The Chromosomes of Orchids: IV: Bulbophyllinae Schltr.

Author(s): M. G. Daker
Source: Kew Bulletin, Vol. 24, No. 1 (1970), pp. 179-184+iii
Published by: Springer on behalf of Royal Botanic Gardens, Kew
Stable URL: http://www.jstor.org/stable/4103259
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 The Chromosomes of Orchids: IV*

BULBOPHYLLINAE SCHLTR.

M. G. DAKER

The subtribe Bulbophyllinae largely comprises the two genera, Bulbophyllum

Thou. and Cirrhopetalum Lindl., both of which have a pan-tropical distribu-
tion. Chromosome numbers for species in these genera have been given by
several authors including Hoffmann (1930), Eftimiu-Heim (I94I), Mutsuura
and Nakahira (1960, I964), Chardard (1963), Pancho (1965) and Sharma
& Chatterji (1966). Apart from the very unusual counts of 2n = 36 (Sharma
& Chatterji, 1966) and 2n = 48 (Eftimiu-Heim, i94I), the species within
the subtribe are predominantly diploid with somatic numbers of 2n = 38
or 2n = 40 (Table I, p. 18o). Many species however, remain to be investi-
gated and as a contribution to this end, chromosome numbers have been
determined in part of the large living collection of both genera at Kew and
are described below.

MATERIALS AND METHODS

All the plants examined were grown in cultivation at Kew. Species were
identified by Mr. V. S. Summerhayes and Mr. P. F. Hunt, and voucher
specimens retained as pressed or spirit material in the Kew Herbarium.
The majority of chromosome counts have been made from squash prepara-
tions of aerial root tips. These were pretreated for four hours in 0o002 M 8-
hydroxyquinoline at a temperature of I4-I6'C, fixed in acetic alcohol (1:3),

**** it to*
b

FIG. I. Karyotype diagrams of the haploid chromosome complements of Bulbophyllum and

Cirrhopetalum (x 4000). a, B. sociale; b, B. lupulinum; c, C. julchrum.

* Continued from Kew Bull. 22: 427 (1968).

'79

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 180 KEW BULLETIN VOL. 24
TABLE I. Chromosome numbers in the Bulbophyllineae given by other
authors

Species n 2n Author

Bulbophyllum Thou.
B. adenopetalum Lindl. 40 Pancho, 1965
B. aeolium Ames 40
B. affine Lindl. 36 Sharma & Chatterji, 1966
B. alagense Ames 40 Pancho, 1965
B. antenniferum (Lindl.) Rchb. f. 40
B. auratum (Lindl.) Rchb. f. 20
B. braccatum Rchb. f. 20
B. canlanoinse Ames 40 ,, ,,
B. careyanum Spreng. 38 Sharma & Chatterji, 1966
B. cornutum (Lindl.) Rchb. f. or Ridl.* 38 Chardard, 1963
B. drymoglossum Maxim. 40 Mutsuura & Nakahira, 1960
B. emiliorum Ames & Quis. 40 Pancho, 1965
B. evrardii Gagnep. 38-42 Chardard, 1963
B. inconspicuum Maxim. 38 Tanaka, 1965
B. japonicum (Makino) Makino 40 Mutsuura & Nakahira, I960
B. japonicum (Makino) Makino 40 Tanaka, 1965
B. lacerata Ames 8o0 ? 2 Pancho, 1965
B. levanae Ames var. giganteum Quis. & 6o
Schweinf.
B. lobbii Lindl. 38-42 Chardard, 1963
B. makinoanum (Schltr.) Masam. 40 Mutsuura & Nakahira, 1964
B. makinoanum (Schltr.) Masam. 38 Tanaka, 1965
B. odoratissimum (Sm.) Lindl. 38 Chardard, 1963
B. reflexiforum H. Perr. 38 .. .
B. saurocephalum Rchb. f. 20 Hoffmann, 1930
B. tenuicaule Lindl. 19 38 Chardard, 1963

Cirrhopetalum Lindl.
C. cumingii Lindl. 38 Chardard, 1963
C. cumingii Lindl. 40 Pancho, 1965
(as B. cumingii (Lindl.) Rchb. f.)
C. mastersianum Rolfe 38-40 Chardard, 1963
C. makoyanum Rchb. f. 38-40
C. ornatissimum Rchb. f. 38-40
C. picturatum Lodd. ex Lindl. 48 Eftimiu-Heim, i941I
C. stramineum Teijsm. & Binn. 38-40 Chardard , 1963
C. umbellatum (Forst. f.) Hook. & Arn. 38-40
(as C. longiflorum (Thou.) Schltr.)
C. umbellatum (Forst. f.) Hook & Arn. 38
(as C. thouarsii Lindl.)

* No authority was given for this name by the author and it is not known whether it is
B. cornutum Ridl., which is a true Bulbophyllum, or B. cornutum (Lindl.) Rchb. f. which is a
Cirrhopetalum.

and stained in Feulgen after 8 minutes hydrolysis. A full account of the

recently modified techniques has been given in a previous paper in this
series (Jones & Daker, 1968). The slides are retained in the collection of the
Cytology Section, having been made permanent by the liquid CO2 method.

RESULTS

Chromosome counts have been made for 24 species of Bulbophyllum and 14

species of Cirrhopetalum growing at Kew (Table 2, p. 182). It can be seen that
there is no variation from a basic number of x = 19 for these plants, all of
which are diploid 2n = 38, (B. lobbii aneuploid, 2n = 38 + I), with the

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THE CHROMOSOMES OF ORCHIDS: IV 181

exception of B. distans which is triploid (2n = 57). Analysis of meiosis in

pollen mother cells of this species showed some multivalent formations. In
B. calamarium and B. velutinum, the two diploid species in which meiosis was
examined, regular bivalent formation occurred.
Karyotypes of Bulbophyllum and Cirrhopetalum are essentially the same. Both
consist of sub-median to sub-terminal chromosomes ranging in size from
approximately I to 2kL. Figure I shows the haploid chromosome complements
of two species of Bulbophyllum and one of Cirrhopetalum. Each has been
arranged so that the chromosomes are divided into three groups based on
the length of the short arm. Photographs of the mitotic chromosomes of
species of Bulbophyllum and Cirrhopetalum are shown in Plates I and 2 (pp. 184
& 185) respectively.

DIscuSSION

All the evidence from plants in cultivation at Kew supports a somatic

chromosome number of 38 and not 40. Of these plants, 18 of Bulbophyllum
and two of Cirrhopetalum are from known localities in tropical Africa or
Borneo. By contast, the Bulbophyllum species examined by Pancho (1965)
were all from the Phillipines, and recorded as having 2n = 40. Unfortunately
of the species counted at Kew, there are only two, B. adenopetalum and
C. cumingii, in common with those counted by Pancho. However, it is
interesting to note that the chromosome numbers of two Kew accessions of
the former from Borneo were both 2n = 38 as opposed to 2n = 40 for the
Phillipine plant of Pancho. Similarly, C. cummingii (as B. cumingii in the list
of Pancho) was found to be 2n = 38 by both the present author and also by
Chardard (1963), and again this differs from the result of Pancho. Such a
discrepancy is unexpected since B. cumingii is restricted to the Philippines,
and all the counts must therefore have been made on material originating
from this one area.
Evidence such as this might be taken as indicative of incorrect counting.
This has certainly occurred before in the family, and is understandable.
Orchidaceous material has long been held to be cytologically difficult
because of a failure to achieve effective pretreatment so that the chromosomes
are fully contracted, and unless the chromosomes are viewed at full
metaphase-contraction it is easy to make errors in counting. The modified
techniques, given in the previous paper in the series (Jones & Daker, 1968),
have largely overcome these difficulties, but although the chromosome
counts given here are stated with confidence and show no variation, the
possibility of the occurrence of different somatic chromosome numbers
within genera should not be overlooked. Thus Tanaka (1965) reports counts
of both 2n = 38 and 2n = 40, and furthermore points out that in certain
Orchid groups including Goodyera R. Br., Bletilla Rchb. f., Calanthe R. Br.
and Eria Lindl., as well as Bulbophyllum, the lower chromosome number was
found in the species that occurred in the more northern localities.
Clearly there is much to be done in the form of careful surveys of chromo-
some number in the Orchidaceae, but if such work were to substantiate this
observation of Tanaka, that there is a correlation between chromosome
number and geographical distribution, then this would be an important
step towards understanding evolution in this family.

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 TABLE 2. Chromosome numbers in the Bulbophyllineae de

Species Origin Access

Bulbophyllum Thou.
B. adenopetalum Lindl. Borneo, Giles
B. adenopetalum Lindl. Borneo, Giles
B. apodum Hook. f. Borneo, L. M. M
B. barbigerum Lindl. Lady Harrison H
B. calamarium Lindl. Nigeria, Cooper
B. cocoinum Batem. cult. Liverpool
B. congolanum Schltr. 6
B. distans Lindl. Uganda, Osmas
B. elatius Ridl. cult. Sande
B.falcatum (Lindl.) Rchb. f. cult. San
B. frostii Summerh. Colman
B. fuscoides J. B. Petersen S. Cameroon, J. O. Wr
B. grandiflorum Bl. cult. J. Spedan L
B. imbricatum Lindl. S. Cameroon, J. O. Wrigh
B. intertextum Lindl. N. Nigeria, D. E. S
B. lobbii Lindl. Borneo, Collenette
B. lupulinum Lindl. N. Nigeria, D. E. S.
B. micholitzii Rolfe cult. Liverpool P
B. minutipetalum Schltr. Papua and New Guinea, C
B. nutans Thou. cult. L. M. M

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 TABLE 2.--continued.

Species Origin Access

B. phaeopogon Schltr. W. Trop. Africa, Greg

B. saurocephalum Rchb. f. 65
B. sociale Rolfe Wyatt 6
B. velutinum (Lindl.) Rchb. f. S. Cameroon, J. O. Wrigh
B. virescens J. J. Sm. Colma
Cirrhopetalum Lindl.
C. acuminatum Ridl. Sabah, Giles
C. andersonii Hook f. 6
C. cumingii Lindl. 6
C. gracillimum Rolfe Colma
C. lasiochilum (Par. & Rchb. f.) Hook. f. cult. L
C. lepidum (Bl.) Schltr.
C. mastersianum Rolfe cult. Sand
C. mundulum Bull cult. Sand
C. mysorense Rolfe cult. Liverpool P
C. ornatissimum Rchb. f. Wya
C. pulchrum N. E. Br.
C. robustum Rolfe Matthews
C. umbellatum (Forst. f.) Hook. & Am. Papua and New Guin
C. vaginatum Lindl. 6

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 I84 KEW BULLETIN VOL. 24
REFERENCES

Chardard, R. (1963). Contribution i l'a tude cyto-taxinomique des Orchiddes.

Rev. Cyt. et Biol. V6g. 26: 1-58.
Eftimiu-Heim, P. (I941). Recherches sur les noyaux des Orchiddes. Botaniste
31: 65-I I .
Hoffmann, K. M. (1930). Beitriige zur cytologie der Orchidaceen. Planta
o10: 523-595-
Jones K. & Daker, M. G. (1968). The chromosomes of Orchids: III. Cata-
setinae Schltr. Kew Bull. 22: 421-427.
Mutsuura, O. & Nakahira, R. (1960). Chromosome numbers of the family
Orchidaceae in Japan (3). Sci. Rep. Kyoto Prefect. Univ. Nat. Sci. Liv.
Sci. 3A series: 83-88.
-- (1964). Chromsome numbers of the family Orchidaceae in Japan (4).
Sci. Rep. Kyoto Prefect. Univ. 15 (4-1): I 1-14-
Pancho, J. V. (1965). Orchidaceae in International Organization of Plant
Biosystematists (I.O.P.B.) Chromosome Number Reports: III. Taxon 14:
52-57-
Sharma, A. K. & Chatterji, A. K. (1966). Cytological studies on orchids
with respect to their evolution and affinities. The Nucleus 9: 177-203.
Tanaka, R. (1965). Chromosome numbers of some species of Orchidaceae
from Japan and its neighbouring areas. Journ. Jap. Bot. 40: 65-77-

European taxonomic literature again.*-For the second year running,

Dr. Brummitt has produced his Index, this time with the assistance of Dr.
Ferguson. This year's production is thicker by a formidable 79 pages than
its predecessor, and such an increase in a single year is sobering in the
extreme. I do not propose to offer any criticism whatever of this publication;
these two collaborators have undertaken, apparently voluntarily, a task the
thought of being assigned to which might well cause me to apply for admission
to the next expedition to Tierra del Fuego. In his notice of the first Index
(Kew Bull. 21: 461), Mr. R. D. Meikle has already said all there is to say
about the diminishing effectiveness of such publications as they increase in
quantity, and one can only reiterate thanks to our two colleagues for doing
a remarkably exacting job remarkably well.
C. C. TOWNSEND

Index of Plant Pathology.-This long needed workt will save much time
formerly spent in looking through back numbers of Rev. Appl. Mycol. There
is a host index giving references to the volume (but not the page) of Rev.
Appl. Mycol. in which each particular parasite is mentioned, and a pathogen
index giving the names of the hosts of each particular pathogen.
D. R. DRING

* Index to European taxonomic literaturefor 1966. Compiled by R. K. Brummitt & I. K. Ferguson.

Pp. 245. Regnum Vegetabile. Vol. 53. Utrecht: International Bureau for Plant Taxonomy
and Nomenclature, 1968. Price ?i 5s. (members), ?2 I3s. (non-members).
t Compiled
Review of by
Applied
HelenMycology: Plant
MacFarlane. Pp.Host-Pathogen Index
viii + 820. Kew: to Volumes I-4oMycological
Commonwealth (1922-ig96)"
Institute, 1968. Price ?Io ios.

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CHROMOSOMES OF ORCHIDS: IV] [KEW BULLETIN, VOL. 24

PLATE I

ao

4' ' *

a 4b

%dlg~ ~iV
f0 4? 4
18~ ~t ~;4
~s sf

CB d

001

'.4

.* 4,
4.

The mitotic chromosomes of Bulbophyllum ( X 1500). a, B. coccoinum (2n - 38); b, B. lupu-

linum (.2n-
(an = 38); f, 38); c, B. grandiflorum
B. distans (an - 57). (an 38); d, B. phaeopogon (an - 38); e, B. sociale

To face page 1841

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CHROMOSOMES OF ORCHIDS: IV] [KEW BULLETIN, VOL. 24

PLATE 2

a ~ lOp.
I.1.
aF

! Iil Q s Qjl i 1 b" a

9% 4 .p 4*srg

a b

8* 4 8

* 8

** 45 it

C d

The mitotic chromosomes of Cirrhopetaluam ( I500). a, C. lepidurm (on = 38); b, C. master-

sianlum (2n = 38); c, C. pulchrum2 (2n = 38); d, C. umbellatum (2n = 38).

[To face page 185

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 THE SEPARATE NUMBERS OF THIS VOLUME
WERE PUBLISHED AS FOLLOWS

No. i Pp. 1-234 2nd March 1970 Price ?4 os. od. [C?4]
No. 2 Pp. 235-378 Ist April 1970 Price ?2 13s. od. [?2-651
No. 3 PP. 379-577 2nd November 1970 Price ?3 ios. od. [?3'50]

CORRIGENDA

Page 98, line 28 for 'rigida' read 'rigidum'

Page 169, index for 'P. hippuridoies' read 'P. hippuridoides'
Page 182, table B. cocoinum after 'Batem.' insert 'ex Lindl.'
Page 184, caption to
Plate I for 'coccoinum' read 'cocoinum'

Page 206, line 24 for 'cardofanum' read 'cordofanum'

Page 221, subheading for 'Pers.' read 'P. Br.'
and line I for 'Pers.' read 'P. Br.'

Page 458, table A. superba for 'H.B.K.' read 'Kunth'

P. elata transpose 'Hook.' after 'elata'
S. grandiflora insert 'Lodd. ex' before 'Lindl.'
S. insignis delete 'ex Hook.'
S. occulata for 'occulata' read 'oculata'

for 'S. tigrina' read 'S. devoniensis'

iii

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