he Knersvlakte is a vast expanse of flat gravelly land T that stretches north from Vanrhynsdorp. During spring it is carpeted with flowers and this is the season when tour buses can be seen traversing the national road in search of the ultimate photo opportunity. We had come to the Knersvlakte during the cold months of winter, long before the veld turns into a kaleidoscope of colour. Our mission was to uncover the polli­ nation mysteries of an unusual lily with the most unprepossessing flowers imaginable. , the object of our attention on this trip, has greenish brown flowers at ground level, nested between a giant pair of flat green leaves. We suspected that these flowers are pollinated at night, not by insects, but by mice, which are, by and large, nocturnal animals. And so, a group of slumbering Karoo sheep were startled by the sight of three botanists picking their way among the vygies in the dead of a winter's night. It is an eerie experience to wander about the Knersvlakte after dark; colder than one might imagine and, in the absence of city lights, luminescent with stars. More than twenty years have passed since John Rourke of Kirstenbosch and his American colleague Delbert Wiens made the sensational discovery that a group of Protea is pollinated by rodents. These proteas have dull flower heads situated near ground level, quite unlike their bird-polli­ nated relatives which have brightly coloured conspicuous flower heads. They also have a strong yeasty odour that makes it possible for mice to find the flower heads OF MICE AND during the night. We have long been intrigued by a small group of lilies in Namaqualand that have features MASSONIAS reminiscent of rodent-pollinated proteas. The flowers of these lilies, The pollination of flowers by rodents was thought to be like Massonia depressa and confined to the Proteaceae but recent discoveries show that Androcymbium dubium, are dull the,Namaqualand lily Massonia depressa, and most and grow low on the ground. Could they represent an as yet undocu­ certainly others like it, are also pollinated by rodents. mented case of rodent-pollination? We had located a large flowering by Steve Johnson, School ofBotany and Zoology, population of Massonia depressa in University of Natal, Pietermaritzburg, late July 1998 and laid out a Anton Pauw and Jeremy Midgley, Department ofBotany, number of rodent traps, baited with University of Cape Town peanut butter and rolled oats. We had also been taking measurements of the nectar, which is unusually viscous and jelly-like and collects in a pool at the base of the bowl­ Above A flower head of Massonia depressa nestles between two large leaves lying flat on shaped flower. We noticed that the the sun baked surface of the Knersvlakte in Namaqualand. Photo: A. Pauw. nectar in each flower increased

166 Veld &- Flora December 2001 towards dusk - an indication that nectar, and did not nibble or pollinators of Massonia, we decided the were preparing for a damage the flowers while doing so. to cover plants with wire mesh nocturnal visit. As evening The rodent's snout was soon heavily cages that would exclude rodents, progressed the flowers developed a encrusted with pollen, which was but allow insect access to the strong scent that permeated the air liberally daubed onto the stigma of flowers. At the end of the flowering around us. The scent is not pleasant each flower that it subsequently season we returned to the site and and was likened by one of us to the visited. Having witnessed this found that almost no seeds were smell of a damp sheep! Now we exciting interaction between produced by the covered plants, waited, shivering as the temperature Massonia and its unlikely polli­ while uncovered plants had plunged to near freezing point. nator, we released the gerbil back at produced plenty of seeds. We now By midnight the first of the traps the field site and retired exhausted, felt that we had enough evidence to had been sprung. These traps were but happy, at 3 am. conclude that Massonia depressa is not lethal spring-loaded 'mouse­ Although the results of this first indeed adapted for pollination by traps', but small tunnels into which season were promising, we wanted rodents. the rodents venture but cannot exit to obtain more data from other Pollination by small 'non-flying until set free by their human populations, and so in the following mammals' (to distinguish them from captors. Inside we were surprised to winter we travelled to the bats) we now know, occurs on discover a hairy-footed gerbil Niewoudtville escarpment which several continents. Marsupials visit Gerbillurus paeba, and not a mouse overlooks the Knersvlakte. We set the flowers of various Australian as we had expected. We dabbed its rodent traps among giant boulders shrubs including Banksia, which are fur with sticky tape to collect on a rocky ridge where plants of cousins of our South African pollen, and most importantly, Massonia depressa are plentiful. proteas. In Madagascar, lemurs have collected the droppings that had Several individuals of the Namaqua recently been implicated in the accumulated in the trap. Three more rock mouse Aethomys namaquensis pollination of several plants gerbils were trapped and later the and spiny mouse Acomys including Ravenalia, a close relative microscope revealed that their subspinosus were found in the traps of Strelitzia. Proteas were thought to droppings were packed with and, on examination, they too be the only plants in southern Massonia pollen. This was the proof proved to have Massonia pollen in Africa that rely on rodents for polli­ we needed to show that they had their fur and faeces. After being nation, but now it seems that a been visiting the flowers. After released in the early hours of one guild of South African lilies can be visiting flowers, rodents become morning, a Namaqua rock mouse added to that list. In all, we predict dusted with pollen which they lick even brazenly lapped nectar from that about a dozen Namaqualand off their fur during the grooming Massonia flowers just a few metres lily species with ground-hugging process. In this way pollen is away from where we stood. flowers are pollinated by rodents.Wi ingested, but as the pollen wall is Insects hardly ever visit the virtually impervious to digestion, flowers of Massonia depressa. it eventually ends up in the One reason might be the jelly-like droppings. constituency of the nectar, which Further reading We took one of the gerbils to our would make feeding difficult for Wiens, D., and J. P. Rourke. 1978. Rodent nearby campsite and placed it insects with their narrow syringe­ pollination in southern African Pratea carefully in a darkened terrarium like mouthparts. (Rodents easily lap spp. Nature 276, 71-73. Johnson, S.D., Pauw, A. & Midgley, J. 2001. containing several flowering plants up the nectar with their broad Rodent pollination in the African lily of Massonia depressa. The first tongues.) To establish with certainty Massonia depressa (Hyacinthaceae). reaction of the gerbil was to sniff whether rodents are the primary American Journal of Botany (in press). the air and then, without hesitation, dash across to a Massonia and begin lapping nectar from the flowers. It pushed its snout in amongst the anthers and stigma of each individual flower when draining the

The ground level position of Massonia flowers makes it easy for rodents like this hairy­ footed gerbil GerbjJlurus paeba to get to the nectar. Photo: Steve Johnson & Anton Pauw.