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Ranunculaceae
HARALD RIEDL and YASIN J. NASIR
Naturhistorisches Museum, Botanische Abteilung, Wien, Austria. and National Herbarium, Pakistan Agricultural Research Council, Islamabad.
Annual or perennial herbs, rarely woody climbers. Leaves undivided or more or less deeply lobed, pinnatisect to pinnatipartite, rarely pedatisect, alternate, rarely opposite. Uppermost leaves sometimes united into an involucre that closely resembles a calyx. Flowers single or united to various types of racemose inflorescences, usually radially symmetric, zygomorphic in Aconitum, Delphinium and Consolida, with a single perigonium either formed by petaloid sepals or true petals corresponding morphologically to nectaries or composed of calyx and corolla. Sepals and petals (nectaries) (2-)3-5(-8), rarely more or 0, sometimes provided with a spur. Intermediates between uppermost leaves and sepals as well as between stamens and nectaries or petals frequently present. Stamens usually numerous, reduced in number to 2, in several whorls or rarely in a single whorl. Carpels 1 to numerous, in the latter case arranged spirally on a convex receptacle. Fruit a follicle, often dry, 2 to many-seeded, with marginal ovules or one-seeded nutlets with a single basal ovule (achenes), very rarely baccate (Aclaea).
The number of genera varies greatly from author to author (35-70) according to wider or narrower generic concepts, as convincing criteria are often lacking. It may be about 50. Approximately 2000 species are present, widely distributed throughout the northern hemisphere, but also in southern temperate regions, in the tropics wh ere they are usually confined to higher altitudes. With a few exceptions found in mesic to humid habitats. Represented in Pakistan by 22 genera and c. 114 species, of which several genera l ike Nigella, Anemone, Aquilegia include plants of ornamental value, others are toxic and used for medicinal purposes. Some of the annual species have become agricultural weeds, and their present day distribution is strongly influenced by man’s activities.
Morphologically the family displays an unusually wide diversity of features in practically every part. Systematically it is fairly close to Magnoliaceae. Number of floral parts are not constant and are rather high in several members but reduced to 1-2 within the family. Functional diversity however, seems to correspond to morphological diversity. Nearly all kinds of seed dispersal known among angiosperms is found, but dispersal by wind, water and by external attachment to animals is most common. The genera Aconitum, Anemone and Delphinium revised by Yasin J. Nasir Acknowledgements: We are grateful to the Director/Curators of the following herbaria for the loan of plant specimens: E, K, KUH, PPFI-B, W. The financial assistance received from the U.S. Department of Agriculture under PL-480 is gratefully acknowledged.
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1 |
Flowers zygomorphic |
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(2) |
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Flowers regular (actinomorphic) |
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(4) |
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2 (1) |
Flowers without a spur. Upper sepal hood-l ike |
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9 Aconitum |
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Upper sepal with a distinct spur |
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(3) |
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3 (2) |
Perennials. Follicles 2-5 |
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10 Delphinium |
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Annuals. Follicles single |
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11 Consolida |
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4 (1) |
Sepals and petals 5, each with a distinct spur, petaloid |
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5 Aquilegia |
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Sepals and petals without spur |
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(5) |
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5 (4) |
Fruit a berry. Flowers small, stamens longer than corolla. Leaves compound |
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2 Actaea |
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Fruit not a berry, dry many-seeded follicles or one-seeded nutlets |
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(6) |
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6 (5) |
Fruit 2-many seeded follicles |
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(7) |
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Fruit 1-seeded achene |
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(12) |
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7 (6) |
Stamens longer than corolla. Numerous small flowers arranged in long, simple or branched racemes |
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1 Cimicifuga |
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Stamens shorter than corolla. Flowers single or arranged in a different kind of inflorescence |
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(8) |
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8 (7) |
Leaves undivided, roundish, dentate to crenate |
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6 Caltha |
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Leaves divided in various ways |
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(9) |
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9 (8) |
Sepals blue to whitish with blue veins, petaloid. Leaf segments filiform |
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8 Nigella |
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Sepals usually white or yellow, rarely lilac. Leaf segments wider |
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(10) |
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10 (9) |
Sepals petaloid yellow to orange, in several circles, conchate |
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7 Trollius |
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Sepals petaloid or lacking, white or rarely lilac |
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(11) |
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11 (10) |
Stem without leaves. Growth-habit distinctly caespitose |
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4 Paraquilegia |
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Stem with few leaves. Growth-habit not caespitose |
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3 Isopyrum |
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12 (6) |
Woody or herbaceous climbers, rarely smaller, erect plants with large, single flowers. Flowers with a simple perianth, strictly tetramerous. Single to a few or numerous flowers in a panicle-l ike inflorescence |
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14 Clematis |
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Neither woody nor climbers. Flowers tetramerous only in Thalictrum |
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(13) |
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13 (12) |
Flowers with a simple perianth |
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(14) |
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Perianth double, composed of calyx and corolla |
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(16) |
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14 (13) |
Achenes with a long plumose style |
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13 Pulsatilla |
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Style short, not plumose |
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(15) |
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15 (14) |
Stem with several ± equally distributed leaves. Inflorescence a simple raceme or a branched panicle |
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15 Thalictrum |
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Stem with only one circle of free or connate leaves forming a kind of involucres. Flowers 1-2 or in an umbel-l ike inflorescence |
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12 Anemone |
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16 (13) |
Floating aquatic herbs, the submerged leaves with filiform leaf-segments |
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20 Batrachium |
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Non floating aquatic herbs but sometimes with flaccid, creeping stems on wet ground |
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(17) |
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17 (16) |
Calyx persistent in the mature fruit |
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18 Oxygraphis |
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Calyx deciduous after flowering |
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(18) |
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18 (17) |
Small annuals. Petals yellowish, shorter or slightly longer than the calyx. Stamens 5-15 |
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22 Ceratocephala |
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Annual or perennials. Petals usually manifestly longer than calyx. Stamens more than 15 |
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(19) |
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19 (18) |
Carpels rugose with prominent veins at least when young (regular and without veins, but with a transverse, prominent, lobed ring at maturity in some species of Adonis). Flowers yellow, red or shade of pink |
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(20) |
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Carpels without prominent veins. Flowers yellow |
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21 Ranunculus |
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20 (19) |
Stems rooting at the nodes, flaccid. Flowers on short leafless stalks, yellow, small. Veins of carpels ± parallel, scarcely branched. Petals 5 |
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19 Halerpestes |
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Stems erect, not flaccid or rooting at the nodes. Flowers terminal. Veins of carpels reticulate. Petals often more than 5 |
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(21) |
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21 (20) |
Flowers large, white or pink. Leaf-segments ovate |
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17 Callianthemum |
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Flowers yellow or red. Leaf-segments narrowly linear |
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16 Adonis |
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List of lower taxa
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