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LANKESTERIANA 19(3): 281–343. 2019.
TO BE, OR NOT TO BE A STELIS
AdAm P. KArremAns
Jardín Botánico Lankester, Universidad de Costa Rica, Cartago, Apartado 302-7050, Costa Rica
Naturalis Biodiversity Center, Endless Forms, Sylviusweg 72, Leiden 2333 BE, The Netherlands.
adam.karremans@ucr.ac.cr
AbstrAct. Despite the availability of multiple sources of evidence and consistency in the support for a broadly
circumscribed Stelis Sw. (Orchidaceae: Pleurothallidinae), some authors continue to be hesitant in its use. It
is certain that the more typical species of Stelis, with their triangular, attish owers with very short eshy
petals and lip, form a monophyletic group that is easily recognized. However, it is likewise undisputed that
they are not an isolated lineage in the subtribe and that several groups of species with a similar vegetative
habit but lacking the typical Stelis ower are in fact very close relatives, sharing a relatively recent common
ancestor. Those species groups need to be classied in a way that also reects their own evolutionary history;
alternatives to a broadly circumscribed Stelis are possible yet neither straightforward or practical at this time.
An infrageneric classication for the whole group is provided here in an attempt to clarify which species
belong where in this highly complex afnity. Emphasis is made on the difculty of diagnosing the less typical
members of each proposed subgenus or section, and on the importance of oral convergence and divergence
as a result of pollinator adaptation. As here dened, Stelis is the largest genus in the Pleurothallidinae, with
1243 species.
Key words: convergence; evolutionary history; oral morphology; generic circumscription; Pleurothallidinae;
pollinator adaptation
doi: http://dx.doi.org/10.15517/lank.v19i3.40082
Received 23 October 2019; accepted for publication 14 December 2019. First published online: 17 December 2019.
Licensed under a Creative Commons Attribution-NonCommercial-No Derivs 3.0 Costa Rica License.
Introduction. What is a Stelis? Or better yet, what isn’t
a Stelis? Some authors may think this is the question we
are still asking ourselves today, but in fact the matter has
been settled for years. We have an indisputable answer.
Rather, what we are still actually asking ourselves is
how can we classify the different groups of species
within the Stelis afnity in a way that both reects their
evolutionary history and satises most users of such a
classication system. That is the only question that still
remains, and for that we may never have an answer that
pleases everyone.
Species of Stelis Sw. (Orchidaceae: Pleurothallidi-
nae) in the tradicional, strict, sense are characterized
by a more or less at, triangular ower, bearing three
subequal, larger, spreading sepals, compact petals
and lip, and a short column, with an apical anther and
stigma. It was one of the rst genera to be recognized
in subtribe Pleurothallidinae, and has been used
relatively consistently for around two centuries. With
few exceptions, members of Stelis s.s. have an easily
recognizable standard ower morphology. DNA data
proves they form a monophyletic group, and there is
no dispute as to which species belong to it, and which
do not. Let’s be clear, there is no doubt that all 1,030
currently accepted species bearing owers with the
classic Stelis morphology are more closely related
to each other than they are to any other species of
Pleurothallidinae. In every sense, Stelis in its traditional
circumscription is a well supported group. Why, then,
don’t we simply recognize them as a genus on their own
and get on with it? Well, because the species of Stelis s.s.
are not an island within the subtribe. They have many
close relatives that need to be classied in a way that
reects their own evolutionary history as well. After all,
there is undisputed evidence that species of Stelis in the
strict sense share an ancestor with many species that
lack the typical Stelis-like owers.
Historically, recognition of genera in Pleurothalli-
dinae has been done by segregating groups of species
that could be easily set aside from all others through
key morphological features (Karremans 2016). Most
other species simply remained in a broadly dened
Pleurothallis R.Br., not for being related to each other
or sharing particular characteristics, but for the lack of
the highly distinctive features of the segregated genera.
Pleurothallis sensu lato had always been expected to
be polyphyletic (Lindley 1859, Luer 1986, Neyland
et al. 1995, Stenzel 2000). DNA based phylogenetic
reconstructions essentially came to demonstrate how
polyphyletic it actually was and to stress the necessity
for a new circumscription of genus Pleurothallis, whose
members were found to be diversely related to most of
the other genera in Pleurothallidinae. What is relevant
to this discussion is that several groups of species
previously assigned to Pleurothallis were proven to
be more closely related to Stelis in the strict sense than
anything else, despite their oral morphology. These
species can no longer be treated as Pleurothallis because
we know for a fact that their ancestors took a different
evolutionary path, which ultimately gave origin to Stelis
in the strict sense.
It is undeniable that the generic circumscription
of Stelis, as dened by Pridgeon (2005) and modied
by Karremans et al. (2013), has not been broadly
accepted. That is most likely due to the ease of orally
recognizing a member of Stelis sensu scrito and the lack
of obvious oral features uniting species of Stelis sensu
lato (Solano-Gómez & Salazar 2013). It is desirable that
genera are diagnosable using morphological features,
and not only through DNA analyses. However, it is also
very important to be accurate about the phylogenetic
relationships among species, establishing groups that
reect the evolutionary histories of its members. If
one were to look past the obvious differences in oral
morphology, which undoubtedly respond to pollinator
pressure, all members of Stelis s.l. are vegetatively very
similar to each other. So much so that without owers it
is difcult to tell them apart.
A partitioning of Stelis s.l. into several smaller,
discrete, morphologically better-dened genera is
possible. It was in fact advocated by Karremans
(2010) and Karremans & Bogarín (2013), and could be
desirable. Nevertheless, to do so one needs to have a
clear evolutionary picture of the whole group. It is not as
easy as separating the most obvious close relatives into
genera, or simply separating Stelis s.s. from everything
else. The whole picture is much more complex and
the reality is that although some relationships within
Stelis s.l. are easily diagnosable, the placement of many
species continues to be a challenge, even with DNA data.
Anyone can diagnose a species as belonging to either
Salpistele Dressler, Stelis s.s., or Physosiphon Lindl., for
example. However, not even the most adamant expert
could’ve predicted that species of Pleurothallis sect.
Petiolatae Luer were sister to those of Salpistele, or
that the Stelis imraei (Lindl.) Pridgeon & M.W.Chase
group belonged in a completely isolated lineage. It is
this lack of predictability that makes classifying these
groups challenging. Unfortunately, there is no easy way
out. The recognition of any segregate genus from Stelis
s.l. requires the recognition of several additional new
genera and the recircumscription of most of the existing
ones (Karremans 2016). The resulting classication
would be neither intuitive or very useful. At this time, it
is preferable to maintain Stelis s.l. rather than to promote
the use of Stelis s.s. together with a series of ill-dened,
non-monophyletic satellite genera.
Some authors may believe that solving the “Stelis
issue” is merely a matter of segregating the members
of Stelis s.s. from the remaining Stelis s.l., but this
is not a viable solution. Several groups within Stelis
s.l. are more closely related to Stelis s.s. than to
other members of Stelis s.l. It is also not a matter of
simply recognizing the more apparent genera like
Crocodeilanthe Rchb.f., Dracontia (Luer) Luer and
Salpistele, as advocated by several authors (Karremans
2010; Karremans & Bogarín 2013; Toscano de Brito
2018a; Damián 2019). Most Crocodeilanthe species
are indeed easily distinguished from other members
of Stelis s.l., but certainly not all of them have those
very evident morphological features of their most
distinctive members. They are closely related to
the species previously assigned to Pleurothallis
sect. Acuminatae Lindl. and those placed in genus
Physothallis Garay, which look nothing like the
Crocodeilanthe morphologically and should be either
included or segregated as a genus as well. Dracontia
may also seem straightforward, but it is not. At least
one species placed in Elongatia (Luer) Luer, another
from Pseudostelis Schltr., a few placed in Effusiella
Luer, and the type of Mystacorchis Szlach. & Marg.
are all intermingled with species of Dracontia. Species
of Salpistele, which have the most divergent oral
morphology among the Stelis s.l. are not only closely
related to species of Dracontia, but they are sister to
two species previously assigned to genus Elongatia
and which are orally completely different. What
do these species groups have in common? Most are
vegetatively similar to each other, but again, this is true
for all members of Stelis s.l.
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Every single phylogenetic study including members
Stelis s.l., as proposed by Pridgeon (2005) and modied
by Karremans et al. (2013), nds the genus to be
monophyletic (Karremans 2010, Chiron et al. 2012,
Ramos-Castro et al. 2012, Karremans et al. 2013,
Wilson et al. 2013, 2017, Pérez-Escobar et al. 2017),
and this continues to be the case in multi-gene genomic
studies (Chumová et al. 2018, Ponert et al. 2019).
Accepting Stelis s.s. as a genus on its own necessarily
entails the recognition of many ill-dened genera that
no user would be happy to adopt. The alternatives to
Stelis s.l. are even less appealing than it itself. Does
a broader concept of Salpistele, which as the oldest
name in the group has priority over all others, be a
more acceptable circumscription for the species of the
Dracontia clade? Perhaps a more inclusive concept of
Physothallis, harboring the species of Pleurothallis sect.
Acuminatae? Or the recognition of monotypic genera
for Stelis carpinterae (Schltr.) Pridgeon & M.W.Chase,
Stelis convallaria (Schltr.) Pridgeon & M.W.Chase, and
Stelis mystax (Luer) Pridgeon & M.W.Chase? Perhaps.
Nevertheless, the fact remains that it is difcult to assign
species to discrete groups in Stelis s.l., any grouping
being more or less articial, and any alternative
classication of this group results in genera that will
not be more accepted and better dened or recognizable
than Stelis s.l. Even though members of Stelis s.s.
are orally very easily diagnosable for anyone, and,
evidently, the owers of other members of Stelis s.l. are
very different, a broader circumscription of Stelis, with
all its defects, still seems preferable over its alternatives.
After all, we need to remember that although owers are
easily comparable with each other, they are under high
selective pressure of pollinators, and may be more or
less similar independently of relatedness (Karremans &
Díaz-Morales 2019).
For the sake of consistency with previous works
in related genera, including Acianthera Scheidw.
(Karremans et al. 2016), Andinia (Luer) Luer (Wilson et
al. 2017) and Specklinia Lindl. (Karremans et al. 2016),
and in the interest of aiding the reader, an infrageneric
classication of Stelis s.l. is provided (Fig. 1).
Figure 1. Relationships among the subgenera of Stelis sensu lato based on diverse phylogenetic reconstructions (Pridgeon
et al. 2001, Solano-Gómez 2005, Karremans 2010, Chiron et al. 2012, Ramos-Castro et al. 2012; Karremans et al.
2013, Wilson et al. 2017, Pérez-Escobar et al. 2017, Chumová et al. 2018, Ponert et al. 2019), showing the proposed
subgenera and sections.
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tAxonomic treAtment
Stelis Sw., J. Bot. (Schrader) 1799(2): 239. 1800, nom.
cons.
Syn. Humboltia Ruiz & Pav., Fl. Peruv. Prodr.: 121.
1794, nom. rej.
Syn. Physosiphon Lindl., Edwards’s Bot. Reg. 21:
t. 1797. 1835.
Syn. Dialissa Lindl., Ann. Mag. Nat. Hist. 15: 107.
1845.
Syn. Crocodeilanthe Rchb.f. & Warsz., Bonplandia
(Hannover) 2: 113. 1854.
Syn. Pseudostelis Schltr., Anexos Mem. Inst.
Butantan, Secç. Bot. 1(4): 36. 1922.
Syn. Physothallis Garay, Svensk Bot. Tidskr. 47:
199. 1953.
Syn. Steliopsis Brieger in F.R.R.Schlechter,
Orchideen Beschreib. Kult. Zücht., ed. 3,
8(29‒32): 457. 1976, nom. nud.
Syn. Apatostelis Garay, Bot. Mus. Lea. 27: 185.
1979, nom. illeg.
Syn. Salpistele Dressler, Orquideologia 14: 6. 1979.
Syn. Condylago Luer, Orquideologia 15: 118. 1982.
Syn. Mystacorchis Szlach. & Marg., Polish Bot. J.
46: 117. 2001.
Syn. Dracontia (Luer) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 257. 2004.
Syn. Unciferia (Luer) Luer, Monogr. Syst. Bot. Miss-
ouri Bot. Gard. 95: 265. 2004, nom. illeg. Non Un-
cifera Lindl., J. Proc. Linn. Soc., Bot. 3: 39. 1859.
Syn. Lomax Luer, Monogr. Syst. Bot. Missouri Bot.
Gard. 105: 88. 2006.
Syn. Effusiella Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 112: 106. 2007.
Syn. Niphantha Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 120: 154. 2010.
Stelis subgen. Stelis (Figs. 2‒4)
Syn. Humboltia Ruiz & Pav., Fl. Peruv. Prodr.: 121.
1794, nom. rej.
Syn. Dialissa Lindl., Ann. Mag. Nat. Hist. 15(96):
107. 1845. Type: Dialissa pulchella Lindl. Ann.
Mag. Nat. Hist. 15(96): 107. 1845.
Syn. Steliopsis Brieger, Orchideen (Schlechter)
8(29-32): 457. 1976, nom. nud. Type: Steliopsis
anneliesae Brieger, Orchideen (Schlechter)
8(29-32): 457. 1976, nom. nud.
Syn. Apatostelis Garay, Bot. Mus. Lea. 27: 185.
1979, nom. illeg. Type: Stelis hylophila Rchb.f.,
Bonplandia (Hannover) 3: 241. 1855.
Stelis cochabambensis Karremans, nom. nov.
Repl. syn.: Stelis dasysepala Luer & R.Vásquez,
Selbyana 32(1,2): 37. 2018, nom. illeg. Non Stelis
dasysepala Luer & R.Escobar, Harvard Pap. Bot.
21(2): 198. 2016.
Stelis luerii Karremans, nom. nov.
Repl. syn.: Stelis marginata Luer & R.Vásquez,
Selbyana 32(1,2): 71. 2018, nom. illeg. Non Stelis
marginata Luer & R.Escobar, Harvard Pap. Bot.
21: 205. 2016.
Stelis peculiaris Karremans, nom. nov.
Repl. syn.: Stelis praecipua Luer & R.Vásquez,
Selbyana 32(1,2): 87. 2018, nom. illeg. Non Stelis
praecipua Luer, Harvard Pap. Bot. 22: 101. 2017.
Stelis subgen. Stelis is synonymous to Stelis s.s. as
dened by Luer (2009). In other words, it includes all
the classical species of Stelis with triangular, attish
owers with very short petals and lip, the very short
column has an apical anther and stigma, and the
pollinaria have a drop-like viscidium attached to the
short caudicles. Many species of Stelis subgen Stelis
have been analyzed genetically and they always group
together into a highly supported clade with low genetic
variation. There is a single exception, and that is an
accession labeled Stelis nexipous Garay in Karremans
et al. (2013) that appeared associated with members of
Stelis subgen. Niphantha. It is surely either a lab mixup
or sequencing mistake.
A comprehensive species list is not yet presented
here. However, of the 1243 species currently accepted
in genus Stelis s.l. (Karremans, in prep.), 1030 belong
to Stelis s.s. The remaining 213 species are listed under
one of the other subgenera hereafter.
Stelis subgen. Crocodeilanthe (Rchb.f. & Warsz.)
Karremans, comb. nov.
Bas. Crocodeilanthe Rchb.f. & Warsz., Bonplandia
(Hannover) 2: 113. 1854. Pleurothallis subgen.
Crocodeilanthe (Rchb.f. & Warsz.) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 20: 34.
1986. Type: Crocodeilanthe xiphizusa Rchb.f.,
Bonplandia (Hannover) 2(9): 114. 1854.
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Figure 2. Lankester Composite Digital Plate (LCDP) of Stelis sp. A. Habit. B. Flower. C. Dissected perianth. D. Column
with lip, lateral view. E. Column ventral and lateral view. F. Anther cap. G. Pollinarium. Photographs by AK and I.
Chinchilla based on Karremans 7293 (JBL-spirit).
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Figure 3. LCDP of a typical Stelis s.s. species. A. Habit. B. Flower. C. Dissected perianth. D. Column with lip, lateral view.
E. Inorescence. F. Lip. G. Anther cap and pollinarium. Photographs by J.S. Moreno based on Moreno 519 (CAUP).
Syn. Pseudostelis Schltr., Anexos Mem.
Inst. Butantan, Secç. Bot. 1(4): 36. 1922.
Pleurothallis subgen. Pseudostelis (Schltr.)
Luer, Monogr. Syst. Bot. Missouri Bot. Gard.
76: 87. 1999. Lectotype: Physosiphon spiralis
Lindl., Edwards’s Bot. Reg. 21: sub t. 1797.
1835 (Garay 1974).
Stelis subgen. Crocodeilanthe includes 85 species,
divided in two sections. Stelis sect. Crocodeilanthe is
composed of 84 species of which 90% are found at high
elevations in the Andes of Bolivia, Colombia, Ecuador,
Peru and Venezuela. Many are local endemics. A few
species are known from Costa Rica and Panama, and
a single species is reported from the Greater Antilles,
another from the Lesser Antilles and yet another from
Brazil. Sect. Pseudostelis includes only one species,
the common and widespread Stelis deregularis Barb.
Rodr. which is found at mid elevations from Mexico to
Brazil, through Central America.
Toscano de Brito (2018a) recognizes Crocodei-
lanthe at the generic level suggesting it may be easily
dened by merging Luer’s Pleurothallis subgen.
Crocodeilanthe and Pleurothallis subgen Pseudostelis.
The species of Pseudostelis, excluding P. rufobrunnea
“which is clearly a member of the genus Stelis”, are
said to “share the same habit and oral morphology
with Crocodeilanthe”. Nevertheless, the placement of
Stelis magdalenae (Rchb.f.) Pridgeon & M.W.Chase
in Crocodeilanthe is not straight forward at all, and
Stelis simplex (Ames & C.Schweinf.) Pridgeon &
M.W.Chase certainly belongs to the Dracontia clade
rather than Crocodeilanthe. Also, even though it may
now seem obvious that Stelis rufobrunnea is in fact
a member of Stelis s.s. and that Stelis deregularis
belongs to Crocodeilanthe, both species were only
faithfully placed on the basis of DNA data.
Furthermore, the suggestion that Stelis gelida
(Lindl.) Pridgeon & M.W.Chase (type species of
Niphantha) belongs to Crocodeilanthe is not supported
genetically or morphologically. The subpandurate,
arcuate lip, elongate column, incumbent anther and
stigma, and whale-tail pollinaria clearly separate it from
other Crocodeilanthe species. Multiple Stelis gelida
accessions analyzed by Karremans et al. (2013) and
again by Pérez-Escobar et al. (2017), were consistently
found only distantly related to Crocodeilanthe. An
accession labeled Stelis antillensis in Karremans et al.
(2013), which was retrieved among those of S. gelida,
is likely misidentied by the original sequence author
(Stenzel) as was stated therein.
The unresolved relationships between Crocodei-
lanthe species and those of the non-monophyletic
Pseudostelis, in addition to the misplacement of
the unrelated Stelis gelida, are evidence that the
denition of this genus is not as straightforward as
suggested. Even though it is possible to recognize most
Crocodeilanthe species morphologically, the short lip
and petals, the stout column with an apical anther, and
pollina with a drop-like viscidium are a step prior to
the typical Stelis s.s. oral morphology. As closest
relative to Stelis in the strict sense, its recognition at
generic level entails the recognition of several other
splinter genera, which is not advised at the time.
Stelis subgen. Crocodeilanthe sect. Crocodeilanthe
Syn.: Crocodeilanthe Rchb.f. & Warsz., Bonplandia
(Hannover) 2: 113. 1854. Pleurothallis subgen.
Crocodeilanthe (Rchb.f. & Warsz.) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 20: 34.
1986. Type: Crocodeilanthe xiphizusa Rchb.f.,
Bonplandia (Hannover) 2(9): 114. 1854.
Stelis aligera (Luer & R.Vásquez) Pridgeon &
M.W.Chase, Lindleyana 16(4): 261. 2001.
Bas.: Pleurothallis aligera Luer & R.Vásquez,
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KArremAns — Stelis
Figure 4. Stelis hualluapampensis Collantes & Karremans,
a non-typical species of Stelis s.s. with subglobose
owers, fused lateral sepals and a lip-like dorsal sepal
that sticks out of the ower. Photograph by B. Collantes.
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Revista Soc. Boliv. Bot. 1(2): 9. 1997. Syn.: Crocodei-
lanthe aligera (Luer & R.Vásquez) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 256. 2004.
Stelis aloisii (Schltr.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 261. 2001.
Bas.: Pleurothallis aloisii Schltr., Repert. Spec. Nov.
Regni Veg. Beih. 8: 57. 1921. Syn.: Crocodeilanthe
aloisii (Schltr.) Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 95: 256. 2004.
Stelis antillensis Pridgeon & M.W.Chase, Lindleyana
17(2): 98. 2002.
Repl. syn.: Pleurothallis domingensis Cogn., Symb.
Antill. 6: 402. 1909. Syn.: Crocodeilanthe domingen-
sis (Cogn.) Luer, Monogr. Syst. Bot. Missouri Bot.
Gard. 95: 256. 2004. Stelis domingensis (Cogn.)
Pridgeon & M.W.Chase, Lindleyana 16: 262.
2001, nom. illeg. Non Stelis domingensis Cogn. in
I.Urban, Symb. Antill. 6: 692. 1910.
Stelis apposita (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 261. 2001.
Bas.: Pleurothallis apposita Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 65: 12. 1998. Syn.:
Crocodeilanthe apposita (Luer) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 256. 2004.
Stelis atwoodii (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 261. 2001. (Fig. 5)
Bas.: Pleurothallis atwoodii Luer, Lindleyana
11(2): 67. 1996. Syn.: Crocodeilanthe atwoodii
(Luer) Luer, Monogr. Syst. Bot. Missouri Bot.
Gard. 95: 256. 2004.
Stelis avirostris (Luer & Hirtz) Pridgeon & M.W.Chase,
Lindleyana 16(4): 261. 2001.
Bas.: Pleurothallis avirostris Luer & Hirtz, Monogr.
Syst. Bot. Missouri Bot. Gard. 65: 13. 1998. Syn.:
Crocodeilanthe avirostris (Luer & Hirtz) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 95: 256.
2004.
Stelis batillacea (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 261. 2001.
Bas.: Pleurothallis batillacea Luer, Selbyana 3(1-
2): 58. 1976. Syn.: Crocodeilanthe batillacea (Luer)
Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95:
256. 2004.
Stelis bracteosa (C.Schweinf.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 261. 2001.
Bas.: Pleurothallis bracteosa C.Schweinf.,
Fieldiana, Bot. 33: 20. 1970. Syn.: Crocodeilanthe
bracteosa (C.Schweinf.) Luer, Harvard Pap. Bot.
16(2): 358. 2011.
Stelis bucaramangae (Luer & R.Escobar) Pridgeon &
M.W.Chase, Lindleyana 16(4): 261. 2001.
Bas.: Pleurothallis bucaramangae Luer &
R.Escobar, Orquideología 20: 38. 1996. Syn.:
Crocodeilanthe bucaramangae (Luer & R.Escobar)
Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95:
256. 2004.
Stelis cassidis (Lindl.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 261. 2001.
Bas.: Pleurothallis cassidis Lindl., Ann. Mag. Nat.
Hist. 15: 384. 1845. Syn.: Crocodeilanthe cassidis
(Lindl.) Luer, Monogr. Syst. Bot. Missouri Bot.
Gard. 95: 256. 2004.
Figure 5. Stelis atwoodii (Luer) Pridgeon & M.W.Chase,
an untypical member of Stelis subgen. Crocodeilanthe.
Photograph by AK.
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Stelis cauliora (Lindl.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 261. 2001.
Bas.: Pleurothallis cauliora Lindl., Companion
Bot. Mag. 2: 355. 1836. Syn.: Crocodeilanthe
cauliora (Lindl.) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 256. 2004.
Stelis choerorhyncha (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 261-262. 2001.
Bas.: Pleurothallis choerorhyncha Luer,
Orquideología 20: 204. 1996. Syn.: Crocodeilanthe
choerorhyncha (Luer) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 256. 2004.
Stelis cosangae (Luer & Hirtz) Pridgeon & M.W.Chase,
Lindleyana 16(4): 262. 2001.
Bas.: Pleurothallis cosangae Luer & Hirtz, Monogr.
Syst. Bot. Missouri Bot. Gard. 65: 18. 1998. Syn.:
Crocodeilanthe cosangae (Luer & Hirtz) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 95: 256.
2004.
Stelis cuatrecasasii (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 262. 2001.
Bas.: Pleurothallis cuatrecasasii Luer, Orquideo-
logía 20: 208. 1996. Syn.: Crocodeilanthe
cuatrecasasii (Luer) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 256. 2004.
Stelis cyathiora (C.Schweinf.) Pridgeon &
M.W.Chase, Lindleyana 16(4): 262. 2001.
Bas.: Pleurothallis cyathiora C.Schweinf., Bot.
Mus. Lea. 15: 90, t. 27. 1951. Crocodeilanthe
cyathiora (C.Schweinf.) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 256. 2004.
Stelis damianii Karremans, nom. nov.
Repl. syn.: Crocodeilanthe chachapoyensis Damian,
Ann. Bot. Fenn. 56: 302. 2019. Non Stelis chachapo-
yensis Rchb.f., Bonplandia (Hannover) 3: 225. 1855.
Stelis dapsilis Pridgeon & M.W.Chase, Lindleyana
17(2): 99. 2002.
Repl. syn.: Pleurothallis maxima Luer, Selbyana
3(1-2): 140. 1976. Syn.: Crocodeilanthe maxima
(Luer) Luer, Monogr. Syst. Bot. Missouri Bot. Gard.
95: 256. 2004. Stelis maxima (Luer) Pridgeon &
M.W.Chase, Lindleyana 16: 264. 2001, nom. illeg.
Non Stelis maxima Lindl., Ann. Mag. Nat. Hist. 15:
106. 1845.
Stelis decurrens Pridgeon & M.W.Chase, Lindleyana
17(2): 99. 2002.
Repl. syn.: Pleurothallis croatii Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 65: 18-19. 1998.
Syn.: Crocodeilanthe croatii (Luer) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 256. 2004.
Stelis croatii (Luer) Pridgeon & M.W.Chase,
Lindleyana 16: 262. 2001, nom. illeg. Non
Stelis croatii Luer, Lindleyana 11: 97. 1996.
Stelis toscanoi Karremans, nom. nov.
Repl. syn.: Crocodeilanthe dewildei Luer &
Toscano (2018: 47). Stelis dewildei (Luer &
Toscano) Karremans, Phytotaxa 406(5): 265. 2019,
nom. illeg. Non Stelis dewildei Luer & R.Escobar,
Harvard Pap. Bot. 22(1): 34. 2017.
Stelis duckei E.M.Pessoa & M.Alves, Brittonia 66(2):
156-157. 2014.
Syn.: Crocodeilanthe duckei (E.M.Pessoa &
M.Alves) Toscano, Harvard Pap. Bot. 23(1): 54.
2018.
Stelis erectiora (Luer) J.M.H.Shaw, Orchid Rev.
122(1308): 77. 2014.
Bas.: Crocodeilanthe erectiora Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 103: 311. 2005.
Stelis expansa (Lindl.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 262. 2001.
Bas.: Pleurothallis expansa Lindl., Fol. Orchid.
~Pleurothallis~ 4. 1859. Syn.: Crocodeilanthe
expansa (Lindl.) Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 95: 256. 2004.
Stelis fons-orum (Lindl.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 263. 2001.
Bas.: Pleurothallis fons-orum Lindl., Fol. Orchid.
~Pleurothallis~ 5, no. 15. 1859. Syn.: Crocodeilanthe
fons-orum (Lindl.) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 256. 2004.
Stelis galeata (Lindl.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 263. 2001.
Bas.: Pleurothallis galeata Lindl., Ann. Mag. Nat.
Hist. 15: 107. 1845. Syn.: Crocodeilanthe galeata
(Lindl.) Luer, Monogr. Syst. Bot. Missouri Bot.
Gard. 95: 256. 2004.
Stelis galerasensis (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 263. 2001.
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Bas.: Pleurothallis galerasensis Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 65: 28. 1998. Syn.:
Crocodeilanthe galerasensis (Luer) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 256. 2004.
Stelis gargantua Pridgeon & M.W.Chase, Lindleyana
17(2): 99. 2002.
Repl. syn.: Pleurothallis gigas Luer & R.Escobar,
Orquideología 20(1): 52. 1996. Crocodeilanthe
gigas (Luer & R.Escobar) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 256. 2004. Stelis gigas (Luer
& R.Escobar) Pridgeon & M.W.Chase, Lindleyana
16: 263 (2001), nom. illeg. Non Stelis gigas Barb.
Rodr., Gen. Spec. Orchid. 2: 89. 1881.
Stelis globosa Pridgeon & M.W.Chase, Lindleyana
17(2): 99. 2002.
Repl. syn.: Pleurothallis popayanensis F.Lehm.
& Kraenzl., Bot. Jahrb. Syst. 26: 438. 1898.
Crocodeilanthe popayanensis (F.Lehm. &
Kraenzl.) Luer, Monogr. Syst. Bot. Missouri Bot.
Gard. 95: 256. 2004. Stelis popayanensis (F.Lehm.
& Kraenzl.) Pridgeon & M.W.Chase, Lindleyana
16: 265. 2001, nom. illeg. Non Stelis popayanensis
F.Lehm. & Kraenzl., Bot. Jahrb. Syst. 26(3-4): 448.
1899.
Stelis heros Karremans, nom. nov.
Repl. syn.: Crocodeilanthe steinbachii Luer &
Toscano, Harvard Pap. Bot. 23: 48. 2018. Stelis
steinbachii (Luer & Toscano) Karremans, Phytotaxa
406(5): 267. 2019, nom. illeg. Non Stelis steinbachii
Luer, Selbyana 32(1,2): 110. 2018.
etymology: The name honors the three larger-
than-life orchidologists that sadly passed away in
2019, Carl A. Luer, Robert L. Dressler and W. Mark
Whitten.
Stelis infundibulosa (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 263. 2001.
Bas.: Pleurothallis infundibulosa Luer,
Orquideología 20: 210. 1996. Syn.: Crocodeilanthe
infundibulosa (Luer) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 256. 2004.
Stelis jurisdixii (Luer & R.Escobar) Pridgeon &
M.W.Chase, Lindleyana 16(4): 264. 2001.
Bas.: Pleurothallis jurisdixii Luer & R.Escobar,
Orquideología 20: 64. 1996. Syn.: Crocodeilanthe
jurisdixii (Luer & R.Escobar) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 95: 256. 2004.
Stelis juxta (Luer, Thoerle & F.A.Werner) J.M.H.Shaw,
Orchid Rev. 122(1308): 77. 2014.
Bas.: Crocodeilanthe juxta Luer, Thoerle &
F.A.Werner, Harvard Pap. Bot. 16(2): 320. 2011.
Stelis laevigata (Lindl.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 264. 2001.
Bas.: Pleurothallis laevigata Lindl., Ann. Mag.
Nat. Hist. 15: 106. 1845. Crocodeilanthe laevigata
(Lindl.) Luer, Monogr. Syst. Bot. Missouri Bot.
Gard. 95: 256. 2004.
Stelis laevis (Luer & Hirtz) Pridgeon & M.W.Chase,
Lindleyana 16(4): 264. 2001.
Bas.: Pleurothallis laevis Luer & Hirtz, Monogr.
Syst. Bot. Missouri Bot. Gard. 65: 33. 1998. Syn.:
Crocodeilanthe laevis (Luer & Hirtz) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 256. 2004.
Stelis laminata (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 264. 2001.
Bas.: Pleurothallis laminata Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 65: 33-34. 1998. Syn.:
Crocodeilanthe laminata (Luer) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 256. 2004.
Stelis lehmanniana (Schltr.) Karremans, Phytotaxa
203(3): 293. 2015.
Bas.: Pleurothallis lehmanniana Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 7: 235. 1920.
Crocodeilanthe lehmanniana (Schltr.) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 95: 256.
2004.
Stelis ligulata (Lindl.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 264. 2001.
Bas.: Pleurothallis ligulata Lindl., Fol. Orchid.
~Pleurothallis~ 29. 1859. Syn.: Crocodeilanthe
ligulata (Lindl.) Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 95: 256. 2004.
Stelis magdalenae (Rchb.f.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 264. 2001.
Bas.: Pleurothallis magdalenae Rchb.f.,
Bonplandia (Hannover) 3: 72. 1855. Crocodeilanthe
magdalenae (Rchb.f.) Toscano, Harvard Pap. Bot.
23(1): 54. 2018.
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Stelis mandonii (Rchb.f.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 264. 2001.
Bas.: Pleurothallis mandonii Rchb.f., Xenia Orchid.
3: 24. 1878. Syn.: Crocodeilanthe mandonii
(Rchb.f.) Luer, Monogr. Syst. Bot. Missouri Bot.
Gard. 95: 256. 2004.
Stelis melanostele (Luer & R.Vásquez) Pridgeon &
M.W.Chase, Lindleyana 16(4): 264. 2001.
Bas.: Pleurothallis melanostele Luer & R.Vásquez,
Phytologia 49(3): 210. 1981. Syn.: Crocodeilanthe
melanostele (Luer & R.Vásquez) Toscano, Harvard
Pap. Bot. 23(1): 54. 2018.
Stelis mendietae (Luer, Thoerle & F.A.Werner)
J.M.H.Shaw, Orchid Rev. 122(1308): 77. 2014.
Bas.: Crocodeilanthe mendietae Luer, Thoerle &
F.A.Werner, Harvard Pap. Bot. 16(2): 321. 2011.
Stelis molleturoi (Luer & Dodson) Pridgeon &
M.W.Chase, Lindleyana 16(4): 264. 2001.
Bas.: Pleurothallis molleturoi Luer & Dodson,
Monogr. Syst. Bot. Missouri Bot. Gard. 65: 38.
1998. Crocodeilanthe molleturoi (Luer & Dodson)
Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95:
256. 2004.
Stelis moritzii (Rchb.f.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 264. 2001.
Bas.: Pleurothallis moritzii Rchb.f., Linnaea 22:
824. 1849. Syn.: Crocodeilanthe moritzii (Rchb.f.)
Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95:
256. 2004.
Stelis neowerneri J.M.H.Shaw, Orchid Rev.
122(1308): 78. 2014.
Bas.: Crocodeilanthe werneri Luer & Thoerle,
Harvard Pap. Bot. 16(2): 323. 2011. Non Stelis
werneri Schltr., Repert. Spec. Nov. Regni Veg. Beih.
27: 42. 1924.
Stelis nivalis (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 264-265. 2001.
Bas.: Pleurothallis nivalis Luer, Selbyana 1(4): 420.
1976. Syn.: Crocodeilanthe nivalis (Luer) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 95: 256.
2004.
Stelis orectopus (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 265. 2001.
Bas.: Pleurothallis orectopus Luer, Selbyana 3(3-4):
356. 1977. Syn.: Crocodeilanthe orectopus (Luer)
Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95:
256. 2004.
Stelis pachypus F.Lehm. & Kraenzl., Bot. Jahrb. Syst.
26: 447. 1899.
Bas.: Pleurothallis pachypus (F.Lehm. & Kraenzl.)
Garay, Canad. J. Bot. 34: 254. 1956. Syn.:
Crocodeilanthe pachypus (F.Lehm. & Kraenzl.)
Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95:
256. 2004.
Stelis patateënsis (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 265. 2001.
Bas.: Pleurothallis patateënsis Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 65: 42. 1998. Syn.:
Crocodeilanthe patateënsis (Luer) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 256. 2004.
Stelis pellucida (Luer & Hirtz) Pridgeon & M.W.Chase,
Lindleyana 16(4): 265. 2001.
Bas.: Pleurothallis pellucida Luer & Hirtz, Monogr.
Syst. Bot. Missouri Bot. Gard. 65: 42-43. 1998. Syn.:
Crocodeilanthe pellucida (Luer & Hirtz) Luer, Mo-
nogr. Syst. Bot. Missouri Bot. Gard. 95: 256. 2004.
Stelis pennelliana (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 265. 2001.
Bas.: Pleurothallis pennelliana Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 65: 43. 1998. Syn.:
Crocodeilanthe pennelliana (Luer) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 256. 2004.
Stelis pilifera (Lindl.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 265. 2001.
Bas.: Pleurothallis pilifera Lindl., Fol. Orchid.
~Pleurothallis~ 9. 1859. Syn.: Crocodeilanthe
pilifera (Lindl.) Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 95: 256. 2004.
Stelis pittieri (Schltr.) Rojas-Alv. & Karremans,
Phytotaxa 406(5): 266. 2019.
Bas.: Pleurothallis pittieri Schltr., Repert. Spec.
Nov. Regni Veg. 3(42-43): 247. 1907.
Syn.: Crocodeilanthe oribunda (Poepp. & Endl.)
Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95:
256. 2004. Pleurothallis oribunda Poepp. & Endl.,
Nov. Gen. Sp. Pl. 1: 48-49, t. 84. 1835 [1836]. Non
Stelis oribunda Kunth, Nov. Gen. Sp. (folio ed.) 1:
362. 1815 [1816].
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Stelis possoae (Luer) Karremans, Phytotaxa 203(3):
293. 2015.
Bas.: Pleurothallis possoae Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 79: 129-130. 2000.
Crocodeilanthe possoae (Luer) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 95: 256. 2004.
Stelis praealta (Luer & Hirtz) Pridgeon & M.W.Chase,
Lindleyana 16(4): 265. 2001.
Bas.: Pleurothallis praealta Luer & Hirtz, Monogr.
Syst. Bot. Missouri Bot. Gard. 65: 45. 1998. Crocodei-
lanthe praealta (Luer & Hirtz) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 95: 256. 2004.
Stelis prolicans (Luer & Hirtz) Pridgeon &
M.W.Chase, Lindleyana 16(4): 265. 2001.
Bas.: Pleurothallis prolicans Luer & Hirtz, Monogr.
Syst. Bot. Missouri Bot. Gard. 65: 45. 1998. Syn.:
Crocodeilanthe prolicans (Luer & Hirtz) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 95: 256.
2004.
Stelis pulchella Kunth, Nov. Gen. Sp. (quarto ed.)
1(4): 364, t. 90. 1816. (Fig. 6)
Syn.: Pleurothallis pulchella (Kunth) Lindl., Exot.
Fl. 2(14): sub t. 123. 1825 [1824]. Crocodeilanthe
pulchella (Kunth) Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 95: 256. 2004.
Stelis reptans Pridgeon & M.W.Chase, Lindleyana
17(2): 100. 2002.
Bas.: Pleurothallis scansor Luer, Phytologia 49(3):
216. 1981. Syn.: Crocodeilanthe scansor (Luer)
Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95:
257. 2004. Non Stelis scansor Rchb.f., Bonplandia
(Hannover) 3(17): 241. 1855.
Stelis retusiloba (C.Schweinf.) Pridgeon &
M.W.Chase, Lindleyana 16(4): 266. 2001.
Bas.: Pleurothallis retusiloba C.Schweinf., Bot.
Mus. Lea. 15: 100. 1951. Syn.: Crocodeilanthe
retusiloba (C.Schweinf.) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 256. 2004.
Stelis rhodotantha (Rchb.f.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 266. 2001.
Bas.: Pleurothallis rhodotantha Rchb.f., Linnaea
22: 825. 1849. Syn.: Crocodeilanthe rhodotantha
(Rchb.f.) Luer, Monogr. Syst. Bot. Missouri Bot.
Gard. 95: 257. 2004.
Stelis rictoria (Rchb.f.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 266. 2001.
Bas.: Pleurothallis rictoria Rchb.f., Linnaea 41:
14. 1877. Crocodeilanthe rictoria (Rchb.f.) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 95: 257.
2004.
Stelis roseopunctata (Lindl.) R. Bernal, Phytoneuron
22: 5. 2015.
Bas.: Pleurothallis roseopunctata Lindl., Orchid.
Linden. 2. 1846.
Syn.: Dendrobium elegans Kunth, Nov. Gen.
Sp. 1: 358. 1816. Pleurothallis elegans (Kunth)
Lindl., Edwards’s Bot. Reg. 28(Misc): 70. 1842.
Crocodeilanthe elegans (Kunth) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 95: 256. 2004. Non Stelis
elegans Luer & R.Vásquez, Phytologia 49(3): 228.
1981.
Stelis rostriformis Zambrano & Solano, Phytotaxa
376(4): 181. 2018.
Stelis sagittata Zambrano & Solano, Phytotaxa 376(4):
183. 2018.
Repl. syn.: Pleurothallis jamiesonii Lindl.,
Edwards’s Bot. Reg. 21: sub t. 1797. 1835. Syn.:
Crocodeilanthe jamiesonii (Lindl.) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 256. 2004.
Stelis jamiesonii (Lindl.) Pridgeon & M.W.Chase,
Lindleyana 16: 264. 2001, nom. illeg. Non Stelis
jamesonii Lindl., J. Bot. (Hooker) 1: 11. 1834.
Stelis salpingantha (Luer & Hirtz) Pridgeon &
M.W.Chase, Lindleyana 16(4): 266. 2001.
Bas.: Pleurothallis salpingantha Luer & Hirtz,
Monogr. Syst. Bot. Missouri Bot. Gard. 65: 50-51.
1998. Syn.: Crocodeilanthe salpingantha (Luer &
Hirtz) Luer, Monogr. Syst. Bot. Missouri Bot. Gard.
95: 257. 2004.
Stelis simplicilabia (C.Schweinf.) Pridgeon &
M.W.Chase, Lindleyana 16(4): 266. 2001.
Bas.: Pleurothallis simplicilabia C.Schweinf.,
Revista Acad. Colomb. Ci. Exact. 5(19): 350. 1943.
Syn.: Crocodeilanthe simplicilabia (C.Schweinf.)
Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95:
257. 2004.
Stelis siphonantha (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 266. 2001.
303
KArremAns — Stelis
Figure 6. LCDP of Stelis pulchella a typical representative of Stelis sect. Crocodeilanthe. A. Habit. B. Inorescence. C.
Flowers. D. Dissected perianth. E. Column ventral and lateral view. F. Lip naturally and expanded. G. Anther cap and
pollinarium. Photographs by AK based on JBL-28245 (JBL-spirit).
LANKESTERIANA 19(3). 2019. © Universidad de Costa Rica, 2019.
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Bas.: Pleurothallis siphonantha Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 65: 52-53.
1998. Crocodeilanthe siphonantha (Luer) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 95: 257.
2004.
Stelis spathosa (Luer & R.Escobar) Pridgeon &
M.W.Chase, Lindleyana 16(4): 266. 2001.
Bas.: Pleurothallis spathosa Luer & R.Escobar,
Orquideología 20: 86. 1996. Syn.: Crocodeilanthe
spathosa (Luer & R.Escobar) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 95: 257. 2004.
Stelis speciosa (Luer, Thoerle & F.A.Werner)
E.M.Pessoa & M. Alves, Brittonia 66(2): 157. 2013.
Bas.: Crocodeilanthe speciosa Luer, Thoerle & F.A.
Werner, Harvard Pap. Bot. 16(2): 321. 2011. Syn.:
Stelis speciosa (Luer, Thoerle & F.A.Werner)
J.M.H.Shaw, Orchid Rev. 122(1308): 78. 2014,
nom. illeg.
Stelis stelidiopsis (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 266-267. 2001.
Bas.: Pleurothallis stelidiopsis Luer, Phytologia
49(3): 218. 1981. Syn.: Crocodeilanthe stelidiopsis
(Luer) Luer, Monogr. Syst. Bot. Missouri Bot. Gard.
95: 257. 2004.
Stelis stergiosii (Carnevali & I.Ramírez) Karremans,
Phytotaxa 203(3): 293. 2015.
Bas.: Pleurothallis stergiosii Carnevali & I.Ramírez,
Harvard Pap. Bot. 3: 247. 1998. Syn.: Crocodeilanthe
stergiosii (Carnevali & I.Ramírez) Carnevali &
I.Ramírez, Nuevo Cat. Fl. Vasc. Venezuela 578.
2008.
Stelis suinii (Luer) J.M.H.Shaw, Orchid Rev.
122(1308): 78. 2014.
Bas.: Crocodeilanthe suinii Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 105: 249. 2006. Syn.:
Pleurothallis suinii (Luer) Pfahl, Internet Orchid Sp.
Photo Encycl. Nomencl. Notes 2. 2013. 2013.
Stelis taxis (Luer) Pridgeon & M.W.Chase, Lindleyana
16(4): 267. 2001.
Bas.: Pleurothallis taxis Luer, Selbyana 5(2): 184.
1979. Crocodeilanthe taxis (Luer) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 257. 2004.
Stelis tepuiensis (Carnevali & I.Ramírez) Karremans,
Phytotaxa 203(3): 294. 2015.
Bas.: Pleurothallis tepuiensis Carnevali & I.Ramírez,
Novon 3(2): 121. 1993. Syn.: Crocodeilanthe
tepuiensis (Carnevali & I.Ramírez) Carnevali &
I.Ramírez, Nuevo Cat. Fl. Vasc. Venezuela 758.
2008.
Stelis tunguraguae (F.Lehm. & Kraenzl.) Pridgeon &
M.W.Chase, Lindleyana 16(4): 267. 2001.
Bas.: Pleurothallis tunguraguae F.Lehm. &
Kraenzl., Bot. Jahrb. Syst. 26: 439. 1899. Syn.:
Crocodeilanthe tunguraguae (F.Lehm. & Kraenzl.)
Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95:
257. 2004.
Stelis uvaegelata Doucette ex L.E.Matthews,
OrchideenJ. 6(3): 13. 2018.
Stelis vargasii (C.Schweinf.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 267. 2001.
Bas.: Pleurothallis vargasii C.Schweinf., Bot. Mus.
Lea. 10: 192. 1942. Syn.: Crocodeilanthe vargasii
(C.Schweinf.) Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 95: 257. 2004.
Stelis vasqueziana Karremans, Phytotaxa 203(3): 294.
2015.
Bas.: Crocodeilanthe vasquezii Luer, Harvard Pap.
Bot. 17(2): 340. 2012.
Stelis vegrandis (Luer & Dodson) Pridgeon &
M.W.Chase, Lindleyana 16(4): 267. 2001.
Bas.: Pleurothallis vegrandis Luer & Dodson,
Monogr. Syst. Bot. Missouri Bot. Gard. 65: 56. 1998.
Syn.: Crocodeilanthe vegrandis (Luer & Dodson)
Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95:
257. 2004.
Stelis velaticaulis (Rchb.f.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 267. 2001.
Bas.: Pleurothallis velaticaulis Rchb.f., Linnaea
22: 824. 1849. Syn.: Crocodeilanthe velaticaulis
(Rchb.f.) Luer, Monogr. Syst. Bot. Missouri Bot.
Gard. 95: 257. 2004.
Stelis velatipes (Rchb.f.) Karremans, Phytotaxa
406(5): 267. 2019.
Bas.: Pleurothallis velatipes Rchb.f., Linnaea 22:
828. 1849. Syn.: Crocodeilanthe velatipes (Rchb.f.)
Carnevali & G.A.Romero, Nuevo Cat. Fl. Vasc.
Venezuela 758. 2008.
305
KArremAns — Stelis
Stelis verbiformis (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 267. 2001.
Bas.: Pleurothallis verbiformis Luer, Selbyana 2:
389. 1978. Crocodeilanthe verbiformis (Luer) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 95: 257.
2004.
Stelis virgata (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 267. 2001.
Bas.: Pleurothallis virgata Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 65: 59. 1998. Crocodeilanthe
virgata (Luer) Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 95: 257. 2004.
Stelis weddelliana (Rchb.f.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 267. 2001.
Bas.: Pleurothallis weddelliana Rchb.f., Xenia
Orchid. 3: 24. 1878. Crocodeilanthe weddelliana
(Rchb.f.) Luer, Monogr. Syst. Bot. Missouri Bot.
Gard. 103: 309. 2005.
Stelis xiphizusa (Rchb.f.) Pridgeon & M.W.Chase,
Lindleyana 16: 268. 2001.
Bas.: Crocodeilanthe xiphizusa Rchb.f., Bonplandia
(Hannover) 2(9): 114. 1854. Syn.: Pleurothallis
xiphizusa (Rchb.f.) Rchb.f., Ann. Bot. Syst. 6(2):
172-173. 1861.
Stelis zunagensis (Luer & Hirtz) Pridgeon &
M.W.Chase, Lindleyana 16(4): 268. 2001.
Bas.: Pleurothallis zunagensis Luer & Hirtz,
Monogr. Syst. Bot. Missouri Bot. Gard. 65: 61. 1998.
Syn.: Crocodeilanthe zunagensis (Luer & Hirtz)
Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95:
257. 2004.
DNA data is available for Stelis atwoodi, S.
galeata, S. pulchella, S. velaticaulis; the latter species
are morphologically highly similar to Stelis xiphizusa,
type species of Crocodeilanthe, of which no DNA
data is currently available. There is no doubt that all
typical species of Crocodeilanthe belong here. They
consistently group together into a well supported clade
that is very closely related to Stelis in the strict sense
(Pridgeon et al. 2001, Solano-Gómez 2005, Karremans
et al. 2013, Pérez-Escobar et al. 2017, Wilson et al.
2017, Ponert et al. 2019). This is consistent with the
Stelis-like morphology of their ower, especially in an
overall reduction in the column and lip, and pollinaria
with a viscidium.
Stelis subgen. Crocodeilanthe sect. Pseudostelis
(Schltr.) Karremans, comb. et stat. nov.
Bas.: Pseudostelis Schltr., Anexos Mem. Inst.
Butantan, Secç. Bot. 1(4): 36. 1922. Syn.
Pleurothallis subgen. Pseudostelis (Schltr.) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 76: 87.
1999. Lectotype: Physosiphon spiralis Lindl.,
Edwards’s Bot. Reg. 21: sub t. 1797. 1835 (Garay
1974).
Stelis deregularis Barb.Rodr., Gen. Sp. Orchid. 2: 94.
1882. (Fig. 7)
Syn.: Physosiphon deregularis (Barb.Rodr.)
Cogn., Fl. Bras. 3(4): 341-342. 1896. Pseudostelis
deregularis (Barb.Rodr.) Schltr., Anexos Mem. Inst.
Butantan, Secc. Bot. 1(4): 38. 1922. Pleurothallis
deregularis (Barb.Rodr.) Luer, Selbyana 2(4): 385-
386. 1978.
Syn.: Physosiphon spiralis Lindl., Edwards’s Bot.
Reg. 21: sub t. 1797. 1835. Crocodeilanthe spiralis
(Lindl.) Toscano, Harvard Pap. Bot. 23(1): 54. 2018.
Pseudostelis spiralis (Lindl.) Schltr., Anexos Mem.
Inst. Butantan, Secc. Bot. 1(4): 38. 1922. Non Stelis
spiralis (Ruiz & Pav.) Pers., Syn. Pl. 2: 524. 1807.
DNA studies consistently nd the accessions of
Stelis deregularis as sister to the remaining species
of Stelis subgen. Crocodeilanthe (Karremans et al.
2013, Pérez-Escobar et al. 2017). No DNA data is
available for Stelis bracteosa, S. magdalenae and S.
melanostele which have been suggested to be close
relatives of S. deregularis (Luer 1999, Toscano de
Brito 2018a). Based on morphology Stelis bracteosa
and S. melanostele are consistent with Stelis subgen.
Crocodeilanthe but their afnity with S. deregularis
is not as clear. They, together with the unresolved
Stelis magdalenae, are excluded from this section until
proven to belong here.
Stelis subgen. Physothallis (Garay) Karremans, comb.
nov.
Bas. Physothallis Garay, Svensk Bot. Tidskr. 47:
199. 1953. Pleurothallis subgen. Physothallis
(Garay) Luer, Monogr. Syst. Bot. Missouri Bot.
Gard. 20: 53. 1986. Type: Physothallis harlingii
Garay, Svensk Bot. Tidskr. 47(2): 199. 1953.
Syn. Pleurothallis subgen. Acuminatia Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 76: 98.
LANKESTERIANA 19(3). 2019. © Universidad de Costa Rica, 2019.
Figure 7. LCDP of Stelis deregularis, type species of Stelis sect. Pseudostelis. A. Habit. B. Flower. C. Dissected perianth.
D. Column with lip, lateral view. E. Column ventral view. F. Anther cap. G. Pollinarium on the stigma. Photographs by
AK based on Karremans 7303 (JBL-spirit).
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1999. Type: Dendrobium acuminatum Kunth,
Nov. Gen. Sp. (quarto ed.) 1: 357. 1816.
The 31 species that belong to Stelis subgen.
Physothallis are mostly found at high elevations in
the Andes of Bolivia, Colombia, Ecuador, Peru and
Venezuela. They are divided in three sections. Stelis
sect. Acuminatae includes 27 species that generally
have rather narrow distributions in the Andean
countries, especially Bolivia and Peru, a single species
from Central America, and a couple are reported from
Mexico, Guyana and Brazil. The three members of
Stelis sect. Physothallis are endemic to Ecuador. The
sole member of Stelis sect. Rubens is widely distributed
from Colombia to Bolivia and Brazil.
Despite being consistently found to be closely
related to Stelis s.s. and Crocodeilanthe in every
single phylogenetic study of the group (Karremans
2010, Chiron et al. 2012, Ramos-Castro et al.
2012, Karremans et al. 2013, Pérez-Escobar et
al. 2017, Wilson et al. 2017), the inclusion of the
members of Luer’s Pleurothallis sect. Acuminatae
in Stelis s.l. is still met with inexplicable resistance
(e.g. Santos et al. 2018, 2019, Toscano de Brito
2018b, Govaerts et al. 2019). The exclusion of
these species from genus Anathallis is not only
evident morphologically (Karremans 2014), but
is highly supported even in multi-gene genomic
studies (Ponert et al. 2019). That they belong
within a broadly defined Stelis is indisputable.
What remains to be proven at this time is how these
species interrelate as the analyses are inconclusive
and the groupings proposed may be artificial. Three
different clades are brought together: a) composed
of the two species previously placed in genus
Physothallis, plus Stelis lennartii (= Pleurothallis
anderssonii Luer); b) the controversial Stelis
montserratii (= Pleurothallis rubens Lindl.); and
c) the remaining members of Pleurothallis sect.
Acuminatae, including the type species Stelis
aurea [= Pleurothallis acuminata (Kunth) Lindl.].
Each one is given sectional status.
The recognition of Stelis s.s. and Crocodeilanthe
at generic level necessarily entails the recognition
of Physothallis and probably of Pleurothallis sect.
Acuminatae at generic level as well. This can only be
done after resolving how the members of these two
groups interrelate. .
Stelis subgen. Physothallis sect. Acuminatae (Lindl.)
Karremans, comb. nov.
Bas. Pleurothallis sect. Acuminatae Lindl. Fol.
Orchid. Pleurothallis 32. 1859. Type: Dendrobium
acuminatum Kunth, Nov. Gen. Sp. (quarto ed.) 1:
357. 1816.
Stelis ariasii (Luer & Hirtz) Karremans, Lankesteriana
13(3): 328. 2014.
Bas.: Pleurothallis ariasii Luer & Hirtz, Lindleyana
12(1): 42. 1997. Syn.: Anathallis ariasii (Luer &
Hirtz) Pridgeon & M.W.Chase, Lindleyana 16(4):
247. 2001.
Stelis asperilinguis (Rchb.f. & Warsz.) Karremans,
Lankesteriana 13(3): 328. 2014.
Bas.: Pleurothallis asperilinguis Rchb.f. & Warsz.,
Bonplandia (Hannover) 2: 114. 1854. Syn.: Anathal-
lis asperilinguis (Rchb.f. & Warsz.) Pridgeon &
M.W.Chase, Lindleyana 16(4): 247. 2001.
Stelis aurea (Lindl.) Karremans, Lankesteriana 13(3):
328. 2014. (Fig. 8)
Bas.: Pleurothallis aurea Lindl., Ann. Mag. Nat.
Hist. 12(79): 397. 1843.
Syn.: Anathallis racemosa Barb.Rodr., Gen. Sp.
Orchid. 1: 24. 1877. Syn.: Pleurothallis racemosa
(Barb.Rodr.) Cogn., Fl. Bras. 3(4): 554. 1896.
Syn.: Dendrobium acuminatum Kunth, Nov. Gen.
Sp. 1: 357. 1816. Anathallis acuminata (Kunth)
Pridgeon & M.W.Chase, Lindleyana 16(4): 247.
2001. Pleurothallis acuminata (Kunth) Lindl.,
Edwards’s Bot. Reg. 28(Misc.): 70, no. 13. 1842.
Non Stelis acuminata Luer & Hirtz, Monogr. Syst.
Bot. Missouri Bot. Gard. 95: 141. 2004.
Stelis bevilacquana (Carnevali & I.Ramírez)
Karremans, Phytotaxa 406(5): 265. 2019.
Bas.: Pleurothallis bevilacquana Carnevali &
I.Ramírez, Orchids Venez. (ed. 2) 1141. 2000.
Stelis candida (Luer & Hirtz) Karremans,
Lankesteriana 13(3): 328. 2014.
Bas.: Pleurothallis candida Luer & Hirtz, Monogr.
Syst. Bot. Missouri Bot. Gard. 76: 107. 1999.
Anathallis candida (Luer & Hirtz) Pridgeon &
M.W.Chase, Lindleyana 16(4): 248. 2001.
Stelis catenata Karremans, Lankesteriana 13(3): 328.
2014.
307
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Figure 8. LCDP of Stelis aurea, type species of Stelis sect. Acuminatae. A. Habit. B. Flower. C. Dissected perianth. D.
Column with lip, lateral view. E. Column ventral and lateral view. F. Lip. G. Anther cap and pollinarium. Photographs
by J.S. Moreno based on Moreno 520 (CAUP).
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Repl. syn.: Pleurothallis ramulosa Lindl., Fol.
Orchid. ~Pleurothallis~ 33. 1859. Syn.: Anathallis
ramulosa (Lindl.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 250. 2001. Non Stelis ramulosa
Luer & Dalström, Monogr. Syst. Bot. Missouri Bot.
Gard. 95: 169. 2004.
Stelis coripatae (Luer & R.Vásquez) Karremans,
Lankesteriana 13(3): 328. 2014.
Bas.: Pleurothallis coripatae Luer & R.Vásquez,
Phytologia 46(6): 362. 1980. Syn.: Anathallis
coripatae (Luer & R.Vásquez) Pridgeon &
M.W.Chase, Lindleyana 16(4): 248. 2001.
Stelis dimidia (Luer) Karremans, Lankesteriana 13(3):
328. 2014.
Bas.: Pleurothallis dimidia Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 76: 109. 1999. Syn.: Anathallis
dimidia (Luer) Pridgeon & M.W.Chase, Lindleyana
16(4): 248. 2001.
Stelis jesupiorum (Luer & Hirtz) Karremans,
Lankesteriana 13(3): 329. 2014.
Bas.: Pleurothallis jesupiorum Luer & Hirtz,
Lindleyana 11(3): 164. 1996. Syn.: Anathallis
jesupiorum (Luer & Hirtz) Pridgeon & M.W.Chase,
Lindleyana 16(4): 249. 2001.
Stelis lagarophyta (Luer) Karremans, Lankesteriana
13(3): 329. 2014.
Bas.: Pleurothallis lagarophyta Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 76: 112-113. 1999.
Syn.: Anathallis lagarophyta (Luer) Pridgeon &
M.W.Chase, Lindleyana 16(4): 249. 2001.
Stelis lauta Karremans, Lankesteriana 13(3): 329.
2014.
Repl. syn.: Pleurothallis concinna Luer &
R.Vásquez, Revista Soc. Boliv. Bot. 2(2): 133. 1999.
Syn.: Anathallis concinna (Luer & R.Vásquez)
Pridgeon & M.W.Chase, Lindleyana 16(4): 248.
2001. Non Stelis concinna Lindl., J. Bot. (Hooker)
1: 11. 1834.
Stelis maguirei (Luer) Karremans, Lankesteriana
13(3): 329. 2014.
Bas.: Pleurothallis maguirei Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 76: 113. 1999. Anathallis
maguirei (Luer) Pridgeon & M.W.Chase, Lindleyana
16(4): 249. 2001.
Stelis mediocarinata (C.Schweinf.) Karremans,
Lankesteriana 13(3): 329. 2014.
Bas.: Pleurothallis mediocarinata C.Schweinf.,
Fieldiana, Bot. 33: 26. 1970. Syn.: Anathallis
mediocarinata (C.Schweinf.) Pridgeon &
M.W.Chase, Lindleyana 16(4): 249. 2001.
Stelis melanopus (F.Lehm. & Kraenzl.) Karremans,
Lankesteriana 13(3): 329. 2014.
Bas.: Pleurothallis melanopus F.Lehm. & Kraenzl.,
Bot. Jaarb. 26: 443. 1899.
Syn.: Pleurothallis stenophylla F.Lehm. &
Kraenzl., Bot. Jahrb. Syst. 26: 442. 1899. Anathallis
stenophylla (F.Lehm. & Kraenzl.) Pridgeon &
M.W.Chase, Lindleyana 16(4): 251. 2001. Non
Stelis stenophylla Rchb.f., Bonplandia (Hannover)
3: 70. 1855.
Stelis meridana (Rchb.f.) Karremans, Lankesteriana
13(3): 329. 2014.
Bas.: Pleurothallis meridana Rchb.f., Linnaea 22:
826. 1849. Syn.: Anathallis meridana (Rchb.f.)
Pridgeon & M.W.Chase, Lindleyana 16(4): 249.
2001.
Stelis papuligera (Schltr.) Karremans, Lankesteriana
13(3): 329. 2014.
Bas.: Pleurothallis papuligera Schltr., Repert. Spec.
Nov. Regni Veg. 10: 453. 1912. Syn.: Anathallis
papuligera (Schltr.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 250. 2001.
Stelis peruviana Damián & Karremans, Systematic
Botany 41(2): 293. 2016.
Stelis poasensis (Ames) Chinchilla & Karremans,
Phytotaxa 406(5): 266. 2019. (Fig. 9)
Bas.: Pleurothallis poasensis Ames, Sched. Orch. 1:
10-11. 1922.
Syn.: Pleurothallis dolichopus Schltr., Repert. Spec.
Nov. Regni Veg. 10(257-259): 394. 1912. Anathallis
dolichopus (Schltr.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 248. 2001. Non Stelis dolichopus
Schltr., Orchis 6: 63. 1912.
Syn.: Pleurothallis lamprophylla Schltr., Repert.
Spec. Nov. Regni Veg. 15(427-433): 205-206.
1918, nom. illeg. Stelis lamprophylla Karremans,
Lankesteriana 13(3): 329. 2014, nom. nov. Non
Pleurothallis lamprophyllum G.Nicholson, Ill. Dict.
Gard., Cent. Suppl. 608. 1901.
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Syn.: Pleurothallis peregrina Ames, Sched. Orch. 6:
67-68. 1923.
Stelis regalis (Luer) Karremans, Lankesteriana 13(3):
329. 2014.
Bas.: Pleurothallis regalis Luer, Selbyana 5(2): 178.
1979. Syn.: Anathallis regalis (Luer) Pridgeon &
M.W.Chase, Lindleyana 16(4): 250. 2001.
Stelis scariosa (Lex.) Karremans, Lankesteriana 13(3):
330. 2014.
Bas.: Dendrobium scariosum Lex., Nov. Veg. Descr.
2(Orchid. Opusc.): 39-40. 1825. Syn.: Pleurothallis
scariosa (Lex.) Lindl., Edwards’s Bot. Reg. 28:
Misc. 71. 1842. Anathallis scariosa (Lex.) Pridgeon
& M.W.Chase, Lindleyana 16(4): 250. 2001.
Stelis schlimii (Luer) Karremans, Lankesteriana 13(3):
330. 2014.
Bas.: Pleurothallis schlimii Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 76: 120. 1999. Syn.: Anathallis
schlimii (Luer) Pridgeon & M.W.Chase, Lindleyana
16(4): 250. 2001.
Stelis sclerophylla (Lindl.) Karremans, Lankesteriana
13(3): 330. 2014.
Bas.: Pleurothallis sclerophylla Lindl., Edwards’s
Bot. Reg. 21, sub. t. 1797 no. 31. 1835. Anathallis
sclerophylla (Lindl.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 250. 2001.
Stelis soratana (Rchb.f.) Karremans, Lankesteriana
13(3): 330. 2014.
Bas.: Pleurothallis soratana Rchb.f., Xenia Orchid.
3: 25. 1878. Syn.: Anathallis soratana (Rchb.f.)
Pridgeon & M.W.Chase, Lindleyana 16(4): 250. 2001.
Stelis spathilabia (Schltr.) Karremans, Lankesteriana
13(3): 330. 2014.
Bas.: Pleurothallis spathilabia Schltr., Repert. Spec.
Nov. Regni Veg. Beih. 27: 56. 1924. Syn.: Anathallis
spathilabia (Schltr.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 251. 2001.
Stelis spathuliformis (Luer & R.Vásquez) Karremans,
Lankesteriana 13(3): 330. 2014.
Bas.: Pleurothallis spathuliformis Luer &
R.Vásquez, Revista Soc. Boliv. Bot. 2(2): 137. 1999.
Syn.: Anathallis spathuliformis (Luer & R.Vásquez)
Pridgeon & M.W.Chase, Lindleyana 16(4): 251.
2001.
Stelis unduavica (Luer & R.Vásquez) Karremans,
Lankesteriana 13(3): 330. 2014.
Bas.: Pleurothallis unduavica Luer & R.Vásquez,
Phytologia 46(6): 372. 1980. Syn.: Anathallis
unduavica (Luer & R.Vásquez) Pridgeon &
M.W.Chase, Lindleyana 16(4): 251. 2001.
Stelis vasquezii (Luer) Karremans, Lankesteriana
13(3): 330. 2014.
Bas.: Pleurothallis vasquezii Luer, Phytologia 49(3):
220. 1981. Anathallis vasquezii (Luer) Pridgeon &
M.W.Chase, Lindleyana 16(4): 251. 2001.
DNA data is available for several species belonging
to this group, including the type of the section Stelis
aurea (as Anathallis angustipetala), as well as S. dimidia,
S. jesupiorum, S. poasensis (as Anathallis dolichopus
and S. lamprophylla), S. sclerophylla (as Anathallis
sclerophylla). They consistently group together into a
Figure 9. Stelis poasensis (Ames) Chinchilla & Karremans,
a typical species of Stelis sect. Acuminatae. Photograph
by AK.
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well supported clade that is closely related to Stelis in
the strict sense (Karremans et al. 2013, Pérez-Escobar et
al. 2017, Wilson et al. 2017, Ponert et al. 2019).
Stelis subgen. Physothallis sect. Rubens Karremans,
sect. nov.
Type: Pleurothallis rubens Lindl., Edwards’s Bot.
Reg. 21: pl. 1797, no. 32. 1836.
Distinguished from sect. Acuminatae by the
subpandurate lip, the long-cucullate, pointed apex of
the column.
Stelis montserratii (Porsch) Karremans, Lankesteriana
13(3): 329. 2014. (Fig. 10)
Bas.: Pleurothallis montserratii Porsch, Oesterr.
Bot. Z. 158. 1905.
Syn.: Pleurothallis rubens Lindl., Edwards’s Bot.
Reg. 21: pl. 1797, no. 32. 1836. Specklinia rubens
(Lindl.) F.Barros, Hoehnea 10: 110. 1984. Anathallis
rubens (Lindl.) Pridgeon & M.W.Chase, Lindleyana
16(4): 250. 2001. Specklinia rubens (Lindl.) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 95: 263.
2004. Stelis neorubens Chiron, Phytotaxa 46: 55.
2012. Non Stelis rubens Schltr., Repert. Spec. Nov.
Regni Veg. 8(191-195): 564. 1910.
Syn.: Anathallis amblyopetala (Schltr.) Pridgeon
& M.W. Chase, Lindleyana 16(4): 247. 2001.
Pleurothallis amblyopetala Schltr., Repert. Spec.
Nov. Regni Veg. 12: 486. 1913.
Syn.: Pleurothallis excisa C.Schweinf., Bot. Mus.
Lea. 16: 48. 1953.
DNA studies conrm that the species previously
known as Pleurothallis rubens is closely related to Stelis
in the strict sense (Chiron et al. 2012, Ramos-Castro et
al. 2012, Karremans et al. 2013, Pérez-Escobar et al.
2017). It is yet unclear how it relates to other members
of Stelis subgen. Physothallis and until that is resolved it
is recognized as a distinct lineage within the group.
Stelis subgen. Physothallis sect. Physothallis
Stelis cylindrica (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 262. 2001.
Bas.: Physothallis cylindrica Luer, Selbyana 3(3-
4): 224. 1977. Pleurothallis cylindrica (Luer) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 20: 53.
1986.
Stelis harlingii (Garay) Pridgeon & M.W.Chase,
Lindleyana 16(4): 263. 2001. (Fig. 11)
Bas.: Physothallis harlingii Garay, Svensk Bot.
Tidskr. 47(2): 199-202. 1953. Pleurothallis
neoharlingii Luer, Monogr. Syst. Bot. Missouri Bot.
Gard. 20: 53. 1986.
Stelis lennartii Karremans, Lankesteriana 13(3): 329.
2014.
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Figure 10. Stelis montserratii (Porsch) Karremans, type
species of Stelis sect. Rubens. Photograph by J.
Meijvogel.
Figure 11. Stelis harlingii (Garay) Pridgeon & M.W.Chase,
type species of Stelis subgen. Physothallis. Photograph
by E. Hunt.
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Figure 12. LCDP of Stelis gelida, type species of Stelis subgen. Niphantha. A. Habit. B. Flower. C. Dissected perianth. D.
Column with lip, lateral view. E. Lip, three views. F. Column, three quarters view. G. Pollinarium and anther cap, two
views. Photographs by G. Rojas-Alvarado based on Díaz-Morales 216 (JBL-spirit).
Repl. syn.: Pleurothallis anderssonii Luer,
Lindleyana 11(3): 145. 1996. Anathallis anderssonii
(Luer) Pridgeon & M.W.Chase, Lindleyana 16(4):
247. 2001. Non Stelis anderssonii Luer & Endara,
Monogr. Syst. Bot. Missouri Bot. Gard. 112. 2007.
Despite their morphological appearances,
accessions of Stelis harlingii and Stelis lennartii form
a well supported clade that appears to be somehow
related to the other members of Stelis subgen.
Physothallis (Karremans et al. 2013, Pérez-Escobar et
al. 2017). Altogether they are sisters of Stelis subgen.
Stelis and Stelis subgen. Crocodeilanthe (Ramos-
Castro et al. 2012, Karremans et al. 2013, Pérez-
Escobar et al. 2017, Ponert et al. 2019)
Stelis subgen. Niphantha (Luer) Karremans, comb. nov.
Bas.: Niphantha Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 120: 154. 2010. Type: Pleurothallis gelida
Lindl., Edwards’s Bot. Reg. 27: Misc. 91. 1841.
Stelis gelida (Lindl.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 263. 2001. (Fig. 12)
Bas.: Pleurothallis gelida Lindl., Edwards’s Bot.
Reg. 27: Misc. 91. 1841. Syn.: Specklinia gelida
(Lindl.) Luer, Monogr. Syst. Bot. Missouri Bot.
Gard. 95: 260. 2004. Niphantha gelida (Lindl.) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 120: 154.
2010. Crocodeilanthe gelida (Lindl.) Carnevali &
I.Ramírez, Smithsonian Contr. Bot. 100: 133. 2014.
Stelis pidax (Luer) Karremans, Phytotaxa 203(3): 293.
2015.
Bas.: Pleurothallis pidax Luer, Selbyana 5(2):
174-175. 1979. Syn.: Anathallis pidax (Luer)
Pridgeon & M.W.Chase, Lindleyana 16(4): 250.
2001. Specklinia pidax (Luer) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 95: 263. 2004. Niphantha
pidax (Luer) Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 120: 154. 2010.
Stelis subgen. Niphantha currently includes two
species of whitish, hirsute owers. Stelis gelida is a
common species with the widest distribution in the
genus, it is found from Florida and Mexico, through
Central America and the Antilles, down to Peru, Bolivia
and Brazil. Stelis pidax is only known from Ecuador.
Accessions of both Stelis gelida and S. pidax
where consistently found to form a clade sister to Stelis
s.s., Crocodeilanthe and Physothallis (Karremans
2010, Karremans et al. 2013, Pérez-Escobar et al.
2017). This is consistent with the highly unusual
morphology of these two species. Some authors
have suggested that Stelis gelida is morphologically
similar to species of Crocodeilanthe, and may be
belong there (Carnevali & Dorr 2014; Toscano de
Brito 2018a). However, the similarities are at best
supercial. The reddish, tightly clasping ramicaul
bracts, pandurate lip, elongate and curved column,
with conspicuous apical teeth, the incumbent anther,
and lack of viscidium, among many other features,
are unlike any Crocodeilanthe species.
Stelis subgen. Physosiphon (Lindl.) Karremans,
comb. nov.
Bas.: Physosiphon Lindl., Edwards’s Bot. Reg. 21:
t. 1797. 1835. Pleurothallis subgen. Physosiphon
(Lindl.) Luer, Monogr. Syst. Bot. Missouri Bot.
Gard. 20: 50. 1986. Lectotype: Stelis tubata
G.Lodd., Bot. Cab. 17: t. 1601. 1830, selected
here. (Fig. 13)
Syn. Lomax Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 105: 88. 2006. Type: Physosiphon
punctulatus Rchb.f., Botanische Zeitung (Berlin)
24(49): 385. 1866.
Stelis asperrima (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 261. 2001.
Bas.: Pleurothallis asperrima Luer, Phytologia
49(3): 201. 1981. Syn.: Physosiphon asperrimus
(Luer) Luer, Monogr. Syst. Bot. Missouri Bot. Gard.
105: 252. 2006.
Stelis emarginata (Lindl.) Soto Arenas & Solano,
Icon. Orchid. (Mexico) 5-6: t. 681. 2002 [2003].
Bas.: Pleurothallis emarginata Lindl., Gen. Sp.
Orchid. Pl. 6. 1830. Physosiphon emarginatus
(Lindl.) Lindl., Edwards’s Bot. Reg. 21: sub t. 1797.
1835.
Syn.: Stelis tubata G.Lodd., Bot. Cab. 17(161):
t. 1601. 1830. Physosiphon loddigesii Lindl.,
Edwards’s Bot. Reg. 21: sub t. 1797. 1836, nom.
inval. Physosiphon loddigesii Lindl. ex Hook. Icon.
Pl. 6: t. 508. 1843, nom. inval. Pleurothallis tubata
(G.Lodd.) Steud., Nomencl. Bot. (ed. 2) 2: 356.
1841. Physosiphon tubatus (G.Lodd.) Rchb.f., Ann.
Bot. Syst. 6(2): 188. 1861.
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Figure 13. Stelis tubata G.Lodd., selected as lectotype of genus Physosiphon Lindl., illustration of type reproduced in the
Botanical Cabinet 17: t. 1601. 1830.
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Stelis greenwoodii Soto Arenas & Solano, Icon.
Orchid. (Mexico) 5-6: , t. 682. 2002 [2003].
Syn.: Physosiphon greenwoodii (Soto Arenas &
Solano) Pfahl, Internet Orchid Sp. Photo Encycl.
Nomencl. Notes 1. 2014. 2014.
Stelis pertusa I.Jiménez, Lankesteriana 15(3): 192.
2015.
Stelis punctulata (Rchb.f.) Soto Arenas, Icon. Orchid.
(Mexico) 5-6: t. 690. 2002 [2003]. (Fig. 14)
Bas.: Physosiphon punctulatus Rchb.f., Bot.
Zeitung (Berlin) 24(49): 385. 1866. Syn.: Lomax
punctulata (Rchb.f.) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 105: 88-89. 2006.
Stelis tacanensis Solano & Soto Arenas, Icon. Orchid.
(Mexico) 5-6: t. 693. 2002 [2003].
Syn.: Physosiphon tacanensis (Solano & Soto
Arenas) Archila & Szlach., Orchid Gen. Sp.
Guatemala 643. 2018.
The six species that belong to Stelis subgen.
Physosiphon are distributed from Mexico and
Guatemala, where the highest diversity is found,
through Central America, and down to Bolivia.
DNA data is available for Stelis emarginata, S.
punctulata and S. tacanensis, they consistently group
together in a clade that is sister to a clade that includes
Stelis subgen. Niphantha, Stelis subgen. Physothallis,
Stelis subgen. Crocodeilanthe and Stelis subgen. Stelis
(Pridgeon et al. 2001, Solano-Gómez 2005, Karremans
2010, Karremans et al. 2013, Pérez-Escobar et al. 2017).
Alrich and Higgins (2011) mistakenly indicate that
Physosiphon spiralis Lindl. (= Stelis deregularis) was
selected as lectotype for genus Physosiphon by Garay
(1974). The author selects P. spiralis as lectotype for
Pseudostelis, not Physosiphon. Here Stelis tubata (=
S. emarginata), which had already been mentioned by
Pfeiffer (1873) as type, is selected as lectotype for this
species as it is clearly what Lindley based his concept
of Physosiphon on.
Stelis subgen. Dracontia (Luer) Karremans, comb.
nov.
Bas. Pleurothallis subgen. Dracontia Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 20: 38.
1986. Dracontia (Luer) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 95: 257. 2004. Type:
Pleurothallis tuerckheimii Schltr., Repert. Spec.
Nov. Regni Veg. 10(251-253): 292. 1912.
Syn. Salpistele Dressler, Orquideologia 14: 6. 1979.
Type: Salpistele brunnea Dressler, Orquideología
14(1): 6-8. 1979.
Syn. Mystacorchis Szlach. & Marg., Polish Bot. J.
46: 117. 2001. Type: Pleurothallis mystax Luer,
Selbyana 3: 146. 1976.
About three fourths of the 40 species that belong
to Stelis subgen. Dracontia are endemic to Costa Rica
and Panama. A few species extend northwards into
Mexico and Guatemala, a couple are known from the
Antilles, and three make it downwards into the Andes.
The oral morphology of this group is highly
variable, lacking apparent diagnostic features. It is made
up of a clade with the species of Luer’s Pleurothallis
subgen. Dracontia, together with Pleurothallis subgen.
Mystax, intermingled with several species placed
in Pleurothallis subgen. Effusia Lindl. and one of
Pleurothallis sect. Elongatae Lindl. It includes a clade
composed of the species of Salpistele and Pleurothallis
sect. Petiolatae. Despite the discrepancy in oral
morphology, there is no doubt that species of Dracontia,
Mystax, Petiolatae and Salpistele are closely related as
suspected from vegetative features and consistently
demonstrated by DNA studies (Pridgeon et al. 2001,
Solano-Gómez 2005, Karremans 2010, Karremans
et al. 2013, Pérez-Escobar et al. 2017, Wilson et al.
Figure 14. Stelis punctulata (Rchb.f.) Soto Arenas, type
species of genus Lomax (= Stelis subgen. Physosiphon).
Photograph by AK.
2017). Future genetic studies are necessary to conrm
the placement of some species listed here that have not
been previously analyzed, especially those placed by
Luer in Pleurothallis subgen. Effusia.
Despite its distinctive oral morphology,
the recognition of genus Salpistele as originally
circumscribed necessarily entails the recognition
of Dracontia, Mystacorchis and several other small
genera. The more distinctive groups are here given
sectional status.
Stelis subgen. Dracontia sect. Dracontia
Syn. Pleurothallis sect. Brobdingnagia Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 72: 66.
1998. Type: Pleurothallis grandis Rolfe, Bull.
Misc. Inform. Kew 1918(7): 234. 1918.
Syn. Pleurothallis sect. Cylindria Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 72: 66. 1998.
Type: Pleurothallis macrantha L.O.Williams,
Ann. Missouri Bot. Gard. 28(4): 417. 1941.
Stelis aenigma Karremans & M.Díaz, Lankesteriana
17(2): 197. 2017.
Stelis alajuelensis Pridgeon & M.W.Chase, Lindleyana
17(2): 98. 2002.
Repl. syn.: Pleurothallis ramonensis Schltr.,
Repert. Spec. Nov. Regni Veg. Beih. 19: 193-
194. 1923. Dracontia ramonensis (Schltr.) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 95: 257.
2004. Stelis ramonensis (Schltr.) Pridgeon &
M.W.Chase, Lindleyana 16: 266. 2001, nom. illeg.
Non Stelis ramonensis Schltr., Repert. Spec. Nov.
Regni Veg. Beih. 19: 176. 1923.
Stelis alta Pridgeon & M.W.Chase, Lindleyana 17(2):
98. 2002. (Fig. 15)
Repl. syn.: Pleurothallis grandis Rolfe, Bull.
Misc. Inform. Kew 1918(7): 234. 1918. Dracontia
grandis (Rolfe) Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 95: 257. 2004. Stelis grandis (Rolfe)
Pridgeon & M.W.Chase, Lindleyana 16: 263. 2001,
nom. illeg. Non Stelis grandis Rchb.f., Bonplandia
(Hannover) 3: 70. 1855.
Stelis carnosilabia (A.H.Heller & A.D.Hawkes)
Pridgeon & M.W.Chase, Lindleyana 16(4): 261. 2001.
Bas.: Pleurothallis carnosilabia A.H.Heller
& A.D.Hawkes, Phytologia 14(1): 9-10. 1966.
Syn.: Dracontia carnosilabia (A.H.Heller &
A.D.Hawkes) Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 95: 257. 2004.
Stelis cobanensis (Schltr.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 262. 2001.
Bas.: Pleurothallis cobanensis Schltr., Repert.
Spec. Nov. Regni Veg. 11(271-273): 42. 1912.
Syn.: Dracontia cobanensis (Schltr.) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 257. 2004.
Rhynchopera cobanensis (Schltr.) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 112: 121. 2007.
Stelis conochila (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 262. 2001.
Bas.: Pleurothallis conochila Luer, Lindleyana 11:
75. 1996. Syn.: Dracontia cobanensis (Schltr.) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 95: 257. 2004.
Stelis convoluta (Lindl.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 262. 2001.
Bas.: Pleurothallis convoluta Lindl., Ann. Mag.
Nat. Hist. 15: 107. 1845. Syn.: Effusiella convoluta
(Lindl.) Luer, Monogr. Syst. Bot. Missouri Bot.
Gard. 112: 106. 2007.
Stelis cylindrata Pridgeon & M.W.Chase, Lindleyana
17(2): 98. 2002. (Fig. 16)
Repl. syn.: Pleurothallis macrantha L.O.Williams,
Ann. Missouri Bot. Gard. 28(4): 417. 1941.
Dracontia macrantha (L.O.Williams) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 95: 257.
2004. Stelis macrantha (L.O.Williams) Pridgeon &
M.W.Chase, Lindleyana 16: 264. 2001, nom. illeg.
Non Stelis macrantha Rolfe, Bull. New York Bot.
Gard. 4: 450. 1907.
Stelis dies-natalis Karremans & M.Díaz, Lankesteriana
17(2): 194. 2017.
Stelis dilatata (C.Schweinf.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 262. 2001.
Bas.: Pleurothallis dilatata C.Schweinf., Bot.
Mus. Lea. 10: 177. 1942. Syn.: Effusiella dilatata
(C.Schweinf.) Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 112: 106. 2007.
Stelis dracontea (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 262. 2001. (Fig. 17)
Bas.: Pleurothallis dracontea Luer, Phytologia
49(3): 204-205. 1981. Syn.: Dracontia dracontea
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Figure 15. LCDP of Stelis alta Pridgeon & M.W.Chase, type species of Pleurothallis sect. Brobdingnagia Luer (= Stelis
sect. Dracontia). A. Habit. B. Flower. C. Dissected perianth. D. Column with lip, lateral view. E. Petal. F. Lip. G.
Column side view. H. Anther cap and pollinarium. Photographs by F. Pupulin based on Bogarín 4604 (JBL-spirit).
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(Luer) Luer, Monogr. Syst. Bot. Missouri Bot. Gard.
95: 257. 2004.
Stelis ferrelliae Pridgeon & M.W.Chase, Lindleyana
17(2): 99. 2002.
Repl. syn.: Pleurothallis ingramii Luer, Lindleyana
11(2): 81. 1996. Dracontia ingramii (Luer) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 95:
257. 2004. Stelis ingramii (Luer) Pridgeon &
M.W.Chase, Lindleyana 16: 263. 2001, nom. illeg.
Non Stelis ingramii Luer, Lindleyana 11: 100. 1996.
Stelis fortunae (Luer & Dressler) Pridgeon &
M.W.Chase, Lindleyana 16(4): 263. 2001.
Bas.: Pleurothallis fortunae Luer & Dressler,
Lindleyana 6(2): 97, 100. 1991. Syn.: Dracontia
fortunae (Luer & Dressler) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 257. 2004.
Stelis gigantea Pridgeon & M.W.Chase, Lindleyana
17(2): 99. 2002.
Repl. syn.: Pleurothallis powellii Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 17: 22. 1922. Dracontia
powelli (Schltr.) Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 95: 257. 2004. Stelis powellii (Schltr.)
Pridgeon & M.W.Chase, Lindleyana 16: 265. 2001,
nom. illeg. Non Stelis powellii Schltr., Repert. Spec.
Nov. Regni Veg. Beih. 17: 16. 1922.
Stelis hydra (Karremans & C.M.Sm.) Karremans,
Phytotaxa 203(3): 292. 2015.
Bas.: Dracontia hydra Karremans & C.M.Sm.,
Harvard Pap. Bot. 17(1): 13. 2012.
Stelis lueriana (Karremans) J.M.H.Shaw, Orchid Rev.
122(1308): 77. 2014.
Bas.: Dracontia lueriana Karremans, Ann. Natur-
hist. Mus. Wien, Ser. B, Bot. Zool. 113: 128. 2012.
Stelis megachlamys (Schltr.) Pupulin, Lankesteriana
4: 74. 2002. (Fig. 18)
Bas.: Pleurothallis megachlamys Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 19: 108. 1923.
Syn.: Pleurothallis tuerckheimii Schltr., Repert.
Spec. Nov. Regni Veg. 10(251-253): 292. 1912.
Dracontia tuerckheimii (Schltr.) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 257. 2004. Non
Stelis tuerckheimii Schltr., Repert. Spec. Nov. Regni
Veg. 8(191-195): 564. 1910.
Stelis megachlamys f. viridiavens (Roeth &
Baumbach.) Karremans, comb. nov.
Figure 16. Stelis cylindrata Pridgeon & M.W.Chase, type
species of Pleurothallis sect. Cylindria Luer (= Stelis
sect. Dracontia). Photograph by AK.
Right, Figure 17. Stelis dracontea, a typical species of Stelis
sect. Dracontia. Photograph by AK based on Bogarín
616 (JBL-spirit).
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Figure 18. LCDP of Stelis megachlamys (Schltr.) Pupulin, type species of Stelis subgen. Dracontia. A. Habit. B. Inorescence.
C. Dissected perianth. D. Column with lip, lateral view. E. Petals. F. Lip. G. Column lateral view. H. Column ventral and
side view. I. Anther cap and pollinarium. Photographs by AK based on Bogarín 2161 (JBL-spirit).
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Bas.: Dracontia tuerckheimii f. viridiavens Roeth
& Baumbach, Orchidee (Hamburg) 58: 98. 2007.
Syn.: Pleurothallis kelloggii Archila, Rev. Guatem.
15(1): 106. 2012.
Stelis montis-mortense (Karremans & Bogarín)
Bogarín & Karremans, Lankesteriana 14(3): 270.
2014.
Bas.: Dracontia montis-mortense Karremans &
Bogarín, Syst. Bot. 38(2): 307. 2013.
Stelis multirostris (Rchb.f.) Pridgeon & M.W.Chase,
Lindleyana 17(2): 100. 2002.
Bas.: Pleurothallis multirostris Rchb.f., Linnaea 41:
49. 1877.
Syn.: Epidendrum racemiorum Sw., Prodr. 125,
1788. Dendrobium racemiorum (Sw.) Sw., Nov.
Act. Upsal. 6: 83, 1799. Pleurothallis racemiora
(Sw.) Lindl. in Hook. Exot. Fl. 2: t. 123. 1825
[1824]. Anathallis racemiora (Sw.) Pridgeon &
M.W.Chase, Lindleyana 16: 250. 2001, nom. inval.
Stelis racemiora (Sw.) Pridgeon & M.W.Chase,
Lindleyana 16: 266. 2001, nom. inval. Non
Stelis racemiora (Lindl. ex Lodd.) W.H.Baxter in
J.C.Loudon, Hort. Brit., Suppl. 3: 643. 1850.
Syn.: Pleurothallis oblongifolia Lindl., Companion
Bot. Mag. 2(24): 355. 1836. Stelis oblongifolia
(Lindl.) Pridgeon & M.W.Chase, Lindleyana 16: 265.
2001, nom. illeg. Dracontia oblongifolia (Lindl.)
Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95:
257. 2004. Rhynchopera oblongifolia (Lindl.) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 112: 121.
2007. Non Stelis oblongifolia Lindl., Fol. Orchid.
8: 12. 1859.
Syn.: Pleurothallis tricostata Cogn., Symb. Antill.
7: 175, 1912.
Even though Epidendrum racemiorum Sw. has
priority over P. oblongifolia and P. multirostris, the
name is already occupied by Stelis racemiora (Lindl.
ex Lodd.) W.H.Baxter which is not based on the same
taxon. The true identity of Epidendrum racemiorum
Sw. has been confused since the nineteen hundreds,
and the taxonomy of this name continues to be highly
confused today. The available type material shows a
species belonging to Stelis sect. Dracontia, which has
generally been known as Pleurothallis oblongifolia.
However, when Lindley, in Hooker, transferred
Swartz’s name to Pleurothallis, he described and
illustrated a different species. That other species is here
treated under the name Pleurothallis quadrida (Lex.)
Lindl., and it is further discussed below.
Pridgeon and Chase (2001) proposed the names
Anathallis racemiora and Stelis racemiora using
the basionym “Pleurothallis racemiora Lindl. ex
Lodd. in Hook., Exot. Fl. 2: t. 123. 1825”. Both
names are invalid for two reasons, under article
36.3 (ICN; Turland et al. 2018) for being published
simultaneously and under article 41.5 (ICN; Turland
et al. 2018) for the erroneous citation and reference
to the publication of the basionym. Although not the
only interpretation, it is more parsimonious to assume
that the authors referred to Pleurothallis racemiora
(Sw.) Lindl. in Hook., Exot. Fl. 2: t. 123. 1825
[1824], which is based on Epidendrum racemiorum
Sw., Prodr. 125, 1788. In Pridgeon and Chase (2002),
the authors placed their S. racemiora under the
synonymy of A. racemiora, but did not validate the
name, for they failed to indicate this was intended,
and again cited the wrong basionym.
Stelis pachyglossa (Lindl.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 265. 2001.
Bas.: Pleurothallis pachyglossa Lindl., Edwards’s
Bot. Reg. 26: Misc. 68. 1840. Syn.: Dracontia
pachyglossa (Lindl.) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 257. 2004.
Stelis papillifera (Rolfe) Pridgeon & M.W.Chase,
Lindleyana 16(4): 265. 2001.
Bas.: Pleurothallis papillifera Rolfe, Bull. Misc.
Inform. Kew 1916(3): 77. 1916. Bas.: Dracontia
papillifera (Rolfe) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 257. 2004.
Stelis pileata (Karremans & Bogarín) Karremans &
Bogarín, Phytotaxa 203(3): 293. 2015.
Bas.: Dracontia pileata Karremans & Bogarín, Syst.
Bot. 38(2): 308, 310-311. 2013.
Stelis platystylis (Schltr.) Solano & Soto Arenas, Icon.
Orchid. (Mexico) 10: t. 1097. 2008.
Bas.: Pleurothallis platystylis Schltr., Repert. Spec.
Nov. Regni Veg. 10(257-259): 395. 1912. Syn.:
Effusiella platystylis (Schltr.) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 112: 107. 2007. Anathallis
platystylis (Schltr.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 250. 2001.
Stelis prolixa (Luer & Hirtz) Pridgeon & M.W.Chase,
Lindleyana 16(4): 265. 2001.
Bas.: Pleurothallis prolixa Luer & Hirtz,
Lindleyana 11(3): 179-180. 1996. Syn.: Effusiella
prolixa (Luer & Hirtz) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 112: 107. 2007.
Stelis simplex (Ames & C.Schweinf.) Pridgeon &
M.W.Chase, Lindleyana 16(4): 266. 2001.
Bas.: Pleurothallis simplex Ames & C.Schweinf.,
Sched. Orch. 10: 37-38. 1930. Syn.: Crocodeilanthe
simplex (Ames & C.Schweinf.) Toscano, Harvard
Pap. Bot. 23(1): 54. 2018.
Stelis tenebrosa (Archila, Szlach. & Chiron)
Karremans, Phytotaxa 203: 293. 2015.
Bas.: Dracontia tenebrosa Archila, Szlach. &
Chiron, Revista Guatemal. 16(1): 30. 2013.
Stelis thymochila (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 267. 2001.
Bas.: Pleurothallis thymochila Luer, Selbyana
3(3-4): 398-399, f. 299. 1977. Syn.: Dracontia
thymochila (Luer) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 257. 2004.
Stelis tintinnabula (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 267. 2001.
Bas.: Pleurothallis tintinnabula Luer, Lindleyana
11(2): 94. 1996. Syn.: Dracontia tintinnabula
(Luer) Luer, Monogr. Syst. Bot. Missouri Bot. Gard.
95: 257. 2004.
Stelis tortilis (Luer & R.Escobar) Pridgeon &
M.W.Chase, Lindleyana 16(4): 267. 2001.
Bas.: Pleurothallis tortilis Luer & R.Escobar,
Orquideología 14(2): 180. 1981. Syn.: Effusiella
tortilis (Luer & R.Escobar) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 112: 107. 2007.
Stelis viridiava (Karremans & Bogarín) Karremans,
Phytotaxa 203(3): 294. 2015.
Bas.: Dracontia viridiava Karremans & Bogarín,
Syst. Bot. 38(2): 311. 2013.
Thirty species belong to Stelis sect. Dracontia.
DNA data is available for Stelis alta, S. carnosilabia,
S. cobanensis, S. conochila, S. cylindrata, S. hydra, S.
ferrelliae, S. gigantea, S. hydra, S. lueriana, S. megachla-
mys, S. multirostris, S. pachyglossa, S. papillifera, S.
pileata, S. platystylis, S. ramonensis and S. viridiava
(Pridgeon et al. 2001, Solano-Gómez 2005, Karremans
2010, Karremans et al. 2013, Pérez-Escobar et al.
2017, Ponert et al. 2019). They have consistently been
shown to belong to Stelis sensu lato, where, despite
the oral appearances, they are most closely related to
species of Stelis sect. Mystax, Stelis sect. Petiolatae and
Stelis sect. Salpistele. Although the owers are quite
unique, the plants of species belonging to this group
are virtually indistinguishable from many typical Stelis
species (Stelis s.s.). So much so, that Stelis simplex has
not been associated with this group given the simple,
very Stelis-like, owers. The same happens with Stelis
platystylis, S. prolixa and S. tortilis which due to their
rather undifferentiated owers have not been placed
here.
Stelis subgen. Dracontia sect. Mystax (Luer)
Karremans, comb. et stat. nov.
Bas. Pleurothallis subgen. Mystax Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 20: 50. 1986. Type:
Pleurothallis mystax Luer, Selbyana 3: 146. 1976.
Syn. Mystacorchis Szlach. & Marg., Polish Bot. J.
46: 117. 2001. Type: Pleurothallis mystax Luer,
Selbyana 3: 146. 1976.
Stelis mystax (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 264. 2001. (Fig. 19)
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Figure 19. Stelis mystax, type species of Stelis sect. Mystax
(Luer) Karremans. Photograph by D. Bogarín based on
Bogarín 2988 (JBL-spirit).
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Figure 20. LCDP of Stelis guttata (Luer) Pridgeon & M.W.Chase, type species of Stelis sect. Petiolatae (Luer) Karremans.
A. Habit. B. Flower. C. Dissected perianth. D. Column with lip, lateral view. E. Column ventral and lateral views. F.
Column apex. G. Anther cap and pollinarium, placed in the stigma. Photographs by AK based on Karremans 7201
(JBL-spirit).
Bas.: Pleurothallis mystax Luer, Selbyana 3(1-2):
146-147, f. 176. 1976. Syn.: Mystacorchis mystax
(Luer) Szlach. & Marg., Polish Bot. J. 46(2): 117.
2001.
A single, aberrant species belongs to Stelis sect.
Mystax. It is endemic to Panama, and morphologically
has no close relatives. The available accessions of
this species have been consistently found to group
with other members of Stelis subgen. Dracontia,
namely the very distinct, and also unique, Central
American species, Stelis carpinterae and Stelis
convallaria (Karremans et al. 2013, Pérez-Escobar
et al. 2017). The plant and flower morphology
are somewhat reminiscent of species belonging to
Stelis sect. Dracontia, but the spathulate lip is quite
unique.
Stelis subgen. Dracontia sect. Petiolatae (Luer)
Karremans, comb. nov.
Bas. Pleurothallis sect. Petiolatae Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 52: 70. 1994.
Type: Pleurothallis guttata Luer, Selbyana 3(1-
2): 116-177. 1976.
Stelis guttata (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 263. 2001. (Fig. 20)
Bas.: Pleurothallis guttata Luer, Selbyana 3(1-2):
116-177. 1976. Syn.: Elongatia guttata (Luer) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 95: 257.
2004.
Stelis janetiae (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 264. 2001.
Bas.: Pleurothallis janetiae Luer, Selbyana 5(2):
169-170. 1979. Syn.: Elongatia janetiae (Luer)
Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95:
257. 2004.
Only two species are known to belong to Stelis sect.
Petiolatae, and DNA data is available for both. Luer
(1994) placed them in Elongatia (= Pleurothallis),
with which they indeed share a very similar oral
morphology. However, these two species endemic to
Costa Rica and Panama belong without a doubt in
Stelis sensu lato (Karremans 2010, Karremans et al.
2013, Pérez-Escobar et al. 2017). They are the closest
relatives of Stelis sect. Salpistele, and even though
their owers are very different, the plants are basically
larger versions of those.
Stelis subgen. Dracontia sect. Salpistele (Dressler)
Karremans, comb. nov.
Bas. Salpistele Dressler, Orquideologia 14: 6. 1979.
Type: Salpistele brunnea Dressler, Orquideología
14(1): 6-8. 1979.
Stelis brunnea (Dressler) Pridgeon & M.W.Chase,
Lindleyana 16(4): 261. 2001. (Fig. 21)
Bas.: Salpistele brunnea Dressler, Orquideología
14(1): 6-8. 1979.
Stelis cymbisepala Pridgeon & M.W.Chase,
Lindleyana 17(2): 98-99. 2002.
Repl. syn.: Salpistele dressleri Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 39: 128. 1991. Non Stelis
dressleri Luer, Phytologia 49(3): 227-228. 1981.
Stelis deutroadrianae J.M.H.Shaw, Orchid Rev.
122(1308): 77. 2014.
Repl. syn.: Salpistele adrianae Luer & Sijm,
Selbyana 30(1): 18. 2009. Stelis adriananijhuisae
Bogarín & Serr., Lankesteriana 14(3): 265. 2014,
nom. super. Non Stelis adrianae Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 88: 36. 2002.
Stelis gnoma Pridgeon & M.W.Chase, Lindleyana
17(2): 99. 2002.
Repl. syn.: Salpistele parvula Luer & Dressler,
Monogr. Syst. Bot. Missouri Bot. Gard. 39: 132.
1991. Non Stelis parvula Lindl., Fol. Orchid.
~Stelis~ (8): 7. 1852‒1855 [1859].
Stelis maculata Pridgeon & M.W.Chase, Lindleyana
17(2): 99. 2002.
Repl. syn.: Salpistele lutea Dressler, Orquideología
14(1): 8-10. 1979. Non Stelis lutea Lindl., Fol.
Orchid. ~Stelis~ 7. 1852‒1855 [1859].
Five species are known to belong to Stelis sect.
Salpistele, and DNA data is available for Stelis brunnea,
S. deutroadrianae and S. maculata. Despite their
Lepanthes-like owers, species of this group have been
consistently shown to belong to Stelis sensu lato based
on DNA analyses (Pridgeon et al. 2001, Solano-Gómez
2005, Karremans 2010, Karremans et al. 2013, Pérez-
Escobar et al. 2017; Wilson et al. 2017). Although not
evident from oral morphology, they are without a doubt
sister to the members of Stelis sect. Petiolatae, with
which they share the small plants with petiolate leaves
and a creeping inorescence with successive owers.
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Figure 21. LCDP of Stelis brunnea, type species of Stelis sect. Salpistele (Dressler) Karremans. A. Habit. B. Flower. C.
Dissected perianth. D. Column with lip, lateral view. E. Anther cap and pollinarium. Photographs by AK based on
Karremans 8260 (JBL-spirit).
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With them they also share the geographical area, both
groups are restricted to Costa Rica and Panama.
Stelis subgen. Dracontia sect. Carpinterae
Karremans, sect. nov.
etymology: The name honors Los Cerros de La
Carpintera, in Cartago, Costa Rica, where the
type material of its only species was collected.
Type: Pleurothallis carpinterae Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 19: 105. 1923.
This section can be easily recognized by the
long, slender ramicauls that bear a significantly
shorter, thin, ovate to sub-cordate, acute leaf. The
successive, few-flowered inflorescence reclines on
the leaf, the glabrous sepals are yellowish-cream
spotted purple. The petals have more or less the
same pattern but are darker, spathulate. The lip is as
long as the sepals, orange, pandurate, unguiculate.
The column slender, clavate, with a thick pedestal-
like base.
Stelis carpinterae (Schltr.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 261. 2001. (Fig. 22)
Bas.: Pleurothallis carpinterae Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 19: 105. 1923. Syn.:
Elongatia carpinterae (Schltr.) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 95: 257. 2004.
The only species belonging to Stelis sect.
Carpinterae is restricted to Costa Rica and western
Panama. Even though Luer (1994) placed it
among the species of Elongatia (= Pleurothallis),
morphologically it has no close relatives. The
accessions of this species were consistently found
to group with other members of Stelis subgen.
Dracontia, namely the very distinct, and also unique,
Central American species, Stelis mystax and Stelis
convallaria (Karremans et al. 2013, Pérez-Escobar et
al. 2017).
Stelis subgen. Dracontia sect. Convallaria Karremans,
sect. nov.
etymology: The name refers to the similarity of its
bell-shaped owers to those of genus Convallaria
L. (Asparagaceae), a terrestrial herb from Europe
and Asia that is known as Lilly of the valley.
Type: Pleurothallis convallaria Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 19: 185-186. 1923.
The habit is similar to other species belonging
to Stelis subgen. Dracontia, except that the multiple
inorescences are clasped basally by the leaf. The
inorescences are semi-erect and bear multiple
drooping bell-shaped owers. The dark purple sepals
are covered in a striking white a pubescens that
trembles in the wind. The dark purple petals are unusual
in that they are widest apically, truncate and bilobed.
The lip is transversally bilobed, long-unguiculate and
tricarinate. The column is elongate, bent, with a broad
clinandrium.
Stelis convallaria (Schltr.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 262. 2001. (Fig. 23)
Bas.: Pleurothallis convallaria Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 19: 185-186. 1923.
Effusiella convallaria (Schltr.) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 112: 106. 2007.
The single species belonging to Stelis sect.
Convallaria is known from Guatemala, Nicaragua,
Costa Rica and Panama. Although morphologically
somewhat aberrant, the accessions of this species are
consistently found to be related to other members of
Stelis subgen. Dracontia, especially two other unique
Central American species, Stelis mystax and Stelis
carpinterae (Karremans et al. 2013, Pérez-Escobar et
al. 2017).
Figure 22. Stelis carpinterae, type species of Stelis sect.
Carpinterae Karremans. Photograph by D. Bogarín
based on Bogarín 7159 (JBL-spirit).
Figure 23. LCDP of an autogamous specimen of Stelis convallaria, type species of Stelis sect. Convallaria. A. Habit. B.
Inorescence. C. Flower. D. Dissected perianth. E. Column with lip, lateral view. F. Column ventral and lateral views.
G. Anther cap and pollinarium, placed in the stigma. Photographs by K based on Karremans 7201 (JBL-spirit).
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Stelis subgen. Unciferia (Luer) Karremans, comb.
nov.
Bas. Pleurothallis subgen. Unciferia (Luer) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 72:
89. 1998. Pleurothallis sect. Unciferae Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 20:
94. 1986. Unciferia (Luer) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 95: 265. 2004, nom.
illeg. Type: Pleurothallis segoviensis Rchb.f.,
Bonplandia (Hannover) 3(15-16): 223-224. Non
Uncifera Lindl., J. Proc. Linn. Soc., Bot. 3: 39.
1859.
Syn. Effusiella Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 112: 106. 2007. Type: Pleurothallis
amparoana Schltr., Repert. Spec. Nov. Regni
Veg. Beih. 19: 23, 104. 1923.
Stelis amaliae (Luer & R.Escobar) Pridgeon &
M.W.Chase, Lindleyana 16(4): 261. 2001.
Bas.: Pleurothallis amaliae Luer & R.Escobar,
Orquideología 14(2): 124. 1981. Syn.: Unciferia
amaliae (Luer & R.Escobar) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 95: 265. 2004.
Stelis ancistra (Luer & Hirtz) Pridgeon & M.W.Chase,
Lindleyana 16(4): 261. 2001.
Bas.: Pleurothallis ancistra Luer & Hirtz,
Lindleyana 11(3): 144-145. 1996. Syn.: Unciferia
ancistra (Luer & Hirtz) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 265. 2004.
Stelis bifalcis (Schltr.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 261. 2001.
Bas.: Pleurothallis bifalcis Schltr., Beih. Bot.
Centralbl., Abt. 2 36(2): 395. 1918. Unciferia
bifalcis (Schltr.) Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 95: 265. 2004.
Stelis brenneri (Luer) Karremans, Phytotaxa 203(3):
292. 2015.
Bas.: Pleurothallis brenneri Luer, Selbyana 3(1-
2): 64. 1976. Syn.: Effusiella brenneri (Luer) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 112: 106.
2007.
Stelis canae (Ames) Pridgeon & M.W.Chase,
Lindleyana 16(4): 261. 2001.
Bas.: Pleurothallis canae Ames, Sched. Orch. 2:
18-19. 1923. Syn.: Unciferia canae (Ames) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 95: 265. 2004.
Stelis chlorina (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 261. 2001.
Bas.: Pleurothallis chlorina Luer, Phytologia
47(2): 75. 1980. Syn.: Effusiella chlorina (Luer)
Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 112:
106. 2007.
Stelis crenata (Lindl.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 262. 2001.
Bas.: Pleurothallis crenata Lindl., Gard. Chron.
6(13): 207. 1846. Syn.: Pabstiella crenata (Lindl.)
Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 112:
119. 2007.
Stelis cypripedoides (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 262. 2001.
Bas.: Pleurothallis cypripedioides Luer, Selbyana
1(1): 70. 1975. Syn.: Effusiella cypripedioides
(Luer) Luer, Monogr. Syst. Bot. Missouri Bot.
Gard. 112: 106. 2007.
Stelis diminuta (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 262. 2001.
Bas.: Pleurothallis diminuta Luer, Phytologia
49(3): 204. 1981. Effusiella diminuta (Luer) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 112: 106.
2007.
Stelis fornicata (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 263. 2001.
Bas.: Pleurothallis fornicata Luer, Lindleyana
11(3): 160-161. 1996. Syn.: Effusiella fornicata
(Luer) Luer, Monogr. Syst. Bot. Missouri Bot.
Gard. 112: 106. 2007.
Stelis immersa (Linden & Rchb.f.) Pridgeon &
M.W.Chase, Lindleyana 16(4): 263. 2001.
Bas.: Pleurothallis immersa Linden & Rchb.f.,
Bonplandia (Hannover) 3(15-16): 224. 1855.
Syn.: Effusiella immersa (Linden & Rchb.f.) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 112: 106.
2007.
Stelis insectifera Karremans, nom. nov.
Repl. syn.: Pleurothallis melicoides Schltr.,
Repert. Spec. Nov. Regni Veg. 19: 24. 1923. Syn.:
Stelis melicoides (Schltr.) Bogarín, Proc. 22nd
World Orchid Conf. I. 354. 2019, nom. illeg. Non
Stelis melicoides Schltr., Repert. Spec. Nov. Regni
Veg. Beih. 9: 66. 1921.
Stelis jalapensis (Kraenzl.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 263. 2001.
Bas.: Masdevallia jalapensis Kraenzl., Repert.
Spec. Nov. Regni Veg. Beih. 34: 117-118.
1925. Syn.: Pleurothallis jalapensis (Kraenzl.)
Garay, Bot. Mus. Lea. 30(3): (58)192. 1985
[1986]. Pleurothallis jalapensis (Kraenzl.)
Luer, Lindleyana 6(2): 103, f.. 1991, nom. illeg.
Specklinia jalapensis (Kraenzl.) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 261. 2004.
Effusiella jalapensis (Kraenzl.) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 112: 106. 2007,
nom. inv. Effusiella jalapensis (Kraenzl.) Archila,
Revista Guatemal. 17(2): 76. 2014.
Stelis kefersteiniana (Rchb.f.) Pridgeon & M.W.Chase,
Lindleyana 17(2): 99. 2002.
Bas.: Pleurothallis kefersteiniana Rchb.f., Bot.
Zeitung (Berlin) 10: 673. 1852.
Syn.: Specklinia exuosa Poepp. & Endl., Nov. Gen.
Sp. Pl. 1: 52, t. 90. 1835. Pleurothallis exuosa
(Poepp. & Endl.) Lindl., Edwards’s Bot. Reg. 28:
Misc. 69, no. 7. 1842. Syn.: Effusiella exuosa
(Poepp. & Endl.) Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 112: 106. 2007, nom. inval. Stelis
exuosa (Poepp. & Endl.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 263. 2001, nom. illeg. Non Stelis
exuosa Lindl., Ann. Mag. Nat. Hist. 12(79): 397.
1843.
Stelis lehmanneptis (Luer & R.Escobar) Pridgeon &
M.W.Chase, Lindleyana 16(4): 264. 2001.
Bas.: Pleurothallis lehmanneptis Luer &
R.Escobar, Orquideología 21: 100. 1998.
Effusiella lehmanneptis (Luer & R.Escobar) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 112: 106.
2007.
Stelis lehmannii Pridgeon & M.W.Chase, Lindleyana
17(2): 99. 2002.
Repl. syn.: Pleurothallis petiolaris Luer,
Orquideología 20: 220. 1996. Effusiella petiolaris
(Luer) Luer, Monogr. Syst. Bot. Missouri Bot.
Gard. 112: 107. 2007. Stelis petiolaris (Luer)
Pridgeon & M.W.Chase, Lindleyana 16(4): 265.
2001, nom. illeg. Non Stelis petiolaris Schltr.,
Repert. Spec. Nov. Regni Veg. Beih. 27: 36.
1924.
Stelis listerophora (Schltr.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 264. 2001.
Bas.: Pleurothallis listerophora Schltr., Repert.
Spec. Nov. Regni Veg. 3(33-34): 107. 1906. Syn.:
Effusiella listerophora (Schltr.) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 112: 106. 2007.
Stelis niesseniae (Luer) Karremans, Phytotaxa 406(5):
265. 2019.
Bas.: Pleurothallis niesseniae Luer, Orquideología
22(1): 59-61. 2001. Syn.: Effusiella niesseniae
(Luer) Luer, Monogr. Syst. Bot. Missouri Bot. Gard.
112: 107. 2007.
Stelis nigriora (L.O.Williams) Pridgeon &
M.W.Chase, Lindleyana 16(4): 264. 2001.
Bas.: Pleurothallis nigriora L.O.Williams, Amer.
Orchid Soc. Bull. 11(5): 168. 1942. Effusiella
nigriora (L.O.Williams) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 112: 107. 2007.
Stelis nonresupinata Solano & Soto Arenas, Icon.
Orchid. (Mexico) 5-6: t. 688. 2002 [2003].
Stelis oestlundiana (L.O.Williams) Pridgeon &
M.W.Chase, Lindleyana 16(4): 265. 2001.
Bas.: Pleurothallis oestlundiana L.O.Williams,
Bot. Mus. Lea. 12(7): 243. 1946. Syn.: Effusiella
oestlundiana (L.O.Williams) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 112: 107. 2007.
Stelis ornata (Rchb.f.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 265. 2001. (Fig. 24)
Bas.: Pleurothallis ornata Rchb.f., Garten Zeitung
1: 106. 1882. Effusiella ornata (Rchb.f.) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 112: 107.
2007.
Stelis oscargrouchii Karremans, Phytotaxa 203(3):
293. 2015.
Repl. syn.: Specklinia ximenae Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 103: 311. 2005. Syn.:
Pleurothallis ximenae Luer & Hirtz, Monogr. Syst.
Bot. Missouri Bot. Gard. 95: 238, 242. 2004, nom.
inval. Specklinia ximenae (Luer & Hirtz) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 95: 265. 2004,
nom. inval. Effusiella ximenae (Luer & Hirtz) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 112: 107.
2007, nom. inval. Non Stelis ximenae Luer & Hirtz,
Monogr. Syst. Bot. Missouri Bot. Gard. 95: 175. 2004.
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Stelis pilosa Pridgeon & M.W.Chase, Lindleyana
17(2): 100. 2002. (Fig. 25)
Repl. syn.: Pleurothallis amparoana Schltr.,
Repert. Spec. Nov. Regni Veg. Beih. 19: 23, 104.
1923. Effusiella amparoana (Schltr.) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 112: 106. 2007.
Stelis amparoana (Schltr.) Pridgeon & M.W.Chase,
Lindleyana 16: 261. 2001, nom. illeg. Non Stelis
amparoana Schltr., Repert. Spec. Nov. Regni Veg.
Beih. 19: 16. 1923.
Stelis pilostoma (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 265. 2001.
Bas.: Pleurothallis pilostoma Luer, Lindleyana
11(2): 89. 1996. Unciferia pilostoma (Luer) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 95: 265.
2004.
Stelis pompalis (Ames) Pridgeon & M.W.Chase,
Lindleyana 16(4): 265. 2001.
Bas.: Pleurothallis pompalis Ames, Sched. Orch. 7:
23-25. 1924. Syn.: Unciferia pompalis (Ames) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 95: 265. 2004.
Stelis pseudocheila (Luer & R.Escobar) Pridgeon &
M.W.Chase, Lindleyana 16(4): 266. 2001.
Bas.: Pleurothallis pseudocheila Luer & R.Escobar,
Orquideología 16(2): 173. 1984. Syn.: Effusiella
pseudocheila (Luer & R.Escobar) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 112: 107. 2007.
Stelis psilantha (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 266. 2001.
Bas.: Pleurothallis psilantha Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 72: 95. 1998. Syn.:
Unciferia psilantha (Luer) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 265. 2004.
Stelis resupinata (Ames) Pridgeon & M.W.Chase,
Lindleyana 16(4): 266. 2001.
Bas.: Pleurothallis resupinata Ames, Orchidaceae 2:
272. 1908. Syn.: Effusiella resupinata (Ames) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 112: 107.
2007.
Figure 24. Stelis ornata (Rchb.f.) Pridgeon & M.W.Chase,
a species with striking appendages on the sepals, but
otherwise a typical member of Stelis subgen. Unciferia
(Luer) Karremans. Photograph by H. Oakeley.
Figure 25. Stelis pilosa Pridgeon & M.W.Chase, type
species of genus Effusiella (Luer) Luer (= Stelis subgen.
Unciferia). Photograph by AK.
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Figure 26. LCDP of Stelis segoviensis, type species of Stelis subgen. Unciferia. A. Habit. B. Inorescence. C. Flower. D.
Dissected perianth. E. Column with lip, lateral view. F. Column ventral and lateral view. G. Anther cap and pollinarium.
Photographs by AK based on Rojas-Alvarado 311 (JBL-spirit).
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Stelis retusa (Lex.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 266. 2001.
Bas.: Dendrobium retusum Lex., Nov. Veg. Descr.
2(Orchid. Opusc.): 40. 1825. Specklinia retusa
(Lex.) Lindl., Edwards’s Bot. Reg. 21: sub t. 1797.
1835. Pleurothallis retusa (Lex.) Lindl., Edwards’s
Bot. Reg. 28: Misc. 81-82. 1842. Syn.: Effusiella
retusa (Lex.) Luer, Monogr. Syst. Bot. Missouri Bot.
Gard. 112: 107. 2007, nom. inval.
Stelis rostratissima (Luer & J. Portilla) Karremans,
Phytotaxa 203(3): 293. 2015.
Bas.: Pleurothallis rostratissima Luer & J. Portilla,
Monogr. Syst. Bot. Missouri Bot. Gard. 88: 108.
2002. Syn.: Effusiella rostratissima (Luer & J.
Portilla) Luer, Monogr. Syst. Bot. Missouri Bot.
Gard. 112: 107. 2007.
Stelis segoviensis (Rchb.f.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 266. 2001. (Fig. 26)
Bas.: Pleurothallis segoviensis Rchb.f., Bonplandia
(Hannover) 3(15-16): 223-224. 1855. Syn.:
Unciferia segoviensis (Rchb.f.) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 95: 265. 2004.
Stelis thomasii (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 267. 2001.
Bas.: Pleurothallis thomasii Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 79: 84, 130. 2000.
Effusiella thomasii (Luer) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 112: 107. 2007.
Stelis trichostoma (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 267. 2001.
Bas.: Pleurothallis trichostoma Luer, Selbyana
5(2): 185. 1979. Effusiella trichostoma (Luer) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 112: 107.
2007.
Stelis trulla (Rchb.f. & Warsz.) Pridgeon &
M.W.Chase, Lindleyana 16(4): 267. 2001.
Bas.: Pleurothallis trulla Rchb.f. & Warsz.,
Bonplandia (Hannover) 2: 114. 1854. Effusiella
trulla (Rchb.f. & Warsz.) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 112: 107. 2007.
Stelis uncinata Pridgeon & M.W.Chase, Lindleyana
17(2): 100. 2002.
Repl. syn.: Pleurothallis kareniae Luer, Lindleyana
11(2): 83, f. 19. 1996. Unciferia kareniae (Luer)
Luer, Monogr. Syst. Bot. Missouri Bot. Gard.
95: 265. 2004. Stelis kareniae (Luer) Pridgeon &
M.W.Chase, Lindleyana 16(4): 264. 2001, nom.
illeg. Non Stelis kareniae Luer, Lindleyana 11(2):
100, f. 31. 1996.
Stelis villosa (Knowles & Westc.) Pridgeon &
M.W.Chase, Lindleyana 16(4): 267. 2001.
Bas.: Pleurothallis villosa Knowles & Westc., Fl.
Cab. 2: 78. 1838. Syn.: Effusiella villosa (Knowles
& Westc.) Luer, Monogr. Syst. Bot. Missouri Bot.
Gard. 112: 107. 2007.
Stelis vinacea (Ames) Bogarín, Proc. 22nd World
Orchid Conf. I. 358. 2019.
Bas.: Pleurothallis vinacea Ames, Schedul. Orchid.
6: 69. 1923.
Stelis wagneri (Schltr.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 267. 2001.
Bas.: Pleurothallis wagneri Schltr., Repert. Spec.
Nov. Regni Veg. 17(8-12): 141. 1921. Unciferia
wagneri (Schltr.) Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 95: 265. 2004.
Stelis werckleana Bogarín & Pupulin, Proc. 22nd
World Orchid Conf. I. 358. 2019.
Repl. syn.: Pleurothallis wercklei Schltr., Repert.
Spec. Nov. Regni Veg. 17: 141. 1921. Non
Stelis wercklei Schltr., Repert. Spec. Nov. Regni
Veg. Beih. 19: 20. 1923.
Stelis xerophila (Schltr.) Soto Arenas, Icon. Orchid.
(Mexico) 5-6: t. 695. 2002 [2003].
Bas.: Pleurothallis xerophila Schltr., Beih. Bot.
Centralbl., Abt. 2 36(2): 398. 1918. Syn.: Specklinia
xerophila (Schltr.) Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 95: 265. 2004.
Stelis zootrophionoides Castañeda-Zárate & Ramos-
Castro, PLoS ONE 7(11): 5. 2012.
Syn.: Effusiella zootrophionoides (Castañeda-
Zárate & Ramos-Castro) Archila, Revista Guatemal.
17(2): 76. 2014.
The 42 species that belong to Stelis subgen.
Unciferia are found only from Mexico to Bolivia and
Peru, they are especially diverse in Middle America
and no records exist for the Antilles or Brazil.
DNA data is available for Stelis canae, S. immersa,
S. jalapensis, S. kefersteiniana, S. listerophora,
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Figure 27. LCDP of Stelis furculifera, one of the two members of Stelis subgen. Condylago (Luer) Karremans. A. Habit.
B. Flower. C. Dissected perianth. D. Flower, oral bract, lateral view. E. Column semi-ventral, side view, with and
without the lip attached. F. Pollinarium. Photographs by AK based on Bogarín 5901 (JBL-spirit)..
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S. nigriora, S. ornata, S. pilosa, S. pompalis, S.
resupinata, S. retusa, S. segoviensis, S. trichostoma,
S. zootrophionoides (Pridgeon et al. 2001, Solano-
Gómez 2005, Karremans 2010, Ramos-Castro et al.
2012; Karremans et al. 2013, Pérez-Escobar et al.
2017, Ponert et al. 2019). They all consistently appear
within Stelis in the broad sense, and mostly as sisters
to the members of Stelis subgen. Dracontia. However,
diverse analyses (using different genes, methods or
taxa) provide contradicting relationships among them.
Stelis pilosa, type species of genus Effusiella, appears
to be a close relative of Stelis segoviensis, type species
of genus Unciferia, but there is also support for a
relationship between some of the members of Stelis
subgen. Unciferia and species of either Stelis subgen.
Condylago and Stelis subgen. Dracontia. It may
therefore not be a monophyletic group and requires
further analysis.
Stelis subgen. Condylago (Luer) Karremans, comb.
nov.
Bas. Condylago Luer, Orquideologia 15: 118. 1982.
Type: Condylago rodrigoi Luer, Orquideología
15(2-3): 118-122. 1982.
Stelis furculifera (Dressler & Bogarín) Bogarín,
Lankesteriana 14(3): 267. 2014. (Fig. 27)
Bas.: Condylago furculifera Dressler & Bogarín,
Harvard Pap. Bot. 12(1): 2-5. 2007.
Stelis rodrigoi (Luer) Pridgeon & M.W.Chase,
Lindleyana 16(4): 266. 2001.
Bas.: Condylago rodrigoi Luer, Orquideología 15(2-
3): 118-122. 1982.
The two species that belong to Stelis subgen.
Condylago are restricted to Panama and Colombia
respectively. DNA data is only available for Stelis
rodrigoi and most analyses nd it as a sister to the rest
of Stelis sensu lato (Karremans 2010, Ramos-Castro et
al. 2012; Karremans et al. 2013, Pérez-Escobar et al.
2017). However, a relationship with members of Stelis
subgen. Unciferia was also found by authors (Pridgeon
et al. 2001, Ponert et al. 2019).
Stelis subgen. Umbralia Karremans, subgen. nov.
etymology: From the Latin umbra, shade or shadow,
in reference to the short twisted inorescence
hidden under the shade of the convex leaf.
Type: Pleurothallis imraei Lindl., Fol. Orchid.
~Pleurothallis~ 9. 1859.
Distinguished from all other subgenera by the large
caespitose plants bearing ovate to suborbicular leaves that
are typically convex. The inorescence is signicantly
shorter than the leaf and twists in such a way that the
owers are frequently hidden behind the leaf blade. The
ovary is strongly bent, causing the owers to be oriented
upwards. The sepals are internally pubescent, the lateral
sepals forming a synsepal with a mentum at the base.
The petals are conspicuously spathulate, obtuse. The
lip is convex in natural position, unguiculate, delicately
hinged to the column foot, lanceolate when extended,
obtuse. The column is cylindrical, incurved, with a
pair of small wings, apically denticulate. Pollinia two,
forming a whale-tail type pollinarium with a pair of
attish caudicles.
Stelis cocornaënsis (Luer & R.Escobar) Pridgeon &
M.W.Chase, Lindleyana 16(4): 262. 2001.
Bas.: Pleurothallis cocornaënsis Luer & R.Escobar,
Orquideología 20: 45. 1996. Syn.: Specklinia
cocornaënsis (Luer & R.Escobar) Luer, Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 259. 2004.
Effusiella cocornaënsis (Luer & R.Escobar) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 112: 106.
2007.
Stelis erucosa (Luer & R.Escobar) Pridgeon &
M.W.Chase, Lindleyana 16(4): 262. 2001.
Bas.: Pleurothallis erucosa Luer & R.Escobar,
Orquideología 21(1): 88. 1998. Syn.: Specklinia
erucosa (Luer & R.Escobar) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 95: 260. 2004. Effusiella
erucosa (Luer & R.Escobar) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 112: 106. 2007.
Stelis imraei (Lindl.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 263. 2001. (Fig. 28)
Bas.: Pleurothallis imraei Lindl., Fol. Orchid.
~Pleurothallis~ 9. 1859. Syn.: Humboldtia imraei
(Lindl.) Kuntze, Revis. Gen. Pl. 2: 667. 1891.
Specklinia imraei (Lindl.) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 261. 2004. Effusiella imraei
(Lindl.) Luer, Monogr. Syst. Bot. Missouri Bot.
Gard. 112: 106. 2007.
Syn.: Pleurothallis umbraticola Schltr., Repert.
Spec. Nov. Regni Veg. 27(1-8): 56-57. 1929.
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Figure 28. LCDP of Stelis imraei, type species of Stelis subgen. Umbralia Karremans. A. Habit. B. Flower. C. Dissected
perianth. D. Column with lip, lateral view. D. Column lateral view. E. Anther cap and pollinarium. Photographs by I.
Chinchilla based on Bogarín 752 (JBL-spirit).
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Stelis tarantula (Luer & Hirtz) Pridgeon & M.W.Chase,
Lindleyana 16(4): 267. 2001.
Bas.: Pleurothallis tarantula Luer & Hirtz,
Lindleyana 11(3): 186-187. 1996. Syn.: Specklinia
tarantula (Luer & Hirtz) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 95: 264. 2004. Effusiella
tarantula (Luer & Hirtz) Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 112: 107. 2007.
Stelis vaginata (Schltr.) Pridgeon & M.W.Chase,
Lindleyana 16(4): 267. 2001.
Bas.: Pleurothallis vaginata Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 19: 197. 1923. Syn.:
Specklinia vaginata (Schltr.) Luer, Monogr. Syst.
Bot. Missouri Bot. Gard. 95: 264. 2004. Effusiella
vaginata (Schltr.) Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 112: 107. 2007. Pleurothallis imraei var.
vaginata (Schltr.) O. Gruss & M. Wolff, Orchid
Atlas 359. 2007.
Five species are currently recognized as belonging
to Stelis subgen. Umbralia. They are mostly found
from Costa Rica to Ecuador, with the highest diversity
in Colombia. This subgenus includes the very variable
Stelis imraei, which is most like a species complex
distributed from Costa Rica to Peru and Bolivia, the
Guyanas, and the Lesser Antilles.
Although Luer (2000) placed these species among
the Effusiella, and the owers are indeed similar,
vegetative morphology is quite unlike any other Stelis.
Not surprisingly, the DNA data available for Stels
imraei places it as sister to all other members of Stelis
in the broad sense (Karremans 2010, Karremans et al.
2013). A multigene analysis of the Pleurothallidinae
including an accession of Stelis cocornaënsis places
it as sister to Stelis subgen. Dracontia (Ponert et al.
2019). It is certainly possible to segregate this group
into a genus of its own, however, such a proposal
would be inconsistent with the current interpretation
that this group is best treated a single genus at this
time.
excluded tAxA
The following groups are still associated with
Stelis in the broad sense, or with certain species groups
therein, in literature. They are here explicitly stated
with hopes that they can be denitively be excluded
from Stelis and no longer be associated with any taxa
belonging to it.
Pleurothallis sect. Alatae Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 76: 99. 1999. Type: Specklinia
obovata Lindl. Edwards’s Bot. Reg. 25: Misc. 75.
1842. (Fig. 29) = Anathallis Barb.Rodr.
DNA data has consistently shown that members
of Pleurothallis subgen. Acuminatia sect. Acuminatae
belong in Stelis, whilst those placed in Acuminatia
sect. Alatae belong to Anathallis (Karremans et al.
2013; Karremans 2014, Pérez-Escobar et al. 2017).
Morphologically this is easily diagnosable as species of
sect. Alatae, like other species of Anathallis, have star-
shaped owers, with acuminate petals that are as long
as the sepals, a attened lip and the sharply winged,
apically fringed column. Whereas species of sect.
Acuminatae, like other species of Stelis, bear obtuse
petals, that are signicantly shorter than the sepals, have
a non-attened lip, and the column is not prominently
winged or fringed (Karremans 2014; 2016).
Pleurothallis subgen. Effusia Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 79: 54. 2000. Syn.: Pleurothallis
sect. Effusae Lindl. Edwards’s Bot. Reg. 28: Misc.
74. 1842. Type: Pleurothallis hypnicola Lindl.
Edwards’s Bot. Reg. 28: Misc. 75. 1842. (Fig. 30) =
Pabstiella Brieger & Senghas
DNA data has consistently shown that P. hypnicola
and its relatives belong in Pabstiella rather than Stelis
(Karremans et al. 2013, Pérez-Escobar et al. 2017).
Even though not closely related, there is a striking
similarity between species of Pabstiella and some
members of Stelis subgen. Unciferia, a convergence
that most likely responds to a similar pollination
syndrome. With few exceptions, species of Pabstiella
are found in Brazil, where the members of Stelis
subgen. Unciferia are absent. The latter instead are
most diverse in Middle America, where very few
Pabstiella species have been recorded.
Elongatia (Luer) Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 95: 257 (2004). Bas.: Pleurothallis
subgen. Elongatia Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 20: 41. 1986. Syn.: Pleurothallis sect.
Elongatae Lindl. Edwards’s Bot. Reg. 28: Misc. 68.
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Figure 29. LCDP of Anathallis obovata, type species of Anathallis. A. Habit. B. Inorescence. C. Dissected perianth. D.
Column with lip, lateral view. D. Column ventral and lateral view. E. Pollinarium. Photographs by AK based on JBL-
28233 (JBL-spirit).
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Figure 30. LCDP of Pabstiella hypnicola, type species of Pleurothallis subgen. Effusia (= Pabstiella). A. Habit. B. Flower.
C. Dissected perianth. D. Column with lip, lateral view. E. Column ventral and lateral view. F. Lip. G. Anther cap and
pollinarium. Photographs by G. Rojas-Alvarado based on HBL960631 (JBL-spirit).
1842. Lectotype: Pleurothallis restrepioides Lindl.
Companion Bot. Mag. 2: 356. 1836. (Fig. 31) =
Pleurothallis R.Br.
DNA data has consistently shown that P.
restrepioides, type species of Elongatia, and its closest
relatives belong in Pleurothallis rather than Stelis
(Karremans et al. 2013; Wilson et al. 2013, Pérez-
Escobar et al. 2017). The owers of Elongatia are
supercially similar to the members of Stelis sect.
Carpinterae and Stelis sect. Petiolatae. From the rst
they are distinguished by very large plants, with thick
coriaceous leaves, from the second by the large plants
with sessile leaves. From both, Elongatia species are
distinguished by the erect, elongate inorescence with
multiple simultaneous owers.
Pleurothallis subgen. Lalexia (Luer) Karremans,
comb. et stat. nov.
Bas.: Lalexia Luer, Harvard Pap. Bot. 16: 358.
2011. Syn. Loddigesia Luer, Monogr. Syst. Bot.
Missouri Bot. Gard. 105: 251. 2006, nom. illeg.
Non Loddigesia Sims Bot. Mag. 24: pl. 965.
1806. Type: Dendrobium quadridum Nov. Veg.
Descr, 2(Orch. Opusc.): 40-41. 1825. (Fig. 32)
Morphologically this taxon could be confused with
a member of Stelis in the broad sense, and in fact many
authors still place it in Stelis rather than Pleurothallis.
Stenzel (2004) doubted the results of his own
phylogenetic reconstruction in which two accessions
of Pleurothallis ghiesbreghtiana A.Rich. & Galeotti (=
P. quadrida) were found sister to Pleurothallis rather
than Stelis. However, except for the phylogenetic
inference presented by Solano-Gómez (2005), all
other DNA based studies consistently show that P.
quadrida, type species of Lalexia, is sister to the
remaining species of Pleurothallis rather than Stelis
(Stenzel 2004, Karremans et al. 2013, Wilson et al.
2013, 2017, Pérez-Escobar et al. 2017). The exclusion
from Stelis is supported by multi-gene genomic studies
(Ponert et al. 2019).
Its only member, Pleurothallis quadrifida, is a
widely distributed and common species without any
close relatives. It is easily recognized by the thick
coriaceous leaves, erect, elongate, simultaneous
inflorescences bearing large bright yellow flowers.
It is unique in the glabrous flowers, with petals
similar in size the sepals, the pandurate lip and the
simple column with sub-apical anther. Unlike the
majority of the members of the subtribe, the sweetly
fragrant bright yellow flowers of this species may
be adapted to pollination by Hymenoptera rather
than Diptera, as a parasitoid wasp was documented
removing pollinaria (Karremans & Díaz-Morales
2019).
Pleurothallis quadrida (Lex.) Lindl., Edwards’s Bot.
Reg. 28(Misc.): 70. 1842.
Bas.: Dendrobium quadridum Lex. in P.de
La Llave & J.M.de Lexarza, Nov. Veg. Descr.
2(Orchid. Opusc.): 40. 1825. Humboltia quadrida
(Lex.) Kuntze, Revis. Gen. Pl. 2: 668. 1891. Stelis
quadrida (Lex.) Solano & Soto Arenas, Icon.
Orchid. 5-6: xi. 2002 [2003]. Specklinia quadrida
(Lex.) Luer, Monogr. Syst. Bot. Missouri Bot. Gard.
95: 263. 2004. Loddigesia quadrida (Lex.) Luer,
Monogr. Syst. Bot. Missouri Bot. Gard. 105: 251.
2006. Lalexia quadrida (Lex.) Luer, Harvard Pap.
Bot. 16: 358. 2011.
Syn.: Pleurothallis racemiora Lindl. ex Lodd.
Bot. Cab. 10: t. 949. 1824 [1825], nom. illeg. Non
Pleurothallis racemiora (Sw.) Lindl. in Hook. Exot.
Fl. 2: t. 123. 1825 [1824]. Stelis racemiora (Lindl.
ex Lodd.) W.H.Baxter in J.C.Loudon, Hort. Brit.,
Suppl. 3: 643. 1850, nom. illeg. Pleurothallis
longissima Lindl., Fol. Orchid. ~Pleurothallis~ 31.
1859.
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Figure 31. Pleurothallis restrepioides Lindl., type species of
Elongatia (= Pleurothallis). Photograph by J. Varigos.
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Figure 32. LCDP of Pleurothallis quadrida, type species of Lalexia (= Pleurothallis subgen. Lalexia). A. Habit. B.
Inorescence. C. Flower. D. Dissected perianth. E. Column with lip, lateral view. F. Lip. G. Column ventral and lateral
view. H. Anther cap and pollinarium. Photographs by AK based on Karremans 6436 (JBL-spirit).
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Great taxonomical confusion surrounds the
name Pleurothallis racemiora in commonly used
databases and literature, warranting the current note.
When Lindley, in Hooker (1824), transferred Swartz’s
Epidendrum racemiorum to Pleurothallis, rather than
describing and illustrating Swartz’s species (treated
above as Stelis multirostris) he presented another
species, treated here as Pleurothallis quadrida.
Lindley later realized his initial mistake and rectied
“P. longissima (P. racemiora, Lindl. in Hook. Exot.
Fl. t 123, nec Swartzii)... It is not the Dendrobium
racemiorum Sw. as I formerly supposed” and that
“P. racemiora (Dendrobium racemiorum Swartz,
Fl. Ind. Occ. 1543. P. oblongifolia Lindl. in Comp.
Bot. Mag. 2. 355)... Original specimens from Swartz
show that this is the plant meant by Swartz”.
However, the damage was already done. In the
Botanical Cabinet, Loddiges (1825) features the
same Pleurothallis racemiora that Lindley had
misinterpreted (thus Pleurothallis quadrida), and
not that of Swartz (= Stelis multirostris). Loddiges’
name has been regarded by various authors as
a combination for the basionym Epidendrum
racemiorum Sw. or simply as a citation of P.
racemiora (Sw.) Lindl. Nevertheless, in the original
publication there is no reference to either, and as
both text and illustration are based on Loddiges’ own
material that actually represents a different species as
that of Swartz, it must be interpreted that the author is
publishing a new taxon. The name is therefore to be
cited correctly as P. racemiora Lindl. ex Lodd. and
it is an heterotypic homonym of P. racemiora (Sw.)
Lindl.
As P. racemiora (Sw.) Lindl. latter was
published a few months prior to P. racemiora
Lindl. ex Lodd., it has priority. Therefore, P.
racemiora Lindl. ex Lodd. is valid, yet illegitimate
under article 53.1 (Turland et al. 2018). The name
Stelis racemiora published by Baxter in the
“Supplement to J.C. Loudon’s Hortus Britannicus”
clearly cites Loddiges as author, and thus should
be correctly cited as Stelis racemiora (Lindl. ex
Lodd.) W.H.Baxter. Although most names from the
Hortus Britannicus are considered invalid, this new
combination is in accordance with articles 35.2, 38.1,
38.2 and 41.4 as it associates the genus and nal
epithet and associating the new combination with
a basionym and earlier description (Turland et al.
2018). However, it is based on an illegitimate name,
and thus illegitimate too. Both are here regarded as
heterotypic synonyms of P. quadrida as they are
based on Loddiges’ material rather than Swartz’s.
Pleurothallis longissima Lindl. is based on the same
type as P. racemiora Lindl. ex Lodd., and thus can
be considered a replacement name.
Conclusions. To be, or not to be a Stelis, that is
the question. For that we hope to have an answer.
Recognizing a member of Stelis in the classic strict
sense is certainly straightforward. Most species
(not all) have a standard and distinctive oral
morphology. If the group was an isolated lineage
within the Pleurothallidinae there would be no need
for the current discussion. However, that is not the
case. We now know for a fact that many groups of
species that lack the typical Stelis-ower are actually
close relatives. An alternative would be not to add
these groups to a broader Stelis but to recognize each
of them as genera as well. But is that alternative more
intuitive or informative? It doesn’t seem to be that
way at all.
Sadly, none of the possible ways in which we
can translate the evolutionary history of this group of
species into a stable classication system appears to be
very appealing. Stelis in the broad sense dened here
is made up of a series of strikingly different species
groups that indisputably share a common ancestor and
a common evolutionary history. In the past, ower
morphology has been the main source for information
regarding evolutionary history between taxa, however
today we know that ower morphology in distant taxa
may appear very similar due to convergence evolution.
Why, despite their indistinguishable owers, are
we happy to accept that Bulbophyllum careyanum
Spreng., B. striatellum Ridl., B. laxiorum Lindl.,
B. maxillare Rchb.f. and B. tremulum Wight, are not
actually species of Pleurothallidinae belonging to
the genera Acianthera Scheidw., Muscarella Luer,
Myoxanthus Poepp. & Endl., Masdevallia Ruiz & Pav.
and Trichosalpinx Luer, respectively? Because it has
been established beyond a doubt that these groups
are unrelated and their oral similarity is merely a
consequence of convergent evolution due to similar
pollination syndromes.
Similarly, we should accept that convergent
morphologies occur within the Pleurothallidinae, the
group with highest diversication rates and species
number in Orchidaceae. It is a fact that species of
Andinia (Luer) Luer are not closely related to species
of Lepanthes Sw., and that neither of them is a close
relative of species of Salpistele Dressler (= Stelis),
despite having almost identical owers. In the same
way, it has been proven that species of Anathallis
Barb.Rodr. are not closely related to the orally
similar of Lankesteriana Karremans. We know that
those oral convergences result from adaptation to
the same pollinators or pollination strategies (Wilson
et al. 2017; Bogarín et al. 2018; Karremans & Díaz-
Morales 2019). Specically, in the case of Stelis s.l.,
Karremans & Díaz-Morales (2019) stress the point
that species of Stelis subgen. Unciferia have been
reported to be pollinated by ies of the families
Phoridae and Chloropidae which are exactly the same
families of ies that pollinate species of Acianthera,
an unrelated genus with owers that are indeed much
more similar than those of Stelis s.s. The authors also
show that the transitional morphology of species
belonging to Stelis subgen. Crocodeilanthe results
in the placement of pollinaria on the top of the head
of their pollinators, which is midway between the
scutellum placement of members of Stelis subgen.
Unciferia and the placement close to the mouthparts
observed in Stelis s.s. (Karremans & Díaz-Morales
2019).
As circumscribed here, Stelis includes 1243
species, making it the most species rich genus in
the Pleurothallidinae, and one of the largest in
Orchidaceae. The most specious group in the genus is
Stelis subgen. Stelis, which harbors some 1030 species
with the more classical Stelis ower morphology. The
other 213 species are divided into eight subgenera
that although orally different are closely related and
share the same common ancestor of Stelis s.s.
AcKnowledgements. Franco Pupulin, Gustavo Rojas-
Alvarado, Juan Sebastián Moreno and Isler Chinchilla were
kind enough to authorize the reproduction of their LCDPs.
Diego Bogarín, Benjamín Collantes, Jan Meijvogel, Henry
Oakeley and Grettel Salguero are thanked for providing
photographs used in this manuscript. Two anonymous
reviewers are thanked for their kind suggestions improving
the manuscript. The Hortus Botanicus Leiden, and staff
associated, is thanked for some of the plant material
reproduced here provided through an exchange with
Lankester Botanical Garden. The staff of the horticulture
department at Lankester Botanical Garden is thanked for
their invaluable support to this study. Kanchi Gandhi is
thanked for his comments regarding nomenclatural issues
discussed herein.
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LANKESTERIANA 19(3). 2019. © Universidad de Costa Rica, 2019.
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