temporal patterns of spring space use by ruffed grouse wild man vol 39 1975... · temporal patterns...

Temporal Patterns of Spring Space Use by Ruffed GrouseAuthor(s): Herbert L. ArchibaldReviewed work(s):Source: The Journal of Wildlife Management, Vol. 39, No. 3 (Jul., 1975), pp. 472-481Published by: Allen PressStable URL: http://www.jstor.org/stable/3800386 .Accessed: 06/02/2012 12:38

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TEMPORAL PATTERNS OF SPRING SPACE USE BY RUFFED GROUSE'

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-tracking system 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 drumming juvenile males. An inverse relationship was found between mean 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 drumming males 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 drumming season. Cumulative ranges of drumming males contained centrally located 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 drumming males.

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of ruffed grouse has resulted from studies in Wisconsin (Hale and Dorney 1963), northern Minnesota (Gullion 1970, unpublished report), and Alberta (Rusch and Keith 1971).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. Fi- nancial support was provided by NIH Training Grant No. 5 TO1 GM01779 and the U.S. Atomic Energy Commission (COO- 1332-106).

STUDY AREA

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 inter-

spersed 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 (Cornus stolonifera). Other lowland vegetation includes cattail (Typha latifolia) marshes and sedge (Carex spp.) meadows.

'Journal Paper No. 5629, Purdue University Agricultural Experiment Station.

2 Present address: Department of Forestry and Natural Resources, Purdue University, West Lafayette, Indiana 47907.

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

SPACE USE BY RUFFED GROUSE * Archibald 473

The upland-lowland edge is dominated by white birch (Betula papyrifera) and aspen (Populus tremuloides). Vegetation on un- cleared 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 re-

port, 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

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 av-

eraging 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 modifica-

tion 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 loca-

tion) 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

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

474 SPACE USE BY RUFFED GROUSE ?

Archibald

no correlation was found between weekly range size and number of fixes. All ranges analyzed fell within the 61.0-m (200-ft) accuracy zone (Heezen and Tester 1967); thus a "worst-possible case" 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 described by the ratio of greatest length (i.e., major axis between the two most distant fixes) to greatest width (i.e., sum of perpendicular distances from the most distant fix on each side of the major axis).

To describe how 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 maximum possible gridsquare use. For the first measure, the center of activity (Hayne 1949) was determined by computing means of the x- and y-coordinates of all fixes for a grouse in a seven-day period. The distances between weekly activity tenters and the mean of all weekly activity centers provided activity center deviations, which were then averaged. For the second measure, a tally was made of the number of seven-day periods in which each gridsquare within the cumulative range (i.e., that for all weeks a grouse was radio-tracked) was occupied one or more times. The ratio of the mean number of weeks in which gridsquares were 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 cumulative ranges of males were delineated and examined for possible territorial significance. The core area was de-

fined as all gridsquares occupied one or more times in 50 percent or more of the total weeks. For an individual radio-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 gridsquares was made on the basis of adjacency to squares with the highest number of weeks of occupation. In- tensity 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 number of fixes obtained which fell in the core area.

RESULTS Locations of 10 males and 3 females were

analyzed for a total of 83,926 fixes and 629 bird-days. Radio-marked males began drumming on 9 April 1970 and had greatly reduced drumming activity by 10 June. Thirteen 7-day data periods, from 12 March through 10 June, arbitrarily were defined, with 9 April as the first day of week 5. With the exception of one adult killed by an avian predator about three weeks before drumming began, all radio-marked males were "active, site-associated" drummers (Gullion 1966:718).

Variations in Weekly Range Size 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 averages of the daily mean temperatures for 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 and females were different, likely reflecting behavioral differences between sexes.



MALES

4

DRUMMING

c/) W

C.J z 3 3 5 5 5 4 5 5 6 6 5 5 4

8 FEMALES

w N 6

(C INCUBATION z 4

S I 2 3 3 3 3 3 ! ! I 2 3 4 5 6 7 8 9 10 11I 12 13

WEEK

Fig. 1. Seasonal variatior in range size by sex. Week 1, 12-18 March; week 13, 4-10 June. Bars depict maxima and

minima; sample size above x-axis.

Following the onset of the spring drum-

ming period in week 5, mean range size of males decreased and did not again reach the week 4-5 size until week 13 (Fig. 1). By this time drumming activity had greatly decreased. An inverse relationship is there- fore suggested between weekly mean range size of males and their level of drumming activity. The comparatively small mean

range sizes during weeks 6-12 were consistent with Eng's (1959) finding that the distance moved from the drumming log was at a minimum in the spring (16 March- 31 May).

Weekly mean range size for females increased in an approximately linear manner from weeks 4 through 9 (Fig. 1). Although other factors may have been involved, the

relatively large mean range sizes in weeks 7-9 are probably a reflection of mate-seek-

ing movements (Brander 1967), nest-site

Table 1. Mean weekly range size (ha) of adult and

juvenile drumming male ruffed grouse (sample size in

parentheses).

Adult Juvenile Week males males Difference

1 2.48 (3) 2 4.30 (2) 3.88 (1) 0.42 3 3.64 (3) 3.07 (2) 0.57 4 4.00 (3) 4.18 (2) -0.18 5 4.47 (3) 3.48 (2) 0.99 6 3.66 (2) 3.37 (2) 0.29 7 2.78 (4) 2.44 (1) 0.34 8 2.90 (4) 2.10 (1) 0.80 9 2.94 (3) 2.35 (3) 0.59

10 3.16 (3) 1.74 (3) 1.42 11 2.24 (2) 2.14 (3) 0.10 12 2.33 (2) 3.06 (3) -0.73 13 4.34 (2) 3.59 (2) 0.75

selection, and increased nutritional requirements prior to and during laying. In con-

trast, ranges during weeks 10-12 were rel-

atively small, indicating that females have a much more restricted movement pattern during incubation.

Mean range size of adult (22 or more months old) males exceeded that for juvenile (10-12 months old) males in 10 of 12 weeks (Table 1). The age difference in

range size of drumming males (ci = 0.45 ha) was significant at the 0.025 level (Wilcoxon signed ranks test, Sokal and Rohlf 1969:400). The propriety of using the test is, however, open to question because the means are based on small, unequal samples. Previous

findings that juvenile males in general are more mobile than adult males (Hale and

Dorney 1963, Rusch and Keith 1971) reflect the greater mobility of non-drumming, non- territorial males which are primarily juveniles (Gullion 1967, Rusch and Keith 1971).

No pronounced sexual differences or seasonal trends in range shape were apparent (Table 2). Large mean length-width ratios

(1.71, 1.68) for two males radio-tracked before and during the drumming season appeared to be a reflection of the linear

configuration of the habitat in which they


476 SPACE USE BY RUFFED GROUSE - Archibald

Table 2. Shape and variation of location in space of ranges of ruffed grouse.

Ratio of mean to maximum Ratio of greatest possible gridsquare use length to greatest Activity center

width of deviation Entire Core area within Total weekly range (km) cumulative range cumulative rangec

No. no. birds weeks Meana SE Meanb SE Mean SE Mean SE

Males Before and during

drumming period 5 36 1.60 0.103 0.045 0.016 0.423 0.055 0.785 0.052

During drumming period only 4 24 1.32 0.159 0.019 0.008 0.542 0.070 0.882 0.041

Total 9d 60 1.47 0.191 0.034 0.019 0.476 0.085 0.828 0.068

Females 3 29 1.62 0.140 0.123 0.041 0.243 0.038

a Average of the mean length-width ratio for each grouse in category. b Average of the mean activity center deviation for each grouse in category. c Not computed for females. d Does 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 re-

markably 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

??

..0.16

KM

01 MILE

Fig. 2. Cumulative range of iuvenile male 1628, weeks 3-12 (26 March-3 June).

.................. ....... .....

X. .;.t';; 8?? ?????? ??

~???2?? ? '?'0 ..;....??5????

X...?.....

Fig 2.Cuulaiverage f jveilemal 128,wees -12(26Mach- Jue)

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

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 "drum-

ming season" (9 April-10 June). Cumula- tive 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


478 SPACE USE BY RUFFED GROUSE * Archibald

20 - 20-

u) w

IS -15

MALES FEMALES

z

N r 10 -10-

w

-5 5- 4

I _ _

DRUMMING

0 -.-.... ......•.....? ?0 v?1?

v v V

0 I 2 3 4 5 6 7 8 9 1011 12 13 0 1 2 3 4 5 6 7 8 9 10 111213

WEEK WEEK

Fig. 3. Development of the ranges of individual male and female ruffed grouse.

and activity center deviations (Table 2) provide documentation of 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 characterized as an area of: (1) 2.26 ? 0.54 (1 SD) ha in size, comprising 34.5 - 10.37 percent of the cumulative

range, (2) consistent occupancy over time, made up of gridsquares used in 82.8 ? 6.79 percent of the total weeks, and (3) intensive use, containing 79.6 + 16.17 percent of the total locations. Similar to the case with cumulative range, there was a tendency for

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

drumming period (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-tracked in weeks 1-5 and then killed by an avian predator. Apparently this bird had not begun to use a central area consistently prior to the onset of drumming.

In all cases, the primary drumming log (Gullion 1967) was contained within the core area, and the core area was centrally located within the cumulative range. With



N J

I-7

-0 0

LOWLAND 0 00W

UPLAND USE-INTENSITY BY 1640 > 1%

0 USE-INTENSITY BY 1641 0 0/

144

-- =

BOUNDARY BETWEEN RANGES

[ EXCLUSIVE USE BY 1640

= EXCLUSIVE USE BY 1641

U = ZONE OF OVERLAP

0.16 KM

0.1 MILE

Fig. 4. Ranges of juvenile males 1640 and 1641, 14-20 May.

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 neigh-

boring males (Aubin 1970). Although de- tailed consideration of these factors is be- yond 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 associ- ation.

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 neigh- bor'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 use- intensity 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 situa- tions. Consequently, females showed an increasing tendency to use upland habitat beginning with the onset of nesting activ- ities.

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 (Glad- felter 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 concen- trated use which was reaffirmed by adver- tisement 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 ta-

pered off markedly during week 13, these data support the hypothesis that drumming is involved in the maintenance of stable

spatial relationships among males.



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. 1967. Selection and use of drumming sites by male ruffed grouse. Auk 84(1): 87-112.

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Accepted 8 January 1975.


temporal patterns of spring space use by ruffed grouse wild man vol 39 1975... · temporal patterns...

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