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IVEW3ITY NO. CXCIII a TAXONO]L'lic STUDY of the GE!\CS

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IVEW3ITY NO. CXCIII a TAXONO]L'lic STUDY of the GE!\CS COi\TI:II)UTIONS FI:O~l THEGI:AY II ERB .~ R lml Of IlAl:VAltD U;\IV EW3ITY J::. lilt".1 b~· lh't',l c. Hl'Jl!i n ~ ull.I Hobtc n c. !-"ht, ~ NO. CXCIII A TAXONO]l'lIC STUDY OF THE GE!\CS ANADE Ai:'i THERA By S ItU Y01'\ REI S ALTSCHl"L Il\FRASPECIFIC VARIATION IN THE GUTlERREZIA SAROTHRAE CmlPLEX (COMPOSITAE-ASTEEEAE) By Orro T. SOLBRIG Published by THE GR AY HERBARIUlII OF HARVARD UKIVERSITY CAMBRIDGE, MASS., U. S. A. , MANUFACTURED BY THE LEXINGTON PRESS, INC. LEXINGTON. MASSACHUSETTS A TAX O:\O,llC STUDY OF TilE GE~ljS AN A DENA:\TI-IEI~A SIR! VO:-i HE!S ALT~l-HUL The "nail, l ropical to suutropical, and strictly :\ew World genu s .AlIlLl!C"1Wllth c,'n formerly was I...'ons ide}' ~d n~ $~c tion NioJ)(I of the genus Pip to.(/ cllia and i$ the cor.',m on It.'gumi­ 11 0 U S sou rce of lhe so-called narcotics known as COIIO/>O, l-iica a nd Yopo, Because of lheir ullusual c"rects upon th~ human nervollS system, the chemical constitutt.'nts of t~.ese material s are of cUlTen t interest in the field of ~xper im " nta l psychia­ lry, Among the chemical compound, which leave u.een iso­ lated from the species of A,wde1U",th ,Ta, may be included some hallucinogenic cl!'ugs designated as psychotomimetics (Hofmann, 1959) , In contrast to otlwr psychotropic drugs, which merely evoke mod ifications in the mood of th~ reci pi­ ent, the psychotomimetics appeal' to produce profound and acute changes in perception_ An interest both in plants of possibl~ medical significance and in species in need of t"xonomic stud,' provided a stimulus for the research whose results are published in part in this paper. An elhnobotanical treatment which a;so has been prepared will be pu blished elsewhere, _-\ detailed s"Hematic treatment of the species of An(L(/.e"'lIIlhcm has been needed for some tim e. While additional work still remains to be carried out on lh is group, as " 'ell as on related species in the genus P ip ladenut, it is hoped that the data presented here from several branches of botany will help bette,. to define Anadenct1!lhem and to stimulate further research in the large and relatively neglected subfamily Mimosoicleae, ACK :.\"OWLEOGE~JEXT5 I would like to express my deep appreciation to Pr,) Ies:::or Reed C. Rollins for guiding the course of my research and for [.is helpful criti­ cism in the pl'epan~.tion of this paper. Professor John H. Welsh aided in the phy~iological aspects of the problem, with regard to cunent research on the effeC'~.s of A.71adenan~ them dui\'ativcs and related compounds upon the r.erVOlli: system. J'll'ofe ssors 1. W. Bailey and Tayior A. Ste-e\'es were kind enough to s ug~est sources of in formation on leaf vcn3tion patterns and foliar anatomy. Professor Gordon R. Willey provided assistance with regard to archaeological data. In particular, I am grateful to Professor SIRI VON REIS A Rid.lI;r.d A. Howard. to Dr. Richard E. Schulles and to D r. Rolla M. Tl'!-.'(.Il for their careful scrutiny of the manllscl"ipt in the last phases. 1 rlso wish to thank the late Dr. C. E. Kobuski, and the other c·~"'3.h·s who made possible the loans of .materials fl'om the following r·:·rb;l·ia: Arnold Arboretum (A); Brit.ish Mu seum of Natural His tory (8M) ; Chicago Natural History Museum (f) j Copenhagen Botanical M ~ fie , .:n and Herbarium (c); Gray .Ficl'bul'iul1l (Gl-I); Kew Herbal'ium ex) i I~ ~ksh crbari um. Leiden (L); Missouri Botanical Garden (:'\IO); .N t" · York Botanical Garden (NY); Museum National d'Histoirc .No..~ Paris (p); Naturhistoriska Riksmuseet, Stockholm (s); I iiAW M- National Herbarium (us); Utrecht Botanical Museum ~ii0k-~). ! ;0{." /2.vbert C. Foster's scllolal'l y interpretations helped to cl arify ~1Wt1&iabou't styli stic forms of early taxono,mic literature. Dr. ~~ E. 'Wood, Jr., offered helpful suggestions as to presentat ion !l! .....wt~rials. Dr. ~i ly M., Pel'TY contributed fl"o m het' understanding -CL .,..,.Cles from Paclfic TcglOns. I \\ ould like to thank DJ'. Joab L. Thomas for instruction ill certain ::-: :F:r at f' l'~' techniques and Dr. Alexander Grobman for exchanging data .froill his Own current research. :.rl .~. 1.. Schwarten's willing assistance in the Herbarium Library. well u,,; that of Miss M. Currier in the Peabady Library, expedited ~ h:l"ldl ing of the literature. Dr. Charles Dills is to be complimented r h.s leaflet photographs, as are Messrs. Arthur G. B. Metcalf and Ken neth J . Wilfert for habit ph6fographs from Puerto Rico and FlOJ':'Ja, respectively, I ,vant to thank :M r, and Mrs. Mulford B, f'o~~e:r, of Ol'l ando, Florida, as well as the kind personnel of the E s!. ; ~ ci6n Experimental de Agricultura, at Rio Piedras, Puerto Rico, for time spent with me in the field, I would li ke also to express my 3!)preciation to Mr. Fl'ed H. Chambel'lin and to DJ', Alice F. Tryon fO l' their help in preparing this paper. Finally, I am deeply grateful to my parents, Gustav and T ~ryne von Reis, for their financial suppur t a nd to my deal' f r iend, Arthul' G. B. Metcal!, for facilitating in countless ways thf' C0l11111et ioll of this work. THE GENUS ANADENANTHERA: HISTORY AND THE CHARACTERS OF DISTINCTIVENESS The species comprising the leguminous genus Anadenal1­ them as here conceived were treated originally as belonging to section Niopn of the genus Piptwlenin Bent ham (1840, 1841-42). The species of Piptndenia. were not well known during Bent ham's lifetime, but that author was quick to recognize that a l1atural distinction cou ld be made between sections E-upiptaclenin and PitYl"Occo'pa with spicate inflores­ cences, and section Niop" with capitate inflorescences. Ben­ tham distinguished five species in section Niopa, though he wondered whether they might not better be considered as A TAXONOMI C STUDY OF THE GE:-':Lf:::: A~ADE~ANTHER ..\ 5 varieties of a single species. The f ruits differed considerably in [orm and proportion, but the flowe r:; and foliage wcre very much alike from one specimen to another. Benthalll later included four of the original speci es in hi s "Revi si,m of the Suborder Mimoseae" of 187~ - 75: Pip/adcnia ]incyri"a (L.) Bentham, Y 11Il1croC((1'}la Bentham, P . fa/clt/a Bentbam and P. col"b,.;'ln (VeiL) I:entham. He placed a fiith spc",ies, P. "llticrophyll" Bentham, in the s ~'lonym~' of P. macn>ca,·pct. Since Bentham's time, three additiL' nal species have been desc ribed for section Niopa. of P ip/ad,."ia. Pip/ad..,li" 'I101}0­ [Jllincensis Warburg "'as published first as the monotypic genu s Sch/cinitzilt microphylla Warburg (1891) alld, for reasons di scussed in this paper, l)1'obabl,- should be returned to its former status. P ip/adr "ia H a8~I< ' riltna Chodat (1904) is I', ferred to synonymy under A"cule" a"fhe1'({ collll>ri"n var. Cl'iJii in the present treatment. P i p/ad c" i(~ amazollica Ducke (1915) ",as recognized by Du cke in 1922 as belonging to Pi/hccellobiwlt Mmtius (1837) . Pip/ndell;{' leptoc/ad..'t Baker (1887) "'as based on a speci­ m ~ n with a globose-spicate inflorescence ",hich could be mis­ tu!(en at first glance for a speci es intermediate between sedions 1 ancl 11 and section III of P i}J /ade ll ic, S(>II$II Ben­ thnm. The specimen was identified b)- Penier de la Bathie in 1938 as D es llumthus COlnmerSOnialllls Baillon (1:383) . In 1923, Spegazzini founded the genus A nadella 11 / hem to accommodate Piptadenia. pc reyrin{( and P . fa/ca /a. He based the genus on the flowers' being "without anther glands", a cha racter of doubtful generic "alue b.,' itself. Spegazzini stated in his description that he did not in clude in Alladellan­ them fi ve southern species he had not seen, but ",hich " 'ere • mentioned in Engler & Prantl (1 892 ). An inspection of those authors' work suggests that Spegazzini must ha" e meant three, not five, othe1' species. For the species named in Englel' & Prantl are: Pill/ac/eni" wr"[J";"a. P. fa/cata, P. macrocarpa, P. cO/llb"ina and P . llol'o-Ylli"eensi.s. The last Lhree have anther glands. In 1927, Britton & n ose proposed to elevate the ",hole of section Niopn to a genus. Th e~' offe red, ho\\'e"er, no reason for the change, added no new in formation and did not recog­ nize the priority of the name Anadcllail fhe1'(1. In 1955, Brenan proposed a r e vi ~ i o n of the genus Pillta.­ ·. 6 SIRI VON REIS ALTSCHUL denia according to a new set of characters involving, in t,articular, the mode of dehiscence of the pod and the struc­ 'tf,re of the seed. The differences he prescribes are compar­ abl(' to those used elsewhere in the Mimosoirleae to ~bl inguish genera. Furthermore, they are not infrequently correlated with vegetative differences. Brenan presents fruit ~~?wer keys to eight genera which have species formerly ~e d in Piptadenia. These keys delimit groupings which ~;W be more satisfactory than have been achieved here­ ~ .
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