Asteraceae in Flora of Pakistan @ efloras.org

Asteraceae Dumortier

M. QAISER

Annual, biennial or perennial herbs, sometimes shrubs, rarely trees, with or without milky latex, sometimes armed with spines. Leaves alternate, opposite or rosulate, exstipulate, entire, toothed, lobed or variously dissected. Individual flowers (florets) usually small and numerous, sessile and aggregated on a receptacle, surrounded by an involucre made up of 1-several series of bracts (phyllaries or involucral bracts) and the whole structure forming a capitulum. Phyllaries may be herbaceous, fully scarious or at the margins only or chartaceous, usually with a sclerified portion in the centre (sterome) which may be undivided or divided into two parallel parts. Receptacle naked (epaleate) or bearing scales (paleate) or long hairs or bristles and receptacular surface with or without scale like ridges. Capitula with all florets of one sexual state, either male, female or hermaphrodite (homogamous) and heterogamous when florets of more than one sexual state are present. Homogamous capitula may be discoid or ligulate. Heterogamous capitula are radiate, radiant or disciform. Florets epigynous, bisexual, female or male (at least functionally) or sterile (neuter). Calyx often represented by a pappus of 1 or more series of bristles or scales on the apex of the ovary or sometimes pappus completely absent. Corolla gamopetalous, tubular, filiform, ligulate or rarely bilabiate, usually 3 or 5-toothed, rarely absent. Stamens 5, epipetalous, anthers connate forming a tube round the style, filaments free, rarely anthers free, dehiscence introrse, calcarate or ecalcarate, caudate or ecaudate or sometimes sagittate, endothecial tissues polar or radial; filaments usually with a collar. Ovary inferior, 1-celled. Style usually divided above into two branches with acute or obtuse sweeping hairs reaching below the furcation or not. Fruit cypsela, with or without ribs, sometimes with distinct carpopodium (basal attachment area). Asteraceae, the largest family of Angiosperms, comprises of ± 1535 genera and c. 23000 species distributed in 3 subfamilies and 17 tribes. The number of genera is rather increasing as more than 10 genera are described each year and several are resurrected from or reduced to synonymy (Bremer, K. 1994. Asteraceae-Cladistics & Classification, Timber Press, Portland, Oregon). It is also the largest plant family in Pakistan, represented by over 650 species distributed in 15 tribes. In view of its massive size it has been decided to publish it in parts covering one or more tribes. Few genera which are not assigned to any tribe are tentatively placed in the nearest tribe. Such cases are marked with an asterisk. The account of three tribes Gnaphalieae, Inuleae and Plucheeae shall be published in the next fascicle of Asteraceae which will also include a complete generic key of all the taxa found in Pakistan. The circumscription of the tribes given below follows the treatment of Bremer (1994). We understand that the placement of the genera in different tribes is not final as our knowledge about this largest and important family is far from complete. As the new evidences are being gathered, there is every possibility of reshuffling of some of the genera from one tribe to the other and even few more new tribes may be erected from time to time.

This volume is funded in part by the US National Foundation and the Andrew W. Mellon Foundation. Anthemideae is the first fascicle of this series.

1 All florets ligulate. Plants with milky latex. Lactuceae

+ At least the central florets tubular, not ligulate. Plants without milky latex (except in Arctoteae but not in our material). (2)

2 (1) Corolla of disc florets bilabiate with 2 adaxial lobes and 3 lobed abaxial limb. Mutisieae

+ Corolla of disc florets radially symmetrical, not bilabiate. (3)

3 (2) Anthers almost ecalcarate, apical appendages often demarcated from the connective and concave. Style hairs restricted to upper portion of branch and very often concentrated at the apices of branches rarely on the shaft below the branches. (6)

+ Anthers almost calcarate, apical appendage not demarcated from the connective and flat. Style hairs scattered on the branches much below the shaft. (4)

4 (3) Leaves mostly unarmed very rarely spiny. Phyllaries neither scarious appendaged nor spiny tipped. Vernonieae

+ Leaves mostly spiny or spinulose. Phyllaries mostly either scarious or spiny tipped. (5)

5 (4) Anthers distinctly caudate, rarely tails short, styles neither thick nor narrowing near the middle, usually with a slight thickening or ring of hairs near the point of bifurcatioin. Cardueae

+ Anthers ecaudate rarely shortly caudate. Styles thick, narrowing near the middle and slender to base, apex shortly divided. Arctoteae

6 (3) Cypselae polymorphic, outer cymbiform, somewhat curved, rostrate or triqueterous; innner distinctly curved, sometimes winged or annual. Calenduleae

+ Cypselae homomorphic, rarely dimorphic, inner ones not curved, ± straight. (7)

7 (6) Receptacle epalaceous rarely palaceous in Plucheeae (but not in our material). (8)

+ Receptacle palaceous. (15)

8 (7) Hairs on the style branches reach far down the shaft below the bifurcation. Plucheeae

+ Hairs on the style branches either present apically or till the middle but never reach down the shaft below the bifurcatioin. (9)

9 (8) Cypselae carbonized ± black or brownish black. (10)

+ Cypselae not carbonized, variously colourd not black. (11)

10 (9) Pappus usually of many scabrid-barbellate bristles, rarely of plumose bristles. Eupatorieae

+ Pappus generally of scales with or without an extended bristle like midrib rarely barbellate or plumose (not in our material). Helenieae

11 (9) Style asteroid type i.e. stigmatic portion prolonged into triangular to lanceolate to subulte sterile appendages. Astereae

+ Style not as above. (12)

12 (11) Stigmatic areas always in two separate lines or partially or wholly continuous ventrally but not confluent apically. (13)

+ Stigmatic areas always confluent apically. Inuleae

13 (12) Filament collar slender or often much dilated with swollen cells. Phyllaries mostly in one row rarely in 2-rows, wholly herbaceous. Senecioneae

+ Filament collar not as above. Phyllaries in several rows, rarely in two rows, either fully scarious or at least with the scarious margin and apex. (14)

14 (13) Anthers ecaudate, rarely shortly caudate. Pappus generally of rather few scarious scales, a crown or an auricle, rarely of many flat bristle-like scales, never of capillary bristles, often epappose. Anthemideae

+ Anthers caudate with short or long tail. Pappus plumose or barbellate, capillary bristles in one to several rows, rarely with scales and bristles or scales only or absent (but not in our material). Gnaphalieae

15 (7) Style asteroid type i.e. the stigmatic portion prolonged into triangular to lanceolate to subulate sterile appendages. Astereae

+ Style not as above. (16)

16 (15) Cypselas carbonized ± black or brownish black. (17)

+ Cypselas neither carbonized nor black. (19)

17 (16) Palea often flat and conduplicate, often with resinous ducts. Heliantheae

+ Palea narrow, not flat and conduplicate without resinous ducts. (18)

18 (17) Pappus generally of scales with or without an extended bristle-like midrib, rarely barbellate or plumore (not in our material). Capitula radiate, discoid or disciform. Helenieae

+ Pappus usually of many scabrid to barbellate bristles, rarely of plumose bristles or of both bristles and scales. Capitula always discoid. Eupatorieae

19 (16) Filaments collar slendar or often much dilated with swollen cells. Phyllaries rarely in 2-rows. Senecioneae

+ Filaments collr not as above. Phyllaries pleuriseriate rarely in two rows. (20)

20 (19) Stigmatic areas always confluent apically. Inuleae

+ Stigmatic areas in two separate lines, never confluent apically. (21)

21 (20) Anthers mostly caudate with short or long tails rarely ecaudate (Phagnalon). Pappus of capillary bristles in one to several rows, rarely with scales and bristles or scales only (not in our material). Gnaphalieae

+ Anthers mostly ecaudate, rarely shortly ecaudate. Pappus not of capillary bristles of few scarious scales or many flat bristle-like scales or pappus absent. Anthemideae

List of lower taxa