Abstract
Investigators
Introduction
Methods
Results
Conclusions
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Landscape-scale patterns of grassland, shrubland, and woodland plant assemblages in relation to environment and disturbance variables, Applegate watershed, SW Oregon

Multivariate techniques were used to investigate how environmental and disturbance factors affect the distribution and species composition of grassland, shrubland, and woodland plant assemblages at the landscape level.  Thirteen plant assemblages were defined and modeled across the landscape based on species composition from 425 stands.  The use of classification, ordination, and indicator species analysis produced relatively discrete and ecologically representative plant assemblages. Although variables related to moisture and soils were most important to plant distribution, vegetation patterns appear to be determined by a complex combination of edaphic, topographic, and climatic variables in addition to site history. Differences between assemblages provide a benchmark for better understanding variation between grassland, shrubland and woodland vegetation types in space and time.

Eric Pfaff and Paul Hosten
Southern Oregon University and Bureau of Land Management, Medford District, respectively

Grassland, shrubland, and woodland vegetation types are a major component of the landscape in the Applegate Valley of southwest Oregon.  The Ashland Resource Area of the Medford Bureau of Land Management is currently implementing thousands of hectares of fuels reduction treatments annually. However, there is a paucity of data regarding these vegetation types and how they are affected by fuel reduction treatments.  The objectives of this study were to gain a better understanding of these vegetation types and the factors responsible for their distribution.

Right: Study Area Map (larger view). Study area shown in red within the Klamath Mountains of southwest Oregon. The study area lies within the Applegate Valley of southwestern Oregon on U.S. Department of Interior (USDI) Bureau of Land Management (BLM) lands in the Ashland Resource Area.

Map of study area

The first author collected data on species composition, woody plant structure (size classes and form), and site characteristics for 425 non-conifer stands using a reconnaissance or releve style methodology. Thirteen plant assemblages based on species composition were classified using clustering techniques in conjunction with Indicator Species Analysis.  NMS ordination was used to investigate patterns of species distribution within assemblages. Nineteen environmental and ten disturbance variables were regressed against ordination axes and used to model assemblages based on landscape characteristics. PC-ORD software was used in analysis.

ASSOCIATION NAME

TYPICAL SPECIES (common names)

PHYSIOGNOMIC GROUP

Fescue grassland

Fescue/St. Johns wort/ balsamroot

Grassland/bald

Buckbrush chaparral

Buckbrush/soft brome

Shrubland

Rabbitbrush/buckwheat bald

Rabbitbrush/arrowleaf and naked buckwheat

Grassland/bald

Manzanita/white oak

Whiteleaf manzanita/Oregon white oak

Woodland/shrubland

Oak woodland

Oregon white oak/Chinook brome/hedgehog dogtail/ spreading hedgeparsley

Woodland

Oat grass grassland

Oat grass/ medusa head/ Lemmon's needlegrass/

California poppy/lupine

Grassland/bald

Mountain mahogany/

Brewer's oak

Mountain mahogany/Brewer's oak/serviceberry/seablush/ yarrow/lambstongue ragwort

Shrubland

Open oak/chaparral

Oregon white oak/grand collomia/California brome/ hedgehog dogtail

Woodland/shrubland

Black oak woodland

California black oak/blue wildrye/yellowleaf iris/beaked hazelnut

Woodland

Disturbance mediated woodland/chaparral

Madrone/white oak/buckbrush/ manzanita/cheatgrass

/madia

Woodland/shrubland

Madrone woodland

Madrone/black oak/deerbrush/ blacksnakeroot

Woodland

Manzanita chaparral

Whiteleaf manzanita/ponderosa pine/white oak/blue dicks

Shrubland

Canyon live oak woodland

Canyon live oak/Douglas-fir/ bearbrush/Oregon grape/ honeysuckle

Woodland

The figure below is an ordination based on the species composition of the 425 surveyed stands. Symbols represent stands, and different symbols represent different assemblages. The farther the distance between symbols, the more dissimilar the stands in terms of species composition. Pictures are from a representative stand of each assemblage (hover the mouse over the picture to see the assemblage name).  

Below: Summary of Plant Assemblages. Images of each of these assemblages follow. Also see the complete plant species list compiled for all stands.

Ordination of plant assemblages
Open oak/chapparal
Mountain mahogany/Brewers oak
Fescue grassland
Rabbitbrush/Buckwheat bald
Oat grass grassland
Buckbrush chaparral
Disturbance mediated woodland/chaparral
Black oak woodland
Manzanita chaparral
Manzanita/White oak
Canyon live oak woodland
Oak woodland
Madrone woodland

Number of significant indicator species for each assemblage from indicator species analysis.

Average P values for each cluster analysis group from indicator species analysis.

Cluster Analysis, coupled with Indicator Species Analysis (ISA), clustered stands into thirteen groups (plant assemblages); this grouping had both a high number of significant indicator species and low average P values. This indicates a high degree of fidelity of indicator species to stand clusters relative to other numbers of groups.

Most of the assemblages were limited to exposed, southerly aspect sites with
shallow, rocky, fast draining, loam soils. Some of the assemblages were fairly
restricted by environmental conditions, while others were found in a number of ecological sites defined by topographic and edaphic conditions.

Disturbance appears responsible for increasing species composition variability within assemblages.  Site data indicated that many of the outlying stands in each assemblage had evidence of relatively recent wildfire or fuel  treatment.  Some assemblages were clearly delineated in ordination space, while other assemblages with more common species and/or habitats tended to overlap considerably.

Correlations (r-squared) of predictor variables with ordination axis

Variable

Axis 1

Axis 2

Axis 3

Elevation

0.004

0.478

0.08

Summer actual evapotranspiration

0.001

0.302

0.081

pH

0.054

0.285

0

Annual precipitation

0

0.251

0.108

Total silt

0.076

0.237

0

Upper soil horizon depth

0.08

0.235

0.004

Total clay

0.062

0.23

0

Annual actual evapotranspiration

0.01

0.181

0.044

Total sand

0.071

0.168

0.016

Cation exchange capacity (CEC)

0.096

0.082

0.013

Fire frequency

0.035

0.008

0.003

Fire interval

0.017

0.005

0.027

Slope Shape (e.g. concave)

0

0.002

0.001

Years since last fire

0.055

0

0.008

Heatload

0.055

0

0.006

Right: Variables associated with moisture
(elevation, summer evapotranspiration, and
annual precipitation) are related to axis 2. Soil
variables (upper soil horizon depth, pH,
cation exchange capacity and total silt, sand,
and clay) are also mostly associated with
axis 2.  Heatload and years since fire were less
important, but were weakly related to axis 1. 

Below: Ordination based on the species
composition of the surveyed stands, as above,
but with an overlay of environmental and
disturbance characterstics measured for each
stand that help predict species distributions.
The lines show associations of
variables with ordination axes. Each line
represents an environmental variable (predictor
variable); lines point toward stands with
relatively high values for each variable, and line
lengths are proportional to the amount of
influence of the variable on species
composition. (Line labels are clarified below.)

Sites at the top of the diagram
(grasslands) have higher elevation, higher
summer evapotranspiration, and higher annual precipitation, with soil that is typically less developed, has a lower pH, lower nutrients, and exhibits soil catena.  Sites are the bottom (chaparral) have lower moisture with soils that are more developed and have a more moderate pH. Heatload (aspect and slope) and associated fire history appear to distribute plants from more open chaparral and grassland sites on the left to higher canopy woodland sites on the right in the diagram; however, low explanation values (r2, as shown in Table above) suggest that other factors are also responsible.   

Line labels, clockwise from top, are elevation, summer evapotranspiration, annual precipitation, years since fire, soil Cation Exchange Capacity, soil %Sand, soil %Clay, soil %Silt, soil pH, heatload, and upper soil horizon depth. Only lines for those variables that most correlate with species distribution are shown.

Ordination of stands with predictor variable overlay

Below: Mean percent cover by native/non-native functional group in the herbaceous layer for each assemblage. Periwinkle = Native Perennial Grass, Maroon = Exotic Annual Grass, Light Yellow = Native Perennial Forb, Light Blue = Native Annual Forb, and Dark Purple = Exotic Annual Forb. Assemblage labels are, from left to right: Fescue grassland, Buckbrush chaparral, Rabbitbrush/buckwheat bald, Manzanita/white oak, Oak woodland, Oat grass grassland, Mountain mahogany/Brewer's oak, Open oak/chaparral, Black oak woodland, Disturbance mediated woodland/chaparral, Madrone woodland, Manzanita chaparral, and Canyon live oak woodland.

Below: Mean percentage of oak size classes for each assemblage Sizes are diameter at breast height (DBH). Light Blue = DBH Greater than 16", Light Yellow = DBH 9-16", Maroon = DBH 4-9", Periwinkle = DBH Less than 1". Assemblage labels are, from left to right: Fescue grassland, Buckbrush chaparral, Rabbitbrush/buckwheat bald, Manzanita/white oak, Oak woodland, Oat grass grassland, Mountain mahogany/Brewer's oak, Open oak/chaparral, Black oak woodland, Disturbance mediated woodland/chaparral, Madrone woodland, Manzanita chaparral, and Canyon live oak woodland.

Mean percentage of oak size classes in each assemblage

Below: Mean stand species richness in each assemblage. Assemblage labels are, from left to right: Fescue grassland, Buckbrush chaparral, Rabbitbrush/buckwheat bald, Manzanita/white oak, Oak woodland, Oat grass grassland, Mountain mahogany/Brewer's oak, Open oak/chaparral, Black oak woodland, Disturbance mediated woodland/chaparral, Madrone woodland, Manzanita chaparral, and Canyon live oak woodland.

Species richness in each assemblage

Meadow death camas (Zigadenus venenosus). Photo by K. Sikes.

  • Variables related to moisture (elevation, summer evapotranspiration, and annual precipitation) and soils (pH, soil horizon depth, and total silt and clay) were found to be the most important determinants of non-coniferous vegetation distribution.
  • Disturbance (wildfires and fuel reduction treatments) appears
    responsible for increasing the variability of species
    composition within assemblages.
  • The high proportion of exotic annual grasses in the
    disturbance mediated woodland/chaparral assemblage
    suggests that disturbance may increase the proportion of
    exotic taxa.
  • Non-native taxa are mostly in the form of exotic annual
    grasses found typically in low elevation, low canopy situations.
  • Regression results indicate that heatload (aspect and slope)
    and fire history are associated, yet neither are highly
    correlated to plant distribution.
  • Classification and Indicator Species Analysis produced
    ecologically representative assemblages, and environmental
    variables elucidated factors predictive for patterns of species
    distribution within assemblages. An exception to the
    classification working well is the fescue grassland
    assemblage, which had a high elevation, mostly native
    component and a low elevation, mostly naturalized
    (non-native) component.
  • Predictive habitat models of assemblages based on ordination scores produced mixed results.  This is possibly due to assemblages occupying multiple ecological sites, individual species responding independently to predictor variables, inaccuracies in base data, limited site history information, unmeasured biotic processes, and variation within stands and associations. 
  • Oak stands are present as mixed size classes. Typically oaks belong to the small size classes with fewer large trees.
  • The rarity, high species richness of native plants, and the potential for the extirpation of the fescue grassland and black oak woodland plant assemblages warrant efforts to ensure their persistence on the landscape.
Zigadenus venenosus

Link to E. Pfaff thesis (PDF).
Contact E. Pfaff or P. Hosten.


Photos were taken by E. Pfaff, unless otherwise noted.

Page by Olivia Duren. Updated 9/2007.

 

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Results from Indicator Species Analysis
Proportion functional groups in each assemblage