 0 = not conde — highly conden 4T. Dem of leaf stipules: 0 = well developed; | reduced to absent. yielded 50 equally parsimonious trees (70 Ms CI = 0.61). АП of these were consistent e sinh cal and evolutionary conclusions the analysis of the full set of taxa а : г asonable is it to assume that all cha a ition. For example, ider charac i the involucellar bracts of the sta vU florescence (pleiochasium) are free, eac E Based lei ial arm, or onnate yee . міт" on outgroup comparisons, the forme Al- o be plesiomorphic and the latter ee i Volume 81, Number 2 Armbruster 307 1994 Early Evolution of Dalechampia Es asain coinage of AS initial EES Н of heee of characters upon the biogeographic/ evolutionary p obtained by cladistic analysis of the reduced list of m le не taxa. pumas ee i hypothesis(es) with which the most parsimonious Table 3). Nüdilen à in ete: Баи the proportion of maximally parsimonious trees consistent with each wee In combination with character Character assumed irreversible Alone 20 41 17, 20 17. Fusion of staminate bracts 2 (9690) 3 (60%) 3 (100%) — 4 (3%) 2, 3 (33%) 3, 4 (190) 5 (4%) 2, 5 (470) 20. Presence of resin gland or its vestige 3 (60%) = 3 (100%) ж 3, 5 (33%) 5 (4%) 2, 5 (4%) 4l. зай of энде arms in staminate 3 (100%) == ps 3 (100%) pleiochas None 1 (100%) E — — "№ = 252 equally parsimonious trees. 7 114 equally parsimonious trees. *М = 42 equally parsimonious tre * М = 50 equally parsimonious trees. T though the shift to the connate condition may not 252 maxi imally parsimonious trees, 9676 of which expected to occur frequently, it seems even i supported H, (Table 2). In thi hypothesis, Dale- likely that the co involucre would split up into сйатріа originated in the Neotropics or western the "correct" number of br. ctlets, in the arrange- Gondwana, a glandless stock coloniz he Paleo ent that recreates the ginal relationship be- tropi y the breakup of Gondwana or dicit tween bracts and pleiochasial arms. Thu e via higher latitudes), the ESS with гез is homoplasy in this character, it seems more likely collecting bees ori in the pics, fol- to reflect convergence than reversal. lowed one or more additional colonizations of The ass E © irreversibility was applied to the Old World from the Ap by gland-bearing each of these T ers in turn and in all possible — (resin-producing) species (Fig. 6). combinations. lad fi the seven dif- other assumptions d combinations of as ferent runs generated КЫҢ ст 3, 2, 5, and 4 sumptio in order of frequency (Tables 2, 3). By far the shortest trees that su of the time Moos 2). Тһиз Г appears = the most commonly supported hypothesis was H,, with : à distant second. For example, the assumption most likel that a sterile pleiochasial arm cannot readily be- are H,, a n- Which of these one ó yoh come fertile a result: equally parsi- depe: to whet assu ar- monious trees that all support H,: Dalechampia acters in general are reversible or irreversible, but originated in ropics or western Gondwana, also on which particu ular character or combination glandless lineage colonized the Paleotropics (by of cha act wi are assumed irove; m mp- ies breaku ndwana or migration via hig tions about cters titudes), mutualisms with resin-collecting polli- have stronge tors evolved se ately in d New than others. That i is, assumptions of spin vena t ominant, in order” ar: Worlds, followed эл recolonization of the Neotrop- of certain characters appear «з by Old Wor ing (resi ucin . The “pecking Species, extinction of the glandless species from apparent in Table 4 is Character 41 > Character most of Africa, and their persistence in Madagascar 20 > Character % a bur of the paganis hypotheses derived from Ssumption that the fusion of staminate The bracts is 15 irreversible, when taken alone, produced TA Dalec hampia originated in the Neotropics o t that parsimonious trees are indicated by polytomies (points of branching of more western Gondwana. (2) The earliest eperies were pollinated by p fromen collecting male ы Bess. Uncertainty re- mains as to when and in which direction dispersal between the Old and New Worlds occurr TESTS OF HYPOTHESES ere are several sources of independent data на Маш Ri used ed evaluate um biogeographic / y ph ylogenetic solu These nios ‘information on the rela- (MA вай Dalechampia, the che mical de- morphological de- tails of the resin gland of Old and New World Spee 308 Annals of th Missouri ن‎ Garden x de Ф = ier du «Гэ, ” « S< a Ыы алт Lu ku «ч =p Sih Ort oe Q а. Іо < am 0D = KF KE ص‎ © cC Ore = оо Оа-—Эх>ш©®©х_ = Z a ш On Has ш ezrpzo-iurmryoS-m Gfici_2 <32 Q22 едш Ыы а ушш ор Ээ UML SO IT O03 9 E "REEDEEEEBREEEESEEERCEDIESEEPECET S $Sz259gUóorouzóot;?smzz0raruzituz ilum agcEzmcuaxus5uOoraouortzomroa5iiuo Ez2 üQr-zzua«lazdao«oozooouvtroorrmrróo x OOD EawBgUudgglBHHuguduHJIHunmimBJuungmgmggrlguug B DISTRIBUTION C] NEOTROPICS AFRICA BN MADAGASCAR is ngth: 11 sieh Lippi: 2 BN PANTROPICAL P f E equivocal 2. fi na : d Plukenetia) RE assumption t all characters are reve: sible, "Tax xa with unstable arrangements in the 436 maximally than two taxa). SPECIALIST HERBIVORES ON DALECHAMPIA f halid b here are several genera o soia iie iei hy- Neotropics and Paleotro (divis Assuming that specialist Pepe Ё ne can make 5e 5 = persal г the continents achieved с afte in the present positions, the but terfly herbivores Volume 81, Number 2 rmbruster 309 1994 Early Evolution of Dalechampia < < 2) СРЕ 2 3 95 = та z= zh Ot = a nos ToO « O4 + к-г 0 ca Mu. ne F отоп acrmcuo«x = = Oopljo Was Ш Eco Du o c Ш z > 5 = O02 EE < шабар О О бошшОо ОБ ш=Ф г HOD OR <1 О e а rootz_f ЕЕ ФЕ: ul uzituzzüzoz- m SOF cOwuzo9« 2 n. rFaóücüomr-ium ESSE «шуша. < < ШОСтхх== 2 IT OO a zm pn Oe.) [6] [9] О ату Ox O10) пш HEBEE и EH H В BBEHEBEBLDDLU TreeLength: PEZ NATION ee POLLEN-COLLECTING INSECTS S м nr ШШ FRAGRANCE-COLLECTING BEES E equivocal FIGURE 3. Same cladogram as is in Figure 2, with pollinati y PI nto the branches. fal 1 dh Old and New Worlds should be completely unre- — ti populati lated. This is because it is extremely unlikely that — into the Old World. (3) p both host plant and herbivore would have become the New World to the Old occurred by a continuous established in the Paleotropics by long-distance dis- population being split into lati Persal from the New World, and the Old World by the rafti t of th | Li I Tf a} АЖ ФА ЭА 14 1 fi it FE (the h | g ap th ( I ) in the Cretaceous, the Old and New members of the loca] butterfly fauna evolving to World herbivores should again be sister taxa, but feed on Dalechampia. (2) If the initial dispersal not as closely related as in the second scenario. à There i from the New World to the Old occurred by mi- ere is some ecological information available &ration of a semi-continuous lation stretching on butterfly herbivores that feed on Dalechampia NC across North America and Europe or Asia in the (Table 4; see Armbruster, 1983; Armbruster & early to mid Tertiary, when these continents had Mziray, 1987; De : 2 the | fly herbi inthe is not yet detailed information available on their ; y А à Old World should be closely related sister taxa of phylogenetic relationships. The systematic (Table the New World species. This is because a semi- 4) and phylogenetic information that is available 215 5 1 ЛИ; d lat hin het n 3 c о» population of host plants across the ind I g s RM igher latitudes should have allowed a semi-con- Old and New World taxa, possibly forming two 310 Annals of the Missouri Botanical Garden TABLE 3. Five most parsimonious biogeographic hy- 1 M tw le. 1 шз 1 1 аг ying I assumptions about character reversibility. Hi: The genus and the resin-collection mutualism originated in the New World, with a subse quent moche. to the Old von fom Ше New and the loss of th mu inb in Ma dics taxa. Н The genus originated in the New World ог western Gondwana; an initial mi: then by a соч migration to the Old World by gl He The genus originated in is [um World o; western Gondwana; an in gration n rafting) to the Old World was followed by the independent evolution of the resin-col- lection mutualism in New Worlds followed by a migration of gland-bearing species to t w d from the Old. Hy. The genus and 9 resin-collection mutualism originated in e Ne ew Wo rld, followe 4 i a migration Ey 1 Old World, los of the m iai sm in и of the п species to the Old World and subsequent recolonization of the New World by gland- bearing species. 1 П FT пири ей manuseript; Armbruster & Мага 1987; Otero, 1 hig no hypothesis of recent long-distance dispersal (see Armbrust & Mziray, 1987, for additional discussion). Th the evidence fro t supports. the pia may som Tragia (Table Ax however, raises another possibility. It is possible that Tra bivores achieved a Mg eene prior to Pass 'hampia. If this , then the lepido be haie pii in the Old pecializing on Dele World could be derived from Tragia-fee estors. In this case phylogenetic relationships mong lepidopteran herbivores may yield insights into the des are history of rie but not of Dal mpia eeding an- EVOLUTION OF THE RESIN REWARD SYSTEM How likely is it that an unusual eui: dt tem such as po ye w resin-collecting could have originated m ки once within Dal. ролат беті еге іп nthe p lant kingdom it has time, si (Guttiferae) AP adi. 1984). T herefore, it seems rst highly unlikely that pollination by in- m a bees would have arisen multiple times in the genus. Any biogeo this would also seem impro ystem arose he owever, makes l origins of Ee. onl seem more probable. ju 4. o opal peream ees р ne. sm lechampia and rela afe and host ublished manuscript), A can (1988), Otero (1990) and D. Lees (unpublish ted ұ genera. Me rela- enkins (un- Neotropics Paleotropics Neptis (T', A, others) Tribe Eurytelini Biblis (T) Mestra (T, D) Byblia (D, TA Mesoxantha (T) Neptidopsis (D, T) Ariadne (T, R, D, e, Ac Eurytela (T, R, D, Te) Tribe Epicaliini Hama ryas Sallya (D, T, others) (A, D, V) Myscelia (D, Ad) Nessaea (P, A) Tribe Dynamini Dynamine (D, T) Abbr eration for pe s: А = Alchornia pa Dalechampia Р: : 2 ЖУА = Adel kenetia, R = veris T =Tragia, Te = Tetracarpium V = Veconcibea. Volume 81, Number 2 1994 mbruster Early Evolution of Dalechampia EC © B c « BD Tr ^ Uu СЕ LI є < КЕ SG c I om EX с.о Ss $ ш Я Es — o S 3 X € s x zu Uu o M X B o o m Oo Eo in o " n BD J BJ Jg M us tree for the singled set of taxa, E 4. Consens id species. distribution ma = RM). Таха pped о with the cladogram and origine of the mutualis hl < < < zi = 21 е) N © Q9 = < Т BÉ Iw а "sS dowd m ow 5 E 5 @ T 0 M M S ыы FG = 53 9 = © т & & S o u un S Ez " RM LOST DISTRIBUTION [__] NEOTROPICS ШЕ AFRICA ШШ MADAGASCAR assuming t that all characters are reversible. The figu with resin-collecting LE e 50 maximally parsimonious trees are indicated 5 (“rein paruum by polytomi n all Dalechampia species that reward polli- si Т : subinflorescence. Compa р holga pes of ны кзн. апа idi deni жа аге "desi | 1 E Use 4 ^ rs (and c ра oo cmb obs.). The bracdets also ete the mixt genated triter- the same compounds creted by glands alon es margins of young leaves, sti pules, e = pie echampia s K & Debevec, sian ae data). Species of Dalechampia identified in cladistic " the analyses as iE ear rly i in the evolution f. H a ence although ня “Character mapping indicates that the r of the rest of the genus also had mmon ancestor sts that resin secretion in a i - this feature. This sugge collect the resin fo plant, secretion, a gland from which the resin could be co . Thus the resin reward inn use place” very early in th it was probably present in the first Dalechampia 312 Annals of the Missouri Botanical Garden < < = =] 4. E О So ss Dor B — о S mor d ue Q.0 . iL И ü ae 8 Ss E o DO ш a 2 MEM Ue m © ~ 3.32 SS 2 oO O O aN ш езеш = * E 2 C de К н GDS Te c S = x 29 3S. = Touda T ао To ou 2 = em = > a Të © [72] oO Oo a = = ” N ü [2| ü ü aj ü [2| 8 a a [2| 0 [2| 0 RM RM „ЖА ям 4 DISTRIBUTION С] NEOTROPICS _ AFRICA ШИШ MADAGASCAR E equivocal :Cad n t character 4l js see е Table Nos uk iind SHARE irreversible. The Ри shows the species distribution Edie onto t the cladogram ig origins of the mutualism неч resin-collecting bees (“resin mutualism the = RM). Taxa with unstable arrangements 42 maximally parsimonious trees are ае by polytomies. to arrive in the Old World, as well as in the N orld species. This is confirmed by the secretion of Bas fense” (non-r reward) resins by many o M as well as several mem section Rhopalos os tylis. Z the stage would have be ше, . Paleotropics and evolut with . Resin- in-collecting ~~ de: once, we might expect to see иэ учы іп D чеш of the gland. If t e of i Pica structure, it would suggest : a Bc origin of the gland. T distinct kinds of resin gland. iiti of арла ROT sn and of‏ رن ew Cr remophyllum have resin glands compo osed 0 esiniferous pr P nd- e llowed by a rec wie pecies bearing the Old World " type (Fig. 5 here are two final aspects of the p e ecology that may have bearing on our ja ol | 9 іс hypot ione First, if Da yen ampia ha relationships with resin-collect pollinato ps э? to its colonization of the Old World ур it spere o Volume 81, Number 2 Armbr 313 1994 Early pO of Dalechampia < < < o 2 £ = ти 22 Mee eS Sob a о г 0 zd = a m EE 3X O0 OQ SS < о 0 R S тох ш E 0 0 ut 2 2 s pco uc g Oo A < < = > = < О МИ С) o Ш < z а. СЕ ES - Gru б оов и 5 X4 EM rl Hd Жак 0 D -g m0 п D 3. "7 8 RM RM RM DISTRIBUTION [1 NEOTROPICS ШШ AFRICA ШШ MADAGASCAR FIGURE 6. Consensu lic a caged Aia gl ш n (“r ualism e 252 четата parsimonious trees аге indicated by т did with lepidopteran ео and it colonized World by some mode other than lo ong- either side ed in the Го нн. ê, not see this; there are not close руне netic ~ Мала between the major pollinators in the e Old. The most likely candi d ; т 5 ates, B poids on neotropical Da lecham- So z z itehe d 90; r, 1992), are not particularly . Mi гет, ш suggests s dispersal Ба nts dista over oceanic barriers to red M to the evolution of the resin » pite ge with H, but not H,), or so: Occurred without the de "following" be mm.). This either was Ene » Or occ; mutualism : ewhat sur rprising feature of H, is that ы t чти Dalechampia that colonized К . The recle — the беж distribution ан onto the delen — RM). Ta resin m reverting to ое y eene cling bees and beetles. This is surprising because pollination by resin-collecting bees appears D be a v ў cessful id pri ion ди danc the genus (some 8076 of Dalechampia int are аа) іп thie s way). р a is occurred, could expect there to b to pills ion by resin-collecting bees in on There might actually be such a гань ас- cording to С. р. Michener (pers. comm.), except e Ln. шола "зр, тнр аге мев too small Arm er & Mzi- ray, 1987; Arahat 988) resin- eating bes are notably uncommon in Mad DISCUSSION AND CONCLUSIONS In this a I have tried to illustrate the ad- ntages of using the results of cladistic analyses to бертын and refine hypotheses about the bio- Annals of the Missouri Botanical Garden geographic and evolutionary history of extant or- = isms for which there are no detailed Fonsi. rec- s. However, many uncertainties remain, and it anl +, ] h hx + ] ae ча x о Ј f Ln EF, : 1 character € The use of molecular data to infer phylogen solve some of the problems E strated here, but even molec- ular bi ( S ders n Å Donogh ue, 1989). T. and po МА hypotheses using inde rin с data (such as оноп оп TENET are dy amd if JE generation. the present study, I used cladistic results to nd heses explaining echa ampia | amg and their р systems. Th potheses seem t likely, and idi aetna’ En are кон d to o distinguish jiu ee ta partie: ular, i of flowers and йййогезсепсе$ would һе у Tabe America in 98, late Cretaceous/early Tertiary, evolved a oil-collecting bees on oil secretion e calyx glands), a more recently colonize e Paleotropics, where the mutualism wit luation of floral morpho ogy of of the PA and Neo рож dio Dou and biogeographical in- к йош is «са y a second tr Malpighiaceae 4: T erori to the primitive лсо states Dalec dues a. It will be in Mee see i d certain ua Ls ша cter: ditional pigen a — to support these earlier p g information was e 1 (3 EL 1:1 J fab г Data {гоа їпзесї m phylogenetic bns lin а best fro herbivores and pol a: T in the New World have evolved only d (Hypo e 1 and 2). The simplest biogeographic scenario is proposed is 1: Dalecha mpia originated in the volve i and Asia, and most recently, Madagascar h * in 1 1 ti h Y g was st. This explanation is similar to explanations of the dispersal history o R & Axelrod at pighiaceae is that the group originated in South origins oge ographic scenarios for NE d Md EE and possibly o oth er plant gro LITERATURE CITED usd: P.R. 1988. w of nymphalid butterflies. Biol 38:9 ее А Hostplants and Am ndens. А a oen scan ма; Pp. 2 а птеп (editor), Costa i Natural ioe Us Chicago Press, Chicago: lino; ова, Therole of resin in angiospern aoe ad Ecological and chemical consideratio! J. Bot. 71: 1149-1160. f mor- ——. 1988. Multilevel “tt ei function, Sd es aa 746-1 а. lossoms. Ecology - & W. Mz sai э 061 clin tion чөк vore ecology of an African ican Dalechenpit e tropica ‘19: 64-1 TEE SENSE A iva K. = STEIN 992. Pollina оп өөө a echampia species (Бр 29: 306- an Natal, South Africa. Amer. J. Bo tion of 1"? & С.І. WEBSTER. 1979. - oo species of Da le зане, HRN pee ie "ne л opica 11: 278 е Pol ——_,, A. \L. Herzic & T. P. CLAUSEN; Volume 81, Number 2 Armbruster 315 1994 Early Evolution of Dalechampia lination of two sympatric species En Dale — WHITTEN, W. M., A. M. с & М. Н. WILLIAMS. (Euphorbiaceae) in Suriname b uglossine bees 1989. Function of AM secretions in fragrance Amer. J. Bot. 79: 1374 esie by male зле bees (Apidae: Euglos- , M. TPs CLA Pollination of Dalechampia nagnoli fl байке) by male граза т” (Apidae: . r. J. Bot. 7 ае 279-1285. ‚кз, E. Epwar Uh se че (U); Lanjouw Lindeman 1114, 180 J.T. Serringa) 12534 (U); Maas et al ifs j nd Prevost 145 (CAY); Reitsma & Reitsma 852 (NY); Sagot 512 (BM, PRS burgk 610 (BM); Service Forestier 3077 (U); Service Forestier 4328 (CAY, P, U); Skog et al. 7427 (CAY, ‚Р, U, US); Solomon 8899 (MO); Solomon & т 24143 (NY, U); W 23523, pedi (ALA). D. schippii Standley: Armbruster 71-303, 78-416, 79-204 (ALA). D. schottii ps eenm.: Armbruster 77-305, 78-409 (ALA). D. shankii (Molina) Huft: Armbruster 79- tig ‚ 91-102, Armbruster & Berg LA 85-128 (ALA); Cuatrecasas ‚ 21512, Davidson 6828, S & Molina à 175. dioses & Valerio 48588 (Р). D. spathulata Baill.: 306, Poeppig 2380, Vigo 20 6480, "im aci lian: 4189 0 Muell. al. 90- D. subternata , 90-150, Arm- bruster bd =%о 158 0.160 90- aon 90-164 (АГА); Croat Winrar. 4. 310 vg 31065, Dorr 3050, Gentry 11801 (MO); p Pac 4181 , US); Lorence 2097 Oy M r & Keating 4527, Phillipson 2492, 3050 (A А, MO у, р. tiliifolia Lam.: Armbruster et al. 85-1 11, 87-111, 87-125, 87-138, 87-141 (ALA); pan: О, N Brit. G 4 (ALA, VEN); Poncy 1808 (CAY, d baies 5IS(By Sad 3248, 7 in 5 э ас & Armbruster 23712 (АГА). D. pie uster & Herzig 85-103 (ALA); Barreto 5058, Henschen үнө н Webster & Armbruster 25182, 5189, 25218 (ALA wom kenetia spp.: pum ster 78-420, Armbruster et al. 85-106, 87- dy: x 113, 87-144, Webster & Arm- bruster 23412 (AL Tragia spp.: Ar ses uster et al. 90-146, Armbruster & Hines 90-157, 90-159, 90-163 "ens Maas 6231 (MO). POLLEN MORPHOLOGY AND Lynn J. Gillespie? PHYLOGENY OF THE TRIBE PLUKENETIEAE (EUPHORBIACEAE)! ABSTRACT ‚ А scanning electron PEE and light microscopy survey of pollen morphology in the Pluk ieae (Eup biaceae) was undertaken to help elucidate phylogenetic relationships within the tribe. Pollen is medium to large, i i or wi defined uboblate, dd olpa characterized by tricolpate pollen with uneven-margined colpi and a perforate to reticulate tectum. Pollen evid supports a division between genera having an aborescent habit TT Astrococcus, and Haemat tostemon) n those with a scandent habit (Plukenetia and Romanoa). Т The synonym stigma with Pl ted. Subtri i i i ll ۲ rphol g аанай а condition ranges diss tricolpate, pias plesiomorphic an к рае ost Өкүш state, to weakly aperturat aes, fragments, or str ands of sexine are usually ш on the apertural membrane, and aperture margins a ^i oveolate, reticulate, үйнө, о ог оган Тһе large к» Tia agia includes seven distinet ^g types, with n и sections (e.g., ун tenomeria, Leptobotrys, Tragia, and Zuckertia, and also d by a uniform and unique pollen erba brc the sectional classification of jeg The other Tragiinae genera have bond үгез from Tragia, with the exception of Tragiella, which closely iov d secti ons O end. faust. Pollen evide псе supports "Cnesmone. eid Ме egis- tostigm а as sister taxa, and sugge chystylidium ll types; th. los INI | Е uggesting t h and the trico olpate species pe rhaps represents a distinct genu. Pollen, E ie pes floral и eine, Supports the hypothesis of section Zuckertia as a plesiomorphic me mber © Tragia, and suggests that Tragia paraphyletic and that the smaller E genera are ded from Tragi € Plukenetieae belong to the Acalyphoideae, fg th largest and least understood of the five eu- lationship are ма nona systematic. iid Phorbiaceous subfamilies. The tribe includes 13 lukeneti genera distributed worldwide in tropical and warm "Р lle n mo RE hs s been invaluable in the temperat regions. Many species are twining vines systematics of th р уана (Punt, 1962; or lianas, both unusual habits in the family; other Köhler, 1965). Light microscopic (LM) observa- ѕ are erect herbs, shrubs, or rarely small tions by Punt (1962), in his pollen survey of the trees. Alth flowers are small and apetalous, Eup orbiaceae, revealed a diversity of pollen types floral morphology is diverse, particularly the style among species belonging to he Pl ieae. The droeciu other m ture isthe present study ollen morphology of the Presence of stinging hairs in many species. Tragi etie ased on scanning electron microscopy L. is the largest genus, with more than 125 species, (SEM ) an d LM w was initiated to help гө polye prob- some of which are commonly known as nose te phy rns. All other genera have fewer than 17 species each; ae relationships. The study 1з the firs of a larger п ; à : : any are monotypic or with few species. Circum- lar project concerning the eaten MAL м warmest thanks to С. Webster for support and encouragement. Much of the work was undertake A rd Ls as a gradu. abi student in his laboratory at the University of Califo bes Davis. The Facility for очна anims entation, University of California, Davis, was ext tremely helpful in SEM we uction and in во solving ms punti; in the SEM work. I tha: nk 1. Nowicke " offering lab ab facilities and p! hotographic supplies, ie V, MO, NY, P, d e Hotton for h The curators at A, “DAV А UC, and US are gratefully б nowledged for gs loan of specimens and for removing or Rai ing pollen to be removed. I thank several individuals the ir SEM photographs, which form part of G. Webster's Systematic 1 Laboratory pollen research collection at tany Dept., UC is. S. Ly! ch was responsible for Figures 23, 24, 9, 39, and 42; А ма 4 for F; , ү, ibl i 2 e оет 19 and 54. All remaining SEM work and photography were isi by the tany Department, Smithsonian Institution, Washington, D.C. 20560, U.S.A. ANN. MISSOURI Bor. GARD. 81: 317-348. 1994. 318 Annals of the Missouri Botanical Garden history of the Plukenetieae, which will include cla- Croizat (1941) and MT Shaw nd Е the two malac- distic analyses based on floral, vegetative, and other species treated v x and BAS characters, lies of ls in ch lution, cense Hook. f. исү а PU of d I lating tl Гапа of BRUM A Baill. E] м. velia s b Sm.) bizarre diversity in floral morphology with age Croizat as the е nation bio! EY: E нарса З, char- genus Ranch “Baill Tus since been i. d ] nonymy under Bocquillonia Baill. (Ac УА а5 Sy: will also be of importance in анс ата deae, tribe Alchornieae of aber genere for whic M Be TS e or its the Plukenetieae (Airy Shaw, 1968, 197 while suc асл Hn s is now jenes red to be a synonym as Dalechampia L. (Armbruster, 1994) and Om- Armbru ee 71091) phalea L. (Gillespie, 1988 » SEE PANE ү classification, Webster (1994) There have been man ions and taxonomic treats Dalechampia as a subtribe, the Dalecham changes in the iis cone since it was first rec- рііпае, of the Plukenetieae, rather than as a distinct ognized istinct group by Bent (1880, as but related tribe (as in Webster, 1975). Although г 7). The circumscription of the genus will not be extensively treated in t the tribe and its genera recognized closely follows present paper, its pollen mop E i that of Webster (1975. 1994) in his classification cussed in relation to the Pluken of the Euphorbiaceae. Two exceptions are recog- In the first attempt at an in pie sien nition of the genera Tragiella Pax & K. Hoffm. and со d relationships, Pax & Hoff- i i the 19 genera of their “Plu- Pachystylidium Pax & K. Hoffm. (following M kenetiinae" into four ноне, groups, Pl Shaw, 1969, 1975), both ET described as formes, Astrococciformes, Tragiiformes, ati € of Тарі ia. кыы Мосор several taxonomic itera чура um ri number and style shape. Webs Webster's 1975 s sis. The mo onotypi qnin the first formal subtribal classification upon de- Vigia Vellozo (which has priority over the mo i ae agiina commonly used generic name Ё) nines A. St. acterized by presence of stinging hairs, a trilocular Hil.) and Fito бине Pax & К. Hoffm. have ovary, and absence of foliar i been synonymized under Plukenetia (Gillespie, found in all habitats but are particularly 1993). Adrien Jussieu’s genus Anabaena has been dry areas. In contrast, the Plukenetii twice renamed, as Romanoa by Trevisan (1848) acterized by abs ingi @ б = 29 5 оа [md = о = ч w 99 Б У and as Anabaenella by Рах & Hoffmann (1919), typically pre n ary due to similarity with its namesake cyanobacteria. 4-locular or less often tr lar, and are Although the cyanobacteria were originally de- found in wet habitats. Style morphology scribed as abaina Bory, ore commonly из used name of Anabaena Bory was re eat Be con- пегіс ES. n. With ж of the gee served against Anabaena Adr. Juss Te Tragia, there На Бееп ап E radiation ^ ter, 1988: 112). Romanoa, th style morphology in the Рен! Within sub- by Punt (1962) and MN Smith (1980), me tribe Tragiinae, many of the genera are charac- becomes the valid name for the e genus. terized and differentiated from 7ragia on the basis The most recent mo нйн of the Plukenetieae of unusually shaped massive styles (e.g-. Sphaero- (as subtribe Рале of tribe Acalypheae) was st ylis, او‎ one Blume, and Tr p by Pax & Hoffmann (1919; with additions and pee KE TUE (1919 1931) based their changes, 1924, 1931). The authors treated a num- ification of Plukenetia on style mor ber of segregate genera in addition to Tragiella о ; and Pachystylidium that were not recognized by Since Tragia is a large and diverse genus it is Webster (1975); these are Gitara Pax & К. Hoffm. necessary to consider an infrageneric classification- considered synonymous with Acidoton Sw. (Web- Eight sections are recognized in the present study s А ч ter, 1967), and урнын Se х (also known The sectional classification of Pax & Hoff г as Angostylidium (Muell pee K. Hoffm.), (1919, 1931) is followed with the following € т ти а апі didam tions. Secti b (Baill.) Muell Р ‚ ай йери rested under Pluken aL. considered distinct from section Tragia, — Gillespie, 1993). A is We rian sections Leucandra (Klotzsch) Muell. Arg - tion recognition of МКК Hook. * following tiga Muell. Arg. are treated as part 9 sec Volume 81, Number 2 1994 Gillespie 319 Tribe Plukenetieae "ow Te Miller & Webster, 1967, and le Romero & Gutiérrez de Sanguinetti, жу "Ako чере 15 кой. 5 eae Mau le Madagascan species. Many of the sections have i in the past been rec- as distinct genera ia Klotzsch, Ctenomeria Harv., Lassia Baill., Leptobotrys Baill., Leucandra Klotzsch, and Zucke Baill.). (e.g., Gillespie, 1 ; Ма de Romero & ed fh Sanguine, 1989). The most extensive survey was that of Punt (1962), who examin M ir genera (ardet to 12 in the lassifi e) in his pollen survey of ЧОК сав types within the tribe ы їп о of Pun - опе M. configuration The diei o configuration is characterized by ob m te or vira grains Lh broad ا‎ a “ru Pachystylidium is ا‎ and includes only that genus. The Cne © pi and lack; Tragia sect. Bia (Klotzsch) Muell. Arg. The Platy- 2. s de Platygyna) type of pollen with an exine at is tectate and Bos reticulate" (defined by f pollen ideni ы the iios i: Nee sect. Bia, the ip matostemon (Muell. Arg.) ce also ) circumscription of gistostigma Miller & We b- ported E S nd сайа 67) кш ройеп of Г Tragia species from nited Sta - evidence from their study of secti d PA uus gy led them to revive na Dara (Baill.) Muell. Arg., which was considered part of section Tragia by Pax & Hoff- mann (1919), and to doubt the taxonomic esed of section Leucandra (Klotzsch) Muell. A METHODS Flowers were removed from herbarium yin of which verifie ET were isolated an n r (1975) was han critical-point dried. Followi ing fia coating with gold or a gold/palladium mix grains were examined and e TAT in a ISI DS130 (equipped with a LaB, filament), Hitachi ridge i ‚ ог Cam 250 scanning electron mi croscope. All pollen was prepared in the above manner unless specified otherwise. In several cases th р, дер ай аї 1 ynology y Department, Smithsonian кб не Washington, “Descriptions of polen | а аге barod ‹ оп ob- ns under І servatio made under LM on 15 gr: medium. The polar axis (Р), 0 x ains mounted in Hoyer’s equatorial axis (E), апо (P/E) о wider than high, and constricted at n base (as distinguished from con- mmae) ical Huic. Twelve of the 13 genera in the Plukenetieae Species of Plukenetieae examined with voucher information, geographical location, р TABLE 1. ies ( equatorial axis (E ie к їп Кн and ре pe es as is " polar ar to леи axes (P/ 5 oras pollen is indicated maximum (mean)) of polar axis (P - and by an asterisk; shortest axis (unde Axis) 1. Pollen with apertures not easily visible under LM is indicated and longest axis (under by “c”: ; shortest and с gus axes are given. Measurements of de pollen à are preceded hy an Species Collection Location P Axis E Axis P/E Plukenetiinae PERRA и Benth. Spruce 2282, NY zil 52-60 (54) 58.5-67.5(62) 0.87 s Benth. Liesner 8693, NY Venezuela 43 2 (48) 0.5-57.5 (54.5) 0.88 = ornutus ы = Liesner 8693, NY Venezuela a 34.5-41.5 (37.5) 41.5 48.5 (4 0.82 Eleutherostigma lehmannianum Pax & K. Hoffm. Cazalet & Pennington 5089, U cuador 48.3-53 (50.5) 3-60 (56.5) 0.89 aematostemon coriaceus (Baill.) Pax & K. Hoffm. ‘urdack & Adderley 43210, NY Venezuela 33.5-37 (35) 3843.5 (40.5) 0.86 Н. guianensis Sandw. Fanshawe 2869, Guyana 41.5-46 (44) 47-50.5 (49.5) 0.89 "ric Ba Sond. Pope et al. 834, M Botswana 34-39 (36.5) 43-49.5 (45.5) 0.80 P. africana Son Wild 5062, Botswana 3440.5 (36.5) 40.5-48 (44) 83 P. leti oes Muell Arg Vargas 18799, US eru 5-57 (53) 60-67.5 (62) 0.85 Р. conophora Muell Zenker 3394, US Cameroon 4-37.5 (35) 41.5-45.5 (42.5) 0.82 conophora Muell. Arg Zenker 3394 Cameroon а 27.(0-32(30.5) 32-41.5 (36 0.84 P. corniculata Smit Koorders 41720, U Indonesia 34.5-37 (35) 41.5-46 (43.5) 0.80 P. loretensis Ule Fosberg 29094, eru 50.5-62 (55) 0-76 (66.5) 0.83 P. lorete. r Maguire & Politi 27371, US Venezuela 48.5-57.5 (53) 57.5-69 (64) 0.83 Р. penninervia Muell. Arg Werff & Wingfield 3173, DAV Venezuela (34) 8 (43) 0.79 Р. penninervia Muell. Arg Standle 708, onduras 35-43 (37.5) 45.5-52 (48) 0.79 Р. polyadenia Muell. Ar, Lindeman 6134, DAV Surinam 50.5-60 (56) 57.5-69 (63) 0.89 Р. madagascariensis Léandri xut o P Madagascar 37.5-40.5 (39) 43-46 (45.5) 0.86 P. multiglandulosa c Cowan & Wurdack 31400, US Venezuela 32.5-37.5 (35) 40.5-47(43.5) 0.80 P. stipellata L. J. Gille Cillesple ы DAV Costa Ric -60 (56 57.5-69 (64.5) 0.87 . supraglandulosa L. 1 “Gillespie Р. verrucosa Smith P. volubili Cowan Prance et al. Y DAV Asplund 14129, US French са Brazil ru 49.5-58.5 (53) -45 (41) 9 0 42.5-50.5 (46.5) 58.5-65 (62) uepuer) eoue; og unossilA 06 eu JO sjeuuy TABLE l. Continued. Species Collection Location P Axis E Axis P/E Romanoa nice (Adr. Juss.) A. Radcliffe-Smith Webster et al. 25436, Brazil 53-55 (54.5 58.5-62 (60) 0.91 pus serrata Vell Bradem et al. im DAV Brazil 36.5-46(38.5) 46-55 (49) 0.79 V. serrata vd Hoehne 29250. Brazil 42-51 Tragiinae Acidoton nicaraguensis open Webster Ortiz 1104, DAV Ni dtm 34.5-39 (37) 39-46 (42) 0.88 A. nicaraguensis (Hemsley) Webster Steyermark & Davidse 116239, DAV Vene 39-41.5 (40) 39-48 (44) 0.91 A. urens Swartz Proctor 36826, MO Jamaica Ф 7-46 (41.5) 8 (44 ЕЗ А. microphyllus Urb. Leonard 5248, US Haiti * 34-40.5 (37.5) 35-45.5 (40.5) — ees iq dem (Merr. & Chun) Croiz. La $ China с 2-57.5 (55) 2-60 (56) — C jen a Blume Morse 5 China с 45.4-50.5 (47.5) 47-52 (49.5) Cn (Merr.) Airy Shaw Ramos С ae 47087, UC Philippines c 48.5-55 (52) 50.5-57.5 (54.5) — ki is (G Pételot 6521, A etnam с 46-49.5 (47.5) 48.5-53 (51 Megistostigna ‘cordate Merr.. amos 17591, US Philippines c 48-58.5 (51) 48-58.5 (53) M. malaccense Ноо! е ahmat roes 13. Indonesia * 46-53 (48.5) 48.5-57.5(52) — Қ ES Ноо Burkhill & rg 15589, MO alaysia * 42.5-53 (48) 46-57.5 (52) — Pachystylidium es ue Pus & K. Hoffm. apey 1748, UC Philippines 27.5-32 (30.5) 30-35.5 (33) 0.92 Р. hirsutum (Blume) Рах & К.Н s& Edaño 49201, UC Philippines 31-33.5 (32) 34.5-39 (36.5) 88 latygyna hexandra (Jacq Л Na RE залга, et ,A Cuba je -44 (40) — P. hexandra (Jac cq.) Muell. A Jack 7 Poir Cuba » 35-41.5 (38.5) 37.5-44 (41) Р. leonis Alain Léon 12176, DAV Cuba * 30-32 (32) 30-34.5 (32.5) — Р. parvifolia Alain Schafer 1427, NY ba * 30-39 (34.5) 35-40 (38.5) — Tragia adenanthera Baill Tanner 672, Tanzania 31-39 (34.5) 34.5-46 (39) 0.88 T. bailloniana Muell. Arg Cowan 2692, DAV Mexico 53-57.7 (56) 60-66.5 (62.5) 0.90 7661 ё JOQUINN ‘1g әшпол aidsalip eeeneueyxnid equ. Lee TABLE 1. Continued. Species Collection Location P Axis E Axis P/E Т. capensis Thunb. Ecklon & Zeyher s.n., US 1170932 South Africa 217.5-32 (31) 0-37 (35) .89 T. capensis Thun Kuntze s.n., NY South Africa a d 3012 27.5-33.5 (29.5) 0.88 T. chlorocaulon Baill iten & Eiten 4285, US Brazil 30-35 (32.5) 34-43 (38.1) 85 T. cordifolia Vah ebster s.n., DAV Kenya 30-37 (34) 37-43.5 (39) .88 T. hispida Wi Nicolson S India 36.5-40.5 (38) 41.5-45.5 (43.5) 0.87 T. involucrata icolson et al. HFP173, US India 30-34 (31.5) -40 (36) .87 1 ivohibe bert 3387, DAV Madagascar 41.5-46(43.5) 44-55 (48) 0.91 Т. blow a, ey s Muell. Arg. Webster 24119, DAV Surinam Е 53 (49) — T. mexicana Muell. A Tuerckheim 7664, US Guatemala 30-35 (33) 35-40.5 (38) 0.87 T. novae-hollandiae Muell. Arg. 56, UC Australia 33.5-38 (36) 39-45 (41 0.88 T. pacifica McVaugh McVaugh 21006, DAV Mexico 27.5-32 (30) 33.5-38 (35.5) . -0.85 T. peltata Kidd dos Santos 1551 Brazil 4-26.5 (25) 1.5-34.5 (30) 0.83 T. polyandr 1 Bresolin 629, US Brazil 31-36.5 (33.5) 36.5-31.5 (38.5) 0.87 T. ramo. TT Ferris & Bacigalupui 8136, DAV U.S.A. 29.5-32 (28) 30-37 (32) .88 1. scandens (Baill: ) Muell; Arg. Humbert 1 ZB Madagascar 31-37.5 (33) 34-41.5(38.5) 0.86 d l. Arg Webster PS DAV razil * 5-57.5(54 — T. smallii Shinners Curtiss s.n., US U.S.A 35-39 (37) 37.5-44 (40.5) 0.92 T. tristis Muell. Arg Anderson d DAV Brazil 32-41.5 (37) 39-48 (43.5) 0.85 T. urens Small Norris 759 S 32-37 (34) 34.5-43.5(38) 0.89 T. volu Webster э л 5325, DAV Јатаіса 25.5-27.5 (27) 30-33.5 (31) 0.87 Tragiella Bar cath (Sond.) Pax & K. Hoffm. Mearns 295, NY Kenya 39-46 (42.5) 43.5-50.5 (46.5) 0.91 T. natalensis (Sond.) Pax & K. Hoffm. Mearns P. NY Kenya a 33.5-39 (36.5) 38-43 (39.5) 2 uapıeo үгошеуод unossijy eee au} Jo speuuy Volume 81, Number 2 Gil 323 1994 lespie Tribe Plukenetieae x pe ү) K | ‘orm 5 i i des and Astrococcus (subtribe Plukenetiinae). “ICURES 1-6, Scanning electron micrographs of pollen of Angostyles and ‚ S see as cor yf Angostyles longifolia. — 1. Polar view.— 2. Close-up of colpus.— 3. e sculpture. 4 : : ату pai ТЕ t. Close up of colpus.—5. Polar view. —6. Equatorial view. Sc ale bar: = 10 um in Figs. 1, 9, 6; ci S st Л 8. 5 = 2 um in Ee Shree to, er EIN Eq e се s -— icher information is in Table 1 unless given in the caption. Annals of th e Missouri Botanical Garden ee ГЕР, Zoe ЛА» ў UN quere : ; a (subtribe FIGURES 7-13.* Scanning electron micrographs of pollen of Eleutherostigma and Haematostemon, bere Plukenetiinae). 7-8. Eleutheros 7. Exi — tT yous E H В ees W. a contin l Лоѕе-ир of colpus in equatorial view lacking a ‹ tigma lehmannianum. —7. Exine structure of mesocolpium of fra 8. Oblique view. 9-13. Haematostemon coriaceus. —9. Cl | Volume 81, Number 2 1994 Gillespie Tribe Plukenetieae ere examined (Table 1). t. endemic to Madagascar, Agirta Baill, was not available. RESULTS—SUBTRIBE PLUKENETIINAE Angostyles (Figs. 1-3). A monotypic genus of small trees known only from the Rio Negro region of Amazonian i “Б @ N E Pollen suboblate (P/E — 0.87), 54 2 um E, tricolpate; amb subcircular to obscurely obtuse-triangular; colpus narrow a , т late Ша, fragmen ted ale irr i vine MN ma rugae with evenly spaced пісготеггосае, С ш o narrow, sinuous to sometimes small anı ircular Astrococcus (Figs. 4-6). A monotypic genus of shrubs or small trees in the u upper Rio Negro region of E la and B Pollen p to obs. E pheroi ] (P/E = 0.88), ms x 54. Е у E s um E, tricolpate; amb ar to curly нче triangular; colpus ie: эи and в d with an en; ex- e et сїаїе- M 1.5- thic ing ic : 2 ег 5-3 um, and 8 tinctly raised at pus margi » with upper and lower exine layers ре ate chamber h arrow, sinuous to small and г des n E od near the colpus. be ii Ontinuous unbroken sexine may sometimes m present over the ا‎ as in polen of PE 'atostemop, (Figs. 11, iscussion п unde I2: refer pt er Hacmatostenon) = Pluke netia) (Figs. "ui as a species of Plukenetia, P. leh- epis. апа (Pax & К. Hoffm.) Huft & L. J. Gil- " Sorian oblate- ee (P/ Е 0 89), 50.5 ий angulaperturate; colpus оаа Nm ends Mi in- di stinct, Жош. 2- -2. 5u m thick, ғ somew hat thinner at us margin; tectum foveolate, surface smooth. Hacia i 7-13). of shrubs or small t Venezuela, both examined. Pollen incen to oblate- veins Mh Е = .86-0.89), 35-44 um P x 40.5 5 um E, tricolpate; iih subcireular colpus narrow, some- with nb n Two species and Amazonas, times covered unbroken sexinous mem- brane, ins uneven; exine tectate-perforate, 5-2 u ick, becoming thicker, um, at P separating оп ап part chamber ш m fir ely fo Vicini ty са А, осии fragmen id: and 4 irregularly gemmate at colpus margin, rugae 192-3 rows Ad evenly spaced кн» intervening muou alla d circular; нк ѕехіпе, when present, Кен and irreg- arly scabrate. Grains within a single SEM abe have apertures that appea e that continuous (Figs. 9, 10) or very а (ог ‚ 12) ing in гон ысу ѕехіпе. Plukenetia зм 1424). 2d pe id 16 trópically with one potus in Asia, fou and азар and 11 in the о о wo species previously treated Vigia are included in the SES > pecies count 13 species examined may be types based on tectum morpho we 1 (Figs. 113) Ро e shine to ob- 1 = 0.80-0.89), 35-56 Р x 42.5-64 4.5 um т te sn into two pollen es becomin, pertur urface smooth or scabrate (e.g., Р. P policia) Of the seven species included in Type 1, the ~ * rtu si ѕехіпе. — 10. Clo 2. CHER -up e col f xis ктай, lacki At T мыла о grain m ae ose-up of colpus of к in Fig. 13; note lack of an арша. sexin -" agus of in Fi —11. ee sexine covering apertu ais, Pola igs. 8, 11, 13; Malo 15.9, 10, 12 Annals d po Garden ; i D pellata oe URES 1 1-19. Scanning electron mic aes of pollen of Plukenetia (pollen Type 1). 14 6. P. M lar vit" Polar i сее ial vie . Exine sculpture.—17. Polar view of P. pe diede -18 = 10 م‎ › с 2 ] afi ana a i 506: ) 19. Equatoria view of ac E grain of P. conophora. Scale ba igs. 15 9; = 2 um in Fig. | olume 81, Number 2 Gillespie Tribe Plukenetieae Ficu Mac ^ Scannir l see ae Se ae m ). A / xg 08 A). hag рр view. — 24. Clogs: -up ы 2 um in gh 24. New W I p orld species, млан оза (Fig. pe Hata (Figs. 1 6), and P. volubilis E) with an ine 2-4 u Гог orld icona E) with a subcircular m that is mostly ipie kened at the colpus margin. l ап exine Ick ¢ Jr Sometimes thie П /» £ L ۰ ‘ ї 20-22. Р. loret 24 р penninervia 10 um in Figs. e im of Plukenetia (pollen Type 2). 22. Equatorial view. 23-2: sculpture. of colpus and exine sculpture. Scale bar: Pollen suboblate (P/ E Figs. 20-24). m 32-55 um Р x 41-66.5 um E, b obtuse-triangular, emer inc Type 2 ( ).78-0.8 lp ite Mrs ёт with margins very unever -tate-reticulate, 2.5-4.5 um | exine semite« thic di muri versely ridged), becoming ‚ їгапу ol- e (i.e fragmentec ү sometimes finely gemmate at ‹ a margin, lumina often smaller near с olpus mar- usually crena six species inc sluded in Type 2 ), Plukenetia Of the brachybotrya and P. loretensis (Figs. 20-22) hav Annals of the Missouri Botanical Garden 3 Clos RES 25-29, е b manoa ато s. — 25. Polar view. sculpture. - Polar view of } large grains (60—7 1.5 um thick, w Ў › ит Е) with an ехіпе са. 4- ме P, аиа шия. Р. pen: 24), P. 7, 8), : and P. шый sized к (39 1 (Gil- verrucosa hav um E) with an exine . Pollen of i Type 1 and Ty c دب с { +] frequently ар реа 1 АК ud & on the 20, 22 small sexinous кы. 1:15 еар апе (e.g., Figs. 14, 2, Romanoa (Figs. 25-27). A monotypic genus up of co gu ol grain less A yi igia serrata (Brade em et al. 8376 DAV).—29. Close-up li) and exine sc abies of V. serrata (Hoehne 29250 A). Scale bar: сеи Ш" 5с maa electron поне of polen of аав апа Vigia neos Pluke 9 nded than in Fig. 2 Р of or: nari igs = to um in es 25. 26, 2 5 pm in eastern ы о southeas of ра woody vines endemic t m El 3 5 pm Pole oblate- bec in (P/E = 54. P x 60 un шы Bias. and ed; exine tectat even jagg um thick, gradually thinning to u ч surfac margin; tectum fossulate- foveolate, surta ; ^os 98 2€ A mono Vigia (= Plukenetia) (Figs. 28, 2 9). | Volume 81, Number 2 1994 Gillespie 329 Tribe Plukenetieae typic genus of vines and lianas of southeastern Brazil, now treated as a species of Plukenetia, P. ek: J, spie. suboblate (P/E = 0.79), 38.5 um P x m E, tricolpate; amb obtuse-triangular, an- ~ colpus prg oad wit ins unev 3 um thick, л coming 1 f. pi. elliptic i in shape with margins uneven, cov- red with exine per quali in (шге к sexine; exine tectate-perforate, 1— 2 pnt t hick: uni- formly thic inal m enm punctate (i.e., having 1 sexine = gmt YS narrow UE separating a and fine ly gemmate at colpus m Tet e strands of sexine were (кибер пеаг tions in SEM (F (Figs. Fh pa but were not visible n LM. RESULTS—SUBTRIBE TRAGIINAE Acidoton ot Five species of shrubs in the West Indies, Central America, and northern South America. Three species were ex- 0 ypes may be ipia css based on aperture presence and exine stru ype 1 (Figs = Pollen p ipis dal (P/E — 0.88-0.91), 37-40 P x 42- E, tricolpate; amb subcircular or obsc ly P б е um thick, y thickened; tectum finely and irregularly зз а -reticulate, lumina round to slitlike and usu р; ч ч © = @ su Sexine islands with surface similar to nonapertural ore irregular. cidoton нави (synonym: A. vene- zolanus), the only species in Central and South America, has зш p Type 1 Pollen spheroidal to Type 2 (Figs. 33-35) cllipsoid-spheroidal, 37.5—41.5 um S, 40.5-44 um dx maperturate; outline circular to pns. elliptic; lat : ine tectate- 208 ate, са. ick; rugae Pride to elongate, sometimes appearir ing ped ан иһ їс sur- usually face «e intervening E OE variable in E two West Indian species examined belong YPe 2. Acidoton mi I (Fig. 35) ap- As to have mor conspicuously beaded rugae n A. urens (Figs. 33, 34) „т one (Figs. 36-39). A genus of ca. 12 les of tw € wem and lianas from south- ‘astern China to abet and Indonesia. => T Species wer, mi Pollen тшен to ellipsoid-sph 5 4T. 5- 55 um S, 49. 3- 56 юа » weakly оа аний circular or elliptic, sionally j irregularly obtuse-triangular; aperture rucate we So Although clearly vi vale gun SEM, the a tures are usually isible und large, poorly define UR od 36, 39); чуй apertural regions of so NAM s are more irregular and do not form distinct кеней (Fig. 37). Megistostigma (Figs. 40, 41). Five species to the ЙҮ and Indonesia. Two species were ined. T Polke dones to i mem spheroidal, 48-51 Sx m L, w eakly redd irregularly te; outline cir r "at eit iter, S 4), large, p aperture or Wreg- ula о а tectàte.perforaté, ca. ] um thickened throughout; tectum punctate and mi- croverrucate, surface often uneven; sexine of ap IBHI f, rs а 1 1 thick, tory o e r islands separated by narrow branched fissures, sur- band sometimes punctate. 1 face microverrucat Т ine are less dense, e areas of fr presumably denkt. areas ie exine that probably r to apertures. In Me- ribu ing patterns such as rings (Figs to he atypical g rain of feo. не. oth In à in me үш е mee leds sometimes di т e (Figs. 36, rly d or in a more random pattern. poa (Figs. 42, 43). А monotyp- ic twining vines distributed from India to the Philippines and Indonesi 330 Annals of the Missouri Botanical Garden Fict 30-35. СПА. 1104 DAV).—30. Equ note presence of sexine islands on d apert 33 E by fossae of variable neg bar: = 10 um in Figs. 31, 30-32. A. Scanning electron mic diss acta T pollen of iine eat ie sur e). 1 релй atoriz | view. : deri vie Close-up of colpus and e EOS per A. Exine sc ulpture showing — 2 сгорћ’ 34. Grain having Pes of monty narrow е — 35. Grain of A. microp 34, 35 2 um in Figs. Volume 81, Number 2 Gillespie 331 1994 Tribe Plukenetieae Pollen oblate- рзы (P/E = 0.88-0.92), late-f. . 47, 48) and 30.5-32 um P x 33-36.5 um E, weakly triporate; т sertim denn a réticidate qom (Fi igs. 49, subcircular to obtuse-triangular, angulaper- turate; aperture circular to broadly elliptic, L/W ae ragia sect. Ctenomeria (Figs m ЛО, d with exine slightly thinner than Section of two species of southern pee н the nonapertural e, but with ted capensis examined her ine; exine tectate-perforate, 1.5-2 Pollen E ii (P/E = 0.89, 31 w E E, Pollen gra (Figs. 36-39), but differ in their circular apertures (Fig. 42) that are usually visible under LM as areas of slightly thinner, more igi exine. In ad- dition, the apertures may sometim appear as a areas under SEM (Fig. 42, ucew on nght Platygyna (Figs. 44—46). Seven species of twining woody vines endemic to Cuba. Three spe- cies were examined. Pollen spheroidal or sometimes ellipsoid-sphe- roidal, 32-41 ит dia erturate; outline cir- = to p liis often irregularly tate-perforate, 3-4 чө tectum E uc u h muri or gae wider than the val slitlike intervening tom surface smooth. " упа б (Fig. 44) апа Р. leonis (Fig. 45) hav a reticulate tectum with broad muri, e^ P. E (Fig. 46) has a rugulate d (Figs. 47-74). Species of herbs, tw wining vine A genus of ca. 130 5, and shrubs found в? етрег па Hen ан around the AB > Particularly abundant in the New World and wenty- pecies une to two sub- : system. sh agia sect. Bia (Figs. 47-5 Neotropical T of six species, of ~ d desertiona and ета меге ех в spheroidal or те т spheroidal, m diam., inapert a wi circular; ^ ү m oig ыен to ticulat ie 9 um t hick; tectum foveolate- e" or reticulate with peius or lumina often "regular mi in size чш sha аре, surface often uneven, ey Basan vaio n in iis i еге is і d^ Perforati i E with 7ragia cau hav- with 3 po rly дле == чш t t, appearing as elliptic depressed area covered with a layer of exine distinctly thinner than the nonapertural ех- h tate-perforate, ca. 2 um thick, muc th цац Кен very finely and о, f umina subcircular “ ойеп narrower than ilie: muri, muri thick, uneven-sur- faced, ponen broken and incomplete, эй о but more irregula r and finer than the nonapertural sexine, sometimes irregularly fragmented. The foveolate-reticulate tectum is usuall the реше, with the aperture ap- lli EM and con- tinuous over pea ring asa Rat thinner area a of exine under ju Ee indistinetly defined areas of thin exine йсй followi e polar axis (Fig. 51). Following acetolysis many P Low apertures split in an irregular manner (Fig. е sect. Lassia (Figs. 59, 60). jos pue section of Madagascar consisting of T. sca. X "Pollen suboblate (P/E — 0.86), 33 P x 38.5 E, tricolpate; amb sica nud pu of narrow to medium width often t attered, irregularly shaped islands of sexine, peed very u and indistinct; exine semitectate- Sha. ca. Es thick, uniformly thickened, fragmented at copus йде sexine islands identical in гау эш to nonapertural sexine Tragia sect. Leptobotrys (Figs -58). The section urat of two species of the southeastern Unit ed States; both species, T urens and T. smallii, were exam Pollen bat sper (P/E = 0.89-0.92), 34- im а m P x 38-40.5 um E, weakly a rcular to us triangular, angula ar to very broadly elliptic wit exine slightly thinn often split in an Bas manner; exine tectate- Annals of the Missouri Botanical Garden Ље Tr? ae’ subtribe Tra" етеп! ] sexi” at the fra te that t у FIG :23 j 3 : ' | | FIGURES 36-41 Scanning electron micrographs of pollen of Cnesmone and Megistostigma iew.—: | 'smone anisosepala. — 36. Polar v 7. Atypical irregularly aperturate grain; no 36-3t : | | | | ; | ; " . а та apertural sexine does not form three apertures. — 38. Close-up of aperture margin with fragmented apertu | | E: Volume 81, Number 2 1994 Gillespie Tribe Plukenetieae perforate, 1-1.5 thick, eue thickened; riable in size; apertu t 55, "ce at top, 57) split in an irre. ты r manner with the underlying mods rane protruding through the resulting hole(s) Figs. 55, aperture at right, 56, 58). ragia i ct. Tagira (Figs ) ion of ca. 60 species, primarily in dry areas of Afric but also found in southwest Asia. Four species Asia. F examined here, Т. adenan AS Т. cordifolia, Т. шоо and T. involuc Polle чынае to Pt spheroidal b Е = 0.87-0. 88), 5-3 m E, tener? inb Subcircular to e е with scattered, regularly shaped islands ағы ѕехіпе, uneve margin very n and indistinct; exine semitec- ri microproje ction: a, nds in rows; apertural sexine s identical in sculpture to nonapertural sex- p agia sect. е йа 64—66). World section о of herbs and t EAE ain. soe a 3 5 š Hanis abu din | in dry tropical areas la ies were d here: T. chloro aul a, T. pel- i. exicana, T. pa iii T. polyandra, T. ramo sa, T. Es and T. M pu (P/E = ud аа 0.83-0.88), 25-37 5 um E, DUM amb obtuse- obscurely so, чү an y broad, often with 1 t veral sc = microverrucae; apertura imilar in sculpture to the nonapertural sex- The ; EN A Bass 15 generally relatively thin (1.5-2 um) uniform thickness throughout, but may sometimes be thicker (ca. 2.5 um) at mid meso- colpium and thinner toward the aperture margins (e.g., T. pecie a). Tragia sect. Zuckertia (Figs. 67-69). Mono- typic section comprising T. bailloniana of Me- soamerica. Pollen oblate- prerada] tA uim - Ss ЭШ 56 3 um шы. obscurely AT parie colpus eo J oo ТНА КА Я ЕА р toward colpus margia: tectum finely reticulate, be- r abrate. t : (Figs. 70, 71). А twining vine endemic to Aus- tralia and the only species of Tragia represented there. Sg oblate- VU UE to suboblate (P/E — 0.88), 36 um P x m E, with three poorly defined kou d ы т use-triangular, e ulap- d aperture circular to broadly еШ .5, with margin indistinct, еі иь a fragmented exine slightly thinner than the nonaper- ural exine; exine tectate- тан, са ит rucate; apertural sexine consisting of small, often ЕРА | ү ое a f. 1 Tragia subg. Mauroya (Figs. 72-74). Mono- typic subgenus Papi of T. ivohibeensis, en- emic to Madagas Pollen oblate- spheroidal (P/E = 0.91), 43.5 P x 48 um E, with - ar t, partially covered е str аі of ѕехіпе, margin very indistinct; 1-1. 5 um pri mostly ак эк kien tectum finely verrucate or microspinulose; dieci No fraguiente ae see similar in sculpture to t The body defined н "t - » ir- regular in size and s t to ern in LM. аіѕсег оа (Figs. Tes m Four E, epocos of twint and оо. ‚А. One species examined. Snapertural sexine at right. mat si Toroes 1389 A).— by fragmented apertural sexine. Scale bar Т ке даан (Ral Mar чод — 39. Equatorial view of Cnesmone tonkinensis. 40-4 — 40. Irregularly speras grain. —41. 10 y 1. Megistostigma Close-up showing nonapertur al m in Figs. 36, 37, 39, 40; = 5 um in in Figs 334 Annals of th Missouri чиле Garden t. Вій Е 42 5с 1chystylidium, Platygyna, and Trag ew t (subiribe Tragii lina e). 42-43. Pa учуй Пит «Rios ij ЕЧайо 49201 UC).—42. Eq uatoria al vie circular apertures, with aperture at right depressed area.—43. Close-up showing apertu ndra- isla nds of sexine at bottom center surrounded Ww nonapertural exine.— 44. Grain of Platygyna icum Bia: T Exine sculpture of Platygyna leonis. —46. Exine sc ulpture of “ena na parv ifolia. 4 47-48. Tragia | m in Figs se vli апа. —47 Whole grain. 48. Exine sculpture. Scale bar: = 10 um in Figs. 42, 44, 417; = ^ e DA 46: 4 Volume 81, Number 2 1994 VEN yr Р; FIGURES 19-50 ect. Ctenomeria. eria: anning electron m > Ti а. — 4 icrographs of pollen of Tragia sect. Bia and Tragia se rtiana. —49. Whole grain. — 50. Exine sculpture. 51 54. Tragia sect. . Equatorial view. 2. Oblique view. $ r 5 ale bar: = 10 ит in Figs. 49, 51, 52, 54; = 2 um in Figs. 5 49-54 5 "ging Tragia sect. Bia: T. less ч е 7 . r t i nsis (Kuntze s.n ) "W of acetol В upti hs Olyze: al 7 ; абы zed grain with ruptured apertural exine. : 336 Annals е алый Garden FIGURES 55-60.* ssp Scanning elec stron mic "rude ч T of Tragia sect. Leptobotry s p Tragia sect. p split 55-58. Tragia sect. Lepto botrys. 55, : Т. иг à Polar view: note a aperture lower right with а 5р f apertural sexine. — 56. Close-up of a ape ture w ^ split spertur s sexine at lower ай on nonapertural е exine at upp® left.—57. Close-up of T. smallii a depr билс сїгсшаг aperture.—58. Equ orial view -i : urens showing circular aperture with ru iptured apertural sexine. 59 . Tragia sect. Lassia: 7 5С scandens, - .0 Ыш eT view. ) 60. Close-up of colpus and exine sc pts Scale Ay = 10 um in Figs. 55, 58, 59; = 5 um in os o Gilles Volume 81, Number 2 illespie 1994 Tribe Plukenetieae T r T. т, Ў Tragia sect. Tragia. 61 i Mo RES 61-66. Scanning electron micrographs of pollen of Tragia sect. — and Zu uis u ©. lragia sect. T, T i 62. Equatorial view. — 65. Exine sculpture + éla sect. Tagira: > > 61. Oblique view. ›2. Equa е vi^ T. adenanthera. —61 1 64. Close-up of colpus in polar view of ittere ! Meven a, xr bs T n and indistinct colpus margin at right. 64-66. Tragia sect. Tragia : ; d el i E 65. Polar view of T. volubilis. —66 Equatorial view of T. volubilis showing coipu E Islands, Se, 1 к i E Y 2 cale Dar: = 10 um in Figs. 61, 62, 65, ӨӨ; = 5 um in Figs. 63, 64 Annals of th = Missouri Ке Р Сагаеп ае. FIGURES 67—71.*_$саппїпр electron mic аа of pollen of Tragia sect. Zuckertia and glad nds novae- i а 67 69. Tragia . Zuckertia: Т. senses na. . Polar view. — 6 . Equato torial view. 9 Eu бач pré e MO . T. novae- -hollandiac.— 70. Polar view. p ose-up of circular aperture Ipt Se ms um in Figs. 68, 70; = 5 um in Figs 69, A е a oblate- куе (Р/Е = 0.91), 42.5 um DISCUSSION Р n E, tricolpate; amb subcircular; colpus ENETIEAE GENERAL POLLEN MORPHOLOGY OF THE PLUK nar > та 1 very uneven; exine semitectate- ште, MATEI OE аки ij ‘ PHYLOGENETIC IMPLICATIONS reticulate, ca. 2.5 um thick, mostly uniformly d in : was found ! thickened; muri microverrucate or sometimes ap- A diversity of pollen morphology Ду арег- еа pearing crenate with ister oblong microver- the Pikes шыра tricolpate, w tructure e str ruraé turate, and inaperturate pollen with exin ly ; Е К to apparently › evidence of : islands was seen anging from tectate, semitectate, in ent, а Ss r LM or in ace stobyze ad bane n in SEM. intectate (Table 2). Apertures, when pres Gilles Volume 81, Number 2 illespie 1994 i Tribe Plukenetieae and Tragiella (subtribe Equatorial view showing 75-77. Tragiella п > Fic Tragiinae), Poorly defined natalensi In Figs, 7: ug Scanning electron micrographs of pollen of T /4. Tragia subgenus Mauroya: T. ivohibeensis. - n: ee strands and fragments of sexine. 3. 76 B nme. 5. Exine sculpture. -76. Polar , 77; = 2 um in Figs. 74, 75. лем 74. Exine sculpture. 7: 7. Equatorial view. Scale bar: = 10 pr ragia subgenus Mauroya 72. Polar view.— 73 view.— 7 BLE 2. Pollen morphology of the Plukenetieae. Aperture condition, tectum morphology (i.e., nonapertura tlined f. Ro genus, including each section of Tragia and ne те type (if more than one). The ones “absent” s includes the condition of very ll f that do not form distinct islands on the Кавли membra Genus Aperture condition Tectum morphology Apertural sexine Figures Plukenetiinae Angostyles tricolpate finely foveolate-rugulate sent 1-3 2 ос tricolpate finely foveolate-rugulate absent—or present and continuous 4-6 эч tricolpate foveolate absent 7-8 matoste. tricolpate finely foveolate-rugulate absent—or present and continuous 9-13 Plukenetia Type 1 tricolpate foveola sent 14-19 tricolpate reticula absent 20-24 Romanoa tricolpate tlt foveolate absent 25- Vigia tricolpate reticulat absent 28-29 Tragiinae Acidoton Type 1 tricolpate finely & irregularly scattered islands 30-32 foveolate-reticulate Type 2 inaperturate rugulate = 33-35 Cn weakly tricolpat punctate mall islan 36-39 Megistostigma weakly tricolpat punctate des лае of small islands 40-41 irregularly aperturate г inaperturate Pachystylidium weakly triporate punctate den: ing of small islands 42-43 Platygyna inaperturate reticulate or rugulate 44-46 Tragia sect. Bia inaperturate foveolate-fossulat finely = 47-50 culat T. sect. Ctenomeria weakly 3-aperturate iy "A dr regularly continuous but thinner, sometimes 51-54 foveolate-reticulate fragmente: T. sect. Lassia tricolpate reticulate А islan 59-60 T. sect. Leptobotrys weakly triporate punctate, uneven-surfaced lense covering ы small islands 59-98 T. sect. Tagira tricolpate reticulate scattered islan 61-63 T. sect. Tragia tricolpate baculate few scattered Suh 64-66 T. sect. Zuckertiana tricolpa finely reticulat absent 67-69 T novae- prse weakly triporate punctate, uneven-surfaced g of Il islands 20=71. L1 He weakly 3-aperturate finely reticulate irregularly fragmented into strands 72-74 Trogiel tricolpate reticulate absent? ord пәрлеку jeoiue3og UNOSSIN ove eui JO sjeuuy —— tS Volume 81, Number 2 1994 Gillespie 341 Tribe Plukenetieae simple, i.e., they lack differentiated endoapertures vera or ora. An unusual feature is the pr different types of weakly defined парце In- variant character states include medium to large size and subobla d sins (i.e., with the polar axis A ben or equal to the equatorial axis). The major 25 of Euphorbiaceae һауе tricolporate pollen; their compound apertures consist of an out- er co pus I an inner os. Colpate, dire and turat ditions are uncommon in the fam enetieae except within the ily outside of the Pluk two subfamilies Oldfieldioideae and Cro fs Aias The ШШЕ аге сһагасї erized by brevicol- porate or porate echinate pm (Levin & Simpson, 1994 P The Cr otonoideae presence of a crotonoid 1994), with the inaperturate Dar a most com- characterized E "n exine аня n is found rarely and only in distantly raed " ADU taxa (e. o RANE thus L., Hymen dia Wall. ex Lind сбн е are characterized рини у tricolporate pollen; however, there are also о е of tricolpate olk ЛК of the Plu. ane tie е ese include three Hy à ers. e à ie and tricolporate aperture dices have ee ribed for pollen. of Ade li bis bauer oton ي‎ ( buy st rman nthus Wall. ex Muell. Arg magia yo mm кае) (Erdtman, 1952; Punt, 1962; pers. obs.). Es ж її ро of the дошу (Dysop- ai defined ap que ollen with three weakly s - t salse ally similar struc- ima different roar би "dr capensis j kara êb comm.) et al, 19 Suarez-Cervera, pers. aol зы бе оге, p on comparison with the dition -— the Acalyphoideae the tricolpate con- Pluken, appea most primitive in th etieae. Given that the most plausible direc- tion о! f evolution of aperture condition in the tribe is Mentes c икан, — weakly Шера, triporate or 3-aper si the vicolpate condition wond ‘stil be considered DI Бош Sah th diti er Aca- lyphoideae, but was separately deed ME the сне condition through loss of the endoap- erture. he genus Dalechampia was treated а tribe of a школе gi Webster (1994) in However, the distinctive pollen of Dalechampia (Punt, 1962; Webster & Webster, 1972: fig. 25)i is very different from any ips. and сагу | (еше The e hypothesis that aa e Plu dieron as suggested other members of the Acalyphoide POLLEN MORPHOLOGY AND PHYLOGENETIC IMPLICATIONS IN SUBTRIBE PLUKENETIINAE Subtribe Plukenetiinae is relatively unifor г terized by tricolpate cat len with well defined ape sometimes jagged margin (Table 2 (Figs. 14-24) has pollen with broad colpi g very uneven, jagged am ob- tuse-triangular and angulaperturate or less often subcircular, and a foveolate or ticulate tectum wo pollen types may distinguished based tectum morphology (Table 2); types corre- spon: pproximately to groups distinguished by Punt (1962), but more nà y defined and restricted to Plukenetia (і.е., fo olate Type pollen with Punt’s Plukenetia volubilis subtype and reticulate Type 2 pollen wi e Plukenetia verrucosa subtype). Species having Type 1 pollen ay be subdivided geographically into a s neotropical group aa erized by large hi that ta apere toward ih ma rgin and a be ed grains exine рие. а a thinner exine that i is ‘uniformly thick or ot de 2 aper ja 342 Annals vide seni Garden a b c А а С0) (@) j i IGURE 78.' Evolution of aperture ore d in the Plukenetieae: a hypothetical character state tree. be that only one aperture is shown in equatori E nd th aly y ill na ture margin ology.— 1 dition (Dalech ost Acalyph mie Бе? Tricolpate (all pipes jE M iie Efe renis deis E: Tricolpate iiie red зе islands on embrane (Tragia sect. m apa sect. Tagira, n A pe = = ы e. apertures w with Че covering of small sexine i sia ds (Сп ne, Megistostigma).— de aperturate. mo cov T ring of small sexine islands (Megistostigma) ks “Tnaperta ae eee pa T 2, Platygyna, Tragia se 1 Typ with thin ae of exine, aa TRE as depressions (Tragia sect. Ctenomeria).— TE Weakly triporate, apertures , Tragia sect. Leptobotr rys, t. Bia). =i eakly 3-aperturate | j. Weakly 3. pec apertura vered areas covered with strands of sexine (Tragia subgenus Mauro oya). thicker at the aperture аа Species havin, 8 m the Neotr ropics, he opical species, pol- OEE og » with thee of чуе nis sion major species gr oups. Pollen morphology of the monotypic genera oo ee (Figs. 7, 8) and Vigia (Figs. 28 2 w of variation found in © = Ф о аа Рае s ctum. Althoug morpho cally very similar to Plukenetia, the genus may AE pistillate me both {урыны “ characters in subtribe. Romanoa tamnoides appears to be ae the sister taxon of Plukenetia, which is сһаг- iz 4 nd 4-part rted calyx, rucate tectum. Colpi are narrower with an even ut not distinctly jagged margin. The three genera shar ilocular ovary and a tre sh es very different from the vine or liana habit of pe kenet d Romanoa. Both Astrococcus Hae mon have unusually thic ed арег separating to form an elongate cha wre um). Fo rnin together wt { ns, a unique characte *, r stam tion- in the subtribe, iui their very dies rela Volume 81, Number 2 1994 Gille 343 espie Tribe Plukenetieae ship. In contrast, ee n lacks the thickened aperture margin and elongate chamber and bear: staminate flower while ш Sls ossia на the ^as two vo related and Ango the ember sharing with сао the plesiomorphic ne racters of пи s stamens and absence of h Astro sa oste- mon a “tectum perforatum" (as ор to a "'psi- late" tectum as described for othe cies in th margin) as in Plukenetia (except P. conophora). POLLEN MORPHOLOGY AND PHYLOGENETIC IMPLICATIONS IN SUBTRIBE TRAGIINAE E abt ribe Tragiinae exhibits much greater diver- in pollen morphology than subtribe Plukene- Pollen is ARA ina ешигин ine wit ih poorly defined apert ctate, foveolate, carats тебе: and bac ulate conditio ons (Table 2). Pa chy. stylidium an з Р laty, are charac у unique pollen types; анон includes two distinct pollen types. The large genus Tragia very distinct E eneric status of Plat a has эя i 1971; Borhidi et iin 1973); 0 characters, a а gl tacle wed ^s ould seem i бетер ip, suggesting that the similarity in pollen x ipn ularly the inaperturate condi- rin due nvergence. e ii PR i was found to contain two distinct pollen types, differing i in aperture presence ra, have colpi with uneven margins and apertural sex- ine islands (Figs. 30-32) dy tips € f th ide 35) ар re si ll its hara than to its mainland cong ingly, P. parvifolia €— — in- termediate between the two gen ha an the rphology, pe penus would s SP 938 un Ф The three Indomalaysian genera of subtribe Tra 6-39 shape and size Расумуйайшт (Figs. 42, 43) has three беши, u smone (Figs. 36, r four) colpus a is char- or with regions, W alaccense is weakl irreg- ularly aperturate or inaperturate. Poll evidence supports the hypothesis based on floral morphol ce morphology and presence of a staminate ap- 344 Annals of th Missouri cle Garden endage appear to break down as more species are d e the genera. Since pollen of a is in- termediate between. pollen nai ne and Me- arc. 5 pat tial ial sexine (and usually not visible under LM). The ly covered Rue D spegies арреата medi simt in b гагы (R: novae-hollandiae and section Leptobotrys), B Pp U either recógnition tylidi typic genus lovin Pax & Hoffmann, 1919; Airy Shaw, 1 1975) or as a species of Tragi owin “soe a кашыр with its oo neigh- agascan section о dial primarily in style morphology. Pollen of Tragia sect. Agirta needs to ka ылар to confirm this Mer relation- ship. econd pollen type unique eee P: Tragiinae i is found in the southern Afric xon Tra, E ie (чаты so nth. wie re- j Ашай v Trage in ahei spite a very y Т staminate flower morpholog The Old World, рн йокы ан; Мне їаха, Tragia sect. Tagira М re a sim mo нбни (ТаЫе mad Polen is pur wi ith a reticulate pon иша t он» fr | apertural membrane Moran absent in Tra- th n Lassia is mor- phologically ny similar to Чорна ani may het ha di by single apomorphic androecial character. Sphaero ostylis (not examined here) also ha colpate with a reticulate tectum similar t the above according to observations by Punt MK "This evidence i is consisten j (Шош асве in calyx and foliar morphology, as — - by Radcliffe ‘Smith (1987), wouid rather ож nis з Megistostigma (following dor & Hoffmann, 19 Two тает taxa of Т) unique pollen types very аай from the ae African Tragiinae. Tragia ivohibeensis is an en- demic Min gascan өрөн for which Léandri (1971) ted us Mauroya, —Ó e he Чоп ught it intermediate betwe Tragiella and араш, i wo sect iong of Tra, agia, AA and {ТЇ тү eap e er, ьн ot sonia side Tra: giella y specie: ragia examined. The iind (sedis д аге very poorly defined Joh eres | | n the Old World), the БП һаз а уегу distinct with лийза; of numerous u ic across the aperture (Figs. 5 в unique in the tribe and cann to other types, emphasizing the Fae. of sec- tion Ситат гіа. 2А somewhat ponian pollen m pes genus Dysopsis (tribe Acalypheae) (Рала ; however, similarit y with sect. CBE is most likely due d con- vergen cese dins pollen Sot are found among neo- tropical species of Tra, ction Bia is the only section of Tra ui ia pollen (Figs. 47—50). мш ана ahi and stamin ens emphasize. the distinctness of section ا‎ ond in its more cke similarity with Romanoa, (1 — E that the species was one 0 a. Since presence e of sti is 3. branched pies ie dE hairs and a sle baia in Tragia and not in s Volume 81, Number 2 1994 Gillespie 345 Tribe Plukenetieae netiinae, pollen evidence supports the и эрез еа d Plukenetiin llen is simila Tragia sect . Ta agira in having apertures with ver ttered lands of se on the aper rtural nie e, b iffers in exine structure and broader colpi ith usually fewer sexine islands. ile d as operculae by Miller & Webster ои оп the bow in shape, size, "m pe and e nnot be considered to Erdman’ s (1952) definition ait an oper ipeum à "thickening of measurable bulk and clearly de. а of an aperture membra ne (circ каен їп роп, elongate in colpi syy of this large EM gl ly gl | were examined, LM obser- d Miller & Webster (1967), and R. oom ap ар р suggest that this pollen type is Tragia and represents the чк оссиггепсе in intectate alá (sensu Walke i s (Pax & > Arg. (resurrected ( ч 989) fo ero & Gutiérrez de Sanguinetti cludin T several So merican species, in- «fins. Polyandra, but treated by Pax and Sections niil sectio: andra) ce these gia ( Share the same pollen type as section Tra- Sensu Pax о 2 ding T. pacifica, Т. E ss and Т. volubilis), pollen N RDA ins not Support oe wirst ае as distinct sec- „= ө pollen type among New World Tra- fid ound in Tra agia sect. Leptobotrys (Figs. 8). Pollen has three к oorly defined ees apertures es with a very indis inct margin and co or baculate onion of €. The distinct apertural condition, tectum morphology, and apertures densely covered with sexine gii pue vs ег & наз "s (1967) 110 rphology and rights bd рее пей Mueller s s sec- tio: in section Tragia (as done by Pax & о, 1919). e pollen of sect. Leptobotrys is most similar to pollen of vae-hollandiae (Figs. 70, 71), the only species of “Tra known from е a, it is anomalous both in that sec «зй; section aE) The two taxa Mure a Med м E © S obtuse- triangular bre iain 2). oar he most closely е the southeast Paci voce d tectum morphology, both apertural and nonaper- ke e Shaw (1202) ош = а = їр be h ун he» - jenes that the species is transitional betwe e lp ai Tragia. In addition, Tragia ect. Leptobotrys and Pachystylidium share stamen пи r of two, an unusual condition in in: tribe. Whether similarity in pollen morphology be tween these three disjunct taxa is due to homology ds to be examined further. or convergence nee APERTURE EVOLUTION IN TRIBE PLUKENETIEAE synapomorphies definin Pollen g the Plukene- tieae are trico Іра ate аре erus Бенин od uneven margins 4 shape, presence of an ае, and presence of ап mbrane here has em exine next to ~ been an unusual apertures in several evolutionary lines. Apertur margins have become more irregular and less E 346 Annals of the Missouri Botanical Garden fined, and fragments or distinct islands of sexine CONCLUSIONS are often present on the apertural membrane. The majority of species of Tragia are tricolpate; The use of SEM in the present ким of pollen i i T. bai morphology of ш Blukenetines has ip ovided w loniana, have scattered islands of sexine on the 1 і 1 ins (Fi RM ‘than was possible with TM акб In [ИУ | J © ` 78c). A single species of Acidoton (Type 1) also ular much more has pollen of this type. highly resolved and enabled detection of unusual ечи е different “= of weakly defined aper- ар Е Sev ral taxa tures are found in the Tragiinae, each presumably originating “independently from the tricolpate con- Punt’s (1902) чш based on LM alone, were istinct when viewed ип M. These include Tragia capensis EY н Punt with the RES volubilis rei Tragiella a nd Tragia ouped w the Plukenetia iid typ ORR m T 2 "o with an exine equal in thickness or = slightly thinner than the nonapertural exine. These areas may be either e (Fig. 78d; Cnesmone, Fig. cosa ORA a i cido ton i ig а hy 1 trys, Figs. 57, 58, a nd Tn no жайлана, Fig. — supports the hypothesis of Dale ере um z d here), rather than being deriv onh within the tribe. Webster’s (1994) treatment subtribe within the Plukenetieae un о б © 3 ge [od бын o dv © =” = © meria (Fi , which has apertures covered with of the genus as a continuous or sometimes fragme омї, кык is consistent with either hypothesis thinner exine, often visible as depressed areas (Fig. Pollen evidence is consistent with Websters i). Tragia subg. Mauroya (Fig. 73) represents (1975) division of tribe Plukenetieae into two sub i no 78 a fure type characterized by very weakly de fined tribes, Plukenetiinae and Tragiinae, but Hoffmann with and identical i lpture to th p l four informal groups. Of these groups o exine (Fig. 78j). isi Inaperturate pollen appears to have evolved at matostemon) is mo twice in ibe Tragiinae. In the Old World, gia) is paraphyletic, while Р inaperturate pollen (Fig. 78f) evolved via weakly — Plukenetiinae plus Acidoton and Pla a) and defined colpate (Fig. 78d) and irregularly apertur- Sphaerostyliformes (all other Tragiinae) appear to ate grains (Fig. 78е) and is found only in Megis- be polyphyletic. tostigma (note that oiu condition is Proud Pollen morphology supports a subdivision of sub- within the genus and within species, e.g., M. ma- tribe Plukenetiinae into an arborescent group (4л laccense ha indi! irregularly jure rns gostyles, Astrococcus, and atostemon) and and inaperturate grains, whereas M. cordata has а lianous group (Plukenetia ares in es Also both weakly tricolpate and irregularly aperturate supported is the recent synon а and grains). Int ld, inaperturate pollen (Fig. Eleutherostigma under Puen (Сезе, 78h) originated опе or more ti st probably 1993), and the relationship of Astrococcus and from tricolpate pollen having scattered sexine is- — Haematostemon as sister taxa. la т\ the apertural membrane (Fig. 78c) and ubtribe T hibits an exceptionally © Ф is found in three taxa, Platygyna (Fig. 44), Acido- verse pollen morphology, with much of the Type 2 oe pe 35), Ex Tragia sect. Bia ation present in the large genus Mice pole бы. 47, 49). А i would be mor tpliology; ior the most part, support s th nde ‘directly; n tricolpate grains lacking ap- f Tragia. The two lar; - 78b). Acidoton Type 2 tions, neotro ia and paleotropical Tag inen гузару Platygyna) к, most likely orig- “з wala ae n hs attered sexine e isl m Aci Type 1, i.e., pollen having оп the apertural membrane, mu are a de ved “ihe reden кше йбн whereas guished ba pe on exine morphology. sections Eu piae үе s Маигоуа аїе way for the origin main ining taxa, of inaperturate pollen i in uis sect. Bia botrys, and Zuckertia, and subgenu Volume 81, Number 2 1994 Gillespie J р! 347 Tribe Plukenetieae each characterized by a unique pollen type, pro- viding evid ndeed distinct natural RU an a ples ьо member por ining genera in subtribe Tragiinae, for EG are characterized 7 unique pollen types distinct from Tragia. The southeast As sian two LL pollen types, EE and tricol- Pate; the close similarity of inaperturate Type 2 pollen to pe of Platygyna suggests that Acid- n may yl Tragiella has pol- en very similar to ые t. Tagira, and would thaps be best con e a section of Tragia Heim to section Tagira. ollen and floral pore evidence indi- ic dod a о reorganization of g c delim- the Tragiinae to better Uu Hec Edel hips. Tragia appears to be a highly aphyletic genus as presently circumscribed. of the remaining Tr ed Nyletic. Вепега appear to be derived with re t to Tragia and defined on the iow of unusual uid минан тее style a al gnize certa and Ctenomeria, as distinct treated in the past, e. Baillo P = п, 1858), since these sections are as distinct ra Benera (as th | L V the Tra Я | giinae genera "M did ia 05 а A сор маа "a of Subgenera a The level а € taxa such as Tragiella, упа are ipee followe exhaus Pans tive; some taxa need to be more thoroughly Tra iA died the two largest sections of Sia, nd Tragia (to verify th ya м pollen type is found in each), while several @8ascan taxa still need to be examined. The — of pigra ие suggested his paper to urther tested by means of койыл, clic using бка, у ve and Lia len characters. Taxa shar hology, but not previously tecto to a. ей relate 1 nee mo oroughly examined to determine if ea sins are homologous or the iue of converg mplete un- du of the Т ee ets e aa is recommended r to making specific changes the generic a ан тыйы. LITERATURE CITED Airy SHaw, Н. К. 1968. Notes оп Malesi d oth Asiatic Euphorbiaceae. XCII. ое on the affinity of Ram е b к 01-403. п and other Asiatic subtribe Pluke- x. Kew Bu E 23:1 Euphorbiaceae АС LN Bocqu а Baill., dte a revised key to Kew Bu "i 729: 315-322 15; The Euphorbiaceae of Borneo. Kew r. IV. the i Bull. A ARMBRUSTER, | Early evolution of Dale- champi biogeography souri Bot. Ga zd ВАш ОМ, H. xw générale du groupe des cec ae Victo son, Paris. BENTHAM, С. 1880. Eu praca. In: G. Bentham & JD H - AE 94. Av dime, and comparative ecology. “Ann. Mis- : 302-31 id m d. Sci. Hung. 19: CROIZAT, L. The tribe Plukenetiinae of the Eu- irr! in eastern tropical Asia. J. Arnold Arbor. 22: 4 e c 1952. Pollen Morphology and Plant Tax- y. Angiosperms. Almqvist & Wikse ll, Stock- нш 1994. sopsis Baill.: Relations "phylogénetiques au с eae (Eu phorbiaceae). Acta Botanica “Calica к >» 41 пуза А со І. J. 1988. revision and phylogenetic analysis of Capai E. тизе sed Unpu ublished Ph.D. Dissertation. University of Cali ornia, Davis. vs synopsis of neotropical Plukenetia new species. Syst. Bot. (Euphorbiaceae) including tw 18: 575-59. GREUTER, tod ET AL. (editors). 1988. нее Code of Botanical ыы Regnum V di Die Polle pmorphologie der e sie g für de Tax. 348 Annals of the Missouri Botanical Garden LÊANDRI, J. 19 Un sous-genre malgache nouveau de Tragia dee не, Adansonia, sér. 2. 1 Levin, G. A. & М. С. SIMPSON. Phylogenetic 1994. implications of pollen ultrastructure in the Oldfie vend deae i Wi one Ann. Missouri Bot. Gard. 203- a Novitates Antillanae IV. Eu- 123- LioGIER H. podia Mem. New York Bot. Gard. 21: Eres. 8 P. & G. L. WEBSTER. 1975. A new technique of | preparing poi for scanning electron microscopy. Grana 15: 127-13 MILLER, K. I. & E dh WEBSTER A preliminary а (Euphorbiaceae) in the United . M. GUTIERREZ DE Ж ЧЕ рч taxonómica de ntina y regiones аа : 77- 1994 palynologieal study of aed e : Euphorbiaceae) Ann. Missouri Bot. Gar: ae Euphorbiaceae- ne K. QUSE dus In: А. Engler (editio. Das Pflanzen- reic S e 147.XI. (Heft 68): 1-108. W. Engelmann, Leip: vod Euphorbiaceae — Addita- ntum IV. dy л, ТА. Engle (editor), Das Pflanzenreich Iv. 147.XV11. (Heft 85): 179-204. W. Engelmann, tanda eipzig. ————,. 31. Euphorbiaceae. In: A gler & K. mee HER Die Natürlichen s zenfamilien, ed. 2, 19c: 11-233. W. Engelmann, Leipzig. PuNT, W 1962. Pollen morphology of the x = ы р special reference to taxonomy. Wi x ас зж А поїе оп Bennen (Eu- ing сев) Ken ^ is 34: 591- s on African Airlines XIII. Kew ae i үт -691. Euphorbiaceae (Part 1). /n: R. M. Polhill (akori Flora of Tropical East Africa. A. A. Balkema, Rotterdam. RADFORD, A. E., W. C. Dickison, J. R. Massey & С. R. BELL. 1974. E edi Systematics. Harper а bie Аа a. rk. REV 1848. Saggio di una monografia delle эше Cocco Padova. ER, J. W. & J. A. DoyLE. 1975. The bases of angiosperm ие Palynology. Ann. Missouri t. Gard. 62: 664-723. d (Euphorbiaceae) in ouri hi rd. 54: 191. ess of Con the Euphorioee Tom 24: Synopsis of the genera si suprage- neric taxa t Ес Ann. Missouri Bot. Gard. 81: 33-144. . S. ARMBRUSTER. synopsis of the neotropical species of FAR (Euphorbi- ae). J. Linn. Soc 105: ШООЛА, Amer. J. Bot. 5 CONTRIBUTIONS TOWARD A MONOGRAPH OF NEOTROPICAL JATROPHA: PHENETIC AND PHYLOGENETIC ANALYSES! Bijan Dehgan? and Bart Schutzman? ABSTRACT Phenetic Lo of the Mor Won Jatropha lis species generally supports | the dm classification of the genus by Dehgan and Webster. Т of section Pe prea phae Pax ciet sections of "s Ms tega) Pax a M ch eam for subgenus i. s tropha, sections у-га LN E and probable paraphyly subgenera, and the clos Жалла nship of sections Mosinna ed. ا‎ E pro ovided evidence of monophyly hyly in subgenus Curcas and severa The ong ge topologies. sectio : o E ant subsections. The cladistic analyses described he erein жиз ы ultiple ۲ ETE ce Hy place r recently d 4 E ES | oY үз Pa ET sg d Ё gh geograF : phylogenetic er a distinct correlation between een d phological f. postulated infra- generic delimitations, and geography o Evid i dt t the tiquity of кч genus and its S pres ent distribution, hinh ж. Br 2.3 {елер ums p * 44 | F р‹ K rican and ] appears to be the norm for the he p S, (x) ыу не orl Pons in concert with migration to areas of increasing pius and cold гасова (1763) was the first to note that Jatro- аз proceeded along two evolutionary lines а is recognized the two genera Jatropha an 68 5, based on Linnaeus’s ا‎ es Plantaru a Although Adanson’s generic delimitation d incompatible with өн of later taxonomists ; Mueller A (Pohl 1827; m illon SE i cations, ee s four ешш into their pu €m and recognized them ' Flo orida oed Experiment Stations Journal Series BAA, ВАВ, » LA, ue ЫАМ. LP , LPB, M М, MAPR, MEXU, мон ‚ MO. mo oans of her ra, ua for P naive and Joannesia, to chael Huft, Walter Judd, Geoffrey le, Florida 32611 сеге danke are ex M in sa, Peri 1 0 Mi Levin, and Alan gs iu Systematics Laboratory, Department of Басайын Ноа, , U.S.A the subgenus Jatropha. Section etin (Ortega) ess equivalent to the a of this study and is presented be Two subgenera Ссн а pads а), ten sec- seme and ten subsections were re to ac- inimodate the Old and New spell era lyx aestivation, corolla coherence or conna plant sexuality (monoecious or dioecious), stamen rr. e, and pres siomorphic, or ve, in the genus, eras its palmately lobed leaves, No. R-00642. We thank the curators of ASU, B, BA, F. GH, GOET, HB, ICN, IJ, JBSD, B D te CEN, CEPEC, CHPA, COL, CORD, DAV, ENCB, F, GH, [ р, SP, TEX, TRIN, UC, UEC, US, USM, , NY, R, RB, SD, SP, tended to Grady L. Webster for contribution 0 of eerow, H Eug and Dena Gar vue for technical "University of Florida, С ANN. Missourt Вот. Garp. 81: 349-367. 1994. 350 Annals of the Азин Botanical Garden Shore roi ebi possessioni of a carat main- lorescence), one occasional hermaphrodite flowers includ ding sev tures support this classification. Both pao and nonarticulated laticifers are presen bers of the и Ja atropha, but anii laticifers all tax eral anatomical features that it shares with mem- f both subgenera. Fyon on from it or a +h habit, pes ш. of — coupled на habit, exemplified by J. EES L. . (eet tropha, subg. Jatropha), or us su shrub habi ti IN ч f abit , Shown b zinna (Ortega) Pax, бе Checda). These ue, were accompanied a series of reductions in Bak omen € of ais "puc iir Cv olutionary бодожо, two RG pathways. On NE one hand, ladia (ind k ey disi int P ( the formation of a highly КЛ compou und di- 0 a chasium, via section Pol rphae Pax ti Peltatae (Pax) Dehgan & Webster (both strictly merican) or the coflorescence was reduce a пз Cur. cas and Plat E Idioblastic- Mas M, mbered Suet imm are char- in subgen SEHE: of t s Curcas but co morphology indicates the presence at Би acytic stomata in subgenus Ji paracytic type in sul a Curcas. БИр hairs f t eu te- urcas are unicellular and verrucate (Dehgan, 1980; Olowokudejo, 1993). The number and ar- ingenio G petiolar vascular bundles vary from (1 a ring, a an in as free traces, medul р er, or "s т free ог medul- late * pit traces (Dehga ybridization ay рона (1984) ке ccd evidence that t taxa furt her erin in the ive y less ie ara of н pho more closely 1T10) ed lenucia ЫР . On the , infl positi rescen > persis markedly reduced to a few or hi p unilater. al compatibility but to be ا‎ ozinna, subgenus Cur- separated by pe ا‎ ntial — whereas the cas, in concert with a gradual shift Жазга monoec more phylogenetically dist taxa were incom- to gynodioecy and dioecy. These modifications were patible. Jatropha curcas ich, in agreement often accompanie п osom with McVaugh (1945) and Wilbur (1954), was mbers, from diploidy to tetraploidy, nearly al- considered the most primitive species on morpho- ways accompanied by acquisition izomatous logical and anatomical grounds, proved most com- nier habit. Evolution of ше flowers also pro- patible with species of sections with pres ably two i nsubgenus closest affinity (e.g tegerrima Jacq. of sect. Jatropha, red ti d g Fe olymorphao. and least segs with taxa 0 as occurred t gl 1 fı ions (e ipae Jr gerer pim to biseriate, connate to free, with no concomitant of sect. Jatropha ioica Sessé, o change in the num ipe " le nd ке of the — Mozinna). It was Bee ria v phy: vary, all with thr and 3-locular Кыныр distance may be reflected by а ability ovary (with the notable exception of two bifurcate е species to interbreed. les of J. martiusii sii (Pohl) Baillon). In contrast espite these convincing lines of evidence SUP s in orting alignment of the taxa, placement of -m Scarry o YP s exhibit a morphological continuum from south to north, ea пе so outhern species the more advanced feat Several anatomical and ME I aes fea- 5р living ts, incomplete herbarium specim nt knowledge of the South American p^ ere of e n& usin vailable information for the neo ор iate ea ors -— also interested in examining the p delimitation and assessment of relationships th Volume 81, Number 2 1994 ehgan & Schutzman 351 D Neotropical Jatropha interpretation of the results of phenetic and cla- distic роге es. rhe: intent ok this paper is not to present a f | the N taxa; we know that Ne constitute a Er monophyletic group, and that Old World ta i о crea cladistic iun tion cladistic m phenetic оше: i — Ror for corroborative purposes о only. With a brief d cussion of geography and н added, at is intended asa P cont pibutión toward a mono- h component in each of the three PCAs, were cal- culated and are listed in Table 3A- in Two discrete groups of OTUs corresponding to s pha and Curcas wi near Ee. А. ORDINATION AND HIERARCHICAL CLUSTER ANALYSIS MATERIALS AND METHODS component ordination and hierarchical variate Statistical program package (version 1. The data matri 77 taxa by 32 char MS is shown in Table а Although thi three-dimensio nal scattergrams, д. ud three- ional scatter, е; were constructed from A fac mputer program w “Т by the pe author. Among NE features, 1S pr = user to identify individual taxa by their iden- tification n number in the data matrix. It also has he a and аА ida e id easily resol Data matrices are input to the p Con wish standardizes raw data (“z-s 973), шы o penc and projects aude onto axes for display. Schutzman's program can read and display the d алы files from NTSYS- Pe with only minor modifica RESULTS AND DISCUSSION The results of PCA and hierarchical cluster ж firme Wa. pene fewer (1979). Factor loadin ings, signifying the ost meaningful RRS for each principal 3 3 ~ eri Gre species placed by рез п £ usn subsect section Polymorp hie, has o been confirm ilio results of the phenetic ата presented here. section was based on ion Hern Webs 1 a in Baja California), J. bullockii Lott (with 8 Rue stamens, endemic to higher elevations of Annals of the Missouri Botanical Garden TABLE 1. Characters and character states for ordination and hierarchical cluster analysis. 1. Growth form: 2. Caudiciform habit: 3. Rhizomatous habit: 4. Plant s - Laticifers: 6. Laticifers: 7. Leaf outline: 8. Leaf base: 9. Leaf basal glands: . Leaf margins: 11. Leaf marginal glands: 12. Leaf sinus depth: 13. Leaf lobe numbers: 14. Leaf veins: . Petioles: 16. Petiolar traces: Stipules: - . Hairs: 19. poner ant gi 20. Flower 21. Flower Reis 22. Petals of 9 flowers: 23. Petals of à flowers: 4. Stamen arrangement; 25. Stamen number: 26. Stamen fusion: 27. on fusion: 2 x of 2 flowers: 29. Calyx lobe margins: 30. Fruit dehiscence: 31. Fruit locule number: 2. Seed: Trees me ties "A used (2). Based on strict b 1 ess of 5 meters, a shrub 1—5 meters, and a subshrub > Pant TM ja Pia typical (0). Plants with thick d sub m (a шдеп audex). (1), Y typical (0), Plants with spreading rhi us (1), monoecious И nd/or arie (1), ое (0). S articulated and nonarticulated laticif ut only some have idioblastie type. Nonarticulated (1), articulated (0). Plants may io both articulated laticifers but nonarti Lanceolate (0), ovate-lanceolate (1), ovate- clipti ps ч ен: reniform (3), ob- ovate ral outline of whole leaves regardless of lobing Peltate (1), ja unii 0). Present (1), absent (0). Basal glands idered p if the entire Ad margin is beset with stipitate dr ed (1), entire (0). dee glandular у eglandular (0). Margins may be toothed ог entire and may or may not have glan set ie not parted (0). Тый» may be shallowly lobed but not necessarily Unie о), 3- (3); 5- (5); 7- (7); 9-lobed (9). Infi t f lobed in plants with typically unlobed 1 been i d (0). 3 (3), 5 (5), 7 (7), 9 (11). Absent or fugacious (0), folia r (1), glandular-ciliate (2), filiform (3). Absent (0), unicellular (1), uniseriate-multicellular (2). Present (1), a t Campanulate (0), rotate e (1), tubular or meager te (2). White (1), other (0). This refers to absolute white flowers as opposed to yellow- ish or greeni $E (2). Distinct (0), imbricate es connate (2. Fett there is no correspondence mine onditi ion of ا ا‎ si The corolla y be distinct in t My male but connate in the male. Monadeiphous (D dbo 0 ). 8 (1), 1 bers have been ignored. Free eis connate (0). Stamens have been dual connate even if only the al group is connate but the outer is free. Fre ree T connate (U). Foliaceous (1), not foliaceous (0). This refers only to the unusually large, leaf- like 2 als. Glandular (1), entire (0). Mais dehiscent dm жу dehiscent (0). Рай а explode upon maturity or remain M 1 (1), 2 a n кал жүнүн (0). а "— area in Jalisco, Mexico), and : purpurea B. CLADISTIC ANALYSIS nic in western Mexico), are also discüese d below m her than these few. iii othe MATERIALS AND METHODS n & basic delineation of Dehga ише ric classification remains intact, bor thg set of ta Webster’s (1979) Cladistic analyses were баве оп the да Us) and 31 char- rtment necessit N < =; Е е 5 "m E = < e Depa yt E w species as J. martiusii will shes be further dnd: of Ecology and Evolution, State University ofr dated. ork, Stony Brook, 1988) version 1.5. 353 the phenetic analysis. 3 Dehgan & Schutzman Characters Neotropical Jatropha р Tat Matrix of character states for 77 Volume 81, Number 2 Taxon 1994 TABLE 2. “OC O9OSCOCOCSCH 9000 OC m m et wt mt mt mt O mt © met et met tet mt mt et et ed ed et et dt Cm COs es OR ORO €0 со со сї c4 сб 05 сб СЧ с© AM AM HMMM =ч 0$ сб сб со C су со со со сб có сб со еб сб сә сб со су 06 со со оу су ез оу су со со сї со са mnonOoOmmmOmMOOOOOnmnnOmnnnOonOnnmnmnmnOmnmnmnnnmnnmnmOono= mono — o Ococoococococc-cocoooocoooc-c-cocooco-oooooc-.-.-c-cooococ-o-oo-—-occ оно ноо но нон ноооооооооорононлооооооооооооооо-ооо-ооо о Ооооо ноор ноооооноооонноонооннлонлрооонна m O mim On Ome BOO Oocoococoococ-ooooooooooo-ooooooooocoo-o-oooooococooococ-.o--o-oco FASHOSOAHASOSOSOSSSHSSCOCHOH OHO HOCH OH HH OH ORO HR HOH OH ROH OH OM Но-о Z00007 00000m ooo OH On OOOO Nm OOH OOH OH OO OOOH OOH OM MOM HOH Occo gOaOOnwdcoooadocooacocoooduooooonunuduooonuo-o-ooooooonununuonuoonuonu © onOWwwaoaocododoodnuacdocoooooonudoouaouduocaooooooo-ooooocoonununouonuoonuo«nu он Јон nr ooomn nt OOH OoOnOooCCOCOOF OOOOH OH OOOO COO oO OH OOOO SS о ~ — © сч сч сз ч сч сї сз © сї сї сї © Сї з NONOOCOONON NONON OH HR OOOOH HH ONO NOMAN с OmO000000mOmMOmOmOmOnmnmmO©OOOOmOmCOOmOOmnmm—m—¬©o0O0000=moo00000° OmOmMmO0O0OOMNOmMOOONONONMNmMOOMNONOOmMOONNNRNOOOmO=nmOoOo=noo m= MMODONMNDSONNDOSTCONSOMONNNDDSCONSONOCNOANMANNNROMONONOMAMAM t OQ io мэ мэ ©\ ©\ С- iD Г- Сс C 00 OQ со C- C- Ст 09 C- C CC МЭ C CC |) C C C C CE O C p CD С n D a D D- oOooOoocooOooOooOocoococoocococoooooocooooooocooooooooooooooooooóoccoc-occoo ille ien n Ln M LL ME Lm LE OSC OC HF HOR BF HB HR RR eR eR eet Ht OOH a te eee eee © аы OcoOoocooo0o0o0o0-ododuonuudu-ooooooooooooooo-.-.oooooooooo-uooo V) мз € C Q t- © о сб с- © ми» 65 мз с© из © © © © из с© e © © © юс с© миз о © сос © © сб сб мз c- мэ мэ c мэ мэз ©\ © о о со OoOooOoocoo-.oo0oc-cooooco-ooooo-ooooooooooooooooo--o-ococo-oooo —"-oOooOooOoooOooOcc-no-cuoooo-ooco---oocoo-coc-cooo-o-c---ooooooo-oooo- Ono ocooOooOooOocdnooococooooooooooooooo-oo-ooooooooo--ooooooooz- —"oOooOooOoo0oo0o-nno-n.-ocoocoooco-cooococ-c--ocooo-oco-ooco-co-c---ococooooo-oooo- ч ооосоооооооооооооооосооооооооонооноооооооооооооооо о MMM CX с© сб Сї сб сб сї сб Ч сї сб С сб N ANNAN N ANM Сї с© 605 сї 06 сї со сб 605 сї СЧ СЧ OI сә с© 0 06 605 СЧ 05 06 0 сб сч СЧ сә со i E e E o M E E LLL NN P P P P M n M M O а-нан n mM © =ч =ч Om m ^ч OOnouwooDrmooomuuom1coooocoooooocoocooooodgoccsaGDuo: „кк aa OaAaAtOnmooonoCcoonOoOmroooecoooeoeooeoooooHOoOmooocoHe Ss CLO Oo ONO OOOO OOo OOK OC ooo ocooceoomocoocoe oO OOO OOo OC SCO OS eB Oo eee oe So OoOooOcooOooc-Qooocoocoocoonooo-oooooooo-oooooo-.-ooo-oooooooooo mG mH NNOm mm OO mg ONAN ON HN mm HN O O mm e S S © g E sS ЕЁ z 2 نه‎ B 4 = = = E = 3 i z = 5 S s ue S > > з 2 РА v X S38 in .8 2 بډ‎ 8 : D e NE M Wee з 8 8 аө п 2 гп es SESS GSES. 3E е 525 Жыз: Ж ups MEET S 33 = VE: sp SP oss = BRC - Е ais US ula nS E ош CIC UC QN IUUD ч с сысы e *$ 8SSUARSSSPESZILSEBLSSSSISÉSSS3GSS3SS 2555.9 5 з Ss еса ооо юе E ы ЧЫ 2-1 o RS Em > SS 3SS E - ЕЗ т) S 6 2 Фф E] ` ° 2 OO: B X BNE S252 5 5 Hrs = с ЧЕ ЖЖ к-К: И Е 5 5,5 9 9 Frese с са Е бО Б-У Т ЕЗ = SSS Sis IIR TORR VISITS OSS AS SSES Sssssesssssgge65s%3 3 8 = DL SS З = Ss کے‎ © = O ы Less 3 & = SSSSSSSCSCSTCSS ss Sse PTT DUO RE g-c-cc2555335EEEEBEBEEESESSB'S 354 Annals of the Missouri Botanical Garden Continued. TABLE 2. Characters Taxon 0-0:0-T 1.3 2:0.1.0,1:0.0 1,105 0.1-.0.2.0.2-2:0 00 ОБОИ ortegae pachypoda 0.00 1.0:0.3:1.0:0:0.0.0.0:1.00.0.0:0:2- 1.2. 2:0. 0:071 2100 АИ platyphylla purpurea rzedowskii standleyi FO O OL I T0 0:0 0.0.0:0:105 031 12.1 OOOO 000111100000011 07000212200 0:0:0:0/0 2D 1:0 0.0:1-1-3.0.1:0:1.0.0-1 1:05. 08), 012: 152 2.0,0:0.0:0] Os lalcozotitlanensis LOO. OO LT 22000:00.50.1:0700010001 11100 POL € lus 000000300000001090010102100000030 yucatanensis O subg. Jatropha * subg. Curcas ic Jatropha species from 32 characters, using F. J. Rohlfs NTSYS-pe program. Infragener (d PCA of FIGURE 1. designations are from Dehgan & Webster (1979). дап & Schutzman 355 Volume 81, Number 2 Dehgan 1994 Neotropical Jatropha TABLE 3A. Qal чн Иа Щщ +, f. 1 1 g f 4 z g 1 ) I I p in analysis of 77 species of Jatropha. PGI PC2 PGS Character Factor Character Factor Character Factor number loading number loading number loading 4 A8 16 =O:327 19 0.384 5 —0.263 6 0.317 26 0:851 32 0.259 1 0.307 18 0.327 23 —0.256 I5 —0.304 9 0.283 25 0.259 14 0.287 20 —0.257 30 0.252 13 — 0.204 11 0.274 * See Table 1. acters are listed in Table 4, and the data matrix in кын hl e (Muell. Arg.) Pax (Webster, is presented in Table 5. Earlier runs made using PAUP Version 2.4.1 (David Swofford, "Illinois Bs Hanc es to be closely related to Jatropha SV s. rior Lear of sister group (McVaugh, 1944; Mi tan & Webster, 1962 аха or on cs er c }; Three sets of analyses were TT the firs Char ve polarities were not explicitly desig- nly Aleur. G. Forster as an кын, ated (nonadditive option of Hennig86), but the the second usin nesia Vell d the selection of Aleurites and Joannesia as outgroups third wit th genera selected as outgroups. Se- ctively polarized the characters (Maddison et lection of these outgroups was based on (1) the he 198 ithin presumed close affinity of Joannesia with Jatropha group selection, two algorithms were used КУ л м. * sect. Jatropha a sect. Peltatae a sect. Polymorphae PC3 10.2% О И t. Polymorphae) from 32 characters, Ficure 2, p : CA of species in Jatropha subg. Jatropha (including species of sec "sing Е. J. Вође NTSYS.pe -pc я гане; designations are from Dehgan & Webster (1979). Annals of the Missouri Botanical Garden * le) f E 3B. in analysis of 39 sp with highest factor loadir 155 (їп or cas with four species of section dem eat PCI PC2 PC3 Character acto Character Fact Character Factor number loading number loading number loading 5 —0.337 22 —0.408 9 0.428 6 —0.333 30 0.373 11 0.415 16 0.329 4 0.328 27 0.286 31 0.296 7 0.295 30 0.273 20 —0.279 28 0.273 15 0.260 14 0.272 26 —0.203 19 —0.243 * Characters 2 and 12 were invariant (see Table 1). to pontine: ssladograms: The ins of these is men- tioned b анешрь to on parsimonious trei Ale а гер- UP and of phylogenetic analysis programs, but is nat dis- P tree and passed to the branch- breaking algorithm, “bb,” which per- parsimonious trees. This causes the branch- Led ith to start with a completely unresolved “‘bush’’ (one lytomy consisting of all the EUs А In i second algorithm, the *mhennig*" option is used to supply the initial tree to the branch- м, which then CLADOS лр was input by the baud Eo d subjected to the “m*; and ; vs uences to ascertain iri Hennig86 could find more parsimonious clado- grams by starting s kai trees from another program RESULTS AND DISCUSSION he CLADOS, PAUP, and Hennig86 data sets sis of different numbers of characters (46 binary in CLADOS, 31 Morc in Hennig86 a nd — W sect. Сигсаз . Lour PC3 10.196 nee Lr PCA of species of ager € сш айм T. |. Rohlf NTSYS-pe program. ters, s and дее 9 lee ne V from 32 charac n & Webster (1979). Volume 81, Number 2 Dehgan & Schutzman 357 1994 Neotropical Jatropha TABLE ЗС. Sel d с} * with highest facto l I I l I in analysis of 34 species of subg Jatropha with bow dh of section eae РС1 РС2 РСЗ Character Factor Character Factor Character Factor number loading number loadin, number loading 19 0.356 т 0.368 12 0.364 18 0.327 25 0.309 23 0.344 9 0.309 6 0.286 24 —0.312 п 0.305 16 0.258 22 0.284 20 0.298 23 —0.254 8 —0.275 29 0.268 17 0.240 13 0.253 * Characters 3, 4, and 28 were invariant (see Table 1). AUP), and taxa (79 in CLADOS and Hennig86, the e Oi index (Kluge & via oe subsets of 4 to 19 taxa in PAUP). Since the con- t. In order to comp: CM sistency index (CI) is highly negatively correlated RO T CLADOS a nd PAUP t to those of to the number of taxa and characters in a study, — Hennig86, we used the in Hennig86 mber S and y the CI. ud Ed VU edid: pr Г” option was then used to calculate the length of TABLE 4. C ^ 12 ЖЕ у paren lysis (31 cl bered 0-30 using th a of the Hennig86 program). à Growth habit: Vanes gentes (1), ле т (0). - Growth habit: pical (0). 2. Plant sex: Dioeci ous monoecious 0) 3. Laticifers ee lated (1), idioblastic (0). * Laticife Nonarticul eo. fake (0). C De тыы), аё авд). : E basal ‘eee Present (1), i p Ы Toothed (1), entire (0). E lat marginal glands: Ciliate-glandular (1), — (0). af sinus depth: arted (1), not parted (0) Leaf lobe numbers: Unlobed (0), 3- (3); 7 (5); 7- (7); 9-lobed (9). ^ Sedis [ш Palmate (1), pon e (0). ET Long (1), subsessile (0). * Pride е 3 (3), 5 (5), 7 0 1 (1). ^ bsent (0), unicellular (1), uniseriate-multicellular (2). 15. lors Present (1), a Flower sha Campa: pegs © guns e (1), tubular or urceolate (2). ЧАА чы color White (1), 0 Peale ч female flowers: Distinct (0), ine (1), connate (2). Eko male flowers: Distinct (0 i imbricate (1), connate (2). ч Slamen arrangement: Monadelph ris diadelphous (0). » Stamen number: 8 (1), 23, New fusion Free (1), connate (0) g on: Free (1), connate (0). 2. * 25 ín rie d а flowers: oliaceous (1), not Urna (0). m n Glandular vis entire б Frun dehiscence: Violently dehiscent (1), tardily dehiscent (0). <í. Fruit locules: 1 (1), 2 (2), y S е Carunculate (1), И (0). 30. r Valvate (1), not valva h yx lobes: Petiolar basa] glands: Present (1), absent (0) Annals of the | Missouri Botanical Garden 0,480 0,400 f т 0,320 T 0,240 T 0,160 T 0,080 T 1 a neopauciflora rzedowskii ciliata = ‘em | hint 0:480 01400 FIGURE 4. Infrag program. g 91320 01240 L 01160 01080 01000 iir c of T Чөө s siue species from 32 characters, using F. 9). n & Webster (197 livacea tlalcozotitlanensis ullockii —O -o — A Sect. Polymorphae —e pu e] Sect. Loureira Sect. Mozinna Sect. Loureira Sect. Platyphyllae kl Sect. Curcas С = = y Sect. Polymorphae lA) Sect. Peltatae 4 Sect. Polymorphae sg Sect. Peltatae Sect. Jatropha‏ چا = Sect. Platyphyllae а Subg. Curcas Subg. Jatropha Subdg. Curcas J. Rohlf's NTSYSP° 359 Dehgan & Schutzman Neotropical Jatropha Volume 81, Number 2 | 1994 р Matrix of character states for 77 Jat TABLE 5. Missing states are indicated by *'?." Characters Taxon 0.0.0.0 2.0 -0:0 04909 0000 00оо 1-0 1101.1 3104 050.041. 010201705 0:007 05151 :13050 11.110050. 0:1:0:3.0 0.0 0010100100500910000100001013100 0011000000500701212000 angustifolia cardiophylla costaricensis cuneata ^ з S з 5 3 = [ue з $ 23 EE Mu CÓ cM DEES ss 3 8 E ee M З 5 S.8 e] Ж. кє ER wu 3g ЕЕЕ» ЕЕЕЗРЕЕ БЕ Е 2883355355252 58 Ss «©©© ОФ D o ® o % andrieuxii canescens oO: Oo о ою ою Фо о 6 oO осо о ecooococecooocooocooooo eooooooooocooooooecooecoecocoocooocooooecoo ooonrroooconror oocecoocoecoocoocooor on OOoe Oo с© со с© =з с© (6 с© с© СЧ с© сс) с© со MO MMMM 00 00 06 сс) 05 з су 05 со 66 05 AN ooonr ooooror ooomonooocoocoocoocoorn Oe OOH Oo ФН es OCC HF ORF HOH SOCOM BF BB OCC OR HF et On OHH Om eooooooornror ooor ococececoeooooeonooor оо о mn OO CCF RF OCH COM BF OR OC OOCO Om mn Ht Om et On One n ea CO oormrocoooocoooooroor onmoooeoeooocoooooor One е aon Oonr OF OCC FOF HR HOH OCH ORF HOF OR FOR OF OR — е na Om OoRmF CoCo eCcoF OCH OCH OR OCC OCO OF OOH OR FT OR — е SO ONSCSTSOSOSONNOSCOCOCH ORF OSC OCOCSCOOCOONNNONOCOON SEAPSNNSTONONNCCONOCOOC ORO OSC OOO ONNNONOON BOR OF TAF FAR OF HR RB ROR mmm ess BROTH OH OHA ~¬_ ANONTSTONONNCH OCH OF HB OCC Omn HH ONC HOR AN SCnm OF OCOFP FRA HOF ONT Omn OOO Omme ee HO n ea нн н NONONANROTOANONHFOHOONNNNOCCORF ORB COON о C= De OO ee De Се Му ee OO) C Бе Bee се (Gb 9b ON eS жае М» ©с©—=—-=—=—=—=—-©5©©—=—©5©©©5©50595—2— 5555—5555 SCmrnmrNwrcoooocoocoooooooe oon E oooocoecoocoessN N — =ч =ч om) CÓ) OC) ч =ч 0 D A 00. О Ш 00 =ч 00 00 з MMO Мз МЗ Г ш о С =ч = mscoooornmooooooomooooeooooor wn On ooo oO axooeonmnr sn Oooo on oocoHnOnnn tn ooocoocoorn OCC е eccooocoocecoeoon Oo Orme ooocoeqcooornaooococe 6 Б чооон TF OCOCOCH OF OCOCOCH Omni exam coocooocooecon ooo Ooococoocooocooococoooooooomnoooooooocooooococoo oocoooqoocoono Coco conor Ono ocooooFon oe © © о ooonoo ces on oooneooooooocomCccSmoC Soo oeooco Ообо м о ооб a Ono OOOO CO 8 OOo OO oO oon oo 5 6 ¬ SSO S.9:'0' 0 SO SO 0:9: А 0:0 OS OOOO оссо еә sacoomocoooqoooqcooonoocoeooconmnmo Comoceooosx ° S S S E = Г {г IS M m v S X © ESE E: ESR шр gaa EC P BCP $3 FSS ESF auis т = SSSHEEDPSSSSSPSESSESSE ка В ассос сссоа-о9 р ss Ps ә А SSIs ес о Ә-ә е әб Ss узго eee ES SS SH ЕЕЕ ЕБЕ Е Е Е Е Е Е 360 nnals of the Missouri Botanical Garden TABLE 5. Continued. Characters Taxon nudicaulis olivacea coooooococoococoooooooor oer ooo Ooo OS wai — OO OO OS کا ا ی‎ oo — ж ا ~ ا‎ ==, wo ucatanensis 1 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 е trees relative to the Hennig86 dat CLA et. The radiis DOS a 0 da i steps. The es ES a ure 5 was selected by calculating character w weights (**xsteps w" command) an o all 100 I f th grams considered. The len of the cladogram were recalculated using weighted characters ("xsteps I” comma The с опе is illus- trated in Fi trees i a using dat , but gives more cre eden nce to Сеге show did without resorting to proximations approach, such as that suggested by Carpenter (1988). Even prior to Archie’ par criticism of the consistency index, Farris ad s ady abandoned it in his 1988 release of H ices: (RI) and escaled Consistency 1 These are constructed to behave a d t ve indi ota: values between О and 1 (Farris, 1989) The most parsimonious cladograms from Henni all - and RI = Stated simply. RC displays a o egree = homoplasy in the буле set, while the high value for RI shows that es cladograms exhibit a ез гае low degree pre’ экелу as compared to the total amount poss or the data set. (Fig. The strict (Nelson) consensus pem o 6 is re similar 10 FiGURE 5. ыы: At Nel (st eed the one selected 5) output by Hennig86 is remark ably ing less hgan & Webster's (1979) evolutionary ya à successive ap- which utilized morphological similarities а KRAUS S o — ites) tree for x и: species and two outgroups (Joannesia and oss 96 vi © program. f. Table 4), using Farris’s Volume 81, Number 2 1994 Deh Neo gan & Schutzman tropical Jatropha 361 Annals of the Missouri Botanical Garden ferences in concert with the principles of evolu- (1 can (1987), and others. Although m Чы of ш sections and 0 ections are u of taxa within groups is advisable, the fundamental infrageneric relationships remain bue intact. BERE Jatropha as a whol м h + к рһу г 69-4 h | fs Sno a WM x ] The strict (Nelson) consensus qnis (Fi ig. 5) Чол lees indicates the relationship between the two s gen m on SM early шупш апа а 1 COrr еѕропаѕ о the previously suggested ку ia wou of sections Platyphyl f the th recognition he group at seal leve |, mor batable. jue. ous to Mozinna, section Lourei- ra includes species with bi- and aedes ied à subsection Canescentes hgan & ile J. gi Webster is mostly triloeular AQUAM higher locule number in some y be the result of reversal (as n J. перане Vd. in nearly all other cases it is ance suggested ee in UN Jatropha эрк transfer as initially n & Web- ster ) to section Polymorphae. This species diff om most m rs of Polymorphae by glandular leaf margins and possession of foliar se- he female flowers. Jatropha macrorhiza » | ict | А ri M } S Desert), which was recognized as distinct subsection M acrorhizae in Polymorphae, would be trans- erred to section Peltatae, together with J. ca- thartica Terán & Berland, J. m rocarpa Griseb., multifida L., wit tifid ERE all of which would be a in pee hs ifidae Pax. Jatr s ran- tha Muell is accurately placed in section Р, o a ade conta m e a ral autapomorphies and also is phenetically distinct. It is the only non audicifo ican species with eight stamens, € fruit, both sections, but in fact does not fi either one. It is abe only Know в gnam m к» locular aisuncuy а се but only two (as ае. to ve) pe branches. No other species in the genus is known to consistently possess a combination of p ts. a purpu ii ate between sections Рр ап‹ ке ec . sect. Loureira), but has red trilocular fruit, and carunculate seeds s those of section Jatropha sections should be er to ассо rtiusii and J. рш" purea. Accordin eun ection Jatropha would in- clude three subsect комур peg taxa, J. purpur and Hee related жеты ты" Sout ‚ы С. COMPARISONS BETWEEN AND ies OF HENETIC AND CLADISTIC ANAL There exists a general concordance еси о ed Ьу phenetic om cladisti ses. Where papa quc exist, a attributed to the inability о ted ps expected eneral agreement ANA. the two m шрген Volume 81, Number 2 1994 n & Schutzman 363 жыз aoe Jatropha with regard to establishment of subgenera, sections, and most subsections. Mo Dehgan & Webster’s (1979) intuitive classification confirms that intimate knowledge of living (as op- dt often essential for in cultivation. The rs of evolutionary changes that have resulted іп p morphological disconti- in the traditional classifica- ost tes mms y the nu- merical analyses Resa) her large numbe tion, an D. RELATIONSHIPS BETWEEN PHYTOGEOGRAPHY, EVOLUTION, AND THE xa NUMERICAL ANAL е fully cognizant of the shortcomings of e f Pcia у and р оар. е ney have a bette the Am n taxa. To under- ee 0 entire genus it is ini that rican and b. species also be included in the in the recent ко о he tatis in (9 yd al. нар ao a subsections dy to the pinosae Pax сос of this «о cime ical dye precludes gens d World species at this time. However, th "eed large ae of neotropical taxa in Jatropha, iin iei) in concert v mim t majority of the taxa, known um : tural к ы and probability of wie 100, all profoundly infl e the results of " ug studies and will be di M a P^ of the genus e it to say that the a merica eed is als a consen nae Vie ee. Several other evo- that were put forth in е wi pH e ur: вап, 1980, 1982, 1984; Dehga '& 1978; Dehgan & Webster, 1978, 1979) have been confirmed. Of particular interest is the fee se progression of infrageneric groups. As evidenced by the ee ee species bP! eness of of section С g rimitiv the section, and of J. c rticular, was noted by McVaugh (1945) sem Wilbur (1954), who maintained that the genus must h ilar to J. c similar ipiis follow progressive mos lution and mosaic шун (Stebbins, 1984) logical pw disi disturbances in Mexico (Toledo, 81; Graham, 1987, 1989b) have probably re- неа in both eliminatio. many taxa related to ho outh rica and seabed opon saic n gene tropics has been m ry regions: єз ш -savanna (соп ado), thorn — b, I n South Americ scru d of sections nee and Peltatae are а Brazil, Раг п drier areas ce from the wetter regions of South America is understandable. Webster (1994) has маку es the Euphorbiaceae ا‎ obably with Afri greatest ne of the prim subfam es. Dehgan (1980 et seq nd Dehgan & 78) enumerated the prim ed its аде ancestral po- uphorbiaceae. This view is further orted T Gilbert а others m Tw issue > ie orig drier regions t Gondwanal land and never E oped the ability to adapt to wet tropic al forests (cf. Retallack & Dilcher, 1981). Annals of the Missouri Botanical Garden Despite диир e distribution of emo and ofte en vast bandes ity o uit spec endem nn which is rep — " Pape: (taney. 1923), no en cs E t dieti enus appears "en т ге нови} of progressive overland dispersal. With the © is 2 i=} 2. : 22 67 n Е > e 9 a Е; Б @ 5 E © ©, — z absence from all oceanie alanis сер! de чиш Г, icata and indication of their inability to disperse by birds The larg emic species i bitat dis- turbance, as has reportedly occurred in Centr: and ss America (Graham, 19 and hence 89a), the npe ben theory (Prance, 1982). uphor- such as Aleurites in Asia, Hevea Aubl. аааз Cnidoscolus, Manihot Miller, and others іп Amer- ica, and G 5 a good and modifications have occurred in conjunction inb migration from south to north in Ma esoamerica a north to scuto in | South Аша migration and ensuing geographical wes was eriod of probably followed by a shorter or lon 2. =” pe о2 оня заса rhizomatous subshrubs short shoots and small to very small leaves, lateral few (8) or solitary т inflorescences, female flowers with a single bifid а, unilocular fruit, and polyploid сы us, species of atropha have endured gradu al mosaic evolution- ary changes, as re vit in their great diversity of adaptive types, which a pie correlated with their geographical distributi zinna originate section (со ога А еа simi milar f garar es a 0 e, lead е. way to evou of taxa in section T shown to have evolved from oon rise ing наор Varo opia is and deld: Grant (1963: 566- 568) has suggested involve that speciation may be inext s cessive spe- gradual s PERENA sien M ds in hen is атаа tuber us-cau- print inflorescences (terminal compo mud di- chasium with a distinct gn to lateral or terminal solitary flowers mber of stamens (10 to 8) and their arrangement vies or — mon- fre d 1) which, with the notable J. martiusii (with 2 styles bu sexuality (from monoe ynodioecy to Progressive diversifying M as a result ti puedo taxa de- ubli- elta seribed after Dehgan & Webster’ s (1979) p " en әрни иы " ас In short, the P = arlier in the phylogenetic scheme. lassification recognized grade taxa ba Е iam plesiomorphic features. Only som к adjus : ts to previously recognized T oup LITERATURE CITED ADANSON, М. 1763. 346-358 XLV. Е milles les and Ti yd li. ARCHIE, J. 9 omoplasy excess Г da indices i. окиб nets of bons m ар ve Volume 81, Number 2 1994 Dehgan & Schutzman Neotropical Jatropha index. E Zool. 38: : ASHLOCK, P. D. IOV: уь ы associated terms. Syst. Zool. 20: 63- Влшом, Н. 1858. Etude générale du groupe des Eu- E pp. 294-296. CARPENTER, J. M. 1988. Choosing among multiple ama parsimonious cladograms. Cladistics 4: 291- 1987. A botanical critique of cladism. 23: l- Dexcan, B. 1980. Application of epidermal morphology to cri delimitations in the genus Jatropha L. (Euphorbiaceae). J. Linn. Soc "Bon 80: 2 257.2 78. PE 1982. Comparative moaned of the petiole nd infi p opha (Euphor- basa imer. i 69: 1283-1208. Ee hylo dari significa p hitb in sj ннд ts aa gis 7-4 ——& CRONQUIST, A. Bot. Rev ME E. CRAIG. 1978. Types of laticifers and M in n Jatropha m their taxonomic implica tion r. J. Bot. 65: 345-352. = fe - WEBSTER. 1978. Three new species fi Mexico. Madroño 25: 30- of Jatroph 39 & —— ——. 1979. Mor phology and infrage- oe ic relationships of the genus Jatropha (Eu Hg Efi Univ. Calif. Publ. Bot. 74: 1-73 P 1970. Genetics of the Evolutionary TOCOess, Columbia Univ. Press, New York. The retention moe and the rescaled 417-419. ropical American bac floras Я paleoenvironmens: M ая ісо, Соз ки апа Panama, Am 1519-153 ЖОЛ а. ны Terry paleo and veg- eru aang in ve "i Central America. Е ag Е Paras ix ратай сһапдез in the Te E of northern Latin America. Rev. Pa- eobot. Palynol. 6 60- 283-293. - 1963. de A of Adaptations. Columbia wd ^^ 19 e Soma ей species M Jatropha бы Vision o ш, Бош) Kew Bull. 4 ud А. С. & J. S. ceci xs MURUS phy- es de the evolution P anurans. Syst. Zool. 18: ТЕ D , 1763. Pp. 1 - 1 i : ‹ tin: ois р p- 1428-1430 in Species Plan ADDI 1 à W. E & R. MADDISON. M. J. DoNoc x Ong analysis жү Dia Syst. Zool. 33: 8 McVat t t: 944. The genus Cnidoscolus: Generic т. Fer oe groups. Bull. Torrey Bot. Club cipal i Ee: The е genus Jatropha in America: Prin- 251-9 itergeneri groups. Bull. Torrey Bot. Club 72: iî ¢ A. & T. Duncan. 1987. The necessity of convex groups in biological classificaiton. Syst. Bot. 12: 78-90. MILLER, = I. E. C. E WEBSTER. 1962. Systematic d Jatropha. Brittonia 14: 174-180. UELLER ARGOVIENSIS, J. 1874. Jatropha. In: Martius ~~ 2 Flo: r gg 486-502 D: i EE in Das Heft 42) Tar von Wilhelm d pese Leipzig. PLATNICK, N. 1989. An empirical comparison of a parsimony programs, II. Cladistics 5: 5-161. Рон, J. E. 1827. аат тее Icones et Des- criptiones roe PRANCE, 982. i. view 4 е phytogeographic evidences a pleistocene climate changes in the Neo- tropics. Ann MM d Gard. 69: 594-624. P SMITH. Euphorbiaceae (part 1). In: us uh hil йш, Fiora of East Africa. ee half o ema, гене Bosto - L. ШНЕК. нав. А costal hy- ce and rise to dominance of flowering plants. /n: K. J. Niklas (editor) Paleobot- any, Paleoecology and Evolution, Vol. 2. Praeger Publishers, M York. SNEATH, LH .&R. dd Бк. 1973. Numerical Tax V. H. Fre de Francisco. Stout TP. "1023. Trees Eod shrubs of Mexico. Con- trib. U.S. Nat'l. Herb. 3: 634-642 STEBBINS, 974. Flowering Plants: Evolution Above he Species Level. The Belknap Press, Cambridge Mosaic evolution, mosaic selection д angiosperm ‘phylogeny. Bot. J. Linn. Soc. 88: 149- TorEpo, V. M. 1 add Pleistocene changes of vegetation in tropical Mexico. /n: G. T. Prance (editor), imd ical Diversification in the Tropics. Colombia Press, e ork. WATROUS, L. > О. D. WHEELER. 1981. The out- group compar method of character eiu Syst. ool. 3 WEBSTER, = 975. se ag of a new classifi- cation р i ЭМА эрка Тахоп 24: 593-601. A new species of нер е irl hice) ков e Ann. Missouri Bot. Gard. e 1087. The of the spurges: A review of classification and relationships i in the Euphorbiales. J. Linn. Soc. Bot. 94: 3-46. ————. 1994. Classification of the Euphorbiaceae. Ann. 2 sud . 8l: 2 32. L. J. POVEDA. 1978 phytogeographically pa * urophe ^ cp E fro Was. R. < gs «ы һа, subsec- tion Eucurcas, with the Чер of two new ao from Mexico. J. Elisha Mitchell Sci. Soc. 70: 92- 101. 366 Annals of the Missouri Botanical Garden APPENDIX. Sp р f the pha sp d Species and authority Herbarium Collector and number Country J. alamanii Muell. Arg. MICH ing 1 Mexico J. andrieuxii Muell. Arg AS Salas 1175 Mexico J. angustifolia Griseb. NY Jennings 147 u J. augustii Pax & Hoffm AL Belshaw 3056 Peru J. bartlettii Wilbur MICH* L. & C. R. Wilbur Mexico 1472 J. bullockii E. i MICH McVaugh 26284 Mexico J. h.) М KEW* Hinds 1841 Mexico ТА ету (Torr. ) m ju DAV Dehgan & Webster US — Arizona B7 3 J. cathartica Terán & Berland BM Fryxell 2943 US — Texas J. chamelensis L. A. Pérez-Jiménez ENCB, MICH Rzedowski & McVaugh Mexico 1400 J. ciliata Sessé CAS, ENCB, F, GH, Pringle 6348 Mexico M, MIN, MO, NY, UC, US, BM J. cinerea (Ор). oe Arg. RSA, GH, NY Gentry 7050 Mexico A HE Mue GM, GH**, NY** Mandon s Bolivia J. cordata (Or kie ub MICH McVaugh 2210 Mexico J. costaricensis Webster & Pools DAV* G. L. Webster ^ L. J. Pove- Costa Rica da 22160 J. cuneata Wiggins & Rollins DAV, MICH, NY Webster & Lynch 17008 Mexico J. curcas L. FLAS Dehgan B86.050 exico J. dioica Sessé hgan a ех! J. dissecta (Chod. & Hassl.) Pax GH*, KEW** Hassler 4 araguay J. divaricata Swartz BM Webster 3 Wien 5073 Jamaica J. eglandulosa Pax BM* Hassler 8 araguay J. elbae J. Jiménez Ramírez FLAS Soto ea. Mexico J. elliptica (Pohl) Muell. Ar BM, KEW, MO Steinbach 6708 Brazi J. excisa Griseb. KEW, BM enturi 7991 Argentina J. flavovirens Pax & Hoffm. KEW*, BM** Hassler 2489 Paraguay J. fr ioides Standley л Fisher 8 Mexico J. galvanii J. Jiménez Ramirez ontreras & Jimén exico ; pre Greenman US, TEX, MICH Lundell & Lundell 7502 Mexico J. giffordiana Dehgan & Webster * Dehgan B74.019 exico J. gossypiifolia L. FLAS hgan & Webster Brazi BD86.30 А состои ves and Hoffm. LPB ck 6442 v" J^ nitica UC* Hassler 10104 Paraguay A coge de [CR KEW Fuertes 359 Dominican Republic J. hieronymii O. К LPB Beck 4970 via J. hintonii Wilbu BM* Hinton 4300 Mexico J. humboldtiana ШО єн d berbauer 6211 Peru J. integerrima MO, NY, F, GH mbs 99 Cuba J. intercedens Pax ICH*, BM**, B** vin 5393, Fiebrig Bolivia J. intermedia (Chodat & Hassler) MICH* Hassler 3795 Fere Pa J. isabellii Muell. Arg. BM* Hassler 4338, 5930 Paraguay J. katharinae Pax BM** ssler 9078 Paraguay J. macrantha Muell. d KEW Sauderman 3983 Pen J. macrocarpa Gri BM, KEW Venturi 5472 res J. macrorhiza Mut. : AV Dehgan B74.076 USA J. malacophylla Standl US* Gentry 1449 Mexico Volume 81, Number 2 1994 Dehgan & Schutzman Neotropical Jatropha 367 APPENDIX. Continued. Species and authority Herbarium Collector and number Country J. martiusii (Pohl) Baillon FLAS, DAV Dehgan & Webster Brazil J. mevaughii Dehgan & Webster DAV ae B74 Mexico ve E. (Pohl) Baillon FLAS Dehgan i Web Brazil BD86.3 J. moranii Dehgan & Webster DAV* Howell о ехісо J. multifida L. MO, KEW Valeur 684 Dominican Republic (Pohl) Baill FLAS n & Webster 312 razil : neopa ша Pax TEX, NY, GH, DAV orae et al. 17319 xico J. nudicaulis Benth MO Dodson & Dodson 12941 Colombia J. olivacea Muell. А g S Pringle 6348 Mexico J. ortegae Standley MIN* tega 6455 Mexico J. pachypoda Pax M* Fiebrig 3042 Bolivia J. pauciflora Gri NY Ekman Cuba J. peiranoi Lou MO O'Donell & Meyer 5308 Argentina J. platyphylla Muell. Arg DAV bster & Lynch 17032 exic J. podagrica MICH Breedlove 11810 Mexico J. pohliana Muell M isa ule 26802 Brazil J. pseudocurcas E RR DAV Webster et al. 11746 Mexico J. pu rpurea Rose BM* Palmer 7 Mexico i ant ы) Baillon KEW* Gardner 2303 razil J. ricinifolia UC Venturi 2350 Argentina Hales s. n F** Miranda 2111 Mexico J. rufescens Brand egee UC* pus 4 Mexico J. rzedowskii f. nei Ramírez PEAS** énez s. Mexico J standleyi Steyerm: MICH Matuda ad Mexico J. stevensii G. L. Web MO Stevens Nicaragua J. sympetala Blake & iue BM, ENCB, GH, wies «pes xico MICH, TEX, UC, NY J. thyrsantha Pax & Hoff ICH T 455 Bolivia X Een $. i ad Ramirez FLAS Lozano 38 Mexico J. tupifolia E AV, GH Wasa A 2 198 Cuba J. vernicosa vate gee UC**, GH Brande: ээн Мехісо J. weddelliana аа CEN Allem 6 Brazil J. yucatanensis Briq CAS Thorn ч dde 40462 Mexico Type. cu ои. A PRELIMINARY CLASSIFICATION OF UPHORBIA SUBGENUS EUPHORBIA Susan Carter! ABSTRACT ring preparation of the Euphorbieae for um Flora of Tropical East diee most of the 250 plus species = cla: Dur e subg. Euphorbia sification. Comments offer n the principal systems to date, all of which have ped based on 1 vegetative I y prop T "ES certain character changes linked from trees, to shrubs, and then herbs, as an adva of the inflorescence, capsules, and seeds a are VON SE ancement in adaptation toward survival in arid conditions. Features be the groups, whil le based on seed «ol is proposed, and the ЕА between eight proposed subsections are discu most impor hin ortant in deciding тар between ment within the рг A of tw oups. A system sections 0 ае e the € recognized specs: Within the sub- t e maj ority chasial cyme, or of a group of cymes. Some Mad- are actually modified stipules, each with an ex pani ey base that flanks the of which are native to Africa (excluding Madagas- car), the to the Canary Bree s Euphorbia in its strictest sense (for- merly section Diacanthium of Boissier, which in- es the type ron si the gren я anm orum within the genus Euphorbia. Its species КЕШ е stems and br иш: with the base of н dest surrounded by a y pad (the spine- zs ر‎ еа and fina modified as Bis The pa ed spines may become fused to produce a bua е spine; and the spun may occasionally be as large as | the spines t apparently obsolete. The extent to which the piik- les are obvious has affected almost all i tions. Above the spine-shield, morphologically the leaf-axil, is the flowering-eye, from vie ie inflorescence develops, consisting of a solitary di- o, e d fu n d [«] - @ n > Ф 8 = ular s ан are often deve ә йер. {тїп, these m belong t Biase E f.) M. G. Gilber rt ^ 987) and are confined to Madagascar, with the exception of i pi ica. dens E. milii Des Mo plenden (Hook.) Ursch & Leandri), now included in m genus Lacanthis (Raf.) M. G. Gilbert. Boissier m: fined two groups within his section Mes wo large prickles and a Рн (fused) spine. He Saat divided Biaculeatae species with cylindrical 1 as opposed to ranches, and z latter into those with сөрү (two-angled) branches versus those with eens ' Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, U.K. Ann. Missourt Bor. Garp. 81: 368-379. 1994. Volume 81, Number 2 1994 Carter 369 Euphorbia subg. Euphorbia more angled branches. Most AE fell within this last group. He thought that Eup a venenifica Kotschy, a eii Spina shrub spiraly; ar- ranged tuberc b I the gen nus; ae * mistook E. ingens Boiss., a large tree species, for one without spines. 862. Mn BY BOISSIER OF GROUPS IN SECTION [s '"ANTHIU. Diacant on. Nd Miis ЫШ Branches sharply sr or ridged чае ез ез = an nada angled ейде ү (fused) бча, 2 prickles Since Boissier’s (1862) treatment, attempts to f classify an increasing num species have re- d nch-angles, including trees, shrubs, and Wf Té; often with ido further delimitatio; In 190 04, Pa ax published a classification for over 80 species in section Diacan thium, based primarily ye. By far the n ^ which he inc cluded i the unrelated species E) E: was the Diaca m CLASSIFICATION BY PAX OF GROUPS IN SECTION 1 (fused) spine leatae Boiss.) 2 spines ous roots or E branches not winged es present Leaves absent Shrubs or trees um es pr Mon Pacem i теа Herbs у tube: Leave: nches Bas -many-angled iiio winged ез absent; branches 2-8- -winged Tr 2 (Triaculeatae Boiss. 1 (fused) spine, 2 Taracantha e spine з, 2 prickles ermedi ediae 2 е кону) prickles, upper stip- т ines etimes s pre А with no third ap using number of branch-angles Berger to no теп of s "d was erected in 1907 by быы, mmodate ї 50 species in cul- Кыша $ at » La Moro a mm кү on the Italian Ber, у; by Sir Thom , wl e Ber T curator. His reo pres formal * only a vague resemblance to those ла pn Pax. Bened included Splendentes (the unrelat branches T ч Es number of species were in this last group. же spined species were not included, as none were i cultivation at La Mor 1907. CLASSIFICATION BY BERGER OF GROUPS IN SECTION DIACANTHIUM Splendentes Branches terete; leaves present, small 1 Branches terete; leaves present, large Scolopendriae Herbs with short thick stem; branches ribbed; leaves absent Compressae Trees; branches 2(-3)winged; leaves ied or n branches 3(-4)-winged or ribbed; leaves abse Polygonae Tee, ra or herbs; branches 4-13- ribbed; leaves absent 1912, N. Euphorbia for bo Flora of Tropical published, in which he fav within the ui. The arrangement for almost 80 spiny species E those „т spines first, followed by those with paired s. The latter were firth loosely arranged by hai dwarf perennial herbs, shrubs, and finall es. In 1915 he followed a similar pattern for 21 species from the regions covered by Flora Ca- nsi E. Brown's و‎ t of Africa ored no e sections n his account zi 1911-1912 AND 1915. TREATMENT BY N. E. BROWN OF SPINY € IN FLORA OF TROPICAL AFRICA AND Pick CAPEN: Flora К deir Africa Spi "si ү e ee leaves large, persistent т shrubs; leaves very small, deciduous dann d ants < 1 foot high; flowering branches < 1 inch thic Plants 7 1 foot high rubs; Doaa branches < 1 inch thick Trees; flowering branches > 1 inch thick Flora Capensis Herbs, <1 foot high HE Shru Trees, 10- 30 fee sented a more formal —€— in pecies own at that lo canthium, he first lis es section, Diacanthae, under countries of origin Annals of the Missouri Botanical Garden (including E. splendens from Majaguensha with Triacanthae and RES os dien absorbing Monacanthae into Triacanthae ‚Теш н contained species dec dt ac u- ation or "o tion of hes pine- shields pee сас of the plan f. | AE 0 D; ed Berger’s Scolopendriae : as a subsection to house them. Th in his former Mon- of the ( Eup rbium Boies- “a862 2)— containing pus Ас species E. clava Jacq. 1921. FORMAL CLASSIFICATION BY PAX OF SECTION Dia4- CANTHIUM INTO SUBSECTIONS Diacanthae spine Flo wert | branches га 5 ст thick "niri чой es 2-3 cm thick Triacanthae 1 vem pe 2 prickles Tetracanthae prickles un pine-shield: Shrubs 1.5 m $ ра») Herbs to 30 cm high ag: 1 m high or more high or more Stems short, Tre Scolopendriae thick i^ eaves present, spirally arranged on Section Treisia t hick, erect ste In 1931, Pax & коиот reduced Diaca thium to subsectional status under sectio iii phorbium Boiss. Qe. n i in the worm all succulen mmed species of the genus. They adopted Berger's system more fully in order to regroup the no wield of disparate species in Pax's original (1921) concept of Mon- 8 o these more as groups without formal ra ES SUBSECTION yena + INTO GROUPS Splendentes Madagascan spec i Spine-shields separate: leaves large Herbs with a short e k stem Trees; branches 2-win Trigonae Trees or shrubs; еня 3(-4)-winged Polver nae gies s goi nches decns i t 9 — Bra ч гапсһез ирили: багад " (fused) spine, 2 prickles Tetracanthae Shrubs or herbs; 2 spines, 2 prickles 8 Dyer & Sloane broke with e n the banz a habit (dart. herois shrubs, 2i kiirii nt of the cyathia were also used as important (тондун in deciding relationship 1941. bk ike BY 6 DYER & SLOANE OF SOUTH AFRICAN SPINY SPE herbs, à des p or less Shrubs, 1-10 feet high Branches < 1 inch thick; cyathia boca arranged E > A inch ied cyat athia vertically lly arranged Cymes soli psules large, subsessile Trees : ч Cymes 1-5 in each flowering-eye; cyathia vertically arrange Cymes solitary i h fl ing tally arranged - cvathia horizon- y two latest treatments reverted to the ers а tor peduncles of the and занал with nin deri sidered, from stipules. This latter group whic wh ed concerns и: -— nae he amalgamated to іт s, sh and he ous roo Trigonae 2 Pol berba clude all other pie ылайын" rubs, à Mon- apart from those in the Compressae. Sections RA acanthium and Triacanthium, with equal $ Volume 81, Number 2 1994 Carter Euphorbia subg. Euphorbia to Euphorbia, he used to accommodate the few ining single-spined species 1954 AND 1960. INFORMAL CLASSIFICATION BY JACOBSEN OF SECTIONS AND GROUPS IN STIPULICANTHA USING BER- GER'S (1907) AND Pax & HoFFMANN’S (1931) ARRANGE- MENTS Euphorbia (Diacanthium) 2 spines Teretes Group 20 (Splendentes) Group 21 (Grandifoliae) Costatae Group 22 рена) Gro эд = (Compre: Gro 24 (Trigonae id Polygonae) onacant ti 1 (fused) 5 spine Triacanthium 1 (fused) spine, 2 prickles (1 967) eie followed this treatment, can- oak нА each spine-sh es 1967. CLASSIFICATION BY RAUH OF SPINY SPECIES INTO SECTIONS NS (Euphorbia) 2 spines; trees, shrubs, and 1 (fused) s spine; s 1 =) spine, 2 ee herbs pines, 2 prickles; herbs nthium NE LL Tetracanthiu ит һезе arrangements have been based pe of the Meis Poe also be : in à cculence has developed as a device for sur- vival in arid re egions. Wi es a decrease in the size of the песна ч stages in this pro- gression da be f in subgenus Euphorbia, jas in habit from trees, to shrubs, and finally dwarf herbs. The weakest d of succu- lenc mcn least advanced s adapta- tion—is бет їп ое! trees get thinly беку, вей Ьга sitat and weder associated with a decrease in size of the t-bod eyi es occur in a red the бр сенбе, бе: а of the cyme, а ге- duction of th in the fe 2 bos. capsule shape a pé. size, and features o ee Com mpane features of 6 Ё M — trees S ро ssible ‚мр а= І greatest “пок during adaptation to an ‘arid en- hi te can be regarded the meee persistent Mover of , hav some trees and shrubs, iven way in most species to УРТ sma all or minute leaves that are го ged angled m pow ridges у to form a continuous horny margin. Stipules, ud though s mall, ee species, whereas in the shrubs, and e cy me at each e gro ring- vertical Ariens se: ше lateral Ba Most rbs a flowerin; 976 considerably smaller and with lateral mong th no mo a ri hrubs, and many ees, with t psule reduced in size and obtusely lobed. Finally, although th to be no and tuberculat ate. ing these features and the way in which they have glas developed within groups of obvi- 372 Annals of the Missouri Botanical Garden Character states, distribution 1 f Е, harh and Te ELL of subgenus Euphorbia. Cyme arrange- Subsection Habit Leaves Branches Spinescence ment Section Euphorbia Euphorbia Trees Large, persistent Terete to winged Shields triangular, Several in separate eac flower- ing-eye Ingentes Trees and shrubs Large, persistent or Winged to angled Shields быу ыз, ]-many small and decidu- e, disin in each 5 fen ating ower: ing-eye Spirales Trees and shrubs Large, persistent Angled to terete Shields triangular, Solitary separate; spines paired or single Scolopendriae Small herbs; roots Obvious, persistent Terete or 2-5-an- Shields triangular, Solitary fleshy or small an gled separate cidu LI Segmentes Trees, shrubs, and Small, deciduous Winged, segment- Shields joined or 1-тапу herbs; r u- ed, 2-8-angled slightly separa- іп eac ted flower- Costatae Trees and large Small, deciduous Shallowly segment- Shields separated 1-3- shrubs ed, 3- 7-angled or joined many in each flower ing-eye Section Tetracanthae essiles кан and herbs; шь deciduous 4-angled, rarely Shields separated, Solitary s fleshy or tu more rarely joined; spines paired gle Pedicellares Small, deciduous — Shrubs and herbs; roots fleshy or fi- 4-angled to 16- ribbed Shields separated or joined Volume 81, Number 2 Carter 373 1994 Euphorbia subg. Euphorbia TABLE 1. Continued. Cyathial arrange- Female Geographical Subsection ment Capsule perianth range Examples Section Euphorbi phorbi Vertical Large, acutely Obvious Thailand west to ctea, E. neri- 3-lobed, exserted India, Arabian jr E. anti- Peninsula, and quorum, E. northern tropi- qarad, E. teke cal Africa Ingentes Vertical or — Globose, fleshy, just Obvious Arabian Peninsula, I. ammak, І. ka- horizon: exserted West Africa, merunica, eastern tropical abyssinica, І. ica from Su- oo 1. dan south to ingens atal Spirales Horizontal ^ Large, acutely Reduced to a rim іне and central 5. desmondii, 5. 3-lobed, exserted tropical Africa sudanica, S. oissonii, S ific Scolopendriae ^ Horizontal Acutely 3-lobed, ex- Reduced to a rim Angola to Mozam- 5. imitata, S. de- erted on a re- bique and south сійиа, 5. knu- curved pedicel to the Ca thii, S. stellata Segmentes Vertical Large, acutely T or reduced Arabia and Soma- S. cactus, S. bal 3-lobed, subses- rim lia south to Mo- lyi, S. nyik sile zambique and S. breviarticu west to Angola, lata, S. coo, Namibia, eri, "is yandi the Ca со enormis Costatae Vertical or Large to small, an- Reduced to a rim Somalia seo to C. robecchii, C. seldom to lobed, ex- Mozambiq wakefieldii, C. horizontal ѕегіеа Angola, Nam nib- confinalis, C. ia, and t parviceps, C. Section Tetracanthae Sessiles Horizontal ua obtusely Reduced to a rim jc and Soma- bed, subsessile lia south to Mo- ы дое, Ап- gola, Namibia, and the Cape Pedicellares Horizontal Small, obtusely lobed, exserted on a recurved pedicel Reduced to a rim Arabia, Socotra, northeast and eastern tropical Africa grandidens, C. griseola S. triaculeata, $. polyacant S. uhligi Р. heterochro- ma Annals of the Missouri Botanical Garden Euphorbia Costatae / Segmentes Scolopendriae EUPHORBIA ET i Euphorbia. F RE 1. Di ti i Куа чүп hi ү ERR T ) 1 d A eius soa Ense J, (eos 22 1 1 1 f p h subsection. ously closely related species, an outline sc cheme for the ane п һе 2 ions E Maps рое groupe or subsections. Inevita roupings, and this has been used to in- а g dope Wende in development. Ше most obscure и species Бау an early, more ге pri imitive eas those with features at йе most Mcd eu are the most easily defined. These ar nent, into two sections: section Euphorbia, co g species with more primitive character Siue and all with smooth, d ipee or "ое seeds; section eire advan ыйы ын teristics, all with 1 or-less ovoid seeds (see Fig. 1 and Tab Among the few representative species Tes 12-15) of the subgenus in India the Far East, all possess features at a ge re- garded here as primitive, with о y slight modifi- e species t oward an adaptation for survival in a drier environment. As an example, Euphorbia neruis bi is a tree, with b branches, roun ally arran pe large leaves i. ps cm gem triangular fein е stipt ules, rom each lowering: -eye, a lobed p indi Ti the ovary, an exserted deeply | lobe deg: psule, s emooth id obose seeds. Eup horbia pee modi y slig its obscurely 3-ang какы ез ә persistent leaves, d — branc c 5 еа о a i G, n دو‎ vident in a g species from pi forest region of Miu westward across northern Zaire to include E. teke Pax, modified only in its Mire 4-angled branc Y with characters at the This group B ‹ species, st primitive stage of development, со i ost nearl h Ber ger’s (1907) grou he Splenden gE EE нон ие ferred to as Euphorbia at whatever ia Several groups (or possible maps ) arr { to have developed from this basic group ("s Volume 81, Number 2 1994 Carter Euphorbia subg. Euphorbia tion Euphorbia”), with different features under- going change. One that I pie calling the “/п- accommodates about 10 species with a vide distribution covering most of th ne ess harden an e be -lo at dehiscence Euphorbia ampliphylla Pax (E. obovalifolia auct. non A. Rich.), ш апа Malawi. At um ana t spe a and decrease in leaf size. Euphorbia in- iss., from southern Africa, has more fleshy, inged nches, and Miseni m oe on seedlings and youn fum drier but has E. са E weinf. on idis "s ian Pen- la : iî West n x, a 4-winged species, st rbia canariensis L., from pho. m Islands, ter s to share most features gro о shows - Шот шеш е tree, бше; ae instead o hia, a psules tgs pene fused together early in the seedling , шы ter which only single spines are produced. These ег deteriorate so that the mature plants sometimes appear to be entirely without spines. a ce distint since рои s (1669) elestificatiin, and and formalized asa subsection | ny F Pax im Tits frica, are small species herbe or geophytes, pT a la arge tu- ous root, extremely short stem branching at or e: "inten намй ee spine ur EF © апа acutely | icel. h ped within the оир. In one a progression can be shown from ne inii with very small spine- ‘shields and spines, as in Euphorbia knuthii Pax, to leafless angled or winged branches with strong spines, as in E. stellata Willd. The second series, in a group of closely related wi dasa ssion Ром such тее as J. E. Br. p egi indes: to E. decidua P. R. O. ncn СЄ Leach, with a development of the tuberous root to a spherical body from which deci ae а develop sep- arately from the cymose inflorescences, Беагіп tiny deciduous leaves жа small rounded spine- shields. The most probable relatives of the group seem to be species in “Spirales,” through a common ч «904 ап sin пея of the any form persistent leaves, с and ao structure and posi tion, and as sai sharply angled exserted capsules with smooth glo- hield se seeds. Another easily defined group within section Eu- horbia is one I propose calling the “Segmentes,” primitive ea ranged cyathia, often large acutely 3-lobed же» and sm sm species, чи most анана ^ interva als ounded or oblong seg- that in almost all species they have joined to a seem уали — the branch-angles, ы уегу stout ionally giving rise to further spines flanking the flowering-eye (Pax' Intermediae), ( leaves are tiny and deciduous, with stipules hard- ened into very short or usually rudimentary prick- 376 Annals of th Missouri Brice Garden les. One of the | 1 1 ies is Euphorbia primitive features, which they share with such spe г 1 nyikae Pax from East Africa, bearing some simi- larity to E. lactea Haw. from Sri Lanka in *'sub- section Euphorbia,” but with decurrent spine- he 40 or more species within the *Segmentes" се; ation to ап in- е: = o shrubs, and eventually psc herbs, {реше үш ап шон in the size At the sa time, the capsule becomes subsessile among most of the sh Pax, and a tuberous-rooted herb E. bur Further south a similar the tree E. cooperi A. Berger, the shru — e E. Br., and the herh Е, Ror N. i. chii Pax from East Africa, and PE i Dy о bear some relationship to E. qarad in “‘subsec- tio Euphorbia,” thro an unnamed interme tree that occur: Djibouti. J t Africa a series of several tree species wii "ie puer sc has developed, including the inged Euphorbia wakefieldii Е. Br eto sm all E spine-shields and E. quinquecostata Volke ith 5-angled branches and decurrent spine-shields that me joined as y lobed capsules that are well exserted. This series culminates in E. classenii P. R. О. Ball hits. a l-m- -high ш with 6- 8- angled boe nal rented суна, схов нег e smo more or less ovoid seeds. 1 i r as arisen in souther! d Angola and чып shrubs a as E. с lescens H he Е. panis Hiern may ba faund. The complex rbs аах їо with the shrub E. cactus Schweinf., and in no dics a wide distribution, has тайкеси сик a, number of t flowering-eye, with тешу ог шара yc Wi evelop ubby habit, ‘and cymes become изгән with the cy m arranged faster Branches are aways a an, gied Age 77), giving. kx group its mun e of “Cos Meat winged ei smaller ones more obtusely lobed on slender re- curved ке edicels. An ther ks tree «ресе from Angola cana to Euph . C. Leach and E. parv ceps L. C. Leach have retained most of the more rica, ЖЫ и the tree species Euphorbia gra di aen s Haw., i J angle d branchi ч apsules and Е. zout arbori A. a r, with 6-anged branches, decurrent spin ds becoming eni and obtusely lobed, e sene epee Shrubby cies such as re avasm a Dinter, with a pe epi iia 5, не. branches, have a SA amid: in the semi-desert regions of DEEN t Africa. Most of these species have a tendency to pro: ае 2 spine-shields on short w ree be st segmented. T ee a superficial resemblance to the reduced n the “Segme. к the кайты, difference : that capsu ies ar and well exserted of large and sessile. Several of the 40 or so spec ies ieri distributed tpe “Сонша ае” сап һе кт with those in the two group* а Һауе ies ds most puis in terms of spec Volume 81, Number 2 1994 Carter 377 Euphorbia subg. Euphorbia umbers and adaptation to arid conditions. These advanced and none of the primi few species regarded her section юр hiat Жая" 5 (0921 ) дешин 9 the ing four s spines on ihe spine- -shields. However, this ithin the more Т © PS N sibepefiangi i х i E Ho owever, within this lanh section, rickles is a ze RP Lon ion S. S c g a small, obtusely lobed peer subsessile ere at the most 1 ust exserted from the involucre. Species e es кт dee group— subsection е also possess a i obtusely lobed ule, E thisi is always well exserted on a slender ч tiny deciduous de aves, decurren 8 is оаа cymes vant ишү я аг- ed cyath The “Ses à includes more than 80 E rde g a link with Se shru by and salle related species e5siles," but despite similarities in spi- nescence Р «а зад апа тайаке characters, the differ- ween ovoid tube bh A UE globose smooth sectional identification. = rbia еи р: . Bally & S. Cangri in the A Cos tae” is a na 2 horizontal cyathia, and only shortly PERS cap- рй na L. C. L he *Sessiles," en- demic in the same area. Sim ROR e | species i southern Africa in the “Cos , the shrub E. w bsec s R. аа е pos- ere ‚ ве lobed capsules, h shrubs such as E. ly- n the “Sessiles.” arge group at such don a number of dis- es evolved. For example, uther: bd iculata R. A. r, fro enlarged spine-shields and appar- ently аена наив sure to be related. Further north, the hubby Е; mala hole » Brod Qu E. ambros eae L. C. Lea ch from Mozambique and rom southern Tanzania. One sma gro inctive "ens with al- most peltate involucra glands, ra d to E. an gustij flora Pax t t and southwest Tanzania. Other subgroups have East Africa, such as E. uhligiana Pax d tospinosa tnan rour Africa, e Е. 1 aokoensis L. С. Leach. In the northeast, as 27 as the Red Sea Hills in Sudan, . polyacantha s. has developed into a 6-7- angled shrub with a stout spinescence . A related named species with more slender й. 5-апр} Annals Мал SOON Garden branches occurs панне the hilly regions from Djibouti eastward across northern Somalia. A feature that has developed RE anong plants of drie е the oe spines in aoe sain stage, when , to produce what appear to e a single s spine. The pinos or áb F n * (e.g., ыы | (1 904) AMD а засн including the shrubs related to E. Rift Valley of northern Tanzania and southern Ke- nya as ves Pei with ч агре ЕБ ан oot PGES vith мы prickles. Another group from к а length between species. Other species dor this nein d *medusoid" habit r to E. тшш Forssk. from Ше subgroup occur in Somalia = eastern тр насан vem with spines only partly fused, are found in Ё. schizacant d Pax, with a short, thick, many-angled stem and thin, trailing, 4-angled bran errs oe E. идиш Pus E. a 2-m- hig h oth s ове t Kenya, southeast Ethiopia, and their relatives occur in the dr чире of northeas and southern Somalia. Within subgenus —— ia, , the most success- in — and кн pasao Islands. At the southern limit of distribution of this group, orbi in rie RAI Manan nia, Ep ii RN roma lassendi in the “ taide, тос differen in ме 6- 8- аар branches wa dis: tly globose seeds. lo: drie каш areas in Kenya, Бакайы, par southern Ethiopia a, E. het A gives чау, шап orion progress шн to shorter spine- shields нр ina ville e margin, culminating the various forms of E. tescorum S. Carter In Somalia and the Og eat region eastern Ethiopia, adaptation to the very specialized semi- desert conditions created by a limestone soil and outcrops of pure gypsum has led to the develop- ment of та SRIOSB br: аа stem with и of up to 30 cymes, maturing su . Euphor ми чыйп гтїз н; that horbia piscidermis M. G. Gilbert also, having developed. a vice body and an ex "scale," with a pair of minute spi id on tiny seedlings There are numerous species from Somalia (many ed) in several series, leading to similar t onlv 7 densely packed nches. S inel. с. spine-s shields, like Mu к lenbeckii Pax, while other margins along the angles, like = hillipsiae i E Br. In common with other examples in Africa of a northeast- northwest pane i a disjunct group with obvi occurs in the cies related to Eupho from the Moroccan с branches and spine-shields } gin along the angles, as well a exserted on recurved Other species of re extend sr the Arabian Peninsula in at 1 st three forms © — in a narrow mar small capsules Balf.f. from Socotra, with 6-8 sad br: ehes, a also belongs here, the ii and a i shields hav rin av the absence of h as Chamae other groups not as diversity a syce?). It shows as sneer if not a greater, Volume 81, Number 2 1994 Carter 379 Euphorbia subg. Euphorbia 0 f for rm as the rest of the ve put wee from erb: d on to identify its relationships positively; ps the scheme roposed here (see Fig more stable features of t capsule and inflores- cence are of p etermining re- ^mi Habit and Negat characters of spi- scence and of development within groups, as well as be them, during the gradual adaptation to an arid environment. Relationships of this very distinct and apparently isolated group of mined. adagascar, small le 1 Rest Кюн fro rom succulent, cylindrical ra 1 Кор ntly ca leaf-scars es surrounded y glandular tissue that co perhap homologous with the spine-shield. In personal discussions with Gilbert, it now seem RM à more likely relationship could lie with a Eu R f fleshy-ste med shrubs related to B. z ч synadenium Ridley m evergre ia ot India and Malaysia. These possess large "dn » With s all, persistent stipules, that are leaf- nd often stiff pointed, pn the E prickles on the е spine-shields of rex ded. in * section Euphorbia.” Thi коюуң Euphorbia (the ا‎ species) may have Sinated in Asia and migrated to northern Africa u os c ame separated. Develop- па ens proliferation within Africa followed as num am r. The relatively few for- ound in the extreme habitat conditions of semi- desert regions, notably in southwest am and especially i n the northeast, in the H f Africa. phorbia is based on ev vidence ng ained from exam- d S ome gaps геше неет among de: groups with and fewer —— "i but thes е gape are not t and it i that, as discoveries cies are will be filled, and of n e er the feasibility of the scheme outlined here will be confirmed. LITERATURE CITED BERGER, 1907. Sukkulente Euphorbien. Ulmer, Stut viri h Е 862. соіа: Euphorbieae Pp. n А. de Candolle (editor), Prodromus SRM ыи Regni Vegetalis 15(2). Masson & Son, Paris. Brown, N E. 1911-1912. Euphorbiaceae. Pp. 448- 608 in W. T. ossa Dyer (editor), Flora of Trop- ical Africa, 6(1). ve, London Lo ABS) Касы: Рр. n W. S етеу Dyer (editor), Flora бич E caus dein Es ` 1988 Euphorbieae. Pp. 4 M. Polhill @ О i of T Euphorbiaceae. Balk е. М. С. orbia with comments on the € group- ee of i its African members. Kew Bull. 4 1-244. LA. 1812. Synopsis Marcii ved dd London Handbuch der Sukkulenten Pflan- en (Ba да M Fischer, Jen lent Plants, Vol. i: , Blandford ven London Pax, 1904. Monographische Übersicht über ДУ баны Arten aus der Sektion Diacanthium = Gattung Euphorbia. Engl. Bot. Jahrb. 34: 61- e Pflanze nwelt Afrikas, Euphorbi- ae. Pp. on n Engler (editor), Die Vegetation der Erde 3(2). ае Leipzig. à M 931 HOFF. . 1931. Euphorbiaceae. Pp. 11-233 in A. Engler et al. (editors), Die Natürlichen Pflanzenfamilien, Aufl. 2, 19C. Englemann, Leipzig. Raun, W. 1967. Die Gr rossartige | Welt der Sukkulen- ten. Parey, Hamburg and Berli „Аз: TA рү R & B L S10 OANE. 1941. The lent E uthern Africa. Abbey Gar- den Press, PHYTOCHEMISTRY AND SYSTEMATICS OF THE EUPHORBIACEAE! David S. Seigler? ABSTRACT Ph 1 l info between ша family p putative ees Within the condary metabolites pr nformation concerning the relationship of th: larities in chemistry to bot e Geraniales an lval that the differences йлы the Ош and Rosidae ma Euphorbiaceae may have arisen fro ormation Sadak erecta both within the Euphorbiaceae and Euphorbiaceae, the prese metabolites support certain infrafamilial groupings, whereas data he seco! i vide info i i ce of several бор Bn -— am Бабак esis related to t Euphorbiaceae to other amis. S are found, supporting the suggestion by ار‎ er (1987) T: not be as great as previously thought, ibd that the those two groups. The chemistry of the Euphorbiaceae is among th tdi d int n fa 1 1 families (Evans, cee b; Hegnauer, 1966b, 1989) and is comparable to the biological diversity of the family (Валоне лета 1989; Mehgten. me tabolites, ahy. species of euphorbiaceous е are poisonous and have been involved in human and livestock poisoning. Plants of this family, es- pecially those of the genus Eu or жер tea to ind me 1980). Others have been use ne, as piscicid i eve а] nda number are important economically as ан plants and as sources of rubber, are known ph Rh + 1 ical ааа and medicinal И (Radcliff Sm ‚ 1986; Rizk, 1987). з principal goal of this article i is to evaluate mation derived from ex- tant ph ytochemical erature iow data iid to tain affi ft ДИМ of subfamilial taxa of this шл famil well. What types of ы on ae x are ШЕШУ to be usefull Во! h th о seconda i hi Aas А $: E are widespread among higher plants. Examples are and and ERE the Mein pelle leading to kaurene, М bly occur in all p a oi PPI a relatively short aceous tax a diterpene eae in seve inogenic ic diterpenes, foun elaeaceae, are de lvin| hat possess these compounds a related is i i mre phylogenetic lant 0 the lar uH num- establishing taxonomic and enano. e Mine ae both morphological and chemical be hip The pr roducte of relatively short pathway morphic characters. Many exotic and int А Ь compounds occur in a single qud or Rs Freie to have oribiuated several times and usually species (Hegnauer, 1989; Rizk, 1987 і ' The author thanks the oe raped Foundation (NSF BSR 82-15274) for in St., S. Goodwi USA, support. ов 61801, 265 Morrill Hall, University of Illinois, Urbana, Illinois ANN. Missouri Вот. GARD. 81: 380-401. 1994. Volume 81, Number 2 1994 Seigler 381 Phytochemistry and Systematics of Euphorbiaceae ae of [рле from кс үрен шаан ponens in OU deant = of the нова СА article makes по at- tem e lite vices ле ш ө the imc ae — Reader for more detailed information. Ap specimens maintained in many iden and 8 pr ay studies. In other instances, th identification of th plant material was not verified by a к» this is especially important for the Euphorbiace Major TYPES OF SECONDARY METABOLITES IN THE EUPHORBIACEAE all chemical classes, th ful for che- otaxonomic stu y of t рези above the level of gen ppear alkaloids, cyano genic glycosides, diterpenes, glucosinolates d and other tannins, and iriterperies (Heg- E. 1966 , 1989). Th several ea Alkaloids, diterpenes, tannins, and triter- e arise from complex pathways; th have limited i E е 9 i ра r subfamily level, these compounds m: ds. as u mu eu at lower taxonomic lev- ALKALOIDS Alkaloids are widely PRPS in RIES These nous c nds very simple, redominance of b, 1989; d 1977). Among the relatively simple and widespread al- Aleuritidae: Aleuritin. N,-methylte ита Spathiostemon javensis ria (Blume) Mull. Arg.) (ead Aaa phea c: (Johns et al., 1970). An ester of ч (3) (а yrrologuinazoline alkaloid) has н reported od Croton o Schltdl. (Cro — "Gritotioi) v some ‘uncertainty exists 1966a, b; Rizk, 1987). Several ларек alkaloids f the Eupho e: Phyllantheae: Astrocasiinae], 2 ноне v- dimethylallyl- Æ- ТЯ ©) Hip rega agellocha r Euphorbioideae: M ^ locroton SpP this pies is only weakly lr 87). Ho EEE virosa а d.) Baill. (syn. 5, sch Pax & Hoffm.) (Phy зуна де) (Hegnauer, 1966b). Pyrrolidine and aint have been reported from depend principle of Jatropha po- oideae: Jatropheae) is tetra- also known from ane ma. OQ, Bu- н; nasi shonkd —— r & Foo 5). Imidazole indus have be poda from the ? See figure at the end of the article. 382 Annals of the Missouri Botanical Garden genera Glochidion (Phyllanthoideae: Phyllan- с коке) and Alchornea ойыр, : Alchor а (Johns & Lam- bern n, 1960 6 Beyer pipat 19995 Riz ч E leyis (15), IV*-oxodecanoylhis sta- mine e (0) gloctiüdioitie a 1), апа аР е — о Alkaloids with th ] struct ral diego groups of plants (Acanthaceae, podea eê aceae, Orchidaceae, ae), but the structures are so diverse e alkaloids яо һе ч кш biosynthetic gin gx ler Th e асас pen — —€— of AL ntain a hem e and a chornea c lso r sent an unusual EERE ia ong ded ve rimidine alkaloids of Alchorne v sis Mull rg. are alchornidine (13) and wis (14) This plant contains /V',/V*-diisopentenylguani- dine and №", Л, №- rišopentenylguanidine (Heg- nauer, 1989; Rizk, 1987 Because of the w caprea occurrence of the the Euphor- biaceae, their presence does not suggest or confirm f. ms 8 Ld ц | 1 p K+ Не е уег, other t ] 1 а. Eds 1:4. E 1 14 be exarhined fer. the p f imidazole, pyrim- 11 tively, as these compounds с ould ciet useful. checabiers at the specific and generic level. In contr: ast, the distribution of benzylisoquinoline alkaloids is restricted among higher plants. Most cur i in families of the Magnoliales and A although sporadic occurrences are known from other i alkaloids are common in the followin, set, 1985; Dahlgren et ки 198 In the Euphorbiaceae, ae (thaliporphine and glaucine), vint E hinandie of benzylisoquin: orotonsine, pronuciforine, priis (17), thali- por phi ine, crotsparine (18 N,O- Pes N-methylcrots sparine, crotonosine (19), flavinan- Haê been reported from Andrach (Decn vu Mil Arg _ Phy pe neae: (K ). urinine alkaloids are a pub group of dice raat han ntine, phy ne (24), suffruticosine, and virosecurinine (25) ll 1987; Seigler, 1977). Their presenc members of the Phyllanthoideae reinforces AU Ra of the genera Phyllanthus, Fluegges 26) (Ricinus commun I ph Acalypheae: Ricinin ам), a асе PER (28), жыр, 3-carboxamide-6- pyridone (Trewia spp m repine (29 ) (Malos дарана Acalyp 4 ae) all appear to be closely related and are pr RE 2 derived from nicotinic acid ( E santi, 1985; Sastry & Waller, 197 Sim à Findlay, 1985; Waller & Nowac ki, 1978). 4 structures of these alkaloids suggest а similar cá ynthetic origin from nicotinic acid as 18 sige x for the cyanogenic glycoside acalyphin (^ i (Acalypha i ndica Г. as alyphoideae: Acalyphe? Acalyphi "198 87). kaloids. such as jar (31) ra May- JE Paa al tenus and | Colubrina of the Rhamnaceae (Rei Е » 1984), but also have been tered in the ge" Trewia (Powell et al., 1981, 1 Volume 81, Number 2 1994 Seigler Phy dein and Systematics of Euphorbiac Peptide alkaloids are known from Hymenocar- (32) ste been isolated ceae, Sterculiaceae (Melochia Waltheri a), iw Urticaceae (Schmidt et al., 985). CYANOCENI Plants from subfamilies resin cie Cro- А һе Eupho Sors, and although опе of this acalyphin (30), is found aay in hé hib. others are widely distributed. veral species of the subfamily Phyllanthoideae oe ii J. Léonard, B. mollis Hutch., їса Merr.*, vata rvilleana B. scleroneura Miill rg.*, B. tomentosa е, Phyllanthus acuminatus Svaki P: &ustifolius (Sw.) Sw. gasstroemii 5 ан сипаг йеп = F hu PP kie s ber ios, gea suffruticosa (Pall). ae * Lis Sons ings Ee (Pall) Rehder, S. ramiflora rg., and Valen, BAT S. flueggeoides. Müll. бнаа g.] СУ a to contain oo (33) as унде суапо- н glycoside; other r species are known to be genic, but the compounds have not bee ch e Plants of Ph hyllanthus gasstroemit m taxiphyllin (34). nid of Bridelia monoi- км : hurrin (an enantiomer of dh eom Her inin ran, xeya — ovii ), but no other с пз have sub- Stantiated fr ind. АВ E species of ف‎ subf: dem (pate hree cyanogens are derived from t * тае of ече of the oe pe nown $ are Prin rites trisperma Вы дене ы naultii (DC.) Baill. апа B. viscosa (Labill.) Miq (Ricinocarpeae: Ricinocarpinae), pee species of е (Maite), sese: atus L., C. ctatu. 8$ i Hook. f. En pe^ um Sene “Blan siena several Heveat species смесен dreae: Неу ‚ тап species of Manihot (Manihot ag эв Quae (Manihoteae), (Adsersen et al., ; Hegnauer, 120a van yaan РЕ Weakly колшо tests [m Jatropha (e. g^ J. © Sond.). Most of the d in this s arin (35) and usuall taustralin (36) (species marked Ф). In ad- ion, careful examination of anogens of aled t corresponding diglycoside, linustatin ro, 1986, 1988). rhe ا‎ lines of еп brasi- and 8-81усоѕ sidases o the Sou th American leaf b ight Ghee | 198 vea орн digircosides, pee as linust фин appear in the plants and are pro ‘obably ne widely несена than realized (schon et al., 1985). Several species from the Ds also are these are Aca- c a pecie. Ricinus commu and 1989; Nahrstedt, 1987; van E ана isolated from Acalypha indica E 1987). ily Eu pee (L.) Millsp. diea irta L.), C. culata (L.) Small чы — (tut ) аас recurva (Hook. f.) Burch mifolia (L.) Millsp. (Euphorbia агт ) С. viminea (Hook. f.) rbieae: зня orbiinae), Euphorbia P ip pestis illie & Hook. (Hippomaneae: Hip Annals of the ag Botanical Garden pomaninae) Macer agi nm. To date, the cya- known from Croton corylifolius Lam. (Schmidt, no, уе іс соп е Euphorbioideae have 1987). t been ee а Cocarcinogenic diterpenes. The powerful ENSUES ES purgative properties of “croton oil," the seed oil A large variety of diterpenes occur in the Eu- of Croton tigliu i ee! (неве 1966Ь, 1 989). чя тау е (Hecker & Schmidt, 1974 pounds, and labdane paranoia irri n oc & Soaps dissi from blu LUN Schmidt, 1980; Evans & Soper, 1978; Hecker, P P ESTANY 86У PY- 1977. 1986; Kinghorn, 1979). These compounds rophosphate. Geranylgeraniol derivatives аге TK ters Aij Shido asd T. A Kore ээ a nee ius t ey are not directly carcinogenic, but pl 1 One simple E 18-hydroxy-6-cis- plication of a Careinpsen following contact wi RN (38 n ШК CUM Rep COesrminagenic diterpenes results in greatly en кк) analogue. T hanced carcinogenic activity. At the same time, i for their com topouad protects die digestive tract from the нее „сотроцпда Deui eie i ck- effects of ulceration and promotes wound healing antitumor properties (Evans & РО з Crot AR (Croteau & Johnson, 1985; Sato et al., 1980). Шш these compounds are derived Cyclic diterpenes from geranylgeran ro- from monocyclic diterpenoid precursors and are ge geranyl py. M Р P phosphate. Cembren e(39), „Беи закс on ch bi y related t as ds of the and the jatro n hyrols, sedie jatro- are fou ost important among these t n hanes, rhamnifolanes, and crotofolanes, as w gena ch as ) (Sch , 1986b), the cocarcinogenic tiglianes, daphnanes, and — daphnane (48) (Schmidt, 1986a), and tigliane (46) ingenanes) are d 0 clization of a gera- (Evans, 198 ) series. nylgeranyl pyrophosphate precursor to a lar, nanes, and daphnane type € Е yclic system (Sch 87). Casbene (41), genic diterpenes have been isolated from the Eu a bicyclic 14-membere g pound, is a phy- phorbiaceae ell as Thymelaeace: a 1 ае. оа анд ip dicun" ишаа, ae :compound арз eim pis оин (tetracyclic with a SIX x AS x : P g) in fab and a MUR. ruens (Th ymelaeaceae) as well a5 era pyrophosphate (corn) (Schmidt, 198) mpound w Aleurites, Croton (Crotonoideae), Euphorbia, ted structure has been Hippomane, Sapium, and Synadenium (р wa from rin itens Sw. ioideae: „Бийин Euphorbiinae) (Ba spies Several unusual structural types of үрүн et 8; Hecker, 1977; Evans & Soper, 197 derived from monocyclic precursors in the Schmidt, iE 6c Euphorbiaceae. ati phanes (such as 42). are aches ngenanes (eg g., ingenol) (eter m bare ae Jat atropha gossypiifoli ia и » J macronbide seven-membered rings) occu in Eup ate Elaeophorbia, considered a feito i^ Eupho m Е mandand e ., E. helio. scopia L., and E. bia by Webster, 1975, 1994) species pee Боемо, 1989; Man- 1977; Evans & Soper, 1978), but not in 0 кай (43) taxa of the Euphorbiaceae and Thymelaeace2e hamnifolius Korihi were examined (Hecker, 1977). (tri быша pian sar (such as 44) are aphnane compounds (е.ё., daphneto xi га nown from Bertya cupressoidea Airy (Cro- cyclic) occur in both the Euphorbiaceae [E tonoideae: Ricinocarpea rtyinae), Euphorbia permum (Crotonoideae Codiaeae), Cunurt ( hia, helioscopia L. E characias ens tonoideae: Micrandreae: Micrandrinae dese E. Meyer, E. jolkini Boiss., E. lathyris L., Excoecaria, Hippoma Hura (Eupho ЙА aranga tan M Arg. (Acalyphoideae: Нигеае), and Stillingia] and th Тули Acalypheae: Macar ae) (Hegnauer, 1989; (Daphne, Daphnopsis, Diarthron, Dirch Schmidt, 1987). а (such as 45) аге Volume 81, Number 2 1994 Seigler Phytochemistry and Systematics of Euphorbiaceae and Wikstroemia) (Adolf et al., 1977; Powell et r 1988; Hecker, opening of the cyclopropane ring (Evans & Soper, г ou in the jim daga bien ra Cham- r aesyce Lun Poi a lack cocarcinogenic diter- penes. th ros and ingenanes are commo in subgenera Euphorbia and Tithy: & Eum. vM Em hs he ah a 3H- SU (ented phorbol receptor edes as- say) untested samples of eaphorbieveous plants. Activ- rypetes (Dry- a (Uapaceae)], that fom i p^ idesma was ney p re et al., tide Cleistan- s found in Cleis- ee schle chteri ee ' Phylanthoidea Bri- delieae e) (McGarry et al., 1971). Onl y weak activity was observed in the к (Alehor rnea and Trewia) and none from the 2 Ж йм activity was found in Cnidosco- 5, M: Ricinodendron (Aleuritidae: Rici nodendrinae), Codi mong fordii (Aleu oy E i АК i н (Schmidt, n the Euphorbioideae, crete! was fou Slostyles (Stomatoc calyce Ho nd in e: Stomatocalycinae), : Carumbiinae), Du- x ou ium, pre Sling 89). A large number of o 2: à were examined and found to lack dii ac- ity, Diter (49) is rpenes derived from kaurene. Probably found in all pl 0 berelli Kaurene W (Bakker et al., 197 72; Ghi any pimárane and abietane derivatives are found in ri ае, EE ib o aceae and (Croteau & Deos ioo Th Med л ү сиг іп the eaf-su P» ace resins bos of the tribe Cr IN бон. А Ficus 1985; Dell & McComb, 1974). Ox china diterpene derivativ xygenated diterpenes pem on de ceae is (Cote au & (кем оп, som d 1981a Е с and restricted ih u e, they are pri arily found in the genus a tie oton pn Seigler 1981) Fı uran formation between C-15 C-16 is mon. Such compounds ylifoline (Burke et al., 1976) and ete on At least two ата or this type are pua in Mb uot ) Benth. | р еї as 1986; folii et iL) 1982), and " compound, mallotucin p price IM ans repandus (Willd.) Mii tical to t in (52) [ crium (itn bea] (Kawashinnn et al., тё iginally isolated from Teu- GLUCOSINOLATES Glucosinolates in the Euphorbiaceae are con- fined to a small number of taxa. Most im opens among these are Drypetes ue ri S. Moo and Putranjiva roxburghii IL. шы with Dry- ae DORE trey is 954). Glucoputranjivin (5 3), glucocochlearin GN and e амр һауе ДЕК, solated from Putranjiv rghii (Rizk, 1987). Although ‘cnt are spe from ni bie MN and nogen at ined of this subfamily. Benzyl ey opha apes = (Ri k & El-Missiry, 1986). ўн igher plants, de distribution of glu- mbe cosinolates is restricted x a small number of fam ili ; Rodman et al., 1993). One group of families sai es the Brassicaceae, Capparidarsas, Resedaceae, and Toren iaceae „A ng mpi families вррейх to be related to мое families ei related to al other, but the беке of be nolate-containing plants (Hegnauer, 1966b; Rod- ‚ 1991a, b; Rodman et al., 1993). Annals of the sees Botanical Garden SEED AND OTHER LIPIDS mber of unusual fatty acids occur iiem of the an ese involve l sites of un saturation and E double bonds. For example, Sapi um sebiferum xb. contains decanoic acid with 2-E-4-Z unsaturation (55) and Sebastiana ichx.) Müll. Arg. contains dodeca- This rapidly pori acid is also fo il of Garcia nutan , Ricino bg ar viti- coides Mildbr., and R. ni nenii Schinz. (Cro- tonoideae: Aleuritidae: Ricinodendrinae), (Heg- auer, 198 obson et al j tillingia texana 1. M. Johnst Si ligustrina (Michx.) Müll. Arg. (Heg- a 3 Е о ix. © Б їпз 12-Һ droxyheptadecanoic i and 12- hy dr голуба a s (Heg 1989). The constituent of du seed t м Matos ME Rp (Lam.) Müll. Arg. and Trewia . is 18-hydroxyoctadec-Z-9-E-11-E- 13. trienoic (a-kamolenic) acid (6 {Benita & Guns (63) ] (Kleiman ét al., Rees Cephialocroton cordofanus Нос hst. Cuiistone, 1956), ме: | Euphorbia lagascae е: (уе rnolic о man et i emis Smith, -1970). The баганада compound ,13-epoxy-11-methyloctadeca- 10,12-dienoic ac 5) occurs in the latex o Hevea id (6 rasiliensis м Juss.) Mill. Arg. (Назта & ane aniam, 1978). y fatty acids are found commonly the Aa as well as in several other PA ilies. These compounds are thought to be derived biosynthetically from epoxy fatty acids. 9,14-dihy- roxyoc de ctadecadien-10,12-oic acid (67) an th. ro-9,10-dihydroxy-1-octadecanyl acetate (66) are foun the see L es fordii and 11,13 acid in Balios r REAR E enoic er TAN lyphenolic compounds known as especially wide among woody plants, but also re found in herbaceous species. Both hydrolyzable and co ins are found in plants ndensed tann the Euphorbiaceae (Rizk & El-Missiry, 1986). Hy- Фуре ere that yed Ale acid 30 оп cies pr im: marily from the genus Euphorbia (Rizk & El- Missiry, 1986). One specific сеза geraniin ( SE is Br ases in family and has been f the Acalyphoideae, Cr Cro- tonoideae, Euphorbioideae, and Phyllan nthoideae (Hegnauer, 1989; Okuda et al., 1980). I [он first de of these subfamili by the Se encountered compound mallotusi ^ (70) but absent iras the Mo. Fagacea Fabaceae. Tannins with quite distinctive raul e been isolated from euphorbiaceous plants, especialy from the genus Euphorbia (Okuda et a T Unfortunately, res; this com ple hs des ec analysis Tannin c is not use the structures of many of these ompounds ч provide poe onal ehemosystematie markers КЫ of plant gro TRITERPENES Many acl of the ape eae possess latex that is rich in eie s. Triterpenes are predomi- pan in 9; 1968; Ponsinet & Our al., 1979; Райна et al., 5). ost of the triterpenes that have been ia from the Euphorbiaceae — e (е.в., В-8ї В-атугіп (71), friedelan, ined ore ere (12) Volume 81, Number 2 1994 Seigle 387 Phytochemistry and Systematics of Euphorbia p (73), taraxerone, B-amyrin acetate. oos Fi establishing tax T taxonomicall e species eneric level ahlberg et al., 1988). Because triterpenes are difficult to isolate, Pur ify, and characterize, many E: those for Tuer re-examined. Euphol Hs tirucallol Mi and РОДЫ [e often are fou latex of members of genus Eu (Rizk & El-Missiry, 1986). m addition, euphol is found in the Burseraceae, Ru- { à : aceae, Simaroubaceae, Cneor nd 1a- ceae, and tirucallol in the Anacardiaceae; either (or both) as precursors for es of met- > Will ompound c ises almost half € sterol mixture (Rizk & El-Missiry, 1986). ere too few 4 ы have be other pieta ese to Pply the data effectively, but olei: plants of m dh other euphorbiaceous genera not accu- ; ate triterpenes to the same xni: as ок species 0 кь. An and distinctive comp ed fro om the us Glochidion k & = Мы, 1986). Be cordes (such as 80), another dem д of “Shenae tributed bers et al., ed among poe are foun ете on genus Mallotus (Rizk, 987; Roberts 66). Cucurbitacins (such as 81) have been denen from the Phy rupicola edd., Clei ; ага, Dry- petes gossweileri, Spondianthus ачак; Engl. (Antidesmeae: бай the геена Саг a "uu Pme сою опеига Prain. (Сагу- l ) ull Arg 1 СІ ), Trewia te pa and the Faphorbiidea EEA задни Mill. Arg., Sapium cornutum Pax) Gian uer, 1989). These extremely bitter compounds ie occur in ), Liliac (Phormium), Begoniaceae (Begonia), Rosaceae (Purshia), D —MÁ civ agb Polemoniaceae (Ipomopsis), and atiscaceae MISCELLANEOUS COMPOUNDS Amino acids. 3,4-Dihydroxy-L-phenylalanine (L-dopa) (82), а ке amino ts is use for тишин. of Pa FE n's disea A a bu t also occurs in posses lathyrus ie (Heg: nauer, 1966b). arins. Many coumarins are essentially letin, and coumarin occu ~ mg сеш ind wn рь Ан адре, Bergenin (83) Riy are found in ue bark of bo T —— Müll. Arg., Iz ta Mill a Arg. (А pores M et al., 1979; Rizk, 1987). Ber in frequently accompanies ellagitannins in eu- phorbiaceous plants, (Чекпе, S 19897 awe Boiss. (Rizk, 1987). Flavonoids. Flavonoids are ubiquitous compo- nens of plants Several types, шоп а s the glyco- teolin ыу the flavonols kaempferol and quercetin are т, ina large percentage of plants examined. Other t of flavonoids, such as какыс are less widely distrib: Rare тене have bons aeta ant (Rizk, 1987; Rizk & El- baroni 1986). É— ke and O-glycosides are {ош as quercetin and quercetin glycosides (e. g. pr чш? tin, vitexin (85), isovitexin, isorhamn ntin, isoorientin, kaempferol e ES rhamnetin, naringen gly-‏ ا 388 Annals of the Missouri Botanical Garden cosides, apigenin and apigenin glycosides, quercit- in, and saponarin, as well as t "mo оз 5 (=) 2 2, = E [nd and also are found in many other plant families. Alih h +h PN wee, ЕДИР. £ sh: ] Ж Pale о © they are involved in attraction of pollinators in several taxa (e.g., Dalechampia). Proteins and peptid members of the te ез. The seeds of several Euphorbiaceae contain toxic pro- eins. For pod the castor река Mou s com- h P Yu 3 £ LI 1 84 i уо! levels, most flavonoids are not particularly useful for study of relationships at higher taxonomic lev- els. The presence of other, more unusual t flavonoids has more pr types of omise in this regard. For oo = честен к Be кореп Ве- enaultii, Ricino- EE stylosus Diels carpeae (subfamily Crotonoideae), inc cluding yeria brevifolia Вай, В. lesc Arg., and (Rizk & El- haberet 1986). ., he veaflavone vA are found e Pinaceae, usiaceae, as monaco, Capnia liaceae, an & 1982). These com- pounds are alas fo in vut mosses, cycads, a L. and in Ginkgo bilob E Anacardiaceae, Casuarinaceae, Clu: NACHRS, bam ардани and waxy materials. Because many euphorbiaceous plants occur in deserts and Pe areas, they have thick cuticles a large amount of 1389; Risk & B -Missiry, 1986). Most of the lng alcohols aldehydes acids, a ÉSLIOD 1 Lh: 3] to those of other каре groups. Candelilla wax of Euphorbia و‎ S Cer, Pedi- Candelilla wax is fepe ia of both Hydro ste fractions and wa erpenes. Monoterpenes are ubiquitous айры. А Беи об ће st нше аге зо сһаг- ma proteins. above the ppodios level. Агаш. genus Croton, Croton s setigera (Eremocarpus se- tigerus), E significant ашары 9 these com- poun g altho t munis defibrinated blood Gad Ad blood cells ui vise As шк chains (MW 63,0 rites оа atropha сигсаѕ id and J. multifida L. a produce toxic proteins that have not been FE pu vely. bber. yos mg Euphoria ines is fond үр imarily ing in the рушса, Oldfieldioidese, P Acalyphoideae (Hegnau 1966b, 9). though latex is a on) mixture of pui ponents, ыу isoprenoid compounds are im portant i el ver: rel plants ‘ol the Еее ЭЕ 8 SLE H ЭРЕ materials | are produced by other Hevea ot gla isolated from many КООШ species (Rizk, 1987 Sesquiterpenes. Although severa ате is re- ported from the family, this class of co unds does not seem to be well represented emer et al., 1979 Rizk, 1987 i of se squi terpene. are found in Oldfieldioideae Ana ulus Reiss "(De ndrobium), poy di Hs ae in (88)] (Cane, 1981; Coscia 1969). Hyaenanc che ds. A series of co ds d luding crotepoxide (89) is thought to o be deri from iso-chorismic acid. This compound is fo in Croton macrostachys H t. A series of Sisi cosidase er including a-homon: ojirim x a (90), h n described from e halea е Le ля еа These co d in legumes, кты Могасеае, i the Polygonaceae and have promise as an itum. and antidiabetic drugs (Kite et al., 1988). Volume 81, Number 2 Seigler 389 1994 етеу and Systematics of Euphorbi SYSTEMATIC VALUE OF CHEMICAL CHARACTERS plants lack the alkaloids characteristic of the B ш poe ДИ, te "iHi is among As entioned above, many of the PR unusual Же of com- ounds are found only in this family and, hence, represent Кер peers Many ot ink ers are of relativel ум Жарга amo ie higher p lan ts. n Although the es mistry of most members of the family E. been i knowledge of the chemistry of this family is be- coming clearer and patterns are emergin n- doubtedly, some NA await additional нЕ а. investiga Triterpenes are ges espread in the Euphorbi- e found i v 8 Top genera of де Phyllant Viu ae Su с. е unite several genera of the phoideae cya nogens of the Phyllanthoideae are ved, that of the Acalyphoidea у, апа He TRE of the е all e nico- P erived dites are foun 7 іп Сенна ss ku iis oidea RENE d we Acalyphoideae terpen Coe dr | Euphorbioideae. Gor emoe enic the BS and ken labd Ps. oxygenated diterpenes based o abdane Bo are found in the Crotonoideae id Acalyp hoidea к жын Es veral chemical features serve to the five Manis The Oldfieldioideae have n oe tly studied. nee are чае similarities between the acera: si the Euphorbiaceae (Hegnauer, Ma ie ba species of the genera Buxus, Pachysan- a nd Sarcococca Крия unusual деда) and oe alkaloid only > "о о аге e distinct from those e t metimes c to be а member of the Euphorbiaceae, ced aceae and E not found in either family (Hegnauer, 1989). The trite erpene alkaloids of the Daphniphyllaceae emble those of the ; Riz no significant chemical калы, between this el and the Euphorbia е Celastraceae and Rhamnaceae both have of. several genera of Euphorbiaceae = egnauer, 190 болун е alkaloids also are et m the ور‎ eae (Rhamnus, Phylica, Col- iei Colu а, Ша, Talguenea, апа Z эри аѕ wel as aom, the a sch аршы (pr imar ily ly га y ү £ ب‎ H it ce erai eie altered ones. quinone pigments sides) is not particularly similar to those of phorbiaceous plants. Some unusual lipids occur in seeds of the eu- Mal- vaceae, Sterculi. шош апа Bombacaceae; о prope а тасу acids аге CSspecially «еи nauer, 1989; “Smi th, 1970). Peptide joues are found in th орлот онен (Melochia and Walthe- ria) as well a epee Borbiacóous Бор: Peers the chemistry vales (Malva бае; E Bombacaceae, Tice Barca arpaceae, etc.) shows some simi- s not strongly resemble that of the try o of members of the Geraniales eae, Limnanthaceae, Tro- paeolaceae, Balsa eae, Linaceae, laceae, Humiriaceae, Zygophyllaceae, Malpighi- aceae, Trigoniacea esemble that the Euphorbiace he Limnanthaceae and opaeolaceae contain glucosinolates, as does the T ig TET eous e Drype es). Hes (and Putranjiva, 1 : f th F dine Linum are cyanogen contain the c nogenic comp mds Seay К linus- , and neolinustatin, as do some mı mbers ompounds amon the Gora nipcene ДООШ monoterpenes: H lip t and glycer ride mixtur пап 390 Annals of th Missouri Botanical Garden though equally unusual floral lipids are found їп Bakker, Н. J., te ISALBERTI & P in m (E enti ceae), and — шщ оїһег А ры" especially the Sol 77). The “Thymrlanene have the strongest chemical link to the Euphorbiaceae. Because of the complex and еа synthetic sepa leading to the for of с genic diterpenes and their highly тааный dici tion (only in t ig Euphor- biaceae an i ly prob able that the families are closely related. This re- ymelaeaceae) lationship has been observed on a mo classical characters as well; both ren et al (1981) and Thorne (1981) placed the Euphorbi- aceae an melaeaceae close t er in their thaps the chemical data speak best for the situation астей by Web: bster (1987), in which e Eupho йу roel are derived from some o the Geraniales а ИУ vale at and ч Rosidae реке 1981) has been exagger зе e thy: пере еасеае rend are closely re- h ig perha ps, th ey sha placed in a dem but distinct NUS and Celastracea "E tho ОЕП chemical m are helpful i in — he final е of the phylogenetic басе t of the Euphorbiaceae remains a goal to be deter- mined PUES future sicud bas s on a com- hi H f 1 1 hw. h > 1 1 1 Жул, logical approaches. LITERATURE CITED ADOLF, W., E. & E. HECKER. H. SEIP 1988. Irritant the mezereon family (Th тамни, oe 4 2 8. = e Dn © gs pt 1 retusa. J. Nat. re) 51: 74, ADSERS| ае м & L. BRIMER. 87. Су- Basi "constituents in plants from the ee Islands. ee hem. Syst. & Ecol. 16: 65- ВАСАУАТНІ, R., RG & E. HECKER. 1988. "Tiglinigo- туре гач ар esters from Synadenium grantii. Pl. 506-510. M R Ж J. CHANG, J. L. MCLAUGHLIN, R. G. POWELL R. SMITH, JR. 1986. кайды ын setigerus. 7 Nat. Prod. (Lloydia) 49: 1174-1175. 972 а of diterpenes in 1 les- chenauli Phytochemistry 1: 2221-2231. BEUTLER, J. A. & A. ЇЧ. BRUBAKER. 1987. The chem- ae and phar macolo ову ot the securinine alkaloids. ; ADO, T. G. McCroup & G. M. CRAGG. Distribution of phorbol ester activity in the Euphorbiaceae. Phytotherapy Research 3: 188- Buanucn, K. E. & F. D. GuNsTONE. 1956. Vegetable oils. V. Component acids of Cephalocroton cordo- i i 06-609. f | alkaloids. д э RL ). Phillipson, M. F. . Roberts & M. H. Zenk (editors), TI of Isoquinoline Alkali кыты Verlag, Berlin. = PRI тч, В.А HAN CEE ee EEO N & J. CLA 1916. eire тула dit rom ton arta lom: Tetrahedron 32: 1881- 1884. M & F. C. GUNSTONE. 1953. Struc- of Уаш | acid. Chem. йш 436- i Con D. bi enes 283-374 in er & S. L. Spurgeon me Biosynthesis of ine Compounds, Vol. 1. Wi- ley, New Yor Coscia, C. 2 1564; vend Pp n W. I. Taylor & A. R. Battersby d rires Terpene Derivatives. ‘De kke CRONQUIST, ife 9 Mer las: ын. of Clas- sification of кы Раа Columbia Univ. Press Жы n wee ra pe of wood ex es. Pp. 3 T. Higu- chi (eid. Biserna and biodegradation of Wood Components. Academic Press, DAHLGREN, R. B. J. NIELSEN. 1981. A NES classification d ihe angiosperms ween ай - in D. A. Young 1983. " riterpenoids. 1-1095. 1974. Hean production Phytochemistry 22: 1071 min . McCol OMB H Lai) Ma мир (рса Austral. E Bot. 25: 195-2 PY Evans, E 986a. Macrocyclic бетен ph ne Ei bati Pp. 139- n F. J. CRC itor, N Natura yo Occurring a Sadi | Esters. a Raton, Florida. D Ph or bol Its esters and de Pp. 1 n F. J. Evans 5 ber siis ele Esters. CRC Р a rivatives. Flor P А comparative es j some ti муы of the Ti. phorbia (be Bo dad 74: 23- NL t. S. Linn. Plants and plant 980. 38: products that induce dermatitis. Pl. Med. 16 . C. J. Soper. 1978. The tigliane, da and ingenane diterpenes , their awg ү tion, and biological activities. A review а (Lloydia) 41: 193- 233 кзг! Prod. Volume 81, Number 2 1994 Seigler 391 Phytochemistry and Systematics of Euphorbiaceae FARNSWORTH, N. R., К. N. BLoMsTER, W. M. MESMER, J. C. Kine, INOS & J. D. WILKES. 1969. А елка). and biological ades E. the genus Croton. J. Nat die (Шоу: DA Fopor, С. B. & B. ke уййне зң piperidine aide. “Che colog Pp. 1-90 in S. W. Pelletier je eg yiii Chen ical and Biological Perspectives, Vol. 3. Wiley, New ork. kon ^n & C. Quinn. 1982. Biflavonoids. P. n J. B. Harborne & T. J. Mabr "i cit, The Bp Advances in Research. bg JEFFRIES & M. A. SEFTO e. Diterpenes from ed ca- lycina Phytochemistry 17: 1961-1965. Croco, S., J, Ban N, J. BECERRA & C. MARTICOREN Alkanes f from Chilean Euphorbiaceae ы К; species. Phyt tochemistry 28: 1254-1256. Hasma, Н. & A. SUBRAMANIAM. 1978 ee occurrence of a furanoid fatty acid in Hevea brasiliensis latex. Lipids 13: 905-907 Hecker, E. 19 toxic, irritant and cocarcino- Я И New genic diterpene esters from Euphorbiac 5 M Г ыйбасёао» Риге Арр1. Chem. 4 1986. Tumour promoters terpene ester type as risk Bot. J. Linn. Soc. 94: 19 EE 1 49:1 423- 14 of "ede irritant x factors of c 7-219. SCHMIDT. 1904, se ene go Sta ants and eens of Croton tigliu m. Organi: fe gee dies a э18- wor. he taxonomic 427 in T. — lie без адет! kaloids. Р тшу o ative Phytochemistry, A (ir lili uphorbiaceae. Pp. emotaxonomie der Pflanzen, Vol. 4. devenue Verlag , Basel. STESTIPI 89. Euphorbiaceae. 440-474 in "DOM der Pflanzen, hs 8. Birkhauser М, W. Н. & R. T. Horman. 1972. Highly opti kay. active кыеш of Sebas tiana ligustri- Я па. Phytoche тіѕігу т E N * Micron du Simple in- dne and quinolizidine alkaloids, Pp. 183- 908 A. Brossi (edito: id = Alkaloids, Vol. 28. H dni с Press, New О? L 54. eli chmalige Durchsicht des i ommlichen Systems dr Chao POS im wei- Sets Sinne. J. Fac. Sci. Univ. Tokyo 6: 209-342. ‚М. M. с 1981. Bar ARYSTAL & Agric. Food Chem. 29°59 € d R. & J. A. LAMBERTON. 1967. a Glochidion oe Family al. J. m J. D. W ARTHEN, Ж rents from tung oil 1-593. New imidazole loids fı Eu- B а Тол, vii A. A. duni з Um Ans tahydroharman E gripe acit javensis ustral. J. Chem. 23: 213 . D. — 1 Ms Dieci cd e IF EAR, M, J. К. COLE E Tempesta . BATES. 1982. Coro. sae, remocarpus setigerus. J. Organic Chem KAWASHIMA, T., T. Nakatsu, Y. Fukazawa & S 1976. Diterpenic lactones of Mallotus E Mi adag 5::227—2832: ‚ М. IKR 5. » Hussain. 1983. Bis- foli УР Med. 47: 191. KINGHORN, A. о Cocarcinogenic irritant Eu- phorbiaceae A -159 in A. D. Kinghorn (ed- itor), Toxic Pla "s ا‎ Univ E iv. Pre: S Na ew 964. koaien Pi of the U.S. nada: 5 ахы Hall, Englewood Cliffs, New lohepitol] from Omph буген inhibition та Іеї. 2 6; KLEIMAN, R D. РА & С. F. ‘Sen att: A rich source of a айога ‘cordifolia а oil: „ epoxide, east 14, fe дызы -cis-1 l-eico- senoic sid Lipids 1 2: 610-612 ——., C. R. SMITH, i. S. С. Yar ES & Q. Jon гар чий E XII. Fify. udin, m. Ass 1986. Cyanogenesis а gore of а brasiliensis pe^ Microcyclus ulei. J. Phy- vh 115: 134-1 88. к. of cyanogenic capacity 3 of the rubber tree Hevea brasiliensis to e y to Microcyclus ulei, the agent causing South American leaf blight. J. Phytopathol. 122: 54- me Bien. 1985. Metabolization b ili is. Plant ————, D. S of pibe Syst. Evol. 130: 49 a . C. Bey The imidazole alkaloids. In A. Bros fei gel Alkaloids, Vol. 22. Academic Pre: hi KA Fk . Furr. 1988. oid profiles ul {ой RT cristata taxa ia and their p as chem omic characters. Monogr. Syst. Bot. Missouri Н Bot. : 623-629. sacking the к вригде v tin rerit Pp. G. R. Waller eret ux Sei in ча and | Forestry o Vol. 330). A shin ingto D.C. apes , L Рнширѕ & Е. S. РЕС 1971. The const ion of the aromatic дете cleistanthol. J. Chem. Soc., Sec. C, 904- NAHRSTEDT, 1982. Strukturelle Beziehungen schen aie und tierischen Sekundarstoffe. "P. Med. 4 ri cen developments in chemistry, distribution, "wh DIT of the cyanogenic glycosides. Pp. 213-234 . Hostettmann a (ed- itors), Biologically Active Natural Products. oa don Press, Ox KUDA, T., К. Mor 1 & T. HATANO. 1980. The distri- Annals of th Missouri ears Garden bution of geraniin and m es a in the order бейш, Vibe eri) 19: 5 xf YOSHIDA, N. SHIOLA & J. жы 1975. An по il rom а seeds = jme fordii. "Plantes iie 14: 2304-2 OunissoN, G., M. ROHMER & R. jt can ae Mac- roevolution and microevolution. Pp. 131-162 in T. n & G. R. Waller (editors), adm in the Bio- iium of Natural ec (Rec. Adv. Phyto- chem., Vol. n Plenum, New Yor Pais, M., J. MAR ito. C Mone UR, F. Јак GOUTAREL. ieee cd Struct de l'Hym anda acida d. He bd. Séances hd. Sci. Compt. R Rastoci. 1979. The triterpenoids. . PIA LP . ent 1.38. “Hydroxybeyer ا‎ -ene- 2,12- dione from Androstachys johnsonii Prain. (Eu- phorbiaceae). Chem. ge a hay 1346-1347. PonsineT, С. & С С. OuRISSON. ‚ 1965. Etud дез chimio- Introduction anh et separation et теке чту, des triterpénes tetracycliques monohydroxylés па- turels. Phytochemistry 4: 799-811. ‚ G. OURISSON & А. С. OEHLSCHLAGER. Syste matic aspects of e distribution of di- and tri terpenes. Pp. 271-302 in T. J. Mabry, R. E. Sitio GV R uneckles (editors), Recent Advances ZA R&C.R udiflora тна и Org from Diarthron I Means J. Nat. Prod od. (Lloydia) 48: 102-1 ‚С. К. SMITH, JR, К. D. Pra & В. ONES. 1983. pee ids fr Trewia Mendes 10- осона апа nortrewiá- pene: sine. J. Nat. Prod. yes уфа) 46: 660-666. RADCLIFFE- калы hu ot review of the fam Euphorbiaceae. js 5 in F. J. Eva ан Nat эш) bi ашы Баа Esters. CRC Pre ss, Boca Raton, RaMAIAH, : re Row, D. АВ ANJANEYULU, R. S. Wan beet Isolation and саи ке of си derivatives rom Macaranga peltata. J. Chem. Soc., Perki Trans. I, pem 2316 Е REIDER, Р. J. & Maytansinoids. D.M.R 4. Pp. 71-156 in A. d eon, The Alkaloids, Vol. 23. Academic Press Rizk, A.M. 1987. Th nomic муе of the Rishon Bot J. Linn. Soc. -326. h & ч. Missiry. 1986. Non de constituents of Жасак ылы and Thymelaeaceae 107-138 in F. J. Evans neis Naturally O Pp. к Phorbol Esters. CRC Press, Boca Raton, Flori ROBERTS, К. D., Е. Weiss & Т. REICHSTEIN. 1966 Die Cardenolide der Samen von Mallotus paniculatus Muell. Arg. (Euphorbiaceae). Helv. Chim. Acta 49: 316- Е RODMAN, J. 1991а. dae te producing Es pa 98 ун taxonomic analysis of glucosinolate- producing ies part 2: Cladistics. Syst. Bot. 16: us taxonomic analysis of glu- art 1: Phenetics. Syst. Bal: 16 E, К. KAROL, E CONTI, KJ SYTSMA & J. D. е 1993. N the mE же ee monophyly of mustard i plan nn. РЕБ К i 80: 686-699. sem, * ud т e . мыйды. "уос ty an ATO. so & H. Kuw 198 80. Асу pens i bois Croton pubs РН еы 19: pi a gear Plants for Man, 2nd Ed. Prentice Hall, "Eglo с, New Je ше ТЛ Р E J The 7-244 FJ. ae Phorbol Esters. CRC Press, Boca Raton Florida. se 6b. bh ingenane polyol esters. Pp. 2 245- ns (editor), bipes Occurring з ^ 86c pects o я t n Phorbol E ———. 1987. e bi ram edi an intriguing problem. Bot. J. Linn 94: 221-230. HMIDT, U., A. LIEBERKN Peptide ‘alkaloids. P. The Alkaloids, н. 26. Academic Press, a SEIGLER, nigh Pp. 1 ke e (editor), The тея ECHT & Е. HASLINGER. р. 299-326 in A. Broes (te New York aloids. TAN Vol. ч {сут ые Press, New Yor 198 1a. iem А үй» pute gen y. Pp. 117-148 in D. A. Young & D. S. Seigler (eir Phytochemistry and jam Phylo geny. К joie No сх metabolites and p b. Se n E. E. Conn cae oo 7. Secon EX Pp. 189-1 PIS $ S ucts. Academic Press, New York. 1970. Occurrence of ий А fatty Situ, C. К., Lip acids in saan Prog. Chem. Fats Othe | ios end 1985. Р S RUNZ, G A. FINDLAY, г prot eit . Ac um Phytochemistry and hos THORNE, R. F 81. phylogeny. A summary statement. Pp. 233- pes in d Y & D.S ler (editors), P c and Angiosperm Phylogeny. Praeger, bete viu F VAN. 1978. арша то Aue: of cyanogenesis in Angiosperms eon Cyanogenesis in Euphorbiaceae. Ei Med. ed. 34: 4 АЗЕ WALLER, С E. К. Nowackr. 1978. а b mii e and Metabolism i Plants. dem rb EBSTER, 1975. n apes ol ei козе a Seigler Phytochemistry and Systematics of Euphorbiaceae Synopsis of the genera and suprage- uri Bot. Gard. 1994. neric taxa of заа Апп. Мїззо! The saga of the spurges: А г in the a be 1987. classification and ў 2 tionshi 3-46. E ome) Novon 2: 269-27 ова. in American Croton (Eu- 3. CO,H N m CH. NH NH 5 L-3-carboxy-1,2,3,4- Nep-methyltetrahydroharman (2) tetrahydrocarboline (1) N(CH3); HO hordenine (7) 4-hydroxyhygrinic acid (7) о М CH; N O HN julocrotine (6) о 2,4-dimethoxy-', Y-dimethyl- allyl-E-cinnamoylpiperidide (5) " N Ся м2 9 М vasicine (3) tetramethylpyrazine (9) < N G NS N N N о м о n-C13H¢ n-C43H, glochidine (12) glochidicine (11) т^. the bad structures found in Euph allo eae g ак cera chemical type. The number of each structure and 31-32 are alkaloids, 30. аву: ide re diterpenes, 5 i 68- sh pags RA tannins, 71-81 чү блар 82 is an amine, 83 is a coumarin, a sesquiterpene, 89 are of other Meque s rigin. 29 35-67 are da a 84-87 are flavo heus 88 394 Annals of the Missouri Botanical Garden De d 2 eel NHCO(CHj;CO(CHj;CH,; N?-oxodecanoylhistamine (10) alchornine (14) Te alchornidine (13) о B о М аА (8) СН; | 4 Y c3 N HN astrocasine (4) securinine (24) norsecurinine (23) allosecurinine (22) virosecurinine (25) FIGURE. Continued. Volume 81, Number 2 Seigler 1994 Phytochemistry and Systematics of Euphorbiaceae (+)-crotonosine (19) trewiasine (31) N(CH3)2 hymenocardine (32) FIGURE. Continued. 396 Annals of the Missouri Botanical Garden OH OH CN o CN ; HO 9 RES HO ГА HO O HO о OH OH А он H^! CN linamarin (35) a, (R)-lotaustralin (36) (0) о HO HO OH OH (R)-taxiphyllin (34) on O H CN HO о OH O O HO HO OH OH linustatin (37) — (0) о 08 о но HC Ln OH OH H OCH, HO OH " CN 24 9 d T CN OH о М он acalyphin (30) triglochinin (33) CH; OCH a c CN o CN = NS CN Tol N о М о ù | [9] М CH, CH; P CH; ricinine (26) ricinidine (28) mallorepine (29) nudiflorine (27) FIGURE. Continued. Volume 81, Number 2 Seigler 397 1994 Phytochemistry and Systematics of Euphorbiaceae OH 18-hydroxy-6-Z-geranylgeranyiol (38) casbene (41) cembrene (39) 4,8,13-duvatrien-1,3-diol (40) jatrophone (42) 13,15-seco-4,12-did lathyrol (44) (a кааныр они (43) CH,0H HO ae CH;OH Hon \сн,он 2 1 (4 daphnetoxin (48) ingenol (47) FIGURE. Continued. 398 Annals of the Missouri Botanical Garden ent-36-hydroxybeyer- 15(16)-ene-2,12-dione (50) isocrotocaudin (51) teucrin (52) 0 о он woo HO A o ellagic acid (68) mallotusin acid (70) geraniin (69) FIGURE. Continued. Volume 81, Number 2 Seigler 399 1994 Phytochemistry and Systematics of Euphorbiaceae OH O vitexin (85) heveaflavone (87) Н OH s отеу юв он glucoputranjavin (53) N — 050,- OH Ae, OH glucocleomin N — OS0;- glucocochlearin (54) FIGURE. Continued. 400 Annals of the Missouri Botanical Garden CH3(CH2)4HC=CHHC=CHCO2H — 2E,4Z-decadienoic acid (55) CH3(CH2)g6HC=CHHC=CHCO2H 2E,4Z-dodecadienoic acid (56) dI T i from Croton tiglium oil (57) Cu.(CH.) 1 t 1 {4 (ER) v 2 сыы woz сазы?) U nno nan 27 E CH3(CH2)5CHCH2CH=CH(CH2)7CO2H ricinoleic acid (60) OH он CH3(CH2)4CH(CH2)19CO2H 12-hydroxyheptadecanoic acid (61) E E Z HO(CH2)4HC—CI 1H HH I(CH2)7CO2H a-kamolenic acid (62) EN 2 CH3(CH2)sCH~CHCH2HC=CH(CH2)9CO2H (+)-2-14,15-ероху-2-11-еісоѕепоіс acid (63) ZN 7 CH3(CH24CH-CHCH2CH-CH(CH2)7CO3H vernolic acid (64) cus ciii pod cw 10,13-epoxy-11-methyloctadeca-10,12-dienoic acid (65) Lice CH3(CH2)7CH-CH(CH)7CH20Ac erythro-9,10-dihydroxy- 1-octadecany bu on аага гатага: 3 e 1 tate (67) Y OH ш! Г = | | 0 14. dihvd =ч » 2. a4 49. зоо СО OH OH , J J эчү FIGURE. Continued. Volume 81, Number 2 Seigler 401 1994 Phytochemistry and Systematics of Euphorbiaceae " HO tirucallol (75) euphol (74) germanicol (77) HO euphorbol (76) phyllanthol (79) d bauerenol (78) HO cycloartenol (73) HO В-атугіп (71) lupeol (72) FIGURE. Continued. Recent Issues of the Annals of the Missouri Botanical Garden Phylogeny of Asteridae Annals of the Mi i Botanical Garden 79(2) (Spring 1992). The HEA sister tad (or groups), and гед of уюш ы of the Asteridae are discussed in the symposium “Phylogeny of organized by Larry Hufford of the University of Minnesota. The ARR was [jointly by the American vu of Plant Taxonomists and the Botanical Society of America and was cm at their annual meeting, 7 Aug. 1990, in Richmond, Virginia. 238 pages. $27.00, plus postag Knowledge Brokering: The Mechanics of ние Annals of the Missouri Botanical Garden 80(2) (Spring 1993). This symposium, the 38th Annual Systematics Symposium of the Missouri Botanical Garden (held 4—5 Oct. 1991, in St. Louis, sith discusses the value and future re of e diverse biological information into ne broader syntheses, taking advantage of с er-based data-management systems. 41 pages ед issue 230 pages). $35.00, plus postage. rbcL Sequence Data and Phylogenetic Reconstruction in Seed Plants Annals of the Missouri Bota wae gees 80(3) (Summer 1993). Fourteen papers plus an appendix containing voucher and sequenc a. This collection of papers, organized by Douglas E. Soltis and Mark W. Chase, Pues RE xd sed pth molecular phylogenetic analysis of seed ante in pu ànd angiosperms in particular. гова throughout ће world involved in rbcL en their unpublished sequences for a broad phylogenetic analysis. 235 Карек. $40. 00, plus postag To place an order, send check or money order i in U. 5. . funds, payable through a US. hank, jun rid orders also 755 US Жагы add $3.00 nd $. 75 va bs oe book. Orders should be prepaid; a $1.00 f SE "n to orders Eid invoices. No shi e made until gemens, is Hover Mail or fax (for кта rens form with Your check or money nar ge y able to Missouri Botanical Ga rden, P, Box 29 Eleven, Missouri Botanical Garden Please send ———— copy(ies) of Annals 79(2) А ` es O 63166-0299, U.S.A. copylies) of Annals 80(2) a IE jonas ү copy(ies) of Annals 80(3) Send book(s) to: 0 Payment enclosed. И TTS ETL АНЫН 0 ind Ue. ү" :00 fee will be . Name WO LET Quid CES LIE AAA ЧР ب‎ н == Û Mastercard d Айе» | 3 9 404 aiu АШИР P Oa EER RE АА Н ج و‎ E I Name as it appears on card ; Postal Code · Сошигу sio Gn g Exp. date Phone number (days) Prices ARE SUBJECT To CHANGE WrrHoUT NOTICE — CONTENTS Systematics of the Euphorbiaceae (Continued from Volume 81, Number 1) The Contribution of eet to the Systematics of the Euphorbiaceae R. N. Kapil & A. hatnagar 145 Leguminlike Proteins and the озеш of the Euphorbiaceae Uwe Jensen, Ina Vogel-Bauer & Marei Nitschke .. 160 Systematic Anatomy of Euphorbi ceae Subfamily Oldfieldioideae. I. Overview W. John Hayden 180 Е Implications of Pollen Ultrastructure in the Oldfieldioideae (Euphorbi- ae) Geoffrey A. Levin & Michael G. Simpson 203 Үзөн Relationships of кич, апа M EU I (Euphorbi- aceae) Geoffrey A. Levin & Michael G. Simp 239 A Palynological Study of Crotonoideae (Euphorbi ) Joan W. Nowicke 245 Laticifers in Crotonoideae (Euphorbiaceae) Homology and Evolution Paula Rudall _— 270 The Relationships of the Euphorbieae (Euphorbi ) M. G. Gilbert . 283 Patterns of Diversification and Evolution in ен odontadenius (Euphorbi- aceae) aicour, L. Rossignol, M. Rossignol & C. Gaisne .... 289 ie Evolution of Dalech ampia a es зо from Phylogeny, Bioge- aphy, and Comparative Ec ott Armbruster .— 302 SN Чыч and Phylogeny of os dd pde Vento Lynn s С, deed ‚ T, + Гот. z d Phyl ERIE Analyses Meet SRE & Bart күч ма dere n ERU hi suc. 349 A Preliminary Classification of Euphorbia Subgenus Euphorbia Susan Carter .. 368 i Phytochemistry and Systematics of the Euphorbiaceae David S. Seigler m- 380 Cover illustration. Petrea sulphurea M. J. Jansen-Jacobs, by Phyllis Bick. Annals of the Missouri Botanical Garden tens VY umber Volume ge Number 3 Annals of th Summer 1994 spinach са Garden The Annals, published quarterly, contains papers, primarily in systematic botany, con- tributed from the Missouri Botanical Garden, St. Louis. Papers originating outside the Garden will also be accepted. Authors should write the Managing Editor for information concerning arrangements for publishing in the ANNALS. 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Four urnal nep is included in the tte price of the ANN issues per vo The mission of the Missouri Botanical Garden is t 5 E 1 jo h 1 “у, o LI environment, in order to preser d enrich life. © Missouri Botanical Garden 1994 NNALS OF THE MISSOURI BOTANICAL a aes 0026-6493) is published quarterly by Missouri on need: 2345 Tower Grove Av 3110 post! Eleven, T6. Box 299, St. elie "MO MO 631600299 ence of Pape? ©) Thi cm r 3 Laien acd q Ф АМС МСГ 730 Arthur Cronquist Volume 81 Number 3 1994, Annals of the Missouri Botanical Garden WZ ORIGIN AND RELATIONSHIPS OF THE MAJOR PLANT GROUPS: DEDICATION AND INTRODUCTION: #1550, Ri P. Mick Richardson? ' POTN Igy AUG ; 0; 994 FARD Ey USRAR, Art Cronquist surely knew more M the origin and relationships of plant groups than has ever you may be surprised E. E 1987), h ію uttered ibo words, “We can use- y think in terms of apomo Pin relationships d if the oe ts во producing the new ides, кке nim ur s deserved to be lis о. t uld have enjoyed the symposium Te Symposium showed how far we po ad- A vanced Since dte Parenti published = appe plants based upon Hennigian principles (Parenti, 980). I Kenrick & Crane (1991), Kenr ships of th h i those put forward by Jim Doyle and his col- mposium of the Missouri the Missouri leri thank Mike D. ti ' This к: "апіса] Gar me Чр апі eoi и. = Major Plant ане The symposium was held 2-3 Oct. 1992 at a Garden in in St. Lou ssouri, U.S.A. "he National Science Foundation under pe number BSR89-18138. I y Kluska, Arthur ан lo jenii rving right чеч апа secretary, for. diti me with the choice of the пои ssouri Botanical Garden, Р.О. Box 299, St. Louis, Missouri 63166, U.S.A. ANN. Missouri Bor. GARD. 81: 403-404. 1994. Annals али punter: Garden laborators. Victor Albert, Mark Chase, and their of the enormous rne ле is сы УЫ presi a continued move ith th + rie у cf rt. Ж: аха. un the а aphyle tic not a P е was certainl + r phytes, gymnosperms," an icots," it em- ph ae ТРА = Аш : Ч sh 2 tiec analysis of data seat in the auditorium for the duration x the con- ference and, with such intense interest in the sub- ject, the conference must surely be judged "d timely and successful LITERATURE CITED — A. 1987. A eo, critique of cladism. Rev. (Lancaster) 53: 1- 0. Present дулат and future trends in systematice. Giorn. Bot. Ital. 124: fas bstract]. Kenrick, P. Alternation of vedi and plants: New phylogenetic e pa dum, res Bi- ological Reviews (in ZI E. 1991. Water-conducting 4 arly fossil land plants: Implications for the ear pies of tracheophytes. Bot. Gaz. Cobol 152: 335-356 PARENTI, L. 1980. A phylogenetic sn of the land plants. Biol. J. Linn. Soc. 13: d Early evolution ES rt origin o d IN, M. L. 1991 eukaryotes. Current Op opment 1: 457-463. YOUNG, D. A. & P. M. RICHARDSON. 1982. А phylo- genetic ате of extant seed plants: The a to Taxon 31: 250-254. PROGRESS AND PROSPECTS IN RECONSTRUCTING PLANT PHYLOGENY’! Michael J. Donoghue? ABSTRACT Phylogeny reconstruction has become respectable science over the last few decades, and trees are магн recognition of s streptophytes, stomatophytes, anthophytes, min) Nevertheless, only a s To ve the toughest problems, especially t those a e ancient, be combined, i Fur ther addressed and some о: of these have resisted solution. ata will need to ith m ll number of pro oblem s have been eon in d ance mngreased use racte: of phylog enies by ecologists, molecular kou and others, which f жазасына taxa. We. can п expect 1 Es in £ M with population biologists promise to be To productive, since there are obvious mutual concerns centered on the analysis of gene trees and reticulation Over twenty- -five years is elapsed since the publication of Willi Henni genetic Sys- n api e, and the — itself docume es signific өт of ^m I challenging LA iei: questions. ТЇ The in general t 3 ا‎ ere е things stand. з plylogenei hie ad a substantial i impac understanding of plant phylogeny, and cibos m we go from here? PROGRESS Although many of the ideas underlying phylo- *netic anal be traced series of devel- °pments in iw. logic of phylogeny reconstruction (e.g., Hennig, 1966; Farris, 1983; see е Sober, 1988; Swofford & Olsen, 1990), to the са of ithms to i i de shies eas have had a signi impact on plant systematics, it is not entirely clear how best to measure the progress that has been made. There are, however, d indicators that bear consideration LEVEL OF ACTIVITY e of pr ogress is simply the rate of use no one trees (Sanderson et al., 199 number of studies Тои a tree а day) is surely ‘Lam grateful to Bruce Baldwin, Peter Aon Jim Devin David Hillis, Toby Kellogg, David Maddison, Wayne rong n, рек Оше ad, Mike Sanderson, and Liz » as well as to Mick Richardson for by the NSF (BSR -8822658), the University of Arizona, and a Instituti eaten me E participate in Aes sym access to unpublished has been | Mellon аа pedes уен the Smithsonia immer for helpful т үт Us Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, Massachusetts 02138, ANN. Missouni Bor. GARD. 81: 405-418. 1994. 406 Annals Фоо а Garden over 471 (53%) of the een studies ka some form of molecular ev; e (inclu zyme data), with the УРИ of such pak increasing from 51% to 56% over the three-year GREEN PLANTS (199) bend, We identified 312 studies of green plant groups oe 27% of the pened) namely organi maii th chlo- Ьа rae d sta Зыр of dr peut and the land "uii (San- an- 882 PHYLOGENETIC STUDIES (1989-91) derson et al., 1993). Hosen, 113 (36%) of these employed phenetic techniques, a greater Pais: of phenetic studies than we found in any her major group of otes. А breakdown of " shined эке) the remaining 199 numerical phylogenape ИШ ANGIOSPERMS ther green plants (10) (168) are 4) other vascular plants (5) is sho own in Figure 1. The vast majority (168, ыт еке) f these were of angiosperm groups. Within а : sperms, Asteraceae (30 phylogenetic ms nd Faba (15 studies) received t t at- A o ORGEN PEANT GUESSED tention, while other large families, as Orchi- Asteraceae (30) de nisi (4 اا‎ are underrepresented. Ти ‘thin orasses. Det Шш Роасеае (14) angiosperm studies, with 14 of tł g SABA orchidaceae (4) To some extent these numbers reflect the number 4 5 of species per group, but historical factors are i 2 u e Asie b e ising г other monocots (31) were stimulat y the other dicots (74) d leagues (see references in Palmer 2 168 ANGIOSPERM STUDIES (1989-91) have re FIGURE l. Pie ch h Figure 1 also show s Hes ort E cud "Eur мга NA EN RE ceived e little attention, — ang rical i i wipe pub- : andi TR — те 1989 and 1991 (compiled fro n updated g thei portance in un under: ЕР a of abe а b radon et al, netic relationships among major line CERE edd 19 93) tudies of green plants (chlorophytes; with For example, we found only five phylogenetic Saa Eo h darchsorge)acou for ок Z3% Ше уш of vaseular plants other than sed plants dir al. - : base i тк 8 th en тай he three-year period, despite their dub ий tudies are of angiosperm groups; the remainder аге of logenetic significance and th pi : wet “о ed plants” (specifically conifers, cycads), “other ^ pteridologists in phylo tic questions (6.8, vascular plants" (ferns, lycopsids), “other embryophytes”” tudies, ding fossi Lye « ao flea nae , ү йд, : ” ап (mosses, liver , "other green plants (various green egies астме Сойһар ee oe mos Вион or i пој ded representatives of “all major lines. ttom: Of 16 9 (ca. 29%) are of ran- ocot groups; Asteraceae and Fabaceae account for Pits to a fect for example, whether ben E ca. 38% of the “dicot” studies iate fern groups are more clos ated to а tosporangiate ferns or perhaps 8 : groups (and hence s nts) Fortunate de an underestimate inasmuch as we did not survey terophylls and lycophytes (including йо) 1992; every journal containing phy be st ben. and receiving more attention (e.g., Bateman man et al., we were unable to include analyses ished in Gensel, 199 stud- books. Nevertheless, our study probably кер Although the absolute number of TT ch a reasonably accurate pic М, what exists in the ies of green plant гүп т increased slightly three- literature. Of the total, 882 (77%) of m е year, the percentage of suc studies over the being loy “cladistic” method (inclu par ear u d around 50%, th ee ony and some distance methods, but not sm based on morphology and/or se secondary ien 4 sim nd netic clustering methods such as UPGMA), a Overall, about half of the molecular $ Volume 81, Number 3 1994 Donoghue 407 Reconstructing Plant Phylogeny based on codi ы. E especially st DNA in chlorop “Те Viii nt studies based on м sequences тй іѕеп dramati- cally since 1991 (e.g., studies of чы chloroplast аре, but mee ps sum TE ig 1989-1991 period. Cites alf of the s se- quence studies Aide мед Aem (see Zim- mer et al., 1989; Hamby & Zimmer, 1992; Wa ters et al., 1992), and in vh near поре, especially at lower taxonomic level u ба of the internal ng sp gions (e.g., Baldwin, such sequ rgely overcome, and I that they will play an increasingly important role. Phylogenetic studies دن‎ include fossils are still sias send hes eir nom MEER PD е et ing (e.g., Golenberg et al., 1990; Soltis et E mr their apes n Eo on developin 18 references in Kenrick & Crane, 1991), E explent sale aa af the е пирог rtance of foss sils B шпоп 01 vascular plant H А . th Agai n, t molecular evidence (e.g., Palmer et ali; 1988; Manhart & Palmer, 1990; Raubeson & Jansen, 1992). orphological phylogenetic analyses of seed 85 et al., 1994, this issue; Nixon et al., 1994, this mtr dg while hot a agreeing in in detail, have consis- most Pigs related to angiosperms (Fig. 2). This pla de i is also supported though not very wrongly) М al., 1993; m & Zimmer, 1992; but see Troiteky et a ; Hasebe et al., 1992). An anthophyt TT consisting of these two living roup s s Bennetittales and Pentoxylon, is a regular feature of the studies that have included ils. Furt ore, with the exception of some атар! o 3 £ DH Я : Jo sci t is abil: DNA - x SUCCESS STORIES While it is obvious that botanists have actively mI in phylogenetic Caa it is A e (ч. п. аен in several cases signe pr progress оез appear to have been made. Some of these Successes are highlighted in 27 затне g (Fig 2). posium papers, so only a few are noted here The idis by Mishler and colleagues (see Mish- s e Oplasts, especially je лыд Ус such as C е е mole -— evi- 994). Kenrick & Crane (1991) have made great к и ш establishing relati rl vascular plants, suggestin, udies of tracheids, coupl & overies of Remy and colleagues gametophyte morphology (e.g., Remy, 1982; have бсш that Gnetales аге mono ; d Welwitschia more closely related to one another than either dra this molecular data are especially compelling (Doyle et it. 1994). There has Es been Paste оп the pigs te of angiosperm n in lingering sea ce to the gre that veia are т (е. z Krassilov, 1991). Within angio- sperms there have also been promising r esults n. Even from this short list it is clear that “success” has not been tied to the use of any one type of evidence. In fect, the 2. impression of success ' of all of th orphology and mole — of veracity,” Hennig BEG їп 66) provided a a gy wh hich reproduced i in Figure 3. In this example, the pro- all of the evidence is likened toa коки ‘attempting to assemble the torn eces of a map so as to bring together all of the 408 Annals of the Missouri Botanical Garden d e $ © © ج‎ д” Sy & 45 S, © CASA AA 2 5, © (0 & $ Ls A A © k4 SESSO ree SRLS G c) Ко FLD о о SY ЖКО МУКУР ES, SEES So Ga d ALD A RT OFS S P S SA LEO. FE SLE AA УО Nd. SG SLES SESE SSE СО AUS STARE SOLIS SPIES SEE EE CHS EE CSS ELS SL SEE Сг OOS FLL OPEL OC OC LOL LLNS MS SD ANGIOSPERMS (flowering plants) ANTHOPHYTES SPERMATOPHYTES (seed plants) LIGNOPHYTES TRACHEOPHYTES (vascular plants) POLYSPORANGIOPHYTES STOMATOPHYTES EMBRYOPHYTES (land plants) STREPTOPHYTES CHLOROPHYTES (green plants) Ficu E 2. А : f 1 ( IDA | "n 1 1 Lnd n2 sa wb showing major dale M seem reasonably well supported and areas of continuing uncertainty (unresolved polychotomies). ped in^ are omitted, and resolution is simplified to highlight familiar clades (see O'Hara, 1992). Only three лай. branches i k which are probably paraphyletic (quotation marks). Some of the best — universally have long been recognized (e.g., land plants, seed plants, flowering plants), though these have not been : ently accepted; others have been re-circumscribed (e.g., tracheophytes) or named (e.g., polys} ngiophytes) iles (Kenrick & Crane, 1991); several others have not yet been named (e.g., Charales + Choleochaetales-- Embry elie R in £ th 1 r ү rs Pgh | " 1: у П f recent phyloge! ше VIII yiy о Recognition of th studies. Note that m ty a number (e.g.. jd d monophyletic, some traditional groups (e.g., “green algae," “bryophytes,” “gymnosperms )aren and are not shown. Volume 81, Number 3 1994 Donoghue Reconstructing Plant Phylogeny roads, rivers, and other landmarks. The corre- sponding operation in phylogenetic systematics is assessing, as Hennig (1966: 130) put it, “whether the differently determined views concerni phylogenetic relationships o y consistent explanation for any one road, or bit of evidence, might turn highly unparsimonious whe tures, howe they the outset, п other fea- may have seemed at are taken into consideration. OUTSTANDING PHYLOGENETIC PROBLEMS For each success story there are, of course, many other I ы. 1 1 h Adi *| or for which a satisfying result has not yet been obtained. Thus, as also shown in Figure 2, rela- ми among the major extant lines of seed ginkgos, conifers, anthophytes) have DER ae 1 і (Doyle & Don- © oghue, 1992; Doyle в мч 1 994). Lik Position of the root of angi CRT agn or amo mained unsettled (but see Doyle et al., 1994). I oth cases, particular dat id ее support for one hypothesis over another ani / or there are conflicts among the results based on different data sets. Even in da 07. реч ample, there is little evidence that ot of angiosperms is among monocots, and + а 1 1 1 l1 е v. 1 nd it the root I ER support rooting the tree in the vicinity of is ranthaceae. Furthermore, it is worth noting Nes conflicts appear to be as significant among а molecular trees (contrast Martin & Dowd, en. E & Zimmer, 1992; Chase et al., a. and among different morphological results a st Dahlgren & Bremer, 1985; Donoghue & yle, 1989: Loconte & St 19 "m , 1991; Taylor ia ickey, 1992; Nixon et al., 1994), as they are ween molecular an gical results. c ` austed. Much of the information present In standard M of 1 E Vi» r 9 Thies been incorporated in phylogenetic analyses. ы п and should , but will require a bn Сн of homology hypotheses (Don- anderson, 1994). Furthermore, new ap- E ONE T: FIGURE 3. panel) from Hennig (1966), illustrating the "criterion of Hennig's ori e se. The remainin| remain isolated. The joining of the map fra wrong." “Figure 38. Criterion of veracity. If queis "n P үр fragments is in the re- a, a’, and a” are interpreted as adjacent sections of a stream, the other elements of the map also join to form a sensible illustration. The joining of the map fragments is correct." proaches to morphology, such as detailed studies of flower development and function (e.g., Endress, 1987; Tucker, 1988; Erbar, 1991; Willi ў iams et provide valuable phyloge- ti lyses that m rt al., 1993), continue to netic characters. Comp f 4 oH ү underlying flower development (e.g., Coen & Mey- erowitz, 1991) promise to be especially useful in elucidating the homology of the structures that i ү p a de Lalaoical nh T 4 logenetic research. TV T, * D nce also с bined and analyzed simultaneously (Kluge, 1989; Barrett et al., 1991; Donoghue & Sanderson, 1992; but see Swofford, 1991; Bull et al., 1993; de Queiroz, 1993). Here the hope is that weak signals Annals of the Missouri Botanical Garden presak in уса oe data sets might com- single aped гей. Obviously, this need not E tus outcome, and pt ovide ut it behooves us to explore such possibilities in particular cases. In th f the osperm root, the NATUS analysis of Doyle et al. (1994) avors a placement among paleoherbs more con nella than any single data set. This is not simply f 1 } 4 1 1 1 4 Y dou s. a2 cause T о r о some elements іп the combined trees аг е may best Donoghue & enu 1992; Doyle ons that w in en" the lengths of a айаш to extant taxa, iereby xau chance SE pego. (the “‘long-branch ОП tion" problem; Felsenstein, 1978). Alre eady there are examples of this : (e.g., “progymno- sperms" relative to seed plants), and the e m of recent paleobotanical dece s (e.g., Cran al., ГР де: A et fin 1991; Gornet, Тн) could g giosp nra & TT 1993). METHODOLOGICAL ISSUES Solving difficult кон problems will re- ‚ but it will also is ез eepecially critical. in view of the hens lim- le com orithms andling podeis of — (rotor d & Oben mes Malis 1991; Maddison et al., 992; Pe puri et se 1992). “Riagh. there i is some nm in the Cha ase et al. (1993) debian of. 499. ei er all (or even any) s trees bes have been recovered, and it is practically impossible to eval- uate alternative hypotheses critically or to explore the robustness of the result or а 7 an «ти (Bremer, 1988; 92). Of cou the а. apply to ine morphologie studies (e.g., Hufford, 1992; Kellogg & Wat 1993). ja view of vi problems, d involving large numbers of taxa must employ carefully de- — (and explicit) search strategies to discover “islands” es (Maddi- 1: ate however, ent iy 8 inadvertent choice of ed well within e longed to one specia which happened to have converged on another terminal taxon in the analysis. А second option is the use of a ee ге Swofford & Maddiso tunately, it may be sufficient to ha e prior hypothesis of relationships, as the sta n taxa nested well within the clade in — З be pig dis *screened-off" from having d d fect on the ssment of basal ond dison et al., 084 ). use А placeholders will mean that more Volume 81, Number 3 1994 Donoghue 411 Reconstructing Plant Phylogeny 1, b Ре 11 а. Ап ате approach, де) by пои t Ds is (1991), i be of lasting value only if all of the evidence un- derlying the results i is made available for нп Otherwise, i into monomorphic terminal units. “Th no av will result in para- or polyphyletic terminal units in many cases. For example, if d E were ai d into two termin a bas e and absence of vessels it is 0 ph tha neither OE be a clad ame would * and absence sels an active cambium, or different types of embryo sac development. Inclusion of ШОк АСД terminal taxa in an ап ires can result даа EF Ab and eer evolution, at least if taxa are LA preted у, namely as representing real Ў branches a a phylogenetic tree. the end, th " ning cautiously on е results of previous TERR "The latter re s that the assumptions used in one analysis be vig- oe tested in other, independent analyses, but 900) oth are ب‎ 4 possible ү 8. таги et among alternative xA Minn or to learn anything at all from experien PROSPECTS What can we look forward to over the next few years? Here I will briefly highlight what I believe to be several i мр tant t trends and эсе r likely con- PoE At the of each o i ore е shift to P iat b: B (1988) his called T thinking”: underst tanding diversity and observed Чщегипоез as having resulted "im That the shift avay from E i linear view of evolution is far fr ed nguage still so often used to шы: evolu utionary history, such as “lower” and “higher” in reference to taxa (O'Hara, 1992) a however, tree-t ing is likely figure more A ominen Чу 0 in e curriculum, dison & Maddis n, 1992) making it easy and fun to explore the ейи of evolutionary trees. A PHYLOGENETIC SYSTEM Although major noe will е p time, I suspect that bota. ts will continue to mo toward a traly | phylogenetic sie p this ы mean at taxa best estimates of oe and will be defined i in th h ct ES S orphi ic, as can be js one in PAUP eru 5 апа MacClade (Maddison & Mad- 92). In the case of multistate characters in as M nucleotides d = pár articular site) this need 'si - Although coding a taxon as unknown a ary, especially rporate incomplete fossils | (Maddison & Mad. dison, 1992), ven in cases that continue to resist resolution, and analyses { hing de this еца е implications of alternative hypothese ‘elles ourse, only to the extent that sources of аш tion (e. especially voucher "spec сйшепв) | аге ste ully doc umented, and onl Ren rns the delimitation of taxa and characters Clearly spelled out. Phylogenetic research с s used to a them (Hennig, 1966; de Queiroz, 1988, 1992; eon & Cantino, 1988; de Queiroz Gauthier, 1992). I rimi terms this will m: ting some ere letic ics V excellent example is половен ннер on Asteraceae, pren inating 1992) in several new names to в reflect conclusions that are well supported by ifferent sources of evidence (e.g., Barnadesioideae as the sister group of the rest of the family). Many more changes of this sort will be made as our understanding of phylogeny ini s. In par- ticular, it is clear Шага some traditional soris yl ith one or more acters g phyletic families adn ni Cantino (199 2 2) has carefully тыы the igin of several lines of Lamiacea e from within Arl benaceae. Judd et al. (1994) mere documented es of other решу vend related £ families. In particular, nested within a para- Annals мня ps dmm Garden phyletic Apocynaceae, Apiaceae within Araliaceae, Brassicaceae within Capparaceae, Valerianaceae plus Dipsacaceae within Caprifoliaceae, Urticace: wit oraceae, and Aceraceae ippocas- This, we hope, will help counteract a bias on the the rank of ma ainly temperate, mainly herbaceous groups, leaving behind a residue of woody tropical plants with which they are less familiar er major changes in some cases, this sh hardl be viewed with dismay. Instead, such changes are a ete sign that we ar ress in understanding phylogeny. Furthermore, although ere g Moria US such ee are 8 likel afi occu useful in elping us understand cade and ecological processes, biogeography, and so on oe: 1966). me oreover, recognition of mono- © diversity 1 and ‘for students to learn about it. It 8 guelia (Gane 1986) relate to modern angio- 5р rem sditüón to better phylogenies, the develop- quires attention vident wy ee to treat taxa assigned the same taxon rank th "aa d based approaches « sei — for — Ü to sys- tematic a diversification (Doyle & Donoghue, 1993). PHYLOGENIES IN USE The of phylogenies by ecologists, paleon- Жыны Морко. molecular biologists, соп- servationists, and nea will ee continue to increase (e.g., Brooks & McLennan, 1991; Harvey & Pagel, 1991). dà the case of ‘iti sperms, for ee An the сходен ч trees has already had 1989), d dioecy PM oe ume double fertiliza- noghue & Scheiner, 1992), myr, 1992), and self-incompatibilit r et PI 199 other ideas on the tion E mor ological evolution (discussed, for example, by Stebbins, 4; Carlquist, 1975; Cronquist, 1988; Takhtajan, 1991) will soon be tested in a - genetic context. Similarly, phylogenies will have an increasing impact on 1 ding of lecular evolution. For instance look f. d of “reciprocal illumination” to a fruitful period between studies of bp phylogeny and greco: cem analyses the mechanisms underlying flower dev Jude (Coen & мое 1991). use of phylogenies іп other es will force phylogenetic systematists to attend t to ү? ben ra reliability: Quite natur ally, tl ER DR to que on phylo nies will bali yabê they are. A variety of methods have 91; Huelsenbe Faith, 1991; i et a 92) and there is a growing (mostly critical) wes on the logic d behavior of such met gs — the meantime, users “ trees should be w of uncertainties and e oura to tak count ambi AHORA) of the phylogeny addiso and T character tenir (e-g.. al., 1992; Maddiso: Weller et al., in pr s in assessing confidence will require tter understanding os ssible sources of wis n (e.g. „ 1994). In particular, w e need to ы cisely how to ish real signal from TR results. Despite their limitations (Hillis et al., 1 " computer s ons will con to be a € xplore a very wide values (e.g., Lanyon, 1988; Hue vine Hills, 1993; Kim et al., 1993). Of course, suc studies Volume 81, Number 3 1994 Donoghue 413 Reconstructing Plant Phylogeny leave open the critical question of exactly where in E space сиу рагісшаг problem is sit- to for malize th f ЖЕП EU th f ical variables Work mh known сее (e.g., Atchley & Fitch, 1991; Hillis et , 1992) and experimen- tation with real data sets 7 g., Smith, 1989; Don- oghue et al., 1989; Allar Miyamoto, 1992) р Mix be useful. An approach ihai e more as benchmarks to help determine whether mist se- ar ample, Dav : id Maddison and I (unpublished oy are Ж ишы approach to over the to the E branch in лш trees based оп outgroup sequences effect they no longer retain phylogenetic infor- campa Our pr ary studies show th t dom" se (genera under « he re so the behavior of real out group = uences vine de- viate from any of the random sequences we have oh apa big a that the real sequences n = ex “et are HERFS as de more RE sta- oo ا‎ in TE also., rines "еор bett about t them Es to e ONE users. We ne idl in connection, for example, 1991). M ation efforts (e.g., Vane- Wright et - Moreover, accessibility of the full range H tudies surely would facilitate the ма for pe patterns; for —— patterns "à els of hom g., Sand Dono- M ue, 1989), or in tree shape (e.g., ا‎ 1983; ds & Slo digo 1991, w i the search for sn patterns oe dieting by all g (at least sa y) ready access to every tree dint taxa pres- ent in particular areas of endemism (e.g., Nelson & Platnick, 1981; Page, 1991). PHYLOGENIES AND POPULATION BIOLOGY Systematic i is ecient piittreyed as disp nea tly E it is К ipie that this gap shoul ain- tained (e e.g., Nixon & Wheeler, 1990). This, I believe, is a mistake. On the рири interactions with population ES should be promoted ac- tively. S 9 | | frui: it fall а in view 0 obvious mutual concerns centered du F coalescence theory (Pamilo & Nei, 1988; Avise, ; Takahata, Med Hudson, arg Dove: vii Page, 1993a; Baum & Sha press; arana in press). application of E ic methods to what have tradition ally been po NS -level aene om ae results already. For aene не devised a geet appr iz oint. It is Pip less obvious how ts, especially those of us primarily concerned wi er аа events, stand to benefit from interacting wi ulation biologists. талы oe одане) t. One such e con ed evolution and the dynamics underlying the es establishment of атыи genes, issues of obvious stematists (e.g., Zimmer et а 1980; арна 1988; тея & Doyle, Ве Better. understanding of mos eea ed E: ill all 1 cases. ө: For heels comparison of denies Annals of the Missouri Botanical Garden ракы on paralogous panes ага ae it possible h this approac allowed Iwabe et al. (1989) to root th tree P life, e the lack of obvious outgro oc to сша treos based on both nu- ], & A vica. Apenucu м to identify кее end specify their нйн e (e.g., Soltis et al., Rieseberg & runsfeld, 1992). ANGE line i have occa- sionally been discussed vé не торе ers in Platnick & Funk, 1983; F 1985), detailed rts (Page, рае case of eee (Hein, 1990), itr transfer of DNA (Val- dez & Piñero, 1992), an n the fusion of entire biotas in ве P Pants 1988). The de- velopment of such a theory depends on breaking at barriers have tended to separate population biologists from systematists. SUMMARY It seems clear that headway has been made in e oblems of taxon sampling sessing the reliability of phy- genet hypotheses. Eventually, we ш see the logenetic t theory, phates reticulation of all and np e boundaries of tradi- id disciplines LITERATURE CITED ALBERT, V. A., A. BACKLUND, К. BRE à J. R. MANHART, ISHLER & К. C. NIXON. 1994, mn E and rbcL evidence з "UN Lo phylogeny. Ann. Missouri Bot. 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Elsevier pu ear N, S. M. PM . W. КАМ вт а » id is ication an Кар f he p oglobin . Proc 1980. А genes uet for the д chains o Natl. Acad. Sci. USA 77: 2158 8-216: INTEGRATION OF James A. Doyle,? И оса AND Michael J. бело OMAL RNA DATA ON and Elizabeth A. XN n E ORIGIN OF ANGIOSPERMS: ABSTRACT whereas f among paleoherbs, with eudicots and woody m: durus forming a clade. Experiments with a revised seed plant ead de data set raise further questions: when ang giospel rms are scored like different angiosperm subgroups, t ШИ, rRNA data | are жен А раво, or rather complementary, with inconsistenci being a funct f of better resoluti f the we experimente and rRNA data sets i six X exant “gymnosperm” and 12 нй дыз taxa. Both analyses again associate angiosperms and osa The angiosperms, bet ut trees er next to Magnoliales ate only one step less parsimonious. . As i рг ious studies, the rRNA ore he sis f th RNA show strong ерон for Bs mon rari of angiosperms and Gnetales en their sister qi p "ааба but гөш Support for groupings within angiosperms. However, опе or another gro up of paleoherbs is basal in most bootstrap trees. A combined analysis favors a мен rooting, but other relationships agree with ds morpho! a results; ш particular, M form a clade. The conclusion that Gnetales are the closest living тше of angiosperms permits a e e range of morphological wieso depending on the arrangement т Bus ates oups. Disc Mid n he ssils on the long К leading to angiosperms, methods of factoring out artifacts ios molecular е жып of floral morphogenesis in angiosperms and Gnetales may be required E: Me Aci in resina origin of angiosperms. Recent cladistic analyses of morphological and r initial sear big ruo analysis of seed plants ribosomal RNA data have led to apparently con- ги & ر‎ ны 1986), which included both flicting results on the position of angiosperms among living and fossil t КЕ. treated angiosperms аз а seed plants, basal relationships iino: Vici is single ie modeled on Magnoliales and Winter- and resulting scenarios for t gin angio- aceae, A the consensus at ae time. This sperms (Crane, 1985; Doyle "i "s uis e, 1986, study was designed especially to st the previous 1992; Donoghue & Doyle, 1989a; Zimmer et al., analysis = Crane (1985) an a current i 1989; Loconte & Stevenson, 1990, 1991; Taylor on seed plant ыч. һу “aching both char- ; Hamby & 2 acters used by Crane and conflicting — had omitt "с їгагу ш our ir sp ions, > | p x dJihat and : е y Tobust conclusions concerning the origin of angio- seed gp are a рош МА group, with Soa rms can be drawn from these data "JAD tha: nks the Department of Ecology and дивні С Biology, ipei] of Arizona, for hospitality during bat tical leav ve in whi fr he NSF (BSR- ch much of this study was carried out. MJD is grateful for er1 rom the 822050) the University of Arizona, and a Mell оп Fou a tion Fellowship from the Smithsonian. пеню, EAZ 861 s C. J. Bult, Re K. Ha amby, and Y. Suh for help with RNA sequencing and the N NSF fo Aana M. W.C uggestions for improvement of the ı manuscript. of Бос and Ecology, U ity of California Dar vis California 95616, 2 gy, University of Cal А , uires ent of Organismic and Evolutionary Biology, Harvard University, Cambri ridge, но 02138, ‘ Bre of Molecular Systematics, Smithsonian Institution, Washington, D.C. 20560, U.S.A. ANN. Missouri Bor. Garp. 81: 419-450. 1994. d 420 Annals «чнай Ма Garden advanced “seed ferns” with platyspermic seeds and Rothwell, sperms form four major cl other groups woody magnoliids, (1) алая includ ing Chlo- ofw ойлы ‘widely dis сие becau: e of their un- , includin Widtefáo "E Illiciales, ind pem ssibly Canell с: (3) dicots with aa но and derived pollen, later designated eudicots (Doyle & mie ШТ (4) fons hori consisting z he 060 г semi- actoris, 8 А Piperales, № haeales) and Somewhat pua ейін ч were obtained by Loconte & Ste- t rt osite leaves and two-trace mt with Calyeanthales. In contrast, analyses partial 18S and 26S quences from angiosperms and other ex- tant sood pran (Ham i m derived clade, within w хоне А аге е роону resolved; These fenus give a would be herbace eous or nearly so, with | palmately- jb leaves and anomocytic stomata, rather than h pinnate This recalls xi views of n (1977, 1981) on primitive status of Piperales and/or monocots, in detail. BAIT More geo the view that the first angio- were “‘paleoherbs” hlor ana pss of pis vogue rý Doyle (19899) es 4 eudicot blade n in Hamby & mm a 992). Th difference may be p eU eating cor- related ovule ress ME to ge (which was assumed to be primitive) as three separate gee! and ці. аг ш of eee] Ден i analys bs P ., Trimeniaceae ich d end t Chl Madii ie with other pei (ies d An 87). apparent conflicts between the mi orpho olog- ical md molecular data, especially regarding “root- were dj an mit ooted a "E only one step pee parsimonious than near Magn oliale i rae a ‚сме where the morphologienl jar e ambiguous an they favo y magnoliid prototype, pix slig htly— о the molecular data point 5 mi ly e direction, toward a paleoherb met e better evidence on cats dar he provide er как in ships. In other cases, it may be the mo Volume 81, Number 3 1994 Doyle et al. Integration of Morphological and rRNA Data that are ambiguous et the — data a may re m are geo Thus the two sort: ntar a t the еен study we have attempted to address these issues by comparing results deri ived from mo mm and molecular data for taxa, probing € би апі weaknesses of of the results with m gainst combinin sets is that reater number of molecular characters will over- whelm the morphological characters. However, thi does not necessarily hold: if the molecular results are poorly resolved, as they often are, even a sma number rphological char. ea at this approach can give instructive re- is not intended to be a унду нага o liids and er rRN quences oy нар а аш en dato є with gymnosperms as а Speer Ме Ж “я Instead of considering all these data sets, beat, E. to address the two that we know M. fly some ргешп Sod жый including rbcL. on INING gig ia ANALYSES SEED PLAN PERMS AND ANGI is : major concern that address first e ч 79 y that previous ачат concern- ‚ rlgin of angiosperms were compro haps the position «е ап- in the seed plant analysis (Doyle & D. oghue, 1986) was a function of the assumption e i oli on basic states within the group. However, we E EE 1 | ^ а Z 1 d ld 11 о о / be necessary to carry out additional analyses de- signed to resolve simultaneously relationships at the point where the two analyses intersect. For р ih pere Jt Corato arpan share тап therefore be directly associated with Gnetales if Chlorantha sumed to be primitive. corr onious position for angiosper what internal конна ра ог Бава! па іп я: the problems are pet due e our em mol is Simply ler( 1994) an compares, This шой oghue & Doyle, 1989a), in | which we did a - inary analysis of Laurales to deter minn css states ina derived subgroup referred to as “соге Lau les.’ These e problems w were recognized and addressed a experim sa reported by d yle ueris E combined nine angio- rm taxa with th ee groups н їп Роу 1е & йозе (1992). Hee we dioit illustrate санау nature of the problem and the tial value of е g both ar altis an molecular eviden TAXA, CHARACTERS, AND ANALYSES ed seed plant and angiosperm matrix, in trees rooted both ра aleoherbs, which: necessitated dividing t he are represented by individual taxa used in Dono- Annals of the Missouri Botanical Garden ghue & Doyle ا‎ дщ i in four id composite taxa Magnoliales, based o ria, Myristicaceae, nnonaceae, and о тағ (“соге Ма ales" of Donoghue & Doyle, 1989a); Piperales (Piperaceae, Saururaceae); mphaeales (Nym- phaeaceae, Cabombaceae); and eudicots (Ranun- culidae, bo, Troch rales, Hamameli- dales) se are scored in terms of estimated sists of two taxa к these include е previous analysis have s P the group was ored as uncertain . In the case Le bibe characters, d isset (0/1) an kno ?) are equivalent in tree a s we usually E A between the mo scor ings s in but we did not try to weed out all cases where “9” дрен Ais а E } d 1 uncertainty I yses. In Magnoliales, usd states were estimated at Degeneria is the ips taxa, which them selves doa an unresolved шер баса ‚ Кап пип е plas A e and Trochodendrales T ame. dal Aus bait which was at or near the base of the ur and lune utu] order revious tree vitiis are ie special interest because they been widely discussed as possible primitive angio- sperms. Use of Austrobaileya to represent Laur- of troba ales might be questionem. etapa s Ачи ENS ad uer: renuit bear out ite wu in: хн vay: since Aus trees obtained, as it was with the larger data set. ing a taxon based oı “prog vite * (Aneurophyton, val ceca, as outgro up. haracters are primarily ed in е need plant analysis of (Done & a of Mi & Doyle Ө that are potentially informative for the taxa under consideration. To he M added кта apomorpluie es that i paien group, either as new v characters or Ms additional ne ree- nucleate RE complete loss of the Ж ДАДЫР wall). In some Слот; features that vary of characters reco ognize ed in seed plants as a whole (e.g., palmate leaf venation). For angiosperm char- ch ther acters in whi ere are no clearly comparable states in other seed plants, we scored the latter as 1, DOW P PR | * fü 21 8 where. we hoped to; angid biasing ‘the results by angiosperms and other g groups: These characters also form the basis as m RN. analysis of Recent taxa presented below as a counterpa' of the rRNA a KUATI for Simpy we > retained nine states that were pot ۴ T CHI tetracytic stomates in | Piperales, -tricolpate pollen in eudicots). ult ies chergeterp were aera pom Po for two e len size wall ASA Two ier characters involving exine structure deserve spe- us consideration, since our decision not to order hem had a RAE impact on the results pre- below‏ اچقا In the‏ e tudy (Donoghue & Doyle, josperms as anular, assuming ы eres structure within the gr ining the tw evolved e group. In co ata кыны our previous polarization of states within s could eserved by adhe атш oneycomb-granular—columel- in which angios ith alv out- roups a ies Cayionia, glossopterids} ae groups al in angiosperms ranular structure in groups a s is a mE gence with Bennettitales, Pentox lon, and ne tales. This scenario involves four steps if the exiné haracter is ordered but three if it is not. Such s bi ылга be ni p colum ra g е (ап prs RS miis within angiosper "mus si Le Thomas, 1980-1981) an ine of alveolar and gr an Cam & be ular structure within conifers assume that a rees with angios daas only ig жы дашы ert of the е & ma y and a мааа from alveolar to granular i5 tainly conceivable on structural gro duction of the side-walls of the alveolae, 1992) data se js cer (e.g ТЄ Volume 81, Number 3 1994 Doyle et al. Integration of Morphological and rRNA Data the es between the walls as columellae: E. Masure, 7 structure was пої included аз a char- e 0 0 р from similar tangential ha and chat i ngiosperms, which is nonlami- nated except sometim кт is a new vs том, 1984). Agi: the origina ы] „ре: 1991). Since there are too many taxa for branch- wl analysis, which andi finding a most parsimonious trees, we used the Me ш ae algorithm, mug 19 replicates of stepwi This increases the probability of finding most | par- simonious trees that belong to different "islands" iia uasa 1991), w bial were in fact found in e experim ents (see res sult ts). Alternative ar- acClade a PE constraints option in ı PAUP. SINGLE-ANGIOSPERM ANALYSES Experiments with single angiosperm taxa re- sulted in severe) diferent potita of the angio: At ar hophy tes (P пио, MN | П larizatio Ка уа Gnetales), and of anthophytes within set by 1 l 1 H 1 : 1 y see кы ted). However, because the ho- tid when Magnoliales are substituted amina mologies involved are rather speculative (cf. Ga- barayeva, 1991), it seems preferable to treat the three states as unordered. We also made a few substantive changes based on new and such as з recognition that pollen 9 а сома Абри 1989), С abomba- сеае ке) have a columellar exine struc- ture (Osborn et al., 991, Myristicaceae (Mag- noliales) have both S and PI type sieve-tube plastids, and Aristolochiaceae жыш basically РП Cond sperm nucleus with a second megaga- EU села This h has been confirmed in phe, 1992: Donoghue & Scheiner, ver. Sets of ER were нат using ew nine-angios; rix e set, v we Yzed the matrix with each of the nine ап s cheated substituted individually for angio- rms. In re moving th uch vel distort ures of homoplasy, E this i is not a emeret are not c rned s ч lasy. acne we did : riant characters with MacClade ipis thon & у ia. ison, 1992). All these data sets w yzed with PAUP (version 3.0L, Sw offord, As expec for angiosperms (Fig. 1), angiosperms are the sister group of other anthophytes, and psc feriis are $ 4 2 1 E Е Еу" а, glossopterids, and corystos in Doyle & pe ghue (1986, 1992). Relationships among other groups also parallel those an by Doyle & Dum ОК clu omg a some with Fé ar- аний by Loconte & ыша: rd in Кае 1; one of the characters that supports dé exine structu: In contrast, when — cenam monocots are also o true of monk trees found sen erids. TRA ia а 2), altho ugh in two such trees the “Bennettit ales Pentoxylon-Gnetales clade, Nymphaeales, дай base imonious arrangement). These trees s break up 1 } р the anthophvtes, al Pent r J с га) 1 J, + О. уе to angiosperms. This result wee be questio oned because it implicitly assumes that Geytonie andy F VUE © 1-5 } 4 L " because they are not preserved or not yet estab- реш i fossils, "A as | ор chemistry, a tunica sio is also true for Bennettitales and Pentoxylon ees of the sort shown in Fig. 1). One reason ч i new isis of these angiosperm taxa is Annals of the Missouri Botanical Garden Ee ا‎ LC | = granular IG um j Popma most drag tree (of 94) found when “core” ` Magnoliales (MAGN) are n. tuted tor angiosperms as a who le e in Шз ше angiosper T branch es shows distribution of the exine structu єз тебер which tends to link Magnoliales with eiue зоре (Pentoxylon, Bennettitales, oneal s). PROG = “pro; per ELKI = Elkinsia; ME = Мары ы = Callistaphyton: С ONI = Coniferales; GINK = Ginkgo CORD = Cordaitales; PELT = Peltasper- CYCA = Cycadales; CORY = Co Duce иы $ CLO = Glosopteridales CAYT = Caytonia; PEN Pentoxylon; — Bennettitales; EPHE = Ephedra: WELW = oon GNET - Gne presumably that they have columellar rather than anthophytes as Mag ar are. e struc- r ended to associate an- кна if it had been wnelar implicitly derived from qu ра t it does when it is treat ted а uno I sperms in general to Caytonits are reticulate’. ve- nation, flat guard cells, and anatropous cupules scored like anatropous bitegmic ovules gne Sr cipated, in n of the trees found when 3a), Chloranthaceae are linked directly with Cne- tales, with which they share such features as op- posite leaves, two-trace nodes, spicate пене cences (scored 1 as com re und strobili), hese trees, the s postion a ssn is highly ава they may be the of other platysperms or ae aia cads, glossopterids, Caytonia, or opteri ra ак are linked with Pentoxylon and Ben- -—-т2-ыс,„ =>» wezzvestosess хыс ошу со = 2 шу осооаосооә оса а E3 E3 E3 C3 C3 OPROG Treelength: 130 66 Chars. ЕЕЗ equivocal FIGURE 2. Representative most parsimonious tre e (of 31 ed when Nymphaeales ( (ЧУМ) аге mieten for angiosperms, tion of the exine struc- ture character. Generally Le trees (some с, angio- sperms linked wi a alone) are e also fo und when is pisne no Other abbreviations as in ats 1: nettitales (Fig. 3b), which maintain the мур such as compound st Anthophytes are not broken u trees. The most varied trees are found when Piperales strobili, as smn up in of these are substituted for angio sperms. As when se tha are the singl perm group, include элец trees where Piperales аге latysper H in trees of the latter т perales are пої d with tales but rather th Pentoxylon and Bennettitales (Fig. 4). ш is presumably because Piperales have f wet orthotropous and b ic ovules that allow ge to be neste t nnettitales and ge nodes te Gnetales and Chloran pe However, Piperales are ve with puts d few ae cally imonious pec "es anthophytes form a «rey group at e : platyaperms. are fees 2a vet "Piperales ar e © asl in anthophyt th mong Caytonia, ger to trees terids, ош corystosperms, analogo ous es Doyle & Donoghue (1992) and trees found Magnoliales are substituted for angiospe linked 1) and a few trees where Piperales are Volume 81, Number 3 994 Doyle et al. Integration of Morphological and rRNA Data = e > ы m] OCALL 4 Treelength: 135 66 Chars. 6-2. Œ 5 > — с Э ЕРЕЕН НЕЕ ЕНЕНЕ Кпуш>=ло—-—оо=ош/7атшш ш >оа сосоооооЗаошсое а эне еле а е] шшшшгшы1г г рер 51 oar ШШ compound v parsimonious t (CHLO) are drin iosperms, БОМ, hg i disibibution of the phyl- ias (9s d rei (b) characters; Bonds isan neo-englerian he ophytes nested among “coniferopsids” an any cordaites). Other ines as in FIGURE 3. ый) o foi id wh Сы direct] with Сау: torts aytonia, thus breaking up t — а 28 мор => VSzZZensSseSsosewess JFI SOE Z52 > ا‎ ш ш > с) со Оо @„ о со о о со о Даша а с [ШШШ еШ Ше е е Ше Шоо а EP C3 Cj Cj шиш LIPROG Treelength: 138 67 Chars. E 4. Representative most parsimonious tree (of iet found when Ier (PIPE) are substituted for a ngiosper © h Ы, those ir in od 1, 2, and 3b. Other abbreviations as in El are linked with Caytonia and/or glossopterids i im- e reproductive structures o i d the cl Ind т Е] = = © = ® = ае = or n ees in which an- ed wit th от оман апа Спе- e are а. imply that angiosperm flowers and floral parts, ие. e ontaining several ovules with t two lah ted £ by ager egation of several M “flowers” a, a pseudanthial interpretation). These results bear out the concern that the Ai > 5 . : 1 ^p EET e © Т r J ` 7 & Donoghue, 1986, 1992) may have been incor- 1 23.252541 4 E „М | etat, . Presumably, Pip an have fewer char sup PP9r t ary O! пе as wth er. The j : ese ciem ts i ant e alone are considered, since in sre "allt trees x re the closest living relatives of angio- ir ptions are the few trees in whi s and Gnetales form a paraphyletic е variations are а ction of differen o Scenari hn M because of the gaps between extant P5- For example, trees in which angiosperms in angiosperms. They that angiosperms а e po csse sin with differe nt *an- giosperm" groups related to different "gymno- sperms." Ho we even, пеше сопс cuman Y fo low y dep whether and how Ч sth h oth © F о t t NINE-ANGIOSPERM ANALYSES To assess the possibilities just raised, we included s and a naly zed the re- andom t which differ only in arrangements consisting of Оши. gan согу- Annals of the Missouri Botanical Garden GNETAL ANGIOSPERMS Wo DEM ee CERE coz. -2- = xum =. Zor = RI XFEFEERFEFHEEHHAHAHHHLEHEE CEuxoookooconburdcoxczill4a2cu Li Oo oe E3 C3 C3 E3 BE EJ EJ E3 C3 C3] HB шсш EJ NS Eee Treelength: 192 Representative most parsimonious tree (of мен found when all nine angiosperm taxa are included in FIGURE 5. the hos рыз distribution of the exine stru cture ee = Nymphaeales; MONO = = monocots; pu with tricolpate and derived pollen); AUST = r. PIPE = Piperales; ARIS = Aristolochiaceae; МҮМ. e" Magnoliales; WINT = = Winteraceae; EUD. Ди. pare e 1 her abbrevi n Fig stosperms, Peltaspermum, and cycads (e.g., Fig. 5). In these trees, angiosperms form eudic paleoherbs were substituted for angiosperms (Fig. i rmor ar i i (t, f, à with tre in angiosper ere associated with Caytonia EE glossopterids, this rearrangement is pre- ture id calis iine as unordered. This h > I 1 : 1 1 1 T J n реп- with gran- nettitales- Pento iso Crete: clade, ar Mag hed between the latter and noliales attac peu angiosperm Although these result cast wine on previous the —— бот! the previons situation i is less radical than s. Essentially the same ore лаа аны: аз presi native re- y parsimo- nious b le & Donoghue (1986, 1992) м (19 апа Mpeg are basa As in the p us analyses, other one- pre tr ees are tales, linear leaves and simple d 2 basal and relat inci plesiomo pce к ant vn М тар ез аге тена ni s aee T A data (Hamby h & Zimmer, 1992); in fact, some of these Ri Я same arrangement of paleoherb taxa мете? rRNA analyses present o Piper: d above N pesce and monoco wi 2 tolochiaceae (Fig. 6c). Given the ences in parsimony, it woul ANGIOSPERMS [- Ө os: = EISfEIIÉ cCousScecz2i BESS Bee a Treelength: 193 uncertain Sporophy Ils 2770008050088 b Mega comp Treelength: 193 Mei © > ES е. о тшу A al = > – we Zaru о — e © exOZe7 505 eosS776S5 222252255 aAuSFOO= u> EKOZSauui C> = HE y25 МК т Кк 53 wa MOO ZA У сш co ggg C3 C3 C3 EP EJ EJ Ed E3 C3 C3 Li NE NE NU NN UN E ШШ с Ficu; Ў P me RS Ue. gom : rm data set: (a) with ang & Donoghue erms p^ sister group of E antho hytes and M х t te d Do CIR paesan pe 14 of the exine structure cha sey e — ĉe, with апо phytes nested among “coniferopsids,” эч distribution of the sporo deg character nd АНА ыт linke d with Caytonia and bine sopterids and Nymphaeales basal in angiosperms, A data, showing distribution of the exine structure character. Abbreviations as in Figures 1 cod 5 Annals of the Missouri Botanical Garden OPROG OCORY 06105 Treelength: 198 RE 7 S. x are forc the base of angiosperms, show = zZ ы a oO GNETAL ANGIOSPERMS Г ПП > a LR زک‎ ЕЕЕ Бае 527352. е EJ E3 икшг г E Fae Representative tree (of 1675) f ound when angiosperms and Gnetales are forced together and О wing distribution id 99 phyllotaxy character and 5. . This tree ed to steps less parsimonious than the shortest trees. Abbreviations as in Figu } f, 4 ak 1! } of taxa and the omission of conne a. d d. ps. ternative topologie ri ts. The most interesting experiments concern proposed links between Gnetales and Chlorantha- ceae. Trees with angiosperms force ther Gnetales using the constraints option in PA re only two steps longer than the shortest trees, but Chloranthaceae are no in sperms, a ight be expected from their gnetalian features; instead, angio: s are arranged as in the mo Д © Ф "O @ fæ] B Ф w © sal in d with ‘Comdial (Fig. ae are six "m Similar the englerian trees (Fig. ge ашны, s are not ar- ed in such a way that the similarities > deg Сасы апа errs are homologous з аге linked with шымы, Өр o unparsimonious, even though ceae are associated with Conde when they is E that ter ant are “screened off” es hav- an influence on the position of angiosperms by beng linked Um Austrobaileya a is di groups, globose pollen with lat Ipture = a ver- rucate sulcus. Similarly, other paleoherbs Бей оп n herbaceous habit, dis- tichous leaves, palmate venation, and globose pol- len. Of the potential homologies 3i “Chloranthaceae d Gnetales, opposite leaves and two-tr ace nodes ios a mong are unequivocally primitive in angiospe ues like Figure 7, since they also occur — baileya. Howe compound strobili would hav to be losi below Austrobaile И 15 independen tly “gneta a Mir" ê morp sistent with a link betwee: Gnetales when Chloranthaceae alo и anra" tuted for angiosperms, but t y do not а Chlor oranthaceae to the base i me Mane ppl angiospe a are in in hough Chloranthaceae are like ueniet а А e more’ like Gnetales a тәр groups. In the tree shown in are unambiguously supported by nine | Volume 81, Number 3 1994 Doyle et al. Integration of Morphological and rRNA Data characters, respectively. i of the angiosperm omorphies are universal in the group (or rep- resented by clearly «нз ore and unknown elsewhere (two pairs of pollen sacs, endothecium, stigmatic pollen germination, loss of versal in angiosperms bu rangement. Four oth e known only in angiosperms are equivocal as angiosperm synapomo phis because the correspon char act Ca ie а and gloseopterid (companion cells, thre ea angi osperms, bi per stayed together as as a clade. Apparently in most magnoliids, and palmate ve- nation in paleoherbs and wA vhs to hold them together. some, this result may seem trivial, but even e some authors have expressed the opinion ansio: that has held back млд іп оздон баі the hn of angios s (e.g., Hughes & McDougall, 990; Krassilov, 1991). Similarly, the view that s still frequen might well un ешо а vara problem their cl modern groups that are separated from a relatives by “long branches”: the more in ers that unite the group, the more certain —. status, but the less certain its position. This h may also apply to —S can ( (ef Wake, 1991). For the same rea- rooting of the group may be pric soe be- 4 vide little ' ‘signal” as ; to which subgroups ere Pn (Wheeler (1290); This probie is кее could not be polarized by out ero up comparison in Donoghue & Doyle (19898), or could not be pd it is ma gnifie d by largo rm characters in fossils. in greate d й Кар (Doyle & eoe ode the problem might be solved i те, of fossils on the lon branch leading angio P dem e., non-angio- opi wu ee ophytes, in the minology of Doyle & Donoghue, 1993). How- still no fossil viti аре states in some 1 1 1 1 ae е } р stem lineage. MORPHOLOGICAL AND rRNA ANALYSES OF EXTANT GROUPS The relationship of these results to those ob- RNA sequences is addressed more 1 ry ла t 1 d rRNA tained from r Je. Д. L directly b у +h t © d with the mor- ince still of the osperms are ambiguous wi lecular data pare the disadvantage | of pong are ы. аше ont fossils: e.g., Gol enberg et ral. 1990), whereas mor- phological data may exist for key fossil taxa that er ad & Doyle, 1989; Donoghue et al., ram over ' interpretation of the pescar homolo- ies of angiosperm structures. TAXA, CHARACTERS, AND ANALYSES e starting point for our rRNA analyses was a successor to the 60-taxon data a of seer da а. Com- Zimmer (1992), enlarged to include plete sequences are available from dienen (Ac- Annals of Missouri S NS Garden cession Nos. M81965- a and the NMNH Gopher Server under ir ” Our e! data йе! is derit ted t th g m JL 1 what different angiosperm taxa and removal. of се groups (Appendix). We encount ove a айыу of probleme in eb rRNA. analyses and combining the two data sets. This requ aired many compromises pei approx f mati but we ssp are unavoidable if progress is to be made at this p (cf. Maddison & ved 1992). The important thing is to spell o assumptions ча so that they сап hs scruti- nized and tested in future analyses. In reducing the pranan and molecular data sets to a f taxa, ae баш was s اظ‎ + b d A ЖҮР Н NM FE uus 1 f H 33 ТС FO | ing ‘the number of taxa small enough for эре тоге wW analyses. which rRNA data are still lacking; this is often unfortunate, since current evidence suggests that some such groups constitute important links. Ex- amples = pren which recent authors have linked wi e (Carlquist, 1990) or Aristo- кеге (Donoghue & Doyle, 1989a; Qiu et al., 1993), and Austrobaileya, Tri iaceae, and Ат- imme many of its morphological characters are difficult н ni an ho ough Ceratophylfum paepe a cl pen of seed е a whole е (Les et al., oon Chase et al., 1993), ирин is unstable in un- rooted rbcL jn of angiosper: alone (Qiu et osi oghue, 1904)" Similarly, we did not include any non- seed plants as outgroups, since ынды in the original data iffere sets were different. In the morphological pier of extant seed plants (Doyle & Donoghue 7 1 ‚ wea ed t a hti dps are vonian "*progymnosperms"' and Carboniferous ко ferns.” There is therefore reason to fear that any той obtained from extant outgroups might be an artifact of spurious long branch attraction (Wheeler, е Maddison w al., 1992). The re- sulting trees are therefore unrooted networks, with t to cycads. However, it will be seen that this pro cedure does provide instructive contrasts, because trees derived from the morphological and rRNA data sets are җе. ч “diffe erent. In combining original taxa into larger groups, age змен even though they were strongly linked in the trees of Donoghue & Doyle (1989a) and заре іп some one-off “RNA in th th icy (1992), ipd we combined Hedycarya ae are not associated in all most parsimonious rRNA trees, because their relat өю is strongly € ported and uncontroversial on morphologica grounds. Despite our efforts, taxa in the mor rphologi cal except perhaps when they are monotypic (Le Ginkgo, 1 Welwitschia), са general, MAE Chlo us; Ranunculidae by R Mi ааа by len a deta the a assume that they are adequately m eee that rescoring them x dendron aa most o — Troc ы ул me: sculptured a сөе e of oil cdi), plus one bres woul if Ranunculidae as a whole were pnia aie Volume 81, Number 3 Doyle et al. 1994 Integration of Morphological and rRNA Data 3 25 5 E 1d 2 LE o uu 5.2 л 2 id t 2.25 КРАМ. DU MEE QM I T є 4 БЕ. 6224$ S228 222,5 $2? AES е Би 5s а Сас Es CDU E A Beis PHEVE, SES mBONAS Pes E * 45529 *?&"^acuwnd2t3o9459Xo£t.L|529Uut9505jo049c£t2££u9edv»sa9»79722 болем 2377 Sacouno6sbtLcogE2Z9SkSasSon5s5ot9n246o9CotHo94 HHATHTEEEEEREHTEHHGEEHEHATRELOIHBHAIBRE Nüo3so&oóosuuurzzmmamoacaoruüu4-óaoóramc-4odzuoáifo«xf«r-o600Fn V ‚а = 2o щт =) Zo z c c cas v r= = ©. ы ® e v © 9 x „2 % z ave о = своте о КЕЕ ҮЕ; ст бот со [33-5 E TO—-59v9cCmX $ S.2£ 58 rEo«cora iid | а b LAUR FIGURE (a) Relationships assumed in reducing taxa in the original 71-taxon rRNA d 8 taxa e in the present rRNA a шау: ses atl text ЖЕ 9 CYCADS = Cycadales; CONIFS = Conk ‘GINKGO Ginkgo; GNET = Geum ELW = EPHE = Ephedra; NYMP = Nymphaeales; PIPE = Piperaceae; SAUR = Saururaceae; e; MAGN = ini in WINT = Winteraceae; CA ALY z Giycanbasean CHLO = Chi “setae AUR = “соге” Laurales; U = Ranunculidae; TROC = Trochodendrales; A Aristolochiaceae; MONO = monocots. (b) Relationships within core Laurales assumed in the morphological dun n лн hi se of variation within that group (stamens with | f prec F vel. на filaments). among grasses that are consistent with rRNA and coring taxa that vary for characters included morphological analyses (Kellogg & Campbell, in te matrix, our goal was to obtain the best 1987), treated Hosta, Sabal, and grasses as a ime of К сы states for the whole taxon. trichotomy, and treated — aroids, and the ^^ the rRNA data se ‚ we accepted internal rela- Hostar Sabale фы т» de а аз ЖЕ Бону, In the d etates for ge that were consistent in the original 71- у сие " 9n rRNA sis and in morphological analyses Magnoliales based on 9 E that Dege- J ourselves and others (Fig ) example, neria is the sister gro of Myristicaceae, Anno- nig d in the gom we as- naceae, agnoliaceae (cf. ); in core чава th 5 is the sister group of Zamia and aurales, we assumed relationships s in Figure sod halartos, and Pinus is the sister jme of 8b, derived from an unpublished analysis of Laura- "nlperus and Cryptomeria, since these relations les used es. the same purpose by Donoghue & re А ; н м found in the whole rRNA analysis and in the = le (19 qo logical analyses of Hart (1987), Crane he use a parsimo ny in optimization of ances- Чаш and Stevenson (1990). In contrast, in es on trees is discussed by Swofford ч «j Phaeales the rRNA data indicate that Bar- Ma (1987). As in the nine-angiosperm ana CN 55 the sister group of the remaining taxa, ysis, when clades а of two taxa that varied т: morphological data (Ito, 1987) imply that Ca- at a given site, we coded them as uncer pose mbaceae occupy this position; therefore we treat- there were three taxa, and the “‘outer’’ taxon a ed the t three © groups as a trichotomy. In estimating one e of the two “inner” taxa had the same state, Annals of the Missouri Botanical Garden es NEE N. s 2252 " | w^ 1 1 Į $ when the outer taxon differed from both inner taxa, we scored th 124 $22. T as ^5 we used MacClade (Maddison & Maddison, 1992) to find the most vi in apie neg — — tak- ing int f trich omies and ctp omie The pr: por of (ar dine Мане. tara mg scor- on & Davis (1991). In nes a и эйс of ho- 5 risks of its ison & ж 1994) However, i in pns ice Nixon к f As illustrated. graphically by the experi- ridden with as many m assumptions and (indels). We included the indels in the analysi nces leave no ambiguity as to their align- e data m obs analyzed with the heu- ristic koe P (Swofford, 1991), with we replicates of stepwise random addition of taxa d TBR branch и g. Alternative topologies were investigated with the constraints option in PAUP and with à ре (Maddison & Maddison, 1992). m o evaluate the relative strength of various results is єн к analysis (Felsenstein, haracters are sampled randomly i = étui data e with replacement ‘ ee calculated for at the frequency of bound bd in which EN occurs is an estimate of its statistical sig- nificance: , if two taxa are united in 95 out of 100 0 replicates, their args Ed hg р > quencies should be interpret ed. in in this wa matter of debate (Carpenter, 1992; Hillis & Bull, Lies) аныл Ы Kishino, 1993), but the crit- 1994) н g up - * | 2 n f - relevant monophyletic taxa might be the ideal so- lution, but pis carried to its logical conclusion this vod quickly lead to computational pude sis. In the а. we prefer t umi — about a states of the sort described, while be tested i in n the fu z Our muti безне iiu set is presented in rcd z — сыч аге "en same as m all into question the value of bootstrap ana x eans of eval the relative robustness of ts s. Actually, tion experiments suggest that the pes errs on the ide of being ervative; under m pe stances, clades see I the conventional limit of 95% are more accurate than the bootstrap n rs would imply bier’ umbers Bull, 1993). In addition, bootstrap analysis ТЫ? be useful in uncovering possible alternative re ‘ tionships, as seen in the aoc ict of “frequent ih occurrence" of groups provided by y PAUP. gıosF seen in the most eat: trees is actua ke К labank r ONEEN ым 1 Дд there also be be minority f fossil dicated in th nitions. characters that reflect he т relationship, ^ Deletion of characters that became финн these should be sampled and amp шеи ез. For after removal of fossil tax - left a kei vil 69 — wit high frequency ч m strap replicates si acters. This data set th 1 tant each data set, we did 1000 "d replica seed plant matrix of Doyle & Donoghue ende To i increase the ا ا‎ f шн ИШ МА 4 is pres Table 3 nious trees in each replicate, we did 10h ‘él (Appendix). Characters from ni те analyses with stepwise random addition of taxe BR branch swa a from iminating characters that became r of characters was re- unin ati "d duced from 411 to 174, of which 167 are base ed to evaluate уй ei 1988; Done ermines s is decay ана Беан ‚ This method det mony has to be relaxed—befor e trees 8 Volume 81, Number 3 1994 Doyle et al. Integration of Morphological and rRNA Data GNETALES ANGIOSPERMS GINKGO a” a c 2 > [а] Su $ 2 ы ج‎ 5 © 3 ш > on 9. T f the f. У 1 AC RES RSEN the ni ; Y ‚ fe UEM umber of unambiguous changes supporting each clade. Abbreviations as in Figure 8. whic h a given clade breaks up. With PAUP, mendis ing all trees of a given length; be- з " Is was not ima with 18 taxa, we use uristic search with 100 replicates of stepwise iss addition and TBR branch swapping. R ESULTS OF MORPHOLOGICAL ANALYSES fou EM of the mor d data set yields Sie Parsimonious trees of 150 steps, two in Figure 9. The variations concern аа Magnoliales are е sister ыс up of th of t bra lauralian taxa or n Fi dit e E the ae — ra е e that | un- be vocally support clades Y at least fifteen бшш rid by iih t indicatie a groups by seven. This gives some lon of th i т level of ме t it is elin dis- ing because it says : no about the f homoplasy— rir these geom the 7 consistency ker is 0.58, xa anderson E 1989); a retention е 0.7 — show the arrange t "i dos gemen per rmous groups found in in analyses of erê taxa alone by Loconte & Stevenson (1990) а: ). Do yle & Don е (1992 у!е & пае (1992) argued that this res an artifact f о g fossils: when fossils are included in the muttin 55815: нс this mecs of living groups is only Ї seve arsimonious. In addition, several of the characters B яе od fossil TA are impie oen bitch they also occur in Caytonia (flat stomata, & F ), Ben- ), or both (pinnate sporophyll organization, integ cellu s). Ths whorled тая апа Ша with Bennettitales. Of the characters uniting an- giosperms and Gnetales, opposite phyllotaxy and 1 kaaf R КУК | а уш F ore fe are interpolated between the two Conary to iome & vg ане the four mious trees root an among к» jees gr ls with Nymphacales plus Ae ipa s ав че sister oop of E 1 } " dad lack of oil cells ovem the latter may not l in alid if Acorus, which has oil cells, is basa onocots, as inferred from rbcL : Duvall et al., 1993). However, conflict with the Dono- he gue & Doyle results is - severe man it repe noliales, ‘only one step longer trees are obtained 434 Annals of the Missouri Botanical Garden GNETALES ANGIOSPERMS S = oni = "z A P = e ec SZZEZEES22Z8zZr5 222 SoEaooeeae SCS ESOS ESSs SESS Boo [Lr] ш ш ш m m gm m m m m m MORPHOLOGV FIGURE BH A i sep lese parsimonious free based son the үү жасу data set mh Марш basal їп of th angiosperms, that are almost entirely consistent with trees in onoghue & Doyle (1989a), except that Laurales are paraphyletic rathe monophyletic (Fig ), and onl e additional step is require ue to possession of PI type sieve-tube plastids, granular exine struc- чик апа а ыса tectum, all features that appear to be convergent when more taxa кы їп- кыз: (Donoghue & Doyle, 1989а). A d Ithough monoco with Nymphaeales rather than Pema ee as in ‚һе rRNA a анау presented be low, t with Aristolochiaceae are pax one step less par- simonious. This shift in rooting may be partly a function of the smaller sampling of taxa, but as we argued 1 pod ope E : 1 1 f T 1 © ysis, it and ende ch han implicitly or ordered. ‘Although the new — results more consistent with the rRN the -— previous conflict. нер еер прозе from the resolution the моа a evidence—actually, arr tei this view. Previously, — data кый A maenol alian rooting, but only weak- ly; now both data sets favor a paleoherb rootin ing, but the T data do so weakly. The fact Abbreviations as in Figur that o sd followed from a rather subtle change erpretation of two characters underlines the h venation, paracytic stomata, cellular e Presu umably, features of this sort are ise for th erms among Une es m some in the trees a Nixon et al. (19 994). m Loconte & Stevenson (1990) arrangement sie cads, Ginkgo, and conifers occurs at а fr agi of 79%. Except for Piperales, gon Di angiosperms that were seen in Donoghue а uch lower ine the rooting ights come - > e bootstrap results also ssi on Der + rezî although only indirectly. Ins Volume 81, Number 3 1994 Doyle et al. Integration of Morphological and rRNA Data E. GNETALES ANGIOSPERMS ш: 1 ве u = a © SEizbaeizeseg222298 P200ontzraocoozES23I555 90 d4 171 3941 9945 4342 1841 1T di 6 di 24 d1 92 d5 100 d>4 MORPHOLOGY [uio roit ВОЙ BOOTSTRAP, DECAY ж 79 42 FIGURE 11. the percent. I Results of bootstrap and decay analyses of the morphologi 1 in whicl h clade is found; the seco cal data set. The first number indicates numbe indi h steps lo; ^ KEE ien axl trees must be before some are found in which the clad possibility th incomplete; clades that had not decayed in that searc y that they do decay іп "islands" of five-off trees tha from examini she f. f 3 lad containing all p" га а Riu Ы ut опе or two taxa, which imply e ta + 1. 1 1 E b without spec- tym; Н i ze their exact arrangement. As expected, = Pings implying that Nymphaeales and mono- 2 аге basal are freq but at only 24%, еы by Nymphaeales al t 17%, and Mag noliales ; e: s : Sat 11%. These results again illustrate the the root, while favori 5 аге basal , ring the ing within angi and ^ia angiosperms are Piperales (Piperaceae T Turaceae) and eudicots (Ranunculidae an roch à odendrales). This result reflects the unstable nd number (91, etc. y ade no longer occurs (decays). The search of five- h are labeled d > 4, because t rul t th t t di red. Abbreviations as in Figure 8. next to paleoherbs as a group in t i angiosperm analysis: Fig an y magnoliids as a group. It also illustrates the fact that strict consensus trees may u timate the amount of structure in the data: if a single taxon (or the root) “jumps” fro e clade to another, f сусадѕ © апа в in two steps, and the relationship between Welwitschia and Gnetum in four. However, angiosperms, Gnetales, the rela- hem ір; 1 till intact 9 t tionship between t in the four-off trees. Fi tr exhaustively because of time and memory limita- tions, but both Piperales and Gnetales decayed in m + ch 436 Annals of the Missouri Botanical Garden GNETALES ANGIOSPERMS ^ wm Or if qe > 25 2 Су 3 A 8 rRNA Treelength: 405 CI: 0.58 FIGUR The two most parsimonious trees found i in analysis a P rRNA data set, showing the number of E 12 Meo d changes supporting each clade. Abbreviations as in Figu several incomplete searches at this length. The eca d of Gnetales reflects trees in which angio- sperms are neste in the group (cf. Nixon et 4) се trees rcing angio of Шш length (155 ше Anginepeimo m an- but because that they do not decay in "'islands" of trees that were not searched, they are labeled d > 4. This decay order closely parallels the relative epe of clades inferred from bootstrap analysis RESULTS OF rRNA ANALYSES Analysis of the rRNA data yield of 405 steps (Fig. Ll differin у Laurales. Т' ith — as Бош the whole rRNA didis set t (ef. Fig and аа magnoliids. The consistency index is the same as in the m .58, orphological analysis; the retention index is 0.66 on this com- Ф Ф ori that ny more o reliable than P »morphdlagisal data, “нона strong conclu- un ed warrant aides of differing ESER missing data. f чыгу conifers, and рин ived from morphology that the group dd to anthophytes is Ginkgo rather than conifers. Angiosperms and Gnet etales e plan is "reae with Equisetum and Psilo by & Zimmer, 1992), it is oir i two steps less parsimonious to associate angiosperms with a clade consisting of Ginkgo, cycads, ies conifers, rather than with Gnetales. The figure of 12 synapomor in holds if seed plants are rooted somewhere cads, conifers, Ginkgo, which v we oyle & Donoghue, 1986, ee record; cycads, mec rs, and pue appear in t e Carboniferous or Per е 2 бонк er toe "s the line leading to d no e Late Triassic o (Cri 1 Do a уч ы, 1993). are 1 п attraction between ‘Nymphscales an ciderations argue groups. How Volume 81, Number 3 1994 Doyle et al. Integration of Morphological and rRNA Data E GNETALES ANGIOSPERMS ы. c = ا‎ = z = ea EES ESS SCBSESEFEABSESBESBEE SES MAGNOLIALES BASAL FIGURE 13. angiosperms and paleoherbs are forced t togethe r as a cla Representative tree found with the rRNA data set when Magnoliales are forced to the base e, as in most rsimonious trees of Donoghue & oye 1 gly а) and Figure 10 above. This tree is 13 steps longer than the most parsimonious trees. суола as in against this. m trees with the positions of ord ales and Nym EH m trees based on morpholo e: al rphology, j «йе g groups are interpolated in various ar lal rangements betw ther woody magnoliids and ов . These are viti pa- leoherb- ат trees with Magnoliales alone pulled to the base. A more analogous tree, obtained by networks on the line leading to Magnoliales (Fig. 13), is 13 steps longer than the most parsimonious trees. Experiments of this: кош fail to перрі "ves current hypotheses Tree with. Chloranthaceae forced to the base of Taylor alycan (cf. Loconte & Stevenson longer. Ae when Magnoliales are forced ш = base, nd other woody mag no! — and eudicots. The shortest trees obt ained by re iem | - most p to Chloranthaceae and теа оя аге lla rees icots as the sister group of other lint ern angiosperms, Annals of the Missouri Botanical Garden FIGURE 14. R Abbreviations as in Figu analogous to trees based Е 1991; Chase еї al., 1993; Qiu et al., 1993), are (Fig. phology. A he monoph is very strongly supported (99.997%). The link het : 1 1 Ч 1 (40071 BIOSE weaker (Об 70), but the monophyly of Gnetales is stronger (99%). This result supports the view that the weaker mor- GNETRLES RNGIOSPERMS 2 гё T = 2 e z — m Szzebeatt2e'b 22358228 0.0 & 5 a.2 8 ££ Ss ЕБ 5.5 ТИТИ 92 d» | 54 d1 E 34 d2 15 d1 17 di 65d2 2311 26 d1 45 41 54 d2 99 d»5 100 d>5 rRNR "2 BOOTSTRRP, DECRY 74 43 esults of bootstrap and decay analyses of the rRNA data set (see Fig. 11 for explanation re 8. h fossil relatives are more closely related to eac are y living gymnospe group, not i related to any living gymno tonia and others to G Void till ave the correct relationships among win. y h a 1 angiosperms would form one branch assoc my o Gnetales and another located one or п} BIOS[ p yiy аррпеѕ ошу to living groups, leaving open the possibility that different “angio- sperm" lines were derived polyphyletically from different fossil *gymnosperm" lines. However, this objection is valid only if all angiosperms and their © E А b t ported than they were with morphology (6576), Би to е m — © nection between monocot weak (23%), monocots are link Volume 81, Number 3 1994 Doyle et al. Integration of Morphological and rRNA Data phaeales in only a negligible 1% of the bootstrap replicates. This may be another case where t a favor one alternative out of two = [7] rRNA dat. that are almost equally Leap in terms of mor- phology; it requires onl a step to associate r y monocots and iara da in the morpholog- i sis. The h esis that Aristolochiaceae monocots may be more plausible if Dioscoreales are basal їп monocots, rather than alismi rooting pro can again be addressed by exa g the frequency of чы 5 containing all . All higher-fre- rt que that one or erb СА ps is basal: ids toe in 22%, etc. E & Hickey, 1992) are bas- onding groupings are observed at КУ mea of less than 0.2%. iai Wint teraceae, Ranun clades left in the consensus of one- Lit € perm th ас" these gro ups decay in two-off trees. The ar- B ngement of cycads, conifers, and Ginkgo breaks Р eg me Жаны ш! Een ss pe Geile s, of Welwit чы ea association dli. schia and Gnetum are still in This deca ay orde e Prov. strength of clades inferred from the boot- ар analysis, roi less precisely than with the morphologica] dat R ESULTS OF COMBINED ANALYSES ез ден of the combined data set ie 15) yields vb tree of 563 steps. The consistency index Which is almost sen to dt in the two a. “og ie (0.58); the retention msn я 1. This re futes опе poss sible | argument a morphological and molecular y are, that adding ns отеран data sets should a in more total n of Figure alternative trees 15. aii oo graphically the complementarity of the two data sets. In non-angiospermous grou ps, иу on tep longer. The strong links inferred from hath * dat + Г tal r о (united by at least 26 characters), among angio- acters), and between angiosperms ted from the ambiguity of the mor- are rooted among the ч @ = ix m 3 ga =: o Ф E Magnoliales forced t е ШШ торе д than the shortest tree; trees } 1 14 and 15 steps Ка respective ely. Despite the cem on number of rRNA char- acters, other results ore consistent with the К olo, iu. Wr in quu with the expec- tation that even a few morphological characters can be decisive when molecular data are a шш: group, as in the morphological trees, rather than the rRNA tre cots interpolated between Giris re and Piperales, although it costs only о tra step to th Aristolochiaceae, as in the rR analysis. Most significantly, Ranunculidae and Winte: rochodendrales and are instead the sister group of Laurales and Magnoli- ales The general result of the bootstrap analysis (Fig. 16) is that the two data sets te y to eei each other in cases where Angio- sperms (100%), Gnetales Y 10022), ‘Piperales (99%), and Welwitschia plus Gnetum (98%) fo clades at bootstrap frequencie: those in the separate analyses. Angiosperms and снаи m united a t the 98% lovely ipe n тке [п a eudicot clade— тп Т' 7 [MET rRNA analysis. Thi i Шеш patterns of homoplasy will ey love memi are detecting the same real phylogene ignal. Con versely, basal seed plant relationships, mares are e less dci as conifers and antho- phytes ed a еу ч of only 54%, bein than 79% in fe morphological analysis Am subtle effect is den the t two data sets bió the most parsimonious trees derived from them Annals of the Missouri Botanical Garden MORPH+rRNA Treelength: 563 Ck FIGURE 15. GNETALES ANGIOSPERMS о 225! a VI i3 = = SEE E E SIS EE Оо Shs St Б ЕЕЕ SC Single most parsimonious tree found in analysis of the combined np = rRNA data set, showing the number of unambiguous changes supporting each clade. Abbreviations as in Figur were not congruent. This involves the link between the two e udicot taxa, which is se ct = o % of бя rather than in the molec reflects ‘he existence af *minori sit gi that this y" rRNA char- lb rbcL data: Chase et al., . Once A oin rac Rie those from Yerba ology. The NONU dde ch ef- fects i pice et al., the 91). is analysis, the relative support for aie inferred iun че decay analya is (Fig. 16) roughly parallels tst the peres clades нага тоге е slowly than they did either individua parsimonious r-off trees are still present in eight-off trees, pene which the шш was abandoned. CONCLUSIONS These exercises clearly show the utility of com- bining molecular and morphological data sets as well as analyzing them separately. This procedure h th PS Se E | E а; } RI г о sets even when опе is much larger, presumably because there are minority characters in each data set that reflect true eee relationships (Barrett et al., 1991 The s dad results of these analyses are that angiosperms, Gnetales, and Piperales (Piperaceae Chlorantha са аге e plus Saururaceae, but not m cl that polyphyly of кү h mined for lack of progress in ce so far we have o у t ве меры бр eid in the pet data set to p Volume 81, Number 3 1994 Doyle et al. Integration of Morphological and rRNA Data Ew GNETALES ANGIOSPERMS A O Lu 2 ы. a © Best tus S 2 ozZzZzuzz£st2co£zizdazc û 5 0 6 3 & эсЕЕЕрЕ5Б5ЕБЕ Е ш 2 а о ш к= 4 Ээ о со шут 98 d» ] 30 d1 5141 5143 99 d58 50 d4 32 d3 53 d3 33 d1 45 di 65 di Es uk 100 d>8 , MORPH+r s BOOTSTRAP TEENS —— Ead 9 54 di { FIGURE 16. Results of bo th ned кыл i о clades. Still, it appears that the rbcL ovules and no perianth are primitive (Burger, 1977; 0 not support some Taylor & Hickey, 1990, 1992). However, if taxa different arrangement so ] th th. д. + Strong] gly that the h ata, even A КАТ are arranged as in Figures 9, 12, or 15, it is more . 8 1 РЕ 1 t res there are more potentially inf aracters than rRNA characters. ts among molecular data (ee в 5 E NA y magnoliids. To discourage potential miscon- s Ptions, we should clarify what a paleoherb rooting ould Say about nrimit; T (of. Taylor that the might seem to support the i+} thotropous r а aed nowers E © t © had flowers like those of Cabombaceae, Lactoris, Saruma (Aristolochiaceae), and monocots, with one perianth parts, a trimerous d gynoecium, and р | т of Nymphaeaceae and the tubular с of most Aris- tolochiaceae would be autapomorphies, although the laminar pl tation of N haeal hol СА 1. g primitive. S would be differentiated into filament and anther, not laminar as in woody magnoliids. The basal position of N ymphaeales and Piperales raises the intriguing possibility that the fici 3 1 : 4, t t l state. not * = derived as usually assumed (D. Haig, pers. comm.). The conclusion that Gnetales are the closest mod- ern relatives of angiosperms permits a wide range of floral prototypes, depending on how fossil taxa Annals of the Missouri Botanical Garden are arranged, from showy flowers, as in Bennet- sed b y (1990), a (semiherbaceous abit bt ih һар explain » why ‘he ї 1 1 p x 1 Ё 1 dekloeftia, Dechellyia, {рери Сгапе, 1988; уап nenburg-van Cittert, 1992). no possibility is «m t eec on the ge- netic ded of floral development in angiosperms ig sd might indirectly Зен among for g aa data is ор сеш rch or seed records, and vegetative remains 4 9 genio in E mag like Аса sce хош (a t) and Үш bos (an aquatic 1 F & Hic key, 1976). Pollen of Nig es = one is гене apomorphic іп being ve oj small, respectively, but 2 iso I atl ACI than granular structure (although t the | жейде are wi hard to recognize cf. Osborn et al., ) and a complete tectum. g this is the Бы polle: TP for angiosperms, it would be difficult impossible to recognize in the dispersed The apparent conflicts among present molecular data sett raise tbe qued sm mar Molecular da ta lem Act. Donoghue et Lal, 1989), although methods (as discussed by Albert et al., 1994) or discoxety d Iwabe et al., ; Ra Шо оп & Jansen, 1992) might permit Pion inferences. It is possible that signif- icant progress on the origin of angiospe rms will le ading to the group. Potential a ن‎ include ‚ a Jura ssic leaf wit rre acters, as required in ees "when ther are linked wit th angios perms— ien. angiosperm нд, Better ter ие ve m — biak риш embers ster: relatives 1 fy whether A of anthophy tes or arose зан а їп з each pm ше; а possibility raised by Late Triassic-Ear big Bos e struc- tures related to Hits groups (Westersheimia, Var- on associated scenarios ‘of floral evo- hon (Doyle, 1 be associated with ip im of us P genes that control development of the angiosperm outer i (Robi Beers et al., 1992). o LITERATURE CITED ALBERT, V. A., A. VERE UND, К. BREMER, M. W. ru T B: MANH , B. D. MISHLER & KE pid 1994. Functional e constraints d Eos Leven 9 land plant phylogeny. Ann. Miss 9. 81: 534-567. 1991. BARRETT, M., M. J. DONOGHUE & E. SOBE а Against consensus. Syst. Zool. 40 pee ai Beck, C. B., R. SCHMID Stelar morphology and the e primary vascular system of seed plan ie Rev. 48: 691- BEHNKE, H.D. 988 Sieve- element к ргоїеїп, luti s; Ш. Mag: noliidae. Taxon 37: +]. 732. he Pye BORNSTEIN, А. J. 1989. omic studies ар. е; acea] e pe edicto. Tod rs of Mexi p be The tral оя Si gad subg. printer Y Arbor. limits of amino o acid sequen нана du c reconstruction. 2 Bae data in angiosperm, npg lution 42: 795-8 te INGHAM, D. A 1993. Pa eges al werner data in phylogenetic analysis. : 384-397. . Ни aad Cw. eem C The Piperita E the er ARN hypotheses for HW VH то! ledonous RN ers Bot: eresy bible The топос! 1 of — origin. Evol. Theory 5: : 345- t theory 189- -225. Volume 81, Number 3 1994 Doyle et al. Integration of Morphological and rRNA Data CARLQUIST, S. 1990. Wood anatomy and relationships Em Amer. J. Bot. 77: 1498-1504. САВР! ELM 992. Random cladistics. Cladistics ск MW 8 41 OTHERS. 1993. Phylogenetics of seed plants: An analysis of nucleotide sequences from the Конна gene rbcL. Ann. Missouri Bot. Сага. 8 CHLONOVA, А. F. & T. D. SUR 1988. Pollen чай ultrastructure of Ctavatipollenites in incisus Chlon two modern species of аана. нер Pollen Spores 30: 29- Corn, E. S. & E. М. MEYEROWITZ 91. the whorls: Ge natio гама controlling flower development. Nature 353: CORNET, B. 1986. The leaf venation aad ee Structures of a Late Triassic angiosperm, Sanm guelia lewisii. Evol. Theory 7: 231-309. ——. 1989a. Late Triassic ime enteric pollen from the Rich жк rift x АША. __Paleeontographica, Abt. 213: 37-8 1989Ь. е reproductive eo and ology of Sanmiguelia lewisii, and its bearing on ue rm evolution in the Late Triassic. "Evol. Trends 51 3: 25- pec P CRANE, Р.В. 1985. joie analysis of seed ars and the prius of uM . Ann. Missouri Bot Gard. 72: 716-7 E 8. Maj г elades and relationships in = her" miel p. 218-272 in C. B. Be (cor Origin and aaa te of Gymnosperms. a lumbia Univ. Press, New bat, id A. 1981. An Integrated System of Classi- з 2 n of үлеп Plants. Columbia Univ. Press, ew 19 do. The Evolution and Classification of DLE ing Plants, ga ed. New York Botanical Gar- den, Bronx, New 6. Syma апей of the An- Wile Sons, N ork. gs тот pe prospects in reconstructing i phyloge eny. Ann. Missouri Bot. zai Gar rd. 81: i De 1989a. Phylogenetic 8 angiosperms i the Adi ips of Hamameli- A Pp. 17-45 in P. R. Crane & fune (lior) Ron ion, Systenaics d You! H of the Hamamelidae, vol. 1. Clarendon Press, rd P iy A. DOYLE. 1989b. hylogenetic studies Of seed plants and angiosperms based on m ical characters. Pp. 181-193 in B. Fernholm, K. remer & Н. died prd The Hiériichy of Life. Elsevier, Amste Mna Trends Ecol. Evol. 6 Door -Angiosperm monophyly. Ae 5 2. The келе. "i те ојесшаг and m Baten Structing plant ph Bie D. E Sub e T T Doa n P. S. New York. s 992. e ымы of е та netic account. Pp. 356-389 R Wyatt (tor, ke and Evolution of Plant New Yo a n: New Appr ia a Chapman & Hall, . Chapman "inel 199 С. OLMSTEAD, J. F. 5мїтн & J. D. PALMER. 2 PHS i relationships of Dipsacales based on rbcL sequences. Ann. Missouri Bot. Gard. 79: 333-345. , J. A. DOYLE, J. GAUTHIER, A. С. KLU А Rowe. 1989. The importance of fossils in phylog- eny reconstruction. Annual Rev. Ecol. Syst. 20: 431- DOYLE, J. A. 1993. Cladistic and paleobotanical per- spectives on the Wed of angiosperm organs. J. Cell. Biochem., Suppl. 1 Seed plant ш: logeny a ad the mental cladistic ی ا‎ cui ev. & 1987. The in elucidating seed plant lution. Rev. Palaeobot. kie 50: 63-95. & . 1990. Reexamination of seed plant and Onde relationships. Amer. J. Bot. 77(6, Su 1. мар ————. 1992. Fossils and wa plant phy- pa reanalyzed. sales 44: 89- ^i oe angio- sperm diversification. Paleobiolo Бу 19: 141 -167. & L. J. Hick 1976. f the mid- rbd oou Potomac c Group a wr pen bearing on early ead erm evolution. Pp. 139-206 in C. itor), Origin and Early Е е of y" mbia Univ. Pres ork. 1991. d. of early en ina en context. Pp. 169- ыл Pollen B. Bec k (ed giosperms. Colu E жеу angiosper 195 in 3. С & S.H. and Spores: Patterns of DBS Clarendon Pros i ford. Duvar, M. R., M. T. CLEGG cipes ;WSD. CLARK, W. J. Kress, Н. G.H Ls, L. E. сая TIES , B. S. GaU TE: үз ныт &.C. ; LEARN, JR. 1993 ннан hypotheses for the monocotyledons constructed age L nce data. Ann. Missouri Bot. Gard. = pe ater F. А 1 tionary significance of chromosomal differentiation patterns in gymno- sperms eem primitive angiosperms. Pp. 220-240 in C. B. Beck (e oe Origi in and =ч cig ay of ceae, Tdio- . Naturwiss. The Chloranthaceae: Reproductive structures and рант position. Bot. Jahrb. Syst. 109: cd 226 FELSENSTEIN, J. 197 Cases in which parsimony or compatibili 1 methode will be positively misleading. Syst. Zool. 2 1985. Ta Games ‘limits on phylogenies: An approach using the bootstrap. Evolution 39: 783- — —— & Н. KisuiNo. 1993. Is there something wron with the ано оп selec ies? A reply to Hillis 3-20 0. and Bull. Sys rie FRIEDMAN, W. Е Eph dra (Ep ) Farther evidence ‹ ч doubl fl nce of a pre-angiosperm Nn of endosperm m: Im те for the « т of flow ering plants. Science 255: 336-3 Annals of the Missouri Botanical Garden GABARAYEVA, N. I awry of development in primitive рь 29175 imiti 268 i Blackmore ARES ditors), Pollen and Spores Patterns of Diversification. Clarendon Press, Oxfor GiBBS, R. аше reaction, lignin, and the relationships between woody plants. Pp. 269-312 . V. Thimann (editor), The Pide of Fore Trees. Ronald Press, ges York. GIFFORD, E. M. & A. S. Fos 1989. EAT = Pan of Wai Plants. W. H. Fre Cnusser, L ni & NowIckE. 1992. "coe . 8t B . ZURAWSKL. las sequence from a Miocene Magnolia mo ore 344: 656-658. HAMBY, R. K. & E. A. ZIMMER. 1992. Ribosomal RNA asa phylogenetic tool in plant systematics. Pp. 50- 91 in P. S. Soltis, D. E. Soltis & J. J. Doyle XH Molecular Systematics of Plants. Chapman & Hall, New York. o SE be 'ggm o 5 A cladistic analysis of conifers: НАЗЕВЕ, M., К . IWATSU & UEDA. j Phyloge eny “of gymnosperms inferred from rbcL gene sequences. Bot. Mag. ( kyo) 105: 673-6 ILLIS, D. М. 1987. Mole cular versus oe ee to systematics. Annual Rev. Ecol. Sys 18: * Н 3 asas empirical test 4 bootstrapping а ethod for assessing confiden in умло йү analysis e Biol. 42: 182-1 192. Hucues, N. F. & A. B. McDo 990. Barremian- Aptian angiospermid p ii from southern England. Rev qoidam "Pa Н по]. 65: 145-151. Iro, M. 1987. Phylogenetic systematics = the Nym- phaeales. Bot. Mag (Tokyo) 100: 17-3 Iware, N., K. Kuma, MIYATA. SEGAWA, S. erem 19 Evolutionary и chaebacteria, eubacteria, and eukar aryotes interred from tee trees of poire Car . Proc. d. Sci. U.S.A. 86: 9355-9 Natl. Aca S. KELLOGG, E. A & C. S. CAMPBELL И енені ттан of the Gramineae. 310-322 in T. R. . Ca ampbell & M. Systematics and Evolu- tion. Smithsonian Institution, Washington, D.C. J e origin of angiosperms: nd od problems. Trends Ecol. Evol. 6: 215- В, D. К. Gar F. WiwPEE. 1991. Mane evolutionary of ancient aquatic ms. Proc. Natl. Acad. Sci. U.S.A. 88: 81. a characters n Annona ru their о! it » nee: .&D. 1990. lasts чы ^er Spermatophyta. Влада 42: 197- —— ——— & — ——. 1991. Cladistics of the Magnoli- idae. Cladistics 7: 267-998 MappisoN, D. К. of m multiple islands of most-parsimonious trees. Syst. Zool. 40: 315-328. . SWOFFORD. 1992. Geo- graphic origins of human mitochondrial DNA: Phy- logenetic evidence from control region sequences. Syst. Biol. 41 ak .R.M 1992. MacClade: An alysi sis of Phylogen ny nes [nn Evolution, Ver- sion 3. Sinauer Associates, Sunderland, Massachu- setts. METCALFE, C. В. 1987. Anatomy of the Dicotyledons, 2nd ed., vol. III. Mags oliales, Illiciales, and Laurales Ox! MEYEN, S. V. 1 ures of gymn systematics and phylogeny as evidenced by the fossil record. Bot. Rev. 50: 1-112. gip i B. D. 1994. The cladi and morphological data. Amer. y ы n Fu 94: 143-15 2: ONEY, L. L., I. W. BAILEY & B. С. L. Swamy. 1950. The morphology ич relationships v the Monimi- aceae. J. Arnold Аг феб SIS NixoN, K. C. & J. LD 1991. E кше taxa, Meer A Buy "ledit analysis. Cladistics 7: 233-2 et CnEPET, D. STE N & E. M. Fris. friend ү sed Ip phylogeny. Ann. Mena Bot. Ga En OSBORN, J. M., T. N. T. Ton EIDER. К, Pales morphology and ultrastru of the baceae: Correlat with polli дее? quee m" J Buc 78: 1367-1378, Qiu, Y.-L., M. W. Cuase, D. H. Les & С. R. PARKS. " 1993. Mo M cular iw hien of the М eri Cladistic analyses of nucleoti ide е sequences of the [rd tid gene rbcL. Ann. Missouri Bot. Gard. 80: 5 ( times). Syst. QUEIROZ, .A.DE. 1993. Fo Biol. 42: 368-372 . ANSEN. 1992. Chloroplast i n the ancient evolutionary m in vascular land plants. Science уы 1697-1699. , R. E. PRUMT Mere N-BE EERS , К. t sm A — 1- and ja TUM sterile mutants. Pl. Cell 4: 1 mess 9. АБД, 2. New interpretations iin conifers. in Palaeobot. Palynol. SANDERSON, M. NOGHUE. 1989. енти іп levels of homoplasy. Erin De of the is А л дут” Characterizing ‚ & С. W. ROTHWELL. 1992. 4 ost primitive seed ferns: Сн А NIC Ellis polymorpha. Int. J. Pl. Sci in SEWARD, A. C. 1904. Catalogue of Mesozoic oe eun British Museum. The Jurassic pis British Museum (Natural History), pe di STEVENSON, D. Morphology and 5 ard. ST: rs hs Cy Айны! Меш: New York Bot. Hone D. L. 1991. PAUP: Phylogenetic / Volume 81, Number 3 1994 Doyle et al. Integration of Morphological and rRNA Data Using Parsimony, Version 3.0. Illinois Natural His- tory eae y pde, re & ү Re rine ancestral d desc nde parsimony. Math me sci. 87: 199-229. TAKHTAJAN, A. L. 1969. Flowering Plants: Origin and Disp AN Smithsonian ША af shington, D.C. TAYLOR, D. . J. Віскі 1990. An Aptian plant with attached leaves ААЛ: flowers: ei os for angiosperm = Science 247: 702 0 DE j^ e the herbaceous origin of angiosperms. Pl Syst. Evol 180: 137-156. F. MELEKHOVETS, С. M. dr BROVA, K. M. VALIEJO-ROMAN - ANTONOV. 1991. 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Jornvall (et itor) The Hivi of Life. | Elsevier, Amsterdam APPENDIX, палос L CHARACTER ney follow Do sim (1 > for char- ol -50 (except 48) = Donoghue & Doyle (1989a) the ni racters 57-82, unless otherwi ын nine-a — er А Т Br. ^ Axilary buds (0) ) sin, ingle, Pavel iple. distichous py (0) spiral, (1) opposite or whorled, (2) ves (0) all yr yee (1) linear or dichoto- Ж. ау mous plus cataphylls, (2 Papias compound plus i A iri (4) primates tino- or acrodromous) plus cataphylls; ч О) simple pinnate, (1) Шо ог меа (2 pm "ue veined. States 0 and 1 of n ae e Doyle constant iio or lower angle at base, vs. ovat е, sal уз ош crowded, at higher angle) are sabe Bis que 7 г nate; the only ta xa with Donoghu e & Doy 1 Cal only one се sh appears in each of the present data sets. 5. Rachis (0) bifurcate, (1) simple. 6. Laminar venation (0) open, (1) reticulate. minar vein pip (0) one, (1) two or more. cell Stomates (0) an 1) m шетел ог variable, o tetracytic. К Lau T огей as unkno noghue & Doyle (1 9892), but Ба state a Pu ingroup tee assumed hes E 8) is paracytic. Apical meristem (0) уно tunica, (1) with tunica. Stele (0) 1 tele with external sec- ondary xylem only, (2) oL. ios internal secondary lem »* = 12. Primary xylem (0) mesarch, (1) епдагс . Nodes (0) unilacunar, one-trace, (1) КМ (тоге ы а three traces from separate primary xylem unde: arcuate in petiole), (2) unilacunar, two- trace, (3) trilac fro id mass or several arcs of primary xylem, scattered in petiole) was trea ted a as a separate state in state and rescored medullosans ave no effect on the we ded eliminated t wn; this lacunar (Cronquist, EG numbers of traces, but these. are usually for x dune dome Bec k et al., gro cally two-trace. y xylem (0) with apenas pitting in the jan th no scalariform pitting (coniferopsid 15. Secondary xylem (0) with circular bordered pute or perforations only, S with at least some sc Motor itting o M ons ing of angiosperm Metall = 16. aie bsent, (1) present. Donoghue & Doyle (1989a) d prem in the roots only as a third state, 0 scored as unknown, but this +h cter, Pec к їп ably to mera the vessel cell type. To pre- e this information, we ha We redefined the character м express this | abili ity a nocots as 1. t biseriate, (1) at least some multe rtical secretory structures (0) absent, (1) cav- es, CM ca nals. in with (0) no Maule reaction, (0) Maule re- censis analysis: sporophylls simple. Chloranthaceae, core due les, тандай Annals of the Missouri Botanical Garden chaeopteris); E MEDU = Me “соге” паро CHLO = Сога: еае; Hamamelid. ; WIN - Wi interacon 1 DE Fen matrix, extant and fossil taxa & Rothwell (1992) = h CORD = * Magnoliales yeu Degeneria, ane Annonaceae) AUST = ae; EUDI = PROG = “рг чоне. (Aneurophyton, A Шол ‘seed fern” of Doy Cordaitales; CONI X IDe © а Cabnbacae М = isis ? = = 0/1; В = 0/2; tn teca fnm dia is unknown; — — ha н. not Mp e 1 3 122456709012 MSG70901224567830123456705012 45678901224567890123456705012346780U ?00? ? 700000000? ? 700700000 0? 2201221 1-000 07001110?30122100110111 111100001-? amet олда mimm 1101 11111100011 Vr CHLO 1012111141 E OLRSBITA CLAN IR. ies 230122 wu кап 100 111113 ee 10047 111 Mes 1024? 11101113011101002017100707001110730122100110711111177171- 701111111 PIPE TTT ЫСКАК teint 110?30122140110111111101001- 1027111018200101! NYMP 10047111011130707070020171007070011107E0122100110011111111000-70A??1 ТН MONO шг ААЛЫМ ганы ое 20111101700100? e scored as unknown to allow equivalence of their uni- AP gece (0) spiral or in mo e than Bis ovulate earpels ne the condition i in Gnetales; Dee кз (1) in a single whorl. Chloran wee are SC 5 Piperal p n dat set are as d ó allo w equivalence of the three-lo and soured b } Iti ls. droecium of Chlor алш n din whorled microspo 21. Ovu » (1) nal. phylls of Bennettitales a nd. Of 22. Cu шаат, (0) rd lobed, (abes netos 28. Strobili a lacking or "ame (1) compoun Doyle (3) йому ee 1 Aa states in the infl e character of Donoghue & ios anatropous, (2) orthotropou (1989a), spikes nid raceme! cored as poten wih 23. ter ae peer from two appendages hom logous with compound — solitary aye DS vun (1) present. simple strobili; а " sented, except ! 4. Ovules pe ешр cupule or potential homolog 0). emi (1) o Mic енен (0) terminal, marginal, or adaxial, Pi a rangia vary from abaxial to M in angiosperms we have scored th hem as unkno because of bis i mens. It is which eed p 26. чаа (0) free, (1) iei at bei basally. sociation with s 29. ade ri (0) ^: (0) radispernie: insi aas extant di et (Gn une cnn radiospermic t i уе € Integ cip (0) ae ^d) with sclerotest@ SP us sta. Sina a tines with (0) lagenostome, rue chambe Angiosperté UN — le ы see (1) tubular. Ang 2), red e : oyle & Donoghue au ve rescor ч аз their ae are so ван) but we һа (1) simple Volume 81, Number 3 Doyle et al. 1994 Integration of Morphological and rRNA Data BLE 2. Morphological matrix, extant taxa. CYCA — Cees SORS idu CONI » Coniferales; EPHE = aii: WELW = Welwitschia; GNET = Gnetum; MAG “соге” egeneria, Myristicaceae, Annonaceae); WINT = Winteraceae; CHLO = a б CALY = Cal canthaceae LAUR = "core" Laurales (Mortonia, Monimiaceae, Atherospermataceae, Siparunaceae, Gomortega, Hernandiaceae, L P= ceae); SAUR = Saururaceae; PIPE = Piperaceae; ARIS = Aristolochiaceae; N = Nymphaeales Lj cava ini Cabombaceae); RANU = Ranunculidae; TROC = Сры DR es; Bo гуш ? — character state unknown = = character not included; A = 0/1; В = 0/2; С = 1/2; D = 1/3; Е = i 2 4 5 6 12345 doge beoe apes 1 34567890123456789012345678901234567800123456789012 CYCA -?00-0-000- -10001201000-1000-1-0-00- - -00-00000000000000000000000? ?? ?010000?? ?? ???? GINK -001- тнт -0-00- - -00-00000000000000000000000? ? ? 2010000? ??????1 CONI -001-0-000--01000200000-100? -0-0-00- --00-00100000100000000000000? ?? ? 40000? ??????1 EPHE -111-0-001--21011010101-0111-0-1-01---11-10100010100000000000000? ???010?00???????0 WELW -111-1-010--21011210101-0111-0-1-00---11-11111111100000000100000? ???010000? ??????? СМЕТ -110-1-011--11?11010101-0?11-0-1-01---1?-11111111100000000700000????120?10???????1 MAGN -020-1-111--D0111011010-?100-0-0-10---10- a RAT DOT, 000 WINT - 1--30101011010-?100-0-0-10---20-2210011011111110000 11001 CHLO -010-1-121--2011101?020-?1?1- 10---20 2 ioeiip7iiiaai7eeis?1277611111201178000 CALY -010-1-111--20111011010-?100-0-0-1?---20-22100170?11111 3000? ? LAUR -01B-1-111--2011101?010-?100-0-0-11---10-22100110?111111000 111 000? SAUR -022-1-131--10?1?011020-?101-0-0-10 100110?111 10010102?1112010200010111 PIPE -022-1-131--10111012020-?101-0-0-10---?0-2211011011111?10??01?102?1112011?07210111 ARIS -022-1-101--30111011010-?100-0-0-10- - -20-22100110?111111??121??011111110071001! NYMP -002-1- 30?0?0?1010-?100-0-0-10---C0-221001100111111110000?0A ?? 100000200D11011 RANU -002-1-101 11011010-?100 -12---20-2210011011111110?00010101??11?1?200??1 TROC -002- -121--30101011010-?101-0-0-12---20-22100110?1 ? 1 20107 МОМО -077- L-1?1--10?1?011010-?10? -0-0-10- --20- 2210011071111111102007011110170010077? them normal, since they certainly have nothing closely 39. Exine structure (0) spongy alveolar, (1) honey- comparable to the gnetalian—bennettitalian state. ir alveolar, (2) granular, (3) columellar; extant anal- 33. Nucell vascularize d s: (0) alveolar, (1) grsnular, E ones ellar. Unordered Е haeales are scored as as (inner) integument (0) fused most of the way diced because Osborn et i (1991) reported that теде to the nucellus and (1 hi ~ Cabo mbac eae are colum ellar and (1) free nearly to the base, whic ставы (1) présent. Doyle ems to have the same distribution but is better doc 40. mented in extant tax —— (1992) scored Gnetum ar avai ‚ on the 35. Pollen with (0) tetrad scar, se — (2)noap- gro hat any striations erre, (3) three ^ (I 1) no ve pe ve been lost; this is now suppor тш А ultrastruc- «m eH ыж - i. if as having a tural observations that the tectal spines e- tetrad scar in the nine- -angiosperm data set, bas ao ‘on semble striations of Ephedra and Welwitschia (Gillespie primitive fossi , but as sulcate in the extant & Nowicke, 1 ta set. The disulculate pollen us Calycanthaceae i is scored 41. —— tetrad (0) апей, (1) linear. as unknown to allow RAY BEDE RATES 4.42 gaspore wall (0 (1) thin, (2) absent (or- inaperturate; they w inn state 1 in Donoghue & ered). уз: эвде of dé carat is in keeping with the Doyle (1989a), but this pon included sulcate, inapertur- ee of thin e absent as one state in Doyle & Ме, and sulculate. Donoghue (1986, 1992). ange ollen symmetry (0) radial, а) — Аз in 43. Microga аточ ( (0) more than four nuclei, “Эз Don hue (1992), we score Gnetum as unknown, (1) four nuclei, (2) three ei In Doyle & Donoghue i еп en has global rather than radial or bilateral (19 е Feri nucleate state of war aoa Rd е also score angiosperms with radial pollen was ted a ngiosperms were scored as unknow ^ Эд not only because their symmetry is unlikely (to a doy. ун eri wein from either state), but this state is a ж homologous with radial symmetry in the sporelike pea synapomorphy of angiosperms in the present А en of primitive seed plants, but also because it is dat sete ne with other characters included in the matrix: 14. учен їгапзїег (0) spe mete (1) sip wen j^ . 26 Shape in Winteraceae, three colpi in eudicots. mous. Conifers are scored as zooidogamous in the nine ninformative in the extant data set. геты: data set, t; ba доа оп lack of a sulcus in primitive п (0) nonsaccate or subsaccate, (1) saccate. f о) Doyle = оле, 1992), but from кеке), (1) p Mesi "mo Mrs € oe serie кн ы aee (0) — MÀ (1) tetrasporic. tectum to be; 46. Egg (0) cellular, (1) free-nuclea Annals of the Missouri Botanical Garden 47. Early embryogenesis (0) free-nuclear, (1) cellular. 48. Fertilization (0) sin mol. M double. See discussion in text. Calycanthaceae lack d & Stevenson, 1991), Cheese h apo: c4 cun 1966) ип. уо (0) without feder Ow eeder. Sond germination е eal, ^ зей Ап- 80) and Endress (1983); contr trary to Loconte & Stevenson (1991), that аена de ompani ies s in phloem (0) absent, ۵ ee 52. Mierosporangia (0) various, (1) in 93: 54. Pollen germination (0) in see рг: E (1) on stigma. ndress reports 55. er SPEI AL S Bu (1) eight-nucleate. e larger едын bryo sacs of E жле аге ore ugh they show eig 56. Endosperm (0) absent m ) present. 57. Radicle (0) persistent, (1) replaced ni adventitious roots. 58. Habit (0) asia ses herbaceous. Groups with in- tetsu eines roducing normal secondary xylem are score 59. E d-wall ae or n perforations (0) multiple, (1) sim PUN Fy ] sle (fM ) PI type, ( (2) PII xt for addition of PII AN scoring of m “ E see (Bebnke, 1988). Core Tareas were scorei unknown in Donoghue & Doyle ü 989a), but sem | state s the i SPI 61. kun (0) ab: assumed h t, (1) pres dks хое ak = ids 10) absent, (1) pres- t. Un здеп ngiosperm data se Tas Stipule = 0) чыш, (1) ately. Other stip- ule types are scored as unknow. 64. Chloranthoid vum on leaf margins (0) absent, (1) derart 65. nth (0) more than two whorls (or spiral or BUE Q) two whorls, (2) absent. Donog hue (1989a) scored Trochod e becau: tate i n the present data wW e group as po 66. Peria АШ. amas try (0) various, (1) at least calyx ege Stamen number (0 (0 ) various, (1) 68. Stamens (0) laminar, (1) with е aan fila we n (0) boat-shaped, (1) globose. сане with saccate 3 рана гта е pollen scored as unk because it is unclear whether their shape conditions can be com- pared w ith those in nonsaccate, basically monosulcate groups. 70. Poll ш (< 20 um) (ord sed only in t (0 1 A“? um), (1) medium, = ered). ‘Prev this character w. m stu ла age ue & Doyle, 9a). assi (except Caytanid) аз uncertain (0/ ids "m ons. First, I ly larger and more suggesting that 55 F © f | lati ab- sence of sacs and size. If so. scoring saccate groups on the angiosperm-based ‘ines саде of Donoghue & Doyle ight excessively weight transitions between saccate and nonsaccate and pr dien spurious groupings among non- angiospermous taxa. Second, it is possible kn. the more ар propriate comparison is between si ze of nonsaccate ро]. ] (nexine) in saccate pollen, which is often much smaller tha an total grain size. . Tectum 0) бош ацша! ог r finely on pret: de reticulate state te- ү! apes ghue & Do yle (1 on reticulate groups than nely жиы rA — 1a Aperture EE (О) (шеле smooth, a ind ut Conifers are score ЧА de. b sperm data set beca ee Res imitive forms. Scoring of Arisclochiaceae i is giar, on 1 SEM inh of Saruma pollen kindly provided by Long Huo (Guang- zhou); scoring of ^ peas i is based on Bornstein (1989) an S wx © ш @ Do © Su tect spines (0) absent, (1) present. Scor- "i s based on Bornstein (1989) and Doyle MN thick, laminated, (1) absent, (2) thin, nonlaminated: , except ї under apertures. Unorder: е б rea- sons di: Fall Chl (1988), we have Жаы Chloranthaceae as hav exine, not u s in s oghue Me re le iP 42 Hypanthium "de t, (1) pr A УШЫ, ene rpel (0) several t one apical. йа state in Donoghue & чен a 89а), one basal, is are therefore scored as unknown Là pepe iH Maio 2 drupe (with E doc on d by mpl е Stevenson (19 91), th к ong "Geni " dehiscent fruits of em E Pusat , scored as dehiscent Vos P — & Doyle NE а better characters M as berr: УК ae oghue & Doyle (19896), fee e rt hee which appears to v ied ares ич family (Vink, 1988), we have rescor as havin Tries. " Testa (0) det heer ein (1) non- c 1) 1 1) sclerot е da peas mnl (eio mus might bec con- redun dant with 8,() n7 ertain aec is nd ех origin, we have rede Р ibe states and scored taxa with n > 16 as u mU third state in Donog yle (19893), н ge is ени iden by nthaceae, whic чац scot nown. Dat pete Ehren менн ). aracters from ianacrens: 1 еа n юув. 132; 6: од 7: 176: 8: 181; 9: 9: 180101 = Fl: eotide not present in soy (betwee: 220.18 . 240; 12: 200; т 220; 14: 222; 15: 236; 16: 18: 241; 19: 242; 20: 244; Volume 81, Number 3 Doyle et al. 1994 Integration of Morphological and rRNA Data TABLE 3. Ribosomal RNA matrix. Order of taxa as in morphological matrix. R = A/G; Y = C/ n M A/C; K = G/T; S = C/G; W = A/T; H = A/C/T; B = C/G/T; V = A/C/G; D = A/G/T; N = A/C/G 12345678901234567890123456789012345678901234567891234567890123436789012345678901234567 TMCASGATAGCYGACACCATATTGCCCRGTTTTTGCCTGTGCGTAATACATTAGCTTAG AATTAGCTTA? GYCYWRCTCTTTC CETGCCTACCCTECACGATA?PTEATACCETTTOCTCGTCOCGECT?27777AATMMATIAGCTIN GATT! GCCCTWGTTC? TCATCGTGTTGAGCCGTCACACTGCA ?2AGGGTTTCCCGGT T TCGCAGTTGATT????? ? TACGCGCCAATTTGG TTTG AGT?CGACT TC?TTGCTTGC CCATACCAC? ?TATGACT?CC C TT CIATATCECCTCTAACCTACTCTCTGATAGTGCGTTGGCCCAT AKCKSGMCTC?YYCCNTCMYBAYAYHGYTCCGCCACTGACGAYNCCATAYCGCTCCYRNCCCGCTCTCKGGTARTRCGTTGGYCCAG 2 3 6 89012 a RUN E EE AU UU O TCGCTCTATATTAG2CCCCRATCGCCTATTWACCTCGCGYCTATGY CTGGCRYTCTWYRTMCGYSSGGGTTAGGGTGATGGERGARR CGAATTGCGCCA TICTRCCTCGTECCACECTECTOCCHACGCA7YCGTCATATACTGCGTATTTCTACCGA GYARTAAGRGAGG Be MINGOR CCG GTATTCTGCGCGCCTCGGCCGGTGTGATAM 2 CAGE TCGAATTGCGC cnc G22???22??? TAGGACGGCGCTGCGC? 22222222?222?22??22?2?2??? ? ? GGCCGGTGTGATAGG??? 2??RWYTGCG MVG??CT cG K rn A A TCGGTCGYCGGA «cac ACE 9521: 245; 22: 247; 23: 251; 24: 257; 25: 263; 26: 1: 759; 92: 769; 93: 784; 94: 790; 95: 791; 96: 215; 27: 276; 28: 280; 29: 281; ; 797; 97: 830; 98: 834; 99: 865; 100: 866 31: 287; 32: 339; 33: 347; 34: 353; 35: 368; 36: 01: 904; 102: 1602; 103: 1612; 104: 1620; 105: 429; 37: 496; 38: 542; 39: 936; 40: 1042; 621; 196: 1624; 107: 1637; 108: 1639; 109: 1650; 165 > : ; 48: í ; 90: 0; 11: tr 112: 1662; 113: 1663; 114: 1683; 115: : 1241; 52: 1245; 53: 1300; 54: 1355; 55: 1357; 17 к 5 1708; 117: 1712; 118: 1716; 119: 1723; ; $ : : Й „ : : 66; 120: 6 : 1370; 62: 1371; 63: 1372; 64: 1376; 65: 1404; 3 dT; 122: 1758; 123: 1760; 124: 1764; 12 Буз 411; 67: 1503; 68: 1514; 69: 1526; 7 : 1527; 1777; 126: 1783; 127: 1796; 128: 1949; 129: Meo: Fo E 1528; 72: 1534; 73: 1555; 74: 1564; 75: 1566; 130: 1951; 76: 1568; 77: 1573; 78: 1606; 79: 1613; 80: 1666; 31: 1958; 132: 1960; 133: 1961; 134: 1962; E 81: 1668; 82: 167 7; 83: 1724; 84: 1729; 85: 1735; 1969; 136: 1971; 137: 1973; 138: 1980; 139: 1982; y т 3: 2000; 144: 2001; 145: 167: Characters f bunit, in terms 41: 1986; 142: 1991; 14 Jor ан from the 265 subunit, in teri iagoa 146: 2081; 147; 2032; 148: 2037; 149: 2040; 87: 740; 88: 741; 89: 748; 90: 750; 150: 2041; Annals of the Missouri Botanical Garden 151: 2056; 152: 2057; 153: 2058; 154: 2059; 155: 2060; 156 2061; 157: 2066; 158: 2072; 159: 2073; : 2077 T6172 078; 162: 2085; 163: 2086; 164: 2088; 165: 2089; 166: 2098; 167: 2102. 168-174: Insertion-deletion events, where gap — 168: ev 1 between 131-132 in cud vod in "hs 2 at soy 454; 170: indel 8 between 15 in soy; 171: indel 9 between 1593-1594 in ie ТА їпде1 т between 768-769 in rice; 173: indel 11 between bus 1750 in rice; 174: indel 12 between 1756-1758 ADDENDUM VOUCHERS OF SPECIMENS USED IN rRNA ANALYSES ence я ае А. gigantea Mart. & Zucc.): у. greenhouse, provided by J. Wen- à tl, z 941 "US. Arundinaria gigantea (Walter) Muehlenb.: Baton Rouge ish, с Asimina RS (D) Dunal; Burden Plantati ea Baton y D. Bryne, Z-8-89, LSU retum, Bato: Chloranthus сая (Thunb.) 2. ee Botany gre enhou e B81 1-804, Doyle 94 3, DAV. l ta (L.) Schott var. antiquorum I Du & bed LSU campus, coll. L. Sim 5, LSU. pud. revoluta (Thunb. ): | LSU се Z-2-85, LSU. Echinodo F. Givens, Z-2-88, LSU. M ferox Bertol. f.: Univ. AT greenhouse, vid ed by D. аан, a MES e 888, Doyle Ephedra fsveediana СА Ne Davis Arboretum A67- 620, Doyle 94-2.09. Yeas Euryale ferox Salisb.: D. i „ CONN Ginkgo biloba L.: LSU c UA "coll. L. Sims, Z-3-85, е тотап m Markgr.: Davis Botany greenhouse B70-116, Doyle 9 bee 2-09-1, DAV. d i, New Caledonia, L. Thien 600, NO. Hosta japónica Tratt.: Baton Rouge, coll. L. Sims, Z-17- Juniperus ashei Buchholz: Travis Co., TX, coll. J. Drost, Z-22-87, LSU. ыыы tulipifera L.: Baton Rouge, coll. С. Knaak, Z-5-89, Magnolia grandiflora U campus, Z-4-85, LSU. Najas ur WIS E Magnus: coll. C. "Knaak, Z-1-89, LSU Nuphar luteum (L.) Sibth. & Sm. subsp. macrophyllum m all) Beal: Jefferson Co., MO, P. H. Raven 27204, [текеге odorata Ait.: Tammany Parish, LA, coll. Ё. Givens, Z-21-87, LSU. Persea d (L. biis b Arboretum, Baton Rou oll. C. Knaak, Z LSU. Piper nigrum ee Towa State бам. pilo provided 439, Ranunculus асгіѕ їй, Жз E Sims, Z3 6, LSU. Sabal minor (Jacq.) Per ae Pannen. Baton Rouge, coll. yi Diod 1:208 m Жене lancifo a L.: i coll. E. Jupe & E.A i enm 2 13- 6, LSU. Saruma henryi Oliver: Us dado Arboretum 49482, . J. Kress, Z-94-4, TARN cernuus L.: San Gabriel Parish, LA, coll. R. Chapman, Z-3-88, 8 х ы Sebi & Zucc.: Taiwan, S.-M. ila MN рее: pho aA : Univ. Illinois greenhouse, pro- vided by D. Niki m 94-5 — ottonis Miq.: Univ. Illinois greenhouse, provided y D. Nickrent, Z- 94. 6:109: SPECIMENS WITHOUT KNOWN VOUCHERS Avena sativa L.: seeds provided: by S. Roux, Univ. Т Cabomba caroliniana A. Gray: San Marcos River, X provided by E. Schne "x Cryptomeria japonica (L.f.) D. Don: Mellingberg a Drimys winteri Forster & Forster ^: Berkeley Botani den 45.307, provided pe à Affolter. : tivar malaya," seeds pro- vided by . Saghai-Maroof. j А Oryza sativa L.: cultiva ia etic Louisiana Rice Re- search Station. Pied e Univ. Illinois greenhouse, provided by D. Nickre Pinus iie del seeds provided by O. Stubbs, Louisiana ept. Wildlife & Forestry. ا‎ E L.: provided by P. Hoch, Missouri Bo- anical Garden. its 0 ви, officinarum L.: line Cavengerie, prov amann, LSU. опи а88 So rghum bicolor (L. IE аме; на Тх 4 че provided Ьу М. Th mas s-Compton, Univ. ‘Neti Tasmannia lanceolata (Po ir.) A АС. ее pe pes tanical | Garden 00 0052, ete by si det ` provided by К. ا‎ re Me ш» aestivum L.: line HW 3022 Rohn and Haas eeds, provided by S. G. Bartlett, 150. d Welwitschia mirabilis Hook. f.: Huntington Botan! Gardens, J. Folsom 13, provided by E. Zea mays L.: : cultivar B73, Pioneer HiBr PHYLOGENETIC Brent D. egent e TUE A. Lewis,? RELATIONSHIPS OF THE i EN ALCAE" AND David А Garbary,* “BRYOPHYTES’”! Charles F. Dehoiohs? Frederick W. Zechman,? Thomas antz,"'" and Russell К Chapman? ABSTRACT assical morphological characters and newly described th tes t algae an of the polymerase chain Considerable progress has been made recently, based. on d ultrastructural features, in understanding the bryophytes. сри, eee "advances in 1 molecular biology, len the advent ү (РСВ), Һа = pecially E TRNA ne sequences (both the 1 e present embong cladistic paa асы of the green за that combine ne and с е tracheop hy e large and small S) from the nucleus and the chlo roplast. ompare ‹ available morphological and certain conclusions molecu lar da ta f j hyleti ntaining the *charophyte" green algae and the land р (i.e., “bryo рһуте other отб аа the bulk of ым е delimited “ргееп algae" Cade nearer m. toph ye. (2) The la nd plants. are pin rted monophyletic Pd but neither the specific outgr for among basa lineages of land plants are clear. In ee шашы, rehin! the combined molecular and cee È cal analysis) the three major pas (1.е., liverworts, hornworts, and mosses) арреа o be paraphyletic with respect to t the tracheophytes, with an indication that the mosses qos pend pues of the tracheophytes; however, in may be the зай E the “bryophytes” are supporte ed as a dete ‘group: (3) The ulvophytes, chlorophyt es, and urastrophvtes siia classificatio on) with the topology: [ulvophytes [chlorophytes + кыно | Combined analyses эн molecular and morphological data offer the greatest potential for resolving these relationships taxa that may Considerable morphological and ultrastructural Reconstruction of the broad-scale phylogenetic о їа һауе ш оуег the last two UNA es relations "m of green plants is оца to our da erstanding of ros evol ry events su ded bear on th ticell Rar diversification о of the sn algae and ШО to the istory area and the соп f land RE E на (e.g., Stewart & Mattox, 1975; He- ( 985; ler & Churchill, 1 йы! ant, 1977; urat ЖД io. Crisi Sent. ler, 1980 d rown & Lemmon, 1 ; d relationships is necessary for purpose ШОР co mparison in studies of е, Phylogeny (Crane, 1990; Gensel, 1992). 1988; Duc lia, 1988; Ligrone y: Gambardella, 1988). More re- cently, comparative molecular data have becom ' We thank Richard Zander and Robe: Wyatt for th -— We are grateful to Angel Maden КЕ information on куруга опа e їп Lycop dium obscurum. "This search was supported in part by рея BSR-9107484 to ВОМ; DEB-9207646 to KSR - DJG; BSR- 8918564 t to MAB and RLC; DEB-9107389 2 үз ment of Botany, ut University, Durham, North Carolina 27708-0338, U.S.A nd University and Jepson rir "University of California, + oust culty of Biological Science, Univers sity of Tulsa, Tulsa, Oklahoma 74104-3189, U.S 03 "nea of Biological Sciences, East Tennessee State University, Box 70703, Тороп jm Tennessee 37601- * Department of Biology, St. Francis Xavier University, Antigonish, bye Scotia, Canada В2С 1C0. z má y of Natural Sciences, 1900 Benjamin Franklin Parkway, Philadelphia, Pennsylyani partment of Botany, Louisiana State University, Baton Rouge, Louisiana resent address: Department of Biology, Coastal Carolina University, Conway аа Carolina 29526, U.S.A. ANN. Missouri Bor. Garb. 81: 451-483. 1994. 7 Department of Bio ology, Indiana University, Bloomington, "eva iana 47405 ао USA Be а . , U.S.A. A Annals of th Missouri Pini Garden EN C , Kantz ч al., 1990; Zechman wis et al., 1992; Mishler et al., 1992; € et al., 1992 "n et al., 1990; Levis stic studies to date Быга ies d ihe gree S ips" nor the oph phyletic. The green ve dies to а ihre fr q h 1 20. f DA E | “ т арена od Roe Mica of = ica зам ана Pleurast rer and Ulvophyceae sensu Mattox & Stewart, 1984). There is an in- dication М the шуо] ls t: are tm to the chlo- rophytes p oned the toodophyly of the ulvoph be- ack of morphological сыш үй to unite ы: roup. This result was supported by a or molecular study pon a al, 1990) а ray have’ beck treated in A died s: (1) a e sister class to the Chloro- п the Chlorophyceae part of the Ulvophycea e Sinan e other m eage of сеа ln includes the Кы Ен green is plus the land plan (се itin пу s plus шалыс The genus 17900 E (3) as to be the cl f land ERES (Graham et al., "180%. The ‚дк чү аге com- of e trach то] ; Waters et al., 1992), apes Fm es, "T tra- logical th r group of tracheophytes. Th as the sister e phylo- genetic vus of the re is not clearly resolved by any o of the ү publish sets. On ы оїһег band; monophyly of the bryophytes (includ ing ee nein was e by a recent cladistic analysis based on sperm ultrastructural data (Gar- эш, et al., Molecular sequence data have shown consid. ph ylogenetic analysis, but they у optimisti li theoretical considera uence char: € (given their quasi-clocklike ue lution and limited number of cha racier states) coul be esp сапу. problematica i in p E 7 рене oth branch ies exists (Felsenstein, 1978; Mishler sulting data s mpa ы: pcd sets of ти ае state че. ing; Miyam 1985; Kluge, Albert & Mishler, 1992; вА et al., 1992; bert et al., 1993) 199 ‚ 1992); апа (3) representing diverse, yet clearly x thod monophyletie clades (e.g., exemplar meth vs. partmentalization" —a new ch ; m jx: tuting an inferred “archetype à hypothetical ancestor for a clade accepted Jj irm i phyletic a priori in an inclusive analysis: it is evident tha ons ferent a are erie сери, at al hierarchical lev: d fully et al., ; Waters et al., 1992; 1992). ae de other e the 5S rRNA Ar SPENS gene appears to be too small, rRNA gene ba m these two genes d Volume 81, Number 3 1994 Mishler е “Green A and “Bryophytes” of some previous morphologically based Maias about the relative ignis order of major lineages of land plants (Mishler et al., 1992). In contrast, ai protein-coding gene rbcL has proven too divergent to be useful iia these lineages rt, ben omm.; Mishler, unpublished data), but it та оны сопзїзїепї учее сһагасїегз within major lineages. For example, within the mos es (with th е bas eae within) m mon AID a n Зур mosses (Mishler, unpublished ta). likely that a robust and ады resolved phylogeny can be produced for the n plants in the next few Sequence data uclear and chloroplast genomes. Careful choice p кин апа appli- cation of proper B. be essen- hal, however. The co-authors dp no imously on the homology of all characters as used here, but have found cladistic analysis to obe hoy, to › frame arguments objectiv ety. This E as a ue (and a target) for ongoing projects in I TE are crucial for future research. The с e MacCLADE files will be seat d on request) vele thus, will provide a basis for future synthetic MATERIALS AND METHODS DATA SETS m na sets were assembled in pairs (focused iin erent phylogenetic levels): two ho et m one lar rge and one small molec ular di. two combined dat s. The sources qa teristics of these reig sets are listed below urposes (see Plants sentative land acters bein [o ve а. Sn: veut a ae and independence (Mishler, 1994); a list of char- ound in Table d & Stewart (1984), Mishler А Churchill (1984, 85), and Graham et al. (19 All 110 char- acters were treated as unorder n А cabin ш Mm coded for root ing purposes, based o The data matrix i s shown in Table 2. : (2) ` more focused land plant. перрі! data set (LP-MORP scored E Garbary et al. (1993) for S onda characters of spermatozoids. A list o d character states can be found i n шу Mais set of Шер 65), we added а set of “general шо шы Сы" e characters (num- bers 66-113), beginning with the Mishler & Chur a modified and (26S) sequen Un Kantz et "i « (1990) oes et al. (1990 . OTUs ebd able 5. xcluded. Those taxa that seemed quen repetitive at this eae level (i.e., multiple, me genus) w take matrix had ЕЕ т including Emiliana me m 1833 are Eds 185 and 346 аге from 26S п. А number of OTUs have considerable this r region wore с) with a question mark. To be conser- vative, gaps w coded as missi 18S data, 17 regions (totaling 206 positions) were yi fro ysis beca unequivocally. The data set is too large to publish, bu t m aligned and — PAUP (4) A smaller “land plant only" molecular - data set (LP-MOLEC) was ex xcerpted from the larger set above (GP-MOLEC); the 16 OTUs (including Annals of the Missouri Botanical Garden TABLE 1 T Les adhue +, 4 hl +, characters and Table 2 for the data matrix. states used for data matrix GP-MORPH. See text for source of these . Habitat of free-living abu stage: 0, freshwater; 1, bracki sh or marine; 2, terrestrial. 2. Life history: 0, haplon m B: E 3 @ = д 28.9 E B zE Ex elie: 0, no; 1, yes. occoid; 1, multicellular; 2, coenobic. . Vegetative cells contiguous in mice mne 0, no; 1, yes. - Multinucleate vegetative cells: - Coenocytic: 0, no; 1, . Distromatic Hu thalli б, беи м КИМҮ, . Plasmodesm: A эме. ; pre . Parenchym T resen with acuminate tips: es. Y cette E or ridi spindle- sha aped: 0, no; 1, yes. . Zoospores аро 1, pre ; 2, present, flattened. TA amete produc tion: 0, тушы H ао дае 0, по; 1, вех: 0, i Е > my ч Б GD 22975: Е da es. ve mai 1, anisogamy; 2, By e of Slee Š Chloroplast shape: cup; 1, reticulated; 2, later яа 3, H-shaped; 4, bi-polar; 5, f tic; iy diplontic; 2, чег n NE 3, heteromorphic ael - Vegetative cell or thallus attached to substrate: 0, no; 1, yes uev an Gna nee у sg l, yes, unbranched; 2, yes, branched; 3, yes, multi-axial. . Filam omplete rings; n multiple disks; 8, spiral; 9, stellate; 10, plate; 11, axile. ше noids: 0, absent; 1, present. raverse pyreno oid: 0, no; 1 , yes. pores: "n 2:1,4: 2, 1; 3, 4+; 4, O. g division: 0, mi Е ipe. Oaks Ha APA S Е Ф, w РЕ < о d M asal bodies distant via m dee in ee ee 0, no; 1, yes. lagella extend to o right А шой cells: 0, no; 1, ye 5. lagellar i sym y: 0, no; 1, yes. bsolute orientation: 0, к. es cui B осм, 2. diet дыл asal body overlap in motile cells: 0, absent; 1, pre: . Basal d re connection: 0, absent; 1, p . Mitotic spindle type: 0, metacentric; 1, cen I] = E pindle lpn at ‘telophase: 0, вор; 1, рен, ч uppin rotubules sur d cent durin no; 1, Ма : Panse AA forming i in plane of cell Ert 0, absent; i, present, D PI I M E cA Rr TE MP АШ y veget pos 0, no; E yes. s yes m II light harbour i oes 0, low molecular we e ie ote MAL ed: d ad porulation: 0, a м. " SE [vs. sporulation} 0) no; 1, йа ight; 1, high molecular weight. es 0, n d yes. on vegetative oe 0, no; 1, e pillae Coyne c ell w. ys Ке ; 1, yes tig : uà i hae ie vacuoles: 0, ы 2+; 2, 1; 3, absent. \pical debeas of flagella: 0, no; m sporangia disi no; l, y oosporangia operculate: 0, no; 1, үк, sporangia ч plu 3 l, yes. me AD AQ AQ N NE ‚ yes. rea Eis e piod 0, no; 1, yes. bodie 0, angled; 1, perpendicular; 2, parallel. lagellar am 0, telling. ae latin, ig; 1, Жып stroke. Volume 81, Number 3 1994 Mishler e 455 “Green fee and ‘‘Bryophytes”’ TABLE 1. Continued. . Terminal cap: 0, absent; 1, p ded 2, Eg Prominent proximal s she ath: ч» : 0, n С Transverse фо. % С dE reel. . Proximal septu resent. absent; 1, present. a 1, present. я іп mile cell: 0, absent; 1, present. : Specialized тее ngia: 0, absent; 1, present. А геѕепі: 0, по; 1, 4 f me п pd 0; f Bo um associated ith ow cells: O, no; 1, yes. 0, no; 1, yes. : rat сей хыя 0, еқ 1, уез. д ES. ids: 0, no; 1, yes. - Oil bodies: 0, no; 1, y | Methionine ЛУ RET 0, no; l, yes. ‚по; 1, Я ical division of zygote: 0, no; 1, yes. . Pseudoelat ‚ no; 1, yes - Xylem: 0, no; es. 4 Pii оет: [1 по; Й уез. ine o s: 0, no; 1, yes. rial sporophyte axis: 0, no; l, yes. olumel , n0; к yes. К: rhizoids: 0 ; 1, ves оп game tophyte moss pe 0, no; 1, yes. 3 Articulated peristome: 0, no; 1, ~ + 0, ne: | : ‚уе. : сага sporophyte: 0, n - Ornamented tracheid та " Med yes. . True lignin: , no; 1, yes д Megaphylls: 0, по; 1; уез. richomes: 0, no; 1 ‚ Vascular cambium: 0, no; 1, yes pm. n 1 . Seeds: ; 1, yes ау branching E no; l, yes. - Flowers: 0, no; a EEA ND are marked in able 5 alignmen usted based on ше inei plant sequen soni. This cP = муш resolution was dior with the acd MOL set in this part of the gree n plants. $i vx nd P ge file is available b e from the authors (5) Ac combined daa se set xt (CP- COMB) was pro- duced by combining GP-MOLEC and GP-MORPH. This data set рад ы outgroups Emiliana and Anemonia from GP-MOLEC and the hypothetical ancestor from GP-MORPH, but otherwise used the full combined data as described separately above for 59 О a th combining morphological data vim LP-MORPH with seque ence data from GP- MOL LEC. These ин nine taxa have data in both sets чеш erem OTUs are constructed pairing p related, but not identical, taxa. The piana of the nine TABLE 2. Data matrix GP-MORPH. See Table 1 for list of characters and states, and Table 5 for list of taxa. Glycine max Oryza sativa Atrichum angustatum Fissidens taxifolius 1 4 4 ig Notothylas breutellii Phaeoceros laevis Porella pinnata Asterella tenella Riccia austinii Coleochaete nitellarum Klebsormidium flaccidum Micromonas pusila Mantoniella squamata Nephroselmis pyriformis Pedinomonas minutissima Tet "wd б Gea EA S 5 Enteromorpha intestinalis Ulva fasciata Ulothrix zonata 5 1 1 1 2 3 4 5 6 7 8 9 0 1 0 0 0 0 0 0 0 0 0 0 0 2310110001 1000000102 70??0????0 ??01000011 0777707710 1007230000 0700077000 071111111? ?11?000110 0111100001 1111111111 2310110001 1000000102 7022077770 2701000011 0777707710 1007730000 0700077000 071111111? 2117000110 0111100001 1111111111 2310110001 1000000102 7027077770 7701000011 0777707710 1007030000 0700000000 0111111111 1112000110 0111100001 1111111100 2310110001 1000000102 70??0????0 7701000011 07777027710 10072730000 0200022000 0111111111 1117000110 0111100001 1111000000 2310110001 1000000102 70??0????0 ??01000011 0?2???0??10 1002030000 0700000000 0111111111 1117000110 0111100001 1110000000 2310110001 1000000102 7077070710 ??01000011 0????20??10 1000230000 0200000000 0111110111 1117000110 0111111100 0000000000 2310110001 1000000102 7027000710 ?101000011 0777707710 1000030000 0200000000 0111111111 1110000110 0111111110 0000000000 2310110001 1000000102 70270007210 ?101000011 0????0??10 1000030000 0700000000 0111111111 1110000110 0111111110 0000000000 2310110001 1000000102 7127000210 ?101000011 0777707710 1000030000 0200000000 0111110111 1110000111 1000010000 0000000000 2310110001 1000000102 7177000710 ?101000011 0????0??10 1000030000 0200000000 0111110111 1110000111 1000010000 0000000000 2310110001 1000000102 7077000710 ?101000011 0??22?0??10 1000030000 0700000000 0111111111 1111111000 0000000000 0000000000 2310110001 1000000102 7022000710 ?101000011 0?2??20??10 1000030000 0700000000 0111111111 1111111000 0000000000 0000000000 2310110001 1000000102 7022000710 ?101000011 0????0??10 1000030000 0700000000 0111111111 1111111000 0000000000 0000000000 2310110001 1000000102 7072007710 ?101000011 0?2???0??10 1000030000 0200000000 0111111111 1111111000 0000000000 0000000000 0010110001 1100100102 1100000010 0101000011 0???707711 1000020000 0700100000 0111100110 0000000700 0000000000 0000000000 0000120001 0100100000 1100000010 0101000001 0??7707711 1000070070 0000000000 0710000000 0000000200 0000000000 0000000000 1200000000 000010120? 010212000? ??01100000 ?????0???1 0000030000 0700011000 0200000000 0000000000 0000000000 0000000000 1200000000 0000101?0? 0102100000 0101100000 7777707171 0000130000 0700111000 0000000000 0000000000 0000000000 0000000000 1200000000 0000101?0? 0100101001 ?101100000 ?????0???1 0000021000 0200110100 0000000000 0000000000 0000000000 0000000000 1200000000 000010120? 0102120201 ??01100000 ?????0???1 0000120000 0200000100 0000000000 0000000000 0000000000 0000000000 1200000000 000010170? 0101121101 0000111100 0??2??0???1 0000101000 0000010000 0000000000 0000000000 0000000000 0000000000 1210110010 0000100011 6101011001 0101100000 0777707001 1010771000 0011010101 0020000000 0000000000 0000000000 0000000000 1210110010 0000100011 6101011001 0101100000 0771707001 1010771000 0011010101 0070000000 0000000000 0000000000 0000000000 0310110000 0100100010 6101011001 0101100000 0707707001 1010101000 0701110101 0000000000 0000000000 0000000000 0000000000 uapsed jeoiuejog unossilA eui Jo sjeuuy 9st TABLE 2. Continued. Cymopolia barbata Batophora oerstedtii Codium decorticatum Cladophoropsis membranosa Blastophysa rhizopus Trentepohlia sp. Cephaleuros parasiticus Characium vacuolatum Dunaliella parva Chlamydomonas reinhardtii Volvox carteri Chlorococcopsis min Uronema belkae Chlamydomonas moewusii Stephanosphaera pluvialis Carteria radiosa Gonium pectorale Chlorella kessleri Chlorella vulgaris Prototheca wickerhamii Ch) 11 у Ba; id Chlorella minutissima Neochloris aquaticus x 2 3 4 5 0 0 0 0 0 1111111100 0300000111 7107071071 ??01100000 0777777701 111111000? 0200100110 7177011071 0101100000 0777777701 1110111100 0300000111 7104011001 0101100000 0777717101 1210111000 0200100110 7101011001 7701100000 0777707001 1210111001 0200100110 7101011001 0101100000 0777717101 2210110001 0200100110 1071011001 0101100070 27777707701 2210110001 0200100110 1071011001 0101100070 ?????0??01 0010000000 0001100010 7110071001 1001110100 7777207701 1000000000 0000101000 0110171701 1007777700 ??2??20???1 0000000000 0000101010 0110001001 1001110100 1771707011 0000200000 0000011112 7170777101 1001110700 ?????0???1 0000000000 0000100010 7110071001 1001110100 7777707701 0010110001 0210100010 6101011001 1001110101 0717707711 0010110001 0110100010 6101011001 1001110101 0717707711 0000000000 0000101010 0170001701 1007777700 77272707711 0000200000 0000011010 2110101101 1001110100 ?????70??11 0000000000 0000101010 0111001001 1007777700 2727707711 0000200000 0000011010 0110721101 1001110100 ?????0??11 0000000000 0000010?1? 717707777? ??????0100 ?0?1000??1 0000000000 000001071? 017707777? 2727770100 ?1?0000??1 0000000000 000001071? 717707777? 27727770100 272777702771 0000000000 000001071? 717707777? 7777770100 71720000771 0000000000 000001071? 271727027727? 2777770100 7070000771 0000001000 0000100010 7100071001 2011110100 27777707701 0000231010 0000131010 0000231000 1000771001 1000731071 1000071100 1000071100 0001771000 0000731000 0001101000 0101771000 0001771000 1010101000 1010101000 0011101000 0111111000 0011101000 0101101000 000?7?37000 0007737000 0007737000 0007737000 0007737000 0000271000 7 0 0?20002??1? 02707000111 0?000???0? 0?0?0???0? 0700000701 170777000? 170770000? 0700010701 0700077701 0100010101 0700077701 0200010001 0100010101 0700010101 0700077701 0700077101 0700010701 0200010101 222202220? 2127202270? 22220272702 222702770? 222202220? 0200010101 8 0 0000000000 0000000000 0000000000 0000000000 2000000000 1170010000 1170010000 0070000000 0000000000 0000000000 0020000000 0070000000 0070000000 0070000000 0020000000 00?0000000 0070000000 0020000000 07270000000 0200000000 0270000000 07270000000 0270000000 00720000000 9 0 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 1 о 0 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 1 1 0 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 #66! € JOQUINN ‘18 әшпюл "J£ 19 әче! .seyAydoiug,, pue , ,eebjy uəə19,, 1S TABLE 2. Continued. Neochloris vigenis Pediastrum duplex A i14 bo Characium hindakii Chlorella fusca Ankistrodesmus falcatus р. Ant hy, 4 + Characium perforatum Friedmannia israelensis Hypothetical ancestor T 2 0 0 0000001000 0000100010 0000210000 0000100010 0000210000 0000010010 0010001000 0001100010 0000000000 0000010?1? 0000000000 0001010?1? 0000000000 000020070? 0000110000 0100200?0? 0010000000 0001100010 00000 0000100010 9,,000000000 0000200002 3 4 5 0 0 0 2100027001 2011110100 ?????0??01 ?1?0011001 2011110100 1????0??11 ?1?40??20? ???1110100 ???1101??1 ?1000?1001 2021110100 ????20??01 01?20????? ??????0100 ?1?1101??1 21220272722 2222920100 ?????0???1 ?100021001 0200111100 0????0???1 ?1000?1001 0200111100 0????0???1 2110071001 010?11?000 ?????0??01 ?1100?1001 0107117000 ?????0??01 6 т 8 0 0 0 0000??1000 0700010701 0070000000 00001271000 0701010101 0070000000 0000277000 01000???0? 0000000000 0000221000 0700010701 0020000000 0007737000 7777077707 0770000000 0007737000 ????0???0? 0770000000 0000231000 0700010001 0070000000 0000231000 0200010001 0070000000 0000271000 0700010701 0070000000 ??1000 0720010001 0070000000 1 9 0 0 0 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 1 1 0 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 ?100011001 0100111100 02?2?20???1 0000731000 0700010001 0000000000 0000000000 0000000000 0000000000 1000000000 0000101010 2000000000 27100000000 0277707001 0000030000 0000000000 0100000000 0000000000 0000000000 0000000000 uapsed |eo9iuejog unossiIN 8S әш JO sjpeuuy Volume 81, Number 3 Mishler e 459 1994 “Green = and “Bryophytes” жык E варгана as outgroup) in this MUL- t is shown in T. ANALYSES À number of pes analyses were carried dile iin ا‎ ions combin ee зе d p тнв netic trees ех acCLADE, version 3.01 (Ma ddis ison e О with 20 megabytes of RAM. With the larger data sets, heuristic search algo rithms were rupe! f thus finding of all the y (MP) trees cannot be guar- апеей. 3 The analyses are presented in eight groups be- low, using standard numb emi-strict consensus (Bre- f the MP trees plus those that are опе step Mise bec decay за 41”), but absent in the semi-strict nsus of the MP trees plus bras E one or two steps longer (called fried ass Analysis 1. Land plant hological data al (LP-MORPH i р morphological data alone т n characters; 100 repetitions of RAN- хоп addition, TBR branch swapping with MULPARS and STEEPEST DESCENT option; de- сау analysis to 3 steps lb. Sper data alone (i.e., characters 1-65); 00 repetitions of an NDOM taxon addition, TBR MULPARS and STEEPEST PAU IN analyses also done (using the same ы Р options), to find the shortest юрю е соп- *nt with [mosses + tracheophytes] and [horn- rts + mosses + tracheo phytes] as monophyletic рз. dat General morphological data alone (i.e., Facters 66-113); 100 repetitions of RA M taxon eic TBR branch swapping with P STEEPEST DESCENT option; decay seriis to 3 steps. устуне зы деп жеге also done (using the tions), to find the shortest ч орав consistent with monophyly of the bry ld. Sperm data alone from the nine OTUs ке аѕ bs of LP-COMB; branch- not bound le. General morphological data alone from the nine OTUs selected as part of LP-COMB; branch- and-bound search lf. All characters from the nine OTUs se lected as part of LP-COMB; iud and-bound rch. Analysis 2. Green plant morphological data alone (GP-MORPH). CLOSEST taxon addition, TBR branch swapping with MULPARS and STEEPEST DESCENT option. ysis 3. Green plant molecular data alone (СР- MO 18S and 26S data combined: All OTUs except two и (59 OTUs); созт taxon sion, i h MULPARS, fi Mas with MULPA by NNI swapping. fes on the shortest trees found о alone plus two charophyte outgroups 160 TUs); 10 repetitions of RANDOM on addition, TBR branch swapping with MUL- PARS. 3c. nine land plant adire FEM as part of LP- COMB: spese and-bound algae minus charophytes, land nemonia (43 OTUs ping with MULPA on the shortest trees found by NNI swapping. Analysis 4. Green per molecular data (GP-MO- LEC). 18S data alon 4a. All OTUs including the two outgroups, swapping with MULPARS, followed by TBR branch ГА with MU ULPA RS on the shortest t trees £ 10 Ab. Ж к we plus Coleochaete as out- 460 Annals of th Missouri eae Garden TABLE 3. List of characters and character states used for data matrix LP-MORPH. See text for source of these characters d Table 4 for the data matrix. All characters are considered unordered, except as noted. b. Apical cell in aig apie: 2 absent; 1, prese 2; "аа не п young antheridia: 0, four. celled; 1, two-celled. 3. End eridia: 0, absent; 1, present. 4. Antheridial ek: "0, absent; 1, present. 5. Due: m cells: р. absent; 1, Tee sent. 6. Sperm in n politi t be: 0, арен il present. Le: Nu sabes of sper 1 ( d character): 0, 1000+; 1, 100-1000; 2, 16-24; 3, 2. 8. Nascent spermii: : 0, paired; ; 1, not paired. 9. Diagonal spindle in final m ne ivision: 0, mo 1, present. 10. Replication of the centrioles: 0, present; 1, a ll. Time of origin of esc лае 0, bi Buon 1, sperm mother cells; 2, sperm mother cell progenitor; 3, earlier. 2. Basal bodies (BB) and a 0, two; 1, more than two. icentrioles: 0, presen absent. а: T been 0, ES angles; 1, mo i side; 2, staggered anterior-posterior; 3, staggered continuous, . Proximal puse nsio 0, absent; 1, long; ort Proxi B: 0, ventral- dorsal; E ven ntral. tell imal e MER ate transition: 0, present; l, absent. . Connecting fibers between BBs: 0, present; 1, absent; 2, fine filaments with centrin. сша сый structure: 0, monomorphic; 1, dim [р ic. 20. BB stagger: ciated th: 0, absent; 1, present. аеро. of lamellar 0, а. b 85 2, partial. mellar strip/an : 0, parallel; 1, perpendicular. nie apert ure: o, "absent 1, present. li P 0, left of (нз, 1; E of center. : 0, adjac ; 1, beneath BBs. ег: stratified betwe алаа o. de nt; 1, present. spline/lamellar strip i d ua А , 45°. n:0, 0051, 4 osterior notch to lamellar strip: 0, absent; 1, present. amellar strip position: 0, under all BBs; 1, ba anterior P "m 2, under some BBs. tray Sam го! tubule: 0, absent; 1, present; 2, develops | Accessory b : 0, absent; 1, present. . Maturational ati nterior кө drion: 0, absent; 1, posterior. Spline shank: уле ыта 1; de than 4 tubules 1. O miophilic crest: 0, pes 1, pr 5. Anterior osmiophilic ridge: 0, absent; 1, present. ; 5. Changes in BBs at Seu y: 0, absent; n 1, dense material at tip; 2, BB cartwheel with plug; 3, BB triplets а їх. 37. Matrix around BBs: 0, homogenous; 1, mottled. 38. Posterior of the stellate pattern: 0, extracellular or partly; 1, entirely intracellular. 40. Late blepharoplast t with transient core: : 0, yes; 1. Di agellar rgence: 0, to чані ЕЯ L p rear; 2, toward anterior. 42. Nuclear shape at m viii 0, ovoid; 1, gate. 3. N 0, حر Ed‏ o‏ 32 © 5 © ,2 th‏ SER‏ P 4 uclear posterior shape: 0, not expan = expanded. 44. Median constriction: 0, absent; 1, Pe 45. Spline attached to nucleus: 0, yes Vin at и 2, never attached. 46. 5 pline ке assi хаса ме) а Mn ii ing: 0, absent; resent. 53, 4 irection of п lear compaction: 0, aye shell: 1, l, anterior sa posterior; 2, at equal rates along nucleus; general increase in densi 43 general 48. je ensed chromatin pore 0, дра d. igh toms to каен 2, spiral-central strand; ^ mpaction; 4, spikes; 5, ena pas edid 15 s from anterior tip 49. DEE during shap absen present. 50. Number of gyres of oan мден d 8, пої dune 1, 0.5-3; 2, greater than 3. 51. Dense body in anterior mitochon drion à . Mitochondri tissue: 0, absent; 1, үи 53. Mitochondrion associated with plastids in young spermatids: 0, absent; 1, present 54. Specialized anterior mitochondrion: 0, present; 1, abse 55. Specialized posterior mitochondrion: 0, present; 1, abse - mitochondrion: 56. Sgro ies mitocho: eie n in anterior of cell: 0, absent; ү row of mitochondria behind anterior mitoc unspecialize 57. Onn of a anterior mitochondrion: 0 0, fusion; 1, elongation. 58. Os ип! derneath anterior mitochondrion: 0, absent; l present... у, йд Volume 81, Number 3 Mishler е 461 1994 “Green pale and ''Bryophytes" TaBLE 3. Continued. ). Change from cristae sacs to baffles: 0, absent; 1, present. ч ро M absent. lastid nt—at ps 1, present —asymmetrical; 2, absent. : areh g grains in Кай, точ 0. ‘more ын опе; 1, astid contact Hd nucleu hu absent; 1, prese i rillerscheide: 0, a 61, pre be EL = 3 ens M "reri 2, complete (or tiny remnant). re 4 @ E a a: ent: . Lunularic acid: 0, abse My T present . Elaters: 0, absent; p e Oil bodies: 0, absen ү nt. D-Methionine: 0, i ag cognize; 1, recognize. Stomates: 0, absent; prese Vertical division of zygote: 0, absent; 1, present. Xylem: 0, absent; 1, presen I oem: 0, absent; 1, p Perine layer on spores: 0, i el absent; Aerial sporophyte axis of Sa "radenphye type: 0, absent; 1, present. - Columella: 0, absent; 1, рге ; ica ios: 0, derit E pre Leaves on game etophyte moss че " D absent 1, present. nd abse p! t; pl 0, absent; 1, present. ы : Тао 0, absent ds present. . n: 0, absent; 1, prese . tw exserted seta of the servo a 0, absent; 1, present. ce pre . y cells: 0, absent; 1, . Spore cells lobed: 0, m ds T presen - Capsule 2-4 valved on a regular basis: ua ж d E1; pre Capsule cells with transverse thickenings: belt vis pE . Thick capsule wall: O, absent; 1, RE а aE dp absent {, aus . abse . filial ү Een er: vis ‘absent ; 1, pre sie sent; 1, present. ] Se pre . mir 0, s nt; E n BE + Microphyll: 0, absent; 1 nt. arch maturation of xylem: 0, hye 1, present ' ‹ hloropla жеп почаи 0, absent; 1, present. . Sporangia borne on leaves: 0, aet 1, present. - lrichomes: 0, tek ; 1, pre lavonoi , absent; 1, radian od etention of zygote: 0, absent; 1, present Sheathed hairs: 0, absent; 1, р t. olyphenolics induced by sex: 0, absent; 1, present. ` tarenchyma: 0, absent; 1, present. Beaked mucilage papillae: 0, absent; 1, present. soup (15 OTUs); 10 repetitions of RANDOM EST taxon addition, TBR branch swapping with ion addition, TBR branch EM н MUL- MULPARS and STEEPEST DESCENT option. оне TT 4d. Land plants € plus Coleochaete and lebsormidium; Pedinomonas as outgroup (17 lants haete and Kleb pigeon OTUs); CLOSEST taxon i кше. TBR branch Land dae! as outgroups (16 OTUs); CLOS- TABLE 4. Data matrix LP-MORPH. See Table 3 for list nd states. Characters 1-65 are from sperm ultrastructure; 66-113 are from general morphology. The asterisk ( *) for character 65, Lyco, podium, оні а ein between ates i and 2 Lo unpublished); the asterisk for Bom 86, Sphaerocarpos, represents a polymorphism between states 0 and 1 within the order sperm characters: general morphology: 1 1 1 2 3 4 5 6 7 8 9 0 1 0 0 0 0 0 0 0 CHARA ?????0??00 307007020? ??0??0???0 070000000? 1100000001 0??1?1?011 20102 00000 0000000000 0000000000 000000 100 000 NITELLA 2777707700 307007020? ??0?000??0 070000000? 1100000001 00012???0?1 20102 00000 0000000000 0000000000 ?000000000 0000200100 000 COLEOC PUL 2222207270 007007070? 000?000??0 000000000? 00000 0000000000 0000000000 2000000000 0000700011 100 COLEOC ORB ?0??10 0000?1 0000 0000000000 0000000000 7000000100 0000700011 110 PHAEOCEROS 0011000001 101111100? 200?101000 010003001? 1101001101 1110000110 11102 11000 1110000100 0000000000 0000000100 0000200010 110 NOTOTHYLAS 0011000001 101111100? 2007101000 010003001? 1101001101 1110000110 11102 11000 1110000100 0000000000 0000000100 0000200010 110 MARCHANTIA 0001000001 1012100111 2011101100 000003001? 1100001011 0010000010 10112 11111 0000000000 0000010000 0000000110 0000000110 110 SPHAEROCARPOS 0001000001 1012100111 2011101100 001003001? 1100001011 0010000010 10??2 11111 0000000000 00000*0000 0000000110 0000200110 110 PELLIA 0101000001 1012100111 2011101000 000003001? 1100101011 0010000010 10112 11111 0000000000 0000101111 0000000000 0000700110 110 BLASIA 0201000001 1012100111 1100001011 0010000010 10112 11111 0000000000 0000101111 0000000110 0000700110 110 JUNGERMANNIA 0101000001 1012100111 1100101011 0010000010 10112 11111 0000000002 0000101111 0000000010 0000700110 110 HAPLOMITRIUM 0101000001 1012170111 2010101000 700703001? 11000015?1 0010000010 ?1??2 11111 0000000000 0000101100 0000000110 0000700110 110 TREUBIA 0101000001 1012100111 ?010101000 2700703701? 11?0001??? 0??0000??0 ?0??2 11111 0000000000 0000101111 0000000??0 0000700110 110 SPHAGNUM 1101000111 1012101111 1011101012 010003001? 1100012201 0110000010 01002 11000 1100011111 0000000000 0100010000 0000200010 110 ANDREAEA 1101100111 1012107111 100012201 0110000010 00002 11000 1001011111 0000000000 1111000010 0000700010 111 POLYTRICHUM 1101100111 1012100111 2011101011 011003001? 1100012201 0110000010 00002 11000 1101111111 0000000000 1111111010 0000700010 110 HYPNUM 1101100111 1012100111 2011101011 010003001? 1100012211 0110000010 00002 11000 1101111111 0000000000 1111111110 0000000110 110 TAKAKIA 1101100111 1012100111 ?01110101? 710703701? 1100?????1 0?10000?10 00??2 11000 1001771171 0000000000 1107000010 0000200110 111 LYCOPODIELLA 0700001711 101407020? 01071 11000 1101111000 1111000000 0000000110 0111000110 110 LYCOPODIUM 0200001011 101221020? 0107101000 010000101? 1000073310 0000107010 0010* 11000 1101111000 1111000000 0000000110 0111000110 110 SELAGINELLA 0??0?01??1 107221070? 2 010007010 70002 11000 1101111000 1111000000 0000000270 0111000110 110 EQUISETUM 0200001111 1001100011 1110003301 0000111001 2?001 11000 1101111000 1111000000 0000000770 0700110110 110 PTERIDIUM 20000211 27001 11000 111111100 1 11110 110 MARSILEA 0770702111 7001 11000 1111111000 11 000211110 1 OSMUNDA 0? 2111 1001110011 1110003301 0000111001 2?001 11000 1111111000 11 00000110 1000111110 110 GINKGO ?7?7713711 2?000 11000 1101171000 71 1000111110 110 TAMIA 77713711 2700 1000111110 110 иәрлеку [Eoiuejog unosstlA c9vr Əy} JO sjpeuuy Volume 81, Number 3 1994 Mishler e 463 “Gr reen Т. and “Bryophytes”’ swapping with MULPARS and STEEPEST DE- option. The nine land plant аге selected as part of LP. COMB; branch-and-bound search. ysis 5. Green ee molecular data (GP-MO- TH 26S data alon a. All OTUs that кее 26S data; Pedin monas as outgroup (37 OTUs); 10 repetitions of RANDOM taxon addition o branch swapping vith MULPARS and STEEPEST DESCENT op- 9b. Land plants that have 26S data; no out- groups (11 OTUs); CLOSEST taxon addition, TBR branch swapping with MULPARS. seven land plant OTUs selected as part of LP-COMB that Меп 26S data; branch- and-bound search Analysis 6. Land plant molecular data alone (LP- MOLEC), Each with 10 as of RANDOM taxon addition, TBR branch s wapping with MUL- PARS and STEEPEST DESCENT an а а; Coleochaete and Klebsormidium as Колы Wo TUs). vet n data; Coleochaete only as outgroup us ү к data; Klebsormidium only as outgroup 6d. All data; no outgroups (14 OTUs). 6e. 18S data only; no outgroups (14 OTUs). 6f. 26S data only; no outgroups (11 OTUs). "rose - Land plant combined ав (LP-COMB). cha [ш and-bound se. Ta. All data. b. A 1 З : oe т neral morphological data plus all mo © All morphological data plus 18S molecular a A morphological data plus 26S molecular ral pla qus ne ы 18S mo- ashes analysis to 2 s hog General morphological Ma s 26S mo- teps. 1 data; decay analysis to Analysis 8. Green plant combined data (GP- COMB). CLOSEST taxon Bir n, NNI branch Vinc: with MULPARS, followed by TBR branch with MULPARS on the shortest trees ix s NNI swapping; das analysis to 3 steps. RESULTS Analysis la. Only one tree island was found, of 9 MP trees at 208 steps (CI — 1, along with the deca branch (obviously, tiere. and n aeter figures, tpe ipie че groups (seed plants, tracheo- уд ides اتوت‎ among these major lineages resolved. Note also that the lyc Кеин меге пої supported as monophyletic Analysis 1b. Only one tree island ты found, of 56 MP trees at 130 js ne step. However, not all trees at 2 and 3 steps less aved with available RAM, 99 669 99 and m qua were well supported by this data set, ve ta irn bryophyte clade rted. Selaginella was plac » adi ide ырны. CONSTRAINT aoe showed that MP topologies with a forced Mishler dies chill (1984, 1985) e i.e., [liverworts [horn- worts [mosses + a һуїез []], were 2155 steps long with this dat was moder ately tree island was found, of 8 MP trees at 67 steps (CI — 0.731; RI — 919). The strict consensus tree is sho Figure 3, along with the dec index for each informative ch. The decay analysis was complete steps. However, not all не аї Э чер - par- ecay classes “3” and “4+” Figu are The Mishler and Churchill topology: was IN yses show ei t topologies with forced bryophyte mono- phyly were 70 steps long with this data set. No that the tracheophytes were strongly үр» as monophyletic, as were the lyc ophytes. Thus it 464 Annals Is of the Missouri Botanical Garden Taxa included in the data sets GP-MOLEC, GP-MORPH , and GP-COMB, with source for rDNA LE 5. sequence data (GENBANK accession number given, if known). The taxa selected for the data set LP-MOLEC are ma k. rked with an asteris Emiliana huxleyi (Lohm.) Hay & Mohler Anemoni ata L. male * Atrichum angus RN n Brid) Bruch & Schimp. *Notothylas breutelii К Жен аме laevis (L. r fh ly hin e innata L. *Con лаві сопісит а ж Lindb. иа tenella Е * Riccia austinii *Klebsormidium o cian ( 6g Br.) Silva, Mattox & Blackwell a m J ) Webe & Mohr) Kutz Cymopolia barbata (L. ) „ры Batho phe ora tee Codium (Wood Pt sis membranacea a (C pa ) Bas ANE rhizopus Rein Trentepo sp. Cepha aniio parasiticus Karsten Characium vacuolatum Lee & Bold Dunaliella parva Lerche hlamydomonas reinhardtii Dangeard Volvox carteri f. nagariensis Chlorococcopsis minut "m iUm minuta (Arce & Bold) Komarek) i h.) A onium pecto torale Muller Beij. Prototheca wickerhamii Soneda & Tubaki Chlorella RR Krug. Chlorella minutissima Fott & Novakova Neochloris aquati tarr Neochloris vigenis Archibald Pediastrum duplex е Bo! Chlorella fusca var. vacuolata Shihira & Krauss Bhattacharya et al., Hendriks et al., ою зн Chapman, unpub Lewis et at ^ 169 (M63001) Lewis et al., 199. = M62998) Gunderson et E Rausch et al., Lewis et al., i992 (15296) Bu im & Chi an, 1992 So d 2 9 M62961) Wilkos et m. eo (M74496) Lewis et al., pa M62997) Huss & Sogin. Lewis et al., Yos ^ M63000) Huss & Sogin Volume 81, Number 3 1994 Mishler e “Green nee: and “Bryophytes” TABLE 5. Continued. Ankistrodesmus урыны (= A. falcatus var. P ges: (Chodat) Lemm.) Huss & Sogin, 1990 Pseudotrebouxia gigantea Hildreth & A cop terrestre Fritsch & John Characium perforatum Log & n arietochloris p & Bold) Watanabe & Floyd Friedmannia Гаури САТ эй & Bold Kantz et Lewis et tt 1992 (M62999) Lewis et al., 2 (M63002) Lewis et al., pe (M62995) clear that a major conflict exists between the sperm data (see E. A analysis lb) and the general I ld Ana ld. Three MP ind were found, at 65 а ‘Cl = 0. 862; RI = 0.870), m strict consensus of which is shown in is ure 10. Note that as in analysis Ib, bryophyte ENGL was supported. Analysis le. Two MP a were found, at 35 steps (cl = 0.857; RI = 0.886), the strict 10. Note sister group to the tracheophytes as in analysis le, the mad position of liverworts and hornworts was not resolved. Апа s lf. Two MP trees were found, = y (d - 0.819; RI = 0. 552. the strict nsensus of which is shown i e 10. Note that bryophyte bl" was OY) unlike the result of апа alys Am lysis 2. 26,300 MP trees were found at l steps (CI — 0.550; RI — 0.895), but the ki ul RA of the Computer was . Thus, the effec- ti exceeded RE of the heuristic search was heda nd ore trees id 2 n оше e exist than could be зау 1 Were say. E à is kde in Figure 4. Note the poo resolution among green deals aud ш ш groups. Analysis 3a. 32 MP trees were baa: at 2245 steps (CI = 0.458; RI = 0.5 cei rict consensus of the trees that igure HH and ies were unresolved d a monophy etic group basal to the chlorophytes plus pleuras- other 1 trophytes. Pleu. the rastrum was grid separated from pleurastrophytes d. 3b. rit pue island of 5 MP trees t 304 — 0.612; RI = 0.600). a In the c phylet DD (not shown), neither the land plants, tracheophytes, nor liverworts were mono- ic. eerie 3c. o MP trees at 127 steps (CI 0.630; RI = 0. Hes were found, the strict con- sensus of Nen is mon Mere. 0. The mosses and I h c group, and ie liverworts heit. M Eri. Analysis 3d. Only one MP e of 1597 steps, was found (CI = 0.501; RI = 0.518; shown in Fig. 6). None of the three тын of green algae nsu Matto: x & Stewart (1984: vet dx been pleurasisophytes and ulvophytes) included in уне ared to be strictly monophyletic. эчә ктар were paraphyletic near the base, and M LE 6. Composition of the OTUs used in data set LP-COMB. Morphological data were taken from selected in LP-MORPH (Table 4) and molecular data were taken from selected taxa in GP- MOLEC (Table 5). OTU LP-MORPH GP-MOLEC COLEOCHAETE C. orbicularis C. nitellarum PHAEOCEROS P. laevi P. laevis NOTOTHYLAS Notot N. breutelei MARCHANTIALES Marchantia Asterella JUNGERMANNIALES ungermanni Porella POLYTRICHALES Polytrichum Atrichu RYAL пи Plagiomnium EQUISETUM Equisetum Equiset ZAMIA Zamia Annals of the Missouri Botanical Garden Analysis 1a. Full morphological data set Decay Index: = 4+ steps E шинни = 3 steps шиини = 2 steps ae | =l —— = 1 step E — E Tur PTERIDIUM OSMUNDA MARSILEA GINKGO ZAMIA EQUISETUM LYCOPODIELLA LYCOPODIUM SELAGINELLA PELLIA JUNGERMANNIA BLASIA TREUBIA HAPLOMITRIUM MARCHANTIA SPHAEROCARPOS ANDREAEA TAKAKIA POLYTRICHUM HYPNUM SPHAGNUM PHAEOCEROS NOTOTHYLAS COLEOC PUL COLEOC ORB CHARA NITELLA аа apte p.MORPH). IGURE l. Semi.stri f ni i ; n n ba The decay i i : 3 t P cay index is shown for each informative branch; thicker branches are better supported. Volume 81, Number 3 1994 Mishler e 467 “Green е апа “Вгуорһуїе$” Analysis 1b. Sperm data only Decay Index: шиша = 3 steps =m = 2 Steps —— =1 step ED =. E- 4+ steps ; POLYTRICHUM HYPNUM TAKAKIA ANDREAEA SPHAGNUM MARCHANTIA SPHAEROCARPOS BLASIA PHAEOCEROS JAN NOTOTHYLAS‏ د PELLIA b JUNGERMANNIA HAPLOMITRIUM TREUBIA SELAGINELLA PTERIDIUM OSMUNDA EQUISETUM GINKGO ZAMIA MARSILEA LYCOPODIELLA M aae L- LYCOPODIUM PUL орот COLEOC PU [с COLEOC ORB oper CHARA ЕА cters 1-65, F n f, (c branch; thicker Feet are better AM. Lp CURE 2. бепан f£ 56 "MORPH). The decay index is shown for Annals of the Missouri Botanical Garden Analysis 1c. General morphological data only Decay Index: = 4+ steps ишш = 3 steps mes = 2 Steps —— = 1 step better suppor n for informative branch; thicker PTERIDIUM MARSILEA OSMUNDA GINKGO ZAMIA EQUISETUM LYCOPODIELLA LYCOPODIUM SELAGINELLA ANDREAEA TAKAKIA POLYTRICHUM HYPNUM SPHAGNUM PHAEOCEROS NOTOTHYLAS PELLIA JUNGERMANNIA TREUBIA BLASIA HAPLOMITRIUM MARCHANTIA SPHAEROCARPOS COLEOC ORB COLEOC PUL CHARA DII МЕША Fic whe Semi-strict consensu & igh ua ical characters only branches 216 Volume 81, Number 3 Mishler е 469 1994 "Green ndm and “Bryophytes” E Cymopolia barbata Analysis 2. Morphological data alone adm iin horopsis jc ER itepohlia sp ( phous parasiticus omorpha intestinalis Г?! [tec Ulothrix zonata — — re EER r— L— Draparnaldia plumosa Uronema belkae Pseudotrebouxia gigantea Friedmannia israelensis Characium perforatum Parietochloris pseudoalveolaris p Ahi lla | lari Rioreia KESsier nhi Chlorella vulgaris m lor ella fusc nkistrodesmus : falcatus(sti) > ~hlamydomonas moewusii ( ;arteria radios sa IE \ folvox | сапеп Кый) onium pectorale unaliella parv etraselmis carteriiformis ( ilycine max t onoceph Г / stra tenella Ricci SSS ae. отуна breutellii س‎ | ha eoceros laevis di " 9n L ала 4 FIGURE 4. MORP H). Mor yp Strict consensus of 26,300 most parsimonious е for the large-scale morphological data set (GP- е trees at this length existed that could not be sa 470 Annals of the | Missouri Botanical Garden Analysis 3a. Molecular data alone (18S plus 26S) тотор vac rococc m ADDO = © © ЕЕ] ш © 3 o n hlamydom r I Junaliella e ! leochloris a aqu / nkistrodesmus гарат arn plum‏ چ İL Uronema belk‏ Cymopolia barba‏ Pariet pse Pseudotreb gig Friedmannia is Codium decort Cladophoropsis — Trentepohlia oo C pus Mantoniel squa‏ سا Nephroselm руг‏ Pedinomonas mi‏ FIGURE 5. Strict consensus of 32 most parsi i for the full molecular data set (GP-MOLEC). Volume 81, Number 3 1994 Mishler e “Green pin and ''Bryophytes" 471 Analysis 3d. Molecular data alone (18S plus 26S) Micromonas pus Mantoniel squa Nephroselm pyr Pedinomonas mi Characium vac Chlorococc min os afa Ulothrix zo Codium decort Cladophoro Trentepohlia Cephaleuro par Blastophysa rh ЕЕ Cymopolia barba Batophora oers the green algal taxa with the full molecular data set ке МОГЕС). fn е for tory of ee я is praise өгү dd the number of changes under ACCTRAN optimization the = е pleurastrophytes were ренин above to chlorophyt tes + Pleurastr Analysis 4а. found (CI 4 MP trees at 1888 steps were m 0. ue RI — 0.589), the strict con- the tracheophytes and the mosses as the sis group to that clade. сяр deep branches іп the tree, Annals of the Missouri Botanical Garden Analysis 4a. 18S data alone Decay Index: EN = 3+ steps = = 2 Steps — z1 step Characium vac и: ар Chlamyd reinha olvox carteri onium pecto eochloris aqu y f 18S — only (GP- MOLEC). The а "ders is ye hen rig di sees donet. thicker e E are сне: виррог Volume 81, Number 3 1994 Mishler e 473 “Green td and ‘‘Bryophytes” of the two major clades of green абы меге Маска; well peepee ed. In- terestingly, the micromonadophytes were support- ed as a monophyletic group in this Serer’ whereas е s were not. Im 4b. Only one MP tree, of 163 st ound (CI — 0.632; RI = 0.641; not ea ] ta basically pea same topology for the land in analysis 4c. Fou Analys r MP trees, of 176 steps, were found (CI = 0. 614; КІ = 0.630; not shown). worts, Porella, allose шу ne co Ht Н forming a tetrachotom Analysis 4d. Six MP ibus of 206 steps, were found (I= 0.607; RI = 0.595; not shown). The consensus = > same Е for the land plants as in analys Analysis 4e. One MP tree was found at 83 Steps (I= D 627; RI = 0. 492; shown in Fig. 10). е bryophytes PF 1 as a monophyletic group 4 a. One tree a was pes e к us pe sail in Figu Iw оону ге she, uda жы "E land Tom d algal classes were all depicted in different а arrangements t inui in previous analyses. п alysis ai Two MP T were found at 83 > ЧС = .699; RI = 0.699), sik strict con- Sensus of ce h (not shown) had an ual to- Pology, with neither the аеру п nor the mosses potentially mono phyleti MUS. f o MP trees at 42 steps were ound (CI = ave an unusual topology, with the ا‎ outgroups widely separated, and tracheophytes polyphyletic Three MP ig а 271 s = 0.6 642; RI = 64), t cde con- Mec of E (not shown) "gd "е topology fro re-rooting о e 9. The two Pelis: the tr. cheophytes were still impo bet and the mosses plus hornworts were monophylet un ^ is бс. Four MP ен at 250 steps were d (CI = 0.636; RI = 0.669), the consensus I3 vis (not shown) had all major groups unre- ved. s 6d. "Three MP trees at 225 steps bens азна but neither liverworts otentially monophylet s 6e. Seven MP trees at 146 steps were found (CI = 0.662; RI = 0.702), the unrooted Ода of which not shown) had the topology Analysis of Two MP pes at 78 steps were found "s = 0.692; RI = 0.700), the unrooted | RS QE A US | had th Consensus or \ very unusual topology of analysis 5b, with neither the mosses nor the tracheophytes potentially mono- phyletic A 7а. Three MP trees at 240 s were found (CI = consensus of which ti shown hat the agent groups of land plants were com- pletely unresolved. Four МЕ sr at 168 steps were 61), the strict con- 10. The horn- worts, mosses, and tracheophytes were placed to- gether in a trichotomy. Analysis 7c. о MP trees at 191 steps were found (CI = 0. ph "RI = 0. «йы the strict con- sensus of which is shown in Figure 10. The bryo- phytes were placed together in a dins eet alysis 7d. One MP tree at 150 steps was und (CI = 0.760; RI = 0.743), shown in Figure . Note that the Mishler & Churchill (1984, qe topology, i.e., [liverworts mg worts [moss- es + tracheophytes]]], was pre nalysis 7e. One MP tree at 124 steps was си [hornworts [liverworts [mosses ytes]]], but it was relatively weakly monu srt 7f. ee at 78 steps was found (CI = 0.744; RI = bri shown i re is in Fi 1985) topo nt and was rela- logy was again prese tively well supported. Annals of the Missouri Botanical Garden Analysis 5a. 26S data alone Stephanos pl Gonium pecto Chlamydom moew FOR т varena raa pr Enteromorpha LL Uva tasci Tetraselm cart е ——— Хх zo n | тарап рип Uronema belk DI ds Fieurasu оп Trentepohlia Cymopolia barba Batophora oers Cladophoropsis Pseudotreb gig Codium decort май роза vfi | p Micromonas pus LLL Mantoniel squa [us Conocephal con Asterella tene Ds Riccia Equisetum ar Еди Atrichum angus Phaeoceros lae Pha Klahsormid fla Klebsor Nephroselm pyr Pedinomonas mi FIGURE 8. Strict consensus of 462 most parsimonious trees for the 26S characters only (GP-MOLEC). Volume 81, Number 3 Mishler et al. 1994 "Green Algae" and “Bryophytes” otha E. Molecular data alone Glycine max (18S pus Ee realignment for ык рап Oryza sativa Zamia pumila Coleochaet nit * 475 Conocephal con Asterella tene Riccia T сакл Psilotum n Equisetum ar Fissidens taxi Plagiomnium cu Atrichum angus ET Notothylas bre [ Phaeoceros !ае Роге!!а рї T Klebsormid fla * 3 А h hyt: lgae. marked with b 5 F t TA FIGURE 9. Sing) t parsimoni for the la es ) g P tree f terisks) with the full molecular data set (LP-MOLEC). ing "le is 8. Fifteen MP trees т 2511 steps The major branches of the tree were relatively well ul ы "Mos = 0.458; RI = the strict — suppo: oe 2 were identical to the Mishler E: of which is shown этш: ure 12, along Chur the decay index for each informative branch. sire" pA (i.e., charophytes -- land plants; e с - © e a — S © 2. © oà i. Б m т © E 5 c PHAEOCEROS NOTOTHYLAS MARCHANTIALES JUNGERMANNIALES POLYTRICHALES BRYALES EQUISETUM ZAMIA COLEOCHAETE Analyses 1d, 1f, 7c. PHAEOCEROS NOTOTHYLAS POLYTRICHALES BRYALES EQUISETUM MARCHANTIALES COLEOCHAETE Analysis 5c. POLYTRICHALES BRYALES EQUISETUM ZAMIA PHAEOCEROS NOTOTHYLAS MARCHANTIALES JUNGERMANNIALES COLEOCHAETE Analysis 1e. PHAEOCEROS NOTOTHYLAS MARCHANTIALES JUNGERMANNIALES POLYTRICHALES BRYALES EQUISETUM ZAMIA COLEOCHAETE Analysis 7a. PHAEOCEROS NOTOTHYLAS POLYTRICHALES BRYALES EQUISETUM sl ZAMIA ——- JUNGERMANNIALES MARCHANTIALES COLEOCHAETE Analysis 3c. PHAEOCEROS NOTOTHYLAS POLYTRICHALES BRYALES EQUISETUM ZAMIA MARCHANTIALES JUNGERMANNIALES COLEOCHAETE Analysis 7b. PHAEOCEROS NOTOTHYLAS MARCHANTIALES POLYTRICHALES BRYALES JUNGERMANNIALES EQUISETUM ZAMIA COLEOCHAETE Analysis 4e. POLYTRICHALES BRYALES EQUISETUM ZAMIA PHAEOCEROS NOTOTHYLAS [ MARCHANTIALES JUNGERMANNIALES COLEOCHAETE Analysis 7d. иәрео [eoiuejog unossiIA 94t au} JO sjeuuy Volume 81, Number 3 1994 Mishler e 477 “Green pene and “‘Bryophytes”’ mer et al., 1987). Unlike previously published to- pologies (Zechman et al., 1990), howev: he ~ @ ^ М ct addition of the problematical genera Trentopohlia and Cephaleuros), while neither T descri sensu Mat Stewart, 84) were supported as REUS Я groups. DISCUSSION In general, it appears that the molecular informative regarding relationships were most mong the green algae. In the combined analys of molecular data (ап alysis 3), resolution of e n algal clades was well supported, whereas resolution of land pl. very po here Q I] а). The realignmen: into account just the ges plants n б), Ф ight have been expected to yield a better as- 1991). marily ultrastructural, making sufficient sampling difficult. These diffi poor t redes obtain Morph al data coded narrowly for the land plants, ho analysis D. ме the latter case, the greater conser- v al dat y have allowed the recovery of a historical signal obscured olecular data. n suggested з, even complex mor- phological агаш Vine may be subject to strong selective dn sis and, es и give те € phy: веке п pe пз because of п nie den oe characters. T is possible that the сойбо! 1 in the mı derived from the E and the general set ganism constraints impos ame sessment of ef poston мз Mindell о of no help in “impr ovine ” the results. ame g te (с f. ба оу et al., 1990) ог оп Le other extinction in the two major t Ultimately, the only means of етет such c Аз it happens, the extant green sided Em d more flicts is by investigation of many different e. evenly sample t tree than xtant acters and cha Е systems. bm one character Streptophytes. Spurious long-branch attractio system (or 1) straints oblems are exp on theo retical dre to bias pl be greater in data t pes with a restricted number or another. The hope i is that if one po at enough of character states, such as (Mishler, character systems the 1994 » and may therefore be seen more frequently to Ш *cancel-out" and а (ша historical $ he green al ith the signal can be detected. Roda is hems one known i po tern across ogical characters broadly across the plants because of the 1 ous the green algal groups be- use the а useful at that level аге ргі- process that can se n pat all Lam widely different aene systems: log pu в dificult to ee үа дшш conclusions fr d шн мч! їп sol 7, beca of characters is ven that only eight land plants oni eh clu Si The combination of both genes together, as well as 18S alone, with the LP- MORPH data set tended to favor a monophyletic bryophyte lineage, ар the gombinstion with 4 t е S alone tended to fav ment of bryophytes in ш вате ран : [liverworts — phytes]]]. The las Spo as neither the а data alone (analysis tern as Figure h D ally intriguing, Ficu trom he "ud available. eed da da ict consensus t ta set 0 COMB) ч that contains land Т4 which had both molecular а d mero d Да Annals авсан Bob Garden Analysis 7e. General morphology EQUISETUM plus 18S molecular data ZAMIA POLYTRICHALES 3+ BRYALES ў Prestan MARCHANTIALES rpm JUNGERMANNIALES PHAEOCEROS — NOTOTHYLAS COLEOCHAETE Analysis 7f. General morphology EQUISETUM plus 26S molecular data т 6+ ZAMIA 2 POLYTRICHALES Em | — 0 BRYALES j TT PHAEOCEROS рени e eee MARCHANTIALES 3 DER JUNGERMANNIALES COLEOCHAETE FIGURE 11. e most parsimonious trees from two different combined analyses of the reduced data n COMB) | en xima land plants which had both molecular and morphological data available. The decay in shown for each informative branch; thicker branches are better supported. 1f) nor the 26S data alone (analysis 5c) gave such a pattern on- a pem that was better resolved than the © The o of the individual ап n of each data < ега] combined analysis of GP-MOLEC lone. " This property is often case with p* and GP- MORPH (analysis 8—Fi ig. 12) produced ее evidence” matics uh. 1989). Volume 81, Number 3 Mishler et al. 479 1994 "Green Algae” and ‘‘Bryophytes” Vol апе Analysis 8. Combined тоа реди and morphological data Chlamyd reinha Decay Index: Chlorophyceae ШЕШ = 4+ steps orella min mmm _ ; : 3 steps Erna PadiastiUm dup c9 op $5 33 ce | = = 2 steps oo 3 © S 3 = о Pleurastrophyceae mec step Ankistrodesmus . | [9] m Ulvophyceae streptophytes orella ар! Pedinomonas mi E © 2 © 8 © m Micromonadophyceae Th GURE 12. Strict consensus of 15 most parsimonious trees for the la: о combined data set (GP-C OMB). * decay index is shown for each informative branch; thicker iere hes are better supported. ssig unen: Э Breen algae to classes accordin ng to the concept of Mattox & ee rt (1984) is dom "streptophyte" indicate " a рһуте green algae plus embryophytes (sensu Bremer et al., 87). The unusual placement of three enigmatic axa is indicated with an asterisk. Annals of th Missouri Reid Garden Chloro. Pleur. FIGURE 13. diagr = oes sensu feng and Stewart (1984), and the four ra whi we ar i siteptaphiptes ^ ad completely теб, їп а to- Мі fishler & Churchill (194, 1985) эш а апа]уѕіѕ а here, despite the fact that п or separate analyses 4 and 5) alone gave such o е observation of a со topology that is different from topologies produced in separate analyses of co ent data sets is not unusual (see discussion of anal above ferent reasons in eac observable n the noise in the two ^e sets cancels out combined a ан (Barrett et а]., 1991). анау. coordinated studies are needed an explanation 0 subanalyses of anal- уа pee the exception. that the ulvophytes - Lie he E Several OTUs with dis pests Bi branches (suc s Pleurastrum and Ankistrodes- mus) “jump Nem in equally parsimonious po- sitions, ves кюне support for the monophyly of both d Chlorophyceae as pasta circumscribed. Major clades within Micromonad. major lineages of la nt a n black, those for ppled, while two en of -kosa certain paraphyly ing of cladistic relationships of the five classes of green : nd plants, sensu Bremer et al., (1987). which Ke support exists had considerable the chlorophytes, however, such as the taxa with ire odies and those with clock- The results of the various analyses illustrate, if nothing else, how difficult nh prios Wee tion is a ruo = EF and systematic studies to a larg that much of the remaining pos ч йл M moved once more А te data sets are produ a There is still a great de а of work to be done, we are ded pro e À summation of ie ursus hypothesis, ge phytes, and ае ytes), the other “ "es "charophyte" green ee: along with Volume 81, Number 3 1994 Mishler е “Green pent and ‘‘Bryophytes”’ phytes plus tracheophytes. A doubtfully monophy- letic assemblage of unicellular micromonadophytes phylogenetically *between" these two major lin- eages. It appears clear t he ulvophytes basal to the chlorophytes plus pleurastrophytes, but the exact circumscriptio t 1 needs revision. In о m alyses, however, including the ined Ad TECH (analysis 8—Fig. 12), the ie E е of ipm галу рага- phyletic o the tracheophytes, with the pam еа; [hornworts e + tra- ia JJJ. [=] tn © cheo) e robust parts of this summary clado- gram ve Él in need of support from future studies sampling TUs and more bass $ a on у БӘМ аѕ well as molecular ns a fra а wie n The habitat еи the т f ev И olutionary issues LITERATURE CITED ALBE si де A. & B. D. MisHLER. 1992. On the rationale Cla _ бане of m nucleotide sequence data reU М. E & B. D. MISHLER. 1993. Gar: codin DN ting for _ 0: а 2 perti уы, Ann. e ae Card. TNR HB. D. Misure vius W. CHASE. acter. stat f 10) 1992, Char- E hs logenetic iunc ith chl а= DNA. Pp. 369-403 in P. S. Solis HE. d 4.4. Doyle editors Molecular e va Chapman & Hall, New York. A ронса НОЕ & "E SOBER. gainst consensus. Syst. Zool. 40: 486-493. B 1991. BHATTACHARYA, D., L. MEDLIN, P. О. WaiNRIGHT, E. V. ARIZTIA, C. BiBEAU, S Kk. Sr TICKEL P M. L. IN. 1992. A + c are para- of amin acid seque data in angiosperm ийетен pean: bs lution 42: 795-8 =. og aie component consensus. Cla- distics 6: 369-372. ‚ C. J. HUMPHRIES, B. D. MISHLER & S. P CHILL. 1987. On cladistic sedan in eee plants. Taxon 36: 339-349. Brown, В. C. & B. E 1988 e nd organismal dat ‚ TUR E “A. ZIMM R & R. 1990. Phylogeny o of Chlamyd L. CHAPMAN. ophy- lear 18S rRNA sequence data. J. йс. 26: 689 CAROTHERS, Z. B. & A. E. RUSHING. 1988. Comparative ophyte ее Аа- . А. BucHHEIM. 1991. Ribosomal A gene | sequen ces: Analysis ue ignificance in the phylogeny and ийе of green algae. С. К. C. Crit. Rev. Pl. Sci. 10: 343- 368. CRANDALL-STOTLER, B. i000, Morphogenetic uer and a theory of изүне origins and diverg BioScience 30: 5 1981. Mo hoo anatomy of hepatics and anthoce rotes. Advances Bryol. 1: 315-398. CRANE, P. В. 1990. E phylogenetic context o crosporogenesis. 11-41 in S. Blackmore à R . Kno: د‎ ahd Microores Evolution and On- togeny. gure mic Press, London. DoNocRUE, M. J. à Mi р a NDERSON. suitability of m in recons cin qn Arana Pp. $03 Itis, . Soltis & J. J. Doyle eas Molecular onm of Plants. Chapman & Ne STEAD, J. F. Ѕмітн & J. D. P‏ پس e asin ‘of e a eme!‏ .1992 on rbcL se . Missouri Bot. Gard. 79:‏ iss‏ Duckett, J. С. & К. S. RENzAGLIA. 1988. Ultrastruc-‏ ture and development i pe in the bryophytes.‏ EAGHER. 1985.‏ EE .B.M ben anpe sequence of soybean nces of ares small- epit :259- j M: Pn m ” 5 > С — о fo > ® Q = о rRNAs. FELSENSTEIN, 1. 78. parsimony and com pa орту will be к irre ai Syst. Zool. 27: 4 GARBARY, Pura . REN A & J. G. Duc 1993. The phylogeny of land plants: А cladistic analysis based on gametogenesis. Pl. Syst. Evol 188: 237- Ae ENSEL, P. G of the zosterophylls an d m ids: Aon om mor- phology, кое ош, and cladistic rs б: їп- ference. Ann. Missouri Bot. Gard. 79: 450-473 kou ee; L. E. 1985 "The er of c; life cycle of and plants. Amer. "Sci. 73: 482 Annals of the Missouri Botanical Garden , С. F. DELWICHE & B. м oer 1991. Ph ogee ret ape betwi har algae” and the “‘bryoj " Advances мн 3-244 GRAUR, D. 19937 eau r phylogeny and e higher mammals. Trends Ecol. ru mis of pis vol. 8: 141-147. HS adi ОНОН: € WOOD, A. INGOLD, К. KIN à IN. 1987. Ph ylogenetic elton ween resi А aara fer nd oo; c. Natl. Acad. Sci. U.S.A 5823-5827, жш "c. 1977. The Conducting Tissues of Bryo- phytes. J. و‎ Vadu HENDRIKS, L., Y. DE Pex EER, M. VAN НЕКСК, J. M. NEEFS & R. HTER. dh The i ribo- somal sequence of the se. mone Zn. sulcata and its evolutionary position be vec peter Fe Ў Let 445-449, Huss, E. I Phylogenetic М. L. Socin. position of some Chlorella species within the Chlo- rococcales based upon complete small-subunit ribo- somal RNA sequences. J. Molec. Evol. 31: 432- 2. KALLERSIO, M., J. S. Farris, А. G. KLUGE & С. Bur 1992. Skewness and tip Cladistics 8 275- 28 E. C. THERIOT, E. A. ZIMMER & R. L. 1990. e Pleurastrophyceae а nd Mi- ae: A cladistic analysis of sd as J. Phycol. "96: ^n Fn KENDRI er-con RICK, P. & l. Wat cells in oe fossil land aop: aena tr ме еа ven E oluti KLUGE, A 1989. A concern for evidence and a phylogenetic suas of relationships among Peg id ool. 38: ox P rr = Ф 5 8 5 2, x s a 78 AMBARDELLA. 1988. The spor oro- рһуте- ‘gametophyte; aopn in bryophytes. Advances : 255 D. 1 92. MacClade: of Phylogen wnat Char er Evolution, 3.0. inauer nerd ates, а Mas achusetts. Mattox, K. R. & K. D Mi iiu елер idi Classification gu UE C vie & 0. M John (editors), The Sy Alga ade i loud. adr aos M. 1990. Chlorophyte orders 2 uncertain a s: Order Microthamniales . Pp. 652-654 in Margui, J. O. Corliss. an (editors), Handbook ‚ Ме loni n & D. J. Cha of Bio Tones and Bare d oras pee" MINDELL, D. . Aligning DNA Vade Ho- melo à reete... nb Pp. 73-89 in oto & J. Cracraft Сач, ng ai netic Analy of DNA Sequences. Oxford Uni New MISHLER, B. D. 1994. Cladistic analysis of molecular and m Vasa. data. Amer. J. Phys. Anthro pol. 43 —— —— & S. Р. (ctn 1984. A cladistic approach ^ + Phylogeny of the “bryophytes.” Brittonia 36: ]0B5: eT‏ ا Phylogenetic Es pe of de Asc ri and‏ bryophytes. е E 305- , M. J. Don RT. 1991. The cay index as a measure of relative robustness wi a cladogram "nd X. Tor ко Pei [Abstract.] ; К. BREMER, C. J. Hum a E CHILL. 1988. 1 sequence data in ее. reconstruction. boar 37 1391- 395. SEPSRERTS S. Hop veis E. DELUNA & R. VIL 1992. A m eve r approach to the phylo; dm 1 чш Cais ЕТ of chloroplast-encoded "pel nd 23S ribosomal RNA genes. ow улг IREL Мїүлмото, M. M. general classifications. Cladistics -180. сао cladograms and 1: 186-189. O’KELLy, C. J. & С. Ктоүр. 1984. Flagellar apparatus, absolute ser "n, Ey phylogeny of the green algae. ag S 107227725 ALMER, lr К. Jis d ЕН. M CHAS &j.R L. MANHART.. = "Ойгор TN са and plant phyl Gard. 75: 1180-12 PICKETT-HEAPS ‘Ele ctron microscopy and t he and land plants. Amer. s. J. SL T was H. 1 A cladisic Meus of the per’ and higher green plants (Viridiplantae). Pl. Syst. E 149: 317-239. The green algal class Лорен Ап ultrastructural survey and classification. Cryptog Bot. 1: 83-94. б STEELE, K. P., K. Hots: Ra на e 1991. Asse praen 6 i reliability of rRNA eee enetic analysis in cum P ME M 0- 2 48. Stewart, К. D. & K. R. Marr 975. Comparative bea evolution and classification of i? йт ееп T viuis with chlorophylls a and b. Bot. ме 4: 104-135. WOFFORD, D. L. 1991. PAUP: Phylogenetic Anes Using eb Ee 3.1. Illinois Nat. Hist. hampaign, Illin TAKAIWA, F., К. SUGUIRA. = edes арте ан изе sequence Wien a rice vi gene. “Аан К 5441-54 THERIOT, 988. of Sluiman es dm sification x green aves ui Ики, refere wat: flagellar A and to land plant origins. Taxon 913-9 Waters, D. a М. A. BUCHHEIM, R. A. DEWEY ‘ite CHAPMAN. 2. Preliminary inference Qu phylogeny of bryophytes from nuclear-enc 9- bosomal RNA sequences. Amer . J. Bot. 79: е 46 GB Witcox, L. W., . Lewis, P. A. ааг m ssessing the rela ei LOYD. 92. tosporic and zoosporic grea Di i gree! Volume 81, Number 3 1994 ZECH Mishler et al. 483 "Green Algae" and “Bryophytes” with 18S rDNA sequence data. J. Phycol. 28: 381- 3 0 ee R & R. L. CHAPMAN. Phylogeny of Pe: Thr (Chloroph p C ds analysis of nuclear-encoded rRNA sequence data. J. Phycol. 26: 700-7 ZIMMER, E. A., R. K. HAMBY, M. L. ARNOLD, D. A. LEBLANC & E. C. THERIOT. 1989. Ribosomal RNA itors), The Hierarchy of Life. Elsevier, Amsterdam А REEVALUATION OF SEED PLANT PHYLOGENY! Kevin C. Nixon, William L. Crepet,? ennis Stevenson, and Else Marie Friis* ABSTRACT Seed plant phylogeny is evaluated using a data set of 46 terminals (taxa) and 103 morphological and anatomical characters. Cladistic Ar u sing the criterion of parsime ony were p erformed on the T data se et as T Tie A ч placement “of the cy well as extant nkgo, conifers, g etopsids, t Gin and angiosper en жле were LM m an Mihephyie Дыр ae included pas MeL and angiosper but all, of in some, = closely associated ae ^e anthop combinir 1g cads as the sister group of a eph group th fossils were pos pos s. Pentox a sister the most parsimonious tree Con onia vas ot ‘ound to these two taxa for all analyses, Ephed clade, ere e consistently outside of the cua pe clade. In all hos parsimoni " 8 N dU 1:333: 09) Спе т, арі Welwitsch, ome previous interpretations. of homolo ogy ‘of the caytonian “ nd s ra Ei gi a d polymo when polymorphic or “summary” * terminals are suggested rphism are explored in sere data, showing how different results may be used. The А” “are ко, The effects of ire be obtained eed for more work on gnetopsids and fossil taxa is INTRODUCTION ent years, several aer кыгы of Е] pna have been published, with explicit emphasis on Em relationship pr angio- sperms to other See plants (Hill & Crane, 1982; Crane, 1985; Doyle & ске ee ts 1987, 1992; Loconte & pul. 1990) refer the r a sum- s paper we present a new cla- tic ana nalis of sed plant p based on morphol e Loconte and Stevenson. mii ош analyses шше ын! (1985) and Doyle & Donoghue рр 1986a, b, 1987 КЕ encompasses two and goals. first, and most obvious, is analysis of seed plants, fo ysis and a phyl sults. While we feel that we h new insights, a few new characters, and what we re more neutra s (Le., fewer phylo- раве hypotheses pibe. of some previously used ее we fully expect readers to focus y on the topologies of the cladograms merous analyses, show the and propo а ever, the over- t users to "compartment result in solutions cm are not se (Nixon & Carpenter, in press ' We thank К. R. Pedersen and M. A. Gando Stevens and Gar Rothwell for helpful and constructive com: * L. H. Bailey Hortorium, Cornell University, iw * New York Botanical Garden, Bronx, New Yor * Department of Paleobotany, Swedish Mus lfo for assistance during various phases of this project, ipt. and Peter nts on the manuscr di an New M 14853 8, U.S eum d xd oy Stockholm, Sweden. ANN. Missourr Bor. GARD. 81: 484—533. 1994. Volume 81, Number 3 1994 xon et al. Seed Plant Phylogeny MATERIALS AND METHODS SAMPLING a were selected for inclusion on the e fossil мара: pap tee are numbered starting with character 0. The majority of characters (80) were binary; of the remaining e ания: соне 15 меге treat ight were treated as additive ERN: as du deis in Ap- endix A. In addition to the complete data set, several modified data sets were also constructed and an- nee. and or uP of term inals “condensed” Тег n e primitive Mv rm, Cer- м dris icing the first branch in morph: al and E anal- ^ (Betula ), and Casuarina (Casuarinaceae). птуз (Winteraceae) represents the modifi s- seyan hypothesis of C nd an meet bee rovid in an effort to үе. major tli and Provide е connecting structure among the other ter- dix. е lower hamamelids (e.g., Trocho- n, oboe and ma agno oliids. 7 ause n о о combine our data with existing rbcL Separate project (Albert et al., 1904, d TI decisions were further restricted a priori e po of rbcL data for taxa in our data set ызы all taxa (46 as presented in g the convention p Henning86, к. чаш юрме” rege ut cóbdenkations were DAN (DADASB6 v ver. т.0. B шоп, ideal и = ape which the new generated * *condensed" terminal was ed as тушрш; i tor all Шага that would be iis the group), DADA translates polymorphic cells to ambiguous (—) upon submission to the Hennig86 daughter process, but retains the poly- morphic coding ML in the working Mame. X Terminal Henning86 uen assuring accuracy by ing a single master copy of the iiio Pe matrix. a sets generated th the above e ons that we are not The da inclu dod numerous combina reporting here because of зене геена The zonis ma we report here are based on the fol- atrices: I; ‚жга analysis: 46 taxa, 103 characters; 23. ا‎ cit of which 1.36 are polymor- i .51% excluding 11 within-an- 92-102). : 31 taxa, 97 characters; oly- VIS R of which 0.32% are poly morphic (ambiguity 7.69% excluding 11 angio- erm characters, 92- e III. Single angiosperm terminal: 29 taxa, 83 characters; 21. 77% сонна ae which 1 93% are polymorphi (rata Fossils excluded, single angiosperm terminal: a, 66 characters; 6.61% ambiguous, of which are ar 3.50% polymor iiem . Fos а single а angiosperm and со- nifer termin: assin T axa, 57 characters; 17. 8% am- biguo us, of which 1. 54% аге polymorp ot intor for reduced data sets varies. No characters were venen or recoded other than for the condensed OW TY гг LJ 1 f 1 Ф. е reduced бзр when taxa ar eig or condensed. , Some characters are d that vary only DE terminals that fc Annals of the Missouri Botanical Garden trix. Characters 0-59. ? = unknown; — = inapplicable; . = uncertain homology; * = TABLE Data m Mas ан All treated as missing dat Aneurophyton Glossopterids Caytonia Williamsoniella Cephalotaxaceae Podocarpaceae Е га Welwitschia netum Chloranthus iper Winteraceae Calycanthus Eupomatia Magnolia Dillenia Caltha Trochodendron Platanus Hamame lis Chrysolepis Betula Casuarina Ceratophyl Lum Q.—5:.:10.- 15.20 425 30.85; 40 5:45.50: 008 о арыр етир s dépit oe a | Í€— Pisos *0?2200200- —— 0595 epo 000?110011210-?? 0.0?110 ben 100000-0-000- -0???? 0.0?2100 0-???200000011110*0000002??0.101010-0-000- -022?? 210100007210: 22210000001 . 110007000000003101110- xr 2200 .27222 ا конатын *00.0?1110-110??--2799?‏ -130101200 -010 ?0227- -10101020010003111110-11000-. 2222222222222-2227 cara ne deat 01102101110-0-171100727?‏ -???010222?222 1110-0-1111002772 0107100100110 ?01102101110-0-121120277 0102100100110 ???02- -00. -00.0001110-0 .0112230011200-2?2000000001110000-000 ???00- -10000 0112110111200-222100000000-0 0000221100210-000 0-000: 0900 0000121100110 00010 0000-21100110-00020000.011110*00000010003101110- pe 000- -00010 нан. 100000100 0-000000010013111110- bi 0000008 10110111200 00110 0110110111200- -000201* *00102- 0- -0000001*003110111- 0- 100--00110 н М шы Dm 0-000--00110 0110110111200-00020100010*-0-0000000100130 -*0000--00110 02011301112110001000110102-0--00000. МАЧ ма 0-1011110100 1110-0-1011111101 11110100100-0- 101100109 41112n4*n Ifi deuil о -— e 102101 > NJ -d o -> — © о о ооо 2 ч го = =] -> =] — =] =d -> =d ч ыш ے‎ d -d =d =d =ò = =ò -= wh a Gh са md = ERS adt E LAM ЛАЛ 120100- 110112010---11111000 oo o о © © © © o = =з а -d о —> a D -d =) — 141201 110 n PIVETEVIV 0101120101110 1001112012000- => = = D NJ „> © > > е c Ot 5 «4 2 2 NJ > o — n141nnn1 4 12020 00 210101010- --0-011000100100-0-00- -00110011120100-0-11.- --02101 Volume 81, Number 3 1994 TABLE 1. xon et al. Seed Plant Phylogeny Continued. Characters 60-102. Aneurophyton Archaeopteris yginopteris Corystosperm Lepidopteris Tatarina Glossopterids Podocarpaceae Ephedra Welwitschia Gnetum Chloranthus iper Winteraceae Calycanthus Dillenia Caltha Trochodendron Platanus Hamamel i Chrysolepis Betula Casuari Ceratophyl| lum e-ec-ccoreri222000-002-0--2---- 24-5» 11.0-12.01001002000001110110110 0111100012121 11.0-12.0100110. 020000011101101101.-- -1 41..0-12.000010720000011101101100011011-1100 Annals of th Missouri i MR Garden For ex- within т РЖ Е d are excluded in the condensed analyses. mole, бача "n are relevant Mir 92 rms( analyses in in vlich h angiosperm taxa were con- nsed into a ec E as a РЭВ (Albert et t с analyses, fossil taxa were scored as ambiguous (missing) for all rbcL dat Cladistic analyses were ene on Intel 486 ennig86 v IBM PC compatibles using He ersion 1.5 (Farris, 1988), run as a 16-bit DOS daughter pro- cess of DADA386 .87. All ified data sets were generated directly within DADA, ensuring were scored identically in all sep- arate аш Alterbative chaxavter рыгар апа trees for publicatio 5 dos ver. 1.4 (Nixon, 1993a y In order m increase the likelihood af finding а & де 1982; Luc dam & Pimentel, 1985), taxon mly P option of of Henni optio үө were с сг еа HIE di 1 set was analyzed with heuristic branch swapping (“ЪЬ*”) T pin command А es Pucci removes redundant trees from diee Ier y filtering tree files through the Hennig86 алын u" optio provides an accurate count of the r of TOOR found, and allows generation of a 1 th 1 Dye generat by replicate runs Some of the daa sets that were generated mi onious data. For eee data sets (V. 2 re replicat ate randoni tax was unnecess. no MI 1-4 ( ophyton, Ar- LR Ч z 1 j а, 1 06 22 is A Ea А | АП рад меге simulta- out m globally mene dn taset (outgroup + ing рКа ams dide. and ingroup. In analyses that excluded {олай terminals, trees were rooted between cycads and the а м the extant seed dene based on the results of the full ei. ses and ү usly pub- ee ana ^d. (Loco e & Stevenson, 1990) It be noted that such кош do not affect i topologies of the trees reported. THEORETICAL CONSIDERATIONS MISSING DATA Problems associated with missing data and/or роот т have пої been discussed a the literature, but interest | in this topic has in n & Davis, 1991) lues The at nown or not collected, and the latter when a cell is scored as a “mi: ue for reasons grams, includin (Swofford, 1991), NONA (Goloboff varaa (Maddison z kna 1992) an os (Nixon, 1993a), treat missing data с oiim tionally the same e durin optimization and dee all irn n en present Some рг ograms or imonious cladograms (e-8- PAUP and NONA) lam the coding of subset poly- e curr though Nixon & Davis (1991) disc lem in the context of polymorphism coded r Hennig86), these propin are odin Davis (1991 rool t poss hough it is Bet na es Proport Volume 81, Number 3 1994 Nixon et al. 489 Seed Plant Phylogeny missing data introduce uncertainty into analyses, it is not yet clear whether this uncertainty is pre- table, how to measure it, or how much missing icular r analysis before Ф у. Data sets include numerous fossil taxa sree have high levels of missing data as a result of эйел. preserved fossil organs iy the difficulty in deter- as ambiguous for some within-a sperm bonn In our analyses, aia vast ma- jority of cells scored as missin, ing were due to lack of information for fossil taxa; this is reflected in ve us data when fossils were excluded from the sancii disce ranging fr rom annt wipe ngio s wii 6 Ms bocanke COMER aOR generate n charac cn ^ citi terminals. И шато, ARTIFICIAL TERMINALS AND POLYMORPHISM of the most serious problems in previous н d plants has been the treatment of Pc diverse groups as single terminals, which ifficulty when these terminals ar ined when analyses are run simulta- neously versus compartmentalized and/or con- lem v various modifications of our seed p a spem iios the er we раш; reduced polymorphism in our data е for each character (Nixon & man 1991). Compartmentalization may rious errors, which are more punta bias, when polymorphic terminals are not c polymorphic or ambiguous, but осей аге e coded as having the “presumed” the group ee nted by the t erminal. In the Doyle & es e (1986a, b, 1987) analyses, ie sp 2 some other diverse groups, suc aving alternate leaves, 1 d arranged anicrosporophyils (= stamens), under the assump- On the е pia ged polymorphic characters of angiosperm red as missing, or ambig- poms dedi. as | presence of тое. Given Mele the ү & монеи (19862, b pen analyses s did n gios permet to C + te vec 1 hot Jes, whorled mic rospor rophylls, a for: m ог simple- mated vessel ешеш); amina order to avoid such bias, angiosperm taxa for observ d characters, instead of presuming an angiosperm мураа беш. ne T А RS : such an approach has other | TES 2 . Of course, "en mostly due to potential undersampli (see Nixon & Wheeler, as е tee these problems to produce less е practice of designating тыы: states for icr terminals. morphic t HOMOLOGY DECISIONS One of the most serious problems in any broad | UE ACC 1. в PLA 1 1 2%. ы, of homology assessment among highly divergent taxa. Typically, the more closely related taxa are the easier such assessments of homology may be 490 Annals of the Missouri Botanical Garden Seed plant analyses of include taxa with of — such eres a уел of fossils disparate eet oop of reproduct ive and vege- n cases where we have tative structures. Some ertainty exists in ho- interpreted eértidd homologies in fossil taxa differ- logy араай for some taxa for almost every ently from authors of previous analyses, we discuss haracter in the rc ars For instance, whether these differences in some detail in both the dis- angiosperm flow ош strobili, and cussion of characters and the discussion of the fossil whether carpels ar are evn docs аге justa taxa. We als cha 8 the terminal few of the p а of units in the analyses іп li th recent fossil dis seed ен: In order to avoid ee deris are coveries so that there is less character polymor — tein phylogenetic hypothe ses, we hav phism in the terminals. In е instances we have as much as pos sible: on direct observation of features. Whether or not we e succeeded or failed in this effort will be left о the Tap of Ше reader, and un- models shold be incorporated into the Mie Ап example of such a complex model used b previous authors is "s en cod integument of angiosperms and Bennettitales by some previous authors (Crane, 1985; Doyle & arii apes ге S PE T Sae our us ing and in terms of our cladistic results, in greater detail below INcLUDED ТАХА See Table 1 for a list of taxa and the complete data matrix. FOSSIL TAXA ossil taxa were selected for inclusion in our phylogeny. While numerous angiosperm fossils are now known, relatively few are Bie gian enough or different enough from modern taxa to be relevant in a broad seed plant: E pit Сера et al., ү ‚ for s). ер Һауе meer of key fossil taxa below ate not intended to be complete, but they emphasize in- sil i a pule of Caytonia). Other differences od re- ot W бын interpretations а ad made them less de- pendent on a priori phylogenetic hypotheses, but in conjunction with establishing new terminal units. FOSSIL PROGYMNOSPERMS Aneurophyton. An. RI is ei includ ed in phylogenetic analyses of the eed plants as teris and Жета, аге bett ter v known but Md d acters, e.g., pla- nation of ды structures мА Banks, 1967; Bonamo, 1977, 1983), sh dos be in- appendages are three-dimens , two- e times RUM branchlets beta are a th th mate branchlets dic пн опсе, branchlets curve inward and bear two rows $ i sporangia the inner de s. All — оруш ales that iach sufficient pices m found to be homosporous (Tay lor & Taylor con Archaeopterids. «Hn i is commonly © di sidered to be a tru omposite genus (reconstr^ s from more than one fossil) that includes rosis runk: ее *progym te the ant gen pec as the basis of the concept of ` osperms.” Beck (1960) was able to o associa | | | | E E | Volume 81, Number З 1994 Nixon et al. реле; Plant Phylogeny coniferous secondary wood o: (Calli. lon) with frondlike planare seep sys ded s (Ar- аии based on similar anatomical details. stem has a nin of scite vascular bundles with pyncnoxylic secondary tinc- tive bands of c r аз x pits on the radial = of the p ds (e.g., k, 1960). €— al. (1966) demon Lesen that the idu; systems webbed leave s of planate, variously A ud rom axial primordia and not from ax- Various hypotheses link the ieri о его- phytes (Meeuse, 1963; Beck, 1971) wath re- al Rothwell (1982) vis suggested. an MANN мао апа 9 an the исгбёрогап ade are borne helically penultim anches E are кзы їп беа. твена "ates bá Mdh n on icum vum ые nod, 1935) FOSSIL SEED PLANTS ulate cupules (Calymmatotheca), com- pan r Pressed fronds (Sphenopteris hoeninghausii aha and several types of arene Tel psis, an eraxo theca, but terminal in Таана апа Telangiop- sis (compression fossils with t gemi Ment iners чуч ang amis Oliver & Scott, 1904, to cop the ferns”). Stems have anastomosing radially ce bands of Miei in the quier co ortex (Oliver & Scott, 1904). T riability within the family — as се is usually circum- scribed (Stidd & Hall, 1970; Taylor & Millay 1981; ош 1988) апа "endi within od outside of the family will erstood a new fossil evidence e availa ble. gap yses we have used a broad the oe as our terminal, that in future analyses this terminal would be di- vided into d based on obs кайым variation the group. ae previous analyses меме in Lyginopteris as axillary to ities (Galtier & Holmes,: within s have ieee в opposed 1982) We as dichotomous, but with occasional close dichot- omies that superficially mimic axillary branching (see Taylor & Millay, 1981). We do not know of niy Пеш 5 (Шоу hrsncbiag in thi group. At any г rate, b Eka di ity within the pan it is possible that referred to as lyginopterids had manam branching se rms of bra nching. Thus, | we have scored o Waage terminal as unknown for the 5 or absence of axillary buds/branching. ans are characterized by к . Medullos & Taylor 1993; Sian & Rothwell, 1993). )) They pr are only preserve: mpres t rifactions and their vegetative and reproductive anatomy and morphology have been care and extensively studied (e.g., Delevoryas, 1955; Stew- art & Delevoryas, 1956; Шау & Taylor, 1979; Taylor, 1965, 1971; Taylor & Eggert, 1 Eg- ert & Rothwell, apes & Rothwell, 1980; Stidd, 1981, 1 & Eggert, 1986), although complete suites - жый for all of the ‘petrified stem taxa. нә Annals of the Missouri Botanical Garden of the stem anatomy and the origin of the s angiate prepollen organs have been su аа controversy (E well & Egg Baxter (Baxter, 1949) a have a different growth habit (vine vs. tree) a mode of sepes bin is the only medullosan a: axillar er & Rothwell, 1988). We ا‎ ilit: tr саше growth habit that do not have branching. In future a pending av of characters, we add the axillary branchin ERN as an gre terminal unit. او‎ А һу ахШагу con- аларым over the origin of the polystelic ish of medullosans or about the homology of the large e ына pollen-bearing organs RR ONS The Callistophytaceae were first and circumscribed by Delevoryas & ант апа ——— "e күс аар been refully described. Stid ) were able " > demonstrate the платио between ovules of & Delevoryas, 196 0) апа Cindi of Callistophyton. Subsequent eias by Rothwell (1972a, 1975, 1980, 1981) e made the genus one of the best known of the RE einer Callist stophyton is reconstructed as r s. Synangia and ovules ме borne on the 1 {. EN 1 x in c nd ith tl 1l free from the inner r integumentary layer except at sti — Synangia ponen vun a ring of latera ally clude Callandrium (Stidd & Hall, 1970) abd Ida- us lateral oom Te Pollen: is une mis poseen ied bh h (Rothwell, ло) and with apparently intact within the walls (Millay & Eg- gert, 1970, 197 74) that are similar to those in extant conifers. These similarities have led to speculation didi in relationship between the Callistophyta- сеа ^ oe of medullosans with sibi ceae and cordaites and even conifers (e.g., Rot h- well, 1982; Stewart & Rothwell, 1993). This is tophytacean-conifer relationship below Cordaites. Cordaites constitutes a conspicuo ~ © < @ Р - ilies are based on Angar: erable structural variation embers h irk regarding | bj тер ті аге (1988) зя Rothwell (1988). The ru ceae include wood ( Corde Mesoxy' lon), pith casts pp ons roots (Amyelo ‚ шш) leaves (Cordaites), male an (Y ones Lew nd male cones Gotan) e Cardiocarpus, Mitr um, Samarops ri) palen AFelixipollenites, "Flo lorinites, "sullsacies) be —Cardi nad present study, and the Meso Gothania— irago yeu ccites trospermum complex, but koc are other ings not yet fully under: ood (e.g. Rothwe Warner, 1984; Trivett & Кон. 1985, Rothwell, 1988 Тһе сои were tall, profusely branc ching trees, rubs basen & We and sp x h, а d be p or uni how. 0 1984), wi while t 1Y1E€ES0. ically yma ا‎ Foliage comprises large тар-аһаре dM or narrow grasslike leaves Cordis a clase some species also small scale or i around buds, at the base of чё ог ех Volume 81, Number 3 1994 Nixon et al. 493 Seed Plant Phylogeny along the branches pe & Warner, 1984; Rothwell, 1988). There is no mid-vein in Cor- daites, but numerous, densely spaced, Vins mizing veins that run almost parallel in the leaf organization and d ovulate detailed account of the t rally pe Each 1 ا our to five or six nuclei are a I in a single row oriented towards the ish Eos are stalklike, and in the Eos single vascular ers the ovule and = rise te one раш of vas- cular bundles that pas: t and к another pair of a, ar Ише that tud into setae The nucellar cuticle is distinct and rom the integument and thick megaspore “ian one rosporangiate cone, Gothania, associ- with Mesoxylon-type wood (Hirmer, 1933), iS similar to the co in tek E ngle row at the spo- Ra E ollen grains found in situ in Gothania rn d nosaccate, a distinct е rilet car, r to dispersed pollen of ат, еши ог Sullisaccites t Millay ч in 74; Trivett & te vi 1985). Ovules n spermum are lin went ne the nce of Felix xipo е е; - pollen ro Pyles (Taylor & Taylor, 1993). They a are piae to Cardiocarpus in most features, but the integ- ument vasculature forms a characteristic tracheal plate, con nucellar vasculature is lacking (Taylor , 1964). & Stewar Glosapierids Тив glossopterids the signal group erstood assemblage of variously preserved fos: Pant, 1977; Schopf, 1976; Gould & Delevoryas, 1977; Retallack & ed 1961; Pigg, | 1990; Таа б Saylor; 1993 of Crane (1985) and due Б ауе кор ا‎ inter- t studies since that tim 0; Taylor & Todos: 1993; Pigg & 993). a multiveined midrib with anas- tomosing esent. venation that is not hierarchical pens espe ^ to vein order diameters (e.g., 90), and our pue ng of venation reflects this dins. a this chatacter, the edis are Чар form of Caytonia). i eat deal of variation in the nature г < pollen bearing sporangia - on ape the маја хаосу, nage to " eid а у borne directly гонене ovulat als r structures. “ig some vules s minating in clusters of micros, ngia that con- ain bisaccate and apparently quasisaccate pollen (Surange & Chandra, 1975; G & Delevoryas, ; Meyen, sarig ere has been some un- ture of these a on t Ing ard these str пешгев аѕ axillary ‘branch Systems Wi h Val iously adnate toa езбе b bract. Rd in s encoding Annals of the Vei Botanical Garden vious analyses (e.g., Crane, 1985), we have inter- preted the pollen апан ae ттш s ax- illary and not foliar or fusion of fertile ae structures to foliar ap- рене is consistent with the: morphology of r re- asilary e such as the conifers, cin is nsistent with n ex. This interpretation of the seeds as borne on axillary structures is suggested by many = "A : б d f : ] ДА] r t | 11 d Nez 3 t n bearing both our interpretation of, and the recent literature on, — Me a seed romii fonde appendages, as axial systems as дррбвей to megasporophylls Me & ете ч, 1985; Antevsia, Friis & Pederse in prep.). Recent discovery and careful analyses of pet Еа axes ieee Glossopteris previd шы mor- been pae in the past (e. g. alternate (spiral) ax are ed (Table 1; Pigg & Taylor, 1993). The шешу pl within the glossop- terids, y understood at the present time, Fi it li that in future 9 the ey ae have to be divided into more than a single minal, edi based on a sospes reconstruction. as — ssed below, “summary” taxa are problem- } ы) 1 j m E ۳ +h 1 аф Peg TEES | А PAS А if any evidence for monophyly. Peltasperms. The family Peltaspermaceae was es- tablished by Thomas (1933) based on the ovulate ер sequently, a variety ôt dispersed mem "x z Comp - sopteris, Kirjamkenia, Meyenopte Phyl. ladoderma, Tatarina), аана ограпз siderable variation in structure among the pelta- sperms, and in the тее analysis we с consider e Pe e Late Triassic epitaph lae HE ae ори s—Peltaspermum complex d, S = = е LE = = E d S баса 1914; Thomas, 1933; Harris, 1937; Pe- dersen, 1981). he pe niue were probably woody plants. and otherwise little is хний about habit and growth form and no wood has been ~~ in association with the pe Depidoperi pla Leaves of Lepido pteris hie generally been in i c a e tube 4 had folded as a result of fossilization. A reexamr ial fr system with the main axis be wes — e lateral branches borne in a P ateral branches bear scale leaves aminar leaves rdin ng to is т teris | indi sem # teris "fro J Leaves imple or once-pinnate. ind e Lepidop comparable to that of an Archaeopte nifers. ith tely dissected lamina (e.£- $ TOP spar e e, and dichotomous bi nia) in the s pinnate fronds "men been ais ian leaves (e.g., редовна (Са у martinsii, Poort & Kerp, 1 The pec typically very thick in the peltasperms, § p dermal features difficult to study in "s The venation pattern is open di pee 2 shaped or pinnate, pinnules typically se me: a tinct ein; the stomata re surface (Antevs, 1914; Meyen, 1986; Poort & Kerp, 1 1990). Volume 81, Number 3 1994 Nixon et al. 495 Seed Plant Phylogeny he microsporangiate organs of Antevsia as- the L found in situ in the microsporangia of Antevsia are similar | to disp prred grains assigned to Mon- osulcit Cycadopites. They are monocolpate with a smooth outer surface, thick | , and a thick homogenous ektexine (Pedersen, 1981). The microsporangiate organs of the Tatarina-complex are only frag- iei preserved, and their structure is not fully un 1929; Gormankov & UM m" 986; Meven 88). The material con- y of dispersed ek of microsporan- while чы a small fragment of an axis was cribed. k are sim AN to Mns laminar unit bearin VOOKSOn ОГ disper: e of Antevsia, although the ngia is not ver t p чу of Tid variabilis Go lii ten (Gorma ank v& Moyen: 1986). A * Vic with оета Севан & ч s Me RI vulate лага, (Petpermam and Pel- КЕ consist of а main axis bearing stalked Peltate and radi i m ed leaves, and Poort & Kerp (1990) indi 9 P r i» ! when treated as a form genus, Pel- taspermopsis should be included in Peltaspermum (see also Durante, 1992). In the Triassic Peltas- ermum-type organs the discs are wid paced, while in some of the Permian forms (e.g., Peltas- 5, т by Радаа. gocarpus is REE vath thg: mucellns free spes The nuce elar mem- zh em] 1 "n spore ШЫ шы In S. bi integ ГЕТ? | OW. ү of Salpingocarpus tl the open уры area is simple (Gormankov & Meyen, 1986). Neto т D A area of plants first ae vulate ite associated dispers microsporangiate leaves of EREK from the Middle Triassic of Afric The disco ae of Dicroidium leaves attached to HE -type stems from the Middle [лөө of woody habit 4 corystosperms Mere т- белый st еї Previous studies have indicated a а ненна ip between Dicroidium апі Rhexoxylon type other parts of Gondwana (South America va n ngel & e "1961; Stir Pu 196 e his ae that several wood types were present in the corys- tosperms. The leaves (Dicroidi ouem are once-pin- nate with dichotomizing mid-vein (e.g., Thomas, 1933; Townrow, 1962; Anderson & Aderson. 983). mic — structures of Pteruchus ntral axis with stalked late ral - s ill rate by Thomas (1933) and ie er Annals of th Missouri кл Сагаеп ribed for material from India (Pant & Basu, r oly- puedo руй mis bn this ie E йе ыч» VANS in some o the en specimens, but istralian material described ue Holmes (1987). Each ves it c ems ed on pulte material j 11 m is E was bas distinct bractlike structures. ‘The REE nature of the lateral appendages in Umkomasia was lat documented by Holmes (1987) tuns ning study of well-preserved specim f alia. Bracts str g structure. This view was followed by Crane (1985) and Doyle & Donoghue (1 ови b, 1902) who characterized the ^am Ше However, the recent discovery of permineralized fragments of Pt y symmet- pps —— нанар of ne central a axis soe for the p leis реак рн (Yao al., 1992). In our analyses we therefore interpreted the re- н structures of both sexes as branching systems, not sporophylls. Caytoniales. Caytonia was originally described as an п чафоро Ьу Thomas (0929), The association ооа tit — a “fleshy saclike тас ше р, which enclosed several seeds, possibly in two rows. Pollen grains were found on the “ір,” ould be take in — M — until the homology of the struc- ly established. The **cupules" of vm are arranged along an axis (as interpreted by Thomas) or ben rachis of a leaf (as interpreted uthors, discussed in detail below). genera that includes ren (Sagenopteris), s angiate pollen-bearing organs (Са Pondus 2 2). Сау pee by dd (1925, 1933). in seed micro- genera © Р їп ies eiit paw pyles (e.g., Harris, 1964 are consistently associa Midi at vı us localities (eg, a 1933; Harris, 1932h, en 1941; = 8; Reymanówna, 1973). T us, there is rav evidence that these organs a the same tax The Mele bei organs of Caytonanthus are according to see also Harris, 1941 pound with —— net venation (Thom: 933) publis o j AE E (1 f Сау and more ester pollen has ne studied Pedersen & Friis, 1986; Zavada & interpretation of peine Thomas’s original yor EE ” e us with the ос of = 0 ussen model, eee duction € several seeds to one seed wi urved “сирше” of dri would give 9? M s is found in ma” Copia would then inc an ise bitegmic ovule, angiosperms. The “cupule” of Volume 81, Number 3 1994 Nixo al. asd ча Phylogeny be homologous with the *'second" or outer integ- cla- such that the results may d confirm the original model. We have rein nre material of Caytonia, Caytonanthus, and Sag eris in order to better m» (on dh hes ¥: Crepet t from the University : ا‎ at к E by Hen n- drews and Maurice Wonnacott and Cie by жу N. ке rews. We have verified features of leaf venation for i e tees iod: acters for к Саул hus, a EU 1 n sh al d laterally Wn ше formed by the adaxial pm (Fig. ai Our studies ү Мол cop. iled to confirm the tubes/ ү пор (1925) and later the micropyles S ihe. ovules with the “channels i in the recurved di 1973) inte f nked ی‎ rows of large parenchy- matous cells (Fig. 1B, C) وان‎ contin- uous with the eer Me and might also av appeared o be tube: es a Reymanówna о) апа Thomas (1925). It i ed other interesting eatery of the Cayton Pay speci mens is an ute extending betw tween the lip of the cupule and рш рейсе! epidermis (Fig. 76, jai This |^ which esults А (SEM) о F aminatio йы т ۶ (Figs. 1, ^d ie ur mm reexamination has ы certai ain characteristic that have: not publis 9f the features reported by Тоо EEE only with light nd ied (LM Eus upule" or seed de is preserved in us orienta- rie with an аа stalk or more than one row is still ч o of spongy parenchymatous cells, а ту ный is по онч smooth nna ermis. ^ The recurved lip of the пета (Fig. parently {ы contin ther no indication of an e UA wall be idermis of the recurved “lip” of the cupule, T precedent exists in | the pollenkitt observ: retaceous in situ angiosperm pollen by Friis et al. 1988) and сарн ће possibility i is kno wn for assoc with a group of seeds instead of the Es of a single Because of the orientation of the lip and pedicel, we do not ik the Ca anatropous structure in the ovules of а engiosperms. in inpcepermous anatr pous ovules, the micr y the uments and is found distal to the point of d. es when either ign or is adnate to on Somi ith 7 — the Thus osperms micropyle i is formed M one or t boh o e integ- f the contras seed envelope is nde e wall of the seed Vaid (lip) only on one si and ge eb rms the other ap t the stoma. In onia Bein a distal estu 1 пеге 498 Annals of the еси Botanical Garden T FIGURE 1. SEM QUA Rs. of L'aytoaig sp. A Fruit compressed dorsiventrally н the E тев n th h d: oved. : oe ss globose form pm fo compron Scale bar = 0. 5 mm, uc lig 6000. TUE A broken fruit Mn a complete eh aboving chan la cale bar = 0.2 т, UCPC 6 6001.—С. f with the abaxial wall removed to show seeds wit thin, Mer ently borne in two rows pr жш i: t lip. Note also the channels in е interior epidermis that аге more Ж less continuous lig the 0| Б P Lateral bottom of the specimen) from the recurved lip- үзө ы ME ale = = 0.5 т, UCPC 6 Er view of the fruit illustrated in C, illustrating the overlapping seeds. pst bar — 0.5 m mm Volume 81, Number 3 Nixon et al. 994 Seed Plant Phylogeny Е 2. M preparations of Cayton — А. View of the pedicel-lip interface Pitt the гузу іп ће m the POR of the M: есы epidermis, a и the me uini py cas - жүзүнө in the о ning me Not hor the nu mero ys Seale! = ‚Т UC i atta lapin i wenn : р t of a “pollen d o other exudate from —D. Hig fi ‘pollen drop” or other exudate magnification view of the remains of a pollen drop illustrated in C. Scale similar in fo orm toa planate ne f envelope is more ag of the the ¢ stalk or a A layer that would represent Continuatio : see interpreted as borne directly on p» foliar append- 500 Annals of the Missouri Botanical Garden age (envelope), or in our estimation a more co sistent nS a would be that the seeds are е оп stem system that has become adnate to Ы Кн bract envelope) At any rate, given the incurved, not a nature of ua structure, the connation to the stalk, and only partial d of ii) Vier wall in tocum of the stoma, it does not арре ar to bes a жые, z aspe much les tegmi iuo E ER г? Susie (1946). W have ides d the ovules ropous and unitegmic, based o ШЫР attachment within the ed copule ” and the lack ré any discussed below, these characters greatly affected the theirs should pay close attention to the treatment of these characters шыт E is Ашырып $ be a iia leaf cushions 9. the breed branches, elliptical ovulate ll are ед! in a whorl on a rim that е а terminal domelike structure (Bose et al., 1985; Crane, 1985) As used in previous cladistic anal- yses, Pentoxylon is a composite Jurassic—Creta- а ceous taxon based on both eae and struc- > Dj " fa? f > 1 1 s 2 organ genera (бакі, 1948; Bose ‘et al 1985). These pressed leaves identical i icm lum), ovulate heads of v. Carnoconites O 1946; “Bos sit al, ded e been va ariously ор (Vishsi: Мир. 5 198 8 ү Bose et al., 5; Taylor, ). Stems (Pentoxylon) are еле eustelic (Вес et al., 1982; Stewart, ‚ 1985; Та i or imilar Medullosaceae Nee 1985. Taylor т Taylor, 1993). They hav fi primar es are mesarch and are surrounded Wee a but unevenly by e develop- nt ker Papers wood with that ar seven cells in anii ug lack s tra cheids быш, 1985; r & о 1993). There is greater жены of secondary wood ward the center of the stem d this and biseriate jg bord arated by par jdn and these are confluent is ш И Pu rtex, both of which h i no resin canals (Taylor & Taylor. im Tie dially leaves né v 2 cm іп | angen ave an m margin insists of кайыше ied veins. Lateral vedi m the midvein at right angles Me and pane occasionally with the branches fusing near the hort originally reported t cm in diameter with a e delicate axes, approxim imately rt, spira aring а whorl о cm 18 уе 5һо гайу агг po appendages (microsporophylls?) ee w clusters of two to four, stalked, t ck-wall 1985, Та Es r, "1988). Vishnu- at the microsporangiophores were сае аї Hom d into a sha sacs (мане М ++. 110۴2) t € еа 3) re rted 1985) concurred. vee Mire A : p "m al. (1985), who ob: b e "n The pollen is monosulcate, about 29 А sud ameter and a exine eor appears endoreticulate w n inner lame == (Osborn et al., 1991). Monosulcate нес d observed in the pelo of seeds of ex 1988: tes (see ; Crane ‚ 198 cedes Бедир organ (C arnoconites) Volume 81, Number 3 1994 Nixon et al. 501 Seed Plant Phylogeny ciated with P. They are borne te с нн оп aont shoots and may be ovoid-ellip- ical or more elongate (Srivastava 1946). ж heri are ое on short stalks and are пої subtended scernible structu by any there re any Einen e distal short shoot, . The seeds are somewhat flattened. The two- layered integument is free fr phe nucellus bem at ви chalazal end, with an inner bicarinat cle pein are difficult t macerations of с lieae Harris, iones. (1962 testa had a thi mpression fossils of C. cranwel- ) noted that the sarco- e was no preserved Hr A lated delicate cuticles “in the bottom of the hn ce poty that he speculated anight BPS Kf. inner nce for two i cause of its unusual nature and the man questions that still surround an interpretation of its тогр pu Pentoxylon is in need of continued investiga tion Bennettitales бане e Bennetti- tales include two aceae and J cadeoidaceae ены & Ver m ents and e an es - ture of fossils that are well known (e.g., me species of Cycadeoidea) and t that are Cant, but ectl la y ielan- diella, Неси, 1909; inanth Stu- E usel, 1948; een OY In some es it is not entirely sen that thag taxa are cen f Wi etn те. elle specimens at the Swedish е stural het (Friis & Crepet, unpublished) revealed that the reproductive structures are not ell enough preserved to d describe; | 2:4 [e nnot be ruled out, nor can the er condition be absolutely ruled out for these c In spite of the uncertainty surrounding sever ef зан the кше һауе been the sobjeote of acters they share with flowering plants. They were first described in the nineteenth century (e e^ Buckland, 1827, 1828; Williamson, 1870; Mac 94) and became ver well link between the angiosperms and M iue ancestors ie i & Parkin, i. This was, a f the most wide preciated мса "n a field of ааа о the botanical sciences. The complex bisporan- giate PERCHE of Bennettitales ше “additionally taxa and E ence of insect ы їп, аї bu Cyeadeoidea (Crepet, 1972, 1974). Bone ше les аге wn from inda of sinon and st and a patafieations from a wide range of ge Sabie bt рган шош ош America, Europe, India, 1963, 1968; irap 1974 Harris, 1969: Бона еї a. 1989; Sahni, 19 ды Bennettitales vary in ue from delicately branched: topps үш persistent at о © ы. -ER bust, , sparsely branch col eaf bases as in s Williameonia (Sahni, 19 columna © ч un © Е = ate within ene ts They may either posed at pnr branch, ite als e — only Мер bra r be borne ter- and subtended 2 two са branches —— or rminal on short lateral hes that the 2 rt shoots bearing the con far enough to elevate the cones beyond t the bases that persist on the trun ks ат. Monanthesia). As currently interpreted, reproductive struc- 1 R ttital iat of alate recep > Bennett р tures in the Ben: 502 Annals of the Missouri Botanical Garden tacles that are fleshy and conical or dome-shaped. They bear hundreds to thousands of ip ovules, sometimes with elongated funicles, among nd the ar- the tangential walls of al scales apices. Tubular micropyles extend beyo mored surface defined by s the ovuliferous portion of the receptacle (Cycadeoidea, Williamsoniella). Sporangia are enclosed in elongate or kidney-shap re n borne in two rows Vos d ae se ea 1906; Delevoryas, 1963; Cre Dove id taxa are well known well ey em (Cj eda idea: Wieland, 1906; =x pet, 1974; hag EP E Thomas, 1915; and Sahni, 1932). Ho non basic nid these involves e a Upper rinse Benn ettitales Кз = there is се: some question ied (cosex sexual) rsus unisexual piae structures in Ben- nettles. ме latter is not as simple a question as r to м In мата differential of the pollen and ids poorly gd so ү association of male and female ar from clear in these diii Nonetheless, it does seem unlikely that of them could have been cosexual. In Var- 1911). In some specimens, however (W. sa lensis е is PS сеў & вие denm, the nature of 1 these microsporophylls выб пена from a den broad cylindrical axis as might be ех pected nettitales have icons lang: bi Le is и. ual (Crane, 1 whe he fossil ele of Berit til if one ех- cludes Wielandiella as too poorly under: to be unequivocally bisexual (and we bee and if i h fossils, but no other seed pla taxa, of necessity, һай а relatia high number of missing уа With r respec to Bennettitales, our analysis dif- fers from Voir us broad seed plant analyses in the explicit t xa (reconstructions) we included instea ead scoring the gr as ambiguous states т Men as was Doyl wae (1986 87). On tainty about cosine structure or affinities (e.g иши fossils, nettitalean or БЕ) poorly understood а 1 sel, 1948; netticarpus, Harris, 1932b); ticular poorly known taxa would ex ex (2) inch sively pont deka, the best known of the Triassic taxa with characters used in our analysis us, 7 аге ci to the ovulate al me ov ulat ate re- — loeftia was excluded from our final analy “a pt РР і cause it generated massive numbers of cladogr a and have therefore d to b митти esulting in deresolved consensus trees by un! ны чя et al., ae Per donum the best evidence with t ny disparate groups (for a d 1991) for unisexual ~~ ttitales is e Williamsoni- of this pna non, see Nixon heeler, aceae f Weltr. ама а form сар We differ from Chine (1988) in our interpr for the petii microsporangium, appea tation of the distribution of the corona WI within cole have been dispersed without ovulate parts. Tus Bennettitales. In contrast to to Crane (1988) " variously ified pinnate microsporop Сувада аз һауїпд а согопа . vulate receptacle, аге only W. whitbiensis Манай st; Nathorst, Cy ilia However, some species Volume 81, Number 3 1994 Nixon et al. Seed Plant Phylogeny eoidea do not have coronas (those with dome- р. п eval ated until own about the structure of е corona in a range of taxa within In Cycadeoidea, the corona is not a fleshy extension u de of мы Wongated ишед interseminal scales that hav to a бам іп ан tissue іп tonifers ome 1974). The nature na out in some species It is in interesting to note that pee interseminal scales similar to those making up the corona of К. are also found at bases of the ovulate receptacles. It is possible that these areas are бея f, J 2 £ 1 Lat isht 18 be associated with pollinator attractior EXTANT TAXA Cycads. The C Cycadales are treated here as three nee S Cycadaceae, Stangeriaceae, and Zami- "ee н а шеше is роды. d a —— t tri e d ancestral states ай the e clade bser Mi atterns, not ideal character th (Nix n & Davis, 1991). Because of е relatively low Жш of s cycads, SA the nature of the characters used in were able to treat the cycad (and ‘oui oe A тЫ s ndis its scored for whole ups) а - ples, as we iia — morphis LEA e ‚Б combinations (N ach of the three cyca - mour s monophyletic. No fossil Kame were inclu our analyses, and the res assigned to the three cycad te als are single cycad terminal as having simple leaves on the presumed cycadalean affinity of a ow kegs fum The шу of many кы fossil wy best trate onifers. As with the cycads, the extant conifers were not treated e a angle terminal but ra ther a excessive scoring of polymorphisms. The selected te ( ph ceae, Podocarpaceae, Taxac and Taxodi- aceea Cupressacene) ai are consistent UR сирр) 1апшу шп aa ee t, 1987; Miller, 1988) at the generic Els hough ku inter ipee relationships ob- ained k Hart (1987) and Miller (198 quite as cue bus T of E Taxodiaceae and Cu- pe ingle s cir inal mscribed y Eckenvalder (976) is is supported by үк ши complete analysis within conifers, and an ny co es on fa ips relationships within the conifer clade are tentat е extant ры of gnetopsids lwi Gnetopsids. The thre ( schia) were treated Ephedra, Gnetum, and We rminals i were fused into a single terminal. Particular inter- pretations of characters for gnetopsids are dis- us: ndix A. below and in Appen E The full data set included 18 an- Th in our ma- yses, the 18 angweperm termi a into a single * ‘angiosperm’ torppa al, which was 1 1 that e» scor р among the original 18 terminals. CHARACTERS “complete” uctive mor- Th characters used in ou analysis i de vegetative and reprodu ose k anatomy, and palynology. an —— rted of characters in recent ae, paria y those d Doyle & "end studies of sud Loconte & Pose (199 a) ао d our analyses, and we lave scored differ- ently several characters that have been used in s. In some cases, the differences previous analyse Annals of the Missouri Botanical Garden from previous treatments involve corrections of errors or the addition of new information, either published or from our reinvestigation of certain fossil tax. nless ihe noted, all multistate characters have been treated as nonadditive (= unordered). For таи" ѕее Арре ndix Obvi y, there is a vast amount of literature source of characters and states for е cgay taxa. Fortunately, there are also exten- sive and seminal compendiums of this literature These works have made the extraction of dat much easier and we owe a great deal to our pre- dec pores: s Amo ong m iners ир, of xps for Бега ( 1935), pese (1985, 988), Сш nquist ( 971), Meyen (1987), Gifford of character data c med to be fro works; as осн аѕ possible, э Һ ргітагу 1 therein, as will often be apparent by the citations in == A and in the section describing fossils above ш шее RESULTS OF CLADISTIC ANALYSES I. COMPLETE ANALYSIS чөн tree PME were found based on ran- domized spins of taxon order with ste and Hennig86. There were a total of 834 h 339 .78, and consistency index (CI) 0 Maren of these cladograms is presented in Fig- ure 3. А representative tree selected mona among Figures with characters mapped. Portions of two grs er trosa are presented in Figures 7 and 8 to illustrate varia ui де of Ginkgo, Pentoxylon, and oth- er see s. If only thee e consider: va ensus insita the lack of a es peo odes group and the A of Ginkgo, which in some trees (e.g., Figs. 4, 8) is outside of the conifer clade + о and in other trees (e.g., Fig. 7) is a member of a broader “conifer” clade. The "higher" or г Mesozoic Sides beg are ин y a la Me that includes Ginkgo as we ds s the e co- ‘anthophyte” clade. Some of the relative anı‏ ا et: ements of taxa — to this ge e illustrated in Fi ; 5, 7, and 8, with Miei mapped to 9 character 9000 П. FOSSILS EXCLUDED When fossils were ed six most parsimo- nious trees were foun зА СІ 0.47 апа ni а ДУ No tree islands were по trees is presented in Ете 9, rooted to the cycads to n reference à the full analysis. The con- i l, with polytomies occurrin ИН the angiosperm clade. саз is most p and the more complete resolution within the extant pl with Podocarpaceae as the first branch. III. SINGLE ANGIOSPERM TERMINAL When t he 18 angiosperm terminals are con- i the is presented in Figure 10. In con plete а the pasion of Ephedra rela to the othe with е to the i The — taxa, which were unstable and joined in a large olytomy in the REE analysis, now show a n t gen a es and ss e ossopterids form phyletic gro IV. FOSSILS EXCLUDED, SINGLE ANGIOSPERM NAL Whe the зони were excluded from the data set in ш III, with a single con 5 otn nly one tree of length пой 0.78, and RI 0.85 was found. This tree oes is identical in topology to thos f ın ا‎ II (fossils excluded). If only extants are consc? it can seen that it differs fro om | — ad 105) condensed) only in having Г ў angiosperms the relative position of Pod alotaxaceae within the conifer clade Volume 81, Number 3 1994 Nixol Seed Plant Phylogeny Aneurophyton | Callistophyton Co orystosperm черинде —Tatar a ——Сауїопі Cordaites +——Ginkgo Cephalotaxaceae Podocarpaceae Taxaceae Taxod Cupress Araucariaceae Ephedra Williamsoniella Cycadeoidea Williamsonia idi ates Спотот found for matrix I (full data set). 506 Annals of the Missouri Botanical Garden ЧЖ-Апеигорһутоп HT-Archaeopteris Lyginopteris 4 ri-Medullosaceae Cycadaceae 22 Stangeriaceae Zamiaceae 5 73 82 - & £i- 5 81 4 81 ГШҤСа!їїзїорһутоп 19 83 eas Corystosperm cetus Lepidopteris Glossopterids Caytonia Tatarina 5 44 63 5 -ШШЕСогдапе$ 40 46 39 57 i Ginkgo 41 62 23 48 17 60 91 inode 75 Fic f the most анат trees found for matrix I (full data set). Сһагас ape numbered кан oe bale hash pe qu м аг улп in some cases where кн 2 character may occur at nodes, but length o pt tion. Solid blac k hashmarks r mee resent unique origins of states; open has тык t d lir d h t q y under tne о The FOSSILS EXCLUDED, SINGLE ANGIOSPERM AND of these two trees is presented i in Figure 12. ALS CONIFER TERMIN о о Condensing the conifer terminal in addition to ОЁ the conifer clade + anthophytes, or h 2 NT | д | RET | а x x with fossils excluded resulted in only oi trees of ered, this analysis produced the same two tof length 97, CI 0.85, and RI 0.87. The consensus gies as were found in the complete analysis. Volume 81, Number 3 1994 Nixon et al. Seed Plant Phylogeny Podocarpaceae Cephalotaxaceae Taxaceae Taxod Cupress 4 23 29 43 59 86 ПИП Chloranthus 17 22 95 (I -node 61 35 Middle crie ga = tree in Figure 4, one of the most parsimonious trees found for matrix I (full data "i Mna! as in Fig DISCUSSION Th SN * general pattern of our results for extant conforms most clos sely with the previous re- in cycad a the — ie "E" sperms. Consid- y extan! ly , Gink- go was ci either a sister group je the боой: or of Ў gnetopsids, and ап- чер Дару por latte Т Loconte & ei: (0001. and by us in the ou i à mainder of manipulations (excluding fossils, Psing angiosperm and conifer clades to single conte itt. on results and those of Lo- favor recognition of a we sites us that includes Ginkgo, coni- fers, gnetopsids, and angiosperms. In our opinion, thes ese results help resolve the pest say i in ree г that аге ин with the conifers, and mus considered AE e i т» topsid— — clade. Beca tail by Crane (1985), — пово Ь, 1987, а ии ee (1989). Annals of the Missouri Botanical Garden data Upper section es tree in Figure 4, one of the most parsimonious trees found for matrix I (full set). {ез as in Fig on some of the sting clades found in our analyses 11) Ае, 4 3 whic К аге outlined i in Appendix our results e in some of the most parsimonious trees. When fossils are kgo removed, these results are not found, and hee. er taxon to the re emainder 0 ormed a remainder p the "anthophyte" cla Sd of previous exception that the position of Pen we able depending on the analysis, ineo is ni P Volume 81, Number 3 Nix 1994 7 509 Sead pes Phylogeny o [-Lepidopteris 55 $ ч Таїаппа 39 57 62 91 2 49 48 Cephalotaxaceae Podocarpaceae Taxaceae 4 a Taxod Cupress 23 68 Araucariaceae [-Pinaceae 23 8 ПШЕРептоху!оп 24 FER ; [-Williamsonia 17 40 76 Williamsoniella 24 А i-Cycadeoidea 1 40 74 nini Ephedra 25 32 55 76 26 45 77 ч д Welwitschia 43 55 2 275 иш Gnetum 29 43 59 86 8 15 31 40 51 56 66 75 91 MIX -node 62 "eus 7 A 13 27 36 42 53 65 70 81 Middle section of another most parsimonious tree from analysis I (full data set). Hashmarks as in Figure 4 as th pe чр sli taxon to фо! соге e Wesce is С i Fi чем (Fig. 8), as reflected in the e in "X . This differs from the placement of Pen- n as a sister clade of the iioc eon topsid group or solely the gnetopsid group in the no nalyses of Doyle & Donoghue (1986a, b, 1987, 1992) and its closer association with nnettitales — is undoubtedly © Б codings acters, especially a гелін alcun of the 510 Annals of th Missouri onion Garden ЧЖЯ-Апеигорһутоп 8 {Archaeopteris Lyginopteris Medullosaceae Cycadaceae 22 Stangeriaceae Zamiaceae 3 4HCallistophyton Glossopterids 2 . [-Caytonia Lepidopteris 24 68 Corystosperm 5 1141 62 9 24 61 67 HHIT-Cordaites pone 17 60 91 -inode 74 FIGURE 8. Basal portion of a third tree from analysis I (full data set). Hashmarks as in Figure 4. pulate st f C all of our analyses, the gnetopsids bes a second integument in P and 1 the an- кыеш mens: were ape en giosperms. Our conclusion that the “second integ- i most parsimonious cladograms. However, ! ment" of Pentoxylon has not been conclusively should ч no eve saa чл еез idi a monophylete Suet, resulted in coding of Pentoxylon as gnet y two steps udin than o uncertain the presence of an outer “seed en- most parsisiorióts s trees in our complete analys? velope." When fossils were excluded and angiosperm* rms con Volume 81, Number 3 1994 Nixon et al. Seed Plant Phylogeny Cycadaceae Stangeriaceae Zamiaceae -—Ginkgo Podocarpaceae Cephalotaxaceae Taxaceae Taxod Cupress Araucariaceae Caltha FIGURE 9. Strict consensus of trees found in analysis II (fossils excluded). 512 Annals of the Missouri Botanical Garden r—Aneurophyton —Archaeopteris Lyginopteris Medullosaceae Cycadaceae Stangeriaceae Zamiaceae Callistophyton Glossopterids Caytonia Tatarina r—Cordaites Ginkgo Cephalotaxaceae Podocarpaceae Taxaceae Taxod Cupress Araucariaceae Pinaceae Pentoxylon Williamsoniella Cycadeoidea Williamsonia FIGURE 10. Strict consensus of trees found for matrix III (angiosperms condensed to single terminal). al 513 Volume 81, Number 3 Nixon et al. 1994 Seed Plant Phylogeny Cycadaceae Stangeriaceae Zamiaceae км ee Angiosperms Welwitschia Gnetum Podocarpaceae Cephalotaxaceae Taxaceae Taxod Cupress Araucariaceae Pinaceae FIGURE 11. Single most parsimonious tree found for matrix IV (fossil tata excluded, angiosperms condensed). Cycadaceae Stangeriaceae Zamiaceae Ginkgo ——Conifers mea Angiosperms Welwitschia Gnetum FicURE 12, rms and conif ndensed, fossils excluded). Strict consensus of trees found for matrix V (angiosp Annals of the Missouri Botanical Garden nsed (analysis IV), trees with a monophyletic o e microgametophyte, absence of the ei b non- bifid i inner integumen t OVE only in ) n e 1 e 1 cellular г early Madii These characters i in are аарыг аз = = s known angio sperms; and E the Case - Wel lwit ehia- o sperm grouping include the presence of a tetra- sporic ا مدا‎ ыыы In all of the complete analyses of Doyle & Don- oghue (1986a, b, 1987, 1992), the gnetopsids have been found to be monophyletic, associated with a RSS Bennettitales. Donoghue & Doyle (19 , list- ed ашушы features” of the three saa gen- as: “opposite leaves, circular bordered pits in dé vessels, simple microsporophylls, one termina ovule, and compound strobili." All of these f are either found in one taxon (sometimes in combinatii; conifers, o or can only realistically be coded as a in gnetopsids, iguou angiosperms, or both (e.g., whether de angiosperm n ower is mee eted asa compound or or simple sitos moak ‘and conifers г are ene! interpreted а as ving compound megastrobili). We r fro o; seat s a the in Ephedra and Wel- ovule” of Gn etales, likewise order to be scored as not homologous with certain angiosperms that have solitary, "terminal" ovules (e.g., Chloranthaceae, Piperaceae, Lara, Hess viens phylla- ceae, to name a few). Such characters are synap- omorphies en the gnetopsid clade only éd he зев а ti observation a broad arr rray o . Thus des uming a pane phylogenetic Aypotbeds b the ойо. ovule es, foliar carpels = * simple strobili, shernate sids and an giosperms, and simultaneously bias their analyses i2 f £ nophyletic Gnetales. This urn has affected ie optimization of the E angiosperm node in their analyses, which was then used as the basis for rooting later analyses within angiosperms (Donoghue & Doyle, 1989; Donoghue, 1989 Based on our eats and the conflict of these results with those of ум studies (includi Loconte & Sone 1990), question of broader amples nd addi onal Pas ice an can be scored ambiguously for gnetopsids and angiosperms. While our study has e be ; to the limited sample of angiosper: includ led, the question of gnetopsid M ог o: pari is now reopen The ене и of whether the seed plants are essed in our sho addre with а э larger sampling of both fossil and extant plants i l s Thus, we f monophyly 0 : n dem ge ed e puis e to shed in and gemere 1987, jn De results are co d ditional rinse of the mien ofm c cads our ea is : contrast to du position of y in Doyle Volume 81, Number 3 1994 Nixon et al. 515 Seed Plant Phylogeny and cycads (e.g., Stewart & Delevoryas, 1956; pns Rothw ell, 1993; Taylor & Taylor, 1993) and m milar to the rene results of Cra 1985) er mor млы t cycads may have orig- or - resem ^is d medullo- ram re e find it difficult to envision е ico ni ideni & ены at 1986a, b, 1987 1992; Donoghue, 1989) that the cycads "PNE scam group — *anthophytes" excluding the ccm and Gin nitially, we се ы, d four Mesozoic seed fern p in our analyses ( asperms," *'corys- tosperms,”” rtl ata and "caytonia"), de- fined in essentially the nner as Doyle Donoghue (19864, b, 1987) In many of those a monophyletic group near the Cordaites-conifer clade (Cre epet et al., 1993). In our final analyses, however, fully de- be specific reconstructions (Caytonia, Peltaspermum, Corystospermum), as 9 umma f grou added Ta- be a *peltasperm"). Of ne- со: re cessity, the dx ssopterid terminal remained a sum- mary of several MUN - we hope that in Mis analyses this can edied. In the anal- y eported , the ^ni Mesozoic seed f: here rminals do not rines form a monophyletic group (e.g., Figs. 3, 4, 7), although they do form a monophyletic атир, outside inkgo, in some th omplete of the most parsimonious t of ou et is. In our complete analysis, the peltasperms and cor ious topologies sometimes in нарсан Крин iie clade and Ginkgo sometimes above conifer dale, аы ding the shes ae пае] clade. Likewise, Pentoxylon is fo associated with t 5, outside rd oM conifer-anthophyte clade, in others Fi Glossopterids and Caytonia formed a mono- hc gro eon outside of the ginkgop ý of t A analy 15 E Ў Tux b d of the most par e trees i analysis cea ondensed s ым erminal (III). This e below” th ча Ginkgo BRS ea ade in т re in all relevant analyses wi anthophyte" clade in some previous us M yle & Donoghue, 1986a, b, 1987, 1992). interpret these differences to be due to different d of structures (e.g., the **cupule" second integument interpretation of Doyle & Don- new data e literatur rom vestigations (e.g., the. axial nature ий the repro- 1 У ssopterids, Е] їп 1 previous ecc end our r restriction оГ. some resent specific reconstructions, + as opposed to syn thetic *composit In the context of a possible relationship between © Ly 1 CL "Pu NE A PERS e | С) a o о the Middle Jurassic of Oaxaca, Mexico, in 1975; Tay dertaken as opposed to treating them as synthetic groups, is long overdue. find the traditional placement of Cordaites with the gnum clade in оше, int not tn dio our fossil analyse h clade, Cordaites is the “n We anc a as often nalyses ols 1985; Doyle & Ф eed cur th Qin-er (1993) that the = reproductive structure of к, сап as an ax- illary se уа keat, these bon a season. that cpa be anor ie the determinate com- pound c xis of conifers (but not as densely aggregated nor ; completely determinate). Thus, the en ended Бы МАНЫ of the “conifer-anthophyte group; a h an interpretation, the carpel of an > spe eriv a se зр (simple сопе = placenta) subtended by a leaf. Such mologous to the bract- a structure would be us vuliferous scale complex of conifers, the leaf-seed- bearing-axis of Ginkgo, and possibly to the leaf- 9 Annals of the Missouri Botanical Garden © А. Ah ] 1 ү 1, uld be general, suggesting that the 'subtending leaf of TO and кр ossibly the ** Сау} are homolog sh of Caytonia and an- nae ashe with i ane ‚рсе infolding of the bract gene iste E ERI günizatiori, dat also, because of aw position of Caytonia and glossoptesids, SAR, that ca рыч їп a © Ж; © iC a 1 1 E: ovuliferous scale complex of conifer: It is in t sal position of Tax- 1988) or Paca (Hart, 1987 the hand, it is interesting bise the Polóvuipicl, "Серина асеае, and respectiv in our IM chum systematists have sug- 1 4 аа 1 к three барди, but without synapomorphies to unite them. A plesiomorphic assemblage perhaps best expresses this situation The position of Calli isp by toi alk of our fossils was consistently below clade, including the Permian- ‚а: above” the с pterids, nes medio. We pain pon de that include *ginkgoph ycads, ly- ot find ê чй as has been proposed by some authors (e.g Rothwell, 1982). However, some characters shared by Mesozoic seed ferns and at least some conifers 7 X E г а callistophytalean-like ancestor; this is consistent with numerous of the trees found in our complete analysis. "Pulse 1 Haba dod 1 uate patterns within E nN and our sample nclusions about — our ir, sample pite Medii itg has been p 0 seen, onsider our results to be of the р eliminary nature of all nalyses up to this point, ect numerous айе ыы from our results relative to other seed plant remains as Gounod belo ow. We seed bine ios and e cupule" of within angiosperms in future analyses. Certainly, m (e.g., Arber & - In more inei mie: studies emphasis h “first branch" of the angiosperitis: While such Ter have iate inted out that: he sister gro branch" is equally the “first im H "hi is probably d ton veleni to such "first" or "basal" 1 32s hes” or **most branches ne o керешүе r terminal” but unfortunately, such erminology is un If the ена group: is Pivi relative to the angiosperms, then ur best estimate of the primitive angiosperm simony analysis within feke. Unfortun affe hes by par- ate- ly popular notion we timate hec = estral state of a group by rence, if ме е: the topology of (Nix sampling within groups and of ingroup and outgroup p panor when samples are of ^ reliable and stable results is still an area of activ rch. rn. How to determine hread t DADA, constrained analyses were inpu ith A, tax: constr: DA a can o be grouped into certain topologies, but tax may also be allowed to “‘float” by assi — pon group membership variables en th yea with constrained to form a monophyletic group er: Maecen and vigencia but allowin are à re "n sit i = o a data set. the tree was two ‘steps ol r. When the t oghue was constrained to the results of Doyle & Don Volume 81, Number 3 1994 Nixon et Seed vie Phylogeny (e. 1992) for ceu and anthophytes (mono- xylon part of gnetopsi a sister group to a monophyletic cdi 0 trees are at least 16 steps longer. f the pteridosperms are allowed to “float” in such they are a basal part of the — trees, surprisingly, 3, se Г Ginkgo I if they are constrained within the cycad—antho- e е 16 steps, depend he extent analysis is constrained. Successive weighting of the complete matrix (Hennig86, xsteps w com results tree: nkgo is placed outside of e fo шер ina condensed 9 а E terminal was Re n jo F © I Malis hel © g? the B ani clade. THEORETICAL ASPECTS The results of various manipulations of our data s matrix, by ex xcluding or including fossil гер апа usin mg mp: Sine into gus olm te ийа" уе ve: 1 yses. It is clear that sample size iid distribution of taxa are extremel rtant. Compartment dimid result in v compartment tied ne phyletic i in is uncompartmenta lized чайна y dien is vpn ese he- Frog are independent of pei а теу тау be; the fact that sr changes in E o ص‎ چ‎ c 4 Ф [LÀ = © et @ 8 & БЯ = E I] m =” [t] ^ [*] = Ф wing more possible à within the angiosperms, fewer ا‎ re found outside of the angiosperms (е.р., toxylon always basal to th lymorphic doe a essarily allow more poss аы for its relation- p to sd taxa, and likewise does not sitim in ie loss of jie resolution. The mbinations of polymorphisms SE va гези ict or P ways that ijp а iste tractable NA Pies сагча of ingroup vari- ation and breaking up of ingroup taxa into mono- thetic units. UNDERSAMPLING rsampling of oe. mp that is not per- esult in increased ҮП Homoplasy in cladograms (error, e.g., the inability in deviation from perfect hierarchic structure; the greater the level of homoplasy, the greater the problem of false hierarchy icum iated wi n. pue sampling. I here, homoplasy any be due to an n inability to о structures, or inability to see that two лийше are actu у homologous in Ms ago, oplas x 1 14 snou nature di in the interpre- tation e characters (the ES a parallelism Mig refer to evolutionary interpre- ). Of necessity, our analyses, like other lyses, suffer from this prob- lem, агып їп our limited sample of angio- sperm taxa. тен within angiosperms, we and ¢ conv e Flour distributions for large groups of taxa, since at this рош nsensus among the results of which s ae йа тыра ry problems here. Thus, оре rns may change and stanly stabilize on particular pat- both within and outside of the angiosperms, when samples become sufficient. It is impos ssible at this point to predict what ae sample sizes MISSING DATA e can be little doubt that missin; date there is no me as tainty, nor to predict its cutn ۸ widely held belief is is that missing data will merely increase the number of equally ae ioe but шч ш оа lution уш Still would be foun initely not ge case (Nixo & Davis, 1991), and is ыа to the problem of missing te were known. This is def- e also Nixon Annals nut Dort Garden ее and homoplasy discussed above. if there is no homoplasy in the data, sub- will marta missing data for real die only re = in less grea trees, the strict consensus of whic will not be in conflict with the orig es boi on full data sets. However, with dines levels of h бон increasing le f missing data in- crease t i that some or all trees found may be in c dun with ^a original trees. In ex- an be extrapolated from the examples n g ET the states for basal groups), when con- sidered ambiguous, may actually restrict the det ly parsimonious тро, by allowing the ter to join another e that reduces homopla gren Wem: pad hare been scored propery. E data is thus severe, and я we have not t escaped it in our analyses, which have level: varying from 8 to 24% e of е The 29 СТР of 259 ambiguity in the data; for istante: we scored pin characters bs WP ad i. within erms (e.g., fl l (represented by a daoni in Table 1) for non- vangio- sperms in ай analyses, such missing data will not have the en they actually е contribute to placement of taxa. This area needs * Hh É f 4 А Bi 1 1 аз inapplicable when all applicable scores form a кешедг? group should be reported 5 T from other kinds of ambiguity in qu dat CONCLUSIONS Our analyses support the Loconte & Steve enson (1990) results in terms of the p throu, various manipulatio of our data. bed we done r connection of cycads with t rm clade psid-a found in ae нехен of D Doyle & паа (1986a, b, 1987, 1992 si nalyses are not definitive in regard to ырдар of angiosperms to gnetopsids and nnettitales, it is apparent that a closer relation- a CARES deg кере © has been pro rear in recent — must be co sidered, and the ibility that the ees group is parap rly a attention. Because of the major role that mo must play in any total evidence analysis, and because vien: data are € ipee to morphology, our eventual understanding of the origin of angio- sperms must be based on farted detailed com- seed bud which were -e in results of seed plant an e (1989) joined ve separately produced р clusions from these data. Any rush to ™ : lu ае scenarios onto existing cladograms se rather fut ithin angiosperms, prie — pe bees in^ LII es tomes wera the “firs eram c pete 1 a DUX h a ini F © PF 1 ters d "у = : ac g the branch under consid- eration. Fin inally, we urge readers to view our results only as a preliminary basis for exa ange of hypotheses about plant relationships. 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Са rd. т jn 55; f the nt cy- ао Bonin 44: 5s 25. 1993. Phylogenetic wave s of the cycads. nd Int. Conf. Cycad Biol., Palm évite n (in press). olystely, primary xylem, and the Pteropsida. Birbal Sahni In st. Paleobot., Luck- now ——— & T. DELEVORYAS.. 1 е medullosan pteridosperms. ved Rev. A 45-8 3. Passus and the Evolution of Тим», ce Univ. Press, New Yor Stipp, B. 'M. 1981. The current status x tnr y seed gehen Rev. Palaeobot. Palynol. 3 7101. 1990. Fur the r documentation of thes т, of D ritique of other ot m M занат Abe B, Palaophytol. 2 ———— & J. W. Haw. 1970. Calla od siis phytoides gen. et sp. nov., the probable pollen- ring organ of the seed fern, Callistophyton. ee. HANDRA. Mo rphology of е gymn ospermous fru oe ‘of the Glosso ор- ic flora and their oreet ете hytol. 149: ORD, D. L. 1991. PAUP \ “ain on 3.0. Program чц Natural History Muse- mpaign, I 1910. The anatomy and тег of the quem SYKES, кы а mirabilis. Philos. Trans., Ser. В 2 Takaso, T. ‚ 1980. A developer ed of the i ond Ginkgo biloba L. J. Jap Bot. 55: 14-27. ——. 1981. A developmental study of the in gymnosperm: s 2. Pinus ейемги Pel, Abies ee Mast. and A. veitchii Lindl. J. Jap Bot. 56: 73-89. 1904. Structural changes in the apex of t female strobilus an и initiation ‹ of a donne ii gi dis. an E. equisetina E pim Bot. Need. 33: "ST- 266. Annals of the Missouri Botanical Garden A developmental study of the integu- rms 3. E, in Спешт gnemon L. Bot. Mag. (Tokyo) 99: 241- 266. & P. ToMLINSON. 1989a. Aspects of cone and ovule ontogeny in Cryptomeria (Taxodiaceae). Amer. J. Bot. 76 692-705. 9b. Cone and ovule devel- opm pa 5 Callitris (Cupressaceae-Callitroideae). Bot. Gaz. Cone and ovule ontogeny in Taxodium an nd oa (Taxodiaceae Co. niferales). Amer. J. Bot. А ont 1221. and ovule ontogeny in o oP (Taxodizceae r Amer. J. Bot. 78: 41 2a. togeny in Metas uoia, uoia кз codd en- ee. (Taxodiaceae Coniferales). Bot. J. Linn. Soc. ers Per 092 чеч of cone mor- phology and — € in POMA paceae (Conif- erales). Int. e zl nt Sci. 153: "588. TAYLOR, T. N h of d erican species of roce a laeontographica, Abt. B, Paláoj hyto| e 04197 1: Synangiate Pennsylvanian pteridosperm phia or- gan. Amer. J. Bot. 58: 300-308. س‎ Poll || £f: 91 nosperms: Phylogeny and reproductive biology. ‚Рр. 177-217 in C. B. Beck (e ditor), Origin and Evolution of Gymnosperms. Columbia Univ. Press, New York. 1985. The Cretaceous preridosperms . uflorinia and ашна чапа ji mpli- tions on carpel 72: 1842-4 1853. . EGGERT. 1969. On the structure and relationships of a new Pennsylvanian species of the seed Pachytesta. wee 12: 382-3 . A. Muray. 1981. Morpholo оріс vari- ability о! an lyginopterid seed ferns. Rev Palaeobot. Pal 3224—62. ‘STEW 1964. The Paleozoic seed Mitr rosperm merican coal balls. e to- graphica, prea B. Pili 15:49) AYLOR. The Biology = Putin of Fossil Plants. шесе, Hall, Englew Cliff: ——À м. A. CICHAN & A. M. s 1984. The ruchus dubius ana noua. Rev. ре Palynol. 41: 319-327. Tuomas, H. H. 1915. On Williamsoniella, a new type Philos. Trans., Ser. B 207: 113- m The Flag uri a new group o ie terc plants from the Jur - — - York. оше, ae Trans., Ser. B 213 299-3 Оп ѕоте атон sei from D mesozoic ды of south Africa. Philos. Trans., Ser. B 222: ne 1958. : idget sopteris. Bull. Brit. ME (Ча, Hist ) em 3: 179- als TOMAR. P. T. Takaso & J. RATTENBURY. 1989, n PE (Podocar- paceae). ead 1 Ши га: "99: 2 perennes On Pter uchus a micr Озунон Bull. В Mus] (Na ы Pw чү A W. ROTHWELL. 1985. Morphol. f Pal, TRIVETT ogy, syst peur Masry priapi, sp. nov. (Cordaitales). Syst. Bot. 223. 988. Diversity among Palaeozoic Corda itales: The vascular чене cture of wu xter. 149: 116-125 k the es of the group. Pale 2: 75-84, Plates Е , J. №. & J. А. Dove. 1975. The Тон angiosperm ины» eren Ann. Misso Bot. Gard. 85 E IELAND, С. 906. puis Fossil Cycads. Car- negie Inst., Washing on. WILDE, M. 944. A new interpretation of coniferous cones I. Podocarpaceae (Podocarpus). Ann. Bot. (дос) 8: WILLIAMSON, ү Contributions toward the history of Zamia gigas Lindl. & Hutton. Trans. Linn. Soc. L 26: 663-674. RE T AYLO E tica. OFP in n P tionale de Palaeobotanique ТУШ eee aris. seda p. 178. ZALESSKY, M. D. 9. Observation sur quelques ve- ito ux fo: = nouveaux. Bull. Soc. Geol. France 39(3- 9 189- ZAVADA, ATE CREPET. 1985 e type material of pec Ж pillatus. Pollen & Spores ultrastructure of th s, P. dubius and P. ра 27. 1271-276 Hen 1986. len wall Caytonanthus arberi (Caytoniales). Pl. pn ch 153: 259-264. APPENDIX A. Character list and discussion tic 0. yo ж (0); herbaceous (1); pac aqua (2); a 2e & Previous investigators (e.g., onte dean d " we bahit (see char? 1990) have segregate: pas shrub an ner san single cti u shrub is at best arbitra polymorphic within angio sperm termina de se n" o include a wood wil of ve and pro; cribe bene te vascular cambium ре Е іп terms of uces secondary wood that can be c Volume 81, Number 3 1994 Nixon et al. 525 Seed Plant Phylogeny wood anatomical features, as discussed below under wood anatomy. This is also applicable to the supernumery cam- bia that occur in some cy сабз аз › well аз ш Gnetum. The ү M » he kno occ taxa in e analysis outside of the iu. T ally, because all known submerge: ШШ» ш. or Saye n Vi egeo (nonwoody), we ed). additive (order 1. AXILLARY Aie absent (0); simple (1); multiple (2); additiv gh coded in terms of the pres- hs ary buds, it is sion to the “branching dichotomous" versus “branching axillary” character of me e previo ous analyses. The character is encoded as ih | are ied It should be noted that a nonadditive coding does not й change our results. Dichotomous branching as u isotomous anı omous pat tna: In some angiosperms, both axillary and dichotomous these instances are found in n» we assume to be liy derived angiosperms such as and ). even have multiple buds in addition to dichotomous and rend pon н states (Troll, mado In 2 = of єк involv: 1 1 + All o e angiosperm terminis used in our iras See in ү fashion. Recently, it bv D. АҢ & Don oghue (1992) that dichotomous focis in extant cycads i y ri from axillary branchi n the e we find no evidence to support axillary branching as occurring in either fossil or extant cycads (Stevenson, 1988), we have ——— the position of Loconte and tevenson and scored all cycad taxa as lacking axillary buds (state 0). 2. SHORT SHOOTS: absent (0); present (1). e defined as gti shoots that differ in by suppression/reduction Short shoots are morphology fr from main shoo of internodes (as in Сіл ad 3. APICAL MERISTEM: w/o tunica (0); w/tunica (1). The cept of t orpus orga CREAN of shoot iis aeri és has "been discussed Бу Loconte & Ste- оп (1 1990): 1992), "The char- h ent. However, as discussed by Loconte & Stevenson (1990) we consider the corpus Li and the presence of a tunica to be a apomor фиг ` The plesiomorphie presence of t is not a ntic issue as pice by Doyle & D u (1992). "Ra I itisa necessary part of the stem apical growth in the seed plants, and its not only by histology but also by cytohistochemistry and cell division patterns in terms of maintaining the conical shape of m qe (Gifford & Corson, 1971; Steeves & Sussex, 1989). STELE: а (0); eustele (1); polystele (2); поп- additiv The on condition throughout extant seed р groupsas is = oustelar Hii Am sip the terminals esie upo! ES le etie morphology, of the lower oae Р apr rotostele, as found Sekido, 1975) and ШЕ axillary branching in Lygi- ene, (appar je Mayaca A ; EE NS hte and medullosans. In the case of bible. epp ue y pnr Aperte ы hee bie ei erue (Galtier & Holmes, di 3 inl y derived Hom. the 1982) is be mia to and bra n Lyginopteris ap- кс conditio f pears t u Some merr on on the pet etiole m кү ат branching ot a polystelic ae iar to hat fou in many m yê ositi osans a enson, 1990). We have also scored Pentox- In [x sa ss etn ү t E L ylon a having a ‘polystele n it differs from that han s ndocentrica, has been reported as having axillary e 6. (На атег & Rothwell, 1988). This points to o the genera, a and the problem of synthate taxa based on as- por iaa of relatedness. We have thus coded both о ullosan and ор rid terminals as — Pad iguous) for the character of axillary bran ме ding to Kim & Sekido (1 975), 3 inclusion -’ssoniocladus within the cycads is base *pidermal cell walls and t presence o n ili ме Е i th of these теа аге plesiomorphic within — based о veral recent —— as well a: tr m fore, the char e pre f the ma fo ssil ca по! exclusively nor coi isin ely in- (ба sections on fossil groups). E NODAL ANATOMY: draco unilacunar (0); 1-trace per bundl (1); т (2); 2-trace unilacunar (8); trilacunar (4% E Nodal a red as in erae & Stevenson (1990). This preis as scored in е & Donoghue d is a composite character of patio anatomy and bios isa. ;. This include ferns in an analysis limited to the of depart the number of a obi de dies i in the stem ould be Laeti true if one were to . This character should be tat cycadalean affinity. In fact, Nilssoniocla ne? has Doyle & Donoghue (1992) нда that cycads hav p E piss with either extant o т known e xtinct mber of traces derived {г solid arc of xylem. [^m 7» phology f Our vi dil is "The leaf trace e Nionoctadu are not known for extant cycads or any cycads is derived fr rom numerous small bun is no that share synapomorphies with the cycads. There t d appear as a solid arc after the ое: сус Я ranchin evi he earliest unequivocal secondary хуіе m bas obecured the integrity of the рг be 1 zamia chinensis Zhu & Du and Сеш 35). We have therefore 0 EM абы Gau & n mas from m: the Early indi hina (Gao & Thomas, 1989 ), w Moose are viral ао from he" Î genus Cycas. Beca scored all three oF d terminals as state e pud pattern of the petiole in Fed les em In contrast, the in the cortex of the reflects the аан of bund! Annals of the Missouri Botanical Garden Thus, the number of bundles departing from those of the агг апрешепі in the petiole. In fact, the number of Чеш їгасез does not even represent a minimum number e of anastomoses that = occur in the cortex. Also, t does natomy of de ераг artin match tha of Nie leaf е ае discussion under character 24—leaf trace xylem E 6. GIRDLING LEAF TRACES: absent (0); present (1). Girdling leaf traces (with multilacunar nodal — thus this ааа; isa ee, for the е Cycadaceae, Stangeriaceae, and Z 1990). 7. PRIMARY XYLEM: mesarch (0); endarch (1). ary xylem character (7, 8) states are describe As fossil taxa under those entries. All o f the extant taxa in this study have endarch primary p (character 7). 8. PRIMARY XYLEM PITS: scalariform (0); conifer type (1). This character is the same as used in Doyle d hi кыне Reel and various "previous analyses; d. der (1955) and Bierhorst (1971 9. WOOD: manoxylic (0); kgs (1). cally the s. onte & Stevenso; asso) and Doyle * Doogie de 92) Dat oxylic versus pyc r be S an iA re дун oft fossil taxa in the literature bel art & Rothwell, 1993; Taylor & Taylor, 1993, а pi Sais cited therein). 10. SECONDARY XYLEM PITS: scalariform (0); mixed (1); circular та additive. ‚ Secondary : 1. ran to ы The state “mixed” refers to i presence of individual Brie elements that have LA scalariform and c in the sabe ат (see Stewart & Rothwell, 1993; & Taylor, 1993; and dienen ci ч therein). Some p e ako di rived from standard wood references vdd as wn & Panshin (1934). 12. VESSELS: absent (0); present (1). In order to reduce bias due to preconceived notions i 1 about the evolution of р foration plates), we have pea the form of the е per- ee fro whether or not vel occ dp 15 ез tha Finde (foraminate) ves sels are ho mologo ‘ous; see i fone & Sattler (1982) for a дон ж. i ш of gnetopsid and angiosperm vessel elem: 13. VESSEL PERFORATIONS: foraminate (0); scalar- iform (1). See Muhammad P Sattler (1982) for a discussion of this character in Gnetum. The simpl ple perforations of some ngiosperms (e.g., Chry. lepis) were assumed to be de- ri kd and there s scal orm. 8 grouping болакай in our analyses, but would п an expanded analysis that included more angiosperms. 14. SIEVE TUBE PLASTIDS: s- ey (0); p-type E^ J 0 "un able to utilize subclasses of th are from Behnke (1988) and references aia therein. 15. COMPANION CELLS: absent (0); present (1). As defined ni шер тее & Stevenson (1990) and Doyle & Donoghue (e.g., 1992). 16. LIGNIN SUBUNITS: vanillin (0); syringal groups (1). vanillin versus syringal subuni ee pe 1985 Onl шы for ех from Logan & Thomas ti A at e reaction) requires intact wood. . CORTICAL SECRETORY STRUCIS: absent (0 cules Rowe have s cored this a an Lis tive character vie (Ty annia (25. qM & Taylor dp poly xed” conditi e from Bierhorst (1971), 5e A a Th 1 ddit it ity f the crc onsist ith b allowing (1998), а from unpublis lishe d p ions. " bere tet р р 18 vities are je secret are independent lacunae that ar е lin longi if “mixed” is scored as an dependen s state in a non- epithelial cells. In со ontrast, secretary € s ae aat additive character. Of course, the c ts circul ar ач scalariform pits suggests that, while tsm nes in 9 same character, they may be independent charact ter: sealariform d circular ine treating this character е have above is logically a to scoring two separate ary pie 11. XYLEM RAYS: uniseriate/bi tiseriate к i) (0); some mul- same as used by оти te & Ste fiver ue Se & dub (1992). a are derived Dat mostly from Greguss (1955) and dei адан of fossil taxa helal Jl. lacuna lined with both с the gen schizo-lysigenous in origin. Cor ied secre dion ie "sid ele armen from DE str deni in secondary xylem or rays. Tes m Ssa E > TE c ij i trea! canals are derived from cavi ot t (1) 18. ETHEREAL OIL CELLS: absent (0): TRE to an Ethereal oil cells (as far as known) ar! Муст are spe pepa ds cells that produce etherea' 0 ist (1981) cialized p sius cells. Da ta are from Cronq Volume 81, Number 3 1994 Nixon et al. Seed Plant Phylogeny Loconte & Stevenson (1991), and general taxonomic references. 19. RESINS: absent (0); present (1). Although resins are the ШЕ a secretory struc- tures, there is not a between type of secretory structure (e.g., canal or cavity) and the type of secreti ion e.g. resin or muc cilage). The presence of 3 t i} T and corystosperms. It should ree aa that while conifers have resin-secreting canals, the corystosperms have resin- secreting ре бузун with in character 18). In the same medullos: and oe inkgo, Calli. stophyton Lep oe ШЕ “Tatarina, and "m d Х аъ a } 1 1 1 ү j^ fossils). 20. NE IET alternate (0); opposite or whorled Phyllotaxy is e as either alternate or opposite/ whorled. As have previous workers e the two states n dip arate groups, as in the case of Ep hedra where di three leaves at node. Pseudowhorls or pseudo- те phyllotaxy ап аге Aces difficult to exem as is the case in Catalpa and Peperom и 1925). 0 the taxa тауы ur ost problematic is Pla. s, which we have ie un ical кш, but which may : оа derived independently through sympodial ran reduction of one leaf at each node to esca the! foliar ный that encircles the stem, much аз the leaf base encircles the axillary @ 21. LEAF BASE: simple (0); sheathing (1). Beca difficulty in E early developmental Stages i in n foi taxa, we have е our definition of sheathing” leaf oa to which express the condition in mature le Ms: tax This character is not strictly D = the xylem pote род in ste bs ndles as might be supposed. For the ave Arta stem үүн with каркар, leaf ie е не 1990). 24. LEAVES: simple (0); compound (1) le a few fossil taxa are difficult to score, this is a idi od uiae character 25. dio uer VEIN FORM: dichotomous (0); aniso- ous (1); solitary/unbranched (2); nonadditive. This character refers to the ШАГУ vein system only, and thus midvein aoe ous” in one taxon, hin midvein “not serine in ni such as Ginkgo with only one order of vein 26. LEAF aed parallel (0); pinnate (1); palmate (2); nonadditi This cha ч was spp only for taxa with more than one order. of vi ven ns (e.g., Ephedra). Тах such as conifers and сав Were coded as api дезе! the leat venation. 27 Mrd со опе (0); two (1); three or more (2); All taxa in our analyses have at least one order of venation. Two vein n orders | occur by branching off from ji ва. . Welwitschia and mono- red as addi ше нер of eaf has we or parallel primary v pound leaves Was pet to be “ргітагу” Һе codings of Zu leaf characters in Du & en hing 1986a, b, 1987). 28. LAMINAR VEIN FORM: dichotomous (0); aniso- р РЕ РЕР 3 (1) Laminar vein form refers to the iore of venation in to have sheathing 1 g leaves ане ane ontogeny ‘ach as in det sro Au (Stevenson, 1990), = ё leaf bases. 22. STIPULES: absent (0); present (1). ules are considered here to be the presence of аззо‹ may be 23. LEAF TRACE XYLEM: mesarch (0); endarch (1). nd * have introduced a new character dealing with vids versus даады (including ren leaf traces. (inapplicable) i in е taxa with only one order of v (e.g. se the form of the primary vein in к іп character 25. ЕЕ mp Е 8 © pon a system of anisotomous branching rather 3.1 fth + Р F gios} 29. VEIN FUSION: nonanastomosing (0); anastomosing P ). usions occur in those forms that are dichotomous in Ё ed adita (e. E cessions nee Gnet tum) as well as eter anas ing f in angio sperms. a = e taxa, а: уртад s occur гоу пеаг фе leaves (e.g., Ginkgo) x: do not margins о such cases, s the taxa sistently ooo the lamina. In e not consistent throughout t the lamina or from leaf to leaf. The character therefore is defined as anastomoses nime as distributed over the leaf blade. Annals of the Missouri Botanical Garden 30. CHLORANTHOID TEETH: absent (0); present ous within-a Mrd Dove, 1999 "We = © made as ааа states within the “hamamelid” termin 31. GUARD CELL POLES: raised (0); level (1). This sangis, AM character 32 have been used rou tinely in seed nalyses (e. E^ Crane 1985; Doyle & Donoghue, foe 7, 1992: 1 , 0 note that it is diel to determine with certainty in e fossil mat S2 ti ere haplocheilic (0); some or all syndeto- ilic (1). See wb & Donoghue (e.g., 1987). 33. ASTROSCLEREIDS IN LEAF: absent (0); present (1 х ты g of this character follows that of ee & aoe n TE 34. STROBILI: tics A bisexual (1); functionally unisexual (2); additiv This character refers to the sex of whole strobilar axes. By: definition, all dioeciou ious plants are unisexual, n жеше of some taxa is of necessity та on the inter- tation of “str obilus ” but ie di fficulty, ttm bilus. Under E^ Ше micros sporoph ylls and a sterile ovule, as in Welwit. bisexual axes, we have encoded this те hi as additive. 35. PERIANTH: absent (0); present (1). s here operationally defined as a set (whorl + pare of sterile Apc subtending carpels and/or of uncertain homology we have not scored the gnetopsids or c ps cadeoids as possessing a eclar s of the “strobili” i positional are unknow. wn; characters such as 8 by their fere relation t to ( | iknown ( d omology in some parts relations of parts anth" are homol carpels, which the angiosperms. 36. RAPER pinnate (0); reted angiospermous sta: Ospo- ТӨЛӨШ 5; qas is a long history of other interpretation (e.g., eer e T с Fd structures to which microsporangia (or Уйа) аге аї- tached. Our аг, интер тос have in general followed tra- i those cases whe: е o polen: characters are s "ton on PENG osans). upon the extensive studies of Martens (1971, and йене cited therein), Ephedr a e таса y! i edul- cored as ambi guous (set abo es vi innate microsporop instead of imple microsporophylls as they have scored in some revio lyses. In Welwitschia, the six "simple" crosporophylls (as interpreted by Doyl 2 9 and tw caller lat eral арра mia in early чч 0 1971) these can be c seen as a opposite three- lobed u further evidence that these are indee their positi Н то subtending bracts, ar on relativ de min if interpreted as oppos e three-lobed pes tures, the micro: peropbyl continue pa ectly the the decus- sate taxis of zn pei ; this ta xis then inues above jas two sterile bracts envel he sterile n ven eie ). The cle t easily be interpreted as having (in some p lobed pinnate microsporo ophylls as opposed E phe Ponal l interpretation that thes e are thre ple ge th рен sed, and the al ей ones аге re orn size (s rendeen et of 1993 for a discussion of the various inter ae Chloranthus stamens and their impli tions) d gun ranthoid sta are int te comp: dol structures, the reduced lateral “glands” found at ip of many L РИ? stam: ight also be quo reduced lateral pin chloranthoi pei ШЫЙ sporophyll. We pres а encoded апу such interp ner ens perms the e суеш of ace structures in ап B / 37. MICROSPOROPHYLLS: spiral (0); whorled/0PP site T. ). This is a straightforward mi А оло k pising fossil prim such as the gloss жат ( Бү, parts or difficulty їп determinin 0 ps! 1987, noted that іп Doyle re Donoghue (doe insi the single angiosperm terminal w аз v ving spi! Volume 81, Number 3 1994 Nixon et al. Seed Plant Phylogeny microsporophylls; in our analysis, by scoring numerous angiosperm taxa as observed, we allow € to select the likely ancestral state for angiosper 38. MICROSPOROPHYLLS: free (0); ку fused (1). This is a difficult character to score within angiosperms in some cases, particularly when fusion of; parts such as Bie ST and carpels may envelop the basal portion of 39. MICROSPORANGIA PER UNIT: many (0); 1-4 (1). This character refers to the number of ара Pd ultimate appendage, or sporangia-bearing “unit.” p" ns that t taxa s such as medullosans can be s number. 40. MICROSPORANGIA: terminal (i marginal (1); ad- axial (2); abaxial (3); nonadditiv: This character is similar in definition and scoring to ШЕ sed i in previous seed dens analyses; we note some form is fil r development of a connective extension € our rig few such taxa were included, but with expanded analyses, this may pose à problem). The “peltate” theory the origi the angiospe: see Eames, 1961) is also germane to this character, for, if such an interpretation is a pted, then all angiosperm stamens are basically ШШ (аз їп gneto sids) and dified t mar- ginal, abaxial, etc. Mis sta ne d on n issue, an acter based on obse: models of КО pat rvation, not 41. о овАНОА, free (0); fused at least basally $ This character is defined in the same manner as pre vious analyses; we find it difficult to score with certainty їп numerous taxa, and our matrix reflects this. 42, coe Bees a _DEHISCENCE: ectokinetic 1(9) endokinetic micro- ffrey & The concept of ec eee versus sporangial dehiscence w. 1916 Teste 6). The ectokinetic mechanism is e om epidermal cells, ds, whereas the endoki- clay "men. is derive d from two or more sul bepidermal asin Gi шо ү, Р Я ntroduced this character here and rt er sate, csc which i isa bE at endokinetie es mal that layer. pos is based u upon observations ee ant and Em taxa (Stevenson, unpublished). Further nd are from ingh (1978) fo many conifers and Martens (1971) for many gnet opsi 43. — (POLLEN) SYMMETRY: radial (0); ateral (1). T Pollen symmet ry has been used previously; it is Moe у hr although in some instances the collapse of globose radial grains could produce a баа. air pue Thus, all scores are based upon uncol- lapsed gr: 44. MICROSPORE TETRAD SCAR: present (0); absent (1). Tetrad scars (character 47) are found only in fossil а. 1. bg s ү ү? ٤ Td ^ i taxa ш OUI J өе entries. It = — e to no e both a proximal tetrad scar and a dist Hy үрп а distinction between characters of the aperture and the tetrad sca 45. MICROSPORE APERTURES: inaperturate (0); po- lar oisi ni ate (1) equatorial multiaperturate (2); n This character has been used in some sip in previous — by oring several pu we have not esumed their State 2, “е uatorial mul- aper urate," encompasses mostly triaperturate (tricol- pat E md taxa. Within angiosperms А Һаг- А, 46. LEPTOMATE APERTURE: absent (0); present (1). “Jeptoma” is a distal oe mm germination pore (thin area in the wall) found in Cordaitalean and extant conifer po ollen: n; it is more radi ally symmet trical i in pollen. 47. COLPUS эр colpate (0); colporate (1); pororate Oy additiv: character is = as additive based on the M er m (€ inaperturate pollen is scored a oodd шйышбей Nolin of the ба multiaper- turate charac te (0); saccate (1). 48. POLLEN SACCUS This character has been used in all previous -a ү! analyses їп a similar manner. See also character ate (1 ) 4 CACCTIS FORM (0) . АССИ q 986) bos a бона infille d PM n & Friis (1 endoreicula) saccus interior, or of Caytonanthus; ы е г ; ЕЕ cian 993). This m cate dpi or transformations within hom pollen types. iologous saccate 50. EKTEXINE STRUCTURE: alveolar or ramifying (0); granul lumellate (1) Some variant of this character has been widely used Arenas BE ARA E g E М and various © “solid” types a si hig ома literature, we slave chose n to lum] structure for all of those put tatively * ‘primitive’ pem in in cycadeoids, gnetopsids, and angiosperms. Annals of the Missouri Botanical Garden 51. ENDEXINE STRUCTURE: lamellate (0); granular (1). Previously d our coding fol lows those. We hou point out, ретте that while al ranular endexine, lay- ered endexine is known especially in the zone of ce and pe rhaps this segs cter needs to be reevaluated t take this into acco EU 52. TECTUM: absent (0); clearly defined (1). As we define the tectum, it is a clearly consolidated outer layer of the ektexine; thus, we do not consider the loosely associated outer granular layer of some conifer- ophyte pollen to be tectate. 98: Канан STRUCTURE: columellate (0); granular solid (1). or those taxa that have tectate pollen, we have guts Pu the tectum is underlain Бу columellae or gra g wall as columellate. we scored t 54. TECTUM FORM: continuous or foveolate (0); semi- tectate or reticulate (1). Scored only for tectate d Terminology follows that of M & Doyle (1975). 55. PARALLEL M STRIATION: absent (0); global (1); proximal (2); nonadditive. ‚ Gross similarity in exine morphology has been noted 1 Welwitschia; the * ‘parallel” exine striations of thes a, however differ in their distribution and orientation (gay dis- йлы залаат Various striate pollen types of some angio- types that are sis supratectal in nature, al- үл gh some axa of striate Bo copiis (e.g., certain 56. VM eye rod PHYTE: more than 4-nucleate (0); 4-nucleate (1); 3-nucleate Gk additive This character has been widely used in seed plant nalyses; we encode it as additive e under the assumption ibat reduced 4- and 3-nucleate types are part re- re o duction series; other a arguments for — coding can be made on the basis of meristic log 57. POLLEN TUBE: suspended (0); penetrating (1). The differences and significance of pe T versus suspended ee tubes are discussed in Loconte & Ste 990 only known for extant taxa, and t us complete corre- lation is not known. The е presence or absence of flagellate male of penetrating Ac tubes is not necessarily dependent upon the latter just as ramiform pollen tubes are not dependent upon gamete motility. 5 ا 58. RAMIFORM POLLEN TUBES: ab (0) (1) amiform дей tubes occur in cycads, Ginkgo, P. conifers but a not known m gne Mui With the e i ollen uo not known in fossil seed eee Within angiosperms, ith chal- appear my in some паа. groups (EARN 1961); ome these data are spotty and inconsisten 59. STALK CELL: di Me absent he tall heridial Meus in the male hdi ا‎ ps ei s eec out by Singh, this cell should probably be called » d prothallial cell," but histo: ove Lm is re tained here.. As talk с ell is fou nd in 1 the т mi- 0- 978). ed this all fossils, altho un hé form fthe acy serait fossil Callistophytaceae (Millay к nae 1974) indicates that it is probably present in PRPS 60. SPERM: flagellate (0); nonflagellate (Dj onflagellate, and their nonmotile refereed to simply as “male cells” t = peg im a Mee prevents scoring this character REN 61. SIMPLE MEGASTROBILUS: absent (0); present (1) 62. a Aia MEGASTROBILUS: absent (0% pres- nt (1). cters 61 and 62 are logically connected since up o зе le all совр. нео by — are made н Гр strobili; by separating the components, we W tonia iig ious niyo with its alternat ad production of — eg ‘yeas, ы structures with ve, ive leaves on the sam огей as t not to have a simple megastrobilus s ear beating structures are produced on wih ome es indetermina m stems м а таппег similar 3l t In е taxa that have a simple megastro m аску it is sae clear a a ыз т ound obilus ыз dl and such taxa have been d as ambi Bo OBA, 1985), of Eames (1952), Hass (1971), Takaso ( nstrates the «ы Takaso & Bouman (1986) clearly demons sence of a compoun d strobilus in um oat à P at P" The * *compound" nature of the microstroblu Volume 81, Number 3 1994 Nixon et al. Seed Plant Phylogeny аротогр phy top 3 ght ue of the ер е- topsid с clade. We have not encoded a character for simple versus compound epit s beso ed дов оп anonymous reviewer). С килче ип tation of ^ ACER floral h score angiosper: “mires ously co: «ы mple; moreover this charac no о elfe сї on our results, because, if the angio: they should be), it is interpreted mpo e, 1944). Aiton, mum OR OME i taxa i in our analy would for 63. WOODY CONES: absent (0); present (1). The рге absen РОК cones is limited to three Setter families Sud. Cordai tales 64. xc NM CONE UNITS: many (0), few (1). aracter reflects the degree of bra This nching in com- каш о cones and is о of value only within the conifer clade. 65. CARPEL: absent (0); present a The arpel is used he & fem d ў angi gios ү n the classical sense ing h h pollen раа this eliminates me Strobes аз the seed envelope of Cay (toni а, since nee ne d EEDS: absent (0); radiospermic (1); platyspermic =e nonadditive. This character i is , commonly and "et nes in seed pla t anal p z note that it is eine: Loy i xui rai larly som diff f opinion al : ог major ө groups (e.g., whether the angiosperms have radiospermic or со oncept ofr di platyspermic seeds). In fact, the trt et ш well established or clea r example, * *platys permy" can arr : d diss of vascular bundles, "n early stages of i 6 ent of ovules, w. ow. netopsids are scored a ic because the х gh the angiosperms in our analyses a Yspermic, it is not clear whether the — isa pla plicable eiie dinis cie studies across the spectrum of angiosper 67. aod gk алаш (0); foliar (1); peltate/ nclosed (2); ni , The angiosperms are scored as nice for this Capa may be simple | b may be compound structures that ‘include a leaf эй logue that subtends an axillary structure еш side nteresting to note that both groups commonly. refer eh to as sister taxa or outgroups of angiosperms (eycadeoide and gn e- foliar ш менш bear subtending eects, dod di 1 РА th Thus xig origin up the pen bun a leaf homologue extending all the way to the coniferophytes and Ginkgo. The ре ү enclosed ovule position is found i have peltate or Шан whic! pel ete structures CER sometimes enfold the ovules. These tui e difficult to меа as to atte yey these as panone? us with angiosperm carpels for reasons outlined above 68. OVULE NUMBER: 1-2 (0); many (1). This character has been widely used in previous anal- yses in similar forms. Undoubtedly, the problem of foliar are homologous over all seed plants. 69. OVULES: orthotropous (0); anatropous (1). This cha r has been used previously, but in our e has ‘been improperly bores on the basis of phy- in eal taxa. a base our Тейиш of orthotropous and natropou the position talk) of in ханды gents without assuming that sine roups ot group itary P uc The two states are thus a s straight phe in н duit is recurved in Caytonia (see discussion in ыга section). 70. MICROPYLE: normal (0); tubular (1). The “normal” micropyle in this case refers to the case Lia of : 3 H 1, А int toh This character bears further investigation within angio- sperms, some of which have extended inner integuments n ht t 71. MICROPYLE ORIENTATION: distal (0); proximal (1). This — refers to the direction in which the ovule int relative to ud structure that bears the oruliferous structure (e.g of the strobilus). It is most useful for differentiating od and agr r repro- ductive st 1988; Stevenson, 1990). Annals of the Missouri Botanical Garden 72. OVULE GROWTH: pachychalazal (0); endochalazal (1). Two contrasting states of ovule growth, pachychalazal and endochalazal, were ssi by Takaso & Bo (19 86). Because this character ‚ it is only applicable a extant groups . The character eg those ovules that have a massive chalaza with їп angie Ey generally attached above the middle of “the ovule. . This is br rought part of the seed ce the ааа end in ra 986). In contrast, gnetopsids oe (the area above the eg Sp attachment) and are termed endochalazal a Takaso —— i m seem: at of a misnomer We hav: velo E a character for various layers aye or Y appendages that envelop the ovule/seed i " phylogenetic hypotheses i in in- ee char defi velope as a laye: dual sé in contact with din of that structure does not evan to ithe micropylar region and is шк]. even after t studies! (Tal 1980). Although the dual Yasculature of the cycad ovule has been in as indicatin not present va entally or in mature ovules, we I. scored it a 74. OUTER SEED ENVELOPE: simple (0); bi-parted (1). Character 76 (outer seed envelope) treats the outer seed envelope r simple or bifid. The bifid condit n of the gnetopsids i is the result of two opposite bracteoles that are fused, as demonstrated ontogenetically by nanter E Takaso (1984, 1985), and Takaso & Boum n (1986). 75. MEGASPORANGIUM: w/lagenostome (0); w/pollen chamber (1); angiospermous (2); nonadditive. This character deals with the structure of th ima i region n) of i I e, and whether the Aeg gig Th 76. INNER INTEGUMENT: not-bifid (0); bifid (1). This character has not been sed in previous seed plant analyses, but was ашу Шз їп numerous extant taxa by various workers. The bifid PAM s p integu- mature ovules of c fossil taxa i aa a in most 1 rok viden bifid in P^ vum inco rn 1871; pec 1984. тз so & Bouman 1986) ог angiosperms (Bouman, 1984). 77. CONE SCALE SEED WING: absent (0); present (1). This character is included to encourage resolution with- in the conifers. 78. SEED COAT: xus (0); sclerotesta/sarcotesta (1). The nature of t ure integument has been used n previous analyses i in pr form. The simple s sores ды! a sclerified lay er fleshy э sclerified layers. 79. INTEGUMENT VASCULATURE: single (0); double (1). This e refers to deo) dual ber system found in the seeds of medullosans and cycads (Stopes, i: integum 80. SEED CORONULA: absent (0); present (1) The seed coronula refers to the specialized MS тиса of the a coat at the micro cea is res чий to the cycad fa mia Me Grako 1990). mun 81. NUCELLAR CUTICLE: thin (0); thick (1). the states Routinely used in analyses of seed plants and s e are given in most morphology and paleobotany tex . MEGASPORE TETRAD: бен (0); linear (1) isobilateral (2); nonadditiv = This character is routinely used in analyses ф ps plants, and SE states Еда tas are given in mos phology and paleobotany te у 1). 83. MEGASPORE WALL: thick (0); thin/absent (1) Widely used in previous seed plant analyses. А r- 84. MEGAGAMETOPHYTE: monosporic (0); tetraspo ic (1). taxa idely spa in seed plant analyses. Most fossil are problemat : eolar 85. MEGAGAMETOPHYTE: alveolar (0); nonalv (1). 1 di d plant analyses. Da Not Wig aber A use side ve 5 : me re ingh (19 т уе, for some fossil taxa, dns << examination of additional published figures. ennt ^ the тагтны. pattern of the alveolar Mei Volume 81, Number 3 1994 Nixon et al. 533 deed «чм Phylogeny 2. in many fossils. 86. ARCHEGONIA: present (0); absent (1). Widely used i in seed plant ponies. Fou in many atio: fossils due f preserv 87. EGG: cellular (0); free-nuclear (1). Not previously used in m plant Seinen This ch acter is observable in mature seeds, and not lim = " extant t i ossil taxa where ссни have been observed (e.g., UM Cordaites). 88. 2 EMBRYOGENY: free-nuclear (0); cellular Data from Singh (1978) and Bierhorst AY Unfor- nu this character can onl red for extant ха beca vm early embryo stages Say are not pre- Biss fossils. 89. EMBRYO MATURITY: postshed (0); preshed (1). viously used in seed дин атат, The states S versus pres| : the timing of embryo b hat embry: y analo logy with living taxa such as repeal and Ginkgo. 90. EMBRYO FEEDER: absent (0); present (1). This tg is scored only for extant taxa and is char known n only in topsi sids. 91. SEED GERMIN on а » ATION: cryptocotylar (0); phaner- ® Н ореа ao This chara ssil t cter has been scored as ambiguo 92. FLORAL CUP: absent (0); present (1). GR те сна asa concave, usually basil _ structure erianth) and b: Каны : aminodes, and possibly db, is found in hd Ta параш taxa such as Calycanthus, various poh a d Eu mate: and may di e much wider o Aus in modifie. cd fo m than is posa ‘oe Й E^ ао » in Laurac ae). We dist uish betw, ween su d S yx ary. We have avoided use of os ypant ium" b бс "une p applied to structures formed mostly of stem *ptacle or mostly of calyx and other floral appendages. 93. CALYX CUP: absent (0); present (1). See previous character (92) for discussion. 94. PERIANTH: indeterminant/spiral (0); determinant/ cyclical (1). This character is scored as ambiguous for those taxa without perianth. 95. TRIMEROUS ANDROECIUM: absent (0); present К ). character is coded as present when one or more es “of the androecium are in threes or even multiples ther 96. STAMEN FORM: filamentous (0); laminar (1). This character is the traditional distinction betwee: бей, үс stamens and thinner, terete Шал» 97. STAMEN CONNECTIVE: absent (0); well developed (1). The occurrence of a well- developed stamen connective, is typical of many he taxa to be considered basal in the angiosperms oe 1981 dem 98. CARPEL ARRANGEMENT: spiral (0); whorled/op- osite (1). Flowers with single carpels have been coded as inap- plicable for this character. Data from standard taxonomic ANA ^h, 4 ы fi 99. CARPEL NUMBER: 1 (0); 2 (1); 3 (2); 4/5 (3); nonadditive. Scored only for those taxa sae vitalibus 100). ith ctahilized mb of 100. CARPEL NUMBER: indeterminant (0); determi- nant (1). Indeterminant carpel refers to spirals or whorls that fixed on particular modal numbers. al 101. OVARY: pad pm ОУ вупсагроиз (1); вупсаг- pus -inferior (2); a har xoa is treate ted as addit үнс {еру а sense use here). Taxa such as thus have an apocarpou gynoecium but г are ein and are scored as ае ovary. In all taxa a ылы 1 cup, n our analyses Ae! as. syncarpous-in inferior, the calyx ку от = appendages are clearly adnate to a syncarpous o 102. Бәнем — laminar (0); apical (1); sal (2); nonadditi haracter is not related to number of ovules; igne apic and or marginal, but in the EM merous ovules were always s ot be the case i more angiosperm taxa were E Thi: inclu FUNCTIONAL CONSTRAINTS AND rbcL EVIDENCE FOR LAND PLANT PHYLOGENY! Ре ictor 4 Albert,?* мий Вани Kare . Cha rex ; Manari; 5 ‘Bren D. Mishler,® and jose C. Nix ABSTRACT though the proportion of “functional” DNA It A in euk: most Sequences g; aryotic genomes is both de ii ae and jor to definition, ipl that ous or functionally labile )F tide- approximately 97. 5% of codon "change on dna branches are function; ally neutral rbe 3 athered for phylo ogenetic purposes are geavi functional. For example, patterns of variation are likely to be , strongly constr шр im ribosoma al RNAs beca of the eir structural por d roles in gs otein translation I gg I " M are usually es Weh d tl f fi traint: plant rbcL sequence ot only do rbcL sequences appear } lly clocklil but nucleo! (i.e., L evolution appe ars to be str ongly constrai ined Liy uncti g to highlight “blocks” ‘of conservative information that m m t be ey constrained. m cladistic analysis of all available or will highlight dd Золи п of congruent information, фер" dn functional constraints g the characters analyzed. We demonstrate the strengt th of this forms of the same rbcL evidence (i.e., чету strings, or amino acids) іп combination RAT the seed- dia. ye of Nixo 1 Diversification of the major reo of extant land pans probably dates from the Silurian to Creta- . During the Sil Devonian stimation of land- plant phylogeny, a research goal spa g over ion years of cladogenesis and extinction, а по — task. For example, many groups lack оо patterns of relati ionshi Recent years have seen an Метр. of i "P pois in seme infor: жыш interpreted similarities for asi ly 1 агре ^ 54 near n] 1 We th = Steve M and Peter Engstróm for ideas, and Mick Richardson f ents on an earlier vhs Stevenson for per mission to use u i ghi ings was written esearch Cou (1 Ng she E BSR- 8906126 to JRM travel arrangem (BSR- 8906496 to MWC, thanks the symp phenotypic gaps. In particular, the plastid rbcL gene (which encodes the large subunit of RuBisCO: ribulose-1 Сна. carboxylase/ rd rima in carbon fixati ө as been sequenced ex ue i prim phasis on ary e the deseen :(Clegg, 1993; Chase et e: 1993). Arguing fi ubstitutions optimized on cladograms; see Albert 1993; Albert & Mishler, 1992 Albert et al., 1993) ы ere omb, Brune! T Marie Friis equ with en ^ advice, and discu ussion. по Lipse Ко and the 1 Тай, VAA f the Missouri Bota nical Garden for their Үт and реет a assistance. 2 Dep с Bo tany, ium University, Monty's 6, S- 752 36 Uppsala, Sweden. 6, $-752 36 rsa nde * Laboratory of Mild: Systematics, Kingd om * Depa sity "ur Jepson Herbaria and Department * Uni California 9472 MESA TE H. rtment of Biology, Texas A&M University, College Station, Texas 77843, Villavä; agen J United Royal Botanical Gardens, Kew, Richmond, Surrey TW9 3AB, U.S.A der of Integrative Biology, University: of California, Berk ailey Weta: Cornell University, Ithaca, New York 14853, U.S.A. ANN. MISSOURI Bor. Garp. 81: 534-567. 1994. Volume 81, Number 3 1994 Albert et al. Functional Constraints and rbcL Evidence we will demonstrate that divergence-time asym- : Geixé 077 +З 11 1 al. (1992) and in ш lati te t f Gaut et al. (1992), but i d diff of land-plant phylogeny far more than do rate asymmetries. We have examined the internal stability of land- plant rbcL evidence through conversion of nucle- otide information into different data forms, includ- ing presence/ Чан of ad hoc eae string Cladogr: oduced from nucleotide, string, and translated a ata a artially con ring trees reveals extensive functional con- каро through t the теч ы of silen 3 replacements. о rbcL eee are no les: functionally constrained than hological д acters (conira Olmstead, 1989; peus et al., 1991; e88, the separation of protein- -functional‏ ر from freiogenetic hip: may not be entirely pos-‏ sible. i reflects‏ phylogeny might be assessed through congr uence‏ studies with characters expected to carry diverse‏ үө at аснова! лш dus ch, we have‏ plant cim ‚ as a "constant," a new meni, ici Ru phological deis (cie et pe: 1994, this ue). 1 ae 1, nucle- ote, amino a nee r string data with t s highly qoe cladistic vb зе studies point to (i ommonality of in- 1991; Donoghue & S son, 1992: Jones et xen 1993; Mishler, 1994). THE RATE **PRoBLEM" As has been pointed out in several recent papers, " the rbcL gensis is not strictly clocklike (Albert et al., 1992a ; Bousquet et a 1992; m et al., 1992; : Clegg, 1993). Here, we provide number of ыны дв پو‎ 1) ristic saisi ae woody taxon r million years; Wen el & Albert (1992) estimated 5-7 x с s ее herbaceous-pair comparisons. Lineage-spe- cific rate differences were found by “чйр аё: et унны чыте, from our own findings. Thus, whereas rbcL data cannot be Дүрт perfectly ultra- ш (ie. ВА ig fying a clock аллар) the et of the clock hypothesis sil a For bet and o ition, apparentl *quasi-ultra- nearly linear. We term this con characterizing rbcL sequence data, * metric.” sry оа дЫ "9 several important im- клы. One is that the extent of sequence di- vergence in a given taxon ae g should roughly е underlyin, metric sequences Ate 1552 ef. Ruins & Sanderson, 1992- 347-349). Oivan that A, T, G, and С are the only character-state alternatives, either scenario is likely to prod atterns of simila at be no ologous and there- fore cladograms that are ahist is ) and oth though Ean r uence change are often invoked to explain branch attraction be havior (see Clegg and Zurawski, 1992: 1 ith ference to rbcL), the problem is er defined in terms о rate and divergence time as their uct, per-character change: the А of A e al. (1992a, 93; Al & Mishler. solute and relative baie of у An additional ination “of аа. -ultrametricity is the near satisfaction hof wine tive pone MY A molecular clock ie огу ot moiecut fixation are the expectation (see Kimura, 1983; Annals of the Missouri Botanical Garden TABLE 1. “Phylogenetic” estimation of total substitution rate for 19 woody-taxon pairs. The rate of sequence divergence buy + calsuleted as 8 per-eite divergence ce (the реті distance, D,, divided by the number of nucleotides (АП a). Average rates for individual ine us half of the veluies shown. Data are from Search П of Chase et al. (1993); Sig estes error associa J exported 19 Ep all C WT ines pe p logi d mpari , which follow "асц the arguments of Wilson et al., 1990). Divergence Divergont rate time а t. /sit Taxon pair Area assumption D, n pair ait) Callitris rhomboidea R. Br. ex Rich. Australia 100 My: 55 3.85 x 107" ааг йена ра Marsh Africa (Cupressa "espe glyptostroboides Hu & Asia 40 My" 16 2.607055 Sequoiadendron giganteum (Lindl.) N. America . Buchholz (Taxodiaceae) kr Illicium parviflorum Michx. ex Vent N. America/Asia 200 My: 54 1.89 40 Austrobaileya scandens C. T. White Australia (Illiciaceae/ Austrobaileyaceae) )rimys winteri J. R. & G. Forst. S. America 100 My 21 1.47 x 10°" Belliolum sp. New Caledonia (Winteraceae) Drimys winteri J. R. & G. Forst. S. America 100 My 14 0.98 x 107" es insipida DC, Tasmania (Winteraceae) ee winteriana (L.) Gaertn. N. America 200 My 78 2.13.1088 Belliolum s New Caledonia (Canellaceae/ Winker bc eae) Canella winteriana (L.) Gaertn. N. Americ: 200 My 67 2.35 x 10°" Tasmannia insipida DC. Tasmania (Canellaceae/ Winteraceae) Liriodendron tulipifera L. N. America 40 My 10 1.75 x 1077. Liriodendron chinense (Hemsl.) Sarg. ^ Asia (Magnoliaceae) Calvan chinensis Cheng & Asia/N. America 200 My 28 0.98 x 107" . Chang сы australiense (Diels) Australia S. lake (( as o СЫ) Chimonanthus praecox (L.) Link Asia 200 My 24 0.84 x 107" Idiospermum australiense (Diels) Australia s T. Blake (Сау h Ta: -i لے اپ‎ Cal -0 (til smod costaricana Онч Americas 60 Му" 15 1.75 х 10 Drymophloeus subdistic S. Pacific (Н. E. Moore) Н. E. edt (Агесасе " Chama je ›геа фе апа Oerst. Americas 60 My 20 2.33 x 107" Мура fruticans Wur S. Pacific/ India (Arecaceae) G Serenoa repens (Bartram) Small Americas 60 My 18 2.10 x 10 Drymophloeus subdistichus 5. Pacific (H. E. Moore) H. E. Moore (Arecaceae) Volume 81, Number 3 Albert et al. 1994 Functional Constraints and rbcL Evidence TaBLE 1. Continued. Divergence Divergence rate time (subst. /site- Taxon pair Area assumption D, taxon pair) ) Small Americas 60 My 23 2.68 x 10-" Nypa tun V Wurb. S. Pacific/ India (Arecac Betula nigra Hemisphere 200 My 35 1.2805 10770 а asi ШАД litorea L. Australia thofagus dombeyi (Mirb.) O S. America 100 My 30 2.10:x:1071* ОЗ У ни (Baill.) Me New Caledonia (Nothofag. Galphimia gracilis Bartl. S.-N. America 100 My 34 2.38 x 10-" der (сири natalitius A. Juss. Africa/ Madagascar / (Malpighiac New Caledonia Dicella est ra Chodat S. America 100 My 33 Өх Acridocar ie natalitius A. Juss. Africa/ Madagascar / Pow New Caledonia Mascagnia stannea (Griseb.) Nied. S.-N. America 100 My 34 2.38 x 10-" rido! pe natalitius A. Jus Africa/ Madagascar / (Malpighiace New Caledonia 33: 3.01 x 10-" Mean 2.05 x 10>" S.D. 2-0,75 x 10-1" * Standard time figure used to represent the breakup of Gondwana (rounded to the nearest 100 My (million years) па 13907 9 estimated using Terra Mobilis® 2.1 by С. К. Denham and С. К. Scotese; see Wendel & Albert, a time figure (ca. early Oligocene) used t t disruption of the boreotropical interchange between North America and Eurasia е Lavin & Luckow, 1993). ee ha figure use e Northern and Southern * Pac € (roun E to the ret 100 My us 160 Tog as estimated using Terra Mobilis? 2.1 by C. R. Denham ps ink otese; see Wen “eigen , 1992: 137). Dive iiid wel by = Ison a: (1990), based on the fossil re * Northe E Ms are eds interpreted as representing ss expansion from South America. Hemispheres upon the breakup UNIT CHARACTERS AND FUNCTIONAL CONSTRAINTS do Quasi-ult d imply se- pices coefficients very close to > neutrality. M phe; As recently reviewed by Clegg (1 993), a number neutrality is the neutral effect of point free of E and evolutionary studies have relied pred clocklike sequence TA should involve sole beL ғ sequence Мы en dnm analyses Pes сурого: of suc h changes, fixed as effec. } L nucleotides че substitutions would аге tape and potently informative ~~ be expected to be italy’ pene (i.e., TEN o Lee LEE n amino ras СК" һ regard to рде to total rates of change, if all EE ra uten conservative events under consideration are Fouad recent, (labile), ree ehr in pm nucleotide se- parsimony UR ay ээ to proce ences is thus an expected m a wt of strong a reduced probability of spurious branch at- Constraints on protein function.’ raction because of the وید‎ fes expected " See Clegg (1993) rbcL; note that only total sul t elevant to cladistic methods becaus Vox de ad. ` Assuming that нты сеа oft tions deleterious to protein function and that fis the fraction ¢ such r mutations, the dod, theory may in je ae as Where S is the total substitution rate p О, te (after Nei, 1987: 52, 411). Annals of th d bonia Garden b — 400 VU Lig Fic Patterns of ne versus spurious similarity resulting from ER ° 10°Yrs a ally ancient and з. In (1), all nodes IGURES 1 AND 2. irae d time-samples. In both cases, ially time- concidit at 400 "My, “true tree” appears lee sae In. (2), t he c E dogenetic E e patt dn MT "E Possible "E of nucleotide acd are indicate cladogenetic the mmetrically with DU o ше, ranging » бе 400 to d by th nuc le otide substitutions. In (1 iL In! 1 tory, whereas the latter represent spurious т, er- iis pisce тез e.g., from ) t these patterns of solis. are ыру q h other multi tely equal in extent (because of nearly er some proportion of ahistorical evidence or even её, Isoetes cles = Sees Eq uisetum, ies of spor ight be the expectation if ы = thr: en E were, st асн com n Ко), whic oximates t ns of convergent similarity eid seed plants, the libens. ms of (1) are LE st oe ж а will result їп most the sly. As divergen s shallower, pe reduced simoni reconstructions that ра ir "uei taxa кашы, e beco: kelihood of will in and E are paired чйр ча Although C is linked with A E) We bos "ало, this ee may uds з false s mae gi, ig tween B a well as kem (C, D ary, comparing ps dn jud d lineages pue st pai of Bes M that mim a hopeless dy even mixtu ure of historically re liable and monreliale mier. Likewise, MUR near the tips. This condition may characterize the rbcL-ba sae EAs and relatively E associ- aed р ‘“‘time-sampling” strategy has been ани сач їп crcumseribed ара заз е from particular an- Kron & as a whole (Chase et al., 1993 Я бан a “tim he п the ter- migm My i as s ene, a time sam- pling is 1 and relative tim- ings of mdr us events in a d atrix. Of course, the nodes of a cl estimated by external criteria (e.g., the fossil record; cf. Nore & Novac 2p Initial EE to analy angiosperms and other seed sme rbcL sequences fro (ie. [n E 99]; 1909. y& Тїш 1992). peii байы. the base sed results shown in this paper. based however, are in conflict with cladistic studies ede | below). Ribosomal RNAs, wit pr re are obvi Li 1 re E NAs та] : Ў exhibit nearly clocklike substitutional n ose positions that are "free" to vary: |. cha ppro i the low va me mig responding patterns of homologous an pia wa йа similar hire structi Sa nder: 1992: al recon 9: 341- To gain insight into the topological элү: of vastly asymmetrical time samples (see Fig- A have со b informat ion from ' pteridophytes,” *'gymnosperm*: rm» (Table 2). If the substitutio hee: css i na] proces angiosp s effectively strings. The rbcL data are examined Volume 81, Number 3 1994 Albert et al. Functional Constraints and rbcL Evidence data transformation ese are listed by taxon and by TABLE 2. rbcL sequences used for cession n GenBank a where available, is given by these sourc The umber and/or pesme reference where sequence data first appeared. Voucher information, Taxon GenBank accession or literature reference Conocephalum conicum (L.) L Lop: fa күш Ца АДЫР ) Dumort. Ж eros punctatus Andreaeobryum macrosporum Steere & um engelmannii Prantl m (L.) P. Be [ nopoda J. Lycopodium digitatum A. Br. ae evecta -— Forst.) Hoffm. Equis: arven Selagi Botrychium ae та (Sav.) Underwood Taxus x media Taxodium distichum (L.) Rich. odocarpus gracilior Pilg. Ginkgo biloba L. Cycas revoluta Pei érioplis wad Baill. Zam ermis Vovides, J. D. Reese & M. ‘Vie rres Ephedra prp С: е Welwitschia mirabilis Hack. б Gnetum gnemon L. Ойы japonicus Siebold Pip. per betle L. ys ۴ Tasmannia insipida Calycanthus chinensis а “г s. T. Chang Eupomatia bennettii F. Mue Magnolia macro pice L Persea ame. a Mill. Trochod, Mao) paraa Siebold & Zucc. Ceratophyllum demersum L. Nymphaea odorata Aiton Lilium superbum L. s. atanus pom Ly altha palustris L. iud ia indica (Kellogg ) Hjelm Betula nigr Casuarina litor Hamamelis rites Oliv. Chrysolepis (Castanopsis) sempervirens mq: Mishler et al., 1994 Mishler et al., 1994 Mishler et al., 1994 Mishler et al., 1994 11058 (J. R. Manhart, in press . R. Manhart, in press . R. Manhart, in press . R. Manhart, in press . R. Manhart, in press . R. Manhart, in press . R. Manhart, in press 13474 (J. R. Manhart, in press al., aay: L11059 111054 L11055 X58135 naa et al., 1992) Chase et а B. Schutzman, s.n., FLAS, (M. W. 1993 Chase et al., Chase, unpublished) L12683 (Chase et al., 1993) L12677 (Chase et al., 1993) Chase et al., 1993 (C. R. ved L12680 (Chase et al., 1993) L12640 (Chase et al., 1993) - Ai v ets leotides 1184-1428 from M 776020 (1 t Dis e et al., 1993 M77035 3 et al., 1991) plus nucleotides 1184-1428 from Qiu et al., 1993 112682 Fret et al., L01943 (Albert et al., 102431 (Albert et al., L01903 (Albert et al., 1992a) 1992c) 1992c) 1992c) Chase et 1993 L01889 ale rt et al., L01893 (Albert et al., L01922 (Albert et al., 1992c) 1992c) 1992c) amino acid leve] for jer Pag seg with the nucleotide and s string e NUCLEOTIDES available in The up is id smallest unit EEE et f -time), becomes s, nu- ber of changes per site, А (= rate large. Unlike some morp hological character onde data are usually — — with Bn os Fitch, 1971). For such aes ures, t al. (1993) examined the poten қ 2t uen A info e s y given aa, EL data are sub- the num ject to ا‎ among sequences when branch teraction under Felsenstein nk wey lifi enario. tour plified fi ities with Jukes-Cantor (Jukes & С dien 540 Annals of the visu Botanical Garden Kimura 2-parameter (Kimura, 1980) с correc- udis for multiple ch culations oom a ver гү e P - de: be ex баб, , provided that А values remained small e e., less than approximately 0.1; see Albert et al., 92a). For quasi-ultrametric data, differences in : ree must principally result from divergence time differenc e bryophyte lineages examined her could easily be nee Aa the ан по pu pons: Devonian; the seed- pleats: ч "m by the y ret ous, followed by their diversification t toh he Ter- tiary—a ише. yange porni ially spanning 500-5 million y t. Thus, even without a priori гонан of precise үк Penn M: is h retention indices (C and R, ee ЫШ & Farris, rate Farris, 1989a) Five hundre d fifteen nucleotide с гоа АШ patterns of similarity among ta Eight equally parsimonious were found (C = 0.362 gE all шш R= "0з is ü The strict and combin trees p mer, 1990) were Мес pes Tie: 3) trees indicate that (i) horn ted inside of all other seed aie (iv) Mies. cycads form the monophylet xt MU giosperms, a and м monocots are basalm Piper. Characteristics (i) lysis of XC dit analyses of Стапи eu. Doy tevenson т КЕ м 4 E fa 1 © divergence. The mea: rate for woody taxa (Table l) averaged for sin in, ivergen bert, 1292) range newer 2.5- я. idu n is . Ав 8- ange 1.0-3.5 x 107 was well, À valu ues are д ыр to lie between 0.05-0.175 (500 My) and 0.0005- 0.00175 e M y). is a fou: axon spurious branch attractions M je et al. ‚ 1993). Here, we are working with 40 taxa and a greater potential for inconsistent rie (scd Penny et al., 1991). Data с. -Nucleotiia sequences {шә ambig 5'-most жойы, se incorporated only ime information for some le 2) were vin with PAUP 3.1.1 EY LN e Fitch criterion (Fitch, 1971; cf. Albert et a ge with N ecole e optimi- zation (Farris, 1970; Swofford & و‎ 1987) he пай до SEN was used w e PSE, MU ULPARS, and m R (tree bisection-re- Кел лайша branch-swappin d Nixon et al. (1994). pa (i) is in confit vi beth mosphelogipe al and molest ar cla- & С d 1985; Mishler teristic (ii) e mer, 1985 al finding et al., 1994, rem ssue). Cha both with m: Kai ae with the chlor Raubeson & pedis (1992) tha phytes е the lycopods, siomorphic (i.e liverworts) state. Characteristic e results of morphological (Don- , 1989; Loconte & Stevenso pe Taylor & Mieke, 1992) and some rDNA н & Zimmer, 1992; cf. Zimmer et al., 1989 me 5 ж gE e I] Function and phylogeny. Ме edless to 54у, ytoge observation ns can x the Ф e e reflect primarily sp than bordes homologies hi possible constraints on rbcL aid hn E amino acid e implied on Ficunes 3-5. Gh the Co mbinable component consensus trees Lor ше Tons of parsimony ' а al; for pe а tide, (4) string, and (5) amino acid data. For iie мерес what trees t would Bi rwise et гіч, Implications of the үш, а аге Кыраа. in the text. Conocephalum lea Conocephalum Angers Sen loss pg. sora ч) ele Ф Ф Ф EFS 3g 3 bu Oz у ae ym ре! трһаеа eratophyllum Жз ГҮЛ 5 Calycanthus Casuarina pede eii A Andreaeobryum (5) g lymphaea imys oratonhvllium eratopiiiynadm OQ. ERODE ES] & zm = v661 € JOqUINN ‘1g euinjoA eouepi^3 71244 pue 5}шедиог) јеиоцоипу ле зә ueqiv Ls 542 Annals of the Missouri Botanical Garden bremobes. of one dios the eight equally most-parsi- D). As summarized in Ta- ics 3, over 84% sof gi eode nucleot ide substi- utio; ches are ns on interna al b to amino acid identit onde ity. “The e perce ея MEME functionally la odds matrix of Dayhof ct etal., 1978: 352; see es Table 3) amount to an additional = 13%. Viewed as a whole, 97.5% percent A all synapomorphous nu- ерше сһапдез аге ке to have little or no ke ER Ар . With um of only 2.596 of these rie nges incurrin e amino d replacements of potential мау functional distinction ieee Tun le 3), rich sequences рреатг heavily burdened by tio tiene conser vation “Тыш, the challenge for land- ae t cladistics is to ae mine how strongly func- nally constrained variation may ^d reflect phy- Eois patterns STRINGS Thei "unit" character in phylogenetic anal- ysis is one that truly evolves as an independent unit, m eaning one RE peau undergoes = I - are racial correlated (i.e., congruent; et. Far my Peery with "— of Saat Jos Жашы £ ers. eet iho a contiguous ct of nucleotide D on/deletion event, several non- ositi that i mosom: a karyological change s of course difficult to assess such possibilities a priori, but it is nonetheless important to begin to develop meth- mine the issue чәр. We have thus examined som ns by which the functionat ‘phylogenetic evidence fe р in y dat ata forms other than nucleotide positions and fee ari states. The nucleotide is indeed the smalles it character in ric evidence, but it is not t necessarily the mort informative ең. m t. First oan racters may restrict potential topological res- olution (e.g., a 4-state, nonadditive character can ost con- т '° Patterns of codon usage intrinsic to the primary nucleotide matrix are also bp anti of anes: con- straints; th paper (Albert, Backl veal & iin in б have minimum homoplasy if optimized а autapomorphies). pepe direct 1 of nucleotide sequences from g genes ig nores Lada impose both ia hs genetic ci codon positions may be both intra- and inter-correlated (Fitch & Markowitz, 1910; Fite h, 1986). A data transformation that may overcome these revolution: pr toic tion o Our pron ene (analagput to > pen ing mapped t y s lined thus: (i) a erate пона of ави А, Т, T and C соп varying г eir da in size between 6 an pairs (so a a minimum and maximum of two and seven гойт аге include), (ii) ium rbcL nay ҮП data for ин record г recognition by both base position and tax- on, (iv) treat multiple positional recognitions by а given E string separately, ү, treat all гес- ognitions found in two or mo a as bina characters for PEN analysis (sequences that h: ion at osition аге n е 1 ring characters, unlike the approach used here a below and Ap- peg sis : effect of transforming seque nor strings i » чеччи 0 " incorpore E in in a larger unit character, and (ii) decrea N ses the that independent ira 1983; De eBry & Slade 1985; Albert et al., historical branch er ted paralle 1989; Therefore, ers much less likely to engage in b — ch occurs because of accumula f. Felsenstein, 1978; Hendy & Penny, Volume 81, Number 3 Albert et al. 1994 Functional Constraints and rbcL Evidence TABLE 3. Analysis of character support for internal ca of tree #1 (of ne rom the nucleotide analysis. "Node" refers to the node numbers on the reference tree of Appendix I. “# changes" refers to the total num of nucleotide changes optimized onto a branch. “Constant” ا‎ аб that P E site bela ange in a codon Кр that codes 19 the ; same mino. ecd throughout the entire matrix. b two о change indicated at this node does not cause a ns i m road acid sequence. Pact means mun the inferred м эй ү } change in acid due to zi observed change : ely to happen by chance or better (according to the PAM-250 log-odds matrix “of Da Кы et at; 52). “Potentially nonlabile" indicates that at least one of the тонні piis. acid changes interrog from a pera nucleotide position is not likely to happen by ra T g are likely to happen by random chance or better. **Nonlabile" 1 ll inferred acid changes (of ly on y с less tl 1 h Potentially Node # changes Constant No change Labile nonlabile Nonlabile 78-77 42 22 4 8 5 3 71-16 24 13 6 4 0 1 76-71 at 13 9 3 2 0 71-70 29 19 9 1 0 0 70-42 40 24 11 5 0 0 42-41 33 26 5 1 0 1 70-69 42 17 16 8 0 1 69-66 29 21 8 0 0 0 66-48 34 15 13 5 0 1 48-44 25 10 12 2 0 1 44-43 29 19 8 2. 0 0 48-47 15 7 8 0 0 0 47-46 24 14 7 3 0 0 46-45 ll 4 4 3 0 0 66-65 56 34 15 Т 0 0 65-64 26 13 10 3 0 0 64-63 18 11 6 1 0 0 63-54 5 2 0 3 0 0 54-53 4 3 0 1 0 0 53-51 10 3 1 5 1 0 51-49 9 4 2 3 0 0 51-50 8 2 1 5 0 0 53-52 11 5 2 4 0 0 63-62 16 11 5 0 0 0 62-61 14 6 Z 1 0 0 61-59 8 2 4 2 0 0 59-58 17 8 © 4 0 0 58-57 13 6 4 3 0 0 57-56 33 20 6 7 0 0 56-55 6 3 2 1 0 0 69-68 58 29 18 8 3 9 68-67 45 24 17 4 0 0 76-75 34 20 7 4 0 3 15-74 38 23 12 2 1 Y 74-73 45 28 14 3 0 0 73-72 65 43 12 9 1 > 529 272 126 13 11 92 00% 55.63% 28.60% 13.25% 1.37% 1.16% 84.23% Annals of the Missouri Botanical Garden Albert et al., T9928 1993) and much more likely to contain “blocks” of evolutionarily correlated inform ation. Nevertheless, bue information co be functionally constrained, as with primary nu- pod pum This possibility can de studied sim- ila Mir M ames infe тей amino acid cintas traced to its y recognized codons and component ucléc- tides One thousa nd random strings After sca 0 quences, 193 posi- tionally dist ring recognitions were recorded (mostl m small strings, the largest being from a 15-mer; see Appendix II). Of these, 112 iden- string recogni independ spect to mm identified (ы Appendix n Therefore, our string data carry an experi 1 о what could occur with restriction site data representing mapped c several endonucleases. The “s transfor J. Farris, sénpublisbed) avoids this of presence/ absence data, would be identical to E r string data should suffice to explore biologic n-indepe ce of nucleotides (functional con- straints); in fact, partial replication of nucleotide "blocks" could enhance detection of conserved n ions. Cladistic pa of the string characters s performed under th —€— ا‎ (Kluge & Te со Farris, 1970; et al., ptions men tioned INVI 165 е ually arsimonious trees e found (C = 0.3 ata), R = mponent consensus ие differs from the strict by ub Fig. The g data provide a different resolution of land- etsi pray netales to ms (with Piper basalmost), (iv) par- port ofa angiotpermdt s comiter s + (Ginkgo, cy- д 1 N Nucleotides, above), Function and phylogeny. hat clad 1 Ir t It could be argued g ۴ g rmed data are better phylogenetic representations than those derived from nucleotides because the unit a r is substantially less subject to parallel гус this pecs is du ure of the his be ment act s differences i in tree to Sms disais paraphyly of angiosperms) may simply result from different representations of func- tional and phylo nt history in string versus тзт Чата We hav ааа им i constraints on чем evolution (as above) by examining the red amin = л, оп a ner branch- es m one equally m rsimonious үш ш Apnd IT). Striking pras from 3) are shown Е the е amino pe replace- ximum amino dee re puce ode amou a max asc ph to internal stop codons, which may ko mee sequencing rrors). This greater number of pre ur pe eo icate a greater chance that dune seri rticular nucleotides may bias tree © different ыле ig patterns between nu- cleotide and string can be explained by in- herent быны of pe latter. Fach string rec ognition s c much m explain rou, recognition identifies a functionally co ed p tif, the larger the motif, the Ed the like that functional preservation need n t require exe Volume 81, Number 3 Albert et al. 1994 Functional Constraints and rbcL Evidence ABLE 4. Analysis к — kp for internal branches of tree #100 наа 165) from the string Asp ма pies to nen —_ n the referenon dre e of A елаш lee! ‘He pang! en to the tot n d on a onstant" indicates ab t de for the same бү acid thr se ae entire matrix. “Labile’” means that the inferred ема in amino м Чие 8 бе perve d АЕН їп string recognition, is likely ep appen by random ‹ chan nce 9r better (ccording t m the ТАМ; E y , 1978: 352). " Potentally nonlabile" amino acid changes inferred from a partic og ring recognition is not likely to happen by random, but that there also are some e changes i in the same бат eid b ad are md to ha appen by random chance or better. “Nonlabile” i ] 1 h “Internal stop” refers о J e)o to string 8 that е | st р а ‚ which тау b q ГА Potentially Internal Node # changes Constant Labile nonlabile Nonlabile stop 0 0 ^ T > > © 4 олм М2 TO RK Фф دن > دب یډ س‎ ОО > O ОО © фә لد > دن‎ Мә (л С > ید ہہ ےی جب حي خي ص‎ Dewey nm QOO س س‎ NRF OR س‎ tl ین سا‎ оо > ке ке ке Q0 I سا ی سم‎ м > > س‎ л фр mer NOF OF О © © юк номоо к о юҥ+ кюк RP © юе NH моо мо —"—O-oO-c-ocoocoocoooc-wooo-oovcovovoow-ooonoc- t0o0-——-0wv-o0-o0-—--—---—€wW-wW--cooouwcococooo-ococo-oc e T е a - 1 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 4 23 15 69 39 21 22 is 100.0075 44.51% 25.16% 13.55% 14.19% 08% 69.67% 27.74% amino acid identity. Strings recognizing regions of Again, these are probably f found in greater pro- non-labile cha ange, indicating potentially TIN portion because the larger size of the unit char- Changes in structure and function among taxa, may acters. Rather han beng co nserved ause of represent another Жз of conserved information gnit Annals of the rebus Botanical Garden es Меру ова markers foe historical Dicas айе (see аот et al., 1990, o rbcL ; ae Nei 270- -271), but they could be of s t DNA r TRIBE r, 1987; Palmer et al., 1988; ; Downie Raubeson & Jensen 1992) if vell. i in n Ro = 8 5 = o: © ay m) HE 87 Es pman et al., 1988; Andere et we “1989: 3). (Cha cf. Clegg, 1 AMINO ACIDS cause rbcL nucleotide substitutions approxi- mate a es hypothesis (see The “Problem,” above), amino acid ch to sociations a characters may bias tree con- struction. Topological resolution may aa im- ited because amino at ata is о POTS kei (Fitch, 1971) and more than iur states could be available for given characters (in ata might be more suitable for bridging large evolutionary time gaps, given a roughly constant rate of substitution combined with ignorance of Hence, we evalu d the amino acid data [о hi- erarchic compatibility with the results of the nu- cleotide and string analyses ta analysis. After ‘ IN the 4 rbcL sequences, 66 (out of the 476) a positions identi r more taxa. Cladistic anal- ft cluding all data), R = 0.554). The combinable бе. опе тоге amino acid yet + aa гезо- lution of land-plant lan (cf. Figs. 3, 4): (1) lycopods are polyphyletic, with Isoetes sister to ыд RR el Anthoceros is embedded a lli erms as a whole (wi ne со- ntes таур) а are е the monophyletic sister haea basalmost), r s hypothesized from арален аге in onflict wi 0 arbiter of cong ‚ lar amino acid data are no more appropriate fo for he: ic time samples le be sensitive to differences in divergence times. Amine ac duis Fy ti. F i Ln 7 аны: о in rbcl ] l constraints бее Nucleotides and Strings, Ee 0 e that amin selective neutrality may be labile amino ac ^ re placement which could simi- mie “wobble”’ evolu e. Only a sm ar pasion: of individu ps rs to be involved in fun evolutionary events (see Mihi. FO PENULTIMATE CONCLUSIONS ve devons. x problematic, func- ined n rbcL markers cul d powe proc ating cladistic history or discrimin: llelism Т pe талаш d par and reversal). Volume 81, Number 3 1994 Albert et al. Functional Constraints and rbcL Evidence TorAL EVIDENCE AND CHARACTER CONGRUENCE (I) ON CHARACTERS Every character in a data matrix showing sim- dass» Ң + 2 Ж) 2 " | 3 / г parsimony аз a discrete and independent piece of rmation. This holds whether or not the char- acter represents a si i " ares analysts is expected to nave a single e; see Riedl, 1978; Donoghue, 1989; Donoghue Ч ү Sanderson, 1992); ns айне ү fue op- at maximum parsimony along with all other char- acters in a matrix. From our argument about shared Маден history (constraints) in ү ps ox of rb hao (eg B^ nuclear v versus cellula endosperm х i2 Ней unlike г iven taxic homology, phas is pecu of Bib: discrete points of information, that is, its ca. pan с То а озшш) unm rithm, h | debate on this issue, see ot ones et al., 1993; Nelson, 1993; Barrett et al., 3; De Qüeirás; 1993). In ct, acceptance ^ parsi rbiter of o E 5 e =" دو‎ - duplication being part of ks кшш регар) коа Ка, use of par alogous Histories (Fitch, 1970). Nevertheless, analysis о tal” evidence (sensu Kluge, 1989) gives each dn Жый spear. i meu ш same HE asa aren mor- nd ph о hier ar chic поп нын taxic homology nitent p tibl pollen unit in the oF chidaceae: monad, tetrad, mas- linium," whatever its underlying com- Нее се, some мАг have found cladistic phi- losophy and m jen with other iy Sigo i te recoding of gene trees (Doy a or Bro eU of compet cn pat ty жен. Legitimate concern over potentially sepa- - м argue below that both approaches — hierarchies, ne par: рез [знн posed Aie quandary. a feature pager ian to the ован: ке Farris, 1919, аг simony i (П) ON EVIDENCE For cladistic «nir evidence is the body of siu پا‎ o at шше patterns of simi rity am казлы а A specific set of evidence "y iun in differe: een we have shown this Коро г fferent data transforma- tons of the rbcL gene (above) ырайда - : € evidence in the form of tree тропе 9 Зо at the cost of information content pi recent fab ^ all ои NP (sensu елен 1969) with other data. (Ш) AN EXAMPLE tent to which rbcL evidence shows hi- The erarehie ‘correlation with othe " paw should ts freedom from bi Sica s ieri conside азаи баду its ка utility. In acter interaction between rbcL ev- п the ee morphological seed-plant matrix of Nixon et al. 1996: Us mg the, set of and consequen- this context, we “constant,” we tested the ability of different re L s (i.e., nucleotides, strings, and а ified representation К E ce, have used the retention index: t б proportion ч MUS V r4 г о Seer 1989a, b, 1991). Although = is not эд е Goloboff, pm oath matrix within our re- spective sets of experiments shares the same “соп- stant.” Additionally, each data transform of rbcL Аппа یر‎ od Garden Homoplasy and character congruence sta- volvi fossil and extant taxa, 101 morphological similarities are relevant (symbolized by “N”); for extant only, th ed by *N,"). The T f relevant similarities for each rbcL dat е cleotides, strings, amino acids) are given in the tex Жо алама накои ossil taxa, rbcL evidence was repre- m ?”). Consis- 5 tency Retention # Trees 7 nl wees 45 М + nucleotides 0.450 0.625 44 N + strings 0.402 0.685 22 N + amino acids 0.467 0.710 309 Extant taxa only N.. + nucleotides 0.464 0.601 3 Be + strings 44 .641 7 N.. + amino acids 0.518 0.670 24 . Finally, not u uggest which analysis(es) may be “better. rna ara otera and cladistic ipsa daga for h (Nixon et al., 1994). We used the same dissi: ethods s out tlined an to examine six combi LF & 1 uy taxa апа 1 x о {атта tions. бо istency and retention ered ir e 5, each analysis are reported in Table topo- logical results are summ d in Figur s 6-8 Character congruence, as Weide gae re- tention, is similar in magnitu 1) across hati ri gnitude (range « 0. h se of experiments, “Although торба сат гез- with заров to the roeL data form used (Figs. 6- 8; s бо a al, 1994), the rbcL he n i consistent statement along with the morphological evidence ith respect to extant taken monophyletic cy- ll erpii conifers "ie or monophyletic ren these taxa (Fig. nt-only analyses, Ginkgo simila arly in- айы е cycads and conifers (Figs. 6b, 7b) or remains sister to conifers (Fig. 8b). Conifers monophyletic in a de a А @ а e апрїозре Ephedra is ister to Des plus Wel- d hp: феа жуш Bennettitales is pro- vided о nalysis with amino ined a si data (Fig. Ва). айн: Пит is placed sister ام‎ et iip i (see Les, 1988; Chase et 3; Qiu et 993) in the combined nu- otide and sring niil qun бм b, 7a, b), but in the bined amino Fi ecisi igure 7a, and all other on this point. cek ngos with triaper- turate or tri pollen; here, Plat- anus, Caltha, a ME Dillenia, Hama- Casuarina) are m e = S 0 © ~ ® i E e w ic] ~ є E 8 string, or amin at different Е of morpholog- ical m a? (of Newel et al., 1994) show con- with these different ы forms. If оп were to o held th the evidential significance of the mor- PORE data qestan on e might Мое е меге incongru ruent a them in h data form p^ pee ору; because congruence is ап voe total interaction, the utility of either set phe idence is always E relative to = er wb ertheles. one subset of total Мес if, as in tn berti rbcL, it is reasonable to assume a single, vestigator were = а al hist tory: if an an =“ — Bh. = eses of correlation bekedi E cons Volume 81, Number 3 1994 Albert et al. Functional Constraints and rbcL Evidence Ó(b) Tot morphological idi a dis 6(a) nd taxon sil a Cycadaceae Stangeriaceae Zamiaceae Ginkgo Podocarpaceae Taxaceae Taxodiac./Cupressac. га Welwitschia otal evidence analyses of morphological and rbcL data for fossil and wind of Nixon et al. ese matrix versions as of 8 Novem Ma Um TRI] Aneurophyton Archaeopteris Cordaites Podocarpaceae axaceae Sl eames Corys poc Tatarina oe Cayton Wiliamsoniella C Williamsonia Ej r Welwitschia netum Ceratophyllum Winteraceae Eupomatia Magnolia Calycanthus ersea Chloranthus {| Casuarina extant seed plants. The ner eee st followed yrs tic rte Я foss combi ий ч опе ‘of three da ta forms: the “b” series). тео ide — (6), e 2). For — Annals of the лам Botanical Garden Aneurophyton [ Агс 000 Lyginopteris Medullosaceae Cycadaceae Stangeriaceae Calistophyion inkgo Gingo " Podocarpaceae axaceae To osperm Caytonia Williamsoniella садеой Williamsonia Ephedra Welwitschia netum Ceratophyllum г Cycadaceae E ag Stangeriaceae Zamiaceae 9 re дй ее 1а саа ахь; Fel character states were scored as Per (ie., “?”; cf. Platnic wofford, база ог the strict со ТаЫе 5). Ѕее € ei sus r fi Mitis Betula a Casuarina me iliu; Chloranthus Piper Casuarina 1 k et al., 1991; 1002) cl nt eithe le tr 24). T ther peni 3.1.1; Swoffor foun се — component consensus in all 2 of all nd -parsimonious trees ШЫ discussion Volume 81, Number 3 1994 Albert et al. Functional Constraints and rbcL Evidence = 8(b) Tascdac /Cupressac. Ephedi Weiwitschia Gnetum Chloranthus Pi Calycanthus ersea Ceratophyllum ilium Platanus Caltha Dillenia Chrysolepis Hamamelis Betula Casuarina Winteraceae Trochodendron Nymphaea Eupomatia Magnolia Aneurophyton Archaeopteris Lyginopteris Medullosaceae Cycadaceae Fe Sh ск Eun [riri 'ycadeoidea Williamsonia phedra Welwitschia Gnetum Winteraceae Calycanthus a Chloranthus Ceratophyllum Annals of the Missouri Botanical Garden and шке history could be generated from the congruence patterns of each rbcL character CONCLUSIONS e phylogenetic informativeness of rbcL vari- su with confidence: (i) rbcL ce substitutional behavior, sdb may either help or hinder бе! о depth end asymmetry of a а given phy- xist over the majority of informative nucleotide танан ЫДЫ 18 5 expected from @ under the i) eviden pakes g Es ri strings, or amino acids) J (e.g., morphological da ta). Although rbcL trees often appear consistent with taxonomi are substantially congruent erns of char- acter раа Among such diverse information tory significantly more powerful than . «ud af rbcL alon LITERATURE CITED v us T A. & B. D. MIsHLER. 1992. On the Кең и: = By pa nucleotide sequence dat Gadel dies D & K. BREMER. DNA char and. сабабе "The optimizatio In Models in Phylog: aracters n of functional rid enetic ep ae an The Sys Systematics Symposium, Missouri Bot. Gard., St. Louis [abstract]. , 5. E. WILLIAMS M. W. CHASE. 1992c. Carnivorous plants: Phylogeny and structural evo- lution. Science € 1491-1495. S T, С. SCHNEIDER, Y. LINDQVIST, . LUNDQVIST, rani RANDEN & С. H. —_ e 337; 229- 2M, ade DONOGHUE. & E SOBER. 12 Against c consensus. Syst. E ee 486-4 gg re A reply to Nelson. Syst. Biol. 42: nes aig TE IA TRA И & R. A. Pri 2. LO variation in bua: rate d tipa: sequen dr: d Pun s. Proc. Natl. Acad. Sci. U.S.A. 89: 344-7 BREMER, В. EK REMER. CHE analysis of blue-gre ocaryote interrelations ships and chloro- plast deem um on 16S rRNA oligonucleotide cat- alogues. J. E ol. Biol. 2: 13-30. BREMER, K. 19 ummary of green ae phylogeny and аа а 1: 369-38 Combinable component consensus. Cla- == BRUNEAU, А., + J. Dovrs & J. D. erre 1990. А ipa oplast DNA inversion as a subtribal dai ead lm (Leguminosae). Syst Bot. 15: 378- TONS SEE W. Sun, P. M. С. CURMI, D. Cascio, ‚ W. SMITH & D. S. EISENBERG. 1988. Tertiary structure n Arek TL Domains and their con- tacts. Scie CHASE, M. W dp ium E ©. OLMSTEAD, D. MORGAN, Н. Les, В. D. Misuter, М. R. Роули, К Price, Н. С. Huts, Y.-L. Qro, К. A. Kron, J. H с, E. , J. D. Pater, J. Е. M , К. J. Syrsma, Н. J. MICHAELS, W. J. KRESS, AROL, CLARK, М. Heprén, B. S. Gaur, R. К. JANSEN, К.-]. KiM, C. F. WIMPEE, J. F. ўа, С. В. FURNIER, S. Н. Strauss, Q.-Y. XIANG, p . PLUNKETT, P. S. S. SwENSEN, S. E. Wil- LIAMS, P. A. GADEK, C. J. Quinn, L. E. EGUIARTE, GOLENBERG, С. Н. LEARN, JR., S. W. y S. C. H. BARR ЕТТ, S. DAYANANDAN & У.А 1993. Phyl nts: Ап m^ ni nucleotide sequences E ds p gene г Ann. Missouri Bot. Gard. 80: 528-5 E CLEGG, M. T. 1993. Chloroplast gene jae S . Proc. Natl. Acad. 50. the study of plant evolution tematics A Oxfo: a ress. U.S.A. 90: 363-367. A and ‚ М. W. CHASE & ar D. Мни нен. Сһаг- G SKI. 1992. Chloroplast DN and acter-state weighting f i the study of gn PIS Present status E coding DNA sequences Ann. ride а Card. future prospects 13 i "^ tics 80: 752-7 Soltis &1. J. Dose (eis Molecular Sy. ‚ B. D. MISHLER & М. W. СНАЗЕ. _1992a. _Char- of Plan eee: єз 3. Tribal acter-state we bue for restriction site data in ph Nn, E., n Езснв ан te y >м 1993 heli logenetic reconstruction, with р implications s rom eh roplast DNA. Pp. 369-403 in P. S. Soltis, D. E. sequence date: Апп. Missouri Bot. Саг Soltis & J. J. Do yle es Pr ic i ене 685. f Plants. Chapman а d Hal ‚ New York. CRANE, P. R. 1985 Padus yid d ‚ К. BREMER, M. W. CHASE, J. MANHART, B. D MISHLER & K. C. NIXON, 1992. Tch gene se. sige and поен diis of vascular plini 13-14 in program booklet “Origin and Rela- ае of the Major p Groups," 39th Annual er per Рр. 345-352 in М. О. Dayhoff peas Atlas Volume 81, Number 3 1994 Albert et al. Functional Constraints and rbcL Evidence Protein Sequence and Structure, Vol. 5, Suppl. 3. биом! Biomedical Research Foundation, Washing- ton, D.C. DeBry, R. W. & N. A. SLADE. 1985. pe pp Am restriction endonuclease cleavage maps within m-likelihood framework. cem Zool. 34: 24- De coim А. 1993. For consensus (sometimes). Syst. Biol. 42: 368-372. UE, M. J. 1989. Phylogenies and Ше айын е evolutionary pi nir s. Evolutio JA 1989 Phylogenetic analysis of dcin end > relationships of Hamameli- dae. Pp. 17-45 . Crane & S. Blackmore (editors), Evolution, асе бе Fossil History of the Hamamelidae, Volume 1: Introduction and “Lower” tone elidae. Clarendon Press, Oxfo & M. J. SANDE 19 The suitability of molecular and morphological evidence in recon- structing ye phylogeny. s 0-368 in P. S. ltis, D. E. Soltis & J. J. Doyle (editors), Molecular of inn. аа nd Hall, New York.‏ ا Downi, S. R. . PALM 92. Use of chlo- za st DNA و‎ in reconstructing plant phylogeny. Pp. 14-35 in P. S. Soltis, D. E. Soltis &xq. р Doyle ens “Molecular фин of Plants. Chapman a nd H mre "nich 6. кук р. E. SoLTIS, Р. S. SOLTIS, J. C. JD MER. 1991. Six indepe: e: UE of aes кы өз DNA r intron in dese tyledons: Molecular and phylogenetic implica "à iege cot -— 1259. Dove, i ^w m ра 1986. Seed plant phylogeny ae the origin Жала bud osperms: Ап exper- imental cladistic Meiste Bot. Rev. 52: 321-431. TOUR Fossils and seed plant phy- а logeny Sauk о 44: 89-106. YLE, J. J. 1992. Gene trees and species trees: lecular systematics as one-character taxonomy. Sys Bot. S 144-16 Farris, J. S. 1969. A successive approximations ap- ns to character weighting. Syst. Zool. 18: 374- 970. eran for computing Wagner trees. S гт 19: 83- ле т 978. Inferring pirate trees from chro- Mun: inversion dat . Zool. 27: 275-284. 79. The nbi rir of the phy- logenetic system. pas Zoo 1. 28: Mm Es em ets ck T e Fu "i Cli pe in ы Vol. 2. oie Univ. s, New Yoi — a. The retention index and the rescaled consistency index. ил, 5: 417-41 7———. 1989b. The retention index and homoplasy ae Syst. Zool. 38: 406- 407. Habes E 991. Excess homoplasy ratios. Cladistics 7: N, J. Cases hich parsimony or esee meld will н, чанка misleading Ase st. Zool. 27: тен, W. M. 19 910, Distinguishing homologous from es proteins. Syst. Zool. 19: 99-113. ~~~. 1971. Toward defining the course of evolu- tion: Minimum снарае for specific tree topology. Syst. a e ee 406-416. The estimate of total nucleotide sub- d trom p Doris is biased. Philos. Tra 316: +) Ser. An improved method application to the r f fix of mutations in evolution. Biochem. Genet s 19-593 Fox, G. E., E. SrackEBRANDT, R. B. HESPELL, J. GIBSON, J. MANILOFF, T. A. DYER, R. S. WoLFE, W. E. BALCH, R. S. TANNER, J. MacRuM, L. B. ZABLEN, R BLAKEMORE, R. Gupta, L. Bo E B К Lewis, Р HRS) N & A 1980. "The phylogeny i sie pa eag бі. епсе 209: 457-463. СА " B. S., S. V. MUSE, RK & M. T. CLEGG. 992. Relati tive rates of n otide substitution at т rbcL locus of RR plants. J. Molec Evol. 35: 292-303 — M. T. Сіксс, C. J. N&G. зз. agnolia s ature LOBOFF, P. A. Homoplasy ind he among dumis Casi 1215-2 Hampy, R. К. & E. A. ZIMMER. bosomal RNA аза phylogenetic tool in plant systema Pp. 91 in P. S. is, D. E. Soltis & J. J. Doyle (editors), Molecular CS of Plants. Chapman and Hall, New Yor Немрү, M. à & D. PEN 1989. A framework for the quantitative ке ^t evolutionary trees. Syst. Zool. 38: 297-309. Hupson, С. S., J. D. MAHON, P. A. ANDERSON, M. J. T EWS & GIBBS, M. R. BADGER, T. J. AND P. WH 1990. Comparisons for rbcL genes for the large subunit of ribulose-bisphosphate carboxy ase from closely related С, and С, plant species. J Biol. Chem, 26 "6 JANSEN, R. K. & J. D. PALMER 198 7. Chloroplast DNA from 1 d Barnadesia borse Struc- ture, gene adici n and characterization of a large rsio ү; -564. Jones, T. R., A. С. KLUGE & A. J. W 1993. When theories and methodologies Шы А Md at amanders. Syst. ngs 42: 92-102. Jukes, Т. С. Тр 1969. iud х ргоїеїп molecul 21- in H. N. (editor), Mammalian E Metabolism. e ork. Press, KIMURA е. A simple method for sc ya evolutionary rate of base substitutions through co parative se Se of nucleotide sequences. J. Molec. Evol. 16: 111-120. 1983. The Меша, se a Maece Evo- lution. Cambridge Uni PW кс. Syst. Zool. FARRIS. 1969. Quantitative phyletics d f Sy st. Zool. 18: 1-32. Kron, К. А. & M. W. СнАЗЕ. 1993. Systematics the Exicacess, Empetraceae, fe дан and г 554 Annals of the Missouri Botanical Garden lated taxa based upon rbcL sequence data. Ann. 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S. ена ELWICHE, Е. W. ZECHMA! р ї. Саг : 4 NEI, М. 1987. "уос г Буйло Genetics. Co- lumbia Univ. et аы ork. NELSON, G. 1993 reply to Barrett, е and Sober. Syst. Bick 42: 215-216 NIXON, К. C., W. L. Си. D: STEVENSON & d. 81: Ais . & M. J. Nov 1992. Congruence en yo uie = phylogenetic patterns: Grape cladist Cla- шер, 8: 319-337 OLMSTEAD, R. С eae); inference Pu mor- phological and phones data. Syst. Bot. 14: 320- 338. Pace, R. D. M. Genes, organisms, and areas: 29 арш. of eee lineages. Syst. Biol. 42: 77- 1968. Molecular pa- [o of human maps di Nature 219: 902- PLATNICK, N. L, С. E. GRISWOLD & J. А. CODDINGTON. 1991. On m . Cla- distics 7: 337- 343, issing Ото, Y.-L., M. W. Cuase, D. Н. L R. PARKS 2: £ em h = E Magnol iidae ana quences of the asi gene rbcL. Ann. Missouri Gard. 80: 587-606. RAUBESON, L. A. 8 R. K. u— 1992. Chloro| roplast INA evidence on the ancient evolutionary split in vascular land plants. Scie 255: -1699, Rr mi 2 пее Order in Living Organisms. Wiley, erate à xu Price, К. С. Како, E. Cont, К. J. SvrsMA & J. D. PALMER. 1993. Nucleotide se- quences of the rbcL gene indicate monophyly y mustard oil plants. Ann. Missouri Bot. Ga rd. 8 1992. An D. E. Sorris & С. J. SMILEY. ri Parsimon пу, Ver er pr ae s manual а b the Шш meds ehe а Champaign, Illinois. W. Р. Mappison. 1987. ا‎ Г ч F. 5мтн & Р. ed MN 1991. x Т systems, — and chloropla ETE Old World species of Fuchsia vss P 57, "269. oe Phylogenetic giosperms vol. 180: pan Phy ogn inference from aps with partic- and the N, A. R. 1983. Phyl apie Ey какы. cleavage m о the evolution of humans 1990. Chl roplast ‘DNA evolves slowly i in the ut family. „Мојес, 14. Zimmer, E. A., R. К. Hampy, М. L. ARNOLD, о LEBLANC & E. С. THERIOT. 1989. ша phylogenies n эса ар р!ап t 214 in B. Fernholm, K. Bremer & H. p itors), "The O of Life: Molecules che phology in Phylogenetic dm ins Elsevier Publishers, Amsterdam Inferred amino es of a тей апію е4 ost -parsimonious) APPENDIX 1 (pp. 554-562).* changes оп the nin branch: qually Ly in- le diei accompan as ош an кач kse t the fui ШО. impact of PU acid as reflected by alterations n Mi uP tide 5 ( enin) Followi ing the apomorphy li T 1 (Swofford, 1993), each internal branc Volume 81, Number 3 1994 Albert et al. Functional Constraints and rbcL Evidence erence tree is identified by the nodes it connects. For ode pair Д prin (= POS,” the e tused); character Ey 1 index -1428 bases of the rbcL Cc; ea aR = f rh consistency of the entire tree; see Farris 1989a), Ше ual change inferred (^NUCA," with arrows follow: dé. conventions in the PAUP 3.1 1 m 00 Sc d labile 3 at the codon involving a given position, and (constant amino acid identity) are лр" a line of the following form 175 1.00 с в R, L, A NL be readily diagnosed: character 175 changes from nucleotide C to nucleotide G (on this particular tree; read “L. constancy of character- state reconstruction among all 8 cof 1 (1.е., no homoplasy), an ich character 175 belongs changes be ino acids А A (using standard ІОВ amino acid ее this does not necessarily те acter-state change gives the indicated changes i ino acid uence; rather, it ely indicates that it might be ed kc in sx (i.e., the cleotide trans tion not affect amino acid identify at all; thus, the ША. amino acid changes аге the “worst” that can happen under the influence "i eie cter 175). The NL designation i transfor- mation between R, L, and A would ioni a | icudabile change. In the line below 486 0.167 a> g a S -= чө is а пис otide 486, t can be post ж are as not ir for да different t amino ac a ide ntities in its associated codon (thus, the SC is giv e * Correction added in proof: P. 560, under “море 62-61," third line from bottom, right hand column, should Annals of the Missouri Botanical Garden 46 7 Lj, Slangeria — Zamia Lilium Piper Chloranthus | 2) iem Calycanthus Е — Persea 53 p— ji |r = Ceratophylum 51 ia БО, ros Nymphaea — Platanus r5 о, ov ELLO Caltha Trochodendron Hamamelis ЖЫЗ = | су => 3 o $ uo etul — Casuarina Conocephalum Volume 81, Number 3 1994 Albert et al. Functional Constraints and rbcL Evidence NODE 78-77 POS с NUCA 68 0.200 a->c 69 0.5 102 0. ae a->t 150 0.231 >t 13460.125 c=>g NODE 77-76 PO 11250.375 a->g 1137 0.231 a-»g S c NUCA AA 120 0.167 t-»c constant R,P constant constant S,C,Y A, i E,V,H,I О; Vest V,M,A V,M,A constant constant L,S HY, SF D,R L,M constant constant constant H, Q constant constant , constant L,M C,S constant T E m o ~ L,S,F,I,M constant vw о 11760.250 а->9 1254 0.429 t=>a 30.167 c-»t NODE 76-71 POS g 327 0.167 o 1398 0. 250 a-»g 795 0.250 t-> 13630.167 t->c E,D constant constant AA constant constant Bh A,W constant D o ` m ms 5 z < m n constant constant > , constant constant V,M,A V,M,A H, Y,S,F constant V, I,Lh,M Ix L,S constant ,„Е,А constant RUE, = AA E,K,Q,T Р ` Е А,Р,5 constant constant 1,8 V,G constant constant L,* L,e constant constant constant z L Pom 2 cf t NODE 70-42 ra s 0.500 t=>c de 0.429 t-»c 453 0. .273 a-»g 543 0.333 t-»c DAS а” id 1345 0.154 a=>t 1350 0.250 a->g 1371 0.200 t->c NODE 42-41 РО .750 а=>9 c NUCA AA P,A A, constant constant , constant S,T prets E,D K,R,E constant E ,S constant 5 L, constant constant 9 ts constant AS T;C constant constant AA constant constant constant constant P,T , constant constant constant 558 Annals of the Missouri Botanical Garden 996 0.400 a=>g constant 279 0.167 =a constant 456 0.222 t->a constant - 10110.429 a=>g constant - 315 0.167 g->a constant = 471 0.500 t=>g A,V - 1021 0.333 а-> ё‚Ъ L 412 0.200 t=>c S,L - 505 0.200 c->t constant - 10950.500 t=>c constan - 505 0.200 t-»c nsta 538 0.400 t=>c L,I - 11370.231 g->a constant = 534 0.200 a=>g constant - 718. 0.33 ->c constant - 11400.300 a=>g constant * 549 0.200 а-> nstant - 759 0.333 g=>a constant - 11640.333 t=>c P,Q,S - 600 0.333 t-»c constant - 768 0.167 t-»c C,F - 12120.429 g-»c constant = 624 0.667 t=>c constant = 8625 0.375 tag ote = 12960.333 t=>c constant - 663 0.500 c->t V,C - 835 0:5 a=>t S,I,T L 771.000 t=>c constant - 687 0.167 a-»g consta = 836 0.222 g=>c S,I,T L 696 0.286 a=>g constant X 837 0.300 c-»g S,I,T + NODE 70-69 780 0.143 a-»g constant = 1023 0.231 с->а I,L, os c NUCA AA sc 813 0.231 a=>g constant = 11220.400 t=>c constant + 10 0.500 с=>а - 861 0.143 ©->с constant + 1128 0.222 >t stant e excluded, (PRIMER) 963 0.182 c->t 5 - 11980.167 c->t 1,5 * 15 1.000 g=>a - 10290.250 a-»g constant + 1224 0.429 a->g солт ы! excluded, (PRIMER) 10470.429 t-»g constant 2 12630.500 a=>g R, б 75 „125 c- YR - 1056 0.167 c-> onstant 13890.143 g->a constant i 84 0.214 t-»g DIE, t 11400.300 a=>g constant - 13971.000 a R,É,I NL 8 0.143 a-»c E,K,Q,T {; 11730.167 t=>c consta - 14131.000 a->t Т,А,5,Е,Р = 108 0.400 t=>c I,T = 1185 0.200 a=>g constant - 124 1.000 a=>g M,V, L 1203 0.200 эшш constant - NODE 44-43 126 1.000 g-»a M,V,L 13980.250 g-» K, os- e AA 165 0.231 c->a A,W =- 10 0.500 a - excluded 201 0.250 t=>c constant - NODE 66-48 18 .333 g->a - excluded, 246 0,333 t=>a constant POS в АА SC 81 0.333 t-»a consta 271 0.250 g- P.A,V,T 33 0.500 t=>c V,S,F,D,A = 258 .333 с=> ,D,E,N,H ke 318 0,250 t-»c consta 84 0.214 g=>a D,E,Q - 284 0.286 a=>g N,D,S,T,E,G L 321 0.333 g-»t constant - 138 0.273 c 318 0.250 c=>t constant 327 0.167 g->a constant - 243 0.200 a=>g constant - 414 0.167 g= ok 388 0.333 t=>c con bed 290 0.125 а ЖУР L 35 0.300 c=> ‚у,5, 397 0.333 S, 297 0.200 t-»c A,V,C = 450 0.214 t-»c consta 486 0.167 g-»a S 309 0.143 t-»c constant - 498 0.333 c->t constant = 504 0.167 +->с nst =- 312 0.182 t->c P,F " 504 0.167 c=>t constan bé 522 6 t-»c constant = 3460.333 а=>с M;i ү; 507 0.333 a=>g constant А 660 0.167 с->© соп t - 498 0.333 t->c consta = 522 0.286 c->a consta 661 1.000 g - 546 0.250 t=>c constant 7 (564. 0021 am у € 662 1.000 t-»g V,C = 552 0.200 c-»t constant p 579 0.375 t-»c constant 663 0.500 a-»c V,C - 570 0.200 t-»c constant - 612 0.111 g->a constant =~ 672 0.300 t-»a constant x: 612 0.111 a-»g constant - 618 0.333 a-»g constant ^ 673 0.111 с=>а 639 0.333 t=>c constant = 702 0.200 a-»g constant ~ 764 0.400 a=>t A, Q,E,V,H,I NL 656 0.500 t-»g L,V,C - 813 0.231 g-»a constant -~ 786 0.500 g-»t V,M,A 657 1.000 a-»c L,V,C NL 952 0.500 t ‚5 810 0.333 g->a constant - 693 0.167 a-»g const - 984 0.182 ,5,7 i 837 0.300 t-»c S,I,T - 771 0.375 t-»c constant - 1045 0.333 c-»t consta 852 0.286 t-»c constant 808 0.167 t-»c constant - 11070.333 a-»c constant y 864 0.333 t-»c constant 810 0.333 a-»g constant 2 "14760,222 amg ВГА T 906 0.286 c-»t - 822 0.143 c-»t constant - 11370.231 g-»a constant 7 0.231 t-»g I,M L 885 0.286 t onst - 11400.300 g-»a consta е 940 0.250 t=>c 1,5 +). 914: 0.148 ‚В L 12151.000 a-»c constant -~ 10170.333 t-»a constant - 954 0.286 a 5 - 12660.429 t=>c constant @, 10230.231 а=>с V,I,L, - 10210.333 a->g V,I,L,M L 13380.333 t-»c constant = 10580.500 a=>t Y,F,C,L L 10420.167 t 1,5 E 460.125 g-»c A,S,T, : 11160.222 t-»a P, - 12210.200 a 156 - . 13590.286 c->t Р,А 2! 11230.250 t=>c L,S,F,I L 12450.200 a-»t constant - 12120.429 g-»a constant - 13200.143 g-»a Q,E,A - NODE 48-47 13300.167 a-»g I,V 13320.500 t-»g I,V RID She WB e 13890.143 ез constant E 13590.286 t-» ALD Ё .333 c=>t constant 0.143 a->g constant = 14160.667 g-»t I,M,V,W L 150 0.231 c=>t A,P,S E bó bud 14220.429 g-»t T,V,L,K - 59 0.167 а=>9 constant 1 0.231 a-»t A,W POS c NUCA AA Sc NODE 48-44 549 0.200 g->a consta y P FEIN n ымы m SC 603 0.143 g-»a constant . g constant = 99 Bente E,K,Q,T £i o 0211 SN р GA sce = - 147 0.154 a-»c constant - 861 0.143 c-»t constant BH oodd s 264 0.333 g->a D,E - 906 0.286 t-»c D,R ii $€ o d ыд » de 276 0.286 g-»a constant - E 429 a=>g cons Ё á nt = 393 0.231 a->g R,P - 2690.600 t-»c constan Volume 81, Number 3 1994 Albert et а E Functional Constraints and rbcL Evidence 14100.429 a-»g 14201.000 a-»g 14210.667 c->t 14250.429 а->9 NODE 47-46 0 N 12570.500 t= >g si 46-45 1362 20. 429 a NODE 66-65 POS c E,D,A,K,P,Q T,V,L,K T,V,L,K L,V,C constant constant M,L constant constant " constant L, ас ane T,M R,C CB K,R,E ,F,A constant constant Е,К,О,Т constant > = ia 2 S,T,E,G A, "e p V:H, I изын: constant D constant constant и a t L L sen 0.286 c-»t 0. 933 0.143 с-> 984 0.182 c- 990 0.500 1005 0.375 t=>g 1017 0.333 a-»c 1020 0.200 а->9 1411 0,600 a~>c NODE 65-64 с NUCA 1425 0.429 а->9 constant - constant ‚5 A,G,T - А4 x constant = A,Q,E,V,H,I L , constant = vL constant - constant - Q,E,D constant 5 F,I,L = T,A,S,E,P L AA sc A,W = constant iz N,S = constant NODE 64-63 POS rd AA 150 0. Т с= «111 .500 c-»g 474 0.250 а=>9 564 0.214 a->g 612 0.111 a->g 13800. 200 a=>g NODE 63-54 POS 10200. "200 g NODE 54-53 POS c NUCA 543 0.333 t=>c 12450. 200 a->t NODE 53-51 b o a n e n constant constant A,V constant АА constant senstant 1408 0. Am g=>a E, D,A,K,P,Q L NODE 51-49 POS с gids 162 0.429 g- 168 0.273 эй 655 0.250 t=>g 11670.200 c=>t 13450.154 a=>t A nstant ули "m K S,T,C 560 Annals of th Missouri ril Garden NODE 51-50 os c P NUCA 57 0.333 t=>g 84 0.214 c=>a NODE 53-52 POS 12390. 14100.429 a-»c NODE 63-62 POS 14220. 429 c NODE 62-61 POS c 13 тора 200 c=>t NODE 61-59 РО 290 0.125 a=>t g->a NUCA AA 108 0. бо c=>t NUCA AA 138 0. zi c=>t s c NUCA 177 0.300 c-»g ,M,T constant = Q,E,A um < ы A,V,C - constant < LIT y Q o 5 и сос pm 3 et LU с constant E,D,A,K,P,Q P,L - constant a constant - L,S, - constant -= constant T,V,L,K a о АА D,E,Q ч A,W constant - R,P I,V - constant = S,A - constant AA R,L,A - Y,F A,V - [э G.T - constant - A sc D,E L D,E, L и a 836 0.222 g->c 12780.500 g=>a 10.250 c=>t NODE 59-58 POS NUCA 153-0, к а->а 177 0.300 g->t eo ч w m 711 0.250 а->9 885 0.286 c=>t 927 0.231 a-»g 982 0.182 t=>g 11370.231 a->g 13200.143 a=>g 13800.200 g->a NODE 58-57 1-07 NODE 57-56 POS UCA 60 . c=t 12090 t=>c 2 I,T атанан АА constant РА и о N,S = N,D,S,T,E,G L , constant constant constant constant L,I constant constant M A,S,T constant Q,E,A Ы кк constant L,V,C 5,А L,M,I SPI; constant constant G,A AA constant constant constant L,V E Уу constant co constant , constant constant 12450.200 g->c ODE 56-55 POS c NUCA 117 0.500 c->g 13350. 167 c=>t NODE 61-60 POS NUCA 225 0. 2а c-»t 1345 0.154 a=>t NODE 69-68 768 0. 167 t-»c C constant * А,5,Т,С L А,8,7Т,6 L A,S,T;,C - E,D - ,D,A,K,P,Q L E,D,A,K,P,Q L AA constant т A,W = constant 3 R,P а A,Q,E,V,H,I L constant = АА constant constant = S,C т A,S,T,C AA - excluded, Volume 81, Number 3 1994 Albert et al. Functional Constraints and rbcL Evidence 711 0.375 t=>a 14140.333 a-»g NODE 68-67 P [7] min z 1122 0.400 t-»c 11760.250 g->a 11790.400 1425 0. 429 a-»g constant I,S constant T,M SIT constant L,P L + an R, constant constant constant S,F,A constant m Oo , constant G,A constant < E,D,A,K,P,Q T,A,S,E,P A V,S,F,D,A K,Q constant A,P,S constant ¥ L,M,I constant tu + o constant G,A E,Q,K constant О,Е,А T,V L,V,C,I sc L L NODE 76-75 POS c WU 28 0.500 а->9 61 1.000 a=>c 66 0.167 a= 90 50 459 0.250 t=>c 600 0.333 t->c 2 1275 0.333 a->g NODE 75-74 =>9 976 0.250 а->9 1008 0.500 а->9 АА - excluded, T < m < 4 vA constant constant AA V,S,F,D,A K,L,S we constant I,L constant constant constant constant ‚M,A constant Q , constant constant K H - X cox OX 5, constant constant pr 1032 0.429 a->c g 712 0.500 a=>c 0 1398 0.250 NODE 73-72 132 148 0.333 c=>g 144 0.333 a=>g А constant constant I,V,L constant А constant constant constant constant R,P L,S,I constant constant K, тр constan N,F constant constant ,N < М R,K constant H,Y,S,F L 562 Annals of the ү Жее Botanical Garden APPENDIX II (pp. 566). Inf branches most- parsimonious) including su stri 0.273 a=>g 1392 0.143 a=>g vex amino rdi constant constant G,D,E,N,H F A,V constant V,A constant constant K,R , constant constant constant V, 1,L,M L, constant constant S,F ` L,S constant A т ы A,S,T,C , constant E,A " constant changes 2 © 2-567; corrections i п ic EM the oof, nter mun string-based cl ойыш (опе of 165 е equally recognitions. mary statistics So the the resultant matrix of a morphic Similar to Appendix I panying c" nce ank aa contain je abou t the functional impact of specific string changes (as re- vided by total steps) is given, (ii) “CHAR” indicates the string character nur hA p the matrix АН the end of this appendix, (iii) “POS.” р tion, but, here, to the Ж. (3’) pos of a string recognition, (iv) “STR., SEQ.” indicates P e the n 5 tide the string itself (divided to show the codon positions of its component nucleotides), and (v) iM shows eac! 1+ f Hi - | p PERES string recognition. Under tł internal st op codons are indicated by * E F2 гаг *3 (f or TAA A, TAG, an Pie CA, respectively), and missing nucleotide E have metimes necessitated m ачыт: (Бу eoo acids. Again, Dayhoff et al. (1978) РАМ. 250 50 log. odds u were determined nondirectionally for cid s e string НУТ. (involving 1000 randomly ре sh t LE nging g in length from 6 to 21 base pairs) are provided below n р ы Me 2 e С. un g. © m Total Mean pork д Total Total Total Total tional nitions String recog- apomor- similar- single- recog- Re length nitions phies ities tons nitions string 6 758 58 129 (77 152 sS 7 204 43 020 23 ЗЕТИ 81559107 14 ^30 4491877600 5 2 1 1 2 100 1 4 2 1 1 2 1.000 1$. 6 p 1 1 0 1 1.000 14 5 1 1 o 1 100 15 8 1 1 o 1100 DP 2112. 393. оа 8 The 1000 strings evaluated contained the fo ie cm endi of "nucleotides," which verify their ran generati У А = 3375 У С = 3309 > С = 3349 > Т = 3297 of 193 string recognitions (including 1 ^ Н тта wily similar ities) is also PF is ion e (f racter à of rbcL nucleotides), and the oe iude following » start position informatio ar the ken “аЬ”; this indicates that adiac string rec Ө pu sa: tart position, and d part (such partial correlation has i in ош р pt nalyses; see text for further details). Ihe ptt sented in t icis, corresponding to p ert с; (500 strings in each, for a t rimer Ге exse urring we% region are shown in and but we ere ignor parsimony analysis Albert et al. Volume 81, Number 3 994 edidi Constraints and rbcL Evidence re Conocephalum E —— Andreaeobryum E Equisetu Er Lophocholea 18 EC Anthoceros {_ Psilotum E r——— Angiopteris L a سے‎ тна 72 Selaginella | — /soetes 71 — Lycopodium 3 r——— Ephedra 70 L-g Welwitschia — Gnetum | Pip 69 Nymphaea Liliu سم‎ mM pee Ceratophyllum — 66 50 Podocarpus Ls. T a Taxus E [ — Taxodium 48 ‘| [rcm Ginkgo —65 47 ee Сус cas EDU шы ла f 5 Т цв E Сш ا Ls _ Chrysolepis —- Betul - 76, relative branch — = 0.0138 NODE 42 - 41, relative branch — = 0.0178 CHARPOS. STR.,SEQ. AA-seq. S . STR., SEQ. AA-seq. 032 313b 7, tta ye t 0. 500 LDL - 044 543 6, t gct aa SAK 091 1254 6, t gc 1.000 VAN, AAN L 052 728 6, ct дса g 0:111 TAG, TSG 100 1344 6, t Mot pea 0.200 ААК, ASK, АСК, АТК, RTK PNL 068 939 7, a ttg gcc 1.000 VLA, VSA 136 465 6, t caa gt 0.250 IQV - 077 1093 8, acc caa ga 0.250 TQD, PQD 142 607 6, gat gaa 0.125 DE - 100 1344 6, t gct aa 0.200 ААК, ASK, АСК, АТК, RTK 172 980 7, ac gct gg 0.100 HAG, HSG, HTG L 113 111 6, а са ос LAA 173 1017 6, t caa gt 0.500 RQV, RDV, REV, REI, RQI, RDL L 136 65 6, t caa gt 0.250 IQV 142 607 6, gat даа 0.125 DE NODE 76 - 75, relative branch am = 0.0138 152 750 6, g atg aa 0.100 MMK, MLK, MIK CHARPOS. STR.,SEQ. AA-seq. sc 033 326 6, aa g 0.125 EEG - NODE 72 - 71, relative branch — = 0.0079 043 87 6, aac aaa 0.143 - CHARPOS. STR.,SEQ. AA-seq. 113 111 6, a gca gc 0.200 LAA - 026 266 6, ct gtt g 0.167 PVA, PVP, PVV, TVT, SVV 121 235 6, cgt tac 0.333 RY - 053 755 7, аа ада gc .200 KRA 137 728 6, ct gca g 0.167 TAG, TSG L 110: 1256 6; eg Act cc 0.500 MTP, VTP, LTP, VSP 152 750 6, g atg aa 0.100 ММК, MLK, MIK L 193 1394 6, tc aag t 0.500 IKF, IRF, IIF 183 1231 6, tgg gga 0.200 WG - NODE 71 - 70, relative branch pu = 0.0138 - 74, relative branch кр = 0.0138 CHARPOS. STR.,SEQ. AA-seq. . STR.,SEQ. AA-seq. sc 034, 333 8,26 tet gtt à 0.167 GSVT 035 345 6, c atg tt 0.200 NMF, NLF 044 543 6, gc 0.143 SAK 100 1344 6, t gct aa 0.200 AAK, ASK, ACK, ATK, RTK PNL 05% 728 6; et gta g 0.111 TAG, TSG 116 162 6, a gca gc 0.250 GAA, GWA NL 092 1259 7, at cga gt 0.167 NRV 150 724 6, gct act 0.125 AT - 142 607 6, gaa 0.1 DE 158 830 8, at act agt 0.167 NTS, NMI, NTT PNL 152 750 6, g atg aa 0.100 ММК, MLK, MIK 166 1259 7, at cga gt 0.143 NRV, N*3V - NODE 70 - 43, relative branch length - 0.0099 74 - 73, relative branch length = 0.0079 CHARPOS. STR.,SEQ. c AA-seq. POS. STR.,SEQ. e AA-seq. sc 006 88 6, aag acc 0,900 FET КЕЁ, КЧ, (DT ОТ, TP 018 152 6, aa gaa g 0.143 EEA - 007 90 6, g acc aa 0.200 ЕТК, EPK, KVS, КТК, DTK, ОТК, 031 313 6, tta gat 0.500 - TPK, PNL 043 487 6, aac aaa 0.143 NK - 114 126 6, act 0.500 MTP, VTP, LTP, VSP 092 1259 7, at cga gt 0.167 NRV - 2138530057 ect p^ 0.250 RPL 158 830 8, at act agt 0.167 NTS, NMI, NTT 73 - 72, relative branch length = 0.0079 P STR. . c “seq. 5С NODE 70 - 69, relative branch length = 0.0039 034 3 8$, t tct gtt a 0.167 GSVT - CHARPOS. STR.,SEQ. с AA-seq. 085 1147 6, cat gtt 0.143 HV - 033 326 6, aa gaa g 0.125 EEG 138 500 7, gt осе tt 0.250 RPL - 183 1231 6, tgg gga 0.200 183 1231 Gs tgg gga 0.200 WG - NODE 69 - 66, relative branch length = 0.0138 NODE 72 - 42, relative branch length = 0.0079 CHARPOS. STR.,SEQ. с AA-seq. CHARPOS. STR.,SEQ. с AA-seq. sc 013 51415 а gtt cc 0.333 -GVP 033 326 6, ал даа g 0.125 ЕЕС - 036 388 6, сїа сда 0.200 LR, LP 115 155 7, aa gca gg 0.167 EAG - 043 487 6, aac aaa 0.143 NK A131 —128 Є: " Є = 0.167 TAG, TSG L 056. 783 б6,-а gtt cc 0.250 GVP, GMP, САР 187 1284 6, а са‹ 0.250 VOA, VEA, VKA L 124 273 6, t ggg ga 0.167 AGE, PGE, VGE, TGE 177 1182 6, t ggg ga 0.333 FGD 193 1394 6, tc aag t 0.500 IKF, IRF, IIF NODE 66 - 65, relative branch length = 0.007 CHARPOS. STR. «Жаз. e AA-seq. 010 129 12. gta act cct ca 0.200 RVTPQ, RMTPQ, RLTPQ, RVSPQ 076 1067 6, gac c 0.200 KFR, EDR 132 395 es ES ipte e Be 143 ALR, ASR, Т?? 150 et 0.125 AT әш jo sjeuuv v9S uapsed jeoiuejog unossiN 65 - 51, relative branch CHARPOS. STR.,SEQ. 085 1147 6, cat gtt 152 750 6, g atg aa 166 1259 7, at cga gt length = 0.0059 c АА-вед. 0.143 HV 0.100 ММК, MLK, 0.143 NRV, N*V - 50, relative branch ego = 0.0059 :, SEQ. CHAR “seq. 124 273 ggg ga 0. 167 AGE, PGE, 130 386 6, ct cta c 0.167 ALR, ALP 142 607 gat gaa 0.125 DE NODE 50 - 49, relative branch — = 0.0118 CHAR . STR.,SEQ. AA-seq. 013 141 6, a gtt cc 0. G 035 345 6, c atg tt 0.200 NMF, N 047 635 6, t = t А 0.333 МАИ 051 684 6, 0.250 AQA, SQA, 053 735 7, да me Fy Ей 0.200 КАА 143 63915, c tgg aga gat cgt tt0.500 RWRDRF - 48, relative branch length = 0.0158 А ,SEQ. C АА-зед. 050 663 6, t дса g 0.500 CAE, VAE, 052 2728 6, ct gca g 0.111 TAG, TSG 085 1147 6, cat gtt 0.143 115 155 7, aa дса gg 0.167 EAG 119 9 6, ac 0.200 TT 130 386 6, ct cta 0.167 ALR, ALP 137 728 6, ct дса 0.167 TAG, TSG 166 1259 7, at cga gt 0.143 NRV, N*V NODE CHAR TR.,SEQ. 031 313 "y tta gat 038 412 6, 2 relative branch — - e 0138 AÀ-seq. 0.100 HAG, HSG, 48 - 47, relative branch — = 0.0079 . STR.,SEQ. 124 273 6, t ggg ga 0. 250 FAR, CAR, 0.167 AGE, PGE, AQT, CAE HTG CAK VGE, 506 TGE zu кее ттеп tator 8 zz э Ж oe NODE 47 - 46, 0 092 1259 132 395 1821207b NODE 65 172 980 Е 64 - os. NOD! CHARPOS 061 856 106 1418 191 1355 NODE 55 CHARPOS. 054 766 152 750 NODE 54 CHARPOS. 0217 198 NODE 54 - CHARPOS. 079 1110 STR.,SEQ. 14, a qat tac aga tta , 7, 6, аа сс at сда gt g atg aa - E relative branch - 64, relative branch STR.,SEQ. 7, ac - 54, relative branch STR.,SEQ. 7, 6, ttt gcc a g atg aa - 52, relative branch STR.,SEQ. 6, g aca ac 53, relative branch Im 073 1135 6, 7, 176 1138 6, NODE 6 CHARPOS. 192 1369 е ганот e gge ggt 63, relative branch SEQ gct gct t relative branch Tenet h =o seq. RDYRL, NLD NLF AA-s 0. 200KDYRL, 0.250 SLD, 0.2 ММЕ, KDYKL, KDYTI, length = 0.0059 с АА-вед. 0.333 GVP 0.143 ALR, ASR, Т?? 0.500 GG len = 0. pete 0.250 ALR, ASC 0.100 HAG, HSG, HTG length = 0.0059 AA-seq. 0.200 DN 0.500 DTL, DVL, DTV, 0.500 SPE, SPD, SAE, ILC, SLE DKL qs - 0.0039 AA-seq. 0. 250 FAR, CAR, САК 0.100 MMK, MLK, MIK length = 0.0020 с AA-seq. 0.250 length = 0.0059 с AÀ-seq. 0.500 sG 1.000 SLP, STP, SLA, SMP 0.500 GG length = 0.0020 с АА-вед. 167 ААС length = 0.007 c АА-вед. 0.200 YTPE, FTPD, ҮТРО, 0.250 FAR, CAR, CAK 0.167 NRV 0.167 AGE, YTPD PGE, VGE, TGE sc KEYKL PNL L 7661 £ JOQUINN ‘8 ӘШпол eouepi^3 729 pue =4шедѕиод үеиоцоипч ‘ye 1e u4eqiv 696 - pel relative branch length = 0.0039 CRARFOS . STR.,SEQ. e AA-seq. iH rs E ы аг ge 0.333 REA, EEA 145 11 6, cta c 0.333 SLP, STP, SLA, SMP NODE 61 - 56, relative branch length - 0.0039 CHARPOS. STR.,SEQ. с АА-вед. 015 146 8, са cct gag 0.100 PPE, PAE, PSE 145 684 6, a ca 0.333 AQA, SQA, 506, AQT DE 60 - 57, relative branch length = 0.0039 POS. STR.,SEQ. с AA-seq. 018 152 6, aa gaa g 0.143 EEA 090 1245 7, g ggt gcc 0.500 PGA, PGG, PVA, PGR, NODE 60 - 59, relative branch — = 0.0099 CHARPOS. STR.,SEQ. AA- 076 1067 6, aa gac c 0.200 KFR, EDR 100 1344 6, t GCT aa 0.200 AAK, ASK, ACK, ATK, 13772 728 `6, дда .167 TAG, TSG 152 750 6, g atg aa 0.100 MMK, MLK, MIK 177 1182 6, t ggg ga 0.333 e “PNL” t Beers in proof: to P. 564, o 7 10-43," right hand column; PRA RTK NODE 59 - 58, S CHARPOS. STR.,SEQ. NL 139. 3501 6, c ccc ct N 146 686 9, ag gct gaa a NODE 69 - 68, sc CHARPOS. STR.,SEQ. PNL 023.269 6; Бб сс Т; 0297543 6, t gct aa 137 728 6, ct gca g 166 1259 7, at с gt sc - NODE 68 - 67, relative branch NL CHARPOS. STR.,SEQ. 025 252 6, с tac: ga 0340333 2% tot gtt.a 041 635 6, tg-cgt t NL 076 1067 6, aa gac c PNL 092 1259 7, at cga gt 116 162 6, a gca gc L 130386 6,:6ct cta c - 150 724 6, gct act 170 966 6, t ggg ga 192 1369 7, gct gct t . 566, under "NODE relative branch relative branch fourth line from bottom, delete comma after d— = 0.0039 AA-seq. 19 c AA-seq. “ТРК” ОСЕТ, TSG CYG, Ф EDR GWA ALP and 68-67," bottom line, right hand column, should read PVV, CYE, QTET TVT, CYN, SVV, CYH PVA sc PNL uapsed jeoiuejog unossiN 99S eu Jo sjeuuy 7661 € JequinN ‘Lg ӘШПоЛ (nucl./string 76 819766676] 6767886666 6776666616 6768666666 6666676666 6677667866 6667667767 6678668677 6676668767 6776676676 6667666 766 787 666676666? 6686676666 6667666766 6616696766 6666766867 7676666666 емне адау 976 ems ات په سم ا‎ IESUS GIN GRO MEUS چ‎ Sig SDSS ue Io FRENTE ardt ca Did А ae i TO, Don چات‎ pal АЫ DER ES coe PEN ые SE LC ESSE эс» RRS کس‎ Imita/string P 35142282 до аре ^oi Viro 9443422241 1212213138 9312242122 E 4131425322 Loh git rasa 2294871 311 321 inne 4837215473 bora de 1382141183 6311281192 2328126467 ume jen 621 зм 2 -94 --0145-528 ~6745-0-0- -52042-—-4 70--7---{ O- 48--06--00 -8-047--8- 40-00~---. 3 3-- ~- ^ 9--10--918 98--- 0 04--7-7--2 -4099-70-- -2--70-9-1 -54-801—-- 0-1-9-7- String start 0000000000 0000000000 0 0000000000 0000000000 0011111111 1111111111 1111111111 111111] 000 000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0011111111 1112111111 111 position 00 OI umm peony UE Bote ED 8899999999 9910000011 1111122222 2222222333 3333444 000 000 0011111112 2222233333 3344444555 5666666677 7777778: 88999999 9900111111 2222222333 333 01 4578899902 3444466577 9112566788 1123489112 2384673556 8256688344 5600011356 7834669912 2233422244 5566689344 7999011 222 467 8912566891 4457800368 9911268004 6038889922 2555661333 5678013566 7817038889 0033558244 569 48 9441480643 60 phe 34 8257267803 3363582272 5673735253 4856835319 6925703934 2950473502 3711735545 4936663814 8044789 335 146 7716525399 5943049626 5529156017 1790464524 9099071022 i iode 8079982495 7715144512 594 = 5 а >л; ————- --—---—— oe oa ---ab----- --ab--- ab- ~- --------——- -=== ART ---------- --ab----ab ---------- ---------- ab-------- -— Character # XX 0000000000 0000000000 0000 0 000 00000 0000000001 1111111 XXX 111 1111111111 1111111111 1111111111 1111111111 1111111111 1 пишип 1211211113 nunn 111 хх 0000000001 1111111112 Hen HR ttis Tees бишен mime DEP 9999999990 0000000 ххх 001 1111111112 pee sini 4444444445 5555555556 6666666667 7777777778 Border ووو‎ 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 123456789 4567890 1234567890 1234567890 1234567 XXX 890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 123 Conocephalum |17?) ??00000000 0000000100 000000100! 1 000! 00000 00000 1000000000 02??? 1???) ??0 0011110000 0000000000 1000011101 0100000001 0001010100 0000010000 0110000000 1000000010 01? Lophocolea [??] ?011000000 ies DHE оин. Meca soam ЕИ ошооо 01000000 Nasr 0000000001 BN [227] 220 oen 1000000000 0000000101 ??20000001 0100111100 0000010000 0000000000 0010010000 01? Anthoceroz [77] 772000100 000100 0000110000 1101100000 0000772227 7100000000 000 0202000000 0000100000 0000000000 022222? [277] 200 0001100000 1000010000 000001010? 0720070000 0100000000 0001000000 0070000000 0110000100 01? Andreaeobryum [1?] 7700011000 0000001007 1000110000 сано 011001000 0110000000 pee 0010000000 0000100000 1000000100 2272722 (???] 220 Е 0000000000 0000011100 0100000001 0000000100 0000010000 0110010000 0010000000 07? Ophioglosaum [10] 1000000000 0000000000 0001010100 00101000 ооо 0010000000 0000000100 0000001000 0000000100 0100000001 0011??? [100] 000 0011000000 1001000000 0100011000 0100000000 0000000000 0000010001 0000000000 0000001000 010 Psilotum (10) 0000000000 0000101000 0000019000 0001100000 0111001000 0110000010 0000000000 0 000 0101100000 0100100000 0000??? [000] 000 1011100010 1001000000 0101100100 0000000000 0100000100 0000000000 0001001000 0010000000 010 Isoetes [20] 1100011000 0000100000 0000000000 0001100000 1010001000 0000000000 0000001000 RS 0000001000 0001000000 SESS 1000) 010 0001100010 1000000000 0100000100 0100000000 0110000100 0000000001 0000000000 0000000000 011 Lycopodium [10] 0000011100 0000000000 0001010000 0000000000 0010001000 0000100000 0100000000 0000000000 0000000000 0000000000 0000??? 1010) 000 0001000000 0100100100 0000000100 0100001000 0000010100 0000000000 0000000000 0000001000 001 inani [10] 1001000000 0000001100 0100010000 0010000000 0001001000 0110001000 0000110000 1000000000 0000000000 0101000000 000072? (001] 000 0001000000 1000000000 0000001000 0000001001 0100000001 реф 0000000000 0000001000 010 Equise 111) 1000000000 0000000100 0100010000 1011001000 0011001000 0110000000 0000000000 0100000000 0000000000 0000000001 0000??? [100] 000 0011110000 0000000001 0000001100 0000000001 0000000100 0000010000 0000000000 0000000100 010 Selaginella [00] 1100100000 0000 n 0000000001 0000100000 0111101000 1100000000 ا‎ 0000100 0010000000 010001100 [000] 001 0000100000 1000001100 0011000000 0000000101 0100000000 0100100010 1110000000 0000000000 001 Botrychius © [10] pem 000000 сре 0 0000001000 0000100000 0010001000 0010000111 0000000100 % epus 3000161009 SERT ue [100] 000 0011000000 1000000000 ap 0100000000 0000000000 0000000010 0000000000 0000001000 010 Taxus [??] 1200002000 0000000000 0000000000 1070010100 0010000001 1110010000 10 er 00 0000010000 0000000000 0000000000 2272222 [000] 000 0000010010 1000000000 0100001101 0110000001 0100000000 0000000000 0100001000 0010000000 ттт Taxodius 110) 1000000001 тие» Batches mod саха TED 0000000000 0000010000 0000000000 0000000000 277772? [000] 000 0000110000 1010000000 оа 0110000001 0100000000 0000000000 0100001000 0010010001 07? Podocarpus [00] 1100001001 0100000 0000000 1010010000 1000000000 0000010000 100010000 0000000 0000000 [000) 000 0000100000 1000000001 000 0010010000 0100000000 0000010000 0000001000 0010000000 01? Ginkgo [??] 1000000001 0000101000 0000000000 0001010000 0000000101 0101010000 0000000000 0000010000 0000000000 0070000000 ? [000] 000 0000000010 1001000000 0100000001 0010000001 0100000000 0000001000 0000001000 002?000000 ?70 т > Cycas im eee 0000001000 0001000000 0011110010 0010000011 1101000000 0000000000 0000001000 0000000000 0100000000 0 Ge [000] 000 0010000010 1001100000 0100001000 0010000001 0000000000 0000000000 0000001000 0010000000 010 c 5 Stangeria pa 1100000000 0100101000 0001000000 0011100000 0010000011 0101010000 0000000000 0000000000 0000000000 0100000000 0000000 (10?] 000 0000000010 1001000000 0000001000 0010000001 0000000000 0000000000 0000001100 0110000000 010 2 g tania 71] 1000000001 0100001 = 0000000000 0011110000 0010010011 1001010000 0000000000 1000000000 CERA 0100000000 0000000 170?) 000 0000000010 1001000000 0000001000 0010000000 0000000000 0000000000 0000001000 0110100000 010 Q 3 Ephedra ne 1000000001 0001000000 0000001010 0011100000 0011001000 0010000000 0000000000 0000000000 0101000000 0000000000 0000001 (200) 000 0000101000 1000000010 0100001100 0100000000 0000000010 0000010000 0000000000 0010000000 011 o Ф Welwitschia (00] 1000000000 0000000000 0010101010 0010101000 0010000001 000001000 000 0000 000 du 0 0100000000 pence [000] 000 0000010000 1000000001 0000001000 0110000001 0000000000 0000010001 0000000000 0010000010 001 a сЕ Cnet us [72] 1000000000 0000000000 0000101010 0000100000 0010000000 0100000000 е dt 10 0000011000 0000100000 0100000000 00000 (000] 000 0000110010 0000000001 0000001000 0100000001 0000000090 0000010001 0000000010 0000001000 772 = Chloranthus |??) 1000000000 0010100000 1000000000 0011110000 0000001000 0101010000 1000000000 0000010000 0000000000 0000000000 im [000] 010 0000100000 1001000000 0100000000 0000000001 0000000000 0000000000 0000001000 0010000000 110 Lon es Piper {??] 1001000000 0010001100 1000000000 аса hi nee tM Ris жнын меде CEPS о обы Se 0000010 [700] 000 0000100100 1001000000 е 0100000000 0000000000 0010000000 0000001000 0010000000 010 o Orinys {27} 1000000001 0010000000 1000010000 0011110000 0000001000 0001010000 10000 0000010000 00000! 0000010 [000] 000 0000100000 1001000000 0100000000 0000100001 0100000000 1000010100 0100001000 0010000000 110 à Calycanthus (??} dee 00001 0010000000 0000000000 0011110000 0000001000 0100010000 1000000000 0000010000 0000000000 0000000000 0000010 [700] 000 0000100000 1001000000 0100000000 1001000001 0000000000 0000000100 0100001000 0010000000 110 = Eupomatia psi 0000001 0010000000 0000000000 0001110001 0000001000 0101010000 1000000000 0010010010 0000000000 0000000000 0000010 [?00] 000 0000100000 1001000001 0100000000 0000000011 0100000100 0000000100 0100011000 0010000000 110 e Magnolia 1001 * ннер 0010100000 0000000000 0011110000 0000001000 0101010000 1000000000 0010010010 0000000000 0010000000 0000010 1000) 000 0000100000 1001000000 0100000000 1000000001 0100000000 1000010000 0000011000 0010000000 110 = Persea 100) (iicet 0010100000 pret SUI bs phe ig Bi рәме, EE кндн 0000010000 0000000010 со 1000) 000 0000100000 рк 0100000000 dei cen 0000000000 0000000100 0100001000 0011000000 000 a frochodendron 1010100000 0 009011 0000000001 0100000100 1700) 000 0010100000 1000000000 0100000000 0000000000 0000000100 0000101000 0010001000 010 X Ceratophyllum [??) 000 0001 0 1110001 0000001000 0001010000 0000000000 0001010000 0000100000 0000000000 0000000 [000] 000 0000100000 1000000001 0000000000 0000100001 0100000000 0000010000 0000001000 0010000000 010 > Nymphaea 9 он ree deme wi 0000001000 0000010000 0000000000 0000010000 0000077227 nmn 2222222 [000) 000 0100100000 1001000000 0100000000 0100000001 0010000000 0000000000 0000001001 0070000000 277 a Liliw 127 он 0010000000 0000000000 0011110000 0000001000 0000010000 1000000001 0000010000 0000100000 0 00000 0100000 [200] 000 0000000000 1001000000 0100000000 0100000001 0100000000 0000010000 0100001000 0010000000 000 E Platanus (2 "i 000000001 н center 0011100000 0000001000 0001010000 0000000000 0000010000 0000000000 91009000 БЕ [200] 000 0000100000 1000000000 0100000000 0000100000 1000000000 0000000100 0100101000 0010000000 100 S Caltha {22} 1000000011 000100 0 0011100001 0000001000 0001000000 0000000000 0000010000 0001000001 0100000100 000000! (000] 100 0000100000 1000000000 0100000000 0000001001 0000000100 0000000100 0000101000 0010000000 000 r- Dillenia [27] 1000000001 0010000100 0000000000 0011100000 0000001000 0101010000 0000000000 1000000 1700) 000 0000100000 1000000000 0100000000 0000011001 0100000000 0000000100 0100000000 0010000000 000 m Chrysolepis [21] 1000000001 B 0010000000 0001100000 0000001000 0101010000 0001000000 00000000090 ам 0165000107 ө 0000000 [700) 000 0000100000 1000000000 0100001010 0000010001 0000000000 0000000000 0000000000 0010000000 000 < Betula n Meth 0010000000 0000000000 0011100000 0010001000 0101010000 0000000000 0000000000 0000000000 0100000101 0000000 [000] 000 0000100000 1000000000 0100001010 0000010001 0100000000 0000000100 0100100000 0010000000 000 a Casuarina zd 100 "e 0010110000 0000000000 0011100000 0001001000 0101010000 0000000000 0000000000 0000100000 0100000101 0000000 [700] 000 0000100000 1000000000 0100001000 0100000001 0100000100 0000000100 0100100000 0010000000 000 Ф Hamanelis 22) quet 0010000100 0000000000 0011100000 0001001000 0101010000 0000000000 0000010000 0000000001 0100000100 0000000 [700] 000 0000100000 1000000000 0100000000 0100000001 0010000000 0000000100 0100001000 0010000000 000 3 [51] o ч STATISTICAL SUMMARY OF SOME OF THE ACTIVITIES IN THE MISSOURI BOTANICAL GARDEN HERBARIUM, 1993 Vascular Bryophyte Total Acquisition of Specimens Staff Collections (includes bryophytes) 27,402 = 27,402 Purcha: 16 0 Exchange 53,377 2,728 56,105 Gifts 9.712 4,781 14,493 Total acquisitions 90,507 7,509 98,016 Mountings Newly mounted 142,737 21,558 164,295 Mounted when received 15,994 0 15,994 Total specimens filed 158/31 21,558 180,289 Repairs Specimens repaired 36,885 n/a 36,885 Tu d aar ed 4,858 n/a 4,858 Total г 41,743 {0 41,743 P sent On exchange 31,807 7,433 39,240 s gifts 23,535 3,656 27,191 Total 55,342 11,089 66,431 Loans sen Total transactions 741 32 773 Total spec 30,309 2,426 32,735 o U.S. institutions Tr tioi 397 23 420 Specimens 15,576 931 16,507 To чк оз institutions sactio! 433 9 442 EE 14,733 1,495 16,228 To student investigators Transactions 65 8 13 Specimens 7,670 485 8,155 To professional investigators Transactions 676 24 700 pe 22,639 1,941 24,580 Loans receive: ransactions 446 14 460 Specimens 32,097 2,500 34,597 From U.S. From abroad Total Visitors 412 131 ŠA 1 December 1993 the total number of mounted, accessioned specimens in the herbarium was 4,292,346 (4, 050, 971 vascular plants and 241,469 bryophytes).—David Brunner, Curator of the Herbarium. 568 NOTICE THE RUPERT BARNEBY AWARD The New York Botanical Garden is pleased to announce that Dr. Andr gentina, orn Anisophylla in South America, and Aem for the floras of Ar- gentina and Parag T w e Yo s nd Garden also invites planning to come to The New Y den to study the rich collection of Leguminosae. ? gem ld амаги shou submit their curriculum vitae, û a letter ее he nd h ы e pr oject for the ai at NYBG will benefit or research: Trave ned between Jan- tor, Institute of Systematic spp The New York Botanical Garden, Bronx, NY 10458-5126, USA, to The New tanical Garden, to Dr. Forero York Bo- Volume 81, Number 3, pp. хе 570 of the ANNALS OF THE MISSOURI BOTANICAL GARDEN as published on 1994. п August 1, Two Important New Titles from the Missouri Botanical Garden Moss Flora of Central Americ | Part 1. ен | Bruce Allen with contributions from Н. Crum, К. A. Pursell, W. D. Reese, №. Salazar Allen Central America possesses one of the world’s richest moss floras, with an estimated 871 species in an area only one-half that of Colombia, which has about as many species. Previous to this publication the onl flora for = Central American country was E. B. Bartram’s now nearly half-century old т The Moss Flora of Central America is based on historic and extensive recently collected material from the area, with — мече s ecran: and oM pod its distribution in Central America mapped on an indiv d, and all taxa are keyed. Extensive synonymy is os e types cited for all п; names. The Flo will appear in four parts during the next few years. t 1 constitutes mm die dum in culte 92 from the Missouri Botanical Garden, volume 49, e pages. April 1994. Index of Mosses Marshall R. Crosby & Robert E. Magill Index of Mosses is a guide to the names published for mosses, including all new taxa from the rank of genus and below. Entries ае full references to places of publication, and, depending on the nature of the oe fisting of basionyms, replaced names, and types. The Index of Mosses, 1963-1989, lists x and fully s about 8. Us place an order, please indicate method of payment below. Checks or money orders should be in s. funds, payable to Missouri Botanical veins through a U.S. bank; U.S. shipments: add $2.00 for = book and $.75 for each additional book; non-U.S. рөн Hs $3.00 for one book, and ‘or each additional book. Orders should be рн a $1.00 fee will be added to orders requiring invoices. No s Shipments are made until payment is received. Mail form, with payment, to: Petpet Eleven, Missouri Botanical Garden Please send copy(ies) of Monograph No. 49 St. Louis, MO 63166-0299, U.S.A. Please send copy(ies) of Monograph No. 42 Please send copy(ies) of Monograph No. 50 О Check/ money order enclosed Send books to: š Send invoice ($1.00 fee will be added to total) Charge card number (Mastercard/ Visa) Name Expiration date Address Name as it appears on card Telephone number (daytime) Postal Code Country PRICES ARE SUBJECT TO CHANGE WITHOUT NOTICE 81(3) CONTENTS Origin and Relationships of the Major Plant Groups Dedication and Introduction P. Mick Richardson 403 Progress and Prospects in Reconstructing Plant Phylogeny Michael J. Donoghue. 405 Integration of Morphological and Ribosomal RNA Data on the еса of Angiosperms James A. Doyle, Michael J. Donoghue & Elizabeth A. Zimmer ..— 419 Phylogenetic SERIA of the “Green Algae" and “Bryophytes” Brent D. Mish- ler, Louise A. Lewis, Mark A. Buchheim, Karen S. Renzaglia, David J. Garbary, х Е ino Frederick W. Zechman, Thomas S. Kantz & Russell L. A Куше. of Seed Plant sued Kevin C. Nixon, William L. ds Dennis Stevenson & Else Marie Frii 484 Evid f ] Plant Р} Victor A. jus Anders Backlund, Kare Bremer, Mark W. Chase, Оне: R. Manhart, Brent D. Mishler & Kevin C. Nixon 534 Statistical Summary of Some of the Activities in the Missouri Botanical Garden Herbarium, 1993 David Brunner REDS PNE The Rupert Barneby Award 569 Cover illustration. Petrea sulphurea M. J. Jansen-Jacobs, by Phyllis Bick. ‘ | a . Annals of the — Missouri Botanical Poci umber Volume 84, Number 4 Annals of th Fall 1994 Missouri Botanical Garden The Annals, published quarterly, contains papers, eid in systematic botany, con- tributed from the Missouri Botanical Garden, St. ou niic originating outside the Garden will also be accepted. Authors should write ihe Manas g Editor for information concerning arrangements for publishing in the ANNALS. ‘iene to Authors are printed in the back of the last issue of each volume. Editorial Committee Henk van der Werff Editor, Missouri Botanical Garden Amy Scheuler McPherson Managing Editor, Missouri Botanical Garden Diana Gunter Editorial Assistant, Missouri Botanical Garden Robin Bruce Magdalen Lampe Publications Staff Ihsan A. Al-Shehbaz Missouri Botanical Garden Gerrit Davidse Missouri Botanical Garden Roy E. Gereau Missouri Botanical Garden Peter Goldblatt Missouri Botanical Garden Gordon McPherson Missouri Botanical Garden P. Mick Richardson Missouri Botanical Garden For nancies epos contact Department leven, Р.О. Box 299, St. Louis, MO 63166-0299. Subscription price is $100 per volume U.S., $105 Canada and Mexico, $125 all other countries. Four issues pr volume. The journal Wa is included in the ption price of the ANNALS е ANNALS OF THE issouR! BOTANICAL GARDEN оз 0026-6493) is qune eb spe Missouri Bo : > A The mission of the Missouri Botanical Garden Mee T 631 discover and share knowledge about plants and Eleven E o. eA х i environment, in order to preserve and enrich life. ИЖ © Missouri Botanical Garden 1994 З i 7 Thi р f ANCI/NICR 720 AR 1 5 anence of ре E Volume 81 Number 4 1994 Botanical Garden MISSOURI BOTANICAL DEC 1 5 1994 REVISION OF HILLIA Charlotte M. Taylor? GARDEN LIBRARY (RUBIACEAE) ABSTRACT Hillia Jacq. is a genus of 24 usually h pecies th inguished | and comose on corolla characters, five subgene recognized he ine nus Hillia p dien cC enus Andin - M. Taylor (one до ubgenus Too c. M. T. кча (five rte subgenus — aylor (seven species), and pin avnia sid ted) С. М. Ta ylor (five. speci ies). Keys to all species of the genus аге provided, and those of the E Hillia Ja Jacq. is a genus of 24 species of usually epiphvti wet Neotropics. These species are easily separated ir seeds with a tuft cculent cous ي‎ е pe are interpetiol aris as sons solitary cylin- drical wood : orm, green to nun or edis and Em or red and tubul k es of Hillia can be co = those of са" Ruiz & Рау., which аге succulent пя with a = geographic range, similar apsules, and salverform white corollas si r to doe of some Senn 4 Hillia (Taylor, 1992). ually three or more flowers per inflorescence. rmark (1974) also distin- ished Hillia by the lack of marginal 5 on e ; however, f Hillia have at least a thin marginal wing, whi ем аге мао in this regard. These genera lly also be separated by their stigmas, which whe hed, hese arian ant n: Move thank t CORD, CR, CHS CUNE. the following institutions. available: A, AAU, CAS, COL, Rap F, G, GB, HAC, " HAJB, Hit HUA, “185 ) MO, “NY, Р, ie, UPR, _UPRRP, _US, Para Plants d Costa Ra inis m ‘the ейн ual to the Foundation (BSR 8r. 0068) and the Dee Scholarship Fund « of the Field ү ктун а he Department of Bio tried of Pd " Sc Rl de Río Piedras, the Man uo» Garden, the National Science aa um of al History for sport for trav W comments, notably ъй Вигрег, - D. Stevens, is Molano, p. Hoc н Missouri dd Garden, P.O. Bo Bera J. Dwyer, L. Andersson, „М. H. and E. Grma I eris P. Berry, ry, " and 1. Pip for T with analyses, 299, St. Louis, Missouri 63166-0299, U.S.A Grayum, W. Haber, ГА Hammel, R. E. Gereau, Z. Mercado, T. Zanoni, B. catae records and fieldwork. ANN. Missouri Bor. Garp. 81: 571-609. 1994. 572 Annals of the Missouri Botanical Garden itate to bilobed or linear, less densely papillose, and Hillia was published : parasitica Jacq., the most decor ано oe рош кнын r о the Antilles. The па: commemorates Sir John u: eles works” (Sims, 1804), and it has been suggested iin the specific uaa was intended as much to describe Hill's nical work as the plant's habit. Hillia has not en Mone comprehensively since de pud rd ized thre . Reviews of its char: ide ا‎ ia tal fos were pre ee a аа 73), Schu- 9, 1891), and Steyermark ipie 1972). re th M the Antilles through Peru, Bolivia, and southern Brazil. Saldanha Vell. was first combined with Hillia ну Spann (1 989) Whon inc chided Ppecjes s with сооло} in this genus. Howe lu variation in 4 been recognized by ^ all authors, particula rly those am America where representatives str чыч d were the present treatment I r nize five subgenera subgenus Hillia е species), subgenus Andin C. M. Ta pec subgenus e M. Taylor (on ies), С. М. Taylor бте species), — enus Kam en species), an genu s). Species of ipee Hillia are dois through most of the e of the genus, with the exception of саг Ameria and Mexico. ilia таш © i£ 1 tral South America. In contrast, species of sub- genus CUM are i imarily fo und in the Great- er Antilles and C i th g poini yete the northwestern coast of South America. Species of subgenus Illustres are found in South w ending orth to Costa Rica. Several species of he iani enis Amer li uggesting that show d 8 they are uncommon or that the epiphytic fl ora of with wens descr range o s subgenus to south sages cese 1991). ern Mexico and MORPHOLOGY The morphology o nus Ravnia has been described Lipid Clay 1997, The ko sion prese including PS of subgenus Кабана, HABIT, STEMS, AND VESTITURE Plants of Hillia are erect or sometimes s clam- frequently rooted in soil, although in Puerto Rico this species is found rooted only on wet mountain the soil is saturated throughout the year oot growth of most are flushed with pink in ini of H. ru Standley has red color a ed or circumferential ridged, р n young twigs. Plants of all species are glabrous оов. c ues are found in all parts, are particularly evident in the stipules, ovaries, abaxial surface of the leaf veins. but and LEAVES The leaves are iso ous (ог апіѕорһу DU some plants of Hillia riora (Oerst.) C. M. d : and petiol ubsessil Н. costanensis А erm.), although the blades ie са attenuate p : base and the petiole difficult to delimit prec шз The blades аге suc , entire, and gen elliptic to oblanceolate or lanceolate. The p E palmate in H. ulei K. Schum. and I1. à C. M. Taylor. ith all of tł 1 rising proximally Volume 81, Number 4 994 Taylor Revision of Hillia to the middle of the blade. The veins are usually pane e, or r the sec: condar ry veins may be prominulous Hillia, and the me order venation is charac c- teristically prominulous as well in H. erm. I the «айу. lacking, agar AERA Re or pocket ac of some species. They are strongly and sweetly esi at night, and are most likely pollinated sg m moths on other шнш. урш The а green sometimes flushed ith yellow, pink, orange, odor or pollinatio ne даре for ba pollination. The tubular ond rophylla Standl. The stipules are interpetiolar, ligulate to oblan- ceolate, d hey a individuals of some species de persist epa an- thesis. Such persistent stipules have been called “bracts” by some authors (e. £-, Steyermark, 1972, 1974). These Structures are here called “stipules subtending flower: FLOWERS The flowers are terminal and one infre- ently three in Н. Жеш К. Schum and Н. saldanhae К. Sch or usually three in H. Meme Whey may p ‘sessile or borne ор stout racts are red and odorless and apparently pollinated by mb; us to thickly carnose; those of all species are glabrous through- out. There are з usually four t 0 si s man Hino Meo The number EE lobes frequently уг e үйр а вре puis to the right, the less common condition in the EE than leftward orientation (Rob- brecht, 8). The stamens are inserted near the mouth of the corolla tube, or near the middle of the tube in subgenus Tetrandrae, subgen species of subgenus ual i шо lobes, шас m may be m dedos nba roma s. The filaments are usua lly sh an nd the anthers ie [ашу included. Ape ане with he filaments attached near ‘the eir SR The pollen is beris x 3- а киле 1988) or & P 52991). are rare and resemble the fei in shape and texture, but are much smaller. These bracts have be ed “‘bracteoles”’ SU 1972, 1974 o stylous and apparently Stand Gas The duration P т flower is unknown. : calyx limb Lou be completely pics into our % six lobes, or as many as nine or ten in some ur of subgenus кй. һауе a well-deve oed and ower bud. 4 esc rym ا‎ nee and often remains ad th 1 shape of 9 calyx lobes fr: чайра Vary widely within a specie: © corollas may be aie with relatively long i. tubes (subg. Hillia, subg. An vg ae subg. Tetrandra е), narrowly be lllustres, subg. RN d or бое о in- x (subg. Ravnia). The salverform ponet have x er tubes and spreading lobes, and are e white T Sometimes flushed with pink or red in individuals wt 12 T syle is straight and alee and is sur- gon at the ba ue у owa umen di M. iei eal positioned at or г just above the йе, ог їп у торсон Teiandrae sd subs p s ANANA ein ппеаг The style d glabrous; the stigmas are pura pa- pillose. The inferior ovary is turbinate to cylindrical, with axile aito that bear numerous ascend- ingly ovules. The ovary is sometimes imbricated narrowed to stipitate at the base FRUITS The mature capsules are cylindrical, pale green oody, and to brown, w У sieut along the naddie of the septum: 1 The with th e fibrous. exocarp and mesocarp separ ating from the papery endocarp. The capsules of most species terminate in a beak 1-10 mm long that is 574 Annals of the Missouri Botanical Garden composed of the enlarged and hardened disk and E ometimes also an a n the flower, but elongates markedly during fruit din lopment. The narrowly rhombic seeds are viec flat- ed with a small e o rhombi ot tened with lliptic to rh 3 mm long, and are partially edged ee a qe membranaceous to papery margin or wing. А tuft e seeds for ne 1 1 u st construction, releasing nu- 1 2 2:2 ۴۳ Í. obs.). CLADISTIC ANALYSIS The classification presented here and for Cos- mibuena (Taylor, 1992) is based on cladistic anal- mea Martinez; (2) the relationship of Hillia to the of Cosmibuena; and (4) the relationships among the species of Hillia. INGROUPS a a analyzed were the four species of Cos- mibuena recognized by Taylor мина м Balmea stormae Martinez of northern M е the twenty-four Pese of Hillia Epp inia present treatm OUTGROUPS our neotropical genera often classified in the uena in stipule, infloresc al, fruit, and he morphology; Е erdinandusa Pohl species = have of Cosmibuena and capsules similar to those of all diria — genera; Cinchona ч) has salve form corollas and capsules similar to those of al iens epiphytic genera, though vith acropeal de- to pink соба with буе to six reflexed we ve pri sine or imbricate apres and w sules with marginally winged s yo vi of Cane from the Canta " Щщ СО Persson, 1991); this genus і is included i in the pres- ent analysis based on the similarity of its corollas assified in other tribes, cytophyllum yotideae), were chosen as outgroups based о on пш еїг e ities outside of the yo 988), fruit, and their меен recent morphological systematic review. a is here regarded in th ele e wide sense as accepted e Lorence (1991); its — hae. ipo heen summar ized һу BAM f 1], is treatment t Mena n CHARACTERS This analysis is based on morphological char- acters. The characters used and their states are Table 1. Several of these ( $ UM In Шо їп > analy: ses. The character states coded for ae spe- cies are shown in Table Because examination ar the relationships among the epiphytic groups was one of the goals of this nalysis, habit was not included as a character. тне characters were ж опей so that the number of polymorphic taxa w Po жес taxa were coded for the condit was presumably less derived, based о com n tion that ANALYSIS Several groups of analyses we both PAUP (Swofford, 1990) and ead (er ris, 1988): 1. considering all species and using ш of the our acters listed in Tables 1 and 2 ristically, with e characters treated ated rario y as ordered or unorde apu rin all s 5 excluding th pomorphic characters, pane’ with the S gerentem treated variously as © or unordered; and m rdered Volume 81, Number 4 1994 Taylor 575 Revision of Hillia TABLE Characters and ERE states E in cladistic analyses en panei in individual species and were elimi ; see text for further explanation. Tí RU or of piante: 0 = green, l = reddish brown. г absent ls ре ubescence: vegetative Lini pier aes er *0- inkled. 5 bark ci Айы Pid as 6 leaf size: 0 — more than 12 cm buds | = ба 12 ст long, 5-5 cm long. 7 leaf 0= e to acuminate, 1 = rounded. 8 af textur E 1= ee 2 огїасе 9 leaf venation prominence: 0 = all prominent abaxially, 1 = primary and secondary veins д abaxially, = primary vein prominent abaxia 10 leaf venation pattern: 0 = praed e 11 number of secondary veins: 0 = 9-1 = " 8 pai 12 anisophylly* 0 = abeent, l= р 13 interpetiolar, 1 = interpetiolar and intrapetiolar. 14 number of flowers: 0 = = 1-3 15 corolla shape: 0 = ake |: = кашы. ds = tubular, 3 = infi: i 16 corolla habe: 0 = white, 1 = green, or green flushed wit — гең or orange, 2 = green flushed with red, 3 red, 4 = bright red. 17 cl aestivation: 0 = convolute, 1 = imbricate, 2 = va 18 corolla lobe be number: 0 = 4, low 2 ‚ 2 = 8-10. 19 corolla lobe length: 0 = 8-15 mm, 1 = 16-2 = more than 25 mm. 20 corolla tube internal epidermis: 0 = glabrous, 1 = papillose. 2 calyx 2 form: 0 = tube present, 1 = absent or vided 22 поя er postion 0 = included to slightly exserted, 1 23 s sigma edd ape: 0 = subcapitate, 1 = with two linear ranches, 2 — Кү two Tt rice flattened branches. 24 Poe position: 0 = below anthers, 1 = above anthers, cluded, 2 = above anthers, exserted. 25 pi on capsule: 0 = absent, 1 = T xs " capsule texture: 0 — capsule furti: 0= mai to spe rigido "s ridged, 1 = lo ae y wing 29 € capsule dehiscence: 0 = basipe em 1= у. mose seeds: 0 = absent, 1 = prese 3 К Considering subsets о ies and excluding the ۴ pe ic carters analyzed heuristi- with со; algorithms, with th ecc treated ae as ordered or unor- dered. TABLE 2. Character state coding used in the cladistic analyses. TM characters and their states are numbered Table k EN apa ШЕ dati; Fam Tac qu e letieae); ARCY Ar сонар тыла (eoi LADE Ladenbergia (Cinchoneae); FERD Fe PA a (Cin- eee ); outarea (Cinchoneae or Condami- ae); CINC Cinchona (Ginchonene); BALM Balmea; dare T T T Co smibuena à ; CVAL Cos oi; HKIL H. killipii; н. macromeris; HMAP HPAR H. parasitica; HWUR Н. wur- crocarpa; CMAT bu tieri; HOAX Н. oaxacana; HRIV Н. CN HTRI H. triflora var. triflora. Characters 12345 67891 11111 11112 22222 2222 0 12345 67890 12345 6789 ROND 00000 00000 00002 01000 00110 0000 RCY 01000 20220 10002 02000 00110 0000 LADE 00000 00100 00000 02101 00210 1000 FERD 00000 00000 00000 00110 00010 1000 OUT 000 10000 00001 00110 01010 1000 CINC 00000 00000 00000 02110 00210 0010 ALM 01100 00 10002 40100 10110 1000 CGRA 01100 00120 10100 01111 00210 1000 CMAC 01100 00120 10100 01111 00210 1000 CMAT 01100 11220 10100 01111 00210 1000 СУА, 11100 11220 10120 00111 10210 1000 HBO 01100 00200 00020 00110 10010 770? HKIL 1111 10010 10020 00120 10010 1001 HMAB 100 20220 10020 00110 10101 1001 MAM 01100 00010 10020 00120 10010 ??0? HMAP 1100 00210 10020 00120 10010 1001 PAR 1100 10120 10020 00120 10010 1001 WUR 01100 10010 10020 00110 10010 770? HPAN 1100 21120 10020 00000 10100 1001 LOR )1100 10220 10020 00010 10100 1001 HMAX 01100 11220 10020 00010 10100 1001 PAL 01100 21220 10020 00000 10100 1001 TET 01100 11120 10020 00010 10100 1101 COS 1100 10220 10021 10100 00021 1001 HFOL 01100 11220 10021 20100 10021 1001 GRA 01100 00220 10021 10100 10010 1001 HILL 01100 00220 10011 10100 10021 1001 HPSA 01100 10220 10021 10100 10011 ??0? HSAL 01100 11120 10011 ?0100 10011 ??0? HULE 01100 00221 10021 10200 10011 1001 ALL 01100 10220 10021 30100 10020 1001 LON 01100 10220 10021 30110 11020 1001 HPIT 01100 10220 11013 40100 10020 1001 HOAX 01100 10221 10021 20100 10020 170? HRIV 01100 10220 10021 20100 10021 1001 TRI 01100 10220 10012 40100 10020 1001 576 Annals of the Missouri Botanical Garden Each ше of "e — — out- groups, 1 1 Jom ecd ofa scm focus group. Cosmibuena and each subgenus of Hillia were taken in turn as focus groups. Auta BAT: characters were eliminated from the second and thir g species. S third group té сауы pm of species вси іп ће full a analyses, 36 in all, exceeds the size at which either cad analysis age is predicted to return y Res Identical cladograms were obta using both PAUP and Hennig86. Selected AREIA hypotheses will be pre: aisi. and evaluated below; their мааа implications will be discussed іп quent sections on classification and one i Tt pots be clea ar from: the hyp coim eses M ai unequivocally indicat ea ч robe phylogenetic relationship bi these "pae le pp las feat ud 1 23 1 is left to future systematist The first group of viis found vis equally ograms. One these sh in Figure ll of these sladograms, Balmea oa grouped with Hs lia subg, Ravnia Hillia ILL ЕШ Senden. as a grade immediately basal to Hillia; Hillia- Balmea-Cosmibuena was monoph and Hilli Tetrandrae was monophyletic The strict consensus of these five clad s i These five cladograms differed i arrang of species o епи е relationships among the species of subgenus Шш. ишь жеге EEE asa а grade of subgenus Illustres with those of rad Rav- nia. Ї these is shown in Figure 4. In all of these Wires ba, Cosmibuena was ar- hylet iak Hillia- Cosinib ena was dives Bal. теа was placed inen tely basal to this group; Hillia- ae almea was monophyletic; е: Tetrandrae was monophyletic. The d in the arrangement of species of ease Hill, which were always arranged in вори relanions hipy ні the ев of Results from the third v of rere are not presented here. These did n p ine and added no information. TRIBAL CLASSIFICATION OF HILLIA The neri s Hillia within the Rubiaceae has ignificant disagreement, and its placement i ix altered as ideas about the reet of the family have evolve tum nn (1891) AES pt m authors ing Hillia in e Cinchoneae DC., disti ilias by capsular nj yb ascendingly mé cated winged us of the oideae Raf, Pesci apts by numerous “ovules in each loc ыб urt (1958) proposed the next, € a in his s guished by the presenc raphides. Hillia oc- cupied an isolated position within ubfamily and was not classified further, although Verdcourt suggested that it might be placed in its own mono- ypic Bremekamp (1966) Pte the вс е n his own and Verdcou acteristics including sculpturing of the e the pr ce or absence of raphides, whic j considered to >have evolved only once іп the fal k illi ubfamily oidea Bre win along "^ Ravnia Oerst and Терра 5с Ae '& Cham on the gie „thr: genera of ae contort a distinctive exotestal ] cell nly in Gea- in the poe дат e DC. (Breme been synonymized with Hillia (Taylor, eni thus Hillia.. occupies an isolated positi mekamp’s classification. Volume 81, Number 4 Taylor 577 1994 Revision of Hillia HMAB HLOR HMAX X HTET е НРАМ 3 9o HPAL р Nö EEE X—— HBON X [—— HMAP 5 HPAR $ H— HKIL Pn X--X——HWwUR E 2 HMAM 8 НОВА HILL we” Б ozo 3 HULE e HSAL | [29 ве — HPSA 2 HCOS $ HFOL 3 X— HRIV & | НОАХ ra ъё ~ FHHI— ном 3 2 HALL H H— HPIT —* — HTRI 152 DX—BALM ae One of fi 1 species obtained from һе б. of analyses as described i in the t, based о e full character set алинин in Table 2. Analysis was ine ane pis mheni onig and Ашуу breaking aa characters were treated as or rdered; polarity was Tmined by m able з -letter Kodi: а ating taxa are defined in Table 2. Black “rectangles indicate Ajo el ume that are not E on the clado; ogram; open recta angl parallel li parallelisms. See also discussion in the text and Figure 2. idw most recent comprehensive classification of that within this subfamily d is = to th Re Rubiaceae was hm орою by Robbr чы (1999) Cinchoneae but "reduced," extreme Phides T case of this trend." He E filli in its Hie 2° зау 1988), pus he pa Ta own tribe, citing “many characters justifying its the Cinchonoideae. Robbrecht indicated separation as a monogeneric tribe," but without Annals о Missouri а Garden Fic Strict co grams кеу in the pi ар Four-letter codes designating taxa are defined а msensus tree of the five туча first set of analyses described gue — dens ME — depends that ibn Structures are ‚млн as as out- growths o т extensions of the testa май, : has d | trichomes. However, he did not otherwise MR жеӊ = pid 1 outgrowths cl as trichom The position of M within м Беа и been examined іп detail by Andersson & Perss (1991) ina wt s analyst of pese that не been proposed by various authors for inclusion in tuin vins of pie radi igl хел) wall, but which — ere pos- -— the sister group m the Cinch "eec eae s. str. rn of the mation may not have es о] und in the Cin reg nu. Pm that group, on which this decision is based do clusive. Канына. characters, а5 ve wall sculpturing and raphides appear 0 h ose t indicate Classific. of Hillia therefore see ide pend on character weighting: this genus © Vna in a rite based on 1 pre rived member е Cinchoneae, based Volume 81, Number 4 Taylor 579 1994 Revision of Hillia FIGURE 3 “One arth В А А Mol А СИИИ; г, T af, the second set of anal , S att an "n in Tabl xcluding autapomorphic characters. d was heuristic СЯ nnig* and br: anch- т pon c dancer Were treated as unordered; polarity was determined чыч, "айо as ancestral. Numbers characters and their ir states are defined in Table k. p rm ignating taxa nd in Table 2. 2. Symbols are the same as in in Figure 1. See also Figure 4 and discussi pn text. quracteristics it shares with that group. Isolating of Hillia and the و‎ shaped, strongly erose Hillia in a monotypic tribe requires the assumption that the characteristics that distinguish it are all filaments ‘that resemble the numerous flamen of independently from the character states Mages need not have arisen independently, wn їп the Cinchoneae, rather than as part of a y be steps in a tra nsformation series. If the transformation бее H owever, the comose ies suis are viewed in this way, isolating Hillia Annals of th Missouri sod GU Garden |. Ficus 4. _ Strict consensus tree of the three. clado- Ssi text. Tan letter codes designating taxa are defined in Table 2. results in an arbitrary separation of the derived end of an evolutionary trend toward reduction, in e istinctness rather than its useful whi ps. Classificatio reflects relationships rather than e s Hillia i mbe Cinchoneae, acies closely — with Bal- теа and Cosmibue GENERIC RELATIONSHIPS OF HILLIA As discussed in the previous section, the rela- tionships of Hillia are vy the Cinchoneae (Rob- brecht, 1988; Andersson & Persson, 1991). Most of the characteristics that distinguish Hillia are 8 are Balmea, Cosmi- buena, a Hymenodictyon Wall. enodictyon includes about 20 occasionally Asia, Indonesia, Mad- г aph Glande 4 EEN does not seem closely related to Hi lia. almea is a monotypic genus found in southern (And pers. ; a cal almost com pletely divided into five lobes; tubular to fun form dark red corolla with five X— lobes that are — reflexed; ‘shortly ine styles; eat beak ed, c cal, apsules. 1 ete im FEES 2 йз small scale i rne in pedunculate ; the presence of r nce у von * нен a al с or fascicle; salverform terminal cym ; j M. Tay or); quincuncial corolla aestivation; Volume 81, Number 4 1994 Taylor Revision of Hillia densely panillgse timeas; каршын of the inner (Taylor, 1992). Cosmibuena seems closely related to Hi Ша апа aba fo Balmea, Cladistic айн lobes. Several species of this subgenus also and flowers, although reduced taxa. om Hillia PRE to be the кор corolla have relatively small leaves a t © th A | 2 f, } J рогурпу М po son o Da Y n (e. 1- pu The ene) ‘of f raphides i in Cosmi- m" 1966; Robbrecht dire the "i en based in part on this one feature aiu cuiii reexamination. How- r, Cosmibuena also shares some biis with d h Ы = ee A £: 1 L ly Б. Дл) white corollas with m to; six ем The flowers are strongly d are presumably pollinated ‘by sphingids or beri xcii "hé bilobed ignes are € subospitate w somewhat S species have leaf domatia and well-marked ondary leaf veins, and the higher-order deii is ae тер matke d in H. bonoi Steyerm. A wide d flower sizes is Beh within о о are not found in Hillia, so the systematics of this group are far from resolved. INFRAGENERIC SYSTEMATICS Five subgenera are recognized here, three newly бейе. The dtliteitetion of these subgenera is d primaril , number of lobes, and = of the corolla. The corolla features are gen- erally аы with similarities in stigma form and En raging to йы ned» йна "ii tap of rolla tube. Т the c E subgenus, including the largest flowers in the genus (Him тас [т una). Subgenus Hillia h den a monophyletic group; the characters avail- able are insufficient to distinguish between these жа Hillia macbridei appears to be intermediate between subgenus Hillia and subgenus Tetrandrae = 2 ü 1 | C z ۳ А h +, E е stigma position and morphology of subgenus Te- Оша while its corollas with five to six long haracteristic of sub Hillia may ja shared in unrelated pegia sith similar о unique іп its reduced Vegetatively this B that are acute and usually stiffly pollinati junction with corolla form appears to be useful for phylogenetic inference in the Rubiac e (Brem & Jansen, 1990 . The geographic SOLES of Bois resulting subgenera and the presence ice ad- 1t1 Support the classification proposed | here genus Te is th ited of these groups. It is kic эн by ita cii коша with four lobes. Tone flowers by sphingids or оше types of moths. “White в al- verform corollas probably represent the ancestral condition in the genus and are also found in sub- е tigmas positioned below the middl. tube and far below the anthers, which are situated at the mouth of the tube. This sti igma a morpho logy ve-lobed corolla. Within 5 Tetran- rae most species have rounded leaf apices and шинчан All cladistic analyses consistently Tetrandr ae. This s species is here ‚ placed i in ité own nus, subpenus E vnia share bonitas corollas with five to ten знан and sub- si anthers. This stigma morphology is also shared with some species of subgenus Hillia. These two sub- enera appear to be more closely rela ted to each ipea ore der red. Subgenus Psi is characterized by corollas that are predomin раму. к e beg may be flushed with yellow, e, pink, . Noth- ing is Сен of the ora ober of don species, be adapted for pollination s. With th ti H grayumii C. M. s of ‘this section are also char- us Illustres in- acterized by nie fruit. Subge ludes the est v on in leaf form, anther position, and n of corolla lobes, although a or trends his nia) may represent a grade rather than a mono- T Y 1 а ЖУ КЫ SES, NN à E! puotog EK о Ly Annals of the Missouri Botanical Garden able are insufficient to distinguish between these possibilities. исе: GIG ed by d co- T ollas This group Fa s discussed и in detail by Taylor ( er treatment ei dern риди the vos subge ot altering its inii sands 2 занн пиз is apparently the m deri he lion of subgenus Ravnia from s pu Illustres is + barod excluso on flower iioi ziven the sim- + between the i floral bove icio. i red iini 6. Subge us Ravnia is here assumed to epresent a monophyletic group, but viene "a may show th sified in subgenus Hura at some TAXONOMIC TREATMENT This study is based on чу of анара ѕрес- niola, i Measurements of soft parts given here are based on dri cimen; planta. б, Hille are fleshy A ahri aes ) accompanies the Prena of herbarium а. mens. Hillia Jacquin, Enum. Syst. Pl. 3. 1760. TYPE: Hillia parasitica Jacq. Fereiria Vell. ex V - Lusit. Bras. Spec. 21, t. 1, . 1788. тү TYPE: gers 4 vellozoana Sch А & t. f., Syst. Veg. 7: 83. 1829. [= Hillia par- perp Jac cq. Saldanha Vell., Fl. Flum . 1825. ТҮРЕ: Saldanha nobilis Vell. inl ИШЕ M Jacq.], lectotype desi gnated her: E биз Viden: Uy eem Naturhist. Foren. av n 18 TYPE: Ravnia tri- [s Hillia ARS ul )C. M. Taylor.] Sch mall trees, suffrutescent herbs, or li- Ee >. тр; thr roughout, fee se ene pim quadrate, iini becoming terete; bark в оо « or ra Mg papyra- Rs (Н. peas gray to brown or sometim own rarely coppery red (H. killipii). a. oppi decussate, isophyllo ous or rarely anisophyllous (H. triflora var. ubsessile n innate or r rarely subpalmate 8, dist al ve alling. Flow olitary o subsessile cyme e (H. "s eee H. saldanhae, H. 21 -—— dn an with ped cels stout, sometimes subtend nded lacking or rarely with short lobes (H. costanensis), green, with lobes 4- 10, sometimes variable in n 1 individual plant; corollas salverform (8 Hillia, subg. Andinae, subg. Tetrandrae) to fun- nelform (subg. Illustres, pe Ravnia), t lust subg. Illustres, subg. Rav flora), white verd ыле а, E Tetran dr ae) to sometim r top or mi on axile placentas; "idi ini stigmas 2, subcapitate to ini Sis abore gated-linear, papillose, positioned at o | r sometim Il bel thers. ê x a cylindrical, woody, sm or pene m te dged t y ed ( ini 00 dinally гі о rarely wing usually with beak formed by old disk and some E also the elo: ema ovary apex, septic basipetally dehiscent; seeds flattened, rhombic, with i" anaceous to ра Volume 81, Number 4 Taylor 583 1994 Revision of Hillia ginal wings, with tufted multiseriate filaments KEY TO THE SUBGENERA OF HILLIA nee ") extending from cell wall at acropetal у, Corollas salverform, hite or pale р ы. Twenty-four species, southern Mexico and An- Croll lobes 5—6; to Peru, Bolivia, and southeastern Brazil, in linear, 0.5-4 mm long, postion above = I wet forest zones at 0-2400 m. f Pen ane frigo 3a. Stamens inserted near top of corolla 8 anha, Vellozo tube; stigmas subcapitate to ares! described the a as Bees parted with REE linear, 0.5-2 mm long, риби lobes, the corolla as five-lobed, and the stamens as above anthers __ 1]. " mas s Hillia six. He named two species, S. illustris Vell. anı » iu pigra middle ram S. nobilis. The identities of both speci 1 ea, 2 am . nobilis. The identities о species are clear long, ond p аы (Steyermark, 1972 th of them typically have 2. subgenus Andinae calyx and corolla lobes and six stamens; the 2b. Corolla lobes stigmas linear, 2-6 mm long, positi ed below anthers generic description thus applies equally ll (or m 5. subgenus Tetran н poorly) to each of them. Although Vellozo's generic — 1 Corollas funnelform to tubular or Ба. description is somewhat weighted toward S. illus- green TS with o pink, orange, or dull tris, his description and illustration of a plant with purple, Men to bright red. 4a. P ecl unnelform, green or e flushed with sii , pink, orang у) т dull purple alee >ы эчи à ubgenus //lustres description of S. nobilis accurately depict H. par- ab. Corollas fuco Seat ir inflat itica, and that species is theref h th salmon to bright red _. 5. subgenus Каота five calyx and corolla lobes are either inaccurate, denict "РЕ ҮН ЖУ ЖЕҢ Oe dE 2 3 lectotype KEY To THE SPECIES OF HILLIA la. Leaf blad rowly elli 1 ‚ 3-4 times as long as wide, 0. аз cm long. ٠ Leaf blades 1 1 ky coriaceous; plants of Peru and Ec uad ie naan ER Н H. macbridei Leaf Ыза, и f M. H. panamensis lb. e Mie Pipa vane о кардын NA ог obovate, less than 3 times as a as wide, 2-22 сш [ew rf. bie ries Secondary and tertiary 4. H. bonoi 4b. Primary and secondary but not prr venation prominent or prominulous 5a. Bark БЕ red, peeling and wrinkled in small circumferential rings, , especially ` young twigs SLES, Н. epii 5b. Bark ey -brown or dull red-brown, not peeling or wrinkled in circumferential ve 6a. Leaf blades 4.5-10 cm long, 1.8-5 cm wide; corolla lobes 12-26 н RET нА . H. wurdackii 6b. I blades 9-22 cm long, 3-10 cm wide; corolla lobes 35-9 90 mm 0 mm long - Corolla dobes:25 54mm Mong аа ees x macrophylla - Corolla lobes 75- -90 mm | long . H. macromeris 3b. al ven. N 2 1 bh :al i f f: сл. lation plar ic Bees f blades ro сва tide at apex. вв salverform, lobes 4; stigmas 5- 15 mm п held below anthers. а. psules tag саан Pn Moa or wings to 1 mm high; Greater Апі northern Nica . H. tetrandra 10b. e а согіасеоиѕ, аі thickly so; capsules men or with r with slight а udinal ern Nicaragu ne Corolla tubes 42-55) mm long; lobes 15-27 mm за LLL 10. Н. maroni llb. Corolla tubes 35-40 mm long; lobes 8-12 long 18 Н. palmana 9b. Corollas funnelform, lobes 5-9; stigmas 1-2 mm long, held above anthers or exse 12a. Leaf blades subcoriaceous; stigmas included, held 2-3 mm below top of corola tube . H. saldanhae 12b. Leaf blades thickl ; styl erted 4-5 mm beyond t = Mea е : Hs 18. H. foldatsii 8b. Leaf blades acute to acuminate a 13a. LA f venation лт pis with hal secondary veins arising below middle of blade. Corolla tubes m long M, 20. H. oaxacana 584 Annals Mi ere ‘ue Garden 14b. Corolla tubes = to yellow-green, 28-39 mm long, lobes 6-7 mm long; HMM of h A 16. Н. Amazonian merica ulei 13b. Leaf venation mon with at least one pair of secondary veins tal t blade. 15a. Leaves strongly anisophyllous, the larger leaf at de at least 1. times as long as the shorter leaf H. triflora var. 15b. Leaves isophyllous or -— anisophyllous, the larger veia ata node less than 1 times as Jong as the zr r leaf. l6a. Calyx limb 36-68 mm long, with tube 33-56 mm long; leaf Bun rounded to slightly once at base Н.е ostanensis 16b. Calyx limb 3-36 mm long and lobed to base or Lt leaf blades "adi to attenuate a 17a. Flowers (1-)3; corollas tubular to eae блай "i چ‎ SEE 24a. H. triflora var. "р Tb Гур solitary(—3); corollas salverform or Рн white or gree! ange- - Ca lyx lobes narrowly triangular to linear, 18-36 mm oe OUR . H. rivalis 18b. ЖНА vul PK or triangular to oblanceolate, iss or 3-35 mm long. pus ‚е salv sfr white or pale green, sometimes pink on surfaces exposed in bud. 20a. Corola mates (5-)6(-9), the tubes (38- аЗ ноя mm g; plants of the Antilles and South Апи Nen . parasitica 20b. co lobes 4, the tubes 45-75 mm long; plants of uthern Mexico and Central bu beu eH Н. loranthoides 19b. ase funnelforni, green to red-orange. la. ; plants of Costa Rica and Pan- 22a. — lobes 13-17 mm long; free porti s 9— an ке long .. 22. H. lo lone 22b. ен: iba mm long; free portion A aments ca afters ng ج‎ ‘le 21b. Corollas ere to eee -green; plants ‚ of Costa Rica to ut! 23a. Leaf GÊ 4.5-6.5 ст long ———————— |, pus йн ТАЕНЕ psammophila 23b. гуча blades 8.5-15 cm long. kii 4a. Calyx limb 9-35 mm gone "u^ -— 24b. Calyx limb lacking o or to 0.5 mm long, truncate to dentate ........... __. 13. H. grayunii 1. Hillia subg. Hillia. types, F, K not seen, MO, NY, UC not seen, Calyx limb E ar with 5-6 bee. согор BS. USMshob SBE) salverform, Shrubs or small trees to 7 m tall; bar k y in bud, sometimes flushed with pink or pad be- brown to red-brown, smooth. Leaf blades elliptic coming yellowed with age, the lobes 5-6; stamens to реч! ovate, 4.5-10 cm long, 16.08 attached near apex of corolla tube; anthers sub ide, acuminate at apex with tip 5-10.mm ma sessile; мыт кашлы to shortly linear, po- cuneate to broadly obtuse at base, chari sitioned just above anther secondary veins pinnate, 4-8 pairs, midrib " Seve geste Antilles andSouth America. secondary veins somewhat impre adi ei prominent abaxially, without domatia; mex l. Hillia wurdackii r^r Mem w petioles 6-11 mm long; stipules s 14- —40 mm 3 York Bot. Gard. 23: 294. 1972. TYPE: ‘ce 6-18 mm wide, ing red e Mug ges. ed on road to cba Rioja 5 сала 7 регеа es 1-1.5 mm long; bra 0 k 0 8 Oct. 1964, Р. С. Hutchinson & JRE or E pale green on surfaces expos Wright 6799 баю, VEN not seen; iso- 40-58 mm long, the lobes 5-6, (12)20- -26 mm Volume 81, Number 4 1994 Taylor 585 Revision of Hillia B. Hillia killiz Fr б: Habit with flower.—D. Flow (US); C, from Mn et al. 6799 (MO); D long, 9-12 mm wide, AE to triangular, acute; саа 5-6, iim 10-11 mm long, with tips mm be v positioned 3 low top of corolla tube; ovar т 10 mm long; stigmas subcapitate, 2—3 mm long, Positioned immediately above anthers. Capsule 6.5-8 cm long including beak 6-7 ng, 1- iniga phenology, and distribution (Fig. З). Ecuador and Peru, in wet forests a +8 2400 т. Collected in flower n to Hillia wurdackii is simil rophy la, Т macromeris, and ái killipii in its d taceous coriaceous leaves with the s secondary veins prom- nt on the abaxial s e, but differs from the uen 0 weir ckii in its relat soe Short corolla Ыы, ey mm long Additional spe. Ret em — s ай to Colonia 24 de Mayo, 2.5 km W on road uyo- “Tena road at элыз 9, К, 775870. " Stein О). U. Am mee peg Bon- rw. of Pom: m on road ?40'S, Knapp et al. 7497 (МО); domo Vil- wer d spak үс A, from , from Woytkowski 8236 (M abit with flower.—B. Capsule. C, D. Hillia wurdackii Steyerm.— Killip & Smith 26085 (NY); Б = rom Camp E-2405 О). All to same 5 caniza, Sagástegui 5997 (US); Mendoza, Woytkowski 8236 Жы = 1-5 do Fe (150°) of e 27 (US, photo NY). Cajamarca: Prov. Cu- ag bes Andrés- voe plas, El Suro, Since: 317 (MO). 2. Hillia e Standley, Publ. Columbian Mus SE AOE 0, 900 m, Jun 0,R. 6186 K, photo E 5 isotype, F). T Pittiera 9 1981. TYPE: ra: ag or ММ а, ада “Са: рае "cater «е rande-Minas de Car- bón de Lo ы 197 79, arcano-Berti = 1 Pea 519- © "edad VEN; isotype, MER en). 11; уен Tach c Suffrutescent — € or small trees to 7 o red- br own, smooth. Leaf пр, Flowers ыу; баен es 1.5-2 mm long; bicis Annals of Missouri Lingle Garden lacking or 1.5- 11 x 1.5-5 mm; calyx limb lack- е, triangular to narrowly nn acute; an- 5 long, with tips a g mm long. Figur Steyermark (1981: fig. 1, as H marcano- E. Habitat, phenology, one distribution (Fig. 6). Costa Rica, Ve la, Colomb; in flower December to July and September to tober, in fruit January, May, July to August, and November. This eg is distinguished by its large leaves and corollas. It is similar to Hillia wurdackii, H. macromeris, d Н. killipii in its chartaceous to subcoriaceous leaves we the secondary j veins as wem as those of н macrophylla. only bg Ye que that would separate Hilli **pustulose" bark (which appears rather lenticellate in Steyermar rk's illus- with flower.— B. Flowe r opened near top of f Срна » нер. —D. Leaf. E-G. Hillia macbridei Standl. fog браћа .—G. Sile (r "M A, B, from Lehmann BT 416 (NY); С to twice G, from Schunke 436 (F). A-F to same scale, Volume 81, Number 4 1994 Taylor 587 Revision of Hillia tration), a *verrucose" abaxial leaf surface, and ut З mm. The to mea- e бооштепсе of раа bark аяп уег- эшо or trees to G m pall or serenely lianas; ba f blades Рд to кн ovate, 10- 19 cm Я 3- cm wide, acute acuminate: E with tip 1— end ceous to subcoriaceous; decem veins innate, на 8 ра irs, in the range of уапаноп у seen in wand species of Hillia. A si j is seen in other species “Hillia, БОН NE spe Therefore, this species is not maintained her Representative specimens examined. COSTA RICA Puntarenas: Upala, El Pilon, cabeceras del Rio Celeste, 1049'N, 84°57 'W, Herrera 1320 (MO). COLOMBIA. р о тере, vereda Santa Rita, Fir inca Mo ontepinar, 6*17'N, 78°08' W. ental de EL Sa acamefio, Jarmil lo ó-Medellín roa Cory d а 24168 or MO). Candinamarea: al S de Santandercito, laguna de Catarnica, 4 de Bogotá, Uribe 3197 (COL, US). = Mees Cones Occidental, vertiente occidental, hoy: о Digu: San Juan, abajo de Queremal a la tlie int rio ps 52-83, йазы 23963 (COL, F, US). Meta: Cordiller: р o NO, macizo Renjif: lrobo & liher 1059 (МО, US). йо: Reserva Planada, 1°10’ №, 77°58'Ұ, Restrepo 449 (MO), de Be navides 9602 ( Ng 1?05'N, 78°01'W, Gentry et 60342 (MO). No an i DOR y Rio Blanco and Rio Norcay on d between Chacanceo and Molleturo, Steyermark 52827 (F). Caüar: quebrada of San Migue! r acienda Monte Negro, Játiva & Epling 285 (NY) Carchi: environs of Chical 12 km below Maldonado on Río San Juan, 1°04’N, OM red et da 4519 n Cotopaxi: 3 k еса: inas on new 4и to Santa Ross Dodson "5 &бешу 9136 (MO). ша о San уо, 2.5 km W оп road fro in Puyo- M Km 9, 1°25’S, 77°58'W, Stein & Tucker 3108 (MO). Pichineha: old Quito-Santo Domingo road, 2-9 Hug , NY). PERU. i pre \ оваа Nacional de paria, Abeja, a 20 km arriba de la confluencia con el río Ucayali, Schunke 5247 (F). 3. Hillia macromeris Standl., Publ. Field Co- ian t Ser 2. 1930. TYPE: pee cs тт м Santa Maria оп mann whe Co Гећ rater die 41 : ней vi бү, A, F; i s, F, K, photo of K sheet NY) prominulous i prominent abaxially, ропа dom- 32- -70 mm long, T- жщ е. Flowers 's solitary; peduncles ca. 2 mm limb lacking; 14.5-15 cm a n lobes 6, 1. 5-9 cm 5 mm wide, very narro attenuate; non 6, bes with tips positioned 12—15 5-1 0 mm ng, positioned i er Gaba ca. 11 cm long inclu ine rae al mm long, ca. 12 mm diam., not stipitate, re or Mos 8-10 low longitudinal ribs; seeds ca. 0.5 mm, with filaments ca. 10 mm long. 6 TA, В. SASIR, phenology, and distribution (Fig. 8). Colombia to Peru, in wet forests at 100-1890 m. сб: in flower in May, September, and No- vember, in fruit in No Теш. This species » (ырш һу its very long pe pee ve an- lobi: stigmas decir linear, immediately of any other species of Hillia. It is ‘similar to — а = 3 8 S 3 p^ BER e mp un v e < t © e, n m la] ese onl size, but it has flowers about half as large. ж specimens examined. ECUADOR. Car- chi: along the Rio Verde, at trail to Rafael’s Mountain Fia. OBEN, 8'08'W, Hoover viri sud Morona- и iago: 6 К а (ипдег рне E Unión, Harling & & stipes 24490 vedi 4 PERU ve Quebra 1 day's s Martin: Prov. Mariscal Cá , distri Río de la Plata, NE of тоо Schunke 8380 (MO). 4. зна bonoi Steyermark, Ernstia 42: 5. 1987. ela. Tà : ito ubio y Las Delicias, 2000 m, 2 Apr. 1984, ^ Bue; 3797 (holotype, VEN not seen, photo MO). th, gray-brown o red-brown. qum blades Т arpa to some- ri ovate, 14.5-16.8 cm long, 9-10.8 588 Annals Puit па Garden h tip 1-1.5 em long, obtuse to rounded at ае stiffly subcoriaceous to coria- ceous; secondary veins pinnate, ca. 10 pairs, mid- ү secon ndary, won er veins бд state жаяр" somewhat aa ieri 15- 28 g a. 38 x not seen А olitary, a: ee calyx limb not seen; corollas salverform, و‎ ike tubes ca. 7 c ү the lobes 5, с cm long, р де, narrowly lanceolate to triangular, ca acute; anthers 5—6, subsessile, with tips positioned 2-3 mm below top of corolla tube; stigmas sub- capitate, ca. 2-3 mm long, Sper ned immediately above anthers. Capsules not see Barrel phenology, and distribution. wn only Kn. the ia ie locality, in cloud forest at ages pee in flower in April. inguished by its leaves, rtionally gre than those most other species of Hillia, with the bu BE ary and higher-order venation н е above and prominent on the lower surfac 5. S Meus ridet pis Field nid ‚+ Bot. Ser. 9. 1930. 900 m, 6 J 929, A. C. Smith ed быы ур; F, dme F; зел NY, US). or small trees to 5 m tall or sometimes 3 ong, 3 wide, narrowly tri мане acute; anthers 5, I 6-7 mm long, with tips " sitioned 6-7 mm below top of ан in оу; 5-10 mm long; stigmas shortly lin ca. vie е positioned imm еі iately above anthers Cap- s 7-10 с 1-5 mm lo ca. e 10 mm i dia ., not s ca. 10 low анана ribs; seeds 2-5 х 0.8-1 with filaments 10-25 mm long. Figure 5A, B. ng, бирө; smooth or with Habitat, phenology, nl dre a7 and Ecuador, in wet for that is circumferentially wrinkled and sometimes Representative specimens examined. ECUADOR. apo: region of Archidona- Tena along Rio Pano at foot of Cordillera de Guacamayos W of Tena, Ownbe (F, US). PERU. Cuzco: Rio Arasa NE of Cuzco, Sande man 3672 (F) reto: Pumayacu between Bal: rto d Moyobamba, urs 5 (A, F, GH, US). San Mar- tin: Prov. Rioja, Pedro Ruíz-Moyobamba road, Km 390 5°50'S, 77°45'W, Су, 4385 (МО). 6. Hillia ombre Jacquin, ru. Pl. Carib. 18. 1760; Select. Stirp. Am 66. oad TYPE: Martinique, КУЛКУ: s.n. Ac. not seen; cited by Moore & Rendle ks Hilig onions Sw., Prodr. 58. 1788, nom. inval., pro > Cosmibuena amine Ruiz & Pav., Fl Peruv. 3: 4,1. 1802. ТҮР! play 1798. d Tafalla 15/2 24 (holotype, MA not seen, photos , MO; isotype, Saldanha nobilis Vell., Fl. Flum. 142. t. 3: 158. 1825. Jell.) Steyerm., Ten Bot. Gard. 23: 292. 1972. TYPE: Brazil. New York Rio de Janeiro моор Flum. t. 3: 158, lectotype designa: т: Еегеігеа vellzonna ‘Schult. & Schult. f., on Veg. 829. TYPE: Fl. Lusit. ^ Spec. t. 1, fig. 8, 1788, EHE ‘designated he: her / gp noi e erm. yocp of H. b ; St 2 ent б pied. nds) in South А Volume 81, Number 4 1994 Taylor 589 Revision of Hillia FIGURE зк А, B. Hillia parasitica Jacq. —A. Habi uth Am —B. Habit with flower, Hillia Bene Cham. & Schltdl., Linnaea 4: 201. iv : Brazil. Brasilia aequino octiali, Sellow в Ач doni B destroyed; isotypes, P, US). "ner montana Mart., Flora 24, Beibl. 2: 80. 1841. Ma razil. Minas Gita: in sylvis H lla Ric ca, April, Oreas s.n. (бор illia boliviana Britton, Mem. To: 1893. TYPE: Bolivia. Cochab ungas, 1890, ^o ti 660 (holo otype, NY; isotypes, F, GH, M M odorata К. Krai , Bot. Jahrb. Т 40: 321. 908. TYPE: Peru. Junin: Pr ov. Tarma, prope Hu- "yp 1800 m Jan. 1903, A. Weberbauer NY we (holo Wd B пес рһоїоз Е, СН, МО L Hillia weberbaueri усу Field Mus. Nat. Hist., Bot. . TYPE: Peru. Without locality, 1909- 1914, 4 Гана 6955 (holotype, Е, photo F Hilli ч арта. Steyerm., Me New York Bot. Gard "up abay 8 May уйн, J. A. Ste Meri pod erint F, photo NY). а S uffrutescent herbs, shrubs. or small trees to 8 it with flower, “typical” form from the Antilles and eastern » Р “nobilis” form from northwestern South America. A, from Otero 639 (МО); Tom Bae, & ШАР, 3564 (MO). А, В to same scale. n tall ]-b mooth. Leaf bade s li clit -oblong, 3.4- 13 c m long, 7-6.5 wide, acute to usually acuminate at wes with tp 5- 10 m m lo ong, cuneate b acute at pinnate, (2-)4-8 gis pairs, plane or sometimes m icd flat; pe s 9-55 mm long, 4-1 ide, sometimes re Flowers solitary; peduncles 1-4 mm long; bracts lacking or 2-7(-9*) x 1-4 mm; жөн limb lacking or divided to base, ns lobes (5-)6(-9), 3-12 mm à triangu! lig- corollas ч. ink o b 0.8-6 mm wide, ulate or orbicular, acute to rounded; TUS white aio — hse green or рїп the tube 6 тт long, the lobes mx 15-6 mm wide, triangular to lanceolate, obtuse to ; pois 4.5- 5 mm long, with a anther. we Psi Шш positioned 2 пиг 5= 10m m long; stigmas Кайсариаа; 1-2 mm long, Figure 9; Хойго & Persson (1991: fig. 2С, 590 Annals of th Missouri жаай! Сагаеп D); Howard (1989: fig. 188); Lamarck (1792: t. 251, fig. 1); (1804: fig. 721); Steyermark (1974: fig. 21); Swartz (1791: t. 5, fig. 1). Habitat, phenology, and distribution (Fig. 10): Thr roast се ean and most of main- coena ч ted in fl 1 in fruit throug} the dn) n common and variable. It is dis- tinguished b its leat blades ш) acute to шу vena tion plane and usually not evident, and Е 6 rollas with tubes 60-125 mm ba and lobes 15— О mm lon, ements c yermark (1972, 1974) but not observed on material studied are designated with asterisks (*) in the description above. Hillia RANG fue in os Кан that eaf size m. ‚ but in general appears to eb correlated with habitat: нан from wet windy ridges and high montane areas tend to have smaller о than plants fro prominul m. plants from South и America, particu- y venation s on some с — dae others, apparently due as much to treatment of the ind imens with fo ee r alcohol as to any inhere feature of the plan The shape of de corolla lobes in Hillia par- asitica varies from narrow in u more Pas ln tilles and a with "^ other Menen "The lengths of the corolla tube a о bes оталу: iu wit rithin- -population range: plants from widely different sites. - Plants fro: ge о „эю горгона m other сахра, Occasional yx limb, as do other species of Hillia. Тера plants with iie short corollas and no c calyx were ere gat ated te у Steyermark as Н. та- guirei, bu ion locality i Jie net kuaing ikke plants h d this pecies is not maintained here. When present, the calyx 10. Distribution of Hillia parasitica Jacq. La circles) а in the Antilles and South America. e 4 in и рондо vary іп shape from er^ geograph n to b y oadly ligulate or orbicular i in some planis from ı scattered I localities in Amazon a pe sn este to orbicular rea ‘lobes! аз Н. ture and rela- 3 zi $ in the shi sis of the calyx species of Hillia from several vienes and Ee flowers fall within the normal r dis H. we asitica. No other features and from 25 to 74 mm in western South A ay Plants of western South America with relativ long floral stipules that tend to persist until a have been seg Hillia boliviana. How regated as E er, the size range for the floral stipul Е boliviana” overlaps that found in plants эп, ү: where, а as does the degree of persistence, speci ined here. ers, and form of the : b ilis peak n Сен gave the range of variety 100! Volume 81, Number 4 1994 Taylor 591 Revision of Hillia as “southern Brazil, Bolivia, Colombia, and possibly Peru,” with the typical variety occupying a more or less complementary range in the “Lesser and Greater и Mexico, Central America, and eae America (Venezuela and Territorio Roraima, Bra: zl." Unfortunately, the ra по of corolla am о FL | 1 Ј E- ts, ms the шш of ine. bracts Antillean quad and SET. of the measurements he Ја ve for these taxa are contradicted іп speci- ns that he annotated contemporaneously. The characters he used to кшен these Pree do vary over the par - om but КЕРА ^ do 50 ab ps the relatively broad corolla age fo und dn plants from the southwes tern p of the range of H. parasitica were probably ds primary ede ractice to delimit va d nobilis. 'This in cor cll lobe shape is continuous and uncorrelated with other fone ae variety no- ili ов P variation i ec may be some incidence of Hiep: or pi Los босу, a pm this has not been tes ed. His E ry the е Кылы ое top of the corolla » Similar to the variation found in other species 5 but the sigma are held above the anthers in all Specimens se n. The flowers are very strongly аха poe ap during the night, begi at ut 6-7 P.M. in Puerto Rico, but are odorless rudi the day Representative s pecimens examined. BOLIVIA. € mba: small power station c. mi. NE of bamba, Brooke 6700 00 (F, NY). La m below La; * N ‚ MO) ta Cruz: Santa Cruz, Williams 1478 $ ы: р. dE e : 8 of Morro do k n : own of Mdoch to W of road tinga, | t 355, guae Mariy 22773 (NY, US). eará: sitio pirito Santo, a da Ara —Maran- бер, Martens & Castro 7076 (М0), Espirito Santo: ta Leopoldina, cabeceira io : hus MO, o de Janeiro: region of Itatiaia, Ginz- ger273(5, "M 11504(F), Plowman 2862 (СН). eie de Lua, gins pd isi in 14"W, Prance et al. 9436 (F, NY). S. Mina Velha, Garura, € Francisco do Sul. ж eK Klein 5789 (NY, US). São Paulo: Boracéa, Salisópolis, Kuhl- mann & Kühn 1750 (P. "COLOMBIA сум — Cor dillera Central 7-15 to Nariño, 5°43'N, 75°15'W, ih & Cogollo 3564 (COL, ve Roraima: summit o yacá: геа western imt of Mt. Chapó Lawrance 35 1 (Е, MO, NY). Cauca: La Depresión, pe of жо Шу, Core 1327 7 (US). Ga dina marca: Cor- diller. ich di ais Rita, Pies T ы uen Аш жу et Me "0 (СО D). ila: Cordillera Oriental E of Neiva, Rusby & Pennell d: La 7389 " мо, о DOMINICAN f ' REPUBLI Bara- hona: : о, 4k del techs rural x de ‘Coke jo (*monteado Nueva 18°07' 30"N, 71°13! 30"0, ые etal. 18867 je Seibo: Los Hai eh tir rques еп Monte Zahon: et al. 35437A (NY al Cas: abito , Liogier & Liogier 215 4.2 km of Laguna, Sam: Rio م‎ n Portove! T di arrivi ev. Puy 54093 (A, F). Imbabura: Rio Li s, Cordillera | Occidental, Steere & Camp 8196 (F). Machalilla National Park, 1°36’S, à 0°42'W, Gentry et al. dd kia Pichincha km old road from Quito to Domingo, Dod. kE Dodson 11852 ү. ceni a tiago: : Plan lagro, ca. 10 km of Indanza, gom b m Su л road in ieie n between Santa Bárbara and La Bonita, Harling & Andersson 12475 Montagn 5856 ^ (MO, ND. 6215 T T ол. Morne Qua- qua, Beard 1225 (A, F, GH, MO, US). GUADELOUPE. Trois-Riviéres, Duss 2546 (F, NY, US). HAITI. Nord: Massif du Nord, Anse-a-Faleur, M e MARTINIQUE. De ain, Mont s 1462 (F, "NY, US). МО ONTSERRAT. Summit А Cone Pesk, English Crater, central они tain d & Howard 17553 (A). PERU. m E of La Peca, Barbour 201 i billo, de Jels 592 Annals of th Missouri “шаш Garden 12530'S, 73°30'W, Dudley 10414 (F). Huánuco: ditto. hinchao, camino a la Hacienda Derrepente, margen 1 (СО derecha del rio er Schunke V. 5317 LR m in: Chanchamayo valley, Schunke 90 (F). L аза, I 1666 (US), 2255 мо Chontabamba val- ne 75°30'W, Smith iscal Caceres, T s Seunke V. 8170 (F, М quillo Ate toni po US) C y, Rte. 184, Cer: sé Tali: 6353 (UPRRP); Ва an ales Pun. Torrecilla, Alvarez & Rodri; ount е: ae Pa: «и s Prov. Oax Sp: Ln. E me of L О, № аг: Аџуап NY), Karehe e p [dat . Falcon ЕМЕ Sani a yen. ta and ps Tra € Л amp: pä on road to La s, Steyerma rk 56156 (F I (3 orn near E end, above Qu son ties Sat Santa cruz, k f Саріга, 10°09'N, 65?48'W, Steyermark & Davidse 116874 (MO). Monaguas: ditto. Acosta, serrania del Turumiquire, altiplanicie la fila I ontana, cabeceras del rio Negro (afluente del rio p Chaba 5 (MO). m N of Salom, top of Са et ity El Am paro, yl et al. 20716 (MO, NY). 2. bus oe Andinae C. M. Taylor, subg. : Hillia ul Standl. A Hillia subg. Hillia Hoot linearibus corolla infra medium positis differ Calyx limb divided to base, lobes 5-6; corollas Кан, salve di white, lobes 5—6; st 6, attached ne le of corolla tube; anthers subsessile; stigmas linear, positioned below an- thers. One species, Ecuador to central Peru. re "eme резне не та Field = Mus., Bot. Ser. 4 . 1929. TYPE ы Fei qoe cienda Schunke, La Месе, са. 4000 +. 11290 т],27 ped 1 Sep! 1923, J. F. Macbride 5670 (holotype, F, photos F, NY). Suffrutescent herbs or shrubs to 1 m tall; bark o rounded at base, coriaceous to usually stiff; ins obscure, th or midrib some- axially, without EM mar- iol mm | positioned 2-3 3-4 oe stigmas 2-4 mm lo Er m including beak ca. 1 mm long, with stipes 3-7 mm Eg idi or low ridges; seeds ca. 3 with filaments ca. 1 cm long. Figure 7E-G. Habitat, phenology, and do ke 8) Ecuador to central Peru, in at 1300-1700 m. Cals in й іп ceni о Octobe by its relatively ades with the flower is de- 8 Б S س‎ Ф m] scribed for the first Additional specimens exam ned. ECU ntón Archidona, faldas al 5 del volcán Suna, 9 km al N de la anms Zak & )). Junin: Hacienda Schunke above Ramón, — A100 (F); Chan- chamayo Valley, PE 436 (F). 3. vice меш Tetrandrae С. М Taylor. subg. TYPE: Hillia берйр #7 Sw. subgeneribus ceteris Hilliae corollaris уре M atu 4- 9 et stigmatibus linearibus infra m positis differ x limb lacking or with lobes 4 5 corollas carnose, Pri ou prie fiam on surfaces exposed in bud, lobes 4; ma attached near top of corolla tube; an of po stigmas linear, LA ges below “middle rolla tube. e species in on Greater vn Mexico: rica. ca America, h ADS Volume 81, Number 4 1994 Taylor Revision of Hillia Hillia loranthoides Standl. . Habit with flower. —H. Stipule (floral) Ficure 11. A-C. Hillia palmana occ flower. —G. Flow opened. — A. Habit Ea Corolla opened. F-I. Hillia pa apsule. A, from Breedlove 57488 (F); B, from dig 3839 ). opened. — C. ur nen D, E. with flower. — В. Flower namensis Standl. — Habit with Фу С, {тот Williams m y; D, E, from жый. 19782 (Е); F, С, Н, from Dressler 1487 (NY). A-G to same e, Н to twice this sc. 8. ея оніо Standley, J. Wash. Acad. 165. 1 . TYPE: Costa Rica. Guana- caste: Quebrada SER SE of Msg ca. dus m, 27 Jan. 1926, P. C. Standley & J. Val 46152 (holotype, US, photo Hillia ma pa Standl. & Steyerm., Publ. Field Mus. Nat. Hist., Bot. 23. 1943. TY mala Quezaltenango lower S-facing slopes of Volcán Santa María hus S Maria de Jesús and Calajuache, along great barr ار‎ Finca Pirineos and San Jua s Pis cito 1300- m, 6 Jan. 1940, J. A. туеш. 33667 are Е, photo Ё). Suffrutescent herbs or shrubs to 1.5 m и ех &ray-brown, smooth. Leaf blades elliptic to what oblanceolate, 2-10 cm long, 1.6- wide, ac es acuminate at apex with ü mm cuneate to attenuate at base, 0 d riaceous А е: thickly so; secondary veins a r te, 3- irs, plane, med domatia; mar- ye "à. petioles 5-1 long; stipules 15-34 a ong, 8-14 mm wide. ers solitary; pe- uncles 1-4 m long; bracts lack IIO -2 mm; calyx limb lacking or eet to base, the lobes 4, 5-10 mm long, 1-4 e, ligulate ^ oblanceolate, acute; Gorilla чедип, white, tube 45-75 mm Ion g, the lobes 4, 15-25 mm long, 10-24 mm wide, P liptic с, UN anthers 4, 7-8 mm 94, — with м". pe 5-8 mm below top of corolla tube; ovary 3— m lng styles 13-15 в ап em stigmas a E M 1! mm li a Dike 30- is mm long including a bea 1 a m long, 5-11 mm diam., m: stipes, smooth; seeds 1. ie ca. 0.5-1 mm, with fila- ments 6-13 mm long. Figure 11A- Co Habitat, phenology, m ваи МАА 12). uthern Mexico to and northern to central Costa т in wet at 700- 2700 m . Collected in Sad = Jun us very کک‎ ша an acute-leaved variant of H. max Acide f bs f the plants of Mexico and Guatemala described as a from those of Costa Rica, and this i Th Н. macrocar e result tant to that of Psychotria (Hamiton, 1989 ). The report Mene from southern Mexico ayn, 1989) was a misidentification of H. lora 594 Annals of the Missouri Botanical Garden Representative agen examine ү” er Alajuela: La Palma de San Ram 839 (CR F), 3847 us T үзе ба т. Nw p p n Lent 1692 (C o, Santa Turrialba, Veri RTT (CR LH Mes pes River voy .9 km upstream from San Luis village, 11787 (CR, hic: GUATEMALA Baja a rapaz: en as Min m S of Purulha, 1600 m, Ac ia ‘a d 41 950 D ). “Huehuetenango km 335; of the Panamerican Hwy. E of the Mexican border beds silla, pra ders. 2 (WIS). Que- zalte: MEXICO. Chi iapas: munic ipio о La Trinit: of Dos Lagos above Santa e ue 1170 m, , Breedlove & Almeda 57488 (CAS, NY). O inity of Con- cordia, Montecristo, 1100 m, Maki 727 (US). 9. Hillia tetrandra Swartz, Prodr. 58. 1788. TYPE: Jamaica. Coldspring, O. Swartz s.n. (ho- lotype, S not seen; isotype, BM not seen, photo NY). Hillia tuxtl, ё & iño ex DC., Prodr. 4: 351. 1830, nom. inval., pro syn. Suffrutescent herbs or shrubs to 4 m tall; bark FIGURE 12. Distribution of Hillia loranthoides Standl. (upper map) and H. 1l. (low southern Mexico and | Central America. gray-brown, smooth. Leaf blades Ка: to a“ er map) in 3 7 5(-10) cm FIGURE 13. = ule. —Е. m F, G. Hillia 0 (Е); C, D, E, from Menéndez oie iie thee Mele, C 1b tek ea ps » F, G, from Maxon et al. 7501 (US). A-D, F A-E. Hillia tetrandra Sw. Habit with flower.—B. Flower opened.— C. S yv (floral. pev Sand. —F. Habit with йок. —G. Stipule bau pi бов орн oe Volume 81, Number 4 1994 Taylor Revision of Hillia 595 FIGURE 14. (-6) pairs, plane, without domatia; margins fla; petioles 1-11 mm lon, ng; stipules 8-30 mm long, 5-20 mm wide. Flowers tary: ко 1-5(-1 dh mm long; bracts lacking o 2 mm; pedicels 0-3 mm long; TUE а uL. ог DAL to base, the lobes 4, 7-19 mm lon : iet oblanceolate do ag rounded; corollas salverform, white, the 2 long, the lobes 4, 10-27 mm long, 8-2 ide, elliptic to b Page broadly roun ; anthers 4, sub: e, 4.5- -3 mm ү тор ol 9-10 mm long; ч 8-18 mm lon, sole ж felon ч wide, flattened, linear, positioned below middle of ا‎ tube. Capsules 25-80 пип lon k оч Б` & ii =. 5 оч e Ey mm, with fila- MOL Figure 13A-E; Standley & Williams (1975: fig. 14). Habitat, phenology, and bri med чы 14. Cuba, deben a iola, and so i moist to we E d e iat at Pr “1350 " 2650) т. сы ted in March to November, in frui throughout | the yea: equently bd species is characterized inii al oriaceous leaf blades MPH ridged capsules. Leaf size in plants from Belize onduras is frequently markedly larger than Distribution of Hillia tetrandra Sw. in southern Mexico, Central America, and the Greater Antilles. in plants from other areas di from Guatemala, нди апа Мехісо lack a a. limb; there is no intermedia o in O Hillia есе s and does Th ес s йн аз 100% ш e on owing studied, Top eg this puer lar gely "шша tetrandra is similar to H. maxonii of 0 A The latter can while H. maxonii is usually re wet forests on ridgetops 800 m, or from coastal mangrove formation: . Representative specimens examined. BELIZE. El o: Mt. Pine beoe banks of Río On, Lundell 6799 aranjo, Webster et al. “209 (A). untains, ^ C€""u-cUo0 C С С A SgEIEETCESTE uw 3 ^ xm n ahiz = Villa ire bii аф; Buenos Aires, Annals of the Missouri Botanical Garden Jack 6812 (A, DS, F, NY). DOMINICAN rane ie и en Se сме ate: за e = Haitises, entre la a de Arena, y Boca de Infierno. POST. be (E 30” "Ww. нан et аі. 35961 (NY maná Peninsula, Laguna, 15099 (GH, US). San Cri. i c tw gston an on N side, Steyermark 39464 (A, F). Peten: 5 E on Pusila River nd ge ntreras iu A CAS, MO), 8893 (CAS, MO). Que near аша igit ч po анов es ey ‘San =i Tajumulco, NW slopes of Volcan erii Sieyermar 36 731 I à E г e slopes of Volcan Santa Clar. f Finca El Naranjo, Steyermark Dooa т MAII Nord: Massif du Nord, e Ms Bonnet Levéque, al SO de Milo t, 19°6'N, 72714" W, Mejía Era aye: ASE, әне : Santa Cruz де Yojoa, ards 654 (A, US). S. arbara: near Mochito, Dickson teat (US). ТАМА AICA. Corn wall: Т. relaw nij. (DUKE, GH). бы: Жз jns, , trail fr rom Bath to ss Gap, j " the gap 3311(DUKE, GH, US). MEXICO ‘Chiapas, mpio. Осо ingo, 5 km SW of Santo Domingo, km SE of Palenque on road to Bonampak, Da ne et al "Mia QM O). Oax aca: road between Valle Nacional, 13 mi. bic Ma Valle National, 17°40'N, 96°22'W, Webster & Breckon 15378 (MO). Tabasco: mpio. Teapa, Puy- acatengo, Ventura 20644 (F, MO). Veracruz: municipio San An б a” xtla, Los — biological station, 18°34- 36'N, 95?04-09'S, Ibarra 202 (MO), 547 (NY), iti (NY), Tob (MO), 1984 мо, Martinez 3075 (F, Ee ا‎ at ber : Los £s гаа е 18 (MO). Zelaya: Сако Waylawas, 13*39'N, 84°48- 9'W, Stevens 7404 (MO). 10. Hillia "m queis J. Wash. Acad. Sci. 18: Mans E: Nicaragua. Ma- nagua: Las N ятан. чануб 5 of Managua, 800- 900 э 28 June W. К. Maxon, . D. Harvey & A. T ae 7501 (ho- Кур, US, o F). Cosmibuena rhi il Hist., meo Sa Ae TYPE: Colombia. Valle: еба ае Da 13 Арг. 1939, Е. Р. Killip 34972 пва Us photo F; isotype, F). Publ. Field Mus. Nat. Suffrutescent p or shrubs to 5(-15) m tall; bark gray-brow ooth. Leaf Pom "nee SECUN or E 2.5-10 cm long, 15- obtuse to rounded at apex, cuneate to attenuate at аур coriaceous to de thickly so eria ry veins pinnate, 4—5 pairs, Mor with- t domatia; margins fat: piis ga p mm long; pud 12-32 mm long, 8-10 mm E Flowers зо mm ке A= M eue lacking or 1-3 х 1-2 mm; calyx limb lacking Pig to tbe, ie e , 5-6 mm lo mm belo mm long; style ca. 15 mm long; stig- mas ca. 7 mm Mp: ae dens Mu ne of corolla tube. Cap ak 1-3 mm lo eget diam., not stipitate, ато) to fen те n s 2- 4 5 s 6-13 mm long. Figure ded c. abitat, phenology, and i Re ama, to Pana ma to o Ecuador Co le eted in loves April, May, Jul r, in fruit January to March, May to Ns emos and December. several hundred miles from those of centr tral Pan- ris (Vell. ) K. FIGURE 15 Taa of T illust Schum. (sol: id circles) and H. m i Standl. ( eode in Central p South pea rica (solid 1 Volume 81, Number 4 1994 Taylor Revision of Hillia ama and are found in coastal formations rather L ^ d Е Т. р 1 1 h fede or p mete truncate at spei cuneate to pin- collected in both habitats. No morphological ее f, | Mer ah a + | 1:6 +h 4 plants are evident. Cosmibuena ma (Benth.) Walp. a a similar E m اس‎ and set of hab Hillia maxonii is to H. tetrandra, and these species have sometimes been combined (Dwyer, 19 PBO): The distinctions рыс them : аге discussed under Hillia maxonii is also similar to H. palm of southern Central America; this latter inia ides tec ide: flowers and usually smaller COLOMBIA. Choe 5 Valle BAe Region: Alto dd d à Minen anuevo-San 76°25'W, Li 3 A " © 5 Estero del Cangrejal, entre los rios Yurumangui y Naya, Cuatrecasas a T (F, La Palma de S ntaren erde Bese serve, Haber 5 (МО), 170) 4501 (MO). Hammel & Ha ber 13933 (МО). S r Finca La Cima above Los fee of E] pia Standley 4 42599 (US), 42771 ЕА EC. ADOR. Carchi: 2 km below Mal- "Ww г ( . Coclé: western slopes and summit Ce е Chiquito, ae 495 (MO, US). Panama: Eod Jefe (Cerro Azul) 9*15'N, 79:23'W, Dwyer & auger 7377 (GH, МО) MePherson 6858 (МО), 7138 ‚ Mori & Kallunki 6500 (MO, NY). Veraguas: Р, of Santa Fe оп road past uate Agricola Alto iedras, Croat 23079 (F, MO, NY). 11. Hillia ia palmana Standley, J. Wash. Acad. Sci. 18: a: ip TYPE: Costa Rica. He- redia: inity of La Palma on road to La Ноза 1500- des m, 17-18 July 1923, - Maxon & A. D. Harvey 8045 (holo- sida US). Hillia depend subsp. grandifolia Dwyer, Ann. Mis- Ven i Bot. Gard. 67: 216. 1980. TYPE: "Paras Taguas: m W of Santa Fe on new road past agriculture heck 2900 ft. [950 m], M. Nee 9698 (holotype, MO). Suffrutescent агар or shrubs to 5 m tall; bark i -brown, sm h. Leaf blades к то оЫап- *olate, 2-7 с нере 1.5-3.5 cm wide, obtuse to nate, 3-4 pairs, plane, Without Doui margins flat; petioles 3-8 mm long; stipules 12-20 mm long, -10 mm wide. Flowers solitary; peduncles 1-2 mm long; bracts lacking or 1-3 x 1-2 m limb lacking or di deb to base, the lobe m long, 1-2 mm wide, ligulate, acute to rounded; corollas salverform, E the tube 35-40 2m ular, mm long, eens with vod potitioned 2-3 mm below top of corolla tube; ovary 3-5 mm long; style 15 mm t long; stigmas ca. C ы, long, linear. iddl Cap- sules s 6 cm cem including beak 1-3 mm long, 5-8 m сосала = шше, coir seeds 2-4 x 0.5-1 g. Figure 11D, E. Ha — phenology, and distribution (Fig. ). ral peerage to central Panama, in wet fort p BG 25 es m. Colle aie in flower in fruit рене to El iced and Y e October. is species is distinguished by its medium-sized leaf ыы es that are usually obtuse to rounded at the nd small flowers. 3 is ntur to Hillia Н. | 'M Aim treatments these р of the last two species. R T specimens exami ined. COSTA RICA. Alaju vc Vo "E cán Br San Carlos, seit pros 11095 с Volcán Irazû, те п pesas ^h (F). Guanacaste: e laguna reg Arenal y el Alto de La Carpintera 4 eredia: entre Limón: entre la quebrada Amubri, margen izquier 9°29' m те = 200 m е area 760. бо, "1156 ү prey м0) d (040) S, CR, MO), of Higuito, ca. 9 km Петра, Wi ilbur ж! ас СЕ, Е, MO, NY, 2 NICARAGUA. Boaco: al N de San José de los Remates, он. 85°45’ . "Moreno 24904 (MO), 99 (MO). G Finca San кары. riek 8510 (МО). Jinotega: ca m Hwy. 3 on to ар 13*02'N, 85*55' v D s5947 (MO). оса ра: ridge along road Danta and La Luna, 12*40'N, E W, esie 9616 (MO P PANA- s del Toro: trail fr uete to e Machin, ыйы BZA, McPherson & tiga 8328 (MO). nity of Fortuna Dam, 79°50" McPherson 10577 (MO). Darién: E ae. of 598 Annals of the Missouri Botanical Garden : ic pe of Hillia age ami Standl. ad а тар) and Н. grayumii С. М. Taylor (right- апі тар) іп esi America. erro Sapo, Hammel 1284 (MO). Panamá: E slope of Cerro Jefe (Cerro Azul), Tyson 3436 (GH, MO). Vera- guas: Cerro Tute, 8°32'N, 81°07'W, Knapp & Dressler 5398 (MO), Mori et al. 7559 (MO). 12. Hillia ee ноду, N. Amer. Fl. 32(2):ET0:51921. > аз Сатр, r. 1911, H. Pittier 3190 (даре US; aaa US). og e Standl., J. Wash. Acad. Sci. 16: 16. 926. TYPE: Mexico. Chiapas: ne сет Арг. 1925, о А. Риг pls 5 262 (holotype, wus Bg woe Hillia у ia кн Sida Sh is 9 . Her "i slope ks Velia Be. | ayl 965, "n H. Hatheway 137 1 (holo а Us Sues F, GH). Suffrutescent herbs or shrubs to 4 m tall; bark h. Leaf blades elliptic to nar- mm lo ide, 7 mm long, 1-3 mm wide, ligulate, rounded; c rollas white, salverform, the tube 24-35 mm v the lobes 4, 8-10 mm long, 3-5 mm de lan- ceolate, acute; anthe rs 4, 1-2 mm lon ile, cM y 2-3 1 tube; o -3 mm long; style 7-12 mm 1 stigm 10 mm long, linear, positioned below middle of corolla tube 5 2-4 g in- cludin ak 1-2 mm long, ca am., not - mm di stipitate, smooth or usually with ca. 10 tia ridges; seeds ы X 0.5 mm, with filaments 6— 13 mm lon 11F-I; Dwyer (1980: i 48, as Hillia aa Habitat, phenology, and distribution 12). Southern Mexico to Guatemala and Beliz and southern учее зыл, forests at 20-2300 Mire in flower Ds cem- rolla vee ilie stipules, mus cap: paile . The Ыл outhern Central Аена in ranthoides and species of Psychotria (Hamilton, 1989). Hillia panamensis is similar to H. palmana; the latter species has elliptic Eir corolla lobes 6-12 mm wide, and usually tise proportionally broader leaf Бл ёс er and capsules. Several sterile and e ing beo s Ho 1, d$. 4 t Пу БЕ S h they have salis breed leaves and ridged capsules. atively low Representative specimens riy ed. ledo: Maya Mountains, Hain 429. Alajuela; Cantón Alen 0 2909 uanacas! Dos de с еш T n 9 (CR), Gon Tet N, 84°55'W, 900 m, m et ad 527 3 (MO). imón om Eo Danto, 10°41'N 2 Se eet He iel 2438 (CR), Stevens & Venu, E ps MOL jy? 2 MO} GUA r az: between Finc ote near CERIS оета vie (F. US). Chiqu ixixi 3 mi. N o ? Steyermark ee T "NY. MEXICO. € -— 25- o de ار با‎ ‚ near Laguna Ocotal Grande са. ауе АА f Monte Líbano a. 45 km E of Oe Dressler 1487 (NY, US). Nica ARAGUA ^ MM 0-33'W, Ome саро, Volcan Maderas, 26-37'N, 85° Stevens o 18806 NM K 9689 ЖО, y A MO, 6552. (MO). P Border: Cerro ча, 8 km sc кеен d 0) im road on the road to Escopeta, Folsom 4 ama: Cerro Jefe, Hous 1021 (MO). ҮЗ е1: те aylor. . 4. wo subg. Illus ` . TYPE: Hillia € qud H Schum. ИШИ; Ili = 5- e en viridibus flavovirentibus vel co pureo AME et stigmatibus subcapitatis me positis differ p x limb eimi very reduced, or with 5- e atropur- a antheras orolla tube, middle or near top of c Volume 81, Number 4 1994 Taylor Revision of Hillia r with filaments well developed; stigmas capsules subsessile o; M s ioc qn above anthers; c usually sti Seven ресе E NU d cie South А Ric 13. Hillia grayumii C. M. Taylor, Selbyana 12: 137. 1991. TYPE: Costa Rica . Heredia: Finca là Selva, the OTS field station near m Viejo de Sarapiqui, near the junction e Rios Puerto Vie jo and Sarapiqui, 100 id May 1980, M. H. Grayum 2793 (ho- E. DUKE: балы, DUKE). Shrubs t ‚ or lianas; bark gray-brown, гн wi bades: бос, 9-16 cm long, 1.5- 6.5 cm wide, acuminate at apex with tip 1-2 cm long, a acute at base, coriaceous; EEE: ry veins pi = ec Ts | =. ide. Flowers solitary; peduncles ез 2-3 тт а fee 1-2 mm 2 тл мете ale green to yell w-green, the 3-50 mm long, the lobes 6, 8-9 mm lon ма, gae to rounded; stamens 6, the filamen: eal long, the anthers ca. 9 mm long, 2 Wm ا‎ Me, ~5 mm below top of corolla tube; ovary 7-8 mm 29 А йал. са. 3 mm long, pe Soo ae apst iles ca kd M ca. 12 mm diam., not Ren oot 7% ед 8-10 йү longitudinal ridges; seeds ca eee filaments 15—16 mm long. Sei et al. og 3). Habitat, 327 and distribution (Fig. Costa Ric. a, 1n. ica, in wet forest at 2 200 m. Col- ected in flower May and June, in fruit March. species is dist y its reduced e o Hi South Jew the dis- the treat- E: illia gra ayumii appears to Be uncommon an oe nf n of Selva понеке) ача vine it rapidly, but within th d ааа grew to flowering s rs was no longer Заги in the tit “Clare: me Pers. comm.; um, pers. comm.). Ane cPresentative Specimens examined. COSTA RICA. Juela: Monteverde Biological Reserve, Río Peñas Blancas, 10°19'N, 84*43'W, Haber & Cruz 8464 m rtago: 83°41'W, Lent 3703 (F, NY). L Pereira, n 47'N, 83°36-37'W, Stevens et al. 251 (CR, MO 14. Hillia psammophila Steyermark, Mem New York Bot. Gard. 19(5): 211. 1963. TYPE: 1100 m, 20 Apr. 960, J. рен & 5. Nilsson 392 (ho- ra EN). Suffrutescent vines, climbing by т» oots; bark smooth. Leaf blades elliptic to lan ca. 6 cm [^ the lobes 6, ca m wide, rounded; anthers 6 sessile, 7— with ie aam oned ca. 5 mm below top of corolla tube; ov a. 7 mm e stigmas ca. 2 1 subcapit: iti 11 li ly b Ї thers г 1 d € 71082. £ an E / ` о gee кю. апа ил. Oe zuela es and anthers cuni eta below the m. Hillia psam- a.45 m тори lai ws similar to H. illustris; he pci latter species. Hillia partici is also similar to H. grayumii of Сона Ric ca and may " coe sel related; H. 9-16 cm long, and its anthers positioned just below the top of the corolla tube. 15. Hillia illustris (Vellozo) K. Marti , Vell., Fl. Flum‏ 4 پک : Brazil. Rio de neiro area, Vell., Fl. Flum. t. 157, 1825, lectotype designated by Taylor (1 Hillia tubaeflora Chus, Linnaea 9: 260. 1834. TYPE: 600 Annals of the Missouri Botanical Garden A-E. Hillia ae (Vell.) K. S. E. Stipule isi A, E, from Dus 26178 (MO). A-C to same пеги T three times t cORQQ (h lot учеа potes F; CH, NYS мерем ). ll illiams & Cheese d& os = 7. 1928. TYPE: Trinida «i Tar Forest, 16 Feb. 1915, W. E. Broadway 7807 (lec. totype, rash by Steyermark (1972: 288), NY; isotype, TRIN not seen). Hillia кодо: Standl., Publ. Field Columbian Mus., Bot. т. 8: 338. 1931. TYPE: Colombia. Valle: Río Sucio, [s T J. Goudot s.n. (holotype, P, photo F; isotype, F). Sulfrutescent herbs, shrubs, or small trees to ou € stipules КЕС mm ges wide, thos "E Na rae 60 mm sae 6-18 mm (-3); peduncles 1-5 mm mm long; iod not seen; Meri limb divided to base, the lobes 6, 9-35 m м, often unequal by с mm thers 6, ут, long, with tips ро- sitioned a mm above top of corolla tube; m.—A. Habit E D d n 15539 (MO, B. from Vásquez & сеа. 4130 AMO, C; D, fioi ncarna: cale, 10 cm t with flower. — —C. Capsul E to twi: y 6-10 mm long ^ Boc 1-5 mm ы са. 2 mm lon сара pst immediately ey ein T sules with pedun- cles 2-10 mm long, (50-)85-11 mis long in cluding beak 1—5 long, 8- diam., supe 5-10 mm 1 mooth or usually ж mae! mm, longitudinal ridges; seeds 2-4 X M filaments 14-22 mm long. Figur (1988: fig. 6b, as Hillia tubiflora). Habitat, phenology, and ане er al South r De March o Jul and September, in fruit January to нр Mone coreg ; the latter he, БОШАП оё S po aeta broader leaf blade s with xm E nation. Hillia illustris also resembles ae pe i ot stipitate, iren "a ie en that йз ks d "10 ml and H. lees sie has smaller leaf filaments ca. 15 mm long, and anthers position below the top of the оода tube. Volume 81, Number 4 1994 Taylor 601 Revision of Hillia There appears to be no morphological difference tween plants of Trinidad and the mainland, and in having flowers bo: a e of three rather P E А similar in flower number in H. saldan- d tł variation, = ith flowers usually г ї rarely one or two, is found in riflora Therefore Hillia goudotii is not maintained here e authors (e.g., Robbrecht, 1988) have cor- ling of H. tuba shaped to * ‘tubiflora” rubs. tne but Фев prae his description on a n, and mo likely intended to apply el ie epithet. N of bridge over ve сах еу, 1 67*03' W, da 13957 (MO) "E 5. : Sete Varras, S entrance to Carlos Botelho е Park, 24°22'S, 46°55'W, Gentry et al. 59 2 COLOMBIA. Amaz а. XS Боген jo Paporis, entre el río Pac rio ана nananari, Sora ame а & CAE pies ie NY). Chocó-El Valle Border egion: carreter anuevo-San José de Palmar, Alto de Семра, Forero et al. 2868 (COL). Vaupés: Mita and vicinity, Rio Vaupés m below Urania, Zarucchi et al. 1885 (COL, F, GH) CUADOR. Morona-Sa ti ision Bomboiza, biológica Jatún Sa carr naca hacia oem Palacios & Iguago 4433 rites -— . nne: region de la Haute Armontabo (Bas Oyapock), de Granville 4362 (COL, P). Sai i: Rivié i bens zco: елы а Га Соп са. ] hour up Río Mapitunuari fre ‘Rio о Apu- es ac, Luisiana, Dudley 1 1426 (F). Huánuco: provincia ncio Prado, distrito Rupa Rupa, al E de Tingo Maria ro Quemado, Schunke 10292 (F, MO, N ). reto: provincia “Mayas, S, rand d Alto Nanay, along a de Nan Ум. pr ren OL. MC 20'N, 56°29-49'W, pens p ai Soo. MO; AY). VENEZU ELA. Anzoátegui: a n NE Tgantin, wee eg m. pe pea summi р, facing basic , Hato de Nuria, iplanicie de Nur куерта}, 88535 "d e Amacuro o Diaz. P. e Anto! , Сайо на arcano et al. 70.2.77 (MO). 16. base dun K. Krause, Verh. Hor i enburg 50: 97. 19 um. as the type col- lection, but this seems to be an error Hillia viridiflora Kuhlm. & Silveira, Arch. Jard. Bot. Rio de here 2: Dt. 4. 1925. TYPE: Brazil. A ‚ Ule RB-15737 (holot sotype lestroyed, из Е, мо, Hillia irwinii i Steyerm. ‚ Mem 287. 1972. TYP "Brazil. ip R о Jari near Ca- hoeira Маарага, 09*53'N; 53°21" 200 m, ug. 1961, Irwin & W. A. Egler 46683 olotype, NY; кое F, GH, NY, US). Hillia schultesii Stey , Ме X Bot 23:287. 1972. a :C mbia. Amazonas- Vaupés border: río AP entre el е Де or Kananari, uvis del P. ,:290 . E. Schultes & E. 13075 icd fj photo NY). Suffrutescent herbs or shrubs to 2 m tall; bar gray-brown, smooth. Leaf blades elliptic to SD тарин us; secondary veins long; stipules 5 mm dua: Flowers EA n 1.5- 2 mm fas nii lacking or 2 to base, the Ry x id 8- 15 mm long, often 1-2 mm =” unequal by 1-3 mm, ligulate to triangular, acute to y е long, rounded; anthe with in positioned at top of пиа үш; оуагу 5-10 mm long, stipes 8 10 mm long, subcapitate, positioned ен above anthers. С 1.5-3 mm wiin реа е 610.2 ст СЕ separ beak 1-5 mm wane 8-10 mm diam., s 8-15 mm long, sm seeds 1-2 cpi et m, with filaments 112 mm long. e 18. Habitat, Phen nology, and distribution (Fig. 6). Spora йанды tropical South America and eastern Pan in wet forests "i 0-900 m, most -— д wis rivers. Collected in flower March to May, July, and August, in E January, March to May, July to September, and November. 602 Annals of the Missouri Botanical Garden . A-F. Hillia ulei K. Krause Capsule.—F. Se "s dab — D, from аы 13075 (US). A-C, Е to same scale; D, Е to This species is distinguished by its subpalmate leaf — otherwise i in Hillia found о ana (subg. о geo- al species rec- sed only on on wide 4 diferen nces in sizes of on its дны size, 3-3.5 cm арн vi in contrast to fs ong by 22 in H. ulei, and its “pie iae ct acute corolla lobes 5 mm wide, i trast to lob 7 mm кие еш Н. ulei. n available to characterize H. viridiflora appears to be immature, and thus probably not fully expanded 5cm with flower. — B. . Flower . Habit ed. A, D, E, F, i McDaniel & Rimachi 25763 a n from ii mes 2 p An twi ice this in 1 either the tube or the throat. The geogr —- unction between the type locality for H. p o doubt influe ever, nolle bro dw, many living co Amazon € to кои іп ће ки 1X4. 5 de Janeiro Silveira stat- plant ap- disap- eie description that the peared Mine mysteriously, lowered, anı peared fro н) Botanical Garden and was seen ни ries Teja ile Hillia irwinii based оп not its ten calyx lobes and corollas ca. 5 cm long т marginate lobes, іп contrast Hi se lobes ve 3-4.5 cm ng with t deir and corollas obes in his ci me rolla of the type collection is 43 ong, ب‎ Ila lobes of this specimen are го vial than emargina e type collection of H. i we has ten c d corolla lobes eight collection has nine, in contrast to seven to 4H calyx and corolla lobes in other pepe a ulei. However, calyx and corolla lobe n Volume 81, Number 4 1994 Taylor 603 Revision of Hillia FicuRE 19. A. Hillia costanensis Steyerm.—A. Hab with flower.—C. Flower opened. A, from Bunting 1 4483 (NY); in most species of Mila; and the "elici in number seen h esci of these speci | rmark m H schultesii based on its a ША blade: -11.5 cm long in contrast to 3.5 ulei, and its S acutish" calyx obtuse" in Н. ul to Er obtusely angled міг sb H. шей vpn mau number available compared to its ary aM large pen phic ra а зерага of H. schu ltesii а у on these features seems unjustified. $ Representative paa s TPT BOLIVIA. La d a ташу Yungas 5 Bopi, cia Bar r- am^ r Calisaya, Кодо 101 je F, MO). BRA- Bc Acre Cruzeiro Sul, Rio Jurua and Rio Moa, x EM manha, P12811 (NY). ECUADOR Аар io Lagarto Cocha, near Redon ha, 0°35’S, БАД Lawesson et al. 4442 ). МАМА. lo om 1 (AAU). PA ên: Sumacate along Tuyra River, Pittier 6594 (US). B. Habit B, C, from Glaziou 12790 (F). All y same scale. it with flower. B, C. Hillia saldanhae К. Schu uitos, Klug 1182 (F, NM 5 Madre Dios: Par N de Manú, Ке Mani, Кар Са Y Station 1 11°50" S, 71°25'W, Fos- ovincia Mariscal 667 (MO, NY). S rtin: pro Chiles: distrito Toc ache rris evo, quebrada de Almendras, margen derecha m río ы, иш 4458 (COL, F, GH, prin US). S kerie: ner slopes id Frederi A 3 km S + нА Top. Wilhelm Gebergte, 3°36-41'N, 56°30-34'W, Irwin et al. 5494 (F) 17. Hillia egent K. Schumann in Mart., Fl. Bras. 6(6): 2 ‚В . 1889. TYPE: Brazil. Rio de Janeiro: No и Glaziou 12790 ones B iste ee F, GH NY; isotypes, F, P n en). Shrubs; bark red-brown, met Leaf blades quss to dien na 6.6 cm long, 1.4-3.2 cm obtuse to rounded at ap cuneate to acute at Viene e, y veins pin- , 7-8 pairs, pes without domatia; margins Pi а 2- 7(-10) mm long; stipules 12-15 ong, 8- mm wide. Flowers solitary(-3); mm peduncles 1— ; mm long; bracts not seen; pedicels Annals of the Missouri Botanical Garden В, С, 20. A-C. Hillia foldatsii Steyerm.—A. Habit with flower.—B. Flower opened.—C. Capsule. A, ii "Holst 3772 (MO). All to same scale. 5-10 mm long; calyx limb divided to base, the lobes 5, 9-14 mm long, triangular, acute; corollas narrowly funnelform, the tu 0-35 mm lon; mm long, suborbicular Hiis td ovary 2-5 mm long; stigmas 1.5-2 mm long, Ft d immediately above anthers. beak 1-5 mm ca. 8 lagi tudinal peo seeds with filaments ca. m long. Figure 19A-C. Habitat, phenology, and distribution jet 6). Southeastern Brazil, in wet forest at ca m. ае month unknown. This е has es d ш HOY with Hil; om H. para rounded | rather than acuminate leaf Sedi tree f tube. It differs from other species of subgenus tinctions between them are disc ent of the latter species. The corollas of the e died: measured here, appear to be imm: 18. Hillia — Steyermark, Mem : York Bot. Gard. 23: 290. 1972. TYPE: Ver B d: rte noro eh танй sat Angel, 1200 m, s 7192 ell VEN). Shrubs m tall; bark red-brown, E Lea blades E 3-6.4 cm Ere 1.5-3. und- wide, obtuse to rounded at a cuneate to го ed at base, wäh coriacea: ea pinnate, 2—5 pairs, plane o "m nulous abaxially, w ces Ba oa flat; petioles 3—6 ng; sti 9 ca. 8 mm wide. Flowers solitary, 5b with peduncles 4—9 mm long; bracts 2—4. 2. 5 mm; calyx Volume 81, Number 4 1994 Taylor 605 Revision of Hillia limb wir to base, the lobes 6, 13-17 mm long, wide, narro did apri to hA acute; sa funnelfor h pink, the stigmas ca. 1 т ng, subcapitate, positioned 4 5 mm above top of corolla tube. Capsules 65-92 mm long ке “ae long, 9-12 mm ong, smooth or n be vies seeds 2.5-3 X 12 mm long. Figure 20. сету phenolo, ogy, and d He antepui in southeastern on roc in wet aen at 1200-1800 m. aee in E and fruit in March and August. This species is distingu аы by its thickly со- riaceous гаа f blades, six-merous flowers, and е "mid stigma: to Hs sa ШЫ with со T st is ап splitting along опе е ыз — 6, 3- IR mm long, often unequal by nelform, bright green to be -green, ‘the tube 51— 70 mm long, the lobes б, 8—10 mm long, rounded; anthers 6, ilios: 10.5-14 mm long, with tips positioned bove Лр of udo "on ovary Q | | NDS 1.5m m long, subcapitate positioned i imme: nediately above anthers. Cap 40-61 mm long i beak 1-5 mm Ыр. mm long, smooth with ca. 10 longitudinal lines ог low жыды seeds ca. 3 X үп m , with m inont 8- mm long. Figure 19A; 4: fig. Habitat, phenology, and esté (Fig. 8). Nor th-central hess to northeastern Co- lombia, wet for 200-1300 m. Col le cted in flower "Ma rch to Ма and July, іп ‘a. March, June, and September. This species is distinguished by its subsessile bl А s ro ounded ae ону: orda te with a leaven im the ers, and included sliginas, Seem digo in the original description that the filaments are 15 mm long, but this has no di bod n se Additional specimens examined. ZUELA. Bolivar: distrito Piar, summit of Auyantepui, 5°51'N, 62°32'W, Holst 3772 (MO). 19. Hillia costanensis Sarthe Mem. New 3: . 1972. TYPE: Ven- 5ай Luis, cerca del о, entre Curimagua у San Luis, О m, 20 July 1967, J. A. Steyermark 99274 (holotype, VEN; isotypes, NY, P). n. Missouri Bot, Gard. vals Hillia zuliaensis Steyerm., А: 1 17 y Ж“ -{ 985. ТҮРЕ: tie vicinity of Rio Guasare between Rancho 505 and Cerro Yolanda, 10°53-56'N, 72°26-28’W, 200- 270 m, 29 May 19 80, J. A. Steyermark 122876 (holotype, MO). i Suffrutescent nate or shrubs to ca. ark gray-brown, smooth. Leaf blades TE ped g, 6-13 cm long, 2.3-6.5 cm midrib sometimes prominulou dil viia domatia; margins flat; petioles 5-3 mm long; stipules ca. 15 long, ca. 6 mm mm mm wide; calyx limb with tube 33- 56 mm long, sometimes Both of these features 5 are unique in Hillio. E ermark distingui ished Th шшш ga H: and reproductive features and disjunct dp hic ranges. However, with more material available the variation in size an » the geographic range seems ntinuo GLORIA. Gua a de la ce ichuao, cerc del dtd то Каа СЕТ e Sugden 85 сог Magda- Parque Nacional Natur. Tayrona, cerro El ls yasa & Barrera 3911 (CO D. VENEZUELA руне Girard Parque Na- cional Henri Pittier, E Cumbre de ncho Grande, 10°21" N, 67°39'W, Davidse а al "672 з (MO; eds er 70 TN. Banting W of Gua Lara-Zulia с m al E de E arriba de la i т, 10°10'N, sa: 30 km of Corpo Zulia Cam 72°20'W, Steyermark e gg 123255 (MO). 5. Hillia деч a Oerst. С ы о: (Oerst aylor]. species, southern Mexico to Venez uela. dies ( 1989) deed this subgenus and Te added two new species (Taylor, 1991), wi 606 Annals of the Missouri Botanical Garden of in geographic range. The individual species are therefore not described or discussed here. The geo- graphic range presented for Hillia triflora var. triflora by Taylor (1989) should be emended to [SE E i4 th Mexi thi | RUSO ешн A REY Ө ДЫЛ red g sp of H. MU DAE lin thi l are Н. allenii Seem. Н. ngifilamentosa (Steye rm.) C. M. Taylor, H. oaxacana C. M. Tay- lor, Н. rivalis С. М. “Taylor, and H. triflora (Oerst.) C. M. Taylor. EXCLUDED SPECIES Hillia chiriquiensis Dwyer, Ann. Missouri Bot. Gard aS 1980. = Co P valerii (Standl.) С. Jh Taylor (Taylor, 1 : Hillia ا‎ = I Ann. Missouri Bot. Gar e osmibuena wes Eo C. ES 92). наша. og for es Fl. Filip. 235. 1837, not Hillia lytranthe ampullacea (Roxb.) еф (Ме iil 1918), Ew. naire Н ,in tte, Fl. гаи 8. 1847, not Hillia longiflora Sw. A 3o нше prasiantha Lem n Houtte, T g Ar bod v E zeylanica Thunb., Hillia t valerii Standl., J: Wash. Acad. Sci. 16: ea 1928. 1l.) C. M. Taylor (Tay- on; 1992). LITERATURE CITED ANDERSSON, L. & C. PERSSO ircumscription of the tribe Cinchoneae aces b. cladistic ap- proach. " Sy pe Evol. 17 D. H BE F. ali за Rui biaceae. /n: 51 b de s othryx. Acta Bot. Aca BREMEKAMP, C. 19 "Ren n the position, delimitation and M subdivision of i. Rubiaceae. ot. Neerl. са "B. А The. sr group of the paleotropical = Ar rgostemmatea энде a Cladistics 3: 35- s „== == Ro JAN Comparative restriction site mapping of pus DNA implies new logenetic relatio. nshi ips within Rubiaceae. Amer. Bot. 78: 198- CANDOLLE, А. P. p 830. Hillia. Prodromus sred matis Naturelle Regn es 4: 350-3 Treuttel & Wiir Dwven, J. D. 1990. Жене п: R. E. Woodso Jr. & R. W. ru & С de Ж Pan: ama— Part IX . Missouri Bot. Саг 529. EwEL, J. S. & J. WHITMORE. 1972. Ecological life gin Islands, zones of Puerto Rico and the U.S. У 3 Forest Service Ме. Рарег U.S. Dept. Agric., ARRIS, " S. 1988. Hennig86, Version 1.5, Program and User’s Reference. State Univ. of New York, a my Br raj ишү, C. 1989 9. A revision of Mesoamerican ds tns i ERN ا‎ o ). Ann. Misso NERA 386-429, hae -916. 190g. vers In: Fl pie d indward Viel 470. Arnold Arbore Sn. He arvard ун бо ‘= maica Pla x Massa oe KiRKBRIDE., J. H., JR. 1968. manian species of дик etia. Ann. Misso Gard. 55: 372-39 KiRKBRIDE, M. C. С. A preliminary phylogeny for the neotropical pure Pl. Syst. Evol. 141: 115-122. Lamarck, J. L. M. ry revision of the Pana- uri Bot. 92. Tableau Enoeet et Méthodique. Botanique. Chez Pancouc te Par LoRENCE, D. 1991. New species а ae Mexican ME Central American "Randeledié mee ace " Am т? 185-151: MENA, P. A revision of the genus Arcytophyllum aca: Hedyotide ae). Mem. New York Bot. Gard. Misi: T A 1918. Nat. Resources, Bureau MOORE, S. LE cs In: W. Face & А endle үз з Flora d d E rn 1825 ee same (Natural Histo- Species blancoanae. Dept. А; E of Sci. p #12, маш R Кошык, E dom. E endi Rubiaceae. Opera Bot. Belg. OEMER, J. J. 17 A pee de а tis Hispanicis, Lusitanicis, Brasiliensibus. ae & J. Porte emburg. SASTR P de ntilles. nee Caribéenes, aris. 889. Hillia. In: C. Martius, Flora Cien O 20 200- viia A. Engler & К Раі MOS Die Natürlichen а nfamilien 4: 1-1 Wilhelm Engelmann, Leip 1804. Hillia бау Bot. Mag. 19: fig- a y 1814. An Introduction to Physi pases and tomate pone Bradford r ai eu STANDLEY, Р. С. & L 2x Gua temala— Part i (Rubiacese). piti 2 . STEYERMA M EZ Hillia. In: B. Maguire & | f the Guayana guire e & Collaborators, The Botany o 11: Highend Part VII. Mem. New York Bot. Gard. a. In: B. Maguire & Collaborat a The Botany oft e е Guayana Hig = Part IX. Men ork Bot. Gard. 23: e Racine тту (өши ), Flora cción i eer Instituto Botánico, Сага Swartz, О. 1791. О Botani Erlangen cae. J. J. Palm Volume 81, Number 4 1994 Taylor 607 Revision of Hillia Sworronp, D. 1990. PAUP— Vries egal Using Parsimony, Version 3. ak and Use М. Guide. Illinois Nat. Hist. Su Urban TAYLOR, С. M. 1989. Revision of fitta ibd т (Rubiaceae: ao e). Selbyana 11: 26-3 1991. new species o of Hill ia subg. Raw nia Гену from Venezuela € Mexico. А Missouri Bot. Gar: 1 1992. idis ud aM (Rubiaceae). Ann. Missouri Bot. Gard. 79: 886-900. 1993. Hillia. In: G ors g & L. Andersson, I of Ecuador 50: 02. Distributed by the for Nordic Раана in Botany, Gothers- M 4 Coun gade 130, coe 1353, Copenhagen VERDCOURT, B. 58. d. Bot. Ae 28: гн "290. in D. idson & rica. Кшышсайе (Part 2). A. A. Balkema, Rotterdam. NUMERICAL List OF SPECIES OF HILLIA = mE So £ > 0 Сол ом; =; . 5 Ру . . m = = = ю ia s жаза ЖЫШ С. М. Taylor А ла ei Standl = м‏ د۵9 ص سے سے سم > ® 3 H‏ ubg. Tetr andrae C. M. Taylor ` loranthoides Standl. a Sw. > . H. pa лапа. lillia уклы Illustres С. М. Taylor B a . ulei ы к= – ы жы н ш اسم‎ л =! = és о HE S cana н Н. ii Ste uH, longifilamentosa pem M. Taylor H. rivalis C. udi deno or 24a. H. triflora O triflora 24b. H. triflora var. aye (Standl.) C. M. Taylor NUMBERED COLLECTIONS оғ Нила EXAMINED Specimens are listed alphabetically by principal eed tor. Identifications are indicated by the numbers llowing th Th .D. I1 9). 3682 6 6034 (9). Allen, P. H. 1872 10) с 0). Alvarez, H. J. 24 (6). Anderson, W. R. 331 У 35844 (6); 35914 (6). Antonio, T. 5078 (10). P i 4239 (6). Arnoldo, Fr. M. 3302 (6). Asanza, E. 41098 (15); 41192 (15). Asplund, E. 10160 (2); J. H. 5185 9. eh 09; ee (95 ир (9). вм апр x 132 (6); 395 (6); 540 (6). Be 4 (8). Be 460 (6) а Ash 1110 (5s. res б 1228 (o is vides 02 (2). Be eem 19). а 5 е 3). Bern ae vit 9) 8 5 (1 x td P. 3468 bee Billiet, F. 1916 05) Blackmore, S. 4104 = e La T ). Boldingh, I. 21 "s ^d 41 ( oe 901 (6); 6933 (6). Bo ге т oe 4 Fernandes, Н. Q. 1830 (6); 2280. o Brade, A. 0 (6). Brandebyge, J. 306 PAL weed uu 33962 (1 6); 42671 (15). Bresolin. (6). Breedlove, D. E. 31213 (8); aaa, Hes ty pi 48354 (9); a (9); 53555 (12); 56648 (12); 57488 (8); 68610 (8). Brenes, A. 3839 (8); + ); den (8); ^. : 9 (6); dus ai 924 (6); 2020 (6); 2203 (б); 2591 (6) (9% ET T ae (6x 5007 гох па sa ) 7 (15). = ont 09,1 de" Brooke, S S чз б 3 7 (9). Buchtien, a 35 е soso ©, ‘nn G. S. MB (19). Burch, D. 1371 (6). B 6-383 (6). Calzada, J. I. 6 FH pent 7144 (9). nae (9); oem (9); "gh (9) 8893 (9). 7 (6). Cosentino, К. 21 (9). Correa, ve E 94 41 010; 1770 (12); 2559 he 1). тлек . Сгоа В. 14801 (10); 20709 (15); 21093 54 ads. do; 26 ne Wm д yai 63462 (8). Cuatrecasas, J. 12 ok 107 (10); 19988 (10); 23963 (2). Totis T 2 (6). Davidse, G. 16723 (19); 20416, ©; 20716 [3 24028 i» nips (11). De 9 (6). Devia, W. 2 у: 7A 4184 52 (9). Dod: 1875 A vies (2); оз 0; 10195 “ee 10285 6% 11852 (6). Donselaar, 15). Do 6 it 1487 (2) 3522 (il). ee W. В. E-553 (2). Dr Nd qno dT (11). Duarte, A. P. 6257 (б); 11481 03 11570 (6). Dudley, T. R. 10414 (6); 11426 (15). Dusén, P. 8109 (6); 8689 (6); 8881 (6); Тууу (6); Veteri 16113 (6). Duss, Pére 342 (6); = ey n 46 (6). Dwyer, J. D. 7377 (10); 10022 m Edwards, J in ds Pages B. p ). Egler, rpg .L. 231715 yr (6); W.A. 46683 7867 (9); 9989 Ori (15). Ernst, W. R. 1 W. ү 393 (6). rreyra, В. 1666 (6); 2255 (8). Folsom, J. P. 4785 озу 9807 (11). Forero, - 2868 (15). Fosberg, F. R. 339 e are: R. B. M. 110 (6). Ga a v^ a L. 167 (6). Eyerdam, G. 204 (2). awe H. 5 (10 v udic 625, 6 be ce ug Ag" ie o 4847 (9) 4930 (9 mS 9 (9). Ge (2); 35166 (2); 40719 (10); Ban (6); 50447 at A (6); 50959 (6); 50959A (9); 52091 (6); 59053 (15); 2 608 Annals of the Missouri Botanical Garden ен du. eh nin 12561 (6); 71121 (9). Ginzberger 3 (6). Glaziou, A. = «s (6); 12790A (15); 13943 95 (11); 11302 (11). G oes 63 (6). Granville, E ]. dé 4362 (15); 5856 (6); 6146 (15); 6215 (6); 6519 (15); 6797 (15); 8563 (6). Grayum, M. 2793 (13). aber, W. 384 (12); 562 (12); 571 (10); 760 (11); 1156 (11); 1342 (11); 1787 (8); 3671 (11); 3672 (12); ); 4530 (11); 4564 (11); 1); 7000 (8); 7083 (11); 7334 co 8910 (б aos Bx. ; 8464 (13); 8914 (11). Lis 9 (6) H | ay + ЕТ My dign io 13930 (11); 13933 10) 17336 X НАТАН arley, R. 9309 (6); 19684 (6); 20860 (6); 27773 (6). Harling 3554 (6); 11639 (6); cesa (6); 24490 (3). Harris W. 5465 (9); а (9). эе n G. 13342 (6). Har € sr TT petes Vise . Hatheway, W. H. 11).H 205 14528 (6); 18509 = ek. (15) pes vm امام‎ (6); 28833 (6); 29669 (15); grape besiege (6x E ur Д, (6). Нау- деп, ч e IR 2 (6). Henrich, 35 Б. үе m. Hernández = ses ); 1244 (9 E Her 6 (9); 13 poe Herrera Ch., m. pes (8); нази T (2); 3910 (10). "i enheide, H. e picts ere eed (6); 986 i ims (9). Шш, Вго. 9 (9). H (6); 1802 (6); 1663 (6); ee (6); m (or 31 10 he Hoehne, Е -Nielsen, L ae uw (6) veu ven = нн S pt 2. 4 (6 xd 6 (9); 14290 (9); 15540 on 16052 a 16093 Я 16098 Anu iot (6); 17553 oe 5 (6); 1 ghton, W. M. 1194 (6). уне Mud. D. R. nu (6). e G. 202 (9); 547 (9); 1025 (9); capes 24 (9), Idrobo, i ч (6). ie G202 (8). INBio-I 181 ms „ЖШ: P E 30209 (6); 54944 (16); Beebe (5): see б P Jack, J. С. 6812 (9); 7211 (9); 8572 (9). eg a (6). Jaramillo A., d 22 usson, iL 62 24A dh Killip, E. P. pe (5); 34972* (10); Had ees Kirkbride, J. H 58 (10); 2398 (6). Klein, R. M. es Ан К, a ign m. Klug, G. 1182 (16); 3 Kna . 2159 (10 2621 (10); 4161 (11); benc pu 5398 (11); inr i T "1 P. з= 10171 (16); 10 MM IM т uhl- *( 0 (6 mann 16). Kuhlmann, M. Kuntze pA M "rr Landrum, L. R. 427 Mes ) Lanjouw, J. 628 (15). Lankester, C. H. 1935 (11). Lanna 1922/1463 (6). wesson, J. E. 44288 из, 44317 ы T (16). Lawrance, A. E. 351 (6). Lawton, R 4 (11 ^ Lehman BT er (3). em pé C 28681 16 t ret 3 (13). I Bro. 6626 (9); 7854 P^ 19805 id: p T i. . Lewis, M. 40471 (16). 9 (9). Liesner, ч; 8170 (6); xs (6); ve rf e 18431 (6). Lindeman, J. C. 13462 (6). Linden 786 (6); 2169 (6). PER A. H. 11207 (6); 14372 (6); 17484 (6); 17770 ө, 18421 (9); 20366 (6); 21526 (6). Little, хә ae вто 13094 (6). lid, F. Е. 164 (6). L (9). 2 (6% ne (6% Eon йэ; ir 9 Lozano C., С. 2 Meo 2 Lu ай Het Q:L 6 y 6372 ‘on hee ae 16 e MO ч ve 2 (9 9). meyn, TE Nov 8738 (2); 11542 (6); 12690 (10% 12 8 (6 Р. J. М 811 Hee Macbride, J. F. 5 (7). "ы gal, " M. Miu (2). € = К (9). Madison, М. Т. 4519 (2); 4700 (10). Мар en 24009 (15); 28522 (6); 35287* (6); 35296 0. M (8). Marcano B. 70-2 192 (6 | С. 9593 (6). Martinez S., E. М. 7969 (9); 15074 (9); 15574 (9); 15589 (9); 16660 (9); 18806 бы ida Vae 22922 ior 22953 fone Martinez аз UR n, W. (10); 10577 (11); 11846 (10); 1340 . Meclinors 35 (15). Mejía, М. 299 (6); 4995 (6); 23929 (9); 35780 (9); 35781 (6). Mello B. 3605 (6); . Menéndez L., F. 85 (9 P. P. 16513 01); rm 19112 (9); тези 19852 (12); 23735 (9); 24904 (11); 24936 (11). М (11); E es A 6500 (10); 7559 (11); 13292 (6). М „ Mort А мш 3 (6). 24 9 хер р. 382 (11); Oldem: 8 (6). Orozco, J. M. 639 io) Totg M. 2669 (2); pe 1: ) - alacios, W. 4433 (15); 6540 (5). Pennell, F. W. 5890 (2); 5898 (6). Pinto E., P. 623 CS) ed 3190* (12); 6594 133 (6). Pounds, W. Z. 221 (11). Po а 1). Prance, С. Т (11); 403 (11). Proctor, 22567 (9); 32634 (9). Purpus, 262* (12). БУ bor n, ut P. 488 (6). а wt qon (6); 2 6 4756 : A 2058 (6); 5789 3 (6). Reitz 1825 (6); non ] sm Pw Aron 6 (9). лаа 449 (2). К (16); 85 8 (15). Riedel 124 0 (6). К (2). Robleto, W. Ap д еы (6); 4460 (6). fat: С Roos M ШУ 7027 (6); 7803 (6). В af N. TT (9). R 6). Sagástegui A., t 09 amp, A. 6896 ( chez, V., J. G. (1). Sandera C. 367210 уз кы Е. s B 2: бу 590 (6). Sastre, С. 1934 (6). peas W. A. S-585 ( eem R 0 (15); 13 ith, D. N. 4139 (6); H. 725 (6); 1654 (6). Solomon, J. C (15). Spellman, D. 1639 (9). Spruce, R. 5079 Volume 81, Number 4 1994 Taylor Revision of Hillia (2). Standley, P. C. 38472 (10); 42187 (10); 42599 00 42771 ао; 44733 (10); 44738 (1 Ik 46152* (8; 0144 (10); 50691 (10); 52736 (9); 54583 (9); 5558 ^ 68706 (9); repe 84893 (9); 91610 (12). Pes W. C. 8070 (6); 8196 (6). Stehlé, H. 187 (6); 334 (6); (6); 6 429 (6); 868 (6); 4688 (6); 6005 060 (6); 7406 (6); 8115 (6). Stein, B. A. x: x (6). Steinbach, J. 996 (6). EM = W. D. 5628 (11); 5947 (11); 6552 9616 (11); л (11); 11679 oem 23919 (125 24189 (12) O 120 (1 2 m 0 (6); 31627 (12); 3 0. XU o Son д poe (8 " 36318 (9); ore 6 4195 (12); 44435 (9); 46048 (9); 46606 T Pap d 54093 (6); 55808 (6); 56156 (6); 56591* (6) 59799 (6); 61433 (15); 62044 (6); 75974 СА 8052 ios 88835 oa Mis T mson, W. s Sur uu ). 2M d E (6). Sut 8 (9 0 (6). Taylor, C. M. let 10445 (6). Taylor, N. ate (9). A d I са Теѕѕепе, - 1377 (11). Tessmann, G 0 (6). Timaná, M. 823 (5) 827 (5). Ton, A. S. € ptt Tonduz, A. 17755 (10). Tuerckheim, H. von 11340 (9); 12254 (9); 7920 (9). Чулу E. L. 3436 (11). Ule, E. 4422 (6); 6305* (16); 15737 19. Uribe U., E чүл (2). Utley, J. 2609 (11); 5223 Valdivia Ae P. 766 (9); 951 (9); 1423 hi Valerio R., J. 1359 (8). ا‎ 4771 (6). Vargas, C. 15615 6). Vasquez, R. 4116 (3); 4130 (15); 6421 (15); 10091 (15); 11879 (16) vous 305 (8). worth, R. K. (6); 1680 (6); 1752 ( ebster, (9); 13239 (6); 15378 (9). Weddell, M. 1520 (6). Wendt, T5271 PE 25 y 5779 erff, H. van der 6722 ( B (11). Wilbur, R. L. 7389 (6); 7816 "t6 7845 (6) ; 7947 (6); 16929 (11); pou. aay d cared ч 99 (15). Williams, L. 13068 (9); 1 hyn 28129 (10); vor o е s, R.S Ure i 3 (6). Wilson, P. 172 (6). Winget R. 6647 (6). пае Е. 8236 (1). Мга л 25B (6); 266 (9); 1256 (6). Wurdack, J: Ld 8 (7). Zamora, N. 619 (11). Zanoni, T. ey Ms 18867 A jan (6); 21127 (9); 22508 (6); 24771 (9); 26660 (6); 35437A (6); 35961 (9). MEE dde pen (15); vipa (2); 5659 (6). SYSTEMATICS AND Ricardo M. Rueda? ELE OF THE GENUS PETREA (VERBENACEAE)! ABSTRACT Fourteen of the 35 previously accepted species of Petrea ree known only as fossils. Mor pae acea and ecological riu ble: of the species of Petrea are presented, along illustra usin, Ciharerylum ч and Duranta T^ E ‘outgroup comparison, win that Petrea is most соу related to Xolocotzia Mira resulting in a cladogram with a consistency index of 0.74 and a retention index of 0. 86, after three rounds of successive weighting RESUMEN El número de aceptadas especies en al finit Petrea L (Verbenaceae) fue reducido de 35 a 14; tres son sólo natomicas, у ec an E cas de las Peporige д Perea son presentadas, ibu Petr j lustraciones, treeae Briq. " se usó como grupo externo 5 Githarxylum n y ees L.; este análisi ló q Хорст Miranda es el mas его 0 Petr a. Xolocotzia fue t polarizar el estado e = EFE PS EE VEA Po 1 А А у de 0. 86. eme de tres id lorización de 1 Petrea L. is a poorly known genus of the Ver- collections and new species described since 1938 benaceae. The most recent taxonomic treatment have rendered the keys obsolete. was written by Moldenke (1938). This monograph Moldenke (1938) included 29 species in Petrea, has long been outdated, making — у of and six additional species were described later, one the species di ficult. A consi siderable namber g dis- as a preliminary part of the present study (Rueda, i 1992). Petrea, as interpreted here, ontains 14 n the keys and those given in the individual species, three of which are known only from fossils. ЕНШЕ e of the species. Moreover, additional The extant species are exclusively Bibs ' This 005,3 was completed as a partial severe of a Ph.D. degree in sets at the pero ane Se of Missour ic Louis. 1 lwyn Н. d the other members of my committee for guidan се ant 90 mments on eis manuscript. I — Richard "Кейш for the use of his laboratory ire at Sout ern Ошту for th SEM facilities at Washingt ton Univera ле Jorge Crisci and Pet i ble hel Га v i ohn Myers Bick x e skillful ae A Peter Raven ‘fellowship and a eaching ass antship from the Univers’ Miseo - Louis and the Comp ton Foundation рг rovided financial support for ‘my ‘studies, The uri Botani u t "d d ie foll grant N внес Logistical Support was provide in rica: n Botáni Pineres"; Jardín Botá joe oaquí n Antonio Urbe": Instit pe ie ee м de ae Universidad d Nacional d de Colombia; Instituto Nació] de eum isas da Amazónica, Manaus; Mobil Oil, Bolivia; and Fundación Jatun "m re grateful to the curators гар се of the v folowing Tala for loans of material: „Аз, C G ' д УТ. LE, NY, P, PORT, PR, R, S, SP, TEFH, TEX, U, UB, U C, UCLA, UEC, UPS, 3, VEN, W, WAG, se а Nacional Autónoma de Nicaragua- yea Facultad de Ciencias, "Departamento de Biología, León caragua ANN. Missouni Bor. Garp. 81: 610—652. 1994, Volume 81, Number 4 1994 ueda 611 Systematics and Evolution of Petrea | е Sew а. УА | © o and savannas and on soil types s uch г as white sand ne. The species range from тп h Indies to Bolivia, Paraguay, and Brazil, being most diverse and concentrated in northern Brazil and the Guianas. Fossils a f the f Сна а апа Аан One spe- cies is widely pure vated in warm and temperate regions of ilie etrea possesses d distinctive features: usu- us Xol а а к», is 0 defined by ic presence of calycinal c History oF THE GENUS st known reference to a plant ii to an was in 1736 when Philip Mi . He name us e,a Been English po lector m living plants (Moldenke, b 1 i 1 g , Casselia Nees & Mart. and Petrea L. Junnel (1934), in one of the most recent studies of the phylogeny of the family, reverted to Schauer's original system and treated Petrea as constituting a separate subtribe in the tribe Verbeneae. ul were E 3 5 et S Ф = [=d Е o $ 5 = Е un additional new рна їп 1938 lished the fi including several new 5р study was ni solely on uar many of the species he described he be variants due to cultivation or environmental factor: Since = Moldenke’ s on monograph, taxonom- ic work on Pet оре stall p local ristic treatments end Bride, 1960; Gibson, 1970; , 1973) and e publications new та siens by Moldenke. scm ais this m wa Xx sola Jax: sen-Jacobs, in the treatment of the Verbenaceae of F Cinta by irem Jacobs (1988). MoRPHOLOGY ies included in Petrea share unique of morphological characteristics of Petrea give below, the degree of variation for each character is presented, as well as an evaluation of the im- t f each с} in the taxonomy of the published d by Linnaeus in ee Genera. Plantarum 19, and Pet lis Specie: f, t niformly in in Verbenaceae, она осе Мит has been deb bated, Schaner (1847) placed ribe, the era Petreae, t. Bentham & Hooker ibe NE a. м diverse genera as Bailloni a & Mart. » Citharexylum L., Dur nia Schreber, and Verbena L. For mea P o) the tribe Petreeae was part of the subfamily Ver- x genus. HABIT ud Petrea species are woody, ranging from li- and shrubs to trees. The habit in Petrea seems to y bó very plastic, and individuals of the same liana, d or tree as in other scandent Verbena 982) found that Ае- giphila i nace Standl. grows as a liana ше fore мі buta asa small br nur tree in the до: species can be a млг лат ibi outing from stumps was con- rmed by field observations. ROOTS of a The root system of P. radiating array of halos hito roots, with an Annals of the Missouri Botanical Garden abundance of a that falls off easily. Ex- cluding tı 9432 a d Hammel 1868) from Panama that report рабай easily; stems that are forced to the ground during t fall + р H + 1 1 сә later become independent individuals. STEMS he stems of Petrea species are mostly slender, with | gr ЖОП, ог ченеп їп n" with con- s lenticels, and may be glabrous or asper- nents. notion, of the stem of ‘Petr ea shows f IOUI OI and this can be used До identify the genus vege- Stems of the tree species tend to be irregular and oval in shape. In lianas the stem is sometimes four- angled and/or furrowed. According to Iquist 1988), Petrea has only a single cambium, situated in the normal position between the xylem and the phloem; rowed stems are because the cambium is equally active аго le perip of xis. In its si cambium irregularly produces axes that are oval in n more extreme forms, the stem be stro О, furrowed, as sometimes occurs in Рен volubilis L. PUBESCENCE Nine mend hair endi tine Table 1) were found in Petrea (1) € duin are june st abundant LEM of e on nthe syrtaces a the leaves. Thess hark d Cha ane and have only bai К in Petrea dd onical- pressions (Fig. p (2) Bulbous s (Moldenke, 1939) are found on upper leaf surfaces and sometimes on bracts TABLE Types of hairs and plant parts and where they occur in Petrea (P) and Xolocotzia (X). Peti- Co- Pedi- Hair type ole Leaf Bract Calyx rolla cel Bulbous PX (PR eth Clavate glandular p P E P P n Clinate Б р В Р Conic PX Falcate X ageniform PX Ribbon PX Subsessile glandular PX Warty P and petioles a Petrea and Xolocotzia. bs hairs are bulbous at the base (Fi rarely eg in older potio, apparently fee they break and fall off with age. (d Clinate hairs (sensu Hewson, 1988) are more n than bulbous on the different structures of Pelton and Xolocotzia. These hairs grow at an angle of 45° with ies AE on which they are oun (4 ) Subsessile glandular hairs (Abu-Asab & Cantino, 1987) are common on lower leaf surfaces and fewer in num on upper su de are sunken. While subsessile glan urface view, a short, usually section (Abu- ubilis, but they were not observed in E T lcate hairs (sensu Payne, 1978) ie e tip йй and are only found on petioles of Xo cotzia (6) Clavate glandular зек (ые & pe! tino, 1987) occur on leave: e more comm olla of several radiall (7) Lageniform fete (sensu pally cells. 1978) are Pubescence in the genera Petrea and is ет АЕ! aspera Tino DT showing conic hairs found on leave: (Мей 7), show PE dy jp und oi bars in A, A.B - 50 и m.) ars in „Р. blanc. n leaves, rng and petioles of Petre ze cteata (Rueda & Ruiz 5 ra hetiana (Martius 1029), D. X. € a and Xolocotzia. (SEM р Annals e Ao Garden с, Д +h f Petrea and Xolocotzia. They are flask-shaped hai $ Ri hban han (sensu уан 1978) аге found of Pet B). They may help retain pollen close to the stigma. LEAVES kne are highly variable in size and texture, hey often feel like sandpaper. This “roughness is due to the presence of special ke, P. campinae Benth., ereas Эб іп Р. rugos unth. All other species the "end ves s RU underneath (Fig. 2D). Le. an texture j is useful as a taxonomic character, although it is in some cases difficult to define except on a comparative basis. Leaves of fea a are uniformly simple and 9p- posite or whor me species are coe cterized by verticils of thee ‹ or ош de eaves. However, paa 10% урый ы. while others show the opposite percentage. Venation poene are рен eful in separating ecies beca аа (Hickey. 1979), with the iG pias ed e secondary veins and the midrib and the anastomosing of the veins near t similar in all species. The P Tere venation і is variable and bed pen: This t na- ceous а, such as Pei ise: Gakara thon. Cornutia, and tomatal patterns bee proven to be useful tax- onomically (Cantine 1990), a f specie: nd three groups o cognized within Petrea. Stomata photograp ps an an S-450 М). Additional imprint 1961), and classi- fication of stomata follows Cantino (1990). Three hasic t f f. RS | ( Pubescence and leaf surface i Jansen Jacobs et al. 341), s showing wa wa ai able 2). The type of stomata is m hein at E. typ NE making Шоп: Thus, the M ae ~~ ЗА) is present in species such as Р. vicalyx and Р. insignis Schauer; о stomata (Fig. ЗВ) occur їп Р. pubes lanchet m haue and P. volubi r, P. maynensis Huber, POLLEN Pollen id am is characte riais of ki genus t has | i P Scanning з n 1 is more or less vil а and ee. The namentation of the of all species is ese or oth without КШ. (Fig. 3D). The pollen р T aue EIU f. that of Petrea о 1 B herical and in having a reticulate in being p exine (Fig. 3E). FRUITS AND SEEDS e fruit of Petrea is drupaceous and enclosed calyx. The accrescent fruiting calyx, which ehe ile fruit, usually has the same ec uU he flowers, b volubilis} The lobes do not change m much in i Héros stiffer and harder with E gea a near right angle with the nies Ne d pi and remnants of the —— on the ia exocarp is leathery or fles y; the endocarp is hard and bony, 2-celled, = forms two stones. The stones are not easy to separate, and each has one seed. However, es, 85 pr sometim in P; Blancheridna, Шоо, is dmm one о found n the fruit lly ached, md lack endosperm. None of these characters are helpful in separating species or groups of species. ANATOMY Anatomical studies were carried out for the dis- tinct в of He pus. dues —— of de stem, w % eth for ашары studies. Three samples ud ent individuals of each species were dehydrated in differ- in the genus Petrea. AZÊ (G ), B. P. т tect i233 ARA D = 500 —C. P. fuos T. ү terior e» the coro! olla. 2 D. P. maynenti (White 860), showing a smooth leaf surface. (SEM pho m.) Volume 81, Number 4 1994 Rueda Systematics and Evolution of Petrea Annals tri Бра Garden a series of increasing concentrations of ethanol, embedded in dicte and secti mic crotome s m. Staining ing aun s (1940) Safranin- Fast n EO apices were sta tained using the zinc chloride tech- nique (Sharman, 1943) to better discern meriste- matically active areas. or taxa available only from herbarium s speci- ia ial with deionized 60°C oven ing a 0.5% solution of The 2b (hansen, 1940). trichomes were ex © and scanning electron microsc opes. Most of the stem кч жеге collected from he кыйры. specimens. ae | | aot thr cc months. T Types of stomata found in extant Petrea and Xolocotzia. Stomatal type Ano- Dialle- Рага- Taxon mocytic locytic суйе Petrea bracteata X P. brevicalyx X P. campinae X P. insignis X P. macrostachya X P. pu ns X P. rugosa X P. sulphurea X Xolocotzia X Petrea blanchetiana M . maynensis X P. volubilis X £ її f, } BAL + dt T o г haracters lor species grouping abl ae Paire be has four кы Р. 5 A p: плута аса, two sing le row. "The de tissue macrostachya, and and aU еы species Вахе only a consisting of = п several layers of bee Bi The е меге frozen. ink a super Eisen Freeze are less dusk ns inch in diameter К. Keating, ers. comm.), and then slides an tangenti 1 o dier. sliding microtome set at 15-25 um. ction stems were then stained by rare the samples with Johansen’s "s safranin for wo hours followed by rinsing in a h 1940) Ё PRIMARY STEM ANATOMY е anatomy of the primary stem was studied n two E of the genu ree) and P. m show ac the same той zones. From ‚ cortical zone, stelar zone, and t ‚ The epidermis is h ypes of and stelar zone do not show any special features. The pith is composed of pa- renchyma cells with b ble special f LEAF ANATOMY In the study of leaf pani three does of each species of Petrea were obtained. The numbers (1972) Scl } fil ent in the mesophyll, and crystals of different sizes are fi e vas E bun- r veins are a T n vascular strand con- ylinder of A sur- rounded by Eagle clos d cylinder: oem аш, fibers e number of petiolar ees pe ong spec ; Thus P nis has n lateral t traces, P. bla nchetiana, P. macr des and P. maynensis have four lateral traces, and the remaining species have only two lateral traces. WOOD ANATOMY are сеа qua: adr ung stems x cept in Petrea the co bir tion "d marginal par dh Ше mbinatio; rg E iH en ions, solitary to oval in dipi жин наде perforation or in clusters of 2-6, 20-40 diameter that varies бош 25 to 250 иш. Volume 81, Number 4 1994 Rueda Systematics and Evolution of Petrea Ficu - Е 3. Stomata апа pollen in the о ну Petreaa nomoc ия stomata. . vie paracytic stoma.— D. 0 species of Petrea. —E. X. um = uniseriate, as observed in P. hurea, or "€ eas (1-12 cells wide), as а іп Р. : ubilis & Hess 41) described the eartw. part с 1 well dilferentiated Кош the yellowish : t olive sapw hey also reported рен iir cipe ү on radial шас: Тһе is mod- stai 4 and heavy, medium textured, with an t to finely undulate grain sg 8 properties and fairly hi em ARTE to i^m & Chalk (1972. 1979), the cork d. and 134), showing Galllocytic stomata. —C. P. volubilis (Gen айп lle и perifolia, with round pollen grains. (SEM photos; bars in А = A. Р. brevicalyx (Ducke 22. tb showing ry 7794), E teristic ,B-E-5 Xolocotzia. 1705); this triangular-shaped m р ad thick d inates superficially and i the outer tangential i The e perieyele contains a боров! ite a and continuous сф 9 scleren- Solitary cape tals in the pericycle are associated with the etimes enclosed within sclerenchyma and are som the stone О. The wood silica in all the elements. The ate in mostly simple in the borders in some specimens of E. ‘ols! ubili Annals of the Missouri Botanical Garden TABLE 3. Журоко features of the leaves of Petrea and Xolocotzia of cell rows Number forming f the palisade Taxon epidermis layers Voucher X. asperifoli 1 1 Neill 5477 (MO) P. blanchetiana 1 1 Rueda 507 (МО) Р. bracteata 1 D Rueda 730 (MO) P. brevicalyx 2 1 Ducke 140 (MO) P. campinae + 2-4 Cordeiro et al. 76 (SP) P. insigni. [ 1 Ducke 688 (MO) кыне" 2 1 Smith 2626 (Е) тен maynensis D 1 Rueda 1129 (МО) P. pubescens 1 1 Rueda 1010 (МО) Р. гивоѕа 1 2 і (COL) P. sulphurea 1 1 Billiet & Jadin 1705 (BR) P. volubilis 1 1 Rueda 431 (МО) CYTOLOGY Number and орношу of dap have cies were studied: Petrea volubilis from es pite at the Missouri Botanical Garden that originally came from Central America and Brazil; and P. mosome nu aoe that Goldblatt diis. Nee. е. Petrea The chro lubilis. nerally ‘varie. The aen ot dus e ranges from 1 to 1.5 um, and the width is less than 1 um (Fig. Pe ECOLOGY HABITAT AND ENDEMISM e e aes g Petrea ae found in the pisc of zm to very specific is E re esulting in narrow endemism. For exa > P. Me and P. inund obvious ecological оеш For nee urs in a wide r. at Rie: range of habit more common within rivers and in open areas SOR е gaps ог оз trails. In general th ecies oft is very patchy even within its habitat, with Е | FIGURE 4. = 10 um.) Petrea volubilis (Rueda & Cuadros 431). sh nus Petrea. (Bat Volume 81, Number 4 1994 Rueda Systematics and Evolution of Petrea clumps of six to ten individuals. Petrea volubilis can also be found on n cliffs. There the rope ste are typically wr) a and e ng 5 m, with the ц он at the end, but the inflorescences are upright. efi POLLINATION BIOLOGY Petrea species seem to have a mass-flowering “big bang” phenolo, gr at the beginning of their flo continue to flower more e flowers at a time for | husk. ce 4000—6000 Fim ig [Пе principal visitors УШ ыга ботев аге PEN cies of Apis, species of Bombus, and Heliconiidae butterflies were als rved. Thrips in pollination becau: r after the pollen is dry. Euglossine bees are the most commonly observed flower visitors an ‚ pre- t amone юп, 1971). Тһе sex ratio of ‘haan bees at Petre is t to the observations of ea Gentry (1976) in th a The Apinae visitors t were species Bombus and the introduced genus Apis. Although a co-evolved one, since Apis only recently reached the Neotropics. SEED SET Fertile fruits appearing swollen or incrassate are пне" {тот infertile ones, which ap- аг Па = n e = B ir { addition to ovule abortion, predation of Tuits init Petrea’s терше крае Little redatio: about a f the pee observed em on ehe т п eaten by Curculionidae larvae DisrRIBUTION extant s species distributed h ern Mexico add through the West Indies and Central Am found throughout th e ge- nus (Fig. 5). Other ER ылы species kde р blanchetiana, found from Panama, Colombia, and Venezuela south to Peru and southeast to Brazil and the Guianas (Fig. 6); ше from Colombia south to ES Brazil, а (Fig. 7); P. pubescens ranges from Sio nda south to Colombia, Ecuador, cat Peru (Fig. 8); P. brac- teata ranges fro Guianas southwest through om the Brazil to Amazonian Peru (Fig. 9 Р, ee is restric azonian Brazil (Fig ee brevicalyx and P: campinae are i to i 11); P. rugosa is endemic to southern Colombia and Шш, Venesuela e. x 1). In addition, paa the former Czechoslovakia (Moldenke, 1938). USES he known uses of Petrea are limited. In flower or fruit the plants are attractive; they were first Lin 1733 when у. Hn MR TN rown as an Qaem ht un brou g Pre Mexico xi 1812). Most of the spe- own 1 ens se focally. med in horti Petrea ipis are sisi propagated by cuttings or layeri Record Hess : 0941) кенеге the wood of ч w ч $ w Lad — Ф Еч 1 and without any y commercial иш. The wood is “very hard to cut when dry and seems to as firewood, edicinal plant in Ecuador, and "e Сали. In the West Indies, Р. i 1 1 Chi lb (1 ) Hitche lA T3 1981). In Еее Р. maynensi s, mixed with three other plants, as a contraceptive (Wilbert & Neill, 1987). In the Gui ianas, the ual pire ыгы the vegetative part о to legs to relieve aches and за (R. DeFilips, pers. comm. The pharmacological. action of Petrea is un- known but Andray et R reported phen lpropanoid glycosid a and in more a The of 1 ا‎ ora the Neotropics. They f 785 Annals of the Missouri Botanical Garden ym oM ERE о Ч L] Ficure 5. Distribution of Petrea volubilis. 40 other genera that are used in folk medicine. These compounds have proved to have good an- tibacterial, antihypertensive, and analgesic prop- erties (Garnier et al., 1989). Finally, Petrea rugosa is used as a melliferous plant for hive honeybees in Medellin, Colombia. Echeverry (1984) al p 1 tł . pubescens and P. rugosa are used as melliferous plants in the department of Tolima. PHYLOGENY More than 1000 species and about 40 genera of Verbenaceae occur in t present on the calyx and by the asperous leaves. Ta thie + + kx 2 1 uc فرت‎ nt, / P 1g with Cas- selia Nees & Martius, Lampaya F. Philippi, Re- h rs originally used and l-seeded pyrenes, characte by Briquet (1895) for tribal delimitation. e Co [ К ai NN METHODS ; ionships of Hypotheses of the phylogenetic relations Pet. аи the oth tae in the Petreeae, as well , a © ; ere as relationships between the species of Petrea, w PSU : alysis generated by a cladistic analysis. For the an jn of the tribal as well as species relationships, ! h; lvzed using distribut f was analyz b parsimony criterion (minimizing omoplasy; са lan, 1984). On the basis of these synapomorpn! | аха were ordered into one or more ae resented by hierarchical branching patterns. ul acter polarity for the characters selected e comparison meth He 1.5; Farris, 1988), applying the implicit pt. for calculating trees. In the an d е E from the best fits to the most EES using rescaled consistency (rc), which is the pr' ا‎ of the character consistency (c) and the Mert in retention index (r). The product is scaled na P the range 0-10. The weighting procedure Volume 81, Number 4 1994 Rueda 621 Systematics and Evolution of Petrea 9 200 400 600 800 1000km ید / 0 100 200 300 400 500 600 miles £: Tirap J FIGURE 6. Distribution of Petrea blanchetiana. © peated on vicodin produced trees until the trees no long ange (Farris, 1989). The pr ET ET am “ 11; (йм ua ia was ИЕ нм Опсе ай Tis ri add оше DE "er nships were Or examinati Production of the [Rod CLADISTIC ANALYSIS OF THE TRIBE PETREEAE т? of the most difficult tasks i in attempting a cladistic analysis ж Ошёгоцрз ( 19 if outgroups are iranê ren the phylo- У hypot thesis can be modified as more data n the relationships of the genera become available. arexylum bois. the tribe Petreeae (whic M Verben eae). Citharexyleae are clearly ted in the subfamily by having fruit composed of two or fo nd 49 ree Cith- кеу nta аге w ГЛ known, making so tribe, where most species et and even entire genera are known only from a few collections the tw 5 synapomorphy that is shared by the two tribes is the spi iform, inde d yoga (Fig. 1C, D) and a genus is defined by its own autapomorphies whereas Petrea is defined by having triangular Annals of the Missouri Botanical Garden 70 T = = = a: eta ч» ee ос х 10 B | : G ° Я : e Ј) дек ы feto, e اک‎ N N rre H N, — oN S | 0 E | Ф." Á e b d Ed e -X| © 2 ti Es 10 1 [ өө 9 (9 E von о 200 400 600 800 1000km Á Соло لد بال‎ о 100 200 300 400 500 600 miles ae ~À 20 FIGURE 7. Distribution of Petrea maynensis. х tained (Fig. 12B). The smooth pollen and a calycinal crest on the calyx. tenti JE poc т g : v P. The use of more than one charac er to infer a resulting tree shows wai Petrea is mon mtr со п ry for this clade does not support and contains the follow g eight » id the possibility that these hme are due to groups: (1) comprises all species iid eun omoplasies rather than to hom which is the only species in the genus witho a traces; (2) P. nsis and P. volubilis v : į ] aborted axillary racemes; (3) Р. pubescen utgro ter three rounds of rsen one fully Bebo tree of consistency index of 0.74, and a re- successive w length 122 26, d the the re aining species except su d d species in this clade with air of brac e Р the pedicel; (5) the remaining species €x ue bracteata, the species in th de with wt one layer of palisade tissue; ( er me P. blanchetiana, the species ы m ве four ms Volume 81, Number 4 Rueda 623 1994 Systematics and Evolution of Petrea 9 200 km е y 100 miles e yi Iri AC E l ON 8 EN T Le FIGURE 8. Distribution of Petrea pubescens. FIGURE 10. ,, Distribution of P. insignis (circle), P. hy and P. sulphurea (square). ee о 200 km برت ا | SS ГА 0 100 miles =" _ ae, TAF С LAL. Ficure 9, Distribution of Petrea bracteata. 624 Annals of th Missouri хын Garden ll. Distribution of Petrea brevicalyx (cir- Le se pue (triangle), and P. rugosa (square). macrostachya, the species in the clade with pitted leaf beneath; (8) a clade formed by P. brevicalyx d ampinae. The cha ccur in northern the main concentration of species in the genus is found. SYSTEMATIC TREATMENT CRITERIA FOLLOWED IN THE SPECIFIC TREATMENTS The delimitation: of taxa was б! оп ошо. 1 the Morphology section. Measurements of the Ке are given for mature leaves ter branches. For dae purposes, only discret characters are included in the tech species. This "e кейе аз much as o TABLE 4. a ale de and character states for analysis of the ps Petre 0. Leaves smooth 0, rough-scabrous 1. . Leaf-scar swollen and c corky 0, neither swollen nor cor . Hairs not bulbous. Calyx lobed 1 _ based 0, hairs bulbous-based 1 "T ааш or truncate 0, calyx ШУ wn e of sepals fused: Aap 0; =0.50, 1. 0, swollen чои Ф , р E л = #2 2 $ costa s ithout ee je ‚диме sagittate 0, not sagittate without basal disk 0, with fa disk 1. o pore X Data matrix for the genera of the tribe The character Mains 13 - - pista ШЫ: a 1= apom ас 709 See Table 4 fo: character states. Taxon O1 2 3 429 6 PT DI Citharexylum/ Duranta 00:0 ОТОО 0 0 DM Casselia Ow 12 vl: 20505020 ES Lampaya OT. 0.0 TOOT Petrea 1:20 01 T7 TES Recordia 0-:0.0:0/0-—0 ОТОО Xolocotzia 171070. Те ee А E as collections represented in a large number of herbaria. However, on the maps provided f for each species өс f the collections exam e been plotted. All the specimens seen are E | in Ap- pendix 1. Petrea L., Sp. Pl., ed. 5 626. 1753. TYPE: P. uita irl: 781.1). Woody у shrubs, or trees. Leaves decus- sate- а: ог анна d often rough- LE 6. ners and character states for analysis of the genus Petr 0. Erect plants 0, climbers ngest peti idate 1. ooth 0, bullate 1, pitted 2. 5. Epidermis with one row of cells 0, with more than one row l. | 6. Mesophyll with опе ya) of palisade tissue 0, wi more than one layer т: 7. Head of subsessile ferma hairs with 74 се <4 cells 1. 8. Clavate бм; hairs absent 0, present 1. 9. in pits 0, in pits 1. 2 10. ytic 0, anomocytic 1, ee Aborte мины dh racemes absent 0, рге: . Inflorescence nal 0, axillary Longest flo: Mg legis oe Pei without a pair of bracteoles 0, d a acteoles сл < o л 2 Ф & Ind o" л B — Noe 1 13 >15 mml. 14. pair of 15. Ca ly x pubescent 0, Morin 1 6. Ого e а l. 17. cas fasion <50% 0, SON. 1 18.-C , united 1. + 19. a corolla tube =15 mm 0, >15 mm 20. Ribbon h lla 0, absent 1. 21. Staminode abton t 0, ne 1; 22. Style pubescent 0, g 23. Stigma subcapitate 0 peu uum Volume 81, Number 4 1994 Rueda 625 Systematics and Evolution of Petrea Ficu - Mos os fi t parsimonious tree for =e of the tribe Petreeae.—B. Most parsimonious ы i a s species of the genus Petrea hs broc nal, race ч sn bracts ten an Owerc: f N beers КЕК Ьу ог тоге аран Inflorescence axillary ог ‚ the rachis with several pairs s in ici sina r portion not sub e brac- r s alternate Mdh Sepals; ку » 3 ары ваи hypocrateriform, usually dee blue or dim than the calyx, some- — white or yellow, occasionally with white or sp wish spot in Г» неш жыдый чаара ма ty роон ке, чач indric infundibular i f lobes van 0 mens 4, didynamous, sometimes hom serted near the middle of corolla-tube, cluded the filaments are wiry or stout nud. орна ог нт dorsifixed near the base, 2-c اا‎ v ning introrse, the connective r dn surpassing the ao 626 Annals of the Missouri Botanical Garden Data matrix for the species of the genus Petrea. The character states 0 = plesiomorphic, 1 & 2 sper See Table 6 for definitions of characters and character states and Table 3 for unabbreviated species pithets Taxa 0:15 227354. .5 "6 7578. 9 10 1E 12.13 14 15.16 17.18 19:20 PI Xolo 0720703 0 —0, "0 0 OO 0. 0 020,-05.0..0 0. 0.0 OO blan Miler 150 0 0 51:0:2 0,055,500 3 p $ 23 8 E brac loD LA TSO WEG A OLD 055040 0.0 4.0 ИТАС brev l1 30.117172 170.407 15 «E 4-40 «0340/70670; 1- 1 25 008 IUE camp 0 5051520 ,32,.1 О T. 1040.0 0 34-1 02 0 insi 1: 130550. 0 0 OO OO 1 204-1 O0 1 0; 0 O 30: ООО macr I 1992. 1 42941. OO 1 lb 005077707 O L| b 030 00 ш ААН mayn 1202501: 0 14.020,20. 0. 2. 141.0 0 LE 0 0 2 1 2 M pube OF 2 E 0. 0'0 O0 0 1 0 0 1 E. 0::0: 0:02:00: ОИ rugo 0:50:12 50. 07 OE 0* 1770 20 0-1 0. 0 0-090 2-0 0D sulp 1 05:15.1. 0 O OSL CO 0. :0.505:0:000H 505205902 15 CDD RE olu Vil РО бы Ө OOOO 2. BETON MED ИЛИ ШОШ staminode sometimes Spore basal "КОО incrassate, its tube slightly accrescent, mostly los- wide; h and изи a m | an y ing its blue color, very oug usually perior, subglobose bl rne on a conspic- ribbed, the lobes accrescent, reticulate-veined, di uous disk, ‚ each cell l-ovulate, compound, — vergent, the c ycinal crest callose, curving rds but 1-carpellate through the abortion of one carpel; and conv: ging, thus completely closing the mouth l l, ascending, hemianat opous, or or- of the calyx-tube; it drupaceous, enc y thotropous; style included in iin r ,usu- the mature calyx, the exocarp leathery or fleshy, ally shorter than the stam e, terminal, the endocarp hard, 2-celled, forming 2 stones, each ay fo sae нып capitate, moa б рор stone 1 -ѕеедей, or by abortion fruit 1-stoned th t 1-seeded; seeds ж ог арїсайу attached. KEY TO THE LIVING SPECIES OF PETREA la. Inflorescence mainly axillary; leaf apex ку m und. » 2a. Pedicels with a pair ss leaflike brac x lobe si ——— 5 P КШ 2b. Pedicels witout a pair of leaflike races calyx sinuses closed. 3a. Climbing plants; зап glandular hairs pre: 4а. Leaves ternate; the longest petiole =10 2 mm E cue CP PU '~ 4b. Leaves opposite; the longest petiole >10 mm .. — M. EN m 3b. Erect trans clavate glandular hairs absent. t H ate; with aborted racemes; staminode present = 8. P. pubescens 5b. Shrub with 1 te; without aborted racemes; staminode ‚ absent ЕЕ SED 9g. P. rugosa 1b. Inflorescence mainly terminal; leaf apex pe Id "E y cuspidate. 6a. Ca uc lobes of 2 sizes of lobes; corolla yellow W; pida winged 10. P. sulphurea 6b. Calyx with equal-si sized lobes; dish blue ; stem not winged. ais = м i "nn stly wit at „the tips of peg qure 2. P. bract 7b. Man leaves mi ог pite hu not nh t the tips of branches. Matur >15 » long — Ж blanchetiana ited: the stomata in pits; the longest corolla tube = озок н 710 mm; calyx not corrugate; calyx fusion 250% aby "obe sin h Fond ed Ê et: ув 9b. on 1 50%; | ses с. 10а. Puig the lon est flo | di l >15 1 è 3. 7 brevicaly* 105. н dhe’ beet Baral аы: Гут a pb 4. P. сатріпаё l. Petrea pamper gouge — in ни Prod (1993), BR; a BM, G(2), 11: 617 1847. Bahia: LE, M(2), MO, NY, S; photo of an Bra: Martius nti pene die » Rueda NY). Figure 627 Systematics and Evolution of Petrea Rueda 5.0. Gm P Bicko Volume 81, Number 4 1994 FIGURE 13. Habit of Petrea blanchetiana Schauer (Nevers 5812). Annals of the Missouri Botanical Garden Petrea peruviana cueva jene kar sid ка бн 1938. TYPE: Реги. Lor 100 m elev., чад 1929. (8, pu Reet pe бе эше NY; isoty ypes HSE uminata Feddes Repert. 43: ко © e ЙИ. 8 "S @ а Е - artí roa 1925, ‘Mor і & Kallunki 5521 hae Usi E type, MO; fruits of isotype and photo of holotype, EX). Petrea ш Moldenke, Phytologia 54: 67. 1983. е . Choco: Río Tagachi, 12 km W of o Atri 19 Jun 1982 Рл try et al. 37075 es m ted b Rueda (1993), MO; isolec- e, MO). Liana; the iei ipe gie ess puberulent. -15 mm long, p E us, smooth to the touch or “ү y slighty roughened, sometimes slightly ‚ sometimes impressed- -punc- tate on both surfaces. Inflorescences termin rminal, ra- cemiform, 8-53 cm lo: ong, 3-9 cm wide s 10—31 mm long, often bearing a bracteole. Flowers with the calyx-tu| = t ase mm wide, apex 4-7 de, corrugated, slightly pubescent, 5-lobed, the lobes membra: “ mm lon - wide at base, 5-9 mm wid int, the 5 асше, glabrous; calycinal crest with de lobes ca. 1.5 long, 2 mm wide at ba corolla fragrant, 13-18 2.5 £g © @ Га у long and 7 "P mm Sie. ee a style 2. 5s mm long, glabrous; stigma subc capita Neblina, Camp on Río Mawarinuma, 140 0 Feb, 1984, 5 esner & Funk 15826 (МО). FRENCH сш Cayen around the runway, 1 1983, E ys Goed 5400 (BR, CAYQ), U); Creek Mulet Mort, S of Saul, 10 Feb. 1966, Oldeman 2012 . PERU. Am - 97 ; Maynas, Tam: al. 28 94 (F, МО NY; Alpa hs: 4509 (MO); V 28 pen 979, Díaz & Jaramillo 1243 ween x Nanay and Rio Itaya, 130 x 29 Nov. 1977, Ca try et al. 20982 (MO); Ris чч un tourist camp, vicinity of Iquitos, 12 2 May 8, Gentry et al. 21 с m E A TE PN Mug 100 , Oct g 170 (F, NY, US); Zungaro Cocha го: ч Quebrada А 160 m, 4 Fel RAZIL. Amazo peque, MENS Nati 5 Feb. 1950, "in "25807 N NY, Sy; São of Cre Oct.-11 ap 1857, Blanchet s.n. (BR(2)). ФА п. (Sy; 1842, Dupre s.n. Р) 28, Sam; aio 5092 (NY a ДЕ? 1 Jan. Dom Silva & pica: 60806 AY). Imt o. cut Im ч ocal names and uses. In Panama, "kwagi in the pas species the bub est pe than 15 mm and in P. bla ncheriana they are much longer dr 15 mm s has the largest flowers kno 5 cm across when fr E . This sp inda genus, up to 2. ee cem Steudel, Sy 6: 7 E: Suriname. Apr. 1842, Hostmann =ч Ean 39 (lectotype, ЖЫ zl Rueda B E (Fig. 6). This is species ranges from (1993), W; isolectotypes, B, BM F, G2) Panama to Ve en French Guiana, Brazil, and K, LE, MO, S, U, W(2) regimen solis Peru. ln Peru it is mon on white sand totype, MO, h of isolectotype: p M growing u m in chamizal forest or Y, TEX; photos of two isolectotypes overl m on more ue soil in varillal forest. It is in the same picture, ). Fi op fone’ in ipe atches; for exa шр Nevers du 2 in DC., Prod. 11: 620. эу тп Par n Antoni e CUM growing within 150 m of each other in Panama. ар s.n. (lectotype, е "elected by x 147 da (1993), М; o Mns 1 ` Petrea schomb iana Schauer i tí road, 24. 5-: 5 is from Panamerican 347 : ыгы 184 0- 1844, enl v inl А Apr. 1975, Mori & ран огуре, (1993,1 Kallunki 5521 (MO, n s COLOMBIA. upes: plicate photos, TEX, US). ologia lto \ upes, near Miraflores, 300 m, 2, 4, 5 Feb. 1944, Petrea iri же na var. glabrescens ee ae cda & Schultes 760 (COL, GH, sed "e VENE- 32: . 1975. TYPE: Brazil. P . Amazonas: Río Ne egro, 1-4 k E of Cerro y. эме зс 1926, gres 14291 (holotype, N Volume 81, Number 4 Rueda 1994 Systematics and Evolution of Petrea NA [LET = Met Ai er. =N ЖОШ SN "ы... 1.5 ст P BICO FIGURE 14, Petrea bracteata Steudel (Feuillet 739).—A. Habit.—B. Flower. Liana; the branchlets tetragonal, short-pubes- short-acuminate, the base acute to broadly round- Cent; o i ften with one or two leaflike bracts below ed, membranous or subcoriaceous, very densely the Petioles or on the outer side of the branchlets. | asperous on both surfaces, bullate beneath when es with petioles 5-20 mm long, short-pubes- mature. Inflorescences mainly terminal, racemi- Cent; blades ovate-elliptic, 3.5-30 cm long, 1.8- form, 19-50 cm long, 4-9 cm wide; ресе. № Th wide, the apex obtusely acute or very 1 ст long; р berulent. Fl = < 630 Annals of th Missouri Bii Garden 4—5 mm long, the base ca. 3 mm wide, 4—5.5 wide at apex, corrugated, short- pubescent 5- lobed g or elliptic, 11.5— the lobes membranous, obl T7 mm long, the base 2-3 wide, the dist po -10 wide, the apex rounded or acute, gla Б 2 [7] = ong, , the apex blunt, glabrous; corolla with throat uet and hairy, 8.5- 20 mm long, the base 2-3 mm wide, ampliate to 5-0 th 6.5 t pubescent, the limb rotate, 5-lobed, the anterior Tobe larger, M. remainin, lobes similar; stamens 4; filaments с mm long, sparsely pilose; anthers ca. 2 mm i iba ob- ovate, ca. 2 mm long and wide, glab 2.5 mm long, glabrous; stigma subcapitate.- Distribution (Fig. 9). This species is found from Venezuela and the Guianas to Brazil and Peru , Selected specimens examined. "VENEZUELA. ‚ between Tama-Tama and Cario is sees ar 19 Apr. 1954, Level 5 (NY); че Cunucunuma, 30 КП from Playa Alta, 200 m, 28 D 9 azaroni 963, Boer 642 (BR, U); оронд ш гезегуе, б ; i ; Sinnama 1988, Billiet & Jadin 4481 (BR, САҮ); 2 ‘gee of Petit Saut road, 40 m, 26 Feb. 1988, Billiet & Jadin 4510 (BR, CAY); S of St. Jean du Mar roni, 19 Feb. 1956, Cremers 7673 (CAY); region of Paul Isnard, Mont du Decou Decou, 400 m, 8 Sep. 1983 3, Cremers 8175 (CAY, . 1984 (MO); mp trail from village to Campamento m, 22 July 0, Gentry et p 28928 (TE ХУ Ro Santa vuya Lores, Tamshiya em Const пзе ia, 50 m, 9 July 1992, Rueda & Ruiz 720 (МО); В ve 4 km NE of Ar- boretum Jenaro Herrera along Rio Ucayalî, Quebrada de Colima, 180 m, 13 Nov ‚1081, Spichiger & Encar- nacion 1039 Cid), MO, NY); Bora native communi о: 0 Cachoeira Re epublica, Rio Curuquete, 25 d Prance ‘ et € 14620 (F, NY, S, TEX); Rio News ў т Сат УЗ CAMBR aracaraí ie km 45, pis ae (BM С, CH, S, U E region, Dec: 19 left margin of fü Mapuera, 3 Aug. 1 985, Almeida 213 (МС); left margin of serm Vira- Mondo, ni ds n МС); ~ d from Cachoeira iaci to Perimetral do Nose 24 | May 1974, Coane et wA piena СЕ = MO, ae Oriximina, Ri tas, left margin across Mineracao Santa E June 1980, Cid a E мв NPA) Rio па bi го, A dud eh Me here pared near Rb umina eh N 0 1977, Prance et al. P2551 19 (F, Е, CH, “HBG, MG, d à Rondónia: basin of Rio Ma dera 8 km Ade NY, Velho, 7 Nov. 1968, саре et al. I On 1977, S, TEX); Cui Santarem, km 118 a NE ien Silva et al. 240 (MG, NY. Uy T Natio: 3939 а IM 25 Nov. 1978, Silva & Rosario G, NY). EE iuvinha": In Brazil ` hes “hierba es and uses u are used to iii out god also soaked in wa k and then Volume 81, Number 4 1994 Rueda Systematics and Evolution of Petrea The p bullat ture leaves (see a 14B) in P. bracteata makes this edd easy t oe ош = en —€— species P. blan r the latter CH 3. Petrea cti мазан Bull. Mus. Hist. Nat. йй ser. 2, 4: 748—749. 1932. TYPE: Brazil. Amazonas: s Mannie, Igarspg da Cache eira Grand shores, Ducke s.n. (lec кү ЕЙ selected М Ў (1993), NY; т A A K, RB, S (2) mS NY, sb qna 15: Liana; the branchlets tetragonal or subterete, with minutely g 5 ndular-based hairs. Leaves with peti m long, short-pubescent; blades elliptic or озы: oblong, 5—16 cm long, 3-9 c wide, the apex obtuse, the base acute or N coriaceous, rough to the touch, minutely pubescent нам оччу pitted beneath with abundant short the pits Euer am ped ter- -59 cm 2-3.5 cm ue PME ü- mm long, АН” cpi anthers oblong; o ary oblong; style ca. long, glabrous; IS capitate, bilobed z Distribution AE This species is only ul from Вга Ap s mazonas), growing in igapó Ck-water E din forest) on sandy river banks. Specimens examined. BRAZIL. — "aus, Estrado do Francés Novo, 31 Jan a 1967, Prance et al. 3 NR. NY, 5); Rio тм Dec. 1901, {Ле 5982 (С, al names and uses. In Brazil * td и ' No use is ka nown. It is P 4. hs. campinae Rueda, Novon 4: 992. TYPE: Brazil. Roraima: along the а fro 1, km ‚ left side, 00*06'N, 60°40'W, 12 Feb oan ), Ro- drigues et al. 10100 б isotypes, , INPA; drawing of holotype, MO). Figure lor 5 m tall; the branchlets tetragonal or or subtree m well wide, the apiculate or truncate, the base obtuse, foriaceous, asperous glab rous abov: l hairs in tl с дорд з; terminal, racemiform, 10-22 cm long 5 cm А bracts setacenns] to Бинго; pedicel 6-10 mm long, pubescent. rugate, dentia 5-lobed, ipu sounded, 4- 8 mm long; calycinal crest with the . 1.5 mm long; corolla 1.2-1.5 cm long, "ie limb cust 5- 8 mm pns the lobes 5, ovate, 3-6 mm long, unequal; straight or curved, ics раа. -ри- scent; stamens 4, the filam ca. 1.5 mm long, the anthers oblong, staminod esent; ovary glo iae the _ 2-3 mm а denk the stigma ca Distribution (Fig. 11). This species is only f Ps teh f h x st Ж ¥ n white sand, near km 350 on the Мапаиз- Caracarai road, in Roraima, Brazil. Speci ined. BRAZIL. prises bees BR 174, between Manaus and Caracarai, 6 km S of equator, in a campina, 00°04'5 60°40'W, 17 June 1985. Cordeiro t al. 76 (INPA, SP). None reported Local names and use. P . bre intricately pitted neath ‘and a cal ays with short rounded dos s It twice as long as those of the lianaceous Р. brevi- calyx 5. Petrea insignis Schauer, in DC., Prod. 11: 620. 1847. TYPE: Brazil. Para: exact locality unknown, 1—21 Aug. 1819, Martius s.n. (ho- lotype, M). Figure 17. ues асе чаве died vee 1: 469-470. 1940. E owe t Rio E Paraná de an Ducke s.n. чыте т isotype, К Liana; puberulent-asperulous. i ud branchlets Leaves with petiole 3.5-12 mm long, with minute dark m irs blades elliptic, 3-18.5 cm m 1.5-8 obtuse, the b 632 Annals of the Missouri Botanical Garden 3.0 cm FIGURE 15. acute or rounded, chartaceous; memb nous, киче ш! asperous, glossy and gland ular ‘above, veins. Inflorescences axillar ry, facenti. 14-37 m long; bracts elliptic, apiculate, 6-10 mm lon ng, мурй leaflike, in pairs on the pedicels; pedicels Р ВІСК? Habit of Petrea brevicalyx Ducke (Frées 20501 ). with 3-15 mm "E е -puberulent. Flowers t- calyx-tube 5 g, 5-lobed, the ше чоч, sinuses bonia a peia opened, obova ай long, 1 uberulent; € 12 6 crest with the folios ts mm long; corolla mm long, the limb 5- xk one of the lobes Systematics and Evolution of Petrea Rueda Усора zx A $5 С) У / [2 od 2 dE AGE i y чату 2 Sy. Gs P. m MC уа «с ` ус US I SOLE a) N — A. Habit.— B. Close up of abaxial leaf surface, D. Flower, cut longitudinally and laid open. inae Rueda (Rodrigues et al. 10100). р Volume 81, Number 4 1994 Ficu sho: . RE 16. Petrea cam wing pits. —C. Flower. Annals of the Missouri Botanical Garden Р BICKS FIGURE 17. larger than the others, short-pubescent; stamens 4, filaments ca. 2 mm long, anthers oblong; ovary oblong, the style ca. 3 mm long, glabrous; stigma subcapitate. Distribution (Fig. 10). This species is endemi to Brazil (Amazonas), growing on banks, along black-water-flooded riv. Selected specimens examined. BRAZI nas: Manaus, Е : thin ESSENT GU S gro Ww & Sano white sand river upstream from Manaus, 13 Aug. 1987, Tsugaru ae B-1001 (NY). Habit of Petrea insignis Schauer (Anderson 229). n. (TEX); Barcelos, gir Ri a Cordeiro 307 (SP); Rio Taruma, 14 Aug. 25053 (NY(2); Igarapé Ponta Negra. Fróes 29568 (IAN); Rio Negro opposite km upstream, 8 Apr. А ТЕХ U). 21 June 1882, Schwacke 3638 (G); er side and small islets of Rio Ne; 5. In Brazil “йог de ж ZIL. Amazo- Local d rded. rancés Novo, 12 Mar. 1956, Chagas Miguel," **viuvinha." No uses have been reco Volume 81, Number 4 1994 Rueda 635 Systematics and Evolution of Petrea This s E is dins distinct and is the sister group to the of t enu: the cladistic analysis thus appearing to dea n Нер ever, cies, such as bractlets on the e as in їр inuses opened as їп Р. g ppor y the cladistic analysis. Perhap this greater E tion of P. volubilis is a PE of increased dispersability of the lighter calyx. 6. Petrea congu ue Bentham, Ann. Nat Hist., ser. 1, 2: . 1839. ТҮРЕ: Guyana Along the brook eed ak, 1835-1839, Schomburgk 158 (lectotype, selected by Rue- da (1993), a dst ў BR FG, vo > КО), EG gment of бише ig NY; ypes, F, MO, TEX). Figure 18. ; the branchlets obtusely tetragonal, mi- г : ovate- "elliptic, 5-21 cm long, 2.5- the apex the Base rounded or acute, coriaceous, as sperous glabrous a ve, intricately pitted beneath wth abundant short hairs in Б с with strongly de- veinlets delimiting the pits. Inflorescences rminal, а many-flow: s 2-8 mm 70. cm p brac ca. 1 mm long, the apex obtuse E rounded, glabrous, corolla blue а violet- e. 13 mm long, the lobes 5-12 Pr than the Ei 4, the apex е оа "AE cent; stamens 4; i filaments ca. 2 mm 1 T. the anthers ca. 1 mm ong; ovary oblong, 1 ong, glabrous; style ca. 2.5 mm long, yT pitate. E Stigma ca Dario disi 10). Petrea eap pde I5 endem d Suri- his climbing up ад of Di Ото; іп сго 800 ^ Plotropis and More stel at tirk up to Selected | specimens. examined. odisse ned runi river, orest Department of Guyana csi d». Potro s paruni region, approximately 1-3 km toi, 650 m, 13 Mar. 1989, Hahn a Уз a e Kanuku mountains along creek, 300 m, 21 Feb. 1985, Jansen-Jacobs et al. 341 (B, CAY, MO, WIS); upper i Sep. 1986, azilian hir 3-5 k ward river valley, Fel b. qM y^ 261 (U); elmina mountains, creek valley at the S base of Juliana Top, 5 Aug. 1963, i hultz 10330 0, US, WIS). FRENCH GUIANA. Cayenne, Е road in he mT near Neyrat, 4 Feb. 1 7, нет 4302 (САҮ п Kourou road near Montsinery road, 18 Арг. no pier 201 (CAY). Local names and uses. vine" in Guyana. No s kno This species is dod. хен to d brevicalyx and P. campinae, which also have pitted leaves. However ah ma Aoc daten is the ey E is called *sandpaper тее half of the total yan and the oe lobe sinuses This species has the longest rachis in the 70 cm, presumably the source 4: 602. 190 TEX). Figure 19 Petrea эмин Moldenke, Feddes Repert. 43: 197. 1938. TYPE: without кон, us te, or иду (ho- lotype, P а Р; fragments of holot F, NY; photos of holotype, NY, TEX). Liana; the branchlets € ай often striate- oe short-pubescent. Ге whor 3s or 4s, ra rled in or 4s, rarely some decus e; petioles -11 mm long, UMP rulent; E E or ovate, E cm wide, the e apex acute or orescenc the racemes 6- 5 cm ns ca. ide at apex, glabrous he lobes ita была je us 11- 15 : mm ee ong, Annals of the 636 Missouri Botanical Garden Volume 81, Number 4 1994 Rueda Systematics and Evolution of Petrea ji | | Ficure 19, Habit of Petrea maynensis Huber (Ferreyra 8100). ~ Ficung 18. Habit of Petrea macrostachya Benth. (Schulz 10330). 638 Annals of the Missouri Botanical Garden —4 mm wide, the apex acute, glabrous; calycinal crest with the lobes ca. 2.5 mm long, and ca. 1 mm wide 8, short-pubescent; style ca. mm long, short-pubescent; stigma subcapitate, lobed. e hE: (Fig. 7). This species occurs from Colombia through or al Peru, and livia ле i nr banks, and стол] hills in tropical rine he plain forest T It has orted 100 t n flood bu altitudes fr Fn 750 m, pure пег еа huge emergent trees of Bertholletia duelo Humboldt & Bonpland. Selected specimens examined. COLOMBI Macarena, TO, Barclay 7146 (MO). Vaupés: near Miraflores, 300 8, 4 Feb. 1914, Gutiérrez & Schultes 783 (COL, бн) Б VM ADOR. Morona-Santiago: Gua id 27 Sep.- t beno verd vini diee il behind the Cuyabeno Seite ч - agp Paz y Mito, si jig Kona PERU. Am s: Rio Cenepa, Hua of Quebrada Chigki Shinuk, 200 m, n Pes m guo MO, TEX); Río Santiago, 180 m, 14 Nov Hua yacu tween Constancia and Serafin, 50 July 1992, Rueda & Ruiz 808 (MO). Madre de Dios Tambopata, Río Tambopata reserve, at mouth of Río D'Orbingy, 250 m, 2 Mar. 1981, Gent y & Young 3187 (MO, TEX); je gru Park, Coc hü uplands, 400 m, 11 Sep. 1986, Nunez 6083 (MO). San Es Tocache No m ia Nue vo airport, 400 82 (F, G, NY, US). BRAZIL. am from Cruziero do Sul, 150 m, 23 Аз 1986, roe 62570 (MO); 12 km "еш Rio Branco, on Rio ied Porto Velho road, bs o 1980, Lowrie et al. 291 (GH, MG, MO, NY). A onas: Sao Paulo de Olivenga, Rio Solimoes, 2 Fe bl 35657 (G, NY, P); ne S ndiatuba, from d to 10 km upstream, 26 Feb. kb no ri et С aes E A US). Mato Grosso ana, 5 Sep. 1 ide dere 3292 "em 200 qe e of Cuiaba, 200 m, ‚ Maguire et al. 56867 (F, NY, TEX pec conta фу» 1 п. 1928, Duck 22543 (NY. Rio de Janeiro: Sao Paulo de Olivença, Rio es 2 Feb. 1937, Ducke s.n. (U). Rondónia: Guajara- Ма, Rio Pacaas Novos, 15 Aug. 1976, Caval- ae 3297 (MG); between Porto Ve Cuiaba vi- cinity of Jarü, 16 Aug. 1968, Forero & Wrigley 7106 (F, INPA, M, MG, L, U). BOLIVIA. Beni: Vaca Diez, разве, 28 к parr Ge pet 22541 oh junc- n of Rio e de Dies, be 188 uec md 25 zd А EI А iri, July- е! xm Bang 14 d pe rm Марі, RM Buchien 1 967 eee Ne Aa White 860 (NY). Pando: Madre о Madre gh Dios, 19 ks y air from m, 18 Aug. 1985, N rr (F, MO, of the Western y 1992, Guaytu forest, 27 Sep. 1917, Stein- bach 347 (G, NY(2)); Río Surutic, 400 m, 2 Oct. 1924, Steinbach 6544 (BM, F, GH). Local names and use In Bolivia, natives чесе pur dias bg flowers of -— cies avenues. а is called ‘‘viuvinha” n Brazil an e calabaza" in Spanish; there th and atlas шы ш with Usnea and T (Wilbert & Neill, 1987). It is called pose raceme: id nensis is ded oce by its v Me ves m volubilis ppo e. In addition, f P. maynensis do not reach more фай 10 тт аѕ іп Р. volubilis 8. Petrea pubescens Turcz., Bull. Soc. Imp Nat. Mose. 36: 211-212. us TYPE: Ven- ezuela. Mérida: near nsr |, 600 m elev., Nov. 1846, ie Schlim a (lectotype, Г аў Ruck (1993), B isolectot types, BR, G(2), Р, U, W; ph uo mise i к Mo, AY PRO NN Petrea A ie Bol. Cient. Tecn. Mus. Ven . TYPE: ween: Рог a near pried on 1924, Peraza 11532 Volume 81, Number 4 1994 Rueda Systematics and Evolution of Petrea ÈS fi) ©) AS N E SS AEN y А W ; d ۱ = Q NV s с RÀ d N Z AN ^ NN i 4 SS \ WO ын СЕД i ИА 4.0 cm | / б \ / BOs el PBICK? \\ a $ Ficure 20. Habit of Petrea pubescens Turcz. (Neill 10014). 640 Annals of the Missouri Botanical Garden ), VEN; isolectotypes, ү: US: photos of a eye, NY, TEX; photos of iso letotypes | эй: pese an a rei M type of ды Г Repert. 43: 188. 1938. n. 1877, André 4707 (holotype, K; y fragment of ауе NY; photos of holotype, ны pubescens var. klugii sup tas e Feddes Repert. 43: TYPE: Peru. San Martín: Río Hu- May 1935, Klug po s, BM, F, GH, MO, S 1938. A fe elev. е, т, (holotype, NY; isoty, US). ша erpita: Moldenke, Е eddes Repert. 43: 177. E: Colombia. dinamarca: near Bogotá, 5, Purdie s.n. (holotype, GH; isotype, ind i holotype, NY; photos of holotype, NY, TEX). EPI: Petrea pie res var. адашуу реч жуы. Mir 18: 72. 1969. TYPE: Venezuela. Táchira: N of San jn. MCA. on hwy: from La Fria to San Cristóbal, from a cultivated phat, 27 he 967, Bunting 2369 (holotype, TEX; isotyp: tu: up to 20 m n eue m usa т or grayish, obtusely tetragonal, densely iu pu- bescent. Leaves decussate-opposite or whorled in 3s or 4s; pee 1-13 mm long, short-pubescen with ellowish hairs like the cen ы трн К «ине ог ар "ешр b p 18. oe 1 wid t its axils; pedicels 1-10 mm асан mn | Beare мее to 12 mm long. whit е or blue, 3-5 mm e, the apex 3.5-5.5 mm ы, 2-3 mm wile at base densel t branous, obovate or elliptic, 11-18 mm 5 mm a “oe widest point, acute or blunt at apex, with the lobes ca. Е 5 mm long and wide, the apex ‚Жеш acute, translucent pubescent; corolla-t 6-7 mm long, the base 1. rs 3 mm wid to 4-5.5 mm at apex, pubescent, the limb es with — dots, rotate, 5-lobed, the an- maining lobes globose mm wide, s ines т siyle с са. 18-25 mm long, pubescent; stigma subcapitat Distribution e 8). from northern America south to northern This species is found th rt Bolivia, and ise tho Brazil. This species has been collected at 2200 m elevation, representing the d altitude for any recorded collection of the Selected specimens examined. | COLOMBIA. Antio- quia: Medellin, 22 Apr. 1927, dies Ve aa US). Со : Rio Quili а , between San San Pedro, 1150 m, 5 Oct 4, Fernandez EE (COE 23 Aug. 1853, [^et 572 (GH). Cundinamarca: " sagasugá, Boquerón de Fusa, 400 m, 20 Aug. 1 Ló, ópez- -Palacios & Idr obo ed Soe BA firn $ ot and Melga: г El Paso, E Cuarrecasas бг и Ae , Sch Sm 1521 (BR, BM, F, G( Jan. 1937, García 5206 (C the mw of the rivers Sansa a "Сие ејаг, m , 22 Feb. 1956, robo E 1 (COL. dns wm Santander: Pa amplona, 1 3, Linden TIO aii CO» GH, W(2)). e. И: mg ria, i . 1930, Klug 1894 (BM, F, GH(2), МО). Буы ег: sailed Cucuta and Pamplona, along Rio Pamplonita, n La ir "m у Tue Сай ano Caribe, са. 19 ai m SW of Elorza, 90 m Mar. iih Davidse & ¢ González 6214 (МО, 5Р, , Ar nr , 6 June R ert. as: Pedraza, Feb. 1953, Aris- te, aui еде 2 A тл 1959, Arise 3864 (NY) 850 m, 4 June 1957, Bernardi 6350 (G(2)). "n Cedeño, near Cai o Villaca, road Cui Y o El Burr pe aes 1984 4, Stergios et al. 8570 res Ак» усе 2288 (ВМ оп road to Villanueva, 1 5 Ма d 5 et al. 1864; 1865 окт) М 4 km NE of yb ong P. (MO road San Fran m A 1722 (BR, U). Portuguesa Feb. 1, Aymar rd ie et PORT); Ospinos, 10 oa 5 June 1984, Aymard M 1510) Río, 1967, Bunting : 2369 ( ТЕХ), “Trujillo: Carache del R betwee: ñas and Las Р: 1 Dec. 20 & dece 10417 (MO). N pe Orellana, 20 Coca, on Via Auca, toward Río Tiputini, 270 m, Volume 81, Number 4 1994 Rueda 641 Systematics and Evolution of Petrea 1991, Neill Rojas 10014 (MO); 15 km S of Coca, 1992, Rusa et al. 2010 (МО). 7а- Chine A pe: Zam slope of the en nuco estal Dani 270 m o Iberia, Río Tahuamanu, 330 m, 4 Sep. 2164 (MO, US). San Marti а Жай Chazuta, Huallaga, 260 m, May 1935, Klug 4155 (F, GH, MO, NY, S, UC). BRAZIL. e: Rio Acre, Aug 1, Ule 9722 ce NYO), US) p. Pando: 54 km SW of Cobija, 250 m, 24 July 1988, РЕ ШОП et al. 35 LPB). Local names and use es. ltisn e orgia and “chaparro” in Colombia and “‘penitente”’ г Venezuela. It is widely cultivated as an aE in Venezuela. is = 18 үш 2 K rugosa, a the E m, as уа to Р. pubescens, a tree у . pubes cussate as rugos leaves in the latter us are bullate as opposed to smooth in D . pubescens. Power ers with ы те ix x es are nanan Or eis 9. Petrea rugosa 1818 Kunth, Nov. Gen. Sp., 2: 282 уре; F Р; E otos of f lectotype TEX, US; F, MO, NY(2), TEX). Figure 21, m e gg es Benth., PL i 1846. M Fig КА (19 93),1 К; isolectotypes, ВМ, С, e t of isolectotype, NY; photos of lectotype, Petrea rugosa var. casta Moldenke, Reena pie: ыч 48. 1938. түрЕ: Рапата. Сап д plant, 2 Dec. 1936, rer ane s ye ы speciall both Medals pubescent e y . Inflorescences eath, mature leaves bullate axilla b d at apex, ам -pubescent, 5-lobed, оте, 10-13 mm long, ca. 3 mm wide at base, 5-7 mm wide е, i mm at apex, the anterior кран largest, body ae ca. 6 mm long, mm wide, the lobe: i ES ng, ca. 0.5 mm wide; .2 mm long and w ide, Ee н . 3 mm long, m stigma subcapita Distribution (Fig. 1 Colombia T Venu, growing in mountain sa- anna with: shrubby growth, or on open rocky UR at altitudes of up t От. spa d specimens gamin . Antio- San Jerónimo, 900 m, July 1968, Barkley 38C494 (ey EX) p ad is ^d T 1979, Be ntería et al. 1630 (COL). Ca rada, 250 m 8 Dec 1936, Бу ght 2103 с р ME US(2 ); Vi EE Gian. ЖЕ ы т, 6442 (Cor, I US). 5, López- Palacios & robo 3831 (COL, Nr LX. еа пеаг ipe ге turn W and pr PPS for : km on гова z A 1400 m, 26 May 2, Barclay et a sagasugá, Feb. yer Lindg a (P). еа raditas Tira 21-26 Feb (UC); Cur n- Zuloaga endl, siege t i 94 (COL). Mag n 8645 E е - МЗ, 7, Triana s.n. (US). М mountain savanna with shrubby рг rowth in pea m, am 1974 ut et al. 5599 (COL, MO, ТЕКТ. үзү 9 рес. US); 400 m, 1851- 1857 M 948 the Magdelena rivi Le igo с phis Valle de си Duque-Jaramillo 4340-A vs rni 2 May 1 90 Sig (US) Valparaiso: Murillo, 1000 m, 2 Mar h e and La Zona Militar, 1650 m, 12 Aug. 1985, Bond 5080 (MO). Distrito Federal: Caracas, Humboldt & Bon pland s.n. (P(2)). Petrea rugosa is Local games ud uses. kno **azulino " "azulito," “cha- arrito,” “chaparrillo,” ies оноро blanco," “ma- moncillo,” **pavita," **pluma del г uma de reina," **plumilla," and “sombre: rito" in Colom- bia, where it d as rnamen It is closely related to P. pubes see dis- white calyces, are known and in culti- vation. 642 Annals of the Missouri Botanical Garden ЎЗ Nr БЫ FIGURE 21. Petrea rugosa Kunth (Garcia 6472).—A. Habit.—B. Close-up of abaxially bullate leaf surface. 10. Petrea sulphurea M. J. Jansen-Jacobs, Flo- primary vein, minutely pubescent and glandular di Guianas 4: 60-63. 1988. TYPE: French dotted beneath. Inflorescences terminal, raceme Guiana. Piste de St. Laurent à Paul Isnard, 9-16 cm long; bracts narrowly ovate, 5-10 mm 15 Nov. 1982, Granville 5341 (holotype, long; bractlets one pair on the pedicels, setaceous, CAY; isotypes, BR, P, U). Figure 22. 1-2 mm long; pedicels 1-3 mm long. Flowers vith { the calyx-tube 6-8 mm long, the lobes rounded or Liana; the branchlets tetragonal, winged, often apiculate, unequal, 2 lobes narr owly elliptic-oblong: twisted, pubescent, some of the hairs glandular. ca. 20 mm, ca. 6 mm wide, acute-obtuse, 3 lobes Leaves with petioles 5-11 mm 1 үл. ue d. , di wide, 5 cm | , 48 iptic, y < 2: md ong, 2.5-7 cm wide, the apex acute — acute-acuminat s and apiculate, the base acute or broadly acute, cinal crest 5-cleft, the lobes ca. 2 coriaceous, asperous labrm h А ong, the lobes 8 pt on the scent; corolla yellow, 2-3 cm long, Volume 81, Number 4 1994 Rueda Systematics and Evolution of Petrea 2, E— EATUR. = э; Sar oS a ВЗА Sen TRIN A Lo VEO кос SN EN es MEM P Bick RT. FIGURE 22. Habit of Petrea sulphurea M. J. J "mu glabrous; ovary oblong, ca. 1 mm long, E» 7—20 mm long, glabrous, stigma e. Capi ed, ca. 0.5 cm long, glandular pubescent; 2 mm gla- ng sub- 10). Petrea sulphurea is area in French Distribution (Fig. 10) nown from a very narrow P onl uiana, growing in clearings in primary forest. y Specimens examined. FRENCH GUIANA. km 80 Piste de St. Laurent a Paul Isnard, 15 Nov. 1982, Gran- 644 Annals of the Missouri Botanical Garden ville 2 MR XU Hi 2 p n Piste de St. Laurent à Paul Le Guet de la Crique R Ман sse, gee ig 1982, и > Jadin 1705 (В i Local names and u None reported. This is a dE E ces in the cladistic anal- ysis but close to the clade where P. bracteata is fo und. Pore ea decies еа is ш to the latter more than 15 mm m long, and petioles with two lateral However, P. sulphurea has calyx lobes of Р only species in the genus with yellow flowers. 11. нв Мер Li, Pu PL, ed. . TYPE uz, Mexico, кум s.n. riu BM, herb. , MO, TEX). Figure he Petrea arborea Kunth, Nov 21^ 2: 2802-283. — @ Z aip Ф S p a 800, Humboldt Bonpland s.n. (lecto: йылы on by Rueda (1993), P). Petrea racemosa Nees, sce 300. = E tes Ba = : Ri an —— оне 3 July- , Wi ad Newent n. (lec Tat x [e xm 993), BR; зенан С; photos of lec- type, MO, NY, TEX). Рана техісапа Cham., To i 367. i M 1829, Berlandier oe Rueda Ит С; 1 ; photo of ГАЯ T Cge ., Linn Sis . TYPE: (lec e чуч ч Rueda (1993), K; pen inolectotype, Petrea kohautiana C. Pre erk. 99. 1844 TYPE: Wi e, 1819-1821, 993), BR; ragment of isolec- a, Aug 94. "TM ted by Rueda (1993), BR; isolectotypes, BR, C(2), GH, LE, US; fragment of isolectotypes, NY; photo of лга апа каеп type, TEX). Petrea aspera Turcz., Bull. Soc. mp. € te osc. 36: =" ра ТҮРЕ: арт MM n Este elev., Apr. 5, Funck ry ich 507 itp “о P Ruck (1993), BR; isolec- totypes, BM, G(2), P fragment of isolectotype, a photos of АА Е, MO, NY(2), ТЕХ, 0, 2 б ч g SN = — +; d 9 i5 - È < Petrea vincentina Turcz., Bull. Soc. Imp. Nat. Mosc. 36: 212. 1863. TYPE: Windward Islands. St. Vincent, / s.n. uw o selected » ee (1993), G; ae otype, G; photo of isolec ple кай s raram Ru E. Mem w York Bo ales 338. 1927. TYPE: Bolivia n ps Rur L4 e S E 1, Rusby s pee NY; isotype, Petrea volubilis var. salir pres Moldenke Re- т. Bot. sot H, MO, NY, S; photo of ho- lotype, TEX). rae amazonica же ыа! E эы b 43: 173. 1938. : Brazil. Am: outh of the Rio Embira "19 dun e 1933, "Кл" 4930 (holotype, NY; isotypes, C. GH, K; photos of isotype, NY, Petrea pm E broadwayi Moldenke, Feddes Re- pert. 43: 26. 1938. TYPE: Trinidad. St. Ann's, 1908, Broadway s. с ы, UC; isotype, MO; photos of holotype, FAR Feddes Repert. 43: 195. . Antioquia: Río Porce, 7 May Petrea airocoerilea id 1938. TYPE: Colombia 1980, Kalbreyer 1634 (lectotype, selected by Rue- 8), fra da (199. gment-isolectotype, NY; photo- eect NY. TEX) Petrea colom a Moldenke, Feddes Repert. 43: 174. 1938. O Santander Sur: Висаг: E: ga, from a cultivated plant, 100 1927, Killip & Smith 19067 (holotype, Te types, GH, NY Petrea kohautiana var. anomala ers» Feddes b vU 43: 31. 1938. ium d x Clair, May 1911, Broadway s.n. (holotype, d nitidula Mo pa nke, Feddes Repert. 43: 168. ma E: Bra: o Uau , BR holotype, NY, TEX; pho isotype, Petrea СИ Moldenke, Pio Shu 43: e 1938 а: São Luis, Rio e s 6 . BM: 1915, Ducke 15 5822 (holotype, G NY, fragmen e, NY; photos of иэ ТЕХ). NM Petrea M Moldenke, P Repert. 43: 19 : Pon a. Waini r, Nov. 1910, Anderso е, Ki ie “К; fragment of holotype, uet holotype, TE Petrea sale Molde nke, Feddes Reperi 43: 192. бе ; TYPE: Brazil. P. o Tapajos, “a US; isotypes, BR, F di d gea x е: Moldenke, aer 1933. Panama. Apr Cooper 2 234 бони ep a, sometimes shrubby with stem са. 0 DBH; br anchlets obtusely a sr cent or glabro some ases petioles 1-16 mm long Ten pubescent; blades elliptic, 3-25 cm pass L cm wide, her apex short-acuminate, the base me ed or acute, entire, sometimes serrate ог den Volume 81, Number 4 Rueda 1994 Systematics and Evolution of Petrea Ficure 23. Petrea volubilis L. (Martinez 11375).—A. Habit.—B. Flower. 646 Annals of the Missouri Botanical Garden late, bear rough to the touch, glabrous or bescent. In miform, 4— О obed, ү La means. Bie wide, the ited: a 25 n mm i Res and 1.5-2 e apex a corolla blue or white, fragrant, 6-15 mm long, dn base 1.5-3 mm sometimes present; ovary 1.2- mm long, .9-2.5 mm wide, glabrous ч iid 2 4 mm long, glabrous; stigma subcapitate. Darum (Fig. 3) This the њеч AT RH southern half of Mexico throughout ille West In- ies to Peru and Paraguay. It grows in seasonal evergreen forest along streams, uius steep limestone walls in dry forest, limesto: one outcrops, pastures on u m, Capparis, Ceiba, Tabebuia, and Trichilia бе. e., a Predani | 49633), and the flowers are visited by honeybees (sub Nee & Taylor 26607). Sel MEXICO. Campe- che: Chasen, 20 km N „of toe a 29 Mar. 1982, C 18 (BM, MEXU). Oaxaca: Tuxtepec, Cerro с Aires, W of la Presa Temazcal, T m, 6 - do. Cortéz et al. 197 (MEXU). Quin- ana Roo tumal e road, 4 Mar. 1958, Schade rt & тск 1686 (СН, MEXU). Tabasco: B. lancán, on road to El Triunfo, 3 Nov. 1976, Méndez 441 (MO). Veracruz: А top road to Cerro de la Mesa, 1 km Ё N boa, 5 , 16 Apr. 1985 s m Mozom 0 m А Acosta & Acosta 245 (CHAPA). Yucatan: Tizimin , 24 km 1 pant et al. 130 ( MEXU) apaz: Rio Cahabon, Fi Sais = eis Izabal: Lake I Sneda е 5). San : Agu rman 77: 28« (F). Zacapa: Loma icach: do iva Rosalia, 1400 m, 15 Jan. 1942, rE 42728 da deor Belize: Gracie Rock, western hwy., 100 m, 21 Jan Doy esee US). Orang e W. Lago Tan and N of San = ntonio S Villa age, ^ ly e Lr 223 (MO). Toledo: trail to Esperanza, beginn N of Columbia ire € E June 1973, ee e M HON Ceiba, 1 May 1985, bados 1 , on road t , 2-11 Mar US) Santa A Chota, ca. 7 km si ‚ 19 Mar te cinity of Jinote ера, 1 К lig 10065 (F). Ze lye 2 vicinity of La 1 Bunting & Licht 649 wt H: US). COSTA RICA. Alajuela: Sa Ses Panamerican hwy., near Nicaraguan frontier, 100 аи а > = £g of its ia with Paine Va n & Ha hn 4247 (MO, TEX). Panamá: Serranê = ч ced min trail alo 0 m2 in Ver San eec rim 1924, UBA. тенни 10 Маг. 1926, /асЁ чыч 18 Арг. 1902, Hamilton 127 (NY). J MAI CA. Ca. 2 mi. NW of iie ‚о Mar : — 38629 (NY). DO Feb. 1946, ду 1028 (US). 1965, ает, betw n Rio Guapo and dr be " pide 300 m, ts 2 June 1977, bete y 1 Volume 81, Number 4 1994 Rueda 647 Systematics and Evolution of Petrea 13511 (MO, U). Zulia: Lagunillas, along Rio Gra nde, 13 km N of Embalse, 600 Е: 55 3 n E. MO, с: GUYANA. Essequibo, Itanime 1937, Smith 2148 (GH, аага f Fer мааа а ея Noy. 1976, маз is чуу ووو‎ ). FRENCH GUIANA. St. Laurent, abo outh of Maroni river, island in Moroni E near Апїесит Pata, 1 E ш, NT sides maque, Rio pires M e 970, Cavalcante 23% (NY, TEX). Rio Tur Ex nis 971, Hatschach ке а. Mania BO- LIVIA. Beni urrenabaque, 3 , Rusby 860A (NY, TEX) "PARAGUAY. May, es 1 1234 (6(3), GH). E DOR t & Pennin ngto ‚ US). Napo: ging National Park, Pozo Amo 2, trail to Rio Бау. 230 m, 14 Jan. 1988, Cerén 3320 (MO). es and uses. Because of the wide ана е Petrea ментте it Рз _known by umerous common names: Brazil, “сіро de Sao Miguel," “flor de Sao Jose," “ог de Say Miguel, е viuva,” “jas Toxo,” “touc e viuva,” b ' Colombia, “chaparro”; Costa Rica choreque”; Honduras, **chaparro," “flor del dia blo y e Jesus," “lengua de vaca," “а, m Š » > pecies is mostly used as an dei ; it is cultivated worldwide. In N are * used to feed domestic animals such as horses and cows, and in Venezuela the wood is used to make toys. Petrea volubilis is closely related to P. may- nensis (see discussion under th flo are known this species in s tivated plant s species is highly variable in leaf and flower т habit ent, the re the оле a iT d ange. nm ati HS when the ШАПА, Пош the entire Ешуа is observe LITERATURE CITED ABU-AsaB, M. S. & Р. " CANTINO. 1987. Go dota hee of leaf anatomy in subtribe 1 (Labiatae) and related taxa. J. Arnold Arbo ні 34. AGARDH, v P 1858. Theoria Systematis Plantarum. C. W iO RA on. AITON, W. T. 1812. Hortus Kewensis, 2nd ed., 4. od Би Rees, 0 rme Ea m London Anpray, C., R. WYL . LAFITTE, G. Privat & F. WINTERN NITZ 1982 of verb ide and oroban choside, caffeic acid s sugar esters from Oro- ban p rapumgenistae. Phytochemistry 21: 1123- 112 AYENSU, n 1981. Medicinal Plants of the West ndies. ые Publications, Michigan . HOOKER. 1876. Genera Plantar- on. 1895. Verbenaceae. Pp. n A. BRIQUET, J Engler & K. Prantl (itor Die cuis a Plan. тошу res 4, А W. Engelman, Leipzig. Cantino, P. D i ae. J. Arnold Arbor. 71: 323-370. uie s S. 1988. Suara Wood Anatomy. Springer-Verlag, Berlin. Ecueverry, В. 1984. Flora Apicola Colombiana. Li- to; dur a Arco, Bogotá. Farris, J. S. 1988. Henning86. Version 1.5. Doc mentation published by the author. State енй of New York, Stony Brook. 1989. The Viger index and ed rescaled consistency ra Cladist = 417-4 GARNIER, J., J. MAHUTEAU, МР & С. Pe RE. 1989. Esters caféiques ien deu eris p des Antilles. Plantes mower et Phytothérapie =н: 1-5. GENTRY, A. Central America: {езе and ecological com. Bio- tropica 8 -131. GIBSON, D. ая в. erbenaceae of Guatemala. Ы 70 у& 1. О 0, Wiliams Аякез Flora ag Fro Bot. 24: GoLpBLarT, Р. . Index to Plant e Num- bers “1985, Monogr. Syst. Bot. Missouri Bot. Саг 214 Hewson, ü т 1988. Plant Indumentum: A Handbook of Termi inology. Australian Government Polling be: 9. revised classification of the archite сше of dice дена leaves. Pp. 25-39 in C. Metcalfe & L. Chalk (editors), о of the 648 Annals of the Missouri Botanical Garden еек 2nd ed., Vol. 1. Clarendon Press, Ох- for: номт, Е. H. A., А. J. Вомрглмр & С. S. Kunta. 18. Nov К Com era et ia Plantarum. Vol. 2 282. ies Parisiorum, Parr Jansen-Jacoss, M. J. 1988. V ая In: A ijn, editor, Flora of Guianas. =н dia tific ‘Books, Koenigstein. 1971. cape in space by Sterculia apetala seeds from the bug Dysdercus fasciata in a Costa Rican deciduous E Ecology 53: 350- JOHANSEN, D. 1940. Plant Microtechnique. McGraw- Hill, New York. e S 1994. Zu r Gynäceummorphologie und vid matik ба понео und Labiaten. Symb. Bo Upsa 1. 4 ne к D. The concept of homology and its entral Bie in 4 pee of pe systematic relationships. Pp. . Ste eussy & T ncan (editors), Cla Axe Р, th Riise fe of КЫШТ History. ‘Columbia Univ. Press, New York. LINNAEUS, C. 1787. L ж i Batavorum ‚ Leiden ў Species Р Plantarum. Vol. 2: 626. Lau- ntii xr Stockholm Machu Т.Е. 1960. Flora of Peru: Verbenaceae. Publ. bog Mus. Nat. Hist. Bot. Ser. 13: 60 E 720. DDISON, W. P., M. J. DoN D. DDISON. 1984. Outgroup analysis and parsimony. Sni. Zool. 3: 83-103. METCALFE, С. R. & L. CHALK. 1972. Anatomy of the ( Dicotyledons, Clarendon on Oxfor . 1979 atomy of dé Dicotyle- dons, 2nd De bed x Clarendon Press, Oxford. e jeer of the genus Petrea. elie му den 1-48, 1 T: A Fifth ameti of s Verbenaceae, ET Sti Dicrastylidacea: Genera Plantarum, ed. 1: 347. eae, ў phoremaceae, Nyctanthaceae, and Eriocaulaceae of the world as to Valid Taxa, and Synonymy. Published ba d 3. Verbenaceae W R. W. Schery йо n lora of а А: issouri. Bot. Gard. NIXON, К.С 92. Clados version 1 Eh Ini ү 0 patible EN fM progra ation published by uthor. Cornell Vei ы са. PAYNE, 19 A glossary of plant hair termi- nology ji us 30: 239-255. аре H 925. Arboles y arbustos nuevos de Ven a Во. Ci. Técn. Mus. Comercial Venezuela. 1: 70. Purr, С. The peculiar trichomes of Petrea vol- mt y rG Galore J. S. African Bot. 44: 119- MR. "s 1982. Natural уени of Lianas and their Influences on Tropical est Dynamics. Unpub- lished Ph.D. thesis. Cornell. ролет Ithaca, New Yor 41. American woods . Wood 65: 4-21. Petrea фый Novon 2: 417 new species of benaceae) from едик Amazonia. ————. 1993. Le fend pg oe in the genus Petrea mre Novon 3: 179-181. prede JC . Verbenaceae. In: А. L. P. Can- ‚ Prodromus roe Naturalis Regi Vege- is 11: 616-62 er B. "Tante acid and iron alum with safranin and orange С i А n studies of the shoot apex. Stain Technol. 18: 10 SINCLAIR, C. B. & D. B. in ool: Surface printing id fae toh wet iride studies. Stain Tech- 36: 2 T m "owe 1979. Taxonomic Lit- Bohn, AR & Holkema d F.A ature, 2nd ed WATROUS, L. & Q. W 1981. The out-group көче ipo dedu of ir analysis. Syst. Zool. 30: E WELLE, B. J. H. & P. DETIENNE. 1988. Von ш timber of Verbenacea e. Pp. 87-99 in A. R. A. Gorts van "e editor, Flora of Guianas. ds Scientific oenigstein. OMS Ww. & D. TE Vids Me dical e he Quijos-Quich upper Río Napo, Am- azonian Ecuador. кб) 3$ the b List ОЕ ExsicCcATAE. Numbers in indic cor- respond to species as numbered in the te: 4239 rai 4840 (2). Acosta, R., 245 0 (11). A pep m , 1212 (1 €— i B., , 2920A, dr qe (11). Acevedo, P., ar, 1. чү ы п. (11). Andra e x G., 9 2) ve E. F., 11). A 7 }} 1674 (8). Arnason, T., 1704 363 (11). Artamanoff, G., d 4, 17 (2). Aviles, S., A (11). iE Е. „ 2894, 4. 4509 (1) Аз С., 99, ‚ 1865, 2585, 4446, 6569 (8); w а dA o = N EIC o со eo с 508 с eo № { pes: iva m, W.N., Ат 1). Basiai 6i м d boza, H. 117 nu ary, A . S., 3461 (9). Panis G., 7146 (7). Barkley, F. A., 17M174, 401 38C494 (9). Barreto, M., 3264, 3267, 3268 (11). ue lett, H. H., 11494, 12133, 12566, 12559 (11). Basurto, F., s.n. (11). Beard, P., 1071, 1475 16699 7). Roles Кыл 7 (11). Belshaw, C. (11). Bena, P., 1227 (11). Bendeck, C., 180 (11, Ben CPE: 150. 2199 (1) std n ). Benson, C., Be „ 180 per Ay т" С.А. 1705 с pied F., Poem (2. Bla ek, fi ә 90, 112 (11). Blackmore, S., 4035 (11). Blanchet, Volume 81, Number 4 1994 Rueda Systematics and Evolution of Petrea (11); s.n. (1). Blanco, C., 145, 499 (11). Blum, К. E., x Boer, P ы Ww. 642, 960 (2). Boke, N. H., 11). Bond, E 0 (9). Boo; Вов; J, Non E аа Bourgeau, M., 212 (11). Во 5 (11 1033 (11). Brade, А. С, жүр 18046 (11) Brana 11). Bravo, H., 11). Breedl 9925, 10681, "HAM posa 33771. aes ri dr 49835, 50472, 50223, 56898 (11). nt 14290, 20462, 20468, 21462, dm er, = = Е б mU و‎ S ا‎ m = 25 (11). B 140 (11 7 (1). Bunting, С. S., 649, 7129, 8808, 11278 (11); =н Вигсһе Il W. 74 7539 (11). Burger, W. C., 5190 4425, 10857 (11). Calderón, eon 115, 2 268 (11). Cald ton, Y., 115 (11). Caley, G., s.n. (11). Callejas, R., 58 (11). Calzada, J. I., 175, 5300, 9604, 9769 (11). Camp, W.H., El (8). С Psa "ER met Meri J Carlson, M. C., 503 ‚ 1940 ( cci-Buzi, V. P., 149 (11). бта, T An diem А., 114 (11). vp ior a ү ер е on 695, 1911 (11). Ege e O3 (11). e P., 289, 3297 To 2302 an. ones P. C., 7649 (11). Cerati, T. M., 11). С а, S. C., 1 (11). Chiang, E, 343 (11). Christ, ы Н. W., 4436 (11). Ga, c A., B 1481, ios (2); 6224 (11). Claussen, P., 625. 32, s.n. ын а. А., 1399 (11). Collins, v N., 11). Converse, on ra 1). mae Gs Yo 5305 m E cg Cooper, G. P., pires T: кч us 1 (5). Cordeiro, M., 220 (2). rni, eL 7., 833 m "ee E 365 ap, Con, es 1). Cowan, В. S 90, 291, 292 ш 3130 (11 Cuatrecasas gi 6577, ve (8). Cuni, A , "М Сип 11). Custodio, A., 3 р W. G., 11388 туў [une Bro., 2665 (9). 947 (2). Darwin, S., 2075 (11). Dauben- Davidse, G. ed (9); iea Davis, H: C., 199, 6370 (11). Defi “dried р до 3 11) Delg x .n. (11). Djoemadi C. W., 6474 (11). Dodson, C. H., Disc n, 2406, diy 3217 (2). E. d ез, J., 197, 520 (11). Dorr, L. J., 5040 (8). Drake, Dick NON 2172, 2536 (1). ses a E., 2303 (9). ске, A., 140 (3); 688 (5); 872, 1133, 1982, 7971, Res TREBE AR ALLE 3267... 3914, : 9303; p 4294, 15451 dina Pr s.n. (7). Duenas, C. C., an, Dupre, M., Duque-Jaramillo, J. 2426 di 4340-A (9). Dusén, P- у» с Here s.n. 11). Duss, P., 1979, 0 (11). D = D7208; аб а, ги ns 12022 a 1). wood, A., s.n. (11). gx 194 (9). Edwall, P58 = Я we (11) E Ве он (11). Egler F. E., 4293 (11). Eijnatten, C., 1102 (11). idus E. L., 13050, H9121 (11). Elias, s. 357; 1201 (11). 11). Emigdio, M., 255, 746 (11 (11). Enriques, O. 396 (11). n ^n 5 (11). Espinal, S., 2273 E 1831, 1870 (7). (9). Espinoza, R. 1 D pruna v ти Fairchild, Bel aie (2). Fendler, ^x 557,8 ae Fernandez, A "s Fernández, 85 (11). E R. Pe бея ge ix: 65 Ci айа, э: 270, 564, 593, 736, 737, a ec Сы "ue ten: 1678 (8). Flores, 2 3 (11). Florschutz, J., 169, -A (11 pr er 25871, 22628 (2); 21121, ў га mm, 2 (11). Frost, S. W., 196 (11). Fryxell, Sn qu Fuentes, M. 53 (11). Funck, N., 507 ay 1504 eotti, H., 193, 5 (11). Gandolfi, S., 6 (2). en 010078358; д (9); 5206, 12344 л урат asi, n 12344, A Eod are 2 (8). Garcia, i 6 (11). Garcia, M. V., 18 (3). Gardner, G., ША v Garwood. WUN ‚ 721, aie (11). Gaudi- cha M., ARE Caumer, F. G., 379, 23585, 23673, Ыы A Gent tle, P. H., 316, 2366, 4972, 8867 (11). Cont , 2164, 3181, 3187, 3228, 3232, 3330, 3331, wily 3289, 5582, 7191, 7794, 12728, 18250, 28575, 28928, 32506, 37844, 57138, 59132 37075, 21721, 22418, 20982 (1): 13325, 54345, (2); 57827, A eis d ul FS О ҮҮ 71866 (8). Gilly, Glaziou, A., 2652, (11), Guedes, T. N., 496 (11). G E ( Guevara, L. C., 2288 a 4420 (11). Cate % 760, 781 (1); 783 (7). Guzman, J., 28 (11). Guzman, M., 108 (11). Haenke, T., 1582 (11). Hahn, L., 270 W., "n (6). Hal Е., 516, 747 (de 585 Е паве Н., С119, С120 (11). Hamilton, S., 127 (11). а В., 1868, 12303 (11). е is ара rriman, 11234 (11). 18074, 24863, 32638, 3 (11). Haught, O., 2103 (9); 1328, 1568, 4046, 4569, 4791 (11). 650 Annals of the п Missouri Botanical Garden 11). Hayes, S., к 1). Heiner, A., 206 (11). Held, J. J., FC88 (11). hs endorff, s.n. (11). Herinberger, E «s 428 (11). Не, ES dr (11). Hernández, A., 195 (11). Her i 3 (11). Hernández, J., n (11). Hernández, p I 4 (11). Herrera, A., 67 (11). Herrera, H. 1186 (1). с уде, Е. 55007. Bin rne Б X" 15, E и жоет Higgins, J. e i n, G. B., . Hite 9 (2 2) fois 984, кы 3102, с ost = yy F. ©, Hh т 28316, s.n. (11). Holt, E. С. A: a 572 (8). Hos ostmann, F. W., з9(2 11303, 11757, 18202, 18203, aig 195 36 (1 1). Ни, S. Y., 12961 (11). sons ДЕА LA AA 2 2 (2). H M., p He Hugh-Jones, D., Sa Humboldt, F. H., s.n. (9); s.n. (11). p ко» D. n. (11). nier A, A 658 (1). "Hurd. RR 72(11) Hutchison Аж 14 Pon ВА М., 2041, 10999 20 i 0x dois (9) m Н. 5, 456 (2). i. Ж Q1) Ji nêz. J E 028 (11). NM EM iu Johnston, J. R., 11 м ie iei 212, А4505, 652 (2): Jud 11). sete Be 4568 (11). А жаен = 28 (i), Kahn, F., 5400 (1). Kalbreyer, W., 4 (11). Kar- winsky, W. T., 709, uns: 1). Ke ieu 7738, .n. 8 (11). T E е (11). Pan ps oyer, L. 6:627). овог, 37068, 43554, pe 19067 174 J. КЗ 96 (11); 2:5 Kuhlmann, M., pm ^ 1). Kumm- 6 (11). ng, H., 87 (2). Lanjouw, J., 852, 1298, 1321, oes (2). Lanna, S., 338 (2). Тан ughlin, К. М., 178 (11). L опа, М., 270 (11). p G., 2525 (11). 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SYSTEMATIC IMPLICATIONS OF POLLEN MORPHOLOGY IN SUBFAMILIES LAMIOIDEAE AND POGOSTEMONOIDEAE (LABIATAE)! Mones S. Abu-Asab? and Philip D. Cantino’ ABSTRACT Pollen din was surveyed in 57 of the 60 genera of Lamioideae iur Mice siye sensu Can din nd tw of uncertain subfamilial affinities: Anisomeles and Eur Pollen all esed ost Lamioideae, Anisomeles, and Eurysolen i is tricolpate with suprareticulate кый Шш! “Variants found in in a few granu Most species | in which pollen was examined in section have simple columellae, but se mingly Gi elonopsis anched colume ellae br With ae to the genera of uncertain affinities, the fin ings. presented here support a Somes between Anisomeles and Pogost emon; the рові tio п of тти unclear. Within шау Pes amio; oidea i ai mis, agopsis Marrubium ach; ys, respec tively. Hexapantocolpate pollen Citi to de a synapomorphy of nies section Empedo: ses ea. одани ties іп e sc ulpturing were noted een Brazoria, Macbridea, and Physostegia, and between Galeopsis and Syna 4 pollen Suey of subfamily Lamioideae sensu Erd it might ght justis our understanding of systematic relationships. rief overview of the study as whole has bee published (Abu-Asab & Cantino, 1992), w , those genera with taxonomic бшнк is needed because we mioideae differ- k РА of the Lamioid Erdt (1 (1945), ane our tier papers adopted Erdt р 1 PCR АК ә es & Cantino, 1992; Сш 1992b) led to an alternative classification (Cantino et al., 1992), шо 15 followed here. Unies тегине цеп, кР the circumscription of Cantino ‘et al. (1992). amily Lamioideae comprises 55 genera, the e a ily Pogostemonoideae is si „Based on morphological and molecular ev- related 1(C tino, 1992a; Downie Palmer, 1992), a Eu- ‚ (бее authors of genus names oad Appe for authors f specie es.) Anisom s been variably — to both subfamilies (see Discussion). ollen studies that members енын and Pogostemonoideae have 2 on particular genera (e.g., Huynh, 1972; Bassett Munro, 1986) or geographical regions (e.g., Waterman, 19 arghese 1968 L/1054 uang, 1972). R conducted broader surveys of the Labiatae, but Risch used only light microscopy, and his descrip- ubmitted to Ohio qi di in We are most С for Joan Nowicke’s This p 1990. The 15 б portions of the st tudy were competed i in the Pali gemein at and encourag per BM, “GH, "k, , MO, US. Financial s support for this е the Smithsonian Ins titu tion: t. We thank the following institutions ke to remove pollen from herbarium imens: A, BHO, rise mt x^ National S Science F а g 'BSR-83 * Dep " requests should be se to the cu au Birzeit University, P.O. Box 14, Birzeit, West Bank, via Israel. Reprint uthor * Department of Bonet and Plant ‘Biology, Ohio University, Athens, Ohio 45701-2979, U.S.A. ANN. Missouri Bor. Garp. 81: 653-686. 1994. Annals of th Missouri i, MSN Garden TABLE l. Genera of Lamioideae and Pogostemonoide- атны, їп quesiti an number of s е cies in the ). N geuus/nuraber 1 (A 1i umbers of TI 1 2 P variety РАС 8 E Mah} , geographic 1 Airy Shaw 1 f 1 (1027 M А (1973). Subfamily Lamioidea delyrermum В Blume (10/6)— Paleotropics Acr e Benth. ex Endl. (8/4)—Tropical & south- ern a ities Ajugoides Makino (1 dom bud Alajja Ikon Soca Ballota /T)— Bua eae, western Asia Basrychantera ipm ed 0)— China Br ay (4/4)— an Oklahoma Chaiturus Wila. ал Tm “Tempera Eur: r& С е: (1/ Fisch ote ral & southwe: рн Mi ig. (1372) наат China, Japan Colquhounia Wallich (5/ "NM Craniotome Rchb. (1/1) — Himalay Eremostachys Bunge ex Ledeb. sensu hes (60/4)— urope to en Asia h. (1/1)—Himalayas (10/ 6)—Temperate Eurasia athe d Wallich ex Benth. (30/7)—South- eastern Asia, Malesia Haplostachys (A. Gray) Hillebrand — Haw. Hypogomphia Bunge (1/ cub mee ну & rit rn one I. pie chilus m ex БО ico ЧЫ Central & southwestern Asi Lagopsis Bunge (4/2)— Western Siberia to Japan Lamiophlomis Kudo (1/1)— Himalayas Lamium L. (16/7 )—Temperate Eurasia, northern Africa Leonotis (Pers.) R. Br. шы 3)—Tropical & southern Africa, 1 pantropica Pier L. (24/2)— т. mperate Eura: rx ii (100/ oe ge Hes Afri- Loxocalyx ae emsley (3/2)—China, Japa: Macbriden mes ex Nutt. (2/2)— Pedir nited Stz Marrubium L (30/5 gua northern Africa, central & southwester: Melittis L. (1/1)— Euro Metastachydium Airy Shaw ex C. Y. Wu & Li (1/ 1)— Central Asia Microtoena Prain (25 /4)— Himalayas, Chin: Жыры. lla L. (2/2)— Southern Europe, AGERE ern А: уба Benth. (1/1)— Himalay ose ia eei i Semere tropical Afri- a to ral A Panz zerina ирей! (2/1)— Western Siberia to Mongo- Paralamium Dunn (1/1)— Southwestern China TABLE 1. Continued. Paraphlomis Prain (8/3)— Eastern Asia, Malesia Phlomidoschema (Benth.) Vved. (1/1)—Iran to western Himal Phlomis L. (ca. 100/7)— Mediterranean region to China Phyllostegia Benth. (2 qu Там = =. hysostegia Benth. (1 —North пла Prasiu. 1/1)— Maca regi Pseuderemostachys Popov (1/1)— сай Asia Pseudomarrubium Popov (1/0)—Central Asia Roylea Wallich ex Benth. ( — Himalayas Sideritis L. (ca. 130/19) gion ti southwestern Asia, Macaron ce dex ee & Vved. (37 туану Asia to wester cle 1. (са /2)— Northern E: southern tem- e& ох tropical mountains E Benth. (20/3)— Hawaii i Rech. f. (1/1)—Pakistan Syn andra Nutt. (1 Thuspeinanta T. Durand wester Ратна Fischer & С. Meyer (2/1)—South- western Asia d St (2/1)—Central & E. Subfamily Pogostemonoideae MT Smith (1/1)—In Com sph е 5. Мооге (1/3) Base Asia s, China Rostrinucula Kudo (2/1)— Eastern Asia Genera of uncertain affinities Anisomeles R. Br. (6/4)— Paleotropics Eur ysolen jan. (17 1)—Southeastern Asia, Indoma- les l 11 Lee hen dum shape, color). Pozhidaev’s (1989) survey used ectron microscopy (SEM), but p в described, and micro: aripa V were provided for less a third of the gener. 1 e are "e of published SEM pollen micrographs for uly o 60 genera of Lamioideae a Pogostemo- posed (Nabli, 1976; Roca Salin 978; Атлап re, 1982; Cantino, 1982; Ubera & Galan, e degree of intrageneric variation in i sculpturing is unknown for most of these ge Volume 81, Number 4 1994 Asab & Cant паа Morphelogr ih in Lamioideae and Pogostemonoideae exceptio e Eremostachys, Paraphlomi [Ай Physostegia, and Stac hys. Transmission ape for only ! three genera (Nabli, 1976; Abu- E Cantino, 19 he objectives of this paper are to documen n te ШО of nh ie Lamioidee ано within most of the larger genera, and | to data. Because е the resulting compilation of SEM the Labiatae MATERIALS AND METHODS udy orien едын 1) comprised 183 of Po five genera goste monoideae = re e pê genera of Lamioideae, and two gen uncertain subfamilial affiniti (Anisomeles and Eurysol n) t 1 © genera, ап effo 5 made to заара а broad sp ect f 1 classifications of Briquet 518392 ne and avail- aphs . Eremostachys, B 8 z © Cantin & Galan, 1983; чын & Munro, 1 ew & Harley, 1992). The monotypic genera Bostrychanthera, Pseudomar- rubium, and Isoleucas were not ex for lack Я, z m л © і) <) © Ф і | Ф a X 5 т = - |ы! = E. ея of рош to equatorial axis leng th (P/E) was nd the mean E ratio was calculated ise om théde individual values. For SEM, pollen was coated with gold- = oom and and examined and photographed n a TEM, 7 pole was treated wh kan wee Stain lead citrate, and examined and photographed with a Hitachi HS-8 transmission electron microscope. RESULTS ults are summarized in Table 2, and е ыш follows Veze our previous pap Asa b n Cantino, 1989, 1992, 1993a, b; АБ Asab et , 1993). Pollen that we previously referred to s elite — is described here as scrobic- te or punc LAMIOIDEAE n of subfamily Lamioideae is е to ais (shape classification follows Walker & oyle, 1975, based on P/E ratios in Table 2), with the polar axis 15- 59 ит and the туч s 13-52 um. The rc dy Hw tetracolpate and h pate grains were found in a few genera (Table 2) The e exine in most species is scrobiculate (some- ing on micro retic culate or dep te). The exine is леу te in species, re- and partially oveolate i in one (Table — sg rugulat ugulat enera warrant a more detailed description das is available in Table 2 Go л emma (Figs. 62-76; Pozhida 1989): Jti ти exceptionally variable: psilate or with suprareticulate, rugulate, or granu ulate sculpturing; columellae branche Lagopsis (Fig. 77): exine rugilete; psilate to- ward the poles, ms nonperforate. Sideritis (Figs. 18 6; Huynh, 1972; Nabli, 1976 6; Roca Selinas, 1978; Poca беи tet 6- nai (the om circ i: or ra disp tii colpate (the amb triangular); exine psilate (ap- proaching a supraretieulate condition in section d irs in e 192, 193), usu- aev, sections were collected о‏ ا ormvar, stained with а uranyl acetate and‏ ый . ally la (i montana, the polar regions са centers of the mae тыры foveolate, the Summary of pollen morphological data for subfamilies eee simple vs. branched. Sculpturing: Upper la >P (or only layer if psilate): Fo, foveolate; Mi, microreticulate; Np, моды 0 Pu, punctate; Re, reticulate; Sc, ЫЗ ш. slash indicates borderline ыан опз. TABLE 2. hexapantocolpate. Columellae: er: с пера | silate; Ки, г апі a and genera of unce ugulate Ln bros ce -rugul in affinities. brun 8; pee: 4, orm жі 4 ate; Su, suprareticulat Lower lay: er Polar axis (um) Equatorial axis (um) P/E ratio Sculpturing Species Mean Range Mean Range Mean Range Colpi Columellae Upper Lower Figures Subfamily Lamioide: Achyrospermum aethiopicum 21.3 (20-22) 18.5 (18-20) 1.16 1:0-1.3 3 Su c K2 Achyrospermum carvalhi 209 — 22-31) 23. 20-29 133 0.8-1.3 3 Su Sc/Mi 3 Achyrospermum cryptanthum 24.0 (22-31) 19.1 18-22 1.25 1-14 3 Su St 4 Achyrospermum densiflorum 24.5 (22-26) 18.4 15-22 1.34 LET 3 Su SD 5 Achyrospermum parviflorum 23.3 (22-24) 18.9 18-20 1.24 1.1-1.3 3 Su 5с 6/7 chyrospermum schimperi 28. (22-33) 20.9 20-22 1.34 1:11-1.5 3 Su Se 8,9 Acrotome angust 20.2 (20-22) 18.0 17-20 145 1.0-1.3 d Su Sc 10, 11 Acrotome flecki 20.7 (20-22) 22.0 20-24 0.94. 0.8-1.1 8 n ? Acrotome hispida 20.0 (—) 21.6 20-22 0.92 0.9-1.0 2 ? ? Acrotome i 22.9 (22-24) 22.9 22-24 1.00 0.9-1.1 3 ? ? Ajugoides humilis 31.9 (31-35) 23.5 22-26 1.36 1.2-1.6 S Su Sc/Mi 12, 13 Alajja rhomboidea 30.1 (29-33) 28.4 26-31 1.06 0.9-1.3 5 Su c 14, 15 llota africa 22.7 (22-24) 20.7 20-22 1.10 1.0-1.2 3 Su с 6 F ota andreuzziana 23:5 (22-24) 20.9 20-22 Las 10951.2 3 Su Sc/Mi Ballota hirsuta 22.0 (—) 8.0 17-20 1:29 1.1-1.3 3 Su Sc 17, 18 Ballots integrifolia 21.6 20-24 2153 20-24 1.01 0.91.1 Б 5и Mi Ballota nigra 23.8 22-26 20.9 18-24 1.14 1.0-1.3 3 Su Sc Ballota pseudodictamnus 29.3 26-31 20.5 18-22 1.43 1.3-1.6 3 Su Sc/Mi 21. 22 Ballota rupestris 21:2 20-22 23.4 22-24 0.90 С) 3 Su 5с Brazoria arenaria 39/9 33-40 2d 31-33 1.09 (1.0-1.2) 3 Su Sc 23 Brazoria pulcherrima 37.0 35-42 317 29-33 IT (1.1-1.5) 3 Su Sc 24 Brazoria scutellarioides 32:5 29-35 33:0 29-35 0.99 (0.8-1.2) 3 Su т 29 Brazoria truncata 39.2 31-42 32.6 29-35 LL (1.1-1.5) 3 Su Se 26, 27 Chaiturus marrubias 212 20-23 20.5 18-22 1.04 (0.9-1.2 = Su Sc 28, 29 Chamaesphacos ilicifolius 30.4 (24-33) 24.0 (20-26) 121 (1.1-1.4 3 Su Sc 0 Chelonopsis lichiangensis 28.4 (24-31) 213 (24-31) 1.04 (0.9-1.2 3 Branched? Su Sc/Mi 31:32 Chelonopsis odontochil 28.0 (26-29) 26.4 (24-29) 1.06 (1.0-1.2 3 Su Sc/Mi 33, 34 unia cocci DU (26-31) 26.6 (26-29) 1.04 (0.9-1.2 3 Su Se uhounia seguinii 2353 (22-26) 21.6 (20-24) 1.09 (0.9-1.3 3 Su Sc 35, 36 Craniotome furcata 16.6 (15-18) 14.9 (13-15) 132 (1.0-1.3 3 Su Sc 37-39 Eremostachys fetissovii 33.4 (31-37) 29.5 (24-33) 1.14 (0.9-1.6) i Su 5с Eremostachys iliensis азыт (31-33) 20k (24-29) Tz (1.1-1.3 3 Su 5с 40, 41 иәрео jeoiueyog HNOSSIN eui jo sjeuuy 999 TABLE 2. Continued. Polar axis (um) Equatorial axis (um) P/E ratio Sculpturing Species Range Mean Range Colpi ^ Columellae Upper Lower Figures Eremostachys rotata 34.5 29-37 32.3 29-35 1.07 0.8-1.2 3 Su Sc 42 Eremostachys speciosa 35.9 33-37 29.5 26-33 1.23 1.1-1.4 3 Su Sc 43 Eriophyt llichi 28.6 26-31 23.5 22-26 1.22 1.0-1.4 3 Su Sc 44, 45 Galeopsis bifida 38.7 33-46 3931 26-37 1:22 0.9-1.4 3 Su Sc 46, 47 opsis ladanum 34.9 31-40 26.7 24-31 1.31 1.2-1.5 3 Su Sc Galeopsis pubescens 32.1 26-35 26.0 22-31 1.24 1.0-1.4 3 Su Sc 48 Galeopsis segetum 35.2 31-40 26.8 24-31 1.32 1.0-1.6 3 Su ? Galeopsis speciosa 37.8 35-40 27.3 26-29 1.39 1.2-1.5 3 Su Sc 49 Galeopsis tetrahit 38.5 35-42 29.0 6-31 1.33 1213125 3 Su Sc 50 Gomphostemma 33.7 31-35 33.4 33-35 1.01 0.9-1.1 3 Gr Sc/Mi 62, 63 Gomphostemma intermedium 32.1 31-35 32.6 31-33 0.99 0.9-1.0 3 Branched Ps Mi 64-66 Gomphostemma javanicu 29.7 26-31 30.8 29-33 0.97 0.9-1.0 3 Ru Sc 69, 70 Gomphostemma leptodon 33:2 29-40) 35.4 29-42 0.94 0.9-1.2 3 Ps Sc 67, 68 Gomphostemma 22.9 29-37 32.6 31-33 0.99 0.8-1.1 3 Branched Ps Sc 71072 Gomphostemma parviflorum 26.6 24-29 25.7 24-29 1.04 OS-t 3 Gr Sc 73,74 ostemma wallichii 299 :(29-31) 31.7 31-33 0.95 0.9-1.0 3 Su Sc 75, 76 Haplostachys haplostachya 51.0 (44-57) 48.4 46-51 1.06 059-152 3, 4 Su 5с 51,52 aplostachys linearifolia 49.7 (46-55 46.2 44-48 1.08 1:0-1.2 3,4 Su Se 53, 54 Hypogomphia turkest 33.4 (29-40 27.4 24-29 1.22 20-15 3 Simple Su Sc 55-57 сг шип: 25.5 (24-26 24.2 22-26 1.06 1.0-1.2 3 Su ? 58, 59 thophy 28 26-2 22.7 20-24 1.25 1.1-1.4 3 Su Sc 60, 61 үре hirtus 25.5 22-29 21.1 20-24 1.22 0.9-1.3 3 Su 5с Lagopsis marrubiastrum 21.2 20-23 19.6 18-21 1.08 0.9-1.2 3 Ru- Ps Np? 16 opsis supina 21.1 20-22 21.8 20-24 0.97 0:011 3 4 ? Lamiophlomis rotata 24.2 22-26 20.0 18-22 121 1.1-1.3 3 Ps Se 78, 79 Lamium album 25.1 22-29 23.8 22-26 1.06 0.8-1.3 3 Gr 5с 80, 81 La exuo 23.5 22-6 21.8 18-24 1.09 0.9-1.3 3 Simple Su Se 82, 83 Lamium к 30.8 29-3 22.9 20-26 1.36 1511.6 3 Su Sc 86, 87 Lamium garg 28.6 26-33 26.8 24-2 1.07 1,0-1.2 3 Simple Su Mi 84, 85 Lamium какы n 34.1 33-35 30.1 26-33 1.14 1.0-1.3 3 Su ? Lamium moschatum 21.2 20-22 20.6 20-22 1.03 Oa а 3 Gr ac 88, 89 Lamium 3 Su Se 90, 91 Leonotis bequaertii 30.7 (29-33) 29.9 (28-33) 1.02 (0.8-1.2) 3 Su Sc 92, 93 Leonotis leonitis 28.6 (26-31) 24.9 (24-26) LI» (1.1-1.3) J Simple Su Sc 94-96 Leonotis mollissima 31.9 (29-35) 28.6 (22-33) EIS (0.9-1.5) 3 Simple Su Sc 97 eeepiououie1soDog pue aeepioiue ш Абоюцаоу иәһоа 7661 ӯ зәашпм “18 eunjoA ounue?) $ qesy-nqv 199 TABLE 2. Continued. Polar axis (um) Equatorial axis (um) P/E ratio Sculpturing Species Mean Range ange Mean Range Colpi ^ Columellae Upper Lower Figures Leonurus cardiaca 22: 22-24) 213 20-24 1.08 1.0-1.1 3 Su 5с 98,99 Leonurus sibiricus 25:5 22-29) 21:8 20-24 1.20 1.0-1.4 3 Su Sc 100 Leucas abyssinica 20.5 18-22) 19.1 18-20 1.07 1.0-1.1 3 Su 5с 107, 108 Leucas alluau 26.0 22-31) 22.9 20-26 1.14 1.0-1.4 3 Su Se 09 Leucas aspera 24.0 22-26) 19.6 18-22 1:28 1.1-1.4 3 Su Sc/Mi 110, 111 Le biflora 29.3 26-31) 23.1 22-26 1:27 1.1-1.4 g Su Sc Leucas calostachys 25.3 24-26) 22.4 20-24 1.14 1.0-1.3 3 Su Sc 112, 113 Leucas ciliata 28:3 22-24) 2241 22-24 1.03 0.9-1.1 3 Su Se Leucas eriostoma 20.9 20-22) 126 15-20 1.20 1.1-1.4 3 Su Mi 114 Leucas hirta 20.5 (20-22) 16.7 15-20 1.24 1.0-1.4 3 Su Se 115 Leucas infl 20.0 (— 18.1 17-20 1.10 1.0-1.1 3 Su? ? 116 Leucas javanica 26.4 (22-31 24.2 22-26 1.09 0.9-1.3 3 Su Sc/Mi 117 Leucas lanata 23:3 26-29 20.5 18-22 1.34 1.2-1.5 3 Su D Leucas eee 26.4 24-29 22.7 20-24 Lely S13 3 Su Sc Leucas marrubioides 22.9 20-24 17.8 15-20 1.29 1.0-1.4 3 ? ? Leucas martinicensis 22.6 22-24 20.1 19-22 1713 1.0-1.2 3 Su Sc Leucas mollissima 29.6 29-31 23.0 22-24 1.29 1.2-1.4 3 ? y: cas rosmarinifolia 22.2 20-24 18.5 18-20 1.21 1.1-1.4 3 Simple Su ? 118 Leucas “р ега 2131 20-24 18.9 18-20 1512 1.0-1.3 Я Su Se Loxocaly. 23.8 22-26 21:3 20-22 1:12 1.0-1.3 8 5и Sc/Mi 101 Loxocalyx utils 24.0 22-26 20.9 20-24 1.16 0.9-1.3 2 Su Sc 102, 103 Macbridea 41.3 42-53 34.5 31-37 и 1.2-1.5 > Su Se 104 Macbridea nu ee 38.1 35-42 40.5 40-42 0.94 0.9-1.1 3 Su Se 105, 106 Marrubium anisodon 2155 24-31 29.0 26-31 0.95 0.8-1.1 4 Ps Sc 1920; 123 bium cuneatum 24.9 24-26 26.8 24-29 0.93 0.8-1.1 3 Ps Sc Marrubium heterodon 26.4 24-29 2791 26-29 0.95 0.8-1.1 3 Ps Sc/Pu 124, 125 Marrubium incanu 52.3 51-55 46.9 44-51 1:12 1.1-1 3 Ps Sc/Pu Marrubium supinu 24.0 (—) 25.9 24-26 0.93 0.9-1.0 3 Simple Ps Sc 126-128 elittis melissophy 45.8 (40-51) 35.6 (31-40) 1.29 1,2-1.5 3i 5и 5с 119, 120 Metastachydium sagittatum 222, (22-24) 23.1 (22-24) 0.96 (0.9-1.1) 3 Su Sc 121 Microtoena delavayi 3 Simple Su Sc/Mi 129-131 Microtoena insuavis 3 Su Mi so Microtoena robusta 32.8 (29-37) 28.8 (24-33) 3.15 (0.9-1.4) 3 Su Mi 133, 134 Microtoena ондо 24.6 (22-26) 28. (24-31) 0.88 (0.8-0.9) 3 Su Mi I35 r396 Moluccella laev 34.0 (33-40) 28.2 (26-31) 1.24 (1.1—1.5) 3 Su Sc/Mi 37 uepJer) [eoiuejog unossiN 899 əy} jo sjeuuy TABLE 2. Continued. Polar axis (um) Equatorial axis (um) P/E ratio Sculpturing Species ean Range Mean Range Colpi ^ Columellae Upper Lower Figures Moluccella spinosa 29.5 29-31 30.4 29-31 0.97 0:9-].] 3 Su Sc 138, 139 Notochaete hamos 30.1 26-33 29.3 26-31 1.03 0.8-1 3 Ps Se 140, 141 ostegia fruticos 21.8 20-22 21 20-22 1.04 TOLI 3 Su Se 142, 143 Otostegia integrifolia 29.0 20-3 23.8 22-26 1.23 1.0-1.4 3 Su Sc/Mi 44 egia тїписсї 24.2 22-26 21-1 20-22 1.15 1.0-1.2 s Su Sc Otostegia repanda 20.9 20-22 17.2 15-18 1:22 1.1-1.4 3 5ч 5с 5 stegia tomentosa 21.1 18-22 18.3 18-20 1.16 1.0-1.3 3 Simple Su Sc 146, 147 Panzerina lanata 23.3 22-26 19.4 18-22 T2 LU-I.5 3 Su Sc 148-1 Paralamium i 16.3 15-20 15.0 15-18 1.10 0.9-1 3 Su 5с 151 Paraphlomis ja 30.1 29-33 32.8 29-35 0.92 0.8-1.0 3 Sr Sc 152, 153 Paraphlomis lanceolata 26.8 26-29 26.0 24-29 1.04 L0-12 B Su Sc/Mi 154, 155 Paraphlomis rugosa 0.0 26-37 31.2 29-37 0.96 0.8-1.1 3 Su Sc 6 Phlomidoschema parviflorum 24.2 22-26 21.6 18-26 1.14 1.0-1.4 3 Ps 5с 157,158 Phlomis agraria 9.5 29-31 25.7 24-29 1.15 1.0-1.3 3 Su Se 159, 160 Phlomis crinita 33.9 33-35 25.1 22-26 1.36 1.3-1.5 3 Su ? Phlomis herba-venti 34.1 31-37 28.4 26—31 1.20 1.1-1.4 3 ? f Phlomis lanata 40.3 31-44 29.0 26-33 1.39 1.2-1.6 3 Su 5с 161, 162 Phlomis maximoviczii 20.9 20-22 18.9 18-20 LTI 1.0-1.3 3 Gr 5с 163 Phlomis tuberosa 27.3 24-29 25.5 24-26 1.07 1:01:2 3 Su Sc 4 Phlomis umbrosa 19.6 18-22 16.1 15-18 1:29 1:1-1:3 3 Gr Sc/Mi 165, 166 Phyllostegia с: 42.5 40-44 45.8 44-48 0.93 0.8-1.0 3 Ps Re 167, 168 Phyllostegia hirsut 39.8 37-44 41.1 35-46 0.97 0.9-1.1 3 Su Sc 169, 170 Phyllostegia hi 33-37 36.3 35-37 0.96 0.9-1.1 3 Su Sc 17:172 Phyllostegia lantanoides 36.3 35-37 38.1 37-40 0.95 0.9-1.0 3 Su Sc Phyllostegia racemos 23.2 33-35 36.1 33-31 0.92 0.8-1.0 3 Su Sc 174 Physostegia longisepala 55.9 53-59 48.4 46-52 1.16 1.1-1.3 3 Su Sc 115.176 Physostegia pulchell 45.8 43-48 42.3 36-46 1.09 1.0-1.3 n: Su Sc sium m 35.9 33-40 30.6 26-44 1.19 0.9-1.4 3 Su Sc 177 әвәріоџошәјѕобоа pue eeepioiue] ш Абојоца;оу иәоа #661 ү JOQUNN ‘1g euinjoA ошиео $ qesy-nqv 669 UN TABLE 2. Continued. Polar axis (um) Equatorial axis (um) P/E ratio Sculpturing Species Range Range е Colpi Columellae Upper Lower Figures Pseuderemostachys sewertzowii 37.0 31-42 29.0 26-31 1.28 (1.0-1.5) 3 Su Se 78, 179 Roylea cinerea 22.2 20-24 22.0 20-24 1.02 (0.8-1.2) 5 Su Sc/Mi 180, 181 Sideritis canariensis 31.7 31-3 35.0 33-31 0.91 (0.8-1.0) 4 Ps Sc 82, 183 S кесе йі {з са око 44.0 42-46 40.3 35-44 1.10 (1.0-1.3) 4 Ps Sc Si hlorostegia 30.4 29-33 30.4 29-33 1.00 (—) 6 Ps Sc 191, 192 $ sAr tis е 26.6 24-29 22.2 20-24 1.20 EIFS) 4 Fs Sc 185 Sideritis euboea 91:7 29-33 31.7 29-33 1.00 (—) 6 Ps Sc 193 Sideritis gomerae 36.3 35-40 33.9 31-37 1.08 (0.9-1.2) 4 Ps Sc Sideritis hirsuta 31.9 31-33 29.7 26-33 1.08 (1.0-1.2) 4 Ps Se Sideri loleuca 85 29-33 91:5 29-33 1.00 (—) 6 Ps 5с 194 Sideritis hyssopifolia 27.5 24-31 25.5 22-29 L:09- .:(0.9-1:4) 4 Ps Sc 186 Sideritis ilicifolia 32.8 31-35 24.9 22-29 1:33 (1.2-1.5) 4 Ps Sc/Pu 187 ideritis in 26.2 24-29 24.6 24-26 1.06 (1:9-1:2) 4 Ps Sc Sideritis lagascana 31.2 29-33 24.0 22-26 1.31 (1.1-1.4) 4 Simple Ps 5с 188 Sideritis lanata 29.7 29-33 26-31 1.07 (0.9-1.6) 4 Ps Sc Sideritis libanotica 28.6 26-31 28.6 26-31 1.00 (—) 6 Ps Sc Sideritis marshchalliana 30.5 29-33 30.5 29-33 1.00 (—) 6 Ps Sc ritis montana 26.4 24-29 25.1 20-29 1.03 (0.9-1.3) 3 Ps Sc+Fo 195, 196 Sideritis pullulans 35.2 31-40 35.2 33-37 L00- 09-1) 6 Ps Sc eritis ro 26.0 24-29 21.3 20-24 1:29 (151.3) 4 Ps Sc 189 Sideritis villosa 30.7 29-33 28.4 26-31 1.07 (0.9-1.2) 4 Ps Sc 190 Stachyopsis oblongata 23.3 22-24 21.6 20-24 1:09. (0:9-]1.2) 3 Simple Su Sc 197-199 Stachys riddellii 3 Su Sc 200, 201 Stachys s t 28.2 (26-31) 24.4 (24-26) 1,19 (1.1-1.3) 3 Simple Su Sc 202-20 Stenogyne haliakal 39.6 (35-44) 36.5 (33-42) 1.09 (1:0-1.2) 3 Branched? Su Sc 205, 206 Stenog kamehamehae 45.8 (44-48) 41.4 (40-44) 1513 (1.0-1.2) 3 Su Sc 207, 208 Stenogyne purpurea 38. (33-44) р (28-35) 1:2 (1.0-1.4) 3 Simple Su Sc 209, 210 Sulaimania otostegioides 21.6 (20-22) 20.0 (19-22) 1.08 (0.9-1.1) 3 Su Sc 11 U@PIED jeo5iuejog HNOSSIN 099 әш Jo sjeuuy TABLE 2. Continued. Polar axis (um) Equatorial axis (um) P/E ratio Sculpturing Species Mean Range Range Mean Range Colpi Columellae Upper Lower Figures Suzukia luchuensis 2451 (24-29) 24.2 (22-26) Eiz (1.0-1.3) 3 Su 5с 212, 213 Synandra hispidula 40.3 (39-42) 30.8 (29-33) 1.31 (1.2-1. 3 Simple Su Sc 214, 215 Thuspeinanta persica 39.6 (33-42) 29.7 (24-35) 1.35 (1.0-1.6) 3 Su Sc 216,217 Wiedemannia orientalis 25.1 (24-29) 18.0 (15-20) 1.40 (1.2-1.7) 3 Su Sc 218, 219 ono ea oppositifolia 10:7 (08-22) 18.9 15-20 0.99 0951.3 3 Simple Su Sc 240, 241 Comanthosphace japonica 23.5 (22-26) 19.4 18-22 1.22 1.0-1.4 3 Su Sc 242-244 Comanthosphace stellipila 24.2 (20-26) 20.0 18-22 029 0.9-1.5 3 Su Se Comanthosph ublanceolata 22.0 (20-24) 18.5 15-22 1.20 1.0-1.4 3 Su Se Leucosceptrum canum 2 (24-29) 24.9 24-26 1.09 1.0-1.2 3 Simple Su 5с 245, 246 Pogostemon brachystachyus 26.0 (24-29) 291 22-24 1.12 LI-IS 3 Su Sc 227, 228 ogostemon cablin 26.0 G=} 24.2 22-26 1.09 1.0-1.2 3 Su Sc 229, 230 Pogostemon cruciatus 24.9 (24-26) 26.4 24-29 0.95 0.9-1.0 3 Su Mi 231 Pogostemon glabe 24.9 (20-29) 19.4 18-22 1.30 0.9-1.5 3 Su Mi 232, 233 Pogostem упеа. 21.9 (26-29) 20.2 18-22 1.35 1.2-1.6 3 Su Sc Pogostemon туоѕи 23.8 (20-24) 17.4 15-20 1:87 1.1-1.6 3 Su Mi 234, 235 Pogostemon plectranthoides 27.1 (24-29) 23.5 20-26 1.16 ®Ъ1-12 3 Su Mi 236, 237 Pogostemon samps 24.0 —) 22: ( a | (— З Su ? Pogostemon stellatus 18.7 (15-22) 16.5 (15-18) 1.15 (0.9-1.4) 3 Su ? Pogostemon yatabeanus 24.2 (22-26) 18.7 (15-22) 1.31 (1.0-1.7) b: Su Mi 238, 239 Rostrinucula dependens 23.1 (22-26) 21.8 (20-22) 1.06 (1.0-1.2) 3 Su Sc 247 Genera of uncertain affinities Anisomeles heynea 27.0 (24-29) 25.1 (22-29) 1.09 (0.9-1.3) 3 Su Mi Anisomeles indica 31.2 (29-35) 32.1 (29-35) 0.98 (0.8-1.2) 3 Su Mi 221,222 Anisomeles i 35.2 (33-40) 34.1 (31-37) 1.04 (0.9-1.3) 3 Su Mi 223 Anisomeles salviifolia 28.2 (26-31) 25.5 (24-26) 1.11 (1.0-1.3) 3 Su Mi 224, 225 Eurysolen gracilis 23.8 (22-26) 18.9 (18-20) 1.26 (1.1-1.5) 3 Simple Su 5с eeepiououije1soBog pue eeepioiue ш ABojoydiow чәцоа Ё661 y ләашпү ‘1g eunjoA ы оициео qesy-nqv 199 Annals of the 662 Missouri Botanical Garden rest of the grain scrobiculate); columellae simple reticulate sculpturing and simple columellae. All of (Nabli, 1976). the Pogostemonoideae, most Lamioideae, and the two genera of uncertain affinities exhibit these ple POGOSTEMONOIDEAE siomorphic conditions, but the wing derived r in some Lamioideae: tetracolpate and Pollen of subfamily Pogostemonoideae (Figs. ели ШЧ: д ме, г Medo lae, on 227-247) is oblate spheroidal to euprolate, with EE ulpt КЫ ing бый ilate, gran ulate Ш the pola —29 um and the equatorial axis tole; xod 15-29 y ains are tricolpate and inoper- t e exine is scrobiculate or microreticulate, a some species of Loca des 281, 2 Бине: б?) 1-331 n y small ones. The cameli are unbranched. DISCUSSION Pollen morphology provides no distinction milies Lamioideae 3 @ 5 @ 4 Ld D, RE @ @ ч Ф nodu are though nt to be derived, шн опа group PETER 1992a; see below), and can Ше. 1991) until they are tested for congruence with other charaich through a a pars оду P M Other and therefore contribute a to an assessment of phenetic d nshi Based o MEHR P ulariales, we iaie (Abu- follo comparison to the Scroph- Asab & Cantino, 1992) 5 у ge clade comprising the gynobasic-styled Labiatae (includ- all taxa under study here), some of the gener: pollen type in subfamilies Lamioideae and Pogoste- monoideae is tricolpate, inoperculate, with supra- NUMBER OF APERTURES ithin len (Figs. 191, bra was f Empedoclea. E has been RE eee th ally assigned to the Verbenaceae but e here as Labiatae, following Cantino et al., 1992): Car- yopteris nepalensis Moldenk в, Сего оа e Ridley, and Huxley. the study para hee pol- ps Sideritis sect. il hs fau trinum 1993). These p are mily Lami oclea pend test for congru snc em eee Although we examined о five species, ae (1972) Ma many others and found this feature to nt within the section and “a ent elsew here in feiss the colpi are е h lpal arran; MEC 182, 186, 189); the colpa "T 8 92), In sid. m- and tetracolpate grains we flower, thus their presence may not feature of the genus n COLUMELLAR STRUCTURE Simple columellae are plesiomorphic a and obasic-styled Labiatae (Abu-A S © = Volume 81, Number 4 1994 Abu-Asab & Cantino Pollen Morphology in Lamioideae and Pogostemonoideae tures are columellae or elements of the tectum. Further TEM ү, of these genera would help clarify the situatio "n is неол; е пане of ae бен іп nigh tenike Gomphostemma, and Steno- of tribe Prasieae sensu Wun- derlich (1967) (= subfamily Prasioideae sensu Bri- quet, 1895-1897), a group six genera distinguished by fleshy nutlets. The c ellar tructure s examined i r thre er r ium), and further study of Chel uld be mellae (if that is indeed worthwhile. Branched colu what they are) are derived within the Fs d and may therefore provide evidence for a rela- tionship between Chelonopsis and tribe Prae DERIVED FORMS OF SCULPTURING f the species studied have supratectal sually -Supratatieulate), cap .122- деа Ph) ll eo tectate cene we apply it here йо of the йе райда bis a serobiculate exine (borde ering on pun d in five di rest of surface is scrobiculate. the Lamioideae sa e suprareticulate sculpturing (plesiomorphic in in “the ami ritos ip ves т 1992 п- psi 6 tino, 1992a). Geis must be cautious erui inferring relationships based on this feature red = transformation of oy pee RE to psilate g has occurred several times e Labiatae (Wag. 992 i staff, 1992). An in pte MR ma represented in one ge of Lamioideae (Meta- SMachydium, 121) and one of Pogostemono- ideae (Rostrinucu 9ng been groupe morphology (Bentham, 1832-1836, 1876; Bri- quet, 1895-1897), but the cn uec: taxa are not thought to be related to The psilate pollen of | Hn Rn supports (at least on phenetic grounds) its segregation from МО 5, as is pointed оп ou tby Asini iei & Minore (1982), its segregation ‘from Stachys. In all such cases in which a genus is segregated on the ba putative apomorphy, its segregation is socii pending е posi of the larger complex it belon; . Ideally, it should be demonstrated that ed tea serene and the taxon it is segre- ted from have synapomorphies; otherwise re- eal of the Ee may create a paraphyletic group The irai of е здан тау be systematically si in Gomphos а Phiom amium, and mis Within the нц it was und only in maximoviczii and umbrosa (is 163, 166) ‘The other peces. of zm Ubera “ беро uen and фен six species ч Phlomis г. (Le., exclu udin wr говна Bent — La- e 9 (1999) ave suprareticulate ne n. The species with grains in th دم Ix]‏ g.‏ 5.6 ]=[ “y‏ > d belo eS, pote 1897). Other species in this section ve suprareticulate pollen (e.g., ria, Figs. = А21 & Moore, 19 hin Lamium, " к id ы йә albuma kien ( mort. sensu Mennema t the case п it where the distribution 9 sculptrin dnd by Briquet (1895-1897) and Wu s Li (1977). G. chi- he species with granulate sculptur nen. parviflorum) fall into голад ѕес- ions. At least three sculpturing types (Figs. 63, 65, 70) are PP within section Gompho- . 68, 74) within section Stenostoma Prain. e ulate nic арня was found in Gompho- a javanicum (Fig. 70) ond agopsis mar- piod (Fig. 77). Altl ribed here a as rugulate, the sculpturing i is ce different e. Могеоу culpturin, à to the пенны the be worth ethe е type occurs in the other three species of Lagopsis, 664 Annals of Missouri АНИ Garden as it may be a synapomorphy for the genus and subfamily Lamioideae are four members of subtribe supporting evidence van its кү from Mar- rubium rubium s. str. е panas polen (Figs. 122-128 3). sculpturing, found in КОРОТ javanica Fie. 153), is marginally Чаш from шашы it differs in that the Ж, £ 1 not e у ‚ so the A 1 s ] £ rugula a single grain; ideis the muri are granulate (more apparent in the micrographs of Azizian & Moore (1982)). wf oblon- goua (Blume): Prain has simile: sculpt uring but & M. 1982); the obs ive species of ат и stud- ied have suprareticulate sculpturing (Figs. 155, 156). VARIANTS OF SUPRARETICULATE SCULPTURING Within the general class of surpareticulate sculpearing, шее is a in en = ape an and the number, size, shape, ud arrangement of the perforations. It is difficult to assess the polarity of the ese ree due = = tion in the out- groups { T taxa nnb commen The similar sculpturing of Lamium galeobdolon Gotorn morphology nets its ан The Japanese endemic Aji ена Ajugoides (Kudo, 1929). Its suprareticulate е sculptur- 8 narpie Epling; t 19, Ld of Bas ii Ajugoides i is unlikely o be тм te to sree ria, flowe dup which i к: ени ке atl on с Quads) “ а NDA to Stachys should P investigat The only endemic North sel genera in y Melittidinae sensu Bentham (1 el i Mac bridea, PEAS and Synandra. Similar- ity in stomatal pt ire d 1990) sug- a, Physostegia, isd Macbridea ever di: is evidenc nce that өөп remaining genera of Melittidinae, M genera. data. Neit зав Cheoropsi (Figs. 31-34) nor Me- 9, 120) resembles any of the North idi ae exine sculptu mini gular m in Physostegia and some species of Brazoria.‘ vila e shalio ow lumina are bounded by рг rad- ually slop Synandra бе. ә 1 м eee in nm Ke Mac ostegia but sten Galop tfe. yet 50) Although a чое relationship betw not previo ously im suggested, they sha re a inctive form of suprareticulate sculpturing, = dites perforations that tend to radiate out from the cen е a. Moreover, Sy” tinct anther thecae; confi phic in the Lamioideae pied 1992a). GENERA OF UNCERTAIN AFFINITIES Two genera of uncertain subfamilial affini Anisomeles and fonds > were in ight study with the hope that pollen morphology might otypic ё sitions in the Labiatae s desc by hee 1898), who placed it in the Prasieae (Гат Wo Gomphostemma e (1940) were’ Li (1977) placed Eurysolen in the Ajug! Ach Keng (1969) noted e resemblance to ^ ospermum (Lamioideae) in general we "d cence, and floral structure. Press p^ e cluded the genus within the ogostemono! eye ointed out similarities with Pogostemon, p P nucula, and Comanthosphace. Unfortundt ately, Volume 81, Number 4 1994 u-Asab & Cant sap езеш EE in Lamioideae and Pogostemonoideae len coe din is of little help in resolving the affinities of Eurysolen. Its sculpturing (Fig. 226) ше specie ies om Ac hyrospermum (Fig. 4) 244), and Ajuga (Abu-Asa & Cantino, к» gned by Bentham she Vis ud pam (1895- 1897) to tribe Lami (corrected pesca EE р, Can- tino, 1984 h ut ical and с К traits у subf: d Laid (Wunderlich, 1967; Abu-Asab & Cantino, 1987; Cantino, 19 din) Cantino ‚ом b) hypothesized between Anisomeles and Po- hre apomorphies: pres- ence of subsessile leaf pbs ean gem nds with a e icellular cap, bearded stamen fila A lustrous pericarp. On the basis of this evidence, ntino е 1992) provisionally assign ni- — to ipsi Po EE EO вов is- large cc OR (F igs. 220-225), is ver to that of Pogostemon (Figs. 227-239), supporting the hypothesized relati m between these gen- era. Within t idea i pturing was , similar scul d in some species of i eih (Figs. 129- LITERATURE CITED ABU-ASaB, 990. Phylogenetic чаў ions of ollen Morphology in T mily Lami Ag pg axa. Ph.D. Disserta nk “Омой . D. 1987. Гирне impli- cations of leaf a anatomy in subt а, Meli devi hata) and em taxa. J. Arnol pesa эи 2 podre di of di Asai m and its systematic implication: Syst. Bot. 14: 359-369. m 92. Pollen morphology in sub- y Lamioideae (Labiatae) a its phylogenetic ee e Pp. 97-112 in R. M. Harley & T. Reynolds (edi гама о Labiate Science. Royal Botanic Garden: w. riaa 1098s. Porini implications of pollen morphology in tribe Ajugeae (Labiatae). 22. Bee. Bot. 18: Mimi Systematic implications morphology i in tribe Prostanthereae (Labia- me pollen tae). Syst. Bot. 18: 563-574 —— P. D. CA o, J. W. Now T. SANG 1993. Systematic implications of ا‎ morphology in Caryopteris Анн» ае). Syst. Bot. 18: 502 таня Any Ѕнду, Н. К. lants and bee э od: . (originally by J. C. Wilis evised by Airy Shaw). Cambr ridge Univ. Press, Cam brid dge, "aei AZIZIAN, D. & D. M. Moore. 1982. Morphological and кы ы їп denis D рше» Bunge and Par s Prain аах tae). Bot. Г ne 85: ae 2a ла. Ваѕѕєтт, I. J. & D. B. phology of the tiem jn (Labiata) i in n North Am with с ico, Central Ey South America and Eurasia. Pollen & Spores 28: 279 BENTHAM, C. 1832-1 686. Labi G ї Sp cies. Ridgway ы Sons, London. Labiatae. Pp. 27-603 in A. de Can dolle (ed ito or), Prodromus Systematis Naturalis Regni Vegetabilis, vol. 12. Treuttel & = Paris. 876. Labiatae. Pp. 1160- in G. Ben- bie & AE D. Hooker (editors), ^na Plantarum, Reeve, ped uoi, т 1895-1 th esent Pp. 183-375 ngler & К tl (editor sh Die Ntirichen aedis dn ^. Abt. За. W. Englema: war d Cantino, P. D. 1982. f th Phy sostegia (Labiatae). e tr. Сг 2 rb. 2 ——. 1990. The эы cin а i A mata d trichomes in the Labiatae and Verbenaceae. F rn Arbor. Т1: 32 23-370. 92a. Evidence for a polyphyletic ae on of ae Ann. Missouri Bot. Gard. 79: ——. 1992. buo a руке classification of the Labiatae. Pp. 27-37 in R. M. Harley & T. Reyn se Neri Десете Science. S Royal ага . WA а-н Сеп- and classifica 522 in vances in in ч к ‘Roy al Botanic Gardens Kew ER . W. SANDERS. 1986. Subfamilial classi- ونا‎ of Labiatae. sang Bot. 11: 163-185. . & M. M. Har Trichome, see mapping of the chlorop last DNA molecular phylogeny of = ижа. Ann. Missouri Bot. Gard. 79: 266-283. ER MN G. 1945. Pollen msi and andis tax- om = e, and Avi -285. ATON. ollen morphological studies in tril ibe Ocimeae рсн Labiatae): І. Ocimum L. Gra . K, N. Н. HoLMGREN & L. C. BARNETT. lex Herbari art I: The Herbaria of the Wor d, 8th ed. New B ical Garden, New . 1972. Pollen yo of Taiwan. National Taiwan Univ. Botany Department Press, Taipei. Ниүмн, K. L. LT Le pel $ 5 зумётанцше du genre Sideritis L. (Labiatae). B age . Nat (Paris), 3rd series, no. 45, Bot. Kenc, Н. 1969. Flora Malesianae A bead XLVIII. 666 Annals of the Missouri Botanical Garden A revision of qus n Labiatae. Gard. Bull. Straits Set T . 24: 13 Kupo. 1929. peer sino-japonicarum prodro- mus | Mem. Fac. Sci. Agr. Taihoku Imperial Univ. 2(2): 1- 1987. The Plant-book. Cambridge Univ. Press SS, Cambridge, England MENNEMA, J. 1989. A Taxonomic Bava sion of Lamium (Lamiaceae). Toldos Bot. Ser., vol. 11. E. J. Brill, eiden MUKERJEE, s. K. 1940. A revision of the Labiatae of the Indian Empire. Rec. Bot. Surv. India 14(1): 1- 228. Nasu, M. 1976. Étude MEAS VORNE ue de l'exine pa PA Put ie 499-525 in I. K. Fer uller (iors The Evolutionary РЫ дии of the sed Academ ` 1991. isi “ag and tests of ho- mology t ladisti Cladistics 7: 367- 394. PozuipAEV, A. E. 1989. Exine S ug rd yere n grains of the Vente family. Bot. Zur The On three new genera of glani pun the Kachin Hils. Sci. Mem. Medical Officers Army oo = jn y tribe Pogostemoneae. Bull. ves Mus. Il aon 10: 1-89. 1956. ve Pollenkórner der Labiaten. Willde- a 1: 617-6 Roca humans A. em eigo morfológicos iniciales del polen de тое L. en la Macaronesia. Bot. Macaronésica 6 SANDERS, R. ie d edd 1984. Nomenclature of the боса of the Lamiaceae. Taxon 33: 64- Bot. — ser 72 TRUDEL, М. D. С. & J. К. Morton. 1992. Pollen er and taxonomy in North American La- biatae. Canad. J. Bot. 70: mind 995. ntribución al conoci- is en la Р ум Ibérica P- -197 in 1 огїа árez Canea (editors), Actas del IV е de Palinologia (APLE, Barcelona, 1982). v. de Barcelona, des RÊS T. 1968. : ASA MA. P morphology of some e Indian pres J. Palynol. Е T- VEZEY, E. L V.P. SHAH & J. J. S 1092. numerical approach d beige uO lerminology. Pl. Syst. Evol. эзе 5-254. WAGSTAFF, S. J. хои bagel iba of poika ж. п tribe Mentheae (Labiatae). Pp. 113-124 in R. М. Harley & T Коке (d itors), RVG in Labiate Science. Royal Botani Gardens, p WALKER, J. W. . рог 975. The bases я angiosperm m ае Ann. Missour Bot. og 723. WATERMA! н red Pollen ае В, ae of the Liane e Michigan E Wu, & H. Flos Replica ian Aib val: sie Sisal Press Beijing WUNDERLICH, R. ü lichen Gliederung der Labiaten auf Grund der Pol- lenkórner, der Sëmenentwieklung u n des reifen Sa- mens. Osterr. Bot. 7. 114: 383-4 APPENDIX. Abbreviated collection data for be. specimens. Herbarium abbreviations follow Holmgren al. (1990). Achyrosperumum а gati Welw.— Malawi, Pum- phi District, Pawek 6606 (M Achyrospermum carvalhi ists Malawi, LaCroix 3058 (MO). Achyrospermum cryptanthum Baker — Tanzania, Frame 44 (MO). Achyrospermum densiflorum Blume— Thailand, ens west Province, Kanchanaburi District, van Beusekom al. 3739 (K). E parviflorum S. Moore— Uganda, Lov- eridge 217 (MO). aA schimper pate ex bis ) Perkins— mm Shoa Province, Ash 172 Acrotome angustifolia G. Ta diets Barotseland, To 0). Benth.—South Africa, Transvaal, Werdermann & E 1281 (A). Acrotome inflata Benth. — Botswana, Galloway 505 (MO) Acrotome fleckii (Gürke) Launert— Namibia, Seydel 2 ds А] got oides humilis (Miq.) Makino—Japan, Yama a-jio, Anonymous (US 350854); Nagasaki, Maxi mowicz 5.1. (BM). daa jon ege (Benth.) Ikonn.-Gal. — Afghanistan, , Aitchison LH (GH). Kur Valley yk HE — India, Mumbra, 2 рес. 1950, Santapau s.n. agra de indica (L.) K ühtze-— India, Hassa pee Saldanha & Gandhi 2170 (MO). China, Hainan, Ansonels malabarica (L.) R. i" he Sims— India, Has P j eis eles Erin i m ad 4 ecu Northern Ter- Bolly ек je — South Africa, Perry € Snij Вара 'androuzziana Pamp. — Libya, Kouf National B. ls hrs Benh Morocco, N of Tafraoute, "B. вайне int КОШ a — Cyprus, Paphos риш hio, Athens County, Ca Larsen КУ Ды, nigra L.— U.S.A., О 2 (BHO) Ya Cicilia; Larsen, 5996 (M Ballota HEE (L.) Be , Cantino 1329 (BH 0). Ballota еи (Biv. ) Vis. oi U.S.A, Ohio, vated), Can o 1310( BH d Lundell—U.S.A, Texas, Aramis S.A, Texas, Lem Wee Ohio, —U.S.A., Athens (cul S кезөө ГА Мы e a 8 3 3 Travis County, Sanders 76179 (TEX). BELA Brazoria truncata (Benth.) Engelm. & A. Gray Volume 81, Number 4 1994 Abu-Asab & Cantino 667 Pollen Morphology in Lamioideae and Pogostemonoideae Texas, Live Oak County, Sanders 76122 (TEX); Burnet ad Correll & Ogden | (GH). aesphacos ilicifolius S Ыр ДАП; Е Кһогазап, ees m (L.) Spenner — U.S.A., Kan- ounty, "McGregor 2 (US). я: lichiang th— China, Forrest 15429 (K). Che cre утре Diels—China, Yunnan, Chung- tien, Yu 1 (рте ота Smith—China, Yunnan, Rock 48 (A). Chi coccinea Wallich—China, Yunnan, Rock Yunnan, Coloni seguinii Vaniot— China, Yunnan, Ten 26 oore— Japan, To- a. s.n. (А). anthosphace japonica (Miq.) S. am qe zm Sep. 7 Ohashi ут Murata Japan, 1862, Anonym Coma anthorphace rte (Miq. 3S: egg er pn Province, Shidzuoka Pref., 30 Aug. 1958, cae n. (A). Hn pan, Makino n. (US ). Co manthosphace sublanceolata a S. Moore— Ja- pan, Hondo, Shiota 1 (GH Craniotome furcata ты —Nepal, Banerjo 2864 (US). Nepal, Riala, Polunin, Sie d & Williams 1297 (A). India, NW Himalayas, Gup US). Eremostachys fetissovii Regel a cia Golosko- kov 4487 (À). UB iliensis Regel— Kazakhstan, Goloskokov ostachys rotata Schrenk ex Fischer & C. Meyer— Каз. Aeon ek 4432 (A). jsp tachys sp ciosa Rupr.— Kirghizia, Armand 961 Eriophyton wallichianum Benth. —China, Tibet, Ludlow d Serif 8859 (GH). Eurysolen gracilis Prain—Indonesia, Java, van Steenis 11118 8 (СН). Fs bifida Boenn.— Denmark, Zealand, Svendsen Аал ladanum L.— Switzerland, 23 July 1953, Lo- hammar эл. (MO). Caleopsi s pubescens Besser—Switzerland, Evolene, Anonymous 3139 (MO). Galeopsis segetum m Haken S.A., Ohio, Athens (cul- a Cantino 1339 (BHO). ateopsis speciosa Miller — Finland, Nyl , Vanheck з моу“ Шег — Finland, Nylandia, Vanhecke шд tetrahit L.— Canada, Ontario, Algoma District, Go mple & B. rammal 2864 (МО). i mmphoena ch chinense Oliver — Thailand, Banang Sta., T zi" intermedium | Craib— Thailand, Kerr mma javanicum (Benth.) Benth. — Malaysia, "tán Shah 2772 (A). omphostemma leptodon Dunn— Indochina, Dec. 1926, nonymous 5002 (A). omphostemma lucidum Wallich ex Benth.— Thailand, Angka, Garrett 409 (K). 0 7 (GH "номад haplostachya (A. Gray) Н. St. John— A., Hawaii, Waimea, Rock 8350 (A). Been linearifolia (Drake) Sherff—U.S.A., Н. aii, Mauna Loa, Molokai, Feb. 1910, онч nd H). Hypogonphia turkestana Bunge— Afghanistan, o Salang Pass, Hewer 10254A (K). USSR, Turgaiskaia, Krasche innikov 5 5127 (A). agochilus aucheri Boiss. — Ігап, Kurdistan, Rechinger 42853 ч Lagochilus eri ie as (Pallas) Benth. — Kazakh- iin Pm kov 5788 (MO). agochilus rus hits & C. Meyer — Kazakhstan, Goloskokov 4292 (M agopsis а (Stephan) Ikonn.-Gal. — India, Koelz 6565 (U La аворзіз ѕиріпа (Stephan) Ikonn.-Gal.— China, Hebei, Beach 17 (K). amiophloni rotata (Benth.) Kudo— China, Tibet, Rock 14406 (GH). писи album L.—Iceland, Knappstadhir, Elseley 55/ 71 (M piana flexuosum Ten. — Italy, Sicilia, Larsen 35657 (MO). Lamium hee (L.) L.— Britain, S Wales, 11 May Pena Price s.n. (M m ga bn nicum id — Turkey, Malatya, Balls B2287 = S rk Ega District а moluccellifolium Fries— D 26 June 1966, си s.n. (МО). fium moschatum Miller — Cyprus, Liveras, Meikle 241 : be am m pu григеит L.— U.S.A, Ohio, Athens County, Santina 1273 (BHO). i De Wild. — Zaire, Katanga Province, за ра Те erritory, I сок big ў otis leonitis К. —Kenya, Machakos District, Verdcourt 2362 ( Gürke— Tanzania, Monduli District, sreenway & E 12500 “ee ne cardia —U.S Minnesota, Houstoi Co , Swanson 811 мо, EH Chiapas, Suus & 1 E 6966 ( onurus sibiricus L.— Hos, Nichols 2140 pea abyssinica yog Briq.—Somalia, Luu п Wieland 1142 ( pe барк Burundi, Muramwya, Teza, 10728 (MO). Leucas тын (Willd.) Link—China, Hainan, Janfen- ling, Chow 78309 (MO). biflora R. Br.—Sri Lanka, Yala Bungalow, Coor- y 69010407R (MO) sc serons Oliver — Burundi, Muyinga Province, Reekmans 697 Pes ). Leucas ы Be nth. — India, Hassan District, Rama- moorthy & Gandhi не (МО). Leucas eriostoma Hook. f.— India, Karnataka Chikma- galur, Saldanha EU (MO). Ган Аа Sprengel— India, Hassan District, Saldan- 17909 (MO). Leucas inflata Benth.— (K). Yemen, Bait al Lakida, Miller TN ucas javanica Benth. — Indonesia, Celebes, Meijer 10953 (MO). ucas lanata Benth.— India, Mussoorie, Dudgeon & [oed r 37 m dulifolia Smith— India, Hassan District, Rama moorthy HFPI aoe коео. India, Bembower 169 (МО). Annals Missouri ALAS Garden Leucas martinicensis R. Br.— India, Hassan District, Mysore, Saldanha 15337 (MO). Leucas mollissima Wallich ex Benth. — Japan, Kagoshi- ma в: ecture, Pyukyu, Ral et al. 104 (MO). Leuc yh ifolia Benth.— India, Kodaikanal, Bem i 1 (MO). pee Sides Wallich — India, Castle Rock, Almeida 57 (MO). ы vago China, Yunnan, Rock 6650 (A). ssi gor ra 96 (US). Loxocalyx оа sel kin) Makino—Japan, Yamato Province, Mur A). Loxocalyx КЛ с Hubei, Henry 6482 Macbridea alba Chapman— U.S.A, Florida, Bay County, pei 70884 (BHO); Apalachicola, Biltmore Herb. а (GH). Mache idea Venegas bonis Blake— U.S.A, North press Bru i os 6334 rcd Mari m aniso: des C. Koch Seybold 36896 (MO). Marrubium cuneatum Russell— Iraq, Kursi, Gillett 10852 (К). Marrubium heterodon (Benth.) Boiss. & Balansa—Tur- key, Adana Province, enr; oe (K). ia incanum Desr ., Ohio, Athens (cul- tivated), peur 1334 (BH Ma on p mL.— e Sierra de Cazorla, Town- sen 6 (K T pics melissophyllum L.—Greece, Mennega 174 achydium sagittatum (Regel) C. Y. Wu & Li— Kirghizia. Tien Shan Mountains, Medvedeva et al. 251 (A). Microtoena delavayi Prain— China, Yunnan, Forrest 15177 (K). Microtoena insuavis (Hance) Prain ex Briq. — China, boss Wang 80582 (A). rotoena robusta Hemsley— China, Hubei, Henry 648 824 (GH). pipera urticifolia Hemsley — China, Yunnan, McLar NT eh чу gat Mo Ше laevis L.— -, Ohio, Athens (cultivated), Cantino 1328 wp ы Moluccella spinosa L.—U.S.A., Ohio, Athens (cultivat- ed), Cantino 1316 (BHO). Notochaete hamosa Benth.—India, Assam, Bor 16154 (A). pti gia fruti Briq.—Saudi Arabia, Humbles 10008 Otostegia integrifolia Bentl Ethiopia, Amshoff 10381 e tegia minuccii Pic.-Serm.— Ethiopia, Addis Ababa, Pri 7472 (MO). Otostegia a Benth. — Ethiopia, Sidamo Province, Ash 1 885 (M Otostegia ene A. Rich.—Ethiopia, Debra Zebit, Boulos M 90 (M M 0) Panzerina lanata (L.) Soják — Mongolia, Altai, ОА E Prochorova 9059 (А). Russia, Krasnojar sk Province asa Mis Dunn— China, 10636 (K). 2 araphlomis ws (Blume) Prain ex Backer & Bakh. — Indonesia, Sumatra, Rahm “ Si Boeea 9772a (СН). RT beans Hand.-Mazz.— China, Hunan, Changning Hsien, Fan & Li 91 (GH). Yunnan, Шу Сосај rugosa Prain—China, Gwangdong, Merrill Phe ени parviflorum (Benth.) Уу Baluchistan, Kurram HMM Sly & pis. 94 v chipczinsky 802 (А). Phlomis crinita Сау.— Algeria, Oran Department, Samu- ae ti L.— France, Aude Department, 12 July 1891, [cse s.n. (GH). Phlomis lanata Willd. — U.S.A., Ohio, Athens (cultivat- m Putin 1365 (BHO). Phi ximoviczii Regel—China, Manchuria, Lit- ш , 15 50 ) (A). mis tuberosa L.— U.S.A, Ohio, Athens (cultivated), Canino шге (ВНО). Phlom е Turez.—China, Shandong Province, Chiao 735 (GH Phylonegia бшк r (Gaudich.) Benth. — aii, Oahu, Palolo Valley, 30 Nov. 1912, Rock Y Des GH). Phyllo ostegia pews Benth. — U.S.A, Hawaii, Oahu, Mt. lympus, Apr. 1918, Rock & Lyon s.n. (A). chen or hispida Hillebr. — U.S.A, Hawaii, Molokai, Rock 7 (A). Ph scite lantanoides Sherff — U.S.A, Hawaii, Oahu, денчи, А єл) Phyllos th. — U.S.A, Hawaii, Molokai, Roe 619 198 (A үк Physostegia longisepala Cantino—U.5 uisiana, qu atiis Parish, Gilmore & Smith 3501 y BH Oh Ph gia pulchell ., Texas, Brazoria ounty, Brown 9892 (BHO). ogostemon уа ھچ‎ ee — India, Khasi Hills, Hooker & Thom n. (G udi Pogostemon cablin (Blanco EEL — Puerta Rico, May guez, Horn 5424 (GH). 4 Pogostemon мамат афар т. Kuntze— Nepal, Stra Assam, rad & Winterbotto mone (GH ре оп d ber — China, Hainan, MeClure д п. (Canton быша lle 1 Herb. 454) (А). йи Pogostem us Bent ‚ Ohio, Athe ушей, Cantino e HO. Kuntze—India, Mal- abar Concan, Stocks, Law & Co. s.n. (GH). __ ч oides Desf.— U.S.A, Ohio, BH ii (H Pr mi en District, Lau 2549 ( untze— China, Kw angs, Kweilin, Wan & Chow 7917 Pogostemon yatabeanus (M si) Press—Japan, не. Но ndo, Shiota 4267 (GH oh Pra s 1. — U.S.A., Ohio, Athens (cultiva бош 1356 T Е Pseuderemostachys УЗУР (Негдег) ra 7 zakhstan, 12 May 1962, Karmyscheva 879982). Chi- Kan оаа лечит (Rehder) Kudo—China, W ‚ Wilson Dis Ro »ylea бисте P on) Baillon — India, Chamba paet 4T Apr. 920, Parker s.n. (A). —Spain, Canary Islands, Palm® oo 125 (A). CEP ROWE di en Aiton— Spain, Canary Islands, T eriffe, Burchard 134 (A). Volume 81, Number 4 1994 Abu-Asab & Cantino Pollen Morphology in Lamioideae and Pogostemonoideae Sideritis chlorostegia Juz.— Ukraine, Crimea, Juzep- czuk 3937 (A). Sideritis curvidens Stapf — Greece, Samuelsson & Zan- der 299 (GH). Sideritis euboea Heldr.— locality unknown, July 1927, urpus s.n. (А А е Noé ех Bolle—Spain, Canary Islands, 25 Ma eh Murray s.n. (K) 1 n, Aragon, Prov. de Huesca, —Spa 92, E Lager e (GH). dem 0 July 18 ideritis my & Heldr.—Turkey, Konya rovince, Davi. ap бзана t E Huesca Province, ideritis EY illd.— Spain, Aragon, Barbastro, 13 July 1892, St. Lager s.n. (GH). a e incana L.— Algeria, 16 July 1894, Cosson s.n. ideritis lagascana Willk.—Spain, Granada Province, everchon 1099 (A). Sideritis Pon L.— Turkey, Bithynia, Bile ik, Born- müller 1 9 (GH ). Sideritis tana L TAN S Shepard s.n. (GH). uns ritis marschallian a raine, Crimea, Juzep- sokoosirovshaja 9 (A). a L.—Algeria, cee TE 18 June 1934, ^y Siderits monta Faure s.n. (С a pullulans Vent.— Lebanon, Samuelsson 6150 Sideritis romana L.—France, 22 Apr. 1905, Raine s.n. GH Sider: nas villosa Cosson— Morocco, 20% В2463 (К). eS la ae E Pop d» Vved.— Af- ghan n, Badaksh Py Саг hg iddellii Hou ounty, ds hio, Yu aids 1229 (BHO); Meigs о, Forked 1 m State ark, Wis осна 64-91 (ВНО). — U.S.A., Ohio, Athens (cultivated), .A., Hawaii, Maui, US. A., Hawaii, Mo- nogyne ki lokai, Degener 5414 Stenogyne purpurea a Mann—U.S.A., Hawaii, Kauai, Degener et al. 1390 7 (K) Sulaimania Ik GF (Prain) ewe & Rech. f.— auc ipo, Mgr 15788 (K Suzukia luchu —Japan, Ryukyu Province, els Isla nds, Tae 3205 (K). Synandra hispidula (Michaux) Baillon — U.S.A, Ohio Morgan Сш. ‘son 1151(BHO); Шке County, Hammer 6 (BHO). Thuspeinanta persica (Boiss.) Вгід. — Afghanistan, Ait- chison 207 (GH). Wiedemannia аааз Fischer & С. Meyer—Turkey, azig Province, Davis & Hedge 28929 (A); Ankara, Kuntay 48 (MO). а е су ouri Bo ( | wt i j J | - WS d p | > er a s 5 NA í | " up o — 2 4 "t д = п L.] ач: 4 : С , n S Ug Tet v wu dt Ж йи» = 3 N af > "n К Р 4 й Volume 81, Number 4 Abu-Asab & Cantino 1994 673 Pollen Morphology in Lamioideae and Pogostemonoideae Ты E : Fict 6-61. Pollen of Lamioideae: — Haplostachys, ORARE and — 2 H Galeopsis. Mid 48. ANN X; pubesc cens. А ae Е 50. Galeopsis tetrahit. — 51, 52 25 T lostachys haplostachya. ingi ами coru n er 102 5A). a 59. Lago chilu s auc Чы; ri. 61. [ і р illus ^» 99, 61); 5 um (46, Б; Py peris 10 pen т 53) ит (47, 48, 49, 50, 52, 54, 56, ATA Zw M ДА, БЕУ Р. P." «ted Р ди ере А эу “eh. et Е 4 А DET еъ? ya Wi ci ү СЮ é ES , LJ L3 re n^ s «t е Sd > A. e е е. е eise ^n ^e B УТЕ * 254 Бө" 2; к PF FP eo aed ote fue, A ey, ~ Missouri Botanical Garden 676 ae » s Se | н AU, } з дәг. PA 31 8 &р NT uo о $255 P. Р A essa « © д. 7е ك‎ ecu 8 "d i: rtl IB # t oss. س‎ 555 "F сї ssa Ф í ean g Я | о BY, а REL ; y 4 88 | € ,r Rar XM E : Қ: & 1 FEET be # i E m. pr р: í А: zee x ЕУ 6 - i as = ama та. e $m mu ww 4 BS W co d NO > 9 3 Зе E d © i co SI DE. EY КӨР | hao w 3 ^ г 9 іс sao. pm ч ~ | 8 ma Be © Ez 4K S S rio 88355 ME gr sce go nu" 5 | БЕ * Sasso ESSE 7 — ON . ee Pod s we uo Y HUE ег 5 5 и © Sats o Маге 59125 Е "oW m ae ee d a À yN Wis Ё 8м a= с so Sesto BSN on mss sy D зї > a о i j : х х: ао а Я Q p 20 а © araphlo As 1 i 6 6 " я fen i D ч 6 8, 160, 10 n Volume 81, Number 4 Abu-Asab & Cantino 681 1994 Pollen Morphology in Lamioideae and Pogostemonoideae * bu AI, а-у 167 181. Pollen of Lamioid Phyllostegia, Physostegia, Prasium, Ps | tachy and Roylea. – Phyl 08. Phyllostegia grandiflora. — 169, 170. Phyllostegia hirsuta. — 171, 172. Ph yllostegia hispida.- 173. ostegia lantanoides. —174. Phyllostegia racemosa. —175, 176. Physostegia longisepala. — 177. Prasium ^ ME Scales = 1 um (168, 170, 172, majus 7f Dip. | 71 dm 17€ seuderemostachys sewertzowii. —180, 181. Roylea cinerea. Sc ales 09, 177, 179. 18 se 75 2.5 um (178, 180); 5 um (174, 176); 10 um (167, 169, 17h 1175). y‏ اا Annals eae ым Garden FIGURES 1 2 E Pollen of сае Sideritis. — 182, . Sideritis canariensis. — 184. Sideritis ^ Side 'rilis curvidens . Sideritis hy. Жазуы ыш. bo Sideritis ilicifolia. — 188. Sideritis 1 “190. Spe tis villosa. —191, 192. Зен chlorostegia. — 193. Sider a 95 ees ^g montana. Scales — m (183, 184, 188, 192, 1 195). ie Sauces 185, 186, Volume 81, Number 4 Abu-Asab & Cant 1994 Pollen Morphobgy 1r in Lamioideae and Pogostemonoidea E NEAN SEM FIGURES 197-211. Pollen of Lamioideae: Stachyopsis, Stachys, Stenogyne, and Sulaim mania. — 197- и. Stachys riddellii (Cantino 1229). — 202-204. = chys sylvatica. — 205, 20 207, 208. Stenogyne kamehamehae. —209, 210. Stenogyne purpurea. 211, Sar tegioides. Scales = ] um (198, 199, 201, 203, 204, 205, 206, 208, 209, 210, 211); 2.5 um (197 202); 5 um (207). 684 Annals of the Missouri Botanical Garden TE "ICHRES 919.99, r es ы D d FIGURES 212- 226. Pollen of Lamioideae (Suzukia, Synandra, Thuspeinanta, Wiedemannia) and genera of uncertain affinities (Anisomeles, Eurysolen).—212, 213. Suzukia luchuensis.—214, 215. Synandra hispidula i Wied 1y 48). —220. Ani- (Hammer 6) 216, 21 Th 1 TO, 217, изреїпата persica. —218, 219 1 1 is (K e: someles hey еап‹ )9 09095 i L ` 1 га етаппіа orientalis (Китау 4 n cu PN, py nb <<<. Anisomeles indica (Chun & Tso 43882).— 223. Anisomeles malabarica. сай 225. Anisomeles salviifolia. — 29€ 5 RG De Sa ET P р кР 917 ә 992. 223, i ene ү? . 6. Eurysolen gracilis. Scales = l um (213, 214, 215, 217, 219, 220, 222, 2 ) o um (218); 5 um (212. 216, 22] 224) THE PHYLOGENY AND CLASSIFICATION OF THE DISEAE (ORCHIDOIDEAE: ORCHIDACEAE)! Н. P. Linder? and H. Kurzweil?’ ABSTRACT cal data. The osi and p The data are Шо; cladistica, and the robustness of the various components s of the d b . Based on the cladistic analysis and the bootstrap analysis, a new classification is assessed by a bootstrap analys proposed for the Diseae. The vote of the bootstrap апа sho recognized. T lassification re cognizes five mon (a new pone, Hutton. е, а тау Бе E closely related to the Diseae than to the Orchide e lysis are agi to establish the "ed at 0 мн c inae, and Coryciinae. It is suggested ce of hybrid oan as it shares i ‘autapomorphies of the Disinae Lad Congas Huttonaea i is aim to be eae, and is conse new classification is formally presented, aid a key to the genera is of ed subtribal анма of the е Diseae (Orchidoideae) i is reviewed i in light of the available morphological, leaf nolog; most parsimonious tree is which formal taxa nae. » Disinae, Brownleein nae subtribes: the Satyr: h еа, the only t Тош nle: included as a subtribe of the Diseae. » The Diseae are a tribe of largely terrestrial Af- rican orchids, ud ue je Ap i em Disa uniflora ‘ange: plicated subtribe Cor e, and ‘the ае Huttonaea rs Schi ) Rolfe. Th available, none of ынын sa ит acter er support. While some of these iden oup- suggested D Linder (1986). A ott of ibit urzweil, bs indicates cepe a o the Bro пета (Lin- well i in prep.) Б ests t The first attempt to subdivide the Orc (tribe Ophrydeae of Lindley, 1830-1840) wa e entham in Bentham & Hooker (1883), who rec- ized four subtribes: Eu тото rapid Haben iud Diseae e, Ex сот former are identical to “the Coryciinae, Disp Disperidinae recognized 3 P ze lin (1899—1900), Schlechter (1898), 1915); Senghas (1973-- 1974), and Dressler (1981): Ther 9 b with li m bases end often cura ddl hel PES Corycium, EE dra, and Dis- peris in the subtribe. i бу ш over the се? cent di to this in the deli y classification, see Kurzweil et al., Th Bentham & uk (1883) have 1981), as well as th ранена Schizochilus, and кус) Етте а Hooker, Ыя named the gro Satyrieae. nadie (1899-1900) removed the up the "ats s the support of the Smuts "hip iis s Herbarium, NEE of Cape Т Afric Supported by the Foundation for Research e Pretoria, South Africa. The second author also ac- Fell wn, ы" 7700, South Africa. urrent urs Compton Herbarium, Na tional Botanical Institute, Kirstenbosch, P. Bag x 7, Claremont 7735, ANN. Missouri Bor. GARD. 81: 687-713. 1994. Annals of the Missouri Botanical Garden TABLE 1. A comparison of the tribal and subtribal classifications of the Diseae sensu lato. entham Hooker (1883) Diseae Corycieae рит (1889) Satyrieae Corycieae fe (1912-1913) Diseae Corycieae head (1927) Disaeinae Disperidinae Schlechter (1926) Satyrieae Diseae Disperideae Senghas (1973-1974) Satyrieae Diseae Satyriinae Disinae Disperidinae Dressler (1981) i d Satyriinae Disinae Coryciinae last three genera and placed them, in our opinion, in ШЫ соггесї groupings. The cba Honi f о the group wa followed by (Geblathter: 1 901; Rolfe, 1912- 191 3; Senghas, ham & morphological information. The recognition that there are two su QUAL } 1 шор within the Шы (ee ensu recognition of separate groups (Senghas, 1973- 1974; y dass The diei to combine the Satyriinae, Disinae, and Coryciinae into a single tribe, the Diseae, was by Dressler S) et has 90; beet enis siuide combination кошы these are sum- marized in Table 1. Huttonaea has h checkered history. Ben- eet & Hooker (1883) placed the genus between holina and Dude (1889) and Kraenzlin (1899-19 cluded the genus in the section 973-1974) and Dressler (1981) sepa- in rated Huttonaea into its own subtri f its appa jer. erect е 008 delimitation of the genera in the tribe has vary greatly. The original generic delimitations were established by Swartz (1800), and were further ela bereng on "de int ey (18905 1840). Subse- quent n any сес ted smaller and Morus genera, a а tendency that finally culminated in the large n r of small genera upheld by Rolfe (1912- 1913) in the Flora Ca- paraphyletic genera were retained, and so e of the segregated genera were poly рды groups based on flower ho and other striking features. In this w ic a letic. The generic limits in the тесеу assessed by pari el " (19918 = are presented there. The subtribe Di sents a problem. Line (1830- d sendy d he difficulty of esta ti m pre- 1966; Linder, 19 2r Linder & Kurzweil, 1990). Although the presently recognized seg gate genera are clearly monophyletic, their rela- ionshi the | е genus is no lear, g In this treatment t nera as establish der 86) sed, despite the clear evidence hat Disa is paraph ne monophyly of the genus (Kurzweil, unpub ку data), an заз еи we treat the two spec! separately he These шыраак = been heavily ме floral morphology. However, until гес Bor hor ral morphology in this је of lies s has orp. 8y Th "im of t ribe Di- Bolus detail scription of the flowers of a large number © Volume 81, Number 4 1994 Linder & Kurz Phylogeny and И айе of Diseae was that x Дил Bolus in his floristic no (Bolus, 1888, 1893-1896, 1911, 1913, 1918). Per E knowledge was contributed (1898, 1901) and Rolfe (1898, 1912- ew The on the floral significant engin were not dealt wit Th and, sequent tly In the past five years there has been substantial progress in the knowledge of the ee and weeny Ө of the Sons of the Dise Ku Leder and Kurzweil (1991) the Coryciinae, Kurse & Linde r (1991) some of the Disperis species, and Lin il (i a. I ources о: ructure tiner FC) шие the of the Disinae p оа . We have sxicdiive ridet data for the Satyriinae. There however, ral structures that have not di ioni adequately iig These include the chro- of the root E stem Tus The oem data suggest that a careful recon- RIS of the s suprageneric плана, on of the tribe Diseae is required. In this stud analysis of all available data is a sound чыйк д. ан classification for the group can be b METHODS Mor eee anatomical, and pone data were collected from vari ons, and in critical cases ne pic V University of Cape To ouchers are given in the figure captions. t were then vibe inco е scored for € genera, and a maximum parsimony analysis performed, usin, the 4 **ie*" routine in the Hennig86 (Farris, 1988). cladistic analysis software package. ed data were outgrouped to a set of genera be- nging to the Orchideae, which differ in the pos- pud of a spurred lip. In order to locate a iip ed tree, successive weighting wa р ied б 1969; Carpenter, 1988). The “indi Vidual character aaa perm for the three Bolus Herbarium fundamental trees were averaged and used to cal- culate a weighting for each character, on a scale procedure was repeat bilized. To test the support for the different t mono- phy le tic gro may be prop the data set was randomly sam- pled 100 times, and the stein, 1985; his was necessary to locate e ux теу i which is sion a 1 pen 1993). ponen ure strap ethods of Felsenstein (1985), a ы i Swofford (1993), it differs in that the component aot a selected issued are еш » rectly, poen byt a 50% majority rule consensus tre of sampled pid an Although t Ше nva do e from the set по pt oduce a limits for the cladogram puse 1991; Werdelin, 1989), th o give a ае indication of the support implicit in the nas used for each node. The methods used are rade by Linder (1991) and Sanderson (1989 The coding of клар taxa poses some prob- be due t th (such as in the Di into its components (s monophyletic taxa are variable, there are two ways of coding the variatio n. If the ancestral condition for the genus has bon e bya cladistic IT. $4 Lurzweil, in prep.) or in the Coryciinae sensu stricto (Kurzweil et al., 1991), hi condition i is E x terminal taxon, pecans of the terminal unit. Where “this detailed infor- oded as unknown (?) for the genus. Characters for which the evolutionary polarity could not be d ined n because they a by outgroup сотр se t small subset of the taxa, but for icri Uc 690 Annals of th Missouri а Garden f the Di Scale b. IGURES 1-4. Leaf арн сарз оп nd vascular bundles sclerenchyma caps s associated ul СА ae hes (Thunb.) Swartz, with the two epidermi: L = H. P. Linder). 1: L 3663, 2 data were available, were coded in exe where the 1 ) ев Pes pillans the ada xi ( (on the left side) and аһах ial sere a and ^" чачы ren e lay milar, and withou' РЕ. 26, 3. PC. 22, 4.1.4 —1. Disa oreophila Н. Bolus; note large eit and & Schinz; note bristle on eae margin and sm ii L. Bolus; n sophyll palisade. radii (PC = P. Chesse 81c) in which the tubers may be lost, and the these iin propagate. by vegetative "чол. a kal char ас te Р infe ed р тот фо one d I 1 h Pu Н. м. Wher ы од species are xe wn to be sse таша to species үр the character, this variation w у Stevens (1991), ignore MORPHOLOGY I. VEGETATIVE MORPHOLOGY There is ак Mass in the vegetative mor- © phology in the tribe. In Ceratandra the perennat- ing organs are d roots, while in the remaining genera they are testicular root-stem tuberoids. The s only exceptions are some species in Disa (Linder, ver. er dis there is variation in the vasc oo incomplete. e leaf anatomy of the Di leaf anato mies ( ж. is no consis T like Кан ра е e leaf anatomi es, but nera. n there are no E that combine gen "Some genera, EC Ficures 5-10. Habit and онан structure of various species in the tribe. Scale 5: 5 p cornuta (L.) Sw.; note robust many 6. Huttonaea Rchb. f. крг к= pn аура broa d. mi Brownleea macroceras Sond., inflorescence of 1- 2 flowers. —8. Cer atandra ee ively 9. Prerygodium leucanthum H. Во =й are inflorescence. эы, Satyrium candidum Lindl., robust, many-flow inflorescence. IT ces ( Kurzweil). 5: HK s $01 HK 1599, 7: HK 1576, 8: HK 1226, 9: HK botas 10: Jackson s. Volume 81, Number 4 Linder & Kurz 1994 Phylogeny and ads of Diseae 692 Annals of the Missouri Botanical Garden may n y in the case of the sclerified leaves of pie o but the relationship with Disa is not clear, and similar leaves occur in some sec- tions of Disa (Fig. 1). The occurrence of hyster- anthy may be related to the anatomy and texture of the leaves and floral Е = ds variation is also Ви to within the Disinae, and с be properly interpreted until i e eek this Ei is better understood (Linder & Kurz- weil, 1990). H. INFLORESCENCES The inflorescences are simple racemes, vary from many- to single-flowered (Figs. 5-10). wever, in most genera some y. eric level (see also the discussion on this character in Kurzw et al., 1991). many-leave y-flow plant (e.g., Fig. 5) is he plesiomorphic condition, and, as this occurs virtually all genera, this does not provid phy- genetic informati di Ill, SEPALS The involvement of the sepals in the attractive Pela a В E o J E ve о information. In the Disinae and Satyrium the lat- eral sepals are brightly colored, | And may be. the 15:20» 21), while i in the Coryciinae s sensu g - 1) they are green and generally insignificant [2% I6; D 18) The modification of the flowers is difficult to oe especially. т Сге petals, аге apiculate (Fig. 12). These apicules de- y earl н of mic Vogal ERD) аеры ү Bo unifacial tips the | remnants of the leaf hue How- re absent in the of the groups. ficult, as theoretica ally the tures, but outgroup comparison inet that they are synapomorphic for the Disi- 5 “The in. pie. are PARA pr ecialized li 11-22). In some о genera they contain import nt autapomorphic соми ог Vago dug: that delimit gr er очред Mo. the ge In оша spurs (Fig. 19), while in Corycium they are often papery, and may be partially hod (Fig. I7): rsal sepal is unspecialized (e.g., without peculiar odi ions) in the Satyriinae (Figs. 20- 22), Huttonaea (Figs. 13, 14), and the Corycunae sensu stricto (Fi 6-18), but differs in the о h these are usually very $ owers are green. sp ies (Linder, 19810, e; Linder & Кшт sepals differ in texture from ei pisi are a weil, t is th at the hen considered modifie were lost secondarily, and all the genera of In Disinae the sepals, and sometimes even the — Disinae are coded as having spurred dorsal se sepals, = Ficures 11- wers of various Diseae. Scale . Disa erubescens Rendle; n mc sepals and the Mean. linear lip. — 12. Н енше АШЫН ike Gawl. ex Spreng.) ee pe: dr broad, ovate ee ue E Ды galeate dorsal se uttonaea ei eke (Sch. ) Rolfe, with a fimbria ms and petals. — ea pulchra Harv., ith е fimbriate реїа . Brownleea macrocert iti with the large eal d. and the petals adnate t to the dorsal se sepal to = the A —16. Са viae atri Durand & Schin: ber he 17. Coryci бк a the large galea constructed largely of the petals. —18. Pierygodium "inversum ( Thunb.) еѕир! pinate. 1 note Sw. with a lar, е ga alea constructed largely fi 19. зр paludosa Har ing t the en ae to the sepals als. sepals fo ae the |, lan алы galeate To e the мема margin гес (HK = Н.К "b Plato dl TL 12: HK : 1069, 1 4, Eid и НК 140. e ee 19: Darling Wild Flower Annals of the Missouri Botanical Garden pecially in Disperis. In following the guidelines ыле ы en d (1991). However, the situation is more complex ome petis ithe sia: are age lorate to linear Disperis, where ap a minority of v species se ое, with the dorsal sepal forming the bulk dorsal sepal spurs. As there is as yet no general of the galea (Fig. 19), while in oth cladogram ae for the genus, it ca fe hether the presence or absence of spurs only fo ing the keel of the galea, while the petals is SERRA nor ca e frequency of occurrence for bos b h u Iu ч arg t beu case, bec t inae and Brownleea fe base the in 0 In the Disin clear » whether these spurs were lost ошаш ог jak d is s fused t o the gynostemium. This or in Disa sect. "eec f Kurzweil gai L 1 f. 1 E oF 1 £ зау eae Res I 8 this Sdn ih the ев of the Coryciinae, the (1990) h b 1 Schlechter’s (1901) op y ga coves Бы althoug that 1} l is derived, at least partially а тау Ь d f from staminodes. The presence of various lobes on with the petals, it is usually the smallest of the the d cm is useful at specific, ees and pos organs involved. It may be that once a ply siblyi е cases generic level, but its presence concave or galeate dorsal sepal evolves, the step is rather making it of little toward a saccate or spurred structure is relatively use for suprageneric analysis. Basal anterior lobes th ll sed to define the sections in the large small, and this may account for the parallel evo- have been u lution of dorsal sepal spurs in the Disinae and in genus Disa d ke 1981c), and pr bilobed Disperis. Comparison with the cladogram would petals occur in the majority of the species of A, also indicate a parallel evolution in Brownleea, in sect. ide (Linder, 1981f; Linder &K which the dorsal sepal is a slender or, rgan. This an exception to the above theory. It is evident that e has very. peculiar pou “Ман аге spurred dorsal sepals evolved several times in the — clawe у iseae, and as this character has generally been (Figs. 13 The apices of the fimbriae appear ter 14). stressed in Djs subtribal classifications, it may have to be кзы аг (Kurzweil, 1989 ib о the persistent misclassification of Biologically, the petals in the Disinae PE f spur, in the E у form the galea, IV. PETALS in the Satyriinae they form part of the landing t- pham det role of the fimbriate el in pesi The petals are very variable in the tribe (Figs. i p 11-22), and have been found useful at the infra- offer a SAAN for the pollinat generic level in the Disinae (Linder, 1981а-е; Lin- requently зине: with this formation of the der & Kurzweil, 1990). In the Coryciinae and petals plus de al sepal galea is the presence of Brownleea the petals are adnate postgenitally inia arily strongly cine petal nerves. ТЕ fused) to the dorsal sepal, and the P structures degree of кше ment of such “petal nerves" is combine to form the е galea (Fi igs. 15-19). This is variable, an they are missing in man ; also found to a lesser ases in ш Orchideae, However, although such “petal | nerves” have been d in Cynorkis. riation recorded in various genera of other subfamilies d n the degree of E of the me and the the orchids, they have a restricted distribu tion in — FIGURE 23. Variation in the pie gynostemium «пены. іп the Diseae, sh ly the gynostemium and lip. ом are indicated «4 м аем, the Ma e black; a endi a pbi: a, 1 = e la — lip appe rape :B, DEG „А, С, Е: a B. Раса bodkinii H. Bolus TA Gynostemium and lip; "Pf the long column-part а кай! three ‘lobed lip.— howing the reflexed “С = oie lobes and the auricles at the apex of ie faa M lobes. C, D. н erectum Swartz. ynostemium and a half-section of the galeate lip, sh one of the two spurs.— D. Details o: (ш s Же pee note the laplike stigma.—E. Disa tripetaloides (L.f.) N.E. Br., Haar шз th the > " showing the vertical scar where the petal was attached to the gynostemium erygodium oculum Vd : and the showing the massive erect lip append X pendage, gynostemium with the anth id 1 тед, to the sides of the lip.—G. еви recurvata Sond., showing the be le мы E the base of the fal * narrow erect limb in front of the stigma, and the large P тіке auricles. Sources. A, B, Burger s.n- ш 031. E, Linder 1670. Е. Tiaia 1589. G, Linder ey‏ س سسس سے Volume 81, Number 4 1994 nder & Kurz icon and ius of Diseae Annals of the Missouri Botanical Garden the ре апа my ео һе E de- in A whieh occur. The d pos the char- ac mra ont re have h пе Е ч Se E uds to four losses. The ы аны у has oe been as = the distribution of the char acter in the outgroup. Y. HF The lip is the most complex perianth member roups. ually ma rgans, a of an es. This hypothesis is use о ерм of sup- 5 in is in that it ected shows ver һа single DE Bid (Fi : 23 The ori e lip is variable. In the Orchideae and in some groups of the Diseae the Т ? , in the : same direction à as E axi ot the Шо This apes on the: lips, where the spur is parallel to the ovary, and essentially in li lip blade. In the Satyriinae the lip is also orthotropous (Fig 23C). In the Coryciinae and eea the lip is E it is deny fused te the кин (Fig. С) Schizodiu joa he lip is reflexed at the base, во that it i is held at n the Disinae the lipi is a simple teste: hout usually without lobing, and not fused to any ғ other шш (Fig. 11). It is usually a linear to lorate organ (e.g., Disa u uniflora), al ачаа arely ovate (Disa ferruginea b.) Sw., Herschelianthe gon inifolia (Ker Сані. ex Spreng.) Rauschert, Fig. 12), variously lacerated (Herschelianthe barbata (L.f.) N.C. A . These spurs are obsolete or highly reduced a few species, but these are almost certainly secondary losses, as the species concerned a clearly related to species with bi-spurred lips. i Pachites the lip simple, as in the Disinae, while in Huttonaea it is broad and fimbriate. In in shape, ranging from linens to triangular to th be d dium-sized to large. In Evotella and Ceratandra the lip blade is Pepe spathulate and anchor-shap 91). The base of the lipi is "E penne -podian as Hes pr. ree us A3 hie s might 1 2 nd 2) 4 188 2 However, ontogenetic evidence не. in cle ашу suggested that the lip is eil 1 pa various {шуш Тһе on lip appe ket of the e initiated late in the atop Apparent С erely an ap age of the growth of de lip as also Pee in numerous pi orchids. This by vei is pa ei primitively bilobed ( 19 zweil & Linder, the bilobed origin, and in massive же over the anther and rostellum (Kurz weil et al., 1991). The ebur. of the lip in the er biology is, like th E ونا‎ мө" ш Sat and Sa- tyridium it for e hood a ee ды! bearing the nectar; i e м аё Es art of ч айна platform together ux He lateral sepals, s the oil-bearing 1989). oryciinae it usually be: аи for the oil-gathering die Steiner VI. GYNOSTEMIUM of nther is basally reflexed in this eom the erect anther is derived from a de and this was supported by their cladistic bat In Brownleea the anther is basally гоб - the apical portion is erect, giving the imp Volume 81, Number 4 1994 Linder & Kurz Phylogeny es с Я of Diseae 697 of an erect anther. The situation in Huttonaea is not clear. The anther appears to be erect, but there are some indications that it has a slight tendency to being reflexe The anther- cells are кеш parallel to each that the anther in these pacta diverge toward their bases, imal to each other. In = енн the anther- arallel to each other, but they th ium (Fig. 23F) other. e Orchideae and Dise eel connecting the эш with the eae um is а present in Brownleea ү ig. ста However, clear ontogenetic evidence was not foun ( 1990; Lin c & Kurz sus. in prep.). Huttonaea species have prominent lateral gynostemium appendages, "a appear to be derive d from “auricles aly. In C connective process, E species where the anther- thee e medi ian part of the con- е rocess, аин Еа їп ап a anther, is found in some Orchideae (e pecially у Serapias) and is considered the ancestral state for the Orchideae and Diseae by Dressler (1990 all species have prominent lateral gy- ost appendages (Fig. 23). The situation wi regard to their derivation from “ E (staminodes) and “auricles” (filament шнен) is less с] th PRE May Eee (Kk °1 as their reis are very obscure in ri ontogeny of most Diseae. However, both stru exist in the tribe Diae and are prominen certain speci In Disa ты Lindl. the basal bulges аге sig- nificant during early r and later develop into a ridge on the k сше op of the connecting keel and аге fused to "s y are very variable ing from small and sculptured to and киш A flat. a e flowers of the Satyriinae have compar- y E lateral gynostemium appendages nex nt чанга ut signi K ordia were not observed in the early — um eil, in prep.). In some species the bilobed mtn pem in the mature Дайе. бх is ald genus, it c of the bin dd that the two- see t a m appendages indicates their o7 £ e - t A ers, and ipit the staminodes are vespa rated into the gynostemium tissue below the уине (Кш е 1991), while the central thick. HEE Many Disperis species have varie ateral dt A appendages (Manning & Lin- dor 1992; ачыш & Landes: 1991), im a кш ке horizontal potions 4 on the side of the mium. It is possible that the ioe erect portion is a staminodial structure, and that the sculptured horizontal portion represents the auri- cle. The primitive — of the Diseae and Or- chideae dip to e presence of prominent staminodes that are и ed to the gynostemium. ince o dn ie probably evolved from ances- tors with three abaxial od bas у, fused pun, n the p be regarded as an ойаны feature, and this ease. the nae can be н rded а Би state, hee the for- mation p e conne ж is эб гыны ран сег- tainly derived. Similar co found in the Brackycoryths er group » (Orchid - hag pun n Disperis Жолай Нагуеу апа Ceratandra а. In the remainder of the Coryciinae, in Huttonaea and possibly in the Satyriinae, the stam- inodes are strongly reduced, which is clearly de- rived. Previously the occurrence of auricles — nt appendage: пи was regarded аз a unique character of the пеки еап = Diseae. However, им stru uses han iam SEEN of the “Diurideae” (Dressler, qai although the rate logy of these structure: ricles 0! ее е Orchideae/ uei uk more а Auri 698 Annals of the Missouri Botanical Garden TABLE 2. Variation in the structure of the rostellum in the Diseae. Number of lobes Taxon mature primitive Central lobe Lateral lobes Viscidium Orchideae 3 3 infrathecal tooth square to elongate 2: lateral lobes Disa 3 3 infrathecal tooth square 2: lateral lobes Herschelianthe 3 aj infrathecal tooth square 2: lateral lobes Schizodium 3 ài infrathecal tooth uare 2: lateral lobes Monadenia 1 1 massive NA 1: central lobe Brownleea 2* 223 lost or reduced* elongate, erect 2: lateral lobes Huttonaea 3 3 massive inute 2: lateral lobes Ceratandra 2 2 lost large, flat, each half cov- 2: lateral lobes ering one theca Pterygodium 3 1? small elongate, d 2: lateral lobes Corycium 3 2-3 small moderately elongate, 2: lateral lobes strongly twisted Disperis 3 P flat and covering elongate, projecting 2: lateral lobes both thecae d 1(-3) 1(-3) variable obscure 2: lateral lobes Satyridiu 1 1 massive NA 1: central lobe Pachites амай 2 ? lost massive 2: lateral lobes Pachites adpressa З ? large U-shaped tape minute 2: lateral lobes * — exceptions occur; ? — unknown; ( ) = rarely occurring; NA = not applicable. are pei "i basic condition of the seam’ — ae, d their reduc tion dn t the Со oryciin ably derive Mas оа which is developed from the me- carpel apex, as is typical of the orchids, is heal пане lobed. Héweser, the lobes: are v n the Disinae the kati lobe is folded between e Ke cells and varies from relatively small to = prominent t. This i is essenti пову" similar t 9 the pork je the basal condition for the eros denia is unique in the Disinae by its massive i rostellum lobe, with minute or ae lateral lobes. The situation in the Satyriinae is less clear. Gen- me the rostellum is less clearly three-lobed than е Disinae and the Orchideae. hee weakly three-lobed or emargi atyrium, there is a wide range of different sizes and shapes of the central rostellum lobe (S erhayes, 1968; Wil- iamson 7), ranging from massive and com- Кы. sa to fingerlike or to short kno The lat | parts bear the viscidia in lateral or subt etailed comparative anal- ysis of the direito: бйнм structures may con- xe Arabi Misit for a phylogeny of the atyriu уф гезеп! lobed (— the "gu ie and bears ars а i terminal vis adult flowers. This is an m teresting eg o Mona cies of the р ing the е aped, straplik central гонаш lobe between as "Pa ane ia inii H. Bol Mesas de sio ipee pe an obsolete — re rge n of a central ros- tellum ia was consequently ien qe such (Bolus, 1893-1896; "Rolfe, oie ; an ion Schelpe, P but has “л d thus pro shown ollen massul. jated ‚ and the main portion is derived байа lobe, while Pe Sah lobes are minu Volume 81, Number 4 1994 nder & Kurzwi 699 menia and НИЕ of Оіѕеае Coryciinae also show evidence of a successive ге- duction of the rostellum three- Кош g. In Cer وا‎ is then developed as a narrow e een them. While the main portion of the strap is derived from lateral rostellum 10 Бев, cially this is similar to the stigma in Satyrium. However, in Satyrium the carpel apices form tw е the fused lateral Pepe apices, ae ше Loser basally. receptive. In Satyridium “and Pachites bodkinii, the stigma is a small, slig is a deep cavity encircled by the U-shaped central rostellum lo! nd is developed on the back side "à the isdem арт n facing the median sepal). 3 Case ыл k а to a central rostellum lobe. mtogeneticaly, the rostellum terygodium and C from a shallowly three-lobed, two-lobed, or unlo median carpel apex. The ro isperis is very different. The lateral rostellum lobes are de- veloped as elongate projecting arms. The central lobe is remarkable in that it is s and flat and covers the er. It was shown i one species that i two-lo a togenetic Коне is available from other Dis- peris speci The E. is some remarkable peculiarities in this pipe. ne Disinae and Bet йе stiomat called pe seudoterminal by Bentham & Hooker (1883), and Pfitzer (1889) also commented on the often terminal дуна of the stigma on the gy- nostemium in the ae and Satyriinae. The авав онар ies clade e development from the probably primitive ntire stigma to two pas stigmas on the ros- 7 E In Hutton on and "Же de sta from a ee structure (Kurzweil ee m most eirmod species, Шш the middle by the ще up: It was apices, which is the солоп situation in the Or- chidaceae (Fig. 23D, E). The median Brest 1 apex contributes from берудеп half 1 mewhat less к опе- Шш of the total mel pee but Є ЖЕ: Ee. in the Coryciinae, as well as in Brown- leea (Fig. 23G) and Huttonaea, the stigma is made pe ntir the median carpel apex, and th lateral carpel apices remain as small, sterile, ves- tigial rgans (Kurzweil, 1989, 1991; Kurzweil Linder, in р is derivation of the stigma from эш Р чоры иез 1982). It Mm de noted that two sp tstrap test shows that the alter- native solution also gets significant support In the Disinae the Mone is distinctly Sabie on a platform in front of di ГЛ Б Е ^ th rim along the argin, so that it faces backwards. Superfi- 1 |. to each other (iis der & кы їп рге ү a some Disperis species, the oo of the ture stigma is very 5 o that of Жаш (Kurzweil & Linder, 1991). In the remainder of Jai іЯ J r О аге — hes ug гусііпае sensu stricto, with their wide , have two separate stigmas Masse 5 зе дайда (Kurzweil, 1991; Kurz- weil et al., 1991). VII. POLLEN SURFACE The surface topology and wall anatomy of th ma cove (Figs. 24—32) provide a ts etal of data; weg ch appear ш. mre eet Ботове. gh, 198 Bore: санай & Funk, 1980) ps palynology о s the group has been studied chil & Pfeiffer (1977), Lin s (1986), ied eig Pen gated in "gena (Lindley, 1830-1840; Burns- Funk, 1986). The eee are combined = mas- sulae, and the m mass кеа аге е easily separated, and the solid body. ium. The pollinium does not form The massulae vary зана іп shape. Some 700 Annals ies а Garden of this variation may be related to the shape of the flower and the les, but е appears to be phylogenetically interesting. Chesselet & Lind (1993) suggest that the massulae are rounded most of the Disinae, while they are fasciculate in he Coryciinae sensu stricto is d massulae were also observed in the fe tgroup taxa stud- ied, and may be the ge aia dide in the Or- boss he organization en the t e. This variation is largely correlated with ame The кпап аге 1 enerally p г and ге ane constant within mane les ^v generad; the appears to have exceptions. In no pollinia studied was the tetrad Organization tetliy constant, but the frequency ied. There are exceptions to the simple pattern described above. Cor е ads Var- cies appear to hav imit p as been coded for tetrahedral tetrads and nonfasciculate pollinia ther problem with the character is that tetrad organization has n en s absent, as is ep of the sub family (В st Balogh 1983). The chidoideae are primitively нң Chesselet & Todes (1993) propose "eu the s , with a a ТЕШ with a 15 s capensis reticulat Ore 1503, 30: HK 1424, 31: HK 1508, 32: HK 1 tectate condition is primitive in the Diseae. This is чане by the analyses here. n-surface ornamentation provides a wealth sensu stricto, and the ornamentation consists of striations on the tectum. Th ind of the Й е semi-tectate walls di are either sinae generally have a rug namentation (Fig. 27). The situation in Satyrium n confused, with г an initi. tial wider mitt re- to ha Йа te ornamentation (Fig. 32). Halll has also rnamentatio been recorded in two spe- cies of Cor racomontana ani ni- grescens) he rest of the Coryciinae have namentation is rare outside the Disinae, its occur- ored when the genera were coded for e “cladistic ren a УШ. SEEDS The seeds are very uniform in the e Diseae (Figs. 33-44, Kurzweil et al., 1991; Kurzweil, 1993b), mi and app tly do carry much taxono cally eful information. The s of the Diseae arê almost always minute an same structur! s in the related tribe Orchideae (Healey et al. 1980; Barthlott, 1976; Ziegler. 1981; Tohda, 1983; Wildhaber, 1972; unpublished data). Th seeds are usually fusiform in gener 1 shape as in most other orchids, although dps and glo- bose seeds occur occasionally (Fig. j па! е hn consists of Seka tells y e^ outer amt inner ne. 1. 27-32. Е orn й i € pcm б). Sw.; note the fasciculate massulae ctum. ose dispen —32. S. striatum Sources (ИК =H. Kurewelly AFTER 1253, 2s «НК 949, 26: HK 1508, 27: HK 1253, 28: HK 1543, 29: — 27-32. 24- -26, 5 La re ‚ vih 0. Satyrium Pee pede айг Ер Volume 81, Number 4 1994 Linder & Kurzweil 703 Phylogeny and Classification of Diseae Character list for hê Diseae. as Hin spices Cobain TABLE 3. characters pee — all same size en grading from large t the ba: small at t sn |7 2; fn ideis m vue 3. Sepals ofte picul 0), dixe 4. Dorsal Ee simil жей to the мй pes ud dorsal sepals galeate (1). 5. Dorsal ea Mec (0), gore (1). 6. Petals similar to th `+» 7. Petals fused to gynostemium (0), free 8. Petals te to dorsal sepal (0), f x 9. Strongly developed pet hen nerves bog (0), petal nerves frequently pr in M T0 о base 5 mium (0), fi ba ini 11. Lip with a voilier hilobed Bien! vx not (1). 12. Lip with two spurs (0), unspurred (1), with one spur (2). 13. Lip galeate (0), not (1). 14. Lip erect at the base (0), patent or descending at the base (1). 15. Lip appendage bilobed (0), single and fused (1). 16. Lip blade simple (0), spathulate and anchor-shaped (1). 17. Column-part well developed (0), not and very short (1). 18. B teral ONDE appendages well developed (0), mall to abse 19, oh erect (Я тейеей an 20. Anther cells adjacent (0), w parated by a wid grum "UO x 21. Pol d | | (0), ош PIN". ssul ДЕУ 23. Pollen surfac sculpturing reticulate (0), bae ог Tn ). е semi-tectate (0), secondarily tectate 25. Rostellum three-lobed (0), only the central lobe de- iym, tooth (0), re and coverin king up inn whole rostellum (3), cq between small Men possim en Ө 21. Rostellum lateral lob minute oes Siri 28. elm "eh elongated 5 Weine late ral lobes which je і (0), projecting (2), twisted (2). 29. Sigma sessile on the Pak ua (0), subterminal Р 26. ай c central lobe an 30. im produced from all three carpels (0), median 31. mAN h сафа stigmatic part from one primordium 0), two primordia (1). ticlina] w ‚ Corycium vandig pee dl.) Rolfe тавал еї БЫ 1991), ^ Lyrium retusum Lindl. (Kurzweil, 1993b). е surface of the periclinal cells is variously smooth to more or less wavy; however, the prominent ginal due of the peri- cells have occasionally small shallow con- колы 3432 35: i ser "vs in бе ear rer = ER intervals as ар as their thickness d rad are too variable to give any phylogenetic eos À rem PME different seed type was found in Disa раина s (L.f.) N.E. Br., D. uniflora, D. „йлн alis тше r, and D. sess Lindl., which larg ег face cells with prs convex periclinal walls (Fig. 37, Kurzweil, 1993b). CLADISTIC ANALYSIS he characters that were ‚ found to be cladisti- sho Diseae (Fig. 45A » В), w i the n tree is a 1 1 and the consensus tree shows a basal polychotomy for the € ig. 45C). din successive weighting had o be run twice before the cladogram stabilized. Tw wo trees were located, cu iffe aa nly in y arrangement of the three outgrou . This the basic topology presented in cin 46 and "m whi ie the subs теш cds is — . The supporting 6 given in Figure 46, and the node numbers and аы. ар койы эбе are indicated in Figure 4 SYSTEMATIC INTERPRETATION DISEAE Dressler’ s (1981) eat of a single tribe, the and this 6 SRP is E at nı рог теа by a RSE 19) whic i reverses cn one group (Disa his an t a rigorous te: nalysis does not pre: cues of in oky for the c as only 704 Annals of th Missouri Poss ERR Garden TABLE 4. Distrib f ch while Mi f D t. Micranthae. Missing or variable characters, or logically inapplicable characters “g are coded as rachycorythis 11100 10111 12100 01000 00000 00700 0 a aiie | 11100 11101 12100 01100 00000 00700 0 Schizochilus 11100 11101 12100 01000 00000 00700 0 Satyrium 10100 01171 10000 00710 0000 71710 0 Pachites bodkinii 10100 01111 11100 00010 00000 10710 0 achites adpressa 10100 01111 11100 00710 00000 41710 0 Satyridium 10100 01111 10000 00110 00001 37710 0 Disa 10011 10101 11110 01010 00100 00710 0 Herschelianthe 10011 10101 11110 01010 00100 00710 0 Micranth 10011 10101 11110 01000 00100 00710 0 Monadeni 10011 10101 11110 01010 00101 31710 0 Schizodium 10011 10101 11100 01010 00100 00710 0 Brownleea 10111 10010 11100 01010 00000 10701 1 Huttonae 11100 11101 11110 01010 10000 41701 0 Disperis ITIT? 11010 01100 01010 00000 22101 1 огусі 11100 11010 01101 01111 11210 12201 1 Ceratandra 01100 11010 01100 IUH 11210 12001 1 Evotell 01100 11010 01100 TELEF} 11210 12001 1 Pterygodium 11100 11010 01101 01111 11210 12201 1 three genera of the Orchideae were the er (1883) and Pfitzer (1889), and e sed in mtg group. vr dt f these genera, Brachyeorythis, йеп f the Diseae, and this probably accounts for de relatively low “bootstrap percentile” for thi node. It d be есу, to ЫЗЫ, ze the whole шу Yý monophyly of the Diea such a global analysis is long overd des are recognized in the Diseae. Ese first ens (node 10) includes the e Satyriinae the Disinae, excluding Brownleea. The second clade (node 12) includes the зо as well as Hut si ee iia cia ih: the Sa- tyriinae are eis an with the Disin e at node 10 (Fig. 46). The characters support: iis is arrange- ment are i die subterminal s “чеп ру эи їег 29) and the brightly colored sepals ( ter 2). The mer character Be: Hooker (1883) and тун (1889) to diagnose this tte while Rolfe (1912-1913) separated his Di- seae from the бшуше һу the li p not реш! fused to th sensu Rolfe is paraphyletic, a as the Diseae are Alei based on the absence of did this finm criticized the characters used by Bentham & Hook- a raised and sharply delineated Ne я саре mon DET DA r define the d a Vere RN It is therefore prie that, al- tvriina ger there i is ect support 2 of т e Disir 83), P as suggested by m & Hooker (18 (1889), С. апа Rolfe (1912- der entile may be affect ed t tributions for directing the pollinator into the 5 dorsal sepal. wn- An alternative hypothesis — be " Br e leea originated as a hybrid betwee кі. inves- and the Disinae. The cladistic iugis Volume 81, Number 4 1994 Linder & Kurzweil 705 Phylogeny and Classification of Diseae OUTGROUP HUTTONAEA DISINAE SATYRIINAE BROWNLEEA CORYCIINAE OUTGROUP SATYRIINAE DISINAE HUTTONAEA BROWNLEEA CORYCIINAE OUTGROUP HUTTONAEA DISINAE SATYRIINAE BROWNLEEA CORYCIINAE undamental tree 1.—B. Fundamental adi The outgroup is as defined in ск text. FIGURE 45. Th fi t Tee 2.— C. Strict consensus tree of the three fund tigated for patterns that might h h hybrids can be recognized cla dis- dr “When th e E Paz ere are 8 length and one taxon € changes, the taxon that is moving may be a and the two taxa which it is TIME may be Де PONI In th patt terr aw sma ныр length t trees th р ts below the 706 Annals of the Missouri Botanical Garden = = > £9 < = c —E ae = wow m ж ч = EX = © 5 oou. a = = E — ui ш с сб ж m ы оо = = э” >с a =. ade - => => < l o = асын а= асч £ Sens sak п Е SESEEESSESSESEBEEL SS HS ES 25° 26°. 18 а | 19 28% 5 13 1 28 12 14 n" 15 22 24 oo im аа 27% 23 20 8 918 23° 7 Uc 17 5 57E 27” 9 4 4= 11 6 3 2 4 10 11 31 27° 10 21 = 9 14 Æ 8 29 2 30 18 ==? E U 19 T x 14 46. One of ihe: two shortest cladograms of the tribe, showing the distribution of the characters numbered CUN to Table 2. Solid " volved several times and dE UR wavy lines id reversals. The larger numbers at the nodes indicate the node а ud referred to in е tex Volume 81, Number 4 1994 Linder & Kurz Phylogeny in س‎ of Diseae £e E sS = A e шы < 5 сс = < сс = ОЕ = Жы лыш. ЖЕ AS Do ru dE IU OS Бы. Б. one m HB 21 сз БЭ 2 c3 eee LE rr ow Eo co RO e eu VENE are PRU CH BS m nb e op n x» сї ££ C. Б Б ае єсє == z c5 dC = HOS c oc uu X EX. МАЗ ap. wp CE (C e COUR Nn cu ee ЖУ oar igs) CD Eun э «9 ©з €. 00 сэ, 4 АЛЕ CO m. ж e ба OO NU um oO 1 2 3 4 82 (83) 20 (25) 93(92) 91(90) 5 6 a 34 (29) 84 (99) 56 (58) 71 (64) 77 (87) 45 12 13 (31) 17 (45) 13 62 (71) 5 Л 3 h A, d 47. Cladogram aps in Figure 46, ries the bootstrap p m ed enm he pres calculated w débain ib | козак igur Annals of th Missouri ^ Garden hybrid are variously arranged; but Brownleea is е assomated with the Coryciinae. ssible to follow a path of char- e should be in this case, as Brownleea mui of the Coryciinae as (WR as ine of the th sinae. bie усш Phipps et al. (1991) in an analysis of the Ma- lo ideae ева wio are кез he. pure hy- brid origin, sugges incon- within the maloids "ih have been the y evolution within the clade, did = the hybrid brisili Within Brown- leea there are also reversals in quite ““fundamen- tal" characters and unusual character combina top (idee id P in prep.), wi might (8), a lip that is зей to ш E (10), a stigma produced only pel (but see the discussion УГ! fae a avs 0), as well as the stigmatic part of the median carpel developing шп two be ig (31). Ee di рш i ‚ 5) and the petals which are d hadd to the gy- nostemium ( ing Brownleea from one lade to the other, the clade without Brownleea i ee rendered реа which it would o course be if Brownl of hybrid orig sole e by asado: ” This applies, Disinae, which are here йш аз np Е If сше жеге to be incl Disinae, would by seven characters, of which five are conflicting. This would identify the Coryciinae as the other parent. (4) “Taxa with reversals may be hybrids." Brownleea shows no reversals (5) “Taxa without ын ا‎ may be par- 3). ^» - E о z = @ Е. Ta Md m zu о Ф 3 3 sus on a strict con- sensus une шау! һе Haud toa hybrid зш Thi is 8 у L 1 1 5C). To test whether Brownleea is OUR the collapse of the —— nho taxon was remov from the data-matr! vith SL © E eet produced a tingle сові ‘parsimonious sree, ssive weight ing d igs. 46, 47), with a cónsistency index of 73 (compared to 67 for the analysis includ ing Brown- элө thus clearly demonstrating that the problem lies with Brownlee Passi сүйөп for hybrid recognition can be introduced. If the hybrid is removed from the creasing from 13% to pides and den уенына from 17% to 45% (Fig by a hybrid origin Although h species have E been demonstrated cladistically (Funk 8 ; igher taxonomic levels (e.g., the Maloideae, Phipps et al., 1991). SATYRIINAE The Sat мү including Pachites, Satyri- dium, and Sat weakly sup- form a rather ported group, with only three synapomorphies and “boots percentile” of 56 (see Fig. 47, but this may well be a id ires Por as it is spe ancestral Org patacgos condition po ay J oo also be used as a character, but this recurs p iven much ” linkin, As the one er feature us ii re is the € gated ане Lia they are probably best ke © Saty m DISINAE e Disinae, even with the removal of ue leea, for pia supported group. 2° di кын. S 6 (Fi me node is sup! = ed by node 6 ( ж-е 7). ea Kies with Brownleea, and на a “bootstrap Рег" , and Volume 81, Number 4 1994 der & Kurz 709 ieee and Mita of Diseae centile" of 71. Neither of the two ARR Toi Eee fo r the group incing. Cha pe the presenca, of apiculate sepals may E is very con that occurs widespread in the subclass (see ibe would be much better Segel discussed above. The included taxa show a great deal of phenetic RONEY, thus роботі ing the ge- neric classificatio hy- letic. This | is indicated by the lack A лы. within the clad CORYCIINAE The Coryciinae form a clearly monophyletic ciinae sensu stricto (Kurzweil et al. ‚ 1991), cheat the genera Evotella, Cera- Мао; Согусіит, апа Pre node 7) are very well s тч рен of 8 x i Riposi is plas (the wed by all previous authors), then only: two Aine are Synapomorphic for the group, but the “bootstra эе cs is на үнү high и Тї: This ee lev be xod адлег кее If n were to apes nailed. from the Cor iinae, it would hav атша ent aining a single genus does not carry any in- пац and the information that Disperis is the e sensu stricto can be y 1). On balance, it would appear to be more informative to include Disperis in the Coryciinae. BROWNLEEA he phylogenetic Pune of Brownleea could ч 0 by sequencing criteria, thus obviating е need к » эрес) subtribe for me e genus. How ever, {го t that there is irs аыр о. for a clade cludi ing Brownleea and the Coryciinae, d ie. oor node at which to establish urther infor- fore this would be a e ay nai Formal taxa should be esta nodes with a high "pootttrap percentile," be- to new data. ition, as was argued бә there is a t at may hav h monophyletic mazon. as it à ERES would не genetic знаци is the single most important criterion yes taxon (Wiley, 1 1981; Bunk, 19856), this would | argue o7 inae. Hybrid taxa are polyphyletic, and the beet approach ma to define them as narrowly as possible, and to give them the same rank as their parents. Placing Brownleea in a distinct subtribe from biologically — ed, the group becomes biologically more diverse, а predictions or gen ral biological + statements ssifications should be “maxi mally inform ative’ Оа 981) ог аѕ зе evens (1985) ane NNNM of e (or all taxa) can be adjusted to serve y purposes, ea. that es E separated Da a pisse mor- phological g We ot argue that because the “bootstrap percentile” is rela tively iw. Brownleea à is pees: ically га iu 1er dues that. it тау be of hybrid origin sanpor aisc it in its own је НОТТОМАЕА If Brownleea is placed in its own subtribe, the it аи that Huttoneae must also Бе гес ognized, мкл at the же. level, in order to maintain nophyly cri r, the Fem does change if the hy- it would be the sister group to the Coryciinae as Annals of th Missouri aca Garden d E vle There i is шше! character dei ed from the Куз; the * “iota percentile" is a low 45 and there is onl single character suppor a oo i is well understood ms of its taxonomic diee baron: As w uh a bye ents зы һе ciinae. First, Huttonaea is in Picea Very porabor: and hok cain pias ed in its o ubtribe iT) (1021) The pollination biology ва not known, but the D temi h little 1 bl h g to the Coryciinae, and the perianth is very different. This pe degree of disanak would be lost if they to be included in the same subtribe. Second, is bootst tile" for the node, even with Dromniven excluded, is a low 45, reheating the for the group. This ae that the node would be А уегу susceptible un new ины " EE is at to retain mis ишш until more кызны is available FORMAL CLASSIFICATION Tribe po ara AE. ip usually spurred, ney erect. This t a few species reachin, Subtribe § pe SATYRINAES нейни, Ха Bot. Gart le 6. Satyrieae на) Szlach., Aes ا‎ nou 26: E tals similar to the lateral S col- ees ted. Ca. species. Packs] Td. robably not mper pios et Lindl., G . Orch.: 345. 1838. 989). Satyrium Swartz in g. Vet. Akad. Handk. 21: 214. 1800. mis genus has et been revised r its whole авик range, and estimates of the number of species range from ca. 100 to 117 (Summerhayes, ; Hall, 198 ut Linder, 1989). 968 ar DISINAE Bentham, Gen. Pl. 3: 464. Dorsal sepal spurred; petals adnate to the gynostemium; sepals MAR pollen e rugose or ul 8 sp urfac зит Lindl., Gen. Sp: Orch.: 358. 838. Six species (Linder, 1981b). Disa Bergius, рен Р]. Cap.: 348. 1767 species (Linder, 1981c, f). посева Rauschert, Feddes Керегі. 94: ide ent problematic Ransche rt A DR ggested that the n eit was originally wn Lindl., is a ognizing t aturally e correct name. 18 species (in der, Monadenia ной ien ede 356. 1838. Subtribe REPE N weil, subtribe nov. TYPE: Brownleea Harv. €x Lindl A Disinis labio erecto, petalis sepalo d nnatis, a Coryciinis petalis gynostemio ion a recedit. 1 1 fused to the base mium; petals adnate 10 ds 58, es and kd: to the gynoste- Lip erect E mium. kamne Tu ex Lindl. in Hook. Lond. 1. Bot. M bdo seven species (Linder, 1981a, 1985) aes пролаз Schlechter Notizbl Bot. G m 9: 568. 1926. Hut rae Got che Sach Folia Gobel Phy’ totax. жоры tigma кро only from the er carpel apex; anthers diverging: clawed and fimbriate. Huttonaea Harv., Thes. Cap. 2 Five species (Stewart et rad po Linder & Kurz 711 Volume 81, Number 4 1994 Phylogeny alah н of Diseae p CORYCIINAE Bentham, Gen. Pl. 3 е Ecklon ex Bauer, Ill. Orch. Plants 464. 16. 1827. 883. Corycieae (Bentham) Szlach., Folia Geo ^ Phyt 6. 1991. Disperidinae = pecies aes et al., 1991). a sai erb. Boiss. 6: 800. 1898 San Karev eil & Linder, Pl. Syst. Evol. s only from the median 175; diis apex; lip with an appendage Es us ono fated hl to ‘he gynostemium. Ca. 1 a dium Sra, Kongl. Vet. Akad. andl. 21: 217. 1800. Disperis Svarz, Kongl. Vet. Akad. Handl. E species (Kurzweil et al., 1991; 21218 Hunt, 1968). me is a wide-ranging npn reach- i ic Saris Kongl. Vet. Akad. Handl. ing from southern Africa ен Guin- 21: 1800. of species - The current estimates are ca. 84 (Verdcourt, 1968; Manning, pers comm.). ae species (Kurzweil et al., 1991). KEY ТО THE GENERA IN THE DISEAE la. Col 1; petals and sepal ү similar (Satyriinae). 2a. Lip notte two spurs i d rarely lost; lip galeate or hood forming. 3a. Viscidium Solar s pie pulvinate Satyridium Viscidia usually paired; stigma flaplike Satyrium 2b. Lip scarcely different Tom the petals and never spurred. Lip with small side Pachites bodkinii ит с А уут ia ie ol Pachites appressa lb. on mn-par ing; p and sepals generally quite different . Petals rua аф kae dee fimbri iate; dues T diverging (Huttonaeinae) ....................... Huttonaea "b . Petals n athulate, entire or lobed (Cory and Brownleeinae). 6a. Pet "uini t to the dorsal sepal to ride a api lip Sanne at the base and appressed to the sti si Lateral Disperis Tb. Lateral Sik never Meg or saccate. Brownleea 8b. We sepal never spurred. 9a. Lip-blade an e "shaped, rarely rhomboid or reniform. 10a, Lip a prominent callus 1... Ceratandra 10b. Lip teet a callus Evotella 9b. I lla. Dorsal sepal and send forming a deeply globose h 2 11b. Dorsal sepal and petals forming a shallow to flat galea „ Pte 6b. Petals free from the dorsal вера; lip never appressed to the stigma and generally patent a the base 122. Lip complex, with an ascending basal part, flat middle part, and toothlike apex ..... Schizodium 12b. s ЖЫ, rarely lobed or fimbriate. scidium s eye? or rarely paired; petals and lip fleshy; rostellum consisting only of a massive ral lo be Monadenia 13b. Viscidia peared « or rarely | һгее jo l4a. Lip ovate, sessile or s viendo the margins often fimbriate or lobed: pet trongly falcate; deis grasslike ~~ Herse ke 14b. Plants not as above EREN EN Ы Disa L ITERATURE CITED 1888. The orchids of the к Peninsula. BARTHLOTT, ыг 1976. Morphologie der Samen von ШЕ i але Philos. Soc. 5: 1-20 chideen eigen ai auf taxonomische un 1896. Icones vesica Austro-Af- tionelle As is sn oceedings of = Eighth World ricanarum s; e tl Vol. 1. Wesley & Son, Orchid Conference 1976: 44 4-4. BENTHAM, G . HOOKER. prt а Plantar- Во, шп, vol. ~ L Gone: London Us, Н. 1882. Notes on р Һа. J. Linn Soc., Bot. 19: 233-238. London 911 Icones Orchidearum Austro-Africana- rum | Extratropicarum, Vol. 2. Wesley & Son, Lon- don. 1913. Icones Orchidearum Austro-Africana- Annals of the Missouri Botanical Garden rum epis Vol. 3. Wesley & Son, Lon- don ado cd xk: lx s А2 ns Cape To Burns- усш, Р 1,983: Рф theor f the exine in d Оса. Amer. J. An 4-1 &V ‚Ким of the буй ом. т Contr. Bot. 61: 1- 79. CARPENTER, 1. М. quality 1988. Been among multiple Cladistics 4: 291- ie MW . & J. S. Pippen. 1988. Seed morphology in the Oncidiinae p Py subtribes (Orchidaceae). Syst. Bot. 13 323. CHESSELET, Р. ر‎ Implications of Leaf rr: and укалы the Disinae and Cory- ae (Orchidaceae). M.Sc. "Thes esis, University of Cape Town. & H NDER. 1993. Pollen morphology of bed Diseae E Orchidaceae). Grana 32: Gnas! Py T. & W. K. Sur 1969. phology sur sification of aa road ا‎ -139. hi ا‎ History and Classification. Harvard Univ. Press, Cambridge, Mas- achusetts, and London. 1 ecent advances in orchid phylogeny. Lindleyana 1: 5-20. 990. The bue ieae in hid classificat m 5: 102-1 Farris, J. 5 196 9. ra successi ve approximations ap- I weighting. Syst. Zool. 18: 374- 85 —. . He ا‎ T 1985. ennig86 Reference. MM jars 1988. Confidence limits on phy- pap 6: b opes using the bootstrap. [tud 39: 7 Funk, V. pe The systematics of Montanoa (As- нас арааран. Mem. New York Bot. Gard 6: 1-13 198 Жы Phylogenetic preter ce done. tin. Ann. Missouri Bot. Gard. 72: Sone . 1985b. Cladistics and generic ки in the Compositae. Taxon 34: 72-80. HALL, A. V. A revision of the southern кен species of Satyrium. Contrib. Bolus Herb. 137. HEALEY, P. L., J. D. MICHAUD & J. Авртт Morphometry of flor seeds, III. Native сар ие related | species «сена ы разе, Ps tanthe- ёз. Bot 67: Horan, C.J. 1981. „Олиро апі cladistic stud n Anacyclus L. (C die J. Bot. 1: 83- = € P» à x dium, Orchidaceae, part E. Milne-Redhead & R. ich ilis Flora a "Tropical East Africa. Crown ыы gus d Е Kaptan, D. R. 1975. Comparative developmental eval- monocotyledons. Bot. J yst. 95: 1- KRAENZLIN, F. 1899 Ў Отаси Сое et Species. Mayer & Miiller, Berlin pesa H. 1987a. Developm std studies in orchid wes ke restent and vandoid orchids. Nordic ede studies in orchid flow- ers. а point species. Nordic J. Bot. 7: 443- Developmental studies in orchid flow- ers. ш: ен species. Nordic J. Bot. 8: 271- 2 ————. 1989. Floral pier m ал ш Huttonaea pulchra. Lindley 1990. Flor: al nice P d in Orchidaceae subtribe Disinae. J. Linn. Soc., Bot. 102: i 1991. The unusual structure of the gynostemi- i Bot. Jahrb. Syst. 112: 273-293. The remarkable anther structure of Pachites bodkini (Orchidaceae). Bot. Jahrb. Syst. 114 Seed morphology in southern African m da (Orchidaceae). Рі. Syst. Evol. 185: 229- 247 & H. P. LiN 1991 А comparative study of the floral Rss ye n the gm. E peris (Or- chida ceae). a pws wt 66: 433-477. CURES LIA & Р. LET. looi. “The phy- log ae огусшт complex (Corina, танта: РІ. Syst. Evol. 175: dtl re Р. y odi Ta euim. x the genus un с Lindl. J. 5. (ons Bot. e Disinae: e Disinae: 2. A revision of ihe. genus dem Lindl. J. 5. жч) Bot. 47: 339-3 ш. 1с. Тахопотіс studies о n the Disinae. A revision of Disa Berg. e: "s pa Micranthae Lindl. Co en. 9 -370. ntrib. Bolus the Disinae. ү. 1981 А avri: at the peri - Monadenia Lindl. Bo EE -363. 81е Taxonomic studies in Ње Disinae. VI. А revision 96 the genus Herschelia Lindl. Bothalia 13: 365-388. " — — —, 1981f. Taxonomic studies іп the Disinae. uk A revision of Disa Berg. sect. и. Micranthae tl. Bel 3 = d «t ps ызын а of southern tropical Eo I: Brownleea and Herschelia. Kew Bull, 4U: 125—129: 86 Notes on the pios of the o" феде, e ing icular reference to the Diseae- dleyana : T poner rostratum. Fl. Pl. Africa E pl. 1993. 1991. Confidence limits in phyloge pe from the African Restionaceae. 253-2 os P" a] morphology Flor е), ]. Linn & Н. Kurzwen. 1990. and phylogeny of the pen i (Orchidace Soc., Bot. 102: 287-3 Volume 81, Number 4 1994 Linder & Kurzw Phylogeny and nisi. of Diseae 713 LINDLEY, J. 1830-1840. Genera and Species of Or- chidaceous Plants. Ridgeways, London. MANNING, . P. LINDER. 1992. TERTI “ evolution i in M (кем, zt why e ther ? S. African J. Sci 38- 49. Blast; R. "895. The fertilisation А din uniflora an + by insects. Trans. S. African Philos. Soc. 8: Boag E. 1889. Orchidaceae. In: i н Die Natiirlichen Mu e m >» mann, MAE sida ^ s . RoB N, J. R. RoHR ius ITH. Origi ns and evolution "s sd Maloideae coii Syst. Bot..16:.303-332. Mos F. i f th t chida. Opera Bot 65: 1-96. rag zur Nomenklatur der : li & P. G. Or . Thiselton- yer ро. A of {Tropica Africa VII: 12-292. п: W. Т. Thisel- ton- Dyer (eto uer Poet Vol. V, Sect. 3. L. Reeve, L Qn omparative leaf morphogenesis in 141-260. W. РЕЕ dns 1977. Untersuchungen an idee enpollinien unter besonderen pie ep a g ihrer F xin Pollen & Spores SCHLECHTER, R. 1898. маш» der Disperideae. _,% Дер erb. Boissier ser. 1, 6: 800-821, 846-859, 1901. Monographie der Diseae. Bot. Jahrb. = 31: 134-313. 1926. Das sien der a Notizbl. +. fot, Gart. Berlin-Dahlem 9: 5 SENGHAS, K. 1973- 1974. и? p К. G. Brie- The pollination of Disperis ( ix. ca. iz] „Р.К. 1985. The е genus concept in practice — ut f. ad ыз practice? Kew Bull. 40: 457-465. 991. Character states, morphological vari- ation, e phylogenetic analysis: A review. Syst. Bot. 16: 553-583. Stewart, JH P L R, E. A. SCHELPE & A. V. H 1982. Wild Orchids ПОО ks. MacMillan South Africa, Johannesb urg S N = Orchidaceae (Part 1 1968. ). In: ead а К. М. Polhill (editors), F z of n , London یو‎ O. 1800. Orchidemes slagter orch arter up- tallde : Kong Vetense. Acad. Nya Handl. 31: 202- SWOFFORD, D. = йү PAUP: dini тено Analysis Using Par: Version 3.1. A User’s Manual. Ilinois Natural ior Survey, Champa 1991. New ZLACHETKO, taxa within the order E ا‎ Ge hytotax 15- Tonpa, Т. 1983. Seed morphology in Orchidaceae I Dactylorchis, Orchis, dad fs iras Н adenia and Galeorchis. Sci. Ned ku Imp. Univ., 4, Biol. 38: 253-26 Tiles R. 1864. On i^ стоя 2 Disa gran- diflora, J. Linn. Soc., Bot. Pp 216 ie Uer боры ылы (part 1). E. Milne-Redhead & R. M. Polhill dir Fre of ет East Africa. Crown Agents ersea Governments and Administrations, Lon- s VOGEL, S. 1959. Organographie der Blüten Kaplän- discher Ophrydeen, mit Bemerkungen zum Koap- tations- Peor . Akad. Wiss. Abh. Math.-Naturwiss. Kl. WAGNER, W. L . STOCKHOUSE & W. M. KLEIN. 1985. systém matics а evolution at ‘the Oe- nothera а за species complex (Onagr: е Mon Syst. Bot. Missouri Bot. Gard. 12: WEALE, M. J. Р. 1873. tive tion of Disa macrantha. J. Linn. Soc., unt 157 45- WEBER, A. 1980. Die Homologie des Perigons der Zingerberaceen. Ein rag zur Morphologie und Phylogenie ior > Monekotylen -Perigons. Pl. Syst. E 133: 149-1 ERDELIN, L. 1989. We are not out of t yet—A ioi from a Nobel symposium. Guages Ё 0. WILDHABER, О. J. 1972. Zur Karpologie von Orchis. In: K. Senghas & H. Peng ей) Probleme der Orchideengatug ipee is 25: Mer Macon ertal WILE 1981. Phy к Һеогу апа Price of Phylogenet vas і yii Wiley & ns, York WILLIAMSON, °С. 1977. The Orchids of South Central Africa. J. M. Dent & Sons, London. ZIEGLER, В. 1981 Mikromorphologie der e unter Berücksichtigung Taxonomischer As- pekte. Ph.D. Thesis, Ruprecht Karls Universitát, Heidelberg TAXONOMY OF ТНЕКМОР515 (FABACEAE) IN NORTH AMERICA! Chia Jui Chen,? M G. Mendenhall? and Billie L. Turner ABSTRACT s to our recognition of 10 species. Three species. occur in the southern “Appalachians: T sae у pe and lead: T. fraxinifolia. The Rocky eas and intermountain specie: ivaricarpa, T. ana, and T. rhombifolia. {е 1 dominated . то . Th { Са by the variable T. ur diee nes three infraspecific iua. which gives. way to T. АТ. їп r pen and western Oregon. Thermopsis macrophylla ai nd Т. robusta Widely distributed throughout the western and pond GREKSA is a con- 3 = б Е LE © Ў @ eedy h it is a her- plant, its ө relatively ү habit and stun- о Е во olden nin er bnght Yellow Sowers and Pit li early and t сч зын eo xm factors, кечуа» should have led derstanding of its systematics, pam a “lack nostic characters within t America, though over 20 spe- eho proposed. е current study is based on over 20,000 km in the western United States. While сэс и Ld Ad into two highly TRIN тес we recognize four m cies in t ‚ three of which d th hp ne c °1 3 1y сїзїтїршїеа tain species in the Rock 1 regions. The three long-recognized species of the southeastern United en are maintain The following herbaria have Miam supplied Ke proximately да. imens hu number y Mountain = ” ARIZ (113), CAS PER P COLO n e F (193), GH (511), JEPS (16), LL (15), MO (444), мо NTU OD, X uo p 9), NEBC (3), Y (715), ORE (78), О PE (52), PH du POM d: RM ait A (302), TENN qu ا‎ (163), UC «и US (419), UT (83), 161), WTU (20 HISTORY OF THE GENUS = lupine + “op- Tee e A k “thermos” sis appe ori sis lanceolata, which is contrary E ode of Колея, Nomenclature (Greater et Mes recognized s still, pr e the genus. In 1818, Nuttall called the g Scholar Fe llowship LS XE thor to chs gee in the silices of the manuscript. 2 [ns of Botany, Ас sd — Xiangshan, Beijing 1000 ent of Botany, U ‚ Departme M “of Botany, University of Texas 3 Au ves ог ба correspondence, Depart * Plant Resources Center niversity of Texas, seni scie Hi. inar We also sp Guy L. Nesom, Саг о iey he gem ity of Texas at pct and Peter C. Hoch at the Missouri Botanical "е 93, Chin ustin, Texas 78713, U.S.A. aes Texas 78713, U.S.A. ANN. Missouri Bor. GARD. 81: 714-742. 1994. Volume 81, Number 4 1994 Chen et al. Taxonomy of Thermopsis Rafinesque erected the genera Scolobus and me based on specimens of e she saw ERI and 18 western speci s to 7. 3 p^ (1941) ig to bones ey's ravinon Though variously treated in regional floras, the t significant work on Thermopsis since its re- in Th marily by lowering these to varietal status. to support the specific rank. мнен е he saw that “all taxa are gly confluent with those geographically contiguous.” His views were not universally тч аї time (Turner, anes support еа of most these taxa as pa arate species. Thermopsis is the largest and most widespread many other morphological a eco chemical dab DUE. In n faet, i (1940) referred to Ther. psis a estern dieci унн h i ore alkalo id denim and тор Since a thé woody erm. id 1 1 ry J 1 lati to the Genisteae (Polhill, 1976; Turner, 1981). BIOGEOGRAPHY Thermops er not se bale orth Ameri 13 species are foun oss China, Mongolia, a 1a, and beyond to Japan, rs eastern oin and the North American coast of the Bering Sea. Most f the Aci: s 4 ALL (1OAL\ udging from 52s relative edge of the North J American specie ааш ancestr al sd ex каан: Ther- d DER p y er On the North American continent, Thermopsis is distributed throughout the western states and in states as T. villo a as well a ia, but at lower Шош, typically be- tween 300 and 800 m x western Жек Ао: Thermopsis occurs the Rocky Mountain and intermountain iM as purge as in 6 Расібс P ne E species; E He fend as two o intergrading, regional varieties in 3200 m. PE тодын and habitat are М ro its range from Montana southward to Arizona and New Mexico and northwestward to Wyoming, Idaho DR: а о сеа Found at ina low elevations, 1000-2000 m, is Т. lecce Although this taxon can be f e fro and e iei К hop Mexico, region from it is ашк restri y grass- lands. Thermops ае occurs at middle to "т elevations, 2000- 3000 m, in the Rocky Mountains from Wyo o New Mexico, in me e conifer ‘aes Most of the mopsis in the Pacific coastal states | is found in California. In central сее, ifornica is со ur egon elevations, to оо, m, in Ї 1 : 4 distributio tions within California. Thermopsis califor- nica var. argentata has a disjun sternmost and southwestern Californi at high elevations, 1200-2200 m a macrophylla is known only from of Santa Ynez Peak in Santa Barbara County. Our 1 Ks h » Stu ка п erroneously applied to the more common 7. Bios var. californica. Unlike the latter, T. marran учар localized, relatively uniform, and strikingly robust. Equally impressive in es is MAINE robusta, which is found only Annals нан и Garden TABLE l. Chro MSN ыр їп ад Эч, Counts listed with voucher only iginal meioti ts by the junior author (T); t TEX. 2n number Reference/ voucher Asian species : эе ui Regel 18 pet N (1969) a Benth. ex Royle 18 en et al. (1992) т cH Benth. ex S. Moore 18 5 et al. (1986) T. chinensis Benth. ex S. Moore 18 Chen et al. (1992) Т. lanceolata В. Br 18 Mesicek & Sojak (1969) T. lanceolata R. Br 18 Chen e 92) T. licentiana E. Pete 18 Chen et al. (1992) T. lupinoides (L.) Link [as T. fabacea (Pallas) DC 18 Federov (1969) T TUA (L.) Link 18 Probatova & Sokolovskaya (1981) T. mongolica Czefr. 18 Mesicek & Sojak (1969) T. przewalskii Czefr. 18 Chen et al. (1992) American species F. серй var. argentata 18 California: Modoc Co., T. 5813, T. 5816 T. divari 36 Colorado: deg on P T. 5636 T. Ss 36 Colorado: Las Animas Co., T. 5772 T. divaricarpa 36 Colorado TEE Co., Tomb 649 Т. gracilis 36 Oregon: Lane Co., Т. 5 T. macrophy 18 California: er Barbara Co., T 5820 montana (var. montana?) 18 Federov (19 T. montana (var. montana?) 18 Colorado: HA Co., pe bes T. montana var. montana 18 Colorado: Saguache Co., T. montana var. montana 18 Idaho: ou d. ers T. montana var. montana 18 Nevada: Elko Co., T. 5795, T. 5796 T. montana var. montana 18 Utah: Wasatch Co., „Т. 5794 T. rhombifolia 18 CA dingham (1957) T. rhombifolia 18 CANADA: Saskatchewan, Taylor (1967) T. rhombifolia 18 CANADA: Saskatchewa T Federov (1969) T. rhombifolia 18 CANADA: Manitoba, Lóve & Lóve (1982) T. rhombifolia 18 Colorado: Huerfano Co., re “т 5773 Т. оШоѕа 18 Zhukova (1967) ally, nica var. semota is e e woodlands of San Diego Coun CHROMOSOME NUMBERS The first chromosome counts for Thermo opsis were ers by Russian pi in 1931 (Fede- 69). These were obtain the species investigated had also been on a base of x = 9 (Table 1). ploids (2n = 36 known atl two species, "T gracilis and T. divaricarpa, both Amer- ican, the former occurring in northern California and adjacent Oregon, the latter occurring in the front range of the Roc ky Mountains, mostly in central Colorado. Polyploidy is believed to have independe in the two taxa, largely be- these two taxa, however, ns it remains to be seen if such counts will hold throughout their re- spective distributions. А base chromosome number of x — 9% found in the closely related Baptisia, a genus onfined to lower Rae about 17 specie tisc 1981 of the southeastern United States (see Isely, 1 к= Ог overview). To ч по polyploids have reported for Baptis SECONDARY CHEMISTRY (MICROMOLECULES) Unlike in Baptisia, {төн and alkaloids kei been of limited и ss in ce of б variation in Therm mopsis the & Gott- lizidine alkaloid distribution, Salatino Volume 81, Number 4 1994 Chen et al. 717 Taxonomy of Thermopsis lieb (0909) меге i ip divide Blusen de into ican specie: А BoP ENP phyloge eny, oe ever, они not yet been proposed since alkaloid and flavonoid data are insufficient (Turner, 1967; De- ment T ‚че 1975) апа нер и (Balan- 1982). bap chemical data ане бв used, попе a piylogenetir er ‘cale, Based on ino & Соні eb, arose Genisteae (Bisby, 1981; C tofolini & Feoli-Chiapella, 1984; Polhill, 1976; Turner, 1981). MorPHOLOGY Thermopsis has been commonly referred to as "false lupine,” “‘buckbean,” **goldenpea," “golden "sw and “bush pea." Though the — m the genus and is not t ord useful for specific delimitation. There a ss r identification iw characters are used to distinguish pai Temopis and Baptisia; compressed Ri are The hos sture and shape of the mature pods vary ely within Thermopsis as well. The fruit erect, ascending, or ipia: from the xi ographical ruits, and nt will also be arcuate or ty (1981) | а that ovule number is dni within regions of the generic distri- sou’ ia and interm cies have 10—18 ovules an the rp a: have 12-16. These figures do overlap, however, and can be called trends at t. etative characters are often quantitative with ан variation around average proportions. Nev- characters тато wide diagnostic t ant height leaf size thor igh taxa, especially varieties. pone the leaf apices may diff sin B & @ .8 a б z 2 oe in size, growing to 9 cm long si 6 cm na. шше гелсе Мас likewise, оз Касет шау bear pm or {зам pedicels, and these stad d t addition, the flowers may be whorls, even where the dense. Within ше flowers, calyx E тем bon Thernopsi species ا‎ metimes, as in 7. mollis and T. rhom- bifolia, pg extensive clonal populations. Oth- erwise, the plants are supported by thick, perennial бошо KG. E is often арі to confirm the es- pecially of з plants, which typically дури just oe rhi- sheet. Yet field pyama е that a given taxon will have a tendency to form rhizomes or not, and this character should bs к during oted for identification. stems, branches, and fruits, the angle of ribe erect, ascending, p n the following descriptions, at the signage Though “рге and Sito od are roughly synonymous, here i ч used for Mida and “divergent” is ie for frui SPECIES AND VARIETAL CONCEPTS he primary difference i in the various ee. * Thermopsi sis has been treatm ment, the te specife rua has been chosen to exp what rceive А EU d above Annals of the Missouri Botanical Garden о be evolutionarily independent taxa. We have productive independence fr pecimens and observations of populations in the field. For example, altho si occasional sympa hybridization is suspected bet ; divaricarpa and both T. сини е a rhom- bifolia, there is evidenc rids have stabiliz and | persisted. e eg Isely occasionally, if n where we, like Larise, see none. Ne attribute the res- д n of па на Pu shat à is the basis of this dp. TAXONOMIC TREATMENT ыште ae R. Br., Hortus Kew., ed. 2, III. 3. 1811. TYPE: Sophora AR E = Ther mopsis lanceolata E. y 5 pinoides (L.) Link s Nutt., Gen. N. Amer. РІ. 1: 282. 1818. TYPE: Scolobus fa Amer. ` Monthly Mag. Crit. Rev. 4: 9. TYPE: Scolobus rhombifolius (Nutt. ex pa pus. Raf., Ne 122552: i ut TYPE: Drepilia rhombifolia (Nutt. ex Pursh) Raf. Perennial herbs, usually rhizomatous; stems erec or ascending, few-branched to well-branch branches ascending or spreading; plants icc to diee pubescent, unarmed. Léaves puce rifoliolate, sent; pedicels 2-17 mm long. Flowers papiliona- ceous; calyx campanulate, appearing 4-lobed, sub- equal, the upper lobe double, broader, and slightly cate to emarginate, the banner very jen ovate, shorter than wings an ke el, Aci f z "We aca claw 4 A 1. asymmetric, i to log rhombi oblong; keel as etrically uet vns lip to car inate, auriculate, posteri iorly fu fused, without a beak, claw narrowly oblong; stamens 1 e, uniform; nthers dorsifixed; pistil short-stipitate; ovary ob- long, usually longer than lous; ovules 5-15; style glabrous; stigma min egumes per istent, erect, ascending, or di to br , tan ‘own, y elliptic, lateral compressed, Moe ar- € or rarely ann cons e of ovule ай бл or lomen- rown-black, oblong to reniform, often with a mi- nute, membranaceous rim-aril. Base chromosome number, x = 9. AE eê ji relationships. Thermopsis is e Thermopsideae Yakovlev (Bot. ГЕ 57: 585- 595, 1972), by virtue of its free stamens and seco trifoliate, репо i шеш foliaceous or nea arly so so, dimorphic, ihe lowermost amplexi t blade- " commonly ew to numerous, scattered or а 1 ‚ Wi ely ov. ate te Ss |‏ ا ا to lanceolate, acute to mna bia télé ab- KEY TO SPECIES la. Lam of southern Appalachian foto. Pes ules on primary axis mo 2b. Sap xis lodi d Ithough Th mops enê, i tis very s similar — to aped e other Lees de orth American member of the tribe. Frui 8 Thermops жыт distinguished readily "us pono The Nor Xi American Therm ipe are et while those of Baptisia are inflated 2 As detailed above, Tero а is found in Asia and North America. In North America four species occur in the Pacific coastal states, three occur in e Rocky Mountains and intermountain regions, and three are found in the southern Appalac ct E e than 1 cm wide; P EM less than 3 mm long; кон erect, densely een sil less than 1 € or pris termin lb. Plants of Rocky Меш; intermountain regio 4a. Plants PINE аа OS 6. ingle woody rootstock; pedicels glabrate and at least as long d e ral; montane, usually dendi 2000 m 3. and Pacific coastal states. в tomentose or sericeous throughout; ба usually in whorls of 3-5; California. TEE is fraxinifolia Volume 81, Number 4 Chen et al. 719 1994 Taxonomy of Thermopsis 5a. Plants mostly А 8 dm tall; leaflets mostly 3-7 cm long; pods ascending, MAD ашы several о more abortive ovules mopsis californica 5b. Plants mostly 10- 20 dm tall; leaflets up to 11 cm long; pods ascending or ee usually wit! hi Teach ovules. be ascending ied — pedicels 2.5-4 mm long; lateral leaf veins 6-8 pairs; Santa z Mts of Califo: ermopsis лл 6b. Pods divergent and ‘arcuate pedicels 4- 78 mm long; lateral leaf veins 9- 12 pairs; Humboldt , and Siskiyo: lif 9. PÎ robusta AT Plants g P h of 40*N latitude, | portions of the western United States and eee Canada. de ight 7. Thermopsis montana a M т? es није» н pi ог annular. ак nches spreading, 5 d g ioles more than 2. 5e cm long; pods straight; western prias d northern California ...... mers rmopsis gracilis 8b. Branches ascending, weakly to derum vigzagged: leaflets less da 2c m wide, emarginate; petioles less than 2.5 cm long; pods usually arcuate or annular; МО. а еазїегп Rocky | Mo untain ins and adjacent Great Plains. a. Plants 3 1 cm long, lateral у eins 7-10 pai ipul 2. Ther: mopsis divaricarpa 9b. Plants mostly 1-3 dm tall; leaflets less than 3 cm long, lateral veins ne a pes stipules usually widely ovate; flowers 1.6-2.1 cm long; pods strongly arcuat nnular zer 8. Ther Raise rhombifolia 1. Thermopsis californica S. Watson, Proc. KEY To VARIETIES OF THERMOPSIS CALIFORNICA Amer. Acad. Arts {5 126. ыр. TYPE: U.S.A. ја, pue widely ovate to ovate, to 5.0 сш ee California: Marin Co., Corte Madera, J. M. racts to 15 mm mor е 1854 sore чом dos NY; than 6 cm long; pod pubescence of matted 6 NY: PIE US) trichom 1b. Thermopsis сие уаг. es abies lb. Stipules ovate to T ee о 2.5 c wide; Pu robust, (2.5)3-8 dm tall, sericeous or br о 6 m arco dens hse "tarii stems slender, erect, clustered, well- vis qa 6 Гар ipta xal aliy es ranched, arising from rhizomes; branches as- appressed nos n cen at 20—45°, weakly to strongly zigzagged. 2a. Silvery sericeous throughout; wing and keel Leaflets elliptic or obovate, 3-7(-9) cm long, 1.2- petals reniform; northeastern and south- = i p - cirea Californ: sick ian erties е т годен эа iba ыле siyay фонны; = Ther ermopsis ; californica var. argentata o villous, lateral veins 6-9 pairs, b. vs i dis conspicuously net-veined beneath; petioles 1-3.5 жир; oblong elliptic; restricted to San Di- cm long; stipules persistent, widely ovate to lan- dm Osborne e ee ceolate, 1.5-6. cm long, 0.7—5 cm wide, cuneat is аи californica var. semota 0 amplexicaul, ac . Racemes 7-30(-35 ng, 10-40 flowers usually horls of la. Thermopsis californica var. argen 3-5; peduncles (2-)4-10 cm long; pedicels 2-6 (E. Greene) C. J. Chen & B. Turner, sa mm long, ous or tomentose; bracts se E nov. Basionym: Thermopsis argent : Sistent, ovate to lanceolate, 6-1 8, 2.5- Greene, Erythea 3: 18 Thermopsis de 1 gracilis var. argentata (E. Greene) Jepson, 5-10 mm long, is mm wide at the limb, lobes Man. Fl. Pl. Calif. 51 Thermopsis 2.5-5 mm lo ong, as long or longer than t pve macrophylla var. argentata (E. Greene) Jep- and keel peat oblong жү to eer ОШ son, Man. Fl alif . x INT: velutin ovules 5- traight, as- U.S.A. California: Modoc Co., “Forestdale, cending ed ularly lenem mam young), Bak r 1893 eig selected here, ND; 6(-7) em long, (0.5—)0.6-0.9 cm wide, sericeous, E CAS, ND, NY, US) 5 Raine pg 1- б T6 LU aad In his on Greene cited a single collection u^ short rostrum, 4-5.5 mm long from m Modoc Coun y, without locality, made by Milo S. Baker Three varieties are es within Thermop- reveals two к collected by Baker in gem ce sis californica as follow Valley Mts." during June-August 1893. Both 720 Annals of th d lo Garden pee ies ое ist uen were subsequen o as to read Asa have designated the latter as lectotype Plants (2.5-)3-5 throughout; branches weakly to moderately zig- мы. сус obovate to со tic, 3—6. эс m long, 1.2-3 wide, obtuse to acute so а dien Sd prie F: гө D 55 +, dat, de t long, with 10-30 flow wers; pone s 4-7(-10) cm раш iiis ovate to lanceolate, 7-15 mm lon 5- е. Calyx 79-10) mm long, 5-8 vile at the limb, lobes 4.5(-5) mm long; banner cleft; wing and keel petals asymmetrically pomis elliptic to optem vules 8-11. Pods 3- 6(-7) cm long, (0.5-)0.7-0.9 cm = Seeds 1- 6, oblong, 4—5(—5. 5) mm me = mm wide. Chromosome number, Distribution and ecology (Fig. _ North- Californi alifornia (Kern Ventura, Santa на апа ро Angeles” бос) mostly interio I from 1200 to 2200 m; flowering May-June. Representative specimens — l- А. Са ifornia: Kern Co., Mt. Pino woe 6378 Ridge, above a Road o i Pollard 1955 (CAS, COLO, TEX). у 25 mi. "NW y Rey Peak, 23 Apr. 1934, Sowder 393 (U С). Thermopsis californica var. argentata differs from the more widespread, more variable variety dm tall, silvery sericeous . californica in its smaller habit and closely ap- pressed, eur vestiture. e are unable to distinguish between the north- ern and southern populations (Fig. i), Isely (1981) Т. eshet) only to the more northern pop- ulations ae included the southern NE. in me ни ma lla var. macrophylla the assessment of Abr rams (1944) that thes o characters by which to distinguish them though both are readily distin guished from varieties rae and semota g habit, vestiture, and petal s lb. Thermopsis californica var. californi- ca Thermopsis ee ee var. velutina E. Greene, Erythea EBE zoe ds t is velutina (E. Greene) E. Greene, pd . Thermopsis pee var. velutina (E. 2а ne) Jepson, Man. Fl. Pl. 515.1 diua macrophylla var. velutina (E. Greene) Larisey, Ann ds ssouri Bot. С: 256. 1940. e U.S.A. California: Santa Clar Co. unt Hamilton, eee 1891 (lecto M d lected Seis ND; isolectotype, ND). In his protologue Watson did not cite specimens, but е Hd to the species as oc n io: a , and prob ably ae : He did say that his Thermopsts californica was the sam t Torrey had re ferred to as “T. macrophylla” (Torrey, 1857) and fie ea" (Torrey, 1859). Watson, in his pio tologue, iden queris a eue with his citat ific Кай d Report in which Torrey cited a y from “Corte Ma- dera. is which is locatedi in Matin. in County, ожа 7 ontending lectotypos si since the Torrey Herbari rium collection at NY has Thermopsis cali- in Wat- at GH, whe were other Mess B lectotype collections anne” tated aks im as belonging to eins ci nic Plants 3- -8(-9) dm tall, sparsely to densely 10 : nches weakly to moderate'y (-9) cm long, 2-4(-4.5) cm de, obtuse г cute; petioles 1.5—3.5 cm long; stipules ит: ovate to narrowly ovate, 2.5-6.5 cm long 5.0 с ез m wide, cuneate to amplexicaul. Racem Volume 81, Number 4 Chen et al. 1994 Taxonomy of Thermopsis o T. californica var. argentata О Т. californica var. californica ӨТ. californica var. semota * Т. gracilis 4 Т. macrophylla A T. robusta MS : f. 111 2 1 S T А, 2 СД о 3 hd г ep Ј =, d les); T. gracilis (small closed circles); Р; f TL "i 1 CURE 1. is in Californi T, саті a (1 ge op ircles); T californica Ре : macrophylla (closed triangle); Т. robusta (open triangle). 10-30(-35) cm long, with 12-40 flowers; pedun- at the limb, lobes 2.5-5 mm long; banner cleft; cles (4-)6-10 cm long; bracts widely ovate to wi i keel petal ically obl llipti nearly lanceolate, 6-15(-30) mm long, 3-15(-20) mm wide. Calyx 5-8(-9) mm long, 6-9 mm wide © Г 4/ 7 © г , ovules 5-8. Pods 3-5.5 cm long, 0.6-0.7 cm wide. Seeds 3-5(-10), oblong, 4 mm long, 2.8 mm wide. Annals of the Missouri Botanical Garden Distribution and ecology (Fi gas Ua cog regions of central California from 35 to lat- rem mostly chaparral from near sea ec to about 000 m; flowering May—June. Representative specimens examined. U.S.A. Cal- ifornia: Alameda ias ‚ hills E of Oakland, Lu LE Tebbe 62 (UC). L e Co. ‚ 9 mi. from Lower Lake 1937, Ewan 10352 (CO 10, п Valley, - May 1920, Jep: fin WTU, UC). Mendocino Co., 5 May 1939, jos dp Wee MO, NY, a E Tni € Valley s, Wigg ris a3 CAS НЫ улы ri ts near е еу, 13 June 1861, Brewer 704 (UC, US) Napa Co., Knoxville, 8 May 1903, Baker 3081 (F, GH, , MO, N PA of Cal T2 . 1924, Heller 1383 7 (C AS, F, MO, NY, US). San Benito Co., ien ee GH, US). San Luis О) 17 кем along old road to Pozo, ay 1928, Eastwood 15140 (CAS, UC); San Luis Obispo, 9 М; 882, Jones 6 (CAS, MO, NY, PH, RM, UC). San Mateo Co., Crystal Springs Lake, "May 1 903, Elmer 4824 (ARIZ, CAS, MO, NY, OSC, UC, US, 9. ота чш Lake, 1 May 1902, Greene ёо (CAS и МО Y. UG, Е meri RI7 CAS, M ue х i i i May 1907, Heller NY, PH, UC, US, WTU); trail to Pine Ridge, N of Miu Springs, 29 May 1895, Smith 4 4075 (GH, OSC, RM, TEX). Santa Cruz Co., Glenwood, Davis ee (cas, GH, RM, UC, US); Bear ak) R d about api Creek. 18 May 1946, Ferris 1 uM (CAS, G RM UT, WS). Sonoma Co., betw and pe AE 22 i 1902, Heller ii Brow Le COLO, GH, MO , US; 6.8 mi. E of S оза on Ee oom ET i Apr. 1949, iris 12058 U). (CAS, UC 08 . the n жошону үп бише more plex. т is бой from both T. ш, var. semota and Т. cali entata by sparsely to densely tomentose vesti Brewer & Watson (1880) с QE noted that ermopsis macr a is distinct from 7. cal- rs have treated mac ine iyi a 1 98 1 ) who treated T. califoruica var. californica within . his concept of is — var. macrophylla, gm owards [T. macrophylla] var. ven a i4 gracilis і in the prono Eee th t eee narrow fruits." This statement was sul fied by the remark that “few specimens posse mature pods." Exa mo of many diverse spec- } d кен hiatus oe by Isely in mapping “the eal, and that the few collections of his T. macro, гэ ра var. нон. c that he maps within d range а T. gracie (of umen ilis йа аге otherwise superficia ally similar to 7. acrophylla var. macrop idenc suggested in Isely's treatment. le. Thermopsis californica var. semota hen & B. Turner, comb. nov. 1986. TYPE: San Spe Volle a vicinity of Julian, paca 1898 There UC). Plants 3-5(-6) dm tall, densely silvery tomen- wi hort mucro; petioles ovate to o lanceolat te, ا‎ 5.5 с п pia mes 7.2 -30) cm lone. wi -30 е peduncles 2-6(-9) cm long; bracts ovate to nearly lanceolate, 6-11 mm long, 3.5-6 mm wide. Calyx 5-8(-9) mm mm long, 5-8 mm wide at the limb, lobes 3-5 mm longs re ~+ mucronate; о п. hl vules 5-7. Pods 2- 2m long. 0.6-0.7 em wie Seeds 1-2 (fully matured not seen Distribution and ecology (Fig. 1)- v рожан of San Diego Co., Ca Morti grassy oodlands, 1000- 1500 m; flowering April-June. ени ѕресітепѕ examined. US wp rn a: San Diego Co., 1.7 mi . NW of Julian 7, Bac igalupi et al. 5799 (ЈЕР, КМ. WTU Jalan, оң 1932 (CAS POM, U е 1946, reed 3576 (RM, CAS. edi Cu aca мл 28 1899, Най 1205 US. Y, UC} v ie of Santa Ysabel, Henshaw 189 ER Volume 81, Number 4 1994 1 723 Taxonomy of Thermopsis , 25 June a Mts. 1924, Munz sig He HENE РОМ); Pine Hills, 1 Jun (G 1917, Spencer 504 (GH, NY, M, UC); inne. bed of Cedar Creek, 27 Mar. 1934, Wieslander 447 (UC). Isely (1981) treated this taxon as a localized B within his concept of gts о. macro- e with his circumscription. Larisey (1940) inexplieabiy omitted e taxon from her 2. Th RS divaricarpa n, Bot. Gaz. 25: 275. 1898. Thermopsis rho Mi var. балот! (Nelson) Icy, "etiani 30: 4 . TYPE: U.S.A. Wyoming: ei $^ fa pia s ranch, Big Laramie River, 8 Aug. E A. Nelson 3903 тавад ү ы , RM; isolectotypes, GH, NY [dated 7 Re 1897], RM). хы я Е. Greene, Pittonia 4: 138. 1900. S.A. Colorado: Saguache Co., below Маг He il Ta , Greene 1896 нера selected “hei ND; isolectotype, ND). wo different collections of Thermopsis divar- n y A.N sims кй: Ise’ d (1978) designated Nelso: 3903 as va e lo ape i he should have BEES this as a 1 In їй y рг э е for Thermopsis pinetorum, Greene cited оа таде by ' himse x in кш below Marshall Pass," 4 Sep. 1 d y C. F. Baker at Ps Pinos (which is near mien in La Plata Co . The on na. ге former is selected а as the SON fe taxa concerned. Plants delicate, 3-6(-9) dm tall, glabrate or Puberulent; stems slender, erect, des ed, mod- erately branched, Pine from or rhi hick woody root- mes; branches ascen ici at 20—35°, p to rod zigzagged. equa elliptic to narrow m long, 0.5 5 s, conspicuously n veined beneath; р a А 2. 5 cm long; stipules Laus lliptic, 2-3.5(-4) cm long, 0.7-3 cm wide, assai чо Pads. acute. Racemes 7-15 cm long, with 5— 45 flowers, maliy in ely of 2-4; peduncles 3 mm long, librata to اا‎ то E de- i ovate to lanceolate, 7-14 mm long, 3-5 mm wide. Flowers 2.5-2.9 cm long; calyx 8-11 mm long, 6-8 mm wide at the limb, lobes 2-5 mm s as inei as or shorter dun tube; wing and (ERES uo wn-black, .5m wi pe pubescent Se Seeds E ird 14 ellipti with a Distribution and (Fig. 2). South-cen- ming (mainly Albany Co.) southeastward Mountains spruce forests, уко бе streams or in moist open areas, 2000-3 000%; j decim (April-)May- June(-July). Judy vitu E examined. S.A. orado: Alamosa Co. e entrance of Great Sand Dos Nationa! МО 8 Мау 19 (TENN). Boulder Co., PR 2r Aug 1884, thills s go Creek, 11 July 1942, gy ee 83 ass, d №. ‚ 30 Ма LO); Sangre de Cristo Cre ek, a ly & Vreeland о аи ‚ RM). р wy. 67, Bear Lake Сат 0, Ste, 42675 (NY, p ud No. W of Walsenburg, 1 J une 1941 R. A. Nelson 4595 (RM). Jackson E uld eius bs 23 Sep. 1944, Ginter 891 (RM). n Co., Ver Canyon, 20 June 19 GH, MO, POM, rimer Co., Estes ; M ME m tg 2581 (COLO, 4 dag WS). Las mi. N of Stonewall, 26 968, Turner TEN). Nea Co., 13 Aug. 1981, ens 1961 724 Annals of the Missouri Botanical Garden T | 25° Fios L ie e ~ _ ri | | l д ij 1 e 77359 | THN m i $ o `i | K [i | VUL ! 300 mi. FIGURE 2. Distribution of Thermopsis divaricarpa. (COLO). Park see kay Junction to Wandcrest Park, 2 R S); ; Turner 5776 (TE idge лз nzie 256 (MO N lear River 30 Jun 19: 0., S side of L ESTE Lar, 71 (GH, NY, RM); Sand Creek, 2 June 1900, N оК ы; (CAS, COLO, GH, MO, NY, POM, RM, UC, US); Mandel, 24 July 1903, haw 8910 (COLO, GH, MO, NY, UC); Kigi Kur: 4 i y 1917, Nelson Yea MO, RM N, UC); Mo. Mt. region, 6 July 1943, Porter men cas, GH, ү М US, WTU); Crow Creek, Larami нен 1985, Rolin 977 (GH, M 0. NY, WS); La in near L , 16 June 1935, Williams 2214 “CH, 5, U US} ne: к Creek, Laramie Mts., 20 June 19 Williams 2236 (GH, MO, UC, WS). d Co., adden, in Medicine Bow Mts., 13 June 1972, рей МО). peus d Isely (1981) treated this taxon within his broa concept of which it Mer Nev ertheless, we hive retained T. ence fo A ECT AE hom, T. NM Pee. ite of s fa е seri do ub appear to intergrade as do the varieties in the T: puc eS e Volume 81, Number 4 1994 Chen et al. Taxonomy of Thermopsis occurs mostly at lower oe in deeper and dri carpa at и. elevations T. served at the present HU with we такоо + + 1 11 ё. 2. E Бы о ils, Т. іп mesophytic pet ни А ү ЫБ i i е ng is true, E that occasional intermediates a Thermo vito divaricarpa and T. rhom- bifolia occur, leri from E Pontes hybridization events. Such “hybridized” in Albany Co ounty, 5 1, A ы. 2 3 41 L Wyoming, of T. a arenosa, which «реа to combine some the features of Т. divaricarpa in the genetic bud orado line, T. montana and T. divaricarpa occur in close nroximit ^T 1 E. г J Р 7 - у? © exchange between the two. The variants from this wrinkles in the broad distributional fabric т Т. e E divaricarpa, and T. do not m: c or their taxonomic subm mergence. If it were аза en many distinct species emend the Жу he genus of ees ould be merged, for oc e or even rampa ана is the ам species of me taxa occur toge (under tween the closely eis d divaricarpa, an ¥ Т. montana is apparently much i Pr lofejd "isla eqq) species Baptisia leucophaea, spha ocarpa, and B. leucantha (Alston & Turner, 1963). * hi 18 эйе ende Мн, that r divaricarpa m T. rhombifolia and a. So far as is known Шш ми ї that extensive likely. t these various hy- 4 molecular Кайа з. Regar ardless е believe that the taxonomic BORÊ i hist unos na, but we ments of 7T. m а var. montana or bifolia In addition. h f ll of T. к-га E from northeastern Wyoming that ү be misidentified elements of ‘arly typical mbifolia Isely icum КОК Thermopsis pinetorum as a POD и E ола var. montana but bo as not ted a уе. shows that the name apparently was typified by mixed collections; the lectotype belongs to his variety divaricarpa. Nelson, in his original description of 7. divari- carpa, iunge о one ей of the fruit- a еп re and y interm ably a hybrid or introgressed individual between T. rhombifolia and T. divaricarpa. 3. Thermopsis exerum Wed ex aid & A. Gray) M. Curtis, Amer. J. S 81. 1843. or [азий ae ex satt ray, Fl. N. Amer. 1: 387. 1840. Ther- mopsis BA var. Fenja ти ех Е геу & А Сгау) Isely 469. TYPE: U.S.A. "Мо nh epe on Co., s e Mountain (“Catawba of Nuttall; т е ‚ 1967), Мына ins (PE NUR m possible bip s, GH, NY). remains a problem with the correct author ri ed by Wilbur (1963) who contended that the correct citation should ex А. Gra y (p ublished in 1848). Isely pow ac- cepted this asinifolia dede ex x Torrey & А. Gray (1840) (beh, 1981). We believe this to be the preferred ation because Torrey x Gray (1930) an " by him in n their бона of T. frazinifolia. Con- trary to Wilbur’s observation, the Nuttall name was not rejected by T orrey & Gray “The ey merely oted that the — had not fallen under their кч ир) and that it was “uncertain whether it H 726 Annals of th Missouri baud Garden 300 бада Ld J 400 (oH VENTES AS) Ii RJ UA A CAL TALLY Xe гаа FIGURE 3. Distribution of Thermopsis fraxinifolia. fraxinifoli ij tf f the spe-‏ ل cies described bova: " We do not view ‘this asa‏ rejection of the name, but rather a form of abey-‏ ance.‏ Plants delicate, 5—10 dm tall, glabrate to sparse- robe stems slender, erect, clustered, well- branched, arising а m а single woody rootstock; branches бал at 45—80°, strongly zigzagged. Leaflets pane t 5-8 cm long, 2-3.5 cm wide, acute to acu e, glabrous to very sparsely pu- berulent, зате veins 9—12 pairs, very conspic- beneath; petioles 1.6-3 cm long; istent, lanceolate to nar- 0. .8 cm wide, terminal o а 12-25 ст ву with 7—25 flowers, scattered; pe duncles 2.5—5.5 ong; pedicels 7—17 mm lone: glabrate; ан, lanceolate, 8-12 т long, 1-3 mm wide. Flowers 6-1. 9 cm ын calyx 7-9 mm BENS 4—5 mm wide at the Ё than tube eliptica to asymmetrically oblong-elliptic; keel bese nt; ovules (6- )12- 16. Pods straight, regu uito divergent, (3—)5.5—7.5 cm long, 0.3-0.5 wide, inconspicuously shea Seats (5- mi 15, brown, reniform with a very short rostrum, 4 8 mm long, 2.5 mm wide. Chromosome number, 2n = 18. Distribution and ecology (Fig. 3). gus an Moun (RM, TENN E Co., near Stanley Сар, 2, Coi ile et al. 2844 x X). 7 (NY). M 7 i. S of the Georgia Hwy. 368 bridge over Savannah Rire? xcd m, 7 Tun е 1979, coeds 863 AE near bridge o a Hwy ^n NCU). Ra Jun нал гузар & Gaile 4 Rabun Bald, 1-4 June 1906, House 2266 (US) "aui Falls, ene 1893 F. N er P^ 0 10 July 1908, Howell 390 (MO, К А mbe Co., Black Mountain, 12 Aug. 1905, Bima erbarium we (F, G M e ) A vicinity o of Kraus 1925 РО Sn کک ا ی‎ 7 d. M Table Rock Mt., аи ug. 1890, Heller А i POM, UC). Henderson Co., Flat Rock, Gibbes 1854 Volume 81, Number 4 1994 Chen et al. 727 Taxonomy of Thermopsis 184 Macon Co., N of the ete а state a n North Carolina Muy: 28, x June 1975, Wood & Boufford 1389 (A); 2.1 mi. N of the Georgia state {чм on North Caroli п North Carolina Route 28, une g uffor 16075 (MO). Polk Co., "m Mt., Townsend 1897 (US) Bn o: hitewater River, just above Upper Falls, 7 Jun 951, Godfrey 51298 aoa Yan ~ т Ж М. Mitchell, laid енн ме М); E o Mitchell, exactly ipis o fandi ге June 1970, Tuner 6070 (ARIZ ges $e MO, NY, TEX). South Carolina: Abbeville Co., 3 mi of Calhoun , near аЬ River, 13 Мау 1957, Radford 22768 (NCU, UC). Greenville Co., Ca ead, Sm 1881 (F, GH, MO, NY, US). Oconee Co., Boynton 1889 (CAS). Pic Co., 0.7 mi. о wy. 178 on Sas safras Mt. Road, 30 June 1970, Turner 6075 (ARIZ, CAS, COLO, F, GH, MO, NCU, TU) Tennessee: Blount Co., Chilhowee Mountain, km W of Hwy. 73 on Foothills Parkway, 650 m, 26 May 1982, Boufford & Spongberg 22881 (A. МО, NCU, А ). Стеепе аа ті. $ Camp Creek School, just N of Hor i , 10 June mountain slope betwee dae желш fre Creek on Green Mt., 30 July e Shake & Shap 7439 pare rs milton C Mt., Can 869 nroe Co., vine Road, 5.8 r 3 «in чайда 23 May 1980, rdum 47 (TENN): ‘Wolf Ridge, 25 June 1983, Z. E. & J. B. Murrell 336 (TENN). Cultivated: Massachusetts, Botanical Garden of Cam bridge, Gray s.n. (NY). armenie, with Ther- p T. fraxi- ecies is partially саа The ermopsis eat al de- 848b), T. ap e arvard Botanical Gar- time. ей pecimens of both e the m d of the his treatment of The ed Sta Ra by Gray (1848b), Wilbur (1963), and der. et n (1968), Thermopsis mollis is mostly à piedmont Species, distributed along the middle and western portions of North Carolina and adja- cent states, occasionally extending into the moun- tains, Meu its range abuts that of T. bios s Indeed, w not know of any specific sites where the two i been pollected together, п nor do we of near contact (or elsewhere) that might justify ir reco species, 1 by Isely (1981). His distribution maps of both taxa | are у оки, sod those at are difficult t he did not cite ns documenting the distributions ios ая (1981) did map a number of specimens from oe Тешен ist appear to be lou Un E a. somewnere anomalo between dns mollis and T. frasinifolia and were designated “T. mollis var.?” o belie this region € the intermediates have ШЕ habit ү habitat preference of Т. frasinifolie, enel г. Р and Т. fraxinifolia have been со ue untain, Hamilton с ве of the putative hybrid intermediates (e.g., En rchill 1911, N Canby 1869, GH, MO, NCU, NY, PH, US). More detailed field investigations may show that TEX a hý Whitten from Monroe Co., Tenn ee (I5, TENN), which seem to o reveal stabilized Some of the characters in at least a qum ok die lants are unique within the American taxa of Мур pen For example, the Bloun ty imens cited above have short, quan gynophores, much а ecies ип in some of how- ever, it is “more sensible to include such plants ih a d г ia dm than to accord these ос it sta 4. Thermopsis СРЕ Howell, Erythea 1 1893. TYPE: U.S.A. Oregon gresa i 5 “Mountains ne sed Ido, ” 5 Jul y 1887, T. J. Howell 669 ies, ORE; isotypes, CAS, GH) D venosa Eastw., Bull. Torrey Bot. Clu ё = 198. 1905. Thermopsis gracilis var. venosa (Ea Jepson, Man. Fl. Pl. Calif. 515. 1925. тузан Annals of the Missouri Botanical Garden montana var. venosa (Eastw.) Jepson, Fl. Calif. 2 245. 1936. Thermopsis macrophylla var. venosa (Eastw.) Isely, Brittoni а 30: 4 469. 1978. TYPE: U.S.A. : Sha Tri of Bohemi. vere as Counties, йш 600 2 Мау 1926, J. R. Patterson 9959 (lectotype, й here, ОКЕ). In his protologue for Therm ‚ How à did not mention a specific type but rather noted in “Mountains of southwes н from the ирек. of the Wills amet, to to the рй ont a number of collections. Actually, aba served as the voucher for Thermopsis grocilis. i A. Vorobik, by annotation, selected this as the holotype: A purist might have called this a lecto- type. Plants robust, 3-8 dm tall, € to sparsely villous; stems slender, erect, sin gle or cluste id well-branched, arising from tock rhizomes, branch 5) cm og stipules р per- asymm n ovate-elliptic, 2.5-3.5 ст Re 3 cm wide ) ong, glabrous to sparsely ie bracts semipersistent, ovate, 6-8 mm 3-5 mm wide. Flowers 2-2 10 mm ы 7-8 mm wide at hei Seb uch shorter than Distribution and ecology (Figs. 1, 4). North- western California and adjacent Or regon, meso- phytic lower a montane evergreen forests, 100-1200 m; pica (April-)May-Jun Representative specimens examined ifornia: Del Norte Со. ‚ Siskiyou lowlands n 22 Apr. 1939, Meola 69 (OSC, UC); N side P Middle Fork of Smith River on Old Gasquet Toll Road, 1 June AS Бнын, 125° 120° / 4 [- 459 1 j NAT b O25 o У к ) | 25° ; Ка we y эс | SRG Ес 200 mi. FIGURE 4. Distribution of Thermopsis gracilis. i Parks & 0 11227 сн GH, UC, ee 2 asquet on French Hill Road, 1 Jun di Tier 5801 (CAS `H, NY, TEX) Humboldt ., Frenc i Tracy 12955 ee 8 74 (MO). Josephine Co., summit o F, Mts. especially N ge 3 July 1902, "Cusick F4 GH C Ws); W 1 UC, US, WS, WTU); Oni A Steward гоз CAS MO, NCU 5 Sexton Mt., 9 Apr. 1934, Poop 10209 chs, (i J POM, WY). Lane Co., Spencer’s Butte, Volume 81, Number 4 1994 Chen et al. 729 Taxonomy of Thermopsis ugene, 5 May 1946, Baker 2791 (ARIZ, CAS, MO, NY, OSC, WTU); near Cottage Grove, Lorane dre 1931, Henderson 13574 (PH). Lincoln Co., Alsea mi 1939 ( Polk Co., 2 mi. W of Grand Mts., Lam Кеф. Brehm 1967 This species is readily distinguished from the oF ра c Thermo nta Sth a names as s ess, we feel that Т. gracilis is = from г. macrophylla, as discussed in det dé чат 5. Thermopsis copi. Hook. & Arn., Bot. Be iun ым . 1840. TYPE: U.S.A. Californi Ba м Co.?, spring of 1833?, p Sade Ynez Mt., near Santa Barbara, Douglas s.n. (holotype, K; photo of holotype, POM, UC; isotype, GH). Ther, mopsis macrophylla var. agnina J. Howell, Leafl W. Bot. 8: 158. Santa Bar C ann Ci 1 mi. non "ii Peak. ca. 1060 m . M. Pollard 1955, plan sheets (lecto otype, , selected here, cer CAS [2 sheets plant A], US[3 sheets). Many workers have localit ty of Thermo pondered over the t unty, California, from where he made "on ce visits to Santa Barbara Mission nta C = f this ma y (Dav s 1907, CA 5; Ф , RM, U ‚ US) reveals xs t robust fo otherwise typic ermopsis californica. How- al Th ell also discussed the possible ы. locality of Ther- иан. eenaphydia: in :соцоесіоп with his e scri of T. agnina, sugges a eim docality for T «anis ie near Aro in San Benito County. But these, too, are merely large-leaved forms 2 otherwise typical californica. Type s ens o rophylla match very closely those of T. ma p ч папа ш had Howell examined ma- we Deneve he not have desacibed the latter. No doubt cach f the fact that T. macrophylla is a relatively rare localized en has con the existence psis robusta, a localized ave recognized The та а as distinct from T. califor: neat both Т. macrophylla and Т. robusta as the ame Plants robust, 12-23 dm tall, tomentose; stems thick, erect, solitary or few, moderately branched, isi: branches ascending at ы. де, amplexicaul to cordate, acuminate. Racemes 25-60 cm long, with peer ovate to жкен 8-10 mm long, 3- e. Flow 1.7- 2.2 cm long; calyx 7-9 e limb, lobes 3-4 “Шо with a short rostrum, 4-5 m » 2.5- 3.0 mm wide. b = 18. Distribution and isis We 1). Known only from the Santa Yne z Mountains, pe Barbara ., California, dis it is locally co the west Je of Santa Ynez Peak near icd summit, in sandy granitic soils, 1000—1400 m; flowering May- g LU Specimens examined. U.S.A. California: Los An geles Co., Claremont, weed on the Ranc Botanic Garden grounds, "apparently introduced," 9 М 1991, Ross 5047 ( г Ba ., S slo} pere Peak, 11 2, Breedlove ] Ma a 5 (CAS, POM); San Ag be Peak, Tark 1977 Ан, Santa Ynez Peak, са. 4000 ft., Pollard 1956 (CAS, Annals of the Missouri Botanical Garden TEX); in chaparral W of Santa Ynez Peak, 17 Oct. 1945, Smith 1620 (POM); just W of Santa Yne z Peak, 12 May 1957, 83 (POM); nak 5022 Bien San ta Ynez Mts., alon Cami no Cie lo Roa: intersection with Refugio Rat 4020 ft., 26 ob, d Junak 5020 (TEX). The relationship of mone Somin and its EERIE aroan, 4 is The early workers, m y S. Wat -— fem - This specimen (СН) is a fairly typical specimen of T. PIRR pres umably c ollected in th haps in the vicinit PS pes its flowering state and vestiture 6. Thermopsis seg go paneer: M. Curtis ex A. Gray, Mem Acad. ie: 3: 47. 1848. Patria то Michaux, Fl. Вог.- Amer. 1: 8: Baptisia n icti (Mi- — sf “Can N Amer. Pl. 1: 281. 1818. Car lini: Mecklenburg es Made without date, Michaux s.n. (holotype, P not seen; probable isotype, PH). yd +» vicini n маг Small, Bull. Torrey Bot. Club 25: 139. . Thermopsis rete Күй Small, Fl. S.E. Us. p 1331. 1903. тү А. Georgia: олй ersham Co., **northern slo; не nt Сг iff, eere A. M. Huger 1897 vitas NY; M Plants delicate, 3-6 dm tall, sparsely Maren pubescent; stems — аср ос br eading, "had or clus tere fro hed, arising in Eee br: acuminate, sparsely appressed pubescent especially on veins, ti g , lateral veins 7-9 pairs, very zen ca net- veined beneath; petioles 0.7-1.7 cm long; stipules caducous, narrowly el- i i 3-0 2 —17 flowers, scattered; pedun- cles 3-7 cm long; pedicels 4-6 mm long, villos- ulous; bracts aa lanceolate, T 13 mm long, —4 mm wide. Flowers 1 1 жес ae © 2- 3 mm ali shorter than tube; wings р К М $ 4 hl t pu 12- 16. Pods arcuate to pe di- 0 pe ie igs (Fig. 5). Mostly mid- G Carolina, western North Car z d qe 300-800 m; ^e (April-)May- Jun Ala- Representative specimens examined. U S.A. ma: De Kalb Co., Mentone, Loving 1899 (US). Geor- gia: Dade Co., 1 mi. W of Durham an Canyon, 12 Men 1953, Hardin 15933 (NCU, US); м 1952, Harper Mos on US). Н id 9 July Carolina: B unco: PH, R = June 1896, Biltmore 102. 5 (F, NY, PH). TENN, UC, US). Caswell Co, Milton, bra ү: Сиш Co., 3 mi. SSE о f Newton. 6822 (NC, NY); 1.25 mi. ide 9 M on 70, 3 May 1953, даз & Wood 6970 net, 5 Chatbam Co., 1i a Co. line on U.S. H ds e 24 Apr. pia ама 23825 (си, NCU Baar Mt 1 pes а а web Rad ord Es на 77 (N Lynn 1933 A Forsyth Co., e hallert 1940 (ОНУ 9 Ё Guilfor E Tot E PON Henderson s ., Flat Rock, Mem ger 1886 NCU ). Iredell Co., 3.3 mi. W of "Dogg Eos rth C a Hwy. 901, 5 he ie RY, 2221 (NCU). pee 4968 (F, be , Р Bi» S OM). Lincoln Co., Hunte: чыч GH, 28 Apr. 1957, Bell 6654 (NCU). Or ы s.n. (NY). Stokes Co., . 1961, usi 1 92 INCUY abo € 1306 ed Hass 1986 (NCU). Surry Co., Pilot Mt., Ashe 18 Volume 81, Number 4 1994 Chen et al. Taxonomy of Thermopsis А ci = jan Nen M з а ne ч А] [| Li [1| " 40 E @ 9 "rx CAS td Lj Gî Беа [ | Wa $ Hy 85° | ok aah DTU ERK ASA REOS Sater ed pease nce n PEDE ® LÀ» 78° FIGURE 5. Distribution of Thermopsis mollis. э, Pilot Mt., 5 May 1935, Oosting 3519 (NCU). EL. Co., Kittrell, Browne 1884 (NY); Kittrell’s Springs, axon 187 H). Wake Co., 5. i. NE of Holly г 9 20 Арг. 1938, а: Сг 1 ., Ridgeway, Sou Greenville th Carolin е Co., Rich Mt., 5.5 mi. , 16 967, Bozeman et al. 8862 (COLO, NCU, POM, num Rest, ca. 0.5 mi. W of West Village О ns Co., 12.5 mi. S of Rosman along U.S. TO. ‚ 1 May 1941, McVaugh 5649 (СН, UC). ennessee: Blount Co., Chilh Apr. 1939, Jennison 299 (TENN); Сар and Walland, H Ов с: 7 Мау 1964, Th 94 (TENN). Virginia: а йм Co., intersection of County Route 711 and Coun- оше 811, 11 Aug. 1985, Wright 469 (MO). Camp- bell Co., Sleepy Hollow Road, 23 May 1967, Freer 5597 (GH, NCU). Charlotte Co., 4 mi. S of Charlotte, Court House township, 16 Apr. 1 6 (NC ‚ MO, NY, US); bridge Co., Balcony Falls, 1 May 1947, Freer 1348 (GH). The relationship of this species to Thermopsis fraxinifolia is discussed under T. fraxinifolia. Fernald (1950) noted that at least one popula- tion of Thermopsis mollis from Essex Co., Mas- sachusetts, had I i 1 d 1 lized i the northeastern United States, although he ig- nored the more widely cultivated 7. villosa. We Essex Co (GH), pre- р sumably that referred to by Fernald, but it appears о be Т. villosa. 7. Thermopsis montana Nutt. ex Torrey & A. Gray, Fl. N. Amer. 1: 388. 1840. TYPE: see Thermopsis montana var. montana. Plants delicate, 2-8(-10) dm tall, glabrous to appressed pubescent or thinly villous; stems slen- der, erect, solitary or clustered, moderately or few- 732 Annals of th ми "ын Сагаеп branched, arising from a woody rootstock or rhi- zomes; branches ascending at 20—45*, weakly to moderately zigzagged. Leaflets narrowly elipti ic or obovate, de 5-8 cm кш 0.7-3(-5) cn or obtuse, sometim 2e sparsely appressed pubescent to дио, lat- ral v ll pairs, conspicuously net-veined long; pedicels 3.5-5 ong, ulous; bracts semipersistent, narrowly elliptic to widely ovate, mm } -6 mm wide. Flowers 1.6-2.2 cm long; calyx 9-11 mm long, 5-8 mm wide at the limb, lobes 3- D mm long, shorter d tube; Шр а ovary velutinous or densely appressed га ovules 10-16. Pods straight, erect, 4.5-6.5 с long, 0. i 0.6 cm wide, vill us or appresse ae uas es M AD ccs s de. me 8, 2.5 wide. 2 B a rostru ‚9—5 mm ognize two Ducem varieties under rec dus aa as follow: KEY TO VARIETIES OF THERMOPSIS MONTANA la. Plants less than 7 dm tall, pubescent; stems diua ten ce and few-branched, often arising hizomes; leaflets not conspicuously net- ut beneath; bracts lpi о widely ovate Her SSE RIOR ermopsis ntana var. montana lb. Plants up to 10 dm tall, gla тың ; stems cl y branched, pine from woody rootstock; leaflets conspi cuously es vind beneat o J elliptic .. —— NE Sidi montana var. ovata 7a. Thermopsis montana var. montana. r. montana (Nutt. ex Tor- rey & A. Gray) Isely, Brittonia 30: vie 197 TYPE: U.S.A. Wyoming: “Rocky Mountains” SA aig: Co.?], Nuttall 1834 быу. not seen; photo of holotype, POM, UK; rne PH; photo of isotype, POM). L8 Зон angustata E. Greene, Pl. Bake: . 1901 E: U.S.A. Nevada: Elko Co., Sar Valey “foot hills p the Ruby Mountains,” E. L. Greene 1896 (holotype, ND). Thermopsis иси. Е. Стеепе, РІ. Baker. 3: ds dg TYPE: July 1901, C. F. Baker 604 | i select e ND» Баура, GH [2 sheets], NY RM [2 od mae UC, US [2 sheets]). According to the ко distribution of Ther- a yoming possible that it was first collec Idaho in early July (cf. Graustein, 196 << = tall, ие pubescent to thin- s stiffly t, solitary or clustered, ulous. vus ome number, - Distribution and ecology (Fig. Vu NE ed States, from southwestern Montana b Arizona and New Mexico and occ to i i to Maguire 11886 ei pz 2618 (RM). Co: y Pass, 30 May 1 ick & у of жы Ee trail to re of th P July 1950, Weber 5894 (COLO). Delta Co. Mule y : 196 Dolores yn " 27 Aug. 1964. foe pe (TEX) Mon Со р Dolores Riv- Volume 81, Number 4 1994 Chen et al. 733 Taxonomy of Thermopsis er, Escalante Sector, Bye et al. 1978 (COLO). ys 0., 30 mi. SW of Montrose, " Aug g- AUS ths llin 978 (CAS, GH , NY, “ыр y Co, lements 1907 pal ny Blanco a i. Е of Mee ker 8 Aug. 1935 е& Pianos ен (МО). Park Co., Jefferson South iod EP 1942, Ewan 1 d Rio Grande Co ES nra rk of Rio еа s River, 27 Aug. 1964, Tuner 509. ). Saguache Co., Marshall р me July 1960, Bae be 60 (US). I Lav. a Hot Spr 1 along а cd 2 July acf H. & C. А eed 19580 (W: { lark Co. К. nas near Birch Creek. ыз О; NY: РИ, U). oak re divide between Marsh and Malad Valleys, Coulter s.n. (PH). Owyhee Co., 9 mi. uneau, 30 May 1968, eee = RM, TEX). Twin А Fal s Co., Twin Falls, M). Montana: н s an E & Armstead, 20 June 1920, E. B. . B. Payson 1734 (CAS, GH, MO, RM). Broadwater ie 4 mi. SE of Town send, 10 June 1960 нш) JES-345 (CAS, RM, WS). Deerl Fishtrap, ca of Anaconda, 20 odge Co. June 1944, Hite hcock 2. С ^ 73 i. GH, MO, NY, RM, UC, WS, WTU). Gallatin lgr. s (RM). 1933 (RM, WTU). Powell Co., de itch ph n. (RM). Silver Bow Co. er Moose Creek, ca 8 Ma ye Co., UT ^ Be uw )N n o Co., 8.5 mi. from San Antonito, 0.7 mi. N of cw vine 21 jn jsen "mer 5649 (CAS, NY, n Co. » Моро ar West Fork of the "23 Aug. E Meal 593 (MO, ND, NY, . Calf lfax Co., 1 mi. M oen IP 0. СОБЕ 11 Kin Jum 4 ( б . E of » Cueva, Stiteler es (PH). б Co. nier summit of pass between Hollywood Mescalero, 26 May 1934, Ferguson & Ottley 5223 (UC). Rio Arriba Co., Ortega Mts of Hopewell u е Pecos River, 9 June 1897, s v E. Ni Heller 3681 (MO, NY, US). Sandoval N of Jemez he 6 June 1974, Atwood SENT. gv Fe Co., near Santa Fe cc Area, just off New M a Route 475, 16 June 1964, Perdue 6047 y? Tao enasco, Santa Ba rbara C г a ene Marceline i (F). O Ar "^ T variety o ta]. M Jun e 1934, Siar 21734 OS, UC) U M 5 mi. < June 1979, " Neese & Wal 7 503 (NY). Garfield Co., t 7 mi. SE o ulder, Deer a 4 May AS Foster 3661 (NY, A Sd; nd Co. Ad of War r Ra Station, La Sal 8 ia 1 3, Мавиіг 313 (C; WTU). Iron one ca. 15 mi. of Ged ar City, just W of e entrance to Dixie sait днае 13 June 1969, Meyer 356 (UT). Millard Co., Simonsen’s Ranch, along irrigation stream, 16 June 1933, Maguire & Becraft 3932 (GH, UC). Morgan Co., flat area below dam by old gauging station, 17 May 1987, Peterson 14 (NY, UT). Piute Co., 5 mi. E ingston on Utah Hwy. 62, 3 May 1977, Foster 36 13 (NY, RM). Rich Co., W shore of Bear Lake, 0.7 mi. E of Pickleville, 5 July | 1 985, Thorne 3938 (GH, MO). $ alt Û ake gne Clements 1908 (F, GH). Sevier Co. - Bank. along stream, 8 May 1937, Hichcock 2047 бн, RM, UC, WS, pie Sa go нав, К а. 80 mi b, NW fl 40 Abajo 2 Jun 1961, Comune p: I ACH, AT NOU. POM, TEX, кб $15 an near Bear River, 30 Jun nc 1926, E B. & L. B. Ux ayson 4833 (MO, PH, POM, RM, UC, US); Uintah Hayden, EA Uintah River, Y "June 1933, qM 8044 (F, GH, N m а i Co. Heber City, along заан Мау 1968, jure 5 ay r 5794B (TEX). Кез 20., 9 mi. terprise at Aspen Park d= ب‎ с е qd 8: — o e دب‎ a > © da 55 5 33 a =~ 2, + = S5 ЕР 5 ® SE ч E... E BS Wee 10720 (GH, M 8 (NY). ec Co. = Uin a Mts., Henrys river Mab 26 June 1902, Goodding 1192 (CAS, F, GH, MO, NY, POM, | RM, UC). U , l mi. S of Lonetree, 27 July 1939, Rollins & gran "2903 (CAS, US). The ie or montana var. montana is wide- spread and highly variable but can usually be dis- hed from variety ovata by its smaller habit, narrower leaflets and brag: ts, and greater ер to spread Ьу rhi Oregon th 734 Annals of the Missouri Botanical Garden — 7 t ix dii 115° поо 105° M v “л ө Top о Г ° e > о ос б) 0 e 83 o S нЕ 95 1259 o о » aos rt | o А 5 ам if а E ^ 89 ә ° o oF o E o РА о тт | o L o [735° "es e o EY ai o o ; o " 200 mi A 2 ^ ed 8 EEE : 5 Е ircles). FIGURE 6. Distribution of Thermopsis montana: variety montana (open circles); variety ovata (closed cir taxa appear to intergrade. Isely (1981) recognized race" (Lemhi Co within th r. g with somewhat oodi leaflets, most of those spec- imens are fairly typical of T. montana var. mon- ta ps Arizona and nee of New Mexico (from below San Juan and R occurs a serie: } iar 1 broad- ranching r plants with leaves more or less like Thermopsis montana var. ovata. Unlike (1981), we do not find such plants in Colorado The strictly erect pods distinguish Ther mio: montana from T. gracilis and from the ы сїайу mgr T divaricarpa. Isely d re tana and T. divaricarpa as varieties 0 S ES zm TR T. rhombifolia. W И have found that Т. г hombifolia, T disi iP : 3 Volume 81, Number 4 1994 Chen et al. 735 Taxonomy of Thermopsis 7b. Thermopsis montana var. ovata (Rob- ins Robinson ex Piper, Contr. U.S. Natl. Herb. 23: 49. 1906. Thermopsis ovata (Robinson ex oe Rydberg, Bull. Torrey Bot. Club 40: 43.1913. Thermopsis rhombifolia var. ovata e selected hue. WS; dnce yide US, WS). Thermopsis xylorhiza ped ушуна өк Bot. Gaz. 52: 265. 1911. e^o ap viui 0., Falk's Store, 667 m 4 Ma (АУЕ Macbride c з>р ы Th isotypes, rae MO, NY, RM WS). Thermopsis macrophylla var. hitchcockii "ede Brittonia 30: 469. 1978. piti: . Wa E : Gra m arbor Co., афа umptulips O July 1931 EN. Thompson 7342 boe PWTU. ibd. GH, OSC, PH, POM, UC) Piper's collection number 1489 was qub à mixture of plants collected on 16 June 1893, 4 July 1893 (in fruit), and 21 June 1894 б {ача аз п: 18 assumed that all of the collections cause some of the labels (21 и 1894, NY) give the locality as orden streams woods." The published type loc n by Peck as “Cedar Mountain"; this loca ico was not on any of the шеге 1489 that we examined. Plants 5— = 10) dm tall, glabrate to sparsely ssed d y elliptic 1.5-3(-5) em wide, -— appressed P ent below, nspicuo ane о m long; bracts narrowly elliptic iot 1 Ps 2. 2 cm long. Pods appressed ан. = and ecology (Fig. 6). Columbia taries; flowering May-June Representative specimens examined. U.S.A. Idaho: Ada oer "Oda rs hi Boise, Moore's Creek Canyon, з er 16858 (NY). Adam: Jun Eo d mt. dots Hornet, 13 May 1915, Ma 39(R i. W of Lowm M). рок Со 5am wman, above Payette rds on between Bank wman, 31 May 1944, Hitc e & Muhli 8563 (NY, WS, WTU) Butte с Gin st 3386 ES Canyon Co., Falk's Store, 10 May RM, m MA WS, ). e Co., Clearw Forest, Lolo Creek, 16 е ті. W of Orofino, 1951, m 241 (WS). cots Rock, Elwood 51 (CAS, NY, POM, UG, WS, WTU); 7 mi. SE of с City, 14 June 1950, Jones 14 (CAS, GH, POM, WS, WTU); 2 mi. S of Grange 29 May 1944, Hitchcock & Muhlick ae (CAS, NY, RM, , WS, WTU). Latah Co., wooded hi 1964, Hitch ӨР 234 c о я 65 (СА Lewis Co., Missio n Creek, 30 May fuse 24, de ы үн 5334 (UC, WS, WTU). Nez Perce Co. Brown 6 (CAS, MO. PH : E well, 30 May 1936, Constance & Rollins 1621 (F, gH, NCU, MO, NY, UG ` ‚ WTU); near n 7 May 1896, A & E. G. Heller 3035 (CAS, alley por үн 13937 (CAS, МО, NY. PO NY, OSC, UC, WTU). Washington Co., S of Cou neil, mi. W of nt Peak, Gilkey 1947 ( OSC, пр 1871, -— e i ille, scia Mts., POM OSC, US); Silver ir ge Fal 19 along summit оаа betw: and Langdon Lake, 28 ae 1950, ‘Kruckeberg vw (CAS COLO, G Walup: i Cany n, 18 Aug 897, re 8724 ГА A Snake Riv: (GH, e, along Grande Ronde Rive Rattlesnake Gack 22 Apr. 1949, Cronquist 3709 y RAS POM, UC, US, WTU). Col ., Dayton, be- n hospital and анн Bre, " May krog aes 51 | (NY) Blue 0 July 1935, Constance * ak 1278 efg oad е. te Annals of the Missouri Botanical Garden Co., N of Aberdeen and 3] May 1987, ig Cn. n P pede along U.S. Hw WTU) open 40, „Thompson 14606 (CAS, MO, NY, PH, POM, RM, UC, US). Lewis Co., Сеш, Oxford Prairie, 12 June 1897, Lamb 11 ipse EI s NY, РН, UC, WS). W Walla, 24 May 1944, Hitchcock & Мишер a ores S, WTU); а Whitman Co., 1 mi. N of Colfax, 18 May 1935, Hitch co. атие1 2594 (OSC, RM); Kamiack, 15 June 1893, Moore 1489 ( U); Dusty, 3 mi. 8 US, W Colfax, 14 May 1922, Parker 355 (NY, OSC, POM, W 5). This De differs from Thermopsis montana var. montana in its relatively robust habit, mod- Mie beatae stems, and broader leaflets and brac Plants similar to Thermopsis montana var. ova- under T. Ош treatment і is like that within include his phyl T. montana var. ovata. We do not believe that the Paci oast pla crophylla v. mbifolia. istri- bution of the Pacific constat pee is ace to represent founder ү by periodic populations of T. cali Preis as sugge: rupted in Oregon by ад quite ү T. grac- ilis, which has diver rgen p ns we have e mpelled by morphological consid- erations to include both T. gracilis and T. cali- fornica also as part of the T. rhombifolia complex, h hological I laries | T. montana var. ovata and T. californica being at least as weak as those between T. Mauro i T. di- varicarpa or T. montan 8. Thermopsis rhombifolia (Nutt. ex Pursh) Richardson, Bot. App Rev. 4: 193. 1819. не he es ex e Raf., 5.92.1 BÀ E: U.S.A rth xs Me ercer Co., near Fort Ma erat Bradbury 1810 E se- lected here, PH B. Lambert herbarium). Thermopsis arenosa Nelson, Bot. Gaz. 25: 276. 1898. Thermopsis rhombifolia var. arenosa (Nelson) Lar- isey, Ann. i Bot er 251. 1940. TYPE: U.S.A. Wyoming: y Co., Laramie Hills, 16 June 1897, A. Melee 3182 (ho lotype, RM; iso- types, ND, RM, [2 sheets] UC, US). Thermopsis иаа Nelso n, Bull. Torrey Bot. Club : 239. 18 ii егар chee ia var. ап. illiams pios un among the rocks м the па те 25 July 1898, те" ors 4971 (ho- lotype, RM; Mena, NY). Larisey Did. designated a Nuttall collection pe of Thermopsis rhombifolia; how- g Cytisus rhombifolius Nutt. as a nomen nu Es from Fraser's Catalogue without citing а Nutt collection. Plants dwarf but robust, 1.2-3 dm tall, glabrate to appressed pubescent; stems slender, erect, ОГ ascending, х or clus f an arising from a woody roots oe сер at 25-50°, weakly zig Leaflets bov: veins 5-7 pair: етен beneath; petioles а elliptic widely e id wide, mucronate. Кареев ра 12 fl 1-4 cm long; pedice el- densely appressed дай: мин s deciduous, Volume 81, Number 4 1994 en et al. 737 Taxonomy of Thermopsis 300 mi. p 99 ө е = e Lm LM DA ٩ : can "T e A ч ° E PAN LÀ Ж b E. ww Le. Zi te ; TX MA mE $ / ( à 1 j^ S~ uw TE - ( У 450 е. 4 Y m Hie © / рот D ode à E EE AAT Ute A Neb «7 zu AD. Jee LS 73 jdn Дао de pe. а, ө | FIGURE 7. Distribution of Thermopsis rhombifolia. liptic to А 5-9 mm Flowers 1.9-2.1 cm long; pod m ж" iC 25 Bu Ls 6 mm wide at the limb, lobes (3 ed 5 mm long, ^ long as or shorter than м i asymmetri ; keel metrically екй: li у е “к nest ° to appressed puberulent; ovules 12-14. Pods arcuate с Distribution ну соку (Fig. I bris Gre. central Alberta and Sas аена to northea: *rnmost New Мех, largely confined to relatively the lower canyons of the front range of the ky Mountains, 1000- yt m, to 3000 m along roadsides; flowering May-Jun Re «fumi ide specimens examined. CANADA. = berta: ca. 26 km NE of Edmonton, 1 km SW of For Saskatchewan, 20 June 1959, Shetler & Stone 3034 (US); m W of Cochrane, 1 Jul PH Here the sand 58(GH _NY).S askatchewan: Cypress Hills Shed tal l Park, 1 Ju ly 1947, eel 4200 (MO, стан 1 һое С р Мау 1 a 1 сз SW TU, NCU, TEX). D Franktown, og e 1937, Beetle Tarr, NY TO rt Co. bey 3 ag ape em ns ker Las Ro эне 3 5664 Го, ТЕХ). Lic Co., 738 Annals of th Missouri ни Garden Lincoln, Чез: m йг ot 23A ag Moffat ue ain, 8.2 mi Iu. 40, 6.5 adis mi. NNE nf Dioni 11 June ur Holmgren P al. 5162 (CAS, NY, US, WTU). Park Co., $ m, ae Vi 8 TE 5774 (TEX). Weld Co. P die Bates, ; July 1906, Dodds 2111 (COLO). Mo mtana: Big visitor center at Yellowtail 883, "dori ). Custe iels Co. 0 Teton Co., 22 of Chouteau, Finley x Collins, Jones 1904 ( Toole Co., Sweetgrass, Recup 1918 (NY). ‚ Fork eos 26 May 1919, Rams. en & TU). Wibaux Co., Hodg- ONTU, RM, TEX, WTU). Yellow- mi. NW of Billings on Lavina Road, 3 June © ewe = = 7 а Hwys. 2 *T Ors 25 Ma 961, Ste evens Misso iver Valley, 8 mi 1941. те 905 NCU UC, hts US), Morton Co., Glen Ullin, May EN Holzinger 5 (GH). Mountrail Co., 12 July 1960. S evens & Moir 2289 (NCU, UC). Slope Co. 5 Atkinson [oss (F, NCU, RM, UC). Stark Co., Dickinson n, Е 949 (US). Ward Co., Haigh 1900 (NY) 4 ton, Stevens 1918 (Е). Oklahoma Cimarron C * Mesa. Ud. А a : e. 566 (US). бон. aM ta: Butt of Belle Fourche, 23 May 1970, Stephens 38357 NY ^ e Li ed of Blocktail Gulch, 12 Apr. 1910, М 4020 (Е, NY). Fall River Со., Ochich, 8 Арг 1, Nelson 2724 , RM). Jackson Ce. Cedar Ра: Ч Tu 1913, Over 6113 (COLO). Harding С 4A 1927, Over IE [E DE Rabbit usi) 1 елү 1912, ester 553 eb e Co., e r Lead, а ‚ Palme = ig US). Mea e Co. Mord mi. NW of Rapid бу 9 iiy 19 1947, тҮ Ч Рита 44 (RM). Pennington eak, 8 July 1927, Hayward ve 1961 0 N М. Perk mmon , Halse 1969 (050). 5 f Dutch John, 7 he 1979, past ud (oes SE Co., ca. 14 ai NW of Vernal along Tridell-Dry Fork Eu М 1 Iie p A nd "iis oN iXX Wyo elephone Canyon, 30 Ma Ont 10 1027 (COLO, MO, MONTU, RM, ws, desi bell Co., ca. 8 air mi. SE of Reno Junction at Little e 1918, Hartman & Dueholm se Co., near the head of La Prele Creek, 27 June 0 1950, eds aue (GAS, FOL. GH, MO, NY, RM, TEX, WTU). C NW of Hulett, 6 June 1935, dh a6 9010 "мо, MONTU, NY, RM, UC, W: ont Co. mi W of Lande: along ah n "28. 2 July 198 3, Evert 5282 (NY, RM). Goshen Co. be Wheatland and Veteran, 19 June 1953, Porter 6235 (RM, WTU). Hot Springs Co., 20 of Thermopolis along Wyo Co., Bighorn Mts. W of Buffalo 1958, C. L. & M. WTU). Laramie Co., 30 and Interstate Hw Dun 1091 (MO). N owder River, 5 July 1979, Dueholm uc RM) Nokes Co., 22. air mi. NW 1 air mi. SW of Lance Cree 2 210% iod Nelson 1591 (RM). Platte Co. o 1979, Рат 17542 NY, RM) a ming Hwy 789, 15 July L 85 (R M. nie Со, Г of Ten ne 251 hy 190 бойле 353 (Е, GH, MO, NY, RM, US). W ei NW of Osage, 22 May 1984, Marriot 62 a vp three varieties prd ed to (1981). Altho sely’s parameters of pce privi ifolia to cl the long-re ed spe I. t i divaricarpa, nich t, we ecognition a arrowe! Volume 81, Number 4 1994 Chen et al. 739 Taxonomy of Thermopsis either 4 Ru cepa t * 2, аш but the oc- between at least T : rhombifolia and T. divaricar- Albany County, Wyoming. The osa ìs probably asta on such hybrid variants Fae. discussion under T. divaricarpa). 9. Thermopsis robusta Howell, Erythea 1: 109. E TYPE: a A. pata eae ds st Range, undary," T. Howel 1884 МЫН - ORE; I F, NY [2 sheets], US, WS, W Some of the apparent isotypes are labeled **Coast H Judge from the specimens we have examine Plants robust, 8-18 dm tall, densely tomentose; stems thick, erect, clustered, well-branched, arisin; from а w ootstock; branches ascen t 30—50°, s ongly zigza. ps кашг Cr to ong, 2.5-6.5 widely iic or ovate, 6.5 cm wi ute to а гар: pus eig veins 9-12 uously net-veined nsely iib, 5 rown, reniform bs rostrum, 3.5-4 m A ong, 2 mm wide. Distribution and кыш Fig. 1). Known only from northwestern California, Humboldt and Sis- kiyou cos., чүү ri yk re 1500 m; flow- ermg May-Jun California: Hum Ores, v ч ick 5 9. . fro on Bar m ty Road, 6 July ea Tillett 620 (POM) E of orbel, Bald Mt., on trail to Blohn’s cabin, June 1951, Vollmer & Beane 146 (CAS, POM, WTU); near Orleans, 19 May 1931, Wieslander 7 (RM, UC). tid. E 11 mi. N of Somes Bar, shady forest near Camp T s by 1954, Нис ga a 20223 (NY, WTU); 9. ba mi, 9 > Ron r Camp Road, 1 y 1950, Hopman 3 (UC); pes t Hill, 1220 m, 18 July 1950, Tracy M e om ect Hill, 20 July 1950, ا‎ 3 (POM, U TU) n near State Creek, 12 ga ae Pu (JEPS). County unknown, E a бана Vasey 1876 (GH). Thermopsis каван is a ea robust en- demic of EE m. Californi Beca ause of it habit large leafl it is supe rficially pieni to T. саайа Both Larisey (1940) and Isely (1981) included 7. ro- busta within their concept of T. l macrophylla. We believe it is more closely related to T. deca to judge from its divergent pods an distributio: 10. Шекер villosa Т Е е & В. . Schubert, Rhodora 50: 90: ОЛУ enh Walter, n. Си. uda iro TYPE: U.S.A. North C not seen). Чур ughly discussed by Fern it transfer of the о from Sophora to rmopsis УР ке о iana M. Curtis, Amer. J. ва 44: 80. 1843 E: U.S.A. North Carolina: ood Co. "ins нн montanis," М. A. Curtis 1839 (holotype, СН; possible rel PH, NY). ‚ Plants delicate, 6-18 dm tall, glabrate cad tary ately zigzagged. 1.8-4.4 cm wide, acute to obtuse, a pes. con- PN 0)cm sparsely \ villous below, lateral veins spicuously net long; stipules posees lip to ovate, i 5-4.5 o 2.4 c e, oblique to cuneate, cm acute to cemes арра long, with —50 flowers, scattered or in whorls of 2-3; pe- Fines 6-13 ong; pedicels 2-3 mm long, pedicels 2 ous; bracts oe widely ovate to тезе) obovate, 5-7 mm 8) mm wide. Flowers 1.7-1.9 cm 6.5 mm wide at the limb, lobes a idis = long, 12-16. Чаш night to i arcuate, erect, 4- ү 5 cm 6 cm wide, densely tomentose aie Seeds 7-12, olive brown, elliptic wit 740 Annals of the Missouri Botanical Garden 90° ase B0? Am us 659 + ае dli on. 9*7. 59 i л... E 3: П : 40? о E + [ S | Distributio: ll thought URE 8. to ine introduced or enia н от rostrum, 3-3.5 mm dee 1.8-2 mm wide. Chro- mosome number, 2 18. Distribution nd ecology (Fig. 8). eastern Tennessee, western A N ative to ations in open ar- ish mostly 1000- О m; flowering ibo. ~June(—July). Representative keen frames. Ala- b Co. ma: DeKalb Rabun Co., Tallulah ock С eef Spáü lding 582 (GH). Ma ا‎ Prince Georges Co., Hermann 18409 (RM). Massachusetts: Hampshire Co., railroad RC -way, Ahles е. ыа‏ اکر پیا Middlesex Co., Framin Pike exit t‏ 27 June 15, 7 Ahles 80370 Ди» E tate Hwy. 93, un JS Ir £j lersey: Sussex Co., Edwards 1971 ga Urian i Moldenke et = 30109 (A ARIZ, pa Au кыл уегу Со., June 1958, Ahi Ра SS mos 19E, iN o Gabe? les & Duke 43574 а МСО) баат N arolina Hwy. 6 et al. 3314 (NCU). Haywood Co. Creek, 5 5 June 1958, Ahles E Duke 42242 2 (NCU) АРУ derson Co. e Logan, a alls, 6 Wilbur 71 42 ( "UO. Jackson с nea 0 W of July 1940, Hood 109 (MO); Soco Gap, ca. 3 mi. Hwy. 19, 1 July 1954, Sargent 6851 i more near SE of Sassafras‏ ,. ا WTU) 13320. F ‘GH, MO, NCU,NY, US; je Volume 81, Number 4 1994 Chen et al. Taxonomy of Thermopsis Gap, 18 June 1969, Leonard & Russ 2512 (COLO, 3 NCU, POM, TENN, UC, WTU); 6.7 mi. W of Wayah Valley Ranch, Arro , 29 June 1970, Turner TU). Madison Co., Ahle ,W . Swain EE along Mone wy. ca Knob » Row 6 Jun une 1956, Bell 3200 Е ая aa e Co IMS 276 near 3 Sep. , Freeman 57819 (NCU); along motor road in Pink Beds Rippe (RM). Pennsyl McKean Co., Fogg & Whe: ry 9818 (PH). T e: Blount Co., Cades Cove, near Crib Gap, Hoss 1954 (TENN). Cumberland Co., of a une 1970, Kral 39741 (GH, МО, NY); тие e near ene ES Gattinger 1879 (GH, MO, NY, US). Polk Co., ca. 1.5 km SE of summit ig Frog Mt., 14 July 1983, n E. & J. B. Murrell 389 (TENN). Vermont: Windham Co., Charette 3345 (MO), and 3349 (NEBC). seem Roanoke Co., Uttal 6228 (NCU). Thermopsis villosa is d ied e, stiffly erect species of the southern achian ridges and upper valleys. SAFER (1970) Ten it en үй caroliniana) as the only m o Caro- linian; i differs markedly from that species in i ias more slender, strict habit and лен radic cally naturalized papye of this п occur from Vi astward to Maine. Fernald 0250 гое his treatment of Thermopsis for the деси United States, nor did ha comme ent on Шш pariy natur; ud ыы (Fig. 5) FH +; Gray (18485) з st tated ei Шү the plant in ae с Garden (where it is perfect] T by Mr. Buck- o do till retained in the occasional garden or has persisted in pue long vacated. One ee tion is known from Hawaii (Oahu, **Mauk of Kawaiiki dies poor ad 30 A d : Degener 17363, GH). W Nashville, first made ALSTON, R. E. & B. L. Turner. 1963. Natural h r. J. Bot. 50: 159-1 BALANDRIN, M. F., E. F. RoBBINS & A. D. KINGHORN. 19 Alkaloid distribution i in some species of the ermo Biochem. Syst. & Ecol. 10(4): 3 ‘ot (Adan ) (RU h. P Bissy, Е. je 1981 425 P. H. Raven (ultor). A vances in diss Systematics. Ron Botanic LA m ga W. H. & S. Watson. 1880. Polypetalae. In: Botany of Californias ird 1. Welch, Bigelow, Cam- bridge, Massach BRowN, R. 1811 Р. T : W. T. Aiton (editor), Hortus Vol. 3. Longman, Hurst, Rees, CHEN, C. J., X. Y. Zou & Y. M. Yuan. 1992. Cyto- logical haha on the tribe Thermopsideae (Fabaceae) I. Report un И ен of eleven species of four genera. Cathaya 4 : 103- 19 . TURN Systematic e lupine а alkaloids ref- erence С ‘Bat a (Leguminosae). Evolution 21: -51 Serological жанр" amo; Sophoreae, adora Geni de ceae). Bot. J. Linn. Soc. 94: 421-432. FEOLI-CHIAPELLA. 1984. e and di- nisteae, Webbia 38: 105-122. SR Series novae generis Sean R. Br. Зан Systematic ae Plantarum 7: 213- DEMENT W. A. & T. J. Masry. 1975. Biological i im- a plications of flavonoid chemistry in Baptisia and ermo, Biochem. Syst. & Ecol. 3: 9 n is. ARIS 0 Chemiotaxinomie des а ées—Genisté Bull. Soc. Bot. France 1965: P102. ies iet ai A (editor). Hug gerente me veis TS of Flowering Plants. i. USSR, "m M. T 1950. car s pem 8th jS 6. ican Book Company, New York ошт, ae . 1975. In: А. Love (editor), IOPB Chro- Number Reports L. Taxon 24: ric des 678. GRAUSTEIN, i E. 007, ода Nuttall, Naturalist, 1808-1841. Harvard Univ. Pre: ess, — м “Chloris idu Americana. Mem. Amer. hs Arts n.s. 3(1): 1-56. ——. 8b. А Man ual of эя dace of the North- ern United States. Poe ston. L. 1988. International Code of Bo- br Ge ttinger in the Harpeth hills in 1875 (MO, ad PS) may also be locally е рор- LITERATURE CITED ue. L. 1944. Fabaceae. Pp. 480-627 in An Il- ustrated Flora of the Pacific States, Washington, Oregon, and Са кой, Vol. 2. Stanford Univ. Press, Stanford, California onoid ао іп i si 8: 14 HoWELL, J. T. 7087. e Ба nez Thermopsis. Геай. T Lag 7(6): 158-1 Se ELY, D. New variet combinations cae acies Thermopsis P. рае Tem gen = onia 30: 466-4 OS of ы United States, III. ET Papilionoideae: . Tribes Sophoreae, Poda- lyriea d Lotea em. New York Bot. Gard. 25(3Y ү E LARISEY, M. M. 1940. A revision of the North American 742 Annals of th Missouri icti Garden species of the genus Thermopsis. Ann. Missouri Bot. Gard. 27: 245-258. LEDINGHAM, G. F. 1957. Chromosome numbers of some Saskatchewan Leguminosae with particular reference to Astragalus and Oxytropis. Canad. J. Bot. 35: 657-666. Linx, Н. F. 1821. Icones Plantarum Selectarum. Pub- . Species Plantarum, ed. 1. Holmiae Fi ейп, .mpesis Laurentii Salvii). A. & р п: A. Love (editor), IOPB "Como me > Neier тен LXXV. Taxon 31: "368. ME J & J. So some Mon en ys EN cobs: Photon 4 NUTTALL, T. 1818. The Genera of North American Plants. Printed for the author by D. Heart, Phila- 1976. Genisteae (Adans.) ге and related tribes ras iso) Pp. 143-368 in V. a Heywood (e or), Botanical Systematics, Vol. l.A ademic е ew York k. PRoBATOVA, N. S. . SOKOLOVSKAYA. 1981. Ka- riologicheskoe issledovanie sosudistykh rastenij pe- trovov Dal’ go sapovednika. Sh. e бузуш ек озїосһподо 92 КАРЕ , Н. E. Anes & C. Малы, a the Vascular Flora of tly n olinas. Univ. ss, Chapel Hill, North Pei ges ааа Ts (819. T "Pini . Chim. Nat. Arts. 8 SALATINO, . GOTTLIEB. 1980. Quinolizidine alkaloids as systematic markers of E e Papilionoideae. Biochem. S ES ud neha ie phical evo- of ч УРЫ ta in Papilionoideae. Pl. Sys Evel, 143: 167-174. leui ud Pp. 30-36 in V. ditor), eat the USSR, Vol. 11. Pro ogram for Scientific SHISHKIN, B. im L. Kom at (e ditor), IOPB Chro- mosome Number Reports X (ee Texon 16: 445- 461. TORREY, L 1857. oad. Perl ке nda 859. gue of poe! ‘ur. In: W Bor (editor), В p US und. Surv., ла WAOE Nicholson "Peiner Washin; 1840. ora of North nos Explo . GRAY iley & Putnam, New York Plant chemosystematics and phy- hem. 14: 189-213. Legume Systematics. Royal Bota . 1982. Review of: Hake the United at Ш. Subfa mily Papilionoideae: Tribes Sophoreae, Podalyrieae, Loteae. Syst. Bot. 350. E N per plants of North Carolina. Technical d No. 151. North оша Agricultural Experiment Statisti Raleigh, North olina. WILLDENOW, C. L. 1799. Species Plantarum. Berlin ome G. C. Nauk). YEH, with а АЗА & Е. MAEKAWA. 1986. Chro- in the Legum inosae. "e Rep. Res. EST s 3: Dru WS ee y G. 1967. Казању of some plants, cul- tivated in the Arctic-Alpine Bot anical Garden. Pp. 139- d 49i nN. A. Arorin itor P Paii um Zonam Polarem Transportatio. II. Lenin, THE BOTANICAL WORKS OF PHILIPPI, FATHER AND SON, IN CHILE! сыс М. Taylor? and lica Murioz-Schick? ABSTRACT „ Rodulfo Amando Philippi (1808-1904) was a leading figure ing the same period and worked closely wi . Bot Philippis Phi i (1838-1910) was a botanist dur orientation to “their works an i е principal field trips are summarized. ir m tributed to ie in Chilean natural history, s j^ dh s son . We nt an arie wi dely i in бше; their oe type: Aparte ited primarily t SGO, but кү шелш ыл of type material is mde дь ted erbar by sparse label data int many piane kso: remain to be selected | for many names. Altho oug h both ai published wid ely i in a variety in Chilean journals. : Rodulfo Amando Philippi (1808-1904; Phil.; ш; ” Stafleu & Cowan, 1983) w as Е vd i H 3 of the country’s s natural richness, the а е е of Chile’ оп of territory "i development. As the leading field biologists of Е ge Copiapó to é. The Philippis published more than 56 lip c s on the natural history of Chile (R.A. Phili i: са de articles; giten 9 of Chilean a = нуз їп Latin, Spanish, aid German. Thee зіч eontology, avifauna entomology, exploration, anthropolo logy, min ammals, mol- & = E е the field and the hé rbarium escribed 3720 new species of TENE from Chile "bs tha: M; Q Tins (Muñoz-Schick, 1973). dori кесиши пера ims 3 w S and Gunckel, 1939), most in publications compiled by his father. He was the first head of the Sección Botánica of he Museo Малаша, "ire h e curate [Ch ent plants (F. Philippi, Ren A he followed his bird as professor of bota he University of Chil те i “ir or of the ва Botanico at the Quinta N In sont 0, ће large number of works pe lished byt them под their work. Heise we concentrate f the combined works, to orient contempor diffi pis’ work an information. kem most comm sparse docum Philippis' very Pide collaboration, to the point that in later years even their individual handwriting became difficult to distinguis A BIOGRAPHICAL SKETCH sienne же igi have been published, most r the other Philippi but docu- e lives if bê both by necessity (Barros- 1904; Gotschlich, 1904, 1910; Gunckel, ы $ © viewers for comments and Селин е tion. pes cial thanks are due to Clodomiro Marticor ena, ipae ón, Museo Nacional de Hist Ишт б 3166 paiak чле Hears and Oscar Leé * Misso! tanical Garden, P.O. Box 299, St. jad | de Con ncepción, for generous help an ria Natural, for the photographs. U. 5А : Louis in Botánica, Museo Nacional de бнт Natural, Casilla 75, Ann Micenirt Bor. GARD. 81: 743-748. 1994. Annals of the Missouri Botanical Garden 1939; шет qum 1991). We present here a brief udis th lives, with an emphasis on botanical acil R. A. Phili ppi was born just outsid [ош = and Berlin. He eventually obtained a medical de- gree but did not practice. He traveled widely in Europe as a young man, notably in Italy. Federico was born a during his father’ E last stay in Каре ich 1910). R. A. Philippi Pies settled into eaching position in the newly gaben а y- tec га ol in Kassel, where he 5 He p ublished ишер: articles Hoi n par- molluscs, and the geology of шеп Italy. He was er known in the contemporary scientific orks were recognized with medals from чы апа Italian rulars. However, he eventually found him t d lat K lí = lh А | Һу СЕХ апа 1аїе б а cold December night; Barros- Arana, made his way ош to m German ONT S near Valdivia. H in 1852, and then together with his brother ac- Legs end Fundo ("Estate") San Juan along the Rio usi arrived i in Chile in late 1853 with the е Quinta Normal de Agri- cultura, where the Jardin Botánico was later es- tablished. = A. ТЫ lived in Santiago most of the year, th y near Valdi San Juan estate until his first wife’s death in 1871, when he traveled to рт to study bota 74 t any and zoology. He returned in to take over his retiring father's teaching duties, and in 1889 be- came head of the newly Higgs Sección Bo- tánica of the Museo Nacional. In 1883, Federico also succeeded his father in the directorship of the Jardin Botánico, and in 1897 he resigned from rope, father and son lived together in Santiago, eventually on the grounds of the Jardín Botánico, and екан closely together. COLLECTING EXPEDITIONS Both Philippis were active is workers through- out their lives, even | thou oth suffered health meningitis mained sind B malformed je dod pos for the rest of his Their ose expen and collecting efforts are outlined in Tal ble Aa aile d list of the names ing latitude and lon- e Philipp coveries has been disc cussed m Gunckel pe Some trips were undertaken by order of ge gov- lished ne of them. These trips brought discovery of many? . А. Philippi" 2 1852 trip oping it as well as its lack of precious metals. n his 1864 trip through the Cordillera de la Costa near Valdivia, Federico Philippi ited the dillera once where he found a mo ж cove a forest composed of millions ias elictual unities that were **nat- ing, ed alerce trees (Fitzr оу cupres (Molina) J alor sprig also re pockets of Magellanic plant commun! disjunct by hundreds of =н тут from their Volume 81, Number 4 1994 Taylor & Mufoz-Schick Botanical Works of Philippi ТАВ Principal travels of Rodulfo Amando Phi- lippi = Е ederico ae in Chile. Pies Jocalits- Pizarro (1960), MulosSchick (1973), and Munoz & Pri- na (19 87). Dates Travels 1852 R. A. Philippi; Río Bueno south es Valdivia, Volcán Osorno. ia Llanquihue, Volcán Cal- co [X Ке uel 1853 (Nov.)-1854 К. ^ " Philippi: by boat from Val- (Feb.) ess o Coquim land 1854-1856 1858 (Dec.) 1859 (Jan.- Mar.) 1860 1860 (Jan.) 1860 (late) 1862 1863 1864 1864 (Oct.) mbo, by Copiapó, Atacama, Eee (sca Desert) [V to . Philippi; various sites around Santiago and the adja cent Cordiller: [Metropolitan Region] R. A. Philippi; Colonias del Sur [X Regio R. A. Philippi Colonias del Sur [X Region R. A. Philippi; Catemu [V Re gion], Laguna Ranco [X Re- gion R. A. & F. Philippi; een LI Dag Andes, Lago Ranco x Re gion R. A. & F. Philippi, Quebrada de San Ramón [Metropolitan Region R. A. Philippi; Nuevo Volcán de Chillán [VIII Region] Pod gl Nuevo Мыш ~ ЛП Re ed A. Philippi; Juan Fernández Islands [V Regi p е оп] 2 sta Hueicolla, Слай Pelada [X Region] . A. Philippi; Cauquenes [VII Region ; = сие теа eses [VI Region] pue Laguna ^s Budi, De- sembocadura (mouth) of Río Tm n [IX Region R.A Г E: E: H n] ppi; San Juan [X Re- pan R. A. & F. Philippi; Arauco, Malleco, Cautin, Cor жы а de Nahuelbuta, Tres Piedr. [VIII to IX Regions] TABLE 1. Continued. Dates Travels 1878 (early) 1878 1879 (early) 1879 (early) 1880 (Jan.) 1880 1882 1883 (Jan.) 1883 (Mar.) 1883 (Oct.) 1884 1884 1885 (Jan.-Mar.) 1885 (Sept.) 1885 1886 1887 (Apr.) Philippi; Termas de Chillán Region R. A. Philippi; Coquimbo Prov- ce [IV Region]; Matanza nd Cahuil in Colchagua Pi. ince [VI Region] . Philippi; Descabezado (head- waters) of Río Maule [VII Re- = v R. A. Philippi; Tomé, Quiriquina, Salto del AM EE Re imi m n] F. P kee Cucao and Ancud: [X Region = "y E. SS 59. — 1 3 < — e к) оч - R. A. Phili agg Arauco pees в оп] К.А рі; бош [V Re- > gion . Philippi; eti area in- cluding Concón and mouth of i, Carlos Rahmer (Sub- director v Vise aro of the useo one бсо Туш Re- R. j* - Ры San Juan [X Re- R. ys Pili Algarrobo [Metro- politan Region] Annals of the Missouri Botanical Garden TABLE 1. Continued. Dates Travels 1887 (Nov.) R. A. Philippi; Araucania [IX буг ion 1888 (Jan.) гои Alfalfal [Metropol- itan Region] 1889 R.A - Philippi; Вапоз Colina [Мег орошап Керїоп] 1890 R. A. Philippi; Concepción to Arauco [V egion] 1891 R. A. Philippi; Victoria, Los An geles, Cur. hue, uco, Valdivia [VIII to X Sa cn 1891 (Oct.) OA. MM Quilpué [V В 1892 к.А A. Philip Banos Chillan VIII Re 1893 (Sept.) F. Philippi, Filiberto Ger (Head, Seccién б б. Museo Nacional); Fray Jorge [IV Region] 1893 R. A. Philippi; Concepcién, Cura- nilahue [VIII п 1894 F. Philippi; Yerbas Buenas, Ca huil, Matanza: Region 1896 (Mar.) R. A. Philippi; Constitución [VII Region] 1896-1908 F. Philippi; various trips to Valdi- via, Fundo San Juan, Cautín, Constitución [VII to x Regio ns] ural" range in the south. The Philippis’ joint trip in 1877 to the Cordillera de Nahuelbuta explored territory recentl d from the control of the war with Peru and Bolivia. ederico on ad several unusual plant comi- munities acquired durin F caria araucana (Molina) K. Koch (Ar- t north of t 10 у Jorge, i miarid region. The гину: extensive otanica collecting and knowl- distribution and provided the basis for inc of our current knowledge of the Chilean flora. THE PHILIPPIS’ SPECIMENS Mos | f the Philippis’ Chilean material, includ tye is depen the Museo Nacional de His toria Natural in Santiago (SGO). This — was variously collected and annotated by on th Philippis ‘Che of Chilean plants sent to the esra by others, Va dne Pearce, Volckmann, Solis de Ovando, andbeck, Fonck, Geisse, Juliet, Bor ipa S, Germain and King, were also in VAs с 1991). All of these other d are n: йж we collections, due to mea атр iple material that wi collec d Federi herba . This personal collection con- arium at home tained duplicate of specimens deposited in the Museo Nacional. It was acquired vi the Museo Nacional i in 1911, after F e of ed labels with black border and the We “Her. at BA, BAF, BM F i SI, and W (Stafleu & (ойло. 1983), and are also a at CORD (Taylor, pers. obs.). hilippis’ handwriting: have been for Schick, Figure 2. ч handwriting was similar, and Є у aged. Although the ' of the specimen is з frequently not noted, e rticularly use by refer- e subsequent head of th was Carlos (Karl) Reiche, who prepared a revised Flora de Chile (1894-1911). Reiche was € "^ the Philippis' specimens, and his comm is work is useful for ae lectotypes- xs ever, some of Federico's collections now 1n in the Museo Nacional were purchased after jagi s de- parture from Chile ayi were not se y Reiche's feet г> tive; his annotation suall established by Mufioz-Pizarro when the speci in the herbarium were , a process ! began in 1942 bua fioz-Pizar- o, 1960). 14 met of Philipp is some times difficult because of the sick n |. specimens, the cleat inclusion of several T heton on o t with no indication of whic plant was acco аат citation of more than опе specimen Volume 81, Number 4 1994 T Botanical Works of Philippi aylor & Munoz-Schick 2. FDR Chee ALO oe X Fano Ley yt Le Ое; ея ads a Car А er: Re. Angeu nuu шийи ИШ HERBARIO DEL MUSEO NACIONAL DE HISTORIA NATURAL, CHILE. Material montado bajo los auspicios de la Oficina del Coordinador de Asuntos САДУ форм Ministerio de pns У Compania Manufacti wrera de Papeles y Cart Handwriting samples: н data b "ila poer with annotation by R A. Philippi dein portion). h PA Uf, Qi + [4 * » SYS. (nd. Жл Ke ИТЕ HERBARIUM FRIDERICI PHILIPPI... sv... deo: HERBARIO «S MUSEO gy og ере DE HISTORIA NATURAL, CHIL! FIGURE 2. Handwriting samples. From top to bottom annotation by Federico нв label by R. А Philip ppi, Philippi’ u d original species n . The known syntypes isthe SGO collecticn shave boen catala ged along with n references, collection мъ NEY an y Munoz-Pizarro (1960) with "kes by Muior- Schick (1973). Neither of these authors attempted to select lec- К Mare ee eee HERBARIO DEL MUSEO NACIONAL DE HISTORIA NATURAL, CHILE. Material montado bajo los avspicios de la Oficina dei Coordinador de Asuntos Inicramert. Pu din iris de VE ME y Compania Мапијаг Pape Cart des y н. Е 1 FIGURE 3. g sample: Label by Karl Reiche. totypes, but concentrated on асир the ma- terial to facilitate ne by sp et ecies dece by the Phi- ered. Photographic negatives of both SGO's type RR 11: QT PER E £ Chl pent Ие deposited in several European herbaria аге av vail- ae on ри from POUR or st т íi ip. meia t be borne bs investigators ойо, т pom THE PHILIPPIS’ PUBLICATIONS Several ee of works Pore by the val in Chi ve been pre- more than once. Th ap to enlarge their list of gpg 5% rather to disseminate their work t wider audi also to help struggling acids by prov siii arti- cles us here on the Philippis’ publications with siio to the flora of Chile, probably a slight +e f 4h 1 fR A Phil : n grecs majority PF most of the publications of Federico. Their publi- 748 Annals of the Missouri Botanical Garden cations can E be divided into се ас- counts, written by опе ог more frequently both Philippis, and аи ions of new dim written almost exclusively by R. A oe R. A. Philippi’s descriptions w species were published principally in odi series, ceti - did n not рызы e ifte these. On a very f p , in different journals. The first of a series of new nted in sectio ee e other sections continue as in one publication. These species are usually based on specimens collected by the Philippis. d some cases a set of consecutive on specimens from one col- ч more new de la ri ia de Chile, with descriptions i in Latin an st series was senes were present ing Gay (1845-1854), which generally follows the sys- em of de Candolle's Prodromus. Fr on, the descriptions were prepared h ppis. cation е second series is detailed by Stafleu & Cowan (1983); all of the series are de- tailed и (199 It w п during the Гир time to rous reprints (t separatas’ j of individual s ar- which wa alè In some cases, the title may also have ares changed slightly. These reprints are commonly encountered in libraries, particularly in euh as separate шше ud in re nd Mri 2н z orks isi tation ide Mun БУРГЕР pas nl pee: (1992) as an aid to de- ermining the ог € publication date, site, and eee of these articles mmon nomenclatural problems seen in treat- hilippi n the inad ed names че were кыныр оп шере but ELT page number, based on the кү rathe r than the original publication; and the names attributed to published by later кай most commonly Reiche. LITERATURE CITED Barros-ARANA, D. 1904. El Doctor Rodulfo Amando hilippi. Su Vida i i Sus Obras. Imprenta Có de Chil Santia : Gay, C. 1845-1854. a: Fi Politi Bota sid Pad 8 "velt ., 1 atlas. 904. Biografi ia del Dr. Rodulfo Aman- do Pii (зов: 1904). арга ‘Central J. Lam- pert, Val CARTES 0. ida i obras de Hook Philippi. s. Nac. Hist. Nat. 1(1): 3 rei н 1939. La labor botani ie p Federico : 13-36. Bibliografía Taxonómica de las Plan tas Уаш d Chile. Monogr. Syst. Bot. Mis- Кр. и Museo Na- [o 1830-19 моа de Educación Püblica, dni: de ed tecas, Аг- useos, Museo Nacional de Historia Nat- ural, Santiago, Chile. Las Especies de Plantas ilii en el Siglo XIX. Ediciones ipei e n a documentación Кы porch Publ. Ocas Complemento de "Las Es- Philippi en el 128: 5-69. 100 s de Museo Nacional de pue Nadu Bol. Mus. Nac. Hist. Nat. Chile 42: 181 erico Philippi en su viaje a Tarapacá. Noticiario МЕИ Mus. Nac. Hist. Nat. Chile 313: PHILIPPI, F. Ey dcos talogus plantarum vascula t A nsium adhuc descriptarum. Anal. Univ. viii, 49-422. [Also published separately. viii, эт P STAFLEU, & R. S. Cowan. 1983. Тай Кыш Vol. IV: P-Sak. W. Junk, The Hagu UNA EVALUACION DEL CONOCIMIENTO FLORISTICO DE MEXICO! Victoria Sosa? y Patricia Dávila? RESUMEN En este trabajo se evalúa el México со herbarios, y estudios floristicos. даь eaten nee que con sus cin d 3 Do de pets ун 71 кеге 133 70% de Dieciséis fl d la superficie del país. E stas pra as, lista al 55% del total de las iridis para Méxi DEUM TUR los (3 5 floristicos y do: оп como áreas Soria s de explor acion botanic a: (1) о. Se pie жешл deciduos de la Vertiente del Pacífico; (2) la Sierra Madre 8 astizales. Grupo ў cu a Depresión del B. Norte con una FM bee en el país, como Rubiaceae, requieren ain estudios taxonómicos. ABSTRACT 151 ENS | This + £ M. е paper and une I n Statistics reveal that the count ry has reached a collection index of 117, ms approximat Sixteen floras (3 concluded a п collections and 71 herbaria. herbaria ely 2.3 nd 13 in process) cover 70% of the national territory. (9) th Species included in floristic studies represent 55% of the total reporte ed for Me xico. Active programs of botanical “Sierra Man del Sur” апі "Depresión del Balsas"; е агеав of the North, where dry а vegetation is present. still required México es un ig con una alta riqueza floristica. extensa superficie territorial, a su diversidad geográfica, " a su ubicación, ya que зе encuentra entr o biogeográfico Ne ártico ya el | Neotropical оон, 1978). B: nümero d MA en en alrededor « de 25,000 apdownkin: 1999) 1 le “чё сото Chiapas, Oaxaca, y Veracruz. Los tipos 1 0 total de especies) fe dowski, 1993). Rz (1991) estimé que aproximadamente el 10% d 1 коз: 35 T 105 gener OS y de México son endémicos, apunta ando que el e demismo es más pronunciado en matorrales xe- rófilos y en pastiz izales. El mayor número de especies de plantas en o en casi чү las floras om age ез, sa . Las especies de pteridofitas sólo parue a 5% d del se pr la r con 1100 ы de las cuales casi 20% son endémicas (Riba, 1993). Las gimnosper- mas con арго oximadamente "160 Ee consti- paoia al género Pinus (Styles, Ааа Entre ias mas div on: Neon , Leguminosae (17 a& Del- gado, 1993), neon he (1200 е es si Hag- Б ` Agradecemos a т Rzedowski у Flora de México 1 Apr LA | proporcionada; а суы en las ы ҮЙ 2 * Flora de Veracruz, Instituto de Ecologia, A.C., Apdo. Postal 63, Isis de Biología, Universidad Nacional Autónoma de México, Méx L. Cabrera su revisión crítica ^ manuscrito; a сонни јо Naci ld a M. iones; al Consejo Nacional de Ciencia y "Tecnologie su apoyo económico (0068. N9106 a V. 91000 Xalapa, Veracruz, Méx Apdo. Postal 70-233, 014510 México, DF., ANN. Missouri Вот. GARD. 81: 749-757. 1994. Annals of the Missouri Botanical Garden TABLA 1. Nú 1974, Holmgren et al. А 1 ү k M. í h : MT] j à E Hal , 1981, 1990; Consejo Nacional de la Flora de México, inf. inédita, 1993). 1974 1981 1990 1993 Colecciones 566,780 1,021,713 2,107,543 2,284,693 Herbari 18 30 41 (70) ral sater & Salazar, 1991), Gramineae (1 226 especies; Hollis, 1978; Bravo-Hollis & San chez -Mejorada, 1991a, b), y Rubia (510 sedo wski, 1993 ). TE суна. del presente БШ ез evaluar el con base en un análisis del nümero de: (1) colec- ciones botánicas; (2) a arios; Fs 2) estudios fo: rísticos s concluídos o n des: е > s. sá infor ee se ntean RR à ais k jeny ies en | Méx ic COLECCIONES BOTANICAS El total я colecciones botanicas en México as- ciende a casi 2.3 millones de ена Su in- cremento a sido notable en las ültimas décadas; 981 casi se duplicaron con qmod a las йырды унйн я рага 1974, у volvieron a мынын рага isti үш 1). El пйшего де también alto y casi iguala al de los herbarios me- хїса сїй asciende a alrededor de 1,800,0 ejemplares, los cuale an albergados en herbarios europeos (B, BM, BR, C, FI, G, HAL, K, LE, M, MA, P, W)y x Ru АСЕ (A, AMES, ARIZ, BH, CAS, F, GH, MO, MICH, MSC, NY, PH, RSA, TEX, VT, UC, WIS, ot TABLA 2. wr] ejemplos de índices de densidad de bbc en Ме 9. (Et numero de писта fup ht id. кофеби de las Floras de las distintas regiones. des par tir Indice Superficie Colec- de co- Estado km? ciones lección pels. oa Norte 70,113 21,000 29 Dur. 119,648 12,600 10 Pike ed 80,137 197,350 246 Méxic 21,461 181,2 844 digo 27,900 ^ 20,000 72 Jaxac 95,354 50,000 52 Tehuacá cán-Cuicatlán 10,00 6,000 00 Veracr 72,815 250,000 343 Yucatán 23,379 37,380 159 Una medida que se ha adoptado para determinar imiento floristico de una zona es un indice ^ P | 1 ж Bet, len БЕ L del кш de (сара depositados € en en her- 1977, Aah rat ot (1989) ha — un indice 00 como un minimo adecuado del co- nocimiento eee México alcanza un indice de Eds el cual mue ec sue minimo. Sin emba argo, + по significa pedi. boténicamente Por ae el estado de Ve ne un indice nsidad de pu un menor per id d pioneer P/ be Pro floristicos, | rentemente son bajos. A pesar de = pon s Durango o Baja California posean рне ЫП grandes áreas ocupadas "pe iia о de vegetación, de matorral xeróflo, con unà 1 stos casos un indice de coleccién pobre no significa necesaria- p E la representación floristic а no ge ade- gunos cua nsideran pir Casos s c t d d E] 1 1 + anjero ۴ ejemplo, el herbario de la riens de Californi 2 Berkeley (UC) alberga una buena coleccion й а Ре souls de Baja California. Del mism нба. ч вечи de la Universidad de Texas en Austm ne una b cada de co- ) co Ni poda is Desierto ibi ii ense (Rzedowski, 1976). HERBARIOS MEXICANOS n la b gs el número de herbarios en Méx- . Su mas £r el Her Nacional (MEXU )ye her "Instituto " Politécnico (ENCB); las ing de estos dos herbarios suman de un barios de México, con base еп el Index Herbariorum de Holmgren et al. (1990): obra re on un as А bn yii xx rai Nacional de la Flora ^u Мако, "1993, y su ride aparece en el apendice T (sus а acrónimos serán de Index Herbariorum de 1990 y TaBLA 3. Herbar La información del nümero de sus xci ашар dot deri ir ell ни te: bee’ e terisco*. de la po de airy numer: Region Norte Region Oeste Región Centro Baja California Norte BCMEX CMMEX Baja California Sur HCIB** Sinaloa Univ. Sinaloa* Nayarit Univ. Nayarit* Aguascalientes CIAN HUAA INEGI Jalisco Normal Jalisco* GUADA IBUG Agric. ape ai Cie Ки Jalisc ZEA* San Luis Potosí Zonas Aridas* SLPM Michoacán EBUM IEB CIMI ** Guerrero niv. Guerrero* 3000 Tlaxcala Univ. Tlaxcala* Hidalgo Univ. Hidalgo* Querétaro Univ. Querétaro* Рие тт Puebla* CODAGEM Cuautitlan* IZTA Univ. Chapingo* XOLO . Chapingo* Distrito Federal A MIZ Univ. 6 Тагар niv. Yucatán* зе (ЕМ СІСҮ) Quintana Roo CIQR** 5000 4000 200,000 30,000 10,000 17,000 5327 3780 2000 6000 5500 26,380 15,000 v66l p JequunN “{8 9unjoA оохәүү әр 021511014 ojuaeiuiouo? еплеа * esos LSZ 752 Annals of the Missouri Botanical Garden TABLA 4. Floras regionales concluídas у en desarrollo en México. Especies Nümero incluídas total de en tratamientos Flora especies taxonómicos Referencia Baja Californi 2705 2705 Wiggins, 1980 ipai bei 2634 2634 Wiggins, 1964 naloa 5000 E Vega, 1990 wem 4300 = González et al., 1990 Nayarit 4500 = Téllez & Flores, 1990 Novogaliciana ? 3523 dir 1983, 1984, 1985, 1987, 1989, 1992 Bajío y regiones adyacentes 5000 Т Rzedowski & Rzedowski (editores) 1991- Valle de México 2071 2071 e & Rzedowski, 1979, 1985, 1989, 1990 Veracruz 7690 1059 Gomez-Pompa bee Alok 1991 Sosa (editor), Tehuacan-Cuicatlan 3000 — Davila et al., Guerrero 6500 15 Di 1990 López-Ferrari, 1989 Oaxaca 10,000 — Dávila et al., 1993 Chiapas 8000 506 Breedlove, 1981 Smith, m Etnoflora Yucatanense 1936 15 die et al., o & Ciau- ti 1992 Jalisco 7000 = А. кыў сот. Mesoamericana ? (3,243) Davidse et al., "1994 En esta zona а han fandado varios herbarios re- (Fig. 1). La mayoría de las floras en desarrollo nte de cientem requieren aún del estudio de más del 50% de los dición de Index Н, iorum de Hol axa distribuidos en sus zonas. Floras como las de et al. (1990) (Tabla 3). De la región oeste, A Sinaloa, Nayarit, Durango. Oaxaca aun mente el herbario del Instituto de Botánica de la no han iniciado la publicación de tratamientos flo- Universidad de Guadalajara (IBUG) alberga más rísticos. Al de reciente inicio como las de 0,000 e a zona sur d el 0 uicatlan у del Bajio у Regiones Ad numero de herbarios y colecciones, aunque el ter- Masc cubren z zonas importantes por кл numero rbario más grande del pais, el del y sad aene er he i reos bri se erae localizado en esta conocidas s foristicamente. ‚ Colima, Cam- Lo taxonómicos que 1n: pe er yG j n por completo de her- y arüp os Шш НД еп México (Tabla 5) ak barios. omo los listados floristicos de áreas des 0 oui botánicos (Tabla 6) son otra fuente de in- EsruDios FLORISTICOS EN MEXICO formación florística. Si se toma el cálculo [D las primeras décadas de este si ; vador d RE ámicas de : «рева estudios floristicos рага Мёх d асаа. йв у si se suman ee et жаз. como monografias ados, canes meis los ih pe ndley 01920- 1926, LIS Rarer ури, 5 ien estudiada o listadas, eli- han sido concluídas: las del oa ierto de "Santen minando las especies comparti (Wiggins, 1964), Baja California (Wiggins, 1980) Valle de México (Rzedowski & Rzedowski, 1979, Discusion 1985, 1990), y trece floras regionales están en ; México ha incrementado notablemente SU in- process (Tabla 4). end 16 floras concluídas o en — fraestructura botánica y sus colecciones en com- el 70% de la pens de Ме exic paración con otros paises de Latinoamérica, como El noreste del pais no cuenta nales por ejemplo Costa Rica (Toledo & Sosa, 1993). Volume 81, Number 4 Sosa & Davila 753 1994 Conocimiento Floristico de México 10 100° 30° id i. 1 3 4 7 420° n 12 Ficu iar AS Bajío y берен adyacentes; 8, Valle 12, Oaxaca; 13, Chiapas; 14, Etnoflora Yucatanense; 15, Asimismo, para alcanzar un indice de colección considerada en n algün es studio Horitico, х ayes de las peo. estimadas qu el país aün no han sido list ada as ni estudiadas taxonómicamente; (3) stán арав ente T iradas gunas areas по e botánicamente; y (4) algunos grupos de plantas Floras de México: 1, Baja California; 2, bee aeree 3, Sinaloa; 4, Durango; 5, Nayarit; 6, de Méxi о; 9, Veracruz; 10, Tel Talise co; 16, Месе 11, Guerrero; „2 1 m ON para su e M taxo onómic Si metros ‘ales como bajos s EM y поо, y las ad tro Ver dd Pacífico, 8, son areas prioritarias de recolección. baas zo n sarrollos turísticos y contienen una alta diversidad TABLA 5. Estudi Cinca did i: Au nd an АА ы Nümero de Especies Grupo de plantas especies estudiadas Referencia Cactaceae 821 821 Brevo Holl, 1978; Bravo-Holis & Sánchez- ian rada, 1991a ^ aprox. 2026 — Turne som (or pace Gramineae 1226 538 fiiis 1983, 1987b, 1991 Malvaceae 372 372 Fryxell, 1988 Orchidaceae 1200 100 Hágsater & Salazar (editores), 1991 Palmae 100 ev 0 { san) Рн (Оаха. 690 690 Mickel & Beitel, 1988 ca Annals of the 754 Missouri Botanical Garden TABLA 6. Listados florísticos de México. Area geográfica o Nümero reportado grupo de plantas de especies Referencia Chiapas 7018 Breedlov Cozumel 542 Télle ORE 1987 Dur 3800 González et al., 1991 Es busier re Biologia Chamela 754 Lott, 1985 Estacion de Biologia Los Tuxtlas 818 Ibarra & Sinaca, 1987 Peninsula de Yucat 1936 Sosa et al., oe étaro 2334 Arguélles et al., Quintana Roo 1300 usa & e лай Tabasco 2147 Cowan, 19 Tehuacan-Cuicatlan 2700 Davila et e Я У" PA ibe i áticas 111 Lot et al., Monocotiledóneas Parte 1 393 Espejo & Verr, 1992 Gramineas de Puebla 427 c et al., Pteridofitas de Guanajuato, 300 z & Palacios-Ríos, 1992 Michoacán, y Guerrero de especies де по езїа Mig: Sites en las colec- ciones. De este modo, flor o las de Sinaloa, nümero de endemikoa у тирга florenen tales la С M o la Depresión м Balsas EM de Guerrero, үчү ia, 9) xaca, у Michoacan) (Lorence & Garcia, 198 Hasta la fe estas areas aun no han sido sufi- cientemente exploradas. Zonas del е del pais, cole énfasis en recolectar ааз рага іпсге- mentar la representacion у variabilidad de las es- arios autores han sugerido estrategias para que son uA por su gran s en el desarrollo de поене del conocimiento flo- i de ёхі Та iu Шер es muy grande. Asimismo € " cuenta con una flora a nivel nacional. Por е0 rma simultánea al pro- gtar ma je? colecció ión y ^d desarrollo de las floras pecies. regionales se inicie una base de da permi En cuanto a grupos botánicos, si se continúa recuperar Paca información florística. Esta con la serie de tratamientos taxonómicos sobre itirá conocer con precisión el numero de or Gramineae y Orchidaceae que se han j iniciado (Ta- = E анат distribuidas en México. д bla 5), sol ubiaceae con base sólo se р a logr S x a alta diversidad en P pais no han recibido un la colaboración de todos los herbarios del país, 3" юз uado tratamiento taxonomico La información de monografias de fa éxico, е por lo Е tanto es muy probable que varios otros grupos botanicos también requieran estudios taxonómicos. 1 5 : me- como de los taxonomos interesados en la flora icana. LITERATURA CITADA ARGUELLES, E., R. FERNANDEZ & S. ZAMUDIO. И: Listado pr a del estado de Querétaro. Pp. Volume 81, Number 4 1994 Sosa & Davila 755 Conocimiento Floristico de México n J. Rzedowski & G. C. Rzedowski оде Flora an l Bajio y Regiones Wd m a c. Com 2: iwe de Ecolo, o - C., i о. В А. А. 1983. Las "Gramí Sas Méx o, Vol. 1 Secretaria de Agricultura y Sedes Hidráulicos, México, D. F. Ta. bero grasses from Mexico XIII. ir s de México, Vol. 2. Seria de Agricultura e у охша Hidráulicos, SDN 1991. Las Gramíneas de México, Vol. Secretaria de Agricultura y Recursos aces Las Cactaceas de México, Vol. id d n H. . Uni a de México, Misi. ` 1978. niv. Nacional Auténom: & Н. SANCHEZ-MEJORADA. 1991а s Cac- táceas de México , Vol. 2. Univ. Мо Же ape de México, M DE. Las Cactáceas de México, А 3. Univ. Miu! А de México, México, BREEDLOVE, D. E. 1981. Introduction to the flora of pest Part 1. California Academy of Sciences, an 1986. Listados Florísticos de México IV. Flora de Chiapas. iun de Biología, Univ. Nacional Autónoma s México, México, CAMPBELL, D. 1989. The “importance of floristic . Pp. 6-30 in D. G. Campbell & H. D. ponen Å (editors), Floristic без ‹ s Tropical Countries k 83. Listados pui fe México I. о de , Univ. Na- ENORIO, E. MANRIQUE, А. MIRANDA & A. Ropnicurz. 1299; Listados! Roristicos de México ҮШ. Listad Puebla. Instituto de Bolg Univ. Nacional Autónoma de éxic o, Méxi Davia, P., J. L ENOR, В. MEDINA, A. RAMI A EAS, J RE KEN & P. TENORIO. 1993 tad i de México X. Flora del Valle de Tehuacán-Cuicatlán. Josue Р pepe Univ. cional Autónoma de Méxic xic .& os-Rios inis preliminar de ез species de ps 2 o gessi de ~ najuato, Michoacan y Querétaro. Pp. 1-57 owski & С. "C. Reed W: L башы), Pora, del jio y Regiones Adyacentes. Fasc. Comp. 3. Ins- átzc 90. Flora del aai er Guerrero, Pp. 494 del XI Congreso o de ae tánica de ee México, F. FET FEREARE ld ү dóneas mexi de к Vol. е o-Univ. isen feci México, FLORES, J. S., H. Narave & A. P. Vovipes. 1992. Mér ida, FRYXELL, P. A. e In J. S. Flores ete па Yucatanense. Fasc. 5. Univ. Autonom: 1290 fM Syst. Bot Monogr. 25: 1-522. Соме7-РомРА, A. GS 1978-1991. Flora racruz. Fasc. 1-66. Instituto Nacional de ae gaciones sobre dicas Rug ticos-Instituto de Eco logía A. C., Xalapa, = e ees БА 2 & H 199]. Li stados floristicos ae Méx o IX. ‘a de1 Durango. rk to de log aie Nacional Autónoma de México, Méx 34i F. . GARCIA. 1990. n Resümenes del XI ad Botánica flora de Durango. Pp. 4 Es Pria Socied 1991. LAZAR (editores) ром c HoLMGREN, P. K. & W. Kid Part 1. The rado of the ab ed. 6 huy eg. 92. ККТ BaRNETT. 1990. Index елет iuto 1. The herbaria of the world, ed. 8. Regn u, W. pee ха Е. ‚К. SCHOFIELD. 1981. Index herbariorum, Pa art 1. world, ed. 7. Regnum ee 106. IBARRA, С. & S. SINA 1987. Listados floristicos de México VII. Estación de Biología Tropical Los shea аз, Veracruz. Instituto de Biologia, Univ. Nacio México, p. F. ERRA Araliaceae. Pp. 1-23 in N. Diego (editor Flora de Guerrero, Vol. 1. Facultad de Ciencias, Univ. Nacional Autónoma de México, México, D. F. Lorenc, D. Н. & A. Garcia. 1989. Oaxaca, Mexico. Pp. 253-269 in D. С. Cam H. D. Hammond P (editors), Floristic Inventory of ун Countries. New York Botanical Garden, New Yor Lorr, o III. La Estac ción de Bielogis Chamela, Jalisco. “Inti de, e Univ. Nacional Autónoma de Méx 3 1983. Gramineae. In W. R. Anderson or), Flora Novogaliciana, Vol. 14. Univ. Mich- д Pres, ма, rbor. mpos n. W. R. Anderson (edi- tor) 1 Fiora "Novos Vol. 12. Univ. Michigan Press apeiron yn Lene In W. R. Anderson (ed itr "m Novogaliciana, Vol. 16. Univ. Michigan Press, Ann Arbor. Legu e. In W. R. Anderson (ed- tr) Fr нате Vol. 5. Univ. Michigan Pre: nn Ar е то Dioscoreaceae. In W. R. Anderson ean Fra tert Vol. 15. Uni DN. ms pir Pteridophytes. In‏ س W.R т vb те Vol. 17. D. conem Press, An Annals ore ы Garden MickeL, J. T. & J. M. BEITEL. 1988. onim hg Flo: ees Oaxaca, Mexico. Mem. New York Bot. -568. Мок, P 1992. Neotropical floristics and inventory: Who will do d pod P: ittonia 44: 372-375. PRANCE, С. T. tic inventory of the or eh Where do we а ы nn. Missouri Bot. Gard 659-684. 78. ш e the tropics: A FE Ann. Missouri Bot. Gard. 65: i-i. ped The present state & D. С. САМ of tr dia ewe Тахо : 519-548. bei n 988. Tr iai Pie tomorrow. Taxon 560. A soot. Mexican ae ee Distribution and ee dug 379-395 . P. Ramamoorthy, R. Bye, A. Lot. RE Fa ОШО), Biological Diversity of MI Le and Distribution. Oxford Uni Press, New Yor| Rzevowsk, 1: 197 A Catálogo de los herbarios insti- onales mexicanos. Sociedad Botánica de México ico, D. F. 1978. Vegetación de México. Limusa, Mé- ke El endemismo en la flora fanerogámica a: Una apreciación analítica preliminar. Acta Botánica x Мени 15: 47-64. 1993. SH iw and origins o ris a үчн. кү he E ni T UP J. Fa (edit bri) ве loge iuis E rear a Org and Distribut Oxford Univ. Press, New & G. C. RZEDOWSKI. 1979. Flora Fanerogá- mica del Valle de México, Vol. 1. Compañía Editorial Continental, México, D. us lora Fanerogámica del le de Méxi o, V ol. 2. E Shine: cional eie] Messen de Ecolo, ogía, A. C. México, D. F. ——— & 989. Sinopsis numérica de la um fanerogimica Valle de México. Acta Bota- a Mexicana. 8: 15-30. 1990. Flora peak кезү. del bue: de México, Vol. 3. Instituto de E a, Pátz- — — — (editores). 1991-1993. Flora del oy raro мин» Fasc. 1-16. Instituto de өзб. a, А. С., Patzcuaro. ЖО А. 981. ендер ны. is : D. Е. Breedlove editor), Flora of Chiapas. Proc : Сай. Acad. Sci. 2: Soss, DU “cr 1992-1993. Flora de тн. ыг 72. Instituto de Ecologia, A. C., Xalap . Fıo RES, V. Rico excom ti eee ORTIZ. 1985 ista F| — а Maya. " V. Sosa (editor), Etno spleen Vol. 1. Instituto Nim de asini iens sobre Recursos Bióticos, Xal Sousa, M. & 1983. —€— bre de México de Quintana В He ‘IL. Biologia, Univ. а Auténoma Pe Ms ёхісо, хісо '& S DELG. 1993. Mexican Legumi nosae: Phyiogeography: endemism, and РЕ " 459-511 in T. P hy, A. Lot & J. Fa (editors), ese Diversity of Meni’ Origins and Distribution. Oxford Univ. Press, New York. Camera SraNpDLEY, P. C. 1920- ы Trees and shrubs of Mexico. Contr. U.S. Natl. Herb. 23: 1-1721. ога of ate Field Mus. Nat. Hist., 57- Pinus: A Mexican poi mamoorthy, R. Bye, А. Lot t&J . Fa (e ез). Biological Diversity of Mox со: ures and Distribution. Oxford Univ. Press, New ne Genus E Tee, 0 bo E. C. CABRERA. i T tados Floristicos co VI. Flórula de a: Isla zumel, Quintana Roo preci de Biologia, bat pce Autónoma e México, México, & A. FLORES. 1990. La flora de Nayarit. Pp. 494 in Resümenes del XI Congreso ме де Вк Sociedad Botanica de México, México, TOLEDO М. & V. Sosa. 1993. Floristics in La America and the Caribbean: An evaluation of te number of plant collections and botanists. Taxon 42 355-364 IAU-UITZ. P In: n Asteraceae. Pp. 5 vog R. Bye, A. Lot e J. = (ойо) E ве loge Diversity of Мей Origins Oxford Univ. Pre ss, New York. (editors). Asteraceae of Mexico. University of х Austin. (En prensa.) i VEGA, В. 1990. Flora de Sinalo: 366 in Resümenes del xt ie greso Mexicano d ánica. Sociedad Mexicana de Botánica, Ме WIGGINS, L Er 1964. Flora of the Sonor an Desert. d F. Shreve & I. L. Wiggins bene: Vegeta en Flora of the Sonoran Des 6 Stanfor pans niv. Press. Stanford, Cali 1980. Flora of cau Салар Stanford Univ. йр, Stanford, Califor n ICE I. gene Mexicanos no registrados € Ni Herbarior de la gric. Jalisco: Herbario, u E de Agricultura TM de Gua 14.5 e Nogales-Las Agujas, Ejido ынан 4522 ыд Jalisco. Cent. Pec. ie rbario Regional del Centro de а 1 uarias del Estado de Sonora, SARH Carbó, Sonora. Facultad de Ciencias, Univer а, Km. 14.5 carretera Nog Las Agujas, Ejido sisi 45220 Gu мша, d om. er VU Brigada II de la Comisión Téc Cons Та Determinación de los Gris de азата SARH, Medicina 9037 А, ua. Córdoba: Нео Jerzy Rzedowski, Unies Vera- ‚ Campus poen Zona Córdol cruzana i camino Peñuela Amatlán s.7., Peru Y Саш: Herbario, Facultad | de Estudios Superiores de en Cua titlán. 1 Autónoma Volume 81, Number 4 1994 Sosa & Davila T5 Conocimiento Florístico de México o, Departamento de Ciencias Biológicas, Km. 3 Dien Cuautitlán- Teoloyucan, Estado de Méxi- Inv. Sureste: pepe del Centro de Investigaciones Eco ureste, Apdo. Postal 63, 29200 0 San Laguna: Herbario, Universidad Auté oma Agraria “Ап- tonio a nidad Laguna, carretera a Santa Fe y Periférico, Apdo. Postal 940. 27000 Torreon, Coa Linares: Herbario, iren de Silvicultura y Manejo de de Recursos Reno ables, bai niversidad Autónoma de Nuev ben Unidad Lin єє Ajo: Postal 41, 67700 Lina o León. Mice: Herbario Micológico de Marei, Uen Ae on a del Estado de Morelos, Fac cias Bolas Av. Universidad 1000 ТЕ Col. Cha- 0. Саала avaca, Morelos. Normal Tes: (rain Escuela Normal Superi Jalisco, Calle Alej sode diss 3317, Vallarta ma ps ge, сува lajara, sali Univ. pres Herbario, ГЭЕ ‘Prof. a "upon Salas," Universidad Autónoma de Chapingo, Km. 38.5 са rretera México- Texcoco, 56230 y. ani Univ. Guerrero: Herbario, ee Superior de Agricul- rsidad Autón ira Guerrero, Periférico s.n., Iguala, Guerre dad Autónoma de Puebla, 4 Sur 104, Puebla, Pueb- la. niv. Querétaro: Herbario, Universidad Autónoma de Que: фра aro, Apdo. Postal 494, 76010 Querétaro, u Univ. Sinaloa: pove rodeo — de Agricultura firn " Uni e Si- pers Pos tal 7 26, ү Univ. же ben. Universi x- c de Investigaciones Biológicas, Apdo o: Нег nida de servicio y apoyo al gna del Departament de Producción Agri- ae imal, Universidad Autonoma Metropoli- a-Xoc himilco Calzada del Hueso 1100, Col. Villa Quie tud, 04960 México Univ. Yucun Herbario d e la Universidad Autónoma de Yuca к tno! sopi Yucatanense, Apdo. Postal 281, es Escuela Nacional de Estudios Pro- sida Nacional Autóno- , Col. e de D.F Мег Zaragora: p Les олив Zar , Univer México, "A C. Bonilla Oriente, 09320 „Iztapalapa, Маб, J estudios de Zonas Aridas y Semiaridas del Cole egio ex Postgraduadoe, Iturbide 13, Salinas de Hidalgo, n Luis Potosi. Zoot. Chapingo: Herbario, Departamento de Zootecnia, ersid de Chapingo, Km. 38.5 ca- Pon Univ. шо bilê Reg nal, e Hidalgo, Centro ie pueden a Pachuca-Tulancingo s.n., 42000 Pachu- idalgo. Univ Nayarit He idi Cien ifica, misa Autónoma de NAE Do- пісі lio Conocido, Tepic, Nayarit. Univ. Puebla: Herbario, Facultad de Ciencias, Universi- rretera (€ Texcoco, Texcoco, Estado de Mé- Zootecnia gripe hogar d de Zootecnia de la U rsidad ue ч a de Chihuahua, Km. 6.5 с tera Chih ua Cuauhtémoc Ардо. Postal F- 28, Сална. тея PALEOBIOGEOGRAPHIC DISTRIBUTION KUYLIS (CYATHEACEAE)! Barbara A. R. Mohr? and David B. Lazarus? ABSTRACT The f, 4 Kuwvli h ub X E 1D. fal. N, species, of which К. mirabilis, K. separatus, and K. scut Th atus gw des waterbolkii, e fourth s a oe that has ‘identical features with the spore | of the extant = ner Cnemidaria, has und exclusively trend to increasing restriction " ure Neotro| t Kuylisporites JER A has been found in the southern Gondwanan realm from the Early Eoc ene through 2 ) е. New of K. u wat terbolkii y area (Antarctica), ier a аба А + whe: ati o cooling took place, y waterbolkii tralia, where climatic dee was balan orthward drift, K. waterbolki persisted зн the led, however, to its extinction. In South A а climatic conditions red were w. xclusively in an a In Au early Moda ne. "Increasing song ht in ted fr a that overlaps wit ced by 1 Austr alia during the Nea rh ree cene. From de Oligoce n: area of extant Cnemidaria, which п К. п found е lives mostly i in (sub)montane forests i in tropical Central South America and the An m Morphological identity and K. w erbo lkii can be « correl ela peur к= of extant t Cnemidaria. If this is the case, the Recent area m лон С о be a relict Broad anéteal biogeographic distributions of a have been known since umber work by e y botanists such as Hooker in the 840s iere . Hooker noticed, for e ample, that the Falkland Islands, South Georgia, Tristan da ll in the th Atlantic), and RO erica, contain plants me related : taxa from Tierra del bec iion observatio d botanists, such as, e.g., 2 material for this study was r given af we former curator о . S. W. Wi ta i y (Florida State Univ., est d bac ickgr round informat A. Tra ries. the manuscript by mprove adding ispo Gard. Zürich) pr H ovided MOF eg uis ce and spore material of e P reviewed an Pedes r dr. raft ities of UAR: to conclude that Antarctica ec have kay ge шс cti an, Minen 1 OS s modern plate ‘tectonics had explained the fragmentation of (ош ч апа v t con tin Son me aps including Fa conifer, and angio- sperm taxa urrently distribut tad i on only one of the Sout dni ела ге continents, Pane the paleobotanical record shows a more wides pr distribution. Examples of this pattern are the © Research Facility, D f the Antarctic Re: ins pare: pa аздер, жарча Ean also £ F р! 57 subspecies of re minas. We thank al ssian lite To ярыш nal Foundation = No. ss Nat r retire uei , ETH, Sonn anonymous Peviem ers valuable ) S. "€ Zürich), the photographic мы уз U, [oer ee pri express 2.654. 087 | and 21-2' eggstr. 5, CH 8092 Züri inancial support for lodged “гє warmes 920.90 i ` С ackno ich, (erai ANN. MISSOURI Bor. GARD. 81: 758-767. 1994. Volume 81, Number 4 1994 hr & Laza е Distribution of Kuylisporites 759 nifer Microcachrys Hook., today restricted to Tas- mania a according to the pollen record, pec distri s in Cretaceo LAN Tert times son & id 1954). 5 кыйа EN was pico dq by a d e соь and, in addition, as а b ridge between үч Gondwanan continents. ng seid E idea of Antarctica 3 n y fer: is nowadays widely abandoned, the кейде ЖЫНЫ» southern continent had been confirmed for erochronous stratigraphic occurrences that the anarchic Peninsula posed з as a dispersal corridor ҺА merica and Aus- tralasia for two fern morphs avifera triplex Bolkhovitina, a жыш Гез and Azolla Spp., a ubiquitous water fern. СЫ кысыш, annulatus Cookson ex Potonié aa +h 1 ££. : 2 1 i f, a г © spore data. This fern spore can be correlated with d aant Чеп genus [ороо (шна 86) k di South Mid Central rire and ds Greater wet (mon ) temperate environments at high altitudes, mostly over 2000 y in uthern South Ameri si o grow at low cepe a pen e times, а , this ge- wn from the tro t ad it 18 0 ntis occurred in southern high lati- des—in southern South America from Cretaceous to the Oligocene, in Antarctica from мам му Cretaceous to the middle Eocene, an mann, 1986). New evidence from Ku uylisporites waterbolkii товар (19501-2 spam. identical in es e the genus might have had a more widespread dis- tribution during the Paleogene in the Southern Hemisphere. The occurrence of Cnemidaria- -type isl dur- tic са used as а уаша 1 эол en a е patterns and ра- gr iens nmental conditions during he oic phe optimum in southern high hides (about 09739). This pias of Сепогою interval of about 58 to 45 Ma. and has been documented in the Weddell Sea and southern In- ian Ocean je ON by microfossil groups such as foraminifera (Sto 0 1990) and tingle d o& ohr, in prem; Mohr & prep. ), ar bl (Hsü et al., 1984; K ennett x статок 1990; Ваг- erra & Huber, THE FossiL SPORES The genus Kuylisporit ес lis (Bolkhovitina) Krutzsch, К. separatus Chlonova, K. s s Newman, = К. water- bo rend ‘Excluded poene ur € consi are the the three sides. Dettmann hat K. lunaris, found in the Uppe southeastern Australia, differs from all the other nown species of Kuylisporites. The oldest species of Kuylisporites clearly be- ene] to the Cyatheaceae is K. mirabilis, kno om be Lippe of Siberia (Bolchovitina, o the sp nem 2^ (С. PSH IS a th t 1 Ты Orkney ге region E adds further credence xa have at least occasionally rd from original high-latitude Southern emisph 1; °1 . . 1 pay a | : o in the iem The number of Cnemidaria species (23 E to Stolze, 1974; plus 2 by Moran, 1990) in northern South, America (сенер M. and Vene- zuela), as well i species, led Stolze sige to the conclu-‏ چ sion tha the genus may have ted in this‏ 1 1 — erly Sites coin America and central Su th America). ccording to the fossil spore evidence, however, -— (ach 1959) = it dene have been m Cre sediments (Krutzsch, i c of Kuylisporites, K. separatus on the surface. In this respect, the spore is similar to the North Ameri tus. e Campanian ا‎ а (Newman ‚ 1964, 973; Beeso ie uA and New Jersey (Traverse, pers. comm m). Th cies is very small (average size 27 um) лода) with the (younger) Southern 760 Annals of the Missouri Botanical Garden 101578-59 South Orkney | Bruce Bank ¥ 0. 101578-59 Weddell Sea 40° 20° Location map of Weddell Sea, Antarctica showing location of Florida State piston core Islas Or ae (IO) 1 59, Ocean Drilling Program Site 696, and other geographic features referred to in the text. Light sil раа ja: edge of yai. td continent indicates ice-shelf. Inset (a) shows approximate location of core 1015 59 in middle Eocene times. Continental reconstruction created using Terra Mobilis program of Denham & Scotese 7). Непизрһеге ra K. waterbolkii and has, ac- Ma. at L4 gw Bank, South Scotia Ridge off th the original 4 cording to t escription, a slightly ru- Antarctic Peninsula (Latitude 50335 gulate surface on rta distal side (Newman, 1965). ткн 13'W; mbsl 2707 m, see Fig. 1). Furthermore, this species lacks any additional scat- Acco g to Toker et al. (1991), the nannofossil tered p ssemblag ese strata can be assigned Kuylioporites e is known from the to the Chiasmolithus gigas Subzone of the Nan- late Early Eoc n from southeastern Australia — notetrina quadrata Zone, which equals late earl (Stover & Partridge, 1973). Kuylisporites water- to mid Middle Eocene. Following Berggren et а]. been pic med Нели eli ia : Ьу variu authors (1985) timescale, an absolute age of about 47 to see below : i bees Ais the Oligocene on. The tre for this study comes from the upper A list of the known fossil occurrences of the 130 cm of a 3.85-m-long piston core (101578- spore yes gio sek is given in an Appendix 59), which contains olive gray to olive brown conr to this acted clay. From this interval six samples por taken, which were processed using 5 dard ow: SPORES FROM BRUCE BANK, OFF THE NORTHERN trifuge preparation techniques. In addition, sieving ANTARCTIC PENINSULA EC ‘eve and oxidation with HNO, were a. Material and method also carried out. The latter procedure was 4 P w evidence has been found from middle Eo- filled a good ori o the abiit T ie Ma.) i Orkney area (Fig. 1). The material was collected jelly were prepar The m during the Islas Orcadas cruise 1578 (Cassidy, reserved ділова" aterial is iis in well-p Volume 81, Number 4 1994 мо & Lazari 761 scbogeegrapni Distribution of oe eelste, pe (Meo & Mohr, in dope мы кай a 2 (Mo cim ep.). From Core 101578- 39 n more than 30 бов; of about 15 to 20 taxa per pep (Mohr, in prep.). е, equaling abou to 1.0% of the total sporomorph content. b. ee С ат a -type spores from the Sout ney a Kuylisporites waterbolkii Potonié (1956), Figure 2A, B, C. IRAM res, amb rounded triangular. Lae тан straight, long (up to 4 of the nod spore dia pneter). Fo ош, ipu fovae (1— gi m) Over Three large а (ТЕЛУ? about 5—7 pes are found symmetrically arranged at the center of the three sides. There, the thickness of the two-layered wall usually increased bim 1.5 to 3 um). Size of the spores: 27-(3 ae Comments: In size neat smaller than spores of most one кы, ipe Cnemidaria, but other- wise identical. GEOGRAPHY AND EcoLocv or EXTANT CNEMIDARIA The geographic range of living Cnemidaria with ran, 1990) covers the area of eate northern part of South America (Tryon 1592). Th The distribution: is generally COBEN, (Stolze, 1974). Ther ere pn bond according to this author, two notable regions of disjunction: en Bra: mnt and а an ree е. the Р eh, E edt province of Alta Тараз Ж. Most Cnemidaria species have a subarborescent habit, a Kava usually less than 1 m in length. These ferns are largely рейт to deeply дей: а 8 some- = Ф moist habitats, in or at t s of forests, some alo eks and ETE at elevations from sea level t m, but typically bermeen 500 ) and 2000 т бое, 1974). Lellinger one of the areas of occurrence of Cnemidaria, in the matos forests of the Rancho Grande” in north- zuela. There, the cloud forest is found at ru xia 1150 m and about 2350 m. vn annual precipitation is about 1750 mm, with mos of the precipitation (1600 mm) falling during ча season from April through November. Daily tem- rature ranges are generally low (16-21°С). THE SPORES OF EXTANT CNEMIDARIA Te features of ше extant genus Cnemidaria partially shared with other (aches: 1974; Tryon & Tryon, 1982), bur lete spores with t} is fer: genus. Cnemidaria pores are died om three pore in the equa atorial «ек. equidistant be. unique to pores (Fig 2).I so special that ah pases of уду of the го of A Vries-Labo um) chose the outline of this taxon as its ogo, pcne of extant Cnemidaria were previ descri detail 2 um. The surface of the e less smooth, but shows vd typical large pores (up to 15 um б. ә, inue des the pier of рп of the three si may be ра аан, over the spore sur- ave examined spores from the following Tryon, Guyana, Pipoly С pec (Kunze) Tryon, French а Сге We ee SUE spore зое ps en pest Bon: with one сасне In very rare of the spores show ы {з гее esed poor (Fig. 2G). This is ee true for fossil material (Fig. n = oa We in this paper that the paleobiogeo- graphic gates pattern of Kuylisporites wa- terbolkii spores fits wit ith me idea that Ew 97 to. del Chocó), where it was ri seen at roadsides at altitudes at or below 500 m arely up to 1000 m. Mágdefrau (1960) gave more кед м Jd for related form of Cnemidaria. These ancestral ipe - cies might have been a species of Cyathea, since many of the spores of Cyathea have a pitted exo- Annals of th Missouri Mond Garden uylisporites waterbolkii; Sample A 59, 26-28 cm; Slide B Fic K ET oo 1578-59, 123-125 cm, Slide D, 26-28 cm; Slide B, x 1000 аА „тше d E. Alsophila decurrens var. ook., decurrens Hook., аң olu bei Fons 1523, x 1000. mith 948, x 1000 spore. However, these pits are irregularly distrib- uted and v e resence of “‘intergeneric”’ hybrids (Holttum & Edwards, 1983 а); PALEOBIOGEOGRAPHIC PATTERNS orted occurrences of species of Kuyli- Upolu (Samoa) Sedge 1523, 1000—B. Kuylispo 39, со Ee Kuylisporites шато Sample D e 1578-59, 123-1 m; Slide A, x10! М x 250.— ” Alsophila dare ча ГЕН: азаа horrida. (L.) Presl, Venezue not been usua since without such negative аст the limits of its past nce, we ca : ve evi bution. Arguments pee partly on neg prei wes dence may be risky, as future discover! of. «e change an area of non-occurrence et one ize this currence. We havet ried to region ا‎ when ede are we li- r cases of occurrence a Kuy sporites Volume 81, Number 4 1994 Mohr & Lazar Paleoblogeographic Distribution of Kuylisporite 763 En 3 Брит of Kuylisporites. waterbolkii Qo and ions ond is Ax +) rh by EE Rae sud indicate interval with no records due to continental glaciation. 8 g £ с с © = 3 У ES Oe зш ч ee ЖЕ < ш = Os oet < < тд Recent m Г] о х х 0 m Quaternary 0 Ll E х Г] О Pliocene m m m E M О L Miocene M| D o Li x Bi m) E х х х L * * Oligocene — E- + -EI- - O - 33,4 -—d-B--3€*-|-— — E * Ё х Еосепе M +? x x x E х Paleocene + + О О о [Maastrichtian * + + 0 Pre-Maastr. + + + U Pre- Paleogene с occurences of ое аге known onl 1, Fig. "ui — pores, even though ш ге- semble К. general aspect, аге sien diferent M. to be сорајсагое we er ancestral о; most likely of c ores о e fon not neces- sarily identical to Cnemidar. d-Eocen exclu ively he therly palaeofloral pr province, the Nothofagidites province Tecognized in Australia, South Ame: and - arctica (Table 1, Fig. 3b) trata con- "a g Cnemidaria-type spores are of (late) early ne age from Australia (Stove Partridge, 1 а Rico). е occurrences of Cnemidaria-type spores (Table 1, Fig. 3c) a are restricted to early Miocene ca, and northern So Has жое цен» сап » Eid as T ys T } +} M. 2328 . Between the late Eocene and Oligocene, 1990), Cnemidaria persisted nati th e early Ma cene ever, led to its сена Бо. In rea shifted from em latitudes à ; s warm-temperate, gocene, | where Cnemidaria lived, lives, р perate conditions. Only a few ees, а as С. па are! still o tem Annals of the Missouri Botanical Garden Volume 81, Number 4 1994 Mohr & Laza Palecbogeogratic Distribution of Kuylisporite: 765 ectabilis ionall und in the low- lend feste of northern South Senate (Kramer, com Kayporites me ж been eo from South Africa (Coetzee 1983, and literature cite over the Sout here жү МОМ had drifted away from Af- is, not earlier than in the late Creta- TEAM the pria پوت‎ in Table sheds light on on the possibility that Cr ا ون‎ 1 suggests that only those species re today restricted to Central America and northern S. as spores of eh xps are not ои from oe ternary to — nts Asus similar wall structures LO & Lugardon se min ial in coll Botanical Garden of Zürich 0 wh (Cyathea) decurrens Hooker, Sledge 1523, Upolu, SUM ) not much bigger than those encountered in A. re addi daria, чп us aired 9 лш decurrens round the exine. This et end the range of mor- me variability pur jos that the 3-pitte ted за occasionally see ila are о tally similar to ves origin o Ее пег}: the occurrence ee 3-pitted ore: xv dire ectly 1 3 Н = ur J of Cnemidar ta spores discussed in this paper. We therefore conclude bas the association of this distinctive spore's distribution in Quaternary үл їз with the ‹ опе аи савы їахоп P PUN the use of Cnemidaria type sp the genus. a aor for CONCLUSIONS e cyatheaceous spore genus Kuylispor- tes, кыс ord four species, vo B ted in the rn Hemisphere in the late Cre = Kuylisporte waterbolkii, closely жы то J is known from 25 the Early Eocene on. It was шша to the иши Не ie ore er realm, ё у & Tryon 197 76: 574). In addition, copies of of SEM pon ographs, generously given by ihe E y P Chan: 5231, Society {д were used for our studies. We note that in A. десш еә var. маа. (Fig. 2E, Е rens (Fi ү th under Ыр. Only SEM pictures reveal the vise e ts surface of the exospore (Gastony i Tryon, 1976). ems too cur r only i m big pores, like in spores of Cnemidaria, to relatively small pores, Afric н From the Middle ви on the biogeographic erbolkii becomes gradually ia occur: y. The bs Antille might be considered a relict area for is genus 3. Some fern taxa associated with K. wate ce such as AES nd ces ш сеае), с m the ecological in- irren s d wat жыш as an indicator of ра шр not tony Te ee е ге- п arm-t E jr, ich the K. w secs pe Jetitudes iren ri. ro aiam Cnemidar a migrat ted n orth when Ашка be- Si i 1 ince came heav Locatio on oe Cnemidaria and Cnemidaria ccurrences on а 4 of Denham & Scotese Пт, а. Cretaceous Paleoc . Neog ns created ne occurrences on a 6 8 Ma. a 15 Ma. map. м е occurrences оп п ТаЫе 1 pnr "t f abe te Annals of the Missouri Botanical Garden most of the species of Cnemidaria seem to avoid bad орна lowland һа bitate, Ciemidori ria mi- the Neogene. LrrERATURE CITED ASKIN, R. A. 1989. Endemism and heterochroneity in the Late о Paleocene pal- ynofloras о { Seymour I Island, Antarctica: јара бащ for origins, ние and palaeoclimates of southern floras. In: J. А. С AM (onmi talus of the Antarctic ‘Biota, Geol. Soc. Special Publ. 47: Bannera, E. . T. HuBER. 1991. Paleogene and arly lealea oceanography of the southern Indian Ocean Leg 119 foraminifer stable isotope ear oe Drilling Program, Sci. Results . C. 1992. High Resolution ае taphy across а Marine SUMA Tertiary Bound- al County, Texas. PhD Y Thesis, Pennsylvania State VM та Park. BERGGREN, W. A., D. T, J. J. FLYNN & J. ۷ ка д СЕХ койдо. Geol. Soc. Amer. сү e BorcHoviTINA, N. A 53. соз. of spor ы p in ond sediments of the centra al ы. Acad. USSR, Geol. Ser. 61, Moskau. [In Fon ee Н. и. Jj: . PERCH-NIELSEN. d e iie Ў — Univ. B Ene рй 5, e en 1980. 8, sedim secl Seen de Anse DM Facili- Geol., Fe Dept. Florida State Univ Aug hassee, Sediment. Res. Lab. Contr. 8: qua A. F. 0. ifi t nd spores in Upper Cretaceous from pus eastern MS of the estin Siberian d ophys., No. 3, Akad. "Ned 4 fin ussia COETZEE, J. A., A. SCHOLTZ & Н. J. DEACO! 983. Palynological бр; Fe the vegetation Мау of е Fynbos. H T. Denton, O, B Herdi S dis Y N. T mbr: weby (editors) [ee Ec NET Trudy Inst. Geol. S.S.S.R. Novosibirsk ology: A Pre liminary Synthesis. бо a National Seni Programmes Repor ( (stk) Not RT Cookson, I. C. & M. E 19: gum trilete spores from Upp үш i Adet ae PR c Ри, Коу. үч ты 70: 95- 128. PIKE. 1954. The fossil occurrence of а Phyllocladus and two other podocarpaceous types in Australia. A J. Bot. 2: CoPELAND, E. В. Antarctica as the source af existing ferns. Proc. 6th P g 625- DENHAM, C. R. & C. R. 987. Terra Mobilis: A Plate Tectonic Program f or pe Macintosh (com- к кон Earth in Motion Technologies, Aus- M 1963. Upper Mesozoic ше‏ ا iib. cantern Australia. Proc. Roy. Soc‏ аа (e 1 Significance of the Cretaceous- Tertia- ry spore genus Cyatheacidites in tracing the origin and migration of tephosoria (Filicopsida). Special edt P tol. s 63-9. ENGLER, A. 1882. ee einer r Ent ntwicklung: te der ler Конен n Florengebiete da тл Hemispha re und ris iride Gebiete ded ki. bc 3. Maestrichtian Mua ЧИШ in s, Maryland, a nd New Jersey. Geoscience and aon & R. TRYON. des Spore morphology in the pies 2. The genera Lophosoria, Me- шуа, S phaeropteris, Alophila and Nephelea. gs 63: 738-758. Mosen ў! 1976. Studies in kagara e. i obotany. II. The Miocene communities of V uz, Mexico. nn. ee ome Gard. 63: 787- 1988. Lower Miocene floras id iggy of iicet America. J. Geol. Soc. Jamaica 25: 8- 91. Studies in neotropical paleobotany. ҮШ. The Pliocene communities of Panama—introduction and ferns, gymnosperms, angiosperms (Monocots). Ann. Missouri Bot. dle E Ea 190 & D. M. Jarz LS Kee in n enia ities of Puert paleobotany. L The O Oligo Rico. Ann. Missouri Bot. "Ced. 56: 308-357. & E. Gonz ALEZ. the Guaya vo ads of H. P is in the years 1839-184 kn os., London. Reprinted 1963, J. Cramer, Hs, es i J. A. MCKENZIE, Н. OBERHANSLI, H. WEISSERT . C. WRIGHT. A South Md ioc Paleoceanography. 1 K. J. Hsii, J. A. А Re nitial Rep. Dey Sea Drilling P am 1%: 77 KENN . F. BARKER. 1990. Latest Cre te y ij^ & hi de taceous to о Cenoroi climate and oceanographic velopments in the Weddell Sea, Ana са: An ocean pm drilling id ective. Proc. Ocean Drilling Program Scr ^de I13-937— Frac KRAMER, . GRE The rong um i Genera of ш Pas. т Preridophyies ei osperms. Springer-V тп, Heide es Kru с W. 1959. Mikropalontologiche (вро Verses Untersuchungen in der ү des yo eltales. Geologie 8(21- 22) 1-425. Kuvr, JE MULL ER Й н. WA зч The а nd ы alyno! px to ae ш t рауы ela. Geol. е il geology ™ ou g п Mijnbouw Li: 49-16. RS ixi D.B. 1975. A physgeograme bns ioe неруе е 10 9. Pay MacrHarL, М. К. & E. M. TRUSWELL. meee патина of the ofi west МЧ ОН isi Mineral Resources, Geol. & Geo . Aus Geol. Сона 11: 301-33 ar Volume 81, Number 4 94 960. Vom Orinoko zu = Bese "тепа Naturf. Ges. Ziirich 10 o, S. & MORE (in press). |! Middle pcysts fr E h Scotia Rid their КУ implications. Rev. Palaeobot. al Morr, В. e R. 1990. Eocene and Oligocene sporo- morphs and dod llate e cysts from Leg 113 dr ill MORAN, о new species of Cnemidaria (Gyatheacea) from anama. Ann. Missouri Bot. Gard. 7: 246-248 NEWMAN, I K. R. асел we ову formatio ons, prins estern Colorado. Soc. zs Paleontol. & Mineral., Special Publ. 11: 169-18 Upper Cretaceous-Paleocene pak mor rphs m Northwestern Colorado. Univ к Stud., Ser Earth Sci. 4(2). OCKNALL, D. T. . MILDENHALL early elo spores d ke бев Southland жш Zealand. New Zealand Geol. Sur Palaeontol. Bull. 51: 1-66. ‚ К. 1956. Synopsis der Gattungen der Sporae dispersae. 1. Sporites. Beih. Geol. Jahrb. 23: 1-103. StOLzE, R. [с Taxonomic revision of the genus ). Fieldiana "Bot. ) 3%: 1- Antarctic Pa- у: Stover, L. E. & Р. В. EVANS. Upper Creta- ceous-Eocene spore-pollen zopation, E one ы e LOU Special Publ. Geol. Soc. А tral. 4: (cdm ARTRIDGE. 1973. Tertiary and Late Cretaceous ton Б diei from the Gip еч Basin, Southeastern Australia. Proc . Roy aN toria 85: 237-286. E THOMAS, 1990. "rà A eben through Des is deep-sea benthic for sgh ud Rise, Weddell Sea, Antarctica). e Drilling Program, Sci. Results 113: 571- Токев, v. S. W. Wise, JR. 1991. "adim d ж Sci., Ca mbri de 1987, Cambridge Univ. Press Camis e. ON, A. F . LUGARDON. 1991. M аш of the ET Springer, N New Mosis lin Tryon, R M. & A. F. TRYON ТАСС, J. са gn HE. Pattern of sl eading (less than 4 ма Уу! cuin breakup [4 ма) їо А20 (44.5 Ma) off the southern т а. Bur pass & Laz 767 отне Distribution of ROME Mineral Resources, wee > Geophys., J. Austral. Geol. & Geophys. 1 9-507. WEGENER, А. 1926. Pillogogaphiche еч der Theorie der Kontinentalverschiebungen. Pp. 174- I in 0. Kende € , Enzyklope MER der Erd- unde. Deuticke, Leipzig APPENDIX. Occurrences and publications of Kuyli- sporites sp. 1953 онаи юн Bolkhovitina: Bolkhovitina (1953: 47, pl. 6, fig. 10); Cenomanian of USSR, eastern slope of the South- Ural, near Nowo-Niko- laew 1955 Cyatheaceae ree des Kuyl, Muller habe тш pl. 1, fig. 7); late Tertiary of 1956 poris waterbolkii Potonié: Potonié (1956: 3, pl. 1959 joined mirabilis (Bolkhovitina) Krutzsch: Krutzsch (1959: 190), base ia mirab- i d Chlon -6 зю s Машка sia 1960 НЕНА: уап der & Gonzalez (1960: ti p. am JD er j meii of Colombia, 1961 "Regio (cf. Hemitelia) sp.: Samoilovitch et al. (19 1. 79, fig. 4); Maastrichtian of the USSR a via om байсу 4); tace 1965 ое scutatus EN an (1965: fig. 1); Campanian of знао Colo- 1969, “emit (Cnemidaria): Graham & Jarzen (1969: 7, fig. 9); Oligoc oo of Puerto Rico. өэ Корте scutatus Newman: Evitt (19 е 1, fig. autera bi nd Paleocen ne of "Tense 1973" Короче waterbolkii d 3, fig. iiis a early Eocene to late southeas та, Коориа. орана т Potonié: Stover & Par- phn el de oni Eocene to early Miocene tern he 1976 ' Heitel Sorts (1976: 812, figs. 23, 24); іосеп ехісо. 1984 ерон waterbolkii dp Lye wigan Mildenhall (1 ah 20 pl. 2, fig. 6); late Oligocene to early Miocene of New Zealand. 1988 болшш: gna (1988: 10, 11); early Mio- cene of Costa Rica 1989 auc yrs waterbolkii Potonié: Macphail & Trus е 30); late Eocene to 7 late Oligoce in, southeastern Australia. 1991 Crema: Graham г 193, 194, без. 12. of Рап 1994 X Kuylisporites нарасте) Potonié: Mohr & Laz- us (1994, Sa ыш er); mid-Eocene, Bruce Bank, CHROMOSOME NUMBERS IN Guillermo 4 Norrmann BOLIVIAN GRASSES TODA иегер (GRAMINEAE)! ABSTRACT Chromosome numbers are ~~ for 59 collections of grasses from Bolivia representing 43 species and 21 genera. First chromosome y are reported for the Imperata tenuis, 2n = 20; Paspalum acumin natum, 2n = 40; P emp fi. 2n = 40; Р. отац 2n = “40; Р. lineare, 2n m M Р. macedoi, 2n = 40 & f. Мыр ж] 2n = 40; Leersia hexandra: 2n = 72; БО: ШЫ ешт, 2n = 45; Setaria vulpiseta, 2п = 54; Тһгазуа petrosa, 2n = 20. Information on chromosome numbers in care- MATERIALS AND METHODS Plants were obtained from seed collected by a opulatio: s T. ERR 3 Ed i * livia. Most accessions originated from the Bra- Davidse et al., 1986). An understanding of the alan Shield region of Santa Cruz (i.e., Prov. Ñuflo Бен еру Pucca de Chavez), but те specio were е с an‏ ا Paspalum , Pan- icum, Picken thin Bothriochloa, Fue Шелл: -— is ier by plant breeders and, ede er T ves- ruz (Prov. Andrés пе) or the dA sa- vannas of the western Beni (Table 1). Conn vasis s Мосо, 1987). y Cou icta Chromosome counts were made Wet me -ti tte г a ш counts, have been obtained: over the er of root-tips collected from po a i e 1 : ү ag 1c. 4) a. АМ. and pretreated for two hours with apte iia dann ын |... monaftalene at room temperature. Sapen E for Bolivian rasses Ку been те orted ош Ь the material was hydrolyzed with a 1 сё ДЕ, ith к P Y " of HCl at 60°C for ten minutes and : Mou a oe 10 aay m F eastern kena while red Messias pus на so oa п Mp EU Д n o EC ERU were scored. А complete list of species, voucher f Сайы specimens for individual accessions, and locality of Santa Cruz. This paper putes complementary nf ided in Table 1; a list of synon information to Killeen’s (1990) study. ee P id Chrom о be ne i T ite Е i Я of all accessions сі я or different by comparison to Killeen (1990), Honfi posited at CTES and ISC; partial sets of illeen et al. (1990), and Dubcosky & Zuloaga (1991) or, diea LPB, MO, SI, US, and USZ. where not specifically cited, from the summary pine рем ingto: i listings of Darl n & Wylie (1956), Ornduff (1967, 1968, 1969), Fedorov (1969), Moore RESULTS AND Discussion (1973, 1977), Goldblatt (1981 1984, 1985, This paper presents the chromosome numbers, " 1988), and Goldblatt & Johnson (1990). as determined by mitotic divisions, for 59 separa 1 This rch was supported de I i Ci y Técnicas (CONIC m the facilities of the аена de pe ncias cae arias, ceni sidad Nacional ea Norde iud Argentina. Fieldwork TEES rted by the Organization of American States and the World Food Institute of -= deis University- * Instituto de Botánica del pose ste, Casilla de im pi den Corrientes Arge! * Missouri Botanical Garden, Bolivian n Program, Cas 4, La Paz, Bolivi ANN. MISSOURI Bor. Garb. 81: 768-774. 1994. Volume 81, Number 4 1994 Norrmann 769 Chro Анадай ae in Bolivian Grasses TABLE 1. Zygotic chromosome number (2л), locality of origin 3 n Ч } LA Б Species 2n Origin, voucher, and figure references Andropogon lateralis Nees 60 Dept. Santa Cruz: Prov. Andrés Iban a Viru Viru E of the International Airport, Доу pee W, 400 m, A leen 1 Arundinella hispida (Willd.) 20 pt. Beni: Prov. Ballivian, gom Biológica del Beni, гт Kuntze Porvenir, 40 km E of San Borja, 14?35'S, 66°30'W, 200 m Killeen 25 93 (Fig. Axonopus chrysoblepharis (Lag 20 Dept. Sant ám Cree Prov. bg de Chávez, Pueblo de Concepción Chase *03'S, 62°10'W, 500 m, Killeen 11A Axonopus compressus (Sw.) P. 40 he Cem Cruz: Prov. Ñuflo de Chávez, Estancia Las Madres, 9 Beauv. "s of Concepción, 16?00'S, 62°00'W, 500 m, Killeen 175 7 Axonopus fissifolius (Raddi) 20 Dept. Santa Cruz: Prov. Nuflo de Chavez, Estancia үн 2 km S Kuhlm. 0 RE 16°08'5, 62°05'W, m, Elionurus muticus (Spreng.) 20 Dept. Pa nta үш Prov. „Мово de Chavez, орн а Мапа, Kuntze k of d LM Lom 16°13’S, 0'W, 500 m, Killeen 2 Eriochloa distachya HBK 18 Dep S ү rt Prov. N uflo te Chive, Estancia Santa Maria, p o Lomerio, 16?10'S, 62°00'W, ees m, Killeen 2441 Gouinia virgata (C. Presl) 40 Dept. Santa Cruz: Prov. Nuflo de Chávez, 17 km N of Concep- Scribn. ción on road to San Ignacio, 15?50'S, 62°00'W, 400 m, Kil- leen 18: Gymnopogon spicatus (Spreng.) 40** шул Santa sana у. Nuflo de Chavez, 3 km S of Concepcién Kuntze oad around reservoir, 16*03'S, 62°10’W, 500 m, Killeen 201 2 Imperata tenuis Hack. 20* -— Баша — Prov. uo de С кару Santa Maria, п to Lomerio, 16°13'5, piss W, 5 00 m, dolls 2482 (Fig. ik Lasiacis sorghoidea (Desv.) 36 Dept. Santa Gus Prov. Nuflo de Chavez, Rancho Puesto Nuevo, Hitche. 40 km S of d to Lomerio, 16°25’S, 62°00'W, 700 е ‘Killeen 2332 (Fig. 3) Leersia hexandra Sw. TOEF Dept. Santa Cruz v. Ñuflo de Chá E Estancia жер, 2kmS of Catania. pa 62°05'W, 480 m, Killeen 2416 Microchloa indica (L. EPE: 12 Dept. Santa Cruz: laser Nuflo de Chive, Comida zd Beauv. Nuevo, т 5 oad to Lomerio, 16°25'5, 62°00'W, 450- 700 m, Killeen 1826 Oryza rufipogon Griffith 24 Dept. Santa Cruz: Pro у. Nuflo de Chavez, Estancia Santa Maria, 8 km S of Concepción on road to Lomerío, 16°10'S, 62°00'W, 500 m, Killeen 2086 Panicum laxum Sw. 40 Dept. Santa bs Prov. Andrés Ibáñez, Pampa Viru Viru, 1 E, c International Airport, 174055, 63*10'W, 400 m, Ed Panicum laxum Sw. 40 Tu [e Cruz: Prov. Nuflo de Chavez, Estancia Salta, 10 km S of C o Lomerio, 16°13’S, 62°00'W, 500 m, ipia 2286 (Fig. Panicum mertensii Roth ex 36 uz: Prov. E de Chávez, 20 km W of Sant Roem. & Schult Е m ^ on road to San Ignacio, 15*50' M 6140'W. 300 m, Killeen 1745 Panicum olyroides HBK 36 Dept. Santa Cruz: Prov. Nuflo de Chávez, Estancia El Recreo, 2 km N of Concepción, 16°02'5, 62°08'W, 480 m, Killeen 2398 (Fig. 5) Panicum trichanthum Nees 36 Dept. Santa Cruz: Prov. Andrés Ibáñez, Pam ampa Viru Viru, 1 k E of the International Airport, 17°40’S, 63*10"W, 400 m, Kil- leen 1556 770 Annals eiiis A) AM Garden TABLE l. Continued. Species 2n Origin, voucher, and figure references Paspalum acuminatum Raddi 40* um п б Prov. Ñuflo de Apicius vndis, 16?20'S, урен oW 0 m, Killeen 2 2. Paspalum erianthum Nees ex 80 T Soma Cruz: Prov. Nuflo de Chavez, Estancia Salta, 10 km Trin. 0 d to Lomerío, 16?13'S, 62°00'W, 460 m, Nm Paspalum erianthum Nees ex 80 i$ heey бл Fier Мово де on Estancia Santa Maria, Trin. omerio, 16?13'S, 62°00'W, 500 т, Killeen 2 ды Paspalum guenoarum Arechav. 40 Dept. vere Cruz: Prov. Nuflo de Chávez, Estancia El Recreo, 2 km N of Concepción, 16?13'S, 62°00’W, 500 m, Killeen 239 Paspalum guenoarum Arechav. 40 Dept. Santa Cruz: Prov Nuflo de Chávez, Estancia El Recreo, km N of Concepción 16?13'S, 62*00'W, 500 m, Killeen 2394 (Fig. Paspalum humigenum Swallen 20 «y Santa su Prov. Nuflo de Chávez, Estancia La Pachanga, km S of icio on road to Lomerío, 16°08'S, 62°05'W, Je "t Killeen 2. ied intermedium Munro 40 cs San a Cruz: tse Мово de Chavez, Estancia Viera, 2 km S ex Morong & Britton d to Lomerio, 16°08'S, 62°05'W, 500 m, К 1631 Бестен E Munr 40 eee na (к Prov. Nuflo de Cet Estancia La Pachanga, ex Morong & Brit o Lomerío, 16°08’S, 62°00'W, 0 m, Kille leen 225 Paspalum kempffii Killeen 40* De. Sa - IUE un Nuflo де ries Estancia La Pachanga, merío, 16°13'S, 62°05'W, 500 m, Killeen 227 Paspalum lenticulare HBK 40* Dept. Santa Cruz drés Ibáñez, Pampa Viru Viru, d of Поета Ани 17°40'5, 63°10'%, 400 m, Ан. Paspalum lenticulare НВК 40 Dept Sant Cruz: Prov. spe de Chavez, Estancia Salta, 1 S of o Lomerio, 16*10'S, 62*05"W, "500 m, Killeen Paspalum lenticulare HBK 40 = сан nA com Prov. Мово ае ee Estanc ia San Sebastián, о, 16°20'5, pie 500 m, Killeen 232, Paspalum Іепіісшаге НВК 40 рер Sana Cruz: Prov. Nuflo de ok Estancia El Recreo, 2 N of Сова 16°13'5, 62*00'W, 500 m, Killeen Paspalum limbatum Henrard 20 Dep. ‘Sant Cruz: Prov. Мово de Chavez, Estancia Viera, 2 km 5 o Lomerio, 16*13'S, 62°05'W, 500 m, Killee Paspalum limbatum Henrard 20 sig Senta on Prov. Nuflo ake — өзөгү? ia La Pa — km o, 16°08'5, 62°05'W 500 m, Killeen 227 276 Paspalum limbatum Henrard 20 ts how: a Cruz: Prov. bue de Chávez, Estancia Salta, S of o Lomerío, 16°08'S, 62:00, 500 m, Killen 2453 (Fig Paspalum lineare Trin. 80* Dept. z а Crus: Prov. [x » Chávez, Estancia La P Pachanga, 5k to Lomerio, 16°08'S, 6205'%. 500 т, K Paspalum macedoi Swallen 40* Dept. San goa 2218 Nuflo de Chavez, ки Las Madres, 9 km N st ncepción on road to San Igna ages aptid, 500 m, Killeen 1796 Paspalum maculosum Trin. 40 Dept. San a Cruz: Pro у. Кий uflo de €— nos Salta, 10 km S of C 13'S, 62*00'W, 500 m, Killeen 2282 (Fig. 8) CS SE Volume 81, Number 4 Norrman al. 771 1994 Chri rehe МА in Bolivian Grasses TABLE 1. Continued. Species 2n Origin, voucher, d fig f Paspalum malacophyllum Trin. 40 үер щш om Prov. Nuflo де eaten im San Josecito, о Lomerío, 16?02'S, 62°05'W, 490 ay Killeen pei Paspalum malacophyllum Trin. 40 e T Cruz: Prov. Nuflo de Chávez, Estancia El Recreo, 2 N of Concepción, 16?12'S, 62°08'W, 500 m, Killeen xs 49 Paspalum minus E. Fourn. 50 Dept. Santa Cruz: Prov. Andrés Ibáñez, Pampa Viru Viru, 1 E uns International Airport, 17°40'S, 63°10'W, 500 m, y leen Paspalum plenum Chase 40 Dept. E Cruz: Prov. Nuflo de ا‎ oo Viera, 2 km SE of Concepción, 16°13'5, 62°00’ m, Killeen 2317 Paspalum plicatulum Michx. 40 Dept. Santa Cruz: Prov. Nuflo de ee т^ cia La Pachanga, 5 f С pció d to Lomerio, 16°08’S, 62*05'W, 480 5 Killeen 244. Paspalum plicatulum Michx. 40 Deni es a Cruz: ve b de Chávez, Estancia Salta, 10 km S of o Lomerío, 16°13’S, 62°00'W, 500 m, Killeen : Pennisetum setosum (Sw.) Rich. 45** Dept. Santa Lo pA Nuflo de Chávez, Comunidad Puesto uevo, 30 km S of Concepción on road to Lomerío, 16°25'S, 62°00'W, m, Killee 2 Pharus lappulaceus Aubl. 24 Dept. Santa pus Prov. Nuflo de Chávez, 17 km S of Concep- ción on Road to San Ignacio, 15?50'S, 62°00'W, 400 m, Kil- leen 18 Saccharum trinii Hack. 60 noni "cy 8 Prov. Nuflo е ien ace La Pachanga, to Lomerio, 16°08’S, 62*05'W, 0 m, Killeen 221 9, са Saccharum trinii Hack. 60 ж Santa Cruz: Prov . Nuflo de Chávez, seid Salta, 10 km S of Concepción, 1613'S, 62°00'W, 500 т, Killeen 2281, cleistogamous form Schizachyrium sanguineum 50 a ak Cruz: Prov. Andrés Ibanez, Pampa Viru Viru, (Retz.) Alston of a oO Airport, 17°40" S, 63°10'W, fiat m, Kil- M Schizachyrium sanguineum 50 Dept. t cR v. Nuflo de Cháve , (Retz.) Alston sr tas west P town, те 62°00'W, 500 m, Killeen Schizachyrium stam: 60 Sese cu Cruz: Prov. Nuflo de Chávez, Pueblo de Concepción, (Retz.) Alsto the laguna west of town, 16*05'S, 62900" W, 500 m, Killeen Sc chizachyrium Du cni 70 Dept. Santa Cruz: Prov. Nuflo de Chávez, Pueblo de Concepción, (Retz.) Ао the laguna west of town, 16*05'S, 62°00'W, 500 m, Killeen Sc true sanguineum 60 Dept. Santa Cruz: Pro v. Nuflo de Chávez, Estancia Salta, 10 km (Retz.) Alston S of Concepción, 16°13'S, 62°00'W, 500 m, Killeen 2095 (Fig. 9 Setaria parviflora (Poir.) Ker 36 Dept. Santa Cruz: Prov. ufo de Chavez, Estancia Salta, 10 km à S of о Lomerío, 16*13'S, 62°00" v, 500 m, Killeen : Setaria 1 lpiseta (Lam.) В 54** Dept. me e "e „Хово де Сан, аа San Sebastian, & Schult 25 km S 0'S, 62°00'W, “500 m, ‘Killeen 2318 aa 10) xi cm minarum (Nees) 20 Dept. Santa Cruz: Prov. Nuflo de Chávez, Esta: eo, 2 Hitch km N of Concepción, 16°02’S, 62°08'W, 490 x m, о 2387 20 en Sante Cruz: Prov. Аиво de Chavez, Estancia Salta, 10 Кт Sorghastrum setosum (Griseb.) Hitche. S d to Lomerio, 16°13'5, 62*00'W, 500 m, ‚ Killeen 2306 (Fig. 1D Annals of th Missouri Botanical Garden TABLE 1. Continued. Species 2n Origin, voucher, and figure references Thrasya crucensis Killeen 20 Dept. ves Cruz: AE Nuflo de Chavez, pee Puesto Nue em o Lomerio, 16?25'S, era: 700 m, , Killeen "23 34 Thrasya petrosa (Trin.) Chase 40 Dept. Santa Cruz: P v. Nuflo de Chávez, Estancia Salta, 10 km S of Mié on а to Lomerio, 16?13'S, 62°00'W, 500 m, Kill Thrasya p (Trin.) Сі 20** Dept. Santa Cruz: Prov. Chávez, Estancia Viera, 2 km 5 £s Nuflo de of Co EA ión on road to tomaid 16°08'S, 62°05'W, 500 m, 418 Killeen * First count for this taxo ** Count different from eiut ones for this taxon. accessions of 43 species in 21 genera and seven f d in Arundinella hispida М (Fig. le in oe with a previous report 8 1967, sub Arundinella con- fn Өл, ) Нее i ice Ап accession of Gym gon spicatus was doc- umented as being tetra e id has or Killeen 7), which contrasts to the as accession (Killeen 2481) reported by Killeen (1990). two accessions were pou within 5 km of one = Ф Imperata tenuis was documented as a diploid with 2n = 20 (Fig. 2); this is the first chromosome count for the species. The base of x = 10 conforms with other reports for the ge Leersia hexandra is a pan d species, and tigators have same population near Concepció leen 2416) was food to contain ан (2п = 72) by the mitotic counts reported he «6 as well as oaase aredi (2n = 48 v by the (1990). aspalum is the most speciose genus in the Noto and is, likewise, the m gen unts are reported for P. na- tum, P. macedoi, P. kempffii, and P. lenticulare, all of which are 2n — 40, as well as for P. lineare, which is 2n — 80. Pennisetum setosum is a neotropical species а previous сш of 2n = 36, 72, and 54 tanloid 2n = 45 cytotype. Nonetheless, plants were , rel- atively fe padi in natu ral po opulations, and pan po lly {ег Presumably the жо or at least this ac coeasion; reproduces via apom a phenomenon that has been reported for vds pes (Dujardin & Hanna, 1984). This is the first salai count p Saec E trinii. À reporiea 60 (Мац, 1981), and iiit or has been v void cleistogamy ос- p SU Both phenotypes — 60 cyto рег Mer г 5с eMsachy ut sanguineum is curred in us d m ed for ‘the ii by Carman & Ж nd this pred scheme may exp these Share polymorphisms Volume 81, Number 4 Norrmann et al. 773 1994 en Numbers in Bolivian rasses г. E у is сл y J — K x ч " 7 РСА ) y уз A „м< à Д 9 ~~ T . ёё, E “у e 4 & yt м + 4 ( ~ - 2 E - ” w к айы 2 * Ө “Ө ‘. Ө ; Tas: ез t [d s ж +” У м 4 Pd mah ia + ري‎ d 4. e ^ " í ao ў 8 m» E. y „к E, Tis Fro рь ^ M ^7 »" i spl ١ S à a * i \ ih. P ' 5 yd oum » “м. г ‚б t ” LÀ By Y iy * (4) s * 2 Ө e Ё РҶ 8 GS "ag = : а | „>, 7 , д e ў * P " ^.. Li -x Lad yr E ^ ME Æ t — M 9 x 7 s „* ы з Beas 6) © № Ө ГД Ф °` = % +. Sy -^ Sr v з Ф ° У ҳ г» „ E ` г v {7 1 rd * on s we x м rs 2% 4 a w %/ъ > ә 05 om \ decr Es ® d o. «= © Ficures 1-11. Photomicrographs of root- -tip squa —1. Arundinella hi tispida, 2n = 3 pue gan zn 2. cvm rata tenuis, 2n — 20 0 (Killeen 2482).— 3. Lasiacis sorghoidea, 2n = 36 (Killeen 2332).—4. Panicum laxum, 2n = 40 (Killeen 2286).—5. Panicum olyroid ys 36 (Killeen 2398).—6. Paspalum кебин 2п (К 0 (Killeen 2394). —7. Paspalum limbatum, 2п = 0, (Killeen 2453).—8. Paspalum maculosum, 40 (Killeen 2 2282).—9. Schizachyrium sanguineum, 2n = 60 (Killeen 2095).—10. Setaria itus, | 2n = 5 (Killeen 2318).—11. Sorghastrum setosum, 2n — 20 eee 2306). Annals Missouri ы Сагаеп Setaria vulpiseta was сосе ю һауе а chromosome number of 2n = 54 (Fig. 10). This differs from previous г mend of 2n — 36 for Bra- zilian populations (de Oliveira Freitas Sacchet, 1980). er species with both Thra. a and tetraploid E (Table By This in the ШЕШ related penus Paspalum i is usually diploids and сок. прера: with sten lg orrmann, ; No orrmann et al. 1989; —€— A Met cytotype is also known for Th. petr from Venezuela (Davidse & Pohl, 1974). It, уч th tet loid fi f } showed t J? d AE Papa eee rs г, . СТС 5 wıtn аро- mixis Several of the included in this study (Axonopus fissifolius, Leersia hexandra, Pani- a antanal savannas of « central Sout h Ameri Kien, 1991a, b). Knowledge of their taxonomy reproductive biology are essential for e improvement yia bre eeding pr programs, as well a Z t © аретепї strategies. LITERATURE CITED BowDEN, W. M. & Н. SENN. 1962. Chromosome bers in 28 grass genera from South America. “berg J. Bot. 40: 1115-1124. & Carman, J. G. E: HATCH 1982. Aposporous apomixis in Sehisach hig (Poaceae: Andropogo- маа : 1252-1255. eeding systems in the grasses: Pie Seo a ш s 131-154. Dae fa D. & A.P. hromosome Atlas of dep Led: pore New es DAVIDSE, G 1 omosome bers, m: s ede and а on tropical oin ican Mine (Ско, Canad. J. Bot. 52: 317- 328 Ene i унаа & В. К. Sm 1986. mosom s of Zimbabwea; nr (Poaceae) and an чыны: E · polyploidy i in the n J. Bot. 52: d ird 8. ZULOAGA. ees Nümeros i-am = ө ш > setum FEDOROV, K 969. Chromosome Numbers of Flowering Plants. mee Botanical Institute “ ы Academy of Sciences of the U.S.S.R. [Керг Scientific Patai, Koeni GOLDBLATT, P. bers, 1975- 1978. Monogr. Syst. Bot. Missouri Bot. © rd. + Index to Plant Chromosom 1984. е Numbers, ` 1979- 1981. Monogr. Syst. t. Bot. Missouri Bot. Gard. ————. 1985. Index to Plant Chromosome Numbers, 1982-1983. Monogr. Syst. Bot. Missouri Bot. Gard. ———. 1988. Index to Plant Chromosome Numbers, 1984-1985. Monogr. Syst. Bot. Missouri Bot. Gard. 25; ———— & D. E. Јонмѕом. 1990. Index to Plant Chro- mosome Nambers, 1986-1987. Monogr. Syst. Bot. . Gard. 3 Gourp, F. W. & T. R. Sop Chromo- ERSTROM. 1967. some numbers of tropical American grasses. Amer. J. Bot. 54: 676-683. Howr, A. I., C. L. N&J FM 1990. Estudios cariológicos еп gramíneas Mw arwiniana 30: 8 d a 30: p KILLEEN, T. J. CHE Cruz, Bolivia. Ann. аве Вот. зы, TU w 201. pump a. Range management and land-use umet in siqua Santa Cruz, Bolivia. Range- lands 13: i x E e Cruz, ur ү Grasslands Es 12-19. A. M. MOLINA, 981. El género Erianthus (Gani) en la fie: y países limítrofes. Darwiniana 559-585. RE y is 1973. Index to Tone gei r 1967-1971. Reg 197 7. Index to Plant E р ЕЕ 1973- 1974. беа Veg. 96. Une Normann, G., C. B. BURSON. pie genetics and reproductive i md 0- me Num 6 тнт umbers Quarin, c. ia 3: 25-35. masc, 07, ا‎ RAVEN, P. H. 1975. The bandi of angiosperm phy hylog- 24- зА у, Апп. Missouri Bot. Gard. 62: 7 CHROMOSOME NUMBERS AND S.M. dere Ahsan, Ahsan A. Vahidy,' INCIDENCE OF POLYPLOIDY IN PANICOIDEAE вое ) FROM PAKISTAN and S. I. A ABSTRACT Ch if x ee Ж] f DLL are reported. С viz. Pan n = - 18) and voee торат * = 10) аге DOW: ботан = icum atrosanguineum (n = 18), disnei t lanat pepe are report rted fe E Buches the 1 4 flora of Pakistan. Моге ep ehe of the species i oth er ғ species are new to t ypl 8 were found rs of Poaceae have been of great interest from both ааыа and cytot omic points view. The rol pores? in the evolution and diversification of grasses has bee stents large. Nearly all genera i a majority of species this family possess chromosome n rs whic are multiple of the original basic icd ai Ace. ing to ges (1950, 1956 ), polyploidy ha ү f id of fee оѕоте number ers As gne s. Further, he estimated nearly 70-75% of grass species to be pues cede EE сои 63.41% poly- ploidy the Hind dayan Lowe pene E ) 7 or of the f. 3 S 4 | Pakist b the present study the ievel ме т dy is discus За in the light of av. information about р. I Some counts jn dE from Pakist MATERIALS AND METHODS For meiotic preparations young, unopened m- 1:3 acetic babel and Stored at —4°C, and the anthers were squashed in 1.8% aceto-orcein. For mitotic preparations young and healthy root yr s from germinating seeds were sage кч 2 М 8-hydroxyquinoline for 4- in acetic те bird in in ү N HCI for 6-12 min. a ed d 18% aceto обе; T lies pus OBSERVATIONS AND RESULTS able 1 lists ck bers for 126 re- cords рове: 58 taxa of 27 genera. Counts new to science and new to the flora of Pakistan are e vey of IPCN (Fe- n the basis of dorov, 1974; Goldblatt, 1981, 1984, 1985, 1988; Moore, 1973, 1974; Ornduff, 1967). Ploidy level NN CJ 1 f, 1 1 1 dala h in the genus. of a of 58 taxa investigated, id (67 | eet eel o be polyploid. The majori e the po ape v were at tie tetraploid 1 ка. x ight hexaploids, taxa were observ In Table 1 ue genera are arranged in a (Cope, 1982), are deposited in Karachi University UH). Herbarium (K DISCUSSION investigated species belong to three tribes The of Panicoideae. In Pabati me tribe Isachneae is himalaica ig 19). Previously the = penta of this species 2n = was г у Mehra (1982) and Parkash (1979). The on ue here i is the J ing iis permanent in Feuparal or Canada balsam. M een and Z. A. Razaq of Department of Botany, B , for DN cytological material. This paper is par - The tribe Paniceae is one so the largest tribes niversity of Karachi, for their help in А. Razaq, Moinuddin, and B. Jahan of the t of the work 1 We thank mrs n of Rb Bs e also ph HOMES S. Omer, T. Ali, partment of Botany, Des Supported by National ee aee gm INT 510 partment of ОШЕН of Kar 0, Pakista ° Department of симе University of Kara rac chi, Karachi- 0 Жыш. ANN. Missouri Bor. GARD. 81: 775-783. 1994. Annals of th Missouri lr Garden TABLE 1. Ch і і ploidy level i leae (P ) fi Ра} K.U. = Karachi University. D. G. Khan = town of Dera Ghazi Khan. Chromosome Ploidy Taxon no. n level Voucher Tribe Isachneae *Isachne himalaica Hook. f. 20 Tetraploid Sargodha: Ghafoor 3848 (Fig. 19) Tribe Paniceae gemis e (Schumach.) 18 Tetraploid Hazara: Omer 2222; Dir: Ghafoor 2331; D. б. ard ex Robyns Khan: Ghafoor 3692 (ы. ig. P *Brachiaria eruciformis (Sm.) 9 Diploid K.U. Campus: Ah 65; S Sal T. Ali ta 17. ш ramosa (L.) Stapf 18 Tetraploid K.U. Campus: Jahan 57 Brachiaria ramosa (Fig. 2) 36+3B Octoploid K.U. Campus: sabes 58 *Brachiaria reptans (L.) Gardner 7 Diploid K.U. Campus: Ahsa дин 48, Razaq 128; Hubbard (Fig. 3) Head Rajkan: Glar 33 Cenchrus biflorus Roxb. (Fig. 4) 17 Aneuploid K.U. Campu tg an 4 Cenchrus biflorus 16 Aneuploid Mianwali: Т. 811 *Cenchrus ciliaris L. 18 iploid met? гуе eu 3747; D. G. Khan: Ghafoor Cenchrus ciliaris 17 Aneuploid K: i oth mpus: Pep 133; Makran: T. Ali 835; Attock: Ghafoor 2268 Cenchrus pennisetiformis Hochst. 18 Diploid K.U. Campus: Siddiqui 2 & Steud. ex Cenchrus setigerus Vahl 17 Aneuploid Safari Park, hoy Siddiqui 63; D. G. Khan Ghafoor 3603; Mi gne T. Ali 1824; Dar- sanochano: тр? iddi 8; K.U. Campus: Sid- diqui 50; Kathore: ie 66, 78; Manghopir: Raz * Digitaria ciliaris (Retz.) Koeler 27 Hexaploid K.U. icu bea 32; Razaq 146 Digitaria ciliaris 36 Octoploid Kashmir: Т. A *Digitaria nodosa Parl. (Fig. 5) 9 iploid K.U. Campus: Moin 68 * Digitaria setigera Roth ex Roem. 36 Octoploid Hazara: Omer 2 ult. *Digitaria stricta Roth ex Roem. 18 Tetraploid Pail: Ghafoor 3791 & Echinochloa colona (L.) Link 9 Diploid Dir: Ghafoor 4093 Echinoc olon 18 Tetraploid K.U. Campus: Jaha Psa ites 27 Hexaploid Kashmir: T. Ali 400; Ed ij Ан 836; Zhob: li 1087; K.U. Campus: Ahsan 7; Baltis- tan: Omer 2539 *Echinochloa crus-galli (L.) Р. 27 Hexaploid Kashmir: T. Ali 46 inochloa crus-galli (Fig. 20) 45 "к: "ey oa frumentacea Link 27 а Lag ensis (Hochst. & 9 Steud. aes Eriochloa с 18 *Eriochloa esen KP C. E:..9 Erio cer 18 Panicum antidotale Retz 9 **Pani trosanguineum x A. шет (Fig. 6) *Pani n Jacq. (Fig.7) 9 * Panic: cum miliaceum L. 1 *Panicum repens L. (Fig. 8) 9 Decaploid Hexaploid Diploid Tetraploid Diploid Tetraploid Diploid Tetraploid Diploid Tetraploid Diploid Thatta: Siddiqui 153 Khushab: Ghafoor 3796 Campus: Moin. 52 U. Campus: Ahsan E U. Campus: Moin. P^ 50 Se ا‎ Ahsan 21; Thatta: Siddiqui 121 . Campus: Razaq 123 ые Ghafoor 3823 6 Hasilpur: Ghafoor ‚ч йк Т. Ali 181 Bahawalpur: Syed Makran: Ed Volume 81, Number 4 1994 Ahsan et a Chromosome Numbers in Panicoideae TABLE 1. Continued. Chromosome Ploidy Taxon no. n level Voucher *Panicum turgidum Forssk. 9 Diploid — T. Ali dei HM: Siddiqui 3: Ka zm vus *Paspalidium flavidum (Retz.) A. 27 Hexaploid ibus Ghafoor un Camus. Paspalidium geminatum 9 Diploid K.U. Campus: Moin. 69, 59 (For set ) Stapf (Fig. 9) *Paspalum dilatatum Poir. 30 Hexaploid -— Vaga 2591, 2271; Rawalpindi: Ghafoor *Paspalum Me (Michx.) 30 Hexaploid K. " Hm mpus: Jah „ 41; Swat: Ghafoor Scribner (Fig. 10 3334; Sukkur: бн 3492; Tuis: Sid- diq *Pennisetum flaccidum Griseb. Hexaploid Dir: Chafoor 24 *Pennisetum glaucum (L.) R. Br. 7 Diploid K.U. Campus: А 73 BR en tum Klotzsch 18 Tetraploid Swat: Ghafoor 3429 (Fig. и orientale Rich. 9 Diploid Soon Sakesar: ^ pos 1624 Pennisetum orientale dde XA 18 Tetraploid Makran: T. Ali Setaria intermedia Roe 18 Tetraploid K.U. Campus: on 76 Schult. (Fig. 21) *Setaria pumila (Poir.) Roem. & 36 Octoploid Vehari: Ghafoor 3598; Chitral: Ghafoor 2493 *Setaria verticillata (L.) P. Beauv. 9 Diploid po Moin. 70; Safari Park, Karachi: gem Tribe ланса Арша m 10 Diploid an: a 3711 * Arthraxon Pc (Trin); 9 Diploid iion T. A Hochst. Arthraxon prionodes (Steud.) 18 Tetraploid Soon Sakesar: T. Ali 1724, 1701 Dandy (Fig. Arthraxon prionodes (Fig. 13) — 10 Aneuploid Hazara: Omer 2265 Bothriochloa ischaemum (L.) 20 Tetraploid Gilgit: Omer 2617; Chitral: Ghafoor 3218 еп Capillipedium parviflorum (R. 20 Tetraploid Kashmir: T. Ali 266 Br.) Stapf Chrysopogon aucheri (Boiss.) 10 Diploid K.U. Campus: Moin. 1; Safari Park, rap Stapf Siddiqu 67; Kathore: Jahan 86; Makr Ali Chrysopogon gryllus (L.) Trin. 10 Diploid Swat: Guo 3431 subsp. echinulatus (Nees) T. G hrysopogon serrulatus Trin. 10 Diploid Hazara: Omer 2267 (Fig. C айай jwarancusa (Jones) 20 Tetraploid Safari pe — Siddiqui 73; K.U. Са Schult. us: Jahan 55; Choa Sayyadan Shah: T. Ali *Cymbopogon martinii(Roxb.) W. 20 Tetraploid Kashmir, T. Ali 207 Watson екен ene 20 Tetraploid cog idee 14; Attock: ard 2275; Gha- (Forssk.) foor 2276; K.U. Campus: Moin. 4, Jahan 54 ан jovclaum (Delile) 20 Tetraploid K.U а Ahsan 57; Kamir T. Ali 35 berty (Fig. 16) Attock: Ghafoor 4232 arte nurus угат Nees ex A. 10 Tetraploid K.U. Campus: Moin. 65 Rich. (Fi, Diploid K.U. Campus: Ahsan 30 ig. 17) *Hockelochióon granularis (L.) 7 Kun E MES E 778 : Annals Missouri ain Garden TABLE l. Continued. Chromosome Ploidy Taxon no. n level Voucher beris t contortus (L.) P. 10 Diploid Pail: Ghafoor 3858; K.U. Campus: Moin. 38 Beauv. ex Roem. & Sc hult. (Fig. 22) Heteropogon contortus 20 Tetraploid : T. Ali Lasiurus scindicus Henr. 9 Diploid Cholistan: Ghafoor 3545; Makran: T. Ali 829; K.U. Campus: Razaq 167, Sid. diqui 46 Phacelurus speciosus (Steud.) С. 10 Diploid Chitral: Ghafoor 2705 E. Hubbard Saccharum m Munro ex 10 Diploid D. G. Khan: Ghafoor 3616 Boiss. (Fig. Saccharum pee 1. 27 Aneuploid Sargodha: T. Ali 1667 (Fig. Pies bicolor (L.) (Fig. 18) 10 Tetraploid kran: Ome a Sorghum halepense (L.) Pers. 20 Octoploid irt Levi ; Muzaffargarh: Ghafoor a Doni eia 4247; K.U. Cam Vetiveria zizanioides (L.) Nash 10 Diploid Sialk ot: Lon va *Zea mays L. 2n = 20 Diploid Darsanochano: туун) 94 * Count new to flora of Pakistan. ** Count new to science of the grass family. In Pakistan it is represented nus, with n = 7 and n = 9. Pennisetum lanatum by 15 үтү а pe 13 species (Cope, 1982). Species (Fig. 11) his been reported for the first time to be of 11 of t era were studied. Brachiaria tetraploid (n = 18) reptans i n (Fig. 3) and B. eruciformi The base number for the genus pcs : x= = 9 are diploids based on an 9. In the present study three species, namı у, 5. respectively. These counts are new to flora of intermedia (п = 18; Fig. 21), S. pumila fe V Pakistan and conform with earlier counts reported and S. verticillata (n = 9) show great cytological from other regions (Malik & Mary, E Chris- variability. Baquar and Saeed (1969) reported п topher & Abraham, 1976; Bas. yam- = 9, while Raman et al. (1959) reported n = 27 ma, ud Sos: 1982). Brachiaria doles for S. intermedia, but we are ne with n — 8 (Fig. 1), is tetraploid on the basis of ploidy level, i.e., n — 18, for the same species - ay ramosa we observed three B-chro- 5 akistan the tribe Andropogoneae is repre- alents mosomes in ipid to 36 biv by 36 genera and 67 species (Cope, 1982). Both n and 8 are Меры ts i in the Ду ^B present b. species belonging to LL enus Cenchrus As far as the 8 consi ‚ Love and Lóve (1961) regarded x = basic number is era of the tribe have been inv vete cytologi- cally. In Arthraxon prionodes, n = 0 (Fig. 13) 17 as dps ng ei number » while Ba quar and Anjum and n = 18 (Fig. 14) are reported he number (196 aber. Thou; which may be hyperancuploid m Mid i^ with number, s fo mdi in other genera ef ‘the tribe, n = respect to the bas 9 is Ый unknown in the genus Cenchrus. The ies count for Колин көрз Therefore, in our opinion, the basic number for (Fig. 17) is reported here for the first е. We this genus is x = 18, r reflecting an early tetraploid believe that it is a tetraploid species (n = 19) This шыу; ‘eee - zi" is an aneuploid descendant from is eios ko the report (Dujardin, 1978) tha xw А A new chromosome count for Panicum atro- The basic ru faethe "gm e Mi. au 8 x n (п = 1 Fig. 6) revealed pe itisa = 9. The present and previous rts (Faruqi tetraploid species. 1979) both indicate the CM S of only pir AL. ennisetum is agronomically an important ge- populations in Pakistan, whereas the count given COIGeae =| Ro AR نہ‎ ^" ~ 5 Ў Eb = = о є 73 3 ees . с So һо v M © Annals бан: ани Garden 7-12. pon Rd gen meiosis in members of Мустела, ا‎ — 1. Panicum maximum ( (Gha pe ap dg Чан п= 8. Pani tum (Moin. repens (Omer Y diakin = 9.—9. Pas palidium geminatu 69), diakines —10. Pa орет Far ашалы (Jahan 40), celal "n ei . Pennisetum lanatum (Ghafoor 342 9), pian п = 18.—12. Pennisetum orientale (Т. Ali 1006), NAM de n = 18. Volume 81, Number 4 Ahsan et al. 781 199 Chromosome Numbers in Panicoideae 3-18. Pollen mother en meiosis in me apre embers of Panicoideae (Poaceae).—13. Arthraxon prionodes (Omer 65) any mee In=10.— sies я онат (T. Ali 1724), diakinesis л = 18. — 15. Chryso, vtr "n lm (Omer 2267), diakinesis n — . Dic —17. Elionurus itas (Moin. 65), مارا‎ п = 10.— a ium foveolatum (Ahsan 57), diakinesis n — 8. Sorghum bicolor (Omer 2043), metaphase-I n = 10. Annals sai d Garden T А 2 jM # ۹ 23 Ро (С infor 3 Poe apace n= 4 24 llen кыс у” Калын їп members of Panicoideae (Poaceae). gis Is Е о мэ. | $ ұу у х. E UE achne himala xS xs —2!. мей ила — dia (Ahsan 76), di Mena n= teropoeon lli (Siddiqui 153), м Жашо d Mea griffithii (Ghafoor 3616), roe emper n= “10, by Celarier (1957) indicates the presence of an- euploidy at hexaploid level. е present investigation of polyploidy in Pan- icoideae from Pakistan demonstr. CELARIER, R. P. 58), à n —24. Saccharum picis (T. Ali 1667), pe LITERATURE CITED tan BAQUAR, S. R. 6. Polyploidy in the Flora of des in relation "s EAM. life form and taxonomic Taxon 25: 621-627 | е behaviour 1969. Chromosome agi Cench pans uthern West Pakistan Fore ari вае XI(3-4): 288-296. a ee RNC 1969. Chro С = the Andro- 957. ы Volume 81, Number 4 1994 Ahsan et al. oo Numbers in Panicoideae рококо ае. П. Subtribes Ischaemineae, Rottboelliinae, d the ve ned Nos) 22: 160-183. Msc on the y VI: Panicoideae, tribe Penis eae. Cie 21 “вэт, EA 982. Poaceae. In: E. Nasir & S. I. Ali т, Fora. E West керд 143: * ades DUJARDIN, M. Chrom: of -— tropical ү кз. grasses from Y Western on. Сап J. Bot. 56: 2 ALAL 1979. Chro- and some morphological aE n. Cy- mosome ristics e And tologia 44: 585-605. FEDOROV, A. A. (editor). 1974. Chromosome Numbers of Б Plants . Otto Koeltz Science Publishers, West any (reprinted). eem i Ж (йш), 1981. Index to Plant Chro- Numbers 197 = 1978. Monogr. Syst. Bot. Missouri Bot. Gard Pl 1-553. ——. 1984. Index Mn bl Maca ies Bot. Missouri Bot. Gerd ine Gd 1985. Numbers EN ipia ou М, Bot. Missouri Bot. Gard. 13: 088. ndex to Plant Chromosome Numbers 1984- 1985. Monogr. Syst. Bot. Missouri Bot. Gard. 1-264. Love, А. & D. Love. 1961. Ch b f Central and Northwest European plant species. Opera Botanica 5: 1-581. C. P. & Т. N. Mary. 1970. Chromosome Number Reports XXVII. Taxon 19: 437-442. EHRA, P. + ee Cytology of East Indian Grasses. Р.М а, Chandigarh. gei: R, T EDS 1973. Index to Plant Chromo- Numbers for 1967-1971. Regnum Vegetabile 9. 90: 15 974. Index to Plant Chromosome Numbers 1972. Regnum Vegetabile 91: 1-108. окчу, R. (editor). 1967. Index to Plant Chromo- е Numbers for 1965. Regnum Vegetabile 50: 1- 12 PARKASH, о. 1979. Cytological Investigations in Some Grasses of Northeastern India (Festucoids, /schae- „ Eulalia Kunth and Saccharum spon- neum Linn.). Ph.D. аг MI. 9. Chromosome numbers in Gramineae. Curr. Sci. 28: ony 454. SHARMA, М. L. 1985. Climato-geographic cid sod "m in the Himalayan Grasses. Cytolog 483- STEBBINS, С d 1950. Variation E к! in Plants. Columbia Univ. Press, New 1956 Cytogenetics aa ик of the grass family. Amer. J. Bot. 43: 890-905. CHROMOSOME COUNTS IN M. Moinuddin,* Ahsan A. Vahidy? and S. І. Al? POOIDEAE (POACEAE) FROM PAKISTAN! ABSTRACT du At { 85 } viz., lad Ses Chloridoideae, and poses {> Ва, аге aan The chromosome counts for iz. ix species, : Leptothrium senegalense (n = 10); Piptatherum gracile mam: Pos sinaica (п = 7); ‘Sporobolus arabicus u = 18), ай nervosus (п = red ЕЕ ошого 18); a (n 4 species are new for the flora of Pakistan. P us cyto types were detected in Aeluro, ji possis (n = 20), pes barbata (n = 21), Diplachne fusca (n = 20), and Ochthochloa compressa (n = The Шен аге опе ОЕ Ше largest a of 000 ering plan species. Its me siis are * widely disperse a in all regions Aud the world, phis de almost every type mina eget Baquar & Saeed, d, 1969; Кап я al., 1979). The rs of 45 species belonging to 29 genera in three subfamilies. MATERIALS AND METHODS or meiotic counts, young unop ened inflores- ences were fixed in Carnoy's Bus (3 part absolute alcohol: 1 part ads acetic acid) for a least 24 hr., then stored at lides were prepared = ordinary squash “он using 1.0% acetocarm .8% aceto-orcein. For mitotic counts, young and healthy Maus upa with 2 M 8- hydroxyquinoline -— 4— б hr. ) or with ids water (at 1—2°C for solution for 1 hr., менне in 1 N HCl for 6. 2 т t 60°C, and squashed іп 1.8% aceto- orcein. ost of the cases photomic crographs were taken from барчы, peper ions. Desired slides were made p manen euparal or Canada da bal- sam. Voucher specimens are deposited in Karachi University Hellas (KUH). OBSERVATIONS AND RESULTS fe the bas: cies mre new to the fy ra pu Meiner as they we IPCN (Fedorov, 1974 Cui 1981, 1984, pes 1988; Moore, 1973, 1974) DISCUSSION In the pea study most of the uen counts that conformity with earlier геро! have not ы, commented upon tribe Aristideae, counts oe 22; п sepe plumosa ч = Fig. 1) are reported here for he first tim lago- In the nre Acbioporieae: Mo > if p (n — 20, Fig. 20) is d to be tetr vid m the basis of x = 10. oe and penta k M. Qaiser A. Raz: nds a of Karachi, i kd dep in the eran of start d N. of the De im hank A. ‘Ghat г, 5. (тае, iz. A pr mt arachi, К scie cting wie material. 318. achi, Karac hi- 75270, Pakistan’ ANN. MISSOURI Bor. Garp. 81: 784-791. 1994 Volume 81, Number 4 1994 mai et al. Counts in стт tee чей and Pooidea ЧЫ isi 1-6. Pollen mother cell meiosis ph via п = 22. aa M gres fusca anion , diakinesis n robolus arabicus MAT А ight D diakin f Poaceae.—1. Stipagrostis plumosa nae r 2776), ne esis n = 20.—3. T ^: li el “metaphase n 68), dia ud n = 10 chloa com, a (T. Ali Spo einig nervosus (Siddiqui 131), Ms n= 18. —6. Leptothri individuals have previously been dir for this a Species (Sindhe, 1980; Baquar & Saeed, 1969). i tr ragrostideae, chromosome n rs fo n species (belonging to 6 genera) were deter- оа. In Зач aegyptium, we found с бы іп оп еѕѕіоп. Оп е basis of саа К numbers (х = 10, 12) poneupl ba ba 20), in two ый specimens of dcum. indicate the presence of cytotypes in this . scin- onm (1963) had reported 2n — 20 (2x) for 786 Annals of the Missouri Botanical Garden TABLE 1. Chromosome numbers in Poaceae from Pakistan. K.U. = Karachi University. D. G. Khan = town of Dera Ghazi Khan. Chromo- some Basic number number Taxon n x Voucher Subfamily Arundinoideae Tribe Arundineae *Schismus barbatus (L.) Thell. 6 6 Chitral: Ghafoor 2693, 3071; Makran: T. Ali 834 (Fig. 13) Subfamily Chloridoideae Tribe Aristideae * Aristida adscens L 11 11 Makran: T. Ali 945 * Aristida funiculata Trin. & 11 K.U. Campus: Siddiqui 4 . (Fig. **Stipagrostis plumosa (L.) 22 22 Chitral: Ghafoor 2776 ex T. (Fig Tribe Aeluropodeae ре lagopoides 2 20 10 Thatta: Siddiqui 150, 119; K.U. Campus: Moin. 71 x Thwaites (Fig. 20) Tribe Eragrostideae Dactyloctenium aegyptium 23 10, 12 = 5 Campus: Raz L.) Willd. 20 U. Campus: Moin is ak Ghafoor 3744 * Dactyloctenium scindicum 10 k U. Campus: Moin. Bois tylocte enium scindicum K.U. Campus: Ahsan 56 мы retroflexa (Vahl) 10 10 Khushab: Ghafoor 3836 CRM ое (L.) P. 20 10 Thatta: Siddiqui 101 Beauv m & Schult. (Fig. 2 Eleusine indica (L.) Gaertn. 9 9 K.U. Campus: paro а Swat: Ghafoor 3977; Islama- bad: Ghafoor 3 Eleusine indica 18 M : Omer 2 P ал, cilianensis (АШ) 10 10 K.U. Campus: Moin. 42 Lutati ex F. T. Hubbard Él japonica (Thunb.) 10 Lahore: Ghafoor 4358 Trin *Eragrostis minor Host 10 Hasilpur: Ghafoor 3583 Eragrostis 20 Sargodha: T. Ali 1719 Eragrostis persed (L.) P. 20 Sakesar: T. Ali 1605 Beauv. *Ochthochloa compressa 10 10 Makran: T. Ali 752 Б, Н Hilu (Fig. 3 hloa compressa 20 Makran: Omer 2166, T. Ali 710; Safari Park, Kun Ahsan 38; RH. Campus: Razaq 144; D. G. I Ghafoor 360 Tribe Chloridese Chloris barbat: 0 10 K.U. Campus: Siddiqui e Sajawal: Ahsan *Cynodon dac ie (L. )Pers. 9 9 K.U. Campus: Siddiqui 31, 76, 90, Moin. is Dir: (Fig. Ghafoor 2491 on nis tenellus (Koe 10 10 K.U. Campus: Moin. 53 xb.) Chiov. (Fig. : 2) "Тш villosus Des 10 D. С. Khan: Ghafoor 3699; Makran: Т. Ali 969 Tr О ШЕ Volume 81, Number 4 нвн еї 787 1994 Chr on unts in din cid сна. апа Рооідеа ТАВІЕ 1. Continued. Chromo- some Basic number п т Тахоп л x Voucher Tribe Sporobolea: Hiper arabicus Boiss. 18 9 Makran: T. Ali 842, 846; Cholistan: Ghafoor 3550 (Fig. "Sporobolus coromandelianus 18 K.U. Campus: Moin. 56 (Retz.) Шш; diander (Retz.) Р. 18 Pail: Ghafoor 3776 **Sporobolus nervosus Hochst. 18 Thatta: Siddiqui 131; Darsanochano: Siddiqui 80 (Fig. 5) Tribe Zoysieae Ee senegal 10 10 K.U. Campus: Ahsan 68 (Kunth) W. D. Gavin ‘Fig. Tragus berteronianus Schult. 10 10 Hazara: Omer 2221; cxt T. Ali 1856 p. roxburghii Panigrahi 10 K.U. Campus: Moin Subfamily Pooideae ibe Poeae me tylis E j 7 7 Eua al: acum ne 3286 actylis glome. 14 Dir: Ghafoor 24 ooi et ula (Е T Chitral: Ghafoor d 7) *Poa annua L. 14 7 Chitral: Ghafoor 2430; Kashmir: T. Ali 573; Kalam: Ghafoor 3422 **Poa sinaica Steud. (Fig. 8) Т Hazara: Omer 2720 Tribe Glyc *Ghyceria ia gie (Fries) Fries 20 5 Chitral: Ghafoor 2850 Tribe Aveneae *Agrostis munroana Aitch. & 21 7 Hazara: Omer 2778 Hemsl. (Fig. * Agrostis viridis Gouan (Fig. 21--2B Dir: Ghafoor 2337 9 Cn viridis 14 Chitral: мра 2698, 2513, 3325, 2608 *Alopecurus arundinaceus 14 7 Hazara: Omer 2722 Poir * Avena н Pott ex Link 21 7 Gilgit: Omer 2559 (Fig “Phalaris minor Retz. (Fig. 14 7 K.U. Campus: Jahan 19 › eg —— (L.) 14 7 ear i at m rid K.U. Campus: Siddiqui Desf. (Fig. 11) Makran: Omer Stipeae **Piptatherum gracile Mez 12 12 Chitral: Ghafoor 3093 (Fig. 12 : *Piptatherum munroi (Stapf) 12 Gilgit: Omer 2464; Swat: Ghafoor 3412; Dir: Ghafoor Mez 2407 Tribe Bromeae iae cau Trin. 7 7 Chitral: Ghafoor 2916 (Fig. ha us pectin Thunb. 7 Gilgit: Omer *Bromus ENE д (Fig. 16) 7 Chitral: eisdem dem Annals of the Missouri Botanical Garden TABLE 1. Continued. Chromo- asic number number Taxon n x Voucher Tribe Triticeae *Elymus dahuricus Turez. ex 21 7 Gilgit: Omer 2570 *Elymus semicostatus (Nees 14 Chitral: Ghafoor 2707, 2975 ex Steud.) Melderis (Fig. 17) *Hordeum vulgare L. (Fig. 2n 214 7 K.U. Campus: Moin. 31 (seed obtained from NARC, Is- 18) lamabad) * Count new to flora of Pakistan. ** Count reported for the first time. ts in Arundinoideae, Chloridoideae, and Pooideae Volume 81, Number 4 Мету еї al. 789 1994 Ficures 13-1 Chromosomes in members of Poaceae.— 13. Schi. ), diakinesis n R = К Cynodon Parution (Siddiqui 90), metaphase a ak = Mee ч» тө danthon Gs na 2916), se-l n = 7.—1 pelis omus tectorum (Ghafoor 2606 г и us pure n (Ghafoor 27 70 т п = » 14 8. Hordeum vulgare (Moin. f 1 ^ € ados 2n = 14. th & CH Cee Diplachne pos whereas our erg showed n ported p ly (Hirema g ъа wis ). This 4x cytotype in D. fusca is a 1982). One о the йө esent reports of л роп. (diploid) for species establishes a new dd Ind Ochthochloa compressa only tetraploids were — level (Fig. 3). poss 7-12. Pollen mother cell meiosis їп members of Poaceae.— 7. Lolium rigidum (Ghafoor = S Ts Te sinaica ind Pride diakinesis n = 7.—9. Agrostis prem d eds r 2337), diakinesis. [Ar indicate Jn B.—10. Phalaris minor eke п = 14.— n. Doiie pe 5 “(Ghafoor 2598), mia is JI 2 = 14.—12. PIN pitas rca 3093), diakinesis n 12. А һе Missouri Botanical Garden JRES 19-23. Pollen gene cell meiosis i Ae b- 55€ үг апарһазе-Ї n = E genus bee eie ics agis goes in two species were observed. The count for rapogon tenellus is con dE е spon п = 1 ngh & Godward, 1960), h т our unt agrees with basic number (x = 1 tay able 1 0) f or the ‚Жш. 21). Сһгот е numbers of four species of Spo- robolus were determined in the tribe Sporoboleae; is f Poacea 5 — 19. ا‎ е (Siddiqui 4), pus es лыны (Siddiqui 150), "ашаны; п 20. pica Agr is munroana (Omer 2 —21. gon te nellus dre e 776), diakinesis n = 21.— a barbata 8. The counts for Spo js - е Т 1, P 4) and S. pee a 1, Fig. 5) are reported as new to scienc 5 In Zoysieae, AME bip міс three эй cies belonging to с unt for Lepto iu um ч елери е (Table 1, hig- oi is reported for the firs In subfamil st t rs y Pooideae, ашы numbe Volume 81, Number 4 1994 Moinuddin Chromos ies ip in loe Chloridoideae, and Pooidea for 19 species (belonging to ee genera) in six tribes were determined. Th or Poa sinaica (Table 1, Fig. 8) is reported for e first time. Chromosome 5 for six species belonging Ч in Aveneae. In = 21) is reported for Pa first ribe е the count for Piptatherum gracile (Table 1, Fig. 12) is reported for the first time. LITERATURE CITED BAQUAR, S. в & V = NJUM. 1 ak Chromosome be- haviour in Cen p rom southern West Pakistan. т чуо дн Асїа Biologica. XI(3-4): 288- Жз M. SAE 1969. Chromosome studies and рор polyploidy soli in pan of West pedis 1: yolog 103-1 Core, T A. T2 Род: In: Е. Nasir & 5. I. Ali (editors), Flora of т гче. е 1-678. FaRUQI, S. A., BON н & N. 1979; o- трое ers aad some елаз olo gical charac- eristics of n еда. acm from Pakistan. Cy- uL 44: FEDOROV, A. A. pens 1974. Chromosome Number of ee Plants. Otto Koeltz Science Publishers, West Germany (reprinted). беи ЗЕ ; ed tor) 1981. Index to Plant Chro- Numbers T Т Monogr. Syst. Bot. н Bot. Gard. 5: 1- . Index to Plans Chrom ~ 1979- 1981. Monogr. Syst. Bot. Missouri Bet Card. 8: 1-427. ——— —. 1985. Numbers жей by i Moog. Seat Be Bot. Missouri Bot. Gard. Index to Plant Chromosome Numbers id vaga 1o. Monogr. Syst. Bot. Missouri Bot. Gard. popa HIREM ATH, S $c & M. S. agen hae 249 togenetical studies in the u of Eleusine (Gramineae). Cirpi Зо: к LARSEN, К. 1963. Studies in the Flora of Thailand. 14. al studies in vascular plants of Thailand. Bot. Ark. 20(3): 211-275. US Lo 1973. Index to Plan o- rs for 1967-1971. Regnum ra 90: baa ————. 1974. Index to Plant upon Numbers for 1972. epe Neral As I= SINDHE, A. N. R. 1980. Chro Кылы Reports 45. . Gopwarp. 1960. tapes studies in the Doni: Heredity 15(2-3): 199. LLIS, J. C. 1973. A Dictionary jd the erate Plants and Ferns. Cambridge Univ. Press, Cam- bridge. (8th Өн revised by Airy Sha aw. ) CHROMOSOME NUMBERS IN SOME TAXA OF FABACEAE MOSTLY NATIVE TO PAKISTAN! Bushreen Jahan, Ahsan А. Vahidy,' nd S. I. Ali? ABSTRACT Chromosome ee are pn for 60 taxa belonging t 33 (ен. of Fabaceae; 44 of bo 94: viz.: Pakistan, while the are cu ultiv: nj: native t Argyrolobium steno 13); Ast тош cuspis (n ulea var. rulea (n — caer reported for D. Hn time. Counts for 35 additional taxa are ragalus ee T$ С yllum hesneya pa arviflora (n — ia Jin 8) o es mollis subsp. ай, Ali (n = 9); and Taverniera етеш plants native to Pakista In Pakistan, Fabaceae, the third largest family of flowering plants, is ене by 104 genera and 514 species (Ali, 1973a, b, 1977) of which 35 species are new records from this country. MATERIALS AND METHODS The meiotic and mitotic materials (i.e., the floral 8 = a e S co [79 > o 8 < @ a. © Ё a | | Ф qs o 5 Dn © 9. "d e x 5 л = > Т меге ргераге t For mitosis, root tips Ron oe — were ааа ith 0.0 ( › hr.), fixed in Carnoy’s solution for 1 hr., hee. aye in 1 N HCI for 6-12 min. and squashed 1.8% aceto-orcein. Most oe were taken from tem- ara ma ^od ~ 7), are deposited in Karachi University Her- barium (KUH). OBSERVATIONS AND RESULTS Ch ts for 60 taxa in 17 tribes of фе аге реле in Table 1. Counts new to nce and new to the flora of Pakistan are spec- ifed on the gi asis of a survey of IPCN (Moore, sie uei: , 1974; or cas men 1984, The the arranged a cording to кы (1981а). Discussion In the present study most of the chromosome counts that a ith earlier reports has n = 9 (Fi ig. E} adel here for th This accords wi dun genus in which n also been found Parle MEE mo Tephrosieae, Milletia peguensis has responding with the basic п rx- Mir by Goldblatt (1981а) for the tribe. Tephro п 1 1 We thank S. Khatoon, Z. A. paper is a part of the work supported b е * Department of Botany, Univ ersity of Karachi, Karachi- chi, Karachi-75270, Pakistan * Department of Genetics, of Kara University Omer Raza ro VT. A of Botany, [o of Kaa chi, for ы il i in "identical of voucher uddin, and N. Ahsan of the Departmen s and collecting cytological material. This 8510318. li, M. Moi ANN. MISSOURI Bor. Garp. 81: 792-799. 1994. Volume 81, Number 4 1994 Jahan et al. 793 Chromosome Numbers in Fabaceae TABLE 1. Chromosome numbers in Fabaceae. K.U. = Khan. Karachi University. D. G. Khan = town of Dera Ghazi Chromosome Basic number number Taxon Voucher Subfamily Caesalpinioideae Tribe Caesalpinieae Parkinsonia aculeata L. (culti- 14 14 K.U. Campus: Siddiqui 36 vated) Tribe Cassiea assia =% osericea Frese 14 14 K.U. Campus: Siddiqu *Cassia occidentalis L. iieri 14 K.U. Campus: Jahan 8 Hasanabdal: ed) Ghafoor 3889 Subfamily Mimosoideae Tribe Ingeae trt LN (Roxb.) 13 13 K.U. Campus: Siddiqui 17 Benth. (cultivated) Subfamily Papilionoideae Tribe Sophoreae **Sophora mollis (Royle) Baker 9 9 Panjgur: Omer 2057 subsp. griffithii (Stocks) Ali Fig. 1) Tribe Tephrosieae * Millettia peguensis Ali (culti- 11 11 K.U. Campus: Siddiqui 35 vated * Tephrosia apollinea (Delile) 22 H Khushab: Ghafoor 3762 k “шона. purpurea (L.) Pers. 11 11 Kashmir: T. Ali 15; Layyah: Ghafoor 3735 Tephrosia purpurea 22 Makran: Omer 2106 Tephrosia pow (Dalzell) 11 11 K.U. Campus: Jahan 32 Sant. & М Tephrosia саз Вакег E1 .U. Campus: Siddiqui 10, 54, 56, 47; ir "m Karachi: Siddi. ui 68, Jaha Tribe Robinieae *Gliricidia sepium (Jacq.) Steud. 11 11 K.U. Campus: Siddiqui 23 n ltivated irri bispinosa (Jacq.) W. 6 6 Kathore: Jahan 71 ight *Sesbania sesban (L.) Merrill 6, 2n ^ 12 Darsanochano: Siddiqui 93; Safari Park, (cultivated) Karachi: Siddiqui 59; Sajawal: Ahsan Tribe Indigofereae Cyamopsis tetragonoloba (L.) 7 7 Penn Ghafoor 3577; шалар T. bert ( le 1 (r: 3 & Ghafoor 3606; K.U. Campus 4) 61 Sipe aai caerulea Roxb. var. 8 8 K.U. Campus: Siddiqui 37 "аула cordifolia Heyne ex 8 K.U. Campus: Ahsan 4; D. G. Khan: Ghafoor 3730 8 Bawana: Ghafoor 3689; Darsanochano: re a hochstetteri Baker (Fig. 6) Jahan 31; Kathore: Jahan 70; K.U. Razaq 140 Campus: Annals of the Missouri Botanical Garden TABLE 1. Continued. Chromosome Basic number number Taxon x Voucher Indigofera oblongifolia Forssk. 8 K.U. Campus: nee 16, 52; Safari Park, Karachi: Sid 66; Darsano- chano: Siddiqui жа Тыш Siddiqui 107 Tribe Desmodie Alysicarpus ECTS (Ba- 8 8 K.U. Campus: Moin. 45 ker) Jafri & Ali (Fig. 7) Alysicarpus monilifer (L.) DC. 8 K.U. Campus: Siddiqui 27, Ahsan 2 * Alysicarpus ovalifolius (Schu- 8 K.U. Campus: Moin. 60 mach.) J. Léonard * Alysicarpus rugosus (Willd.) 8 Ghotki: Ghofoor 3515 «Геза: elegans DC. 11 11 Kashmir: 7. Ali 236 *Desmodium gangeticum (L.) DE Kashmir: Т. Ali 130 «родий juncea (L.f.) Pers. 10 10 Kashmir: Т. Ali 471 var. sericea (Thunb.) Lace & Hemsl. Tribe Phaseoleae Marice cajan (L.) Millsp. (cul- 11 11 K.U. Campus: Moin. 11 vat Chorin ternatea L. 8 8 Malir, Karachi: Moin. gia fruticulosa Wall. ex 11 11 Murree: Ghafoor s "em *Glycine max (L.) Merrill (culti- 20 20 K.U. Campus: Razaq 137 vated *Phaseolus vulgaris L. (cultivat- 11 11 D. G. Khan: Ghafoor 3690 ed *Rhynchosia capitata (Heyne 11 11 K.U. Campus: Razaq 131 ex Roth) DC. Rhynchosia minima (L.) DC 11 K.U. Campus: Siddiqui 18 Rhynchosia Баните 11 K.U. Campus: Siddiqui 38 Stocks *Vigna mungo (L.) Hepper (cul- 2n = 22 1% K.U. Campus: Moin. 29 tiv. *Vigna trilobata (L.) Verdc. 11 K.U. Campus: din 129; Manghopir: dide | Razaq 155 Vigna unguiculata (L.) Walp. 11 K.U. Campus: Razaq 136 subsp. 8 H | ( 1+. 1) Tribe Aeschynomeneae *Arachis hypogaea L. (cultivat- 20 10 K.U. Campus: Razaq 130 ed) Tribe een hagi maurorum Medikus 8 8 K.U. Campus: Moin. 14; Sajawal: Ahs- an n ds Thatta: Siddiqui 96, 128, nens bicuspis Fisch. 8 8 ilgit: 2470 (Fus. S & E Puce Omer - т leucocephalus Gra- 8 Shangla, N.W.F.P.; Ghafoor 4008 t ** Astragalus ptilocephalis Baker 8 Chitral: Ghafoor 2567 Volume 81, Number 4 1994 Jahan et al. 795 A Numbers in Fabaceae TaBLE 1. Continued. Chromosome Basic number number Taxon x Voucher **Chesneya parviflora Jaub. & 8 8 Panjgur: Omer 2076 Spach (Fig ia j А (Wahl) 8 8 Chitral: Ghafoor 3154 Tribe de E cus (Roth) 8 8 K.U. Campus: Ahsan 54 Arn. (Fig. 10 pee glabra Boiss. (Fig. 8 8 Panjgur: Omer 2073 11) Tribe Vicieae *La athyrus а phaca L. (Fig. 12) 7 7 K.U. Campus: Razaq 149 *Vicia faba E ор, 6 6 Chitral: Chior 3200 Tribe Cicereae *Cicer arietinum L. (cultivated) 8 8 K.U. Campus: Moin. 28 Tribe Trifolieae *Medicago falcata L. 16 8 Astor: Omer 2 *Medicago lupulina L. 8 U. Campus: Siddiqui 30; Dir: Gha foor 2426; Swat: Ghafoor 4007 Medicago sativa L 8 Thatta: Siddiqui 104 Medicago sativa 16 Chitral: Ghafoor 2848, 3108; K.U. Campus: Jahan 16; Mustuj: Ghafoor 3003; Arkari: Ghafoor 2799 Melilotus alba Desr. 8 8 K.U. pus: Siddiqui 28; Thatta: Sid diqui 127, 1 13 Melilotus indica (L.) All. 8 K.U. р ui 6; Thatta: Sid- diqui 125; as diu: Siddiqui Melilotus officinalis (L.) Pall. 8 Astor: Omer 2428; m Omer 2582; Avarik: Ghafoor A Uo resupinatum L. (Fig. 8 8 peg Ghafoor E Thatta: Siddi- onella rion -graecum L. 2n = 16 8 K. б. bee Moin. 24 а (Fig. 14) Tribe Crotala talari Crotalaria burhia Buch.-Ham. 8 8 K.U. Campus: Siddiqui 47, 51; Cholis- ex Benth tan: Ghafoor r 3543; Darsanochano: Siddiqui *Crotalaria juncea L. (Fig. 15) 8 Khushab: Cher 3813 *Crotalaria medicaginea Lam. 16 Kashmir: 390 var. medicaginea (Fig. 16) Tribe Genisteae nl mes stenophyllum 13 13 Khushab: Ghafoor 3829 Boiss. (Fig. 17) is * Count new to flora of Pakistan. Count being Ыы Hs the first time. sia subtriflora and T. strigosa have n = 11, while 7. sie! ate has both n = 11 and ži = 22, rvations of LN (1986). In T. Aena we find n — 22 ac 2), jo cogendi reported as a Aelius cytotype in n Rol = 11 PU z Raina 86); therefore the present report is first frst of in this species. bineae, Gliricidia Am has n = 1l. 796 Annals of the Missouri Botanical Garden Р me voiced ж уфа взе. E eme apollinea (Ghafoor 3 gina (Ghafoor туң met a ase І, n = —5. Indigofera са pu Bi Т Both n — 11 and 10 have been reported for many cultivated species (cii tt, 19813). ndigofe ur species of Indigofera ex- ies in Phaseoleae ii n І, pt Glycine max with and C TNCS Nro with n — 8 (Table D. ‘Cutie of yamopsis eoo, (Ghafoor 3606), m a var. caerulea (Sidaign? 37), diakines wa (Omer a saison e r m e tam. 52) à n usq d" 6. Indigofera ATE Me х= 11 are the most со base number ге + in the "ie for this ba ute = ai 198 н in e x= is almost cert С. ef of our accessions of Alhagi maurorum whee n= acco records eu 1979; Кыс & 990 ( F observations of n — 8 in Astragalus bicuspts (Fig 8a & b a d n = 8 in eme ри ы 9) are first EXT number determi these specie Volume 81, Number 4 1994 Jahan et al. Chromosome Numbers in Fabaceae Ficures ae . Pollen mother We I, n = 8.—Ba. rages E one taphase I, n = 8.—9. Che cuneifolia (Ahsan 54), late anaphase I, Chrom numbers of two poas < of Tav- erniera тейи have п = 8, co nt with the basic number for the ii (Goldblatt, 1981), The present count for ra (п = 8; Fig. 11) is new to sci Н в In the monogeneric Cicereae our count of л = or um confirms basic number (x — 8) for the species and А (Goldblatt, 1981а). rifolieae are important in agriculture and al- ready well studied u barey Medicago falcata £F. 70), "m atta votes . Taverniera glabra (Omer 2073), ы cn de T: Ая heterophyllus (Moin. 45 i = 8.—8| ге averniera = 8. b. Се —— ams 2076), pi anaphase d M. sativa have polyploid races. Medica, po has diploid : and tetraploid eyto pet, but кар) seed has been extensively шей by Mein rear in the literature. We iem two cy- totypes, i.e., 4x, in our accessions. We did not find any sedes in the rest of bé d genera of In Crotalariese our counts for three species of n mother meta phase I, n ria medica Crotalaria (Table 1) ror the earlier pore for the species (Bir & , 1978; Khatoo Ali, 1991). In C. me oe (Fig. 16) folk dip- loids and Сасы are recorded (Bir & Sidhu, 1975; Bir & Kumari, 1978). Сан nisteae the count for Argyrolobium sten- ophyllu = 13 (Fig. 17),:is the first for the species. Goldblatt (1981a) suggested x = 8 as the bas genus and regards the gametic numbers 15 and 13 as hypotetraploid. cell meiotic and root tip mitotic chromosom aphase talar aginea var. асин (Т. Ali 390) prophase П, п 17. Argyrolobium E Ама (Ghafoor 3829), diakinesis, n = 13. Annals paiet id Garden 1 3. Trifolium eri edges n ieri metaphase, 2n = 16.— 15. Crotalaria juncea (6 T LITERATURE CITED ALI, 5.1 imosacese. In: E. Nasir & S- I- A! 6: 1 " Genét. Ibér. 22: 41- Volume 81, Number 4 1994 Jahan NE MEA Numbers in Fabaceae & A. Husain. 1967. Chromosome studies in some flowering гов of West Pakistan. І. Phyton (Buenos Aires) 24: 49-55. 68. Chromosome Number . 1965. c chro- some vascular plano of de Indus Dif т Bor [5 "l8 : 289-29 А. Be AKHTAR. ieee. Meiotic chro moso deat some vascular plants of the indie из delta. "Т. ТЕ Not. 119: 24-32. died Chromosome Number worm, A n J. m Ms 71-72 FEDORO . (edito: ES he Number: of of Flowering Plants. One Kod сазы, Yon. West Germany (reprinte GOLDBLATT, P. 1981a. Cytology а the eT of Le; guminosae. Pp. 4 . M. Polhill & P. H. чети (editors), тиа іп үөр Systematics, Part 2. Royal Botanic Gardens, Kew. —— (editor). 1981Ь. iof to Plant Chromosome gr. Syst. Bot. Missouri Numbers ae 1978. Mon Bot. Gard. 5: 1-553. Index to Plant Chromosome N Р gui umbers 1979- 1981. Monogr. Syst. Bot. Missouri Bot. Gard. 8: .1- kr 1985. Index to Plant Chromosome Numbers б 1982- 1983. Monogr. Syst. Bot. Missouri Bot. Gard. 13: 1-224 198 88. Index to Plant Chromosome Numbers О 1984- 1985. Monogr. Syst. Bot. Missouri Bot. Gard. 23: 1-264. Ja 966. The Flora of Karachi. Book KHATOON, . I. AL. 198 2. Chromosome numbers jm ome plants of Pakistan. Pakistan J. Bot. 14: 117- 129. ————. 1991. Chromosome numbers mily Papilionoid leae (Leguminosae) Mu Paki- stan. Willdeno owia 20: 159-165. oe Е J. (editor). 1973. Index to Plant Chromo- Numbers for 1967-1971. Regnum Veg. 90: ——— & subfa: 1-5 ho URAISH, H. B А. FARUQI. E bears та іп Rhyncosia minima апа К. ота (Legu nosae). Pakistan J. Bot 2:057 82. Srivastav, Р. K. & S. N. К 1986. етен AINA. of Tephrosia УІ: ne systems in some taxa. Cy- tologia 51: 359-3 CHROMOSOME NUMBERS IN Zeenat A. Razag.* Ahsan A. Vahidys COMPOSITAE FROM and S. 1. Ali PAKISTAN! ABSTRACT Chromosome numbers of 82 taxa, belonging to 48 genera in ten tribes of the family Соки are reported from Pakistan. The chromosome numbers of 13 taxa the first time, including one new generic count ; 6 for L salsolioides, Conyza stricta var. pinnatifida, Grantia aucheri, apio pee E altaicus var. canescens, Launaea —€— and Phagnalon pycnophyllon; n = 10 for Blumea bovei and Sonchus lacnocephalus; and n = d for Scorzonera корана and Scorzonera tortuosissima. The ето counts for 31 other taxa аге п for the flora of Pakist Compo aH pas approximately 20,000 once preis were taken from Mee rary species and a 40% of these have been inves- unts, in all cases. Later on slides were made bon “rol Ne 1977). Very little acit in euparal or Canada balsam. tologic research has been carried out on the qc where to date chromosome OBSERVATIONS AND RESULTS niher for o only es (i. e., ca. 11.0%) of the ca. 30 78) fos the family hav А total of 140 chromosomal counts on ; been counted (Ba aqua r & е. 1970; а representing 62 Jata BLE а. а " & Mi, 1988; Басы al, ; BO RI Compositae have been dete piu E ts for . ere contribution ел ш үзган numbers for taxa are reported for the first tim found to be reported in IPCN (Fondi 1974; dem 1981, 1984, 1985, 1988; Moore, 1973, 1974; Ornduff, 1967). The classification adopted MATERIALS AND METHODS here follows that of Heywood et al. (1977). Meiotic material qari of immature c capit- 82 taxa in 48 gen ula, collected mostly in the wild (those which were DISCUSSION cultivated are specified d Table 1), was fixed in пер Vernonieae аге the least known acetic alcohol (1:3) for 24 hr. and stored at —4°C. tribe aie with fewer than 100 (out J slides were prepared by conventional squash 1450) s pue so far counted (Mathew & Mathew, technique using aceto- or propionic-carmine. Counts 1983). Available ctl ogical чун on the genus were made from pollen mother cells, except in Vernonia Жыш. 1974; Jones, 1974; Mathew v xU tenuiloba, where both somatic ii ga- & Mathew, 1976; СШ, 1978; Kei & rone metic counts were made. 1979; Gupta & Gill, Hm) reveal that species 9 sis, root ti ting seeds this genus are based on x = 9, 10, and 17. ме were даць with 8-hyd Yigg for 4hr., of the four species of onia, the only genus 0 5 h ern acetic alcohol (1:3) for 1 hr., hydrolysed ^ Vernonieae in Pakistan (Stewart, 1972), E IN HCI for 6-12 min. and squashed in 1.8% amined. According to = (1977), Old Wor n кёз, onia have x = 9 or 10 and a little poly, yploidy- m ! We k M. Qaiser, S. Khatoon, A. Ghafoor, = gs sir of the песна " Rance for — of Bde: of vouchers. We also thank A. Gh. T T. Ali, N. Ahsan, M. M uddin, and В National the University of Karachi for collecting Prosa ud de paper is a part of the vh . supported by ience Rooters: fo INT 851031 я Тир е " tany, University i Karachi, Karachi-75270, Pakistan. Department tet Le etics, University of Karachi, Karachi-75270, Pakistan. ANN. Missouni Bor. Garp. 81: 800-808. 1994. Volume 81, Number 4 1994 azaq et al. Chromosome Numbers in Compositae TABLE 1. Ch b Comp . K.U. = Karachi University; D. G. Khan = town of Dera Ghazi Khan. Chromosome Basic number number Taxon n x Voucher Tribe Eupatorieae * Ageratum conyzoides L. (Fig. 1) 10 10 Sialkot: Ghafoor 4311 * Ageratum houstonianum Mill. 20 K.U. Campus: Siddiqui 43 (cultivated) Tribe Verno Vernonia cinerascens у ad 20 10 K.U. Campus: Moin. 3 бекіт cinerea (L.) Les 9 9 Kashmir: T. Ali 112; p Ghafoor 4278; Sargodha: T. Ali 1 Tribe Astereae Сопуга aegyptiaca А 9 9 D. G. Khan: Ghafoor 3652 Conyza bonariensis ^N ) Cronquist 27 K.U. Campus: Ahsan 70; Kathore: Jahan 75; Hasan Abdal: Ghafoor 2270 *Conyza japonica (Thunb.) Less 9 Sargodha: Т. Ali jd **Conyza stricta Willd. var. pinna- 9 Kass T. Ali 2 aie (D. р it. ropappus on (Willd.) 9 9 Dir: Ghafoor 2324; Gilgit: Omer 2412, 2304, чаңы Fi ** Hetero, рор. dem illd 9 Chitral: Ghafoor 3224 Novopokr. . canescens (Nees) Serg. (Fig. с. holohermaphrodi- 9 Quetta: T. Ali 1414; D. С. Khan: Ghafoor 3643 tus Grierson Myriactis wallichii Less. 18 18 Rawalpindi: Ghafoor 4119 Tribe Inuleae **Blumea bovei (DC.) Vatke (Fig. 4) 10 10 Mr T. Ali 9. Blumea lacera DC. 10 .U. Campus: Si di ui 2 Blumea obli (L.) Druce 10 KU. со Siddiqui 5 Gnaphalium luteo-album L. 7 7 Dir: Ghafoor oe p тата 3351 **Grantia aucheri Boiss. (Fig. 5) 9 9 Makran: Omer 2 *Inula cuspidata (DC.) C. B. 10 10 Hazara: Omer 27 ey Clarke Iphiona g ioides (Boiss.) An- 9 9 cerned Omer pie E Ali 905; Safari Park, derb. (Fig. 6) Karachi: Siddiq Phagnalon niveum Edgew. 9 9 Шай: yrs 2262; з С. Khan: Ghafoor 3700 dfe pycnophyllon Rech.f. 9 Chitral: Ghafoor 3157 t a arguta. Boiss. X» 8) 10 10 Makran: T. Ali 8 eis indica (L.) Le 0 Makran: Т. Ali 4) K.U. Campus: Jahan 7, 8, 9 Pluchea reos d a ) Clarke 10 K.U. Campus: Jahan Pulicaria angustifolia DC. T 7 K.U. Campus: Sd Fa 49 ** Pulicaria gnaphalodes (Vent.) 7 Quetta: Т. Ali Boiss. (Fig Pulicaria “ole Jafri ? Super Highway, Karachi: T. Ali 1437 Tribe Helianthea *Bidens EDGE (Lour. Merr. & 36 12 Hazara: Omer 2233 er ies acmella (L. Кин 17 LT K.U. puni e 41 Согео, atkinsoniana Doug 12 12 Malir: Moin Жн ылы 10 10 K.U. Campus: Jahan 14, 25 Coreopsis lanceolata L. (cultivat- ed) (Fig. 10) 802 Annals of the Vide Botanical Garden TABLE l. Continued. Chromosome Basic number numbe Taxon n Voucher *Cosmos bipinnatus Cav. (cultivat- 12 12 K.U. Campus: Siddiqui 44 e * Dahlia variabilis (Willd.) Desf. 32 16 K.U. Campus: Jahan 15 (cultivate Eclipta prostrata (L.) L. ll ll K.U. Campus: Siddiqui 9; Sajawal: Ahsan 1 Thatta: TRER 106, А Khushab: Pet 3835; r pater Flaveria trinervia (Spreng.) C. 18 18 Kathore: pus mi ore pulchella Fouger (culti- 17 17 K.U. Campus: Siddiqui 24 козы oga parviflora 8 8 Kashmir: as Ali 312; D. G. Khan: Ghafoor 3650 *Helianthus annus L. на, 17 17 «Ба харти *Parthenium hysterophorus 18 18 Сш: Ghafoor *Rudbeckia maxima Nutt. (culti- 18 18 K.U. Campus: безе 10 vated) Tridax procumbens L. 18 9 K.U. Campus: Siddiqui * Xanthium strumarium L. 18 18 K.U. Campus: Razaq s Ghotki: Ghafoor 3512; Kashmir: T. Ali 198; Kathore: Jahan 74 Zinnia elegans Jacq. (cultivated) 12 12 K.U. Campus: Moin. 5 Tribe Tageteae eee tenuiloba (DC.) B. L. 12 12 K.U. Campus: Ahsan 55 Robinson (cultivat Dyssodia tenuiloba (Fig. in 2n = 24 K.U. Campus: Jahan *Tagetes minuta L. (Fig. 12) 24 12 Jhelum: Ghafoor mes а Т. Ali 349 Tribe MEO E Hertia int ia (Boiss.) Kuntze 10 10 Quetta: T. Ali i и analogus DC. 20 10 Swat: Ghafoor 345 ecio desfontanei Druce 10 — Omer 26 Chitral: Ghafoor 2551; Dh; Ghani Senecio krascheninnikovii 10 Chitral: Ghafoor vf Gilgit: Omer 2520 Schischk. Tribe Anthemidea: Achillea miletin L 9 9 Gilgit: Omer 2326; Kashmir: T. Ali 55 59 * Anthemis co 9 9 K.U. Cam mj Jahan 36; Kashmir: T. Ali 511; Chitral: Ghafoor эге 2976, 2518 * Artemisia capillaris Thunb. (Fig. 8 8 Sargodha: Ghafoor 38 *Artemesia persica Boiss. (Fig. 14) 9 9 Chitral: Ghafoor 3225, 2571, 2581 Artemisa rutaefolia Spreng. (Fig. 9 Chitral: Ghafoor 3219 15 ** Artemisia salsoloides Willd. (Fig. 9 Gilgit: Omer 2430 d emisia siversian 9 Chitral: Ghafoor 3111 “andl wichophylle( Баец) 9 9 Chitral: Ghafoor 3234 eem fruticulosum Ledeb. 9 9 Gilgit: Omer 2548 SM ird P е disciforme (C. 9 9 Chitral: Ghafoor 2846 Mey -) Sch. Bip. eer re E Volume 81, Number 4 Razaq et al. 803 1994 Chromosome Numbers in Compositae TABLE l. Continued. Chromosome Basic number number Taxon n x Voucher Tribe Cynareae Centaurea cyanus L. (cultivated) 12 12 K.U. Campus: Jahan *Oligochaeta ramosa (Roxb.) Wag- 14 14 Darsanochano: зайцы 82; Макгап: Отег enitz 2090, Т. Ali 9 Tribe Lactuceae **Cephalorrhynchus picridiformis (Boiss.) Tuisl (Fig. 18) со © Quetta: Т. Ali 1394 ium intybus L. 9 9 Thatta: Siddiqui 105 *Crepis multicaulis Ledeb. subsp. 5 5 Swat: Ghafoor 3376 а (Regel) Babe. герїз sancta (L.) Babe 5 Chitral: Ghafoor ped actuca dissecta D. Don 8 8 pe 2 *Lactuca serriola L. 9 9 Attoc € зв Zhob: Т. Ali 1059 *Launaea capitata (Spreng.) Dan- 9 9 SEER T. Ali 1 dy Launaea nudicaulis (L.) Hook.f. 9 Rawalpindi: Ghafoor 4175; Faisalabad; Ghafoor 4377; Makran: Т. es 708; Kathore: Jahan 63; Sajawal: Ahsan 15; Thatta: Siddiqui 98, 114, 118, 12 **Launaea oli la (Haussl 9 Makran: T. Ali 953 "ede ex роп: ) Во (Fig. *Launaea procumbens (Roxb.) Ra- 9 sre Omer 2097, 2040; Hasilpur: Ghafoor mayya & Rajagopal (Fig. 20) Launaea remotiflora (DC.) Amin 9 baa vm Karachi: Siddiqui 71; K.U. Campus: ex Rech.f. Moin. 6; Kashmir: T. Ali 271; Bahlolpur: *Launaea resedifolia (L.) Kuntze 8 8 Safari Park, Karachi: Ahsan 39; Bahawalpur: Ghafoor 3574 *Launaea secunda (C. B. Clarke) 9 9 Chitral: Ghafoor 2410 Hook.f. pow tomentella Rech.f. (Fig. 6 6 Zhob: T. Ali 1167 21) Picris hieracioides L. 5 5 Chitral: Ghafoor 2583 *Scrzonera koelpinioides Rech.f. 14 7 Makran: 7. Ali 992 (Fig. 22) е tortuosissima Boiss. 14 Makran: T. Ali 991 (Fig. oss asper (L.) Hill 9 9 Kashmir: T. Ali 578 **Son we lachnocephalus Rech.f. 10 10 Kashmir: T. Ali 174, 493 (Fig. 2. Sonchus oleraceus L. 16 8 K.U. Campus: Jahan 29; D. G. Khan: Ghafoor Sonchus wightianus DC. 9 9 Kashmir: T. Ali 331; Rawalpindi: T. Ali 1955 H mes new to flora of Pakistan. Count new to science Our counts for V. cinerascens (n = 20) and V. Anderson et al., 1974). Out of eight taxa of As- cinerea (n = 9) confirm the above statement. tereae studied, chromosome counts of all the spe The жузе пре ат pre various members of Às- cies except Myriactis wallichii are based on x = tereae is reported t = 9 with polyploidy 9. All the species ка the genus Myriactis are re- (Raven et ks 1960; вад et al., 1964, 1969; рогіей to have n = 18. 804 Annals of the UA Botanical Garden 9 „б Eire I "e Chromosomes in members о f Compositae.— 1. Ager joran i e yn (Ghafoor 4311). дай, Hispana кини (Ghafoor 2324), metaphase-I, n = 9.—3. Heteropappus alta rigs — 6 (Chor. tem metaphase he Blumea boves (T. Ali 9. vhs diakinesis, m m . Gra ntia ice (Omer 2118), diakinesis, n = 9, — ap s grantioides (Omer 2052), diakinesis, n — 9. The members of tribe Inuleae je a spot conclusion (1977). In rons the most fre- inance of the basic numbers x = 9 a 0 (Merx- quent basic numbers are multiples of ten — muller et al., 1977). In this tribe, we зан mined stam, 1977). Our observations on an chromosom 15 species in eight igs Of these, four species numbers of three species of Senecio (Table 1) are (belonging to d on x = 7, four based on x = 10. The count for S. k —— species (belonging to three genera) on x — 9, and e: n = 10, differs from the previous report seven species (belonging to three genera) on x — — 9 (Khatoon & Ali, 1988). 10. Our data thus agree with Merxmuller et al.'s Chromosome counts have been reported for is Volume 81, Number 4 1994 Razaq et al. 805 q Chromosome Numbers in Compositae -12. Chrom m aoe 8. Pluchen arguta (Т. li 87 7) F = 7.—10. Cor ag 1 иен (/аһап ), one. Ta a Bo 2n = 24. 50% of the genera of arabe dee peni & Ten tie 1977). Ten of the of An- "AR i = 9; рыбы бар, ris, ieee n= g^ (Fig. 13), a co 1 nn to he e findings (2n = 18: daas : Peng & Hsu, 1978) e Present ets may represent aneuploid variation. of Composita: metap reg че tae.— 7. Phagnalon оку (Ghafoor 3157), ana ЖЕЗ, Pulicaria gnaphalodes (Т. Ali 1243), a н tenuiloba (Ahs . Tagetes e i ra УГ 3886), “diakinesis, n= The most frequent ечен misia is п = 9 and probably th т for the genus was also x = 9 eh & Wojtas, 1987 Chro: me am for 21 taxa of Lactucea are reported her e count for Cephalorr: Eran chus micridiformis (n = 8, Fig. 18) does not agree Annals = iuh Mon Garden 17 FIGURES 13-18. Chrom members of Compositae.— 13. Artemisia capillaris (Ghafoor 3801), TD n = 8.— 19, Artemisia persica (с 3225), diakinesis, ne den Зн 5. CLE rutaefolia (Ghafoor diakinesis, n = 9.—16 2430), diaki es nacetum vii. sum ( 2548), башпы, п = 9.— 18. Cephalorrhynchus feti mera (Т. Ali 7900. denar with basic number x — 9, proposed by ee the genus. Chromosomally, the Lactuceae are ae & Wylie (1955), tc this genus. In Launaea, x — haps the best known tribe in hac fa n 6, 7, 8, and 9 are reported. However, n = 6 has mosome numbers are known 87. 0% of i en reported for L. asplenifolia Hook. f. by Sar- genera of the tribe (Tomb, on Stebbins ae kar et al. (1975). We have observed the same ны) Ыз x = 9 as the ancestral base € re number for L. tomentella (Table 1 Fig. 21). For mo number for the tribe, on es basis 0 the genus Sonchus, x = 8 and 9 are r reported. Our {ге а псу of occurrence ot presence in Lear count of n = 10 for Sonchus lac носер (Fig. considered to be т E the p P esent а 24) establishes а new basic number (x = 10) for ten taxa out of 2 Volume 81, Number 4 1994 azaq et al. Chromosome Numbers in Compositae i ms 19-24. Chromosomes in pee Papae d» eu х = 9, whereas other taxa are based on x = 5, 6, 7, and 8. LITERATURE CITED Ап, S. I. 1978. The flora of Pakistan: Some general and analytical Doubs Notes Roy. Bot. Gard. Ed- A inburgh 36: 427-439 NDERSON, L. C., D. W. Күноз, T. Mosguin, А. M. de Compositae. — 19. L L 97), Lern n (T. Ali 953), diakinesis, aunaea S toan la (T. Ali 1167), —23.5 PowE Р.Н. Raven. 1974. Chromosome num- bers in Compositae. IX. Haplopappus and other Astereae 1: 665-671. ARANO, H aryotypes and the spec „> BAQUAR, S. R ASKA 19 Chromosome numbers in some viget pan of West Pakistan — n Genét. Ibér. 22: Annals of the Missouri Botanical Garden DARLINGTON, C. D. & A. P. WYLIE. 1955. Chromosome Atlas = Flowering Plants. George Allen & Unwin Ltd., оп. FEDOROV, 3 ^ (editor). 1974. Chromosom of Flow рыз Meer Otto Koeltz Science Publishers, d poca dde Number Reports LX. 9-230. GOLDBLATT, P. (editor). 1981. Index to Plant Chro- mosome Nu ar 1975-1978. Monogr. Syst. Bot. rae Bot. Gard. 5: ——. 1984. to Fes Chromosome Numbers i 1961. Monogr. Syst. Bot. Missouri Bot. Gard. -427. —— ———. 1985. Index to Plant Chromosome Numbers 1982-1983. Monogr. Syst. Bot. Missouri Bot. Gard. 3: 1-224. 1988. Index to Plant Chromosome Numbe NU 1985. Monogr. Syst. Bot. Missouri Bot. Card. Rn Gupta, В. C. & B. S. GILL. 1979. eee Number Reports LXIV. Tassa 28: 401-40 V.H. & UMPHRIES. а ~ Systematic r review. In: V. ood, PENES & B.L ner (editors), The Боов ad кары of e Vol. II. Academic Press, ondon & New HEY i 977. Anthem- H Же B Halo & B. L. TURNER (editors). positae, w Y nieae Me npe chromo some e numbers Bull. Ton Bot. Club -84. ———. nie nur ору а апд eye ы ац mic гате п & New York. J E 213; . & 5.1. Аш. 1988. Ch I in Compositae from Pakistan. Candollea 43: 455- 462 MATE, A. .& P. M. MATHEW. 1976. oe Du ен ©7 X 41: 401- 406. MATHEW, Р н. . МАТ 983. Studies on the South Indian Compositae Y V. pie xonomic consid- tribes Vernonieae xd Eupatorieae. aia ай p een bath In «КОҢ od, J. В. arborne P BELT тану eich th Biology and Chemisty of сте Vol. I. Academic Press, lon & New York. ‚ В. J. (editor). 1973. Index to Plan some Numbers for 1967-1971. Regnum Vere 90: 1-53 1974. Index to Plant Chromosome Numbers r 1972. Regnum ont 911-108: ‘eeu B. 1977. Senecioneae and Li Systematic н 1п: В? e & В. L. Turner (editori) The Biology id Chemist try баране Vol. П. Academic Press. Lond w York. RNDUFF, R, CUTE 1967. Index to Plant Chromo- Numbers for 1965. Regnum Vegetabile 50: ru 1-12 8. PENG, C..I. & C.-C. Hsu. 1978. Chromosome numbers in Taiwan Compositae. Bot. Bull. Acad. Sin. 19: 53- 66. Raven, P. H., T зоо D. br Күноѕ & R. SNo 1960 omosome dece I. ke tereae. Amer. J. Bot. 47: ee AZAQ, Аз; Severs & S. І. Ац. 1988. contribution - the chromosome numbers of Com- үе ositae from Pakistan. Pars | Bot. 20: 177- 189. SARKAR, A. K., CHATTERJEE. N. Datra & U. 1975. (ибо Мыш Reports XLVIII. ee 24: 367- Soxsric, О. T. 7. Chromosomal m and evo- lution in the family сор In: V. Н. Heywood, J.B 3. Ha rbor rne & В. L. Turner оли Тһе Biology cademic Press, London & New ‚ L. C. ANDE ү. W. Күноѕ & P. Н. RAVEN 1 Richtee numbers in Compositae VII: Айе Ш. Amer. J. Bot. 56: 348-353. . RUDENBERC. 1964.. Chromosome numbers wee As- tereae II. Amer. d d ddr 513-519. ۷ E. . WOJTAS. d NE АД I) 1987. Chro- moore es J some North ache species of Artemisia (Asteraceae). Canad. J. Bot. 6: 612- Srensins, б. L., Jr. ‚ J. A. Jenkins & M. S. War 953. Chromosomes ап ny Эн ири in the Co Suit Tribe Cichorieae, Univ. Calif. Publ. Bot. 16: -43 . R. 1972. An annotated yum of the scular plants of West Pakistan an hmir. In: . Nasi & S. I. Ali (editors), Flora of puro Kar: e А: "à “1977. Taemin Gr review. In: .H. ood, J. B. Har L. Turner m The Biology and tie of Compositae, Vol. II. Academic Press, London & New Yor BOOK REVIEWS Р. М. Hyland & Т. Whiffin. 1993. Australian by D. C. Christophel and B to be ordered from CSIRO а бргу East Melbourne, Victoria 3002, : 613-419-0459). ISBN 0-643- U.S. $195 outside, plus shipping and handlin pou ithin Australia, U.S. $25 Minis Aus- lia). ve Müstrülian Thapisak Rain Forest Treni an components. The main part is the computer- -based which is su MS-DO c o 4 [e] ч m. Ф explanation 2 a list of gonen with Таншу name used in th a species list e fa amily, a species n by genius; a list шпон names and a glo ossary. Volutne 2 provides f iud; and genus, with a shaw description ini p of additional features and distribution. Volum a leaf atlas, which presents excellent Hog of the leaf venation of all included L^ main att f this identification system e is the comput r ke еу. used, аши) haracters. For identification one simply indicates which of the listed character states are — on 1 the specimen 86: be ide ent P M: The - ae ^" ration are lacking and in this way the ак of possible species becomes smaller any time One can check the list of remaining species and rmine which characters have been scored а: resent. This type of key has several advantages wer the traditional keys found in ns. On » аз a starting point. Th Каа, Eo the presence in the key of easy to observe vegetative features (such as leaf position, oes of domatia) make the key very user- endly. The brief descriptions pna оуан fea- tures listed in V onfirm па шоу E r in ere fol- ted in ossible identificati ons. Iti is b poil to n de ме of species by including only species from a wane area a (the ares covered by the flora is divided bse Another option is 5 to list all the features of a a species, a sort of identification in re- ve This entire Cahn package i is Mreoped көө, not polely botanists. Г have used the Key on ince features. I found the key a pleasure to e. Frequently the woh leaves one with a handful “ рене identific ati x as and the additio: nal notes listed in | Voume. very helpful. Also, when lef names, it is relatively ка D compare the speci- mens with | 1. The entire process of identifying is quick and ей, Homeron, even with this Ар enc on b her the specim . As pointed out in j die instructions, not "n каше аге о end me For instance, leaf margins entire oothed is more reliable темин “than Tength/ vid ratio 4 choose с anth whorls? Bec possible, the authors scored Lauraceae as а bar indy à difficult choices easy to ma nake. Still, because of the enormous amount of i е resin oblems, d, they relatively eye to correct, and updated versions of n be sent to the key с users With a Lin of U.S. $200, this кин» syste m will be too So for mo ost e of staff members. Time ANN. Missouni Bor. Garp. 81: 809-811. 1994. 810 ri Bot. rici of the Mis: | Garden equate; having the key available only on a PC in the library is on not practical. Very few insti- tutions o the degree that one co is accessible to all co анх users, although that seems g oo solution ору у йй аге attractively © << scriptions and diagnoses in Volum obably be the most aspi used of the three, and арыу the author names included here would have been very practical. I ras recommend this publication to anyone intérested i in Au stra Маж rain forest trees. Its key M tot. A new пет опе new инин two OMM nece sum- ized i in a short appendix. The organization of separate summary of climate, geography (in- dole a simple stad pem d vegetation, history je me iino use by eer and s raphy, as well a to families and a useful table s species vts РМа ет ing an peter of mics and introduced taxa. These o involved and a catia ing features, and 1 think hi р blication off — г, tralian rain forest trees. o Kass question to the er Werff, Miss Botanical Garden, P.O. Box 299, St. Louis, Mis- souri 63166-0299, U.S.A. Flora of rg dann Vol. 50, Oceanic eerie 2: 1993. Australi ublishing Ser- , Canberra, ACT "ee 6 pp. рон d 644-14446-7 йал, — $444. The fourteenth lr o aes volume published in the Flora of Australia s s is somewhat o E Island, Cartier А and th coast, the Coral Sea Islands Territory to the east and northeast, Macquarie Island, west olu rulas for Lord Howe and Norfolk Islands, the re- maining islands. Volume 50 includes approximately 500 species of vascular plants in 113 families. It contains one or vegetational нее for at least the Е ‘nil should have been in- clude The bulk of the volume contains a combined treatment of a taxa within ves family that o on any of the islands. This includes keys, ries descriptions at =: taxonomic levels, e , Б ch цо and a citation of repre tati taxonomic Met This uni unique organization re- om in a volume that is taxonomically cohesive, the species easily located in the text, while аа allowing for convenient access to information specific toa particular r island. "i gr and the percentage of each island's à dan з рге- sent on the continent would be useful once the remaining ave have been treated in the com- panion volum Eleven ан wrote the text for volume 50. As has been the case with previous volumes in the Flora of Australia series, the information е is wonde nology r contractions throughout it text, (n marized at the end of the and which do not detract from its ease es use. ate — of see ine rise taxa are beautifully ere of graphics. There a e 42 black- pem an gea dude about 100 ) species, the work of three talented artis A beautiful color over and frontispiece b na Pele anus tectorius. There are graphs (by 11 arabe E of vogetatió n types and species that are uniformly exc cellent in clarity and herde Volume 81, Number 4 1994 Book Reviews 811 The Flor ontrib- utors to this book are to be vane on the t for elegance and detail in the presentation of flo- ristic data. I am eager to see the companion volume that will complete the island florulas, as well as future volumes on the continent's flora.—George Yatskievych, Miren Botanical Garden, Р.О, Box 299, St. Loui 63166-0299, U.S.A ANNALS OF THE MISSOURI BOTANICAL GARDEN: CHECKLIST FOR AUTHORS 1. General Instructions O Text is in English or Spanish on numbered a Masri is typed on one side ofa nonglossy B x x d 1 рарег. [m] "s "FERA l in. is left as argin a all around, except on the first page, which ty 3 in. left blank at the top. ble- or triple-spaced printed manu- жы including abstract, notes, legends, tables, spec- n lists, Literature Cited, and footnotes, are en- o Manuscript is also submitted on MS- ig 3% in. dis- е margin is not justified, sn QE are not set in italics are underlined instead of e or other special typefaces are not used. 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Co cation with the ент at all points of iis salit Ld PAGE CHARGES Page costs are $80 per page, although charges may be will be billed nonnegotiably to the author at $3 per line ed. CORRESPONDENCE pesh eger FE begs ging Edito Siete Publications ouri ess anical Garden s 0. B i Louis MO 63166-0299, Tel: (314) 577-5112 Ках: One in 9594 Electr FE E org Volume 81, Number 4, в.) 571-814 of the ANNALS OF THE MISSOURI BOTANICAL GARDEN lished on November 16, 1 VOLUME 81 1994. ABU-AsAB, Mones S. & PHILIP D. Cantino. Systematic Implications of Pollen Morphology in Subfamilies Lamioideae and Pogostemonoideae (Labiateae) 653 ALBERT, Victor A., ANDERS BACKLUND, KARE BREMER, MARK W. CHASE, AMES R. MANHART, BRENT D. MISHLER & Kevin C. Nixon. Functional Constraints and rbcL Evidence for Land Plant Phylogeny -.................... a 534 Ап, S. I. (See M. Moinuddin, Ahsan A. Vahidy & S. I. Ali) mm 184 Ап, S. I. (See S. M. Nadeem Ahsan, Ahsan A. Vahidy & S. I. Ali) ........... RTT Alı, S. I. (See Zeenat A. Razaq, Ahsan A. Vahidy & S. I. Ali) nm E7800 Ап, S. I. (See Bushreen Jahan, Ahsan A. Vahidy & S. I. Ali) -..................... Өд ARMBRUSTER, W. Scorr. Early Evolution of Dalechampia = кегени Insights from Phylogeny, Biogeography, and Comparative Ecology ..... 302 BACKLUND, ANDERS. (See Victor A. Albert, Anders Backlund, Kare си Mark W. Chase, James К. Manhart, Brent D. Mishler & Kevin С. Nixon 534 BHATNAGAR, A. K. (See В. N. Kapil & A. К. Bhatnagar) еи Я 145 BREMER, ves (See Victor А. Albert, Anders Backlund, Káre ко Mark W. Chase, James R. Manhart, Brent D. Mishler & Kevin ia 534 BRUNNER, DAVID. Statistical Summar y of io of the Activities in the Missouri Botanical Garden ова 93 568 BucHHEIM, MARK A. (See Brent D. ed Louise A. Lewis, Mark A. Buchheim, Karen S. Renzaglia, David J. Garbary, Charles F. Delwiche, Frederick W. Zechman, Thomas S. Kantz & Russell L. Chapman) 451 Cantino, PHILIP D. (See Mones S. Abu-Asab & Philip D. Cantino) ~ 653 CARTER, ine A Preliminary Classification of Euphorbia Subgenus Eu- phorbia 368 CHAPMAN, RUSSELL L. (See Brent D. Mishler, Louise A. Lewis, Mark A. Buchheim, Karen S. Renzaglia, David J. Garbary, Charles F. Delwiche, Frederick W. Zechman, Thomas S. Kantz & Russell L. Chapman) ... 451 CHASE, e W. (See Victor A. Albert, Anders Backlund, Kare Bremer, Mark W. Chase, James В. Manhart, Brent D. Mishler & Kevin С. ixon Оа 534 Снід Jur CHEN, MEGHAN С. MENDENHALL & BILLIE L. TURNER. Taxonomy of Thermopsis (Fabaceae) in North America 714 CREPET, WILLIAM L. (See Kevin C. Nixon, William L. Crepet, Dennis Ste- & Else Marie Friis) 484 Davita, Patricia. (See Victoria Sosa & Patricia Dávila)... 749 Dencan, BIJAN & Вакт ScHUTZMAN. Contributions Toward a Monograph af of Neotropical Jatropha: Phenetic and Phylogenetic Analyses Shae ae ee F. (See Brent D. Mishler, Louise A. Lewis, Mark A. Buchheim, Karen S. Renzaglia, David J. Garbary, Charles F. Delwiche, ee W. Zechman, Thomas S. Капі & Russell L. Chapman) .... DoNocHUE, MicHAEL J. Progress and Prospects in Reconstructing Plant Phylogeny DONOGHUE, MICHAEL J. (See James A. Doyle, Michael J. Donoghue & lizabeth A. Zimmer) DovLE, James A., MICHAEL J. DONOGHUE & ELIZABETH A. ZIM- ein si lo and Ribosomal RNA Data on the Origin of Angiospe FRIIS, e MARIE. (See Kevin C. Nixon, William L. Crepet, Dennis Ste- п & Else Marie Friis) Ыы C. (See R. Haicour, L. Rossignol, M. Rossignol & C. Gaisne) — GARBARY, Davip J. (See Brent D. Mishler, Louise A. Lewis, Mark A. Buch- heim, Karen S. Renzaglia, David J. Garbary, Charles F. Delwiche, Frederick W. Zechman, Thomas S. Kantz & Russell L. Chapman) — GILBERT, M. С. The Relationships of the Euphorbieae (Euphorbiaceae) ..... GILLESPIE, LYNN J. Pollen Morphology and Phylogeny of the Tribe Pluke- netieae ka ali ) Haicour, R., L. RossicNor, M. ROSSIGNOL & C. GarsNE. Patterns of Di- versification red Evolution in Phyllanthus odontadenius к сеае) HAYDEN, W. JOHN. sini Anatomy of Euphorbiaceae Subfamily Old- fieldioideae. I. Over JAHAN, BUSHREEN, AHSAN A. VaHIDY & S. I. ALI. Солине Numbers Taxa of Fabaceae Mostly Native to Pakistan .................. JENSEN, ее INA VocEL-BAUER & MAREI NITSCHKE. a Proteins he Systematics of the Euphorbiaceae Kantz, THOMAS S. (See Brent D. Mishler, Louise A. Lewis, М А. Висһ- eim, Karen S. Renzaglia, David J. Garbary, Charles F. Delwiche, Frederick W. Zechman, Thomas S. Kantz & Russell L. a артап) .— Kapit, R. N. & А. К. BHATNaGAR. The Contribution of Embryology to the Systematics of the Euphorbiaceae KILLEEN, ear: ү (See Guillermo А. Norrmann, Camilo L. Quarin & Timothy J. Killeen) . ire H: a Н. P. Linder & H. Kurzweil) ...... Lazarus, Davip B. (See Barbara A. R. Mohr & David В. Lazarus) m d A. & MICHAEL С. Simpson. Phylogenetic Implications of a E in the Oldfieldioideae (Euphorbiaceae) ....—— —— LEVIN, GEOFFREY A. & MICHAEL С. SIMPSON. Phylogenetic Relationships of Didymocistus and Hymenocardia Е д —— LEwis, LOUISE A. (See Brent D. Mishler, Louise A. Lewis, Mark A. Buch- heim, Karen S. Renzaglia, David J. Garbary, Charles F. Delwiche, Frederick W. Zechman, Thomas S. Kantz & Russell L. Chapman) — 792 160 Linper, H. P. & Н. KURZWEIL. The Phylogeny and Classification of the Diseae (Orchidoideae: Orchidaceae) MANHART, Seog R. (See Victor A. Albert, Anders Backlund, Kare Bremer, Mar . Chase, James R. Manhart, Brent D. Mishler & Kevin C. б MENDENHALL, MEGHAN С. (See Chia Jui Chen, Meghan С. Mendenhall & Billie L. Turner) MISHLER, BRENT D. (See Victor A. Albert, Anders Backlund, Kare Bremer, Mark W. Chase, James R. Manhart, Brent D. Mishler & Kevin C. Nixon) MIsHLER, BRENT D., Louise A. Lewis, Mark A. BUCHHEIM, KAREN S RENZAGLIA, DAVID J. GARBARY, dius F. DELWICHE, FREDERICK ZECHMAN, THOMAS S. Kantz & RUSSELL L. CHAPMAN. Phylogenetic iS of the “Green ye. d "“Bryophytes” ас. Monr, BARBARA А. R. & Davin B. Lazarus. Paleobiogeographic Distribution of Kuylisporites and its Possible Relationship to the Extant Fern Genus Cnemidaria (Cyatheaceae) Mornuppin, M., AHSAN A. VAHIDY & S. I. ALI. Chromosome Counts rundinoideae, Chloridoideae, and Pooideae (Poaceae) from Pakistan . MuRoz-Scuick, MÉLICIA. (See Charlotte M. Taylor & Mélicia Munoz-Schick) NADEEM AHSAN, S. M., AHSAN A. VAHIDY & S. I. ALI. Chromosome Numbers and Incidence of Polyploidy in Panicoideae (Poaceae) from Pakistan .. NrrscHKE, Maret. (See Uwe Jensen, Ina Vogel-Bauer & Marei Nitschke) NIXON, "dp C. (See Victor A. Albert, Anders Backlund, Káre Bremer, Mar . Chase, James R. Manhart, Brent D. Mishler & Kevin C. Nixon, KEVIN RS WILLIAM L. CREPET, DENNIS STEVENSON & ELSE MARIE FRIIS. A Reevaluation of Seed Plant Phylogeny ....— — NorRMANN, GUILLERMO A., Сампо L. Quarin & TIMOTHY J. Ki- LEEN romosome N rs in Bolivian Grasses (Gramineae) — NowıckE, Joan W. А Palynological Study of Crotonoideae (Euphorbiaceae) REN os L. (See Guillermo A. Norrmann, Camilo L. Quarin & Timothy J. Killeen) еен Razag, sla A., AHSAN A. — & S. I. Аш. Chromosome Numbers in Compositae from Pakist 02 eee RENZAGLIA, KAREN S. (See Brent D. Mishler, Louise A. Lewis, Mark A. Buchheim, Karen S. Renzaglia, David J. Garbary, dd» F. Delwiche, Frederick W. Zechman, Thomas S. Kantz & Russell L. Chapman) — RICHARDSON, P. Mick. Origin and Relationships of the Major Plant Groups: Dedication to Arthur Cronquist and Introduction ы RE RossicNoL, L. (See R. Haicour, L. Rossignol, M. Rossignol & C. Gaisne) . Rossicnot, M. (See К. Haicour, L. Rossignol, M. Rossignol & C. Gaisne) 687 451 RUDALL, PAULA. Laticifers in Crotonoideae (Euphorbiaceae): Homology and Evolution Ruepa, Ricarpo М. Systematics and Evolution of the Genus Petrea (Ver- benaceae) ScHUTZMAN, BART. (See Bijan Dehgan & Bart Schutzman) „mm SIEGLER, Davip S. Phytochemistry and Systematics of the Euphorbiaceae Simpson, MICHAEL G. (See Geoffrey A. Levin & Michael С. Simpson) —. Simpson, MICHAEL G. (See Geoffrey A. Levin & Michael С. Simpson) ..... з арни & А DáviLA. Una Evaluación del Conocimiento Flor- o de Méx LN Dennis. (See Kevin C. Nixon, William L. Crepet, Dennis Ste- n & Else Marie Friis) TAYLOR, CHARLOTTE M. Revision of Hillia (Rubiaceae) mu TAYLOR, CHARLOTTE M. & MELIcIA Muñoz- SCHICK. The Botanical Works of Philippi, Father and Son, in TURNER, BILLIE L. (See Chia Jui Chen, Meghan C. Mendenhall & Billie L. Turner) VAHIDY, AHSAN A. (See M. Moinuddin, Ahsan A. Vahidy & S. I. Ali) —— VAHIDY, AHSAN A. (See S. M. Nadeem Ahsan, Ahsan A. Vahidy & S. I. Ali) VaHIDY, AHSAN A. (See Zeenat A. Razaq, Ahsan A. Vahidy & S. I. Ali) ~ VaHIDY, AHSAN A. (See Bushreen Jahan, Ahsan A. Vahidy & S. I. Ali) — VocEL-BAUER, INA. (See Uwe Jensen, Ina Vogel-Bauer & Marei Nitschke) WEBSTER, GRADY L. Systematics of the Euphorbiaceae. Introduction to the Symposium WEBSTER, GRADY L. Classification of the Euphorbiaceae . WEBSTER, GRADY L. Synopsis of the Genera and Suprageneric Taxa of Euphorbiaceae .... WERFF, HENK VAN DER. Book review: Australian Tropical Rain Forest Trees YATSKIEVYCH, — Book review: Flora of Australia, Vol. 50, Oceanic Islands ZECHMAN, nidis W. (See Brent D. Mishler, Louise A. Lewis, Mark A. Buchheim, Karen S. Renzaglia, David J. Garbary, Charles F. Delwiche, EEE Y, Zechman, Thomas S. Kantz & Russell L. Chapman) — ZIMMER, ELIZABETH A. (See James A. Doyle, Michael J. Donoghue & Eliz- abeth A. Zimmer) 419 Two Important New Titles from the Missouri Botanical Garden Moss Flora of Central America Part 1. Sphagnaceae—Calymperaceae Bruce Allen with contributions from H. Crum, R. A. Pursell, W. D. Reese & N. Salazar Allen Central America possesses one of the world’s richest moss floras, with an estimated 871 species in an area only one-half that of Colombia, which has about as many species. Previous to this publication the only moss flora for any Central American country was E. В. Bartram’s now nearly half-century- old уе ae Guatemala. s Flora of Central America is based on historic and extensive recently collected material ба E. area; each. species is кашу apse and eed AEn distribution in Central America ivid synonymy is given with types cited for all names. The Flora will алея in four parts during the next few years. Part 1 constitutes Monographs in Systematic Botany from the Missouri Botanical Garden, Volume 49. 242 pages. May 1994. $20.00. Index of Mosses Marshall R. Crosby & Robert E. Magill dex of meister ss isa уйе to anes names ме published for роз, ЖОЕ) ub new es ipo the rank of genus and below fi of the name, ising of parce acd names, хой types. ‘The Уй ar Mosses, 1963- 1989, lists ut 8500 na and contains appendices giving full authors’ nam ully spelled journal and book titles with da: to their entries in 77-2 and B-P-H. Ма уруу Mo: osses, 1990-1992, contains "im 800 names and a UM) e us рее іп hich nova for 1990-1992 appeared. onogr. in Systematic Botany fro e Missouri al Garden, Ap sna 42 (1992) and 50 "e 1994). 656 and 87 pages Mura ra, Vol. 42 $55 00; Vol. 50 $7. To place an order, please indicate method of payment below. Checks or money orders should be in US. funds, payable to Missouri ME 1 nig through a U.S. bank; U.S. ae add $2.00 for one book and $.75 for each additional book; non-U.S. shipments: add $3.00 for one book, and $.75 lor each additional book. Orders җые be repaid; a $1.00 fee will be A to orders requiring invoices. No shi pments are made until payment is received. Mail form, with payment, to: copy(ies) of Monograph No. 49 Ул Eleven, Missouri Botanical Garden Please send i No. 42 St. Louis, MO 63166-0299, U.S.A. copy(ies) of Monograph No Please send Please send copy(ies) of Monograph No. 50 D Check / money order enclosed | Send books to: Û Send invoice ($1.00 fee will be added to total) o Charge card number (Mastercard/ Visa) Name Expiration date __ А = = dares Name as it appears on card Telephone number (daytime) Postal Code Country | PRICES ARE SUBJECT TO CHANGE WITHOUT NOTICE 81(4) CONTENTS Revision of Hillia (Rubiaceae) Charlotte M. Taylor Systematics and Evolution of the Genus Petrea (Verbenaceae) Ricardo M. Rueda ... Systematic Implications of Pollen ee in Subfamilies Lamioideae and ная noideae (Labiatae) Mones S. 9 sab & Philip Р. Cantino 1 Classificati hidoideae: Orchid ) Н. P. Linder & H. КЕРЕР Taxonomy of Thermopsis (Fabaceae) in North America Chia Jui Chen, Meghan G. Mendenhall & Billie L. Turner The Botanical Works of Philippi, Father and Son, in Chile Charlotte M. Taylor & Mélicia Munoz-Schick noz-S Una Evaluación del Conocimiento Floristico de Mexico Victoria Sosa & Patricia Dávila EES Distribution of Ki it its Possible Relationship to the Extant s Cnemidaria e Barbara A. R. Mohr & David Dd E ААА REE ОООО io e O AO DI FO M Ede Ch N li (G ) Guillermo A. Norrmann, Camilo L. Quarin & Pie NS жт то V iA RM ET ARN A E IO E AE gl Chromosome Numbers and tees of Polyploidy іп Panicoideae (Poaceae) from Pakistan S. M. Nadeem Ahsan, Ahsan A. Vahidy & S. L Ali Chromosome Counts in Aids oideae, Chloridoideae, and Pooideae (Poaceae) from Pakistan M. did nuddin, Ahsan A: Vahidy-& S.J. Ali ас Chromosome Numbers in Some Taxa of sree Mostly Native to Pakistan Bushreen ahan, Ahsan A. Vahidy & S. I. Chromosome Numbers in йы x Pakistan Zeenat A. Razaq, Ahsan А. ahidy & S. 1. Ali .... Book Reviews Australian Tropical meus s Trees by B. P. M. Hyland & T. Whiffin, reviewed b nk van der Flora of Australia, RE si та Islands 2, reviewed by George AES LU 8 Checklist for Authors Cover illustration. Petrea sulphurea М. J. Jansen-Jacobs, by Phyllis Bick. 571 653 687 784 192