Chapter IV — 59 — Nepenthes 



pitchers of N. Rafflesiana" (which is similar to N. hicalcarata in all 

 respects except that it lacks the canine-like thorns), "and empties 

 them of their prey, but not those of A^. hicalcarata, in which the very 

 sharp spurs are so arranged that the tarsius is certainly held and 

 pierced when he inserts his head to see what there is in the pitcher." 

 GoEBEL remarks of this idea that more study of the matter in the 

 habitat is required. In N. intermedia the spurs are interesting because 

 they are broad, and are quite obviously made up of a group of corru- 

 gations; they are not sharp and tooth-Hke, and could not act in the 

 manner described by Burbidge for N. hicalcarata. Yet so far as we 

 know, the latter shows no superiority over the former or over N. 

 Rafflesiana in the struggle for existence. /V. intermedia is a hybrid of 

 horneensis and Rafflesiana (the former parent is uncertain, Mac- 

 farlane). If this occurred in nature it would be doubtful if the 

 specialized tooth-Hke portion of the peristome could act adaptively as a 

 beginning for the condition seen in N. hicalcarata. 



The several interpretations of the morphology of the Nepenthes 

 leaf, as resumed in part by Troll (1932, 1939), are the following: 



1. The Hd is the lamina of the leaf, the rest is the petiole with 

 highly specialized regions, phyllodial at the base. This view is trace- 

 able to A. P. DE Candolle (1827). Among others Goebel took this 

 position in his earlier writings (1884). The recognition by Hooker 

 that the spur is the true organic apex of the leaf threw this out of 

 court. According to Bower, Goebel regarded the lid as only a part 

 of the lamina, the rest appearing in modified form as the pitcher, 

 tendril, etc. 



2. Instead of regarding the laminar portion of the leaf as petiolar, 

 WuNSCHMANN (1872) preferred to see in it the "lower part of the leaf 

 blade", and therefore that the leaf is non-petiolate. The evidence 

 from development denies this. 



J. The pitcher has arisen phylogenetically as an apical gland, 

 which through enlargement and specialization became the complex of 

 organs which we now know. This, Hooker's interpretation, was 

 based in part on embryological observations and by comparison with 

 such leaves as that of Flagellaria, Gloriosa which have a cirrhus, a 

 terminal tenuous apex serving as a tendril. Faivre held a somewhat 

 similar view that the pitcher arises in the elongated midrib. But the 

 spur is, as said above, the organic apex of the leaf (Hooker). 



4. The leaf arises as a peltate one. According to this view the 

 pitcher is a peltate leaf in which the margin is contracted so that the 

 upper surface lines a hollow organ, the pitcher. Its outer surface is 

 the lower leaf surface. Dickson, receiving his impulse from Baillon's 

 examination of the embryology of the Sarracenia leaf, and impressed 

 by the analogy supplied by the interrupted leaf of Codiaeum sp., wrote 

 "it seems highly probable that in Nepenthes we have to deal with a 

 leaf, the lamina of which is interrupted in the middle of its course by 

 becoming reduced to a midrib and that, while the proximal portion of 

 the lamina retains its typical form of a flat expansion, the distal por- 

 tion becomes peltately expanded into a funnel or pitcher. " But 

 Troll, though conceding the outward resemblance, one which strikes 

 anyone who has made the comparison, even to the peltation of the 



