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C. J. Potgieter et al.: Pollination in Plectranthus 109<br />
I /<br />
Fig. 7. Comparisons <strong>of</strong> floral tube length with proboscis length <strong>of</strong> visiting dipterans for two species <strong>of</strong> Plectranthus. a P.<br />
hilliardiae above with Stenobasipteron sp. below, b P. zuluensis above with Psilodera sp. A below; note the close correlation<br />
between proboscis and floral tube length. Bars: 5 mm<br />
visiting bees and flies in most cases correspond well<br />
with that <strong>of</strong> the corolla-tube, style and filaments <strong>of</strong> the<br />
Plectranthus species in question (Fig. 4). Insects with<br />
shorter mouthparts tend to concentrate on pollen<br />
collection.<br />
Pollen loads. Voucher specimens <strong>of</strong> anthophorid<br />
bees and nemestrinid and other flies that yielded a<br />
high percentage <strong>of</strong> Plectranthus pollen show that<br />
although pollen transfer is achieved for Plectranthus,<br />
these vectors are not exclusive to the genus. A few<br />
observations <strong>of</strong> vectors moving between Plectranthus<br />
flowers and those <strong>of</strong> other genera, e.g. Lobelia spp.<br />
(Campanulaceae) and IsogIossa spp. (Acanthaceae),<br />
have been made and this is supported by the<br />
placement <strong>of</strong> foreign pollen in patches inaccessible<br />
to Plectranthus stigmas on some insect specimens. It<br />
is difficult to distinguish between pollen grains from<br />
different species <strong>of</strong> Plectranthus, but it is seldom that<br />
more than two species grow in close proximity.<br />
Differences in filament and style lengths place pollen<br />
in different areas on the insect body, thus isolating<br />
pollen from different Plectranthus species. The lack <strong>of</strong><br />
pollen on the voucher specimens <strong>of</strong> Lepidoptera<br />
suggests that butterflies are not important polinators<br />
<strong>of</strong> P. madagascariensis. The Pieridae were present <strong>of</strong><br />
a limited period <strong>of</strong> the flowering season and were<br />
seldom encountered.<br />
Nectar robbing. The proboscis <strong>of</strong> Allodape pernix<br />
is too short to reach nectar legitimately, hence the<br />
behaviour <strong>of</strong> robbing nectar through a hole pierced at<br />
the base <strong>of</strong> the corolla tube. This corresponds to<br />
Chapter 2/ 25<br />
primary nectar robbing according to the system<br />
suggested by Inouye (1980). Glandular trichomes are<br />
abundant on the calyces <strong>of</strong> Plectranthus and in other<br />
members <strong>of</strong> the Lamiaceae have been shown to<br />
secrete essential oils that have insecticidal properties<br />
(see Levin 1973 and Konstantopoulou et al. 1992).<br />
Individuals <strong>of</strong> A. pernix do not appear to be deterred<br />
by these oils, as it is the corolla tissue, containing<br />
fewer terpenoid-containing trichomes, that is pierced<br />
above the calyx and nectary.<br />
The generally small body size, short proboscis<br />
length and pollen-collecting habit <strong>of</strong> the insects listed<br />
in Table 5 make them unlikely primary pollinators <strong>of</strong><br />
Plectranthus. In his account on floral larceny Inouye<br />
(1980) refers to pollen theft which is the collection <strong>of</strong><br />
pollen directly from the anthers by small bees, without<br />
contacting the stigma. This is probably the case for<br />
these species and that <strong>of</strong> the nectar robber A. pernix,<br />
but the possiblility that some contribution is made to<br />
intra-specific pollen transfer cannot be discounted.<br />
Comparison <strong>of</strong> tube length with proboscis<br />
length. This study records several new groups <strong>of</strong><br />
visitors to Plectranthus: flies <strong>of</strong> the families Nemes-<br />
trinidae, Acroceridae and Tabanidae, and lepidopter-<br />
ans <strong>of</strong> the families Lycaenidae and Pieridae. The<br />
occurrence <strong>of</strong> pollination by long-tongued flies<br />
(primarly <strong>of</strong> the family Nemestrinidae, but also one<br />
species <strong>of</strong> Tabanidae) is confirmed for both longer-<br />
and shorter-tubed species <strong>of</strong> Plectranthus. Species<br />
with intermediate tube length (P. ecklonii and P.<br />
zuluensis) allow for visits by shorter-tongued insects -