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).
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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
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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).
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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
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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
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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
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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
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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).
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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.)
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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
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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.
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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-
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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V. tricolor var. suavis1,4
2n