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THE BULLETIN OF THE AMERICAN ORCHID SOCIETY
VOL. 89 NO. 8 AUGUST 2020
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CONTENTS
August 2020
Volume 89
Number 8
The Bulletin of the American Orchid Society
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602
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FEATURES
618 Rhyncholaelia digbyana
The Frilled Wonder
Stephen van Kampen-Lewis
626 orchids in watercolor
Isotria verticillata
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Fieldwork in the Most Mysterious and Dangerous Mountains of Ecuador
Kelsey L. Huisman
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Epiphytic Orchids North of the 49th Parallel
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Lindleyana 656
Studies on Oberonia 7
Ten New Synonyms of Oberonia equitans (G. Forst.)
Mutel Indicated by Morphology and Molecular
Phylogeny
Daniel L. Geiger, Benjamin J. Crain, Melissa K.
McCormick and Dennis F. Whigham
FRONT COVER
Those massive, frilly lips that we have come to love in complex Rhyncholaeliocattleya hybrids (often socalled “brasso-lips”) owe their origin to Rhyncholaelia digbyana. This incredible species has proven to
be a goldmine for cattleya breeders over the years. Pictured here is Rl. digbyana ‘Springwater’ AM/AOS,
photographed by Ernest Walters, was grown by Thanh Nguyen, Springwater Orchids.
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Lindleyana
Studies on Oberonia 7
Ten New Synonyms of Oberonia equitans (G. Forst.) Mutel Indicated
by Morphology and Molecular Phylogeny
By Daniel L. Geiger, Benjamin J. Crain, Melissa K. McCormick and Dennis F. Whigham
ABSTRACT A combined morphological-molecular investigation identifies ten new synonyms of Oberonia equitans (G.Fost.)Mutel:
Oberonia affinis Ames & C.Schweinf. in O.Ames, Oberonia aurea Schltr., Oberonia ciliolata Hook.f., Oberonia enoensis J.J.Sm.,
Oberonia equitans var. chaperi Finet, Oberonia lampongensis J.J.Sm., Oberonia mcgregorii Ames, Oberonia murkelensis J.J.Sm.,
Oberonia oxystophyllum J.J.Sm., and Oberonia palawensis Schltr. We designate here as neotype for Oberonia palawensis Crain 198
US 3737901. The synonymies are supported by floral characters and habit of each named taxon, ecology and phenological data,
and lack of differentiation in a molecular phylogeny. The species is shown to be widely distributed from Malaysia through French
Polynesia based on examination of some 300 herbarium specimens. If the magnitude of over-naming in Oberonia is indicative of
other intrageneric groups that are cryptic and/or difficult to identify in the field or from pressed herbarium specimens, diversity
could be over-estimated by a third in such groups.
INTRODUCTION Oberonia is a genus of
tropical and subtropical orchids ranging
from Central Africa eastwards to French
Polynesia. On the order of 470 names have
been introduced for 200–300 species. The
number of currently recognized species
is by far too high, with large numbers of
synonymies to be addressed (Bunpha et
al. 2019; Geiger 2016, 2019a, in press a,
unpubl. data).
Difficulties in sorting out identities
of names arise from species that lack
types and that were never illustrated.
Furthermore, verbal descriptions are
frequently difficult to match to biological
species, particularly from geographic
regions that have limited material in
herbaria, such as the Republic of Palau in
the Caroline Islands of Micronesia.
Fieldwork in Palau provided the
impetus for the current study (Crain
2018). While various sources suggest that
Palau is home to at least three species
of Oberonia (Costion 2009, Costion and
Lorence 2012, Hillman-Kitalong and
Uesugi 2017), herbarium specimens for
most of the species are sparse, and some
of the type specimens are missing and/
or questioned (Fosberg and Oliver 1991,
Natural History Museum (NHM) 2014).
Oberonia palawensis Schtr. is a case in
point. Ledermann’s type specimen for the
species (Ledermann 14475) is missing and
only a few other specimens of the species
are known to exist (Fosberg and Oliver
1991). Several specimens are housed
in US: Hosokawa 9100; Canfield 367,
666; Timberlake 3168; Fosberg 25740;
Bowden-Kerby LR5796; Evans 619. Two
recent collecting trips to Palau yielded two
additional Oberonia specimens that were
preliminarily identified as O. palawensis.
Each of these specimens was examined in
detail to confirm its identity.
While we worked out the identity
of the Palau specimens, it became clear
that a much more far-reaching taxonomic
issue needed to be tackled. We used
a combined morphological–molecular
approach to demonstrate that multiple
described species all refer to one and the
same biological species. We identified
10 new synonyms of Oberonia equitans
(G.Forst.) Mutel in addition to the three
already accepted ones.
MATERIALS AND METHODS Light
Microscopy. Flowers on a herbarium sheet
were imaged by applying z-stacking on
a Zeiss Discovery V20 stereomicroscope
with motorized focus and a 1.5x
planapochromatic objective lens. Files of
the stack were processed in ZereneStacker
using the Pmax algorithm and then further
adjusted in AffinityPhoto. For details see
Geiger (2013, 2017).
Electron Microscopy. For scanning
electron microscopy (SEM), preserved
flowers were brought to 100 percent
ethanol through three changes of 100
percent ethanol. The flowers were
then critical-point dried in a Tousimis
Autosamdri 815A (Tousimis, Rockville,
Maryland) using default settings,
mounted on double-sided carbon tabs
(PELCO Image Tabs, Ted Pella, Redding,
California) on aluminum stubs and coated
with gold in a Cressington 108Auto with
rotary planetary stage (Watford, United
Kingdom). The flowers were imaged on a
Zeiss EVO 40XVP or Zeiss EVO 10 LS SEM
in variable pressure (30 Pa) at 20 kV and
50–500 pA depending on magnification
and working distance.
Gatherings found in multiple
institutions are cited with the most
complete data. Standard repository
abbreviations are used: DLG/HOAG
indicates Daniel Geiger living collection/
Geiger herbarium and spirit collection,
Santa Barbara, California. For elevation
and phenology information, data plots
with local regression least square fitting
were generated with DataGraph 4.3
(Visual Data Tools).
Molecular Analysis. Vegetative tissue samples were collected from four
naturally growing individuals of Oberonia
in the Republic of Palau to facilitate
molecular identification. Each leaf and
root tissue sample was stored in a sealed
2-ml microcentrifuge tube filled with
CTAB buffer solution to preserve DNA,
and then frozen at −20 C until processing
in the laboratory. For DNA extraction,
each tissue sample was removed from
the tube with buffer and clipped into
small fragments into a well of a 96-sample
deep-well plate. For herbarium plant
samples, a small (~1 cm2) sample of leaf
was similarly clipped into small fragments
into a plate well. We extracted DNA from
approximately 25 mg of plant tissue using
a BioSprint 96 automated DNA extraction
workstation with a BioSprint96 Plant DNA
kit (Qiagen, Inc., Valencia, California)
following the supplied protocol. After
DNA extraction, we amplified the matK
region of the chloroplast using matk1F/
matK1R (Parveen et al. 2017) in a BioRad C1000 Touch Thermal Cycler (Bio-Rad
Laboratories, Inc., Hercules, California).
We also amplified the internal transcribed
spacer of the nuclear ribosomal repeat
(ITS) and chloroplast trnH/psbA. These
two additional loci amplified poorly from
the herbarium specimens and gave similar
phylogenetic patterns, so only the matK
is presented here. We prefer an analysis
based on a complete data matrix of fewer
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markers to one with more markers but
significant missing data.
Polymerase chain reaction (PCR)
success was evaluated by electrophoresis
on a 1 percent agarose gel, stained using Gel
Red, and visualized on a transilluminator.
All successful PCR products, defined as
those producing a single band, were
sequenced using Sanger Sequencing.
PCR product was cleaned using ExoSapIT (Applied Biosystems, Inc., Foster City,
California) and subjected to sequencing
reactions using BigDye v.3.1 (Applied
Biosystems, Inc.) in quarter-strength half
reactions, using 1 μl Big Dye 3.1, 2 μl Big
Dye buffer, 1 μl 3.2 μmol primer, 5 μl H2O
and 5 μl (50–80 ng DNA) of PCR product,
and otherwise followed manufacturer
protocols. Sequencing reaction products
were cleaned using Sephadex G-50
Fine (GE Healthcare Systems) and dried
and analyzed on an ABI 3100 Sanger
Sequencer. Sequences were manually
checked for quality and trimmed. Forward
and reverse sequences were combined
into a single contig and discrepancies
resolved manually in Sequencher v.5.4.6
(Gene Codes Corp.).
Phylogenetic analysis. We
downloaded all Oberonia matK sequences
from GenBank and aligned our sequences
and the downloaded sequences using
MUSCLE, implemented in Geneious Prime
2020.0.4 (BioMatters Ltd.) and checked
manually. We retained a subset of the
downloaded sequences to represent each
clade and to demonstrate the amount
of variation within a species. We used
this approach to allow us to focus on
phylogenetic identity of our focal samples,
rather than on possible synonymies and
identification issues in the genus as a
whole. Malaxis tenuis (AY907196), Liparis
liliifolia (AY907156), and Stichorkis gibbosa
(KJ459318) were used as outgroup taxa.
We then estimated phylogenetic trees
using MrBayes, implemented in Geneious
Prime 2020.0.4 (BioMatters Ltd., Auckland,
New Zealand) and using a burn-in of
110,000, sample frequency of 200, total
chain length of 1,100,000, four heated
chains, a heated chain temperature of
0.2, GTR substitution model and gamma
rate variation.
M O L E CU L A R R E S U L T S T h e
molecular
results
support
the
morphologically derived synonymies (Fig.
1). The Palau specimens are interspersed
with those from Samoa and French
Polynesia, mostly in a major polytomy
with rather short terminal branches. Two
samples are separated in a distinct clade.
However, the cumulative branch lengths
1
of less than 0.5 percent within O. equitans
are as long as or shorter than the terminal
branch length in Oberonia cavaleriei Finet
and Oberonia rufilabris Lindl. Those two
species are easily identified and distinct.
Accordingly, the cumulative branch lengths
within O. equitans are compatible with a
single species. Furthermore, the largest
genetic differences are encountered
within the rather small islands of the
Palauan Archipelago, while the much
more distant samples from Samoa and
French Polynesia show no phylogenetic
patterning at all, as they are found in
a large polytomy. The voucher for O.
equitans from Cameron (2005) T. Motley
& K. Cameron 2255 was most likely from
New Caledonia (K. Cameron, pers. comm.)
but the specimen cannot be found at the
New York Botanical Garden herbarium
(A. Weiss, pers. comm.) and is presumed
lost. This sample was also positioned
within the same polytomy as the Palauan
samples and the more distant samples.
[1] Oberonia spp. matK phylogenetic tree
inferred using MrBayes. The tree shows
the relationships between field-collected
samples in bold, herbarium specimens,
and selected GenBank sequences for
references. Values at each node are
posterior probabilities. Insert: In situ photograph of Oberonia equitans in Palau
courtesy of B.J. Crain.
We consider this detail of the topology a
random result of limited sampling.
Sample Oberonia sp. B.J.Crain 160L
was conservatively not included under O.
equitans. The specimen was not in flower,
for which reason the identity could
not be positively confirmed. Given the
extensive vegetative phenotypic plasticity
of Oberonia spp., the overall rather poor
understanding of distributional limits of
species in the genus, and the fact that
three species of Oberonia have been
reported from Palau (none of which
have molecular data for comparison),
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geiger, et al.
we take the conservative approach of
referring to it as Oberonia sp. The internal
and cumulative branch lengths are
compatible with either a single sample of
a distinct species, or a further sample of
O. equitans.
The tree is well supported at the
relevant nodes with posterior probabilities
>0.94. Some internal nodes are less well
supported, with posterior probabilities as
low as 0.55, which should be considered
unresolved. Those low support values
are most likely due to incomplete taxon
sampling of a genus with approximately
200 species.
The marker matK may be considered of
insufficient discriminating power by some
who prefer a set consisting of rbcL + matK
+ ITS. The clean results with samples from
the same species forming well-supported
clades in every single case (O. cavaleriei,
Oberonia delacourii Gagnep., O. rufilabris,
Oberonia recurva Lindl.) indicate that for
the purpose of this investigation matK
provides the necessary information.
Additionally, taking into account withinand between-species branch length and
the associated morphological data further
supports our conclusion.
Li et al. (2016) found a similar short
branch length polytomy for 15 samples of
Oberonia jenkinsiana Griff. ex Lindl. and
its not yet formally recognized synonym
Oberonia austro-yunnanensis S.C.Chen &
Z.H.Tsi in their Bayesian phylogeny based
on matK + ITS, while other species such
as the 13 samples of Oberonia caulescens
showed more structuring. Terminal and
internal branch lengths varied in a similar
way as in our analysis, and support values
were similarly strong. Relative branch
lengths were similar in species common
to both studies. The terminal and internal
branch lengths of O. rufilabris were
longer than those of O. cavaleriei, which
were longer than those of O. delacourii,
suggesting that matK is an adequate
estimator for the assessment of species
boundaries.
As an aside, the spelling of O. austroyunnanensis should retain the hyphen
in accordance with ICN Art. 60.11 Ex. 41
(contra World Checklist of Selected Plant
Families 2020).
SYSTEMATICS
Oberonia equitans (G. Forst.) Mutel,
1837
Basionym. Epidendrum equitans
Forster, 1786: 60. Type. G. Forster 170
lectotype BM (not seen; designated by
Kores 1989: 57). G. Forster 192 isotype/
isolectotype BM000084364, P [not seen
in 2019, not in P database]. Possible
isotype BM 000082099 Forster s.n., sine
loc. Isolectotype Moscow State University
Herbarium, fide US herbarium sheet
photo. Tahiti.
Oberonia
glandulosa:
Lindley,
1859: 6. [nomen illeg.]. Type. Kartalsky
(Prescott) s.n. (K), Matthews 158 (syntype
E00373990, K s.n.), Otaheite [=Tahiti].
No lectotype has been designated (Kores
1991).
Oberonia aurea Schlechter in K.
Schumann & Lauterbach, 1905: 109.
Type. Schlechter 14673 (syntype: B
lost). Mountain forest near Punam, Neu
Mecklenburg, 600 m. Syn. nov.
Oberonia ciliolata Hooker, 1890:
181. Ridley s.n.; Ridley [375] (syntype K
000943006: basis of Hooker (1895: pl.
2318), SING 0047511); not BM000088559:
is lectotype of O. dissitiflora, see remarks.
Krangi, Singapore. Syn. nov.
Oberonia oxystophyllum J. J. Smith,
1905: 237–238. Type. Herb. Lugd. Bat.
904, 84–127, now [Korthals s.n.], syntype
L 0091768, [Hallier s.n.] K 000942986),
Gede, near Tjibodas; Patoeba, Java,
[Indonesia] [6.788S 106.982E]. Reference
by Smith to —128—130? cannot be
considered type material, because it was
not unambiguously assigned to the taxon.
Syn. nov.
Oberonia flexuosa Schlechter, 1906:
62. Type. Schlechter 15496 (syntype
B: lost). On trees along streams in
the mountains near Ou Hinna, New
Caldeonia.
Oberonia mcgregorii Ames, 1907:
321–322. Type. R. C. McGregor 291
(syntype AMES 9919/Harvard Barcode
00101991), Balete, Baco River, Mindoro,
Phillippines. Syn. nov.
Oberonia equitans var. chaperi Finet,
1908: 337, pl. 10, figs 29–30. Type.
(iconotype), ubi?
Oberonia lampongensis Smith, 1917:
22. Type. H. A. Gusdorf living Culture In
Hort Bogor. sub numero 62 pro parte
(syntype). Lampong near Menggala,
Sumatra. Syn. nov. of O. equitans.
Oberonia affinis Ames & C.
Schweinfurt in Ames, 1920: 79–81, pl. 89,
figs. II, 2. Type. Clemens J. 102 syntypes
AMES 16978, 16979; Harvard Bar Code
00101935, 00101936, BM000088335,
E00394089, F 493793, K 000942995, MO
68377, MO 6837753, P02291942, SING
0043926, US 1170971, Lobang, Cave,
5,000 feet (= 1,666 m). Clemens 275,
syntype, Marei Parei Spur. Clemens 380,
syntype, Kiau. Syn. nov.
Oberonia palawensis Schlechter,
1921: 462–463. Type. C. Ledermann
14475 (syntype B: lost: Fosberg and Oliver
1991), Palau, near Ngatkip on Babelthaob,
Palau, 50 m (7.382N 134.513E). Neotype
Crain 198 US 3737901 here designated.
Ngardok Nature Reserve, Melekeok,
Babeldaob, Palau. Syn. nov.
Oberonia enoensis Smith, 1928a: 454–
455. Type. Toxopeus 94 syntype L0061740,
Boeroe: Wai Eno, 600 m (protologue).
Buru, Maluku, Moluccas, Indonesia (type
label). Syn. nov.
Oberonia murkelensis Smith, 1928b:
125. Type. Kornassi ex Ruten 1457,
syntype L 0061788, Goenoeng Moerkele
[= Gunung Murkele], Central Seran,
[Sumbawa], Moluccas, Indonesia, 1000–
1900 m. Syn. nov.
MATERIAL EXAMINED
Thailand. Kerr 610 K 0000596120.
Bangkok.
Malasysia. UNESCO 420, 421 SING
0141326. Ulu Kelantan, Gua Musang,
Malaya. UNESCO Limestone expedition
1962 421 K s.n. Gua Muang, Ulu. Native
Collector/Synge 447 K s.n. Mount Dulit,
Dulit Ridge, Sarawak, 1,240 m. Haviland
s.n. SING 0141407. Sarawak. Hewitt
15 SING 0141408. Sarawak. Carr s.n.
SING 0141415. Mount Kinabalu, Koung,
Sarawak, 400 m. Henderson 332 SING
0141327. Kot Glanggi, Pahang. Henderson
22446a SING 0141321. Kota Glanggi,
Pahang. Mot Nur s.n. SING 0141322.
Bukit Sagu, Pahang, 300 m. E Rostado
s.n. SING 0141323. Bundi, Tringganu.
R Denny 313 SING 0141324, 0141328.
Sungli Rambai, Malacca. Ridley 313 BM
000088324. Malacca. R Derry s.n. SING
0141326. Sungei Kesang. Lugas 1489 K
s.n. Sabah, Kampung Melangkap Tomis,
Sekitar Kampung Melangkap Tomis, 400
m. Carr 3659, SNF 27982 SING 0141402,
SING 0022049 [spirit] Sabah, Bundu
Tuhan, 1,400 m. Carr SFN 26371 SING
0141401 Sabah, Dahobong River on
label, Tahubant River in citation, 1,100
m. Clemens 40924 B s.n., E 00616203,
K s.n. Sabah, Penibukan near Pinokkok
Falls, 1,600–2,300 m. Clemens 40822 BM
000088329, K s.n. Sabah, Penibukan, belos
Pinokkok falls, 1,500 m. Clemens 50240 B.
s.n., BM 000088331, K s.n Sabah, Mount
Kinabalu, Tenomopok, 1,650 m. Clemens
40861 BM 000088330 Sabah, Mount
Kinabalu Penibuka, Side Ridge E of camp,
1,350 m.
Papua New Guinea. Carr 10494 CANB
61947, SING 0141467. Lala River, 1,770
m. Carr 17128 BM 000088388. Kokoda.
Carr 17084 BM 000088410. Kokoda. Carr
10494 BM 000088415. Lala River, 1,800
m. Millar NGF 38328 CANB 212585, K
s.n., SING 0141492. Watabung, Goroka
subdistrict, Eastern Highlands, 2,300
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m. Millar NGF 38328 K s.n. Watabung,
Goroka subdistrics, Eastern Highlands,
2,300 m. Millar & Dockrill 22868 K s.n.
Island of Kui Morobe, 15 m.
Singapore. Ridley s.n. BM 000088324.
Krangi. Ridley 2034 BM 000088337, MEL
s.n., SING 0010910. Cahn ctran Kang,
23 m. Ridley 2034a BM 000088377, K
s.n., MEL s.n., SING 0010908. Changi. JJ
Smith s.n. SING 0010913. Suna Murai?.
JS Goodenough s.n. SING 0010912.
Krangi. Ridley 10153 SING 0010907. Bukit
Tumiak? Road. Sinclair 5129 E00616464.
Seletar forest behind Kee Soon Village.
Indonesia. Balgooy 3669 K s.n. Lake
Matano south, between Soroako and
Matano, Sulawesi, 400 m. SBGO 3291
SING 0141504. Sulawesi, Desa Baruppu,
Kampong Bubuk, 1,600–1,700 m. de
Vogel 6062 K s.n. Selatan, N shore of
Lake Matano, E of Nuha, Sulawesi, 450
m. Vermeulen & Dustermaat 972 K s.n.
interior zone, along trail Long Pa Sia–Long
Samado, near crossing with S Malabid,
Sabah, 1,300 m. SBGO 3392 SING 0141501.
Lower slopes of E flank Rantepao-Paloppo
divide, Sulawesi, 300–400 m. SGBO 3393
SING 0141503. Lower slopes of E flank
Rantepao-Paloppo divide, Sulawesi,
300–400 m. Alston 13852 BM 000088423.
Kambahan, near Lubuksikaping, Sumatra,
300 m. Haviland 841 K s.n., Borneo. JJ
Smith 22 SING 0141440. Tiibodas?, Java.
Zollinger s.n. W Reichenbach 12598. Java,
500 m. S. Darutan 355 BM 000088341.
Borneo, 1 mile of Kahung, downstream,
430 m. Everard Im Thurn 34 P 00310614.
[Java] Drake Range, W of Matakin, Colo
North, 1,300 m. Comber 1395 K s.n. Java,
Arjuno Weliran, NW above Trawas, 1,370
m. Comber 1559 K s.n. Java, G Lamongan
S of Probolinggo, 510 m. Comber 1557
K s.n. Java, G Raung S., 1,100 m. SFN
8155 SING 0141439. Java, Tjibodas on G
Godela, 1,500 m. Chaper s.n. P 00364386.
Borneo. de Vogel & Vermeulen 7185 K
s.n. Celebes, Utara Bolaang Mongondow,
Gunung ambang Nature Reserve, Danau
Mooat area, 1,000 m.
Phillippines. Reillo 16 MO 799956. Luzon,
Laguna.
Palau. Canfield 666 US 3293387. Oreor
[= Koror]. Timberlake 3168 US 3293383.
Aimeliik, Babeldaob (note: specimen is
listed as coming from the island of Oreor
[= Koror]; however, the State of Aimeliik
is on the island of Babeldaob, which is
just north of Koror). Fosberg 25740 US
3293386. Babelthuap [= Babeldaob], 0–
20 m. Bowden-Kerby Lr 5796 US 3293385.
Babelthuap [= Babeldaob], 0–10 m. Evans
619 US 3293384. Aulupse’el = [Ulebsechel]
?, Risong Bay, Koror, 0–50 m. Canfield
367 US 3293388. Ngatpang, Babelthuap
[= Babeldaob], 5 m. Hosokawa 9100 US
3726606. Babelthuap [= Babeldaob]. Crain
134 US 3694789. Ngeruktabel, Koror.
Crain 198 US 3737901. Ngardok Nature
Reserve, Melekeok, Babeldaob. Canfield
366 US 3296687. Ngelobel (Ashakasengu)
Island, S Koror Municipality, inlet on NW
side of island, 1 m. Timberlake 3168 US
32993383. Skillang’s Paddok (Tulau),
Imutsubech, Aimeliik State.
New Caledonia. Vieillard 3296 P
00081696, 00081697. Whitmee s.n. BM
000088469. Loyality Islands Lifu. MacKee
21978 P 00081685. Oue Koura, haute de
Dothio 40 m. Le Rat s.n. P 00081684. Sable
Unio. MacKee 26296 P 00081686. Kone,
Mount Tandji, 800–900 m. MacKee 31023
P 00081687. Col d’Amieu, mont Pembai,
800 m. Hurliman 1163 P 00081683. Paoué
valley, south flank, Tipindje, 500 m. MacKee
33678 P 00081688. Hienghene, Kavatch,
200 m. MacKee 42529 P 00081680.
Noiumea, Haute Amoa, Pomanhou, 500
m. Veillon 2123 P 00081690. Katrikoin,
Launay, 300 m. Dagostini & Barriere 1281
P 02102974. Tiebahi, 500 m. Guillaumin
9866 P 00081682. Between Ponerihonen
and Honaiton. McPherson 2562 P
00081701. Mount Panié, 20 air km NW of
Hienghénee, 500 m.
Vanuatu. Macdonald 4 MEL 569474.
Aneitum. HF Moore 277 US 00241464.
Maewo. Morrison s.n. K s.n. Efate Nudine
Bay. Morrison s.n. K s.n. Hills between
Unum & Anilgnkot/Anitzem. Morrison s.n.
K s.n. Efate Hills Nudum Bay. Whatley 104
K s.n. Pentecost Village, Ena, 290 m. Smith
1485 K s.n., P 00310616. Vanua Mbalavu,
0–200 m. Schmid 3624 P 00310683.
Anatom. Raynal 15994 P 00310684.
Ikouroup Tanna.
Wallis and Futuna. Veillon 5108 P
00310685 Futuna, Alofi, north slope 300
m. Hoff 4053 P 00310687. Futuna, Mount
Puke, 450–500 m. Morat 7084 P 00310686
Mount Lulu.
Niue. Williams 9598 MICH s.n. S of
Alofi village, 20 m.
Samoa. Le Guillou s.n. P 00310618.
Sin. coll., s.n. W 3066. Savaii approx.
100 m. WA Whistler W2638 K s.n., US
00241470. Savai’i, W of Mauga Mu
1550 m. Rechinger 1589 W Reichenbach
3063. Savaii Aopo, 300 m. Rechinger 16
W Reichenbach 3064. Savaii, Lalatelle.
Rechinger 127 W Reichenbach 3062.
Savaii, between Sassina and Aopo. Graeffe
s.n. W Reichenbach 8372 W68800. Upolu.
HE Parks 16237a US 00241481. Upolu,
near Tiai, 720 m. WA Whistler W204 US
00241469. Upolu. F Reinecke 184 US
00241483. Upolu. WA Whistler 3946
US 00241456. Namua. Reinecke 184 E
00616504, WU s.n. Upolu, Vailale ridge.
Whistler 2906 K s.n. Tutuila, Mount Tau,
350 m. Mansfeld 172 K s.n. Upolu, swamp
near Tiavi, 720 m. K & L. Rechinger s.n. W
Reichenbach 3065. Tutuila, Pago-Pago.
Tonga. Parks 16237 MO 1035159.
Eua, Plateau, exposed rocks. Parks 16237A
BM 000088471, US 00241482. Eua,
Plateau. Yucker 15402 BM 000088470, US
00241473. Eua, above Riechelmann’s Fuai
plantation near center of island, 245 m.
Cook Islands. A. Whistler 5825
US 01232921. Mangaia. Sin. coll., s.n.
Reichenbach 37743 W68916. Mangaia
Island. WA Whistler W5038 US 00447248.
Rarotonga, E of Ikurangi. WR Phillipson
10132 US 00445241. Luttril 350 K s.n.
Rarotonga. Cheeseman 675 K s.n.
Rarotonga, central part of island. Wilder
8 K s.n. Rarotonga.
Fiji. Seemann 588 MEL s.n., BM
000088468, K s.n., P 00310615, W
Reichenbach 37814. Navua and Manar.
Parham 1823 K s.n. Naitasiri, above
Wamawaya, 50 m. Daanu 82 K s.n.
Kurovisilou, Serau, 100 m. AC Smith
7075A US 00241472. Viti Levu. AC Smith
7469 US 00241471. Ovalau. AC Smith
7551 US 00241465, K s.n. Ovalau, Valley
of Mbureta and Lovin Rivers, 20–50 m. AC
Smith 8586 US 00241466, K s.n. Viti Levu,
Namosi Hill, bordering Winavindrau Creek
in vicinity of Wainimakutu, 150–200 m. AC
Smith 1485 US 00241479. Eastern Division,
Vanua Balavu (Mbalavu). AC Smith
5241 US 00241476. Viti Levu, Western
Division. AC Smith 5528 US 00241475, K
s.n. Viti Levu, Western Division, Nandroga
and Navosa, N portion of Rairaimatuku
Plateau, between Nandrau and Nanga,
725–825 m. AC Smith 5751 US 00241474,
K s.n., P 00310613 [top specimen only].
Viti Levu, Central Division, Naitasiri, N
portion of Rairaimatuku Plateau between
mount Tomanivi/Victoria and Nsasonggo,
870–970 m. Greenwood 1112 K s.n. Viti
Levu, Naitasiri, near Nasinu. Vaughan
3181 K s.n. Viti Levu, Tamavna Ridge, mile
6, 200 m. Meebold 16813 K s.n. Suva, 9
miles above Suva. Greenwood 1155 K s.n.
Viti Levu, Lautoka, Mount Evans, 750 m.
Smith 1731 K s.n., P 00310617. Vanua
Levu, Mbua Lower Winunu river valley,
0–200 m Greenwood 209 K s.n. Mount
Lautoka, 260 m Tothill & Tothill 511 K s.n.
Suva, Central Road. Tothill & Tothill 883 K
s.n. Viti Levu, Suva, Central Road. Parham
147 K s.n. Viti Levu, Rewa Province,
Mount Korobaba, 260 m. Vaughan 3181
BM 000088472. Tamavua Ridge, Mile 6,
200 m.
French Polynesia. Tahiti. J Banks &
www.AOS.org august 2020 © American Orchid Society Orchids 659
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DC Solander 430 US 00241463. Banks
& Solander s.n. F 694765. Expedition
Novara 117 W68918. Sin. coll., s.n. W
Reichenbach 11734 W68802. Setchell
& Parks 430 P 00310665. Richard s.n.
P 00310674. Pancher 3 P 00310663.
Vieillard 3 P 00310668. Vesco s.n. P
00310667. Vesco s.n. P 00310666.
Ribiourt 82 P 00310676. Vieillard s.n.
P 00310672. Vieillard s.n. P 00402800.
Vieillard s.n. P 00310673. Morrenhout
s.n. P 00310677. Lepine s.n. P 00310678.
Lay & Collie s.n. BM 000088474. Sin. coll.
s.n. BM 000088473. Shuttleworth s.n. BM
000088473. Grandjean s.n. P 017712107.
WA Setchell & HE Parks 430 US 00241462.
Florence 2482 P 00310669. Mount
Marau, Sentier du Pic Vert 1,240 m. Hallé
6931 P 00310680. Tahiti, route of Mount
Marau 700 m. Hallé 6937 P 00310679.
Riv. Vaihiria valley 50–60 m. Hallé 6993 P
00310671. Punaahuia, S of Papeete, 300
m. Raynal & Taureau 16546 P 00310664.
Road of Aorai, between Fare Hamuta
and le Belvedere 950 m. Savatier s.n. P
00310675, P 00310682. Vallée de la reine.
ML Grant 3613 US 01104522. Punaauia
Diadem district, 491 m. FR Fosberg 63613
US 00619673. 5–20 m. FR Fosberg 62965
US 00619672. Papeete, headwater of
Tipaerui River, 950–1,000 m. Florence &
Sykes 11342 US 00689598. Papeete, 80
m. Balgooy 1677 MO 2228544. Mahina
Valley, 700 m. ML Grant 3613 P 00310653.
Punaauia Diadem district, 491 m.
Bora Bora. D’Urville 1061 P 00310645.
D’Urville s.n. P 00310650. MMJ Balgooy
1942 CANB 00583505, K s.n. W slope of
Pahia, 100 m. Florence 12107 P 00360516.
Faanui, Mount Pahia, E flank, 420 m.
Huahine. L Dunn 511 US 01076801.
Mouly 496 P 00592106. Flancs of the
Vaiparao valley, 200 m. J Florence &
Tahuaitu 11583 P 00310656, US 00445241.
Fare, Mount Mato Ereere, N crest, 425 m.
Florence & Tahuaitu 11566 P
00310655. Fare, Pipihaua valley, right
flank, 130 m.
Moorea. HM Smith 8 US 00241458. FR
Fosberg 63663 US 00619701. M-H Sachet
2542 US 00619665. FR Fosberg 60957 P
00310569, US 00619674. Ridge above
Le Belvedere, head of Hpunohu Valley.
Cronk et al. T62 E s.n. SW valleys, Patiri
valley, 300 m. Raynal 17930 P 00310648.
Moo Afaretaitu. Jolinon 1334 P 00310654.
Vaiare hill summit. Smith 8 P 00310660.
Opunohu–Roroie District. Florence 8318
P 00310661. Haapiti, crest between the
high valleys of Vaianae and Atiha, 350
m. Birnbaum 75 P 00310657. Afareiatu,
Haute Nauroa 700 m.
Makatea. GP Wilder 1141 K s.n.,
US 00241480. 60 m. Jones 861 K s.n., P
00310620. 100 m.
Meetia. St. John 14198 F 1438060,
K s.n., MO 1638487, P 00310643, US
00241478, W Reichenbach 5315 W68919.
Fatia-po to Fareura, 200 m.
Raiatea. GH Gagné 1786 US
00619667. 475 m. Welch et al. 26616 MO
5796911. Trail to Temehani Ute Ute, 450
m. Welch et al. 26619 MO 5796913. Trail
to Temehani Ute Ute, 400 m. Moore 129
MO 2196179. Valley W of Uturoa, 280 m.
Moore 410 MO 2196301. Opua Valley, 100
m. Morat 9683 P 00310649. Temehani
ute, 150 m. Florence 10351 P 003110658.
Uturoa, Mount Tapioi, 230 m. Mouly 377
P 00591985. Vaiaou valley, 200 m.
Funaauia. Florence 8195 P 00310652.
Punaruu valley, Fare Anani, 430 m.
Raupiti. Raynal 17854 P 003106466,
P 003106467. Mount Tiriano, 260 m.
Tahaa. Florence 10631 P 00310662.
Mount Purauti, SE crest, 245 m.
Austral Islands. Ruruto. Hallé s.n. B
s.n. N part. Hallé 7284 P 00310630. N of
Avera. Forence 5567 P 00310635. Unaa,
N of Tetuanui plateau, high Vaire, 205 m.
Hallé 7163 P 00310628, US 00241452.
Naairoa. Hallé 7082 P 00310631. Prise
d’eau de Tetuanui. H St. John 16741 F
1438056. Arei, 75 m. N Hallé 7239 P
00310629, US 00241451. Middle Vaitotai.
Florence 9240 P 00310623. Rurutu, Rairiri,
East, below T Anaoeva, 110 m. Florence
9180 P 00310622. Tetuanui Plateau, NW
sector, 210 m. Hallé 6648[b?] K s.n. Middle
Saitotai. Forence 5703 P 00310636.
Raivavae. Anatonu, foot of the cliff
below Mount Hiro, 160 m. Florence &
Sykes 11342 P 00310621. Vaiiuru, Turani
valley, 80 m. N Hallé 7082 CANB 8602862,
K s.n., US 00241453. N of island, spring of
Cetuanin.
Rapa. G Paulay 85 US 00619675. 200
m. Hallé 7694 P 00310632. SSE Pukumia,
185 m. Hallé 7619 P 00310633. SSE
Pukukia, 200 m. Sykes 161 CHR 495761 K
s.n. Valley behind Vairua. John & Fosbert
15976 K s.n., P 00310634. Pic Rouge, S
slope, 80 m.
Tubuai. SH Shomer 6761 US 00241468.
SH Shomer 6692 US 00241467. 300–
325 m. Hallé 6648 K s.n., P 00310627,
US 00241455. Below coffee plantation
of Tamatoa, 25–35 m. Hallé 6884 P
00310624. N of Mahu, 250 m. Hallé 6673
P 00310626, US 00241454. Mahu. Hallé
6755 P 00310625. Mont de la Prise d’eau
de Mataura, 60–80 m. St. John 16349 MO
1638532. Taitaa NE slope, 330 m.
REMARKS
In the genus Oberonia, the species
epithet equitans is highly confused
because it was introduced by several
authors, with subsequent authors creating
primary and secondary homonyms:
• Epidendrum equitans Forster =
Oberonia equitans (Forster) Mutel, correct:
Malaya to Western Pacific Islands.
• Cymbidium equitans Thouars =
Oberonia equitans (Thouars) Lindley,
synonym of Oberonia disticha (Lam.)
Schltr.: East Africa.
• Malaxis equitans Blume = Oberonia
equitans (Blume) Lindley, junior synonym
of Oberonia padangensis: Malaya.
Subsequent authors have not
always been aware of those homonyms,
commonly adding the wrong authority
to the record. The same also applies to
identifications on herbarium specimens.
This has led to widespread confusion
regarding the identity and distribution
of Oberonia equitans. Sometimes range
indications of “equitans” were combined
from multiple discrete taxa, leading to
a cited distribution from East Africa to
French Polynesia (e.g., Finet 1908).
The lectotype of O. equitans is labeled
Forster 170, but in the publication it is
erroneously referred to as Forster 316
(Kores 1991). The lectotype designated
by Kores (1989) at BM and the cited
isolectotype at P could not be found in
those collections during visits in 2016
and 2019, respectively. They are not
recorded in the online databases either.
The whereabouts of those specimens is
currently unresolved. The only confirmed
type known to exist is a Forster s.n. sheet
at BM, which agrees with the commonly
accepted species concept of O. equitans
(Forster) Mutel.
Oberonia glandulosa is a wellestablished synonym of O. equitans (e.g.
Cribb and Whistler 1996, 2011) and is not
further discussed here.
Oberonia aurea is a synonym of O.
equitans. The common attributes include
the habit, the hairy rachis, the hairy
pedicelled ovary, the hairs on the external
surface of the floral bract and the sepals,
the lip with small auricles, constricted
mesochile, bilobed and incised epilobe,
flower color from pale-green yellow
turning to dull orange with maturation
(Fig. 2).
Oberonia ciliolata is a clear synonym
of O. equitans (Forster) Mutel. The type of
O. ciliolata is confusing and requires some
clarification. Hooker (1890) designated a
specimen collected by Ridley without
gathering number (Ridley) or repository
as type. The specimen cited at Kew is the
basis for Hooker’s (1895, pl. 2318) figure.
Accordingly, this gathering (Ridley
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375) can be considered the type gathering,
with the Kew specimen constituting a
syntype. The same gathering number was
used twice by Ridley, and that is the type
gathering of Oberonia dissitiflora Ridley.
The overall caulescent habit, the
papillose rachis, papillose pedicelled
ovary and papillose back of tepals, the
deeply incised acute, acuminate bract,
and the identical shape of sepals, petals
and bract clearly show conspecificity of
O. ciliolata and O. equitans. Oberonia
equitans was applied to specimens from
the Western Pacific Islands, while O.
ciliolata was applied to specimens from
the Malayan Archipelago.
Oberonia oxystophyllum has the
same habit and floral characteristics as O.
equitans, making the two synonyms. The
hairy bracts, hairy column, hairy back of
the sepals, the oblong shape of the petals,
the lip with moderate auricles, constriction
of the mesochile, and bifurcated epichile
with incised apical lobes are all shared
characters.
Oberonia flexuosa is a well-established
synonym of O. equitans (e.g. Cribb and
Whistler 1996, 2011) and is not further
discussed here.
Oberonia mcgregorii shares the habit,
the hairy backs of the flowers, the shape
of the petals and every aspect of the lip
shape with O. equitans, which makes the
two synonymous. Oberonia mcgregorii
has been cited for the Phillippines.
The illustration of Oberonia equitans
var. chaperi clearly represents O. equitans
s.s. (G. Forst.) Mutel. Finet (1908)
thoroughly mixed the two equitans in his
treatment (see above) and his diagnostic
characters (hairs on pedicelled ovary, all
tepals, bract) address the distinction of
Oberonia disticha from East Africa and
O. equitans from the Pacific region. Due
to this elementary confusion, he did not
describe a variety of O. equitans s.s.,
hence, his variety is unjustified.
Oberonia
lampongensis
has
previously been recognized as a synonym
of O. ciliolata (e.g., Comber 1990). With
Oberonia ciliolata as a synonym of O.
equitans, O. lampongensis becomes a
new synonym of O. equitans.
Oberonia affinis is described in the
protologue as having hairs on rachis,
pedicelled ovary and back of sepals,
all indicative of conspecificity with O.
equitans; those characters unfortunately
were not shown in the drawings. The
shapes of the floral elements as well as
the habit of the type is equally compatible
with O. equitans. Accordingly, the two
species are considered as synonyms.
2
It is important not to confuse the type
gathering of Oberonia affinis Clemens 275
syntype of O. affinis with Clemens 275A
syntype of Oberonia triangularis.
Oberonia palawensis is an overlooked
taxon that has rarely been used after
the introduction by Schlechter (1921).
A neotype is here designated for O.
palawansis. The syntype in B has been
destroyed (Fosberg and Oliver 1991); no
other syntypes are known to exist. The
species has never been illustrated and it
is difficult to ascertain the identity of a
taxon from verbal descriptions alone. This
is particularly true for species-rich groups
of microfloral species such as Oberonia
spp. We here clarify the identity of the
name by designating a gathering from as
near as possible from the locus typicus.
The neotype matches the protologue
(Schlechter 1921) in every detail.
Oberonia palawensis is a clear
synonym of O. equitans. The protologue
indicates the erect habit with triangular
leaves, the papillose rachis of the
inflorescence,
papillose
pedicelled
ovary, papillose flowers, and papillose
lanceolate-acuminate bract, and also
specifies the narrow oblong petals.
The cited commonalities offer
abundant evidence for conspecificity.
The shape of the petals shows some
intraspecific variability from more narrow
oblong so somewhat broader triangular.
Once a large number of specimens have
been examined, it becomes clear that
those are merely the extremes of a
continuous character state distribution,
and accordingly, are of no taxonomic
importance. It is merely intraspecific
variability.
The drawing of Oberonia enoensis
is indistinguishable from those of O.
oxystophyllum, O. murkelensis, and O.
ciliolata, all drawn by J.J. Smith. The
[2] Synonyms of Oberonia equitans.
Oberonia affinis: Drawing from protologue and habit of O. affinis syntypes
AMES barcode 00101935. Oberonia
enoensis: Flower illustration by J.J. Smith
and habit of syntype. Oberonia murkelensis: Illustration of flower of O. murkelensis by J.J. Smith. Dissected floral parts
digitally rearranged to match live position,
and habit of O. murkelensis syntype.
Oberonia oxystophyllum: Flower illustration by J.J. Smith and habit of syntype.
Oberonia ciliolata: Stylized figure from
Hooker (1895), drawing from type by J.J.
Smith. Oberonia mcgregorii: Drawing
from Ames (1908), and habit of AMES
syntype. Oberonia aurea: Drawing of
flower, bract and column from Schlechter
(1923). Oberonia equitans var. chaperi:
Drawing of lip and bract from Finet
(1908). Scale bar for habits = 5 cm.
habits of the type specimens of those
respective taxa is equally congruent.
Smith (1928a) noted the similarity of his
O. enoensis with O. lampongensis [= O.
ciliolata]. Congruent characters cited in
the protologue are the hairy pedicelled
ovary, bracts, and sepals, as well as the
light yellow color of the flowers. Oberonia
enoensis is a synonym of O. equitans.
Smith (1928b) placed O. murkelensis
in section Otoglossum (= Menophyllum),
but the spreading epichile of the lip and
the hairs on the pedicelled ovary place
the species in Adenorachis.
As noted above, Oberonia murkelensis
is indistinguishable based on floral shape
and habit of the type. The protologue
compared the new species to Oberonia
subanajamensis (= Oberonia punctata: see
Geiger 2019a) and noted the hairy bracts
as well as the yellow color of the flower,
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3
[3] Oberonia equitans flowers from throughout its distributional range, demonstrating intraspecific variability. Scale bars: flowers = 1 mm.
Scale bar seeds = 100 μm. For details see text.
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all compatible with O. equitans. The
species was described from a herbarium
specimen only. The drawing of the flower
indicates that the specimen was not in the
best condition, which explains why some
of the finer details of flower morphology
were not given. For instance, the sepals
were not described as hairy, but not as
glabrous either, hence, the absence of
additional confirming characters cannot
be taken as disconfirming.
INTRASPECIFIC VARIABILITY With
the combined morphological–molecular
approach we can demonstrate extensive
intraspecific variability. Flowers and
sequence data from Crain 134 from Palau
and the sequences from French Polynesia
serve as anchor points; both are found
in a large polytomy with short terminal
branches. The flowers of Crain 134 (Fig.
3) are typical O. ciliolata morphs with a
rather narrowly constricted mesochile,
more flared epichile lobes, and extreme
pubescence all over the flower. From
French Polynesia, only O. equitans is
known, with a less constricted mesochile,
a less flared epichile with fewer points,
and somewhat less pubescence overall
(Fig. 2: K45717 Vanuatu, K 21015), and
K21015 showing additionally the turgid
flower on an already developing seed
capsule commonly seen in O. equitans.
The few samples with multiple flowers
(K12099 Malaysia and L17887 Fiji),
demonstrate within sample variability in
number and degree of point formation on
the epichile, and degree of constriction of
the mesochile. Absolute size also appears
to be somewhat variable as shown by the
smaller flowers of K20999 from Malaysia.
While those flowers were not entirely
open, they appear to be approximately
one-third smaller compared to the average
size of the remainder. L17887 from Fiji is
somewhat intermediate between the
above-cited sample from Malaysia and
the others.
Extensive intraspecific variability
is increasingly recognized in Oberonia.
Bunpha et al. (2019) correctly
synonymized
two
well-established
names, Oberonia falcata King & Pantl.
under Oberonia anthropophora Lindl.
Those were distinguished based on sizedependent habit, flower size, and length
of epichile lobes. The floristic characters
vary extensively, and additional synonyms
will be added to O. anthropophora.
Horticultural data from plants grown over
multiple years further support extensive
intraspecific variability (Geiger 2018,
unpubl. data). While minute differences
are generally considered significant in the
4
systematics of orchids including Oberonia
(e.g., Averyanov et al. 2019), it becomes
increasingly clear that Comber’s (1990, p.
149) remarks on the “annoying variability”
of Oberonia were correct.
That variability extends to the
vegetative portion of the plant. Cameron
(2005) demonstrated that major lineages
within Malaxideae can be separated
using vegetative characters. However,
that discriminating power does not
extend to the species level in Oberonia.
Our illustrations can only show a limited
sample of the range of morphologies.
The extensive material examined (see
above for listing) has shown every
conceivable intermediate between those
few samples shown and even further
variation. The variability includes size of
stem at flowering (5–20 cm), proportion
of leaf length to leaf width (wider in
smaller specimen, more narrow in larger
specimens), leaf curvature (almost
straight to distinctly falcate), and angle of
spread of leaves (~10°–35°). No groupings
can be formed, and there are no trends
of covariation between vegetative and
floral morphology. The simplest answer
of a single species supported by a
polytomy in our molecular analysis is the
best explanation for all observations. We
acknowledge that significant intraspecific
variability in Orchidaceae goes against
commonly held views, yet the available,
multipronged, data-rich evidence clearly
supports our conclusion.
COMPARISON There are several
species with bifid epichile lobes bearing
points. We include select comparisons to
species with available material.
[4] Oberonia punctata habit (Andy’s Orchid
plant), portion of inflorescence (DLG 680/
HOAG 196) and SEM images of flowers
(E00233059).
Oberonia punctata J.J.Sm. (Fig. 4) is
overall smaller with more leaves of more
or less equal lengths along the stem; the
bract lacks hairs on the dorsal surface and
along the margin, the pedicelled ovary
and the back sides of the tepals lack hairs,
and the epichile lobes separate at an
acute angle from the mesochile.
Oberonia aporophylla Rchb.f. (=
Oberonia longirepens J.J.Wood; Fig. 5)
has much shorter leaves on the elongated
stems, forms creeping stolons, and has
white flowers.
Oberonia dubia J.J.Sm. (Fig. 6) has an
acaulescent, fan-shaped habit with much
more elongated leaves, auricles with
some projections, and more elaborately
fringed epichile lobes.
Oberonia disticha (Lam.) Schltr. (Fig. 7)
is restricted to the East African province,
is generally smaller, has more fleshy
leaves, lacks hairs on rachis and back of
tepals, and has an only slightly erose bract
without distinct hairs.
Lewis and Cribb (1989) suggested that
Oberonia kaniensis Schltr. [junior synonym
of Oberonia imbricata (Blume) Lindl.] may
be a synonym of O. equitans. The former
is in the section Labidous, characterized
by revolute sepals and a dual sac, while
the latter is in section Adenorachis with
spread sepals and a single sac. Accordingly,
that suggested synonymy is in error.
ECOLOGY The ecological data are
consistent with a broad synonymy of the
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taxa in question. The available elevation
data are shown in two groups, one for
the Western Pacific Islands records, the
traditional O. equitans, and one for the
Malayan region, traditionally referred
to as O. ciliolata. For the remaining
taxa, there are too little data available
to make meaningful frequency-based
assessments. Those minor species are
all known from the range of elevations
covered by O. equitans s.l. For instance,
O. palawanesis was described from 50 m
elevation (Schlechter 1921).
The frequency distributions are
remarkably similar. The observations near
sea level are somewhat less frequent
than at lower to intermediate elevations
(~1,000 m), after which they drop
rather steeply. There are fewer reports
and records of the species at higher
elevations in the Western Pacific. This can
be explained in part by the overall lower
elevations of the Western Pacific islands,
particularly French Polynesia.
The phenology data show the species
flowering throughout the year in both
broad areas of its distribution. Phenology
data in other species can show marked
seasonality (Geiger 2016, unpubl. data).
Oberonia equitans occurs on both sides
of the equator, which may obscure
seasonality patterns. However, seasonality
is less pronounced around the equator,
which lends credence to a true continuous
phenology pattern. With respect to the
Palau specimens (Crain 134, neotype Crain
198), they were found flowering in May
and October, respectively; Schlechter’s
type was in bud in March and most likely
would have flowered in April, the relative
peak flowering time of O. equitans in the
Malayan region.
Oberonia equitans has a distribution
range that includes Thailand, Singapore,
Malaysia, Indonesia, New Guinea,
Palau, New Caledonia, Vanuatu, Niue,
Wallis and Futuna, Tonga, Cook Islands,
Fiji, French Polynesia, and the Austral
Islands. Dispersal abilities seem to be
pronounced as evidenced by samples
from Palau, French Polynesia and Samoa
all being found in a major polytomy in our
molecular phylogeny. Oberonia has the
smallest seeds in the Orchidaceae (~100
μm; Barthlott et al. 2014; Geiger 2014,
unpubl. data), facilitating wind dispersal
and making microendemism less likely.
The seeds of O. equitans are slightly larger
at 150–200 μm, but still small enough to
permit wind dispersal over extensive
distances (Fig. 3).
DISCUSSION The recent work on the
alpha taxonomy of the overlooked genus
5
6
Oberonia (Bunpha et al. 2019; Geiger,
2016, 2019a, in press, unpubl. data) shows
that critical global revisions are needed to
clean up the accumulation of duplicate
names in minor orchid groups. If the scale
of overnaming in the genus Oberonia
alone is taken as an indicator for orchids
in general, then global orchid diversity
could be overestimated by around
one-third. The issue more likely affects
diminutive or cryptic species, however,
due to difficulties in field identification.
It is important to note that the issues
stem not only from poorly known older
names, but also from contemporary
authors that continue to introduce a
plethora of excess names. As a recent
example, Oberonia khuongii Aver. &
V.C.Nguyen in Averyanov et al. (2019) is
identical to Oberonia griffithiana Lindl.
(Geiger, in press), a fact communicated
to one of the coauthors well prior to
publication. They compared their O.
[5] Oberonia aporophylla habit (DLG
394/HOAG 49), portion of inflorescence
(DLG 599/HOAG 50) and SEM images
of flowers (DLG 394/HOAG 49, DLG
599/HOAG 50).
[6] Oberonia dubia habit (DLG 692/HOAG
272), portion of inflorescence (DLG 692/
HOAG 272) and SEM images of flowers
(DLG 692/HOAG 227).
khuongii only to O. cavaleriei, a very
different species with terete leaves, while
O. griffithiana was only mentioned by
name as another similar species without
any further discussion of characters.
A critical problem is that new species
are not compared to the most similar
species, but to seemingly random species
occurring in the same geographic area,
which is most likely due to the authors
not being sufficiently familiar with the
group in question. Formal synonymies for
recent names will be detailed elsewhere
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(Geiger, in press).
Another contributing factor, particularly in microfloral groups, is the
outdated reliance on drawings and
single-shot photographs to document
morphology. While in systematic zoology
(e.g., entomology, malacology), z-stack
photography and electron microscopy
have been considered standards for <5mm specimens for at least two to three
decades, those 21st century techniques
are only rarely employed in systematic
botany. Misleading drawings are a serious
problem as shown with the case of
Oberonia carprina Gilli, a synonym of O.
punctata J.J.Sm. (Geiger 2019a).
Orchid systematics is still too 18thcentury typological, where species are
considered immutable and minute
differences of individual specimens are
considered diagnostic at the species
level. Population thinking and explicit
consideration of intraspecific variability is
hardly taken into account when assessing
biodiversity. Bunpha et al. (2019) doubted
Geiger’s (2019a) synonymization of
Oberonia fungumolens Burkill under O.
padangensis Schltr. due to slight differences
in leaf shape and lip-to-sepal length ratio
of 1.4× vs. 1.6×, which Geiger (2019a)
was faulted for not explicitly addressing.
Differences in vegetative shape were
explicitly discussed under O. rufilabris
by Geiger (2019a) as a proxy for range of
variability in the genus overall. Variability
in vegetative parts of O. equitans is clearly
documented here and further supported
by examination of the cited herbarium
specimens. Proportional differences of
floral parts can easily reach one-quarter
to one-third among flowers on the same
inflorescence (e.g., Oberonia acaulis Griff.,
Oberonia ferruginea C.S.P.Parish ex Hook.
f., O. cavaleriei; Geiger, unpubl.), hence,
proportional differences of <15 percent
are clearly meaningless for taxonomy
in Oberonia. Accordingly, the crystalclear synonymy between O. padangensis
and O. fungumolens is maintained. The
complementary study of cultivated
specimens and wild-collected material
was noted by Geiger (2018), particularly
considering
intraspecific
variability
and genotype × environment effects.
Intraspecific variability in Oberonia
was discussed by Geiger (2019b). The
“annoying variability” of Oberonia spp.
had previously been noted by Comber
(1990, p. 149).
Last but not least, in today’s scientific
climate of measurements of academic
achievements, synonymizations are
possibly the least recognized activity
7
8
[7] Oberonia disticha habit (DLG 635/HOAG 14: yellow variant), portion of inflorescence
(DLG 424/HOAG 12: orange variant) and SEM images of flowers (DLG 424/HOAG 12).
[8] Elevation and phenology data for Oberonia equitans. The y-axis shows frequency of cited
occurrence. Elevation data for O. equitans from Western Pacific Islands from Cribb and
Whistler (1996), Hallé (1977), Lewis and Cribb (1989), Wood and Cribb (1994), Schlechter
(1906: as O. flexuosa), Fleischmann and Rechinger (1910), Brown (1931), Christophersen
(1935), Kores (1989), Streitmann (1983), Sykes (2016), and various herbarium records.
For Malayan region (as O. ciliolata) from O’Byrne (1994: as O. aurea, 2001), Beaman et
al. (2001), Comber (1990), Wood and Cribb (1994), Carr (1930, 1935: as O. lampongensis), Schuiteman and de Vogel (2006: as O. aurea), Schlechter (1911: as O. aurea, 1921:
as O. palawensis) and various herbarium records. Phenology data for O. equitans from
Western Pacific Islands from Hallé (1977), Anonymous (1995), Guillaumin (1957), Kränzlin (1898), Schlechter (1906, 1910), Ames (1933), Brown (1931), Christophersen (1935),
Yuncker (1959) and various herbarium records. For Malaysian region from Carr (1930,
1935: as O. lampongensis), Schuiteman and de Vogel (2006: as O. aurea), O’Byrne
(1994: as O. aurea), Schlechter (1911: as O. aurea, 1921: as O. palawensis), Schlechter in
Schumann and Lauterbach (1905: as O. aurea), and various herbarium records.
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geiger, et al.
in systematics. While the introduction
of a new name, including a synonym,
and even the transfer of a species to a
different genus, adds recognition, the
cleanup of excessive taxonomic splitting
garners far less recognition. No malus is
attached to introducing synonyms either.
Cleanup of superfluous names can have
further implications beyond the raw
tally of names. Recognizing Oberonia
attenuata Dockrill, considered endemic
to Queensland, Australia, and either
extinct or at least critically endangered,
as a southern range of the widespread
Oberonia insectifera Hook.f. changes
the outlook of the species’ survival and
conservation needs (Geiger 2019a).
A significant number of clear
synonymies remain to be formally
addressed in Oberonia. Furthermore,
there are several names that are extremely
difficult to assess due to limited verbal
descriptions, lack of any illustrations,
and unknown (lost or destroyed?) types.
Examples include Oberonia tahitensis
Lind., Oberonia werneri Schltr., and
Oberonia zimmermanniana J.J.Sm. Those
nomina dubia are still listed as correct
names in databases and inventories.
Accordingly, the cleanup and revision of
the genus will continue for several more
years to help alleviate many of the issues
discussed above.
Acknowledgments
Collection visits were facilitated by the
respective staff: B (Robert Vogt and Nils
Köster), BM (Ranee Prakash), CANB (Mark
Clements, Brendan Lepschi), E (Hanna
Atkins, Suzanne Cubey), F (Christine
Niezgoda), K (André Schuiteman), MICH
(Brad Ruhfle), MEL (Wayne Gerber), MO
(James Solomon and Donna Herrera), P
(Marc Pignal), SING (Balizah Mohd Ibrahim,
Hubert Kurzweil, Serene Lee, Paul Leong),
W (Ernst Vitek), WU (Walter Till), US
(Rusty Russell). André Schuiteman (Kew
Botanical Gardens), Barbara Gravendeel
(Naturalis, Leiden) and Sean Lahmeyer
(Huntington Botanical Gardens) facilitated
loans of spirit material. Fieldwork in Palau
was supported by a 2019 Research Award
of the American Orchid Society to B.
Crain, a Scholarly Studies Grant through
the Smithsonian Institution, and support
from the US Forest Service Institute
of Pacific Islands Forestry. Laboratory
and herbarium work was supported
by the Smithsonian Environmental
Research Center and National Museum
of Natural History. Fieldwork in Palau
was also supported by multiple state and
federal government agencies, the Belau
National Museum, staff at the Ngardok
Nature Reserve, the Coral Reef Research
Foundation, and an exceptional group
of citizen scientists including R. Leidich,
G. McKlinay, and J. and L. Miles. Amy
Weiss (NYBG) and Kenneth M. Cameron
(WIS) provided additional information.
The reviewers and editor Ron McHatton
helped to improve the manuscript.
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— Daniel L. Geiger is Curator of
Malacology at the Santa Barbara Museum
of Natural History, 2559 Puesta del Sol,
Santa Barbara, California 93105. (email
geiger@vetigastropoda.com, dgeiger@
sbnature2.org). He is also a Visiting
Research Scholar, Huntington Library, Art,
Museum and Botanical Gardens — Botany
Division, 1151 Oxford Street, San Marino,
California 91108. Benjamin J. Crain
(email bcrainium@yahoo.com), Melissa
K. McCormick (email mccormickm@
si.edu) and Dennis F. Whigham (email
whighamd@si.edu) are ecologists with
the North American Orchid Conservation
Center,
Smithsonian
Environmental
Research Center, 647 Contees Wharf
Road, Edgewater, Maryland 21037.
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