FRUIT DIVERSITY OF FEW MEMBERS OF THE TRIBE HELIANTHEAE (ASTERACEAE)-MORPHOLOGICAL OVERVIEW

BIODIVERSITY

Interrelation between Flora, Fauna and Human

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Mrinalini Datta Mahavidyapith Birati, Kolkata-51

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Biodiversity: Interrelationship between Flora, Fauna and Human ©2014, Mrinalini Datta Mahavidyapith

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Proceedings on Biodiversity: Interrelationship between Flora, Fauna and Human Pp. 11-15 (ISBN: 978-81-929410-0-4)

FRUIT DIVERSITY OF FEW MEMBERS OF THE TRIBE HELIANTHEAE (ASTERACEAE)-MORPHOLOGICAL OVERVIEW

Tulika Talukdar* and Sobhan Kumar Mukherjee** 'Dept. of Botany, Krishnagar Govt. College, Krishnanagar-74110L, Nadia, West Bengal.

"Dept. of Botany, University of Kalyani, Kalyani, Nadia-741235, West Bengal E-mail: talukdartulip 12@gmail. com

Abstract

The sunf lower family, Asteraceae being the largest family of f lowering plants possesses end numbers of specialities including its unique type of fruit known as 'cypsela' . Cypse la is a dry one seeded fruit deve loped f rom inferior bicarpel late ovary. Perusal of l iterature reveals that very little attention has been devoted to cypsela for optimising it as a potent taxonomic tool. Within the scope of the present communicat ion, a critical and detail observat ion on fruits of few members of Hel iantheae like Acmel la Rich, ex Pers., Echinacea Moench., Gal insoga Ruiz & Pav. and Tridax L. has been undertaken which showed a wide range of variation in macro and micro-morphological fruit characterist ics. Present investigation clearly indicates that cypsela is a very rich source of morphological diversity and could be uti l ized for bet terment of Asteraceae taxonomy.

Key words: Cypsela, morphology, pappus, carpopodium, trichome, taxonomy.

Introduction

The, tribe Hel iantheae is morphological ly most d iverse t r ibe of the As te raceae and proved to be a useful source of various valid controversies in many taxonomic treatments. The tribe was establ ished by Cassini (1819) based in part on m i c ro - synan the ro l og i ca l characters. Bentham (1873) fol lowed Cassini's s y s t e m w i t h a d e p a r t u r e f r o m p r e v i o u s classification. Bentham recognized Helenieae ("Helenioideae") for epaleate taxa with scaly p a p p u s tha t w e r e p l a c e d in C a s s i n i ' s Hel iantheae and also moved epaleate taxa with bristly pappus of Cassini 's Hel iantheae to Senecioneae. According to Cronquist (1955), this tribe is the most primit ive among the all

tribes and suggested as the ancestral stock of the ent i re A s t e r a c e a e . S tuessy (1977) reviewed Hel iantheae from systematic point of v iew and recognized 15 sub-tr ibes.

Based on cpDNA f indings of Panero et al. (2001), Panero & Funk (2002) proposed a new tribal classif ication that redelimit Hel iantheae in a more l imited sense (Hel iantheae S.s.) consist ing of majority paleate taxa and a wide diversity of woody l ineages in the Hel iantheae a l i i a n c e . H e l i a n t h e a e (S . s . ) is l a r g e l y monophylet ic consist ing of 113 genera, ca. 1461 species (Panero & Funk 2002). The tribe is m o s t l y d i s t r i b u t e d in t he n e w w o r l d , especia l ly in Mex ico, Centra l Amer ica and South America.

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To unders tand phy logeny of any d iverse group like Hel iantheae it is necessary to study all the organs of a plant be longing to all the species wi thout except ion. Detail invest igat ion on fruits or cypsela in di f ferent members of Hel iantheae will be thus immense ly helpful for bet terment of Hel iantheae taxonomy. In this context , present commun i ca t i on dea ls wi th d e t a i l a n a l y s i s of m a c r o a n d m i c r o -morphological features of cypseia in four taxa namely Acme l la Rich, ex Pers. , Ech inacea Moench. , Ga l insoga Ruiz & Pav. and Tridax L. using Light Microscope. Special emphas is has b e e n g i v e n o n s u r f a c e c h a r a c t e r s , carpopodium, s ty lopodium and pappus, with a purpose to create a reference set of micro-morphological features wh ich could be useful for identif ication of the studied taxa in absence of f lower ing stage.

Materials and Methods

Plant mater ia ls (cypselas) for the present invest igat ion were col lected by the author and obta ined in the fo rm of rece ived herbar ium spec imens f rom the fo l lowing herbar ia of the world which are ment ioned in Index Herbar ium (Holmgren et al., 1981).

DK : H o r t u s b o t a n i c u s H a u n i e n s i s , Denmark .

Z : Botanischer Gar ten der Universitat Zur ich,

Zol l ikerstrasse 107, CH8008 Zurich, Switzer land.

Macro-morphologica l studies of cypsela

In c a s e s , w h e r e in tac t c y p s e l a s w e r e avai lable, the first and foremost step was to mark the pos te r i o r a n d an te r i o r (abax ia l ) sur face of the cypsela. Then 10 dry and 10 FAA preserved mature cypsela were randomly taken in glass sl ides and graphed sl ides and observed under O l y m p u s stereo d issect ing

m i c r o s c o p e ( D M ) a n d O l y m p u s b i nocu la r m i c roscope (No. -611062) . Su i tab le images were taken using Ze iss Stemi DV4 camera equ ipped microscope.

Colour, shape , d i rect ion of cypse la w a s noted carefully. Length and width of the cypsela were measured visUally by g raphed sl ides, in few cases they were counted by ocular and stage micrometer. The length of the cypsela in the present study is def ined as the length of the body of cypsela f rom basal mer is temat ic zone (carpopodium) up to apical end excluding pappus. The width was measured at the widest par t of t he c y p s e l a . O u t l i n e d i a g r a m s of comp le te cypse la and d i f ferent parts we re drawn by the Mirror type camera lucida.

Micro-morphologica l studies of cypsela

M a t u r e c y p s e l a s w e r e d i p p e d ' i n 1 - 5 % N a O H solut ion for 2 -7 days depend ing upon the hardness. Then they we re t ransferred into s a t u r a t e d ch lo ra l hyd ra te so lu t i on for few hours , r epea ted l y w a s h e d w i th wa te r and properly stained in 0.2-0.5 % aqueous Safranin solution. After staining, spec imens were placed in 70 % phenol glycerine solution and dissected c a r e f u l l y f o r s t u d y i n g d i f f e r e n t p a r t s of cypselas. Sui table pho tog raphs were taken using Olympus C-310 zoom digital camera (3.2 Megapixel ) and Zeiss-stereo microscope.

N a t u r e of r ibs, t y p e s , d i s t r i b u t i o n a n d or ientat ion of hairs, nature of sur face cells, other epidermal st ructures, carpopodia l cells etc . all w e r e c r i t i ca l l y o b s e r v e d . P a p p u s characters such as nature of pappus brist les, their number, a r rangement , apex organizat ion etc. were also examined.

Results and Discussion

C o m p a r a t i v e m o r p h o l o g i c a l fea tu res of cypsela in the s tud ied taxa are presented in T a b l e 1. D e t a i l e d m a c r o a n d m i c r o -

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morphological studies of cypsela in the present invest igat ion (F ig.1-4) reveal cons iderab le va r ia t i ons and h a v e t a x o n o m i c po ten t ia l . A m o n g t h e d i f f e r e n t m o r p h o l o g i c a l parameters , shape of cypse la was not so va r i ab l e ; it w a s e i t he r o b l o n g or na r row obovate. Cypsela colour was also supposed to have not much impor tance in ass is t ing re lat ionship as it changes wi th matur i ty of cypsela. Size of cypsela ranged from 1.0-1.5 x 0.5-0.7 mm in Gal insoga to 4.0-5.0 x 2.0-2.5 mm in Echinacea.

After cleaning, the cypsela surface showed a l v e o l a t e m a r k i n g s in G a l i n s o g a a n d scorb icu la te mark ings in Tr idax (Fig. 3,4). A m o n g the s tud ied taxa, r ibbed g lab rous surface was noted in cypsela of Echinacea. In all other studied taxa cypsela were pubescent without any rib. Among the surface features, t r i chome or hair t ype p r o v e d to be mos t valuable feature. In the present investigation, biseriate or twin type of hair was most prevalent along with simple branched fil iform hair (Fig.4) as in Tridax. Short conical hairs were noted in Cosmos by Mukher jee & Sarkar (1998). Such t r i chomes var ia t ion may be under genet ic control as Mehta et al. (1979) indicated that in guayu le p lants ( P a r t h e n i u m a rgen ta tum) , presence of 'mariola' genes has resulted in an i n c r e a s e in leaf t r i c h o m e s l eng th and a decrease in the rubber content.

Very little variat ion is noted in sty lopodium and carpopodium in all the studied taxa and probably for the tr ibe Hel iantheae they have lesser taxonomic values.

Diversity of pappus as depicted by various different forms like awned, scaly, coronate or bristly pappus noted in present studied taxa could be treated as valuable taxonomic marker and has considerable systemat ic implication (Fig.1-4). Different forms of pappus structure

in Heliantheae have also have been mentioned by Bremer (1994). Robinson (1981) has given too much importance of pappus features in higher level of taxonomic decisions. Among the different forms,""Galinsoga posses primitive scaly pappus (Fig.3) and it can be said that Tridax by having bristly pappus (Fig.4) is more advanced than any other present investigated taxa, as bristle or bristle-like pappus characters have evolved repeatedly f rom scales or awns (Baldwin 2009). However, stiff awns of Acmella in our investigation (Fig. 1) probably are not a true pappus but are separate outgrowths of the cypsela as ment ioned by Robinson et al. (2009).

A l l t h e s e m a c r o a w e l l as m i c r o -morphological features of cypsela could be s u c c e s s f u l l y u t i l i z e d f o r b e t t e r m e n t of Hel iantheae taxonomy.

Acknowledgements

Author is immensely greatful to University G ran t C o m m i s s i o n ( U G C ) for e x t e n d i n g f inancial suppor t for imp lementa t ion of the project [Sanction No.F.PSW-068/11-12(ERO), Da ted 3rd Augus t , 2011] , Au thor ex tends special thanks to Curator, Denmark and to Curator, Zur ich for their active assistance in despatching the identif ied mature cypselas for our studies.

References

Baldwin, B. G. 2009. Hel iantheae all iance. In : Funk, V.A., Susanna, A., Stuessy, T.F. & Bayer, R.J. (eds.). Systematics, Evolution, and Biogeography of Composi tae, pp. 689-711. Smi thson ian Inst i tut ion, Wash ing ton , D.C., U.S.A.

B e n t h a m , G. 1 8 7 3 . N o t e s o n t he c l a s s i f i c a t i o n , h is to ry , a n d g e o g r a p h i c a l d is t r ibut ion of Compos i t ae . Journa l of the Linnean Society, Botany 13 : 335 - 577.

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Bremer, K. 1994. As te raceae . Cladist ics and Classif icat ion. T imber Press, Port land.

Cassini , H. 1816-1830. In: King, R.M. & Dawson, H. W. (eds.): Cassini on Composi tae, col lected from the Dict ionnaire des Sciences Naturel les. New York : Or iole Edit ions, 1975.

C r o n q u i s t , A . 1 9 5 5 . P h y l o g e n y a n d taxonomy of the Compos i tae . Amer. Midi. Nat. 53 : 478 - 511.

Holmgren, P. K., Keuken, W. & Schof ield, E. K. 1981. Index Herbar iorum. Part I. The Herbar ia of the Wor ld . Ed i t ion 7 (Haque : Regnum Vegetabi le, 106).

Mehta, I. J., Dhil lon, S . P & Hanson, G. P. 1979. Tr ichome morpho logy as an indicator of high rubber bear ing guayu le (Par thenium argentatum Gray) plants in native populat ions. Amer. J. Bot. 66(7) : 796-804.

Mukher jee , S. K. & Sarkar, A. K. 1998b (2001). Comparat ive morpho-anatomical study of c y p s e l a s in s o m e s p e c i e s of the t r ibe Heliantheae (Asteraceae). Bull. Bot. Surv. India 40 (1-4) : 34-46.

Panero, J. L., Baldwin, B. G., Schi l l ing, E. E. & C l e v i n g e r , J . A . 2 0 0 1 . M o l e c u l a r phy logenet ic s tud ies of m e m b e r s of t r ibes H e l e n i e a e , H e l i a n t h e a e a n d E u p a t o r i e a e (Asteraceae).2. Tribal/generic relationships. In: O s b o r n , J. M . ( p r o g . d iv . ) , B o t a n y 2 0 0 1 , Abs t rac t s , par t 3, S y s t e m a t i c s . Bo tan i ca l Society of Amer ica, St. Louis, Missouri .

Panero, J. L. & Funk, V. A. 2002. Toward a phylogenet ic subfami l ia l c lassif icat ion for the Compos i tae (As te raceae) . Proc. Biol. Soc. Wash. 115 : 909-922.

Robinson, H. 1981. A revision of the tribal a n d s u b t r i b a l l im i t s of t he H e l i a n t h e a e (Asteraceae). Smi thson ian Cont r ibut ions of Botany 51 : 1-102.

Robinson, H., Schi l l ing, E. & Panero, J. L. 2009. Eupator ieae. In: Funk, V.A., Susanna, A . , S t u e s s y , T.F. & B a y e r , R . J . ( e d s . ) . Systemat ics, Evolut ion, and B iogeography of C o m p o s i t a e ^ S m i t h s o n i a n I n s t i t u t i o n , Washington, D.C., U.S.A.

0.2mm

0.1mm q

O . O S m m E

Figure 3. Cypsela morphology of Galinsoga sp. A-complete cypsela, B-base, C-apex, D-surface, E-surface hair, F-part of apical corona.

Figure 4. Cypsela morphology of Tridaxsp. A-complete cypsela, B-base, C-apex, D-surface, E&F-surface hairs, G-surface cells with hair base, H-base of pappus bristle, l-middle part of pappus bristle, J-apex of pappus bristle.

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Stuessy, T. F. 1977. Hel iantheae-systematic of the C o m p o s i t a e , Vo l . II. pp . 6 2 1 - 6 7 1 . review. In : Heywood, V. H., Harborne, J. B. & Academic Press, London. Turner, B. L.(eds.), The Biology and Chemistry

Table 1. Comparat ive morphologica l features of cypsela in four studied taxa.

Name of the Taxa Acmel la sp. Echinacea sp. Gal insoga sp. Tr idax sp.

Characters

Size

excluding

pappus (mm) 1.8-2.0x0.8-1.0 4 . 0 - 5 . 0 x 2 . 0 - 2 . 5 1 . 0 - 1 . 5 x 0 . 5 - 0 . 7 2.5-3.2 x 1.0-1.4

Shape Ob long Narrow obovate Narrow obovate Ob long

Colour Black Dirty white Black Deep brown

Sur face mark ings - - Alveolate Scorbiculate

Sur face hair Twin hairs Absent Twin hairs Twin & branched hairs

Ribs Absent Present Absent Absent

Carpopod ium Symmet r ic , r ing like

Symmetr ic , ring like

Symmetr ic , ring like

Absent

Sty lopodium Wel l deve loped, tubular

Weak ly deve loped

Weak ly deve loped

Absent

Pappus Represented by 2 awns

Coronate Scale like Represented by p lumose bristles

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