The Cytology and Taxonomy of the Genus Pleïone D.

Don (Orchidaceae)
Author(s): P. F. Hunt and C. G. Vosa
Source: Kew Bulletin, Vol. 25, No. 3 (1971), pp. 423-432
Published by: Springer on behalf of Royal Botanic Gardens, Kew
Stable URL: http://www.jstor.org/stable/4103191
Accessed: 21-06-2016 04:08 UTC

REFERENCES
Linked references are available on JSTOR for this article:
http://www.jstor.org/stable/4103191?seq=1&cid=pdf-reference#references_tab_contents
You may need to log in to JSTOR to access the linked references.

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at
http://about.jstor.org/terms

JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted
digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about
JSTOR, please contact support@jstor.org.

Springer, Royal Botanic Gardens, Kew are collaborating with JSTOR to digitize, preserve and extend
access to Kew Bulletin

This content downloaded from 129.89.24.43 on Tue, 21 Jun 2016 04:08:44 UTC
All use subject to http://about.jstor.org/terms
The cytology and taxonomy of the genus Pleione
D. Don (Orchidaceae)
P. F. HUNT & C. G. VOSA*

Several years ago, at the request of the Royal Horticultural Society, one of us
(P. F. H.) commenced a taxonomic study of the genus Pleione D. Don with
especial reference to those species found in cultivation. The preliminary
results of this work, including a key for the identification of the species with
horticultural notes, has already been published (Roy. Hort. Soc., Dict.
Gard., Suppl., 1969) but the purpose of the following contribution is to
reduce formally to synonymy many of the concepts hitherto considered
distinct. An originally independent survey into the cytology of the cultivated
species was begun in 1965 by one of us (C. G. V.) but as the results of the
two approaches, cytological and taxonomic, were so inter-related it was
agreed to collaborate and publish them in this joint paper.
About forty specific epithets have been published in the genus Pledone and
considerable confusion exists among the names given to plants in cultivation.
Understandable and acceptable differences of taxonomic opinion were
confounded with outright mis-identifications. Minor colour variations with,
occasionally, very small differences in labellum shape, size and number of
crests have been used in the past as differential characters. To complicate
the problem further, the species in cultivation were usually represented by
a very few clones which often represented only the extremes of variation in
colour, etc. Few intermediate plants have been seen by horticultural authors,
who, naturally, base their opinions on the living material with which they
are familiar.

MATERIALS AND METHODS

The localities from which living material was originally collected are
shown in Table I. The sources were mainly commercial but were also from
Botanic Gardens and private collections (see acknowledgments at the end
of the paper).
For chromosome counts, actively growing root tips were pre-treated for
3-4 hours with an aqueous solution of 8-hydroxyquinolin (o-oo2M), fixed
overnight in 1: 3 acetic-alcohol and stained in Feulgen after 6-8 minutes
hydrolysis (Jones & Daker, 1968). The same method of preparation, with
the exclusion of pre-treatment, was used for determining the relative D.N.A.
content of metaphase or late prophase nuclei. The measurements were taken
with a Deeley's integrating microdensitometer, type GN2 (Barr & Stroud,
Glasgow).
As much herbarium material as possible has been seen from several of the
larger herbaria. A very considerable number of living specimens has also
been seen and opportunities taken to discuss their characters with various
growers. Quite independently of the herbarium studies carried out at Kew,

* Botany School, Oxford.

423

This content downloaded from 129.89.24.43 on Tue, 21 Jun 2016 04:08:44 UTC
All use subject to http://about.jstor.org/terms
 424 KEW BULLETIN VOL. 25(3)
TABLE I. Original localities for plants of Pleone

Names Localities

Section Pleione
P. bulbocodioides
(as P. limprichtii (H. Smith I3020)) Kangting (Szechwan, China)
(as P. limprichtii (H. Smith 13018)) Kangting (Szechwan, China)
(as P. pogonioides) Am Whei (Western China)
(as P. formosana) Mt. Syakaroo (Formosa)
(as P. pricei) Central Mountain Range (Formosa)
(as P. formosana 'Alba') Central Mountain Range (Formosa)
P. forrestii Banks of Shweli River (Burma)
P. hookerana Tanglu, Sikkim
P. humilis Tanglu, Sikkim
,,,, (as P. humilis var. tricolor) Sikkim
. ,, yunnanensis
P. 'Frank Kingdon Ward'WesternMt. Victoria
China (Burma)
Section Dictyopleione
P. praecox Tanglu (Singalina Range, Darjeeling)
Sikkim, Khasia Hills (Assam)
,,,, (as P. praecox var. wallichiana) Tanglu
P. maculata Tanglu, Sikkim, Nepal, Khasia Hills

Drs. Wang and Tang of the Institute of Botany, Academia Sinica, Peking,
reached the same general conclusion, namely that the species of Pleione
could be reduced to less than a dozen.

CYTOLOGICAL OBSERVATIONS

The chromosomes of Pleione are very small and show very little differentia-
tion in size or morphology. (Fig. I, below.) Chromosome counts of 16 species
and forms and of one hybrid are summarized in Table 2. On the basis of the
present study the basic chromosome number of Pleione could be considered
to be x = 20, derived from an ancestral x = IO. The count of 2n 20 + IB
(La Cour, 1951) for P. pricei may relate to a haploid plant or possibly to a

FIG. I. Pleione bulbocodioides

This content downloaded from 129.89.24.43 on Tue, 21 Jun 2016 04:08:44 UTC
All use subject to http://about.jstor.org/terms
CYTOLOGY AND TAXONOMY OF PLEIONE 425

relict diploid. There are no gross differences in chromosome morphology

between the two sections of the genus. B-chromosomes, very small and dot-
like in appearance, have been found so far only in three species (Table 3,
p. 426), all belonging to section Pleione.
The polyploid species, which includes forms of P. bulbocodioides (grown as
P. limprichtii (H. Smith 13018)) (4x = 80), P. pogonioides (4x = 80), the grex
P. Versailles ('P. pogonioides x P. limprichtii') (4x = 80)) and P. yunnanensis
(6x 12o) have smaller chromosomes than the diploids.
The relative values in D.N.A. content in arbitrary units, as determined by
microdensitometry of mitotic metaphase nuclei for all species investigated, is
summarized in Table 4 (p. 426). These values show that no appreciable
differences in the D.N.A. content per nucleus have arisen during the evolution
and speciation of the genus Plefone at the diploid level. Variations in D.N.A.
values between species of the same genus are known for most angiosperm
families (John & Hewitt, 1966; McLeish & Sunderland, 1961; Rees et al.,
1966). The differences, which in many cases are considerable, are in the
main independent of any change in chromosome number, and are due to
deletions or to changes in polynemy or, in the case of some Allium species,
to duplications, which may often be large and involve most chromosomes in
the complement (Jones & Rees, 1968).

TABLE 2. Chromosome counts in Pledone

Species 2n References

Section Pleione
P. bulbocodioides
(as P. formosana) 40 Miduno, 1940
40 ? I-2B's
(as P. formosana 'Alba') 40
(as P. pricei) 2o + IB La Cour, 195I
40
(as P. limprichtii (H. Smith 13020)) 40
(as P. limprichtii (H. Smith 13018)) 80
(as P. pogonioides) 80
(as P. Versailles) 80
P. forrestii 40
P. hookerana 40 + I-2B's
P. humilis 40 + I-3B's
,,,, (as P. humilis var. tricolor) 40
.P.
,, 'Frank Kingdon Ward'
yunnanensis 12040
Section Dictyopleione
P. praecox 40
,,,, (as P. praecox var. wallichiana) 40
P. maculata 40

The variations found between the diploid species of Pleione are consistent
with very small deletions and duplications of chromosome material during
the course of speciation. In the polyploids, however, the chromosome size
and the D.N.A. content per nucleus are smaller and are about 65 per cent. of
the expected value. This reduction is proportionally the same in the tetra-
ploid and in the hexaploid species. As in most polyploid plants the numerical
increase of the chromosomes has been accompanied by a decrease in size,
with a reduction in the level of polynemy (Darlington, 1955, I963). In the

This content downloaded from 129.89.24.43 on Tue, 21 Jun 2016 04:08:44 UTC
All use subject to http://about.jstor.org/terms
 426 KEW BULLETIN VOL. 25(3)
TABLE 3. B-chromosomes in Pleione

No. of B's
Species No. of ------- Original locality
plants o I 2 3
P. bulbocodioides (as P. formosana) 24 18 4 2 - Mt. Syakaroo, Formosa

P. hookerana 24 21 3 I - Sikkim
,, ,, 6 4 I I - Tanglu, Darjeeling

P. humilis 21 14 3 3 I Tanglu, Darjeeling

,, ,, 18 Io 4 2 2 Khasia Mts.

TABLE 4. Relative D.N.A. content of mitotic metaphases in Pledone

Note: Measurements taken on Ioo nuclei for each plant.

Species 2n mean relative value

(in arbitrary units)

Section Pleone
P. bulbocodioides
(as P. formosana) 40 15"50
(as P. pricei) 40 15'58
(as P. limprichtii (H. Smith 13020)) 40o 1575
(as P. limprichtii (H. Smith 13018)) 8o I9-48
(as P. pogonioides) 80 o1945
(as P. Versailles) 8o 19'45
P. forrestii 40 15'4o
P. hookerana 40 15'44
P. humilis 40 15'75
P. yunnanensis 120 2890o
Section Dictyopleione
P. praecox 40 15'56
P. maculata 40 15'35

case of Pleione, and no doubt of many other organisms (Lewis & John, 1963),
this reduction has not been achieved by a linear halving of the D.N.A.
content.

A preliminary investigation has shown that two forms of P. bulbocodioides

(grown as P. pogonioides and P. limprichtii (H. Smith 13018)) have a regular
meiosis with a high percentage of good pollen. This indicates that they are
most probably allopolyploids. The percentage of good pollen in P.
yunnanensis 2n = 120, is very low. The species may have arisen through the
fertilization of an unreduced gamete in a tetraploid form or through the
crossing of a probably transient octoploid with an old established tetraploid.

TAXONOMIC CONCLUSIONS

As a result of the morphological and cytological examinations carried out

we have no hesitation in concluding that the number of good species to be
recognized in Plefone should be reduced to nine. Their full synonymy
together with comments on their taxonomic status are given below.
The nomenclatural types are cited in the taxonomic conclusions below
and have been examined, unless otherwise stated. In those taxa where we
have seen all the syntypes the most appropriate specimen has been selected

This content downloaded from 129.89.24.43 on Tue, 21 Jun 2016 04:08:44 UTC
All use subject to http://about.jstor.org/terms
CYTOLOGY AND TAXONOMY OF PLEIONE 427

as a lectotype. In a few cases where no type has been seen a drawing of the
original material, usually coloured, has been examined. The abbreviations
holo., iso., isosyn., lecto. and syn. are used throughout for holotype, isotype,
isosyntype, lectotype and syntype respectively.

Section Pleione

Pleione bulbocodioides (Franch.) Rolfe in Orch. Rev. II : 291 (I903).

Coelogyne bulbocodioides Franch. in Nouv. Arch. Mus. Paris, s&r. 2, o: 84
(1888). Type: Tibet, Moupine, David s.n. (K, iso.).
C. delavayi Rolfe in Bull. Misc. Inf. Kew, 1896: 195 (1896), synon. nov.
Type: China, Yunnan, Delavay 4739 (K, holo.).
C. henryi Rolfe, l.c. (1896), synon. nov. Type: China, Hupeh, Henry 6o68a
(K, lecto.).
C. pogoniofdes Rolfe, op. cit.: 196 (1896), synon. nov. Type: China, Hupeh,
Patung, Henry 3785 (K, lecto.).
Pogonia pleionoides Kraenzl. in Engl., Bot. Jahrb. 29: 267 (I9oI), synon.
nov. Type: China, Szechuan, v. Rosthorn 2131 (not seen).
Pleibne delavayi (Rolfe) Rolfe in Orch. Rev. II: 291 (1903), synon. nov.
P.formosana Hayata in Journ. Coll. Sci. Tokyo 30, art. 1: 326 (191 1), synon.
nov. Type: Formosa, Nan6, Hyah6sha, Mori 16 (not seen).
P. pricei Rolfe in Bot. Mag. 143, t. 8729 (1917), synon. nov. Type: Formosa,
Bonbonsan, Giran, Price s.n. (K, holo.).
P. mairei Schltr. in Fedde, Rep. Sp. Nov. Beih. 4: 61 (1919), synon. nov.
Type: China, Yunnan, Maire 634I (not seen).
P. speciosa Ames & Schltr. in Fedde, Rep. Sp. Nov. Beih. 4: 6i (i919), synon.
nov. Type: China, Hupeh, Wilson 1761 (K, iso.).
P. amoena Schltr., op. cit.: 185 (1919), synon. nov. Type: as for Pogonia
pleionoides Kraenzl.
P. henryi (Rolfe) Schltr., op. cit.: 186 (I919), synon. nov.
P. limprichtii Schltr., op. cit. 12: 346 (1922), synon. nov. Type: China,
Szechuan, Limpricht 1598 (K, isosyn.).
P. hui Schltr. in Fedde, Rep. Sp. Nov. 19: 377 (1924), synon. nov. Type:
China, Kiangsi, An-Fu, Wu-Kungshan, Hu 718 (not seen).
P. smithii Schltr. in Act. Hort. Goth. I: I49 (1924), synon. nov. Type: China,
Szechuan, Teng-hsiang-ying, H. Smith, 1883 (GB, holo.).
P. fargesii Gagnep. in Bull. Soc. Bot. France 78: 25 (I93i), synon. nov.
Type: China, Tchen-Keou-Tin, Farges 534 (K, iso.).
P. rhombilabia Hand.-Mazz., Symb. Sin. 7: I348 (1936), synon. nov. Type:
China, Yunnan, Bei-Lidjiang, 'v. E.' 3989 (not seen).
GEOGRAPHICAL DISTRIBUTION: Tibet, China, Formosa.
On the basis of their cytology and associated morphological and geo-
graphical difference it is perhaps better to separate the polyploids from the
diploids at subspecific level, but until a chromosome count is made of the
true P. bulbocodioides subsp. bulbocodioides it has been agreed that this separa-
tion shall not be formalized because of the possibility of it being nomen-
claturally premature. Similar subspecific treatment should probably also be
given to P. formosana which is geographically isolated from the centre of
differentiation, and which, with its very many forms, is probably in a phase
of active evolution.

This content downloaded from 129.89.24.43 on Tue, 21 Jun 2016 04:08:44 UTC
All use subject to http://about.jstor.org/terms
 428 KEW BULLETIN VOL. 25(3)
P. forrestii Schltr. in Not. Roy. Bot. Gard. Edinb. 5: io6 (1912).
Type: China, Yunnan, Tali range, Forrest 4859 (E, holo.).
GEOGRAPHICAL DISTRIBUTION: China (Yunnan), Burma.
Closely related to P. bulbocodioides but probably worthy of specific status.

P. grandiflora (Rolfe) Rolfe in Orch. Rev. I 1: 291 (Oct. Igo3).

Coelogyne grandiflora Rolfe in Journ. Linn. Soc. 36: 22 (Jan. 1903). Type:
China, Yunnan, Mengtze, Henry I I I 6 (K, holo.).
GEOGRAPHICAL DISTRIBUTION: China (Yunnan).

P. hookerana (Lindl.) B. S. Williams, Orch. Grow. Man., ed. 6: 548 (1885).

Coelogvne hookerana Lindl., Fol. Orch. Coelog.: 14 (1854). Type: Sikkim,
Darjeeling, Hooker 74 (K, holo.).
C. hookerana Lindl. var. brachyglossa Rchb. f. in Gard. Chron., ser. 3, I: 833
(1887). Type: Sikkim, cult. Lawrence (W, holo.).
Pleione hookerana (Lindl.) B. S. Williams var. brachyglossa (Rchb. f.) Rolfe in
Orch. Rev. II : 291 (1903)-
P. laotica Kerr in Journ. Siam Soc. Nat. Hist. Suppl. 9: 235 (1933), synon.
nov. Type: Laos, Pu Bia, Kerr 0977 (K, holo.).
GEOGRAPHICAL DISTRIBUTION: India (Assam, Sikkim), Nepal, Bhutan,
Thailand, Laos.

P. humilis (Sm.) D. Don, Prodr. Fl. Nep.: 37 (1825).

Epidendrum humile Sm., Exot. Bot. 2: 75, t. 98 (I8o6). Type: Nepal,
Buchanan Hamilton s.n. (not seen).
Coelogyne humilis (Sm.) Lindl., Coll. Bot., sub. t. 37 (1821).
C. humilis (Sm.) Lindl. var. tricolor Rchb. f. in Gard. Chron., ser. 2, 13: 394
(188o). Type: origin unknown, cult. Bull (W, holo.).
C. humilis (Sm.) Lindl. var. albata Rchb. f. in Gard. Chron., ser. 3, 3: 392
(1888). Type: origin unknown, cult. Sander (W, holo.).
Pleione humilis (Sm.) D. Don var. adnata Pfitz. in Engl., Pflanzenr. Orch.
Coelog.: 121 (1907). Type: not known.
P. humilis (Sm.) D. Don var. purpurascens Pfitz., op. cit.: 122 (1907). Type: not
known.
P. diantha Schltr. in Orchis 9: 44 (1915), synon. nov. Type: Burma, cult.
Hennis (not seen).
P. humilis (Sm.) D. Don var. pulchella E. W. Cooper in Roy. Hort. Soc., Dict.
Gard. 3: I6o6 (1951). Type: not known.
GEOGRAPHICAL DISTRIBUTION: India (Sikkim), Nepal, Burma.

P. yunnanensis (Rolfe) Rolfe in Orch. Rev. II: 292 (Oct. 1903).

Coelogyne yunnanensis Rolfe in Journ. Linn. Soc. 36: 23 (Jan. I903). Type:
China, Yunnan, Mengtze, Henry 111 3 (K, lecto.).
Plezone communis Gagnep. in Bull. Soc. Bot. France 78: 25 (1931 ), synon.
nov. Types: China, Yunnan, Ducloux 2057, 5645, Delavay 3141; Tibet,
Monbeig s.n. (P, syn.).

This content downloaded from 129.89.24.43 on Tue, 21 Jun 2016 04:08:44 UTC
All use subject to http://about.jstor.org/terms
CYTOLOGY AND TAXONOMY OF PLEIONE 429
P. communis Gagnep. var. subobtusum Gagnep., l.c. (1931), synon. nov.
Types: Tibet, Gnia-pa-tong, Souli I1385 & Tse-Kou, Souli' s.n. (P, syn.).
P. ganchuenensis Gagnep., op. cit.: 26 (1931), synon. nov. Type: China,
Kouytcheou, Gan-chouen, Cavalerie s.n. (P, holo.).
P. chunii Tso in Sunyatsenia I: I48 (I933), synon. nov. Type: China,
Kwangtung, Chun 43047 (PE, holo.).
GEOGRAPHICAL DISTRIBUTION: China (Yunnan).

P. scopulorum W. W. Sm. in Not. Roy. Bot. Gard. Edinb. 13: 218 (1921).
Type: China, Yunnan, Forrest, 14230 (K, iso.).
Bletilla scopulorum (W. W. Sm.) Schltr. in Fedde, Rep. Sp. Nov. 19: 375
(1924).
GEOGRAPHICAL DISTRIBUTION: Tibet, China (Yunnan).

Section Dictyopleione
P. praecox (Sm.) D. Don, Prodr. Fl. Nep.: 37 (1825)-
Epidendrum praecox Sm., Exot. Bot. 2: 73, t. 97 (1806). Type: Nepal,
Buchanan-Hamilton (not seen).
Coelogynepraecox (Sm.) Lindl., Collect. Bot., sub. t. 37 (1821).
C. wallichiana Lindl., Gen. Sp. Orch. P1.: 43 (May 1830) & in Wall., P1. As.
Rar. i: 46, t. 54 (Jul. 1830). Type: India, Pundua, Wallich 1965 (K, holo.).
C. wallichii Hook. in Bot. Mag. 76, t. 4496 (I850).
Pleione lagenaria Lindl. in Paxt., Fl. Gard. 2: 5, t. 39 (1851), synon. nov.
Type: Paxt. Fl. Gard. 2: 5, t. 39.
P. wallichiana (Lindl.) Lindl., op. cit.: 66 (I851).
Coelogyne lagenaria (Lindl.) Lindl., Fol. Orch. Coelog.: 15 (1854), synon. nov.
C. praecox (Sm.) Lindl. var. sanguinea Lindl., op. cit.: 16 (1854). Type: Sikkim,
Hooker 73 (K, holo.).
C. praecox (Sm.) Lindl. var. wallichiana (Lindl.) Lindl., 1.c. (1854).
C. reichenbachiana T. Moore & Veitch in Gard. Chron. 1868: 1210 (1868),
synon. nov. Type: Burma, Moulmein, Benson s.n. (not seen).
Plefone reichenbachiana (T. Moore & Veitch) B. S. Williams, Orch. Grow.
Man., ed. 4: 252 (1871), synon. nov.
Coelogyne birmanica Rchb. f. in Gard. Chron., ser. 2, 18: 840 (1882). Type:
Burma, Boxall s.n. (W, holo.).
C. praecox (Sm.) Lindl. var. tenera Rchb. f., op. cit. 20: 294 (1883). Type:
origin unknown, cult. Bull (W, holo.).
Pleione concolor Hort. ex B. S. Williams, Orch. Grow. Man., ed. 7: 681 (1894).
Type: not known.
P. birmanica (Rchb. f.) B. S. Williams, l.c. (1894).
P. praecox (Sm.) D. Don var. birmanica (Rchb. f.) Grant, Orch. Burm.: 167
(1895).
P. praecox (Sm.) D. Don var. sanguinea (Lindl.) Pfitz. in Engl., Pflanzenr.
Orch. Coelog.: 126 (190o7).
P. praecox (Sm.) D. Don var. candida Pfitz., l.c. (I907). Type: not known.
P. praecox (Sm.) D. Don var. alba E. W. Cooper in Roy. Hort. Soc., Dict.
Gard.: I606 (I951). Type: not known.
P. praecox (Sm.) D. Don var. wallichiana (Lindl.) E. W. Cooper, l.c. (1951).

This content downloaded from 129.89.24.43 on Tue, 21 Jun 2016 04:08:44 UTC
All use subject to http://about.jstor.org/terms
 430 KEW BULLETIN VOL. 25(3)
GEOGRAPHICAL DISTRIBUTION: China, India (Assam, Sikkim), Nepal,
Burma.

P. maculata (Lindl.) Lindl. in Paxt., Fl. Gard. 2: 5 (1851).

Coelogyne maculata Lindl., Gen. Sp. Orch. P1.: 43 (May 1830) & in Wall., P1.
As. Rar. I: 45, t. 53 (July 1830). Type: India, Pundua, Wallich 1964 (K,
holo.).
Plefone diphylla Lindl. in Paxt., Fl. Gard. 2: 66 (1851). Type: India, Khasia,
Grifith 29/138 (K, holo.).
Coelogyne diphylla (Lindl.) Lindl., Fol. Orch. Coelog.: 15 (1854).
C. arthuriana Rchb. f. in Gard. Chron., ser. 2, 15: 40 (1881). Type: origin
unknown, cult. Veitch (W, holo.).
Plefone maculata (Lindl.) Lindl. var. virginea Rchb. f. in Gard. Chron., ser. 3,
2: 682 (1887). Type: origin unknown, cult. W. H. Scott (W, holo.).
P. maculata (Lindl.) Lindl. var. arthuriana (Rchb. f.) Rolfe ex Kraenzl. in
Engl., Pflanzenr. Orch. Coelog.: 128 (1907).
Gymnostylis candida Wall. ex Pfitz., op. cit.: 127 (1907).

GEOGRAPHICAL DISTRIBUTION: India (Assam, Sikkim), Bhutan, Burma,

Thailand.

KEY TO THE SPECIES OF PLEIONE

To assist identification the following key is reprinted from the Roy. Hort.
Soc., Dictionary of Gardening Supplement (1969), to which reference should
be made for brief descriptions and cultivation notes.

I. Pseudobulb flask-shaped, gradually narrowed into a beak occasionally

somewhat flattened horizontally .......... 2
Pseudobulb barrel-shaped, abruptly contracted into a beak . 8
2. Keels of lip barbate or irregularly incised-crested. 3
Keels of lip undulate or straight ........... 6
3. Keels of lip barbate, 6-7 in number ...... 4
Keels of lip irregularly incised-crested, 4-5 in number . . 5
4. Flowers appearing at same time as young leaves, lip reniform, about

2'5-4 X 3'5-4 cm..... . . .. ...... . hookerana

Flowers appearing when no leaves present, lip elliptical, about 4 x 3 cm.
humilis
5. Lip broadly elliptical, about 5'5-6 x 3'5-4'5 cm.; leaves solitary;
pseudobulb large, about 4 cm. high and I 5 cm. broad . grandiflora
Lip reniform, about 2 x 3-5 cm.; leaves 2 at flowering time; psuedo-
bulbs small, less than 4 cm. high and I -5 cm. broad scopulorum
6. Dorsal sepal and petals acute at apex; flowers appearing at same time as
immature leaves; mature leaves 3-6-5 cm. broad . bulbocodioides
Dorsal sepal and petals sub-obtuse at apex; flowers appearing when no
leaves present; mature leaves 1.5-2 cm. broad . . . . . . 7
7. Flower varying shades of pink with deep red blotches on lip; immature
ovary as long as or longer than floral bract; pseudobulb flattened
horizontally; scape more than 6 cm. high, usually considerably longer
yunnanensis

This content downloaded from 129.89.24.43 on Tue, 21 Jun 2016 04:08:44 UTC
All use subject to http://about.jstor.org/terms
CYTOLOGY AND TAXONOMY OF PLEiONE 43I

Flowers yellow to orange with brown blotches on lip; immature ovary

much shorter than floral bract; pseudobulb not flattened horizontally;
scape 7 cm. maximum height. ........ forrestii
8. Basal sheaths of flowering stem warty; flowers larger with lip 3-4'5 X
3'5-5'5 cm., keels extending only two-thirds of length of lip. praecox
Basal sheaths of flowering stem non-warty; flowers smaller with lip
2-3 X 2-5-3-5 cm., keels extending to apex of lip . . maculata

INDEX TO NAMES IN PLEIONE

To complete this taxonomic section a list of all binomials ever published in

the genus is given below (except those by Kuntze, Rev. Gen.: 679-681 (1891)
which are members of the genus Coelogyne) and the current systematic position
indicated.

P. amoena Schltr. A synonym of P. bulbocodioides.

P. birmanica Rchb. f. A synonym of P. praecox.
P. bulbocodioides (Franch.) Rolfe.
P. chiwuana Tang & Wang. The affinity of this species is uncertain but it
could be a form of P. maculata.
P. chunii Tso. A synonym of P. yunnanensis.
P. communis Gagnep. A synonym of P. yunnanensis.
P. concolor Hort. ex B. S. Williams. A synonym of P. praecox.
P. delavayi (Rolfe) Rolfe. A synonym of P. bulbocodioldes.
P. diantha Schltr. A synonym of P. humilis.
P. diphylla Lindl. A synonym of P. maculata.
P. forgesii Gagnep. A synonym of P. bulbocodioides.
P. formosana Hayata. A synonym of P. bulbocodioides.
P. forrestii Schltr.
P. ganchuenensis Gagnep. A synonym of P. yunnanensis.
P. grandiflora (Rolfe) Rolfe.
P. henryi (Rolfe) Schltr. A synonym of P. bulbocodioides.
P. hookerana (Lindl.) B. S. Williams.
P. hui Schltr. A synonym of P. bulbocodioides.
P. humilis (Sm.) D. Don.
P. lagenaria Lindl. A synonym of P. praecox.
P. laotica Kerr. A synonym of P. hookerana.
P. lauterbachiana (Kraenzl.) Kuntze. A synonym of Hologyne lauter-
bachiana Kraenzl.
P. limprichtii Schltr. A synonym of P. bulbocodioides.
P. maculata (Lindl.) Lindl.
P. mairei Schltr. A synonym of P. bulbocodioides.
P. mandarinorum (Kraenzl. ex Diels) Kraenzl. The affinity of this species is
uncertain.
P. pogonioides (Rolfe) Rolfe. A synonym of P. bulbocodioides.
P. praecox (Sm.) D. Don.
P. pricei Rolfe. A synonym of P. bulbocodioides.
P. reichenbachiana (T. Moore & Veitch) B. S. Williams. A synonym of P.
praecox.
P. rhombilabia Hand.-Mazz. A synonym of P. bulbocodioides.
P. schillerana (Rchb. f.) B. S. Williams. A synonym of Coelogyne schillerana
Rchb. f.

This content downloaded from 129.89.24.43 on Tue, 21 Jun 2016 04:08:44 UTC
All use subject to http://about.jstor.org/terms
 432 KEW BULLETIN VOL. 25(3)
P. scopulorum W. W. Sm.
P. smithii Schltr. A synonym of P. bulbocodioldes.
P. speciosa Ames & Schltr. A synonym of P. bulbocodioldes.
P. wallichiana (Lindl.) Lindl. A synonym of P. praecox.
P. yunnanensis (Rolfe) Rolfe.

ACKNOWLEDGEMENTS

We wish to thank Professor C. D. Darlington, Dr. K. Jones and Mr. V. S.

Summerhayes for their helpful advice and criticisms. In providing us with
the indispensable living plants for cytological and morphological study we
are grateful to Mr. B. N. Ghose (Darjeeling, India), Rev. R. J. Blakeway-
Phillips, Mr. H. F. Siggers, Mr. R. B. Burbidge, Mr. B. Mathew, Mr. E.
Hodgkin, Mr. G. B. Rawinsky and Mr. D. B. Milne and to the following
Directors of Botanic Gardens: Sir George Taylor (Kew), Dr. H. R. Fletcher
(Edinburgh) and Professor P. Wendelbo (G6teborg, Sweden). For informa-
tion about various species our thanks are due to Professor G. Morel
(Versailles), Drs. T. Tang and F. T. Wang (Peking), Mr. I. Butterfield,
Mr. G. B. Rawinsky and Mr. R. M. Adey. Finally we should like to thank
the Curators or Keepers of various herbaria who have kindly lent
herbarium material for study.

REFERENCES

There are innumerable references to the genus Pleione in the popular and
semi-popular horticultural and orchid literature. The majority of the
important references are cited in the articles on P. limprichtii, P. formosana
and P. forrestii in Curtis, Botanical Magazine 174, tt. N.S. 397 & 421 and
176, t. N.S. 501 respectively. The major botanical references are given
by Pfitzer & Kraenzlin in Engler, Das Pflanzenreich, Orchidaceae-
Coelogyninae: 119 (1907): there have been no monographic accounts of
the genus since that time.

Darlington, C. D. (I955). The chromosome as a physico-chemical entity.

Nature 176: I1139-1144.
(1963). Chromosome botany and the origins of cultivated plants.
John, B. & Hewitt, G. M. (1966). Karyotype stability and DNA variability
in the Acrididae. Chromosoma 20: 155-172.
Jones, K. & Daker, M. G. (1968). The chromosomes of orchids: III.
Catasetinae Schltr. Kew Bull. 22: 421-427-
Jones, R. N. & Rees, H. (1968). Chromosome DNA variation in Allium.
Heredity 23: 58i-605.
La Cour, L. F. (1951). Annual Rep. John Innes Hort. Inst. 42:50.
Lewis, K. R. & John, B. (I963). Chromosome markers.
McLeish, J. & Sunderland, N. (196I). Measurements of deoxyribonucleic
acid (DNA) in higher plants by Feulgen photometry and chemical
methods. Expl. Cell Res. 24: 527-540.
Miduno, T. (1940). Chromosomenstudien an Orchidazeen. IV. Chromo-
somenzahlen einiger Arten und Bastarde bei Orchideen. Cytologia I I : 179.
Rees, H., Cameron, F. S., Hazarika, M. G. & Jones, G. H. (1966). Nuclear
variation between diploid angiosperms. Nature 211: 828-830.

This content downloaded from 129.89.24.43 on Tue, 21 Jun 2016 04:08:44 UTC
All use subject to http://about.jstor.org/terms