MOVEMENTS, HABITAT USE, AND ACTIVITY PATTERNS
OF A TRANSLOCATED GROUP OF ROOSEVELT ELK
by
Matthew McCoy
A Thesis
Presented to
The Faculty of Humboldt State University
In Partial Fulfillment
of the Requirements for the Degree
Master of Science
June, 1 986 MOVEMENTS, HABITAT USE, AND ACTIVITY PATTERNS
OF A TRANSLOCATED GROUP OF ROOSEVELT ELK
by
Matthew McCoy
A
F gam
Director, Natural Resources Graduate Program
86/W-66/06/10 Natural Resources Graduate Program Number
Ala M. Gillespi9( ABSTRACT
In March 1982, 17 Roosevelt elk (Cervus elaphus roosevelti) were
captured at Gold Bluffs Beach, Humboldt County, California and
translocated to an enclosure 4 km east of Shelter Cove, Humboldt County.
The animals were released from the enclosure in November 1982. Data were collected for eight radio-collared animals (seven adult females and one 4-year old male) January to December 1983.
The number of herds varied from one (January through March) to
four (October through December). Three female herds moved 45 km, 58 km,
and 84 km south of Shelter Cove, while the radio-collared male remained within 16 km of Shelter Cove. Home range locations varied seasonally.
Home range sizes were largest during the summer reflecting migrational
and exploratory movements.
Habitat use was disproportionate to habitat availability at the home range level during each season. Cultivated grasslands and riparian
areas were used in proportions greater than their availability. Coastal
Prairie use was greater than that available except in the Shelter Cove
area. Shrub and forest habitat types were generally used less than that
available. Animal distances to nearest road and water varied
seasonally, but were greatest for radio-collared females during the
spring. Use of cover was greatest during spring and summer and was
correlated with increased temperature and decreased cloud cover.
Activity followed a diurnal pattern with extended periods of
activity (two or more hours) occurring at sunrise and sunset and shorter
iii iv periods (about one hour) occurring at night and during the day. The amount of activity decreased significantly (p < 0.02) during the year.
Animals were most active during the winter (43 percent of a 24—hour period) and least active during the fall (36 percent of a 24—hour period). ACKNOWLEDGEMENTS
I would especially like to thank Dr. R. Golightly, my major professor, for his advice, support, patience, and friendship. His efforts are largely responsible for the completion of this thesis. I would also like to thank the members of my graduate committee: Drs. D.
Kitchen and K. Fulgham, for their advice and criticism.
The Bureau of Land Management provided a vehicle, maps, and equipment. The California Department of Fish and Game provided radio-telemetry equipment and the animals. Humboldt State University provided additional equipment and computer facilities. Residents in the study area provided useful information and support.
Special thanks to M. Bivin for offering encouragement, helpful criticism, and for being a friend. B. Hardenbrook and J. Grenier also provided much appreciated manuscript reviews. B. Burger was more helpful than he could have imagined. Thanks to all the people who remained friends and offered support through the ups and downs. I am a better person for knowing them.
Finally, I wish to thank my family for their support and encouragement. This thesis is dedicated to the memory of my father, I regret he could not be here to enjoy it. TABLE OF CONTENTS
Page
ABSTRACT iii
ACKNOWLEDGEMENTS
LIST OF TABLES ix
LIST OF FIGURES xi
INTRODUCTION 1
STUDY AREA 5
Habitat Types 5
Grassland 5
Shrubland 7
Forest 7
Riparian 8
MATERIALS AND METHODS 10
Capture and Release 10
Animal Data Collection 11
Habitat Data Collection 13
Animal Data Analysis 14
Definition of Season, Herd, and Movement 14
Test for Independence 15
Home Range Area 15
Habitat Use 15
Comparison of Distance Variables Between Seasons 16
Slope and Climate 16
vi vii
TABLE OF CONTENTS (CONTINUED) Page
Activity Pattern 17
Deliniation of Habitat Types 17
RESULTS 18
Movements and Home Ranges 18
Habitat Types 22
Habitat Use 22
Use of Grassland Habitats 32
Use of Shrub and Forest Habitats 32
Use of Riparian Habitats 33
Distance to Road 33
Distance to Water 33
Distance to Cover 36
Slope 36
Climate 36
Activity Pattern 40
Time Spent Active 40
Specific Behaviors 42
DISCUSSION 45
Movements and Home Ranges 45
Habitat Use 49
Use of Grassland Habitats 49
Use of Shrub and Forest Habitats 50
Use of Riparian Habitats 50
Distance to Road 51
Distance to Water 51 viii
TABLE OF CONTENTS (CONTINUED) Page
Distance to Cover 52
Slope 52
Climate 53
Activity Pattern 53
Time Spent Active 54
Time Allocation to Specific Activities 56
Conclusions 58
REFERENCES CITED 60
APPENDIX A. Animal Capture 68
APPENDIX B. Animal Movements 70
APPENDIX C. Habitat Sampling 72
LIST OF TABLES
Table Page
1 Herd Movements of Translocated Roosevelt Elk, Humboldt and Mendocino Counties, California, January-December 1983. 19
2 Seasonal Averages of Individual Home Range Areas (ha) for Translocated Roosevelt Elk, Humboldt and Mendocino Counties, California, Winter-Fall 1983. 21
3 Total Hectares and Total Number of Hourly Locations of Translocated Roosevelt Elk by Activity for each Habitat Type in the Study Area, Humboldt and Mendocino Counties, California, January-December 1983. 27
4 Habitat Use by Bull #201RW, Humboldt County, California, Winter-Fall 1983. 28
5 Habitat Use by the Red Herd (Cows #226R and #233B), Humboldt and Mendocino Counties, California, Winter-Fall 1983. 29
6 Habitat Use by Cow #231W, Humboldt and Mendocino Counties, California, Winter-Fall 1983. 30
7 Habitat Use by the Orange Herd (Cows #2290, #236RWB, #237Y, and #240G), Humboldt and Mendocino Counties, California, Winter-Fall 1983. 31
8 Yearly and Seasonal Breakdown of Animal Distances to Road for Translocated Roosevelt Elk, Humboldt and Mendocino Counties, California, Winter-Fall 1983. 34
9 Yearly and Seasonal Breakdown of Animal Distances to Water for Translocated Roosevelt Elk, Humboldt and Mendocino Counties, California, Winter-Fall 1983. 35
10 Yearly and Seasonal Breakdown of Animal Distances to Cover for Translocated Roosevelt Elk, Humboldt and Mendocino Counties, California, Winter-Fall 1983. 37
11 Slope Use by Translocated Roosevelt Elk, Humboldt and Mendocino Counties, California, January-December 1983. 38
12 Correlation Between Habitat Type Used (Grassland, Riparian, and Forest) and Climatic Variables (Ambient Temperature, Cloud Cover, Precipitation, and Wind Speed) for Translocated Roosevelt Elk, Humboldt and Mendocino Counties, California, Spring-Fall 1983. 39
is LIST OF TABLES (CONTINUED)
Table Page
13 Percentage Time Spent in Specific Active and Inactive Behaviors for Several Herbivores. 57
14 Dominant Plant Species in the Seven Grassland Subgroups Determined Using Cluster Analysis, Humboldt and Mendocino Counties, California. 72
15 Dominant Plant Species in the Three Shrubland Subgroups Determined Using Cluster Analysis, Humboldt and Mendocino Counties, California. 73
16 Dominant Plant Species in the Seven Forest Subgroups Determined Using Cluster Analysis, Humboldt and Mendocino Counties, California. 74
17 Dominant Plant Species in the Three Riparian Subgroups Determined Using Cluster Analysis, Humboldt and Mendocino Counties, California. 75
LIST OF FIGURES
Figure Page
1 Initial Trap Site and Study Area for the Translocation of Roosevelt Elk, Humboldt and Mendocino Counties, California, February 1982 through December 1983. 6
2 Study Area for the Translocation of Roosevelt Elk, Humboldt and Mendocino Counties, California, January-December 1983. 20
3 One-hundred, 90, and 50 Percent Home Range Estimates (Convex Polygon), with Number of Observations Used in the 100 Percent Estimate, of Bull #201RW, January-December 1983. 23
4 One-hundred, 90, and 50 Percent Home Range Estimates (Convex Polygon), with Number of Observations Used in the 100 Percent Estimate, of the Red Herd (Cows #226R and #233B), January-December 1983. 24
5 One-hundred, 90, and 50 Percent Home Range Estimates (Convex Polygon), with Number of Observations Used in the 100 Percent Estimate, of Cow #231W, January-December 1983. 25
6 One-hundred, 90, and 50 Percent Home Range Estimate (Convex Polygon), with Number of Observations Used in the 100 Percent Estimate, of the Orange Herd (Cows #2290, #236RWB, #237Y, and #240G), January-December 1983. 26
7 Percentage Time Collared Roosevelt Elk Were Observed Active during each Hour, Humboldt and Mendocino Counties, California, Winter-Fall 1983. 41
8 Monthly Breakdown of Specific Active Behaviors Based on 5-minute Scan Data for Translocated Roosevelt Elk, Humboldt and Mendocino Counties, California, January-December 1983. 43
9 Monthly Breakdown of Specific Inactive Behaviors Based on 5-minute Scan Data for Translocated Roosevelt Elk, Humboldt and Mendocino Counties, California, January-December 1983. 44
10 Percentage Time Spent Active (during a 24-hour Period) as a Function of Body Weight (log w( kg )) of Several Herbivores. 55 11 Location of Prebaited Trap Sites in Prairie Creek Redwoods State Park and Redwood National Park, Humboldt County, California, February 1982. 69
xi INTRODUCTION
Historically, Roosevelt elk (Cervus elaphus roosevelti) were found in the Pacific states from the Olympic Peninsula, Washington to
San Francisco Bay, California and inland to the Cascade Mountains (Murie
1951). Overhunting and habitat loss have reduced California's Roosevelt elk population numbers and restricted their distribution (Orr 1937,
Mandel 1979).
Translocations have long been a popular method of re-establishing animals in their former ranges (Bryant and Maser 1982). From 1892 to
1967, approximately 13,500 Rocky Mountain elk (C. e. nelsoni) from
Yellowstone National Park and the surrounding area were translocated into many parts of the United States, Canada, and Mexico (Robbins et al.
1981). Roosevelt elk were successfully translocated to Afognak Island,
Alaska from Olympic National Park, Washington (Troyer 1960). In contrast, an attempt in 1964 and 1965 to translocate elk from Gold
Bluffs Beach in Prairie Creek Redwoods State Park, California to an area
48 km northeast of the park partially failed when three of the 14 translocated animals returned to their former home range (Franklin et al. 1975).
Studies of Roosevelt elk in Prairie Creek Redwoods State Park and
Redwood National Park have been conducted on unmarked and on eartagged animals. The studies have been concerned primarily with behavior (Lieb
1973, Bowyer 1976), habitat use (Bentley 1959, Franklin 1968), and general ecology (Harp 1958, Harper 1961, Logsdon 1965, Stevens 1965,
1 2
Lemos 1971, Mandel 1979). Information on movements, home ranges, habitat use, and activity patterns is either qualitative in nature or
limited in scope.
Animals react to their environment on a variety of levels
(Johnson 1980). For example, animals establish home ranges within a
geographical range, and within these home ranges certain habitats and
habitat components are used to meet behavioral and physiological needs.
Knowledge of an animal's movements is an important aspect in
understanding a species' ecology. Movement patterns are useful in
predicting how a translocated group of animals may react to a new
environment (Sanderson 1966). Daily movements define an animal's home
range, whereas exploration, migration, and dispersion result in
movements between disjunct home ranges.
Roosevelt elk home ranges have been described for open meadow
habitats (Franklin et al. 1975), silviculturally managed forests (Graf
1955, Witmer 1982), and old-growth conifer forests (Jenkins and Starkey
1984). Comparisons between different home range sizes have been used as
an indicator of habitat quality. However, comparisons of home range
sizes may be misleading because of the biases of the specific method
used to determine home range size (Mohr 1947, Anderson 1982, Laundre and
Keller 1984). Adams and Davis (1967) and Laundre and Keller (1984)
stated that spatial and temporal use patterns within a home range were
more important than the actual size of the area used.
Seasonal habitat use patterns depend on a variety of factors such
as traditional habitat use patterns, food and cover availability and
quality, type of activity, climate, animal physiology, group size, and 3 topography. These factors operate either alone or in combination.
Habitat use patterns have been described for a variety of elk populations (Beall 1976, Marcum 1976, Irwin 1978, Morgantini and Hudson
1979, Pederson et al. 1980); however, only recently have habitat use patterns been discussed in relation to availability as defined by an animal's biology (Witmer and deCalesta 1983, Jenkins and Starkey 1984).
Diurnal activity patterns have been reported for a variety of ungulates (Craighead et al. 1973, Collins et al. 1978, Georgii 1981,
Georgii and Schroder 1983 for elk and red deer, C. elaphus; McMillan
1954, Geist 1963, Belovsky 1981 for moose, Alces alces; Ozoga and Verme
1970, Kammermeyer and Marchinton 1977 for white-tailed deer, Odocoileus virginianus; Ellis and Travis 1975 for pronghorn antelope, Antelocapra americana; Jarman and Jarman 1973 for impala, Aepvceros melampus).
However, detailed studies of 24-hour activity patterns for an entire year are limited (Georgii 1981, Georgii and Schroder 1983, Jingfors
1982). While studies have documented seasonal anomalies in elk activity patterns (i.e. Bowyer 1981 on breeding), direct observational studies
for an entire year are lacking.
Most elk ecology studies have been conducted on established herds
(i.e. Knight 1970, Craighead et al. 1973). Conversely, few studies have been conducted to determine the result of efforts to translocate
animals. The translocation of radio-collared elk to the King Range
Conservation Area, Humboldt County, California provided an opportunity
to study movements, home range dynamics, habitat use, and activity
patterns of translocated elk. Objectives of the study were to determine
or describe: 1) temporal changes in movement patterns; 2) seasonal 4 changes in home range sizes and use patterns; 3) if habitat use was in proportion to habitat availability at the home range level; 4) daily and seasonal changes in habitat use patterns; and 5) daily and seasonal changes in activity patterns. STUDY AREA
The study area for the translocated elk extended from Shelter
Cove, Humboldt County south to Fort Bragg, Mendocino County and as far east as Whitethorn, Humboldt County (Figure 1). The term "study area" is used to denote the total area traversed by all animals during 1983.
Elevations in the study area ranged from sea level to 780 m. Aspects in the study area were primarily flat or west facing.
Ambient temperatures at Shelter Cove averaged 16 ° C during the study, with a maximum of 30 °C and a minimum of 3 ° C (NOAA 1983). A total of 281.2 cm of rain was recorded in 1983 at Shelter Cove, 124 cm above the previous 10 year average. January through April, August, and
October through December accounted for 96 percent of the annual precipitation (NOAA 1983).
Habitat Types
Four major physiognomic categories of vegetation were present on the study area: grassland, shrubland, forest, and riparian.
Grassland
Grasslands were represented by two major types: cultivated grasslands and Coastal Prairie (Munz 1959). Cultivated grasslands (i.e. golf courses, residential home sites) were dominated by bluegrass (Pon sp.), fescue (Festuca sp.), perennial ryegrass (Lolium perenne), and clover (Trifolium sp.). Coastal Prairies consisted primarily of velvet grass (Holcus lanatus), California oatgrass (Danthonia californica), and
5 6
Figure 1. Initial Trap Site and Study Area for the Translocation of Roosevelt Elk, Humboldt and Mendocino Counties, California, February 1982 through December 1983. 7 to a lesser extent thin grass (Arrostis diegoensis), orchard grass
(Dactvlis glomerata), perennial ryegrass, blue wild-rye (Elvmus glaucus). Dogtail (Cvnosurus echinatus) was the most abundant annual grass on less well-developed soils. Scattered coyote brush (Baccharis pilularis) provided limited shelter from wind. Common forbs included bull thistle (Cirsium arvense), common dandelion (Taraxacum officinale), plantain (Plantago lanceolata), sheep sorrel (Rumex acetosella), and bracken fern (Pteridium aauilinum).
Shrubland
Shrublands were characterized by a dense shrub overstory.
Blueblossom (Ceanothus thvrsiflorus) and tanoak (Lithocarpus densiflorus) or hairy manzanita (Arctostaphylos columbiana), evergreen huckleberry (Vaccinium ovatum), and golden chinkapin (Castanopsis
chrysophvlla) characterized Chaparral type shrublands with few understory forbs or grasses (Munz 1959). California hazel (Corylus cornuta) or coyote brush and California blackberry (Rubus vitifolius) dominated North Coastal Scrub types and had understories including strawberry (Frageria chiloensis), lupine (Lupinus sp.), hedge-nettle
(Stachys rigida), bracken fern, sword fern (Polystichum munitum), and velvet grass (Munz 1959).
Forest
Forest vegetation types were represented by Douglas-fir Forest,
Redwood Forest, and Mixed Evergreen Forest types (Munz 1959). Overstory
cover and composition and understory productivity and composition depended to a large extent on past logging practices at the site. 8
The Douglas-fir Forest type was dominated by a Douglas-fir
(Pseudotsuga menziesii) overstory. Understory species included evergreen huckleberry, wood rose (Rosa gymnocarpa), iris (Iris sp.), vetch (Vicia sp.), bracken fern, sword fern, and Bromus vulgaris. An understory tree layer of tanoak, Pacific madrone (Arbutus menziesii), and California laurel (Umbellularia californica) differentiated the
Mixed Evergreen Forest type from the Douglas-fir Forest type.
The Redwood Forest type was composed of a coast redwood (Sequoia sempervirens), Douglas-fir, and, south of Usal Creek, grand fir (Abies grandis) overstory. Dominant understory trees included tanoak and
Pacific madrone. Understory shrub and forb species included evergreen huckleberry, coast rhododendron (Rhododendron macrophvllum), salal
(Gaultheria shallon), bracken fern, sword fern, redwood-sorrel (Oxalis oregana), and sphagnum moss (Sphagnum sp.). Additional understory species in selective cut Redwood Forest types included coffeeberry
(Rhamnus californica), coyote brush, thimbleberry (Rubus parvaflorus), fireweed (Erectites prenanthoides), hedge-nettle, bull thistle,
strawberry (Fragaria californica), vetch, Yerba de Selva (Whipplea modesta), starflower (Trientalis latifolia), velvet grass, and blue wild-rye. Recent clear cuts consisted primarily of sapling hardwoods,
seedling conifers, and a variety of shrubs, forbs, and grasses similar to those found in selective cuts. Older clear cuts were dominated by tanoak and madrone overstories, scattered conifers, and rhododendron,
evergreen huckleberry, salal, and bracken fern understories. 9 Riparian
Riparian areas were dominated either by red alder (Alnus rubra) or willow (Salix spp.) in exposed areas near the coast. Understory species included thimbleberry, California blackberry, Himalaya berry
(Rubus thyrsanthus), horsetail (Equisetum sp.), hedge-nettle, sword fern, bracken fern, figwort (Scrophularia californica), cow-parsnip
(Heracleum lanatum), velvet grass, rush (Juncus spp.), and bullrush
(Scirpus spp.). MATERIALS AND METHODS
Capture and Release
On 10 March 1982, ten adult and three yearling females, one yearling male, and one male calf were captured in a trap located 14 km north of Orick, Humboldt County (Figure 1, Appendix A). In addition, a three-year old male and a yearling male were darted and immobilized
(with etorphine hydrochloride), then processed with the other animals.
Numbered ear tags were placed on each individual. Color-coded radio-collars (159MHz; Telonics Inc., Mesa AZ 85201) were placed on the three-year old male and seven adult females. The animals were transported in horse trailers to the Bear Creek enclosure and released the morning of 11 March 1982 (Figure 1). One uncollared adult cow was euthanized because of injuries suffered during transport.
The animals were held in an enclosure to promote site fidelity
(Wilson et al. 1973). The Bear Creek enclosure was constructed of two tiers of sheep fencing and two strands of smooth wire for a total height of 3.5 m. The enclosure area was approximately 28.5 ha, including a
16.3 ha meadow and two small creeks. The elk were monitored in the enclosure during the summer of 1982 by Bureau of Land Management personnel. In early November 1982, gates at the southern and northern ends of the enclosure were opened and two additional openings were made along the west side of the enclosure. This allowed the elk free access in and out of the enclosure.
10 1 1
Animal Data Collection
The elk became habituated to my presence during November and
December 1982. The animals were considered to be habituated to my presence when they no longer reacted to my presence. Observations were made from a vehicle or on foot at distances ranging from 5 m to 750 m.
When possible, animals were observed from a vehicle to minimize possible disturbance. Data collection was restricted to radio-collared animals which assured recognition of individuals and some data (location and activity) collection when visual contact was impossible (night-time or in cover). Each radio-collared animal was sampled at least one 24-hour period each month. Observations usually occured over a three day period between Thursday and Monday of each week. Only data collected from
January through December 1983 were used in analyses.
A focal animal (Altmann 1974) was observed for a 24-hour period starting during the activity period just before sunset. Animal locations were recorded as X and Y coordinates which corresponded to
4.05 ha grids on topographic maps. When visual (daylight) contact could be maintained, data from an individual were collected every five minutes. Parameters recorded during each instantaneous sample (Altmann
1974) included time of day, location, and activity. Animal activity was based on the following classification: grazing; browsing; standing; standing-grooming; standing-interacting; urinating/deficating; walking; walking-interacting; walking-ruminating; bedding; bedding-ruminating; bedding-grooming; bedding-feeding; bedding-interacting; and sleeping
(Craighead et al. 1973, Bowyer 1976, Clutton-Brock et al. 1982). Habitat use was recorded as presence in one of the habitat types described for 12 the study area. Distances between the focal animal and the nearest neighbor, road, water, and cover (any vegetation or structure that could provide shelter from wind) were qualitatively estimated or measured on aerial photographs and recorded in interval classes. Percentage slope was measured with a clinometer. Wind direction and speed, percentage cloud cover, and type of precipitation were qualitatively estimated and recorded in interval classes. Ambient temperature was measured in the shade. Herd size and human influence (presence of humans on foot, motorcycle, automobile, plane or presence of dogs) were also recorded.
Daytime observations were aided by the use of 7 by 35 mm binoculars.
When visual contact was not possible, data were collected at least once an hour. Non-visual animal locations were made using a receiver (model TR1 or model TR2 with digital processor, Telonics Inc.) with a two-element Tag/ antenna. Locations were based on compass bearings of maximum signal strength measured from two known locations.
Two measurements were probably adequate because non-visual locations were generally in homogenous forest types and the distance between observer and animal was relatively short (usually less than one km).
Heezen and Tester (1967) found a significant loss in accuracy of location (based on two bearings) at distances greater than 1.6 km between observer and animal; consequently, locations of elk based on measurements from distances greater than 1.6 km were not used in analyses. Animals were classified as either active or inactive based on the amplitude variation in the received signal. Night-time locations and activities were verified with a spotlight when possible. 13
Habitat Data Collection
Habitat types were mapped on 1:24,000 scale topographic maps using infrared (1978 flight, 1:32,000 scale, Humboldt State University,
Arcata CA 95521 and California Coastal Commission, Eureka CA 95501) and color (1981 flight, 1:20,000 scale, Bureau of Land Management, Arcata CA
95521) aerial photographs. A zoom transfer scope (Bausch and Lomb,
Rochester, NY 14602) was used to correct for differences between photo and map scales. A compensating polar planimeter (model 3651-30, Lietz,
Overland Park, RA 66201) was used to determine areas of habitat types from the maps.
Habitat types were sampled in proportion to their occurrence in the study area. Field habitat sampling was conducted in November 1984 using the Relevé method (Mueller-Dombois and Ellenberg 1974). Circular plots of 50 m 2 (grassland), 100 m2 (shrubland) and 405 m2 (forestland) in size were used. Percentage ground cover of each species present in a plot was recorded in one of seven classes (solitary plant, few plants, less than 5%, 5-25%, 25-50%, 50-75%, or greater than 75% cover). Tree cover was measured using a spherical densiometer (Forest Densiometers,
Arlington, VA 22207). Tree density was estimated using a Bitterlich prism [either a 15 basal area factor (BAF) for smaller trees, or a 40
BAF for larger trees] (Forestry Supplies, Jackson, MS 39204).
Percentage slope was determined using a clinometer. 14
Animal Data Analysis
Data were analysed using nonparametric statistical programs from the Statistical Package for the Social Sciences (Hull and Nie 1981) or
Biomedical Data Package (Dixon 1981). Variables were classified as either nominal (habitat type; activity) or ordinal (season; time of day; distances to road, nearest neighbor, water, and cover; percent slope; wind speed; cloud cover; precipitation; temperature). Choice of statistical methods used was based on this classification.
Definition of Season, Herd, and Movement
The division of a year into seasons has an important affect on the results of data analyses. Seasons should relate to the reproductive cycle of the animal being studied (Laundre and Keller 1984). Because no females calved in 1983 and no males were with females during most of the rut, traditional calendar seasons [winter (January through March), spring (April through June), summer (July through September) and fall
(October through December)] were used in seasonal data analyses.
Groups of animals that remained in a cohesive unit during a specific time period (month or season) were called herds. Herds were identified by their geographic location or the eartag number of the dominant radio-collared animal.
Movements of less than 14 km within home ranges and between seasonally disjunct home ranges were classified as exploratory. The return of animals to a previously used home range was classified as migration. Movements of more than 14 km to new home ranges were classified as dispersal if the animals did not return to former home ranges during the study. 15
Test for Independence
The runs test (Sokal and Rohlf 1981) was used to examine continuous dichotomous data for randomness. The activity pattern variable was chosen as the indicator of randomness in the data because all other animal related parameters were dependent on activity.
Therefore, if a series of observations were independent for the activity variable, then they were probably independent for all animal related variables. Runs tests were performed on the activity pattern variable to determine the minimum time interval between independent observations.
Observations made at hourly intervals were found to be independent of each other. Subsequent statistical analyses were based on data collected at this time interval.
Home Range Area
Home range areas were calculated using a program from Harestad
(1981) based on the minimum area (convex polygon) method (Mohr 1947).
The 100 percent home range areas were calculated using all locations for the given animal(s) and time. The 90 and 50 percent home range calculations were based on the omission of 10 and 50 percent of the locations (outermost from the geometric center) respectively.
Individual home range areas were compared by season using a single factor analysis of variance test (Sokal and Rohlf 1981). Monthly home ranges of herds were calculated using locations of all animals within a specific herd.
Habitat Use
Chi-square tests (Sokal and Rholf 1981) were used to determine if seasonal habitat use was nonrandom for each herd. Habitat availability 16
(expected frequency) was compared with actual use (observed frequency) by each herd at the seasonal home range level. Habitat available to a herd included all habitat types within both the seasonal home range and a buffer strip around the home range. The buffer strip extended either
0.4 km beyond the outermost locations or to natural barriers (i.e. drainages, ridgetops). Habitat types of limited availability (small size) were pooled so that expected frequencies were greater than five.
Available habitat types that were not used by a herd were not included in the analyses (Johnson 1980). Significant relationships were tested using a technique described by Neu et al. (1974) to determine which specific habitats were used more or less frequently than expected.
Comparison of Distance Variables between Seasons
Observed distances (to road, water, and cover) were compared between seasons for each herd using the Kruskal-Wallis test (Sokal and
Rholf 1981). The Scheffe test was used to distinguish between similar and dissimilar seasonal distributions.
Slope and Climate
Chi-square tests were used to determine if slope use by all animals was nonrandom in either grassland or forest habitat types.
Frequencies of slopes available to animals were derived from slope measurements made during habitat sampling.
Relationships between climate variables and habitat use were determined using Kendall rank correlation (Sokal and Rholf 1981).
Correlations were determined for all animals combined for each season. 17
Activity Pattern
Active-inactive observations were compared for eight three-hour time intervals during a 24-hour period. Log-likelihood ratio (or G) tests (Sokal and Rholf 1981) were used to determine if activity patterns varied by time of day and season.
The effects of habitat (including habitat type, and distances to road, water, and cover) and climate on activity were determined using
Kendall rank correlation tests. The proportion of time spent in specific behaviors was determined using the five-minute scan data; however, no statistical analyses were performed on these results because the data were not independent.
Deliniation of Habitat Types
Habitat data were analysed according to major physiognomic groups
(grass, shrub, forest, and riparian). Subgroups of similar Releve plots were delineated by cluster analysis using the Clustan program package
(Wishart 1982). The hierarchy method of pairwise comparisons of Releves was used to break a physiognomic group into a series of subgroups. The
Ward's (error sum of squares) method was the specific hierarchical method used (Ward 1963). Species composition and cover were used to describe subgroups. RESULTS
From January through December 1983 a total of 12,890 individual scans were collected from observations of radio-collared elk. Two calves were born in the enclosure in June 1982. One calf was killed by an automobile in December 1982, the remaining 17 animals (12 females, four males and one male calf) survived through the end of the study.
Movements and Home Ranges
In January 1983 all 17 elk were at Shelter Cove. However, by
October 1983 the animals were in four separate herds on home ranges located at Shelter Cove (male herd), Usal Creek (red herd), Westport
(white herd), and Tenmile River (orange herd) (Table 1, Figure 2,
Appendix B). Distances moved between home ranges were greatest in spring and summer. Females from the orange and white herds moved the greatest distance (approximately 58 km) from Chemise Creek to Howard
Creek in June. In late July the orange herd moved from Westport to Fort
Bragg (approximately 26 km) before returning to the Tenmile River area.
The red herd moved approximately 45 km from Bear Creek to the Usal Creek area in early September. Movements by bull #201RW were restricted to the Shelter Cove/Bear Creek/Chemise Creek area. The male's movements were greatest during the rut in September.
Individual home range areas varied significantly (p < 0.03) between seasons (Table 2). Seasonal home ranges were largest in summer because of movements between different monthly home ranges. Monthly
18 Table 1. Herd Movements of Translocated Roosevelt Elk in Humboldt and Mendocino Counties, California, January-December 1983. Herd Names Denote Groups of Animals That Were in a Cohesive Group in December (Male Herd = Bulls #201RW a , #242, #243; Red Herd = Cows #226R, #233B, #234, Male Calf; White Herd = Cows #231W, t227, #238; and Orange Herd = Cows #2290, #236RWB, #237Y, #240G, #230, #235, Bull #202 ). Dates Animals Were Initially Observed in an Area Are Indicated by Location Codes (SC=Shelter Cove, EN=enclosure, CC=Chemise Creek, BC=Bear Creek, BH=Bear Harbor, UC=Usal Creek, WP=Westport, TMR=Tenmile River).
Herd Month
J F M A M J J A S 0 N D
Male herd SC EN-CC---SC BC SC
Red herd SC EN-CC EN SC BC UC
White herd SC EN-CC BH- WP
Orange herd SC EN-CC BH WP- -TMR a Letter denotes a collared animal. b Bull #202 was transferred to the enclosure in August 1983. 20
Figure 2. Study Area for the Translocation of Roosevelt Elk, Humboldt and Mendocino Counties, California, February 1982 through December 1983. Table 2. Seasonal Averages of Individual Home Range Areas (in ha) for Translocated Roosevelt Elk in Humboldt and Mendocino Counties, California, Winter-Fall 1983. Areas Were Calculated Using the Convex Polygon Method (Harestad 1981).
Percentage of Home Range Area (ha) by Season Locations Used in Estimate (x + SD number of animals range of values
Winter Spring Summer Fall
100 Percent 17+4.4 108+152.9 2369+3130.6 178+181.4 7 7 8 8 12-24 22-85 148-8029 18-576
95 Percent 13+5.8 51+18.7 2302+3097.5 102+71.1 7 7 8 8 6-24 22-59 210-8252 12-192
50 Percent 3+1.0 7+4.3 151+25.3 12+4.7 2 2 7 6 2-4 4-10 10-657 6-20 22 home range areas of herds were largest during July (orange and white herds), August (red herd), and September (bull #201RW) (Figures 3-6).
Movements by bull #201RW during the rut resulted in large monthly home range areas (100 and 90 percent) during August and September (Figure 3).
Movements by the red herd at Usal Creek resulted in large monthly home range areas (100 and 90 percent) during September, October, and December
(Figure 4).
Habitat Types
Sixteen major habitat types were delineated from aerial photographs of the entire study area (Table 3). A total of 183 Relevé plot analyses were conducted: 48 in grasslands; 118 in shrub and forest habitat types; and 17 in riparian areas. Cluster analysis resulted in the identification of seven grassland habitat types, 10 shrub and forest habitat types, and two riparian habitat types (Appendix C). The six
Coastal Prairie habitat types distinguished by cluster analysis could not be delineated on aerial photographs; therefore, they were treated as one habitat type in habitat use analysis. Grass and forb cover was
sparse along Usal Creek and Wages Creek which were subjected to periodic flooding.
Habitat Use
Habitat use was nonrandom for all herds in each season (Tables
4-7). The breakdown of percentage of seasonal observations for herds in each habitat type was: grassland, 51-100 percent; forest and shrubland,
2-26 percent; and riparian, 1-26 percent. Figure 3. One-hundred (----), 90 (--) and 50 (....) Percent Home Range Estimates (Convex Polygon), with Number of Observations Used in the 100 Percent Estimate, of Bull #201RW, January-December 1983. Figure 4. One-hundred (-), 90 (--), and 50 (****)Percent Home Range Estimates (Convex Polygon), with Number of Observations Used in the 100 Percent Estimate, of the Red Herd (Cows #226R and #233B), January-December 1983. Figure 5. One-hundred (---), 90 (--), and 50 (....) Percent Home Range Estimates (Convex Polygon), with Number of Observations Used in the 100 Percent Estimate, of Cow #231W, January-December 1983. Figure 6. One-hundred (-..m), 90 (--), and 50 (****) Percent Home Range Estimates (Convex Polygon), with Number of Observations Used in the 100 Percent Estimate, of the Orange Herd (Cows #2290, #236RWB, #237Y, and #2400, January-December 1983. Table 3. Total Hectares and Total Number of Hourly Animal Locations by Activity for Each Habitat Type Over the Study Area, Humboldt and Mendocino Counties, California January-December 1983.
Habitat Type Area Number of Hourly (ha) Animal Locations by Activity
Active Inactive
Cultivated/disturbed grassland 42 137 172 Coastal Prairie 3218 550 781 Riparian shrub/grass 96 36 35 Riparian tree/shrub/grass 502 37 70 Upland shrub 570 3 5 Chaparral (manzanita/chinkapin) 549 Chaparral (blueblossom) 548 3 Douglas-fir: low density 77 moderate density 382 31 28 high density 46 5 5 Mixed Conifer: low density 636 9 5 3 moderate density 15212 28 26 high density 3142 1 8 Clearcut 487 8 5 8 307 8 Hardwood b Bare Ground a 872 178b 1
Totals 36 810 866 1150
aIncludes sand, however no attempt was made to estimate areas covered by asphalt and dirt roads.
bIncludes locations on asphalt and dirt roads. Table 4. Habitat Use by Bull #201RW in Humboldt County, California, Winter-Fall 1983. Percentage of Animal Observations in each Habitat Type Was Greater Than (>), Less Than (<), or Not Significantly Different from (=) Expected Use. Asterisks (*) Denote Expected Values That Were Combined in Chi-square Analysis. Chi-square values and their Respective Significances (p <) Are Shown for Comparisons within each Season.
% Observed Use: X Expected Use By Season
Habitat Type Winter Spring Summer Fall
* * Cultivated Grassland 41:16 39:2 * 31:1 43:13 > > > >
Coastal Prairie 59:84 44:60 31:39 57:86 < < = < * * Riparian Shrub/Grass 6:1 11:0.5 > > * * Riparian Tree/Sh./Gr. 3:7 25:5 < >
Moderate-Density 7:19 Douglas-fir <
Low-Density 1:33 Mixed-Conifer <
Moderate-Density 1:23 Mixed-Conifer <
Bare Ground 1:12 <
Chi-square 24.25 269.27 532.45 56.79 P < 0.001 0.001 0.001 0.001 Table 5. Habitat Use by the Red Herd (Cows #226R and #233B) in Humboldt and Mendocino Counties, California, Winter-Fall 1983. Percent of Animal Observations in each Habitat Type Was Greater Than (>), Less Than (<), or Not Significantly Different From (=) Expected Use. Asterisks (*) Denote Expected Values That Were Combined in Chi-square Analysis. Chi-square Values and their Respective Significances (p <) Are Shown for Comparisons within each Season.
X Observed Use: % Expected Use By Season
Habitat Type Winter Spring Summer Fall
* Cultivated Grassland 27:16 8:0.5 > >
Coastal Prairie 63:82 76:39 43:11 61:7 < > > > * Riparian Shrub/Grass 10:2 5:0.5 > > * Riparian Tree/Sh./Gr. 5:4 17: 2 * 13:2* > > ** Upland Shrub 2:34 0.5:1 < =
Hardwood 3:11 4:2** < = * * Moderate- and High- 14:12 1.5:2 1:1 Density Douglas-fir = = =
Low- and Moderate- 13:63 16:72 Density Mixed-Conifer < <
High Density 2:11 4:16 Mixed Conifer < < * Clearcut/Bare Ground 6:7 5:2 = =
Chi-square 30.74 46.39 954.53 767.19 p < 0.001 0.001 0.001 0.001 Table 6. Habitat Use by Cow #231W in Humboldt and Mendicino Counties, California, Winter-Fall 1983. Percentage of Animal Observations in each Habitat Type Was Greater Than (>), Less Than (<), or Not Significantly Different From (=) Expected Use. Asterisks (*) Denote Expected Values That Were Combined in Chi-square Analysis. Chi-square Values and their Respective Significances Are Shown for Comparisons within each Season.
% Observed Use: % Expected Use By Season
Habitat Type Winter Spring Summer Fall
* Cultivated Grassland 49:16 1:2.5 > =
Coastal Prairie 49:82 94:83 77:78 92:91 < = = = * * Riparian Shrub/Grass 2:2 3:3:11 * 6:0.4 4:0.5 = = > * * Riparian Tree/Sh./Gr. 1:5 3:6 < =
Low-Density 3:10 Mixed-Conifer <
Moderate-Density 13:6 Mixed-Conifer > * Bare Ground 3:16 =
Chi-square 35.82 2.63 12.60 0.03 p < 0.001 0.105 0.002 0.856 Table 7. Habitat Use of the Orange Herd (Cows #2290, #236RWB, #237Y, and #240G) in Humboldt and Mendocino Counties, California, Winter-Fall 1983. Percentage of Animals Observations in each Habitat Type Was Greater Than (>), Less Than (<), or Not Significantly Different From (=) Expected Use. Asterisks (*) Denote Expected Values That Were Combined in Chi-square Analysis. Chi-square Values and their Respective Significances (p <) Are Shown for Comparisons within each Season.
% Observed Use: % Expected Use By Season
Habitat Type Winter Spring Summer Fall
* Cultivated Grasslands 38:16 1:1:11 * 3:0.3 > = >
Coastal Prairie 61:82 75:41 79:79 95:82 < > = > * * Riparian Shrub/Grass 1:2 6:0.3 1:0.5 = > = * Riparian Tree/Sh./Gr. 5:3 =
Upland Shrub 2:42 2:13 1:13 < < <
Hardwood 3:0.2** > * Moderate-Density 17:13 2:0.5** 0.4:1 Douglas-fir = = =
High-Density 6:4 Douglas-fir = ** * Bare Ground 2:3 0.4:3 = <
Chi-square 43.44 14.92 51.77 36.71 P < 0.001 0.001 0.001 0.001 32
On a yearly basis, animals were more active than expected (total active-inactive observations) in moderate-density forest types (X 2 =
3.95, p < 0.05) and bare ground areas (X 2 = 19.47, p < 0.001). Animal activity in all other habitat types was not significantly different from the expected value.
Use of Grassland Habitats
Ninety percent of available cultivated grasslands occurred in the
Shelter Cove area. Use of cultivated grasslands by bull #201RW was greater than expected in each season (Table 4). Female herds used cultivated grasslands more than expected in the Shelter Cove area, but not in other home ranges (Tables 4-7).
Coastal Prairie use varied seasonally for each herd (Tables 4-7).
Use was less than that available in the Shelter Cove area. Use was greater than that available in the Chemise Creek area (by all females) and the Usal Creek area (by the red herd). Coastal Prairie use in the
Westport and Tenmile River areas was not significantly different from its availability to the white and orange herds.
Use of Shrub and Forest Habitats
Use of shrubland habitats was less than that available in the
Chemise Creek area (Tables 4-7). Shrubland use was less than expected or did not occur in other areas.
Spring and summer use of forest habitat types by bull #201RW in the Shelter Cove area was significantly less than expected (Table 4).
Female use of forest types increased during the spring (Tables 4-7).
Use of forest types in the Chemise Creek area was not significantly different from that available. Forest habitat use during summer and 33
fall by the red herd increased in the Usal Creek area; however, use was
significantly lower than that available in both seasons. Forest habitat
use in the Westport area was limited to the summer. The white herd used
low-density mixed-conifer areas less than expected and moderate-density mixed-conifer areas more than expected in the Westport area.
Use of Riparian Habitats
Riparian habitat use by bull #201RW was greater than expected in
the Shelter Cove and Bear Creek areas during the summer (Table 4). The red herd used riparian areas in the Usal Creek area more than expected
(Table 5). Use of riparian shrub/grass areas in the Westport and
Tenmile River areas was greater than expected, while use of riparian tree/shrub/grass areas was equal or less than expected (Tables 6 and 7).
Distance to Road
Elk were observed generally within 420 m of a road in 1983 (Table
8). Animal distances to roads were shortest (9-15 m average) in the
Shelter Cove area and were greatest in the spring when the females were in the Chemise Creek area. Road densities varied from 6.5 km of road /km2 in the Shelter Cove area to less than 1.0 km of road/km 2 in the
Chemise Creek area.
Distance to Water
Elk were observed 95 percent of the time within 420 m of water in
1983 (Table 9). Animal distances to water were greatest for all females in the Chemise Creek area during the spring. Animals were nearest water in the Shelter Cove area during the winter (bull #201RW and all females) and summer (bull #201RW). Distances to water during summer and fall
Table 8. Yearly and Seasonal Breakdown of Animal Distances to Road for Translocated Roosevelt Elk in Humboldt and Mendocino Counties, California, Winter-Fall 1983. Percentage of Observations in Each Distance Interval (m) Are for All Herds Combined. Seasonal Differences Between Observed Distances to Road for Each Herd Were Determined Using the Scheffe Test. Seasons (Wi=Winter, Sp=Spring, Su=Summer, Fa=Fall) That Were Not Significantly Different from Each Other Are Shown with a Common Underline. An Average Distance (X) Was Determined for Each Season.
Distance Interval (m) = Percent Observations
0 = 1 1-6 =4 6-15 = 19 15-30 = 30 30-60 = 22 60-120 = 8 120-240 = 4 240-420 = 2 420-950 = 10
Herd Season (Average Distance to Road)
Male Herd Wi Fa Su Sp
1 11 12 15 27
Red Herd Wi Fa Su Sp
I 12 26 29 192
White Herd Wi Su Fa Sp
X 11 51 54 156
Orange Herd Wi Su Fa Sp
x 9 26 29 312 Table 9. Yearly and Seasonal Breakdown of Animal Distances to Water for Translocated Roosevelt Elk in Humboldt and Mendocino Counties, California, Winter-Fall 1983. Percentage of Observations in Each Distance Interval (m) Are for All Herds Combined. Seasonal Differences Between Observed Distances to Water for Each Herd Were Determined Using the Scheffe Test. Seasons (Wi=Winter, Sp=Spring, Su=Summer, Fa=Fall) That Were Not Significantly Different from Each Other Are Shown with a Common Underline. An Average Distance (x) Was Determined for Each Season.
Distance Interval (m) = Percent Observations
0 = 1 1-6 = 7 6-15 = 17 15-30 = 20 30-60 = 16 60-120 = 11 120-240 = 8 240-420 = 19 420-950 = 1
Herd Season (Average Distance to Water)
Male Herd Su Wi Sp Fa
2 21 24 26 96
Red Herd Wi Su Fa Sp
1 18 27 42 102
White Herd Wi Su Fa Sp
1E 21 29 36 84
Orange Herd Wi Fa Su Sp
1 27 36 48 192 36
were intermediate to those observed in winter and spring and varied with
herd and location.
Distance to Cover
Elk were observed 14 percent of the time in cover and 78 percent
of the time within 30 m of cover in 1983 (Table 10). Animals were
nearest cover in the spring (all females in Chemise Creek area) and
summer (all animals). Animals were observed farthest from cover during
the winter (all females in the Shelter Cove area) and the fall (bull
#201RW in the Shelter Cove area). Animals were not observed moving to
cover in response to human activity.
Slope
Animals were observed 78.6 percent of the time on slopes of 15 percent or less, 17.5 percent on slopes between 16 and 30 percent, 3.4 percent on slopes between 31 and 45 percent, and 0.5 percent on slopes greater than 45 percent in 1983. Slope use was significantly different
(p < 0.001) from availability in both grassland and forest habitat types
(Table 11). Animal activity was negatively correlated with slope
(Kendall t = -0.11 to -0.26, p < 0.05) for both sexes (except for cows
#236RWB and #240G). Animals bedded primarily on slopes less than 15 percent, but were active on slopes as steep as 75 percent.
Climate
Habitat use during the winter was restricted to grassland and riparian shrub/grass habitat types; therefore, correlations between climate and habitat use were not determined. Habitat use during spring, summer, and fall was correlated with climatic variables (Table 12). Use
Table 10. Yearly and Seasonal Breakdown of Animal Distances to Cover for Translocated Roosevelt Elk in Humboldt and Mendocino Countiers, California, Winter-Fall 1983. Percentage of Observations in Each Distance Interval (m) Are for All Herds Combined. Seasonal Differences Between Observed Distances to Cover for Each Herd Were Determined Using the Scheffe Test. Seasons (Wi=Winter, Sp=Spring, Su=Summer, Fa=Fall) That Were Not Significantly Different from Each Other Are Shown with a Common Underline. An Average Distance (R) Was Determined for Each Season.
Distance Interval (m) = Percent Observations
0 = 14 1-6 = 21 6-15 = 34 15-30 = 23 30-60 = 7 60-120 = 1
Herd Season (Average Distance to Cover)
Male Herd Su Sp Wi Fa
R 4 6 11 15
Red Herd Su Sp Fa Wi
1 3 3 3 10
White Herd Sp Su Wi Fa
27 3 7 10 10
Orange Herd Su Sp Fa Wi
X 5 5 6 8 Table 11. Slope Use by Translocated Roosevelt Elk in Humboldt and Mendocino Counties, California, January-December 1983. Percentage of Observed Slope Use Was Greater Than (>), Less Than (<), or Not Significantly Different From (=) Expected Use.
%Observed Use : % Expected Use by Habitat Type
Percent Grassland Forest and Shrub Slope
0 - 15 % 81:65 27:3 > >
16 - 30 % 15:12 49:8 = >
31 - 45 % 3:15 20:20 < =
46 - 60 % 0.5:6 3:24 < <
61 + % 0.5:2 1:45 = < Table 12. Correlations Between Habitat Type Used (Grassland, Riparian, and Forest) and Climatic Variables (Ambient Temperature, Cloud Cover, Precipitation, and Wind Speed) for Translocated Roosevelt Elk in Humboldt and Mendocino Counties, California, Spring-Fall 1983. Kendall Rank Correlation Values (0 and Significance Levels (p) were Computed for All Collared Animals During Each Season.
Climatic Variable Season
(Kendall 1) k n < Spring Summer Fall
Temperature 0.3152 0.3072 0.0557 0.001 0.001 0.196
Cloud Cover -0.3080 -0.1983 -0.0931 0.001 0.001 0.014
Precipitation -0.0542 -0.0615 -0.1436 0.356 0.170 0.001
Wind Speed 0.0272 0.0872 -0.1334 0.560 0.025 0.001 40 of forest types increased during spring and summer as ambient temperature increased and percentage cloud cover decreased. Use of forest types was only weakly correlated with decreased cloud cover during fall (Table 12). Use of grasslands increased during fall as wind speed and precipitation increased.
Activity Pattern
The proportion of time spent active versus inactive during the eight three-hour time intervals in a 24-hour period of observation
(24-hour activity pattern) was significantly different between seasons
(X2 = 50.4, p < 0.001) and within each season (X 2 = 33.9, p < 0.001) for all individuals combined (Figure 7). The activity pattern was generally diurnal with longer periods of activity (about two hours in duration) starting at sunrise and 1.0 hour before sunset and shorter periods of activity (about one hour in duration) at night and during the day.
Time Spent Active
The proportion of time spent active and inactive decreased significantly (X2 = 10.7, p < 0.02) from winter to fall. Animals were active 43 percent of the time (n = 11 24-hour activity patterns with three or less missing observations) in the winter, 42 percent of the time (n = 11) in the spring, 40 percent of the time (n = 23) in the summer, and 36 percent of the time (n = 25) in the fall (Figure 7).
Animals were more active in close proximity to roads (p < 0.05) and when disturbed by humans (p < 0.05). No significant relationships existed between activity and habitat type, distance to water, distance 41
Figure 7. Percent Time Collared Roosevelt Elk Were Observed Active during each Hour, Humboldt and Mendocino Counties, California, Winter (A), Spring (B), Summer (C), and Fall (D) 1983. Ranges of Time of Sunrise and Time of Sunset Are Shown for each Season. 42 to cover, temperature, wind speed, percentage cloud cover, and precipitation.
Specific Behaviors
When specific active behaviors could be determined during the year, animals spent an average of 72.8 percent (SD = 4.47) of the time grazing, 4.3 percent (SD = 3.02) browsing, 6.5 percent (SD = 1.45) walking, 12.1 percent (SD = 2.84) standing, 2.6 percent (SD = 2.0) grooming, and 2.2 percent (SD = 1.95) in other activities (walking- and standing-interacting, walking- and standing-ruminating, and deficating)
(Figure 8).
When specific inactive behaviors could be determined during the year, animals spent an average of 53.4 percent (SD = 4.42) of the time bedded, 33.8 percent (SD = 5.29) bedded-ruminating, 7.4 percent (SD =
5.32) sleeping, 3.0 percent (SD = 1.21) bedded-grooming, and 2.2 percent
(SD = 1.11) bedded-feeding (Figure 9). Figure 9. Monthly Breakdown of Specific Inactive Behaviors Based on 5-Minute Scan Data for Translocated Roosevelt Elk, Humboldt and Mendocino Counties, California, January-December 1983. Figure 8. Monthly Breakdown of Specific Active Behaviors Based on 5-Minute Scan Data for Translocated Roosevelt Elk, Humboldt and Mendocino Counties, California, January-December 1983. DISCUSSION
Movements and Home Ranges
Animal movements have been classified as exploration, migration,
or dispersion depending on their distance and duration (Sanderson 1966).
The type of movement, the distance moved, and the stimulus to move are
regulated by both internal (physiological) and external (habitat and
climatic) factors.
The translocated elk made exploratory, migratory, and dispersal
movements during the study. The initial movements to the Shelter Cove
and Chemise Creek areas by all animals, movements by the red herd at
Usal Creek, and movements by bull #201RW during the rut were probably
exploratory movements (less than 14 km). The return of bull #201RW and
the red herd to Shelter Cove during the spring and summer respectively were migratory movements. The white and orange herds made dispersal movements to the Westport and Tenmile River areas. The red herd
returned to the enclosure area sometime during 1984.
The movement of all animals from the Shelter Cove area to the
Chemise Creek area coincided with the loss of antlers by bull #201RW.
Cow #226R apparently became the group leader at the onset of antler
drop. The Chemise Creek area provided an isolated area prior to the
calving period.
Females and older males (bulls #201RW and #243) were in separate herds in May through June and October through December. Franklin et al.
45 46
(1975) found that bulls older than two or three years were generally not
associated with cow herds except during the rut. Georgii and Schröder
(1983) also found that seasonal separation of sexes in red deer was age
dependent.
Annual grasses were a major component of the Chemise Creek meadow (Appendix C). A change in phenology of plants (especially
annuals) may have led to a reduction in food availability that caused
the female herd to divide into two smaller herds and abandon the Chemise
Creek meadow. Clutton-Brock et al. (1982) reported that social ties within female groups became strained when forage became scarce. Group
sizes varied seasonally with food availability in Tule elk (McCullough
1969) and impala (Jarman and Jarman 1973, Waser and Jones 1983). Cow herds at Gold Bluffs Beach were most stable November through May with divisions of cow herds increasing during calving, post calving, and rutting (Franklin 1968).
Movements by bull #201RW in the Shelter Cove area and bull #202
in the Fort Bragg area increased during the rut. Male Roosevelt elk wandered as much as 40 km from their home ranges during the rutting season in southwestern Oregon (Harper 1964). Movements of male ungulates generally increased before and during the rut as efforts to locate females increased (Dasmann and Taber 1956, Van Ballenberghe and
Peek 1971, Phillips et al. 1973, Grubb and Jewel 1974, Leslie and
Douglas 1979). Although both bull #201RW and bull #202 encountered female groups during their wanderings, only bull #202 was able to maintain a harem. The combination of the relative dominance of cow
#226R and the inexperience of bull #201RW may have resulted in his failure to hold the red herd during the rut. Lowe (1966) found that 47 larger or older red deer females appeared to dominate all other classes of either sex.
In mammals, females are less likely to disperse than males
(Greenwood 1980). Bull #202 dispersed with the female group from the
Chemise Creek area to the Westport area. However, the movement in
September of the red herd, without an adult male, was unusual because it coincided with the rutting period of bull #201RW. Possible reasons for female dispersion include intraspecific crowding (Howard 1960) and increased frequency of encountering members of the opposite sex (Bunnell and Harestad 1983, Waser and Jones 1983).
Bulls in the Shelter Cove area returned to a former home range and formed a bachelor herd after the rut. Franklin et al. (1975) found that older bulls were rarely solitary except during the rut. Male deer and moose returned to their traditional home ranges when sexual competition was reduced in the post rut period (Hamann and Taber 1956,
Phillips et al. 1973, Kammermeyer and Marchinton 1976).
Home range estimates based on the minimum area method are affected by a variety of biases including: 1) the area estimated depends on the definition of home range being used; 2) the area estimated varies with sample size (sample-size bias) (Anderson 1982, Laundre and Keller
1984); and 3) the area estimated varies with the grid size chosen (Mohr
1947). These biases became evident in comparisons of seasonal and monthly home range estimates. The summer home ranges for some herds were actually the combined area of two disjunct home ranges and the travel corridor between them. Home ranges calculated in this study represented a variety of home range defintions. The one-hundred-percent home range areas corresponded with Walther's (1972) definition of action 48
area in that they included travel corridors and explorations, but they were calculated on a monthly and seasonal basis rather than over an
individual's lifetime. The ninety-percent home range areas corresponded with Burt's (1943) classic definition of home range as "that area traversed by an animal during its normal activities of food gathering, mating, and caring for young." The fifty-percent home range areas represented the core area defined by Kaufmann (1962) as "that area used more frequently than any other." Sample-size bias became a factor from
August through December for small herds (i.e. male herd, white herd) that were sampled only once a month. Differences in monthly estimates of home range areas during the fall may have been the result of sample-size bias or differences in habitat patchiness between use areas.
Usal Creek and Tenmile River had large patches of homogenous habitat,
Westport had moderate patches of homogenous habitat, while only small patches of homogenous habitat existed at Shelter Cove. Grid size constraints are a problem primarily in heterogenous habitats (Johnson
1982). Habitat types within home ranges were generally homogenous and grid sizes were small enough that animals could easily traverse them between sample periods. Seasonal and monthly comparisons of home range sizes in this study primarily reflected animal movements. Home ranges were largest during July, August, and September when movement within and between home ranges was greatest. Interactions between habitat productivity, climate, and herd size also may have influenced home range size during the study. 49
Habitat Use
Roosevelt elk probably evolved in a stable habitat and mild climate, occupying patchy old—growth conifer forests and alluvial areas dominated by grasses and deciduous forests (Starkey et al. 1982).
Therefore, when animals are translocated, their closed program (Mayr
1974) responses to environmental features and group traditions should result in use of areas that closely resemble traditional home ranges.
Habitat use by Roosevelt elk at Gold Bluffs beach was greatest in the
Coastal Prairie community (specifically the perennial grass and forb type) (Franklin et al. 1975, Bowyer 1981). Therefore, animals translocated from Gold Bluffs Beach were expected to use primarily
Coastal Prairie habitats with use of other habitat types restricted to special needs.
Use of Grassland Habitats
Grasslands were the most used physiognomic group throughout the year. Elk food habits (Church and Hines 1978) and social systems (Geist
1982) were adapted to continuous, open grassland habitats. Cluster analysis results showed a variation in plant species composition between different grassland areas. Seasonal changes in home range use may have been the result of animals using different components of the Coastal
Prairie type to meet different nutritional or physiological requirements.
Cultivated grasslands (i.e. golf courses) represented a habitat type not previously encountered by the translocated elk. The elk were probably able to graze more efficiently on such cultivated grasslands because vegetation litter was removed by mowing. 50
Use of Shrub and Forest Habitats Shrubland habitat types were used in proportions less than their availability. Shrublands were characterized by dense shrub overstories and limited grass and forb cover. Restricted mobility and lack of forage probably limited elk use of shrubland types.
Forest habitat types were used in proportions equal to or less than their availability. Douglas-fir and mixed-conifer stands of moderate density were probably used the most because they had the greatest grass and forb cover. Animals were more active than expected in moderate-density forest habitats, suggesting that those habitats provided important sources of forage in addition to thermal cover.
Forest habitat types with at least 50 percent canopy closure were important sources of thermal cover (Black et al. 1976) during spring and summer. Low-density and moderate-density forest types were used primarily for thermal cover. These forest types were characterized by less than 50 percent combined grass and forb cover, whereas, adjacent grasslands generally had greater than 90 percent combined grass and forb cover.
Use of Riparian Habitats
Use of riparian areas varied with season and herd; however, all herds used riparian habitats during the summer in proportions greater than their availability. Riparian areas provided green forage during drier months. Thomas (1975), Marcum (1976) and Pederson et al. (1980) found that riparian areas provided necessary forage and thermal cover during summer and early fall. Red alder dominated riparian areas in the 51
Usal Creek and Westport areas were used primarily for thermal cover because the sparse grass and forb cover provided limited forage.
Distance to Road
Rost and Bailey (1979) and Starkey et al. (1982) reported that, because of human influence, elk and deer generally did not use areas adjacent to roads; however, Witmer and deCalesta (1983) stated that distances to roads were not important to Roosevelt elk in determining habitat use in the Oregon coastal range. Distances to roads were generally small because of the many roads in the study area, especially near Shelter Cove. The presence of roads should have had little influence on habitat use because the translocated animals had been conditioned to high levels of human influence at Gold Bluffs Beach
(Bowyer 1981). Although no females calved in 1983, females were farthest from roads during the spring coinciding with the normal calving period.
Roads in forested areas (Usal Creek) were used as travel corridors allowing animals to move quickly and with relative ease.
Overgrown skid roads were also used as foraging areas. Use of logging roads and skid trails as travel routes and foraging sites has also been reported for elk in Montana (Marcum 1975).
Distance to Water
Witmer and deCalesta (1983) reported that female elk were closest to water during calving season and were generally within 190 m of water throughout the year. Swanson (1970) found that Roosevelt elk habitat use was frequent within 400 m from water with greatest use 52 occurred within 100 m of water. Distances to water were greatest during spring in the Chemise Creek area indicating that forage availability, cover, or isolation from human influence were probably more important than water availability. Shorter distances to water in summer corresponded to the increased use of riparian areas to meet forage, cover, or water needs associated with an increase in temperature.
Distance to Cover
Availability of cover for escape from predators and human influence was an important factor in determining elk habitat use in many areas (Pedersen 1976, Irwin and Peek 1979, Lyon 1979, Morgantini and
Hudson 1979, Waldrip and Shaw 1979). Lack of hunting pressure in the study area and habituation to human presence reduced the use of escape cover by the translocated elk.
Increased use of cover by elk during spring and summer has been related to calving (Witmer and deCalesta 1983), increased availability of green forage (Pedersen et al. 1980, Jenkins and Starkey 1984), and thermal cover (Pedersen et al. 1980). Throughout the year the elk spent
92 percent of their time within 30 m of cover. This was probably a function of the broad definition of cover used in this study. Animals were usually in heterogenous habitats where shrubs or trees were well dispersed in grasslands. Shorter distances to cover during spring and summer reflected increased use of cover to avoid increased temperatures and reduced cloud cover.
Slope
Slope use was similar to that reported for Roosevelt elk (Witmer and deCalesta 1983) and Rocky Mountain elk (Julander and Jeffrey 1964, 53
Mackie 1970, Hershey and Leege 1982). Use of relatively flat areas reduced energy expenditures involved in obtaining nutrients (Belovsky
1981). Use of steeper slopes in the Chemise Creek and Usal Creek areas suggested that percentage slope may not have been an overriding factor in determining habitat use, as long as relatively flat bedding sites were readily available.
Climate
Elk in the study area responded to the combination of decreased cloud cover and increased ambient temperature by using forest and riparian habitat types for thermal cover. Changes in habitat use patterns of elk in response to thermoregulatory needs during the summer have been described in several studies (Swanson 1970, Beall 1976, Marcum
1976, Pedersen 1976, Collins and Urness 1983).
Swift et al. (1980) stated that energetic demands were greatest for elk during winter. Red deer generally utilized shelter during periods of low temperatures or high winds (Staines 1976, Clutton-Brock et al. 1982). The translocated elk were primarily in open grasslands during periods of low temperatures, high winds, or precipitation suggesting that winter conditions in the study area were not stressful.
Activity Pattern
Daily active-inactive cycles of animals are controlled by exogenous factors (Montgomery 1963, Erriksson et al. 1981, Georgii and
Schroder 1983), endogenous factors (Aschoff 1966), or a combination of both (Ozoga and Verme 1970, Jarman and Jarman 1973, Georgii 1981,
Grobler 1981). Daan and Aschoff (1975) found that the onset of activity 54
was usually more precise than activity cessation; and, in the course of
the year, tended to stay closer to sunrise in day-active animals. The
onset of the morning and evening activity periods was probably related
to sunrise and sunset. Morning peaks usually occurred about 0.5 to 1.0
hours after sunrise. Evening peaks usually occurred 1.0 hour after
sunset. The onset of daytime and night-time activity periods was
probably related to the amount of food ingested and digested in the
previous active-inactive cycle.
Time Spent Active
Owen-Smith (1982) noted a general decrease in feeding time with
a decrease in body size in ungulates. Similar differences within and
between species have been attributed to seasonal variations in habitat
quality (Ellis and Travis 1975) and changes in animal physiology (Dunbar
1979). Relatively high habitat quality and lack of major physiological
demands such as calving or breeding may have explained the comparatively
low amount of activity relative to body weight by elk in this study
(Figure 10).
The elk were most active during the winter and least active
during the fall. Seasonal variations in activity levels indicated responses to changes in habitat quality and animal behavior. Georgii
(1981) found that winter activity of red deer was reduced to 50 percent of the summer activity level, while most other studies reported similar, but less dramatic decreases in winter activity of ungulates (Skogland
1972, Thompson et al. 1973, Yahner 1980, Erriksson et al. 1981,
Clutton-Brock et al. 1982, Georgii and Schröder 1983, Chao and Brown
1984). Increased feeding in winter has been reported for roe deer 55
Figure 10. Percent Time Spent Active (during a 24—Hour Period) as a Function of Body Weight (log W(kg)) of Several Herbivores. Species Shown Include: a = Klipspringer (Oreotragus oreotragus), Grobler (1981); b = Impala (Aepyceros melampus), Jarman and Jarman (1973); c = Pronghorn Antelope (Antilocapra americana), Ellis and Travis (1975); d = Red Deer (Female) (Cervus elaphus elaphus), e = Red Deer (Male), Clutton—Brock et al. (1982); f = Wildebeest (Connochaetes taurinus), Berry et al. (1982); g = Roosevelt Elk (C. e. roosevelti), THIS STUDY; h = Domestic Cow (Bos taurus), (Loxodonta Ellis and Travis (1975); African Elephant africana), Wyatt and Eltringham (1974). 56
(Turner 1979), soay sheep (Grubb and Jewell 1974), and southern white-tailed deer and domestic animals (Short et al. 1969). Wheaton and
Brown (1983) cited inherited adaptations to winter food shortages, decreased metabolic requirements, endocrine level changes, and reduced day length as possible causes for decreased winter food intake. Short et al. (1969) stated that forage quality and climatic variables
(specifically temperature) were responsible for fluctuations in forage intake. Seasonal changes in activity should have been minimal because of mild climatic conditions and abundant food. There are several possible explanations for the observed decrease in activity from winter through fall. These include: 1) time required for the animals to become habituated to a new environment after translocation and release from the enclosure; 2) the animals were in poor to fair condition prior to translocation and may have been taking advantage of the resources at
Shelter Cove to improve their condition during the winter; 3) amount of activity may have been correlated with herd size; 4) high quality habitat at Shelter Cove allowed increased activity; or 5) changes in food quality between different home ranges may have affected the ratio of feeding time to rumination time.
Time Allocation to Specific Activities
The amount of time spent in specific active behaviors was a function of habitat type and level of human influence. A guaranteed food source and lack of human influence allowed Rocky Mountain elk to spend a high proportion of time feeding when in an enclosure (Collins et al. 1978). The amount of time spent standing or alert increased as the level of human influence increased (Table 13). Vigilance by antelope Table 13. Comparison of Percentage Time Spent in Specific Active and Inactive Behaviors.
Species Active Behavior Inactive Behavior
Feed Walk Stand Groom Bed Ruminate Source
Impala 66-72 10-16 11-15 6-7 62-71 29-38 Jarman and Jarman 1973 Aepyceros melampus White-Tailed Deer LaGory et al. 1981 Odocoileus virginianus Indiana 50-60 13-20 20-33 Ohio 15-40 22-25 30-55 Mule Deer 68 32 Browman and Hudson 1957 Odocoileus hemionus Red Deer 90.3 3.9 5.8 32.2 67.8 Clutton-Brock et al. 1982 Cervus elaphus elaphus Rocky Mountain Elk 89 6 1 4 Collins et al. 1978 C. e. nelsoni Roosevelt Elk 77 6.5 12 2.5 66 34 THIS STUDY C. e. roosevelti Moose 71 21 Geist 1963 Alces alces 58
individuals increased as habitat openess and group size decreased
(Underwood 1982).
The increase in browse use from June through September
corresponded with the increased use of ripairian areas. Increased browse use may have allowed the animals to meet specific nutritional needs during that period. Nutritional values for different forage items peaked during different seasons of the year (Short 1975). Seasonal shifts in diet composition have been reported for other elk populations
(Harper 1962, Kufeld 1973, Hobbs et al. 1981).
Rumination time was greatest during January and February and coincided with the increased amount of feeding. Hanley (1982) stated that rumination time increased as the amount of dietary fiber increased.
The percentage of time spent ruminating is fairly constant for a variety of ungulates (Table 13) suggesting that animals may have been selective during feeding. The amount of time spent sleeping was greatest in May and June when the elk were in an isolated area. The recovery of bull
#201RW from a sparring injury accounted for 83 percent of the increase in time spent sleeping during October.
Conclusions
The use of an enclosure to enhance site fidelity in the translocated animals was only partially successful. Nine cows and one bull dispersed from the release site and did not return during the course of the study. Translocating animals habituated to human activity caused some problems. Use of areas adjacent to houses by the orange herd in the Tenmile River area was not compatible with use by local residents. Six animals were translocated back to the enclosure in 1983 59
and 1984 resulting in the death of one female. Animal movements and patterns of home range use were probably dependent on resource quantity
and quality. Because the elk used several different areas during the
study, follow-up studies of habitat-use patterns should indicate which areas are most important to the elk.
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APPENDIX A. Animal Capture.
Potential trap sites in Prairie Creek Redwoods State Park and
Redwood National Park were pre-baited with alfalfa hay, apples and prepared feed mixtures in early January 1982. Temporary traps were built on three sites (Figure 11) by National Park, State Park and
California Department of Fish and Game personnel during 15 to 20
February 1982. 69
Figure 11. Location of Prebaited Trap Sites in Prairie Creek Redwoods State Park and Redwood National Park, Humboldt County, California, February 1982. 70
APPENDIX B. Animal Movements.
Month Movements
Jan-Mar All animals were in a single herd in the Shelter Cove area.
Apr All animals had moved to the enclosure by April 2. They moved
through the south end of the enclosure on April 4 and were in
a meadow north of Chemise Creek by April 9. Bulls #201RW and
#243 were separated from the main herd by April 24.
May-Jun Bulls #201RW and #243 had returned to Shelter Cove by May 15.
The females and bulls #202 and #242 remained in the Chemise
Creek area.
Jun The female group split into the red and orange herds on June
11. The red herd (with bull #242) moved to the enclosure
where they remained for approximately two weeks. The orange
herd (with bull #202) moved south through Whale Gulch (Figure
3) and spent approximately two weeks in the area between Bear
Harbor and Jackass Creek. During this time, Cow #234 moved
back to Shelter Cove and had joined the red herd by June 20.
Jul-Aug The orange herd moved south to Howard Creek arriving sometime
before July 8. The females remained in the area between
Howard Creek and Wages Creek, while bull #202 moved farther
south. Bull #202 returned to the orange herd on July 23 and
the group moved south. The white herd became separated from
the orange herd in Westport (July 23). The white herd
remained in the area between Dehaven Creek and Wages Creek
through December. The orange herd moved as far south as 71
Pudding Creek during August, before settling in the area
between Bruhel Point and Tenmile River (August 25). Bull #202
was translocated to the enclosure in August.
Aug-Sep By August the red herd was in Shelter Cove with bulls #201RW,
#242, and #243. The red herd (without cow #234) separated
from the bulls on August 5 and had moved to the enclosure by
August 7. On August 28 the red herd (with cow #234 and bulls
#242 and #243) were at Bear Creek. Bull #201RW had also moved
to the Bear Creek area, but apparently did not join the red
herd.
Sep The red herd (without bulls #242 and #243) moved from Bear
Creek east to Whitethorn. During September the animals moved
south from Whitethorn and reached Usal Creek on September 11
travelling primarily on roads. The red herd remained
primarily in the Usal Creek area, but travelled east to Hales
Grove and south to Rockport on December 2. Bull #201RW
remained in the Bear Creek area until October 1; however, the
bull did move toward Whitethorn on September 4, but returned
to Bear Creek the same day. Bulls #242 and #243 returned to
Shelter Cove by approximately September 10.
Oct Bull #201RW returned to Shelter Cove by October 1 where all
three bulls remained through December.
1984 Four cows from the orange herd were translocated from the
Tenmile River area to the enclosure. One cow (#230) died
during the capture of the animals. The white herd moved south
and joined the lone female in the Tenmile River area. The red
herd eventually returned to the Bear Creek area. APPENDIX C. Habitat Sampling.
Table 14. Dominant Plant Species in the Seven Grassland Subgroups Determined Using Cluster Analysis, Humboldt and Mendocino Counties, California. Frequency (Number of Plot in Which a Species Was Found) and Average Cover Class (1 = solitary plant; 2 = few plants; 3 = <5%; 4 = 5-25%; 5 = 25-50%; 6 = 50-75%; and 7 = >75%) Are Given for the Most Frequently Occurring Species. Habitat Subgroups A-F Are Coastal Prairie Grasslands, Subgroup G Is the Cultivated Grassland. Locations of Habitat Subgroups Are: A = Shelter Cove; B = Shelter Cove/Bear Harbor/Westport; C = Bruhel Point/Westport; D = Enclosure/Whitethorn/Usal Creek; E = Tenmile River/Westport; F = Chemise Creek.
Species Common Name Habitat Subgroups
A B C D E F G
Holcus lanatus velvet grass 100:6 100:5 100:4 63:4 100:6 75:4 29:3 Danthonia californica California oatgrass 86:3 100:5 88:5 63:4 29:2 100:5 0 Cynosurus echinatus dogtail 14:4 14:3 0 88:5 0 100:6 0 Dactylis glomerata orchard grass 29:4 0 50:4 13:3 86:3 0 29:3 Lolium perenne perennial ryegrass 14:2 29:2 38:3 38:4 86:5 0 29:4 Elymus glaucus blue wild-rye 0 0 25:2 50:5 0 75:4 0 Festuca sp. fescue 0 0 0 0 0 0 100:5 Plantago lanceolata plantain 86:4 100:4 100:4 63:4 86:4 25:3 86:4 Rumex acetosella sheep sorrel 86:3 71:3 50:2 88:3 71:3 50:3 29:3 Cirsium arvense bull thistle 57:2 57:2 63:2 88:3 86:2 50:2 43:2 Pteridium aquilinum bracken fern 43:3 57:4 50:3 74:4 14:4 100:5 0 Trifolium sp. clover 43:2 29:2 13:2 50:3 43:3 0 86:6 Erectities prenanthoides fireweed 43:2 43:3 50:2 0 0 50:2 0 Taraxacum officinale common dandelion 43:2 43:3 38:2 75:4 57:2 75:3 100:4 annual 86:3 86:2 63:3 13:3 57:3 25:4 0 Baccharis pilularis coyote brush 25:1 71:3 100:3 13:2 0 100:3 0 Rubus vitifolius California blackberry 43:3 43:2 88:3 38:2 57:2 0 0 APPENDIX C. Habitat Sampling.
Table 15. Dominant Plant Species in the Three Shrub Dominated Subgroups Determined Using Cluster Analysis, Humboldt and Mendocino Counties, California. Frequency (Number of Plots in Which a Species Was Found) and Average Cover Class (1 = solitary plant; 2 = few plants; 3 = <5%; 4 = 5-25%; 5 = 25-50%; 6 = 50-75%; and 7 = >75%) Are Given for the Most Frequently Occurring Species. Habitat Subgroups Include: A = Ceanothus Dominated Shrubland; B = Chaparral Shrubland; C = North Coastal Scrub.
Species Common Name Habitat Subgroup (Freg:Cover Class)
A B C
Lithocarpus densiflorus tanoak 100:4 25:4 22:2 Ceanothus thyrsiflorus blueblossom 100:7 0 0 Corylus cornuta California hazel 17:3 0 56:6 Rubus vitifolius California blackberry 0 0 78:5 Baccharis pilularis coyote brush 0 0 100:5 Vaccinium ovatum evergreen huckleberry 50:3 100:4 11:3 Arctostaphylos columbiana hairy manazanita 0 100:6 0 Castanopsis chrysophylla golden chinkapin 0 100:4 0 Lupinus sp. lupine 0 0 56:4 Stachys rigida hedge-nettle 50:2 0 78:3 Fragaria chiloensis strawberry 0 0 67:3 Pteridium a.uilinum bracken fern 33:3 100:4 100:4 Holcus lanatus velvet grass 17:3 0 56:3
Percent Slope (average) 61 36 66 APPENDIX C. Habitat Sampling.
Table 16. Dominant Plants in the Seven Forest Subgroups Determined Using Cluster Analysis, Humboldt and Mendocino Counties, California. Frequency (Number of Plots in Which a Species Was Found) and Average Cover Class (1 = solitary plant; 2 = few plants; 3 = <5%; 4 = 5-25%; 5 = 25-50%; 6 = 50-75%; and 7 = >75%) Are Given for the Most Frequently Occurring Species. Habitat Subgroup Labels Are: A = Low- to Moderate-Density Douglas-fir; B = Moderate- to High-Density Douglas-fir; C = Low-Density Mixed-Conifer; D = Moderate- to High-Density Mixed-Conifer; E = Moderate-Density Mixed-Conifer (Selective Cut); F = High-Density Mixed-Conifer; G = Low-Density Mixed-Conifer (Clear Cut).
Species Common Name Habitat Subgroup
A B C D E F G
Pseudotsuga menziesii Douglas-fir (tree) 94:5 75:5 36:4 71:4 82:5 100:6 71:5 Pseudotsuga menziesii Douglas-fir (sapling) 88:4 100:3 54:3 43:4 35:3 50:2 86:4 Sequoia sempervirens coast redwood (tree) 18:4 25:2 57:4 100:5 82:5 50:5 43:3 Sequoia sempervirens coast redwood (sapling) 18:3 0 64:3 100:4 82:4 50:3 29:2 Lithocarpus densiflorus tanoak (tree) 88:6 100:4 96:5 95:5 29:3 25:3 71:5 Lithocarpus densiflorus tanoak (sapling) 94:4 100:5 100:6 100:4 71:4 25:4 71:6 Arbutus menziesii Pacific madrone (tree) 53:4 50:5 75:4 29:3 0 25:1 29:6 Arbutus menziesii Pacific madrone (sapling) 12:4 75:4 68:3 0 6:4 0 29:4 Vaccinium ovatum evergreen huckleberry 100:4 75:5 89:5 67:4 65:3 75:2 43:5 Rhododendron macrophyllum coast rhododendron 0 0 39:5 82:3 71:4 100:3 29:5 Pteridium aquilinum bracken fern 65:2 50:2 39:2 82:3 71:4 100:3 29:3 Polvstichum munitum sword fern 94:4 25:2 29:2 95:4 100:5 100:6 14:3 Sphagnum sp. sphagnum moss 41:3 75:3 25:3 71:3 18:3 25:3 43:3 Oxalis oregana redwood-sorrel 53:3 0 0 53:3 76:4 50:3 100:3 Gaultheria shallon salal 29:2 0 50:3 48:3 65:3 25:3 29:2 Holcus lanatus velvet grass 12:2 0 4:2 5:2 65:3 0 14:3
Percent slope 66 49 61 58 56 30 49 APPENDIX C. Habitat Sampling.
Table 17. Dominant Plant Species in the Two Riparian Subgroups Determined Using Cluster Analysis, Humboldt and Mendocino Counties, California. Frequency (Number of Plots in Which the Species Was Found) and Average Cover Class (1 = solitary plant; 2 = few plants; 3 = <5%; 4 = 5-25%; 5 = 25-50%; 6 = 50-75%; and 7 = >75%) Are Given for the Most Frequently Occurring Species.
Species Common Name Habitat Subgroup (Freg:Cover Class)
Shrub/ Tree/Shrub/ Grass Grass
Alnus rubra red alder 78:6 100:6 Pseudotsugs menziesii Douglas-fir 0 50:3 Sambucus sp. elderberry 22:2 62:4 Salix spp. willow 67:5 0 Rubus parviflorus thimbleberry 44:3 100:3 Rubus vitifolius California blackberry 89:5 25:4 Rubus thyrsanthus Himalaya berry 44:5 63:4 Scrophularia californica figwort 67:3 38:3 Cirsium arvense bull thistle 67:2 38:2 Equisetum sp. horsetail 56:4 100:4 Stachys rigida hedge-nettle 56:2 88:3 Urtica californica stinging nettle 11:4 75:3 Pteridium aquilinum bracken fern 66:3 38:4 Polystichum munitum sword fern 100:3 100:4 Holcus lanatus velvet grass 67:3 25:2 Juncus spp. juncus 78:3 50:4 Scirpus spp. bullrush 56:2 63:5