Temporal Patterns of Spring Space Use by Ruffed Grouse

Temporal Patterns of Spring Space Use by Ruffed Grouse Author(s): Herbert L. Archibald Reviewed work(s): Source: The Journal of Wildlife Management, Vol. 39, No. 3 (Jul., 1975), pp. 472-481 Published by: Allen Press Stable URL: http://www.jstor.org/stable/3800386 . Accessed: 06/02/2012 12:38 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected].

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TEMPORAL PATTERNS OF SPRINGSPACEUSE BY RUFFEDGROUSE' HERBERT L. ARCHIBALD,Department of Ecology and Behavioral Biology, University of Minnesota, St. Paul 551012

Abstract: During spring 1970, locations of 10 male and 3 female ruffed grouse (Bonasa umbellus) were determined with an automatic radio-trackingsystem on the Cedar Creek Natural History Area in east-central Minnesota. Mean weekly range size of drumming adult males was significantly greater than

that of drummingjuvenilemales. An inverserelationshipwas found betweenmean weekly range size

of males and their level of drumming activity. Mean range size of females increased in an approximately linear manner during the six weeks prior to incubation but became much smaller (ca. 2 ha) during incubation. Cumulative ranges of drummingmales were more fixed in space than, and approximately half as large as, those of females. Male ranges became more fixed in space after the onset of

the drummingseason. Cumulativerangesof drummingmales containedcentrallylocated core areas

(k = 2.26 ha) of intensive use and consistent occupancy over time. Habitat quality and neighboring males are probably the two major factors influencing spring space use of drummingmales. J. WILDL. MANAGE. 39(3):472-481

Temporal variations in an animal's pattern of space use result from changes in its physiological and ecological requirements and from responses to changes in features of its environment, including presence or absence of conspecifics. Much of the present knowledge of the movement and space use patterns of ruffed grouse has resulted from studies in Wisconsin (Hale and Dorney 1963), northern Minnesota (Gullion 1970, unpublished report), and Alberta (Rusch and Keith 1971). This paper reports on temporal changes in space use patterns of ruffed grouse during the breeding season of 1970. For advice and encouragement in all phases of the study, I thank the following University of Minnesota personnel: J. R. Tester, G. W. Gullion, W. H. Marshall, D. B. Siniff, and D. F. McKinney. The field assistance of R. A. Huempfner and others at the Cedar Creek Natural History Area is gratefully acknowledged. Technical assistance was provided by University of Min-

nesota Bioelectronics Laboratory (UMBL) personnel including V. B. Kuechle, R. A. Reichle, and R. J. Schuster; and by R. K. Maxwell. C. M. Kirkpatrick, G. L. Storm, and H. P. Weeks offered useful suggestions on earlier versions of the manuscript. Financial support was provided by NIH Training Grant No. 5 TO1 GM01779 and the U.S. Atomic Energy Commission (COO1332-106).

STUDYAREA

This investigation was conducted at the Cedar Creek Natural History Area, about 48 km north of Minneapolis, Minnesota. Pierce (1954) and Bray et al. (1959) described the soils, topography, and vegetation of the area. Generally, the topography is flat to gently rolling sandy uplands interspersed with lowland swamps, bogs, and marshes. Lowland vegetation in the area inhabited by study grouse consists primarily of alder (Alnus rugosa), white cedar (Thuja occidentalis), tamarack (Larix laricina), willow (Salix spp.), and red osier dogwood 'Journal Paper No. 5629, Purdue University Agricultural Experiment Station. (Cornus stolonifera). Other lowland vegePresent address: Department of Forestry and tation includes cattail (Typha latifolia) Natural Resources, Purdue University, West marshes and sedge (Carex spp.) meadows. Lafayette, Indiana 47907. 2

472

J. Wildl. Manage. 39 (3) :1975

SPACE

The upland-lowland edge is dominated by white birch (Betula papyrifera) and aspen (Populus tremuloides). Vegetation on uncleared uplands is primarily oak (Quercus spp.) forest with an understory of hazel ( Corylus spp.). During the study, temperatures ranged from -23 C on 27 March to 34 C on 29 June. The last substantial snowfall occurred on 20 March, with an accumulation of 8.9 cm. Most of the snow was gone in open areas by 8 April. All weather data were recorded at the U.S. Weather Bureau Station on the Cedar Creek Natural History Area.

METHODS Grouse were captured in lily-pad and mirror traps (Gullion 1965, 1966, unpublished report), weighed, and leg-banded. Central rectrix length was the primary criterion for determination of sex (Dorney and Holzer 1957, Gullion 1964, unpublished report, Minnesota P-R Quart. 24:26-137). Age was determined by comparison of the amount of sheathing at the base of the 8th and 9th primaries (Hale et al. 1954). Grouse were fitted with continuously broadcasting radio-transmitters (53 mHz) built by UMBL. Polyvinyl chloride tubing was used to attach the transmitter package (21-25 g) to the bird (Brander 1968). All grouse were released at the capture site immediately after being equipped with transmitters. The general behavior of the study birds was not noticeably affected by the transmitter package during the study. Males appeared to drum normally in terms of both frequency of drums and sound production, and females nested and began incubation normally. One nest was destroyed by a predator, and another was deserted after accidental capture of the hen in a mammal live-trap. Location data were not analyzed after abnormal termination of incubation. Two males were killed by avian J. Wildl. Manage. 39(3):1975

USE BY RUFFED

GROUSE *

Archibald

473

predators, and another died accidently during the study. An automatic radio-tracking system (Cochran et al. 1965) was used to record location azimuths which were read to the nearest degree at a five-minute sampling interval and then converted to x-y coordinates (Siniff and Tester 1965). Seven-day data periods were chosen to provide averaging of day-to-day variability in movements and a sufficient number of periods for studying temporal changes in space use patterns. Ranges were calculated with a modification of the "computer-fill"method described by Phillips et al. (1973) and Rongstad and Tester (1969). An 0.026-ha (0.016 km on a side) gridsquare size was selected so that use of an entire gridsquare could be assumed, given a fix (radio-tracking location) recorded at any point within, and all gridsquares contained at least one possible fix (Siniff 1966, unpublished report, Minnesota Mus. Nat. Hist. Tech. Rep. 12). Range size was estimated by summing gridsquares containing one or more fixes plus certain empty gridsquares assumed to have been occupied according to two criteria. First, if two fixes were located less than 0.097 km apart in space and 5.1 minutes apart in time, a linear travel route was assumed, and all empty squares crossed by the travel route were assumed to have been occupied. Second, if two squares, each containing one or more fixes or assumed (travel route) locations, were separated in a vertical or horizontal direction within the grid by two or fewer empty squares, those empty squares were assumed to have been occupied. The maximum possible number of fixes for a grouse in a 7-day period was 2,016. Weekly sample sizes obtained ranged from 404 to 1,520, with a mean of 931. Sample sizes were sufficiently large so that

474

SPACE

USE BY RUFFED

GROUSE

?

Archibald

no correlationwas found between weekly range size and numberof fixes. All ranges analyzed fell within the 61.0-m (200-ft) accuracy zone (Heezen and Tester 1967); thus a "worst-possiblecase" azimuth error of 1 degree would result in a distance error not exceeding 61.0 m and averaging considerably less. Range shape (Mohr and Stumpf 1966) was describedby the ratioof greatestlength (i.e., major axis between the two most distant fixes) to greatest width (i.e., sum of perpendiculardistances from the most distant fix on each side of the majoraxis). To describehow much a grouse changed the location of its range over time, two measures of variation of range location in space were developed: (1) mean activity center deviation, and (2) ratio of mean to maximumpossible gridsquareuse. For the first measure,the center of activity (Hayne 1949) was determinedby computingmeans of the x- and y-coordinatesof all fixes for a grouse in a seven-day period. The distances between weekly activity tenters and the mean of all weekly activity centersprovided activitycenter deviations,which were then averaged. For the second measure,a tally was made of the numberof seven-day periods in which each gridsquare within the cumulativerange (i.e., that for all weeks a grouse was radio-tracked)was occupied one or more times. The ratio of the mean numberof weeks in which gridsquareswere occupied to the maximum possible (i.e., the total number of weeks) provided the second measure. Its value varied from 1/n (where n is the number of weeks) in the case of use of completely discrete areas to 1.0 if exactly the same space was used in each week. Core areas (Gullion 1953,Weeden 1965) within the cumulativeranges of males were

fined as all gridsquares occupied one or more times in 50 percent or more of the total weeks. For an individualradio-tracked an odd number of weeks, half of the gridsquares occupied for the modal number of weeks were included in the core area. Selection of such gridsquareswas made on the basis of adjacencyto squares with the highest numberof weeks of occupation.Intensity of use (Siniff 1966, unpublished report, Minnesota Mus. Nat. Hist. Tech. Rep. 12, Adams and Davis 1967) of the core area was computed as the percentage of the total numberof fixes obtainedwhich fell in the core area.

delineated and examined for possible territorial significance. The core area was de-

and females were different, likely reflecting behavioral differences between sexes.

RESULTS Locationsof 10 males and 3 females were analyzed for a total of 83,926 fixes and 629 bird-days. Radio-markedmales began drummingon 9 April 1970 and had greatly reduced drumming activity by 10 June. Thirteen7-day data periods,from 12 March through 10 June, arbitrarilywere defined, with 9 April as the first day of week 5. With the exception of one adult killed by an avian predatorabout three weeks before drumming began, all radio-markedmales were "active, site-associated" drummers (Gullion 1966:718). Variationsin Weekly RangeSize and Shape Several trends were evident in mean weekly range size over the study period (Fig. 1). For both sexes, the variation in range size among weeks 1 through 3 may be partially attributable to differences in air temperature.The 7-day averagesof the daily mean temperaturesfor these periods were -6.4, -1.4, and -7.9 C, respectively, suggesting that colder weather may have resulted in reduced movement. Beginning with week 4, mean range sizes for males

J. Wildl. Manage. 39(3):1975

SPACE

DRUMMING

Week

1

c/)

2 3

W

4 3 8

3

475

Mean weekly range size (ha) of adult and juvenile drumming male ruffed grouse (sample size in parentheses).

4

z

Archibald

Table 1.

MALES

C.J

GROUSE *

USE BY RUFFED

5

5

5

4

5

5

6

6

5

5

FEMALES

w

N 6

(C

INCUBATION

z

4

4

5

6

Adult males

Juvenile males

Difference

4.30 (2) 3.64 (3)

3.88 (1) 3.07 (2)

0.42 0.57

4.47 (3)

3.48 (2)

2.48 (3) 4.00 (3) 3.66 (2)

7 8 9 10

2.78 2.90 2.94 3.16

11 12 13

2.24 (2) 2.33 (2) 4.34 (2)

(4) (4) (3) (3)

4.18 (2)

-0.18

3.37 (2)

0.29

2.44 2.10 2.35 1.74

(1) (1) (3) (3)

2.14 (3) 3.06 (3) 3.59 (2)

0.99 0.34 0.80 0.59 1.42

0.10 -0.73 0.75

selection, and increased nutritional requirements prior to and during laying. In con2 3 3 3 3 3 S I ! ! trast, ranges during weeks 10-12 were rel4 I 2 3 5 7 6 8 9 10 11I 12 13 atively small, indicating that females have WEEK a much more restricted movement pattern Fig. 1. Seasonal variatior in range size by sex. Week 1, during incubation. 12-18 March; week 13, 4-10 June. Bars depict maxima and Mean range size of adult (22 or more minima; sample size above x-axis. months old) males exceeded that for juveFollowing the onset of the spring drum- nile (10-12 months old) males in 10 of 12 ming period in week 5, mean range size of weeks (Table 1). The age difference in males decreased and did not again reach the range size of drumming males (ci = 0.45 ha) week 4-5 size until week 13 (Fig. 1). By was significant at the 0.025 level (Wilcoxon this time drumming activity had greatly signed ranks test, Sokal and Rohlf 1969:400). decreased. An inverse relationship is there- The propriety of using the test is, however, fore suggested between weekly mean range open to question because the means are size of males and their level of drumming based on small, unequal samples. Previous activity. The comparatively small mean findings that juvenile males in general are range sizes during weeks 6-12 were consis- more mobile than adult males (Hale and tent with Eng's (1959) finding that the Dorney 1963, Rusch and Keith 1971) reflect distance moved from the drumming log was the greater mobility of non-drumming, nonat a minimum in the spring (16 March- territorial males which are primarily juve31 May). niles (Gullion 1967, Rusch and Keith 1971). No pronounced sexual differences or seaWeekly mean range size for females increased in an approximately linear manner sonal trends in range shape were apparent from weeks 4 through 9 (Fig. 1). Although (Table 2). Large mean length-width ratios other factors may have been involved, the (1.71, 1.68) for two males radio-tracked relatively large mean range sizes in weeks before and during the drumming season 7-9 are probably a reflection of mate-seek- appeared to be a reflection of the linear ing movements (Brander 1967), nest-site configuration of the habitat in which they J. Wildl. Manage. 39(3):1975

- Archibald

476

SPACE USE

Table 2.

Shape and variation of location in space of ranges of ruffed grouse.

BY RUFFED

Total

Males Before and during drummingperiod During drumming period only Total Females

GROUSE

Ratio of greatest length to greatest width of weekly range

Activity center deviation (km)

Ratio of mean to maximum possible gridsquare use Entire cumulative range

Core area within cumulative rangec

No.

no.

birds

weeks

Meana

SE

Meanb

SE

5

36

1.60

0.103

0.045

0.016

0.423

0.055

0.785

0.052

4 9d

24 60

1.32 1.47

0.159 0.191

0.019 0.034

0.008 0.019

0.542 0.476

0.070 0.085

0.882 0.828

0.041 0.068

3

29

1.62

0.140

0.123

0.041

0.243

0.038

Mean

SE

Mean

SE

Average of the mean length-width ratio for each grouse in category. ba Average of the mean activity center deviation for each grouse in category. Not for females. dc Does computed not include an adult male radio-tracked only in week 1.

lived. In a given week, the shape of a grouse's range seemed to be determined largely by the extent to which it visited seldom-used, peripheral portions of its cumulative range. The among-week variability in the shape of the ranges of most birds (average of the standard deviation of ratios for each grouse was 0.414, n = 12) suggested that range shape was transitory. Differences in ratios among birds suggested the lack of a common explanation of range shape. Variations in the Location of Weekly Ranges in Space It is important to determine not only changes in the size of an individual's range over time but also temporal changes in the location of its range in space. Both measures of variation of range location in space (Table 2) indicated that the ranges of drumming males were considerably more fixed in space (stationary) than those of females, as might be expected. The low values for mean activity center deviation (Table 2) showed that the center of a male's range changed little over time. Ratios of mean to maximum possible gridsquare use showed that males used the area within

their cumulative ranges more consistently over time than females. Yet, the fact that ratios for males were considerably lower than the maximum of 1.0 (Table 2) indicated that males did not use parts, mainly the peripheral portions, of their cumulative ranges consistently (Fig. 2). Evidence of two kinds suggested that ranges of males were more stationary during the drumming period (weeks 5-13). First, smaller activity center deviations were found for males radio-tracked only in the drumming period (Table 2). Second, of the 60 weeks of range data on males, 5 activity center deviations exceeded 0.08 km. All of these larger deviations occurred in or prior to period 5, the first week of drumming. For example, male 1628's activity center deviation in week 3 was 0.12 km, but after the onset of drumming (period 5) its weekly centers of activity fell in a tight cluster around the location of its primary drumming log (Fig. 2). During active drumming or incubation, the geometric centers of activity of grouse generally fell remarkably close to their activity centers of biological significance-the drumming log and the nest. J. Wildl. Manage. 39 (3):1975

SPACE

USE

BY

RUFFED

GROUSE

- Archibald

477

NUMBER OF WEEKS OF OCCUPATION S= PRIMARY DRUMMING LOG

S=1-2

S

=3-5

6-8

= 9-10 X.

.;t';

......... 8? ? ? ?

...

.

?

'?'0

~? ?2? ?

. ;. . ??5????

X.. ?. . .

Fig 2.Cuulaiverage

f

jveilemal

-12(26Mach-

128,wees

Jue)

??

KM ..0.16

Fig. 2.

01

MILE

Cumulative range of iuvenile male 1628, weeks 3-12 (26 March-3 June).

Characteristics of Cumulative Ranges Sexual differences were found in both the pattern of development and the ultimate size of cumulative ranges. Rates of increase in cumulative range size of males tended to be large in the early weeks and again in the final week of the study (Fig. 3). In contrast, hectarage additions to cumulative ranges were relatively small during weeks 6 through 12, suggesting the areas occupied by males then were substantially the same each week. Curves for females (Fig. 3) showed a pattern of continual increase that did not reach a plateau until just prior to the start of incubation. The size of cumulative ranges of males was found to be highly correlated (r = -0.90, P < 0.001) with the week in which radio-tracking was initiated, reflecting relatively large weekly ranges J. Wildl. Manage. 39 (3):1975

early in the study. Thus, curves are not comparable unless they originated in the same week. No male was radio-tracked either for the entire study period or only the drumming portion of the study. A regression equation (Y -9.504 - 0.553 X1 - 0.006 X2; 0.01< P < 0.025) relating overall range size (Y) to initial week (XI) and number of weeks radio-tracked (X2) was used to estimate range size of males for these two periods. Resulting estimates of range size (SE = 1.18) were 8.9 ha for the entire period (12 March-10 June) and 6.7 ha for the "drumming season" (9 April-10 June). Cumulative range sizes for females (2 =16.5 ha) were roughly twice as large as the regression estimate for males during the entire study period. The large cumulative range sizes

SPACE USE BY RUFFEDGROUSE* Archibald

478

20 -

20-

u)

w IS -15 MALES

FEMALES

z N

r

10 -10-

w

-5

5-

4

I

_

_

DRUMMING 0

v -.-....

0 I 2 3 4 5 6 7 8 9 1011 12 13 WEEK Fig. 3.

......•.....?

?0

v

V

0 1 2 3 4 5 6 7 8 9 10 111213 WEEK v?1?

Development of the ranges of individual male and female ruffed grouse.

and activity center deviations (Table 2)

provide documentationof the tendency for greater female mobility reported by Hale and Dorney (1963) and others. The core area of a male grouse's range can be characterizedas an area of: (1)

greater consistency of use of the core area in males radio-tracked only during the

drummingperiod (Table 2). Much of the variation in use-intensity of the core area was due to a low value (40.6 percent) for an adult male which was radio-trackedin in 2.26 ? 0.54 (1 SD) ha size, comprising weeks 1-5 and then killed by an avian pred34.5 - 10.37 percent of the cumulative ator. Apparentlythis bird had not begun range, (2) consistentoccupancy over time, to use a central area consistentlyprior to made up of gridsquaresused in 82.8 ? 6.79 the onset of drumming. In all cases, the primary drumming log percentof the total weeks, and (3) intensive use, containing 79.6 + 16.17 percent of the (Gullion 1967) was contained within the total locations. Similar to the case with core area, and the core area was centrally cumulative range, there was a tendency for located within the cumulativerange. With J. Wildl. Manage.39(3):1975

SPACE

N

J

I-7 0 -0 LOWLAND

0

00W

UPLAND

USE-INTENSITYBY 1640 0

0

> 1%

USE-INTENSITYBY 1641

0/

144

[

BOUNDARYBETWEEN RANGES --= EXCLUSIVEUSE BY 1640 = EXCLUSIVEUSE BY 1641

U

= ZONE OF OVERLAP

KM 0.16 0.1 MILE

Fig. 4. May.

Ranges of juvenile males 1640 and 1641, 14-20

the exception of one non-connected square in the case of one male, all core areas were single regions in space. However, as might be expected, consistency and intensity of use of gridsquares were not uniform within core areas. For example, male 1628 used an area comprised of 32 gridsquares (0.83 ha) in at least 9 of the weeks in which it was radio-tracked (Fig. 2), whereas its core area (occupied in 6 or more weeks) was 1.61 ha in size. Factors Influencing Use of Space by Males The two major factors which influence the spring space use of drumming males probably are habitat quality and neighJ. Wildl. Manage. 39 (3):1975

USE

BY RUFFED

GROUSE *

Archibald

479

boring males (Aubin 1970). Although detailed consideration of these factors is beyond the scope of this paper, Fig. 4 provides graphic illustration of their influence. Upland-lowland classification of habitat provides insight into the question of what determines the range selected by a male grouse. Fig. 4,A shows that males 1640 and 1641 both had a pronounced affinity for the upland-lowland edge. The dominant plants within their ranges included speckled alder in the lowland, oaks and hazel in the upland, and trembling aspen along the edge. The northwesternmost extent of both birds' ranges was approximately aligned with a transition zone to a sedge-willow association. For most of its length, the boundary between the ranges of males 1640 and 1641 was found to be a region of overlap (0.47 ha) rather than an absolute line separating zones of exclusive use (Fig. 4,B). The boundary line was drawn on the basis of greater intensity of use within individual gridsquares. Each grouse was found to confine nearly all (93-96 percent) of its use of the overlap zone to "its" side of the boundary line. Although the behavior involved in the establishment and maintenance of the boundary region is not known, the apparent effect is restriction of the range of each male in the direction of the neighbor's range. DISCUSSION Knowledge of a species' pattern of space use is an essential part of our understanding of its ecology and management. Insight is needed on the factors which interact to determine the size and location of the animal's range as well as the distribution of useintensity within it. In this study, the temporal variations in range characteristics which have been documented can at best be explained only partially.

480

SPACE USE BY RUFFED GROUSE - Archibald

The larger cumulative range sizes found for females (Fig. 3) may be partially attributable to a change in their pattern of habitat use. Early in the study, both sexes were noted to use lowland and upland-lowland edge habitat almost exclusively. For males, this trend continued throughout the study period; however, although females occasionally nest in lowlands on the study area (S. Maxson, personal communication), all three of the radio-marked females in the present study nested in oak-upland situations. Consequently, females showed an increasing tendency to use upland habitat beginning with the onset of nesting activities. On the same study area, Meslow (1966) found a tendency for drumming logs to be located in lowland habitat (84 percent) and near the upland-lowland edge (mode = 18.3 m). The preference for lowland drumming sites at Cedar Creek may reflect lack of suitable upland habitat (Gullion, personal communication) due to former agricultural use. The approximately inverse relationship between mean range size of males and females during weeks 5-9 (9 April-13 May) is consistent with the view that the more mobile female, attracted by drumming, seeks out the territorial male at its drumming log (Brander 1967). Courtship and copulation take place during a brief period in the vicinity of the drumming log (Gladfelter and McBurney 1971), and no further intentional contact occurs between the sexes (Gullion 1970, unpublished report). One of the essential characteristics of a territory is that it is a "fixed area, which may change slightly over a period of time" (Brown and Orians 1970:242). Due to inconsistency in the use of the peripheral portions over time, the cumulative ranges of males did not seem sufficiently fixed to satisfy this criterion; however, the core areas

within males' ranges did appear to meet the "fixed area" criterion due to their high intensity and consistency of use. Since it contained the primary drumming log in all cases, the core area seems synonymous with Gullion's (1967) "activity center" and Fowle's (1953) "primary area." Similar to the results presented here, Weeden (1965) found that total activity spaces (i.e., ranges) of tree sparrows (Spizella arborea) generally had a central core of more concentrated use which was reaffirmed by advertisement and an outer cortex where visits were less frequent. Another characteristic of a territory is that it is a "defended area" (Noble 1939). Pitelka (1959:253) emphasized that the fundamental importance of territory lies "in the degree to which it is in fact used exclusively by its occupant." The example of the ranges of males 1640 and 1641 (Fig. 4) suggested a high degree of exclusiveness; however, there was no direct evidence of defense of the boundary demonstrated in this example. The age difference found in range size of drumming males (Table 1) suggested that adults generally are able to defend larger territories than juveniles. Drumming is thought to be a defensive act (Gullion 1967), and one of its functions commonly is regarded as facilitation of spacing among males (Fowle 1953). The relatively large size and high within-week variability in size of male ranges during weeks 4-6 may reflect a period of unstable spatial organization within the male population (Fig. 1). Weeks 7 through 12 were characterized by consistently low means and within-week variability, suggesting a period of relatively stable spatial organization. Since drumming began in week 5 and tapered off markedly during week 13, these data support the hypothesis that drumming is involved in the maintenance of stable spatial relationships among males. J. Wildl. Manage. 39(3):1975

SPACE USE BY RUFFED GROUSE *

LITERATURE CITED

HALE,

L., AND S. D. DAVIS. 1967. The internal anatomy of home range. J. Mammal. 48(4):529-536. AUBIN, A. E. 1970. Territory and territorialbehavior of male ruffed grouse in southwestern Alberta. M.S. Thesis. Univ. of Alberta, Ed-

monton.ll0pp.

B. 1967. Movements of female ruffed grouse during the mating season. Wilson Bull. 79(1):28-36. 1968. A radio-package harness for game birds. J. Wildl. Manage. 32(3):630632. BRAY, J. R., D. B. LAWRENCE, AND L. C. PEARSON. 1959. Primary production in some Minnesota terrestrial communities for 1957. Oikos 10(1):38-49. BROWN, J. L., AND G. H. ORIANS. 1970. Spacing patterns in mobile animals. Pages 239-262 in R. F. Johnston, ed. Annual review of ecology and systematics. Vol. 1. Annual Reviews, Inc., Palo Alto, Calif. COCHRAN, W. W., D. W. WARNER, J. R. TESTER, AND V. B. KUECHLE. 1965. Automatic radio-tracking system for monitoring animal

movements. BioScience 15(2):98-100. DORNEY, R. S., AND F. V. HOLZER.

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aging methods for ruffed grouse cocks. J. Wildl. Manage. 21(3):268-274. ENx, R. L. 1959. A study of the ecology of male ruffed grouse (Bonasa umbellus L.) on the Cloquet Forest Research Center, Minnesota. Ph.D. Thesis. Univ. of Minnesota, St. Paul. ll0pp. FOWLE, C. D. 1953. An analysis of the territorial behaviour of the ruffed grouse (Bonasa umbellus L.). Ph.D. Thesis. Univ. of Toronto, Toronto, Ontario. 139pp. GLADFELTER, H. L., AND R. S. McBURNEY. 1971.

AND

R. S.

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movements of ruffed grouse in Wisconsin.

ADAMS,

BRANDER, R.

J. B.,

Archibald

Wildl. Manage. 27(4):648-656. --

J.

, R. F. WENDT, AND G. C. HALAZON. 1954. Sex and age criteria for Wisconsin ruffed Wisconsin

grouse.

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Wildl. Bull. 9. 24pp. HAYNE, D. W. 1949. Calculation of size of home range. J. Mammal. 30(1):1-18. HEEZEN, K. L., AND J. R. TESTER. 1967. Evaluation of radio-tracking by triangulation with special reference to deer movements. J. Wildl. Manage. 31(1):124-141. MESLOW,E. C. 1966. The drumming log and drumming log activity of male ruffed grouse. M.S. Thesis. Univ. of Minnesota, St. Paul.

86pp.

AND W. A. STUMPF. 1966. Comparison of methods for calculating areas of animal activity. J. Wildl. Manage. 30(2): 293-304. NOBLE, G. K. 1939. The role of dominance in MOHR, C. 0.,

the social life of birds. Auk 56(3):263-273. PHILLIPS, R. L., W. E. BERG, AND D. B. SINIFF.

1973. Moose movement patterns and range use in northwestern Minnesota. J. Wildl. Manage. 37(3):266-278. R. L. 1954. Vegetation cover types and PIERCE, land use history of the Cedar Creek Natural History Reservation, Anoka and Isanti Counties, Minnesota. M.S. Thesis. Univ. of Minnesota, Minneapolis. 137pp. PITELKA, F. A. 1959. Numbers, breeding schedule, and territoriality in pectoral sandpipers of northern Alaska. Condor 61(4):233-264. RONGSTAD, O. J., AND J. R. TESTER. 1969. Movements and habitat use of white-tailed deer in Minnesota. J. Wildl. Manage. 33(2):366379. RUSCH, D.

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