or low woodlands; of the eastern species, B. integrifolia and B. marginata occur in forests; many south-western species such as B. grandis, B. sphaerocarpa, B. sessilis, B. nobilis and B. dallanneyi grow as understorey plants in jarrah (Eucalyptus marginata), wandoo (E. wandoo) and karri (E. diversicolor) forests, with B. seminuda being one of the forest trees in suitable habitat. Most species do not grow well near the coast, notable exceptions being the southern Western Australian species B. speciosa, B. praemorsa and B. repens. Only a few species, such as B. rosserae and B. elderiana (swordfish banksia), occur in arid areas. Most of the eastern Australian species survive in uplands, but only a few of the Western Australian species native to the Stirling Ranges – B. solandri, B. oreophila, B. brownii and B. montana – survive at high altitudes. Studies of the south-western species have found the distribution of Banksia species to be primarily constrained by rainfall. With the exception of B. rosserae, no species tolerates annual rainfall of less than 200 millimetres, despite many species surviving in areas that receive less than 400 millimetres. Banksia species are present throughout the region of suitable rainfall, with greatest speciation in cooler, wetter areas. Hotter, drier regions around the edges of its range tend to have fewer species with larger distributions. The greatest species richness occurs in association with uplands, especially the Stirling Range. Evolution and fossil record There are many fossils of Banksia. The oldest of these are fossil pollen between 65 and 59 million years old. There are fossil leaves between 59 and 56 million years old found in southern New South Wales. The oldest fossil cones are between 47.8 and 41.2 million years old, found in Western Australia. Although Banksia is now only native to Australia and New Guinea, there are fossils from New Zealand, between 21 and 25 million years old. Evolutionary scientists Marcell Cardillo and Renae Pratt have proposed a southwest Australian origin for banksias despite their closest relatives being north Queensland rainforest species. Ecology Banksias are heavy producers of nectar, making them an important source of food for nectivorous animals, including honeyeaters and small mammals such as rodents, antechinus, honey possums, pygmy possums, gliders and bats. Many of these animals play a role in pollination of Banksia. Various studies have shown mammals and birds to be important pollinators. In 1978 Carpenter observed that some banksias had a