ORCHIDS www.aos.org THE BULLETIN OF THE AMERICAN ORCHID SOCIETY VOL. 89 NO. 8 AUGUST 2020 Prepared for download exclusively for Benjamin Crain ORCHIDS CONTENTS August 2020 Volume 89 Number 8 The Bulletin of the American Orchid Society RON MCHATTON Chief Education and Science Officer Editor, Orchids Magazine rmchatton@aos.org AWARDS REGISTRAR Laura Newton laura@aos.org ADVERTISING Kevin Hall Advertising Sales Executive Allen Press 810 East 10th Street Lawrence, Kansas 66044 khall@allenpress.com 785-865-9143 SUBSCRIPTIONS AND MISSING ISSUES Membership Services Department Tel 305-740-2010 Fax 305-747-7154 membership@aos.org EDITORIAL BOARD Jean Allen-Ikeson, Chair Greg Allikas, Sue Bottom, Mark Chase, Phillip Cribb, Nile Dusdieker, Wes Higgins, Carol Klonowski, Judith Rapacz-Hasler, Larry Sexton Send electronic submissions to jean.ikeson@gmail.com or rmchatton@aos.org PROOFREADERS Laura Newton, Larry Sexton, Olga Skoropad, Susan Wedegaertner FORMER EDITORS Dr. David Lumsden (1932–1940), Dr. Louis O. Williams (1940–1943), Gordon Dillon (1943–1967; 1970–1973), Merle Reinikka (1968–1969), Richard Peterson (1973–1984), Stephen R. Batchelor (1984), Alec Pridgeon, PhD (1984–1988; 1989–1991), Chuck McCartney (1988–1989), James B. Watson (1991–2013) Volume 89, Number 8 August 2020 Orchids (ISSN 1087-1950) is published monthly by the American Orchid Society, Inc., at Fairchild Tropical Botanic Garden Editorial Office: 10901 Old Cutler Road, Coral Gables, Florida 33156 (telephone 305-740-2010; fax 305-7477154; email theaos@aos.org; website www.aos.org). ©American Orchid Society, Inc. 2017. Printed by Allen Press, 810 East 10th Street, Lawrence, Kansas 66044. Subscription price of Orchids is $79 a year within the US, $99 Canada and Mexico and $119 for all other countries. Single copies of current issue cost $8.50 (plus shipping and handling). Prices are subject to change without notice. Although Orchids endeavors to assure the reliability of its advertising, neither Orchids nor the American Orchid Society, Inc. can assume responsibility for any transactions between our advertisers and our readers. Periodical postage paid at Miami, FL and additional offices. POSTMASTER: Send address changes to: Orchids, PO Box 565477, Miami, FL 33256. The American Orchid Society follows the World Checklist of Selected Plant Families with regard to questions of botanical nomenclature and synonymy in orchid species names and the International Orchid Register for hybrid nomenclature and parentage in editorial. The opinions and recommendations that appear in Orchids regarding the selection and use of specific plant-care products, including but not limited to pesticides, fungicides and herbicides, are those of the individual authors, and not those of the American Orchid Society, which neither adopts nor endorses such opinions and recommendations and disclaims all responsibility for them. When selecting and using such products, readers should seek and obtain the advice of the manufacturer and of responsible government agencies. Mail date: July 27, 2020. Printed on 10 percent post-consumer recycled paper. 602 618 628 636 FEATURES 618 Rhyncholaelia digbyana The Frilled Wonder Stephen van Kampen-Lewis 626 orchids in watercolor Isotria verticillata Marcia Whitmore 628 In pursuit of teagueia Fieldwork in the Most Mysterious and Dangerous Mountains of Ecuador Kelsey L. Huisman 636 who were these guys: part 11 Hugh Low (1824–1905) David Rosenfeld, MD DEPARTMENTS In This Issue Tom’s Monthly Checklist 600 August: The Month of the Fireflies Thomas Mirenda AOS DIRECTORY OF SERVICES 594 AOS MEMBERSHIP INFORMATION 594 pronunciation guide  595 Collector’s Item 602 Bulbophyllum maxillare Charles Wilson AOS NATIONAL VOLUNTEERS 596 gifts of note  597 New Rufugium Botanicum 606 Coelogyne lawrenceanum Melania Fernández and Franco Pupulin Watercolor by Sylvia Strigari Conservation Committee Cypripedium guttatum Thomas Mirenda president’s message  598 webinars  605 Useful tips Selected botanical terms  609 610 The 2020 Phillip E. Keenan Awards Thomas Mirenda Calendar  667 612 Species Identification Task Force 613 Cattleya bradei ‘Elizabeth Anne’ CHM/AOS Ron McHatton Orchids Illustrated 614 de Vriese Peggy Alrich and Wesley Higgins Awards Gallery 605, 668, 670 ORCHId mARKETPLACE 669 ORCHIDS CLASSIFiEDs  669 Ad Index 671 Parting Shot  672 Epiphytic Orchids North of the 49th Parallel Sasha Kubicek 640 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. SUBSCRIBE TO ORCHIDS TEL 305-740-2010 EMAIL THEAOS@AOS.ORG WEBSITE WWW.AOS.ORG Prepared for download exclusively for Benjamin Crain 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 656 Orchids august 2020 © American Orchid Society   www.AOS.org Prepared for download exclusively for Benjamin Crain geiger, et al. 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), www.AOS.org  august 2020 © American Orchid Society Orchids 657 Prepared for download exclusively for Benjamin Crain 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, 2,300 658 Orchids august 2020 © American Orchid Society   www.AOS.org Prepared for download exclusively for Benjamin Crain geiger, et al. 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 Prepared for download exclusively for Benjamin Crain geiger, et al. 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 660 Orchids august 2020 © American Orchid Society   www.AOS.org Prepared for download exclusively for Benjamin Crain geiger, et al. 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, www.AOS.org  august 2020 © American Orchid Society Orchids 661 Prepared for download exclusively for Benjamin Crain geiger, et al. 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. 662 Orchids august 2020 © American Orchid Society   www.AOS.org Prepared for download exclusively for Benjamin Crain geiger, et al. 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 www.AOS.org  august 2020 © American Orchid Society Orchids 663 Prepared for download exclusively for Benjamin Crain geiger, et al. 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 664 Orchids august 2020 © American Orchid Society   www.AOS.org Prepared for download exclusively for Benjamin Crain geiger, et al. (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. www.AOS.org  august 2020 © American Orchid Society Orchids 665 Prepared for download exclusively for Benjamin Crain 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. References Ames, O. 1907. Orchidaceae Halconenses: An Enumeration of the Orchids Collected on or Near Mount Halcon, Mondoro, Chiefly by Elmer D. Merrill. 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Plants of Tonga. Bernice P. Bishop Museum Bulletin 220:1–283. — 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. DUE TO CORONAVIRUS CONCERNS, AOS JUDGING WAS SUSPENDED AND SHOWS CANCELED OR POSTPONED. 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