460 ANGIOSPERMAEDICOTYLEDONES 



Knuth, Bot. Centralbl., Cassel, xl, 1889, p. 273. Pachypleurum : Herm. Miiller r 

 ' Alpenblumen,' p. 120. Sanicula: Herm. Mttller, ' Weit. Beob.,' I, p. 303 ; Schulz, 

 op. cit., I, p. 40. Scandix : Henslow, Trans. Linn. Soc. (Bot.), Ser. 2, i, 1880, 

 P- 365; Schulz, op. cit., I, p. 61. Turgenia: Schulz, op. cit., I, p. 60. As regards 

 Astrantia major Z., Eryngium campestre Z., and Sanicula europaea Z., it is expressly 

 stated by Schulz that the styles project from the hermaphrodite flowers at an early 

 stage, giving the appearance of protogyny, although the stigmas are still immature. 

 Kerner goes too far in his conception of protogyny, when, for example (op. cit., 

 p. 312), he asserts that the Rosaceae and Cruciferae are exclusively protogynous, 

 and (p. 310) even speaks of protogyny when the anthers dehisce ten to fifteen minutes 

 after the flower opens. 



Kirchner (op. cit.) discusses protogyny among the Umbelliferae. The first 

 accounts were given by A. F. Foerste (Bot. Gaz., Chicago (111.), vii, 1882, pp. 70-1), 

 and W. Trelease (loc. cit.), and have reference to Erigenia bulbosa Nutt. The 

 protogyny of this species was subsequently confirmed by C. Robertson (op. cit., xiii. 

 1888, p. 193), and that of four other North American Umbelliferae was also 

 established, i.e. Sanicula marilandica Z., Zizia aurea Koch, Pimpinella integerrima 

 Benth. et Hook. /., and Polytaenia Nuttallii DC. During the autumn of 1891 

 Kirchner had the good fortune to observe on the Lido near Venice a case of distinct 

 protogyny in one of the European Umbelliferae, Echinophora spinosa Z., the stigmas 

 of which are mature before any of the anthers dehisce. 



In protandrous hermaphrodite flowers the stamens ripen successively. When 

 the flower opens, one anther springs up so as to occupy the middle of the flower, its 

 filament being curved. When it has shed its pollen the filament bends back towards 

 the corolla, a second stamen assuming the position of the first, and so on. The 

 styles usually begin to grow when all the stamens of a flower, or even of an umbel, 

 are withered; they then diverge so that the terminal stigmas occupy the middle of 

 the flower. 



In correspondence with the exposed position of the nectar in most species, the 

 large majority of guests are short-tongued insects (flies, beetles, wasps, and some 

 bees). Lepidoptera with their long proboscis are only seen occasionally as visitors, 

 but seek out more frequently the flowers of those Umbelliferae which belong to 

 class S. The more specialized bees (honey-bees, humble-bees, and the like) also 

 resort in greater numbers to flowers of the latter class, and generally only collect 

 pollen from those of class E; sometimes, however, they lick the nectar of such 

 flowers. Many species are andromonoecious. 



Warnstorf makes the following remarks (Verh. bot. Ver., Berlin, xxxviii, 1896.) 

 A definite line of adaptation is noticeable in our native Umbelliferae as regards 

 flower pollination. In order to appreciate this fully, all the inflorescences of a plant 

 or branch have to be considered. It then appears that in by far the greater number 

 of cases, the primary umbel only bears hermaphrodite flowers in its umbellules, there 

 being rarely a few male flowers in the centre, while very rarely all the flowers have 

 become female by the degeneration of the anthers. The last condition is associated 

 with the presence of relatively long styles. In most smaller secondary umbels, 

 as a general rule, the marginal flowers of the umbellules are hermaphrodite only, 

 while the central ones are male: more rarely all are hermaphrodite, as in the 



