On the types of Maxillaria grandiflora and Heterotaxis valenzuelana (Orchidaceae: Maxillariinae)

On the types of Maxillaria grandiflora and Heterotaxis valenzuelana(Orchidaceae: Maxillariinae)

Mario A. Blanco1,2 & Fred W. Stauffer3

Summary. The type specimen of Maxillaria grandiflora (Kunth) Lindl. was collected by Alexander von Humboldt andAimé Bonpland during their scientific expedition to Tropical America. Its type locality, which has been the subject ofconfusion, is identified as a place on the eastern part of La Cruz municipality in the province of Nariño, Colombia.The type of M. grandiflora is shown to be a mixed collection and a lectotype is designated; the protologue illustrationrepresents an amalgamation of both species. The type of Heterotaxis valenzuelana (A. Rich.) Ojeda & Carnevali wascollected by José María Valenzuela but has been erroneously cited as Wright 3314 in the recent literature.

Key Words. Charles Wright, Colombia, Cuba, Humboldt and Bonpland, José María Valenzuela.

This paper aims to clarify the confusion surroundingthe collecting locality and vegetative morphology ofthe type specimen of Maxillaria grandiflora (Kunth)Lindl., and the confusion about the type specimen ofHeterotaxis valenzuelana (A. Rich.) Ojeda & Carnevalicited in the literature.

Maxillaria grandifloraThis species was first published in 1816 as Dendrobiumgrandiflorum Kunth, based on a collection made by AiméBonpland and Alexander von Humboldt during theirfamous expedition to the New World (1799 – 1804).Although correctly identified by Lindley as a species ofMaxillaria (and transferred by him to that genus in 1832),both the type locality and the morphology of the typespecimen (and thus the circumscription of the species)have been a source of confusion up to the present day.

The information provided in the protologue forDendrobium grandiflorum is presented below:

“6. DENDROBIUM GRANDIFLORUM. † Tab. LXXXVIIID. bulbiferum; foliis lanceolatis, acutis; scapis unifloris,foliolis calycinis ovato-oblongis, acutis; lateralibus apicerevolutis.

Crescit in radicibus Andium Puruguayensium properupem El Pulpito, villam La Erre et planitiem montanamSacondonoensium, alt. 1060 hex. 4[?] Floret Octobri.

HERBA parasitica. RADICES fibrosæ, teretes, albidæ,Bulbus ovato-oblongus, compressus, fuscescens, nitidus,subtripollicaris. FOLIA lanceolata, acuta, plana, basivalde angustata et carinata, striato-nervosa, subcoriacea,glabra, pedalia, sesquipollicem lata. SCAPI sexpollicares,erecti, glabri, spathellis pluribus vestiti, uniflori.SPATHELLÆ membranaceæ, acuminatæ, apice laxiusculæ,striatæ, pollicares et longiores. CALYX patulus, albus,glaber; foliola carnosa, striato nervosa; exteriora ovato-oblonga, acuta, pollicaria; superius erectum, concavum;lateralia apice revoluta, basi ungui labelli adnata; duointeriora exterioribus duplo breviora, erecta, oblonga,acuta, apice oblique revoluta. LABELLUM unguiculatum;unguis carnosus, cum foliolis calycinis exterioribuslateralibus connatus; lamina semipollicaris, ovata,obtusa, carinato-concava, margine undulata, rubescens,unguem versus transversaliter plicata. OVARIUM costa-tum, glabrum. GYNOSTEMUM arcuatum, foliolis interior-ibus duplo brevius, canaliculatum, apice triangulare,interius rubro-punctatum. ANTHERA terminalis; ejusstructura mihi tamen ignota. CAPSULA haud visa.”

Type localityThe route followed by Humboldt and Bonplandduring their expedition through the Americas is wellknown: Venezuela, Colombia, Ecuador and Peru werethe only South American countries visited (Stearn1968; Lack 2009). Despite this, some authors misinter-preted the phrase Andium Puruguayensium in the

Accepted for publication January 2012.1 Department of Biology, University of Florida, 214 Bartram Hall, Gainesville, FL 32611-8526, USA.2 Present address: Escuela de Biología, Universidad de Costa Rica, 11501-2060, San José, Costa Rica. e-mail: [email protected] Herbarium, Conservatoire et Jardin botaniques de la Ville de Genève, Université de Genève, Laboratoire de Systématique Végétale et Biodiversité,

case postale 60, CH-1292 Chambésy/Genève, Switzerland. e-mail: [email protected]

KEW BULLETIN VOL. 66: 529Y536 (2011)

© The Board of Trustees of the Royal Botanic Gardens, Kew, 2012

protologue of Dendrobium grandiflorum as “the Andes ofParaguay.” For example, while referring to Maxillariagrandiflora, Veitch (1893: 154 – 155) stated the follow-ing: “As no such mountains as the Andes of Paraguayare known in modern geography, and the other placesmentioned are not found on any map to which we haveaccess, the habitat given by Humboldt is virtuallyvalueless […] If we assume the word Puruguayensium ofHumboldt and Kunth to be a clerical error forPeruvianorum, the habitat given by them can to a greatextent be reconciled with that given by [ÉdouardFrançois] André” [who received plants from Perucollected by GustaveWallis, supposedly ofM. grandiflora].

Several horticultural and floristic works propagatedthis error, citing the country of origin as Paraguay, orsimply stating that the species was collected in Peru(e.g., Watson & Chapman 1903; Sander et al. 1927;Foldats 1970; King 1984). Humboldt and Bonplandnever visited Paraguay together. Bonpland alone spentten years (1821 – 1831) as a prisoner in Paraguay(Lourteig 1977), but this was well after the publicationof Dendrobium grandiflorum. Moreover, there are nospecies of Maxillaria known from Paraguay.

Herman (1977: 137), without citing any evidence,asserted that Humboldt and Bonpland collected thetype of Dendrobium grandiflorum along their way fromCañar to Loja (in Ecuador). That leg of their tripoccurred during July and August of 1802 (Sandwith1926). Herman’s assumption can also be refuted, aswe demonstrate below.

The protologue of Dendrobium grandiflorum does notspecify the year in which the plant was collected, but itstates that it was done in the month of October. Thecollection number (2113) indicates that the year ofcollection must have been 1801 (see Table 1 in Lack2004). Humboldt and Bonpland travelled throughpresent-day Colombia on their way to Quito fromSeptember to December 1801 (Sprague 1926). Threelocality names mentioned in the protologue (Páramode Puruguay, Hacienda La Erre and Sacandanoy),although not featured on modern maps, are listed bySprague (1926) as being part of the Cordillera de Pasto,which the collectors crossed at the end of Novemberand early December (they left Popayán on their way toPasto on November 29, 1801). The protologue statesthat the plant was collected in October, but other plantswith numbers near 2113 were collected in the samegeneral area (e.g., Besleria bicolor Kunth; type: Bonpland& Humboldt 2118; Isochilus graminifolius Kunth; type:Bonpland & Humboldt 2115; both at P-Bonp).

By comparing the place names in the protologueand in Sprague (1926) with maps of southernColombia, we conclude that the type locality ofMaxillaria grandiflora must be somewhere in theeastern part of La Cruz municipality in the depart-ment of Nariño (c. 1°35'N, 76°54'W), or possibly on its

border with the department of Cauca. This area isincluded in the recently created Doña Juana-CascabelNational Park. There is a prominent, isolated mountcalled Cerro Púlpito (2,750 m, 1°38'45"N, 76°59'25"W,datum: WGS84) in the northwestern part of La Cruzmunicipality. This mount is visible from the highermountains of the cordillera to the east, where webelieve the plant was collected.

Humboldt relied on his barometers to estimate theelevation of localities in the mountains; he recordedthe elevation in toises or “hexapoda” (abbreviated“hex.” or “hexap.” in Kunth’s descriptions). One toiseor hexapodarum roughly corresponds to 1.95 m(Stearn 1968). Thus, the 1,060 hexapoda indicatedfor the type collection of Maxillaria grandiflora corre-sponds to roughly 2,067 m.

Type collectionThe type collection of Maxillaria grandiflora consists ofthree sheets in the Bonpland herbarium in the MuséumNational d’Histoire Naturelle in Paris (P-Bonp). Noduplicates exist in the Willdenow herbarium inBerlin (B-Willd).

The first sheet (Fig. 1A) has a plant specimenconsisting of a young pseudobulb with a single apicalleaf (folded on top of the petiole and part of thepseudobulb), a new bud developing at its base, andtwo inflorescence peduncles, one of them with adetached flower placed near the apex (the otherinflorescence lacks a flower). On its lower left corner,this sheet has a label with the annotations “Dendrobiumgrandiflorum” and “numr. 2113” handwritten by Kunth.

The second sheet (Fig. 1B) has a single pseudobulbwith two apical leaves and no flowers nor inflorescen-ces. The third sheet has a line drawing; the illustrationthat appeared in the protologue (Fig. 1C) is a mirrorimage of this drawing. The two last sheets do not havelabels attached to them (other than the printed ticket“HERB. MUS. PARIS, Herbier Humboldt & Bonpland.AMÉRIQUE ÉQUATORIALE”), and none have herbariumaccession numbers.

This is, however, a mixed collection. Upon closeexamination, it is clear that the elements in the firstand second sheets do not belong to the same species.This is not necessarily obvious from the images ofthese specimens in the microfiche of the Bonplandherbarium in Paris (Inter Documentation CompanyAG 1972). As mentioned before, the pseudobulb onthe first sheet has a single apical leaf while the one onthe second sheet has two.

Upon drying, the plant mounted on the first sheet(Fig. 1A) turned dark olive green and the flowerturned almost black, a characteristic feature of speciesin the “Maxillaria platypetala Ruiz & Pav. alliance”(sensu Whitten et al. 2007, which includes all thespecies in the “Maxillaria grandiflora complex” sensu

530 KEW BULLETIN VOL. 66(4)


Fig. 1. Type specimens and illustration of Dendrobium grandiflorum (A – C) and Pleurothallis valenzuelana (D). A first sheet of thetype collection of D. grandiflorum (Bonpland & Humboldt 2113, P-Bonp); B second sheet; C illustration published as part of theprotologue (“Tab. LXXXVIII”), DRAWN BY P. J. F. TURPIN; D holotype of P. valenzuelana (Valenzuela s.n. [or 23?], P). Pen length in A andB = 14 cm.

531ON THE TYPES OF MAXILLARIA GRANDIFLORA AND HETEROTAXIS VALENZUELANA


Horich 1988; Christenson 2002a, b). This pseudobulbis oblong, its surface has a matt texture, and the leafabscission layer is raised on an apical extension that isvisibly narrower than the rest of the pseudobulb (aphyllopodium, sensu Whitten et al. 2007).

In contrast, the leaves of the plant mounted on thesecond sheet (Fig. 1B) turned a lighter shade ofyellowish grey, while the pseudobulb turned brownish-orange. Even more importantly, the shape of thispseudobulb is ovate, its surface has a thick, shinytexture and the leaves are sessile on the apex of thepseudobulb. This plant is most likely a species ofCyrtochilum Kunth, Odontoglossum Kunth or OncidiumSw. (Orchidaceae: Oncidiinae). In fact, this pseudo-bulb is quite similar to those of the type collection ofOncidium pictum Kunth (type: Colombia, Cauca,Popayán, Bonpland & Humboldt s.n., P-Bonp); Lack(2009: plate 32) published a copy of the originalcoloured illustration of that species. This pseudobulbappears to have been yanked from the rest of the plant(as suggested by the torn base with no trace of therhizome), and lost the subtending foliaceous sheaths(characteristic of members of the above mentionedgenera) in the process. It probably belonged to a plantthat grew in close proximity to the Maxillaria, or itcould have been a loose piece from a differentcollection that was erroneously added to the M.grandiflora specimen when the collectors pressed theplants at the end of the day, or when the dryspecimens were sorted at a later date.

The leaves present in both herbarium sheets havesimilar shapes and dimensions, which probably con-tributed to the confusion.

Protologue illustrationThe illustration of Dendrobium grandiflorum presentedin the protologue (“Tab. LXXXVIII”, Fig. 1C) wasprepared by Pierre Jean François Turpin (1775 – 1840;Stafleu & Cowan 1979; Lack 2009) from the driedspecimens. The original drawing mounted on thethird sheet at P-Bonp. has a tiny annotation near thelower left corner that reads “Turpin Del.” (délinée,drawn).

The illustration aimed to recreate the appearanceof the plant in life, and represents an amalgamation ofthe two discordant elements of the mixed collection.The pseudobulb and leaves correspond to the plantmounted on the second sheet (Fig. 1B), but theflower, inflorescences, developing new growth, andeven the bracts at the base of the pseudobulbcorrespond to the plant mounted on the first sheet(Fig. 1C).

Field notesOne of us (FWS) has been able to study the entry for thecollection number 2113 in Humboldt and Bonpland’s

botanical field notes conserved in the BibliothèqueCentrale of the Muséum National d’Histoire Naturellein Paris (the so-called “Journal Botanique,” shelfmark“MS 2534”; see Lack 2004).

Entry 2113 was written by Humboldt, and isassociated with the name “Epidendrum roseum.” However,this combination was never published by Humboldt,Bonpland nor Kunth; it was validly published bySchlechter (1913: 497) for an unrelated species. Someother person, probably Kunth, later added the anno-tation “Epidendrum grandiflorum;” this name was even-tually published by Bonpland (in Humboldt &Bonpland 1808: 94 – 97, t. 27) for a different species,now called Stanhopea jenischiana Kramer ex Rchb. f. Astypical of other field notes of Humboldt, the descrip-tion is quite long and complicated, and has lateradditions and corrections by Humboldt himself. Thelocality “Montaña de Puruguay”, however, is expresslyindicated. The third paragraph of Kunth’s descriptionin the protologue was evidently based on Humboldt’snotes.

This description does not mention whether thepseudobulbs had one or two apical leaves. Thecollectors, being unfamiliar with this group of orchids,probably did not regard this as a taxonomicallyimportant character.

Humboldt normally included very detailed linedrawings of the flowers in the descriptions of orchidshe made in the Journal Botanique (F. W. Stauffer, pers.obs.). Unfortunately, this is not the case for the entrynumber 2113.

Taxonomic implicationsFor most orchids with pseudobulbs, the number ofapical leaves per pseudobulb is a character of taxo-nomic importance. Thus, the existence of a pseudo-bulb with two apical leaves as part of the typecollection of Maxillaria grandiflora has been a sourceof confusion, leading some authors (e.g. Christenson2002a,b, 2009) to conclude erroneously that thisspecies is not a member of the M. platypetala Ruiz &Pav. alliance (which includes M. augustae-victoriae F.Lehm. & Kraenzl., M. elegantula Rolfe, M. fletcherianaRolfe, M. fucata Rchb. f., M. huebschii Rchb. f., M.irrorata Rchb. f., M. lehmannii Rchb. f., M. molitor Rchb.f., M. sanderiana Rchb. f., M. striata Rolfe, amongothers; see Whitten et al. 2007). Phylogenetic analysesbased on various DNA regions (Whitten et al. 2007)indicate that these species are closely related to eachother; the relatively low genetic differentiation amongthe members of this group suggests that these speciesare the product of a recent radiation.

Many species in this alliance are superficiallysimilar to each other (see Teuscher 1959; King 1984;Dunsterville 1984; Horich 1988; Withner 1996), butcan be easily distinguished when live plants are



compared side by side. Some of the distinguishingfeatures, especially colour patterns, disappear or areobscured in dried herbarium specimens; the flowerstend to become dark brown to almost black when dry,regardless of their original colour. Christenson (2002a,b) listed diagnostic features for several species in thiscomplex, but the entire group (theMaxillaria platypetalaalliance sensu Whitten et al. 2007) still awaits a thoroughtaxonomic revision.

Because the type collection of Maxillaria grandifloraconsists of plant parts that belong to two differenttaxa, a lectotype is chosen below so that it correspondsmore closely to the original description (as perArticles 9.2 and 9.12 of McNeill et al. 2006). Data inbrackets are inferred by us (see above).

Maxillaria grandiflora (Kunth) Lindl. (Lindley 1832:147). Type: [Colombia, Nariño, La Cruz], foot of thePuruguayan Andes near Mt El Púlpito, La Erre ranchand the plains of Sacandonoy [Sacandanoy?], 1,060hex. [c. 2,067 m], Oct. [most likely Dec. 1801], Bonpland& Humboldt 2113 (P-BONP!, mixed collection of twosheets and a third sheet with the original illustration;lectotype selected here: sheet with flowering specimen,photograph of which is presented in Fig. 1A); Dendrobiumgrandiflorum Kunth (1816: 359, Tab. LXXXVIII);Broughtonia grandiflora (Kunth) Spreng. (Sprengel1826: 735); Lycaste grandiflora (Kunth) Beer (1854:265); Callista grandiflora (Kunth) Kuntze (1891: 654).

No attempt is made to assign heterotypic synonyms atthis point. We believe that Maxillaria eburnea Lindl.(type: Schomburgk s.n., K-Lindl.!), frequently assignedto the synonymy of M. grandiflora, is a different,closely-related species (see Sauvêtre 2010).

The specimens identified by Lindley as “Dendrobiumgrandiflorum H.B.K.” or Maxillaria grandiflora in hisown herbarium (Jameson 21, Linden 631, Spruce 5103and Warscewicz s.n., K-Lindl!) do not correspond to thisspecies either. Jameson 21 and Spruce 5103, bothmounted on the same sheet (with an additional sheetfor another part of Spruce 5103), were collected inEcuador; their label notes indicate that their flowerswere orange to pale yellow, and probably representMaxillaria molitor. Linden 631 was collected in Vene-zuela and matches the drawing erroneously identifiedas M. grandiflora by Dunsterville & Garay (1972, 1979)and Dunsterville (1984); this same drawing was lateridentified asM. augustae-victoriae by Romero & Carnevali(2000). The Warscewicz specimen (collected in Peruaccording to Christenson 2002a, b), which consists of asingle flower and a crude sketch of the plant, isprobably M. platypetala.

The flower in the type specimen of Maxillariagrandiflora has a labellum that is subequal in length

to the column and is markedly curved toward theabaxial side (not apparent in the original illustration,Fig. 1C). The morphology and size of the differentfloral parts agree well with the illustrations of M.grandiflora presented by Hoehne (1953) and Dodson& Dodson (1982; also reproduced in Dodson 2002:554). Dodson & Dodson (1982) note that M.grandiflora appears to be a clearly distinct species,with relatively low variability between individuals andpopulations.

Heterotaxis valenzuelanaThis species was described in 1850 as Pleurothallisvalenzuelana A. Rich. Some recent treatments (e.g.,Atwood 1989, 1999; Nir 2000; Ojeda et al. 2005) citethe type information as “Cuba, Valenzuela, Wright3314 (holotype P?).” However, the publication ofPleurothallis valenzuelana predates the beginning ofCharles Wright’s fieldwork in Cuba by six years(Howard 1988).

It seems that the authors cited above misunder-stood “Valenzuela” to mean a locality in Cuba, butthere are no Cuban localities with that name accord-ing to Gannett (1902). The GeoNames website(http://www.geonames.org) lists a locality namedValenzuela in the province of Granma; however, thisis not the type locality either, because the protologueexplicitly cites the locality of Vuelta de Abajo, and J. M.Valenzuela as the collector.

The poorly known José María Valenzuela collectedplants in Vuelta de Abajo (Pinar del Río province,municipality of Viñales; an area south of the Sierra de losOrganos, c. 22°37'N, 83°43'W; Gannett 1902: 10) inOctober of 1833 (Urban 1898: 143; Vegter 1988). Richard(1850) cited Valenzuela as the collector in several of thespecies he described as new in Ramón de la Sagra’sHistoria Física, Política y Natural de la Isla de Cuba.

The type specimen (Fig. 1D) is deposited in thegeneral vascular plant collection at the MuséumNational d’Histoire Naturelle in Paris (P). There is asmall, square piece of paper with the number 23attached to the base of the plant. On the lower leftcorner there is a handwritten label with the annota-tion “Pleurothallis Valenzuelana Nob. (Achille Richardscrip.) Cuba. J. M. Valenzuela,” all apparently inRichard’s hand. There is a second small piece ofpaper glued to this label with the number 91 (or 21?)written by a different hand. Above this label, H. G.Reichenbach annotated the specimen as “Maxillariairidifolia Rchb. f.” It is possible that 23 is the collectionor species number assigned by Valenzuela, but none ofthese numbers (23, 91 or 21) was mentioned in theprotologue.

On the lower right corner there is the standardprinted label “Herbier de LOUIS CLAUDE et d’Achille



http://www.geonames.org

RICHARD,” with the annotation “Tab. 98” written on it.This is perplexing, as no figure 98 was published (noreven mentioned) in de la Sagra’s Historia volume 12(Sagra 1855), which included all plant illustrations. Itmight have been one of the parts reported by de laSagra to have been lost or stolen at the printer (seeAguayo 1946).

The correct citation of the type specimen is asfollows. The data in brackets are inferred by us.

Heterotaxis valenzuelana (A. Rich.) Ojeda & Carnevali(in Ojeda et al. 2005: 581). Type: Cuba, [Pinar del Río,Viñales], Vuelta de Abajo, [Oct. 1833], J. M. Valenzuelas.n. [or 23?] (holotype P no. 463348!; isotype W-Reich-Orch. no. 50132); Pleurothallis valenzuelana A. Rich.(Richard 1850: 234 – 235); Maxillaria valenzuelana (A.Rich.) Nash (1907: 121); Marsupiaria valenzuelana (A.Rich.) Garay (1952: 183); Heterotaxis valenzuelana (A.Rich.) F. Barros (2005: 427).

Ojeda et al. (2005) provide a complete heterotypicsynonymy.

It should be noted that the numbers associated withCharles Wright’s specimens correspond to species, not toindividual gatherings; this was caused in part by AsaGray’spractice of replacing Wright’s field numbers with his own“distribution” numbers (Howard 1988; Gottschling &Miller 2007). Thus, two or more of Wright’s specimenswith the same number are not necessarily duplicates, asdefined by McNeill et al. (2006).

The first association of Wright 3314 with the namePleurothallis valenzuelana can be traced to Grisebach(1866), who identified the specimen as Maxillariairidifolia Rchb. f. (type: Poeppig s.n. (W-Reich.-Orch)).Grisebach merely listed P. valenzuelana as a synonym ofM. iridifolia, but the succinct text could have beenmisinterpreted by some authors as a type designation.For example, Cogniaux (1909 – 1910: 606) mentionedcollections by Poeppig, R. de la Sagra (probably aconfusion with Valenzuela’s specimen cited byRichard) and Wright 3314 for M. valenzuelana, butfailed to mention Valenzuela’s name.

A similar situation has occurred with Vernoniavalenzuelana A. Rich. (Compositae), whose type isValenzuela s.n. (P) but has been wrongly cited as Wright2785 (e.g., by the W3TROPICOS database). Misled bythe erroneous type citation, one of us (MAB)annotated specimens of Wright 3314 as isotypes ofHeterotaxis valenzuelana in several herbaria (e.g., G, K,MO, P); these annotations should be disregarded.

It is notable that Lindley (1858) did not mentionPleurothallis valenzuelana (under this or any othergenus) in his list of orchids collected by CharlesWright in eastern Cuba, despite his assertion that hereceived a complete set of them from Asa Gray

(Lindley 1858: 325). In fact, there are no “dupli-cates” of Wright 3314, nor any specimen of thisspecies, in Lindley’s personal herbarium at Kew.Lindley would have probably included this speciesunder the genus Dicrypta Lindl., which he favouredagainst his own earlier genus Heterotaxis Lindl. (Ojedaet al. 2005).

AcknowledgementsWe thank the curators and personnel of P and K fortheir attention and assistance during our visits. Fund-ing from a Kew-Latin American Research Fellowshipallowed MAB to visit these and various other Europeanherbaria to study types of subtribe Maxillariinae(Orchidaceae). MAB thanks James Ackerman (UPRRP)for discussions about Charles Wright. FWS would liketo thank Pascale Heurtel (Bibliothèque Centrale duMuséum National d’Histoire Naturelle, Paris) forkind access to the Journal Botanique of Humboldtand Bonpland. The present study is part of the project"Systematics of Maxillariinae (Orchidaceae): Genericdelimitation, pollinator rewards, and pollination,"supported by a grant (No. DEB-0234064) from the U.S.National Science Foundation to Norris H. Williams andW. Mark Whitten (FLAS).

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On the types of Maxillaria grandiflora and Heterotaxis valenzuelana (Orchidaceae: Maxillariinae)

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