VIR Jl~, UPPEAI~DRA DHAR and P. KACHROO

Department of Botany, University of Kashmir, Srinagar - - 190 006, India

Contribution to the Phytogeography of Kashmir Himalaya I. Ranunculaceae and Paeoniaceae

K e y w o r d s

Floristic composition, Extant links, Endemism, .Ranunculaceae, Kashrnir Himalaya

A b s t r a c t

J~.E V., DHA~ U. et KACHROO P.: Contribution to the Phytogeography of Kashmir Himalaya I. Ranunculaceae and Paeoniaceae. -- Folia Geobot. Phytotax., Praha, 24: 387--402. -- The affini- ties and endemism of Kashmir Himalaya t~anunculaceae and Paeoniceae reveal some interesting distribution patterns in some taxa. Isoflor maps, revealing the affinities of Aconitum, Aquilegia, Delphinium and Ranunculus, are given. A high degree of endemism exists in the study area. About 41 ~ of the total taxa show restricted distribution which is discussed. The floristic relation- ships of the wldes show close links with Central Asia than any other phytogeographical unit. The significance of altitudinal amplitude of some taxa vis-a-vis their distribution pattern is discuased.

I N T R O D U C T I O N

K a s h m i r H i m a l a y a is p r e d o m i n a n t l y a h igh m o u n t a i n o u s a r ea a n d affords in te res t ing scope for p h y t o g e o g r a p h i c s tudies . I t is s i t ua t ed be tw e e n 3 3 - - 3 6 ~ N l a t i t u d e a n d 7 2 - - 9 0 * E long i tude ; occupies an a r ea of a b o u t 222.8 sq. k m in which mos t of the l a n d lies over 3,000 m (Fig. 1). The h ighes t peak ascends to 5,500 m, t h e r e b y offering s t rong i so la t ion which a p a r t f rom being geograph ica l also seems to be ecological as well (STOTT 1980).

The p h y t o c h o r i o n ( K a s h m i r H i m a l a y a ) cons idered in t h e p re sen t s t u d y is confined to the phys i ca l bounda r i e s o f K a s h m i r and L a d a k h ; however , i t a lso inc ludes the a rea o f n o r t h P a k i s t a n on a c c o u n t of i ts v e g e t a t i o n a l s imi lar i t ies (STEwAgT 1972). The a r ea is su r rounded b y China in the east , P a k i s t a n in t he west , U S S R - - China in t h e n o r t h and the I n d i a n P la ins in the south .

Cl imat ica l ly , t he a r ea exh ib i t s a g r e a t d i v e r s i t y r ang ing f rom sub t rop ica l -

388 F O L I A G E O B O T A N I C A E T P H Y T O T A X O N O M I C A 24, 1989

-temperate and high alpine tundras to the cold arid conditions of Ladakh. Two high altitude passes separata the valley from these climatic extremes, namely the Zoji La Pass (3,400 m) situated in the northeast and the Pir Panjal Pass (3,200 m) in the southwest direction. The former checks the influence of cold dry conditions while the latter that of subtropical north Siwaliks. Perhaps the marked climatic diversity provides ample opportunities for speciation in this geographically young area.

No doubt, during the past two decades data on floristics and phtogeography has accumulated fast yet certain areas are underexplorcd. Ladakh Expedition Report (1982) sufficiently proves this point by putting on record 20 species new to Ladakh. Besides, many species have been added to the general flora of this area from time to time. Thus, on the one hand, new endemics are constantly being added while thorough systematic research and exploration keeps on reducing the endemic status of many other taxa. This insertion/elimination of endemics usually maintains a balance in almost every active phytochorion.

Although a fairly good account of the post glacial vegetational history of Kashmir is available (S~.H~I 1963; WADIA 1975; PURI 1933, 19~:7; VIS~Ntr-MITTTR~ 1963, 1964) yet the knowledge of the extinct flora is too limited to be of any major significance in assessing the nature and extent of endemism.

ENDEMISM IN KASHMIR HIMALAYA

The only detailed treatise on the extant endemics of the Indian subcontinent is that of CHATT~,RX~. (1939), who puts on record 61.5 % of the dicots as endemics. The highest concentration of endemics is encountered in the Himalaya (28.8 %) which is attributed to geographical isolation caused by the lofty mountains. TAKH- TAJAX (1969) listed about 150 endemic genera in the Indian Floristic Region (which includes peninsular India upto the flanks of the Himalaya and to Sri Lanka); RAo (1972) recognised 164 endemic genera in the same region (including Burma and Ceylon) and NAu (1980) records 141 endemic genera exclusively within the boundaries of India in which 111 genera are monotypic. Of these monotypic genera, 68 are confined to the Himalaya (40 in the eastern, 11 in the western and 17 throughout the Himalaya). MAxI (1978) believes that 3,000 dicot and 1,000 monocot species are endemic to the Himalayan region.

KAXAI (1963, 1966) published an extensive review of the phytogeography of gapano-Himalayan elements and of the eastern Himalaya respectively. BANERJI (1963) analysed the flora of Nepal Himalaya which was later supplemented by ])O~ERMEZ (1972). I-I~DG]~ and WW~DELBO (1972) recognised 23 endemic genera in the Afghanistan flora and specific endemism studied in 9 important families ranged from 23 % (Brassicaceae) to 100 % (Ericaceae). Later BRV.CKI~ (1974) recorded 20.2 ~ endemic species from alpine and nival flora of the Hindukush mountains. BHARGAVA (1983) reports 54 endemic species in Poaceae of northwest Himalaya, many of which are endangered. Recently DEXR et al. (1987) in their comprehensive review on endemism in Kashmir Himalaya emphasised the con- servation and preservation of many endangered taxa and their habitats.

The detailed phytogeographie studies on west Himalayan flora are still frag- mentary, however, some notable contributions include: M~USEL (1971) and M_EUS]~L

V I R J E E E T AL.: P H Y T O G E O G R A P H Y O F K A S H M I R H I M A L A Y A 389

and SCHVa~RT (1971) on the coUine, montane and subalpine vegetation of the western Himalaya followed by MAgi (1978) on the high alti tude flora of northwest Himalaya. DHxa and :KACHROO (1982; 1983a, b) dealt this aspect critically with respect to the alpine-subalpine flora of l ~ h m i r Himalaya (Poaceae excluded). H~tTMAN~ (1966, 1968, 1972) contributed significantly to the floristic analysis and ecological peeularities of the Karakorum flora. S ~ W ~ T (1916), KAcmr et el. (1977) and HARTMA~ (1984) investigated the flora of Ladal~h while SEYBOLD and KULL (1985) confined their studies to the floristics and vegetation of Zanaskar. t t ~ T M A ~ (1983) also a t tempted synecological studies en route from Kashmir to Ladakh and more recently (HARTMA~ 1987) he studied the community characteristics of xerophytic subalpine and alpine species in the southern and eastern part of Ladakh. Cytological studies have also been initiated to provide a bet ter understanding of the biogeographical character of Kashmir Himalayan taxa (Vm J ~ et el. 1983, 1984, 1985, 1987). The present a t t empt is, however, aimed at providing a family wise precise account of the extent of endemism and affinities of this phytoehoriou.

T a b l e 1 . 1 ) i s t r i b u t i o n o f e n d e m i c t a x a (Ranunculaceae) i n K a s h m i r H i m a l a y a

I~ )ca l a r e a .No. of endemic taxa

Kashmir 3 Ladakh 3 Kashmir -- Ladakh 0 Kashmir-- north Pakistan 8 Ladakh-- north Pakistan 3 Kashmir-- Ladakh-- north Pakistan 3 Total 20

ENDEMISM IN .RANUNCULACEAE

Ranunculaceae is practically confined to the north temperate hemisphere (GoOD 1964) whereas in India, it is mostly distributed in the Himalaya. In Kashmir Himalaya the family is represented by 98 taxa in 17 genera ranging from 350--5,500 m (Table 3). Ranunculus is the largest genus (28 taxa) followed by Thalictrum (12 taxa). Six genera are monospecifie in the s tudy area, namely, Actaea, Caltha, Cimicifuga, Consolida, Pulsatilla and Trollius. Of the 98 taxa, 20 are endemic (20.4 %) in which 13.25 ~o shows specific and 7.14 ~ infra-specific endemism. These endemic taxa further reveal the following distribution pat tern (Table 1).

I t is interesting to note that only three taxa are represented throughout the natural area while the remaining 17 lie confined to one or two localities (Figs. 1, 2). Of the 78 wide taxa, 20 (25.64 %) are distributed in the Himalaya including parts of east Afghanistan and Tibet. In general 40 taxa (40.71%) of the total Kashmir :Himalayan Ranunculaceae remain confined to the Himalaya (including Afghanistan

390 FOLIA GEOBOTANICA ET PHYTOTAXONOMICA 14, 1989

a n d T ibe t ) whereas the res t serve as wides b e y o n d th i s range. I n t he 20 r e s t r i c t e d t a x a m e n t i o n e d a b o v e t h e fol lowing local d i s t r i b u t i o n p a t t e r n is d i s ce rnab le (Table 2). The e x t e n t o f e n d e m i s m in Ranunculaceae a n d Paeoniaceae of K a s h m i r H i m a l a y a a n d the ad jo in ing a reas is g iven in Tab le 3. Aconitum shows m a x i m u m d o m i n a n c e i n Cent ra l Asia (51 t a x a ) t h a n E u r o l ~ (14 t a x a ) b u t t he reverse is t r u e i n so fa r as the endemic pe rcen t age is concerned (Table 3). However , i t is no te -

Fig. 1. Total known distribution o f . A~ruilegia nlvalis FAI~. ex JACKSON - - a characteristic endemic species of the Kashmir Himalaya.

Table 2. Distribution of restricted taxa (Ranunculaceae) of the Kashmir Himalaya

Local range No. of restricted taxa

Kashmir Himalaya--western Himalaya Kashmir Himalaya--western and central Himalaya Kashmir Himalaya-eastern Himalaya

Total

12 4 4

20

V I R J E E E T AL. : P H Y T O G E O G R A P H Y O F K A S H M I R H I M A L A Y A 391

worthy tha t 58.82 % of the endemic Central Asian species of A c o n i t u m are either new species or new combinations. In Kashmir Himalaya the endemic percentage is no less significant (50.00) while it reaches max imum in the eastern Himalaya (100 %). Genera like Actaea, Geratocephalu~ and Cimic i fuga are more or less widely distributed and hence not represented by endemics. Conversely Consolida and Pulsat iUa reveal 100 % endemism in Kashmir Himalaya. The endemics in

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A d o n i s (25 %) and Caltha (14.28 %) are known only in Europe and Central Asia respectively. The m a x i m u m number of t axa of Aqui leg ia (29) ,in Europe corresponds to its max imum endemic percentag e (72.41). The same is the case with A n e m o n e in Central Asia, but the reverse for Cal l ian themum. The only species of D e l p h i n i u m in eastern Himalaya is endemic, however, specific endemism of the genus is signifio cant in Soviet Asia (61,72 %). In Kashmir Hima laya 25 % endemics exist in this genus. Paraqui leq ia - - described relatively recently - - has a m a x i m u m of five t axa in Afghanistan (40 % endemics). The genus is nat known from southeast Asia and Europe. Clematis , despite being well represented in Central Asia, has insignificant endemics (5.55 %).

392 F O L I A G E O B O T A N I C A ]~T P H Y T O T A X O N O M I C A 24, 1989

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A F F I N I T I E S IN R A N U N C U L A C E A E

The interesting features of affinities revealed by Kashmir Himalayan Ranun- culaceae are given in Fig. 4. Aconitum shares 80 % wides with Pakistan; 40 % with USSR and 20 % each with Afghanistan and Europe. No doubt 60 % of the wides extend up to China but none beyond this area (Fig. 5). Actaea, Caltha, Cimicifuga, Adonis and Ceratocephalus range widely in all the adjoining phyto- geographic units; however, the representatives of the lat ter two genera are not known from the Sine-Japanese region. In Anemone the affinities are more marked with West-Central Asia (71.42 %) than with Sine-Himalaya (42.85 %) while only 14.28 % taxa are common with the remaining phytochoria. Amongst restricted genera, Consotida and Pulsatilla lie confined to the s tudy area; Trolliu~ extends up to the west H ima laya while Aquilegia -- up to western and central Himalaya (Fig. 6). Of the two species in Callianthemum, C. alatavicum FR~.YN is distributed in west-central Asia while C. pimpinelloides (D. DoN ex ROYLE)H. et T. shows a Sine-Japanese link. Clematis leans more heavily towards the Sine-Japanese region (71.42 ~ ) and beyond the western limits of Afghanistan, the relationship tends to decline (28.56--14.28 %). The wides in Delphinium manifest an almost similar decreasing trend in their extensions (100--16.66-%) in all neighbouring regions

394 F O L I A G E O B O T A ~ I C A E T P H Y T O T A X O N O M I C A 24, 1989

(Fig. 7). Paraquilegia displays 100 % affinities with west-central Asia and parts of Sine-Japanese Region (China). Ranunculus and Thalictrum reveal almost similar extensions with the adjoining parts of the study area; however, the former shows stronger links with Central Asia (Fig. 8) and the latter with China (83.33 %).

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Fig. 8. Affinit ies o f K a s h m i r H i m a l a y a n t a x a o f Ilanunculus.

396 F O L I A G E O B O T A N I C A E T P H Y T O T A X O N O M I C A 24, 1989

ENDEMISM AND A F F I N I T I E S IN PAEONIACEAE

The genus Paeonia is monotypic in the western Himalaya and is represented by 15 and 17 taxa in Soviet Asia and Europe respectively. The endemic percentage is maximum in Central Asia (46.66 %) followed by Europe (23.52 %); whereas in the Kashmir Himalaya the lone species is a broad range endemic.

DISCUSSION

The s tudy of endemism is not only useful in assessing the origin and ant iqui ty of a taxon but also plays an important role in realising the status of the past vegetational history. PoLuN~ (1960) believed tha t relic endomics are useful in indicating antiquity, isolation and the diversification of habitats, while Good (1964) held that endemics may be more useful in the recognition of different floristie units. A~sARI (1984) stressed the occurrence of restricted taxa in a bio- logical co!nmunity in view of their sensitivity to even minor habitat differences. The geological and geographical changes axe, however, also essential for bet ter understanding of the nature of endemics in a particular region. The changes brought about by the Pleistocene glaciation have largely affected the distribution pat tern of many t a r a in the Kashmir Himalaya.

Ranunculaceae shows dominance in the U.S.S.R. (502 ta ra ) followed by Europe (418 taxa). The family is almost equally represented in the western Himalaya (77--105 ta~xa) while in the eastern Himalaya, the representation is very feeble (37 tara) . A comparative account of the number of taxa in various genera of Kashmir Himalaya vis-a-vis their areas of major preponderance implies: Aconitum (10 : 51); Actaea (1 : 3); Adonis (3 : 12); Anemone (11 : 46); Aquilegia (5 : 29); CaUianthemum (2 : 4); Caltha (1 : 7); Ceratocephalus (2 : 3); Cimicifuga (1 : 5); Clematis (8 : 18); Consolida (1 : 16); Delphinium (8 : 81); Paraquilegia (3 : 5); Pulsatilla (1 : 26); Ranunculus (28: 170); Thalictrum (12 : 28); and TroUius (1 : 11). Afghanistan represents the area of dominance for two genera (Ceratocephalu8 and Paraquil~ia); Europe for 4 (Adonis, Aquilegia, Thalictrum and Consolida) while USSR for the remaining 11 genera of the study area.

Being a young mountain system, the Kashmir Himalaya, as expected, harbours a relatively lower concentration of taxa. However, it is noting that Ranunculu~% although it is a well adapted genus with a varied altitudinal range (450--5,500 m) shows meagre representation (28 ta~xa; 16 ~ of the maximum area of occurrence) as against other weedy genera -- Ceratocephalus (66 ~ Adonis (25 %) and Anemone (25 ~ etc. Fur ther the monotypic nature of Consolida, Pulsatilla and Trolli~ perhaps indicates their recent arrival, or an unfavourable environment for their speciation. Among the other genera of the neighbouring areas, Paroxy- graphis is distributed in the eastern Himalaya; He~emone, Enemion, Semiaquilegia, Trauvetteria and Leptopyrum, in the USSR and HeUeborus, Hepatica, Myosurus, Eranthis and Niqella, in the USSR and Europe (the lat ter two genera are also present in Afghanistan). Further, Coptis, Batrachium, Ficaria, Oxygraphis and Itelerpestes represent valid genera in the USSR but are covered under Ranunculu~ in Europe and Kashmir Himalaya. Similarly Atraqene considered valid in the USSR, is reported as a synonym of Clematis in Europe and Kashmir Himalaya

V I R J E E ET AI,.: P H Y T O G E O G I ~ A P H Y OF K A S H M I R H I M A L A Y A 397

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400 F O L I A GEOBOTA~NICA E T P t t Y T O T A X O . ~ 0 ~ [ I C A 24,1989

and of Naravelia in the eastern Himalaya. GarideUa is monotypic in Europe but included under ~Viqella in the USSR. Most of these genera are mono- or bitypic and do not display any significant endemism. The overall degree of endemicity in the Kashmir Himalaya and adjoining areas reads: 20.40 % (Kashmir Himalaya); 3.80 % (Pakistan); 20.77 % (Afghanistan); 31.47 % (USSR); 34.92 % (Europe) and 37.83 % (Eastern Himalaya).

The family, essentially quite old and diverse, envisages many phytogeographic pecularities within the s tudy area. These considerations are usually discernible at both intra and intergeneric levels with the result tha t no positive correlation is ordinarily conceivable. In some genera -- Aconitum (2,100--4,800 m); Aquilegia (1,800--4,800 m); Delphinium (600--4,800 m), etc. altitudinal amplitude appears to have direct bearing on the distribution pat tern of their constituent taxa irrespective of their occurrence in different ecological zones. Thus one might logically at tr ibute the endemic status of taxa like Aconitum deinorrhizum (3,400 m) ; A. kashmiricum (2,400--3,300m); A. moschatum (3,300--3,900m); Aquilegia nivalis (3,300--4,200m); Delphinium pyramidale (2,100--3,000m); D. roylei (1,800--2,700 m); C ol~solida schl agintweitii (2,300--2,500 m); PulsatiUa wall ichiana (3,300--3,900m); and TroUius acaulis (3,000--,~,900m), etc. to their narrow altitudinal range which offers them limited scope of diversification. Conversly Actaea (2,100--3,300 m); Adonis (300--3,900 m); Caltha (2,400--3,900 m); Cerato- ccphalus (350--3,000 m); Cimicifuga (2,100--3,600 m); Clematis (900--4,200 m) and Paraquilegia (2,400-5,100 m) owing to their wide ecological range are characterised by greater adaptibility and tolerance and are, therefore, widespread in distribu- tion. Perhaps this may also be responsible for their significant representation in the study area vis-a-vis neighbouring floristic units. In addition, habitat specificity also appears to be a characteristic feature of many typical endemics and such a correlation becomes apparant even at the interspecific level within a genus. Thus, for example, Aconitum laeve (2,400--4,000 m) grows on diverse habitats and is widely distributed whereas A. kashmiricum (2,400--3,300 m) is a peculiar plant of alpine tundra and hence an endemic.

To sum up, it could be safely said that owing to the remarkable degree of phytogeographic pecularity in the family, no single factor can be considered as decisive in determining the "potential area" occupied by a particular taxon. The degree of endenficity is quite considerable, having evolved in the course of t ime following speciation in an invading flora. The extant links reveal more pronounced affinities with West-Central Asia than with the Sine-Japanese and other neighbour- ing phytogeographic units.

A c k n o w l e d g e m e n t

The authors are thankful to Prof. (Dr.) S. FAROOQ for his useful comments.

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R e c e i v e d 30 Oc tobe r 1987