Published In:
Genera Plantarum 117–118. 1789. (4 Aug 1789) ( Gen. Pl.)
(Last Modified On 6/9/2015)
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Acceptance
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Synonym
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(Last Modified On 6/16/2015)
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Contributor Text:
*R.R. MILL, **M. QAISER, T. SIDDIQUI, G. R. SARWAR, S. I. ALI, S. KHATOON, SULTANUL ABEDIN, HAMIDULLAH, ***S. A. GHAZANFAR
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Contributor Institution:
*Royal Botanic Garden, Edinburgh, ** University of Karachi, Karachi-75270, Pakistan, *** KEW
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General/Distribution:
Scrophulariaceae s. l. is represented in Pakistan by 37 genera and 187 species.
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Comment/Acknowledgements:
We are grateful to the Directors/ Curators of the following herbaria for herbarium and library facilities: A, AR, BM, CAL, DD, E, GH, K, KUH, MICH, NA, NY, O, PH, RAW, TI and TZ. We thankfully acknowledge financial support for this publication from Pakistan-U.S. Science and Technology Cooperation Program. We are grateful to Prof. Steven Blackmore, former Director, Royal Botanic Garden, Edinburgh for permitting Mr. R. Mill to generously contribute to Flora of Pakistan. We are thankful to Dr. D. Albach (Gutenburgh Universität, Meinz) for kindly reviewing the account of the genus Veronica. Grateful thanks are due to Professor Anjum Parveen, Director, Centre for Plant Conservation, University of Karachi for providing working facilities to the project and for understanding and encouragement. It is a matter of pleasure for us to thank Dr. Shahina A. Ghazanfar for kindly helping us from time to time.
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Herbs, subshrubs, shrubs or rarely small trees, autotrophic rarely hemiparasitic. Leaves exstipulate, alternate, opposite or whorled. Inflorescence racemes, spike, solitary axillary or thyrsoid panicles. Flowers bisexual, usually zygomorphic bilabiate, rarely actinomorphic. Calyx (2)-4 or 5 lobed or parted. Corolla gamopetalous zygomorphic and often bilabiate or actinomorphic sometimes with a spur or pouch at base. Stamens epipetalous, mostly 4, didynamous or 2 rarely 5; sometimes 1 or 2 staminodes are present; anther locules 1 or 2 equal or subequal, free or confluent. Nectary often present at the base of the ovary, ringlike, copular or reduced to a gland. Ovary superior, bilocular, axile placentation, ovules numerous or sometimes few, style simple, stigma capitate. Fruit a capsule. Seeds numerous, variously ornamented. The family is being considered here in a broader sense. However, the molecular systematic studies (Angiosperm Phylogeny Group 2009*) have amply demonstrated that the family is grossly polyphyletic. Many genera have been shifted to Orobanchaceae, Phyrmaceae, Plantaginaceae, Linderniaceae and Paulowniaceae. As we have already published the accounts of Orobanchaceae (1976), Phyrmaceae (1973) and Plantaginaceae (1974) according to the formally accepted circumscription it seems best to treat the family Scrophulariaceae sensu lato. *Angiosperm Phylogeny Group 2009. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Bot. J. Linn. Soc. 161: 105-121. Excluded Genera 1.Anarrhinum Desf., Fl. Atl. 2: 51. 1798. Anarrhinum orientale Benth. in DC., Prodr. 10: 289. 1846. According to R. R. Stewart (l. c. 645) Blatter reported the above mentioned taxon from Harboi, Baluchistan, 9000 ft on the authority of Hotson, who collected in Persian Baluchistan and the Makran Coast in 1916-18 and his plants were named by Blatter. This taxon has never been collected since then. D. A. Sutton (Rev. tribe Antirrh. 258. 1952) lectotypified Anarrhinum orientaleBenth. by the Ethiopian material. 2.Kashmiria D. Y. Hong in Bot. Notiser 133 (4): 565. 1980. Falconeria himalaika Hook. f. in Hook. Icon. Plant. 16: 30, pl. 1438. 1883 non Falconeria Royle (1839). Though it is named after ‘Kashmir’, but it does not occur there. The genus is endemic in Utterkhand, India.
1. |
+ |
Corolla with distinct abaxial spur (or 2 spurs) or at least gibbous or sacate at base.
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2 |
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– |
Corolla neither spurred nor with gibbous base.
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9
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2. |
+ |
Palate on the lower lip of corolla absent. Capsule dehiscence loculicidal or septcidal.
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23. Nemesia
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– |
Palate on the lower lip of corolla present. Capsule dehiscence usually porate, sometimes operculate or valvate.
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3 |
3. |
+ |
Fertile stamens 4, anthers marginally coherent and forming a ring-like structure.
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4 |
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– |
Fertile stamens 2 or 4, anthers either free or marginally coherent in pairs not forming a ring-like structure.
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5
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4. |
+ |
Anthers ciliate; capsule dehiscence operculate.
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10. Kickxia
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– |
Anthers glabrous; capsule dehiscence valvate.
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22. Nanorrhinum |
5. |
+ |
Abaxial base of corolla tube with distinct long spur.
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6 |
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– |
Abaxial base of corolla gibbous or sometimes saccate.
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7 |
6. |
+ |
Leaves sessile. Spur cylindrical or narrowly conical sometimes linear or filiform, often arcuate.
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16. Linaria |
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– |
Leaves petiolate, at least in the lower portion. Spur conical or fusiform, straight.
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6. Chaenorrhinum |
7. |
+ |
Seeds with a narrow wing.
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21. Misopates |
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– |
Seed without a wing.
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8 |
8. |
+ |
Calyx with adaxial lobe usually conspicuously exceeding 4-abaxial lobes, exceeding corolla tube. Capsule few seeded, indehiscent.
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30. Schweinfurthia |
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– |
Calyx with adaxial lobe equalling or more rarely exceeding 4 abaxial lobes, shorter than corolla tube. Capsule many seeded dehiscent.
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2. Antirrhinum |
9. |
+ |
Trees.
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25. Paulownia |
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– |
Shrubs, subshrubs or herbs.
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10 |
10. |
+ |
Inflorescence cymose, usually composed of main axis and many lateral cymes.
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11 |
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– |
Inflorescence racemose or spikate, often ramified.
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13 |
11. |
+ |
Corolla actinomorphic rotate to subrotate, stamens 4-5, filaments densely woolly.
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35. Verbascum |
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– |
Corolla zygomorphic, stamens 4 or 2, sometimes a fifth median staminode present, filaments not woolly.
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12 |
12. |
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Corolla tube inflated, ventricose.
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32. Scrophularia |
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– |
Corolla tube neither inflated nor ventricose.
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29. Russelia |
13. |
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Upper lip of the corolla galeate.
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14 |
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– |
Upper lip of the corolla variously shaped but never galeate.
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16 |
14. |
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Galea bilobed, lobes recurved, filaments glabrous.
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9. Euphrasia |
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– |
Galea entire, never bilobed. Filaments papilose or hairy rarely glabrous.
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15 |
15. |
+ |
Lower lip of corolla equal or shorter than galea, tube equal to galea.
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24. Odontites |
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– |
Lower lip of corolla and tube longer than galea.
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26. Pedicularis |
16. |
+ |
Anthers with equal (rarely subequal) thecae, usually rounded at base, often confluent at apex, rarely monothecous.
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17 |
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– |
Anthers with subequal to unequal thecae, usually mucronate to apiculate at base, never confluent at apex, if unequal then thecae sometimes oblong and rounded, often monothecous.
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33. Striga |
17. |
+ |
Leaves alternate.
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18 |
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– |
Leaves opposite.
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24 |
18. |
+ |
Plants rosulate.
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19 |
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– |
Plants with leafy stem, sometimes also rosulate leaves present at base.
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22 |
19. |
+ |
Corolla longer than or equal to 50 mm, weakly 2-lipped to subrotate, cylindrical or inflated, sometimes globose, stigma bilobed.
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7. Digitalis |
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– |
Corolla shorter than 50 mm distinctly 2-lipped, stigma capitate.
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20 |
20. |
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Stamens 4; leaves basal and rosulate.
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28. Picrorhiza |
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– |
Stamens 2; leaves rosulate or cauline.
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21 |
21. |
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Calyx spathaceous or of 2 sepals, fruit indehiscent, drupe-like with 2-cocci at maturity. Seed 1 per locule. |
11. Lagotis |
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– |
Calyx deeply 5-lobed, fruit a capsule, seeds numerous.
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37. Wulfeniopsis |
22. |
+ |
Prophylls present.
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1. Anticharis |
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– |
Prophylls absent.
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23 |
23. |
+ |
Stamens 2, shrubs.
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4. Campylanthus |
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– |
Stamens 4, annual, biennial or perennial herbs or subshrubs.
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7. Digitalis |
24. |
+ |
Abaxial stamens usually curved or twisted or Z-like with geniculate or clavate appendages, if reduced then staminode clavate curved or with appendages.
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25 |
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– |
Abaxial stamens straight or curved but without appendages or if reduced then staminodes filiform.
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28 |
25. |
+ |
Corolla with subequal lobes; anthers unilocular.
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27. Peplidium |
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– |
Corolla with distinct lips; anthers with separate thecae.
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26 |
26. |
+ |
Plants with a basal, decussate leaf rosette and lacking the terminal main inflorescence; all flowering shoots axillary.
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18. Lindernia |
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– |
Plants without basal leaf rosette; the stem with decussate leaves and long internodes; flowers solitary axillary or terminal or pseudoaxillary racemes or subumbellate clusters.
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27 |
27. |
+ |
Dehiscence of capsule poricidal, opening on both sides of the septum, at the same time as dehiscence of calyx, ovary with apical and internal hair.
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34. Torenia |
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– |
Dehiscence of capsule septicidal; ovary without apical and internal hairs within the locule.
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18. Lindernia |
28. |
+ |
Fertile stamens 2, abaxial or adaxial stamens reduced to staminodes or lacking.
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29 |
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– |
Fertile stamens 4.
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30 |
29. |
+ |
Leaves dimorphic, emerged leaves linear-lanceolate to ovoid, entire to palmately divided, submerged leaves usually capillary-multifid; abaxial stamens reduced to staminodes; stigma bifid; seeds with smooth testa.
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14. Limnophila |
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– |
Leaves monomorphic, entire submerged leaves usually linear or lanceolate; adaxial stamens reduced to staminodes; stigma peltate; seeds with reticulate testa.
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8. Dopatrium |
30. |
+ |
Leaves with inconspicuous veination (often only mid vein visible) or distinctly palmately veined, usually linear-ovoid to pinnatisect.
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31 |
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– |
Leaves pinnately veined and ovoid-lanceolate to orbicular or deeply pinnatifid to pinnatisect.
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32 |
31. |
+ |
Usually rosulate herbs, rarely with cauline leaves; corolla subrotate, 5-lobed, small; style short; incurved.
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15. Limosella |
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– |
Herbs with cauline leaves; corolla campanulate, equally or subequally 5-lobed or subrotate; style straight, filiform.
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3. Bacopa |
32. |
+ |
Calyx distinctly winged, 5-dentate, accrescent in fruit; corolla with distinct palate on lower lip.
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20. Mimulus |
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– |
Calyx neither winged nor accrescent in fruit; corolla without palate on lower lip.
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33 |
33. |
+ |
Corolla subrotate, plane, 4-lobed.
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31. Scoparia |
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– |
Corolla bilobed, upper lip 2-lobed, lower lip 3-lobed.
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12. Lancea |
34. |
+ |
Fruit indehiscent.
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12. Lancea |
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– |
Fruit a capsule.
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35 |
35. |
+ |
Plants rosulate.
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19. Mazus |
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– |
Plants only with cauline leaves.
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36 |
36. |
+ |
Thecae separated by the stipitate, 2-armed connective.
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17. Lindenbergia |
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– |
Thecae separate, but connective rounded, not stipitate.
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37 |
37. |
+ |
Fertile stamens 2, staminodes present or absent.
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36. Veronica |
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– |
Fertile stamens 4.
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13. Leptorhabdos |
38. |
+ |
Plant holoparasitic, lacking chlorophyll. Anther with one theca.
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33. Striga |
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– |
Plant hemiparasitic or autotrophic with green leaves. Anther with two unequal thecae, one fertile and one sterile, reduced to a small appendage.
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5. Centranthera |
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