The hare, the tortoise and the crocodile: the ecology of angiosperm dominance, conifer persistence and fern filtering


David A. Coomes (tel. +44 1223 333911; fax: +44 1223 332418; e-mail:


  • 1Angiosperm trees often dominate forests growing in resource-rich habitats, whereas conifers are generally restricted to less productive habitats. It has been suggested that conifers may be displaced by angiosperms except where competition is less intense, because conifer seedlings are inherently slow growing, and are outpaced by faster-growing angiosperm species. Here we investigate whether competition with ferns and deeply shading trees also contributes to a failure of conifers to regenerate in resource-rich habitats.
  • 2We examined how changes in soil nutrient availability and drainage affected vegetation along the retrogressive stages of a soil chronosequence in southern New Zealand. Vegetation composition shifted from angiosperm-tree dominance on ‘recent’ alluvial terraces (< 24 ky), via coniferous-tree dominance on older marine terraces (79–121 ky), to coniferous-shrub dominance on the oldest marine terraces (291 ky). Soil drainage deteriorated along the sequence, and N : Pleaves and N : Psoil indicate increasing P-limitation. Conifer species appear to be adapted to persistence on infertile and poorly drained soils.
  • 3The floor of the relatively fertile alluvial forests was deeply shaded (∼1% light transmission) by dense groves of tree-ferns and ground-ferns, and by large-leaved subcanopy trees. Few seedlings of any type were found on the forest floor, even in tree-fall gaps, and establishment was restricted to rotting logs and tree-fern trunks. Angiosperms were particularly successful at colonizing these raised surfaces.
  • 4Less shade was cast by the conifer-dominated forests on infertile marine terraces (∼5% light transmission), which lacked tall ferns. There were many opportunities for conifer establishment, with high seedling densities recorded on the forest floor and on logs. By contrast, angiosperm seedlings were mainly restricted to logs.
  • 5Our results suggest that several mechanisms act in concert to reduce regeneration opportunities for conifers in productive habitats. In particular, we suggest that tall ferns and deep shade are responsible for a restriction of regeneration opportunities in relatively productive forests in New Zealand, diminishing the opportunities for conifers to escape the competitive effects of fast-growing angiosperms. Thus ‘crocodiles’ may alter the outcome of the race between ‘hares’ and ‘tortoises’.