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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.

Ecological Impacts and Threats

Succession is neither good nor bad; its ecological impact depends on
context and the resources at risk. Succession eventually leads to local
losses and even severe reductions of some species, and can affect the
composition of entire landscapes. For example, succession of coastal
grasslands into native shrubs and trees is a natural process, but if the
grassland supports endangered species or a particularly rich native flora,
it may compromise biodiversity conservation goals.
In the Bay Area, where many rare species are grassland obligates, the loss
of open grassland habitats and associated species can have significant
impacts on biodiversity. Native brush succession on San Bruno Mountain,
for example, has emerged as a major threat to the rich coastal grasslands
and associated endangered species (TRA Environmental Sciences 2007).
The invasion of oak woodlands by Douglas-fir creates dense thickets of
Douglas-fir that crowd the forest floor, eventually over-topping the oaks,
leading to losses of oaks and understory shrubs and herbs. After several
decades of dense canopy closure, natural mortality thins out the Douglasfir trees, letting more light reach the forest floor and creating a complex
multi-story forest.
The biomass accumulation that typically accompanies ecological
succession can lead to increased fire risks a serious problem at the
urban-wildland interface where shrub cover can increase potential fire
intensity and spread rates.

Network Design and Management

Responses
Many successional changes are inevitable, and it is not possible or
desirable to manage succession across the entire Conservation Lands
Network. Selected stewardship actions are necessary to maintain
biodiversity in a dynamic landscape.
Successional vegetation transitions require careful consideration in
conservation planning. In the design phase, the Upland Habitat Goals
Project strategy of capturing large areas of extant vegetation provides
some buffer against short-term change.
On existing or potential conservation lands, local land managers should
inventory key resources likely to be affected by succession. The basic
questions to ask include:
1. Is there on-the-ground evidence of incipient or ongoing succession?
2. What might this landscape look like in 10, 20, or 30+ years if these
trends continue with no intervention?
Can rates of succession be inferred from historical data such as aerial
photographs and surveys such as the Wieslander maps?
4. Can feasible management actions help achieve conservation goals?
Managing succession requires clear local goals for maintenance of
vegetation types, combined with adequate resources to implement
management actions such as grazing, mechanical removal, chemical
application, or prescribed burning. Management actions need to be
carefully planned and documented in an adaptive management process,
so that effectiveness can be evaluated and appropriate adjustments made
through time.
Early treatment of invasive species (native and non-native) is essential to
maintain open early successional habitats. Once certain shrubs (e.g.,

coyote brush) become established in grasslands, they may prove difficult

to remove because of resprouting. In the case of Douglas-fir invasion of
oak woodlands, treatment of trees is best done in the sapling or pole stage
the larger the trees to be removed, the more expensive and disruptive
the management treatments. For more information on non-native
invasives, see the Invasive Plants Viability Summary.
Oak recruitment need occur only intermittently to sustain populations, and
individual trees can live for a century or more. In some cases, small
established oaks may get browsed down each year; protection from
browsing will allow them to grow to a larger, less-vulnerable size. Loss of
oaks can be mitigated by planting, as well as by management to
encourage local recruitment.

Policy and Institutional Responses

Recognition of the dynamic nature of vegetation is central to developing
policies for succession management. On many conservation lands, a
hands-off approach is the default policy because of assumptions about
what is natural and a lack of stewardship resources. In extensive
wildlands, doing nothing may be appropriate. In some areas, natural fires
may provide the successional mosaic desired for biodiversity. However,
over time, such hands-off approaches may lead to undesirable outcomes,
especially where locally-rare conservation targets are at risk or where fire
is not an option. In smaller habitats or urban settings, policies that support
management practices such as grazing, mowing, or planting can be
essential for biodiversity conservation.
Education of both the land management community and the general
public can improve support for hands-on management of natural lands; in
addition, funding for ongoing stewardship is critical. Policies regarding the
use of fire and other treatments for succession management can be
unwieldy and delay, stop, or increase costs of management. Streamlining
the environmental review process and developing programmatic
Environmental Impact Reports can reduce these barriers.

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.

2. Succession will continue in the absence of disturbance, and may result

in desirable or undesirable changes to vegetation and impacts on
species.
3. Explicit decisions about succession are usually necessary to meet
resource management goals.
4. Anticipation of undesirable changes and rapid response to incipient
successional dynamics are always easier than trying to reverse
succession once it is well along.
5. Compilation of vegetation type conversions since the 1930s through
detailed analysis of the Wieslander vegetation maps could provide
some baselines on rates of change.
6. Succession is affected by many of the other viability factors. The
interactions among these factors should be evaluated and elucidated.
7. Climate change, in particular, will alter succession because aridity will increase and
directly limit many species. Increased fire frequency and intensity will accelerate type
conversions. add another sentence: Better locally calibrated climate/fire/succession
models will enhance the understanding of the potential rates of conversion under
climate change scenarios.