H eringeriana - 14(2): 157 191. 2020. Jardim Botânico de Brasília http://revistas.jardimbotanico.ibict.br/ Taxonomic and nomenclatural notes on infraspecies ()

Thiago E. C. Meneguzzo* ABSTRACT

Zygopetalinae: This Orchidaceaearticle presents nomenclatural notesBatemannia and keys on speciesParadisanthus and infraspecies of thePescatoria subtribe ( ), specifically the generaZygosepalum Lindl., Rchb.f., Rchb.f, Lindl., Hook. and Rchb.f., for which are proposed 15 new combinations,Key words: biodiversity, 44 new synonyms, neotropics, and 63 nomenclatural typifications. novelties, .

RESUMO (Notas taxonômicas e nomenclaturais em infraespécies de Zygopetalinae (Orchidaceae)

: NesseZygopetalinae artigo são Orchidaceae apresentadas notas nomenclaturais e chaves paraBatemann espéciesia e infrasespéciesParadisanthus da subtribo ( Promenaea), especificamenteZygopetalu m para os gênerosZygosepalum Lindl., Rchb.f., Rchb.f., Lindl., Hook. e Rchb.f., para os quais sã o propostas 15 novasPalavras combinações,-chave: biodiversidade, 44 novas sinonimizações neotrópico, e novida63 tipificações.des nomenclaturais, taxonomia.

Zygopetalinae Orchidaceae

The subtribe ( ) taxonomic morphological features. Their mainly occurs in the Neotropicset al. and has 37 generaet al. nomenclature had also been revised and whenever and ca. 400 species (Pupulin 2009, Chase necessary types were designated in order to fix the 2015). Based on molecular data, it is comprisedZygopetalum of use of the names, as well as keys for identification four lineages and groups: the have been provided. Since unpublished thesis et are al. clade/grade, theR.B HuntleyaLindl., the genus not considered effective publication (Turland et al. r., and theZygopetalum clade/grade 2018: Art. 30.9), the author himself hereby (Whitten 2005). The clade/grade publishes part of his findings concerning the matter had been subject of recent taxonomic revision by of infraspecies (Meneguzzo 2018). Meneguzzo (2018). It is basal to the subtribe and is MATERIAL AND METHODS composedeastern Brazil of 13 genera which primarily occur in and secondly in the Guianas Shield This study uses standard methods of alpha and the (Meneguzzo 2018). Several species taxonomy and was based on specimens examined were recircumscribed by the inclusion of new since 2008 from 98 collections in 86 herbaria synonyms and the combination of species to (acronyms according to Thiers 2020); those infraspecies for cases of constant but less relevant personally visited are in italic:- A (includes AMES, * Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Rua Pacheco Leão 915, Jardim Botânico, 22460 030, Rio de Janeiro, Rio- de Janeiro, Brazil; Universidade de Brası́lia, Instituto de Ciências Biológicas, Departame-nto de- Botânica,- caixa postal 4457, 70919 970, Brası́lia, Distrito Federal, Brazil. [email protected]. https://orcid.org/0000 0003 4760 9588.

157

ALCB BHCB, BM, BR Meneguzzo,CEN, hort. T.E.C. Colley sub J. Bateman s.n.

CEPECGH), AAU,, CESJ , ARIZ, B, , BRIT, C, , August- 1834, E , COAH,G COL, CR, G- DCCRI, GENTCTES, CUZ, CVRD, (lectotypeBatemannia K L barcode lepida K0003954 68!). Figurein Moore 1A. & HB, ESA, F, FUEL,HEPH (includes, HERBAM, HJ), , HPUJ,, GOET, HRB HAL, Reichenbach , HBG,HUEFS HBR, IAN, IBGE, HTO, Reichenbach— (1878:S. loc. , ex 558). hort. Type (designateds. leg. HUA,INPA ,K HUEM, HUFU,K-L HUTU,L U , ICN, here):s.n. BRAZIL. -R , January 1877, , JAUM,MBM (includes, MBML ), MG(includes , WAG), (lectotype W 38148s.d., E. onlyMorris bottom s.n. icon!). LE, LPB, M,OXF , MEXU,R, RB , MPU,des GFJP MO, ,MOL, GUA, Amazonas.- Negro River,S. loc., ex hort. E.K(excluded. Bailey NDG,ITA and NY, RUSU , P, PACA, PH, (incluSP, SPF UB, s.n.as type W R 30516!). -R , 1882, S. loc., UEC, UFMT ), RENZ,UPCB S, SBT, SEL, -, TO, s.d., (excludeds. leg. s.n. as type W 38148-R up icon!). , UFP, W, UPS (includesW-R UPS THUNB),WU (possible syn.syntype nov. W 38148 envelope US, USM, VEN, VIES, (includes ) and . The and bottomPetronia !), regia herbarium material identified by the author for this petroniaBarbosa Rodrigues (1877: study can be found at REFLORA (2020) and 107). nom. superfl. et illeg.Barbosa Rodrigues speciesLink (CRIA 2020). International (1891:— 131), Type (designated Code of NomenclatureNomenclature for follows Algae, theFungi and here): BRAZIL. Amazonas:J. Barbosa Manaus Rodrigues [“Manáos”], s.n. et al. Negro River, March, (Turland 2018). Types are designated when it (lectotype is the original illustration!in that Iconographie was to be was not possible to find internal or externalnal despublished Orchidées by J. duBarbosa Brésil Rodrigues evidence of a single element from the origi t. 299, deposited in the material and in a specific collection that strictlyet library of Jardim Botânico do Rio de Janeiro, bound correspondedal. to holotype (McNeill 2014, Turland in volume 5 of the book with same title published by 2018: Rec. 9A.1). Species and infraspecieset Barbosa Rodrigues 1996: 350, t. ’ 225!; typeonal al.definitions follows the discussion in Meneguzzo specimen João Barbosa Rodrigues pers (2015). New synonyms are proposed for cases on herbarium Batemannia destroyed). yauaperiensis Barbosa which there were evidences of discontinuation or — overlap of a state or set of character states amongst Rodrigues (1891: 131). Type (designated here): two or more previously accepted species. BRAZIL.June, J. Barbosa Amazonas: Rodrigues Jauaperi s.n. [“Yauapery”] River, RESULTS AND DISCUSSION (lectotype is theJ. original illustration!in that Iconographie was to be des published Orchidées by du Batemannia colleyi BarbosaBrésil Rodrigues 1. t. 854, deposited in the library of Jardim Batemannia colleyi colleyi in Botânico do Rio de Janeiro, bound in volume 6 of the LindleyLycaste subsp. colleyi book with same title published by Barbosa Lindley (1834: t. 1714).nom. illeg. Maxilaria(Lindl.) colleyi RodriguesBarbos 1996: 380,’ t. 252 C!; type specimen João Planchon (1858: 70), nom. illeg. aBatemannia Rodrigues personalwolteriana herbarium destroyed). (Lindl.) Planchon — (1858: 72), s. loc.Type, ex Batemannia wolteriana Schlechter (1915a: (designated here): GUYANA. Demerara: nom.28). superfl. Schlechter (1915b:—S. loc. 52),, ex Type (designated here): H eringeriana - 14(2): 157 191. 2020. 158 Notes on Zygopetalinae. hort. P. Wolter s.n.

, February, (neotype is the and petals light greenBatemannia to brown with lepida no maculae and illustration– later published by Schlechter 1915b: t. a white labellum. Rchb.f. has 5, fig. 7 14!; type B destroyed). been accepted in the taxonomic literature since its Batemannia colleyi peruviana publication, although it had not virtually been used , comb. etsubsp. stat. nov. Batemannia(Mast.) in herbarium specimens. The study of its Meneguzzoperuviana Masters Batemania nomenclatural type B. and colleyi protologue indicates the peruviana Batemannia(1895: peruviana 551), as “ differences of it with could be attributed to ”. Batemania peruviana nom.Rolfe superfl. (1895: the labellum midlobe which is somewhatB. lepida longererein and 193),Zygopetalum as “ peruvianum ”, narrower. Notwithstanding, is h (Mast.)— NicholsonS. loc.(1901:, ex formalizedB. colleyi as a synonym since it neatly fits within hort.747). Type (designatedMessrs. here): F. SanderPERU. and Sons s.n. the morphological range. Batemannia , March 1895, colleyi In the eastern distribution of (lectotypeBatemannia K barcode leferenzii K000589110!). Figure 1B. , in PeruB. and peruviana Bolivia, a biological entity firstly — Senghas (1993: 171). described as ibed as B. leferenziiMast. occurs that was Type: BOLIVIA. La Paz: betweens.d., P. CoroicaLeferenz andsub latersol redescr Senghas, based HeidelbergCaranavi, 50 Botanical km before GardenCaranavi, O -20702 ely on differences in the colouration. It is (holotype characterized by light green sepals and petals with HEID, isotypesyn. Roberto nov. Vásquez Chávez’s private red maculae that are confluent or not, light rose herbarium), labellum with small dark red spots, and midlobe BATEMANNIA COLLEYI whitere to light rose. Since this colour variation is KEY TO THE INFRASPECIES OF adily diagnosable, constant within its populations, i.e., there are no populations with 1. Flowers with sepals and petals light green to intermixed types ofB. flower colleyi colouration between this brown and no maculae, labellumB. colleyi white ……………….colleyi… eastern form and , it is herein proposed to ……………………………….…...….. subsp. be considered an infraBatemanniaspecific taxon colleyi at level of 1’.no Flowers with sepals and petals light green withae, subspecies,peruviana hence subsp. maculae or with confluent dark red macul (Mast.) Meneguzzo.Batemannia peruviana is labellum lateral lobes light rose with small dark red The authorship of spots, midlobe white toB. light colleyi rose ………peruvianum…………….…… herein attributed to Maxwell Tylden Masters (1895)

….………………….……….. subsp. who was editoral The and Gardeners’ writer of Chroniclethe unsigned matters Batemannia colleyi colleyi is an of the journ at that time Lindl. subsp. (Britten 1907). It is worthy noting that NicholsonB. epiphyte species that mainly grows in rainforests peruviana(1901) had to alsoThe Gardeners’ attributed Chronicle the publication of throughout northern South America, in Colombia, publication, - Venezuela, Guyana, Suriname, French Guiana, even though he omitted the authorB. as peruvi he alsoana did for northern and central eastern Brazil, Ecuador, Peru, otherbeen works. Notwithstanding had Bolivia, and on the South American continental shelf misattributed to a publication about three island of Trinidad. Its flower morphology is months latter by Robert Allen Rolfe (1895), which Hrelatively eringeriana homogeneou s -along its range with sepals 14(2): 157 191. 2020. 159

et al. Meneguzzo, T.E.C. turns to be a superfluous name (Turland 2018: destroyed). Rio de Janeiros.d., J. :Barbosa [Casimiro Rodrigues de Abreu,] s.n. Art. 52.1 and 52.2). Barra de São João, bahiensis (syntype J. Barbosa . Rodrigues’s personal herbarium 2. destroyed). Paradisanthus mosenii Paradisanthus bahiensis Paradisanthus paulensisReichenbach (1881a: bahiensis in Rei Reichenbach subsp. 298). nom. superfl. et illeg.Barbosa Paradisanthus Rodrigues bahiensis Hortulanorumchenbach in (1852b: 931). (1882:mosenii 215),var. paulensis non valid. publ. Reichenbach (1854: 30),et non valid. publ.(Barb. Paradisanthus Rodr.) Hoehne geraënsis (1953: al. —Type (designated by Meneguzzo 153),Barb.Rodr., in sched. 2015: 26): BRAZIL.- Bahia: Valença, road J.L. Hage,from et al. Type— (designated by L.Valença Anderson to road & BRM. 101,Hagberg 25 February 1958 1986, Meneguzzo 2015: 27): BRAZIL. São Paulo:C.W.H. (neotype MBM Santos,Mosén 3486 Burutoca River, 10- January 1875, 117077!, isoneotypes CEPEC 38252!, K barcode— (lectotype W R 38119!, isolectotypes K000293788!, s. VIES loc. , ex 1929!). hort., s.d. Original, M.J. Jenisch material: Junior BR 658479!, P - barcode P00447842!,- , Psyn. barcode nov. BRAZIL.sub F. Kramer Bahia: s.n. - P00447843!, Paradisanthus S 7 7768!, neglectus S 6 6246 spirit)

(W R not found). Figure 1C. Schlechteret al. (1918: Paradisanthus bahiensis 34). — Type (designated by Meneguzzos. loc. , ex hort.2015: micranthus comb. et stat. Reichenbach subsp. 27): C.BRAZIL. Grossmann South subern A. region: Malmquist s.n. , ca. nov. Zygopetalum micranthum (Barb.Rodr.) Meneguzzo, 1901, (lectotype Paradisanthus paranaënsis Barbosa Barbosa Rodrigues is the illustration published in the protologue by nom. superfl. et illeg. (1877: 109). Schlechter , syn.1918: nov. 31, t. 3!; type specimen B Paradisanthus micranthus Rodrigues (1882: 215), destroyed) espiritosantensis et al. (Barb. Rodr.) Schlechter . Paradisanthus espiritosantensisRuschi — S. loc., s.d., J. Barbosa Rodrigues (1918: 36). Type (designated by Meneguzzo (1954: 551) s.n. 2015: 27): BRAZIL. (Ruschi) Ruschiet al. (1964: 1). Type— (designated by Iconographie des Orchidées (the lectotype is the original illustration! that Meneguzzo 2015: 27): BRAZIL. Espı́ritoA. du Brésil t was to be published in Santo:Ruschi Santas.n. Teresa, Canaan Valley, 22 April 1951, Instituto . 502, deposited in the library of the (lectotype is the original illustration! de Pesquisas Jardim Botânico do Rio de deposited the library of Instituto Nacional da Mata Janeiro, bound in volume 5, and later published in Atlântica and published in the protologue, Ruschi, the book with same title by Barbosa Rodrigues 1954:syn. nov. 550, t. s.n.!; holotype MBML 1431 destroyed) 1996: 357, t. 232 and 358, t. 233, fig. A!; type J. Paradisanthus mosenii var. virens in — Barbosa Rodrigues personal herbarium destroyed). sched. — Ghillány, s.d., J. Barbosa Rodrigues s.n. Original materials: BRAZIL. Paraná: Curitiba Material: BRAZIL.-not Bahia: Una, Santa Rosa ’ [“Curityba”], (syntype farm, 7 km eastE.B. Santosherneast & E.J. of Judziewicz São José, 4044 27 s.d., J. Barbosa Rodrigues s.n. João Barbosa Rodrigues personal herbarium February 1986, destroyed). Paranaguá, (CEPEC 39443!, K barcode K000293787!, MBM

(syntype J. Barbosa Rodrigues’s personal herbarium 117076!, RB 554744!, VIES 1931!). H eringeriana - 14(2): 157 191. 2020. 160 Notes on Zygopetalinae.

Figure 1. Batemannia and Paradisanthus. Batemannia colleyi colleyi. B.

BatemanniaPictures colleyi of livingperuviana specimens Paradisanthus of bahiensis bahiensisA. Paradisanthus subsp.bahiensis micranthus. subsp. . C. subsp. . D. subsp. Biodiversit A, D. by user ‘Orch’ facilitated by Wikimedia Commons. B. published in Masters (1895) and facilitated by

y Heritage Library. C. by A. Popovkin. PARADISANTHUS BAHIENSIS Paradisanthus

KEY TO THE INFRASPECIES OF Rchb.f. and accepted with reservationsoldest name, a singleP. bahiensis biological entity under the 1. Flowers with sepalsP. and bahiensis petals white tobahiensis light pink P. micranthus Rchb.f., henceP. …………………………....… subsp. synonymizingmosenii Barb.Rodr., P. neglectus(Barb.Rodr.) Schltr., P. 1’. Flowers with sepalsP. bahiensis and petals whitishmicranthus green ….… espiritusantensis Schltr., and …….……………..…….… subsp. (Ruschi) Ruschi. At that time it was et al. clear the existence of two distinct flower In a previous study, Meneguzzo (2015) morphologies which differed only by flower colour, Hrevised eringeriana all published - names for the genus but with a not much known about the limits of the 14(2): 157 191. 2020. 161

Meneguzzo, T.E.C. tyrianthina distribution of each one. Later, Meneguzzo (2018) ReichenbachHortulanorum (1 in852a: 668). non delineated that the variation with white to lightted pink in valid. publ. ZygopetalumReichenbach violaceum (1856b: 187), sepals and petals is geographically distribu violacea(Lindl.) the Brazilian state of Bahia, in the countryside in Reichenbach (1863a:in 650). Chapada Diamantina and in the coast from its (Lindl.) Rollisson —Burbridge (1874: ra, 160). ex hort. Type, northernmost distribution point towards the (designateds.d., R.H. Schomburgk here): GUYANA. sub Messrs. Demera Loddiges s.n. southern municipality of Una.P. bahiensis Therefore, the only namevar possibly linked to it is . The second (neotype is the illustration published- by Lindley iation with sepals and petals green is totally 1839d: t. 26!; original material K L not found). allopatric, native from the coastal Bahia southern of Figure 2A. Una and a few localities in the Brazilian countryside Pescatoria violacea alba towards the southern state of Santa Catarina. All the comb. nov. f.Bollea violacea(Christenson) alba remaining names have their type specimens with Meneguzzo, — f. green sepals and petals and can be linked to this ChristensonS. loc., ex hort. (1992:, s.d., C. 440). Pavillowski Type: s.n.FRENCH GUIANA. morphological type. Hence, the latterParadisanthus is herein - -du- (holotype AMES proposedbahiensis to be a subspecies,micranthus 64900!). Saint LaurentP.A. SagotMaroni 1141 [“Maroni”], ı̂le subsp. (Barb.Rodr.) Portal, October 1857, (paratype P Meneguzzo,Paradisanthus which is based micranthus on the oldest. name of barcode P00612157!). the group, Warrea bahiensis Hort. is a not PESCATORIA VIOLACEA The name KEY TO THE INFRASPECIES OF validly publishedParadisanthus name because bahiensis it is merely citedet al.as a synonym of Paradisanthus(Turland 1. Flowers with sepals and petals purple, callus 2018:paranaënsis Art. 36.1(b)Barb.Rodr. and and 36 P. Ex. paulensis 7). Barb.Rodr. yellowish to purplish with orP. without violacea dark violacea purple keels ………………….…..………….. f. are respectively superfluousZygopetalum and micranthum illegitimate 1’. Flowers with sepals and petalsP. vio white,lacea callusalba Barb.Rodr.homotypic and names P. moseniiof yellowish ………...…………..….……….. f. Paradisanthus moseniiRchb.f. (l.c.:var. Art. paulensis52.1 and Pescatoria violacea 52.2). violacea (Lindl.) Dressler f. (Barb.Rodr.)P. mosenii Hoehne is an illegitimate homotypic is a species widespread in the Guiana name of because it is basedate name on the P. Shield in northern South America. It is characterized homotypicpaulensis superfluous and illegitim by flowers with sepals and petals purple with callusout (l.c.: Art. 6.10, 35.1, 52.2, and 52 Ex. 2). of the same colour or yellowish, with or with 3. Pescatoria violacea dark purple keels. It is presumable that sporadically within populations with typical purple flowers Pescatoria violacea violacea in specimens with white flowers and yellowish callus et al. (Lindl.)Huntleya Dressler violacea f. appear,named Bollea therefore violacea categorized alba at form rank and Whitten (2005: 95).Bollea violacea Lindley enus Bollea f. Christenson. Since the (1839b: misc. 19). (Lindl.) g Rchb.f. was synonymized under H eringeriana - 14(2): 157 191. 2020. 162 Notes on Zygopetalinae.

Pescatoria in et al.

Rchb.f. (Dressler Whitten 2005), (Rchb.f.)—S. Nicholson loc., ex hort. (1901:, s.d., s. 747). leg. s.n. Type (designated-R and this infraspecies therefore lacks a combination here): —S. (lectotypeloc., ex hort. W , innamed the latterPescatoria genus, violacea it is herein alba transferred and 30809!).Messrs. H. Low Remaining & Co. s.n. syntype: -R not f. (Christenson) syn. nov. (remaining syntype W Meneguzzo. - found), Promenaea dusenii At Herbarium Lindley there is a sheet (K L Schlechter— (1921: 476). Huntleyabarcode K000718360!)violacea identified by Lindley-annotated as TypeMorretes, (designated d here): BRAZIL. Paraná: composed of one non eviationP.K.H. Ipiranga Dusén 13872[“desvio Ypiranga”], 9- flowering specimen, and a reproduction of the February 1912, syn. nov. (lectotype S R illustration published of the same species (Lindley 5276!; typePromenaea B destroyed), malmquistiana 1839d: t. 26). There is no internal or external — Schlechter evidence that would allow recognizing this (1921: 475). Type (designated here): BRAZIL. Rio specimen as a type, so it remains a specimen with de Janeiro:M. Bocayuva Nova Iguaçu, 150 pipeline Orbel II, 11 July unidentified origin, and the designated neotype is 2005, —S. loc.(neotype, ex hort. RB, s.d. 464, s. leg. spirit!). s.n. the illustration of the specimen used in the Original material: syn. nov. description, but not publishedHuntleya tyrianth in the inaprotologue Hort. is (type B Promenaeadestroyed), stapelioides var. macrantha (Lindley 1839d: t. 26). — not validly publishedBollea violacea name as it was merely cited a Hoehne (1952: 126). Type: BRAZIL. Rio de etsynonym al. of (Lindl.) Rchb.f. (Turland Janeiro:C. Spann half wayagel on113 Petrópolis Range, December. 2018: Art. 36.1(b) and 36 Ex. 7). 1927, (holotype SP 25447!) Promenaea stapelioides Promenaea stapelioides stapelioides

4. viridiflora subsp. f. Promenaea stapelioides comb. et stat.(F.Barros nov. Promenaea & Barberena) viridiflora Meneguzzo, Barros & stapelioides stapelioides(Link in & Otto) Lindl. subsp. Barberena in Cymbidiumf. stapelioides Lindley (1843: misc. —BR Barberena & Barros (2015: 206). 13). Maxillaria stapelioides Link & Otto (1821: Type: AZIL. São ex Paulo: hort. Cananéia, Ilha do 111). Peristeria stapelioides(Link & Otto) Lindley Cardoso State Park, F. ,Barros 1988, s.n. flowered sic recte in (1832: 146).in (LinkZygopetalum & Otto) F.cultivation Barros 3248 10 January 2014, [ ] [ Loudonstapelioides Baxter (1850: 604). ] (holotype SP 465171!). Santo André, (Link & Otto) Reichenbach (1863a: Altoex hort. da Serra de Paranapiacaba Biological Reserve, 658). — Type (designated by Christenson 2013:ex , 23 JulyF. 1982,Barros flowered s.n. sic in recte cultivation F. Barros 10 hort.793):, s.d.BRAZIL., H.K. RioBeyrich de Janeiro: s.n. Rio de Janeiro, January3249 2014, [ ] [ (lectotype is the ] (paratype SP 465170!). illustration published in the protologue, Li nk & Otto Promenaea stapelioides nigricans 1821: t.Promenaea 52!; type B destroyed).stapelioidesFigure var. 2B.heteroptera nothosubsp. comb. et in (Königerstat. nov. & Promenaea J.G.Weinm.bis) × Meneguzzo,nigricans ReichenbachZygopetalum N.E. stapelioides Brown & Reichenbach var. heteropterum (1883: inmann in Königer— & 70). We Königer (1995: 113). Type: BRAZIL. H eringeriana - 14(2): 157 191. 2020. 163

s. loc., ex hort., s.d.Meneguzzo,, J.G. T.E.C.— Rio d

ParanáWeinmann or & São E.B. Paulo: Pfeifer sub W. Königer WK-55 here): BRAZIL. e Janeiro: Mangaratiba,E. Saddi 66 margin of Grande River, 20 April 2003, —S. loc., (holotype M not found, isotypes HB not found, K not (neotypeex hort., s.d. RB, s. 462 leg. spirit!). s.n. Original material:syn. nov. found, Willibald Königer's private herbarium—Promenaea not Promenaea albescens(not found), seen).stapelioides Parent taxa (postulated here): stapelioides Promenaea paranaënsis var.Schlechter albescens (1919: stapelioides(Link. × &Promenaea Otto) Lindl. stapelioides subsp. 327). (Schltr.) xanthinumf. . subsp. Hoehneet al. (1953: 81).— TypeS. loc. (designated, ex hort., s.d. by, Barberenas. leg. s.n. Promenaea(Lindl.) Meneguzzoovatiloba robertii 2016: 326): —nothovar. (neotypesic recte is the illustration published by Mansfeld Menezes s. loc. (1995:, ex hort. 12). Type: BRAZIL.P.H.S. Robert Espı́rito s.n. [ ] [ —Schlechter]S. loc., ex hor 1930:t., s.d. t., Grossmann50, n. 197!). s.n. Original Santo:sic recte L. Robert, Januarys.n. 1995, material: (type [sic] [recte ] (holotype MBM 170073 B destroyed).Promenaea acuminata [— ] [ MBM 170894]!; epitype designated here —Schlechter (1919: the original picture published in the protologue,ed 327). Type (designated here): BRAZIL. Paraná:G. Menezes—Promenaea 1995: 13, t.stapelioides s.n.!). Parent taxa (postulat Morretes,Hatschbach Floresta & C. Koczicki Colony, 20 249 07 January 1969, here): stapelioides stapelioides(Link. × & Otto)Promenaea Lindl. — (neotype MBM subsp.stapelioides xanthinumf. 12694!).s. loc. Original, ex hort. material: BRAZIL. GrossmannSouthern syn. nov. subsp. (Lindl.) Meneguzzo, region,s.n. , Novembersyn. nov. 1918, (typePromenaea B destroyed), catharinensis Promenaea stapelioides rollissonii —Schlechter (1921: comb. et stat.subsp. nov. Maxillaria rollissonii(Lindl.) 479). Type (designated here): sicBRAZIL.recte Paraná: Meneguzzo, Promenaea rollissonii São José dos Pinhais, Garuva [ ]G. [ HatschbachSanta Lindley (1837: sub t. 1986). Zygopetalum Catarina:1849 Garuva], 6 January 1950, — (Lindl.)rollissonii Lindley (1843: misc. 13). (neotype MBM 50058!).s. loc. Original, ex hort. material:, s.d., von (Lindl.) Reichenbachet (1863a:al. 659). Type— BRAZIL.Fürstenberg Santa s.n. Catarina: syn. nov. (designatedS. loc. by, ex Barberena hort., s.d. , Messrs.2016: Rollisso 326):n s.n. Promenaea(type paulensis B destroyed), BRAZIL. Schlechter (1922b:— (neotype is the published illustration- by Lindley 63). Type (designated by Christenson 1996: 22): 1838: t. 40!; original material K L not found). Figure BRAZIL. São Paulo: Iguape, PedrasA.C. Peak,Brade Iguape7758 2C. Maxillaria lentiginosa River Valley, January 1918, Promenaea lentiginosa Lindley (1839c: misc. (lectotype US 1208125!, isolectotypes AMES 61). Zygopetalum (Lindl.)lentiginosum Lindley (1843: 30646!, HB 8540!, R 29940!, SP 8255!; type B misc. 13). (Lindl.) destroyed). Promenaea riograndensis Reichenbach— (1863a:S. loc., ex 659). hort. , s.d. Type, Messrs. (designated Loddiges Schlechteret al. (1925: here):s.n. BRAZIL.- 87). Type— (designated by Barberena 2016: (lectotypePromenaea K L barcode rollissonii K000857188!). var. obtusa in 326): BRAZIL. SantaL.C. Catarina: Oliveira Araranguá, s.n. Veado et al. Regel River, 14 April 2011, — (neotype CRI von Kuester (1856: 23). Type (designated 8945!). Original material: BRAZIL. Rio Grande do H eringeriana - 14(2): 157 191. 2020. 164 Notes on Zygopetalinae.

L. Burger sub F. Aquino Zygopetalum micropterum

33Sul: Torres, February 1922, (1881b: 134). Zygopetalum (type B destroyed). Reichenbachmicropterum (1881b: 134). Promenaea stapelioides xanthina Bois in (Rchb.f.) Bentham &nom. Hooker illeg. filius ex comb. et stat.subsp. nov. Maxillaria xanthina(Lindl.) Bois (1893:—S. loc. 133),, ex hort. Type Meneguzzo, Zygopetalum xanthinum d'Haede(designated sub here):H. Veitch s.n. , July- 1881, Lindley (1839a: sub t. 17). (lectotype W R 40606— (Lindl.) Reichenbach— (1863a: 659). Type pencilS. loc., iconex hort. on left!)., s.d. ,Remaining B.S. Williams original s.n. material: (designated here): BRAZIL.G. Gardner Rio de652 Janeiro: Organ- -R syn. nov. (remaining Mountains, May 1837, (lectotype K syntypePromenaea W not found), citrina in L barcode K000857191!, isolectotypes BM barcode nom. illeg Bull ex Regel Regel— BM000533456!, BM barcode BM001122605!, G (1887b: 694), . Type (designated here): barcode G00359929!, P barcode P00447850!,— SP BRAZIL. Rio deD. Janeiro: Miller s.n.Nova Friburgo, Sitio Baccus, 114332!, OXF s.n.!). Remaining syntype: BRAZIL. October 1998, —S. loc., ex(neotype hort., s.d. RB, s. 729 leg. spirit!).s.n. RioJune, de J.T. Janeiro: Descourtilz Angra s.n. dos Reis, Island Ilha Grande, Original material: (LE (remaining syntype is the not found).Zygopetalum ovatilobum original illustration thatin was Epidendres to be published des Forestes by Jean Promenaea ovatiloba Klinge (1898: 144). ThéodoreVierges du DescourtilzBrésil (Klinge)— Cogniaux S. (1906a:loc., ex 3 (16): t. 63!, deposited at the 468).hort. Type (designated here):Lietze s.n.BRAZIL. library ofMaxillaria l’Institute decitrina France). Figure 2D. , 25 April 1897, syn. nov. (lectotype LE Promenaea citrina Lyons (1845: 76). barcode Zygopetalum LE00001454!), xanthinum var. major Masters Zygopetalum citrinum(J.Lyons) Don (1845: 720). Promenaea xanthina var. major (J.Lyons)— Nicholson (1887: (1901: 106). (Mast.)— 245). Type (designated here): BRAZIL. São Paulo:G. Cogniaux (1906a: 465). Type (designated here): Martinelli,São Luiz T. do Barbará, Paraitinga, G.M. Souza 13 October & R. Azoury 1999, 15930 BRAZIL. São Paulo: Piquete, AÁrea de Proteção —S. loc., ex AmbientalL.N. Gonçalves Serra da & Mantiqueira, P. Duffles 293 28 December (neotypehort., s.d., RBs. leg. 429828!). s.n. Original material: 2013, —S. loc., ex(neotype hort. RB Maxillaria (notguttata found). 597001!).H.A. Tracy Originalsub I. Bradt material: s.n. , 1901, nom. nud. PromenaeaReichenbach guttata (1852b: Promenaea fuerstenbergiana(not found). 672), Zygopetalum guttatumReichenbach Schlechter— (1856a: 323). (Rchb.f.) (1921: 481). Type (designated here): BRAZIL. Reichenbach—S. loc. , ex (1863a: hort., s.d. 659)., s. leg. Types.n. (designated-R Santa Catarina:A.C. Cervi Camburiú, 7008 Guarita Peak, 9 March here): (lectotype W 2000, (neotype MBM 256016!,—S. 40602 specimen by— S. side loc. of, ex red hort. label!)., s.d. , Remainings. leg. s.n. isoneotypeloc., ex hort. UPCB, s.d ., 42082!).von Fürstenberg Original material:s.n. original material: - S. syn. nov. (type B (remainingloc., ex hort. syntype, s.d., s. W leg.R 40602 s.n. two envelopes!). destroyed), Promenaea paranaënsis - (remainingsyn. nov. original —Schlechter (1921: material Promenaea W R 40601 twomicroptera icons!), 477).Morret Type (designated here): BRAZIL. Paraná: Reichenbach es, Parque Estadual do Marumbi, 6 February H eringeriana - 14(2): 157 191. 2020. 165

C. Giongo 101 Meneguzzo, T.E.C. syn. nov.

1999, Pico Facãozinho, — (neotype UPCB SP 247598!),Promenaea chautinana in 47310!). Originex alhort. material: –idem, [“Fazenda sched. —S. loc., ex hort.Reichenbach, A. Morumby”],P.K.H. Dusén 1581 , October Novembersyn. nov. 1914, ChautinMaterial: s.n. - , June 1888, Promenaea(type sincorana B destroyed), o Neto & Promenaea(W R 40607!). xanthina var. minor in —Castr sched. — Hoehne, Campacci (1993:sic recte 10). Type: BRAZIL. Bahia:ex Material: BRAZIL. A.C. Rio Brade de 9465 Janeiro: hortItacoatiara. [ ] [ Ibicoara],V.P. SincoráCastro Range,Neto s.n. Teresópolis, December 1929, (R , 15 November 1990,syn. nov. 32293!). Promenaea xanthina concolor Pabst, in (holotypePromenaea SP 333604!), silvana sched. — f. — Barros & CatharinoS. loc., Material: BRAZIL. Espı́rito Santo:R. Domingos Kautsky ex(1995: hort. 94). Type: BRAZIL. SoutheE.P. Silvarn s.n.Bahia: 310Martins, Azul Peak, 3 December 1970, , 20 November 1980, (holotype (HB 57258!).

Figure 2. Pescatoria and Promenaea Pescatoria violacea violacea. B. Promenaea stapelioidesPictures stapelioides of living specimens stapelioides of Promenaea stapelioides . A. rollissonii Promenaeaf. stapelioides xanthina. –subsp. f. . C. subsp. . D. subsp. A C. by K. Senghas and facilitated by Swiss Orchid Founda tion at Herbarium Jany Renz. D. by R. Jenny and facilitatedH eringeriana by Swiss Orchid -Foundation at Herbarium Jany Renz. 14(2): 157 191. 2020. 166 PROMENAEA STAPELIOIDES

KEY TO THE INFRASPECIES OF specimens cultivated in Europe. The authors that ... 2 published on this genus (Rolfe 1905, Cogniaux 1. Labellum callus with distal portion rounded... … 4 1906a, Hoehne 1953, Pabst & Dungs 1977) more or 1’. Labellum callus with distal portion obtuse …… less accepted nearly all published species at the 2. Sepals, petals, and labellum light green with dark..... time, probably due to the lack of access to brownP. stapeliodes maculae, labellum stapelioides more densely stapelioides maculate representative samples of herbarium collections, …. subsp. f. difficulties in understanding species delimitation, 2’. Sepals, petals and labellum dull coloured (no3 and also almost certainly by not having studied the maculae) …………………………….………………………….… type materials. It led to a substantial taxonomic 3. Sepals and petals dull dark crimson to reddish inflation which does not reflect this genus in the purple, labellum dark purple with the proximal natural world by the use of tenuous morphological portion of a darkerP. stapelioides hue than the distal portionnigricans …….… characteristics which were commonly …………………. nothosubsp. superimposed on the formerly accepted species 3’. Sepals and petals dull light green, labellum light circumscriptions. greenP. with stapeliodes distal portion ofstapelioides midlobe white viridiflora ………….… The study of all type specimens …..... subsp. f. accompanieds, as by their original illustrations and 4. Sepals, petals, and labellum light green with light protologue well as of a good amount of brown spots; labellum callus lateral- margin.. herbarium specimens and observations of living irregularly denticulate P. stapelioides and distal margin 2rollissoniilobed … specimens, led to the recognition that in fact there ………………………….. subsp. are constant morphological differences that 4’. Sepals, petals, and labellum light to dark yellow separate the genus into a small number of and dull coloured, or with few and sparse crimsonin to recognizable biological entities. Although, when spots on proximal portion, or rarely with th those biologicalZygopetalinae entities are accessed within the thick coloured radial bars on proximal portion;- scope of the , it is clear that their labellum callus marginP. stapelioides entire, distal marginxanthina 2 or 3 characterization are very subtle in a way that do not lobed …………...……... subsp. merit the recognition as distinct species. The differences between them are comprised of the variations in flower colour and subtle variations in Promenaea labellum callus. Since there are no populations The genus had been comprised intermixed, except by a case latter discussed, they ofBrazil 16 specieserts widespread et al. in eastern and southern are classified as subspeciesPromenaea under stapelioides oldest described (Gova 2020) which grow in shady name for the genus, (Link & areas under the forest canopy and seem to prefer Otto) Lindl. A significant quantity of typifications moister niches. By the end of the 20th century less are proposed to ascertain the placement of than half of the 27 names had been described. Most synonyms and to fulfil the requirement of a type of them were described over a short period by material for each published name. Schlechter (1919, 1921, 1922b, 1925) based on wild -

* Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Rua Pacheco Leão 915, Jardim Botânico, 22460 030, Rio de Janeiro, Rio- de Janeiro, Brazil; Universidade de Brası́lia, Instituto de Ciências Biológicas, Departame-nto de- Botânica,- caixa postal 4457, 70919 970, Brası́lia, Distrito Federal, Brazil. [email protected]. https://orcid.org/0000 0003 4760 9588.

167

Promenaea stapelioides Meneguzzo, T.E.C. n P.

stapelioides (Link & Otto) Lindl. sporadicstapelioides colour shift stapelioides in some specimens withi subsp. is characterized by sepals and subsp. , which is most petals light green with dark brown maculae on probably less frequent, as commonly happen in which the labellum is more densely maculate, and orchids. Thus it is accordinglyPromenaea reduced stapelioides to the the distal margin of the- labellum callus distal infraspecificstapelioides rank of form:viridiflora portion is rounded and P. 2 duseniilobed. and The P study. stapeliodes of the subsp. f. (F.Barros & var.type heteropteraand protologues of Barberena) Promenaea Meneguzzo. stapelioides rollissonii P.let stapeli to theoides conclusion stapelioides they are subsp. d as a conspecific with subsp. (Lindl.) Meneguzzo is herein propose since their colour variation and labellum callus subspecies characterized by flowers with sepals, shape neatly fit within the morphological range of petals and lip light green with light brown maculae, the subspecies.Promenaea viridiflora Barros & Barberena labellumirre callus distal portion obtuse, lateral margin- F. lobed.gularly Barberena denticulate, et al. and terminal portion 2 was describedP. stapelioide as as distinct species which differed P. lentiginosa(2016) proposedP. al besce the n s from only by the dull green flower synonymizationP. paulensis of P. riograndensisLindl., colour and by what is tentatively interpreted from Schltr., Schltr., and Schltr, the protologue as an erect labellumberena callus & Barros with a which are here scrutinized andP. rollissonii accepted asvar. good. obtusa To conic trunk like protuberance (Bar that listP. the acumi newnata synonyms P. catharinensis 2015). The last morphologic feature is not Regel, Schltr., and et al. diagnostic because it is not exclusive to the taxon, Schltr., which were overlooked in by Barbarena but neatly falls within theP. morphologicalstapelioides range (2016) have also been added. The designation of stapelioidescommonly found in subsp. theirt neotypes in accordance to the description in labellum . Moreover, subtle differences in he protologue and original type localities well Zygopetalinaemorphology do not differentiate characterize them to reinforce the proposed species (Meneguzzo 2018). Flower synonymy. Promenaea stapelioides xanthina colour is not a good distinctive feature at species subsp. level since P. it stapelioides clearly has the same light green (Lindl.) Meneguzzo is the only yellow flowered perianth of but with a lack of spots. subspecies of the genus. It is characterized by Contrary to what was mistakenly cited in the sepals, petals, and labellum light to dark yellow and protologue, the holotype and paratype bear dull coloured, or with few sparse crimson spots on collection numbers of the senior author which are the proximal portion, or rarely thin to thick coloured herein rectified. Both specimens were prepared radial bars on the proximal portion. The labellum from cultivated specimens collected over 30 yearstions callus- margin is entire, and the distal margin is 2 orP. ago; the authors did not access their popula 3ovatilobalobed. Specimens commonlyP. guttata identified as andPromenaea the provided viridiflora data is of a speculative nature. (Klinge) Cogn. and Rchb.f. have occursP. stapelioideswithin the geographic somewhat narrower petals and sepals and distribution and niche ofP. viridiflora . All these commonly have thick coloured radial bars on its evidences indicate that is merely a proximal portion (including personal field H eringeriana - 14(2): 157 191. 2020. 168 Notes on Zygopetalinae.

Promenaea xanthina var. major observations in IlhaPromenaea Grande andsilvana surro unding areas, 41.3). Similarly “ Rio de Janeiro). was described Cogn.” had been mistakenly considered a new for a group of specimens from southern Bahia with variety by Cogniaux (1906a) in the botanical golden yellow flowers (orange in the protologue) indexes.Zygopetalum The xanthinu authorm var. explicitly major indicated and petals and sepals sometimes tending to be more b“ Hort.” as acute. The complicated matter of accepting the last asionym along with its full reference. The name as a good and separate species is that plants authorship of the basionym itself is hereby with the typical yellow colouration are found identified as Maxwell Tylden Masters (1901), who amongst them in natural populations (including was editor and writer of the unsigned matters of orethe personal field observations in Arataca Range and Promenaeajournal at thatcitrina time (Britten 1907). Theref surrounding areas, Bahia), as well as some with Maxillaria citrina (J.Lyons) P.N.DonPromenaea is basedxanthina on narrower or more acute - sepalsorrelated and petals;and not thus an var. major J.Lyons, and Zygopetalum these characters are non c xanthinum var.(Mast.) major Cogn. Mast. is based on exclusive but rather common feature which Maxilaria guttata comprisereasons P. the ovatiloba variability, P. ofguttata this subspecies., and P. si lvanFor athese are Reichenbach (1852b: 672) is a denomination or naked name due toet theal. hereinP. citrinasynonymized . The designationP. citrina of a neotype lack of a diagnosis or description (Turland for P. xanthina(J.Lyons) var. P.N.Don,major Bull exP. 2018: Art. 6.3, 38 Ex. 1, and Rec. 50B.1); therefore,Promenaea it Regel,fuerstenbergiana P. paranaënsis(Mast.) Cogn., mustguttata not be cited as the basionym of Schltr., and Schltr. Reichenbach (1856a: 323) as frequently along with the study sof under their protologuesP. stapelioides led to the found in botanical. Promenaea literature; citrina the later name is the xanthinanew synonymization subsp. basionym itself P. citrinaBull ex Regel is P. microptera. In the same way P.the sincorana study of the types of an illegitimate later homonym of (J.Lyons) Rchb.f. and V.P.Castro & P.N.Don (l.c.: Art. 52.1 and 52.2).in Campacci led to the conclusion they do not differ KönigerPromenaea & Weinmann nigricans Königer (1995) from this species and are also synonymized. Promenaea published as a new species citrina It has been found that the name “ from a specimen obtained from cultivation from the P.N.Don”, as cited in the nomenclatural state of Paraná or São Paulo. It is unique in the genus indexes, had been incorrectly treated as a new due to its immaculate reddish purplend flowers labellum on species attributed to Patrick Neill Don (1845). which the lateral sepals are darker a However,Maxillaria this citrina name is hereby traced to its basionym, dark purple with the proximal portion of a darker J.Lyons by John Charles LyonsP. hue than the distal portion. MenezesP. ovatiloba (1995) var. (1845).citrina Don (1845) himself clearly indicatedM. citrina that . robertiireported in the protologue of is a combination based on that only a single specimen of it was foundP. Although, he did not cite the reference and did this amongststapelioides many wildxanthina collected specimensP. ovatiloba of for other names throughout his publication, whichons subsp. (cited as was not required for validly publishedet combinatial. (Klinge) Cogn.) from an unreported provenance in publications at that time (Turland 2018: Art. the state of Espı́rito Santo. It was intentionally H eringeriana - 14(2): 157 191. 2020. 169

Meneguzzo, T.E.C. et al. described as a mere flower colour variation current Code (Turland 2018). Lindley (1839c) characterized by dull dark crimson flowers with the didMaxillaria not designate lentiginosa the holotype in the protologue of labellum callus of a darker hue. Lindl., hence the lectotype is The studyP of. nigricans the original and materials P. ovatiloba led to var.the herein designated since the protologue lacked the robertiiconclusion that indication of a unique specimenet inal. a specific are conspecific, and completely spotless as collection (McNeill 2014, Turland 2018: Art.e confirmedP. nigricans by the and iconography P. ovatiloba va in r. the robertii protologue. are not 9.1).Promenaea The set of albescens illustrations from which the neotyp Both P. stapelioides of Schltr. was designated was xanthinastrictly conspecificdue to with subsp. posthumously published by Schlechter (1930), not the distal margin of the labellum Mansfeld who edited the publication. callus having rounded instead ofP. obtusestapelioides lobes. They colorata stapelioidesare not strictly conspecificstapelioides with subsp. 5. f. - either, because of the Warreopsis colorata darker, and sparsely spotted flowers instead of light colorata in (Linden & Rchb.f.)Zygopetalu Garaym green and densely spotted, although the obtuse coloratumsubsp. Garay (1973a: 51).in distal margin of the labellum callus is a shared Linden & Reichenbach —Reichenbach diagnostic character. Empirical observations on the (1863a: 662). Type (designated here):s. loc.COLOMBIAL.-J. hybrid crossings by horticulturists and widely Schlim[“New Granada”].51 Eastern- region: , July, presented in the horticulture literature and in . (lectotype W R 40578 only left icon!). orchid nurseries show that whenever the last Figure 3AZygopetalum pardinum species is crossed with a yellowish specimen thatP. Warreopsis pardina Reichenbach bearstapelioides spotless flowersxanthina or with some spots (like (1863a: 662). (Rchb.f.) Garayet al. subsp. or a yellowish hybrid), (1973a: 51).— Type (designated by Meneguzzo the results are siblings with flowers of the same 2015: 30):s.d., W.ECUADOR. Jameson AndesJamieson forests, s.n. western colour pallet of the spots. DueP. stapelioides to that, the two declivity:- [“ - ”] (lectotype postulatedstapelioides parentalstapelioides species and ( P. stapelioides subsp. K L 364389!, isolectotype W R 40577- fragment!); xanthina f. - subsp. idem (remaining, syn. nov. original material W R 40577 right ) mostly co occurs in its marginal icon!) hirtzii in distribution areas and the empirical observationsP. C.H. Dodson —C.H. Dodson onnigricans hybridization, it is herein proposed to combine et & P.M. Dodson (1984: t. 976). Type: ECUADOR. al. the status of nothosubspecies (Turland Pichincha: old road betweenA. Hirtz San & J. JuanLeon and201 2018: Art.P. 50.1 ovatilba and H.10var. robertii Note 1), and thereafter Chiribonga, 7 March 1982, , syn. nov. synonymize under it. For the (holotype SEL 44290!, SEL spririt s.n.!) last name, the original picture published in the Warreopsis colorata parviflora protologue is selected as the epitype since the comb.subsp. et stat. nov. holotype Barberena lacks the labellumet al. (l.c.: Art. 9.9). (L.O.Williams)Zygopetalum parviflorumMeneguzzo, in (2016) made some L.O. WarreopsisWilliams parvifloraWoodson nomenclatural notes that must be revised under the Junior & Schery (1941: 424). H eringeriana - 14(2): 157 191. 2020. 170 Notes on Zygopetalinae.

—

(L.O.Williams) Garay (1973a:– 51). Type: PANAMA.R.E. geographical range, hence expended as Chiquirı́:Woodson Junior Bajo & Chorro, R.W. Schery 20 22605 July 1940, aforementioned. Warreopsis parviflora (holotype AMES (L.O.Williams)ater Garay also 90660!, isotypes F 93942!, MO 102762!). was initially described for Panama,W. colorata but l Warreopsis colorata purpurea found in Costa Rica. It differs from by the comb. etsubsp. stat. nov. Warreopsis(P.Ortiz) dull brown sepals and petals and magenta labellum. Meneguzzo,purpurea — Since the differences are solely in the flower colours Ortiz Valdivieso (1994: 19). Type: P. and both biological entities are disjunctW. parviflora and clearly be COLOMBIA.Ortiz V. 927 Putumayo: Sibundoy, April 1977,. distinguishable, it is proposed thatWarreopsis colorata (holotype HPUJ 10617!). Figure 3B reducedparviflora to subspecific level: subsp. (L.O.Williams) Meneguzzo. WARREOPSIS COLORATA WarreopsisIn the southernmost purpurea areas of the northern s or KEY TO THE INFRASPECIES OF Andes, occurs; it differs from .. the former species by the dull purple flower 1. Labellum midlobe widelyW. colorata obovate, apexpurpurea obtuse … sepals and petals with darker maculae and labellum obe ………………llum………...…… midlobe transversesubsp.- midlobe widely obovate with the apex obtuse, transverse- 1’. Labe oblong, apex2 whilst the others subspecies have the midl Warreopsis purpurea rounded ……………………………….………………….…….. oblong with a rounded apex, and 2. Sepals and petals yellow to green with brown different colours. inhabits maculae, labellum dull whiteW. colorata …………………………... colorata…… western Colombia, northern Ecuador, and western ……….………………….…….. subsp. Bolivia, but has not so far been collected in Peru. It 2’. Sepals and petals dullW. colorata brown, labellum parviflora magenta is reduced to infraspecific level because it is … …………….………..……. subsp. allopatric, and although its morphological Warreopsis colorata differences are readily identified, they are not Warreopsis colorata (Lind. & Rchb.f.) Garay strong enough to characterize it as a distinct colorata purpurea subsp. inhabits the northern part of the species. Therefore it is recognized as Warreella Andes in the Ecuadorian, Colombian, and subsp. (P.Ortiz) Meneguzzo. In W. Venezuelan cloud forests. Its flowers are yellow to this way the four formerly accepted colorata. green with brown maculae and labellum W.dull pardina white. species are accepted as three subspecies of

Senghas & Gerlach (1993) considered (Rchb.f.) Garay as a good Otostylis specie shirtzii endemic to

Ecuador and synonymizedW. colorata Dodson 6. Zygopetalum crinitum crinitum under it, whilst was considered endemic in Zygopetalum mackaii to Venezuela and perhaps ColoW.mbia. colorata The, W. Loddiges subsp. var. crinitum pardinaexamination, and ofO. thehirtzii type specimens of Loddiges (1831: t. 1687). Zygopetalum mackaii var. convexum let to the conclusion that they (Lodd.) Lindley (1833: 187). nom. superfl. et illeg. Eulophia crinita comprise a single biological entity that do not differ Mutel (1842: 9), in the flower shape, size, and colours along all of its (Lodd.) H eringeriana - 14(2): 157 191. 2020. 171

in ZygopetalumMeneguzzo, coeruleum T.E.C. Rollisson in microtusLoudon Steudel (1840:in 605). (B.S.Williams) — Burbridge nonHoffmannsegg valid. publ. Reichenbach (1863a:— (1874: 160). Type (designated here): BRAZIL: Rio 661), TypeS. loc. , (designatedex hort., s.d. , here):F. Warre de Janeiro: Nova Friburgo,A. Bonnet Parqu & E.e CaglioniEstadual 12001 de Três78 BRAZIL.sub C.L. Rio Loddiges de Janeiro: s.n. - Picos, 25 July 2011, —S. loc., ex . (lectotype K L barcode (neotypehort., s.d., RBs. leg. 530496!). s.n. Original syn.material: nov. K000458550!). ZygopetalumFigure stenochilum 3C Zygopetalum(not crinitum found), var. rubellum in non valid. publ. ZygopetalumLoddiges stenochilum (1833: t. Regel 1923), in von Kuester— & Regel (1855: 23). Type (designated LoddigesEulophia stenochila ex Drapiez Drapiez (1834: s.n.). here): BRAZIL. Rio de Janeiro: Nova Friburgo, (Lodd. ex Drapiez)— SteudelS. Reserva Ecológica Municipal de Macaé deC. Cima,Baez, loc.(1840:, ex hort. 605). Type F. (designated Warre s.n. here): BRAZIL. trilhaP. Rosa, para D. PicoMaurenza, da Bicuda, L. Moraes 11 August & R. 2016, Overney 793 , 1828, (the lectotype is the original illustration! deposited at the library of (neotype RB 660907!,—S. loc. isoneotype, ex hort., HUEFSs.d., s. leg. 231950!). s.n. Natural History Museum, London, and published by Originals material:yn. nov. (not Loddiges Zygo 1833:petalum t. 1923! mackaii). var. nanum Harrison found), Zygopetalum crepeauxii mackayi — —Carrière Rio (1887: de (1835:S. loc., ex72), hort. as ,“ s.d., Cooper”. Type s.n. (designated here): 420). Type (designated here): BRAZIL. (the lectotype is the Janeiro: Nova Friburgo, Reserva EcológicaM. illustration published in the protologue, Harrison MunicipalLeitman et deal. M267acaé de Cima, 26 August 1987, 1835:Zygopetalum between crinitum page 48 andsyn. 49nov. mistakenly named as — (neotypeS. loc., ex RBhort. 295035!)., s.d., P. Binot Original sub Zygopetalum ), pubescens Crepeauxmaterial: s.n.BRAZIL. syn. nov. —HoffmannseggRio Zygopetalum(not found), mackaii var. macranthum (1843: 63). Type (designated here): BRAZIL. Rollisson in nom. nud. deM. LeitmanJaneiro: Macaé,et al. 20 Pico do Frade, 22 October 1985, Zygopetalum Burbridgemackaii var. (1874: major 160), Rollisson in — (neotype RB 241568!).S. loc., exOriginal hort., nom. nud. Zygopetalum s.d.,material: s. leg. s.n.BRAZIL. Rio de Janeiro:syn. nov. Burbridgemacranthum (1874: Hortulanorum 160), , in sched. —S. Zygopetalum(type B destroyed),velutinum loc., ex hort., s.d., s. leg. s.n. Material: Zygopetalum mackaii var.Hoffmannsegg velutinum Zygopetalum mackaii(BR var. s.n.!). roseum Rollisson (1843: 62). Rollisson in in nom. nud. Zygopetalum (Hoffmanns.) —Burbridge (1874: 160). crinitumBurbridge var. (1874:roseum 160), in Type (designated here): BRAZIL. Rio de Janeiro: Rollisson ex — Stein Stein Nova Friburgo, Reserva Ecológica MunicipalA.L.V. Toscano de (1892: 599). Type (designated here): BRAZIL. Sã o deMacaé Brito de et Cima, al. 664 28 November 1990, Paulo: SerraA.C. Brade da Bocaina, 20971 Mata da Garrafa, 14 May — (neotype RB 294440!).S. loc., exOriginal hort., 1951, —S. loc., ex(neotype hort., s.d., s. RB leg. 74174!). s.n. s.d.material:, s. leg. s.n.BRAZIL. Rio de Janeiro:syn. nov. Originalsyn. material: nov. (not Zygopetalum(type B destroyed),crinitum var. coeruleum found), Zygopetalum ghillanyi Zygopetalum mackaii B.S.var. — Pabst (1976:in, ex hort. 86)., Williams (1852: 98). Type: BRAZIL. São Paulo: Serra da Boca H eringeriana - 14(2): 157 191. 2020. 172 Notes on Zygopetalinae.

sic recte A.

Ghillány1 November sub 1974R. Windish [ ] [ s.n. 3 November 1975], from southern state Bahia to Rio Grande do Sul, syn. nov. (holotype HB 63061!), Brazil and northeastern . Its habit is Zygopetalum reginae epiphytic and the populations are not aggregated as — Pabst (1976: 87).ex commonly occurs in other species the genus, hence Type:hort. BRAZIL. São Paulo:R. Windish Serra s.n. da sic Bocaina,recte A. fewunique specimens in t are seen during fieldwork. It is Ghillány, 1 November sub R. Windish 1974, s.n. [ ] [ he genus and easily distinguished by the — ] (holotype HB 63017!). labellum with hirsute veins and callus with the Original material:ex BRAZIL.hort. São Paulo:A. Ghillány Santo distal portion deeply fissured. The Z. revision crinitum of and the Antônios.n. do Pinhal, ,syn. 5 April nov. 1975, typeZ. maculatum specimens and protologues of (paratypeZygopetalum HB 63171!), crinitum var. triumphans (Kunth) Garay, the latter as discussed , in sched. —S. loc., ex hort., s.d., s. below, led to the recognition of the herein proposed L.Lindenleg. s.n. Material: new synonyms. Most of the synonyms comprise (BRZygopetalum s.n.!). panduratum , in minor variations in flower colour and labellum sched. —S. loc., ex hort., s.d.Reichenbach, s. leg. s.n. -R veiningstatus. which does not merit distinct taxonomic Material: (W Zygopetalum pabstii 28108!). T A variation named Zygopetalum crinitum pabstii oscano is found in the central region of Espı́rito comb. et stat.subsp. nov. Zygopetalum (Toscano) pabstii Santo, Brazil. Its habitZ. crinitum is terrestrial and the Meneguzzo, — itself is larger than and consequently all Toscano de Brito (1980: 115). Type: BRAZIL. the organs are consistently larger. The flowers Espı́rito Santo:ex hort. Alfredo Chaves, TodosR.A. Kautsky os Santos, 666 mostly haveZygopetalum the same characteristics of other Pico do Sal, , 31 March 1980,. species of , except for a much larger (holotype HB 71173!). Figure 3D side and sepals and petals strongly convex insteadZ. ZYGOPETALUM CRINITUM crinitumof convex. and Since Z. pabstii the only differences between KEY TO THE INFRASPECIES OF are the habit, the flower – measurements are consistent, and populations do 1. Epiphyte; blade short (20.9 38.0 cm long); not intermix, it is hereinZygopetalum proposed that crinitum it be reduced shorter than pseudobulb– plus leaves; topabstii subspecies level as subsp. labellum narrow (midlobeZ. crinitum 2.5 3.3 cm wide);crinitum sepals (Toscano) Meneguzzo.Zygopetalum mackaii var. and petals convex ….… subsp.– convexumThe Mutel names and Z. microtos 1’lon. Terrestrial; leaves blade long (45.6 105.0 cm Hoffmanns. are not as g); inflorescences longer than pseudobulb– plus validly publishedZ. crinitum because theyurland were et merely al. cited leaves; labellum wide (midlobe 4.3 7.1 cm wide); synonyms of Zygopetalum(T mackaii2018: Art.var. sepals and petals stronglyZ. convex crinitum ……… ………….pabstii……… 36.1(b)roseum andRollisson, 36 Ex. Z. 7). mackaii var. macranthum ………………………………….... subsp. Rollisson, and Z. mackaii var. major are denoted as Zygopetalum crinitum crinitum

Lodd. subsp. denominations or naked names by lack of a proper is a species distributed in the Atlantic Rain Forest diagnosis or description (l.c.: Art. 6.3, 38 Ex. 1, and H eringeriana - 14(2): 157 191. 2020. 173

Zygopetalum stenochilum Meneguzzo, T.E.C.

Rec. 50B.1). Lodd. is not present an analysis (l.c.: Art. 6.3, 38.7, 38.8, 38 Ex. 1, validly published due to the lack of a diagnosis or and Rec. 50B.1), but was latter validated by Drapiez description and the fact that its illustration does not (1834).

Figure 3. Warreopsis and Zygopetalum. . Warreopsis colorata colorata.

B. WarreopsisPictures colorata of living specimenspurpurea of. . Zygopetalum crinitum A crinitum. . Zygopetalumsubsp. crinitum pabstii. subsp. C B subsp. D subsp. A. by A. Kay and facilitated by Flickr. . by M. Stickrod .and facilitated by iNaturalist. C. by user ‘Orch’ and facilitated by Wikimedia Commons. D. by T.E.C. Meneguzzo Maxillaria maculata

7. Sprengel (1826: 735). (Kunth)— Zygopetalum maculatum Lindley (1832: 149). Type (designated here): maculatum maculatum in(Kunth) Garay subsp. PERU. Jaén: Between JaénF.W. H.A.de Bracamoros, Humboldt &Sagique A.J.A. Dendrobium maculatumf. Garayin Humboldt (1970: 189).et al. andBonpland Pucura, 3621 August, BroughtoniaKunth maculata (lectotype P QR code P00669292!). (1816: 359). (Kunth) Figure 4A. H eringeriana - 14(2): 157 191. 2020. 174 Notes on Zygopetalinae.

Zygopetalum mackaii Fraga, M.M. Saavedra & M.O.O. Pellegrini 3326

Eulophia mackaiana Hooker (1827: t. —S. loc., ex nom.2748). s uperfl. et illeg. EulophiaLindley mackaii (1831: t. 1433), (neotypehort., s.d., RBs. leg. 599865!). s.n. Original material:syn. nov. (Hook.)— Zygopetalum(not rivierii found), Steudel S. (1840: loc., ex 605). hort. Type (designatedJ.T. here): Mackay — Carrière (1873: 191).S. BRAZIL.s.n. , February 1827, Typeloc., ex (designated hort. here):Rivièri BRAZIL.s.n. Minas Gerais. (lectotypeZygopetalum K barcode mackaii K000589006!). var. intermedium , 1870, (the lectotype is the mackayi original illustration published in the protologue, Mutel — (1842: 9), as “ ”. Type (designated Carrièresyn. nov. 1873: s.n. between pages 190 and 191!), here): BRAZIL. Rio de Janeiro: Rio de Janeiro, Zygopetalum mackaii var. superbum Parque NacionalE.M. da Saddi Tijuca, & ridgeL. Cardoso of Pedra 275 da Gávea, Rollisson in nom. nud. 10 June 2007, — (neotypeS. loc., ex —S. loc.Burbridge, ex hort. , (1874:s.d., s. leg. 160), s.n. RBhort. 466296!)., TaffinOriginal s.n. material: BRAZIL. Material: Zygopetalum messangei Moore (not& Masters found). August,Zygopetalum brachypetalum(not found). — — LindleyS. (1844: loc., ex (1876: 603). Type (designated here): BRAZIL. Sã o hort.misc. 9). Type (designatedWaterhouse here): s.n.BRAZIL. - Paulo: SãoO. Paulo,Handro Campo 504 Grande station,RB 21 August, , July 1840, syn. nov. (lectotype K L 1955, (neotype 728512! barcode Zygopetalum K000857174!), intermedium isoneotypes HB 85133!,—S. loc. SP, ex 93159!,hort. SPF 65077!). in LoddigesZygopeta lum ex Messrs.Original Jacob material:-Makoy & Co. s.n. , 30 Aprilsyn. 1876, nov. Lindleymackaii var.Lindley intermedium(1844: misc. 9). Zygopetalum intermedium(not found),var. peruvianum nom. illeg.(Lodd. ex Lindl.) — S. Nicholson— (1887: S.246), loc. , ex hort., s.d.Type, C.L. (designated Loddiges loc.Rolfe (1894:A. 71). Matthews Type (designated 1896 here): PERU. here):1136 BRAZIL. - , 1835, (lectotype—S. loc. K, ex barcode hort., (lectotypeZygopetalum K L barcode mackaii K000857176!). var. parviflorum K000880334!).s.d., L. Linden s.n. Original material: in ZygopetalonRegel Zygopetalum(remaining protheroeanum syntype not found). mackayivon Kuester var. parviflorum & Regel (1855:Zygopetalum 22), as “ mackaii var. — Rolfe Rio (1899: de minor Rollisson in ”. nom. 287). Type (designated here): BRAZIL. superfl. et illeg. Burbridge (1874: 160),— Janeiro: Itatiaia, Parque Nacional do Itatiaia, Type (designated here): BRAZIL. Macieiras,J.M.A. trek Braga to Macieiras4510 shelter, 3 December Espı́rito Santo. Castelo, arredores da sede do ParqueR.C. 1997, —S. loc., ex(neotype hort. RB 414990!). EstadualForzza, J.A. do Amaral, Forno D. G rande,Monteiro 6 & August R.G.B. 2003,Silva 7654 W.Original Cobb s.n.mat erial: syn. nov. , 15 August 1899, —S. loc., ex Zygop(notetalum found), mackaii var. pallidum Rollison hort.(neótipo, s.d. ,RB s. leg 585452!).. s.n. Original material:syn. nov. in nom. nud. Zygopetalum Zygopetalum(not mackaii found), var. pictum in pallidumBurbridge (1874: 160), nom. nud. Zygopetalum ZygopetalonRegel brachypetalumRolfe (1900:var. pallidum 320), vonmackayi Kuester var. & pictum Regel (1855: 23), as “ Zygopetalum pallidum Cogniaux (1901: t. 5). ”. Type (designated here): (Cogn.)—S. loc. Rolfe, ex hort. (1905:, s.d. 27)., A. BRAZIL. Rio de Janeiro: Petrópolis, Araras, Vale C.N.das TypeBuchan (designated s.n. here): Videiras, ridge of Morro do Cuca, 26 June 2011, (lectotype is the original illustration! H eringeriana - 14(2): 157 191. 2020. 175

Meneguzzo, T.E.C. publishedsyn. nov. in the protologue, Cogniaux 1901: t. 5!), Botânico do Rio de Janeiro, bound in volume 5, and Zygopetalum bolivianum later published in the book with same title by — SchlechterS. loc. (1922a: Barbosa Rodrigues 1996: 361, t. 236!; type J. 50).M. Bang Type 435 (designat ed here): BOLIVIA , 1890, Barbosa Rodrigues personal herbarium destroyed). (lectotype US 814920!, isolectotypes Figure 4D.Zygopetalum brachypetalum var. AMES 75624!, BM barcode BM000074603!, F stenopetalum 77447!, G s.n.!, GH 90652!, K s.n.!,- MO 2158877!, NY — Regel (1887a: 374).S. loc. Type, ex (d hortesignated, 1477510!, Zygopet NY 1477511!,alum sincoranum W 1890 1548!). Lietzehere): s.n.BRAZIL. Minas Gerais. ., April — Castro Neto & (the neotype is thesyn. illustrationnov. published Campacci (2000: 36). Type: exBRAZIL. hort. Bahia. Barra by Regel 1888a: t. 1277!), V.P.da Estiva,Castro Serra s.n. dosic Sincorá,recte S. Koehler, 4 March 11/00 2000, ZYGOPETALUM MACULATUM [ ] [ syn. nov. ] KEY TO THE INFRASPECIES OF (holotype UEC 111157!), – Zygopetalum maculatum maculatum 1. Leaves blade short (6.8– 22.4 cm long); charlesworthii subsp. comb. etf. pseudobulbs short (2.4– 3.6 cm long);ls stat. nov. Zygopetalum(R.H.Pearson) mackaiiMeneguzzo, inflorescences short (19.4 25.6 cm long); sepa charlesworthii [unranked and petals light green just on proximal portion and infraspecies]Zygopetalum mackayi charlesworthiiPearson (1912: 83), as eventually on apex, distal portion dark brown, “ — S. loc., ex ”.hort. Type rarely light green background with large and almost (designated here):Messrs.BRAZIL. Charlesworth & Co. s.n. , 23 completely confluent dark brown maculae; labellum January 1912, (the purple with dark purple confluent veins, commonly. lectotype is the original illustration! published in– fading towards the apex Z.……………………………………. maculatum triste the protologue, Pearson 1912: 83, t. 37!). Figure 4B ………………………………..... – subsp. C. Zygopetalum mackaii var. albicans in. 1’. Leaves blade long (28.9– 65.8 cm long); sched., as “mackayi — Dronk, pseudobulbs long – (3.9 12.5 cm long); ”. Material: BRAZIL. Paraná:A.G. inflorescences long (25.1 78.9 cm long); sepals and DrankBalsa Nova. sic Sãrecte o Luiz A.G. do Dronk Purunã, 1 1 October 1998, petals light green with maculae confluent or not, or [ ] [ ] (MBM 235052!). light green to light brown with- no maculae; labellum Zygopetalum maculatum triste white with purple and non confluent veins, rarely2 , comb. subsp.et stat. nov. light rose …………………………………………………….…..... (Barb.Rodr.)Zygopetalum triste Meneguzzo 2. Sepals and petals light green to lighteral brown lobes with and Barbosa— Rodrigues (1877: 108). no maculae; labellum white, rarely lat Type (designated here): BRAZIL. MinasJ. Barbosa Gerais: Z.midlobe maculatum with light rosemaculatum veins ………………………….. charlesworthii…. CaRodriguesldas, Pedra s.n. Branca, March 1859, subsp. f. (the lectotype is the original 2’. Sepals and petals light green not; with labellum light brownlateral illustration!Iconographie des that Orchidées was du to Brésil be published in maculae either confluent or . t. 11, deposited lobes Z. andmaculatum midlobe with purplemaculatum veins ……………………. maculatum at the library of the Instituto de Pesquisas Jardim …... subsp. f. H eringeriana - 14(2): 157 191. 2020. 176 Notes on Zygopetalinae.

Figure 4. Zygopetalum. Zygopetalum maculatum maculatum maculatumPictures. B– Zygopetalum of living specimens maculatum of maculatumA. charlesworthii Zygopetalumsubsp. maculatumf. triste. C. – subsp. f. . D. subsp. A, D. by T.E.C. Meneguzzo, B C. by W. Pisoni. H eringeriana - 14(2): 157 191. 2020. 177

Zygopetalum maculatum Meneguzzo, T.E.C.

maculatum maculatum (Kunth) Garay adjacents areas of loweret al. altitude, but neverZ. subsp. f. is the species of its maculatumympatrically. an dCampacci Z. triste (2017) found that genus with the widest distribution and that has the freely cross and produce most extensive list of synonyms. In Brazil it occurs viable seeds under controlled experiments, which from central Bahia to Rio Grande do Sul, and in the reinforces their affinity. These two biological Bolivian and Peruvian Andean. It may also be entities present significant discontinuity- in possible to occur in Ecuador. The species is morphological features and do not co occur in same relatively common in high altitude regions usually habitat. Therefore, it is proposedZygopetalum that two maculatum distinct bordering thickets and Z.rupestrian brachypetalum fields. The studyZ. subspecies be recognized:maculatum and Zygopetalum ofbrachypetalum the protologue var. of pallidum Z.Lindl., rivierii maculatum(Kunth) Garay subsp.triste Z. sincoranum Cogn., subsp.Zygopetalum (Barb.Rodr.) maculatum Meneguzzo. Carrière, and V.P.Castro & Campacci maculatumAs for subsp. leddistr to their inclusion as new synonyms since their , there is a specimen in herbarium MBM, ibution and morphology agreesZ. maculatum with that of the in which the flower colour differs from the typical well established and accepted . For the by the sepals and petals being light brown with no same reasons, the study of the protologues andZ. maculae and the labellum with light pink veining. It mackaiineotype var. designation intermedium led toMutel, the inclusionZ. mackaii of var. was collected in the district of São Luiz do Purunã,tion parviflorum el, Z. mackaii var. pictum Z. municipality of Balsa Nova, Paraná. This varia messangei T.MooreReg & Mast., and Z. protheroanumRegel, hasZygopetalum been known mackaii for some time among orchidists as ‘Dronk’ (cultivar) in reference Rolfe as new synonyms. It is worth noting that the to its collector and grower Alessandro Garrett last names. is currently absent from all botanic name Dronk. It was reported (Valter Zomer Pisone, pers. indexe comm.) that a similar cultivar named- ‘Valter Pisone’ In the states of Minas Gerais, São Paulo, and was also obtained from the self crossing of the Paraná, in rupestrian fields, at high altitude areas cultivar ‘Dronk’. Among other siblings obtained, a like the top and ridgesZygopetalum of ranges, triste a Barb.Rodr. variation new cultivar hence named ‘Orleans’ presents a traditionally named Z. maculatum different colour variation that widens the occurs. It differs from by being plants circumscription of the taxon by the sepals and considerable shorter in stature with small petals being light green with no maculae and having pseudobulbs, which are frequently buried in the a white labellum with no Zygopetalum veining. During a substrate, and by the petals and sepals being bibliographic review of the , it was predominantly brown, rarely with a green retrieved that this variationZygopetalum had already mackaii been background with large almost confluent maculae, described and named ascharlesworthii , and and the purple labellum in which veining is very [unranked infraspecies] that faint and fades towards the apex. The author himself this was very similar indeed to those describedBatemannia for and others (Thiago Vinicius Silva Campacci, pers. theperuv cultivariana and ‘Dronk’. Zygopetalum As it occurs xanthinum with var. major, comm.)Z. hadmaculatum reported that it is not uncommon to Zygopetalum find blooming simultaneously in the authorship of the debatable H eringeriana - 14(2): 157 191. 2020. 178 Notes on Zygopetalinae.

nom. superfl. et illeg. — infraspeciesThe Gardeners’ is attributed Chronicle to the editor of the journal 83), s.Type loc. ,(designated ex hort. here): at that time, in case Robert BRAZIL.F. RioWarre de Janeiro:s.n. , September Hooper Pearson (1912) (Anonymous 1918). 1831, (the lectotype is the original Because this variation is sporadic,Z. andmaculatum of low illustration! deposited at the Natural History frequency amongst populationsmaculatum of Museum, London, and published in. the protologue, (Kunth) Garay subsp. , itZygopetalum is hereby Loddiges Zygopetalum 1832: t. 1772!). gautieriFigure 5A maculatumrecognized as a form by the namecharlesworthii Zygopetalum maxillare var.Lemeire gautieri (1867: t. subsp. maculatum f. 535). in (Lem.) (R.H.Pearson)Eulophia Meneguzzo. mackaiana dl. and Zygopetalum Regel Regel & Maximowicz— (1870: 34). Type mackaii var. minor Lin (designated here): BRAZIL. Santa Catarina:– Rollisson are respectivelyZ. mackaii Florianópolis [“IÎleA. Verschaffelt de Ste. Catherine”], s.n. September superfluousZ. mackaii and illegitimate var. parviflorum names of et al. October 1867, (the lectotype is Hook. and Zygopetalum(Turland mackaii var. the original illustration published in the protologue, 2018:intermedium Art. 52.1 and 52.2). Lemeire Zygopetalum 1867: t. 535!). obtusatum Nicholson (1899) is an illegitimate Reichenbach—S. loc., ex name of the earlier heterotypicZygopetalum name mackaii by Mutel var. (1878:hort 300). Type, T. (designated Lawrence s.n. here): -R superbum(1842) (l.c.: Rollisson, Art. 53.1). Z. mackaii var. pallidum ., 11 June 1878 (lectotype W Rollisson, and Z. pallidum 40643 syn. , nov. upper left and bottom left Rolfe are denoted as icon!), Zygopetalum graminifolium in denominations or naked names due to the lack of a Rolfe Rolfe & proper diagnosis or description (l.c.: Art.Zygopetalum 6.3, 38 Ex. Baker (18— 92: 179). Types. (designatedloc., ex hort. here): 1,mackayi and Rec. 50B.1). The spelling “ BRAZIL.Messrs.Southern F. Sander region, & Co. s.n. , April ” has been massively favoured in the 1891, syn. nov. (lectotype K botanicalZygopetalum literature mackaii in spite of the original spelling barcode Zygopetalum K000589009!), hasslerianum “must st ”. However, the first spelling — Kränzlin (1906: and not only because it is the original one 389). Type (designated here): PARAGUAY.É. HasslerGuairá. (l.c.: Art. 60.1), but also because it resulted from the Ca9020raguazú [“Caaguazú”],- February 1905, intentional Latinization of the surname Mackaymackaii to (lectotype S 07 7834!, isolectotypes BR s.n.!, G Mackaius, therefore the correct epithet is 87511/1!,- K barcode K000880339!, NY 9458!, W (Nicolson 1974). 1906 1439!).Zygopetalum rigbyanum Zygopetalum maxillare — Ruschi (1975: 1).

8. Type:, A.BRAZIL. Ruschi s.n.Espı́rito Santo. Castelinho, 11 April Zygopetalum maxillare maxillare in 1974 —S. (holotypeloc. MBMLs. leg. spirit s.n. 2004!). Eulophia maxillaris Loddiges f. Other material: , April, (living in Loddiges (1832: t. 1772). specimen and orchid greenhouse F.C. Hoehne— Zygopetalum maxillare var. typicum in (Lodd.) Loudon Steudel (1840: 605). accession number 775 at MBML, not seen non valid. publ. Regel Regel & misnamed as “paratype”) MaximowiczZygopetalum mandibulare (1870: 34), H eringeriana - Hoffmann segg (1844: 14(2): 157 191. 2020. 179

Zygopetalum silvanum Meneguzzo, T.E.C.

—Castro Neto & morphological dissimilaritiesZ. maxillare. fit in the Campacci (1991: 27). Type:ex hort.BRAZIL. Bahia. circumscription of Itororó,E.P. Silva Serra s.n. de Ouricana, , syn.17 Aprilnov. 1984, By the endZyg ofopetalum the 19th sanderianum century, Regel (holotype SP 333601!), (1888b) described Regel Zygopetalum maxillare sanderianum from a specimen of unknown provenance which is comb. et stat.f. nov. Zygopetalum(Regel) characterized by green sepals and petals with small Meneguzzo,sanderianum Zygopetalum dots in the proximal portion and light lilac labellum. maxillare var.Regel sanderianum (1888b: 657). Later, Cogniaux (1898a)Z. maxillare attributed it with a new (Regel)— S. Cogniaux loc., ex status Z.as asanderianum variety of . It is proposed toZ. (1898a:hort., s.d., 577). H.F.C. Type Sander (designated s.n. here): keepmaxillare asZygopetalum an infraspecies maxillare of (the lectotype is the sanderianum, therefore as f. original illustration published in. the protologue, (Regel) Meneguzzo. It is proposed as a

Regel 1888b: t. 1287!). Figure 5B form because the only difference is the variation in ZYGOPETALUM MAXILLARE

flower colour and only a few individuals with this KEY TO THE INFRASPECIES OF flower colour sporadically appear among 1. Sepals and petals light green with light brown populations with the common coloured flowers. maculae, which are- confluent in the proximal The circumscription is somewhat expanded to portion and non confluent in the distal one; include specimens with sepals and petals green Zygopetalum maxillare var. labellum lilac to purple occasionally attenuate. dotted or not and labellum light lilac to white. typicum towards the apex, callus darkeZ. maxillarer ………………………. maxillare… … The name ….……………………..………...…. f. Regel is not a validly published name et al. 1’ Sepals and petals light green with no maculae, because its final epithet might be equal to the Zygopetalum mandibulare rarely with small light brown dots in the proximal second one (Turland 2018: Art. 24.3 and 26.2). Z. maxillare portion; whole labellum light lilac to white. or Hoffmann. is a eventually light lilac inZ. the maxillare proximal portisanderianumon …… …… superfluous and illegitimate name of Zygopetalum rigbyanum ……………..….………….... f. Lodd. (l.c.: Art. 52.1 and 52.2). The living Zygopetalum maxillare nomenclatural type of maxillare Lodd. subsp. Ruschi is not a preserved specimen, therefore it can is a widely spread species in the Atlantic not be accepted as a type (l.c.: Art. 8.4). Rain Forest from southern Bahia to Rio Grande do labiosum

Sul, Brazil, and northeastern Argentina and eastern 9. ts labellum is Paraguay. Typically its sepals and petals are light Zygosepalum labiosum green with light brown maculae and i labiosum in (Rich.). Epidendrum Garay labiosum subsp. lilac to purple, occasionally with an attenuated hue Garay (1967:Menadenium 6) labiosum Z. obtusatum towards the apex. The study of the type specimens Richard (1792: 112). Zygopetalum labiosum(Rich.) Z. graminifolium Z. silvanum accompanied by the protologue of Cogniaux (1898b: 582). Zygosepalum Rchb.f., Rolfe, and V.P. (Rich.)labiosum L.O. Williams (1942: 16). nom. Castro & Campacci, led to their being newly (Rich.) Schweinfurth (1967: 184), Hproposed eringeriana synonyms, - since their putative 14(2): 157 191. 2020. 180 Notes on Zygopetalinae. superfl. — lindeniae Menadenium lindeniae

S. loc. Type –(designatedJ.B. here):Leblond FRENCHs.n. GUIANA. Rolfe (1891: 73). Zygosepalum , 1787 1789, . (lectotype P (Rolfe)lindeniae Cogniaux (1898b: 584). in barcode Zygopetalum P00612111!). rostratumFigure 5C (Rolfe) Garay & Dunsterville Menadenium rostratum Hooker (1828: t. Dunsterville—S. loc. &, Garayex hort. (1965: 336). TypeBungeroth (designated sub 2819). non valid. publ. (Hook.)Eulophia Rafinesque rostrata here):Messrs. Linden s.n. , April 1891, (1838: 45), Zygosepalum .(lectotype K barcode (Hoorostratumk.) Steudel (1840: 605). K000589026!). ZygosepalumFigure revolutum 5D - (Hook.) — Reichenbach (1859: 330).s. loc.Type, ex — Garay & Romero (designatedhort. here): GUYANA.C.S. Parker Demerara: s.n. González (1999: 487). Type:s.d., G.COLOMBIA. Escobar RestrepoVaupés: , October 1827, (lectotype K Miraflores,1039 Vaupés River, syn. nov. barcode Zygopetalum K000589024!). kegelii (holotype AMES 287162!), Zygosepalum kegelii Reichenbach (1852a: Zygosepalum labiosum marginatum 668). non valid. publ.(Rchb.f Zygosepalum.) Reichenbach kegelii nothosubsp.comb. et stat. nov. (1859: 330), Menadenium (Garay)Zygosepalum Meneguzzo, × marginatum kegelii(Rchb.f.) Reichenbach (1863b: 666). Zygopetalum × marginatum Garay (1967: 8). (Rchb.f.) Cogniaux— (1898b: 584). Type nom. illeg. Mendoncella (Garay)× marginata Garay (1973: Tourtonne,(designated ex here):hort. SURINAME. H.A.H. Geiersvlyt Kegel 1427 and 32), × marginata (Garay) sub van Houtte s.n., January 1849,- Garay (1973: 32). in (Garay) H.A.H. Kegel(lectotype 1427 WsubR van 40623!); Houtte idem, s.n. Dressler— & ChristensonS. loc. ,Chris ex hort.tenson, s.d. ,(1989: C. Mark 222). sub May 1846, - syn. nov. Type:Joseph COLOMBIA.Redlinger Orchids sub W.C. Cornett s.n.

(remaining syntype W R 17644!), Zygosepalum labiosum ballii (holotype— Galeottia AMES 102311!). grandiflora Parent taxa (postulated comb. et stat. nov. Zygopetalum × ballii nothosubsp. (Rolfe) here):Zygosepalum labiosum A.Rich.lindenia & Galeottie × Zygopetalum × ballii Meneguzzo, . . subsp. (Rolfe) nom. superfl. Menadenium × ballii Rolfe (1900: 149). Rolfe (1906: Meneguzzo Figure 5E Zygosepalum × ballii 33), (Rolfe) ZYGOSEPALUM LABIOSUM nom valid. publ. Zygopetalum × Cogniaux (1906b: 576). (Rolfe) KEY TO THE INFRASPECIES OF ballianum in sched. d Garay (1967: 8), - 2 — sic recte H.J.Veitch, Type (designate 1. Anther cap apex short- rostellate …………………….. 3 ex hort. here): BRAZIL. [ ] Pernambuco [ northern 1’. Anther cap apex long rostellate ……………..………. Messrs. J. Cowan & Co. sub G.S. Ball s.n. South America, possibly Brazil], , June 1898, 2. Labellum callus hippocrepiform and canaliculate (lectotype K with margin smooth, Z.midlobe labiosum margin entire … …….ballii… —Batemannia colleyi barcode K000880371!, isolectotype AMES 4693!). ……………..…..………..... nothosubsp. colleyi × Zygosepalum labiosum Parent taxa (postulated here): 2’. Labellum callus widely hippocrepiform and lindeniae . Lindl. subsp. subsp. concave with margin digitate, midlobe margin

(Rolfe) Meneguzzo irregularly. serrateZ. labiosum …………………………………….. marginatum………... Zygosepalum labiosum lindeniae ………………and nothosubsp. comb. et stat.subsp. nov. Zygopetalum(Rolfe) 3. Sepals petals light green to light brown, HMeneguzzo, eringeriana - 14(2): 157 191. 2020. 181

labellum white Meneguzzo, T.E.C.

, callus and proximal portion of the Meneguzzo.Zygos epalum marginatum midlobe with a few crimsonZ. labiosum veins ……………… labiosum…….… was described by Zygopetalum Mendoncella, ……………………...... …….. subsp. Garay (1967) and shortly after combined by the 3’. Sepals and petals light brown, dark brown to dark author to , then to red (rarely light green), whole labellum white to showing how unusual is its morphology. It was not Zygopetalum light crimson and denselyZ. labiosum veined ……………lindeniae……..….….. a perfect fit to the generic concepts as stipulated by

…………………………….. subsp. Garay himself in his overview of the Zygosepalum labiosum complex (Garay 1973). It is herein postulated that labiosum (Rich.) Garay subsp. thisGaleottia entity grandiflora is a nothospecies with parent taxa is a widespread epiphyte species from rain ZygopetalumRichard & Galeotti labiosum (1845: 33) forests along northeastern South America, (lindeniaeFigure 5F) and subsp. occurring in Venezuela, the Guianas, and eastern due to the intermediate nature of its Brazilian Amazon. It apparently grows in the morphology in some aspects and equal to one or the proximity of water bodies and lower parts of tree other parent in others, and sympatry of both trunks under dense forest canopy. Its flower parents on its country of origin. It has a reptant and morphology does not vary significantly, being the rhizomatose habit, pseudobulbs sheaths, subtend sole variation that the labellum has is more or less on base of pseudobulb, base margin hyaline, prominent lobes or that the base articulate, leafy, flower colour, lip callus widely cordate.Zygosepalum Cogniaux rostratum (1898b) under synonymizedZygosepalum hippocrepiform, and lateral lobes length between labiosum as Menadenium apexes of the lateral lobes narrowerZygosepalum than the ( ZygosepalumRaf. ex Cogn.), kegelii and he also midlobelabiosum width lindeniae inherited from proposed the inclusion of in the subsp. . Whilst the lip callus is synonymy. widely hippocrepiform, wider than width, In northwestern South America, in composed of several longitudinal carinae, and the Venezuela, Colombia, Ecuador, Peru, and western lip midlobe marginZygopetalum is denticulate, grandiflorum characters BrazilianZygosepalum Amazon, lindeniae a biological entity named inherited- from . The (Rolfe) GarayZygosepalum & Dunst. shortintermrostellate column anther cap apex is clearly occurs,labiosum which only differs from ediate, as in the former- parent it is rounded, by sepals and petals that are usually and in latter parent it is long rostellate. The rarity of darkerZygopetalum and revolutum the densely veined labellum. nothotaxa in natural habitats is a feasible is only know from the type explanation why it has never been recollected. Since specimenZygosepalum and has a morelindeniae deeply cordate labellum onera of its postulated parents is of lower taxonomic than in , but this sole feature nk than the other, the nothotaxonand et al. must equally be does not support its recognition as a good species at the lowest rank (Turl 2018: Art. H.5 Ex. arand therefore it is synonymized. Since the two taxa 2under and H.11.2).Zygosepalum Hence, itlabiosum is herein stipulated to be e allopatric and only morphological difference is Zygosepalumand the labiosum new the flower colour, and mainly in the labellum, it is combination marginatumis proposed too: proposedZygopetalum it labiosum be reduced to subspecificlindeniae rank: nothosubsp. (Garay) Meneguzzo. H eringeriana - subsp. (Rolfe) 14(2): 157 191. 2020. 182 Notes on Zygopetalinae.

Figure 5. Zygopetalum, Zygosepalum and Galeottia. Zygopetalum maxillare maxillare.

B. ZygopetalumPictures maxillare of living specimenssanderianum of . Zygosepalum labiosum labiosumA. Zygosepalum labiosumf. lindeniae Zygosepalum labiosumf. C. marginatum Galeottiasubsp. grandiflora. . D. subsp. . E. nothosubsp. . F. A. by T.E.C. Meneguzzo. B. by P. Fabro. C. by user ‘Orch’ facilitated by Wikimedia Commons. D. by K. Senghas. and facilitated by Swiss Orchid Foundation at Herbarium Jany Renz. E. by L.A. Garay and facilitated by J. Pfahl F. by Ecuagenera and facilitated by Swiss Orchid FoundationH eringeriana at Herbarium Jany- Renz. 14(2): 157 191. 2020. 183

Zygopetalum balliiMeneguzzo, as T.E.C.

Rolfe (1900) described conspicuous, and the anther cap short rostellate vs. a new species based on a cultivated specimen rounded. Both parent species and the hybrid shares without precedence. He had access only to the column with peculiar morphology for the genera preservedve flowers and on the report of the constituted by the clinandrium margin projectedurs, getative part that was written by the owner of the over theZygopetalum anther cap. labiosum On the grounds oflindeniae the colo and plant,mate which is reflected on the state of the original bothBatemannia colleyi colleyisubsp. rials. Later Rolfe (1906) redescribed the taxon subsp. have petals and and indicate to be from the state of Pernambuco, sepals brown with lighter hue towards the margins, Brazil.Menadenium Cogniaux (1898b) combined it to the genus the lip in the first is white to light crimson and ZygosepalumRaf. ex Cogn. and Garay (1967) to the denselyZygopetalum veined × ballii and on the second white. genusbetter unde ; none of them contributed to a has sepals and petals with rstanding of the biological entity and same colour, but in the original material the petals merely repeated the limited previously published diffused and discreet blotched with dark rose data. Schlechter (1919: 86) cited its occurrence to instead of brown, the lip white with the proximal Bolı́var, Venezuela, without giving any extra half portion crimson. information to track neither provenance, nor the ZygopetalumThe locality× ballii of the type specimen of specimen. The identity of this putative species had is disputed as Pernambuco not been elucidated and Zygopetalum recollected. × It ballii is herein as a since none of the parent taxa occurs there, but postulated the nature of Batemannia colleyi instead a putative locality in northwestern north nothospeciescolleyi wiandth parentZygosepalum taxa labiosum America area of the Amazon forest where both subsp.lindeniae subsp. parents does occur. It concurs with the observation based on the unique combination of of a specimen confidentially identified as this taxon characters that does not neatly match any of both from São Gabriel da Cachoeira, Amazonas, Brazil, genera, but are quite intermediate and on which and kept under cultivation (Cássio van den Berg, certain characteristics can clearlyZygopetalum be picked ballii from pers. comm.). This specimen does not bears the parents.Zygosepalum The similarities of ulbs blotchedmaterial marksdoes. on the perigonium as the originalame with is inevitable: pseudob Hence, following the s laterally compressed and 3‒4 leaved, sepals and nomenclatural rule of nothospeciesZygosepalum with labiosum parents petals subsimilar in shape and slightly convex, lip different ranks,marginatum as in et al. ovate- with inconspicuous lateral lobes, callus high, nothosubsp. (TurlandZygopetalum ×2018: ballii Art. is Batemanniaerect patent, and canaliculate. When compared to, H.5 Ex. 2 and H.11.2), , sepals and petals subsimilar in size combinedZygosepalum tolabiosum the andsubspecies level under convex and about 60º between them vs. sepals subsequentlyZygosepalum as labiosum it status longer than petals, flat and about 30º, the lip lacks changed to aballii nothospecies: lateral lobes, the callus has no resemblance as nothosubsp. (Rolfe) Meneguzzo. already described vs. distally free and long aciculaten ZygosepalumSome nomenclatural commentsMenadenium about to denticulate, but instead with sessile apex, colum Rchb.f. and its synonym with inconspicuous stigmatic wings vs. Raf. ex Cogn. are necessary to clarify their tortuous H eringeriana - 14(2): 157 191. 2020. 184 Notes on Zygopetalinae.

. Zygosepalum labiosum history and Zygosepalum labiosum (Rich.) Garay (1967: (1859: 330) is not a validly published Znameygosepalum as it is 6) (Rich.) Schweinfurth merelyrostratum cited as a synonym of (1967: 184) were published in different journals. et al.(Hook.) Reichenbach (1859: 330) Surprisingly both bear the same year and month (Turland 2018: Art. 36.1(b) and 36 Ex. 7), but cover date, and thderefore et al. they are equal priority- later validlyZygopetalum published lindeniae by Reichenbach (1863b: homonyms (Turlan 2018: Art. 53.5). Romero 666Zygopetalum). lindenii Rolfe (1891: 73) and González (2005) literally stated that Garay's name Rolfe (1892: 5) are legitimate undoubtedly preceded Schweinfurth's, but failed by names, not homonyms as commonlyet al. stated in not presenting such evidence or rejecting the latter. botanical literature (Turland 2018: Art. 53.2, Notwithstanding there is no internal evidence in 53 Ex. 12, and 60 Ex. 23),Warczewiczella being the latter amazonica currently bothZygosepalum works labiosum of which was earlier published. accepted as synonym of (Rich.) Garay (1967: 6) is Rchb.f. & Warsz. hereinZygosepalum adopted labiosum simultaneously rejecting Contrary to whatZygopetalum Rolfe (1906) × believed,ballii the (Rich.) Schweinfurth (1967: original publication of (Rolfe 184), whose legitimate statuset al. is kept but turned to 1900) is indeed validly published because it fulfilsin bean superfluous (Turland 2018: Art. 52.1, 53.5, all the requirements, even though it lacks a Lat d 53.5 Note 4). Menadenium diagnosis which is mandatory fromet al. 1 January 1935 The genus Rafinesque (1838: to 31 December 2011 (Turland 2018: Art. 32.1 45) was not validly published because it was not and 39.1). Hence, the second of Rolfe’s publicationet accepted byet al.the author at the time of its publication onal. the name is superfluous (Rolfe 1906; Turland (Turland 2018: Art. 36.1), i.e. by not citing the 2018: Art. 52.1 and 52.2). It has direct names in its own paragraph with unique numbering consequenceZygosepalum × onballii the fact that the combination and by the absence of the genus in the index of the (Rolfe) Garay (1967: 8) turns otto volume. The genus itself was later validly published be a not validly published name because it is n by Cogniaux (1898b:M. kegelii 581) who also publishedM. labiosum the based on the original publication (Rolfe 1900: 149), combination M. (Rchb.f.)lindeniae Cogn., but instead on the later publication that has a directet (Rich.) Cogn., andZygopetalum rostratum(Rolfe) Cogn. A al.reference to the first one (Rolfe 1906: 33; Turland combinationMenadenium of Hook. to 2018: 35.1 and 41.8). never was validly published because, ACKNOWLEDGMENTS as said, Rafinesque (l.c.) did not validly published the genus, and also he did not associatend et the al. genus re I am grateful to C.E.B. Proença for critical name with the specific epithet (Turla 2018: view of this manuscript; C. van den Berg and V.Z. Art. 35.1 and 35.2). SubsequentZygopetalum authors rostratum accepted Pisoni for discussions on the biology and theunder synonymi Menadeniumzation oflabiosum or Zygosepalum distribution of some of the species; J. Prado for labiosum nomenclatural advise; Biodiversity Heritage , hence the combination to any of those last Library, Herbarium Jany Renz, P. Fabbro, Flickr, L.A. genera wouldZygosepalum be unuseful. kegelii Garay, iNaturalist, R. Jenny, A. Kay, ‘Orch’, A. (Rchb.f.) Reichenbach Popovkin, V.Z. Pisoni, J. Pfal, M. Rosin, Swiss Orchid H eringeriana - 14(2): 157 191. 2020. 185

Meneguzzo, T.E.C.silvana Orchidaceae

Foundation, L.F.K. Varella, Wikimedia Foundation, F.Barros & CatharinoHoehnea ( – ), nova and A. Zaslawski for permission to use their images, espécie para o Brasil.Supplement to21: Loudon’s 93 96. Hortus L.C. Meneguzzo for the layout design, and cited Baxter,Britannicus W.H. (1850). herbaria for access to their collections.- This paper Longman, Brown, Green & was supported by a post doctoral researcher Longmans, London, 263Les pp. orchidées: manuel de scholarship from CAPES/JBRJ. Bois,l'amateur. D.[G.J.M.] J. - (1893) REFERENCES B. Baillère & Fils, Paris, 323 pp. The Brittehttp://dx.doi.org/10.5962/bhl.title.125414 – AnonymousGardeners’ (1918) Chronicle Robert Hooper Pearson. n, J. Journal (1907) of Maxwell Botany, Tylden British Mastersand Foreign (1833 , ser. 3, 63: 246. [anonymous 1907).– 45: authorship, but likely to be authored by that time 257 258. editor Charles Henry Curtis.] Brown, N.E.The & Reichenbach,Gardeners’ Chronicle H.G. (1883) New garden Barberena,Promenaea F.F.V.A. & Barros, F. (2015); A new species; plants. Cool orchids,, n.s., 20:and 70. how to ofOrchidaceae ( Burbridge,grow them F.W.[T.] (1874) Phytotaxa ) from Southeastern Brazil.– . R. Hardwicks, London. 160 pp. 203: 205 209. http://dx.doi.org/10.5962/bhl.title.15546 https://doi.org/10.11646/phytotaxa.203.2.11 Campacci, T.V.S., Castanho, C.T., Oilveira, R.L.F., Barberena, F.F.V.A., Baumgratz, J.F.A. & Barros, F. Suzuki, R.M., Catharino, E.L.M. & Koehler, S. (2016) UnvePromenaeailing the enigmaticrollissonii Pabst and (2017)Zygopetalum Effects mackayi of pole orchidsn origin. Flora on apomixis(Jena) in Dungs’sOrchidaceae Novon alliance– – 226: ( ). 24: 325 332. 96 103. https://doi.org/10.3417/2015004Genera et Species Castro Neto, V.P.Zygopetalum & Campacci, M.A. (1991) Uma nova BarbosaOrchidearum Rodrigues, Novarum J. (1877) espécieBoletim CAOB de – do estado da Bahia. 1. C. & H. Fleiuss, Rio de 3 (4): 26 31. Janeiro. 206 pp. CastroPromenaea Neto, V.P . & Campacci, M.A. (1993) Uma nova http://dx.doi.org/10.5962/bhl.title.585Genera et Species – do estado da Bahia. Boletim CAOB 5 BarbosaOrchidearum Rodrigues, Novarum J. (1882) (3): 9 12. 2. Typographia Nacional, CastroZygopetalum Neto, V.P. & Campacci, M.A.Orquidário (2000) Novo Rio de Janeiro. 295 pp. – do estado da Bahia. 14: http://dx.doi.org/10.5962/bhl.title.585 36 37. Zygopetalum rivieri. Revue Barbosa Rodrigues, J. (1891)Vellosia Genera et species– Carrière,Horticole EÉ .[A.] (1873)– orchidearum novarum. , 2nd. ed., 1: 115 45: 191 192. Zygopetalum crepeauxii. 133. Carrière,Revue Horticole EÉ .[A.] (1887) Barbosa Rodrigues, J. (1996) Sprunger,Iconographie S.; Cribb, des P. 59: 420. Orchidées& Toscano du de Brésil Brito, A. (eds.). Chase, M.W., Cameron, K.M., Freudenstein, J.V., 1. Friedrich Reinhardt, Basel. Pridgeon, A.M., Salazar [Chávez], G. [A.], van den 186 pp. Promenaea Berg, C. & Schuiteman, A. (2015) Botanical An Journal updated of Barros, F. & Catharino, E.L.M. (1995) classification of Orchidaceae. H eringeriana - 14(2): 157 191. 2020. 186 Notes on Zygopetalinae.

the Linnean Society –

177: 151 174. http://www.splink.org.br/ (accessed: 1 Dec http://dx.doi.org/10.1111/boj.12234 2020). Otostylis hirtzii. Christenson, E.A.Orchidaceae (1989) Nomenclatural. Lindleyana changes– in. Dodson,Icones C.H. Plantarum & Dodson, Tropi P.M.ca rum(1984) Neotropical 3: 221 223 Hortus Cantabrigiensis10: t. 976. [issue cover y ear 1988, issue effectively Don, P.N. (1845) . 13th ed. published 1989.] Longman & Co., London, 772 pp. Christenson,Orchidaceae E.A.. Phytologia (1992) Notes on – Neotropical http://dx.doi.org/10.5962/bhl.title.160219Encyclographie du Règne 71: 440 444. [issue Drapiez,Végéta [P.A.J.]l (1834) cover date December 1991, effectively published 1. Encycographie, Brussels. 575Venezuelan pp. 18 February 1992.] Dunsterville,orchids illustrated G.C.K. & Garay, L.A. (1965) Christenson,Orchidaceae E.A. II. Lindleyana (1996) Notes on– Neotropical 3. Deutsch, London,Zygosepalum 348 pp. . 11: 12 , 26.an unfinished Garay,Orquideología L.A. (1967) Sinopsis– del género Christenson,monograph. E.A. (2013) Maxillaria 2 (3): 5 9. Zygopetalum - 2 vol. In: Hardin, P.A., McIllmurray, M Garay,Maculatum L.A. (1970) LaOrquideología identidad del & Blanco[ Coto], M.[A.] (Eds.). Editors's edition, H. B. K. 5:Zygopetalum 184, 189. . Lebanon, 936 pp. Orchidaceae Zygopetalum. Garay,Orquideología L.A. (1973) – El complejo Cogniaux,In: Mar C.A. (1898a) : 8: 15 51.- Flora Brasiliensistius, C.F.P., Eicher, A.G. & Urban, I. (Eds.). Garay, L.A. & Romero HarvardGonzález, Papers G.A. in (1999)Botany 3 (5). Typographia Regia,– Schedulae– Orchidum II. Munich. pp. 563 580. 4: 475 488. http://dx.doi.org/10.5962/bhl.title.454Orchidaceae Menadenium. Harrison, J. (1835)The Floricultural Reference Cabinet to andthe Cogniaux,In: A.[C.] (1898b) : embellishments.Florist’s Magazine FloraMartius, Brasiliensis C.F.P., Eicher, A.G. & Urban, I. (Eds.). 3: 72. 3 (5). Typographia Regia,– Hoehne, F.C. (1952) EspéciesArquivos e variedades de Botânica novas dedo Munich. pp. 5 81 585. OrchidáceasEstado de S. Paulo do Brasil. – – http://dx.doi.org/10.5962/bhl.title.454Orchidaceae Promenaea. , n.s., 2: 121 136,In t. 52 58. Cogniaux,In A.[C.] (1906a) : Hoehne, F.C.Flora (1953) Brasílica Orchidáceas. : Hoehne, F.C. Flora: Martius, Brasiliensis C.F.P.; Eicher, .A.G. & Urban, I. (eds.). (Ed.) 12 (2). Secretaria de Muni 3 (6) –Typographia Regia, Agricultura do Estado de São Paulo, São Paulo. ch, pp. 461 469, t. 95. 397 pp, 181 t. Verzeichniss der http://dx.doi.org/10.5962/bhl.title.454Orchidaceae In Hoffmannsegg,Orchideen J.C. (1843) Cogniaux, [C.]A. (1906b) : addenda. : . 2 ed. H.M. Gottschalk, Dresden. 64 pp. Martius,Flora Brasiliensis C.F.P., Eichler, A.G. & Urban, I. (Eds.) http://dx.doi.org/10.5962/bhl.title.87286Verzeichniss der 3 (6).– Typographia Regia,– Hoffmannsegg,Orchideen J.C. (1844) Munich, pp. 525 584, t. 107 120. . 3 ed. H.M.Zygopetalum Gottschalk, mackaii Dresden.. Curtis's 69 pp. http://dx.doi.org/10.5962/bhl.title.454– Hooker,Botanical W.J. Ma (1827)gazine CRIA Centro de ReferênciaspeciesLink em Informação Zygopetalum54: t. 2748. rostratum. Curtis's Ambiental (2020) . Available from: Hooker, W.J. (1828) H eringeriana - 14(2): 157 191. 2020. 187

Botanical Magazine Meneguzzo, T.E.C.Botanical Register

55: t. 2819. Maxillaria23: t. 1986. stapelioides. Edwards’s Humboldt, Nova F.W.H.A., genera Bonpland, et species A.J.A. plantarum & Kunth, K.S. Lindley,Botanical J. (1839a) Register (1816) 1. Huntleya25: t. 17. violacea. Edwards’s Lutetiae Parisiorum, Paris. Lindley,Botanical J. (1839b) Register http://dx.doi.org/10.5962/bhl.title.640 25: misc. 19. [issue published Klinge, J. (1898) Diagnosis Orchidaceum novarum March 1839.] Maxillaria lentiginosa. Edwards’s in caldariis HortiTrudy Imperiali Imperatorarsk Botanici Petropolitaniago S.- Lindley,Botanical J. (1839c) Regist er cultarum.Peterburgsk Botaniceskago Sada – 25:Huntleya misc. 61. violacea. Sertum – 17 (1): 133 Lindley,Orchidaceum J. (1839d) 144, t. 1 3. : t. 26. [issue published December Königer,Cyrtochilum W. (1995), Masdevallia Neue, Oncidium arten der, Promenaea gattung 1839.] http://dx.doi.org/10.5962/bhl.title.204Maxillaria. Edwards’s Botanical und Sigmatoslix. Arcula – Lindley,Register J. (1843) – 4: Orchidaceae90 121. andinae 29: misc.Zygopetalum. 10 16. Edwards's Botanical Kränzlin, F.W.L. (1906) auerianae III. Lindley,Register J. (1844) imprimisBotanische peruvianaeJahrbücher Weberb für Systematic, 30: misc. 9. stapelioides. Pflanzengeschichte und Pflanzengeographie Link,Icones H.F. &Plantarum Otto, F.[F.]. Selectarum – 37: : 111, t. 52. 381 398. Index seminum http://dx.doi.org/10.5962/bhl.title.51952Zygopetalum crinitum. The von1855 Kuester, K. & Regel, E.[A.] (1855) Loddiges,Botanical C.L. Cabinet (1831) . Imperatorskaya Akademiya Nauk, Saint 17:Zygopetalum t. 1687. maxillare. The Petersburg. 20 pp. Loddiges,Botanical C.L. Cabinet (1832) von Kuester, K., Regel, E.[A.],Index Körnicke, seminum F.[A.] & 1856Rach,. 18:Zygopetalum t. 1772. stenochilum. The L.[T.] (1856) Loddiges,Botanical C.L. Cabinet (1833) ImperatorskayaPetersb Akademiya Nauk, Saint 20:A practicalt. 1884. treatise on the urg, 40 pp. Zygopetalum gautieri. Lyons,management J.C. (1845) of Orchidaceous plants. Lemeire,L'Illustration [A.]C. Horticole (1867) J. Ridgway, Eulophia14: mackaiana.t. 535. Edwards's Hodges & Smith, LondonBatemania & Dublin, peruviana 234 pp.. The Lindley,Botanical J. (1831) Register Masters,Gardeners’ M.T. Chronicle (1895) Maxillaria17: t. 1433.. In: The genera and , ser. 3, 17: 551. [published 4 Lindley,species J. of (1832) orchidaceous plants. May 1895]. – Ridgways, London. Masters,The Gardeners’ M.T. (1901) Chronicle Orchid notes and gleanings.– Pp. 142 151. http://dx.doi.org/Zygopetalum. In10.5962/bhl.title.499 The genera and , ser. 3, 30: 106 107. Lindley,species J. of(1833) orchidaceous plants : McNeill, J. (2014) HolotypeTaxon specimens and– type – . Ridgways, London. citations: general issues. 63: 1112 1113. Pp. 187 188. http://dx.doi.org/10.5962/bhl.title.499Batemannia colleyi. Edwards's http://dx.doi.org/10.12705/635.7Systematics of Lindley,Botanical J. (1834) Register Meneguzzo,Zygopetalum T.E.C. (Orchidaceae) (2018) 20:Maxillaria t. 1714. steelii. Edwards’s . PhD thesis. Escola Lindley, J. (1837) Nacional de Botânica Tropical, Instituto de H eringeriana - 14(2): 157 191. 2020. 188 Notes on Zygopetalinae.

Genera Orchidacearum 5.

Pesquisas Jardim Botânico do Rio de Janeiro, Rio Rasmussen,Epidendroideae, F.N. part(Eds.) two de Janeiro. 528 pp. – . Oxford University, New Meneguzzo, T.E.C., Baumgratz, J.F.A. & van denAganisia Berg, York. Pp. -456 546. Flora C. (2015)Orchidaceae Taxonomic, Zygopetalinae studies in theP hytotaxa Rafinesque[TellurianaScham . altz], C.S. (1838) complex ( ). – 4 Author's edition, Philadelphia, 135 238: 1 39. REFLORApp. http://dx.doi.org/10.5962/bhl.title.7751 – http://dx.doi.org/10.11646/phytotaxa.238.1.1 Resgate Histórico e Herbáriora Brasileira para o Menezes,Boletim L.C. CAOB (199 5) Novas– orquı́deas brasileiras. Conhecimento e Preservação da Flo 5 (1): 8 13. (2020) Available from: Moore, T. & The Masters, Gardeners’ M.T. Chronicle (1876) Reports of– http://reflora.jbrj.gov.br/reflora/herbarioVirtu a societies. , n.s., 5: 600 al/ ( ccessed: 1 Dec 2020). 604. Regel, E.[A.] (1887a) Descriptiones plantarum Moore, T. &The Reichenbach, Gardeners’ Chronicle H.G. (1878) New garden nonnullarum Hortio. TrudyImperialis Imperatorskagobotanici in stato S.- Mutplants. , n.s., 9: 588. Peterburgskagovivo examinatum Botanicheskago Sada – el, [P.]A.[V.] (1842) Mémoire sur plesieurs 10: 367 orchidées nouvelles ou peu connues, avec des 377. observationsMémoires de la sur Société les caractéresdu Muséum génériques. d'Historie Regel, E.[A.] (1887b) Descriptiones et Naturalle de Strassbourg – – emendationes plantarum in Horto ImperialiTrudy Zygopetal3 (9):um. 1In:35, The t. illustrated 1 5. ImperatorarskagoBotanico Petropolitano S.-Peterbur Cultarum.gsk Botaniceskago Nicholson,dictionary G. (1887)of gardening Sada – – 4. L. Gill, London. pp. 10: 685 697. Zygopetalum brachypetalum 244 248. http://dx.doi.org/10.5962/bhl.title.247Zygopetalum. In The century Regel,var. E.[A.]stenopetalum. (1888a) Gartenflora – Nicholson,supplement G. (1901) to the dictionary of gardening: 37: 385 386, t. – . L. 1277. Zygopetalum. Upcott Gill, London. Pp. 744 747. Regel,Zygopetalum E.[A.] (1888b) sanderianum. Ein Gartenflora neues http://dx.doi.org/10.5962/bhl.title.13794 37: 657, t. Ortiz Valdivieso,Orquideología P. (1994) Nuevas – especies de 1287. Index Colombia. 19 (2): 13 24. Regel,Seminum E.[A.] &1868, Max imowicz,supplement C.J.[I.] 1869 (1870) (Saint Pabst, G.F.J. (1976) AdditamentaBradea ad– orchideologiam Petersburg) brasiliensem- XXII. Hortus2: 79 90.Donatensis . Eduyarda Praca, Saint Petersburg. Planchon, J. E. (1858) . W. 34 pp. Renquet, Paris, 112 pp. Zygopetalum mackayi Reichenbach,Botanische Zeitung H.G. (Berlin) (1852a) Gartenorchideen.– Pearson,charlesworthii R.H. . (1912)The Gardeners’ Chronicle 10: 665 674. , ser. 3, Reichenbach,Botanische Zeitung H.G. (1852b) (Berlin) Gartenorchideen– V. P 51: 83. Paradisanthus10: 927 937. bahiensis. upulin, F.,- Pridgeon, A.M., Veitch, N.C., Grayer, R.J.In & Reichenbach,Xenia Orchidacea H.G. (1854). – Blanco[ Coto], M.A. (2009) Zygopetalinae. : 1: 30 31, t. 14. Pridgeon, A.M., Cribb, P.J., Chase, M.W. & http://dx.doi.org/10.5962/bhl.title.462 H eringeriana - 14(2): 157 191. 2020. 189

OrchidaceaeMeneguzzo, T.E.C. Zygopetalum intermedium .

Reichenbach, H.G. (1856) Stipulae Rolfe,var. R.A. peruvianum (1894) Lindenia Lodd. ReichenbachianaeBonplandia intra (Hannover) “folia” Lindleyana – Rolfe. 7: 71 intraaxillares. 4: 321 http://dx.doi.org/10.5962/bhl.title.64391– Bulletin 330. Orchideae Rolfe,of R.A.Miscellaneous (1895) New Informations, orchids decade Royal 14. Botanic Reichenbach, H.G. (1859) Gardens, Kew – Splitgerberianae surinamensis e Museo- 1895: 191 195. [issue published SplitgerberianoBatavo. Nederlandsch Horti Kruidkundig Academici Archief Lugduno August 1895.] – 4: http://dx.doi.org/10.2307/4114968 319 335. Orchides Zygopetalum. Rolfe, R.A. (1899)The Orchid Orchids Review at the Royal– Horticultural Reichenbach,Annales Botanices H.G. (1863a) Systematicae : (Walpers) Society. Zygopetalum7: ballii 285. The288. Gardeners’ – 6: Rolfe,Chronicle R.A. (1900) 650 662. http://dx.doi.org/10.5962/bhl.title.7556Orchides Zygosepalum. , ser. 3, 27: 149. – Bulletin Reichenbach,Annales Botanices H.G. (1863b) Systematicae : (Walpers) Rolfe,of R.A.Miscellaneous (1906) New Informations, orchids decade Royal 26. Botanic – 6: Gardens, Kew – 665 666. http://dx.doi.org/10.5962/bhl.title.7556The 1906: 30 34. Reichenbach,Gardeners’ Chronicle H.G. (1878) New garden plants. Rolhttp://dx.doi.org/10.2307/4115506 , n.s., 10: 300. The Thefe, R.A. Gardeners’ & Baker, Chronicle J.G. (1892) New garden– plants. ReichenGardeners’bach, H.G.Chronicle (1881a) New garden plants. Romero- , ser. 3, 12: 178Orchidaceae179. , n.s., 15: 298. The González, G.A. (2005) Reichenbach,Gardeners’ H.G.Chronicle (1881b) New garden plants. Schomburgkianae: the orchids collected by R. H. , n.s., 16: 134. SchomburgkHarvard in South Papers American in Botan y and the– Richard, L.[C.M.] (1792) Catalogus plantarum, ad Caribbean. 10: 231 - societatem,missarum a d ineunte anno 1792,Actes de e la Cayenna Société 268. http://dx.doi.org/10.3100/1043 d'historie naturelleomino de Le Paris Blond. – 4534(2005)10[231:OSTOCB]2.0.CO;2Bromeliaceae et Orchidaceae 1: 105 114. Ruschi, A. (1954) Bulletin du Muséum Richard, A. & Galeotti, H. (1845) Orchidographie novaeNational Espirito d’Histoire Santensis. Naturelle – Mexicaine, d’apres les echantillons, notes et , sér. 2, 26: 244 251. dessins de Annales MM. des Galeotti, Sciences Linden, Naturelles Funk, Ruschi, A. Boletim(1964) Orquidaceas do Museu de do Biologia Estado Prof. do Espı́rito Mello- Ghiesbreght.– , sér. Santo.Leitão, Botânica 3, 3: 15 33. Zygopetalum lindeniae Zygopetalum, 18: 1. rigbyanum. Boletim Rolfe,Lindenia R.A. (1891)– Rolfe. Ruschi,do Museu A. (1975) de Biologia Prof. Mello Leitão, Botânica, 6: 73 74, t. 275. [volume cover– date – 1890, issues effectively published 1890 1891, 83: 1 4. this issue 1891.] Schlechter,Orchis (Berlin) [F.R.]R. (1915a)– Kleine Mitteilungen. http://dx.doi.org/10.Zygopetalum5962/bhl.title.64391 (Warscewiczella) 9: 27 32. Rolfe,lindenii R.A. (1892) Lindenia Schlechter,- [F.R.]R. (1915b)Orchis Neue (Berlin) und seltene– Rolfe. 8: 5, t. 337. Garten– Orchideen VIII. 9: 49 54, http://dx.doi.org/10.5962/bhl.title.64391 t. 4 5. H eringeriana - 14(2): 157 191. 2020. 190 Notes on Zygopetalinae.

Aganisia

Schlechter, [F.R.]R. (1918) Die Gattung OrchisLdl. 16. Dieterichianae, Gothenburg. 936 pp. und(Berlin) ihre Verwandten,– Fortsetzung. http://dx.doi.org/10.5962/bhl.title.822Nomemclator botanicus 12: 24 42. Orchidaceae novae, in Steudel, E.T. (1840) 1. 2 ed. Schlechter, [F.R.]R. (1919) Notizblatt Cottae, Stuttgart. 468 pp. caldariisdes Botanischen Horti Dahlemensis Gartens und cultae.Museums II. zu Berlin- http://dx.doi.org/10.5962/bhl.title.655Index Herbariorum Dahlem – Thiers, B. (2020) . Available 7: 323 330. Promenaea from: http://sweetgum.nybg.org/science/ih/ Schlechter,Notizblatt [F.R.]R. (19des21) Botanischen Die Gattung Gartens und (accessed: 1 Dec 2020). Ldl.Museums zu Berlin-Dahlem – Toscano de Brito, A.L.V. (1980)– I. Bradea Species novae– in 7: 467 482. Brasiliae Orchidacearum 3: 115 117. Schlechter, F.R.R. (1922a) Orchideenfloren der Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter, W., südamerikanischenBolivia. Repertorium Specierum Kordillerenstaaten, Novarum Regni V. Hawksworth,- D.L., Herendeen,- P.S., Knapp, S., Vegetabilis, Bei Kusber, W. H., Li, D. Z., Marhold, K., May, T. W., hefte 10: 1‒80. McNeill, J., Monro, A.M., Prado,International J., Price, Code M.J. of & Schlechter, [F.R.]R. (1922b) Contribuições ao Smith,Nomenclature G.F. (eds.) for(2018) algae, fungi, and plants conhecimentoOrchidaceae Bradeanae das Orquidáceas Paulensis. do Anexos Brasil, das II. (Shenzhen Code) adopted by the Nineteenth Memórias do Instituto de Butantan, Secção de International Botanical Congress Shenzhen, Botânica China, July 2017. Regnum Vegetabile

1 (4): 1‒68, t. 1‒14. 159. Koeltz Schlechter, [F.R.]R. (1925) Repertorium Die Orchideenflora Specierum von Botanical Books, Glashütten. NovarumRio Grande Regni do Vegetabilis Sul. http://dx.doi.org/10.12705/Code.2018 , Beihefte 35: 1‒108. Whitten, W.M., Williams, N.H., Dressler, R.L., Gerlach, Schlechter, [F.R.]R. (1930) Blünanalysen neuer G.Zygopetalinae & Pupulin, F. (2005)Orchidaceae Generic relationshipsCymbidinae of Orchideen.Repertorium I: Specierum Südarikanische Novarum Orchideen. Regni ( : Lankesteriana ): Vegetabilis – combined– molecular evidence. 5: , Beihefte, 5Orchidaceae8: t. 1 60. 87 107. http://dx.doi.org/10.15517/lank.v5i2.19799- Schweinfurth,Memoirs C. (1967) of the New Yorkof the Botanical Guyana Williams, B.S. (1852) The orchid growers manual. 1 GarHighland.den – ed. London, Chapman & Hall. 108 pp. 14 (3): 69Orchideen:214. Pflanzen der Extreme, http://dx.doi.org/10.5962/bhl.title.84830 Senghas,Gegensätze K. (1993) und Superlative. Williams, L.O.Caldasia (1942) Algunas– orquidaceas Paul Parey, Berlin & americanas. 1 (5): 11 16. Hamburg, 182 pp. Woodson Junior, R.E. & Schery, R.W. (1941) Senghas,Zygopetalinae K. & Gerlach,Warreopsis G. . (1993)In Subtribus Contributions towards a Flora of Panama. ert V. : : Die Brieger, Orchideen F.G.,. Collections chiefly by Paul H. Allen and by Rob Maatsch, R. & Senghas, K. (Eds.) – E.Annals Woodson of the and Missouri Robert Botanical W. Schery Garden (Orchidac eae).– 3rd. ed., Paul Parey, Berlin,Systema pp. vegetabilium 1701 1703. 28: 409 Sprengel, C.P.J. (1826) 3. ed. 491.

H eringeriana - 14(2): 157 191. 2020. 191