Delivered by Publishing Technology to: Alice Calvente IP: 200.19.175.8 on: Wed, 31 Oct 2012 22:27:32 Copyright (c) American Society for Plant Taxonomists. All rights reserved. O 10.1600/036364412X656455 DOI © Botany Systematic hpaieei h ags ao feihtccci tencom- It 2006). cacti. genera: al. epiphytic four of et passes taxon (Hunt largest species the cacti is Rhipsalideae epiphytic most include and Cactoideae subfamily to belong tribes Rhipsalideae The regions. and Hylocereeae humid of epiphytes are some species but areas, arid epiphytic and and lithophytic species 1993). Hunt and terrestrial winged (Barthlott columnar or and family, cylindrical 2001). the globular in (Anderson variation leaves morphological including widest reduced the greatly leaves presents character- or It is taxa, subfamily absent persistent of by number diverse highest ized cylindrical, the most and tribes the nine small, with Cactoideae, and 2001). (Anderson stems spe- culent 2001). shrubby genus two the (Anderson with of and subfamily cies cylindrical aril, small a often bony is a and Maihuenioideae by flattened deciduous covered stems seeds leaves have glochids, species cylindrical, per- regular or Opuntioideae present and leaves. terrestrial are sistent 2005). al. group et this (Edwards in Cactaceae Species in the diverge to includes lineages Pereskioideae two 2006). first four Asia al. and into et and divided Cactoideae, (Hunt currently Africa Maihuenioideae Opuntioideae, is in family Pereskioideae, occurs The subfamilies: also 2006). al. which et Stearn, (Hunt Muel.) S. for (J. except Neotropics, the throughout uted h ags.Arcn oeua hlgn fRhipsalideae of phylogeny molecular of monophyly recent the supported A largest. the Rhipsalis reduction. habitat threat- to are due sev- and ened and ranges distribution Brazil, restricted to have of endemic species majority are eral tribe The this 2011a). to belonging al. species et (Calvente circumscribed ously that cated Rhipsalis .Schum., K. alr19) h ugnrccasfcto of classification subgeneric The 1995). Taylor Berger, urn drs:Dpraet eBota de Departamento address: Current oyih 02b h mrcnSceyo ln Taxonomists Plant of Society American the by 2012 Copyright otCcoda aescuetsesadihbtdyor dry inhabit and stems succulent have Cactoideae Most distrib- species 1,438 and genera 127 includes Cactaceae h otrcn n opeesv aooi ramn of treatment taxonomic comprehensive and recent most The Laborato ecito n ito pce nlddaepeetd e o h dniiaino ugnr sas provided. also is subgenera of identification the for key A presented. are included species of list a and description pce rmsubgenus from species h omrsbeei lsiiain rpsdfrtegns he oohltcsbeeaaerecognized, are in included subgenera previously monophyletic species Three of genus. inclusion the re-evaluate for to used proposed is Erythrorhipsalis classifications data morphological subgeneric with combined former information phylogenetic the molecular Here, morphology. stem by mainly characterized otoesa.Bten13 n 95 ih ifrn ugnrccasfctoshv enpooe for proposed been have subgenera, classifications five recognized subgeneric genus different the eight of treatment 1995, taxonomic and 1837 Between controversial. eu feihtccci pce of Species cacti. epiphytic of genus Keywords— Abstract— Phyllarthrorhipsalis arn,and Gaertn., nldd3 pce nfv subgenera: five in species 33 included Hatiora 21) 74:p.983–988 pp. 37(4): (2012), ´ i eSistema de rio Epallagogonium e ugnrcCasfcto of Classification Subgeneric New A otCcaeehv ucln tm n nai r rai ra,btsm r ppye fhmdregions. humid of epiphytes are some but areas, arid or dry inhabit and stems succulent have Cactaceae Most hc r anycaatrzdb lrltat.Tecagspooe nld xadn h icmcito of circumscription the expanding include proposed changes The traits. floral by characterized mainly are which , tatcFrs,eihtccci Rhipsalideae. cacti, epiphytic Forest, Atlantic and Maihuenia Hatiora Schlumbergera Schlumbergera Epallagogonium ´ ub,and Buxb., iaVgtl eatmnod Bota de Departamento Vegetal, tica ihclnrclo lbs suc- globose or cylindrical with , rto Rose, & Britton Rhipsalis .Schum., K. oNre apsLgaNv,CP 97-0,Ntl N Brazil. RN, Natal, 59078-900, CEP: Nova, Lagoa Campus Norte, do Rhipsalis u oMata do Rua e. fwhich of Lem., h icmcito of circumscription The . r aahltca previ- as paraphyletic are and Rhipsalis uhrfrcrepnec ([email protected]) correspondence for Author ˆ ia clgaeZooi,Cnr eBiocie de Centro Zoologia, e Ecologia nica, Phyllarthrorhipsalis r ooiul ifcl oietf,adtesbeei lsiiaino h eu a remained has genus the of classification subgeneric the and identify, to difficult notoriously are Lepismium Erythrorhipsalis Rhipsalis Lepismium hpai baccifera Rhipsalis omnctn dtr nrwHipp Andrew Editor: Communicating ˜ Calamorhipsalis Brhotand (Barthlott ,27 E:05800 Sa 05508-090, CEP: 277, o, Rhipsalis proposed u indi- but Pfeiff., lc Calvente Alice Phyllarthrorhipsalis Erythrorhipsalis and A. is Epallagogonium 983 ˆ ia nttt eBiocie de Instituto nica, Rhipsalis subg. Erythrorhipsalis as cassytha R. transferred huhGete 18)ue h aetp aeilof material type same the ing used baccifera Cassytha (1788) Gaertner though recognized also he as described nrgnrccasfcto Tbe1). (Table classification infrageneric of ment 73.Gete 18)sbeunl described subsequently (1788) Gaertner previ- 1753). the to due of level publication generic ous the at name illegitimate an was ti loipratt oeta dno 16)dsrbdthe described (1763) Adanson that note genus to important also is It published and combination baccifera equivocal this noticed nnmas(e akno 97 98 e oteconservation the to of led 1988) 1987, Adansonian Parkinson of (see application uninomials problematic the However, Muell. opie in comprised Rhipsalis within placed previously expanding subg. species include some of mainly inclusion of changes circumscription proposed the monophyletic The Only nized. recog- 2011b). are characters, al. data morphological by diagnosed et phylogenetic subgenera, Calvente molecular 2010; and (Calvente morphological on based for classification subgeneric new a Here, previously as ( paraphyletic circumscribed are subgenera three that indicated of phylogeny species 35 most recognized the which in in Cactaceae, adopted the also of was account (1995) recent Taylor and Barthlott by aooi History— Taxonomic eCnol 12)peetdtefrtcmrhnietreat- comprehensive first the presented (1828) Candolle De Rhipsalis ean nhne.Frec ugnsals fsnnm,abrief a synonyms, of list a subgenus each For unchanged. remains Rhipsalis .cassutha R. .cassytha R. , Rhipsalis Epallagogonium Epallagogonium Hariota ˜ al,S,Brazil. SP, Paulo, o J .Mel)Ser;ti stecrettp of type correct the is this Stearn; Muell.) S. (J. n eonzn broader a recognizing and subg. Rhipsalis J .Mel)Gete oiial nhspublication his in (originally Gaertner Muell.) S. (J. arn against Gaertn. Hn ta.20) oee,arcn molecular recent a However, 2006). al. et (Hunt astabaccifera Cassytha astabaccifera Cassytha , u hsi rsmbyatpgahclerror); typographical a presumably is this but , Ccoda,Cactaceae) (Cactoideae, Epallagogonium Rhipsalis Rhipsalis eoetepbiainof publication the before J .Mel)Gete leiiae tan(1939) Stearn illegitimate. Gaertner Muell.) S. (J. Rhipsalis ean unchanged. remains Rhipsalis eaotdadfeetseii pte,mak- epithet, specific different a adopted he , ˆ ca,Uiesdd eea oRoGrande Rio do Federal Universidade ncias, ihsvnseisrcgie without recognized species seven with , Cassytha ˆ .cassytha R. ca aUiesdd eSa de Universidade da ncias h icmcito of circumscription The . n ieigtecrusrpinof circumscription the widening and subg. Clet 00 avnee l 2011b) al. et Calvente 2010; (Calvente Rhipsalis , yteicuino pce previously species of inclusion the by Calamorhipsalis h is pce of species first The , Hariota Calamorhipsalis .wti h arca (Linnaeus Lauraceae the within L. Rhipsalis. Phyllarthrorhipsalis .S ul.However, Muell. S. J. .S ul.into Muell. S. J. stetp of type the as Calamorhipsalis Rhipsalis subg. Adans. Rhipsalis h otcomprehensive most The Rhipsalis Calamorhipsalis ,and , ,and astabaccifera Cassytha Calamorhipsalis Rhipsalis loincluding also , a subsequently was Erythrorhipsalis Rhipsalis Epallagogonium stelargest the is Rhipsalis Rhipsalis Rhipsalis ˜ Rhipsalis and Paulo, o Rhipsalis sproposed is .baccifera C. ythe by Rhipsalis Rhipsalis Rhipsalis Rhipsalis and ythe by Even . subg. , was and .S. J. as ). . Delivered by Publishing Technology to: Alice Calvente IP: 200.19.175.8 on: Wed, 31 Oct 2012 22:27:32 Copyright (c) American Society for Plant Taxonomists. All rights reserved. subg. into of division subgeneric and rec- also ognized who (1850), Salm-Dyck by adopted subgeneric this was classification of version modified A 1837). (Pfeiffer stems the Articuliferae .Schum., another of K. as taxa morphology infrageneric ( subgenera flower eight of described feature use diagnostic to key and subgenera into genus to recognize not of did (species and 1890), Schumann in (cited fifr13 Divided 1837 Pfeiffer Salm-Dyck, oiindwithin positioned 8 YTMTCBTN Vlm 37 ( [Volume series four into subdivided BOTANY SYSTEMATIC Divided for species seven Listed 1850 Salm-Dyck 1828 Candolle De 984 aplrfr otelwrpr fteccu lwrincluding Zappi within flower and Taylor cactus 2004), tissue; stem the in embedded of peri- is which part term ovary, lower the (the the pericarpel to immersed refers with carpel species all placed Epallagogonium Schum., cuan19 Divided 1890 Schumann cuan19 Divided 1899 Schumann Lo ute l 06Floe h ugnrccasfcto fBrhotadTyo 19)adrcgie iesubgenera: five recognized and (1995) Taylor and Barthlott of classification subgeneric the Followed 2006 al. et Hunt atlt 97 atlt and Barthlott 1987; Barthlott rpsdb cuan(89,btwdndtecircumscrip- of the tion widened but (1899), Schumann by proposed ubu 90Rcgie h rb yoeea,sbrb hpaia ihtre“ia” fifre clmegra,adRhipsales. and Schlumbergereae, Pfeifferae, “linae”: three with Rhipsalinae subtribe Hylocereeae, tribe the Recognized Divided 1970 Buxbaum 1923 Rose and Britton Lo lcdwti genera within placed and Loefgr. rb hpaiaa oehrwt h el ecie genera described newly the Erythrorhipsalis with together Rhipsalidanae tribe n oe(93 i o olwtecasfctosproposed classifications Lo the or Rhipsalis follow (1899) not Schumann did by (1923) Rose and ¨ ¨ ge 95 97DsrbdsvrlBaiinnwseisadpoie aooi ramn dpigtesbeeapooe by proposed subgenera the adopting treatment taxonomic a provided and species new Brazilian several Described 1917 1915, fgren cuan(89 a h is uhrt iietegenus the divide to author first the was (1899) Schumann Table ge 11,11)sbeunl cetdteegtsubgenera eight the accepted subsequently 1917) (1915, fgren alr1995 Taylor Perpetuae Articuliferae Erythrorhipsalis Rhipsalis .Hsoyo h nrgnrccasfctosof classifications infrageneric the of History 1. Acanthorhipsalis no1 e eis hyincluded They series. new 16 into Sarmentosae Hariota Author ,caatrzdb h hp n ipsto of disposition and shape the by characterized ), Articuliferae Hariota Acanthorhipsalis Ophiorhipsalis Calamorhipsalis and egr(urnl oiindwithin positioned (currently Berger .Shm,and Schum., K. ). oicuetoadtoa ugnr ( subgenera additional two include to cuan(80 anandtesame the maintained (1890) Schumann . Terminatae Schlumbergera ofr) ohicuigseiscurrently species including both Loefgr.), ), nadto to addition in Rhipsalidopsis Rhipsalis eetaserdt series to transferred were Rhipsalis .Schum., K. Eurhipsalis ¨ rnfre pce from species Transferred ge 11,11)addivided and 1917) (1915, fgren , .Schum., K. K cu. rto Rose, & Britton Schum.) (K. Epallagogonium, Alatae , Alatae species), Sarmentosae h subgenus the Rhipsalis Teretes oeseis.Fu pce eelf s“netesedis.” “incertae as left were species Four species). (one Eurhipsalis Calamorhipsalis 1 pce) and species), (10 subgenera: oespecies), (one in included Current genera: new two and (1899) Schumann oeseis rvosyin previously species, (one codn oG .Lindberg A. G. to according Hatiora, nsre u divided but series in n h genera the and ) Lepismium Rhipsalis Rhipsalis Rhipsalis Rhipsalis Calamorhipsalis Rhipsalis rto oe(currently Rose & Britton Alatae , sxspecies), (six 1 species), (11 .Schum., K. Angulosae Rhipsalis Phyllarthrorhipsalis sdvddi orsubgenera: four in divided is 1 pce,icuigseismsl rmprevious from mostly species including species, (18 Rhipsalis and oeseis and species) (one Angulatae , Rhipsalis Rhipsalis 4 pce,including species, (47 5 pce)it 6sre salgop fmrhlgclsmlrspecies). similar morphological of groups (small series 16 into species) (57 1 pce)it orsre ae nse hp icuigseiscretypae in placed currently species (including shape stem on based series four into species) (16 3 pce)it orsre ae nse hp icuigseiscretypae in placed currently species (including shape stem on based series four into species) (36 Calamorhipsalis Phyllorhipsalis , Angulosae 1 pce)it ieseries: five into species) (19 Epallagogonium .Shm)and Schum.) K. Erythrorhipsalis Pfeiffera and pce r oiindi genera in positioned are species Angulatae Angulosae , sxspecies), (six Rhipsalis Goniorhipsalis Rhipsalis Rhipsalis Teretes, ), Articuliferae nofu series: four into Teretes Rhipsalis .Schum., K. Lepismium ntesub- the in Ophiorhipsalis , Britton . Rhipsalis Angulatae Pfeiffera Pfeiffera Teretes, =emnte current (=Terminatae, Alatae nn species), (nine ihu subdivisions without trespecies), (three Articuliferae sxseiswt oeseisof species some with species (six . .He , and and Phyllarthrorhipsalis, svnspecies). (seven to K. Calamorhipsalis . Lepismium Lepismium ) Lepismium Ophiorhipsalis tospecies), (two Pfeiffera cu. n hpae:genera Rhipsales: and Schum.; Rhipsalis Mesembryanthoides subgenus the subdivided and species) winged cies), Phyllorhipsalis divided and Hatiora Acanthorhipsalis genera (Pfeifferae: “linae” three with Rhipsalinae Rose. & Britton Pseudorhipsalis 20)divided (2006) to to assigned transferred previously were species several although urnl lcdi h genus the in placed currently avnee l 21a eosrce oeua phylogeny (1995). molecular Taylor a recognizing and reconstructed Rhipsalideae for (2011a) Barthlott al. and et (1987) Calvente Barthlott by posed Erythrorhipsalis hltc(scrusrbdb ute l 06.I hssame this In 2006). al. et Hunt by circumscribed (as phyletic iiainas eonzdfv ugnr within subgenera five ( recognized also sification of part Lepismium middle ind the and segment) branching from segment) the the primarily of (branching base mesotonic the of and from species primarily all (branching almost basitonic transferred (1995) Taylor classification. Buxbaum’s under itself oefoe hrces h aetteteto Cactaceae ( of genera treatment four recognized 2006) latest al. The et (Hunt characters. flower some and treseis nldn n pce rvosypstoe in positioned previously species one including species, (three Goniorhipsalis Rhipsalis Alatae oercnl,Bxam(90 eonzdtesubtribe the recognized (1970) Buxbaum recently, More h lsiiain fBrhot(97 n atlt and Barthlott and (1987) Barthlott of classifications The Teretes Alatae iie h eann pce of species remaining the Divided . pce)it ih ugnr ae nse n lwrmorphology: flower and stem on based subgenera eight into species) Erythrorhipsalis and Phyllarthrorhipsalis fv species). (five trespecies), (three , 1 pce,dvddin divided species, (11 Rhipsalis and ,and Hariota fv pce)and species) (five , , sxspecies), (six Rhipsalidopsis Notes: Erythrorhipsalis recognizing , Mesembryanthoides Phyllorhipsalis Calamorhipsalis , Erythrorhipsalis Phyllorhipsalis Rhipsalis . rto Rose, & Britton Shubreee n in and (Schlumbergereae) =Preue current Perpetuae, (= igdseis and species) winged ;Schlumbergereae:genera Rhipsalis Schlumbergera , Teretes ,mil hrceie yse hp and shape stem by characterized mainly ), Epallagogonium Cereusculii, Teretes Pfeiffera Angulosae Rhipsalis 1 pce,previous species, (11 Shubreee and (Schlumbergereae) , Rhipsalis ub =Triaa,current Terminatae, (= Buxb. Articuliferae nofu subgenera: four into . , Schlumbergera, , n te from other and Rhipsalis notesm iesbeeapro- subgenera five same the into =Preue current Perpetuae, (= ), Cereusculii, Epallagogonium Perpetuae Goniorhipsalis nti lsiiainHn tal. et Hunt classification this In . Lepismium fu species), (four ihntieRhipsalideae, tribe within ) nadto,Bxam(1970) Buxbaum addition, In . triaegot otegenus the to growth eterminate Lepismium inanarrowersense.Thisclas- and svnspecies), (seven Rhipsalis Rhipsalis nfu series: four in tospecies). (two and Rhipsalis Rhipsalis and Rhipsalis Lepismium Terminatae Angulatae and treseis previously species, (three Rhipsalis efrhrsubdivided further He . , and Alatae Rhipsalis Hatiora Phyllarthrorhipsalis, fif and Pfeiff. eeicue in included were 3 pce)it five into species) (33 Teretes Rhipsalis Rhipsalis and Erythrorhipsalis Lepismium otspecies Most . Phyllarthrorhipsalis ), . Zygocactus )and ), Goniorhipsalis Rhipsalis Ophiorhipsalis Acanthorhipsalis igdspe- winged Epallagogonium Pfeiffera , fv species), (five Lepismium (Rhipsales) Lepismium Lepismium (Rhipsales). Sarmentosae Lepismium Lepismium Hatiora smono- as Rhipsalis Rhipsalis Rhipsalis Hatiora winged with and ) K. ): ): ) , , , , , Delivered by Publishing Technology to: Alice Calvente IP: 200.19.175.8 on: Wed, 31 Oct 2012 22:27:32 Copyright (c) American Society for Plant Taxonomists. All rights reserved. pce ae.Cae ,2 ntelws ato h iuewr eoee rmteaayi fIS+cDAdt e n eosrc h oiinof position the reconstruct and set followi data asterisk cpDNA an + by ITS indicated of species analysis type the subgenera from names. with recovered column, were species and first figure following ITS the the cpDNA, in of C the tree, part or in the lowest included B of the not right in grey) A, termina 3 the (in letters single species 2, to indicated a 1, with indicated by are Clades are represented indicated probabilities names. (2006) are are species species al. posterior monophyletic et species Bayesian bold; Hunt non-monophyletic in and to highlighted branches trimmed); according are above sampled were were indicated specimens specimens are multiple (additional which bootstrap for likelihood Species branches. maximum below and bootstrap parsimony Maximum icmcito of circumscription 02 AVNE YTMTC FRISLS985 RHIPSALIS OF SYSTEMATICS CALVENTE: found (2011a) al. et Calvente work 2012] h ergto of supports segregation 2011a) al. the et (Calvente Rhipsalideae of phylogeny ular Hatiora in positioned previously species the transferring paraphyletic, Fig. e ugnrcCasfcto fRhipsalis— of Classification Subgeneric New A .Mjrt uecnesste of tree consensus rule Majority 1. subg. Rhipsalidopsis Lepismium Rhipsalis to spooe yBrhot(1987), Barthlott by proposed as Schlumbergera from Hatiora Rhipsalis Rhipsalis . and MS eie rmteBysa nlsso pN ( cpDNA of analysis Bayesian the from derived n narrower a and Schlumbergera aast oiidfo avne(00 n avnee l (2011b). al. et Calvente and (2010) Calvente from Modified set. data h molec- The Clet ta.21b using 2011b) al. et analyses (Calvente phylogenetic molecular recent the However, exceeding never pericarpel. or absent trans- tubes actinomorphic, flower small, with these flowers and lucent cm In seven (2006). than al. longer ments et Hunt and classifications (1995) Taylor and Barthlott T and ITS MS Fg )d o upr nifaeei iiinof division infrageneric an support not do 1) (Fig. Rhipsalis psbA-trnH nldsol pce ihse seg- stem with species only includes psbA-trnH , trnQ-rps16 , , trnQ-rps16 rpl32-trnL Rhipsalis ,ISand ITS ), , rpl32-trnL subgenera MS ng , l . Delivered by Publishing Technology to: Alice Calvente IP: 200.19.175.8 on: Wed, 31 Oct 2012 22:27:32 Copyright (c) American Society for Plant Taxonomists. All rights reserved. ihrprpyei rntsrnl upre (bootstrap supported strongly not or values paraphyletic either orbrtdb h lwrmorphology. flower the by corroborated species “core all subg. includes within subgenera 0.88) within Epallagogonium = maxi- placed 0.81, bootstrap = previously bootstrap likelihood parsimony maximum mum 1.00, = bility not “core all pericarpel of composed subgenus a morphologically with recognize defined to well flowers propose and we broader lateral Therefore, areole. the rotate they in immersed patent as all together, hand, have mor- clear maintained other all a if the present characterization clade In Rhipsalis” phological 2011). “core al. the from et species Korotkova clas- proposed of the clades sification regarding smaller as bellow these discussion genus in detailed consistent the (see not in is confusing morphology the be overall because would practical distinction not is subgeneric choice this However, supported. previously species all includes 0.99) in = included like- maximum bootstrap 1.00, = lihood bootstrap parsimony maximum 1.00, rmte“uknprcre”caeaehr nlddi a in included here are all clade subgenus characterize pericarpel” broader “sunken hand, the other species from All the clade. pericarpel” “sunken on the to stem, belonging species “core the the in deeply not pericarpel of a immersed and species flowers pericarpel lateral remaining rotate patent The the the clade. in with found morphol- flower ogy the flowers perfectly matching stem, the rotate in immersed patent presents ugnrclvladapast emr sfli species the in useful in more consistent be not to is taxonomy. appears level it and genus, a level is the subgeneric shape in stem character Although clade). variable pericarpel” (=”sunken are l. that s. lineages main (1) 1): three (Fig. (“core traits in flower genus by mainly the characterized of subdivision a Rhipsalis lastria.Tefu eann ugnr ( subgenera remaining case, four this Calamorhipsalis in The and, rotate terminal. or terminal; always or lateral either and ulate to 0.99, belonging = Species 1.00). Erythrorhipsalis bootstrap = parsimony bootstrap likelihood maximum maximum 1.00, posterior corrobo- = (Bayesian is data probability 2006) phylogenetic al. molecular et the Hunt by rated 1995; Taylor and Barthlott 1987; While omaintain recognize to to “core be the in would clades possibility smaller One 1). bootstrap (Fig. 0.93, maintained parsimony is = probability and Rhipsalis posterior likelihood (Bayesian maximum support low with subg. yohssfrRislda ae tityo chloroplast on strictly ( based markers Rhipsalideae for hypothesis 8 YTMTCBTN Vlm 37 [Volume BOTANY SYSTEMATIC 986 psbA-trnH ifrn cnrowt ifrn oiinn of positioning different a with scenario different h “core The h sne eiapl ld Bysa otro proba- posterior (Bayesian clade pericarpel” “sunken The h rvoscrusrpinof circumscription previous The ookv ta.(01 eetypbihdaphylogenetic a published recently (2011) al. et Korotkova Rhipsalis Rhipsalis Rhipsalis < Rhipsalis ae nse hp.Ised hs nlsssuggest analyses these Instead, shape. stem on based .5 otro rbblt values probability posterior 0.75, subg. and , trnK Rhipsalis Rhipsalis Epallagogonium Phyllarthrorhipsalis ld ntemlclrpyoey htis what phylogeny, molecular the in clade ” , xetfor except , plus , ) (2) ”); Rhipsalis aepnetfoes hc a ecampan- be can which flowers, pendent have intron, trnQ-rps16 Epallagogonium subg. Calamorhipsalis ld Bysa otro rbblt = probability posterior (Bayesian clade ” Rhipsalis subg. Erythrorhipsalis .pentaptera R. matK Phyllarthrorhipsalis sprpyei if paraphyletic is .Terrslsidct slightly a indicate results Their ). .pentaptera R. Rhipsalis , Phyllarthrorhipsalis , .pentaptera R. species. ” rbcL and , . vntog ti weakly is it though even , Erythrorhipsalis ld ssbeeaand subgenera as clade ” rps3-rpl16 n (3) and ; (subg. Phyllarthrorhipsalis rvosyincluded Previously . Calamorhipsalis mre ihnthe within emerges hpai pentaptera Rhipsalis < Phyllarthrorhipsalis smonophyletic is 0.95). Epallagogonium , Calamorhipsalis and rpl16 Rhipsalis (Barthlott Rhipsalis Rhipsalis Rhipsalis Rhipsalis Rhipsalis < intron, are ) 0.5), and .l. s. ). ” , a rvosycniee useisof subspecies a considered previously was Goniorhipsalis of subspecies (including a sidered including excluding subgenus into .pacheco-leonis R. si h excluded 14 the ca. in genus in as entire occur also the (that of stems species branching terete most acrotonical in occur strictly also this by (that of characterized form “typical” is the subgenus that stated they subgenus nevertheless work, the for tion oe of posed Erythrorhipsalis yaoopis sdsusdaoe ti rfrnilto the preferential at wider is taxon it a good above, consider a discussed distinctive make As by characterized not synapomorphies. not is would it because clade level, subgeneric this so classifica- previous Even (in work their tions in classifications specific and sequencing) DNA shaferi it for efforts, R. material obtain despite to collections; possible private not in was available only is and sulcata R. “core inside of species all almost in shoots” occur extension basal also “indeterminate (which of presence the and hi ok ookv ta.(01 lobifypooe a proposed briefly also (2011) for division al. subgeneric et different Korotkova work. their ta.(01 characterize (2011) al. et or to subgenera addition In these and here) identification. of circumscribed their (as each for for key consis- diagnosis a a explicit present not and do tent authors which these However, in subgenera. inference, logenetic Korotkova since (2011b), al. included et not Calvente did (2011) and al. et (2011) the al. for reason et the of be positioning must same al. conflicting The et clades. Calvente these in shown for support and (2011b) and relationship resolution this spe- better in the change these for the among for responsible relationships were of cies) resolution better offered nldsti clade. this includes shvn pce ihtrt tmsget n woolly and other all segments exclude they stem while includestems, terete but with areoles sunken species having as ugnsfor subgenus excluding but here of sampling complete including more (especially a with regions together chloroplast species all for sso morphology. of ysis of separation The Calamorhipsalis areoles. sunken woolly 14 (ca. (2011a) al. et Calvente found clade.” also pericarpel “sunken the to than paradoxa iiy=08.W eiv htteadto of addition proba- the posterior that 0.51, believe We = 0.8). (bootstrap = support bility low with ever ookv ta.(01 i o rvd omldescrip- formal a provide not did (2011) al. et Korotkova utemr ookv ta.(01 udvd cr Rhipsalis” “core subdivide (2011) al. et Korotkova Furthermore .shaferi R. Rhipsalis hc smr lsl eae o“core to related closely more is which , ti pce a ecie rmcliae material cultivated from described was species (this sasprt ao ste i o xlr infra- explore not did they as taxon separate a as .sulcata R. .baccifera R. .grandiflora R. .grandiflora R. .paradoxa R. Rhipsalis hpai paradoxa Rhipsalis (including .Clet ta.(01)ddfn ld com- clade a find did (2011b) al. et Calvente ). sntspotdb u eut rb u anal- our by or results our by supported not is ). a osdrdasbpce of subspecies a considered was Rhipsalis pce)a elas well as species) .grandiflora R. and Rhipsalis .paradoxa R. ld Fg ) u hyddntinclude not did they but 1), (Fig. clade ” , oerltdt “core to related more .teres, R. , Calamorhipsalis , .shaferi R. Rhipsalis .lindbergiana, R. .ewaldiana, R. .lindbergiana R. a rvosycrusrbdbut circumscribed previously (as .paradoxa R. Calamorhipsalis .baccifera R. .trigona R. MS subgenus , ,te eonz separate a recognize they ), .lindbergiana, R. and ( Rhipsalis Rhipsalis hc a rvosycon- previously was which , Epallagogonium and scrusrbdi their in circumscribed as Rhipsalis Rhipsalis .mesembryanthemoides R. .paradoxa R. rpl32-trnL hc a trigonous has which , (excluding hw yKorotkova by shown and ), and , Rhipsalis ae nterphy- their on based .pentaptera, R. sdvddi six in divided is Phyllarthrorhipsalis rpl32-trnL a circumscribed (as .paradoxa R. ihtrt stems terete with .pittieri R. .ewaldiana R. .floccosa R. Rhipsalis, pce,a well as species, Rhipsalis Erythrorhipsalis MS and .Korotkova ). eune in sequences hc has which , .baccifera R. .paradoxa R. sequences Rhipsalis Rhipsalis htalso that .shaferi R. which , ,which how- ” ,and ), subg. from and and ), ). ” ) ) Delivered by Publishing Technology to: Alice Calvente IP: 200.19.175.8 on: Wed, 31 Oct 2012 22:27:32 Copyright (c) American Society for Plant Taxonomists. All rights reserved. nlddin of included circumscription the expanding subgenus changes include The mainly presented. is proposed features morphological by terized Rhipsalis clas- the practical. make more to sification Rhipsalis” “core of clades smaller remaining giino broader a of ognition hpai, oee,hr hyaepstoe nawider a in positioned are they Rhipsalis here however, Rhipsalis,” nnmu eiwr n h soit dtrAde ipfrcom- for Hipp Andrew editor manuscript. the associate on the ments and reviewers anonymous .Prcre o mesdi h roe. areole the in immersed not Pericarpel 1. ap,Load esex ebr fLu Lu of support; members Versieux, financial Leonardo for Zappi, IAPT and FAPESP thank R unchanged. remains subg. “core other oerltdto under related assigned more is Calamorhipsalis and taxa of inclusion infra-specific the follows it therefore, atlt,W 97 e ae nRisldne(Cactaceae). Rhipsalidinae in names New 1987. W. Barthlott, 2001. F. E. Anderson, 1763. M. Adanson, areole the in immersed Pericarpel 1. 987 RHIPSALIS OF SYSTEMATICS CALVENTE: 2012] ic idppltost ofr hspsto n the and as taxon position subsp. this of classification this previous the confirm maintain to with populations relationship dis- from wild specimens different tinct and markers includ- DNA species, nuclear this also with ing conducted it be as should study surprising, detailed is resembles (2011) and areoles al. sunken has et Korotkova of by placement indicated The character. ical circumscrip- proposed new of tion this that recognize authors these ( floccosa R. pittieri R. included also stems but winged group was or this flattened subgenus in with this species the for maintained (2011) character of diagnostic classifications further this in followed and of 1970), use (Buxbaum stems the winged or flattened with species .ewaldiana R. HIPSALIS eo natraienwsbeei lsiiainof classification subgeneric new alternative an Below Acknowledgments. Phyllarthrorhipsalis .Sesclnrclo lvt;foespnet apnlt adltrlo pcli tmsegments) stem in apical or lateral (and campanulate pendent, flowers clavate; or cylindrical Stems 2. .Sesclnrclo igd lwr aet oaeadltrli tmsegments stem in lateral and rotate patent, flowers winged; or cylindrical Stems 2. 97–100. Hariota Epallagogonium Phyllarthrorhipsalis pittieri rrtt adecuieyaia nse segments) stem in apical exclusively (and rotate or rrl pclbti hscs aea lwr r lopresent) also are flowers lateral case this in but apical (rarely htol eonzsmnpyei ugnr charac- subgenera monophyletic recognizes only that subg. arn,Fut e.p.1 3.18,nm cons. nom. 1788, 137. 1: pl. Sem. Fruct. Gaertn., wihwspeiul osdrdasbpce of subspecies a considered previously was (which )in Rhipsalis Rhipsalis Phyllarthrorhipsalis dn. a.P.2 4.1763. 243. 2: Pl. Fam. Adans., lohsage tmsget) Nevertheless, segments). stem angled has also Bitn&Rs)Brhot&N .Tyo and, Taylor P. N. & Barthlott Rose) & (Britton .Calventeetal.(2011b)alsofound Rhipsalis ailsdsPlantes des Familles Phyllarthrorhipsalis Phyllarthrorhipsalis h atsfamily cactus The .floccosa R. aooi Treatment Taxonomic pce and species ” yteicuino pce previously species of inclusion the by Calamorhipsalis .Tecrusrpinof circumscription The ). soiial ecie otie only contained described originally as hswr spr fteP.D hsso .C I C. A. of thesis D. Ph. the of part is work This L including ITERATURE sntspotdb n morpholog- any by supported not is .Inthisworkitisproposedto and .grandiflora R. ai:Vincent. Paris: . otad ibrPress. Timber Portland: ...... C .ewaldiana R. .floccosa R. l aigtrt stems terete having all , ITED Epallagogonium ´ .pittieri R. Phyllarthrorhipsalis i omn’ a ru,two Group, Lab Lohmann’s cia as nldsseisfrom species includes (also ld Fg )ta othe to than 1) (Fig. clade Rhipsalis ...... 2 ...... floccosa R. Cassytha ´ i omn,Daniela Lohmann, cia .Thereforeamore , ookv tal. et Korotkova . .ewaldiana, R. nti ld as clade this in e oteSbeeaof Subgenera the to Key eae o“core to related Rhipsalis Erythrorhipsalis n l fits of all and .grandiflora R. n h rec- the and .S Muell., S. J...... floccosa R. Bradleya n the and subg. and 5: ...... avne .2010. A. Calvente, 1–5 Pp. Cactaceae. der System phylogenetische Das 1970. F. Buxbaum, 1923. Rose. N. J. and L. N. Britton, Steud., atlt,W n .P alr 95 oe oad oorp of monograph a towards Notes 1995. Taylor. P. N. and W. in Barthlott, 161–196 Pp. Cactaceae. 1993. Hunt. R. D. and W. Barthlott, .micrantha R. Rose, Zappi, .pulchra R. juengeri R. Dyck, Schum., Pfeiff., ex Dyck et.Foesptn,rtt,ltrlo ultrli stem in sublateral seg- areole. or the stem in lateral immersed in conspicuously pericarpel rotate, continuous segments; patent, not Flowers wings, narrow ments. with or angled Salm-Dyck. Pfeiff.) ex apnlt,aia rltrli tmsget rrotate or immersed segments areole. not the pericarpel stem in segments; stem in in apical lateral strictly and or apical campanulate, .R 1. Barthlott, & grandiflora R. Schum., K. ex crispimarginata R. .cereoides R. rto Rose, & Britton .R 2. Rhipsalis Rhipsalis .R 3. 1.19,sn nov.—TYPE: syn. 1898, 615. Rhipsalis eiaplntimre nteareole. the in immersed segments; not stem pericarpel in apical also rarely sublateral, or lateral rotate, pce Included— Species pce Included— Species Included— Species tmsget yidia,wt eldvlpdpodaria, well-developed with cylindrical, segments Stem tmsget yidia ocaae lwr pendent; Flowers clavate. to cylindrical segments Stem tmsget yidia r26wne.Foespatent, Flowers winged. 2–6 or cylindrical segments Stem atek 0 .1920.—TYPE: 4. 30: Kakteenk. at:65 1898.—TYPE: 615. Kakt.: rnk’ceverlagshandlung. Franckh’sche in family cactus Washington. of the Institution of plants of hpaiee(Cactaceae). Rhipsalideae Springer-Verlag. Berlin: plants Bittrich. vascular V. of genera and lies h .dsetto,Sa dissertation, D. Ph. HIPSALIS HIPSALIS HIPSALIS 44:17,sn nov.—TYPE: syn. 1970, 44–45: ad it,e.8 78 o . 1753.—TYPE: L., non 1768, 8. baccifera ed. Dict., Gard. 1.19,sn nov.—TYPE: syn. 1898, 615. .shaferi R. .puniceodiscus R. .triangularis R. .pachyptera R. R .pacheco-leonis R. i Kakteen Die HIPSALIS Loefgr., subg. subg. subg. atlt .P Taylor, P. N. & Barthlott Knh DC., (Kunth) Haw., .lindbergiana R. Bce.&Vl)Backeb., Voll) & (Backeb. .ewaldiana R. ug R subg. ug C subg. ug E subg. J .Mel)Stearn. Muell.) S. (J. rto Rose, & Britton .hoelleri R. .campos-portoana R. Epallagogonium Goniorhipsalis Loefgr., .cereuscula R. Phyllarthrorhipsalis ioei oeua,evoluc molecular, Filogenia .hileiabaiana R. .agudoensis R. .dissimilis R. Werderm. .aurea R. Pfeiff., d .Kan.Sutat Kosmos-Verlag, Stuttgart: Krainz. H. ed. , .Lindb., G. HIPSALIS ALAMORHIPSALIS RYTHRORHIPSALIS Loefgr., ˜ .cuneata R. al:Uiesdd eSa de Universidade Paulo: o atlt .P Taylor, P. N. & Barthlott atlt .P Taylor, P. N. & Barthlott .oblonga R. .pentaptera R. .Schum., K. .F ria .M .Braga, A. M. J. & Freitas F. M. Bradleya Haw., .sulcata R. oue4 ahntn h Carnegie The Washington: 4. Volume . .trigona R. hpai neves-armondii Rhipsalis G id. .Schum., K. Lindb.) (G. .paradoxa R. h atca:dsrpin n illustrations and descriptions Cactaceae: The N .Tyo atlt)N Korotkova N. Barthlott) & Taylor P. (N. .Shm,Gsmbsh.Kakt.: Gesamtbeschr. Schum., K. .P Taylor, P. N. .Shm,Gsmbsh.Kakt.: Gesamtbeschr. Schum., K. d.K uizi .G owr and Rohwer, G. J. Kubitzki, K. eds. , hpai paradoxa Rhipsalis rto Rose, & Britton Loefgr., .pilocarpa R. .ormindoi R. Loefgr., hpai pentaptera Rhipsalis 3 43–79. 13: hpai pilocarpa Rhipsalis ub nKan,Kakteen Krainz, in Buxb. , .Shm,Gesamtbeschr. Schum., K. hpai pachyptera Rhipsalis .Dietr., A. .mesembryanthemoides R. .A .Weber, C. A. F. Pfeiff. Rhipsalis Rhipsalis ¸a .Bre,Monatsschr. Berger, A. .crispata R. ˜ (Salm-Dyck .clavata R. sistema e o .olivifera R. .baccifera R. Loefgr. Rhipsalis .P alr&Zappi, & Taylor P. N. .goebeliana R. subg. subg. .russellii R. .elliptica R. .neves-armondii R. ˜ Paulo. o ´ iade tica .A .Weber, C. A. F. .floccosa R. ex Hw)Pfeiff., (Haw.) .P alr& Taylor P. N. Erythrorhipsalis Calamorhipsalis subg. .teres R. Ml. Stearn, (Mill.) (Salm-Dyck fif)Salm- Pfeiff.) .Schum. K. Loefgr. .burchellii R. .Dietr. A. Rhipsalis Rhipsalis. rto & Britton .Lindb. G. h fami- The Rhipsalis Backeb., Pfeiff. (Vell.) Haw., Salm- K. Delivered by Publishing Technology to: Alice Calvente IP: 200.19.175.8 on: Wed, 31 Oct 2012 22:27:32 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 8 YTMTCBTN Vlm 37 [Volume BOTANY SYSTEMATIC Molecular 2011a. Lohmann. G. L. and Forest, F. Zappi, C. D. A., Calvente, 988 avne . .C ap,F oet n .G omn.21b Molecular 2011b. Lohmann. G. L. and Forest, F. Zappi, C. D. 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