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Published In: O Prirozenosti Rostlin 254. 1820. (Jan-Mar 1820) (Prir. Rostlin) Name publication detail

Project Name Data (Last Modified On 6/2/2011)
Acceptance : Accepted
Project Data     (Last Modified On 4/11/2022)
Contributor Text: M. QAISER
Comment/Acknowledgements: 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.

we are thankful to the Directors and Curators of the following herbaria for providing herbarium and library facilities: B, DMHN, BM, E, GZU, HUP, K, KUH, RAW, SWAT. The financial support of Pakistan Academy of Sciences (PAS) for the publication of this volume is gratefully acknowledged. Thanks are also due to Prof. Dr. Khalid Mahmood Iraqi, Vice–Chancellor, University of Karachi for providing working facilities for the completion of this work and also to Messrs M. Rafiquddin (Karachi), Abrar Ali (Karachi), M. Y. Saleem (Islamabad) and S. Hameed (Islamabad) for executing the illustrations and to Dr. Jan Alam (Hazara University) and Nazar Khan (Zhob) for providing coloured plates.


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Annual, biennial or perennial herbs, sometimes shrubs, rarely trees, with or without milky latex, sometimes armed with spines, usually terrestrial, rarely aquatic herbs. 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. Capitula usually solitary or variously arranged in corymbose or paniculate synflorescences. 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, 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 having only disc florets or ligulate bearing only ligulate florets. Heterogamous capitula are radiate having disc florets in the centre and peripheral ray florets or radiant bearing disc florets in the centre and peripheral florets with ± dilated, radical to strongly bilateral corolla or disciform bearing perfect or functionally male disc florets in the centre and peripheral filiform florets. Florets are epigynous, bisexual, female or male (at least functionally) or sterile (neuter). Calyx often represented by a pappus of 1 or more series of bristles, scales or awns on the apex of the ovary or sometimes pappus completely absent. Corolla gamopetalous, tubular, fililform, 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 bifurcation or not. Fruit unicellular indehiscent, cypselas (achene) with or without ribs, sometimes with distinct carpopodium (basal attachment area), very rarely a drupe.

Asteraceae, the second largest family of Angiosperms after Orchidaceae in terms of number of species but largest in number of genera, comprises of ca 1600 genera and ca 24000 species, present almost every where with the exception of Antartica.

Asteraceae is also the largest family in Pakistan, represented by 188 genera and 770 species including some commonly cultivated taxa also. Volume 224 (Asteraceae – VI) of the Flora of Pakistan is the last volume/ part of the family In the first volume of Asteraceae (I) – Anthemideae, a infra-familial key pertaining to the tribes present in Pakistan was provided. The present volume VI is now being published almost after 18 years, since then lot of changes have taken place particularly based on molecular and phylogenetic studies resulting into reshuffling of number of genera from one tribe to another. The circumscription of quite a few tribes is greatly changed. Some old and forgotten tribes and genera are resurrected even some new tribes are recognized/ described. New genera are being described continuously and many polyphyletic genera are splitted into a number of satellite and smaller genera. Therefore, it seemed necessary to provide a new key for the tribes of Asteraceae occurring in Pakistan. In all 19 tribes in our area are recognized. There is a controversy in the recognition of subfamilies for instance Bremer (Asteraceae: Cladistics and Classification, 1994) recognized 3 subfamilies and 16 subtribes; Panero & Funk (Proc. Biol. Soc. Washington 115: 900-922. 2002) recognized 10 subfamilies whereas Kadereit & Jeffery (The Families and Genera of Flowering Plants Vol. VII. 2007) recognized 5 subfamilies. Not going in the details of the classification, we have followed Jeffrey in Kadereit & Jeffrey, l. c. in general outline and arrangement of tribes.

Key to the tribes, treated in Flora of Pakistan along with the synopsis of the genera are provided. Genera within each tribe are alphabetically arranged. Volume number against each genus is also given in parenthesis.


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1 All florets ligulate (strap–shaped), equally 5–toothed at the apex. Plants with milky latex. Cichorieae
+ At least the central florets tubular, not ligulate (not strap–shaped) if radiate then with 4 or fewer equal or unequal apical teeth. Plants without milky latex (except in Arctotideae) or with watery sap or not. (2)
2 (1) Plants thistle – like. Leaves mostly spiny or spinulose. Phyllaries spiny-tipped. (3)
+ Plants not thistle like. Leaves mostly unarmed, very rarely spiny. Phyllaries not spiny tipped. (4)
3 (2) Capitula heterogamous, rarely homogamous. Anthers ecaudate rarely shortly tailed. Style thickened apically, narrowing near the middle and slender to base, usually with longer style hairs in a ring below bifurcation. Nectary absent. Receptacle epaleate, deeply foveolate. Arctotideae
+ Capitula homogamous. Anthers distinctly caudate, rarely shortly tailed. Style neither thick nor narrowing near the middle, usually with a papillose – pilose thickening below the branches. Nectary present below the style. Receptacle paleate with long bristles, rarely epaleate. Cardueae
4 (2) Cypselas carbonized, black or brownish black, with a phytomelanin layer in the pericarp. (5)
+ Cypselas not carbonized, variously coloured but not black, without a phytomelanin layer in the pericarp. (12)
5 (4) Disc floret corollas with conspicuous reddish or orange resin canals. Neurolaeneae
+ Disc floret corollas without conspicuous reddish or orange resin canals. (6)
6 (5) Corolla of disc florets with twin glandular trichomes. Cypselas ribbed or with thickened striations or with ciliate lateral ridges and glandular trichomes. Pappus a crown formed from apically arcuate, twin trichomes or with recurved apices. Athroismeae
+ Corolla of disc florets glabrous or if pubescent then without twin glandular trichomes. Cypselas unribbed or without thickened striations or laterally ciliate ridges. Pappus not as above or absent. (7)
7 (6) Plants caulirosulate or megaphytic rosulate. Leaves sometimes with expanded leaf bases (cupule), lamina linear – ovate or suborbicular, unlobed. Ligule of ray florets sometimes extremely short or lacking, apices conspicuously trilobed, rarely shallowly trilobed. Millerieae
+ Plants neither caulirosulate nor megaphytic rosulate. Leaves without expanded leaf bases (cupule). Ligule of ray florets not as above. (8)
8 (7) Capitula always discoid. Style usually conspicuous and strongly exerted from corollas. Style arm appendages longer than stigmatic lines. Pappus composed of bristles, never of barbed awns. Florets never truly yellow. Eupatorieae
+ Capitula radiate, discoid or disciform. Style not strongly exerted from corolla. Style arms appendages usually as long as or shorter than the stigmatic lines. Pappus scaly or of few barbed awns (rarely bristly in Tageteae and Millerieae but remaining characters otherwise). Florets variously colored, including yellow. (9)
9 (8) Phyllaries dimorphic, outer ones herbaceous, inner ones membranous or wholly scarious. Coreopsideae
+ Phyllaries not dimorphic, outer and inner phyllaries mostly membranous (10)
10 (9) Receptacles epaleate, rarely with few scales or bristles. Plants aromatic. Leaves and / or phyllaries with pellucid glands or pustules, if without glands or pustules then cypselas strongly 9 – 10 (–12) ribbed. Tageteae
+ Receptacles paleate, paleae partially or strongly conduplicate. Plant not aromatic. Leaves and / or phyllaries without pellucid glands or pustules. Cypselas either unribbed or faintly few ribbed (less than 9). (11)
11 (10) Style arms usually with a tuft of papillae or hairs at distal end. Ray and disc cypselas similar, compressed, rarely obcompressed, biconvex, narrowly obovate to suborbicular in outline. Pappus of awns or squamellae or both, sometimes absent, borne on cypsela neck. Heliantheae
+ Style arms usually glabrous. Ray and disc cypselas different. Disc cypselas obconic to obpyramidal, sometimes incurved, quadrate to round in outline. Pappus of bristles or awns or both, not borne on cypsela neck. Millerieae
12 (4) Phyllaries with a thickened cartilagi-nous, basal portion (stereome), generally brightly coloured or hyaline, greenish or greenish brown. Style arms apices conical. Gnaphalieae
+ Phyllaries without thickened cartila-ginous basal portion. Style arms apices acute, acuminate, globose, rarely conical. (13)
13 (12) Most of the florets zygomorphic. Marginal florets usually bilabiate with an expanded limb, rarely without adaxial lobes. Corolla of disc florets bilabiate with 2 adaxial lobes and 3 abaxial lobes. Mutisieae
+ Florets mostly actinomorphic. Marginal florets mostly not bilabiate. Corolla of disc florets not bilabiate. (14)
14 (13) Indumentum of T – shaped or stellate hairs, sometimes mixed with simple or glandular hairs. (15)
+ Indumentum of various types but not of T–shaped or stellate hairs or rarely glabrous. (16)
15 (14) Anthers mostly rounded, rarely shortly tailed at the base, with an ovate, triangular to subtriangular appendage. Style with a slender or a bulbous base usually situated on a cup – shaped, stylopodium (nectary). Tectum not lophate. At least central disc florets 3 – 6 – lobed. Anthemideae
+ Anthers mostly long tailed, with a flat, thin apical appendage. Style not as above. Tectum usually lophate. Disc florets 4 – lobed. Vernonieae
16 (14) Style branches often with long hairs in a ring, well below the bifurcation (arctotoid). Leaves and phyllaries usually spinulose or spiny tipped. Latex rarely present. Lobes of disc florets generally with continuous veins. Arctotideae
+ Style branches not as above. Leaves and phyllaries neither spinulose nor spiny tipped. Latex absent. Lobes of disc florets without continuous veins. (17)
17 (16) Phyllaries cartaligineous. (18)
+ Phyllaries herbaceous or chartaceous. (19)
18 (17) Anthers ecaudate. Ectexine of pollen 2 – layered, outer columellate layer and an inner irregularly interlaced basal layer (Gnaphalioid type). Hairs on style branches not sweeping, stigmatic rows generally separated. Gnaphalieae
+ Anthers caudate basally with long or short and branched or unbranched tail, rarely ecaudate. Ectexine not as above. Style branches more often with acute or obtuse sweeping hairs, stigmatic rows apically confluent, basally separated. Inuleae
19 (17) Anthers appendages several times as long as wide, usually acuminate or apiculate. Style base, arms and shaft glabrous. Mutisieae
+ Anthers appendages not as above. Style arm and shaft with sweeping hairs or at least papillate at the apex or a tuft of papillae or sweeping hairs. (20)
20 (19) Style arm of ray florets supinate, of disc florets pronate at maturity. Style asteroid type i.e. stigmatic portion prolonged into triangular to lanceolate to subulate sterile appendages. Astereae
+ Style arm of ray florets and disc florets not as above. Stigmatic portion not prolonged into triangular to lanceolate to subulate sterile appendages. (21)
21 (20) Phyllaries usually prolonged into a membranous spiny or unarmed appendage, inner ones whitish to pinkish– violet to purple or with a long pink or purple apex. Anthers ending apically into a rigid, lignified, lanceolate appendage. Style with a papillose–pilose thickening (functioning as a pollen brush) below the branches, nectary usually present at the base of style. Stamens strongly thigmotropic. Cardueae
+ Phyllaries not prolonged into an appendage, inner ones otherwise. Anthers not ending into a rigid, lignified appendage. Style not as above, nectary absent at the base of style. Stamens not or slightly thigmotropic. (22)
22 (21) Filament collar either dilated with swollen cells or the collar slender, hardly wider than basal part of the filament. (23)
+ Filament collar not as above. (24)
23 (22) Phyllaries usually 1 – seriate rarely 2 – seriate. Style not situated on a cup – shaped stylopodium (nectary). Anthers appendages flat, basally obtuse, sagittate or caudate. Senecioneae
+ Phyllaries 2 – 7 – seriate. Style with a slender or bulbous base usually, situated on a cup – shaped stylopodium (nectary). Anthers appendages ovate, triangular or subtrianglar to subulate, basally rounded. Anthemideae
24 (22) Cypselas polymorphic rarely homomorphic within the same capitulum, outer corymbiform, somewhat curved, rostrate or triquetrous; inner mostly curved, sometimes winged or sometimes with a fleshy exocarp (drupaceous). Pappus absent. Calenduleae
+ Cypselas homomorphic, rarely dimorphic, not curved, or fruit never drupaceous. Pappus present, of various forms of few to many bristles, scales, awns or squamellae or very rarely absent. (25)
25 (24) Cypselas ribbed or distinctly striated or ridged. (26)
+ Cypselas neilther ribbed nor distinctly striated or ridged. (28)
26 (25) Disc florets with twin glandular trichomes. Pappus of fertile floret cypselas a crown of twin trichomes, sometimes with recurved apices or lacerate fused squamellae. Anthers usually ecaudate or rarely shortly caudate. Athroismeae
+ Disc florets without twin glandular trichomes. Pappus without a crown of twin trichomes, usually of scales or bristles. Anthers sagittate or caudate. (27)
27 (26) Phyllariles uniseriate, rarely 2–seriate. Ray florets not conspicuously 3–5 lobed. Anthers appendages flat. Senecioneae
+ Phyllaries in 2-several series. Ray florets conspicuously 3–5–lobed. Anthers appendages narrowly ovate to round, strongly carinate. Helenieae
28 (25) Anthers appendages present. Style arms with two parallel stigmatic surfaces. Ray florets prominently radiate, ligule mostly attenuate to oblique into tube, rarely obtuse to truncate. Helenieae
+ Anthers appendages absent. Style arms with two apically confluent and basally separated stigmatic surface. Ray florets minutely radiate, tubular or filiform. Inuleae

Lower Taxa
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