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An example of Secondary Succession by stages:
1. A stable deciduous forest community
2. A disturbance, such as a wild fire, destroys the forest
3. The fire burns the forest to the ground
4. The fire leaves behind empty, but not destroyed, soil
5. Grasses and other herbaceous plants grow back first
6. Small bushes and trees begin to colonize the area
7. Fast growing evergreen trees develop to their fullest, while shade-tolerant trees develop in the
understory
8. The short-lived and shade intolerant evergreen trees die as the larger deciduous trees overtop
them. The ecosystem is now back to a similar state to where it began
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Process
Ecological succession is the long-term process in which plant species
replace others in a vegetative community over time, changing both
vegetation composition and structure. Vegetation types are dynamic;
competition and disturbance cause continual changes in vegetation types
and dominant species. The pace of succession can be rapid open
grassland can become a solid stand of coyote brush within five years; live
oak seedlings can create dense thickets within 15 - 20 years. The full
dynamic range of ecological succession plays out over decades and even
centuries.
New species and thus succession can be introduced via interfaces
between different vegetation types. Other introductions can result from
dispersal that allows plant species to become established far from seed
sources thanks to transportation by animals (e.g. oaks) or wind (e.g.
coyote brush). Climate change, nitrogen deposition, weed invasions,
disease, and other site conditions can all alter natural successional
pathways.
Some successional series produce simplified vegetation structure; for
example, the steady loss of oaks in woodlands over time leads to
conversion to annual grassland. More typically, however, ecological
succession is accompanied by higher biomass and greater complexity,
exemplified by the grassland-shrubland-woodland-forest transition. The
dominant climax species present in the final stages of succession and
maximum biomass at a site is ultimately determined by water balance and
soil fertility.
Distribution
Natural and anthropogenic disturbances have shaped the Bay Area
landscape and distribution of vegetation for thousands of years. Extensive
burning and localized digging by California Native Americans (Anderson
2005) was followed by timber harvesting, fires, grazing, land clearing,
plowing, and urbanization after the arrival of Europeans. These activities
have left a mosaic of different-aged stands of vegetation interspersed with
permanently disturbed urban and agricultural land.
Extensive reviews for succession in many vegetation types can be found in
Barbour et al. 2007 and Sawyer et al. 2009. Some key successional
dynamics of concern to the Conservation Lands Network include:
1. In the absence of grazing and/or fire, open grasslands convert to
shrublands, especially dense coyote brush. This process occurs
throughout the coastal belt (including the Berkeley Hills) and has led to
the loss of rich coastal prairies represented by Cool and Moderate
Grasslands. Coyote brush can die back in some areas, allowing the
return of open grassland, but most of the succession is unidirectional.
2. Open grasslands convert to conifer forest, especially Douglas-fir and
pines. This process is most frequent in the coastal zone, but can occur
in grasslands near Douglas-fir stands in inland locations.
3. Shrublands convert to woodland and forest as trees overtop the shrubs.
For example, manzanita stands on Mt. Tamalpais are gradually
converting to Douglas-fir forest.
4. Oak woodland and montane hardwoods are invaded by Douglas-fir, and
is occurring in many coastal areas. In Sonoma County, Pepperwood
Preserve and Annadel State Park are both experiencing a conversion to
Douglas-fir forests.
5. Open oak woodlands become denser and convert to closed canopy
woodlands, resulting in the loss of understory grassland.
6. Many oak woodlands and montane hardwood forest will eventually be
dominated by bay laurel as a natural process of succession. Sudden Oak
Death can accelerate this transition because bay trees harbor the
pathogen that kills live oaks and tanoaks.
7. The senescence of mature oaks and the lack of recruitment leads to the
conversion of oak woodlands to open grassland.
Monitoring
Monitoring succession is an essential part of a management program. Field
observations can provide early detection of the establishment of transition
species as seedlings and young plants. Ground-based and aerial photo
monitoring and satellite remote sensing are invaluable for documenting
long-term changes. In small habitat areas of reference sites, repeat
sampling of permanent plots can track fine-scale successional dynamics,
but in broader landscapes, extensive surveys are necessary. Posttreatment monitoring is a critical part of adaptive management plans
(Elzinga et al. 1998).
Conclusions, Management
Recommendations, and Research Needed
The dynamics of vegetation succession pose ongoing management issues
across the Conservation Lands Network. Some key conclusions and
recommendations to keep in mind are:
1. The landscape at present is a mosaic of different successional stages
that reflect historic and prehistoric disturbance regimes.