Thank you to Brian aka aeranthes@Flickr for sharing the photograph today (original image via BPotD Flickr Pool). The illustration is by John Nugent Fitch, and is now a public domain work after having been originally published in the 1882-1897 publication, The Orchid Album (link to additional illustrations).
Angraecum sesquipedale has a bevy of common names, including Star of Bethlehem orchid, comet orchid and Darwin’s orchid. The latter name is a reference, of course, to Charles Darwin, who wrote the following in 1862 in On the various contrivances by which British and foreign orchids are fertilised by insects, and on the good effects of intercrossing.:
“I fear that the reader will be wearied, but I must say a few words on the Angræcum sesquipedale, of which the large six-rayed flowers, like stars formed of snow-white wax, have excited the admiration of travellers in Madagascar. A whip-like green nectary of astonishing length hangs down beneath the labellum. In several flowers sent me by Mr. Bateman I found the nectaries eleven and a half inches long, with only the lower inch and a half filled with very sweet nectar. What can be the use, it may be asked, of a nectary of such disproportional length? We shall, I think, see that the fertilisation of the plant depends on this length and on nectar being contained only within the lower and attenuated extremity. It is, however, surprising that any insect should be able to reach the nectar: our English sphinxes have probosces as long as their bodies: but in Madagascar there must be moths with probosces capable of extension to a length of between ten and eleven inches!”
“The rostellum is broad and foliaceous, and arches rectangularly over the stigma and over the orifice of the nectary: it is deeply cleft, with the cleft enlarged or widened at the end. Hence the rostellum pretty closely resembles that of Calanthe after the disc has been removed. The under surfaces of both margins of the cleft near its end are bordered by narrow strips of viscid membrane, easily removed; so that there are two distinct viscid discs. To the middle of each disc a short membranous pedicel is attached; and each pedicel carries at its other end a pollen-mass. Beneath the rostellum a narrow, ledge-like, viscid stigma is seated.”
“I could not for some time understand how the pollinia of this Orchid were removed, or how it could be fertilised. I passed bristles and needles down the open entrance into the nectary and through the cleft in the rostellum with no result. It then occurred to me that, from the length of the nectary, the flower must be visited by large moths, with a proboscis thick at the base; and that to drain the last drop of nectar even the largest moth would have to force its proboscis as far down as possible. To effect this, whether or not the moth first inserted its proboscis by the open entrance into the nectary (as is most probable, from the shape of the flower) or through the cleft in the rostellum, it would ultimately force its proboscis into this cleft, for this is the straightest course, and by slight pressure the whole foliaceous rostellum can be depressed: the distance from the outside of the flower to the extremity of the nectary can be thus shortened by about a quarter of an inch. Hence I took a cylinder, one-tenth of an inch in diameter, and pushed it down through the cleft in the rostellum: the margins readily separated, and were pushed downwards together with the whole rostellum. When I slowly withdrew the cylinder the rostellum rose from its elasticity, and the margins of the cleft upturned and clasped the cylinder. Thus the viscid strips of membrane on the under sides of the cleft rostellum came into contact with the cylinder, and firmly adhered to it; and the pollen-masses were withdrawn. By this means alone I succeeded in each case in withdrawing the pollinia; and it cannot, I think, be doubted that a large moth must thus act; namely, by driving its proboscis up to the very base, through the cleft of the rostellum, so as to reach the extremity of the nectary; and then withdrawing its proboscis with the pollinia attached to it.”
“I did not succeed in imitating the fertilisation of the flower so well as I did in withdrawing the pollinia, but I effected it twice. As the margins of the cleft rostellum must be upturned before the discs adhere to the cylinder, they become, during its withdrawal, affixed some little way from its actual base. The two discs did not always adhere at exactly corresponding points. Now, when a moth inserts its proboscis, with the pollinia affixed to it near the base, into the mouth of the nectary, the pollen-masses will probably be first inserted beneath the rostellum; and during the final exertion, when the moth pushes its proboscis through the cleft of the rostellum, the pollen-masses will almost necessarily be placed on the narrow, ledge-like stigma projecting beneath the rostellum. By acting thus with the pollinia attached to the cylinder the pollen-masses were twice torn off and left glued to the stigmatic surface.”
“If the Angræcum in its native forests secretes more nectar than did the vigorous plants sent me by Mr. Bateman, so that the nectary becomes filled, small moths might obtain their share, but they would not benefit the plant. The pollinia would not be withdrawn until some huge moth, with a wonderfully long proboscis, tried to drain the last drop. If such great moths were to become extinct in Madagascar, assuredly the Angræcum would become extinct. On the other hand, as the nectar, at least in the lower part of the nectary, is stored safe from depredation by other insects, the extinction of the Angræcum would probably be a serious loss to these moths. We can thus partially understand how the astonishing length of the nectary may have been acquired by successive modifications. As certain moths of Madagascar became larger through natural selection in relation to their general conditions of life, either in the larval or mature state, or as the proboscis alone was lengthened to obtain honey from the Angræcum and other deep tubular flowers, those individual plants of the Angræcum which had the longest nectaries (and the nectary varies much in length in some Orchids), and which, consequently, compelled the moths to insert their probosces up to the very base, would be fertilised. These plants would yield most seed, and the seedlings would generally inherit longer nectaries; and so it would be in successive generations of the plant and moth. Thus it would appear that there has been a race in gaining length between the nectary of the Angræcum and the proboscis of certain moths; but the Angræcum has triumphed, for it flourishes and abounds in the forests of Madagascar, and still troubles each moth to insert its proboscis as far as possible in order to drain the last drop of nectar.”
Darwin’s 1862 prediction of a coevolved moth with a proboscis of 35cm was ridiculed by some as being impossible. Others suggested the long nectaries were proof of supernatural creation (read: Darwin’s Madagascan Hawk Moth Prediction (PDF)). Wallace and Darwin responded by detailing how evolutionary processes could develop both a long-nectaried orchid species and a co-evolved moth. Though Darwin did not live to see it, in 1903 a moth was discovered in Madagascar with the characteristics as predicted by Darwin.
For more reading / photographs:
- Angraecum sesquipedale via the University of Connecticut’s Ecology & Evolutionary Biology Greenhouses
- Angraecum sesquipedale via Encyclopaedia angraecorum
- Angraecum sesquipedale via Wikipedia
- Darwin the Botanist and Origins of Life Research podcast (and text) via Scientific American
Happy Birthday, Charles.