BULL. BOT. SURV. INDIA

Vol. 27, NOS.1-4 : pp. 177-196, 1985

SYSTEMATICS OF THE GENUS L. - ( JUSS.) - ITS INF'RAGENERIC CLASSIFICATION AND EVOLUTIONARY TRENDS

Botanical Sum of India, Howrah

ABSTRACT

The genera J. Hill, Pmtaph@oi&s Duhamel, m. ilkeg* (=D&@a Rafin.1 and Dwksnea Smith are treated as congeneric synqnyms of Potantilla L. smr. strict0 ; nine sub- genera are recognised with keys to their ~denhficat~onand nomenclatural set up. The origin of the genus Potmtilla L. is discussed and evolutionary trends within the subgencra hypothesized.

INTRODUCTION ARGENTINAJ. Hill vis-a-vis POTENTILLAL. Linnaeus (I753, I 754) described the genus Rydberg (1906, 1go8), followed by Hut- Potentilla L. (accredited with 22 species), and chinson (1964) treat Argentina J. Hill (as together with two other related genera, 'Lam.'), lectot~pified by the perennial Comarum L. (with I species) and Fragaria herbaceous species A. anserina (L.) Rydberg L. (with 3 spp.), attributed them to his (Potentilla anserina L.) as generically &is- class "Icosandria Polygynia". Simultaneouslv, tinct from PotentiEEa L. he assigned another related genus ~ibbaldia The discovery of primitive phenolic con- L. (with z spp.) to his class "Pentandria stituents (both elagitanins and the trihyd- Pentagynia7'. These genera are assigned roxy phenolics p~ricetin and leucodel- to-day to the family Rosaceae Juss. (1789), phinidin) in PotentiZla anserina L. (Bate- subfarn. Rosoideae, -tribe Potentilleae Juss., Smith, 1962) led Challice (1973) to visualise subtrib. Potentillinae. Rydberg (1908) and P. anserina as the most primitive member of Hutchinson (I 964) recognised 16 genera the subfam. Rosoideae in terms of chemical within the tribe Potentilleae, of which characters. Wolf (1908) also considered his species belonging to eight genera, viz. pex Anserinae Wolf (with P. anserina, Argentina J. Hill, Chamaerhodos Ledeb., amongst other taxa) as an old group from Comarum L., Duchesnea J. E. Smith, its circumpolar distribution extending way Frogaria L., pentuphylloides Duhamel nom. back to the Tertiary period. These may illeg. superfl. ( =Wpbora Rafin.), Potentilla justify segregation of Argentina J. Hill as a L. and Sibbaldia L., occur in India. While distinct genus based on A. anserina (I,.) we treat Sibbaldia L.,as generically distinct Rydberg. (Dikshit and Panigrahi, 1~81b),we reduce But in terms of comparative morphological Argentina J. Hill, Pentaphylloides Duhamel, characters such as primitive versus advanced and Duchesnea J. E. Smith as congeneric characters, Potentilla anserina L., with its with and synonymous to ~otentillaL., for herbaceous habit, rod-shaped styles, glabrous reasons as discussed hereunder : achenes coupled with its higher polyploid --- cytodemes (2n=28, 35, 42) and apomictic Date of receipt : 24.6.86. Date of accmtance : 11.8.86 nature, must be treated as an evolut~onary *Present address : Central Sericulture Institute, Kora~ut* advanced group within the genus Pot~tillt~L. Bhudan Colony, Koraput, Orissa. 178 BULLETIN OF THE BOTANICAL SURVEY OF INDIA [Vol. 27

As a member of the subgenus Cheno- entire leaflets and hairy ...." and that potentilla (Focke) Juzepczuk, P. anserina "these shrubs are completely unable to cross L., has affinity with P. leuconota and with any other section ...." This group is P. coriandrifolia, amongst others, in its isolated from other taxa of Potentilla for a ~erennial herbaceous habit, im~aripinnate long time and the present distribution of , rod-shaped styles and glabrous the taxa indicates that they are old well- achenes ; and with P. indica, in its perennial established types. Sojak (1969) supported herbaceous creeping habit, rod-shaped styles the idea of treating the shrubby species of and glabrous achenes. PotentiZZa as a distinct genus, because they possess alternate cotyledonary leaves, the DASIPHORARafin. vis-a-vis POTENTILLAL. primary leaves are never rosette and the stipules are ochreate, whereas, in other Schwarz (I 949), Lijve (I 954). Hutchinson species of Potentilla L., they first form roset- (1964) and Sojak (1969) considered Penta- tes and the stipules are not ochreate. phylloides Duhamel (1755) as a genus However, the genus Dasiphora Rafin., distinct from Potentilla L. and made several shares its (i) shrubby habit and absence of combinations at species rank. But the rosette leaves with Potentilla palustre (L.) genus has to be rejected as a superfluous Scop., P. salesoviana Stephan, P. sericophylla illegitimate name, in as much as Duhamel Parker and P. xylorhiza Boiss. & Hohen., (c.), (although he referred only to the but the leaflets of P. palustre and P. saleso- woody species of Potentilla L., under Penta- viam are serrate ; (ii) its entire leaflets with phylloides Duhamel) included Potentilla L. P. biflora and P. armerioides ; (iii) its hairy (not Potentilla L. p.p.) as an obligate syno- achenes with P. salesoviana, P. sericophylla, nym. (Art. 7. I I ; Art. 63. I, ICBN 1983). In P. cuneifolia, P. eriocarpa, P. curmeseta, P. the circumstances, Dasiphora Rafin. (1838) pteropoda and P. collettiana and in P. typified with D,+ria Rafin., nom. illeg., biflora, P. palustre and P. armerioides the is the next available correct name at generic achenes are glabrous ; (iv) its basic chromo- rank, if one were to consider the genus as some number x=7 (2n=14, 28) with all distinct from Potentilla L. sens. stricto. species of Potentilla L., studied to-date. Schwarz (LC.), Love 1 c.), Hutchinson The geographical distribution of various (1.c.) and Sojak (I.C.), as stated above, con- species of Dasiphora Rafin., is no more dis- sidered the shrubby Potentilla species with continuous than some other species of woolly hairy receptacles and hairy achenes Potentilla, such as P. biflora L., which as constituting a separate genus, Penta- occurs in Europe, Asia and North America. Phylloides Duhamel ( =Ihdphra Rafin.) on We, therefore, opt here to follow Wolf morphological, anatomical, chemical, cyto- (1908) in treating the Fruticosae Potentillas logical evidence and on geographical distri- as belonging to the genus Potentilla L., but bution. Liive (Z.C.), who follows Schwan segregate the species involved into two sub- (c.), observed that "taxonomically the genera, viz. subg. Dasiphora and subg. Fruticosae group doubtlessly deserves the Tricho'thalamus. rank of a separate genus, distinct from Potentilla even in its wider sense ...." and DUCHESNEAJ. E. Smith vis-a-vis that of all sections of Potentilla, the most distinct, . both morpholo~callyand anatomi- POTENTZLLAL. cally, is the group Fruticosae which includes Wolf (I go8), Bate-Smith (I g61), Kalkman pkennial shrubs, pinnately leaved with (1968) and Acharga Goswami (1972) adduce 19gsI PANIGRAHI AND DIKSHIT ; SYSTEMATICS THE GENUS PO~WTILLA L. 119 morphological, -chemical and genetical J. D. Hooker (1878) which conform with all evidence in support of reduction of Duches- the above characters are segregated and nea J. E. Smith (1811) as with treated as Sibbaldia L. pixit and Panigrahh PotentzUa L. (1753), because : 198I b).

(i) The morphological difference between INFRAGENERIC CLASSIFICATlON OF the two genera concerns only the character Yo-rmriLLAL. WNSU STRIGTO of the torus : torus subcarnose and more or After a review of literature and critical ; less edible-Duchesnea torus not very study oi specimens we have iormulated the much enlarged and not edible-Potentilla. following classlhcation of Potentzlla L. sens. However, such a difference in the dimension s~rzcto,which recognlses g subgenera and a of the torus attained only after anthesis in number of sectlons to lnclude tne 80 lndlan Duchesnea, is merely cpantitative in nature. taxa dealt wirh, withln the systematic (ii) Chromato~raphically,Bate-Smith (1961) account of the genus : grouped Duchesnea indica with many other species of Potenrilla L., because he did I. Subg. DASIPHORA(Rafin.) Panigr. et Dik- not find any consistent differences between shit, comb. nov., et stat. nov. Dasi$hora the two genera. Rafin. in Aut. Bot. : 167. 1838 ; Rydberg (iii) Genetically, Potentilla and Duches- in N. Amer. F1. az(4). 366. 1908 ; nea belong to one compari~~m(Wolf, 1908 ; Juzepczuk, in Komarov's F1. URSS ro : Acharya Goswami, 1972) ; when Duchesnea 52. 1941 (Eng. edn. : 68. 197 I). Lectotype indica (male) was artificially crossed with sp. : Daszphora riparia Rahn. nom. illeg. Poientilla repans (female), the F, , [selected by Rydberg, 1908 ; = PotentlllP although remained infertile and reproduced hticosa L. ; = Dasiphora fruticosa (L.) only vegetatively, had very luxurious Rydberg]. growth and flowered extensively. In view of such evidence, we treat 2. Subg. TRICHOTHALAMUS(Lehm.) Reich., Duchesnea Smith as a subgenus of Potm- Consp. Rep. Veget. : 167. 1828. Tricho- till22 L. thalamus [Spreng. Anleit. ed. 2, 2 : 864. 18181 Lehm. in Nova Acta Acad. Leap.- Carol. 10 : 585. 1821. Type species : SIBBALDIAL. vis-a-vis POTENTILLAL. Trichothalamus lignosus (Wllld. ex Linnaeus (1753, 1754). Wolf (~gd),Ryd- Schlecht.) Lehm. ( = Potentilla lignosa berg (1g08), Juzepczuk (I 94 I) and Hutchin- Willd. ex Schlecht.). son (1964) treat Sibbaldia L., as generically Potentilla L. subg. Lasiocarpa DiHit et distinct from Potentilla L. and differentiate Panigrahi in Journ. Sc. Club 33-34 : 39. them on the number of stamens : 4, 5 or 10 1983, Type species : P. salesoviana starnens4ibbaldia whereas 2e30 st amens- Stephan. Poten~iNaL. Except the number of stamens, no other consistent difference can be discern- 3 Subg. J?RPCARUSTRUM (Heist. ex Fabr.) Reich, cit. : 167. I 828 ; Ball er al., Fl. ed between the two genera. However, op. Sibbaldia L., is only herbaceous, sexually- Europ. 2 : 45. 1968. reproducing and shows no enlargement of Fragariastrum Heist. ex Fabr., Enum. torus after anthesis, In porentilla, some Method. P1. 64. 1759. Lectotype species : species are fruticose, are obligate apomicts F. sterilis (L.) Schur. Fragaria sterilis L. and exhibit torus enlargement after anthesis. (selected by Panigrahi and Dikshit, Taxon The Indian species of Potentilb sensu 35 : 350. 1986). ~80 BULLETIN OF THE BOTANICAL SURVEY OF INDIA [Vol. 27

Sect. Trldenhtae Dikshit et Panigrahi in Subg. CLOSTEROSTYLIS(Torrey & Gray) J. Orissa Bot. SOC.3(1) : 13. 1981. Juzepczuk, op. cit. : 93. 1941 (Engl. ed. : Potentilla L. sect. Trichocarpae Wolf sub- 71- 197'). sect. Nematostylae Wolf grex Tridentatae Potentilla L. sect. Closter~stylisTorrey & Wolf, Bibl. Bot. Heft 71, Band 16 : 46, 77. Gray, op. cit. : 445. 1840; Janchen, Cat. 1908, nom. invalid. (Art. 4, Art. 5. I, F1. Austr. Tiel. I Pteridophytes and ICBN, 1983). LT : Potentilla jridentata Sol. Anthrophytes I 78. 1957. (selected here). Lectotype species : Potentilla arguta Sect. Ehaupe Juzepczuk, in Komarov, Pursch (selected here). N. URSS 10 : 88. 1941 (Eng. ed. : 1971) ; Subg. CHENOPOTEN~LLA(Focke) Juzepczuk, Grossan. FI. Kawkass 5 : 68. 1952. op. cat. : 164. 1941 (Engl. ed. 220. 197 I). Potentilla L. Sect. Trichocarpae Wolf Potentilla L, sect. Chenopotentillu Focke, subsect. Nemadostylue Wolf ser. Suffruti- op. cit. : 415. 1899. culosae Wolf grex Eriocarpae Wolf, op. Type species : Potentilla anserina L. cit. : 46, 79. 1908, norn. invalid. (Art. 4. I ; Sect. Chenopotentilla Focke Type Art. 5. I, ICBN, 1983). Species : Potentilla anserina L. Lectotype species : Porentilla eriocarpa Argentina J. Hill, op. cat. : 6. I 756 ; Wall. ex Lehm. (selected here). Larnarck, op. cit. : I 18. 1778, p.p. ; Hut- Sect. CQRrisetae Schiman-Czeika in K. H. chinson, op. cit. : 198. 1964. Rechinger, FZ. Iranica 66 : 87. 1969. Lectotype species : Argentina anserina Potentilla L. sect. Trichocarpae Wolf sub- (L.) Rydberg ( = Potentilla anserina L.). sect. Nematostybe Wolf ser. Herbaceae Potentilla L. sect. Gymnocarpae Wolf Wolf grex Curvisetae Wolf, op. cit. : 47. subsect. Leptostylae Wolf grex Anserinue 1908, norn. invalid. (Art. 4.1 ; Art. 5.1, (Lehm.) Wolf, op. cit. : 52, 667. 1go8, p.p. ICBN, 1983). Potentilla L. sect. Herbaceae Lehm. ser. Lectotype species : Potentilla curviseta Anserinae Lehm. op. cit. : 8, 188. 1856. J. D. Hooker (selected here). Potentilla L. sect. Anserinae (Lehm.) Subg. SCHISTOPHYUIDIUM Juzepczuk, ofi Grossh. op. cit. : 82. 1952. cit. : 81. (1941) (Engl. ed. : 61. 197 I) Ball Sect. Lencondae Dikshit & Panigrahi, et al., op. cit. : 39. 1968. J. Orissa Bot. Soc. 3(1) : 33. 1981. Lectotype species : PotentiUa bifurca L. Type species : Potentdla leuconofa D. (selected here). Don. SUbg . MICROPOCON(Bunge) Juzepczuk, op. Potentilla L. sect. Gymnocarpae Wolf n't. : S4. 1941 (Eng. ed. : 64. 1971). subsect. Nematostylae Wolf grex Anserinae Micropogon Bunge, Suppl. F1. Alt. : 40. Wolf, op. tit. : 52, 667. 1908, pap., excl. P. anserinn I,. nom. invalid Art 4. 1, 5. I, ICBN I 836. Type species : Potent& biflora Willd. ex 1983. Schlecht. Sect. Duw~neSojak in Preslia 41 : 349. Potentilh L. sect. Trichocarpae Wolf sub- 19%. sect. Nematostylae Wolf grex BifEorae Type species : Potentilla dumosa (Franch.) (Rydberg) Wolf, op. cit. : 46, 70. 1908, Hand,-Mazz . p. p., comb. invalid. (An. 4.1 ; Art. 5.1, Potentilla L. sect. Gymnocarpne Wolf ICBN, 1983). subsect. Conostylae Wolf ser. Orthotrichue 19~51 PANIcRAHI AND DIKS~T : SYS~ATTCSOF THE GENUS POTENT~LLA t. 181 Wolf grex Tanacetifoliue (Lehm.) Wolf, op. Subg. HYPARGYRIUMJuzepczuk, op. cit. : it. : 670, 743. 1908, p-p. 101. rgdr (Engl. ed. : 76. 1971). Lectotype species : Potentilla argentea L. Subg. POTE~LLASyme in Smith & Sower- (selected here). by, English ~otai~,ed. 3, 3 : 143. 1864. Lectotype species : Potentilla reptans L. Potentilla L. sect. F ragariastrum Ser . ex DC. Prodr. 2 : 583. 1825 incl. P. (selected by Rydberg, I 908). P.P. leschenaz~ltiunaSer. Potentilla L. sect. Gymnocarpae Wolf subsect. Gomphostylae Wolf op. cit. : Hypargy rium Fourr. Ann. Soc. Linn. 671, 786. 1904, et op. cit. : 52, 641. 1908. Lyon 2, 16 : 371. 1888, nom. invalid. (Art. Potentilla L. subg. Dynamidium Juz., op. 36.1, ICBN, 1983). tit. : 194. 1941 (Engl. ed. : 144. 1971). Sect. Multifidae Juzepczuk, op. ch. : Sect. Pdeutilla Lectotype sp. : Potentilla 1 13. 1941 (Engl. ed. : 85. 197I) ; Schiman- reptans L. Potentilla L. sect. Tormentilla Czeika, op. cit. : 68. 1969), [as '(Rydberg)']. Focke, op. cit. : 415. 1889 ; Schiman- Lectotype species : Potentzlla multzfidu L. Czeika, op. cit. : I I I. 1969. (selected here). Sect. Dacbeamw (J. E. Smith) Dikshit et Potentzlla L. group Multzfidae Ryd berg Panigrahi, comh, nov. in Bull. Torr. Bot. Club 23 : 262. 1896, nom. invalid. (Art. 4.1, Art. Duchesnea J. E. Smith in Trans. Linn. 5.1). Soc. London 10 : 372. 18I r ; Valentine in Sect. Haematachroae Schiman-Czeika, op. Tutin et al., F1. Europ. 2 : 36. 1968. tit. : 91. 1969. Type species : Duchesnea f ragiformis Lectotype species : Potentilla atrisanguinea J. E. Smith, nom. illegit. [=Potentilla Lodd, (selected here). indica (Andrews) Wolf 1. Polentillo L. sect. Gymnocarpae Wolf Fragariu sensu J. D. Hooker, op. cit. : subsect. Conostylae Wolf ser. Eriotrichae 343. 1878, p.p., incl. F. indica Andrews, non Wolf grex Haematochroae Wolf, op. cit. : 49, 2 19. 1908, nom. invalid. (Art. 4.1 ; L. ('753)- Art. 5.1, ICBN, 1983). Potentilla L. sect. Gymnocarpae Wolf subsect. Gomphostylae Wolf grex Tormen- Sect. Hypaqyrium tillae (Lehm.) Wolf, op. cit. : 52, 641. 1908, Potentilln L. sect. Herbaccae Lehm. ser. p.p. incl. Potentilla indica (Andrews) Wolf. Digitatae Lehm. tribe Multiflorae Lehm. : Sect. Aurae Juzepczuk, op. it. : 197. subtribe Argenteae Lehm., op. cit. 6, 93. 1856 nom. invalid. (Art. 4.1 ; Art. 5.1, 1941 (Eng. ed. : 147) [as '(Lehm.) Juz.']. Lectostype species : Potentilla aurea L. ICBN, 1983). (selected here). Potenn'Na : sect. Gymnocarpae Wolf Potentilla L. sect. Herbace~e Lehm, ser. subsect. Conostylae Wolf ser Eriotrichae Digitatae Lehm. tribe Aureae Lehm., op. Wolf grex Argenteae Wolf, op. cit. : 49, 252. 1908, nom. invalid. (Art. 4 ; Art. 5.1). tit. : 6 : 112. 1856, p.p., nom. invalid. (Art. 4. I ; Art. 5.1, ICBN, 1983). Potentilla L. subg. Hypargyrium Juz. Potentilla L. sect. Gymnocarpae Wolf sect. Argenteae Juz., op. cit. : 144. 1941 subsect. Gomphostylae Wolf grex A~-eae (Engl. ed. : 106. 197I) ; Schiman-Czeika, op. cit. : 92. 1969. Wolf, op. it. : 787. 19q, op. tit. : 523. 1908. nom. invalid. (Art. 4.1 ; Art. 5.1, Sect. Tanacetifolhe Juzepczuk, op. ICBN, 1983). it. : 152. 1941 (Engl. ed. I 13. 197 I) ; 182 BULLETIN OP THE BOTANICAL SURVEY OF INDIA pol. 27

Schiman-Czeika op. cit. : 94. 1969 [as Sect. Redae Juzepczuk, op. cit. : I&- '(Lehm.)' Juz.]. 1941 (Eng. ed. : I tg. 197 I) ; Schiman- Lectotype species : Potentillo tenacetifolke Czeika, op. cit. : 94. 1969 [as '(Lehm.) Juz.']. Lindl. ex Lehm. (selected here). Lectotype species : Potentilla recta L. Potentilla L. sect. Herbaceae Lehm. ser. (selected here). Pinnatae Lehm. tribe Tenacetifoliae Lehm., Potentilla L. sect. Herbaceae Lehm. ser. op. it. : 55. 1856, p.p., nom. invalid. Digitatae Lehm. tribe Multifiorae Lehm. (Art. 4.1). subtribe Rectae Lehm., op. cit. : 5, 8 I. 1856, Potentilla L. sect. Gymnocarpae Wolf nom. invalid. (Art. 4.1 ; Art. 5.1, ICBN, subsect. Conostylae Wolf ser. ~rthotrichae 1983). Wolf grex Tenacetzfoliae Wolf, op. cit. : Potentilla L. sect. Gymnocarpae Wolf 670, 743. 1908, nom. invalid. (Art. 4.1 ; Art. subsect. Conostylae Wolf ser. Orthotrichae 5.1, ICBN, 1983). Wolf grex Rectae Wolf, op. cit. : 50, 302. 1908, nom. invalid. (Art. 4.1 ; Art. 5.1). SPlRAEOiDE AE ROSOlDEAE X39 X 7,849 Sect. Persicae Juzepczuk, op. cit. : 175. E aC lost 1941 (Eng. ed. : 130. I); Grossh., op. cit. : 76. 1952 [as '(Wolf) Juz.']. Lectotype species : Potentilla persica Regel & Schmalh. (selected here). AMYGDALOTEY MYIDEAE Potentilla L. sect. Gymnocarpae Wolf x a8 x'17 C lost subsect. Conostylae Wolf ser. Orthotrichae Wolf grex Persicae Wolf, op. cit. : 67 I, 786, 1904, et op. cit. : 51. 1908, nom. invalid. (Art. 4.1 ; Art. 5.1). Sect. Rhales Juzepczuk, op. cit. : 165. ROSACEAE 1941 (Engl. ed. : 123. 197 1) ; Schiman- Cieika, op. cit. : 99. 1969 [as '(Wolf) Juz.']. Er Ellogltannlns Ct Flavone-c-glycosidcs Lectotype species : Potentilla rivalis L. (selected here). Hypothetical Scheme-1 . Potentilla L. sect. Gymnocarpae Wolf ;Sect. Cbryamthae Juzepczuk, ~p.cit. : ser. Orthotrichae Wolf grex Rivales Wolf, 180. I 941 (Eng. ed. : I 34. I 971) ; Schirnan- op. cit. : 670, 744. 1904, et op. cit. : 50, 384. Czeika, op. cit. : 108. 1969. 1908 nom. invalid. (Art. 4.1 ; Art. 5.1, Lectotype species : PotentiZla chrysantha ICBN, 1983). Trevir ex Ledeb. (selected here). Potentilla L. sect. Herbaceae Lehm. tribe Potentilla L. sect. Herbaceae Lehm. ser. MultifEorae Lehm. subtribe Rectae Lehm. Digitatae Lehm. tribe Multiflorae Lehm. op. tit. : 5, 81. 1856, p.p., nom, invalid. subtribe Chrysanthae Lehm., op. cit. : 5, (Art. 4.1 : Art. 5.1). 75. 1856 norn. invalid. (Art. 4.1 ; Art. 5.1, ECONOMlC UTILISATION ICBK, 1983). Economically, Potentilla are not very im- PotentilZa L. sect, Gymnocarpae Wolf portant to human beings, although accord- subsect. Conostylae Wolf ser. Orthotrichae ing to the old herbals of the 16th-18th Wolf grex Chrysanthae Wolf, op. cit. : 51, century by Fuchs, Bauhin, Book and 455. 1908, nom. invalid. (Art. 4. r ; Art. 5. x). Tabaernamentanus, the healing powers of I~~SJ PANIGRAHI AND DMSHIT : SYSTEMA~SOF THE GENUS POTJWTIUA L.

KEY TO THE NINE SUBGENERA OF POTENTILLA L. la. Receptacle after anthesis becoming densely villous ; achenes clothed with and con- cealed by long woolly hairs 2a. Herbs ; bark not peeling off; caudex aerial, with a rosette of radical leaves ; leaves chiefly radical, ternate ;styles lateral or subterminal . . Fragariastnmr (6 taxa) 2b. Shrubs ; bark peeling off; caudex subterranean ; without a rosette of radical leaves ; leaves chiefly cauline, imparipinnate (except Potentilla rigida with ternate leaves) ; styles subbasal or lateral 3a. Leaflets entire ; androdynamic or gynodynamic ;petals yellow (white in same Chinese taxa) ;styles claviform, a8 long as ovary .. Dasipkra (6 taxa) 3b. Leaflets finely serrulated or dented ; flowers bisexual ; petals white ; styles filiform, two (three-) four times longer than ovary . . ~ruhot~mus(3 taxa) lb. Receptacles after anthesis becoming glabrous-shortly tomentose ; achenes 4a. Flowers a-drodynamic or gynodynami~; styles claviform . . S~hist@~Eliditmr(1 tam) 4b. Flowers bisexual ; styles atherwise, nct claviform 5a. Leaflets entire ; styles filiform (2-3 mm long) .. Micrapogon (2 taxa) 5b. Leaflets to~thed/serrulated/pinnatelypartite ; styles otherwise, not filiform 6a. Styles spindle- or rod-shaped 7a. Leaves interruptedly pinnate with one or more pairs of reduced leaflets, styles spindle-form . . Closteros~lis (3 taxa) 7b. Leaves regularly pinnate without reduced leaflets (except in P. anminu and cften in P. leuconota) ; styles rod-shaped . . ClraMpo&ntilla (13 taxa) 6b. Styles claviculiform or coniform 8a. Styles claviculiform . . Potentdla (7 taxa) 8b. Styles coniform .. HypargWum (39 taxa]

Potentilla for many human ailments were geographical areas and showing a tendency eulogised. However, P. reptans sometimes for further spread, often occupying larger is applied as astringent in domestic medi- areas than their sexual relatives. Cosmo- cines and for tanning of hide and to impart politanism with significant intraspecific ply- red colour for the woollen fabrics. Thick morphism, without much evidence for roots of P. anserina serve as the food ecological localisation (oiz. ecotypes, ecads stuff to poor inhabitants of Northern etc.) are characteristic of larger genera of region. In an earlier communication apomictic nature. Apomicts show an (Dixit & ~ani~rahi,1980) the cconomic absence of stability in chromosome num- utilisation of 12 species of Potentilla occurr- bers : polyploid and aneuploid series are ing in India has been exhaustively dealt almost the rule. Yet, a relative constancy with. in the morphology of the apomictic progeny and even of mutants in the succeeding Apomiwis and its impact on the delimita- generations, is ensured by the absence of tion of taxonomic categories in Potenfilla meiosis and fqrtilisation, aided by the uni- Darlington (1939) considers a~ornixisas an versal presence of agarnospermy and vegeta- escape from sterility, because it ~ermitsper- tive apomixis, 'While this is true for obligate petuation of well-adapted but sexually apomicts, as in Alchemilla L., in which sterile phenotypes, resulting in a large ca. rooo 'microspecies' are recognised population of gknetica~l~similar individuals (Panigrahi & Purohit, I 983), taxonomic for colonization. Apornixis has been re- delineation in facultatively-apomictic genera, Ported in about 300 genera, including such as Potentilla L. poses problems of a Potentilla L., belonging to about 80 families different, more challenging, nature. Here, of angiosperms (Khokhlov, 1~70). The in the progeny of facultatively apomictic highest rate of occurrence of a~omixisis in parents, biotypes resulting from normal the phylogenetically young and progressive sexual processes, with an introgression of taxonomic groups occupying widespread characters of both the parents occur simul- 184 BULLETIN OF THE BOTANICAL SUKVEY OF INDIA Lv01. 27 taneously, with parthenogenetically produced within P. recta L., a collective species, four 'uniparental' progeny, thus blurring up the segregate species are recognised in Europe originally phenotypically distinguishable (Ball et at., 1968). Of the Indian apo- parents. Recognition of 'microspecies' a micts, P. reptans is represented by an addi- taxonomic category amongst obligate apo- tional variety var. microphylla, P. argyro- micts or subspecies or varieties which are Phylla by an additional var. leucochroa, P. taxonomic entities recognised in the sexually- multifida by two additional varieties var. breeding taxa, is rather difficult, among glabrata and var. minor and P. supina by facult atively apomictic taxa. However, an additional subsp. paradoxa. Since P. apomixis is not found in relics which are neglecta, P. recta and P. nirea are represent- at the point of extinction or represent evolu- ed by less than 5 specimens in Indian her- tionary blind-ends. baria, it is not possible to comment on the Judged in this context, amongst the nine degree of variations amongst their Indian subgenera recognised in Potentdla, four sub- populations. P. atrisanguinea, and P. nepa- genera, uiz. Dasiphora (6 taxa), Schisto- lensis, each with a number of cytotypes, phyllidium (I sp.), Trichothalamus (3 taxa) represented by a large number of speci- and Fragariastrum (6 taxa) and even mens in CAL, DD, BSD, do not contribute Micropogon (2 sp.) and Closterostylis (3 distinctive phenotypes sufficient to recognise taxa), which represent the basal stocks or any additional variety in any of them. P. evdutionarily blind-ends or occupy mid- multifida, on the other hand, with 2n= 28, points in the evolutionary grades, sharing 42, presents distinctive populations not suffi- one or more primitive traits, are strictly ciently geographically isolated differing from sexually-breeding and no evidence of apo- each other in the degree of pubes- mixis in any species of the six subgenera is cence of the upper surfaces of leaflets and in yet available. the cutting of the, leflets. By and large, then, Even amongst the remaining three sub- the Indian apomicts do not pose a challenge genera, comprising 59 (80-2I) taxa apomixis to orthodox taxonomists, bit conform with is recorded only in 10 species : subg. the theoritical expectations that, despite Potentilla (P. reptans L. with 2n=28), subg. genotypic variations, the facultative apomicts Chenopotentilla (P. anserina L. with 2n=28, do not throw up distinctive phenotypes suffi- 35, 42) and subg. Hypargyrium with the cient to accord them taxonomic status. remaining eight apomictic taxa, viz. P. Of the sexually-breeding species, P. arbus- neglecta Baung. (2n= 42), P. argyrophy lla cula and P. microphylla, etc. produce dis- Lehm. (2n= 14, 56, 63), P. atrisanguinea tinctive morphological types, in different Lodd. (zn=56, 63, 84), P. nepalensis W. J. ecological habitats, to enabl; one to recognise Hooker (2n=42) and P. nivea L. (2n=~6, four-five distinct varieties, not subspecies, 70), P. multifida L. (zn =28, 421, P. recta L. because of the geographical overlapping (2n=28, 42) and P. supina L. (2n = 28). These between the distinct phenotypic populations. taxa exhibit cuploid- and aneuploid series Thus, the original apprehension that faculta- based on x= 7 and are usually considered as tive apomixis in Potentilla would blur taxo- facultatively apomictic with pseudogarnous nomic judgements Droves false at least reproduction. P. neglecta usually an obli- amongst the Indian taxa. gate apomict with 2n=42, produces in Europe progeny with 2n =35 which are mor- Distribution : The distributional range of ~holdgically indistinguishable from the Potentillo L. lies mainly between the Tropic hexaploid parents. On the other hand, of Cancer and the everlasting ice of the Polar 19~~1 PANICRAHI AND DIICSHIT : SYSTEMAnCS OF THE GENUS POTENTLLLA L. I@

region in the Northern Hemisphere, although ties with 24 additional varieties) of potentiZJa the cold temperate high mountain peaks in L. and seven species to Potentilla L. the Southern Hemisphere harbour some spe- sect. Sibbaldia (L.) J. D. Hooker. A critical cies. The great Himalayan ranges in the study of the tribe Potentilleae Jussieu north, serve as the gene sanctuary for the in India has led us to delimit the occurrence wealth of Potentillas in India. Airy Shaw of 80 taxa of Potentilla sensu strict0 (Map) (197~) species to potentilla L., 20 and 14 taxa (12 species and 2 varieties) of species to Sibbaldh L, and one species to Sibbaldia L. (Dixit 6Panigrahi, 198 1b) with Duchesnea Smith, in the world flora. J. D. Indian distribution. Of the 80 taxa of Hooker (1878) reported onlyij6 taxa (32 spe- Potentilla, 54 taxa are indigenous[endemic PTtilla Taxa Indigenous~~1danicto the Himalayas and to South India

Serial Names of the 8 Number

Subg. DASIPHORA * 1. Pot&&, ri@ 2. P. albnscrJ3 var. 3.. ,, var. glabrata 4. 9, var.PIDdkr 5. ,r Var. Subg. TRICHO~S-

6. P-mkophyU. var. 7. ,. var. &ngijilia Subg. FRAGARIASTRUM Sect. TRIDENTATAE

9. P. lxfn@lia Sect. BRTWAE 9. P. niawrpo var. niwarprr 10. rr VW. ~~

Sect. CURVISETAE

Subg. MICROPOOON Table 1 ( Contd. ) I \S)m

A 8 3 m zSr Serial 8 Number Names of the taxa

Subg. CLOSTEROSTYLIS

15. P. lineata var. lincata 16.; ,, var. in&nncdia

Subg, CHENOPOTENTILLA Sect. LEUCONOTAE 17. P. lcuconola 18. P. p~dwt~ularisvar. pcdundaris 19.* ,, var. clarkei 20.* ,, var. obscura 21. P. microphylh var. microphylla 22. ,, var. achillewia 23. ,, var. glab~iuscula 24. ,, var. latifolia 25. ,, var. latilobn 26. P. lalsimluenris

Sect. DUSlOS;\E

Subg. POTEPiTlLL.4 Sect. POTESTILL.-\ 29. P. hemkyona Sect. AUREAE 30.; P. khasiana Subg. HYPARGYRIUM Swial Names of the taxa Numbet

Sm. MULTIPIDAE

31.' P. mdb~rdovar. gldnotcr 32.* P. omithopauk var. iw 33. P. konaOR(i 34.* .P. karahnnka 35.* P. 6rtuiscirsa 36.. P. luchemdw

Sect. HAEMATOCHROAE 37. P. -. 38. P. ~UUWI 39. P. argyrojhyh var. 40. var. 41. P. so&i

Sect. NIVEAE 42. P. saundcrsimm 3 P. &nioccr var. ccrarpifosa 44. P. $nrpstii 45. P. pau&ia?nl Sect. TANACETIFOLIAE 46. P. spodioklma 47. P. #arardioM . 4%. P. bllm&hm 49. P. &i 50.. P. cLukei. . 51. P. tmtabdu

San. WALES 53. P. nmJmmhsvar. z4elmih H. ,, v-ar.swV@i& PANICRAM AND DIKSHIT : SYSTEMA'I'ICS OF THE GENUS POTE~YTIU L. i 89 to the temperate-alpine regions of the Hima- ORIGIN OF THE GENUS POTEflfILLA L. layas between 1830-6100 m, extending from The epoch when the genus originated : the Baluchistan arc to the Burmese arc, The exact epoch for the origin of Poteatilla enduring severe cold and frost (Table I). L. and its close allies, cannot be stated with They are distributed from the Northwestern certainty, although the occurrence of unmis- to the Eastern Himalayas, through the takable remnants of ~otektilbfnrticosa in mountain ranges of the Himachal Pradesh, Pliocene deposits, seen together with the Wttar Pradesh, Nepal, ~arjeeling-dkkim doubtful findings of Potentilla L. seeds both region, Bhutan, Arunachal Pradesh, Mani- in the Miocene and Pliocene deposits (Wolf, pur and Burma and extending to Megha- 1908)~is a pointer to the probable origin of laya-Nagaland sector, the latter belonging to Potentilla and Potentilla-like ancestors earlier geological age than the Himalayas. earlier to the Miocene period. Some of the higher hills of the Western Place of origin and di@rentiation : It is Chhota Nagpur plateau in Bihar enjoying well recognised that the recent arctic and a mild subtropical climate congenial to the boreal flora of the Northeast- Asia has growth of Potentillas, harbour four taxa viz. survived to a great extent up to the present- Potentilla bannehalensis, P. indica, P. sun- day because of their escape from complete daica and P. supina subsp. supina. It is glaciation during the Pleistocene. That the interesting to note that the latter three ex- modern Potentillas and their allies occur tend their range further South to the Nilgiri preponderantly in the arctic boreal region, Hills, through the tropical belt of India and probably led Wolf (1908) to assume, with grow luxuriantly, together with POten tiNa certainty, that the Tertiary Circumpolar land leschenaultiana, endemic to the Nilgiri Hills. might have been the native land of earlier Of the 53 Potentinu taxa endemic to the Potentillas and that its major evolution Himalays, only 15 taxa are endemic to must have taken place in the temperate and India proper (asterisked in table I). Of the cold regions of the Northern Hemisphere. remaining 26 taxa having a wider geogra- One may justifiably start with the presump- phical distribution, nine extend as far west tion that the original stock in the arctic as the Western Asia, three species (viz. P. Tertiary flora, might have already split up sundoica, P. polyphylla and P. indica), to into many species-groups, of which some of Japan and Sri Lanka, and, 13 taxa reach as the remnants still survive in that region. far as the cold temperate belt of Europe and Wolf (1908) regarded the groups, viz. the Sibiria. Of these 13 taxa, only five species Fruticosae (= Potentilla L. subg. Dasiphora), reach as far as the North ~merica,beyond the Paltrstres (= Potentilla subg. Comarum), their Asiatic range (Table 2). According to the Rupestres ( = Potentilla L. Closterostylis), Wolf (~gos), no species of ~otentilla are the Niveae (Potentilla shbg. Hypargyrium common to Europe and Arctic America sect. Niueae) and the Anserinae ( = Potentilla (including Greenland). subg. ChenopotentiNa sect. Chenopotentilla) Only Potentilla anserina L. of the North- distributed in all the three northern conti- ern Hemisphere occurs in the saline habitats nents (Asia, Europe and North America), as of Chile, South America and New Zealand very old within the genus. He (Z.C.)believed as P. anserina L. subsp. ~mserinoides(Rous~, that these groups distributed originally in 1965) Not a single species of ~otentillais the Tertiary Circumpolar region migrated reported from Australia and from Africa southwards, differentiated and split up into beyond the Mediterranean region of Nonh the present-day northern continents. And, Africa. that, with this separation, a new phase of Table 2 9 Europe U.S.S.R. &A Names ofthe taxa (includ- (indud- E. A& W. Asit Pakia- Nepal Bhutan Burma S. E. China Sri N. Arne- S. +I- Number ing N. +g Sibi- tan Mi hka rica am Afirl~a) rb gion) Subg. DASIPHORA 1. PoUntiUa @mtBIia f + + + Sub. TRICHOTHALAMUS 2. P.&k + Subg. SCHISTOPHYLLIDIUM 3. P. bifiuca + + 4. P. a;$ha + Subg. CLOSTEROSTYLIS 5. P. poluph-~lla Subg. CHENOPOTENTILLA sect. CHENOPOTENTILLA 6. P. ansrnmnasub. SD. anmina + + Subg. POTENTILLA Sect. DUCHESNEAE P. indica Sect. POTENTILLA P. rcptans var. rcptans var. &&Uo P. &:w Sect. AUREAE

var. minor P. bnlcto~ P. pdyskista P. hddcuca P. ~wngaico P. saricco w. NIVEAE P. Riosa Sect. HYPARGYRIUM P. wglacta Sea. RECTAE P. ncta Sect. PERSICAE p- Sect.- WALES SI . Jammu Hima- Uttar Pun- No. N3mc.s of the tau & Kash chal Pra- jab Raja%- Guja- Tamil Maya Biiar West Sikkim Aqsam Megha- Aruna- Naga Mani- mir Pra- dah than rat Nadu Pca- Bcn~al la- chal land DW

Subg. DASIPHORA 1. PotcntillnmYdie + Subg. TRICHOTHALAMUS 0 2. P. snlcsouiana + + Subg. SMISTOPHYLLIDIUM E 3. P. ~CO + ++ * Subg. MICROPOGON 4. P. ba@ra ++f 8 Subg. CLOSTEROSTYLIS 5. P. pol*ylk, + ++ Subg. CHENOPOTENTILLA kt.CHENOPOTENTILLA 6. P. anserina sub sp. ansnitur + i- Subg. POTENTILL.4 m Sect. DUCHESNEAE ce 7. P. in&a + ++++ ++ +++4- 4- ++ Sect. POTENTILLA 8. P, rqjtans var. nptans + ++ 1=I 9. var. ++I + + lo. P.~I&W HIMALAYAN REGION 2 Sect. AUREAE 0 11. P. gslido + ++ *1 Subg. HYPARGYRIUM Sect. MULTIFIDAE 12. P. df&var. nylh'f& + ++ P 13. var. o~m + + 0 14. P. dlicto~ + 15. P. pd'&& + 16. P. S + 'u 17. P. mngonca i- 0 18. P. saricca + + Sect. NIVEAE 19. P. niza + + I Sea. HYPARGYRIUM 20. P. ~gban + E Sect. FUETAE 21. P. wci% + + r Sect. PERSICAE 22. P.j-l&Wa Sea. RIVALaS 23. P. drsatarrrn 192 BULLETIN OF THE BOTANICAL SURVEY OF INDIA [Vol. 27 evolution for Potentillas began ; many spe- salesoviana in the Central Asia and P. cies perished, leaving a few survivors in the bijlora in the circumpolar region. It .is Circumpolar region. And, consequent to interesting to record that major part i,e. the disappearance of original Circumpolar 75% of Potentilla common to North source and spatial separation of the three America-Europe-Sibiria-Indian sector are continents, further diversification and sepra- apomictic species. While the secually free- tion occurred, resulting in specific Asiatic, breeding species of Potentilla have in general specific European and specific American restricted distribution, the apomictic taxa groups of species. Despite such specificity, comprise a number of 'microspecies' and about 16 taxa of the European continent ex- cover wider geographical area and present a tend their range to the temperate Asiatic network of morphologically intergrading mountains through the higher mountain forms. ranges of the Mediterranean-West Asia and Afghanistan (Wolf, 1908). However, ~iriAncestral relationship of the genus Potentilla Sojak comments (Personal communication) Of the four subfamilies recognised that Wolf's theory of the arcto-tertiary within the family Rosaceae Jussieu. the evolutionary region as the centre of Poten- Spiraeoideae with basic chromosohe till#, is rather improbable. number x=8, g and the Amygdaloi- deae with x=8 comprising trees and Centre of diflerentiation shrubs represent two ancient and distinct The principal centre of origin of Po~entilla evolutionary lines. The subfamily Maloi 'L: Wolf in India may be ,conjectured dene with x=,17, on the other hand, ' is from the fact that as many as 54 taxa considered as an allopolyploid arising from from among a total of 80 taxa (66%) are hybridization between the Spireoideae with endemic to the termperate-alpine zone of x=g and the Amygdaloideae with x=8, the Himalayas. Critical analysis of the followed by chromosame doubling. Accord- distributional data of these taxa suggests ing to Gajewski (1959) and Challice (1974) that within this broad centre of origin, two the present-day Rosoideae evolved from the endemic subcentres may be recognized : Spiraeoideae, because : (i) the 'basic chromo- (a) the North-West Himalayan subcentre some numbers x=7, 8, g of Rosoideae pro- with 66 taxa, of which only rjare restricted bably developed through one of its tribes, to this region and @) the East Himalayan the Kerrieae (x=g) from the tribe subcentre with 47 taxa, of which only ro S@reae of the subfamily Spiraeoideae. taxa are restricted to the Sikkim-Burmese (ii) Ellagitanins (E) and flavone c-glycosides arc of Himalayas. (c) present in the Spiraeoideae ancestors It is significant to note that of the 80 are also present in the Rosoideae genera Indian taxa of Potentilla L., only 10 are with x=7 and x=8 ; (iii) the Spiraeoi- known to be apomicts : three species uiz. P. deae and the Rosoideae are linked by nepalensis, P. atrisanguinea and P. argyro- 6-0 substitution of flavones ; Scutellerin phylla, endemic to the Himalayas ; four 7-rhamnoside and scutellerin 7-methyl ether species presumed to have originated in the in Sorbaria of the Spiraeoideae vis-a- Central Europe and remaining three species, vis Scutellerin 6, 4 dimethyl ether in Kmia in Central-Asia-Arctic region. Of the three of the Rosoideae,

non-apomictic species common to ' India, Challice (I.c.) speculates that the 6-hydro- Europe and Asia, P. bifurca is postulated to xylated flavonols came from the original have originated in the Arctic region ; P. rosaceous stock and that the dhydroxylited 19851 PANIGRAHI AND DIKSHIT : SYSTEMAmCS OF THE GENUS PoTGNTIUA L. I93 flavonols did not appear until later and sides of the Pacific ocean. The tribe, prior to the separation of primitive Potentilleae, comprising 16 genera, shares Spiraeoideoc into present-day Spi~aeoideae the basic number of chromosome (x=./) and Rosoideae. On the other hand, Bate- with a few other allied tribes, such as the Smith (1961) considers, on the basis of plant Geeae, the Rose6 the Sanguisorbeae and chemistry, that the Rosoideae represent the the Ulmarieae amongst dthers, and may be oldest surviving form of the primitive considered as one of the youngest evolu- Rosaceae. The view is countered by the tional branches of the subfamily Rosoideae argument that, although the Rosoideae may (Gajewski, 1957) and in which active evolu- be primitive in respect of phenolic consti- tionary differentiation is discernible (Hypo- tuents, in comparative morphology, it is thetical scheme I). more advanced. Within the subfamily Rosoideae, the Evolutionary trends within the nine sub- ancient ancestral group comprising trees and genera of Potentilla L. shrubs belong to the tribe Kerrieae, In discussing the evolutionary tendencies Lkyadeae and Cercocarpae, each with detected in the different subfamilies or x=9. The majority of .the genera in tribes or genera of the family Rosaceae, these tribes are monotypic with small relic botanists generally consider the following geographic ranges lying primarily on both traits as primitive or advanced :

--r . Primitive Traits Advanced Traits -- , . * 1. Plants perennial Plants annual or biennial 2. shrubs or undershrubs without a rosette of radical Herbs with a rosette of radical leaves leaves 3. Leaflets with entire gin Leaflets with toothed/dented/pinnatifid margin. 4. Flawers dioecious Flowers bisexual 5. Styles persistent, hairy and gmiculate Styles deciduous, glabrous and entire 6. Achenes hairy Achenes glabrous 7. Receptacles saucer-shaped and becoming woolly Receptacles more or less enlarged after anthesis, dry or after anthesis fleshy and becoming shortly pubescent or glabrous after anthesis 8. Anemochoric dissemination of 8eeds Epizo~~horicdissemination of seeds

Amongst the g subgenera of Potentilla, and further southwardly to central Asia and subgenus Dasiphora stands out as the Mongolia, the discovery of both diploid most primitive, as much as all of its forms in North America-Europe and tetra- species represent shrubs with woody bran- ploid forms in the Far East, may be a pointer bhes, ensuring perennation, lacking rosette to its ancestral and still evolving nature. In leaves, entire leaflets, almost dioecious its southward migration to the temperate- flowers, woolly receptacles, daviform st@s alpine Himalayas, Potentilla fruticosa the and long-hairy achenes conducive to anemo- most primitive species of the genus Potentilla, choric dissemination. Such mor- is substituted by the polymorphic species phol.ogical trends, seen against the palaento- P. arbuscula with four varieties and P. logical evidence of its occurrence in the rigida, both endemic to the =malayas ; pleistocene of the tertiary (Kirchheimer, whereas P. pa*vifoth, mainly distributed in 1957) ; its wide geographical distribution the high mountains of Central Asian region ranging from North America to Europe of U.S.S.R., extends to the Himalayas. 194 BULLETIN OF THE BOTANICAL SURVEY OF INDIA pol. a7 In possessing serrated leaflets and bisexual S~histoph~llidiumappears to have a parallel flowers, the species belonging to the sub- development with the subgenus Trichb genus Trichothalamus exhibit evolutionarily thalamus from Dasiphora-like ancestors. advanced traits in relation to the species of Similarly, in the subgenus Micro- the subgenus Dasiphora. pogon, the paleotypic species P. biflora The subgenus Fragariastrum with its re- originating in the circumpolar land, has markable distribution in the high moun- gradually advanced in all directions and is tains of Central Europe and of the Alps presently distributed over a wide range, shows that probably the ancestor of the from North America, U.S.S.R., Mongolia, whole group originated here and as the Iran and to the whole of the Himalayas. little known or small groups (sections), it However, in the Eastern Himalayas P. extended to and/or developed in due course armerioides might have developed indepen- in the Himalayas. These Himalayan taxa dently. Subgenus Micro@gou (with P. are not only morphologically, but also gene- biflora and P. armerioides in the Indian tically, nearly related to the other species region) approaches the subgenus Fraga- within the groups or other groups (sections) riastrum in its perennial herbaceous of the subgenus Fragmiustrztm. The sub- habit, radical leaves (ternate or irnpari- genus Fragaiastrum shares its serrated pinnately pfoliolate), bisexual flowers and leaflets, bisexual flowers, filiform styles and filiform styles, and represents a further stage hairy achenes with the subgenus Tricho- in evolutionary advance in possessing gla- thalamus. But in producing a rosette of brous achenes. Yet, the presence of entire radical leaves and herbaceous aerial branches leaflets, a primitive character met with in it scores over the subgenus Trichothalamus Dasiphora, shows that evolutionary deve- as more advanced. lopment of different organs of the same The subgenus Schistoph y llidium repre- plant body is not synchronous. sented by the single polymorphic species P. Subgenus Cheno@tentilla is represented bifurca, probably originated in the upper by four sections. Of this, the section North, and radiated eastwardly to Central Chenopotentilla with the centre of origin Europe and to U.S.S.R. as distinct varieties/ in the mountains of central Asia, com- formas. But only as var. bifisrca it has ex- prises a single polymorphic and apomic- tended southwardly through Western Asia tic species, P. unserina, with a number to the Himalayas. The subgenus Schisto- of geographical races : as subspecies anse- 'phylEidium shares with the subgenera rim, it attained most extensive range of Dasiphora and Trichothalamus its peren- distribution and the broadest morphological nial habit and imparipinnate leaves, variation and occupies most of the temperate but it shows more affinity with the former zone in the Northern Hemisphere and in possessing entire leaflets, dioecious flowers, Southern Hemisphere due to its obligate out hairy ovaries, when young, claviform styles, breeding, combined with a vigorous vegeta- yet, it differs from them in developing a tive reproduction, aided by frequent migra- rosette of radical leaves with toothed tions through human agency. However, leaflets and glabrous achenes. The affinity the other sections tiz. Leuconotae and of 'the subgenus SchistophyRidium with Dumosae are neot~picin origin and are the subgenus Fragariastrum appears re- mainly distributed in the Himalay as. mdte despite the rosette of radical leaves In the subgenus Chmopotentilla, the sec- 'Common to both. In weighing the charac- tion Chenopotentilla shows its affinity with ter& t-on to the other subgenera, the subgenus Micropogon i~ its perennial herbaceous ha bit, rosette of radical leaves, But in coniform styles, it is possible to have bisexual flowers, filiform styles and glabrous been derived either from the filiform styles achenes, but represents a further stage of of Micropogon or the rod-shaped style evolution in having interruptedly irnparipin- of Chenopotentilla by the swelling of nate leaves, dentate or serrulated leaflets, and the lower basal ,portion of the style, skck prostrate and creeping flowering stems. Sec- it is difficult to visualise the development of tion Leuconoiae, in having perennial herbs a coniform style from a claviculifo~or with a rosette of imparlpinnate leaves, even spindle-shaped style, reduced (in P. leuconotrr) and serrated - lets, bisexual flowers, rod-shaped styles and glabrous achenes, approaches the section Chenopotentilla. But, in possessing a rosette of leaves without reduced leaflets (as in P. $endunculnric, P. microphylla and P. tatsienluensis) and erect or ascendent flowering stems, it is Jess advanced than the section Chenopotentilla; and, in having pinnatisect leaflets (as in P. micro- phylla), it is more advanced than the section Chenopotentilla. Whether Leuconotae with rod-shaped styles is a direct offshoot from Chenopotentilta or is derived from Micro- mn may only be conjectured at present. Subgenus Closterostylks in having peren- nial herbs with a rosette of interrupt- Hypothetical Scheme-!? edly imparipinnate radical leaves, serrated leaflets, bisexual flowers and glabrous The evolutionary tendencies within Poten- achenes, is related to the subgenus Cheno- tilla L. in India, as visualised above, may be potentilla. But it shows an advanced stage represented by a schematic diagram (Hp of development in possessing spindle-shaped thetical scheme 2). styles over the subgenus ChenopoteniiUa. ACKNOWLEDGEMENT Subgenus Potentilla with perennial We are grateful to the Director, Botanical herbs, rosette of radical leaves ternate or Survey of India, Howrah for facilities and digit at el^ 5-foliolate, bisexual flowers, sub- encouragements and for awarding a Seoior terminal styles, glabrous and smooth achenes, Research Fellowship to one of us (B.K.D.). is nearer to the subgenus Micropogon, but, in its creeping flowering stems (as in REFERENCES section Potent&) and serrated leaflets ACHARYAG~~~AMI, D. Biosystematic studies in the genua (in both the section Potentilla and Aureae), Potdntilla L. section G'carpdc. Ph. D. Thesis (abstract), University of London, London. 1972. it has greater affinity with the subgenus BALL, P. W., B. PAWL^ AND S. M. WALTEBB.Genus Chmopotentilla. Potmtilla L. Flora Europaea. Edited by T. G. Tutin In having leaves ternate, digitate or even et al., 2 L 36-47. Cambridge University Press. 1968. BATE-Sm,E. C. Chromatography and in inparipinnate with serrated leaflets, subg. the Rosaceae, with special reference to Potentilia and H@rgyrium appears to be a synthetic Prunus. Journ. Linn. Soc. Bot. 58 : 39-54. 1961. pup combining with the characters of CHACLI~,J. S. Rosaceae chemotaxonomy and the origins of the Pomoideae, Bet. Jem. Lim. Sec. 69: 219- subgenera Micropogon and Potentilla. 259, fig. 2. 1974. 196 BULL~ETWOF BOTAMCAL SURVEY OF I~IA [tiof. 27

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