Botanical Journal of the Linnean Society, 2010, 163, 234–278. With 5 figures Variation in chromosome number and the basic number of subfamily Epidendroideae (Orchidaceae) boj_1059 234..278 LEONARDO P. FELIX1* and MARCELO GUERRA2 Departamento de Fitotecnia, Universidade Federal da Paraíba, Campus II, Areia, PB 58.397-000, Brazil 2 Departamento de Botânica, Universidade Federal de Pernambuco, Recife, PE 50.670-420, Brazil Received 10 November 2009; revised 2 March 2010; accepted for publication 12 May 2010 The chromosome numbers of 21 genera and 44 species of subfamily Epidendroideae belonging to tribes Sobralieae, Epidendreae, Malaxideae and Vandeae, and subtribe Dendrobiinae, were determined. Chromosome numbers varied from 2n = 24 in Malaxis pubescens to 2n = c. 240 in Epidendrum cinnabarinum. A revision of the chromosome numbers known for the subfamily was also performed, aimed at determining the basic numbers of the genera, subtribes and tribes. The first counts for 31 species and six genera of tribe Sobralieae and subtribe Ponerinae are presented. The basic number for each genus was evaluated. A predominance of x = 20 in genera of Epidendreae and Arethuseae, x = 19 in Vandeae and subtribe Dendrobiinae and x = 15 and 21 in Malaxideae was observed. Other tribes were more variable. A wide occurrence of x = 19 and 20 in Epidendroideae and of x = 21 in at least one genus of all tribes suggests that disploidy of one or a few chromosomes has played a decisive role in the establishment of the basic karyotypes. The karyotype variability observed in the subfamily is discussed in light of current phylogenetic proposals for the family. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278. ADDITIONAL KEYWORDS: basic chromosome number – cytogenetics – disploidy – polyploidy. INTRODUCTION The analysis of chromosome numbers has been used to evaluate evolutionary and taxonomic relationships in diverse groups of plants. Studies of chromosomal variations in Orchidaceae have contributed to the better understanding of the taxonomy of this family at many hierarchical levels. Arditti (1992) reviewed some of the important aspects of the cytology of the orchid family, with emphasis on chromosomal variation. Brandham (1999) reviewed chromosomal variation in each genus of Apostasioideae Garay and Cypripedioideae Garay, but did not consider the probable basic numbers of higher taxonomic categories. Felix & Guerra (2000, 2005) examined the chromosomal variability in the cymbidioid clade (sensu Dressler, 1993) and in Cypripedioideae, Orchidoideae (including Spiranthoideae Dressler) and Vanilloideae Szlach. and *Corresponding author. E-mail: lpfelix@hotmail.com 234 concluded that x = 7 is the most probable basic number for the family Orchidaceae. The basic number corresponds to the haploid number encountered in a given taxon that explains in the most parsimonious manner the variation in chromosome numbers seen in that and related taxa (Guerra, 2000). According to Chase, Freudenstein & Cameron (2003), orchids are subdivided into five subfamilies: Apostasioideae, Cypripedioideae, Vanilloideae, Orchidoideae and Epidendroideae Lindl. Epidendroideae comprise an estimated 18 000 species and 650 genera (Cribb & Chase, 2005), mainly epiphytic plants, distributed in tropical and subtropical regions throughout the world. Epidendroideae are highly diversified and no morphological synapomorphy is shared by all members of the group. They are subdivided into 16 tribes, of which Epidendreae Kunth, Cymbidieae Pfitzer, Dendrobieae Lindl. and Vandeae Lindl. account for approximately 15 000 species (Chase et al., 2003). © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 1 CHROMOSOMAL VARIABILITY OF EPIDENDROIDEAE MATERIAL AND METHODS The large majority of the plants analysed here were collected in north-eastern Brazil. All of the material studied was cultivated at the Federal Rural University of Pernambuco, the Federal University of Paraíba or the Federal University of Pernambuco, except for some plants donated by private collectors. The species analysed, chromosome numbers counted here and previous counts are listed in Table 1. Samples of all materials were deposited in the PEUFR, HST and EAN herbaria. Identifications were based on Pabst & Dungs (1975, 1977), but the binomials were updated according to Govaerts et al. (2009; http:// www.kew.org/wcsp/monocots/). Chromosome analyses were undertaken using root tips pretreated with 0.002 M 8-hydroxyquinoline at 4 °C for 24 h. The material was then fixed in absolute ethanol/glacial acetic acid (3 : 1 v/v) for 3–24 h at room temperature 25 °C and stored at -20 °C. To prepare slides, root types were hydrolysed in 5 M HC1 at room temperature, frozen in liquid nitrogen to remove the coverslip and stained with 1% acetic haematoxylin (Guerra, 1999) or 2% Giemsa (Guerra, 1983). The diversity in chromosome numbers in the epidendroid clade was examined by undertaking a revision based on the chromosome numbers compiled by Tanaka & Kamemoto (1984) and the indexes prepared by Fedorov (1969), Moore (1973, 1974, 1977), Goldblatt (1981, 1984, 1985, 1988), Goldblatt & Johnson (1990, 1991, 1994, 1996, 2000, 2003, 2006) and more recent papers. A separate list of chromosome numbers was then prepared, including the data from the present analysis (see also Appendix 1). In this table, the names of the genera, species and authors are cited as in the World Checklist of Monocotyledons (Govaerts et al., 2009) and previous names indicated in parentheses. The data from Appendix 1 were synthesized in Table 2, showing the chromosome number variation in each genus. The numbers were ordered from the most to the least frequent, and the probable basic number for each genus was underlined, based on the frequency of each number in the genus and in the closest related genera. The chromosome numbers that were considered questionable (as they differed significantly from information available in the literature), and the occasional case of monosomy and trisomy were excluded from Appendix 1 and Table 2, and from the discussion, and are presented separately in Appendix 2. RESULTS The interphase nucleus structure and the chromosome number of a total of 44 species belonging to 21 genera, seven subtribes and four tribes of Epidendroideae (sensu Chase et al., 2003) were analysed. The structure of the interphase nuclei was quite variable among the species analysed. According to the classification of interphase nuclei suggested by Tanaka (1971) for Orchidaceae, the structural types varied from diffuse nuclei in Campylocentrum pernambucense Hoehne (Fig. 1B) and Cattleya walkeriana Gardner to complex chromocentric nuclei in Prosthechea caetensis (Bicalho) W.E.Higgins and Malaxis excavata Kuntze (Fig. 1C). The species analysed here generally had small chromosomes, varying slightly in length, all species displaying meta- to submetacentric chromosomes and © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 Cymbidieae have corms or pseudobulbs and two to four pollinia bodies (Chase et al., 2003). The chromosome numbers in this tribe are quite variable, ranging from n = 6 in Erycina pusilla (L.) N.H.Williams & M.W.Chase (as Psygmorchis pusilla (L.) Dodson & Dressler) to n = 84 in two species of Oncidium Sw. (revised by Felix & Guerra, 2000). Epidendreae are subdivided into two main subtribes: Pleurothallidinae Lindl. and Laeliinae Benth. The former has 28 genera and approximately 4000 species, but chromosome numbers are available only for four genera and 36 species, with a predominance of n = 20 in Pleurothallis R.Br. s.l. and n = 16 in Stelis. Chromosome numbers are known for 14 out of 45 genera and 94 of 1788 species of Laeliinae, with a predominance of n = 20 in the majority of the genera (Tanaka & Kamemoto, 1984). Vandeae are comprised of two monophyletic subtribes having monopodial growth (Aeridinae Pfitzer with x = 19 and Angraecinae Summerh. with x = 19, 20, 21, 23, 24 and 25) (Carlsward et al., 2006) and subtribe Polystachiinae Schltr. which has sympodial growth and x = 20 (Jones, 1966). Subtribes Agrostophyllinae Szlach., Dendrobiinae Lindl. and Collabiinae Pfitzer do not have welldefined positions in Epidendroideae (Chase et al., 2003). Although the molecular phylogenetics of Epidendroideae has been relatively well studied, it has never been cytotaxonomically evaluated. In the present study, we analysed 44 Brazilian species of Epidendroideae (sensu Chase et al., 2003) and critically reviewed the chromosome numbers for each genus and subtribe in order to establish the probable basic numbers of the subfamily. These data were compared with the classification system developed by Dressler (1993) and Chase et al. (2003), and with recent phylogenetic proposals by Pridgeon, Solano & Chase (2001), Freudenstein et al. (2004), Carlsward et al. (2006), Cameron (2005), van den Berg & Chase (2004a), van den Berg et al. (2005, 2009), among others. 235 236 L. P. FELIX and M. GUERRA Table 1. List of species analysed with respective provenances, voucher numbers, chromosome numbers (n and/or 2n), figures in the text, herbarium where each material was deposited, previous counts and sources Taxa Subfamily Epidendroideae Tribe Sobralieae Elleanthus brasiliensis (Lindl.) Rchb.f. Sobralia liliastrum Lindl. B. nodosa (L.) Lindl. Cattleya amethystoglossa Linden & Rchb.f. ex R.Warner C. elongata Barb. Rodr. C. granulosa Lindl. C. nobilior Rchb.f. var. amaliae Pabst C. nobilior var. nobilior C. walkeriana Gardner C. walkeriana var. princeps L.C.Menezes Cattleyella araguaiensis (Pabst) Van den Berg & M.W.Chase Dimerandra emarginata (G.Mey.) Hoehne Encyclia advena (Rchb.f.) Porto & Brade E. flava (Lindl.) Porto & Brade E. oncidioides (Lindl.) Schltr. Subtribe Laeliinae (cont.) Epidendrum avicule Lindl. E. cinnabarinum Salzm. ex Lindl. E. E. E. E. E. difforme Jacq. ellipticum Graham latilabrum Lindl. nocturnum Jacq. rigidum Jacq. Jacquiniella globosa (Jacq.) Schltr. Laelia marginata (Rchb.f.) L.O.Williams Prosthechea caetensis (Bicalho) Pabst P. fragrans (Sw.) W.E.Higgins P. vespa (Vell.) W.E.Higgins Scaphyglottis fusiformis (Griseb.) R.E.Schult. Sophronitis lobata (Lindl.) Van den Berg S. purpurata (Lindl. & Paxton) Van den Berg & M.W.Chase Subtribe Pleurothallidinae Acianthera ochreata (Lindl.) Pridgeon & M.W.Chase Stelis sp. Tribe Malaxideae Malaxis excavata (Lindl.) Kuntze M. pubescens (Lindl.) Kuntze Tribe Vandeae Subtribe Polystachiinae Polystachya estrellensis Rchb.f. Subtribe Angraecinae Campylocentrum amazonicum Cogn. C. crassirhizum Hoehne C. pernambucense Hoehne Unplaced subtribes within Epidendroideae Subtribe Dendrobiinae Bulbophyllum cribbianum Toscano B. sanderianum Rolfe Bulbophyllum sp. 1 Bulbophyllum sp. 2 Voucher no. 2n/n Herbarium Figs Maranguape, CE Camocim do São Felix, PE Palmeiras, BA Morro do Chapéu, BA L.P.Felix, L.P.Felix, L.P.Felix, L.P.Felix, 50 48 48 48 PEUFR PEUFR PEUFR EAN 1A, 2A 2B 2C Taquaritinga do Norte, PE L.P.Felix, 12715 40 EAN 2D 8284 8213 9124 7394 Previous counts (2n) Sources† 40 DA09 40 TK84 40 TK84 40 40 GJ90 TK84 Gilbués, PI L.P.Felix, S/N 40 HST 2E Camocim do São Felix, PE Puxinanã, PB Cultivated L.P.Felix, 9650 L.P.Felix, 12728 L.P.Felix, 12725 40 40 40 PEUFR EAN EAN 2F 2G Morro do Chapéu, BA Morro do Chapéu, BA Natal, RN Alcaçuz, RN Cultivated Cultivated, Ostetto Orquídeas Cultivated, St Cruz 3263 Cultivated, St Cruz 3149 L.P.Felix, 9651 L.P.Felix, 12976 L.P.Felix, 12977 L.P.Felix, 12978 Unvouchered Unvouchered Unvouchered Unvouchered 40 80 40 40 42 42 40 c. 80 PEUFR EAN EAN EAN – – – – Rio Araguaia, TO Carmópolis, SE Bonito, PE Brejo da Madre de Deus, PE Araguaina, TO Brejo da Madre de Deus, PE Esperança, PB L.P.Felix, 9360 L.P.Felix, 9652 L.P.Felix, 9638 L.P.Felix, 12725 LPFelix, 12723 L.P.Felix, 12727 L.P.Felix, 12731 40 40 40 40 40 40 40 EAN PEUFR HST EAN EAN EAN EAN 1E, 2M 2N 3A Brejo da Madre de Deus, PE Camocim do São Felix, PE Mamanguape, PB Serraria, PE Esperança, PB Cabo, PE Camocim do São Felix, PE Taquaritinga do Norte, PE Belém, PA Brejo da Madre de Deus, PE Bonito, PE Taquaritinga do Norte, PE Bezerros, PE Morro do Chapéu, BA Brejo da Madre de Deus, PE Alcaçuz, RN São Felix do Xingu, PA Bezerros, PE Cultivated L.P.Felix, L.P.Felix, L.P.Felix, L.P.Felix, L.P.Felix, L.P.Felix, L.P.Felix, L.P.Felix, L.P.Felix, 40 c. 240 c. 240 n = 108–124 c. 240 40 68 40 80 EAN PEUFR EAN EAN EAN PEUFR PEUFR EAN EAN 3G 3D 40 c. 240 TK84 G00, OL06 3F 3E 3H 39–40 – – 40, 80 – TK84 L.P.Felix, 12735 L.P.Felix, 8385 L.P.Felix, 12721 L.P.Felix, 1038 L.P.Felix, 8644 L.P.Felix, 12733 L.P.Felix, 12732 L.P.Felix, 12979 L.P.Felix, 9640 Unvouchered 40 38 38 40 80 40 40 40 40 40 EAN PEUFR EAN EAN PEUFR EAN EAN EAN PEUFR 3I 3J – 3K 4A 4B 40 – 40 4C 4D – – – – Cultivated Unvouchered 40 – 40 Bezerros, PE Ibateguara, AL L.P.Felix, 9641 L.P.Felix, 8271 40 32 PEUFR EAN 4E 4F – – Bonito, PE Rio Grande, RS L.P.Felix, 8464 L.P.Felix, 9022 30 24 PEUFR PEUFR 4G 4H – – Itapororoca, PB L.P.Felix, 798 80 EAN – 80 Acará, PA Bezerros, PE São Vicente Ferrer, PE Areia, PB L.P.Felix, L.P.Felix, L.P.Felix, L.P.Felix, 8490 8919 S/N 12982 80 38 38 EAN PEUFR EAN EAN 4I 4J 4K – – – Rio de Contas, BA Brejo da Madre de Deus, PE Campo Formoso, BA Morro do Chapéu, BA L.P.Felix, L.P.Felix, L.P.Felix, L.P.Felix, 8831 12981 11802 11701 38 38 38 80 PEUFR EAN EAN EAN 4L 4M 4N 4O – – – – 8355 9653 2165 11489 12989 8324 9435 12736 12667 2H 2I 2J 2K 2L 3B 3C TK84 TK84 TK84 M77 TK84 *Brazilian state abbreviations: AL, Alagoas; BA, Bahia; CE, Ceará; Pa, Pará; PB, Paraíba; PE, Pernambuco; PI, Piauí; RN, Rio Grande do Norte; RS, Rio Grande do Sul; SE, Sergipe; TO, Tocantins. †Sources: TK84, Tanaka & Kamemoto (1984); GJ90, Goldblatt & Johnson (1990); G00, Guerra (2000); M77, Moore (1977); OL06, Conceição et al. (2006); DA09, Daviña et al. (2009). © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 S. sessilis Lindl. Tribe Epidendreae Subtribe Ponerinae Isochilus linearis (Jacq.) R.Br. Subtribe Bletiinae Bletia catenulata Ruiz & Pav. Subtribe Laeliinae Brassavola tuberculata Hook. Provenance* CHROMOSOMAL VARIABILITY OF EPIDENDROIDEAE 237 Table 2. Chromosome numbers and probable base numbers (underlined) of tribes, subtribes and genera of subfamily Epidendroideae (sensu Chase et al., 2003), except those of the cymbidioid phylad previously published by Felix & Guerra (2000), including the number of genera and species in brackets Subfamilies, tribes and subtribes, number of genera/species cytologically known and probable base number Genera with the number of species known/analysed Chromosome numbers*† and most probable base numbers (underlined) Aphyllorchis Blume (15/1) Cephalanthera L.C.Rich. (14/10) Epipactis Sw. (21/19) Limodorum L. (1/1) Neottia L. (Listera included) (63/30) c. 18 16–18, 17, 22 20, 18–30, 16–19 28–32 18, 19–20–21, 17, 14–28–23 Elleanthus Presl. (106/1) Sobralia Ruiz & Pavon (120/2) 25 24 Gastrodia R.Br.(41/4) 20, 19 Calypso Salisb. (1/1) 28 Isochilus R.Br. (12/1) 20 Bletia Ruiz & Pavon (30/2) 20, 30 Acianthera Scheidw. (131/7) Anathallis Barb. Rodr. (89/1) Masdevallia Ruiz & Pavon (380/2) Pabstiella Brieger & Senghas (8/1) Pleurothallis R.Br. (1120/10) Scaphosepalum Pfitzer (41/1) Specklinia Lindl. (90/1) Stelis Sw. (370/7) 20 21 18–22 16 19–20, 18, 34–36–42 32 10 16–19, 17–21–32–38–39 Brassavola R.Br. (20/4) Broughtonia R.Br.(6/1) Cattleya Lindl. (54/26) Catleyella Van den Berg & Chase (1/1) Caularthron Rafin. (4/1) Dimerandra Schltr. (6/2) Encyclia Hook. (154/7) Epidendrum L. (1125/38) Guarianthe Dressler & W.E.Higgins (5/1) Jacquiniella Schltr. (6/1) Laelia Lindl. (s.l.) (11/9) Leptotes Lindl. (6/1) Prosthechea (93/14) Psychilis Raf. (17/1) Rhyncholaelia Schltr. (2/1) Scaphyglottis Lindl. (63/4) Sophronitis Lindl. (57/19) 20 20 20, 40, 21–27–30 20 20 20 20 20, 14–40, 30–34, 12–15–19–26–35–60, 21–24–45–80–(108–124)–120 20 19 20, 21, 22–30 20 20, 28–40 20 20 20, 19 20, 40, 60 Ceratostylis Blume (145/1) Cryptochilus Wallich (4/2) Eria Lindl. (404/48) Mediocalcar J.J.Sm. (24/4) Porpax Lindl. (13/3) Pseuderia Schltr. (19/1) Trichotosia Blume (73/2) 20 19 19, 20, 18–22, 33–38 19 12, 21 20 19–22 Podochilus Blume (60/1) 19 Phreatia Lindl. (201/1) Thelasis Blume (23/1) 16 16 Anthogonium Lindl. (1/2) Arethusa L. (1/1) Arundina Blume (1/1) Calopogon R.Br. (5/5) Eleorchis Maek. (2/1) 20–19–21 20 20, 16–19 21, 13 20 TRIBE NEOTIEAE (6/191) TRIBE GASTRODIEAE (6/70) TRIBE CALYPSOE (13/70) TRIBE EPIDENDREAE (86/5870) Subtribe Ponerinae (22/3) Subtribe Bletiinae (3/48) Subtribe Pleurothallidinae (28/3021) x = 20 Subtribe Laeliinae (van den Berg et al., 2000) (45/1788) x = 20 TRIBE PODOCHILEAE (21/1232) Subtribe Eriinae (11/725) Subtribe Podochilinae (4/208) Subtribe Thelasiinae (6/299) x = 16 TRIBE ARETHUSEAE (24/701) Subtribe Arethusinae (2/3) x = 20 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 TRIBE SOBRALIEAE (4/237) Subtribe Sobraliinae (4/237) 238 L. P. FELIX and M. GUERRA Table 2. Continued Subfamilies, tribes and subtribes, number of genera/species cytologically known and probable base number Subtribe Coelogyninae (20/274) x = 20 TRIBE VANDEAE (157/1960) Subtribe Polystachyinae (4/228) x = 20 Subtribe Aeridinae (98/1352) x = 19 Chromosome numbers*† and most probable base numbers (underlined) Bletilla Rchb.f. (9/3) Coelogyne Lindl. (100/19) Dendrochilum Blume (100/2) Neogyna Rchb. f. (1/1) Otochilus Lindl. (4/3) Panisea Steudel (7/3) Pholidota Hook. (30/11) Pleione D.Don (15/11) Thunia Rchb.f. (5/1) 16, 32–38 20, 19, 21–40 15–19 21 20 20 20, 19, 21–22 20, 19–40 20, 19 Liparis Rich. (418/52) Malaxis Sw. (395/13) Oberonia Lindl. (308/30) 21, 15, 19, 10–20, 14–18–40, 11–13–28–34–38 21, 15–30, 14, 18–c. 20–22 15, 30 Polystachya Hook. (150/42) 20, 40, c. 60 Acampe Lindl. (7/4) Aerides Lour. (25/10) Arachnis Blume (11/1) Ascocentrum Schltr. (13/5) Biermannia King & Pantl. (10/1) Calymmanthera Schltr. (5/2) Chilochista Lindl. (20/3) Cleisostoma D.Don (87/17) Cleisostomopsis Seidenf. (2/1) Cottonia Wight (1/1) Diplocentrum Lindl. (2/2) Diplopora Hook. (2/1) Drymoanthus Nicholls (4/3) Esmeralda Rchb.f. (2/1) Euanthe Schltr. (1/1) Gastrochilus D.Don (55/9) Gunnarella Senghas (9/1) Holcoglossum Schltr. (10/2) Hygrochilus Pfitzer (1/1) Luisia Gaudich. (39/9) Micropera Lindl. (18/2) Neofinetia Hu (2/1) Ornitochilus Lindl. (3/1) Papilionanthe Schltr. (10/6) Paraphalaenopsis Hawkes (4/2) Pelatantheria Ridl. (7/2) Phalaenopsis Blume (62/23) Pomatocalpa Breda (35/3) Pteroceras Hasselt ex Hassk. (24/2) Renanthera Lour. (33/7) Rhynchostylis Blume (3/3) Robiquetia Gaudich. (38/5) Sarcochilus R.Br. (25/2) Sedirea Garay & H.R.Sweet (2/1) Schoenorchis Blume (26/2) Seidenfadenia Garay (1/1) Smitinandia Holttum (3/2) Smithsonia C.J.Saldanha (3/2) Staurochilus Ridl. (14/1) Stereochilus Lindl. (7/1) Taeniophyllum Blume (185/5) Thrixspermum Lour. (144/6) Trichoglottis Blume (64/4) Tuberolabium Yamam. (12/2) Uncifera Lindl. (6/3) Vanda Jones (57/21) Vandopsis Pfitz. (5/3) 19, 18 19, 38 19 19 19 19 19 19, 20 19–20 19–20 19 8 19, 38 21 19 19, 17–20 19 19 19 19, 20, 18 19 19 19 19, 20–38 19 19 19, 38 19 19 19 19 19 19–38 20 19 19 19 19 19 19 19–20, 28 19 19 15–19 19 19, 38, 57 19 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 TRIBE MALAXIDEAE (7/1158) x = 15, 21 Genera with the number of species known/analysed CHROMOSOMAL VARIABILITY OF EPIDENDROIDEAE 239 Table 2. Continued Subfamilies, tribes and subtribes, number of genera/species cytologically known and probable base number Subtribe Angraecinae (18/445) x = 19 UNPLACED SUBTRIBES Subtribe Agrostophyllinae (8/196) x = 20 Subtribe Dendrobiinae (17/3332) x = 19 Subtribe Collabiinae (19/435) x = 20 Chromosome numbers*† and most probable base numbers (underlined) Angraecum Bory (219/28) Calyptrochilum Kraenzl. (2/2) Campylocentrum Benth. (73/4) Cryptopus Lindl. (4/1) Dendrophylax Rchb.f. (9/1) Jumellea Schltr. (58/1) Listrostachys Rchb.f. (2/1) Oeoniella Schltr. (2/1) 19, 25, 21, 20–23–24, c. 38–46 19 19–22–40 38 44 19–22 23 19 Aerangis Rchb.f. (490/15) Ancistrorhynchus Finet (16/9) Angraecopsis Kraenzl. (21/5) Bolusiella Schltr. (6/1) Chamaeangis Schltr. (10/2) Cribbia Senghas (4/1) Cyrtochis Schltr. (15/9) Diaphananthe Schltr. (24/5) Eggelingia Summ. (3/1) Eurychone Schltr. (2/1) Microcoelia Lindl. (29/14) Microterangis Senghas (7/1) Mystacidium Lindl. (9/1) Podangis Schltr. (1/1) Rangearis Summ. (7/3) Rhipidoglossum Schltr. (37/6) Solenangis Schltr. (6/2) Sphyrarhynchus Mansf. (1/1) Tridactyle Schltr. (43/3) 25, 21, 23–26–27–100 24, c. 25–36–48 25, 24 25 25–50 25 23, 25–46–69–75 25 23 25 24 25 24 23 23–46–50–54 25, 50 25 25 50, 25 Agrostophyllum Blume (91/5) Earina Lindl. (6/3) Glossorhyncha Ridl. (80/2) 20, 19 20 20 Bulbophyllum Thouars (1784/166) Dendrobium Sw. (1184/251) Diplocaulobium Kraenzl. (99/5) Epigeneium Gagnep. (38/5) Flickingeria A.D.Hawkes (69/3) Geniorchis Schltr. (7/1) Saccoglossum Schltr. (5/1) Trias Lindl. (12/1) 19, 20, 18, 21, 10–40 19, 20–18, 38, 10–21–22 19 20 19 19 c. 20 19 Acanthephippium Blume (12/6) Calanthe R.Br. (187/48) Cephalantheropsis Guillaumin (5/2) Chrysoglossum Blume (6/1) Gastrorchis Thouars (8/4) Nephelaphyllum Blume (12/1) Pachystoma Blume (1/1) Phaius Lour. (48/11) Plocoglottis Blume (39/2) Spathoglottis Blume (45/8) Tainia Blume (29/9) 24, 20, 20, 18 20 18 20 21, 19 20, 20, 21–23 21, 19–22, 23, 10–30 21 22, 14–19–23–24 18–19 15–16–18–36–38 *Chromosome numbers are ordered from the more to the less frequent. †Numbers connected with a rule (–) have equal frequencies. symmetrical karyotypes. Chromosome numbers varied from 2n = 24 in Malaxis pubescens Kuntze to 2n = c. 240 in Epidendrum cinnabarinum Salzm. ex Lindl. In tribe Sobralieae Pfitzer, subtribe Sobraliinae Pfitzer, Elleanthus brasiliensis Rchb.f. had 2n = 50 (Fig. 2A), whereas Sobralia liliastrum Lindl. and S. sessilis Lindl. had 2n = 48 (Fig. 2A–C). The proportion of condensed chromatin per chromosome was more variable in the most asymmetric karyotypes, generally being greater in the largest chromosome pairs, as for example in E. brasiliensis (Fig. 1A). In tribe Epidendreae, Isochilus linearis (Jacq.) R.Br. (Fig. 2D) and Bletia catenulata Ruiz & Pav. (Fig. 2E), from subtribes Ponerinae and Bletiinae had 2n = 40, whereas subtribe Laeliinae, the most extensively © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 Subtribe Aerangidinae (32/315) x = 25 Genera with the number of species known/analysed 240 L. P. FELIX and M. GUERRA studied subtribe, exhibited the greatest numerical chromosomal diversity, varying from 2n = 38 to 2n = c. 240. These species had generally symmetrical karyotypes, with metacentric and submetacentric chromosomes, and prophase chromosomes with a similar condensation pattern. All species investigated of Brassavola R.Br. (Fig. 2F, G), Encyclia Hook. (Fig. 3A–C), Sophronitis Lindl., Laelia marginata (Lindl.) L.O.Williams (Fig. 3K) and Scaphyglottis fusiformis (Griseb.) R.E.Schult. (Fig. 4D) had 2n = 40. In the genus Cattleya Lindl., C. amethystoglossa Linden & Rchb.f. ex R.Warner (Fig. 2H), C. granulosa Lindl. (Fig. 2J) and the accession Santa Cruz 3263 of C. walkeriana had 2n = 40, whereas C. nobilior var. nobilior Rchb.f. and C. nobilior var. amaliae Pabst (Fig. 2K) had 2n = 42. Cattleya walkeriana accession Santa Cruz 3149 (Fig. 2L) and C. elongata Barb.Rodr. were polyploids with 2n = c. 80 (Fig. 2I). Prosthechea fragrans (Sw.) W.E.Higgins (Fig. 4B) and P. vespa (Vell.) W.E.Higgins (Fig. 4C) had 2n = 40, whereas P. caetensis (Fig. 4A) had 2n = 80. In Epidendrum L., 2n = 40 was observed in most species [E. difforme Jacq. (Fig. 3F), E. avicula Lindl. (Fig. 3G), E. latilabre Lindl. (Fig. 3H) and E. rigidum Jacq. (Fig. 3I)], whereas E. ellipticum Sessé & Moc. (Fig. 3E) and E. cinnabarinum Salzm. ex Lindl. (Fig. 3D) displayed 2n = 68 and 2n = c. 240, respectively. Meiotic analysis of a sample of E. cinnabarinum collected in Serraria, Paraíba, revealed the formation of up to five tetravalents during diakinesis, and the occurrence of bridges and lagging chromosomes during anaphases I and II. Haploid numbers ranged from n = 105 to n = 122, in prometaphase II cells, to n = 108 to n = 124 in pollen mitosis (in a total of 66 cells analysed). Jacquiniella globosa (Jacq.) Schltr. with 2n = 38 had large and small chromosomes. In several species of subtribe Laeliinae, including Jacquiniella globosa (Figs 1D, 3J), Cattleyella araguaiensis (Pabst) Van den Berg & M.W.Chase (Figs 1E, 2M) and Epidendrum cinnabarinum (Fig. 3D), some entire chromosomes were late condensing. These chromosomes were generally small (approximately 0.5 mm), difficult to visualize during prophase and apparently did not have © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 Figure 1. Prometaphase chromosomes and interphase nuclei of representatives of Sobraliinae, Laeliinae and Angraecinae. A, Eleanthus brasiliensis showing more condensed chromatin in two larger and 10 smaller chromosomes. B–C, interphase nuclei of the diffuse type in Campylocentrum pernambucense (B) and the complex chromocentric type in Malaxis excavata (C). D–E, Jacquiniella globosa (D) and Catleyella araguaiensis (E) showing late-condensed chromosomes (arrows). CHROMOSOMAL VARIABILITY OF EPIDENDROIDEAE 241 the condensed proximal chromatin observed in all other chromosomes. In subtribe Pleurothallidinae, Acianthera ochreata (Lindl.) Pridgeon & M.W.Chase had 2n = 40 (Fig. 4E) and Stelis sp. had 2n = 32 (Fig. 4F), both with small chromosomes. The latter had a slightly asymmetric karyotype. Malaxis excavata, tribe Malaxideae, had n = 15 in the first microspore mitosis (Fig. 4G) and 2n = 30 in somatic mitosis. Malaxis pubescens Kuntze showed 2n = 24 and chromosomes larger than those of M. excavata (Fig. 4H). In the tribe Vandeae, Polys- tachya estrelensis Reichb.f. had 2n = 80, and, in subtribe Angraecinae, Campylocentrum crassirhizum Hoehne (Fig. 4J) and C. pernambucense (Fig. 4K) had 2n = 38, and C. amazonicum Cogn. had 2n = c. 80 (Fig. 4I). In tribe Dendrobieae, Bulbophyllum cribbianum Toscano, B. sanderianum Rolfe and Bulbophyllum sp. 1 had 2n = 38 (Fig. 4L–N), whereas Bulbophyllum sp. 2 (Fig. 4O) had 2n = 80. In B. sanderianum and Bulbophyllum sp. 1, two of the four small chromosomes were only partially condensed during metaphase (Fig. 4M–N, arrows). © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 Figure 2. Chromosome complements of subtribes Sobraliinae, Ponerinae, Bletiinae and Laeliinae. A, Eleanthus brasiliensis (2n = 50). B, Sobralia liliastrum (2n = 48). C, S. sessilis (2n = 48). D, Isochilus linearis (2n = 40). E, Bletia catenulata (2n = 40). F, Brassavola tuberculata (2n = 40). G, B. nodosa (2n = 40). H, Cattleya amethystoglossa (2n = 40). I, C. elongata (2n = 80). J, C. granulosa (2n = 40). K, C. nobilior var. amalieae (2n = 42). L, C. walkeriana var. princeps (2n = c. 80). M, Cattleyella araguaiensis (2n = 40). N, Dimerandra emarginata (2n = 40). Scale bar in (O), 5 mm. 242 L. P. FELIX and M. GUERRA Appendix 1 lists the chromosome numbers of 115 genera and 844 species of subfamily Epidendroideae, compiled from the chromosome number indexes and our own data. Counts for the cymbidioid clade (sensu Dressler, 1993) published previously (Felix & Guerra, 2000) were not included here. Subfamily Epidendroideae (Table 2) shows a predominance of genera with the basic numbers x = 19 (tribes Podochyleae Pfitzer and Vandeae) and x = 20 (tribes Epidendreae, Arethuseae and Colabieae Pfitzer). Some disparate basic numbers are also observed, including x = 16 in Stelis (subtribe Pleurothallidinae), Bletilla Rchb.f. (Coelogyninae), Phreatia Lindl. and Thelasis Blume (Thelasiinae), and principally x = 23, 24 and 25 in various genera of Aerangidinae Summerh. Other tribes, such as Sobralieae and Neottieae, are currently insufficiently sampled or have variable basic numbers among genera. DISCUSSION The present work provides the first chromosome counts for 31 species, including the first counts for six genera (Elleanthus C.Presl, Sobralia Ruiz & Pav., Isochilus R.Br., Cattleyella Van den Berg & M.W.Chase, Jacquiniella Schltr. and Acianthera Scheidw.) and for subtribe Ponerinae and tribe Sobralieae. Previous chromosome counts were confirmed for Dimerandra emarginata (G.Mey.) Hoehne (Guerra, 1986), Sophronitis purpurata (Lindl. & Paxton) Van den Berg & M.W.Chase [as Laelia purpurata Lindl. & Paxton (Moore, 1977)], Brassavola tuberculata Hook. (Daviña et al., 2009), Encyclia advena (Rchb.f.) Porto & Brade (as E. megalantha (Barb.Rodr.) Porto & Brade), Epidendrum avicula (as Lanium avicula Lindl. ex Benth.), Laelia marginata (Lindl.) L.O.Williams (as Schomburgkia crispa Lindl.) © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 Figure 3. Chromosome complements of subtribe Laeliinae. A, Encyclia advena (2n = 40). B, E. flava (2n = 40). C, E. oncidioides (2n = 40). D, prometaphase of Epidendrum cinnabarinum (2n = c. 240). E, E. elipticum (2n = 68). F, E. difforme (2n = 40). G, E. avicula (2n = 40). H, E. latilabre (2n = 40). I, E. rigidum (2n = 40). J, Jacquiniella globosa (2n = 38). K, Laelia marginata (2n = 40). Scale bar in (I), 5 mm. CHROMOSOMAL VARIABILITY OF EPIDENDROIDEAE 243 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 Figure 4. Chromosome complements of tribe Malaxideae, subtribes Laeliinae, Pleurothallidinae, Polystachyinae, Angraecinae and Dendrobiinae. A, Prosthechea caetensis (2n = 80). B, P. fragrans (2n = 40). C, P. vespa (2n = 40). D, Scaphyglottis fusiformis (2n = 40). E, Acianthera ochreata (2n = 40). F, Stelis sp. (2n = 32). G, metaphase of first mitotic division of microspore of Malaxis excavata (n = 15). H, M. pubescens (2n = 24). I, Campylocentrum amazonicum (2n = c. 80). J, C. crassirhizum (2n = 38). K, C. pernambucense (2n = 38). L, Bulbophyllum cribbianum (2n = 38). M, B. sanderianum (2n = 38). N, Bulbophyllum sp. 1 (2n = 38). O, Bulbophyllum sp. 2 (2n = c. 80). Scale bar in (P), 5 mm. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 244 L. P. FELIX and M. GUERRA insufficiently investigated. The numbers of n = 24 and 25 for tribe Sobralieae diverge from the lower chromosome numbers observed in the majority of species of Epidendroideae. The count of 2n = 54 for Sobralia ¥ amesiana Hort.Sander (Vij & Shekhar, 1985) from West Bengal, India, was transferred to the table of excluded numbers (see also Appendix 2) as it appears to be a misidentification. This genus is exclusively American (Bechtel, Cribb & Launert, 1992) and this binomial was included by Govaerts et al. (2009) in the World Checklist of Orchidaceae as an ‘unplaced name’. Tribe Epidendreae, the largest group of subfamily Epidendroideae (Chase et al., 2003), has karyological reports for 27 genera and 163 species, corresponding to 31% of the genera but less than 3% of the species. These data indicate n = 20 as the most widely occurring number among the three subtribes. Pleurothallis and Stelis display an exceptionally high variation in chromosome numbers, with 13 different numbers among the 17 species investigated here for these genera. The number 2n = 20 reported for Specklinia grobyi (Bateman ex Lindl.) Pridgeon & M.W.Chase (Nakata & Hashimoto, 1983) and for Liparis cordifolia Hook.f and L. glossula Rchb.f. (Goldblatt & Figure 5. Partial cladogram of subfamily Epidendroideae, based on Chase et al. (2003), with the probable basic number indicated for each tribe and subtribe. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 and Prosthechea fragrans (Sw.) W.E.Higgins (as ‘Hormidium fragrans’), all having 2n = 40 (Tanaka & Kamemoto, 1984), Polystachya estrellensis Rchb.f and Epidendrum nocturnum Jacq. with 2n = 80 (Blumenschein, 1960a), and E. cinnabarinum, with 2n = c. 240 (Guerra, 2000; Conceição, Oliveira & Barbosa, 2006). We found 2n = 40 for Epidendrum difforme, previously reported as 2n = 39–40 (Tanaka & Kamemoto, 1984). The count of 2n = 40 for Cattleya walkeriana (accession Santa Cruz 3149) confirmed the only previous count for the species (Blumenschein, 1960a). Additionally, 2n = 80 was found for another accession of C. walkeriana, indicating that this species has at least two ploidy levels. There are records of chromosome numbers for five of the six recognized genera of tribe Neottieae, but its basic number is still unclear. According to Chase et al. (2003), this tribe comprises an early branching group of Epidendroideae. In Neottieae, x = 20 is the basic number in Epipactis Sw. and the orthoploid series x = 14, 21 ± 1 occurs in Neottia L., Limodorum L. and Cephalanthera Rich., supporting the hypothesis of an ancestral basic number related to this series in the tribe and to the entire subfamily Epidendroideae (Fig. 5). Tribes Sobralieae and Calypsoe are also CHROMOSOMAL VARIABILITY OF EPIDENDROIDEAE habit because of the water stress associated with the epiphytic habit. As species with large genome size usually have large guard cells and a slower response to water stress, they are better adapted to the terrestrial lifestyle, whereas species with small genome size are free to occupy both habits. The only population analysed here of E. elipticum, from the Epidendrum secundum complex showed 2n = 68. However, an analysis of six populations of this complex displayed 2n = 28, 40, 48, 52, 56, 68 and 80 (Pinheiro et al., 2009). These data indicate that E. secundum is a highly polymorphic complex. It also showed intra- and inter-populational morphological variants (Pinheiro & Barros, 2007). It belongs to the Amphyglottidae group of Epidendrum (Pabst & Dungs, 1975), which comprises species with different chromosome numbers (n = 12, 14, 18, 19 and 28) (Hágsater & Arenas, 2005). In E. cinnabarinum, the count of 2n = c. 240 in four populations from Pernambuco and Paraíba and at least in one population from Bahia (Conceição et al., 2006) indicates that its ploidy is stable, in spite of some meiotic instability, as the occurrence of ring tetravalents and aneuploid spores, observed here, and anaphase bridges and lagging chromosomes, reported by Conceição et al. (2006). In tribe Podochileae, n = 19 occurred in the majority of the genera of subtribe Eriinae and in the sole species of Podochilinae analysed, whereas, in subtribe Thelasiinae, n = 16 was found in the only two species studied. In subtribe Eriinae Benth., Eria Lindl. had n = 19 in 27 of the 54 species investigated. The other four genera, with few species investigated, had n = 19 (Cryptochilus Wall. and Mediocalcar J.J.Sm), n = 20 (Ceratostylis Blume), and n = 12 and 21 (Porpax Lindl.). The tribe Arethuseae is monophyletic (Goldman et al., 2001), but do not have well-defined relationships with other Epidendroideae (Chase et al., 2003). In this tribe, n = 20 was the best represented haploid number in the genera of both subtribes that are cytologically known, with the exception of Bletilla with n = 16, and Dendrochilum Blume with n = 15 and n = 19. Tribe Malaxideae has a great diversity in chromosome numbers, with n = 21 predominating in Liparis and Malaxis and n = 15 in Oberonia. The other genera are quite small and insufficiently studied. The most probable basic numbers for the tribe are x = 15 and x = 21, n = 21 being the most frequent haploid number and n = 15 the second most frequent and the only one that occurs in all three genera. The occurrence of n = 14 in some species of Liparis and Malaxis, and the high frequency of n = 21 in these two genera, suggest that these numbers may be related to the polyploid series x = 7, 14, 21 that is quite common in orchids (Felix & Guerra, 2000). Liparis and Malaxis are arti- © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 Johnson, 1994) is probably the lowest diploid number for the subfamily. It seems to represent two independent examples of intense dysploid reduction rather than the ancestral chromosome number of these genera (Guerra, 2008). Similarly low haploid numbers, not representatives of the basic number, have also been found in a few genera of subtribe Oncidiinae (Felix & Guerra, 1999, 2000) and in other families, for example n = 6 in Aristolochiaceae (OhiToma et al., 2006) and n = 2 in Cyperaceae (Vanzela, Guerra & Luceño, 1996). Likewise, n = 10 in Specklinia Lindl. and Liparis Rich. can be a dysploid remnant of the ancestral number x = 7 or of a tetraploid lineage with n = 14. In subtribe Laeliinae, n = 20 is observed in all of the genera, except Jacquiniella, a small genus previously included in subtribe Ponerinae (Dressler, 1993), which also has n = 20. Small variations around this number have been observed in some species of Cattleya, Epidendrum, Laelia, Scaphyglottis, and Sophronitis. Cattleya nobilior Rchb.f., for example, which is morphologically close to C. walkeriana (Pabst & Dungs, 1975), differs from the other species of Cattleya by having 2n = 42, a rare number in this genus. Jacquiniella globosa (Jacq.) Schltr. and Scaphyglottis reflexa Lindl. (Sau & Sharma, 1983), both with 2n = 38, were included recently in the Scaphyglottis alliance by van den Berg et al. (2009). Tetraploid species with 2n = 80 occur in Cattleya, Epidendrum, Prosthechea, and Sophronitis. Intraspecific polyploidy occurred in C. bicolor Lindl. (Blumenschein, 1960a), in Sophronitis (Blumenschein, 1960b; YamagishiCosta & Forni-Martins, 2009) and in some species of Epidendrum (Blumenschein, 1960a; Pinheiro et al., 2009). This ploidy change seems to be related to morphological variants placed in distinct subspecies, as in C. bicolor ssp. bicolor with 2n = 40 and C. bicolor ssp. minasgeraensis Fowlie with 2n = 80 (Blumenschein, 1961). In C. walkeriana var. princeps (accession Santa Cruz 3149), the number 2n = 80 reported here was observed only in a cultivated specimen. In some lithophytic species of Sophronitis, the intraspecific polyploidy observed seems to be related to the acquisition of this habit, as has been reported for Sophronitis longipes (Rchb.f.) Van den Berg & M.W.Chase (Blumenschein, 1960b), S. rupestris Cogn. (Yamagishi-Costa & Forni-Martins, 2009), Oncidium aff. flexuosum, and O. varicosum Lindl. (Felix & Guerra, 2000). This tendency was also observed in the polyploids Epidendrum cinnabarinum, C. walkeriana var. princeps, and C. elongata, which have the lithophytic or terrestrial habit. Terrestrial species of subfamilies Vanilloideae and Cypripedioideae also have genome sizes notably higher than most epiphytic orchids. Leitch et al. (2009) suggested that orchids with large genomes are restricted to a terrestrial 245 246 L. P. FELIX and M. GUERRA presence of one or more pairs of late-condensing chromosomes. Among orchids, this type of chromosome was recorded for the genus Habenaria (Felix & Guerra, 1998), in some species of subtribe Oncidiinae (Felix & Guerra, 2000) and in the genus Spiranthes of New Zealand (Dawson, Maloy & Beuzember, 2007). Late-condensing chromosomes have also been observed in other angiosperms, e.g. in some Velloziaceae (Melo et al., 1997) or in species of Arachis L. (Fabaceae) with the A genome, where they are used as a cytogenetic marker (Fernandez & Krapovickas, 1994). Cattleyella araguaiensis, a species recently separated from Cattleya (van den Berg & Chase, 2004b) stood out from the other species of Cattleya that were analysed by having at least one pair of late-condensing chromosomes. The data presented here reaffirm the extensive variation in chromosome numbers encountered in subfamily Epidendroideae. Only five of the 128 genera with chromosome records for more than five species have stable chromosome numbers. As the large majority of these genera have haploid numbers between 15 and 25 (principally 19 and 20), each of these numbers may have arisen independently several times during the evolution of the family. The principal karyological difference between tribes Epidendreae, Cymbidieae, Podochileae and Vandeae, together comprising the large majority of the Epidendroideae orchids, is the predominance of n = 20 in Epidendreae, n = 21 in Cymbidiae (Felix & Guerra, 2000) and n = 19 in the last two tribes. Multiples of n = 7, especially n = 21 or nearly 21, also predominate in the majority of the genera of subfamily Orchidoideae (Martínez, 1985; Yokota, 1990; Pridgeon et al., 1997; Felix & Guerra, 2005). Among orchids in general, n = 21 is a recurring chromosome number that appears to be related to the probable basic number of the family x1 = 7 (Felix & Guerra, 1999, 2000, 2005). The principal polyploid series has been hypothesized to be n = 7, 14, 21, with a variation of ± 1 at each ploidy level (Felix & Guerra, 2005). According to this assumption, dysploid reduction occurred in the large majority of the tribes of Epidendroideae (x = 21, 20, 19), whereas ascending dysploidy occurred only in the subtribe Angraecinae sl. (x = 25). ACKNOWLEDGEMENTS The authors would like to thank CAPES (Coordenação de Aperfeiçoamento de Pessoal de Nível Superior) and CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico) for their financial support and Celso Pires from Taquaritinga do Norte, Pernambuco, for granting us access to his private collection to perform the karyological analyses. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 ficial cosmopolitan genera (Cameron, 2005), which may explain their large numerical chromosomal variations. Similarly, the large and chromosomally variable genus Habenaria (Orchidoideae) (Felix & Guerra, 1998, 2005) was also recognized as polyphyletic (Bateman et al., 2003). Oberonia, which is exclusively epiphytic (a type of habitat that is considered primitive in this subtribe), has n = 15 or 30 and is related to a group of ‘primitive’ epiphytic Liparis (Cameron, 2005). At least one of these Liparis species (L. viridiflora) has n = 15 (Mehra & Sehgal, 1980), indicating the karyological affinity between these two genera. The occurrence of n = 15 in Malaxideae further supports the hypothesis that the karyological evolution of the family was principally through dysploidy of approximately one chromosome in the three principal ploidy levels, n = 7, 14, 21 ± 1 (Felix & Guerra, 2000, 2005). In tribe Vandeae, a group of orchids principally known from the Old World, n = 19 predominates in subtribes Aeridinae and Angraecinae, whereas n = 25 predominates in Aerangidinae and n = 20 in Polystachyinae (sensu Chase et al., 2003). The latter is considered the sister group of the other subtribes of Vandeae (Chase et al., 2003; van den Berg et al., 2005; Carlsward et al., 2006). Polystachya, the largest genus of Polystachyinae and the only one with known chromosomal counts, had n = 20 in 35 of the 41 species analysed. This haploid number also occured in various genera of Aeridinae and in some Angraecinae, suggesting that x = 20 is the basic number for the tribe Vandeae and that x = 19 represents a secondary basic number. Subtribes Aerangidinae and Angraecinae are considered individually to be polyphyletic, but together they form the more widely circumscribed monophyletic clade Angraecinae s.l. (Carlsward et al., 2006). Cytologically, the two tribes have shown a distinct base number (x = 19 and x = 25) and at least Aerangidinae, with x = 25, seems to represent a welldefined branch of Epidendroideae. Among the subtribes with undefined taxonomic positions (Chase et al., 2003), Agrostophyllinae and Collabinae predominantly have n = 20, whereas in Dendrobiinae n = 19 predominates in all genera except Epigeneium and Saccoglossum (both with n = 20 in the single species analysed). Therefore, Dendrobiinae may be cytologically related to tribe Vandeae, which includes some subtribes with x = 19. Among Dendrobiinae, the two genera largely investigated, Bulbophyllum and Dendrobium, showed relatively little variation. In contrast, in Collabiinae, the two genera with many species investigated, Calanthe and Phaius, exhibited a large variation in chromosome numbers. A karyotypic aspect observed in four genera and seven species of subfamily Epidendroideae is the CHROMOSOMAL VARIABILITY OF EPIDENDROIDEAE REFERENCES Conceição LP, Oliveira ALPC, Barbosa LV. 2006. Characterization of the species Epidendrum cinnabarinum Salzm. 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Index to plant chromosome numbers 1967–1971. Regnum Vegetabile 90: 1–539. Moore RJ, ed. 1974. Index to plant chromosome numbers 1972. Regnum Vegetabile 91: 1–108. CHROMOSOMAL VARIABILITY OF EPIDENDROIDEAE 249 Appendix 1. Chromosome numbers in epidendroid orchids, organized according to Chase et al. (2003), except the cymbidioid phylad sensu Dressler (1993). TAXON TRIBE NEOTTIEAE Aphyllorchis caudata Rolfe ex Downie Cephalanthera damasonium (Mill.) Druce (as C. alba) C. damasonium (Mill.) Druce longifolia (L.) Fritsch (as C. ensifolia) epipactoides Fisch. & Meyer erecta (Thunb.) Blume erecta (Thunb.) Blume (as C. shizuoi) erecta f. subaphylla (Miyabe & Kudô) M.Hiroe (as C. subaphylla) falcata (Thunb.) Blume longibracteata Blume C. longifolia (L.) Fritsch(as C. grandiflora) C. longifolia (L.) Fritsch (= C. ensifolia) 2n Source 18 ca. 36 36 36 TK84 F69 TK84, G84, 88, GJ91, GJ96 TK84, G84, GJ90 G85, G88 TK84, G88, GJ91 F69, TK84 G85 TK84, G88, GJ91 TK84, G88, GJ91 GJ94 TK84, G88 TK84, G84, G85, G88, GJ91 TK84, G84, G88, GJ91 G85 F69 TK84, G84, 85, 88, GJ03 F69 GJ03 16, 17 18 16 32, 34 32 34 32 34 34 32 34, 36 36 32 C. rubra (L.) Rich. 36 Epipactis. atrorubens (Hoff.) Schult. (as E. atropurpurea) 44 32, 36 30, 38, 40, 60 E. E. E. E. E. E. atrorubens (Hoff.) Schult. (as E. rubiginosa) exilis P.Delforge (as E. gracilis B.Baumann & H.Baumann) fageticola (C.E.Hermos.) Devillers-Tersch. falcata Sw. gigantea Dougl. ex Hook. helleborine (L.) Crantz E. E. E. E. helleborine subsp. latina Rossi & Klein helleborinae subsp. tremolsii Klein helleborine subsp. orbicularis (K.Richt.) E.Klein (as E. distans) latifolia Sw. E. E. E. E. E. leptochilla (Godf.) Godf. leptochila subsp. futakii (Mered’a & Potucek) Kreutz (as E. futakii) microphylla (Ehrh.) Sw. muelleri Godfrey palustris (L.) Crantz E. E. E. E. E. E. E. E. E. papillosa Franch. & Savat. papillosa Franch. & Savat. (as E. sayekiana) persica (Soó) Hausskn. ex Nannf. phyllanthes G.E.Sm. phyllanthes var. phyllanthes (as E. confusa) placentina L.Bongiorni & P.Grunanger purpurata Sm. (as E. pseudopurpurata) royleana Lindl. veratrifolia Boiss. & Hohen. (as E. consimilis) 20 40 40 20 20 40 40 40 36 40 38 40, ca. 40 40 40 21 20 18 56, 64 36 50 40 40 (34 + 6B) 42 40 36 TK84 TK84 G81, G84, G84, GJ91, GJ96, GJ03 GJ06 GJ91 G88, GJ91 GJ03 G81, TK84, G84, GJ90, GJ94 TK84 GJ03 TK84, GJ03 GJ03 G81, TK84, G84, GJ91, GJ00, GJ03 TK84, GJ94 TK84 TK84 TK84 TK84 GJ03 G88, GJ96, GJ03 TK84, G84, GJ90 TK84, G85, GJ90, GJ91, GJ94 GJ03 TK84 A08 G88 G85 TK84, G84, GJ90 TK84, GJ90, GJ94 TK84, G88, GJ91, 50 48 PW PW 20 20 18, 18 + 2B 19 camtschatea (L.) Rchb. cordata (L.) Rich. (as Epipactis cordata) inayatii (Duthie) Beaverd listeroides Lindl. nidus-avis (L.) Rich. TRIBE SOBRALIEAE Elleanthus brasiliens Rchb.f. Sobralia sessilis Lindl. 32, 38, 40 20 40 40 40 20 36 40 40 38 40 20 20 20 Limodorum abortivum (L.) Sw. Neottia asiatica Ohwi N. N. N. N. N. 24 40 40 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 C. C. C. C. C. C. C. n 250 L. P. FELIX and M. GUERRA Appendix 1. Continued TAXON n Source TRIBE GASTRODIEAE Gastrodia cunninghamii Hook.f. G. minor Petrie G. aff. sesamoides Gastrodia sp. 40 40 38–40 38–40 D07 D07 D07 D07 TRIBE CALYPSOEAE Calypso bulbosa (L.) Oakes 28 G84 40 PW 40 40 60 PW TK84 TK84 40 40 PW G85 43 40 40 40 G85 G85 G85 G85 TK84 G85 G85 TK84 TK84 G85 G85 G85 G85 G85 G85, GJ94 G85 G85 G85 G85 G85 TK84 G85 G85 TK84 G85 G85 G85 G85 G85 TRIBE EPIDENDREAE Subtribe Ponerineae Isochilus linearis (Jacq.) R.Br. Subtribe Bletiinae Bletia catenulata Ruiz & Pavon B. catenulata (as B. rodriguesii Cogn.) B. purpurea (Lam.) DC. (as B. verecunda) Subtribe Pleurothallidinae Acianthera ochreata (Lindl.) Pridgeon & M.W.Chase A. aurantiolateritia (Speg.) Pridgeon & M.W.Chase (as Pleurothallis aurantiolateritia) A. brevipes (H. Focke) Pridgeon & M.W.Chase (as P. brevipes) A. aff. casapensis (Lindl.) Pridgeon & M.W.Chase (as Pleurothallis aff. cofeicola) A. luteola (Lindl.) Pridgeon & M.W.Chase (as Pleurothallis luteola) A. teres (Lindl.) Borba (as Pleurothallis teres) A. pubescens (Lindl.) Pridgeon & M.W.Chase (as Pleurothallis vittata) Anathallis obovata (Lindl.) Pridgeon & M.W.Chase (as P. obovata) Masdevallia civilis Rchb.f. & Warsc. M. coccinea Linden ex. Lindl. Pabstiella tripterantha (Rchb.f.) F. Barros (as P.procumbens) Pleurothallis alopex Luer P. bivalvis Lindl. P. bivalvis Lindl. (as P. chanchamayoensis) P. carinata Schweinf. P. matudana C.Schweinf. P. revoluta (Ruiz. & Pav.) Garay P. ruscifolia (Jacq.) R.Br. P. saccatilabia Schweinf. P. tridentata Klotz P. xanthochlora Rchb.f. Pleurothallis sp. Scaphosepalum verrucosum (Rchb.f.) Pfitzer (as Masdevallia ochthodes) Specklinia grobyi (Bateman ex Lindl.) F.Barros (as Pleurothallis grobyi) Stelis argentata Lindl. S. ciliaris Lindl. S. gelida (Lindl.) Pridgeon & M.W.Chase (as Pleurothallis gelida) S. pachyglossa (Lindl.) Pridgeon & M.W.Chase (as Pleurothallis pachyglossa) S. restrepioides (Lindl.) Pridgeon & M.W.Chase (as Pleurothallis restrepioides) S. segoviensis (Rchb.f.) Pridgeon & M.W.Chase (as Pleurothallis segoviensis) S. velaticaulis (Rchb.f.) Pridgeon & M.W.Chase (as Pleurothallis velaticaulis) Subtribe Laeliinae Brassavola cucullata R.Br. B. grandiflora Lindl. B. nodosa Hook. B. tuberculata Hook. B. tuberculata Hook. (as Brassavola perrinii) Broughtonia sanguinea (Sw.) R.Br. Cattleya amethystoglossa Linden & Rchb.f. ex R.Warner C. bicolor Lindl. C. bicolor var. bicolor (as C. bicolor var. measuresiana) C. dormaniana Rchb.f. C. dowiana Bateman C. dowiana var. aurea (Linden) B.S.Williams & Moore C. elongata Barb. Rodr. C. forbesii Lindl. C. gaskelliana (N.E.Br.) B.S.Williams 21 43, 42, 45 36 44 16 36 77 84 40 68, 72 40, 80, ca. 80 38 38 36 40 38 64 20 38 16 32, 64 38 76, 78 42 34 40 20 20 20 20 20 40 40 40 40 40, 80 80 40 40 40 80 54–60 40 TK84 TK84 TK84, PW DA09, PW TK84 TK84 PW TK84 TK84 TK84 TK84 TK84 PW TK84 TK84 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 2n CHROMOSOMAL VARIABILITY OF EPIDENDROIDEAE 251 Appendix 1. Continued TAXON Cattleya granulosa Lindl. C. guttata Lindl. C. harrisoniana Bateman ex Lindl. C. intermedia Graham ex Hook. C. intermedia var. alba C. labiata Lindl. E. E. E. E. avicule Lindl. (as Lanium avicula) blepharistes Barker ex Lindl. (as E. funckii) calanthum Rchb.f. & Warsc. ciliare L. E. E. E. E. E. E. cinnabarinum Salzm. cochlidium Lindl. cooperianum Bateman (as E. longispathum Barb. Rodr.) cristatum Ruiz & Pavon (as E. raniferum Lindl.) denticulatum Barb. Rodr. difforme Jacq. E. diffusum Sw. E. elipticum Grah. E. E. E. E. E. E. E. E. E. E. E. elongatum Jacq. flexuosum G.Mey fulgens Brongn. fulgens Brongn. (as E. mosenii) ibaguense Kunth lanipes Lindl. latilabrum Lindl. loefgrenii Cogn. magnoliae Muhl. (as E. conopseum) myrmecophorum Barb. Rodr. munroeanum 20 20 20, 21 20 20 20 20 20 20 20 20 108–124 20 20 20 2n Source 40 40 40 40 41 + 1f 40 46 40, 41 40 40 42 42 40 40 40 40 40 80 40 40 46, 56, 76 40 40 40 40 40 40 40 40 40 40 40 36 38 38, 40 PW TK84 TK84 TK84 TK84 M73, TK84 TK06 TK84 TK84 TK84 PW PW TK84 TK84 TK84 TK84 TK84, PW PW TK84 TK84 GJ91 PW M73 G88, PW TK84 PW TK84 PW TK84 PW TK84 TK84 GJ94 GJ94 TK84, G, J94, DA09, PW 40 40 30 40 40, 80, 160 ca. 240 28 40 40 40 39–40 40 40 56 68 GJ94 PI09 TK84, G85 TK84 PW PI09 TK84 TK84 TK84, PI09 TK84 PW TK84 TK84 PW 24 28 24 70 40 40 40 40 120 40 PI09 PI09 TK84 PI09 G85 PW TK84 TK84 PI09 TK84 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 C. labiata var. amesiana C. lueddemanniana Rchb.f. C. mossiae C.Parker ex Hook. C. nobilior Rchb.f. Cattleya nobilior var. amaliae Pabst C. percivaliana (Rchb.f.) O’Brien C. tigrina Rich. (as C. leopoldii) C. trianae Linden & Rchb.f. C. velutina Rchb.f. C. walkeriana Gardn. C. walkeriana var. princeps L.C.Menezes C. warnerii T.Moore ex Warner Cattleya warscewiczii Rchb.f. (as C. gigas) Cattleya sp. Cattleyella araguiensis (Pabst) van den Berg Caularthron bicornutum (Hook.) Raf. Dimerandra emarginata (Meyer) Hoehne D. stenopetala Schltr. Encyclia advena (Rchb.f.) Porto & Brade Encyclia advena (as E. megalantha) E. flava (Lindl.) Porto & Brade E. patens Hook. (as E. odoratissima) E. oncidioides (Lindl.) Schltr. E. serroniana (Barb. Rodr.) Hoehne (as E. odoratissima var. serroniana) E. tampensis (Lindl.) Small (as Epidendrum tampense) Epidendrum angustatum (T.Hashim.) Dodson (as Neolehmannia angustata) Epidendrum appendiculatum T.Hashimoto E. avicula Lindl. n 252 L. P. FELIX and M. GUERRA Appendix 1. Continued TAXON n E. nocturnum Jacq. 20 19 40, 70 60 40 28, 40, 48, 52, 68, E. rigidum Jacq. E. secundum Jacq. E. secundum Jacq. (as E. brachyphyllum) E. secundum Jacq. Epidendrum aff secundum (as Epidendrum aff. brachyphyllum) E. xanthinum Lindl. Guarianthe aurantiaca (Bateman ex Lindl.) Dressler & W.E.Higgins (as Cattleya aurantiaca) G. bowringiana (O’Brien) Dressler & W.E.Higgins (as Cattleya bowringiana) Jacquiniella globosa (Jacq.) Schltr. Laelia albida Bateman ex Lindl. L. anceps Lindl. Laelia anceps subsp. dawsonii (J.Anderson) Rolfe (as L. anceps var. sanderiana) L. autumnalis (Lex.) Lindl. L. gouldiana Rchb.f. L. peciosa (Kunth) Schltr. (as L. grandiflora) L. marginata (Lindl.) L.O.Williams L. marginata (Lindl.) L.O.Williams (as Schomburgkia crispa Lindl.) L. rubescens Lindl. L. rubescens Lindl. (as L. peduncularis) Leptotes unicolor Barb. Rodr. Prosthechea brassavolae (Rchb. f.) W. E. Higgins (as Epidendrum brassavolae) P. caetensis (Bicalho) W.E.Higgins P. calamaria (Lindl.) W.E.Higgins (as Hormidium calamarium) P. campylostalix (Rchb. f.) W.E.Higgins (as Epidendrum campylostalix) P. citrina (Lex.) W.E.Higgins (as Cattleya citrina) P. cochleata (L.) W.E.Higgins (as E. cochleatum) P. fragrans (Sw.) W.E.Higgins P. fragrans (Sw.) W.E.Higgins (as Hormidium fragrans) P. glumacea (Lindl.) W.E.Higgins (as Hormidium glumaceum) P. lindenii (Lindl.) W.E.Higgins (as E. lindenii) P. linkiana (Klotzsch) W.E.Higgins (as Epidendrum linkianum) P. mariae (Ames) W.E.Higgins (as E. marieae) P. ochracea (Lindl.) W.E.Higgins (as Epidendrum ochraceum) P. prismatocarpa (Rchb.f.) W.E.Higgins (as Epidendrum prismatocarpum) P. vespa (Vell.) W E Higgins Prosthechea vespa (Vell.) W E Higgins (as Hormidium variegatum) Psychilis atropurpurea (Willd.) Sauleda (as Epidendrum atropurpureum) Rhyncholaelia digbyana (Lindl.) Schltr. (as Brassavola digbyana) Scaphyglottis reflexa Lindl. (as Hexisea reflexa) S. prolifera (R.Br.) ex Lindl.) Cogn. S. fusiformis (Griseb.) R.E.Schult. Sophronitis briegeri (Blumensch. ex Pabst) Van den Berg & M.W.Chase (as Laelia briegeri) S. caulescens (Lindl.) Van den Berg & M.W.Chase (as Laelia calulescens) 30 56 14 ca. 80 40 28, 30, 60 40 20 40 38 42, ca. 63 40 40 41, 42 40, 60 20 20 20 20 20 ca. 20 20 20 20 19 40 40 40 40–44 40 40 80 40 40 40 40 40 40 56 Source TK84 TK84 TK84 PW TK84 G81 TK84 TK84 TK84 PI09 TK84 PI09 M73 TK84 PI09 TK84, PW PI09 TK84 TK84 TK84 TK84 G88 PI09 TK84 TK84 PW TK84 TK84 TK84 TK84 TK84 TK84 PW TK84 TK84 TK84 TK84, DA09 TK84 PW TK84 TK84 40 40 40 40 40 40 40 40 38 40 40 80 TK84 PW TK84 TK84 TK84 TK84 TK84 TK84 TK84 TK84 PW TK84 TK84 TK85 G85 GJ94 PW TK84 80 TK84 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 ca. 80 40, 80 80 42–48 40 40 40 40 52 56 120 Epidendrum nocturnum var. guadeloupense E. obrienianum Rolfe E. paniculatum Barb. Rodr. (as E. floribundum Kunth) E. patens Sw. E. propiquum Rich. & Gall. E. puniceoluteum F.Pinheiro and F.Barros E. purpureum Barb. Rodr. E. radicans Pav. 2n CHROMOSOMAL VARIABILITY OF EPIDENDROIDEAE 253 Appendix 1. Continued TAXON TRIBE PODOCHILEAE Subtribe Eriinae Ceratostylis subulata Blume Cryptochilus lutea Lindl. C. sanguineus Wall. Eria acervata Lindl. E. alba L. E. amica Rchb.f. (as E. confusa) E. E. E. E. bambusifolia Lindl. biflora Griff. brachystachya Rchb.f. bractescens Lindl. E. E. E. E. E. E. E. bulbophylloidea Tang & Wang clemensiae Leavitt corneri Rchb.f. corneri Rchb. f. (as E. yakuschimensis) coronaria (lindl.) Rchb.f. crassicaulis Hook.f. crassipes Ridl. (as E. reptans) E. microchilos Lindl. E. muscicola Lindl. 40 40 TK84, DA090 TK84 80 40 80 40 TK84 TK84 TK84 TK84 40 40 40 40, 60, 80 40 40 TK84 TK84 PW TK84 TK84 TK84 TK84 TK84 PW M77 TK84 40 40 40 80 40 19 19 38 40 20 18 19 21 18 19 38 42 44 40 38 42 44 36 36 36 40 76 42 ± 2 24 + 5–7B 24 + 10B 24 + 5B 42 20 38 38 38 38 38 44 38 E. exilis Hook.f. E. feddeana Schltr. E. ferruginea Lindl. E. fitzalanii F.Muell. (as E. solomonensis) Eria floribunda Lindl. E. floribunda Lindl. (as E. giungii) E. gigantea Ames E. graminifolia Lindl. E. hyacintoides (Blume) Lindl. E japonica Maxim.(as E. arisanensis) E. javanica (Sw.) Blume Eria lasiopetala (Willd.) Ormerod (as E. flava) Source 20 E. cymbidifolia Ridl. E. dalzellii Lindl. E. excavata Lindl. 2n 20 19 66 38 38 40 36 40 38 24 + 5–11B 18 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 G88 M73 M73 G85, G88, GJ91, GJ96 G81 M77 G88, GJ94 G81, GJ91 G81, G84 TK84 TK84 G88 GJ91, GJ96 TK84 TK84, G88 TK84 M73, G88 G81 TK84 G88 TK84 TK84 G84 G84 G84 M73 G84 TK84 G88 G84 G88 TK84 G88 TK84 TK84 M73, G84 G88 TK84 G88 G81 G84 TK84, G84 G81 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 S. cernua Lindl. S. cinnabarina (Bateman ex Lindl.) Van den Berg & M.W.Chase (as Laelia cinnabarina) S. crispata (Thunb.) Van den Berg & M.W.Chase (as Laelia crispilabia) S. crispata (Thunb.) Van den Berg & M.W.Chase (as Laelia flava) S. crispata (Thunb.) Van den Berg & M.W.Chase (as Laelia rupestris) S. esalqueana (Blumensch. ex Pabst) Van den Berg & M.W.Chase (as Laelia esalqueana) S. fournieri (Cogn.) Van den Berg & M.W.Chase (as Laelia oestemayeri) S. harpophylla (Rchb. f.) Van den Berg & M.W.Chase (as Laelia hapophylla) S. lobata (Lindl.) van den Berg & M.W.Chase S. longipes (Rchb. f.) Van den Berg & M.W.Chase (as Laelia longipes) S. milleri (Blumensch. ex Pabst) Van den Berg & M.W.Chase (as Laelia milleri) S. mixta (Hoehne) Van den Berg & M.W.Chase (as Laelia mixta) S. perrinii (Lindl.) Van den Berg & M.W.Chase (as Laelia perrini) S. pumila (Hook.) Van den Berg & M.W.Chase (as Laelia pumila) S. purpurata (Lindl. & Paxton) van den Berg & M.W.Chase S. purpurata (Lindl. & Paxton) van den Berg & M.W.Chase (as Laelia purpurata) S. tereticaulis (Hoehne) Van den Berg & M.W.Chase (as Laelia tereticaulis) n 254 L. P. FELIX and M. GUERRA Appendix 1. Continued TAXON mysorensis Lindl. ovata Lindl. ovata Lindl. (E. luchensis) ovata var. retroflexa (Lindl.) Garay & H.R.Sweet (as E. nudicaulis) pachystachya Lindl. paniculata Lindl. pannea Lindl. philippinensis Ames E. E. E. E. E. E. queenslandica T.E.Hunt reticosa Wight retusa (Blume) Rchb.f. ringens Rchb.f. spicata (D.Don) Hand-Mazz. spicata (D.Don) Hand-Mazz. (as E. convallarioides) E. stricta Lindl. E. tomentosa Hayata E. woodiana Ames E. yakushimensis Nakai Eria sp. Eria sp. Eria sp. Mediocalcar agathodaemonis J.J.Sm. M. latifolium Schltr. M. paradoxum subsp. latifolium (Schltr.) Schuit. (as M. latifolium) M. aff. pygmaeum M. uniflorum Schltr. (as M. sepikanum) Porpax fibuliformis King & Pantl. P. jerdoniana (Wight) Rolfe P. meriax King & Pantl. Pseuderia smithiana C.Schweinf. Trichotosia lagunensis (Ames) Schuit. & de Vogel (as Eria lagunensis) T. pulvinata (Lindl.) Kraenzl. (as Eria rufinula) Subtribe Podochilinae Podochilus cultratus Lindl. 2n Source 18 38 38 36 40 38 38 40 38 44 TK84 G88 M77, TK84 G85 G88 G81, GJ91 G88, GJ91 TK84 TK84 G88 TK84 G88 TK84 G81, GJ94 M73 TK84, G88 G88 G88 TK84 38 38 38 38 G84 GJ91 GJ96 G88 38 85 38 24 42 G88 G88 G88 G85 G84 TK84 GJ03 TK84 G88 19 + 0–2B 38 40 38 38 42 38 44 38 20 19 12 40 44 38 19 Subtribe Thelasiinae Phreatia elegans Lindl. Thelasis capitata Blume TRIBE ARETHUSEAE Subtribe Arethusinae Anthogonium gracile Lindl. 20 19 Anthogonium sp. Arethusa bulbosa L. Arundina graminifolia (D.Don) Hochr. A. graminifolia (D.Don) Hochr. (as A. bambusifolia) A. graminifolia (D.Don) Hochr. (as A. chinensis) Arundina sp. Arundina spp. Calopogon barbatus (Walter) Ames C. multiflorus Lindl. C. pallidus Chapm. C. pulchellus (Salisb.) R.Br. Calopogon tuberosus (L.) Britton, Sterns & Poggenberg C. tuberosus (L.) Britton (as C. pulchellus) Eleorchis japonica (A.Gray) Maek. 20 20 16 ca. 13 Subtribe Coelogyninae Bletilla formosana (Hayata.) Schltr. B. hyacinthia 18 16 M73 32 32 G81 G88 42 40 38 32 40 42 42 42 M73 M73 M73, G84 G84 G84 M73, G81, G84, G85, GJ90 G84, G85 GJ91, GJ96 TK84 G84 M77 M77 M77 42 ca. 26 40 M77 TK84 M73 36 M73 TK84 TK84 38 42 40 40 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 E. E. E. E. E. E. E. E. n CHROMOSOMAL VARIABILITY OF EPIDENDROIDEAE 255 Appendix 1. Continued n 2n Source B. striata (Thunb.) Rchb.f. 16 32 32, 64 32, 76 32 M73, TK84 G88, GJ90 GJ90 GJ91, GJ96 TK84 M73, G84 G88 M73 M73, M77, G88, GJ94 G81, GJ90, GJ91, GJ96 M73, G84, G85 G88 M73, G85, GJ06 M73, G84, GJ91 GJ03 G88 M77 G85 M73, G85 M73 G81 M73, G84 G88, GJ90, GJ94 GJ96 M77 M73 M77, GJ91, GJ96 GJ91 M73 G88 G88 M74 M77 M73 M73, G85 M73, G85, G88 G85 GJ91 M73 M73, TK84, GJ96 GJ96 G81, GJ91 G85 M73 TK84 M73, G84, GJ94 G85 G84 M73, G85 M73, TK84 G85 G88, GJ91, GJ96 M73 GJ96 M73, GJ90, GJ91 GJ06 GJ91, GJ96 M73, GJ91 M73, G84, GJ91 M73, G84, GJ91 M73, GJ91 M73, G84, GJ91 M73, G84, GJ91, GJ94 G81 GJ91 B. striata (Thunb.) Rchb.f. (as B. striata var. albomarginata) Coelogyne barbata Griff. C. beccarii Rchb.f. C. corymbosa Lindl. C. cristata Lindl. C. fimbriata Lindl. C. flaccida Lindl. C. fragrans Schltr. C. fuscescens Lindl. C. longipes Lindl. C. macdonaldii F.Muell. & Kraenzl. C. macdonaldii (as C. lamellata) C. micranta Lindl. C. nitida Lindl. C. nitida (Wall. ex D.Don) Lindl. (as C. ochracea) C. ocultata Hook.f. 21 20 20 20 C. ovalis Lindl. 20 C. prolifera Lindl. C. prolifera Lindl. (as C. flavida) C. punctulata Lindl. C. speciosa (Blume) Lindl. (as Pleione speciosa) C. stricta (D.Don) Schltr. (as C. elata) C. viscosa Rchb.f. Dendrochilum longifolium Rchb.f. (as D. bartonii (Ridl.) Schltr.) D. uncatum Rchb.f. (as D. formosanum) Neogyna gardneriana (Lindl.) Rchb.f. (as Coelogyne gardneriana) Otochilus albus Lindl. O. fuscus Lindl. O. porrectus Lindl. Panisea demissa (D.Don) Pfitzer (as P. parviflora Rchb.) P. tricallosa Rolfe P. uniflora (Lindl.) Lindl. (as Coelogyne uniflora) Pholidota articulata Lindl. 20 20 19 + (0–2B) P. articulata Lindl. (as P. articulata var. griffithii Lindl.) P. articulata Lindl. (as P. griffithii Hook.) P. chinensis Lindl. P. imbricata Lindl. (as P. conchoidea) P. imbricata (Roxb.) Lindl. P. imbricata Lindl. (as P. calceata) P. pallida Lindl. P. protracta Hook.f. P. recurva Lindl. P. rubra Lindl. Pleione bulbocodioides (Franch.) Rolfe P. bulbocodioides (Franch.) Rolfe (as P. pogonioides) P. coronaria P.J.Cribb & C.Z.Tang P. formosana Hayata 20 20 40 20 20 20 + 1f 20 20 20 40 40 40 40 40 40 40 40 38 40 80 P. formosana Hayata (as P. priscei) P. forrestii Schltr. P. hookeriana (Lindl.) B. S. Williams Pleione humilis Lindl. P. limprichtii Schltr. P. maculata Lindl. P. praecox Lindl. P. scopulorum W.W.Sm. 38 80 20 40 38 30 21 20 20 20 20 20 + O-3B 40 40 40 40 40 38 40 40 20 20 20 20 20 20 20 20 + 0–3B 40 40 40 40 40 40 80 40 40 + 0–2B 40 38, 40, 42 40 40 40 + 0–1B 40, 80 40 40 40 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 TAXON 256 L. P. FELIX and M. GUERRA Appendix 1. Continued TAXON n P. yunnanensis (Rolfe) Rolfe Thunia alba Rchb.f. 20 ca. 40 Source 40 40 GJ91, GJ96 M73, GJ90, GJ94 G81 G85 M73 G81 G84 G85 G81 ca. 40 T. alba Rchb.f. (as T. marshalliana Rchb.f.) 20 38 42 40 T. alba (Lindl.) Rchb.f. (as Phaius albus) T. alba var. bracteata (Roxb.) N.Pearce & P.J.Cribb (as T. venosa Rolfe) TRIBE VANDEAE Subtribe Polystachyinae Polystachya adansoniae Rchb.f. P. affinis Lindl. P. albescens Summerh. P. albescens subsp. imbricata (Rolfe) Summerh. P. bancoensis Van der Burg P. bennettiana Rchb.f. (as P. stricta) P. caespitifica subsp. hollandii (L. Bolus) P.J.Cribb & Podz. (as P. holandii) P. caloglossa Rchb.f. P. campyloglossa Rolfe P. concreta (Jacq.) Garay & H.R.Sweet (as P. extinctoria) P. cooperi Summerh. P. coriscensis Rchb.f. P. cultriformis Lindl. P. dendrobiiflora Rchb.f. (as P. tayloriana) P. doggettii Rendle & Rolfe P. dolichophylla Schltr. P. estrellensis Rchb.f. P. eurygnatha Summerh. P. falax Kraenzl. P. foliosa (Hook.) Rchb.f. (as P. clavata) P. fulvilabia Schltr. P. fusiformis Lindl. P. galeata (Sw.) Rchb.f. P. isochiloides Summerh. P. laxiflora Lindl. P. mauritiana Spreng. (as P. tessellata) P. modesta Rchb.f. P. mukandaensis De Wild. P. nyanzoensis Rendle P. odorata Lindl. 20 40 P. polychaete Kraenzl. 40 40 40 ca. 40 40 40 40 40 ca. 80 80 40 40 38, 39 40 81 40 80 40 40 ca. 80 40 ca. 40 40 40 40 40 ca. 40 40 40 40, 80 40 40 ca. 20 P. pubescens Rchb.f. P. ramulosa Lindl. P. rhodoptera Rchb.f. ca. 120 40 40, 41 19 P. ruwenzoriensis Rendle P. stauroglossa Kraenzl. P. stuhlmanii Kraenzl. P. subulata Finet P. supfiana Schltr. P. transvaalensis Schltr. P. zambesiaca Rolfe (as P. hislopii) Polystachya sp. Subtribe Aeridinae Acampe ochracea (Lindl.) Hochr. A. pachyglossa Rchb.f. A. praemorsa (Roxb.) Blatter & McCann A. praemorsa (Roxb.) Blatt. & McCann (as A. papillosa) 40 40 40 40 40 40 80 40 18 18, 19 Acampe praemorsa (Roxb.) Blatter & McCann (as A. wightiana) 38 38 38 38 36 38 M73, G84 M73, G84 M73 TK84 G84 TK84 TK84 M73 TK84 TK84 TK84 M73 TK84 TK84 TK84 M73 TK84, PW TK84 TK84 TK84 TK84 TK84 M73, G84, TK84 TK84 M73, TK84 TK84 TK84 M73 M73 M73 TK84 M73, G84 TK84 TK84 M73, TK84 M73 TK84 TK84 TK84 TK84 TK84 M73 TK84 TK84 M73 TK84, G88 G85 TK84 M73 G81, GJ94 TK84 TK84 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 2n CHROMOSOMAL VARIABILITY OF EPIDENDROIDEAE 257 Appendix 1. Continued TAXON 19 19 19 19 19 19 19 A. A. A. A. A. maculosa Lindl. multiflora Roxb. multiflora Roxb. (as A. affine) odorata Lour. odorata Lour. (as A. suavissima) A. odorata Lour. (as A. odorata var. immaculata) A. ringens Fischer A. rosea Lodd. ex Lindl. & Paxton (as A. fieldingii) A. rosea Lodd. ex Lindl. & Paxton (as A. williamsii) Arachnis labrosa (Lindl. & Paxton) Rchb.f. (as Armodorum labrosum) Ascocentrum ampulaceum (Roxb.) Schltr. A. curvifolium (Lindl.) Scltr. A. himalaicum (Deb, Sengupta & Malick) Christenson (as Holcoglossum junceum) A. miniatum (Lindl.) Schltr. A. rubescens (Rolfe) P.F.Hunt (as Saccolabium rubescens) Biermannia bimaculata (King & Pantl.) King & Pantl. Calymmanthera major Schltr. C. paniculata J.J.Sm. Chilochista lunifera J.J.Sm. C. usneoidese (Don) Lindl. C. lunifera (Rchb.f.) J.J.Sm. (as Sarcochilus luniferum) Cleisostoma appendiculata (Hook.f.) (as Sarcanthus appendiculatus) C. discolor Lindl. (as Sarcanthus termissus) C. duplicilobum (J.J.Sm.) Garay (as Sarcanthus. carinatum) C. filiforme (Lindl.) Garay (as S. filiformis) C. fuerstenbergianum Kraenzl. (as Sarcanthus flagelliformis) C. paniculatum (Ker Gawler) Garay C. peninsularis (Dalz.) (as S. peninsularis) C. recemiferum (Lindl.) Garay C. recemiferum (Lindl.) Garay (as S. recemiferum) Cleisostoma racemiferum (Lindl.) Garay (as Sarcanthus palidus) C. recurvum (Hook.) ined. (as C. rostratum) C. striathum (Rchb.f.) Garay C. striatum (Rchb.f.) N. E. Br. (as Cleisostoma brevipes) C. strongyloides (Ridl.) Garay as (S. strogyloides) C. subulatum (Blume) as (S. subulatus) C. williamsonii (Rchb.f.) Garay (as S. williamsonii) Cleisostoma sp. as (Sarcanthus sp.) Cleisostomopsis eberhardtii (Finet) Seidenf. (as Saccolabium eberhardtii) Cottonia peduncularis (Lindl.) Rchb.f. C. peduncularis (Lindl.) Rchb.f. (as C. macrostachya) Diplocentrum congestusm Wight D. recurvum Lindl. Diploprora championii (Lindl.) Hook.f. (as D. uraiense) Drymoanthus adversus (Hook.f.) Dockrill D. flavus St.George & Molloy D. minimus (Schltr.) Garay Esmeralda clarkei Rchb.f. (as Arachnis clarkei) Euanthe sanderiana (Rchb.f.) Schltr. (as Vanda sanderiana) Gastrochilus calceolaris (Sm.) D.Don G. dasypogon (Lindl.) Ktze. G. dasypogon (as Saccolabium dasypogon) 19 19 2n Source 38 M73 G85 G81, TK84 G84 M73 TK84 TK84, G85 TK84 38 38 38 38 40 38 38 38 38 38 ca. 38 38 76 38 38 38 38 38 38 38 38, ca. 38 19–20 18 38 38 38 38 19 38 38 38 19 38 38 38 38 19 38 38 19, 20 36 19 19 38 38 38 19–20 19 19 19 19 38 40 38 38 16 76 38 38 42 38 38 38 38 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 TK84 TK84 M73, TK84, G84 M73, TK84 G85 M73,TK84, G85, GJ94 M73 TK84 TK84 TK84 G85 M73, G88 G88 M73 M73 GJ96 M73 TK84 G81 G88 G88 TK84 M73 M73 M73, TK84 M77 M73 M73 M73 GJ91 G81, G85 G88 G81 M73 GJ91 TK84 M77 M73 M73, G88 M77 TK84 TK84 G88, GJ90 G84 TK84, G88, GJ90 TK84 M74 D07 D07 D07 GJ91, GJ96 TK84 M73, GJ94 M73 TK84 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 A. praemorsa (Roxb.) Blatter & McCann (as Saccolabium papilosum) A. rigida Buch.-Ham. ex Sm.) P.F.Hunt A. rigida (Buch.-Ham. ex Sm.) P.F.Hunt (as A. longifolia) A. rigida (Buch.-Ham. ex Sm.) P.F.Hunt (as Aerides rigida) A. rigida (Buch.-Ham. ex Sm.) P.F.Hunt (as Saccolabium longifolium) Aerides crassifolia Par & Rchb.f. A. crispa Lindl. A. falcataLindl. & Paxt. A. houlettiana Rchb.f. A. lawrenceae Rchb.f. n 258 L. P. FELIX and M. GUERRA Appendix 1. Continued TAXON n G. G. G. G. 19 19 38 distichus (Lindl.) Kuntze (as Saccolabium distichum) inconspicuus (Hook.f.) Kuntze (as L. inconspicua Hook.f.) intermedius (Griff. ex Lindl.) Kuntze (as Saccolabium calceolare) japonicus Schltr. Papilionanthe tricuspidata (J.J.Sm.) Garay (as Vanda tricuspidata) P. uniflora (Lindl.) Garay (as A. longicornu) Papilionanthe vandarum (Rchb.f.) Garay (as Aerides vandarum) Paraphalaenopsis denevei (J.J.Sm.) A.D.Hawkes P. serpentilingua (J.J.Sm.) A.D.Hawkes Pelatantheria ctenoglossa Ridl. P. scolopendrifolia (Makino) Aver. (as Sarcanthus scolopendrifolium) Phalaenopsis amabilis Blume P. amabilis var. grandiflora P. amboinensis T.J.Sm. P. aphrodite Rchb.f. P. buyssoniana Rchb.f. (as Doritis bruysoniana) P. cornu-cervi Blume & Rchb.f. P. deliciosa Rchb.f. (as Doritis wightii) P. equestris Rchb.f. (=P. rosea) P. equestris (Schauer) Rchb.f. (as P. rosea) P. esmeralda Rchb.f. (=Doritis pulcherrima) P. fuscata Rchb.f. P. gigantea J.J.Sm. P. lindenii Loker P. luedemanniana Rchb.f. P. luedemanniana var. boxallii P. luedemanniana var. ochracea P. mannii Rchb.f. P. mannii Rchb.f. (as P. boxallii) P. mariae Burbidge P. parishii Rchb.f. P. parishii Rchb.f. (as P. decumbens) P. pulcherrima (Lindl.) J.J.Sm. (as Doritis pulcherrima) P. pulcherrima var. buyssoniana P. sanderiana Rchb.f. P. schilleriana Rchb.f. Source 40 38 30 M73 M73, G85, G88 M73 TK84 TK84 M74 34 19 19 38 38 38 38 40 20 19 18 19 19 19 19 19 19 19 19 19 19 19 19 19 19 19 19 19 19 19 38 38 38 38 38 38, ca. 38 38 38 38 38 38 38 38 40 76 38 38 38 38 38 38 38 38 38 38 76 38 38 38 38 38 38 38 38 38 38 38 38 38 38 38 76 38 38 TK84 M73 G88 TK84 TK84 M73, TK84 TK84 M73 M73 TK84, G88 TK84, G88 G88 TK84 M73, G81, G88 M73, G88 G85 G85 M73, TK84 TK84 G81, TK84, G88 M73 TK84 G88 TK84, G85 GJ91, GJ96 TK84 M73, G85 G85 G84 G84 M73 TK84 M73, G81, TK84 TK84 M73, G81 M73, G81 TK84 G81 M73 M73, G81 G81 TK84 G81 G81 TK84 M73, G81 G81 G81 M73, G81 TK84 G81 G81 TK84 TK84 TK84 M73, G81 G81, TK84 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 G. japonicus (Makino) Schltr. (as G. somai) G. maculatus Kuntze G. matsuran (Makino) Schltr. G. pseudodistichus (King & Pantl.) Schltr. (as Saccolabium pseudodistichus) Gunnarella robertsii (Schltr.) Senghas (as Chamaeanthus robertsii) Holcoglossum amesianum (Rchb.f.) Christenson (as Vanda amesiana) H. kimballianum (Rchb.f.) Garay (as V. kimballiana) Hygrochilus parishii (Veitch & Rchb.f.) Pfitzer (as Vanda parishii) Luisia boninensis Schltr. L. brachystachys (Lindl.) Blume L. liukiuensis Schltr. L. macrantha Blatt. & McC. L. tenuifolia Blume L. tenuifolia (as L. tenuifolia var. evangelinae) L. teres Blume (= L. teretifolia) L. trichorhiza Blume L. tristis (G. Forst.) Hook.f. (as L. teretifolia) L. volucris Lindl. Micropera mannii (Hook.f.) Tang & F.T.Wang (as Sarcochilus manni) M. rostrata (Roxb.) N.P.Balakr. (as Sarcochilus purpureus) Neofinetia falcata (Thunb.) Hu Ornithochilus difformis (Wall. ex Lindl.) Schltr. (as Ornithochilus fuscus Wall.) Papilionanthe biswasiana (Ghose & Mukerjee) Garay (as Aerides biswasianum) P. hookeriana (Rchb.f.) Schltr. (as Vanda hookeriana) P. teres (Roxb.) Schltr. P. teres (Roxb.) Schltr. (as Vanda teres) 2n CHROMOSOMAL VARIABILITY OF EPIDENDROIDEAE 259 Appendix 1. Continued TAXON 19 19 19 19 19 19 19 2n Source 38 38 40 38 38 38 38 38 38 38 38 38 38 G81 M73, G81 M73, G81, GJ94 G81 G81 G88 TK84 G85 TK84 TK84 TK84, GJ96 TK84 TK84 TK84 TK84 TK84 M73, TK84 M73 TK84 TK84 TK84 G81, TK84, G88, GJ03 G88 G88 TK84 M73 G88 GJ96 D07 TK84 M73 G88 G81, M73 M73 G88 TK84, G88, GJ94 GJ90 GJ90 M73 TK84 TK84 TK84 G88 TK84 TK84 M73 G88 GJ96 G88 TK84 G81 M73 M73 M73 TK84 M74 G88 G84 M73 TK84 TK84, G85 TK84, G85, GJ91, GJ96 38 38 38 38 38 19–20 19 38 38 38 38 38 38 38 38 76 38 20 19 19 38 38 38 38 38 38 38 38 38 38 19 40 56 38 40 38 38 38 38 38 19 38 38 ca. 38 19–20 30 38 19 19 19 19 38 38 38 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 P. speciosa Rchb.f. P. stuartniana Rchb.f. P. taenialis (Lindl.) Christenson & Pradhan (as Doritis taenialis) P. tetraspsis Rchb.f. P. violacea Teijsm. & Binn. Pomatocalpa marsupiale (Kraenzl.) J.J.Sm. P. kunstleri (Hook.f.) J.J.Sm. (as Sarcanthus kunstleri) P. spicatum Breda (as Cleisostoma mannii) Pteroceras longicalcareum (Ames & Rolfe) Garay (as Sarcochilus longicalcareum) P.teres (Blume) Holttum (as Sarcochilus palawanensis) Renanthera coccinea Lour. R. elongata Lindl. R. histrionica Rchb.f. R. imschootiana Rolfe R. matutina Lindl. R. monachica Ames R. storiei Rchb.f. Rhynchostylis coelestis Rchb.f. R. gigantea Ridl. R. gigantea (as Saccolabium albo-lineatum) R. gigantea (as V. densiflora) R. retusa (L.) Blume Robiquetia bertoldii (Rchb.f.) Scltr. R. bertoldii (as R. mimus (Rchb.f.) Garay) R. mooreana (Rolfe) J.J.Sm. R. spathulata (Blume) J.J.Sm. R. succisa (Lindl.) Seidenf. & Garay (as R. paniculata) R. wassellii Dockr. Sarcochilus fitzgeraldii F.Muell. S. hartmannii F.Muell. Schoenorchis gemmata (Lindl.) J.J.Sm. (as Cleisostoma gemmatum) Sedirea japonica (Rchb.f.) Garay & H.R.Sweet (as Aerides japonica) Schoenorchis gemmata (Lindl.) Sm. S. micrantha Reinw. ex Blume (as Ascocentrum micranthum) Seidenfadenia mitrata (Rchb.f.) Garay (as Aerides mitrata) Smitinandia micrantha (Lindl.) Holttum S. micrantha (Lindl.) Holttum (as Cleisostoma micranthum) Smithsonia maculata (Dalzell) C.J.Saldanha (as Gastrochilus maculatus) S. viridiflora (Dalzell) C.J.Saldanha (as Gastrochilus dalzelianus) Staurochilus fasciatus (Rchb.f.) Ridl. (as Trichoglottis fasciata) Stereochilus erinaceus (Rchb.f.) Garay (as Sarcanthus erinaceus) Taeniophyllum crepidiforme King. & Pantl. T. elmeri Ames T. fasciola (G.Forst.) Rchb.f. T. glandulosum Blume (as T. aphyllum) T. philippinensis Rchb.f. Thrixspermum acuminatissimum (Blume) Rchb.f. T. carinatifolium Ridl. T. centipeda Lour. T. graeffei Rchb.f. T. japonicum Rchb.f. (as Sarcochilus japonicum) T. pygmaeum (King & Pantl.) Holttum (as Sarcochilus pigmaeum) Trichoglottis cirrhifera Teijsm. & Binn. T. rosea (Lindl.) Ames T. tenera (Lindl.) Rchb.f. (as Saccolabium tenerum) T. triflora (Guillaumin) Garay & Seidenf. (as Saccolabium triflorum) . kotoense Yamam. (as Saccolabium kotoense) T. rhopalorrhachis (Rchb.f.) J.J.Wood (as Saccolabium rhopalorrhachis) Uncifera obtusifolia Lindl. U. obtusifolia (as Saccolabium obtusifolia) Uncifera sp. Vanda alpina Lindl. V. coerulea Griffith n 260 L. P. FELIX and M. GUERRA Appendix 1. Continued n 2n Source V. coerulescens Griff. 19 38 19 76 38, 76 ca. 38, 38 38 38, 76 76 38 38 38 38 38 38 76 114, 115 38 38 38 38 38 38 38 38 38 38 38 57 38 38 38 38 M73, TK84, GJ91, GJ96 TK84 GJ91, GJ96 M73, G81, TK84, G85, GJ00 GJ00 TK84 TK84 GJ91, GJ96 TK84, GJ91, GJ96 TK84 TK84 TK84 TK84 GJ91, GJ96 TK84, G88 TK84 M73, G85, G88 GJ91, GJ96 TK84 G81, TK84, G85 M73, GJ90 M77 G81, G88, G85, GJ94 TK84 TK84 TK84 GJ96 GJ96 M73 TK84 G84, GJ06 GJ96 38 48 38 50 38 38 38 38 42–48 38 50 92 40 38 M73 G85 M73 G50 G84 M73, G85 M73 G85 TK84 G85 G84 G85 TK84 M73 38 63 42 46 50 38 38 40 ca. 38 42 50 46 ca. 76 38 TK84, GJ91 M73 G85 G85 TK84 M73 G85 TK84 M73 M73 G85 G85 M73 M73 V. concolor Blume V. cristata Lindl. V. V. V. V. V. V. V. V. V. V. dearei Rchb.f. denisoniana Benson & Rchb.f. denisoniana (as V. denisisoniana var. hebraica Rchb. f. flabellata (Rolfe ex Downie) Christenson (as Aerides flabellata) lamellata Lindl. lamellata var. boxallii Rchb.f. lilacina Teijsm. & Binn. (as V. laotica) luzonica Loher ex Rolfe pumila Hook.f. spatulata Spreng. V. stangeana Rchb.f. V. subconcolor Tang & Wang V. tessellata (Roxb.) Hook. V. tessellata (Roxb.) Hook. ex G.Don (as V. roxburghii) V. testacea (Lindl.) Rchb.f. V. testacea (Lindl.) Rchb.f. (as A. wrightiana) V. testacea (Lindl.) Rchb.f. (as V. parviflora) V. tricolor Lindl. V. tricolor Lindl. (as V. suavis) V. tricolor var. suavis Vanda sp. Vanda sp. Vandopisis gigantea (Lindl.) Pfitzer V. lisochiloides Pfitzer V. undulata (Lindl.) J.E.Sm. V. undulata (Lindl.) J.J.Sm. as Stauropsis undulatum) Subtribe Angraecinae Angraecum aichlerianum Kraenzl. A. aporoides Summerh. A. arachnites Schltr. A. bancoense Burg A. birrimense Rolfe A. calceolus Thouars A. calceolus Thouars (as A. onocentrum) A. caricifolium Perrier A. compressicaule H.Perrier A. cultriforme Summerh. A. distichum Lindl. A. doratophyllum Summerh. A. eburneum Bory Angraecum eburneum subsp. giryamae (Rendle) Senghas & P.J.Cribb (as A. giryamae) A. eichlerianum Kraenzl. A. erectum Summerh. A. A. A. A. gabonens Summerh. guillauminii H.Perrier infundibulare Lindl. leonis (Rchb.f.) Vietch A. A. A. A. A. A. moandense De Wild. (as A. chevaleri) multinominatum Rendle podochiloides Schltr. pungens Schltr. sacciferum Lindl. scottianum Rchb.f. 19 19 38 19 19 19 19 19 19 19 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 TAXON CHROMOSOMAL VARIABILITY OF EPIDENDROIDEAE 261 Appendix 1. Continued TAXON n Source A. sesquipedale Thouars A. subulatum Lindl. Calyptrochilum christianum (Rchb.f.) Summerh. C. emarginatum (Sw.) Schltr. Campylocentrum amazonicum Cogn. C. crassirhizum Hoehne C. neglectum (Rchb.f. & Warm.) Cogn. C. pernambucense Hoehne 38 50 38 38 80 38 38 38 G85 G84 G84 G84 PW PW DA09 PW Cryptopus elatus (Thouars) Lindl. 95 76 44 M73 G85 G88 38–40 46 38 M73 G84 TK84 50 50 50 46 ca. 50, ca. 50 51 50 52 50, ca. 50 M73 TK84 G85 G85 M73, G85 M73 G85 G85 M73 42 42 G85 M73, G85 TK84 G85 M73, G85 M73 Dendrophylax monteverdi (Rchb.f.) Ackerman & Nir (as Campylocentrum moteverdii) Jumellea walleri (Rolfe) la Croix (as Jumellea filicornoides) Listrostachys pertusa (Lindl.) Rchb.f. Oeoniella stachys Schltr. Subtribe Aerangidinae Aerangis biloba (Lindl.) Schltr. A. biloba (Lindl.) Schltr. (as Angraecum bilobum) A. brachycarpa (A.Rich.) Durand & Schinz A. calantha (Schltr.) Schltr. A. citrata (Thouars) Schltr. A. collum-cygni Summerh. (as A. compta) A. cryptodon Rchb.f. A. kirkii (Rchb.f.) Schltr. A. kotschyana (Rchb.f.) Schltr. A. luteoalba var. rhodosticta (Kraenzl.) J. Stewart Aerangis luteoalba var. rhodosticta (Kraenzl.) J.Stewart (as A. rhodosticta) A. modesta (Hook.f.) Schltr. (as Angraecum sanderianum) A. somalensis (Schltr.) Schltr. A. ugandensis Summerh. Aerangis sp. Ancistrorhynchus capitatus (Lindl.) Summerh. A. clandestinus (Lindl.) Schltr. A. crystalensis Crib & Van der Lan A. metteniae (Kraenzl.) Summerh. A. ovatus Summerh. Ancistrorhynchus recurvus Finet A. straussii (Schltr.) Schltr. A. tenuicaulis Summerh. Ancistrorhynchus sp. Angraecopsis amaniensis Summerh. A. breviloba Summerh. A. gracillima (Rolfe) Summerh. A. pusilla Summerh. A. trifurcata Schltr. Bolusiella batesii (Rolfe) Schltr. Chamaeangis odoratissima (Rchb.f.) Schltr. C. vesicata (Lindl.) Schltr. Cribbia brachyceras (Summerh.) Senghas (as Rangaeris brachyceras) Cyrtorchis arcuata (Lindl.) Schltr. C. arcuata subsp. arcuata (as C. arcuata subsp. variabilis) C. arcuata subsp. whytei (Rolfe) Schltr. C. aschersonii (Kraenzl.) Schltr. C. brownii (Rolfe) Schltr. C. chailluana (Hook.f.) Schltr. C. hamata (Rolfe) Schltr. C. monteiroae (Rchb.f.) Schltr. C. praetermissa Summerh. C. ringeus (Rchb.f.) Summerh. Cyrtochis spp. Diaphananthe fragrantissima (Lindl.) Schltr. 25 25 54 ca. 50, 50 50, ca. 50, 200 72 48 48 48 48 ca. 50 48 48 96 48 50 50 50 50 50 50 95–100 50 ca. 150 46 138 46 46 22–23 46 46 46, 92 46 46, 50 50 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 G85 M73, G85 GJ91 G85 G85 M73 GK91 GJ91 G85 G88 M73 G85 G85 GJ91 G84 M73, G85 M73 M73, G85 M73 G85, GJ91 G85 G84 G85 TK84 G84 G85 G85 G85 M73 G85 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 2n 262 L. P. FELIX and M. GUERRA Appendix 1. Continued TAXON n 54II Rhipidoglossum brevifolium Summerh. (as Diaphananthe brevifolia) R. cuneatum (Summerh.) Garay (as Diaphananthe cuneata) R. densiflorum Summerh. (as Diaphananthe densiflora) R. pulchellum (Summerh.) Garay (as Diaphananthe pulchella) R. rutilum (Rchb.f.) Schltr. (as Diaphananthe rutila) R. xanthopollinium (Rchb.f.) Schltr. (as Diaphananthe xanthopollinia) Solenangis clavata (Rolfe) Schltr. S. scandens (Schltr.) Schltr. Sphyrarhynchus achiliebenii Mansf. Tridactyle anthomaniaca (Rchb.f.) Summerh. T. tridactylites (Rolfe) Schltr. Tridactyle sp. UMPLACED SUBTRIBES Subtribe Agrostophyllinae Agrostophyllum brevipes King & Pantl. A. callosum Rchb.f. A. myrianthum King & Pantl A. planicaule (Wall. ex Lindl.) Rchb.f. (as A. khasianum) A. uniflorum Schltr. Earina aestivalis Cheeseman E. autumnalis (G.Forst.) Hook.f. E. mucronata Lindl. Glossorhyncha chlorantha van Royen G. macdonaldii Schltr. 20 20 20 20 Subtribe Dendrobiinae Bulbophyllum acuminatum (Ridl.) Ridl. (as Cirrhopetalum acuminatum) B. aff. acuminatum B. acutibracteatum var. rubrobrunneopapillosum (De Wild.) J.J.Verm. (as B. fuscoides) B. acutissepalum De Willd. B. adenopetalum Lindl. B. aeolium Ames B. andersonii J.J.Sm. (as C. andersonii) B. affini Lindl. 20 Source 50 50 ca. 50 50 46 50 48 48 48 48 48 48 48 48 48 48 48 48 48 48 50 48 46 92 ca. 100 ca. 108 46 50 ca. 50 50 50 100 50 50 50 50 ca. 100 ca. 100 ca. 50 G85, GJ91 M.73 M73 GJ91 GJ91 M73, GJ91 G84 G84 G84 G84, G85, G88, GJ91 G84 G84 G84 G84 G84 G85 G88 G84 G84 G84, G88 G85 G85 GJ91 GJ91 M73 M73 G84 G85 M73 M73 G85 M73, G85 G85 G84 G84 GJ91 M73 M73 M73 40 38 40 40 40 40 40 M73 M73, G85 TK84 M73 M73, G85 G88 D07 D07 D07 G88 GJ06 38 38 ca. 38 TK84 G88 M73 38 38 40 38 36 G88 M73 TK84 TK84 TK84 M73 40 38 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 D. pellucida (Lindl.) Schltr. D. plehniana (Schltr.) Schltr. Diaphananthe sp. Diaphananthe sp. Eggelingia gabonensis Cribb & Van der Lan Eurychone rothschildiana (O’ Br.) Schltr. Microcoelia bispiculata L.Jonsson M. bulbocalcarata L.Jonsson M. caespitosa (Rolfe) Summerh. M. exilis Lindl. M. gilpinae (Rchb.f. & S.Moore) Summerh. M. globulosa (Hochst.) L.Jonsson M. hirschbergii Summerh. M. koeleri (Schltr.) Summerh. M. macrantha (H.Perr.) Summerh. M. macrorhynchia (Schltr.) Summerh. M. megalorrhyza (Rchb.f.) Summerh. M. obovata Summerh. M. physophora (Rchb.f.) Summerh. M. stolzii (Schltr.) Summerh. Microterangis hildebrandtii (Rchb.f.) Senghas (as Chameangis hildebrandtii) Mystacidium capense (L.f.) Schltr. Podangis dactyloceras (Rchb.f.) Schltr. Rangaeris amaniensis (Kraenzl.) Summerh. R. musicola (Rchb.f.) Summerh. R. rhipsalisocia (Rchb.f.) Summerh. 2n CHROMOSOMAL VARIABILITY OF EPIDENDROIDEAE 263 Appendix 1. Continued TAXON alagense Ames ambrosia (Hance) Schltr. andersonii Kurz. anteniferum Rchb.f. apodum Hook.f. apodum Hook.f. (as B. ebulbum) argyropus (Endl.) Rchb.f. (as Adelopetalum argyropus) aurantiacum F.Muell. auratum Rchb.f. aureolabellum T.P.Lin (as C. gracillimum) baileyi F.Muell. barbigerum Lindl. biantennatum Schltr. biflorum Teijsm. & Binn., bisetum Lindl. boninense J.J.Sm. (as C. boninense) braccatum Rchb.f. caespitosum Thou (as B. Caespitosum) canlanoense Ames caulflorum Hook.f. careyanum Spreng. cauliflorum Hook.f. cerinum Schltr. clandestinum Lindl. (as B. sparsifolium) clavatum Thouars cochleatum var. tenuicaule (Lindl.) J.J.Verm. (as B. tenuicaule) cocoinum Bateman ex Lindl. congolanum Schltr. cornutum Ridl. cornutum (as C. cornutum) crassifolium Thouars crassipes Hook.f. cribbianum Toscano cumingii (Lindl.) Rchb.f. cumingii (Lindl.) Rchb.f. (as C. cumingii) cumingii (Lindl.) Rchb.f. (as C. stramineum) cylindraceum Lindl. B. daloaense Cribb & Perez-Vera B. denisii J.J.Wood. B. densiflorum Rolfe B. drymoglossum Maxim. B. elassonotum Summerh. B. elliottii Rolfe (as B. malawiense) Bulbophyllum elisae F.Muell. B. emiliorum Ames & Quisumb. B. encephalodes Summerh. B. eublepharum Rchb.f. B. evrardii Gagnep. B. exiguum F.Muell. B. falcatum (Lindl.) Rchb.f. B. falcatum var. velutinum (Lindl.) J.J.Verm. (as B. melanorhachis) B. falcatum var. velutinum (Lindl.) J.J.Verm. (as B. rhizophorae) B. fascinator (Rolfe) Rolfe B. flavidum Lindl. B. B. B. B. B. fletcherianum Rolfe formosum Schltr. fritillariflorum J.J.Sm. frostii Summerh. gamblei Hook.f. B. gibbosum (Blume) Lindl. B. grandiflorum Blume 2n Source 40 38 38 40 38 38 36 38 TK84 G88, GJ91, GJ96 G85 TK84 M73 G85 D07 G85 TK84 TK84 G85 M73, G85 G85 G88 G84 TK84 TK84 TK84 TK84 TK84 M73, TK84 G81 G88 G85 G88 TK84 M73, G85 M73, G85 TK84 TK84 G88 G88 PW TK84 TK84 TK84 G84 M73 G85 G88 M73 TK84 G85 G85 G88 TK84 G85 M73 TK84 G88 M73 G88 G88 G88 M73 G84 G88 G88 G88 M73 M73 G88 G85 M73, G85 20 ca. 38 38 38 38 38 40 20 20 20 40 21 19 + 0–1B 21 38 19 38 38 38 38 38 38 38 19 38 38 38 40 38 38–40 19 20 38 38 19 40 38 38 38 40 38 19 38–42 ca. 38 38 38 38 38 40 38 38 38 38 38 20 19 38 38 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. n 264 L. P. FELIX and M. GUERRA Appendix 1. Continued TAXON n B. grandiflorum Blume B. griffithii Rchb.f. B. gymnopus Hook.f. 19 19 imbricatum Lindl. imbricatum Lindl. (as B. linderi) inconspicuum Maxim. intertextum Lindl. intricatum Seidenf. ipanemense Hoehne japonicum Makino khasyanum Griff. lasiochilum Par. & Rchb.f.(as Cirrhopetalum lasiochilum) leopardinum Lindl. leptanthum Hook.f. lepidum (Blume) J.J.Sm. lepidum (as C. lepidum) levanae var. giganteum Quisumb. & C. Schweinf. aff. levatii lobbi Lindl. logiflorum Thouars logiflorum Thouars (as C. longiflorum) longiscapum Rolfe lupulinum Lindl. B. B. B. B. B. B. B. B. B. B. B. B. macraei (Lindl.) Rchb.f. macranthum Lindl. maculosum Ames (as C. maculosum) mahonii Rolfe makoyanum Rchb.f. mandibulare Rchb.f. mastersianum J.J.Sm. (as C. mastersianum) maximum (Lindl.) Rchb.f. maximum (Lindl.) Rchb.f. (as B. oxypterum) medusae (Lindl.) Rchb.f. miniatum Hort. ex F. Moore minutipetalum Schltr. B. morphologorum Kraenzl. B. mundulum (W. Bull) J.J.Sm. (as C. mundulum) B. mysorense (Rolfe) J.J.Sm. (as C. mysorense) B. neilgherense Wight B. nutans Thou. B. odoratissimum (Sm.) Lindl. B. odoratum var. odoratum (as B. elatius) B. oreonastes Rchb.f. B. oreonastes Rchb.f. (as B. zenkerianum) Bulbophyllum ornatissimum J.J.Sm. (as C. ornatissimum) B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. orthoglossum Kraenzl. oxychilum Schltr. (as B. buntingii) pallidiflorum Schltr. parvulum Lindl. (as C. parvulum) patens King pahudii (de Vriese) Rchb.f. (as B. virescens) penicillium Par. & Rchb.f. peninsulare Seidenf. phalaenopsis J.J.Sm. picturatum Rchb.f. picturatum (as C. picturatum) pipio Rchb.f. polyrhizum Lindl. protractum Hook.f. pulchrum (N.E.Br.) J.J.Sm. Source 38 M73 G81 M77 38 38 38 38 38 38 40 M73, G84 G88 M73 M73, G85 G88 G85 M73 G84 TK84 M73 G81 G85 TK84 TK84 20 38 19 19 38 38 60 19 39 38 38–40 38 38 40 38 38 38 38 38–40 38 38 38 38 38 38 38 36 38 38 ca. 38 40 38 38 38 38 38 38–40 ca. 38 38 38 38 19 57 38 19 38 38 38 48 38 19 19 38 38 M73 G85 TK84 G88 M73 M73 G88 G85 TK84 G85 TK84 G88 TK84 G85 G85 G85 G85 M73 GJ96 G88 TK84 TK84 TK84 M73 TK84, G88 M73 G85 G85 TK84 TK84 G85 G88 G88 TK84 G85 M73 G81 G88 G85 G85 TK84 G84 M73 G84, G85 G85 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. 2n CHROMOSOMAL VARIABILITY OF EPIDENDROIDEAE 265 Appendix 1. Continued TAXON pulchrum (as C. pulchrum) purpureorhachis (De Willd) Schltr. raui Arora reflexiflorum H.Perrier refractum Rchb.f. refractum Rchb.f. (as C. refractum) reptans Lindl. retusiusculum Rchb.f. retusiusculum Rchb.f. as B. micholitzii) rigidum King & Pantl. robustum Rolfe (as C. robustum) rothchildianum (O’Brien) J.J.Sm. roxburghii (Lindl.) Rchb.f. rufinum Rchb.f. rugosibulbon Summerh. saltatorium Lindl. saltatorium var. calamarium (Lindl.) J.J.Verm. (as B. calamarium) saltatorium var. albociliatum (Finet) J.J.Verm. (as B. distans) saltatorium var. albociliatum (Finet) J.J.Verm. (as B. nudiscapum) sanderianum Rolfe sandersonii (Oliv.) Rchb.f. saurocephalum Rchb.f. schinzianum Kraenzl. schiazianum Kraenzl. (as B. phaepogon) secundum Hook.f. B. sociale Rolfe B. stenobulbon E.C.Parish & Rchb.f. B. stenobulbon E.C.Parish & Rchb.f. (as B. clarkeanum) B. sterile (Lam.) Suresh (as C. caudatum) B. striatum Rchb.f. B. tentaculgerum Rchb.f. B. tetragonum Lindl. (as B. wrightii) B. trachyantum Kraenzl. B. tricanaliferum J.J.Sm. B. tridentatum Kraenzl. B. trimeni (Hook.f.) J.J.Sm. B. triste Rchb.f. B. tseanum Hu & Barr. B. tuberculatum Colenso (as Adenopetalum tuberculatum) B. umbellatum Lindl. B. umbellatum Lindl. (as C. umbellatum) B. unicaudatum Schltr. B. unifoliatum De Wild. B. vagans Ames & Rolfe B. vaginatum (Lindl.) Rchb.f. (as C. vaginatum) B. velutinum (Lindl.) Rchb.f. 19 19 + 0–1B 19 19 2n Source 38 38 40 38 38 TK84 G85 TK84, G88, GJ94 TK84 G88 G88, GJ94 G81, G88, GJ94 G84 M73, G85 G85 TK84 G88 G88 G88 G88 G85 M73, G85 G85 G85 PW G85 TK84 G85 M73, G85 M73 TK84 M73 G88 M73 TK84 M73 G84 G84 G85 G88 G88 G88 M73, G81 G88 38 38 38 38 38 38 38 38 38 38 38 38 38 38 20 38 38 19 20 38 38 19 19 19 38 38 38 38 38 38 19 39 38 38 38 38 38 95 38 19 40 Bulbophyllum viridiflorum Lindl. B. viridiflorum (as C. viridiflorum) B. wallichii Rchb.f. (as C. wallichii) B. wallichii Rchb.f. (as B. refractoides) B. wendlandianum (Kraenzl.) J.J.Sm. B. winkleri Schltr. B. zenkerianum Kraenzl. Bulbophyllum sp. Bulbophyllum sp. Bulbophyllum sp. Bulbophyllum sp nv. Bulbophyllum sp nv. Dendrobium acinaciforme Roxb. D. acerosum Lindl. D. acuminatissimum Lindl. 19 + 1B 19 + 0–3B 18 19 19 57 38 38 40 38 80 38 + Bs 38 38 38 40 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 D07 TK84, GJ94 TK84, G88 G88 G85 G85 TK84, G88 M73 G84 G88 TK84, GJ94 TK84 M73 G88 G84 G85 TK84 PW PW G88 G88 G81, G84, G88 GJ90 G84, G88 TK84 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. B. n 266 L. P. FELIX and M. GUERRA Appendix 1. Continued TAXON n D. acuminatum (Rolfe) Kraenzl. (= D. lyonii) D. adae F.M.Bayley D. aduncum Wall. ex Lindl. D. aemulum R.Br. D. agrostophyllum F.Muell. alaticaulinum Royen albayense Ames albosanguineum Lindl. & Paxton anosmum Lindl. (as D. leucorhodum) anosmum Lindl. (as D. superbum) antennatum Lindl. (as D. dalbertisii) aphyllum (Roxb.) C.E.C.Fisch. (as D. macrostachyum) aphyllum (Roxb.) C.E.C.Fisch. (as D. pierardii) atroviolaceum Rolfe aurantiroseum P.Royen ex T.M.Reeve baileyi F.Muell. barbatulum Lindl. bellatullum Rolfe bensoniae Rchb.f. bicallosum Ridl. bicameratum Lindl. D. D. D. D. D. D. D. D. D. D. D. D. bifalce Lindl. biflorum (Forst.) Sw. bigibum Lindl. bigibbum Lindl. (as D. phalaenopsis) bilobum Lindl. brymerianum Rchb.f. bulbophylloides Schltr. bullenianum Rchb.f. bullenianum Rchb.f. (as D. topaziacum) calcaratum A.Rich. camaridiorum Rchb.f. canaliculatum R.Br. capilipes Rchb.f. capituliflorum Rolfe capra J.J.Sm. cariniferum Rchb.f. catenatum Lindl. (as D. tosaense) D. cathcartii Hook.f. D. chameleon Ames D. chrysanthum Wall. D. D. D. D. 19 + 0–4B 19 19 19 19 D. candidum Wall. D. D. D. D. D. 19 Source 38 40 38 38 40 38 38 40 38 40 40 38 38 38 38 38 38 76 38 38 38 38 40 38 TK84 GJ90 G88 G85, GJ90 G88, GJ90 G84, GJ90 GJ90 G84 G85, G88 TK84 G88, GJ90 TK84 G84, GJ90 M73 M73, M77 M73, G84, G85, GJ90 M73, G88 G88 G85 G84, GJ90 G85 G88, GJ90 TK84 G81, G85, G88, GJ90, GJ94 G85 G85 TK84, G84, GJ90 TK84, G84, GJ90 G85 TK84, GJ90 G88 M73, GJ90 TK84 G85 G85 M73, GJ90 G84 G81, TK84, G84, G88, GJ90 TK84 G85, GJ90 G84 M73 M73, G84 TK84 M73 G85 G81, TK84, G88, GJ94 G81, TK84, G85, GJ90 GJ06 GJ06 M73, G81, G85, G88 G88, GJ90 M73, TK84, G84 G88, GJ90 TK84 GJ90 G85, GJ90 G84, GJ90 TK84 G85, G88 G88 TK84, G84, GJ90 38 38 38 38 38 38 38 38 38 38 38 38 38 38 38 38 19 19 38 38 40 19 20 19 chryseum Rolfe chryseum Rolfe (as D. clavatum) chryseum Rolfe (as D. denneanum) chrysotoxum Lindl. Dendrobium chrysocrepis E.C.Parish & Rchb.f. ex Hook.f. D. clavator Ridl. D. coelogyne Rchb.f. D. compactum Rolfe D. compressum Lindl. (as D. platycaulon) D. connatum Schltr. D. crassifolium Schltr. D. crassinodes Benson & Rchb.f. 19 19 19 20 38 40 38 76 38 38 38 38 ca. 76 38 40 40 38 38 38 38 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 20 D. D. D. D. D. D. D. D. D. D. D. D. D. D. D. D. 2n CHROMOSOMAL VARIABILITY OF EPIDENDROIDEAE 267 Appendix 1. Continued TAXON n D. crepidatum Lindl. D. crispilinguum Cribb. D. cruentum Rchb.f. crumenatum Sw. crumenatum Sw. (as D. kwashotense) crystallinum Rchb.f. chryseum Rolfe (as D. flaviflorum) cruttwellii T.M.Reeve cucumerinum Maclay ex Lindl. cumulatum Lindl. cunninghamii Lindl. cunninghamii Lindl. (as Winika cunninghamii) cuthbertsonii F.Muell. cuthbertsonii F.Muell. (as D. sophronites) cyanocentrum Shltr. cymbidioides (Blume) Lindl. delacourii Guill. (= D. ciliatum) 20 D. ¥ delicatum (F.M.Bailey) F.M.Bailey D. denneanum ¥ D. moschatum (Buch.-Ham.) Sw. D. densiflorum Wall. 20 + (0–2B) D. denudans D.Don D. devonianum Paxton D. dicuphum F.Muell. D. D. D. D. D. D. D. D. D. D. D. D. D. D. dichaeoides Schltr. dilonianum Hawkes & Helter distichum Rchb.f. dixanthum Rchb.f. draconis Rchb.f. engae Reeve epiphedum Lindl. equitans Kraenzl. equitans Kraenzl. (as D. batanense) falconeri Hook.f. farmerii Paxt. farmerii var. aureoflava Hook.f. fellowsii F.Muell. (as D. bairdianum) fimbriatum Hook. D. D. D. D. fimbriatum Hook. (as D. normale) findlayanum Par. & Rchb.f. finetianum Schltr. finisterrae Schltr. 20 19 19 formosanum Roxb. ex Lindl. friedericksianum Rchb.f. gibsoni Paxton goldfinchii F.Muell. goldschmidtianum Kraenzl. (as D. miyakei) goldschmidtianum Kraenzl. (as D. victoriae-reginae var. miyakei) D. gordonii S.Moore Dendrobium gouldii Rchb.f. D. gracilicaule F.Muell. D. gracilicaule var. howneanum Miden 38 38 40 38 + 1B 38, 38 + 0–2B 38 38 38 38 38 40 38 40 76 38 38 40 38 40 38 44 38 40, 40 + 1–3f 38 38 40 38 + 2B 40 38 38 38 + 2B 40 38 38 38 38 38 M73, TK84, M77, G88 G85 M73, TK84 G81 M73, TK84, G84, G85 M74 M73, G88 M74 G85 G84, GJ90 G85 G85 D07 G88 G84, GJ90 G88 G84, GJ90 M73 G84, G88 G85, G88 GJ06 G81, G85, G88 M73, G81, TK84, G84, G85, GJ90 G81, G84, GJ90 M77, TK84, G88 TK84 G84, GJ90 G88 G88 M73, G84, GJ90 M73, G84, G85, GJ90 M73, G85 G85, G88, GJ90 G81 G84, GJ90 G85 G84, G88, GJ90 M73, M77, G84 M73, G85 G88 M73, G85, G88 G81, TK84, G88, GJ90, GJ94 G81, GJ90 M73, G84, JG90 GJ96 G85 G84, GJ90 TK84 G88 G85 G84 M73, M77, G84, GJ90 M73, G84, G85, GJ90 G81, G88, GJ90 G85 M74, GJ90 38 38 38 38 TK84 M73, G85, GJ90 G85 TK84 40 38 38 39 38 38 38 40 + 2–4f 38 36 20 19 20 19 D. flamula Schltr. D. fleckeri Rupp. & C.T.White D. forbesii Ridl. D. D. D. D. D. D. Source 19 19 38 38 38 40 40 36 40 38, 38 + 0–2B © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 D. D. D. D. D. D. D. D. D. D. D. D. D. D. 2n 268 L. P. FELIX and M. GUERRA Appendix 1. Continued TAXON n gratiosissimum Rchb.f. griffithianum Lindl. guerreroi Ames & Quisumb. haemoglossum Thwaites (as bambusiaefolium) hancokii Rolfe heishanaense Hayata helix Cribb hellwigianum Kraenzl. hendersonii A.D.Hawkes & A.H.Heller herbaceum Lindl. hercoglossum Rchb.f. hercoglossum Rchb.f. (as D. wangi) heterocarpum Wall. D. D. D. D. D. D. D. D. D. D. D. D. D. hookerianum Lindl. indivisum var. indivisum (as D. porphyrophyllum) infudibulum Lindl. infudibulum Lindl. (as D. jamesianum) insigne (Blume) Rchb.f. jenkinsii Wall. ex Lindl. jenkinsii Wall. ex Lindl. (as D. aggregatum var. jenkinsii) johanis Rchb.f. johnsoniae F.Muell. jonesii var. jonesii (as D. fusiforme) jonesii var. jonesii. (as D. ruppianum) kauldorumii T.M.Reeve kingianum Bidw. D. leucocyanum T.M.Reeve D.aff. leucohybos D. lichenastrum (F.Muell.) Kraenzl. D. linawianum Rchb.f. D. lindleyi Steud. D. lindleyi Steud. (as D. aggregatum) D. lineale Rolfe D. lineale Rolfe (as D. grantii) D. lineale Rolfe (as D. veratrifolium) D. linguella Rchb.f. D. linguiforme Sw. D. D. D. D. lituiflorum Lindl. lodgesii Rolfe longicalcaratum Hayata longicornu Lindl. D. D. D. D. lohohense Tang & Wang lyoni Ames macranhum A.Rich. macrophyllum A.Rich. D. D. D. D. D. D. D. D. macrophyllum A.Rich. (as D. musciferum) macrophyllum A.Rich. (as D. polysema) mannii Ridl. mayandyi T.M.Reeve & Renz microbulbon Blatter & McCann microchilos Dalzell mirbelianum Gaudich. mohlianum Kraenzl. 20 19 19 19 20 D. moniliforme (L.) Sw. (as D. monile) D. monophyllum F.Muell. 38 40 40 38 40 38 38 38 38 38 38 38 38 38 38 38 36 76 ca. 76 38 38 38 38, 40, 76 38 38 38 38 38 38 38 40 38 38 38 38 43 38 40 38 38 38 + 2f 38, 38 + 3f 38, 38 + B 38 36 38 38 38 38 G85 G85 G84 G85, GJ90 G88 M74 G85 G85 TK84 G84, GJ90 G85 G88 M73, M77, TK84 G84, G85, GJ90 G81, G85, GJ90 TK84 M73, TK84, G88, GJ90 TK84 G88, GJ90 TK84 G85 TK84 G88 TK84 G85, GJ90 G85, G88 TK84 TK84 G88 G88 G85 G88 G85, G88, GJ90 M73, TK84, GJ90 G85 M73 TK84 M73, G85 G85, GJ90 G84 M73, GJ90 G85, G88 M74 G81, G85, GJ90 G84 G88 G84 G85 G85, GJ90 G88 TK84 G85, GJ03 GJ90 G85 G84, GJ90 TK84 M73, TK84 G85, GJ03 ca.38 38 38, 38 + 1-ª3f 38 38 M73 M73, G84, G85, G88 TK84 M73, GJ90 G85, GJ90 40 38 38 36 + 2f 38 38 19 19 D. moniliforme (L.) Sw. Source 19 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 D. D. D. D. D. D. D. D. D. D. D. D. D. 2n CHROMOSOMAL VARIABILITY OF EPIDENDROIDEAE 269 Appendix 1. Continued TAXON D. mooreanum Lindl. D. morrisonii Schltr. Dendrobium mortii F.Muell. D. moschatum Sw. mutabile (Blume) Lindl. nanum Hook.f. (as D. mabelae) nebularum Schltr. nindii W.Hill (as D. toffti) nobile Lindl. 2n Source 19 36 38 38 38 G88 GJ03 G88 M73, G81, G84, GJ90 19 40 38 38 38 38 G85 G84 G88 TK84 M77, TK84, G85, G88, GJ90, GJ06 TK84 TK84 TK84 GJ03 G85, G88, GJ90 G85 G85 TK84 GJ90 TK84, GJ90 TK84 G85 M73, G84, G88, GJ90 G84 G88 G85 G88 TK84 G88 G85, GJ90 G84 TK84 M73, G88, GJ90, GJ94 G88 G81 G88, GJ90 G88 G85, GJ90, GJ96 M73 TK84 G85 D07 G84, GJ90 TK84 M73, G84 G88 G85 G85, G88, GJ90 TK84 G85 G85 TK84, GJ90 M73, G84, GJ90 G88, GJ90 G85, GJ90 G84 TK84 TK84 M73, TK84, G84, GJ90 M73, G88 M73, TK84, G88 TK84, G88, GJ90 G84 ca. 57 38–40 57 38, 57, 76 38 38 40 40 38 40 D. nobile Lindl. misc. cultivars D. ochreatum Lindl. D. oreodoxa Lindl. D. ovatum (Willd.) Kraenzl. D. D. D. D. D. D. D. D. D. D. D. D. D. D. D. D. D. D. D. D. palpebrae Lindl. parcum Rchb.f. (as D. parcoides) papilio Lohner parishii Rchb.f. parishii Rchb.f. (as D. rhodopterigium) patentilobum Ames & Schweif. pendulum Roxb. petiolatum Schltr. philippinensis Ames pinifolium Ridl. platygastrium Rchb.f. plicatile Lindl. polyanthum Wall. ex Lindl. (as D. cretaceum) polyanthum Wall. ex Lindl. (as D. primulinum) polyschistum Schltr. porphyrochilum Lindl. praecinctum Rchb.f. (as D. pauciflorum) pseudoglomeratum T.M.Reeve & J.J.Wood pugioniforme A.Cunn. pulchellum Roxb. ex Lindl. D. D. D. D. D. D. D. D. D. D. D. D. D. D. D. punamense Schltr. pygmaeum Sm. (as Ichthyostomum pygmaeum) ramosii Ames regium Prain revolutum Lindl. rhytidothece Schltr. rhodostictum F.Muell. & Kraenzl. rigidum Lindl. ruckeri Lindl. (as D. ramosum) ruginosum Ames salascense (Blume) Lindl. sanderae Rolfe scabrilingue Lindl. schneiderae F.M.Bailey schoeninum Lindl. (as D. beckleri) D. D. D. D. D. D. D. schuetzei Rolfe schulteri J.J.Sm. secundum (Blume) Lindl. senile Par. & Rchb.f. signatum Rchb.f. (as D. hildebranditii) smillie F.Muell. smillie F.Muell. (as D. ophioglossum) 20 19 38 38 38 38 38 38 38 40 40 38 38 ± 1f 38 38 19 20 19 20 20 38 38 38 40 38 38 38 40 38 40 38 36 38 40 36 38 40 38 38 38 38 40 38 40 38 38, 38 + 1f 38 38 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 D. D. D. D. D. n 270 L. P. FELIX and M. GUERRA Appendix 1. Continued TAXON n D. terrestre J.J.Sm. D. tetragonum var. giganteum P.A.Gilbert D. thyrsiflorum Rchb.f. ex André D. D. D. D. tokai Rchb.f. toressae (F.M.Bailey) Dockrill tortile Lindl. transparens Wall. ex Lindl. D. D. D. D. D. D. D. D. trigonopus Rchb.f. undulatum R.Br. undulatum var. broonfieldii unicum Scidenf. uniflorum Griff. vagans Schltr. (as D. semanii) vannouhuysii J.J.Sm. ventricosum Kraenzl. D. D. D. D. D. D. D. venustum Teijsm. & Binn. (as D. ciliatum) verruciferum Rchb.f. vexillarius J.J.Sm. victoria-reginae Loher violaceum Kraenzl. violaceum Kraenzl. (as D. quinquecostatum) wardianum R.Warner D. D. D. D. wassellii S.T.Blake wightii A.D.Hawkes & A.H.Heller (as D. graminifolium) williamsianum Rchb.f. williamsonii Day & Rchb.f. D. woodsii P.J.Cribb Dendrobium sp. Dendrobium sp. Dendrobium sp. Dendrobium spp. Diplocaulobium aratriferum (J.J.Sm.) P.F.Hunt & Summerh. 19 19 20 19 20 19 19 19 19 Source 38 38 38 38 38 40 40 38 38 38 38 40 38 57 76 38 40 38 38 38 38 38 76 40 40 38 36 38 40 G85 TK84 TK84 M73, G85 G88 TK84 G85, G88, GJ90 M73, TK84 M73 TK84 G85 GJ90 G85, G88, GJ90 G85 G85 GJ90 GJ90 G81, TK84, GJ90 M73, GJ90 G85 TK84 G88 G85 G84 G88 G84 G85 G88 M73, TK84, G85, G88, GJ90 TK84 G85 M73, G85, GJ90 M73, TK84, GJ90 G84, G85 M73, TK84, G88 M73, TK84 TK84 G85, G88 GJ90 G85 G85 TK84 TK84 GJ90 GJ96 G85 M73, G85 G85 G84, GJ90 G81, TK84, GJ90 TK84 TK84 G85, GJ90 TK84 G88, GJ90 G81, GJ90 G85 G85 G81 G81 G85 GJ90 G88 38 38 38 40 38 38 38 38 40 38 38 38 20 40 41 38 38 38 38 38 ca. 57 40 38 38 38 57 38 36 40 38 40 40, 42 38 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 D. speciosum Sm. D. speciosum Sm. (as D. speciosum var. fusiforme) D. speciosum var. hillii Mast. D. spectabile (Blume) Miq. D. sphenochilum F.Muell. & Kraenzl. (as D. confusum) D. spurium (Blume) J.J.Sm. D. stuposum Lindl. Dendrobium stratiotes Rchb.f. D. strebloceras Rchb.f. D. striaenopsis M.A.Clem. & D.L.Jones (as D. phalaenopsis var. schroederianum) D. striolatum Rchb.f. D. stuposum Lindl. D. subclausum var. phlox (Schltr.) J.J.Wood (as D. phlox) D. ¥ suffusum Cady D. sulphureum Schltr. D. subulatum (Blume) Lindl. D. sulcatum Lindl. D. ¥ superbiens Rchb.f. D. sutepense Rolfe ex Downie D. tangerinum P.J.Cribb D. taurinum Lindl. D. tenuissimum Rupp D. teretifolium R.Br. D. teretifolium var. fasciculatum Rupp D. terminale E.C.Parish & Rchb.f. 2n CHROMOSOMAL VARIABILITY OF EPIDENDROIDEAE 271 Appendix 1. Continued TAXON n Subtribe Collabiinae Acanthephippium bicolor Lindl. A. pictum Fukuy A. striatum Lindl. A. striatum Lindl. (as A. sinense) A. splendidum J.J.Sm. A. sylhetense Lindl. Acanthephippium sp. Calanthe actinomorpha Fukuy. C. alismifolia Lindl. C. C. C. C. C. C. alismifolia Lindl. alismifolia Lindl. alismifolia Lindl. alpina Hook.f. ex alpina Hook.f. ex arcuata Rolfe (as (as C. faurieii) (as C. japonica) (as C. okinawaensis) Lindl. Lindl. (as C. schlechteri) C. caudatilabella) C. C. C. C. C. C. argenteostriata C.Z.Tang & S.J.Cheng arisanensis Hayata aristulifera Rchb.f. aristulifera Rchb.f. (as C. eliptica) aurantiaca Ridl. biloba Lindl. Source 38 38 38 38 38 G88 TK84 G88 G88 G88 G81 G88, GJ90 G85, GJ90 G84 M73, G88, GJ90 TK84 M73, G81, GJ90 GJ90 G85 G88 TK84 20 20 19 24 II 22 20 40 40 40 40 18 38 38 38 ca. 40 38 48 48 48 42 46 48 48 40, 38 44 40 40 40 20 42 40 38 45 40 40 40 40 40 38 C. brevicornu Lindl. C. cardioglossa Schltr. 20 C. C. C. C. C. C. C. C. C. C. chevalleri Gagnep. chloroleuca Lindl. clavata Lindl. coreana Nakai conspicua Lindl. cremeoviridis J.J.Wood davidii Franch. (as C. bungoana) davidii Franch. (as C. matsudae) densiflora Lindl. discolor Lindl. 20 20 20 C. C. C. C. C. C. C. discolor Lindl. (as C. discolor var. kanashiroi). graciliflora Hayata graciliflora Hayata (as C. hamata) hancockii Rolfe hattorii Schltr. hennisii Loher herbacea Lindl. ca. 44 46 20 21 C. hirsuta Seidenf. 40 40 40 40 46 42 40 40 40 40 40 40 40 40 42 40 + 0–2B 42 46 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 G84 G85 G84, G85 GJ96 GJ06 G85 G84 G85 M73, G81 G81 TK84 G84, TK84, G85 G84 M73 G84 G85 GJ94 GJ94 G85, GJ94 TK84, G84, GJ94 G85 G84, G88 M73, TK84 M73, G81 M73 TK84 GJ94 TK84 G81, G88 M77, G85, GJ94 GJ91 GJ94 GJ94 G84 G85, GJ94 G84, G85, GJ94 M73, G84, G88, GJ91, GJ96 G84, TK84 G85, GJ94 G88, GJ94 GJ94 G81, G84 GJ94 M73G81, GJ94 G81, GJ94 GJ94 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 D. chrysotropis (Schltr.) A.D.Hawkes D. chrysotropis (Schltr.) A.D.Hawkes (as Dendrobium chrysotopis) D. aff. fariniferum (Schltr.) Carr D. hydrophilum (J.J.Sm.) Kraenzl. D. mekynosepalum (Schltr.) Kraenzl. Epigeneium amplum (lindl.) Summerh. E. fuscescens (Griff.) Summerh. (as Dendrobium fuscescens) E. nakaharaei (Schltr.) Summerh. (as D. nakaharaei) Epigeneium nakaharaei (Schltr.) Summerh. (as D. sanseiense) E. rotundatum (Lindl.) Summerh. (as D. rotundatum) Flickingeria comata (Blume) A.D.Hawkes (as Dendrobium fimbriatolabellum) F. macraei (Lindl.) Seidenf. (as Dendrobium macraei) F. scopa (Lindl.) Brieger (as D. scopa) Genyorchis pumila (Sw.) Schltr. Saccoglossum verrucosum L.O.Williams Trias stocksii Benth. ex Hook. 2n 272 L. P. FELIX and M. GUERRA Appendix 1. Continued TAXON izu-insularis (Satomi) Ohwi & Satomi lyroglossa Rchb.f. lyroglossa Rchb.f. (as C. foertermannii) lyroglossa Rchb.f. (as C. liukiuensis) mannii Hook.f. musca (D.Don) Lindl. nipponica Makino oblanceolata Owi & T.Koyama plantaginea Lindl. puberula Lindl. puberula Lindl. (as C. reflexa) C. pulchra (Blume) Lindl. Calanthe rosea (Lindl.) Benth. C. rubens Ridl. C. rubens Ridl., (as C. elmeri) C. speciosa (Blume) Lindl. (as C. formosana) C. speciosa (Blume) Lindl. (as C. yushuni) C. striata R.Br. C. striata R.Br. (as C. striata var. sieboldii) C. striata R.Br. ex Lindl. (as C. bicolor) C. striata R.Br. ex Lindl. (as C. sieboldii) C. succedanea Gagnep. C. sylvatica (Thouars) Lindl. C. sylvatica (Thouars) Lindl. (as C. longicalcarata) C. sylvatica (Thouars) Lindl. (as C. masuca) C. tricarinata Lindl. C. C. C. C. C. C. C. triplicata (Willemet) Ames triplicata (Willemet) Ames (as C. triplicata var. angraecifolia) triplicata (Willemet) Ames (as C. furcata) triplicata (Willemet) Ames (as C. veratrifolia) trulliformis King & Pantl., ventilabrum Rchb.f. (as C. langei) vestita Wall. ex Lindl. Calanthe sp. Calanthe sp Cephalantheropsis calanthoides (Ames) T.S.Liu & H.J.Su (as C. koshunensis) C. obcordata (Lindl.) Ormerod (as C. gracilis) C. obcordata (Lindl.) Ormerod (as Calanthe venusta) C. obcordata (Lindl.) Ormerod (as Phaius gracilis) Chrysoglossum ornatum Blume Gastrorchis francoisii Schltr. G. humblotii (Rchb.f.) Schltr. G. humblotii humblotii var. schlechteri (H.Perrier) Senghas ex Bosser & P.J.Cribb G. lutea (Ursch & Toill.-Gen. ex Bosser) Senghas G. tuberculosa (Thouars) Schltr. Nephelaphyllum cordifolium (Lindl.) Blume Pachystoma pubescens Blume (as Pachystoma senile) Phaius elatus P. flavus (Blume) Lindl. P. flavus (Blume) Lindl. (as P. maculatus) P. flavus (Blume) Lindl. (as P. minor) P. flavus (Blume) Lindl. (as P. minor f. punctatum) P. luridus Thwaites P. mannii Rchb.f. P. mindorensis Ames P. mishmensis (Lindl. & Paxton) Rchb.f. 20 20 20 20 20 2n Source 40 40 40 40 40 40 38 40 40 40 + 0–1B 40 G84 G84, G85, GJ94 TK84 M73 M77, G81, G84, G88 G84 G84 G84 M73, GJ94 G81, G88, GJ94 M73, G84, G85, GJ90, GJ94, G84, G88 GJ94 G84, G88, GJ94 GJ94 G85, GJ94 M74 M77 G85 G84 G84, GJ90, GJ94 GJ94 GJ94 G85 M73, GJ94 M73, G85 GJ94 TK84 G84, G85, G88, GJ94 GJ96 M73 TK84, G84, G88 M73 GJ96 TK84 GJ94 M73 TK84 GJ94 M73, G88, GJ94 TK84 G85 40 44 40 + 0–2B 44 40 20 40 40 40 40 44 40 40 40 40 40, 60 40 40 40 40 + 0–2B 20 40 20 21 20 42 20 40 40 40 40 42 20 36 40 40 40 40 40 36 + 3–7B 40 42 42 GJ91 GJ06 GJ06 GJ06 GJ06 GJ06 G81 G81, G88, GJ94 TK84 G85, GJ90, GJ96 G81 44 44 28 42 42 TK84 GJ90 G84 TK84 G81 G88 G85 GJ06 GJ06 21 22 + 2f P. mishmensis (Lindl. & Paxton) Rchb.f. (as P. gracilis) P. pulchellus Kraenzl. P. pulchellus var. sandrangatensis Bosser 42 40 40 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 C. C. C. C. C. C. C. C. C. C. C. n CHROMOSOMAL VARIABILITY OF EPIDENDROIDEAE 273 Appendix 1. Continued TAXON n P. tankervilliae (Aiton) Blume 23 2n 32, 40 76 G81 G84 GJ96 22 + 2B 38 38 48 21 Phaius sp. Phaius sp. Plocoglottis javanica Blume P. lowii Rchb.f. Spathoglottis affinis de Vries S. aurea Lindl. (as S. microchilina) S. pacifica Rchb.f. S. paulinae F.Muell. (as S. rivularis) S. petri Rchb.f. S. plicata Blume S. plicata Blume (as S. spicata) S. pubescences Lindl. 20 38 38 38 38 40 40 40 40 40 40 40 36 19 S. unguiculata (Labill.) Rchb.f. Tainia bicornis Benth. T. latifolia Benth. 15 + 1–3B T. T. T. T. T. 20 latifolia (Lindl.) Rchb.f. (as T. khasiana) laxiflora Makino minor Hook.f. penangiana Hook.f. penangiana Hook.f. (as T. hookeriana) T. viridifusca (Hook.) Benth. ex Hook.f. Tainia spp. Tainia sp. (as Ania sp.) 40 20 F69 = Fedorov, 1969; M73 = Moore, 1973; M74 = Moore, 1974; M77 = Moore, 1977; G81 = Goldblatt, 1981; TK84 = Tanaka and Kamemoto, 1984; G84 = Goldblatt, 1984; G85 = Goldblatt, 1985; G88 = Goldblatt, 1988; GJ90 = Goldblatt and Johnson, 1990; GJ91 = Goldblatt and Johnson, 1991; GJ94 = Goldblatt and Johnson, 1994; GJ96 = Goldblatt and Johnson, 1996; Goldblatt and Johnson, 2000; Goldblatt and Johnson, 2003; Goldblatt and Johnson, 2006; FG98 = Felix and Guerra, 1998; DA09 = Davinha et al. (2009); PI09 = Pinheiro et al. (2009); D07 = Dawson et al. (2007); PW = Present work. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 40 32 32 36 + 0–9B 40 ca. 72 40 G81 G85 G84, G88 GJ90 GJ90, GJ96 M73 M73 M73 GJ90 GJ96 G84, G88 G84 G84 G84, G88, GJ96 GJ06 G88 GJ06 TK84, G84 G88 G85 M77 TK84 GJ06 G88 G81 G84 G84 TK84 G85 TK84 G84, GJ91, G96 42 48 P. tankervilliae (Aiton) Blume (as P. grandifolius) P. wallichii Lindl. Source 274 L. P. FELIX and M. GUERRA Appendix 2. Species with chromosome numbers uncertain or unimportant to evolutionary interpretation of the group. Abbreviation according to Table 1. TAXON n 2n Acampe papillosa (L.) Lindl.1 A. papillosa (L.) Lindl.1 36 18 18, 19 36 A. suavissima Lindl.1,4 Anthogonium gracile Lindl.1 Arundina bambusifolia Lindl.1 A. graminifolia (Don) Hochr.1 Bletilla formosana (Hay.) Schltr.6 B. striata (Thumb.) Rchb.f.4 Bulbophyllum cylindraceum Lindl. B. distans Lindl.2 B. leopardinum Lindl.1 B. lobbi Lindl.1,2 B. mastersianum J.J.Smith (as C. mastersianum)3 B. odoratissimum (Sm.) Lindl.1 B. oreonastes Rchb.f.1 B. patens King2 B. reptans Lindl.1 B. vagans Ames & Rolfe2 B. wendlandianum (Kraenzl.) U.Dammer2 Calanthe argenteo-striata C.Z.Tang & S.J.Cheng2 C. brevicornu Lindl.1,3 18 20 51 40 40 (22) 36 40 ca. 38 27 21 32 42 16 ca. 38 42 57 18 38–42 38–40 29 80 57 42 95 57 45 24 38(57) C. brevicorum9 C. chloroleuca Lindl.1 C. discolor var. amaniana (Fukuyama) Nasamune5 C. longicalcarata Hay.6 C. masuca Lindl.1 24 28 60 20 52 21 C. C. C. C. matsudai Hayata1 puberula Lindl.1 reflexa Maxim.1 tricarinata Lindl.1,5 C. triplicata (Willemet) Ames6 Calanthe sp. Calanthe sp. (Thailand)9 Cattleya bicolor Lindl. var. measuresiana5 C. forbesii Lindl.3 C. intermedia Grah.1,3 C. intermedia var. alba7 C. labiata Lindl.1 C. labiata var. amesiana3 Cattleya sp.1,3 Cephalanthera damasonium (Mill.) Druce1 20, 21 44 42 42 42 60 20 20 58 80 54–60 46, 55, 76 41 + 1f 42 40, 41 46, 56, 76 54 32 G88 G81, GJ94 TK84 M73 TK84 TK84 G84, GJ94 TK84 M73 M73 G81 TK84 TK84 M74 M73 G84 M73 G81 TK84 TK84 G81 M73 G85 G81, GJ90 G85 G88 GJ94 M73, GJ94 TK84 G81 M73, G81 G84 M74 M73 G81 G85 G81 G85 M73 TK84 G85 M74 TK84 TK84 TK84 GJ96 TK84 TK84 TK84 GJ91 G85 TK84 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 Aerangis compta Summ.2 A. hitchongii9 Aerides multiflorus Roxb.3 A. odoratum Lour.1 Source CHROMOSOMAL VARIABILITY OF EPIDENDROIDEAE 275 Appendix 2. Continued TAXON 24 D. D. D. D. D. D. D. D. D. D. D. D. chrysanthum Wall.5 chrysotoxum Lindl. crumenatum Sw.1 densiflorum Lindl.1 delicatum Bailey5 dicuphum F. Muell.2 digibum var. compactum C.T. White3,5 distichum Rchb. f.5 dixanthum Rchb. f.2 fimbriatum Hook.1 fimbriatum var oculatum1,2 gibsoni Lindl.1 TG84, G88 G84, G88 TK84, G91 M74 TK84 G84 M73 G81 G81 M77 G85 M73 TK84 GJ90 GJ90 G88 M73, GJ90 TK84, G88 G88 38 43 44 44 76 138 95 32–35 80 40 40 40 57 20 76 40 40 42 ca. 57 39 ca. 57 57 41 18 43 40 D. kingianum Bidw.4, 20 C. bowringiana Weitch ex Gardn.1,5 21 D. kingianum var. album4 leonis Rchb.f.1 lodgesii Rolfe1 longicornu Lindl.2 longispicatum9 macrostachyum Lindl.1 moschatum Sw.1,2 nagasaki9 nakaharaei Schltr.1 nobile Lindl.1 nobile var. nobilius4,5 nobile var. virginale5 30, 42 16–34 + 2B, 33 48 36 18 + 0–6B D. hercoglossum Rchb.f.5,1 D. D. D. D. D. D. D. D. D. D. D. Source 16 22 C. micranthum Lindl.1 Cyrtochis arcuata subsp. variabilis Summ.5 Cryptopus elatus (Thouars) Lindl. Dendrobium agregatum Roxb.3 D. amoenum Wall.5 D. anceps Sw.1 D. aphyllum (Roxb.) Fischer1 D. bicameratum Lindl.1 D. brymerianum Rchb. f.1 D. candidum Wall.5 2n G88 TK84 TK84 GJ90 TK84 G84, GJ90 TK84 TK84, G84 TK84 GJ90 G88 G85 57 36 ca.57, 74, 112–114 41 42 60 ca.57, ca. 76 G88 G81 TK84 40 40 43 38 M73 TK84 G84 TK84 M73 M73 G84 M74 G85 TK84 TK84 20 39 38 30 40 ca. 57 57 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 TK84 TK84 TK84 TK84 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 C. longibracteata Blume3 C. longifolia (L.) Fritsch3 C. rubra (L.) Rich.1 Cleisostoma micranthum (como Sarcanthus micranthus)1 C. pallida (como S. palidus Lindl.)1 Coelogyne corymbosa Lindl.1 C. elata Lind.1 n 276 L. P. FELIX and M. GUERRA Appendix 2. Continued TAXON n Source 40 76 38 40 38, 57 40 57 38 38 40 ca. 80 57 ca. 36 (30) 20 41 40 ca. 57 57 57 80–90 39–40 45–85, ca. 80 42–48 48–57 57, 62 20 + 2B G81 GJ90 G84 G81 TK84 G81 G81, GJ90 G85 M77 G84 TK84 G85 G88 TK84 TK84 GJ96 G85 TK84 GJ90 G88, GJ94 TK84 TK84 TK84 TK84 TK85 GJ91, GJ96 M73 TK84 G81 M77 G88 G84 TK84 G84 G84 TK84, G84 G88 TK84 TK84 M73 G88 Holcoglossum junceum Tsi5 Laelia peduncularis Lindl.1 57 40–44 GJ96 TK84 Listera cordata (L.) R.Br.3 Listera ovata (L.) R.Br.3,4 39–42, 44 34–36 ca. 38 17, 20, 32, 35, 37, 39, 40, 42, 44 G85 G85, GJ91 GJ00 TK84, G85, G88 D. parishii Rchb.f.1,5 D. phlox var. flava9 D. pierardii Roxb.1,3 19–20 D. primulinum Lindl.1,5 D. pitcherianum Rchb.f.9 D. sawianum Lindl.9 D. senile Par. & Rchb.f.1 D. sophronites Schltr.4,5 D. suffusum Cady5 D. toressae (Bailey) Dockr.4 D. transparens Wall.1 D. ventricosum Kraenzl.1 D. verruciferum Rchb.f.2 D. wardianum Warm.1 D. wardianum var.album4,5 D. williamsonii Day & Reich.5 Doritis taenialis Benth.1,5 Epidendrum atropurpureum Willd.3,5 E. difforme Jacq. (= Neolehmannia difforme)3 E. nocturnum Jacq.3,4 E. nocturnum var. guadetoupense3 E. radicans Pav.3 20 19 19 40, 70 Eria acervata Lindl.1 E. alba L.1 E. biflora Griff.1 E. coronaria Rchb.f.3 E. dalzellii Lindl.1 E. E. E. E. E. E. 20 34 46 17, 19, 22, 26 microchilos Lindl.1 graminifolia Lindl.1 noodiana9 ovata Lindl.1 pannea Lindl.1 spicata (D.Don) Hand-Mazz.1 24 + 5–7B 24 + 10B 24 + 5B 24 + 5–11B 42 44 44 18 16 Luisia teres Blume (= L. teretifolia)1 L. trichorhiza Blume1 Mediocalcar aff. pygmaeum Schltr.5 21 20 85 TK84 M73 G88 © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 2n CHROMOSOMAL VARIABILITY OF EPIDENDROIDEAE 277 Appendix 2. Continued TAXON Microcoelia caespitosa (Rolfe) Summ.2 Neotia listeroides Lindl.1,3 N. nidus-avis (L.) Rich.1 n Source 47 36, 46 G84 TK84 TK84 gj90 GJ94 G81 G84 GJ90 TK84 M73 M73 G81 TK84 TK84 TK84 G85 M77 M73 GJ91 GJ91, GJ96 GJ91 M73 M73 TK84 TK84 TK84 M73, TK84, GJ96 TK84 G85 TK84 M73 M74 G84 TK84 G84 M73 16 60 60 36 + 4–10B 18 + 4–10B N. nigra (L.) Rchb.f.5 Nephelaphyllum cordifolium Lindl.1 18 + 3–8B Ornithochilus fuscus Wall.1 Phaius albus Lindl. P. mishmensis Rchb.f.1,4 (36) 44 31 ca. 50 114 69 + 3f 76 44 Phalaenopsis amabilis Blume5,2 P. schilleriana Rchb.f.5 Pholidota bicolor Lindl.9 P. calcarata Rchb.f.9 P. yunnanensis (Rolfe) Rolfe Pleione bulbocodioides (Franch.) Rolfe5 P. forresttii Schltr.1,2 P. humilis Lindl.5 P. praecox var. wallichiana9 P. versailles9 Polystachya cultriformis Lindl.1,2 P. galeata (Sw.) Rchb.f.4 P. rhodoptera Rchb.f.1 Renanthera coccinea Lour.5,2 Saccolabium calceolare Lindl.3,1,2,6 S. papilosum Lindl.1,2,6 S. japonicus Miq. Sarcanthus crinaceous9 Spatoglottis plicata Blume1,6,5 21 120 120 38, 39, 42, 44 60 40 80 38, 39 ca. 40 19 54 + 1 Taeniophyllum aphyllum Makino1 Tainia spp.8 Thunia alba Rchb.f.1,4 44 36 ca. 40 ca. 40 38 ca. 40 Tunia alba Rchb.f.1,4 T. marshaliana Rchb.f.4 Vanda alpina Lindl.1 V. coerulea Griffith1 V. coerulescens Griff.1 V. cristata Lindl.1 V. densiflora Lindl.1 V. luzonica Loher ex Rolfe1 V. parviflora Lindl.1,4 V. pumila Hook.f.1,5 V. polyantha (W.W.Sm.) Tang & Wang9 V. roxburghii R.Br.1 ca. 114, 115 19–20 19 36 38 18 60 24 32, 40 42 10 (36) 40 (36) 36(42) 42 ca. 40 40 72, 76 38 (42) © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 G81 G81 G81 G85 G88 G81 M73 TK84 G85 TK84 TK84 TK84 M73 G85 GJ91, GJ96 GJ91, GJ96 TK84 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 2n 278 L. P. FELIX and M. GUERRA Appendix 2. Continued TAXON n V. spatulata Spreng.5 V. suavis Lindl.9 V. teres Lindl.1 V. tricolor Lindl.1,4 Vanda sp.5 1 2 3 4 5 6 7 8 9 Source 114 114, 115 38 (36) ca. 16 ca. 18, ca. 20 28 TK84, G88 TK84 TK84 TK84 TK84 TK84 TK84 TK84 TK84 GJ96 ca. 16 ca. 18 57 Chromosome number clearly conflicting with previous counts to the species or the genus; Odd diploid numbers; Poorly defined count; Imprecise counts (ca.) substituted by another more precise record; Occasional polyploids in this species; Apparently haploid number; Occasional aneuploids; Different chromosome numbers reported for several undetermined species. Misidentification. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163, 234–278 Downloaded from https://academic.oup.com/botlinnean/article-abstract/163/2/234/2418561 by guest on 25 May 2020 V. tricolor var. suavis1,4 2n