Chimaeras and ghosts: solving a chimaeric specimen
and two neglected orchid names
Authors: Meneguzzo, Thiago E. C., and Den Berg, Cássio Van
Source: Willdenowia, 50(1) : 139-146
Published By: Botanic Garden and Botanical Museum Berlin (BGBM)
URL: https://doi.org/10.3372/wi.50.50113
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Willdenowia
Annals of the Botanic Garden and Botanical Museum Berlin
THIAGO E. C. MENEGUZZO1* & CÁSSIO VAN DEN BERG2
Chimaeras and ghosts: solving a chimaeric specimen and two neglected orchid names
Version of record first published online on 27 March 2020 ahead of inclusion in April 2020 issue.
Abstract: We present nomenclatural and taxonomic studies of selected species and names of Neotropical Orchidaceae
currently placed in Bifrenaria, Cleistes, and Pleurothallis, but once classified in ten different genera. Several nomenclatural and taxonomic actions are proposed, including changes in nomenclatural status, typifications, and taxonomic
rearrangements by indication of the correct name to be used, re-evaluation of previously proposed synonyms, and
new synonyms. The accepted names remaining after the study are: Cleistes rosea Lindl. f. rosea (relevant synonyms:
C. angeliana Campacci, C. castaneoides Hoehne, and Epistephium monanthum Poepp. & Endl.); Cleistes rosea f.
augusta (Hoehne) Meneguzzo & Van den Berg, comb. nov. (for Pogonia rosea var. augusta Hoehne); Cleistes speciosa Gardner [relevant synonyms: C. caloptera Rchb. f. & Warm., C. metallina (Barb. Rodr.) Schltr., and C. monantha
(Barb. Rodr.) Schltr.]; Bifrenaria harrisoniae (Hook.) Rchb. f. (for Maxillaria spathacea Lindl.); and Pleurothallis
quadrifida (Lex.) Lindl. [for the homotypic pair Gomesa stricta Spreng. and Rodriguezia stricta (Spreng.) Steud.].
Key words: Bifrenaria, biodiversity, botanical literature, Cleistes, Neotropics, nomenclature, Orchidaceae, orchids,
Pleurothallis, typification
Article history: Received 2 October 2019; peer-review completed 22 January 2020; received in revised form 30
January 2020; accepted for publication 3 February 2020.
Citation: Meneguzzo T. E. C. & van den Berg C. 2020: Chimaeras and ghosts: solving a chimaeric specimen and two
neglected orchid names. – Willdenowia 50: 139 – 146. doi: https://doi.org/10.3372/wi.50.50113
Introduction
The investigation of taxonomic and nomenclatural issues
may take considerable time, not only for the researcher to
feel confident about its results, but also to get acquainted
with the extensive existing botanical literature and the
intricate rules that govern botanical nomenclature. To
obtain reasonable expertise on the biology of the study
group itself, to understand its diversity and variability, is
also time consuming. How former botanists found and
interpreted the organisms, what their working and publishing methods were, and to what extent subsequent
researchers and users perpetuated or rejected such re-
sults is almost a study in itself. It is not rare to find taxonomic and nomenclatural problems that have persisted
for several years before being satisfactorily investigated
and resolved. The degree of complexity of some cases
directly reflects the availability of original literature and
herbarium specimens, which until recently were accessible only through personal visits to scattered collections
distant from home institutions, but which are now gradually becoming available on the internet.
Over the course of our research for the REFLORA
project (a digitizing programme for specimens of Brazilian plants at the Royal Botanical Gardens, Kew and
Muséum national d’Historie Naturelle, Paris, from 2012
1 Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Diretoria de Pesquisa, Rua Pacheco Leão 915, 22460-030, Rio de
Janeiro, Rio de Janeiro, Brazil; *e-mail: botanica@meneguzzo.net.br (author for correspondence).
2 Universidade Estadual de Feira de Santana, Departamento de Ciências Biológicas, Av. Transnordestina s.n., 44036-900, Feira de
Santana, Bahia, Brazil.
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140
Meneguzzo & van den Berg: Solving a chimaeric specimen and two neglected orchid names
to 2014) we found some complex taxonomic and nomenclatural issues that deserved attention. Here we present
our results in three cases concerning names and species
currently classified in Bifrenaria Lindl., Cleistes Rich. ex
Lindl., and Pleurothallis R. Br. These names were once
described or combined in ten different genera, as follows:
Cattleya Lindl., Epistephium Kunth, Gomesa R. Br.,
Humboltia Ruiz & Pav., nom. rej. (non Humboldtia Vahl,
nom. cons.), Lalexia Luer, Loddigesia Luer, nom. illeg.
(non Sims), Pogonia Juss., Rodriguezia Ruiz & Pav.,
Specklinia Lindl., and Stelis Sw.
=
=
Material and methods
This study follows standard methods for alpha taxonomy.
It is based on material examined since 2008 from 98 collections in 86 herbaria, as follows (herbarium codes are
according to Thiers 2020+; italics indicate herbaria personally visited): A (includes AMES, GH), AAU, ALCB,
ARIZ, B, BHCB, BM, BR, BRIT, C, CEN, CEPEC,
CESJ, COAH, COL, CR, CRI, CTES, CUZ, CVRD,
E, ESA, F, FUEL, G (includes G-DC), GENT, GOET,
HAL, HB, HBG, HBR, HEPH, HERBAM, HJ, HPUJ,
HRB, HTO, HUA, HUEFS, HUEM, HUFU, HUTU,
IAN, IBGE, ICN, INPA, JAUM, K (includes K-L), L (includes U, WAG), LE, LPB, M, MBM, MBML, MEXU,
MG, MPU, MO, MOL, NDG, NY, OXF, P, PACA, PH, R,
RB (includes GFJP, GUA, ITA, RUSU), RENZ, S, SBT,
SEL, SP, SPF, TO, UB, UEC, UFMT, UFP, UPCB, UPS
(includes UPS-THUNB), US, USM, VEN, VIES, W (includes W-R), and WU. Nomenclature follows Turland &
al. (2018). Lectotype or neotype designations are made
whenever necessary, especially in cases when we found
no internal or external evidence of an element among
the original material that could be the holotype (McNeill
2014; Turland & al. 2018: Rec. 9A.1). Specimens are
cited with the herbarium code (e.g. K) followed by the
barcode where available (e.g. K000079712); a character
string following a herbarium code is a barcode unless indicated otherwise (e.g. “accession no.”).
Epistephium monanthum and Barbosa Rodrigues’s
and Schlechter’s Pogonia monantha homonyms
Cleistes rosea Lindl., Gen. Sp. Orchid. Pl.: 410. 1840 [f.
rosea] ≡ Pogonia rosea (Lindl.) Rchb. f., Xenia Orchid.
2: 89. 1865 ≡ Pogonia rosea (Lindl.) Hemsl., Biol.
Centr.-Amer., Bot. 3: 304. 1884, later isonym. – Lectotype (designated here): Guyana, “Savannahs adjacent to
the lake Capooey, Arabisee Coast of Essequibo” [from
protologue], 1836 – 1837 [see Romero-González 2005:
239], R. H. Schomburgk s.n. (K K000079712!; isolectotype: P P00367101!).
= Epistephium monanthum Poepp. & Endl., Nov. Gen.
Sp. Pl. 1: 53. 1836 ≡ Pogonia monantha (Poepp. &
Endl.) Schltr. in Repert. Spec. Nov. Regni Veg. Beih.
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=
=
=
9: 121. 1921, nom. illeg. [non Pogonia monantha
Barb. Rodr., Gen. Spec. Orchid. 1: 167. 1877]. – Lectotype (designated here): Peru, Loreto, Huánuco,
between Cuchero and Chihuamecala, Dec 1829, E. F.
Poeppig 1601 B (W accession no. 7715!).
Pogonia venusta Schltr. in Repert. Spec. Nov. Regni
Veg. Beih. 7: 42. 1920 ≡ Cleistes venusta (Schltr.)
Schltr. in Arch. Bot. São Paulo 1: 180. 1926. – Holotype: Cauca, c. 1500 m, s.d., M. Madero s.n. (B destroyed). – Neotype (designated here): Colombia,
Valle del Cauca, near Buenaventura, 24 May 1939, A.
H. G. Alston 8620 (P P00441796!).
Cleistes castaneoides Hoehne in Arq. Bot. Estado
São Paulo 1: 42. 1939. – Lectotype (designated
here): Brazil, São Paulo, São Paulo, Indianópolis, ao
lado da estrada para Santo Amaro, 14 Jan 1932, F. C.
Hoehne s.n. (SP SP001994 [accession no. 28697]!;
isolectotypes: GH 00056707 [AMES accession no.
17832]!, NY 00414849!, US 00093314 [accession
no. 2790601]!). – Syntype: Brazil, São Paulo, São
Paulo, Ipiranga [“Ypiranga”], 27 Jan 1908, H. Luederwaldt s.n. (SP SP001995 [accession no. 28995]!).
Cleistes latiglossa Hoehne in Arq. Bot. Estado São
Paulo 1: 42. 1939 ≡ Cleistes latiplumis Hoehne in
Orquídea (Rio de Janeiro) 2: 113. 1940, as “latiplume”, nom. illeg. superfl. – Lectotype (designated
here): Brazil, Mato Grosso, Salto Augusto, River
Juruena, Feb 1912, F. C. Hoehne sub Commissão
Rondon 5328 (SP SP001999 [accession no. 29001]!).
– Syntype: ibidem, F. C. Hoehne sub Commissão
Rondon 5329 (R accession no. 3220!).
Cleistes rosea f. pallida Carnevali & I. Ramírez in
Ann. Missouri Bot. Gard. 77: 551. 1990. – Holotype:
Venezuela, Amazonas, Átures, palm bog 1 – 2 km E
of Piedra Tortuga, 8 km S of Puerto Ayarucho, 30
Jun 1988, G. Carnevali, I. Ramírez & G. A. Romero
2661 (VEN accession no. 231152!; isotypes: AMES
not found, INPA not found, MO not found, MY not
found, PORT accession no. 30669!, TFAV not found).
Cleistes angeliana Campacci, Colet. Orquídeas Brasil. 9: 324. 2011, syn. nov. – Holotype: Brazil, Minas
Gerais, Diamantina, Prata River, Jan 2010, E. Menezes EM-001 (ESA accession no. 114454!).
Remarks — The tribe Pogonieae contains five well-defined genera according to both morphological and molecular characters (Chase & al. 2015). Historically, names
in Cleistes, Epistephium, and Pogonia were often recombined in these genera. Cleistes was considered a section
of Pogonia by Ames (1922). Schlechter (in Schlechter &
Hoehne 1926) re-established Cleistes as an autonomous
genus. A treatment of names in both Cleistes and Pogonia
is necessary in any taxonomic revision of this group, as it
is crucial to establish which names belong in which genus.
The initial motivation for investigating this nomenclatural imbroglio started in late 2012 while we studied the collections at the Naturhistorisches Museum
Willdenowia 50 – 2020
Wien (herbarium W), and found the original specimen
of Epistephium monanthum collected in Peru by Eduard Friedrich Poeppig, and described by Poeppig &
Endlicher (1836: 53). It immediately reminded us of a
comment by Hoehne (1945) in the revision of Brazilian
species of Epistephium that he did not have the opportunity to study this specimen. On revising E. monanthum,
we concluded that it is an earlier heterotypic name for
Cleistes rosea, along with other synonyms. And on revising Pogonia monantha Barb. Rodr., we established that it
is a new synonym of C. speciosa Gardner together with
several other names.
The clarification of the taxonomy of Epistephium
monanthum was probably hindered by the fact that most
botanists did not to have the opportunity to study its
original specimen. Lindley (1840) kept the name for an
accepted species as originally proposed. Reichenbach
(1861 – 1864) neglected it in the treatment for Cleistes,
Epistephium, and Pogonia. Shortly after, when he summarized and commented on Pogonia species (Reichenbach 1865), he made no mention of E. monanthum.
Cogniaux (1893) presented some progress by expressing
doubt on its classification, and suggesting it belonged
to Pogonia. Latter, Hoehne (1945) kept the same doubt
because he was not able to draw any conclusion on its
taxonomic status in relation to the Brazilian species,
solely analysing the protologue, which contained only a
crude illustration. Despite this, he pointed out that the
name would correspond instead to a species in Cleistes.
Schweinfurth (1958) simply treated it as an Epistephium.
Finally, Garay (1978) studied the original specimen and
published his finding that E. monanthum was a synonym
of C. rosea. Garay (1978) additionally cited two synonyms: P. lenheirensis Barb. Rodr. and P. venusta. We
checked the matter in Pansarin’s (2005) unpublished PhD
thesis (widely available on the internet) on the systematics of Cleistes. Surprisingly, neither E. monanthum nor
its combination P. monantha (Poepp. & Endl.) Schltr.
was cited in it. Additionally, we found a combination
by Schlechter that is a later homonym of a new species
name published by Barbosa Rodrigues 44 years earlier.
The names we are dealing with here have been omitted in
Pansarin’s work (Pansarin 2005).
Regarding Garay’s (1978) list of synonyms, we agree
that Epistephium monanthum and Pogonia venusta are
heterotypic synonyms of Cleistes rosea. These two names
were not cited by Pansarin (2005). On the other hand, we
disagree that P. lenheirensis is conspecific with C. rosea.
We concur with Pansarin (2005) that P. lenheirensis is
conspecific with C. metallina, but in the present study the
latter name is proposed as a new synonym of C. speciosa
Gardner. Pansarin (2005) pointed out that C. speciosa (as
C. metallina) and C. rosea share similarities, but can easily be distinguished because the distal leaf is smaller and
the lip slightly trilobed on the former, whereas the distal
leaf is considerable larger and the lip entire in the latter.
The conspecificity of C. rosea with C. castaneoides, C.
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141
latiglossa, and C. rosea f. pallida, synonyms proposed
by Pansarin (2005), is followed here. To this synonymy
we add C. angeliana, which was not compared with other
species in its protologue, but our study of the protologue
and the holotype showed that it does not differ from C.
rosea.
Pogonia monantha (Poepp. & Endl.) Schltr. is an
illegitimate name because it is a later homonym of P.
monantha Barb. Rodr. (Turland & al. 2018: Art. 53.1),
and P. rosea (Lindl.) Hemsl. may be disregarded because
it is a latter isonym of P. rosea (Lindl.) Rchb. f. (Art. 6
Note 2). A neotype for P. venusta is designated here because the type at the herbarium B was destroyed in 1943.
An illustration of the name was posthumously published
by Schlechter (1929: t. 10, fig. 38). However, in that publication there was no indication that it is part of the original material that could potentially be designated as the
lectotype (problem extensively discussed by Meneguzzo
& al. 2013). In this case we designate as the neotype a
complete specimen instead of an illustration solely consisting of a dissected flower. Cleistes latiplumis is an illegitimate superfluous name of C. latiglossa because the
same type specimen is cited in protologue of both names
(Art. 51.1 and 51.2). Cleistes castaneoides has been most
frequently cited in the literature in its misspelled form
“C. castanoides”. The protologue of C. latiglossa cited
P. rosea var. augusta as a synonym, but because a name
has no priority outside its published rank (Art. 11.2), the
former name is not made superfluous and illegitimate by
that synonymy.
Finally, even though the name Epistephium monanthum is conspecific with Cleistes rosea and was published
four years earlier than the latter, it cannot be used in the
genus Cleistes because the specific epithet is already occupied by the heterotypic name C. monantha (Barb. Rodr.)
Schltr., which is hereafter synonymized under C. speciosa.
Additional specimens examined — Brazil: Distrito
Federal: Poço Azul, 23 Dec 2008, T. E. C. Meneguzzo
& al. 44 (UB). Goiás: Serra Dourada, 29 Jan 1966, E.
P. Heringer 10938 (HB, K, UB). Mato Grosso: Ribeirão
Cascalheira, 10 Jan 1968, D. Philcox & A. Ferreira 3957
(K, UB). Minas Gerais: Uberlândia, 6 Dec 2004, A. A. A.
Barbosa s.n. (HUFU). Pará: Cachimbo Range, 12 Dec
1956, J. M. Pires & al. 6084 (IAN, NY). — Colombia:
Camana, 1843, N. Funk s.n. (P). — French Guyana:
Road to Kaw, 5 Mar 2002, M. Pignal & N. Charrier-Arrighi 1968 (P, RB). — Venezuela: Orinoco River, Jun
1854, R. Spruce 3603 (K, P).
Cleistes rosea f. augusta (Hoehne) Meneguzzo & Van
den Berg, comb. & stat. nov. ≡ Pogonia rosea var.
augusta Hoehne, Com. Lin. Telegr., Bot. 4: 9. 1912 ≡
Cleistes latiglossa var. alba Hoehne, Fl. Bras. 12(1):
219. 1940, nom. illeg. superfl. – Lectotype (designated
here): Brazil, Mato Grosso, Augusto Fall, River Juruena,
Feb 1912, F. C. Hoehne sub Commissão Rondon 5353
142
Meneguzzo & van den Berg: Solving a chimaeric specimen and two neglected orchid names
(R R000002498!). – Syntype: ibidem, F. C. Hoehne sub
Commissão Rondon 5352 (R accession no. 44784!).
Remarks — Pogonia rosea var. augusta was included in
the revision of Cleistes for Brazil by Hoehne (1940), but
neither this name nor C. latiglossa var. alba was cited by
Pansarin (2005). The name C. latiglossa var. alba is an
illegitimate superfluous name for P. rosea var. augusta
because the protologue of both names cited the same
type specimen, hence the earlier legitimate name should
have been used for eventual combinations (Turland & al.
2018: Art. 11.4 and Art. 52.1). Therefore, its proper combination is made here in Cleistes, but instead of a variety
we choose the rank of form, as we have been adopting for
infraspecific taxa in which the only morphological difference is a sporadic mutation in flower colour (as discussed
and used by Meneguzzo & al. 2015). Cleistes rosea f.
augusta differs from C. rosea f. rosea by possessing dull
white sepals and petals and a white labellum with a pinkish veined interior.
Additional specimen examined — Brazil: Mato Grosso:
Salto Augusto, River Juruena, Feb 1912, F. C. Hoehne
sub Commissão Rondon 5584 (R).
Cleistes speciosa Gardner in Hooker’s Icon. Pl. 5: t.
473–474. 1842 ≡ Pogonia speciosa (Gardner) Rchb. f., Xenia Orchid. 2: 90. 1865. – Lectotype (designated here):
Brazil, Tocantins [“Goyaz”], Natividade, between Natividade and Arraias [“Arrayas”], Feb 1840, G. Gardner 4003
(K K000463766!; isolectotypes: F F0046279F [accession no. 1025428]!, GH 00056702 [AMES accession no.
70959 & GH accession no. 8037]!, K K000463767!, NY
00009270!, NY 00547506!, P P00367096!, P P00367097!,
SP SP003575 [accession no. 114345]!).
= Pogonia monantha Barb. Rodr., Gen. Spec. Orchid. 1:
167. 1877 ≡ Cleistes monantha (Barb. Rodr.) Schltr.
in Arch. Bot. São Paulo 1: 179. 1926. – Lectotype
(designated here): Brazil, São Paulo, Cajurú, Serra
das Lajes, 14 Mar 1849, A. F. Regnell III.1158 (S 077195!; isolectotypes: S 07-7196!, P P00367126!,
P P00367127!), syn. nov.
= Cleistes caloptera Rchb. f. & Warm., Otia Bot. Hamburg. 2: 82. 1881, syn. nov. ≡ Pogonia caloptera
Rchb. f. & Warm. in Otia Bot. Hamburg. 2: 82. 1881.
– Lectotype (designated here): [icon] (W 0079662
[Rchb. Herb. Orchid. No. 48046] [left-hand sheet of
drawings]!).
= Pogonia metallina Barb. Rodr. in Revista Engen. 3:
74. 1881, syn. nov. ≡ Cleistes metallina (Barb. Rodr.)
Schltr. in Arch. Bot. São Paulo 1: 179. 1926. – Lectotype (designated here): [icon] original illustration
that was to be published by Barbosa Rodrigues in
Iconographie des Orchidées du Brésil, deposited at
the library of Jardim Botânico do Rio de Janeiro (accession no. OR/584.150981); reproduced in Barbosa
Rodrigues (1996: vol. 1: 92, t. 39).
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= Pogonia lenheirensis Barb. Rodr. in Contr. Jard. Bot.
Rio de Janeiro 1: 47. 1901, syn. nov. ≡ Cleistes lenheirensis (Barb. Rodr.) Hoehne, Fl. Bras. 12(1): 232.
1940. – Lectotype (designated here): [icon] “Pogonia lenheirensis”, Barbosa Rodrigues in Contr. Jard.
Bot. Rio de Janeiro 1: t. VI, fig. D: 1 – 8. 1901.
– “Bacamania speciosa” (Gardner, in sched.), nom.
inval.
Remarks — During the study of Epistephium monanthum we found additional nomenclatural problems in the
heterotypic Cleistes monantha. Initially we thought that
the latter name should be the correct heterotypic name
to be used instead of C. metallina, as indicated by Pansarin (2005). This author pointed out that C. caloptera
and Pogonia lenheirensis are conspecific, a position we
agreed with. However, we studied the original specimens
of C. speciosa collected by George Gardner in the Brazilian central plateau. The study of this material led us
to the conclusion that this name also is conspecific with
P. monantha Barb. Rodr. Because C. speciosa was published 35 years earlier than P. monantha Barb. Rodr., the
former should be used for this species. Consequently, C.
caloptera, P. lenheirensis, P. metallina, and P. monantha
Barb. Rodr. are heterotypic synonyms of C. speciosa.
The original specimens of Cleistes speciosa were labelled by Gardner as being from the province of Goyaz,
near Natividade. However, Goiás State (the current spelling) has been divided into two states in 1988 and currently the specific locality is in Tocantins State. “Bacamania
speciosa Gardn.” was merely a name in schedula, i.e. an
unpublished name handwritten by Gardner on the label
of the lectotype specimen of C. speciosa in K. Neither
“Bacamania” nor “B. speciosa” is a validly published
name because a handwritten label on a herbarium specimen does not constitute effective publication (Turland &
al. 2018: Art. 29 – 30, 32.1(a)).
Pansarin (2005) stated that only the name Pogonia
caloptera was proposed in the protologue, and the combination under Cleistes was made later by Schlechter (in
Schlechter & Hoehne 1926). However, both names C.
caloptera and P. caloptera are alternative names by simultaneous publication in the protologue (Turland & al.
2018: Art. 36.3).
The illustration of Pogonia metallina published in
the protologue (Barbosa Rodrigues 1881: t. II C) depicts only a dissected perianth. However, the original illustration of the complete specimen, which includes the
identical elements illustrated in the protologue, was kept
unpublished until it was later reproduced by Barbosa Rodrigues (1996: vol. 1: 92, t. 39). Hence, the original illustration of the complete specimen is designated here as
the lectotype of P. metallina. The illustration published in
the protologue was miscaptioned as “Pogonia montana”.
We speculate that this mistake was induced by the publication of a second name with an identical specific epithet,
i.e. Galeandra montana Barb. Rodr, in the same article
Willdenowia 50 – 2020
(Barbosa Rodrigues 1881: 73). As for P. lenheirensis, the
whereabouts of its original illustration is unknown, therefore the one depicted in the protologue is designated here
as the lectotype.
Additional specimens examined — Brazil: Bahia: Serra
dos Lençóis, 26 May 1980, R. M. Harley & al. 22654
(CEPEC, K). Distrito Federal: Brasília, 25 Jan 1997, J.
A. N. Batista 672 (CEN). Goiás: Chapada dos Veadeiros,
16 Mar 1969, H. S. Irwin & al. 24496 (IAN, NY, UB).
Minas Gerais: Miguel Burnier, 19 Apr 1957, E. Pereira
& G. F. J. Pabst 3014 (RB).
Gomesa stricta and its identity
Pleurothallis quadrifida (Lex.) Lindl. in Edwards’s Bot.
Reg. 21: sub t. 1797. 1835 ≡ Dendrobium quadrifidum
Lex. in La Llave & Lexarza, Nov. Veg. Descr. 2(Orchid.
Opusc.): 40. 1825 ≡ Humboltia quadrifida (Lex.) Kunth,
Rev. Gen. Pl. 2: 668. 1891 ≡ Stelis quadrifida (Lex.)
Solano & Soto Arenas, Icon. Orchid. 5 – 6: 11. 2003 ≡
Specklinia quadrifida (Lex.) Luer in Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 263. 2004 ≡ Loddigesia quadrifida (Lex.) Luer in Monogr. Syst. Bot. Missouri Bot.
Gard. 105: 251. 2006 ≡ Lalexia quadrifida (Lex.) Luer in
Harvard Pap. Bot. 16: 358. 2011. – Neotype (designated
here): Mexico, Michoacan, Coalcoman, 4 Apr 1939, G.
B. Hilton I3643 (ARIZ acession no. 12480!).
= Gomesa stricta Spreng., Syst. Veg. 3: 730. 1826 ≡
Rodriguezia stricta (Spreng.) Steud., Nomencl. Bot.,
ed. 2, 2: 463. 1841. – Lectotype (designated here):
Jamaica, s. loc., Jun 1821, C. L. G. Bertero 1676 (TO
s.n.!).
– “Physanthera callistachys” (Sprengel, Syst. Veg. 3:
730. 1826), nom. inval.
Remarks — Gomesa stricta. is a name proposed for a
Jamaican species that has been cited for a long time in the
literature as being of unknown application to a biological
entity. It drew the attention of one of us (T.E.C.M.) during the revision of the genus Gomesa, which is restricted
to South America and has its main diversity centred in
eastern Brazil. We confirm the identity of G. stricta as a
heterotypic synonym of Pleurothallis quadrifida.
Gomesa stricta was described by Sprengel (1826:
730) solely based on a specimen from “Jamaica” and
collected by “Bertero”. We contacted all herbaria cited
in Index herbariorum (Thiers 2020+) which claimed to
have Carlo Luigi Giuseppe Bertero’s specimens in their
collections. Ackerman (2014) cited the type specimen as
deposited at herbarium G, although he had not studied
it. It seems it had been a misattribution because it was
not located. The only herbarium that returned a positive
answer for a specimen labelled as G. stricta or any of its
denominations was the Università degli Studi di Torino
(TO). This specimen is designated here as the lectotype
and seems to have been signed and dated by Sprengel
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143
himself, or at least it could be attributed as a reference to
him because we were not able to confirm the handwriting was by Sprengel. The lectotype consists of two stems
bearing inflorescences, but the leaves are lacking.
“Physanthera callistachys Bert.” was cited as a
synonym in the protologue of Gomesa stricta, and later “Physanthera Berter.” and “P. callistachys Bert[er].”
were cited as synonyms of Rodriguezia and R. stricta,
respectively, by Steudel (1841: 330, 463). These were
subsequently frequently cited in botanical literature incorrectly as validly published names, e.g. by Govaerts &
al. (2020), but they are not validly published because they
were merely cited as synonyms (Turland & al. 2018: Art.
36.1(b)). The only validly published names are the basionym G. stricta and the new combination R. stricta, the
latter mostly probably following Lindley’s (1827) treatment uniting Gomesa and Rodriguezia.
Cogniaux (1909 – 1910: 621) considered Rodriguezia stricta as a doubtful name in his treatment for the
Antilles. However, he did cite a specimen collected
by Bertero under Pleurothallis longissima Lindl. (Cogniaux 1909 – 1910: 398), and a corresponding specimen
was virtually untraceable by the supplied internal data.
Fawcett & Rendle (1910) and Adams (1966) made no
mention of this debatable name in their treatments of the
orchid flora of Jamaica and neither did Nir (2000) for the
whole of the Antilles. In the orchid flora of the Greater
Antilles, Ackerman (2014: 573) kept it as an excluded
taxon and believed that it could be a mislabelled specimen from somewhere else that Bertero had collected in
the Neotropics.
Ormerod (2006) made a risky assumption based
on circumstantial evidence on the identity of Gomesa
stricta as a new synonym of Pleurothallis quadrifida.
He referred to the just-mentioned Bertero specimen by
Cogniaux (1909 – 1910), considered P. longissima a synonym of P. quadrifida, and Sprengel’s (1826) diagnosis as
a good and reliable enough description. From a modern
perspective this diagnosis would be considered too vague
and could be applied to numerous orchid species. Despite
this, our study of the lectotype of G. stricta agrees on its
conspecificity with P. quadrifida. We revised the protologues, the list of synonyms presented by Luer (2000) and
Ackerman (2014), and their respective types to confirm
the synonymy. To consolidate the synonym status and because the type designation for Dendrobium quadrifidum,
the basionym of P. quadrifida, was not done by previous authors, we designate here a neotype from a locality
as close as possible to the one of the original specimens
because it has not been collected from the type locality
again (López Ferrari & Serna 2000; Luer 2000; Nir 2000;
Ackerman 2014; Ormerod 2016).
The generic circumscriptions in Pleurothallidinae
have been rather controversial and unstable over the last
couple of decades. For the species discussed here, the
monotypic genus Loddigesia Luer (2006: 251), nom. illeg. (non Loddigesia Sims; Leguminosae), was described
144
Meneguzzo & van den Berg: Solving a chimaeric specimen and two neglected orchid names
and later renamed as Lalexia Luer (2011: 358). In the current study, we follow a more conservative circumscription
that places this species in Pleurothallis because P. quadrifida is phylogenetically placed as a sister group of Pleutothallis (Karremans & al. 2013). Hence, “Physanthera”
is a synonym of Pleurothallis, not of Rodriguezia as listed
by Steudel (1841). However, under a splitting approach,
“Physanthera” would be a synonym of Lalexia.
Additional specimens examined — Costa Rica: Guanacaste, 3 Jan 1964, L. O. Williams & al. 26550 (F). —
Jamaica: Fairfield, s.d., H. R. Wullschägel 1383 (M). —
Venezuela: Portuguesa, 15 Nov 1951, O. Renz 7359
(RENZ).
The chimeric artefact Maxillaria spathacea
Bifrenaria harrisoniae (Hook.) Rchb. f. in Bonplandia (Hanover) 3: 217. 1855 ≡ Dendrobium harrisoniae
Hook., Exot. Fl. 2: t. 120. 1824 ≡ Maxillaria harrisoniae
(Hook.) Lindl., Bot. Reg. 11: t. 897. 1825 ≡ Colax harrisoniae (Hook.) Lindl. ex Spreng., Syst. Veg. 3: 727. 1826
≡ Stanhopea harrisoniae (Hook.) P. N. Don in Donn,
Hortus Cantabrig., ed. 13: 607. 1845 ≡ Lycaste harrisoniae (Hook.) P. N. Don in Donn, Hortus Cantabrig.,
ed. 13: 721. 1845. – Lectotype (designated by Koehler &
Amaral 2004: 327): [icon] “Dendrobium harrisoniae” in
Hooker, Exot. Fl. 2: t. 120. 1824.
= Maxillaria spathacea Lindl., Gen. Sp. Orchid. Pl.:
151. 1832. – Lectotype (designated here): Brazil,
s. loc., s.d., Boaz [sic] W. Boog s.n. (K K000879747!
flowers only, excluding vegetative part [Cattleya sp.]
of K K000879700!).
– “Bifrenaria harrisoniae var. typica” (Hoehne in Arq.
Bot. Estado São Paulo 2: 116. 1950), nom. inval.
Remarks — In late 2012, while handling specimens at K,
we faced a very peculiar and curious specimen stored as
an unidentified species of Bifrenaria. Our first examination along with annotations on the herbarium sheet, and
later literature inspection, revealed it to be the chimeric
original specimen of the name Maxillaria spathacea. It
consists of an assemblage of flowers of B. harrisoniae
and vegetative parts of an unidentified species of Cattleya
subg. Intermediae (Cogn.) Withner sensu van den Berg
(2014). Maxillaria spathacea was not cited by Koehler &
Amaral (2004) in their revision of Bifrenaria.
The specimen of Maxillaria spathacea was part of
William Jackson Hooker’s personal herbarium later incorporated into K in 1867. The only presented information is a locality “Brazil” and a collector whose name
is discussed below. It is annotated by John Lindley as
“Maxillaria”, and by a third person as “Max. spathacea
Lindl. Mss. [?] Orch. p. 151”, and by Reichenbach filius, more or less as quoted from his work (Reichenbach
1856): “Maxillaria spathacea: est artefactum ex Bifrenaria Harrisoniae et Cattleya quadam. (Vid. sp. typ. in
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hb. Hook.).” [M. spathacea is an artefact of B. harrisoniae and a Cattleya (I have seen the type specimen in
the Hooker herbarium); our translation and adaptation].
Additionally, Eric A. Christenson in April 1994 annotated the specimen as “Plant = Cattleya [;] Flowers =
Bifrenaria”.
Lindley (1832) described the name Maxillaria
spathacea with a diagnosis including both vegetative
and reproductive parts. On the specimen at K he credited the collector as being “Mr. Boog”, but the handwritten name is slightly unclear and could easily be read as
“Mr. Boaz” or “Mr. Booz”. We consider that it refers to
William Boog, who extensively sent botanical specimens
to Hooker (Kew Archives, Director’s Correspondence,
item KMDC1677). Lindley also expressed doubt by including a question mark next to the new species name,
but the nature of such uncertainty has not been detailed.
Apparently, Reichenbach (1856) was the first to indicate the chimeric nature of the specimen, but he did
not undertake any taxonomic action. Cogniaux (1898)
resolved it by synonymizing M. spathacea under Bifrenaria harrisoniae. Curiously, the vegetative part is
here confirmed by us as belonging to some species of
Cattleya subg. Intermediae. The specimen also carries
an inflorescence enclosed by a spathaceous bract (from
which the original flowers were removed), after which
the specific epithet was probably coined. Exactly in the
axil of each floral bract of the Cattleya inflorescence,
the B. harrisoniae flowers identified by Reichenbach
(1856) have been skilfully attached as to appear somewhat natural. Because the preparation is undoubtedly
an admixture, we designate as the lectotype the flowers that can be unequivocally identified at specific level
(Turland & al. 2018: Art. 8.2 and 9.14). Consequently,
M. spathacea is confirmed as a heterotypic synonym of
B. harrisoniae, and the vegetative Cattleya part is excluded from the lectotype.
Among the synonyms of Bifrenaria harrisoniae
we detected an additional nomenclatural matter to be
clarified. “Bifrenaria harrisoniae var. typica” (Hoehne
1950: 116) is not a validly published name (Turland &
al. 2018: Art. 24.3 and 26.2) because it was applied to
a variety that contained the typical element of B. harrisoniae, yet its final epithet was “typica” rather than
“harrisoniae”, which would have been correct for the
autonymic variety.
The specimen itself as an artefact received much attention and opened room for questions. The specimen
could simply be regarded as an unwanted result of a mixture of specimens of two different and phylogenetically
unrelated species. Or perhaps, it could be seen as a trick
played on botanists, that caught Hooker and Lindley unawares, and just remained without solution.
Additional specimens examined — Brazil: Minas Gerais, Caldas Range, 17 Oct 1876, A. F. Regnell III 4410
(S). Paraná, Balsa Nova, 4 Oct 2003, J. M. Silva & al.
Willdenowia 50 – 2020
3792 (ALCB, MBM). Rio de Janeiro, Rio de Janeiro, Nov
2015, G. Cattan 117 (RB). São Paulo, Bocaina Range, 11
Aug 1968, D. Sucre & P. I. S. Braga 3497 (RB).
Acknowledgements
We are grateful to Nicholas Turland for his significant
advice on the manuscript; Jefferson Prado for his constant support on nomenclatural advice to our work on
Orchidaceae; Mark A. Nir for a valuable discussion
on Gomesa stricta; herbarium ESA for pictures of the
original specimen of Cleistes angeliana, herbarium
TO for G. stricta, herbarium W for C. caloptera, and
other cited herbaria for access to their collections. This
work was funded by grants from Conselho Nacional
de Desenvolvimento Científico e Tecnológico, CNPq,
processes
MCT/CNPq/FNDCT/MEC/CAPES/FAP’s
REFLORA 563556/2010-2 and 563535/2010-5. TECM
was granted a post-doctoral researcher scholarship from
CAPES/Brasil/Herbário Virtual da Flora e dos Fungos
88887.351871/2018-00. We also thank Adam P. Karremans (Jardín Botánico Lankester, Universidad de Costa
Rica) and Andre Schuiteman (Royal Botanic Gardens,
Kew) for their comments on an earlier version of this
article.
References
Ackerman J. D. 2014: Orchid flora of the Greater Antilles. – Mem. New York Bot. Gard. 109: 1 – 625.
Adams C. D. 1966: A checklist of the orchids of Jamaica.
– Amer. Orchid. Soc. Bull. 35: 995 – 999.
Ames O. 1922: A discussion of Pogonia and its allies in
the northeastern United States with reference to extralimital genera and species. – Orchidaceae 7: 3 – 38.
Barbosa Rodrigues J. 1881: Resultado botanico de uma
breve excursão a S. João d’El-Rei, Minas Geraes
[conclusão]. – Revista Engen. 3: 73 – 75, t. II.
Barbosa Rodrigues J. 1996: – In: Sprunger S., Cribb P. &
Toscano de Brito A. [L. V.] (ed.), Iconographie des Orchidées du Brésil. 2 vols. – Basel: Friedrich Reinhardt.
Chase M. W., Cameron K. M., Freudenstein J. V., Pridgeon
A. M., Salazar [Chávez] G. [A.], van den Berg C. &
Schuiteman A. 2015: An updated classification of Orchidaceae. – Bot. J. Linn. Soc. 177: 155–174.
Cogniaux [C.] A. 1893: Orchidaceae: Epistephium. – Pp.
135 – 144, t. 28 – 31 in: Martius C. F. P. de, Eichler A.
G. & Urban I. (ed.), Flora brasiliensis […] 3(4). –
Monachii: Typographia Regia.
Cogniaux [C.] A. 1898: Orchidaceae: Bifrenaria. – Pp.
476 – 498, t. 95 – 96, 108 in: Martius C. F. P. de, Eichler
A. G. & Urban I. (ed.), Flora brasiliensis […] 3(5). –
Monachii: Typographia Regia.
Cogniaux [C.] A. 1909 – 1910: Orchidaceae. – Pp.
293 – 696 in: Urban I. (ed.), Symbolae antillanae 6.
Downloaded From: https://bioone.org/journals/Willdenowia on 23 Jul 2020
Terms of Use: https://bioone.org/terms-of-use
145
– Lipsiae: Fratres Borntraeger; Parisiis: Paul Klincksieck; Londini: Williams & Norgate.
Fawcett W. & Rendle A. B. 1910: Flora of Jamaica […]
1. Orchidaceae. – London: Trustees of the British
Museum.
Garay L. A. 1978: Orchidaceae: Cypripedioideae, Orchidoideae, Neottioideae. – In: Harling G. & Sparre
B. (ed.), Flora of Ecuador 225(1). – Stockholm: Göteborgs Universitet & Riksmuseum.
Govaerts R., Bernet P., Kratochvil K., Gerlach G., Carr
Junior G. F., Alrich P., Pridgeon A. M., Pfahl J., Campacci M. A., Baptista D. H., Tiggers H., Shaw J. M.
H., Cribb P., Georg A., Kreuz K. & Wood J. 2020
[continuously updated]: World checklist of Orchidaceae. Facilitated by the Royal Botanical Gardens, Kew.
– Published at http://wcsp.science.kew.org/ [accessed
6 Jan 2020].
Hoehne F. C. 1940: Orchidaceae: Selenipedium – Triphora. – In: Hoehne F. C. (ed.), Flora brasílica 12(1).
– São Paulo: Secretaria de Agricultura do Estado de
São Paulo.
Hoehne F. C. 1945: Orchidaceae: Vanilla – Corymborchis. – In: Hoehne F. C. (ed.), Flora brasílica 12(2).
– São Paulo: Secretaria de Agricultura do Estado de
São Paulo.
Hooker W. J. 1824: Dendrobium harrisoniae. – Exot. Fl.
2: t. 120.
Karremans A. P., Bakker, F. T., Pupulin, F., SolanoGómez, R. & Smulders M. J. M. 2013: Phylogenetics of Stelis and closely related genera (Orchidaceae: Pleurothallidinae). – Pl. Syst. Evol. 299:
151 – 176.
Koehler S. & Amaral M. C. E. 2004: A taxonomic study
of the South American genus Bifrenaria Lindl. (Orchidaceae). – Brittonia 56: 314 – 345.
Kuntze [C. E.] O. 1891: Revisio generum plantarum […]
2. – Leipzig: Arthur Felix; London: Dulau & Co.; Milano: U. Hoepli; New-York: Gust. E. Stechert; Paris:
Charles Klincksieck.
Lindley J. 1827: Orchidaceous plants. – Trans. Hort. Soc.
London 7: 67 – 71.
Lindley J. 1832: Maxillaria. – Pp. 142 – 151 in: The genera and species of orchidaceous plants. – London:
Ridgways.
Lindley J. 1836: Pleurothallis grobyi. – Edward’s Bot.
Reg. 21: t. 1797.
Lindley J. 1840: Epistephium. – Pp. 432 – 434 in: The
genera and species of orchidaceous plants. – London:
Ridgways.
Lindley J. 1842: Pleurothallis. – Edward’s Bot. Reg.
28(misc.): 67 – 84.
López-Ferrari A. R. & Serna A. E. 2000: Nuevas combinaciones en monocotiledóneas mexicanas III (Orchidaceae, Poaceae). – Acta Bot. Mex. 51: 61 – 70.
Luer C. A. 2000: Systematics of Pleurothallis subgenus
Effusia. – Monogr. Syst. Bot. Missouri Bot. Gard. 79:
53 – 103.
146
Meneguzzo & van den Berg: Solving a chimaeric specimen and two neglected orchid names
Luer C. A. 2006: Miscellaneous new taxa in the Pleurothallidinae (Orchidaceae). – Monogr. Syst. Bot. Missouri Bot. Gard. 105: 245 – 259.
Luer C. A. 2011: Miscellaneous new species in the Pleurothallidinae (Orchidaceae) excluding species from
Brazil. – Harvard Pap. Bot. 16: 311 – 360.
McNeill J. 2014: Holotype specimens and type citations:
general issues. – Taxon 63: 1112 – 1113.
Meneguzzo T. E. C., Baumgratz J. F. A & van den Berg
C. 2015: Taxonomic studies in the Aganisia complex
(Orchidaceae, Zygopetalinae). – Phytotaxa 238: 1–39.
Nir M. A. 2000: Orchidaceae antillanae. – New York:
DAG Midia Publishing.
Ormerod P. 2016: Neotropical orchid miscellanea. – Harvard Pap. Bot. 21: 231 – 245.
Pabst G. F. J. & Dungs F. 1977: Orchidaceae brasilienses
2. – Hildesheim: Kurt Schmersow.
Pansarin E. R. 2005: Sistemática filogenética e biologia
floral de Pogoniinae sul-americanas, e revisão taxonômica e análise das ceras epicuticulares do gênero
Cleistes Rich. ex Lindl. (Orchidaceae). – Campinas:
unpublished PhD thesis, Universidade Estadual de
Campinas.
Reichenbach H. G. 1856: Stipulae Orchidaceae Reichenbachianae intra “folia” Lindleyana intraaxillares. –
Bonplandia (Hanover) 4: 321 – 330.
Reichenbach H. G. 1861 – 1864: Orchides. – Ann. Bot.
Syst. 6: 167 – 933.
Reichenbach H. G. 1865: Zu Pogonia. – Pp. 88 – 92 in:
Xenia orchidacea. Beiträge zur Kenntnis der Orchideen 2. – Leipzig: F. A. Brockhaus.
Romero-González G. A. 2005: Orchidaceae schomburgkianae: the orchids collected by R. H. Schomburgk
in South America and the Caribbean. – Harvard Pap.
Bot. 10: 231 – 268.
Schlechter [F. R.] R. 1929: Figuren-Atlas zu den Orchideenfloren der Südamerikanischen Kordillerenstaaten. – Repert. Spec. Nov. Regni Veg., Beih. 57: t.
1 – 142.
Schlechter [F. R.] R. & Hoehne F. C. 1926: Contribuições
ao conhecimento das Orchidáceas do Brasil. – Arch.
Bot. Estado São Paulo 1: 165 – 349.
Schweinfurth C. 1958: Orchids of Peru. – Fieldiana, Bot.
30: 1 – 260.
Sprengel C. [P. J.] 1826: Systema vegetabilium 3. –
Gottingae: sumtibus librariae Dieterichianae.
Steudel E. T. 1841: Nomenclator botanicus […], ed. 2,
2. – Stuttgartiae et Tubingae: typis et sumptibus J. G.
Cottae.
Thiers B. 2020+ [continuously updated]: Index herbariorum: a global directory of public herbaria and associated staff. New York Botanical Garden’s virtual
herbarium. – Published at http://sweetgum.nybg.org
/science/ih/ [accessed 27 Jan 2020].
Turland N. J., Wiersema J. H., Barrie F. R., Greuter W.,
Hawksworth D. L., Herendeen P. S., Knapp S., Kusber
W.-H., Li D.-Z., Marhold K., May T. W., McNeill J.,
Monro A. M., Prado J., Price M. J. & Smith G. F.
(ed.) 2018: International Code of Nomenclature for
algae, fungi, and plants (Shenzhen Code) adopted
by the Nineteenth International Botanical Congress
Shenzhen, China, July 2017. – Glashütten: Koeltz
Botanical Books [Regnum Veg. 159].
van den Berg C. 2014: Reaching a compromise between
conflicting nuclear and plastid phylogenetic trees: a
new classification for the genus Cattleya (Epidendreae; Epidendroideae; Orchidaceae). – Phytotaxa
182: 75 – 86.
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