Bignoniaceæ.—This family contains many tendril-bearers, some twiners, and some root-climbers. The tendrils always consist of modified leaves. Nine species of Bignonia, selected by hazard, are here described, in order to show what diversity of structure and action there may be within the same genus, and to show what remarkable powers some tendrils possess. The species, taken together, afford connecting links between twiners, leaf-climbers, tendril-bearers, and root-climbers.
Bignonia (an unnamed species from Kew, closely allied to B. unguis, but with smaller and rather broader leaves).—A young shoot from a cut-down plant made three revolutions against the sun, at an average rate of 2 hrs. 6 m. The stem is thin and flexible; it twined round a slender vertical stick, ascending from left to right, as perfectly and as regularly as any true twining-plant. When thus ascending, it makes no use of its tendrils or petioles; but when it twined round a rather thick stick, and its petioles were brought into contact with it, these curved round the stick, showing that they have some degree of irritability. The petioles also exhibit a slight degree of spontaneous movement; for in one case they certainly described minute, irregular, vertical ellipses. The tendrils apparently curve themselves spontaneously to the same side with the petioles; but from various causes, it was difficult to observe the movement of either the tendrils or petioles, in this and the two following species. The tendrils are so closely similar in all respects to those of B. unguis, that one description will suffice.
Bignonia unguis.—The young shoots revolve, but less regularly and less quickly than those of the last species. The stem twines imperfectly round a vertical stick, sometimes reversing its direction, in the same manner as described in so many leaf-climbers; and this plant though possessing tendrils, climbs to a certain extent like a leaf-climber. Each leaf consists of a petiole bearing a pair of leaflets, and terminates in a tendril, which is formed by the modification of three leaflets, and closely resembles that above figured (fig. 5). But it is a little larger, and in a young plant was about half an inch in length. It is curiously like the leg and foot of a small bird, with the hind toe cut off. The straight leg or tarsus is longer than the three toes, which are of equal length, and diverging, lie in the same plane. The toes terminate in sharp, hard claws, much curved downwards, like those on a bird’s foot. The petiole of the leaf is sensitive to contact; even a small loop of thread suspended for two days caused it to bend upwards; but the sub-petioles of the two lateral leaflets are not sensitive. The whole tendril, namely, the tarsus and the three toes, are likewise sensitive to contact, especially on their under surfaces. When a shoot grows in the midst of thin branches, the tendrils are soon brought by the revolving movement of the internodes into contact with them; and then one toe of the tendril or more, commonly all three, bend, and after several hours seize fast hold of the twigs, like a bird when perched. If the tarsus of the tendril comes into contact with a twig, it goes on slowly bending, until the whole foot is carried quite round, and the toes pass on each side of the tarsus and seize it. In like manner, if the petiole comes into contact with a twig, it bends round, carrying the tendril, which then seizes its own petiole or that of the opposite leaf. The petioles move spontaneously, and thus, when a shoot attempts to twine round an upright stick, those on both sides after a time come into contact with it, and are excited to bend. Ultimately the two petioles clasp the stick in opposite directions, and the foot-like tendrils, seizing on each other or on their own petioles, fasten the stem to the support with surprising security. The tendrils are thus brought into action, if the stem twines round a thin vertical stick; and in this respect the present species differs from the last. Both species use their tendrils in the same manner when passing through a thicket. This plant is one of the most efficient climbers which I have observed; and it probably could ascend a polished stem incessantly tossed by heavy storms. To show how important vigorous health is for the action of all the parts, I may mention that when I first examined a plant which was growing moderately well, though not vigorously, I concluded that the tendrils acted only like the hooks on a bramble, and that it was the most feeble and inefficient of all climbers!
Bignonia Tweedyana.—This species is closely allied to the last, and behaves in the same manner; but perhaps twines rather better round a vertical stick. On the same plant, one branch twined in one direction and another in an opposite direction. The internodes in one case made two circles, each in 2 hrs. 33 m. I was enabled to observe the spontaneous movements of the petioles better in this than in the two preceding species: one petiole described three small vertical ellipses in the course of 11 hrs., whilst another moved in an irregular spire. Some little time after a stem has twined round an upright stick, and is securely fastened to it by the clasping petioles and tendrils, it emits aërial roots from the bases of its leaves; and these roots curve partly round and adhere to the stick. This species of Bignonia, therefore, combines four different methods of climbing generally characteristic of distinct plants, namely, twining, leaf-climbing, tendril-climbing, and root-climbing.
In the three foregoing species, when the foot-like tendril has caught an object, it continues to grow and thicken, and ultimately becomes wonderfully strong, in the same manner as the petioles of leaf-climbers. If the tendril catches nothing, it first slowly bends downwards, and then its power of clasping is lost. Very soon afterwards it disarticulates itself from the petiole, and drops off like a leaf in autumn. I have seen this process of disarticulation in no other tendrils, for these, when they fail to catch an object, merely wither away.
Bignonia venusta.—The tendrils differ considerably from those of the previous species. The lower part, or tarsus, is four times as long as the three toes; these are of equal length and diverge equally, but do not lie in the same plane; their tips are bluntly hooked, and the whole tendril makes an excellent grapnel. The tarsus is sensitive on all sides; but the three toes are sensitive only on their outer surfaces. The sensitiveness is not much developed; for a slight rubbing with a twig did not cause the tarsus or the toes to become curved until an hour had elapsed, and then only in a slight degree. Subsequently they straightened themselves. Both the tarsus and toes can seize well hold of sticks. If the stem is secured, the tendrils are seen spontaneously to sweep large ellipses; the two opposite tendrils moving independently of one another. I have no doubt, from the analogy of the two following allied species, that the petioles also move spontaneously; but they are not irritable like those of B. unguis and B. Tweedyana. The young internodes sweep large circles, one being completed in 2 hrs. 15 m., and a second in 2 hrs. 55 m. By these combined movements of the internodes, petioles, and grapnel-like tendrils, the latter are soon brought into contact with surrounding objects. When a shoot stands near an upright stick, it twines regularly and spirally round it. As it ascends, it seizes the stick with one of its tendrils, and, if the stick be thin, the right—and left-hand tendrils are alternately used. This alternation follows from the stem necessarily taking one twist round its own axis for each completed circle.
The tendrils contract spirally a short time after catching any object; those which catch nothing merely bend slowly downwards. But the whole subject of the spiral contraction of tendrils will be discussed after all the tendril-bearing species have been described.
Bignonia littoralis.—The young internodes revolve in large ellipses. An internode bearing immature tendrils made two revolutions, each in 3 hrs. 50 m.; but when grown older with the tendrils mature, it made two ellipses, each at the rate of 2 hrs. 44 m. This species, unlike the preceding, is incapable of twining round a stick: this does not appear to be due to any want of flexibility in the internodes or to the action of the tendrils, and certainly not to any want of the revolving power; nor can I account for the fact. Nevertheless the plant readily ascends a thin upright stick by seizing a point above with its two opposite tendrils, which then contract spirally. If the tendrils seize nothing, they do not become spiral.
The species last described, ascended a vertical stick by twining spirally and by seizing it alternately with its opposite tendrils, like a sailor pulling himself up a rope, hand over hand; the present species pulls itself up, like a sailor seizing with both hands together a rope above his head.