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systems within the Proteaceae have favoured bird or mammal-pollinated taxa. For 
example, Collins and Rebelo (1987) add new information on functional morphology 
and floral behaviour of Proteaceae in Australia and southern Africa. However, they 
concentrate on vertebrate-pollinated genera and remark that information on 
entomophilous systems remains 'rudimentary'. 
Second, Sussman and Raven (1978) offered a novel interpretation of the wiry 
protostigmas found in most genera in the Proteaceae. They suggested that there had 
been an early association between angiosperm flowers and small, wingless mammals. 
The tough, curved style served both as a pollen presenter and as a 'rung' for a 
climbing mammal. Arboreal rodents and marsupials in Australia do feed on the 
nectar and pollen of native Proteaceae (Turner 1982). However, their role as agents 
of cross-pollination remains controversial since Hopper and Burbidge (1982) accuse 
these mammals of consuming or grooming away pollen deposits in fur and whiskers 
before it is transferred to the stigma of a second genotype. There has been additional 
evidence that large bats (Megachiroptera) may also pollinate some Banksia spp. (Law 
1994) as first predicted by Johnson and Briggs (1975). However, research directed 
exclusively towards vertebrate pollination of Australasian Proteaceae will always 
result in a biased and incomplete interpretation of floral evolution of this family. 
If pollination by vertebrates is an ancestral feature of the Proteaceae why do so 
many rainforest relicts (e.g. Placosperiuum, Eidothea, Carnarvonia, Sphalmium, Neoritcs, 
Cardwellia, Buckinghamia, Opisthiolepis, Floydia, Musgravea) lack the full suite of 
reproductive characters associated with vertebrate-pollination? In fact, pollination 
by birds or wingless mammals has been recorded far more frequently in the Proteaceae 
of sclerophyllous woodlands and shrublands (Hopper and Burbidge 1982, 1986; 
Paton 1986; Turner 1982; Collins and Rebelo 1987). Similarly, why do most basal 
lineages in the Proteaceae, such as the Persoonioideae, Bellendenoideae, 
Carnarvonioideae, Sphalmioideae, Eidotheoideae and the tribe Conospermeae 
(Proteoideae) lack the suite of characters associated with vertebrate pollination? 
Based on patterns of character distribution in the Proteaceae, Johnson and Briggs 
(1975) inferred that the family was primitively entomophilous and restricted to closed, 
mesothermic forests. Evolutionary shifts to vertebrate pollination and to xeric habitats 
was inferred to be a secondary and often recurrent process. Primitive occurrence in 
rainforests has been supported, in part, by the fossil evidence (Truswell 1990). 
The genus Persoonia would appear to be the most logical choice to help close the 
information gap on the role of insects in floral evolution within the Proteaceae. 
Comparatively few Persoonia species occur in rainforests; most are shrubs or small 
trees of sclerophyll woodlands and shrublands. Consequently, many Persoonia species 
form far denser and more readily accessible populations (see Weston 1991, 1994; 
Weston and Johnson 1991) than those rainforest relicts listed above. 
Of greater importance, Persoonia belongs to the Persoonioideae, the only subfamily that 
completely lacks proteoid roots and thus is likely to be one of the most basal lineages in 
the family. The Persoonioideae also shows no evidence of protostigma development 
considered synapomorphic in at least three different lineages that include most of the 
genera in the family. Persoonia may then provide a model system for understanding 
evolutionary trends in the functional morphology of the flowers of Proteaceae. 
Bees in the long-tongue families Anthophoridae (Allodapitla, Amegilla, Exonettra), 
Apidae (Apis, Trigona) and Megachilidae {Chalicodoma, Megachile) have been reported 
to collect nectar and/or pollen on Persoonia species (Armstrong 1979). Pollination 
mechanisms in Persoonia were first addressed by two amateur entomologists in a 
little known paper (Payment 1950). Payment described three new species of short- 
tongue bees in the genus Cladocerapis (now Leioproctus subgenus Cladocerapis) including 
