Bidyut Kr. Jana and Sobhan Kr. Mukherjee. / Journal of Science / Vol 4 / Issue 5 / 2014 / 327-338.

e ISSN 2277 - 3290 Print ISSN 2277 - 3282 Journal of Science Botany

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DIVERSITY OF TESTAL STRUCTURE AMONG SOME TRIBES OF COMPOSITAE

*Bidyut Kr. Jana and Sobhan Kr. Mukherjee

Department of Botany, University of Kalyani, Kalyani-741235, West Bengal, India,

ABSTRACT Seed coat of mature cypselar wall of Compositae is usually termed as testa. It is usually found towords the inner side of pericarp of cypsela. In majority of the taxa, this testal layer exists either attached with the pericarp or secondarily separated from the pericarp. In most cases it is represented by a narrow to wide zone of crusted parenchyma cells or as a dense narrow layer, where cells are not clearly visible, termed as pellicle. Sometimes testal zone is clearly represented by outer and inner zone. Among the tribes of Compositae, the structure of testal zone is very characteristic in Astereae, Arctoteae, Cardueae etc. The structure of testa is less specialized in , , Tageteae, Anthemideae, Lactuceae etc. For this study, cypselas of 82 of the family Compositae have been under taken.

Keywords: Testal structure; 82 species; Compositae.

INTRODUCTION Grau [1] had recognized the seed coat of wide zone, which can be designated as middle zone of Compositae as testa. He suggested the outer layer of seed testa but actually middle layer of seed coat and inner coat as testa epidermis (TE) and inner part of seed coat as testal zone which is actually inner zone of seed coat. inner testa (T). Reese [2] had coined the term ‘sporoderm’ Outer layer of seed coat i.e. usually represented in the structure of seed coat. Here, the term testa has been by single layer of intact cells or collapsed cells or testal followed after Grau [3] in the structure of seed coat with epidermal cells, which are associated with inner collapsed some modification by others. Here, outer part of seed coat cells of many layers. There is some exception, where seed is recognized as testa epidermis (TE), middle part of seed coat is constitute by outer well organized testa, followed coat designated as testal middle part (TM), whereas inner by middle zone of intact or partially intact cells and inner part clearly differentiated from middle part has been zone of seed coat is constituted by another type of coined as testal inner part (TI). Testa is a term to qualify parenchyma cells or non-cellular pellicle like structure as the seed coat. Actually it is a protective outer coat or an in Helipterum sp. In case of Vernonia bainesii, Mikania integument of a seed of flowering . trinervis etc, testal inner zone is well organized i.e. inner According to Cabrera [4] outer layer of seed coat layer of cells have been maintained their structure in is designated as testa, which is usually found in mature mature cypsela. cypselas in cross sectional configuration. Usually in Majority of taxonomist have been included the mature cypsela, testa is represented by a single layer of structure of testa during the description of anatomical dimersified type of cells or it is represented by a narrow features of cypselas, but there are a few papers in or wide zone of collapsed tissue. However in some cases, Compositae, which deals with the structure of testa. testa is associated with inner two narrow or moderately Therefore there is a scope to analyze

Corresponding Author:- Bidyut Kr. Jana Email:[email protected]

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the characeteristic features of testa in some taxa of primitive one in comparison to advanced taxa, which Compositae. Since testal features have significant value have disorganized testal epidermis i.e. testal epidermal for the betterment and upgradation of Compositae cells lack sclerification and wall thickening. This view systematic. was adapted on the studied of fruit development in Calenduleae and Arctotideae as well as in two species On the basis of testal structure, the studied taxa of of Anthemideae. On the perspective of modern idea, Compositae can be divided into following types: regarding Compositae classification [5-8], it has been Type: 1. Testal layer is made up of crusted layer of indicated that these two tribes have been placed in parenchyma cells, uni-seriately arranged. E.g.- advanced position. So, present study support the alpina,Crepis aspera, Crepis pyrenaica, Leontodon placement of these two tribes in advanced group instead muelleri etc. of ancient position in earlier system of Compositae Type: 2. Testal layer is made up of thin-walled, classification. horizontally placed, parenchyma cells. Testal cells usually lacks Ca-oxalate crystals, E.g.-Hypochaeria uniflora, Tragopogon porrifolius, but the presence of crystals in testal region has been wightianus, Sonchus oleraceous, Leontodon reported in Pulicaria dysenterica of the tribe Inuleae by crispus, Hieracium racemosum etc. Jana, Bar and Mukherjee [9]. In the cypselas ofInula Type: 3. Testal layer is made up of thick-walled, ensifolia, helenium, Pulicaria dysenterica, Telekia horizontally placed, parenchyma cells. E.g.- speciosa, Solidago gigantea, Solidago Canadensis, albanicus, Helianthus debilis etc. Solidago virgaurea, Aster alpinus, Lactuca serriola, Type: 4. Testal layer is made up ‘U’-shape parenchyma Pieris hieraciodes etc. seed coat are attached with cells, uniseriately arranged. pericarp, but in some cypselas (Inula Britannica, E.g.- Cichorium endivia, Aster alpinus, Leontodon muelleri etc.), seed coats are secondarily heterophylla, ramulosa, krebsiana , separated from the pericarp. Secondarily separated seed Leptorhynchos elongates etc. coat from pericarp has been considered by Grau as Type: 5.Testal layer is made up of bi-seriately fairly advanced taxa than the seed coat is attached with arranged, parenchyma cells. E.g. Tragopogon the pericarp. orientalis, Tragopogon pratensis, Vernonia galamensis The structure of mature testal layer may be etc. developed in the following ways. In some taxa Type: 6. Testal layer is made up of palisade (Leontodon muelleri,Crepis palaestina, Crepis parenchyma cells. E.g. aspera, Centau dioscoridis, Crepis alpina, Crepis foetida,etc.), entire sabiosa, Centaurea stobe, Arctium lappa , seed coat layers disintegrate and form a testa of arvanse , Circium candelabrum, Cirsium heleniodes , collapsed cells only. In some other taxa, seed coat or Montisalea salmantica etc. outer coat or integumental epidermis, remain as such as Type: 7. Testal layer is biseriately arranged and is made previous state and form a cellular layer, which are up of both normal and crusted layer of parenchyma formed in majority of the studied taxa, while in some cells. E.g. Carpesium cernuum other taxa, entire or part of seed coat remain as such as Type: 8. Testal layer is biseriately arranged and is made in younger stage or more or less younger.So, the mature up of both the normal and palisade parenchyma cells. cypselas (In early state), cypselar coat is clearly E.g. Serratula tinctoria differentiated into outer testa of single layer, middle zone of testa and inner zone of testa. Actually the MATERIALS AND METHODS middle and inner zone of seed coat are represented by Cypselas were procured from different foreign inner part of testal epidermal cells. herbaria of the world as gift. The names of studied taxa Testa is highly specialized and thick walled in with their sender address are given in the table-1. The some tribes (Astereae, Cardueae etc.) and less specific studied taxa are arranged in alphabet order of the and less distinct in the tribes like Vernonieae, genera. Senecioneae, Inuleae, Lactuceae etc. For the observation of testal features, cross sections In majority of taxa (Chondrilla juncea, have been taken from the middle part of cypselas with Lactuca pseudoumbrella, Cichorium intybus, the help of sharp razor blade to observe the different dalechampii, Lapsana communis, anatomical regions, mainly the testal structure in mature Tragopogon porrifolius, platyglossa, cypselas. elegans etc.), testal epidermal cells are arranged in tangential fashion. There are some taxa (Elephantopus RESULTS AND DISCUSSION carolinianus,Pieris hieraciodes, Centaurea aspera, On the basis of the nature of tissue organijation Centaurea sabiosa, Centaurea stoebe etc.), which have and layers of arrangements of cells, different types of radially oriented testal epidermal cells. Such type of testal structures have been noted, which are given in the taxa may be more advanced in comparison to table-2. tangentially oriented testal arrangement of cells. According to Reese (1989), taxa having well Therefore, it is essential that testal structure organized testa epidermis has been considered as has significant role for characterization of taxa along

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with pericarpic features of Compositae as advanced in by Briquet, Schuttz-Bipontinus and some others [10- earlier time, on the basis of cypselar anatomical studies 11].

Table 1. Name of taxa and Sender address Sl.no Name of taxa Sender address Acanthospermum hispidum DC. Instituto De Investigacao Cientifica De Angola, Herbarium 1 (Millerieae) (LUAI). Col: A. Menezes, Dated 28. 6. 1972. No-4223

Anthemis maritima L. Botanischer Garten der Universitat Zurich. XXOBRISS- 2 (Anthemideae) 20053261 Arctium lappa L. Humboldt- Universitat zu Berlin. Institut fur Biologie, Spezielle 3 (Cardueae) Botanik u. Arboretum, Berlin, Germany. Collection Number- 704. Artemisia vulgaris L. Botanic Garden Of The University, Jyeangontie 2, SF- 00550 4 (Anthemideae) Helsinki, Finland. Aster albanicus Degen. Botanic Garden of the University of Copenhagen, Danmark. 5 (Astereae) 211S2000-0862*A Humboldt- Universitat Zu Berlin, Institut fur biologie, Spezielle 6 Aster alpinus L.(Astereae) Botanik u. Arboretum, Spathstra βe 80/81, D-12437, Berlin, Germany.

Bahia absinthifolia Benth. 7 Swedish Museum of Natural History, Stockholm, Sweden. (Bahieae) purpurea (DC.) Mast. Botanischer Garten der Universitat Zurich (Z). Collection 8 (Arctotideae) Number: XXOZ-20041189

Bidens frondosa L. 9 Humboldt- Universitat Zu Berlin, Institut fur biologie, Spezielle (Coreopsideae) Botanik u. Arboretum, D-12437, Berlin, Germany.

Carpesium cernuum L. Botanischer Garten der Universitat Zurich, Zollikerstrasse 107, 10 (Inuleae) CH-8008, Zurich, Switzerland. XXOZ-20051457 Centaurea sabiosa L. Humboldt- Universitat Zu Berlin, Institut fur biologie, Spezielle 11 (Cardueae) Botanik u. Arboretum, D-12437, Berlin, Germany. 72.

Centaurea spinosa L. 12 Botanischer Garten der Universitat Zurich, Switzerland. (Cardueae) Centaurea aspera L. Botanischer Garten der Universitat Zurich, Zollikerstrasse 107, 13 (Cardueae) CH- 8008, Zurich, Switzerland, XX0BRISS- 20093677

Centaurea stobe L. Humboldt- Universitat Zu Berlin, Institut fur biologie, Spezielle 14 (Cardueae) Botanik u. Arboretum, D-12437, Berlin, Germany. 72.

Chaenactis fremontii A. Gray 15 Swedish Museum of Natural History, Stockholm, Sweden (Chaenactideae) Humboldt- Universitat Zu Berlin. Institute fur Chondrilla juncea L. 16 Biologie spezielle Botanik u. Arbor-etum, Spathstra (Lactuceae) Be 80/81. 12437 Berlin,Germany. Cichorium endivia L. Institut fur pflanzengenetik und Kulturpflanzenforschung 17 (Lactuceae) Gatersleben FLORA of . Cichorium intybus L. Lower Austria, Thermenlinie, Wineyards east of 18 (Lactuceae) the Heberlberg 2 km southwest of Gumpoldskirchen, 275 m s.m. 48°02'N, 16" 16' E, 23 September 2001, Along roadside Botanic Garden and Museum of the University of Copenhagen Circium candelabrum Griseb. 19 (Natural History Museum of Denmark). Collection Number- 268 (Cardueae) S2000-0876 A W Kit Tan 24698. Cirsium arvanse (L.) Scop. Conservatoire Et Jardin Botaniques Geneve- Geneva, Switzerland. 20 (Cardueae) Collection Number- 107. Cirsium heleniodes (L.) Hill Botanischer Garten der Universitat Zurich. Collection Number- 21 (Cardueae) CHOZ-20031567 Cirsium spinosissimum (L.) Scop 22 Botanischer Garten der Universitat Zurich, Switzerland. (Cardueae) Crepis alpina L. Botanic Garden of the University of Copenhagen, Denmark. 23 (Lactuceae) 288 E 3022-0024* AG.

Crepis pulchra L. Botanic Garden of the University of Copenhagen, Denmark. 24 (Lactuceae) 298 E 3022-0011* AG.

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Crepis pyrenaica (L.) Greuter Botanischer Garten der Universitat Zurich, Switzerland (Z). 25 (Lactuceae) Collection Number: CHOZ-20110589 Crepis aspera L. Botanic Garden of the University of Copenhagen, Denmark. 26 (Lactuceae) 289 E 3022-0021* AG.

Crepis dioscoroidis L. Botanic Garden of the University of Copenhagen, Denmark. 27 (Lactuceae) 294 E 3022-0022* AG.

Crepis foetida L. Botanic Garden of the University of Copenhagen, Denmark. 28 (Lactuceae) 295 E 3022-0027* AG.

Crepis neglecta L. Botanic Garden of the University of Copenhagen, Denmark. 29 (Lactuceae) 296 E 3022-0004* AG.

Crepis palaestina (Bois.) Bornm Botanic Garden of the University of Copenhagen, Denmark. 30 (Lactuceae) 297 S1921-1474* AG.

Felicia heterophylla (Cass.) Grau 31 Botanic Garden of the University of Copenhagen, Denmark. (Astereae) Gazania krebsiana Less. Botanischer Garten der Universitat Zurich (Z). Collection 32 (Arctotideae) Number: XXOZ-19810083 Hieracium racemosum W. & K. ex Willd. Institut fur Pharmakog nosie der Universitat Wien, 33 (Lactuceae) Pharmaziezentrum, Althanstra Be 14, A- 1090 Wien, Austria.

Hypochaeria uniflora Vill. Botanischer Garten der Universitat Zurich, Switzerland (Z). 34 (Lactuceae) Collection Number: CHOZ-20120603 Inula britannica L. Botanischer Garten der Universitat Zurich, Switzerland (Z). 35 (Inuleae) 2008 Drawer. XX0Z- 20060389.

Inula ensifolia L. Botanischer Garten der Universitat Wien Rennweg 14, A- 1030 36 (Inuleae) Wien, Austria (HBV).

Inula helenium L. 37 Botanischer Garten der Universitat Zurich, Switzerland (Z). (Inuleae) Lactuca pseudoumbrella D.Maity & Maiti From Sikhim Himalaya, supplied by the Authors of 38 (Lactuceae) the Taxon Lactuca serriola L. Humboldt- Universitat zu Berlin. Institut fur Biologie. Spezielle 39 (Lactuceae) Botanik u. Arboretum. Berlin, Germany. Lapsana communis L. Botanischer Garten der Universitat Zurich (Z). 40 (Lactuceae) XXOZ 19964888 Layia platyglossa (Fisch. & C. A. Mey.) Botanischer Garten der Universitat Zurich, Switzerland (Z). 41 A. Gray (Madieae) XX0Z- 20050466.

Leontodon crispus (Wild.) Finch et 42 Botanic Garden of the University of Copenhagen, Denmark. P.D.Sell (Lactuceae) Leontodon muelleri (Sch.Bip) Fiori 43 Botanic Garden of the University of Copenhagen, Denmark. (Lactuceae) Leptorhynchos elongates DC. 44 Herbarium CBG, CBG- 8906276 (Gnaphaleae) Madia elegans D. Don Botanic Garden of the University of Copenhagen, Denmark. 45 (Madieae) 387 SI 972- 0309 * AG.

Microseris lanceolata L. 46 South : Wilpena pound 310 30’ s, 1380 33’ E (Lactuceae) Botanic Garden and Museum of the University of Copenhagen ( 47 Montisalea salmantica (L.) Brig. Et casill. Natural History Museum of Denmark). Collection Number- 389 E 2867-B001 A G Botanic Garden of Adelaide, South Australia: South-east Olearia ramulosa Benth. 48 Region, Little Dip Conservation Park road, side by Lake edge. (Astereae) (370 10’S, 1390 45’E)

Picris hieraciodes L. 49 Institut fur Pharmakognosie der Universitat Wien, Austria. (Lactuceae) angustifolia (Labill.) Less. 50 Botanic Garden of Adelaide, South Australia. (Gnaphaleae) Prenanthes purpurea L. Botanischer Garten der Universitat Zurich. 51 (Lactuceae) CHOZ-20100751 Pulicaria dysenterica (L.) Bernh. 52 Botanischer Garten der Universitat Zurich. (Inuleae) 53 Saussurea fastuosa (Decne.) Hand.-Mazz. Sikhim. Collection Number- 52.

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(Cardueae) Schkuhria pinnata (Lam.) Kuntze 54 Swedish Museum of Natural History, Stockholm, Sweden. (Bahieae) Serratula tinctoria L. 55 Botanischer Garten der Universitat Zurich, Switzerland. (Cardueae) Humboldt- Universitat Zu Berlin, Institut fur biologie, Spezielle Solidago canadensis L. 56 Botanik u. Arboretum. Spathstra Be 80/ 81, D- 12437 Berlin, (Astereae) Germany. 736.

Humboldt- Universitat Zu Berlin, Institut fur biologie, Spezielle Solidago gigantea Aiton 57 Botanik u. Arboretum., Spathstra Be 80/81, D- 12437 Berlin, (Astereae) Germany. 737.

Solidago virgaurea L. 58 (Astereae) Botanischer Garten der Universitat Zurich. CH0Z- 20080796

Sonchus oleraceus L. 59 Botanischer Garten der Universitat Wien Rennweg 14, A- 1030 (Lactuceae) Wien (Austria).

Sonchus wightianus DC. 60 Kalyani Township. Kalyani, Nadia, WB, India. B.J- 201 (Lactuceae) lucida L. Botanischer Garten der Universitat Zurich With collection 61 (Tageteae) number-XX0Z-20031315 Tagetes tenuifolia L. Botanischer Garten der Universitat Zurich With collection 62 (Tageteae) number- XX0Z-20110213 Tanacetum parthenium Sch. Bip. 63 Botanischer Garten der Universitat Zurich. XXOZ 19965019 (Anthemideae) Tanacetum vulgare L. Humboldt- Universitat zu Berlin. Institut fur Biologie. Spezielle 64 (Anthemideae) Botanik u. Arboretum. Berlin, Germany. Telekia speciosa (Schreb.) Botanischer Garten der Universitat Zurich, Switzerland. 2009, 65 Baumg (Inuleae) Drawer (Z). XX0BRISS- 0000983.

Tragopogon orientalis L. 66 Institut fur Pharmakognosie der Universitat Wien, Austria. (Lactuceae) Tragopogon porrifolius L. Botanischer Garten der Universitat Zurich, Switzerland (Z). 67 (Lactuceae) Collection Number: XXOZ- 20010072 Tragopogon pratensis L. Humboldt- Universitat zu Berlin. Institut fur Biologie. Spezielle 68 (Lactuceae) Botanik u. Arboretum. Berlin, Germany. Tridax procumbens L. Swedish Museum of Natural History, P.O. Box 50007, SE- 69 (Millerieae) 10405, Stockholm, Sweden.

Tripleurspermum inodorum (L.) Sch. Bip. Botanic Garden of The University, Jyrangontie 2, SF- 00550 70 (Anthemideae) Helsinki, Finland Tripleurspermum maritimum (L.) W. D. J. Humboldt - Universitat zu Berlin. Institut fur Biologie. Spezielle 71 Koch (Anthemideae) Botanik u. Arboretum. Berlin, Germany. Urospermum dalechampii (L.) 72 F.W.Schmidt Botanischer Garten der Universitat Zurich (Z).XXOZ 19950008 (Lactuceae) Vernonia galamensis Less. 73 North Central Regional, Introduction Station, Ames. (Vernonieae) Vernonia hymenolepis Vatke 74 North Central Regional, Plant Introduction Station, Ames. (Vernonieae) Vernonia stenolepis Oliv. 75 North Central Regional, Plant Introduction Station, Ames. (Vernonieae) Vernonia anthelmintica (L.) Willd. 76 Kalyani Township. Kalyani, Nadia, WB, India. B.J.-33 (Vernonieae) Vernonia cinerea Less. 77 Kalyani Township. Kalyani, Nadia, WB, India. B.J.-30. (Vernonieae) Villanova oppositifolia Lag. Swedish Museum of Natural History, P.O. Box 50007, SE- 78 (Perityleae) 10405, Stockholm, Sweden.

Xeranthemum annuum L. Botanischer Garten der Universitat Zurich. Collection Number- 79 (Cardueae) XXOZ 20040897 80 haageana Regel Botanic Garden of Copenhagen, Copenhagen, Denmark.

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(Heliantheae) Zinnia pauciflora Regel 81 Botanic Garden of Copenhagen, Copenhagen, Denmark. (Heliantheae) 82 Helianthus debilis Nutt. North Central Introduction station. Ames.

Table 2. Testal structure in mature cypselas in different taxa Nature of tissue Layer of Sl.No. Name of Taxa Tribe organization arrangement Crusted layer of 1 Crepis alpine (Figure-1 D) Lactuceae Uni-seriately arranged parenchyma tissue Crusted layer of 2 Crepis aspera (Figure-1 E) Do Uniseriately arranged parenchyma tissue Crusted layer of 3 Crepis dioscoridis (Figure-1 B) Do Uniseriately arranged parenchymatous Crusted layer of 4 Crepis foetida (Figure-1 F) Do Uniseriately arranged parenchymatous 5 Crepis neglecta (Figure-1 G) Do parenchymatous Uniseriately arranged Crusted layer of 6 Crepis palaestina (Figure-1 A) Do Uniseriately arranged parenchymatous Crusted layer of 7 Crepis pulchra (Figure-1 C) Do Bi-seriately arranged parenchymatous 8 Leontodon crispus (Note attach) Do Parenchymatous Uniseriately arranged 9 Hieracium racemosum (Figure-2 A) Do Parenchymatous Uniseriately arranged Crusted parenchyma 10 Leontodon muelleri (Figure-2 B) Do Uniseriately arranged cells Crusted parenchyma 11 Crepis pyrenaica (Figure-3 A) Do Uniseriately arranged cells 12 Hypochaeria uniflora (Figure-3 B) Do Parenchymatous Uniseriately arranged Tragopogon porrifolius 13 Do Parenchymatous Uniseriately arranged (Figure-3 C) 14 Sonchus wightianus (Figure-4 B) Do Parenchymatous Uniseriately arranged 15 Sonchus oleraceous (Figure-4 A) Do Parenchymatous Uniseriately arranged 16 Chondrilla juncea (Not attach) Do Parenchymatous Uniseriately arranged 17 Cichorium intybus (Not attach) Do Parenchymatous Uniseriately arranged Lactuca pseudoumbrella 18 Do Parenchymatous Uniseriately arranged (Not attach) 19 Lapsana communis (Not attach) Do Parenchymatous Uniseriately arranged Urospermum dalechampii 20 Do parenchymatous Uniseriately arranged (Not attach) Crusted layer of 21 Lactuca serriola (Fig.5-A) Do Uniseriately arranged parenchyma cells 22 Picris hieraciodes (Fig.5-B-C) Do Parenchymatous Uniseriately arranged 23 Tragopogon orientalis(Fig.5-D-E) Do Parenchymatous Bi-seriately arranged 24 Tragopogon pratensis (Fig.5-F) Do Parenchymatous Bi-seriately arranged U-shape parenchyma 25 Cichorium endivia(Fig.6-A) Do Uni-seriately arranged cells Crusted parenchyma 26 lanceolata (Fig.6-B-C) Do Uni-seriately arranged cells Crusted parenchyma 27 Prenanthes purpurea (Fig.6-D) Do Uni-seriately arranged cells 28 Bidens frondosa (Not attach) Coreopsideae Parenchymatous Uniseriately arranged U-shaped 29 Solldago virgaurea (Fig.7-A) Astereae Uniseriately arranged parenchymatous U-shaped 30 Solidago canadensis(Fig.7-B) Do Uniseriately arranged parenchymatous 31 Solidago gigantean (Fig.7-C) Do Parenchyma cells Uniseriately arranged

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U-shaped parenchyma 32 Aster alpines (Fig.8-A) Do Uniseriately arranged cells U-shaped parenchyma 33 Felicia heterophylla (Fig.8-B) Do Uniseriately arranged cells U-shaped parenchyma 34 Olearia ramulosa (Not attach) Do Uniseriately arranged cells 35 Aster albanicus (Fig. 9 A-B) Do Parenchyma cells Uniseriately arranged Palisade like 36 Centaurea aspera (Fig. 23-A) Cardueae Uniseriately arranged parenchyma cells Palisade like 37 Centau sabiosa (Fig. 23-B) Do Uniseriately arranged parenchyma cells Palisade like 38 Centaurea stobe (Fig. 23-C) Do Uniseriately arranged parenchyma cells Palisade like 39 Arctium lappa (Fig. 24-A) Do Uniseriately arranged parenchyma cells Palisade like 40 Cirsium arvanse (Fig. 24-B) Do Uniseriately arranged parenchyma cells Palisade like 41 Circium candelabrum (Fig. 24-C) Do Uniseriately arranged parenchyma cells Palisade like 42 Cirsium heleniodes (Fig. 24-D) Do Uniseriately arranged parenchyma cells Palisade like 43 Montisalea salmantica (Fig. 24-E) Do Uniseriately arranged parenchyma cells 44 Saussurea fastuosa (Fig. 24-F) Do Parenchyma cells Uniseriately arranged 45 annuum (Fig. 24-G) Do Parenchyma cells Uniseriately arranged Palisade like 46 Centaurea spinosa (Fig. 25-A-B) Do Uniseriately arranged parenchyma cells Palisade like 47 Cirsium spinosissimum (Fig. 25-C) Do Uniseriately arranged parenchyma cells Normal and palisade 48 Serratula tinctoria (Fig. 25-D) Do Bi-seriately arranged parenchyma cells 49 Vernonia anthelmintica (Fig. 22-A) Vernonieae Parenchymatous Uniseriately arranged 50 Vernonia cinerea (Fig. 22-B) Do Parenchymatous Uniseriately arranged 51 V. galamensis (Fig. 22-C) Do Parenchymatous Bi-seriately arranged 52 V. hymenolepis (Fig. 22-D) Do Parenchymatous Uniseriately arranged 53 V. stenolepis (Fig. 22-E) Do Parenchymatous 4-6 layer arrangement Acanthospermum hispidum 54 Millerieae Parenchymatous Uniseriately arranged (Not attach) 55 Tridax procumbens (Fig. 21-C-D) Do Parenchymatous Uniseriately arranged 56 Inula britannica (Fig. 19-A) Inuleae Parenchymatous Uniseriately arranged 57 Inula ensifolia (Fig. 19-B) Do Parenchymatous Bi-seriately arranged 58 Pulicaria dysenterica (Fig. 20-A) Do Parenchymatous Uniseriately arranged 59 Telekia speciosa (Fig. 20-B) Do Crsted parenchyma cells Uniseriately arranged 60 Inula helenium (Fig. 19-C) Do Parenchymatous Uniseriately arranged Both the normal and 61 Carpesium cernuum (Fig. 9 C-D) Do Bi-seriately arranged crusted parenchyma cells 62 absinthifolia (Fig. 16 C-D) Bahieae Parenchymatous Uniseriately arranged 63 Schkuhria pinnata (Fig. 16 A-B) Do Parenchymatous Uniseriately arranged 64 Berkheya purpurea (Fig. 18 A-C) Arctotideae Parenchymatous Bi-seriately arranged U-shape parenchyma 65 Gazania krebsiana (Fig. 18 D-E) Do Uniseriately arranged cells 66 Layia platyglossa (Fig. 17 A-B) Madieae Parenchymatous Uniseriately arranged Crusted parenchyma 67 Madia elegans (Fig. 17 C-D) Do Uniseriately arranged cells 68 Tagetes lucida (Fig. 15 A-B) Tageteae Parenchyma cells Uniseriately arranged

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Crusted parenchyma 69 Tagetes tenuifolia (Fig. 15 C-B) Do Uniseriately arranged cells 70 Helianthus debilis (Fig. 9-E) Heliantheae Parenchyma cells Bi-seriately arranged 71 Zinnia haageana (Fig. 10 A-B) Do Parenchyma cells Bi-seriately arranged 72 Zinnia pauciflora (Not attach) Do Parenchyma cells Bi-seriately arranged Leptorhynchos elongates U-shape parenchyma 73 Gnaphaleae Uniseriately arranged (Fig. 14 C-D) cells Crusted and palisade 74 Do Bi-seriately arranged (Fig. 14 A-B) parenchyma cells 75 Anthemis maritime (Fig. 13-A) Anthemideae Parenchyma cells Uniseriately arranged 76 Artemisia vulgaris (Fig. 13-B) Do Parenchyma cells Uniseriately arranged 77 Tanacetum parthenium (Fig. 13-C) Do Parenchyma cells Uniseriately arranged 78 Tanacetum vulgare (Fig. 13-D) Do Parenchyma cells Uniseriately arranged Tripleurspermum inodorum 79 Do Parenchyma cells Uniseriately arranged (Fig. 13-E) Tripleurspermum maritimum 80 Do Parenchyma cells Uniseriately arranged (Fig. 13-F) 81 Villanova oppositifolia (Fig. 12) Perityleae Parenchyma cells Uniseriately arranged 82 fremontii (Fig. 11) Chaenactideae Parenchyma cells Uniseriately arranged Fig. 1. A-G-Cypselar anatomy showing the testal Fig. 2. A-B- Cypselar anatomy showing the testal structure structure

A. Crepis palaestina, B. Crepis dioscoridis, C. Crepis pulchra, D. Crepis alpina, E. Crepis aspera, F. Crepis A. Hieracium racemosum , B-Leontodon muelleri foetida, G. Crepis neglecta.

Fig. 3. A-C- Cypselar anatomy showing the testal Fig. 4. A-B- Cypselar anatomy showing the testal structure structure

A- Crepis pyrenaica, B- Hypochaeris uniflora, C- Tragopogon porrifolius A- Sonchus oleraceous, B-Sonchus wightianus 334

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Fig. 5. A-F- Cypselar anatomy showing the testal Fig. 6. A-D- Cypselar anatomy showing the testal structure structure

A-D- 0.05 mm A- Lactuca serriola, B - C – Pieris hieraciodes , D- A- Cichorium endivia, B-C- Microseris lanceolata (B- ETragopogon orientalis, F- Tragopogon pratensis Through rib, C-Through furrow), D- Prenanthes purpurea Fig. 7. A-C- Cypselar anatomy showing the testal Fig. 8. A-B- Cypselar anatomy showing the testal structure structure

A-Solidago virgaurea, B-Solidago canadensis, C-Solidago A-Aster alpinus, B-Felicia heterophylla gigantea Fig. 9. A-E- Cypselar anatomy showing the testal Fig. 10. A-F- Cypselar anatomy showing testal structure structure

1mm B,C,E. 0.05 mm A,D,F. A,C. 1mm A-B-Zinnia haageana: A-Diagramatic view, B-Cellular A-B- Aster albanicus: A-Diagramatic view, B-Cellular view; view; C-D Leptorhynchos elongates: C-Diagramatic view, C-D- Carpesium cernuum: C-Diagramatic view, D-Cellular D-Cellular view; E-F- Podotheca angustifolia: E- view; E- Helianthus debilis (Cellular view) Diagramatic view, F-Cellular view. Fig. 11. Cypselar anatomy of Chaenactis fremontii Fig. 12. Cypselar anatomy of Villoneva oppositifolia showing testal layer showing testal layer

0.05 mm

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Fig. 13. A-G-Cypselar anatomy of studied species Fig. 14-A-D-Cypselar anatomy of the studied species showing testal layer showing testal layer A-B-Podotheca angustifolia: A-Diagramatic view, B- Cellular view; C-D-Leptorhynchos elongates: C- Diagramatic view, D-Cellular view.

0.05 mm A,B,C,D,E,F,G. A- Anthemis maritime, B-Artemisia vulgaris, C-Tanacetum parthenium, D-Tanacetum vulgareE-Tripleurspermum inodorum, F-Tripleurspermum maritimum 0.05 mm B,D. 1 mm A,C. Fig. 15-A-D- Cypselar anatomy showing testal structure Fig. 16-A-D- Cypselar anatomy showing testal structure

1mm A,C. 0.05 mm B,D. A-B-Tagetes lucida: A-Diagramatic view, B-Cellular view; 1mm A,C 0.05 mm B,D C-D- Tagetes tenuifolia: C-Diagramatic view, D-Cellular A-B--Schkuhria pinnata: A- Diagramatic view, B-Cellular view. view; C-D- Bahia absinthifolia: C-Diagramatic view, D- Cellular view. Fig. 17-A-D- Cypselar anatomy showing testal structure Fig. 18 A-E- Cypselar anatomy showing testal structure

1 mm A,D. 0.05 mm B,C, E 1 mm A,C 0.05 mm B,D A-C-Berkheya purpurea: A-Diagramatic view, B-Cellular A-B-- Layia Platyglossa: A-Diagramatic view, B- Cellular view (Through the rib), C-Cellular view (Through the view; C-D- Madia Elegans: C-Diagramatic view, D-Cellular furrow); D-E- Gazaniakrebsiana: D-Diagramatic view, E- view. Cellular view.

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Fig. 19. A-C- Cypselar anatomy showing testal structure Fig. 20. A-B- Cypselar anatomy showing testal structure

A-Inula Britannica, B-Inula ensifolia, C- Inula helenium A-Pulicaria dysenterica, B- Telekia speciosa Fig. 21. A-D- Cypselar anatomy showing testal structure Fig. 22. A-E- Cypselar anatomy showing testal structure

1 mm A,C. A-B- Melampodium perfoliatum: A-Diagramatic view, B- 0.05mm B,D Cellular view; C-D-Tridax procumbens: C-Diagramatic A-Vernonia anthelmintica, B- Vernonia cinerea, C- view, D-Cellular view Vernonia galamensis, D-Vernonia hymenolepis, E- Vernonia stenolepis Fig. 23- A-C-Cypselar anatomy showing testal structure Fig. 24. A-G- Cypselar anatomy showing testal structure

A- Centaurea aspera, B-Centaurea saobiosa, C-Centaurea 0.05 mm A-G. stobe A: Arctium lappa, B : Cirsium arvanse , C: Circium candelabrum , D: Cirsium heleniodes E: Montisalea salmantica , F: Saussurea fastuosa , G: Xeranthemum annuum

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Fig. 25-A-D- Cypselar anatomy showing testal structure

0.05 mm A, C, D. 0.05 mm B. A- Centaurea spinosa, B- Magnified view of testal layer of Centaurea spinosa, C- Cirsium spinosissimum, D- Serratula tinctoria

ACKNOWLEDGEMENT We are thankful to Dr. Teresa Martins, Walter Till (Professor and Curator of the Herbarium Curator,(Institute De Investigacao Cientifica De (WU), Faculty Center Botany, Department of Plant Angola, Herbarium (LUAI), Luanda Herbarium); Dr. Systematics and Evolution, Faculty of Life Science, Peter Enz, Curator (Botanischer Garten der Universitat Universitat Wien); Dr. HBS Womersley, Chair in Plant Zurich (Z), Switzerland); Dr. Robert Vogt, Curator ( Systematics (State Herbarium of South Australia, Humboldt- Universitat Zu Berlin, Institut fur Biologie, Adelaide, Australia (AD);Dr. Marko Hyvarinen, Spezielle (B), Germany); Prof. Bertil Nordenstum, Director, Botanical Museum, Finnish Museum of Retd. Emeritus Professor (Swedish Museum of Natural Natural History (H), University of Helsinki, Finland; History, Stockholm, Sweden); Prof.(Dr.) Hans Vilhelm Director of the herbarium of CBG; Curator of Institut Hansen, Curator (University Botanic Garden of fur pflanzengenetik und Kulturpflanzenforschung Copenhagen (DK); Director of ( North Central Regional Gatersleben for sending mature, identified seeds for this Plant Introduction Station, Ames, IA.50011, USA); Dr. study.

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