Succession

Primary successions occur after disturbances that have been intense enough to obliterate any living organisms from the site, and even to have wiped out all traces of previous biological influences, such as soil development. Natural disturbances of this intensity are associated with glaciation, lava flows, and very severe wildfires, while human-caused examples might include the abandonment of a paved parking lot, or an above-ground explosion of a nuclear weapon. In all of these cases, organisms must successfully invade the disturbed site for succession to begin.

A well-known study of primary succession after deglaciation has been conducted in Alaska. This research examined a time series of substrates and plant communities of known age (this is called a chronosequence) at various places along a fiord called Glacier Bay. In this study, historical positions of the glacial front could be dated by various means, including written records and old photographs. The primary succession begins as the first plants colonize newly deglaciated sites and, in combination with climatic influences, begin to modify the post-glacial substrate. These initial plants include mosses, lichens, herbaceous dicotyledonous species such as the river-beauty (Epilobium latifolium), and a nitrogen-fixing cushion plant known as mountain avens (Dryas octopetala). These pioneers are progressively replaced as succession continues, first by short statured species of willow (Salix spp.), then by taller shrubs such as alder (Alnus crispa, another nitrogen-fixing plant), which dominates the community for about 50 years. The alder is progressively replaced by sitka spruce (Picea sitchensis), which in turn is succeeded by a forest of western hemlock (Tsuga heterophylla) and mountain hemlock (T. mertensiana). The primary succession at Glacier Bay culminates in the hemlock forest—this is the climax stage, dominated by species that are most tolerant of stresses associated with competition, in a mature habitat in which access to resources is almost fully allocated among the dominant biota.

The plant succession at Glacier Bay is accompanied by other ecological changes, such as soil development. The mineral substrate that is initially available for plant colonization after the meltback of glacial ice is a fine till, with a slightly alkaline pH of about 8, and as much as 7-10% carbonate minerals. As this primary substrate is progressively leached by percolating water and modified by developing vegetation, its acidity increases, reaching pH 5 after about 70 years under a spruce forest, and eventually stabilizing at about pH 4.7 under a mature hemlock forest. The acidification is accompanied by large reductions of calcium concentration, from initial values as large as 10%, to less than 1%, because of leaching and uptake by vegetation. Other important soil changes include large accumulations of organic matter and nitrogen, due to biological fixations of atmospheric carbon dioxide (CO₂) and dinitrogen (N₂).

Other studies of primary succession have examined chronosequences on sand dunes of known age. At certain places on the Great Lakes, sandy beach ridges are being slowly uplifted from beneath the lakewaters by a ponderous rebounding of the land. The process of uplifting is called isostasy, and it is still occurring in response to meltback of the enormously heavy glaciers that were present only 10,000 years ago. The initial plant colonists of newly exposed dunes that were studied on Lakes Michigan and Huron are short-lived species, such as sea rocket (Cakile edentula) and beach spurge (Euphorbia polygonifolia). These ephemeral plants are quickly replaced by a perennial dunegrass community, dominated by several grasses (Ammophila breviligulata and Calamovilfa longifolia). With time, a tall-grass prairie develops, dominated by other species of tall grasses and by perennial, dicotyledonous herbs. The prairie is invaded by shade-intolerant species of shrubs and trees, which form nuclei of forest. Eventually a climax forest develops, dominated by several species of oaks (Quercus spp.) and tulip-tree (Liriodendron tulipifera). This series of successional plant communities is accompanied by soil development, with broad characteristics similar to those observed at Glacier Bay (although, of course, the rates of change are different).