I n s e ct - p l a n t r e l a tionships

Pollination of Adromischus
Adromischus alstonii has evolved to produce two sources of nectar to attract pollinators and protectors
by Greg Nicolson

On

a recent trip to the Namibian Richtersveld last November, I had the good fortune to observe the pollination system of Adromischus alstonii. I found A. alstonii growing on the Kuamsib Berge (1160 m), which was an interesting find in itself as this species has not been recorded this far north. (According to Pilbeam, Rodgerson and Tribble, see further reading.) Adromischus is a succulent plant that belongs to the Crassulaceae family and is closely related to the genus Cotyledon. There are twenty-eight species in the genus although, with recognized subspecies and considerable variation in some species, there are closer to fifty different plants.

The specimens I saw had large leaves in varying colours of red, cream and green, many of them speckled with red spots. During the ascent of the mountain we encountered a number of specimens. Ernst van Jaarsveld mentioned that he had seen a bee pollinating a flower, and as I have a particular interest in pollination biology, I decided to try and capture the pollinator on camera while we were breaking for lunch on the peak of the Kuamsib Berg. After a few minutes of waiting I heard the buzz of an approaching bee. It visited all three open flowers on the plant and with some luck I managed to capture the bee on camera while it was pollinating (see photo opposite). The bee has

been identified as probably belonging to the genus Amegilla, family Apidae. On closer examination it was clear that the plant was also being visited by many ants that serve as the plants protectors. The plant has evolved to produce two sources of nectar to attract both pollinators and protectors. The nectar in the flower attracts the bee as the pollinator, and the plant also produces nectar on the outside of the flower that attracts ants. These aggressive and territorial insects, in return, protect the plant from other destructive insect visitors. Many plants make use of nectaries away from the flower (known as extrafloral nectaries) in order to attract

This Adromischus inflorescence continued to secrete minute drops of nectar on the outside of the flower for a number of days after being removed from the plant, and attracted invasive Argentine ants. The ants are not the pollinators of the plant, but protect it from unwanted insect visitors. Photo: Greg Nicolson.

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ABOVE LEFT: The waxy succulent leaves of Adromischus alstonii showing the red speckles. ABOVE RIGHT: The inflorescence of Adromischus alstonii was constantly patrolled by ants that feed off the extra-floral nectar in return for protecting the plant from unwanted visitors. RIGHT: The Amegilla bee visited each open flower on the Adromischus alstonii plant for a short period (two seconds) before flying off. BELOW RIGHT: Nectar from nectaries on the inside of the flower is exuded through the sepals to form small drops of sweet, sugary nectar on the outside of the calyx in this Adromischus. Photos: Greg Nicolson.

insects for various reasons including protection, as in this case. On return from the trip Ernst provided me with some more Adromischus (but not A. alstonii) plants in flower to try to determine the source of the extra-floral nectar. Examining the inflorescence using a stereo-microscope showed that the nectar is exuded through the sepals to form minute drops on the calyx, as shown in the photo. It was not possible to determine the exact method that the plant uses to move nectar from the inside of the floral tube to the calyx on the outside, and further anatomical research is needed for this genus. However, it is clear that Adromischus species do not have extra-floral nectaries on the outside of the flower but have evolved to allow nectar to permeate through the floral tube or have minute glands on the outside of the flower that serve the same purpose as extra-floral nectaries. The inflorescence continued to produce extra-floral nectar for a number of days after it was removed from the plant. The invasive Argentine

ants enjoyed the nectar, as the accompanying photo shows. There seems to be no record of extrafloral nectar in the genus Adromischus or in the family Crassulaceae; therefore making this interesting discovery a first record. This is a good example of the highly evolved plant-insect relationships so common in southern Africa. It is amazing to think that the production of minute drops of nectar on the outside of the flower can play such a significant role in ensuring the survival of this beautiful plant in some of the most arid areas of southern Africa. This is certainly a most interesting and amazing plant-insect relationship that deserves further investigation.

What does that mean? calyx collective term for all the sepals, particularly when they are more or less joined together inflorescence the flowerhead or collection of flowers on a plant nectary the organ in which nectar is secreted sepals the outer floral segments in flowers

Acknowledgements Ernst van Jaarsveld for his help and enthusiasm. Margie Cochrane of Iziko South African Museum for identifying the bee. Further reading Pilbeam, J., Rodgerson, C. & Tribble, D. 1998. The Cactus File Handbook 3: Adromischus. The Cactus File, Southampton. The author Greg Nicolson is an environmental science graduate from UCT with a passion for fynbos. He is now a botanical photographer with a keen interest in botany and pollination biology. He can be contacted at gregnicolson@gmail.com.
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