CYTOLOGICAL STUDIES ON SOME EASTERN
HIMALAYAN PLANTS AND THEIR
RELATED SPECIES

by
Sachiko KUROSAWA




This report comprises the results of my cytological studies on flowering plants collected by the Botanical Expeditions to Eastern Himalaya organized by the University of Tokyo in 1967 and 1969. It also includes some additional data based on the collections of our previous Expeditions to Eastern Himalaya since 1960, and of my trip to Darjceling, Simla and Kashmir in 1970. Some supplementary data which, are closely related to those of the Eastern Himalayan ones were also given in this paper.

The materials were fixed either at the habitats or in Tokyo or Karuissawa where the plants were cultivated. The methods used (OQ, & W) are the same as those described in my paper in Flora of Eastern Himalaya pp. 658-659 (1966).

The author wishes to express her deep gratitude to Professor Hiroslii Hara for his valuable advise and constant encouragement. Thanks are also due to all the members of the Expeditions for their kind help.


MATERIALS STUDIED

Aconogonum campanulatum (Hook. f.) Hara
(Polygonum campanulatum Hook. f.)
2n=ca. 64, 60. Bhuspate Danra, E. Nepal, 2500 m (Dec. 1, 1963). [Fig 57. a (OQ)]

Aconogonum molle (D. Don) Hara (Polygonum molle D. Don)
2n=32. Kakani, C. Nepal, 2200 m (Sep. 24, 1963). [Fig. 57. b (OQ)]

var. frondosum (Meisn.) Hara
(Polygonum paniculatum Blume)
2n=32. Meghma-Chitrey, Singalila Range, 2700 m (Jul. 22, 1969).
The only report on the chromosome number of this group is that Pauwels (1959) found 2n=ca. 30 in Polygonum rude Meisn. cultivated in Botanic Garden at Kew.

Achyranthes bidentata Blume
2n=24. Tonglu, Singalila Range, 3150 m (Jul. 11, 1969). [Fig. 57. c (OQ]
This number has been reported by Pal (1964), and agrees also with that of var. japonica Miquel of Japan.

Caltha Govaniana Wall. ex Royle
2n=ca. 80. Phalut, Singalila Range, 3850 m (Jul. 18, 1969).

Caltha palustris L. var. alba Hook. f. & Thorms.
n=ca. 24 (PMC). Gulmarg, Kashmir, 2400 m (May 14, 1970).

Caltha palustris L. var. himalensis (D. Don) Mukerjee
2n=32. Kalingchok, C. Nepal, 3400 m (Sep. 10, 1970).
It has been well-known that the C. palustris group is very polymorphous and also very variable in the chromosome number ranging from 16 to 80. C. Govaniana has generally rob'ust erect stems, leaves with numerous small teeth and a closed sinus, larger flowers, and erect follicles with longer (2-3 mm) erect style. 2n==80 has hitherto been reported only in C. minor Miller and C. arctica subsp. sibirica (Regel) Tolm. But morphologically C. Gosvaniana widely differs from these two plants. It is noteworthy that three morphologically distinct wild Himalayan races mentioned above have different chromosome numbers.

Clematis napaulensis DC.
2n=16. Ratsoo, Bhutan, 2000 m (Apr. 11, 1967).

Ranunculus Brotherusi Freyn
2n=32. Sandakphu, Singalila Range, 3970 m (Jul. 4, 1969). [Pl. 21. c (OQ)]

Ranunculus adoxifolius Hand.-Mzt.
2n=ca. 48. Sandakphu, Singalila Range, 3900 m (Jul. 14, 1969). [Pl. 21.b (OQ)]

Ranunculus pulchellus C. A. Meyervar. Stracheyanus (Maxim.) Hand.-Mzt.
2n=48. Sabargan, Singalila Range, 3300 m (Jul. 19, 1969). [Pl. 21. a (OQ)]


Podophyllum hexandrum Royle (P. Emodi Wall.)
2n=12. Barshong, Bhutan, 3400 m (May 25, 1967). [Pl. 21. d (OQ); Fig. 58]

2n=12+0-2f. Gulmarg, Kashmir, 2400 m (May 14, 1970). [Pl. 21. e (OQ)]
As previously reported by Litardière (1921), Langlet (1928) and Kuroki (1965), this species has basically 12 somatic chromosomes. But the karyotype seems to vary considerably by population. It has been known that a pair of large submedian chromosomes has submedian secondary constrictions, and in our materials, another pair of smaller subterminal chromosomes also has often submedian or subterminal secondary constrictions. In a wild population collected in Kashmir, one or two very small metacentric chromosomes or B-chromosomes are sometimes found. It is highly probable that in this population occur the similar chromosomal aberrations especially translocations to those observed in P. peltatum by Newman (1967).

Agrimonia pilosa Ledebour var. nepalensis (D. Don.) Nakai
2n=56. Gokarna, Kathmandu, C. Nepal, 1900 m (Sep. 1963).
2n=56. Dhunche, Gosainkund, C. Nepal, 2100 m (Jun. 5, 1,969).
2n=56. Akasay, E. Nepal, 2600 m (Dec. 1, 1963).
2n=56, Mane Bhanjang, Singalila Range, 2250 m (Jul. 22, 1969).
The chromosome number and the relationships between the Hirnalayan plants and the Japanese ones were discussed in detail by Hara & Kurosawa in Journ. Jap. Bot. 43: 392-400, pis. 20 & 21(1968).

Fragaria Daltoniana J. Gay
2n=14.Sandakphu, Singalila Range, 3900 m (Jul. 13, 1969).
This number agrees with that reported by Darrow (1937). It became apparent that all the four species occuring on the Singalila Range, F. nubicola lindley, F. nilgerrensis Schlecht., F. rubiginosa Lacaita and F. Daltoniana are diploid (cf, Kurosawa in Fl. E. Himal. 661, 1966).

Desmodium podocarpum DC, subsp. oxyphyllum (DC.) Ohashi
2n=22. Orchid Sanctuary, Gangtok, Sikldm, 1700 m (Jun, 27, 1969). [Pl.21. f (OQ)]
All the species of the genus so far examined cytologically have 22 somatic chromosomes, and Funabiki (1958) suggested the same number for the Japanese species but has never studied them. Sokolovskaya (1966) found 2n==22 in D. podocarpum subsp. oxyphyllum var. mandshuricum (Maxim.) Ohashi, and the author confirmed the same number in subsp. oxyphyllum.

Sarcococca Hookeriana Baillon
2n=28. Bhuspate Danra, E. Nepal, 2900 m (Dec. 2, 1963).

Berchemia flavescens (Wall.) Brongn.
2n=24. Langtang Gorge, C. Nepal, 2500 m (Jul. 7, 1970).

Stachyurus Sigeyosii Masamune
2n=24. Taroko, Formosa (Mar. 1966). [Fig. 59. a (OQ)]

As I have reported in 1966, both S. himalaicus Hook. f. et Thorns. of E. Nepal and S. praecox Sieb. et Zucc. of Japan have also 2n=24 chromosomes. Morphologically the Formosan plants are similar to S. himalaicus in some respects, but resemble S. praecox var. lancifolius (Koidzumi) Hara of southern Japan in the characters of leaves.

Panax pseudo-ginseng Wall. subsp. pseudo-ginseng
2n=24. Shioupuri, Kathmandu, 2500 m (Jun. 14, 1969) [Pl. 22. a (OQ)]

subsp. himalaicus Hara
2n=24. Phalut, Singalila Range, 3800 m (Jul. 18, 1969).
2n=24. Tonglu, Singalila Range, 3100 m (Jul. 12, 1969).
2n=24. Chitrey-Mane Bhanjang, Singalila Range, 2300 m (Jul. 22, 1969).
2n=24. Western Bhutan (Apr. 1967).

var. angustifolins (Burkill) Li
2n=24. Darjeeling, 2200 m (Apr. 1967) [Pl. 22. b, c (OQ)]
2n=24. Darjeeling, 2200 m (Jun. 23, 1969).
The Himalayan races have 2n=24 chromosomes, as reported by Kurosawa (ex Hara 1970). In my previous report (1966), 2n=ca. 48 for a plant from Tonglu was recorded by mistake. But I confirmed that the Japanese plants have 48 somatic chromosomes.

Aucuba chinensis Benth.
2n=16. ShishikyÔ, KÔshun, Formosa (Mar. 28, 1966).

Aucuba japonica Thunb.
2n=16. Mt. Katsadake, Okinawa (Jan. 12, 1969). [Pl. 22. d (OQ)]
2n=16. Mt. Yuwan, Amami-Õshima (Apr. 6, 1970).
2n=16. AnbÔ, Yakushima Is., 300 m (Feb. 1971).
A. himalaica Hook. f. et Thoms, and A. chinensis (Meurman 1930, Kihara & Yamamoto 1935) have been known as diploid, while A. japonica Thunb. as tetraploid (2n=32). It was unexpected that the plants of Okinawa in the Ryukyus, AmarniÕshima and Is. Yakushima are diploid, because they are rnorphologically more similar to typical A. japonica than to A. chinensis. Further studies will reveal the detailed relationship between diploid and tetraploid races in Japan.

Helwingia formosana Kanehira et Sasaki
2n=38. KeitÔ Expcrim. Forest, Formosa, 1300 m (Nov. 13, 1968), (Fig. 59. b (OQ)]
2n=38. Mt. Nankotaisan, Formosa, 1700 m (Aug. 20, 1969).
In my previous reports (1965 & 1966), it was proved that H. himalaica Hook. f. et Thorns, from Darjeeling has 2n=38 chromosomes, while H. japonica (Thunb.) Dietr. of Japan is a high polyploid with. 2n=ca, 114 chromosomes. It is noteworthy that the Formosan plants which bear black fruits have the same number as that of H. himalaica with red fruits, and clearly differ from the Japanese ones.


Lysimachia prolifera Klatt
2n=24. Sandakphu, Singalila Range, 3900 m (Jul. 15, 1969). [Pl. 22. e (OQ)]
2n=24. Garibas-Tonglu, Singalila Range, 2900 m (Jul. 21, 1969).

Lysimachia ramosa Wall.
2n=20. East Birch Hill Road, Darjeeling, 2100 m (Jul. 5, 1969). [Fig. 60. b]

Lysimachia evalvis Wall.
2n=30. Daijeeling, 2200 m (Jun. 23, 1969). [Fig. 60. a (W)]

Lysimachia debilis Wall.
2n=84. Chimakhothi, Bhutan, 2350 m (Apr. 21, 1967).
L. prolifera belongs to Sect. Gandidae Hand.-Mzt. which has never previously been studied cytologically. In Sect. Apodanthera, L. sikokiana Miquel was proved to have 2n=60 chromsomes (Jinno 1956), and taking the numbers of L. ramosa and L. evalvis above mentioned into consideration, the basic number of this section appears to be 5.

Ellisiophyllum pinnatum (Wall.) Makino
2n=18. Ritang Valley, Bhutan, 2000 m (Apr. 17, 1967).

Wulfenia nepalensis Yamazaki
2n=16. Phulchauki, Kathmandu, 2600 m (Jun. 26, 1967). [Fig. 60. c (OQ)]


Yamazaki considered that the species is closely allied to W. Amherstiana Bentham of Western Himalaya (see page 120). It is interesting that the chromosome number of this species is also the same as that of W. Amherstiana (n=8, Verma & Dhillon 1967; Mehra & Gill 1968), but differs from that of W. carinthiaca Jacquin of E. Europe (2n=18, Favarger & Huynh 1964; Kurosawa ex Yamazaki 1968).

valeriana Jatamansi Jones
2n=32. Yusmarg. Kashmir, 2100 m (May 16. 1970). [Fig. 60. d (OQ)]
2n=32. Godavari, Kathmandu, 1600 m (May 29, 1969). [Fig. 60. e (OQ)]

Valeriana pyrolaefolia Decaisne
2n=16. Gulmarg, Kashmir, 2400 m (May 14, 1970).
Mehra & Sobti (1955) reported 2n=32 under the name V. Wallichii DC., but Kishore (1951) found 2n=28 in the same species.

Peracarpa carnosa (Wall.) Hook. f. et Thoms.
2n=ca. 30. Phalut, Singalila Range, 3850 m (Jul. 18, 1969).
The plants of Phalut have roundish leaves, and agree with the Japanese ones in this respect.

Adenocaulon himalaicum Edgeworth
2n=46. Phulchauki, Kathmandu, 2800 m (Jun. 10, 1969).
2n=46. Ritang Valley, Bhutan, 2000 m (Apr. 16, 1967).
This is the first record on the Himalayan plants, and the chromosome number agrees with that reported on the Japanese plants by D. M. Moore (1963), Arano (1965), and Huziwara (1968). But different numbers such as 2n=20 (Arano 1960), 2n=38 (Arano 1962) and 2n=48 (Koyama 1966) have hitherto been published based on the Japanese plants.

Artemisia indica Willd.
2n=34. Darjeeling, 2150 m (Apr. 1960).
2n=36. Kalapokhari, Singalila Range, 3100 m (Jul. 21, 1969). [Pl. 22.f (OQ]
2n=36. Tiger Hill, Darjeeling, 2450 m (Jul. 26, 1969).

Artemisia dubia Wall. ex DC.
2n=54, Birch Hill Road, Darjeeling, 2100 m (Mar. 16, 1963).
Stoloniferous 'Mugworts' found very common in the lowlands of Japan, A.. indica and A. princeps Pamp., have 2n=34 chromosomes. It is notable that one population from Darjeeling, as already reported by Koyama ex Kitamura (1966) under the name A.. princeps, agrees with the Japanese plants in chromosome number, but two other populations from the same district have 2n=36 chromosomes. A. dubia with caespitose semiwoody stems is quite distinct from A. indica morphologically, and has 2n=54 chromosomes, which number agrees with that of Suzuka (1956). The Indian plants of the A. vulgaris complex with n=18 chromosomes reported by Khoshoo & Sobti (1958) are probably identical with A.. indica, and those with n=27 & 2n=54 chromosomes correspond with A. dubia Wall., as this species often occurs at lower altitudes.

Arisaema ostiolatum Hara
2n=28. Phalut-Sabargan, Singalila Range, 3400 m (Jul. 19, 1969).

Arisaema Jacquemontii Blume
2n=28. Tonglu, Singalila Range, 2800 m (Jul. 12, 1969).

Arisaema intermedium Blume
2n=28. Phulchauki, Kathmandu, 2900 m (May 29, 1969). [Pl. 23. a (OQ)]

Arisaema consanguineum Schott
2n=28. Kalingchok, C. Nepal, 2000 m (Sep. 10, 1970).

Aletris gracilis Rendle
2n=ca. 52. Pele La, Bhutan, 3000 m (Apr. 12, 1967).


Allium Wallichii Kunth
2n=14. Gosainkund, C. Nepal, 3000 m (Aug. 1969). [Pl. 23. f (OQ), Fig. 61]

There is a pair of large submedian chromosomes with distinct secondary submedian constrictions, and those chromosomes are liable to be detached at the constriction.

Clintonia udensis Trautv. et Meyer var. alpina Hara
2n=28. Sandakphu, Singalila Range, 3950 m (Jul. 15, 1969).

Disporum cantoniense (Lour.) Merrill
2n=14. Tonglu, Singalila Range, 3100 m (Jul, 12, 1969).
2n=14. Chimakhothi, Bhutan, 2350 m (Apr. 21, 1967).
2n=16. Phulchauki, Kathmandu, C. Nepal, (May 29, 1969).
2n=16. Tendong, Sikkim, 2900 m (Jun. 29, 1969). [Pl. 23. b (OQ)]
It became apparent that three cytotypes with 2n=14, 16 and 30 chromosomes occur in Eastern Himalaya.

Fritillaria cirrhosa D. Don
2n=24. Sandakphu, Singalila Range, 3900 m (Jul. 14, 1969).
This number agrees with that reported by La Cour (1951).

Paris polyphylla Smith
2n=20. Singhik, Sikkim, 1500 m (Jun. 30, 1969).
2n=20. Badam La, near Kalimpong, 1500 m (Jul. 6, 1969).
2n=20. Ritang Valley, Bhutan, 2000 m (Apr. 17, 1967).
2n=20. Samtengang-Ritang, Bhutan 2700 m (Apr. 11, 1967).
2n=20. Chmakhothi, Bhutan 2350 m (Apr. 21, 1967).
2n=20. Darjeeling, 2150 m (Apr. 1967).
2n=20. Tonglu, Singalila Range, 3100 m (Jul. 11, 1969).

subsp. marmorata (Steam) Hara
2n=10=2Am+2Bsm+2Cst+2Dst+2Esm. Pele La, Bhutan, 3000 m (Apr. 17, 1967). [Pl. 23. d (OQ)]
2n=20. Pele La, Bhutan, 3000 in (Apr. 17, 1967).
It is interesting that tetraploid races prevail in Eastern Himalaya, as suggested before (Kurosawa 1966; Hara 1969).

Polygonatum graminifolium Hooker
2n=30. Gosainkund, C. Nepal, 2900 m (Jun. 4, 1969). [Pl. 23. e (OQ)]


Polygonatum singalilense Hara
2n=ca. 46. Phalut, Singalila Range, 3900 m (Jul. 18, 1969).
The Himalayan species especially those of the P. verticillatum group are exceedingly polymorphic, and are also variable in chromosome number (see Hara on page 170).

Plate 12. c. Polygonatum singalilense


Streptopus simplex D, Don
2n=16. Sandakphu, Singalila Range, 3900 m (Jul. 15, 1969). [Pl. 23. c (OQ)]
Streptopus simplex [Pl. 11. a] characterized by axillary filiform peduncles and cam panulate white flowers is the only representative species of the genus in the Hima layas, and has never previously been studied cytologically. It is a diploid with. 2n= 16 chromosomes, and its karyotype is similar to that of S. roseus (Bent & Smith 1969).

Plate 11. a. Streptopus simplex

Tricyrtis maculata (D. Don) Macbride
2n=26. Gosainkund, C. Nepal, 2800 m (Aug. 1969).
This number agrees with that reported by Miller, E. W. (1930) and Subramanyan & Kamble (1967) under the name T. pilosa Wall. All the species of the genus so far examined cytologically are diploid with 2n=26 chromosomes.

Trillium Govanianum Wall. ex Royle
2n=20=4Am+4Bsm+2C1st+2C2st+4Dst+4Esm. Sandakphu, Singalila Range, 3900 m (Jul.14, 1969). [Pl.24. a (OQ)]

2n=20. Gulmarg, Kashmir, 2400 m (May 14, 1970).


Trillium Tschonoskii Maxim. var. himalaicum Hara
2n=20=4Am+4Bsm+4Cst+4Dst+4Esm. Pele La, Bhutan, 3200 m (Apr. 17, 1967). [Pl. 24. b (OQ)]
The genus Trillium is represented in the Himalayas only by the two species above mentioned. Their chromosome number and karyotype agree with those reported by Haga & Watanabe ex Hara (1966 & 1969) based on the collections of our pre vious Expeditions. It is noteworthy that in T, Govanianum a pair of the chromosomes with a subterminal centromere are satellited.

Iris Clarkei Baker
2n=38. Tonglu, Singalila Range, 3150 m (Jul. 22, 1969). [Pl. 24. f (OQ)]
Although Simonet (1934) found 2n=40 in I. Clarkei, I confirmed the number 2n=38, as reported in 1966.

Plate 5. a. Iris Clarkei

Iris decora Wall.
2n=ca, 36. Near Kyapcha, Bhutan, 2400 m (Jun. 1, 1.967).
It is remarkable that I decora (I. nepalensis) belonging to subgen. Nepalensis is variable in chromosome number and pollen type. For the group including I. Collettii, such numbers as 2n=24, 28, 30, 40 have been published (Larsen 1963).

Iris Hookeriana Foster
2n=24. Sonamarg, Kashmir, 2600 m (May 13, 1970). [Pl. 24. d (OQ.)]

Iris sp.
2n=32. Near Lingshi, Bhutan, 3400-4000 m (May 22, 1967). [Pl. 24. e (OQ)]
I. Hookeriana and I. kamaonensis belong to subsect. Pseudoregelia of sect. Hexapogon, and Simonet (1952), La Cour (1955) and Malik (1961) reported 2n=22 and 24 for I. kamaonensis.

Iris japonica Thunb.
2n=36. Darjeeling, 2150 m (Apr. 1960). [Pl. 24. c (OQ)]
It is well-known that typical I. japonica of Japan is triploid with 2n=54 chromo somes. The plants cultivated at Darjeeling are morphologically similar to I. japonica, but have 2n=36 chromosomes and pollen grains with a different mode of reticulum on the surface from both I. japonica and I. formosana (cf. Chuma in Journ. Jap. Bot. 45; 281, t. 23, 1970). This interesting race may be the same as that described as 'Iris sp. (Chinese origin)' by Yasui (1939), and also identical with that reported by Sharma & Talukdar (1960).


LITERATURE CITED


Arano, H. 1962.
Bot. Mag. Tokyo 75: 356-367.
——— 1963.
Bot. Mag. Tokyo 76: 459-465.
——— 1965.
Jap.Joum. Bot. 19: 31-67.
Bent, A, E. and E. C, Smith 1969.
Canad. Journ. Gen. Cytol. II; 409-413.
Haga, T. and H. Watanabe ex Hara 1.966.
Flora of Eastern Himalaya 418
——— and ——— ex Hara 1969.
Journ, Jap. Bot. 44: 374-375.
Hara, H. 1969. Journ. Fac. Sci. Univ. Tokyo Sec. III, 10 (10); 141-180.
——— 1970.
Journ.Jap. Bot. 45; 197-212.
Hara,H. and S. Kurosawa, 1965.
Journ. Jap. Bot. 40; 36-40.
——— and ——— 1968.
Journ. Jap. Bot. 43: 392-400.
Huziwara, Y. 1968.
Kromosomo 72-73; 2360-(2363).
Jinno, T. 1965.
Jap. Jour. Genet. 31; 87-88.
Khoshoo, T. N. and S. M. Sobti 1958.
Nature 181: 853-054.
Kihara, H, and Y. Yamamoto. 1935.
Agric. and Hortic. (Tokyo) 10; 2485-2496.
Koyama, S. 1966.
Act. Phytotax. Geobot. 22: 80.
Kuroki, Y. 1965.
Mem. Ehime Univ. Sect. II, Ser. B(Biol.) 5; 97-102.
Kurosawa, Sachiko. 1966.
Hara, Flora of Eastern Himalaya 658-670.
La Cour, L. F. 1951.
Heredity 5; 42.
——— 1955.
Darlington & Wylie, Chromosome Atlas of Flower. Pls. ed. 2, 387.
Langlet, O. F. J. 1928.
Svensk Bot. Tidskr. 22: 169-184.
Larsen, K. 1963.
Dansk Bot. Ark. 20: 261.
Meurman, O. 1930.
Mem. Soc. Fauna Flora Fennica 6; 95-100.
Moore, D. M. 1963.
Madroño 17: 52-53.
Newman, L. J. 1967.
Chromosoma 22: 258-273.
Pauwels, L. 1959.
Bull. Soc. Roy. Bot. Belg. 91: 301.
Sharma, A. K. and C. Talukdar. 1960.
Genetica 31: 340-384.
Simonet, M. 1934.
Ann. Sci. Nat. Bot., Ser. 10, 16: 229-388.
———1952.
Compt. Rend. Acad. Sci. (Paris) 235 (21): 1244-1246.
Subramanyam K. and N. P. Kamble, 1967.
Taxon 16: 341-350.
Suzuka, O. 1956.
Land and Crops of Nepal Himalaya 2: 243, pls. 1-2.
Yamazaki, T. 1968.
Journ. Jap. Bot. 43: 406-408.
Yasui, K. 1939.
Cytologia 10: 180-188.


ERRATA


Kurosawa in Hara, Flora of Eastern Himalaya (1966)
Page 669, line 1.   read   2n=36=6Am+12Bst+10Cst+8Dm
for   2n=36=3Am+6Bst+5Cst+4Dm




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