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Pakistan | Family List | Liliaceae

3. Gagea Salisb. in Köenig & Sims in Ann. Bot. 2: 555. 1806. Boiss., Fl. Or. 5: 203. 1882; Hook. f., Fl. Brit. Ind. 6: 355. 1892; E. M. Rix in P. H. Davis, Fl. Turk. 8: 312. 1984; S. Dasgupta & Deb in J. Bomb. Nat. Hist. Soc. 83(1): 78. 1986; Wendelbo & Rech. f. in Rech. f., Fl. Iran. 165: 13. 1990; M. N. Tamura in Kubitzki, Fam. Gen. Vasc. Pl. 3: 351. 1998; Xinqi & Turland in Wu Zhengyi & P. H. Raven, Fl. China 24: 117. 2000.

S. I. Ali and I.G. Levichev

Department of Botany, University of Karachi, Karachi-75270 Pakistan

Herbarium, Komarov Botanical Institute of the Russian Academy of Sciences, Saint Petersburg

Gagea punjabica

Credit: I.G. Levichev

Small bulbous plants. Roots short lived, 2-4 (-7) months; two types, first ageotropic thick sclerified roots, generally followed by thin geotropic feeding roots. Bulb consists of one fleshy tunicate basal scale. Sometimes bulbils present inside or outside the bulbar tunic or in axils of leaves on the peduncle. Basal leaves two, more often one, very rarely none. Leaves on the peduncle generally present. Inflorescence branched, quite often umbellate, sometimes uniflorus. Bracts often present. Tepals free, 3+3, frequently yellow inside, more or less green, less often with a purple shade outside. Stamens free, unequal, filaments filiform or thickened at the base. Anthers oblong or linear, yellow, very seldom purple. Ovary superior, sessile or stipitate; style distinct, trilobate to trifid. Fruit a loculicidal capsule. Seeds small terete, faceted or lobule-form or flattened, with more or less expressed aril

A genus with about 250 species (Levichev, Komarovia 1: 45-57. 1999) distributed in temperate regions of N. Africa, Asia and Europe, mainly in Central Asia and Mediterranean regions; represented in Pakistan by 36 species.

1 Inflorescence umbellate: leaves and pedicels arise from distal end of the peduncle, forming a whorl, or inflorescence nearly umbellate; lower subinflorescence leaf is, in old and large plants, situated somewhat below the whorl or 1-2 pedicels are located slightly above the whorl, and several smaller leaves develop on pedicels   (2)
+ Inflorescence branched; leaves located by turns, only in immature or young specimens (uni-, biflorous) the inflorescence may seem almost umbellate   (14)
2 (1) Peduncle quadrangular in T.S. Basal leaf always single, linear, from above angularly-grooved, from below carinate. Inflorescence 2-3(-5) flowered, at base with 2 opposite leaves, lower leaf narrowly lanceolate, as long or slightly longer than the perianth. Vegetative reproduction in generative plants is absent. Seeds terete   13 Gagea bashoensis
+ Peduncle circular in T.S., smooth or more or less ribbed   (3)
3 (2) Peduncle longitudinally ribbed. Plants xeromorphic, rigid due to collenchyma developed in the peduncle and in leaf angles. Tunic of bulbs fiber-l ike, forming short fibrous collar   (4)
+ Peduncle smooth, cylindrical, plants mesomorphic, elastic, soft   (12)
4 (3) Basal leaf single, narrowly-linear, ? pentahedral in T.S. Peduncle well developed, rising above soil surface. Tepals extended into a long tip   (5)
+ Basal leaves grass-l ike, usually 2 (in mature specimens 1), wider than thick, in T.S. triangular-grooved. Peduncle short, rising slightly above the soil surface or often hidden in soil. Tepals apiculate   (9)
5 (4) Juvenile and immature plants develop a long stolon (up to 4-5 cm), with a single vegetative bulbil at the end. Simultaneously there may also be a short stolon for further deep pushing of the replacing bulb. Bulb tunics thin, papery, light or faintly brownish. Roots numerous, hair-l ike, not braiding bulb. Basal leaf usually arcuately bent   18 Gagea quettica
+ Vegetative bulbil sessile and formed in juvenile period only. Incrassate sclerified roots also develop at early stages of an ontogeny, may be present in older plants also   (6)
6 (5) All parts of plants very thin. Basal leaves less than 1 mm wide. Pedicels of different lengths, may be up to 3 times or more surpassing the tepal length   (7)
+ Plants not very thin, breadth of basal leaf up to 1 mm, the peduncle is slightly wider. Pedicels shorter or slightly longer than the tepals   (8)
7 (6) Pedicels widely directed in different directions. Base of lower subinflorescence leaf expands to 1.5 mm. Tepals up to 12 mm long, apiculate   22 Gagea wallichii
+ Pedicels forming a very narrow, vertical bunch. Base of lower subinflorescence leaf expands to 3 mm. Tepal length up to 18 mm   20 Gagea punjabica
8 (6) Basal leaf arcuate in upper half. Lower subinflorescence leaf as long as the inflorescence. Capsule wide, oviform, largest on longest pedicel (of first flower), subsequent capsules - up to twice as small   19 Gagea baluchistanica
+ Basal leaf directed upwards. Lower subinflorescence leaf longer than inflorescence. Capsule narrowly-lanceolate   21 Gagea setifolia
9 (4) Plants solitary, bearing a few flowers, inflorescence base hidden below the soil surface. Leaves curved. Pedicels of flower buds almost absent, in fruiting plants equal to tepals. Anthers yellow   (10)
+ Plants in dense stands, bearing numerous flowers, inflorescence base not hidden below the soil surface. Pedicels of flower buds short, in fruiting 2-3 times longer than tepals. Anthers always or temporarily brown-violet tinged   (11)
10 (9) Inflorescence 2-4-flowered or flowers solitary. Bulb with thick and dense sclerified roots. Leaves bent in different directions. Tepals narrowly lanceolate with a long tip   16 Gagea conjungens
+ Inflorescence 2-4- flowered. Bulb with poorly weaved thin sclerified roots. Leaves arcuate. Vegetative reproduction, l ikely by stolon. (In immature stages the replacing bulb forms a stolon for further deep pushing)   17 Gagea mergalahensis
11 (9) Inflorescence 3-6-flowered. Basal leaf 0.6-0.8mm broad. Lower subinflorescence leaf equal to or longer than inflorescence, 1.5 mm broad, shortly lanceolate, linear elongated. Anthers dark-brown, tepals pale yellow inside, greenish outside with narrow yellowish margin   15 Gagea rawalpindica
+ Inflorescence 3-20(-30) - flowered, forming a narrow bunch. Basal leaf 1.5-2 mm wide. Lower subinflorescence leaf sometimes located at the neck of the bulb, usually accreting to a short peduncle, at the base up to 3.5 mm broad, very gradually narrowed to tip, frequently with a vegetative bulbil in its axil. Anthers in flower buds completely or along the margin violet, at anthesis yellow, tepals yellow within white-pale-bordered   14 Gagea pakistanica
12 (3) Plants multiply vegetatively in all ontogenetic stages, group of vegetative bulbils develops laterally on parent bulb in juvenile plants, in immature plants (in form and colour looks l ike a compound fruit of blackberry) it develops on a short peduncle, wrapped up by a wide subinflorescence leaf and born on soil surface, generative plants with a single basal bulbil. Two basal leaves present at the beginning of generative stages, later only 1 basal leaf is present. Pedicels elongated, 2-4 times longer than tepals. Tepals obtuse, often sinuate or rounded. Seeds terete   25 Gagea liotardii
+ Vegetative reproduction absent. Basal leaf always 1, inflorescence 1-2(3)-flowered. Pedicels shorter than perianth. Seeds thin-flat   (13)
13 (12) Flowers broadly bell-shaped, tepals broadly elliptic, rounded at the tip (at the end of flowering their tips roll-up and seem pointed). Inflorescence with 2(-3) leaves at the base. Bulb 10-15 mm in diameter, bearing numerous thin roots   23 Gagea aitchisoniana
+ Flowers narrowly bell-shaped, tepals elongated narrowly-lanceolate, apiculate. Inflorescence with 4-5 leaves at the base. Bulb 8-10 mm in diameter. Anthers orange tinged   24 Gagea alexii
14 (1) Basal leaves more or less wide, lanceolate or lan ceolate-l inear, flat or faintly concave, always 1 (second leaf is reduced)   (15)
+ Basal leaves filiform or linear, concave, angularly grooved, or cylindrical, 2, 1 or absent   (18)
15 (14) Basal leaf smooth. Inflorescence branched, lower internodes longer than pedicels. Vegetative bulbils light brown, densely compressed, forming a spherical group situated laterally to parent bulb. Tepals up to 8 mm long, tip rounded. Capsule narrow, prismatic, apically convex   9 Gagea dschungarica
+ Basal leaf longitudinally furrowed. Inflorescence shortly branched, lower internodes shorter than pedicles. Vegetative bulbils black-brown, surrounding about half the perimeter of parent bulb. Tepals up to 16 mm long. Capsule three-rayed, height equal to width, apically faintly widely-concave   (16)
16 (15) Lower subinflorescence leaf is much broader than the basal leaf, the lower subinflorescence leaf is longer than the inflorescence   11 Gagea davlianidzeae
+ Basal and lower subinflorescence leaves have the same width   (17)
17 (16) Basal leaf and lower subinflorescence leaf broadly lanceolate (up to 13-15 mm wide). Tunic of bulb light brown. Bulbils colaterally surrounding parent bulb, mainly semi-dr op-shaped (sometimes fa ceted), black-brown. Tepals widely oblanceolate, blunt   10 Gagea lowariensis
+ Basal leaf and lower subinflorescence leaf narrowly lanceolate (up to 3-5 mm wide). Tunic brown-black. Bulbils a little pulled together, forming a compact group surrounding the basal bulbil, usually polyhedral (group also can be named colaterally located). Tepals elongated, lanceolate, apic ulate   12 Gagea elegans
18 (14) Subinflorescence leaf with widened lanceolate base, twice or more wider than the basal leaf. Basal leaves 1 or 2 (always 2, or 2 only in immature plants and at onset of generative stages, in mature plants 1, second basal leaf reduced). Seeds faceted, lobule-form, thick, flat   (19)
+ Subinflorescence leaf linear, not expanded at the base, gradually narrowed to the tip, as broad as the basal leaf or slightly broader. Basal leaves grooved, 1 or 2 (always 2, or 2 only in immature plants and at the onset of generative stages, and in mature plants 1, second basal leaf accreted to peduncle). Seeds flat (thin-flat). Tunic more or less elongated forming a short neck   (29)
19 (18) Basal leaves 2 (in older plants 1) filiform, grooved abaxially, carinate. In immature plants on the surface of the soil, in the axils of twisted hair-l ike leaves, the capitate group of bulbils is present at the end of the peduncle. In generative plants only a semi-teardro p shaped basal vegetative bulbil is present   26 Gagea tenera
+ Basal leaves cylindrical, not carinate abaxially, slightly concave adaxially, usually 1, rarely 2 or absent, in that case a tiny rudiment is preserved on top of vegetative bulbil. Seeds faceted, lobuleform, flattened (thickly)   (20)
20 (19) Basal leaf absent or reduced to a more or less rudiment. Small group of vegetative bulbils develop in axils of all leaves of the inflorescence   36 Gagea gageoides
+ Basal leaves (1 or 2) always present   (21)
21 (20) Basal leaves always 2, filiform. Vegetative bulbils in dense group, blackish, basal narrowly cochleariform, sister bulbils hypobasal lobuliform or semilunar   33 Gagea spumosa
+ Basal leaf always 1   (22)
22 (21) Delicate, tiny plants (5-10 cm high), more or less forming groups. Leaves hair-l ike, bent   (23)
+ Larger plants (10-20 cm high), usually solitary   (24)
23 (22) Plants usually not in dense groups. Vegetative bulbils in a compact group at the base of peduncle, very small (basal bulbil not differentiating) fine-cellular. Flowers golden yellow inside, greenish outside   34 Gagea bowes-lyonii
+ Plants in dense groups which include numerous juvenile plants. Single, rather large, vaguely finely cellular basal bulbil develops at the base of parent bulb. Group of elongated fine bulbils formed on truncate peduncle of immature plants. Basal and lower inflorescence leaves shorter than the inflorescence. Tepals and capsule narrowed towards the base. Flowers white inside, greenish outside   35 Gagea delicatula
24 (22) Internal tunics light, pale-brownish, contrasting with the colour of vegetative bulbil, which is single, almost black, wrinkled outside   28 Gagea leosii
+ Colour of tunicated vegetative bulbils the same or slightly different (colouring from brown to black)   (25)
25 (24) Bulbils fine-cellular, with top attachment (a small depression is preserved in the narrow upper part of the bulbil)   32 Gagea rubinae
+ Vegetative bulbils with lateral vertical light strip or bottom attachment (forming a small depression at the base of expanded part of bulbil)   (26)
26 (25) Basal leaf fistular, 2-2.5 mm in diameter. Tepals up to 4 mm wide, outer almost rhombic, shorter than the inner, inner obovate, rounded at the top. Tunic dark-brown. Group of brown cellular vegetative bulbils develops at the base of the peduncle, largest bulbil is basal, semi-dro p-shaped or faceted with a vertical strip of attachment, sister bulbils are ± numerous attached at the top   27 Gagea absurda
+ Basal leaf not fistular in middle part, about 1 mm in diameter   (27)
27 (26) Tepals white, narrowly-lanceolate, 6-8 mm long. Tunics dark brown with short neck (5-10 mm). Vegetative bulbils very small (0.6 ? 0.9 mm), in dense group, faceted, very finely cellular, are attached to the parent plant at the base, surrounding basal bulbil of the similar size. Lower subinflorescence leaf from narrowly-lanceolate base (2 mm) gradually narrowed to long cylindrical tip. Peduncle 3-5 cm long. Capsule not narrowed to stipe   29 Gagea kunawurensis
+ Tepals yellow. Plants >10 cm in height, peduncle >5 cm   (28)
28 (27) Tunica brown, continued into a neck, about 1.5 cm long. Basal leaf c. 1 mm wide. Vegetative bulbils in groups, small, trihedral-spherical, fine-meshed, the basal bulbil because of accretion with the base of peduncle, is separated with a long fragment of its superficial tissues. Tepals 7-8 mm long, golden yellow inside, greenish outside   30 Gagea jispensis
+ Tunica brown-black, does not form a neck. Basal leaf 1 mm wide or slightly wider. Attachment of vegetative bulbils (both large basal and smaller sister bulbils) forms a light vertical strip, remaining surface composed of large cells. Tepals up to 11 mm long, white or pale yellow, or yellow. Capsule narrowed into a stipe   31 Gagea ova
29 (18) Peduncle not branched, terminated by a single flower. Numerous leaves get reduced in size from base upwards, bulbils develop in their axils   8 Gagea jaeschkei
+ Inflorescence 1-2 - multi- flowered, without bulbils or bulbils present only in the axils of several lower leaves   (30)
30 (29) Only single bulbils present in the axils of basal leaf and several lower inflorescence leaves. Pedicels forming a narrow bunch, very unequal in length, elongating at anthesis, having the same length in the fruiting plants   (31)
+ Bulbils not present on the peduncle   (32)
31 (30) Pedicels up to 6 cm long. Capsule narrowly oval, up to 3.5 mm wide, about half the length of tepals   6 Gagea drummondii
+ Pedicels up to 4 cm long. Capsule of the first flower largest, widely-oval (10 ? 15 mm), almost reaching the length of tepals   7 Gagea utriculosa
32 (30) Basal leaf single   (33)
+ Basal leaves two   (35)
33 (32) Inflorescence 6-14 flowered. Tunic and bulbils brown. (In generative plants, single vegetative bulbil forms thick sclerified roots)   1 Gagea alii
+ Inflorescence 1-3-flowered. Tunic dark grey   (34)
34 (33) Inflorescence branched, 1-3-(-5)-flowered, pedicels during flowering get elongated. Generative plants do not reproduce vegetatively, only in juvenile stage a group of dark grey longitudinally wrinkled bulbils are formed   2 Gagea vvedenskyi
+ Inflorescence usually uniflorous, very rarely 2-3-flowered. In juvenile plants one flattened brown-black longitudinally cellular bulbil is formed   5 Gagea uliginosa
35 (32) Plants up to 2-4 cm high, forming individual groups of different ages. Pedicels equal to tepals or slightly longer   3 Gagea chitralensis
+ Plants up to 8-10 cm high. Pedicels 3-5 times as long as tepals   4 Gagea grey-wilsonii

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