CORRELATION
BETWEEN HABITAT CHARACTERISTICS
IMPLICATIONS
FOR RESTORATION
***
Catherine
S. Linden
Wildland/Range
Science
ABSTRACT
This study took place in
spring 1999, in
INTRODUCTION
Currently less than 5% of
Field testing of different seeding
methods at the Ranch has produced a successful method of seeding Nassella pulchra (Needlegrass),
which is commonly thought to have been the dominant native perennial grass of
the area. Previous studies in
restoration of native grasses in
The study was conducted at
the Gray Davis Dye Creek Preserve (the Ranch) in
The Ranch is located on an
ecological subregion of
The soils in the study area consist of three associations
with eight phases represented (USDA, 1967).
The Tuscan-Inks association includes the Tuscan, Inks, and Keefer soil
phases. The association is characterized
by nearly level to steep, cobbly soils that are
shallow to moderately deep to hardpan.
The soil was formed in stratified deposits of alluvium derived from
volcanic rocks of andesite, basalt and rhyolite. The Toomes-Guenoc association includes the Toomes
soil phase. This association is
characterized by shallow or moderately deep, rocky, gently sloping to steep
soils underlain by volcanic rock. The
underlying rock in the Toomes-Guenoc association is
predominantly andesite with inclusions of
basalt. The Columbia-Vina
association includes the Berendos soil phase. The association is characterized by very
deep, nearly level, moderately fine textured to moderately coarse textured
soils on flood plains of the
The agricultural history of the
study area is long and varied (Stechman, 1996). The area was part of a land grant received by
Job Dye in 1845. Cattle were grazed on
the land from 1845 to 1868. From 1868 to
1884, new owner Joseph Cone grazed sheep and dryland
farmed wheat on the acreage. In 1906,
Cone switched to raising cattle.
Ownership changed hands many times until in 1940, Stover and Conrad
bought the property and it was given the name Dye Creek Ranch. Recent historical records show that for 35
years 2,350 cattle per year have been grazing the Ranch property. Presently cattle are grazed in the study area
in the month of November and the month of May.
This year 1720 acres in the study area were not grazed at all due to a
prescribed burn scheduled for June.
During the spring and early summer, 1999, a survey was taken on 8,000 acres of the Ranch to determine the areas that contained remnant populations of native perennial grasses. Before each pasture was surveyed, a review of the Tehama County Soil Survey was done to determine what soils were in the pasture and where changes in soil phase occurred. Once in the field and a relict stand was found, a photograph was taken of the stand including the site in which it was found. All data regarding site characteristics were noted in a field journal. Data were recorded on the following characteristics: species, soil phase (taken from Tehama County Soil Survey), topography, elevation, amount of shade (full sun, partial shade, full shade, canyon shadow), percent slope, aspect, associated species, number of plants, and size of stand. Species found on the preserve in numbers large enough for which data analysis was possible are Nassella cernua and Nassella pulchra (Needlegrass), Melica californica (Melic grass), Muhlenbergia rigens (Deer grass) and Elymus glaucus (Blue Wild Rye). There were not enough sightings of two other species (Poa secunda and Aristida hamulosa) to perform data analysis. Data for Nassella cernua and Nassella pulchra were pooled as Nassella.
Indirect gradient analysis, or
vegetation ordination, is the technique commonly employed to order vegetation
data (Kent & Coker, 1992). This
technique is employed independently of environmental data. Once the vegetation data have been described
and summarized the environmental data are compared and summarized. Thus the environmental interpretation is
indirect. The three environmental
variables compared and summarized in this analysis were elevation, shade, and
slope. The results of the indirect
gradient analysis were used to run a principal component analysis to calculate
a correlation matrix between all species and variables. The correlation matrix was collapsed into two
major trends or components. Pearson
correlations were calculated between each of the three environmental variables
and the two principal components.
Proportional distributions were constructed for each native grass
species across levels of each of the three classificatory variables (nominal
or, without numerical value or rank) (soil phase, topographic position, and
associated species) and the ordinal variable (shade). A plot was prepared of each species’ aspect
distribution and the median for the species’ distribution was calculated
(Holmes, 1999).
Figure 1 shows the results of the principal component analysis. This is a biplot of
the data on the first two principal components, each component being a weighted
combination of variables.
Results of the species ordination analysis were not
especially strong as the first two components only account for 63% of the total
variation in the data. Elymus and Nassella are separated by
the first component and Muhlenbergia
and Melica
are separated by the second component.
This suggests that species distributions are dissimilar for each of
these two pairs (Holmes, 1999).

Figure 1. Biplot from principal component analysis of Dye Creek data. Species are represented by arrow,
numbers represent observations (Holmes, 1999).
Table 1 provides Pearson
correlations between each environmental variable and each principal
component. None of these correlations is
large, although it does appear that a weak negative correlation exists between
slope and the second principal component that separates Muhlenbergia and Melica. Melica
occurred on steeper slopes than Muhlenbergia.
Table 1. Pearson correlations
between each environmental variable and each principal component (Holmes,
1999).
|
|
Elevation |
Shade |
Slope |
|
1st Component |
-0.19 |
0.21 |
-0.07 |
|
2nd Component |
-0.15 |
0.19 |
-0.41 |
Figure 2 provides aspect distribution of relict sites by
species. Nassella and Melica
occur on all aspects, while Elymus
and Muhlenbergia
are sites with southwest aspects. Median
aspects are 236° for Nassella,
195° for Melica,
210° for Elymus,
and 210° for Muhlenbergia. Thus, the centers of their aspect
distributions all tend to face southwest.

Table 2 shows the species distribution for each of the three
classificatory variables, plus shade.
Marked deviations from the median distribution are highlighted. Review of the results of the analysis shows
that all species preferred the Toomes very rocky
loam, 10 to 30% slope soil phase (TgD) except for Muhlenbergia, which
prefers Keefers complex, channeled, 0 to 3 &
slope soil phase (Kn). Of the four species, all had distinct
preference for topographic position except Nassella, which is equally distributed on four
topographic positions. Nassella and Muhlenbergia prefer full
sun whereas Melica prefers partial oak shade and Elymus prefers full shade. Nassella
is commonly found with wild oats & YST. Melica
is most frequently found with short annuals, Elymus with no associated species, and Muhlenbergia is associated
with Cynodon dactylon (Bermuda grass) and Lolium multiflorum (annual rye
grass).
Table 2. Species distribution on the three classificatory variables, plus shade. Numbers indicate the percent that each species was found with a particular variable. Marked deviations from the median distribution are highlighted (Holmes, 1999).
Soil phases are: TgD, Toomes very rocky loam, 10 to 30% slope; IcD, Inks cobbly loam, 3 to 30% slope; TxC, Tuscan very stony loam, 3 to 15% slope; Mr, Millrace complex, channeled, 0 to 3% slope; Bg, Berrendos clay loam, 0 to 3% slope; Rr, Riverwash; Kn, Keefers complex, channeled, 0 to 3% slope; and TuB, Tuscan cobbly loam, 1 to 5% slope.
Associated species: 1&3, Short Annuals in addition to Oats & YST.
|
|
Soil
Phase |
|
|
|
|
|
|
|
|
|
|
|
|
TgD |
IcD |
TxC |
Mr |
Bg |
Rr |
Kn |
TuB |
|
|
|
Nassella |
0.70 |
0.20 |
0.02 |
0.01 |
0.02 |
0.00 |
0.01 |
0.02 |
|
|
|
Melica |
0.85 |
0.10 |
0.00 |
0.05 |
0.00 |
0.00 |
0.00 |
0.00 |
|
|
|
Elymus |
0.78 |
0.11 |
0.00 |
0.11 |
0.00 |
0.00 |
0.00 |
0.00 |
|
|
|
Muhlenbergia |
0.31 |
0.00 |
0.00 |
0.08 |
0.00 |
0.08 |
0.54 |
0.00 |
|
|
|
|
Median |
0.74 |
0.11 |
0.00 |
0.06 |
0.00 |
0.00 |
0.01 |
0.00 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
Topographic
Position |
|
|
|
|
|
|
|
|
||
|
|
SeasCrkBank |
Roadside |
CrkBot |
RockOut |
Draw |
Seep |
RiparBank |
RockLawn |
Alluvium |
OakSavan |
|
Nassella |
0.28 |
0.02 |
0.01 |
0.21 |
0.06 |
0.01 |
0.00 |
0.20 |
0.21 |
0.01 |
|
Melica |
0.60 |
0.00 |
0.07 |
0.19 |
0.02 |
0.00 |
0.02 |
0.02 |
0.07 |
0.00 |
|
Elymus |
0.00 |
0.00 |
0.78 |
0.00 |
0.00 |
0.00 |
0.11 |
0.00 |
0.11 |
0.00 |
|
Muhlenbergia |
0.00 |
0.00 |
0.23 |
0.00 |
0.00 |
0.00 |
0.15 |
0.00 |
0.62 |
0.00 |
|
|
Median |
0.14 |
0.00 |
0.15 |
0.10 |
0.01 |
0.00 |
0.07 |
0.01 |
0.16 |
0.00 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
Shade |
|
|
|
|
|
|
|
|
|
|
|
|
Oak-Full |
Oak-Part |
Full Sun |
CanyonShadow |
|
|
|
|
|
|
|
Nassella |
0.08 |
0.36 |
0.58 |
0.00 |
|
|
|
|
|
|
|
Melica |
0.08 |
0.56 |
0.36 |
0.00 |
|
|
|
|
|
|
|
Elymus |
0.67 |
0.33 |
0.00 |
0.00 |
|
|
|
|
|
|
|
Muhlenbergia |
0.08 |
0.08 |
0.69 |
0.15 |
|
|
|
|
|
|
|
|
Median |
0.08 |
0.35 |
0.46 |
0.00 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
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|
|
|
Associated
Species |
|
|
|
|
|
|
|
|
||
|
|
Shrt Ann |
BrushAnn |
OatsYST |
None |
Buckwheat |
1 & 3 |
SedgeRush |
Clover |
Medusa |
|
Lolium |
Nassella |
0.25 |
0.00 |
0.33 |
0.02 |
0.05 |
0.14 |
0.00 |
0.05 |
0.14 |
0.00 |
0.03 |
Melica |
0.45 |
0.02 |
0.25 |
0.02 |
0.05 |
0.16 |
0.00 |
0.05 |
0.00 |
0.00 |
0.00 |
Elymus |
0.00 |
0.00 |
0.00 |
0.67 |
0.00 |
0.11 |
0.00 |
0.00 |
0.00 |
0.22 |
0.00 |
Muhlenbergia |
0.00 |
0.00 |
0.26 |
0.15 |
0.00 |
0.04 |
0.04 |
0.00 |
0.00 |
0.26 |
0.26 |
|
Median |
0.12 |
0.00 |
0.26 |
0.09 |
0.02 |
0.12 |
0.00 |
0.02 |
0.00 |
0.11 |
0.02 |
The data analysis of the four most
commonly found relict populations of native perennial grasses in the surveyed
area of the Ranch shows Nassella
to be a generalist. It grows in two soil
phases (TgD, IcD), four
topographic positions (seasonal creek bank, rock outcrop, rocky lawn, and
alluvium), in full sun and partial sun, in deeper soils with other deep rooted,
tall species such as wild oats & YST, or in shallower soils with a
combination of non-native short statured annuals
(Table 2). This conclusion is supported
by the principal component analysis (Figure 1).
A review of Figure 1 shows that the analysis of the first principal
component (which accounts for the most variance and describes the dominant
pattern among the species) is weighted positively by Nassella and negatively by the other species. This indicates that the distribution of Nassella across sites
contrasts with the distribution of the other species. Figure 2 shows that Nassella
can be found over a wide range of aspects, but the median aspect is 236°,
facing southwest.
Nassella prefers Toomes
very rocky loam, 10 to 30% slope (TgD)(70%); and Inks cobbly loam, 3 to
30% slope (IcD)(20%).
Both soils are well drained with low available water holding capacity
and low fertility. Permeability is
moderate and runoff is slow to medium.
The difference between the soils is mainly soil depth. The Toomes is 8-15
inches deep while the Inks is 10-24 inches deep. Nassella
prefers full sun (58%) but will tolerate partial shade (36%).
Melica is almost exclusively found on the Toomes very rocky loam, 10 to 30% slope (TgD) (85%). It prefers seasonal creek banks (60%), but may be found as often as Nassella on rock outcrops (19%). It prefers partial shade (56%), but tolerates full sun (36%), the opposite of Nassella. Melica is most commonly associated with short non-native annuals (45%), but tolerates the presence of wild oats & YST (25%).
The second principal component (Figure 1) shows that Melica and Muhlenbergia have different distributions in terms of slope. The Pearson correlation (Table 1) provides correlation between each environmental variable and each principal component. The negative correlation between slope and the second principal component suggests that sites that have lower slopes contained more Muhlenbergia, conversely, Melica occurs on steeper slopes. This information concurs with Melica’s 60% preference for seasonal creek banks. The median aspect for Melica was 195°, tending towards the southwest.
Data analysis indicates that Melica has similar habitat requirements to Nassella. Both are highly correlated with the Toomes very rocky loam, 10 to 30% slope, (Nassella, 70%, Melica, 85%). Both are correlated with seasonal creek banks
(Nassella,
28%, Melica,
60%) and rocky outcrops (Nassella,
21%, Melica
19%). Preference for sunlight is the
opposite, with Nassella
preferring full sun and tolerating partial shade (58% and 36%) while Melica prefers partial shade and tolerates full sun (56%
and 36%). They are both associated with
the same non-native annuals and forbs. Nassella is found with
short annuals (25%), wild oats & YST (33%), a combination of the two (14%),
and with Medusa head (14%) while Melica
is found with short annuals (45%), wild oats % YST (25%), and a combination of
the two (16%). Melica was not found
with Medusa head.
Elymus glaucus is highly
correlated with the Toomes very rocky loam, 10 to 30%
slope (TgD) (78%), creek bottoms (78%), full shade (67%)
to partial shade (33%). It occurs
primarily without associates (67%), or with Bermuda grass (22%). Elymus prefers a southwest facing aspect, the median being
210°.
Muhlenbergia is correlated with Keefers complex, channeled,
0 to 3% slope (Kn) (54%) and Toomes
very rocky loam, 10 to 30% slope (TgD) ( 31%). Muhlenbergia
is found on alluvium deposited in the channels (62%) and creek bottoms
(23%). It is highly correlated with full
sun (69%) but also occurs in canyon shadow (15%). Muhlenbergia occurs with wild oats & YST (26%), Bermuda grass (26%),
and Lolium multiflorum (26%) reflecting a
more mesic habitat.
The median aspect for Muhlenbergia
is 210°.
Data analyses indicate that Nassella is correlated with more habitat variables
than the three other species. The
ability to grow in various habitats coupled with the development of a
successful seeding method and high nutritive value (Wrysinski
et al., 1997) makes this an appropriate native perennial grass for restoration.
No seeding method has been developed for Melica californica. Since this species parallels many of Nassella’s habitat
requirements, Melica
may be a species of consideration for restoration work in the future. It is known to be nutritious and highly
palatable (Wrysinski et al., 1997; Stebbins, 1999).
Developing a seeding method for the highly specialized sites
that Elymus glaucus prefers in the studied
portion of the Ranch may be of consideration as well. Nutritive value for Elymus glaucus has been tested with positive results
and its value to wild life is documented (Wrysinski
et al., 1997; Strait, 1999).
Muhlenbergia rigens is
important to wild life and for stabilizing creek banks but grazers do not
utilize it. Therefore it would not be an
economically important species for restoration.
This study examined the correlation between habitat characteristics and the distribution of native perennial grasses. Any possibility that relict stands may be environmentally distinct from the rest of the landscape was not addressed by this study. Furthermore, this study reflects data taken from only a portion of the Ranch. When additional acreage is surveyed information regarding species/habitat correlations should be further refined.
REFERENCES:
Barbour, M.G., and W.D.
Billings. 1988. North American Terrestrial
Vegetation.
Bugg,
R.L.,
Holmes, T.H. 1999. Data analysis on bunchgrass
survey for Dye Creek Field Office.
Unpublished report. 5 pp.
Menke, J.W. 1992. Grazing and fire management for native perennial grass restoration
in
Stebbins, G.L. 1999. The genus Melica
in
Stechman, J.V.
1996. Unpublished
report. Rangeland conservation and management plan: The Nature
Conservancy, Gray Davis Dye Creek Preserve,
Strait, D. 1999. Native grasses
and their value for wildlife habitat enhancement. Grasslands.
Stromberg, M.R. and P. Kephart.
1996. Restoring native grasses in
Wrysinski,
J., P. Robins and G. Veserat. 1997. Native
grass forage quality pilot study. Yolo County Resource
Conservation District,