P1. Syst. Evol. 188:1 - 16 (1993)
--Hant
Systematics
and
Evolution
© Springer-Verlag 1993
Printed in Austria
Pollination biology and sexual differentiation of
Osyris alba (SantMaceae) in the Mediterranean region
G. ARONNE, C. C. WILCOCK, and P. PIZZOLONGO
Received February 27, 1991; in revised version January 20, 1993
Key words: Santalaceae, Osyrls alba. - Phenology, dioecy, floral morphology, floral mim-
icry, pollination biology. - Mediterranean shrublands.
Abstract: Osyris alba L. is a widespread dioecious hemiparasitic shrub of S Europe, N
Africa, and SW Asia. Male inflorescences are multiflowered whereas each female inflo-
rescence is reduced to a single flower with persistent enlarged bracts. Pollination is a
prerequisite for fruit and seed development and wind is unlikely to be an effective means
of pollen spread. In southern Italy pollen is transported by small unspecialized flies and
beetles. Both male and female flowers produce an indistinguishable sweet odour. Male
flowers are produced in large numbers and over a larger period than the females and provide
pollen, nectar, and staminal hairs as rewards for pollinators. The presence and function
of stamina1 hairs with tip cells in Osyris alba has been reported for the first time. Female
flowers are rewardless, producing neither mature pollen, nectar nor stamina1 hairs, but
possess three modified yellow indehiscent anthers containing no viable pollen which may
provide a strong visual feeding stimulus for pollinators. It is suggested that pollinators are
attracted by deceit to female flowers by mimicry of the males and the floral mimicry is,
therefore, intraspecific and intersexual. The floral characteristics and flowering phenology
of male and female plants are consistent with this kind of mimicry. The female flower
possesses a tricarpellary ovary with three ovules of which only one develops. The single
seed, containing a small embryo and a large, rich endosperm, is borne in a red fleshy bird-
dispersed fruit. The reduction in seed number per flower to one highly nutrient-invested
seed, together with a reduction of the multiflowered inflorescence to a solitary flower and
the sequential production of ripe fruits over an extended fruiting season, suggest that the
female function is markedly resource-limited. It is suggested that, although all the repro-
ductive characteristics present in Osyris alba, as well as hemiparasitism, had probably
evolved before the end of the tropical Tertiary, they are of adaptive advantage in the
nutrient and water-limited environment of the Mediterranean maquis.
Osyris alba L. belongs to the Santalaceae, a family distributed in the tropics and
temperate regions but being concentrated in relatively dry areas (HEYWOOD 1978).
It is a widespread, perennial, dioecious evergreen shrub of S Europe, N Africa,
and SW Asia and a frequent member of the shrub community of the Mediterranean
maquis. It has a ribbed photosynthetic stem with narrow leaves, often caducous
in the summer, and is capable of extensive vegetative spread. As is typical of the
family (KuJ3:1969), Osyris alba is an unspecialized hemiparasite, producing haus-