Turner Review No. 2 Southern Conifers in Time and Space

The three southern conifer families, Araucariaceae, Cupressaceae and Podocarpaceae, have a long history and continue to be an important part of the vegetation today. The Araucariaceae have the most extensive fossil record, occurring in both hemispheres, and with Araucaria in particular having an ancient origin. In the Southern Hemisphere Araucaria and Agathis have substantial macrofossil records, especially in Australasia, and Wollemia probably also has an important macrofossil record. At least one extinct genus of Araucariaceae is present as a macrofossil during the Cenozoic. Cupressaceae macrofossils are difficult to identify in older sediments, but the southern genera begin their record in the Cretaceous (Athrotaxis) and become more diverse and extensive during the Cenozoic. Several extinct genera of Cupressaceae also occur in Cretaceous and Cenozoic sediments in Australasia. The Podocarpaceae probably begin their macrofossil record in the Triassic, although the early history is still uncertain. Occasional Podocarpaceae macrofossils have been recorded in the Northern Hemisphere, but they are essentially a southern family. The Cenozoic macrofossil record of the Podocarpaceae is extensive, especially in south-eastern Australia, where the majority of the extant genera have been recorded. Some extinct genera have also been reported from across high southern latitudes, confirming an extremely diverse and widespread suite of Podocarpaceae during the Cenozoic in the region. In the Southern Hemisphere today conifers achieve greatest abundance in wet forests. Those which compete successfully with broad-leaved angiosperms in warmer forests produce broad, flat photosynthetic shoots. In the Araucariaceae this is achieved by the planation of multiveined leaves into large compound shoots. In the other two families leaves are now limited to a single vein (except Nageia), and to overcome this limitation many genera have resorted to re-orientation of leaves and twodimensional flattening of shoots. The Podocarpaceae show greatest development of this strategy with 11 of 19 genera producing shoots analogous to compound leaves. The concentration of conifers in wet forest left them vulnerable to the climate change which occurred in the Cenozoic, and decreases in diversity have occurred since the Paleogene in all regions where fossil records are available. Information about the history of the dry forest conifers is extremely limited because of a lack of fossilisation in such environments. The southern conifers, past and present, demonstrate an ability to compete effectively with angiosperms in many habitats and should not be viewed as remnants which are ineffectual against angiosperm competitors.