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Published In: Genera Plantarum 128. 1789. (4 Aug 1789) (Gen. Pl.) Name publication detailView in BotanicusView in Biodiversity Heritage Library

Project Name Data (Last Modified On 6/2/2011)
Acceptance : Accepted
Project Data     (Last Modified On 6/3/2011)
Contributor Text: YASIN J. NASIR
Contributor Institution: National Herbarium,Pakistan Agricultural Research Council, Islamabad.
General/Distribution: A family of c. 100 genera and 2000 species distributed in temperate, especially mediterranean and tropical regions. Represented in Pakistan by 32 genera and 135 species of which c. 5 species of Cordia, Echium and Anchusa are cultivated.
Comment/Acknowledgements: As the flower and especially the fruit are most important diagnostically, it is helpful to elaborate and explain some of the terms used in the text. The corolla size includes the length of the tube and limb. Faucal appendages, as used in some literature is replaced here by “throat scales”. The style which is diagnostic, may sometimes be. split once or twice (Arnebia, Cordia, Ehretia etc.) and hence the term 2-fid or 4-fid. There are generally as many stigmas as there are styles.

Various terminology has been used to describe the characters of the nutlets-sometimes of an elaborate nature (M. Popov in Schischkin, Fl. URSS 19:98-100. 1953-English translation, Jerusalem, 1974). The attachment scar (cicatrice) of the nutlet is referred to here as the areole. The gynobase is that part of torus (receptacle) on which the nutlets are borne; its shape varies from flat to elongate and may bear the persistent style at its apex. When the gynobase is flat (Castrocotyle, Lycopsis, Nonea, Echium etc.) the nutlets are said to be basally attached, if ± pyramidal (Cynoglossum, Paracaryum, Solenanthus etc.) then the nutlets are ventrally attached. The nutlets may be variously oriented in relation to their plane of attachment (viz. erect, transverse or oblique). The dorsal middle area (disc) is the side that is turned towards the apex or faces the style (adaxial); it may be rimmed or crested by a thickened ring or flatmargined. The margin may be formed by the fused bases of the appendages, or the appendages may be free. Appendages are glochidiate if they are stiff and apically barbed (Heterocaryum, Cynoglossum, Lappula).

Acknowledgements: I am grateful to the Directors/Curators of the following for the loan of herbarium specimens: BM (NH), E, G, GH K, KUH, M, PES, PPFI-B, US, W. I am also thankful to Mr. I. C. Hedge (EdinbuRGH) for going though the manuscript and for his helpful advice. The financial assistance received from the United States Department of Agriculture under P1,480 is gratefully acknowledged.


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Annual, biennial or perennial herbs or less often shrublets or trees, usually with coarse hairs that arise from a tuberculate base; hairs appressed, subappressed or setose. Leaves alternate, simple, exatipulate. Inflorescence initially a scorpiod or helicoid cynic that uncoils at anthesis, sometimes spicate, loosely cymose or with solitary flowers. Flowers generally regular and bisexual or less often irregular, sometimes heterostylous. Calyx 5-lobed, often enlarging in fruit. Corolla 5 -lobed, salverform, funnelform or campanulate. Throat glabrous or with scales (appendages) or a zone of hairs, a nectiferous ring or collar sometimes present at base of corolla tube. Stamens 5, epipetalous, alternating with the corolla lobes, equal or unequal. Filaments exserted beyond the corolla tube or not. Ovary superior, 4-locular at maturity. Style gynobasic or terminal, usually simple; stigma entire or by splitting of style, 24-lobed. Fruit of 2 or 4 nutlets or drupaceous. Nutlets borne on a gynobase that is flat or raised and fused to the style base, keeled or not, attachment scar (areole) narrow or not; often differentiated into a disc and margin; margin smooth of appendiculate; surface smooth to hairy or variously ornamented.

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1.Trees or shrubs
2.Style 4-fid
2.Style 2-fid.
1.Herb, sometimes with a suffruticose base, rarely shrubs
3.Style terminal.
4.Styles 2, not dilated towards apex or base.
4.Style 1, dilated towards apex or base.
3.Style gynobasic.
5.Fruit with 2 nutlets.
5.Fruit with 4 nutlets, at least in young stage.
6.Corolla distinctly zygomorphic
7.Gynobase flat. Style usually 2-fid at apex, rarely entire. Nutlets without a lateral attachment
7.Gynobase pyramidal. Style simple. Nutlets with a sessile lateral attachment
6.Corolla actinomorphic
8.Tips of anthers elongated and contorted
8.Tips of anthers obtuse or apiculate, not contorted
9.Stamens unequal, with 1 anther larger than the other four
9.Stamens of equal size
10.Corolla throat with scales absent; throat sometimes villous
11.Flowers nodding at anthesis; corolla lobes reduced. Anther connective elongated
11.Flowers ± erect; corolla lobes distinct. Anther connective not elongated
12.Corolla throat vinous. Style simple
12.Corolla throat glabrous. Style divided
13.Corolla throat glabrous. Style 24-fid, rarely undivided; stigmas 2-4
13.Corolla throat glabrous to hairy; style 2-fid; stigmas 2
14.Annuals. Corolla throat hairy. Nutlets ± pear shaped, surface tuberculate or rugose
14.Perennials. Corolla throat not hairy. Nutlets ovoid
10.Corolla throat with scales present; sometimes sparsely hairy or with small glands
15.Margin and middle dorsal area of nutlets glabrous to hairy or variously ornamented but never prickly or appendiculate
16.Calyx irregular; much enlarged (12-14 mm) and saucer-like in fruit: lobes unequal, dentate in between
16.Calyx regular, not much enlarged (up to 10 mm) in fruit; lobes equal or subequal, not dentate in between
17.Throat scales fimbriate or densely pubescent. Areola prominently hollowed, with a thick girdle
17.Throat scales not fimbriate, glabrous or puberulous. Arcola not prominently hollowed out
18.Calyx in fruit stellately spreading
18.Calyx in fruit not stellately spreading
19.Flowers axillary (at least lower ones
19.Flowers in terminal cymes
20.Areole of nutlet with surface of attachment concave, margin prominently thickened
20.Areole of nutlet with surface of attachment flat, margin not thickened
21.Perennials. Corolla infundibuliform or cylindrical-campanulate, blue to deep purple-blue
21.Annual or perennial. Corolla rotate or hypocrateriform, blue or white
22.Flowers ebracteate. Nutlets smooth
22.Flowers (at least the lower ones) bracteate. Nutlets minutely tuberculate
15.Margin and middle dorsal area of nutlets prickly or appendiculate
23.Nutlets attached laterally to the gynobase and decurrent on style
24.Dorsal middle area of nutlet partly covered by the incurved margin
24.Dorsal middle area of nutlet not covered by the margin
23.Nutlets not as above
25.Middle area of nutlet ovate; inner face above areola keeled throughout its length
25.Middle area of nutlet not ovate; inner face above areola not keeled throughout its length
26.Nutlets inseparable from gynobase; base of
26.Nutlets separable from gynobase; bases of marginal appendages usually not confluent
27.Nutlets ± globose or napiform
28.Filaments longer than anthers, attached above throat scales. Stamens distinctly exserted
28.Filaments equalling anthers, attached below throat scales. Stamens included or barely exserted
29.Corolla rotate, the tube not exceeding calyx length. Stamens included
29.Corolla funnel-shaped, tube exceeding calyx length. Stamens exserted
27.Nutlets pyramidal to truncate-pyramidal-oblong
30.Nutlets ± equalling the gynobase length, attach-ed for nearly the whole length;, but not over topping the calyx
30.Nutlets exceeding the gynobase and style, attached obliquely
31.Tall plants with broad leaves. Fruiting calyx reflexed
31.Generally caespitose plants with small narrow leaves. Fruiting calyx not reflexed
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