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diff --git a/29688-8.txt b/29688-8.txt new file mode 100644 index 0000000..bff6c3d --- /dev/null +++ b/29688-8.txt @@ -0,0 +1,1302 @@ +The Project Gutenberg EBook of Home Range and Movements of the Eastern +Cottontail in Kansas, by Donald W. Janes + +This eBook is for the use of anyone anywhere at no cost and with +almost no restrictions whatsoever. You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: Home Range and Movements of the Eastern Cottontail in Kansas + +Author: Donald W. Janes + +Release Date: August 14, 2009 [EBook #29688] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK THE EASTERN COTTONTAIL *** + + + + +Produced by Chris Curnow, Joseph Cooper and the Online +Distributed Proofreading Team at http://www.pgdp.net + + + + + + +UNIVERSITY OF KANSAS PUBLICATIONS +MUSEUM OF NATURAL HISTORY + +Vol. 10, No. 7, pp. 553-572, 4 pls., 3 figs. +May 4, 1959 + + + + +Home Range and Movements +of the Eastern Cottontail in Kansas + +By + +DONALD W. JANES + +UNIVERSITY OF KANSAS +Lawrence +1959 + + + +UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY + +Editors: E. Raymond Hall, Chairman, Henry S. Fitch, +Robert W. Wilson + +Volume 10, No. 7, pp. 553-572, 4 pls., 3 figs. +Published May 4, 1959 + +UNIVERSITY OF KANSAS +Lawrence, Kansas + +PRINTED IN +THE STATE PRINTING PLANT +TOPEKA, KANSAS +1959 + + + + +Home Range and Movements +of the Eastern Cottontail in Kansas + +By + +DONALD W. JANES + + +INTRODUCTION + +A knowledge of the home range and movements of the cottontail +(_Sylvilagus floridanus_) is one of the most important prerequisites +for estimating effectively its numbers and managing its populations. By +comparing results obtained from different methods, previously used, for +determining the size of the home range I have attempted to develop a +more valid procedure. + +The study here reported upon was made on the University of Kansas +Natural History Reservation (Sec. 4, T. 12S, R. 20E), the +northeasternmost section of Douglas County, Kansas, approximately 6-1/2 +miles north-northeast of the University campus at Lawrence. The +590-acre reservation, situated in the ecotone between the eastern +deciduous forests and the prairie of the Great Plains near the north +edge of the Kansas River Valley, has been protected as a "natural area" +since 1948 (Fitch, 1952). It consists of tree-covered slopes, and flat +grass-covered hilltops and valleys. Two limestone outcrops follow the +contours about five and 20 feet below the tops of the hills. + +The 90-acre study area consists of a valley bordered on the north by a +wooded slope and on the southeast by another wooded slope adjacent to a +narrow hilltop, east of which is another wooded slope. The area is thus +an alternating series of three wooded slopes and two grass-covered, +relatively level areas. + +The wooded slopes rise from the valley for about 125 feet at a grade of +approximately 16 per cent. There is a sharp increase in grade to 36 per +cent 100 feet below the top of the hills. A natural terrace 50 feet to +100 feet wide parallels the hilltop at the base of the 36 per cent +incline. + +The vegetation of the northwest-facing wooded slopes has been described +by Packard (1956). It consists of American elm (_Ulmus americana_), +shagbark hickory (_Carya ovata_), chestnut oak (_Quercus +muehlenbergii_), black oak (_Quercus velutina_), and black walnut +(_Juglans nigra_), in that order of dominance. Honey locust (_Gleditsia +triacanthos_) and hackberry (_Celtis occidentalis_) are also present. +Shrubs and herbs of the lower story include greenbriar (_Smilax +hispida_), wild grape (_Vitis vulpina_), Virginia creeper +(_Parthenocissus quinquefolia_), coralberry (_Symphoricarpos +orbiculatus_), gooseberry (_Ribes missouriense_), bluegrass (_Poa +pratensis_), sedges (_Carex_ sp.), poison ivy (_Rhus radicans_), and +white snakeroot (_Eupatorium rugosum_). + +The flat hilltops are covered by a mixture of grasses and forbs but are +dominated by awnless brome (_Bromus inermis_). Foxtail (_Setaria +glauca_), false redtop (_Triodia flava_), and panic grass (_Panicum +clandestinum_) also occur commonly. Awnless brome is dominant in the +valley (Pl. 46, fig. 1; Pl 47, fig. 2) except in the eastern end where +bluegrass is dominant (Pl. 45). + +Near the tops and bottoms of the slopes barbed wire fences separate the +woodlands from the grasslands, which were grazed until 1948. The +borderline between woods and grasslands is well defined but woody +plants are rapidly encroaching into the grasslands. Young Osage orange +(_Maclura pomifera_), American elm, and hackberry are common trees +encroaching on the grasslands. The edge vegetation between woods and +fields (Pls. 45 and 47) includes smooth sumac (_Rhus glabra_), +coralberry and wild plum (_Prunus americana_). The lowland edges are +characterized by blackberry (_Rubus argutus_), greenbriar and +elderberry (_Sambucus canadensis_). Plates 45, 46 and 47 all show local +habitat in situations where traps were actually operated. Fitch +(1952:8-22), Leonard and Goble (1952:1015-1026) and Martin +(1956:366-372) have described parts of the Reservation that include the +study area. + +I am grateful to Professor Henry S. Fitch for guiding my work, to +Professor Rollin H. Baker for suggestions and encouragement in the +early part of the study, to Mr. Robert L. Packard for certain trapping +records that supplemented my own, and to Professor E. Raymond Hall for +valuable suggestions. Norma L. Janes, my wife, typed the manuscript. +Photographs were taken by me. The State Biological Survey of Kansas +provided funds, equipment, and transportation. + + +METHODS AND TECHNIQUES + +Schwartz (1941), Dalke and Sime (1938), Dalke (1937 and 1942), +Hendrickson (1936), and Allen (1939) estimated the home range of the +cottontail by drawing, on a map, straight lines that connected all +marginal points of capture in live-traps. The resulting home ranges +were polygonal figures. Haugen (1942) altered this method by drawing +lines that connected points midway between the actual points of capture +and the next outermost traps in the grid. Fitch (1947) used a method +for enclosing all points of capture in a circle or ellipse that +represented the home range boundaries and expressed home range as the +diameter of these figures. Another method, which has been used to +determine the home range of birds, is to map the movements of an +individual as it is observed. Stebler (1939) suggested the use of +tracking records to determine home range. Connell (1954) expressed home +range of cottontails as the average distance traveled from a computed +center of activity. The method was originally proposed by Hayne (1949). + +The methods used by other investigators to calculate the home range of +the cottontail have yielded estimates varying from 0.1 acre to 100 +acres. Such wide variations in the estimated size of home range may +result from the use of different methods and from insufficient data. +The data obtained from live-trapping are not fully adequate because +traps cannot sample, in time and space, the entire home range of an +individual. Also, "trap habit" or "trap shyness" may distort the +apparent shape of the home range. In order to compare these methods I +have calculated home range from my data by each of five different +methods. The results are shown in Table 1. + +No two methods yielded exactly the same results. To utilize all +available data for each individual, I recorded on a map the locations +of capture in live-traps, nests and forms, locations where the animal +was observed in the field and the routes that it took between them. At +the end of the study a line was drawn on the map to enclose the areas +where the cottontail was known to have been. + +Live-traps were operated intermittently at 130 stations between +December 8, 1954, and February 10, 1956. Sixteen cottontails were +marked in the same area by Robert L. Packard in 1954. Data from 7850 +trap nights were used in this study. The traps were set at fifty-foot +intervals and the pattern approximated that of a grid in habitat +favorable for cottontails such as at edges of woodland. In wooded areas +traps were placed at fifty-foot intervals parallel to the edges. Traps +were not set on areas of poor habitat where neither cottontails nor +their sign were observed. + +The traps were operated eight to twelve days and closed for two to five +days intermittently throughout the trapping period except in the months +of April to August, 1955, when trapping was unsuccessful because the +cottontails then were not attracted by bait. + +Two kinds of traps were used in my study: those made by the National +Live Trap Company and those described by Fitch (1950). Both types +performed well and were serviceable under ordinary field conditions. +Experiments were made periodically throughout the trapping period to +determine which bait was most attractive to cottontails and least +attractive to birds, rodents, skunks, raccoons, and opossums. All of +these animals hindered operations by stealing bait and springing traps. +Corn, scratch-feed, carrots, parsnips, tomatoes, lettuce, apple, +cabbage, raisins, sorghum, sugar candy, and onions were used as bait. +Corn and scratch-feed attracted cottontails best in all seasons. Corn +was superior to scratch-feed, which was quickly stolen by small birds +and rodents. Eighty-nine cottontails (40 females, 49 males) were +captured in the course of the study. + +Cottontails were marked individually at the time of first capture. When +necessary, the markings were renewed at later captures; in all such +instances the same color codes and numbers were retained for each. +Markings were of four types: numbered ear tags, colored ear ribbons, +colored tail fur, and colored feet. It was intended to make each +individual cottontail recognizable in a trap or in the field. +Occasionally when a predator had killed and eaten a cottontail the tail +and feet remained and, when dyed, they provided important clues to the +identity of the individual. However, the color of the feet is not +ordinarily discernible in the field while the rabbit is alive. + +Monel metal ear tags (size No. 4, National Band and Tag Co.) were +punched through the lateral or posterior fold of the ear close to its +base (Pl. 48), one in each ear as insurance against possible losses. +However, only three tags were pulled out of the ears and lost in the +course of this study. In no instance was identity of an individual +cottontail lost. The tags caused no damage to the ears over a period of +21 months. + +Trailing in snow is an effective method of studying the daily +activities. The record preserved by the tracks becomes somewhat +confused after the snow has lain on the ground for more than one night, +and after the third night it is impossible to read the surface of the +snow. The first day of thaw usually ends tracking because the +investigator loses the trail when it crosses a patch of bare ground. +The use of a dye on the feet of the individual to be trailed eliminates +much of the difficulty of determining which tracks are to be followed. +One or more feet can be dyed when the investigator handles the animal +in releasing it from the trap. The trail of dyed footprints is +distinguishable from all other rabbit tracks in the area. Even when +only patches of snow remain, the animal can be followed by checking the +edges of the snow for the emerging footprints of the marked rabbit. The +same dye is used to color the tails. The color persists in the +footprints for about three hours, over a distance of 600 to 800 yards. +The animal leaves only a small spot in each footprint, but when it +pauses, the mark is large. Red dye makes the most conspicuous mark in +the snow. + +Thirty-one rabbits were trailed 68 times with one to six records for +different individuals. Almost two-thirds of the trailing records +pertain to males, which were caught more easily than females while snow +was on the ground. + +The trail, for both sexes, in those individuals released and followed +immediately was longer than in individuals released and allowed to move +away before they were followed. The area ranged over by a trailed +individual was not significantly greater in either case. The area +ranged over was greater by day than by night. Individuals continued to +move while being followed in the day, but stopped to forage and look +about at night. + +Records were obtained by identifying cottontails that I flushed from +forms as I walked through the study area, sometimes using a +noise-making device or dragging a rope. Regular search was made along +the hilltop rock outcrops, under which hiding cottontails could be +identified with the aid of a flashlight. Forms in brush piles, and +thickets were visited and the inhabitants identified. Other persons, +working on the study area, supplied some of the records of cottontails +that were seen alive or found dead. Also through binoculars or a +telescope I watched movements of undisturbed individuals. Twenty-three +individuals were identified 59 times. Nine females were seen 28 times +and 14 males were seen 31 times. Sixty-five other individuals were +seen, but could not be identified in the field. + + +MOTIVATION AND EXTENT OF MOVEMENTS + +The home range is an area in which an animal carries on its normal +activities of eating, resting, mating, caring for young, and escaping +from predators. The cottontail establishes a definite home range and +may live its entire life within this area, which permits familiarity +with food sources, hiding places, and escape routes. + +The cottontail usually establishes its home range in the area where it +was born, being semi-gregarious and tolerant of crowding. Eight +cottontails that were captured and marked as young remained in the area +of original capture after becoming adults. Two of them lived 17 months +in the same area, two lived 14 months, two lived 13 months, one lived +12 months and one lived eight months. No young were observed to have +moved to another home range after they matured, although some may have +moved off the study area and thereby escaped observation. + +Young become independent and are seen foraging and moving about by the +time they weigh 200 to 300 grams, at an age of four to six weeks. They +associate with other young of the same litter and neighboring litters, +and frequently frolic together. When two to three months old and +weighing 400 to 700 grams they begin to live a more solitary life and +usually rest alone in forms. Fourteen young between one and six weeks +of age never were recorded to have moved more than 150 feet. + +The population reaches its peak in August or September; home ranges +varying in size from one-half acre (in young ranging in size from 150 +grams to 800 grams) to 12 acres, in adults, are superimposed upon each +other. In a woodland area of approximately 21 acres 33 cottontails were +living together in September, 1955. + +As the growing season ends and winter approaches, the amount of food +available to the cottontail decreases and the cover becomes sparser +(Pls. 45 and 46); predators, disease, and weather take their toll of +the young. The survivors must move farther to find adequate food and +cover. The home range of the cottontail in the first winter is +overlapped by the home ranges of the other members of the same litter, +and members of other litters, as the home range is enlarged to +approximately its full size. By April the population reached its annual +low point; nine of the original 33 cottontails were known to have +survived on the 21-acre area of northwest-facing wooded slope south of +the pond. + +Foremost among the needs of the cottontail are food and cover. Daily +movements motivated by these needs are the most frequent and most +extensive that it makes. Movements such as are associated with courting +and mating, escaping severe weather, escaping from predators, and +caring for young are seasonal or irregular in occurrence. + +Because the abundant vegetation of summer provides adequate food and +cover, movements made while foraging and seeking concealment are less +extensive than those made in winter when leafy vegetation is absent and +food is scarce. The average length of trails of foraging cottontails +was 175 feet per day in summer (11 individuals observed without +disturbance) and 325 feet per day in winter (22 individuals trailed or +observed without disturbance). + +In the spring and summer cottontails forage mostly near woodland edges +for grass and herbs, and usually wander no more than 40 feet into the +grasslands from the protection of woodland edges and thickets. In +autumn and winter cottontails forage in woods and along woodland edges +for bark of trees and shrubs and for fallen fruits of trees. Ninety-two +per cent of all fecal pellets found in grassland were within 40 feet of +cover suitable for cottontails. + +Movements made by the cottontail while foraging appear aimless; typical +behavior consists of progression with a hesitant gait of two or three +hops, a stop to eat, another series of hops and another stop. +Footprints made by this movement are about 12 inches apart. With +occasional spurts of hopping the individual moves perhaps ten to twelve +feet where it stops and begins to eat again. The area in which the +individual forages is usually elongated with its long axis parallel to +the edge except in areas of uniform habitat (such as large patches of +coralberry) where the area covered tends to be more nearly circular. +Cottontails observed foraging were estimated to utilize 10 to 20 per +cent of the home range area in one evening. Paths or runways are not +ordinarily utilized by foraging cottontails. + +In seeking protection from predators or from the weather, cottontails +move farther in winter than in summer. The average length of trails of +cottontails flushed by me in the study area was 80 feet in summer and +210 feet in winter. + +When cottontails were released from live-traps they ran an average of +30 feet before stopping to look about. Cottontails always ran toward +the densest cover within 50 feet of the point of release. Ten per cent +of the cottontails released from live-traps did not stop running until +out of sight (always more than 30 feet). + +Movements made by cottontails escaping from predators differ from +movements made while foraging. The gait is a full run, often eight to +ten feet between footprints in snow; the trail is either straight or +slightly zig-zag. If possible, the individual will take refuge in a +hiding place such as a rock outcrop, brush pile, or thicket. Eight +cottontails emerged from such hiding places an average of 22 minutes +after the disturbance ceased. + +If unable to find a hiding place a pursued cottontail will run 600 to +1200 feet while circling and returning to the area from which it ran. +If not closely pursued, it will usually (in 68 per cent of the +instances) not enter hiding places, but continue to run ahead of the +pursuer. Of 70 released from live-traps and followed, 23 sought refuge +in hiding places. The others ran slowly (4 to 7 feet between footprints +in the snow) with frequent pauses to look and listen; they usually +succeeded in keeping out of my sight. Twelve times the trails of +cottontails followed in this manner passed near a form, or other +resting place; always the cottontails had paused at the resting place, +and twice the individual went into the resting place and ran out again +when I approached. Resulting trails were almost circular, covering most +(70-90 per cent) of the home range; sometimes three or four complete +circuits were made. The trails made when I pursued cottontails ranged +in length from 800 to 3,000 feet. A trail recorded for an individual on +one night was almost identical with another trail for the same +individual recorded another night. The cottontail is not easily driven +out of its home range. Paths or runways are used by cottontails +escaping from predators in dense vegetation along fence lines, in +thickets, or brush piles, or in snow that is eight or more inches deep. + +Most of the year cottontails rest in forms of grass or brush near +woodland edges but in extremely cold or hot weather they seek the +greater protection of the woods. Movements are limited to the woodlands +in severe weather, especially when deep snow makes travel difficult. + +Hilltop rock outcrops on the area provide excellent protection for the +cottontail especially from low temperatures and freezing rain or +blowing snow. Eighty per cent of the cottontails resting under the rock +outcrops were found in severe winter weather. Fifteen per cent were +found in severe summer weather, and five per cent were found at times +when the weather was not severe. Undercut creek banks and exposed tree +root-systems in eroded gullies were favorite hiding places. Brush that +had accumulated in the ravines and stream beds also was used for cover +by the cottontail. + +In heavy rain a cottontail may move along, neither hopping nor running, +with its body close to the ground, head low, ears laid back. Losing its +customary alertness it may pass a person without seeing him. At times, +I have been able to approach almost close enough to seize one of the +miserable creatures. In deep snow cottontails may progress with long +bounds carrying them high enough vertically, to clear the surrounding +snow. + +Courting activities were seen only in evening. Four male cottontails +and three females were observed to move an estimated 1200 feet in an +evening (1-1/4 hours) while chasing each other around in an area of +less than an acre. It is presumed that this activity was in addition to +normal movements made while foraging. + +Seven females known to be pregnant or lactating were not observed to +move more than 100 feet away from their centers of activity. Their +nests were never found. At the same time the males were moving over +much larger areas. + +The cottontail is most active in the evening or early morning. Of those +for which time of capture in live-traps was known 70 per cent were +captured in the evening between dusk and 11:00 P.M., 10 per cent were +captured between 11:00 P.M. and dawn, and 20 per cent were captured +after dawn. Nocturnal rodents and carnivores often stole bait in the +night; the percentage of capture of cottontails after dawn might have +been larger had bait remained in all the traps. + +Except for those flushed, cottontails were seldom seen by day. In walks +through the study area approximately three times as many cottontails +were flushed at night as in the daytime. On cloudy days cottontails +were active longer than on bright days. On dark nights more cottontails +were captured in live-traps between 11:00 P.M. and dawn than on bright +moonlight nights. + +Cottontails were more active (as determined by trap success and +frequency of observation) at temperatures between 0° F. and 33° F. than +at temperatures between 33° F. and 55° F. Activity of the cottontail +increased as the temperature of the air decreased. Activity increased +in proportion to the percentage of ground covered by snow. Activity of +the cottontail decreased as precipitation increased; there was less +activity in rain than in snow and less activity in wet snow than in dry +snow. Activity did not vary significantly with depth of snow until snow +was more than 8 inches deep, when activity decreased abruptly. + +The average of the longest distance traveled between captures for +cottontails whose entire home ranges were thought to have been sampled +was 900 feet for males and 684 feet for females. The average of the +maximum distance across the home range for cottontails whose home range +had been thoroughly sampled was 1019 feet for males, and 936 feet for +females. + +The home range is used by the cottontail in different ways, depending +on the needs of the individual and the condition of the habitat. In +uniform habitat the home range is roughly circular and is used most +near its center and least toward its periphery. The entire home range +is traversed every four or five days. The center of the home range has +been called the "center of activity" (Connell, 1954). + +In habitat of alternating woodland and grassland, such as that on the +University of Kansas Natural History Reservation, two centers of +activity often developed in the home range of a cottontail, at opposite +edges of a tract of woodland. The individual concentrated its movements +near one center for three to five days then moved to the other center. +Pursuit by a predator, random movement, or other cause may be +responsible for shift from one edge to another. + +PLATE 45 + +[Illustration: FIG. 1. "House Field" viewed from the northeast corner, +looking southwest, March 3, 1956, showing the condition of vegetation +in winter. Traps were operated on a 50-foot grid throughout this area.] + +[Illustration: FIG. 2. Same area as above, on July 10, 1956, showing +the condition of vegetation in summer.] + +PLATE 46 + +[Illustration: FIG. 1. View southeast from the north edge of "Picnic +Field" showing condition of the vegetation on December 3, 1954. Trap +lines were placed along woodland edge from which this picture was +taken, along road where vehicle is parked, along creek beyond road, +along edge of field beyond creek, and along edge of woods in +background.] + +[Illustration: FIG. 2. Summer aspect of the vegetation, on July 14, +1955; same view as shown in Fig. 1.] + +PLATE 47 + +[Illustration: FIG. 1. Condition of vegetation along woodland border +northeast of Reservation headquarters on December 3, 1954. Camera was +facing southwest. Traps were operated along this edge and in woods to +right and in background.] + +[Illustration: FIG. 2. Condition of vegetation at edge of "House Field" +on July 14, 1955. Scale is shown by 4-1/2 foot tripod. Brome grass was +approximately two feet high. Traps were operated in a grid in this +area.] + +PLATE 48 + +[Illustration: FIG. 1. Cottontail in bag with ear protruding, ready for +marking with nylon ribbon and metal ear-tag shown in upper right hand +corner. × 1/4.] + +[Illustration: FIG. 2. Cottontail bearing ear-tags and ribbon. × 1/4.] + +When moving undisturbed through the woods cottontails usually do not +pause to forage or associate with other cottontails, but keep to a +straight route except in severe weather, when, as noted above, they +find resting places in the woods. Ninety-two per cent of the +cottontails captured in live-traps were captured within 100 feet of a +woodland edge; six per cent were captured in the woods, more than 100 +feet from an edge, and two per cent were captured in grassland more +than 100 feet from the edge. In winter, when the air temperature was +less than 20° F., 22 per cent of the cottontails were captured +in the woods more than 100 feet from the edge. + +The maximum distance between two centers of activity of an individual +was 700 feet, average 550 feet. If two centers of activity were +maintained, the cottontail usually traversed the entire home range +every seven to 11 days. + +In no case was a cottontail known to have lived in two woodland edges +which were separated by open grassland. Cottontails usually did not +move more than 75 feet from suitable cover. In winter when herbaceous +vegetation was dormant cottontails did not cross open fields. + +Forms in grass clumps were the usual resting place for cottontails, but +others in brush piles, rock outcrops, and tree stumps were also used. +On the average a cottontail maintained 3.5 forms. If disturbed +repeatedly at a form, a cottontail would permanently desert it. On +seven occasions a cottontail used a form that had been used by another +within 24 hours. Three cottontails used the same shelter under a rock +ledge in five days; one was under the ledge on December 17, 1955, and +another was there on December 18. The first was there again on December +20 and a third was there on December 21. The original cottontail had +returned by January 2, 1956. There may be 20 to 30 resting places used +by cottontails within a single home range area since five to seven +cottontails may live there as co-occupants at one time. Two +cottontails, both males, lived together in a _Smilax_ thicket for three +weeks, resting within 15 inches of each other. Occasionally a female +was present in the same thicket, and rested about three feet from the +males. + +A male trapped on land adjoining the Reservation and confined overnight +at the Reservation headquarters escaped the next day and was seen 32 +days later, 1800 feet from the point of escape, back in the area where +it was originally captured. + +A female confined for observation, escaped and ran in the direction +opposite from her home. Subsequently she was seen on four different +occasions, over a period of one month, in her original home range, +1,100 feet from the point where she escaped. Both these animals which +made homing movements had been removed in cloth bags from their homes. + +Another cottontail removed from its home range and taken to the +laboratory building to be marked, escaped and ran to a nearby wooded +hillside without pursuit where it could be observed because of snow on +the ground and lack of leaves on the trees. The animal ran and hopped +about over a one-half acre area. Its movements seemed to be unoriented +and it frequently stopped and stood on its hind legs in order to look +about. After 10 minutes of this behavior, a red-tailed hawk (_Buteo +jamaicensis_) screamed as it flew overhead. The cottontail, stimulated +by seeing and/or hearing the hawk, ran faster, moving in circles until +it disappeared from view five minutes later. When last observed the +cottontail was 1,700 feet from its home range and was headed in the +opposite direction. It had passed several potential shelters but had +not attempted to use them, presumably because it was not familiar with +the area. Although for several months afterward traps were operated in +the cottontail's home range area and in the area where it escaped, the +animal was never recaptured. + + +SIZE AND SHAPE OF HOME RANGE + +Of the 89 cottontails observed in the study, 35 were captured in +live-traps only once and were never seen in the field or trailed. The +remaining 54 served for calculation of home range by one or more +methods. The minimum, maximum and average home ranges for these 54 +individuals were calculated by each of five methods. All individuals +for which any area was recorded were included in the average. +Incomplete home ranges lower the averages. According to the most +reliable method (Composite Method) 30 male cottontails had home ranges +of between 0.46 acre and 12.19 acres and 24 female cottontails had home +ranges of between 0.46 acre and 12.62 acres. The average for males was +5.05 acres and the average for females was 4.81 acres. The average for +all 54 cottontails was 4.86 acres. + +Because of irregularities in live-trapping and field observation some +cottontails were more intensively studied than were others; one +cottontail was followed one time only, while another was trapped 26 +times, followed three times, and seen in the field six times. It was +necessary to determine which cottontails had been studied sufficiently +to determine the approximate extent of their home ranges. + +[Illustration: FIG. 1. Correlation between average size of home range +and number of "peripheral points" (marginal records, which form angles +when the range is outlined). On the average, a sufficient number of +records to yield approximately nine peripheral points must be obtained +before the full extent of the home range is revealed.] + +The average area, in acres, of home range was plotted, on a graph, +against the number of peripheral points (Figure 1). When a home range +had nine or more peripheral points, on an average, the size of home +range did not increase significantly with additional captures, +observations in the field, or records of trails. Home ranges with less +than nine peripheral points were likely to be increased in size with +each new observation. A similar situation has been shown by previous +authors who have plotted the size of home range against number of +captures in live-traps, where only live-trapping was used to gather +data. Therefore, in my study, home ranges with nine or more peripheral +points were considered to be adequately studied. + +Data for eighteen cottontails that had been studied sufficiently to +determine the full extent of their home ranges were used to calculate +minimum, maximum, and average home range by each of five methods (Table +1). The methods used by Schwartz (1941), Dalke and Sime (1938) and +Allen (1939) yielded results which were lower than any others, +presumably because only live-trap data were used and because straight +lines were used to connect traps in which cottontails were captured. +The "composite method" was considered the most reliable because it +utilized all data gathered for each individual and because with this +method the home range boundaries were drawn to enclose all areas in +which the cottontail lived and excluded all areas in which the +cottontail was not known to have been. The method used by Fitch (1949) +agreed most closely with the "composite method" and suggests to me that +the home range of animals can be estimated with reasonable accuracy by +this method when field observation or trailing are not feasible. The +composite method is superior to others for studying the home range and +movements of cottontails. + +Of the individuals whose entire home ranges had been thoroughly +studied, nine males had home ranges of between 4.72 acres and 12.19 +acres with an average of 8.92 acres; nine females had home ranges of +between 2.42 acres and 12.62 acres with an average of 7.76 acres. The +average size of home range for both sexes was 8.34 acres (Table 1). + +TABLE 1.--HOME RANGES, IN ACRES, OF 18 COTTONTAILS ON THE +RESERVATION IN 1956, COMPUTED BY FIVE DIFFERENT METHODS. + +======================================================================= + | | Number of | Average | Maximum | Minimum + Method | Sex | individuals| area | area | area +----------------------+-----+------------+---------+---------+--------- +Allen (1939), | (M) | 9 | 2.00 | 6.78 | .30 +Dalke and Sime (1938) | (F) | 9 | 2.54 | 7.20 | .35 +and Schwartz (1941) | all | 18 | 2.27 | 7.20 | .30 + +-----+------------+---------+---------+--------- + | (M) | 9 | 4.01 | 12.89 | 1.05 +Fitch (1947) | (F) | 9 | 5.68 | 11.50 | 1.84 + | all | 18 | 4.85 | 12.89 | 1.05 + +-----+------------+---------+---------+--------- + | (M) | 9 | 7.20 | -- | -- +Fitch (1949) | (F) | 9 | 9.00 | -- | -- + | all | 18 | 8.40 | -- | -- + +-----+------------+---------+---------+--------- +Tracking and | (M) | 9 | 8.74 | 11.15 | 3.54 +field observations | (F) | 9 | 8.62 | 12.18 | 5.51 + | all | 18 | 8.69 | 12.18 | 3.54 + +-----+------------+---------+---------+--------- +Tracking and field | (M) | 9 | 8.92 | 12.19 | 4.72 +observations plus | (F) | 9 | 7.76 | 12.62 | 2.42 +live-trapping | all | 18 | 8.34 | 12.62 | 2.42 +----------------------+-----+------------+---------+---------+--------- + +[Illustration: FIG. 2. Maps showing home ranges of cottontails in +relation to woodland and open fields on the study area. One inch equals +approximately 470 feet. Each home range is shown slightly rounded from +the polygonal figures obtained by connecting actual points where the +animal was recorded. _Upper Left._ An area of 4.6 acres occupied by a +cottontail in winter, increased to 6.5 acres in summer by the animal +crossing a narrow grassland strip to another woodland edge. _Lower +Left._ Showing increasing size of home range of a female cottontail; in +July, 1954 (at age of three weeks), she had a range of .25 acre; +September, 1954, 1.5 acres; December, 1954, 8 acres; and December, +1955, 11 acres. _Right Half._ Two home ranges of a cottontail which +moved from its original area to occupy a new one 410 feet farther north +across a field in September and October, 1955.] + +Cottontails range over a larger area in summer than they do in winter +because suitable cover and food is more abundant in summer. One +cottontail (Figure 2, upper left) lived in a woodland home range of 4.6 +acres in the winter but increased the range to 6.5 acres in summer by +crossing the narrow overgrown end of a field to another woodland area. +Another cottontail (Figure 3, top part) lived in a woodland home range +of 7.9 acres in winter but in summer increased the home range to 9.5 +acres by including also a part of an adjacent field. Other cottontails +increased their home ranges in summer by five to 15 per cent. + +On the average, male cottontails had a larger (by 13 per cent) home +range than females probably because of the increased activity of males +in the breeding season and the decreased activity of females when +pregnant and caring for young. Nevertheless, some of the largest home +ranges measured were those of females. + +[Illustration: FIG 3. Diagrams showing home ranges of cottontails in +relation to woodland and open fields on the study area. One inch equals +approximately 545 feet. Each home range is shown slightly rounded from +the polygonal figures obtained by connecting actual points where the +animal was recorded. _Upper._ Winter range of 7.9 acres (solid line) +increased to 9.5 acres in summer by area in dashed line. _Lower._ +Overlapping home ranges of four of the many cottontails living on the +study area. Each of the four cottontails occupied approximately the +same home range throughout the year.] + +The size of the home range in immature cottontails varies between 0.1 +acre and 4.0 acres, depending on the age and size of the individual. +Fourteen young cottontails between three and six weeks of age did not +leave areas of approximately one acre in each instance. Nine +cottontails between six weeks and 18 weeks of age lived in areas of +about two acres. By the time cottontails are four to five months old +they inhabit a home range of four to eight acres. + +One cottontail (Figure 2, lower left) born in July, 1954, was estimated +to have wandered over approximately 0.25 acre at an age of three weeks. +In September this cottontail occupied a home range of one and one-half +acres. By December it was five months old and occupied an area of about +eight acres. In the next year it enlarged its home range to 11.5 acres. + +The cottontail usually settles down in one area and stays there until +it dies. Changes from one home range to another are unusual, but minor +shifts, in response to changes in vegetation and weather, are common. +In one exceptional instance (Figure 2, right) a male cottontail, +occupying a home range of 11.2 acres in a woodland, suddenly shifted to +a new area that barely overlapped its former home range at one edge. +Two months after the change was first noticed the cottontail was living +in a new home range of 6.6 acres 300 feet from its original home range. +In changing from one home range to the other the cottontail traveled +along a dry stream bed and was captured there three times. + +Maps of the home ranges of four of those 18 cottontails for which +sufficient data were collected to determine the size of home range are +shown in the lower part of Figure 3. + + +SUMMARY AND CONCLUSIONS + +The home range and movements of the cottontail were studied on a +90-acre area of the University of Kansas Natural History Reservation +from February, 1954, to March, 1956. Eighty-nine cottontails were +identified in the field 59 times, trailed 70 times and captured in +live-traps 326 times in 7,850 trap nights. Home range of the cottontail +was calculated by five methods, using the same set of data, and the +results were compared. A composite method was used, which permitted the +use of more data than any other one method. + +The maintenance of a home range is of survival value to the cottontail. +Knowledge of the home range is of value to man when control or +propagation of cottontail populations is desired. Cottontails establish +a home range where they are born and enlarge it to nearly full size the +first winter. Home ranges of cottontails are overlapped by those of +others regardless of sex or age. No territory is maintained. + +The cottontail makes movements to forage, to seek cover from predators +and the weather, to reproduce, build nests, care for young, keep pace +with changes in vegetation through the year, and escape unusually +severe climatic conditions. Movements may be caused by desire for +acquaintance with surroundings and other animals, escape from +undesirable surroundings or animals, or merely release of nervous +energy. + +When foraging, cottontails moved 175 to 325 feet per day, mostly near +woodland edges, and used from 10 to 20 per cent of the home range. When +escaping from predators cottontails moved 30 to 1200 feet and used 5 to +70 per cent of their home ranges, depending on the type of pursuit. +Some cottontails that were followed, ran in almost circular courses for +as far as 3000 feet and covered as much as 90 per cent of their home +ranges. Paths or runways were not used except in deep snow or very +dense vegetation. Movements were limited by deep snow. When +temperatures were unfavorably high or low, cottontails sought cover +deep in the woods or under rock outcrops, and in dry stream beds. In +moderate weather resting places in grass forms, brush piles and +thickets were used. + +Both males and females moved farther in the breeding season than in the +rest of the year, but females that were caring for young in summer and +late spring moved shorter distances than they did when not so engaged +in autumn and winter. + +Cottontails were most active at dawn and especially, dusk, and were +more active on dark nights than on moonlight nights. Cottontails were +most active when the air temperature was between 0° F. and 33° F. and +when rain was not falling. Activity increased as the percentage of +ground covered by snow increased and as the abundance of food +decreased. Activity did not vary with physiological condition except +that as body weight decreased activity increased--probably because of +lack of food. + +The home range is used most intensively near centers of activity that +are near woodland edges or in other areas of good cover. Cottontails +often ranged through the woods and along edges but did not cross open +areas more than 75 feet wide. Cottontails use their home range most +intensively in winter when they are forced to move long distances in +poor cover, searching for food. More than one cottontail may use sites +of good cover at the same time and two or three used the same resting +place at different times. + +Two instances of homing were observed; cottontails moved 1,100 and +1,800 feet to return to their home ranges, but one cottontail that +escaped 1700 feet from home failed to return. + +The average home range of 18 cottontails for whom adequate data were +gathered was 8.34 acres. The home ranges of males averaged 1.16 acre +larger than those of females. In summer, cottontails increased their +home ranges 5 to 15 per cent by taking advantage of cover provided by +the more abundant vegetation. Cottontails three weeks to five months of +age lived in home ranges of between 0.1 and 4.0 acres and enlarged +their home ranges almost to their ultimate full size in the first +winter. + + +LITERATURE CITED + +ALLEN, D. L. + + 1939. Michigan cottontails in winter. Jour. Wildl. Mgt., + 3(4):307-322, 6 half-tone pls., 7 tables. + +CONNELL, J. H. + + 1954. Home range and mobility of brush rabbits in California + Chaparral. Jour. Mamm., 35(3):392-405, 6 figs., 2 tables. + +DALKE, P. D. + + 1937. A preliminary report of the New England Cottontail studies. + Trans. Second North Amer. Wildl. Conf., 542-548. + + 1942. The cottontail rabbits in Connecticut. State of Connecticut + Public Document No. 47, State Geological and Natural History + Survey Bull. No. 65. 1-97 pp., 22 figs., 43 tables. + +DALKE, P. D., and SIME, P. R. + + 1938. Home and seasonal ranges of the eastern cottontail in + Connecticut. Trans. Third North Amer. Wildl. Conf., 659-669 + pp., 4 figs., 9 tables. + +FITCH, H. S. + + 1947. Ecology of a cottontail rabbit (_Sylvilagus auduboni_) + population in Central California. California Fish and Game, + 33(3):159-184, 48-53 figs., 8 tables. + + 1949. Study of snake populations in Central California. Amer. Midl. + Nat., 41(3):513-579, 11 figs., 28 tables. + + 1950. A new style live-trap for small mammals. Jour. Mamm., + 31(3):364-365, 1 fig. + + 1952. The University of Kansas Natural History Reservation. Univ. + Kansas Mus. Nat. Hist, Misc. Publ. No. 4:1-38, 4 pls., 3 + figs, in text. + +HALL, E. R. + + 1951. A synopsis of the North American Lagomorpha. Univ. Kansas + Publs., Mus. Nat. Hist., 5(10):119-202, 68 figs. in text. + +HAUGEN, A. O. + + 1942. Home range of the cottontail rabbit. Ecology, 23(3):354-367, + 6 figs., 9 tables, 1 graph. + +HAYNE, D. W. + + 1949. Calculation of size of home range. Jour. Mamm., 30(1):1-18, 2 + figs., 2 tables. + +HENDRICKSON, G. O. + + 1936. Summer studies on the cottontail rabbit; _Sylvilagus + floridanus_. Iowa State Coll. Jour. Sci., 10:367-372. + +JANES, D. W. + + 1957. Body temperature in the eastern cottontail. Jour. Mamm., 38 + (1):137. + +LEONARD, A. B., and GOBLE, C. R. + + 1952. Mollusca of the University of Kansas Natural History + Reservation. Univ. Kansas Sci. Bull., 34:1013-1055. + +MARTIN, E. P. + + 1956. A population study of the prairie vole (_Microtus + ochrogaster_) in northeastern Kansas. Univ. Kansas Publs., + Mus. Nat. Hist., 8(6):361-416, 19 figs. in text. + +PACKARD, R. L. + + 1956. The tree squirrels of Kansas: Ecology and economic + importance. Univ. Kansas Mus. Nat. Hist., Misc. Publ. No. + 11:1-67, 10 figs, in text, 2 pls. + +SALMAN, D. H. + + 1948. On the home range of cottontails. Physiologia Comparata et + Oecologia, 1(2):95-109. + +SCHWARTZ, C. W. + + 1941. Home range of the cottontail in central Missouri. Jour. + Mamm., 22(4):386-392, 1 fig., 2 tables. + +STEBLER, A. M. + + 1939. The tracking technique in the study of the larger predatory + mammals. Trans. Fourth North Amer. Wildl. Conf., 203-208. + + + + + + + +End of the Project Gutenberg EBook of Home Range and Movements of the +Eastern Cottontail in Kansas, by Donald W. 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