ACHENE MORPHOLOGY; AN AID TO OF INDIAN . I. COMPOSITAE. LIGULIFERAE

G urcharan Singh, Bimai, Misri and P. Kachroo

Department o f Botany, Kashmir UnivcrsUy, Srinagar

A bstr a c t

The morphology of achcncs of hguliferous composilae of India has been discussed with special reference to their use in identification of taxa. A key to the genera based on achene characters is presented. Diagnostic characters of the species found in India are also given. A representative sample of each genus is illustrated.

I ntroduction drupaceous Chrysanlhemoicles of Africa, fruit is an achene, and shows sufTicient Plasticity is an attribute of nature which consistency within a species and diversity enables living beings to survive over dis­ of characters in dilTerent taxa; yet reliance crepancies of environment. The mainaimof’ merely on achene structure for such a large taxonomy has been to sort out least plastic group would obscure the very value of characters so as to form the basis of classi- characters. This is, however, a valuable lication. The somatic bodies arc more plas­ tool in regional floras. In fact, characters tic as compared to gametic bodies and as of achene supplcniented by other charac­ such latter have been greatly exploited for ters have been extensively used in several taxonomic purposes. The characters of taxonomic works dealing with Compositae seed and fruit, in this connection, have re­ (Hooker, 1872-97; Rydberg, 1954; Ferris, ceived more attention in most of the fami­ 1960; Gupta, 1968, etc). lies of flowering plants. In the proposed Indian Compositae are represented by ca series, of which the present treatise is the 697 species (Datta, 1965). Though treated first communication, the detailed morpho­ variably, there is general agreement in logical characters of achene are being ex- recognizing two subdivisions in the family: r>loitedfor taxonomic studies independent Tubiflorae withca 12 tribes and Liguliflorae if other characters. This in addition, may with a single tribe . The latter serve to identify the seeds of the concerned characterised by presence of only ligulate nlants. florets and milky latex is so distinct from Compositae, easily the largest family of other Compositae that some authors

lowering plants, embraces nearly 2 0 , 0 0 0 prefer to remove the ligulate genera into a Decies distributed in about 950 genera separate family, Cichoriaceae (Rydberg, Lawrence, 1951). In all of them, excepting 1954), It is represented in India by ca 22 genera with nearly 80 species, barring Accepted for publication on December 7 ,1972. numerous species of Taraxacum recognised byV onSoest (1963). stra ig h t...... 9 Twenty two liguliferous Compositae Pappus uniseriate achenes slightly genera are studied for their aehene charac­ curved upwards and narrowed into ters. To determine the range of flexibility inconspicuous (less than a mm) if any of achene structure large number beak ...... Pier is. of fresh as well as herbarium specimens 9. Achenes short, 5-6 mm long usually were examined. The representative species b ro a d e r...... Epilasia

are described in detail and illustrated, for Achenes longer than 8 mm, usually other Indian species only distinguishing narrow...... Scorzonera characters are given. The characters which 10. Achenes beaked conspicuously... 11 were most static were selected for diagnosis Achenes beakless or inconspicuously

of taxa. In a very few cases where actual so (beak less than a m m )...... 2 0 specimens were not available for exami­ 11. Achenes with toothed ring below the nation help was taken from literature. b e a k ...... ChonJrilla Achenes without a toothed ring below

KEY TO SEEDS OF THE INDIAN the beak...... 1 2 LIGULI FERAE 12. Achenes echinate near apex, strongly 4-,‘5 angled, (at least lower ones are so) 1. Pappus a b s e n t...... 2 ...... Taraxacum Pappus p resen t...... 3 Achenes not echinate ab o v e...... 13 2. Achene straight or slightly curvcdbut J3. Achenes less than 4 nim long (incl. not sickle shaped, without prickles... beak) ...... 14 ...... Lapsana Achenes more than 4 m m long (incl. Achene strongly curved and sickle beak) ...... 15 shaped, with rows of strong prickles 14. Achenes usually sm ooth...... Ixer/s ...... Koflpinia Achenes usually with muricate ribs 3. Pappus pJaeaceous ...... 4 ...... Laduca (pp) Pappus hairy...... 5 15. Achenes strongly compressed...... 16 4. Achene short straight...... Cichorium Achenes terete or slightly compressed Achene curved almost sickle shaped . and 4 angled ...... (ppj ...... Garhadiolus 16. Achenes with beak less than 0.75 the

5. Hairs plumose ...... 6 length of body...... 17 Hairs not plumose, sim ple...... 10 Achenes with beak more than 0.75 the

6 . Beak (at least on inner achene) conspi­ length of b o d y ...... 18 cuous...... 7 17. Achenes usually more than 5 mm long Beak absent or inconspicuous (less ...... Cicerbita (pp)

than a mm long)...... 8 Achenes usually less than 5 mm long 7. Outer achenes beak less, inner with a ...... Lactuca (pp) very slender beak, pappus hairs atta­ 18. Body of achenes cleft at top into which

ched individually...... Hypochaeris base of beak fits i n ...... 1 ^ All achenes beaked, beak gradually Body of achcne entire at top, conti­ narrowed from body and stout, pappus nuous into beak ...... Lactuca (pp) hairs forming a deciduous ring...... 19. Beak more than 1.5 times the length ...... Tragopogon of b o d y ...... Lactuca (ppS

8 . Pappus multiseriate, achenes almost Beak almost equalling or shorter than b o d y ...... Cicerhiia (pp) collar. Pappus absent. 20. Achcacs strongly compressed, faces Koelplnla.—Koclpinia linearis Pall. usually strongly 3 ribbed... (Fig.2).—Achenes curved sickle shaped, Achenes terete or angled or slightly 1.2-1.8 cm long, cylindrical, faintly ribbed,

com pressed...... 2 1 dorsal surface with prickles arranged in 21. Pappus uniseriatc, rigid, brittle, persis- rows. Prickles 1.5-3 mm long, hooked at tent, usually tawny, achenes dark apex, those at apex of achene spreading coloured...... Hieracium stellately, 6-9 in number. Achene glochidi- Pappus multiscriate, hairs usually soft, ate all over. Pappus absent, deciduous silvery...... 22 Cichorium.— Cichorium infyhus Linn. 22. Pappus hairs forming a deciduous ring (Fig. 3).— Achene short, 2-3 mm longca 1.5 ...... 23 mm broad, mottled, 5-anglcd, angles dark- Pappus hairs individually deciduous greyish. Achenes broadest nearapex which ...... 24 is truncate, gradually narrowed towards 23. Achenes 4 angled, constricted above, base. Pappus paleaceous, pales 2-3 seriate, ribs 4 strong...... Picridium few, 0.4-0.6 mm long. Achenes subterete or slightly com- Garhadlolu*.— Garhadiolus minutis- pressed, ribs 4-5, usually not constric- sinm.—(Bunge) kit. (Fig. 4).—Achenes ted above...... Launea 5-9 mm long, curved, cylindrical. Outer 24. Achenes slightly comprcssed, narrowed achenes glabrous gradually narrowed, at both ends, pappiferous disc distinct abovefrom the base, enclosed in involucre ...... 25. bracts. Inner with more prominent pappus Achenes not compressed, terete or than in outer achenes. Pappus palaeaceous, angled, pappiferous disc not distinct.. pales fringed, usually less than a mm long...... 26 All achenes faintly ribbed. 25. Achenes narrowly linear, less than 5 Hypochaaris.— Nypoc/iacris glahra mm long...... Youngia Linn.—Achenes of two types: outer 4-5 Achenes narrowly oblong usually more mm long, nearly ) mm broad, linear oblong, than 4 mm long...... Prenanihes truncate, beakless, pappus plumose short- 26. Achenes very slender, curved, broader ly exceeding the achene; inner 7-10 mm towards base...... Phaecasiutv long, beaked, beak slender longer than the Achenes slender, straight, narrowed body, pappus as long as beak, towards both ends, usually narrowly Tragopogon.— Tragopogon kashmiriam linear...... (pp). Singh (Fig. 5).—Achenes with beak. 2-3 cm long including the beak, body narrowly Briei- Descriptions oi Species lanceolate oblong, gradually narrowed above to a shorter or as long beak, ribs Laptana.—Lapsana communis Linn. muricate. Inner achenes smoothly ribbed; ('^•ig, 1).—Achenes pale brown, glabrous pappus 2-3 cm long, plumose, hairs rigid ' 'lip.soid to obvoid straight or slightly connate at base into a deciduous ring, a ' Lirved, compressed, 3-5 mm long, 1-1.5 few hairs much longer than others and with "m broad, much narrowed towards base, simple tips, tawny, hairs ashy. ' 'Oadest just above middle. Ribs 10-13 on T. gracile D. Don.—Achenes hardly 1.2 t ich face, faint but conspicuous near apex cm long (incl. beak), smooth and with very ' liich is narrowed to form a rounded short beak. T. porrifolius Linn, has more F ig s. 1-10. Achcnes of ligulifcrous Compositae of India,. Fig, 1. Lapsana communis. Fig. 2. Koe/pinia linearis. Fig. .1. Cic/ioriiim intybus. Fig. 4. Garhailiolus minuti.ssima. Fig. 5. Tragopogon kashmiriana. Fig. 6. Picri.'i hieracoiiks. Fig. 1. Scor:onfra divaricata. Fig. 8. ChondriUa graminea. Fig. 9. Taraxacum officinale. Fig, 10. polycephala. strongly muricate achenes with beak much nate above. Beak capillary, nearly 8-14 longer than body. mm long, broadened above into a pappi- Picris.— Picris hieracoides Linn. (Fig. ferous disc. Pappus 5-9 mm long, soft

6 ).—Achenes almost cylindrical, narrowed white, pappus hairs simple. at both ends, more so and curved near I. wattii Hook f. has white achenes apex into an inconspicuous beak, achenes with the beak shorter than body. 2.5-4 mm long, dark brown, ribbed, ribs Ixtris.—/xeris polyccphala Cass. (Fig. transversely rugose, pappus deciduous, 10).—Achencs with beak 2.5-3.5 mm,

white soft and plumose, 8 - 1 2 mm long. long, body lanceolate, glabrous, narrowed E p ila tia ..-EpHasia ammophila Bge.— into as long or shorter beak, brownish. Achcnes short almost cylindrical, 5-6 Ribs 4-5 on each face, strong, paler than mm long, sometimes turbinate and as rest of the body which is reddish brown.

broad, closely nearly 2 0 ribbed. glabrou.s, Pappus .silvery equalling the achenes. Hairs tip raised and dome shaped within the softs, simple. pappus, pappus dark brown, multiseriate, I. sagittarioUk’s (Clarke) Stebbins has hairs plumose, a few with simple tips. smaller (ca 2.5 mm long) achenes with Scorzonera.— Scorzonera Jivaricata white beak; while /. gracilis (DC.) Stebbins

Turcz. (Fig. 7 ).- Achenes linear, l.5-2cm has pale red smaller achenes (ca 2 mm long) long, cylindrical or angled, many ribbed, and dirty white pappus. pale brown, equally broader throughout Lactuca.—Lactuca serriola Linn. (Fig. or slightly narrowed above, pappus less 11).—Achenes oblanceolate, pale brown, than I cm long, mutiseriate, hairs unequal, strongly compressed, 3-5 mm long plumose, tips usually simple. excluding beak, faces many ribbed, ribs In S. purpurea Linn, achenes arc shorter scabrid. Beak very slender, white, directly than pappus. continuous from body and nearly equalling In S. Stew art a Hook. f. achenes are it. Pappus soft, silvery, 4-7 mm long, winged with echinate ribs. pappiferous disc prominent. Chondrilla. - ChonJrilla f’raininca M. Other species with similar achenes are:

Bieb. (Fig. 8 ).—Achenes oblong-lanceolate, L. longifolia DC. (Fig. 12) with dark brown 3-5 mm long, narrowed towards both achenes, 3-5 ribbed, base of beak white: ends, more so towards base, prominently L. brcvirostris Champ, with black achenes

10-16 ribbed, pale coloured with 1 - 2 rings having a strong midrib and other slender of teeth below the beak. Beak very slender, ribs on each face, base of beak black; L. nearly equalling the body, broader clarkei Hook. f. with black achenes having towards two ends, pappus 5-8 mm long, a coloured beak with pale tip. white, persistent, soft, simple. L. dissecta Don (Fig. 13).—Achenes Achenes are more strongly toothed in oblanceolate, 5-8 mm long (incl. beak), C. setulosa C. B. Clarke. strongly compressed, faces 3 ribbed, T araxacum .— Taraxacum officinale margins slightly thickened, scabrid, tip of Wigg. (Fig. 9).— Von Soest (1963) has body cleft to accomodate base of the beak. isolated many species from this complex Beak capillary, white, twice the length of Indian . body, broader towards two ends, thickened Achenes narrowly ovoid, 4-5 angled, above into a pappiferous dise. Pappus greyish or greenish brown. Body of achene soft, silvery, 3-6 mm long, simple. 3-5 mm long, strongly ribbed, ribs echi­ L. undulata Ledeb.—With similar ache- nes has larger sized achencs, (13-18 mm C. nndticaulis Ledeb. subsp. congesta long), and beak 3-4 times as long as body. (Rgl.) Babe. — achenes narrowly fusci- Shorter beaked species fall into two form, 4-6 mm long, red brown. categories: (a) with smooth achenes in L. C. glomcrata Dene., C. himalaica Kit., orientalis Boiss., pappus equalling the ache- C. tlioinsoni Babe., C. nanifornia Babe, ne; L. larlanca Mey., pappus twice the have essentially similar achenes. length of achenes (b) with muricate ribs C. fheiida Linn, has achenes unlike the of achenes in L. heyneana DC., achenes rest species; they have a beak (at least the 2-3 mm long, pappus twice as long; L. inner one) almost equalling the body. ramoiifolia DC., achenes 3-5 mm long, Sonchus.— Sonclius aspcr'ViU. (Fig. 16).-— pappus as long. Achenes ellipsoid, 2-3 mm long, strongly Cicerblta.— Cicerhiia decipiens (Hk. f & compressed, narrowed above but not T. ex Clarke) Beauv. (Fig. 14).- Achencs beaked, edges serrulate, brownish, faces 7-10 mm long (incl. beak), elliptic to strongly 3-ribbcd, obscurely rugose on ribs lanceolate, strongly compressed, dark and intervals. Pappus soft, while, multi- brown, dorsal face with several faint ribs, seriale. Hairs few, 8-14 mm long, simple. ventral with a strong midrib and faint S. otcraceous Linn, achencs2 mm long, lateral ribs, slightly undulating, puberulus. strongly transversely rugose. Beak 2.5-3.5 mm long, upper part slightly 5. arvcnsis l.inn.— achenes 2.5-3.5 mm whitish, apex enlarged into a pappiferous long, transversely rugose. disc. Pappus 6-9 mm long, soft, white, S. muritimus Linn, has midrib much hairs simple individually deciduous. thicker than the laterals. Another long-beaked species, C. cyanea Hieracium.— Hieracium Yidgatum Koch. (Don) Beauv. has apex of body cleft as in (Fig. 17).— Achenes 2.2-3 mm long, cylind­ some species of Laciuca accommodating rical, black, shinning, ribbed, narrowed the base of the beak, which nearly cqualls towards base, truncate above. Pappus uni- Ihe body. seriate of rigid tawny coloured hairs latter Rest of the species have a very short puberulus nearly equalling the achenes. beak, usually less than 0.75 the length of //. virosum Pall, has similar red brown body. C, rapwicuhides (DC.) Beauv. has achenes while species listed below, have slender, 7-9 mm long achenes with slender larger achenes. ribs. C. macrorhiza (Royle) Beauv. has H. crocatum Fries (Fig. 18)— achencs 4-6 mm long achenes, black in colour. 4-5 mm long, black, faintly ribbed. Crepis.— Crepis sancla(L.) Babe, subsp. //. prenanthoides Vill.— achenes 4-5 mm bifida (Vis.) Thell ex. Babe. (Fig. 15).—^ long, red brown, strongly ribbed. Achenes narrowly linear, 3-5 mm long, less //. tiinhcllaliim Linn.— achenes 3-3.5 mm than a mm broad, slightly narrowed at long, reddish. two ends, pale brown, many ribbed, pap- PIcridium.— Picridium lingilatiumDesL- piferous disc not distinct. Pappus soft, Achenes oblong, 4 angled 2-3 mm long, white, simple, almost equalling or smaller pale, truncate at both ends., slightly cons­ than achenes. tricted towards apex. Ribs 4, rugose; pap­ C. sihirica Linn.— achencs 10-15 mm pus multiseriate, soft, white, connate below long, ribs slender. into a deciduous ring, pappus 10-15 mm C. kashmirica ^a.bc.— achenes 7-10 mm long. long, ribs strong. Launea.— Launea failax (Jaub. et spach F ig s. 11-21. Achenes of liguliferous Conipositac of India. Fig. 11. Lactuca scrriola. Fig. 12. L. longifotia. Fig. I.V L. dissccta. Fig, 14. Ciccrhita decipicns. Fig. 15. Crepis sancta. Fig. 16. Sonchus asper. Fig. 17. Hicracium viilnatiim. Fig. 18. H. crocalum. Fig. 19. Launea fallax. Fig. 20. Yoimgia lemiifolia. Fig. 21. Prenamhes hrunoniana. Ktze. (Fig. 19).— Outer achenes often brown, glabrous, pappus white; achenes compressed, inner cylindrical narrowly, 2-5 larger. mm long, truncate at both ends. Ribs 4-5 y. silhetensis (DC.) Babe, and stebbins stout. Upper end of achenes suddenly has achenes 2-3 mm long, pale brown, broadened into a pappiferous disc. Pappus pappus brown; achenes smaller. multiseriate, soft, white 10-15 mm Y. japonica (Linn.) DC. has achenes

long, hairs simple, connate below into a nearly 2 mni long, glabrous, red brown deciduous ring. pappus white; achenes smaller, Other species having same sized achenes y. graciUpcs (Hook, f) Babe, and are L. asplenifolia DC., pappus 4-6 mm stebbins. long; L. sarmentosa (Willd.) Alston, y. racemifera (Hook f.) Babe, and pappus 6 - 8 mm long; L. secunda Clarke, y. nilgiriensis Babe, are other Indian spe­ achenes ca 3 mm pappus as long. L. micro- cies of the genus. Their achenes were not cepliala Hk. f.. achenes ca 3 mm long, available for study. pappus 2 mm or less. — Prcnanthes hniiiotiiana L. chondriUoides DC. has larger (5-6 mm Wall. (Fig. 21).— Achenes narrowly oblong long, pappus 10-15 mm long) achenes. 4-5 angled, 4-6 mm long, sligthtly com­ Achenes of L. glonierala Cass, are different pressed, dark brown, minutely pubescent, in being winged, ca 4 mm long with 5-6 mm ribs strong, achenes more prominently nar­ long pappus. rowed towards base, obscurely so above, Youngia.— Youngia tenuijoHa (Willd.) pappus 2-3 seriate, hairs soft, while, 7-9 mm Babe, and Stebbins (Fig. 20).— Achenes long, individually deciduous, puberulous. small dark brown, 2.5-4 mm long, narrowed Achenes in the rest of the species are at both ends, more so above, enlarged similar sized, being very narrow in P. above into a pappiferous disc, ribs distinct. scandens Hook, f., less so in P. violaefolia Pappus soft, nearly 1.5-2 times the length Dene., P. khasiana Clarke., broader more ofachene, hairs simple white, individually darker in P. hookeri Clarke. deciduous. P. sikkimensis Hook. f. has larger nearly

Y. glauca Edgew.—achenes 4-5 mm long, 6 mm long achenes which are more narrow­ pale, obscurely scabrid. pappus white, ed above. y. gracilis (Hk. f. ex Benth.) Hook. f. P haecaiium .— Phaecasium lampsanoides . achenes 3-4 mm long, brown, minu­ Cass.—Achenes 4-6 mm long, terete, broa­ tely scabrid, pappus grey. dest towards base, very slender, slightly cur­ Y. depressa (Hk. f. and T.) Babe, and ved, pappus soft, white, multiseriate, hairs stebbins.— achenes ca 5 mm long, dark simple, deciduous, shorter than achenes.

R e f e r e n c e s

D a t t a , s. C. 1956. A Handbook of Systematic L a w r f .nc'F, G. H. M. 1951. Ta.xonomy o f Vas­ Botany. Asia Publishing House, Bombay. cular plants. The Macmillan C o ., New York. F erris, R. S. I960. Illustrated Flora o f Pacific R y o b e r u , p. a . 1954. Flora o f Rocky Moun­ States. Vol. IV. Stanford University Press, tains and Adjacent Plains. Hafner Publ. Co., California. New York. G u p t a , R. K. 1968. Flora Nainilalensis. Nava- SoEST, Von 1963. Taraxacum species of India, yug Traders, New Delhi. Pakistan and neighbouring countries. tVenlici H o o k e r , J. D. 1872-97. Flora o f British India. JO: 1-91. Vols. I-VIl. L. Reeves Co., London.