HABITAT USE BY FEMALE MULE DEER

IN NORTH CENTRAL CALIFORNIA

by

Terri A. Weist

A Thesis

Presented to

The Faculty of Humboldt State University

In Partial Fulfillment

of the Requirements for the Degree

Master of Science

August 1993 HABITAT USE BY FEMALE MULE DEER

IN NORTH CENTRAL CALIFORNIA

by

Terri A. Weist

Approved by the Master's Thesis Committee

Archie S. Mossman, Chairman Date

Date

Gerald M. Allen Date

Director, Natural Resources Graduate Program Date

93/W-273/05/31 Natural Resources Graduate Program Number

Approved by the Dean of Graduate Studies

Susan H. Bicknell Date ABSTRACT

Summer habitats used by 17 female mule deer (Odocoileus hemionus hemionus) near McCloud, California, were estimated using radio telemetry. Deer from two vegetatively distinct sub-regions within the study area displayed differences in home range behavior. Six out of 9 deer from the Porcupine Butte sub-region moved 3.7 to 19 km to second summer home ranges. In contrast, 7 out of 8 mule deer in the Hambone Butte sub-region occupied single home ranges throughout the summer. Home ranges estimated from minimum convex polygon method, averaged 197 ha for the 8 deer at the Hambone Butte sub-region and 130 ha for 9 deer at the Porcupine Butte sub-region for first home ranges. The mean pooled home range size (areas for each home range were added together into one overall home range for each deer that exhibited this behavior) for all deer within each sub-region was 221 ha. The difference in behavior patterns between the two sub-regions may have been in response to the habitat types available within each sub-region. The first home ranges of the 6 deer in the Porcupine Butte sub-region that moved to a second summer home range consisted of pine plantations and manzanita shrub habitats and later moved to habitats consisting of mixed conifer or the mixed trees/shrub habitat

iii types. Results from the Chi-Square analysis showed that deer occupying the Porcupine Butte sub-region selected tree plantations > 15 years old and avoided mixed conifer habitats, plantations < 15 years old and shrub habitats (when including dual home ranges together). The deer from Hambone Butte selected mixed conifer over other habitat types and avoided the mixed trees/shrub and shrub habitats. The Hambone Butte sub-region had significantly greater habitat diversity than the Porcupine Butte sub-region (P=0.035) based on the Shannon-Wiener index. The differences in the number of habitat edges within deer home ranges were also compared between sub- regions. The Hambone Butte sub-region had a significantly greater number of habitat edges than the Porcupine Butte sub-region (P=0.027). Vegetation characteristics did not differ significantly between plot types ("Use" plots and random plots) measured within sub-regions. However, four out of five variables differed significantly between sub-regions. Percent canopy cover was significantly greater at the Hambone Butte sub-region (P=0.029), while percent shrub cover (P=0.004), vegetation height (P=0.001) and percent low cover (P<0.001) were significantly greater at the Porcupine Butte sub-region. Deer movements from Porcupine Butte may indicate a deficiency in some needed habitat elements. Further

iv research is needed to determine what elements are missing from the Porcupine Butte sub-region so that land managers conducting habitat improvements to benefit deer populations could then incorporate this knowledge when designing their management strategies.

v ACKNOWLEDGEMENTS

I would like to thank the California Department of Fish & Game, particularly Tim Burton, for entrusting this opportunity to me. I am especially grateful for the patience they have displayed in waiting for this report. This project was funded by the California Department of Fish and Game Hill Bill project funds. Additional funds that helped get me through the school year came from a scholarship from the Marin Rod & Gun Club. Their support of graduate students is most appreciated. I am also grateful to the folks at the McCloud Ranger District for their cooperation in supplying me with aerial photos, compartment and topographic maps, information and housing. Those that were extremely helpful and important to this project are: Karen Austin and Debbie Selby, Ralph Phipps who provided me with information regarding plantations, Steve Clauson who provided the sorely needed amenities of housing and shower facilities. Also, thanks go to Larry Grant and Yvonne Studinski on the Modoc Ranger District, who provided the Medicine Lake aerial photos and the GIS maps of that area. In addition, I am grateful to Boyd Turner of the Hat Creek Ranger District for responding so quickly to my request for vegetative maps of a portion of their district (these deer got around!). Also to

vi vii

Arlene Angelides of the Supervisor's office in Redding for providing me with all the GIS maps of the McCloud study area. Thanks also go to Greg Schmidt, Jack Kahl and Lynn

Roberts at the Six Rivers National Forest for allowing me to use the digitizer at their office.

To my major professor, Dr. Archie Mossman, who took a big chance on me. His patience and guidance on this project helped me to get beyond the "statistical razzle dazzle" and try to understand why critters do what they do, which seems to be a never-ending process. To my committee members, R.J. Gutierrez and Jerry Allen whose criticisms and input were most appreciated.

To Lisha, my four-legged field companion, whose presence helped keep me from loneliness. And most of all, to my friend and confidant, Pat "Ranger" Ward. His constructive criticisms, advice, and dedication to the wildlife profession have truly been my inspiration. He has been the "wind beneath my wings."

Naturally, I have to acknowledge my study subjects, who epitomized the philosophy that for every question answered, two more questions will inevitably arise.

Finally, I would like to dedicate this thesis to my mother, Martha Van Ness. She may not have quite understood my calling in life, but advised me, nevertheless to "do my own thing." I regret she will never know that I finally completed this thesis. TABLE OF CONTENTS Page ABSTRACT iii ACKNOWLEDGEMENTS vi LIST OF TABLES x LIST OF FIGURES xiii INTRODUCTION 1 STUDY AREA 4 METHODS 8 Radio Telemetry 8 Home Range Analysis 12 Habitat Analysis 12 Vegetation Sampling 13 Habitat Diversity 16 Edge Effects 17 Elevation 19 RESULTS 20 Home Range 21 Habitat Analysis 24 Vegetation Analysis 28 Habitat Diversity 34 Edge Effects 38 Elevation 39 Deer Migration 39

viii ix

TABLE OF CONTENTS (CONTINUED) Page DISCUSSION 41 Home Range 41 Habitat Use 42 Vegetation Analysis 43 Habitat Diversity 43 Edge Effects 44 Deer Movements 45 REFERENCES CITED 53 APPENDIX A 57 APPENDIX B 71 APPENDIX C 72 APPENDIX D 73 APPENDIX E 75 LIST OF TABLES

Table Page 1 Number of deer on Hambone Butte and Porcupine Butte that occupied one vs two home ranges, Siskiyou County, California, 1987-1988 . . . 22 2 Summer home range sizes (ha) of female mule deer within each region in Siskiyou County, California, 1987-1988 (Hambone Butte: n=8; Porcupine Butte: n=9) 23 3 Chi-Square analysis of habitat selection using the combined home ranges of deer at Hambone and Porcupine Buttes, Siskiyou County, California, 1987-1988 26 4 Bonferroni confidence intervals for habitats selected by deer at Hambone Butte, Siskiyou County, California, 1987-1988 27 5 Bonferroni confidence intervals for habitats selected within combined home ranges by deer at Porcupine Butte, Siskiyou County, California, 1987-1988 29

x xi

LIST OF TABLES (CONTINUED) Page

6 Chi-Square analyses of habitat selection within

the first home ranges of the Porcupine Butte

deer that occupied two summer home ranges . . . 30

7 Bonferroni confidence intervals for habitats

selected within first home ranges by deer that

moved from Porcupine Butte, Siskiyou County,

California, 1987-1988 31

8 Chi-Square analyses of habitat selection within the

second home ranges of the Porcupine Butte

deer that occupied two summer home ranges

32

9 Bonferroni confidence intervals for habitats

selected within second home ranges by deer at

Porcupine Butte, Siskiyou County, California

1987-1988 33

10 Two-way ANOVA on vegetative characteristics

within habitats used by deer in Siskiyou County,

California, 1987-1988 35 xii

LIST OF TABLES (CONTINUED) Page

11 Means and standard errors of vegetation

characteristics sampled within home ranges of

deer at Hambone and Porcupine Buttes, Siskiyou

County, California, 1987-1988 36

12 Habitat diversity (H'), relative diversity (J')

and heterogeneity (1-J') indices within the

home ranges for deer on Hambone and Porcupine

Buttes, Siskiyou County, California, 1987-1988

37 LIST OF FIGURES

Figure Page

1 Location of the McCloud Study Area located

in Siskiyou County, California 5

2 Delineated sub-regions (Hambone Butte and

Porcupine Butte) within the McCloud Study

Area 6

3 Deer capture locations within the McCloud

Study Area 9

4 Frequency of use for each habitat type used by

deer at Hambone Butte and Porcupine Butte

within the McCloud Study Area 25

xiii INTRODUCTION

Rocky mountain mule deer (Odocoileus hemionus hemionus) and Columbian black-tailed deer (Odocoileus hemionus columbianus), as well as hybrids of the two sub- species, share their summer range on the McCloud Flats of northeastern California. In autumn, these populations migrate to nine distinct winter ranges (Ashcraft 1961).

Most deer researchers focus on winter range as the limiting factor in deer survival (Tierson et al. 1985).

Thus, the importance of the quantity and quality of summer range has been underestimated. In fact, the decline in reproductive rates for deer has been attributed to inadequate summer forage (Julander et al. 1961). High quality forage on summer range is essential to assure good body condition and a higher winter survival (Garrott et al.

1987). For example, the winter survival of moose was found to be dependent on the quality of the summer and fall diet

(Belovsky 1978). Deer require a variety of plant species in order to achieve the proper synergistic effects of a diverse diet (Church 1977, Mooty et al. 1987). Therefore, it is important for resource managers to discern those habitat characteristics that deer select. With this knowledge, managers can enhance the quality of summer range through vegetative management.

1 2 Home range size is determined by habitat condition and the distribution of such factors as food, cover and water and not by density of deer (Dasmann and Taber 1956). Movements within the home range are influenced by the distribution and the quantity and quality of these elements. Deer inhabiting more diverse areas maintain smaller home ranges which result in higher deer densities (Loft et al. 1984). Also, deer numbers fluctuate more in marginal habitat than on optimum ranges (Leckenby et al., 1982). High density mule deer populations normally occur in diverse and complex habitats where resources are abundant and their availability is relatively stable (Mackie 1983). Thus, habitat diversity appears to be an important element of mule deer habitat (Mackie 1983, Loft et al. 1984 and Wallmo 1978). Energy expenditure required to travel to areas with suitable habitat reduces the amount of energy available for reproduction and survival (Leckenby et al., 1982). Habitat evaluation is a form of carrying capacity assessment and should be a basic function in management of mule deer (Connolly and Wallmo 1981). To properly manage mule deer populations, it is necessary to determine habitats selected by deer. The purpose of this study was to aid in the management of the mule deer residing in northeastern California. 3

Consequently, the following primary (p: a priori) and secondary (s: a posteriori) objectives were undertaken:

p: 1. Estimate the summer home range size for

radio-collared female mule deer.

p: 2. Compare the home range sizes and habitats

used by the female mule deer between two

vegetatively distinct sub-regions.

p: 3. Identify the vegetative components within the

habitat types occupied by the female mule

deer.

p: 4. Describe summer movements of radio-collared

deer.

s: 5. Investigate differences in habitat diversity

between the Hambone Butte and Porcupine Butte

sub-regions. STUDY AREA

The study area is located about 113 km northeast of

Redding, California encompassing Township 40N and Range 2E to Township 43N and Range 3E (Fig. 1). There are two sections within the study area (referred to here as "sub- regions") that are easily distinguished from each other due to the dissimilar distribution of habitat types available in each sub-region. The Hambone Butte sub-region is approximately 227 km2 and the Porcupine Butte sub-region is approximately 322 km2 (Fig. 2). Elevations range from 1220 to 2130 m. Topography is generally flat with slopes less than 40%. Soils consist mostly of weathered volcanic rock and are very porous (McCloud Flats Deer Herd Management

Plan, Calif. Dept. Fish & Game Rept., 1985). The mean temperature for May-October 1988 was 27°C.

The Hambone Butte sub-region is less volcanic and slightly lower in elevation than the Porcupine Butte sub- region. The term 'volcanic' used here denotes the presence of lava flows with lava tubes, and ice caves. All recognized habitat types (Appendix A) occurred at Hambone

Butte, thus creating a greater habitat diversity than was present at Porcupine Butte. Habitat diversity in this context refers to beta diversity which is the number of different habitat communities occurring along an

4 5

Fig. 1. Location of the McCloud Study Area located in Siskiyou County, California. 6

Fig. 2. Delineated sub-regions (Hambone Butte and Porcupine Butte) within the McCloud Study Area. 7 environmental gradient (Kimmins 1987, Raphael 1992). By contrast, Porcupine Butte consists mostly of conifer plantations and manzanita shrub habitat types. Included within the Porcupine Butte sub-region is the "Lava Crack Springs" area which provides a greater number of habitat types but still maintains the volcanic component. Dominant tree species present on both study areas include ponderosa pine (Pinus ponderosa), Jeffrey pine

(Pinus jeffreyi), sugar pine (Pinus lambertiana), incense cedar (Calocedrus decurrens) and white fir (Abies concolor). Common shrub species include green leaf manzanita (Arctostaphylos patula), bittercherry (Prunus emarginata), bitterbrush (Purshia tridentata), serviceberry (Amelanchier alnifolia) and tobacco brush (Ceanothus velutinus). METHODS

Habitat use by female mule deer was studied by monitoring deer movements using radio telemetry. Vegetation was sampled within deer home ranges in order to estimate habitat characteristics at these locations.

Radio Telemetry

Twenty female mule deer were captured using drive nets near pre-selected water holes. They were ear-tagged and fitted with radio transmitters by California Department of Fish & Game personnel and volunteers. Two to four animals were captured and radio-collared in the vicinity of eight dispersed water sources to increase the likelihood of getting a representative sample of deer in the sub-regions (Fig. 3). The radio transmitters (159.275-159.466 MHz:

Telonics, Inc. Mesa, AZ) were equipped with mortality sensors. Radio collared deer were located using a hand-held, two element yagi ("H") antenna. Locations were obtained during July through August 1987 and May through November 1988. A minimum of three compass bearings were used to triangulate these locations and the locations were entered onto 7.5-minute topographic maps (1:24,000 scale). Only locations obtained within 15 minutes were considered reliable. The mean distance from telemetry stations (taken

8 9

Fig. 3. Deer capture locations within the McCloud Study Area. 10 from a random sample of deer locations) to the estimated location for deer from Hambone Butte was 0.42 km (n=42, s.e.=0.009) and 0.45 km (n=48, s.e.=0.014) for deer from the Porcupine Butte sub-region. Locations were recorded using the Universal Transverse Mercator grid system (Grubb and Eakle 1988).

Telemetry accuracy was estimated either by locating transmitters secretly placed at various locations throughout the study area by another person or by obtaining visual

confirmations of deer locations once triangulation was completed (a minimum of three bearings were used to locate transmitters). The true bearings were then compared with the estimated bearings (Springer 1979). The linear distance between the estimated location and the actual location was

measured on a map (1:24,000 scale) by using a ruler and converting the ruler distance to actual ground distance.

These distances were added together for all test cases and a

mean distance was calculated. A random-systematic approach was used to select the sub-region and the deer to be observed. The first sub-

region sampled was chosen by flipping a coin. Subsequent

coverage of the sub-regions alternated each day. For

example, if Hambone Butte was chosen for day one, then Porcupine Butte was observed the following day. Deer near Hambone Butte would be observed on day three and so forth. Thus, each doe usually was not located more than once every 11

24 hours in an attempt to maintain statistical independence between locations (Dunn and Gipson 1977).

Individual deer were identified by the frequency numbers of their radio collars. The first deer observed for a given day, was selected. The order of deer observed after the first chosen individual was based on spatial convenience

(that is, the deer in closest proximity to the deer preceding it). Locations that took longer than 15 minutes to obtain were eliminated from habitat selection analysis in order to minimize triangulation error (Amlaner and Macdonald

1980; Mech 1983).

The day was divided into six sampling periods:

TIME (24hr) SAMPLING PERIOD

0001-0400 1 0401-0800 2 0801-1200 3 1201-1600 4 1601-2000 5 2001-2400 6

The first sampling period during which deer were to

be observed, was chosen randomly using the random numbers generator on a scientific calculator ("Sharp" model EL-506).

To avoid observer fatigue, time periods were sampled in subsequent order following the first randomly chosen time

block. That is, if the first sampling period chosen was sampling period 3, then the following day would begin at sampling period 4. Observations for the third day would

begin at sampling period 5 and so on. Since habitat use was observed repeatedly throughout the summer and at any point 12 during a 24-hour day, I assumed that this method produced an unbiased estimate of habitat use.

Home Range Analysis

The minimum convex polygon was the home range estimator (Mohr 1947, Tinkle et al. 1962). Home range boundaries were digitized and the areas estimated using the program Designcad 4.0 (American Small Business Computers

Inc., 327 So. Mill St. Pryor, OK 74361).

An average estimate of home range size for all deer was calculated for each sub-region. To detect differences in home range size between the two sub-regions, a two-sample t-test (Zar 1984) was performed. In addition, the behavioral differences observed (i.e., dual home ranges) between deer in both sub-regions, was tested using a 2X2 contingency table to estimate whether this pattern was independent of sub-region. That is, did deer originating from a particular sub-region demonstrate the dual home range behavior more frequently than deer from the other sub- region?

Habitat Analysis

Seven habitat types were recognized for this study and each type is described in Appendix A. These habitat types are: Mixed conifer, mixed trees/shrub, shrub, plantations < 15 years old, plantations 15 years old, recent clearcut and ecotone. 13 Habitat boundaries were defined by using aerial photographs and USFS vegetation maps and were field- verified. Deer home ranges were marked on the USFS vegetation maps (1:15,840 scale), and a digitizer was used to input home range boundaries. The area of each habitat type within each deer's home range was estimated using the Designcad program. The proportion of each habitat type present within an individual's home range was calculated. The Chi-square goodness-of-fit test was used to test the null hypothesis that habitat types were used in proportion to their availability for deer in each sub-region

(Neu et al. 1974). The Bonferroni Z-statistic was used to calculate confidence intervals around the proportions of used habitats to estimate which habitat types were used more or less frequently than expected (Neu et al. 1974, Byers et al. 1984). This analysis was performed separately for

Hambone and Porcupine Buttes sample populations. In addition, the Chi-square and confidence intervals were calculated for both the first home ranges and second home ranges for deer in Porcupine Butte that utilized dual summer home ranges.

Vegetation Sampling

Vegetation sampling was conducted using circular plots (0.04 ha) (Noon 1981). The circular plots had a radius of 11.3 m. In order to detect vegetative characteristics that might influence deer in their selection 14 of habitat types, two types of vegetation plots were established: "Use" and "random." "Use" plots were measured at telemetry and visual deer locations. "Use" plots were selected among locations observed throughout the summer.

This was accomplished by selecting a minimum of two locations per deer per month. Preference was given to visual confirmations due to the accuracy of the location. Each individual deer's locations were numbered sequentially and those numbers were randomly selected as vegetation plot locations. "Random" plots were chosen from a list of UTM coordinates that were randomly generated by computer. Only those coordinates that were contained within the home range boundaries of radio-collared deer were selected. If a deer is selecting one or more habitat types over other habitats within its' home range, then one would expect to find differences between "use" plots and "random" plots. To test for differences between "use" versus

"random" plots within each sub-region as well as between the sub-regions, a two-way ANOVA was performed on the variables measured in vegetation plots. Canopy cover was estimated using a spherical densiometer at plot center and at the end points of the four cardinal directions that marked the perimeter of the plot

(i.e., north, east, west and south) (O'Connor 1988) resulting in five readings per plot. The observer faced 15 towards plot center when looking at the densiometer to reduce the influence of vegetation present on the outside of the plot on the estimation of canopy cover. The average percent canopy cover for each plot was calculated.

Shrub cover was estimated by a line transect technique. The line transect method involved counting the number of centimeters either obscuring or obscured by shrubs in a 10-meter section of the transect tape with the plot center as the mid-point of this 10-m section. This number could then be converted directly to a percentage to estimate shrub cover within the vegetation plot. Thus, if 5 meters were obscured by shrubs, then the shrub cover was estimated to be 50% (5m/10m X 100%). Either the north/south direction of the transect tape (assigned a '1') or east/west direction (assigned a '2') was used to estimate shrub cover.

The direction by which shrub cover was estimated was chosen randomly (by using the random numbers generator on a scientific calculator) for each plot.

Height of the understory was estimated using a Robel pole (Robel et al. 1970). The wooden pole (1 m in height)

was erected at plot center and at the end points marking the perimeter of the plot at the four cardinal directions. The pole was marked at 1 decimeter intervals using masking tape and an indelible marking pen. The observer stood at a distance of 4 meters from the pole and estimated vegetation height to the nearest decimeter. 16

A foliage volume board 3 m in height, was used to estimate vertical stratification inside each plot (Noon

1981). The board was erected at the four cardinal end points of the vegetation plot (Noon 1981). The observer stood at plot center and recorded the total number of squares obscured by foliage for two vertical strata: low shrub cover (0-1 m) and high shrub cover (1-3 m). The mean

low shrub and mean high shrub cover were calculated and

converted to percentages for each plot (O'Connor 1988). In addition, shrub and tree species that were observed within the plots were noted so that relative frequencies for most species could be estimated. This

information was then used to compare the two sub-regions in

terms of species composition present within the vegetation plots.

Habitat Diversity

Precursory field observations led me to hypothesize

that the home ranges of deer in the Porcupine Butte sub- region were less diverse than those of deer in the Hambone Butte sub-region. The Shannon-Wiener index (H') (Zar 1984)

was used to calculate habitat diversity using the

proportions of habitat types within each deer's home range.

This index was calculated for all deer within each sub- region. The mean indices of diversity for the first home ranges of deer within each sub-region were calculated and compared using a one-tailed t-test. 17

To determine if there was a lower habitat diversity within home ranges of deer that moved to second summer home ranges than those deer that did not move, I compared the mean diversity indices of the two groups with a one-tailed t-test.

If the original home ranges of deer that moved to second summer home ranges were less diverse than those of deer that did not move, then perhaps the two home ranges pooled together would have habitat diversity indices similar to those for the home ranges of deer that occupied single home ranges. To test this hypothesis, I calculated the mean habitat diversity index for the combined home ranges of each deer that exhibited this behavior and compared it with the mean diversity index for deer that occupied single home ranges using a two-tailed t-test.

Edge Effects

The "edge effect", or that edge that exists between two different habitat types, is considered important to wildlife species (Leopold 1933, Dasmann 1964, Thomas et al.

1979). As with habitat diversity, I hypothesized that the number of edges within deer home ranges between the two sub- regions differed. To examine this hypothesis, an index of habitat edge or interspersion index (I) (modified from

Baxter and Wolfe 1972) was used to evaluate the differences in the number of habitat edges between Hambone Butte and

Porcupine Butte. Baxter and Wolfe's method involved 18 dividing the area of interest into square sample blocks and then drawing diagonal lines across the squares from each corner to the opposite corner. The number of habitat changes were counted along both lines. Thus, 'I' was expressed in terms of the number of habitat changes per unit area or as the total changes of habitat types for the area of interest. I modified this method by drawing a line

between the two furthest points delimiting the boundaries of the home range for each deer. Another line was drawn through its' center and perpendicular to the first line extending to the edges of the home range boundary. The

number of habitat type changes that occurred along each line

was tallied and divided by the distance in meters that the

lines represented to give an index of the number of habitat changes/meter. The mean edge index for all deer for each

study area, using only the first home ranges, was calculated

and compared using a one-tailed t-test in order to determine

if the number of habitat edges for Hambone Butte is greater than or equal to the number of habitat edges for Porcupine

Butte. Similarly, a one-tailed t-test was also conducted to

compare the difference in mean number of habitat edges

between the first and second home ranges of the 6 deer that

had two home ranges. This was done to see if the number of

habitat edges within the first home range is greater than or

equal to the number of habitat edges of their second home

range. 19 Elevation The highest elevation within each deer's home range was estimated from topographic maps. These elevations were then used to calculate the mean maximum elevation for each sub-region. One-tailed t-tests were used to test for differences in the mean elevations between 1) sub-regions and 2) between first and second home ranges of deer that moved.

The single deer from Hambone Butte that exhibited the dual home range behavior was excluded from all analyses involving the comparisons of deer that relocated to second summer home ranges because deer at the Hambone Butte sub- region and deer at Porcupine Butte were considered two different populations (J. Allen, pers. comm., Humboldt State Univ. Arcata, CA. 95521). RESULTS

No locations were obtained from two of the radio- collared deer from Hambone Butte due to transmitter failure or some other unknown cause. One deer from Porcupine Butte died on her winter range between 1987 and 1988. Three hundred sixty five radio locations were obtained for the remaining 8 deer at Hambone Butte and 314 radio locations were obtained for 9 deer at Porcupine Butte. For Hambone Butte, the number of radio locations per deer ranged from 43 to 51 (X=47, s.e.=1.16). At Porcupine Butte, the number of radio locations per deer ranged from 19 to 44 (X=35, s.e.=3.27). The average angular bias of the three bearings taken for each deer was ±2.01°. The mean bias was not significantly different from zero (t=0.840, p>0.50, to.05(2),6=2.056), and therefore was not included within the confidence limits. The resulting confidence limits were

±4.79°. The bearing error (pooled standard deviation) was

12.4°. Distances between estimated and true locations ranged from less than 10 m and up to 175 m from polygon center (occularly estimated) to the actual deer location with a mean error of 80.7 m (n=22, s.e.=11.6).

20 21

Home Range

Six out of 9 deer in Porcupine Butte demonstrated greater movement than the Hambone Butte deer and established

"dual" home ranges (Table 1). These 6 deer remained at their initial capture sites until mid-summer at which time they moved to their second home ranges. These movements would often be completed within a 48 hour period (Appendix

D). One deer from Hambone Butte occupied two home ranges.

Home range area estimates ranged from 64 ha to 371 ha in

Hambone Butte (n=9) and 25 ha to 343 ha in Porcupine Butte

(n=15) when considering dual home ranges separately (Table

2). Mean home range size between deer occupying the Hambone and Porcupine Butte areas were not significantly different

(t=1.5, p=0.15). Home ranges for deer exhibiting the dual home range behavior were combined into a single estimate of home range and a t-test conducted to compare the mean home range size estimates of the two sub-regions. Again, there was no statistically significant difference between mean home ranges (t=0.01, p=0.99). Home range sizes for deer for each sub-region are listed in Appendixes B and C.

Six of 9 deer at Porcupine Butte occupied two summer home ranges whereas only one of the 8 deer at Hambone Butte had a second summer home range. The contingency table analysis that tested whether the two sub-regions differed in this deer behavior showed a significant difference (X2=7.61, p<0.005, 1 d.f.). 22

Table 1. Number of deer on Hambone Butte and Porcupine Butte that occupied one vs. two home ranges, Siskiyou County, California, 1987-1988.

Number of Number of Deer With One Deer With Two Sub-region Home Range Home Ranges N

Hambone Butte 7 1 8 Porcupine Butte 3 6 9

Total 10 7 17

23 Table 2. Summer home range sizes (ha) of female mule deer within each sub-region in Siskiyou County, California. 1987-1988 (n= number of home ranges)

Hambone Butte Porcupine Butte (for 8 deer) (for 9 deer)

Range X s.e. (n) Range X s.e. (n)

64 to 371a 196.6 36.5 (8) 25 to 338 130.0a 36.1 (9)

64 to 371b 196.7 32.2 (9) 25 to 343 132.3b 27.1(15)

110 to 371c 221.2 31.7 (8) 46 to 498 220.5c 53.3(9)

aEstimates using the first home range areas only.

bEstimates treating each dual home range separately.

cAreas for each dual home range summed into one home range. 24 Habitat Analysis

Deer at Hambone Butte used mixed conifer relatively more frequently than other habitat types whereas deer at Porcupine Butte frequented tree plantations (15 years old) and shrub habitats more than other habitat types (Fig. 4). The fact that mixed conifer existed within the home ranges of deer at Porcupine Butte was primarily a function of the dual home range behavior exhibited by these deer: the deer moved to areas where mixed conifer was present and plantations were absent. The portion within the Porcupine

Butte sub-region, called Lava Crack Springs (Fig. 2), had mixed conifer habitat along with plantations, mixed trees/shrub and shrub habitat types. Two deer were radio- collared at this location. One of these deer did not have mixed conifer habitat present within her home range but did occupy a second summer home range that consisted primarily of mixed coniferous forest. The other deer at the Lava

Crack location used mixed conifer within her home range and did not move to a second summer home range. All deer did not use habitat types in proportion to the availability of the habitats within their home ranges at either sub-region (Table 3). Deer at Hambone Butte used mixed conifer significantly more than expected, while mixed trees/shrubs and shrub habitat types were used significantly less than expected in proportion to their occurrence within deer home ranges (Table 4). Deer at Porcupine Butte, used Fig. 4. Frequency of use for each habitat type used by deer at Hambone Butte and Porcupine Butte within the McCloud Study Area. 25 26

Table 3. Chi-square analysis of habitat selection using the combined home ranges of deer at Hambone and Porcupine Buttes, Siskiyou County, California. 1987-1988.

Hambone Butte Porcupine Butte

Habitat Types X2 D.F. X2 D.F.

Mixed Conifer 26.12* 1 9.16* 1

Mixed Trees/shrub 26.67* 1 3.13 1

Shrub 22.15* 1 17.06* 1

Plantation < 15 0 1 13.92* 1 Plantation > 15 0.31 1 38.40* 1

Clearcut 0.88 1 1.21 1

* Indicates statistical significance at α=0.05

Chi-square= 76.13 with 5 d.f. P< 0.001 (Hambone Butte) Chi-square= 82.88 with 5 d.f. P< 0.001 (Porcupine Butte) Table 4. Bonferroni confidence intervals for habitats selected by deer at Hambone Butte, Siskiyou County, California, 1987-1988.

Habitat Total Prop.of Number Expected Prop.of Conf.int. on Types Area home range of number observ. proportion of (ha) ) (pio observ. observ.b occurrencec

Mixed Conifer 564.6 0.319(s) 177 109 0.517 0.466

Trees/ pio Shrub 517.0 0.292(a) 60 100 0.175 0.121

Shrub 257.0 0.145(a) 26 50 0.076 0.03915 55.7 0.031(n) 13 11 0.038 0.011

Total 1769.8 342 342 a Symbols within parentheses indicate: a=avoidance; s=selection and n=neither. b Calculated by multiplying proportion pio X n; i.e.,0.319X342=109 c pi represents theoretical proportion of occurrence and is compared to corresponding pi to determine if hypothesis of proportional use is accepted or rejected, i.e., Pi=Pio 27

28 plantations 15 years old significantly more than expected and used mixed conifer, shrub, and plantations < 15 years old significantly less than expected (using combined home ranges) (Table 5). Within the first home ranges of deer that had two summer home ranges (exclusive of the Hambone Butte doe with dual home ranges), habitats were not used in proportion to their availability. However, expected observations were fewer than 5 for the mixed conifer and clearcut habitat types (Table 6). Bonferroni confidence intervals for first home ranges of deer that moved showed that plantations > 15 years old were selected while mixed trees/shrub and plantations < 15 years were avoided (Table 7). Plantations were absent within the second home ranges of these deer.

Also, the proportions of clearcut and mixed trees/shrub habitat types within the second home ranges were so low they yielded expected observations of fewer than 5 in the chi- square analysis. Therefore, only mixed conifer and shrub habitat types could be included in the analysis (Table 8).

Within the second home ranges of deer that moved, mixed conifer was selected and shrub habitats were avoided (Table 9)

Vegetation Analysis

The two-way ANOVA analysis used to test for vegetative differences within deer home ranges comparing the two sub-regions indicated significant differences with Table 5. Bonferroni confidence intervals for habitats selected within combined home ranges by deer at Porcupine Butte, Siskiyou County, California, 1987-1988.

Habitat Total Prop.of Number Expected Prop.of Conf.int. on Types Area home range of number observ. proportion of (ha) a (P) io observ. observ.b occurrence

Mixed Conifer 366.7 0.193(a) 39 58 0.130 0.079

Shrub 756.5 0.398(a) 82 119 0.273 0.205

Plant. <15 138.1 0.073(a) 10 22 0.037 0.00815 436.6 0.230(s) 143 69 0.473 0.397

Table 6. Chi-square analysis of habitat selection within the first home ranges of the Porcupine Butte deer that occupied two summer home ranges.

Habitat Types Observed Expected X2 D.F.

Mixed Conifer 0 2 36.1 1

Trees/Shrub 11 22 11.0* 1

Shrub 47 38 1.7 1

Plantation < 15 6 20 32.7* 1

Plantation > 15 79 62 3.7* 1

Clearcut 1 0.1 0.8 1

• *Indicates statistical significance at a=0.05

Chi-square=85.96 with 5 d.f. P<0.001 Table 7. Bonferroni confidence intervals for habitats selected within first home ranges by deer that moved from Porcupine Butte, Siskiyou County, California, 1987-1988.

Habitat Total Prop.of Number Expected Prop.of Conf.int. on Types Area home range of number observ. proportion of (ha) (Pio)a observ. observ.b occurrence

Mixed Conifer 8.5 0.011(n) 0 2 0 0 Trees/ Shrub 115.0 0.151(a) 11 22 0.076 0.018

Plant. >15 328.8 0.430(s) 79 62 0.549 0.439

Clearcut 0.1 0.0001(n) 1 <1 0.007 -0.011

Table 8. Chi-square analysis of habitat selection within the second home ranges of the Porcupine Butte deer that occupied two summer home ranges.1

Habitat Types Observed Expected D.F.

Mixed Conifer 30 17 9.94* 1

Shrub 6 19 8.89* 1

1 Other habitats either had expected observations less than 5 or were non-existent within the second home range boundaries.

* Indicates statistical significance at a=0.05 X2=18.83 with 1 d.f. P<0.001 Table 9. Bonferroni confidence intervals for habitats selected within second home ranges by deer at Porcupine Butte, Siskiyou County, California.

Habitat Total Prop.of Number Expected Prop.of Conf.Int. on Types Area homea io p range of number of observ. proportion of (ha) observ. observ.b occurrence c

Mixed Conifer 333.6 0.427(s) 30 21 0.612 0.42815 0 0 0 0 0 0

Clearcut 11.3 0.014(n) 3 1 0.061 -0.029

Total 780.5 49 49 a Symbols with parentheses indicate: a=avoidance; s=selection and n=neither. b Calculated by multiplying proportion pio X n; i.e.,0.427X49=21 c pi, represents theoretical proportion of occurrence and is compared to corresponding piopi =ptoio determine if hypothesis of proportional use is accepted or rejected, i.e., 1 33 34 respect to 4 of the 5 vegetation variables measured (Table

10). Only the percent high cover was not significant between the two sub-regions (P=0.103). In addition, results for the 5 variables sampled in vegetation plots, showed no significant differences between use plots and random plots within deer home ranges at either Hambone Butte or Porcupine

Butte (Table 10). Thus, while the vegetative characteristics within deer home ranges for the two sub- regions were quite distinct from each other, there appeared to be no selection by deer for any of the variables that were measured within either sub-region (Table 11).

Habitat Diversity

Hambone Butte was significantly more diverse in terms of proportions of different habitat types within deer home ranges than were available within home ranges of deer at Porcupine Butte (t=1.95, P=0.035, n1=8, n2=9) (Table 12).

Also, home ranges for deer at Hambone Butte had significantly greater relative diversity values (J') than did the home ranges of deer at Porcupine Butte. That is, given the maximum number of habitat categories possible

(e.g, 6 habitat types), home ranges of deer at Hambone Butte were more diverse than those at the Porcupine Butte sub- region (t=1.957, 0.025

SUB-REGION PLOT TYPE Hambone vs Porcupine Use vs Random

Variable F-ratio df P F -ratio df P

Shrub Cover(%) 8.664 1,151 0.004 0.117 1,151 0.737 Canopy Cover (%) 4.885 1,151 0.029 1.482 1,151 0.225 Vegetation Height 10.873 1,151 0.001 0.060 1,151 0.809 Percent Low Cover 15.541 1,151 <0.001 0.869 1,151 0.363 Percent High Cover 2.693 1,151 0.103 2.204 1,151 0.140 36 Table 11. Means and standard errors of vegetation characteristics sampled within home ranges of deer at Hambone and Porcupine Buttes, Siskiyou County, California, 1987-1988.

Hambone Butte Porcupine Butte PLOT TYPE PLOT TYPE Use Random Use Random (n=46) (n=31) (n=44) (n=34)

Variable X s.e. X s.e. X s.e. X s.e.

Shrub Cover(%) 20.68 3.33 29.67 4.85 39.41 4.31 33.62 4.81 Canopy Cover(%) 49.23 4.42 44.30 5.91 39.35 4.19 32.32 5.37 Understory Vegetation Height(m) 0.24 0.04 0.31 0.05 0.45 0.04 0.36 0.05 Low Cover 36.6 4.65 39.52 6.00 63.68 5.05 51.15 5.45 (%) High Cover(%) 21.6 3.26 22.57 4.78 33.53 4.06 21.21 3.38 37 Table 12. Habitat diversity (H'), relative diversity (J') and heterogeneity (1-J') indices within the home ranges for deer on Hambone and Porcupine Buttes, Siskiyou County, California, 1987-1988.

Hambone Butte Porcupine Butte X n s.e. X n s.e.

H''a 0.514 8 0.0497 0.335 9 0.0744

b H 0.561 7 0.0206 0.359 6 0.0877 H'c 0.514 8 0.0497 0.496 6 0.0505 J'd 0.661 8 0.0642 0.430 9 0.0955

1-J'= 0.339 (Hambone Butte) 1-J'= 0.570 (Porcupine Butte)

a Habitat diversity indices for the first home ranges of deer from each sub-region. b Habitat diversity indices for deer from Hambone Butte that occupied single home ranges and the first home ranges of deer that moved from Porcupine Butte. c Habitat diversity indices for the Hambone Butte deer with single home ranges and for the combined home ranges of deer that moved from Porcupine Butte. d The relative diversity indices for first home ranges of deer from each sub-region. 38 habitat type than was the case for the home ranges of deer at Porcupine Butte (Table 12). Consequently, the heterogeneity value (1-J') was less for deer at Hambone

Butte than for deer at Porcupine Butte. The mean habitat diversity index for deer at Hambone Butte was significantly greater than the mean habitat diversity index for the first home ranges of the deer that moved from Porcupine Butte (t=2.41, 0.010.50; n,=6, n2=8) (Table 12).

Edge Effects

Deer at Hambone Butte had significantly more changes between habitat types (edges) existing within their home ranges than occurred within the first home ranges of deer at Porcupine Butte (t=2.09, P=0.027; n1=8, n2=9). The number of edges within the first home range of the doe that moved from

Hambone Butte was four with an interspersion index (I) of

3.41 habitat changes/meter (c/m). The number of edges within her second home range was nine with an interspersion index of 3.64 c/m. The mean interspersion index for all deer at Hambone Butte was 6.221 c/m (n=8, s.e.=0.526). The 39 doe from Hambone Butte that moved to a second summer home range had the lowest interspersion index of the Hambone Butte deer (Range: 3.41 to 8.30 c/m) (Appendix E).

The doe that moved from Lava Crack to a second summer home range, had a lower interspersion index (1=2.55 c/m) than the other Lava Crack doe which did not move (1=7.60 c/m). The mean interspersion index for the first home ranges of all the deer from Porcupine Butte was 3.43 c/m (n=9, s.e.=3.48). While the mean interspersion index for the second home ranges for deer that moved was greater (X=5.624 c/m; n=6, s.e.=1.54) than for the first home ranges (X=3.324 c/m; n=6, s.e.=1.50), the difference was not significant (t=1.07, P=0.15).

Elevation The mean elevations within the home ranges of deer at Hambone Butte and Porcupine Butte were approximately equal (t=1.603, P=0.06). However, the second home ranges of the Porcupine Butte deer were significantly higher in elevation than their first summer home ranges (t=-2.13, P=0.03).

Deer Migration

The earliest departure of a radio-collared deer from the study area was a doe (frequency #415) from Porcupine Butte which left the area on 27 August 1988 (Appendix D).

She was found dead on 19 September near Garner Mountain 40 presumably enroute to her winter range. The cause of death was unknown. The earliest migratory movement by a doe at

Hambone Butte, occurred 21 September 1988 (that is, she could not be located on the study area following her last location on 20 September 1988). All radio-collared deer departed from the study area by 20 October 1988 (Appendix D). Winter locations of these deer were obtained by Fish and Game personnel. DISCUSSION

Home Range

Home range sizes calculated for these deer were

probably underestimates of true home range size (Bekoff and

Mech 1984). Sample sizes for location data were small,

particularly for deer at Porcupine Butte. This is due to the

fact that when these deer relocated to their second home

range, the number of locations for each home range was

reduced (Appendix C). Since the number of radio locations

for both sub-regions were equally low, comparisons between

the mean home range sizes could be made.

The mean summer home range size for female mule and

black-tailed deer near McCloud was similar to that of deer

found elsewhere. Loft et al. (1984) found black-tailed deer

in Trinity County, California to have mean home range sizes

of 138 ha at elevations 910-1520 m and 185 ha at elevations

> 2070 m. Tierson et al. (1985) studied white-tailed deer in

New York that had a mean home range size of 221 ha. Non-

migratory mule deer studied in Montana had mean home ranges

of 121.5 ha (Mackie 1970). In contrast, migratory deer

studied east of Fresno, California, had relatively high mean

home range sizes of 300 to 400 ha at lower elevations and up

to 1000 ha in higher elevations (Bertam and Rempel 1977).

Migratory mule deer in Arizona had summer home ranges of up

41 42 to 7850 ha primarily due to the dry summer conditions

(Rautenstrauch and Krausman 1989).

Home range composition is influenced by inter- and intraspecific competition, predation and habitat quality.

These factors in turn influence the use of habitats within the home range (Loft et al. 1991). Large home range sizes may be indicative of low habitat diversity (Loft et al.

1984). Deer at McCloud Flats have comparable home range sizes to deer from other areas, however, their unique movements may be the result of missing habitat elements, drought, disturbance, or some other unknown factors.

Habitat Use

Five of 9 deer from Porcupine Butte moved to areas of mixed conifer. The sixth deer that moved to a second home range, went to an area that consisted of mixed trees/shrub, and shrub habitat types. There was only 8.5 ha of mixed conifer available within the first home ranges of the 6 Porcupine Butte deer that had dual home ranges. On the basis of habitat selection within the pooled home ranges, these deer appear to avoid the mixed conifer habitat type but this probably was misleading. The relocation to areas of mixed conifer found within the second home ranges could be viewed as habitat selection for this habitat type.

It is also possible that the unknown elements the deer were seeking in the mixed conifer habitats were not available to deer in the early summer due to seasonal influences. 43

Vegetation Analysis

Variables measured in vegetation plots appeared not to differ between use and random plots. Thus, there appeared to be no discernible vegetative features that were a typical characteristic of deer habitat. However, the analysis indicated a difference in species composition and structure observed between the two sub-regions. Percent shrub cover was greater at Porcupine Butte than at Hambone

Butte, which frequently resulted in impenetrable conditions at Porcupine Butte. Hambone Butte had a greater percent canopy cover than was present at Porcupine Butte with understory conditions that were more open and easily traversed.

Habitat Diversity

It has long been recognized that the quantity of habitat components is seldom the important factor in limiting wildlife populations, but rather the spatial relationships between those components (Dasmann 1964).

Homogeneous habitats do not provide the variety of plant communities that deer require. One reason that deer from

Hambone Butte did not move to other areas may be that habitat diversity within home ranges of deer at Hambone

Butte was significantly greater than for deer from Porcupine

Butte. 44 Edge Effects Edges between different habitat types are considered to be reflections of total diversity (Thomas et al. 1979). Deer have been shown to use edges disproportionately more than they will use other habitats (Dasmann 1964, Thomas et al. 1979). In 1933, Aldo Leopold declared that "game is a phenomenon of edges" and went on to name this phenomenon the

"law of interspersion". This concept implies that the density of game is proportional to the amount of edge for game species of low mobility that require more than one habitat type (Leopold 1933). The number of edges within deer home ranges in Hambone Butte was greater than the number of edges within the first home ranges of the

Porcupine Butte deer. Therefore, Hambone Butte may provide the desirable interspersion of plant communities for deer and that may explain why all but one of them occupied single home ranges.

Most of the changes between habitats within the first home ranges of the Porcupine Butte deer occurred between plantations and shrub habitat types. In contrast, edges within the home ranges of deer near Hambone Butte encompassed several different habitat types. Therefore, there were more edges between different combinations of habitats at Hambone Butte than at Porcupine Butte. Consequently, Hambone Butte probably was more desirable in terms of edge effects than Porcupine Butte. The exception 45 to this was the individual in Hambone Butte that relocated to a second home range which had the fewest number of edges of all the Hambone Butte deer.

In essence, there were 15 possible combinations of edges between habitat types available to deer at Hambone

Butte. At Porcupine Butte, only 5 edge combinations existed within the first home ranges.

Deer Movements

Bertram and Rempel (1977) discovered deer on spring or fall holding areas often made "test trips" to their summer or winter ranges and would subsequently return to the holding areas until final movement to their summer or winter range. Seven deer from Porcupine Butte occupied plantations

(>15 years old) and shrub habitats. Two of these deer remained at Porcupine Butte for the duration of the summer while 5 deer shifted to different habitats.

Deer 315 traveled the greatest distance leaving her first home range at Porcupine Butte to go to her second home range which was located 19 km north of Porcupine Butte near Medicine Lake (Fig.2). This area was dominated by lodgepole pine and red fir forest. This deer made two separate trips away from Porcupine Butte prior to her final departure to her second home range. She was absent from

Porcupine Butte on 25 June 1988 but returned by the 28 of

June. I did not determine where she was during that absence. Then on 4 July, she was located near Six Shooter 46

Butte. She returned once again to Porcupine Butte by 6 July and remained until her final departure to Medicine Lake where she was located on 15 July. Deer 436 moved from Porcupine Butte, north 11.2 km, to Six Shooter Butte. This area was primarily white fir forest and clearcuts. Both the Medicine Lake and Six Shooter Butte forests had little shrub understory (<3%).

Deer 395 spent most of the summer at Porcupine Butte. Her second home range was located approximately 8.1 km south of Porcupine Butte and was approximately 244 m lower in elevation. This area, called Julia Glover Flat, primarily consisted of shrub and mixed trees/shrub habitat types. She traveled to Julia Glover Flat shortly after 9 June 1988 and subsequently returned to Porcupine Butte by 12

June 1988. The most feasible explanation for this movement was the occurrence of a snowstorm on 9 June 1988. She remained at Porcupine Butte until 10 September 1988 when she returned to Julia Glover Flat. She remained at Julia Glover

Flat until she migrated to her winter range in Mid-October. Deer 335 and 456 moved 4.7 and 3.7 km respectively northwest of Porcupine Butte to Stud Hill. Stud Hill was adjacent to a lava flow and was rocky with steep terrain.

The vegetated areas were mixed conifer and patchy shrub habitat types. These deer made occasional trips between their dual home ranges. Deer 335 made one trip from Stud

Hill back to her first home range (31 July 1988) where she 47 remained until she returned to Stud Hill by 18 August 1988.

Similarly, deer 456 left Porcupine Butte for Stud Hill by 18

August but returned to Porcupine Butte on 13 September until a final move to Stud Hill by 3 October. She was not located on the study area again following that date. Deer 355 migrated to her winter range after 29 August.

The two deer within the Lava Crack Springs area (295 and 415) occupied mostly plantations (< 15 years old), mixed trees/shrub and mixed conifer habitat types. Deer 295 moved

12.7 km north after 22 July to Red Hill (northwest of Six

Shooter Butte). This area was mostly fir forests with low shrub understory. Deer 415 remained at Lava Crack throughout the summer. She was found dead on 19 September

1988 near Garner Mountain, presumably en route to her winter range.

In investigating habitat shifts by mule deer in relation to cattle grazing, Loft et al. (1991) found that deer did not travel to other areas of preferred habitats

probably because those other areas were already occupied by other deer or cattle. They believed reproductive fitness of

maternal does would be sacrificed by expanding home ranges

beyond their traditional home ranges to include a greater

proportion of preferred habitats. They concluded that the

energetic costs of enlarging home ranges or locating new

home ranges and of establishing new inter- and intraspecific

relationships would be detrimental to developing fawns. 48 However, optimal use of forage cannot be attained if animals remain sedentary, because the quality and quantity of forage is dynamic (Garrott et al. 1987). As availability of nutritious forage declines, greater movement is necessary for mule deer to meet their needs (Mackie 1970, Cornett et al. 1983 and Garrott et al. 1987). Browse species, such as bittercherry (Prunus emarginata), lose palatability in late summer and deer will move to other areas to find suitable habitat (Cornett et al. 1983, Riley and Dood 1984). Garrott et al. (1987) observed deer movements of distances between

6-14 km within their summer range, but these deer returned to their original territories.

Woodland caribou moved from open stands occupied in early summer to closed canopy forests late in the season where forage was still succulent and thermal cover was available (Servheen and Lyon 1989). They suggested that caribou moved into dense forests where deciduous forage was less susceptible to an early frost. Black-tailed deer near

Mineral King, California increased their use of red fir forests and meadows in late summer shifting from mixed brush habitat (Cornett et al. 1983). Although little understory was apparent in the fir forests occupied by some of the

Porcupine Butte deer, Cornett et al. (1983) found white fir to be one of the most important late-summer browse species of the Mineral King deer. Thus, there may have been sufficient white fir available to the Porcupine Butte deer 49 to account for their seasonal shifts to these forests.

Seasonal shifts in habitat types were also observed by Mackie (1970) and Dusek (1975). These movements were attributed to desiccation of the forage species within the original ranges of the deer.

Coniferous forests also provide thermal cover for deer during the hot summer months (Leckenby et al. 1982).

High summer temperatures can reduce productivity and lower the reserves necessary for deer to survive the coming winter (Leckenby et al. 1982). A forest can reduce air temperatures in the summer by -12.2° C below temperatures usually found in open areas (Reifsnyder and Lull 1965). While shrub communities also provide thermal cover, they are not as effective at reducing heat stress as coniferous forests (Leckenby et al. 1982). Optimum microclimates for deer usually require canopy closures greater than 50 percent (Verme 1965). An adequate overstory not only protects deer against high summer temperatures, but also allows soils to retain soil moisture (Steigers 1981). This may result in forage that retains nutrient content and palatability longer than in arid environments.

Elevations were between 610 m to 914 m higher in the

Six Shooter Butte and Medicine Lake area, where deer had second home ranges, than at Porcupine Butte. Thus, temperatures were also cooler due to the elevational difference. In contrast, Julia Glover Flat was 50 approximately 305 m lower in elevation than Porcupine Butte.

Perhaps the nature of the mixed trees/shrub habitat type provided better thermal cover for the doe (#395) that moved from Porcupine Butte, than was provided by the plantations and manzanita habitat types. Mixed trees/shrub habitats are characteristically less dense in terms of shrub understory and foliage volume, but still provide adequate canopy cover, thus air circulation might provide some relief from heat exposure (Thomas et al., 1979).

Conifer forest also provides excellent security cover (Carson and Peek 1987). Fawns have been observed to seek refuge (for hiding cover) in Douglas fir and juniper forests as the summer progressed, especially when precipitation levels were lower than normal (Riley and Dood

1984). Therefore, another motive for the Porcupine Butte deer to travel to areas of more coniferous habitat might have been to seek refuge from the disturbance associated with hunting season (Cornett et al. 1983).

Archery deer season in 1988 began on 20 August and lasted 23 days and a 2-week rifle season began on 1 October.

Deer 436, 315 and 295 relocated to areas with road closures,

which prevented access to vehicles and thus decreased disturbance to deer in that area. However, deer 315 and 295 moved to their second home ranges before the start of the archery hunting season (Appendix D). Thus, it was unlikely that an increased presence of vehicles and humans caused the 51 initial movement of these deer to second summer home ranges.

Pregnant does initially establish small fawning territories in early summer but will expand home range size later in the season and move to higher elevation meadows and forests when fawns are more mobile (Miller 1970, Cornett et al. 1983). The Porcupine Butte area may provide adequate fawning habitat for deer. This area consists mostly of impenetrable shrub and plantation habitats which provide thick cover suitable for fawning areas. Optimal fawning habitat is characterized by dense, multilayered vegetation of tall shrubs, succulent forage, and nearby water (Loft et al. 1984, Loft et al. 1987, Fielder and McKay 1984, Leckenby et al. 1982, Carson and Peek 1987). Four man-made water sources at the Porcupine Butte area, probably provided the necessary water for lactating does.

In contrast to Porcupine Butte, only one deer from the Hambone Butte area traveled to a new location late in the summer. Deer 365 vacated a home range consisting of coniferous forest and shrub habitats and relocated 6 km away to an area primarily consisting of open shrub, clearcut and plantation (< 15 years) with small portions of mixed conifer and trees with shrub habitats. Habitats within the second home range of deer 365, were similar in species composition and characteristics to Hambone Butte (Appendix A). Thus, it appears that deer 365 was using habitat types for her second home range that were opposite to those used by deer from 52

Porcupine Butte. The Porcupine Butte area may provide suitable forage and fawning habitat for early summer habitation, but may fail to meet other needs for these deer as the summer progresses. Thus, it would be advantageous to gather more information on the seasonality of the nutritional value of the forage species in both sub-regions. It would also be beneficial to compare reproductive rates and/or abundances of deer between sub-regions to see if there is any relationship between habitat diversity and reproductive success. REFERENCES CITED

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Mech, L.D. 1983. Handbook on radiotracking. Univ. of Minn. Press. Minneapolis. 97pp. Miller, F.L. 1970. Distribution patterns of black-tailed deer (Odocoileus hemionus columbianus) in relation to environment. J.of Mammal. 51:248-260.

Mohr, C.O. 1947. Table of equivalent populations of North American small mammals. Am. Midl. Nat. 37:223-249.

Mooty, J.J., P.D.Karns and T.K. Fuller. 1987. Habitat use and seasonal range size of white-tailed deer in northcentral Minnesota. J. Wildl. Manage. 51:644-648.

Neu,C.W., C.R. Byers and J.M. Peek. 1974. A technique for analysis of utilization-availability data. J. Wildl. Manage. 38:541-545.

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Reifsnyder, W.E., and H.W. Lull. 1965. Radiant energy in relation to forests. USDA For. Serv. Tech. Bull. 1344, 111 p. Washington, D.0 Riley, S.J. and A.R. Dood. 1984. Summer movements, home range, habitat use and behavior of mule deer fawns. J. Wildl. Manage. 48:1302-1310. Robel, R.S., J.N. Briggs, A.D. Dayton and L.C. Hulbert. 1970. Relationships between visual obstruction measurements and weight of grassland vegetation. J. Range. Manage. 23:295-297. Servheen, G. and L.J. Lyon. 1989. Habitat use by woodland caribou in the Selkirk Mountains. J. Wildl. Manage. 53:230-237. Springer, J.T. 1979. Some sources of bias and sampling error in radio triangulation. J. Wildl. Manage. 43:926-935. Steigers, W.D. 1978. Mortality of mule deer fawns in south- central Washington M.S. thesis. Brigham Young Univ., Provo, Utah. Thomas, J.W., C. Maser and J.E. Rodiek. 1979. 4 edges. Pages 48-57 in, J.W. Thomas (ed.), Wildlife habitats in a managed forest, the Blue Mountains of Oregon and Washington. USDA For. Serv., Agricult. Handbook. 553 pp.

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APPENDIX A. Tree and Shrub Species Composition Within Each Habitat Type Used by Radio-Collared Female Mule Deer During the Summer Near Hambone and Porcupine Buttes, Siskiyou County, California, 1987-1988.

Hambone Butte Porcupine Butte (n=29) (n=13)

No. of plots species % of % of No. of % of % of Species occurred plots' occur. plots plots occur.

Mixed Conifer Habitats Tree Species

White fir 21 72 32 10 77 40 (Abies concolor)

Ponderosa pine 18 62 28 1 8 4 (Pinus ponderosa) Lodgepole pine 10 34 15 4 31 16 (Pinus contorts)

Incense cedar 7 24 11 0 0 0 (Calocedrus decurrens)

Grand fir 4 14 6 4 31 16 (Abies gararandis

Sugar pine 4 14 6 1 8 4 (Pinus lambertiana) Calif. red fir 0 0 0 5 38 20 (Abies magnifica)

Shrub Species

Bitterbrush 11 38 13 0 0 0 (Purshia tridentata)

Bittercherry 1 3 1 0 0 0 (Prunus emarainata)

Chinquapin 0 0 0 0 3 23 (Castanopsis chrysophylla)

57 58 APPENDIX A. Tree and Shrub Species Composition Within Each Habitat Type Used by Radio-Collared Female Mule Deer During the Summer Near Hambone and Porcupine Buttes, Siskiyou County, California, 1987-1988. (continued)

Hambone Butte Porcupine Butte (n=29) (n=13)

No. of plots species % of % of No. of % of % of Species occurred plots' occur.b plots plots occur.

Mixed Conifer Habitats

Chokecherry 1 3 1 0 0 0 (Prunus viainiana) Manzanita 6 21 7 0 0 0 (Arctostaphvlos patula)

Rabbitbrush 15 52 17 0 0 0 (Chrvsothamnus viscidiflorus)

Tobacco brush 12 41 14 1 8 33 (Ceanothus velutinus)

Squawcarpet 17 59 19 0 0 0 (C. prostratus)

Squaw current 8 27 9 2 15 33 (Ribes cereum)

Western 16 55 18 0 0 0 serviceberry (Amelanchier alnifolia)

'This represents the number of plots in which the species occurred divided by the number of plots taken for that habitat type for each sub- region.

bThis represents the number of plots in which the species occur red divided by the total species occurrence. For example, there was a total of 64 occurrences of tree species present within mixed conifer vegetation plots for Hambone Butte. White fir was present within 21 plots; therefore, 21 divided by 64 and multiplied by 100 equals 32 percent occurrence for white fir. 59 APPENDIX A. Tree and Shrub Species Composition Within Each Habitat Type Used by Radio-Collared Female Mule Deer During the Summer Near Hambone and Porcupine Buttes, Siskiyou County, California, 1987-1988. (continued)

Hambone Butte Porcupine Butte (n=10) (n=12)

No. of plots species % of % of No. of % of % of Species occurred plots' occur.b plots plots occur.

Mixed Trees/Shrub Habitat Type

Tree Species

White fir 3 30 21 1 8 6 (Abies concolor)

Ponderosa pine 9 90 64 9 75 53 (Pinus ponderosa)

Lodgepole pine 0 0 0 0 0 0 (Pinus contorta)

Incense cedar 2 20 14 4 33 23 (Calocedrus decurrens)

Grand fir 0 0 0 2 17 12 (Abies grandis)

Sugar pine 0 0 0 1 8 6 (Pinus lambertiana)

Calif. red fir 0 0 0 0 0 0 (Abies maqnifica)

Shrub Species

Bitterbrush 6 60 19 9 75 22 (Purshia tridentata)

Bittercherry 0 0 0 4 33 10 (Prunus emarainata)

Chinquapin 0 0 0 3 25 7 (Castanopsis chrysophylla) 60 APPENDIX A. Tree and Shrub Species Composition Within Each Habitat Type Used by Radio-Collared Female Mule Deer During the Summer Near Hambone and Porcupine Buttes, Siskiyou County, California, 1987-1988. (continued)

Hambone Butte Porcupine Butte (n=10) (n=12)

No. of plots species % of % of No. of % of % of Species occurred plots' occur." plots plots occur.

Mixed Trees/Shrub Habitat Type

Chokecherry 0 0 0 2 17 5 (Prunus viainiana)

Manzanita 3 30 10 7 58 17 (Arctostaphvlos patula)

Rabbitbrush 8 80 26 2 17 5 (Chrvsothamnus viscidiflorus)

Tobacco brush 3 30 10 4 33 9 (Ceanothus velutinus)

Squawcarpet 8 80 26 4 33 9 (C. prostratus)

Squaw current 2 20 6 2 17 5 (Ribes cereum)

Western 1 10 3 4 33 9 Serviceberry (Amelanchier alnifolia)

'This represents the number of plots in which the species occurred divided by the number of plots taken for that habitat type for each sub-region. bThis represents the number of plots in which the species occ urred divided by the total species occurrence. For example, there was a total of 14 occurrences of tree species present within trees with shrub vegetation plots for Hambone Butte. White fir was present within 3 plots; therefore, 3 divided by 14 and multiplied by 100 equals 21 percent occurrence for white fir. 61 APPENDIX A. Tree and Shrub Species Composition Within Each Habitat Type Used by Radio-Collared Female Mule Deer During the Summer Near Hambone and Porcupine Buttes, Siskiyou County, California, 1987-1988. (continued)

Hambone Butte Porcupine Butte (n=7) (n=16)

No. of plots. species % of % of No. of % of % of Species occurred plots' occur.b plots plots occur.

Shrub Habitat Type

Tree Species

White fir 1 14 11 2 13 11 (Abies concolor) Ponderosa pine 6 86 67 9 56 50 (Pinus ponderosa)

Incense cedar 2 28 22 4 25 22 (Calocedrus decurrens)

Lodgepole pine 0 0 0 0 0 0 (Pinus contorta) Sugar pine 0 0 0 1 6 5 (Pinus lambertiana) Grand fir 0 0 0 2 13 11 (Abies arandis)

Shrub Species

Bitterbrush 6 86 22 7 44 13 (Purshia tridentata)

Bittercherry 2 29 7 10 63 18 (Prunus emarainata)

Chinquapin 0 0 0 1 6 2 (Castanopsis chrysophylla) 62 APPENDIX A. Tree and Shrub Species Composition Within Each Habitat Type Used by Radio-Collared Female Mule Deer During the Summer Near Hambone and Porcupine Buttes, Siskiyou County, California, 1987-1988. (continued)

Hambone Butte Porcupine Butte (n=7) (n=16)

No. of plots species % of % of b No. of % of % of Species occurred plotsa occur. plots plots occur.

Shrub Habitat Type

Chokecherry 1 14 4 5 31 9 (Prunus viginiana)

Manzanita 2 29 7 15 94 27 (Arctostaphylos patula)

Rabbitbrush 5 71 19 2 13 4 (Chrvsothamnus viscidiflorus)

Tobacco brush 4 57 15 8 50 14 (Ceanothus velutinus) Squawcarpet 1 14 4 7 44 13 (C. prostratus)

Squaw current 1 14 4 0 0 0 (Ribes cereum)

Western 5 71 19 1 6 2 Serviceberry (Amelanchier alnifolia)

This represents the number of plots in which the species occurred divided by the number of plots taken for that habitat type for each sub-region.

bThis represents the number of plots in which the species occurred divided by the total species occurrence. For example, there was a total of 9 tree species present within shrub vegetation plots for Hambone Butte. White fir was present within 1 plot; therefore, 1 divided by 9 and multiplied by 100 equals 11 percent occurrence for white fir. 63

APPENDIX A. Tree and Shrub Species Composition Within Each Habitat Type Used by Radio-Collared Female Mule Deer During the Summer Near Hambone and Porcupine Buttes, Siskiyou County, California, 1987-1988. (continued)

Hambone Butte Porcupine Butte (n=7) (n= 8)

No. of plots species % of % of No. of % of % of Species occurred plots' occur.b plots plots occur.

Plantations < 15 Years Old Habitat Type Tree Species White fir 1 14 11 0 0 0 (Abies concolor) Ponderosa pine 7 100 78 8 100 100 (Pinus ponderosa) Lodgepole pine 0 0 0 0 0 0 (Pinus contorta) Incense cedar 1 14 11 0 0 0 (Calocedrus decurrens) Grand fir 0 0 0 0 0 0 (Abies grandis) Sugar pine 0 0 0 0 0 0 (Pinus lambertiana) Shrub Species Bitterbrush 0 0 0 2 25 8 (Purshia tridentata) Bittercherry 3 43 16 0 0 0 (Prunus emarqinata) Chinquapin 0 0 0 0 0 0 (Castanopsis chrysophylla) 64 APPENDIX A. Tree and Shrub Species Composition Within Each Habitat Type Used by Radio-Collared Female Mule Deer During the Summer Near Hambone and Porcupine Buttes, Siskiyou County, California, 1987-1988. (continued)

Hambone Butte Porcupine Butte (n=7) (n=8)

No. of plots species % of % of No. of % of % of Species occurred plots' occur.b plots plots occur.

Plantations < 15 Years Old Habitat Type

Chokecherry 1 14 5 2 25 8 (Prunus viginiana)

Manzanita 2 29 11 8 100 31 (Arctostaphvlos patula) Rabbitbrush 4 57 21 1 13 4 (Chrvsothamnus viscidiflorus)

Tobacco brush 5 71 26 5 63 19 (Ceanothus velutinus)

Squawcarpet 0 0 0 7 87 27 (C. prostratus) Squaw current 0 0 0 1 13 4 (Ribes cereum)

Western 4 57 21 0 0 0 Serviceberry (Amelanchier alnifolia)

aThis represents the number of plots in which the species occurred divided by the number of plots taken for that habitat type for each sub-region.

bThis represents the number of plots in which the species occu Arctostaphylosyatulaotal species occurrence. For example, there was a total of 9 tree species present within plantations less than 15 years old vegetation plots for Hambone Butte. White fir was present within 1 plot; therefore, 1 divided by 9 and multiplied by 100 equals 11 percent occurrence for white fir. 65

APPENDIX A. Tree and Shrub Species Composition Within Each Habitat Type Used by Radio-Collared Female Mule Deer During the Summer Near Hambone and Porcupine Buttes, Siskiyou County, California, 1987-1988. (continued)

Hambone Butte Porcupine Butte (n=4) (n=23)

No. of plots species % of % of No. of % of % of Species occurred plots' occur.b plots plots occur.

Plantations > 15 Years Old Habitat Type Tree Species White fir 0 0 0 0 0 0 (Abies concolor) Ponderosa pine 4 100 100 23 100 100 (Pinus ponderosa) Lodgepole pine 0 0 0 0 0 0 (Pinus contorta) Incense cedar 0 0 0 0 0 0 (Calocedrus decurrens) Grand fir 0 0 0 0 0 0 (Abies grandis) Sugar pine 0 0 0 0 0 0 (Pinus lambertiana) Shrub Species Bitterbrush 0 0 0 1 4 1 (Purshia tridentata) Bittercherry 0 0 0 17 74 20 (Prunus emarginata) Chinquapin 0 0 0 0 0 0 (Castanopsis chrysophylla) 66 APPENDIX A. Tree and Shrub Species Composition Within Each Habitat Type Used by Radio-Collared Female Mule Deer During the Summer Near Hambone and Porcupine Buttes, Siskiyou County, California, 1987-1988. (continued)

Hambone Butte Porcupine Butte (n=4) (n=23)

No. of plots species % of % of No. of % of % of Species occurred plotsa occur.b plots plots occur.

Plantations > 15 Years Old Habitat Type

Chokecherry 2 50 17 7 30 8 (Prunus viginiana)

Manzanita 4 100 33 20 87 23 (Arctostaphylos patula)

Rabbitbrush 1 25 8 10 43 12 (Chrysothamnus viscidiflorus)

Tobacco brush 4 100 33 17 74 20 (Ceanothus velutinus)

Squawcarpet 1 25 8 12 52 14 (C. prostratus) Squaw current 0 0 0 0 0 0 (Ribes cereum)

Western 0 0 0 1 4 1 Serviceberry (Amelanchier alnifolia)

'This represents the number of plots in which the species occurred divided boccur.bumber of plots taken for that habitat type for each sub-region. bThis represents the number of plots in which the species occurred divided by the total species occurrence. For example, there was a total of 4 tree species present within plantations greater than 15 years old vegetation plots for Hambone Butte. Ponderosa pine was present within all 4 plots; therefore, 4 divided by 4 and multiplied by 100 equals 100 percent occurrence for ponderosa pine. 67 APPENDIX A. Tree and Shrub Species Composition Within Each Habitat Type Used by Radio-Collared Female Mule Deer During the Summer Near Hambone and Porcupine Buttes, Siskiyou County, California, 1987-1988. (continued)

Hambone Butte Porcupine Butte (n=13) (n=2)

No. of plots species % of % of No. of % of % of Species occurred plots' occur.b plots plots occur.

Clearcut Habitat Type Tree Species White fir 1 8 17 0 0 0 (Abies concolor) Ponderosa pine 4 31 67 1 50 100 (Pinus ponderosa)

Lodgepole pine 0 0 0 0 0 0 (Pinus contorts)

Incense cedar 1 8 17 0 0 0 (Calocedrus decurrens)

Grand fir 0 0 0 0 0 0 (Abies grandis) Sugar pine 0 0 0 0 0 0 (Pinus lambertiana)

Shrub Species

Bitterbrush 3 23 18 0 0 0 (Purshia tridentata) Bittercherry 0 0 0 0 0 0 (Prunus emarginata)

Chinquapin 0 0 0 0 0 0 (Castanopsis chrysophylla) 68 APPENDIX A. Tree and Shrub Species Composition Within Each Habitat Type Used by Radio-Collared Female Mule Deer During the Summer Near Hambone and Porcupine Buttes, Siskiyou County, California, 1987-1988. (continued)

Hambone Butte Porcupine Butte (n=13) (n=2)

No. of plots species % of % of No. of % of % of Species occurred plots' occur.b plots plots occur.

Clearcut Habitat Type

Chokecherry 1 8 6 0 0 0 (Prunus viginiana)

Manzanita 2 15 12 0 0 0 (Arctostaphvlos patula) Rabbitbrush 8 61 47 0 0 0 (Chrysothamnus viscidiflorus)

Tobacco brush 1 8 6 0 0 0 (Ceanothus velutinus) Squawcarpet 1 8 6 0 0 0 (C. prostratus)

Squaw current 0 0 0 1 50 100 (Ribes cereum)

Western 1 8 6 0 0 0 Serviceberry (Amelanchier alnifolia)

'This represents the number of plots in which the species occurred divided by the number of plots taken for that habitat type for each sub-region.

bThis represents the number of plots in which the species occurred divided by the total species occurrence. For example, there was a total of 6 tree species present within clearcut habitat type vegetation plots for Hambone Butte. Ponderosa pine was present within 4 plots; therefore, 4 divided by 6 and multiplied by 100 equals 67 percent occurrence for ponderosa pine. 69

APPENDIX A. Tree and Shrub Species Composition Within Each Habitat Type Used by Radio-Collared Female Mule Deer During the Summer Near Hambone and Porcupine Buttes, Siskiyou County, California, 1987-1988. (continued)

Hambone Butte Porcupine Butte (n=7) (n=4)

No. of plots species % of % of b No. of % of % of Species occurred plotsa occur. plots plots occur.

Ecotone Habitat Type

Tree Species White fir 3 43 30 1 25 25 (Abies concolor) Ponderosa pine 6 86 60 3 75 75 (Pinus ponderosa)

Lodgepole pine 0 0 0 0 0 0 (Pinus contorta)

Incense cedar 1 14 10 0 0 0 (Calocedrus decurrens) Grand fir 0 0 0 0 0 0 (Abies grandis) Sugar pine 0 0 0 0 0 0 (Pinus lambertiana)

Shrub Species

Bitterbrush 2 29 13 0 0 0 (Purshia tridentata) Bittercherry 0 0 0 1 25 9 (Prunus emarainata)

Chinquapin 0 0 0 0 0 0 (Castanopsis chrysophylla) 70 APPENDIX A. Tree and Shrub Species Composition Within Each Habitat Type Used by Radio-Collared Female Mule Deer During the Summer Near Hambone and Porcupine Buttes,Siskiyou County, California, 1987-1988. (continued)

Hambone Butte Porcupine Butte (n=7) (n=4)

No. of plots species % of % of b No. of % of % of Species occurred plots' occur. plots plots occur.

Ecotone Habitat Type

Chokecherry 0 0 0 1 25 9 (Prunus viainiana)

Manzanita 1 14 6 3 75 27 (Arctostaphylos patula) Rabbitbrush 5 71 31 0 0 0 (Chrvsothamnus viscidiflorus)

Tobacco brush 3 43 19 3 75 27 (Ceanothus velutinus)

Squawcarpet 3 43 19 1 25 9 (C. prostrates)

Squaw current 3 43 19 1 25 9 (Ribes cereum)

Western 2 29 13 0 0 0 Serviceberry (Amelanchier alnifolia

aThis represents the number of plots in which the species occurred divided by the number of plots taken for that habitat type for each sub-region.

bThis represents the number of plots in which the species occurred divided by the total species occurrence. For example, there was a total of 10 tree species present within ecotone habitat type vegetation plots for Hambone Butte. Ponderosa pine was present within 6 plots; therefore, 6 divided by 10 and multiplied by 100 equals 60 percent occurrence for ponderosa pine. 71

APPENDIX B. Summer Home Range Sizes (ha) for Female Mule Deer Near Hambone Butte, California. 1987-1988.

Deer Number of Size of Home I.D. Locations Range (ha)

285 51 186.244 305 43 125.278

326 47 370.616 345 44 312.252 365' 43 64.417

365 b 4 197.036

425 50 220.932 445 43 110.447

466 44 182.676 aFirst summer home range. bSecond summer home range. 72 APPENDIX C. Summer home range size (ha) for female mule deer at Porcupine Butte, near McCloud, California, 1987- 1988.

Deer Number of Size of home I.D. Locations Range (ha)

275 44 46.052 295a 14 338.262 295 b 5 159.910 315a 11 220.656 315 12 109.167 335a 26 56.093 335 b 3 27.042 355 38 98.496 395a 37 68.882 395 b 11 342.691 415 31 237.165 436a 28 25.477 436 b 14 89.587 456a 34 78.654 456 b 6 86.452 aFirst summer home range. Second summer home range. 73

APPENDIX D. Duration of summer occurrence for radio- collared female mule deer at Hambone and Porcupine Buttes near McCloud, California, 1988.

Deer ID. Dates Location Fawn

HAMBONE BUTTE 285 25 May-13 Oct. Mud Well Yes 305 25 May-24 Sept. Sand Flat Unknown

326 25 May-19 Oct. White Deer Lake Yes

345 25 May-13 Oct. Mud Well Unknown 365 25 May-13 Sept. Hambone Well Unknown 14 Sept-20 Sept. So. of Porcupine Unknown Lake 425 25 May-19 Oct. Mud Well Yes

445 25 May-3 Oct. Sand Flat Unknown

466 25 May-8 Oct. Toad Lake No

PORCUPINE BUTTE

275 24 May-17 Sept. Porcupine Butte Yes

295 24 May-22 July Lava Crack Unknown 3 Aug.-10 Sept. Red Hill

315 24 May-6 July Porcupine Butte Yes 15 July-17 Sept. Medicine Lake

335 24 May-31 July Porcupine Butte Unknown 1 Aug-18 Aug. Stud Hill

355 24 May-29 Aug. Porcupine Butte Unknown

395 24 May-3 June Porcupine Butte Unknown 4 June-9 June Julia Glover Flat 12 June-10 Sept. Porcupine Butte 13 Sept.-13 Oct. Julia Glover Flat 74

Appendix D. Duration of summer occurrence for radio- collared female mule deer at Hambone and Porcupine Buttes near McCloud, California, 1988. (continued)

Deer ID. Dates Location Fawn

415 24 May-27 Aug. Lava Crack No 436 24 May-13 Aug. Porcupine Butte Yes 18 Aug.-19 Oct. Six Shooter Butte

456 24 May-13 Aug. Porcupine Butte Unknown 18 Aug.-10 Sept. Stud Hill 13 Sept.-10 Sept. Porcupine Butte 3 Oct. (migrated) Stud Hill 75

APPENDIX E. Number of edges and interspersion indices calculated for radio-collared deer (after Baxter and Wolfe 1972) at Hambone Butte, Siskiyou County, California. 1987- 1988.

Deer Number of Interspersion I.D. Edges Index (# edges/m)

285 13 6.256

305 9 4.902

326 21 6.621

345 18 6.776

365a 4 3.413

365b 9 3.645

425 14 6.222

445 14 8.296

466 19 7.281

First home range bSecond home range 76

APPENDIX E. Number of edges and interspersion indices calculated for radio-collared deer (after Baxter and Wolfe 1972) at Hambone Butte, Siskiyou County, California. 1987- 1988. (continued)

Deer Number of Interspersion I.D. Edges Index (# edges/m)

275 0 0 295a 7 2.546

295b 11 4.191 315a 1 0.435

315b 0 0

335a 6 4.923

335b 8 8.678

355 5 3.368

395a 0 0

395b 13 4.078

415 18 7.604

436a 9 9.931

436b 9 6.128

456a 3 2.110

456b 15 10.667

'First home range

bSecond home range