Ungulate-Forest Relationships in Olympic National Park: Retrospective Exclosure Studies

Metadata:


Identification_Information:
Citation:
Citation_Information:
Originator:
Woodward, A. , Schreiner E. G., Houston, D.B., and Moorhead, B.B.
Publication_Date: 1994
Title:
Ungulate-Forest Relationships in Olympic National Park: Retrospective Exclosure Studies
Series_Information:
Series_Name: Northwest Science
Issue_Identification: Vol. 68, No. 2
Publication_Information:
Publication_Place: Pullman, Washingtom
Publisher:
Published by the Washington State University Press with collaboration of Northwest Scientific Association.
Description:
Abstract:
25 ungulate exclosures were established in Olympic National Park (23) and Olympic National Forest (2) in the 1930s and 1950s. We gleaned information from the early studies and, where possible resampled vegetation within and adjoining exclosures from 1987-1990 to describe effects of ungulates on forest vegetation. Exclosures were established on low elevation (< 350 m) ungulate ranges; 21 were located on river valley floors in the Sitka Spruce or Western Hemlock Forest Zones, 2 were on valley walls or other upland sites in the Western Hemlock and Silver Fir Zones, 2 were in clearcuts. We provide examples where ungulate understory, shrubs, lower tree canopy, and overstory canopy). In communities initially dominated by grasses or grasslike plants on valley floors, exclusion of ungulates resulted in decreased cover of grasses and forbs (usually), decreased species richness of forbs, and sometimes increased height and abundance of ferns. Shrub size and density increased following exclusion, particularly salmonberry and huckleberry. Ungulates influenced the recruitment and morphology of vine maple. Ungulate effects on establishment of overstory species were variable; browsing appeared to affect recruitment of Pacific silver fir and western red cedar following clearcut logging, but effects on other species were unclear. The importance of ungulate herbivory in determining forest composition is discussed in relation to the scale of other processes.
Purpose:
To evaluate the long-term effects of ungulates on the coniferous "rain" forest ecosystem unique to the Olympic Peninsula.
Supplemental_Information:
The geographic coordinates for this study are the coordinates of the Olympic Peninsula, estimated from as USGS 1:24000 map. For exact coordinates of the study sites, please contact the author.
Time_Period_of_Content:
Time_Period_Information:
Range_of_Dates/Times:
Beginning_Date: 1987
Ending_Date: 1990
Currentness_Reference: Observed
Status:
Progress: Complete
Maintenance_and_Update_Frequency: None Planned
Spatial_Domain:
Description_of_Geographic_Extent: Olympic National Park
Bounding_Coordinates:
West_Bounding_Coordinate: -124.7
East_Bounding_Coordinate: -122.6
North_Bounding_Coordinate: 48.4
South_Bounding_Coordinate: 46.9
Keywords:
Theme:
Theme_Keyword_Thesaurus: None
Theme_Keyword: ungulate
Theme_Keyword: Forest
Theme_Keyword: exclosure
Theme_Keyword: herbivory
Theme_Keyword: forest composition
Theme_Keyword: forest morphology
Theme_Keyword: standing crop of forest vegetation
Theme_Keyword: overstory species establishment
Place:
Place_Keyword_Thesaurus: None
Place_Keyword: usA
Place_Keyword: Washington
Place_Keyword: Olympic Peninsula
Stratum:
Stratum_Keyword_Thesaurus: None
Temporal:
Temporal_Keyword_Thesaurus: None
Access_Constraints:
Contact author for data. Reprints of article may be obtained from Northwest Science or the author.
Use_Constraints: Must cite author if using data in published work.
Point_of_Contact:
Contact_Information:
Contact_Person_Primary:
Contact_Person: Andrea Woodward
Contact_Organization: USGS
Contact_Address:
Address_Type: Mailing and Physical Address
Address: Western Fisheries Research Center,
Address: 6505 NE 65th St.
City: Seattle
State_or_Province: Wa
Postal_Code: 98115
Country: USA
Contact_Voice_Telephone: 206/526-6282 xt.332
Contact_Electronic_Mail_Address: andrea_woodward@usgs.gov
Security_Information:
Security_Classification_System: None
Security_Classification: Unclassified
Security_Handling_Description: None

Data_Quality_Information:
Lineage:
Methodology:
Methodology_Type: Field
Methodology_Identifier:
Methodology_Keyword_Thesaurus: None
Methodology_Keyword: vegetation sampling
Methodology_Keyword: herbivory analysis
Methodology_Description:
Twenty-three exclosures were established in Olympic National Park during two periods when there was concern about elk-vegetation relationships: 1934-1936 (18 exclosures) and 1952-1958 (5). J. E. Schwartz constructed 12 permanently closed exclosures and two series of three "take-down" exclosures which could be left open for predetermined amounts of time. These exclosures were actually established on Olympic National Forestlands that became Olympic National Park in 1938.The second pulse of exclosure construction was by C. C. Newman, who also repaired and measured some of the Schwartz exclosures. In addition, two exclosures were established in 1957 in Olympic National Forest. Although our work focused on the park, exclosures in the national forest provided an opportunity to compare the effects of ungulate herbivory in managed forests. From 1987 to 1990 we gleaned information from the early studies and, wherever possible, resampled vegetation within and adjoining exclosures. The names and numbers used to identify exclosures changed over time which resulted in considerable confusion.

Schwartz Permanent Exclosures (Nos. 3-6,14-16, 19-23) J. E. Schwartz constructed 12 permanent exclosures from 1934 to 1936 in the Hoh, South Fork Hoh, Queets and Quinault river valleys. Two have never been relocated or sampled (Nos. 6, 20). The remaining 10 were either 0.09 ha (n=6) or 0.19 ha (n=4) and were constructed of split rails and barbed wire (n=8) or woven wire (n=2).Schwartz was concerned about the level of browsing on three species he considered "key browse species," salmonberry (Rubus spectabilis), vine maple (Acer circinatum), and huckleberry (Vaccinium parvifolium, and V. alaskense) (Schwartz 1939,Schwartz and Mitchell 1945, Schwartz pers.comm. 1990). Schwartz described vegetation by estimating "density of ground cover" on a scale of 1-10 in one (sometimes two) 9.3 mē plots inside exclosures. Estimates included only herbaceous species and omitted mosses. On different 9.3 mē plots he charted aerial cover of shrubs and also estimated percent cover of "grass" (including sedges), and "weeds" (i.e., forbs including ferns). Unfortunately only preliminary data from seven exclosures remain(Schwartz and Mitchell 1945).

Schwartz Takedown Exclosures (Nos. 7-12) Schwartz constructed six "takedown" exclosures in 1936 in addition to the permanent exclosures. These were designed to measure the effects of regulated browsing on vine maple (Nos. 7-9) and huckleberry (Nos. 10-12). The three exclosures forming each series were less than 50 meters from one another. The original objective was to allow wintering elk to browse shrubs to predetermined levels (25, 50, 75% utilization of annual production), close the exclosures and monitor shrub response (Schwartz 1939). All exclosures were opened in spring 1937 and closed permanently by 1938 (Schwartz 1990 pers. comm.). All data have been lost. C. C. Newman (1953) repaired six of the Schwartz exclosures and sampled vegetation inside and outside four in 1952 and 1955. Percent cover was estimated as <1% or as midpoints in 10 subsequent classes at 10% increments. Shrub stems were counted on 2.3 mē circular plots.

Newman Exclosures (Nos. 1, 2, 13, 17, 18) Newman constructed five variously sized exclosures of woven wire from 1953 to 1958. Three were in Sitka spruce stands in different river valleys (Bogachiel, No. 2; Hoh, No. 13; Queets, No.18), one in a red alder stand (Elwha, No.l) and one along an old-field/vine maple ecotone (Queets,No. 17). Three exclosures were established at particularly heavily grazed sites (Nos. 1, 13, 17). Apparently, No. 17 was to evaluate the ability of vine maple to recolonize an abandoned hayfield in the absence of browsing. No. 13 was built mainly to determine if vine maple and big leaf maple require mineral soil to reproduce from seed; one half of the exclosure and a corresponding outside plot were spaded in 1953. Percent cover by species was estimated on eight (four inside, four outside) 2.3 mē circular plots. Plots were established systematically, one in each quarter of the exclosure with a similar pattern (usually) outside. Data were collected by park personnel regularly through 1965 and again in 1982, although methods and seasons varied.

Olympic National Forest Exclosures (Nos. 24,25) Two 0.8 ha exclosures made of woven wire were established in 1957 on clearcut western hemlock/Pacific silver fir stands logged in 1955; both were essentially intact in 1990. Logging slash was left unburned at No. 24 and burned at No. 25. One-half of each exclosure was planted with 2-year old Douglas-fir (Pseudotsuga menziesii) seedlings and the other half was allowed to regenerate naturally. Mortality and growth rates were measured from 1957 to 1962 on 100 planted Douglas-fir seedlings inside and outside each exclosure (Harrison 1963). Species composition and cover of shrubs, grasses and forbs were measured inside No. 24 during 1957 (Brent 1957).

Recent Samples Schwartz Permanent Exclosures. We examined eight exclosures from 1986-1990; none was intact. Tree composition was sampled in six exclosures (Nos. 3, 14, 19, 21, 22, 23) using one or two 100-400 mē plots inside and outside (the two remaining sites had been altered by extensive treefall and were not sampled). Cover of large organic debris (LOD) was estimated and tree age was obtained for all individuals in each plot, either from increment cores taken close to the ground or basal sections of saplings 1-4 cm in diameter. Overstory cover was estimated overall and for each species to the nearest 10 percent. Inside and outside plots were comparable with respect to overstory composition and cover of LOD. Plots inside exclosures were placed a minimum of 2 m from fencelines, while outside plots were at least 10 m and usually much further from fencelines. Quinault River exclosures (Nos. 19-23) were sampled by Pfister and Pfister (1989) under our direction. Sampling focused on vine maple age and morphology in No. 4 (see takedown exclosures).

Schwartz Takedown Exclosures. We relocated the six takedown exclosures in 1990. While examining the vine maple series, we noticed that apically dominant stems originated either directly from the ground or from branches of older stems, and stem orientation varied from vertical to horizontal. Therefore, we measured age, stem length (rather than height above ground), number, diameter, and morphology (originating from branch or ground rooted) of stems > 1 cm. Samples were taken inside Nos. 7 and 9; we also included No. 4 (permanent) because vine maple density was comparable. Takedown No. 8 was not sampled because vine maple was sparse. Plot sizes ranged from 9.5 x 14.3 m to 20 x 20 m for a total area of 675 mē inside and 850 mē outside. Because exclosures were placed specifically around clumps of vine maple, we deliberately established outside plots in stands having comparable composition and stem density. We measured dimensions of huckleberry clumps (stems arising from a common root crown) in 8.5 x 14 m plots inside and outside Nos. 11and 12. Exclosure 10 was omitted because huckleberry was uncommon. Outside plots were chosen which had huckleberry-dominated shrub layers, and overstory types, slopes, aspects and geomorphic surfaces similar to the exclosures. We counted the number of stems and measured height of the tallest stem, length and width of each clump. Canopy area was computed as an ellipse from the length and width. Several of the largest stems at each site were cut for determining age.

Newman Exclosures. We examined all five Newman sites from 1987-1990; two exclosures were intact (Nos. 1 and 2). We measured cover of shrubs(1 x 4 m plots) and herbaceous species (1 x 1 m plots) in classes of <1, 1-5, 6-25, 26-50, 51-75,76-95, and 96-100%. Fourteen subplots were placed in a single 8 x 20 m macro plot inside exclosures and distributed seven each in two 8 x 20m macro plots outside. As with the Schwartz exclosures, cover of LOD was measured, and ages of all trees >1 cm in diameter at the base were determined from cores or basal disks.

Olympic National Forest Exclosures. We sampled species composition, age and growth (i.e., dbh and height) of trees on two (burned, No. 25) or three (unburned, No. 25) 225 mē plots inside and out of each exclosure. We sampled the unplanted site at No. 25 and the planted site at No. 24. We avoided sampling other sites because vegetation heterogeneity prevented us from locating paired plots in and out of exclosures.

Photographs. In addition to sampling vegetation, we examined historic photographs. We located 48 photos of Schwartz exclosures from construction through 1980, 349 photos or slides of Newman exclosures, and 2 photos of the Forest Service exclosures.

Process_Step:
Process_Description:
Analysis, interpretation, and presentation of data from all four groups of exclosures posed certain difficulties. Although the total number of exclosures is large, only four vegetation types were replicated. Within these groups, some exclosures were not found, or appeared to be atypical for reasons other than herbivory, or included too few individuals (particularly trees) to provide meaningful conclusions. Therefore, our presentation is mostly narrative and focuses on replicated comparisons; statistical treatments are limited to robust tests with the fewest underlying assumptions. Percentage data was transformed by the arcsine of the square root before analysis. Interpretations from measurements were also compared to the photos.

Process_Date: 1993
Process_Time: Unknown
Process_Contact:
Contact_Information:
Contact_Person_Primary:
Contact_Person: Andrea Woodward
Contact_Organization: USGS
Contact_Address:
Address_Type: Mailing and Physical Address
Address: Western Fisheries Research Center,
Address: 6505 NE 65th St.
City: Seattle
State_or_Province: Wa
Postal_Code: 98115
Country: USA
Contact_Voice_Telephone: 206/526-6282 xt.332
Contact_Electronic_Mail_Address: andrea_woodward@usgs.gov

Spatial_Data_Organization_Information:
Indirect_Spatial_Reference: Olympic National Park and Olympic National Forest

Spatial_Reference_Information:

Entity_and_Attribute_Information:
Detailed_Description:
Entity_Type:
Entity_Type_Label: Characteristics of 25 exclosures
Entity_Type_Definition:
Characteristics of 25 ungulate exclosures established in Olympic National Park and Olympic National Forest, 1934-1958.
Attribute:
Attribute_Label: Site name
Attribute:
Attribute_Label: Date established
Attribute_Definition: Date the exclosure was established
Attribute:
Attribute_Label: Original Biologist
Attribute_Definition: Name of biologist who established the exclosure
Attribute:
Attribute_Label: Years sampled
Attribute_Definition: Years each exclosure was sampled
Attribute:
Attribute_Label: methods used
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: A
Enumerated_Domain_Value_Definition: Cover in 9.3m2 plots
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: B
Enumerated_Domain_Value_Definition: Cover and stem count in 2.3m2 plots
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: C
Enumerated_Domain_Value_Definition: tree mortality and growth
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: D
Enumerated_Domain_Value_Definition: Examination and/or measurements, this study
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: P
Enumerated_Domain_Value_Definition: Photo
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: X
Enumerated_Domain_Value_Definition: Known to be no longer functional
Attribute:
Attribute_Label: Comments
Attribute:
Attribute_Label: Vegetation type
Attribute_Definition: Vegetation inside the exclosures
Attribute:
Attribute_Label: Area (ha)
Attribute_Definition: Size of the exclosure
Entity_Type:
Entity_Type_Label: Vegetation cover changes
Entity_Type_Definition:
Summary of vegetation cover changes for exclosures set up by Schwartz and Newman at several post-establishment time intervals.
Attribute:
Attribute_Label: Exclosure site
Attribute:
Attribute_Label: Intervals (year)
Attribute_Definition: Intervals at which cover changes were sampled
Attribute:
Attribute_Label: Change in vegetation cover
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: grasses
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: forbs
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: ferns
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: shrubs
Entity_Type:
Entity_Type_Label: Huckleberry clumps
Entity_Type_Definition:
Dimensions of huckleberry clumps in take-down exclosures Nos. 11 and 12 and control plots established by Schwartz.
Attribute:
Attribute_Label: Clump density (no/m2)
Attribute:
Attribute_Label: Stem density (no./m2)
Attribute:
Attribute_Label: Canopy area (area/m2)
Attribute:
Attribute_Label: Height (cm)
Attribute_Definition: Height of clumps
Attribute:
Attribute_Label: Canopy area (m2/m2)
Attribute_Definition: canopy area of clumps
Attribute:
Attribute_Label: Stems (no.)
Attribute_Definition: Number of huckleberry stems in a clump
Entity_Type:
Entity_Type_Label: Establishment and survival
Entity_Type_Definition:
Comparison of establishment and survival of Pacific silver fir, western red cedar, and western hemlock in and out of two exclosures established in clearcuts.
Attribute:
Attribute_Label: Sp. Name
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: Pacific Silver fir
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: Western Red Cedar
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: Western Hemlock
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: Douglas-fir
Attribute:
Attribute_Label: Exclosure number
Attribute:
Attribute_Label: Living (stems/ha)
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: In
Enumerated_Domain_Value_Definition: Inside the exclosure
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: Out
Enumerated_Domain_Value_Definition: Outside the exclosure
Attribute:
Attribute_Label: Dead (stems/ha)
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: In
Enumerated_Domain_Value_Definition: Inside the exclosure
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: Out
Enumerated_Domain_Value_Definition: Outside the exclosure
Entity_Type:
Entity_Type_Label: Establishment of vine maple
Entity_Type_Definition:
Establishment of vine maple inside and outside the exclosures since construction in 1936.
Attribute:
Attribute_Label: Total stems
Attribute_Definition: Number of stems
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: In
Enumerated_Domain_Value_Definition: Inside the exclosure
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: Out
Enumerated_Domain_Value_Definition: Outside the exclosure
Attribute:
Attribute_Label: ground rooted
Attribute_Definition: Number of stems that are rooted in the ground
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: In
Enumerated_Domain_Value_Definition: Inside the exclosure
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: Out
Enumerated_Domain_Value_Definition: Outside the exclosure
Entity_Type:
Entity_Type_Label: Distribution of stem length
Entity_Type_Definition:
distribution of stem length classes for three age cohorts of vine maples measured 54 years after establishment inside and outside the exclosures.
Attribute:
Attribute_Label: Stem Length Class
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: In
Enumerated_Domain_Value_Definition: Inside the exclosure
Attribute_Domain_Values:
Enumerated_Domain:
Enumerated_Domain_Value: Out
Enumerated_Domain_Value_Definition: Outside the exclosure

Distribution_Information:
Distributor:
Contact_Information:
Contact_Person_Primary:
Contact_Person: Andrea Woodward
Contact_Organization: USGS
Contact_Address:
Address_Type: Mailing and Physical Address
Address: Western Fisheries Research Center,
Address: 6505 NE 65th St.
City: Seattle
State_or_Province: Wa
Postal_Code: 98115
Country: USA
Contact_Voice_Telephone: 206/526-6282 xt.332
Contact_Electronic_Mail_Address: andrea_woodward@usgs.gov
Resource_Description: journal article
Standard_Order_Process:
Non-digital_Form:
Contact author for data or reprints, or obtain reprints from local library

Metadata_Reference_Information:
Metadata_Date: 19980722
Metadata_Review_Date: 19980929
Metadata_Contact:
Contact_Information:
Contact_Organization_Primary:
Contact_Organization: Olympic Natural Resources Center
Contact_Position: GIS Specialist
Contact_Address:
Address_Type: Mailing Address
Address: PO Box 1628
City: Forks
State_or_Province: WA
Postal_Code: 98331
Country: USA
Contact_Voice_Telephone: 360.374.3220 x258
Contact_Facsimile_Telephone: 3603743336
Contact_Electronic_Mail_Address: onrc@u.washington.edu
Hours_of_Service: M-F 8-5
Contact_Instructions: Email preferred
Metadata_Standard_Name:
Content Standards for National Biological Information Infrastructure Metadata
Metadata_Standard_Version:
NBII Draft of December 1995, Based FGDC of June 8, 1994
Metadata_Access_Constraints: None
Metadata_Use_Constraints: None
Metadata_Security_Information:
Metadata_Security_Classification_System: None
Metadata_Security_Classification: Unclassified
Metadata_Security_Handling_Description: None

Generated by mp on Tue Sep 29 14:07:04 1998